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Full text of "Proceedings of the Indiana Academy of Science"

Digitized by the Internet Archive 

in 2012 with funding from 

LYRASIS Members and Sloan Foundation 



http://archive.org/details/proceedingsofin791969indi 



PROCEEDINGS 

of the 

Indiana Academy 
of Science 

Founded December 29, 1885 



Volume 79 
1969 



Marion T. Jackson, Editor 

Indiana State University 

Terre Haute, Indiana 47809 



Spring- Meeting- 
April 25-26 
Hanover College, Hanover, Indiana 47243 

Fall Meeting- 
October 23-24 
Hanover College, Hanover, Indiana 47243 

Published at Indianapolis, Indiana 
1970 



1. The permanent address of the Academy is the Indiana State Library, 140 N. 
Senate Ave., Indianapolis, Indiana 46204. 

2. Instructions for Authors appear at the end of this volume, P. 484-485. 

3. Exchanges. Items sent in exchange for the Proceedings and correspondence con- 
cerning exchange arrangements should be addressed : 

John Shepard Wright Memorial Library of the Indiana Academy of Science 
c/o Indiana State Library 
Indianapolis, Indiana 46204 

4. Proceedings may be purchased through the State Library at $5.00 per volume. 

5. Reprints of technical papers can often be secured from the authors. They cannot 
be supplied by the State Library nor by the officers of the Academy. 

6. The Constitution and By-Laws reprinted from v. 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 care of the State Library. Papers published in the Proceedings of the Academy 
of Science are abstracted or indexed in appropriate services listed here : 

Annotated Bibliography of Economic Geology 

Bibliography of Agriculture 

Bibliography of North American Geology 

Biological Abstracts 

Chemical Abstracts 

Chemisches Zentralblatt 

Current Geographical Publications 

Geological Abstracts 

Metallurgical Abstracts 

Pesticides Documentation Bulletin 

Psychological Abstracts 

Review of Applied Entomology 

The Torrey Bulletin 

Zoological Record 



TABLE OF CONTENTS 

Part 1 

THE WORK OF THE ACADEMY 

Page 

Officers and Committees for 1969 3 

Minutes of the Spring Meeting 6 

Minutes of the Fall Meeting (Executive Committee) 9 

Minutes of the Fall Meeting (General Session) 12 

Annual Financial Statement 14 

Annual Report, Junior Academy of Science 16 

Biological Survey Committee Report 23 

Necrology 27 

New Members for 1969 39 

Part 2 

ADDRESSES AND CONTRIBUTED PAPERS 

Presidential Address 45 

"The World of the Honeybee", Howard R. Youse 
"To Ruine a World", John B. Patton 49 

Anthropology 

E. Dolan — Use of Historical and Archaeological Evidence to 

Reconstruct the Cherokee Past* 57 

R. E. Pace — Exploratory Examination of the Oxendine Site* 57 

E. V. McMichael and S. J. Coffing — Test Excavations at the 

Daughtery-Monroe Site (12-Su-13) * 57 

R. J. Ferguson — The Continued Excavation of the Van Nuys Site: 

A Probable Late Woodland Occupation* 58 

G. K. Neumann — A Re-examination of the Question of the Middle 

Western Origin of the Delaware Indians 60 

K. B. Hunter and G. K. Neumann — Origins and Racial Affiliations 

of the Illinois Hopewell Indians 62 

W. H. Adams — Some Possible Causes for Late-Pleistocene Faunal 

Extinctions 65 

G. K. Neumann and T. A. Murad — Preliminary Report on the 

Crania from the Island Field Site, Kent County, Delaware .... 69 
K. D. Vickery — Preliminary Report on the Excavation of the 

"Great Mound" at Mounds State Park in Madison County, 

Indiana 75 



: Abstract or Note only 



iv Indiana Academy of Science 

Botany Page 

L. and Adele Beesley — Trilliums of Franklin County, Indiana* .... 83 

K. E. Nichols and W. W. Bloom — Report of a Carotenoidmutant 

of Cyanidium caldariwm* 83 

W. W. Bloom and K. E. Nichols — Responses of Megagametophytes 
of Marsilea to Growth Substances with Respect to Rhizoid 
Formation* 83 

B. O. Blair — Phenology Studies of Ten Species at Eleven Locations 

in Indiana* 83 

L. Ford — Chromosome Associations in Corn Monoploids* 84 

J. Yemma — A Microspectrophotometric Analysis of DNA in the 
Heterothallic Species of Slime Mould, D. iridis, which Some- 
times Exhibits Apogamy* 84 

Fay K. Daily — Some Charophytes from the Pleistocene* 84 

P. Weatherwax — Some "Atypical" Stem Structures in the 

Gramineae 85 



Cell Biology 

J. R. Welser and W. W. Carlton — Response of the Duck Thyroid 

to the Administration of Thiouracil* 91 

J. F. Van Vleet, B. V. Hall and J. Simon — Regeneration of 

Skeletal Muscle in Vitamin E-Deficient Rabbits* 91 

I. Watanabe and G. Bingle — Deficient Myelination in the Brain 

of "Quaking" Mouse: An Electron Microscopic Study* 92 

J. F. Schmedtje — The Identification of Gamma-A Globulin in 

Human Enteric Epithelial Cells* 92 

N. E. Weber and P. V. Blair — Effects of Adenosine Diphosphate 

on the Morphology of Heart Mitochondria* 93 

A. N. Siakotos — The Isolation of Nuclei and Endothelial Cells from 

Brain* 93 

S. Yumoto and S. Ishikawa — Ultrastructural and Enzymological 

Observations of Isolated Kidney Microvilli* 93 

J. B. Whitten — Ultrastructural Features of the Calcifying 

Epithelial Odontogenic Tumor* 94 

D. J. Morre, J. C. Roland and C. A. Lembi — Comparisons of 

Isolated Plasma Membranes from Plant Stems and Rat Liver . . 96 
R. D. Cheetham and D. J. Morre — Di- and Tri-nucleotidase 

Activities of Rat Liver Cytomembranes 107 

T. F. Chuang, Y. C. Awasthi and F. L. Crane — A Model Mosaic 

Membrane: Cytochrome Oxidase 110 



*Abstract or Note only 



Table of Contents v 

Chemistry Page 

L. G. Ong and E. Schwartz — Determination of the Hydrolysis 
Constant of the Stannous Ion by an Electromotive Force 
Method- 121 

J. Nowak, R. Williams and J. George — The Infrared Spectra of 

Coordination Compounds* 121 

R. L. VanEtten and D. S. Page — pH Dependent Isotope Effects 

on the Flavin Enzyme L-Amino Acid Oxidase* 121 

J. T. Snow and G. R. Barker — A Kinetic Study of the Decar- 
boxylation of Duroic Acid in Sulfuric Acid Solutions* 122 

D. J. Cook — Molecular Complexes of Bromine and Various Sub- 
stituted Carbostyrils and their Hydrolysis Products* 122 

Sr. Barbara Buckbee, R. E. Surdzial and C. R. Metz — Temperature 

Dependence of Eo for the Daniell Cell 123 

W. E. Hoffman, M. Jacobs, G. Kennepohl, D. W. Parrott, P. 
Reed, T. R. Stout and J. Sundy — The Study of Complexes of 
Di-n-butyloxamidine with Transition Metals 129 

Ecology 

T. S. McComish and R. 0. Anderson — The Annual Growth Cycle 

of the Bluegill* 135 

P. G. Davidson and T. S. McComish— The Effect of Photoperiod on 

Growth of Bluegill* 135 

J. R. Gammon — The Response of Fish to Heated Effluents* 13G 

T. E. Mangum, III and T. S. McComish — Preliminary Experiments 

on Growth of Bluegill with Varied Feeding Frequency and 

Constant Ration* 136 

Ruth A. Wilsey and T. S. McComish — The Relationship Between 

Growth and Social Hierarchy in the Green Sunfish* 136 

D. R. Goins — A Taxonomic Survey of the Ostracods of Delaware 

County, Indiana* 137 

R. D. Hart and W. B. Crankshaw — The Influence of Environmental 

Factors on the Concentration of Hydrocyanic Acid in Manihot 

esculenta* 137 

M. T. Jackson and R. 0. Petty — A Comparison of Dominance 

Expressions for Tree Species in Foley Woods, Edgar County, 

Illinois* 137 

D. Schmelz — Testing the Quarter Method against Full Tallies in 

Old-growth Forests* 138 

W. J. Gulish — Bluegill Predation by Three Fish Species 139 

H. E. McReynolds — Practicality of Endrin as a Fish Toxicant .... 148 
H. P. Weeks, Jr. — Courtship and Territorial Behavior of Some 

Indiana Woodcocks 162 

G. S. Jones — Foods of the White-footed Mouse, Peromyscus leucopus 

noveboracensis, from Pike County, Indiana 172 



*Abstract or Note only 



vi Indiana Academy of Science 

Page 
J. P. Muehrcke and C. M. Kirkpatrick — Observations on Ecology 

and Behavior of Indiana Ruffed Grouse 177 

R. D. Kirkpatrick — Fox Bounty in Indiana During the Years 1961 

through 1968 187 

M. E. Heath — Naturalized Big Trefoil (Lotus pedunculatus Cav.) 

Ecotypes Discovered in Crawford County, Indiana 193 

A. A. Lindsey and D. V. Schmelz — The Forest Types of Indiana 

and a New Method of Classifying Midwestern Hardwood 

Forests 198 

L. A. Krumholz, R. L. Bingham and E. R. Meyer — A Survey of 

the Commercially Valuable Mussels of the Wabash and White 

Rivers of Indiana 205 

Entomology 

Mildred G. Ware and H. L. Zimmack — The Use of Heartbeat as a 

Potential Screening Technique for Insect Pathogens* 227 

F. N. Young — Further Studies on the Interbreeding of an Insular 
Form of Tropistemus collaris (Castelnau) with Mainland 
Forms* 227 

L. Chandler — The Second Record of Coelioxys obtusiventris Craw- 
ford (Hymenoptera, Megachilidae) . 228 

L. Chandler — Indiana vs. Indian Territory: Misinterpreted Locality 

Citations 229 

Gertrude L. Ward — The occurrence of Chalybion zimmermanni 

Dahlbom (Sphecidae) in Indiana 231 

Patricia M. Arnett — A Taxonomic Key to the Collembola in Four 

Serai Stages Leading to the Beech-Maple Climax 234 

R. E. Siverly — Factors Influencing the Species Composition of 

Mosquito Populations in Indiana 238 

J. W. Hart— A Check List of Indiana Collembola 249 

Geology and Geography 

C. T. Statton — Faunal Assemblage Study of the Maryland 

Miocene* 253 

A. H. Siddiqi — Urbanization in Asia* 253 

R. F. Boneham — Acritarchs (Leiosphaeridia) in the New Albany 

Shale of Southern Indiana 254 

C. E. Wier — Factors Affecting Coal Roof Rock in Sullivan County, 

Indiana 263 

N. I. Johansen and W. N. Melhorn — Proposed Origins for the 

Hadley Lake Depression, Tippecanoe County, Indiana 270 



* Abstract or Note only 



Table of Contents vii 

Page 
R. L. Powell — Base Level, Lithologic and Climatic Controls of 

Karst Groundwater Zones in South-Central Indiana 281 

L. A. Schaal — Cooling Degree Days in Indiana 292 

E. M. Agee — The Climatology of Indiana Tornadoes 299 

R. M. Dinkel and A. J. Cantin — The Changing Location Patterns 

of the Neighborhood Grocers in Terre Haute, Indiana: A 

Geographic Analysis 309 

T. F. Barton — Population and Settlement Decline in Southwestern 

Indiana 318 

N. V. Weber — A Comparison of the Central Place Hierarchy Pattern 

of Central Indiana to the Walter Christaller Model 325 

D. M. Coffman — Parameter Measurement in Fluvial Morphology . . 333 

Microbiology and Molecular Biology 

R. Ramaley and R. Kindig — Stream Pollution from Coal Mine 

Waste Piles: Effect of Sulfur and Iron Oxidizing Bacteria* . . 345 

D. A. Cotter — Fine Structure Changes during Germination of 

Dictyostelium discoideum Spores* 345 

S. S. Lee — Thermal Induction of Bacteriophage PI* 346 

A. R. Schulz and D. D. Fisher — Computer-based Derivation of 

Rate Equations for Enzyme-catalyzed Reactions* 346 

S. Ochs, M. I. Sabri and N. Ranish — Axoplasmic Transport of 

Materials in Nerve Fibers* 346 

T. A. Cole and W. F. Middendorf — Development of a Clear, Photo- 
polymerizable Acrylamide Gel and Its Use in Immobilization 
and Staining of Nucleic Acids 348 

K. Schwenk and Alice S. Bennett — The Effect of Avidin on the 
Biosynthesis of Fatty Acids in Aspergillus niger and Asper- 
gillus flavus 351 

Physics 

W. D. Mueller — Measurement of Ionization in Nuclear Emulsion by 

Lacunarity Technique* 357 

M. A. Burkle and L. M. Reynolds — Operation and Flux Deter- 
mination of a Neutron Generator* 357 

L. K. Steinrauf, 0. Seely, J. A. Hamilton and J. M. H. Pinker- 
ton — Molecular Structure of Thyroxine by X-Ray Crystallog- 
raphy* 357 

J. F. Houlihan and L. N. Mulay — Electron Paramagnetic Reso- 
nance Studies on the Magneli Phases of the Titanium-Oxygen 
System* 358 

D. L. Steinert — An Evaluation of Relativistic Thermodynamics* . . 358 



'Abstract or Note only 



viii Indiana Academy of Science 

Page 

R. A. Llewellyn — Physical Oceanography in Indiana: A Study of 

Horse Shoe Lake* 359 

J. H. Hettmer — Polarized : *He+ Ion Source for the Indiana Uni- 
versity Cyclotron* 359 

U. J. Hansen and J. L. Hazelton — Non-local Contributions to the 
Cyclotron Absorption Spectrum for a Single Valley Semi- 
conductor Model" 360 

C. C. Sartain — Semiconductors Produced by Doping Oxideglasses 

with Ir, Pd, Rh or Ru 361 

Plant Taxonomy 

N. E. Lambert and D. L. Dilcher — Eocene Euphorbiaceous Fruits" 375 
M. V. Sheffy and D. L. Dilcher — Morphology and Taxonomy of 

Fossil Fungal Spores* 375 

C. B. Heiser, Jr. — The Cultivated Solanaceae of Ecuador* 376 

Winona H. Welch — Hookeriaceae Species and Distribution in 

Africa, Europe, Asia, Australia and Oceania 377 

Jeanette C. Oliver — Biosystematic Studies of the Beech and Marsh 

Ferns 388 

D. N. Moore, T. R. Mertens and Joyce E. Highwood — Cytotax- 

onomic Notes on Genus Polygonum, Section Polygonum 396 

Soil Science 

R. K. Stivers — Distribution of Corn (Zea mays L.) Roots in Two 

Soils in Relation to Depth of Sampling and Type of Sampler . . 401 

J. V. Mannering and D. Wiersma — The Effect of Rainfall Energy 

on Water Infiltration into Soils 407 

M. F. Baumgardner, S. Kristof, C. J. Johannsen and A. Zachary 
— Effects of Organic Matter on the Multispectral Properties of 
Soils 413 

L. B. Hughes and H. W. Reuszer — A Two-year Study of Bacterial 

Populations in Indiana Farm Pond Waters 423 

N. L. Meyers, J. L. Ahlrichs and J. L. White — Adsorption of 

Insecticides on Pond Sediments and Watershed Soils 432 

Zoology 

E. Whittle — Some Modifications in Rat Ovaries and Uteri Follow- 

ing Aminoglutethimide Treatment* 439 

J. B. Cope and R. Mills — Big Brown Bat (Eptesicus fuscus) Move- 
ment in Tunnel Cave, Clifty Falls State Park, Indiana* 439 



♦Abstract or Note only 



Table of Contents ix 

Page 

J. O. Whitaker, Jr. — Parasites of Feral Housemice, Mus musculus, 

in Vigo County, Indiana 441 

R. E. Brechner and R. D. Kirkpatrick — Molt in Two Populations 

of the House Mouse, Mus musculus 449 

R. E. Zimmerman and W. J. Eversole — Some Effects of Amino- 
glutethimide on Water and Electrolyte Metabolism in the 
Female Rat 455 

F. J. Zeller and W. R. Rathkamp — The Effect of Steroids on the 
Follicle Stimulating Hormone (FSH) Content of Chicken 
Anterior Pituitary Glands 462 

J. B. Cope, D. R. Hendricks and W. B. Telfair — Radiotelemetry 

with the Big Brown Bat (Eptesicus fuscus) 466 

J. B. Cope and D. R. Hendricks — Status of Myotis lucifugus in 

Indiana 470 

R. L. Richards — Vertebrate Remains from an Indiana Cave 472 

R. E. Schaffer and R. W. Bullard — Comparisons of Rewarming 
from Natural Torpidity and Induced Hypothermia in Chip- 
munks (Tamias striatus) with Reference to Heart Rate and 
Temperature Relationships 476 

Instructions for Contributors 484 

Index 486 



* Abstract or Note only 



PART 1 

THE WORK 

OF THE 
ACADEMY 

1969 



Howard R. Youse, President 



Officers and Committees for 1969 

OFFICERS 



President Howard R. Youse, DePauw University 

President-elect Frank A. Guthrie, Rose Polytechnic Institute 

Secretary James R. Gammon, DePauw University 

Treasurer Damian Schmelz, St. Meinrad College 

Editor William R. Eberly, Manchester College 

Director of Public Relations Paul E. Klinge, Indiana University 

DIVISIONAL CHAIRMEN 

Anthropology Robert E. Pace, Indiana State University 

Botany Robert L. Kent, Indiana Central College 

Cell Biology Edward J. Hinsman, Purdue University 

Chemistry James E. George, DePauw University 

Ecology Thomas S. McComish, Ball State University 

Entomology Jack R. Munsee, Indiana State University 

Geology and Geography. . Wilton N. Melhorn, Purdue University 

History of Science B. Elwood Montgomery, Purdue University 

Microbiology and Molecular Biology D. S. Wegener, 

Indiana University Medical Center 

Physics Richard L. Conklin, Hanover College 

Plant Taxonomy Jack E. Humbles, Indiana University 

Soil Science James E. Newman, Purdue University 

Zoology James C. List, Ball State University 



EXECUTIVE COMMITTEE 

(Past Presidents,* Current Officers, Divisional Chairmen, 
Committee Chairmen) 



*Baldinger, L. H. 

Behrens, 0. K, 

Brooker, R. M. 
*Christy, 0. B. 
*Cleland, R. F. 

Coats, N. 

Conklin, R. L. 

Daily, F. K. 
*Daily, W. A. 
*Day, H. G. 

Eberly, W. R. 
*Edington, W. E. 
*Edwards, P. D. 

Gammon, J. R. 

George, J. E. 
*Girton, R. E. 
*Guard, A. T. 

Guthrie, F. A. 
*Haenisch, E, L. 



Heniser, V. 

Hinsman, E. J. 

Humbles, J. E. 
*Johnson, W. H. 

Kaufman, K. L. 

Kent, R. L. 

Kessel, W. G. 

Klinge, P. E. 
*Lilly, Eli 
*Lindsey, A. A. 

List, J. C. 

McComish, T. 
*Markle, C. A. 

Melhorn, W. M. 
:i: Mellon, M. G. 
*Meyer, A. H. 
*Michaud, H. H. 

Montgomery, B. 



"Morgan, W. P. 

Moulton, B. 

Munsee, J. R. 

Newman, J. E. 

Nisbet, J. 

Pace, R. E. 

Petty, R. O. 
*Powell, H. M. 

Schmelz, D. 

Stockton, Sister M. R. 
*Wayne, W. J. 
*Weatherwax, P. 

Webster, J. D. 

Wegener, D. S. 
*Welch, W. H. 
*Welcher, F. J. 

Winslow, D. 

Youse, H. R. 



4 Indiana Academy of Science 

BUDGET COMMITTEE 
President, H. R. Youse; President-elect, F. A. Guthrie; Secretary, 
J. R. Gammon; Treasurer, D. Schmelz; Editor, W. R. Eberly; Director of 
Public Relations, P. E. Klinge; Retiring President, W. J. Wayne; Di- 
rector of Junior Academy, D. R. Winslow; Library Committee, Nellie 
Coats; Program Committee, R. L. Conklin; Relation of Academy to State, 
W. A. Daily. 

COMMITTEES ELECTED BY THE ACADEMY 
Academy Foundation: W. P. Morgan, 1969, Chairman; W. A. Daily, 1970. 
Bonding: R. M. Brooker, 1969, Chairman; H. H. Michaud, 1969. 
Research Grants: 0. K. Behrens, 1969, Chairman; J. E. Newman, 1973; 
J. B. Patton, 1972; W. K. Stephenson, 1971; W. H. Welch, 1970. 

COMMITTEES APPOINTED BY THE PRESIDENT 

(President an ex officio member of all committees) 

Academy Representative on the Council of A.A.A.S.: W. H. Johnson. 

Auditing Committee: W. G. Kessel, Chairman; L. Guernsey; F. A. 
Guthrie. 

Youth Activities Committee: V. Heniser, Chairman; Sr. Mary Alexandra; 
R. Brooker; J. Colglazier; J. Davis; E. Haenisch; K. Kaufman; W. 
Kessel; G. Kirkman; R. Lefler; R. Settle; D. Winslow. 

Indiana Science Talent Search: V. Heniser, Director; L. H. Baldinger; 
R. L. Henry; A. Kahn; A. R. Schmidt; H. L. Zimmack. 

Indiana Science Fairs State Coordinator: K. L. Kaufman. 

Library Committee: Nellie Coats, Chairman; Lois Burton; J. W. Klotz; 
Eli Lilly. 

Program Committee: R. L. Conklin, Chairman; S. C. Adams; J. D. 
Webster. 

Publications Committee: W. R. Eberly, Chairman; J. A. Clark; D. A. 
Frey; W. N. Melhorn; J. F. Pelton; W. J. Wayne. 

Relation of Academy to State: W. A. Daily, Chairman; J. A. Clark; C. F. 
Dineen; W. R. Eberly. 

Membership Committee: Sr. M. Rose Stockton, Chairman; G. R. Bakker; 
W. L. Burger; G. H. Bick; K. H. Carlson; R. H. Coleman; R. L. 
Conklin; J. P. Danehy; F. K. Edmundson; Olive Forbes; R. E. Hale; 
W. E. Hoffman; W. B. Hopp; J. F. Hayden; W. R. Hurt; E. R. 
Johnston; R. L. Kent; H. P. Leighly; Marie Mayo; J. W. McFarland; 
D. E. Miller; G. R. Miller; E. Nussbaum; P. A. Orpurt; J. B. Patton; 
J. F. Pelton; R. 0. Petty; S. N. Postlethwait; A. F. Schneider; Rev. J. 
Siegrist; L. A. Willig; F. J. Zeller; W. A. Zygmunt. 

Fellows Committee: B. Moulton, Chairman; M. F. Baumgardner; R. L. 
Conklin; S. Crowell; F. K. Daily; H. E. Driver; D. Fraser; C. B. 
Heiser; D. E Miller; B. E. Montgomery; K. M. Seymour; W. H. 
Welch. 



Officers and Committees 5 

Resolutions Committee: J. E. Newman, Chairman; A. T. Guard; J. M. 
Smith. 

Invitations Committee: J. Nisbet, Chairman; W. B. Hopp, J. C. List; W. K. 
Stephenson, J. D. Webster. 

Necrologist: F. K. Daily. 

Parliamentarian: P. Weatherwax. 

SPECIAL COMMITTEES APPOINTED BY THE PRESIDENT 

Biological Survey Committee: J. D. Webster, Chairman; L. Chandler; C. B. 
Heiser; G. C. Marks; R. Mumford, W. H. Welch; F. Young. 

Emeritus Members Committee: R. E. Cleland, Chairman; R. Cooper; E. L. 
Haenisch; H. H. Michaud; W. H. Welch. 

Preservation of Scientific Areas Committee: R. 0. Petty, Chairman; R. C. 
Gutschick; C. H. Krekeler; C. Markle; B. Moulton; D. Schmelz; R. C. 
Weber; W. H. Welch. 

Science and Society Committee: W. Johnson, Chairman; 0. K. Behrens; 
M. Burton; J. E. Christian; R. Cleland; H. Day; W. R. Eberly; P. 
Klinge; H. Kohnke; A. A. Lindsey; R. Miles; R. Rogers; R. Menke; 
H. Wells. 



SPRING MEETING 

Hanover College, Hanover, Indiana 

MINUTES OF THE EXECUTIVE COMMITTEE MEETING 

April 25, 1969 

The meeting was called to order in the Senate Room of the J. Graham 
Brown Campus Center at 4:00 pm., by President Howard R. Youse. 

The minutes of the General Session, Ball State University, October 19, 
1968, were approved as read. 

Treasurer: Rev. Damian Schmelz submitted the financial report for 
the period January 1, 1969, through April 22, 1969. A summary is as 
follows: 

Academy Accounts 

Balance as of January 1, 1969 $ 8,719.46 

Receipts through April 22, 1969 4,147.40 

Expenditures through April 22, 1969 2,855.02 

Balance as of April 22, 1969 10,011.84 

Administered Accounts 

Balance as of January 1, 1969 . $14,817.13 

Receipts through April 22, 1969 12,125.00 

Expenditures through April 22, 1969 10,441.66 

Balance as of April 22, 1969 16,500.47 

Academy Foundation Committee: Mr. W. A. Daily reported that the 
Academy Foundation is in very satisfactory condition. 

Bonding Committee: Dr. R. M. Brooker, Chairman, presented the idea 
that perhaps the amount of the bond should be increased. President Youse 
suggested that the matter be considered at the October Executive 
Committee Meeting, and a recommendation be made at that time. 

Research Grants Committee: Chairman O. K. Behrens reported that 
eight grants-in-aid-of-research had been made since the last Academy 
Meeting, a total of $2,956.40, and that there are funds available for 
additional grants. 

Academy Representative on the Council of A.A.A.S.: Dr. W. H. 

Johnson, our representative, attended the 1968 meeting. 

Auditing Committee: Dr. F. A. Guthrie reported that the committee 
found the financial records in fine condition. 

Youth Activities Committee: Dr. Howard R. Youse stated that the 
Indiana Science Talent Search and Science Fairs have had very good 
results. 

Library Committee: Mrs. Lois Burton reported that there are 330 
copies of "Natural Features of Indiana" available. 

6 



Minutes of the Executive Committee 7 

Program Committee: Chairman R. L. Conklin announced that the Fall 
Meeting will be held at Hanover College, October 24-25, and that the pre- 
liminary notice will be mailed in June. Dr. J. D. Webster has served as 
chairman of the Spring Meeting. 

Publications Committee: President Youse read a letter from Editor 
W. R. Eberly, stating his desire to resign upon the completion of the 1968 
volume of the Proceedings and monograph now in press. Dr. Youse asked 
for suggestions regarding a successor to Dr. Eberly. 

Relation of Academy to State Committee: Chairman W. A. Daily 
announced that the Academy will receive $4,000 per annum for the next 
two years. 

Membership Committee: Chairman Sister Mary Rose Stockton has 
had 2,000 membership blanks prepared and distributed to her committee 
members. Fifteen applications for membership have been received since 
the October 1968 meeting. 

Invitations Committee: Chairman J. Nisbet extended the invitation 
of St. Mary's College for the 1972 Fall Meeting. The invitation was 
accepted with thanks. 

Parliamentarian: Paul Weatherwax read the following: Indiana 
Academy of Science Constitutional Amendment, authorized by Executive 
Committee at Annual Meeting, 1968, and submitted to Executive Com- 
mittee at Spring Meeting, 1969. Art. II, Sec. 8 to be added: 

Sec. 8. Sustaining Members. Any person or corporation making a 
substantial annual contribution to The Academy in an amount to be deter- 
mined by the Executive Committee may be elected to sustaining mem- 
bership. 

This proposal is to be acted upon at the 1969 October Meeting after a 
copy has been distributed to each member by mail, 30 days before the 
Fall Meeting. Dr. Weatherwax made the motion that this Constitutional 
Amendment be distributed to the membership and acted upon at the 1969 
October Meeting. The motion was seconded by Dr. Behrens. Motion 
carried. 

Emeritus Members Committee: Chairman R. E. Cleland presented a 
list of five requests for Emeritus Membership: Dr. Thomas DeVries, Dr. 
Naomi Hougham, Dr. David T. Jones, Mr. Elmer Sulzer, and Mr. William 
A. Wilier. These members of the Academy were voted Emeritus 
Membership. 

Preservation of Scientific Areas Committee: The report is one of 
continuation of activities. 

Science and Society Committee: Chairman Willis H. Johnson gave a 
favorable report on the requests for the Speaker's Bureau and for Science 
in Government, and recommended the continuation of this committee 
because of its benefit to the legislators as well as to the State. Dr. Johnson 



8 Indiana Academy of Science 

announced that the Academy had received a grant of $20,000 for each of 
the next two years for the activities of this committee. 

Dr. R. L. Bernhardt asked for information regarding the formation 
of a new section in Science Education at the October Meeting. Parlia- 
mentarian Weatherwax read instructions regarding procedure from Art. 4 
of the Constitution. 

Dr. J. S. Ingraham presented a request from The Indiana Branch of 
the American Society of Bacteriology regarding the change of its name 
as an Academy Section from Bacteriology to Microbiology and Molecular 
Biology. The Section so voted at the 1968 October Meeting. Dr. Paul 
Weatherwax moved that the request be granted. Dr. Willis Johnson 
seconded the motion. The vote was favorable. 

Adjournment at 5:35 pm. 

Approved October 23, 1969. 

Howard R. Youse, President 
Winona H. Welch, Secretary pro tern. 



FALL MEETING 

MINUTES OF THE EXECUTIVE COMMITTEE MEETING 

Hanover College, Hanover Indiana 

October 23, 1969 

The meeting was called to order at 7:30 p.m. by Dr. Howard R. Youse, 
President of the Academy, in the Student Senate Room of the Campus 
Center. 

The minutes of the Executive Committee and General Session of 
the Spring Meeting of the Academy held at Hanover College on April 
25, 1969, were read and approved. 

Treasurer: Rev. Damian Schmelz reported the Academy funds as 
follows : 

January 1, 1969 balance $23,536.59 

Income to October 17, 1969 26,126.28 

Expended to October 17, 1969 17,738.06 

Balance, October 17, 1969 31,924.81 

Trustees of the Academy Foundation: William A. Daily reported 
Academy Foundation Funds as follows: 

October 1, 1968 balance $ 1,050.20 

Receipts through September 30, 1969 725.32 

Disbursements through Sept. 30, 1969 1,302.46 

Balance, September 30, 1969 473.06 

In the John S. Wright Fund: 

Balance, October 1, 1968 $ 1,466.33 

Receipts to September 30, 1969 11,351.42 

Disbursements to Sept. 30, 1969 12,605.72 

Balance as of September 30, 1969 212.03 

Bonding Committee: Dr. R. M. Brooker, Chairman, stated that there 
seemed to be no need to increase the amount of the bond at this time. 

Research Grants Committee: Dr. O. K. Behrens reported the approval 
of nine grants thus far this year totalling $3,456.30. 

Publications Committee: Dr. W. R. Eberly reported that the Proceed- 
ings are in the final steps of publication. The number of published papers 
has been increasing each year and that a system of review and selection 
may soon become necessary. Dr. Eberly announced the publication of 
Monograph No. 1, "Distribution of the Mammals of Indiana," by Russell 
Mumford. Dr. Howard Youse suggested that this publication be mailed 
to Academy members without cost. Dr. Weatherwax moved the adoption 
of the suggestion and the motion was seconded and approved. 

Youth Activities Committee: Mr. Gil Turpin reported that Dr. 
Wendell McBurney has assumed the duties of Dr. Virgil Heniser who 

9 



10 Indiana Academy of Science 

has retired after many fruitful years with the Youth Activities Com- 
mittee. He also discussed briefly the activities planned for the Annual 
Meeting- of the Indiana Junior Academy of Science which will be held 
November 1, 1969, at Purdue University. The Indiana Science Talent 
Search was reported to have had a busy and successful year. 

Library Committee: Miss Nellie M. Coats reported several major 
acquisitions purchased from a Lilly Endowment Fund. The process of 
listing new library titles for the National Union List of Serials was 
continued and was listed in a Serials Data Bank being compiled by 
computer at Purdue University. Increased use of the Library's materials 
was noted for 1969. 

Relation of the Academy to the State: William A. Daily stated that 
.00 will again be provided by the State. 



Membership Committee: Sister M. Rose suggested that the applica- 
tions of new members be forwarded to her after processing by the 
Treasurer and Secretary so that the appropriate member of the Com- 
mittee can be notified. 

Fellows Committee: Dr. B. Moulton recommended the following 
members for Fellows of the Academy: 

Robert I. Fletcher 

John A. Leighty 

A motion was approved to accept these members as Fellows of the 
Academy. 

Invitations Committee: Dr. Jerry J. Nisbet reported the schedule of 
fall meetings include 1970— Indiana State University; 1971— Earlham 
College; 1972— St. Mary's College; 1973— open. 

Emeritus Membership Committee: Dr. Ralph E. Cleland recom- 
mended four members for Emeritus status: 

May S. Iske 
Harry R. Mathias 
Elmer Sulzer 
Darl F. Wood 

A motion was approved to accept these members as emeritus members of 
the Academy. 

Scientific and Natural Areas Committee: Dr. Robert Petty reported 
that 251 areas are on the list of natural areas. Information concerning 
their location was sent to several companies, institutions, other organiza- 
tions and individuals. An evaluation of specific threatened areas were 
offered to the Audubon Society, the Izaak Walton League and the Indiana 
Conservation Council, Inc., this year. 

Committee on Science and Society: Dr. W. H. Johnson reported that 
representatives of the Committee have held two meetings with the Gover- 



Minutes of the Executive Committee 11 

nor and his Administrative Assistant, were received cordially and were 
promised that different agencies of government would call on the Com- 
mittee for assistance. Dr. John Leighty is now serving as Executive 
Director for the work of the committee under the NSF grant. Favorable 
publicity was received in news media through Dr. Leighty's introduction 
in his new capacity at the second meeting in the Governor's office. Dr. 
Leighty opened an office in Columbus, Indiana, in September at 641 Wash- 
ington Street, Room 5. The Public Service Commission requested as- 
sistance in connection with problems of pipeline safety and were pro- 
vided with a list of consultants. Dr. Leighty has already attended a 
meeting planning a midwest conference on science and technology in 
relation to state and local government and another meeting with the 
State Drug Education Committee which had asked for information on 
the work of the committee for inclusion in their newsletter. 

The disposition of several inactive Academy Sections was discussed 
and a motion recommending the placing of Sections Mathematics and 
Psychology on inactive status was approved. 

The formation of a new section of Science Education was also dis- 
cussed. Mr. Virgil Imel reported a new group, "Hoosier Association of 
Science Teachers, Inc." which is meeting on October 25, 1969, and is 
for elementary, junior high school and high school science teachers. Some 
discussion ensued about the need for a duplicate, competing Section in 
Science Education in the Academy of Science. Dr. Eberly suggested the 
formation of an ad hoc committee which would consider ways in which 
the Academy might become associated with science teaching at these 
levels. Following a suggestion by Dr. Youse, Dr. Brooker moved the 
provisional establishment of a Section in Science Education for a trial 
one-year period. This was seconded and approved. 

Dr. Howard R. Youse recommended that serious consideration be 
given to establishing a permanent position of Executive Director who 
would carry out much of the continuing work of the Academy. 

The meeting was adjourned at 9:20 pm. 

Approved October 24, 1969. 

James R. Gammon, Secretary 



MINUTES OF THE GENERAL SESSION 

October 24, 1969 

The General Session of the Eighty-fifth Fall Meeting of the Indiana 
Academy of Science was held in Parker Auditorium on Friday, October 
24, 1969, at 11:00 am. Dr. Howard R. Youse, President, called the meet- 
ing to order. 

Dr. Harold J. Haverkamp, Academic Dean and Acting President of 
Hanover College, bid the Academy members welcome. 

Dr. Richard S. Westfall, Department of the History and Philosophy 
of Science, Indiana University, delivered a scholarly and interesting ad- 
dress entitled "Problems of Biological Thought in the 17th Century." 

The minutes of the Executive Committee of Thursday, October 23, 
1969, were read and approved as read. 

President Howard R. Youse presented the following Constitutional 
Amendment to the Academy Membership for action. The Amendment had 
been approved at the 1969 Spring Meeting by the Executive Committee 
and circulated to the members more than 30 days prior to the Fall Meet- 
ing. 

Article II, Section 8 to be added: 

Section 8. Sustaining Members. Any person or corporation making a 
substantial annual contribution to The Academy in an amount 
to be determined by the Executive Committee may be elected 
to sustaining membership. 

This amendment was approved. 

President Howard R. Youse then introduced Dr. John Leighty, the 
new Executive Director of the Committee on Science and Society, who 
discussed some of the activities planned for the near future. 

Fay Kenoyer Daily read a biographical sketch of each member who 
had died since the 1968 Fall Meeting. These are printed under Necrology. 

The Annual Dinner Meeting was held in the Dining Hall, J. Graham 
Brown Campus Center, Dr. Frank A. Guthrie, President-Elect presiding. 
Dr. Arthur T. Guard, Resolutions Committee, submitted the following 
resolution: "That the Academy members here assembled express their 
appreciation to Hanover College for all the courtesies which have been 
extended to the membership of the Academy during this meeting. We 
are indebted especially to Dr. Richard L. Conklin, Chairman of the 
Program Committee, for his efforts in arranging facilities for this an- 
nual meeting. Further, the Academy is appreciative of the warm wel- 
come extended by Dr. Harold J. Haverkamp, Academic Dean and Acting 
President, Hanover College." The resolution was approved. 

12 



Minutes of the General Session 13 

The Secretary presented 65 applications for membership to the 
Academy. A motion was approved accepting these applicants as mem- 
bers. 

Dr. Alton A. Lindsey, Nominating Committee, presented the follow- 
ing slate of officers and elected committees: President, Frank A. Guthrie, 
Rose Polytechnic Institute; President-Elect, Samuel N. Postlethwaite, 
Purdue University; Secretary, J. Dan Webster, Hanover College; Editor, 
Marion T. Jackson, Indiana State University; (Treasurer, Damian 
Schmelz, St. Meinrad College, and Director of Public Relations, Paul E. 
Klinge, Indiana University continue in office for two more years); Bond- 
ing Committee, Robert M. Brooker, Indiana Central College and Howard 
H. Michaud, Purdue University (both 1970); Research Grants Committee, 
Nelson Easton, Eli Lilly & Co. (1974); Academy Foundation Trustee, 
W. P. Morgan, Indiana Central College (1971). A motion was made to 
accept the officers and committee members and was approved unanimously. 

The meeting was concluded by an informative and thought-pro- 
voking talk delivered by Dr. Howard R. Youse entitled "The World of 
the Honey Bee." 

The meeting was adjourned at 9:00 pm. 

Approved May, 1970. 

James R. Gammon, Secretary 



FINANCIAL REPORT OF THE INDIANA ACADEMY OF SCIENCE 
January 1-December 31, 1969 

I. ACADEMY ACCOUNTS 

Income Expenditure Budgeted 

Dues $ 3,906.00 

Reprints 1,847.10 $ 1,828.73 $ 150.00 

Publication 476.17 

Editor $ 400.00 400.00 

Mailing 76.17 100.00 

Secretary 360.55 300.00 

Clerical 354.28 

Postage, etc. 6.27 

Treasurer 222.23 225.00 

Clerical 61.25 

Postage, etc. 160.98 

Office 120.56 175.00 

Travel, Dues 170.10 180.00 

President's Fund 100.00 

Membership Committee 18.00 75.00 

Junior Academy 22.18 150.00 

Program Committee 472.43 650.00 

Chairman 66.08 

Printing 212.09 

Mailing 194.26 

Transfer to Administered 

Accounts: 2,100.00 

Science and Soc. 600.00 600.00 

Library Binding 1,000.00 1,000.00 

Publication 500.00 500.00 

Transfer from Administered 

Accounts: 22.03 

Interest on Savings 1,070.70 

Miscellaneous 43.05 43.05 

$ 6,888.88 $ 5,834.00 $ 4,605.00 



14 



Financial Report 



15 



II. ADMINISTERED ACCOUNTS 



Jan. 1 
Balance 

Sciene Talent $ 1,995.80 

Science Fair 7,515.99 

Science and Society 466.42 

Research 307.67 

J. S. Wright Library 134.28 

Lilly III Library 4,374.94 

Publication 

Proceedings 

Natural Features 

Academy Budget 

Library Binding 

NSF Funded Project: 

Science to People 

Transfer to Academy 

Account from Miscellaneous 22.03 





1969 




1969 


Expen- 


Dec. 31 


Income 


diture 


Bal. 




$ 1,860.45 


$ 135.35 


$10,730.00 


5,545.99 


12,700.00 


600.00* 


554.53 


511.89 


6,659.35 


6,533.50 


433.52 
134.28 


34.88** 


1,487.00 


2,922.82 
1,029.50 


217.00* 






312.50* 






500.00* 






1,000.00* 




1,000.00 


11,663.33 


4,492.61 


7,170.72 



22.03 



$14,817.13 $31,717.06 $20,496.11 



$26,038.08 



♦Transfer from Academy Accounts 
** Payment cancelled on check 



III. SUMMARY 

Academy Administered Total 

1969 Income $ 6,888.88 $31,717.06 $38,605.94 

1969 Expenditure 5,834.00 20,496.11 26,330.11 

Net Gain 1969 1,054.88 11,220.95 12,275.83 

1968 Balance 8,719.46 14,817.13 23,536.59 

December 31 Balance 9,774.34 26,038.08 35,812.42 

BANK BALANCES 

Terre Haute First National Bank, Terre Haute, Ind. $15,087.86 

Equitable Savings & Loan, Los Angeles, California 5,392.10 

First Western Savings & Loan, Las Vegas, Nevada 15,332.46 

Total Assets in All Accounts $35,812.42 

Rev. Damian Schmelz, Treasurer 
December 31, 1969 



January 30, 1970 

We the undersigned have audited the Treasurer's records for the Indiana Academy 
of Science for the year 1969 and have found them to be accurate and in order. 

W. G. Kessel 
J. L. Guernsey 



INDIANA JUNIOR ACADEMY OF SCIENCE 
OFFICERS 

President: Dennis Waltke, Division of University Schools, Bloomington, 
Indiana 

Vice-President: Timothy O'Leary, Brebeuf Preparatory School, Indian- 
apolis, Indiana 

Secretary: Rachel Koontz, New Haven Senior High School, New Haven, 
Indiana 

JUNIOR ACADEMY COUNCIL 
Prof. Howard Michaud, Honorary Chairman, Purdue University 
Mr, F. Ray Saxman, Cascade High School, Clayton 
Mr. Charles Souers, Division of University Schools, Bloomington 
Miss Helen Reed, Manual High School, Indianapolis 
Mr. David Blase, Arlington High School, Indianapolis 
Dr. Mary J. Pettersen, Morton High School, Hammond 

STATE DIRECTOR 

Prof. Donald R. Winslow, Division of University Schools, Bloomington, 
Indiana 47401 



PROGRAM 

Thirty-Seventh Annual Meeting 
Saturday, November 1, 1969 

8:00 am-10:00 am 

Registration of members, sponsors, and guests — East Foyer registra- 
tion counter, Memorial Center. 

9:30 am 

Tours of several academic facilities. Check registration area. 

9:30 am 

INTERVIEWS for "Best Boy" and "Best Girl" Awards. Memorial 
Center — Room 111. 

11:30 am-1:00 pm 

Lunch — Purdue Union Cafeterias. 

1:00 pm 

Academy General Session. Room 302 of the Memorial Center. Dennis 
Waltke, presiding. 

16 



Junior Academy of Science 17 

1:30 pm 

Presentation of papers. Rooms 302 and 306, Memorial Center. 

3:00 PM 

Closing Session. Room 302, Memorial Center. Announcements; pres- 
entation of awards. 



PROGRAM OF PAPERS 

1. The Origin of the Universe. 

John J. Farrell, Brebeuf Science Club, Brebeuf Preparatory School, 
Indianapolis. 

2. Morphogenesis of the Chick Embryo Lung under the Influence of 
Nicotine. 

Mary Jane Oliver, Kennedy Research Center, Roncalli High School, 
Indianapolis. 

3. Physiological Factors Involved in Acid-Base Balance Disturbances. 
Barbara Konkle, Madison Consolidated High School Science Club, 
Madison. 

4. Infrared Study of the Environment of Exchangeable Ammonium Ions 
on Clay Surfaces. 

Ken Morse, Chemistry Club, Oliver P. Morton Senior High School, 
Hammond. 

5. The Use of Matrices and Determinants in the Balancing of Redox 
Equations. 

Janet Marshall, Phi Chi Science Club, New Haven High School, 
New Haven. 

6. The Characterization of RE Virus. 

Mark Coates, Madison Consolidated High School, Madison. 

7. Comparative Analysis of Commercial Dentrifices Containing Fluoride 
Compounds. 

W. Craig Lannin, Oliver P. Morton High School, Hammond. 

8. Reduction of Cancerous Tumors through Helianthemum annum Ex- 
tractions. 

Joan Field, Kennedy Research Center, Roncalli High School, 
Indianapolis. 

9. Drugs Cause Damage. 

Steve Marietta, Pius X Science Club, Schulte High School, Terre 
Haute. 

10. Experiments in Artificial Intelligence Using a Digital Computer. 
Kevin McGill, Brebeuf Science Club, Brebeuf Preparatory School, 
Indianapolis. 



18 Indiana Academy of Science 



ACKNOWLEDGMENT 



The officers, members of the Junior Academy Council, and all others 
concerned with the program wish to thank Prof. Ralph Lefler of the 
Purdue Physics Department, and Mr. Mark Ocker of the Purdue Division 
of Conferences and Continuation Services for arranging the fine tours 
and meeting facilities for our convention. We are indeed indebted to 
these members of the Purdue campus for so generously giving of their 
services. We also are indebted to the Purdue President's Office for making 
this meeting financially possible. 



MINUTES OF THE THIRTY-SEVENTH ANNUAL 

MEETING OF THE 

INDIANA JUNIOR ACADEMY OF SCIENCE 

For the first time, in order to meet in a central location, the Junior 
Academy met separately from the Senior Academy. The thirty-seventh 
annual meeting of the Indiana Junior Academy of Science was held 
Saturday, November 1, 1969, on the Purdue Campus at Lafayette, Indi- 
ana. There were 274 students and sponsors present, representing 51 
chapters. 

President Dennis Waltke called the meeting to order at 1:00 pm. 
in Room 302 in the Purdue Student Union Center. Previous to this 
session members and their sponsors enjoyed tours and lectures in 
various parts of the campus. 

Following a brief welcome, Dennis introduced the 1969 officers as 
following: 

Dennis Waltke President 

Timothy O'Leary Vice-President 

Rachel Koontz Secretary 

Next Mr. Charles Souers of the Junior Academy Council explained 
and presented the awards for 1970. Officers for 1970 are as following: 

Rachel Koontz President 

Mark Coates Vice-President 

John Farrell Secretary 

The 1969-70 "Best Scientist" awards were presented by Mr. Souers 
and they went to Rachel Koontz, "Best Girl Scientist" and to Steve 
Marietta, "Best Boy Scientist." Each received a certificate and a year's 
membership in the American Association for the Advancement of 
Science. 



Junior Academy of Science 19 

Dennis then broke the group into two parts for presentation of 
papers. Vice-President Timothy O'Leary presided over the second group 
in Room 306. Each person was then allowed seven minutes in which to 
present his paper. Nine papers were given. 

Following the presentation of papers the entire group met in Room 
306, and Dr. Mary Jane Pettersen presented certificates to all those who 
presented papers. Dennis then introduced Mr. Winslow, the State Di- 
rector, who introduced the new Council members for 1970 and extended 
the Academy's thanks for the outstanding co-operation provided by 
Purdue University. Particular acknowledgment was extended to Prof. 
Ralph Lefler who made arrangements for the excellent tours. 

Following a short closing message President Dennis Waltke 
adjourned the meeting at 3:15 PM. 

Next year's meeting is to be held with the Senior Academy at 
Indiana State University, Terre Haute. 

Respectfully submitted, 

Rachel Koontz, Secretary 
Dennis Waltke, President 



INDIANA JUNIOR ACADEMY OF SCIENCE 
1969-70 

Town Club and School Sponsor 

Acton Sigma Mu Chapter of FSA, Frank- Margaret Richwine 

lin Central H. S. 

Bedford Bedford Science Problems Research Paul Hardwick 

Group, Bedford H. S. 

Bloomington National Scientific Honor Society, Orville Long 
Bloomington H. S. 

Bloomington E. Wayne Gross Academy, Univer- Billie Stucky 
sity H. S. 

Bloomington MSE Academy, University Junior Charles Souers 
High 

Clarksville Clarksville H. S. Science Club, Gerald K. Sprinkle 
Clarksville Junior-Senior H. S. 

Clarksville Phy-Chem, Our Lady of Providence Sr. Jean Marian 
H. S. 

Columbus Science Club, Columbus Senior L. N. Carmichael 

H. S. 

Crawfords- Up-N-Atom, Crawfordsville H. S. David Wells 

ville 

Evansville Reitz Memorial Chapter of FSA, Charles Hames 
Reitz Memorial H. S. 

Fort Wayne Albertus Magnus Science Club, Sr. Winifred 
Central Catholic H. S. 

Fort Wayne Phy-Chem Club, Elmhurst H. S. Ruth Wimmer 

French Lick Springs Valley Science Club, D. L. Clark 

Springs Valley H. S. 

Gary Andrean Biology Club, Andrean Sr. Marie Antoine 

H. S. SS.C.M. 

Sr. Marie Carmel 
SS.C.M. 

Gary Mu Alpha Theta, Andrean H. S. Sr. M. Nadine 

SS.C.M. 

Gary Biology Club, Lew Wallace H. S. Lola Lemon 

20 



Junior Academy of Science 21 

Town Club and School Sponsor 

Griffith Griffith Junior High Science Club Fred Meeker 

Griffith Junior H. S. 

Griffith Griffith Junior High Science Club, Geraldine R. Sherfey 

Griffith Senior H. S. 

Hammond Chemistry Club, Oliver P. Morton Mary J. Pettersen 

H. S. 

Hartford City Hartford City H. S. Science Club, 

Hartford City H. S. 

Highland Science Club, Highland H. S. Jon Hendrix 

Hobart Hobart Senior High Science Club, Stanley J. Senderak 

Hobart Senior H. S. 

Huntington Aristotelian, Huntington Catholic Sr. M. Petrona 
H. S. 

Huntington Science, Huntington H. S. Robert Diffenbaugh 

Indianapolis Arlington Science Club, Arlington Robert McClary 
H. S. 

Indianapolis Nature Club, Arsenal Technical Michael Simmons 
H. S. 

Indianapolis Brebeuf Science Club, Brebeuf Pre- Donald G. Maines 
paratory School Harold J, Sommer 

Indianapolis Science Club, Howe H. S. Jerry Motley 

Indianapolis Mendelian Science Club, Ladywood Sr. Helen Jean 
H. S. 

Indianapolis North Central H. S. Science Club, Robert Prettyman 
North Central H. S. 

Indianapolis Roncalli H. S. Sr. Mary Alexandra 

C.S.J. 

Indianapolis Science Club of Westlane, West- John Van Sickle 
lane Junior H. S. 

Indianapolis Science Club, George Washington William Baldwin 
H. S. 

Jamestown Science Club of Granville Wells, Cecil O. Bennington 
Granville Wells School 

LaPorte Bi-Phi-Chem Club, LaPorte H. S. Frances M. Gourley 

Byron Bernard 

Lebanon Junior Explorers of Science, Leba- Tom Evving 

non Junior H. S. 



22 



Indiana Academy of Science 



Town Club and School 

Logansport Lewis Cass H. S. Science Club, 
Lewis Cass H. S. 

Madison Madison Science Club, Madison 

Consolidated High 

Muncie Muncie Central Science Club, Mun- 

cie Central H. S. 

New Albany Science Club, New Albany Senior 
H. S. 

New Haven New Haven Science Club, New 
Haven H. S. 

Portland Science Club, Portland- Wayne 

Township Junior H. S. 

Portland Portland Senior H. S. Science and 

Mathematics Club, Portland Sr. 
H. S. 

South Bend Junior Izaak Walton League, John 
Adams H. S. 

South Bend JETS Junior Engineering Technical 
Society, Central H. S. 

South Bend Second Year Biology Class, Clay 
H. S. 

South Bend IONS Club, J. W. Riley H. S. 

South Bend LaSalle High School 

Terre Haute Pius X Science Teens. Schulte H. S. 



Tipton Tipton H. S. Science Club, Tipton 

H. S. 

Trafalgar Indian Creek School 

Vincennes Sigma Tau Science Club, St. Rose 

Academy 



Sponsor 
Raymond T. Kozer 

David Dunkerton 

William Beuoy 

Roger Moody 



Keith Hunnings 
E. H. Sanders 

Mary Zehner 



Ralph Settle 
Robert Freemyer 

Ernest Litweiler 



John V. Davis 
John Marker 



Sr. Marie Barbara 
S.P. 

Richard Garst 
Frederick Calhoun 



Sr. Anna Margaret 
Sr. Aloyse 



BIOLOGICAL SURVEY COMMITTEE 

J. Dan Webster, Chairman, Hanover College 

Publications of 1968-1969 
Dealing- with the Flora and Fauna of Indiana 



Virales : 



Whitaker, J. O., JR., W. A. Miller and W. L. Boyko. 1969. 
Rabies in Indiana Bats. Proc. Indiana Acad. Sci. 78 : 447-456. 



Tracheophyta 



Beesley, Adelle, and L. Beesley. 1970. Trilliums of Frank- 
lin County, Proc. Indiana Acad. Sci. 79 :83. 

Heath, M. E. 1970. Naturalized big trefoil {Lotus peduncula- 
tus Cav. ) ecotypes discovered in Crawford County. Proc. In- 
diana Acad. Sci. 79:193-197. 

Jackson, M. T. 1969. Hemmer Woods: an outstanding old- 
growth lowland forest remnant in Gibson County, Indiana. 
Proc. Indiana Acad. Sci. 78: 245-254. 

Jackson, M.T., and P. R. Allen. 1969. Detailed studies of 
old-growth forests in Versailles State Park, Indiana. Proc. 
Indiana Acad. Sci. 78:210-230. 

Lindsey, A. A. and D. V. Schmelz. 1970. The forest types of 
Indiana and a new method of classifying midwestern hard- 
wood forests. Proc. Indiana Acad. Sci. 79 :198-204. 

Moore, D., T. Mertens and Joyce Highwood. 1970. Cytotaxo- 
nomic notes on the genus Polygonum, Section Polygonum. 
Proc. Indiana Acad. Sci. 79 :396-400. 

Oliver, Jeanette C. 1970. Biosystematic studies of the Beech 
and Marsh ferns. Proc. Indiana Acad. Sci. 79 :388-395. 



Protozoa : 



Worms and Arthropods 



Crustacea : 



Insecta : 



Tamar, H. 1968. Observations on Halteria bifurcata sp. n. and 
Halteria grandinella. Acta Protozoologica. 6:175-184. 

Whitaker, J. O., Jr. 1970. Parasites of feral housemice, Mus 
muaculua, in Vigo County. Proc. Indiana Acad. Sci. 79:441- 

448. 

Coins, D. R. 1970. A taxonomic survey of the Ostracods of 
Delaware County, Indiana. Proc. Indiana Acad. Sci. 79:137. 

Arnett, Patricia M. 1970. A taxonomic key to the Collem- 
bola in four serai stages leading to the Beech-Maple climax. 
Proc. Indiana Acad. Sci. 79 :234-237. 

Arnett, R. H., E. C. Mignot and E. H. Smith. 1969. North 
American Coleoptera fauna: notes on Pyrophorinae, Elate- 
ridae (Coleoptera), Coleoptera Bull., 23:9-15. 

Chandler, L. 1970. Indiana vs. Indian Territory: Misinter- 
preted locality citations. Proc. Indiana Acad. Sci. 79 :229- 
230. 

Chandler, L. 1970. The second record of Coelioxya obtusiven- 
tria Crawford (Hymenoptera, Megachilidae). Proc. Indiana 
Acad. Sci. 79:228. 

Hart, J. W. 1970. A checklist of Indiana Collembola. Proc. 
Indiana Acad. Sci. 79 :249-252. 

Siverly, R. E. 1970. Factors influencing the species compo- 
sition of mosquito populations in Indiana. Proc. Indiana 
Acad. Sci. 79 :238-248. 

Ward, Gertrude L. 1970. The occurrence of Chalybion zim- 
mermanni Dalbon (Sphecidae) in Indiana. Proc. Indiana 
Acad. Sci. 79:231-233. 

23 



24 Indiana Academy of Science 

Acarina: Whitaker, J. O., Jr., and N. Wilson. 1968. Mites of the 

small mammals of Vigo County, Indiana. Amer. Midland 
Natur. 80:537-542. 

Pisces: Zimmerman, C. J. In press. Ecology of the brook silverside 

(Labidesthes sicculus) in Lake Lemon, Stephens Creek, and 
Monroe Reservoir, Indiana. Trans. Amer. Fish. Soc. 

Aves : Baker, Mrs. H. A. Breeding bird census — grazed, brushy fields 

and tree-bordered creek. Audubon Field Notes 22:701. 

GUTH, R. W. Winter bird population study — shrubby field. 
Audubon Field Notes 23:550-551. 

Guy, R. E. Winter bird population study — shrubby field. 
Audubon Field Notes 23 :541-542. 

Guy, R. E. Winter bird population study — corn stubble. Audu- 
bon Field Notes 23 :549-550. 

Guy, R. E. Winter bird population study — pasture. Audubon 
Field Notes 23:550. 

Indiana Audubon Society Members. 1969. Many titles in Indi- 
ana Audubon Quarterly 46 :1-152. 

Smith, Shelia L. Breeding bird census — suburban edge. Audu- 
bon Field Notes 22 :721. 

Webster, J. D. Winter bird population study — soybean stubble. 
Audubon Field Notes 23 :550. 

Mammalia : Cope, J. B. and D. Hendricks. 1970. Status of Myotis luci- 

fugus in Indiana. Proc. Indiana Acad. Sci. 79:470-471. 
Mumford, R. E. 1969. Distribution of the mammals of Indiana. 

Indiana Acad. Sci. Monograph No. 1. Indiana Acad. Sci., 

Indianapolis. 114 p. 
Mumford, R. E. 1969. Long-tailed weasel preys on big brown 

bats. J. Mammal. 50:360. 
Whitaker, J. O., Jr., and G. R. Sly. 1970. First record of 

Reithrodontornys megalotis in Indiana. J. Mammal. 51:381. 

Aquatic animals: GAMMON, J. R. 1968. Aquatic life survey of the Wabash River, 

with special reference to the effects of thermal effluents on 
populations of macroinvertebrates and fish. Second year 
progress report to Public Service Indiana. 48 p. 
Gammon, J. R. 1968. The effect of inorganic sediment on 
stream biota. Second year progress report to FWPCA. 84 p. 

All organisms: Lindsey, A. A., D. V. Schmelz and S. A. Nichols. 1969. 

Natural areas in Indiana and their preservation. Indiana 
Natural Areas Survey, Lafayette, Indiana. 594 p. 



Theses Completed and Placed on File Dealing with the Flora 
and Fauna of Indiana 

Arachnida: Parker, T. A. 1969. Spiders of the Wabash-Tippecanoe river 

system. Ph.D. Purdue. 

Insecta: Bell, Susan C. 1969. The Effects of Thermal Pollution on the 

Macroinvertebrate Population of the Wabash River. M.A. 
DePauw. 
FlNNl, G. R. 1969. Developmental patterns in winter stonefly 
species (Insecta: Plecoptera: Allocapnia: Taeniopterys) . M. 
Sc. Purdue. 
Lawrence, V. M. 1968. The composition and dynamics of 
Odonata populations in farm pond ecosystems. Ph.D. Purdue. 



Biological Survey Committee 



•if. 



Pisces 



Mignot, E. C. 1969. Taxonomic revision of the tribe Aspicelini 
and Disonychini ( Coleoptera, Chrysomelidae, Alticinae) 
north of Mexico. Ph.D. Purdue. 

Smith, E. H. 1969. Taxonomic revision of the genus Systena 
Chev. (Coleoptera: Chrysomelidae, Alticinae) north of Mex- 
ico. M.Sc. Purdue. 

Young, R. M. 1969. Polyphylla Harris in America north of 
Mexico. Part II: the Hammondi, Decimlineata, and Occi- 
dentalis complexes, and additions to the Diffracta complex 
(Coleoptera: Scarabaeidae, Melolonthinae) . Ph.D. Purdue. 

Zimmerman, C. J. 1969. Fish populations of Stephens and 
Brummett Creeks, Monroe County, Indiana: A Comparative 
Study. M.A. Indiana U. 



Work in Progress, but not yet Published, Dealing with the 
Flora and Fauna of Indiana 



Algae : 



Brown, R. D. Indiana State. Variations in the species mor- 
phology of certain rheophilic diatoms. 



Reptilia and 
Amphibia : 
Mammalia : 



Rubin, D. Indiana State. Continued studies on the amphibians 
and reptiles of Vigo County. 

Whitaker, J. O., Jr., Indiana State. Continued studies on the 

mammals of Vigo County. 
Tuszynski, R. C. Purdue. Ecology of the Pocket Gopher 

(Geomys bursarius) in Indiana. 



NECROLOGY 

Fay Kenoyer Daily, Butler University 

Merrill T. Carr 
West Terre Haute, Indiana Terre Haute, Indiana 

March 6, 1912 November 7, 1969 

Mr. Merrill T. Carr was born in West Terre Haute, Indiana, and his 
early education was obtained in that vicinity. He also received the B.S. 
and M.S. degrees at Indiana State University. His professional career 
included teaching at Fayette, Prairie Creek and West Terre Haute public 
schools. He was a science teacher at Gerstmeyer High School, Terre 
Haute. He was very effective in his teaching and was regarded very 
highly by former students. Several Science Fair participants were spon- 
sored by Mr. Carr. They were regional winners against the stiff compe- 
tition which they encountered. Mr. Carr took part in a research partici- 
pation program at Indiana University for several years. Despite the 
handicap of heart trouble for the last twenty to thirty years, Mr. Carr's 
career was very successful. 

Rebecca Carr, his daughter, joined the Academy in 1961 and was a 
member until her marriage and subsequent change of address to Florida 
(1967). Merrill T. Carr joined the Academy in 1967 and was looking 
forward at the Hanover College Spring Academy meeting in 1969 to 
becoming better acquainted and to pleasant associations at our meetings. 
His pleasant, quiet charm would have assured this if it had not been 
for his untimely death, November 7, 1969. 

Howard O(wen) Deay 
Eudora, Kansas Lafayette, Indiana 

March 5, 1896 June 30, 1969 

Dr. Howard 0. Deay, Professor Emeritus of Entomology at Purdue 
University died June 30, 1969, at Lafayette, Indiana, after a rewarding 
career. 

He was born in Eudora, Kansas, on March 5, 1896, and was educated 
in that state. He started his professional career by teaching in the Eudora 
City Schools in 1917. He was then a private in the United States Army 
1918 to 1919, but returned to Eudora as a school principal from 1921 to 
1924. 

His college education at the University of Kansas began by com- 
pletion of some correspondence courses, so when he entered the school 

27 



28 Indiana Academy of Science 

in 1924, it was as a sophomore. He received an A.B. degree in 1926 and 
became a Graduate Fellow in Entomology. He received an M.A. degree 
in 1927 continuing as a Graduate Fellow until 1928 when he was made 
an Instructor. During his student years, summer employment included 
being supervisor of European corn borer scouts for the U. S. Department 
of Agriculture in 1926 and 1927 and nursery inspector for the State of 
Kansas in 1925 and 1928. 

Dr. Deay came to Indiana in 1929 to teach Entomology at Purdue 
University continuing his graduate research at the University of Kansas 
in absentia. He would work for the State Entomologist's office in Indi- 
ana one summer, then return to Lawrence, Kansas, the next summer to 
continue his research until 1934 when he received his Ph.D. degree. Our 
former state entomologist, Frank Wallace, described him as "the best 
entomologist in the world." 

Dr. Deay was a very successful teacher at Purdue becoming full 
professor in 1950. Besides this, he counseled and registered all students 
in entomology, maintained the departmental library, conducted and di- 
rected research — doing all of these things well. In addition to regular 
classes, he also taught operators when the pest control industry held 
its conferences at Purdue. He maintained a continuing interest in the 
many business men who attended, some of whom returned annually for 
a refresher course in insect identification. He retired as Professor Emeri- 
tus in 1964. 

His early research interest was in systematics of insects. Some of 
the papers on this subject presented before the Indiana Academy of 
Science included: Cicadellinae of Indiana, Membracidae of Indiana, 
Hemiptera unrecorded from Indiana, and Cicadidae of Indiana. Later 
he was interested in insect control by ultrasonics and radiant energy. His 
research in cooperation with the Department of Agricultural Engineering 
and the U. S. Department of Agriculture produced the yellow insect-free 
light bulb, the standard survey trap, the concept of insect-free lighting, 
and the first recommendation for the use of light as a control measure. 

Dr. Deay joined the Indiana Academy of Science in 1929 and was 
elected to Fellow in 1934. He served as Divisional Chairman of the 
Entomology Section in 1957 and was Chairman of the Fellows Committee 
for 1959. Accordingly, he was a member of the Executive Committee for 
these years. He was also a member of the Biological Survey Committee 
for about 8 years. He was also a member of several other societies: Ento- 
mological Society of America, Association of Economic Entomology (vice- 
president, 1948). He was a member of Phi Beta Kappa, Sigma Xi 
(honoraries), Phi Delta Kappa, Phi Sigma (honorary) and Pi Chi Omega 
(honorary member). He was a 50-year member of the Eudora (Kansas) 
Masonic Lodge and First United Methodist Church. He served as editor 
of the Journal of Economic Entomology. He is listed in American Men 
of Science, Indiana Scientists, Who's Who in the Midwest (1955 and 
1958), and Who's Who in Indiana by Hepburn, 1957. 

In a memorial resolution for Howard Owen Deay prepared by G. E. 
Gould G. E. Lehker and L. Chandler, Chairman of the committee, an 



Necrology 29 

inscription is given from a plaque presented to Dr. Deay at his retire- 
ment. It reads: "To Dr. Howard 0. Deay, Professor of Entomology, for 
his tireless efforts to educate, counsel and befriend his students. Presented 
by his students May 19, 1964." No better evidence of the esteem and 
respect for him held by colleagues and students can be presented. 



Leslie Willard Freeman 
Escanaba, Michigan Indianapolis, Indiana 

August 17, 1915 July 7, 1969 

Dr. Leslie Willard Freeman, born August 17, 1915, at Escanaba, 
Michigan, was an internationally known neurosurgeon, professor and 
director of the surgical experimental laboratories at the Indiana Uni- 
versity School of Medicine when he died July 7, 1969. 

He received an A.B. degree at Augustana College at Rock Island, 
Illinois, in 1937. At the University of Chicago, a Ph.D. degree in 1940 
and an M.D. in 1943 were obtained. He was a laboratory instructor of 
physiology at the University of Chicago from 1937 to 1940; instructor 
of the College of Medicine, University of Illinois from 1940 to 1941; 
intern in general and neurological surgery at the Chicago Memorial 
Hospital from 1943 to 1944 and then carried on research in neurological 
surgery there from 1944 to 1945. He served with the chief paraplegic 
section of the U. S. Veterans Administration and the U. S. Department 
of Army from 1946 to 1947. He built centers and directed them under 
this program. He was research associate and assistant professor of 
surgery at the School of Medicine at Yale University from 1947 to 
1948. 

Dr. Freeman came to Indiana in 1948 as Assistant Professor in 
Surgery at the Indiana University School of Medicine in Indianapolis. 
He was associate professor from 1950 to 1953 when he became a full 
professor. He joined the Indiana Academy of Science in 1950. 

Dr. Freeman's treatment of impending paraplegia in soldiers with 
spinal injuries by immediate operation was helpful to many soldiers and 
constituted a new approach. He was a Founder of the National Para- 
plegia Foundation, and served as chairman of the advisory committee 
since 1950. 

At the Indiana University Medical School, the neurological research 
which he directed gained international recognition. Numerous contribu- 
tions were made in repairing spinal injuries and especially in regenera- 
tion of severed nerve tissue of the spinal cord. He was also interested in 
lymph and lymphatics, hemolysis and cardiovascular dynamics. He was 
named the first Betsy A. Barton Professor when the memorial chair was 
made possible in 1967 through the establishment of the fund for neuro- 
logical research. His work was also supported by the John A. Hartford 



30 Indiana Academy of Science 

Foundation, Inc. and the continuing support of the National Paraplegic 
Foundation from whom he received a distinguished service award. He 
was author of around 70 articles and chapters in several books. 

In 1963, the Paralyzed Veterans of America awarded Dr. Freeman a 
plaque as the person doing the most for paraplegia in a 10-year period 
and he delivered an honorary lecture at the 10th Latin American Con- 
gress of Neurosurgery at Buenos Aires, Argentina. 

In 1966, he received an outstanding achievement award from 
Augustana College. He was elected to Fellow in the American Physio- 
logical Society. He was a member of the American Association for the 
Advancement of Science, Society of Experimental Biology in Medicine, 
American Academy of Neurology, American College of Cardiology, Mar- 
ion County Medical Society, Indiana State Medical Association, Indiana 
Neurological Society, American Medical Society, New York Academy of 
Sciences, charter member of the Southern Neurosurgical Society, Ameri- 
can Neurological Association, Pan-Pacific Surgical Association, American 
Institute of Physiological Sciences, Association of Military Surgeons, 
Irvington Historical and Landmarks Society, Illinois State Historical 
Society and Irvington Presbyterian Church. He is listed in Whos Who in 
the Midwest, American Men of Science and Indiana Scieiitists. 

Dr. Leslie Willard Freeman achieved much in a relatively short life 
span and leaves a great gift of knowledge benefitting mankind. 



Vaclav Hlavaty 
Louny, Czechoslovakia Bloomington, Indiana 

January 27, 1894 January 11, 1969 

A distinguished service Professor Emeritus of mathematics at Indi- 
ana University, Vaclav Hlavaty died January 11, 1969, at his home in 
Bloomington, Indiana. Initially having been a recognized authority in 
differential geometry, he was author of several books and many articles 
in that field. He then turned his attention to problems in physics about 
1950 after Einstein proposed the unified field theory. In 1958, Vaclav 
Hlavaty was credited with one of the great intellectual accomplishments 
of the century when his The Geometry of Einstein's Unified Field Theory 
was published in Holland. His solution to Einstein's equations had been 
considered next to impossible. It involved a tremendous intellectual ex- 
ercise involving 64 unknowns. His work had world-wide impact and led 
the way to further scientific discovery. 

Dr. Hlavaty was born in Louny, Czechoslovakia, January 27, 1894. 
He was educated at the Universities of Prague (Ph.D., 1921), Paris, 
Rome, Oxford (1924 and 1927-1928), and Delft. He served in World War 
I from 1915 to 1919 in the Austrian Army. He was a professor of 
mathematics at Prague University from 1930 to 1948, and was an ex- 
change professor at Sorbonne (Paris) for the spring term of 1948. He 



NECROLOGY 31 

was a member of Parliament and teaching at Charles University at 
Prague when the communists took control of the country in 1948. He 
escaped to this country that year seeking refuge at Indiana University 
where he taught mathematics. He made a "poignant appeal for free men 
to support the people of Czechoslovakia in their struggle against the 
tyranny of communism" (Indianapolis News Editorial, January 14, 
1969.) 

He joined the Indiana Academy of Science in 1951 and was honored 
as a fellow in 1959. He had been chairman of the Physics Section in 
1951 when he presented a paper at the fall Academy meeting. 

He took a sabbatical leave from Indiana University in 1961 for a 
lecture tour of 19 cities in 14 countries. His subjects included relativity, 
the unified field theory and geometry. 

Dr. Hlavaty belonged to the Royal Society of Science of Liege, Inter- 
national Free Academy of Science and Letters at Paris, the Czechoslo- 
vakian Society of Arts and Scientists, American Mathematical Society 
and Sigma Xi. His life and works have been the subject of a number of 
articles and editorials in Indiana newspapers and the Indiana Alumni 
Magazine. He is also listed in Whos Who in Indiana, 1957, and American. 
Men of Science. 

Combined with the scientific genius of Vaclav Hlavaty was a deep 
humanity, love of music and the arts and a charm which won him many 
friends. He brought great honor and the warmth of his friendship to his 
adopted land, state, university and this society. 

Ruth Jordan 
Danville, Indiana Lafayette, Indiana 

July 4, 1891 July 14, 1968 

Miss Ruth Jordan was an Assistant Professor in the Home Economics 
section of the Agricultural Experiment Station at Purdue University 
and taught also in the Foods and Nutrition Department of the Home 
Economics School. Her work in foods research pioneered in this field as 
Indiana had one of the first departments in Home Economics in the 
Experiment Station. 

Born in Danville, Indiana, July 4, 1891, her early education was 
obtained there where she also attended Central Normal College. She was 
a teacher in public schools of Indiana from 1911 to 1914 and 1915 to 
1918. She then finished undergraduate work at Purdue and received a 
B.S. degree in 1920. After teaching at Central Normal College from 
1920 to 1921, she returned to Purdue in September of 1921 as an Assist- 
ant in Home Economics in the Agricultural Experiment Station. Her 
education also continued with study during the summers of 1922 and 
1935 at Chicago University and she received an M.S. degree in Foods 
from Purdue in 1929. 



32 Indiana Academy of Science 

In 1930, she was given the rank of Assistant Professor at Purdue, 
and in 1939 was made an associate in Home Economics. 

Miss Jordan was an able teacher, meticulous researcher and directed 
many research projects. Her abilities attracted many students from her 
own and other departments. Her research on the effect of various process- 
ing methods on the quality of eggs gained national recognition. Other 
interests included the effect of cooking procedures on the calcium con- 
tent of vegetables, effect of frozen storage on meat palatability, effect 
of hydrogenation of lard on culinary properties, problems associated with 
the use of homogenized milk in cookery, and calcium matabolism in 
adults. In all, she was author of over thirty research papers, seven Ex- 
periment Station publications and a number of popular articles. After 
retiring in 1961, she continued active until her death on July 14, 1968. 

Miss Jordan joined the Indiana Academy of Science in 1924. Other 
organizations to which she belonged included: the honor societies of 
Omicron Nu, Kappa Delta Pi and Sigma Xi. She was elected to Fellow 
in the American Association for the Advancement of Science in 1950. 
She was editor of the official publication of the Indiana State Home 
Economics Association, and member of Sigma Delta Epsilon, American 
Home Economics Association, American Chemical Society, Institute of 
Food Technologists, First Methodist Church, American Association for 
University Women, Alpha Xi Delta and Sigma Psi Omicron. 

She was honored in 1960 for 38 years of dedicated service to Purdue. 
Her name was included in the American Men of Science and the National 
Register of Scientific and Technical Personnel. Also, taken from a me- 
morial resolution by Professors Vianna D. Bramblett and Helen E. 
Clark of Purdue, the following excerpt gives a fitting tribute to Miss 
Jordan: "Among Ruth Jordan's outstanding qualities were a spirit of 
humility, service and independence, intellectual curiosity and integrity 
and loyalty to Purdue and Indiana. Faculty members, former students 
and others remember her with respect and sincere appreciation." 



Richard M. Rogers 
Washington, District of Columbia Marion County, Indiana 

January 13, 1924 October 6, 1969 

Mr. Richard M. Rogers was born in Washington, D. C, January 13, 
1924. His early education was obtained in that city where he attended 
Western High School. During World War II, he served in the Marine 
Corps. After that, his education resumed at the University of North 
Carolina from which he received a B.S. degree in 1951. 

His professional career began as exploration geologist for the Gulf 
Oil Company in Colorado and Oklahoma from 1952 to 1954. He was af- 
filiated with the Skaggs Oil Company at Oklahoma from 1954 to 1956. 
He became a consulting geologist in 1956 at Norman, Oklahoma, and 



Necrology 33 

was geologist and property manager for the Slick Urschel Company 
from 1960 to 1962 and Martin Marietta from 1962 to 1964 in Oklahoma. 
He came to Indianapolis, Indiana, in 1965 as a geologist for the Standard 
Materials Company. 

Mr. Rogers joined the Indiana Academy of Science in 1967 and had 
been appointed to the promising new committee, Science and Society. 

He was also a member of the first Congregational Church, Assistant 
Leader of Boy Scout Troop 56 and had served as Assistant Cub Scout 
Master and Cub Scout Master. 

Richard M. Rogers was the victim of a three-car highway accident 
in northeast Marion County, Indiana, October 6, 1969. He was only 45 
years old, married and the father of three children. What a tragic loss in 
the senseless slaughter on our highways! 



Robert L ( avere ) S h elle y 
Bluffton, Indiana Muncie, Indiana 

December 9, 1903 January 26, 1969 

Dr. Robert L. Shelley was a native of Indiana born December 9, 
1903, at Bluffton and his education was obtained in this state. He gradu- 
ated from Indiana University where he obtained an A.B. (1925), M.A. 
(1928) and Ph.D. (1929) degrees. 

His career was quite varied and very fulfilling. He began teaching at 
the Shady Springs High School in Oxley, West Virginia, from 1925 to 
1926. He returned to Indiana to further his education and to teach in 
the Clay Township High School at Kokomo, Indiana, from 1926 to 1927. 
After receiving his Ph.D. degree, he went to New York as a chemist in 
product development for the Roessler and Hasslacher Chemical Com- 
pany, Niagara Falls, New York from 1929 to 1931. Then back in Indiana 
from 1933 to 1943, he was a teacher at the Lew Wallace School in Gary, 
Indiana. He went to Ball State Teacher's College at Muncie, Indiana, in 
1943 as an Assistant Professor. He became Head of the Chemistry De- 
partment in 1945 and was a full professor from 1954 until he died Janu- 
ary 26, 1969. 

He joined the Indiana Academy of Science in 1928 while in graduate 
school and was honored by election to Fellow in 1953. He presented a 
joint paper with 0. W. Brown at a fall Academy meeting on Chlorine in 
a lead storage battery. His chief research interest was paste composi- 
tions for lead storage batteries and his Ph.D. thesis was on expansion 
as a controlling factor in positive plate paste compositions for lead 
storage batteries. 

Dr. Shelley was a member of Sigma Zeta (science, undergraduate 
honorary society) and served as national vice-president in 1949 and 
president in 1950. He belonged to Phi Lambda Upsilon (Chemistry honor 



34 Indiana Academy of Science 

society), Sigma Xi (science honor society), Alpha Chi Sigma (Chemistry 
Society), Sigma Gamma Epsilon (Earth Sciences, honorary), Sigma Phi 
Epsilon, the American Chemical Society, Indiana Chemical Society, and 
was former Director of the Muncie Technical Society. 

His interests at Ball State included membership in the University 
Senate, American Association of University Professors of which he was 
past president, the varsity athletic committee of which he was formerly 
head. He was also Elder and Trustee of the Hazelwood Christian Church. 
He is listed in American Men of Science, Indiana Lives, Indiana Scien- 
tists, Whos Who in the Midwest (V. 8), and Whos Who in American 
Education (V. 1). 

Dr. Shelley led a full life with honor. It is regrettable that such a 
capable man did not reach the average age of three score and ten. 



William Lowell Toms 
Hancock County, Indiana Greenfield, Indiana 

December 8, 1896 August 20, 1969 

William Lowell "Tubby" Toms, outdoor editor and columnist for 
the Indianapolis News died at Greenfield, Indiana, August 20, 1969, not 
far from his birthplace. He had retired in December of 1967 because of 
ill health. His informative and popular newspaper column, "Out in the 
Open" had appeared since 1946 and covered many subjects such as: 
outdoor sports (hunting and fishing especially), natural areas, nature, 
the cooking of game and oldtime recipes, remarkably accurate weather 
forecasts, conservation subjects, and sometimes anecdotes about people 
including Academy members Frank Wallace and Richard Lieber. 

Born in Hancock County, Indiana, December 8, 1896, Tubby Toms 
developed an early interest in outdoor activities and nature in the Green- 
field-Morristown area. It was during his youth that the nickname "Tubby" 
was acquired, a name by which he was known and loved throughout 
Indiana. He could quote from the Bible at length as a result of his early 
training in a devout Quaker family. He graduated from Greenfield High 
School in 1915. He then attended DePauw University, but interrupted 
his education in 1918 to serve in the Army Air Corps. He was stationed 
in France where he was one of the founders of the first air service news- 
paper, Flights and Landings. When he came home in 1919 he worked 
briefly for the Richmond Palladium Item, then he returned to DePauw 
and received the B.A. degree in 1920. He spent another brief period on 
the Indianapolis News staff, but received a government scholarship for 
study at Columbia University School of Journalism. There he received 
a Bachelor of Literature degree in 1922. Before rejoining the Indianapolis 
News staff, he was a school teacher in Rockford, Illinois, reporter for 
the Indianapolis Times, Indianapolis Star, and United Press and Interna- 
tional News Service. He rejoined the Indianapolis News staff serving 



Necrology 35 

from 1926 to 1946 as state house reporter and member of the legislative 
staff. He was reporter on the presidential trains with Landon, Roosevelt, 
Willkie and Dewey. He covered the 500-mile race for 20 years and in 
1932 wrote articles which helped win for his newspaper the Pulitzer 
Prize for Public Service in promoting- tax reduction. He wrote the column, 
Out in the Open, from 1946 until retirement. He also was state cor- 
respondent for Time, Life and Fortune magazines, and was political 
correspondent for several midwestern newspapers. He owned and operated 
three farms. 

He received many honors for his outstanding work, particularly for 
publicizing conservation matters and our natural resources. Among these 
were an award from the Izaak Walton League (Sept., 1956), and Wood- 
man of the World (April 21, 1955). He received a distinguished alumni 
award from DePauw University, May, 1966, and was made Sagamore of 
the Wabash by Harold W. Handley in 1961. Also in 1966 he was named 
"Newspaperman of the Year" by the Indianapolis Press Club. 

He was a member of Delta Kappa Epsilon, Masonic Lodge, the Mor- 
ristown Lions Club and Indiana Historical Society. He was birthright 
member of the Westland Friends Church, a founder and life member 
of the Indiana Press Club, and a member of about twelve conservation 
clubs. He joined the Indiana Academy of Science in 1940. 

Mr. Toms rebuilt a century old log cabin on 60 acres of southern 
Hancock County land which he used as a retreat. It was in a forest 
traversed by a creek. Ten acres of land were given to a boys club to 
establish the Nameless Creek Youth Camp so that the youngsters could 
enjoy nature, too. He was also interested in Earlham College and spon- 
sored student trips to Canada and Alaska. He also contributed to several 
scholarship funds anonymously. 

Several biographical articles about Mr. Toms appeared in the 
Indianapolis News (Mar. 10, 1959, Aug. 21 and 22, 1969). In these writ- 
ings, the fondness for the man, appreciation of his generosity, efforts on 
behalf of conservation, remarkable memory, amazing knowledge of a 
wide range of subjects and great good humor are expressed eloquently 
by his associates. He was, indeed, a well-loved, talented man. 



Glen W(ones) Warner 
Adams County, Indiana Portland, Oregon 

October 23, 1883 December 28, 1968 

Dr. Glen W. Warner, an internationally known educator, was born 
in a two-room log cabin four miles northeast of Decatur in Adams County, 
Indiana, October 23, 1883. Dr. Warner's education began in an Adams 
County country school. He then attended a two year high school at Mon- 
mouth, Indiana; Valparaiso Normal College, Valparaiso, Indiana; Marion 
Normal College and Business University, Marion, Indiana, where he 



36 Indiana Academy of Science 

attained a B.S. degree in 1910. Majoring in physics, he received an A.B. 
degree (cum- laude) from Indiana University in 1913. At the University 
of Chicago, he obtained an A.M. degree in education in 1919. Then major- 
ing in physics with a minor in mathematics he received a Ph.D. degree 
from Indiana University in 1937. He worked his way through school 
using the commercial training he had received. 

He taught in one-room rural schools, country high schools and was 
teacher and principal of various town and city high schools. These 
included District School, Adams County, for two years; Principal of the 
Junior High School, Decatur, Indiana, for two years; Principal of the high 
school, Peterson, Indiana, for four years; Assistant Principal of a high 
school, Goshen, Indiana, for four years; Principal of a high school, Globe, 
Arizona, one year; physics teacher, Englewood High School, Chicago, 
Illinois, for six years; physics teacher, Crane Junior College, Chicago, 
Illinois, for five years; physics teacher, Chicago City College, Wilson 
Branch, for thirty years (to Jan. 30, 1949). 

His many writings appear in a number of journals including The 
American Journal of Physics, Journal of Acoustical Society of America, 
Science, Measurement, The Chicago Schools Journal, School Science ayid 
Mathematics and others. 

Dr. Warner joined the Academy in 1949 when he lived at Lakeville. 
He was interested in both the Mathematics and Physics Sections. His 
research interests included the effect of frequency and temperature on the 
velocity of ultrasonic waves in gases; quartz, the magic mineral in ultra- 
sonics; the small linear units; and wrote his M.A. degree thesis in Educa- 
tion on "An analysis of the content of high school courses in Physics." 

Membership in other professional societies included: the American 
Physical Society, American Association for the Advancement of Science, 
American Association of Physics Teachers (charter member); honorary 
member of the Central Association of Science and Mathematics Teachers 
of which he was secretary from 1920 to 1924, Director, 1926 to 1930, 
President, 1931; Illinois Academy of Science; and Chicago Physics Club. 
He was elected to Sigma Xi, 1936. He is listed in headers in Education 
for 1941, Who's Who in American Education (1941 to 1942); and 
American Men of Science in 1944 and as late as the 9th Edition, 1955 
(Physics). Church and fraternal memberships included the Methodist 
Church, 50-year member of the Masonic Lodge, Scottish Rite, Knights of 
Pythias and 50-year member of the Eastern Star at Lakeville, Ind. 

Other activities included being president of the Farm Bureau of 
Union Township, St. Joseph County, Ind., and he was a member of the 
Chicago Teacher's Union. 

He was editor of School Science and Mathematics from 1926 to 1957, 
and was honored by the dedication of an article about him in that journal 
in October, 1957, when he retired from editorship. It stated: "It is doubt- 
ful whether an editor of any similar journal has served so long and with 
such eminence. Many tasks associated with the journal were the responsi- 



Necrology 37 

bility of his wife, Alice Koos Warner. He served above and beyond the 
editorship." He lived at Lakeville, Indiana, after retirement where his 
wife died February 2, 1962. He married Lucy M. Wright, October 3, 
1963, and moved to Portland, Oregon, where he lived until his death on 
December 28, 1968. During his last illness he wrote a letter to his pastor 
which was printed in a church news letter. In this year of man's foot- 
prints on the moon, his comments are as revealing as they are timely. 
First, referring to his 85 years of age, he remarked that he needed 
another 85 years "just to get a few things straight." Then he wrote: 
"I learned that this little system in which we live and we are trying so 
hard to get even out to the moon, is only a minimum part of what we call 
our great orbit of the milky way, and all that is only one of the great 
revolving universes that our telescopes show, so I cannot think of a God 
that looks only over our own little planet." His humility and continuing 
search for truth were sustaining virtues in this kindly man. 



Howard Ford Wright 
Wharton, Ohio Indianapolis, Indiana 

December 27, 1904 October 25, 1969 

Mr. Howard F. Wright was an assistant senior bacteriologist in 
Biological Research at Eli Lilly and Company at his death October 25, 
1969. He was born at Wharton, Ohio, December 27, 1904, and was educated 
in that state. He received an A.B. degree in zoology and chemistry in 1928 
at Ohio Wesleyan University. He also obtained an M.A. degree in zoology 
and botany from the University of Michigan in 1936, did graduate work 
at Indiana University, Bloomington, Indiana, and studied at Butler 
University, Indianapolis, Indiana. 

His professional career included teaching at Shortridge High School, 
Indianapolis, Indiana, from 1929 to 1946. He had worked at Eli Lilly and 
Company in the summer for four years when he became a permanent 
employee in 1946. His latest research interest at Lilly's was in leukemia 
and he had been co-author of several papers in that field. 

Howard Wright not only followed a career in science, but natural 
history and conservation were fascinating pursuits in his leisure time. 
His whole family enjoyed with him activities relating to these subjects. 
His daughter, Melinda, is interested in entomology and was an interested 
partner to her father in insect collecting. His son, Bruce, is chiefly inter- 
ested in physical science and mathematics. His wife, Virginia, is a biology 
teacher and counselor. 

Howard Wright was particularly interested in birds and their habits. 
He was president of the Indianapolis chapter of the Audubon Society and 
edited the state magazine, The Audubon Quarterly. About twenty years 
ago, he established a film and lecture series dealing with wildlife and 
conservation. The program has included among other guests the inter- 



38 Indiana Academy of Science 

nationally famous Roger Tory Peterson, who attracted a large, enthusi- 
astic audience. A scholarship has been established at Shortridge High 
School in his honor. The recipient can choose from four Audubon summer 
camps in Connecticut, Maine, Wisconsin and Wyoming and take courses 
dealing with conservation and natural sciences, some of which offer col- 
lege credit. The Howard F. Wright Camping Scholarship will be presented 
each year to a graduating senior of Shortridge interested in these 
subjects. 

Howard Wright joined the Indiana Academy of Science in 1943. He 
also was a member of the Indiana Historical Society; former president of 
the Watson Road Park Association; treasurer, of the Dad's Club of 
Shortridge High School and honorary Boy Scout firecrafter. He also 
belonged to the First Congregational Church. 

This quiet, gentle, accommodating man was well-liked and respected 
by students and associates. 



NEW MEMBERS FOR 1969 

The following list contains the names and addresses of all new mem- 
bers who joined during 1969. The letter (s) following the address indicates 
the Division of the Academy in which the member has indicated his major 
interest, according to the following code: 

A — Anthropology 

B — Botany 

C — Chemistry 

E — Entomology 

G — Geology and /or Geography 

H — History of Science 

L — Ecology 

M — Mathematics 

O— Cell Biology 

P — Physics 

R — Bacteriology 

S — Soil Science 

T— Plant Taxonomy 

Y — Psychology 

Z — Zoology 

D — Secondary Science Teaching 

Dr. Kraig K. Adler, Dept. Biology, Univ. of Notre Dame, Notre Dame, Ind. 46556 Z 

Mrs. Judith F. Anderson, Box 88, North Salem, Ind. 46165 ORD 

MR. William B. Barnes, 6149 Primrose Ave., Indianapolis, Ind. 46220 Z 

Dr. Paul E. Baronowsky, Dept. Biochemistry, Mead Johnson Research Center, Evans- 
ville, Ind. 47721 OCR 

Miss Karen S. Belcher, Dept. of Life Sciences, Indiana State Univ., Terre Haute, Ind. 
47809 ZOE 

MR. Ron Bitner, Entomology Hall, Purdue Univ., Lafayette, Ind. 47907 ELZ 

Mr. Harvey L. Candler, 1118 Chestnut St., Vincennes, Ind. 47591 E 

MR. Frank Cardinali, 518 S. Sixth St., Fulton, N. Y. 13069 ZLC 

Mr. Roland Chapdelaine, Biology Dept., Ball State Univ., Muncie, Ind. 47306 ZLB 

Dr. Dennis R. Christian, Dow Health Labs, Box 10, Zionsville, Ind. 46077 CPM 

MR. Dennis E. Clark, Dept. of Life Sciences, Indiana State Univ., Terre Haute, Ind. 
47809 ZLE 

MR. Howard W. Clark, P. O. Box 141, Pendleton, Ind. 46064 LBS 

MRS. E. C. Coppess, R. R. 2, Sheridan, Ind. 46069 GD 

39 



40 Indiana Academy of Science 

Dr. William W. Davis, Eli Lilly & Co., Indianapolis, Ind. 46206 CP 

Dr. Melvin W. Denner, Dept. of Life Sciences, Indiana State Univ., Evansville, Ind. 47712 
ZLE 

Mr. Frank H. Dunbar, 7401 E. Troy Ave., Indianapolis, Ind. 46239 DZB 

Mr. Roger Ferguson, 1513 Pleasant Dr., Kokomo, Ind. A 

Mr. C. T. Fletcher, Dept. Anat. & Physiol., Indiana Univ., Bloomington, Ind. 47401 ZCL 

Miss Marjorie Fuller, Mead Johnson Co., Evansville, Ind. 47721 HC 

Dr. James E. George, Dept. Chemistry, DePauw Univ., Greencastle, Ind. 46135 CGP 

DR. Robert B. Gibson, Dept. Phys. & Health Sci., Ball State Univ., Muncie Ind. 47306 Z 

MR. Daniel R. Goins, Daleville High School, Box 525, Daleville, Ind. 47334 ZLB 

Miss Dagnija Grins, Clinical Lab, Indiana U. Med. Center, 1100 W. Michigan, Indian- 
apolis, Ind. 46220 RDO 

MR. William J. Gulish, Driftwood Experiment Station, Vallonia, Ind. 47281 LZB 

MR. Larry M. Hagerman, 134 S. Boeke Rd., Evansville, Ind. 47714 

MR. Donald W. Hamilton, Vincennes Univ., Vincennes, Ind. 47591 E 

Dr. George D. Hanks, Dept. Biol. Sciences, Indiana Univ., NW, Gary, Ind. 46408 ZOL 

DR. William B. Hebard, 3402 Deerwood Dr., New Albany, Ind. 47150 ZOB 

MR. Donald R. Hendricks, Box 6, Earlham College, Richmond, Ind. 47374 ZLA 

Dr. Eldon L. Hood, Dept. Agronomy, Purdue Univ., Lafayette, Ind. 47907 SCG 

Dr. Robert J. Hosley, 5001 N. Illinois St., Indianapolis, Ind. 46208 G 

MR. Luther B. Hughes, Jr., 101-15 MSC, Purdue Univ., Lafayette, Ind. 47906 SRC 

DR. William T. Jackson, Eli Lilly & Co., Indianapolis, Ind. 46206 RC 

MR. D. P. Jerrett, 421 Anderson St., Greencastle, Ind. 46135 OZE 

Dr. Hollis R. Johnson, Astronomy Dept., Indiana Univ., Bloomington, Ind. 47401 PHL 

Dr. William S. Klug, Dept. Biology, Wabash College, Crawfordsville, Ind. 47933 OZ 

MR. Robert K. Landes, 2002 O St., Bedford, Ind. 47421 LBZ 

MR. Henry R. Lawson, Entomology Hall, Purdue Univ., Lafayette, Ind. 47907 ELZ 

MR. Terry N. Lewis, Dept. Biology, Indiana Central College, Indianapolis, Ind. 46227 
ODZ 

MR. Richard Lopez, R. R. 3, Box 351, Muncie, Ind. 47302 ERL 

MR. Thomas S. McComish, Dept. Biology, Ball State Univ., Muncie, Ind. 47302 L 

Mr. Charles F. McGraw, Hayes Regional Arboretum, 801 Elks Rd., Richmond, Ind. 47374 
LBZ 

Rev. Basil Mattingly, St. Meinrad College, St. Meinrad, Ind. 47577 YAG 

MR. Richard S. Mills, 103 SW 6 Street, Richmond, Ind. 47374 ZLR 

Mr. Daniel L. Mollaun, 416 E. Pearl St., Batesville, Ind. 47006 DRL 

MR. Donald N. Moore, Biology Dept., Ball State Univ., Muncie, Ind. 47306 BTZ 

MR. Donald I). Pascal, Jr., 57 Westmont, West Hartford, Conn. 06117 ZL 



New Members 41 

MR. James L. Pease, 799 E. Jefferson St., Franklin, Ind. 46131 ZCL 

MR. Anthony J. Perzigian, Campus View House, A 513, Bloomington, Ind. 47401 A 

Mr. Randal B. Richardson, Box 131, R. R. 7, Bloomington, Ind. 47401 LZR 

Dr. Eugene P. Schwartz, Dept. Chemistry, DePauw Univ., Greencastle, Ind. 46135 C 

Mr. Karl Schwenk, Anthony Apt. 32, Muncie, Ind. 47304 ORB 

MR. Frank S. Sterrett, Box 1176, Earlham College, Richmond, Ind. 47374 ZL 

Dr. William L. Stoller, 1901 S. Park Rd., Apt. A-202, Kokomo, Ind. 46901 ZY 

MR. William E. Stovall, Dept. Physiol. & Health Sci., Ball State Univ., Muncie, Ind. 
47306 RE 

Mr. Ronald E. Surdzial, 436 N. Indiana St., Griffith, Ind. 46319 CDM 

Dr. Robert Van Etten, Dept. Chemistry, Purdue Univ., Lafayette, Ind. 47907 C 

MR. Kent D. Vickery, Dept. Anthropology, Indiana Univ., Bloomington, Ind. 47401 ALB 

Dr. Gary L. Walsh, Dept. Biol. Science, Indiana Univ., NW, Gary, Ind. 46408 LZS 

MR. Neil V. Weber, Dept. Geog. & Geol., Indiana State Univ., Terre Haute, Ind. 47809 
GH 

MR. Harmon P. Weeks, Jr., Dept. Forestry & Cons., Purdue Univ., Lafayette, Ind. 47907 
LZ 

PROF. Paul P. Weinstein, Dept. Biology, Univ. Notre Dame, Notre Dame, Ind. 46556 Z 

MR. Rex Wells, R. R. 6, Columbia City, Ind. 46723 ZLE 

Dr. Jack M. Whitehead, Dept. Soc. & Anthropology, Ball State Univ., Muncie, Ind. 
47306 A 

Mr. Egerton Whittle, Dept. of Life Sciences, Indiana State Univ., Terre Haute, Ind. 
47809 Z 

MR. Donald L. Wilbur, Dept. of Life Sciences, Indiana State Univ., Terre Haute, Ind. 
47809 Z 

MR. Frank T. Willis, 390 S. Home St., Apt. A-4, Franklin, Ind. 46131 LZE 

Mr. Robert C. Wingard, Jr., P. O. Box 173, Paoli, Ind. 47454 S 

MR. & Mrs. Edgar I. Winger, R. R. 1, Bennett St., Paoli, Ind. 47454 GAH 

Dr. Walter E. Wright, Eli Lilly & Co., P. O. Box 618, Indianapolis, Ind. 46206 

MR. David D. Wynne, 1431 N. Jordan Ave., Bloomington, Ind. 47401 AZL 

Mr. Ronald E. Zimmerman, 513 S. Jefferson St., Brownsburg, Ind. ZCO 



PART 2 

ADDRESSES 

AND 

CONTRIBUTED 

PAPERS 



Hanover, Indiana 
October 24, 1969 



The address, 'The World of the Honey Bee" was presented 
by retiring president, Dr. Howard R. Youse, at the annual 
dinner meeting of the Academy at the J. Graham Brown 
Campus Center at Hanover College on Friday evening, Oc- 
tober 24, 1969. The address by Dr. John B. Patton, Chairman, 
Department of Geology, Indiana University, and State Geolo- 
gist, Indiana Department of Natural Resources, was given at 
the Spring Meeting dinner at the J. Graham Brown Campus 
Center at Hanover College on Friday evening, April 26, 1969. 
His subject, "To Ruine a World," deals with the problems of 
solid waste accumulation and the accelerating consumption 
of non-renewable resources. 



PRESIDENTIAL ADDRESS 



The World of the Honey Bee 

Howard R. Youse 

DePauw University, Greencastle, Indiana 

This has been a very eventful year for science. The climax was prob- 
ably reached this summer when the Apollo astronauts set foot on the 
moon, thus fulfilling a dream of man for centuries. At the same time we 
hear many reports among scientists that all is not well on earth. Several 
of our recent presidential addresses have been concerned with ecological 
problems that face us today. We have a Science and Society Committee 
which is exploring the ways in which the Academy might play a more 
active role in attempts to solve many of the problems of society. 

It may seem strange to some of you that a botanist would choose as 
his topic The World of the Honey Bee; my only qualification is that I 
have worked on pollen grains utilized by honey bees. I thought it might 
be a topic that would be of interest to many of you in various branches 
of science, since scientists from various disciplines have found the study 
of the honey bee to be a source of lasting wonder. I personally am con- 
vinced that the study of the honeybee may give us important biological 
information that may be helpful in understanding some of our problems. 

The honey bee is one of the few insects that has been domesticated 
by man. The species we are most concerned with is Apis meUifera of 
which there are many varieties. Life in the hive centers around the queen. 
She is larger, lives longer, and forms the hub around which all activities 
move. Most of the routine activities in the hive are carried out by numer- 
ous workers. An average hive might have about 30,000 workers, but this 
may vary widely, and man may manipulate the hive to have 100,000 
workers. Then from time to time a few drones may be observed in the 
hive. 

When the geneticist examines the hive, he finds that the drones are 
males with a haploid set of chromosomes. Thus they must develop by 
parthenogenesis. The queen and workers are females with a diploid set of 
chromosomes and thus must have developed from fertilized eggs. The 
chromosome complement thus determines if we have a male or female 
bee. When we examine the growth and development of the larvae emerg- 
ing from the eggs, we find that all the larvae receive royal jelly, honey, 
and pollen the first 24 hours of their lives. The larvae that develop into 
queens are continually fed royal jelly and mature in about 16 days. The 
other larvae are fed only honey and pollen as they are developing, and it 
takes about 21 days for workers to mature and 24 days for drones. Royal 
jelly is a glandular extract from young worker bees, and it must be a 

45 



46 Indiana Academy of Science 

remarkable material, as it determines whether the diploid eggs will 
develop into a functional female capable of laying eggs or a non- 
functional worker unable to lay eggs. One might conclude that both 
genetics and nutrition are important in the growth and development of 
the honeybee and perhaps of all living organisms. 

If we examine a hive early in the spring, we often find a number 
of queen cells developing and a relatively large number of drones. The 
hive is usually crowded with workers at the same time. The society then 
goes through a stage known as swarming. At this time the old queen 
takes off from the hive with some of her workers. This usually occurs just 
as the young queens are ready to mature. As the new queens emerge from 
their cells, they engage in mortal combat until just one queen is left. In 
case the old queen has not left, she in turn is killed. When the new queen 
of the hive is about 5-7 days old, she is ready for her nuptial flight. 
There is quite an air of excitement around the hive at this time. Mating 
of the queen takes place outside the hive, and it is the one big day for 
the drones. The virgin queen takes off from the hive with all the drones 
in pursuit. Presumedly the drone that is strongest and fastest mates with 
the queen. Enough sperms are deposited in the queen to last her lifetime. 
In the process the drone loses his life. When they return the other drones 
are usually killed by the workers. This drama in the hive might lead us 
to conclude that nature is often very ruthless. 

Within 48 hours after the queen returns from her nuptial flight she 
settles down to the main activity of her lifetime, laying eggs. The queen 
can lay unfertilized eggs, although she most often lays fertilized eggs. 
She may average about 2,000 a day, but at the peak of the honey flow, 
she may lay 5,000 a day. She will lay over a million in her lifetime. She is 
constantly attended by a host of workers as she goes about her business. 
The workers also go about a wide variety of jobs. Anyone who has 
observed a teaming mass of thousands of bees has probably asked how do 
bees communicate with each other. 

An examination of a hive will reveal some bees busy building comb 
by secretion of thin scales of wax from their wax glands which are then 
moulded into almost perfectly hexagonal comb cells. Others may be mend- 
ing damaged comb structure. Other young workers are engaged in giving 
royal jelly, pollen, and honey to the developing larvae. Some are engaged 
in cleaning the hive and removing waste from the hive. The largest num- 
ber of workers are involved in collecting pollen and nectar. As they 
return to the hive other workers receive the nectar and deposit it in 
the comb where it is evaporated to the right consistency and then capped. 
Other workers strip pollen and stamp it into cells. As all these activities 
are going on, a number of guard bees stand at the entrance of the hive 
to see that no bee gets in except a working member of the hive. 

These observations might lead one to conclude that each bee special- 
izes in a particular function, but if a bee is tagged and observed over a 
period of time, it will be evident that this is not true. Each worker per- 
forms a succession of each of the various tasks. Some of these may be 
more or less synchronized with the development of the bee, but one finds 



Presidential Address 47 

great variation depending on the environmental conditions inside and out- 
side the hive. How is an individual bee informed of the tasks that need to 
be done? Lindauer (1) tagged individual bees and observed them over a 
period of time. He is convinced that each bee gathers her own information 
by extended inspection tours around the hive. Then she seems to do what 
needs to be done. This seems to be a rather good way to get the job done 
in the event that each bee is willing to do something. 

The general activity in the hive is the collection of nectar and pollen. 
Nectar is used for the production of honey, which is the main carbo- 
hydrate food. Pollen is the main source of proteins and vitamins, and it is 
stored directly as it is brought into the hives. For many years it was a 
source of much debate as to how bees were able to communicate rich 
sources of nectar and pollen to the workers. It was known that bees were 
very sensitive to different colors. They also were very sensitive to differ- 
ent odors or essential oils found in flowers. Von Frisch (2) observed that 
bees seemed to go through a curious dance when returning to the hive 
with a load of nectar or pollen. When sources were close by, the bee per- 
formed a "round dance." This seemed to indicate that the bees need only 
to go out of the hive and collect. If the source was at some distance, the 
bees went through a "tail-wagging" dance. The direction is conveyed by 
the orientation of the dance in relation to the sun. The rhythm and inten- 
sity of the dance as well as certain sounds indicate the distance. There are 
records of bees flying ten miles to a nectar source, although they seldom 
will fly more than three miles. On one of these flights, the honey bee col- 
lects a load of nectar equal to half its own weight, then cruises non-stop 
back to the hive at about 15 miles per hour. This would suggest that the 
bee is indeed a superior flying insect both from the point of view of design 
and use of energy. 

Individual workers may average about ten trips a day, and bees from 
a single hive may visit over a quarter of a million blooms a day. Thus 
they are our most effective pollinators of flowering plants. It has been 
estimated that over 100,000 species of flowering plants would disappear 
from the earth if bees were eliminated. When DDT was first applied over 
orchards by airplanes, it was observed that both fruit production and 
the bee population dropped in the area. Since honey bees work only dur- 
ing the day, they tried to get around this by dusting at night or on cloudy 
days when bees were not active. Now we are beginning to have some 
additional concern of the effect of DDT on other organisms such as fish 
and birds. 

The product from bees that man is most interested in is honey. If 
one were able to collect nectar from flowers and evaporate the water, one 
would still not have honey. As soon as nectar is sucked into the honey 
bee's crop, enzymes are mixed in that convert the sugars to dextrose and 
levulose. At the hive the load is transferred to the crops of young work.- 
ers The young bees thoroughly mix it with more enzymes. It is finally 
deposited in open cells of the combs. Other workers then gently fan it 
to evaporate the excess water. When it is just the right consistency, it is 
capped. A chemical analysis of the honey at this point would show about 



48 Indiana Academy of Science 

40% levulose, 35% dextrose, and 18% water. Thus honey is a very con- 
centrated carbohydrate food, and anyone familiar with the complexities 
of sugar chemistry might decide to just let the honey bee go on doing 
the job. 

Honey is stored in wax combs. The comb is only about 0.002 inch 
thick, but it can support 25 times its own weight. The wax comes from 
the wax platelets on the abdomen of young bees, and it is estimated that 
six to seven pounds of honey are consumed to produce one pound of wax. 
The wax has a high melting point of around 140° F and can be used in a 
wide variety of products. It is used to make candles and to insulate wires. 
It is often used in furniture waxes and varnishes as well as in phonograph 
records and lubricants of various types. 

In concluding, I would like to give you a few tips on how to get along 
with bees. For some reason bees do not like black; so it is a good policy to 
never wear black clothes around them. They are very sensitive to vibra- 
tions so one should avoid jarring them, especially their hive. They are 
very sensitive to odors, and I am convinced that many people just have 
B.O. as far as bees are concerned. Smoke seems to quiet bees, so they are 
frequently smoked before a hive is opened. A good pipe or cigar tends to 
keep them away from your face in case you do not want to bother with a 
veil. If you are stung, you may rest assured that it hurt the bee more 
than it does you, as the honey bee dies. When stung by the honey bee, one 
should quickly cut off the stinger with a sharp knife, as the stinger acts 
as a little syringe that gradually forces the venom into your skin as it 
contracts. One should also wash after being stung, as the venom tends to 
excite other bees. Bee stings usually lead snake bites as a cause of death 
each year, and thus one might decide that they should be eliminated in 
order to save human lives. However, we might find that more human lives 
would be lost due to starvation. The problem of what to do with honey- 
bees is rather typical of many ecological problems that face us today. 
There seem to be several courses of action, and we are uncertain what 
path will be best in the long run. Perhaps if each problem were studied 
in a variety of ways, we might gain additional insight as to the best 
course to follow. It is my hope that the Indiana Academy of Science will 
be a leader in attacking the many problems we face, and I hope each of 
you will keep busy as a bee doing what needs to be done. 



Literature Cited 

1. LlNDAUER, M. 1961. Communications Among Social Bees. Harvard University Press, 
Cambridge, Mass. 143p. 

2. von FRISCH, K. 1950. Bees, Their Vision, Chemical Senses, and Language. Cornell 
University Press, Ithaca, N.Y. 119p. 



To Ruine a World 

John B. Patton 

Chairman, Department of Geology, Indiana University, 

and 

State Geologist, Indiana Department of Natural Resources 

My text and title are taken from Fontenelle's Plurality of Worlds, 
written in 1686, from which I quote. . . . 

"How," cried the countess, "can suns be put out?" "Yes, without 
doubt," said I, "for people some thousands of years ago saw 
fixed stars in the sky which are now no more to be seen. These 
were suns which have lost their light and certainly there must 
be a strange desolation in their vortexes." "You make me 
tremble," replied the countess. "0 madam," said I, "there is a 
great deal of time required to ruine a world." 

My thesis is that Fontenelle was wrong, or at least that he would be 
wrong if he said the same words today, and that a great deal of time is 
not required to ruin a world. Man, in just the short part of this earth's 
history that he has been present, has gone far toward creating ruin, and 
especially in those parts of the earth where educational opportunities and 
material prosperity would appear to offer the greatest opportunity to 
improve on rather than to detract from Nature's handiwork. 

The views that I express are subjective and stated from the anthropo- 
centric viewpoint that what is bad for mankind is bad for the world. On 
the other hand, the objective view of the geologist must be that mankind 
has come and will go, leaving little evidence of his presence or handiwork, 
and leaving the earth little the better or worse for his ephemeral presence 
or for his passing. I mean by this that man's activities, wondrous as they 
may have been at times and at places, and catastrophic as they have 
been and are now over much of the earth's land area, are insignificant 
in comparison with the inanimate changes, and even some related to the 
plants and animals, of the past. That past, we must presume, permits 
prognosis for the future, just as the doctrine of uniformitarianism states 
that the present is the key to the past. Within the part of geologic history 
recorded reasonably well in the interpretable rock record, mountain- 
building and continent-building forces have caused vast expansions and 
contractions in both the area and the height of the lands. Surface vol- 
canism has buried tremendous land areas, increasing the amount of land 
in some instances, and leading to the terranes and soils that have per- 
mitted the flowering of economies and cultures. In the geologic past the 
evolution and proliferation of certain plant and animal strains have 
changed the earth's surface, or at least substantial parts of it, to a degree 
that would have occasioned the outcry "this means the end of the world 
as we know it" had there been a voice to speak. 

49 



50 Indiana Academy of Science 

To return, however, to the view that the world is being ruined, in the 
sense that the more desirable parts of it are becoming less acceptable as 
an abode for mankind, the ruin of which I speak is taking place in three 
ways: first, we are strewing the surface and filling the shallow subsur- 
face with waste to an extent that further constricts the amount of 
usable land and makes even that area less tolerable, and we are making 
much of the air and fresh water unfit for human consumption. Second, 
we are destroying the accomplishments of the past. Third, we are con- 
suming our irreplaceable natural resources at a rate that suggests sub- 
conscious acceptance of the view that man's occupany of this planet will 
be short. 

The air is being spoiled with noxious gases and dusts. Surplus water 
vapor and carbon dioxide are being added to the atmosphere at a rate 
that will affect world climates. 

Water is being contaminated with chemicals — some of them poison- 
ous — ,sewage, industrial wastes, and sediment, and the heat balance is 
being changed by thermal pollution. 

Our soils are being destroyed, covered, or made unusable by paving 
and stripping, by being covered with trash and garbage, and by being 
buried under subsoil materials in our determination to alter the natural 
topography. 

Biologists are particularly familiar with those factors that are 
acting to disturb natural environmental balances, and they have been for- 
tunate in having such persons as Rachel Carson to speak against tamper- 
ing with the environment before adequate study of the consequences. 
How great our debt to those who force us to review the effects of our 
errors! And how fortunate if the warnings come in time! 

But the aspect of ruin in which I may be best qualified to speak is 
that related to geology. The exploitation of ores for their essential pur- 
pose of yielding metals has led to extensive destruction and pollution at 
both the mining and the smelting levels of development. Landscape alter- 
ation is inherent in removal and processing of mineral raw materials, and 
the problem thus becomes one of utilizing the mineral commodities for 
man's benefit without, in the process, creating damage that will result 
in a net loss to man's long-term welfare. Many pit, quarry, and mine 
operations are conducted in a manner that is efficient in terms of 
present-day dollar profit and loss but inefficient in terms of long-range 
land use. Few deposits are worked in a manner that will recover the 
largest amount of usable material from the minimum number of acres 
of land used or ruined. Some encouragement is to be found in the fact 
that an increasing number of mineral producers, principally in the non- 
metals construction materials and numbering very few among the metal- 
mining companies, plan their land use programs with an eye toward 
rehabilitation and even improvement of the terrane as their acreage is 
worked out. 

It is a curious paradox that as we have more demand for water we 
have less respect for its beauty. It is also a paradox that as personal and 



Address 51 

domestic cleanliness increases we should tolerate a landscape in which 
trash is more and more evident. To a considerable extent this results from 
a complete breakdown in the scavenging system — or the salvage system, 
to use a more polite term. In all earlier societies, and in many societies 
today, waste had too much value to remain a blight on the landscape. 
Mayhew recorded (London Labour and the London Poor, 1851) the degree 
of specialization that characterized salvage in the Victorian London of his 
day. For each type of discard material, different scavengers came to the 
doors, some buying old iron and various others buying grease, drippings, 
broken metal, old umbrellas, rabbit skins, waste paper, glass, old bottles, 
and old clothes. Other scavengers, gathering waste from the streets and 
river flats, were classified as bone grubbers, rag gatherers, trampers, 
mud-larks, pure-finders, and other named specialists, and in addition to 
the activities implied by their names they collected waste metal — the 
most valued material — ,rags (which they sorted into lots of white or 
colored for the paper makers and into canvas and sacking for other pur- 
poses), cigar ends, old wood, chunks of coal, waste paper, and every 
salvagable discard. Mayhew estimated their number at 800. Dust con- 
tractors, who numbered 80 or 90, were paid by the city to remove ashes 
and cinders, and they sifted the ash for resale as soil conditioner and raw 
material for brick. 

No generation younger than mine has heard the street cry (or alley 
cry, to be more precise) "Any rags, any bones, any bottles today?" In 
many systems all the trash, garbage, and other discarded materials are 
loaded into one truck and hauled to some spot at which fuel is consumed 
to incinerate them or land is used to stack them or bury them — 
generally without regard to the possible polluting effect on ground 
water. The industries that once depended, entirely or in part, on waste 
and scrap have ceased or gone to other materials. As an example, an 
essential ingredient of the glass industry is the material called cullet, 
whi h is broken glass. It was formerly salvaged and sold as clear, colored, 
or mixed cullet. Its function in the glass furnace is to provide nuclei of 
vitrification that speed and improve the melting process. Most glass 
companies make their own cullet today. Whatever they are making when 
the need arises — fancy decanters, pickle bottles, or anything else, feeds 
off the conveyor belt onto a concrete floor to provide cullet, while the 
waste glass of our society fills acres of our landscape. Not only will it 
defy the process of weathering for thousands of years; it makes the soil 
untillable and dangerous to walk on or work with. The Romans avenged 
themselves on Carthage by plowing the site and sowing it with salt. The 
way modern society treats itself and future generations makes the 
Romans' treatment of their vanquished enemies seem tender-hearted. 

The first step in the war against solid waste would be a strictly- 
enforced program of presorting by users. Trash would be sorted into 
newsprint, magazines, glass, cans, waste metal, and burnables (penalty 
for non-compliance: a day at the dump, sorting). Refuse would have to 
be separately collected, on a rotating schedule if necessary, and separately 
processed. The cost of collecting and policing would be balanced by the 



52 Indiana Academy of Science 

saving: in land-acquisition costs, by salvage, and, of course, by aesthetic 
appreciation in land values. 

The second step in the war would involve large-scale reforms in pack- 
aging. Our present methods use resources at an inexcusable rate and 
compound the damage by contributing excessive waste. 

Among the accomplishments of the past, man has, over the centuries, 
adorned the earth with gardens and groves, bridged rivers, laid roads, and 
built homes and public buildings, the most splendid of which have been 
ecclesiastical. The skill, patience, and effort they cost have been pro- 
digious, and the artistic achievements they represent humble us in these 
late untalented days. The apprentice system, in spite of its severities, 
raised up craftsmen, which our kinder modern society cannot replace. 
Their handwork is irreplaceable and deserves our protection. This country 
has been even more careless about its historic buildings than European 
countries, but here and there an aesthetic conscience is beginning to 
inspire preservation programs. The State of Indiana needs more civic 
efforts of the kind typified by Historic Madison, Inc., and the State and 
private activities at New Harmony in the fields of historic preservation 
and restoration. 

As a total environment for mineral resources we could say this about 
the earth: by chance we live on one of nine known planets in a minor 
solar system that forms an obscure part of one of the lesser galaxies. 
We are concerned with a very small fragment of the total matter in 
the universe. We are inhabiting this planet only transiently, our entire 
history as a human race having occupied but one million years of the 
four billion years of known geologic time. My point in mentioning this 
transience is that the earth's mineral resources are different from period 
to period in geologic time. There would have been a place in the geologic 
time scale, undoubtedly, when the earth's mineral resources would not 
have included oil and gas. Organisms either were not present or were 
not abundant enough to furnish the hydrocarbons that constitute oil and 
gas. We know quite precisely the place in geologic time at which there 
would have been no coal resources — no members of the coal family that 
includes lignite, peat, bituminous coal, or anthracite — because the first 
coals appeared with the development of vascular tissue in plants late in 
the Devonian Period. 

Similarly, before a certain point in geologic time, there would have 
been few iron ores of the kind we regard as commercial now, and con- 
versely, because the greater part of our commercial iron ores are Pre- 
cambrian, meaning that they are 550 million years old or more, there 
was a time in earth history when iron ores of the type that we consider 
commercial today were vastly more abundant than they are now. Most of 
the iron ores of this kind have undoubtedly disappeared through the 
processes of weathering and erosion since that time which was most 
favorable for their accumulation. 

All mineral deposits are theoretically exhaustible, as sufficient use 
would ultimately consume all of anything at the earth's surface. Materials 
that can be removed from the sea are almost limitless so far as their 



Address 53 

reserves are concerned. One other type of mineral resource may also be 
classed as practically limitless, meaning that man's use will not exhaust 
the supply, and these are the materials that occur as common rock 
types. We shall never run out of an adequate supply of granite to keep 
us all in tombstones. We shall never lack an adequate supply of basalt, 
limestone, dolomite, and other such materials to crush for concrete aggre- 
gates and road metal. We shall never run out of salt because it is a 
relatively common rock type, and even if we could not extract limitless 
quantities from the sea, we could mine salt virtually forever without 
exhausting the world's supply. 

Even after using these dangerous words, limitless and inexhaustible, 
I feel obliged to tell you that most of the mineral deposits that have been 
called inexhaustible have long since been exhausted. The gas supply that 
caused northern Indiana and western Ohio to be settled and industrialized 
in the 1880's and 1890's was called inexhaustible in every county news- 
paper in the two states, but specifically it lasted as a good source of 
supply for less than 25 years, in large part because it was wasted, but 
if the best of conservation measures had been applied to it, the life 
expectancy would still have been less than 50 years. Most inexhaustible 
mineral resources have been exhausted, and most of the ones we now call 
inexhaustible are likely to be exhausted unless they are fairly common 
rock types or obtained from the sea. 

The fossil fuels offer some of the most striking examples of the 
interrelationship between mineral resource needs, the pattern of our 
society, and man's prospects for a tolerable future. The problems involved 
in the removal and use of the solid fuels are no greater than those in the 
liquid, gaseous, and nuclear fuels, but they are more visible. 

Coal mining, whether by stripping or by underground workings, 
necessarily disrupts the surficial environment. The surface effects of 
stripping are the more apparent, and the subsidence effects of shallow 
underground mining more delayed. Land reclamation is extensive and 
growing in our own State and certain other regions in which the strata 
are flat-lying and the natural topography subdued. In Indiana more strip 
coal land undergoes some reclamation treatment per year than is newly 
mined, but in regions of steeply dipping coal beds and rugged topography 
it is inevitable that many acres of land will be destroyed for every acre 
of coal recovered. Reclamation can accomplish little in those terranes, and 
the only answer, if there is an answer, is rigorous restriction and regula- 
tion of surface and near-surface coal mining. 

There is no more striking example of the enormous increase that has 
taken place in per capita consumption than in the domestic use of fuel or 
heat energy from fuel. Less than a century ago the average household 
heated a scant three rooms, of perhaps a much larger housing space, with 
small open fires, in many instances burning wood, in which case the 
summer's growth supplied the winter's warmth. In many social orders 
a community oven provided most of the heat for baking, and one hot 
meal a day was the accepted pattern. Now, in our society at least, 



54 Indiana Academy of Science 

enormous cubages of space are kept at summer temperature all winter 
by fossil fuel, and at spring's temperature all summer, also by fossil fuel, 
and empty houses — weekend houses as the fashion calls them — are kept 
at 55 or 60 degrees to be instantly usable should whim suggest their 
temporary occupancy — an extravagance with few parallels in history. 

In the energy field we have laid out our homes, our towns, and our 
communications on the premise that cheap petroleum products will be 
available indefinitely. By analogy the British laid out their social order 
in the cheerful delusion that cheap domestic help would always be 
abundant. Their economic dislocation, when things changed, was and 
continues to be drastic. Our dislocations, when the energy sources dwindle 
or price themselves out of our market, will be catastrophic. 

By way of review in this field of energy sources, let me say that one 
of the most respected estimates calculates 200 billion barrels of total 
recoverable petroleum. Of this amount 85 billion barrels have been pro- 
duced and used to date. A reasonable projection indicates that the 
remainder will last 65 to 70 years, at progressively increasing cost. 
Naturally the day will not come when all oil has been found and produced 
and used, but by the time the indicated date arrives the discovery- 
production-consumption ratio will have reached that point at which the 
liquid and gaseous fossil fuels will no longer supply a significant part of 
our energy needs. 

Coal reserves will suffice for several more centuries, even consider- 
ing the inevitable shift to coal for energy needs filled now by other 
materials. 

Nuclear energy will be required in increasing amount to phase out 
both solid and liquid-gaseous fossil fuels, but present methods of develop- 
ing nuclear energy cannot keep up with the required increase. Only 
breeder reactors, which are estimated to be 20 years away so far as 
extensive development is concerned, can provide the needed energy sup- 
plement for short-term purposes, and they will produce toxic wastes 
beyond our ability to cope with them. Only fusion reactors, which are 
estimated to be 40 years away, but which will not produce toxic wastes, 
can give long-term energy security. 

In the non-fuel mineral industries, both reserves and outlook vary 
tremendously, and the two are not the same and are, in fact, not entirely 
correctable. Reserves are known supplies recoverable by mining, and the 
known reserve divided by annual consumption gives an indicator, ex- 
pressed in years, called the reserve-consumption index. An index as low 
as 25 years is no occasion for concern in the case of a commodity for 
which we believe that the reserves can be extended through exploration 
programs; numerous mineral substances fit this category. A reserve- 
consumption index of 25 years is alarming in the cases of those commodi- 
ties for which the prospects of significant additional reserve discoveries 
appear to be dim. The answer to a supply failure for some minerals may 
be found in substitution. Tin, as an example, had a 25-year reserve- 
consumption index in 1964, and the prospects for additional major dis- 



Address 55 

coveries are poor. Yet the major use of tin — plating — is one for which 
substitutes, not as good and not as economical, but still acceptable, are 
possible. 

Both mercury and silver, with 1964-65 indices of 14 and 23 years 
respectively, are metals for which the index is likely to be extended 
through additional discovery, but even so an adequate supply is probably 
short-lived, and each has essential uses for which no substitute is avail- 
able at an economic level. 

All reserve-consumption indices are likely to change with new dis- 
coveries and with alterations in uses and technology, and so I prefer to 
deal with our overuse of mineral resources on a basis that cannot, 1 
believe, be denied: we waste them prodigiously. As a person who has 
spent a considerable number of years in searching for mineral resources, 
I am distressed by the upsurgence of such things as the throw-away 
bottle. When my children bring home soft drinks in these containers, I 
wonder whether the two minutes of pleasure derived from their consump- 
tion has justified the exploration and production effort for high-silica 
glass sand, feldspar, borax, and other ingredients of a rather high- 
quality container which must have greater intrinsic value than the 
syrupy mixture it contains. The aluminum beer can that is discarded 
after use would be the most valued household possession in some under- 
developed societies. 

To compound the difficulties presented by these examples, both items 
become particularly durable elements in our growing amount of waste 
materials, and each will survive the process of weathering better than 
man himself. One of the principal reasons that life has expanded and 
survived so successfully on this planet is that organisms are disposable 
and in fact decay to enhance prospects for living things in the future 
rather than the reverse. Many organisms have, in a sense, left their 
bodies to science and have contributed to the limestones, clays, shales, 
coal beds, and oil and gas accumulations on which we now depend, but 
man is the first organism to leave artifact offal that may act to stifle life 
on the planet. 

Water is everyone's problem, and the geologist's role is to locate 
ground water, establish sites for surface reservoirs, assist in matters of 
drainage and flooding, and do his part in preventing pollution of both 
surface waters and subsurface waters. 

Water differs from all the other inorganic materials we require in 
that it alone has ethical implications. Present-day Western morality 
judges countries and people with respect to cleanliness of their persons 
and households — in other words on the basis of the amount of water 
available for plumbing, laundering, and bathing. Neither the courtesy and 
competence of the population, the harmony of the architecture, or the 
order and tidiness of housekeeping are as important as available surface 
or subsurface water and the means of conserving and supplying it. One 
may categorize a populace as arrogant, parsimonious, drunken, lazy, dis- 
honest, immoral, or even cowardly and still encounter only an agitated 
rebuttal, but call it dirty and relations are terminated. 



56 Indiana Academy of Science 

Some sort of ceremonial washing- and purification has been a feature 
of most religions. Basins or fountains for ritual ablutions are architectural 
features of their places of worship, as in the great European baptisteries, 
and they hold sacred certain rivers, springs, or lakes. I think it possible 
that some of the revulsion we feel at today's dirt-cults among the young 
is not just a generation gap but a natural response to a perversion of a 
deep human instinct. The urge to feel clean is innate, perhaps phylogenetic 
and related to the fact that life came from the sea, but most societies 
until ours have been, or have been obliged to be, realistic about personal 
water consumption. We are not realistic. On a few occasions in my 
youth I broke ice in a pitcher to wash in the morning — a commonplace 
occurrence one generation earlier — my son is incensed if a 20-minute hot 
shower is criticized. 

This assumption that moral superiority and personal cleanliness are 
mutually dependent causes great international strain. In much of Asia, 
for example, if all the available water in the populated regions could be 
collected and piped to users, there still would not be enough per capita 
to operate a sanitary sewage system — much less provide water for 
industry or irrigation. 

That society is most enlightened that best lives in harmony with its 
environment and secures that harmonious relationship for future gen- 
erations ad infinitum. It is time to adjust our personal requirements to 
realistic per capita use of energy and raw materials, and we are overdue 
for a disinterested examination of our environment and establishment of 
a regimen to insure our national health and longevity. A great annual 
outcry is heard about the national deficit. The need to balance our 
resource budget is of far greater importance to this nation than is the 
balancing of the fiscal budget. 

Peter Blake concluded his eloquent book God's Own Junkyard with 
these sobering words: "The inscription on Sir Christopher Wren's tomb 
in St. Paul's Cathedral contains these famous words: 'If thou seek his 
monument, look about thee.' God forbid that this should ever become our 
epitaph. . . ." 



ANTHROPOLOGY 

Chairman, Robert E. Pace, Indiana State University 

Edward V. McMichael, 

Indiana State University, was elected Chairman for 1970 

ABSTRACTS 

Use of Historical and Archaeological Evidence to Reconstruct the 
Cherokee Past. Edward Dolan, DePauw University. — Archaeologists fre- 
quently use historical documents to identify remains of proto- and early 
historic sites. By combining archaeological and historical evidence, errors 
in interpretation of some local histories may be corrected. An example is 
discussed from the early historic Cherokee area. 

Exploratory Examination of the Oxendine Site. Robert E. Pace, Indiana 
State University. — The Oxendine Site is located along a 250-yard 
sand rise, adjacent to a vestigial finger of the Greenfield Bayou 
in Vigo County. Information derived from surface and controlled excava- 
tion shows it to be a late summer and fall campsite. Recovered materials 
suggest that it was occupied off and on by peoples of the Riverton Cul- 
ture, and later Woodland cultures. Except for brief Adena-Hopewell and 
Mississippian interludes, seasonal exploitation of an Oxendine type of 
microenvironment appears to represent a major form of adaptation along 
the middle Wabash Valley. 

NOTES 

Test Excavations At The Daughtery-Monroe Site (12-Su-13). Edward 
V. McMichael and Stephen J. Coffing, Indiana State University. — 
In late June and early July, 1969, the authors conducted test excava- 
tions at the Daughtery-Monroe Site, in northern Sullivan County, 
Indiana. The site is located on the east bank of the Wabash River, oppo- 
site Hutsonville, Illinois, and approximately 35 miles southwest of Terre 
Haute, Indiana. A 5-by-15-foot trench was excavated on the north side 
of this circular village and two 5-by-5-foot squares were dug on the east 
and west sides respectively. Below a 9-inch thick plow zone, a general 
village midden extended 15 to 20 inches below the surface, and below this 
eight pits were found. The latter were of two types: three basin-shaped 
shallow refuse pits; and five deeper, circular to oval, straight-walled 
pits, assumed to be used originally for storage, but also eventually used 
for refuse. Artifacts and non-artifactual debris were plentiful, including 
3267 pottery sherds of the Embarrass Ceramic Series, much animal bone, 
some worked bone, ground stone, and chipped flint. Of the pottery, 
simple stamped surfaces account for 53% of the sample; cordmarking 
26%; and plain surface 19%. Chipped stone included one Lowe Flared 
Stem projectile point and one triangular type; ground stone included 3 
grooved sandstone abraders, 1 unfinished pendant and a bi-pitted stone; 
and worked bone included 1 bi-pointed pin, 4 awl tips, 1 cut and per- 

57 



58 Indiana Academy of Science 

f orated deer toe bone, 6 turtle shell cup fragments, plus many unworked 
bone fragments. This village is definitely one of the La Motte Culture 
which is well represented, although largely uninvestigated, in the middle 
Wabash Valley. It is a later Woodland culture and probably dates between 
A.D. 500 to 1000, and in some way reflects prehistoric Southeastern 
United States cultural influences upon Wabash Valley prehistoric cultures. 

The Continued Excavation of the Van Nuys Site: A Probable Late Wood- 
land Occupation. Roger J. Ferguson, Ball State University. — The Van 
Nuys Site, Hn-25, (IAS-BSU) was recorded as a possible Woodland 
occupation site in 1967, and there was speculation that it was related to 
the Commissary Site, Hn-2, (IAS-BSU) which is believed to be a Late 
Woodland cemetery. The Van Nuys site is located in Henry Township, 
Henry County, Indiana. The 1969 excavation was continued north and 
west of the 1968 grid in order to further delineate the site. Various exca- 
vation techniques were used to ascertain the profile of the 1968 post hole 
assemblage. The depth of the excavation exceeded the 1968 depth by at 
least 12 inches in order to not miss evidence of multiple components. 

One hundred-fifty body sherds were excavated in 1968, and one rim 
sherd identified as Late Woodland and similar to Albee Mound material. 
There were but 15 fragmented sherds recovered in 1969. The 1969 exca- 
vation of the perimeter of the previous profile did not yield a single post 
hole between the 22-inch to 30-inch level. The 1968 excavation yielded 
approximately 500 post holes that ranged in size from 1 inch to 9 1 /£ 
inches in diameter. The post holes were found between the 22- and 24-inch 
level and had a 5-degree tilt in an easterly direction. The holes that were 
recorded in 1969 were found at the 10- to 20-inch level. They were 2 to 2V 2 
inches in diameter with no tilt. 

The hypothesis of multiple structures and a permanent settlement 
was formulated in the 1968 excavation. The findings of the 1969 excava- 
tion do not support this hypothesis. The lack of occupational rubbish and 
the limited number of post holes found at the 22- to 24-inch level cast 
doubt upon the 1968 hypothesis. The post holes found in 1969 are few in 
number and the interpretation that they actually are post holes is 
debatable. The vast number of post patterns found in 1968 were the same 
diameter (2 inches) and each had a 5-degree easterly tilt. It is unlikely 
that such posts could support a sizeable structure. They were shallow 
and none were tapered and since all holes tilt in an easterly direction, it 
is unlikely that they were supportive. The possibilities of their being the 
remnants of trees is offered. 

The decision prior to the excavation of Hn-25 in 1969 was (1) to 
delineate further the profile, and (2) to establish the components. This 
was adequately accomplished. It was recommended that no further 
excavation be undertaken at the Van Nuys Site but further site surveys 
should be continued to definitely establish an occupation area for the 
Commissary Site, Hn-2. 



Anthropology 59 

OTHER PAPERS READ 

Who Were the Oneota People of Minnesota, Iowa, and Wisconsin? 

Elizabeth J. Glenn, Ball State University. 

Environmental and Racial Factors in Ostoporosis: A Design for an 
Investigation. Anthony J. Perzigian, Indiana University. 



A Re-examination of the Question of the Middle 
Western Origin of the Delaware Indians 

Georg K. Neumann, Indiana University 



Abstract 

The locating of the traditional homeland of the Delaware Indians along the White 
River in Indiana aroused great interest not only of the historian and archaeologist but 
also of the layman of the Middle West. This interest culminated in the publication of the 
Walam Olum or Red Score (1) — an interdisciplinary examination of the migration legend 
of the Delaware, under the sponsorship of Eli Lilly in 1954. The excavation, twelve years 
later, of the Island Field site in Delaware with abundant skeletal material and the 
assigning of an approximate date to the site, provides substantiating evidence that the 
arrival of the tribe on the Atlantic coast can indeed be placed into the proto-historic 
period. 

During the past summer the writer had the opportunity to do field 
work with the help of four students — Richard D. Hill, Randall J. Mar- 
mouze, Turhon Murad, and David Parman — on the skeletal materials at 
the Island Field site, Kent County, Delaware. Also the known Delaware 
cranial series from the Montague site, and the known Nanticoke crania 
from the Townsend site, both in the collections of the U. S. National 
Museum, were re-examined. 

The Island Field site, 7-KF-17, is a Middle Woodland cemetery, dated 
about A.D. 900, which is being developed as a museum exhibit by the 
Delaware Archaelogical Board. To date, over 100 burials have been 
exposed and left in situ under the protection of a corrugated iron shed. 

The purpose of the field work was to remove the skulls of 16 of the 
skeletons — 8 male and 8 female — reconstruct, measure, and photograph 
them in order to determine whether this population could possibly be 
identified as ancestral to the Delaware tribe, which inhabited New Jersey 
and northern Delaware in historic times. If this were the case it would 
place the Delawares in the East about 500 years before the date sug- 
gested in the Walum Olum (1), the migration record of the Delaware 
and related Algonquian-speaking tribes. If, on the other hand, the affilia- 
tion of this population could be shown to be with the skeletal material 
from the historic Townsend site, date ca. 1400-1500, the Island Field 
population could be identified as Nanticoke, whose ancestors inhabited 
the Delmarva Peninsula since Archaic times — a circumstance that would 
confirm the Delaware migration account. 

Actually the latter proved to be the case, for all but one of the 16 
crania were clearly of the long-headed Lenid variety associated with 
the Algonquian-speaking coastal populations of the Middle Atlantic and 
New England states. The other, an individual who was buried in an 
extended position and accompanied by Late Woodland pottery, was 
Ilinid, like most of the Delaware crania from the Montague site on 
Minnisink Island. At the Townsend site, just south of the Island Field 
site, 9 of 12 males could be clearly classified as Lenid, 1 as Ilinid, and 
2 seemed to exhibit Muskogid affiliations, a variety that is wide-spread 

60 



Anthropology 61 

in the southern states. At the Montague site, in contrast, 7 of 10 males 
were Ilinid, and 3 Lenid; and of the 9 females, 7 were Ilinid, 3 Lenid, 
and 1 Muskogid. 

In conclusion, these findings substantiate an earlier migration of 
Algonquian-speaking people to the Atlantic coast, followed by a second 
migration with Central Algonquian relationships in proto-historic times. 
Since both are derived from the same gene pool and spoke languages of 
the same linguistic family, considerable overlapping is to be expected, 
but the isolation had evidently been of long enough duration to differen- 
tiate the two populations in a number of morphological characteristics, 
characteristics which can be used to distinguish most of their members 
from each other (2). 



Literature Cited 

1. Lilly, Eli (Ed.). 1954. Walam Olum or Red Score — The Migration Legend of the 
Lenni Lenape or Delaware Indians. Indiana Historical Society, Indianapolis. 379 p. 

2. Neumann, Georg K. and Turhon A. Murad. 1970. Preliminary Report on the Crania 
from the Island Field Site. Kent County, Delaware. Proc. Indiana Acad. Sci. 79 :69-74. 



Origins and Racial Affiliations of the Illinois Hopewell Indians 

King B. Hunter and Georg K. Neumann, Indiana University 

Abstract 

The present paper which constitutes a revision of an earlier preliminary report on a 
Middle Woodland population clarifies its origin, morphological changes, and relationships 
to neighboring contemporary groups. The study of the material based on a multivariate 
descriminant analysis demonstrates that the Hopewellian people of the lower Illinois Valley 
are primarily derived from a long-headed Early Woodland population of the Lenid variety 
native in the Great Lakes area since Archaic times. By A.D. 200 this population had 
undergone a number of changes, such as some brachycephalization, which characterized 
many of the later groups of the Middle Woodland period, designated as the Ilinid variety. 
Parallel changes occurred since Late Archaic times in the Red Ocher people and groups 
in the Middle Mississippi area. 

With the completion of a detailed morphological and metrical analy- 
sis of the skeletal materials from the Klunk Site in Calhoun County, 
Illinois, and comparisons with Fulton County and Ohio Hopewell crania 
of the same culture, we are now in a position to answer some questions 
of the origins, racial composition, physical changes, and perhaps the fate 
of the people that have been archaeologically grouped into the Hope- 
wellian category. 

The material from the Klunk Site is of special importance since it 
represents a complete population sample of over 300 burials, extending 
from an Archaic to a Late Woodland level, spanning a possible 800 year 
period. 

As a background, the broad outline presented by Neumann (6) will 
be followed here. The most ancient populations in eastern North America 
which are reasonably well documented appear in early Archaic times, as 
early as 4000 B.C., and perhaps earlier in a few cases. Two widely dis- 
tributed and quite homogeneous populations dominate the whole of the 
eastern United States at this time, both seeming to have considerable 
antiquity, and showing perhaps, a generalized common ancestor. In the 
northern area, primarily associated with a Great Lakes area Archaic 
cultural pattern, is the Lenid variety, described elsewhere by Neumann 
(6). An example is Old Copper. The area south of the Ohio Valley is 
populated by groups which may easily be distinguished from the Lenids, 
and which are generally associated with the southern riverine Shell 
Mound Archaic cultural pattern. These populations are consistently of 
the Iswanid variety, described by Neumann. The largest sample of this 
variety to date comes from Indian Knoll, which has a radiocarbon date 
of 3352 B.C. ± 300 years (3). The earliest material from Modoc Rock 
Shelter is also Iswanid. 

Neumann (6) proposes a differentiation from Lenid to the Ilinid 
variety, and from Iswanid to the Muskogid variety, beginning in Medial 
Archaic times, perhaps as early as 2500 B.C. or earlier. An Ilinid popu- 
lation is associated with Glacial Kame. This process of Lenid to Ilinid 
differentiation may also be observed in Classic Hopewell times. The se- 

62 



Anthropology 63 

quence of Modoc Rock Shelter reflects the progressive differentiation 
from Iswanid to Muskogid. The Muskogid variety, especially, is quite 
widespread by Terminal Archaic, and Early Woodland times, with the 
Iswanid becoming marginal or peripheral by Early Woodland times, 
migrating both to the east and northward. 

During Terminal Archaic and Early Woodland times the Muskogids 
apparently enjoyed a considerable extension of their range, moving up 
the Ohio Valley into the New York and eastern Great Lakes area, and 
into the Illinois Valley. In the Illinois Valley they are associated with 
Red Ocher and Morton, and in the Northeast, with Laurentian and Point 
Peninsula. The Adena people of Ohio were probably also originally 
Lenid, possibly receiving some Muskogid admixture as they spread into 
Kentucky. In general, the Lenids and Ilinids appear to be peripheral 
to the distribution of these groups during this time. With the appearance 
of Hopewell in Middle Woodland times, however, the Lenids once again 
became the dominant population north of the Ohio River and in the 
Illinois Valley (4). The Middle Woodland population in the Southeast 
is predominantly Muskogid, associated with such cultures as Copena 
and Marksville, while the differentiated Ilinid populations become 
dominant in the north. A final northern expansion of the Muskogid popu- 
lation during the Hopewell breakdown in Late Woodland times takes 
place in connection with the spread of Middle Mississippi culture. In 
most of the cases discussed above, little admixture is indicated. As a 
consequence, the populations associated with these culture complexes 
ought to be reasonably homogeneous. 

The material from the Klunk mound group is rather significant when 
considered in this framework, in that it offers evidence which may be 
used to test certain of the foregoing hypotheses. The site may be briefly 
described as follows. The mounds are located on the western bluff over- 
looking the Illinois River, in Calhoun County. Archaeological evidence 
in the form of log tombs, subfloor burial pits, zoned Hopewell pottery, 
Hopewell stamped rocker-dentate pottery, and so on, indicates that the 
main body of the material is Classic Illinois Hopewell. The burials as- 
sociated with this culture pattern constitute the largest sample of the 
Illinois Hopewellian population extant. One Hopewellian mound overlies 
a low Archaic or Early Woodland mound, which may have Red Ocher 
affiliations. This material offers an interesting comparison to the Hope- 
well series, and to Early Woodland populations from other sites. Finally, 
a few intrusive Jersey bluff burials are found in the mounds. 

The investigation of the population associated with Classic Hopewell 
at the Klunk site yields an interesting and significant statistical pattern. 
Although it is primarily Lenid in character, certain morphological 
characteristics diverge from the expected Lenid pattern, and approach 
those of Ilinid groups. 

There is no indication of any tendencies toward Muskogid character- 
istics, ruling out the possibility of significant admixture from this 
population. This is entirely consistent with Neumann's (6) proposal that 
Classic Hopewell is a period of Lenid to Ilinid differentiation. 



64 Indiana Academy of Science 

Furthermore, those burials which are associated with Bluff culture 
ere consistently Ilinid, and compare very nicely with the material from 
the Schild site. This last is under preparation in the same laboratory, 
and promises to be a large and very homogeneous sample of the Jersey 
Bluff Late Woodland population. 

The Archaic Early Woodland material which has been examined and 
given a radiocarbon date of 908 B.C. comprises a small but very homoge- 
neous sample that is unquestionably Muskogid. This material is tenta- 
tively classified as Red Ocher. This is not particularly surprising since 
it is during this period that a differentiation of Iswanid to Muskogid is 
hypothesized. Consequently, some Red Ocher groups would be expected 
to be clearly Iswanid, some clearly Muskogid, and some in an inter- 
mediate condition between the two. 

Conclusions 

1. The hypothesis that the Illinois Hopewell population is essentially 
Lenid is strengthened by a preliminary investigation of the Hopewellian 
skeletal material from the Klunk Mound group. 

2. The proposed Lenid to Ilinid differentiation during Classic Hope- 
well times is also substantiated in this investigation. 

3. The Lenid population associated with this Illinois Hopewell 
group appears to be essentially identical to the population associated 
with Ohio Hopewell, which is contemporaneous. 

4. An examination of the presumed Red Ocher burials indicates 
a definite Muskogid association, which is expected. 

5. The Late Woodland burials recovered are consistently Ilinid, 
and compare very well to other populations of the same period. 

6. A northern source for certain of the characteristics that influ- 
ence the development of Hopewell out of the indigenous Early Wood- 
land culture is indicated by the appearance of certain Point Peninsula 
ceramic characteristics, such as dentate stamping. This helps to explain 
the predominance of Lenids in Hopewell, since their distribution at this 
time was to the north and west of the Illinois and Ohio Valleys. 

Literature Cited 

1. Deuel, Thorne. 1958. American Indian Ways of Life. Illinois State Museum, Spring- 
field. 80 p. 

2. , (Ed.) 1952. Hopewellian Communities in Illinois. Illinois State Museum, 

Springfield. 271 p. 

3. Criffin, James B. 1952. Culture Periods in Eastern United States Aehaeology, p. 352- 
364. In J. B. Griffin [ed.l Archaeology of Eastern United States. Univ. Chicago Press, 

Chicago. 597 p. 

4. . 1958. The Chronological Position of the Hopewellian Culture in Eastern 

U. S. Anthropology. Papers Mus. of Anthropol., Univ. of Michigan No. 12. Univ. 
of Michigan, Ann Arbor. 

5. Neumann, Ceorg K. 1952. Archaeology and Race in the American Indian, p. 13-34. in 
J. B. Griffin [ed.l Archaeology of Eastern United States. Univ. Chicago Press, Chicago. 

6. . 1960. Origins of the Indians of the Middle Mississippi Area. Proc. Indiana 

Acad, of Sci. 60:66-68. 



Some Possible Causes for Late-Pleistocene Faunal Extinctions 

William H. Adams, Indiana University 

Abstract 

While Indiana has thus far not yielded a stratified early man site, the distribution 
of early type projectile points indicates that not only was he in Indiana but that he 
hunted the Pleistocene megafauna and contributed to their extinction. This paper examines 
the causes for these faunal extinctions and why, ultimately, the blame rests on man. 

Beginning around 15,000 years ago there was a radical reduction in 
the Pleistocene megafauna. By 9,000 B.P. most had become extinct. These 
late Pleistocene extinctions were widespread both geographically and 
interspecifically. They involved areas of the world which were in equal 
stages of human development: nomadic hunting and gathering. These 
extinctions were limited to land mammals weighing over 50 kg, the 
majority being herbivores and their predators (3). Many of these 
animals have been found in Indiana associated with a tundra-like environ- 
ment. Since elsewhere in North America under the same conditions these 
fauna have been found with Clovis projectile points, and Clovis points 
have been found throughout Indiana, we should eventually find sites 
which yield the same associations. 

A number of forces, biological, geological, and environmental, could 
have been responsible for these extinctions. Their interrelationships 
reveal that no single cause was totally responsible. 

The biological forces at work are complex. Natural selection will 
eliminate the unfit, leaving only the best adapted organisms in an en- 
vironment. Fitness would include: birth of offspring at the right time, 
length of maturation, adaptation to climate, feeding methods, and 
predator defense. 

If a species fails to adjust the time of the birth of its progeny to 
meet changing climatic conditions, it may not survive. During periods 
of lengthening winters, species which cannot adjust their birthing will 
be selected against; while those that can will have a better chance for 
survival. Thus a gradual change in climate should cause variation in the 
time of birth unless another factor overrides the selection process. 

Two such factors exist; length of gestation and the size of the 
individual. The longer the time between mating and birth, the less the 
assurance of environmental conditions into which the young will be born. 
In species where mating occurs at the first sign of improving environ- 
mental conditions (as in the spring warm-up) and birth shortly after, 
there will be a better chance of survival than those species that have 
long gestation periods. 

Since larger animals have longer gestation periods, and greater 
body size usually requires proportionately more time for maturity to 
be reached, a change in climate would have greater impact upon the 
larger animals. 

65 



66 Indiana Academy of Science 

An organism must be able to adapt to a new or changing environ- 
ment. During this period, between 15,000 and 9,000 B.P., the climate 
was slowly changing and with it the available food supply. Thus, in order 
to survive, the existing fauna had two courses available; a change in 
diet, or migration. If the species were physically able, they would follow 
the changing ecological zones, but if trapped by local physical features 
(such as mountains or rivers) they would have to change their diet. For 
some species, in some areas, the ability to migrate would be cut off by 
the new vegetation cover which, unless they were preadapted, would be 
as formidable a barrier as a mountain. 

There exists in nature a predator-prey balance that could, if upset 
enough, result in the extinction of one or both. Some of the causes which 
could upset this balance include: extinction of the major predators, en- 
vironmental forces which cause gregarious fauna to scatter into small 
groups less able to protect themselves and their young, change in the 
individual method of predator defense, and the introduction of a highly 
efficient predator. 

By approximately 15,000 the saber-toothed cat, Smilodon, was ex- 
tinct. If Smilodon played an important role in holding the herbivore 
population to a level compatible with available vegetation, then what 
would be the result if this predator pressure were released? If the 
extinction of Smilodon was gradual, other predators would increase in 
numbers, but since these smaller predators would not be able to fill 
Smilodon's niche completely, the result would be an increase in the 
population of herbivores. A large increase would cause the cycle of 
overgrazing — starvation — low population — vegetation recuperation — over- 
population — ad infinitum. This cycle would occur in a more severe man- 
ner if Smilodon were exterminated in a short period by an epidemic 
disease and thus, lacking that predator's pressure, the herbivore popu- 
lation would increase to numbers which could not be maintained by the 
food supply. 

A vegetation change which resulted in the desired food plants becom- 
ing scarce would limit the number of animals and would be selective 
against large herds, scattering them. This would have disadvantaged 
species that were dependent upon group defense. Those species that 
could fight or outrun their foes increased their chances for survival. 

The addition of an efficient predator to an environment usually has 
wide effect. Naked man as a predator was not very efficient. He lacked 
the claws, teeth, and stamina of a predator. With tools he increased the 
efficiency but not until he possessed a projectile point did he become a 
very efficient killer. Man has been a significant force in the selection of 
some fauna over others. Man's ability to kill a particular animal at a 
certain time varied with his ingenuity; but, in general, he tended to kill 
the smaller fauna. Until the advent of the projectile point, he was a 
selective force towards larger and harder to kill animals since his limited 
hunting ability forced him to select the smaller individuals. With the 
projectile point, the larger animals were more easily within his grasp 
and the forces of selection favored a smaller and fleeter fauna. 



Anthropology 67 

If we assume that climatic conditions were about the same during 
this period (15,000-9,000 B.P.) as during the other post-glacial periods, 
then climatic changes, per se, cannot be the cause of these extinctions. 
These climatic conditions were important as contributing factors. By 
limiting the land area by covering it with glaciers and changing the 
flora, they no doubt decreased the animal population. Since conditions 
in post-glacial periods were similar, then the late Pleistocene post-glacial 
faunal extinctions must be the result of an additional factor not present 
in earlier periods: MAN. 

Man has been in the New World for over 35,000 years, yet until 
about 11,000 years ago did not possess a stone projectile point. Before 
that date he used bone points or none at all. With the addition of the 
highly efficient fluted Clovis point man could become the major force in 
the extinctions. While the origin of the Clovis point is still unknown its 
effects are widely known. Either by migration or diffusion the Clovis 
points had spread in only a few decades throughout most of the habitable 
portions of North America. These points reflect not only the game which 
were killed but also the distribution of the various species at the time 
they were killed (W. H. Adams, unpublished data). 

Dorwin lists 87 Clovis-like fluted points found in Indiana (2). All 
were surface finds and were distributed throughout the state. The Ohio 
River area from Posey County upstream to Clark County accounts for 
1/3 of the points. Another 1/3 are found in the remaining southern half 
of Indiana. Thus there is a decrease in the frequency as one moves 
northward. For the later Folsom points a proportionately greater num- 
ber (11/26) are found in the northern half of the state than were 
Clovis points. This would indicate that at least two migrations of 
different cultures occurred; one sometime after 15,000 B.P. and the 
other sometime afterwards as the tundra was leaving Indiana, since 
the animals with which Clovis and Folsom are usually associated, 
mammoth, mastodon, bison, would have been available at that time 
in Indiana. We can assume that the hunting of those animals was the 
cause for the presence of those points in Indiana, since man would 
have followed these megafauna northward across the tundra until the 
end of the tundra — the ice sheet — was reached; also that the northern- 
most distribution of fluted points marks the place where man and animal 
could go no further north and soon killed off the few remaining mega- 
fauna. This line, the Mason-Quimby Line, marks the end of a biological 
era and technological stage. 

However, man is often given too much credit for these extinctions. 
While his many methods of killing animals, such as snares, pitfalls, 
stampedes, and fire drives could kill off all available animals, it was only 
with a projectile point that man could affect a species quickly enough 
to cause extinction, in the period under examination here. 

Perhaps the most overrated possible causes for extinction has been 
the fire drive. Whether or not the fire drive was used at an early date 
in the New World, it has been pointed out that, at least in the savanna in 
southern Africa's Kruger National Park, the overall effect of fire 



68 Indiana Academy of Science 

would be helpful to the grazer. There Brynard (2) found that when 
fire was prevented the amount of brush increased and left smaller areas 
of grazing land. The same situation should have occurred on the North 
American savanna. By using fire drives man would have aided the 
grazers, mammoth and bison, and hurt the browsers, sloth and mastodon. 



Conclusions 

A combination of biological and environmental forces reduced faunal 
populations during the glacial and post-glacial times. By 9,000 B.P. 
many of those species were extinct. The changing climate was not the 
cause since in previous glacial periods there had been no extinctions and, 
at the time of these major extinctions, the environment was actually 
improving for these species. The only differences between this post- 
glacial period and the others was man, not his presence, but by his 
possession of weapons which could affect these large animals. These 
late-Pleistocene extinctions would appear to have no single cause but 
instead to be the result of an untimely combination of factors. Thus not 
only was man in Indiana in the period between 15,000 and 9,000 B.P. 
but also he was most likely a significant factor in the extinction of 
several Pleistocene megafauna. For that reason we must concentrate 
our search for early man sites in Indiana and the Midwest which will 
yield artifacts which are in association with those extinct fauna. 



Literature Cited 

1. Brynard, A. M. 1964. The influence of veldt burning on the vegetation and game of 
Kruger National Park. In D. H. S. Davis [ed.] Ecological Studies in Southern Africa. 
The Hague. Junk. 

2. Dorwin. John T. 1966. Fluted Points and Late-Pleistocene Geochronology in Indiana. 
Prehistory Research Series. Vol. 4, No. 3. Indiana Historical Society, Indianapolis. 

3. Martin, P. S., and H. E. Wright (Eds.). 1967. Pleistocene Extinctions, The Search 
for a Cause. Yale University Press, New Haven. 



Preliminary Report on the Crania from the Island Field Site, 
Kent County, Delaware 

Georg K. Neumann and Turhon A. Murad, Indiana University 

Abstract 

The report deals with a study of the skeletal population from the Island Field Site, 
carried out jointly by the two writers during the summer of 1969. It includes demo- 
graphic data on 95 burials of the late Middle Woodland group that inhabited the Del- 
marva Peninsula about A.D. 900 according to typological artifact dating. Sixteen of the 
crania, those of 8 males and 8 females, were reconstructed and measured. The report 
deals with a brief description of this material, pointing out the relationships of this 
population to that of other sites in the middle Atlantic States area. 

The Island Field Site (7K-F-17) is located near the mouth of the 
meandering Murderkill, at South Bowers, Delaware, on a sandy knoll 
only a few hundred yards from Delaware Bay. On a typological basis 
the Island Field Site has been given the relative date of about A.D. 900. 
The culture represented at this site is obviously of the Middle Woodland. 
However, its placement into one of the subdivisions of this broad period 
presented some difficulties. For that reason the site has been assigned 
to the Webb Phase — a newly established phase of the Middle Woodland 
complex denned at the Island Field Site. 

Skeletal material was first encountered at the site in the 1930's 
when, during the construction of a nearby road, the knoll was used for 
fill. At that time the material was sealed in kegs and reburied in the 
surrounding marsh by the workmen. As yet, that material has not been 
relocated. It was not until 1953 that the archaeological importance of 
the site was recognized by an amateur archaeologist, Frank Austin, who 
with the aid of the Sussex Society of Archaeology, recovered from a 
shell pit artifacts characteristic of the Late Woodland Period. Since 1966 
there has been intensive excavation carried on at Island Field by the 
Delaware Archaeological Board under the direction of Mr. Ronald A. 
Thomas, State Archaeologist. Nearly 90 burials have been uncovered at 
the site with no fewer than 7 burial types present, although the ma- 
jority of the burials are either flexed or semi-flexed. Extended burials, 
two types of cremation, in situ and redeposited, disarticulated and skull 
burials, and both single and mass burials are found at Island Field. 

Many of the graves contain grave offerings. Such materials as bone 
and antler have been used to produce awls, needles, and harpoons while 
stone artifacts have been encountered as in the case of waste flint, 
arrowheads, knives, stone pipes, celts, and pendants. Both clay and 
steatite pipes have been recovered along with clear quartz pebbles, mica, 
a conch shell cup, shell beads, two shark teeth, and a bowl made from a 
human skull. 

Whereas many of these artifacts are not uncommon to the region, 
there is an interesting array of foreign materials, whose styles and 
composition indicate contact between the Island Field Site and such areas 
as New England, the Midwest and the Southeast. 

69 



70 Indiana Academy of Science 

At the time the data were collected, the skeletal population from 
this site consisted of 27 adult males, 24 adult females, 11 infants (age 3 
or below), 10 children, 12 known but thus far unexcavated individuals, 
and 5 adults of undetermined sex. The poor condition of the material 
combined with the desire of the Delaware Archaeological Board to keep 
the material in situ for a possible future museum, allowed only the best 
skulls to be reconstructed. This resulted in the measurement, and ob- 
servation of 16 adult crania — 8 males and 8 females. From the analysis 
of the 8 adult male crania it has been determined that the population is 
primarily Lenid with the exception of one individual which has been 
classified as Ilinid. 

Generally, the Island Field population displays a large degree of 
muscularity, is quite dolichocephalic or long-headed, with an average 
cranial index of 71.39. The average length-height index is 73.35, placing 
the group within the ortho-cranial index class. In no instance was there 
a case of platybasia; and the position of basion is high. Face size is 
medium with a tendency toward being large. The average total facial 
index is 91.12, or leptoprosopic in character. 

The superior facial index is 76.96, or mesene. The orbits are rhomboid 
in 62.5% of the individuals and oblong in the remaining 37.5%. Their 
inclination tends to be small. The overall orbital index is 77.43, or meso- 
conch. The nasal bones are medium in size and the root is medium to 
high. Nasal bridge height is also high in % of the individuals observed. 
The nasal index is 49.14, or mesorrhine in classification. The height of 
the palate is from high to very high while the average maxillo-alveolar 
index is 141.41, or mesuranic in character. The average mandibular index 
is 86.42. However, the size of the mandible is from medium to large, 
each displaying a frequency of 50%. Chin projection is usually neutral. 

The purpose of this summer's field work with the skeletal material 
from the Island Field Site was to remove the skulls of 16 of the skeletons 
of adults — 8 male and 8 female — reconstruct, measure, and photograph 
them in order to determine whether this population could possibly be 
identified as ancestral to the Delaware tribe, which inhabited New Jersey 
and northern Delaware in historic times. If this were the case it would 
place the Delawares in the East about 500 years before the date sug- 
gested in the Walam Olum, the migration record of the Delaware and 
related Algonquin-speaking tribes. If, on the other hand, the affiliation 
of this population could be shown to be with the skeletal material from 
the historic Townsend site dated ca. 1400-1500, the Island Field popula- 
tion could be identified as Nanticoke, whose ancestors inhabited the 
Delmarva Peninsula since Archaic times — a circumstance that would con- 
firm the Delaware migration account. The morphological data seem to 
substantiate the latter. The metric, indicial, and morphological descrip- 
tion of the crania is given in Tables 1 through 3 for comparative 
purposes. 



Anthropology 

Table 1. Cranial measurements — means for 8 males. 



71 



Measurement 



Abbreviation 



Average 



Cranial Vault 

Cranial module 

Mean thickness of left parietal 
Glabello-occipital length 
Maximum breadth 
Minimum, frontal 
Frontal chord 
Basion bregma height 
Porion-apex height 
Basion-porion height 
Cranial base length 

Face 

Total facial height 
Upper facial height 
Total facial breadth 
Midfacial breadth 
Internal biorbital breadth 
Subtense biorbital breadth 
Biorbital breadth 
Ant. inter orbital breadth 

Nasal structure 
Nasal breadth 
Nasal height 
Dacryal chord 
Dacryal subtense 
Minimum nasal breadth 
Subtense nasal breadth 
Breadth of nasal bridge 
Height of nasal bridge 

Orbit 

Left orbital height 

Left orbital breadth (mf ) 

Left orbital breadth (d) 

Dental arch and profile 

Maxillo-alveolar breadth 
Maxillo-alveolar length 
Facial length (pr) 
Facial length (alv. pt. ) 

Angles 

Facial profile angle 
Midfacial profile angle 
Alveolar profile angle 
Gonial angle 

Mandible 

Length of mandible 

Bicondylar breadth 

Height of mandibular symphysis 

Biangular breadth 

Minimum ramus length 



CM 

TP 

L 

B 

MF 

FC 

H 

PAH 

BPH 

LB 



TFH 

UFH 

TFB 

MFB 

IOB 

SIOB 

BOB 

AIM 



NB 

Nil 
DC 
DS 

MN 
SMN 
BNB 
HNB 



LOH 

LOBM 

LOBD 



MB 
ML 
FL 
FLA 



FP< 
MP< 
AP< 

G < 



LM 
BCB 
SH 
HA 

HI, 



140.2 

5.4 

191.5 

136.5 

93.1 
116.4 
141.0 
118.1 

26.1 
107.3 



125.1 
73.9 

137.5 
97.5 
97.6 
20.6 
99.3 
19.4 



25.5 
52.6 
22.2 
13.8 
9.3 
5.0 
59.8 
24.8 



32.8 

4 2.4 



63.6 

55.6 

101.0 

99.0 



84.2° 

88.5° 

69.8° 

124.8° 



108.7 
124.9 

37.9 
107.3 

34.9 



72 



Indiana Academy of Science 

Table 2. Cranial indices — means for 8 males. 



Indices 



Abbreviation 



Average 



Crania] Vault 
Cranial 
Length-height 
Breadth-height 
Mean height 
Length-auricular 
Flatness of cranial base 
Trans. Fronto-parietal 
Frontal 

Face 

Total facial 

Upper facial 

Midfacial 

Trans, cranio-facial 

Zygo-frontal 

Fronto-mandibular 

Zygo-mandibular 

Facial flatness 

Ant. interorbital 

Nasal 

Nasal 

Nasal root height 
Nasal bone height 
Nasal bridge height 

Orbit 

Left orbital (mf ) 
Left orbital (d) 

Dental Arch 

Maxillo-alveolar 

Mandible 

Mandibular 



B/L 


71.39 


H/L 


73.35 


H/B 


102.76 


H/(L + B/2) 


85.48 


PAH/L 


61.79 


BPH/H 


18.56 


MF/B 


68.24 


MF/M'F 


80.75 


TFH/TFB 


91.12 


UFH/TFB 


53.77 


UFH/MFB 


76.96 


TFB/B 


100.82 


MF/TFB 


70.62 


BA/MF 


114.32 


BA/TFB 


77.51 


SIOB/IOB 


21.45 


AIB/BOB 


19.27 


NB/NH 


49.19 


DS/DC 


62.28 


SMN/MN 


55.98 


HNB/BNB 


41.76 


LOH/LOB 


77.43 


LOH/LOBD 


82.31 



MB/ML 



LM/BCB 



114.41 



86.42 



Anthropology 

Table 3. Morphological observations — percentages and frequencies 





sm* 


m 


1 


vl 






Muscularity 


(0) 
It 


(0) 
m 


87 (7) 
h 


13 (1) 






Weight 


(0) 

sm 


71 (5) 
m 


29 (2) 

1 








Face size 


(0) 


62 (5) 


38 (3) 










med 


div 


con 


tor 






Brow ridge shape 


(0) 


100 (8) 


(0) 


(0) 








tr 


sm 


m 


1 


vl 




Brow ridge size 


(0) 
sm 


12 (1) 
m 


25 (2) 

1 


50 (4) 
vl 


13 (1) 




Glabella 


12 (1) 


38 (3) 


38 (3) 


12 (1) 








vlo 


lo 


m 


hi 


vhi 




Frontal height 


(0) 


13 (1) 


75 (6) 


12 (1) 


(0) 






n 


bul 


si 


m 


P 


vp 


Frontal slope 


(0) 
sm 


(0) 
m 


13 (1) 

1 


87 (7) 


(0) 


(0) 


Post-orb. constr. 


75 (6) 
sm 


25 (2) 
m 


(0) 

1 








Frontal eminences 


75 (6) 
n 


25 (2) 
sm 


(0) 
m 


1 






Med. front, crest 


75 (6) 


25 (2) 


(0) 


(0) 








n 


sm 


m 


1 


vl 




Sagittal elev. 


37 (3) 

sm 


50 (4| 
sm 


37 (1) 

m 


(0) 

1 


(0) 




Parietal eminences 


(0) 
fl 


100 (8) 
m 


(0) 

1 








Temp, fullness 


25 (2) 

sm 


38 (3) 
m 


37 (3) 

1 


(0) 
vl 






Mastoids, size 


(0) 


50 (4) 


25 (2) 


25 (2) 








n 


sm 


m 


P 


vp 




Occip. curve 


(0) 
hi 


(0) 
m 


25 (2) 

1 


62 (5) 


13 (1) 




Occip. position 


87 (7) 


13 (1) 


(0) 










bun 


na 


m 


wi 






Occip. breadth 


13 (1) 


87 (7) 


(0) 


(0) 








n 


srn 


m 


P 






Lamb, flattening 


(0) 
sm 


13 (1) 
m 


50 (4) 
1 


37 (3) 
vl 






Elev. occ. condyles 


(0) 


14 (1) 


57 (4) 


29 (2) 








lo 


m 


hi 


vhi 






Basion 


(0) 
n 


71 (5) 

s 


15 (1) 
m 


14 (1) 

1 






Platybasia 


100 (8) 
vsm 


(0) 
sm 


(0) 
m 


(0) 

1 






Styloids 


25 (2) 
sm 


50 (4) 
m 


13 (1) 
1 


12 (1) 






Lacerate foramen 


17 (1) 

sm 


17 (1) 
m 


66 (4) 
1 








Glenoid fossa 


(0) 
sm 


25 (2) 
m 


75 (6) 

1 








Post-glen, process 


(0) 


75 (6) 


25 (2) 










th 


m 


tk 


vtk 






Tympanic plate 


87 (7) 
s 


13 (1) 
m 


(0) 
1 


(01 






Petrous depr. 


(0) 


50 (4) 


50 (4) 










obi 


rhm 


sq 


ell 


id 





74 



Indiana Academy of Science 



Table 3. (Continued) 





sm* 


m 


1 


vl 




Orbit shape 


38 (3) 


62 (5) 


(0) 


(0) 


(0) 




n 


sm 


m 


P 


vp 


Orbit inclin. 


25 (2) 


38 (3) 


37 (3) 


(0) 


(0) 




i) 


si 


m 


dp 




Suborbit. fossa 


(0) 


63 (5) 


37 (3) 


(0) 






sm 


m 


1 


vl 




Zygomatic size 


25 (2) 
sm 


63 (5) 
m 


12 (1) 

1 


(0) 




Zygo. lat. proj. 


(0) 
sm 


50 (4) 
m 


50 (4) 

1 






Zygo. ant. proj. 


(0) 


87 (7) 


13 (1) 








lo 


m 


hi 






Zygo. bone ht. 


(0) 


100 (8) 


(0) 








sm 


m 


1 






Size of nasals 


(0) 


100 (8) 


(0) 








lo 


m 


hi 


vhi 




Nasal root height 


13 (1) 


50 (4) 


37 (3) 


(0) 






srn 


m 


1 


vl 




Nasal root breadth 


37 (3) 


38 (3) 


25 (2) 


(0) 






lo 


m 


hi 


vhi 




Nasal bridge height 


17 (1) 
sm 


17 (1) 
m 


66 (4) 

1 


(0) 




Nasal bridge breadth 


50 (3) 


50 (3) 


(0) 








str 


cone 


ccv 


cvx 




Nasal profile 


(0) 


(0) 


100 (5) 


(0) 






n 


sm 


m 


dp 




Nasion depr. 


13 (1) 


62 (5) 


25 (2) 


(0) 






par 


hyp 


ell 


sU 


III 


Palate shape 


50 (4) 


(0) 


13 (1) 


37 (3) 


(0) 




lo 


m 


hi 


vhi 




Palate height 


(0) 


(0) 


38 (3) 


62 (5) 






n 


sm 


m 


1 




Inframax. notch 


33 (2) 


33 (2) 


17 (1) 


17 (1) 


r 




sm 


m 


1 


vl 




Mandib. size 


(0) 


50 (4) 


50 (4) 


(0) 






nb 


wb 


int 


med 




Chin form 


25 (2) 


75 (6) 


(0) 


(0) 






neg 


neu 


sm 


m 


1 


Chin proj. 


(0) 


87 (7) 


(0) 


13 (1) 


(0) 




ti 


sm 


m 


1 




Gonial eversion 


50 (4) 


37 (3) 


13 (1) 


(0) 





* Abbreviations: bul bulging, bun bun-shaped, ccv concavo-convex, con continuous, 
cone concave, cvx convex, div divided, dp deep, ell elliptical, fl flat, h heavy, hi high, hyp 
hyperbolic, int intermediate, 1 large, lo low, It light, 1U large U-shaped, m medium, med 
median, n none, na narrow, nb narrow-bilateral, neg negative, neu neutral, obi oblong, p 
pronounced, par parabolic, id round, rhm rhomboid, si slight, sm small, sq square, str 
straight, sU small U-shaped, th thin, tk thick, tor torus, tr trace, vhi very high, vl very 
large, vlo very low, vp very pronounced, vsm very small, vtk very thick, wb wide-bilateral, 
wi wide. 



Literature Cited 



1. Lilly, Eli (Ed.) 1954. Walam Olum or Red Score — The Migration Legend of the 
Lenni Lenape or Delaware Indians. Indiana Historical Society, Indianapolis. 379 p. 



Preliminary Report on the Excavation of the 

"Great Mound" at Mounds State Park in 

Madison County, Indiana 

Kent D. Vickery, Indiana University 



Abstract 

A summary of two season's excavations of the "Great Mound" at Mounds State Park is 
presented. Two major phases in the construction of the mound were apparent. The primary 
mound was a "platform" consisting of three superimposed burned clay floors, each covered 
with a layer of ash. Over this had been placed a capping of earth which covered a subfloor 
log tomb adjacent to the primary mound. Interpretations are given concerning the pres- 
ence of two distinct post hole patterns related to the mound, and the results of a test 
trench in another mound are summarized. The "Great Mound" is compared with other 
excavated "sacred circles" and its chronological and cultural relationships are discussed. 



Introduction 

The earthwork complex at Mounds State Park near Anderson, Indi- 
ana, has long been recognized as one of the more unusual archaeological 
sites in the Ohio Valley. Within the park are found five circular enclo- 
sures; two panduriform or "fiddle-shaped" enclosures; one earthwork 
shaped like a figure-8 open at both ends; and one rectangular enclosure. 

The largest and best preserved of these earthworks is a circular 
enclosure known as the "Great Mound." It consists of an embankment 
averaging 6 feet in height; an interior ditch; an entranceway to the 
south; and a small mound about 45 feet in diameter on the central plat- 
form. 

During the first field season, i a contour map was made of the "Great 
Mound," and most of the mound on the central platform was excavated. 
The excavation of this mound was completed during the second season, 
after which a bulldozer was used to clear the topsoil from the sur- 
rounding central platform. This revealed a number of post holes in a 
roughly circular pattern. The final project of the season was the exca- 
vation of a test trench in a small mound on the western end of the 
larger of the two panduriform earthworks. 



Mound Structure 

Two major phases in the construction of the mound were apparent. 
The primary mound was a "platform" consisting of three superimposed 
burned clay floors, each covered with a layer of ash. Over this had been 
placed a capping of earth which covered a subfloor log tomb adjacent 
to the primary mound (4, C. F. White, unpublished data). 



1 The excavations at Mounds State Park were directed by Claude F. White in 1968 and 
by Kent D. Vickery in 1969. The project was financed by the Indiana Department of 
Natural Resources with the cooperation of the Glenn A. Black Laboratory of Archaeology, 
Indiana University. 

75 



76 Indiana Academy of Science 

The primary mound platform was oval and measured about 25 feet 
by 28 feet. It was underlain by a prepared floor of fine-grained silt, 
which was probably obtained from the nearby White River. A depression 
had been excavated into subsoil for the reception of this silt layer, but 
the fact that it was found at a higher elevation than the subsoil in the 
surrounding central platform suggests that the entire mound may have 
been built on a natural knoll. 

The thickness of the platform and its underlying silt layer was about 
2.25 feet. It was relatively flat, but terminated at the edges in a wedge- 
shape, thus indicating that each successive layer of burned clay and ash 
covered an area slightly less extensive than the one below it. 

The ash covering the upper two burned clay layers was white and 
relatively "pure," a condition which could have been caused by total 
incineration of the material burned or the intentional removal of foreign 
matter such as charcoal and cremated bone fragments. The earth of 
which the upper two burned clay layers was composed was relatively 
soft in consistency and burned to a dull red color. The lower burned 
clay floor was generally level; of uniform thickness; and was baked very 
hard throughout. This suggests that intense fires had been built re- 
peatedly over its surface. The lower ash layer was dry and compacted, a 
condition which could have been brought about by the deposition of 
earth on top of it, thus sealing it off and inhibiting the percolation of 
water down to it. The compaction of the ash may be explained by the 
overlying weight of two more layers of burned clay and ash, as well 
as the final mantle of earth constituting the mound capping. 

Occasional bands of hard black burned material, mostly ash, were 
noted on the lower floor of the platform. This may indicate that burn- 
ing took place in a reducing atmosphere, thereby resulting in incomplete 
combustion. The nature of the bottom burned clay layer, however, sug- 
gests sustained firing in the open and complete combustion. It is likely 
that fires were built on this floor for some period of time, and that the 
surface was scraped clean of ash and other debris periodically. During 
the final burning, earth was thrown over the platform, thereby causing 
the fire to smolder and turn the earth and ash black in spots. The dirt 
thus deposited then became the middle burned clay layer. 

The original function of the platform is unknown, but Warren K. 
Moorehead speculated that it might have been a "dance floor" when an 
auger test made by Moorehead and Glenn A. Black in 1931 disclosed 
". . . an 8 inch ash and burned earth bed . . . [with] a possible trace of 
calcined bone in the ash removed" (2). Nothing was noted in the struc- 
ture of the platform that could either confirm or refute Moorehead's 
observation. 

Another theory is that the primary mound platform served as the 
central crematorium for the entire earthwork complex, in which case 
one would expect to find redeposited cremations in the other mounds in 
the complex, but without evidence that they were burned in situ. It is 
always possible that the primary mound platform served both of these 
purposes or other, as yet unknown, purposes. 



Anthropology 



77 



Post Hole Patterns 

A number of post holes were found at the edges of the primary 
mound platform (Fig. 1). Five of these were large, and were located 
near the eastern edge. Three smaller ones were found in back of them. 
This pattern was nearly duplicated at the other end of the platform, 
where six large post holes were found — four at the edge and two more 
in back of them. The large post holes were filled with loose black soil 
containing an abundance of charcoal. They were about 1 foot in diameter 
and from 1-2 feet deep, with the exception of 2 shallow post holes, 1 at 
each edge of the platform and in identical positions with relation to 
the other post holes in alignment with them. Several others were also 
noted at the northern edge of the mound, but they were generally small, 
shallow, and were not placed in any noticeable arrangement. The size 
and placement of the larger post holes suggests that they may have held 
the vertical support posts for some type of roofed structure over the 
primary mound platform. 




Figure 1. Central platform of "Great Mound" showing primary mound, mound capping, 
post holes, cntranccway, and inside edge of ditch. 



78 Indiana Academy of Science 

Approximately 450 small post holes were revealed on the central 
platform in a roughly circular pattern surrounding the mound. In the 
eastern portion of the platform, they were confined to a narrow zone 
which tended to follow in a straight line near the inside edge of the 
ditch. They were scattered elsewhere on the platform, perhaps suggesting 
periodic reconstruction of the fence, but the generally circular arrange- 
ment was apparent all around the periphery of the mound. With the 
exception of one place where a test trench had previously obliterated 
some of the post holes, there was no obvious break in the pattern. The 
post holes were very shallow and small, averaging about 0.2 foot in 
diameter. They were filled with light brown earth which was difficult to 
distinguish from subsoil. Most of them were pointed at the bottom, in 
contrast with the larger post holes at the edge of the primary mound 
platform, which were rounded or flat at the base. 

The post holes on the central platform probably represent small 
stakes or saplings, sharpened to a point at one end and placed in such a 
way that branches could have been woven between them in wattle-like 
fashion. The resulting brush fence or screen would have isolated the 
primary mound platform and prevented outsiders from observing any 
activities which might have taken place within. There was no clear evi- 
dence of an opening through the fence, but suggestions of one or 
possibly two pathways were noted to the north and south, where post 
holes in both locations led from the ditch to the edge of the mound 
capping in relatively straight lines. Since no large gap in the pattern 
was observed, however, the possibility of a baffled entranceway cannot 
be dismissed. 

Features 

A rectangular subfloor log tomb was found adjacent to and south 
of the primary mound, and was apparently the central feature of the 
later mound capping. The tomb was about 5 feet wide and 7 feet long, 
and was constructed of logs which had been placed in a "lean-to" fashion; 
burned; and then covered with earth while the structure was still burn- 
ing. Two burials had been placed on the floor of the tomb, and associated 
with them were some fragments of mica and a platform pipe. The 
burials consisted of a redeposited cremation and a secondary or "bundle" 
burial, the latter of which was an adult male. A total of 13 deer bone 
awls placed upright around the edge of the tomb suggests that they 
may have been used to tack down a covering of cloth or animal skin. 
This trait has also been noted at the Seip mound in Ohio (7). 

A roughly circular feature about 5 feet in diameter and a rectangu- 
lar basin about 3 feet by 3.5 feet were found within the primary mound. 
Evidence of burning on the interior and the presence of a baked clay 
ridge surrounding each of these features suggests the possibility that 
they once served as crematory basins, but no concentrations of bone 
fragments or artifacts were found in them. 



Anthropology 79 



Burials 



A total of six burials were excavated in the "Great Mound." With 
the exception of the two burials in the log" tomb, however, all of them 
were apparently intrusive. Two of these burials, one adult male and one 
adult female, were flexed inhumations which were found near the surface 
of the mound. There was clear evidence that one of these was buried in 
an intrusive pit. A concentration of burned human bone fragments repre- 
senting a redeposited cremation was found in disturbed earth near the 
center of the primary mound, and another re-deposited cremation of a 
single individual was present in a pit which had been intruded through 
all three floors of the platform. Although there were no artifacts found 
in association with any of these burials, the practice of intruding burials 
into mounds is typically a Late Woodland trait, and has been docu- 
mented for several Late Woodland cultures in the Ohio Valley. 

Artifacts 

With the exception of the mica, bone awls, and platform pipe as- 
sociated with the log tomb, most of the artifacts recovered from the 
"Great Mound" were found in disturbed fill dirt. 

All of the deer bone awls which had been placed around the tomb 
were made from split metatarsals, some of which had been burned. 

The platform pipe was made from material resembling limestone. 
It was approximately 4V 2 inches long and 1V2 inches high. The base was 
slightly curved, and the bowl was constricted near the out-flaring rim. 
A ridge was present around the middle of the bowl, which expanded 
slightly from this point downward to its juncture with the base. The 
pipe was not keeled. 

Ten of the 13 sherds recovered from the "Great Mound" were plain. 
Three sherds show portions of the "nested-diamond" design character- 
istic of New Castle Incised (3). 

Eleven fragmentary bone artifacts were also found, most of which 
had been burned and polished. All but two were drilled completely 
through from both sides. Objects of this type frequently have two 
holes drilled through them, and evidence for this was noted on the nearly 
complete specimens. Three of the artifacts were in the shape of split 
bear canine teeth. Tentative identification of the material from which 
several of the bone artifacts were made revealed one of deer; one of 
snapping turtle; and three of bear, including two of the bone imitations 
of bear canines. Several are polished on one side only, as if they had 
originally been attached to a garment rather than worn around the 
neck as gorgets or pendants. Effigies in bone of split bear canine teeth 
have been noted in several Ohio and Illinois Hopewell sites, including 
Mound 25 of the Hopewell group (6). 

Other artifacts from the "Great Mound" include one rectangular 
gorget fragment of slate and several ground stone and chipped flint arti- 
facts, including hammerstones, scrapers, knives, and projectile points. 



80 Indiana Academy of Science 

Some of the projectile points are corner-notched; others came from the 
subsoil underlying the mound and from the central platform. One has 
the bifurcated base typical of Archaic points. 

Test Trench 

A 5-foot by 10-foot test trench was excavated to a depth of 8 
inches in a small mound on the larger of the two panduriform earth- 
works. The trench yielded a great quantity of rocks, flint chips, deer 
bone, chunks of burned clay, pottery and other debris. All of the material 
appeared to be characteristic of a village midden deposit. Approximately 
200 sherds were recovered, at least 25 of which have incised designs. 
Cremated human bone fragments were scattered throughout the fill. 
Two secondarily deposited lenses of ash were noted, but there was no 
indication of in situ burning. This evidence tends to support the theory 
that the primary mound platform of the "Great Mound" may have 
served as a central crematorium. 

Conclusions 

The excavation of the "Great Mound" was undertaken because the 
excavations of other "sacred circles" have failed to provide a clear 
definition of the structural features or cultural affiliations involved. 
The first "sacred circle" to be excavated was the Mt. Horeb site in 
Kentucky, where Webb (10) found a very regular arrangement of 
paired post holes in a circular pattern measuring 97 feet in diameter. 
No break in the post hole pattern was noted, however, nor was there 
any evidence of a structure inside the fence. The "sacred circle" at Mt. 
Horeb did not have a mound on the central platform. 

The next circular enclosure to be excavated was the Dominion 
Land Company site in Ohio, where Baby and Goslin (1) found the re- 
mains of a house outlined by a circular pattern of outsloping post holes 
underneath one of two mounds on the central platform. This house 
measured 40 feet in diameter, and additional post holes suggesting roof 
supports were located inside the pattern. No post holes were found en- 
circling the mounds, however. 

The Bertsch site in Wayne County, Indiana, was recently excavated 
by J. M. Heilman {unpublished data), who found a centrally located 
burial pit, a portion of a wall trench, and what appeared to be three 
parallel lines of post holes flanking these features, all within a circular 
burned structure about 30 feet in diameter on the central platform of a 
"sacred circle." Since plowing had defaced the surface of the central 
platform, it was undetermined whether or not the "sacred circle" origi- 
nally enclosed a mound. 

The "Great Mound" has some features in common with each of these 
sites, but also has some characteristics which appear to be unique. On 
the basis of pottery, geographical proximity, and occurrence within an 
earthwork complex, the closest affinities of the "Great Mound" seem to be 
with the New Castle site in Henry County, Indiana, from which radio- 



Anthropology 81 

cardon dates of A.D. 10 and A.D. 40 were obtained (9). As far as 
structural features are concerned, however, its ties are with the Ginther 
Mound in Ohio, which was an isolated mound adjacent to a "sacred 
circle." In his excavation of the Ginther Mound, Shetrone (5) noted a 
"highly specialized floor" of burned clay, as well as post holes around the 
edge. The fact that no burials were found which could be attributed to the 
people responsible for constructing the mound led Shetrone (5) to the 
conclusion that ". . . the impressive tumulus was erected to mark the 
spot where some event or occurrence of great moment and significance 
to its builders transpired — rather than as monument to the dead." 

As far as cultural relationships are concerned, the "Great Mound" 
has traits considered typical of both Adena and Hopewell. Circular en- 
closures and incised pottery with the "nested-diamond" design have 
traditionally been considered characteristic of Adena, but the occurrence 
of a platform pipe and bone artifacts in the shape of bear canine teeth 
suggests Hopewellian influence. The Mt. Horeb and Dominion Land 
Company sites are considered to be Adena by Webb (10) and by Baby 
and Goslin (1). Shetrone (5) regards the Ginther Mound as basically 
Hopewell, but recognizes some anomalous traits. Swartz (8) and J. M. 
Heilman (unpublished data) are noncommittal about the cultural affilia- 
tion of the New Castle and Bertsch sites, but both tend to emphasize the 
Hopewellian aspects of each. 

It is possible that the "Great Mound" represents a marginal persist- 
ence of Late Adena at a time when Hopewell was fully developed else- 
where in the Ohio Valley. The presence of several characteristically 
Hopewell traits, however, is clearly in evidence. In the final analysis, 
it is artificial to assign either the Adena or the Hopewell label 
exclusively to the situation at Mounds State Park. It appears, rather, 
that the blending of several cultural expressions produced a distinctive 
regional tradition during the Middle Woodland period which may be 
restricted to the upper Whitewater and White River drainages. The 
excavation of the "Great Mound" has contributed to our knowledge of 
this little-known cultural complex, and to our general understanding of 
Ohio Valley prehistory as well. 



Literature Cited 

1. Baby, Raymond S., and Robert M. Goslin. 1953. Archaeological Field Work, 1953. 
Museum Echoes, Ohio Hist. Soc. 26 :79-80. 

2. Black, Glenn A. 1931. Report of Trip of Glenn A. Black and Dr. Warren K. Moore- 
head. Manuscript on file at Glenn A. Black Laboratory of Archaeology, Indiana 
University, Bloomington. 

3. Buchman, Randall L. 1968. A Preliminary Report of the Pottery from the New 
Castle Site, p. 10-14. In B. K. Swartz, Jr. [ed.] Archaeological Reports, No. 3. 
Ball State Univ., Muncie, Indiana. 

4. Kellar, James H. 1969. New Excavations at Mounds State Park — Life in Indiana 
2,000 Years Ago. Outdoor Indiana 34 :4-9. 



82 Indiana Academy of Science 

5. Shetrone, H. C. 1926. Exploration of the Ginther Mound, p. 61-70. In W. C Mills 
[ed.] Certain Mounds and Village Sites in Ohio. Vol. 4, Pt. 3. 

6. . 1926. Exploration of the Hopewell Group, p. 77-305. In W. C. Mills 



[ed. ] Certain Mounds and Village Sites in Ohio. Vol. 4, Pt. 4. 

and E. F. Greenman. 1931. Explorations of the Seip Group of Prehis- 



toric Earthworks. Ohio Archaeol. and Hist. Quart. 40(3) :343-509. 

Swartz, B. K., Jr. 1967. Tentative Observations on the Placement of Archaeological 
Materials Recovered from Mound Four, New Castle Site, p. 11. In B. K. Swartz, Jr. 
[ed. ] Archaeological Reports, No. 2, Ball State Univ., Muncie, Indiana. 

. 1968. Radiocarbon Dates from the New Castle Site, p. 15. In B. K. 



Swartz, Jr. [ed. ] Archaeological Reports, No. 3, Ball State Univ., Muncie, Indiana. 

10. Webb, William S. 1941. Mt. Horeb Earthworks, Site 1, and the Drake Mound, Site 
11, Fayette County, Kentucky. Univ. of Ky. Rep. in Anthropol. and Archaeol. 5 
(2) : 135-218. 



BOTANY 

Chairman: Robert L. Kent, Indiana Central College 
Robert Simpers, Crawfordsville, Indiana, was elected Chairman for 1970 

ABSTRACTS 

Trilliums of Franklin County, Indiana. Lloyd Beesley and Adele 
Beesley, Cedar Grove, Indiana. — According to Deam's Flora of Indiana, 
there are seven species of Trillium found in Indiana. In our search in 
Franklin County we have found all seven species: Trillium sessile, T. 
sessile f. luteum; T. recurvatum; T. nivale; T. grandiflorum ; T. 
cernuum; T. gleasoni; T. gleasoni f. Walpolei. 

Report of a carotenoid-mutant of Cyanidium caldarium. K. E. Nichols 
and W. W. Bloom, Valparaiso University. — Wild-type cells of C. cal- 
darium have been reported to produce J5-carotene, zeaxanthin, and an uni- 
dentified xanthophyll. Chlorophyll a, C-phycocyanin, and allophycocyanin 
also characterize the wild form. A new mutant has recently been isolated 
from a previously described chlorophyll-less form. Spectroanalysis of 
an ether extract of ground cells of the new mutant reveal an absence of 
the carotenoids of wild-type cells. Light absorption maxima are at 
378, 400, and 425 m/*. and are similar to those reported for ^-carotene. 
If the identification is correct, the finding would appear to support the 
belief that ^-carotene may be one of several hydrolycopene precursors 
of the carotenoids. 

Responses of Megagametophytes of Marsilea to Growth Substances with 
Respect to Rhizoid Formation. William W. Bloom and Kenneth E. 
Nichols, Valparaiso University. — High concentrations of indole acetic 
acid and napthalene acetic acid inhibit rhizoid formation in both light and 
darkness in non-pregnant megagametophytes of Marsilea, but lower con- 
centrations stimulate rhizoid growth. Gibberellic acid stimulates cell 
division but has limited effects on rhizoid production. 

Phenology Studies of Ten Species at Eleven Locations in Indiana. Byron 
O. Blair, Purdue University. — In 1964 with support from the NC-26 
(Regional Committee on Climatology Studies) and the Purdue Agri- 
cultural Experimental Stations, 10 semi-shrub perennial species were 
planted at 9 locations in Indiana. Plantings in each case were located 
near a functioning weather station and, in most instances, at experi- 
mental farms where weather data have been taken for several years. 
The species vary in blooming habit, varying from early spring until 
early fall. All species have been developed from cuttings or clonal ma- 
terial as a means of controlling genetic variability. This is an essential 
feature of phenology studies which was neglected in most past records 
and studies. 

At each location, in addition to daily weather data which are avail- 
able, 4-inch soil cores have been taken for physical and chemical analysis. 

83 



84 Indiana Academy of Science 

Three years of satisfactory flowering data have now been collected and 
with analysis of soil profiles which vary from muck, to sands, to im- 
perfectly drained clays, we are ready to evaluate seasonal influences 
and diurnal effects of the local climate on growth and development of 
these species. 

Chromosome Associations in Corn Monoploids. L. Ford, Butler, Indiana. — 
Today there is good evidence that haploids occur spontaneously in most 
Angiosperms, including Zea maize. The study of cytological aspects and 
interpretations of univalents, bivalents, secondary associations, restitu- 
tion nuclei, and somatic doubling have become important not only in 
evolutionary and species relationships studies, but also from the point 
of view of practical plant breeding. There have been only a few studies 
of maize monoploid microsporocytes in the literature, and because of 
the importance of corn monoploids in modern hybrid corn production, 
this study is offered. Microsporocytes from 50 maize monoploid plants 
were identified, collected, and prepared by aceto-carmine squash tech- 
niques, and examined cytologically. A rather high amount of non- 
homologeous pairing was found. In 43 cells with chiasmata or bivalent 
association, 31 also had secondary association. In addition, there were 
87 cells in the same material with secondary association only. There does 
appear to be evidence, therefore, for a type of secondary association in 
corn due either to basic homologies of chromosomes; presence of homo- 
logous parts in non-homologous chromosomes; or residual attraction 
between chromosomes. 

A Microspectrophotometric Analysis of DNA in the Heterothallic Species 
of Slime Mould, Didymium iridis. Which Sometimes Exhibits Apogamy. 

J. Yemma, Pennsylvania State University. — Data are presented which 
show that selfing (or apogamy) sometimes takes place in individual 
clones of known mating types. These conclusions were arrived at through 
appropriate use of the microspectrophotometer for Feulgen-DNA nuclear 
content analysis, and IBM 360 computer for data analysis. 

Some Charophytes from the Pleistocene. Fay Kenoyer Daily, Butler 
University. — The occurrence of specimens belonging to the genus Lato- 
chara in glacial deposits of New York and Indiana was reported in Daily 
(1961). This extended the range of the genus from the Eocene. Repre- 
sentatives were again discovered in the late Wisconsin till of South 
Dakota. Specimens were so abundant that sectioning of lime-shells was 
possible, allowing confirmation of identification and providing new in- 
formation about the species, Latochara Way net Daily. Other specimens 
in the lacustrine deposits were referred to the modern species, Chara 
delicatula Ag. emend A. Br. A mineral incrustation on the exterior of 
the whole plant provided casts of several structures rarely preserved in 
fossil charophytes. 

Dr. F. V. Steece (1961, 1966) examined the charophytes from two 
of the sites and provided these specimens and additional material as 
well as stratigraphic data for the present study. Dr. Steece reported 
Clavatorites and Chara from these sites. The Clavatorites are considered 
to be Latochara in the present study, although eventually they may be 
found to be synonymous. 



Some "Atypical" Stem Structures in the Gramineae 
Paul Weatherwax, Indiana University 



Abstract 

To regard the stems of such grasses as wheat or corn as "typical" of the Gramineae is 
to overlook some interesting and significant deviations in vascular and parenchymatous 
pattern. Some of these, occurring in the culms and rhizomes of representatives of such 
genera as Andropogon, Olyra, Zizaniopsis, Arrhenatherum, Phleum. Melica, Cinna. and 
Hymenachne, are pointed out and described. 

Many of the brief general descriptions of the grass family, such 
as those given in encyclopedias or as introductions to taxonomic trea- 
tises, cite the hollow stems of such plants as wheat, rye, or bamboo as 
"typical" for the family. It is true that they usually mention the solid 
stems of corn or sugar cane, but in stressing the mechanical advantages 
of the hollow one they often leave the impression that the solid one is 
of rare occurrence. This treatment does not do full justice to the solid 
stem, and it overlooks some interesting deviations from these two types. 

Formation of the hollow internode can easily be traced downward 
from the apical meristem. The young internode is solid, but the pro- 
vascular strands appear only in the outer part. As the internode grows 
older and elongates, the cells of the peripheral region continue to divide, 
and some of them elongate, this activity finally being limited to the 
intercalary meristem at the lower end of the internode. However, the 
parenchyma cells in the middle soon cease to divide, but grow larger, 
become highly vacuolate, are finally torn apart by the elongating action, 
and ultimately disintegrate. Maturation of the vascular tissues and fibers, 
often accompanied by extensive lignification of parenchyma cells, forms 
a firm outer layer, giving to the internode its characteristic rigidity and 
strength. The inner wall of the hollow cylinder thus formed usually 
bears a few ragged remnants of the parenchyma cells destroyed in the 
process. 

The solid type of stem, that is, the one in which the body of the 
fully developed internode is completely filled with tissue, is to be found 
in one or more species of probably one-fifth of the 500 or more genera 
now recognized by systematists. In most of these the vascular bundles 
are scattered over the cross section of the internode, as in the well- 
known corn stem, being larger and more widely spaced in the middle 
of the section. Since this type of stem is found in some members of the 
Bambuseae and in most, if not all, of the Maydeae, thus at opposite ends 
of the phylogenetic spectrum, it would seem to have little evolutionary 
significance. 

Between the hollow and solid forms there is an intermediate condi- 
tion in which the middle of the internode is devoid of vascular tissue 
and the parenchyma breaks down irregularly, leaving a ragged lacuna 
which may vary in size and shape in different specimens of a species or 
even in different parts of a single individual. An interesting variation of 

85 



86 



Indiana Academy of Science 



^STO^, 



% 






3 4 



/: 



i%Jllr' 





.* 







Figures 1, 2. Cwi!m a?id rhizome of Euchlaena perennis. 

Figures 3, 4. Culm and rhizome of Agropyion repens. 

Figures 5, 6, 7. Culm, rhizome, and stolon of Cynodon dactylon. 

Figures 8, 9. Culm and rhizome of Zizaniopsis miliacea. 

Figure 10. Clustered corms of Arrhenatherum. 

Figure 11. Corm of Cinna. 

FIGURE 12. Ornamental bird, made from dyed parenchyma cores of Hymenachne. 

Figure 13. Section of stem of Hymenachne pseudointenupta. 

Figures 14, 15. Transverse and longitudinal sections of stem parenchyma of Hymenachne. 



Botany 87 

this type was reported many years ago in a South American species of 
Olyra. In it the central part of the parenchyma of the internode dis- 
integrates, leaving one or usually more vascular bundles free, extend- 
ing from node to node in the cavity (6). 

Whatever the structure of the culm may be, we usually find a 
very different picture on examining the rhizome or stolon of the same 
plant if it has either of these structures. Although the internode of the 
upright stem may be hollow, there is a strong tendency for that of the 
horizontal structure, especially the rhizome, to have a solid pith. And, 
while the vascular bundles of the culm tend to be concentrated toward 
the periphery, the horizontal structures usually show a more or less 
definite cortical region having few or no vascular bundles. 

One of the simplest illustrations of this difference between culm and 
rhizome may be seen in the perennial teosinte plant, Euchlaena perennis 
Hitchc. The culm of this grass is so much like a corn stem in structural 
pattern that the two are almost indistinguishable (Fig. 1). But if a 
series of cross sections are cut, one from each internode, continuing 
downward as the vertical stem becomes a horizontal rhizome, a progres- 
sively widening cortical zone is seen (Fig. 2). The common weed, 
Sorghum, halepense (L.) Pers. (Johnson grass) has the same structure, 
but with a more variable expression. In this species, vertical rhizomes, 
which often grow to considerable depth in porous soils, also show this 
cortical region. 

The well known quack grass, Agropyron repens (L.) Beauv., is 
fairly representative of a large number of species in which, although 
the internodes of both culm and rhizome are hollow, the rhizome tends 
toward the solid form (Figs. 3, 4). The vascular bundles of the culm 
are arranged in circles toward the outside, the outer circle consisting of 
very small bundles set in a ring of sclerenchyma just beneath the epi- 
dermis. Although the rhizome may have a diameter at least twice that 
of the culm, its lacuna is much smaller than that of the culm. Here the 
main vascular bundles are located in a ring of sclerenchyma which has 
a diameter about half that of the rhizome. The cortical region consists 
mostly of parenchyma with a few vascular bundles which come in from 
the leaves and will enter the central cylinder at a morphologically lower 
level. 

Some species such as Bermuda grass, Cynodon dactylon (L.) Pers., 
have both rhizomes and stolons, the latter usually standing somewhere 
between the culm and the rhizome in structure (Figs. 5-7). 

A much more striking difference between culm and rhizome is found 
in the aquatic grass, Zizaniopsis miliacea (Michx.) Doell. & Aschers. The 
culm internode is hollow and not very different from that of other hollow 
stems (Fig. 8). But a cross section of the rhizome (Fig. 9) shows a 
clearly defined cortical region set off from the central cylinder by a row 
of cells having much the appearance of an endodermis. Thus far, how- 
ever, tests have failed to show any suberization of the cell walls of 
this layer. The central pith is solid, with vascular bundles scattered over 



88 Indiana Academy of Science 

it much as in the corn stem. The cortical region has many rudimentary 
vascular bundles, some of them consisting merely of strands of fibers. 
Adventitious roots arising from the rhizome have their origin in the 
outer part of the central cylinder, giving to the cross section much the 
appearance of a section of a root at the place where a branch root arises. 

These marked differences between culm and rhizome lead us to 
ask what factors cause the apical meristem to leave behind it one pattern 
of structure when moving horizontally to form the one and then to 
change the pattern as it turns upward to form the other. The difference 
cannot be wholly due to the horizontal position of the one structure and 
the vertical position of the other, because in some species, such as 
Sorghum halepense, segments of rhizomes in both positions are alike in 
structure. Neither can it be attributed wholly to the underground as op- 
posed to the aerial environment since stolons show much the same pattern 
as rhizomes. The answer to the question must lie somewhere in that 
relatively unexplored region between the extensive studies on tunica 
and corpus on the one hand and what is known of the gross morphology 
of the mature stem on the other. 

Another deviation from what is regarded as the typical stem 
structure is the formation of bulb-like swellings in the basal internodes 
of culms. Frequently only one internode is involved in each culm, but 
sometimes there may be as many as three or four. These do not fit neatly 
into any one of the conventional categories of modified stems, but the 
term corm seems to be the least objectionable. The frequent reference to 
them as bulbs is acceptable only if this is taken to refer to their external 
appearance. They are not at all to be confused with the bulblets produced 
regularly in Poa bulbosa L. and occasionally in many other species where 
spikelets are modified into bulbs or small plants. 

The occurrence of these corms is known in one or more species of 
Arrhenatherum, Hordeum, Phleum, Alopecurus, Molinia, Phalaris, Pani- 
cum, China, Holcus, Poa, Beckmannia, Colpodium, Ehrharta, and 
Melica, and they are probably found in other genera. Their distribution in 
seven of the 15 recognized tribes is such that no phylogenetic significance 
can be attached to them (2). Neither is there any clear correlation 
between corm formation and ecologic conditions. The species which display 
it are well represented in the Mediterranean region and in the 
moist climates of Northern Europe, but some are found in other environ- 
ments, and in any area the expression of the condition is erratic. 

In such genera as Phleum, China, and Melica there is usually only a 
single swollen internode, at about the ground level (Fig. 11), but in others 
such as Arrhenatherum elatius, var. bulbosum (Willd.) Spenner and 
Panicum bulbosum H.B.K., there may be as many as three or four, and 
the culms having them may grow in clusters of 15 or 20 or more (Fig. 
10). Because of the peculiar appearance of such plants, some of them are 
popularly known as onion grasses. One report (3, 4) indicates that the 
variety of Arrhenatherum may even produce rhizomes whose swollen 
internodes give them the appearance of short strings of beads. 



Botany 89 

The internal anatomy of the corm is much like that of a rhizome. 
The corm is solid although the culm internodes above it may be hollow. 
In cross section it shows a cortical region surrounding a circle of vascular 
bundles set in a ring of sclerenchyma. The central parenchyma often con- 
tains vascular bundles, and its cells remain alive and display much sea- 
sonal physiological activity, indicating that food storage is involved (1). 
In some genera the predominant food is ordinary starch; in others, such 
carbohydrates as phlein, graminin, triticin, or inulin may be found. In a 
species of Molinia the parenchyma cells in the corm develop thick walls 
in autumn, and the following spring these walls dissolve and are probably 
used for food (1). This suggests that the reserve food is a hemicellulose. 

The last stem anomaly to be mentioned is included more as a curiosity 
than as a thing of any particular morphological importance. It occurs in 
Hymenachne, a genus of aquatic grasses with about eight species in the 
tropics of both hemispheres. H. amplexicaiilis (Rudge) Nees, of the 
American tropics, and the very similar H. pseudointerrwpta C. Muell., of 
Southeast Asia, have been examined in some detail. 

A cross section of the spongy submersed stem (Fig. 13) shows a thin 
layer of sclerenchyma next to the epidermis and then two rings of 
parenchyma separated by a ring of fibers. The vascular bundles are 
arranged in circles in the inner of these layers of parenchyma. All these 
tissues make up a shell around a central cylinder of parenchyma, which 
is devoid of vascular bundles. This core of parenchyma is the object of 
unique interest. 

When the parenchyma cells are young, they are closely fitted 
together, with very small intercellular spaces. But as they mature during 
elongation of the internode, they are pulled apart until they assume the 
stellate form characteristic of the tissue of many other submerged 
aquatics (5) and (Figs. 13-15). These stellate cells form a series of trans- 
verse plates with enough longitudinal connections between the plates to 
hold the entire mass together so that the core of parenchyma can be 
punched out of the internode as an intact cylindrical mass. These long, 
terete cores are pliable and firm enough to be bent into various shapes. 
They can also be colored with various dyes. In the markets of Bangkok, 
and probably other cities of Southeast Asia, there can frequently be 
found on sale various ornamental objects in the form of flowers, birds, 
or animals made from these cylinders of pith (Fig. 12). 

The late Dr. Agnes Chase, of the Smithsonian Institution, once told 
me that, in the hinterlands of Brazil, she had seen stems of Hymenachne 
used for wicks in crude oil lamps. A simple experiment with pieces of 
stems of the Asian species shows that they could be put to the same use. 
The unique structure of the central parenchyma provides an excellent 
capillary pathway for movement of the oil between the stellate cells. At 
present we know of no grass species outside the genus Hymenachne 
whose stems could be used in this way. 



90 Indiana Academy of Science 

Literature Cited 

1. Arber, Agnes. 1934. The Gramineae. The Macmillan Co., New York, N. Y. 480p. 

2. Burns, W. 1946. Corm and bulb formation in plants, with special reference to the 
Gramineae. Trans. Proc. Bot. Soc. Edinburgh 34 :316-347. 

3. Chase, Agnes. 1911. The subterranean organs of China arundinacea. Rhodora 13 :9-10. 

4. Chase, Agnes. 1911a. Arrhcnatherum elatius, var. tuberosum in America. Rhodora 

13 :207-8. 

5. Haynes, James L. 1935. The anatomy of an anomalous grass, Hijmenachne amplexi- 
caulis. Proc. Indiana Acad. Sci. 44 :69-72. 

6. MCller, Fritz. 1889. Freie Gefassbiindel in den Halmen von Olyra. Flora 72:414-420. 



CELL BIOLOGY 

Chairman : Edward Hinsman, Purdue University 
Edward Hinsman, Purdue University, was re-elected Chairman for 1970 

ABSTRACTS 

Response of the Duck Thyroid to the Administration of Thiouracil. J. R. 

Welser and W. W. Carlton, Purdue University. — The response of the 
thyroid gland to thiouracil was studied in day-old Pekin ducks fed a diet 
of duck mash supplemented with 0.2 7r thiouracil for 6 weeks. Thyroid 
glands were fixed with gluteraldehyde and formalin at days 1, 7, 15, 22, 
28, 31, 38, 42 of the experimental period for observation with the light 
and electron microscopes. The initial response was a marked hypertrophy 
of thyroid epithelium with the normally squamous to cuboidal thyroid 
cells becoming tall columnar. Ultrastructural features of the response to 
thiouracil included a marked distention of the endoplasmic reticulum, 
proliferation of Golgi apparatus, increase in the number of mitochondria 
and microvilli and the formation of numerous large colloid containing 
vacuoles in thyroid cells. Hypertrophy was followed by a marked hyper- 
plasia of the thyroid epithelium. The ultrastructural morphology of the 
hyperplastic cells was similar to the hypertrophic cells. In thyroids from 
ducks fed longer than 21 days, distention of the cisternae of the endo- 
plasmic reticulum was greater and was accompanied by the formation 
of more numerous and larger colloid containing vacuoles. 

Regeneration of Skeletal Muscle in Vitamin E-Deficient Rabbits. J. F. 

Van Vleet, B. V. Hall and J. Simon, Purdue University. — Weanling 
rabbits fed a semi-synthetic vitamin E-deficient diet developed hyaline 
degeneration of skeletal muscle in 20-30 days. Light and electron micro- 
scopic study of the subsequent events of regeneration of damaged muscle 
fibers in the diaphragm of affected rabbits showed the discontinuous type 
of regeneration to predominate. Myoblasts developed from surviving 
sarcolemmal nuclei and adjacent sarcoplasm. Myoblastic proliferation 
produced multi-nucleated giant cells that fused to form a syncytium or 
cords of cells that lay within the sarcolemmal tube of the regenerating 
fiber. Short fragments of thin (50 A) myofilaments were observed adja- 
cent to polysomes in proliferating myoblasts. Thick (100 A) myofilaments 
were subsequently found in small bundles 2-3,u long in the sarcoplasm 
between a central row of myoblast nuclei and the sarcolemma lining the 
sarcolemmal tube. Sarcomere formation followed as thick and thin fila- 
ments become interdigitated and Z-band material formed discs at the 
ends of the sarcomeres. Numerous myofibrils were soon recognizable and 
sarcomere bands became aligned transversely to restore longitudinal and 
transverse striation to the regenerated fiber. Rows of nuclei surrounded 
by numerous mitochondria remained in the centers of regenerating fibers. 
Later, these nuclei were found in their normal position against the 
sarcolemma. 

91 



92 Indiana Academy of Science 

Deficient Myelination in the Brain of "Quaking" Mouse: an Electron 
Microscopic Study. Itaru Watanabe and Glenn Bingle, Indiana 
University Medical Center. — The mutant gene "quaking" is inherited as 
an autosomal recessive trait. The homozygotes exhibit a life-long 
neurologic disorder manifested by tremulousness and frequent tonic 
seizures. The brain is small in size due to lack of myelin sheaths. 

To obtain further information of the mechanism of the deficient 
myelin formation, electron microscopic studies were performed on the 
corpus callosum of homozygotes aged 10, 17, 19, 21, 35, and 40 days, 
namely at the period of physiological myelination in the normal mice. 

The major alterations were restricted in the myelin-forming cells or 
oligodendrocytes. A number of unique lipid bodies were already present 
in the perinuclear cytoplasm as early as 10 days of age when myelination 
was not evident in the adjacent tissue. Furthermore, the process of 
myelination was markedly disturbed. Redundant oligodendroglial processes 
surrounded a single axon in an irregular fashion and eventually com- 
pacted to form a structure reminiscent of a myelin sheath. This resulted 
in a sheath of variable thickness and often a segment of the axon was 
not at all covered by these abnormal sheaths. The cytoplasm of the 
oligodendroglial processes harbored spherical vacuoles, which increased 
in size by liquefaction of oligodendrocytic cytoplasm and, further, dis- 
integrated the abnormal myelin-like membranes. 

These morphological changes suggest an inborn-error of metabolism 
in the oligodendrocyte. 

The Identification of Gamma-A Globulin in Human Enteric Epithelial 
Cells. John F. Schmedtje, Indiana University Medical Center. — There 
have been discordant reports on the presence of gamma-A globulin in 
enteric epithelial cells — particularly in mucous type cells. In the present 
investigation, gamma-A globulin was identified in the mucous type epi- 
thelial cells of the human appendix. 

Tissue blocks from normal human appendixes were quick frozen 
immediately after surgical removal. Frozen sections were cut on a 
cryostat. Some sections were set aside for H & E staining. Goat anti- 
human gamma-A globulin, conjugated with fluorescein isothiocyanate was 
used according to Coon's direct method. Non-specific reactivity of the 
conjugated antiserum was removed by adsorption with mouse liver and 
ox marrow powders. Unconjugated antiserum was used for control 
reactions. 

Positive fluorescence occurred in numerous plasma cells located 
beneath the basement membranes of luminal epithelial cells and cells 
that lined the crypts of Lieberkuhn. Positive fluorescence also occurred 
in all epithelial cells, except those at the bottom of the crypts of 
Lieberkuhn. In mucous type epithelial cells, the mucinogen areas and 
nuclei were negative. However, positive fluorescence occurred in the 
cytoplasmic areas around the mucinogen. These results are interpreted as 
supportive evidence that gamma-A globulin is secreted into the 
appendiceal lumen. 



Cell Biology 93 

Effects of Adenosine Diphosphate on the Morphology of Heart Mito- 
chondria. N. E. Weber and P. V. Blair, Indiana University Medical Cen- 
ter. — Ultrastructural studies have been made of beef heart mitochondria 
in various metabolic steady states. Adenosine diphosphate promotes a 
highly condensed morphological arrangement of the inner mitochondrial 
membrane. Although oxidation rates are altered (and adenosine triphos- 
phate is produced) upon the addition of oxidizable substrate and inor- 
ganic phosphate, the appearance of the inner mitochondrial membrane 
is essentially unchanged. The effect of adenosine diphosphate was 
observed not only at 30°C but also at 0°C when mitochondria were 
exposed to rotenone, potassium cyanide and anaerobic conditions. The 
adenosine diphosphate promoted the condensed morphology in all cases 
with the possible exception of mitochondria exposed to rotenone. These 
observations suggest that large ultrastructural transformations are not 
required for energy transduction and that the morphology of the inner 
mitochondrial membrane may depend primarily on osmotic changes cre- 
ated by experimental conditions and nucleotide binding during steady 
state metabolism. The width of the most condensed double-layered 
cristae is approximately 110 A. Thus a reinterpretation of the negative 
staining of mitochondrial membranes consisting of 'tripartite elementary 
particles' must be considered with these measurements in mind. (Sup- 
ported by USPHS Grant HE060308 and the Indiana Heart Association.) 

The Isolation of Nuclei and Endothelial Cells from Brain. A. N. SlAKOTOS, 
Indiana University Medical Center.— Pure preparations of organelles and 
individual cell types are required for determination of their chemical 
composition and metabolic characteristics. A procedure is offered for pre- 
paring highly purified endothelial cells (capillaries) and nuclei from 
human and bovine brains. Phospholipid compositions of typical prepara- 
tions demonstrate, in particular, a lack of species variability for capillary 
structures. 

The isolation procedure employs modifications of differential and 
density gradient centrifugation commonly used for subcellular separa- 
tions. Special features of the procedure are: large-scale preparation; 
Sephadex G-25 used for the separation of nuclei and endothelial cells; 
and foam concentration used for further purification of endothelial 
cells. 

Ultrastructural and Enzymological Observations of Isolated Kidney 
Microvilli. Sakae Yumoto and Shinji Ishikawa, University of Tokyo. — 
Essentially pure microvilli of rat kidney cortex were isolated by isopicnic 
centrifugation on a discontinuous sucrose density (0.72-0.92 M). Electron 
microscopic examination of the obtained fraction showed the presence 
of cylindrical structures of 1.3 to 2.0^ in length and 0.1ft in width. When 
negatively stained with phosphotungstic acid, globular particles of 40 to 
60a in diameter were seen on the margin of microvillar membrane. 
These globular subunits were also observed en face on the surface of 
the membrane. This fraction showed phosphatase activity against ATP, 



94 Indiana Academy of Science 

AMP and p-nitrophenylphosphate as a substrate. Presence of Mg ion 
is required for ATP hydrolysis but Ca can replace Mg. This adenosine 
triphosphatase activity is not stimulated with the addition of Na and K 
ions and not inhibited by ouabain. The microvilli fraction also hydrolyzes 
other nucleotide triphosphates such as GTP, CTP, UTP and ITP. The 
activation energy of adenosine triphosphatase is 9.200 cal/mole. It is 
concluded that kidney microvilli do not have (Na+ + K + ) — stimulated 
adenosine triphosphatase but possess very high specific activity of Mg+ + 
adenosine triphosphatase. This Mg++ adenosine triphosphatase might 
play an important role on the active transport mechanism of kidney 
microvilli. 

NOTE 

Ultrastructural Features of the Calcifying Epithelial Odontogenic Tumor. 

J. B. Whitten, Jr., Indiana University Medical Center. — The calcifying 
epithelial odotogenic tumor (CEOT) was described in 1955 by Pind- 
borg. This is a benign, locally aggressive, epithelial tumor which probably 
arises from the odontogenic apparatus. The lesion most often occurs in 
the mandible usually in the posterior aspect, does not exhibit a sex 
predilection, and involves the same age group as the ameloblastoma. 
Radiographically, this disease presents as an ill-defined radiolucency 
usually with multiple radiodensities. Histologically, the tumor is com- 
posed of sheets or cords of polyhedral cells which have dense eosinophilic 
cytoplasm and large single or multiple nuclei often with multiple 
nucleoli. Focal droplet calcification resembling Liesegang's rings is often 
found interspersed within the epithelial cells as well as from calcification 
of the surrounding connective tissue. 

The tissue for this project was secured from a 68-year-old Caucasian 
male with a large 3.5 x 4 cm. expanding lesion of the anterior mandible. 
The tissue was hemisected, a portion processed for light microscopy and 
the remaining tissue prepared for electron microscopy. The tissue for 
electron microscopy was fixed in phosphate buffered 4% glutaraldehyde, 
post-fixed in phosphate buffered 1% osmic tetraoxide, dehydrated in 
ascending concentrations of ethyl alcohol and propylene oxide, embedded 
in epoxy resin, sectioned at about 600 A, stained with lead citrate and 
uranyl acetate, and examined with a RCA EMU 3H electron microscope. 

The fine structure of the CEOT showed the lesion to be composed of 
two cell types. The outer (Type A cell) was papillary in outline demon- 
strating many microvilli and closely resembled the papillary layer in 
developing teeth. The Type B cells which resembled the stratum inter- 
medium, were more regular in outline and were "packed" with mitochon- 
dria. These Type B cells were remarkably similar to the previously 
reported structure in oncocytes. The Type B cells were responsible for the 
bulk of the neoplastic process. In association with both the Type A and 
Type B cells were large quantities of fibrous protein with the period and 
width of amyloid. This amyloid appeared to be produced by the Type B 
cells. In many odontogenic tumors there has been reported an inductive 
effect. In this example of CEOT this inductive effect is found to be com- 
posed of elaborate arrays of basal lamina. 



Cell Biology 95 

OTHER PAPER READ 

Methacrylate Embedding: With Good Results. Frank Padgett, Indiana 
University Medical Center, Indianapolis, Indiana. 



Comparisons of Isolated Plasma Membranes from 
Plant Stems and Rat Liver 1 

D. James Morre, Purdue University, J.-C. Roland and C. A. Lembi, 
Laboratoire de Biologie Vegetale, Faculte des Sciences, Paris. 

Abstract 

A fraction enriched in plasma membrane was prepared from plant stems by low 
shear homogenization and differential and sucrose density gradient centrifugation. The 
plasma membrane fragments were identified by electron microscopy after differential stain- 
ing with a mixture of phosphotungstic acid and chromic acid which specifically and 
characteristically stained the plant cell plasma membrane. Chemical and enzymatic analyses 
comparing plasma membrane-rich cell fractions from rat liver and onion stems showed 
the characteristics of the surface membrane of the plant to be different from that of its 
mammalian counterpart. Whereas rat liver plasma membranes were characterized by high 
levels of the enzymes 5'-nucleotidase and Mg++ adenosine triphosphatase and a lipid 
content high in sphingomyelin, these plasma membrane markers for the rat could not be 
demonstrated in the plant preparations. 

The physiological response of any plant or animal tissue to an 
external stimulus involves the transmission of the stimulus across the 
barrier between the cell's interior and its external milieu. This barrier is 
the plasma membrane (or plasmalemma) plus any surface coat such as 
the plant cell wall. Three principal functions are usually attributed to 
plasma membranes: transport (both uptake and secretion); synthesis 
and/or assembly of surface coats; and transfer of information between 
the external environment and the cell's interior. Examples of such 
processes include uptake of ions, metabolites and growth regulators (8); 
secretion of enzymes (15); cell wall synthesis and assembly (23, 31); and 
the hormonal control of plant growth through altered cell wall mechani- 
cal properties (6, 16, 18). 

Studies of the role of the plasma membrane in influencing plant 
processes have been largely limited to indirect methods of analysis of 
whole cells and tissues (4, 10). In general, this appears due to the limita- 
tions set by the lack of cell free systems suitable for the analysis of such 
a complex and delicate structure. 

Cell membranes have been isolated from mammalian cells (2, 7, 
22) but not from plant cells. Progress has been hampered by two factors: 
(1) the inability to recognize isolated plasma membrane fragments in 
cell homogenates; and (2) the lack of techniques that rupture the rigid 
plant cell walls without destroying the fragile plasma membrane. 

This report summarizes results of a study directed toward the 
isolation, identification, purification and characterization of a plasma 
membrane-rich cell fraction from plant stems. 



1 Portions of these studies were conducted at the Institute de Biologie Moleculaire de la 
Faculte des Sciences de Paris in the laboratory of Prof. E. L. Benedetti during a sabbatical 
leave of absence from Purdue University by the senior author. We thank Drs. T. W. 
Keenan and L. Marcel Merlin for use of unpublished information. Supported in part by a 
grant from the NSF GB 7078 and a contract with the U. S. Army Biological Laboratories, 
Frederick, Maryland. Purdue University AES Journal Paper No. 3909. 

96 



Cell Biology 97 

Materials and Methods 

Biological material. Green onions (Allium cepa) were purchased 
locally and used on the day of purchase or stored at 4°C. Stem explants 
were harvested by cutting roots and lignified stem regions from the 
onion base. A cone of tissue 0.5 to 1 cm diam at the base and 0.5 to 1 cm 
high was then removed from the central portion of the onion bulb using 
a scalpel fitted with a narrow blade. Included in the explant were stem, 
meristematic region and leaf bases. Scale leaves and the green (top) 
portions of the onion were discarded. 

Livers were obtained from male rats (200 to 250 g) of the Wistar or 
Holtzman strains fed Purina Laboratory Chow or fasted 24 hr prior to 
sacrifice. 

Preparation of cell fractions. Approximately 5 g of stem explants 
from 30 to 50 onions were collected and weighed. Homogenates were pre- 
pared using a Polytron 20ST homogenizer (Kinematica, Incerne, Switzer- 
land) operated at slowest speed for about 60 sec (19) or a loose fitting, 
all-glass homogenizer of the Potter Elvehjem type. The homogenates were 
then squeezed through a single layer of miracloth (Chicoppe Mills, 
New York) to remove cell walls and tissue fragments and to break addi- 
tional cells. Centrifugations were for 30 min using the SW-39 L rotor 
for the Spinco Model L ultracentrifuge operated at 4°C. 

A nuclei fraction, containing occasional wall fragments, proplastids 
and mitochondria was obtained by low speed centrifugation (3,000 rpm 
for 10 min). A fraction enriched in proplastids was obtained at 5,000 rpm 
and a fraction enriched in mitochondria at 10,000 rpm. The fraction sedi- 
menting between 10- and 20,000 rpm contained dictyosomes of the Golgi 
apparatus and a variety of smooth membrane vesicles suspected of being 
a mixture of plasma membrane and tonoplast (vacuole membrane) frag- 
ments. Everything sedimenting between 20- 30,000 rpm was included in 
the microsome fraction and consisted largely of fragments of rough 
endoplasmic reticulum. The 30,000 rpm supernatant is referred to as the 
soluble fraction of the cytoplasm. 

The following homogenization media were employed: 

(1) 0.5 m sucrose containing 10 mM sodium phosphate, pH 6.8 and 
1% (w/v) dextran (Sigma average mol. wt. 225,000) (20); 

(2) 0.5 M sucrose containing 37.5 mM Tris-maleate, pH 6.5; 1% 
dextran and 5 mM MgCL (19); 

(3) Medium II minus MgCl 2 (for estimation of phosphatide acid 
phosphatase which was inhibited by Mg+ + ) ; 

(4) 1 mM sodium bicarbonate (22). 

Media (1), (2) and (3) gave smiliar results with plant preparations. 
Medium (4) was used in the rat liver preparations. 

Further fractionation of the 10- to 20,000 rpm fraction from plant 
stems was obtained by centrifugation in a layered sucrose gradient yield- 



98 Indiana Academy of Science 

ing 5 bands (21). The uppermost band contained lipid droplets and was 
discarded. The lowest bands contained mitochondria and endoplasmic 
reticulum. The intermediate bands yielded dictyosomes and smooth mem- 
branes free of dictyosomes. 

Plasma membrane fractions from rat liver were obtained by a modi- 
fication (7) of Neville's procedure (22). Golgi apparatus, endoplasmic 
reticulum and other cell fractions from rat liver were obtained as 
described previously (3, 14, 19). 

Chemical assays. Protein was determined by the Lowry procedure 
(13) or by the biuret method. Inorganic phosphorus was determined by 
the method of Fiske and Subbarow (9). Polar lipids were separated by two 
dimensional thin layer chromatography (12) and analyzed for phosphorus 
by the procedure of Rouser et al. (27) as modified by Parsons and 
Patton (24). Identity of the separated lipids was established by co- 
chromatography with authentic reference compounds (Applied Science 
Laboratories, State College, Penn.). Total sterols were measured by the 
method of Jorgenson and Dam (11). 

Morphological assays (electron microscopy). Portions of isolated pel- 
lets were fixed in 67r buffered glutaraldehyde (0.1 M potassium phos- 
phate, pH 7.2) for 18 to 20 hr with or without post fixation for 1 to 24 hr 
in V/ ( osmium tetroxide (in 0.1 M sodium phosphate, pH 7.2). Specimens 
were dehydrated through an acetone series; embedded in vestopal and 
sections were stained by one of the following procedures: 

(1) (with osmium post fixation) — Sections were stained with 
aqueous uranyl acetate and/or lead citrate (25). 

(2) (with osmium post fixation) — Sections were treated with 1% 
periodic acid for 30 min followed by 5 washes of 10 min each with dis- 
tilled water. The sections were then treated with a mixture of 1% phos- 
photungstic acid plus 10% chromic acid in water (pH less than 1) for 
2 to 5 min (PTA-CA procedure) (2). Finally, the sections were washed 
to remove excess stain and mounted on copper grids. 

(3) (no osmium) — Sections were treated directly with the mixture 
of 1% phosphotungstic acid plus 10% chromic acid in water for 2 to 5 
min; washed free of excess stain and mounted on copper grids (26). 

Enzyme assays. Enzyme assays used 0.1 to 0.8 mg protein in a final 
volume of 1 to 3 ml. The following enzymatic activities were determined 
according to the procedures referenced: 5'-nucleotidase (7); Mg++- 
ATPase (7); Na + -K + -Mg+ +-ATPase (7); phosphatide acid phospha- 
tase (28); and invertase (5). Assays were carried out at the pH optima 
for the total particulate fraction determined separately for liver and for 
cnions (see Fig. 5, for example) and at near optimum temperatures 
for each system (37°C for liver and 25°C for onions). 

Results 

When onion stem homogenates were fractionated, the fraction sedi- 
menting between 20- and 30,000 rpm contained smooth (ribosome-free) 
membrane vesicles of various sizes, many of which were suspected of 





"*\ 



W- 





PM 



y 



*. 



w 



I - 





Figure 1. Electron micrograph of an isolated cell fraction containing numerous smooth 
(ribosomc-frce) vesicles of onion stem. Obtained by centrifugation between 20- and 30,000 
rpm and fixed for 18 hours in 6</<< buffered glut ar aldehyde with post fixation for 24 hours 
in l''/o osmium tetroxide. Section stained with lead citrate and uranyl acetate (Method 1). 
Isolation Medium 3. X 32,600. 

Figure 2. As in Figure 1 except section staining Method 2 using phosphotungstic acid 
plus chromic acid which differentiates the darkly staining plasma membrane vesicles (PM) 
from other smooth vesicles (V) presumed to represent tonoplast fragments. X 32,600. 



100 Indiana Academy of Science 










i 

1 t 



xw t 






© 



Figure 3. Onion stem tissue fixed in 6% glut ar aldehyde for 20 hours and section stained 
using Method 3. This electron micrograph shows that in the whole cell, the plasma mem- 
brane (PM) is the only cell component staining with the phosphotungstic acid-chromic 
acid mixture except for cell wall which occasionally stains (CW). Nuclei (N), tonoplast 
(T) and mitochondria (M) are unstained. X 6,400. 



being- derived from plasma membrane (Fig. 1). Dictyosomes and mito- 
chondria were also present. However, using ordinary lead and uranyl 
ion-based staining methods (Method 1), vesicles tended to look alike; for 
example, those derived from plasma membrane could not be distinguished 
from those derived from the vacuolar membrane (tonoplast). 

To identify plasma membrane-derived vesicles, it was necessary to 
use a staining procedure specific for plasma membrane. The PTA-chromic 
acid procedure of Roland (26) was adapted for use with isolated cell 
fractions (Fig. 2). When applied to whole plant cells, the plasma mem- 
brane appeared as a dense, darkly-stained line (Fig. 3). The only other 
cell component staining with the phosphotungstate-chromic acid procedure 
(Method 2 or Method 3) was the cell wall. Nuclear membranes, mitochon- 
dria, proplastids, Golgi apparatus, endoplasmic reticulum and tonoplast 
did not stain. 



Cell Biology loi 



GA 




f 




% 



Figure 4. Plasma membrane fragment (PM) at higher magnification showing dark-light- 
dark trilamellar pattern of the membrane. Isolated dictyosome of the Golgi apparatus 
(GA) showyi for comparison is imstained. Conditions as in Figure 1. X 100,000. 



A similar range of specificity was encountered with the isolated cell 
fractions. Using- staining methods (2) or (3), it was easy to recognize 
which of the fragments of the pellets were plasma membrane-derived 
(Fig. 2). Figures 1 and 2 were obtained from sections of the same tissue 
block. At higher magnifications (Fig. 4), the stained membranes retained 
the dark-light-dark trilamellar staining pattern characteristic of the 
plasma membrane when fixed in situ and stained by conventional methods. 
No differences in staining quality could be detected among the cell frac- 
tions isolated in each of the three homogenization media used for onion 
stem. 

We estimate our plant fraction to be no more than 50% plasma 
membrane-derived, but a comparison of some of the properties of the 
crude preparations with purified plasma membrane fractions from rat 
liver shows them to be different (Table 1). A striking difference con- 
cerns the presence of the so-called marker enzymes for plasma mem- 
brane from mammalian sources. These include 5'-nucleotidase, Mg++- 
adenosine triphosphatase and the Na + -K + -stimulated Mg+ + -adenosine 
triphosphatase (2) which are concentrated in rat liver plasma membrane 
but appear to be absent from the corresponding plant fraction. The results 
are exemplified by studies of 5'-nucleotidase using AMP as substrate. 
When examined at pH 7.0, a near optimum pH for the rat liver enzyme, 



102 



Indiana Academy of Science 



total homogenate of onion stem had a specific activity of 0.06/umoles 
inorganic phosphorous /hr/mg protein as compared with 1 to 2 
^moles/hr/mg protein for rat liver homogenates. In contrast, the plant 
enzymes exhibited a pH optimum at about pH 5.5 with a maximum spe- 



0.3 



0.2 - 



— 0.1 



h- 




v O 






0.0 


> 




— CD 




o* 
<£ 




y^ 


1.0 


u_ ._ 




— CL 




O 




CO . 


0.8 



- A. ONION STEM 


- B. RAT LIVER 


\ A\ Total 


x ' X ^x Total 
/ \ Particulate 


1 J \ Homogenate 


1 / V 


\/ 


- / x\ 




x \ 




/ \ 


~~x v Total Particulate 


' — y 

' • TOtQl 

^ — ^•^^•^Homogenate 


x x~x x 


V ^x 


s ^*^ 


1 1 1 1 x 1 1 


Ir^l 1 1 1 1 1 - 



o 

^ 0.6 
0.4 




0.2 - 



0,0 

TH N PP M PM ER S TH N M ER GA PM S 

CELL FRACTION 

Figure 5. Distribution of phosphatidic acid phosphatase among cell fractions of onion 
stem and rat liver: 

A. pH relationship of total homogenate and particulate fraction of onion stem. 

B. pH relationship of total homogenate and particulate fraction of rat liver. 

C. Distribution of activity among cell fractions of onion stem. 

D. Distribution of activity among cell fractions of rat liver. 

(Key to labeling: TH = total homogenate. N = nuclei-; PP .-_ proplastid-; M — mito- 
chondria-; PM = plasma membrane-; ER = endoplasmic reticulum (microsomes)-; and 
GA — Golgi apparatus-rich cell fractions. S = soluble fraction of the cytoplasm.) 



Cell Biology 103 

cific activity at that pH of 0.16 /-orioles /hr/mg protein. With both the 
5'-nucleotidase and the adenosine triphosphatase, 93'/- of the activity 
was in the soluble fraction. The remaining !'/< of the combined particu- 
late fractions was removed by washing and no activity was detected in 
purified plasma membrane fractions. 

Phosphatidic acid phosphatase is a fourth enzyme associated with 
the rat liver plasma membrane (17). This enzyme showed a distribution 
in the plant similar to that of the 5'-nucleotidase with more than 90% 
of the activity being soluble, but the particulate fraction was active 
(Fig. 5A). In contrast, the rat liver phosphatidic acid phosphatase 
activity was largely particulate (Fig. 5B). The rat liver enzyme showed 
a single pH optimum at pH 6.25, whereas the plant preparations showed 
three optima: in the vicinity of pH 5.5; pH 6.0 to 6.25 and pH 8.0 to 
8.5. The phosphatidic acid phosphatase activity at pH 6.25 for the plant 
was more or less equally distributed among the various cell fractions 
and not concentrated in the plasma membrane fraction as in rat liver 
(compare Figs. 5C and 5D). 

Invertase is one enzyme associated with the cell surface of plant 
cells (30). Specific activities of various fractions in rmxmoles reducing 
sugar/hr/mg protein were: 0-10,000 rpm fraction = 1.8; 10 to 20,000 
rpm fraction (plasma membrane rich) = 0.4; and 20 to 30,000 rpm 
fraction (microsomes) = 0. 

The purity of the fractions precluded a complete phospholipid analy- 
sis, but sphingomyelin, a major constituent of the rat liver plasma mem- 
brane, was absent from the plant fraction (Table 1). Both fractions 
were sterol rich although the predominant sterols of plants (stigmasterol 
and /^-sitosterol) are different from those of rat liver (largely choles- 
terol). 

Discussion 

Junctional complexes, hexagonal subunit patterns, 100 A knobs and 
sialioproteins, all reliable markers for the animal plasma membrane (1, 
2, 14), were absent from the plant cell surface. Thus, the first problem 
was to devise a way to recognize plasma membrane in an isolated cell 
fraction in the absence of positional relationships to other cell com- 
ponents. 

By treating glutaraldehyde-fixed pellets with or without osmium post 
fixation with a mixture of phosphotungstic acid and chromic acid (PTA- 
CA procedure), the plant plasma membrane stained specifically and 
characteristically (26). This staining pattern appears to be applicable 
to plasma membranes from plants other than onion (26). 

Certain enzymes that have proven to be reliable markers for the 
animal cell membrane appear to be absent from the plant cell surface. 
Histochemical studies have revealed that certain nucleoside mono-, di- 
and triphosphatases may be present in the plasma membrane of some 
plants (29), but even when present they vary from cell to cell type and 



104 



Indiana Academy of Science 



Table 1. Comparison of the properties of plasma membrane-rich cell 
fractions isolated from plant and mammalian sources. 



Constituent 



Specific Activity or Amount 
Rat Liver Onion Stem 



5'-Nucleotidase 

Mg++ -adenosine 

triphosphatase 
Mg + +-Na + -K + -adenosine 

triphosphatase 
Phosphatidic acid 

phosphatase 
Sphingomyelin 

Sialic acid 
Sterols 
Buoyant density 



50 /xmoles/hr/mg 

protein 
45 ^moles/hr/mg 

protein 

10 /umoles/hr/mg 

protein 

1 //mole/hr/mg 

protein 

18% of total lipid 

phosphorous 

32 mum/mg protein 

5-6% of dry weight 

1.16-1.18 



not detected 

not detected 

not detected 

0.1 /xmoles/hr/mg 

protein 

not detected 
not detected 

present 

est. 1.4* 



*Determined for membranes stabilized by the addition of 50 mM glu- 
taraldehyde to the initial homogenization medium. 



are often difficult to demonstrate. We found low levels of AMP hydro- 
lyzing activity in the soluble fraction with a pH optimum in the acid 
range. They probably represent unspecific acid hydrolases. A specific 
membrane-associated 5'-nucleotidase was not demonstrated. With rat 
liver plasma membrane, Widnell (32) showed that 5'-nucleotidase is a 
lipoprotein specifically complexed with sphingomyelin. When the sphingo- 
myelin is removed, enzyme activity is lost. It is of interest that the 
plant plasma membrane fractions contain neither sphingomyelin nor 
5'-nucleotidase. 

With phosphatidic acid, the substrate is not only hydrolyzed by 
the acid hydrolases of the soluble fraction from onion stem but also by 
an activity of the particulate fraction. This activity has a pH optimum 
in the range 6.0 to 6.25, similar to that for rat liver. A small fraction 
of this activity appears to be associated with the plasma membrane- 
rich fraction, but is present in all cell fractions and cannot be used as 
a marker enzyme as it is with rat liver. 

Our results support the contention of Emmelot and Benedetti (2) 
that the plasma membrane is a highly differentiated membrane system 
with its characteristics and composition varying according to species, 
cell type and perhaps even the metabolic state of the cell. Our com- 
parisons of rat liver vs. onion stem show the plasma membranes to be 
different in many ways and that species differences are reflected in the 
characteristics of the surface membrane. 



Cell Biology 105 

Literature Cited 

1. Benedetti, E. L., and P, Emmelot. 1967. Studies on plasma membranes. IV. The 
ultrastructural localization and content of sialic acid in plasma membranes isolated 
from rat liver and hepatoma. J. Cell Sci. 2 :499-512. 

2. Benedetti, E. L., and P. Emmelot. 1968. Structure and function of plasma membranes 
isolated from liver, p. 33-120. In A. Dalton and F. Haguenau [eds. 1 The Membranes. 
Academic Press, New York. 

3. Cheetham, R. D., D. J. MORBE, and W. N. Yunghans. 1970. Isolation of a Golgi 
apparatus-rich fraction from rat liver. II. Enzymatic characterization and com- 
parison with other cell fractions. J. Cell Biol. 44 :492-500. 

4. Clowes, F. A. L., and B. E. Juniper. 1968. Plant Cells. Blackwell Scientific Publica- 
tions, Oxford and Edinburgh. 546 p. 

5. Edleman, J., and M. A. Hall. 1965. Enzyme formation in higher-plant tissues. 
Development of invertase and ascorbate-oxidase activities in mature storage tissue of 
Helianthus tuberosus L. Biochem. J. 95:403-410. 

6. Eisinger, W. R., and D. J. MORRE. 1968. The effect of sulfhydral inhibitors on plant 
cell elongation. Proc. Indiana Acad. Sci. 77:136-143. 

7. Emmelot, P., C. J. Bos, E. L. Benedetti and Ph. Rumke. 1964. Studies on plasma 
membranes. I. Chemical composition and enzyme content of plasma membranes 
isolated from rat liver. Biochim. Biophys. Acta 90:126-145. 

8. Epstein, E. 1960. Spaces, barriers, and ion carriers : Ion absorption by plants. Amer. 
J. Bot. 47 :393-399. 

9. FlSKE, C. H., and Y. Subbarow. 1925. The colorimetric determination of phosphorus. 
J. Biol. Chem. 66 :375-400. 

10. Frey-Wyssling, A., and K. Muhlethaler. 1965. Ultrastructural Plant Cytology. 
Elsevier Press, Amsterdam, The Netherlands. 

11. Jorgensen, K. H., and H. Dam. 1957. An ultra micromethod for the determination of 
total cholesterol in bile, based on the Tschugaeff color reaction. Acta Chemica Scand. 

11:1201-1208. 

12. Keenan, T. W., and D. J. Morre. 1970. Phospholipid class and fatty acid composi- 
tion of Golgi apparatus isolated from rat liver and comparison with other cell 
fractions. Biochemistry 9 : 19-25. 

13. Lowry, O. H., N. J. Rosenbrough, A. L. Farr and R. J. Randall. 1951. Protein 
measurement with the Folin phenol reagent. J. Biol. Chem. 193 :265-275. 

14. Middleton, A. E., R. Cheetham, D. Gerber and D. J. Morre. 1969. Adenosine 
mono-, di- and trinucleotidase activities of rat liver eytornembranes. Proc. Indiana 
Acad. Sci. 78:183-188. 

15. Morre, D. J. Cell wall dissolution and enzyme secretion during leaf abscission. 
Plant Physiol. 43:1545-1559. 

16. Morre, D. J., and W. R. Eisinger. 1968. Cell wall extensibility: Its control by auxin 
and relationship to cell elongation, p. 625-645. In F. Wightman and G. Setterfield [eds.] 
Biochemistry and Physiology of Plant Growth Substances. Runge Press, Ottawa, 
Canada. 

17. Morre, D. J., T. W. Keenan and H. H. Mollenhauer. In press. Golgi apparatus 
function in membrane transformations and product compartmentalization : Studies 
with cell fractions from rat liver. Proc. 1st Intern. Sym. Cell Biol. Cytopharm., 
Venice, Italy, 1969. 

18. MORRE, D. J., and J. L. Key. 1967. Auxins; pp. 575-593. In F. H. Wilt and N. K. 
Wessels [eds.] Methods in Developmental Biology. Thomas Y. Crowell Co., New York. 

19. Morre, D. J., R. L. Hamilton, H. H. Mollenhauer, R. W. Mahley, W. P. Cun- 
ningham, and V. S. Lequire. 1970. Isolation of a Golgi apparatus-rich fraction 
from rat liver. I. Method and morphology. J. Cell Biol. 44 :484-491. 

20. Morre, D. J., and H. H. Mollenhauer. 1964. Isolation of the Golgi apparatus from 
plant cells. J. Cell Biol. 23 :295-305. 



106 Indiana Academy of Science 

21. Morrk, D. J., H. H. Mollenhauer and J. E. Chambers. 1965. Glutaraldehyde stabili- 
zation as an aid to Golgi apparatus isolation. Exp. Cell Res. 38 :672-675. 

22. Neville, D. M. I960. The isolation of a cell membrane fraction from rat liver. J. 
Biophys. Biochem. Cytol. 8:413-422. 

23. Ordin, L., and M. A. Hall. 1967. Studies on cellulose synthesis by a cell-free oat 
coleoptile enzyme system : Inactivation by airborne oxidants. Plant Physiol. 42 :205-212. 

24. Parsons, J. G., and S. Patton. 1967. Two dimensional thin-layer chromatography of 
polar lipids from milk and mammary tissue. J. Lipid Res. 8 :696-699. 

25. Reynolds, E. S. 1963. The use of lead citrate at high pH as an electron-opaque stain 
in electron microscopy. J. Cell Biol. 17:208-212. 

26. Roland, J. C. 1969. Mise en evidence sur coupes ultrafines de formations poly- 
saecharidiques directement associees au plasmalemme. C. R. Acad. Sc. Paris 269D :939- 
942. 

27. Rohser, G., A. N. Siakotos and S. Fleischer. 1966. Quantitative analysis of phospho- 
lipids by thin-layer chromatography and phosphorous analysis of spots. Lipids 1 :85-86. 

28. Smith, S. W., S. B. Weiss and E. P. Kennedy. 1957. The enzymatic dephosphoryla- 
tion of phosphatidic acids. J. Biol. Chem. 228 :915-922. 

29. Poux, N. 1967. Localisation d'activites enzymatiques dans les cellules du meristeme 
radiculaire de Cuctimis sativus L. I. Activites phosphatasiques neutres dans les cellules 
du protoderme. J. Microscopie 6:1043-1058. 

30. Straus, J. 1962. Invertase in cell walls of plant tissue cultures. Plant Physiol. 
37:342-348. 

31. Villemez, C. L., J. M. McNab and P. Albersheim. 1968. Formation of plant cell 
wall polysaccharides. Nature 218:878-880. 

32. Widnell, C. C, and J. C. Unkeless. 1968. Partial purification of a lipo-protein with 
5'nucleotidase activity from membranes of rat liver cells. Proc. Nat. Acad. Sci. 
61:1050-1057. 



Di- and Tri-nucleotidase Activities of Rat Liver Cytomembranes 1 

R. D. Cheetham and D. J. Morre, Purdue University 

Abstract 

Enzymatically active fractions of endoplasmic reticulum, Golgi apparatus and plasma 
membrane were isolated from rat liver. Without exception, the di- and trinucleotide phos- 
phatase activities of the Golgi apparatus fraction were intermediate between those of the 
endoplasmic reticulum and plasma membrane. These data support the proposal that the 
Golgi apparatus serves as a site of endomembrane differentiation from endoplasmic 
reticulum-like to plasma membrane-like. 

Morphological evidence has provided the basis for a proposal that 
Golgi apparatus function as sites of cytomembrane differentiation in 
the formation of membranes which are plasma membrane-like, beginning 
with an input of membrane constituents from endoplasmic reti2ulum 
and implying a progressive change in the composition or arrangement 
of constituents within the membrane (5, 10). An understanding of the 
functional significance of the transitional nature of the Golgi apparatus 
requires information on the chemical and enzymatic composition of 
Golgi apparatus relative to membranes of the endoplasmic reticulum 
and plasma membrane. 

In this report, we compare nucleoside di- and triphosphatase ac- 
tivities of endoplasmic reticulum, Golgi apparatus and plasma mem- 
brane. With every nucleotide tested, the activity of the Golgi apparatus 
is intermediate between that of the endoplasmic reticulum and plasma 
membrane. 

Materials and Methods 

Methods for isolation (2, 9, 10), yield and purity (8) of the endo- 
plasmic reticulum, Golgi apparatus and plasma membrane fractions have 
been described. Enzyme assays were at 37° C under conditions where 
activity was proportional to time of incubation and protein concentration. 
Protein was determined by the method of Lowry et al. (7) and inorganic- 
phosphate by the method of Fiske and Subbarow (4). All assays were 
with nucleotides as sodium salts in medium A-l of Emmelot et al. (3), 
pH 7.4. 

Substrates were of the highest purity obtainable from the suppliers 
indicated: inosine-5'-diphosphate (IDP); thy midine-5'-diphosphate 
(TDP); thymidine-5'-triphosphate (TTP) and guanosine-5'-diphosphate 
(GDP) [Calbiochem]; adenosine-5'-diphosphate (ADP), cytidine-5'-di- 
phcsphate (CDP), uridine-5'-diphosphate (UDP), inosine-5'-triphosphate 
(ITP), uridine-5'-triphosphate (UTP) [Sigma]; adenosine-5'-triphosphate 
(ATP), guanosine-5'-triphosphate (GTP) [Nutritional Biochemicals]. 



1 Purdue University AES Journal Paper No. 3910. Work supported in part by grants 
from the NSF GB 1084 and 707S. 

107 



108 



Indiana Academy of Science 



Results and Discussion 

The relative nucleoside di- and triphosphatase activities of the 
isolated fractions are shown in Figure 1. The specific activity (/xmoles 
Pi/hr/mg protein) of each fraction was divided by the specific activity 
of the corresponding total homogenate. This corrects for fluctuations in 
feeding and age of the animals and for variations in the isolation pro- 
cedures. The activity of the Golgi apparatus-rich fraction is intermediate 
between that of endoplasmic reticulum- and plasma membrane-rich frac- 
tions even though the activity is increasing, as with uridine triphos- 
phatase; decreasing, as with guanosine diphosphatase; or unchanging, as 
with inosine diphosphatase; in going from endoplasmic reticulum to 
plasma membrane. 

Morphological evidence for a transitional nature for the Golgi ap- 
paratus has come from studies with the fungus Pythium ultimum (5) 



12 



UJ 

|s 

LU 
CD 

o 
o 

X 

_l 

< 4 
o 



go 



I 



1 



m 






1 



I 



^ 



i 

I 



H_i 



II 



m m 



I 



5 



I 



s 






o 



4- 



ERGAPM 
ATPase 



ER GAPM 

UTPase 



ER GA PM 

GTPase 



ER GA PM 

ITPase 



ER GAPM 

CTPase 



i 



IZZl 



m 



m 



m 



m. 



11 




L. 



I 



ERGAPM 

TTPase 




ERGAPM 
ADPase 



ER GA PM 
UDPase 



ER GA PM 

GDPase 



ER GA PM 

IDPase 



ER GA PM 

CDPase 



Figure 1. Relative specific activities of hydrolysis of nucleoside di- and tri- 
at pH 7.-4 by endoplasmic reticulum (ER)-, Golgi apparatus (GA)- and plasma 
(PM)-rich cell fractions from rat liver. Relative specific activity is the ratio 
activity (umo/es iP/hr/my protein) of each fraction to that of the total h 
(Abbreviations for substrates are explained in the text.) 



ER GA PM 

TDPase 

phosphates 

membrane 
of specific 
omoociiatc. 



Cell Biology 109 

and rat liver (10). These studies show a progressive change in both 
membrane staining and membrane thickness across stacked cisternae 
from a morphology which resembles that of the endoplasmic reticulum 
to a morphology which is plasma membrane-like. The transitional nature 
of the Golgi apparatus is also reflected in the lipid composition of these 
fractions where the Golgi apparatus is again intermediate between the 
endoplasmic reticulum and the plasma membrane (10). The enzyme ac- 
tivities presented here lend further support to our proposal that the 
Golgi apparatus functions in the transformation of membranes from 
endoplasmic reticulum-like to plasma membrane-like. 

Golgi apparatus are not known to be sites of protein synthesis (6) 
but enzyme proteins might be added to or removed from the membranes 
during transformation. If transfer of proteins from endoplasmic reticu- 
lum to Golgi apparatus occurs in the formation of plasma membrane- 
like secretory vesicles, then it appears that the process must involve a 
selective transfer so that certain proteins become concentrated in the 
secretory vesicle membranes whereas others are not incorporated (1). 
As an alternative, certain activities of enzyme proteins derived from 
endoplasmic reticulum might be progressively activated or inhibited con- 
current with the changes in lipid and carbohydrate composition of the 
membranes (1). Hopefully, pulse labeling studies involving one or more 
of the protein constituents of plasma membrane can be used to provide 
a direct test of the concept of membrane flow in rat liver. 

Literature Cited 

1. Cheetham, R. D., T. W. Keenan, S. Nyquist, and D. J. Morre. 1969. Biochemical 
comparisons of endoplasmic reticulum-, Golgi apparatus-, and plasma membrane-rich 
cell fractions from rat liver in relation to cytomembrane differentiation. J. Cell Biol. 
43:21a. 

2. Cheetham, R. D., D. J. Morre, and W. N. Yunghans. 1970. Isolation of a Golgi 
apparatus-rich fraction from rat liver. II. Enzymatic characterization and comparison 
with other cell fractions. J. Cell Biol. 44 :492-500. 

3. Emmelot, P., C. J. Bos, E. L. Benedetti, and P. H. Rumke. 1964. Studies on plasma 
membranes. I. Chemical composition and enzyme content of plasma membranes isolated 
from rat liver. Biochim. Biophys. Acta 90:126-145. 

4. Fiske, C. H., and Y. Subbarow. 1925. The colorimetric determination of phosphorus. 
J. Biol. Chem. 66:375-400. 

5. Grove, S. N., C. E. Bracker, and D. J. Morre. 1968. Cytomembrane differentiation 
in the endoplasmic reticulum-Golgi apparatus-vesicle complex. Science 161:171-173. 

6. LeBlond, C. P. 1965. General conclusions, p. 321. In: C. P. LeBlond and K. B. 
Warren [eds. ] The Use of Radioautography in Investigating Protein Synthesis. 
Academic Press, New York. 

7. Lowry, O. H., N. J. Rosenbrough, A. L. Farr, and R. J. Randall. 1951. Protein 
measurement with the Folin phenol reagent. J. Biol. Chem. 193:265-275. 

8. Middleton, A. E., R. D. Cheetham, D. Gerber, and D. J. Morre. 1969. Adenosine 
mono-, di- and trinucleotidase activities of rat liver cytomembranes. Proc. Indiana 
Acad. Sci. 78:183-188. 

9. Morre, D. J., R. L. Hamilton, H. H. Mollenhauer, R. W. Mahley, W. P. Cun- 
ningham, R. D. Cheetham, and V. S. Lequire. 1970. A Golgi apparatus-rich cell 
fraction isolated from rat liver. I. Method and morphology. J. Cell Biol. 44:484-491. 

10. Morre, D. J., T. W. Keenan, and H. H. Mollenhauer. In press. Golgi apparatus 
function in membrane transformations and product compartmentalization : Studies 
with cell fractions isolated from rat liver. Proc. 1st Intern. Symp. Cell Biol, and 
Cytopharmacol., Venice, Italy, 7-11 July 1969. 



A Model Mosaic Membrane: Cytochrome Oxidase 1 

T. F. Chuang, Y. C. Awasthi and F. L. Crane, 
Purdue University 



Abstract 

During purification of a lipid-free cytochrome oxidase, beef heart mitochondrial mem- 
branous structure was broken down through use of the nonionic detergent Triton, 
together with KC1. After removal of lipid, the purified cytochrome oxidase appears as 
90A diameter globules or as assemblies of rod-like structure with the same thickness. 
Upon addition of mitochondrial phospholipid, structural transformation of the enzyme 
occurs and enzyme activity is enhanced. Three phases of transformation in structure 
depending on the amount of phospholipid added are observed: 1) at 0-20 g Atom P of 
phospholipid/mole cytochrome oxidase there is transformation from swollen particles or 
rod-like elements (200A) to sheets made up of 50A globules; 2) at 20-65 g Atom P/mole, 
the 50A globules are evenly dispersed to form mosaic membrane vesicles; 3) at higher than 
65 g Atom P/mole, there is excess phospholipid around the membrane vesicle, and the 
subunits appear to be 30-50A in diameter. During this transformation there is a regular 
increase in activity which attains maximum at 65 g Atom P/mole in oxidase and this 
activity remains unchanged even at higher phospholipid concentration up to 235 g Atom 
P/mole. We conclude that the oxidase protein globules form a mosaic membrane with 
phospholipid interspersed between the globules. 

Introduction 

Purified cytochrome oxidase has been shown to be able to form 
membranes. Deoxycholate will disperse the membrane structure and after 
removal of deoxycholate the solubilized cytochrome oxidase is organized 
to form membrane vesicles (8). In the study of the relationship between 
membrane formation and ionic strength, we observed that the membrane 
formation could be accomplished in the presence of high concentration 
of nonionic detergent (10). By employing nonionic detergents, Tritons, 
together with KC1, a lipid-free cytochrome oxidase has been prepared 
(2, 10) and it appears as 90 A globules. In the investigation of phospho- 
lipid function in the cytochrome oxidase reaction system (2) we found 
that this lipid-free preparation is rather inert to its substrate, cytochrome 
c. Maximal activity can not be obtained using even lipid-cytochrome c 
(5) as its substrate if phospholipid is absent on the enzyme side (2). 
Enzyme activity toward either cytochrome c or lipid-cytochrome c in- 
creases on addition of phospholipid micelles to the enzyme. In the pres- 
ent study we discuss the relationship between the enzyme activity and 
amount of phospholipid present as well as the structural transforma- 
tion of the enzyme on addition of mitochondrial phospholipids. 

Materials and Methods 

Mitochondria were isolated by the method of Low and Vallin (7) and 
were stored in concentrated suspension at — 20° C before use. Enzyme 
and phospholipid were prepared from mitochondria which were less than 



1 Supported under grant AM04663 from the National Institutes of Arthritis and 
Metabolic Diseases. F. L. Crane is supported by Career Grant K6-21.839 from the National 
Institutes of General Medical Science. 

110 



Cell Biology 111 

two weeks old. Lipid-free cytochrome oxidase was prepared as described 
by Sun et al. (10 with some modification (2). One mM succinate was 
added before sonication and 0.02 m Tris pH 7.4 was used instead of 
phosphate buffer. This enzyme contained only 2-3 g Atom P/mole cyto- 
chrome oxidase or 2-3% phospholipid (w/w) and it has heme a concen- 
tration of 8-9 rn.fi moles /mg of protein as determined by its differential 
spectrum using a millimolar extinction coefficient AA at 605-630 m/x at 
13.1 (12). 

Phospholipid micelles and the lipid-free cytochrome oxidase were 
soluble and could not be centrifuged out at 108,000 x g for 1 hr. To 
study the amount of phospholipid bound to cytochrome oxidase during 
membrane formation, different amounts of phospholipid were added to 
40 mg cf lipid-free cytochrome oxidase and the mixture was diluted 
with 0.25 m sucrose, 0.02 M Tris, pH 7.4 to 3 mg of protein per ml. The 
mixture was sonicated at maximal output with a Branson's Sonifer for 
five 30 sec intervals and centrifuged at 108,000 x g for 30 min. The pellet 
was resuspended in sucrose-Tris buffer and centrifuged at the same 
speed. After three washes all free lipid and free cytochrome oxidase 
which were not in the membrane were washed out. 

Protein was determined according to Yonetani (13). Molecular 
weight of cytochrome oxidase was based on the value of 72,000 (3). 
Phospholipid micelles were prepared by the sonication method of Flei- 
scher and Fleischer (6). Cytochrome oxidase activities were assayed 
polarographically in the following mixture at pH 6.5: 16 mM potassium 
phosphate; 10 mM potassium citrate; 0.80 mM EDTA; 13 mM potassium 
ascorbate; 1.11 mM TMPD; 15 ^M cytochrome c in a total volume of 1.8 
ml using a Gilson oxygraph at 37° and the enzyme in the range of 
5-20 M g. 

Tritons and cytochrome c Type III were purchased from Sigma 
Chemical Company. Other chemicals were reagent grade. 

Samples negatively stained with phosphotungstate, pH 6.8 were 
prepared for electron microscopy according to the procedure of Cunning- 
ham and Crane (4) and observed with a Philips EM 300. 

Results 

Membranes of mitochondrial electron transport particles (ETP) 
(Fig. 1) were split into two membranous fractions by using Triton X 114 
and KC1 as described by Prezbindowski et al. (9). Further purification 
of the green fraction results in a lipid-free cytochrome oxidase which 
appears as 90a diameter globules or assemblies of rod-like structure 
with the same thickness (Fig. 2). This lipid-free enzyme has low activity 
which can be recovered upon addition of phospholipid. Figure 3 shows 
the relationship between the enzyme activity and the amount of phos- 
pholipid added. Three phases of the enzyme activity and enzyme struc- 
ture related to the amount of phospholipid added can be observed. At 
low amount of phospholipid (0-20 g Atom P/mole cytochrome oxidase), 
the 90a globules and rod-like elements began to swell to the size of 



112 



Indiana Academy of Science 







'■,: 









MM^Sm. mam 

photung state. X 103,700. Marker 



WmmmBBMW 

FIGURE 1. Mitochondrial membrane fragments. Phi 
1000.1. 

Figure 2. Purified lipid-frec cytochrome oxidase showing 90 A globules and rod-like struc- 
ture made of 90.1 subunits. Phosphotung state. X 103,700. Marker 1000.1. Arrow A indicates 
90 A enzyme globules; arrow B, a rod-like element with 90 A subunits. 



Cell Biology 



113 



200a in diameter and gradually to transform to a small sheet of mem- 
brane made up to 50a globules (Fig. 4). Addition of phospholipid 
rapidly changed the activity of the enzyme (Fig. 3). With addition of 
more phospholipid to the enzyme, (20-65 g Atom P/mole cytochrome 
oxidase), the lipoprotein globules became organized to form a larger 
sheet of membrane with the same size (50a) globules visible in surface 
views (Fig. 5). 




50 100 150 200 250 

PHOSPHOLIPID ADDED (g ATOM P/MOLE CYTOCHROME OXIDASE) 

Figure 3. Enzyme activity of a purified lipid-free cytochrome oxidase preparation and 
the amount of phospholipid added. Phospholipid was added and incubated at cold for 5 
minutes before dilution for assay. This enzyme contained 1.5 mg of Triton x 100 per mg 
of protein. 



At phospholipid concentrations higher than 65 g Atom P/mole cyto- 
chrome oxidase, excess of phospholipid was observed around the mem- 
brane vesicles and the summit was reduced to about 30-50A diameter 
(Fig. 6). Activity at this stage remains at the maximum and addition of 
more phospholipid (up to 235 g Atom P/mole cytochrome oxidase) did 
not increase activity. 

When the lipid-free enzyme was sonicated with increasing amounts 
of phospholipid micelles (up to 235 g Atom P/mole cytochrome oxidase) 
membrane obtained by centrifugation followed by several washes to re- 
move unbound phospholipid and cytochrome oxidase showed regular in- 
crease in its phospholipid content up to a constant value of 65 g Atom 
P/mole cytochrome oxidase which corresponds to about 40% of lipid 
in the membrane (Fig. 8). Any excess phospholipid was washed out and 
in no experiment did the value of bound phospholipid in the washed mem- 
brane increase beyond this constant value. No phospholipid micelle struc- 



114 



Indiana Academy of Science 




Figure 4. Cytochrome oxidase with 10 g Atom P of phospholipid per mole of enzyme 
(or 10% phospholipid, w/w) showing swollen particles, rod-like elements and small mem- 
brane with 50 A subunits. Phosphotung state. X 103,700. Marker 1000. 1. Arrow points on 
a swollen rod-like element with 50 J subunits. 

Figure 5. Cytochrome oxidase with .11 g Atom P of phospholipid per mole of enzyme 
(or 25% phospholipid, w/w) showing vesicle membrane with 50A subunits phosphotung- 
slate. X 103,700. Marker 1000A. 



Cell Biclogy 



115 




Pil 



■-i*.,? 











Figure 6. Cytochrome oxidase with 235 g Atom P of phospholipid per mole of enzyme 
(70% phospholipid, w/w) showing excess of phospholipid micelles around membrane 
vesicles. Phosphotwng state. X 103,700. Marker 1000.1. 

Figure 7. Membraneous cytochrome oxidase after washing away the free phospholipid 
micelles. Excess of phospholipid (235 g Atom P of phospholipid per mole of cytochrome 
oxidase) was added to lipid-free cytochrome oxidase as described in the text and centri- 
fuged at 108,000 x g for 30 minutes and 3 washes to remove unbound phospholipid. Phos- 
pholipid content decreased to 65 g Atom P of phospholipid }>< r mole of cytochrome oxi- 
dase. No micelle is seen around membrane vesicles. Note 50.1 subunits. Phosphotung state. 
X 103,700. Marker 1000A. 



116 



Indiana Academy of Science 



uj 

>- ^ 

N uJ 
2 CO 

UJ < 

o 

2 x 
H o 

UJ 

o 

X 

o 

o 

h- 
>- 
o 



I < 

O cp 



80 



60 



40 



20 



50 100 150 200 250 

PHOSPHOLIPID ADDED (g. ATOM P/M0LE CYTOCHROME OXIDASE) 

Figure 8. Amount of phospholipid bound to cytochrome oxidase. 



ture was observed around the edge of the membrane vesicles after wash- 
ing (Fig. 7). Subunits of the membrane consistently showed a diameter 
of 50a and completely filled the sheet surfaces. 

Enzyme with phospholipid added (vesicle membraneous form) was 
found to be much more stable to heat treatment than the lipid-free en- 
zyme. Figure 9 shows the activity change of both lipid-free and lipid- 
added enzymes after incubation at different temperatures. After incuba- 
tion at the temperature and time indicated, lipid-free enzyme was incu- 
bated with phospholipid at 4° C for 10 minutes before enzyme assay. 
Lipid added cytochrome oxidase has 90%, 81% and 19% of the original 
activity after 20 minutes incubation at 30° C, 38° C and 50° C respec- 
tively while the lipid-free enzyme had only 75%, 40% and 0% of original 
activity. The cytochrome oxidase of washed beef heart mitochondria was 
also found to be more stable to heat treatment (Fig. 10). 



Discussion 

The capability of cytochrome oxidase to form membrane was demon- 
strated earlier by McConnell et al. (8) and our laboratory (10). Mem- 
brane formation demonstrated in the present study requires the presence 
of both protein and phospholipid. Phospholipid content is also a critical 
factor for enzyme activity. Deficiency in lipid content causes the enzyme 
molecules to cluster together and may prevent access of substrate to the 
active site. As can be seen in Figures 2 and 4, on addition of phospholipid 



Cell Biology 



117 



100 




TIME OF INCUBATION (MIN) 

Figure 9. Stability of Upid-free and lipid-added cytochrome oxidase activity. Enzyme 
was incubated at temperature indicated at protein concentration of 1 mg/ml. 120 g Atom 
P of phospholipid/mole enzyme was added before or after incubation. Solid lines are the 
lipid-added cytochrome oxidase activity; dotted lines the Upid-free cytochrome oxidase 
activity. Control activity as 100% was it 1.2 ninoles Q-i uptake /min/mg. 



micelles the enzyme swells and 50a lipoprotein globules are loosely 
packed into a sheet to form membrane vesicles. In this sheet form the 
substrate has ready access to active sites on the enzyme as shown by 
the high cytochrome oxidase activity. 

Further addition of phospholipid micelles (20-65 g Atom P/mole 
enzyme) allows a more even dispersion of the enzyme in the form of 
mosaic membrane vesicles with 50a globules evenly dispersed as shown 
in Figure 5. Thin sectioning of this membrane shows unit membrane 
structure which is 50-55A thick. Once the phospholipid requirement for 
the membrane formation and maximal activity is reached, further addi- 
tion of phospholipid does not further increase the activity. The negative 
staining data show excess phospholipid micelles around the edge of 
membrane vesicles. The globule size on the membrane is somewhat re- 
duced to 30a. This reduction of apparent globule size may indicate an 
excess of phospholipid among the lipoprotein globules. 

Sonification facilitates the interaction of cytochrome oxidase pro- 
tein and phospholipid to form tightly bound membrane. The unbound 
lipid is still in the state of micelles and can be easily removed by wash- 



Hi 



Indiana Academy of Science 













38° 


inn * 








n 




IUU 




o 






o 






^ 


A__^ 






V 










-^^^45° 


h- 80 


— y 










> 












h- 












o 












< 












- 1 ,-~ 












O 60 


— 










(T 












h- 












Z 












o 












o 












fe 40 


- 








____^ 50° 


h- 












z 












LJ 












u 












cr 












IH 20 


— 










Q_ 

n 




1 1 


i 


1 


1 1 



10 20 30 40 

TIME OF INCUBATION (MIN) 



50 



60 



Figure 10. Stability of cytochrome oxidase in a beef heart mitochondrial preparation. 
Mitochondria were adjusted to protein concentration of U mg/ml and incubated at indi- 
cated temperature. Control activity as 100% was 5:20 ^motcs 02 uptake / min j mg protein. 



ing. A consistent value of 60-70 g Atom phosphorus of phospholipid per 
mole of cytochrome oxidase was obtained after washing when an excess 
of phospholipid was added (Fig. 8). The same saturated value of phos- 
pholipid was obtained for the maximal activity of cytochrome oxidase 
(Fig. 3). The electron micrograph (Fig. 7^ showed 50a globules on the 
washed membrane surface and no phospholipid micelles were observed. 

The 90a particles characteristic of the lipid-free enzyme are equiva- 
lent to a molecular weight of 290,000 daltons if the protein density is 
assumed as 1.25. Molecular weight of monomer cytochrome oxidase is 
72,000 (3). Thus, our lipid-free cytochrome oxidase may exist as a 
tetramer. Ball et al. (1) suggested that cytochrome oxidase might be a 
tetrapolymer consisting of four identical hemoprotein submolecules. 
Takemori et al. (11) also suggested that their Emasoln.to solubilized 
cytochrome a as a pentamer based on sedimentation experiments. After 
cytochrome oxidase is reorganized into membrane the globule size ap- 
pears to be 50-60A which is equivalent to the molecular weight of 
50,000-85,000. This value agrees with the molecular weight of 72,000 
suggested by Criddle and Bock (3). Takemori et al. (11) suggested that 
cytochrome oxidase existed as a monomer with only one heme group 



Cell Biology 



119 



per molecule in mitochondrial particles, it tended to polymerize after 
extraction from mitochondria with bile salt and purification with am- 
monium sulfate. Similar polymerization probably also occurs in our 
preparation and depolymerization occurs again during reconstitution. 






50 A ~j~~ 







Q PROTEIN GLOBULE 
T PHOSPHOLIPID 



Figure 11. A model mosaic membrane: cytochrome oxidase. 



In a study of the heat stability of the enzyme we found that the 
membraneous form of cytochrome oxidase (made by addition of phos- 
pholipid to lipid-free cytochrome oxidase) was more stable than the 
lipid-free enzyme. Figure 9 shows the cytochrome oxidase activity after 
incubation at a series of temperatures for enzyme to which phospholipid 
was added before or after incubation. The membraneous cytochrome 
oxidase has 90%, 81% and 19% of the control activity while the incu- 
bated lipid-free enzyme has only 75%, 40% and 0% of the control ac- 
tivity after incubation for 20 minutes at 30° C, 38° C and 50° C re- 
spectively. Stability of cytochrome oxidase activity in mitochondria is 
much greater than in the purified lipid-free state (Figs. 9 and 10). 

Wallach and Gordon (14) proposed that in membrane formation the 
various subunits were assembled to form a protein lattice penetrated by 
cylinders of lipid. We propose that a model membrane made from inter- 
action of purified membrane protein, cytochrome oxidase, and phospho- 
lipid may simulate in part the membrane structure in mitochondrial 
cristae as in Figure 11. Upon addition of phospholipid to lipid-free cyto- 
chrome oxidase a complex forms between these two components, and 
structural transformation occurs parallel to and in as pronounced a 



120 Indiana Academy of Science 

fashion as the enhancement of enzyme activity. Protein molecules, as 
membrane subunits, are assembled to form mosaic structure with phos- 
pholipid. After formation of protein-lipid complex, the enzyme is much 
more insensitive to heat treatment than the lipid-free enzyme. The mem- 
brane of cytochrome oxidase may represent a model for some biological 
membranes in vivo. Further comparison between this membrane model 
and the mitochondrial cristae membrane, with respect to sensitivity to 
enzyme attack (e.g., phospholipases and proteases) as well as com- 
parison of molecular conformations will be necessary before we can tell 
how closely the model membrane reflects the structure in cristae. 



Literature Cited 

1. Ball, E. G., C. F. Strittmatter, and O. Cooper. 1951. The reaction of cytochrome 
oxidase with carbon monoxide. J. Biol. Chem. 193 :635-647. 

2. Chuang, T. F., F. F. Sun, and F. L. Crane. 1969. Proteolipids VI. The dual role of 
phospholipid in cytochrome oxidase reaction. J. Bioenergetics. 

3. Criddle, R. S., and R. M. Bock. 1959. On the physical-chemical properties of water- 
soluble cytochrome oxidase. Biochem. Biophys. Res. Commun. 1:138-142. 

4. Cunningham, W. P., and F. L. Crane. 1965. Identification of isolated cellular mem- 
branes by electron microscopy. Plant Physiol. 40:1041-1044. 

5. Das, M. L., and F. L. Crane. 1964. Proteolipids I. Formation of phospholipid- 
cytochrome c complexes. Biochemistry 3 :696-700. 

6. Fleischer, S., and B. Fleischer. 1967. Removal and binding of polar lipids in 
mitochondria and other membrane systems, p. 406-433. In R. W Estabrook and M. Z. 
Pullman [eds. 1 Methods in Enzymology, Vol. X. Academic Press, New York. 

7. Low, H., and I. Vallin. 1963. Succinate-linked diphosphopyridine nucleotide reduc- 
tion in submitochondrial particles. Biochim. Biophys. Acta 69:861-374. 

8. McConnell, D. G., A. Tzagoloff, D. H. MacLennan, and D. E. Green. 1966. Studies 
on the electron transfer system LXV. Formation of membranes by purified cytochrome 
oxidase. J. Biol. Chem. 241 :2373-2382. 

9. Prezbindowski, K. S., F. J. Ruzicka, F. F. Sun, and F. L. Crane. 1968. A double 
layer of protein in mitochondrial cristae. Biochem. Biophys. Res. Commun. 31:164-169. 

10. Sun, F. F., K. S. Prezbindowski, F. L., Crane, and E. E. Jacobs. 1968. Physical 
state of cytochrome oxidase, relationship between membrane formation and ionic 
strength. Biochim. Biophys. Acta 153:804-818. 

11. Takemori, S., I. Sekuzu, and K. Okunuki. 1961. Studies on cytochrome a. Physico- 
chemical properties of purified cytochrome a. Biochim. Biophys. Acta 51 :464-472. 

12. Vanneste, W. H. 1966. Molecular proportion of the fixed cytochrome components of 
the respiratory chain of Keilin-Hartree particles and beef heart mitochondria. Biochim. 
Biophys. Acta 113:175-178. 

13. Yonetani, T. 1961. Studies on cytochrome oxidase III. Improved preparation and 
some properties. J. Biol. Chem. 236:1680-1688. 

14. Wallach, D. F. W., and A. Gordon. 1968. Lipid protein interactions in cellular mem- 
branes. Fed. Proc. 27:1263-1268. 



CHEMISTRY 

Chairman: James E. George, DePauw University 
Joseph R. Siefker, Indiana State University, was elected Chairman for 

1970 

ABSTRACTS 

Determination of the Hydrolysis Constant of the Stannous Ion by an 
Electromotive Force Method. Lusan G. Ong and Eugene Schwartz, 
DePauw University. — The formal hydrolysis constant 

K= [SnOH + ] [H + ] 
Sn2 + 
for the hydrolysis reaction 

Sn^+ + H 2 = SnOH+ + H + 

was obtained at 25° C and at an ionic strength of unity by an electro- 
motive force method employing the concentration cell 

/ HC1O 4 (1.00m) / HC10 4 (XF) / 

Sn(Hg) / / Sn(ClO 4 ) 2 (0.020F) / (Hg)Sn 

/ Sn(ClO 4 ) 2 (0.020M) / NaClO 4 (1.0-XM) / 

in which x was varied from 0.04 to 0.96. The liquid junction potentials 
for the above cell were obtained from voltages of the cell 

Ag, AgCl/HCl (1.0m) / HCI(Xm) / AgCl, Ag 



/ HCI(Xm) / 

/ NaCl(l.O-XM) / 



The hydrolysis constant of the stannous ion was found to be 1.5± 0.5 X 
10 2. 

The Infrared Spectra of Coordination Compounds. James Nowak, Robert 
Williams and James George, DePauw University. — The infrared spec- 
tra of [Co(NH 3 )-Cl]Cl 2 , [Co(ND H ) n Cl]Cl 2 , [Cr(NH ;s ) 5 Cl]Cl 2 and a 
series of related compounds containing ions of the form [M(NH a ) 5 X] p + 
(where M = Co(III) or Cr(III) and X = H 2 0, F", Cr, Br N0 2 ", SCN" and 
C0 3 2 ) have been obtained. The spectra of these compounds are very 
similar to those of other metal-ammonia systems which have been more 
extensively studied. The absorptions arising from the vibrations of the 
coordinated ammonia molecules occur in the regions: 31,500 cm 1 ; 1,600- 
1,650-1; 1,300-1,350 cmi; and 800-850 cnri. The metal-ammonia stretch- 
ing frequencies are near 500 cm* 1 . The coordinated polyatomic ligands 
could, in all cases studied, be distinguished from their ionic counterparts. 

pH Dependent Isotope Effects on the Flavin Enzyme L-Amino Acid Oxi- 
dase. Robert L. VanEtten and David S. Page, Purdue University. — 
The effects of changes in pH and of deuterium substitution upon the 
kinetic parameters of the reaction of L-amino acid oxidase with L-leucine 
have been examined. The Michaelis-Menten parameter K IH was measured 
for the L-amino acid oxidase — L-leucine system as a function of pH in 
H 2 at 25, 30 and 35° C in the pH range 5.5 to 9.5. Treatment of the 

121 



122 Indiana Academy of Science 

data according- to Dixon's rules yields pK values for a group situated in 
the active center (i.e., in the ES complex) and for a group in the free 
enzyme. The temperature dependence of these pK values allows the 
calculation of an enthalpy of ionization of 7-8 kcal/mole for this ioniz- 
able group. This value together with the observed pK values lead to the 
conclusion that a histidyl group is involved in the catalytic interaction 
between enzyme and substrate. Studies of the pK dependence of the 
system in D 2 at 25° C reveals substantial D 2 solvent isotope effect, 
particularly at lower pD values. A major part of this D 2 effect is at- 
tributable to effects upon the ionization constant of one or more cata- 
lytically active groups. Such an interpretation is consistent with the ob- 
served shift of 0.7 units in the pK value of the catalytically important 
group described previously, as well as the changed pKa of free imida- 
zole in D 2 0. When DL-[a--H]leucine is employed as a substrate there is 
observed a strikingly pH dependent kinetic isotope effect, with a value 
of kn/k D of 4.0 being observed at pH values less than 6.5, but approaching 
1.0 at pH values above 8.5. Stopped-flow experiments have been con- 
ducted to locate the particular steps in the action mechanism which are 
affected by deuterium substitution. 

A Kinetic Study of the Decarboxylation of Duroic Acid in Sulfuric Acid 
Solutions. John T. Snow and Gerald R. Barker, Earlham College. 
— The decarboxylation of diortho-substituted benzoic acids in strong 
mineral acids has long been known and is of some synthetic use, but the 
mechanism of the reaction is not well understood. Kinetic data are pre- 
sented on the decarboxylation of duroic acid in various concentrations of 
sulfuric acid and at several different temperatures. The reaction is first 
order in duroic acid, but the rate dependence on sulfuric acid concen- 
tration allows no simple interpretation. Several possible mechanisms 
are presented. 

Kinetic data were obtained by measuring evolved carbon dioxide. 
The development of the gasometric technique is discussed and a com- 
parison is made of the gasometric method and the infrared and ultra- 
violet spectrophotometric methods of analysis. 

Molecular Complexes of Bromine and Various Substituted Carbostyrils 
and their Hydrolysis Products. Donald J. Cook, DePauw University. — 
A number of 1:1 molecular compounds between bromine and various 
carbostyrils have been prepared. These compounds are fairly stable at 
room temperature and are inert in nonpolar solvents. However, in the 
presence of water, hydroxide ion, or pyridine, the molecular compound 
is destroyed resulting in the substitution of a bromine on the three or 
six position of the carbostyril. Identification of the substituted bromine 
carbostyrils was made by infrared studies and by independent synthesis 
of the compound by known methods. The position of the bonded bromine 
molecule on the carbostyril is not known but some studies with the 
nuclear magnetic resonance have been initiated to determine the struc- 
ture. 

The molecular compounds have also been shown to be brominating 
agents for alkenes and ketones. 



Temperature Dependence of E° for the Daniel] Cell 

Sister Barbara Buckbeei, Ronald E. Surdzial^ and Clyde R. Metz-% 

Indiana University 

Abstract 

EMF measurements for the cell Zn/ZnSOj (1m)//CuS04 (lM)/Cu were made at various 
temperatures over the range of 0-50 °C. The equation describing the voltage as a function 
of temperature is 

E°(volt) = (1.1028 ± 0.0026) — (0.641 ± 0.425)10- 3 t + (0.72 ± 0.87)10- 5 t 2 
where t is the Celsius temperature. This result compares favorably with electrochemical 
measurements reported in the literature for similar cells, but the derived values of ^G°, 
AS° and /\H° differ considerably with accepted thermodynamic values. This disagreement 
probably results from the presence of a residual voltage and corresponding temperature 
coefficient inherent in the cell from incomplete elimination of the liquid junction potential. 

The conventional Daniell cell, Zn/Zn- + //Cu2 + /Cu, is often used in 
general chemistry courses to demonstrate the calculation of overall cell 
voltage by combining half-cell potentials. Using the data commonly 
found in textbook tables (3) for the Zn/Zn-+ and Cu/Cu 2 + couples, 
-0.763 v and 0.337 v, respectively, the predicted value of E° at 25 °C is 
1.100 v. Upon careful construction of the cell using 0.5m solutions of 
the nitrates or sulfates of the metals, the observed voltage is somewhat 
lower than the predicted value (11). 

For a cell containing the sulfates of the metals, the overall cell 
potential E(exp) is given by 

RT 

E(exp)=E° — lnQ + E(LJ) [1] 

nF 

where Q, the thermodynamic activity quotient, is denned as 

Q =: a Zn -+ a Cu /a Cu 2+ a Zn = a2 ZnS0 ^ a Cu /a2 CllS0 ^ a Zn [2] 

and E(LJ) is the liquid junction potential arising from the ZnS0 4 -KCl- 
CuS0 4 interfaces. The activities of the metals in Equation [2] are unity 
by convention and the product of concentration and mean activity co- 
efficient gives the mean activities for the salts. Using the values given 
by Robinson and Stokes (8) for the activity coefficients, the emf contri- 
bution in Equation [2] for the activity term is negligible for the 1m 
concentrations. 

As written above, the cell contains a salt bridge to minimize the 
liquid junction potential. Maclnnes (5) states that E(LJ) is negligible 
provided the ionic mobilities of the cation and anion in the bridge are 
equal. If a saturated KC1 bridge is used, this equality is nearly achieved 
at 25° C. Thus any observed voltage should represent E° for the 
chemical reaction of interest: Zn + Cu-+ = Zn- + -f Cu. 



1 Current address : St. Joseph High School, Brooklyn, N.Y. 

2 Current address: Highland High School, Highland, Indiana. 

3 Current address : Indiana University-Purdue University at Indianapolis. 

123 



124 Indiana Academy of Science 

Experimental 

The half cells consisted of strips of Zn and Cu metal (10x1x0.1 cm) 
dipping into 1M solutions of ZnS0 4 and CuS0 4 , respectively. To reduce 
heat transfer from the solutions to the air, the electrodes were thermally 
insulated using 1-inch thicknesses of plastic foam. The half cells were 
joined by a salt bridge consisting of a saturated KC1 solution suspended 
on a strip of chromatographic filter paper. To reduce possible interaction 
between the metals and KC1, the electrodes and a small quantity of 
solution were isolated from the bulk solution by enclosing them within 
glass tubing which was drawn to a capillary tip. At each temperature 
fresh solutions and a newly-constructed salt bridge were used. All chemi- 
cals were of reagent grade quality and standard quantitative procedures 
were used in preparation of the solutions. 

The complete Daniell cell was placed in a constant temperature bath 
and the temperatures of the half cells were monitored by separate 
thermometers. Once temperature equilibrium was reached, the tempera- 
ture was recorded to the nearest 0.05° C (corrected for stem immersion) 
and the open-cell voltage was read from a Honeywell Potentiometric 
Voltmeter, Model 852, to the nearest 0.1 mv. At the sensitivity used, the 
input impedance was 10 Mil, so negligible current was drawn from the 
cell. 

Results 

The data appear in Figure 1. Using the method of Bennett and Frank- 
lin (1), the regression coefficients for a linear and a quadratic dependence 
on temperature were determined. An analysis of variance indicated the 
quadratic term to be significant and the corresponding equation is 

E° (volt) = (1.1028 ± 0.0026) — (0.641 ± 0.425)10 3 t+ 
(0.72 ± 0.87)10-5 t2 [3] 

where t is the Celsius temperature and varies from 0° to 50° C. The 
standard deviation is 1.36 mv and the estimated errors in the regression 
coefficients are calculated on a 95% probability limit basis. 

The maximum random error in the voltage resulting from tempera- 
ture measurement is negligible, ± 0.02 mv. The error resulting from 
temperature gradients is estimated as ± 0.5 mv based on observations 
made with nonisothermal cell conditions. 

The important thermodynamic properties AG , AS° and AH can be 
derived from Equation [3]. These are 

AG°(kcal/mole) = -nFE° = -50.863 + 0.0296 t — 0.33x10 -3 t-' [4] 

AS° (gibbs/mole) = -d(AG°)/dT = -29.6 + 0.66 t [5] 

AH°(kcal/mole) = AG° + TAS° = -58.948 + 0.1813 t + 

0.33x10-3 t2. [6] 

Table 1 contains values of these properties and accepted thermodynamic 
values (10). Although the quadratic equation for AG° allows the esti- 
mation of AC° p , the confidence limits in the regression coefficients are 
too large to provide reliable values. 



Chemistry 



125 




% L 



10 20 30 40 50 
TEMPERATURE (°C) 



Figure 1. Plot of experimental emfs for the cell Zn/ZnSO',( 1m)//CuSOj,( lu) /Cu. The 
curve is the least squares equation. 



Table 1. Thermodynamic values for the Daniell cell calculated from 

Equations [3] -[6]. 

t (°C) E° (volt) AG° (kcal/mole) AS° (gibbs/mole) AH° (kcal/mole) 






1.1028 


—50.863 




—29.6 




—59.948 




10 


1.0971 


—50.600 




—23.0 




—57.168 




20 


1.0929 


—50.403 




—16.4 




—55.354 




25 


1.0913 


— 50.329 (- 


-50.71)* 


— 13.1(- 


-3.73)* 


— 54.621(- 


-51.82)* 


30 


1.0901 


—50.272 




—9.8 




—53.806 




40 


1.0887 


—50.207 




—3.2 




—52.224 





* The values given in parentheses are accepted thermodynamic values 
(10). 



Discussion 

The calculated value of E° at 25° C from Equation [3] of 1.0913 ± 
0.0026 v is lower than the predicted value of 1.100 v by roughly 9 mv. 
Reported values for E° at 15° C of 1.09337 by Cohen, Chattaway and 
Tombrock (2) for a cell consisting of amalgamated electrodes in satu- 
rated solutions and 1.0962 v (average) by Jahn (4) give similar differ- 
ences between experimental and calculated values. 



126 Indiana Academy of Science 

The value of dE°/dT at 25° C calculated from Equation [3] is 
— 0.289 mv/deg. This value compares favorably to — 0.429 mv/deg 
reported by Cohen, Chattaway and Tombrock (2), to — 0.2 mv/deg re- 
ported by Rosset (9), and to — 0.182 mv/deg listed by deBethune and 
Loud (3) as an experimental determination. These results are consider- 
ably more negative than — 0.083 mv/deg as predicted by combining half 
cell values of dE°/dT based on thermodynamic values given by deBethune 
and Loud (3). The thermodynamic value for AS° in Table 1 corresponds 
to — 0.0809 mv/deg (10). Considering the similarities of the species in- 
volved in the cell, the smaller thermodynamic values appear to better 
express AS for the reaction than do the electrochemical values. 

The source of the discrepancies, 9 mv and 0.2 mv/deg, probably 
results from the last two terms in the expressions for the overall cell 
potential, Equation [1], and the corresponding temperature coefficient 

dE(exp) dE° R d In Q dE(LJ) 

-dT- = ^-^ (lnQ + T -l^ )+ "^ [7] 

which are present because of experimental conditions. 

As mentioned earlier, the emf contribution of the second term in 
Equation [1] is negligible at 25° C and so the 9 mv must be the result 
of the two liquid junction potentials for the ZnS0 4 -KCl and KCl-CuS0 4 
interfaces. Maclnnes (5) gives the following general equation 

soln II 

soln I 

which must be applied at each interface. Unfortunately Equation [8] 
cannot be integrated directly, but Maclnnes (5) describes two approxi- 
mate methods for obtaining values of E(LJ). 

The first approximation is based on the assumptions that ionic 
mobilities are independent of concentration and that activities and 
concentrations are equal. This results in 



E(LJ)=f ±l ^ fli/z ' i >^ i -^> In f-^f- [9] 



RT n Wz,) (Ci — Ci) nC./ii 

n Mi (Ci"— c!) nC ^i 

where /m is the ionic mobility. Using data from Milazzo (7) for Equation 
[9], one obtains 

E(LJ) = E(LJ) + E(LJ) = —1.3 + 1.6 = 0.3 mv 

ZnSO, — KC1 KC1— CuSO^ 

which is considerably less than the 9 mv and has the incorrect sign. 

The second technique for obtaining E(LJ) values from Equation [8] 
is by graphical integration of values of t-f/Cf plotted against Cf between 
the appropriate concentrations. Following the procedure given by 
Maclnnes (5) and assuming that the activities of the sulfate ion in 1m 



Chemistry 127 

ZnS0 4 and in lM CuS0 4 are equal and that the changes in the activities 
of Cl~, SOj' - and K 2 S0 4 passing from the salt bridge to 1m ZnS0 4 and 
lM CuS0 4 to be identical, the total liquid junction potential is given by 

sat KC1 

— RT r O ' Zn ~ + d(C f ) 

E(LJ) = ~- I Vj q~ ~t~ ZnS0 4 ZnSO, 



/ V ZnS0 4 ZnS0 4 

" 1m ZnS0 4 

sat KC1 

S { Cu^+ d (C 

C f 

CuS0 4 CuS0 4 
1m CuS0 4 Llt'J 



■ • >7 

CuS0 4 CuS0 4 / 



Using available data (5)-(8) and making the approximation that the 
activity coefficients for the components of mixtures to be equal to the 
activity coefficients for the pure components at the concentration of the 
component, the graphical integration of Equation [10] gives 

E(LJ) = —38.7 + 38.3 = —0.4 mv. 

Although the magnitude is low, the sign of E(LJ) is correct and if 
proper data were available, Equation [10] should predict values more in 
line with the 9 mv. It is felt, therefore, that the 9 mv difference can be 
accounted for by the value of E(LJ). 

From data reflecting the temperature dependence of the limiting 
ionic conductances (7), Equation [9] can be used to estimate dE(LJ)/dT. 
For a 2% increase in the ionic mobilities for an increase in temperature 
of one degree, the value of E(LJ) increases by 0.1 mv. Thus a significant 
portion of the 0.20 mv/deg discrepancy is accounted for by the dE(LJ) /dT 
term of Equation [7]. No attempt was made to estimate dE(LJ)/dT 
from Equation [10] because of the uncertainty in the assumptions regard- 
ing the choice of data. 

In conclusion, the values of E°, AG°, AS° and AH given in Table 1 
are valid for the experimental Daniell cell which includes, by necessity, 
a liquid junction. 



Acknowledgment 

Two of the authors (SBB and RES) were participants in the Institute 
for High School Chemistry Teachers sponsored by the National Science 
Foundation at Indiana University. 



128 Indiana Academy of Science 

Literature Cited 

1. Bennett, C. A., and N. L. Franklin. 1954. Statistical Analysis in Chemistry and 
the Chemical Industry. John Wiley and Sons, New York. 724 p. 

2. Cohen, E., T. D. Chattaway, and W. Tombrock. 1907. Zur thermodynamik der 
normalelemente. Z. Physik. Chem. 60:706-727. 

3. deBethune, A. J., and N. A. Swendeman Loud. 1964. Standard Aqueous Electrode 
Potentials and Temperature Coefficients at 25°C. Clifford A. Hampel, Skokie, 111. 19 p. 

4. Jahn, H. 1886. Uber die Beziehung von chemischer Energie und Stromenergie 
galvanischer Elemente. Wied. Ann. 28 :21-28. 

5. MacInnes, D. A. 1961. The Principles of Electrochemistry. 2nd. ed. Dover Publica- 
tions, New York. 478 p. 

6. Maron, S. H., and C. R. Prutton. 1958. Principles of Physical Chemistry. The 
Macmillan Company, New York. 789 p. 

7. Milazzo, G. 1963. Electrochemistry. Elsevier Publishing Company, New York. 708 p. 

8. Robinson, R. A., and R. H. Stokes. 1949. Tables of Osmotic and Activity Coefficients 
of Electrolytes in Aqueous Solution at 25°C. Trans. Faraday Soc. 45 :612-624. 

9. Rosset, G. 1904. Daniell Cell as Standard Cell for Technical Purposes. Electrochem. 
Ind. 2:246. 

10. Rossini, F. D., D. D. Wagman, W. H. Evans, S. Levine, and D. Jaffe. 1952. Selected 
Values of Chemical Thermodynamic Properties. Nat. Bur. Stand. Circ. 500. 1268 p. 

11. Young, J. A., and J. G. Malik. 1969. Chemical Queries. J. Chem. Educ. 46:227-228. 



The Study of Complexes of Di-n-butyloxamidine with 
Transition Metals 

Warren E. Hoffman, Maurice Jacobs, Gerhard Kennepohl, Dennis 

W. Parrot, Phillip Reed, Theodore R. Stout and James Sundy, 

Indiana Institute of Technology. 

Abstract 

The determination of solution complexes of di-n-butyloxamidine with Cr(III), Mn(II), 
Fe(III), Co (III), Ni(Il), and Cu(II) was made using Job's method of continuous variation. 

Upon investigation of the chemical reactions of aliphatic oxamidines 
(3, 4), we found that Cu(II) and Ni(II) salts formed solid metal com- 
plexes of the general formula, 

( R-NH-C-C-NH-R ) • MCI, • 2H,0, whereas the Mn(II) salt formed 

II II 
HNNH 

( R-NH-C-C-NH-R ) • MCI, • 4H,0. Attempts at that time to obtain 

II II 
HNNH 

solid complexes with Co(II), Co(III), Fe(II), Fe(III), and Cr(III) failed. 
However the color changes with these salts pointed to the formation of 
solution complexes. Subsequently, a brown Fe(III) solid complex was 
isolated using benzene as solvent (1). 

Recently, we have re-examined the solutions involving the di-n- 
butyloxamidine with Cr(III), Mn(II), Fe(III), Co(II), Ni(II) and Cu(II). 
Applying Job's method of continuous variation (2), we have attempted to 
establish the ligand to metal ion ratio for any complexes that might be 
formed in these solutions. All the aforementioned metal (II) ions form 
complexes in solution with the ligand to metal ion ratio of 2:1. The 
Cr(III) and Fe(III) ions appear to form more than one complex in 
solution. Thus, Job's (2) method is not applicable. 

Assuming a reaction in solution to be pM -f qN = MpNq and deter- 
mining p and q, Job's (2) method shows a plot of change in the absorb- 
ance versus mole fraction to yield a maximum or a minimum at the 
stoichiometric composition of the complex. If the measurements are made 
at several wavelengths, all should yield the same result. The method of 
continuous variation can yield reliable results only when the following 
conditions are observed: 

1) Only a single chemical reaction occurs in the solution. There is 
no association, protolysis, solvolysis, etc., of either the reactants or the 
products. 

2) The law of mass action is applicable in terms of concentration. 

3) The reactants form only one complex. 

129 



130 Indiana Academy of Science 

Methods 

1) A stock solution of di-n-butyloxamidine of 0.01 M (0.001 for 
Fe(III)) was made in anhydrous alcohol, either methanol or ethanol. The 
choice of alcohol made little or no difference in the results. In the case 
of Mn(II), a water solution of about 0.1 m was prepared. Due to the 
ready facility of hydrolysis of the oxamidines, water solutions should 
he avoided except where they can be used immediately. 

2) The metal salts were dissolved in the same solvent and made up 
to the same strength as the corresponding oxamidine solution. The 
Cr(III), Co (II) and Ni(II) salts were all hydrated. The Mn(II) salt was 
dissolved in water. 

3) The spectra of solvent, metal salt solution, and oxamidine solution 
were determined using a Beckman DB spectrophotometer. Comparison of 
these spectra with one of the solution containing both metal ion and 
oxamidine led to a selection of wavelengths at which Job's (2) method 
would be applied. The absorbance of the mixtures as the mole fraction of 
oxamidine varied from 0.00 to 1.00 was read using a Beckman DU 
spectrophotometer. 

4) The room in which the experiments were performed and the in- 
struments housed was held at 68 °F and under 30% relative humidity. 

5) The wavelengths at which Job's method for each of the salts was 
applied are listed in Table 1. 

6) The plot of A versus wavelength for the Ni( II) -oxamidine system 
using methanol as solvent is shown in Figure 1. 

7) The plot of A versus mole fraction of oxamidine is shown in 
Figure 2. 

Figures 1 and 2 are representative of the types of curves obtained 
for all the salts with the exception that for Cr(III) and Fe(III), Figure 
2 had more than one maximum or minimum or both. 

Table 1. Wavelengths for application of Job's Method to salts of 
transition metals. 



Salt 


Wavelengths (mu) 


Cr(III) 


370, 380, 540, 550 


Mn(II) 


370, 380, 400 


Fe(III) 


355, 365, 375 


Co(II) 


390, 400, 410 


Ni(II) 


460, 480, 500 


Cu(II) 


530, 560, 590 



Summary 

Figure 2, which is typical of the data obtained with the M-+ ions, 
indicates a solution complex of oxamidine /M-+ of 2:1. Work is continu- 
ing using other methods to determine the nature of the M ;5 + complexes 
formed in solution. 



Chemistry 



131 



2.4r 

2.2 



2.0- 



ujl.4 

I' 2 
$ 1.0- 

< 

0.8 
06- 
0.4- 
02- 



NiCI 2 Solution 
■Oxamidine Solution 
■Complex Solution 



200 



280 3 20 3 60 400 440 

Wavelength, mu 



480 



520 



560 



600 



Figure 1. Absorbance vs. wavelength for NiCk system. 



132 



Indiana Academy of Science 



uj 
o 



1.3 
1.21- 
U 
1.0 
0.9- 

0&' 
0.7 • 



4$0mu 

-480 mu 
•500mu 




0.1 02 Q3 0.4 05 Q6 07 08 0.9 

Mole Fraction Oxamidine 



Figure 2. Ni-\-2-oxamidine complex. 



Chemistry 133 

Literature Cited 

1. Jay, Theodore. 1964. The Reaction of Ferric Chloride with Di-n-Butyloxamidine. M.S. 
Thesis. Indiana Institute of Technology, Ft. Wayne. 

2. Job, P. 1928. Method of Continuous Variation, Ann. de Chimie 9:113(10). 

3. Woodburn, H. M., and W. E. Hoffman. 1958. The Chemistry of Oxamidines I. J. Org. 
Chem. 23 :262-268. 

4. Woodburn, H. M., R. R. Salvesen, J. R. Fisher, W. E. Hoffman, E. L. Graminski, 
and R. L. Van Deusen. 1967. Metal Complexes of Cyanoformamidines, Oxamidines and 
Oxalimidates. J. Chem. and Eng. Data 12 :615-617. 



ECOLOGY 

Chairman: Thomas S. McComish, Ball State University 
James R. Gammon, DePauw University, was elected Chairman for 1970 

ABSTRACTS 

The Annual Growth Cycle of the Bluegill. Thomas S. McComish, Ball 
State University, and Richard 0. Anderson, University of Missouri. — 
Continuous growth experiments were conducted from fall 1965 through 
1967 with bluegill, Lepomis macrochirus, under similar natural seasonal 
photo-periodicity and temperature fluctuations to those occurring in ponds 
near Columbia, Missouri. Each fish was isolated in an aquarium and fed 
frozen chironomid larvae in excess. Growth in length and weight; food 
consumed; conversion efficiency of live, dry, and protein weights; and 
calories energy were measured monthly. 

Growth of yearling bluegill increased steadily from March to a peak 
in June and July followed by a steady decline to a low in December. 
Food consumed followed a similar pattern. Growth of two-year-old 
sexually mature bluegill started in March, peaked in May, decreased 
steadily to a low in July, followed by a steady increase to a second peak 
in September, and decreased to a low point in November. Food con- 
sumed and conversion efficiencies followed similar patterns. 

The mid-summer slump in growth for two-year-old bluegill was 
correlated with temperature. The May and September growth peaks 
occurred at 20 to 22° C and the mid-summer low at about 27° C. Meta- 
bolic rate was implicated in the growth cycle as a function of tempera- 
ture, season, size, and perhaps sexual maturation in preparation for 
spawning. 

The Effect of Photoperiod on Growth of Bluegill. Paul G. Davidson and 
Thomas S. McComish, Ball State University. — Growth experiments 
were conducted with bluegill, Lepomis macrochirus, for 102 days under 
3 different photoperiods at 26° C. One photoperiod increased from 15.50 
to 19.78 hours daily, another decreased from 15.50 to 12.25 hours daily, 
and a third was held constant at 15.50 hours daily. Growth, food con- 
sumption (mealworms, Tenebrio molitor) , and food conversion efficiency 
were evaluated for bluegill in each set of conditions. 

Under the conditions used in this experiment there was no apparent 
effect of photoperiod on the growth of bluegill. This was true for all 
measurements of growth in length, growth in weight, food consumption 
and food conversion efficiency. It was also true when males and females 
were compared for each of these measurements. 

These results were not expected. A possible explanation is that the 
relatively high temperature of 26° C increased metabolism enough to 
mask the effects of photoperiod. 

135 



136 Indiana Academy of Science 

The Response of Fish to Heated Effluents. James R. Gammon, DePauw 
University. — The specter of temperature elevation of surface waters due 
to electric generating- stations has caused great concern to aquatic 
ecologists because of projected construction estimates. The distribution 
and abundance of adult fish in two three-mile segments of the Wabash 
River have been studied since 1967. One site is bisected by an 860 mega- 
watt station, while the other will receive heated effluents beginning in 
1970. 

During normal summer flows the temperature of the effluent was 
about 7° C higher than the river water. Complete mixing was achieved 
about % mile below the effluent jetty at which point the temperature was 
about 4° C higher. Some species populations were estimated by mark- 
and-recapture techniques using hoop nets and electrofishing apparatus, 
but for most species only relative indices of abundance based on catch 
data could be obtained. 

Consistent differences in the distribution patterns were noted for 
some species which seemed related to temperature. Species which tended 
to avoid the heated water included northern river carpsucker (Carpiodes 
carpio carpio), golden redhorse (Moxostoma erythrurum) , shorthead 
redhorse (M. breviceps), spotted bass (Micropterus punctulatus) , long- 
ear sunfish (Lepomis megalotis megalotis), and sauger (Stizostedion 
canadense). Species which were significantly more abundant in the 
heated water included carp (Cyprinus carpio), buffalo fish (mostly 
Ictiobus babalus), gar (Lepisosteus osseus and L. plato stomas ) , channel 
catfish (Ictalurus punctatus) and flathead catfish (Pilodictis olivanis) . 

Preliminary Experiments on Growth of Bluegill with Varied Feeding 
Frequency and Constant Ration. Thomas E. Mangum, III and Thomas 
S. McComish, Ball State University. — Growth of bluegill, Lepomis 
macrochirus, in length and weight, and food conversion were compared 
between groups of fish fed once and three times daily. Daily ration and 
evironmental conditions were constant between groups. Groups fed 
three times daily showed greater growth and conversion efficiencies in 
most cases. Differences, however, were small and not significant. 

The Relationship Between Growth and Social Hierarchy in the Green 
Sunfish. Ruth A. Wilsey and Thomas S. McComish, Ball State Uni- 
versity. — Growth in weight and length, and food conversion efficiencies 
were compared between similar sized control and paired female green 
sunfish {Lepomis cyanellus) in aquaria. Growth was related to hier- 
archy. Prior residence and size were major factors in hierarchy estab- 
lishment. Interaction between paired fish stimulated growth and food 
consumption to an optimal level beyond which a decline was observed. 
Control and paired fish grew more than subordinate fish. Control fish 
converted food more efficiently than paired fish. Social interaction re- 
sulted in the death of four subordinate fish. This was related, at least 
in part, to elevated maintenance levels and food consumption reductions. 
Growth and conversion efficiency of dominant fish after the death of 
subordinates improved in direct relation to the length of the recovery 
period. 



Ecology 137 

A Taxonomic Survey of the Ostracods of Delaware County, Indiana. 

Daniel R. Goins, Ball State University. — A survey of ostracods (Class 
Crustacea, Order Ostracoda) was taken throughout Delaware County, 
during May and June, 1968. One hundred random samples were taken 
from creeks, drainage ditches, farm ponds, temporary streams and ponds, 
and rivers in this county. A bottom dredge net was used to sample bodies 
of water, and collections were preserved in 50% alcohol in pint jars. 

Sixty-two % of the samples contained ostracods. Many of these 
ostracods were dissected in the laboratory and mounted in permanent 
media on microscope slides. The ostracods were then identified and re- 
corded. Comprehensive surveys of ostracods have been made in Ohio 
and Illinois, but only one collection previously had been reported from 
Indiana. This study increased the known Indiana genera from 3 to 19 
and revealed several new species. Names and descriptions of these new 
genera and species will be published in the near future. 

The Influence of Environmental Factors on the Concentration of Hydro- 
cyanic Acid in Manihot esculenta. Robert D. Hart and Wm. B. Crank- 
shaw, Ball State University. — Manioc or yuca (Manihot esculenta), a 
common cultigen of milpa agriculture in the tropics, varies from site to 
site with respect to the concentration of hydrocyanic acid in the tubers. 

A short term study was conducted to determine the possibility of a 
correlation between acid concentration and microclimatic factors. A 
site was selected on the edge of the Amazon Basin in the rain forest 
of eastern Ecuador to conduct the study. 

Microclimatic factors were measured at 7 stations along a 60 m 
transect through a milpa of manioc. These factors included insolation, 
precipitation, air temperature, soil temperatures, and soil moisture. At 
the termination of the field study, the manioc plant closest to each 
station was removed and assayed for hydrocyanic acid content. Soil 
samples were also taken at each station and analyzed for percent nitro- 
gen, phosphate and potassium. 

Regression analysis indicated that the highest correlation existed 
between insolation and temperatures and acid content. Enzyme action 
converting cyanogenetic glucoside to hydrocyanic acid seems to be in- 
hibited by the stress situation of high air and soil temperatures. 

A Comparison of Dominance Expressions for Tree Species in Foley 
Woods, Edgar County, Illinois. M. T. Jackson, Indiana State University, 
and R. 0. Petty, Wabash College. — Foley Woods is a 120-acre stand 
occupying a till plain depression just north of the Wisconsin terminal 
moraine. The diverse stand (39 tree species) is notable as a western 
edge outlier of American beech near the prairie border. Sugar maple, 
red oak, bur oak, white oak, shellbark hickory, white and green ash, and 
basswood lead in importance. 

Canopy openings created by selective cutting of large trees have 
allowed sugar maple and elm to increase greatly in density in the smaller 
size classes. High densities of small trees give these species a two- 



138 Indiana Academy of Science 

factor importance value index (the average of relative density and 
relative basal area) disproportionately higher than their actual contribu- 
tion to stand dominance. 

To evaluate this disparity, a 10% sample of the south 80 acres was 
tallied by taking forty 1/5-acre line strips (200 X 43.56 feet). All trees 
over 2 inches dbh were recorded and line-intercept crown cover was 
determined for each plot. Density, frequency, basal area, cover, average 
diameter and average basal area were determined for each species. 
Various two-, three- and four-attribute importance value indices, and 
other synthetic indices were examined. The indices were weighed against 
relative cover to determine which integrating expressions best reflect 
the contribution of each species to total stand dominance. A Density- 
Double Dominance Index in which relative density plus relative basal 
area weighted twice were averaged 



/ D 3 + B 3 + B 3 \ 



gave the best approximation of importance. Relative basal area alone 
was the next most accurate expression, followed by the four factor 
index (the average of the relative values of density, basal area, fre- 
quency and cover). 

Testing the Quarter Method against Full Tallies in Old-growth Forests. 

Damian Schmelz, St. Meinrad College. — The quarter method, although 
used with good results by workers interested in the general characteris- 
tics of many stands, has been criticized for its bias for species tending 
toward clumped dispersal, for the amount of work involved in office 
computations of vegetational attributes, for the difficulty in computing 
standard error, and for its relative field-inefficiency when compared 
with other rapid sampling methods. A further criticism of the accuracy 
of the quarter method results from a comparison of data obtained by 
this method with that obtained by full tallies in seven old-growth stands 
in Indiana. The stands ranged in size from 4 to 21 acres. Two or three 
quarter point samples were obtained for each stand, giving a total of 
17 samples for comparison. The number of points in each sample aver- 
aged two per acre. The quarter method consistently overestimated the 
stand basal area, from 1.0% to 99%, averaging 39%. Stand density was 
overestimated in most samples, by as much as 60%. Of the total number 
of species in each stand, from 43% to 73% were recorded by the samples. 
One of the five most important species was missed in five samples, and 
the order among them was different in ten samples. In one sample the 
most important species was different, and in six samples the second most 
important species was different. The quarter method is not judged use- 
ful for any detailed analysis of hardwood forests. 



Bluegill Predation by Three Fish Species 

William J. Gulish, Indiana Department of Natural Resources 



Abstract 

Seven ponds at Driftwood Experimental Station were used to evaluate the ability of 
three species of predaceous fish to control bluegill populations. These were the largemouth 
bass, northern pike and white catfish (Ictalurus catus) . The ponds were stocked in April, 
1967, with 100 predators and 1,000 bluegill per acre. Two control ponds were stocked 
only with bluegill at 1,000 per acre. Four ponds were drained in October, 1967, and the 
populations evaluated. Three of the ponds were refilled and the fish returned. All ponds 
were drained in August, 1968, and the populations evaluated. 

The largemouth bass was the most effective bluegill predator followed by the northern 
pike and white catfish. The effects of intraspecific competition on the dynamics of the 
bluegill populations and its relation to the predator population is discussed. 



Introduction 

The tendency of the bluegill to overpopulate has long been a major 
problem for the fisheries manager. The problem is particularly acute in 
farm ponds, as the stocking of bluegill in these bodies of water is wide- 
spread. This study was conducted to evaluate the ability of three species 
of predaceous fish to control bluegill populations. 



Methods 

Seven ponds at the Driftwood Experimental Station were used in 
the experiments. The ponds range from 0.62 to 2.05 acres in size, have 
maximum depths of 5 feet and are supplied with water from Starve 
Hollow Lake. Screens placed over the inlet valves to prevent the en- 
trance of lake fish were not completely effective. While contamination 
did occur, it was not serious in most of the ponds. 

Three species of predaceous fish, white catfish, northern pike and 
largemouth bass were used. Two ponds each of white catfish-bluegill and 
northern pike-bluegill were stocked while one pond with largemouth 
bass-bluegill and two ponds with only bluegill were used as controls. The 
ponds were stocked during April and May, 1967. A rate of 100 preda- 
tors to 1,000 bluegill per acre was used in all the ponds. Stocking is 
summarized in Table 1. 

Ponds 2, 4, 6 and 7 were drained during October, 1967. Pond 7 con- 
tained a large bass (1.2 lbs.) and a large number of green sunfish and 
crappies so the data were discarded. All predaceous fish and bluegills 
from the three remaining ponds were counted, weighed and measured. 
Both the white catfish and largemouth bass had spawned, but the young 
were easily separated from the original stock by size. The same was 
true for bluegill in all ponds. 

All bluegill were separated by year class. The survivors of the 
original bluegill stock (age 1) were counted and lengths and weights 

139 



140 



Indiana Academy of Science 



Table 1. Stocking of seven experimental ponds at Driftwood Experi- 
mental Station, April 19-May 11, 1967. 



Pond 


Area 




Bluegill 


Northern 


White 


Largemouth 




(Acres) 






Pike 


Catfish 


Bass 


1 


0.62 


No. 
Size 


620 
1.0-2.0" 








2 


1.07 


No. 
Size 


1070 
1.0-2.0" 




107 
2.5-7.0" 




3 


1.07 


No. 
Size 


1070 
1.0-2.0" 




107 
2.5-5.0" 




4 


1.23 


No. 
Size 


1230 
1.0-2.0" 


123 

1.5-4.0" 






5 


2.05 


No. 
Size 


2050 
1.0-2.0" 


205 
1.5-4.0" 






6 


1.37 


No. 
Size 


1370 
1.0-2.0" 






137 
6.0-7.0" 


7 


1.4(1 


No. 
Size 


1400 
1.0-2.0" 









were recorded. Young-of-the-year bluegill were bulk weighed and the 
total number calculated from counts of weighed random samples. 

After processing, all fish (except for Pond 7) were returned to the 
refilled ponds. As many as possible of the contaminating species, such 
as green sunfish, were sorted out and discarded. Handling mortality 
among the large fish was quite low. A high percentage of the young- 
of-the-year bluegill were lost, however. 



Table 2. Species compositioyi and standing crop of fishes in six ponds 
at Driftwood Experimental Station, August, 1968. 







Pond 1 


Pond 2 


Pond 3 


Pond 4 


Pond 5 


Pond 6 


White Catfish 


No./A. 
Lbs./A. 




117 
31.4 


7 
1.7 








Northern Pike 


No./A. 
Lbs./A. 








10 
14.0 


10 

15.4 




Largemouth Bass 


No./A. 
Lbs./A. 












274 
87.1 


Bluegill 


No./A. 


72,684 


29,147 


8,923 


13,429 


17,914 


8,054 




Lbs./A. 


368.2 


109.6 


131.1 


330.6 


177.5 


52.3 


Other Sunfish 


No./A. 


5 


2 


1,785 


2,776 


6,183 


2 




Lbs./A. 


1.1 


0.3 


17.4 


14.5 


54.0 


0.7 


Crappies 


No./A. 
Lbs. /A. 




1 


186 
14.9 




4 
1.1 




Bullheads 


No./A. 
Lbs./A. 


2 
0.6 








1 
0.9 




Total 


No./A. 


72,691 


29,267 


10,901 


16,215 


24,112 


8,330 




Lbs./A. 


369.9 


141.3 


165.1 


359.1 


248.9 


140.1 



Ecology 141 

In August, 1968, all ponds were drained. All bluegill of 4 inches 
total length or longer were sorted into 1-inch groups, counted and 
weighed. A minimum of 100 fish from each 1-inch group were measured 
to the nearest 0.1 inch. The relative abundance of bluegills below 4 
inches was calculated from random samples because of the high num- 
bers of these fish. A minimum of 100 fish from the 2 and 3-inch groups 
were measured to the nearest 0.1 inch. The remaining bluegills were 
then bulk weighed. No measurements were taken from fish in the 1-inch 
group, which included all fish less than 2.0 inches long. However, aver- 
age weights of the 1-inch group fish were determined by counting and 
weighing a minimum of 500 fish. All predaceous fish were sorted into 
inch groups, weighed and measured. 

It was possible to separate the different year classes by their length 
distributions. Scale samples taken as cross checks showed this method 
to be sufficiently accurate. 



Results 
Bluegill Mortalities 

Mortality rates for the original stock of bluegill in the study ponds 
were calculated from the tables in Ricker (4) and are given in Table 3. 
These rates were similar in both white catfish ponds. The same was true 
in both northern pike ponds. 

Table 3. Survival, seasonal mortality rate (a) and instantaneous rate 

of natural mortality (i) for the original bluegill stock in six ponds at 

Driftwood Experimental Station, 1967-68. 









Predator 








None 
Pondl 


White Catfish 


Northern Pike 


Bass 




Pond 2 


Pond 3 


Pond 4 


Pond 5 


Pond 6 


No./A. stocked 


1000 


1000 


1000 


1000 


1000 


1000 


No./A. 10-67 




936 




751 




140 


a (4-67 to 10-67) 




0.064 




0.249 




0.860 


i (4-67 to 10-67) 




0.07 




0.29 




1.97 


Days (4-67 to 10-67) 




L83 




174 




1S4 


No./A. 8-68 


SS4 


86] 


951 


347 


444 


104 


a (4-67 to 8-68) 


0.116 


0.139 


0.049 


0.653 


0.556 


0.896 


i (4-67 to 8-68) 


0.12 


0.15 


0.05 


1.06 


0.81 


2.26 


Days (4-67 to 8-68) 


475 


470 


467 


454 


453 


490 


a (10-67 to 8-68) 




0.080 




0.538 




0.257 


i (10-67 to 8-68) 




0.08 




0.77 




0.30 


Days (10-67 to 8-68) 




287 




280 




306 



Table 3 indicates that there was little predation by the white cat- 
fish on the original bluegill stock. Change in bluegill mortality rate in 
Pond 2 between drainings was negligible, indicating a uniform predation 
rate over the study period. 



142 Indiana Academy of Science 

A Saprolegnia infection in March, 1968, drastically reduced the white 
catfish population in Pond 3, since 41 were found dead from March 19 
to 23. When drained in August, 1968, only seven white catfish were 
found in the pond. Several white catfish also died of the infection in 
Pond 2 along with a few bluegills in both ponds. The fungus infection 
probably caused the decrease in the 6-inch bluegill group in Pond 2 
(Table 4). 

The lack of Age I fish in Pond 2 (Table 5) is a result of the 1967 
draining which almost completely eliminated the 1967 bluegill year class. 

Table 4. Length distribution of white catfish and bluegill from Ponds 

2 and 3, 1967-68. 









Pond 


2 






Pond 


3 






10-19-67 






8-2-68 




7-30-68 


Inch Group 


Catfish Bluegill 


Catfish 


Bluegill 


Catfish 


Bluegill 


1 




6,500 




33 


30,260 






7,541 


2 


7 






6 








17 


3 


1 


12!) 






6 






379 


4 




768 




1 


500 






1,162 


5 


6 


r,4 






402 






354 


6 


9 


50 




10 


18 


1 




92 


7 


14 






5 


1 


1 




3 


s 


17 






15 




3 






9 


19 






20 




1 






Ki 


17 






17 




1 






11 


6 






5 










12 


X 






Id 










13 








•>> 











Table 5. Number per acre, average length and percent total weight 

of bluegill year classes from six ponds at Driftwood Experimental 

Station, 1968. 







Age II 






Age I 






Age 




Pond 


No./A. 


Ave. L. 


% wt. 


No./A. 


Ave. L. 


% Wt. 


No./A. 


Ave. L. 


% wt. 


1 


SS4 


5.3 


24.7 


* 


* 


* 


* 


* 


* 


2 


861 


5.0 


73.7 


6 


3.2 


o.l 


28,280 


1.3 


26.2 


3 


95 1 


5.1 


68.3 


925 


4.1 


29.9 


7,048 


1.3 


1.8 


4 


347 


5.9 


16.4 


10,938 


3.4 


82.3 


2,143 


1.3 


1.8 


r> 


444 


5.0 


21.1 


12,817 


2.7 


74.4 


1,325 


1.3 


4.5 


6 


104 


7.1 


56.9 


102 


5.4 


28.3 


7,853 


1.3 


14.S 



* Not separable with available data. 

Total bluegill production in the white catfish ponds (Table 2) was 
the lowest in any of the ponds except Pond 6, which was overpopulated 
with bass. 



Ecology 143 

Despite a high level of mortality, the northern pike significantly 
reduced the original bluegill stock. This stock declined from 55 to 65% 
over the study period. However, data from Pond 4 indicate that predation 
did not occur at a uniform rate (Table 3). Pike predation on the original 
bluegill stock shows a marked increase with time. Early in the study, the 
small size of the pike made the bluegills relatively invulnerable to them. 
However, the pike grew faster than the bluegill, increasing the latter's 
vulnerability accordingly. Beyerle and Williams (1) reported that north- 
ern pike fed in aquaria showed a preference for the smaller size groups 
of centrarchids. Results from Pond 4 indicate the opposite. Pike in this 
pond appeared to feed selectively on the larger sizes of bluegill, despite 
the fact that the pond contained a huge quantity of smaller size bluegill 
(Table 6). Total weight of the original bluegill stock decreased from 75 
to 68 lb. in the time between the first and second drainings. Average 
increment in length of the original stock in this same time period was 
only 0.9 inch. 

Despite impressive predation on the original bluegill stock, northern 
pike are poorly adapted for controlling bluegill populations. This is 
particularly true in ponds, where their spawning requirements are un- 
likely to be met. A major problem in the pike-bluegill combination is the 
difference in hatching dates between the two species. The pike is an 
early spawner. The young-of-the-year become piscivorous (in this area) 
in late April or early May. 

There are seldom young-of-the-year bluegill available for forage 
before June. During this interval the young pike must feed on the 



Table 6. Length distribution of northern pike and bluegill from Ponds 
U and 5 at Driftwood Experimental Station, 1967-68. 





oup 








Pond 


Jt 








Pond 


5 






10-31 


-67 






8-7-68 






8-5-68 




Inch Gi 


Pike 




BlueKill 


Pike 




BlueKill 


Pike 




Bluegill 


1 








22,218 








2,636 






9,540 


•1 
















1,515 






22,980 


3 








31 








11,603 






3,294 


5 








537 








274 






365 


6 








L6 








150 






39 


7 
















3 






3 


8 






















1 


13 




5 




















14 




2 




















15 




5 














1 






16 












4 






1 






17 


















2 






18 












4 






4 






1!) 












2 






10 






20 












1 






3 






-n 












1 













144 



Indiana Academy of Science 



preceding year class (age I) of bluegill. As is indicated by Table 3, 
vulnerability of this size bluegill to the pike is low. As a result, where 
other forage species are not available, the size of a given year class of 
pike may well be determined by the abundance and size of the pre- 
ceeding year class of bluegill. Where other species are present, pike seem 
to feed on them in preference to the bluegill (5). 



Table 7. Length distribution of largemouth bass and bluegill in Pond 

6, 1967-68. 



10-25-67 



8-27-68 



Inch Group Bass 



Bluegill 



Bluegill 



60,463 



3 

87 
1 02 



112 

1 



18 
121 
23 
22 
73 



10,718 
31 

10 
25 
107 
09 
84 



In terms of both bluegill control and size of fish produced, the large- 
mouth bass-bluegill combination provided the best results (Table 7). 
Because of the relatively large size of the bass in relation to the blue- 
gill at stocking (Table 1), the vulnerability of the bluegill was high. 
The result was a high mortality rate among the bluegill and good growth 
by both species. Total bluegill production in this pond was low (Table 
2) as a result of a large bass overpopulation (274 bass/acre) coupled 
with the destruction of a large portion of the 1967 bluegill year class 
during the fall draining in 1967. 

Bluegill Dynamics 

In most cases, the bluegill carrying capacity of a body of water is 
determined by the food supply. The degree of intraspecific competition 
within the bluegill population is determined by the difference between 
the standing crop and the carrying capacity (2). 

The amount of recruitment into a bluegill population is a function 
of the degree of food competition. If carrying capacities were uniform, 
many problems could be solved easily. Unfortunately, in addition to 
the differences in basic fertility among bodies of water, carrying capacity 
for bluegill may be influenced by relationships within the population. 
Swingle (6) reported an inverse relationship between the percentage 
of large fish in largemouth bass-bluegill populations and the standing 
crop. 



Ecology 145 

Gerking (3) has shown that small bluegill are more efficient in 
using protein for growth than are large bluegill. This gives the smaller 
bluegill an advantage over the larger one in food competition. It also 
suggests that the potential carrying capacity is greater for small blue- 
gill than larger ones, increasing the likelihood of overpopulation. 

Evaluation of Results 

Pond 1 is a good example of what happens in the absence of preda- 
tion and all bluegill control is intraspecific. No length frequency dis- 
tribution or scale samples were taken from the 1-inch group bluegill, 
making positive year class assignment impossible. However, the length 
distribution curve from the 2-inch group, if extended, reaches a peak 
between 1.5 and 2.0 inches. The large average individual size of the 
1-inch group bluegill (321 /lb.) indicates that most of them were members 
of the 1967 year class. Intraspecific competition appeared to severely 
limit bluegill reproduction in Pond 1 in 1968. 

The difference between the standing crop and the carrying capacity 
in Pond 1 was originally very large, resulting in the production of a 
large 1967 year class. This year class expanded to fill the carrying 
capacity, thereby limiting further reproduction or growth. 

The results from Ponds 2 and 3 indicate that the white catfish 
is more of a bluegill competitor than a predator. Both catfish ponds 
remained turbid throughout the growing season. This probably re- 
duced the carrying capacity for bluegill. If white catfish do, in fact, 
reduce the carrying capacity for bluegill, the difference between the 
standing crop (original stock) and carrying capacity was much lower 
than in Pond 1 resulting in less recruitment. 

The northern pike mortality in Ponds 4 and 5 was so high that they 
never achieved a population level capable of controlling the bluegill. As 
in Pond 1, the difference between the standing crop and carrying ca- 
pacity was large, resulting in high bluegill recruitment. The apparent 
selectivity of the pike for larger bluegill tended to aggravate the prob- 
lem. 

Since northern pike rarely reproduce in ponds, their applicability 
for bluegill control in ponds seems very limited. With sufficient effort, 
a pike population might be maintained at a level that would control the 
bluegill population. However, this can be said of almost any predator. 
At present, it appears that the largemouth bass is the best adapted 
species for controlling bluegill population in ponds. 

Pond 6 developed an overpopulation of bass. Bass predation kept 
the bluegill population far below carrying capacity. The standing crop 
of bluegill actually decreased slightly between 1967 and 1968. While 
the production of bluegill recruits under these conditions had to be 
high, there were sufficient bass present to limit their survival to a very 
low level. 



146 Indiana Academy of Science 

Discussion 

Since recruitment into the bluegill population is such an important 
component of population dynamics, the effect of various management 
procedures on it should be kept in mind. Preoccupation with growth 
rate often results in failure to consider the effect of various procedures 
on the entire system. 

A case in point is the stocking rates used for farm ponds. A com- 
monly used rate is 100 fingerling largemouth bass and 100 fingerling 
bluegill per acre. In ponds of average fertility, the original bluegill 
stock will rapidly grow to a large size. They will spawn at age I, pro- 
ducing a tremendous number of young. This very high level of bluegill 
recruitment results in good growth of the original bass population. 
However, it exceeds the ability of the bass to reduce the number of 
bluegill recruits enough to maintain good bluegill growth. If the bass 
are unable to reduce the numbers of the first bluegill year class suf- 
ficiently, this single year class will tie up most of the ponds carrying 
capacity causing low bluegill recruitment the following year. The young 
bass produced at this time find very few bluegill of a vulnerable size 
and a high degree of competition with the bluegill for other food organ- 
isms. Since the original bass stock will have reached desirable size at 
this stage, they are subject to fishing harvest, reducing the number of 
effective predators on the oversize bluegill year class. While the original 
stock of bluegill has also reached desirable size and are being harvested, 
they make up a relatively small portion of the total standing crop and 
their removal does not appreciably increase the growth of the remaining 
bluegill. 

This stocking rate (100 to 100) results in good growth of the 
original fish but makes a bluegill overpopulation likely in short order. 
Since a population that will maintain a high equilibrium yield is desir- 
able, stocking rates must be adjusted so that the bulk of the standing crop 
will be composed of harvestable fish. If a situation producing a high level 
of recruitment is set up, there should be enough predators stocked to 
control it. 

It is nearly impossible to recommend a good stocking rate without 
some knowledge of the carrying capacity of the body of water involved. 
However, if a single fingerling stocking ratio is to be used, it would 
be advantageous to increase the number of bluegill stocked. This would 
slow the growth rate somewhat, and reduce recruitment to a more 
manageable level. 

An alternative is to decrease the survival of bluegill recruits by 
increasing the number of bass stocked. Which method would produce 
the highest fishing yield over an extended time period cannot be pre- 
dicted. 

Another area where factors affecting recruitment are important is 
that of partial kills. Nothing seems more ridiculous to me than attempt- 
ing to destroy nests or young- of -the-y ear in a stunted bluegill popula- 
tion. Since recruitment into a stunted population is low to begin with, 



Ecology 147 

the food made available by their destruction will not appreciably alter 
the growth of the remaining fish. Besides, this reduces the supply (low 
anyway) of vulnerable fish to the young-of-the-year predators. 

A more common practice is the large scale reduction in the bluegill 
population concentrated on the smaller size groups. It should be remem- 
bered here that the same factors that increase growth also increase 
recruitment. Where a large reduction in standing crop is made, precau- 
tions against the recruitment of a massive bluegill year class should be 
taken. This, again, could take different forms. One way would be to 
increase the predator population to accommodate the higher level of 
bluegill recruitment. The other would be to replace the destroyed year 
class (es) with a number of bluegill more suited to the carrying capacity. 



Literature Cited 

1. Beyerle, George B., and John E. Williams. 1968. Some observations of food selec- 
tivity by northern pike in aquaria. Trans. Amer. Fisheries Soc. 97(1):28-31. 

2. Carlander, Kenneth D. 1966. Relationship of limnological features to growth of 
fishes in lakes. Verh. Internat. Verein. Limnol. 16:1172-1175. 

3. Gerking, Shelby D. 1962. Production and utilization in a population of bluegill sun- 
fish. Ecol. Monogr. 32:31-78. 

4. Ricker, W. E. 1958. Handbook of computations for biological computations of fish 
populations. Fisheries Res. Board Canada Bull. 119. 

5. Seaburg, Keith G., and John B. Moyle. 1964. Feeding habits, digestive rates and 
growth of some Minnesota warm water fishes. Trans. Amer. Fisheries Soc. 93(3) :269- 



6. Swingle, H. S. 1961. Some relationships within fish populations causing fluctuations 
in productions. Proc. Pacific Sci. Cong. 10 :43-45. 



Practicality of Endrin as a Fish Toxicant 1 

H. E. McReynolds, U. S. Forest Service 

Abstract 

When fisheries bioassays documented the lethality of endrin to fish at almost ineoiv- 
ceivably low concentrations, the question of the suitability of this pesticide as a replacement 
for rotenone naturally followed. Fisheries workers immediately noted two characteristics of 
this chemical that would make it superior to rotenone as a piscicide : (1) it was cheaper; 
and (2) it was more persistent, thereby more likely to effect a complete kill. 

However, many other characteristics and effects of endrin as a fish toxicant were 
unknown or poorly understood. Experiments were set up in selected southern Indiana ponds 
to test this chlorinated hydrocarbon. Laboratory bioassays with test fish established an 
LCno (Lethal Concentration, 50 r /f ) or TL m (Median Tolerance Limit) of about 1.3 parts 
per billion. Field tests indicated highly disparate Lc's. After testing in a diversity of 
aquatic situations, it was postulated that the amount of suspended particulate matter 
exerted a considerable effect on the lethality of the chemical in field tests. It was surmised 
that the mechanism of lowered toxicity in more tui'bid situations was the adsorption 
of the chemical to organic (and possibly inorganic) suspensoids. Verification of these 
postulations was not pursued, since other endrin experiments of a public health nature 
were beginning to point out the potential dangers of endrin in potable water supplies. 
In addition, the duration of toxicity of the pesticide showed evidence of extending beyond 
that desired in the ideal fish toxicant, especially in the sediments. Later, more thorough 
experiments have generally tended to establish the validity of these surmisals. It was 
concluded that despite the economy and kill efficiency of endrin, other undesirable and 
dangerous characteristics outweighed the kill-cost attributes. 



Introduction 

At the time of these experiments, fisheries biologists were — and 
still are — looking for a cheaper, more effective replacement for rotenone 
as a fish toxicant. The use of piscicides for eliminating undesirable fish 
populations has become an important tool in fish management. While 
rotenone, a ketone of botanical origin, has certain advantages of low 
toxicity to mammals and low-level of danger to the applicator, it also 
has a number of limitations that fisheries technicians have found frus- 
trating (difficulty in killing resistant species such as bullhead catfish; 
the problem of getting effective vertical dispersal through metalimnetic 
barriers; relatively quick oxidation to non-toxic levels; high toxicity for 
many fish food organisms; and relatively high cost). 

When biologists began to notice that organochlorine pesticides were 
displaying piscicidal potencies many times greater than rotenone, first 
reactions were dismay and apprehension. After the first shock waves 
had passed, however, some biologists began to wonder if this chemical 
lightning might be harnessed for fish management use. Toxaphene, a 
chlorinated camphene, was the first to receive extensive testing as a 
fish toxicant, and was even used — somewhat surreptitously — in a com- 
mercial fish toxicant product. Unfortunately, a toxicity of extended du- 
ration (up to 8 years in Michigan lakes) made its use unfeasible in 



1 This study was supported in part by Federal Aid to Fish Restoration funds (Project 
F-4-R-8, Indiana). 

148 



Ecology 149 

most situations. Interest turned to other members of the generically 
similar chlorinated hydrocarbon pesticides. While Canadian biologists 
began testing thiodan, some American fisheries people became intrigued 
with endrin, an insecticide and rodenticide with toxicity thresholds for 
fish measured in low or fractioned parts per billion! This paper reports 
one of these earlier experiments with endrin. 

Laboratory Aquaria Tests 

Methods 

The preliminary tests were conducted in seven 15-gallon metal frame 
glass aquaria (Fig. 1) having aeration and filtration systems. Diluent 
waters were obtained from the two ponds on the Crosley State Fish and 
Game Area which were to be used for the field tests. These are desig- 
nated merely as Pond 1 and Pond 2 in this paper. Two species of cen- 
trarchids were used as test animals: the bluegill sunfish (Lepomis m. 
macrochirtis Rafinesque), and the redear sunfish (Lepomis microlophus 
[Gunther]). Fish were obtained by seining and were held in the aquaria 
for 10 days before testing began. The fish ranged in size from approxi- 
mately 1 inch to 3 inches. Although this range does not meet size- 
uniformity recommendations (12) no grading was done since these pre- 
liminaries were merely exploratory in nature. Actually, the larger and 
smaller fish were purposely combined in each aquarium to observe any 
difference in size-susceptibility. Two bluegills and 2 redears were placed 
in each of 6 aquaria and 10 fish, a mixture of both species, were held in 
the 7th aquarium as controls. 

The endrin was obtained as a 75% wettable powder and weighed on 
an analytical balance in the following amounts: 0.19, 0.38, 0.75, and 1.50 
g. These weights are for the active ingredient and assume that the manu- 
facturer's assay (75%) is correct. The validity of this assumption was 
not investigated. 

The minute concentrations at which these tests must be conducted 
present a problem in dilution. It was decided that 10 gal of water would 
be used in the aquaria, and calculations of dosage began from there. To 
get the lowest concentration, 0.26 parts per billion (ppb), it was neces- 
sary to mix 0.19 g active ingredient of the endrin powder with 5 gal of 
water, take 1 g of this solution and introduce this amount into the 10 
gal of aquarium water. Addition of another 0.19 g of endrin to the 
original solution gave the 0.52 ppb dosage, and so on up the scale. The 
test solution was drawn off by pipette from the center of the 5 gal 
bottle after extended agitation. A shell vial was weighed on a triple 
beam balance having a sensitivity of 0.01 g, and the weight recorded. 
The balance was set up one more gram and the test solution was pipetted 
into the vial until the correct weight was reached. The contents of the 
vial were then dripped into the aquarium and the aquarium water 
stirred with a clean glass rod. Hourly checks were made by the study 
leader or biologist aides, and notes were made on the reactions of the 
fish. Observations were made from 8:00 AM until 4:30 pm (the work 



150 



Indiana Academy of Science 




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Ecology 151 

day) each day from Monday through Saturday. Water temperatures 
were not controlled during these tests and ranged between 63° F and 
75° F. 

Results i 

Concentrations of 0.26 ppb and 0.52 ppb were run concurrently for 
a period of 96 hours, and at neither level were there any mortalities. 
There were no indications of even slight distress in any fish. These fish 
were observed for several weeks after completion of the 96-hour test, 
and no reaction to the chemical was observed. The next dosage tested 
was 1.0 ppb. During a 48-hour test no mortalities occurred and the fish 
showed no observable evidence of discomfort. At the termination of 
the 48-hour test, another 1.0 ppb was added to Aquarium 3, and the 
concentration in Aquarium 6 was augmented by 2.0 ppb. Assuming that 
the original 1.0 ppb dosage had not been appreciably dissipated during 
the previous 48 hours, concentrations at this time were upwards to 2.0 
ppb in Aquarium 3 and 3.0 ppb in Aquarium 6. After 18 hours, the first 
indication of distress was noted. A small bluegill in Aquarium 6 showed 
the initial signs of disquietude. Symptoms of distress in this fish became 
more intense at each 30-min observation period, and 90 min after the 
first manifestations the fish was dead. Soon afterward, other fish in 
both aquaria began to show reactions to the test solution, and 42 hours 
after increasing the concentration all fish in both aquaria were dead. 

The physiological reactions were similar to those described by 
Public Health Service biologists (14). An increase in respiration and fin 
movements was first noted. Later, fish exhibited annoyed wanderings to 
the surface or into bottom corners of aquaria. As distress became more 
intense, darting movements were noted, with fish occasionally breaking 
the surface and in one instance jumping clear of the aquarium. Later, a 
loss of hydrostatic equilibrium was evidenced by fish swimming abruptly 
to the surface and settling slowly to the bottom. At this time, the test 
animals were oriented at an angle to the horizontal. The body axis was 
a line diverging from the horizontal by 20° -35°, caudal end down. Fish 
near expiration showed little control of their movements; some were 
attempting feeble swimming efforts while on their sides or backs. 

These results indicated that the toxicity threshold of endrin lay 
between 1.0 ppb and 1.52 ppb in these experimental waters and with 
these test animals. However, since the first dosage of 1.0 ppb was fol- 
lowed for only 48 hours, there was a possibility that a more extended 
test period would have resulted in some mortality. Another test at this 
level (1.0 ppb) was run for 72 hours but no mortalities resulted. Again, 
1.0 ppb was added to each aquarium and once more no fish survived an 
additional 72-hour test. From these exploratory tests, it was assumed 
that the toxicity threshold was a value between 1.0 ppb and 2.0 ppb, 
indicating that the field tests should begin with a concentration of 1.0 
ppb (apparently a sublethal dose) and continue upward until concentra- 
tion at which all test fish were killed was reached (Lethal Concentration, 
100%, or simply LC 100 ). It should be noted here that where a chemical's 



152 Indiana Academy of Science 

use as a piscicide is concerned, the LC 100 becomes a more important figure 
than the more commonly used LC 50 (also commonly termed LD 50 Lethal 
Dosage, 50% or tl ih ). This stems from the fact that many chemicals 
show a non-linear regression from the concentration producing total 
extermination to those of sublethal levels (LC 10ft to LC ). Thus, where the 
economic feasibility of a product is one of the important criteria, it be- 
comes of greater concern to establish this level by direct testing rather 
than to estimate it by projection from intermediate levels (lc 50 ). 

Later checks of the 2.0 ppb solutions were made to determine if they 
were still toxic. Two fish were placed in each of 2 aquaria 15 days after 
the solutions were made. All fish failed to survive until the 19th day (96- 
hour test) indicating that dissipation had not progressed to any great 
degree. 

An interesting sidelight which supports results of a Japanese study 
(19) was the presence of cladocerans and planarians in the 3.0 ppb 
solution. Two days after all fish had been killed, these organisms were 
noted in large numbers on the sides of the aquarium. 

Crosley Pond Tests 

During 1960, the endrin study suffered from a low priority, and there 
was no time available for this project until mid-October. On October 13, 
a small pond on the Crosley State Fish and Game Area was treated 
with endrin calculated to produce a concentration of 1.0 ppb. After 120 
hours without any mortality of test fish, another 1.0 ppb of endrin was 
added. This presumably raised the endrin concentration in the pond to 
2.0 ppb, a level that had killed all test fish in laboratory experiments. 

After 144 hours of testing at this level, there were no mortalities, 
nor even discomfort among test fish. It was theorized that this puzzling 
situation resulted from lower temperatures and /or higher turbidities than 
had been present in laboratory tests. This study was not continued dur- 
ing 1961. 

Driftwood Pond Tests 

In 1962 the study was given a higher priority and the testing site 
was moved to the Driftwood Farm Pond Experiment Station where four 
ponds were available. It was felt that the common water source, depth, 
basin morphometry, and soil types, and similar thermal gradients would 
eliminate some of the variables encountered in the heterolimnetic en- 
virons of dissimilar farm ponds. 

Three ponds were used for testing and the fourth served as a con- 
trol. The study here was designed to bracket the minimum LC 100 with a 
concentration below the LC 100 (at 1 ppb) and one calculated to give a 
total kill (4 ppb). These figures were based on the preliminary lab tests 
which showed LC 50 of 1 ppb or more, and no mortality in the Crosley 
pond at an assumed 2 ppb level. It was reasoned that 4 ppb in waters 
of higher temperature would effect a complete kill. A third pond was 
treated with a tremendous dose to observe the duration of toxicity in 



Ecology 153 

cases of miscalculated dosages. Treatment in this pond was at 46 ppb. 
No treatment was made on the control pond. Ponds ranged in size from 
0.62 acres to 1.37 acres. Actual testing began in mid-September when 
water temperatures were near 16° C. 

Pond 1 

Pond 1 (0.62 acres) received the initial treatment. This pond was 
sprayed with a solution calculated to give a 4 ppb concentration of 
endrin (active ingredient). Twenty-four hours previous to the spraying 
of the toxicant, 2 test cages each containing 10 bluegills were placed in 
the pond. One cage was placed in water of 18-inch depth, the other in 
5 feet of water. Test cages were approximately 36" X 36" X 22". Since 
bluegills used as test fish were from adjacent ponds, the 24-hour acclima- 
tion period seemed sufficient. 




Il|lllijililt« 







&00* ' 



v. ~a . 



ill 



mm 



Figure 2. One of the endrin test ponds at the Driftwood, Station showing deepwatt 
cage raised for examination. Dead test fish are visible in cage. 



Test cages were checked periodically to observe fish reactions, but 
for the purpose of recording, data were considered only on a 24-hour 
interval basis (i.e., fish were recorded as having died only at the termi- 
nation of a 24-hour test period). Although the last fish in a test group 
may actually have expired at 42 hours, the LC 100 was recorded as having 
occurred at the 48-hour level. (There seems to be little justification for 
carrying the recording of results to a precision beyond the sophistication 
of the basic technique itself.) 

After 24 hours, 2 fish in the deep water cage had died, but there were 
no mortalities in the shallow water cage although 2 bluegills were in 
extreme distress. At the 48-hour check, all fish in the deep cage were 
dead and there was but one survivor in the shallow cage. The following 
day (72-hour test) the last fish had expired. 



154 Indiana Academy of Science 



Pond 2 



The same procedure was followed in Pond 2 testings as had been 
used in Pond 1 (a deep and a shallow cage each with 10 test fish). This 
pond (1.07 acres) was sprayed with a dosage calculated to give a 
concentration of 1 ppb of endrin (active ingredient). After introduction 
of the toxicant, no mortalities were recorded during the first 4 days (the 
24-, 48-, 72-, and 96-hour tests), nor was any distress noted in any of 
the test fish during this time. In view of subsequent results, the signifi- 
cance of this lack of mortality during the 96-hour period will be discussed 
in some detail later. 

On the fifth day, one bluegill showed the first signs of agitation 
and symptoms of distress became more intense. Before the end of the 
120-hour test, this fish had become the first mortality in Pond 2. The 
following day another fish died (144-hour test) and sporadic mortalities 
continued until the last fish expired 3 weeks after introduction of the 
toxicant (504-hour test). These events, although instructive, were some- 
what disconcerting since this level (projected from aquaria and Crosley 
Pond tests) was intended to be a sublethal concentration. It seemed un- 
tenable to suspect disease, parasitism, debility from feeding inter- 
ference, etc. as the causative agent of the mortalities. Observations 
showed no external parasites or disease and body forms of the dead 
fish displayed no noticeable emaciation. In addition, during this period, 
the control fish in an adjacent pond had suffered no mortalities under 
very similar, if not identical, conditions — with the exception of the test 
toxicant, of course. I was forced to the conclusion that the LC 100 concen- 
tration in these Driftwood waters was somewhat lower than it had been 
in the Crosley pond waters. The major differences in the ponds were 
size, shading, temperature, and transparency. The Driftwood ponds were 
larger, unshaded, warmer, and had greater clarity. All of these physical 
factors have been suspect in influencing the toxicity of various chlori- 
nated hydrocarbons in solution. These will be discussed later at some 
length. 

Pond 4 

Pond 4 was a 1.23-acre pond used for testing the residual effect 
of an extremely high dosage. This pond was sprayed with a quantity of 
endrin calulated to give a concentration of 46 ppb. The same procedure 
(two cages with 10 bluegills each) that had been used in Ponds 1 and 2 
was used in Pond 4. 

At the termination of the 24-hour test (8:00 AM, September 18, 
1961) all fish were dead. Until the pond froze over, test fish (usually 
at 2-week intervals) were placed in the cages, and each time died within 
the 24-hour period. After ice cover sealed the pond, no tests were made 
until a warmer period opened it in January. At this time, test fish again 
died within a 24-hour period. It was decided to replace the water and 
see if the residual endrin in the sediments would produce a toxicity in 
previously-uncontaminated water. 

The pond was drained February 1-3, 1962, and allowed to stand idle 
for about 3 weeks. Then it was refilled with uncontaminated water 



Ecology 155 

(March 1) and allowed to stand until March 13. On this date, a new 
group of 10 bluegills was placed in a single cage in the pond. Two of 
these subsequently escaped from encagement and one was later found 
dead along the pond edge. Of the eight fish remaining in the test cage, 
the first mortalities occurred on March 26. At this time, four fish were 
found dead. One more bluegill was dead on March 28, another on March 
29, and the final two expired on April 1. 

The pond was then drained a second time, and refilled on April 13. 
Nine test fish were put in the cage on April 25. The first mortality was 
recorded on May 2 and by May 9 all remaining test fish were dead. The 
pond was immediately drained again and had been refilled on May 15. 
Ten new fish were placed in the pond on May 20, and on June 2 the 
first death was recorded. Another died on June 5, and two more on 
June 7. Although five fish were still alive, it was obvious that a long- 
term lethal toxicity still existed. 

Once more the pond was drained and refilled, and was ready for 
the continuation of testing by June 15. Fish again were put in the cage, 
and these fish showed no mortality in a subsequent month of observa- 
tions. Tests were suspended at this time since the pond appeared to 
have become non-toxic and was needed for another study. Apparently, 
the dissipation of the toxicant was finally complete since the other fish 
stocked in this test pond were not affected. 

It should be stressed that the water source for these ponds was 
piped by gravity flow from adjacent Starve Hollow Lake which is lo- 
cated upstream from the station. Ponds drained into a small cresk 
which also drained the lake. The source water was entirely uncontami- 
nated by endrin and no other insecticides were known to have been used 
in the lake itself. In addition, no fish kills were noted in the lake. Thus 
it appears that the toxicity was of an autochthonous nature, and is as- 
sumed to have been recycled or brought into re-solution from the con- 
taminated sediments. 



Discussion 

Only in 1962 were the endrin tests given precedence over other 
duties. By 1962 it was becoming apparent from personal experiences and 
those of other investigators that this chemical was too dangerous for 
use as a piscicide except in extremely isolated instances. 

As data accumulated from various studies of endrin, certain prob- 
lems became apparent. It has been noted that other chlorinated hydro- 
carbons were present in fish flesh at higher levels than were present in 
solution (6, 7, 8, 18, 20). This accumulative effect is indicated for 
endrin also in Bridges' (3) study of a Colorado fish kill, and by implica- 
tion (22) in aquaria tests. The concentrating potential of lower forms 
in the food chain is well documented for hydrocarbons: moss (15); 
Potamogeton (8); vegetation (3, 4); earthworms (2); and plankton, 
frogs and fish (18). It is generally assumed that these accumulations in 



156 Indiana Academy of Science 

fish flesh are dietary translocations of residues through ingestion of 
contaminated food items. The role of direct absorption of the materials 
from the aquatic medium through integumentary tissues has not been 
as convincingly demonstrated, although there are data (22) that imply 
direct absorption of endrin. Hoss (17) has effectively shown that both 
pathways (absorption and ingestion) may be important in his studies 
of zinc65 accumulations in the flounder, and Williams and Pickering (28) 
have shown that bluegills accumulate cesium 1 -^ and strontium^ by both 
ingestion and absorption. Though not conclusive, these data are insinua- 
tive. 

Regardless of whether the accumulative deposition in fish flesh re- 
sults from ingestion or from absorption, it forces us to re-evaluate our 
reliance on short term bioassays (24- to 96-hour tests) where chronic 
exposures to chlorinated hydrocarbons are involved. One of the two 
major points arising from this present study is that short term bio- 
assays are insecure evidence of non-toxic levels in situations of chronic 
or long-persisting contamination and/or with materials accumulatively 
stored by metabolic processes. The need for studies of the effects of 
long term exposure has been pointed out (14), but there is a rather 
dramatic example in the study of Driftwood Pond 2. Had testing in 
this pond been suspended after a common 48-hour, 72-hour, or even 
96-hour period, the assumption of a sublethal level would have ap- 
peared valid. However, the first mortality occurred on the 5th day and 
all test fish had expired within 3 weeks from time of exposure. There 
were no mortalities among the control fish during this period. There- 
fore, one must carefully examine over extended periods those agents 
of an accumulative or an insidious nature to ascertain that seeming sub- 
lethality is not actually slow response. This is particularly true as 
testing approaches the toxicity threshold. 

The second point of interest deduced from this and other studies 
involves the variance in toxic limits for fish established by several 
workers (Table 1). Katz and Chadwick (22) obtained a 96-hour LC 50 
(their TL m ) of 0.27 ppb for coho salmon, and 0.60 ppb for bluegills. 
Henderson et al. (14) established a 96-hour LC 5 o (their TLm) of 0.44 ppb 
for bluegills in hard water using the emulsible concentrate. In the present 
study, I failed to get any kill on test fish in the Crosley test at an 
assumed 2 ppb. On the other hand, there was a complete kill of test fish 
over a 3-week period in one Driftwood pond at a 1 ppb concentration. A 
resume of Michigan lake and stream rehabilitations (16) notes a sur- 
vival of some fish (Fundulus) at 8 ppb. Bridges (3) points to a partial 
survival of largemouth bass, bluegills, pumpkinseed sunfish, and black 
crappies in a Colorado pond adjacent to a beet field which had been 
sprayed with an emulsible formulation of endrin. At the time of investi- 
gation (4 days after the spraying), water analyses indicated a concen- 
tration of 40 ppb of endrin. Fish were not dying at this time. Bridges, 
noting the disparity of his results with those of the USPHS team, points 
a suspicious finger at the 9.1 pH of the Colorado pond and the fact that 
the highest pH tested by Henderson's group (14) was 8.2. 



Ecology 



157 



Temperature has been shown to exert a tremendous effect on the 
toxicity of endrin. The studies of a Japanese group (19) observed that 
endrin toxicity to fish increased at higher temperatures, and Katz and 
Chadwick (22) found 96-hour LC 50 levels for bluegills of 8.25 ppb at 
1.0°-4.5°C and 0.33 ppb at 25 °C. The data of the latter study indicate a 
25-fold increase in toxicity with an increase of 20.5°-24.0°C. 

There are wide differences between the toxic thresholds of endrin, 
DDT and dieldrin in diverse animal groups (1, 5, 9, 10, 13, 14). An 
instance of this is obvious in Table 1. Species susceptibility to endrin has 
been pointed out by numerous investigators (14, 21, 22, 25, 27). Even 
within a particular species there is evidence that susceptibility to endrin 
changes with age, with embryonic stages being more resistant than larval 
stages (19). 



Table 1. The varying toxicity levels of three chlorinated hydrocarbons in their 

effect on several classes of organisms (measured in mg/kg of body weight or 

in ppb of active ingredient) . 



Organism 


DDT 




Dieldi 


in 


Endrin 


Estimated fatal dose for a 


30 g (5, 13) 


( = an 


5 g (13) ( = 


an esti- 


? 


man weighing 70 kg 


estimated 428 


mg/kg) 


mated 71 mg 


/kg) 




Pheasants (LD50) 


300 mg/kg 




50 mg/kg 




14 mg/kg 




(9, 10) 




9, 10) 




(9, 10) 


Bluegills (LC50 for 


8.8 ppb (14) 




9.1 ppb (14) 




0.44 ppb 


emulsible concentrate) 










(14) 


Daphnia (50-hour 


1.4 ppb (1) 




330 ppb (1) 




352 ppb 


Immobilization test) 










(1) 



It has been pointed out that the tolerance of some species is different 
in different volumes of water even though the concentration of the active 
ingredient is the same. This so-called volume effect has been noted for 
DDT (24) and for endrin (22). This increase in toxicity in greater vol- 
umes of the same concentration as smaller volumes can be logically cor- 
related with the previously discussed accumulative nature of this hydro- 
carbon. In addition to fish as accumulators of pesticidal hydrocarbons, 
studies have indicated the importance of macrophytes as concentrators 
of these chemically related insecticides (4, 8). Bridges and his co-workers 
(4), by use of isotope-labeled DDT (Cu-DDT), have found that storage 
of this relative of endrin is about Vz external (i.e., adsorption to external 
portions of the plant) and % internal (absorption into the tissues). This 
could be presumably true for endrin also. If such a presumption has any 
basis in actuality, it is quite conceivable that in aquatic ecosystems, 
where macrophytes form an important part of the biocoenosis, with- 
drawal of hydrocarbons by these plants would have a depressing effect on 
the toxicity. The result would be a balancing or a suppression of the 
volume effect, if one assumes the plants' retention of endrin until 



158 Indiana Academy of Science 

degradation of the chemical. The volume effect has been reported for 
aquaria studies, and seldom, if ever, in natural waters. Similarly, lab- 
oratory studies have generally established lower toxicity levels than field 
studies, sometimes with great disparities. My experience with 1 ppb in 
Driftwood Pond 2 is a notable exception, but as previously pointed out 
this would have been considered an lc„ at the 96-hour level. 

Fish bioassayists have noted different toxicity levels produced by 
various formulations of the same chemical compound. These are most 
graphically depicted by Henderson, et al. (14). They compared 96-hour 
LCsos for bluegills when tested in 2 formulations (acetone solution and 
emulsible concentrate) of a number of insecticides. DDT was considerably 
more toxic in emulsible concentrate, dieldrin was less toxic in this form, 
and endrin showed little difference. From other experiments, they con- 
clude that the toxicity of wettable powders (used in the present study) 
was similar to that of the acetone solution. Other chemical and physical 
factors such as the effects of light, dissolved mineral content, suspended 
particulate matter, etc., are not well-understood and could have an 
important or contributing influence. 

In spite of all these possible contributory influences, the distribution 
of variously-established toxic thresholds seems implausibly disparate. 
From these doubts is born a second — or at least, an accessory — explana- 
tion. This is simply that a lack of standardization of analytical techniques 
exists. Extraction and analysis of almost infinitesimal hydrocarbon resi- 
dues in tissues has been a tedious, painstaking job with interfering sub- 
stances having unknown effects on the results. It appears that the more 
exacting gas chromatography-infra-red spectrophotometric method now 
in use will narrow the divergence of results. However, even this advance 
did not produce incontrovertible evidence in the lower Mississippi River 
incidents. 

I suspect that higher temperatures and water transparency were 
responsible for the total kill in Driftwood Pond 2 at a level which, in 
laboratory tests, was sublethal. Conversely, the failure to get a kill in 
the Crosley Pond at a high dosage (2 ppb) is attributed to the high 
turbidity in this pond, with a possible assist from the decreasing water 
temperatures. 

Different investigators have used various designations for their 
toxicity measurements (ld, tl, and lc). Lethal dosage (ld) has been 
used for some time as the index of relative toxicity to terrestrial or 
avian animals, and the method generally has used milligrams of a sub- 
stance per kilogram of body weight as its comparative measurement. The 
Subcommittee on Toxicity, of the Federation of Sewage and Industrial 
Wastes Associations (12) recommended the term Median Tolerance Limit 
(tl,,,) to express the concentration at which just 50% of the test animals 
die. This designation was aimed at measurements of chemicals in a 
liquid medium with fish as the test animals. It is usually derived by a 
straight line graphical interpolation from tests producing higher and 
lower percentages of mortality. More recently, several investigators have 



Ecology 159 

shown an increasing preference for Lethal Concentration (lc). Usually LD 
and lc tests, like tl„,, have attempted to establish the 50% level (ld 51 „ 

lc 5 o). 

For a piscicide, as briefly mentioned previously, the minimum con- 
centration which produces 100% mortality of the fish population (LC100) 
constitutes a more important figure than the LCr.o. Desirably, for field 
use as a fish toxicant, a material should be effective in eradicating the 
population within a specific, preferably short, time (LC 100 during x time 
interval). It is obvious then that the LC100 is not a precise unity but 
rather a fluctuating value between time parameters (e.g., the 24-hour 
LC100 may be 4 ppb whereas the LC100 for a 96-hour test may be 2 ppb). 

Lennon and Walker (23) have used the term Effective Concentra- 
tion (EC100) in their delineative screening program. If we define an effec- 
tive concentration (EC) as the minimum concentration that produces a 
100% mortality to problem species, this can be written without the mor- 
tality-percentage subscript numeral. This would allow contraction of the 
abbreviate symbols by inclusion of the time interval as a superscript 
numeral as has Dorris et al. (11) in refinery waste bioassays (tl„, 48 = 
tl,„ for 48-hour test). This would avoid complicating the formula 
with both a subscript and a superscript (LC100 48 ), and it would be simpler 
to use. While EC (or LC100) is a varying quantity with time-influenced 
parameters, the EC 34 , the EC 4S , etc., are precise figures for particular test 
situations. 



Recommendations 

Among a number of desirable characteristics of a fish toxicant (low 
cost, easy application, homogeneous dispersal, relatively persistent tox- 
icity, etc.), the mandatory nature of one trait makes it the primary con- 
sideration. This is its effect on human health. It must first meet regula- 
tory standards set up to guard the public (against itself?). At the time 
these tests were initiated, endrin was merely the latest and most potent 
of an exponentially-increasing number of chlorinated hydrocarbon in- 
secticides. Its entry into the commercial market was accompanied by the 
usual — at that time — incomplete testing of its biological field effects. 
(Erstwhile tests by chemical manufacturers might be paraphrased as the 
development of broad spectrum pesticides with narrow spectrum 
evaluations.) 

It soon became apparent that endrin (for fish, at least) was the 
most lethiferous substance to come out of the chemists' cauldron. When 
certain doubts concerning its effect on human health arose, its status 
became immediately suspect, in spite of its use in harvesting food fish 
in Malaysia (26). All the ramifications of its use are still not under- 
stood, and it appears possible that authorities may belatedly prohibit its 
sale. In view of its possible health hazards and the probability of its 
imminent withdrawal from the commercial market, it seems that health 
considerations and unavailability have negated whatever potential endrin 
may have had as a fish toxicant. One cannot with any integrity abhor 



160 Indiana Academy of Science 

the fallout from atomic testings, deplore the dispersal of heptachlor to 
kill some insignificant bug, and then put on the Hydean mask to espouse 
the widespread dissimination of endrin in our waters. 



Literature Cited 

1. Anderson, Bertil G. 1960. The toxicity of organic insecticides to Daphnia. Robert A. 
Taft Sanitary Eng. Center, U. S. Public Health Serv. Tech. Rep. W60-3. p. 94-95. 

2. Barker, Roy J. 1958. Notes on some ecological effects of DDT sprayed on elms. 
J. Wildl. Mgt. 22 :269-274. 

3. Bridges, W. R. 1961. Disappearance of endrin from fish and other materials of a 
pond environment. Trans. Amer. Fish. Soc. 90 :332-335. 

4. Bridges, W. R., B. J. Kallman, and A. K. Andrews. 1963. Persistence of DDT and 
its metabolites in a farm pond. Trans. Amer. Fish. Soc. 92:421-427. 

5. Brown, A. W. A. 1951. Insect control by chemicals. John Wiley and Sons, Inc., 
New York. 817 p. 

6. Burdick, G. E., E. J. Harris, H. J. Dean, I. M. Walker, Jack Skea, and David 
Colby. 1964. The accumulation of DDT in lake trout and the effect on reproduction. 
Trans. Amer. Fish. Soc. 93:127-136. 

7. Cope, Oliver B. 1960. The retention of DDT by trout and whitefish. Robert A. Taft 
Sanitary Eng. Center, U. S. Public Health Serv. Tech. Rept. W60-3. p. 72-75. 

8. Cope, Oliver B. 1961. Effects of DDT spraying for spruce budworm on fish in the 
Yellowstone River system. Trans. Amer. Fish. Soc. 90 :239-251. 

9. DeWitt, J. B., C. M. Menzie, V. A. Adormaitis, and W. L. Reichel. 1960. Pesticidal 
residues in animal tissues. Trans. 25th N. Amer. Wildl. and Natur. Resources Conf. 
p. 277-285. 

10. DeWitt, J. B., and J. L. George. 1960. Bureau of Sport Fisheries and Wildlife 
Pesticide-Wildlife Review: 1959. Fish and Wildlife Circular No. 84, Revised. U. S. 
Dept. Interior, Washington, D. C. 

11. Dorris, Troy C, William Gould, and Charles R. Jenkins. 1960. Toxicity bioassay 
of oil refinery effluents in Oklahoma. In Biological Problems in Water Pollution. 
Trans, of the 1959 Seminar Eng. Center, Cincinnati, Ohio. p. 276-285. 

12. Doudoroff, P., B. G. Anderson, G. E. Buddick, P. S. Galtsoff, W. B. Hart, 
R. Patrick, E. R. Strong, E. W. Surber, and W. M. Van Horn. 1951. Bioassay 
methods for the evaluation of acute toxicity of industrial wastes to fish. Sewage and 
Ind. Wastes 23:1380-1397. 

13. DuBois, K. P. 1958. Insecticides, rodenticides, herbicides, household hazards. Post- 
grad. Med. 24 :278. 

14. Henderson, C, Q. H. Pickering, and C. M. Tarzwell. 1959. Relative toxicity of ten 
chlorinated hydrocarbon insecticides to four species of fish. Trans. Amer. Fish. Soc. 
88 :23-32. 

15. Hoffman, C. H., and A. T. Drooz. 1953. Effects of a C-47 airplane application of 
DDT on fish-pond organisms in two Pennsylvania watersheds. Amer. Midland Natur. 
50:172. 

16. Hooper, Frank F., John E. Williams, Mercer Patriarche, Fred Kent, and James 
C. Schneider. 1964. Status of lake and stream rehabilitation in the United States and 
Canada with recommendations for Michigan waters. Mich. Inst, for Fish. Res. Rep. 
1688. 56 p. 



Ecology 161 

17. Hoss, Donald E. 1964. Accumulation of zinc-65 by flounder of the genus Paralichthys. 
Trans. Amer. Fish. Soc. 93 :364-368. 

18. Hunt, Eldridge G., and Arthur I. Bischoff. 1960. Inimical effects on wildlife of 
periodic DDD applications to Clear Lake. Calif. Fish and Game 46 :91-106. 

19. Iyotomi, Kisabu, Tomotsu Tamura, Yasno Itazawa, Isao Hanyu, and Syotoro 
Sugiura. 1958. Toxicity of Endrin to Fish. Prog. Fish Cult. 20:155-162. 

20. Kallman, B. J., O. B. Cope, and R. J. Navarre. 1962. Distribution and detoxication 
of toxaphene in Clayton Lake, New Mexico. Trans. Amer. Fish. Soc. 91:14-22. 

21. Katz, Max. 1961. Acute toxicity of some organic insecticides to three species of 
salmoids and to the three-spine stickleback. Trans. Amer. Fish. Soc. 90 :264-268. 

22. Katz, Max, and George G. Chadwick. 1961. Toxicity of endrin to some Pacific 
Northwest fishes. Trans. Amer. Fish. Soc. 90 :394-397. 

23. Lennon, Robert E., and Charles R. Walker. 1964. Investigations in fish control : 
(1) Laboratories and methods for screening fish-control chemicals. U. S. Dept. of the 
Interior, Bureau of Sport Fisheries and Wildlife, Circular 185. 15 p. 

24. Prevost, G., C. Lannouette, and F. Grenier. 1948. Effect of volume on the determi- 
nation of DDT or rotonone toxicity on fish. J. Wildl. Mgt. 12 :241-250. 

25. Rudd, R. L., and R. E. Genelly. 1956. Pesticides: their use and toxicity in relation 
to wildlife. Calif. Dept. Fish and Game, Game Bull. No. 7. 209 p. 

26. Soong, MlN Kong. 1960. Shell "Endrex" used as a fish toxicant. Prog. Fish-Cult. 
22:93. 

27. Screenivasan, A., and M. V. Natarajan. 1962. Use of endrin in fishery manage- 
ment. Prog. Fish-Cult. 24:181. 

28. Williams, Louis G., and Quentin Pickering. 1961. Direct and food-chain uptake of 
cesium 137 and strontium* in bluegill fingerlings. Ecology 42 :205-206. 



Courtship and Territorial Behavior of Some Indiana Woodcocks 1 

Harmon P. Weeks, Jr., Purdue University 

Abstract 

Woodcocks were observed during evening flight performances from their beginning 
on March 15 until conclusion on May 24 at Shidler Forest in western Tippecanoe County. 
Aspects of flight performances were noted, timed when possible, and recorded. Territorial 
observations were recorded. Three birds were mist-netted for banding and identification 
purposes. Taped woodcock calls were used to attract males for netting and for testing 
territorial extent. 

The greatest number of performing males was observed during the last half of 
March. This was probably the peak migration period. A few birds remained on the study 
area throughout the breeding season. Light intensity controls the onset and cessation 
of performance events. Low temperatures and precipitation had little observed effect on 
performances. Longer flight times and greater ground area coverage by flights were 
noted in this study than in previous studies. These two parameters are interrelated. 

With the aid of the taped woodcock call, a woodcock was found to defend an area 
of at least 4.38 acres. Most territorial defense consisted of threats and retreats with no 
physical contact observed between combatants. When attracted by taped calls, one bird 
did attack the author and struck him several times. One singing site appeared most 
attractive because of the sequential appearance there of three males coinciding exactly 
with disappearances of males from other sites. 



Introduction 

The American woodcock (Philohela minor) is an uncommon summer 
resident in Indiana, but because they commonly nest farther to the north 
and winter to the south, large numbers may be seen here during fall and 
spring migrations (10). By the time woodcocks reach this area in the 
spring, they have already begun their courtship and this performance 
may be seen almost anywhere in the state if one searches the correct 
habitats. 

The courtship performance consists of a rather spectacular spiraling 
flight by the male and a harsh, nasal call, generally described as a 
"peent" (8), which is given from the ground between flights. A true vocal 
song is given in the flight during the first part of a rather direct descent. 
The performing area is defended against other males by the territorial 
bird, and females move into this area to be mated. 

Although this behavior has been described by many observers (1, 8, 
9), the territorial and courtship behavior of the woodcock are poorly 
described and little work has been done on this species in Indiana. The 
objectives of this study were to attempt to answer some of these 
questions and to establish some performance parameters for Indiana 
woodcocks. 

The study area was a 200-acre tract known as Shidler Forest, owned 
by the Department of Forestry and Conservation, Purdue University, and 
located 10 miles west of Lafayette, Indiana, in Tippecanoe County. It 



Journal Paper No. 3876 from Purdue University Agricultural Experiment Station. 

162 



Ecology 



163 



consists of bottomlands along Indian Creek, ridges, and ridgetops. There 
are openings in the bottomlands and on the ridgetops which are 
used as woodcock singing fields. The area has a small resident woodcock 

population. 

Methods and Materials 

Beginning in early March, the open fields of the study area were 
checked periodically for performing birds. When birds began to perform, 




LEGEND 
Field 

Woodland 
Stream 
Census Route 

500 



1000 



Scale (feet) 

Figure 1. A major portion of Shidler Forest showing performing areas which were 
occupied by singing woodcocks at times during 1969. 



164 Indiana Academy of Science 

a route was established to include visits to all occupied and other likely 
singing sites on the area. These sites were numbered for identification, 
incorporating woodcock territorial numbers of previous studies. This 
accounts for the non-sequential numbering (Fig. 1). 

This route was walked several evenings a week, and a record was 
kept of sites being used by singing woodcocks. No morning censuses were 
made. When a site was in use on two consecutive observations, it was 
assumed to have been used during the period between those visits. 
Records were kept of numbers of birds on and off territories, behavioral 
observations, and timing of performance events. 

On some days all field time was spent observing complete perform- 
ances of individual territorial males. The chronological order and duration 
of performance events, the number of flights and peents, and behavioral 
observations were recorded. 

In an attempt to identify individual birds performing on an area, 
three males were mist-netted and banded with U.S. Fish and Wildlife 
Service bands. In the first netting attempts the net was simply set in the 
usual path of flight of the bird. Later a portable cassette tape recorder 
was used to play a tape recording of the peenting of a territorial 
woodcock to lure the territorial bird into the net. 

The size of the territory (defended area) of resident birds was 
tested with the taped peenting call. This tape was played at various 
distances from the landing site of a territorial male and a positive 
response was considered to be an overflight with the issuance of a 
cackling threat note. The tape was played at the same volume at which it 
was recorded. This technique also allowed the observation of various 
reactions of territorial birds to intruding males. 

Weather conditions were recorded each day on the area. Temperature 
was measured with a mercury thermometer. Wind speed was estimated. 
In the consideration of light intensity effects, only completely clear or 
completely cloudy days were used. 

Data were tested for significance when necessary with Students'-t 
test. Differences were considered significant at the 95% confidence level 
and highly significant at the 99% confidence level. 

Results and Discussion 

Occupancy of Singing Grounds 

The first woodcock was observed performing on Area 1 on March 15, 
1969. The number of birds performing and the total number of birds 
observed on the area increased from that date until about March 25 and 
then decreased, more or less stabilizing during the first week in April 
(Fig. 2). Individual singing fields were occupied for periods of from 1 day 
to 2 months. The last half of March would seem to be the time of the main 
migratory flight in this area. Migratory males obviously occupied per- 
forming areas during short pauses in their northward movement. 



Ecology 



1(55 



15 
14 
13 
12 
Number 11 



of 



10 



Woodcocks | 

6 

5 
4 
3 
2 

1 



^- Total birds performing 
|- Total birds observed 



J 




Milan 



15 16 17 18 21 24 25 26 28 29 3 6 8 10 14 16 17 20 22 23 25 26 29 4 7 9 14 21 
March April May 

Figure 2. Number of performing woodcocks and total woodcocks recorded during cen- 
suses at Shidler Forest, 1969. 

The requirements for a singing field are fairly rigid, for even 
transient birds performing along their migratory routes prefer certain 
areas (11). A few conditions seemed to be necessary on the study area. 
A site was always an area with no tall herbaceous vegetation, either an 
area of bare ground, of matted vegetation, or of short grasses (mowed 
or otherwise checked), and small rather widely scattered woody plants. 
Acceptable heights of woody vegetation ranged from 2-15 feet. 

Temperatures did not seem to affect the performance level even 
when coupled with precipitation; birds performed at normal levels on 
March 29 when the temperature was at 27 °F and on March 25 when the 



8 

7 
Number 

of Birds 5 
4 

". 3 
2 

1 



Performing 



H - Temperature 

P~ No. of Birds Performing 



60 



50 


°F 


at 


Time 




of 


Census 


40 








30 








20 









17 18 21 24 25 26 28 29 

Clear Clear Clear Rain Snow Snow Cdy. Clear 

Date (March) 

Weather 
Figure 3. Comparison of temperature and the number of woodcocks performing in 
late March, 1969, at Shidler Forest (one inch of snow on ground, drifts to four inches). 



166 Indiana Academy of Science 

temperature was at 31 °F and a light snow was falling (Fig. 3). On 
March 26, however, with the temperature at 28 °F, a light snow falling 
and 1 inch of snow on the ground, only a few very irregular flights 
occurred (Fig. 3). Although other studies showed that performance levels 
decreased as temperatures reached 35° to 41°F (2, 5, 7), no decrease was 
evident in this study at these temperatures. Snow cover was apparently 
the influencing factor. Mendall and Aldous (7) observed that birds hesi- 
tated to land on snow and Pettingill (8) reported that a snow cover cur- 
tailed performances and that those that did occur were irregular. 

Fairly strong winds also failed to alter performance levels. On March 
28, a 10-20 miles per hour (mph) wind, with gusts to 30 mph, blew without 
inhibiting performances (Fig. 2). Light rain had no depressing effect, but 
during periods of heavy rain no flights were made, although the birds 
continued peenting from the ground. Song flights resumed when intensity 
of the rain decreased. Sheldon (11) found that high winds with or with- 
out rain and heavy downpours without wind curtailed breeding activity. 
It seems that the drive to perform courtship flights is so great that only 
the most extreme environmental conditions would cause elimination of a 
performance. 

Display Behavior 

Territorial birds usually began peenting from their diurnal areas 
5-15 min before moving onto their singing grounds. Some individuals 
gave no preperformance peenting. Mendall and Aldous (7) found it 
unusual for distances from the diurnal area to the singing ground to 
be greater than 100 yards. Although the same was found in this study, 
one bird had its diurnal area in the bottomland, about 0.4 miles from 
its singing ground on the ridgetop, Area 4-5. This bird, however, after 
11 days shifted to a performing territory within 75 yards of its diurnal 
area, when that territory, Area 1, was abandoned by its performing bird 
(Fig. 1). 

On clear days woodcocks arrived on their singing grounds an 
average of 16 min after sunset and began the first flight an average of 
21 min after sunset. Cloudy conditions caused the birds to occasionally 
move onto the areas well before sunset and greatly increased variability 
in time of first peent and of first flight relative to sunset (Table 1). 
Light intensity is the major influencing factor on time of first on-area 
peent and first flight. The greater variability in time of first peent than 
in time of first flight on clear days is undoubtedly because of the 
variable light intensities on the individual diurnal areas (Table 1). All of 
singing grounds have very similar light conditions, for none have over- 
head cover. This first flight begins when the light intensity reaches about 
2 foot-candles (4, 7). The greater variability evident on cloudy days in 
the times between sunset and the first peent and flight comes from the 
fact that even a complete cloud cover can let through extremely variable 
amounts of light (Table 1). 

A total of 89 flights in 9 complete performances were timed. There 
was much variation within and among performances. The average flight 



Ecology 167 

Table 1. Time after official sunset of first peent from performing area 
and of first flight of male woodcocks at Shidler Forest, 1969. 





First Peent 


First Flight 


Measurement 


Clear Sky 


Cloudy Sky 


Clear Sky 


Cloudy Sky 


Mean Length (min) 
Range (min) 
Standard Deviation 
No. of Observations 


16.29 
7 to 24 
4.37 
24 


—0.08 
— 17 to 20 
12,03 
13 


21.18 

16 to 27 
2.28 
22 


8.00 
—13 to 22 

12.44 
11 



length was 61.5 sec with a range of 52-76 sec. The time intervals between 
flights were also very variable. 

Three component parts of a total of 29 flights from 3 separate 
performances were timed — the ascent (period before vocal song), the 
song, and the silent descent. By far the most variable component among 
the three was the ascent with the other two being relatively invariable. 
The mean length of time for the ascent was 45.2 sec with a standard 
deviation of 3.30; of the song 9.7 sec with a standard deviation of 0.81; 
and of the silent descent 7.0 sec with a standard deviation of 0.63. Almost 
all of the variability in the flights was due to the variation in ascent 
times. Because the song and silent period occurred during fairly direct, 
though somewhat zigzagged, groundward plunges, the altitude attained 
during such flights must have been fairly uniform. Estimates of heights 
attained vary from 200-300 feet (7, 8); Sheldon (11) measured the 
altitude of 3 flights of the same bird and found all to be 275 feet. If this 
constancy of song and silent descent length hold for woodcocks elsewhere, 
then variation evident in flight length can be attributed to variation in 
ascent times. Brewster (3) reported the length of 2 songs as 11 and 12 
sec. The average and the range of flight times in this study were longer 
than those of other studies, and the area covered in one flight was 
also larger, averaging over 2 acres. Pitelka (9) reported average flight 
coverage of only V 3 acre and flight times of 29 to 60 sec with an average 
of 43 sec (calculated by author). The majority of the flights timed by 
Mendall lasted 50 to 55 sec with a range from 44 to 63 sec (7). The 
author suggests that the time of an ascent, and thus a flight, is deter- 
mined by the time required to reach a certain altitude, and that this time 
is directly proportional to the amount of area encompassed in a flight. 
This area may in turn be determined by several factors including size of 
singing field, height of surrounding vegetation, and juxtaposition of other 
territorial males. 

The number of peents per minute varied greatly, ranging from 7 to 
28 per minute. The woodcocks occasionally walked or ran short distances 
between peents but most often remained in one spot, usually pivoting a 
little after each peent. Some birds turned 360° in one direction while 
others pivoted 90-180° in one direction and then reversed directions. This 
rotation caused the volume of the peenting to seem to vary to an 



168 Indiana Academy of Science 

observer and probably served to make territorial birds conspicuous for 
the maximum distance possible in all directions. 

The average length (from first peent on the area to the last) of 26 
performances not influenced by moonlight was 34.2 min, ranging from 
16 to 53 min. On a moonlit night, April 29, a performance lasted 74 min. 
It was found that performances were highly significantly longer on cloudy 
than clear days, 41.3 and 31.5 min, respectively. The lengths of per- 
formances were also more variable on cloudy than on clear days. Total 
performance length was greatly influenced by light conditions. Cloudiness 
extended performances because of earlier starting times and fairly com- 
parable stopping times; the greater variability in lengths of perform- 
ances on cloudy days was undoubtedly caused by differential light pene- 
tration through a full cloud cover. A long performance on a moonlit night 
as observed in the study has been reported by many investigators. It is a 
good example of the degree to which various aspects of the performance 
are light intensity dependent (7, 8, 11). 

The manner in which performances ended differed among individuals. 
A bird left an area after the last flight either immediately, after a long 
period of silence, or most commonly after several minutes of decelerating 
peenting. 

Territorial Behavior 

Territorial birds tolerated no territorial intrusions by other males. 
Cackling was the most common threat exhibited. When an intruder 
peented within an occupied territory, the territorial bird usually would 
fly immediately toward the source of the strange peenting and cackle. 
This generally discouraged the intruder which hid and became silent or 
departed. The peent was the major communication in woodcock relation- 
ships and served the dual purpose of advertising occupancy and of dis- 
closing the presence of intruders. 

Several chases were observed and in all cases both birds seemed to 
flush from nearly the identical spot. When first sighted, the birds were 
usually 10 to 15 feet above the ground with one about 5 feet ahead of the 
other. This interval was kept throughout the chase as the birds climbed 
at a 45 to 90° angle from the horizontal. Usually the only sound emitted 
during these chases was wing-twitter although cackling occasionally 
occurred. About half of these chases ended with the bird giving chase 
veering off and beginning a flight song. At other times the chase con- 
tinued upward until both birds were lost from view and several minutes 
elapsed before the territorial bird returned to the area. No physical con- 
tact between birds was ever observed. Similar chases were observed by 
Pitelka (9) who attributed them to two males simultaneously starting 
their flights. From the observations in this study, this does not seem 
likely. It is also unlikely that this is a courtship flight including male and 
female, because in none of the 11 chases observed did the bird being 
chased ever return to the field. It seems most probable that it is a very 
ritualized form of chase which has developed evolutionary to maximize 



Ecology 169 

survival by preventing injury to combatants and still maintaining terri- 
torial integrity. 

An attempt was made to determine the area defended by a terri- 
torial bird. On May 5 and 6, as the peenting tape was played from 
various locations on the area, the territorial male on Area 1 reacted by 
flying toward it and cackling over a total area of 4.38 acres. The 
greatest distance from which a response was drawn was 330 feet. It 
had been thought previously that the birds defended at most the area 
covered by their flights, estimates of which varied from V 3 to 2 acres in 
different studies (8, 9). Because this bird reacted at all points tested, 4.38 
acres is the minimum estimate of the territorial size. If the greatest dis- 
tance from which a response was drawn, 330 feet, is assumed to be the 
territorial limit and a circle circumscribed about the landing site with 
this as the radius, the area included is 7.85 acres, a considerably larger 
territory than anyone had previously suspected. 

Several aggressive reactions of territorial birds to the presence of 
the investigator were encountered. On May 1, I was lying about 100 feet 
from the usual landing site of the bird on Area 1. The taped peenting 
was played to test the bird's reaction to it. The territorial bird cackled, 
flew toward the sound and lit within 20 feet of the recorder, which was 
on the ground about 3 feet from my head. After flying closer to the 
recorder and peenting several times, it flushed and returned to its usual 
landing site. The complete approach process was repeated twice more. 
On the third approach, the bird landed 5 feet from the recorder and 
approached it on foot. It stood silently next to it for a few seconds and 
then began searching through the grass around the recorder. Suddenly it 
moved through the weeds and appeared about a foot from my face. 
Almost immediately the woodcock lunged toward me and struck my eye 
with its bill and then grabbed a tuft of hair in its bill and yanked vigor- 
ously several times. The bird then released its hold, jumped back about a 
foot, and stood watching me. I remained as still as possible. The whole 
attack was then repeated with the bird first striking my eye and then 
yanking my hair. After this attack an attempt was made to catch the 
bird by hand, but its reactions were much too quick and it flushed. How- 
ever, it immediately responded again to the peenting tape and attacked 
my hand several times as I shuffled it in the grass. These attacks indicate 
how strong a role peenting plays in the territorial behavior of the wood- 
cock. This woodcock and others were invariably attracted by the peenting 
from the tape even seconds after unsuccessful attempts to capture them. 
After a woodcock is attracted to an area by alien peenting, any movement 
is evidently attributed to the intruder and is attacked. In both face-to-face 
encounters the woodcock first struck my eye, the only place in which 
movement occurred. It also attacked my hand only when it was shuffled 
in the grass. This would be the expected reaction sequence in a species 
which defends territory only during periods of marginal light intensity 
and against somber colored intruders. 

In an attempt to band birds so that homing could be tested in subse- 
quent years, the bird on Area 1 was mist-netted and banded on April 25. 



170 



Indiana Academy of Science 



Soon after this a different behavior pattern for the bird on this area 
was noted (i.e., changes in landing site, flight pattern, diurnal area, 
reaction to taped peenting, and various other performance parameters) 
and a male caught on May 7 proved to be a different bird. The subse- 
quent appearance soon afterward of a third behavior pattern lead to the 
capture and banding of a third male from Area 1 on May 16. The appear- 
ance of the second bird on Area 1 was closely linked with the discontinu- 
ance of the performances of the bird on Area 4-5, and the appearance of 
the third bird with the cessation of the performances of the bird on Area 
10 (Fig. 4). Woodcocks from adjacent areas undoubtedly replaced previ- 
ous territorial birds as they disappeared from Area 1. Both birds which 
were replaced on Area 1 disappeared soon after netting and banding. It 
is impossible to determine what caused this disappearance, but the 
trauma of the netting and banding operation may have caused the birds 
to move off the study area. However, the subnormal post-banding per- 
formance by bird "A" illustrated that banded birds may have ceased 
performing while remaining on the study area (Fig. 4). Because of this 
replacement pattern, Area 1 was interpreted as being the prime per- 
formance site on the study area, and thus, the one most competed for. 
The fact that it had at least three males on it with not a day of non- 
occupancy is indicative of its attractiveness. 

Sheldon (11) suggested that competition for open spaces was very 
important in the evolutionary shaping of the territorial behavior of the 



B 



March 
15 20 25 

I 



30 1 



April 
10 15 20 



May 



25 30 

H«,i— 



H ' h 



First performing bird netted J 

Subnormal performance - 

Study area not checked - 

New pattern noted - 

Bird on area 4-5 no longer performing - 

Performing bird "B" netted - 
Normal performance of "B" pattern 
New pattern noted 
Bird on area JO no longer performing 



15 20 24 



Performing bird "C" netted 

Normal performance of "C" pattern 

Subnormal "C" performance 

No flights— only peenting 

No bird on area 



FIGURE 4. Chronological sequence of events on Area 1 in woodcock study at Sh idler 
Forest, 1969. 



Ecology 171 

woodcock, for in primeval times very few openings existed in the eastern 
United States and Canada. It is probable that even now strong competi- 
tion for preferred sites exists. This competition and territorial defense 
appears to be principally a ritualistic type which involves little physical 
contact. Lack (6) said that avoidance by others of occupied areas was the 
primary factor in maintaining territorial integrity in birds. Tinbergen 
(12) said that his previous view that hostility was the primary factor as 
well as Lack's that avoidance was primary were both one-sided and that 
the two factors operate in tandem. The cofunctioning of these two factors 
may offer a partial explanation for the tenacious drive to perform which 
has evolved in the woodcock. When a bird is on territory, it is at an 
advantage since its defense is nearly always successful because of the 
natural avoidance of occupied areas by unattached males. However, if a 
bird fails to occupy its territory for a day or more due to the weather 
or some other factor and another bird occupies the area in the interim, 
the former territorial bird is then at a disadvantage. Those birds which 
establish a territory in a favorable location and perform there daily, 
regardless of conditions, will be selected for evolutionarily, for they have 
a greater chance of being on territory throughout the breeding season and 
thus more chances to procreate. 



Literature Cited 

1. Bent, A. C. 1927. Life histories of North American shorebirds, part 1. U.S. Nat. Mus. 
Bull. 142. 420 p. 

2. Blankenship, L. H. 1957. Investigations of the American woodcock in Michigan. 
Mich. Conserv. Dep., Game Div., Rep. No. 2123. 217 p. 

3. Brewster, William. 1894. Notes and song flight of the woodcock {Philohela minor). 
Auk 11:291-298. 

4. Duke, Gary E. 1966. Reliability of censuses of singing male woodcocks. J. Wildl. 
Mgt. 30 :697-707. 

5. Goudy, William H. 1960. Factors affecting woodcock spring population indexes in 
southern Michigan. Mich. Conserv. Dep., Game Div., Rep. 2281. 44 p. 

6. Lack, David. 1954. The natural regulation of animal numbers. Oxford University 
Press, London. 343 p. 

7. Mendall, Howard L., and Clarence M. Aldous. 1943. The ecology and management 
of the American woodcock. Maine Coop. Wildl. Res. Unit, Orono. 201 p. 

8. Pettingill, Olin Sewall, Jr. 1936. The American woodcock, Philohela minor 
(Gmelin). Mem. Boston Soc. Natur. Hist. 9(2) :167-391. 

9. Pitelka, Frank A. 1943. Territoriality, display, and certain ecological relations of 
the American woodcock. Wilson Bull. 55(2) :88-114. 

10. Robbins, Chandler S., Bertel Bruun, and Herbert S. Zim. 1966. Birds of North 
America. Golden Press, New York. 340 p. 

11. Sheldon, William G. 1967. The book of the American woodcock. The University of 
Mass. Press, Amherst. 227 p. 

12. Tinbergen, N. 1957. The function of territory. Bird Banding VIII il> :l-8. 



Foods of the White-footed Mouse, Peromyscus leucopus 
noveboracensis, from Pike County, Indiana 1 

Gwilym S. Jones, Purdue University 

Abstract 

During a fauna] study of a strip-mined area in Pike County, Indiana, the stomachs 
of 489 white-footed mice (Peromyscus leucopus noveboracensis) were examined to de- 
termine food preferences. The stomach contents were identified by comparison with 
seeds, arthropods, and other materials collected on the study plots where the mice were 
trapped. The average % volume of each food was estimated for each stomach. Of the 
68 different materials identified, undifferentiated starchy substance and invertebrates were 
the most common. Wild cherry (Prunus sp.), blackberry (Rubus sp.) and wood son-el 
(Oxalis sp.) were the most abundant identifiable plant foods. The average % volumes of 
food types (i.e. plant, animal, and "other") were computed for each season. The results 
showed that plant foods are dominant throughout the year, but increase noticeably during 
the fall and winter. 

Introduction 

As part of a faunal study of a strip-mined area (6), stomachs of all 
white-footed mice captured during a trapping- program were examined 
to determine food preferences of the species. This paper summarizes the 
findings of the study. 

Study Area 

The study area covers approximately IV2 square miles in Monroe 
and Lockhart Townships (R7W and R8W, T3S), Pike County, Indiana. 
About V2 of the area has been strip-mined for coal. The topography 
resulting from the strip-mining is either pyramidal, parallel ridges rang- 
ing in height to 75 feet, or extensive flat areas with intermittent small 
hills. The stripping began in 1921 and ended in 1961. The unstripped half 
of the area has rolling topography with intermittent low, wet areas. 

The vegetation was divided into nine distinct cover-types based on 
dominant species. Pines, mixed pine-hardwoods, pine-hardwood saplings, 
and black locust occurred only on the stripped portions. Hardwoods, brush 
and weeds occurred on both stripped and unstripped land, and bottom- 
land hardwoods and crops occurred only on the unstripped land. Except 
for brush and weeds, all cover types on the stripped land had been 
planted. 

Methods 

The food habits study was based on mice that were captured during 
a population study which employed 100 plots randomly selected from a 
gridded map of the area. The plots were 125 feet square and contained 
18 trapping sites of 3 traps each. Each plot was set for three nights. The 
stomachs of the captured mice were removed as soon as possible and 
preserved in 70% alcohol. During trapping all plants found on each plot 
were recorded and samples of their seeds and fruits collected. 



1 Work was supported by the Indiana Dept. of Natural Resources, Midwest Coal 
Producers, Central States Forest Experiment Station and Purdue Agricultural Experiment 
Station. 

172 



Ecology 173 

After the trapping program was completed, the stomachs were dis- 
sected and the contents washed into a dish of alcohol. The contents were 
then identified by comparison with the sample collections of seeds and 
animal matter. The volume of each food was estimated according to the 
methods of Hamilton (4) and Whitaker (9). Every effort was made to 
identify all stomach materials. However, some unidentifiable contents 
were listed as unidentified plant or animal materials, or unknown, depend- 
ing upon their nature. The data were programmed and analyzed by 
computer. Because the study emphasized plant materials, arthropod 
remains were identified only to general classifications. 

Results 

The most prevalent year-round food eaten by the white-footed mouse 
was undifferentiated, starchy material. Insects and Lepidoptera or 
Hymenoptera larvae were other preferred foods. Unidentified seeds ranked 
below starchy material and insects. Of the identifiable seeds, those with 
the highest volumes were wild cherry (Prunus serotina) , blackberry 
(Rubus sp.), and yellow wood sorrel (Oxalis sp.). All stomach items 
except those in the next paragraph are listed in Table 1. 

The following foods were found in quantities of less than 0.1% 
volume: Compositae seeds, 0.8% frequency; crowfoot (Ranunculus) 
seeds, Annelida, and touch-me-not (Impatiens) seeds, 0.6 % frequency; 
pupae cases, pokeberry (Phytolacca) seeds, tick trefoil (Desmodium) 
seeds, and metal, 0.4% frequency; trumpet creeper (Campsis) seeds, 
Coleoptera larvae, flower petals, bones, redbud (Cercis) seeds, oxeye 
daisy (Chrysanthemum) seeds, avens (Geum) seeds, sedge (Scirpus) 
seeds, daisy fleabane (Erigeron) seeds, lettuce (Lactuca) seeds, bluets 
(Houstonia) seeds, St. John's-wort (Hypericum) seeds, and plantain 
(Plantago) seeds, 0.2% frequency. 

The seeds of planted trees consumed by the white-footed mouse were 
maple (Acer), pine (Pirius) , ash (Fraxiyius), elm (Ulmus), and black 
locust (Robinia). Of the planted species, maple had the highest average 
volume and % frequency, but these values were only 1.2% volume and 
8.3% frequency. 

A consideration of the seasonal preferences indicates that plant foods 
are dominant throughout the year but increase noticeably during the 
fall and winter, after the plants have dropped their fruits and arthropods 
have become scarce. 



174 Indiana Academy of Science 

Table 1. Average percent volume and percent frequency of foods found in 489 white- 
footed mice (Peromyscus leucopus) stomachs from September, 1965, to August, 1966, in 
Pike County, Indiana. 

Food Volume Frequency 

Starchy material 

Insects 

Lepidoptera or Hymenoptera larvae 

Unidentified seeds 

Wild cherry (Prunus) seeds 

Rubus (sp. ) seeds 

Yellow wood sorrel (Oxalis) seeds 

Unidentified plant 

Hair 

Chilopoda 

Green vegetation 

Indian hemp (Apocynum) seeds 

Cranesbill {Geranium) seeds 

Mollusca 

Maple (Acer) seeds 

Unknown 

Arachnida 

Flesh 

Pine (Pinus) seeds 

Endogone 

Ash {Fraxinus) seeds 

Sumac (Rhus) seeds 

Elm iUlmus) seeds 

Bush clover (Lespedcza) seeds 

Rose (Rosa) seeds 

Rumex (sp. ) seeds 

Bittersweet (Celastrus) seeds 

Cottonwood (Populus) seeds 

Poison ivy (Rttus) seeds 

Sassafras (Sassafras) seeds 

Pebbles 

Honeysuckle (Loniccra) seeds 

Unidentified grass seeds 

Panic grass (Panicum) seeds 

Dogwood (Cornus) seeds 

Unidentified animal 

Everlasting pea (Lathyrus) seeds 

Unidentified fungus 

Grape (Vitis) seeds 

Greenbriar (Smilax) seeds 

Sedge (Car ex) seeds 

Feathers 

Peppergrass (Lcpidium) seeds 

Foxtail (Setaria) seeds 

Black locust (Robinia) seeds 

Sweet clover {Melilotus) seeds 

Apple (Pyrus) seeds 



30.1 


77.3 


14.5 


7S.fi 


9.2 


42.2 


7.5 


41.0 


5.:-! 


21.4 


3.7 


10.0 


3.6 


9.4 


•A. -A 


23.3 


2.2 


41.2 


1.9 


18.7 


1.8 


12.7 


l.fi 


7.0 


1.5 


5.0 


1.2 


8.9 


1.2 


8.3 


0.!) 


8.3 


0.8 


9.8 


0.8 


4.8 


0.8 


4.8 


0.7 


6.0 


0.7 


4.4 


<u; 


2.3 


0.5 


A.A 


0.5 


A A 


0.5 


2.0 


0.4 


3.7 


0.4 


2.0 


0.4 


2.5 


0.4 


2.5 


0.4 


1.5 


0.3 


7.:-! 


0.3 


1.5 


0.2 


3.3 


0.2 


2.7 


0.2 


2.1 


0.2 


1.5 


0.2 


0.6 


0.1 


3.5 


0.1 


1.5 


0.1 


1.2 


0.1 


1.2 


0.1 


1.0 


0.1 


1.0 


0.1 


0.6 


0.1 


o.i; 


0.1 


o.t; 


0.1 


0.2 



Ecology 175 



Discussion 



I agree with Hamilton (4) that the starchy material, called mast by 
Whitaker (9), is composed of many different materials (such as mast, 
root tips, and seeds) which cannot be separated by observation. Hamilton 
(4) found stomachs containing nothing but starchy material. Such was 
the case in this study. Undoubtedly some of the starchy material was 
composed of acorns (Quercus) , hickory nuts (Carya) , and other such 
fruits. Whitaker (personal communication) indicates that he has found 
portions of the exocarp of such fruits still clinging to the material. Such 
was not the case in this study. 

Green vegetation was separated from unidentifiable plants because 
this material is composed of leaves, buds, or other vegetative plant parts. 
This is supported by Hamilton's (4) contention that the remains of buds, 
and Jameson's (5) conclusion that cotyledons of sprouting seeds, when 
eaten by mice, retain their green color. By my methods it was impossible 
to determine the origin of the plants or the parts. The unidentified plant 
material was thought to be fragments of xylem or phloem (2). 

The fungus Endogone was a common item in the stomachs of many 
small mammals by a number of investigators (1, 2, 3, 4, 7, and 11). 
However, it was not a common food of the white-footed mouse in this 
study, ranking 20th in volume with a frequency of 0.7%. 

Although insects were a major food, they never surpassed plant food 
in volume or frequency. Similar observations were reported by Hamilton 
(4), Jameson (5), Williams (10), and Whitaker (8, 9). 

Hair, found in 41.5% of the stomachs, was often in the form of balls, 
suggesting that it resulted from the grooming habits of the mice. Some 
hair undoubtedly accompanied ingested flesh which was found in 4.8% of 
the stomachs. 

Stomach analyses admittedly provide only an indication of food 
preferences. However, they provide some information concerning the 
ecology of the mammal and indicate what plants it eats. Although some 
believe that enough food habits work has been done for Peromyscus, of 
the 68 foods identified, seeds of 27 plant species were new foods for the 
species. To the author's knowledge none has been reported before. The 
most common of these were Indian hemp (Apocynum) and cranesbill 
{Geranium) with frequencies of 7.9% and 5.0%, respectively. 



Acknowledgements 

Thanks are extended to Dr. R. E. Mumford, Purdue University, 
Dr. J. O. Whitaker, Jr., Indiana State University; and William Crawford, 
Enos Coal Mining Company. 



176 Indiana Academy of Science 

Literature Cited 

1. Bakerspigel, A. 1958. The spores of Endogone and Melanogaster in the digestive 
tracts of rodents. Mycologia 50 :440-442. 

2. Calhoun, J. B. 1941. Distribution and food habits of mammals in the vicinity of 
the Reelfoot Biological Station. J. Tenn. Acad. Sci. 6:177-225. 

3. Dowding, E. S. 1955. Endogone in Canadian rodents. Mycologia 47 :51-57. 

4. Hamilton, W. J., Jr. 1941. The food of small forest mammals in eastern United 
States. J. Mammal. 21 :250-263. 

5. Jameson, E. W., Jr. 1952. The food of deer mice, Peromyscus maniculatus and 
P. boylei, in the northern Sierra Nevada California. J. Mammal. 33 :50-60. 

6. Jones, G. S. 1967. Vertebrate ecology of a strip-mined area in southern Indiana. 
M. S. Thesis, Purdue Univ. 158 p. 

7. Whitaker, J. O., Jr. 1962. Endogone, Hymenog aster, and Melanogaster as small 
mammal foods. Amer. Midland Natur. 67:152-156. 

8. . 1963. Food of 120 Peromyscus leucopus from Ithaca, New York. J. 

Mammal. 44 :418-419. 

9. - — . 1966. Food of Mus musculus, Peromyscus maniculatus, and Peromyscus 
leucopus in Vigo County, Indiana. J. Mammal. 47:473-486. 

10. Williams, O. 1959. Food habits of the deer mouse. J. Mammal. 40:415-419. 

11. Williams, O. and B. A. Finney. 1964. Endogone — food for mice. J. Mammal. 
45:265-271. 



Observations on Ecology and Behavior of Indiana Ruffed Grouse 1 

John P. Muehrcke and Charles M. Kirkpatrick, Purdue University 



Abstract 

On an 840-acre study area in Monroe County, vegetation types consisted of second- 
growth hardwoods including a 19-year-old burn, old fields, pine plantations, and wildlife 
openings. Flush surveys showed that grouse used hardwoods most consistently (especially 
the burn), pines for winter and hunting season protection, and other types very little. 
Age ratios of summer-flushed and summer-trapped grouse were essentially even in 1968. 
Eight grouse broods found in summer of 1968 averaged 3.1 young, less than noted in 
1965. Brood hens showed protective behavior until young were 10-12 weeks old. Extent 
of brood separation movements varies among individual young. Fifteen drumming males 
were found in 1969 compared with 12 in 1968. In 1969, drumming began February 28, 
the number of drumming males increasing until April 10 when all 15 males drummed, 
and decreasing afterward until only 2 were heard April 23. The daily period from 
about % hour before sunrise until y 2 hour after sunrise marked the most intense drum- 
ming activity. The intervals between drumming performances were shortest just before 
and just after sunrise. The drumming act averaged 5+ seconds. Males occasionally roosted 
on drumming logs, but rarely drummed at night. Besides their main drumming logs, all 
males had one or more alternate logs. 

A century ago ruffed grouse (Bonasa umbellus) were probably found 
throughout Indiana woodlands, being reported in 58 counties (13), but 
the effect of man has limited this species to the less intensively used 
hills of south-central Indiana. Evidence mounts that the bird is holding 
its own if not actually thriving in the managed public forest lands. 
Thurman (25) started a study on ruffed grouse ecology in Monroe County 
to which a report by Muehrcke (19) represents a continuation. The work 
of these students, supervised by the junior author, proposes to accumulate 
data for the better understanding of Indiana ruffed grouse ecology to 
aid in management practices that will benefit this fine bird. 

The study concentrated on an 840-acre part of the Hoosier na- 
tional Forest in southeastern Monroe County locally known as Geiger 
Ridge. It lies within the western mesophytic and oak-hickory climax 
vegetation (17) in which man has wrought many changes to the forest. 
Most of the wider valley bottoms and ridge tops were cleared and 
farmed and remaining timber was extensively logged. Now the most 
common species in the overstory include chestnut oak (Quercus spp.), 
black oak (Quercus velutina Lam.), red oak (Quercus spp.), white oak 
(Quercus alba L.) , and hickory (Carya spp.). Species nomenclature fol- 
lows Deam (5). In 1935 the U. S. Forest Service began buying the 
worked-out farms, allowing some cleared fields to revert to forest suc- 
cessional stages and planting some sites with pines. This history of 
land use has resulted in some distinct vegetation types consisting of 
second-growth hardwoods including a 19-year-old burn, old fields, pine 



1 Journal Paper No. 3856 from Purdue University Agricultural Experiment Station, 
based on a master's thesis by John P. Muehrcke with research support from the Indiana 
Division of Fish and Game and cooperation from U.S. Forest Service personnel, Hoosier 
National Forest. 

177 



178 Indiana Academy of Science 

plantations, and wildlife openings. Thurman (25) describes these and 
other physical and historical features of the area in detail. 

Methods 

To locate grouse broods, the observer made intensive searches in 
all cover types in early summer. Brood locations later became focal 
points for trapping with cloverleaf traps (4). Trapping was done pri- 
marily to gather information about age ratios, grouse movements, and 
to provide a basis for population estimation. Sex and age were deter- 
mined for each trapped bird, which was then color-banded and released. 

Throughout the study, major effort was devoted to observing be- 
havior of grouse in their natural environment. Hens with broods and 
drumming males provided most such contacts. Whenever a brood was 
discovered, usually by flushing, the observer concealed himself to watch 
the behavior of hen and young as they regrouped from hiding. Ten 1/10- 
acre vegetation samples were taken to describe the plant communities 
where broods were flushed. Muehrcke (19) used a line transect method 
as the sampling technique. 

The observer located all drumming males on the area by walking 
along ridge tops from x k hour before sunrise to Vz hour after sunrise. 
Each ridge top was covered at least twice during the clear calm days of 
March and April. The sound of drumming revealed drumming log loca- 
tions and offered chances to study the performing males. Certain drum- 
ming males were watched intensively for recording behavior. Drumming 
logs of all cocks were periodically examined for signs of use as a clue 
to drumming male population. Cover analyses were made of 10 drumming 
log sites by the method of Lindsey et al. (16). 

Results and Discussion 

The study area divided itself roughly into the part burned over in 
1951 and the unburned hardwoods. The burn is an interspersion of pines, 
field edges, and open canopy hardwoods with a great deal of brush, sap- 
lings, and pole timber. The unburned area is predominantly oak-hickory 
with little understory. The burned hardwoods in the southern half of 
the area obviously held more grouse than the unburned portion. Even 
though grouse were not uniformly distributed in the burn, most field 
work was concentrated there. In general, most grouse appeared where 
brushy vegetation was densest and along ridge tops in the burn and 
near pine plantations. Ridge tops with older unburned timber were not 
used extensively except by drumming males. 

From June 1968 to April 1969, 250 grouse were flushed in the 
study area. Since the 1968 estimated fall population based on marked 
birds was 42, obviously many birds were flushed more than once. Al- 
though grouse appeared to be most abundant in the burned section, when 
Thurman (25) more completely searched the whole area by randomized 
routes, his recorded flushes in unburned hardwoods led in all seasons 
except fall. In the present study, of all broods flushed in summer, 86% 



Ecology 179 

came from burned over hardwoods, the rest from pines. Slopes and 
valley bottoms, as opposed to ridge tops, produced 70% of brood flushes. 
Temperature and humidity data from valley bottom (670-foot contour) 
and ridge top (850-foot contour) showed that the valley had a wider 
range of temperature and was more humid than the ridge. The ap- 
parent preference of grouse for the lower levels in summer may be re- 
lated to temperature and moisture conditions more comfortable to the 
birds, especially as the same conditions associate with denser vegetation 
affording protection and food supplies. Crop contents from four summer 
juveniles were strong in occurrence of fungi and touch-me-not (hnpatiens 
spp.), both abundantly found in bottoms. 

In September, the first noticeable difference occurred in grouse loca- 
tions with birds increasingly found on ridge tops. During the period of 
most movement in the first week of October, grouse often appeared on 
roads and in areas where previously unseen. The October movement was 
most frequently away from dense hardwood thickets of slopes and bot- 
toms, the high use area of summer, to edges around pine plantations. In 
October, hunting pressure was associated with increased grouse use of 
pine plantations, the densest protective cover on the area. Thurman (25) 
also noted increased use of pines in fall and winter. In midwinter, grouse 
use in and around pines remained high, but they also spread back into 
dense hardwood vegetation, often feeding there on the abundant crop of 
sumac (Rhus spp.) fruit. Groups of grouse were more commonly seen in 
winter than in fall, indicating a tendency to concentrate at feeding sites. 

During early spring, grouse again used slopes and ridge tops, males 
moving upward to more mature timber to establish drumming territories. 
Males flushed from drumming logs showed no unusual behavior. Even 
those flushed from logs on moonlit nights flew normally. They had no 
special escape routes but usually flew in a direction away from the ob- 
server. 

In three summer months of 1968, a total of 80 flushed grouse gave 
nearly an even adult: juvenile age ratio (38 juveniles, 37 adults, 5 un- 
known). Of 26 different grouse captured from July to October inclusive, 
the age ratio was even. This is unusual as a healthy post-breeding popu- 
lation of small game birds should include 60-80% young of the year (1). 
In 1968, broods accompanying hens averaged 3.1, substantially less than 
the 6-8 found by Thurman (25) on the same area in 1965. Bump et al. (3) 
reported 6-8 young in other states. Obviously, 1968 was a poor year for 
recruitment of ruffed grouse on the study area. A few reasons of circum- 
stantial nature suggest themselves. 

Rainfall amounting to 9.4 inches occurred in 14 days in May, more 
than twice as much as a previous 100-year average for the period. Ex- 
treme wet conditions at the time when ruffed grouse are hatching may 
cause high chick mortality (7). To underline the possibility of wet 
weather mortality, 54% of summer trapped grouse were males. There- 
fore, by interpolation, 17 of 37 adult grouse flushed in summer 1968 may 
have been hens. There were actually eight known females with broods, 



180 



Indiana Academy of Science 



leaving a possible nine broodless hens. Torrential rains could easily 
destroy whole broods of young chicks, clutches near hatching, or decimate 
either to account for small brood size and broodless hens in 1968. (A 
better brood year probably occurred in 1969. In a survey for cover 
mapping on a nearby area, John D. Vanada found 10 broods during the 
period June 11-August 22. They averaged 6.9 young per brood and no 
broods had less than 5 chicks). 

On the basis of recaptured banded grouse, the 1968 fall population 
of the study area was estimated at 42±14.8 birds at the 95% confidence 
interval (18). Meager data make this only an approximation, but other 
methods of crude estimating give about the same number. 

Composition and arrangement of forest ground cover and overstory 
determine ruffed grouse distribution and influence productivity (7). Va- 
riety among food producing plants is essential to counteract irregulari- 
ties in food production because of weather or alternate seeding years. 
Because of the sedentary nature of ruffed grouse, good habitat meets 
food and cover requirements within a relatively small area (24). Sum- 
mer brood range poses the most critical requirements. Broods must have 
a complex understory of woody and herbaceous plants of proper height 



Table 1. Vegetation analysis of canopy and shrub strata for ten l/lO-acre samples at 
brood locations (1968) and ten l/25-aere samples at drumming log sites (1969), Geiger 
Ridge Study area. (Only the 10 highest ranking species for importance value are shown 

in each case.) 







Relative 












Trees 


Stem den- 


stem 


Basal 


Relative 




Relative 


Impor- 


and 


sity pei- 


density 


area 


basal area 


Fre- 


fre- 


tance 


shrubs 


acre 


per acre 


per acre 


per acre 


quency 


quency 


value 






Brood 


Locations 










Hickory spp. 


341 


9.1 


5.9 


22.6 


100 


3.5 


11.7 


Shortleaf pine 


83 


2.1 


7.S 


30.3 


20 


0.7 


11.0 


Sumac spp. 


783 


20.7 






100 


3.5 


8.6 


Black oak 


i7<; 


4.6 


4.S 


IS. 5 


SO 


2.S 


8.6 


White oak 


74 


1.9 


2.S 


L0.8 


90 


3.2 


5.3 


Sassafras 


450 


11.9 


0.08 


0.3 


100 


3.5 


5.2 


Ironwood (Ostrya) 


203 


5.3 






90 


3.2 


4.2 


Sugar maple 


101 


2.6 


1.5 


0.0 


100 


3.5 


4.0 


Dogwood 


269 


7.1 


0.08 


0.3 


100 


3.5 


3.6 


Hazelnut ( Corylus ) 


L73 


4.5 






70 


2.5 


3.5 






Drumming Log Sites 








Black oak 


272 


5.4 


20.4 


37.2 


100 


6.4 


16.3 


Sumac spp. 


592 


11.9 






100 


6.4 


9.1 


Red oak 


32 


0.5 


9.4 


17.1 


30 


1.9 


S.5 


Sassafras 


627 


12.6 


3.4 


6.2 


100 


6.4 


S.4 


Hickory supp. 


462 


9.2 


3.1 


5.S 


100 


6.4 


7.1 


Dogwood 


460 


9.2 


L.5 


2.S 


90 


5.S 


5.9 


Grape (Vitis) 


11)7 


3.9 






100 


6.4 


5.1 


Blue beech (Carpinus) 


i 345 


6.9 






50 


3.2 


5.0 


Ironwood 


215 


4.3 






90 


5.S 


5.0 


Tulip tree 
















( Liriodendron ) 


27 


0.5 


6.1 


11.1 


50 


3.2 


4.8 



Ecology 181 

and density to allow easy movement of chicks as they forage (24). The 
fact that grouse have persisted on the study area without any special 
attention indicates that the range is meeting at least the minimum 
requirements for their survival. It is likely that extensive interspersion 
of woodland successional stages on Geiger Ridge similarily provide the 
mixture of types found necessary for ruffed grouse in Missouri (14). 

The study area supported a minimum of 8 broods, usually found 
in relatively open canopy situations with dense underbrush. In a com- 
parison of basal areas of different species, hickory, black and white 
oaks, sugar maple (Acer saccharum Marsh.), and shortleaf pine (Pinus 
echinata Mill.) dominated the overstory of brood locations. In stems 
per acre, sumac, sassafras (Sassafras albidiim), hickory, and dogwood 
(Cornus spp.) led in that order of understory dominants (Table 1). 
Ground cover consisting of grasses, greenbrier (Smilax spp.) tickclover 
(Desmodium spp.), cinquefoil (Potentilla spp.), goldenrods (Solidago 
spp.), and other species was thinner where broods were found than in 
other spots. 

As noted by others (2, 23), the grouse brood is held together by the 
hen's vocalizations. When a brood is flushed, the hen typically feigns 
injury, attempting to decoy the intruder away from the hiding young. 
After a more or less intensive display, the hen flushes to rejoin the 
brood, responding to their shrill peeping whistle with her own rasping 
peep or clucking. The intensity of the bond between hen and chicks ap- 
pears strongest when chicks are less than 10 weeks old (up to July 30) 
and gradually weakens as they mature. This was shown by timing the 
return of flushed hens to their broods (Fig. 1). By September, hens did 
not try to hold broods together, and at this time break-up of broods and 
wandering of some individuals occur. 

A relatively small number, 26, of different grouse were trapped and 
marked in this study. Marked grouse were recaptured or identified by 
sighting their color bands 11 times. The recovery data are insufficient to 
suggest more than considerable variation in the tendency to move. One 
juvenile captured in July was recovered in August, 1,800 feet away in 
the same valley; and was recovered a second time in October, 1,600 feet 
from the last site, this time in another valley. This bird was with a brood 
on the first two occasions, but was alone the last time. Another mem- 
ber of the same brood was recaptured the same day in October as the 
first juvenile mentioned, but in the same trap where it was originally 
caught. These records from two individuals in the same brood suggest 
that young grouse move up and down a valley system and sometimes 
wander into other valleys as brood separation occurs. In 18 resightings 
of marked grouse frcm the Geiger Ridge area, Thurman (25) found 9 
juveniles less than 300 yards between points of observation, although a 
10th was recovered 2% miles away and the 11th, 5 Ms miles away. 
Adults on the average were even more sedentary than the shorter- 
ranging juveniles. Although our data, and those of Thurman (25) as 
well, are slight by comparison, they fit the general conclusion of Hale 
and Dorney (11) that juvenile ruffed grouse are more mobile than adults. 



182 Indiana Academy of Science 

50n 



40 - 



c/> 30 
Id 

Z> 

i 



20- 



10 



Figure 1. 
broods. 



i n i i i i I i i i 

16 23 30 7 14 21 28 4 II 18 
JUNE JULY AUGUST 

Time required for ruffed grouse hens flushed with broods to return to the 



In this respect, Indiana grouse resemble northern birds. The phenomenon 
of juvenile mobility implies a sexual difference as discussed by Gullion 
(10) on the basis of his extensive experience with Minnesota grouse. 
Young cocks evidently prefer to claim territories within the familiar 
brood range, whereas young hens disperse more erratically to new 
habitats. 

Drumming Males and Breeding Populations 

During the last week of March and the first week of April 1968, 
drumming males found on the study area totaled 12 or 1 per 70 acres. 
In 1969, the count was 15 or 1 per 56 acres. Thurman's (25) count for 
this area in 1966 was 25 males or 1 per 32 acres. Wise (personal com- 
munication) found 13 drumming logs on an 835-acre area 6 miles from 
Geiger Ridge. Lewis et al. (15) estimated 2 grouse per 100 acres in 
Missouri, half of them males. 

The number of drumming males on an area from one year to the 
next is an indicator of breeding male population trends although not a 



Ecology 183 

precise one (6). If a total census of drumming males can be obtained, 
it is often assumed that a sex ratio of 1:1 exists, hence the total spring 
population is at least twice the number of drumming males. This is a 
conservative estimate since nondrumming males are overlooked and since 
selective predation is heavier on males than on females in late winter 
and early spring (8). In spite of caution recommended in estimating 
breeding populations or trends by this method, the difference between 
Thurman's 25 drumming males and Muehrcke's 15 appears significant. 
As discussed above, 1968 apparently was a poor production year. 

Further evidence of a reduced 1969 population is seen in the dis- 
tribution of drumming males. Thurman found 13 drumming territories 
in the unburned hardwoods. Muehrcke found only 2 there, but found 13 
territories in the burned section, which is believed to be the better grouse 
habitat. In 1969 the population was small enough to allow most terri- 
torial males to establish themselves in the better habitat. In view of the 
wet weather catastrophe in 1968, and the attachment of sexually ma- 
turing males to their brood territories, the situation also suggests better 
survival of young in the burn. 

Drumming Log Sites 

Biologists (9, 20) have observed that grouse use certain logs year 
after year. Gullion (9) called them perennial logs. In the present study, 
Muehrcke found 3 of the 25 drumming logs identified by Thurman in 
1966 still used in 1968. Eight of 12 logs used in 1968 were also used in 
1969. In their analyses of drumming log relationships, neither Palmer 
(20) nor Gullion (9) found a single site quality factor or other good 
explanation for the phenomenon of traditional use. 

Detailed data were collected for the physical features of drumming 
logs in 1969. On 6 of 15 territories, alternate logs were also used. De- 
composition of logs varied from moderate to very much. Their diameters 
ranged 9-16 inches, averaging 12.8. Total length ranged 8-38 feet, averag- 
ing 23.2. Distance the log lay from ridge top usually ranged from less 
than 10 to more than 600 feet, although 4 logs were fairly well down 
slope. Distance from field edge was less than 100 feet except those on 
slopes which were 210 to more than 600 feet away. Hardy (12) located 
18 drumming sites on his eastern Kentucky study area, all on or near 
ridge tops; and Thurman (25) tabulated the exposure of 25 sites on the 
Geiger Ridge area, noting that some were on ridge tops and others on 
slopes. The distance of the latter from ridge tops is not on record but 
CMK remembers that many were near the tops. The higher levels for 
display sites may offer some survival value in permitting flushed males 
to fly downward quickly into denser escape cover. Otherwise, a ridge 
top location may play a part in natural selection to the extent that 
sound may travel farther from a higher than a lower level, and hence 
attract more distant hens. 

A vegetation analysis of drumming log and brood location sites is 
summarized in Table 1. According to basal areas of hardwood species, 



184 Indiana Academy of Science 

drumming male grouse use more mature timber than broods. Otherwise, 
stems per acre and importance values show the essential nature of young 
growth and shrub species in both habitats. This coincides with Palmer's 
(20) observation that woody vegetation over 8 feet high is more dense 
near drumming sites than in surrounding cover. We believe that our 
grouse generally inhabit dense vegetation in the valleys during inclement 
weather of spring, and that males move upward to drumming logs in 
brushy areas dominated by older trees. 

1969 Drumming Season 

On January 15 at least two grouse visited drumming logs as shown 
by tracks in snow and droppings around logs. Snow cover shortly dis- 
appeared and no further activity around logs was apparent until 
February 24. Droppings showed a number of logs visited, but undisturbed 
leaves below the stages (spots on the logs where the grouse habitually 
stand to drum) hinted that drumming had not yet begun. First drum- 
ming was heard February 28 after a week of mild weather when tempera- 
tures ranged from a high of 41° F to a low of 28° F. Drumming activity 
then increased in the following days with 4 males drumming on March 7, 
and droppings showing that other males were visiting their logs. From 
March 8 to 17 all males were silent during which time temperatures 
ranged from 10° F to 21° F. After that the number of different males 
drumming increased until April 10 when all 15 males drummed. 

The peak of drumming intensity, as measured by the largest num- 
ber of different active males, extended from April 7 to 12 inclusive. No 
less than 8 and as many as 15 of the 15 different males performed daily. 
Thurman (25) gives the first week in April for this intense activity 
period in 1966. In 1969, drumming intensity tapered off until only 2 males 
were heard on April 23. No observations were made after that. 

As also noted by Petraborg et al. in Minnesota (22), Muehrcke (19) 
found the daily duration of drumming was longest in the period of peak 
activity, with drumming beginning earlier before sunrise and ending 
later after sunrise as the peak period approached. On April 8 one grouse 
started drumming at 0515 hours EST, the earliest time noted, and 
stopped at 0614. On April 6, grouse drummed sporadically on the study 
area until 1710 hours exept for a silent period from 1200 to 1410 hours. 
Usually the early morning drumming period ended about x k hour after 
sunrise, but some males returned to drum intermittently for another 2 
hours. Bent (2) also reported this behavior. 

During the peak period of drumming activity, Muehrcke, from a 
hide, observed one cock intensively to time the frequency of successive 
drumming performances. His raw data are less extensive and precise than 
those of Palmer (21) but do show that the time interval between drums is 
shortest just before and just after sunrise. The raw data further show 
that the intervals were shorter when a second grouse is present. It is to 
be expected that the presence of a hen stimulates an increased rate of 
drumming (8). The data are summarized in Table 2. They show that 



Ecology 185 

drumming" behavior was more erratic when a second grouse was present; 
drumming continued longer in the morning; the intervals between drums 
were extended as the cock left his log frequently to react with the 
visitor; and length of drums ranged up to 3 sec longer than when the 
cock was alone. 

One aberrant cock often drummed between 1500 and 1800 hours in 
the same area where a grouse drummed in evenings in 1988. Gullion (8) 
observed that some males are predominantly afternoon or evening drum- 
mers. Night drumming apparently occurs commonly (2). On calm, moon- 
lit nights, grouse drummed at 2200 hours on 2 occasions, and at 0345 
on 3 occasions on Geiger Ridge. Thurman (25) also heard night drum- 
ming on the area. 

Table 2. Time data for ruffed grouse drumming performances on Geiger 
Ridge study area, 1969. 





Drumming male 


Second grouse 




was alone 


was present 


Date 


March 23 


April 4 


First drumming 


0555 EST 


0535 


Last drumming 


0655 


0727 


Number of drums 


24 


39 


Interval between drums 






Range (minutes) 


1-5 


1-10 


Average 


2.3 


2.9 


Length of drums 






Range (seconds) 


4-6 


3-9 


Average 


5.3 


5.3 



Only certain males roosted on their drumming logs and even then 
uncommonly except for the week of most intensive activity. During that 
7-day period, 6 logs were used 1-3 nights and 4 logs were used 4 nights, 
Muehrcke actually flushing grouse from these logs. One grouse flushed 
from a drumming log at 2100 on September 19. More intensive field work 
in the fall would probably show that this is not unusual. Other studies (9) 
have shown that many drumming logs are closely attended in summer as 
well as in fall. Young males, seeking to become established on logs in fall, 
probably cause increased activity in older, established males. 

Male grouse are especially wary when drumming and react to 
unfamiliar sounds or movement by remaining silent or taking flight. 
Grouse leave their logs by walking if undisturbed, or by running or 
flushing apparently according to their level of fright. All drumming males 
habitually performed from a main log but had one or two alternative logs. 
At least 7 males drummed on alternate logs. If a bird ran from its main 
log before sunrise during the peak drumming period, it invariably 
drummed on an alternate log within the half hour. Males flushed from 
their main logs would not use alternate logs the same day but the follow- 
ing day. 



186 Indiana Academy of Science 

Literature Cited 

1. Allen, D. L. 1962. Our wildlife legacy. Funk and Wagnalls Co., New York. 421 p. 

2. Bent, A. C. 1932. Life histories of North American gallinaceous birds. U. S. National 
Mus. Bull. 162, Washington, D. C. 490 p. 

3. Bump, Gardiner, R. W. Darrow, F. C. Edminster, and W. F. Crissey. 1947. The 
ruffed grouse — life history — propagation — management. New York State Cons. Dept. 
Albany. 915 p. 

4. Chambers, R. E., and P. F. English. 1958. Modifications of ruffed grouse traps. 
J. Wild! Mgmt. 22:200-202. 

5. Deam, C. C. 1940. Flora of Indiana. Dept. of Cons., Indianapolis. 1236 p. 

6. Dorney, R. S., D. R. Thompson, J. B. Hale, and R. F. Wendt. 1958. An evaluation 
of ruffed grouse drumming counts. J. Wildl. Mgmt. 22 :35-40. 

7. Edminster, F. C. 1954. American game birds of field and forest. Chas. Scribner's 
Sons, New York. 490 p. 

8. Gullion, G. W. 1966. The use of drumming behavior in ruffed grouse population 
studies. J. Wildl. Mgmt. 30:717-729. 

9. Gullion, G. W. 1967. Selection and use of drumming sites by male ruffed grouse. 

Auk 84 :87-112. 

10. Gullion, G. W. 1969. The ruffed grouse in northern Minnesota. Processed report. 
Forest Research Center, U. Minn., Cloquet. 20 p. + 18 append. 

11. Hale, J. B., and R. S. Dorney. 1963. Seasonal movements of ruffed grouse in Wis- 
consin. J. Wildl. Mgmt. 27:648-656. 

12. Hardy, F. C. 1950. Ruffed grouse studies in eastern Kentucky. Ky. Div. Fish and 
Game, Frankfort. 26 p. 

13. Haymond, Rufus. 1856. Birds of southeastern Indiana. Proc. Acad. Natur. Sci. 
VIII :286-298. 

14. Korschgen, L. J. 1966. Foods and nutrition of ruffed grouse in Missouri. J. Wildl. 
Mgmt. 30:86-100. 

15. Lewis, J. B., J. D. McGowan, and T. S. Baskett. 1968. Evaluating ruffed grouse 
reintroduction in Missouri. J. Wildl. Mgmt. 32:17-28. 

16. Lindsey, A. A., R. O. Petty, D. K. Sterling, and Willard Van Asdall. 1961. Vege- 
tation and environment along the Wabash and Tippecanoe Rivers. Ecol. Monogr. 
31 :105-156. 

17. Lindsey, A. A., W. B. Crankshaw, and S. A. Qadtr. 1965. Soil relations and dis- 
tribution map of the vegetation of presettlement Indiana. Bot. Gaz. 126:155-163. 

18. Mosby, H. S. (editor). 1963. Wildlife investigational techniques. The Wildlife Society, 
Washington, D. C. 419 p. 

19. Muehrcke, J. P. 1969. Observations on ruffed grouse ecology and behavior in south- 
eastern Monroe County, Indiana. Unpub. master's thesis, Purdue University, Lafay- 
ette, Indiana. 98 p. 

20. Palmer, W. L. 1963. Ruffed grouse drumming sites in northern Michigan. J. Wildl. 
Mgmt. 27:656-663. 

21. Palmer, W. L. 1969. Time frequency between successive drumming performances of 
ruffed grouse. Wilson Bull. 81 :97-99. 

22. Petraborg, W. H., E. G. Wellein, and V. E. Gunvalson. 1953. Roadside drumming 
counts a spring census method for ruffed grouse. J. Wildl. Mgmt. 17:292-295. 

23. Sawyer, E. J. 1923. The ruffed grouse with special reference to its drumming. 
Roosevelt Wildl. Bull. 1 :355-386. 

24. Sharp, W. M. 1963. The effects of habitat manipulation and forest succession on 
ruffed grouse. J. Wildl. Mgmt. 27:664-671. 

25. Thurman, J. R. 1966. Ruffed grouse ecology in southeastern Monroe County, Indiana. 
Unpub. master's thesis, Purdue University, Lafayette, Indiana. 101 p. 



Fox Bounty in Indiana During the Years 1961 through 1968 

Ralph D. Kirkpatrick, Ball State University 



Abstract 

Fox bounty records for 1961 through 1968 were obtained from each county auditor 
in Indiana. The number of Indiana counties paying fox bounties declined from 63 in 
1961 to 51 in 1968. Numbers of fox bountied ranged from 20,066 in 1963 to 23,863 in 1966. 
Fox bounty records may reflect gross fox population fluctuations. Estimates of relative 
abundance of foxes in Indiana, derived from the annual Sportsman's Questionnaire and 
the annual Furbuyer's Report are compared to data from fox bounty records. During this 
8-year period, the number of foxes bountied peaked at 1.036 foxes per square mile in 
1966. The number of fox pelts purchased by Indiana fur buyers also peaked in 1966 at 
0.258 pelts per square mile. Numbers of fox pelts purchased annually indicated a greater 
correlation with total numbers bountied than with economic value of pelts. A lesser 
degree of correlation was obtained between numbers of foxes bountied and numbers of 
hunter-killed foxes as determined by the annual Sportsman's Questionnaire. 

Laporte County paid fox bounties from 1875 through 1877. Fox 
bounties were not paid in Indiana again until 1911. One or more counties 
have expended funds for fox bounties each year since that date (2). 

Early Indiana fox bounty records through 1948 are given and dis- 
cussed by Haller (2). Fox bounty records for the years 1949 through 
1960 are listed and discussed by Brooks (1). No Indiana state agency 
compiles fox bounty records by county or by year. Both Haller and 
Brooks comment on the difficulty of compiling an accurate and complete 
fox bounty record. County records, including fcx bounty information, 
tend to be stored in courthouse attics or basements where they are 
occasionally damaged or lost. 

Fox bounty records for each Indiana county for the years of 1961 
through 1968 were secured in the present study. Numbers of foxes 
bountied were compared with numbers cf fox pelts purchased by Indiana 
furbuyers and with numbers of foxes supposedly killed by Indiana 
hunters. 



Methods 

Fox bounty records for the years 1961 through 1968 were secured 
by a personal contact with every County Auditor's Office in northeastern 
Indiana. A questionnaire was mailed to all other County Auditors. County 
Auditors that did not return to questionnaire were contacted in person 
or by telephone. 

Records of the number of fox pelts purchased by Indiana furbuyers 
and average fur prices were secured from the Annual Furbuyers Reports 
filed with the Indiana Department of Natural Resources (William E. Ginn, 
personal communication). 

Estimates of annual hunter kill of fox were obtained from the 
Annual Sportsman's Questionnaire compiled by the Indiana Department of 
Natural Resources (William E. Ginn, personal communication). 

187 



188 



Indiana Academy of Science 



Results and Conclusions 

The number of counties paying fox bounty has declined from 63 in 
1961 to 51 in 1968, with the majority of bounty-paying counties being 
located in northern Indiana (Table 1, Fig. 1). States north and east of 
Indiana have also experienced a decline in the number of counties paying 



Table 1. Number of foxes bountied in Indiana 1961 through 1968. 



County 


1961 


1962 


1963 


1964 


1965 


1966 


1967 


1968 


Total 


Adams 


270 


17ti 


1 95 


218 


206 


242 


203 


252 


1,762 


Allen 


408 


808 


241 


444 


297 


8S1 


525 


390 


2,989 


Benton 


407 


318 


2S1 


310 


233 


306 


240 


346 


2,441 


Blackford 


224 


IBS 


174 


207 


205 


223 


184 


141 


1,526 


Boone 


584 


496 


546 


299 


347 


425 


00 


00 


2,647 


Carroll 


458 


467 


482 


42S 


377 


519 


442 


236 


3,354 


Cass 


308 


8X1 


500 


400 


49S 


500 


500 


500 


3,587 


Clinton 


62S 


421 


8S7 


36S 


321 


627 


407 


332 


3,491 


Decatur 


75 


89 


XS 


94 


1 05 


00 


00 


00 


451 


DeKalb 


454 


308 


810 


320 


321 


366 


352 


307 


2,741 


Delaware 


450 


405 


880 


271 


80S 


435 


306 


309 


2,814 


Dubois 


769 


767 


00 


00 


00 


00 


00 


00 


1,536 


Elkhart 


1S1 


228 


203 


2S3 


3S7 


422 


3S3 


366 


2,448 


Fayette 


133 


4X9 


250 


1S9 


192 


240 


207 


234 


1,934 


Fountain 


842 


8 IS 


227 


213 


1S3 


325 


390 


312 


2,310 


Fulton 


491 


8S0 


872 


416 


500 


S2S 


46S 


442 


3,897 


Grant 


850 


870 


876 


319 


351 


435 


271 


267 


2,739 


Hamilton 


400 


897 


824 


309 


301 


311 


2S4 


262 


2,588 


Hancock 


405 


842 


321 


236 


211 


361 


303 


299 


2,478 


Hendricks 


701 


500 


472 


391 


429 


579 


500 


493 


4,065 


Henry 


487 


857 


800 


196 


274 


404 


462 


425 


2,905 


Howard 


500 


4S0 


460 


430 


890 


3S0 


400 


223 


3,263 


Huntington 


844 


857 


2S8 


297 


273 


3S7 


270 


279 


2,490 


Jasper 


762 


658 


655 


599 


519 


672 


774 


543 


5,182 


.) a y 


281 


895 


812 


335 


221 


2S0 


243 


200 


2,217 


Jefferson 


820 


800 


801 


345 


00 


00 


00 


00 


1,266 


Jennings 


167 


100 


816 


263 


00 


00 


00 


00 


846 


Johnson 


00 


00 


00 


00 


226 


252 


2S3 


36S 


1,129 


Knox 


00 


00 


00 


170 


200 


220 


00 


00 


590 


Kosciusko 


494 


550 


5 OS 


677 


6S7 


S33 


594 


565 


4,908 


LaGrange 


248 


2S0 


2S9 


266 


264 


3S9 


413 


331 


2,480 


Lake 


479 


548 


400 


465 


313 


315 


292 


330 


3,137 


LaPorte 


5 t;t; 


624 


665 


696 


667 


7S3 


929 


732 


5,662 


Madison 


408 


488 


293 


323 


320 


389 


414 


307 


2,882 


Marion 


248 


287 


143 


22S 


97 


123 


171 


00 


1,247 


Marshalll 


47X 


857 


33S 


552 


471 


6S9 


54S 


517 


3,950 


Miami 


871 


821 


362 


347 


863 


42S 


3S0 


347 


2,919 


Monroe 


401 


2S0 


314 


395 


325 


500 


590 


520 


3,325 


Montgomery 


441 


504 


455 


350 


440 


00 


00 


00 


2,190 


Morgan 


425 


850 


242 


2S5 


266 


333 


330 


571 


2,802 


Newton 


682 


874 


43S 


46S 


353 


524 


371 


421 


3,631 


Noble 


490 


801 


4S6 


533 


672 


6S6 


471 


376 


4,015 


Ohio 


81 


87 


45 


65 


00 


00 


00 


00 


178 


Owen 


00 


00 


402 


399 


269 


00 


00 


00 


1,070 


Parke 


888 


81 S 


27S 


196 


213 


342 


331 


332 


2,343 


Porter 


00 


00 


00 


495 


500 


500 


400 


446 


2,341 


Posey 


840 


840 


00 


00 


00 


00 


00 


00 


680 


Pulaski 


600 


560 


496 


664 


551 


691 


612 


5 1 5 


4,689 











Ecology 








189 


Table 


. Numb 


er of foxes boun 


ied in Indiana 1961 thron 


gh 1968- 


-Contim 


ed. 


County 


1961 


1962 


1963 


1964 


1965 


1966 


1967 


1968 


Total 


Futnam 


425 


380 


380 


325 


396 


466 


466 


466 


3,304 


Randolph 


00 


00 


00 


00 


00 


00 


537 


442 


979 


Ripley 


460 


357 


329 


277 


00 


00 


00 


00 


1,423 


Rush 


391 


331 


295 


273 


298 


442 


333 


00 


2,363 


St. Joseph 


200 


228 


232 


392 


300 


366 


352 


405 


2,475 


Scott 


L67 


174 


157 


85 


83 


17 


17 


00 


700 


Shelby 


442 


387 


328 


293 


268 


366 


442 


499 


3,025 


Starke 


2f>r> 


230 


215 


228 


298 


348 


246 


228 


2.048 


Steuben 


365 


222 


290 


374 


499 


500 


500 


655 


3,405 


Sullivan 


00 


00 


00 


00 


250 


300 


250 


00 


800 


Switzerland 


85 


77 


116 


00 


111 


00 


00 


00 


389 


Tippecanoe 


47(1 


301 


263 


36,7 


321 


398 


403 


332 


2,855 


Tipton 


369 


300 


274 


328 


300 


278 


268 


210 


2,327 


Union 


229 


250 


249 


223 


218 


245 


273 


00 


1,687 


Vermillion 


190 


174 


163 


195 


178 


206 


267 


266 


1,639 


Vigo 


284 


333 


358 


335 


313 


384 


332 


00 


2,339 


Wabash 


253 


377 


292 


391 


362 


400 


400 


325 


2,800 


Warren 


400 


299 


276 


176 


169 


194 


211 


194 


1,919 


Wells 


269 


336 


236 


302 


342 


405 


324 


306 


2,520 


White 


576 


472 


477 


423 


503 


428 


467 


405 


3,751 


Whitley 


1X9 


366 


326 


420 


385 


475 


359 


273 


2.793 


Total 


23,843 


22,168 


20,066 


20,751 


20,243 


23,863 


21,690 


18,842 


171.676 


Counties 




















Paying 


63 


63 


62 


63 


60 


56 


54 


51 


69 


Bounty 




















Mean No. 




















Bountied/ 


378.5 


348.7 


323.6 


335.9 


337.4 


426.1 


401.7 


369.5 


2,488.1 


County 























fox bounty. The number of Ohio counties paying fox bounty declined from 
66 in 1961 to 52 in 1968 (5) and Michigan stopped paying fox bounty 
entirely in 1965 (3), 

The custom of fox chasing persists in southern Indiana but has prac- 
tically died out in northern Indiana where it was practiced in Grant 
County as recently as 1931 (4). Fox chasers have discouraged the expendi- 
ture of funds to pay fox bounty in southern Indiana counties. A reason 
for the discontinuance of bounty payments in the few southern counties 
that appropriated bounty funds is the import of dead foxes from nearby 
counties. 

The mean number of foxes bountied per Indiana county peaked in 
1966 when an average of 426.1 foxes (or 1.036 foxes per square mile) 
were bountied per county (Fig. 2). 

Brooks (1) reported peak Indiana fox bounty payments in 1926, 1934, 
1946, 1954, and 1960. It appears that, if bountied foxes actually are an 
index to total fox numbers, the Indiana fox population fluctuates with 
high populations occurring each 6 to 8 years. 



190 



Indiana Academy of Science 



• 'H-H I I I I M° 




PAID BOUNTIES 
IN 1968 



• • ceased paying 

o* # BOUNTIES 

PRIOR TO 1968 



Figure 1. Map of Indiana showing counties reporting bounty payments during the 
years 1961 through 1968. 



Fox pelts sold may reflect fur price to a greater extent than it 
reflects fox numbers, but Figure 3 indicates that this is not true during 
the last 8 years in Indiana. 



Ecology 191 





1.5 




1.4 




1.5 




1.2 




1.1 


4) 
rH 


1.0 


a 




u 

CO 

cr 

CO 


0.9 
0.8 


U 
ft 


0.7 


CD 

X 

o 


0.6 


O 


0.5 


u 

1 

S25 


0.4 
0.5 




0.2 




0.1 












\____ 



1961 1962 1965 1964 1965 1966 1967 1968 

••••fox kill by hunters* 

11 ■ foxes bountied 

— — fox pelts sold to furbuyers 

* fox kill data for 1968 is not available; validity of kill 
data for I966 is questionable 

Figure 2. Comparison of three measures of fox population levels in Indiana for the 
years 1961 through 1968. 



Bounty amount paid per fox during the years of this study has been 
$3.00 per animal. Brooks (1) points out that the amount paid per fox does 
not directly affect numbers of foxes bountied per year. Fluctuations in 
numbers of foxes bountied during the years of 1961 through 1968 
parallel fox kill data supplied by the Indiana Sportsman's Questionnaire 
(Fig. 2). These data may be an index of gross populations fluctuations. 



192 



Indiana Academy of Science 



u 

CO 

C 
00 


1.10 
1.05 


u 


1.00 


a, 




CO 


0.95 




0.90 


T3 
■•-> 

c 
o 


0.85 
0.80 


<3 


0.75 


,0 


0.70 


53 


O.65 




6.5c 

6.00 
5.50 
5.00 
4.50 
4.00 
3.50 
3.00 
2.50 
2.00 
1.50 
1.00 

0.50 

0.00 



1961 1962 1963 1964 1965 1966 1967 1968 



foxes bountied 
• • •• price paid per red fox pelt 
— — price paid per gray fox pelt 

Figure 3. Comparison of average prices paid for fox pelts by Indiana fur buyers with 
numbers of foxes bountied per square mile for the years 1961 through 1968. 



Literature Cited 

1. Brooks, D. M. 1962. A study of the fox population. Ind. Dept. of Cons., Wildl. Res. 
Rep. 23(2) :68-118. 

2. Haller, F. D. 1950. The bounty system in Indiana. Ind. Dept. of Cons., Wildl. Res. 
Rep. 11(2) : 93-125. 

3. Laun, C, ed. 1968. 68 bounty survey (United States). Bounty News, 2(1) :n. p. 

4. Leopold, A. 1931. Report on a game survey of the North Central States. Sporting 
Arms and Ammunition Manufacturers' Institute, Madison, Wisconsin. 299 p. 

5. Russell, K. R. 1968. Fox bounty payments in Ohio, 1944-1967. Wildlife In-Service 
Note No. 97, Ohio Dept. of Natural Resources, Division of Wildlife. 10 p. 



Naturalized Bi*j; Trefoil (Lotus pedunculatus Cav. ) Ecotypes 
Discovered in Crawford County, Indiana 

Maurice E. Heath, Purdue Agricultural Experiment Station 



A bstract 

In 1964, eight naturalized ecotypes of big trefoil, (Lotus pedunculatus Cav. ; for- 
merly L. ugilinosis Schkuhr., L. major Sm.) were discovered growing in a tall fescue 
(Festuca arundinacea Schreb. ) meadow on the Clarence E. Kaiser farm in Crawford 
County. Subsequent evaluations show three of the ecotypes superior in forage and vegeta- 
tive characteristics for use in tall fescue pastures on fragipan soils in southern Indiana. 
The environmental model under which the big trefoil ecotypes have become naturalized 
is described. 

Big trefoil (Lotus pedunculatus Cav.), where adapted, is a long 
lived perennial forage legume. It is of European origin and is a recog- 
nized forage plant in France, Italy, Denmark, Germany and other 
countries. 

It is not definitely known when big trefoil was introduced into the 
United States. However, it has been grown in this country for over 95 
years (5). It has become naturalized in western Oregon, western Wash- 
ington and northwest California where it is used for pasture and hay. It 
has shown good adaptation on wet soils in Georgia, North Carolina and 
Florida (3,4). 

In 1947, Roland McKee (personal communication), Senior Agrono- 
mist for the Bureau of Plant Industry, U.S. Department of Agriculture, 
reported, ". . . some work with Lotus uliginosus for the southern part of 
the Plains Corn Belt region would be justified as I am inclined to believe 
that by selection, a sufficiently hardy strain can be obtained for use in 
combination with some of the grasses." Several accessions of big trefoil 
were tested on the Southern Indiana Forage Farm, Dubois County, Indi- 
ana, in 1954-55 but failed to survive the winter. 



Ecotypes Discovered 

On August 13, 1964, Mr. Clarence E. Kaiser showed me big trefoil 
growing in eight different areas in a vigorous tall fescue (Festuca 
arundinacea Schreb.) field on his farm (Fig. 1). Mr. Kaiser thought the 
plants to be a particularly adapted form of birdsfoot trefoil (L. cornicu- 
latus L.) since the areas were gradually increasing in size. Upon exami- 
nation of the rhizomatous and stoloniferous type of growth and spread- 
ing, I immediately recognized these plants to be big trefoil. Mr. Kaiser's 
farm is in Crawford County, Indiana, located 1 mile east and 1 mile 
south of the junction of Indiana Highways 164 and 145. 

193 



194 



Indiana Academy of Science 



^IbsSPl 








- - - #..*" 



Figure 1. Naturalized big trefoil (Lotus pedunculatus Cav.) discovered August 
growing on fragiyan soils on Mr. Clarence E. Kaiser's farm in Crawford County, 
Mr. Kaiser is second from the right. 






IS, 1964, 

Indiana. 



Mr. Kaiser believes that the big trefoil was a contaminant in a 
grass-legume seed mixture he frost seeded over the field in the late winter 
of 1939. The seed mixture included: 

Rate per acre 
Species in pounds 

Birdsfoot trefoil (L. corniculatus L.) 1 

Ladino clover (Trifoliwm repens L.) 1 

Alsike clover (T. hybridwm L.) 1 

Timothy (Phleum pratense L.) 4 

Smooth bromegrass (Bromus inermis Leyss.) 5 

Redtop (Agrostis alba L.) 2 

In 1941, five pounds per acre each of tall fescue and orchardgrass 
(Dactylis glomerata L.) seed were drilled into the meadow. Because of 
the excellent adaptation of tall fescue the field is now a heavy fescue sod. 



How Field Was Managed 

Each year since the 1939 seeding, the spring growth of grasses and 
legumes has been harvested for hay. The summer and fall growth has 
been used for beef cow pasture from late September following calf 
weaning until late January. 

From 1939 to 1951, a mid-summer Ladino clover seed harvest was 
made annually. In 1964, red clover (T. pratense L.) was frost seeded in 
late winter. The red clover has been allowed to produce and shatter seed 



Ecology 



195 




Figure 2. Primary area of adptation of tall fescue (Festuca arundiacsa Schreb.) in 
the east humid area of the United States including the southern third of Indiana (1). 



on the ground every other year. Thus, the red clover has behaved as a 
reseeding biennial legume growing in the tall fescue sod. The meadow 
has been fertilized every other year with phosphorus and potassium 
according to soil test and limed as needed. 

The Environmental Model (2) 

The hill soils on Mr. Kaiser's farm belong to the Zanesville-Wellston 
soil association commonly found throughout the unglaciated sandstone 
shale region. The soil fragipan common to many of these soils restricts 
water movement and root penetration. The shallow root zone depth may 
range from 18 to 34 inches. Although rainfall averages 46 inches 
annually, 20 inches are lost as run-off, and droughts are common in 
mid-summer. The soils are often water-logged in winter and early 
spring. Severe winter-heaving is a common occurrence. The soil moisture 
is one of extremes from wet to dry. Tap rooted alfalfa is considered a 
high risk crop on these soils when compared to tall fescue and biennial 
red clover. The native pH of these soils ranges from 5.3 to 5.5. 



Tall Fescue Adaptation 

The southern third of Indiana is in the primary area of adaptation 
of tall fescue (Fig. 2). Since 1950, tall fescue has been increasingly grown 
on the fragipan soils of southern Indiana where it currently excells all 
other perennial grasses in adaptation. 

It is believed that big trefoil may have its greatest promise for use 
in southern Indiana as a perennial pasture legume in combination with 



l ;m; 



Indiana Academy of Science 






1 ^ v ^iO.-. '* • ., 




I'x.'- 




/.- 








FIGURE 3. One of the eight ecotypes of big trefoil observed to be associating compatabily 
with a heavy stand of tall fescue on the Kaiser farm (Photographed June 17, 1966) 




Figure 4. One of many big trefoil nursery plants growing on fragipan soils on the 
Southern Indiana Purdue Agricultural Center, Dubois County, Indiana. It has spread 
three and a half feet through a good stand of tall fescue during a period of three growing 
seasons without evidence of any heaving. Tall fescue was seeded when the big trefoil 
seedlings were transplanted from the greenhouse April 15, 1967 (Photographed October 
3, 1969). 



Ecology 197 

tall fescue on fragipan soils (Fig. 3). Big trefoil appears to be a very 
efficient supplier of readily available nitrogen. Tall fescue has a dark 
green vigorous growth during the spring season when growing with big 
trefoil. 

Agronomic Progress 
Approximately 15 two-inch plugs were obtained from each of the 
eight ecotypes in the fall of 1965 for propagation and evaluation pur- 
poses. These were planted on the Southern Indiana Forage Farm in the 
spring of 1966. Two years of observations showed three of the ecotypes 
to be superior in vigor of growth and seed production. A total of 10.3 
pounds of seed of the 3 ecotypes were produced in 1968 for experimental 
purposes by special seed production facilities of the Minnesota Agricul- 
tural Experiment Station at Rosemont, Minnesota. Seed germination 
ranged from 85 to 90%. Excellent stands from seeding resulted in field 
plots at the Southern Indiana Purdue Agricultural Center (known as the 
Southern Indiana Forage Farm prior to 1969) in the spring of 1969. 
Seed produced from the three superior ecotypes (#1015, #1016 and 
#1019) equally blended together will be known as 'Kaiser' trefoil in honor 
of Mr. Clarence E. Kaiser who introduced, observed and nurtured these 
plants for many years on his farm. 

Significance of Discovery and Summary 

1. The naturalized ecotypes of big trefoil from the Clarence E. Kaiser 
farm in Crawford County, Indiana, are strongly perennial and appear 
to fit the environmental model of the hill land fragipan soils of the 
unglaciated sandstone shale soil region of southern Indiana. 

2. Five years of observation have not shown any disease or heaving 
tendencies among the eight big trefoil ecotypes discovered. 

3. Several of the big trefoil ecotypes compete aggressively and com- 
patibly with tall fescue (Fig. 4) which is the dominant perennial grass 
of the unglaciated sandstone shale soil region. 

4. The big trefoil ecotypes not only produce seed but they can also be 
readily established from seed. 

5. It is hypothesized that the ruminant response from the tall fescue-big 
trefoil mixture will be superior to that of the ruminant response from 
tall fescue alone. 

6. The seed produced from the three agronomically superior big trefoil 
ecotypes when equally blended together will be known as "Kaiser" 
trefoil in honor of Mr. Clarence E. Kaiser who introduced, observed 
and nurtured these plants for many years. 

Literature Cited 

1. Cowan, J. Ritchie. 1962. Forages. Iowa State University Press, Ames, Iowa. 300 p. 

2. Heath, Maurice E. 1969. Fitting plants to fragipan soils in southern Indiana. Proc. 
Indiana Acad. Sci. 78 :429-434. 

3. Henson, Paul R., and H. A. Schoth. 1962. The trefoils — adaptation and culture. 
ITSDA-ARS Agric. Handbook 223. 

4. McKee, Roland and H. A. Schoth. 1941. Birdsfoot trefoil and big trefoil. USDA Cir. 
625. 13 p. 

5. McKee, Roland. 1948. Big trefoil proved valuable as a forage crop for the West and 
South. USDA-ARS Res. Achiev. Sheet 101. 



The Forest Types of Indiana and a New Method of Classifying 
Midwestern Hardwood Forests 

Alton A. Lindsey and Damian V. Schmelz,i Purdue University 



Abstract 

Evidence from the plotting of modified importance sums for 3 species-groups of 
dominant forest trees on a 3 axis, triangular graph supported the classification of 58 
outstanding old-growth, relatively undisturbed stands in Indiana into four major forest 
types : Oak-hickory, Beech-maple, Lowland-depressional and Mixedwoods. Two clearly 
recognizable subtypes of the latter were well drained mixedwoods versus poorly drained 
mixedwoods. 

Other lines of evidence, such as the species association gradient presented herein, 
and a 3 dimensional ordinational model published elsewhere, corroborate this interpre- 
tation. 

This classification appears applicable to the hardwood forests of all nearby mid- 
western states where the forest stands also seem to be dominated in different sites by 
essentially the same 3 general species-groups, and intermediate mixtures. 

The Indiana Natural Areas Survey of 1967-9 provided an opportunity 
for extensive field work on the old growth forest stands of the state. 
Detailed descriptions of many such stands were presented in the book on 
the survey results (5). The best 36 forest stands (i.e., those least dis- 
turbed and mest nearly in equilibrium with their specific environment) 
were used for ordination and an intensive comparison of forest types. 
After completion of this work, the authors jointly developed several sim- 
pler, more readily applicable approaches for the classification and com- 
parison of midwestern hardwood forests, using 58 relatively undisturbed 
stands, including the 36 stands of the Schmelz and Lindsey analysis (7). 
The best one of these approaches appears worthy of recommendation 
for use in all midwestern states where 3 essentially similar groups of 
hardwood tree species attain high importance values; it therefore forms 
the basis of the present brief paper. The method depends on importance 
percentage sums of the respective species-groups, plotted on a special 
triangular graph. It does not require tediovs computations, ordination 
or the construction of three-dimensional models. 

The potential usefulness of the present method is not restricted to 
application by professional forest ecologists. Because of current and 
probable continuing interest in preservation of natural areas by citizens' 
groups, state conservation departments, federal agencies and others, an 
accurate but relatively simple approach to comparison and classification 
of forest stands should aid in determining priorities for official protection. 

Methods and Materials 

The 58 forest stands we used (Table 1) were either subjected to full 
tallies of substantial portions (up to 23 acres complete census) or were 
intensively sampled by a number of Vs acre strips or % acre strips, each 



Present address: Biology Department, St. Meinrad College, St. Meinrad, Indiana. 

198 



Ecology 199 

400 feet in length. The unusually thorough gathering of basic data was 
prompted by the demonstration by Lindsey, Barton and Miles (3) that 
a satisfactory level of adequacy depends on quite high intensity of 
sampling. 

In each stand, the following attributes were computed for each 
species represented by individual stems exceeding 4 inches dbh: density 
per acre (D-), relative density (D. f ), basal area per acre (B L .), relative 
basal area (B. { ) and importance (V 3 ). The latter was obtained by aver- 
aging relative density and relative basal area figures. The V3 for separate 
selected spe:i:s within a given stand were summed, to obtain the total 
importance for each of three basic species-groups which dominated and 
characterized stands representing particular forest types. 

The oak-hickory species-group included the species of upland Quercus 
and Carya characterizing rather xeric forest sites in Indiana, i.e., white 
(Q. alba), chestnut (Q. prinus) and black (Q. velutina) oaks, and 
pignut (C. glabra) and shagbark (C. ovata) hickories. The lowland- 
depressional species-group included 13 species on flood plains or poorly 
drained depressions — red and silver maples (Acer rubrum, A. saccha- 
rinum) , big shellbark hickory (C. laciniosa) , hackberry (Celt/is occi- 
dentalis), green ash (Fraxinus pennsylvanica) , sweet gum (Liquidam- 
bar styraciflua) , black gum (Nyssa sylvatica) , sycamore (Platanus 
occidentalis) , swamp white oak (Q. bicolor) , bur oak (Q. macrocarpa) , pin 
oak (Q. palustris) , Shumard oak (Q. Shiimardii) and American elm 
(Ulmus americana) . 

The beech-maple group typical of upland mesic sites consisted of 
Fagus grandifolia and Acer saccharum. 

The use of a graph in the form of an equilateral triangle made it 
possible to plot, by one dot for each stand, the selected population impor- 
tance sum for each of the 3 species-groups in a single plane. This graph, 
reproduced as Figure 1, shows values for the beech-maple species-group 
increasing upward, i.e., the base of the triangle stands for zero and apex 
of the triangle would represent 100% beech and /or sugar maple. How- 
ever, the plotted figures cannot be simply the straight importance per- 
centage sums. The total importance for a stand is actually 1007r, but 
we disregard some species, namely, those not included in any of the 
three species-group lists above. Because the use of the three-axis graph 
depends on the 3 values for each plotted point adding up to 100, we con- 
verted each species-group figure to a new per cent value, on the basis of 
a full 100 % for the sum of only the 3 species-group importance figures. 
These adjusted values, then, were plotted for Figure 1. 

Such figures for the oak-hickory species-group were plotted on the 
axis that has values increasing from the entire right-hand side (zero for 
OH) to the lower left point (100% for OH). Conversely, all along the 
left side is the zero value for the lowland-depressional species group 
component of a given stand, while this component increases rightward 
and downward to 100% LD species at the lower right-hand point. Upon 



200 



Indiana Academy of Science 




OAK-H ICKO RY 



An BB IS.LC.PI 
PO.SI.TI 



Figure 1. Location of 58 hardwood forest stands on a three-axis graph, plotting sums 
of the 8 species-group importance percentages per stand (see text) on the triordinates 
to determine the stand type. The 60 per cent importance level (heavy line) delimits the 
3 corners where beech-maple, oak-hickory and lowland depressional stands occur, while 
the stands falling centrally represent the mixedwoods type. (Abbreviated stand names 
are explained in Table 1. In several cases, one dot represents more than one closely 
similar stand). 



this 3-axis graph, then, we plotted the 3 major components of each of the 
58 stands selected for minimal history of disturbance by human activity. 

For example, the Bear Creek Valley slope forest (BV in Table 1) 
contained no lowland-depressional species, hence its dot (BV) fell on the 
zero LD line that forms the left side of the Figure 1 graph. The figures 
for the other axes were approximately 20% beech-maple and 80 % oak- 
hickory, as the BV dot in Figure 1 indicates. 



Results and Discussion 

Since the abovementioned position of stand BV on the graph closely 
approaches the oak-hickory (lower left) point, this stand doubtless rep- 
resents the oak-hickory forest type. It would be classed as an oak- 
hickory stand by the criteria of Crankshaw et «/. (2) or on any other 
basis of which we have heard. But how should stand BU (with the same 
[20%] beech-maple percentage but only 63% oak-hickory, and with 17% 



Ecology 



201 



Table 1. Names, symbols, forest types and locations of the 58 stands considered herein. 





County 


U.S.G.S. quad 


Town 


Range 


Section 


BEECH-MAPLE 












Allee (Al) 


Parke 


Montezuma 


16N 


xw 


3 


Bendix (Bx) 


St. Joseph 


Lydick 


37N 


1W 


11 


Caster (Ca) 


Montgomery 


Shannondale 


19N 


3W 


34 


Cring (Cr) 


Jay 


Portland 


23N 


14E 


10 


Hayes (Ha) 


Wayne 


New Paris 


14N 


1W 


35 


Hoot (Ho) 


Owen 


Patricksburg 


9N 


4W 


6 


Jackson (Jk) 


Ripley 


Milan 


7N 


12E 


18 


Logansport (Lo) 


Cass 


Anoka 


27N 


2E 


33 


Manlove (Mu) 


Fayette 


Connersville 


15N 


12E 


29 


Meltzer (Crosby) (MC) 


Shelby 


Rays Crossing 


12N 


8E 


7 


Nature Conservancy (NC) 


Montgomery 


Alamo 


17N 


6W 


2 


Officer's North (ON) 


Jefferson 


Volga 


4N 


9E 


22 


Pine Hills ( PH ) 


Montgomery 


Alamo 


17N 


6W 


1 


Pioneer Mothers (PM) 


Orange 


Paoli 


IN 


IE 


7 


Potzger (Po) 


Ripley 


Milan 


7N 


12E 


20 


Rocky Hollow (RH) 


Parke 


Wallace 


17N 


7W 


27 


Rosbrugh (Ro) 


Kosciusko 


Leesburg 


33N 


BE 


30 


Rush (Ru) 


Montgomery 


Alamo 


18N 


6W 


27 


S. LaPorte (SL) 


LaPorte 


LaPorte East 


36N 


3W 


2 


Spurgeon (Sp) 


Noble 


Ligonier 


35N 


9E 


18 


Warren (Wr) 


Berrien (Mich) 


Three Oaks 


7S 


1W 


27 


Weaver (Wv) 


Fayette 


Connersville 


15N 


12E 


30 


Wygant (Wy) 


Huntington 


Majenica 


27N 


10E 


3 


OAK-HICKORY 












Beall (Upland) (BU) 


Wabash (111.) 


Mt. Carmel 


2S 


13W 


11 


Bear Creek Plateau (BP) 


Fountain 


Stonebluff 


21N 


8W 


33 


Bear Creek Valley ( BV ) 


Fountain 


Stonebluff 


20N 


8W 


4 


Dunes (Xeric) (DX) 


Porter 


Dune Acres 


37N 


<;w 


L3 


Fox Island (FI) 


Allen 


Fort Wayne W. 


30N 


he 


25 


Johnson (Jo) 


Posey 


Wabash Is. 


17S 


14W 


32 


Lilly-Dickey (Ly) 


Brown 


Nashville 


9N 


3E 


8 


Ross Reserve (RR) 


Tippecanoe 


Otterbein 


23N 


tiW 


26 


Wing Haven (Wi) 


Steuben 


Angola E 


38 N 


13E 


35 


LOWLAND-DEPRESSIONAL 










Andrus (An) 


Knox 


E. Mt. Carmel 


IS 


12W 


11 


Beall (Bottom) (BB) 


Wabash (111.) 


Mt. Carmel 


2S 


13 W 


11 


Beckville (Bk) 


Montgomery 


Shannondale 


18N 


3W 


11 


Davis 1 (Dv) 


Randolph 


Redkey 


21N 


12E 


23 


Giants (WG) 


Vermillion 


Perrysville 


18, 19N 


9W 


3, 34 


Hemmer (Hm) 


Gibson 


Lynnville 


3S 


9W 


24 


Independence (IB) 


Tippecanoe 


Otterbein 


22N 


i;w 


3 


Kramer (Kr) 


Spencer 


Owensboro W. 


SS 


7W 


12 


Little Cypress (LC) 


Knox 


E. Mt. Carmel 


IS 


12W 


14 


Meltzer (Brookston) (MB) 


Shelby 


Ray's Crossing 


12N 


8E 


7 


Paramecium Is. (PI) 


White 


Buffalo 


28N 


3W 


1 


Pin Oak (PO) 


Gibson 


E. Mt. Carmel 


IS 


12W 


34 


Sigmoid Is. (SI) 


Carroll 


Brookston 


25N 


3W 


15, 16 


Terrace Is. (TI) 


Tippecanoe 


Brookston 


24N 


3W 


17 


Tippecanoe (Upper) (TU) 


Pulaski 


Bass Lake 


31N 


1W 


18, 19 


Wesselman (Ws) 


Vanderburgh 


Evansville 


6S 


10W 


22, 23 


MIXED WOODS 












Big Walnut (BW) 


Putnam 


Roachdale 


15N 


3W 


29, 31, 32 


Botany Glen (BG) 


Grant 


Gas City 


23N 


8E 


11, 14 


Bradford (Br) 


Morgan 


Mooresville W 


13N 


IE 


r> 


Clifty (CI) 


Montgomery 


Alamo 


17N 


<;w 


l 


Conboy (Cy) 


Jennings 


Vernon 


6N 


9E 


10 


Donaldson's (Dn) 


Lawrence 


Mitchell 


3N 


IE 


4 


Dunes (Mesic) (DM) 


Porter 


Dunes Acres 


37 N 


6W 


13 


McCormicks Cove (McC) 


Owen 


Gosport 


ION 


3W 


22 


Officer's (South) (OS) 


Jefferson 


Volga 


4N 


9E 


22 


Tippecanoe (Lower) (TL) 


Pulaski 


Bass Lake, 
Winamac 


31N 


1W 


18, 19 



202 Indiana Academy of Science 

lowland-depressional), be typed? An inspection of the distribution of all 
the stands on the graph aids in establishing improved criteria for stand 
classification. 

The 58 original or very old growth stands utilized do not appear as 
a continuum throughout Figure 1 since intergradation between the two 
sides (OH versus LD) appears lacking not only along the zero BM (base) 
line but throughout a large central region and high up toward the BM 
apex. The intergradations can be traced instead from LD with high 
available moisture supply, through BM with median moisture and drain- 
age (mesic sites), to OH with limited moisture and excessive drainage. 
Although these old stands must at present be relatively stabilized and in 
balance with their sites and overall environments in accord with current 
polyclimax interpretation, reading the graph upward from either the OH 
or LD corner parallels the successional trend in Clementsian theory from 
moisture extremes to optimum conditions for the beech-maple type on 
mesic sites. This conforms with the results shown by the much less direct 
approach (through 3-axis ordination technique) of Schmelz and Lindsey 
(7) based on a more stringently selected 36 stands of these 58. 

An area of some selected size subtending each point of the triangle 
will serve to delimit or define the major forest types for Indiana in a 
revised type classification. Any criteria adopted are somewhat arbitrary, 
as were those of Crankshaw et al. (2) and Lindsey et al. (4) in earlier 
papers. The present authors consider that the information in Figure 1 
supports recognition of 4 major hardwood forest types, when each "point" 
is delimited at the 60% cross line of the graph, as shown by heavier 
lines. The 3 forest types occupying the extreme site conditions, near the 
points of the triangle (Lowland-depressional, Beech-maple and Oak- 
hickory) are defined by having at least a 60 (adjusted) importance per- 
centage sum for the one dominant species-group, hence 40% or less for 
the other two species-group sums taken together. Stand BU, having 63% 
upland OH species, is therefore placed in the oak-hickory type. Obviously, 
the three importance sums are sufficient to designate forest types (with- 
out plotting stand positions on a graph) once a quantitative criterion is 
accepted. The graph facilitates broad comparisons. 

Stands that are not conspicuously dominated by a single species- 
group fall within the remaining central hexagonal figure. We term this 
hardwood type as Mixedwoods. This fourth type is subdivided rather 
clearly (within the stands examined, at least) into well drained and 
poorly drained subtypes, at the left-center and right-center respectively, 
and without intergradation between them within the Mixedwoods Type. 
The latter includes, basically, both the Mixed Woods and Western Meso- 
phytic Types of Schmelz and Lindsey (7) (both these types representing 
the well-drained Mixedwoods subtype) plus the poorly-drained mixed- 
woods subtype represented here by TL, OS and Cy at the right. The 
latter show very low oak-hickory representation, because low-ground oaks 
like pin and swamp white oaks are not included in the (upland) oak- 
hickory species-group. However, these 3 stands have moderate amounts 
of beech and /or sugar-black maple; this raises them above the lowland 



Ecology 203 

depressional type, into the Mixedwoods Type, as the subtype of the latter 
which lies intermediate between Beech-maple and Lowland-depressional 
rather than between Beech-maple and Oak-hickory. 

Since about 87% of Indiana was covered by forest in presettlement 
times (6), all substrate conditions paralleling the forest type triangle in 
Figure 1 probably supported forests in this state originally. The large 
central blank space appearing on the graph represents, by and large, 
the vegetation of sites without extremes of ponding or droughtiness, 
steep slope or excessive internal drainage. Since these median sites have 
been chiefly cleared for agricultural utilization, fine present-day forests 
tend to be restricted to sites less favorable for farming. It seems reason- 
able that mixedwoods were somewhat more prevalent in the original 
vegetation of Indiana than in that of today, not so much as an extensive 
mappable type as on some isolated median sites within the regions 
mapped (4, 5, 6) as beech-maple or oak-hickory. The portions of southern 
Indiana mapped previously (4, 5, 6) as "western mesophytic" we now 
prefer to consider Mixedwoods, in part because Aescuhis octandra and 
Tilia heterophylla which Braun (1) considered mesophytic indicators do 
not extend very far northward in Indiana. 

Data on soil requirements-tolerances of certain Indiana tree species 
were published by Crankshaw, Qadir and Lindsey in 1965 (2). We have 
arranged those species according to the ecological affinities shown in 
Table 2 into a gradient from black oak at the xerophytic end through 
black ash at the hydrophytic extreme. They fall clearly into the 3 
groups roughly corresponding with the 3 apical regions of our Figure 1 
graph and with the 3 extreme positions on the ordination model (of 36 
stands) published py Schmelz and Lindsey (7). The upland oak-hickory 
type (upper third of Table 2) prefers high per cent sand, whereas 
high silt content characterizes the sites of the dominant tree species of 
beech-maple forests, and high per cent clay favors lowland-depressional 
stands. Highly leached soils, low pH and low available nitrogen are 
compatible with oak-hickory. The lowland depressional species are 
favored by high available water, neutrality of reaction, high clay con- 
tent and at least 4 of them by high per cent nitrogen in the soil. 



204 



Indiana Academy of Science 



Table 2. Gradient of influence of the most important soil factors on tree 
species in the presettlement Indiana forest. Plus sign indicates high value 
favors species under natural competition. Minus sig?i indicates that low 
value is favorable or well tolerated. A reworking of certain data from 
Crankshaw, Qadir and Lindsey (2). 





Avail. 


Leach- 




% 


% 


% 


'/< 




H,0 


ing 


pH 


N 


Sand 


Silt 


Clay 


Black Oak 





+ 








+ 






Chinquapin Oak 


— 


+ 




— 


+ 






White Oak 






6.1 


— 


+ 


— 




W. B. Cherry 




— 


6.1 




+ 




— 


Shingle Oak 






— 




+ 






Post Oak 




+ 


— 


— 


— 






Upl. Hickories 




+ 




— 


— 


+ 




Red Oak 




+ 












Tulip Tree 




+ 








+ 




Sugar Maple 






+ 


+ 




+ 




Basswood 


+ 


— 


+ 


+ 








White Ash 


— 


< » 




+ 






+ 


Beech 




< > 






— 


+ 




Bur Oak 




+ 








+ 




Sweet Gum 




+ 


— 








+ 


Amer, Elm 






7 






— 


+ 


Bl. Walnut 


+ 




7 




+ 






Buckeye 


+ 




+ 


+ 


• 




+ 


Hackberry 


+ 





7 


+ 






— 


Cottonwood 


+ 





7 


+ 








Honey Locust 


+ 


< — > 


7 


< > 








Sycamore 


+ 




7 








+ 


Black Ash 


+ 


— 


7 


+ 






+ 



Literature Cited 

Braun, E. Lucy. 1950. Deciduous forests of Eastern North America. Hafner Pub. Co., 

New York. 610 p. 

Crankshaw, W. B., S. A. Qadir, and A. A. Lindsey. 1965. Edaphic controls of tree 

species in presettlement Indiana. Ecology 46 :688-698. 

Lindsey, A. A., J. D. Barton, Jr., and S. R. Miles. 1958. Field efficiencies of forest 

sampling methods. Ecology 39 :428-444. 

— , W. B. Crankshaw, and S. A. Qadir. 1965. Soil relations and distribution 
map of the vegetation of presettlement Indiana. Bot. Gaz. 126:155-168. 

— , D. V. SCHMELZ, and S. A. Nichols. 1969. Natural Areas in Indiana and 
their Preservation. Indiana Natural Areas Survey, Purdue University, Lafayette. 
606 p. 

Petty, R. O., and M. T. Jackson. 1966. Plant communities, p. 264-296. /» A. A. 
Lindsey |ed. 1 Natural Features of Indiana, Indiana Sesquicentennial Volume. Indiana 
Acad. Sci., Indianapolis. 630 p. 

Schmelz, D. V., and A. A. Lindsey. 1970. Relationships among the forest types of 
Indiana. Ecology (in press). 



A Survey of the Commercially Valuable Mussels of the 
Wabash and White Rivers of Indiana 1 

Louis A. Krumholz, Roy L. Bingham,-' and Edward R. Meyers 
University of Louisville 

Abstract 

The shells of several unionid mussels of the Wabash River system in Indiana are of 
considerable value in the manufacture of nuclei for cultured pearls in Japan. Because of 
increased exploitation of that fauna for commercial purposes, the Indiana Department of 
Natural Resources sponsored a survey to determine the status of the mussel populations 
over about 500 stream miles of the Wabash, White, and East Fork of the White rivers, 
and to gather data upon which to base recommendations for the preservation of that 
resource. 

The survey, conducted by the University of Louisville, extended through 1966 and 
1967 and included 99 collections made either with a crowfoot bar or by diving and hand- 
picking with or without auxiliary air supply. In those collections, 30 species of unionids 
were represented, but only 10 were considered important in the commercial market; 
representatives of those 10 species made up 77.1% of the total catch of the survey. The 
most abundant mussel in the commercial market as well as in the survey collections was 
the mapleleaf, Quadrula quadrula (Rafinesque). Data from the survey as well as from the 
commercial market indicated that the mussel stocks in the rivers were depleted seriously 
in 1966, mostly through the very efficient efforts of divers with auxiliary air supply. 
Those data led to the passage of Discretionary Order No. 136 by the Indiana Department 
of Natural Resources prohibiting the use of such gear in commercial musseling. 

Mussels of commercial value grow very slowly ; it requires from 4 to 5 years for a 
mapleleaf mussel to reach the legal size of 2.5 inches. The serious depletion of breeding 
stocks in 1966 probably will result in a very low yield of marketable shells at least until 
1970 and perhaps later. 

Introduction 

Freshwater bivalve mussels of the family Unionidae have been in 
existence since early Mesozoic times in the lakes and rivers of the north- 
ern temperate and subtropical areas of the world (29). At present, 
there are between 500 and 600 living species of unionids within the con- 
tinental United States (10), and of these, several are commercially valu- 
able because of the quality of the material in their shells. Although the 
soft parts of these mussels are edible, no commercial market has ever 
been established for freshwater clams comparable to that for marine 
clams. 

The shells of the freshwater mussels have become valuable at two 
different times in recent history of the United States, for two quite dif- 
ferent reasons. During the last decade of the nineteenth century, the 
pearl button industry was established at Muscatine, Iowa, by J. P. 
Boepple, a German button cutter (8). That industry flourished for about 



1 Contribution No. 126 (New Series) from the Department of Biology, University of 
Louisville, Louisville, Kentucky 40208. 

J Present Address: Department of Oceanography, Texas A&M University, College 
Station, Texas 77843. 

3 Present Address : Department of Zoology, Arizona State University, Tempe, Arizona 
85281. 

205 



206 Indiana Academy of Science 

50 years, but by the beginning of the twentieth century, the mussel beds 
in the Mississippi River began to show signs of depletion (28). 

Even though the use of mussel shells in the manufacture of pearl 
buttons essentially disappeared in 1953, new demands were put upon the 
mussel populations by the cultured pearl industry. 

Freshwater mussel shells from rivers in the United States are used 
in the manufacture of the spherical nucleus that is implanted in the 
oyster by the Japanese. Shells from the Wabash River and Tennessee 
River systems in the United States have an extraordinary translucence 
that makes them particularly valuable as nuclei. The diameter of the 
spherical nucleus prepared from mussel shells usually ranges from Vs to hi 
inch, but may be even larger on occasion. In preparing the nuclei, blemish- 
free mussel shells are cut into strips, then into squares, and then into 
cubes. The cubes are placed between rotating discs similar in shape to 
the stone of a grist mill, together with a slurry of abrasive material. The 
discs are rotated and the nuclei are worn down until they become spheri- 
cal. At this point the spheres are selected on the basis of their perfection 
and are placed within the visceral mass of the living oyster. The yield 
of cultured pearls of superior quality using such nuclei is very high. The 
time required for a cultured pearl to be formed ranges from 1 to 5 years, 
depending on the desired thickness of the nacreous covering on the 
nucleus (27). 

Persons associated with the mussel industry realized that the mussel 
supply is exhaustible and efforts were made to conserve this valuable nat- 
ural resource. In April 1967, the Indiana Department of Natural Resources 
passed Discretionary Order No. 136 which restricted the methods for 
taking mussels for commercial purposes to handpicking, short forks, 
tongs, or brail (crowfoot) bar. The use of mechanical dredges and diving 
with auxiliary air supplies was prohibited. 

Because of the very great demand for mussel shells of high quality 
in the cultured pearl industry, the beds of commercially valuable mussels 
in the Tennessee River have been virtually wiped out. Similarly, the 
mussel beds of the Wabash River system are in danger of serious deple- 
tion. In many instances, populations of desirable species of shells have 
been extirpated. However, it should be pointed out that the cultured pearl 
industry and their demands on mussel shells of the Wabash River is not 
the sole factor operating in the depletion of those shells. Pollution, 
whether it be industrial, domestic, or agricultural, has played a leading 
part in the decline of mussel populations in most Indiana streams. Also, 
the invasion of the Asiatic clam (Corbicula fluminea Muller) has become 
widespread in Indiana streams. Since the Asiatic clam does not form 
glochidia, and hence needs no fish host in its life history, it has a decided 
competitive advantage over other mussels. By late 1989, many sections 
of the Wabash and White rivers were heavily infested with Asiatic clams. 

Materials and Methods 

All together, 99 collections were made either by crowfoot bar or 
by diving and handpicking at 63 sites in the Wabash River, the White 
River, and the East Fork of the White River. 



Ecology 



207 




Figure 1. Sections of the Wabash and White rivers included in the study area. Collections 
were made at approximate 10-mile intervals from Delphi, Indiana, to the mouth of the 
Wabash River and from Tunnelton, Indiana, on the East Fork of the White River to the 
mouth of the White River. 



208 Indiana Academy of Science 

Sampling Stations 

The area under investigation extended from Delphi, at Wabash River 
Mile 331, to the Ohio River, and in the East Fork of the White River 
from Tunnelton, at White River Mile 158, to its mouth at Mt. Carmel, 
Illinois, a total of about 500 miles of stream (Fig. 1). In sampling the 
study area for distribution and relative abundance of commercially im- 
portant mussels, the river was divided into 10-mile stretches, and 1 mile 
of each stretch was marked off and sampled intensively using crowfoot 
bars either alone or in conjunction with diving and handpicking. The 
location of each sampling station by river mile was obtained from flood- 
plain charts published by the U. S. Army Corps of Engineers. Rather 
than sampling every tenth mile, a 1-mile section out of each 10 miles, suit- 
able for operation of the crowfoot bar, was selected. If no such section 
was available in any 10-mile stretch, a 1-mile section either immediately 
above or below that stretch was sampled. All together, 63 such sampling 
sites were selected and the crowfoot bar operated over those 63 miles of 
stream. 

Many of the sites were visited more than once to determine whether 
the population had changed either because of changes brought about by 
the activities of the mussel fishermen, or by some other change such as 
increased pollution. 

The Crowfoot Bar 

The crowfoot bar or brail bar (Fig. 2) has been used for collecting 
mussels since 1897 in the upper Mississippi River and its tributaries (8, 
10). Crowfoot bars are fabricated from sections of iron pipe 0.75 to 1.0 
inch in diameter and 10-20 feet in length. At intervals of 1 to 3 inches 
over the length of the bar, 3-foot lengths of seine cord or trot line are 
attached by loose half hitches. Three crowfoot hooks are attached at 
equal intervals to each strand of cord, and one end of a loose wire or rope 
bridle is attached to each end of the bar. When in use, the bar is towed 
downstream by a length of rope attached by a snap hook to an iron ring 
mounted in the center of the bridle. Crowfoot hooks are constructed from 
two 10- or 12-inch lengths of No. 9 or No. 11 steel wire bent into loops, 
then twisted together in such a manner that the ends of the loops may 
be bent at equal intervals into 4 adjacent curved hooks (Fig. 2). The 
crowfoot bar in use today is constructed essentially as it was 70 years ago. 

Crowfoot bars are carried in pairs in johnboats, each bar carried on 
two iron stanchions, mounted one on either end of each side of the boat. 
When in use, a 0.5-1.0 inch manila rope is affixed to the bridle ring and 
the whole bar comes to lie on the river bottom, perpendicular to the 
shoreline with the strands of hooks spread out behind the bar on the 
bottom. The connecting line is allowed to assume an angle of approxi- 
mately 45° to the plane of the river bottom. Then it is attached to a 
cleat mounted on the bow of the boat. 

The boat usually is carried downstream by the current with the aid 
of a sea anchor, commonly referred to as a mule, attached by cords to the 



Ecology 



209 





Figure 2. A. Johnboat with crowfoot bar on stanchions. B. Detailed view of crowfoot 
hooks. 



210 Indiana Academy of Science 

stern of the boat. Sea anchors are made of canvas, wood, cr metal about 
3 feet deep and 5 feet long. The boat may be steered by adjusting the 
cords that attach the mule to the boat. Whenever there is insufficient cur- 
rent for the mule to propel the boat, an outboard motor is used. 

In the stream bottom, mussels orient themselves with the open 
posterior portion of the valves facing upstream. As the crowfoot bar 
hooks trail down the river bottom, they may become lodged between the 
open valves of the mussel. The presence of a hook between its valves irri- 
tates the mussel and it immediately closes its shell, thereby effectively 
attaching itself to the hook. 

Mussel fishermen using crowfoot bars choose an area of clean river 
bottom known to support large populations of mussels. A bar is lowered, 
pulled along for an appropriate time, and then returned to the boat. As it 
is returned to the boat, its counterpart is lowered to the river bottom 
so that fishing is continuous. The speed of such dragging is slow, usually 
not more than 0.5 mph. While one bar is fishing, the other bar is cleaned 
of its clams and any debris and made ready for fishing again. As fishing 
proceeds, the two bars continue to be alternated over the entire stretch 
of stream. 

In the present study, we used a 14-foot johnboat carrying 2 crowfoot 
bars, each 10 feet long, and each fitted with 150 crowfoot hooks, affixed 
in groups of 3 to each of 50 three-foot lengths of seine cord (Fig. 2). To 
move about at will, the boat was powered with an 18-horsepower out- 
board motor. 

Crowfoot bars are most effective in water more than 6 feet deep and 
on bottoms of clay or sand with little large gravel. Crowfoot bars were of 
little use in water less than 3 feet deep or in areas having bottoms 
covered with dead twigs, mats of dead leaves, or other such debris. In 
such situations, the hcoks became entangled shortly after they touched 
the river bottom. 

Each collection made during this study routinely used a pair of crow- 
foot bars, and on some occasions auxiliary samples were taken with 
SCUBA (self-contained underwater breathing apparatus) gear. At each 
station, one crowfoot bar was dragged downstream for about 20 min and 
then replaced by the second bar and the process repeated over the 1-mile 
section. 

Diving and Handpicking 

Diving was performed with SCUBA gear similar to that used by 
sportsmen and by commercial mussel fishermen. A tank of compressed 
air carried on the back enables the diver to stay under water for 30-45 
min. In some instances, a portable compressor is fixed to a small raft, and 
two or more divers are supplied with air continuously. 

Mussels are placed in a bucket and taken to the surface for unloading. 
SCUBA gear is a very effective technique for sampling mussel populations 
since it can be used any time except during flood conditions, and hence 



Ecology 211 

virtually no area of the Wabash or White river systems of Indiana is 
impregnable to such diving methods. On several occasions, SCUBA gear 
was used to collect in an area that had been collected previously with 
a crowfoot bar to determine the relative efficiencies of the two methods. 

In addition to diving, many samples were collected by handpicking 
mussels in shallow water. In some areas, this was the most productive 
method of collecting. 

Reproduction 

The cultured pearl industry of today is just as dependent upon the 
continuous production of freshwater mussels as the pearl button industry 
was a half century ago. Effective practices for the conservation of any 
resource necessitates a knowledge of the complete biology of the organ- 
ism concerned. 

For this part of the study, the gonads of 369 individual mussels, 
representing 15 species, were examined (Table 1). The reproductive 
processes of Quadrula quadrula (Rafinesque) were studied intensively, 
and the extents of the breeding seasons were determined tentatively for 
that species and for Quadrula metanevra (Rafinesque), Quadirula pustu- 
loses (Lea), Obovaria, olivaria (Rafinesque), Tritogonia verrucosa 
(Barnes), Megalonaias gigantea (Barnes), Amblema costata Rafinesque, 
and Actinonaias carinata (Barnes). Principal emphasis was placed on 
those 8 commercially valuable species, and limited data were available on 
6 others as follows: Lampsilis anodontoides (Lea), 2 males, 2 females; 
Lampsilis ovata ventricosa (Barnes), 2 males, 1 female; Obliquaria 
reflexa Rafinesque, 1 male, 2 females; Elliptio crassidens (Lamarck), 2 

Table 1. Numbers, kinds, and sexes of freshwater mussels from the 

Wabash and White rivers, Indiana, used in studies of reproduction. Only 

those species for which at least 15 specimens were available for study 

are listed. Data for the other species are given in the text. 

Number of Specimens 
Species 

Actinonaias carinata 
Amblema costata 
Megalonaias gigantea 
Obovaria olivaria 
Quadrula metanevra 
Quadrula pustulosa 
Quadrula quadrula 
Tritogonia verrucosa 
Others 

Totals 185 184 369 



Male 


Female 


Total 


9 


14 


23 


11 


9 


20 


11 


6 


17 


18 


21 


39 


7 


8 


15 


24 


14 


38 


78 


89 


167 


13 


7 


20 


14 


16 


30 



212 Indiana Academy of Science 

males, 2 females; Fusconaia ebenus (Lea), 1 male, 3 females; Fusconaia 
undata (Barnes), 5 males, 5 females; Plethobasus cyphyus 
(Rafinesque), 1 male, 1 female. 

It is not practicable to attempt identification of the sex of fresh- 
water mussels without aid of a compound microscope. In this study, sex 
was determined by inserting a hypodermic needle through the incurrent 
siphon and into the gonad, withdrawing a small amount of gonadal tissue, 
and examining it under a compound microscope. The differences between 
the male and female tissues were obvious, and it is believed that unionid 
mussels can be sexed in this manner with complete confidence. So far as 
could be determined, this method for sex determination had no deleterious 
effects on any of the species studied. 

Sexually mature males and females of Q. quadrula, collected in 
March, were studied histologically to determine whether there were con- 
current areas of active and inactive gametogenesis within a single individ- 
ual. The complete foot, along with the inner and outer gills were excised, 
cut midsagittally, and the central and peripheral areas compared by 
studying stained histological sections from the various midsagittal areas. 
The gills were fixed in Bouin's fluid (15) and preserved in 70% ethanol. 
Serial sections, 6 microns thick, were prepared and stained with Heiden- 
hain's iron-haematoxylin and orange G. 

Breeding seasons were determined by correlating the degree of 
gametogenic development in the gonads of both sexes with the presence 
or absence of eggs in the water tubes or marsupia of the females. A 
female was considered gravid if the gills had become modified into a 
marsupium and contained eggs. In determining the extents of the breed- 
ing seasons, the condition of the development of the ova was considered 
the best indicator, and the next best indicator was the condition of the 
male gonad. In some species, such as O. olivaria and L. ovata ventricosa, 
the marsupium is very pronounced whereas in Q. quadrula the appear- 
ance of the marsupium is not typical even though it may contain eggs. 
In that species, the eggs are in the inner and outer gills, and the water 
tubes had to be examined at various times of the year to determine the 
extent of the breeding season. 

Relative Abundance and Distribution of Unionid Mussels 

Although no attempt was made toward a revision of unionid mussels, 
it is important here to provide a brief account of the various attempts 
to delineate the systematics of those animals. The first attempt was that 
of Stein (26) and was followed by several preliminary lists by Call (4, 5, 
6), who presented generalized geographic distributions for the aquatic 
forms known to occur within the boundaries of Indiana. In addition to 
these preliminary studies of Call, a rather superficial list of endemic 
species was published by Simpson (25). 

The first detailed study of the unionid fauna of Indiana was Call's 
Descriptive Illustrated Catalogue of the Molltisca of Indiana (7), which 
included detailed descriptions and excellent drawings of each species 



Ecology 



213 



known to exist in the state at that time. Blatchley and Daniels (3) pro- 
vided a supplement to Call's catalogue adding a number of species, mostly 
gastropods. Daniels (11, 12) provided up-to-date checklists of the mol- 
luscan fauna of Indiana. 

The most recent comprehensive study of the unionid fauna of Indiana 
is Goodrich and van der Schalie's (14) Revision of the Mollusca of Indi- 
ana. The latter paper was used in this study for species identification. 



Table 2. Distribution and abundance of unionid mussels m the 
Wabash, White, and East Fork of the White rivers of Indiana based on 
99 collections in 1966 and 1967. R, rare; — , not present; C, common; 
A, abundant. Upper Wabash River: Delphi to Terre Haute, Indiana; 
Middle Wabash River: Terre Haute to Mount Carmel, Illinois; Lower 
Wabash River: Mount Carmel to Ohio River (22). 







Wabash River 


White 


River 




Main 
Stream 


East 


Species 


Upper 


Middle 


Lower 


Fork 


Subfamily Anodontinae 












Alasmidonta marginata 


R 


— 


— 


— 


— 


Anodonta grandia 


— 


— 


R 


— 


— 


Anodontoides j ' eruaaacianua 


R 


— 


— 


— 


— 


Laamigona complanata 


C 


C 


C 


C 


C 


Laamigona compressa 


R 


— 


— 


— 


— 


Laamigona coatata 


K 


K 


— 


— 


— 


Strophitua rugoaua 


C 


— 


— 


— 


— 


Subfamily Lampsilinae 












Actinonaiaa carinata* 


A 


A 


c 


c 


c 


Cyprogenia irrorata 


— 


R 


— 


— 


— 


Lampailia anodotitoidea 


C 


C 


V 


— 


— 


Lampailia ovata ventricoaa 


c 


C 


c 


c 


(' 


Leptodea fragilia 


c 


c 


C 


V 


(' 


Leptodea laeviaaima 


— 


— 


R 


— 


— 


Obliquaria reftexa 


K 


R 


R 


R 


c 


Obovaria olivaria* 


A 


A 


A 


C 


c 


Obovaria aub rotunda 


K 


— 


— 


— 


H 


Proptera alata 


c 


C 


C 


C 


c 


Truncilla truncata 


R 


U 


R 


R 


R 


Subfamily Unioninae 












Amblema coatata* 


C 


c 


C 


C 


A 


Cyclonaiaa tuberculata 


— 


— 


— 


— 


R 


Elliptio craaaidena 


— 


— 


— 


— 


(.' 


Fuaconaia ebenua* 


R 


R 


R 


c 


C 


Fuaconaia undata* 


R 


R 


— 


R 


C 


Megalonaiaa gigantea* 


R 


C 


— 


R 


(' 


Plethobaaua cyphyua 


R 


— 


— 


— 


K 


Pleurobema cordatum 


— 


— 


— 


— 


R 


Quadrtda metanevra* 


R 


R 


R 


K 


R 


Quadrula puatuloaa* 


A 


A 


A 


A 


A 


Quadrula quadrula* 


A 


A 


A 


A 


A 


Tritogonia verrucoaa* 


C 


C 


C 


— 


— 



The 10 species of greatest commercial value. 



214 Indiana Academy of Science 

The distribution, relative abundance, and locations of collections are 
shown in Table 2 (22). The species are arranged in alphabetic order under 
three subfamilies. Many species taken were of little or no commercial 
value, but their occurrence is listed. Particular emphasis is placed on 10 
species of known commercial value (Table 2). 

All species collected in this study were listed by Goodrich and van der 
Schalie (14) and most were described and illustrated by Call (7). How- 
ever, some rare and less abundant species listed by those authors, such as 
Amblema peruviana (Lamarck), were not encountered here, and some, 
once common in Indiana rivers, such as Elliptio dilatatus (Rafinesque) 
and Quadrula cylindrica (Say), were in evidence only through isolated 
dead shells. Those two species apparently have been extirpated from the 
study area within the last two decades. Other species, such as Pleurobema 
cordatum (Rafinesque), Cyclonaias tuberculata (Rafinesque), Cyprogenia 
irrorata (Lea), and Obovaria sub7'otunda (Rafinesque) are much less 
abundant and have much more restricted patterns of distribution than 
those recorded by Goodrich and van der Schalie (14). No species 
encountered in this study exhibited wider ranges of distribution since 
the earlier studies, and most species appeared much less abundant. Only 
two species, Q. pustulosa and Q. quadrula, were as abundant or more 
abundant than indicated by earlier authors. 

Such information indicates a definite trend toward restriction in 
range and reduction in absolute abundance of most unionid mussels in the 
Wabash and White rivers. Although some species may have been extir- 
pated from the areas under investigation there may still be limited pop- 
ulations in other parts of the drainage system. If there are, hopefully 
the populations in the mainstreams of the Wabash and White rivers may 
be re-established when conditions permit. 

This pattern of reduction and elimination of unionid mussel popula- 
tions may be attributed to two principal factors: 1) the disturbance and 
destruction of mussel beds through overexploitation for commercial pur- 
poses; and 2) deterioration of the environment as suitable habitat for 
mussels through increasing burdens of pollution. 

The devastating effects of industrial and organic pollutants on unionid 
mussels were first pointed out by Ortmann (23) in his report on the effects 
of such pollution in the Allegheny, Monongahela, and Ohio rivers in 
western Pennsylvania. Ten years later, Forbes and Richardson (13) noted 
a direct correlation between increasing levels of pollution and decreasing 
ranges and numbers of mollusks in the Illinois River. More appropriate 
to the present work was the study by Baker (1) in which he reported in 
detail the elimination of all mussels from certain areas of the Big 
Vermillion River, a tributary to the Wabash River, by heavy loads of 
municipal organic wastes. More recently, Wurtz (30) stated that unionid 
mussels were quite intolerant to pollution of any kind and reported 
unequivocally that freshwater mussels disappear quickly from streams 
carrying moderately heavy burdens of pollutants. 

Polluted streams exhibit remarkable patterns of recovery down- 
stream from sources of pollution or following the abatement of pollution 



Ecology 215 

(1, 16, 18, 19). Perhaps the first detailed description of the biological 
recovery of a river system downstream from a source of pollution was 
that of Kolkwitz and Marsson (17) who reported on the changes in the 
species composition of aquatic communities, and pointed out that the 
abundance and diversity of organisms increased as the effects of pollution 
diminished. 

Disruption of the stream bottom by mechanical means may be just 
as devastating to a mussel population as pollution. One of the methods 
for collecting mussels that is most destructive of the streambed is 
mechanical or hydraulic dredging. In the operation of such a dredge, about 
the uppermost 2 feet of stream bottom is lifted out of place, carried to 
a barge where the mussels are removed, and then dumped back into 
the river. In this operation, the stream bottom, with all its biotic com- 
munities, is completely discommoded. Any benthic organisms that survive 
must become re-established rather quickly or those segments of the 
stream ecosystem that depend on benthos for their livelihood will not 
survive. 

Tongs, crowfoot bars, and other such equipment disturb the stream- 
bed to a limited extent, but they usually are used only in localities where 
they are known to be effective. At most, those tools disrupt small areas 
of bottom to a depth of a few inches, instead of upheaval of the entire 
streambed. 

Perhaps the method least destructive to the stream bottom is hand- 
picking. Here, the musseler simply walks along the stream bottom or 
swims over it and picks up whichever mussels he desires. 

Numbers and Kinds of Mussels Collected 

Data for the 20 species of mussels taken in at least 6 different col- 
lections are arranged according to method of collection in Table 3. 

From the data in Table 3, it is obvious that about % (34.0%) of the 
total catch was made up of mapleleaf (Quadrula quadrula) , one of the 
most highly sought-after commercial mussels for the cultured pearl 
industry. The mapleleaf was followed in order by the sandshell (Obovaria 
olivaria) which contributed 10.2% and the pimpleback (Q. pustulosa) 
with 9.0%. Thus, these three species made up 53.2% of the total catch. 
The remaining 7 of the 10 commercially important species listed earlier 
comprised another 23.9% of the catch, and all together the 10 species 
made up more than % (77.1%) of all mussels taken during the study. 

Of the 20 species listed in Table 3, several have shells that are too 
thin and fragile or are too highly colored to be of value as nuclei for 
cultured pearls. Others were taken in such small numbers that they could 
not be considered important in the commercial market. Only 15 of the 20 
species were represented by more than 25 specimens. 

The white heelsplitter, Lasmigona complanata (Barnes), is used 
sparingly in the cultured pearl industry, but has some commercial value 



216 Indiana Academy of Science 

Table 3. Numbers of individuals of the 20 species of the unionid mus- 
sels represented in six or more collections taken by crowfoot bar and by 
handpickifig, together with the numbers of collections in which they 
occurred, from the Wabash and White rivers during 1966 and 1967. The 
averages are for the numbers of individuals per collection. 









All 


Collections 








Crowfoot Bar 


Handpicking 




Total 


Species 


No. 


Coll. 


No. 




Coll. 


No. 


Coll. 


Lasmigona complanata 


24 


5 


40 




11 


04 


10 


Lasmigona costata 


1 


1 


5 




5 








Others* 


12 


11 


3 




2 


15 


13 


Actinonaias carinata 


XX 


21 


21 




6 


109 


27 


Lampsilis anodontoides 


36 





1 




1 


37 


7 


Lampsilis ovata ventricosa 


34 


20 


54 




15 


XX 


35 


Leptodea fragilis 


54 


22 


3 5 




15 


X9 


37 


Obliquaria reflexa 


10 





3 




2 


13 


8 


Obovaria olivaria 


162 


35 


32 




10 


194 


45 


Obovaria subrotunda 


9 


4 


2 




2 


11 


6 


Propter a alata 


9 


7 


14 




6 


23 


13 


Truncilla truncata 


4 


3 







6 


10 


9 


Others* 


— 


— 


2 




2 


2 


2 


Amblema costata 


41 


If) 


08 




10 


104 


25 


Elliptio crassidens 


35 


3 


22 




3 


57 


6 


Fusconaia ebenus 


4 


4 


51 




8 


55 


12 


Fusconaia undata 


S 


6 


13 




7 


21 


13 


Megalonaias gigantea 


30 


X 


08 




it 


93 


17 


Quadrula metanevra 


35 


12 


12 







47 


18 


Quadrula pustulosa 


XX 


25 


84 




14 


172 


39 


Quadrula quadrula 


359 


39 


288 




17 


047 


56 


Tritogonia verrucosa 


27 


12 


— 




— 


27 


12 


Others* 


6 


2 


10 




12 


22 


14 


Totals 


1076 


02 


830 




37 


1906 


99 


Average 


17.4 




22.4 






19.3 



* The total number of specimens for each of the species included in the category of 
"Others" are: Alasmidonta marginata (Say), 2 specimens; Anodonta grandis Say, 2; 
Anodontoides ferussacianus (Lea), 2; Lasmigona compressa (Lea), 4; Strophitus rugosus 
(Swainson), 5; Cyprogenia irrorata, 1; Leptodea laevissima (Lea), 1; Cyclonaias 
tubcrculata, 3 ; Plethobasus cyphyus, 8 ; Pleurobema cordatum, 4 ; and 7 unidentified 
individuals. 



because of its relative abundance. The shell is fairly heavy and the 
nacre is creamy white. The 64 specimens taken made up 3.4% of the total 
catch; however, most of those shells were collected by hand in the East 
Fork of the White River in water less than 6 feet deep. 

Lasmigona costata (Rafinesque), the fluted shell, was rare in the 
collections and is of very little commercial value. 

The mucket, Actinonaias carinata, was fourth in numerical abundance 
in our collections and made up 5.7% of the total catch. The shell of this 
species is commercially important and is of high quality in the manufac- 
ture of nuclei for cultured pearls. 



Ecology 217 

The yellow sandshell, Lampsilis anodontoides, is not of commercial 
value because usually it does not have enough thick portion of the shell 
to warrant the expense of cutting it for nuclei. 

Although the pocketbook, Lampsilis ovata ventricosa, and the 
floater, Leptodea fragilis (Rafinesque), comprised 4.5 and 4.7%, respec- 
tively, of the total catch, neither is important because of the quality of 
the shell. The pocketbook usually is pale green to brown in color and the 
shell of the floater is so fragile that it can easily be crushed between the 
fingers of one hand. 

With the exception of Obovaria olivaria, all other members of the 
Lampsilinae were taken in such small numbers that they are not of 
commercial importance. The sandshell is one of the most highly sought- 
after shells and is relatively abundant, particularly in the Wabash River, 
where it consistently makes up a significant part of the commercial 
market. 

The threeridge, Amblema costata, ranked fifth in the number of 
specimens collected in this study and made up 5.5% of the total catch. 
This mussel was common throughout the study area and was particularly 
abundant in the upper reaches of the East Fork of the White River. In 
that area, in contributes consistently to the commercial market. 

Elliptio crassidens, the elephant's ear, is of no commercial value in 
the cultured pearl industry because of the purple color of its nacre. 

Although the two representatives of the genus Fusconaia, F. ebenus 
and the pigtoe, F. undata, contributed only 2.9 and 1.1%, respectively, to 
the total catch in this study, they are highly desirable as shells in the 
cultured pearl industry. Fusconaia ebenus is particularly desirable and 
actually makes up a significant portion of the commercial market. 

The largest freshwater mussel taken in the study was the wash- 
board, Megalonaias gigantea. In this study it contributed 4.9% of the 
total catch and was particularly important in the East Fork of the White 
River. The washboard has a relatively high commercial value because of 
its size and thickness of the shell, but the quality of the nacre is not as 
desirable as that of some other shells. 

The most important genus of freshwater unionids, so far as the 
contribution of its representatives to the cultured pearl industry is con- 
cerned, is Quadrula. In the Wabash River system, three members of the 
genus, the monkeyface, Q. metanevra, the pimpleback, Q. pustulosa, and 
the mapleleaf, Q. quadrula, contributed 45.4% to the total number of 
mussels taken during this study. These 3 shells, together with Obovaria 
olivaria and Fiisconia ebenus are the 5 shells of greatest commercial value 
for pearl nuclei and they made up 56.7% of the total catch in this study. 
When these 5 shells are considered together with the other 5 of the 10 
most important commercial species listed earlier, the contribution to the 
total number collected in this study was 77.1%. Thus, the 10 most sought 
after species are in aggregate the 10 most abundant species. 



218 Indiana Academy of Science 

The only other species of commercial importance is the pistolgrip, 
Tritogonia verrucosa, which has a distribution limited almost entirely to 
the Wabash River upstream from Vincennes, Indiana. Although this shell 
contributed only 1.47c to the overall catch, in the area above Vincennes in 
1966, it contributed 3.7%. 

Comparisons of Catches in Different Localities 

In the present study, it was assumed that the sites for collections by 
crowfoot bar and by handpicking were distributed randomly over the 
river systems investigated and that those methods of collection were 
non-selective in the kinds of mussels taken. Thus, the number and kinds 
of mussels in the collections should be a good indication of their distribu- 
tion throughout the study area (Table 3). As is true of most animals in 
the world, the distribution of unionid mussels in the Wabash and White 
river systems was not uniform. Rather, some of the species were not 
taken in the Wabash River and others were not taken in the mainstream 
or the East Fork of the White River (Tables 2 and 3). 

Among the 20 species mentioned in Table 3, only one, Elliptio crassi- 
dens, was not taken in the Wabash River at any time, and another, 
Obliquuria reflexa, was represented by a single female specimen taken 
on a crowfoot bar near Cayuga, Indiana, 24 August 1967. Conversely, 
E. crassidens was represented by 57 specimens collected by crowfoot bar 
and handpicking from the East Fork of the White River, and 12 speci- 
mens of O. reflexa were collected from the mainstream and East Fork 
of the White River in 1967. In that part of the White River system 
under study, 5 of the 20 species were not present in the collections from 
the main stem and 3 were not taken in the East Fork of the White 
River. However, of all of those species, only Tritogonia verrucosa is of 
commercial value. 

It is also obvious from the data in Table 3 that some species are 
relatively much more abundant in one area than in another, whereas 
others may be fairly evenly distributed. Several species, Actinonaias 
carinata, Obovaria olivaria, Quadrula metanevra, Q. quadrula, and Tri- 
togonia verrucosa apparently are much more common in the Wabash 
River than in the White River. By the same token, Amblcma costata, 
Fusconaia ebenus, and Megalonaias gigantea appear to be more common 
in the White River. Only one of the 10 most commercially valuable spe- 
cies, Quadrula pustulosa, seems fairly evenly distributed over the study 
area. 

Another datum that may give an indication of the distribution of 
mussels in the rivers investigated is the number of collections in which 
each kind of mussel occurred. Only four species, Lampsilis ovata ventri- 
cosa, Obovaria olivaria, Quadrula metanevra, and Q. quadrula, were 
taken by crowfoot bar and by handpicking from the Wabash River in 
both 1966 and 1967, and from the main stem and East Fork of the 
White River (Table 3). One other species, Propter® alata (Say), was 
taken by handpicking from each area and by crowfoot bar from the 



Ecology 219 

main stem and East Fork of the White River and from the Wabash 
River in 1966, but not in 1967. Similarily, Truncilla truncata Rafinesque, 
although rare, was collected in each of the areas. 

For the entire study, 19 of the 20 species were represented in the 
61 collections from the Wabash River, 17 were represented in the 27 
collections from the East Fork of the White River, and 15 were taken 
in the 11 collections from the White River proper. Here, it should be 
pointed out that the 61 collections from the Wabash River were taken 
over a distance of about 330 miles, the 27 from the East Fork from 
about 110 miles of stream, and the 11 from the main stem of the White 
River from about 50 miles of stream, so that the intensity of sampling 
was about the same in each area. 

In considering the occurrence of only the 10 species of greatest 
commercial value in all collections made during this study, only 1, 
Q. quadrula, was represented in more than half those collections. Four 
others, O. olivaria, Q. pustulosa, A. carinata, and A. costata, were repre- 
sented in more than %, and each of the remaining 5 was represented 
in from 10 to 20% of all collections. In the Wabash River alone, 2 
species, Q. quadrula and O. olivaria were represented in more than half 
the 61 collections, 5 were represented in from 20 to 35%, and 3 were 
present in less than 10% of the collections. In the main stem of the 
White River, where only 11 collections were made, no species was taken 
in more than four collections, and one, T. verrucosa, was not repre- 
sented. In the East Fork of the White River, 2 species were present in 
more than half the 27 collections, 2 others were represented in more 
than %, 3 others in more than hi, 2 others in from 15 to 20%, and 1, 
T. verrucosa, was not represented. From these data it is apparent that 
Q. quadrula and O. olivaria are ubiquitous and that the other eight 
species, with the exception of T. verrucosa, are fairly widespread 
throughout the Wabash River system. 

Comparisons of Catches in 1966 and 1967 

During the course of this study, 10 one-mile sections of the Wabash 
River between Delphi (Mile 331) and Terre Haute (Mile 220), Indiana 
were sampled with a crowfoot bar during June or July 1966, and again 
in August 1967. All collections were made with the same equipment 
operated by the same personnel, and the data are believed comparable. 

In 1966, 297 specimens referable to 21 species were taken from the 
10 collecting sites (Table 4), whereas in 1967, only 56 mussels referable 
to 11 species were collected. This amounts to a reduction in numbers 
of specimens of 81% and a reduction in the diversity of species of 
48%. The only instance in which more individuals were collected in 
a single station in 1967 than in 1966, was at Wabash River Mile 253-252, 
where 24 individuals were taken the second year as compared with 19 
the first. Mussels referable to 9 species were taken during the 2-year 
period, of which 7 were collected in 1966 and 6 in 1967. The numbers 
of individuals of each species taken in 1966 were: Lasmigona complanata, 
8; Lampsilis anodontoides, 2; Proptera alata, 2; Obovaria olivaria, 2; 



220 Indiana Academy of Science 

Table 4. Collections of mussels from each of 10 1-mile sectio?is taken 
with a crowfoot bar in 1966 and 1967, showing the numbers of mus- 
sels a?id numbers of species taken in each collection. 

1966 1967 



50 


12 


22 


9 


42 


7 


2 


1 


45 


8 


1 


1 


19 


5 


2 


1 


7 


2 


■ — 


— 


27 


9 


2 


2 


39 


7 


— 


— 


19 


7 


24 


(5 


3 


2 


— 


— 


46 


10 


3 


2 



No. of No. of No. of No. of 

River Mile Specimens Species Specimens Species 

320-319 
311-310 
294-293 
285-284 
281-280 
271-270 
259-258 
253-252 
239-238 
230-229 

Totals 297 21 56 11 



Quadrula pustulosa, 1; Q. quadrula, 3; and Tritogonia verrucosa, 1. In 
1967, the numbers were: Actinonaias carinata, 4; Obliquaria reflexa, 1; 

0. olivaria, 6; Q. pustulosa, 1; Q. quadrula, 11; and T. verrucosa, 

1. An analysis of these catches indicates that the mussels taken 
in 1967 had a much higher commercial value than those taken in 1966; 
of the 10 commercially important mussels listed earlier, there were 23 
taken in 1967 and only 7 taken in 1966. The reasons for these differences 
in the locality under discussion are not readily apparent. In the light 
of the data from the other nine localities where collections were made 
both years, it seems most likely that the chance of a crowfoot hook 
coming in contact with a particular mussel was very important in the 
makeup of that part of the study. Still, the consistently low catches 
in 1967 in the other 9 localities indicates that the overall populations 
of mussels in the upper Wabash River had declined rather spectacularly. 

The numbers of individuals of each species taken each year in all 
10 collections are listed in Table 5. Of the 22 species listed, 21 were 
taken in 1966 and 11 were taken in 1967. A single specimen represented 
the total catch for each of 8 species in 1966, and there were 5 species 
represented by a single individual in 1967. Although 9 of the 10 com- 
mercially desirable species listed earlier were taken in 1966 com- 
pared with 6 such species in 1967, the percentages of the total catch 
made up by those species for the 2 years was about the same. However, 
the actual drop in the numbers of such highly desirable species as A. 
carinata and Q. quadrula cannot be regarded lightly. This is even more 
impressive when it is considered that all of the specimens of Q. quadrula 
collected in 1967 were taken in the single collection at Mile 253-252. 



Ecology 



221 



Table 5. Numbers of each species of mussels collected by crowfoot bar 

from the same 10 one-mile sections of the Wabash River in 1966 and 

1967 (see Table U for location of collecting sites). 



Species 



No. Taken 


No. Taken 


in 1966 


in 1967 


1 





1 


__ 


8 


2 


1 


1 


5 


— 


43 


9 


2 


— 


9 


4 


16 


1 


— 


1 


44 


15 


3 


— 


5 


— 


1 


— 


1 


1 


1 


— 


1 


— 


1 


— 


15 


_ 


24 


4 


110 


17 


5 


1 



Alasmidonta marginata 
Anodontoides terussacianus 
Lasmigona complanata 
Lasmigona compressa 
Strophitus rugosus 
Actinonaias carinata 
Lampsilis anodontoides 
Lampsilis ovata ventricosa 
Leptodea f?'agilis 
Obliquaria reflexa 
Obovaria olivaria 
Obovaria subrotunda 
Proptera alata 
Truncilla truncata 
Amblema costata 
Fusconaia ebenus 
Fusconaia undata 
Plethobasus cyphyus 
Quadrula metanevra 
Quadrula pustulosa 
Quadrula quadrula 
Tritogonia verrucosa 



Totals 



297 



5(5 



Only two species, O. olivaria and Q. quadrula, were represented in 
each of the 10 collections in 1966, whereas O. olivaria was represented 
in 5 of the 1967 collections and Q. quadrula was represented in 4. Seven 
other species were represented in 4 or more collections in 1966, but only 
2 species other than those mentioned above were represented in more 
than a single sample. The lack of representation in more than a single 
collection in 1967 lends credence to the decline in species diversity as 
well as actual abundance of mussels during the period under considera- 
tion. 

In comparing the catches in 1966 with those in 1967, it is important 
to compare them with changes in the numbers of shells sold for use in 
the cultured pearl industry. In 1965, 2000 tons of Indiana shells were sold 
for use in Japan; in 1966, that number was 4200 tons; in 1967 it was 
1080 tons; and in 1968 the total number probably did not exceed 250 
tons. Using these data for comparison, the percentage drop between 



222 Indiana Academy of Science 

1966 and 1967 was essentially the same as for our samples taken in 
the survey and there was another substantial drop between 1967 and 
1968 for which we have no data. 

It is important here to note, however, that in 1969 the com- 
mercial musselers have reported very large numbers of small mussels 
in the Wabash River, and from the estimated size of these young ani- 
mals they must be in their second or third year of life. On examination, 
it was found that most of these small mussels were Asiatic clams. 

Efficiency of Different Kinds of Gear 

Only two methods for collecting mussels were used in this study, 
the crowfoot bar and handpicking with or without SCUBA gear. When 
a crowfoot bar is dragged over an area, only those mussels at the 
surface of the stream bottom with their valves open are capable of 
being caught. With diving or handpicking, each mussel seen or touched 
by the musseler is capable of being captured. Thus, it is obvious that 
the opportunities for collecting mussels by handpicking, including div- 
ing with or without auxiliary air supply, are much greater than with a 
crowfoot bar. 

On 13 September 1966, in the Wabash River near Americus, Indiana, 
we roped off an area of stream bottom 10 feet wide and 170 feet long. 
That area was then dragged with a crowfoot bar, and a total of 8 
mussels representing 3 species and weighing 8.4 lbs. was collected. 
Following the study with the crowfoot bar, 2 divers equipped with 
SCUBA gear covered the same marked-off area and picked up 220 
mussels of legal size including 201 specimens of commercial value as 
follows: 8 Actinonaias carinata, 3 Obovaria olivaria, 13 Amblema costata, 
7 Quadrula metayievra , 6 Q. pustulosa, and 164 Q. quadrula that weighed 
a total of 138.8 lbs. The other 19 specimens represented 4 species of 
no commercial value and weighed 8.7 lbs. 

On the basis of the data in Table 3, where the average catch per 
drag of the crowfoot bar over a 1-mile section of river yielded 17.4 
mussels and the average catch per collection by handpicking was 22.4 
mussels, the difference does not seem great. It should be pointed out 
here that each collection with a crowfoot bar required 2 men more than 
2 hours of work whereas the average time for the collections by hand- 
picking was 2 men for less than 30 min. Thus, it is apparent that the 
latter method is more than four times as efficient on the basis of time 
alone. 

Handpicking with the aid of an auxiliary air supply is so efficient 
that an entire mussel bed can be wiped out in a few hours. We know 
of 1 instance in which 3 men with auxiliary air supply collected more 
than 3000 lbs. of commercially valuable mussels from a single bed in 
less than 2 days. Of course, they collected only mussels of legal size 
and left the smaller individuals behind. That act did assure the survival 
of the population, but the growth rate of most thick-shelled mussels is 
slow; it requires about 4 or 5 years for a mapleleaf or threeridge mussel 



Ecology 223 

to attain the legal size of 2.5 inches. Still, by removing all the larger 
individuals, the reproductive capacity of the population is seriously im- 
paired, and it may take another 4 or 5 years or even longer for the 
population to recover its reproductive potential. 

Reproduction 

The cell lineage in unionid mussels has been adequately described 
and illustrated by Lillie (20), and atypical spermatogenesis in mollusks 
has been reported by Coe and Turner (9) and Loosanoff (21). No at- 
tempt has been made in the present study to describe the details of 
meiosis in either the male or female mussel, but atypical spermatogenesis 
was demonstrated in Quadrula quadrula (2). 

Atypical sperm cells develop from the same type of spermatogenic 
cells as normal sperm cells, but the atypical cells undergo several mitot- 
ic divisions while still surrounded by the original cytoplasm within the 
original cell membrane. The number of cells undergoing atypical sperma- 
togenesis usually are much less numerous than those undergoing normal 
spermatogenesis, but there are occasions when the atypical cells out- 
number the typical ones. 

Spermatogenesis is a continuous process in Q. quadrula, the most 
active period being in the seasons immediately before and during spawn- 
ing. In March and April, before the breeding season commences, 
spermatogenesis is active and continuous until late June. 

In the ovary of Q. quadrula, the primordial wall becomes differen- 
tiated and the ovum is formed following the usual maturation divisions. 
The ovum moves from the lumen of the ovary through ducts to the 
suprabranchial chamber and down into the water tubes of the inner 
and outer gills. The spermatozoa move through ducts from the lumina 
of the acini to the suprabranchial chambers and cloacal chamber and out 
the excurrent siphon into the water. The sperm are drawn into the in- 
current siphon of the female by respiratory action, and are carried 
through the mantle cavity to the gills where they enter the water tubes 
through the ostia. 

Syngamy and cleavage take place in the water tubes and the gills 
become modified into a marsupium to accommodate the developing em- 
bryos. Q. quadrula is larviparous, the homolecithal ova are incubated 
within the interlamellar spaces of the modified gills. In about 2 to 4 
weeks, glochidia are formed and discharged from the water tubes into 
the suprabranchial chambers, through the cloacal chamber, and out the 
excurrent siphon. Since glochidia are obligate temporary parasites of 
fishes, if they do not find a host fish within a few days they perish. 

If there is a large or dense adult population of mussels or if 
there is a sparse population of suitable fishes to serve as hosts for 
the developing glochidia, the survival of young individuals is low. If, 
however, there is an adequate breeding population of mussels but they 
are scattered rather sparsely over a large area, the survival of glochidia 



224 Indiana Academy of Science 

will be high if enough suitable host fishes are present. Thus, if the legal- 
sized mussels are harvested after they have reproduced, any glochidia 
that survive the host-parasite relationship with fishes, will have a much 
better rate of survival but they will not be able to add to the harvest- 
able crop for about 5 years. 

According to Pennak (24), all members of the Subfamily Unioninae 
are short-term breeders and are gravid sometime between April and 
August; whereas, all members of the Subfamilies Anodontinae and 
Lampsilinae are long-term breeders among which fertilization takes 
place in mid or late summer and the embryos are carried until the 
next spring. 

Among the species reported here (Table 1), only Actinonaias car- 
inata and Obovaria olivaria belong to subfamilies other than the Union- 
inae. Histological studies of the gonads of 21 specimens of Actinonaias 
carinata indicated that the ovary was not in breeding condition until 
late July and continued in that condition until mid-October. Spermato- 
genesis became active during the late spring, but no mature sperm were 
encountered until late July. Thus, based on the relatively few specimens 
studied, it is apparent that fertilization could not have taken place 
much before August and could continue on through early October. No 
observations were made on the time of release of glochidia in this spe- 
cies, but the evidence at hand indicates that it is a long-term breeder. 

In O. olivaria, the gonads of 39 specimens were examined in detail. 
Uncleaved but mature ova were observed in the ovaries of specimens 
sacrificed in August, and at that time the ovary appeared to be in 
typical breeding condition. Mature spermatozoa were present in the 
lumina in late July, indicating that active spermatogenesis was in 
progress. These conditions in the gonads of each sex persisted through 
September, typical for the long-term breeder. No observations were 
made on the release of glochidia. 

All other species for which there were adequate samples for de- 
tailed study belong to the Unioninae and would be expected to be typi- 
cally short-term breeders. In Amblema costata, the breeding seasons 
extend from early May to early July. The breeding season for Mega- 
lonaias gigantea extends from early June through August. The breed- 
ing seasons for the three species of Quadrula studied here extend 
through May, June, and July. 

Of the 19 specimens of Tritogonia verrucosa reported here, there 
were 7 females and 12 males. Examination of the testes from specimens 
collected in July indicated that they were in prebreeding condition and 
specimens collected in November and early May contained mature sperm. 
The ovary of a specimen taken in November was in typical breeding 
condition. The walls of the primordium were thin, the vitelline mem- 
branes were well developed, amphinuclei were present, and the ova were 
crowded in the lumen. Ovaries from females collected in June and 
July were characteristically in prebreeding condition. Thus, it appears 
that T. verrucosa, although a member of the Unioninae, is a long-term 



Ecology 225 

breeder. However, more data will have to be gathered in order to estab- 
lish the precise limits of the breeding season, and observations must be 
made on the time of release of glochidia. 

The only evidence of sexual dimorphism among the mussels studied 
was in individuals of T. verrucosa. In that species, the valves of the 
females have long posterior extensions not present in the valves of 
the males. There is no evidence, based on this study, of any relation- 
ship between degree of obesity and maleness or femaleness. 

Acknowledgements 

This study was supported by the Indiana Department of Natural 
Resources, Division of Fish and Game, under Sub-Project No. 4-10-R, in 
cooperation with the U. S. Department of the Interior, Bureau of Com- 
mercial Fisheries. We are grateful to Messrs. Woodrow M. Fleming and 
R. Eugene Bass of the Indiana Division of Fish and Game; to Darrell 
Christensen and William W. Oakes for field assistance; to Joe K. Neel 
for initiating the project; and to E. Nelson Cohen, Terre Haute, Indiana, 
for providing crowfoot bars and information on commercial aspects of 
mussel shells in the Japanese cultured pearl industry. 

Literature Cited 

1. Baker, F. C. 1922. The molluscan fauna of the Big Vermillion River, Illinois, with 
special reference to its modification as a result of pollution by sewage and manu- 
facturing wastes. 111. Biol. Monogr. 7:105-224. 

2. Bingham, R. L. 1968. Reproductive seasons of eight freshwater mussels from the 
Wabash, White, and East Fork of the White rivers of Indiana. Unpublished M.S. 
Thesis. Univ. Louisville, Louisville, Ky. 102 p. 

3. Blatchley, W. S., and L. E. Daniels. 1902. On some mussels known to occur in 
Indiana. Annu. Rep. Ind. Geol. Surv. 26 :557-628. 

4. Call, R. E. 1892. A contribution to a knowledge of Indiana Mollusca. Proc. Indiana 
Acad. Sci. 3 :140-160. 

5. . 1896a. Second contribution to a knowledge of Indiana Mollusca. Proc. 

Indiana Acad. Sci. 6:135-146. 



1896b. The hydrographic basins of Indiana and their molluscan fauna. 



Proc. Indiana Acad. Sci. 6 :247-258. 



7. . 1900. A descriptive illustrated catalogue of the Mollusca of Indiana. 

Annu. Rept. Geol. Surv. Ind. 24 :335-535. 

8. Carlander, H. B. 1954. A history of fish and fishing in the upper Mississippi River. 
Upper Miss. R. Conserv. Comm. 96 p. 

9. Coe, W. R., and H. J. Turner, Jr. 1938. Development of the gonad and gametes in 
the soft-shell clam, (Mya arenaria) . J. Morph. 62:91-111. 

10. Coker, R. E. 1921. Freshwater mussels and mussel industries of the United States. 
Bull. U. S. Bur. Fish. 36:13-89. 

11. Daniels, L. E. 1903. A check-list of the Indiana Mollusca, with localities. Annu. 
Rep. Ind. Geol. Surv. 26 :629-652. 

12. . 1914. A supplemental check-list of Indiana Mollusca, with localities and 

notes. Annu. Rep. Ind. Dept. Geol. Natur. Resources 39:318-326. 



226 Indiana Academy of Science 

13. Forbes, S. A., and R. E. Richardson. 1919. Some recent changes in Illinois biology. 
Bull. 111. State Lab. Natur. Hist. 13:137-156. 

14. Goodrich, C, and H. van der Schalie. 1944. A revision of the Mollusca of Indiana. 
Amer. Midland Natur. 32(2) :257-326. 

15. Guyer, M. F. 1953. Animal micrology. 5th ed. The University of Chicago Press, 
Chicago. 327 p. 

16. Hynes, H. B. N. 1960. The biology of polluted waters. Liverpool Univ. Press, Liver- 
pool, England. 202 p. 

17. Kolkwitz, R., and M. Marsson. 1909. okologie der tierische Saprobien. Beitrage zur 
Lehre von der biologische Gewasserbeurteilung. Int. Rev. ges. Hydrobiol. 2 : 126-152. 

18. Krumholz, L. A. 1946. Repopulation of the West Fork. Outdoor Indiana 13 ((3) :12. 



19. - — , and W. L. Minckley. 1964. Changes in the fish population in the upper 
Ohio River following temporary pollution abatement. Trans. Amer. Fish. Soc. 93 

(l):l-5. 

20. LlLLlE, F. R. 1895. The embryology of the Unionidae. J. Morph. 10(1) : 1-100. 

21. Loosanoff, V. L. Reproductive cycle in Cyprina islandica. Biol. Bull. 104(2) :146-155. 

22. Meyer, E. R. 1968. The distribution and abundance of freshwater mussels of the 
family Unionidae (Pelecypoda) of the Wabash, White, and East Fork of the White 
rivers of Indiana. Unpublished M.S. Thesis. Univ. Louisville, Louisville, Ky. 68 p. 

23. Ortmann, A. E. 1909. The destruction of the freshwater fauna in western Pennsyl- 
vania. Proc. Amer. Phil. Soc. 48:90-110. 

24. Pennak, R. W. 1953. Freshwater invertebrates of the United States. Ronald Press. 

New York. 769 p. 

25. Simpson, C. T. 1900. Unionidae of Indiana. Nautilus 14 :95-96. 

26. Stein, F. 1881. The molluscous fauna of Indiana. Annu. Rep. Ind. Geol. Surv. 16 :451- 
467. 

27. Stephens, W. M. 1963. Man-made pearls. Sea Frontiers 9 (5) :299-308. 

28. van der Schalie, H. 1938. Hitch-hiking mussels and pearl buttons. Mich. Conserv. 
7:4-5, 11. 

29. Walker, B. 1917. The method of evolution in the Unionidae. Occas. Pap. Univ. Mich. 
Mus. Zool. 45:1-10. 

30. Wurtz, C. B. 1956. Freshwater mollusks and stream pollution. Nautilus 69 (3) :97-102. 



ENTOMOLOGY 

Chairman: Jack R. Munsee, Indiana State University 
John W. Hart, Earlham College, was elected Chairman for 1970 

ABSTRACTS 
The Use cf Heartbeat as a Potential Screening Technique for Insect 
Pathogens. Mildred G. Ware and Harold L. Zimmack, Ball State 
University. — The purposes of this study were: 1) to develop a rapid 
and accurate screening technique which will determine, by observing 
changes in heartbeat, whether or not a specific species of bacteria is 
pathogenic to a specific insect and 2) to determine criteria which would 
make it possible to apply this technique to other insects. 

On the basis of these data, it is possible that the heartbeat of the 
European corn borer, Ostrinia nubilalis (Hubner), can be used to detect 
bacterial pathogens. 

Results obtained indicate that the non-pathogenic species of bac- 
teria, Escherichia coli (Migulo), increased the mean post-ingestion heart 
rate 4.36 heartbeats/minute; whereas mean post-ingestion heart rates in 
the larvae ingesting the pathogenic species of bacteria, Bacillus thurin- 
giensis Berliner, Serratia marcescens Bizi and Bacillus subtilis Cohn, 
decreased 19.44 to 24.21 heartbeats/minute. 

The investigators believe that age had a definite influence on the 
heart rates and the susceptibility of the larvae to pathogenicity. In the 
test group using B. thuringiensis, reported to be the most vigorous of 
the experimental pathogens, all late fifth instar larvae were used. The 
mean pre- and post-ingestion heart rates for this group were 69.68 
heartbeats /minute and 50.24 heartbeats /minute, respectively. These fig- 
ures are significantly lower than the mean pre- and post-ingestion heart 
rates of early fifth instar larvae used to test the effects of the moderate 
pathogenic species, S. marcescens, and the controversial pathogen, B. 
subtilis. The mean figures in the S. marcescens test group were 96.39 
heartbeat/minute and 72.18 heartbeats/minute and in the B. subtilis 
test group the mean pre- and post-ingestion heart rates were 96.77 
and 74.78 heartbeats/minute. 

Further Studies on the Interbreeding of an Insular Form of Tropis- 
ternus collaris (Castelnau) with Mainland Forms. Frank N. Young, 
Indiana University. — Experimental crosses of a form of Tropisternus 
collaris from Puerto Rico indicate that the color pattern shows partial 
or complete dominance over that of forms from Florida, Indiana, and 
Mexico. Such crosses are, however, highly sterile or infertile. Crosses 
with a melanic form from Colombia (Ayapel) show dominance of color 
pattern but nearly complete fertility of progeny. Backcrosses have 
produced interesting examples of particulation of the elements of the 
color pattern. 

OTHER PAPER READ 
On the Nature of Communication of Bees. Harold Esch, University of 
Notre Dame (by invitation). 

227 



The Second Record of Coelioxys obtusiventris 
Crawford (Hymenoptera, Megachilidae) 1 

Leland Chandler, Purdue University 

Abstract 

The parasitic bee, Coelioxys obtusiventris Crawford, was described in 1914 from 
Florida and has been known only from the unique female holotype. A second specimen has 
now been collected at Lafayette, Indiana, and is recorded. 

The parasitic megachilid, Coelioxys obtusiventris Crawford, was 
described on the basis of a single female located in the C. F. Baker 
Collection (1). The specimen bore a label "Florida; Palm." without 
additional data. The holotype is Type Specimen Catalog Number 18217, 
U.S. National Museum (1, 2). Mitchell (2) redescribed the species and 
indicated a possible relationship with C. modesta Smith. 

Females of C. modesta and C. obtusiventris characteristically have 
the apex of the sixth metasomal tergite upturned abruptly; however, 
a number of individuals of C. modesta do not exhibit this condition. In 
either case, the apical third of this tergite (in C. modesta) bears nu- 
merous long, erect, black hairs, but these are not sufficiently dense to 
obscure the upturned portion when present. Comparatively, this por- 
tion of the apical tergite of C. obtusiventris is covered so extensively 
with brownish-black hairs that the upturned portion is nearly hidden. 

The most distinguishing structure of C. obtusiventris is the sixth 
metasomal sternite, described by Mitchell (2) as ". . .; sternum 6 
slightly flared apically, with a long apical spine, apical margin with a 
prominent fringe of long brownish hairs, nearly equally the spine in 
length." This feature is so striking and differs so greatly from the 
apical sternite of other species that its diagnostic value has probably 
been under-emphasized within longer descriptions. 

The second specimen of C. obtusiventris was identified among ma- 
terial in the Purdue Entomological Research Collection. The specimen, 
a female, bears the label "Lafayette, Ind. VI • 16 • 59." No other 
information is available nor was the bee associated with other speci- 
mens of Coelioxys or of the host genus Megachile. 

The status of species based upon unique specimens usually creates 
a puzzling situation, especially when localities are indefinite. A second 
record from a remote region does not reduce the puzzle, but it does 
confirm a continued existence. 

Literature Cited 

1. Crawford, J. C. 1914. Some species of the bee genus Coelioxys. Ann. Entomol. Soc. 
America 7(2) :148-159. 

2. Mitchell, T. B. 1962. Bees of the eastern United States. Vol. II. North Carolina Agric. 
Exp. Sta. Tech. Bui. No. 152. 557 p. (esp. 214-215). 



1 Journal Paper No. 2963 of the Purdue University Agricultural Experiment Station. 

228 



Indiana vs. Indian Territory : Misinterpreted Locality Citations 

Leland Chandler, Purdue University 



Abstract 

A number of insect species have been erroneously recorded from Indiana, whereas the 
localities are actually in Oklahoma. An examination of locality labels on some of these 
specimens show the abbreviation [Ind.]. Whether some designation of Territory was 
included originally and removed, or perhaps not printed originally, is not known. Several 
zoogeographic interpretations are based on these resulting in erroneous conclusions. 

Modern systematic treatments include data from numerous disci- 
plines. Thence, by analysis of these data, certain zoogeographic, eco- 
logical, and phylogenetic concepts are derived. A basic source of these 
data is the locality label. 

This paper is devoted, in part, to a clarification of one series of 
locality labels which has been misinterpreted. Fortunately, reinterpre- 
tation does not result in major changes for most of the species involved. 

Mitchell (4) recorded the leaf-cutter bee, Megachile p. parallela 
Smith, from South McAlister, the site listed as an Indiana locality. 
Since this bee does occur in the state, no significance was attached to 
this specific place. Stephen (6) recorded the silk bee, Colletes mandi- 
bularis Smith, from Macalester, also credited to Indiana. This species, 
likewise, occurs throughout the state. La Berge (3) recorded three spe- 
cies of bees from Indiana as follows: Svastra (as Melissodes) o. obliqua 
(Say) from McAllister; Svastra (as Melissodes) p. petulca (Cresson) 
from South McAlester; and Melissodes c. communis Cresson from South 
McAllester. Of the three species, S. petulca is the only one not known 
to occur, being of more southern and western distribution. Deleting the 
Indiana record from the distributional map [(3), fig. 9, pg. 1010] re- 
sults in a somewhat different configuration. On this basis, it would be 
predicted that this species is, however, likely to occur within the state in 
the Lower Wabash Valley or "Pocket" Biotic Unit (2). 

As the number of references to this locality increased, in addition 
to the various spellings, it became important to discover the exact 
location and the reasons for the interpretations. With but meagre facts 
to substantiate the conclusions, it was reasonable to accept the idea 
that all records should be attributed to McAlester, Oklahoma. This 
would mean that labels read as "Indiana" were in fact "Indian Territory." 

Confirmation that such an idea was correct has been found in the 
recent publications by Campbell (1) and Quate and Thompson (5). 
Campbell recorded Lobopoda yiigrans (Melsheimer) from South McAles- 
ter and Atoka, both localities given as being in Indiana. The inclusion 
of Atoka, also an Oklahoma locality, further represents a label mis- 
interpretation. 

The reference by Quate and Thompson strangely does not have 
reference to Indiana, but to Arkansas. It does, however, give complete 

229 



230 Indiana Academy of Science 

label information as follows: "Vinita, Ind. T., June 7-8, '99, Wickham." 
The species referred to is Melanotus lanei Quate and the locality is 
cited as being- in Arkansas. 

It has become apparent that one or more collections of beetles and 
of bees contains materials with labels that can be misread. Assuredly, all 
references to Atoka, McAlester (and its derivative spellings), and Vinita 
are Oklahoma localities, not Indiana or Arkansas. 



Literature Cited 

1. Campbell, John M. 1966. A revision of the genus Lobopoda (Coleoptera :Alleculidae) 
in North America and the West Indies. Illinois Biol. Monog. 37, Univ. Illinois Press, 
Urban a. 

2. Chandler, Leland. 1966. The origin and composition of the insect fauna, p. 345-361. 
In A. A. Lindsey [ed.] Natural Features of Indiana. Indiana Acad. Sci. Sesquicenten- 
nial Volume, Indianapolis. 600 p. 

3. La Berge, Wallace E. 1956. A revision of the bees of the genus Mclissodcs in North 
and Central America. Part 1 (Hymenoptera, Apidae). Univ. Kansas Sci. Bull. 37, Pt. 2 
(18) :911-1194. 

4. Mitchell, T. B. 1937. A revision of the genus Megachile in the Nearctic Region. Part 
Vi. Taxonomy of subgenera Argyropile, Leptorachis, Pseudocentron, Acentron and 
Melanosarus. (Hymenoptera :Megachilidae). Trans. American Entomol. Soc. 63:45-83. 

5. Quate, Laurence W., and Sarah E. Thompson. 1967. Revision of click beetles of 
genus Melanotus in America north of Mexico. Proc. U. S. Nat. Mus. 121 (3568) :l-83. 

6. Stephen, W. P. 1954. A revision of the bee genus Colletes in America north of 
Mexico. Univ. Kansas Sci. Bull. 36, Pt. 1 (6) :149-527. 



The Occurrence of Chalybion zimmermanni Dahlbom (Sphecidae) 

in Indiana 

Gertrude L. Ward, Earlham College 

Abstract 

Chalybion zimmermanni Dahlbom (Hymenoptera, Sphecidae) is added to the list of 
insects of Indiana. The northern extent of this insect's range was shown by Bohart in 1963 
as Tennessee. It was found nesting in holes in wood in Wayne County, Indiana, in 1968, 
and in 1969 evidence of nesting was found in six other counties. Nests are provisioned with 
small spiders, generally Argiopidae or Theridiidae. An unusual two-layered plug is made 
of mud and uric acid. The white uric acid contrasts distinctly with the old wood around 
the nest hole. A small number of nests were located in deserted nests of the yellow-legged 
mud dauber, Sceliphron caementarium. 

During a recent study of three mud-using wasps, Sceliphron cae- 
mentarium (Drury), Trypargilum politum (Say), and Chalybion cali- 
fornicum (Saussure), another large mud-using wasp appeared in my 
study area near Centerville, Wayne County, Indiana. This was Chaly- 
bion zimmermanni Dahlbom. On 1 August 1968, an assistant, Jay 
Myers, called my attention to the metallic blue wasp which was plugging 
a hole in the wooden plate of an old tool shed. The original borer was 
probably a beetle. 

We watched this wasp clean the debris out of a second hole, gather 
mud from old Sceliphron caementarium nests and form a barricade 
deep in the hole. She collected small spiders, laid an egg on the abdomen 
of one of the first spiders, and, on two occasions, built a wall about 
halfway down the hole. The remaining space was used for a second cell. 
When this was filled, the female formed a countersunk plug of dark 
mud. This sometimes required three or four loads of mud, apparently 
all of it gathered from old nests. 

After the dark mud was dry, she made a level or slightly concave 
seal of white material. We collected some of this white plaster from one 
cell, and analysis by infra-red spectrophotometry showed that it was 
mainly uric acid. The wasp disappeared after completing six holes. 

In May, 1969, six small cylindrical traps made of fiberglass screen- 
ing were fastened over the white-plugged holes. Emergence of wasps 
from these cells was noticed first on 28 June. Nothing emerged from one 
cell. Two males were collected and are in the Joseph Moore Museum of 
Earlham College. Five wasps, including both males and females, were 
released. A summary of the adults which emerged is shown in Table 1. 

Several of the wasps which emerged were chilled in a glass jar in a 
food freezer for about 4 minutes, and then were measured before their 
release. The clypeal teeth were examined to determine the sex. The male 
has a median tooth which is longer than either of the side teeth, and the 
female has three small flaps with the middle one not shorter than the 
others, as it is in Chalybion calif or nicum. 

231 



232 Indiana Academy of Science 

Table 1. Adults emerging from cells of Chalybion zimmermanni Dahl- 

bom, Indiana, 1969. 





Number 


Body 




Cell 


emerging 


length (mm) 


Sex 


A 


1 


18 


F (?) 


B 


2 


19, 15.5 


F, M 


C 





— ■ 


— 


D 


1 


16 


M 


E 


2 


18, 16 


F (?), M 


F 


1 


large 


F (?) 



Two females started to make nests in old holes in the wooden plate 
on 16 July 1969. Eight holes were filled, four of which had been occupied 
in 1968. 

An attempt was made in 1969 to determine the extent of C. zimmer- 
manni penetration into Indiana. In 1963, Bohart and Menke (1) reported 
the northern extent of the range as Tennessee. Although no other 
specimens were collected, the distinctive two-layered, two-colored plugs 
were found in the following counties: Clark, Crawford, Dearborn, Ohio, 
Ripley, and Switzerland. An equal amount of collecting yielded negative 
results in Decatur, Jackson, Lawrence, Rush, and Washington counties. 

Most of the cells were in the old timbers of barns and sheds, but in 
Clark County four of these plugs were found in a mass of deserted 
Sceliphron caementarium cells. Rau (5) reported that in Mexico C. 
zimmermanni uses the old cells of Sceliphron assimilis (Dahlbom). 

A total of 40 spiders was examined from C. zimmermanni nests. 
The majority, 60%, were in the family Araneidae, and the remaining 
40% were in the Theridiidae. These are shown in Table 2. 

Table 2. Prey of Chalybion zimmermanni Dahlbom in Indiana, 1969. 





No. 


% 


Total % 


Araneidae 








Araneus spp. 


VI 


30.0 




Argiope aurantia Lucas 


9 


22.5 




Argiope trifasciata (Forskal) 


2 


5.0 




Cyclosa conica (Pallas) 


1 


2.5 


60.0 


Theridiidae 








Theridion frondeum Hentz 


15 


37.5 




Asagena americana Emerton 


1 


2.5 


40.0 




40 




100.0 



Entomology 233 

Whether or not the uric acid serves as a deterrent to parasites 
which might invade the completed cell through the plug has not been 
determined. The use of two colors of material in the final plug was 
reported by Williams (6) for Chalybion violaceum (Fabricius) in the 
Philippines. He said that this wasp ". . . stores her small spiders in 
some convenient hollow, as a rung socket, penholder base, old mud nest, 
etc., and simply plugs up the aperture, first with mud or moist earth, and 
finishes this off with a mixture of the excreta of geckos (lizards), giving 
the plug a whitish or plaster-like appearance." 

Williams (6) also reported that d'Herculais in 1882, observing 
Chalybion chalybeiis (Smith) at Port Natal, Africa, noticed "this curious 
habit" of using a light-color final plug. The material used at Port Natal 
was bird excrement. 

Iwata (2) reported that he saw Sceliphron (Chalybion) inflexum 
Sickmann on Taiwan plaster the mud seal of her nest with white 
material, and also (3) reported this in Thailand. Yamamoto (7) said 
that he saw this wasp in Japan collecting bird droppings which were 
still damp, and using them for plastering. 

In India, Jayaker and Spurway (4) observed Chalybion bengaleyise 
Dahlbom make a plug of brown mud and then cover it with a plug of 
white. One wasp collected the white material from the feces of a pet 
tortoise and another wasp used bird feces. 

Summary and Conclusions 

Chalybion zimmewnanni, having the northern edge of its range 
reported in 1963 as Tennessee, appears to be moving northward in 
Indiana. The food stored for the young is small spiders from the 
families Araneidae and Theridiidae. It most often nests in borings in old 
timbers, but has been seen to use the deserted mud nests of Sceliphron 
caementarium. 

Acknowledgments 

The author wishes to express appreciation to Alan Rushton for 
spectrophotometric analysis, to National Science Foundation for an 
undergraduate assistant under Grant GY 4495, and to Research Corpora- 
tion for a portion of the institutional grant to Earlham College. 

Literature Cited 

1. BOHART, R. M., and A. S. Menke. 1963. A reclassification of the Sphecinae with a 
revision of the nearctic species of the tribes Sceliphronini and Sphecinae. Univ. Calif. 
Berkeley. Pub. Entomol. 30:19-182. 

2. Iwata, K. 1939. Habits of some solitary wasps in Formosa (IV). Trans. Natur. Hist. 
Soc. Formosa 29:161-178. 

3. Iwata, K. 1964. Bionomics of non-social wasps in Thailand. Nature and Life in 
Southeast Asia 3 :323-383. 

4. Jayaker, S. D., and H. Spurway. 1963. Use of vertebrate faeces by the sphecoid wasp 
Chalybion bengalense Dahlb. J. Bombay Natur. Hist. Soc. 60 :748-749. 

5. Rau, P. 1943. The nesting habits of certain Sphecid wasps of Mexico, with notes on 
their parasites. Ann. Entomol. Soc. Amer. 36 :647-653. 

6. Williams, F. X. 1919. Philippine wasp studies. Bull. Expt. Sta. Hawaiian Sugar 
Planters' Assoc, Entomol. Ser. No. 14 :19-181. 

7. Yamamoto, D. 1942. Habits of Sceliphron (Chalybion) inflexum Sickmann. Kontyu 
16 :69-75. 



A Taxonomic Key to the Collembola 
in Four Serai Stages Leading to the Beech-Maple Climax 

Patricia M. Arnetti, Indiana State University 



Abstract 

From April through July, 1968, 96 leaf litter samples were taken from an old field, 
oak and maple-oak dominated serai stages, and a beech-maple climax in Parke County, 
Indiana. Collembola were extricated by a modified Tullgren funnel apparatus, collected, 
and identified. A key was based on morphology and color for 32 species, which represent 
20 genera and 5 families. A table of the distribution of each species by serai stage was 
included. 

Introduction 

From April through July, 1968, 96 leaf litter samples of 1.0 dm^ 
each were taken from an old field, oak and maple-oak dominated serai 
stages, and a beech-maple climax in Allee Woods, Parke County, 
Indiana. Collembola were extricated by a modified Tullgren apparatus 
and identified (2, 3). With current keys, not all individuals could be 
identified to species. The key represents 1,533 individuals, 32 species, 
20 genera and 5 families. The purpose of this paper was to present a 
simplified key to the species in the four serai stages leading to the 
beech-maple climax. The changes in Collembola populations as influenced 
by plant successional patterns was previously described (1). 

Method 

After all species were identified, the most obvious external char- 
acteristics were selected for these 32 species and a key was constructed. 
The key was based on color and morphology. The primary morphological 
characteristics were: 1) length and shape of the body; 2) degree of 
fusion and length of abdominal segments; 3) nature of prothorax; 
4) presence of scales, body hair, and setae; 5) number of eyespots; and 
6) number of segments and length of antennae. The distribution of each 
species by serai stage (Table 1) shows the relative abundance of each 
species per ecological area and hence could be helpful in confirming an 
identification of an individual from a comparable sere. 

Taxonomic Key 

1. Body elongate; abdominal segments distinct although IV, V, and VI or V and VI 
may be ankylosed 2 

suborder Arthropleona Borner 

1'. Body globular ; abdominal segments not distinct ; the first four abdominal seg- 
ments fused with thorax 26 

suborder Symphypleona Borner 
family Sminthuridae 

2. Prothorax reduced and membraneous 3 

superfamily Entomobryoidea Womersley 

2'. Prothorax similar to other segments 22 

superfamily Poduroidea Womersley 



Present address: Northwestern High School, Kokomo, Indiana 46901. 

234 



Entomology 235 

3. Scales absent ; body segments equal to subequal in length ; antennae with 4 
simple segments ; last abdominal segments may be ankylosed 4 

family Isotomidae 

3'. Scales and/or brush-like setae present ; third or fourth body segment elongate ; 
antennae with 4-6 segments, the third and fourth sometimes annulated ; abdominal 

segments always distinct 11 

family Entomobryidae 

4. Fourth, fifth, and sixth abdominal segments ankylosed 5 

Folsomia 

4'. Fourth, fifth, and sixth abdominal segments not ankylosed 6 

5. E>es absent; pigment absent 

Folsomia fimentaria L. 

5'. Eyes 2 and 2 ; pigment gray to black 

Folsomia quadrioculata Tullberg 

6. Body with bothriotrichia (long sensory body hairs) 

Isotomurus palustris Muller 
6'. Body without bothriotrichia 7 

7. Manubrium (single part of furcula ; is attached to abdomen) much shorter than 
dentes (middle part of furcula; is forked), with many ventral setae; Abd. IV 
usually shorter than Abd. Ill 8 

Isotoma 
7'. Manubrium often longer than dentes with few or no ventral setae ; Abd. IV 

usually longer than Abd. Ill 10 

Proisotoma 

8. Eyes 4 and 4 on round patches connected by an inverted V-shaped mark 

* Isotoma eunotabilis Folsom 
8'. Eyes 8 and 8 on elongate patches without an inverted V-shaped mark 9 

9. Length 0.6 mm 

Isotoma viridis Bourlet 

9'. Length 1.5 mm 

*Isotoma olivacea Tullberg 

10. Dentes shorter than manubrium 

Proisotoma minuta Tullberg 

10'. Dentes longer than manubrium 

*Proisotoma immersa Folsom 

11. Abd. Ill longer than Abd. IV; mucrones (tip of furcula) hairy; Ant. Ill longest 
segment and annulated; antennae 4-segmented 12 

subfamily Tomocerinae 

Tomocerus 

11'. Abd. Ill shorter than Abd. IV; mucrones not hairy; antennae 4- to 6-segmented __ 15 

subfamily Entomobryinae 

12. Maxilla bearded 13 

12'. Maxilla not bearded 14 

13. Antennae longer than body 

*Tomocerus elongatus Maynard 

13'. Antennae shorter than body 

* Tomocerus flavescens Tullberg 

14. Dental spines tridentate; Th. II overlapping but not obscuring Th. I dorsally 

*Tomocerus minor Lubbock 

14'. Dental spines simple; Th. II obscuring Th. I dorsally 

*Tomocerus vulgaris Tullberg 

15. Antennae with 6 segments 

Orchesella ainsliei Folsom 
15'. Antennae with 4 segments 16 

16. Body without scales 17 



236 Indiana Academy of Science 

16'. Body with scales 21 

Lepidocyrtus 

17. Eyes not on daik patches 

*Isotobryoides ochracius Maynard 

17'. Eyes on dark patches 18 

Entomobrya 

18. Body unicolorous without crossbands of contrasting color 19 

18'. Body with dark dorsal and lateral spots or bands or both on light ground 

color 20 

19. Color gray to olive green to bluish purple 

Entomobrya marginata Tullberg 

19'. Color yellow to yellow-orange 

^Entomobrya atrocincta f. 
pseudopcrpulchra Mills 

20. Transverse bands on every segment 

* Entomobrya multifasciata Tullberg 

20'. Transverse bands on most segments ; Abd. I with 2 dark dorsal spots 

Entomobrya assuta Folsom 

21. Purple pigment on Abd. IV 

* Lepidocyrtus unifasciatus James 

21'. Purple pigment on antennae and legs. 

Lepidocyrtus curvicollis Bourlet 

22. Eyes absent 

family Onychiuridae 
*Onychiuru8 armatus Tullberg 

22'. Eyes present 23 

family Poduridae 

23. Pigment present 24 

23'. Pigment absent 

*Neanura barberi Handschin 

24. Furcula well developed 25 

24'. Furcula reduced 

*Xenylla welchi Folsom 

25. Color brown and yellow mottled ; abdomen not considerably distended 

*Hypogastrura tigrina Harvey 

25'. Color dark gray to black ; abdomen considerably distended 

*Pseudachorutes simplex Maynard 

26. Antennae shorter than head; eyes absent 27 

Neelus 
26'. Antennae longer than head ; eyes present 28 

27. Color white 

*Neelus albus Maynard 

27' Color yellow with red speckles 

*Neelus maculosus Maynard 

28. Color purplish red 

*Arrhopalites binoculatus Borner 
28'. Color not purplish red 29 

29. Black pigment spots present on abdomen 30 

29'. Black pigment spots absent; eyes orange 

Katiannina maegillivrayi Banks 

30. Antennae pale basally 

*Dicyrtomina variabilis Maynard 

30'. Antennae dark basally 31 

Sminthtirinus 



Entomology 



237 



31. Body with much dark purplish black pigment ; antennae and legs banded with 

dark pigment 

*Sminthurinus radiculus Maynard 

31'. Body with black pigment reduced in form of lateral polygons; buff and orange 

spots present 

*Sminthurinus radiculus f. pictus Maynard 



Species first reported for Indiana. 

Table 1. Species distribution by serai stage. 





Beech- 


Maple- 




Old 




Species 


maple 


< »ak 


Oak 


field 


Total 


Arrhopalites binoculatus 


2 


1 


3 


1 


7 


Dicyrtominia variabilis 





1 





o 


1 


Entomobrya assuta 


42 


7 


it; 


s 


73 


Entomobrya atrocincta 












f. pseudoperpulchra 


L6 


8 


41 


2 


67 


Entomobrya marginata 


13 


8 


10 


11 


42 


Entomobrya multij asciata 


44 


69 


121 


19 


253 


Folsomia fimentaria 


10 


31 


is 


4 


63 


Folsomia quadrioculata 


6 


1 








7 


Hypogastrura tigrina 


o 





1 


1 


2 


Isotobryoides ochracius 


35 


49 


42 


2 


L28 


Isotoma eunotabilis 


1 


2 


3 


o 


6 


Isotoma olivacea 


8 


ir, 


46 


16 


85 


Isotoma viridis 





1 








1 


Isotomurus palustris 


2 


2 


3 





7 


Katiannina macgillivrayi 


2 


1 


2 


1 


6 


Lepidocyrtus curvicollis 


5 


3 


1 





9 


Lepidocyrtus unifasciatus 





2 








2 


Neanura barberi 








2 





2 


Neelus albus 


1 


3 


2 





6 


Neelus maculosus 





o 


1 





1 


Onychiurus armatus 


94 


124 


43 


4 


265 


Orchesella ainsliei 


(i 





o 


I 


1 


Proisotoma immersa 


2 


6 


14 


1 


23 


Proisotoma minuta 


(l 





5 





5 


Pseudachomtes simplex 


1 


1 


\ 


o 


<■. 


Sminthurinus radiculus 


2 


1 








3 


Sminthurinus radiculus 












f. pictus 


5 


3 


17 





25 


Tomocerus elongatus 


1 


1 


1 





3 


Tomocerus flavescens 


2 


2 


2 





<; 


Tomocerus minor 


i:\ 


74 


L15 


36 


298 


Tomocerus vulgaris 


1 


3 


38 


5 


47 


Xenylla welchi 


20 


10 


33 


20 


S3 



Literature Cited 

1. Arnett, Patricia M. 1969. A Study of Collembolan populations associated with four 
serai stages leading to the beech-maple climax. Proc. Indiana Acad. Sci. 78 :231-240. 

2. Folsom, J. W. 1937. Nearctic Collembola or springtails, of the family Isotomidae. 
Bull. U. S. Natur. Mus. 168. 145 p. 

3. Maynard, E. C. 1951. The Collembola of New York State. Comstock Publishing 
Company, Inc., Ithaca, New York. 388 p. 



Factors Influencing the Species Composition of 
Mosquito Populations in Indiana 1 

R. E. Siverly, Ball State University 



Abstract 

Climate, and natural features such as forests, bodies of water, soils, vegetation and 
shade are location-dependent factors which influence species composition of mosquito 
populations in Indiana. Other factors include: urbanization, suburban development, de- 
forestation, drainage, and tillage of soil. 

The mosquito fauna of Indiana has both northern and southern climatic elements. 
Indiana is the northern range limit for certain southern species (e.g. Psorophora cyanc- 
scens, P. howardii), and the southern limit for certain northern species (e.g., Aedcs 
abserratus, A. excrucians). 

Indiana's mosquito fauna is characteristically sylvan, plains species being few and 
limited in distribution. Aedes stimulans, a northern forest mosquito, is well established 
north of the Wisconsin glacial boundary. South of this boundary it is largely limited to 
beech-maple tracts in the Southwestern Till Plain. 

About 12 of the 50 species of Indiana mosquitoes are produced in permanent water. 
Included are species of Mansonia, Anopheles, Culex, and Uranotaenia. Temporary pools 
in depressions produce both univoltine and multivoltine Aedes and Psorophora. Recurrent 
rains during summer disposes continual production of multivoltine members of these two 
genera. 

Some species are directly dependent on certain plants either for production or as 
sources of carbohydrate. Indirect effects of vegetation include shade and humidity. 

Soils underlying depressions used by early spring mosquitoes contained clay loam and 
were poorly drained. In one instance, soil disturbance appeared to enhance mosquito pro- 
duction. The need for further studies of relationships between soil characteristics and 
mosquito production is stressed. 

Urbanization tends to substitute one set of mosquito problems for another, rather 
than eliminate all problems. Species diversity decreases, with attendant increase in 
density of adaptive species. Domestic species (e.g., Culex pipiens, the house mosquito) can 
attain dominance as a result of improper liquid waste and solid waste disposal practices. 
Aedes triseriatus, A. vexans, A. sticticus, A. trivittatus and Psorophora confinnis are 
some of the para-domestic mosquitos which annoy suburban dwellers. 

Because of diverse conditions in Indiana, one set of recommendations for mosquito 
abatement will not suffice for all localities, and control recommendations for a given 
community will require periodic revision. 



Introduction 

The species composition of mosquito populations varies from one 
part of Indiana to another. Presently, 50 species are listed for Indiana. 
Twenty-seven species were reported from Delaware County (8) and 
30 species are listed for Wayne County (J. H. Hart, personal communi- 
cation). 

Some species are limited to one or more of the northernmost tier 
of counties. Other species are known to occur only in the extreme 
southern parts of Indiana. Species composition even varies between 
adjacent counties. Evidently, not one factor (e.g., climate), but a com- 

1 This investiyiration was supported in part by a Riant from the Indiana State Board 
of Health, funded by PL 89-749, Section 314(d). 

238 



Entomology 239 

plex of factors influences these patterns of distribution. This paper 
identifies some of the factors responsible for variations in species 
composition of mosquito populations in Indiana, and makes some pre- 
dictions, based upon these observations, regarding species composition 
of mosquito populations in future years. Factors are considered as 
location-dependent and location-independent. 

Location — Dependent Factors 
Climate 

Members of the genus Psorophora generally are considered as 
southern mosquitoes which develop rapidly in temporary bodies of water 
following summer rains. In years with cool or dry summers, some of 
the eight species which represent this genus in Indiana may be absent, 
at least in the northern part of the state. Psorophoi^a ferox, P. varipes 
and P. horrida were not found in Delaware County during an intensive 
survey in 1964 (8). These species were taken in biting collections during 
the summer of 1969 at one of the sampling sites used in the 1964 survey. 
Thus, climatic variations from year to year will influence the species 
make-up of mosquito populations in a given area. 

Neither P. cyanescens nor P. howardii has been reported north of 
Bartholomew County. From state distribution records (1, 2) it appears 
that both of these species are rare or absent north of 40° latitude. 

Conversely, some northern species attain the southern limit of 
their ranges in northern Indiana. Aedes abserratus, one of the black- 
legged, univoltine, cold-hardy mosquitoes, had not been reported as 
occurring south of LaPorte, Lagrange, and Steuben Counties. Nationwide, 
it has not been reported south of 40° latitude. One of the band-legged 
Aedes — A. excrucians — is found in kettles and bogs in northern Indiana, 
but it fades out in numbers, and its occurrence is infrequent or rare in 
central Indiana. As in the case of A. abserratus, Illinois and Indiana 
appear to be the southern limit of its range. 

The necessity for water in mosquito development is axiomatic. Rise 
and decline in number of some species is directly dependent upon patterns 
of precipitation in summer. Production of Aedes and Psorophora species 
from forest tracts during August often is prevented — not because of 
insufficient warmth — but because of insufficient rainfall. Rainfall of 
three inches or more within a 24-hour period may be required to inundate 
soil depressions containing eggs, and to provide standing water for a 
week or more thereafter. With other species, numbers produced in a given 
year remains about the same, regardless of whether summers are wet 
or dry. This applies both in Indiana and Wisconsin. (8, 9). 

Although other factors undoubtedly play a role, climate probably is 
the overriding influence in the distribution of sub-tropical Psorophora 
and certain Nearctic Aedes in Indiana. Mean daily minimum tempera- 
tures and mean daily maximum temperatures vary 10° F or more between 
northern and southern Indiana (6). This suggests sufficient gradient for 
differences in species composition in the two areas; such is true with 
mosquitoes. 



240 Indiana Academy of Science 

Natural Features 

Natural features which influence the species make-up of local mos- 
quito populations include forests, bodies of water, soils, and vegetation. 

Forests. In northern, and in certain parts of central Indiana, the 
immature stages of one forest mosquito, Aedes stimulans, occur in 
cattail ponds and roadside ditches, as well as in woodland pools. With 
some exceptions, the most recent Wisconsin glacial boundary (10) is the 
southern limit of its range. 

A. stimulans was collected south of this boundary in southeastern 
Indiana in the spring of 1969, but only in relict populations in forested 
tracts over wet soils. On the other hand, this species was absent in wet- 
soil areas, which formerly were forested, just north of the Wisconsin 
glacial line. In such habitats it apparently is displaced with competitive 
species such as A. canadensis or A. vexans, which tolerate exposure in 
unshaded areas. The distribution of A. stimulans is discussed further in 
connection with soils and vegetation. 

Aedes triseriatus is another forest mosquito, widely distributed in 
Indiana. It is commonly known as the treehole mosquito since it utilizes 
rot holes containing water in stumps, as well as cavities which develop 
on trunks and limbs and hold water as a result of natural processes. 

Stumps need to be in the right stage of decay to hold water. With 
extensive decay, the stump becomes entirely hollow and no water is re- 
tained. This suggests a time span for a stump hole as a larval habitat. 
Over a 10-year period, a stump hole in Delaware County yielded Ano- 
pheles barberi and Cnlex restuans as well as Aedes triseriatus. After 10 
years the cavity rotted through, and no longer held water. 

Aedes triseriatus lately has come into prominence because of its 
implication as a potential vector of California encephalitis. Small forest 
mammals, such as squirrels, are believed to act as a reservoir for the 
causative arbovirus. This vector is not limited to forest tracts, however, 
since immature stages also are found in artificial containers, such as old 
tires. The negative association between California encephalitis and urban 
environment may result from limited numbers of reservoir hosts instead 
of limited numbers of suitable vectors in towns and cities. 

Bodies of Water. The simplest classification of bodies of water is 
permanent or temporary. Permanent water describes a site where water 
stands all year during most years. Temporary bodies of water contain 
water for shorter periods. 

Contrary to popular belief, the size of the mosquito crop is not 
directly proportional to the expanse of water. Only about a dozen of 
Indiana mosquitoes can utilize permanent water as production sites. Such 
species have adaptations which permit them to complete development 
in a habitat which is alive with predators. Larvae and pupae of Mansonia 
perturbans attach anal siphons to the subterranean tracheal systems of 
such aquatic plants as cattails and sedges, and acquire oxygen in this 
way. They never need to surface except for the brief period required 
for emergence of the adult. 



Entomology 241 

Steuben County has about 7% of its area in wetlands, much of it 
in dry marshes, open water, shallow and deep shrub swamps, all in the 
permanent water category (4). Adult trap collections made in this 
county in 1969 contained relatively high percentages of Mansonia per- 
turbans (42%) and Anopheles species (8%). 

Larvae of Anopheles mosquitoes are commonly found in emergent 
vegetation and in mats of floating vegetation where fish and other 
predators do not easily gain access. The resemblance of these larvae 
to floating sticks may have some protective value. The same general 
habitat used by Anopheles is also used by Culex species. 

Larvae of Culex territans, a rather ubiquitous mosquito which feeds 
on cold blooded animals, also may be found along with Uranotaenia 
sapphirina, in permanent bodies of water. 

Farm ponds seldom serve as production sites for mosquitoes when 
clean shore lines are maintained, where there is little if any emergent 
vegetation, and when fish and other predators are present. 

Many kinds of depressions — some man-made — hold temporary pools 
of water which serve as production sites for mosquitoes after spring 
thaw, snow melt or rains. There may be a predator problem for those 
which develop slowly. For the most part, mosquitoes which develop in 
pools and puddles either have a lower temperature threshold for develop- 
ment than their predators, or they simply are geared for more rapid 
development. The univoltine Aedes which develop during early spring 
while the water is 40-50°F include: A. abserratus, A. aurifer, A. cana- 
densis, A. cinereus, A. excrucians, A. fitchii, A. flavescens, A. grossbecki, 
A. stimulans, and A. thibaulti. These are off the water and on the wing 
before crayfish, predaceous beetles, dragonfly nymphs and other preda- 
tors become established in the temporary pools and puddles. 

Multivoltine Aedes and Psorophora which undergo rapid develop- 
ment in temporary water impoundments following spring and summer 
rains include: A. trivittatus, A. sticticus, A. vexans and all of the 
Psorophora. A week to 10 days is sufficient for these mosquitoes to com- 
plete development, assuming normal summer temperatures. They may 
utilize the same habitats as the univoltine Aedes which developed earlier 
in the year. 

The importance of temporary bodies of water should not be under- 
rated. One woodlot pool in Delaware County was estimated to produce 
more than 600,000 Aedes stimulans (8). Flood plains, such as the Wabash 
in Vigo County, are notorious for mosquito production. Aedes vexans is a 
prime contributor to this kind of mosquito problem. 

Soils. Probably no one factor is more important in mosquito produc- 
tion than soil, yet no factor is more poorly understood. Soil character- 
istics and numbers and kinds of immature mosquitoes present in the 
water overlaying these soils were studied in the spring of 1969. Soil 
samples were taken with a soil auger at the approximate center of the 
depressions where mosquito collections were made. Soil cores approxi- 
mately IV2, inches long and 1% inches in diameter were transported in 
plastic bags to the Soil Conservation Office in Muncie for soil descriptions. 



242 



Indiana Academy of Science 



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Entomology 243 

Soil results are summarized in Table 1. One characteristic shared by 
all soil samples was poor drainage. With the exception of the samples 
taken in Steuben County, all soils had a clay component. Amounts of 
organic matter in the samples were quite variable. In cases where 
development had passed the larval stage, pupae or adults were collected. 
With the exception of Steuben County samples, all soil samples were 
taken from forest soils. 

Samples collected in Franklin County are noteworthy. One soil sample 
was taken in a water-filled depression where no mosquitoes were present; 
another soil sample was taken in a ditch nearby, where larvae were 
present. According to the Soil Conservation Service, the ditch sample 
revealed more evidence of disturbance than the soil taken from the 
depression. Conceivably more minerals and nutrients were available in 
the ditch, having been brought to the soil surface as a result of excava- 
tion. This same phenomenon was observed in northern Wisconsin where 
greater density of larvae was observed in a borrow pit adjoining a 
logging road than in an adjacent swamp. 

Aedes fiavescens is rare in Indiana (8). Although peat overlaying 
marl probably is not prerequisite for production of this mosquito, the 
unusual density of both immatures and adults near Orland, in Steuben 
County, suggests that A. fiavescens may be produced over Edwards or 
Warner Muck in other areas. 

Less definite is the association between A. stimulans and Brookston 
soils. These are timber soils formed from fine textured till from recent 
(Wisconsin) glaciation. South of the Wisconsin glacial boundary A. 
stimulans was found in minority numbers in Clark and Ripley Counties. 
The associated soil series were Avonburg and Clermont, respectively. 
These are timber soils formed on old (Illinoian) glacial till. 

From Table 1, one could generalize that mosquitoes are likely to be 
produced in depressions in poorly drained forest soils which have clay 
loam components. Undoubtedly, some species are tolerant of a wide 
variety of soils, others less so. More precise information regarding soil 
characteristics of habitats will have to be obtained before predictions can 
be made relative to species occurrence. This subject warrants further 
investigation. 

Vegetation. The association of Wyeomyia smithii with the pitcher 
plant, Sarracenia purpurea, is an example of direct influence. W. 
smithii is not known to occur in the absence of this pitcher plant. W. 
smithii was reported in LaPorte County in 1964 (8) and subsequently 
collected in Steuben County. The direct dependence of Mansonia per- 
turbans on plants such as cattail, water lily and sedge has been 
mentioned. 

Plants may be of direct benefit to adult mosquitoes as a source of 
nectar for food. Preliminary observations in Delaware County indicate 
that Aedes stimulans feeds on nectar of Phlox divaricata (wild phlox). 
The occurrence of nectar-yielding plants near its sources of production 
may influence its distribution. 



244 Indiana Academy of Science 

Aedes stimulans is associated with both oak-hickory and beech- 
maple climax forests in the northern part of the state but is limited in 
its occurrence to beech-maple tracts south of the Wisconsin glacial line. 
Aedes canadensis and A. vexans usually are dominant in oak-hickory 
forests on Illinoian drift. 

Some of the most influential effects of vegetation on species composi- 
tion are the indirect effects of shade and humidity. Some mosquitoes are 
more tolerant of open, well-lighted conditions than others. A given species 
may seek shade in the southern part of its range, but tolerate unshaded 
conditions in the northern part of its range. This has been observed with 
Aedes aurifer. Larvae were found under dense shade along the Muscata- 
tuck Flats in Washington County, but also in a peat bog, near button 
bush, in fairly open water in LaPorte County. Aedes aurifer were also 
collected from very open, unshaded pools in a bog near Tomahawk, Wis- 
consin. 

Culiseta melanura and Culex territans larvae were collected from the 
same pools in Delaware County. Culex territans larvae occupied the 
open portions of the pools. Culiseta melanura larvae were found in the 
darker recesses, close by hollow rotting bases of trees or in densely 
shaded root holes in the forest floor. However, in northern Wisconsin, 
C. melanura larvae were collected in sphagnum-leatherleaf bogs, quite 
similar in appearance to habitats utilized by A. aurifer. 

During warm weather the tolerance for exposure to light is an 
indication of capacity for rapid development, since lighted pools are also 
exposed to the warmth of the sun's rays. Several Psorophora species and 
some Aedes (especially A. trivittatus) , which develop in open shallow 
sunlit pools, are able to complete development before depressions 
become dry through drainage and evaporation. 

Because of their small size and high surface: volume ratio, adult 
mosquitoes are particularly subject to desiccation. The association of 
adult mosquitoes and shrubbery is well known. Here again, the effect of 
vegetation for mosquitoes is an indirect one, through providing more 
humid conditions for harborage than would be encountered in unshaded 
areas. In general, Indiana's mosquito fauna reflects its history of a pre- 
dominant forest vegetation. Plains mosquitoes are infrequent or absent 
in the state. 

Location-Independent Factors 

Urbanization 

In a strict sense, whether towns eventually become cities may 
depend upon location, but cities of 50,000 or more occur in all regions of 
the state. Regardless of location, the most significant feature of urban 
mosquito problems is their striking similarity. Mosquito problems in 
these urban communities arise mainly from solid waste and liquid waste. 

Litter is solid waste, including artificial receptacles which hold water 
such as cans, buckets, and old tires. Old tires hold water for surprisingly 



Entomology 245 

long periods, and are utilized as production sources by Culex pipiens, C. 
restuans, Aedes triseriatus, and other species of mosquitoes. Used tires, 
stockpiled in the open at junk yards and service stations, often serve as 
production foci for domestic mosquitoes. Urbanization and heavy traffic 
do not appear to be a deterrent, especially for C. pipiens. The implication 
of this species, the northern house mosquito, in the transmission of 
St. Louis Encephalitis will be discussed later. 

The yellow fever mosquito, Aedes aegypti, was reported from 
Charlestown, in Clark County, in 1941 (3). A visit to Clark County in the 
spring of 1969 revealed no trace of A. aegypti. There was no litter along 
the creek where early collections were made. Other production sites were 
plentiful in adjacent areas, and tires were positive for house mosquito 
larvae. Later in 1969, tires at an open dump in Posey County were 
checked for A. aegypti, but none were collected. Other mosquitoes, 
chiefly Culex pipiens, were present. The availability of artificial con- 
tainers, particularly in areas within its former range, poses a threat of 
recurrence of A. aegypti within the state. 

Not all artificial water-holding receptacles are litter. Bird baths, eave 
troughs and garden pools also may serve as production sources for 
domestic mosquitoes. These tend to be less important than litter recep- 
tacles, and more residential than commercial. 

The discharge of household sewage into streams is a common prac- 
tice in towns and cities in the midwest. Often, the water in these streams 
moves slowly or tends to stagnate. Organic enrichment and stagnation 
provide conditions which are ideal for production of Culex pipiens. 
Heavy house mosquito production, in proximity to concentrations of both 
human and bird populations are the basic ingredients for epidemics of 
St. Louis encephalitis. The isolation of St. Louis encephalitis virus 
from Culex pipiens in Evansville and Boonville in 1964 was reported by 
Newhouse and Siverly (5). 

Water polluted by human sewage is not the only medium created 
by urbanization and used by domestic mosquitoes. Liquid waste from 
meat packing plants may be a potent source of mosquitoes, whether run 
into standing ponds, lagoons, or other bodies of water. 

Obviously, the solution to these mosquito problems created by 
urbanization is through sanitation programs and adequate waste disposal 
systems. My purpose is not to discuss how this can be done, but to 
show that species composition of mosquito populations does change with 
industrialization, urbanization, and concentrations of human population. 

Suburban development, and the tendency for families to live in or 
near wooded tracts has intensified certain mosquito problems. Such 
species as Aedes vexans, A. trivittatus, A. sticticus, Psorophora confinnis 
and Anopheles punctipennis — once regarded as country mosquitoes — 
now may also be regarded as para-domestic, coming onto lawns, porches, 
and even inside dwellings to obtain blood meals. Residents inadvertently 
aggravate this situation by watering lawns and providing favorable condi- 



246 Indiana Academy of Science 

tions for mosquito harborage in their lush, ornamental shrubs. Residents 
who are subjected to this kind of mosquito annoyance often strongly 
assert there is no standing water in their entire neighborhood. Usually 
this is not the case. On the other hand, there is insufficient information on 
flight range and dispersal habits of many para-domestic species. Until 
such evidence is obtained, one should not rule out the possibility that 
such mosquitoes might have travelled several miles from production 
sites. 



Agricultural Practices 

Agricultural practices, also, can alter species composition. Draining 
and deforestation are examples. 

Decatur County offers good examples of the effects of drainage on 
mosquito production. Much of the land in this county is wet soil, and 
conceivably was well infested with mosquitoes when it was in native 
forest. Now, only remnants of forest remain, mostly as drained woodlots. 
In some cases, the drainage leads into farm ponds. Farms are well tiled 
and crop production is high. The few depressions found positive for mos- 
quito larvae in these wooded tracts in 1969 were producing Aedes vexans 
and A. canadensis. These mosquitoes develop more rapidly, and in more 
temporary type pools than A. stimulants, which probably was present, if 
not dominant, in the native forests of Decatur County. 

Shelby County, also, shows effects of drainage and deforestation. 
Much of the land is in cultivation and few forest tracts remain. Aedes 
stimulans was collected in Meltzer Woods in association with A. vexans 
and A. canadensis, and as a single species in a woodlot in the southern 
part of the county. Mosquitoes in the rural parts of this county appear 
to have been largely eliminated through deforestation and draining. 
Psorophora and summer Aedes may be produced in sites where 
drainage is blocked. Thus, both Decatur and Shelby Counties afford 
examples of alteration of species composition through agricultural 
practices. 

Discussion 

It is apparent that species composition of mosquito populations in 
any given locality in Indiana is influenced by a complex of many 
factors. Superimposed upon such location-dependent factors as climate, 
natural features and soil are the effects of urbanization and agricultural 
practices. 

A characteristic pattern of species succession in rural areas is as 
follows: a rather explosive peak of univoltine Aedes occurs in late April 
or early May. This is shortly followed by a succession of overlapping 
peaks of multivoltine Aedes and Psorophora, the occurrences of these 
peaks varying from year to year depending upon patterns of precipita- 
tion. These peaks are concurrent with steady but sustained populations 
of such permanent water breeders as Mansonia and Anopheles. 



Entomology 247 

This pattern is likely to change quite markedly with urbanization. 
The univoltine Aedes are drastically reduced or entirely eliminated. Some 
of the multivoltine, para-domestic species are able to survive, especially 
those which are versatile in utilizing such sites as blocked drainage ditches 
or artificial containers. There is likely to be a domestic mosquito problem 
unless municipal ordinances requiring solid waste and liquid waste dis- 
posal keep apace with the increase in urban populations. 

Thus, a result of severe environmental disturbance associated with 
construction of streets, highways, shopping centers, schools, housing 
developments, factories and other trappings of urbanization is reduction 
in the number of species and increase in the numbers of members of a 
few species. The effect is similar to that observed in streams. As pollu- 
tion increases, diversity of species decreases until through selection all 
that remain are those which tolerate pollution. Representatives of these 
few species are in much greater abundance than previously. 

Can any predictions be made regarding the picture as a whole for 
Indiana? If current trends continue toward movement of human popula- 
tions from rural to urban areas, and increased use of land for crop 
production, one might predict that, for mosquitoes: 1) the number of 
species will tend to decrease; 2) that reduction will most likely occur in 
numbers, if not in species of sylvan mosquitoes and permanent water 
breeders; 3) the numbers of specimens of domestic mosquitoes likely 
will increase, with attendant likelihood of transmission of diseases com- 
monly associated with domestic mosquitoes; 4) that practices which 
preserve or maintain the undisturbed environment will tend to dampen 
the foregoing effects. 

The day is rapidly approaching when communities in Indiana will 
want organized mosquito control. Oddly enough, the recognition of this 
need is contemporary with increasing sensitivity to the use of pesticides. 
It should be recognized that any method of control — especially such a 
sophisticated method as radio-sterilized male techniques, or chromosomal 
incompatibility, or biological control — can only succeed in a given locality 
if species composition in that particular locality is known and at least 
partly understood. The factors mentioned, and undoubtedly others, are 
worthy of studies in depth. 

Acknowledgments 

Thanks are extended to A. A. Lindsey, H. P. Ulrich and A. L. 
Zachary of Purdue University for information received. Also, apprecia- 
tion is expressed to Keith Huffman and John Hart of the Soil Conserva- 
tion Service for assistance and suggestions. 

Literature Cited 

1. Carpenter, Stanley J. 1968. Review of recent literature on mosquitoes of North 
America. Calif. Vector Views 15(8) -.11-97. 

2. Carpenter, Stanley J., and Walter J. LaCasse. 1955. Mosquitoes of North 
America (north of Mexico). University of California, Berkeley and Los Angeles. 360 p. 



248 Indiana Academy of Science 

3. Christensen, G. R., and F. C. Harmston. 1944. A preliminary list of the mos- 
quitoes of Indiana. J. Econ. Entomol. 37:110-111. 

4. Hamilton, Max. 1962. Wetlands of Steuben County. Indiana Department of 
Conservation, Indianapolis. 

5. Newhouse, V. F., and R. E. Siverly. 1966. St. Louis encephalitis virus from mos- 
quitoes in southwestern Indiana, 1964. J. of Med. Entomol. 3(3-4) :340-42. 

6. Schall, Lawrence A. 1966. Climate, p. 156-170. In A. A. Lindsey [ed.] Natural 
Features of Indiana. Indiana Acad, of Sci. Sesquicentennial Volume, Indianapolis. 
600 p. 

7. Siverly, R. E. 1964. Occurrence of Wyeomyia smithii in Indiana. Proc. Indiana 
Acad, of Sci. 73:144-145. 

8. Siverly, R. E. 1966. Mosquitoes of Delaware County, Indiana. Mosquito News. 
26(2) :221-229. 

9. Siverly, R. E., and G. R. DeFoliart. 1968. Mosquito studies in northern Wisconsin 
II. Light trapping studies. Mosquito News. 28(2) : 162-167. 

10. Wayne, William J. 1966. Ice and Land, p. 21-39. In A. A. Lindsey [ed.] Natural 
Features of Indiana. Indiana Acad, of Sci. Sesquicentennial Volume, Indianapolis. 
600 p. 



A Check List of Indiana Collembola 
John W. Hart, Earlham College 



Abstract 

Several entomologists have studied Indiana Collembola. Most of their findings have 
not been published. This checklist includes all species known from the state at this time. 

Hoosier entomologists have shown little interest in the Collembolan 
insects. Eight species were known from the state when Wilkey (15) 
reported 21 new records in his taxonomic study of a small area in central 
western Indiana. Pedigo (8, 9, 10, 11, 12) contributed substantially to 
both the bionomic and taxonomic knowledge of the Collembolan fauna. 

Springtails are flightless insects and restricted for the most part to 
a life on or in damp soil, under bark, or on water surfaces. In a few 
instances they are found on growing plants. Only a few different kinds of 
habitats, involving a limited number of soil types and a small geographi- 
cal area, were investigated in finding the 69 species and forms and 40 
genera reported herein. Far more study is indicated if we are to know 
the true distribution of this order in Indiana. 

Considering the multitude of possibilities for introduction of Col- 
lembola from other states (or even other nations) one may conclude that 
given satisfactory growing conditions, any species might be found in 
almost any place. In fact, introduction is often effected in a package of 
"home soil" making survival quite probable. 

Scott (14), in his study of springtails of New Mexico, reported 28 
of the species and forms noted in this check list. Of the 28 reported by 
Scott (14), 26 were also found in New York by Maynard (5). When 
Maynard (5) made his study, 63 of the species listed in this paper had 
already been described. He found 47 of them in New York. Mills (6) 
studied Iowa springtails at a time when only 59 of the 69 Collembola 
we now know from Indiana were recognized. He identified 49 of them as 
occurring in Iowa. 

There is a close relationship between some species of springtails 
and the soil. These have interesting possibilities as indicator species. 
Environmental disturbance by pollution with wastes or chemicals (includ- 
ing insecticides) could be reflected by variations in densities of one or 
more species or by variation in composition of Collembolan populations. 
There appears to be much of both importance and interest yet to be 
studied. 



249 



Checklist of Indiana Collembola 



Mesaphorura clavata (Mills 
granulata (Mills), 



System of classification, Salmon, 1964 

Order Collembola 

Suborder Arthropleona 

Superfamily Hypogastruroidea 

Family Onychiuridae 

Subfamily Tullbergiinae 

3), 1934 

1934 



Hart, 1969 
Hart, 1969 



Subfamily Onychiurinae 
Hymenaphorura subtenuis (Folsom), 1917 

Paronychiurus ramosus (Folsom), 1917 

Protaphorura encarpata (Denis), 1931 

Family Hypogastruridae 
Xenylla humicola (Fabricius), 1780 

Hypogastrura armata (Nicolet), 1841 
lucifuga (Packard), 1888 
matura (Folsom), 1916 
packardi (Folsom), 1902 

Suborder Neoarthropleona 
Family Brachystomellidae 
Brachystomella stachi Mills, 1934 

Family Anuridae 
Aphoromma granaria (Nicolet), 1847 

Friesea claviseta Axelson, 1900 

Family Neanuridae 
Pscudachorutes aureo-fasciatus (Harvey), 1898 
saxatilis Macnamera, 1920 

Neanura barberi (Handschin), 1928 

Superfamily Entomobryoidea 
Family Tomoceridae 
Maynardia elongata (Maynard), 1951 

Pogonognathellua flavescens (Tullberg), 1871 

Family Isotomidae 
Subfamily Proisotominae 
Folsomia fimetaria (Linne), 1758 

quadrioculata (Tullberg), 1871 

Proi80toma minuta (Tullberg), 1871 

Subfamily Isotominae 
Isotomurus palustris (Miiller), 1776 
palustroides Folsom, 1937 

Pscudisotoma sensibilis (Tullberg), 1871 

Isototniella minor (Schaffer), 1896 

Isotomina constricta (Folsom), 1937 

Isotoma trispinata MacGillivray, 1896 

violacea form caerideatra (Guthrie), 1903 
viridis form catena (Guthrie), 1903 

Heteroisotoma andrci (Mills), 1934 

Vertagopus arborea (De Geer), 1740 
cincrca (Nicolet), 1842 



Hart, 1969 

Wilkey, 1950 

Hart, 1969 

Pedigo, 1969b 

Wilkey, 1950 

Packard, 1888 

Mills, 1934 

Wilkey, 1950 



Pedigo, 1969b 

Hart, 1969 
Hart, 1969 

Pedigo, 1969a 
Pedigo, 1969b 

Hart, 1969 



Pedigo, 1969a 
Folsom, 1913 



Wilkey, 1950 
Hart, 1969 

Pedigo, 1969b 



Mills, 1934 

Wilkey, 1950 

Wilkey, 1950 

Hart, 1969 

Hart, 1969 

Pedigo, 1969a 

Wilkey, 1950 

Folsom, 1937 

Mills, 1969 

Wilkey, 1950 

Wilkey, 1950 



250 



Entomology 



251 



Family Entomobryidae 
Subfamily Entomobryinae 
Orchesella ainsliei Folsom, 1924 

hexfasciata (Harvey), 1895 
zebra Guthrie, 1903 

Parasinella cavernarum (Packard), 1888 

Entomobrya assuta Folsom, 1924 

marginata (Tullberg), 1871 
nivalis (Linne), 1758 

Entomobryoides purpurascens (Packard), 1873 

Pseudosinella petterseni Borner, 1901 
rolfsi Mills, 1932 
violenta (Folsom), 1924 

Lepidocyrtus curvicollis Bourlet, 1839 
cyaneus Tullberg, 1871 
cyaneus form cinercus Folsom, 1924 
lanuginosus (Gmelin), 1788 

(near) jmllidus Reuter, 1890 
paradoxus Uzel, 1890 

Salina banksii MacGillivray, 1894 



Subfamily Paronellinae 



Suborder Metaxypleona 
Family Poduridae 



Podura aquatica Linne, 1758 



Suborder Symphypleona 
Family Sminthuridae 
Subfamily Sminthuridinae 
Sminthurides hyogramme Pedigo, 1966 
lepus Mills, 1934 
malmgreni (Tullberg), 1876 
pseudassimilis Stach, 1956 

Sphaeridia pumilis (Krausbauer) , 1898 

Subfamily Sminthurinae 
Tribe Katiannini 
Sminthurnius aureus (Lubbock), 1862 
elegans (Fitch), 1863 
similitortus Maynard, 1951 



Tribe Sminthui 



Sminthurus mcdialis Mills, 1934 
trilineatus Banks, 1903 



Sphyrotheca minnesotensis (Guthrie), 1903 

Tribe Bourletiellini 
Katiannina macgillivray (Banks), 1897 

Bourletiella hortensis (Fitch), 1863 
millsi Pedigo, 1968 

Pseudobourletiella chandleri Pedigo, 1968 

Deuterosminthurus yumanensis Wray, 1967 

Subfamily Dicyrtominae 
Ptenothrix marmorata (Packard), 1873 
unicolor (Harvey), 1893 



Wilkey, 1950 
Wilkey, 1950 
Wilkey, 1950 

Packard, 1888 

Wilkey, 1950 
Wilkey, 1950 
Wilkey, 1950 

Wilkey, 1950 

Pedigo, 1969a 

Wilkey, 1950 
Wilkey, 1950 

Pedigo, 1969a 

Pedigo, 1969a 

Pedigo, 1967 

Pedigo, 1969a 

Pedigo, 1969b 
Pedigo, 1969b 



Pedigo, 1968 



Folsom, 1916 



Pedigo, 1966 
Pedigo, 1969a 
Pedigo, 1969b 
Pedigo, 1969a 

Pedigo, 1969a 



Wilkey, 1950 
Pedigo, 1969a 
Pedigo, 1969a 

Pedigo, 1969a 
Pedigo, 1966 

Pedigo, 1969a 

Wilkey, 1950 

Gould, 1945 
Pedigo, 1968 

Pedigo, 1968 

Wray, 1967 

Pedigo, 1969a 
Wilkey, 1950 



252 Indiana Academy of Science 



Literature Cited 

1. Folsom, J. W. 1913. North American springtails of the subfamily Tomocerinae. 
Proc. U. S. Nat. Mus. 46 :460. 

2. — . 1916. North American Collembolous insects of the subfamilies Achoru- 



tinae, Neanurinae, and Podurinae. Proc. U. S. Nat. Mus. 50:515. 

3. - — . 1937. Nearctic Collembola or springtails of the family Isotomidae. Bull. 
U. S. Nat. Mus. 168:1-472. 

4. Could, G. E. 1945. Insect pests of cucurbit crops in Indiana. Proc. Indiana Acad. 
Sci. 53:169-170. 

5. Maynard, E. A. 1951. A Monograph of the Collembola or Springtail Insects of 
New York State. Comstock Publishing Co., Ithaca, N.Y. 388 p. 

6. Mills, H. B. 1934. A Monograph of the Collembola of Iowa. Collegiate Press, 
Ames, Iowa. 

7. Packard, A. S. 1888. The cave fauna of North America. Mem. Nat. Acad. Sci. 
4:1-156. 

8. Pedigo, L. P. 1966. A new Sminthurid from northwestern Indiana with a rede- 
scription of Sminthurus trilrneatus Banks. (Collembola: Sminthuridae) . J. Kansas 
Entomol. Soc. 39(1) :90-98. 

9. . 1967. Selected life history phenomena of Lcpidocyrtus cyancus f. cincrcus 

Folsom with reference to grooming and the role of the collophore (Collembola: 
Entomobryidae) Entomol. News 78 (10) :263-267. 

10. . 1968. Pond shore Collembola: a redescription of Salina bankaii Mac- 



Cillivray (Entomobryidae) and new Sminthuridae. J. Kansas Entomol. Soc. 
41(4) :548-556. 

11. - . 1969a. Activity and local distribution of surface-active Collembola (In- 

secta) : I. Woodland populations. Amer. Midland Natur. 83 :107-118. 

12. - — . 1970. Activity and local distribution of surface-active Collembola (In- 
secta) : II. Pond-shore populations. Annals Entomol. Soc. Amer. 63:753-760. 

13. Salmon, John T. 1964. An index to the Collembola. Bull. 7, Royal Soc. New Zeal. 
651 p. 

14. Scott, H. G. 1960-1965. The Collembola of New Mexico, I-III, V-XII and XIV-XV. 
Entomol. News 71 through 76. 

15. Wilkey, R. F. 1950. Collembola of Tippecanoe and surrounding counties. Unpub- 
lished B.S. Thesis, Purdue University. 

16. WRAY, D. L. 1967. Some new North American Collembola. Entomol. News 
78(3) :53-62. 



GEOLOGY AND GEOGRAPHY 

Chairman: W. N. Melhorn, Purdue University 
Robert Drummond, Indiana State University, was elected Chairman 

for 1970 



ABSTRACTS 

Faunal Assemblage Study of the Maryland Miocene. C. Tom Statton, 
Hanover College. — The location is approximately eight miles north of the 
mouth of the Patuxent River on the shore of the Chesapeake Bay. The 
area is known as the coastal plain, and the study was that of the faunal 
assemblages of the Miocene deposits. Emergence and submergence of the 
coastal plain eastward of the fall line are recorded by subsequent 
advances and retreating of the Miocene seas. The three Miocene seas 
under study resulted in the Calvert, Choptank, and St. Mary's Forma- 
tions. Included in this study are faunal lists, block counts, and measured 
sections of these sediments. Determination of the boundaries of the 
Miocene deposits are either shown in measured section or described as to 
the nature of the boundary, and should be readily apparent. The zonation 
of the Miocene sediments is rather arbitrary, based upon interpretive 
analysis of the earlier zonation set forth by Shattuck in 1904. Following 
the presentation of field exploration, some interpretive data dealing with 
ecology and climatology are presented. This is an attempt at conclusion 
analysis of data. 

Urbanization in Asia. Akhtar Husain Siddiqi, Indiana State University. 
— The study brings out the comparative degree of urbanization among the 
Asian countries. Briefly reviewing the historical background of urbaniza- 
tion in Asia, in the light of the economic growth of the Asian countries, 
the current (1950-60) structure of the urban population was discussed. 
Finally, with the help of the component analysis, an attempt was made 
to measure the levels of urbanization in Asia. 



OTHER PAPER READ 

Fact and Fancy. Mary E. Cedars, Roosevelt High School, Kokomo, 
Indiana. 



25:? 



Acritarchs (Leiosphaeridia) in the 
New Albany Shale of Southern Indiana 

Roger F. Boneham, Indiana University at Kokomo 

Abstract 

The Devonian-Mississippian New Albany Shale of southern Indiana is one of the 
numerous black shale facies in central and eastern North America. The shale is 
divided into five lithologically distinct members. These members contain varying per- 
centages of two acritarchs Leiosphaeridia plicata Felix and L. linebacki sp. n. The mean 
percentages of these two species are sufficiently different in the uppermost Clegg Creek 
and Camp Run Members to separate them from the lower members. Leiosphaeridia may 
be a useful genus for stratigraphic separation of other black shales. 

This study was made to determine whether the green alga Tasmanites 
and the acritarch Leiosphaeridia (of unknown biological affinity) might 
be used to differentiate the various members of the New Albany Shale in 
Indiana. 

The New Albany Shale was originally named by Borden (2) from 
exposures along the Ohio River in Floyd County. The lithology is charac- 
teristic of the Devonian-Mississippian dark shales which outcrop in many 
parts of the eastern United States and Ontario. 

The New Albany Shale contains intermittent layers of dolomite and 
occasional silty layers high in quartz. The formation contains pyrite 
scattered throughout its layers although the pyrite, for the most part, is 
minute crystals seldom being larger than a few millimeters. The shale has 
two aspects: a brownish-black to black carbon-rich facies and a greenish- 
gray to gray carbon-poor facies. In the area of southern Indiana 
from which the samples of this report came, the brownish-black to black 
carbon-rich facies is the predominant one. 

The New Albany Shale is continuous over much of north-central 
United States. Its equivalents in Ohio are the Huron, Ohio, Cleveland, 
Bedford and Sunbury shales and in Michigan the Antrim and Ellsworth 
shales (4). 

Campbell (3) divided the New Albany Shale into a number of forma- 
tions and members. A recent study by Lineback (15) has subdivided the 
New Albany Shale into five members. At the top of the uppermost mem- 
ber (Clegg Creek), Lineback has four beds which are only recognizable 
locally. 

The New Albany Shale contains Tasmanites and Leiosphaeridia 
specimens scattered throughout its various members (3, 15). The genus 
Tasmanites has been enigmatic for many years being classified as worm 
eggs, trilobite eggs, land plant spores, hystrichospheres, algal spores, and 
green algae. I believe that Wall (19) has offered convincing arguments 
for placing Tasmanites in the class Chlorophyceae as single cell algae, 
The same diverse origins have been proposed for Leiosphaeridia as for 
Tasmanites. Indeed, Leiosphaeridia was not recognized as a distinct genus 

254 



Geology and Geography 



255 





New Providence Shale 


Lower Mississippian 


Rockford Limestone 




Clegg Creek Member 








Upper Devonian 


Camp Run Member 


OS 

m 


Morgan Trail Member 


< 




Selmier Member 






Blocher Member 


£ 






Middle Devonian 


North Vernon Limestone 





Figure 1. Column showing stratigraphic position of the Neiv Albany Shale and its 
members in southern Indiana. 



until fairly recently (8). Eisenack (8) placed Leiosphaeridia with the 
hystrichospheres while others (6, 7, 10, 16) have placed the genus in the 
group Acritarcha (a group of unknown and possible diverse biological 
affinities). 

The Devonian-Mississippian boundary is apparently present near 
the top of the New Albany Shale. Huddle (12) thought the uppermost 
strata might be Mississippian from his study of the conodont faunas. 
Read and Campbell (17) placed the whole formation within the 
Devonian. However, Campbell later (3) decided that the uppermost strata 
contained a fauna of Kinderhook age. Cross and Hoskins (5) also placed 
the Devonian-Mississippian boundary at the uppermost strata of the New 
Albany Shale. 

Excellent discussions on the earlier reports of Tasmanites (20) and 
Leiosphaeridia (6) are readily available and need not be repeated here. 
There are two published studies of attempts to use Tasmanites for corre- 
lation of rock units. The first study (13) was done on subsurface samples 
from the Williston Basin of southwestern Manitoba. I did the second study 
(1). My attempt was unsuccessful mainly because the correlation was 
attempted over too large an area using too many formations. 

This paper is the first attempt to use Leiosphaeridia, as a strati- 
graphic marker. The New Albany Shale contains both Tasmayiites and 
Leiosphaeridia but the number of Leiosphaeridia specimens is far greater 
than the Tasmaiiites specimens. In this study I found that the most sig- 
nificant results were obtained by concentrating on the two most common 
species, Leiosphaeridia plicata and L. linebacki, and excluding all other 
specimens. Since the other specimens in all cases totaled less than 2% of 



256 



Indiana Academy of Science 



the entire count, I do not believe their inclusion in this paper would have 
altered my final conclusions. 

In this study, I used only one formation in a relatively small area 
(Fig. 2). I was fortunate to have the outcrop localities of Lineback who 
has spent much time in the field studying and mapping the New Albany 
Shale (14, 15). 




I oVernon 

2 



JEFFERSON 



k 6 Han 



? CLARK 

Henry- ° 

ville £ 




9 oSellersburg <J 




FLOYD 

New 

Albany y £ 

II ^L ? 

•Jefferson- 

Ville N 




J L 



25 



MILES 



Figure 2. Map of southern Indiana showing collection localities. 



Geology and Geography 257 

Section Localities 

Section 1 (Lineback, (14) sec. 23). Stream bank, Six-Mile Creek, T6N, 
R7E, sec. 11, NW *4, Jennings Co. Morgan Trail Mbr. (top) 3 m brownish- 
black, fissile shale. Selmier Mbr. 2 m olive-gray to greenish-gray, blocky 
shale containing dolomitic septarian concretions up to 1 m in diameter. 

Section 2 (Lineback (14) sec. 18). Road cut, State Highway 3, 1 mile 
south of Vernon, T6N, R8E, sec. 11, SE 1 ^, NW 1 /*, Jennings Co. Blocher 
Mbr. 2 m brownish-black, fissile shale interbedded with thin, grayish 
dolomitic layers. 

Section 3 (Lineback (14) sec. 19). Road cut, State Highway 3, iy 2 
miles south of Vernon, T6N, R8E, sec. 14, NE % , NW %, Jennings Co. 
Selmier Mbr. 2 m greenish-gray, blocky shale. 

Section 4 (Lineback (14) sec. 20). Stream bank and road cut along 
Quick Creek, T4N, R7E, sec. 15, NE hi, NE %, Scott Co. Selmier Mbr. 
(top) 1 m dark greenish-gray, blocky shale. Blocher Mbr. 2 m brownish- 
black, fissile shale. 

Section 5 (Lineback (14) sec. 1). Along State Highway 56 beginning 
at bridge in T3N, R8E, sec. 9, SW X A, SW V±, thence along north line sec. 
16, NW *4, Jefferson and Scott Cos. Camp Run Mbr. 3 m brownish-black 
to grayish-black, fissile to blocky shale. 

Section 6 (Lineback (14) sec. 11). Standard Materials Quarry, 4 miles 
west of Hanover, T3N, R9E, sec. 16, SW &, NE %, Jefferson Co. Morgan 
Trail Mbr. (top) 1 m brownish-black, fissile shale. Selmier Mbr. 1 m 
greenish-gray, blocky shale. Blocher Mbr. 1 m brownish-black to greenish- 
gray, fissile shale. 

Section 7 (Lineback (14) sec. 9). Road cut State Highway 203, 1 mile 
northwest of Lexington, T3N, R8E, sec. 33, NW %, NE %, Scott Co. 
Morgan Trail Mbr. 2 m brownish-black, thinly bedded shale. 

Section 8 (Lineback (14) sec. 7). Road cut State Highway 160, 
3 miles southeast of Henryville, Clark Grant, lot 223, N %, Clark Co. 
Clegg Creek Mbr. 5 m brownish-black, fissile shale. 

Section 9 (Lineback (14) sec. 3). Road cut State Highway 31-W, just 
west of Sellersburg, Clark Grant, lot 110, center southwest line, Clark Co. 
Clegg Creek Mbr. 6 m brownish-black to greenish-gray, fissile shale. 
Lineback (14) considers the lower 4 m to be the Camp Run Mbr. but the 
Leiosphaeridla linebacki/L. plicata ratio indicates the lower part is Clegg 
Creek (Table 1). 

Section 10 (Lineback (14) sec. 13). Slate Run just west of Blackiston 
Mill at north edge of New Albany, Clark Grant, lot 63, Floyd Co. Camp 
Run Mbr. SV2 m brownish-black to olive-gray, fissile shale. 

Section 11 (Lineback (14) sec. 14). Falling Run, just below bridge to 
Silver Hills, west side of New Albany, T3S, R6E, Sec. 3, NW %, SE %, 
Floyd Co. New Providence Shale (top) 1 m greenish-gray, blocky shale. 



258 Indiana Academy of Science 



Chapel Shale io m greenish-gray, glauconitic shale. Clegg Creek, Mbr. 
1 m brownish-black to brownish-gray, fissile shale. 

Section 12 (Lineback (14) sec. 15), Atkins Quarry, north of Jefferson- 
ville, Clark Grant, lot 10, center southwest line, Clark Co. Morgan Trail 
Mbr. (top) 1 m brownish-gray, soft shale. Selmier Mbr. 1/20 m greenish- 
gray, blocky shale. Blocher Mbr. 3 m brownish-black to brownish-gray, 
soft to fissile shale. 

Section 13 (Lineback (14) sec. 8). Road cut, State Highway 160, 
V 2 mile east of Henryville, Clark Grant, lot 255, West y 2 , Clark Co. Rock- 
ford Ls. (top) 8/10 m light brownish-gray, fossiliferous limestone. Jacobs 
Chapel Bed 2/10 m greenish-gray, calcareous shale. Clegg Creek Mbr. 
3 m brownish-black, fissile shale. 

Field and Laboratory Methods 

Each outcrop was sampled vertically at 1 m intervals. The sample 
spacing was closer if a particular shale member was less than 1 m thick. 

The samples were crushed in the laboratory with a rotary crusher. 
It was necessary not to crush the material too finely since the leiospheres 
are rather large and might be damaged. A working sample of approxi- 
mately 10 g was obtained by repeatedly quartering the crushed field 
sample. 

This laboratory sample was boiled in concentrated hydrofluoric acid 
(52%) for approximately 30 minutes. The residue was centrifuged and 
washed. Then it was placed in a 2% solution of sodium hypochlorite over- 
night. The residue was again centrifuged and washed, then stained in a 
1% aqueous solution of methyl green. The Tasmanites and leiospheres 
usually did not take the stain. However, the amorphous, organic matter 
which is universally present in the shales did take the stain. The color 
contrast between the green-stained amorphous material and the light 
yellow to dark orange color of the Tasmanites and leiospheres made it 
much easier to conduct specimen counts under the microscope. 

The stained material was placed in glycerine jelly and from there 
mounted on glass slides. A minimum of 200 grains were counted for 
each sample which contained fossils. The results of these counts are 
summarized in Table 1. 

Stratigraphic Separations 

Table 1 illustrates the fact that Leiosphaeridia linebachi and L. 
plicata are present in differing quantities in the various members of the 
New Albany Shale. Note that at any outcrop the L. linebacki/L. plicata 
ratio may vary, in some cases, over rather wide limits. However, when 
the specimen counts of all the exposures of a given member are averaged 
we arrive at a meaningful conclusion. Namely, that the Blocher, Selmier 
and Morgan Trail Members contain similar mean values of L. linebachi 
and L. plicata and it follows that the L. linebacki/L. plicata ratios fall 



Geology and Geography 



259 



o ^ 



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260 Indiana Academy of Science 

within similar ranges. The Camp Run and Clegg Creek Members can 
easily be separated from each other and from the underlying members 
either by their difference in means or the L. linebacki/L. plicata ratios. 

Conclusions 

The significance of the differences in the means and L. linebacki/L. 
plicata ratios of some members of the New Albany Shale is that they are 
most likely a reflection of a portion of the changing biota of the floating 
algal mat which covered areas of the sea during the deposition of the 
New Albany Shale. Some species of Leiosphaeridia do have at least 
limited stratigraphic value within the New Albany Shale and similar 
studies of other black shales may prove the value of these fossils in help- 
ing stratigraphers with some of their problems in correlating various 
black shale exposures. 

Systematic Descriptions 

Group Acritarcha Evitt 1963 

Subgroup Sphaeromorphitae 

Downie, Evitt, and Sarjeant 1963 

Genus Leiosphaeridia 

Eisenack 1958 

Leiosphaeridia plicata Felix 1965 
Figure 3, Illustrations 4 and 5 

Diagnosis: Spherical to subspherical. Diameter 100-(110-225)-245 fi 
(50 specimens measured) 96% of measured specimens are in the size 
range 110-225 fi. Wall thickness 3-7 fi. Surface laevigate with no pylome 
or pores. Wall often, but not always, folded. Lunate folds common on 
thinner walled specimens. Straight or slightly sinuous folds common on 
thicker walled specimens. 

Remarks: Felix (11) found L. plicata in Neogene age sediments of 
southern Louisiana. This would indicate an extremely long time span for 
the species. There is a possibility that Felix (11) was working with 
reworked specimens. Another possibility is that the New Albany L. 
plicata are not the same species as the Louisiana L. plicata but resemble 
each other due to convergence. Speciation within the leiospheres is difficult 
since they have so few diagnostic characters. 

Felix did note that of the hundreds of specimens he observed only 
one was larger than 200 /*. It was 245 fi. Felix gives a size range of 
120-200 fi for the Louisiana specimens. The Indiana material extends 
this range slightly. However, I do not feel the great age range or slightly 
different size range is sufficient to believe the Indiana specimens are a 
different species than L. plicata. Leiosphaeridia plicata is similar in 
many respects to Tasmanites plicatilis Boneham 1967 which I have found 
in the Upper Devonian black shales of Michigan, Ohio, and Ontario (1). 
The size range and lunate-shaped wall folds are similar. However, T. 



Geology and Geography 



2(\\ 



plicatilis has wall pores, the diagnostic feature which separates the two 
genera Tas?nanites and Leiosphaeridia. 

Leiosphaeridia linebacki sp. n. 
Figure 3, Illustrations 1-3 

Diagnosis: Spherical to subspherical. Diameter 35-(40-90)-95 /* 
(50 specimens measured) 96% of measured specimens are within the 
size range 40-90 t*. Wall thickness ca. 1 /*. Surface laevigate with no 
pylome or pores. Wall often but not always, folded. Folds straight or 
slightly sinuous. 

Remarks: Most of the specimens of L. linebacki are easily distin- 
guished from L. plicata by their size difference. However, there is some 
suggestion of an intergradational population since the largest specimens 
of L. linebacki outwardly resemble the smallest specimens of L. plicata 
if the latter have sinuous rather than lunate folds. However, there is a 
distinct size difference between the two species. L. linebacki is distin- 
guished from L. ralla Felix mainly by a size difference. L. valla, has a 
diameter of 87-100 (i and a wall thickness of 1-3 fi. L. linebacki is dis- 
tinguished from L. tenuissima Eisenack by the diameter, the latter has 
a diameter of ca. 100 p. Eisenack (9) shows figures of L. tenuissima 
that are somewhat larger than 100 /u. Also L. tenuissima has lunate folds 








FIGURE 3. 1) Leiosphaeridia linebacki, holotype, x900, IU 12129G40/2; 2) L. linebacki, 
x900, IU 12129J26/0; 3) L. linebacki, x200, IU 12129J26/0; 4) L. plicata, x 
200, IU 12130J26/3; 5) L. plicata, x200, IU 12131Y29/3. (All slides are deposited toith 
the Department of Geology, Indiana University, Bloomington. Individual figures are 
located on a given slide using an England Finder Slide.) 



262 Indiana Academy of Science 

which is not the case with L. linebacki. The size of L. linebacki is 
similar to Tasmanites decorus Boneham from the Devonian black shales 
of Michigan, Ohio and Ontario. However, T. decorus has wall pores. 
L. linebacki bears a close resemblance to Leiosphaeridia (Protoleios- 
phaeridium) major (Staplin) Downie and Sarjeant 1963. L. major 
(Staplin) (18) has a more restricted size range (55-85 /jl) and is rela- 
tively thick walled. These two differences lead me to believe that 
L. major and L. linebacki are two distinct species. 



Literature Cited 

1. Boneham, R. F. 1967. Devonian Tasmanites from Michigan, Ontario, and northern 
Ohio. Papers Michigan Acad. Sci. 52:163-173. 

2. Borden, W. W., 1874. Report of a geological survey of Clark and Floyd Counties. 
Indiana Dept. Geol. and Natur. Resources Ann. Rept. 5:133-189. 

3. Campbell, Guy. 1946. New Albany Shale. Geol. Soc. Amer. Bull. 57:829-903. 

4. Collinson, Charles. 1968. Devonian of the north-central region, United States. 
p. 933-971. In International Symposium on the Devonian System. 

5. CROSS, A. T. and J. H. Hoskins. 1951. The Devonian-Mississippian transition flora 
of east-central United States. C.R. 3 erne Congres Strat. et Geol. du Carbon., 
Heerlen: 113-122. 

6. Downie, Charles and W. A. S. Sarjeant. 1963. On the interpretation and status 
of some hystrichosphere genera. Palaeontol. 6(1) :83-96. 

7. . 1964. Bibliography and index of fossil dino-flagellates and acritarchs. 

Geol. Soc. Amer. Mem. 94:180 p. 

8. Eisenack, Alfred. 1958a. Tasmanites Newton und Leiosphaeridia n.g. als Gattungen 
der Hystrichosphaeridea. Palaeontographica. (A) 110:1-19. 

9. . 1958b. Mikrofossilien aus dem Ordovizium des Baltikums. Senckenberg. 

leth. 39 :389-405. 

10. Evitt, W. R. 1963. a discussion and proposals concerning fossil dinoflagellates, 
hystrichospheres, and acritarchs, II. Proc. Nat. Acad. Sci. 49 :298-302. 

11. Felix, C. J. 1965. Neogene Tasmanites and leiospheres from southern Louisiana, 
U.S.A. Palaeontol. 8(1) : 16-26. 

12. Huddle, J. W. 1934. Conodonts from the New Albany Shale of Indiana: Bull. 
Amer. Paleontol. 21 : 136 p. 

13. Jodry, R. L. and D. E. Campau. 1961. Small pseudochitinous and resinous micro- 
fossils: new tools for the subsurface geologist. Amer. Assn. Petrol. Geol. Bull. 
45(8) :1378-1391. 

14. Lineback, J. A. 1964. Stratigraphy and depositional environment of the New 
Albany Shale (Upper Devonian and Lower Mississippian) in Indiana. Unpublished 
Ph.D. Thesis. Indiana University, Bloomington. 136 p. 

15. . 1968. Subdivisions and depositional environments of New Albany Shale 

(Devonian-Mississippian) in Indiana. Amer. Assn. Petrol. Geol. Bull. 52:1291-1303. 

16. NORRIS, G. and W. A. S. Sarjeant. 1965. A descriptive index of genera of fossil 
Dinophyceae and Acritarcha. New Zeal. Geol. Surv. Paleontol. Bull. 40 : 72 p. 

17. Read, C. B. and Guy Campbell. 1939. Preliminary account of the New Albany Shale 
flora. Amer. Midland Natur. 21 :435-453. 

18. Staplin, F. L. 1961. Reef-controlled distribution of Devonian microplankton in 
Alberta. Palaeontol. 4(3) :392-424. 

19. Wall, David. 1962. Evidence from recent plankton regarding the biological affini- 
ties of Tasmanites Newton 1875 and Leiosphaeridia Eisenack, 1958. Geol. Mag. 
94 :353-363. 

20. Winslow, M. R. 1962. Plant spores and other microfossils from Upper Devonian 
and Lower Mississippian rocks of Ohio. U.S. Geol. Surv. Prof. Paper 364 : 93 p. 



Factors Affecting Coal Roof Rock in Sullivan County, Indiana 

Charles E. Wier, Indiana Geological Survey 



Abstract 

Maintaining- a safe and stable roof is both an economic and geologic aspect of an 
underground mine operation. Roof falls occur when the rock is too weak to support the 
overlying pressure across the open span where coal has been removed. Roof falls are 
related to characteristics of the rocks. They may be described as dust, lenticular, con- 
cretion, slate, clay squeeze and massive falls. Lithology and thickness of beds, jointing, 
strength of bedding plane bond, and the effect of moisture are important considera- 
tions. No single criterion seems to be adequate for practical roof evaluation. 

Introduction 

Probably no single aspect of underground coal mining is less under- 
stood than the evaluation of roof strength before actual mining begins. 
Certain areas of good roof and certain areas of poor roof may be recog- 
nized but the roof in most of the area is likely to be of questionable 
strength. Before mining begins the geologist can, by using cores of the 
roof rock, run physical and chemical tests that characterize the roof 
rock. During the mining operation the engineer can devise a system of 
mining that leaves approximately 50% of the coal as supporting pillars 
and a system of roof support using roof bolts, supplemented by posts, 
rails, or bars. If the roof collapses, the engineer can design a method 
of cribbing to stabilize the roof in the roof fall area. Coal mines, how- 
ever, should operate at a profit and if too much material and too many 
man hours are required for roof control this cost is more than the 
margin of profit and the mine has a deficit operation regardless of the 
efficiency of actual coal removal. The mine superintendent must see 
that the roof is supported well enough that it will not endanger the 
miners nor collapse in rooms or entries and restrict the flow of fresh air 
through the passages, or the flow of coal out on conveyor belts. On 
the other hand, his job is to keep costs down and put the minimum 
amount of money into roof control. 

In an attempt to characterize good versus poor roof condition, in- 
formation has been accumulated intermittently during the past 15 years. 
Data for this report was gathered from drill holes and from the under- 
ground workings of the Minnehaha, R. S. and K., and Thunderbird Mines, 
all in Sullivan County. 

Kinds of Roof Falls 

The roof in an underground mine collapses when the rock is too weak 
to support the overlying pressure across the area where the coal has 
been removed. The kinds of rocks are most important in an evaluation 
of the roof. Thick homogeneous beds are more competent than thin 
heterogeneous ones. In general, thick massive sandstone or limestone 
provides the best roof, and shale or mixed sand, silt, and shale beds are 
the poorest. However, a fairly homogeneous gray shale may be a satis- 

263 



264 Indiana Academy of Science 

factory roof. Vertical joints across a bed and abundant mica and carbon 
films in a bedding plane allow slippage and decrease the competence of 
the rocks. Expandable clays in the rock swell in the presence of moisture 
and cause disruptive pressures. Water in the rock also adds weight to 
the roof, tends to weaken some rocks, and lubricates the moving surface. 
A combination of the above parameters may produce a variant of the 
six different kinds of roof falls described below. 

A dust roof fall (Fig. 1) is related to the thin soft dark gray shale 
that overlies the coal in some places. This shale contains finely dissimi- 
nated pyrite, calcareous shells, and carbon films. As soon as the coal is 
removed and moist air comes in contact with the pyrite in the shale, 
the iron sulfide (pyrite) changes to an iron sulfate and swells, and the 
shale literally falls apart. Alternate moist and dry air that circulates 
through the mine may hasten the dissociation of the shale. This soft 
shale crumbles into dust and falls out between the roof bolts even if 
large metal or woods plates are used at the bottom of the bolt. Not much 
can be done about holding this part of the roof, but such falls are not 
a serious problem because this shale commonly ranges only a few inches 
to a foot in thickness and the dust that falls to the floor creates only 
a minor nusiance. 

The lenticular roof fall is related to a sandstone roll (Fig. 2). In 
this situation, the bottom surface of the sandstone is quite irregular 
and the upward concave areas are filled with dark gray to black shale 
that at that place forms the roof of the coal. This shale is fairly soft 
and is not well cemented to the sandstone. Thus when the coal is re- 
moved, this lenticular unit of shale will come down. 

Coal beds, such as the Springfield coal (V), that are overlain by 
black fissile shale (called slate by the miners) that contains ironstone 
concretions have two special kinds of roof falls — the concretion fall 
(Fig. 3) and the "slate" fall (Fig. 4). In areas where the ironstone con- 
cretions are developed they are from an inch to four feet in diameter 
and commonly occur at the top of the coal and the base of the overlying 
black shale. When coal is removed these concretions may hang down 
from the roof into the passage (Fig. 3). They are called pots or kettles 
by the miners who generally pry them out of the roof before they fall. 

In areas where the black slaty shale does not contain concretions it 
commonly makes a good roof. If a break occurs, a large slab of rock may 
pull partly loose from the roof (Fig. 4) and hang there for days before 
falling unexpectedly. 

The fifth type of fall is related to weakness through jointing or 
fracturing, in the roof rock and in the coal. This weakness is most 
obvious when it is also a clay squeeze (Fig. 5). An underclay squeeze 
occurs where the clay beneath the coal flows, probably by a combination 
of shear and plastic flowing, from beneath the coal into a vertical 
crack or joint in the coal. In some cases the clay moves through the coal 
and several feet above the coal into a joint in the roof rock. During 
mining the clay may flow in mined-out entries or rooms. Although clays 



Geology and Geography 



265 




1 — 


— 


— = 


Blnrk Shale ^^ 


H r^ H 


3 





Figure 1. Dust roof fall. 
Figure 2. Lenticular roof fall. 
Figure 3. Concretion roof fall. 

Figure 4. Slate roof fall. 

Figure 5. Clay squeeze and fall. 

Figure 6. Massive roof fall. 



266 



Indiana Academy of Science 



that readily squeeze have somewhat different physical properties than 
those that do not, the trigger that starts the process is differential 
weight on different parts of the coal bed and thus on the underlying 
clay. Thus if a clay squeeze occurs across a mined-out area, commonly 
the clay moves out from under the coal on one side of the joint lowering 
the coal and the overlying roof rock relative to the mostly undisturbed 
coal and roof rock on the other side of the joint. The roof then is con- 
siderably weakened and the probability of a roof fall is increased tre- 
mendously. 

Fractures or joints in the roof rock either in bedding planes or 
across bedding planes may be present and not easily recognized. It is 
difficult to predict the probability of a roof fall in this case. The roof 
may look the same as in Figure 1, but a massive roof fall occurs un- 
expectedly filling the entry and extending 20 or 30 feet upward, mostly 
through thin-bedded and interlaminated shale, sandstone, and siltstone. 
This material is full of weak bedding planes that consist mostly of 
carbonaceous films and mica. When the coal is removed, the weight of 
the overlying material begins to pull these bedding planes apart; the 
individual beds start breaking and form nearly vertical cracks. If 
water seeps in through the cracks, it weakens the shale further, acts 
as a lubricant to bedding plane movement, and adds weight making the 
chances of roof fall much greater. This interbedded material ordinarily 
does not contain water in the natural state. The shale beds effectively 
seal off the sand beds and lenses from each other and make the whole 
material more or less impermeable, that is, before cracking takes place. 

Two conditions are favorable for the accumulation of water: 1) a 
thick-bedded porous sandstone on top of the coal or close enough to be 
reached by the 4- to 8-foot long roof bolt holes, and 2) swags or struc- 



000 feet 




Figure 7. Cross section near the east side of Thundcrbird Mine showing variations in 
dip of the coal and in the distribution of kinds of roof rock. 



Geology and Geography 267 

tural lows in the coal. These two conditions are well illustrated at the 
east end on Main East in the now abandoned Coal VI workings of the 
Thunderbird Mine (Fig. 7). Where a porous sandstone rests on top of 
the coal (and may cut into the coal) it likely will cause a water problem 
even without being* in a structural low. On the other hand a structural 
low that has a less porous laminated sandstone may allow the roof to 
be saturated with water so that it becomes quite heavy. Water may 
weaken the bonding ability of the clay and increase the probability of 
collapse. Another minor factor in some roof falls may be gas pressure. 
At places where mining progresses up hill, gas in the top of the coal 
and in the roof may exert some lateral pressure on the roof where it is 
slightly downhill and where the coal has just been mined. 

Evaluation of the Rocks 

In an attempt to try to predict areas of massive roof falls in the 
Hymera coal (VI) in the Thunderbird Mine, 43 cores were studied in 
detail. The common sequence for the 20 feet of rocks above Coal VI is, 
from the top down: 1) thick-bedded sandstone, 2) laminated sandstone, 
3) laminated shale, 4) dark gray non-laminated shale, and 5) dark 
gray shale containing plant fossils and pyrite. 

1) Sandstone: light-gray, medium-grained, thick-bedded; locally may 

be replaced by siltstone, laminated sandstone, or 
laminated shale; locally cuts down into coal. 

This medium-grain, thick-bedded sandstone, should theoretically be 
an excellent roof where it is close to the coal, because it is structurally 
strong. But in some areas it carries much water and completely saturates 
the underlying laminated sandstone and shale such that they fall from 
as high as the base of the sandstone. 

2) Sandstone: light- to medium-gray, laminated with 10 to 50% 

shale; contains chlorite, mica, and carbonaceous ma- 
terial in bedding planes. 

3) Shale: dark- to medium-gray, laminated with 10 to 50% fine- 

grained sandstone; contains mica and carbonaceous ma- 
terial in bedding planes. 

The laminated shale and laminated sandstone differ mostly in 
amount of sandstone versus the amount of shale. Both units contain weak 
bedding planes composed of lineated mica flakes and carbonaceous films 
(from fossil plants). The sandstone has a greater capacity to contain 
water (more porous) and a greater capacity to absorb water (more 
permeable) through the rock if drill holes connect it to water-bearing 
rock. This water not only makes the roof heavier but tends to react 
electrochemically with some of the clays and chlorite (in shale laminae) 
in some bedding planes. The sandstone laminae act as a conduit for the 
water but remain solid, but shale laminae desintegrate. Montmorillinite 
clays that expand when wet were not found in the samples tested. Sawed 



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Indiana Academy of Science 



samples of the more clayey bands of these two units readily break into 
small pieces when immersed in water for several hours. 

4) Shale: dark-gray, not laminated with standstone; locally con- 

tains in lower part calcite or pyrite brachiopod shells 
and crinoid columnals. 

This shale, although not particularly strong, is homogeneous and 
seems to hold fairly well where it is 2 feet or more thick. The strength 
of this shale is adversely affected by water, but it has a low enough 
permeability that ordinarily, water is not a problem. 

5) Shale: dark-gray; contains abundant plant remains, some of 

which contain pyrite and some carbon (both vitrain and 
fusain). 

The shale is commonly less than a foot in thickness and is absent 
in many areas. Where present it commonly falls down on to the floor of 
the entries as small fragments or as dust. 

Physical Tests 

In the sequence of rocks discussed the key to predicting good versus 
poor roof conditions seems to be hidden in the physical and chemical 
variations of units 2 and 3, the laminated sandstone and laminated shale. 
Several tests were tried. A few look promising. 




IOOO 2000 

Modulus of Rupture 



3000 



4000 



Figure 8. Graph showing vertical variation in modulus of rupture for samples taken 
from two cores. Solid line represents core from Thundcrbird Mine area; dashed line 
from R. S. & K. Mine area. 



Geology and Geography 269 

Modulus of rupture was determined for blocks cut both from hand 
samples and from cores. In general roof rock known to be strong had 
high shearing strength and poor roof rock had low shearing strength. 
Where the first 20 feet of rock immediately above the coal was laminated 
sandstone and shale the readings were irregular. Cores were obtained 
from an area of poor roof rock in the Thunderbird Mine and from an 
area of satisfactory roof rock in the R. S. & K. Mine (Fig. 8). Modulus 
of rupture tests do not show significant difference. 

Clays were separated from the shale laminae. Illite is the most 
abundant clay mineral. X-ray patterns indicate that the 001 peak for 
illite is usually symmetrical in the area of good roof and asymmetrical 
in the area of poor roof. The asymmetry indicates a degraded illite that 
will take water readily and will swell. 

Viscosity tests were run on clays by mixing 50 g of clay with 100 ml 
of water and measuring the resistance to a rotating paddle. The viscosity 
of the clay slurry is related to internal friction and cohesion of particles. 
In general the good roof rocks had the highest viscosity, but viscosity 
varies vertically over short distances. 

Conclusions 

In evaluating roof rock conditions the geologist must look at the 
rocks from many viewpoints. No single physical or chemical parameter 
tells the whole story. Not only are lateral variations in the rocks im- 
portant but vertical variations are also. Not only may the rock be 
significantly different from one foot to the next but, in some cases, from 
one centimeter to the next. 



Proposed Origins for the Hadley Lake Depression, 
Tippecanoe County, Indiana 

Nils I. Johansen and Wilton N. Melhorn, Purdue University 



Abstract 

This paper concerns a study of an unusual linear topographic depression in north- 
western Tippecanoe County, Indiana. This depression contains Indian Creek, Hadley 
Lake, a branch of Burnett Creek, and a reach of the Wabash River. 

Four hypotheses are presented for the origin and development of the topographic 
depression : 

1. A partly abandoned glacial sluiceway. 

2. An esker trough or rinnentaler. 

3. A segment of the "original" Wabash River channel. 

4. Stream capture. 

After evaluating available evidence, it is concluded that hypotheses 1 and 4 may 
be combined and appear to provide a reasonable solution to the problem. Hypothesis 2 
is an exciting possibility and hypothesis 3 is not totally implausible. 

Additional fieldwork, particularly shallow exploration geophysical methods to de- 
termine bedrock configuration and altitude, is needed to verify one of the hypotheses, a 
combination of more than one of them, or provide evidence that none are reasonable 
explanations of the observed phenomenon. 



Introduction 

The most striking" topographic feature in Tippecanoe County is the 
Wabash River and its tributary valleys. The Wabash enters the county 
near the northeastern corner and leaves near the middle of the western 
boundary. All other streams drain to the Wabash River making it the 
local base level of erosion. Changes in gradient or level of the Wabash 
thus will change the erosive and transportive power of streams entering 
it. 

The Wabash flows, for the most part, in a wide, alluviated valley 
bounded by relatively steep, terraced valley walls (Fig. 1). The modern 
stream may be classed as "underfit" in the sense of volume of flow and 
channel width and configuration compared with total valley width. Pres- 
ent-day flow did not carve this wide valley; thus, proglacial and immediate 
postglacial flow were much greater than now. Two distinct levels of 
paired terraces along the valley sides, the Mississinewa and Maumee 
terraces of Thornbury (8) and an indistinct third terrace level, whose 
place in the late glacial history of the valley is uncertain, suggest the 
importance of the Wabash valley as a spillway for meltwater draining 
away from the retreating Wisconsinan glacier. 

But did late-glacial Wabash River, in the Lafayette vicinity, always 
follow its present-day course ? The reason for posing this question lies 
in examining the topographic maps of Tippecanoe County and neighbor- 
ing areas to the northeast and southwest. Overall, the Wabash flows from 
northeast to southwest; the river enters the county flowing southwest- 
ward, gradually changes course to south, but at Lafayette makes a 
nearly right-angle bend and flows in a nearly due west direction for 

270 



Geology and Geography 



271 




Figure 1. Oblique topographic model of the Lafayette area, Tippecanoe County, Indiana 
(After A.K.F. Turner). 



several miles. It then resumes a southwestward course along nearly the 
same vector followed when entering the county from the northeast 
(Fig. 2). 

The near right-angle bend at Lafayette is also marked by a sig- 
nificant increase in width of an already wide valley. This has been at- 
tributed to the fact that at this location the present river intersects the 
glacially-filled valleys of the preglacial Teays River and Anderson Valley, 
the latter a tributary of the ancient Teays which entered it from the 
east along the general line of present-day Wildcat Creek (9). Thus we 
have an apparently adequate explanation for the great bend of the 
Wabash, but another problem arises. It is generally accepted that the 
present drainage of the glaciated portion of Indiana is, except for 
reaches of the upper middle Wabash drainage, independent of lines of 
preglacial drainage. Why, then, should the modern Wabash intersect 
the buried Teays Valley at this particular location? 

The key to the question may lie in an explanation of an unusual 
topographic lineament, just northwest of West Lafayette, where a small 
natural lake, Hadley Lake, and several other closed, undrained depres- 



272 



Indiana Academy of Science 




sions, lie athwart the crest of the present drainage divide. Two creeks 
drain away from Hadley Lake, but the lake has no outlet. These creeks 
are Indian Creek flowing: southwestward and the northeastward flow- 
ing* Burnett Creek. These creeks also appear "underfit" in terms of cross- 
section profile versus flow, indicating that both valleys carried a greater 
flow of water in the past. 

The question now becomes, "Is the apparent alinement of Indian 
Creek, Hadley Lake, Burnett Creek, and the Wabash River significant, 



Geology and Geography 273 

and if so what is its meaning in respect to development of the modern 
Waabsh?" There may be several acceptable answers, considering the 
paucity of currently available data, but before elaborating the problem 
a brief review is required of the late glacial history of the area as now 
interpreted. 

Late Glacial History 

By late Tertiary time, bedrock subcrop data, especially abundant 
from Illinois, indicates regionally mature topography (in the Davisian 
sense). Lithologies of the region are Paleozoic sedimentary rocks — sand- 
stones, shales, and limestones — on which polycyclic episodes of regional 
planation during the Tertiary produced smoothly rounded uplands, broad 
valleys of meandering streams, and local rock benches or straths mark- 
ing the record of the polycyclic events. The master drainage system 
was the Teays River and its tributaries. The flow direction of the Teays 
was westward to join the preglacial Mississippi River in central Illinois. 
Preglacial Wabash River was tributary to preglacial Ohio River, as 
today, but the Wabash drainage basin was much smaller than now, and 
separated from Teays drainage by a watershed divide located some- 
where south of Lafayette, probably along a line intersecting the modern 
Wabash bedrock gorge somewhere between Attica and Covington. The 
drainage of Tippecanoe County was entirely within the Teays basin. 

Multiple Pleistocene glaciations effected many topographic changes 
and drainage diversions in the northern % of Indiana. The principal 
effects of ice invasion were to smooth terrain by filling of valleys in 
existing topography with glacial till or outwash. During interglacial 
stages, progressive erosion occurred, with many streams re-establishing 
themselves in their original valleys partly filled with glacial debris. The 
present topography of the Wabash bend area, however, is principally a 
function of late Wisconsinan deposition and subsequent erosion after 
the ice departed perhaps 12,000± years ago. Minor deposition of loess 
on uplands since glacial retreat has effected small topographic modifica- 
tions but these are irrelevant to the problem at hand. 

Retreat and melting of ice produced great volumes of flood waters 
to be drained, but drainage was complicated by obstructions of various 
kinds — local ice lobation, ice-damming to create lakes, different erosional 
properties of surficial materials, and existing surface topography. Glacial 
drainage lines, or sluiceways, are commonly occupied by modern streams; 
however, these valleys are too large for the present flow volume, and 
are in this sense "underfit." This is a key factor in attempting to re- 
establish drainage systems as they existed during retreat and melting 
of the ice. Indeed, underfit streams (in the sense used in this paper) are 
common in Tippecanoe and adjacent counties. Essentially all the larger 
streams fit into this category. 

Glaciation also made broad-scale changes in drainage-basin organi- 
zation in the Midwest. Teays River disappeared and the upper Wabash 
River occupied in part, and by sequential events not yet understood, the 



274 Indiana Academy of Science 

old Teays drainage basin in Indiana. Thus Tippecanoe County now is 
totally incorporated within the Wabash River drainage system. 

In the northeastern part of the state, drainage is toward the lower 
Great Lakes, or more specifically, is part of the St. Lawrence River 
drainage basin. During later stages of Wisconsinan glaciation, there is 
evidence that part of the St. Lawrence drainage was reversed for some 
time. The eastern glacial Great Lakes used the upper Wabash River as 
an outlet, and a significant flow of water (Maumee flood) was added to 
the Wabash Valley sluiceway. 

In summation, the picture of what happened to drainage systems 
during ice melting and shortly after the glaciers disappeared from the 
region is by no means totally clear. Climatic and vegetation conditions 
are unknown or only inferred. These two factors have great significance 
in determining type and rate of erosion and landscape modification. Ero- 
sion and/or change in climate and vegetation may obscure or destroy the 
evidence of former location of a stream, and we must accept the fact 
that a stream is where it is for no good reason that we can ascertain. 
The stream is where it is because it started out in this location for 
reasons not now apparent. These unknowns probably explain why many 
areas in Indiana look peculiar to the geomorphologist or airphoto inter- 
preter; they can not be fully explained, and look more like a "geo- 
morphic happening" than something that developed by rational ordering. 
But the reasons are there. We have not yet found them. 

Previous Work 

Tippecanoe County and the great bend of the Wabash appears to 
have been a popular area for geographic field investigations in the 
early part of this century. McBeth (5) investigated many features re- 
sulting from glaciation, as well as development of the Wabash drainage 
system, but his papers make no reference to a Hadley Lake-Wabash River 
lineament; very probably it escaped his notice because it is not a topo- 
graphically prominent feature as compared to eskers and terraces, and 
of course he lacked topographic maps or air photos on which the linea- 
ment immediately attracts attention. F. J. Breeze (1) apparently recog- 
nized the peculiar relations of Burnett Creek and Indian Creek in a 
paper presented before the Indiana Academy in 1916 but no abstract or 
paper was published and thus details of his observations or interpreta- 
tions are lost. The early report on Tippecanoe County by Gorby (3) 
likewise fails to mention the Hadley Lake depression. Gorby's report is 
not particularly admirable even by geologic standards of his time, though 
his statement that ". . . The assertion may be ventured that no county 
in the Union affords a more varied exhibit of this puzzling deposit (i.e., 
the Glacial drift) than Tippecanoe . . ." might draw plaudits from 
modern workers. Leverett's classic work (4) also lacks mention of the 
Hadley Lake lake and trench. 

No further geomorphic work was done in the Tippecanoe region 
until recent years. Schneider and others (6) describe three sets of linear 



Geology and Geography 275 

features in western Indiana, which consist of straight, shallow troughs 
and discontinuous low ridges or elongated knobs of sand and gravel. They 
relate these linear features (one of which is Hadley Lake trough) to sub- 
parallel northeast-southwest drainage developed in accord with ice move- 
ment from the northeast. West and Barr (11) describe groundwater prob- 
lems in the area immediately west of Hadley Lake and infer a substantial 
thickening of glacial drift eastward into the depression. Schneider 
and Wayne (7) suggest that the depression is an esker-trough that is 
a downstream continuation of that segment of the Wabash River 
extending from near Lafayette upstream to Georgetown in Cass County. 

We are now in a position to again ask the question, "Is the aline- 
ment of Burnett Creek, Indian Creek, and Hadley Lake significant, and 
in what relation to the modern course of the Wabash River?" Let us look 
at four preliminary hypotheses in an attempt to answer this question. 

Hypotheses of Origin 

We suggest four possible hypotheses for development of this peculiar 
alinement of land forms. 

1. An abandoned glacial sluiceway 

2. An esker trough, or perhaps a rinnentaler 

3. The "original" post-glacial Wabash river channel 

4. Stream capture 

All hypotheses deal with fluvial processes as an erosional factor. 
What is not taken into consideration are bedrock topography, joint 
control, faulting (if any), and general structural geology of the area. 
As the buried extension of the Knobstone cuesta must pass almost di- 
rectly beneath the linear surface depression, bedrock cannot be elimi- 
nated as a determinant in developing a satisfactory explanation for 
surface drainage adjustments despite a rather thick covering blanket of 
glacial till or outwash. Too few bedrock data are available, however, to 
assess its importance at this time, although Wayne (10) produced a 
preliminary bedrock physiography map of the area based on limited 
information. 

Let us look at each hypothesis in some detail. 

Glacial Sluiceway 

This is the most popular local explanation of the landforms by 
geomorphologists and air photo interpreters. The assumptions behind 
this interpretation follow: 

1) The Wabash River has not changed its course. 

2) The sluiceway received large volumes of water from a 
stagnant glacier or glacier receding primarily by vertical 
ablation. 

3) The glacial source of meltwater was nearby, perhaps as close 
as the northern border of Tippecanoe County because the 



276 



Indiana Academy of Science 



sluiceway is not traced farther northeast; if it did extend 
farther, post-glacial erosion or colluviation has destroyed 
evidence of former existence. 

4) The time of origin is thus placed as late-glacial, when the 
glaciers left the area about 12,000 ± years ago, or about the 
time of formation of the Iroquois ( ? ) moraine. 

5) The sluiceway carried water for only a short time, and later 
creeks occupied the depression and began to cut new valleys. 



NW 



SE 




Typical (7) 



N.W.-S.E. Sec. 9, T.23N., R.5W. 




Near headwater© 



N.W.-S.E. Sec. 3, T. 23 N., R.5W. 



SE 




Near the Wabosh 
River @ 



N.W.-S.E. Sec. 24, T.23N., R.6W. 



Ficuke 3. Cross-section of present Indian Creek (See Fig. 2 for locations). 



Geology and Geography 277 

Indian Creek and Burnett Creek thus formed. Hadley Lake 
was left as a ponded local segment of the trench because the 
bedrock high southwest of the lake prevented free drainage 
in that direction. 

The conclusion is thus reached that the trench is essentially a 
temporary overflow distributary of the Wabash. With further retreat 
of the ice, the source of water was gone and the sluiceway was occupied 
by small, newly-formed streams. 

Cross-sections of Indian Creek valley (Fig. 3) show clearly at least 
two cycles of erosion, with a wide valley representing the sluiceway stage, 
and a smaller valley of the modern creek. Thus the sluiceway hypothesis 
appears adequate to explain the landform feature of the trench; how- 
ever, it explains neither the reason for location or alinement. The 
hypothesis fails to take into account either bedrock topography or 
correlation between this, the present Wabash Valley, and the preglacial 
Teays River Valley. The glacial sluiceway interpretation suggests that 
the landforms are independent of these latter factors and dependent only 
on an abundant source of melt-water. 

Esker Trough or Rinnentaler 

This hypothesis implies subglacial or englacial scour creating a 
bedrock trench of linear section. Leverett used the term esker trough for 
similar linear features in Montgomery County, near Muncie, and else- 
where in the state, limiting the term to linear depressions now contain- 
ing only relicit patches or knobs of sand and gravel, and as noted previ- 
ously, Schneider has continued usage of this term in describing three 
sets of linear features in west-central Indiana. 

As no relict patches remain in the Hadley Lake trench there is no 
proof that an esker was ever present. The suggestion is tenable that 
the trench is a rinnentaler (channel-valley). These valleys may be cut 
uphill beneath the ice because of pressure; terminal base-level of 
streams may raise and streams flow englacially or even superglacially 
over collapsed ice filling the rinnentaler (2) 

Acceptance of this hypothesis requires formation in a glacial rather 
than proglacial environment as suggested by the glacial sluiceway theory. 
The fact that the Hadley Lake trench cuts uphill and across a bedrock 
divide appears to favor the rinnentaler over the esker trough hypothesis 
of origin. 

"Original" Post-glacial Wabash River Channel 

This is a bold hypothesis but is worth consideration. 

1) The hypothesis is not dependent on a particular source of 
meltwater, but rather a large flow for a longer period of 
time, as could be expected during the time of glacial melt- 
ing. 

2) The hypothesis assumes a correlation between alinement of 
the landform, bedrock topography, the Teays valley, and the 
modern Wabash River. 



278 



Indiana Academy of Science 



Let us project a sequence of events which could produce the aline- 
ment of the Indian Creek-Hadley Lake-Burnett Creek system and the 
Wabash River (Fig. 4). 

When the ice retreated and the land surface was re-exposed, pre- 
existing topography was buried by glacial drift. Preglacial drainage 
channels were buried and running water commenced re-dissecting the 
new surface. The ample water available gave birth to a Wabash River 





L Wabash encounters obstruction. 



2. Downcutting ceases, possible ponding. 

Tributary eroding in unconsolidated 
debris partly filling the buried Teays 
channel. 



3. Tributary reaches the Wabash. 
Overflow with rapid downcutting. 




4. Reversal of drainage in the 
future Burnett Creek segment. 



5. Modern distribution of streams. 



Figure 4. Projected sequence of events creating the Hadley Lake trough. 



Geology and Geography 279 

flowing in almost a straight southwestern direction. The new stream was 
unaffected by buried topography; soon, however, downcutting encountered 
the bedrock "high" located southwest of present Hadley Lake. This 
produced a temporary base level; the rate of downcutting could have 
slowed upstream from the obstruction and ponding possibly took place. 
Downstream, Indian Creek continued its work of grading this segment 
to the next lower temporary base level control at some unknown point 
further down the Wabash valley. 

In time, overflow from the upstream segment may have occurred 
approximately where Tippecanoe River enters the modern Wabash. Over- 
flow may have combined with headward erosion of tributaries from the 
east and south; these tributaries in part were re-occupying an only partly 
filled segment of the old Teays Valley. Once diversion occurred, upstream 
erosion was accelerated, for the stream was now completely cutting in 
glacial drift filling the old Teays and Anderson valleys and no longer had 
to contend with the bedrock high athwart its previous course. As the 
softer fill was exposed to less overall consolidation by glaciation, it 
provided no obstruction to lateral corrasion. In time, a marked widening 
of the Wabash Valley occurred southward, in the upstream direction of 
the old Anderson Valley. 

The "original Wabash" valley was thus abandoned. The small 
streams in this valley continued slow development. Hadley Lake was 
left behind as a ponded depression blocked by the bedrock divide. Some 
factors needing verification if this hypothesis is to be supported are: 

1) The presence of lacustrine sediments in the Hadley Lake 
basin, provided the original lake existed long enough. 

2) Geophysical profiles or drill holes along the axis of the 
trough to determine depth to bedrock, gradient of the postu- 
lated river, and evidence, if any, of former terraces along 
this postulated course. 

3) Composition of soil materials along the postulated course 
to determine if they were deposited by running water or 
if the till has been reworked by fluvial action. 

There can be, admittedly, substantial objections to this hypothesis. 
Nevertheless, it deserves consideration until negative proof is forth- 
coming. 

Stream Capture 

This hypothesis describes a chain of events to explain the linear 
depression. Some fundamental assumptions used in suggesting a simple 
case of stream capture are: 

1) The Wabash River did not change its course, but did have a 
much greater flow volume. 

2) Notation of the peculiar alinement of Burnett Creek, with 
a northeastward, or opposite, direction of flow. 



280 Indiana Academy of Science 

3) The presence of a bedrock high or divide southwest of Hadley 
Lake. 
The sequence of events may have been as follows. Wabash River 
and its tributaries were carrying- a tremendous volume of meltwater, 
whereas the north fork of Burnett Creek followed the Hadley Lake- 
Indian Creek channel southwestward. As flow was obstructed by the 
bedrock high, the upper reaches of the system (Burnett Creek) lost 
most of its erosive power. Wabash River and the Indian Creek segment 
continued downcutting. Meanwhile a small tributary was eroding north- 
ward from the Wabash River in section 22, T. 24 N., R. 4 W. This 
tributary eventually reached Burnett Creek in the E V 2 section 21, T. 24 
N., R. 4 W., and at this time a more favorable outlet to the Wabash 
River was provided. Consequently, Burnett Creek reversed its flow di- 
rection and Hadley Lake was left as a local depression behind the 
bedrock dam. Indian Creek and Burnett Creek became two independent 
drainage systems, and a wide abandoned valley, or col, was left across 
the bedrock divide. Proof of this hypothesis of drainage reversal awaits 
identification of terrace deposits on Burnett Creek and determination of 
whether the gradient of the terraces correspond to a supposed flow in 
the opposite direction from that of the modern stream. 

Conclusions 

Four hypotheses have been presented to explain the origin of the 
Hadley Lake depression and related fluvially carved landforms. Two of 
these (abandoned glacial sluiceway and stream capture) may be com- 
bined and appear to offer a reasonable explanation. All hypotheses, 
however, must remain in consideration until sufficient evidence or proof 
is provided to explain the observed phenomena. 

Literature Cited 

1. Breeze, F. J. 1916. Diversion of drainage, Indian Creek to Burnett Creek. Proc. 
Indiana Acad. Sci. for 1916:49. 

2. Embleton, C, and C. A. M. King. 1968. Glacial and Periglacial Geomorphology. 
Edward Arnold, Ltd., London. 608 p. 

3. Gorby, S. S. 1886. Geology of Tippecanoe County. Indiana Dept. Geol. and Natur. 
Resources 15:61-69. 

4. Leverett, F., and F. D. Taylor. 1915. The Pleistocene of Indiana and Michigan 
and the history of the Great Lakes. U. S. Geol. Surv. Mongr. 53 :529 p. 

5. McBeth, W. A. 1901. Wabash River terraces in Tippecanoe County, Indiana. Proc. 
Indiana Acad. Sci. for 1901 :237-243. 

6. Schneider, A. F., G. H. Johnson, and W. J. Wayne. 1963. Some linear glacial 
features in west-central Indiana. Proc. Indiana Acad. Sci. 72:172-173. 

7. Schneider, A. F., and W. J. Wayne. 1968. Segmentation of the Upper Wabash 
River across Indiana. Second Ann. Meeting North-Central Section, Geol. Soc. 
Amer. Iowa City, Iowa. Program Abstr. :36-37. 

8. Thornbury, W. D. 1958. The geomorphic history of the upper Wabash Valley. 
Amer. J. Sci. 256 :449-469. 

9. Wayne, W. J. 1952. Pleistocene evolution of the Ohio and Wabash valleys. J. Geol. 
60:575-585. 

10. Wayne, W. J. 1956. Thickness of drift and bedrock physiography north of the 
Wisconsin glacial boundary. Indiana Geol. Surv. Rep. Prog. 7:70 p. 

11. West, T. R., and D. J. Barr. 1965. Economic groundwater problems encountered 
in the development of a housing area near West Lafayette, Indiana. Proc. Indiana 
Acad. Sci. 74 :259-267. 



Base Level, Lithologic and Climatic Controls of Karst Groundwater 
Zones in South-Central Indiana 

Richard L. Powell, Indiana Geological Survey- 
Abstract 

Nearly all groundwater movement with the carbonate bedrock of south-central 
Indiana has been within a karst groundwater zone that occupies the lower part of the 
vadose groundwater zone above the zone of permanent saturation (phreatic zone) or 
local base level and that is characterized by a highly fluctuating water table. 

Variations of precipitation within climatic cycles and the local relief above base 
level control the vertical range of the water table and subsequent solution within the 
karst groundwater zone during that time period corresponding to each subaerial erosion 
level. Variations in carbonate solubility, thickness of beds, and intensity of jointing 
determine the texture and shape of the integrated subterranean conduits. The volume of 
water transmitted through the karst groundwater zone and its acidity determine the 
size. 

Limestone is relatively soluble compared with dolomite and clastic sediments, but 
it is less permeable than some dolomites or sandstones. Initial permeability in limestone is 
along joints and bedding planes and varies greatly from bed to bed. Shales and silty 
carbonate units may form perched water bodies that may be breached by joints or 
erosion. Sandstone, dolomite, and intensely jointed limestone may form perched water 
bodies or aquifers within relatively impermeable limestone strata and release the water 
to less permeable, subjacent limestone. 

Introduction 

Solutionally-enlarged openings and caverns in dynamic karst ground- 
water zones carry the greatest amount of groundwater in the non-glaci- 
ated portion of south-central Indiana (Fig. 1). Base level, lithology and 
climate are the three major controls of karst groundwater zones in 
carbonate rocks of Mississippian age that underlie the Mitchell Plain 
and Crawford Upland physiographic units of south-central Indiana. 

Karst Groundwater Zone 

The karst groundwater zone is characterized by a highly fluctuating 
water table within the confines of any integrated openings, such as joints 
and bedding planes, in carbonate bedrock (13) (Fig. 2). It is above 
the zone of permanent saturation (phreatic groundwater zone) and there- 
fore includes the lower part of the zone of aeration (vadose ground- 
water zone). This zone is somewhat the same as the fluctuating water 
table zone described by numerous hydrologists, such as Finch (4) and 
Meinzer (8), and defined by Swinnerton (14) as the zone within which 
limestone caverns are developed. 

Water table fluctuations within karst terrains have been ignored 
by most theorists concerned with formulating a universal theory of 
origin of limestone caverns, even though the most intense and frequent 
water table fluctuations documented have been within karst or cavern 
areas. A few modern hydrologists argue that a true water table does 
not exist within the widely spaced openings in carbonate rocks. The fact 
that the relative grain size between the interstices, usually open joints 

281 



282 



Indiana Academy of Science 



Outcrop Area 
of Middle j 
Mississippian 
Limestones 




1 000 -i 



CRAWFORD UPLAND 
Stephensport and 



MITCHELL 
PLAIN 



600- 



200 




FIGURE 1. Generalized map and cross section of south-central Indiana showing the 
location of the Mitchell Plain and Crawford Upland and their relationship to rocks of 
M iss iss ippian ag e . 



Geology and Geography 



28.'] 




Figure 2. Idealized diagram showing the range of fluctuations of a normal toater table 
in a homogeneously permeable medium. 



and bedding planes, is measured in feet rather than in microns does not 
alter the fact that groundwater level fluctuates within the integrated 
openings. The fluctuations are commonly measurable in feet and tens 
of feet in Indiana, but rises of the water table up to a few hundred feet 
are known elsewhere. 

A fluctuating water table exists in all permeable strata. The fluctua- 
tions of the water table are caused by the difference in frequency and 
amounts of precipitation, infiltration rates, transmission rates, storage 
capacity of the bedrock, and discharge rates from the permeable strata 
or bedrock. Precipitation is variable seasonally, annually, or within cli- 
matic cycles and geologic episodes, causing corresponding fluctuations 
of the water table. The rate of infiltration is dependant upon the presence 
of exposed surface openings and is affected greatly by the type and 
density of vegetation. There may be no surface run-off from a forested 
area while from a denuded area it may be total. The transmission of 
karst groundwater is dependent upon the size, shape and sinuosity of 
integrated openings between an intake area and discharge point. The 
larger the openings, the greater the potential groundwater movement. 
The amount of groundwater movement and discharge is controlled by 
size and abundance of integrated openings, and by storage capacity within 
the fluctuating zone. The rate of groundwater discharge to surface out- 
lets is controlled by the size of the integrated openings and the hydro- 
static head that may be developed within the subterranean tributaries to 
the outlets. 

The greatest volume of groundwater movement within carbonate 
bedrock of a karst terrain occurs within the zone that is most affected 
by infiltration of acidic meteoric waters and discharge of the accumulated 
karst groundwaters. The shallow zone just above the zone of permanent 
saturation within the normal range of water table fluctuation that is 
affected by even slight infiltration transmits the greatest volume of 



284 Indiana Academy of Science 

fresh karst groundwater (Fig. 2). Therefore, it is the zone of the 
greatest amount of solution. The amount of karst groundwater trans- 
mitted above or below the normal range of the water table decreases 
proportionally to distance from the normal zone. A decrease in infiltra- 
tion results in a decrease in the amount of acidic karst groundwater, 
therefore, a significant decrease in solutional enlargement and a lower- 
ing of the water table. An increase of precipitation causing a rise of 
the water table above the zone of normal fluctuations would increase the 
hydrostatic head, rate of flow and transmitted acidic karst groundwater 
in pre-existing openings developed within the underlying zone of normal 
fluctuations. The normal range of water table fluctuations should de- 
crease to shallower limits during a climatic or geologic episode as open- 
ings within the karst groundwater zone enlarged by solution to cavernous 
routes of proportions capable of containing normal amounts of infiltration. 

Heavy precipitation or snow melt may flood integrated openings 
causing a rise in the water table, an increase in hydrostatic head, and a 
high velocity within the integrated openings with a corresponding in- 
crease in solution rates. The greater the velocity of a solvent, the 
greater the rate of solution and the greater the amount of solvent, the 
greater the amount of solution. Thus a large volume of solvent becomes 
partly saturated much more rapidly than a small volume becomes com- 
pletely saturated (6). Swinnerton (14) stated that, "A solution of low 
concentration dissolves more rapidly than a highly concentrated solution. 
Four volumes of water become one-fourth saturated with CaCO ; > ( or any 
similar solute under ordinary conditions in far less time than one volume 
becomes completely saturated." 

Recent papers indicate that groundwater in carbonate rocks which 
is nearly saturated with calcium carbonate is incapable of a significant 
amount of solution unless it changes temperature or velocity, or is 
mixed with other saturated groundwater of a different temperature (1, 
15). They have also based nearly all of their cave stream analysis on 
samples taken during normal flow conditions and none during flood stages, 
yet the latter condition fills the cavern and produces ceiling solution 
features that are cited as evidence for development below the water 
table. Analysis of groundwater taken at spring in Indiana that are 
discharging from caverns indicate that from about 150 to 300 ppm of 
carbonate and sulphate ions are removed by solution during normal to 
low flow stages (9). The concentration of the solution decreases during 
flood stages, but the great increase of volume of groundwater transmitted 
results in a net increase in solution of the carbonate bedrock. Nearly 
saturated groundwater has previously accomplished the significant amount 
of solution of which it was capable. 

Most theorists have noted that many caverns contain evidence of 
complete water filling, at least in early stages of cavern development, 
such as the presence of solution features on the upper walls and the 
ceiling of cavern passages (2). Consequently, they have based their 
theories on the presumption that the caves were formed at a time when 
the water table was stable and permanently above the zone of solution 



Geology and Geography 285 

and cavern formation (15). Solution features on the roofs of cavern 
passages are not proof of permanent saturation nor stable water table 
conditions. They prove only that the passages were filled when these 
features were developed. Stable water table conditions are probably 
non-existent. Climates are not stable, and if the water table were to 
stabilize, it would indicate that there is a lack of imbalance between 
intake and discharge within the carbonate bedrock which would suggest 
a lack of significant groundwater movement. Thus, solution features on 
cavern ceilings were most likely developed when the passages were 
water filled with moving groundwater during a temporary high position 
of the water table. 

Base Level Control 

Base level has been cited classically as a control of surface geomor- 
phic erosion cycles, as well as a control of levels at which caverns may 
develop. Ultimate base level is commonly considered as a landward 
extension of sea level, while regional or local base level is defined in 
terms of strata or materials resistant to downcutting. Base level in 
regard to cavern development has been designated as the base level 
of solution, generally referring to non-soluble rocks below the carbonate 
host. 

Base level is here defined at the top of the zone of permanent satu- 
ration (phreatic groundwater zone) or the base of the karst groundwater 
zone whenever the zone of permanent saturation was relatively stable 
during any definable episode of cavern development (Fig. 2). Such 
episodes are usually contemporary with surficial geomorphic episodes. 
Thus base level is not a flat planar surface, but rather is a gently sloping- 
surface graded in a downstream direction. Surface erosion or downcut- 
ting controls the position of the somewhat permanent water table. Sub- 
terranean tributaries do not develop significantly below this permanent 
water table, or base level of significant groundwater movement, or base 
level of significant solution. Caverns that develop at a designated base 
level may exhibit irregularities caused by differential solution of various 
carbonate or non-carbonate lithologies, but generally they reflect the 
position of the karst groundwater zone that existed when they formed. 

Perched water bodies caused by lithologic differences are not re- 
garded as base levels. However, a karst groundwater zone may become 
perched following surface rejuvenation if the surface stream cuts through 
an impervious layer or aquitard. 

Lithologic Control 

A few theoretical papers have considered groundwater flow within 
a carbonate bedrock as transmission of phreatic groundwater within a 
homogenously permeable medium hundreds of feet thick (2, 3). Stratified 
rocks, including the carbonates, are seldom of uniform lithology, thick- 
ness, porosity or permeability for more than a few feet vertically and 
a few hundred feet laterally. Thick sequences of carbonate strata are 
usually composed of various thick, medium, and thin-bedded types of 



286 Indiana Academy of Science 

limestone, shaly and sandy limestones, calcareous shales and sandstones 
and dolomites, and perhaps including non-calcareous (non-soluble) 
strata, which are all of different solubility rates. The intensity of 
jointing- within each type of lithology also varies greatly from bed to 
bed and according to the location in relation to local structural features. 
Thus, any general theory which assumes a homogeneously permeable 
carbonate bedrock for groundwater movement or cavern development, for 
other than one bed in a very small area, is ignoring the most important 
geologic problem concerned. 

Most limestones are relatively impermeable rock types, although 
most are very soluble and some have a high porosity. Most limestones 
owe their high permeability to integrated openings along bedding planes 
and joints and subsequent solutional enlargement of these openings. 
Coarse-grained and porous limestones appear to be more readily soluble 
than fine-grained or compact limestones. The more dense, compact, thin 
limestone beds of the lower part of the Sanders Group and the litho- 
graphic limestone beds of the upper part of the Blue River Group and 
some of the limestone units of the West Baden and Stephensport Groups 
(Table 1) commonly project as resistant ledges into the larger cave 
passages, or have a comparatively narrow slot dissolved through them 
if they are thick-bedded. Conversely, large cavern passages formed by 
solution are more common in thick-bedded, coarse-grained or porous 
limestones, as in the Salem Limestone Member. Most coarse-grained and 
porous limestones, even if thin-bedded, have been more extensively dis- 
solved and form recesses in the cave passages formed in strata of the 
upper part of the Blue River Group and the lower part of the Sanders 
Group. The compact, denser limestones offer less surface exposure and 
allow less penetration of solvents than more porous and coarse-grained 
limestones; therefore, the latter rocks are more easily removed by solu- 
tion. Rocks low in carbonate content are generally less soluble than 
those with a higher carbonate content. Purity of the limestones in 
respect to high calcium carbonate content is of questionable importance. 
Calcite veins, pure calcium carbonate deposited as crevice fillings, com- 
monly protrude as resistant ledges and dikes in cave passages. Caverns 
within the St. Louis Limestone Formation, which is generally a hetero- 
genous unit of argillaceous limestones, are generally unknown, except 
for small solution tubes. Care should be taken to distinguish between 
large solutionally-formed cavern passages and large passages which 
have resulted from collapse enlargement of a small solution passage 
and solutional removal of the debris at stream level. 

Dolomites are less readily soluble than many types of limestone, but 
they are an important zone of groundwater movement within the Ste. 
Genevieve Limestone because of their high inherent permeability. Sig- 
nificant amounts of solution occurs in some dolomite beds simply because 
the dolomite is the groundwater transporting medium which comes into 
contact with the acidic groundwater. Cavern passages developed in 
association with dolomite beds are known in several Indiana caverns, 
notably in Wyandotte Cave, where solution has occurred within or 



Geology and Geography 287 

Table 1. Sequence of important carbonate stratigraphic units of 
Mississippian age in south-central Indiana. 

Group Formation 

Glen Dean Limestone 
Hardinsburg Formation 
Stephensport Golconda Limestone 

Big Clifty Formation 
Beech Creek Limestone 

Elwren Formation 
Reelsville Limestone 
West Baden Sample Formation 

Beaver Bend Limestone 
Bethel Formation 



Paoli Limestone 
Blue River Ste. Genevieve Limestone 



St. Louis Limestone 



Salem Limestone 
Sanders Harrodsburg Limestone 



through dolomite beds as well as the underlying limestone beds owing 
to discharge of water from the dolomite beds into the underlying lime- 
stone (12). Very permeable porous and vuggy dolomites may project 
as ledges because the groundwaters pass through them easily, while 
some very fine-grained, loosely-cemented dolomites are easily removed, 
perhaps partly by mass wasting and mechanical erosion. Medium beds 
of dolomite in several Indiana caverns have been widened within cave 
passages more easily than adjacent limestone beds. Some thick beds of 
dolomite, in now abandoned passages, have apparently caused passage 
enlargement by exfoliation or disintegration by weathering. 

Calcareous shales that are particularly susceptible to solution occur 
stratigraphically within the upper part of the Blue River Group. These 
shales are very soluble and passages within several caves have developed 
partly by solutional removal of the shale, perhaps aided by some me- 
chanical erosion. Shales, including calcareous and non-calcareous varieties, 
are generally compact and appear to lack significant open joints; thus, 
they are commonly barriers to vertically moving groundwater. Perched 
water bodies occur on the top of the shale within its unbreached limits. 
However, once breached the perched water is discharged into the under- 
lying carbonate strata. If the relative relief between the perched water 
body on the shale and the normal water level of the karst groundwater 
in the underlying carbonate strata is sufficient and if the descending 
water is transmitted through a vertical joint or set of joints that pene- 
trates several beds, a vertical shaft or pit may result from water running 
down the walls. Progressive upstream migration of the breached edge 



288 Indiana Academy of Science 

of the shale may result in a laterally elongated shaft, or successive 
breaching at different points may cause successive abandonment of the 
former horizontal cave passage and each preceding shaft. 

Vertical solution shafts or pits are formed below any lithology such 
as shale, sandstone, limestone, or dolomite which is relatively resistant 
to solution in comparison to carbonate strata below, and which is over- 
lain with relatively permeable strata or material. The overlying perme- 
able unit serves as a perched water body reservoir for groundwater to be 
discharged through an opening in the resistant bed. An opening, such 
as a joint, through the resistant bed allows somewhat regulated discharge 
into the underlying carbonate rocks. The character of the reservoir rock 
and the opening in the resistant bed determines the rate of groundwater 
discharge into the underlying strata. Vertical shafts or pits are enlarged 
primarily by regulated flows or seepage down the walls of the vertical 
opening rather than by complete flooding and filling which is probably 
possible only in the initial small openings. Some vertical solution shafts 
are now fed directly by diverted surface drainage and they may have 
originated by this means. 

Thin sandstone beds, particularly the Popcorn Sandstone Bed at the 
base of the Paoli Limestone Member, serve as resistant beds to form 
perched water bodies within overlying limestones, discharging water 
through restricted outlets into underlying strata, commonly forming 
vertical solution shafts, or laterally elongated shafts. Perched ground- 
water within thick sandstone units, such as those within the West Baden 
and Stephensport Groups, is the prime source for the groundwater which 
develops caverns in underlying limestones (11). Groundwater movement 
from a perched water body in a permeable sandstone into a limestone 
as a means of cavern development is probably the major exception to the 
concept that most caverns are formed by groundwater solution within 
a karst groundwater zone. 

Groundwater movement, and subsequent cavern enlargement, in 
south-central Indiana is primarily along the joints and bedding planes 
in the carbonate strata. Indiana caverns commonly have developed 
parallel to the dip of the strata following sets of joints that allow the 
steepest gradient to the nearest or lowest surface outlet. Most Indiana 
cave passages trend with the regional structure which has average dips 
of about 25 feet per mile to the southwest. There are local structures, 
however, which deviate from this pattern, and caverns situated on these 
structures follow the local dip. Some caverns follow the strike of the 
structure where the karst groundwater zone within the particular cavern- 
ous strata has no outlets in a downdip direction along streams which 
dissected the cavernous strata or karst groundwater zone. 

Each bed, depending upon its lithologic type and thickness, has a 
unique pattern of joints or joint sets. Generally any particular set of 
joints does not extend into overlying or underlying strata, but some joints 
are common to several strata, and some master joints apparently extend 
through many beds of dissimilar lithologies. Thin-bedded strata are 



Geology and Geography 289 

usually more intensely jointed than thiek-bedded units. In general it 
seems that there is a joint spacing- of a particular bed that is related 
to the unit thickness and type of lithology. Although the joints from bed 
to bed may not be superposed, they do cross and this allows vertical 
groundwater movement without necessitating significant flowage along 
bedding planes. 

Multi-level cavern passages that have developed in progressive stages 
as tributaries related to adjacent surface drainage levels or geomorphic 
episodes commonly reflect the effect of joint patterns shifting from bed 
to bed. Passages formed along a master joint which dissects numerous 
beds, usually have a canyon-like appearance with wide places repre- 
senting temporary halts in solutional deepening or more soluble strata. 
In addition, passages in one stratum may form an orientation pattern 
related to somewhat different intake and discharge areas than passages 
at a different level within another stratum. Goss Cave, in Washington 
and Harrison Counties, is an excellent example. The upper levels have 
developed within one set of joints in the upper part of the Salem lime- 
stone owing to a general infiltration of meteoric water, while the lower 
level within the lower part of the Salem Limestone has developed pri- 
marily by surface run-off diverted into sinkholes in addition to water 
diverted from the upper levels. Similar subterranean diversions in 
Wyandotte Cave have been described (12). 

An important cause of the down dip development of caverns and 
groundwater movement is the dissection of permeable strata by en- 
trenched surface drainage in places down dip from intake or recharge 
areas. Gardner (5) based his "static water zone" theory of cavern de- 
velopment on the concept that dissection of permeable strata was re- 
sponsible for initiating groundwater movement and solution of caverns. 
Although much of his theory is correct, he places too much importance 
on structural control of cavern development. Other aspects which he 
mentions, such as climate, are equally important. 

Climatic Control 

Numerous authors have either suggested that the misfit streams 
in modern caverns are not large enough to have dissolved the cavern, 
and that therefore they were either formed in the past when the passages 
were water filled below a permanently higher water table (3), or they 
have been formed by diversion of surface streams to subsurface routes 
at flood stages (7) or by subterranean stream piracy (16). 

Although the streams in most Indiana caverns are now misfit, it is 
obvious that each level was essentially filled with water at times during 
its period of solutional enlargement. The thick and extensive cave sedi- 
ments present in Indiana caverns are further evidence of drastic changes 
in cave stream regimen that have taken place, particularly where the 
deposits are within abandoned upper levels which no longer contain a 
stream. Some of the passages which contain misfit streams now flood 
with even light precipitation, but most are known to flood only in- 



290 Indiana Academy of Science 

frequently during exceptionally heavy periods of precipitation. Some 
water filled caverns now exist only because they are flooded owing to 
alluvial sediments which have dammed their outlets (10). 

Three factors, misfit cave streams, abandoned cave levels and ex- 
tensive cave sediments, are proof that the carbonate bedrock area of 
south-central Indiana has been subjected to multiple cycles of wetter 
and drier conditions. Further proof of significant climatic change is 
available in the interpretation of the Pleistocene history of the immedi- 
ately adjacent areas. At least three ice sheets of the Kansan, Illinoian 
and Wisconsinan glaciations were close enough to the area to effect 
moister conditions than during the interglacial stages. These wetter 
epochs undoubtedly caused more frequent temporary filling of the solu- 
tional openings within the karst groundwater zones existing during 
each glacial episode, with solutional enlargement taking place at a 
corresponding rate. Waning of the glacial climatic conditions would 
result in a decrease of precipitation, a change to drier conditions and 
the subsequent misfit stream conditions. 

Ancient climatic changes would also have caused changes in the 
amount and type of vegetation in south-central Indiana. A dense wood- 
land cover during a wet climatic episode would have caused a lack of 
surface run-off with a corresponding increase in groundwater infiltration 
and an increase in groundwater acidity, whereas a drier climatic episode 
would cause a decrease in vegetal cover, an increase in surface run-off 
and perhaps in sedimentation in surface and subsurface drainage routes. 

Although some evidence is available to correlate certain cave levels, 
cave sediments or karst groundwater conditions with specific geologic 
events, data are generally lacking to devise an exact correlation of all 
cavern development episodes with known geologic events. Data available 
at this time suggest that episodes of cavern solution and deposition per- 
haps are the only evidence of geologic or climatic episodes which are yet 
to be detected as surface geomorphic features. That is, evidences of 
past climates and geomorphic events may perhaps be best interpreted 
from evidence obtained in caverns where the evidence has not been de- 
stroyed by subsequent events. Multiple cavern levels in south-central 
Indiana and the cave sediments deposited within them appear to record 
a sequence of events ranging in age from late Tertiary to Recent times. 
Each of the cavern levels was obviously initially formed by solution within 
a karst groundwater zone and perhaps later filled with cave sediments 
as each succeeding cave level or karst groundwater zone became re- 
established at another level, higher or lower than the preceding level, 
owing to changes in base level and climate. 



Geology and Geography 291 

Literature Cited 

1. BOGLI, A. 1965. The role of corrosion by mixed water in cave forming. Problems of 
the Speleological Research. Prague, p. 125-131. 

2. Bretz, J. H. 1942. Vadose and phreatic features of limestone caverns. J. Geol. 
50:675-811. 

3. Davis, Wm. M. 1930. Origin of limestone caverns. Bull. Geol. Soc. Amer. 41 :475-628. 

4. Finch, J. W. 1904. The circulation of underground aqueous solutions. Proc. Colo. 
Sci. Soc. 7:193-252. 

5. Gardner, J. H. 1935. Origin and development of limestone caverns. Bull. Geol. Soc. 
Amer. 46:1255-1274. 

6. Kaye, C. A. 1957. The effect of solvent motion on limestone solution. J. Geol. 
65:35-46. 

7. Malott, C. A. 1938. Invasion theory of cavern development (Abstract) p. 323. In 
Geol. Soc. Amer. Proc. for 1937. 

8. Meinzer, O. E. 1923. Outline of groundwater hydrology, with definitions. U. S. 
Geol. Surv. Water-Supply Paper 494. 71 p. 

9. Powell, R. L. 1961. A geography of the springs of Indiana. Unpublished M.A. 
Thesis. Indiana University. 

10. . 1963. Alluviated cave springs in south-central Indiana. Proc. Indiana 

Acad. Sci. 72:182-189. 

11. . 1966. Groundwater movement and cavern development in the Chester 



Series of Indiana. Proc. Indiana Acad. Sci. 75:210-215. 

12. . 1968. The geology and geomorphology of Wyandotte Cave, Crawford 

County, Indiana. Proc. Indiana Acad. Sci. 77 :236-244. 



13. . 1969. Cavern development in a karst groundwater zone (Abstract) 

p. 38. In Geol. Soc. Amer. Absti-acts with programs for 1969. Part 6. 

14. Swinnerton, A. C. 1932. Origin of limestone caverns. Bull. Geol. Soc. Amer. 
43 :663-694. 

15. Thrailkill, J. W. 1968. Chemical and hydrologic factors in the excavation of 
limestone caverns. Bull. Geol. Soc. Amer. 79:19-46. 

16. Woodward, H. P. 1961. A stream piracy theory of cave formation. Nat. Speleol. 
Soc. Bull. 23:39-58. 



Cooling Degree Days in Indiana 

Lawrence A. Schaal, 1 Purdue University 

Introduction 

Abstract 

Cooling degree days are used by the air-conditioning industry to estimate power 
requirements for cooling buildings as temperatures vary from day to day, seasonally and 
geographically. Normal cooling degree days are calculated for various climatic stations 
in Indiana and the appropriateness of the base temperature used will be related to 
daily data recorded from a typical Indiana home. 

Cooling degree days is a meteorological statistic derived from 
accumulating the excess of the daily mean outdoor temperature above a 
chosen base temperature. They closely parallel the energy necessary to 
cool for human comfort the interior of buildings exposed to a hot exterior 
environment. The chosen base temperature has ranged from 60 to 75 °F. 
This study will hopefully add to the apparent meager work done on this 
question for ordinary homes, or small buildings. Normal cooling degree 
days are calculated for some weather stations in Indiana from monthly 
mean temperatures and a chart for the state is constructed from the 
data. 

One cooling degree day is a day when the mean temperature exceeds 
the base temperature one degree. Lower mean temperatures are given a 
value of zero. The resulting sum for a day, a week, or season enables com- 
parisons from one area to another, within various time periods, and in a 
locale to assess cooling energy requirements of structures. Heating 
degree days which are calculated from mean temperatures below 65 °F 
have a wide use in the heating industry, a fine example of profitable use 
of climatology. 

Interest in this study started as the author considered the appropri- 
ateness of 65 °F as the standard base temperature for cooling degree 
days. The Glossary of Meteorology (3) gives the common base of 75 °F. 
In selected Climatic Maj)s of the United States (2) the base temperature 
used is 65 °F. The Climatological Handbook, Columbia Basin States (4) 
suggests 60 °F. In Indiana, a daily mean temperature of 65 °F results 
from minimum temperature for the day of about 54 °F and a maxium 
temperature of about 76 °F. An air temperature of 76 °F is still in the 
comfortable range for non-working humans under common humidity, 
wind and solar conditions. But as described in this paper, the solar 
radiation load on a small building seems to require a lower base 
temperature at least when the maximum interior temperature is set 
at 78 °F. 

The correlation of cooling requirements for several types of com- 
mercial buildings was reported by Thorn (7). Correlations in a 15-day 



1 Agronomy Department and Environmental Science Services Administration, United 
States Department of Commerce. 

292 



Geology and Geography 



293 



study in Washington, D. C. ranged from 0.405 (office building) to 0.918 
(hotel). The base temperature was 65 °F. 

Methods 

In this low budget study the author compared the minutes the air 
conditioner ran in his own home to the measurements of air temperature 
and solar radiation at the Purdue University Agronomy Farm 6 miles 
west-northwest of the house. Readings, taken for 42 days at about 8 AM, 
consisted of the total minutes the home thermostat set at 78 °F caused 
the coolant compressor to run and cool the house (Fig. 1). 




58 



One-tenth of day's solar 
radiation given at points, 



Line equation: 



31.75 T - 1888.9 



200 



300 400 500 
COOLING MINUTES 



600 



700 



800 



Figure 1. Daily observations of outdoor mean temperature are related to minutes 
of cooling to maintain a temperature of 78° F or lower in a typical dwelling. The 
day's solar radiation in tens of calories per square centimeter is given at the point. 
Cooling minutes accumulated when the daily mean temperature exceeded 59° F generally. 



The circulating fan of the furnace ran continuously and the ther- 
mostat was set to turn on the coolant compressor when 78 °F was 
exceeded during the 42 days. A simple electric clock was connected to 
the circuit on the furnace which actuated the coolant compressor and the 
coolant fan outside the house. The room thermostat closed the circuit for 
running the cooling equipment. 

Cooling minutes used in this paper could be converted directly to 
electrical energy by using the electrical specifications of the electric 
motors. 



294 Indiana Academy of Science 

Of some interest was the frequency and time periods when the 
cooling- system ran. These were monitored by placing a hygrothermo- 
graph beside the outlet of the furnace. Thus the rise and fall of outlet 
air temperature and humidity were charted. These data made it quite 
obvious how the radiation load on the house had to be removed by the 
air conditioner. Cooling began on many days at about noon and con- 
tinued until evening or midnight. 

Results and Discussion 

Linear regression was calculated for the 42 daily observations using 
C = aT + k where: C was cooling minutes; T equalled the mean tem- 
perature for the 24-hour period (the average of the maximum and 
minimum temperature); a was the regression coefficient of T; and k 
was the constant. The equation of the regression line was: 
C = 31.75 T— 1888.9 

1888 9 

when C = then T = — = 59.5 

31.75 

The intercept on T when cooling minutes were zero was 59.5 °F. This 
data suggests a base temperature of about 60 °F. The simple correlation 
coefficient was 0.880. 

During the period of observation it was obvious that solar radiation 
had an effect on the minutes of cooling. For this reason solar radiation 
observed at the Agronomy Farm became the second factor in a multiple 
regression problem. The effect of showers was not studied. There were 
very few showers during the 42 days but during one evening a shower 
cooled the house and environs reducing the cooling minutes compared to 
other days at the same temperature. Showers favor shifting the base 
temperature higher since cooling minutes are less. A sunny day favors 
a lower base temperature because the interior temperature of the house 
increased by solar radiation greatly exceeds exterior air temperature. The 
multiple correlation equation using both temperature and solar radiation 
was C = aT + bS + k where: C = cooling minutes; T = mean tempera- 
ture of day; S = solar radiation (daily total gram calories on a square 
centimeter of horizontal surface). The coefficients calculated were: 
a = 30.15; b = 0.288; resulting in C = 30.15 T + 0.288 S — 1939.24. If 
the first equation for C (obtained when the solar radiation term was not 
included) is placed equal to the above equation including the solar radia- 
tion term, we obtain: 31.75 T — 1888.87 = 30.15 T + 0.288 S — 1939.24; 
1.6 T = 0.288 S — 50.37; T = 0.18 S — 31.48. This shows a relation of 
1°F temperature change equal to 0.18 Langley where the two linear 
equations intersect. In the above equation the constant 1939.24 is a little 
greater than the 1888.9 of the first equation which did not have the 
solar radiation term. The intercept of T with zero minutes of cooling 
turns out to be 61.1 °F. 

The coefficient of T and S may also be normalized to show a relative 
relationship for one standard deviation of C by calculating Beta Coeffi- 
cients (1). This is done by dividing the coefficients a and b by the 



Geology and Geography 295 

standard deviation of C which was calculated to be 172.48. They were 
next multiplied by their respective standard deviations as follows: Cal- 
culated standard deviations: for C 172.48; for T 4.781; for S. 155.38. 

30.15 X 4.781 
Beta for T = — = 0.8357; 

172.48 

0.2884 X 155.38 
Beta for S = = 0.2598. 

172.48 

Therefore one standard deviation of C relates to 0.84 standard deviation 
of T and 0.26 standard deviation of S. 

The significance of solar radiation to cooling minutes was calculated 
as follows (N = 42): 

Source of Variation Degrees of Freedom Sum of Squares 
C on T 40 275,116 

C on T and S 39 195,118 

Reduction due to adding S 1 79,928 

79 928 79 928 
For the well known F test, = — ■ = 15.97 

195,188/39 5005 

From F-table values (5), an F of 8.87 or greater indicates that the 
chance is less than 1 in 200 that this set of observations accidentally 
showed such a contribution from solar radiation increasing cooling 
minutes after the effect of temperature. 

The correlation coefficients based on additional tests and computa- 
tions are listed in Table 1. 



Table 1. Correlation coefficients. 

Daily mean temperature and cooling minutes 0.880 

Daily highest temperature and cooling minutes 0.881 

Daily mean temperature and solar radiation with cooling minutes 0.917 

Daily highest temperature and solar radiation with cooling minutes 0.886 



Linear correlation of the maximum temperature with cooling min- 
utes was about the same as with the mean temperature and did not 
improve much with the addition of the solar radiation term. This seems 
to indicate that the maximum temperature reflected the effect of solar 
radiation with very little contribution by the minimum temperature or 
night temperature. 

The importance of interaction between temperature and solar radia- 
tion (product of T and S) to cooling was computed. The addition of this 
third term to the equation became significant at the 5% level but not at 
the 1% level using the F test. 



296 



Indiana Academy of Science 



The one-story house of about 1200 square feet plus a full basement 
was insulated by 4-inch-thick bats of fiber glass insulation in ceilings 
and walls and is believed fairly typical of the contemporary 3-bedroom 
home. The roof was grey with a slope of 3 to 12. The window area was 
more than the average house, and storm windows were in place. Con- 
siderable attic ventilation came from openings under the eaves. It 
seems that the solar radiation load was proportionately larger than for 
a large apartment or office building having large interior areas away 
from exterior walls and roof. Afternoon relative humidity varied little 
during the experiment. Daily lows ranged from 39 to 86 at the 
Agronomy Farm and averaged 53% relative humidity. 



It r\ j-\ " "' 


900 A 




i— L - 1 




1000 
















• 900 I 

1 
1 

*100C 


! 




V 1 






1100 
























— X_ 




— — 




r 








1200 


















1 ^ 




13( 


)0> 












^w^ 


^y 


S T 






SJ 




44-noo 

1— J*»1200 

COOLING 
DEGREE 
DAYS 
NORMAL 

ANNUAL . 


14 




i"" 






00 


kh 


1/1300 




D<>140C 


) 


BASE 65° 





Figure 2. Normal annual cooling degree days calculated from daily mean temperatures 
above C5°F. 



Geology and Geography 297 

I conclude that in a sunny climate and for small houses the base 
temperature for cooling is closer to 60 °F than 65 °F, but perhaps for 
large multistoried buildings the base temperature may be more appro- 
priately 65 °F. Since 65 °F is becoming the most frequently used base 
temperature, a normal cooling degree day map for Indiana has been 
drawn (Fig. 2). Monthly mean temperatures for the period of 1931-1960 
were used since the World Meteorological Organization and Environ- 
mental Science Services Administration, Environmental Data Service 
use this average as the normal. 

The monthly mean temperature of the mid-summer months readily 
convert to cooling degree days with practically no error by subtracting 
65 from the mean and multiplying by the number of days in the month. 
However, for the peripheral months of the season when the mean is 
near 65 we used the standard deviation of the monthly mean tempera- 
ture to estimate the additional degree days needed. Thorn's unpublished 
nomogram (6) was used to estimate the increment. 

For an example in the use of Figure 2, consider a location in Indiana 
where the normal of cooling degree days is 1200 for a season. A sum- 
mer has just passed when the cost of cooling a new building was $300 
and the cooling degree days totalled 1500 as calculated from daily tem- 
peratures nearby. What is the normal expected expense ? Since 1500 
cooling degree days is 300 more than the normal 1200 and 300 is 20% 
of 1500, then 20% of $300, or $60, is the above normal cost of cooling the 
building. A normal or average summer would result in cooling costs $60 
less than $300, or $240. Supposing the next summer averaged near normal 
in cooling degree days and costs were appreciably different from $240. It 
would then be evident that changes in the efficiency of cooling had 
occurred. 

The Indiana map relates to the standard exposure of official ther- 
mometers generally found in rural or village areas. These localities are 
generally cooler than the brick and concrete areas of the cities. There- 
fore city islands of heat are not shown unless included unknowingly in 
some substation data used. 

Conclusions 

In conclusion, this experiment favored a base temperature nearer 60 
than 65 °F for the computation of cooling degree days as they relate to 
home cooling. The test period, however, did cover a sunny period and a 
humidity much higher than found in the western plains, both of which 
should be inversely related to the base temperature. 

Acknowledgments 

The author is indebted to James E. Martin and Oscar Luetkemeier 
for Agronomy Farm observations, to Walter L. Stirm, Advisory Agricul- 
tural Meteorologist for instrumentation, and Robert May for computer 
work. 



298 Indiana Academy of Science 

Literature Cited 

1. Arkin, Herbert, and Raymond R. Colton. 1946. An Outline of Statistical Methods, 
4th Edition. Barnes and Noble Inc., N. Y. 224 p. 

2. Data Information. 1968. Selected Climatic Maps of the United States. Environ- 
mental Science Services Administration, U. S. Department of Commerce. 32 p. 

3. Huschke, Ralph E. 1959. Glossary of Meterology. American Meteorological Society. 
638 p. 

4. Meteorology Committee, Pacific Northwest River Basins Commission. 1969. Climato- 
logical Handbook, Columbia Basin States, Temperature, Vol. 1, Part B. 540 p. 

5. Snedecor, George W. 1965. Statistical Methods, Fifth Edition. Iowa State University 
Press, Ames, la. 534 p. 

6. Thom, H. C. S. 1956. Monthly Degree Days derived from Monthly Mean Tempera- 
ture, nomogram. Environmental Data Service, Environmental Science Services Ad- 
ministration, U. S. Department of Commerce. 

7. Thom, H. C. S. 1966. Cooling Degree Days and Energy Consumption. Air Condition- 
ing, Heating and Ventilation. 63 :53-54. 



The Climatology of Indiana Tornadoes 

Ernest M. Agee, Purdue University 

Abstract 

Various tornado statistics for Indiana have been determined based upon climato- 
logical records from 1916 through 1968. Yearly, monthly, and diurnal variations of 
tornado frequencies, injuries, and deaths are examined on a state-wide and county 
basis. An attempt has been made to remove the problem of the population bias in 
tornado reporting. Also, the effect of terrain on the areal distribution of tornadoes is 
noted. Further study into the matter of orographic effects is suggested. 

Introduction 

A national summary of tornado statistics by Flora (1) has provided 
the scientific community with considerable information on tornado fre- 
quencies and occurrence patterns. It is recognized, however, that the 
scope of study in a national summary does not provide the detailed 
information that is often desired on a state-wide basis. Precedence for 
the value of state summaries of tornado statistics and related climato- 
logical inferences is evident in the work by Darkow (unpublished data) 
on Missouri tornadoes. In fact, the study by Darkow provided consider- 
able incentive to do a similar study for Indiana, especially since it is 
well above the national average as a tornado-producing state. Indiana 
has been involved in some of the most dramatic tornado outbreaks of 
this century. Most notable are the tri-state tornadoes of 1925 and the 
Palm Sunday tornadoes of 1965. 

The primary objectives of this investigation were to determine 
yearly, monthly, and diurnal variations of tornado frequencies, injuries, 
and deaths for Indiana. In addition to obtaining an areal distribution of 
tornadoes, a limited effort was made to examine the effects of population 
and topography. Tornado data for the United States (2) do show a 
tendency for decreasing tornado frequencies toward the southeast corner 
of Indiana, which could be attributed to population and /or terrain 
effects. It was one of the intentions of this study to substantiate or 
dismiss this observation by examining the data in greater detail on a 
state-wide basis. Finally, it was hoped that this work would also provide 
a suitable format and method for examining the climatology of tor- 
nadoes for other states. A generalized computer program has been 
compiled and is available to other investigators. 

Methods 

The period of this study dates from 1916, the beginning of official 
tornado records, through 1968. Data assembled from United States 
Weather Bureau records (3, 4, 5, 6, 7) were checked against the records 
maintained by Indiana's State Climatologist, Mr. Lawrence A. Schaal. 

Data extraction required that certain decisions be made in making 
the tornado tallies. Funnel clouds aloft were not counted but water- 
spouts over Lake Michigan were. Also, tornadic winds were counted 
when accompanied by path damage with tornado characteristics. In 

299 



300 Indiana Academy of Science 

nearly all cases, however, the storm was logged in the records as a 
tornado. 

The format of the data reduction was prescribed by the computer 
program developed simultaneously to handle the data analysis. Infor- 
mation recorded included the date, time of day, and place of origin of 
tornado by county (in some cases out of state). Also, counties affected, 
width and length of damage path, direction of movement, injuries and 
deaths, and estimated property damage were recorded. All available 
information was placed on one computer card per tornado. Problems in 
handling the data included such matters as broken storm paths, chang- 
ing directions, large time intervals, varying damage paths, and different 
translational speeds. 

Other data collected for the study include rural population figures 
per Indiana county (9) for 1920, 1940, and 1960. These three sets of 
values were averaged to yield a mean rural population per county and 
for the state during the period of the study. Also obtained were the 
number of square miles in each county and the state. 

Results and Discussion 

During the period from 1916 through 1968 a total of 551 different 
tornadoes were observed and recorded in climatological data. Of this 
total, 20 came from Illinois, 2 from Kentucky, and 529 originated in 
Indiana. Some of those tornadoes only touched down briefly in a single 
county while others had continuous paths on the ground through several 
counties and into the adjacent states of Michigan and Ohio. 

Figure 1 shows the number of tornadoes affecting each Indiana 
county from 1916 through 1968. Twenty-four tornadoes were observed 
in Porter County, more than any other. Close behind are Elkhart with 
22, Lake with 20, and Marion and St. Joseph with 19 each. These coun- 
ties are highly populated and certainly reflect the effect of population 
density on tornado reporting. No tornadoes have been officially recorded 
for Ohio and Crawford counties. Also shown are the number of tor- 
nadoes originating in each county. A maximum of 20 tornadoes origi- 
nated in Lake County. On the other hand none have originated in 
Crawford, Dearborn, Ohio and Union counties. Of course counties are 
political boundaries and do not represent equal geometrical areas. But, 
a county-by-county analysis has been made for the sake of convenience. 
Figure 1 also gives the number of tornado days per county. Porter 
County has the highest value with 21. 

In Figure 2 the effects of population and unequal county areas 
have been considered. To reduce the number of tornadoes affecting each 
county to a more meaningful statistic a tornado index has been formed. 
This dimensionless quantity is defined as, 



Tornado Index = 
f Tornadoes /Unit Area"! f Rural Population 



I Rural Population J I Tornadoes/Unit Area 

county state 



(1) 



Geology and Geography 



30] 



ELKHART 

16 



4© 2 



STEUQEN 

4© 5 




Figure 1. Tornadoes affecting Indiana counties (circled), tornadoes originating in 
county (underlined) , and the number of tornado days per county from 1916 through 1968. 



302 



Indiana Academy of Science 




Figure 2. Tornado indices per Indiana county for 1916 through 1968. 



Geology and Geography 303 

The tornado index for each county is therefore based on a common 
denominator of 1.00, the state average. Counties with indices greater 
than 1.00 are above the state average and those less than 1.00 are below 
the state average. The use of rural population was observed to more 
aptly handle the problem of population bias than total population. Fur- 
ther, it seems logical that a given area should only be inhabited to a 
certain extent, beyond which additional population would not improve 
the chances of tornado detection. Rural population figures were also 
used by Darkow in his Missouri study, supporting the approach in this 
investigation. The largest tornado index, 2.60, was computed for Porter 
County. Other high values include Jasper with 2.09 and Pike with 2.06. 
The northwest quadrant and the southwest corner of the state are well 
above the state average. As was expected the southeast portion of the 
state is considerably below the state average. It was not the objective 
of this study to examine in detail the effect of topography on tornado 
distribution but these data are indicative of terrain effects and do 
suggest further study into the problem. It should be mentioned, how- 
ever, that the method used to handle the population bias in tornado 
reporting may not be the most accurate and the tornado indices could 
be slightly misrepresentative. 

Figure 3 shows the yearly distribution of tornadoes and tornadoes 
causing injury or death. Most notable is the increase in tornado occur- 
ences since 1954. This can be partially attributed to the development of 
the severe local storm (SELS) forecasting and warning program by the 
U. S. Weather Bureau. This program also improved and updated the 
efforts in the tornado recording network which should account for much 
of this increase. Figure 3 is also indicative of more tornadoes causing 
injury and death in recent years. The most tornadoes for any single 
year was the 52 recorded in 1965. Nineteen of these caused injury or 
death. However, the Palm Sunday tornado outbreak in 1965 did make 
this an exceptionally high tornado year for Indiana. Also shown is the 
yearly distribution of tornado days and tornado death days for Indiana. 
Again, the initiation of the program by the U. S. Weather Bureau is 
reflected in the data. 

Figure 4 shows the monthly distribution of tornadoes. More tor- 
nadoes occurred in April than any other month and 78% of all tornadoes 
occurred during the period March through July. The maximum number of 
tornado days occurred in June but tornado days in early spring produced 
more tornadoes. For instance, 45 tornado days produced 63 tornadoes 
in July, but 30 tornado days produced 71 tornadoes in March. Another 
interesting feature is that a larger portion of the tornadoes occurring 
in late winter and early spring caused injury and /or death. Notice that 
March had 71 tornadoes, 30 of which caused injury and /or death. But, 
April had 124 tornadoes and only 29 caused injury and/or death. This 
seems to indicate that the tornado season must be in progress for awhile 
before people begin to take it seriously. The data also suggest a small 
second maximum of tornado occurrences in early fall. This is associated 
with increased frontal activity which is not as violent as that associated 
with the storms that develop in the spring. 



304 



Indiana Academy of Science 




Geology and Geography 



305 



130 




120 




110 




100 




90 




80 




70 




60 


• 


50 




40 




30 




20 




10 





•78% 



□ TORNADOES 
BS3 TORNADO DAYS 
■i TORNADOES CAUSING 
INJURY AND/OR DEATH 




JFMAMJ J ASOND 



FiGURi: 4. Indiana tornado occurrences by month from 1916 through 1968. 



306 



Indiana Academy of Science 



Table 1. Indiana tornado deaths and tornado death days by month 
from 1916 through 1968. 

MJJASOND 
3 2 10 10 





J 


F 


M 


A 


Tornado 










Death Days 


1 





10 


4 


Tornado 










Deaths 


1 





207 


157 











Table 1 gives the number of Indiana tornado deaths and tornado 
death days by month for the period. More deaths have occurred in 
March which interestingly enough is not the month of maximum tornado 
activity. 




00 02 04 06 08 10 12 14 16 18 20 22 24 

Figure 5. Diurnal variation of Indiana tornado activity from 1916 through 1968. 



The diurnal variation of tornado activity is given in Figure 5. 
Tornadoes can occur any hour of the day just as they could any day of 
the year, but late afternoon is the preferred time. More tornadoes 
occurred between 5:00 and 6:00 pm than any other hour of the day. A 
second maximum appears between midnight and 1:00 am which is prob- 
ably attributed to the midwest's nocturnal thunderstorm activity. Also, 
tornadoes causing injuries appear to be less in proportion to the num- 
ber of tornadoes for nighttime compared to daytime. If such is true this 
could be related to the fact that more people are at rest or asleep in 
their homes and are therefore not quite as vulnerable to injury. 



Geology and Geography 307 

Table 2. Diurnal variations of Indiana tornado deaths from 
1916 through 1968. 



00-01 


01-02 


02-03 


03-04 


04-05 


05-06 


06-07 


07-08 


08-09 


09-10 


10-11 


11-12 








3 


2 








1 


















12-13 13-14 14-15 15-16 16-17 17-18 18-19 19-20 20-21 21-22 22-23 23-24 
2 24 66 118 174 40 73 30 5 



Table 2 shows the diurnal variation of tornado deaths. The maxi- 
mum is associated with peak tornado activity. Again it is evident that 
nighttime deaths are much less than for daytime in proportion to the 
number of tornadoes occurring. Since death producing tornadoes did 
occur at the beginning and end of adjacent hourly periods, the tabular 
data indicate more deaths than actually happened (Table 1). 

The direction of movement in percentages for Indiana tornadoes 
has also been determined. Over 81% of the tornadoes traveled in an 
east through northeast direction with 47% moving toward the north- 
east. Tornadoes have been observed to move in all directions but from 
southwest to northeast is the general tendency. 

Summary 

Yearly, monthly, and diurnal variations of tornado frequencies, 
injuries, and deaths for Indiana have been presented based upon data 
assembled for the period 1916 through 1968. The data are summarized 
in Figures 1-5 and Tables 1 and 2. An attempt has been made to remove 
the problem of population bias in tornado reporting. Also, the effect of 
topography on the areal distribution of Indiana tornadoes has been 
touched upon and further study is intended along these lines. A gen- 
eralized computer program has been developed, and is available, to do 
similar climatological analyses for other states. 

Acknowledgments 

I am grateful to Dr. P. J. Smith for his review of the manuscript 
and to Mrs. Carol Gilliom for her assistance in the data analysis and 
computer programming. This work was partially supported by NASA 
Engineering System Design Traineeship Program Project No. NGR 15- 
005-069. 

Literature Cited 

1. Flora, Snowden D. 1954. Tornadoes of the United States. University of Oklahoma 
Press, Norman, Okla. 194 p. 

2. U. S. W. B. Technical Note No. 20, 1960: Tornado Occurrences in the United States. 
United States Weather Bureau, Washington, D. C. 

3. Report of the Chief of the U. S. W. B., 1916-1931. U. S. Weather Bureau, Wash- 
ington, D. C. 

4. U. S. Meteorological Yearbook, 1931-1949. U. S. Weather Bureau, Washington, D. C. 



308 Indiana Academy of Science 

5. Monthly Weather Review, 1921-1949. United States Weather Bureau, Washington, 
D. C. 

6. National Climatological Data, 1950-1961. U. S. Department of Commerce, Weather 
Bureau, Washington, D. C. 

7. Indiana Climatological Data, 1950-1961. U. S. Department of Commerce, Weather 
Bureau, Washington, D. C. 

8. Storm Data, 1959-1968. U. S. Department of Commerce, Weather Bureau, Washington, 
D. C. 

9. U. S. Bureau of the Census, 1920, 1940, 1960. Census of Population, Washington, U. S. 
Government Printing Office. 



The Changing Location Patterns of the Neighborhood Grocers in 
Terre Haute, Indiana : A Geographic Analysis 

R. Michael Dinkel and A. J. Cantin, Indiana State University 



Abstract 

In today's corporate society, the small businessman is becoming less and less 
common. A prime example of this presently can be found in the food marketing 
industry. Since the year 1900, there have been many significant changes in the grocery 
business throughout the United States. This study specifically deals with the causes 
and effects of changing location patterns of the small grocer in Terre Haute, Indiana. 

The location of the small grocer with respect to major thoroughfares, does appear 
to be instrumental in the survival of the small grocer against the onslaught of various 
supermarket chains. Therefore the geographic location seems to be a major influence on 
the survival of this particular type of small business establishment. 



Introduction 

The changes in the location and distribution of retail grocers within 
the corporate limits of Terre Haute, Indiana, is an example of the 
changing economic environment of man. This study was undertaken 
during the spring of 1969 when both authors, in personal discussion, 
observed the rather rapid rate at which certain neighborhood grocers 
were being eliminated, due perhaps to strong competition of the 
numerous supermarket chains. 

We questioned whether the elimination of the neighborhood grocers 
was due to the location of supermarket chain stores in shopping centers. 
Since in Terre Haute there are only a relative few such centers, it could 
not, by itself, account for the change. We, therefore, decided to test his- 
torically, the hypothesis of location with respect to major thorough- 
fares. 

It might be assumed that the retail grocer would tend to polarize 
around the major thoroughfares within the city. This was found to be 
true in the early 1900's and in the past two decades. However, in the 
intervening years, a negative trend was apparent with regard to locat- 
ing on major thoroughfares. 

Data for this study were gathered from the Terre Haute City Di- 
rectory for the years 1900 to 1967. Recognizing that time would bring 
considerable change in the location pattern, we decided that a sample 
of one year in a decade would demonstrate such change. The years 
selected were drawn randomly, and if data for that year were lacking, 
the next year was arbitrarily selected. These data were then plotted 
on a city map (using the city limits for the year selected). Stores out- 
side of the city limits were not included in the study and were not 
plotted. For purposes of finding trends in recent years, data for both 
1962 and 1967 were included (Table 1). 

309 



310 



Indiana Academy of Science 



Otiwwt* 




Fig 



URE 1. The location of retail grocery stores in Tcrre Haute, Indiana, in 1! 



Geology and Geography 311 

Table 1. Retail grocers on major traffic arteries 





1900 


1908 


1918 


1927 


1937 


1944 


1954 


1962 


1967 


Wabash Ave. 


27 


22 


28 


31 


12 


12 


11 


5 


3 


Lafayette Ave. 


16 


14 


16 


34 


21 


17 


13 


8 


5 


Poplar St. 


16 


18 


10 


12 


12 


9 


7 


6 


6 


Thirteenth St. 


20 


20 


25 


27 


22 


17 


18 


10 


7 


Third and 




















Fourth Sts. 


29 


27 


39 


56 


39 


26 


27 


13 


13 


Twenty-Fifth St 














9 


7 


6 


Harding Ave. 


21 


13 


14 


15 


10 


12 


— 


— 


— 



Analysis of Patterns, 1900-1967 

1900 Map 

In 1900, location with respect to major thoroughfares was ex- 
tremely important (Fig. 1). Four major concentrations were observable: 
along Lafayette, Wabash, Poplar, and Third and Fourth Streets. Concen- 
trations were present on Thirteenth Street and on Harding Avenue (Sec- 
ond Street), which formerly was the major north-south route in the 
city. The heavy concentration on Fourth Street was an enigma until 
it was learned of a Farmers Market in the Court House area. Along 
Fourth Street, a huge "hay rack" had been installed by farmers to 
mass feed their horses when they brought in their produce for sale 
and grocers located with respect to this potential market. 

Located along all streets mentioned, with a heavy emphasis in 
the Central Business District (CBD), were 129 of 191 grocers in Terre 
Haute, nearly 68 % of the total. Concentration along major thorough- 
fares in 1900 was extremely important. Note, by contrast, the sparse 
pattern in the NE and SE neighborhoods of Terre Haute. One can sum- 
marize the city for 1900 as having very few neighborhood grocers. 

1908 Sample 

Vast changes occurred between 1900 and 1908. The number of 
neighborhood grocers increased, particularly in the NE and SE sec- 
tions of the city. Now only 114 of 224 (or about 50% of the total) were 
located on the 6 main arteries of the city — a drop of 18% from 1900. 

As the city grew toward the NE and SE and construction of many 
homes of the working class were completed, neighborhood grocers ap- 
peared within these areas. 

However, the major distribution of stores was still dominated by 
location with respect to points of major traffic flow. A major redistribu- 
tion was taking place. The heavy concentration near the "hay rack" 
had diminished, and the Third and Fourth Street concentration had 
spread toward the city limits. These two streets had but one additional 
store in 1908 as compared to 1900, but those stores pre-dating 1908 



312 Indiana Academy of Science 

were no longer concentrated near the CBD to the degree they were in 
1900. 

Between 1900 and 1908 a decline, to remain unchecked for nearly 
half a century, began for stores located along major thoroughfares, 
with consequent growth in importance of neighborhood stores. 

1918 Sample 

By 1918 the trend toward greater numbers of neighborhood grocers 
continued with a drop of nearly 5% of the thoroughfare-oriented stores 
from 1908. Note the relatively dense pattern of neighborhood stores in 
NE and SE Terre Haute. 

The traffic-flow locations are realigned, with growing importance 
for Thirteenth, Wabash, and Third and Fourth Streets, plus a 50% loss 
of the grocers located on Poplar. Reappearing is a very heavy con- 
centration on Third and Fourth Streets between Ohio and Chestnut 
Streets. These stores were largely small stores owned by newly arrived 
Syrian immigrants in a low-income area of the city. 

Between College and Hulman and Twelfth and Fourteenth Streets, 
a Negro district referred to as "Baghdad," had nearly a 250% increase 
from 7 to 18 stores. Again, as in the Third and Fourth Street complex, 
this illustrates a growing tendency for large numbers of small stores 
to locate in low-income areas. 

1927 Map 

This was the peak year for grocers in Terre Haute, as there were 
489 stores located in the city, up from 295 in 1918 (Fig. 2). A large per- 
centage of this was due to the 36-store Oakley Chain that was opened 
during this period, and years later, sold out to Kroger Supermarkets, Inc. 
These stores were spaced evenly throughout the entire city and were 
supplied, in part, through Oakley's farm. 

This locational trend continued. The major traffic arteries accounted 
for only 40% of the stores, despite a noticeable growth of stores in the 
Third and Fourth Street area and Lafayette Avenue which increased 
from 16 stores to 34. 

Major traffic arteries had a higher number of stores than ever 
before, but a smaller percentage of the total. Store coverage was most 
uniform in the 68 year period. More stores were located in neighbor- 
hoods, except in the "Baghdad" area where the number of stores fell 
from 18 to 7, the number originally found in 1908. 

1937 Sample 

The effect of the depression is demonstrated by the rapid decline in 
grocer numbers, from 439 stores to 338, or a 23^ decrease. In 1937 
only 116 of 338 stores (or 34%) were located on major traffic routes, 
down 6% from the previous decade. 



Geology and Geography 



313 




Figure 2. The location of retail grocery stores in Terre Haute, Indiana, in 1927. 



The remaining stores had even distribution, with only a few cluster- 
ings. A new cluster in the Twelve Points area, which began in 1927, 
continued through 1937, due perhaps to the rebuilding of Garfield High 
School and a resultant increase of residences in north and northeast 
Terre Haute. 



314 



Indiana Academy of Science 




FIGURE 3. The location of retail grocery stores in Terre Haute, Indiana, in 195U- 



Of the 101 stores that failed from 1927 to 1937, 59 of these were 
located on major thoroughfares, indicating the declining importance 
of this location factor. 



Geology and Geography 



315 




Figure 4. The location of retail grocery stores in Terre Haute, Indiana, in 1967. 



1944 Sample 

The number of grocers declined from 338 to 276, a drop of 62 
stores. However, an important trend change occurred at this time. Al- 
though only 93 stores of the 276 were located on traffic-flow routes the 



316 Indiana Academy of Science 

percentage of stores so located remained at 347c, the same figure as 
for 1937. This marks the first time the decrease was stayed. Stores lo- 
cated on traffic-flow had fewer failures than neighborhood stores, indi- 
cating a trend away from the neighborhood stores for the first time. 
No particular section of the city had a disproportional share of grocer 
failure. 

1954 Map 

By this date new thoroughfare alignments had been made in Terre 
Haute (Fig. 3). Harding Avenue (Second Street) was no longer a major 
north-south route, but was replaced by Seventh, Eighth, and Ninth 
Streets. Also, Twenty-fifth Street had grown to major status. 

The percentage of stores on major traffic arteries had grown to 
41%, up considerably from 1944. This marks the first time in 50 years 
that the pattern had definitely reversed, the gain for major traffic 
locations came from a net loss for neighborhood stores. 

1962 Sample 

The rate of decline was now accelerating as there were but 161 
grocers left in Terre Haute, a drop of nearly 100 stores in an 8-year 
period. Of those remaining, 68 or 43% were on major traffic locations, 
up 2% since 1954. This indicates the weakening position of the neigh- 
borhood grocer and that traffic-flow and shopping centers (as a location 
factor) were making inroads. 

1967 Map 

All trends continued, but now at an accelerated rate (Fig. 4). Of the 
107 remaining stores, 53 (or about 50%) are located with respect to 
major traffic, up 7% in 5 years. In addition, at least six major super- 
markets were established in shopping centers, leaving relatively few 
neighborhood grocers in Terre Haute. 

The net number of stores in the city is declining rapidly. Obviously 
the supermarket era which began after World War II is now establish- 
ing itself, and larger stores have spread farther apart to areas of easy 
automobile access. The decline of the neighborhood grocer is apparent 
both in relative importance and in total numbers. 

Conclusion 

It is now apparent that there are many reasons for small grocers 
locating where they do in Terre Haute, but location within a neighbor- 
hood is no longer significant. Location with respect to shopping centers 
is a growing factor; nonetheless, the most important factor at present 
is major avenues of transportation. 

The future of the neighborhood grocer in the city appears to be 
rather precarious. Although the total volume of each retail store was 
unattainable, that of the small neighborhood grocer doubtless is very 



Geology and Geography 317 

low. The authors estimated, by using a small sample of shoppers, that 
the major chain stores plus allied independent grocers (i.e., IGA) con- 
trol in excess of 90 ( /< of the total retail volume. 

It would appear that the average consumer, due to the higher prices 
charged by the neighborhood grocer, will most often patronize the 
larger chain for purely economic reasons. The convenience of a neigh- 
borhood store appears, therefore, to remain secondary to the savings 
effected by shopping at a larger volume establishment. 



Literature Cited 

1. Dinga, Carl F. 1968. Analysis of retail site locations in Terre Haute, Indiana. Proc. 
Indiana Acad. Sci. 77:321-25. 

2. Guernsey, Lee. 1961. Characteristics of the Terre Haute Central Business District. 
Proc. Indiana Acad. Sci. 71 :203-09. 

3. Terre Haute, Indiana City Directories, 1900-1967. R. L. Polk Publishing Company. 



Population and Settlement Decline in Southwestern Indiana 

Thomas Frank Barton, Indiana University 

Abstract 

The largest contiguous and historically continuous area of population and settlement 
decline in Indiana is found in the southwestern part of the state. This mobile and 
declining population influences many facets of the economy and life style. Reasons for, 
and potentialities of, a Southwestern Indiana Regional Planning Commission are pre- 
sented. Population shifts are not a disaster if proper short- and long-ranged plans are 
adopted to readjust to the situation. 

Introduction 

The largest continuous area of population and settlement decline 
in Indiana is found in the southwestern part of the state. Except for 
Vermillion County, there are 17 counties located in the crotch of the 
Wabash and Ohio Rivers in which populations declined between 1950- 
1960, and 4 counties which gained population only because births ex- 
ceeded the combined number of deaths and emigration. The counties 
designated in this paper as southwestern Indiana are listed in Table 1. 
Population mobility is characteristic of this area with some people 
moving from: 1) one county in the area into another in the area, 2) 
the countryside into the settlements, 3) smaller settlements to the 
largest cities, and 4) from the area. This moving and declining popu- 
lation influences many facets of the society, such as: taxation and 
debts on formerly fully-used public and private facilities; property 
values; need and support for commercial establishments and services; 
land-use requirements; and future economic development. Population 
shifts in a state are not a disaster if proper short-ranged and long- 
ranged plans are adopted to re-adjust to the situation. 

County Population Decline 

During 1950-1960, 21 Indiana counties decreased in population and 
17, or over 4/5 of these, are located in southwestern Indiana. The four 
counties outside of this region that declined in population are Warren, 
adjacent to the Indiana-Illinois boundary and adjacent to this region 
on the north, and Union, Ohio and Switzerland counties, adjacent to 
the Indiana-Ohio boundary in southeastern Indiana. 

Not only did counties in this region decline in population between 
1950-1960 but also the number of counties with such declines steadily in- 
creased in the past 30 years, 1930-1960 (Table 1). During the 1930-1940 
decade, 14 counties increased in population and only 7 decreased; the 
highest percentage loss was 6.2 in Vermillion County. However, dur- 
ing the decade of 1940-1950 when employment opportunities and 



Statistics in this paper preceding the centerhead "Some Chain Reactions" are taken 
from the 1930, 1940, 1950 and 1960 Census of Population of the United States or secured 
by using the data from these volumes. 

318 



Geology and Geography 



319 



Table 1. Population percentage changes in Southwestern Indiana. 



County- 



Estimated 
1930-1940 1940-1950 1950-1960 1966-1985 



Clay 

Crawford 

Daviess 

Dubois 

Gibson 

Greene 

Knox 

Lawrence 

Martin 

Orange 

Owen 

Parke 

Perry 

Pike 

Posey 

Spencer 

Sullivan 

Vanderburgh 

Vermillion 

Vigo 

Warrick 



. — ■ 


4.2 


— 


5.7 


+ 


1.2 


— 4.1 


+ 


0.1 


— 


8.7 


— 


9.8 


— 25.0 


+ 


L.3 


+ 


2.3 


— 


0.5 


— 15.3 


+ 


9.9 


+ 


5.3 


+ 


15.5 


+ 20.6 


+ 


5.2 


+ 


* 


— 


2.5 


— 10.3 


— 


0.5 


— 


11.0 


— 


5.6 


— 25.9 


+ 


0.4 


■ — 


1.3 


— 


4.3 


— 21.4 


— 


1.5 


— 


2.0 


— 


6.5 


— 5.2 


r 


1.9 


+ 


3.7 


— 


0.7 


— 9.0 


— 


0.8 


— 


2.5 


— 


* 


— 6.2 


\ 


6.5 


— 


2.7 


— 


3.1 


— 16.6 


+ 


4.8 


— 


9.7 


■ — 


5.6 


— 14.2 


+ 


6.9 


— 


2.3 


— 


0.8 


- 16.6 


f- 


4.2 


— 


12.0 


— 


14.7 


— 33.3 


+ 


7,1 


+ 


3.3 


— 


3.0 


— 23.8 


— 


3.0 


— 


0.2 


— 


0.6 


— 17.6 


— 


4.0 


— 


12.4 


— 


8.2 


— 23.8 


f 


15.4 


+ 


22.7 


+ 


3.3 


+ 29.0 


— 


6.2 


— 


9.5 


— 


10.3 


— 18.7 


+ 


0.9 


+ 


5.5 


+ 


3.1 


+ 3.6 


+ 


6.6 


+ 


10.8 


+ 


9.5 


+ 12.0 



Change too small to compute percentage. 



employment surpassed the previous decade, 13 counties declined in 
population, 7 gained and Orange County had a small net loss. During 
1950-1960 only 5 counties gained in population and 3 of these only 
slightly— Clay 1.2%, Vigo 3.1% and Vanderburgh 3.0%. 

Although 11 counties in this region had both gains and declines 
during the 30-year period from 1930-1960, 10 counties experienced 
consistent declines or gains. Four counties (Dubois, Vanderburgh, Vigo 
and Warrick) registered continuous gains and six counties (Greene, 
Lawrence, Orange, Spencer, Sullivan and Vermillion) had continuous 
losses. The six declining counties are located adjacent to the four 
gaining counties. Two of the four counties with continuous population 
growth, Vanderburgh and Vigo, contain the two largest cities in this 
area, Evansville and Terre Haute, respectively. Three counties with 
continuous gains (Vanderburgh, Warrick and Dubois) are adjoining, 
but the three-county cluster is surrounded by Indiana counties where 
one or two decades of population decline has taken place. 

The severity of population decline was greater during the more 
prosperous decades of 1940-1960 than during depression years of the 
1930's. The population declines during the 1930-1940 period in the 7 



320 Indiana Academy of Science 

counties ranged between 1.5 and 6.2%. In contrast, the population de- 
clines during the decade of 1940-1950 in 13 counties ranged between 
1.3 and 12.0 % with 6 counties having declines greater than 6.2%. Dur- 
ing the 1950-1960 decade the decline in 16 counties ranged between 0.7 
and 14.7%. 

Population Growth in the Region 

The 21 counties in this region have had a small population gain 
during 1940-1960. The gain was 2.8% between 1940-1950 and dropped 
to about 0.5% during the 1950's. 

This region is characterized by emigration. None of the 21 counties 
escaped emigration. Any gain in population was chiefly due to births 
outnumbering the total deaths and emigration. Not one of these coun- 
ties registered an increase of 18.5%, which was both the state and na- 
tional growth rate during the 1950-1960 decade. In contrast, 22 counties 
in Indiana had a population growth greater than 18.5% with increases 
ranging between 19.7% in Tippecanoe and 66.3% in Hendricks counties. 

Growth in Settlements 

Settlements with over 1,000 inhabitants increased in 15 counties 
and decreased in 4 during 1930-1960. The increase ranges between 
3.3% in Spencer County to 39.7% in Parke County, and the rate of 
decrease between 3.0% in Knox County and 23.9% in Vermillion County. 
Crawford County does not have a settlement with 1,000 inhabitants, and 
in Sullivan County, settlements with over 1,000 decreased by 7 persons. 

During this 30-year period the number of settlements with popu- 
lations over 1,000 increased from 41 to 46. Six of these 46 did not have 
a population of 1,000 in 1930. And, only one settlement with 1,000 or 
more inhabitants in 1930 dropped below that level before 1960. 

While it is true that the population in settlements of over 1,000 
has increased in 15 counties during 1930-1960, 18 of these were smaller 
in 1960 than they were in 1950 (Table 2). Moreover, 12 of these 18 were 
cities with a minimum population of 2,000 and the largest, Vincennes, 
had 18,046 in 1960. The 18 settlements with population declines totaled 
78,312 in 1960, as compared with 83,605 in 1950, or a loss of 5,293. 
This loss is relatively small when compared with the gain of 12,907 
persons by Evansville in the same decade. At the same time, 3 of the 
small cities lost over 10% of their inhabitants: Oakland City, 14.8%, 
Jasonville 17.0% and Bicknell 37.0%. During the preceding decade, 
1940-1950, Oakland City's population gained while Bieknell's declined 
by 10.5%, and Jasonville lost 14.1% (2). 

More settlements of 1,000 or more declined in population during 
1940-1950 than during the following decade. There were 41 settlements 
of this size in 1950; 20 of these had a population decline during the 
1940's. 

The cities with populations of over 5,000 registered a smaller de- 
cline during 1950-1960 than cities of less than 5,000. Of the 12 cities 



Geology and Geography 



321 



Table 2. Population trends in county seats and /or largest cities. 



County 



County Seat and/or 
Largest Cities 



1930- 
1940 



1940- 1950- Population greater in 1960 
1950 1960 than any other decade 



Clay 


Brazil 


Crawford 


English 


Daviess 


Washington 


Dubois 


Jasper 


Gibson 


Princeton 


Greene 


Bloomfield* 




Linton 


Knox 


Vincennes 


Lawrence 


Bedford 


Martin 


Loogootee* 




Shoals 


Orange 


Paoli 


Owen 


Spencer 


Parke 


Rockville 


Perry 


Cannelton 




Tell City* 


Pike 


Petersburg 


Posey 


Mount Vernon 


Spencer 


Rockport 


Sullivan 


Sullivan 


Vanderburgh 


Evansville 


Vermillion 


Clinton* 




Newport 


Vigo 


Terre Haute 


Warrick 


Boonville 



+ 
+ + 
+ 
+ 
+ 



+ 

+ 
+ + 



1 



* Largest city in county but not the county seat. 

with over 5,000, 7 had a larger population in 1960 than in 1950, and 5 
had less. The two largest cities, Evansville and Terre Haute, with over 
70,000, gained in population during the 1950's while the next 2 largest 
cities, Vincennes and Bedford, lost. 



Summary of Population Decline 

Counties, towns and cities have lost population in Southwestern 
Indiana. Some of these declines existed during only 1 decade, some 
during 2 and in others, declines were continuous over 30 years, between 
1930-1960. Projections of population growth between 1966 and 1985 pre- 
dict that 17 of these 21 counties will decline by approximately 57,000 
and 4 counties, Dubois, Vanderburgh, Vigo and Warrick, will gain about 
61,000 inhabitants during the same period (4). The total gain would be 
only 4,000, a very small gain (less than 1%). Should the projection be 
correct, since the total population gain was only 0.5% during 1950- 
1960, it is obvious that this region has problems with population 
declines, almost stagnant growth and emigration. In contrast, other coun- 
ties in Indiana will rapidly gain in population due to births and immi- 
gration. For example, according to John M. Huie, Monroe County's 
population increased by 35.5% between April 1, 1960, and July 1, 1966, 
and immigration of more than 14,000 was the highest of any county in 
the state. During the same period Vanderburgh's emigration reached 
11,800. 



322 Indiana Academy of Science 

Some Chain Reactions 

As the population of an area declines over a period of several 
decades a chain of economic, social and political reactions bring about 
complications. Some of these interrelated reactions are briefly described 
in the following paragraphs. 

1) Local employment opportunities either stagnate or decline. 
In Southwestern Indiana employment in mining, agriculture, com- 
merce and services has declined in recent decades. Technological change 
and new techniques have influenced need for employment in all these 
occupations. New machinery and techniques in coal mining have dras- 
tically cut the need for a large number of miners. Machinery, farm 
consolidation and corporate farming have resulted in rapid declines in 
population and farm employment opportunities. With decline of mining 
and farming employment, fewer people are needed to supply goods and 
services resulting in some becoming temporarily or permanently un- 
employed. As work opportunities appeared in adjacent counties or in 
nearby cities people were forced either to commute to work or to move. 

2) Areas with declining populations generally have a larger num- 
ber of their people in the upper age brackets. Emigration generally con- 
sists of the younger workers and those more economically "foot loose." 
And the older people who are: unable to liquidate their property; will- 
ing to accept less affluent living conditions; and /or incapable of leaving 
life-long friends, remain behind. 

3) Employees who remain in the area spend more time and money 
in commuting to work in cities such as Evansville, Terre Haute, and 
Vincennes and to factories in Dubois County or in counties outside the 
area such as Monroe. 

4) As the population declines in townships, counties, incorporated 
villages and small cities, school districts and special districts, it often 
becomes more expensive per person or family to support the existing 
services and to provide new facilities. Moreover, as commercial and/or 
small industrial establishments go out of business or move from the 
area, heavier taxes fall upon residential and agricultural land and 
structures. 

5) In attempting to reduce costs, new services (typical of more 
prosperous counties and cities) are often postponed and the quality of 
the service already established may decline. For example, the size of 
police and fireman forces may be cut, new equipment may not be pur- 
chased and that on hand may not be adequately repaired. 

6) With the resulting population decline, public services retarded 
and /or restricted, and tax rates increased, it becomes very difficult for 
settlements to attract manufacturing and service industries which could 
give the community a more varied employment mix. Industrial manage- 
ment makes careful investigations of the quality and quantity of serv- 
ices available before moving into a city. 



Geology and Geography 323 

7) Residential property values may decline in villages and small 
cities thus putting owners with large debts in a financial squeeze and 
preventing people, if they must sell, from receiving a reasonable price 
for their land and buildings. 

8) The number and quality of commercial and service establish- 
ments in small cities may decline which may: reduce customer selection; 
raise prices; and /or influence customers to shop in other cities. 

9) The number, kinds and quality of professional services may 
also decline and some may disappear. Some villages and smaller cities 
which formerly had several doctors, dentists, lawyers and bankers now 
have only one or none of each profession. 

10) As cities become dormitory settlements where most of the 
taxes are levied on residences, the taxing unit cannot afford to provide 
the services considered necessities in a twentieth century community 
or, if it does, it becomes a burden. 

11) Vacant urban structures and land provide a poor economic 
image to potential developers. 

12) Commuters who live outside the corporate limits may be sub- 
ject to payroll taxes. Mayor Frank F. McDonald, Evansville, estimates 
that 15,000 persons who work in Evansville live outside the political 
city (1). 

As these and other actors interact the economic situation spirals 
downward aggravating forces already in operation, sometimes intro- 
ducing other degrading forces, and stimulating additional emigrations 
to more prosperous areas in Indiana and other states. According to 
Huie (4), Indiana's employment grew by approximately 206,000 jobs 
between 1950-1960 but, according to the United States Department of 
Commerce, if Indiana's employment had grown at the national rate there 
would have been an additional 29,000 jobs. 

Some Suggested Action 

What has been done in the last decade or two to help the county 
leaders and people to meet the problems associated with declining 
populations? Have conferences been held, study groups organized and/ 
or a program of investigations been launched? Is there literature on 
these problems that could be made available to the high schools and 
universities in this area? Does the Legislature or the State have study 
groups for research and to offer suggestions and alternate courses for 
action? ,..;.._.,.,_ 

It seems that not too much has been written either about the 21 
counties under consideration or the problems associated with declining 
populations and settlements. And one of the best publications a propos 
to this area, entitled Regional Development and the Wabash Basin (3), 
either has not been read or understood or, if so, has not stimulated much 
action or reaction in Southwestern Indiana. 



324 Indiana Academy of Science 

It seems that most of the American literature on settlements, urban- 
ism and urban problems and solutions to problems is about cities with 
populations over 100,000. Moreover, the literature on problems created 
by rapid population and city growth far exceeds that on problems and 
alternate solutions for areas of population and settlement decline. There 
is need for material written on smaller settlements. Southwestern 
Indiana is a region of small settlements with only 5 of the 46 having 
populations of 10,000 or more. Twenty of the 46 have populations in the 
1,000 to 2,000 bracket. And in Indiana where a settlement qualifies as 
a city if it has a population of 2,000, there are 26 cities with 14 in the 
2,000-5,000 bracket and 7 in the 5,000-10,000 bracket. 

The government provides information on surveys and studies deal- 
ing with the Wabash and Ohio River watersheds and has made or will 
make suggestions for economic and social development. But most of 
these studies focus on areas of both rapidly-expanding and declining 
counties. 

Is it possible for the 21 counties in this area and Warren (adjacent 
and to the north) to create a regional planning organization which could 
focus directly on the problems, potentialities and beneficial adjustments 
for this area ? Such an organization could provide a forum wherein the 
problems of the area could be aired, freely discussed and courses of 
action agreed upon in a non-partisan atmosphere. Conferences could be 
held for high school students and their parents, university audiences, 
and in settlements throughout the region to discuss economic and so- 
cial problems and alternative solutions. A Southwestern Indiana Re- 
gional Planning Commission could outline and start a program of 
study investigations. It would seem that such a program would be 
better than the present situation. This is serious and action is needed 
now. 

Literature Cited 

1. Anonymous. 1969. Politics in perspective: eminent mayors favor payroll tax. The 
Indianapolis Star, Section 2, September 28, p. 8. 

2. Barton, Thomas Frank. 1952. Cities with a population decline in southwestern 
Indiana, 1940-1950. Proc. Indiana Acad. Sci. 62 :250-255. 

3. Boyce, Ronald R. (Ed.) 1964. Regional development and the Wabash basin. Univer- 
sity of Illinois Press, Urbana. 

4. Huie, John M. 1968. Population changes pressure local governments. Economic and 
Marketing Information for Indiana Farmers. December 31. p. 2-4. 



A Comparison of the Central Place Hierarchy Pattern of 
Central Indiana to the Walter Christaller Model 

Neil V. Weber, Indiana State University 



Abstract 

Walter Christaller's theory on the hierarchy of central places was used as a model 
for the analysis of the distribution of agglomerated centers in central Indiana. The 132 
study centers were analyzed for potential hierarchy breaks relating to the degree of 
centrality of the individual centers. The t test method was used to check whether or not 
significant differences existed between the spacing of Indiana centers and the theoretical 
Christaller pattern. Finally, the Near-Neighbor Analysis was used to test whether or not 
the distributional pattern of the study centers conforms to the maximum dispersal of the 
hexagonal framework. Conclusions drawn from the study affirm the hypotheses examined. 

The purpose of this study was to empirically analyze the basic 
principles of the central place theory. The problem is focused on the 
interpretation of the spatial relationships which are found among the 
central places on the Tipton Till Plain of central Indiana — namely the 
size, spacing and distribution of central places within the study area. 

The method of analysis was concerned with testing two primary 
hypotheses. The first being that certain ordering principles of settle- 
ment exist in nature; second, that there is a distinct relationship be- 
tween the organization and distribution of agglomerated centers in 
central Indiana and the Walter Christaller pattern for southern Ger- 
many. 

Delimitation of Study Area 

The study area consists of the 52-county, 20,362 square mile central 
portion of Indiana. These counties reflect homogeneous unity in that 
they are all directly associated with the physiographic sub-province of 
Indiana known as the Tipton Till Plain. All counties either border on or 
lie within this physiographic unit. 

The Tipton Till Plain is a nearly flat to gently rolling glaciated 
surface (6) with virtually featureless topographic expression (relief 
seldom greater than 50-100 feet). The monotony of the flat-lying land- 
scape is evident throughout the central portion of the state with only 
the extreme southeastern section (the gradational boundary zone) re- 
flecting any topography related to the underlying bedrock (8). 

The Tipton Till Plain exemplifies a homogeneity of economic ac- 
tivity as well as topography. It is an agricultural region (4) contain- 
ing three predominant types of farming: cash-grain in the west, grain- 
livestock in the central portion, and general farming in the east. 

Some portions of the study area deviate from the normal pattern 
described in this section. These particular areas will be discussed in 
detail later in the study. The vast majority of central Indiana, how- 
ever, is a flat-lying agricultural region. 

325 



32(5 



Indiana Academy of Science 



This study is concerned with the distributional pattern of 132 ag- 
glomerated settlements within the study area. These settlements repre- 
sent all centers within the region having populations of 1,000 or more. 
The cut-off limit was set at 1,000 due to the problem of acquiring 
comprehensive data for centers below this figure. 

Evaluation of the Hierarchy Pattern for Indiana Centers 

Population is a common measurement of the importance of place 
(3). It may, therefore, serve as a major criterion for determining the 



,000,000- 



First order canter 



Second order centers 



Third order centers 



Fourth order centers 



*•*. 



'• 000 t — i 1 1 1 1 1 1 1 1 1 1 r 

10 20 30 40 50 60 70 80 90 100 110 120 130 140 

RANK 



Figure 1. Rank-size relationship of urban centers in Central Indiana. 



Geology and Geography 327 

functional grade of trade centers. Population gaps in the progression 
from small to large centers would reflect major distinctions in the size 
groups (1). Theoretically, a hierarchy of centers would develop re- 
lating to the degree of centrality of the groupings. 

By plotting the 132 centers on a semi-logarithmic graph according 
to their rank-size value, distinct groupings within the series are ob- 
served (Fig. 1). The progression from largest to smallest is geo- 
metrically constant from 80,000 to 30,000; 23,000 to 11,000; and 9,600 
to 1,000. Distinct gaps in the trend line are evident between 400,000 to 
80,000; 30,000 to 23,000; and 11,000 to 9,600. This pattern would imply 
four well developed hierarchy groupings for the central places dis- 
tributed within the study area. 

The first order or primary group descends to approximately the 
100,000 level. This would include only one center, that of Indianapolis. 
The secondary grouping ranges from 80,000 to 30,000. This grouping 
contains 8 cities. There are 13 tertiary centers ranging from 23,000 to 
11,000; and 110 centers of quaternary level with a population of 9,000 
to 1,000. 

Description of Areas Excluded from Study 

There are two factors which tend to upset the normal pattern of 
homogeneity within the study area. Both have had a noticeable effect 
on the development and distribution of central urbanized centers within 
the region. 

In the south-central portion of the study area, there is a hilly, 
unglaciated surface located within the Norman Upland physio- 
graphic region (5). This is a rugged area with the greatest local relief 
in the state. The natural agglomerating principles on which this study 
is based are restricted to a great extent in topographic regions as com- 
plex as this upland section. Second, in the late 19th century, natural 
gas was discovered in the Muncie- Anderson area (7). The locations 
within this region never developed in such a way that they can be 
classified trade centers within the criteria set forth in this study. 
These centers developed as a result of proximity to a natural resource 
rather than in the context of having a functional service location. Since 
these two elements are not suitable to space and distance calculations, 
they will henceforth be segregated from the rest of the study area by 
suitable boundaries of their own. 

Model Pattern for Indiana Centers 

Having once established a distinct hierarchy pattern for the study 
area, the next step is to analyze the spacing of the centers in light of 
the various groupings. 

Christaller ordered the pattern of development in the form of 
hexagons. Six centers within each net are located at equal distances 
from one another as well as from their focal place. Each center within 



328 Indiana Academy of Science 

the group represents the same magnitude of centrality. The distance 
between centers of each successive group increases by the \/ 3. 

The model from the Tipton Till Plain region is composed of 46 type 
cast centers from the 4 hierarchy groupings. The centers were selected 
on the basis of the best-fit rule. The theoretical distribution based on 
the Christaller scheme portrays the centers symmetrically placed about 
the landscape. 

The primary (first order) city is Indianapolis. It is the most cen- 
tralized of the major cities in the state. Indianapolis was designated 
the state capitol in 1825 largely because of the central location and easy 
accessibility. It continues to function as the major focus of all adminis- 
trative and consumer activity within the study area. 

There are six secondary centers (excluding those in the indus- 
trialized area) fairly evenly spaced around Indianapolis. These are 
Terre Haute-West Terre Haute, Bloomington, Lafayette-West Lafayette, 
Marion, Richmond and Columbus-East Columbus. Each occupies a focal 
point on an apex of the hexagon. 

Between the primary city and the six secondary centers are 
grouped six tertiary centers: Crawfordsville, Frankfort, New Castle, 
Shelbyville, Franklin and Greencastle. Although Greencastle and Frank- 
lin were previously classified fourth order centers, their strategic loca- 
tion with respect to serviceability warrants a third order trade dis- 
tinction. 

Around each of the first, second and third order centers are quater- 
nary centers in clusters of 6 (33 centers in all). These centers range in 
population from 1,000 to a little over 10,000. Although centers such as 
Bedford were previously ranked as third order places, their true trade 
status is more comparable to the quaternary level. Therefore, a few 
such centers have been adjusted to fit the model more correctly. 

Comparison of Spacing Patterns 

In actuality the centers are distributed about the landscape in a 
more random fashion than the model prescribes. However, a statistical 
analysis concerning the spacing of the trade centers gives impressive 
support to the hexagonal theorem. 

The theory states that the mean distance between centers within 
the four groupings is directly proportional with the distance to func- 
tional groups of the next higher order. The mean distance between 
secondary centers is 38.6 miles, while the mean distance to the primary 
city is 58.0 miles. The tertiary cities have a mean distance of 35.5 miles; 
the distance to the secondary cities averages 35.1 miles. A mean dis- 
tance of 20.7 miles separates quaternary centers with the nearest places 
of higher order approximately 20.9 miles away. 

Christaller further projected that each successive hexagonal net 
would increase by the radical of three. This would postulate that by 



Geology and Geography 329 

ascribing a theoretical value to the secondary centers, the lower group- 
ings would approximate the \/3 rule in descending order. 

Based on the mean distance figure, the distance between secondary 
centers is ordered at 58.3 miles. This places the tertiary centers at a 
model distance of 33.7 miles; the actual computed distance is 35.1 miles. 
Quaternary centers should average a distance of 20.4 miles; in reality 
they are 20.9 miles. 

When observed distances and theoretical distances are analyzed 
quantitatively, the results reflect a minimal aberration among the units 
tested. The average variation is computed to be 3.18% with a correla- 
tion coefficient of 0.9984 (Table 1). 

Table 1. Comparison of spacing patterns. 



Hierarchy Groupings 



2nd to 1st 58.0 58.3 —.52 (.0032) 

2nd to 2nd 58.6 58.3 +.51 

3rd to 2nd 35.1 33.7 +4.15 

3rd to 3rd 35.5 33.7 +5.34 



Mean 


Theoretical 


Variation 


Measured 


Distance 


from 


Distance 


( Miles — Based 


Theoretical 


(Miles)* 


on V 3 > 


(Percentage) 





4th to 3rd 


22.6 


20.4 


+ 10.78 




4th to 2nd 


20.5 


20.4 


+ .49 


(nearest) 


4th to 1st 


19.6 


20.4 


—3.92 


(around 3rd) 


4th to 4th 


20.7 


20.4 


+ 1.47 


(around 2nd) 


4th to 4th 


20.7 


20.4 


+ 1.47 



* The distance measurements were taken from a 1968 Indiana Chamber of Commerce 
road map. Calculations were based on straight line distances. It was found that straight 
line distances between centers averaged from 77,-19% less than road distances. Therefore 
a mean correction factor of 13% should be applied to the straight line distances to obtain 
the approximate read distances between centers. 



Therefore one may conclude that the centers of this region tend 
to lie in a uniform pattern about the landscape, evenly spaced within 
their own unique groupings as well as externally between groupings. 
The distance between the various hierarchies conforms directly with 
Christaller's observed tendency toward W3 as the set norm of increase. 

Analysis of Deviations from Theoretical Locations 

The theoretical hexagon system was rotated to a position of best 
fit with reference to the observed location of the model centers (Fig. 2). 
Best fit was determined by calculating the position which warranted the 
lowest sum departure of the observed secondary centers from their 
theoretical locations. After the theoretical pattern had once been fixed 



330 



Indiana Academy of Science 



in position, departures were calculated for each of the model centers. 
These figures were then analyzed to determine to what degree each 
hierarchy grouping correlated with the Christaller framework. 

A "t" test was used to test the significance of the difference be- 
tween the observed and the theoretical distances to adjacent centers. 
The test was computed for each of the three groups in an attempt to 
determine the amount of uniformity within units. The result from the 
"t" test showed that in all 3 groupings the observed distances and the 
theoretical distances were not significantly different at the 0.05 level. 





LEGEND 

Primary center 

Secondary canters 

Tertiary centere 
Quaternary centers 



FIGURE 2. Comparison, of theoretical pattern and model centers. 



Geology and Geography 331 

Therefore, one may conclude that even though the centers do devi- 
ate to a certain degree from their theoretical location, the spatial 
patterns are similar enough to be considered statistically related. 

Near-Neighbor Analysis 

To test if the centers were located in a pattern of maximum dis- 
persal (which is in essence what the hexagonal theorem prescribes), 
the spacing of each unit with respect to its adjacent settlements must 
be analyzed. 

Examination of this pattern can be accomplished by the use of 
the Near-Neighbor Analysis. This technique indicates the degree to 
which any observed distribution of points deviates from what might be 
expected if the points were distributed in a random manner within the 
same area (2). The theory states that there are three distinct patterns 
of settlement: Aggregate (clustering), Random and Uniform. 

The mathematical test for the Near-Neighbor pattern assigns an 
"R" value for the observed density of points in the area under con- 
sideration. An "R" value of less than 1.00 represents a tendency toward 
clustering (0.00 equals a perfect aggregate pattern). An "R" value of 
greater than 1.00 reflects a tendency toward uniform spacing (2.15 
equals maximum dispersal or the perfect hexagonal pattern). 

The following results were obtained by use of Near-Neighbor analy- 
sis. The "R" for the second order grouping is 2.011. Deviation is 0.139 
(6.4%) from the perfect hexagonal framework. The tertiary centers 
have an "R" of 2.126. This pattern's deviation is less than 2% (0.024) 
from the maximum hexagonal dispersal. The "R" for the fourth order 
centers is 2.060 with a departure from uniformity of only 0.090 (4.1%). 

Therefore, one may conclude on the basis of the aforementioned 
data, that there is a distinct distributional pattern of settlement for 
this region. The model centers do in effect reflect a near perfect rela- 
tionship with the uniform spacing (2.15) tendency of the Near-Neighbor 
equation. 

Conclusion 

The results of this study clearly support the two hypotheses tested. 
Nature dictates a certain degree of order to the development of central 
places. The central places of central Indiana have a tendency to conform 
to such an ordering principle; and the pattern for this area is very 
much similar to the K-3 network of Walter Christaller. 



Literature Cited 

1. Berry, B. J. L., and W. L. Garrison. 1958. Functional bases of the central place 
hierarchy. Econ. Geogr. 34:145-154. 

2. Berry, B. J. L., and D. F. Marble. 1968. Spatial Analysis : A Reader in Statistical 
Geography. Prentice-Hall, Inc., Englewood Cliffs, N.J. 512 p. 



332 Indiana Academy of Science 

3. BRUSH, J. E. 1953. The hierarchy of central places in southwestern Wisconsin. Geogr. 
Rev. 43 :380-402. 

4. Hart, J. F. 1968. Field patterns in Indiana. Geogr. Rev. 63:450-471. 

5. Kingsbury, R. C. 1966. An Atlas of Southern Indiana. Occasional Pub. No. 3. Dept. of 
Geogr. Indiana Univ., Bloomington. 60 p. 

6. Schneider, A. F. 1966. Physiography, p. 40-56. In A. A. Lindsey [ed.] Natural Fea- 
tures of Indiana. Indiana Acad. Sci. Sesquicentennial Volume. Indianapolis. 600 p. 

7. Visher, S. S. 1923. Economic Geography of Indiana. D. Appleton and Co., N.Y. 511 p. 

8. Wayne, W. J. 1956. Thickness of drift and bedrock physiography of Indiana north of 
the Wisconsin glacial boundary. Progress Rep. No. 7. Indiana Geol. Surv., Bloomington, 
Ind. 



Parameter Measurement in Fluvial Morphology 

Daniel M. Coffman, Purdue University 



Abstract 

Because of the quantitative significance of stream order, an accurate, ordered drain- 
age map has become a base for much research in fluvial morphology. These maps are 
often prepared from U. S. Geological Survey 1:24,000 topographic maps and doubt is 
created about their usefulness as sources of primary data. This study indicates that 
blue lines of such topographic maps represent third- and fourth-order streams. Even 
interpretation of topographic maps by Strahler's method of "V's" does not necessarily 
result in an accurate drainage map. Relief, age, and geology appear to control the 
quality of drainage maps which are produced. The relative relief of apparent first-order 
streams may be used to predict the quality of a drainage map which will be produced 
from a topographic quadrangle and permits classifications of these quadrangles as 
excellent, marginal, and unacceptable. 



Introduction 

For nearly 25 years attempts to quantitatively define river systems 
and landforms derived from their development have stemmed from Hor- 
ton's concept of an ordered stream system (3). The advantage of stream 
order, in a given physiographic region, is its relation to the number, 
average lengths, gradients, drainage area, and perhaps discharge of 
stream segments of varying size. A modification of Horton's work 
whereby the smallest stream tributaries are assigned the lowest order 
was introduced by Strahler (8) and resulted in the system of stream 
order which is widely used today. An example of a stream ordered ac- 
cording to Strahler's system is shown in Figure 1. Because of stream 
order's quantitative significance, an accurate, ordered drainage map 
has become a base from which research into fluvial processes normally 
begins. 

The results of any research are no better than the quality of 
original input data, and this is especially true in the study of stream 
patterns, geometry, and mechanics. Here the raw data generally in- 
volve measurement of the number, length, and gradient of stream 
segments; width, depth, and velocity of water in the channel; and 
area, main stream length, and shape of the drainage basin. If these 
parameters are measured and mapped in the field the quality of the 
data is generally excellent. Unfortunately, few researchers have time 
or money to devote to this procedurally difficult type of data collection, 
resulting in the use of topographic maps as a major source of original 
data for studies in fluvial morphology. 

The purpose of this paper is to examine the problems involved in 
obtaining an accurately ordered drainage map from commonly available 
U.S.G.S. topographic maps. This study was sponsored by the Purdue 
University Water Resources Research Center, Office of Water Resources 
Research, but this paper has not had the benefit of its review. The 
author also acknowledges the assistance of Dr. W. N. Melhorn and 

333 



334 Indiana Academy of Science 

Prof. R. D. Miles in discussions relating to aspects of fluvial mor- 
phology. 

Measurement from Maps 

In the field, first-order stream channels (defined as the smallest 
unbranched tributaries of a river system) are easily recognized result- 
ing in the production of a very accurate drainage map. The U.S.G.S. 
topographic maps on which many studies rely, however, do not gen- 
erally show all stream channels. Apparent first-order stream segments 
identified from these maps thus become functions of map scale and the 
type of map interpretation employed, and in the published literature 
may not represent from article to article, the same size stream channel. 
As a result, parameters such as average length may be correlated with 
stream orders which are incorrectly numbered. 

Because of this common practice of compiling stream data directly 
from topographic maps, much effort has been devoted to determining 
the validity of ordering streams from the various types of maps com- 
monly available. Morisawa (5) concluded that U.S.G.S. topographic 
maps scaled 1:62,500 are not reliable for measuring drainage basin 
characteristics other than area. She also stated that data on numbers 
and lengths of stream segments show great variation when taken from 
maps. Coates (2), in working with streams in southern Indiana, found 
that on U.S.G.S. maps scaled 1:24,000 first- and second-order streams 
were rarely shown and that most channels interpreted as first-order 
were actually third-order. Strahler (9) proposed an improvement in 
interpreting topographic maps, by adding to those stream channels 
shown in blue, segments where "V's" in contour lines indicate that 
valleys are present. This method greatly aids in adding otherwise over- 
looked stream segments to the drainage net, but produces an un- 
certainty as to the size of the smallest segment now shown. 

Because of the ever increasing need for basic data, some research- 
ers have ignored published warnings about the questionable validity 
of a drainage map prepared from topographic maps. One example is 
provided by Stall and Fok (7) who ordered drainage basins from 
U.S.G.S. topographic maps scaled 1:62,500 without interpretation by the 
method of "V's." Their correlation of discharge, drainage basin area, 
length, and slope which they ascribe to low order segments must con- 
servatively belong to fourth- and fifth-order streams. Although some 
authors merely bypass the question of their data's validity, Scheidegger 
(6) suggested what the techniques of other investigators appear im- 
plicitly to assume; that if, for instance, second-order streams are 
treated as first-order streams because of map inaccuracy the results 
obtained in a study are proportional to the parameters of the actual 
stream system. If this is true, after a field check the values of the 
actual system may be obtained from the study by using Horton's law 
of drainage composition or some other mathematical tool of extrapola- 
tion. 



Geology and Geography 335 

Objectives of this Study 

Publication of the Atlas of County Drainage Maps for Indiana in 
July, 1959, permits large areal studies of stream ordering- to be con- 
ducted while maintaining a high level of confidence in the data (1). 
These maps were prepared by the staff of the airphoto laboratory of 
the Engineering Experiment Station at Purdue University from 1940 
A. A. A. aerial photographs. Depending on the time of year that photo- 
graphs were taken these maps are estimated to show an average of 
90% of all existing stream channels (R. D. Miles, personal communica- 
tion). A field check in the vicinity of Tippecanoe County demonstrated 
that these maps show almost all first-order segments. Those stream 
channels not shown appeared young, probably having developed since 
the date of photography. 

Using these maps as a base, a study was conducted to answer the 
following questions about drainage maps prepared from U.S.G.S. topo- 
graphic maps scaled 1:24,000 with 10-foot contour interval: 

1) What portion of a drainage basin and what order stream seg- 
ments are actually shown by the blue lines of these maps ? 

2) What increase in accuracy is obtained by interpreting these 
maps using Strahler's method of "V's" ? 

3) Are drainage maps produced by analysis of data in the above 
two cases proportional to the actual stream net, and if not, what 
effect do lost segments have on ordering the basin? 

4) Does accuracy of these maps vary with location in the state of 
Indiana ? 



Procedure 

Drainage basins up to sixth-order in size were selected randomly 
from the different physiographic provinces of the state (4). Topographic 
maps for these basins were obtained and drainage maps were produced, 
initially by tracing only those streams shown by solid or dashed blue 
lines, and secondly by adding all stream segments which could be 
recognized using the method of "V's." Experiments with different oper- 
ators indicated that more than a general knowledge of topographic 
maps was required to apply the method of "V's" and that some practice 
was necessary before any operator made the best, consistent use of this 
method. Once familiar with the method, however, different operators 
could produce reasonably identical maps of the same drainage basin. 

Once the two tracings were prepared they were ordered and com- 
pared with an ordered base map taken from the Atlas of County Dram- 
age Maps. The data for this segment-to-segment comparison was 
punched on IBM cards and analyzed by a CDC 6500 computer. 



336 



Indiana Academy of Science 



■8-2 

* * 

Q P 

-■ 

o 

E 



<U ft 



<J EC 



83 2 

< 



C c to « 

S I B W 

Sis S» 

* m P 



U ft 

Ph ft 



ft 

E 



O M 

0) 



ft X 

ft H 
<! 



w P 



+> GO ° g 

Slew 
« S ai a 

££ £ £ 



15 

Sg. 

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Geology and Geography 



337 



Table 2. Accuracy of U.S.G.S. maps according to physiographic province. 



Physiographic Province 



Range of 

General 

Relief 



Geology 



Percent of Various 


Se K 


men 


ts Shown 




on 


Map 




1 


2 


3 


4 


its 


LOO 


LOO 


100 


97 


LOO 


LOO 


100 


97 


LOO 


LOO 


100 


89 


96 


100 


100 


79 


90 


100 


100 


(57 


90 


99 


100 


19 


4X 


91 


100 


15 


45 


100 


100 



Crawford Upland 
Norman Upland 
Dearborn Upland 
Scottsburg Lowland 
Muscatatuck Slope 
Wabash Lowland 
Tipton Till Plain 
Lake and Moraine 
Mitchell Plain 
Indiana 



220-350 Mississippian SS, LS, SH 

170-260 Mississippian SS, SH 

120-250 Ordovician LS 

100-150 Devonian & Mississippian SH 

100-150 Silurian LS 

100-150 Pennsylvanian SH, SS 

100-150 Glacial Till 

80-120 Fluvial Glacial 

Underground Drainage 
All Provinces Using Only the Blue Lines 



Results of Computer Analysis 

The percentage of each order of stream segments shown by blue 
lines on the topographic maps was found to be about the same for all 
physiographic provinces of the state (see "Indiana" in Table 2). Table 1 
shows results for basins from the Tipton Till Plain. These findings 
agree well with those of Coates (2) as almost no first- and second-order 
streams are shown. From one-third to one-half of the third-order seg- 
ments appear and almost all fourth-order segments, are shown. As the 
breakdown into apparent order indicates, nearly all third-order streams 
and one-half of the fourth-order streams would be classed as first-order 
if only blue lines were used to construct a drainage map. Of 4,498 seg- 
ments from various basins used in this analysis, only 153 or about 4% 
of all segments were shown in blue on the 1:24,000 U.S.G.S. topographic 
maps. 

An increase in the number of stream segments located using the 
1:24,000 U.S.G.S. topographic maps was found to be significant in every 
physiographic province if interpretation by Strahler's (9) method of 
"V's" was employed. Table 1 shows that in the Tipton Till Plain about 
Vs of the first-order, V 2 of the second-order, and almost all of the third- 
order channels are shown. The proportion of total segments shown in- 
creased from 4<7r to 28% in this province, whereas in other physiographic 
provinces of the state the method of "V's" resulted in almost 100% re- 
covery of all stream segments. Even in areas where this method proved 
most successful, however, it still contained two major uncertainties. First, 
the correct length of an added segment was always in doubt, commonly 
by a factor many times the contour interval. Second, many segments 
were recognized whose point of junction with other channels was 
uncertain, which ultimately could result in incorrect ordering of the 
stream net. 

In provinces where almost all stream segments in a basin are 
obtained using the method of "V's," only the uncertainty of junction 



338 



Indiana Academy of Science 




Us 


N 


L 


Us 


N 


L 


1 




236 


.2 


4 


--. 


9 


1.6 


2 




81 


.4 


5 


-== 


3 


2.6 


3 





27 


.8 


6 


1 — - 


1 


- 



R = 3 



Figure 1. Hypothetical stream. 



locations will cause lack of proportionality between actual and traced 
stream nets. However, if a portion of segments is missed, either 
because of the map or inaccurate procedures used in tracing a stream 
net, the ordered stream system which results will be in no way propor- 
tional to the real stream system. To emphasize this point, consider the 
stream system shown in Figure 1. It has a perfect bifurcation ratio (R) 
of three, and the numbers (N) and average lengths (L) for the various 
orders are indicated. Figure 2 is the same drainage basin without first- 
and second-order segments. This remains a reasonable representation 
of the original basin and provides an example of the stream net which 
some researchers assume is obtained by ordering stream systems from 
large-scale topographic maps. The bifurcation ratio is unaltered and 



Geology and Geography 



Mi) 




Us 


N 


L 


Us 


N 


L 


1 




27 


.8 


3 





3 


2.6 


2 




9 


1.6 


4 


. 


1 


- 



Figure 2. Hypothetical stream without 1st and 2nd orders. 



after a field check the laws of drainage composition would allow calcula- 
tion of the numbers and average lengths of stream orders not shown by 
this map. 

Figure 3, however, shows what actually happens when a stream net 
is ordered directly from the blue lines of a U.S.G.S. topographic map 
scaled 1:24,000. Since approximately only V 3 of the third-order stream 
segments are shown, the ordered basin is no longer proportional to the 
real stream net. The bifurcation ratio is altered and the average seg- 
ment lengths become distorted. It is impossible to extrapolate the prop- 
erties of the original stream net from this ordered basin. Using the 
data for the Tipton Till Plain, Figure 4 shows a theoretical drainage 
map produced from a U.S.G.S. topographic map using the method of 
"Vs." It shows more segments than the stream net in Figure 3, but the 



340 



Indiana Academy of Science 




Us 


N 


L 


Us 


N 


L 


1 , 




10 


2.0 


3 





1 


6.0 


2 




4 


.9 


4 








- 



Figure 3. Stream as it might appear on USGS map. 



bifurcation ratio is still altered and the measurement of average lengths 
is meaningless. 

The ability of interpretation using the method of "V's" to add seg- 
ments to the blue lines of topographic maps was found to vary consid- 
erably depending on location in the state. This variability appears to be 
a function of 1) basin relief, 2) age of development of topography 
and 3) geology. Basin relief (the difference in elevation from the 
drainage divide to the mouth of the stream) is an indication of slope. 
The greater the slope the more contour lines a given length of stream 
crosses and thus the easier it is to map. As the length of time during 
which the topography has been evolving increases, low order streams 
develop more distinct channels, commonly rectangular in shape, result- 
ing in more distinct "V's" in the contour lines. Geology effects resistance 



Geology and Geography 



341 




Us 


N 


L 


Us 


N 


L 


1 




43 


.7 


3 





4 


2.1 


2 





12 


1.5 


4 


•m^» 


1 


— 



Figure 4. Stream as it might be mapped using method of V 



to erosion and thus controls the rate of gully development. Some geologi- 
cal materials are characterized by shallow channels whereas others 
possess steep, deep channels more likely to produce pronounced "V's" in 
contour lines. 

Because physiographic divisions of the state reflect differences in 
relief, age, and geology some generalizations about the topographic 
maps covering these areas may be made. Table 2 lists the divisions and 
indicates the percent of various order streams which would likely be 
obtained by interpreting a 1:24,000 U.S.G.S. topographic map using 
Strahler's (9) method of "Vs." This table intends to show only average 
conditions and because of variations within any province it should not 
be used as an accurate guide for all topographic quadrangles within this 
physiographic division. No data were taken for the Mitchell Plain 



342 



Indiana Academy of Science 



because drainage in that province is essentially underground. Data for 
the Lake and Moraine Province are poor and inconclusive because of 
extensive development of ditch systems over most of the area, which are 
mapped in detail as cultural features on U.S.G.S. quadrangles. 

Estimation of Map Accuracy 

As most first- and second-order streams are tributary to third-order 
segments, and because they account for the majority of those segments 
not shown on the topographic maps, a method to estimate their presence 
or absence was sought. It has been demonstrated that the smallest 
dashed, blue lines on a U.S.G.S. quadrangle represent third- and 
fourth-order channels and that relief is of prime importance in the 
ability to interpret a map using the method of "Vs." Thus, the average 
relative relief of apparent first-order streams was compared with the 
quality of interpretation using the method of "Vs." Ten apparent first- 
order segments were chosen from a basin (or quadrangle if the basin 
was small) and their relative relief calculated. Highest and lowest 



100 



KEY 




• 




LAKE AND 




MORAINE 




PROVINCE 




O 




TIPTON 




TILL 


a 

< 
2 


PLAIN 


A 


(0 


WABASH 


© 


LOWLAND 


u> 




Z> 


X 


z 




o 


CRAWFORD 


z 


UPLAND 




o 




I 


D 


v> 


NORMAN 


t- 


UPLAND 


z 

kJ 




2 


© 


Ui 


SCOTTSBURG 


V) 


LOWLAND 


z 




< 




UI 


A 


a: 

l- 


MUSCATATUCK 


V) 


SLOPE 


u. 
o 




*- 


■ 


z 

ID 


DEARBORN 


o 
a: 


UPLAND 


Ui 

a. 




50 100 150 50 100 

AVERAGE RELATIVE RELIEF OF APPARENT FIRST ORDER STREAMS 

Figure 5. Plot of average relative relief versus percent of streams. 



Geology and Geography 343 

values were discarded and the average relative relief of the eight 
remaining segments was plotted against the percent of first-, second-, 
and third-order segments shown on the map. Results as shown in 
Figure 5 indicate that the average relative relief provides a good 
estimate of the accuracy of a single U.S.G.S. quadrangle map. 

Average relative relief of apparent first-order streams has been 
used to divide U.S.G.S. quadrangles into three separate classes; excel- 
lent, marginal, and unacceptable. Quadrangles with an average relative 
relief of 150 feet or more are excellent, showing all stream segments of 
all orders, and may be used with confidence to produce an accurate 
drainage map. Quadrangles with an average relative relief less than 80 
feet are unacceptable because they fail to show first-, second-, and some 
third-order segments, making any drainage map produced from them 
unproportional and unreliable. 

If average relative relief is between 80 and 150 feet the topographic 
map is marginal as a base for production of an accurate drainage map. 
Assuming random failure to detect "lost" stream segments the mini- 
mum acceptable omission of first-order streams may be calculated using 
the formula 

2 

(1 ) • (100) where R = bifurcation ratio 

R 

and is compared to the probable percentage of first-order streams as 
obtained from Figure 5 after calculating the average relative relief. If 
enough first-order segments are shown the quadrangle should produce 
a drainage map proportional with the real system. If R is not known it 
is recommended that three be used because it is the worst possible value. 

Conclusions 

Unless drainage maps are produced by field mapping or from 
interpretation of large-scale aerial photographs their reliability as 
accurate models of the real stream net must be initially questioned. No 
acceptable drainage map can be made from a U.S.G.S. topographic map 
scaled 1:62,500 without considerable field verification. If drainage maps 
are traced from 1:24,000 U.S.G.S. topographic maps, using only the 
streams shown by blue lines, the apparent first-order segments actually 
will represent third- and fourth-order streams. If these quadrangles 
are interpreted using the method of "V's" uncertainty about order, 
length, and junction location of added segments is introduced. 

The quality of interpretation by the method of "V's" varies as a 
function of 1) basin relief, 2) age of topography, and 3) geology. An 
approximate guide to the quality of a drainage map produced from a 
given topographic map is thus the physiographic province from which it 
comes. It has been demonstrated that the average relative relief of 
apparent first-order stream segments shown by blue lines can be used 
to predict the quality of a topographic map for use in preparing an 
accurate drainage map. Based on this average relative relief the 



344 Indiana Academy of Science 

1:24,000 U.S.G.S. topographic maps for Indiana may be classed as 
excellent, marginal, and unacceptable. 

Literature Cited 

1. Atlas of County Drainage Maps of Indiana. 1959. Purdue Univ. and State Highway 
Dept. Ext. Ser. Rep. No. 97. 

2. Coates, D. R. 1958. Quantitative geomorphology of small drainage basins of southern 
Indiana. Columbia Univ. Dept. Geol. Tech. Rep. 10 : 67 p. 

3. Horton, R. E. 1945. Erosional development of streams and their drainage basins ; 
hydrophysical approach to quantitative morphology. Geol. Soc. Amer. Bull. 56 :275- 
370. 

4. Malott, C. A. 1922. The physiography of Indiana, p. 57-257. In W. N. Logan and 
others. Handbook of Indiana geology. Pub. 21 Indiana Dept. Cons., Indianapolis. 

5. Morisawa, M. E. 1959. Relation of quantitative geomorphology to stream flow in 
representative watersheds of the Appalachian Plateau Province. Columbia Univ. Dept. 
Geol. Tech. Rep. 20 : 94 p. 

6 Scheidegger, N. A. 1965. The algebra of stream-order numbers. U. S. Geol. Surv. Prof. 
Paper 525-B : B187-B189. 

7. Stall, J. B., and Yu-Sl Fok. 1968. Hydraulic geometry of Illinois streams. Univ. of 
111. Water Resources Center Rep. 15 : 47 p. 

8. Strahler, A. N. 1952. Hysometric (area-altitude) analysis of erosional topography. 
Geol. Soc. Amer. Bull. 63:1117-1142. 

9. Strahler, A. N. 1957. Quantitative analysis of watershed geomorphology. Amer. 
Geophys. Union Trans. 38 :913-920. 



MICROBIOLOGY AND MOLECULAR BIOLOGY 

Chairman: Warner S. Wegener, Indiana University Medical Center 
Larry Day, Eli Lilly & Co., was elected Chairman for 1970 

ABSTRACTS 

Stream Pollution from Coal Mine Waste Piles: Effect of Sulfur and Iron 
Oxidizing Bacteria. Robert Ramaley and Richard Kindig, Indiana 
University. — Analysis of 100 samples of stream water taken during 
hydrological studies of the Patoka (Pike County, Ind.) and the 
Busseron (Sullivan County, Ind.) watersheds showed a correlation 
between the acidity of the samples and the number of sulfur or iron 
oxidizing bacteria (Thiobacillus-F 'errobacillus) . 

Data obtained during local washouts of surface mined areas and 
experiments with sterilized mine waste material are consistent with the 
production of the acid pollution and perhaps some of the oxidized iron 
by the action of sulfur and iron oxidizing bacteria on sulfur bearing 
mine wastes and a subsequent washing out of both the acid and some 
of the bacteria into the streams following rainfall. 

Core samples taken of land reclaimed in accordance with the 1967 
Indiana Mining Act showed that such reclamation was effective in 
reducing the number of sulfur and iron oxidizing bacteria in the covered 
mine maste material and thereby decreasing the acid, production to 
negligible amounts. 

Fine Structure Changes during Germination of Dictyostelium dis- 
coideum Spores. David A. Cotter. Indiana University Medical Center. 
—The earliest developmental stage in the life cycle of the cellular 
slime mold, Dictyostelium discoideum, is spore germination. Spores of 
this cellular slime mold can be induced to germinate by exposure to a 
mild heat shock of 45° C for 30 minutes. The spore germination process 
occurs in four well-defined stages: 1) dormant spore activation, 2) 
post-activation lag, 3) spore swelling, and 4) myxamoeba emergence. 

Electron microscopy revealed significant changes in the fine struc- 
ture of germinating spores during stages three and four. The mito- 
chondria progressively became less dense, lost their peripherally 
attached ribosomes, and revealed more pronounced tubuli as germina- 
tion proceeded. During spore swelling, the three-layered spore wall 
broke down in two stages: 1) the outer and middle layers were rup- 
tured as a unit; and 2) the inner wall was breached. Crystals and dark 
(lipid) bodies seemed to disappear shortly before or during emergence 
of the myxamoebae. Autophagic vacuoles were found in dormant spores 
and throughout the entire germination process. 

The addition of cycloheximide to germinating spores inhibited the 
loss of the crystals and dark (lipid) bodies. In addition, the drug 
inhibited the breakdown of the inner wall layer. Cycloheximide did not 

345 



346 Indiana Academy of Science 

prevent the formation of the water expulsion vesicle or the apparent 
function of the autophagic vacuoles. A hypothesis is offered to explain 
the inhibition of myxamoebae emergence by this drug. 

Thermal Induction of Bacteriophage PL S. S. Lee, Indiana University 
Medical Center. — The induction of a triply auxotrophic (thy , ura", met) 
Escherichia coli lysogenic for PI was studied under different nutritional 
and temperature conditions. At 37° C, as reported by others, induction 
occurs only under thymineless conditions. At 45° C, induction occurs 
under thymineless conditions, but also under conditions of amino acid 
deprivation. Although thermal induction appears to be a different 
process than thymineless induction, the fact that the two interact 
synergistically suggests that both involve repressor inactivation. 
Thermal induction is generally less complete than thymineless induction 
at 37° C. The fact that pre-treatment at 37° C under condition of 
amino acid deprivation lowers subsequent thermal induction under con- 
ditions where thymine, uracil and methionine are all withheld suggests 
that cells that have just completed one round of DNA replication and 
have not initiated another round are relatively insensitive to thermal 
induction. Finally, non-induced cells undergo little, if any, thymineless 
death at 45° C, although induced cells slowly lose plaque-forming 
ability at this temperature. 

Computer-based Derivation of Rate Equations for Enzyme-catalyzed 
Reactions. Arthur R. Schulz and Donald D. Fisher, Indiana Univer- 
sity Medical Center. — An algorithmic process has been developed which 
provides for derivation of rate equations for enzyme-catalyzed reactions 
by a digital computer. An explicit notation system is employed which 
is adaptable to computer processing techniques. The sequence of an 
enzymic reaction is represented by two matrices. A connection matrix 
is used to determine the valid paths which connect the enzyme species, 
and a matrix of substrate and product names is used to append 
reactant symbols to the proper path vectors. 

The denominator of a rate equation is given by the sum of the 
valid paths which connect the enzyme species. The algorithm provides 
for derivation of the numerator of the rate equation and for alphabetical 
sorting of the numerator and denominator terms. Reformulation of the 
rate equation from the "coefficient" form to the more useful "kinetic" 
form is accomplished by expressing each denominator term as a vector 
of reactant concentration exponents. This provides for computer-based 
definition of the Michaelis constants and for all possible inhibition con- 
stants, and for the reformulation of the equation using the definitions 
selected by the investigator. 

Axoplasmic Transport of Materials in Nerve Fibers. S. Ochs, M. I. 
Sabri, and N. Ranish, Indiana University Medical Center.— Axoplasmic 
flow is present in nerve fibers. Our recent studies have shown that after 
lumbar seventh ganglion injection with HMeucine, a crest of labelled 
activity is present in the sciatic nerve with the position of the crest 



Microbiology and Molecular Biology 347 

moving outward at a rate of 400 mm per day. Evidence that the labelled 
activity is intra-axonic was gained by freeze block. Extraction of labeled 
material showed it to be present in particulate and soluble form, as high 
molecular weight protein (450,000 and 65,000) and polypeptide 
(4-13,000). The precursor P :i2 -orthophosphate is incorporated into a 
slower moving material. The possible molecular basis of the fast trans- 
port system and other studies regarding the mechanism underlying 
intra-axonic transport were discussed. 



Development of a Clear, Photopolymerizable Acrylamide Gel and Its 
Use In Immobilizing and Staining Nucleic Acids. 

Thomas A. Cole and Wayne F. Middendorf, Wabash College 

Abstract 

A clear, photopolymerizable gel of a poly acrylamide has been developed. The com- 
ponents will form a gel in presence of high concentrations of cesium chloride. DNA 
immobilized in such a gel can be stained and the blank gel subsequently destained. The 
application of this technique to isopycnic density gradient centrif ligation of cesium 
chloride solutions was discussed. 

Introduction 

Polymerization of acrylamide and N,N'-methylene-bis-acrylamide 
has been widely used in the preparative, routine and analytical versions 
of disc electrophoresis (1). Additionally, photopolymerization of these 
compounds have been used in conjunction with sucrose density gradient 
centrifugation for immobilization of components separated by centri- 
fugation (2, 3, 4). This report extends the immobilization technique to 
dense solutions of cesium chloride and details the development of a 
clear photopolymerizable gel. The techniques of staining and destaining 
the immobilized gel are reported also. 

Methods and Materials 

Preparation of Cesium Chloride-Containing Gels 

Cesium Chloride (SC11352) was purchased from the Sargent-Welch 
Company. Cesium chloride was dissolved in water to give a solution 
with a density of around 1.875 gm/ml. Such a solution requires about 
1.200 gms CsCl/ml solution at 20 °C. Three parts of the cesium chloride 
solution were mixed with one part of polymerizable solution. 

Polymerizable solution is made of three parts, Solutions B and C and 
riboflavin. Solution B consists of 11.4 gm Tris buffer, 1.2 ml N,N,N',N'- 
tetramethylethylenediamine (TEMED, practical grade, Matheson Cole- 
man and Bell 8563) and distilled water to a final volume of 33 ml. The 
pH is adjusted to 6.9 with 85 percent phosphoric acid. Solution C is 
24 gm of acrylamide, 0.735 gm N,N'-methylene-bisacrylamide (Bis, East- 
man 8383) and distilled water to give a final volume of 50 ml. Approxi- 
mately 0.5 mg riboflavin/5 ml aliquot is dissolved in the polymerizable 
solution. Solutions B and C are stable and may be stored at room tem- 
perature. After addition of riboflavin the polymerizable solution must be 
protected from light. 

Layers of gel were successively polymerized in a cellulose nitrate 
centrifuge tube. Alternate layers contained DNA. Each layer was 
polymerized from 0.4 ml of solution (0.1 ml of Cesium chloride solution, 
0.3 ml of polymerizable solution) with a thin (2 mm) overlay of water. 
Those layers which contained DNA were made by diluting stock DNA 

348 



Microbiology and Molecular Biology 349 

solution with cesium chloride-polymerizable solution. Polymerization was 
completed for each layer in 10 to 20 minutes at a distance of 4 inches 
from a 15 watt fluorescent bulb. 

Staining and Destaining 

After polymerization the gels were removed from the cellulose 
nitrate tubes by shaking or rimming with a microspatula and were 
stained by one of a variety of procedures. Three procedures gave the best 
results and they are reported here. A methyl green procedure (5, 6) gives 
a green stain, pyronin B (5) gives a lavender stain and methyl green and 
pyronin B together (7) give a blue-green stain. The gels were immersed 
in the staining solution for 1 hour or more. The excess stain was re- 
moved by repeated washing in 0.2n acetate buffer of an appropriate pH 
(4.0 for methyl green, 4.5 for pyronin B and 4.25 for methyl green-py- 
ronin B). Electrophoretic destaining may be done in the same buffers. 
A suitable destaining apparatus may be constructed from lucite (8). The 
washing procedure takes about 2 days with occasional changes of buffer. 
The electrophoretic procedure requires about 24 hours at 3-4 ma of 
current per tube. 

DNA Samples 

Calf thymus DNA (Sodium Salt, Type I, lot 115B-1690) was pur- 
chased from Sigma Chemical Company, St. Louis, Missouri. A stock 
solution of DNA was prepared by dissolving DNA in dilute NaCl-citrate 
(0.015m NaCl, 0.0015m sodium citrate, pH 7). 

Results 

The results show that within the range of approximately 100 to 250 
/xg calf thymus DNA/ml good contrast bands were obtained. At 
500fig/ml the DNA partially inhibited the polymerization process. 
Consequently, the 500^g/ml band was small due to loss of material. 

Discussion 

The photopolymerizable gel of disc electrophoresis (1) and sucrose 
density gradient techniques (2, 3, 4) is translucent. The transparent gel 
reported here is the result of lowering the Bis/acrylamide ratio. This 
means that the transparent, photopolymerizable gel is not as highly 
cross-linked as the translucent one. 

The results reported here show that DNA can be immobilized, 
stained and destained in a gel containing cesium chloride at concen- 
trations comparable to those used in isopycnic density gradient centri- 
fugation. Thus, cesium chloride does not interfere with the polymeriza- 
tion process. 

The methyl green stain has been used successfully for staining 
native DNA in acrylamide gels (5); since these gels are much thicker 
than those of disc electrophoresis the primary exposure time to the 
stain was 1% more hours rather than 1 hour as reported by Boyd and 



350 Indiana Academy of Science 

Mitchell. The pyronin B method has been used for denatured DNA (5). 
Presumably, our DNA samples were not denatured and the pyronin B 
method gave the least satisfactory results. The methyl green-pyronin 
B method gave the best contrast but the methyl green alone could be 
completely removed from non-DNA containing areas of the gel. 

With the techniques developed here, it should be possible to carry 
out isopycnic cesium chloride density gradient centrifugation with DNA 
and stain the samples after polymerization. With disc electrophoresis 
many staining procedures have been quantitated with a microdensi- 
tometer scanning of the gel (1). This instrument costs $3900. Recently, 
an inexpensive high-resolution densitometer for disc electrophoresis 
has been developed (9). The total cost for materials for this instrument 
are reported not to exceed $300. Thus, a guantitative and analytical 
data-acquisition system for density gradient centrifugation should be 
possible without the use of an analytical ultracentrifuge (Model E, 
Spinco Division, Beckman Instruments, Inc., Palo Alto, California 94304) 
or drop collecting techniques. The requirements for such a system are 
a preparative ultracentrifuge, the immobilization and staining pro- 
cedure reported here, and a microdensitometer for gel scanning. 



Acknowledgements 

This work was supported by Grant NIH-GM-11860, U. S. Public 
Health Service, and an NSF-COSIP grant to Wabash College. 



Literture Cited 

1. Davis, Baruch J. 1964. Polyacrylamide Gel Electrophoresis. Ann. New York Acad, 
of Sci. 121 :404-427. 

2. Jolley, Weldon B.. H. W. Allen, and O. M. Griffith. 1967. Ultracentrifugation 
Using Acrylamide Gel. Anal. Biochem. 21:454-461. 

3. Prins, H. K., and D. D. Smink. 1965. Zone-Centrifugation. Bibl. Haematol. 23:1186. 
(Proc. 10th Congr. Intern. Soc. Blood Transfusion, Stockholm, 1964.) 

4. Cole, Thomas A., and Thomas W. Brooks, Jr., 1968. Density Gradient Centrifu- 
gation: Fixation of Bands by Photopolymerization of Acrylamide. Science: 161:386. 

5. Boyd, James B., and H. K. Mitchell. 1965. Identification of Deoxyribonucleases in 
Polyacrylamide Gel Following Their Separation by Disc Electrophoresis. Anal. 
Biochem. 13:28-42. 

6. Kurniok, N. B. 1950. The Quantitative Estimation of Desoxyribosenucleic Acid 
Based on Methyl Green Staining. Exp. Cell Res. 1:151-158. 

7. Jensen, W. A. 1962. Botanical Histochemistry. W. H. Freeman and Company, San 
Francisco. 251 p. 

8. Maurer, H. R. 1966. Einfacher Entfarbeappararat fur die Disk-Electrophorese. Z. 
Klin. Chem. 4:85-86. 

9. Petrakis, Peter L. 1969. An Inexpensive High-Resolution Densitometer for Disc 
Electrophoresis. Anal. Biochem. 28:416-427. 



The Effect of Avidin on the Biosynthesis of Fatty Acids in 

Aspergillus niger and Aspergillus flavus 

K. Schwenk and A. S. Bennett, Ball State University 

Abstract 

Submerged cultures of Aspergillus flavus and Aspergillus niger were grown in a 
medium containing avidin, a substance which inhibits the conversion of acetate to 
malonate. Control cultures were grown without avidin. 

Mycelium samples were taken at time points over a 70-hour incubation period, 
harvested by centrifugation, washed with distilled water and re-suspended in absolute 
methanol. After saponification, the fatty acids were extracted with hexane, methylated, 
isolated by thin layer chromatography, and separated and identified by gas liquid 
chromatography. 

An increase in Cm fatty acids and a decrease in Cis fatty acids by cultures grown 
with avidin suggest that malonate plays an important role in the elongation of long- 
chain fatty acids in these organisms during the time interval from 5 to 15 hours. 

In control cultures, the initially high percentage of stearic acid decreased while the 
percentage of oleic acid and linoleic acid increased, further indicating the conversion of 
stearate to oleate. 

Several monoenoic acids are present in these organisms, but the only dienoic acid 
found is linoleic acid. This suggests that the conversion of oleate to linoleate involves a 
highly specific desaturase. 

The first step in the biosynthesis of long chain fatty acids by way 
of the malonate pathway involves the enzyme, acetyl-CoA-carboxylase. 
This enzyme is one of the biotin-containing carboxylases and catalyzes 
the overall reaction. 

Acetyl-CoA + HCCV + ATP ?± Malonyl-CoA + ADP + P, 
This reaction is followed by the successive addition of 2-carbon units in 
the form of malonyl-CoA or malonyl-ACP to acetyl-CoA (4). The end 
product of this sequence of reactions is dependent upon the type of 
organism and is either palmitate (9) or stearate (3). 

Although the malonate pathway is thought to be the predominant 
pathway for the biosynthesis of fatty acids, evidence for the existence 
of alternate pathways has been presented (4). Mattoo et al. (5) reported 
that the biosynthesis of fatty acids in intact mycelium of Aspergillus 
flavus was only partially inhibited after increasing amounts of avidin, 
a biotin inhibitor, had been added to the medium. Their results sug- 
gested that the formation of malonate was not essential for the synthesis 
of fatty acids in this organism. The fatty acid distribution of the prod- 
ucts recovered from the inhibited cultures was not determined. 

In the present experiments avidin was used to study the effect of 
the inhibition of malonate formation on the types of fatty acids synthe- 
sized by A. niger and A. flavus. 

Materials and Methods 

Aspergillus niger and Aspergillus flavus, obtained from Dr. K. B. 
Raper, University of Wisconsin, were maintained on slants of Czapek 

351 



352 Indiana Academy of Science 

medium and stored at 4° C. Spores were lightly scraped and washed 
from the slants with sterile medium. Five ml of spore suspension 
(A = 0.1 at 525 m^) was added to 25 ml of sterile (autoclaved; 15 psi, 
15 min) culture medium (glucose, 40 g; NfLNO.,, 1 g; MgS0 4 «7H 2 0, 
0.3 g; KH,P0 4 , 0.3 g; H,0, 1 liter) in 250 ml Erlenmeyer flasks. The 
inoculated cultures were incubated for 24 hours at 28 °C on a recipro- 
cating shaker. At the end of this period, stir cultures were prepared by 
adding the contents of the flask to 420 ml of culture medium in a 
2-liter Erlenmeyer flask, aerating (500 ml/min) and stirring for 60 
hours. Forty-five units of avidin (1 U=amount capable of inactivating 
1 A biotin) had been added to the experimental culture medium; no 
avidin was added to control cultures. 

Aliquots of the mycelial suspension were removed from the stir 
culture at various time points. After centrifugation, the mycelial mat 
was washed with distilled water, placed in 15 ml absolute methanol, and 
sonified for 3 min (J-17A Sonifier, Branson Sonic Co. Power). The 
sonified material was diluted with an equal volume of 15 % KOH in 
methanol, refluxed for 2 hours, acidified with concentrated HC1, and 
extracted with hexane. After washing the extract with distilled water, 
drying over anhydrous sodium sulfate and removing the solvent on a 
rotary evaporator, the resulting fatty acids were methylated with 
diazomethane (7). The methyl esters were isolated on silica gel G thin 
layer plates and separated and identified by gas liquid chromatography 
using a 10-foot glass column packed with 5% poly(diethylene-glycol 
adipate) on 60/80 mesh Chrom GA/W; column temperature, 190° C; 
gas flow, 70 ml/min; Varian Aerograph 90-P instrument. For further 
identification of unsaturated fatty acids, methyl esters were separated 
by thin layer chromatography on 10% AgN0 3 -impregnated silica gel G 
with chloroform :acetone (99.5:0.5) as the developing solvent prior to 
GLC analysis. 

Results and Discussion 

Members of the class Ascomycetes have a fatty acid composition 
similar to that of plant seeds (2) and like the growing leaves and seeds 
of plants produce the stearate to linolenate series of unsaturated fatty 
acids. Shaw (8) reported that the only Cis fatty acids produced by the 
ascomycetes are the A9-unsaturated fatty acids. Analysis of the products 
in the present experiments revealed that linoleic acid was the only 
dienoic acid synthesized by A. niger and A. fla/vus, although the 
monoenoic form of C 9 through Cis acids were identified, suggesting that 
the desaturase which is responsible for the conversion of oleate to 
linoleate is highly specific. This finding makes members of the genus 
Aspergillus particularly well-suited as experimental organisms for 
studying the biosynthesis of unsaturated fatty acids in plant-like 
organisms. 

Cultures of Aspergillus -niger grown in a medium containing 
inhibitory amounts of avidin did not differ from those grown in the 
absence of avidin as to the relative rate of palmitate synthesis (Fig. 1). 



Microbiology and Molecular Biology 



353 



25 h 



20 



1 5 



1 




1 



20 30 40 50 

Age of Culture (hours) 



6 



Figure 1. Percent distribution of palmitic and stearic acids in mycelium of Aspergillus 
niger grown in media with and without avidin. 



a) r) 

PS -H 



■P -P 

G -P 



50 |- 



4 



A/ith Avidin 



18:2 




2 3 4 5 

Age of Culture (hours) 

Figure 2. Percent distribution of fatty acids in mycelium of Aspergillus niger grown in 
media with and without avidin. 



354 



Indiana Academy of Science 



However, the relative amount of stearic acid produced was substantially 
lower in the mycelium grown in the avidin containing medium. In the 
control, the relative amount of stearic acid recovered increased from 
12% in the inoculum to 22% after a 5-hour incubation period; with 
avidin, stearic acid decreased from 12% to 7%. This difference in 
stearate concentration is evident during the first 30 hours of the incu- 
bation period. 

At the 5-hour time point only 23% of the fatty acids recovered 
from the mycelium grown in the avidin-containing medium was of the 
Cis series, compared to 33% in the control (Fig. 2). 

These data suggest that malonate plays an important role in the 
elongation of palmitate to stearate. Although Mattoo (5) reported a 
partial overall inhibition of fatty acid synthesis upon the addition of 
avidin to the medium, our data show that all steps in the biosynthetic 
pathway were not affected to the same degree. In both A. niger and 
A. flavus, the addition of avidin to the culture medium inhibited the 
formation of long chain fatty acids to a greater degree than short 
chain. The importance of malonate in the elongation of palmitate to 
stearate in Aspergillus is in agreement with the report of Nagai and 
Bloch (6) who found that elongation to stearate in bacterial and plant 
extracts was dependent upon the presence of malonate. Barron (1) 
found that malonate was used for the elongation of preformed fatty 
acids by the soluble fraction of rat liver homogenates, while acetate was 
used by the mitochondrial fraction. 



-P -U 

O tu 

U 

Q) 




1 



20 30 40 50 

Age of Culture (hours) 



Figure 3. Percent distribution of fatty acids in the mycelium of Aspergillus niger at 
various times in the incubation period. 



Microbiology and Molecular Biology 355 

Bennett and Quackenbush (2) previously reported studies on the 
biosynthesis of fatty acids in Penicillium chrysogenum which indi- 
cated that endogenous palmitate was rapidly elongated to stearate and 
desaturated by the sequential pattern of oleate to linoleate to linolenate. 
In the present experiments, relatively large amounts of palmitate and 
stearate were synthesized at early time points in the growth period 
(22% and 12%, respectively, at 5 hours), followed by increases in oleate 
and linoleate concentration at 45 hours (Fig. 3), to give additional 
support for this pattern of elongation, then desaturation. 



Literature Cited 

1. Barron, E. J. 1966. The Mitochondial Fatty Acid Synthesizing System : General 
Properties and Acetate Incorporation into Monoenoic Acids. Biochim. Biophys. Acta. 
116:425-435. 

2. Bennett, A. S., and F. W. Quackenbush. 1969. Synthesis of Unsaturated Fatty 
Acids by Penicillium chrysogenum. Arch. Biochem. and Biophys. 130 :567-572. 

3. Lynen, F. 1961. Biosynthesis of Saturated Fatty Acids. Fed Proc. 20:941-951. 

4. Majerus, W. P., and P. R. Vagelos. 1967. Fatty Acid Biosynthesis and the Role of 
the Acyl Carrier Protein. Adv. Lipid Res. 5 :2-33. 

5. Mattoo, A. K., V. V. Modi, and R. N. Patel. 1966. Biotin and Fatty Acid Biogenesis 
in Aspergillus flavus. Experimentia. 22 :436-437. 

6. Nagai, J., and K. Bloch. 1966. Elongation of Acyl Carrier Protein Derivatives by 
Bacterial and Plant Extracts. J. Biol. Chem. 242 :357-365. 

7. Schlenk, H., and J. L. Gellerman. 1960. Esterification of Fatty Acids with 
Diazomethane on a Small Scale. Anal. Chem. 32:1412-1413. 

8. Shaw, R. 1965. Occurrence of Linolenic Acid in Fungi. Biochim. Biophys. Acta. 
98 :230-236. 

9. Wakil, S. J. 1961. Mechanism of Fatty Acid Snythesis. J. Lipid Res. 2:1-24. 



PHYSICS 

Chairman: Richard L. Conklin, Hanover College 
Ralph A. Llewellyn, Rose Polytechnic Institute, was elected Chairman 

for 1970 

ABSTRACTS 

Measurement of Ionization in Nuclear Emulsion by Lacunarity Tech- 
nique. W. David Mueller, Ball State University. — Lacunarity was 
studied as a technique of measuring ionization and energy loss of a 
charged particle passing through nuclear emulsion. Lacunarity is a 
measure of track density and is defined as the linear fraction of a 
track segment that consists of gaps. Measurements were made of blob 
density and lacunarity of segments of tracks of particles emanating 
from nuclear disintegration in nuclear emulsion. Both blob density and 
grain density as determined by lacunarity were plotted versus residual 
range of track segments to compare the two methods of measuring 
ionization. 

Except for tracks of very heavy ionization and tracks of light 
ionization, grain density as determined by lacunarity is a better measure 
of ionization than blob density. Both theoretical and experimental re- 
sults indicate that over a considerable range of ionization, blob density 
does not vary greatly with the degree of ionization. Also, particles 
could be distinguished by lacunarity which could not be distinguished by 
blob density. 

Operation and Flux Determination of a Neutron Generator. Martin A. 
Burkle and Leon M. Reynolds, Ball State University. — A neutron gene- 
rator-accelerator consisting of a 150 kv Cockroft-Walton accelerator which 
makes use of the 3 H(d, n) 4 He reaction was placed in operation by 
the Department of Physics, Ball State University. Experiences as- 
sociated with the installation and startup of the equipment were de- 
scribed along with projected uses. Preliminary results for neutron flux 
determinations using 2.8 Mev neutrons produced by the 2 H(d, n)3He 
reaction were given. The usefulness of the device for experimentation 
in the advanced undergraduate or beginning graduate laboratory was 
considered. 

Molecular Structure of Thyroxine by X-Ray Crystallography. L. K. 

Steinrauf, 0. Seely, J. A. Hamilton and J. M. H. Pinkerton, Indiana 
University Medical Center. — The molecular structure in the crystal 
form of the thyroid hormone thyroxine was determined by single crystal 
x-ray diffractometry using the Supper-Pace Autodiffractometer. The 
structure was found to consist of planar layers of the hormone sepa- 
rated by layers of water molecules, all connected by a highly complicated 
network of hydrogen bonds. A possible charge transfer bond exists be- 
tween molecules of thyroxine. The hydrogen bonds and the charge 

357 



358 Indiana Academy of Science 

transfer bond provide the means for speculation on the manner in which 
thyroxine can bond to blood serum proteins. 

Electron Paramagnetic Resonance Studies on The Magneli Phases of the 
Titanium-Oxygen System. 1 John F. Houlihan and L. N. Mulay, 
Pennsylvania State University. — Several transition metal oxides exhibit 
a number of stable phases with variable stoichiometry, different from 
that predicted by simple valence considerations of the cation. These 
oxides have well-defined crystal structures and are known as Magneli 
phases. 

Magnetic susceptibility studies by Mulay and others on the phases 
of the titanium-oxygen system described by Ti n 2 n-i and recent elec- 
trical conductivity data by Bartholomew of the Materials Research 
Laboratory have revealed several interesting semi-conductor to metal 
transitions. 

In this paper, typical exploratory electron paramagnetic resonance 
spectra studied as a function of temperature are presented for the fol- 
lowing oxides: TLO5, T^Ot, TioOu, and Ti T Ow. The electron paramag- 
netic resonance data, in general, have confirmed the transitions previously 
observed by other means. 

Proposed lines of interpretation correlating electron paramagnetic 
resonance parameters (such as g values, asymmetry of line shapes, etc.) 
with the magnetic data and the observed transitions were presented. 

An Evaluation of Relativistic Thermodynamics. Darryl L. Steinert, 
Hanover College. — The problem of the accuracy of the formulations of 
relativistic thermodynamics proposed by Planck, Eckart, Ott, and Lands- 
berg was examined. Due to the lack of experimental data on relativistic 
thermodynamic systems, it is not possible to compare predictions made 
by the various formulations with experimental data. But, using the 
process of evaporation as a model, I found that it is possible to study 
the consistency between the transformation laws for temperature and 
for mechanical energy. The result obtained was that only Ott's formula- 
tion is free of contradiction. 

Ott's proposed transformation laws were further evaluated in 
terms of their compatibility with relativistic formulations of fluid dy- 
namics and statistical mechanics. Compatibility is to be expected be- 
cause thermodynamics, fluid dynamics, and statistical mechanics are 
compatible in their non-relativistic formulations. 

The lack of contradiction between Ott's formulation and the trans- 
formation law for mechanical energy, and its compatibility with formu- 
lations of relativistic fluid dynamics and statistical mechanics, gave 
support to a conclusion that Ott's formulation of relativistic thermo- 
dynamics is correct. 



1 We acknowledge Prof. W. B. White for providing samples and Mr. W. J. Danley for 
assistance. This work was initially sponsored by AEC contract AT(30-1 )-2581. 



Physics 359 

Physical Oceanography in Indiana: A Study of Horse Shoe Lake. Ralph 
A. Llewellyn, Rose Polytechnic Institute. — During the spring of 1969, 
a thorough experimental oceanographic study was conducted of a large 
artificial lake in west-central Indiana. Initiated as an educational 
exercise for a group of 67 science and engineering students, the study 
developed into an integrated recording and analysis of 7 parameters of 
the lake over a several week period. 

The results of the study included a computer-drawn contour map 
of the bottom, the discovery of an unexpected region of "dead" water, 
evidence for a sub-surface current, and the formulation of plans to moor 
continuous recording gear in the lake. This concentrated study could 
well serve as a prototype for such investigations by other colleges. 



NOTES 

Polarized "He+ Ion Source for the Indiana University Cyclotron. J. H. 

Hettmer, Indiana University. — An optically pumped polarized Heli- 
um-3 ion source, similar to that designed and constructed at Rice Uni- 
versity (1), is being built for use with the cyclotron now under construc- 
tion at Indiana University (2, 3, 4). This cyclotron will be particularly 
well-suited to this purpose due to its unique property of accepting 
low-energy positive ions at ground potential for acceleration to high 
energies. This is of interest because experimental data concerning the 
scattering of, and nuclear reactions induced by, these particles are 
extremely difficult to obtain by any other technique. 

Previous work on this type of ion source, together with adaptations 
and improvements associated with this application was described. 



Literature Cited 

1. Findley, D. O., S. D. Baker, E. B. Carter, and N. D. Stockwell. 1969. A Polarized 
3 He+ Ion Source. Nucl. Instr. and Method. 71 :125-132. 

2. Sampson, M. B., M. E. Rickey, B. M. Bardin, and D. W. Miller. 1967. Planned 200- 
MeV Indiana University Cyclotron: Properties and Unique Features. Proc. Indiana 
Acad. Sci. 77:347-348. 

3. Rickey, M. E., M. B. Sampson, B. M. Bardin, and D. W. Miller. 1966. Proposed 
Indiana University 200-MeV Multi-Particle Variable Energy Cyclotron Facility. 
I.E.E.E. Trans. Nucl. Sci. NS-13 :464-468. 

4. Rickey, M. E., M. B. Sampson, and B. M. Bardin. 1969. General Design Features of 
the Indiana University 200-MeV Cyclotron. I.E.E.E. Trans. Nucl. Sci. NS-16 :397-414. 



360 Indiana Academy of Science 

Non-local Contributions to the Cyclotron Absorption Spectrum for a 
Single Valley Semiconductor Model. Uwe J. Hansen, Indiana State 
University, and James L. Hazelton, Oklahoma State University. — In 
a material like PbTe for which at 35 Ghz the skin depth is of the same 
order of magnitude as the radius of carrier orbits at cyclotron reso- 
nance, semiclassical calculations of the cyclotron absorption coefficient 
indicate that absorption maxima should be observed for dielectric anom- 
aly, rather than the expected cyclotron resonance conditions (1). 
Experiments indicate, nevertheless, that some structure is observed at 
the cyclotron frequency (1). A non-local calculation, adapted from 
Hebel's calculations for Bismuth (2), for a single ellipsoidal Fermi 
Surface with PbTe parameters was carried out for the limited case of 
the magnetic field orientation along the major symmetry axis and the 
microwave electric field parallel to the magnetic field. This calculation 
indicated an absorption maximum at the cyclotron frequency (3). More 
extensive calculations are in progress. 



Literature Cited 

1. Hansen, U. J., J. H. Gardner, and K. F. Cuff. 1966. Analysis of cyclotron absorption 
in PbTe. Bull. Amer. Phys. Soc. 11(5) :755. 

2. Hebel, L. C. 1965. Cyclotron resonance in Bi with slightly anomalous skin effect. 
Phys. Rev. 138(6A) :1641-1649. 

3. Hazelton, J. L. 1969. Semiclassical calculation of cyclotron absorption in lead 
telluride. Unpublished M.S. Thesis, Indiana State Univ., Terre Haute. 



Semiconductors Produced by Doping Oxide-glasses 
With Ir, Pd, Rh or Ru 

C. C. Sartain, Indiana State University- 



Abstract 

Semiconductors were produced by diffusion doping oxide glasses with more than 
1 wt % of Ir, Pd, Rh or Ru and by implanting 40 kilovolt Ir ions into several oxide 
glasses. Hall mobility at 300° K and 78° K was less than 0.005 cm 2 /volt sec. Mobility 
from impurity concentration and conductivity was 0.001 cm 2 /volt sec. Conductivity was 
not ionic. Enough direct current was passed through one sample to have plated out a 
million times as many Ir ions as were in the sample with no change in conductivity. 
X-ray small angle scattering indicates that conductivity was not due to electron hopping 
between conducting islands. Conductivity was ohmic at 300° K and was field dependent 
below 4° K. Thermoelectric power of 12 to 25 microvolts per degree relative to copper 
indicated hole conduction. The material absorbed throughout the visible region. For con- 
stant firing temperature and time at a constant oxygen pressure conductivity increased 
as impurity concentration increased and varied from 10- 3 to 10 J reciprocal ohm-cm. 
For a constant concentration of platinum-metal ions and for equilibrium firing, the 
conductivity was directly proportional to the oxygen pressure. Firing in hydrogen to 
remove oxygen reduced conductivity at the rate of two carriers per oxygen atom 
removed. These data indicate that the frozen-in valence states of the platinum-metal ions 
serve as the source of the carriers. 



Introduction 

Oxide glasses were caused to become amorphous semiconductors 
by heavily doping them with iridium, ruthenium, rhodium or palladium. 
The conductivity is between 10 ~' 5 and 10 2 ohms -1 cnr 1 . 

Most metals will oxidize, dissolve into and become a part of molten 
oxide glass. They are so chemically active with oxygen that all of their 
electrons form binding orbitals. They are unable to retain charges which 
can act as donors or acceptors. Thus, no one has reported doping oxide 
glasses with ordinary dopants such as boron, phosphorous, etc. 

Certain members of the platinum metal family, while not inert, are 
relatively inactive chemically. These metals have more than one oxi- 
dation state and they can retain charges which contribute to the elec- 
trical conductivity. 



Conductivity vs. Temperature 

Sample resistance was measured as a function of temperature 
from 1.48°K to 1150°K. For several hundred samples, the d-c resistance 
was measured by the volt-amp method for about 20 different tempera- 
tures between the triple point of nitrogen and 200 °C. 

Typical data are shown in Figure 1. There exists a striking simi- 
larity between the shapes of these graphs and those shown in Pearson 
and Bardeen's Figure 4A, "Resistivity of Silicon-Boron Alloys as a 
Function of Inverse Absolute Temperature'' (7). 

361 



362 



Indiana Academy of Science 



10 Meg 




Figure 1. Logarithm of resistance vs. reciprocal absolute temperature for various con- 
centrations of iridium in an oxide glass. These curves are typical examples of platinum 
metals doped into any one of many kinds of oxide glasses. 



At high temperatures the intrinsic resistance of one sample of 
oxide glass was shown. For low doping and low temperature the resist- 
ance is given by R = A exp (W/kt). For high doping the carriers seem 
to be degenerate. At low temperature as the doping increases, the slopes 
of the curves decrease monotonically as far into the "degenerate" re- 
gion as it was carried. 

It is obvious from the shape of these curves that at low doping, the 
conduction cannot be metallic. 

The conduction cannot be ionic because there was no transport of 
ions. Ionic conduction in glass usually leads to a depletion of ions from 



Physics 



363 



some volume and a change in resistance as current is passed through 
the unit. Tests of many hours duration on many samples showed that 
the resistance did not change as much as one part in 10 r> after enough 
charge had passed through the samples to have plated out a million 
times as many iridium atoms as were in the sample. 

There was no electrolysis of metals nor of oxygen. In some cases 
the only element common to different samples was oxygen. For example 
a lead-borate-oxide glass doped with iridium conducted, yet so did an 
alkali-silicate-oxide glass doped with ruthenium. In every case at least 
half of the atoms in the system were oxygen. In no case was there any 
evolution of oxygen at the terminals. 

The resistance at 300 °K was ohmic over more than 6 orders of 
magnitude between 100 /*v per cm and 500 v per cm. Conductivity did 
depend on the field below 4°K. The data here are too meager to estab- 
lish the relationship between conductivity and electric field with accu- 
racy. 



5.080 



5.070 



5.060 



5.050 



5.040 



5.030 



5.020 




1000/T 

Figure 2. Logarithm of resistance vs. reciprocal absolute temperature for sample 
I - 1 - C - D of Figure 1. Note how the experimental points fit a smooth curve even at 
greatly expanded vertical scale. 



364 



Indiana Academy of Science 



The conduction was not due to electron hopping between conduct- 
ing islands with dimensions of the order of 100 A as described by Neu- 
gebauer and Webb (6). Many X-ray small angle scattering experiments 
were performed in search of this effect. In some cases particles of this 
size were found, but there was no correlation between particle size 
and resistivity of the sample. Based on their model, Neugebauer and 
Webb (6) predicted the magnitude of the resistivity and its temperature 



Meg. 



1 Meg. 



100K 









1.80 x 


10- 3 ev 






a 


0- 3 ev. 













.3 

1 / T 



Figure 3. Logarithm of resistance vs. reciprocal absolute temperature for a sample with 
a room temperature resistance of 100 kilo ohms. The slope of this curve in the liquid 
nitrogen is 2.73 X lO- 3 ev. The slope in the liquid helium range appears to be 
1.80 XJ0- 3 ev. The point at 148° K is below the extrapolated straight line. This prob- 
ably indicates that the curve is not truly straight anywhere, but drops off continuously 
as predicted by Mott (5). See text. 



Physics 365 

dependence. The resistivity in this study was several orders of magni- 
tude too low to have been due to this effect and the temperature depend- 
ence differed from that predicted. 

Ir 2 and Ru 2 single crystals are metallic conductors (8), but they 
play no part in the conduction in these Cermets. 

The conduction was due to the presence of metals belonging to the 
platinum family. The glasses did not conduct (that is, the conductivity 
was many orders of magnitude less) unless these metals were present. 

There were no compensating metals present. 

Figure 2 is the curve which appeared flat in Figure 1. The points 
fit a smooth curve and at low temperature the line is straight with a 
slope of 1.90 x 10" 8 ev. 

Figure 3 shows how one sample behaves at liquid helium tempera- 
tures. The resistance of this sample was 10 5 ohms at 300 °K. In the 
liquid nitrogen range the slope was 2.73 milli-electron volts. The curve 
as drawn shows a slope of 1.80 milli-electron volts below 4°K. 

However, the point at 1.48 °K is definitely below the curve. 

This curve exhibits the fall-off in slope of the In p vs 1/T at low 
temperature as predicted by Mott (5). Probably at no point is the 
curve truly a straight line. It appears to fit a straight line over limited 
temperature ranges, but its slope probably decreases continuously. This 
change in slope may occur for all of our samples. 



Hall Effect 

In no sample could a Hall voltage as large as 10' 9 volts be detected 
at any temperature. For some samples, 10~ 9 volts would represent a 
mobility of about 3 x 10" 8 cm 2 per volt sec. The carrier mobility in all of 
our samples was less than this amount. 



Seebeck Effect 

Thermoelectric emf was measured in two ways. In the first method, 
one end of a long, slender sample was fixed in an ice bath while the 
other end was heated to various temperatures. In a variation of this 
method, the cold end was kept in a liquid nitrogen bath. 

In the second method, pt-pt 10 rh thermocouples were welded to 
each end of a Cermet sample about 5 cm long. The sample was inserted 
to a series of positions of increasing temperature in a tube furnace so 
that the ends of the sample were at different temperatures. The tem- 
perature of each end could be measured. The Seebeck voltage could be 
measured between the two platinum wires. 

In every case, the Seebeck Voltage indicated hole conduction. 



366 



Indiana Academy of Science 



Conductivity vs. Metal Concentration 

The conductivity as a function of concentration of ruthenium in one 
kind of glass fired at one temperature for a fixed time in air is shown 
in Figure 4. Each point on the curve represents five samples. The bars 
cover the maximum and the minimum resistance of the group. The 



300 - 



LU 

o 

< 
co 

CO 
UJ 



ID 

o 



100 - 




2 5 10 

Wt % Ru in Glass " A 



FIGURE 4. Logarithm of conductivity at :i00° K vs. logarithm of weight percent 
ruthenium in glass A. The Cermets were fired in air through a tunnel kiln so that each 
sample was exposed to the same temperature-time profile. For these samples 
a = CW*. 44 up to about 6 wt% where devitrification occurs. 



Physics 



367 



curve is fitted by a power law up to the point where devitrification 
begins. The data will not fit an exponential dependence of conductivity 
on concentration. 

For Glass "A" containing lead, silicon, Cadmium and aluminum 
oxides, devitrification occurs at about 6 wt % Ru. Devitrified samples 
look like black emery paper. 




Log CONCENTRATION GLASS B 



100 



Figure 5. Logarithm of conductivity at 300° K vs. logarithm of concentration of 
ruthenium in glass B. The samples were fired in air for constant times, but for 
different temperatures. The exponents of the different curves vary from 2.6 to IS. 3. 



368 Indiana Academy of Science 

For a different glass, labeled B, the effect of varying the firing 
temperature is shown in Figure 5. Firing time was a constant. Devitri- 
fication occurred at higher Ru concentration for this glass. 

Resistance vs Firing Time in Air 

If the number of metal atoms, the external oxygen concentration 
and the firing temperature were held constant and a number of similar 
samples are fired for different times, the resistance appeared to decrease 
exponentially with time to approach a constant value. 



Oxygen Concentration 

Figure 6 shows the conductivity of a series of similar samples fired 
for the same time and temperature at different pressures of oxygen. The 
arbitrary line with a slope of one drawn on the graph indicates that the 



100 




0.3 



1.0 

OXYGEN 



3.0 
PRESSURE , 



10.0 
INCHES Hg 



300 



Figure 6. Logarithm of conductivity at 300° K vs. logarithm of oxygen pressure. The 
samples were fired under a controlled atmosphere of commercial oxygen for a constant 
temperature and time. The arbitrary straight line has a sloj)e of 1. Each bar on the 
graph covers the measured range of 10 samples which were not precisely uniform in 
thickness. Later techniques improved the reproducibility of the sample dimension, but 
this experiment was not repeated. 



Physics 



M) 



conductivity varied directly as the number of oxygen atoms diffusing 
into the sample and reacting with the ruthenium. 

Firing in Hydrogen 

An especially large sample was made for thermogravimetric analy- 
sis. A Cermet mix containing 3.60 wt % Ru was screened and fired on 
a substrate. The measured mass of the sample was 0.303 g and its re- 
sistance at 300 °K was 23 kilo ohms. The sample was fired in hydrogen 
for 30 minutes at 500 °C. 

The sample lost a mass of 760 jxg and its resistance at 300 °C 
increased to 704 kilo ohms. 

A previous test showed that the hydrogen firing at 500° C did not 
affect the glass. 

At 500 °C, apparently hydrogen converted Ru 2 to Ru-O:.. The water 
formed evaporated from the sample. Each Ru O2 presumably contributes 
one hole for conduction. Conversion of each Ru (X to V2 (Ru20 ;f ) or 
removing one-half of an oxygen atom from each Ru atom removes one 



Conductance 

Micromhos 

47.8 



Measured 43.5 



Oxygen Wt. 



No. of 
Holes at 



Measured 1.4 



r u 07 Micrograms 300 K 
864 — 6.47xl0 19 




5.91x10 



19 



5.72xl0 19 
Holes 



0.19x10 



19 



Ru 2 3 



— 



Figure 7. Schematic diagram showing how removal of oxygen from one sample of 
Cermet removes holes and correspondingly reduces conductance. 



370 Indiana Academy of Science 

hole thus removing one charged carrier. The Ru in the sample was 3.6% 
of 0.303 gm or 10.9 mg, or 6.47 X 10 19 atoms. The loss of 760 fig of 
oxygen represented a loss of 2.86 X 10 19 atoms of oxygen, or 5.72 X 10 19 
holes (two holes per oxygen atom removed). The conductance became 
1/704 kilo ohms or 1.4 micromhos. The loss in conductance was 43.5 — 
1.4 or 42.1 micromhos. 

In Figure 7, the quantity of oxygen required to convert Ru 2 3 to 
Ru 2 is plotted schematically on the vertical scale. For this sample, 
which contained 10.9 mg of Ru, 864 fig of oxygen was required for the 
conversion. 

On the left, conductance of the sample in micromhos was plotted, 
and on the far right the number of holes in the sample at 300 °K. 

When the sample was originally fired in air, the ruthenium was 
oxidized to the level such that its measured conductance was 43.5 mi- 
cromhos. The sample was then fired in hydrogen. This firing removed 
a measured quantity of 760 fig of oxygen or 5.72 X 10 19 holes and re- 
duced the conductance to the measured value of 1.4 micromhos. The 
assumption that removing 5.72 X 10 19 holes removed 42.5 micromhos 
of conductance gave the scale of the graph. For this sample one mi- 
cromho was equivalent to 0.1345 X 10 lfi holes, or 10'" holes was equiva- 
lent to 7.44 micromhos. Now the measured level of 1.4 micromhos above 
zero was set to be 0.19 X 10 10 holes. 

Firing this sample in air at the temperature and time did not 
oxidize all of the Ru to Ru Oz. Only 5.91 to 10 19 atoms of Ru out of a 
possible 6.47 X 10 19 or 91.3% were combined as Ru O2. The remainder 
was oxidized only to the Ru 2 On level. 

For this sample the carrier concentration was 11.2 X 10 20 Ru atoms 
per cm 3 times 91.3%- or 10.2 X 10 30 holes per cnr. <r = 0.204 fi J cm 1 . The 
mobility fi = a/ne — 1.25 X 10~ 3 cmVvolt sec. 



Other Physical Properties 

No magneto-resistance was observed up to 14 kilogauss. No photo- 
conductivity was observed at liquid nitrogen temperatures or above. 
A very small increase in conductivity was noted when light from an 
incandescent bulb (microscope illuminator) was focused on a sample 
bathed in liquid helium. This was attributed to the heating of the sample, 
not to photoconductivity of the sample. The samples absorbed energy 
throughout the visible region. 

The kind of glass does affect the conductivity, suggesting that the 
conductivity might be proportional to the square of the dielectric con- 
stant. However, in these experiments, the effect of the dielectric con- 
stant was not isolated from other effects, particularly that due to the 
temperature of firing, i.e., that of quenching the oxide into a particular 
frozen-in valance state. Obviously this latter effect depends on the 
softening point of the glass. 



Physics 371 

No effects such as rectification were observed which could be inter- 
preted as due to the contacts or terminations. 

Noise was proportional to 1 /frequency for these samples at room 
temperature. The noise was not measured at any other temperature. 

An unsuccessful attempt was made to measure mobility directly (2, 
3) by injecting carriers and measuring the time to travel a microscopic 
distance. This experiment indicated the lifetime was very short. If it is 
assumed that /* r= € r/m* with p, = 10 -3 and m* = m, T is indeed very 
small. Smaller values of m* require even smaller values of T . 

Conductivity in alternating current below 4°K is independent of 
frequency up to 10 kc/sec. 

It is interesting to compare Sb-doped Ge (1) with Ir-doped glass. 

I noted that at the doping level just high enough to make Fritzsche's 
Sb-doped Ge become degenerate, N = 2.7 X 10 17 atoms per cm 3 , p 300 = 
0.015 ohm cm and fi = 1500 cnrVvolt sec. At the doping level required to 
make Ir in glass "degenerate," N = 1.2 X 10"', p = 2.1 and ix — 1.5 X 
lO -3 . The ratio of N is about 5000, that of p is about 200 and that of y, 
is about 10- 6 . 

Ion Implantation 1 

Forty kilovolt ions of Ir-193 were implanted into fused Si0 2 and five 
other glasses. The doping levels were 10 14 , 10 ]5 and 10* 16 ions per cm a . 
The Lindhard depth of penetration and the deviation of the range for 
40 kv Ir-193 into SiO- were computed by Gibbons and Johnson (4) to be 
222a with a spread of ~ 22a. The electrical properties of the ion im- 
planted material is about the same as that of the diffused material. If it 
is assumed that half of the Ir-193 ions do stop in a layer only 45a 
thick, if the quantum mechanical effect of such a thin conductor is 
neglected, and if the conductivity based on the behavior of the same 
number of ions diffused into the glass is estimated, the measured con- 
ductivity is obtained. This is good evidence that the holes do not reflect 
or scatter from the surface of the conductor and therefore they hop 
short distances in short lifetimes, Also, the conductivity of the various 
glasses were crudely proportional to the square of the dielectric constant. 

It appears that when the 40 kv Ir ions stop in oxide glass, local 
heating and other effects permit the ions to form chemical bonds with 
neighboring oxygen atoms and that Ir introduced by ion bombardment 
forms the same bonds as Ir diffused into the glass at high temperature. 

Discussion 

These experiments indicated that each tetravalent atom of one of 
the platinum metals in an oxide glass can contribute a hole which 
participates in the conduction mechanism. The trivalent atoms seemed 



1 We are indebted to Mr. G. Alton and the Stable Isotopes Division of the Oak Ridge 
National Laboratory for implanting the ions. 



372 Indiana Academy of Science 

to take no part in the conduction. The valence state was fixed at high 
temperature by adding or removing oxygen from the platinum metal 
ion. When the sample was cooled this valence state was frozen-in. Since 
the rate of cooling was rapid, this may be considered a quenching 
effect. The number of platinum metal ions frozen in the tetravalent state 
determined the number of holes available to carry the current, thus fix- 
ing the resistance (or conductance) of the sample. The number of car- 
riers was easily changed by changing the number of atoms in the 
tetravalent state. This could be done only at high temperatures. 

The resistance could be changed by changing the total number of 
platinum metal atoms in a sample. However, in a sample already manu- 
factured which contains a fixed number of platinum metal atoms, the 
number of tetravalent atoms could be changed by increasing or de- 
creasing the oxygen content of the sample. This could be done by firing 
for a different time at a given oxygen pressure and temperature, by 
firing in an oxidizing or reducing atmosphere or even by firing at a 
different temperature. 

At high temperatures the oxides of the platinum metals were vola- 
tile, so only a limited temperature range was available for work. 

Under certain conditions semi-conducting glass could be made 
from osmium. However, osmium oxide was very volatile and poisonous, 
and it is difficult to process the glass without losing osmium. 

In these experiments the conductivity apparently depended on 
the state of one of the electrons belonging to the d-shell of the platinum- 
metal atom. If this electron was bound to an oxygen atom, there existed 
a hole in the d-shell (band?) which may act as a trap or otherwise par- 
ticipate in conduction. If there was no oxygen atom nearby to attract 
the electron, it occupied its ground state in the d-shell and no hole 
was available. 

The conductivity was a discontinuous function of concentration. If 
the concentration was less than about 1 wt % or about 3 X 10 30 Pt-metal 
atoms per cm 3 the sample conductivity decreased by many orders of 
magnitude to that of the intrinsic conductivity of the glass. At this con- 
centration the average volume occupied by only one ion was about 3 
X 10 -* cm 3 . Thus, at the average distance between ions of about 15a, the 
samples change from the conducting to the non-conducting state. 



Physics 373 

Literature Cited 

1. Fkitzsche, H. 1958. Resistivity and Hall coefficient of antimony-doped germanium at 
low temperatures. J. Phys. and Chem. of Solids. 6 :69-80. 

2. Haynes, J. R., and W. Shockley. 1951. The mobility and life injected holes and 
electrons in germanium. Phys. Rev. 81 :835-843. 

3. Haynes, J. R., and W. C. Westphal. 1952. The drift mobility of electrons in silicon. 
Phys. Rev. 85 :680. 

4. Johnson, W. S., and J. F. Gibbons. 1966. Statistical range distribution of ions in 
single and multiple element substrates. Applied Phys. Letters 9 :321-322. 

5. Mott, N. F. 1968. Conduction in glasses containing transition metal ions. J. Non- 
Crystalline Solids. 1 :36-52. 

6. Neugebauer, C. A., and M. B. Webb. 1962. Electrical conduction mechanism in ultra- 
thin, evaporated metal films. J. Appl. Phys. 33 :74-82. 

7. Pearson, G. L., and J. Bardeen. 1949. Electrical properties of pui-e silicon and silicon 
alloys containing boron and phosphorus. Phys. Rev. 75 :865-883. 

8. Ryden, W. D., A. W. Lawson, and C. C. Sartain. 1968. Temperature dependence of 
the resistivity of RuO- and Ir02. Phys. Letter 26A :209-210. 



PLANT TAXONOMY 

Chairman: Jack E. Humbles, Indiana University 
Jeanette C. Oliver, Ball State University, was elected Chairman 

for 1970 

ABSTRACTS 

Eocene Euphorbiaceous Fruits. Neal E. Lambert and David L. Dil- 
cher, Indiana University. — A recent study of a population of 20 well- 
preserved fossil fruits has shown that these fruits have probable af- 
finities with the Euphorbiaceae. These fruits were collected from Eocene 
clay deposits in Henry County, Tennessee. In 1922, E. W. Berry de- 
scribed similar fruits collected from Eocene deposits in western Ten- 
nessee and assigned these to the genus Monocarpellites. He tentatively 
referred the genus to the Malvaceae but expressed uncertainty concern- 
ing its botanical affinities. M. E. J. Chandler described similar fruits 
from Lower Tertiary deposits of Egypt, Isle of Wight, Sussex of 
England. Chandler referred her material to two genera, Wetherellia and 
Palaeow ether ellia, and placed the genera in the Euphorbiaceae. The 
fruits from western Tennessee most resemble the genus Palaeowether- 
ellia; however, there are some consistent differences. Both the external 
features and internal anatomical structure of the fossil material indi- 
cate an affinity with some of the large capsules found in extant woody 
genera of the Euphorbiaceae. The fossil fruits are septicidal capsules, 
24-39 mm in diameter with 7-10 locules which dehisce radially exposing 
the seeds. Sections showing internal cellular detail have been prepared 
and will be discussed in addition to the overall aspects of the fruits. 

Morphology and Taxonomy of Fossil Fungal Spores. M. V. Sheffy and 
D. L. Dilcher, Indiana University. — The Eocene clay deposits of west- 
ern Tennessee and Kentucky contain large numbers of preserved dis- 
persed fungal spores. Extensive research has been done on the fossil 
leaves associated with these clays and one study by Dilcher (1965) re- 
ports numerous epiphyllous fungi found on these fossil leaves. A large 
assemblage of dispersed fungal spores was isolated from the sediments 
by zinc bromide flotation and mounted in glycerine jelly. Camera lucida 
line drawings and photographs were made of each spore type to ac- 
company the spore descriptions. Distinct morphological characters such 
as shape, size, sculpture, and number of cells and pores were used to 
delimit 14 genera and 81 species. They were classified according to an 
artificial system of nomenclature followed by von der Hammen, Rouse, 
Clarke and Elsik. This preliminary taxonomic work is necessary for 
any further work with fungal spores as part of a complete organic 
assemblage or as marker fossils in stratigraphic correlations. Many of 
the spores found are similar to dispersed spores recorded from Cre- 
taceous-Recent sediments of North America, Europe and Africa. Al- 
though some spores have been tentatively assigned to modern taxa by 

375 



376 Indiana Academy of Science 

Bradley, Dilcher, Wolf and Graham, this continues to be a difficult task 
until extensive modern reference collections are required. 

The Cultivated Solanaceae of Ecuador. Charles B. Heiser, Jr., Indiana 
University. — The family Solanaceae is well represented among the plants 
cultivated in Ecuador. Several members of the family are important 
food plants, including such well known ones as the potato (Solanum 
tuberosum) and peppers {Capsicum spp.) , as well as a number of lesser 
known ones, including the naranjilla (S. quitoense), the pepino (S. 
muricatum) and the tree tomato {Cyphomandra crassi folia) . Most of the 
ornamentals of the family grown in the temperate zone are also culti- 
vated there, and in addition a number of shrubs are grown for their 
ornamental value (Solanum spp., Iochroma fuchsioides, Streptosolen 
jamesonii, and Datura spp.). Various species of the genus Datura are 
also employed for use as narcotics. Chromosome counts were obtained 
for the following: Datura arborea, n=12; D. aurea n=12; D. Can- 
dida, n=12; Iochroma fuchsioides, n=ca.24; and Streptosolen jame- 
sonii, n=ll. 



Hookeriaceae Species and Distribution in Africa, Europe, Asia, 
Australia and Oceania 

Winona H. Welch, DePauw University 

Abstract 

This is the third and last in the series of papers on the distribution of the 
Hookeriaceae. The first (4) pertained to North and Central America and West Indies, 
and the second (6) to South America. 

No Antarctic or cosmopolitan species of Hookeriaceae have been noted. Endemism 
occurs frequently in the family. An observation of the world distribution of genera 
shows that nine genera are known only from one area and twelve genera occur in more 
than one area but have a hemisphere or a continental distribution, such as in the 
Americas, in the West Indies and the Americas, or in Asia. 

In the progress of the monographic studies in the Hookeriaceae and 
due to the publication of Volume 5 of Index Muscorum (7), the follow- 
ing changes and additions in synonymy, authors of epithets, and distri- 
bution have been made since publications (4) and (6)' Callicostella 
depressa var. rubella (Besch.) Wijk & Marg. = C. depressa (Hedw.) 
Jaeg. — Am 2-5; *C. filescens (Besch.) Jaeg. — Am 3; C. leonii Ther. = 
Pilotrichidium leonii (Ther.) Crum & Bartr. — Am 3; *C. merkelii 
(Hornsch.) Aongstr. — Am 5; C. strumulosa (Hampe & Lor.) Jaeg. — 
Am 4, 5; *C. subdepressa (Besch.) Kindb. — Am 5; C. wallisii Fleisch. 
r= nom. nud. — Am 2. Crossomitrium crugeri C. Mull. = C. patrisiae 
(Brid.) C. Mull. — Am 1-6; C. herminieri (Besch.) Jaeg. — Am 2, 3. 
Cyclodictyon blandum (Lor.) Kuntze = C. varians (Sull.) Kuntze — Am 
1, 3, 5; C. pattens (Mitt.) Kuntze = C. varians (Sull.) Kuntze — Am 1, 
3, 5; *C. regnellianum (C. Mull.) Broth, ex Par. — Am 5; C. varians 
(Sull.) Kuntze — Am 1, 3, 5. Distichophyllum densirete Broth. — *Les- 
keodon densiretis (Broth.) Broth. — Am 5; *D. fasciculatum Mitt. — 
Afr 4, Am 6, Oc. *Eriopus haitensis Crum & Steere — Am 3. Hemiragis 
anrea (Brid.) Kindb. — Am 2-5. Hookeria viridula Mitt. = Cyclodictyon 
viridulum (Mitt.) Kuntze — Am 4, 5; H. wallisii C. Mull. = nom. nud. 
— Am 2. Hookeriopsis diffusa (Wils.) Jaeg. — Am 2, 4; *H. fissideyitoides 
(Hook. & Wils.) Jaeg. — Am 3; *H. heteroica Card, has recently been 
collected in Mississippi, previously known from Mexico — Am 1 ; *H, 
luteo-viridis (Besch.) Kindb. — Am 5; *H. perfulva (C. Mull.) Broth. 
ex Par. — Am 5; *H. websteri Crum & Bartr. — Am 3. Hypnella diversi- 
folia (Mitt.) Jaeg. = Neohypnella diversifola (Mitt.) Welch & Crum 
— Am 3-5. Isodrepanium lentulum (Wils.) Britt. — 1-5. Lepidopilidium 
divaricatum (Doz. & Molk.) Broth. — Am 2-5. Lepidopilum calomicron 
Broth. = * Crossomitrium calomicron (Broth.) Welch — Am 3; L. dia- 
phanum (Hedw.) Mitt. — Am 2-4; L. pumilum Mitt. — Am 2, 4; L. sub- 
tortifolium Bartr. — Am 1-2. *Leskeodon parvifolius Bartr. — Am 4; *L. 
wallisii (C. Mull.) Broth, ex Par. — Am 4. Neohypnella chrysophyllopodia 



1 It is assumed that the introductory pages of the first (4) and second (6) papers will 
be reviewed before reading this treatise. 

* The asterisk before the epithet in the following lists indicates the species, varieties, 
and forms which are not known to occur in geographical areas other than the one cited. 

377 



378 Indiana Academy of Science 

(C. Mull.) Bartr. = N. diversifolia (Mitt.) Welch & Crum— Am 2, 4, 
5; N. mucronifolia Bartr. = N. diversifolia. Pseudohypnella mucroni- 
folia Bartr. ex Welch & Crum in error for Neohypnella mucronifolia. 
Thamniopsis killipii (Williams) Bartr. — Am 4, 5. 

Africa, America, Asia, Australia, Europe, and Oceania are abbre- 
viated, respectively, as Afr, Am, As, Austr, Eur, and Oc. The numbers 
following the abbreviations refer to specific areas in these countries, as 
used in Index Muscorum and by previous papers on the distribution of 
Hookeriaceae. 

An observation of the world distribution of genera shows that the 
following are known from one area or one continent only: Archboldiella 
and Leskeodontopsis, As 4; Orontobryiim, As 3; Bellia, Austr 2; Cal- 
licostellopsis, Stenodesmus, and Thamniopsis, Am 4; Lamprophyllnm, 
Am 6; and Tetrastichium, Afr 1. 

Further observations show that additional genera occur in more 
than one area and have a hemispheric or a continental distribution: 
Adelothecium, Am 1-5; Amblytropis, Am 3-4; ChaetomitHopsis, As 2-4; 
Crossomitrium, Am 1-6; Dimorphocladon, As 3-4; Helicoblepharum, Am 
4-5; Hemiragis, Am 2-5; Isodrepanium, Am 1-5; Neohypnella, Am 2, 
3, 5; Philophyllum, Am 2, 4, 5; Rhynchostegiopsis, Am 2-5; and Steno- 
dictyon, Am 2-4. 

The areas of distribution in the earlier publications (4, 6) are re- 
peated for reference. America 1: North America (Canada, United 
States, Mexico), Greenland, Aleutian Islands, Bermudas; America 2: 
Central America and Cocos Island ; America 3 : West Indian Islands 
(Greater and Lesser Antilles, Bahamas) except Trinidad and Tobago; 
America 4: Venezuela, Colombia, Peru, Bolivia, Ecuador, Galapagos 
Islands: America 5: Brazil, Paraguay, Guiana, Trinidad, Tobago; and 
America 6: Chile, Argentina, Uruguay, Falkland Islands, and Hermite 
Island. 

The following islands have been included with Africa: the Azores, 
Canary Islands, Madeira Islands, Saint Helena Island, Madagascar 
Island, Mauritius Island (formerly Isle de France), Reunion Island 
(formerly Bourbon) and Kerguelen (Desolation Island). 

If tropical Africa is considered as a distribution center, we find 
species which occur in Africa also occurring in Europe and Asia, as well 
as Australia to the east and the Americas to the west, particularly 
South America. Cyclodictyon laete-virens has been reported from 
throughout Africa and in Europe. Daltonia angustifolia occurs in 
Africa, Asia, and New Zealand, and D. strictifolia in Africa and Asia. 
D. splachnoides has been recorded in America, Africa, Europe, and 
Australia. Collections of Eriopus cristatus have been reported from 
Africa, South America, Australia, New Zealand, and Oceania. Ap- 
parently Hookeria lucens grows in cooler areas than the great majority 
of the species since it occurs in northern Africa and adjacent Asia, 
Europe, and northern North America. 



Plant Taxonomy 379 

Endemism in African Hookeriaceae is greatest in Afr 2, consist- 
ing of 67 species and varieties. Afr 3 has 35 species, and Afr 1 and 
Afr 4, two and six, respectively. The Hookeriaceae are represented in 
Afr 1 by four species; Afr 2 by 89 species, varieties, and forms; Afr 
3 with 56 species and varieties; and Afr 4 by 14. 

The European species of Hookeriaceae have extensive distribution 
records. Cyclodictyon laete-virens occurs also in Afr 1-4; Daltonia 
splachnoides in Afr 2, Am 2, Austr 1, 2; Distichophyllum carinatum in 
As 2; Eriopus apiculatus (1) (introduced and established in England), 
in Austr 1, 2, Am 6; and Hookeria lucens in Am 1, Afr 1, and As 5. 

One hundred and sixty-eight species, subspecies, and varieties have 
been reported from As 4, 57 from As 3, 27 from As 2, one from As 5, 
and one from As 1. It is evident that the center of distribution of 
Asiatic species is As 4. Ten endemic species, subspecies, and varieties 
of Hookeriaceae are known in As 2, 28 in As 3, and 126 in As 4. 

Distichophyllum carinatum occurs in Eur as well as in As 2. As 2 
has in common with Oc: Callicostella papiUata, Cyclodictyon blume- 
anum, Distichophyllum mittenii, and Hookeria acutifolia which also 
occurs in Am 1-5 and As 3-4. In addition to As 2-4, Daltonia angusti- 
folia also occurs in Afr 2-3 and Austr 2, and var. strictifolia in Afr 3. 
Twenty Asian species occur in Oc, six in Afr 3, four in Afr 2, three in 
Austr 1, and three in Austr 2. Hookeria lucens has been reported from 
As 5, Eur, Afr 1, and Am 1. H. acuti folia has been recorded in As 2-4, 
Am 1-5, and Oc. The following species in As 3 also occur in Oc : Calli- 
costella papiUata, C. prabaktiana, Cyclodictyon blumeanum, Daltonia 
contort a, Distichophyllum 'mittenii, and Hookeria acutifolia (also in 
As 2, 4, and Am 1-5). Hookeriopsis pallidifolia has been reported from 
Afr 2 as well as As 3. In addition to As 4, Callicostella kaernbachii, 
Chaetomitrium tahitense, var. deplanchei, and Cyclodictyon lepidum 
occur in Austr 1 ; Callicostella papiUata, C. prabaktiana, Chaetomitrium 
tahitense, var. deplanchei, Cyclodictyon blumeanum, var. vescoanum, Dal- 
hitense, var. deplanchei, Cyclodictyon blumeanum, var. vescoanum, Dal- 
tonia contorta, Distichophyllum mittenii, D. undulatum, Eriopus re- 
motif olia, Hookeria acutifolia (also As 2, 3, Am 1-5), and Leskeodon 
acuminatus occur in Oc; Daltonia, angustifolia in Austr 2 and Afr 2, 3, 
and its var. strictifolia in Afr 3. One species, Hookeria lucens (Hedw.) 
Sm., is the only species known from As 5. This species also grows in 
Eur, Afr 1, and Am 1, a northern distribution in comparison with the 
subtropical and tropical habitats of the great majority of species. 

Australia and Tasmania have 34 species of Hookeriaceae, 19 of 
which are endemic; and New Zealand, etc., 26 species, varieties, and 
forms, 15 of which are endemic. In the distribution of species, Australia 
and Tasmania have 9 species in common with New Zealand, and 7 in 
common with Am 6, 3 each with As 4, Afr 3, and Oc, and 1 each with 
Eur, Afr 2, and Am 2. New Zealand has 9 species which also occur in 
Australia and Tasmania, and 6 in common with Am 6, 2 each with 
Afr 2 and 3, and 1 each with As 3 and 4, Eur, Am 2, and Oc. It is ap- 



380 Indiana Academy of Science 

parent that Australia, Tasmania, and New Zealand have more species 
of Hookeriaceae in common with each other and with Am 6 than they 
have with other countries. 

Pterygophyllum balantii Broth. (Hepaticina balantii C. Mull, in 
Hedwigia 41:128. 1902) was described in his Symbolae ad Bryologicam 
Australiae III, without citation of locality as Australia, Tasmania, or 
New Zealand. 

Austr 1 has 19 endemic species in the Hookeriaceae. Austr 1 and 2 
have in common nine species. Distichophyllum rotundi folium, Eriopus 
apiculatus, Pterygophyllum dentatum, and Sauloma tenella occur in 
Austr 1, 2, and Am 6; Distichophyllum assimile and Pterygophyllum 
obscurum have been reported from Austr 1 and Am 6; Callicostella 
kaernbachii, Chaetomitrium tahitense, and var. deplanchei have habitats 
in Austr 1 and As 4. Austr 1 and Oc have in common : Chaetomitrium 
tahitense, var. deplanchei, and Eriopus cristatus. Species with a very 
extensive range of distribution are Daltonia splachnoides in Austr 1, 
2, Afr 2, Eur, Am 2, and Eriopus cristatus in Austr 1, 2, Afr 3, Am 6, 
and Oc. 

Austr 2 has seven endemic species, six varieties, and two forms in 
26 representatives of the Hookeriaceae. Nine species occur in both 
Austr 1 and 2, and six of those in Austr 2 also grow in Am 6; Disti- 
chophyllum rotundi folium, Eriopus apiculatus, E. cristatus, E. flexi- 
collis, Pterygophyllum dentatum, and Sauloma tenella. Three species of 
Austr 2 have an extensive range of distribution. Daltonia angustifolia 
has been reported from As 2-4 and Afr 2,3; D. splachnoides from Austr 

2, Afr 2, Am 2, and Eur; and Eriopus cristatus from Oc, Austr 1, Afr 

3, and Am 6. 

Endemism in the Hookeriaceae of Oceania consists of 45 species, six 
varieties, and one form. Hookeria acutifolia occurs in As 2-4 and 
Am 1-5, as well as in Oc. Eriopus cristatus has been reported from Oc, 
and also from Austr 1, 2, Afr 3, and Am 6. The distribution range of 
four species from Oceania extends into As 2: Callicostella papillata, 
Cyclodictyon blumeanum, Distichophyllum mittenii, and Hookei'ia 
acutifolia; six into As 3; Callicostella papillata, C. prabaktiana, Cyclo- 
dictyon blumeanum, Daltonia contorta, Distichophyllum mittenii, and 
Hookeria acutifolia; and 12 into As 4: Callicostella papillata, C. 
prabaktiana, Chaetomitrium tahitense, var. deplanchei, Cyclodictyon 
blumeanum, var. vescoanum, Daltonia contorta, Distichophyllum mit- 
tenii, D. undulatum, Eriopus re motif olius, Hookeria acutifolia, and 
Leskeodon acuminatus. Austr 1 and Oc have two species and a variety 
in common: Chaetomitrium tahitense, var. deplanchei, and Eriopus 
cristatus; and Austr 2 and Oc, 1: E. cristatus. 

Hookeriacae species of Oceania have a distributional relationship of 
11 species with As 4, six with As 3, four with As 2, two with Austr 2, and 
one each with Austr 2, Afr 3, and Am 1-6. 



Plant Taxonomy 381 

Africa 1: North Africa, Madeira, Azores, Canary Islands. 

Cyclodictyon laete-virens (Hook. & Tayl.) Mitt.; Hookeria lucens 
(Hedw.) Sm. ; *Lepidopilum virens Card.; '• Tetrastich ium fontanum 
(Mitt.) Card. 

Africa 2: Central Africa, Saint Helena Island, Fernando Po. 

*Actinodontium dusenii Broth.; *A. streptogoyieum Broth.; Cal- 
licostella africana Mitt.; *C. ascenionis (C. Mull.) Kindb.; *C. attenuata 
(C. Mull.) Kindb.; C. brevipes (Broth.) Broth.; ::: C. chevalieri Broth. 
in Corb.; *C chinophylla (C. Miill.) Kindb.; C. constricta (C. Mull.) 
Kindb.; *C. emarginatula Broth, in Corb.; C. eroso-truncata Card.; 
C. fissidentella (Besch.) Kindb.; *C. gabonesis Broth. & P. Varde; 
C. lacerans (C. Miill.) Jaeg. ; C. leptocladula (Broth.) Broth.; *C. 
maclaudii (Par. & Broth.) Broth.; *C. papillosula Broth. & P. Varde; 
*C. perpapillata Broth. & P. Varde; C. pusilla Broth, ex Demar. & 
P. Varde, C. salaziae (Besch.) Kindb.; C. seychellensis (Besch.) Kindb.; 
*C. sub emarginatiila Broth. & P. Varde; C. tristis (C. Miill.) Broth.; 
:!: C. usambarica (Broth.) Broth.; *Chaetomitrium dusenii C. Miill. ex 
Broth.; *C. dusenii var. brevinerve (P. Varde) P. Varde; ^Cyclodictyon 
bidentatum Demar.; C. borbonicum (Besch.) Broth.; *C. brevifolmm 
Broth, in Mildbr. ; :;: C. crassicaule Broth, in Mildbr.; *C. delicatum P. 
Varde; *C. dixonianum Demar.; *C. filicuspis P. Varde; C. hildebrandtii 
(C. Miill.) Kuntze; :|: C. immersum Broth. & P. Varde; *C krebedjense 
Broth.; *C krebedjense var. argutidens P. Varde; C. laete-virens 
(Hook, and Tayl.) Mitt:; *C. lebrunii Demar. & P. Varde; *C. per- 
limbatum Broth.; *C. preussii (Broth.) Broth.; : ' : C. recognitum De- 
mar. ■& P. Varde; :|: C. spectabile Broth, in Mildbr.; *C. subbrevi- 
folium Broth.; :|: C. subobtusifolium Dix. ex Demar. & P. Varde; C. 
vallis-gratiae (C. Miill.) Kuntze; C. vallis-gratiae f. breutelianivm 
(Hampe) Demar. & P. Varde; C. vesiculosum (Brid.) Kuntze; 
Daltonia angustifolia Doz. & Molk.; : D. constricta P. Varde; 
*D. dusenii C. Miill. ex Broth.; *Z). euryloma P. Varde; *D. 
longinervis Mitt.; *Z). mildbraedii Broth, in Mildbr.; *D. mildbraedii 
var. laevis Demar. & Leroy; *D. minor Besch.; *Z). minuta Ther. ; *D. 
plicata P. Varde; D. splachnoides (Sm.) Hook. & Tayl.; *D. tortifolia 
Demar. & P. Varde; *Distichophyllidium africanum Demar. & P. Varde; 
*Distichophyllum procumbens Mitt.; *D. rigidicaule (Dus.) Broth.; *D. 
rigidicaule var. gabonense (P. Varde) P. Varde; * Hookeria contracta 
Gepp in Hiern.; *Hookeriopsis ambigna P. Varde; *H. angolensis 
(Welw. & Dub.) Broth, ex Par.; *H. cheiloneura (Broth.) Broth.; *H. 
gabonensis Broth. & P. Varde; *H. mittenii P. Varde; H. pallidifolia 
(Mitt.) Geh. & Herz.; *H. papillosula Broth. & P. Varde; H. pappeana 
(Hampe) Jaeg. ; *H. staudtii (Broth.) Broth.; *Hypnella abrupta 
(Mitt.) Jaeg., *H. guineensis Broth. & Par., ■'Lepidopilidium cyrtoste- 
gium (Ren. & Card.) Card, in Grand.; *L. devexum (Mitt.) Broth.; 
*L. hanningtonii (Mitt.) Broth.; *L. subdevexum (Broth.) Broth.; 
*L. subdevexum var. intermedium P. Varde; *L. theriotii Nav. in Dix. 
& Ther.; *Lepidopilum callochlorum (C. Miill.) ex Broth.; L. dusenii 



382 Indiana Academy of Science 

C. Mull, ex Broth.; *L. hirsutum (Besch.) Broth, var. tuberculatum 
Ther.; *L. lastii Mitt.; L. niveum (C. Mull.) Kindb.; *Sauloma afri- 
cana Dix. & Ther.; *S. tisserantii P. Varde. 

Africa 3: Madagascar, Mauritius, Reunion Islands. 

* Actinodontium hirsutum Besch. var. ramosum Besch.; Callicostella 
af vicuna Mitt.; C. brevipes (Broth.) Broth.; C. constricta (C. Mull.) 
Kindb.; C. erosotruncata Card.; C. fissidentella (Besch.) Kindb.; C. 
lacerans (C. Mull.) Jaeg. ; *C. laeviuscula Mitt.; C. leptocladula 
(Broth.) Broth.; C. papillata (Mont.) Mitt.; C. papillata var. brevifolia 
Fleisch.; *C. parvocellulata Demar. & P. Varde; *C perrotii (Par.) 
Broth.; C. pusilla Broth, ex Demar. & P. Varde; C. salaziae (Besch.) 
Kindb.; C. seychellensis (Besch.) Kindb.; * Chaetomitrium borbonicum 
Besch.; *C. comorense Hampe ex C. Mull.; *Cyclodictyon aubertii (P. 
Beauv.) Kuntze; C. borbonicum (Besch.) Broth.; C. hildebrandtii (C. 
Mull.) Kuntze; C. laete-virens (Hook. & Tayl.) Mitt.; *C. perrottetii 
Demar. & P. Varde; C. vallis-gratiae (C. Mull.) Kuntze; C. vallis-gratiae 
f. breutelianum (Hampe) Demar. & P. Varde; C. vesiculosum (Brid.) 
Kuntze; Daltonia angustifolia Doz. & Molk.; D. angustifolia var. 
strictifolia (Mitt.) Fleisch.; *D. forsythii Broth, ex Card, in Grand.; 
:|: Z). latimar ginata Besch.; *D. latimarginata var. madagassa Ren.; 
*Z>. stenoloma Besch.; *Distichophyllum mascarenicum Besch.; *Eriopus 
asplenioides (Brid.) Besch.; E. cristatus (Hedw.) Brid. in C. Mull. ; 
*E. fragilis C. Mull.; *Hookeria splachnif olia (Brid.) Arnott; *Hooker- 
iopsis darntyi (Besch.) Broth, in Card, in Grand.; *H. diversifolia 
(Ren. & Card.) Broth, ex Card, in Grand.; *Hypnella semi-scabra Ren. 
& Card.; *H. viridis Ren. & Card.; '■Lepidopilidium attenuatum Ther.; 
*L. branneoleum (C. Mull.) Broth.; L. cespitosum (Besch.) Broth.; 
L. chenagonii (Ren. & Card.) Card, in Grand.; *L. corbieri (Ren. & 
Card.) Card, in Grand.; *L. flexuosum (Besch.) Broth, ex Par., *L. is- 
leanum (Besch.) Broth.; :|: L. parvulum Card, in Grand.; *L. subrevolutum 
(Ren. <& Card.) Card, in Grand.; *Lepidopilum carrougeauanum Ther. 
& P. Varde; L. hirsutum (Besch.) Broth.; 'L. hirsutum var. ramosum 
Besch.; :|: L. hirsutum var. tuberculatum Ther.; :;: L. humblotii Ren. & 
Card.; *L. verrucipes Card, in Grand. 

Africa 4: South Africa, Kerguelen Island, Tristan da Cuhna. 

^Callicostella applanata Broth. & Bryhn; C. fissidentella (Besch.) 
Kindb.; C. salaziae (Besch.) Kindb.; C. tristis (C. Miill.) Broth.; 
Cyclodictyon laete-virens (Hook, and Tayl.) Mitt.; C. vallis-gratiae 
(C. Mull.) Kuntze; C. vallis-gratiae f. breutelianum (Hampe) Demar. & 
P. Varde; * Daltonia tristaniensis Dix. in Christ.; -Distichophyllum 
fasciculatum Mitt.; D. imbricatum Mitt.; W. nniiifoliuni (Hornsch.) 
Sim; *Z>. taylorii Sim; *Eriopus mniaceum (C. Miill.) Broth.; 
Hookeriopsis papeana (Hampe) Jaeg*. 

Europe, Iceland, the Caucasus. 

Cyclodictyon laete-virens (Hook. & Tayl.) Mitt.; Daltonia splach- 
noides (Sm.) Hook. & Tayl.; Distichophyllum cariiiatum Dix. & Nich.; 
Eriopus apiculatus (Hook. f. & Wils.) Mitt, (introduced and established 
in England) ; Hooker ia lucens (Hedw.) Sm. 



Plant Taxonomy 383 

Asia 1 : North Asia including Sakhalin Island. 

Hookeria lucens (Hedw.) Sm. 

Asia 2: China, Mongolia, Japan, Korea, Taiwan (Formosa). 

Callicostella papillata (Mont.) Mitt.; Chaetomitriopsis glaucocarpa 
(Schwaegr.) Fleisch.; Cyclodictyon blumeanum (C. Mull.) Kuntze; 
Daltonia angustifolia Doz. & Molk. ; D. angustifolia var. strictifolia 
(Mitt.) Fleisch.; D. aristifolia Ren. & Card.; *Distichophylhim breviro- 
stratum Ther.; D. carinatwm Dix. & Nich.; *D. cavaleriei Ther.; *D. 
collenchymatosum Card.; D. cuspidatum (Doz. & Molk.) Doz. & Molk.; D. 
gracilicaule Fleisch.; D. jungermannioides (C. Mull.) Bosch & Lac; " : D. 
maibarae Besch.; D. mittenii Bosch & Lac; D. montagneanum (C. Mull.) 
Bosch & Lac; D. nigricaule Mitt, ex Bosch & Lac; *D. obtusif olium 
Ther.; *D. osterwaldii Fleisch.; *D, stillicidiorum Broth.; *Eriopus ja- 
ponicus Card. & Ther. ; E. parviretis Fleisch. ; *E. spinosus Nog. ; Hook- 
eria acutifolia Hook. & Grev.; Hookeriopsis pappeana (Hampe) Jaeg. ; 
H. utacamundiana (Mont.) Broth.; *H. yakushimensis Toyama. 

Asia 3: India, Pakistan, Ceylon, Burma, Thailand (Siam), Vietnam 

(Indochina). 

Actinodontium ascendens Schwaegr.; A. rhaphidostegum (C. Mull.) 
Bosch & Lac; Callicostella papillata (Mont.) Mitt,; C. prabaktiana (C. 
Mull.) Bosch & Lac; Chaetomitriopsis glaucocarpa (Schwaegr.) Fleisch.; 
Chaetomitrium ciliatum Doz. & Molk. ex Bosch & Lac; *C. confertum 
Thwait. & Mitt.; C. cucullatwm Dix.; C. leptopoma (Schwaegr.) Bosch & 
Lac; *C. nervosum Dix.; C. papillif olium Bosch & Lac; C. philippinense 
(Mont.) Bosch & Lac; C. torquescens Bosch & Lac; *C. volutum Mitt.; 
Cyclodictyon blumeanum (C. Mull.) Kuntze; Daltonia angustifolia Doz. & 
Molk.; D. angustifolia var. strictifolia (Mitt.) Fleisch.; *Z). apiculata 
Mitt.; *D. aristifolia Ren. & Card. ssp. leptophylla Fleisch.; *D. 
brevipedunculata Mitt.; D. contorta C. Mull.; D. contorta ssp. inac- 
gregorii (Broth.) Fleisch.; *D. flexifolia Mitt.; *D. gemmipara Dix.; *D. 
marginata Griff.; *D. perlaxiretis Dix.; *D. reticulata. C. Mull.; *D. 
semitorta Mitt.; *D. sub angustifolia Ren. & Card.; Dimorphocladon 
borneense Dix.; *Distichophyllum ceylanicum (Mitt.) Par.; D. cuspi- 
datum (Doz. & Molk.) Doz. & Molk.; *£>. griffithii (Mitt.) Par.; *D. 
heterophyllum (Mitt.) Par.; *D. humifusum (Mitt.) Par.; *Z>. limpidum 
Thwait. & Mitt.; *D. madurense Ther. & P. Varde; D. mittenii Bosch & 
Lac; D. montagneanum, (C. Miill.) Bosch & Lac; D. nigricaule Mitt, ex 
Bosch & Lac; *D. obovatum (Griff.) Par.; D. schmidtii Broth.; D. 
sinuosulum Dix.; 'D. succulentum (Mitt.) Broth.; *Eriopus bonianus 
Besch.; *£\ lucidus Thwait. & Mitt.; E. parviretis Fleisch.; E. remoti- 
folius C. Miill.; Hookeria, acutifolia Hook. & Grev.; Hookeriopsis pallidi- 
folia (Mitt.) Geh. & Herz.; *H. purpuvata (Mitt.) Broth.; *H. secunda 
(Griff.) Broth.; 'H. thwaitesiana, (Mitt.) Broth.; H. utacamundiana, 
(Mont.) Broth.; *Lepidopilidium furcatum (Thwait. & Mitt.) Broth.; 
*Orontobryum hookeri (Mitt.) Fleisch.; Pseudohypnella verrucosa (Doz. 
& Molk.) Fleisch. 



384 Indiana Academy of Science 

Asia 4: Indonesia, Malaya, Philippine Islands, New Guinea. 

*Actinodontium ascendens Schwaegr. ; A. rhaphidostegum (C. Mull.) 
Bosch & Lac; *Archboldieila pilifera Bartr.; *Callicostella aiomensis 
Bartr. ; *C. armata Herz.; *C. beccariana (Hampe) Jaeg\ ; *C. chloneura 
C. Mull.; *C. eberhardtiana Broth. & Par.; C. kaernbachii Broth.; *C. 
loriae Fleisch. ; C. papillata (Mont.) Mitt.; *C. papillata var. brevifolia 
Fleisch.; *C papillata var. viridissima Dix.; :|: C. paupera (C. Mull.) 
Kindb.; C. prabaktiana (C. Mull.) Bosch & Lac; *C. prabaktiana var. 
acuminata Baumg. ; *C. pterygophylloides (Broth.) Broth.; Chaetomitri- 
opsis glaucocarpa (Schwaegr.) Fleisch.; *C diversifolia Zant. ; *Chaeto- 
mitrium acanthocarpum Bosch & Lac; *C auriculatum Dix. & Herz.; 
*C. beccarii Dix.; *C. bornense Mitt.; *C. brassii Bartr.; C. ciliatmn 
Doz. & Molk. e:r Bosch & Lac; *C. crispifolium Bartr.; *C. ctenidioides 
Broth.; C. cucullatum Dix.; *C. divergens Dix.; *C. elegans Geh.; *C. 
elmeri Broth.; *C. elongatum (Doz. & Molk.) Doz. & Molk.; *C. 
everettii Mitt, e:r Dix.; ::: C. fimbriatum (Doz. & Molk.) Bosch & Lac; 
*C. finisterrae Dix. & Herz.; *C. horridulum Bosch & Lac; *C. integri- 
folium Bartr.; *C laevifolium Dix.; ::: C. laevisetum Dix.; :;: C. lanceolatum 
Bosch & Lac; *C. lancifolium Mitt.; :;: C. lauterbachii Broth.; C. lep- 
topoma (Schwaegr.) Bosch & Lac; *C. leptopoma var. massartii Ren. & 
Card.; :|: C. longisetulum Bartr.; *C. macrohystrix C. Mull. ; ::: C. madan- 
gense Bartr.; *C. nanohystrix C. Miill.; C. nematosum Broth.; C. 
orthorrhynchum (Doz. & Molk.) Bosch & Lac; *C. paleatum (Hampe) 
Fleisch.; C. papillifolium Bosch & Lac; *C. papuanum Bartr.; *C. 
parcesetulosum Bartr.; *C. perakense Broth.; ::: C. perarmatum Broth.; 
:i: C. perlaeve Dix.; C. philippinense (Mont.) Bosch & Lac; *C. plicatum 
Bartr.; ::: C. poecilophyllum Dix.; ::: C. pseudo-eiongatum Broth.; *C. 
pseudo- papillifolium Bartr.; ::: C. recurvifolimn Fleisch.; *C. rigidulum 
Broth.; *C. robbinsii Bartr.; :|: C. roemeri Fleisch.; *C. seriatum Broth.; 
:!: C. setosum Broth.; :;: C. spinosum Bartr.; ::: C. sublaevisetum Dix.; ::: C. 
subplicatum Bartr.; C. tahitense (Sull.) Mitt.; C. tahitense var. 
deplanchei (Besch.) Wijk & Marg. ; C. torquescens Bosch & Lac; *C. 
torquescens var. barbatum Dix.; *C vrieseanum Bosch & Lac; *C. 
warburgii Broth.; C. weberi Broth.; *C. werneri Herz.; *C. wildei Zant.; 
Cyclodictyon blumeanum (C. Miill.) Kuntze; *C blumeanum var. 
morokae Fleisch.; C. blumeanum var. vescoanum (Besch.) Fleisch.; C. 
lepidum (Mitt.) Broth. & Watts; Daltonia angustifolia Doz. & Molk.; 
*D. angustifolia var. gemmiphylla Fleisch.; *D. angustifolia var. longi- 
pedunculata (C. Miill.) Fleisch.; *D. angustifolia var. rcvoluta (Broth.) 
Bartr.; D. angustifolia var. strictifolia (Mitt.) Fleisch.; D. aristifolia 
Ren. & Card.; *D. armata Bartr.; *Z). baumgartneri Froehl.; D. contorta 
C. Miill.; D. contorta ssp. mac-gregorii (Broth.) Fleisch.; :|: Z). contorta 
var. humilis Fleisch.; *D. himalayeusis Dix. & Herz.; D. pseudosteno- 
phylla Bartr.; *D. schiffneri Froehl.; : D. tuberculosa Bartr.; Dimorpho- 
cladou borneense Dix.; *Distichophyllidium antarense Zant.; *D. junger- 
manniaceum Fleisch.; - : D. nymanianum Fleisch.; :]: D. rhizophorum 
Fleisch.; *Distichophyllum aciphyllum Dix.; *D. angustifolium Dix.; *D. 
angustissimum Dix.; *D. borneense Broth.; :|: Z). brevicuspidatum Bartr.; 
*D. brevicuspis Fleisch.; *D. catinifolium Froehl.; 'D. cucullatum Bartr.; 



Plant Taxonomy 385 

D. cuspidatum (Doz. & Molk.) Doz. & Molk. ; *D. denticulatum Dix.; *D. 
dixonii Herz.; *D. elmeri Broth.; *D. evanidolimbatum Fleisch.; D. graci- 
licaule Fleisch.; D. jungermannioides (C. Mull.) Bosch & Lac; -D. leio- 
pogo7i Dix.; *Z>. lixii (Broth.) Ther.; -D. longipes Broth.; *D. longobasis 
Fleisch.; *D. lorianum Fleisch.; *D. macropodium Dix.; D. mittenii Bosch 
& Lac; D. montagneanum (C. Mull.) Bosch & Lac; *D. nidulans Herz.; D. 
nigricaule Mitt, ex Bosch & Lac; *Z>. nigricaule var. cirratum (Ren. & 
Card.) Fleisch.; *D. nigricaule var. complanatum Fleisch.; *D. perundu- 
latum Dix.; *D. pullei Dix.; *D. santosii Bartr. ; D. schmidtii Broth.; 
D. sinuosulum Dix.; *Z>. spathulatum (Doz. & Molk.) Doz. & Molk.; *D. 
stipitati folium C. Mull.; *Z). submucronatum Fleisch.; *D. subnigricaiile 
Broth.; *D. tortile Doz. & Molk.; *D. turgidum Bartr.; D. undulatum 
Doz. & Molk.; *Eriopus cristatus (Hedw.) Brid.; *E. flaccidus Broth.; 
■ f E. microblastus Broth.; E. parviretis Fleisch.; *E. perlimbatiis Dix.; 
*£'. ramosus Fleisch.; E. remotifolius C. Mull. ; E. subremotifolius Broth.; 
Hookeria acutifolia Hook. & Grev.; *Hookeriopsis gemmidens Broth.: 
*H. ?nacropus (Bosch & Lac) Broth.; *//. majurei Bartr.; H. utacamun- 
diana (Mont.) Broth.; : H. wichurae Fleisch.; Leskeodon acuminatus 
(Bosch & Lac) Fleisch.; *L. acuminatus var. laeviseta Zant. ; *L. pan- 
durifolius Fleisch.; *L. philippinensis Broth.; *L. robbinsii Bartr.; 
*L. rotundifolius Bartr.; *Leskeodontopsis pustulata Zant.; Pseudo- 
hypnella verrucosa (Doz. & Molk.) Fleisch.; *Pterygophyllum javense 
Dix.; *P, novae-gaineae Bartr.; *Sauloma tenuis C. Mull. 

Asia 5: Asiatic part of the Middle-East, including Cyprus. 

Hookeria lucens (Hedw.) Sm. 

Australia 1 : Australia and Tasmania. 

■ f CallicosteUa bailey i (Broth.) Kindb.; C. kaernbachii Broth.; *C. 
rugiseta Dix.; *Chaetomitrium entodontoides Broth. & Watts; C. tahi- 
tense (Sull.) Mitt.; C. tahitense var. deplanchei (Besch.) Wijk & Marg. ; 
Cyclodictyon lepidum (Mitt.) Broth. & Watts; *Daltonia pusilla Hook. f. 
& Wils.; D. splachnoides (Sm.) Hook. & Tayl.; DistichophyUum assimile 
Broth.; *D. baileyanum C. Mull. ; *D. beccarii (Harape & Geh.) Par.; 
*D. complanatum (Hampe) Mitt.; D. crispulum (Hook. f. & Wils.) Mitt.; 
*jD. levieri (Geh.) Broth.; *D. longicuspis Broth.; D. microcarpum 
(Hedw.) Mitt.; *D. minutif olium C. Mull.; D. pulchellum (Hampe) 
Mitt.; D. rotundif olium (Hook. f. & Wils.) C. Mull. & Broth.; *£>. 
sub minutif olium (Broth. & Geh.) Fleisch.; *D. whiteleggeanum C. Mull.; 
Eriopus apiculatus (Hook. f. & Wils.) Mitt.; *E. brassii Bartr.; E. 
cristatus (Hedw.) Brid. in C. Mull.; *E. tasmanicus Broth.; *Pterygo- 
phyllum bryoides Broth.; P. dentatum (Hook. f. & Wils.) Dix.; 
*P. flaccidissimum Broth.; P. obscurum Mitt.; *P. subrotundum (Hampe) 
Jaeg. ; *P. wattsii Broth.; Sauloma tenella (Hook. f. & Wils.) Mitt.; 
*S. zetterstedtii (C. Mull.) Jaeg. 

Australia 2: New Zealand, etc. 

*Bellia nervosa (Hook. f. & Wils.) Broth.; Daltonia angustifolia 
Doz. & Molk.; D. splachnoides (Sm.) Hook. & Tayl.; DistichophyUum 
crispulum (Hook. f. & Wils.) Mitt.; *Z). crispulum var. adnatum (Hook. 



386 Indiana Academy of Science 

f. & Wils.) Dix.; D. microcarpum (Hedw.) Mitt.; f D. microcarpum var. 
homodictyon Sainsb.; *D. microcladum (Col.) Broth.; D. pulchellum 
(Hampe) Mitt.; *D. pulchellum, var. elliptic? folium Sainsb.; *D. pul- 
chellum var. parvirete Sainsb.; D. rotundifolium (Hook. f. & Wils.) 

C. Mull. & Broth.; Eriopus apiculatus (Hook. f. & Wils.) Mitt.; *E. 
brownii Dix.; E. cristatus (Hedw.) Brid. in C. Mull.; E. flexicollis 
(Mitt.) Jaeg.; *Hookeria flava Col.; Pterygophyllum dentatum (Hook. f. 
& Wils.) Dix.; *P. dentatum var. latifolium (C. Miill.) Wijk & Marg.; 
*P. dentatum var. robustum (Hook. f. & Wils.) Dix.; *P. distichophyl- 
loides Broth. & Dix.; *P. quadrif avium (Sm.) Brid.; *P. quadvif avium f. 
mavginata Sainsb.; *Sauloma macvospova Sainsb.; S. tenella (Hook. f. & 
Wils.) Mitt.; *S. tenella f. pvopagulifeva Sainsb. 

Oceania: Pacific Islands. 

*Callicostella bisexualis (Besch.) Jaeg.; :;: C. caledonica Ther.; *C. 
coMipbelliana (Hampe) Jaeg. ; *C. chlovina (Besch.) Broth.; :|: C. fvatevi 
Broth. & Watts; *C. melanotheca (Besch.) Jaeg.; *C. melanotheca var. 
scabviseta Ther.; :i: C. nukahivensis (Besch.) Broth.; *C. oblongifolia 
(Sull.) Jaeg.; C. papillata (Mont.) Mitt.; C. prabaktiana (C. Miill.) 
Bosch & Lac; *C. vesiculata (C. Miill.) Jaeg.; *Chaetomitvium- aneitense 
Broth. & Watts; *C. callichvoum Besch.; *C. densum Dix. ex Bartr. ; 
*C. depvessum Mitt.; :!: C. frondosum Mitt.; :;: C. hebridense Dix.; C. 
ovthovvhynchum (Doz. & Molk.) Bosch & Lac. var. vitense Bartr.; 
*C. vugifolium (Sull.) Mitt.; :!: C. smithii Bartr.; *C. speciosum (Sull.) 
Mitt.; C. tahitense (Sull.) Mitt.; C. tahitense var. deplanchei (Besch.) 
Wijk & Marg.; C. weberi Broth.; *C. wheeleri Hampe; *Cyclodictyon 
bescherellei (Par.) Broth.; C. blumeanum (C. Miill.) Kuntze; *C. blumea- 
nwm f. gvaeffeana (C. Miill.) Fleisch.; C. blumeanum var. vescoanum 
(Besch.) Fleisch.; *Daltonia baldwinii Broth.; D. contorta C. Miill.; 

D. pseudostenophylla Bartr.; *D. vufescens Broth.; *D. sphaevica Besch.; 
*Distichophyllidium maticiim Broth. & Par.; *Distichophyllum apicidi- 
gevum Broth. & Par.; *D. capillatum Mitt.; D. cuspidatum (Doz. & 
Molk.) Doz. & Molk.; D. fasciculatum Mitt.; *D. flavescens (Mitt.) Mitt. 
in Seem.; :] D. fossombvonioides Ther.; *Z). fvancii Ther.; *Z>. fveycinetii 
(Schwaegr.) Mitt.; *D. fveycinetii var. crassetuvgescens C. Mull.; *D. 
graeffeanum (C. Miill.) Broth.; D. imbvicatum Mitt.; *D. koghiense 
Ther.; *D. limbatulum (C. Miill.) Par.; 'D. lingulatum Bartr.; D. mit- 
tenii Bosch & Lac; D. montagneanum (C. Miill.) Bosch & Lac; :]: D. 
nadeaudii Besch. ; V D. pavadoxum (Mont.) Mitt.; *D. samoanum Fleisch.; 
■•'D. samoanum var. bvevipes Bartr.; ' : D. semimavginatum Ther.; *D. 
tahitense Besch.; *D. tovquati folium Dix.; D. undulation Doz. & Molk.; 

D. vitianum (Sull.) Mitt.; Eriopus cristatus (Hedw.) Brid. in C. Mull.; 
*E. marginatus Ther.; *E. pacificus (Besch.) Fleisch..; E. remotifolius C. 
Miill.; *E. subvemotifolius Broth.; Hookevia acutifolia Hook. & Grev. ; 
*H. sandvicensis Reichdt. ; *Hookeviopsis purpurea (C. Miill.) Broth.; 
*H. purpurea var. acuminatula, (C. Miill.) Bartr.; *H. purpurea var. 
ligulacea (C. Miill.) Bartr.; Leskeodon acuminatus (Bosch & Lac.) 
Fleisch. 



Plant Taxonomy ::x7 

Literature Cited 

Paton, Jean A. 1968. Eriopus apiculatus (Hook. f. & Wils. ) Mitt, established on 
Tresco. Trans. Brit. Bryol. Soc. 5(3) :460-462. 

Welch, Winona H. 1962. The Hookeriaceae of the United States and Canada. The 
Bryologist 65(1) :l-24. 

— . 1966. The Hookeriaceae of Mexico. The Bryologist 69(1) :l-68. 



— . 1968. Hookeriaceae Species and Distribution in North and Central 
America and West Indies. Proc. Indiana Acad. Sci. 77 :351-356. 

. 1969. The Hookeriaceae of Cuba. The Bryologist 72(2) :93-136. 

— . 1969. Hookeriaceae Species and Distribution in South America. Proc. 



Indiana Acad. Sci. 78 :396-405. 

7. Wijk, R. van der, W. D. Margadant, and P. A. Florschmtz. 1959, 1962, 1964, 1967, 
1969. Index Muscorum. Vol. 1-5. Utrecht, The Netherlands. 



Biosystematic Studies of the Beech and Marsh Ferns 1 

Jeanette C. Oliver, Ball State University 



Abstract 

The taxonomic status of five species of ferns common to the Northeastern United 
States has been in a state of confusion for many years. The broad beech fern, the long 
beech fern, the marsh fern, the New York fern, and the bog fern have been placed in 
the genus Thelypteris by some authors. Others consider them members of the large 
cosmopolitan genus Dryopteris. In other interpretations the beech ferns are separated 
into the genus Phegopteris. 

Evidences from morphological, cytological, and biochemical studies do not warrant the 
inclusion of these species in the genus Dryopteris. Significant differences were noted 
between the beech ferns and the marsh ferns which would appear to justify their 
separation into different genera. 

This study was undertaken in an attempt to clarify the confusion 
regarding the taxonomic status of five species of ferns common to the 
northeastern United States, These ferns, and their counterparts from 
other regions, have been variously classified generically. 

Some have considered the broad beech fern, the long beech fern, 
the bog fern, the marsh fern, and the New York fern as being of com- 
mon generic rank. These ferns were treated as members of the large 
cosmopolitan genus Dryopteris by Christensen in his monograph (4, 5). 
This interpretation was followed by Fernald in his Gray's Manual of 
Botany (9) and by Deam in his Flora of Indiana (8) (Table 1). 

Christensen, in a later work, set the genus Thelypteris, with the 
marsh fern, Thelypteris palustris as its type, apart from the genus 
Dryopteris (6). Morton (12) in his contributions to Britton and Brown's 
Illustrated Flora (10) recognized the differences between the "thelyp- 
teroid" and the "dryopteroid" ferns (Table 1). 

On the basis that Thelypteris Schmidel was not validly published, 
Copeland (7) in his Genera Filicum used the name Lastrea Bory. The 
Nomenclature Committee at the International Botanical Congress in 
Stockholm decided, however, that the name Thelypteris is correct and 
voted down a proposal to conserve Lastrea (13, 14). 

Cytologically the beech ferns differ from both the dryopteroids and 
the thelypteroids; further, their sori lack indusia in contrast with the 
latter. Therefore, some feel that these ferns should be placed in a 
separate genus, Phegopteris (11, 21) (Table 1). 

Methods and Materials 

Extensive collections of the five species were made throughout 
Indiana and Ohio. A total of 422 herbarium specimens from the Field 
Museum of Natural History and 432 specimens from the Missouri 
Botanical Garden was examined. 



1 This work was supported by grants from the Indiana Academy of Science, Sigma 
Xi, and Ball State University. 

388 



Plant Taxonomy 



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390 Indiana Academy of Science 

The external morphology was carefully studied. Drawings and photo- 
graphs were made to record distinguishing characteristics. 

Rhizomes and stipes were cross sectioned at 10> and stained with 
safranin-fast green. 

Spore samples were obtained from both living and moribund speci- 
mens. The spores were mounted in Hoyer's medium for clearing and 
preservation. 

For cytological study, meiotic materials were fixed in 3 parts ethyl 
alcohol : 1 part glacial acetic acid. Squashes were made in aceto-orcein. 

Biochemical comparisons were made by means of paper chromatog- 
raphy. Extracts were prepared by powdering dried fronds and soaking 
the materials in 80% methanol for 48 hours. Fifty microliters of each 
sample were applied to Whatman #1 paper using the spot method. Both 
1 and 2-dimensional chromatograms were run descendingly using bu- 
tanolracetic acidiwater (12:3:5) as the solvent. 

Dried chromatograms were examined in the presence of ultra- 
violet light. The chromatograms were sprayed sequentially with ninhy- 
drin and Ehrlich reagent for the detection of free-amino acids and re- 
lated substances. Some chromatograms were sprayed with Pauly's 
reagent or alkaline silver nitrate for the detection of phenolic com- 
pounds (16). 

Morphological Observations 

The frond of the beech ferns consists of a deltoid blade borne by 
a pilose stipe. The lanceolate apex of the blade is pinnatifid. 

The blade of the long beech fern has 10-12 pairs of separate pinnati- 
fid pinnae below the apex. The length of the blade exceeds its breadth at 
the widest point. Nine to 18 pairs of surcurrent pinnae are noted in the 
broad beech fern. The lower 3-4 pairs are bipinnatifid and the upper ones 
are pinnatifid (Fig. 1, 2). 

Both species bear rounded, supramedial sori which lack indusia. 

The blades of the New York, bog, and marsh ferns are lanceolate in 
outline and are composed of pinnatifid segments below a pinnatifid apex. 
The dark stipe is glabrous in the marsh fern and scaly in the other two 
species. 

The blade of the New York fern is composed of 18-40 pairs of 
oblong-lanceolate pinnae. The lower 4-5 pairs are greatly reduced in 
size (Fig. 3). The sori are located submarginally and have a kidney- 
shaped indusium. 

The marsh fern possesses 13-25 pairs of linear-lanceolate pinnae 
which bear medial indusiate sori (Fig. 4). 

The blade of the bog fern is composed of 10-38 pairs of oblong- 
lanceolate pinnae. The lowermost are basioscopic (Fig. 5). The indusiate 
sori are borne submarginally. 



Plant Taxonomy 



8i)l 






I 



/m T , t Pwt T aphS ° f herba ™™ specimens. 1) Phegopteris hexagonoptera 
(Mich*.) Feet broad beech fern; 2) Phegopteris polypodioides Fee> long beech fcrn . 

Thelypteris noveboracensis (L.) Nieuwl, New York fern; 4) Thelypteris palustris Schott 
marsh fern; 5) Thelypteris simulata (Davenp.) Nieuwl., bog fern 

Figures 6-10. Fern spores X 675. 6) Broad beech fern; 7) Long beech fern; 8) Bog 
fern; 9) Marsh fern, 10) New York fern. 



392 Indiana Academy of Science 

Spore Morphology 

The spores of all five species are bilateral in shape and bear an 
adherent perispore. Spores of the beech ferns are light greenish-brown 
in color. Their perispore does not form conspicuous ridges or wings. 
The spores of the broad beech fern are 20-27 x 30-50/a. The surface is 
tuberculate in texture. The perispore is not winged (Fig. 6). Spores of 
the long beech fern are 28-30 x 55-60/a. The spore surface is tuberculate. 
The perispore varies, some spores being wingless and ridgeless and 
others bearing short broken ridges and narrow discontinuous wings 
(Fig. 7). 

The remaining species are yellow-brown in color and possess a 
tuberculate winged and ridged perispore. The spores of the marsh fern 
are 34-45 x 53-63/u. Ridges are few and broken (Fig. 9). The New York 
fern spores are highly sculptured with wide apical wings. The spores are 
25-30 x 33-45/* (Fig. 10). The spores of the bog fern are sculptured 
similarly to those of the New York fern, but approach those of the marsh 
fern in size (Fig. 8). 

Cytological Observations 

It is apparent from this and other studies that different chromosome 
numbers exist among the thelypteroid ferns. 

Broad beech fern materials collected in Indiana showed a chromosome 
number of n = 30 (Voucher Specimen #0159 Oliver). Counts made of 
Virginia specimens by Wagner (18) and by Britton (1, 2, 3) of materials 
from Ontario confirm this number. 

Studies of the long beech fern have been made by Manton (11) in 
Great Britain and by Britton (1, 2, 3). Both have reported numbers of 
n = 90 and In = 90. The long beech fern is apogamous. A doubling of 
the diploid number of 90 chromosomes occurs just prior to meiosis; thus 
the sporophytic and the gametophytic numbers are the same. 

Counts of the marsh fern (Voucher #0172 Oliver) and the New York 
fern (Voucher #0195 Oliver) yielded numbers of n = 35 and n = 27, 
respectively. These numbers are in agreement with those reported by 
Wagner (17, 18). Britton reported a number of n = 35 for the marsh 
fern; however, his tentative studies of the New York fern indicated 
n = 29 (1). 

Meiotic material of the bog fern was unavailable for this study. 
Wagner reported a number of n = 64 for specimens collected in 
Maryland (17). 

Biochemical Observations 

Chromatograms were prepared from dried specimens of ferns col- 
lected throughout the northeastern United States. Materials had been 
preserved for periods of 1 month to 25 years. Remarkable consistency 
was obtained from materials of different ages and localities. 



Plant Taxonomy 393 

Examination of chromatograms in the presence of ultraviolet light 
revealed one compound with a rf value of 0.67 common to all species. 
Another substance, rf 0.80, was common to the marsh, the bog, and the 
New York fern. Several additional spots appeared to be species specific. 

Subsequent spraying revealed all ninhydrin reacting compounds to be 
species specific. The compound common to all and that common to the 
three species were indicated by their positive reactions with Pauly's 
reagent and alkaline silver nitrate to be phenolic (Fig. 11). 

Negative results were obtained with Ehrlich's reagent in all cases. 







* / 



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On 

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L . _ i 

Figure 11. One-dimensional descending chromatograms. A. Marsh fern, B. Bog fern, 
C. Long beech fern, D. Broad beech fern, E. Netv York fern (n =z ninhydrin postive 
substance p = phenolic compound) . 



Discussion and Summary 

All aspects of this study indicate exclusion of the beech and marsh 
ferns from the genus Dryopteris. 

The blades of these ferns are light green, thin-membranous, and non- 
evergreen; whereas, those of true Dryopteris are deep green, subcori- 
aceous and evergreen. The vascular bundles reach the margins of the 
pinnae in contrast to those of the dryopteroids which terminate in 
hydathodes. 



394 Indiana Academy of Science 

The thelypteroids bear slender rhizomes; the stipes contain two 
vascular bundles which unite at the base of the blade. Massive rhizomes 
and stipes bearing 3-7 bundles are characteristic of the dryopteroids. 

The spores are distinguished with some difficulty. Basically, the 
spores of the thelypteroids have a tuberculate surface; perispore ridges, 
if present, are quite discontinuous. Dryopteroid spores tend to be cristate 
with closed, anastomosing ridges (15). 

Extensive cytological studies have been made of the American 
species of Dryopteris sensu stricto (19, 20). A base chromosome number 
of n = 41, accompanied by various levels of ploidy, was noted. This 
number has not been seen in any thelypteroid fern. 

There appears to be much basis for maintaining the beech ferns as 
a separate genus Phegopteris. Some special characters common to the 
beech ferns as opposed to the marsh ferns are as follow: deltoid blades; 
exindusiate sori; spores wingless or bearing much reduced broken wings; 
chromosomes extremely small. Chromatographically, fewer common 
phenolic substances were detected in the beech than in the marsh ferns 
(Fig. 11). 

In conclusion, the results of this study indicate the exclusion of the 
thelypteroid ferns from Dryopteris; further basic differences appear to 
justify the delimitation of the beech ferns as a distinct genus. A number 
of outstanding problems are inherent in these ferns. The relationship of 
the oak-fern to the beech ferns is one such problem. Another is the rela- 
tionships of the thelypteroids of the western United States to those of 
the east. Extensive studies of Western North American, Asian, and 
European counterparts are needed before the taxonomy of the group can 
be understood fully. 

Literature Cited 

1. Britton, D. M. 1953. Chromosome studies on ferns. Amer. J. Bot. 40 :575-583. 

2. Britton, D. M. 1961. The problems of variation in North American Dryopteris. 
Amer. Fern. J. 51 :23-30. 

3. Britton, D. M. 1965. The cytology and distribution of Dryopteris species in 
Ontario. Can. J. Bot. 44 :63-78. 

4. Christensen, Carl. 1913. A monograph of the genus Dryopteris Part I. The tropical 
American pinnatifid-bipinnatifid species. Kgl. Dansk. Vid. Selsk. Skr., VII. 10 :55-582. 

5. Christensen, Carl. 1920. A monograph of the genus Dryopteris Part II. The 
tropical American bipinnate-deeompound species. Kgl. Dansk. Vid. Selsk. Skr., 
VIII. 6:1-123. 

6. Christensen, Carl. 1938. Manual of Pteridology. Chronica Botanica Press. 550 p. 

7. Copeland, E. B. 1947. Genera Filicum. Chronica Botanica Press. 247 p. 

8. Beam. C. C. 1940. Flora of Indiana. Department of Conservation, Indianapolis. 
1236 p. 

9. Fernald, M. L. 1950. Gray's Manual of Botany. American Book Co., New York. 
1632 p. 



Plant Taxonomy 395 

10. Gleason, H. A. 1952. Illustrated Flora of the Northeastern United States and 
Adjacent Canada. New York Botanical Garden, New York. Vol. 1. 482 p. 

11. Manton, I. 1950. Problems of cytology and evolution in the Pteridophyta. University 
Press, Cambridge, Eng. 316 p. 

12. Morton, C. V. 1950. Notes on the ferns of the eastern United States. Amer. Fern 
J. 40:213-225. 

13. Morton, C. V. 1958. The Californian species of Thelypteris. Amer. Fern J. 
48:136-162. 

14. Morton, C. V. 1963. The classification of Thelypteris. Amer. Fern J. 53:149-154. 

15. Oliver, J. C. 1968. A study of spore characteristics of the ferns of Indiana. Amer. 
Fern J. 58 :5-12. 

16. Smith, I. 1958. Chromatographic techniques: Clinical and biochemical application. 
Heinemann Medical Books, Ltd. London. 419 p. 

17. Wagner, W. H. Jr. 1963. A biosystematic survey of United States ferns — pre- 
liminary abstract. Amer. Fern J. 53 :1-16. 

18. Wagner, W. H. Jr. 1966. Pteridophytes of the Mountain Lake area, Giles Co., 
Virginia: Biosystematic studies, 1964-5. Castanea 31:121-140. 

19. Walker, S. 1961. Cytogenetic studies in the Dryopteris spinulosa complex II. 
Amer. J. Bot. 48 :607-614. 

20. Walker, S. 1962. Further studies in the genus Dryopteris; the origin of D. clintoni- 
ana, D. celsa and related taxa. Amer. J. Bot. 49 :497-503. 

21. Wherry, E. T. 1937. Guide to eastern ferns. Science Press, Lancaster, Pa. 252 p. 



Cytotaxonomic Notes on Genus Polygonum, Section Polygonum 1 

Donald N. Moore, Thomas R. Mertens and Joyce E. Highwood, 
Ball State University 



Abstract 

The most accurate identification of species in genus Polygonum, section Polygonum 
is based on fruit and perianth characteristics. Preliminary investigations suggested that 
these morphological features could be effectively correlated with specific chromosome 
numbers. The achenes of five species of Polygonum collected in Indiana, Nova Scotia and 
New Brunswick are briefly described and chromosome numbers for these species are 
reported as follows: P. aviculare L. sensu stricto 2n = 60; P. buxijorme Small In = 
60; P. arenastrum Bor. In = 40; P. fowleri Robinson 2n = ca. 40; and P. erectum L. 
2« = 40. 

Although numerous taxonomic investigations of genus Polygonum, 
section Polygonum (Avicularia) have been conducted, there is little 
agreement as to species identity in North America or elsewhere. Much of 
the taxonomic confusion comes from the tremendous morphological 
variation within the individual species of the section. At present the 
most accurate identification of Polygonum specimens is based on fruit 
and perianth characteristics. The purpose of this investigation was to 
attempt to gain evidence to support this basis of identification by showing 
that morphological features can be correlated with chromosome numbers. 
Plants identified as a given species on the basis of morphological charac- 
ters may be expected to have identical chromosome numbers. 

The fruit of Polygonum is a small, dry, indehiscent achene with a 
relatively thin wall. According to Styles (5), Mertens and Raven (2), 
Savage and Mertens (4), and Mertens (3), fruit and perianth character- 
istics are the most consistent morphological features used in the 
identification of species within section Polygonum. The criteria employed 
for identification in this investigation included: achene color, texture, 
shape, and size; the depth of perianth sinuses; and the position of 
inflorescences on the stem. Inflorescences are located either at the apices 
of branches or in the axils of the leaves along the stem, depending on the 
species. 

Cytological investigation by Love and Love (1) has revealed that the 
chromosomes of species within section Polygonum are uniformly small 
in size with centrally located centromeres. The present investigation sup- 
ports the same conclusion. Love and Love claim that the basic chromo- 
some number of the section is 2n = 20 with tetraploid and hexaploid 
plants being quite common. Mitotic chromosome counts of 40 and 60 
for species in section Polygonum have been frequently reported in the 
literature (2, 5, 6). 

This report consists of a brief discussion of achene characteristics 
and chromosome numbers for several species of section Polygonum 
collected in Indiana, Nova Scotia and New Brunswick. 



1 This study was supported by grants to Dr. Mertens from the Indiana Academy of 
Science and The Society of Sigma Xi. 

396 



Plant Taxonomy 



897 



# 



**A 





I 

a 



w •*%%> 



B 



1 *t 



f 




f 



t 



' 







Figure 1. Typical achenes. A. Polygonum aviculare L., B. P. buxifoime Small, and C. P. 
fowleri Robinson. 



398 Indiana Academy of Science 

Polygonum aviculare L. sensu stricto, has achenes (Fig. 1A) which 
are dull and heavily striated (5). Their color ranges from dark brown to 
black. The achene has two more-or-less equal concave sides and a third 
flat side, and is completely enclosed in the perianth, which is divided for 
three-fourths or more of its length. The achenes of P. aviculare from 
Nova Scotia and New Brunswick varied from 3.58 to 4.08 mm in length 
(mean: 3.93 mm) and from 2.16 to 2.92 mm in width (mean: 2.54 mm). 

Chromosome counts of 2n — 60 and 2?? — ca. 60 were obtained for 
P. aviculare from specimens collected in Nova Scotia and New Brunswick. 
Counts of 2n = 60 for this species are well documented in the literature 
(2,5,6): 

1) P. aviculare L., rocky beach on Sand Beach south of Yarmouth, 
Yarmouth Co., Nova Scotia, August 14, 1968, T. R. Mertens NS-2. 
(BSU). 2n = 60. 

2) P. aviculare L., Yarmouth Co., Nova Scotia, August 16, 1968, 
T. R. Mertens NS-16. (BSU). 2m = ca. 60. 

3) P. aviculare L., St. John Co., New Brunswick, August 19, 1968, 
T. R. Mertens NB-35. (BSU). 2w = ca. 60. 

4) P. aviculare L., Deer Island, Charlotte Co., New Brunswick, 
August 19, 1968, T. R. Mertens NB-42. (BSU). 2n = ca. 60. 

The achenes (Fig. IB) of Polygonum buxiforme Small are small in 
size in comparison to those of the closely related P. aviculare. Specimens 
from Nova Scotia and New Brunswick measured 2.58 to 3.16 mm in 
length (mean: 2.87 mm) and 1.58 to 2.93 mm in width (mean: 2.25 mm). 
The achene is dull to mildly shiny, dark brown in color and heart-shaped 
with two equal, concave sides and one flat side. The achene is enclosed 
in the perianth, which has a distinctive flanged edge (Fig. IB). The 
perianth is divided for % to % of its length. 

Chromosome numbers of 2n =z 60 and 2n — ca. 60 were obtained for 
the following specimens of P. buxiforme: 

1) P. buxiforme Small, Dearborn Co., Indiana, July 9, 1966, A. D. 
Savage 17-2. (BSU). 2n = 60. 

2) P. buxiforme Small, Dearborn Co., Indiana, July 9, 1966, A. D. 
Savage 17-3. (BSU). 2n = 60. 

3) P. buxiforme Small, Halifax Co., Nova Scotia, August 18, 1968, 
T. R. Mertens NS-26. (BSU). 2n = 60. 

4) P. buxiforme Small, Queens Co., Nova Scotia, August 17, 1968, 
T. R. Mertens NS-22. (BSU). 2n = ca. 60. 

5) P. buxiforme Small, Queens Co., Nova Scotia, August 17, 1968, 
T. R. Mertens NS-23. (BSU). 2n = ca. 60. 

6) P. buxiforme Small, Lunenburg Co., Nova Scotia, August 17, 
1968, T. R. Mertens NS-24. (BSU). 2n = ca. 60. 



Plant Taxonomy 399 

Polygonum arenastrum Bor., is characterized by achenes which are 
dark brown with two more-or-less equal convex and one narrow concave 
side. Savage and Mertens (4) report that the achenes of this species 
range from 1.58 to 2.50 mm in length and from 1.00 to 1.75 mm in 
width. The achene surface is dull but shiny along the edges. 

A chromosome count of 2n — 40 was obtained from the following 
specimen collected in Indiana: 

1) P. arenastrum Bor., Delaware Co., Indiana, September 18, 1968, 
Joyce Highwood 1-2. (BSU). 2n — 40. 

Another specimen, identified on the basis of morphological features 
as P. buxiforme, was found to have a diploid chromosome number of 40 
thus suggesting that it was, in fact, P. arenastrum. 

2) P. arenastrum Bor., (?) Warrick Co., Indiana, September 17, 
1966, A. D. Savage 62-1. (BSU). 2n = 40. 

Styles (5) and Mertens and Raven (2) also report 2n = 40 for 
P. arenastrum. 

Polygonum fowleri Robinson (= P. allocarpum Blake) produces 
highly distinctive achenes having a granular texture and a "beak-like" 
apex (Fig. 1C). A high incidence of biconvex fruits are encountered in 
this species (2). The length of the achenes determined from the study of 
specimens from New Brunswick, Canada, ranged from 2.92 to 3.67 mm 
with an average of 3.34 mm. The width of these achenes ranged from 
1.63 to 2.08 mm with the average of 1.85 mm. 

The color of the granular achenes varied from a light to a medium 
brown. Although normally three sided with two sides convex and one 
side concave, some achenes have only two sides (Fig. 1C). 

The light green perianth completely encloses the achene of P. 
fowleri. The perianth is divided for % to % of its length, varying with 
individual specimens. The inflorescences appear in the axils of the 
leaves along the stem. 

Taylor and Mulligan (6) report n = 20 for P. fowleri specimens 
from Graham Island in the Queen Charlotte Islands, British Columbia, 
Canada. Supporting their data are the following chromosome counts of 
2n r= ca. 40 obtained from specimens of P. fowleri collected in New 
Brunswick: 

1) P. fowleri Robinson, Lord's Cove, Deer Island, Charlotte Co., 
New Brunswick, August 19, 1968, T. R. Mertens NB-36. (BSU). 
2n = ca. 40. 

2) P. fowleri Robinson, Lord's Cove, Deer Island, Charlotte Co., 
New Brunswick, August 19, 1968, T. R. Mertens NB-38. (BSU). 
2n = ca. 40. 

The achenes of Polygonum erectum L. are light brown to tan in 
contrast to the much darker fruits exhibited by the other species of 



400 Indiana Academy of Science 

section Polygonum. The P. erectum achene is dull and granular with 
two convex and one concave side. It has been reported that the achenes 
range from 2.33 to 2.92 mm in length and 1.58 to 2.17 mm in width (4). 
The perianth is characteristically bottle-shaped and divided for less 
than one-half of its length. 

The chromosome number of 2n — 40 was established for the 
following specimens from Indiana: 

1) P. erectum, L., Porter Co., Indiana, August 29, 1966, A. D. 
Savage 58-1. (BSU). 2m = 40. 

2) P. erectum L,, Porter Co., Indiana, August 29, 1966, A. D. 
Savage 58-2. (BSU). 2n - 40. 

Voucher specimens for the plants for which chromosome numbers 
are reported in this paper are housed at Ball State University. 

This investigation supports the importance of fruit and perianth 
characteristics and chromosome number in the identification of species 
within Polygonum, section Polygonum. Members of a given species, 
identified on the basis of morphological features, may be expected to 
have identical chromosome numbers. This was generally found to be 
the case in the present investigation. Chromosome numbers reported 
herein agree with those reported in the literature in those cases where 
such reports are known. On the other hand, the fact that morphologically 
distinct species {e.g., P. arenastrum, P. fowleri, and P. erectum) have 
identical chromosome numbers {i.e., 2n = 40) indicates that chromo- 
some number alone cannot be used in identifying species in section 
Polygonum. 



Literature Cited 

1. Love, A., and D. Love. 1956. Chromosomes and taxonomy of eastern North American 
Polygonum. Can. J. of Bot. 34:501-521. 

2. Mertens, T. R., and P. H. Raven. 1965. Taxonomy of Polygonum, section Polygonum 
(Avicularia) in North America. Madrono 18(3):85-92. 

3. Mertens, T. R. 1968. Polygonum — A knotty problem in plant taxonomy. Amer. Biol. 
Teacher 30(10) :832-840. 

4. Savage, A. D., and T. R. Mertens. 1967. A taxonomic study of genus Polygonum, 
section Polygonum (Avicularia) in Indiana and Wisconsin. Proc. Indiana Acad. Sci. 
7 7:357-369. 

5. Styles, B. T. 1962. The taxonomy of Polygonum aviculare and its allies in Britain. 
Watsonia. 5:177-214. 

6. Taylor, R. L., and G. A. Mulligan. 1968. Flora of the Queen Charlotte Islands Part 
2. Cytological aspects of the vascular plants. Research Branch, Canada Department of 
Agriculture. Monogr. No. 4 Part 2. Ottawa, Canada. 148 p. 



SOIL SCIENCE 

Chairman: James E. Newman, Purdue University 
Clyde W. Hibbs, Ball State University, was elected Chairman for 1970 

Distribution of Corn (Zea mays L .) Roots in Two Soils in 
Relation to Depth of Sampling and Type of Sampler 1 

Russell K. Stivers, Purdue University 13 

Abstract 

A high yielding field containing both Crosby silt loam and Ragsdale silty clay loam 
was selected for this study. The roots of the crop were sampled while the corn was in 
the maturation or ear filling stage. Soils cores in 15 cm (6 in.) vertical samples were 
taken in each soil type with a bucket auger and with a Pit-O-Matic core sampler to a 
depth of 152 cm (60 in) where possible. Gravel in the Crosby profiles made it im- 
possible to sample deeper than 107 cm (42 in.) in 4 of the 5 subsampling areas. Roots 
and soil were separated. Dry weights of roots decreased highly significantly with depth. 
The 0-30 cm (0-12 in.) depths contained from 75 to 82% of the total weight of roots in 
the sampled profiles. Less than 2'/ ( of the roots were below 107 cm (42 in.) The bucket 
auger took more soil and with it more roots per 15 cm vertical sample than did the 
Pit-O-Matic core sampler. Sample holding diameters of these two tools were the same, 
but sample cuttings diameters were different. 

Distribution of corn roots in soil is important because corn plants 
obtain water and nutrients from soil. Fehrenbacher (2, 3) has shown 
that physical properties of soil and their genetic horizons are related 
to available water-holding capacity and to root penetration and total 
weight. The purpose of this study was to compare vertical distribution 
of corn roots by using two different soil sampling tools on two different 
adjoining high-yielding soils. 

Methods and Procedures 

Hand soil sampling tools were used in this study because it was not 
possible to use a Kelley (1) or similar type mounted soil core sampler 
while the corn was growing. The hand type soil sampling tools used were 
a standard bucket auger' and a Pit-O-Matic core sampler 4 of approxi- 
mately the same size, 5.5 cm (2i% in) inside diameter. The cutting edge 
diameter of the bucket auger was 7.0 cm (2% in), and that of the core 
sampler was 5.2 cm (2i 1 6 in). 

A Crosby silt loam (Aerie Ochraqualff) and a Ragsdale silty clay 
loam (Typic Argiaquoll) adjacent to each other in the same field of 
William Franklin, Jamestown, Boone County, Indiana, were used for 
this study. The soils were described by D. P. Franzmeier and P. W. 
Harlan (personal communication). Some properties of these soils are 



1 Journal Paper No. 3875, Purdue University Agr. Exp. Sta. 

J The author acknowledges the help of C. D. Raper, Richard Fletcher, Ethel Tudor, 
and Enola Ruff in conducting this research. 

:! Manufactured by Art's Machine Shop, American Falls, Idaho. 

4 Manufactured by the Soil Testing Company, Smithville, Tennessee. 

401 



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Soil Science 403 

listed in Table 1. Ragsdale is higher in organic matter, more alkaline in 
subsurface horizons to parent material, and has a higher strong acid 
phosphorus test value in the B horizon to 56 cm (22 in) than does the 
Crosby soil. Ragsdale is found in lower areas to which runoff comes 
from the higher Crosby soil. 

Soil and root samples were taken between August 9 and August 14, 
1968, after midsilk date of this corn crop, while the ears were still 
filling and while the brace roots were still growing. The variety was 
Northrup King 610 planted April 27. For the soil and root sampling and 
for yield determinations, a nested design suggested by W. E. Nyquist 
(personal communication) was used. Soils were main treatments. Five 
replications or subsampling areas approximately 15.3 m (50 ft) apart, 
located in the center and on the points of the compass, were used in each 
soil type. Within each sublocation, depths were sampled to 152 cm (60 
in) where possible, in 15 cm (6 in) vertical increments starting at the 
surface. At each sublocation two samplers, the standard bucket auger 
and the Pit-O-Matic core sampler, were used, one on each side of a 
corn plant located 18 cm (7.1 in) from another corn plant in the row and 
96.5 cm (38 in) from plants in adjoining rows. Surface increments in 
the row touched the stalk. 

Soil and root subsamples for each 15 cm increment were put in 
paper bags and air dried at room temperature. Prior to separating roots 
from soil, the samples were dried at 38° C (100° F) for 24 hours and 
weighed. After the dried sample was broken up into smaller pieces with 
a mortar and pestle, the roots were separated from the soil by the 
method of Raper (5). This consisted of a wet screening procedure with 
oscillation in a saturated calgonite solution. Tweezers and flotation in 
water were used where soil particles containing roots failed to break 
down. Roots were placed in steel cans and dried at 38° C for 24 hours, 
weighed, and placed in labeled plastic bags for reference. 

Corn grain yields were taken in 1968 in the same five sub-locations 
on each soil where the soil and root samples were taken. Yields are 
reported with 15.5% moisture in the grain. 

Weights of roots, soil plus roots (roots were less than 1% of the 
total weight), and yields were subjected to analyses of variance and F 
tests of significance. 

Results and Discussion 

Weight of roots per 15 cm depth of soil sampled decreased highly 
significantly from the 0-15 cm depth to the 15-30 cm depth. The average 
root weight of the 0-15 cm depth was 1.33 grams or 67% of the total 
root weight to 152 cm (60 in) depth and that of the 15-30 cm depth was 
0.22 grams or 11% of the total root weight to 152 cm (60 in) depth. 
Even though average root weights declined as soil depth increased 
below 30 cm depth, these differences in root weights were not significant 
at 19 to 1 or greater odds. These averages are shown in Table 2 for all 
depths sampled, both soils, and both soil sampling tools. Part of the 



404 



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Soil Science 405 

wide ranges in weight per subsample in the 0-15 cm and 15-30 cm 
samples can be attributed to presence of brace roots. There were brace 
roots in some samples and not in others. However, after drying it was 
impossible to determine rapidly what was a brace root and what was 
not. Foth (4) found that brace root growth increased total root weight 
of corn nearly 50% between the 67th and 80th days after planting. 
Harvests in this experiment were made between the 104th and 109th 
days after planting — probably before total brace root growth was 
completed. 

Another possible explanation in the wide range in root weights is 
related to the method of sampling. Samples of soil and roots were taken 
out of the hole after each successive 15 cm depth was reached. This 
allowed extra soil and roots which fell into the hole to be brought up. 
Coefficients of variation of the dry weights of soil samples (with very 
small weights of roots in them) varied from 14% up to 18%. However, 
coefficients of variation of corn root weights found in these soil samples 
ranged from a low of 62 % to a high of 125%. These are extremely high 
coefficients of variation. When variation in weights of soil samples and 
root samples is compared, it is found that root weights were four to 
seven times as variable as weights of soil samples. This indicates that 
corn roots were not uniformly distributed in the soils sampled (Table 2). 

Root weights from the bucket auger averaged 27% heavier than 
those from the core sampler in samples taken to 91 cm (36 in). This 
difference was significant at 4 to 1 odds and was expected because of 
the wider cutting edge of the bucket auger. The cutting diameter of the 
bucket auger was 35% wider than that of the core sampler. However, 
the average weight of soil and roots per 15 cm increment taken by the 
bucket auger was 699 g. This compared to 465 g for the core sampler 
increment or 50% more. The difference of 234 g was highly significant. 
Root samples taken by the bucket auger were macerated and harder to 
separate than those taken with the core sampler. However, the bucket 
auger was able to by-pass stones and sample to a greater depth in one 
sublocation than was the core sampler in Crosby soil. Gravel in pockets 
probably laid down by water prevented sampling any deeper than 122 cm 
(48 in) with the core sampler in Crosby soil. In the one sublocation 
where it was possible to sample to the desired depth (152 cm or 60 in) 
on the Crosby soil, root weights were less than the averages found in 
Ragsdale soil at the same depths. However, it was possible to sample 
all depths to 152 cm at all sublocations of the Ragsdale soil. There were 
no significant differences between the two soils in root weights found to 
91 cm. Below this, other sampling procedures are needed to determine 
whether or not Ragsdale has more roots at greater depths than has 
Crosby. The methods used in this study to sample corn roots in Crosby 
soil below 91 cm were not satisfactory. 

Yields of corn grain averaged 10,057 kg per ha (160 bu per A) on 
the Ragsdale soil in 1968. On the Crosby soil, yields were 8809 kg per 
ha (140 bu per A). The difference in yields of 1248 kg per ha (20 bu 
per A) was significant at 19 to 1 odds. Since the rainfall of 2.77 cm 



406 Indiana Academy of Science 

(1.09 in) in July and 7.29 cm (2.87 in) in August was deficient, the less 
gravelly Ragsdale soil must have supplied more water to the crop — ■ 
probably because of deeper root penetration. 

Summary 

Roots were sampled from five sublocations of both a Crosby silt 
loam and a Ragsdale silty clay loam in the same field. Two types of 
sampling tools were used to obtain vertical soil cores. Subsamples were 
taken by 15 cm (6 in) increments from the surface down to 152 cm 
(60 in) where possible. The roots were separated from the soil. Highly 
significant differences were found between root weights from the 0-15 cm 
(0-6 in) depth subsamples and root weights from subsamples taken 
below that. About 67 r /<- of the total weight of roots to a depth of 152 cm 
was found in the 0-15 cm subsamples. Weight of roots rapidly declined 
from the 0-15 cm depth to the 137-152 cm (54-60 in) depth. Gravel in 
the Crosby profiles made it impossible to sample deeper than 107 cm 
(42 in) in 4 of the 5 sublocations. The standard bucket auger with 
wider effective cutting diameter took more soil and with it, 27% more 
roots than did the Pit-O-Matic core sampler. The bucket auger macerated 
the roots while the core sampler did not. The very wide range in root 
weights taken in subsamples to 30 cm depth (12 in) was partially 
explained by the presence of brace roots in some subsamples and not in 
others. It was shown that weight of roots in soil subsamples was four to 
seven times as variable as the weights of the soil from which they 
came. Grain yields of corn in 1968 of 10057 kg per ha (160 bu per A) 
on the Ragsdale soil were significantly greater than the 8809 kg per ha 
(140 bu per A) on the Crosby soil. Since July and August were relatively 
dry, the less gravelly Ragsdale soil apparently supplied more available 
water — probably because of deeper root penetration. 



Literature Cited 

1. Fbhrenbacher, J. B., and J. D. Alexander. 1955. A method of studying corn root 
distribution using a soil-sampling machine and a shaker-type washer. Agron. J. 
47:468-472. 

2. Fehrenbacher, J. B., P. R. Johnson, R. T. Odell, and P. E. Johnson. 1960. Root 
penetration and development of some farm crops as related to soil physical and 
chemical properties. Transactions of the 7th Inter. Congress of Soil Science, Madison, 
Wise, U.S.A., 1960. Vol. 111:243-252. 

3. Fehrenbacher, J. B., B. W. Ray, and J. D. Alexander. 1967. Root development of 
corn, soybeans, wheat, and meadow in some contrasting Illinois soils. Illinois Re- 
search, University of Illinois Agricultural Experiment Station, Spring 1967:3-5. 

4. Foth, H. D. 1962. Root and Top Growth of Corn. Agron. J. 54 :49-52. 

5. Raper, Charles David, Jr. 1970. Significance of Root Characteristics in Nutrient 
Uptake and Growth of Soybeans. Ph.D. Thesis, Purdue University. 



The Effect of Rainfall Energy on Water Infiltration into Soils 1 

J. V. Mannering and D. Wiersma, Purdue University 



Abstract 

Simulated rain was applied in the field to seven Indiana soils ranging in texture 
from a sand to a silty clay. Each site was divided so that one half of the plot was bare 
fallow while the other half was protected by a layer of screenwire suspended 10 cm above 
the surface. A comparison of the infiltration characteristics of these soils under these two 
conditions when exposed to high energy rain showed rainfall energy to be the principal 
causative factor in surface sealing. The magniture of these differences was greatly 
influenced by soil texture, however, medium textured soils were affected most severely. 
After 30 minutes of rainfall, infiltration rates on medium textured, bare soils were 20 to 
30% of those on protected soils. 

The formation of a layer at the soil surface that reduces water 
intake has been recognized for over a century (1). The principal factor 
responsible for the formation of this surface seal has been shown to be 
the impact of raindrops on the soil surface (5). Duley and Kelly (2), 
for example, showed that when the surface was protected by cover, a 
broad range of infiltration rates occurred between different soil types. 
The differences diminished, however, when these same soils were bare 
and subjected to rain. Other workers (3,7) have reported permeabilities 
for surface seals of 1/5 to 1/2000 of those of underlying materials. In 
the solution of a mathematical model, Swartzendruber (6) showed that 
the conductivity of the least permeable layer does not of itself control 
flow but is dependent on the physical characteristics of both the least 
permeable layer and the other material in the system. 

Although much effort has been directed to understanding water 
intake into sealed soil surfaces, the magnitude of changes in infiltration 
rates of Indiana soils brought about by rainfall energy remains 
unsolved. 

It is the purpose of this study to show the accumulative influence 
of rainfall energy on infiltration rates for a wide range of soil 
textures. 

Methods and Procedures 

Infiltration rates were determined by using the rainfall simulator 
described by Meyer and McCune (4). Two storms of 1-hour duration at 
an intensity of 7.0 ± 0.5 cm per hr were applied on successive days to 
soils that had been maintained in a fallow condition for several months. 
Two treatments or conditions were used to measure the influence of 
high energy rainfall on infiltration rates of fallow (bare) soils: 1) 
rainfall of high kinetic energy was applied to a bare, unprotected soil, 

1 Contribution from Purdue University Agronomy Department, Lafayette, Indiana. 
Published with the approval of the Director of the Purdue University Agricultural Experi- 
ment Station as Paper No. 3894. The authors wish to acknowledge the Soil and Water 
Conservation Research Division, Agricultural Research Service, USDA for their con- 
tribution in this study. 

407 



408 Indiana Academy of Science 

and 2) rainfall with low kinetic energy was applied to this same soil. 
This was accomplished in the following manner. Prior to the application 
of simulated rainfall, one plot (12 x 35 feet in size) was covered with 
a double-layer of 18 x 14 mesh screenwire. This screenwire was sus- 
pended 10 cm above the soil surface to reduce drop size and velocity 
(kinetic energy) of the rain without obstructing overland flow or 
runoff. A second adjacent plot was left bare and unprotected. 

This study was conducted on seven Indiana soils (Table 1) ranging 
in texture from a sand to a silty clay. Some of the relevant soil 
properties are shown in Table 1. 

Table 1. Properties of soils tested. 











Organic 


Aggre- 




Sand 


Silt 


Clay 


Matter 


gation 




(%) 


(%) 


(%) 


(%) 


Index 


Oakville sand 


94 


4 


2 


0.5 


0.065 


Fox gravelly sand loam 


80 


12 


8 


1.1 


0.181 


Warsaw sandy loam 


62 


25 


13 


3.3 


0.408 


Fox silt loam 


22 


57 


21 


1.3 


0.494 


Zanesville silt loam 





72 


10 


1.3 


0.160 


Cincinnati silt loam 





72 


19 


1.3 


0.205 


Markland silty clay 


4 


55 


41 


3.0 


0.986 



The kinetic energy of the rainfall from the simulator is approxi- 
mately 800 ft tons per acre inch (4). The kinetic energy of the same 
intensity rainfall falling through a double-layer of screenwire is not 
known. However, it has been observed by means of high-speed pho- 
tography that the screenwire effectively disperses large drops. As a 
consequence, the rainfall energy of single drops is expended over a 
much larger area and the velocity of a sizable portion of the falling 
drops is reduced. Both of these factors would greatly reduce the 
kinetic energy of the falling rain and, therefore, its dispersive power in 
the formation of surface seals. 

Results and Discussion 

The effects of the rainfall energy on infiltration rates of the various 
soils over a 2-hour period are shown in Figures 1 through 7. 

The protective screenwire is shown to have very little influence on 
the infiltration on the Oakville sand during the first 60-minute storm 
(Fig. 1) since on both the protected and unprotected plots essentially 
all of the rain applied entered the soil. This was true for the protected 
plot throughout the 2-hour storm. However, the infiltration on the bare 
plot was reduced to 70 % of that on the protected plot at the end of 
2 hours. 



Soil Science 



401) 



More evidence of surface sealing- occurred on the Fox gravelly 
sandy loam (Fig. 2) when the protective cover permitted essentially all 
water applied during the 2-hour storm to enter the soil. On the other 
hand, infiltration rates on the unprotected plots were only 55% and 32%, 
respectively, of that on the protected plots at the end of 1 and 2 hours. 
The formation of the surface seal on the bare plot had largely occurred 




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410 Indiana Academy of Science 

after only 20 minutes of rain as evidenced by the rapid drop in 
infiltration rate. 

Figure 3 shows the effect of rainfall energy on the infiltration rate 
of a Warsaw sandy loam. Again, the infiltration rate remained high on 
the protected plot throughout the 2-hour test while the infiltration rate 
on the sealed plot had decreased 48% and 24%, respectively, at the end 
of 1 and 2 hours. Although the soils studied in Figures 2 and 3 are both 
sandy loam in texture, the Warsaw soil had approximately twice the 
silt + clay content of the Fox soil which probably accounted for the 
appreciably lower infiltration rate at the end of the 2-hour storm. 

After only 10 minutes of rain, a seal had formed on the Fox silt 
loam soil and little further reduction in infiltration rate occurred after 
30 minutes of rain during the first 1-hour storm (Fig. 4). Infiltration 
rates on the bare plots were 35% and 80%, respectively, of those on the 
protected plots after 1 and 2 hours. The swelling of the clay present is 
thought to be responsible for the rapid decrease in infiltration on the 
protected plot during the second-day test. 

Figures 5 and 6 illustrate the influence of rainfall energy on the 
infiltration rates of soil high in silt. When not protected these two soils, 
Zanesville and Cincinnati, containing 72% silt, both exhibited a marked 
reduction in infiltration after only 15 minutes of rain. Further reduc- 
tions in infiltration rate after 30 minutes of rain were relatively small. 
Infiltration rates on the bare plots after the first hour of rain were 
approximately 35 % of those on protected plots for both soils. After 2 
hours, the differences between rates on the bare and protected plots 
were greater on the Zanesville (61%) than on the Cincinnati (34%). 
The reduced infiltration with time on protected plots indicate that 
either internal soil properties are controlling infiltration rate or that 
the rain falling through the screenwire contained sufficient energy to 
produce seals on these particular soils. 

Rainfall energy did not result in a major reduction in infiltration 
on the Markland silty clay (Fig. 7). After 60 minutes of rain, the rate 
on the bare plot was still 60% of that on the protected plot and there 
was essentially no difference between treatments at the end of 2 hours. 
The high level of water stable aggregation resulting from high clay 
content and relatively high organic matter levels prevented the forma- 
tion of a severe surface seal on this soil. 

These results show that when exposed to high energy rainfall, all 
of the soils tested are subject to surface sealing, thus reduced water 
infiltration. The magnitudes of these differences vary greatly among 
soils and are to a great extent controlled by soil texture and structure. 
The soils most seriously affected are the medium textured soils where 
infiltration rates into exposed soils can drop to as low as Ys those of 
protected soils after only a few minutes of rain. Organic matter and 
clay contents when sufficiently high to produce stable structure greatly 
reduce the soil sealing effect of rainfall energy. 

The realization that the rate of water infiltration into soils can be 
reduced as much as four or five-fold because of surface sealing, is 
essential in many forms of hydrological planning. 



Soil Science 



411 







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412 Indiana Academy of Science 

Literature Cited 

1. Baver, L. D. 1956. Soil Physics, Third Edition, John Wiley and Sons, Inc., New York. 

2. Duley, F. L., and L. L. Kelly. 1939. Effect of soil type, slope and surface conditions 
on intake of water. Nebr. A.E.S. Res. Bull. 112:1-16. 

3. McIntyre, D. S. 1958. Permeability measurements of soil crusts formed by raindrop 
impact. Soil Sci. 85:185-189. 

4. Meyer, L. Donald, and D. L. McCune. 1958. Rainfall simulator for runoff plots. Agr. 
Eng. 39 :644-648. 

5. Neal, J. H., and L. D. Baver. 1937. Measuring the impact of raindrops. J. Amer. 
Soc. Agron. 29 :708-709. 

6. Swartzendruber, D. 1960. "Water flow through a soil profile as affected by the least 
permeable layer. J. Geophys. Res. 65(12) :4037-4042. 

7. Tackett, J. L., and R. W. Pearson. 1964. Some characteristics of soil crusts formed 
by simulated rainfall. Soil Sci. 99 :407-413. 



Effects of Organic Matter on the Multispectral Properties of Soils 1 

M. F. Baumgardner, S. Kristof, C. J. Johannsen, and A. Zachary, 

Purdue University 



Abstract 

The use of data obtained from an airborne optical-mehanical scanner in determining 
the organic matter content of surface soils is presented. Multispectral data covering the 
electromagnetic spectrum from 0.32 to 14 microns was recorded on magnetic tape in 
analog form. After conversion to digital form, the data were analyzed by computer to 
provide pictorial printouts of soils areas. The computer was trained to recognize the 
spectral responses of different levels of organic matter of surface soils and produce maps 
showing the locations of organic matter content. The computer maps were compared 
with results from approximately 200 surface soil samples analyzed for organic matter. 
These samples were collected in a 25-hectare field in the center of a flight line in Tippe- 
canoe County, Indiana. 



Introduction 

For many years the soil scientist has been using" a color designa- 
tion as part of his description of the various horizons of the soil profile. 
The color designation commonly used in many countries in the world is 
the Munsell notation system (5). 

From a certain color designation the soil scientist can make a 
number of inferences about other properties of the soil. He relates 
variability of the soil color to: 1) organic matter content, 2) the 
presence or absence of oxidized or reduced iron compounds, 3) the 
internal drainage characteristics of the soil, and even to a certain 
extent 4) the potential productivity of the soil. 

Attempts have been made to quantify the soil color measurements 
and to relate these measurements quantitatively with other soil 
properties. Many of these attempts have met with little success because 
of the difficulties encountered in making quantitative color measure- 
ments. Variations of soil color are found when measuring undisturbed 
soil samples versus disturbed samples. Surface roughness and variations 
in soil moisture content also affect soil color (2). 

During the past 4 years research has been conducted at the Labora- 
tory for Applications of Remote Sensing (LARS) at Purdue Uni- 
versity to develop techniques and instrumentation for measuring* and 
characterizing earth surface features with remote sensing devices from 
aerospace platforms. The use of an airborne optical-mechanical scanner 
for obtaining earth resources data and the use of computer techniques 
for reducing the data have been described in numerous papers (2, 3, 
6,7). 



1 Agricultural Experiment Station Journal No. 3939. This research was supported 
jointly by the U. S. Department of Agriculture and the National Aeronautics and Space 
Administration, and conducted at the Laboratory for Applications of Remote Sensing. 

413 



414 



Indiana Academy of Science 



Kristof (4) and Baumgardner et al. (2) describe the use of these 
techniques in the automatic identification, separation, and mapping' of 
soil categories based on the differences of spectral properties of the 
surface soils. 

The initial results of a study of the effects of soil organic matter 
on the spectral properties of soils are reported in this paper. Data 
essential to this study were obtained with an airplane containing an 
optical-mechanical scanner flown by the University of Michigan. The 
data were analyzed by computer techniques at LARS. 



Materials and Methods 

General soils studies have been conducted on data obtained from 
flight lines flown in Tippecanoe County, Indiana, during May, 1969. 

TIPPECANOE COUNTY, INDIANA 
21 22 23 24 



25 ttrfcR 




Figure 1. FUghtlincs fioivn for scanner data collection and location of Soil Area D. 



Soil Science 



415 



Sample Locations - Dieterle Farm 




it 



Figure 2. Soil sampling locations on Test Site D. 



f" C ° U " t ?' heS m a transitional zone between prairie and forest 
so s. In the northwestern part of the County, soils were formed under 
tall prarne grasses. Soils of the southeastern portion were formed 
under deciduous hardwood forest. M 

A 25-heetare field was selected for more intensive soil studies This 
field shown as test site D in Figure 1, is located in the native fore 
yegetat.on area and has surface soil patterns typical of the Hapludalfs 
(gray-brown podzols). The field is a portion of the SEV 4 , Sec. 6 T21N 
R3W, Tippecanoe County, Indiana. 



416 Indiana Academy of Science 

Scanner and photographic data were obtained on the flight lines 
shown in Figure 1 on May 26, 1969, between 1100 and 1300 hours. 
Ground truth data showed that test site D had been plowed and disked 
in preparation for planting corn and soybeans. Since there was no sur- 
face vegetative cover, the soil patterns were clearly defined and easily 
discernible by visual means. 

Within a few days after the scanner flight, a sampling grid plan 
was drawn (Fig. 2) and 1-kg surface soil samples were obtained at 
32m intervals. A total of 197 samples was taken from the surface to a 
maximum depth of 2 cm. 

One of the objectives of this research was to study the relation- 
ship between the organic matter content of surface soils and the spec- 
tral response patterns in the spectral range from 0.40ft to 2.6/x. Ac- 
cordingly, organic matter determinations were made on each of the 
197 samples by the Walkley and Black method (8). 

It was theorized that if the correlation between these two variables 
was good, then the scanner data could be used in the rapid preparation 
of computer printouts or maps of fields showing different levels of 
organic matter content in the surface soils. 

The scanner data used in this study was obtained at an altitude of 
approximately 1300 m (4000 feet). The scanner data were digitized at 
an interval such that each sampling point on one scan line would repre- 
sent the average energy of a specific area on the ground. At the altitude 
of 1300 m, each sampling point or resolution element represented ap- 
proximately 16 to 24 m 3 (150 to 200 ft 2 ). 

To study the correlation between organic matter content and spec- 
tral response, it was necessary to obtain a quantitative spectral value 
in each wavelength channel which was representative of the area from 

Table 1. Wavelength ranges in 12 channels of the University of 
Michigan optical-mechanical scanner. 

Channel Number Wave length Range (microns) 



1 0.40-0.44 

2 0.46-0.48 

3 0.52-0.55 

4 0.55-0.58 

5 0.58-0.62 

6 0.62-0.66 

7 0.66-0.72 

8 0.72-0.80 

9 0.80-1.00 

10 1.00-1.40 

11 1.50-1.80 

12 2.00-2.60 



Soil Science 417 

which each surface sample of soil was obtained. On a gray-scale com- 
puter printout of the entire field, four resolution elements were selected 
to represent each soil sample location. The average radiance values of 
the 4 resolution elements for each surface sample in each of 12 wave- 
length channels were used to plot against the organic matter content. 
Table 1 shows the wavelength frequencies measured in each of the 12 
channels used in this study. 

Alexander (1) developed a color chart for estimating organic 
matter in mineral soils in Illinois. Correlations were made of soil 
colors (Munsell notations) with laboratory analyses for organic matter 
determined from a large number of Illinois soils. 

Table 2 indicates the five levels of organic matter which Alexander 
uses in his color chart. Soils of the test site, which were used for 
analysis with the multispectral scanner data, contained similar amounts 
of organic matter. 

Table 2. Levels of soil organic matter content determined in Illinois (1). 

Average (%) Range (%) 

5 3.5-7.0 

3.5 2.5-4.0 

2.5 2.0-3.0 

2 1.5-2.5 

1.5 1.0-2.0 



In this study the desired task was to obtain a computer printout 
(or map) of the test site showing the locations of soils having five 
levels of organic matter similar to the Illinois study (1). The organic 
matter content and corresponding spectral responses of 12 wavelength 
bands for each soil sample location were used to train the computer. The 
samples were divided into five levels of organic matter content (Table 
3). The computer classified each resolution element of each scan line 
in the test site using a pattern recognition technique which used the 
data provided by the training samples. Less than V± of the area in the 
test site was used to train the computer. 

Table 3. Number of samples in each of five levels of soil organic 
matter used for training the computer. 



Percent 


Number of 


Organic Matter 


Samples 


>3.5 


46 


2.5-3.5 


37 


2.0-2.5 


18 


1.5-2.0 


63 


0-1.5 


46 



418 



Indiana Academy of Science 



Results and Discussion 

To provide a laboratory model of the soil patterns on the test site 
area a soil mosaic was constructed with the use of portions of each of 
the 197 surface soil samples. Figure 3 contains a photograph of this 
mosaic and provides a good general representation of the soils pattern 



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Figure 3. General soil patterns of Test Site D. 



Soil Science 



419 




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Figure 4. Photograph of computer printout of general soil patterns at Test Site D. 



420 



Indiana Academy of Science 



of the field. A photograph of a computer printout (Fig. 4b) showing 
five different levels of organic matter in the field is shown for compari- 
son with the soil mosaic. This organic matter map compares very well 
with the color patterns observed in the soil mosaic. The depressional 
soils which have the greatest amount of organic matter lie near the 
broad drainage ditch. In the lower right hand area of the illustration a 
high organic matter content is also indicated. This is a slight depression 
and natural drainageway. 



190 



170 



150 



130 



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70 



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Organic Matter Content (Percent) 



5.7 6.2 



P^igure 5. Correlation between average radiance level (relative response) and organic 
matter content for 197 samples from Test Site D. 



The test site contains undulating land. The lighter areas to the 
left and right of the center of the illustration represent sloping topog- 
raphy. In some cases, these slopes are as much as 3 to 4 m higher in 
elevation than the high organic matter soils along the drainageways. 
It was observed that with more detailed examination of the organic 
matter classes, severely eroded and moderately eroded spots in the 
field could be identified on the computer printouts. In these instances, 
erosion has been sufficiently severe that subsoil was exposed. 

Average radiance levels of the 12 wavelength channels were 
plotted against soil organic matter content of each sample location. 
Most of the plottings or correlations were quite similar. Figure 5 pre- 
sents data for channel 6 (0.62-0.66*0 . Linear regression analysis gave 



Soil Science 421 

an r value of — 0.74. However, the plotted data seem to indicate that 
perhaps a linear relationship may not be valid over the range of organic 
matter used. It appears that above 2.0 or 2.5% organic matter, there 
may be a linear correlation with a much higher r value. Below 2.0% 
organic matter, the curve becomes much steeper. 

It appears that organic matter plays a dominant role in bestowing 
spectral properties upon soils when the organic matter content exceeds 
2.0%. As the organic matter drops below 2% it becomes less effective 
in masking out the effects of other soil constitutents such as iron or 
manganese on spectral response of soils. 

Training samples which are used for the test site were also used 
for obtaining a 5-level organic matter map of the entire north-south 
flight line across Tippecanoe County. The area covered was approxi- 
mately 24 miles long and 1 mile wide. The results look very promising. 
More research is necessary, however, to check the accuracy of the 
results and to determine the proper methods for determining organic 
matter levels with the use of the computer. Size of the training samples 
is also being evaluated. 

Summary and Conclusions 

Results of this research have well established the feasibility of 
using multispectral scanner data and computer analysis to prepare 
maps which outline surface soils with varying levels of organic matter. 
Surface soil samples were obtained from a test site area and organic 
matter content was determined. Linear regression of organic matter 
content and wavelength response gave a r value of — 0.74. 

More research is needed to define more precisely the effects of 
and interactions between organic matter, iron, and aluminum com- 
pounds, surface roughness, and moisture as these factors affect the 
spectral properties of soils. 

An operational capability to map five or more levels of soil organic 
matter in a rapid manner would provide a very useful tool to the soil 
surveyor, the urban and regional planner, the agricultural chemical 
industry, the drainage engineer, the farm manager, and many others. 



Literature Cited 

1. Alexander, J. D. 1968. Color chart for estimating organic matter in mineral soils in 

Illinois. University of Illinois, Urbana. 

2. Baumgardner, M. F., C. J. Johannsen, and S. J. Kristof. In press. Multispectral 
studies of soils and clay minerals. Proc. German Soil Sci. Soc. Meetings. Hannover, 
W. Germany. September, 1969. 

3. Johannsen, C. J., and M. F. Baumgardner. 1968. Remote Sensing for planning 
resource conservation. Proc. Soil Conserv. Soc. of Amer. Athen, Ga., August, 1968, p. 
149-155. 

4. Kristof, S. J. In press. Preliminary Multispectral Studies of Soils. J. Soil and Water 
Conserv. 



422 Indiana Academy of Science 

5. Munsell, A. H. 1947. A color notation. Ed. 10. Munsell Color Company, Baltimore. 

6. Laboratory for Agricultural Remote Sensing. Remote Multispectral Sensing in Agricul- 
ture. 1967. Vol. 2 (Annual Report). Res. Bui. No. 832, Purdue University. 

7. Laboratory for Agricultural Remote Sensing. Remote Multispectral Sensing in Agricul- 
ture. 1968. Vol. 3 (Annual Report). Res. Bui. No. 844, Purdue University. 

8. Walkley, A., and I. A. Black. 1934. An examination of the Degtjareff method for 
determining soil organic matter and a proposed modification of the chromic acid titra- 
tion method. Soil Sci. 37 :29-38. 



A Two- Year Study of Bacterial Populations in Indiana 
Farm Pond Waters 1 

L. B. Hughes and H. W. Reuszer, Purdue University 



Abstract 

A two-year study of bacterial populations in farm pond waters was conducted on 
three farm ponds on the Southern Indiana Purdue Agricultural Center. Samples of water 
from the surface and near the bottom were taken approximately once per month from 
each pond. Bacterial numbers were determined by colony counts. Great differences in 
bacterial numbers were found between ponds and large seasonal variations in bacterial 
numbers were found within ponds. Maximum bacterial numbers occurred in late spring 
or early summer in all three ponds. Consistently higher numbers of bacteria were 
present in Pond C and lowest numbers were present in Pond B. Water temperatures 
were quite similar in all three ponds, although a significant correlation between water 
temperature and bacterial numbers was found only in Pond C. Organic carbon content 
was consistently highest in Pond C and lowest in Pond B. Significant positive correlation 
coefficients between organic carbon and baterial numbers were found in Ponds A and B, 
but a significant negative correlation was found in Pond C. 



Introduction 

Farm ponds have many uses in our day including serving as sources 
of recreation, sport and commercial fishing, livestock water, and even 
water for human consumption and general household use. Consequently, 
the chemical and biological nature of farm pond waters is of extreme 
importance. A thorough search of the literature revealed few micro- 
biological studies in farm pond waters. The purpose of this study was 
to determine the changes in numbers of bacteria in farm pond waters 
over a 2-year period and to investigate some of the factors possibly 
influencing the bacterial numbers. 

Literature 

Many workers have reported large fluctuations in bacterial num- 
bers in bodies of water. Fred et al. (1) and Snow and Fred (4) found 
large fluctuations of bacterial numbers in Lake Mendota, Wisconsin, 
in studies covering 4 years and extreme fluctuations could not always 
be explained (1). In a brief study of Flathead Lake, Montana, Graham 
and Young (2) found maximum bacterial numbers below 8,000 per ml. 
Lower numbers of bacteria were found in the surface water than in 
water below the surface. Stark and McCoy (5) reported striking dif- 
ferences in bacterial numbers in the surface water at different loca- 
tions on the same lake. 

Irwin and Claffey (3) found wide variations in numbers of bac- 
teria in the waters of 20 ponds in Oklahoma with fewer bacteria found 
in ponds with higher turbidity. Wilson et al. (7) reported 5,000 to 



1 Journal Paper No. 3889, Purdue University Agricultural Experiment Station, 
Department of Agronomy, Lafayette, Indiana. This study was supported in part by the 
Office of Water Research, U. S. Department of the Interior. 

423 



424 Indiana Academy of Science 

30,000 bacteria per ml in 6 West Virginia ponds. Little differences in 
numbers of bacteria were detected at different depths. Water tempera- 
ture and pH showed no correlation to the bacterial numbers. 

Procedures 

The three farm ponds studied were constructed in 1953 or 1954 
on the Southern Indiana Purdue Agricultural Center. The ponds were 
arbitrarily labeled A, B, and C. Pond A was the largest with a surface 
area of 0.96 acre, a maximum depth of 12 feet, and a watershed area 
of 7 acres. Substantial aquatic plants in the edge of Pond A con- 
tributed organic matter to the water. Pond B had a surface area of 
0.66 acre, a maximum depth of 12 feet and a watershed area of 2 
acres. Very little aquatic plant growth occurred in this pond. Pond C 
had a surface area of 0.30 acre, a maximum depth of 15 feet and a 
watershed area of 3 acres. Dense growth of aquatic plants extended 
several feet into the edge of the water. Substantial algal growth was 
noted in Ponds A and C in late spring and early summer, but little 
algal growth was noticed in Pond B. A mixture of alfalfa and orchard 
grass was the common forage crop growing on the three watersheds and 
provided good protection against erosion of the soil. 

Samples of water were taken from a raft at the surface and 6 
inches from the bottom at 4 different locations on each pond approxi- 
mately once per month over a 2-year period (April, 1967 to March, 
1969). Samples of water were immediately placed in ice and trans- 
ported back to the laboratory (4 hours travel time). The water samples 
were then stored overnight at 5° C and plated the following day on an 
agar medium containing 1.0 gm glucose, 1.0 gm peptone, 0.5 gm yeast 
extract, 0.25 gm K 2 HPO,, 12.0 gm agar, and 1,000 ml deionized water. 
Bacterial numbers were determined by colony counts made after 14 
days incubation in the dark. 

Water temperatures were obtained four times daily by E. J. Monke 
and P. R. Goodrich of the Purdue University Agricultural Engineering 
Department using thermocouples at various depths. Data used in this 
paper are the means of the four temperatures recorded on the day of 
sampling. Water temperatures were available only from April to De- 
cember, 1967. Aliquots of water were evaporated and organic carbon 
was determined using the manometric procedure described by Van Slyke 
and Folch (6). Data on pH, nitrate concentration, and water turbidity 
were obtained from the Indiana State Board of Health. 

Results and Discussion 

The factors possibly influencing the bacterial numbers that were 
studied include organic matter content, water temperature and to a 
lesser extent, pH, nitrate concentration, and turbidity of the water. 

Nitrate concentration, pH, and water turbidity of the three ponds 
are shown in Table 1. The highest average pH found in Pond B would 
seem to be farther from the optimum pH for maximum bacterial 



Soil Science 425 

growth than the pH in either Ponds A or C. Water turbidity was also 
highest in Pond B. The small variations in nitrate concentrations would 
not be expected to have significant effect on bacterial numbers. 



Table 1. Turbidity, pH and nitrate concentration of the pond water, 

at different times." 





April 


August 


December 


April 


July 


Avg. 




1967 


1967 


1967 


1968 


1968 










pH 








Pond A 


7.4 


6.8 


8.0 


7.1 


7.0 


7.3 


Pond B 


7.4 


8.0 


7.4 


7.7 


7.8 


7.7 


Pond C 


7.3 


7.1 


7.4 


7.6 


7.3 


7.3 






Nitrate Concentration (ppm) 






Pond A 


0.1 


0.1 


0.3 


0.1 


0.1 


0.14 


Pond B 


0.1 


0.2 


0.2 


0.1 


0.1 


0.14 


Pond C 


0.1 


0.2 


0.3 
Turbidity 


0.1 


0.2 


0.18 


Pond A 


0.3 


0.1 


10 


2 


0.3 


2.5 


Pond B 


15 


0.1 


3 


15 


3 


7.2 


Pond C 


0.7 


0.2 


3 


1 


0.0 


1.1 



2 The authors express gratitude to the Indiana State Board of Health for use of 
these data. 



Water temperatures of the 3 ponds for a 9-month period are shown 
in Figure 1. Surface water temperatures were quite similar in all 3 
ponds, reaching a maximum near 80° F in June, and a minimum near 
40° F in December. The maximum bottom temperatures in Ponds A and 
B were above 70 °F in August. Cooling began one month earlier in Pond 
C. Minimum bottom water temperatures occurred in December and 
were about equal to surface temperatures at that time. 

Seasonal variations in organic carbon content are shown in Figure 
2. The highest organic carbon content generally occurred in late spring 
or early summer in both years. Pond B had the least seasonal variation 
of organic matter content, with a range of 7.1 to 14.8 mg of organic 
carbon per liter of water with similar quantities of organic carbon in 
the surface water and in the water near the bottom. Pond A showed 
larger seasonal variation of organic matter at both depths with organic 
carbon quantities ranging from 8.0 to 24.0 mg per liter. Higher organic 
matter content generally was prevalent near the bottom than at the 
surface in Pond A. Both depths of Pond C contained the highest or- 
ganic matter content and the largest seasonal variation, ranging from 
11.1 to 29.2 mg of organic carbon per liter of water. 



426 



Indiana Academy of Science 



80 








POND C 
• "Surface Water 






/°^ 




~~\ °-—o Water 6" From Bottom 


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/ 


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POND A 
—•Surface Water 
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'A M 
1967 



J J A S 

TIME OF SAMPLING 



Figure 1. Seasonal variations in water temperature for a nine-month period. (The 
authors express gratitude to P. R. Goodrich and E. J. Monke for use of these data.) 



Bacterial numbers are given in Table 2. Seasonal variations in 
bacterial numbers are shown in Figure 3. The highest numbers of 
bacteria generally were present in late spring or early summer in both 
years of the study. High numbers of bacteria seemed associated with 
the active growth period of aquatic vascular plants rather than the 
autumn period of plant death and decay. The role of algae with respect 
to numbers of bacteria was not clear. Pond A had bacterial maxima at 
the same time each year. The surface water usually had fewer bacteria 
than the bottom water. Bacteria in Pond A ranged from 4,400 to 102,700 



Soil Science 



427 



POND C 

• 'Surface Water 

0---0 Water 6" From 
Bottom 




AMJJASONDJFMAMJJASONDJFM 

POND B 

• • Surface Water 

0---0 water 6" From 
Bottom 




AMJJASONDJFMAMJJASONDJFM 
1967 1968 1969 

TIME OF SAMPLING 

Figure 2. Seasonal variations in organic carbon content of the pond waters for a 
two-year period. 



per ml in the surface water and from 5,100 to 118,100 in the bottom 
water. Pond B had only small variations in bacterial numbers with 
similar numbers of bacteria at both depths throughout the 2-year 
period. Bacteria in Pond B ranged from 4,200 to 48,000 per ml in the 
surface water and from 3,400 to 28,300 per ml in the water near the 
bottom. Pond C had the highest numbers of bacteria and the largest 
seasonal variations of bacterial numbers. In Pond C, the bacteria ranged 



428 



Indiana Academy of Science 



Table 2. Seasonal variations of bacterial number 
for a two-year period. 



in thousands per ml 





Pond A 


Pond B 


PondC 


Month 


Upper 


Lower 


Upper 


Lower 


Upper 


Lower 


April 1967 


20.7 


28.3 


14.2 


10.4 


7.7 


3.2 


May 


94.1 


110.6 


48.0 


10.8 


17.0 


18.2 


June 


70.6 


40.1 


11.8 


12.5 


160.0 


___ 


July 


11.1 


22.4 


6.2 


11.6 


9.9 


17.6 


August 


4.8 


9.7 


5.3 


10.2 


13.3 


19.2 


September 


4.4 


7.8 


5.0 


4.2 


10.9 


24.5 


October 


10.4 


12.2 


9.6 


6.9 


15.0 


3.6 


December 


15.1 


16.3 


5.7 


6.4 


25.2 


31.3 


February 1968 


15.9 


20.1 


15.9 


15.4 


77.8 


75.1 


April 


6.8 


20.1 


16.9 


20.6 


64.6 


75.6 


June 


102.7 


118.1 


10.6 


28.3 


109.3 


48.4 


July 


46.7 


42.8 


25.2 


12.0 


102.5 


37.3 


August 


16.6 


32.7 


25.1 


24.1 


68.4 


56.4 


September 


11.6 


14.3 


7.1 


27.7 


18.8 


26.7 


November 


5.3 


15.5 


10.2 


9.6 


13.1 


41.3 


December 


4.5 


5.1 


4.2 


3.4 


10.4 


8.8 


January 


45.2 


47.2 


5.8 


6.5 


111.2 


121.0 


March 


42.7 


44.1 


9.2 


8.5 


25.7 


45.8 



from 7,700 to 160,000 per ml in the surface water and from 3,200 to 
121,000 per ml in the bottom water. 

Many chromogenic bacterial colonies appeared on the plates in- 
cluding red, pink, shades of yellow and orange, white and cream colored. 

A statistical analysis of variance of the bacterial numbers is given 
in Table 3. In each of the ponds, the numbers of bacteria varied sig- 
nificantly with time of sampling, location of sampling, and depth of 



Table 3. Analysis of variance of bacterial numbers. 



Source of 




F Values 




Variation 


Pond A 


Pond B 


Pond C 


Time of Sampling 


859.41** 


276.68** 


681.51** 


Location of Sampling 


55.58** 


49.72** 


23.15** 


Depth of Sampling 


74.58** 


3.49** 


20.73** 


Time x Location 


40.72** 


55.07** 


19.76** 


Time x Depth 


24.61** 


186.74** 


100.26** 


Location x Depth 


24.87** 


47.92** 


6.76** 


Time x Depth x Location 


18.67** 


61.62** 


14.79** 



** Significant at the 1% level. 



Soil Science 



42'.) 



POND C 
• Surface Water 
° Water 6" From 
^o v Bottom 




r= AMJJASONDJFMAMJJASONDJFM 



en 



POND B 
-• Surface i Water 
-o Water 6" From 
Bottom 




A M J 
1967 



JASONDJFMAMJJASONDJFM 
1968 1969 

TIME OF SAMPLING 



Figure 3. Seasonal variations in bacterial numbers for a two-year period. 



sampling. Table 4 shows correlation coefficients for organic carbon and 
water temperature with bacterial numbers in each of the ponds. Corre- 
lation coefficients between water temperature and bacterial numbers were 
not significant in Ponds A and B but were highly significant in Pond C. 
The correlation coefficients between organic carbon and bacterial num- 
bers were highly significant in both Ponds A and B while a negative 
correlation was significant at the 57c level in Pond C. 



430 



Indiana Academy of Science 
Table 4. Correlation coefficients 





Organic 


Watsr 


Bacterial 




Carbon 


Temperature 


Numbers 






Pond A 




Organic Carbon 


1.000 


.445 


.440** 




(580) 


(260) 


(580) 


Water Temperature 


.445 


1.000 


.042 




(260) 


(320) 


(320) 


Bacterial Numbers 


.440** 


.042 


1.000 




(580) 


(320) 


(720) 






Pond B 




Organic Carbon 


1.000 


.122 


.299** 




(580) 


(260) 


(580) 


Water Temperature 


.122 


1.000 


—.005 




(260) 


(320) 


(320) 


Bacterial Numbers 


.299** 


—.005 


1.000 




(580) 


(320) 


(720) 






Pond C 




Organic Carbon 


1.000 


.355 


—.115* 




(560) 


(260) 


(560) 


Water Temperature 


.355** 


1.000 


.325** 




(240) 


(300) 


(300) 


Bacterial Numbers 


—.115* 


.325** 


1.000 




(560) 


(300) 


(700) 



* Significant at 5% level. 

** Significant at 1% level. 

Values in parentheses indicate number of samples. 



Summary 

Seasonal variations of bacterial numbers were found in a 2-year 
study of three Indiana farm pond waters. Statistical analysis showed 
significant variation of numbers of bacteria in each pond with time 
of sampling, location of sampling, and depth of sampling. In addition, 
correlation coefficients for water temperature with bacterial numbers 
were significant in Pond C and not significant in Ponds A and B. Cor- 
relation coefficients for organic matter content with bacterial numbers 
were significant in Ponds A and B, but a significant negative correlation 
co-efficient was found in Pond C. 



Soil Science 431 

Literature Cited 

1. Fred, E. B., F. C. Wilson, and A. Davenport. 1924. The distribution and significance 
of bacteria in Lake Mendota. Ecology 5 :322-339. 

2. Graham, V. E., and R. T. Young. 1934. A bacteriological study of Flathead Lake, 
Montana. Ecology 15:101-109. 

3. Irwin, W. H., and J. Claffey. 1968. Soil turbidity, light penetration and plankton 
populations in Oklahoma ponds and lakes. Proc. Okla. Acad. Sci. 47 :72-81. 

4. Snow, Laetitia M., and E. B. Fred. 1926. Some characteristics of the bacteria of 
Lake Mendota. Wis. Acad. Sci., Arts and Letters. 22:143-154. 

5. Stark, W. H., and Elizabeth McCoy. 1938. Distribution of bacteria in certain lakes 
of northern Wisconsin. Zentrbl. Bakt. Abt. II. 98 :201-209. 

6. Van Slyke, D. D., and J. Folch. 1940. Manometric carbon determination. J. Biol. 
Chem. 136:509-551. 

7. Wilson, H. A., T. Miller, and Rosa Thomas. 1966. Some microbiological, chemical 
and physical investigations of farm ponds. West Virginia Agriculture Experiment Sta- 
tion Bull. 522T. p. 1-17. 



Adsorption of Insecticides on Pond Sediments and Watershed Soils 1 
N. L. Meyers, J. L. Ahlrichs and J. L. White, Purdue University 

Abstract 

Adsorption of malathion, phorate, and carbaryl was studied on pond sediments and 
watershed soils. Mineralogical characterization showed the clay fractions of both the 
sediment and soil to contain kaolinite, micaeous minerals and vermiculite. Adsorption 
studies with the soils revealed malathion was adsorbed to the greatest extent followed 
by carbaryl and then phorate. Since adsorption tends to retain pesticides in soil, the 
probability of contamination of pond water following application of these insecticides is 
slight. 

Introduction 

The pollution of water resources by pesticides and other organo- 
toxicants has received much attention since contamination of the Ten- 
nessee River by toxaphene in 1951 as reported by Young and Nicholson 
(7). Entrance of a pollutant into water from use on agricultural land is 
regulated by the factors controlling the fate of pesticides in the soil. 
Removal of the pesticide from the soil might occur through leaching, 
volatization, or runoff while adsorption of pesticides tends to retard 
or prevent removal. In addition, pesticides may undergo alterations in 
the soil as the result of chemical, biological, or photochemical processes. 
The ultimate fate of a pesticide depends on a combination of these 
parameters. However, adsorption on colloidal surfaces of the soil ap- 
pears to determine to a greater extent than any other single factor 
the ultimate fate of pesticides. The nature and extent of adsorption 
has been discussed by Bailey and White (2) and Meyers (4). 

This study deals with the adsorption of three insecticides on water- 
shed soils and their corresponding pond sediments. Generally, adsorp- 
tion of a pesticide to a significant degree will greatly reduce, if not 
eliminate, movement into surface waters. To facilitate the study, three 
small farm ponds were selected on the Purdue University Southern 
Indiana Forage Farm in Dubois County. The watershed soil types were 
Zanesville (6-18% slope) and Welston (12-18% slope) silt loams which 
have developed from sandstone and shale. The soils are similar except 
for the fragipan formation in the Zanesville soil. Each of the water- 
sheds was devoted to alfalfa production. Prior to the application of in- 
secticides, soil samples were collected from the watersheds and sedi- 
ment samples were collected from the pond bottoms. The mineralogical 
characterizations and the adsorption studies described below were con- 
ducted using these materials. Since the mineralogical and physical 
properties of the three ponds were similar, only results from one of 
the ponds are reported. 



1 Contribution from Purdue University Agronomy Department, Lafayette, Indiana. 
Published with the approval of the Director of the Purdue University Agricultural 
Experiment Station as Paper No. 3888. This study was supported by the Office of Water 
Resources Research, Department of Interior. 

432 



Soil Science 



4)5.", 



Results 
Mineralogical Characterization 

The mineralogical composition of the clay fraction of pond sedi- 
ment in relation to its watershed soil has not previously been reported 
in Indiana. Therefore, the mineralogical composition of both soil and 
sediment was determined to provide information on this relationship 



SOIL CLAY 



POND I5A 
< 2.0 u 



SEDIMENT CLAY 




Mg-SATURATED 



GLYCEROL SOLVATED I 



K- SATURATED 



I i / K- SATURATED 
HEATED 300°C 



K-SATURATED 
HEATED 55CTC 




Figure 1. X-ray diffractogram of the soil and sediment clay. 



434 



Indiana Academy of Science 



as well as to provide a basis for meaningful interpretation of the 
adsorption studies. The mineralogical composition was determined by 
x-ray diffraction and infrared techniques. The x-ray diffractograms of 
the soil and sediment clay fraction (<2.0/x) from the pond are shown 
in Figure 1. The positions of intensities of the peaks show the min- 
eralogical composition of the soil and sediment to be essentially identi- 
cal. The 14 A peak present on Mg saturation, glycerol solvation, and 
mild heating was interpreted to indicate the presence of vermiculite. 
The 10 A peak is characteristic of micaeous minerals and is greatly 
enhanced by collapse of the 14 A material on strong heating (550° C). 
The presence of kaolinite is confirmed by the 7 A peak remaining on 
K saturation and mild heating and by the disappearance of the 7 A 




SOIL CLAY 
POND I5A 



SEDIMENT 
CLAY 



SOIL CLAY 
POND I5B 

SEDIMENT 
CLAY 



SOIL CLAY 

POND I3B 

SEDIMENT 
CLAY 



1020 



780 



3800 



COO 



OOO 



800 



FREQUENCY (CM'') 



FIGURE 2. Infrared absorption spectra for soil and sediment clays. 



Soil Science 



435 



peak on heating to 550° C which destroys the kaolinite structure. De- 
tailed procedures for the identification of clay minerals are given by 
Whittig (6). 

The infrared patterns of the clay fractions are shown in Figure 2. 
The presence of kaolinite is confirmed by the weak band at 3690 cm " 
and a strong band at 3620 cm'. Montmorillonite or vermiculite would 
also exhibit a strong band at 3620 cm J in addition to a broad band in 
the 3400 cm J region. Details concerning the application of infrared 
spectroscopy to clay mineral systems has been given by Ahlrichs 
etal. (1). 

Although the mineralogical composition of the clays appears to be 
identical, the quantity of clay is much higher in the sediment than in 
the soil (Table 1). This is in agreement with the work of Kohnke (3) 
and is in the order expected. Thus, clay and silt are preferentially 
eroded into the pond at the expense of sand but no differential erosion 
of clay types occurs. 





Table 1. Texture of the watershed 


soil 


and pond sediment. 




% Sand % Silt % Clay 




Coarse clay 

( /c of total 


Fine clay 
% of total 


Soil 
Sedin 


15.6 68.4 16.0 
lent 4.6 70.4 25.0 




63.1 

58.7 


36.9 
41.3 



600 


i i 1 1 1 1 1 1 


- 


ADSORBENT 

O O 
O O 




- 


E 

v. 
Q 


MALATHION 


- 


w 300 

CD 

QL 

O 

O) 

§ 200 




y' 


- 


E 


/ CARBARYL 


- 


100 


/ ^^^ PHORATE 




(r i i i i i i i i 



10 20 30 40 50 60 70 

EQUILIBRIUM CONCENTRATION figm/ml 

Figure 3. Adsorption isotherms for the three insecticides. 



80 



436 Indiana Academy of Science 

Adsorption Studies 

Adsorption of malathion, phorate, and carbaryl was studied using 
watershed soils and the pond sediments as adsorbents. Selection of 
these materials was based largely on their current use on alfalfa for 
weevil and spittlebug control. Adsorption of the insecticides is repre- 
sented as Fruendlich isotherms in Figure 3. It can be noted that mala- 
thion is strongly adsorbed followed by carbaryl, with phorate showing 
limited adsorption. 

Comparison of adsorption on soil with adsorption on sediment 
showed little difference for malathion and phorate while carbaryl ad- 
sorption was greater on the sediment. Examination of Table 1 would 
lead one to expect greater adsorption on the sediment due to the in- 
creased clay content. However, the expected increase occurred only 
with carbaryl. 

Absolute interpretation of adsorption data is of limited value but 
comparison of the relative extent of adsorption is worthwhile. If a 
pesticide is adsorbed it would be less likely to enter a pond as com- 
pared to a non-adsorbed counterpart. We might expect then that the 
degree of pollution for these insecticides following application to the 
soil would be in the order phorate > carbaryl >> malathion. In addi- 
tion, adsorption in the soil under field conditions should be more com- 
plete since concentrations used in laboratory studies represent an appli- 
cation of 10-300 times the normal rates of application. 

Conclusions 

Little difference was found between the clay mineralogy of a pond 
sediment and the soil from which it is derived. The soils and sediments 
used in this study contain kaolinite, micaeous minerals, and vermiculite. 

Adsorption studies showed that malathion is adsorbed to the 
largest extent followed by carbaryl and then phorate with significant 
adsorption by the soil of all three. If a pesticide is adsorbed, it should 
not be subject to movement into a pond by leaching or runoff; thus, we 
would conclude that contamination of a farm pond probably would not 
occur following application of malathion, carbaryl or phorate to the 
watershed, and that phorate would probably be the first to reach the 
pond if contamination did occur. These conclusions are based on the 
assumption that pesticide applications are made according to the manu- 
facturers suggestion and applied at the recommended rate. 

Further support of these observations is given by the work of 
other cooperators on the project (5). Continuous monitoring of the 
pond water for 8 months following application of phorate and carbaryl 
at levels 4 times their recommended dosage showed no trace of carbaryl 
at anytime in the water and only a slight temporary trace of phorate. 
When recommended rates were applied, no trace of either insecticide 
was found in the pond. 



Soil Science 437 

Literature Cited 

1. AHLRICHS, J. L., J. R. Russell, R. D. Harter, and R. A. Weismiller. 1965. Infrared 
Spectroscopy of Clay Mineral Systems. Proc. Indiana Acad, of Sci. 75 :247-255. 

2. Bailey, G. W., and J. L. White. 1964. Review of adsorption and desorption of organic 
pesticides by soil colloids, with implication conerning bioactivity. J. Agr. Food Chem. 
12:324-332. 

3. Kohnke, H. 1950. The reclamation of coal mine spoils. Advances in Agron. 2:318-349. 

4. Meyers, N. L. 1968. Adsorption of organic insecticides on well characterized watershed 
soils and their corresponding pond sediments. M.S. Thesis. Purdue University. 

5. Office of Water Resources Research. 1969. Effect of pesticide residues and other 
organic-toxicants on the quality of surface and ground water resources. Annual 
Report. Purdue University, Lafayette, Indiana. 

6. Whittig, L. D. 1965. X-ray diffraction techniques for mineral identification and 
mineralogical composition. In Methods of Soil Analysis. Part 1. Ch. 49. Number 9 in 
the series Agronomy. Amer. Soc. of Agron. Inc., Madison, Wisconsin. 

7. Young, L. A., and H. D. Nicholson. 1951. Stream pollution resulting from the use 
of organic insecticides. Prog. Fish Cult. 13:193-198. 



ZOOLOGY 

Chairman: James C. List, Ball State University 
Ralph D. Kirkpatrick, Ball State University, was elected Chairman 

for 1970 



ABSTRACTS 

Some Modifications in Rat Ovaries and Uteri Following Aminoglute- 
thimide Treatment. Egerton Whittle, Indiana State University. — 
Immature female albino rats (Charles River Strain) were injected 
subcutaneously with aminoglutethimide phosphate (AGP) for a 15-day 
period beginning on day 25 post partnm. The experimental animals were 
given dosages of either 50, 100, or 200 mg/kg body weight per day. At 
the end of the period of treatment the ovaries and uteri were excised 
and the weights compared with the weights of ovaries and uteri of 
control animals to determine whether AGP produced a definite effect on 
the size and weight of these organs and the general stature of the 
animals. 

The overall body weight of the treated groups displayed an in- 
creasing retardation with increasing dose levels of AGP. The amount 
of weight gain decreased almost linearly with increased amounts of 
AGP injected. 

The effect of AGP on ovarian weight in the treated animals was 
that of an apparent intial stimulation to growth at the lower dose level 
but an abrupt reversal of this trend when higher dose levels were ad- 
ministered. It would seem likely that AGP effects involve more than 
a single mechanism of action. 

The growth of uteri was inhibited increasingly with increasing 
dosage of AGP. At the highest dose level the uterine weight was only 
about Vz that observed in control animals. 

The evidence from this study indicates some relationship between 
presence of AGP in a system and the amount of estrogen present in the 
system, as circulating estrogen is the main governing factor for de- 
velopment and growth of the organs observed. 

Big Brown Bat Eptesicus fuscus Movement in Tunnel Cave, Cliffy 
Falls State Park, Indiana. James B. Cope and Richard Mills, Earl- 
ham College. — The senior author and students have studied the bat 
population in Tunnel Cave for the last five winters. During this time, 
the disturbance by banding and the subsequent reading of bands caused 
unnatural movements in the cave. Color banding techniques were tried 
which reduced the disturbance considerably after the initial banding. 
Movement was greater than expected and further refinement of tech- 
nique was employed. 

439 



440 Indiana Academy of Science 

Every 6 hours ( 6 AM, 12 Noon, 6 pm, and 12 Midnight) for a 5-day 
period during the last week in January, the cave was monitored for 
banded bats. Flashlights were used and only color bands that were 
visible were recorded. No bats were disturbed even if the band could 
not be read. There is overwhelming evidence that some bats come out 
of torpor and fly during this period even when the outside temperature 
isO°F. 



Parasites of Feral Housemice, Mus musculus, in 
Vigo County, Indiana 

John 0. Whitaker, Jr., Indiana State University 

Abstract 

The major external parasites of the housemouse, Mus musculus, in Vigo County, 
Indiana, are Myobia musculi, Radfordia affinis, Myocoptes musculimis, Ornithonyssus 
bacoti, Androlaelaps fahrenholzi, and Dermacarus heptneri (all are mites), in approxi- 
mate order of decreasing abundance. Among the internal parasites, Heligmosomoides 
polygyrus is the most abundant, followed by cestodes (presently unidentified and listed as 
a group), Protospirura sp., and Syphacia sp. Males and females harbored similar parasite 
infestations, but there was a definite increase in internal parasite load with increased 
age of the mouse. The same trend was apparent in one species of mite, Radfordia 
affinis, but was not evident in the rest of the species of ectoparasites. Heligmosomoides 
is primarily a spring and winter form and occurred at its greatest abundance in the 
single habitat present only at that time, winter wheat, while cestodes were most common 
during the summer and fall. The data are scanty, but suggest Syphacia to be a spring 
and summer form. Myocoptes was primarily a fall form. Most external parasites reached 
their greatest abundance in the fall and summer. Myobia was most abundant in winter 
wheat and corn, Radfordia in soybeans and corn, Myocoptes in corn and Ornithonyssus 
bacoti in sorghum. No relation was found between internal and external parasites ; their 
distributions seemed independent of each other. 

Introduction 

A number of housemice, Mus musculus, were taken during studies 
of the mammals of Vigo County, Indiana (2). Of these, 470 from 
randomly selected plots were examined for external parasites, and 503 
for internal parasites. The present study was initiated to determine 
if there were seasonal, sex, age or habitat differences in parasite in- 
festations. 

Some of the information has been presented previously, in a paper 
on the fleas of the mammals of Vigo County (3), in a paper on the mites 
of the small mammals of Vigo County (4), and in a paper on the Labi- 
dophorine mites of North America (1). 

Donald Norris was kind enough to make the nematode identifica- 
tions. 

Methods 

External parasites were obtained by searching the fur with dissect- 
ing needles and a 10 to 60X zoom binocular dissecting microscope. Mites 
were cleared and stained overnight in cold Nesbitt's Solution, mounted 
in Hoyer's Solution and ringed with asphaltum. Fleas were run through 
the alcohols and mounted in permount. Internal parasites were preserved 
in 70 % alcohol or formal-acetic acid (FAA). 

Data concerning incidence of parasitism were presented in terms 
of percentage of mice infested, with Chi-square being used to test for 
significant differences using the actual numbers of parasitized mice in 
the different categories. Abundance information was presented in terms 
of average number of parasites per mouse, but this number often seemed 

441 



442 



Indiana Academy of Science 



of less value than the incidence values because of the great amount 
of variation in the numbers of parasites infesting individual mice, with 
a few individuals harboring large numbers. For this reason "t" tests 
were not run. 

External parasites 

Other than some of the species of mites, there were few external 
parasites on the housemouse. One mouse yielded about 15 larval ticks, 2 
mice each had 1 flea, 1 had 2 lice, while 124 mice had mites (Table 1). 

Table 1. Comparison of parasites of male and female Mus musculus. 







270 Males 






219 Females 






Infestation 


No. /Mouse 


Infestation 


No./M 
Total 


ouse 


Parasites 


No. 


% 


Total 


Avg. 


No. 


% 


Avg. 


Internal Parasites 


















Heligmosom oides 


















polygyrus 


35 


L3.0 


482 


1.70 


21 


9.5 


171 


0.78 


Cestodes 


21 


7.8 


93 


0.34 


22 


9.6 


04 


0.43 


Syphacia 


5 


1.9 


27 


0.10 


3 


1.4 


40 


0.22 


Protospirura 


5 


Lit 


6 


0.02 





2.7 





0.04 


Cuterebra larvae 


1 


0.4 


1 


0.004 





0.0 





0.0 


Ascarid larvae 





0.0 





0.0 


1 


0.5 


1 


0.005 


Heterakis 





0.0 





0.0 


1 


0.5 


1 


0.005 






253 Males 






214 Females 




External Parasites 


















Myobia musculi 


2 4 


9.5 


132 


0.52 


12 


5.6 


29 


0.14 


Radfordia affinis 


23 


9.1 


51 


0.20 


12 


5.6 


22 


0.10 


Ornithonyssus bacoti 


16 


6.3 


35 


0.14 


2 


0.0 


3 


0.01 


Androlaelaps fahrenholzi 


11 


4.3 


25 


0.10 


3 


1.4 


3 


0.01 


Myocoptes musculinus 


7 


2.8 


10 


0.04 





4.2 


27 


0.13 


Dermacarus heptneri 


2 


o.s 


57 


0.22 


1 


0.5 


1 


0.005 


Dermacarus hypudaei 


2 


o.s 


2 


0.01 





0.0 





0.00 


Hirstionyssus talpae 


1 


0.4 


3 


0.01 


l 


0.5 


1 


0.005 


Eulaelaps stabularis 


1 


0.4 


1 


0.004 





0.0 





0.0 


Larval ticks 


1 


0.4 


ir> 


0.06 





0.0 





0.0 


Hoplopleura cajjtiosa 


1 


0.4 


2 


0.01 





0.0 





0.0 


Misc. Mites 


13 


5.1 


IS 


0.07 


11 


5.1 


15 


0.07 


Listrophorus leuclcarti 





0.0 





0.00 


1 


0.5 


1 


0.005 


Orchopcas leucopus 





0.0 





0.00 


1 


0.5 


1 


0.005 


Ctenojnhalmus pseudagyr. 


tea 


0.0 





0.00 


1 


0.5 


1 


0.005 


Androlaelaps morlani? 





0.0 





0.00 


1 


0.5 


1 


0.005 



Mites, then, were the dominant form of external parasites, with 11 
species being found in all. Seven of these, Androlaelaps morlani (?), 
Dermacarus hyjmdaei (not previously reported), Eulaelaps stabularis, 
Haemogamasus longitarsus and Listrophorus leuckarti were repre- 
sented by only one or two specimens each, and Hirstionyssus talpae by 
only four, hence these species were not considered as important para- 
sites of the housemouse in the area under consideration. 



Zoology 443 

The hypopial form of the mite, Dermacarus heptneri, was found on 
3 housemice, totaling; 58 individuals. This species is very tiny and can be 
easily overlooked, hence it may be more abundant than indicated. Mites 
of this type cling tenaciously to the individual hairs; one must separate 
the hairs with dissecting pins and examine their bases to find the 
hypopi. D. heptneri was found on no other species of Vigo County 
mammal, but the hypopi of another species of Dermacarus, D. hypudaei, 
were found on two housemice. D. hypudaei is primarily a species of 
Zapus, but is also found on other species, especially Microtus ochro- 
gaster and M. pennsylvanicus. 

Androlaelaps fahrenholzi, totaling 29 individuals on 14 mice, was a 
parasite of the housemouse, as was the case with most species of small 
mammals examined. It occurred at a lower rate, at 0.07 individuals per 
housemouse, than on most of the other species (4). 

Fifteen housemice yielded 56 specimens of the tiny Listrophorid 
mite, Mycoptes musculinus. This species of mite was not restricted to 
the housemouse, but all except seven specimens taken were from that 
host. 

Ornithonyssus bacoti was an important mite on Mus musculus, 34 
specimens being taken from 18 mice. This mite occurred at a similar 
rate on Peromyscus maniculatus bairdi, and three specimens were 
taken from P. leucopus. 

Radfordia affinis was taken almost entirely on Mus, with 65 individ- 
uals being taken from a total of 36 different mice. Two specimens were 
taken from Peromyscus maniculatus in the area under consideration (4). 

Myobia musculi, a tiny white form similar in general appearance to 
Radfordia affinis, and like that species a myobiid mite, was the most 
common species of external parasite on Mus musculus in Vigo County. 
It was taken on 9.1% of the housemice, but was not found on any of the 
other species of small mammals examined (4). 

The housemouse, in Indiana, appears to be relatively flealess, only 2 
fleas being taken on the 470 housemice examined during the present 
study, 1 each of Orchopeas leucopus and Ctenopthalmus pseudagyrtes. 
A third species, Epitedia wenmanni, again one specimen, was pre- 
viously reported from a Vigo County housemouse (3). Likewise Wilson 
(5) took only two fleas from Indiana Mus, one an Epitedia wenmanni, 
the other a specimen of Orchopeas leucopus. 

One of the housemice examined yielded two lice, Hoplopleura 
captiosa, the only louse recorded from Mus in Indiana. Wilson (5) 
took four specimens from the housemouse from Carroll and Tippecanoe 
Counties. 

One housemouse yielded about 15 larval ticks, which were pre- 
served, but subsequently lost. Wilson (5) reported three different 
individuals of the tick, Dermacentor variabilis, from Mus from three 
Indiana counties. 



444 Indiana Academy of Science 

Internal Parasites 

No trematodes or acanthocephalans were taken from any of the 503 
housemice examined for internal parasites, while both nematodes and 
cestodes were found to be relatively common. 

Among the internal parasites the most abundantly represented 
group overall was the Nematoda, especially Heligmosomoides polygyrus. 
This species was generally tightly coiled when seen among materials 
from the intestinal tract, and was red. Fifty-six of the mice examined, 
or 11.1% contained from one to over 100 individuals of this species, 
totaling 653 worms, and averaging 1.30 worms per mouse. Eight of the 
mice yielded over 25 worms per host. This species was found mostly in 
Mus, but a few individuals, apparently H. polygyrus were found in 
Peromyscus maniculatus bairdi. 

Eleven mice had 15 nematodes of the genus Protospirura, nearly 
all of which were in the stomach rather than in the intestine. Female 
oxyurids, Syphacia, were found in nine mice, totaling 96 specimens. 

Cestodes were important as parasites of Mus musculus, but unfor- 
tunately these have not yet been identified and are treated here as a 
group. Forty-three housemice yielded 187 cestodes. 

Cuterebra, sp., a botfly larva, was found in one mouse; one mouse 
yielded a nematode of the genus Heterakis; one harbored five objects 
from under the skin of the head which appeared to be larval cestodes; 
and one yielded a larval Ascarid nematode. 

Parasites in Relation to Sex of Mice 

A total of 270 male housemice were examined for internal parasites, 
of which 62, or 22.9%, yielded 593 parasites, or 2.20 per mouse. Of the 
210 females examined, 47, or 22.3%, yielded 325 parasites, or 1.55 per 
mouse. Thus, males and females were about equally infested in terms 
of incidence, but the rate of infestation in terms of average number of 
parasites per mouse was higher in males than in females in this 
sample. 

The two main types of internal parasites were nematodes, 
Heligmosomoides polygyrus, and cestodes. Of the 270 male housemice 
examined, 35, or 13.0%, harbored 482 nematodes, Heligmosomoides, 
averaging 1.79 worms per mouse for all mice, or 13.77 worms per mouse 
for those parasitized. Twenty-one of the 219 females, or 9.5%, yielded 
171 Heligmosomoides , averaging 0.78 worms per mouse overall, and 8.1 
in those infested. The higher average number in the males was pri- 
marily because of three males with particularly large infestations of 
40, 58 and 100 worms. 

A total of 93 cestodes were taken from 21, or 7.7% of the male Mus 
examined, averaging 0.34 cestodes per mouse overall, or 4.43 per para- 
sitized mouse. In the females, 22 of 219, or 10.0%, harbored 94 cestodes, 
for an average of 0.43 per mouse, or 4.27 in the mice infested. Thus 



Zoology 445 

males and females harbored relatively similar internal parasite loads. 
(Comparisons for the remainder of the individual kinds of parasites can 
be found in Table 1.) 

Parasite Infestation and Age of Animal 

To determine the relationship between age of the animal and para- 
site infestation, the mice were divided into 4 groups based on weight, 
those under 10 g, 10.0 to 14.9 g, 15.0 to 19.9 g, and those over 20 g. 

There was a significant increase (Chi-square = 28.75**, 3 df) in the 
incidence of animals with internal parasites with increased age of the 
animals, going from 9.1 through 16.5, 23.5 and 47.8% of the mice being 
parasitized in the four size groups (Table 2). This trend was apparent in 
both the most important parasite groups, the nematode H elig mosomoides , 
and in the cestodes. For Heligmosomoides, the largest class again har- 
bored the most worms per mouse, on the average, while the three 
smallest classes were about the same, but for cestodes there was an 
increased mean number of individuals per mouse with increased age. 



Table 2. Major parasites of four size classes of housemice, Mus musculus. 





Tinder 10 g 


10.0-14.9 g 


15.0-19.9 g 


20 g 


and over 




% 


Avg.#/ 


% 


Avg.#/ 


% 


Avg.#/ 


% 


Avg.#/ 


Parasites 


Infest . 


Mouse 


Infest. 


Mouse 


Infest. 


Mouse 


Infest. 


Mouse 


Interna] Parasites 


















Number examined 


(66) 


(206) 


(162) 


(69) 


(503) 


All internal parasites 


9.1 


1.32 


16.5 


1.32 


23.5 


1.32 


47.8 


5.58 


Heligomosomoides polygyrus 


4.5 


1.02 


9.2 


0.86 


11.7 


0.90 


21.7 


3.86 


Cestodes 


0.0 


0.00 


4.9 


0.17 


8.6 


0.35 


26.1 


1.44 


Syphacia sp. 


3.0 


0.32 


i.r. 


(1.24 


1.2 


0.03 


2.9 


0.30 


Protospirura 


1.5 


0.02 


0.9 


1.94 


3.7 


0.05 


2.9 


0.03 


External Parasites 


















Number examined 


(61) 


(194) 


(153) 


(62) 


All external parasites 


26.2 


0.51 


21.6 


0.79 


31.5 


1.14 


29.0 


1.10 


Myobia musculi 


3.2 


0.03 


5.2 


0.16 


11.7 


0.66 


9.7 


0.44 


Radfordia affinis 


3.2 


0.0c 


4.6 


0.11 


7.S 


0.1S 


1(1.1 


0.25 


Myocoptes musculinus 


4.9 


0.K 


2.6 


0.04 


3.3 


0.10 


3.2 


0.08 


Ornithonyssus bacoti 


0.0 


0.0( 


4.1 


0.07 


4.6 


0.10 


4.8 


0.10 


Androlaelaps fahrenholzi 


4.9 


0.2« 


2.1 


0.04 


2.6 


0.03 


4.S 


0.06 



Among external parasites as a group there was no such apparent 
trend in incidence (Table 2). Respective values for the 4 size classes are 
26.2, 21.6, 31.5 and 29.0%. In most cases there was no definite relation- 
ship between parasites and age, or else the data were too scanty to draw 
conclusions. One species, however, Radfordia affinis, did show the same 
tendencies as the internal parasites. It is more apt to be taken in the 
older mice (Chi-square r= 10.10, 2 df). 



440 



Indiana Academy of Science 



Parasite Infestation and Season 

Data were summarized (Table 3) on a seasonal basis as Spring 
(Mar. through May), Summer (June through Aug.), Fall (Sept. through 
Nov.), and Winter (Dec. through Feb.). 

Table 3. Seasonal distribution of major parasites of Mus musculus. 





Spring 


Summer 


Fall 


Winter 




% 


Avg.#/ 


% 


Avg. #/ 


% 


Avg.#/ 


% Avg.#/ 


Parasites 


Infest. 


Mouse 


Infest. 


Mouse 


Infest. 


Mouse 


Infest. Mouse 


Internal Parasites 
















Number examined 


(79) 


(88) 


(194) 


(142) 


All internal parasites 


40.5 




19.3 




18.0 




19.0 


Heligmosomoides 


34.2 


4.48 


5.7 


0.89 


5.2 


0.24 


9.9 1.23 


Cestodes 


1.3 


0.06 


11.4 


0.65 


12. !) 


0.57 


4.2 0.10 


Syphacia 


2.5 


0.51 


4.5 


0.35 








2.1 0.18 


Protospirura 


5.1 


0.09 


2.2 


0.02 


1.5 


0.02 


1.4 0.02 


External Parasites 
















Number examined 


(77) 


(85) 


(168) 


(140) 


All external parasites 


19.3 




34.1 




42.3 




12.9 


My obi a 


3.8 


0.43 


8.2 


0.46 


13.1 


0.41 


2.9 0.15 


Radfordia 


2.6 


0.17 


10.6 


0.27 


12.5 


0.1S 


1.4 0.01 


Myocoptes 








2.4 


0.02 


7.7 


0.32 





Omithonyssus 


1.3 


0.04 


11.8 


0.24 


3.0 


0.07 


1.4 0.02 


A. fahrenholzi 


1.3 


0.01 


1.2 


0.01 


5.4 


0.14 


2.1 0.02 



The greatest percentage of housemice harbored internal parasites 
during the spring, at 40.5%, while just under 20% of those taken were 
parasitized during the other 3 seasons. This difference was significant 
(Chi-square = 14.45, 1 df). The parasite load is heaviest during the 
spring because the principle internal parasite, Heligmosomoides 
polygyrus, is primarily a spring parasite, reaching by far its greatest 
percentage infection of mice, and its greatest average number of worms 
per mouse at that time. This nematode had its second greatest occurrence 
during the winter. Cestodes, as a group the second most important of 
the internal parasites, were least common during the spring and winter, 
and most common during the summer and fall. For Syphacia, the data 
are scanty, but this form would appear to be a spring and summer 
form. It was not taken at all in the fall sample of mice, even though 
this was the largest sample, at 194. Protospirura seemed to be most 
abundant in the spring, but data are too few concerning this species to 
be reliable. 

External parasites, as a group, were most abundant in the fall and 
summer, and least abundant in the winter (Chi-square = 29.47, 3 df). 
Myobia, musculi, Radfordia ajfinis and Myocoptes mus&ulinus were fall 
and winter mites, with Myocoptes occurring only during that season. 
Omithonyssus bacoti was taken at its greatest rate in the summer, and 
Androlaelaps fahrenholzi was taken at its greatest rate in the fall. With 
the exception of A. fahrenholzi, all species occurred at their lowest 
abundance in the winter. 



Zoology 



447 



Parasites of Mus as Associated with Habitat 

There were enough data concerning: Mus parasites in eight habitats 
to make a meaningful presentation (Table 4). Late stage winter wheat 
(over 6 inches high) was the one habitat available only during the 
spring. As seen previously, Heligmosomoides was primarily a spring 
parasite, and as one might expect, winter wheat was the habitat in 
which the greatest incidence and abundance of this nematode occurred. 
The second greatest abundance occurred in cut corn, primarily a winter 
habitat, although some cut corn areas were available in the fall, and 
some were still present in the spring. The major habitats for cestodes 
were corn and cut wheat. Syphacia was most abundant in the winter 
wheat over 6 inches, and in winter wheat which had been cut. This latter 
habitat was available in late spring and early summer. 



Table 4. 



Relationship of Mus parasites to habitat. (Numbers in parentheses are the 
numbers of plots in the habitats.) 















Internal Parasites 








Helimosomoides 


Cestodes 


Syphacia 
Avg.#/ 


Prot 


ospirura 






Avg.#/ 




Avg.#/ 


Avg.#/ 


Habitat 






% 


Mouse 


% 


Mouse 


% 


Mouse 


% 


Mouse 


Weedy field (58) 






3.4 


0.05 


5.2 


0.17 












Grassy field (69) 






5.8 


0.72 


4.3 


0.35 







2.9 


0.06 


Soybeans (25) 






12.0 


0.12 


4.0 


0.04 


4.0 


0.04 


4.0 


0.08 


Winter wheat 






















over 6" (37) 






29.7 


5.22 


5.4 


0.27 


5.4 


1.08 


5.4 


0.08 


Corn (118) 






3.8 


0.18 


15.4 


0.65 







2.3 


0.02 


Sorghum (31) 






0.7 


0.77 







3.2 


0.03 







Wheat, cut (79) 






10.1 


0.71 


12.7 


0.65 


3.8 


0.62 


2.5 


0.03 


Corn, cut (69) 






27.5 


4.35 


4.3 


0.07 


2.0 


0.07 


1.4 


0.01 












External Parasites 












Myobia 


Radfordia 


A. fahrenholzi 

Avg.#/ 


Myocoptes 

Avg.#/ 


0. 


bacoti 






Avg.#/ 




Avg.#/ 




Avg.#/ 


Habitat 


% 


Mouse 


% 


Mouse 


% 


Mouse 


% 


Mouse 


% 


Mouse 


Weedy field (54) 


5.6 


0.37 







3.7 


0.04 







3.7 


0.07 


Grassy field (65) 


4.6 


0.05 


7.7 


0.12 


6.1 


0.06 


1.5 


0.05 


t.5 


0.02 


Soybeans (23) 


4.3 


0.04 


17.4 


0.39 







4.3 


0.04 







Winter wheat 






















6" (37) 


8.1 


0.89 


2.7 


0.03 

















Corn (118) 


14.4 


0.64 


16.1 


0.26 


5. 1 


0.17 


10.2 


0.42 


2.5 


0.08 


Sorghum (25) 


8.0 


0.12 

















20.0 


0.24 


Wheat, cut (78) 


7.7 


0.28 


3.8 


0.10 


1.3 


0.01 


1.3 


0.04 


7.7 


0.21 


Corn, cut (67) 


1.5 


0.04 


3.0 


0.19 


1.5 


0.02 


o 




1.5 


0.03 



Myobia was most abundant in the winter wheat and in the corn 
while Radfordia was taken at its greatest rates in soybeans and corn. 
Myocoptes was quite definitely a form of the cornfields while 
Ornithonyssus bacoti was found at its greatest rate in sorghum. 



448 Indiana Academy of Science 

Discussion 

Season, habitat and age of the host all seemed to influence the para- 
site population of the housemouse, Mus muscuhis, while the fourth 
factor under consideration, sex of the animal, seemed to have little or no 
effect. Some factors appeared interrelated in such a way that it was 
difficult to determine which of two factors was affecting- the mice. For 
example, the nematode parasite, Heligmosomoides polygyrus, was most 
abundant in the spring in winter wheat fields in which the wheat was 
at least 6 inches high. Hence, one can conclude that the best situation 
in which to look for this species is in winter wheat fields during the 
spring, but I was unable to evaluate the relative effects of the 
particular season as opposed to those of the habitat. 

The relationship of internal and external parasitism was assessed 
in an attempt to determine whether animals became parasitized because 
of their generally poor physical condition, or if the animals simply 
happened to be in the right place at the right time to become para- 
sitized. If animals were infected because of a general overall poor 
physical condition, then the same animals that were parasitized inter- 
nally should also tend to be parasitized externally. On the other hand, 
if parasitism was strictly a chance happening, then one should be able 
to compute the approximate number of mice expected to have both 
internal and external parasites by multiplying the percentage of mice 
with external parasites times the percentage with internal parasites. 
For this calculation, only the 470 mice examined for both internal and 
external parasites were used. Of these, 130, or 0.277 were found to have 
external parasites, and 102, or 0.217 were found to harbor internal para- 
sites. One would expect, by chance, that 0.277 X 0.217 = 0.060 of the 
470, or 28.20 mice would have both internal and external parasites. The 
actual number of mice with both internal and external parasites was 30, 
hence we can conclude that the relationship between internal and 
external parasites of the housemouse in Vigo County is strictly a 
chance one. 



Literature Cited 

1. Rupes, V., and J. O. Whitaker, Jr. 1968. Mites of the subfamily Labidophorinae 
(Acaridae, Acarina) in North America. Acarologia 10 :493-499. 

2. Whitaker, J. O., JR. 1967. Habitat and reproduction of some of the small mammals 
of Vigo County. Indiana, with a list of mammals known to occur there. Occas. Papers 
Adams Ctr. Ecol. Studies. 16 :l-24. 

3. Whitaker, J. O., Jr., and K W. Corthum, Jr. 1967. Fleas of Vigo County, Indiana. 
Proc. Indiana Acad. Sci. 76 :431-440. 

4. Whitaker, J. O., Jr., and N. Wilson. 1968. Mites of small mammals of Vigo County, 
Indiana. Amer. Midland Natur. 80 :537-542. 

5. Wilson, N. 1961. The ectoparasites (Ixodides, Anoplura and Siphonaptera) of 
Indiana mammals. Unpublished thesis. Purdue University. 527 p. 



Molt in Two Populations of the House Mouse, Mus musculus 

Ronald E. Brechner and Ralph D. Kirkpatrick, 
Ball State University 

Abstract 

This study correlated molt in two population samples of the house mouse, Mus 
musculus, with sex, age, geographic origin and time of year. Sample A consisted of 
273 house mice trapped in 1964 and 1965 on Sand Island, Johnston Atoll, central 
Pacific Ocean. The 272 mice of Sample B were caught over a 6-month period in Delaware 
County, Indiana, in 1968 and 1969. 

Incidence of pigmented areas (molt) on the fleshside of the pelts of both samples 
was recorded and statistically analyzed to find patterns and percent molt. There was 
no positive correlation between incidence of molting and origin and age of sample mice, 
nor between incidence of molting and sex or month of collection. 

Introduction 

The present study correlated molt (as indicated by pigmentation of 
the fleshside of pelts) in two feral populations of the house mouse, Mus 
musculus, with sex, age, geographic location and time of year. The term 
"feral" as used here implies mice which are either living out-of-doors or 
in unheated buildings. 

Sample A represented an insular population from the central 
Pacific Ocean. Sample B was taken from Delaware County, Indiana. 
Percentage molt and areas of molt were recorded and each specimen 
was assigned to an arbitrary age class. 

Related Literature 

Osgood (11) noted three different phases of pelage color in 
Peromyscus which corresponded to age — juvenile, adolescent, and adult. 
Allen (1) described molt in this same genus as generally beginning in 
the feet and around the nose and extending dorsally and also proceed- 
ing anteriorly from the base of the tail. Collins (3) agreed that this is 
the general pattern of spring molt in Peromyscus maniculatus, but that 
it is subject to much irregularity. He found that the juvenile pelage was 
complete at 4 to 5 weeks and transition from juvenile to post juvenile 
pelage occurred between the ages of 6 and 8 weeks. Molt on the ventral 
surface was usually completed before the dorsal surface and usually 
proceeded from anterior to posterior. Collins (3) checked for molt by 
parting the hair and checking for the presence of new hair. But as 
pointed out by Golley et al. (5), the molt is more readily determined 
from the underside of the skin and may be well underway before it is 
detected on the surface. 

Gollschang (6) stated that some Peromyscus leucopus, both captive 
and wild, molt over an indefinite period of time, and that body weight 
had no direct relationship to onset of pelage change. 

Most mammals have similar pigmented areas as illustrated by 
Kopenen (9) who examined the sequence of pelages of the Norwegian 

449 



450 INDIANA ACADEMY OF SCIENCE 

lemming, Lemmus lemmus, by using" the pigment patterns of the 
fleshside of the skin and Skoczen (12), who studied seasonal changes of 
pelage of the mole, Talpa europaea, as measured by the planimetric 
measuring of pigmented areas on the underside of the skin. 

Golley et al. (5) studied skins of Peromyscus polionotus by pinning 
the skins out to dry on cardboard for 6 months with the furside down 
and then using a planimeter to measure total area exhibiting molt. He 
found that molt was influenced by increasing age; however, he also noted 
it may be influenced by sex, body size, trauma, and other environ- 
mental factors. The percentage of the pelt involved in the molt process 
progressively declined with age. He found 80 to 90% of total pelage 
involved in molt in juveniles, but it never exceeded 45% at later stages 
of development. There was a 35% maximum in post-juvenile molt. 
Golley et al. (5) also noted areas of gray pigmentation preceding or 
following the areas of most active molt. 

The pigmentation of flat-skin mounts of 33 specimens of known age 
Microtus calif ornicus used in an experiment by Ecke and Kinney (4) 
revealed a close age-molt correlation from 17 to 60 days of age. Animals 
could be aged by degree of molt to within 4 days of their actual age. 
The method was 88% accurate with laboratory-reared mice. Evidence 
from skins of old animals indicated that all adult molts are irregular 
after the molting of post-juvenile pelage. These occurred as mottled 
patterns of dark and light areas on the underside of the skin with no 
apparent consistency of design. No significant molt variation was 
found between the two sexes. 

Methods 

Sample A was collected on Sand Island, Johnston Atoll, central 
Pacific Ocean, by the junior author while working with the Pacific 
Ocean Biological Survey Program in 1964 and 1965. The atoll covers less 
than 800 acres and this insular population has developed since 1923 when 
the U.S.S. Tanager Expedition biologists found no mammals on the 
island (8). 

Sample B was taken from Delaware County, Indiana. A minimum 
of 30 mice per month was collected during a 6-month period from 
September 27, 1968, through March 22, 1969. The mice came from 
2 county locations: 1) Delaware County Fairgrounds, Muncie, Indiana; 
2) Earl Southworth's farm — 1 mile west of Tillotson Avenue on Bethel 
Pike, Muncie, Indiana. 

All but 6 of the 272 mice in Sample B were assigned to arbitrary 
age classes based on molar wear after Lidicker (10). The remaining 6 
were not aged because the skulls were destroyed during trapping. 
Data collected from the 266 usable specimens were compared with data 
from the 273 mice in Sample A. 

Mice were collected using live-traps and snap-traps. The pelt was 
removed from the carcass, tagged, and pinned with the furside down to 
dry. Facia and excess fat were removed as suggested by Clark (2), 
to prevent their masking of molt pattern. 



Zoology 



151 



Percentage of molt was estimated. A grid system composed of 
Vs-inch squares was imposed on a plastic transparency. This was then 
placed over the skinside of each pelt, and the percentage of molt was 
calculated by comparing the total number of squares covered by the 
pelt to the number of squares covered by pigmented areas. 

We decided that a modified version of the pelt pattern used by 
Hendricks (7) would best demonstrate the possible patterns (Fig. 1). 
A map with 14 areas was prepared for each pelt. To designate an area 
as molting, a minimum of Vs square inch of that area must be molting 
as demonstrated by the presence of pigmentation. This indicator was 
arbitrarily chosen. 

The areas of molt on each pelt were marked on the map of the 
pelt, along with other pertinent information available for that speci- 




Figure 1. Pelt map of house mouse illustrating areas of molt. 



452 



Indiana Academy of Science 



men. A combination of Chi-square and co-factor analysis was run to 
find the most frequently occurring patterns. 

After these patterns were determined, each pelt map was again 
examined and classified as representing one of the main patterns, as 
irregular, or as displaying no molt. 

A pelt was recorded as having a certain molt pattern if it dis- 
played molt in 50% or more of the areas comprising that pattern. It 
was possible for more than one pattern to be represented on the same 
pelt. This information was analyzed for correlation of molt pattern with 
age, sex, geographic origin, and time of year. 

Results and Discussion 

Population Samples A and B were analyzed separately for repre- 
sentative molt patterns. Five patterns were found in Sample A and four 
patterns in Sample B (Fig. 2). The molt patterns were lettered A 
through I; J was used to indicate an irregular unclassified molt pattern; 
and K represented no molt. 



SAMPLE A 

3 A5A1 ! 



2, 12,13 




SAMPLE 



Figure 2. Predominant molt pattern in samples from two house mouse populations. 



Each pelt was again checked to determine which pattern or patterns 
it represented. It was possible for several patterns to be represented on 
one pelt. When this occurred, the composite of patterns was considered 
as one pattern. For example, many of the pelts represented one clear-cut 
pattern, such as I or H; but, some of the pelts had patterns F, G, H, I 
represented on the same pelt. Such pelts were considered as representing 
one pattern, the pattern of FGHI. 

Males and females within the same sample were considered jointly, 
since they were characterized by the same distribution and frequency 
of molt patterns. The molt patterns were randomly distributed through- 
out the age classes and the months of collection, and did not show 



Zoology 453 

positive correlation with either of these. The samples were represented 
by different patterns, with the