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of the 

Indiana Academy 
of Science 

Founded December 29, 1885 

Volume 78 

William R. Eberly, Editor 

Manchester College 
North Manchester, Indiana 

Spring Meeting- 
April 26-27 
The Honeywell Center, Wabash 

Fall Meeting 

October 18-19 

Ball State University, Muncie 

Published at Indianapolis, Indians 

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

2. Instructions for Authors appear at the end of this volume, P. 450. 

3. Exchanges, Items sent in exchange for the Proceedings and corre- 
spondence concerning 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 

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 relatives and friends of deceased members 
by the Secretary. 

7. Officers whose names and addresses are not known to correspond- 
ents may be addressed care of the State Library. Papers published in the 
Proceedings of the Academy of Science are abstracted or indexed in appro- 
priate 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 


Part 1 



Officers and Committees for 1968 9 

Minutes of the Spring Meeting 12 

Minutes of the Fall Meeting (Executive Committee) 15 

Minutes of the Fall Meeting (General Session) 18 

Annual Financial Statement 20 

Annual Report, Junior Academy of Science 22 

Biological Survey Committee Report 29 

Necrology 32 

New Members for 1968 43 

Part 2 


Presidential Address 49 

"Urban Geology — A Need and A Challenge", William J. Wayne 

"Science, Communication, and the Critical Mass" 65 

Robert E. Gordon 


R. L. Michael— Stratigraphy of the White Site* 71 

R. E. Pace, S. Coffin, and J. Richardson — A Preliminary Report 

on the Welsh-Dunlap Site, Vigo County, Indiana* 71 

B. J. Morris — The Initial Excavation of the Van Nuys Site* 71 

T. Curren— Mound Four New Castle Site* 71 

M. Salovesh — Culture Change in a Maya Community* 71 

R. L. Blakely, R. J. Marmouze, and D. D. Wynne — The Incidence 
of the Perforation of the Coronoid-olecran Septum in the Middle 
Mississippian Population of Dickson Mounds, Fulton County, Illi- 
nois 73 

P. L. Walker — The Linear Growth of Long Bones in Late Woodland 

Indian Children 83 

R. W. Alexander, Jr. and G. K. Neumann — On the Origin of the 

Tutelo— An Eastern Siouan Tribe 88 

L. M. Robbins and G. K. Neumann — The Origin of the Shawnee 

Indians 93 

B. R. Huelsman — Urban Anthropology and the Southern Moun- 
taineer 97 

M. S. Weiss — A Systems Approach to the Study of Complex 

Society 104 


B. H. Peterson, Z. Brahmi, J. S. Ingraham, and A. S. Levine — De- 
velopment of a Modified Antibody Plaque Technique for the De- 
tection of Single Cells Making Anti-viral Antibody* 109 

K. Carlson and R. Bochrath — Amber Streptomycin-resistant Mu- 
tants of Escherichia colv f 109 

B. Peri and M. Wagner — Immune Response to Streptococcus faecalis 

in the Rat* 110 

*Abstract or Note only 

2 Indiana Academy of Science 

A. C. Raitano and A. S. Levine — Enzymatic Effect of Cobra Venom 

on Rauscher Leukemia Virus (RLV) * 110 

W. F. Campbell and A. S. Levine — Suppression of Rauscher virus- 
induced Murine Leukemia by L-Asparaginase* 110 

P. C. Morgan and R. F. Ramaley— Physiological Studies of the In- 
corporation of 5-Bromouracil During Growth and Sporulation in 
Bacillus subtilis-16&* Ill 

D. R. Brannon, M. Gorman, B. B. Molloy, W. M. Stark, and J. 

Mabe — Biosynthesis of Thiadiketopiperazine Antibiotics* Ill 

R. H. Williams, J C. Cline, R. E Holmes, and M. J. Sweeney — 

Mycophenolic Acid: Studies on Biological Activities* 112 

P. A. Lemke — Genetic Evidence for Resistance of Cephalosporium to 

Specific Compounds* 112 

J. H. Nuner and R. J. Downey — Effect of Oxygen on the Synthesis 

of Nitrate Reductase in Bacillus stearothermophilus* 113 

S. H. Kendall and S. A. Minton— The Serum Profiles of Certain 
Reptile Sera and preliminary Observations on Antibody Forma- 
tion in Snakes* 113 


D. L. Dilcher and C. A. Zeck — A Study of the Factors Controlling 

Variation of Cuticular Characters* 115 

G. E. Dolph and D. L. Dilcher — An Eocene Discovery of Dendro- 

panax* 115 

R. E. Girton — Effects of Selenium on the Respiration of Excised 

Root-tip Segments of Maize* 116 

A. T. Guard — Some Disappearing Plant Species* 117 

A. J. Ullstrup — Weather and Corn Diseases in Indiana in 1968* .... 117 
J. E. Rahe and J. Kuc' — Induced Resistance of Phaseolus vulgaris to 

Bean Anthracnose* 118 

L. and A. Beesley — Lobelias of Franklin Co. and Indiana* 118 

K. K. Curtis and D. E. Smith — Daily Variation in Chlorophyll Con- 
tent of Corn Seedlings* 118 

R. W. Judd and J. J. Nisbet — Pennsylvanian Coal Ball Flora of 

Indiana 120 

C. M. Palmer — Algal Records for Three Indiana Sewage Stabilization 

Ponds 139 

D. J. Morre', B. Rau, R. Vieira, T. Stanceu, and T. Dion — Environ- 

mental Regulation of Experimental Leaflet Abcission 146 

Cell Biology 

E. J. Hinsman and K. Moe — The Fine Structure of the Ventral 

Horn Neuron in the Calf Spinal Cord* 161 

I. Watanabe, S. Donahue, and W. Zeman — An Intranuclear Struc- 
ture in Neurons of Human Cerebral Cortex* 161 

W. Yunghans and D. J. Morre' — Chemical Composition of Mem- 
brane Fractions Isolated from Rat Liver in Relation to Membrane 
Differentiation During Secretion* 161 

K. M. Mak and R. A. Jersild — An Electron Microscopic Study of 
Zinc Iodide-Osmium Staining of the Golgi Apparatus of Rat 
Intestinal Epithelial Cells* 161 

J. B. Whitten, Jr. — The Ultrastructural Features of Intraoral 

Lichen Planus, Simplex* 162 

D. J. Niederpruem and R. A. Jersild — Direct Studies of Nuclear 

Movements in Schizophyllum commune* 163 

J. F. Schmedtje — Plasma Cell Antibody Against Bovine Serum 
Albumin in the Rabbit Appendix as Revealed by the Fluorescent 
Antibody Technique* 163 

: Abstract or Note only 

Table of Contents 3 

W. E. Stovall and G. L. Rosene — Tumor Cell Mitotic Activity in 

Mice Treated with Antigenic Materials* 164 

S. F. Smalley — Microimmunoelectrophoresis of Human Blood in 

Regard to the Study of the Gc System* 164 

G. Barski, J. W. Butler, and R. J. Thomas — Quantitative Measures 
of In Vitro Cell Mobility by Use of a Pattern Recognition Com- 
puter* 165 

D. J. Morre' and H. H. Mollenhauer — Studies on the Mechanisms 

of Gluteraldehyde Stabilization of Cytomembranes 167 

A. E. Middleton, R. Cheetam, D. Gerber, and D. J. Morre' — Adeno- 

sine Mono-, Di- and Trinucleotidase Activities of Rat Liver Cyto- 

mebranes 183 

J. D. Hall, J. W. Stiles, Y. Awasthi, and F. L. Crane — Membrani- 

fibrils on Cristae and Grana Membranes 189 


R. M. Lawrence— X-Ray Diffraction Study of Aqueous Thallium 

(III) Chloride* 199 

F. J. Holler and R. D. Joyner — Studies of a Cyanamide Complex 

of Iron (II)* 199 

R. L. Stover and R. D. Joyner — Phtalocyaninogermanium (II) and 

Other Phthalocyaninogermanium Compounds* 199 

L. A. McGrew and S. C. Robling — A Kinetic Study of the Reaction 

of Phenyl Isocyanate with Water* 199 

B. N. Storhoff and J. R. Doyle — Organonitrile Complexes of Rheni- 

um (I) and Manganese (I)* 200 

R. F. Copeland — Use of Computers in Undergraduate Physical 

Chemistry* 200 

A. G. Cook and T. A. Hecht — Synthesis and Properties of [2.2.2] 

Bicyclooctyl Enamines* 200 

A. G. Cook and D. J. Schultz — Reduction of Selected Enamines 

with Lithium Aluminum Hydride* 200 


R. M. Dinkel and F. Rothwell — Temperature and Moisture Rela- 
tionships of Green County, Indiana Strip Mine Areas* 201 

A. G. Craske, Jr. — Ecological Site Preference and Taxonomic Differ- 
ences within Two Acer sacchanim-Acer nigrum Complexes found 
in Parke County, Indiana* 201 

J. L. Gerwig and W. B. Crankshaw — Effects of Thermal Discharge 
on the Phytoplankton and Macroinvertebrates of the Wabash 
River* 201 

F. Morgan— Effects of Effluent on the Fish Population of Mill Creek, 

Rochester, Indiana* 202 

P. T. McKelvey and C. E. Smith, Jr.— A Study of Selected Physio- 
chemical Properties of Two Indiana Main-Stream Reservoirs* . . 202 

R. O. Petty — Pattern of Mesic Forest Succession at the Western 

Border* 203 

J. R. Gammon — The Effect of Inorganic Sediment on Macroinverte- 

brate and Fish Populations of a Central Indiana Stream* 203 

W. B. Crankshaw — The Effect of Ground Cover on the Soil Moisture 

Regine in a Mixed Mesophytic Woods 204 

M. T. Jackson and P. R. Allen— Detailed Studies of Old-Growth 

Forests in Versailles State Park, Indiana 210 

P. M. Arnett — A Study of Collembolan Populations Associated with 

Four Serai Stages Leading to the Beech-Maple Climax 231 

W. B. Crankshaw, J. A. Smith and R. D. Kirkpatrick — Wood- 
cock Singing Ground Descriptions for Two Indiana Sites 241 

Abstract or Note only 

4 Indiana Academy of Science 

M. T. Jackson — Hemmer Woods: An Outstanding Old-Growth Low- 
land Forest Remnant in Gibson County, Indiana 245 


D. L. Schuder — A Japanese Weevil Discovered in Indiana* 255 

J. W. Hart — A Checklist of the Mosquitoes of Indiana with a 

Record of the Occurrence of Aedes infirmatus D&K 257 

F. N. Young — Crosses of Tropisternus from Central America with 

Other Color Forms of the Tropisternus collaris Complex (Cole- 

optera: HYDROPHILIDAE) 260 

T. A. Parker — An Annotated List of the Spiders of Indiana 266 

Geology and Geography 

J. H. Cleveland and C. F. Tiefel — Gypsum Resources of the Mid- 
western United States* „ 315 

A. F. Schneider — A Significant Exposure of Pleistocene Drift in 

South-Central Indiana* 315 

R. L. Powell — Unconsolidated Deposits on the Mitchell Plain of 

Indiana* 316 

L. A. Lewis — Analysis of Surficial Landform Properties: The Re- 

gionalization of Indiana into Units of Morphometric Similarity. . 317 

R. F. Boneham — Earth Science Teaching in the Secondary Schools 

of Indiana 329 

R. W. Orr — Stratigraphy and Correlation of Middle Devonian 

Strata in the Logansport Sag, North-central Indiana 333 

T. F. Barton — Planning for and Utilization of the Web Pattern of 

Physical Urban Development in Cities 342 

R. R. French — Transportation of Mineral Aggregates in Indiana . . . 348 

R. M. Dinkel and L. Guernsey — An Economic Appraisal of Recla- 
mation Practices on a Strip Coal Mine Site in Greene County, 
Indiana 355 

L. I. Dillon — Popcorn Production in Indiana 363 

History of Science 

D. Howard and T. R. Mertens — Academic Origins of Members of 

the Genetics Society of America 370 

N. G. Sprague — Original Science Apparatus Preserved in Science 

Museums and Universities in Free Europe 378 

R. H. Cooper — A Half Century of Science at Ball State University. . 381 


R. Callis and E. Craig — Preliminary Results of a Muon Energy 

Study* 387 

J. F. Houlihan — Determination of Absolute D* for Photovoltaic, 

Infra-red Detectors* 387 

R. D. Burgess and M. Hults — Possible Methods for Observing 

Shadow Bands at the Next Solar Eclipse in North America* . . 387 

D. E. Koltenbah — Nuclear Electric Quadrupole Resonance Analysis 
of Chemical Bonds in C135-Containing, Straight-Chain Hydro- 
carbons* 388 

G. P. Thomas — Low Energy Elastic Scattering of K-Mesons off 

Protons* 388 

T. V. Blanc, a. G. Danemar, and D. E. Koltenbah — The Design 
and Construction of a System for Direct Measurement of Atomic 
Lifetimes 389 

* Abstract or Note only 

Table of Contents 5 

Plant Taxonomy 

W. H. Welch — Hookeriaceae Species and Distribution in South 

America 396 

F. K. Daily — Some Late Glacial Charophytes Compared to Modern 

Species 406 

J. Humbles — Indiana Plant Distribution Records, XX, 1966-68 413 

G. C. Marks — The Flowering of Lemna minor and the Establishment 

of Centaurium pulchellum in Northwestern Indiana 414 

Soil Science 

L. E. Hughes and H. W. Reuszer — Fluctuations of Bacteriological 

Numbers in Farm Ponds* 417 

J. M. Smith — Earth Mound in Eastern Indiana* 417 

H. Kohnke and S. A. Barber — Tillage Techniques on Indiana Prairie 

Soil 418 

M. E. Heath — Fitting Plants to Fragipan Soils in Southern Indiana 429 
R. K. Stivers — Fertilizer Experiments with Corn on Several Soils in 

Indiana, 1963-1965 435 


D. C. Kramer — Temperature Preferences in the Eastern Garter 

Snake (Tharnnophis sirtalis sir talis) * . 445 

W. J. Brett — Relationship between Emergence Rhythm and Meta- 
bolic Rhythm in Drosophila melanogaster* 445 

J. M. Burns — The Effects of Chicken Luteinizing Hormone on the 

Pullet Ovary* 445 

A. W. Gruenholz and H. Tamar — Tracheal Mucous Velocities in the 

Rabbit, Dog, and Rat* 446 

R. E. Geyer, Jr. and W. B. Hopp — Studies on the Growth Rate of 

the Juvenile Piolt Black Snake (Elaphe obsoleta)* . . . 446 

J. 0. Whitaker, Jr., W. A. Miller and W. L. Boyko — Rabies in 

Indiana Bats 447 

M. W. Wagner and W. C. Gunther — Preference for Toxic and Non- 
toxic Artificial Sweeteners in Rodents 457 

R. H. Cooper — -Melanoma in Heloderma suspectum Cope 466 

G. P. Pollock — A Comparative Study of Some Effects of Amino- 

glutethimide Phosphate on Serum Potassium and Sodium 468 

B. L. Pickard and A. E. Reynolds — Aspects of Water Loss Physiol- 

ogy in Certain Plethodontid Salamanders 472 

H. W. Wendt — Rapid Approximations for some Chi Square and De- 
rived Correlational Statistics Used in the Social and Biological 

Sciences 482 

F. Don Fulk and J. O. Whitaker, Jr. — The Food of Rana catesbei- 

ana in Three Habitats in Owen County, Indiana 491 

R. E. Mumford — The Hoary Bat in Indiana 497 

Instructions for Contributors 502 

Index 505 

* Abstract or Note only 





William J. Wayne, President 

Officers and Committees for 1968 


President William J. Wayne, Indiana University 

President Elect Howard R. Youse, DePauw University 

Secretary James R. Gammon, DePauw University 

Treasurer Frank A. Guthrie, Rose Polytechnic Inst. 

Editor William R. Eberly, Manchester College 

Director of Public Relations James A. Clark, Indiana Department 

of Natural Resources 


Anthropology Ben K. Schwartz, Ball State University 

Bacteriology. Hayward Campbell, Jr., Eli Lilly & Co. 

Botany . Thomas Mertens, Ball State University 

Cell Biology Ralph Jersild, I. U. Med. Center 

Chemistry LeRoy A. McGrew, Ball State University 

Ecology. Wm. B. Crankshaw, Ball State University 

Entomology Leland Chandler, Purdue University 

Geology & Geography Lowell Dillon, Ball State University 

History of Science Robert Cooper, Ball State University 

Physics Edwin C. Craig, Ball State University 

Plant Taxonomy Carrolle A. Markle, Earlham College 

Soil Science Marion Baumgardner, Purdue University 

Zoology John 0. Whitaker, Indiana State University 


(Past Presidents*, Current Officers, Divisional Chairmen, 

Committee Chairmen) 

♦Baldinger, L. H. 

Behrens, 0. K. 

Campbell, H. Jr. 

Chandler, L. 
* Christy, 0. B. 

Clark, J. A. 
*Cleland, R. E. 

Coats, N. 

Cook, D. J. 

Cooper, R. 

Crankshaw, W. B. 

Craig, E. C. 

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

Dillon, L. 

Eberly, W. R. 
♦Edington, W. E. 

* Edwards, P. D. 

Gammon, J. R. 
*Girton, R. E. 
♦Guard, A. T. 

Guthrie, F. A. 
♦Haenisch, E. L. 

Heniser, V. 

Jersild, R. 
♦Johnson, W. H. 

Kaufman, K. L. 
♦Lilly, Eli 
♦Lindsey, A. A. 

List, J. C. 

McGrew, L. A. 
♦Markle, C. A. 

Markle, M. S. 
♦Mellon, M. G. 

Mertens, T. 

♦Meyer, A. H. 
♦Michaud, H. H. 
♦Morgan, W. P. 

Moulton, B. 

Petty, R. O. 
♦Porter, C. L. 
♦Powell, H. M. 

Schwartz, B. 

Stockton, Sister M.R. 
♦Wallace, F. N. 

Wayne, W. J. 
♦Weatherwax, P. 

Webster, J. D. 
♦Welch, W. H. 
♦Welcher, F. J. 

Whitaker, J. 0. 

Winslow, D. 

Youse, H. R. 

10 Indiana Academy of Science 

President, W. J. Wayne; President-elect, H. R. Youse; Secretary, J. 
R. Gammon; Treasurer, F. A. Guthrie; Editor, W. R. Eberly; Director 
of Public Relations, J. A. Clark; Retiring President, A. A. Lindsey; 
Director of Junior Academy, D. R. Winslow; Library Committee, Nellie 
Coats; Program Committee, J. C. List and L. A. McGrew; Relation of 
Academy to State, W. A. Daily. 


Academy Foundation: W. P. Morgan, 1969, chairman; W. A. Daily, 1970. 
Bonding: D. J. Cook, 1968, chairman; R. M. Brooker, 1968. 
Research Grants: O. K. Behrens, 1969, chairman; H. H. Michaud, 1968; 
J. B. Patton, 1972; W. Stephenson, 1971; W. H. Welch, 1970. 

(President an ex officio member of all committees) 

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

Auditing Committee: J. C. List, chairman; R. H. Cooper. 

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; B. Malin. 

Program Committee: J. C. List and L. A. McGrew, chairmen; R. H. 
Cooper; E. C. Craig; C. Parish; H. Paschall; D. F. Richmond; H. H. 
Roepke; B. K. Swartz, Jr. 

Publications Committee: W. R. Eberly, chairman; J. A. Clark; D. G. 
Frey; W. N. Meihorn; J. 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; 
O. Behrens; J. F. Hayden; G. H. Bick; M. Burne; K. H. Carlson; 
N. Coats; R. H. Coleman; G. B. Cummins; J. P. Danehy; F. K. 
Edmundson; H. Feldman; Olive Forbes; F. Frieders; J. R. Gammon; 
W. C. Gunther; F. A. Guthrie; R. E. Hale; W. E. Hoffman; W. B. 
Hopp; W. R. Hurt; E. R. Johnston; R. L. Kent; H. Kohnke; H. P. 
Leighly; Marie Mayo; J. McFarland; D. E. Miller; G. R. Miller; 
M. A. Moussa; M. Murphy; E. Nussbaum; P. A. Orpurt; J. B. 
Patton; R. Petty; S. N. Postlethwait; L. M. Reynolds; A. F. Sch- 
neider; M. C. Shanks; J. Siegrist; H. K. White; H. G. Wilhelm; L. 
Willig; F. J. Zeller; W. A. Zygmunt. 

Officers and Committees 11 

Fellows Committee: H. E. Driver; W. H. Welch; D. E. Miller; B. E. 
Montgomery; F. K. Daily; K. H. Carlson; R. L. Conklin; C. B. 
Heiser; E. J. Asker. 

Resolutions Committee: L. H. Baldinger, chairman; J. E. Newman; J. M. 

Invitations Committee: H. R. Youse, chairman; R. H. Cooper; W. B. 
Hopp; W. K. Stephenson; J. D. Webster. 

Necrologist: F. K. Daily. 

Parliamentarian: P. Weatherwax. 


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

Academy Conference Representative (President-elect): H. R. Youse. 

Emeritus Members Committee: R. E. Cleland, chairman; E. L. Haenisch; 
M. S. Markle; H. H. Michaud; W. H. Welch. 

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

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


The Honeywell Center, Wabash, Indiana 


April 26, 1968 

The meeting was called to order at 4:30 p.m. by Dr. William J. 
Wayne, President of The Academy, in the library of the Honeywell 

The minutes of the Executive Committee and the General Session 
of the Fall Meeting of The Academy held October 20, 1967 at Indiana 
University were read. 

Treasurer, Dr. Frank A. Guthrie reported Academy funds as fol- 

Balance January 1, 1967 $21,685.67 

Income for 1967 32,365.22 

Expenditures for 1967 37,435.45 

Balance, December 31, 1967 17,568.47 

The question of instituting an "Affiliate" status for groups such as 
the Indiana Chapter of the Society of American Microscopists was dis- 
cussed. It was generally agreed that there should be no official "Affiliate" 
status, but that if most of the members of a group were also Academy 
members then any meeting of the group could be considered an official 
function of the Academy. 

A motion was unanimously carried to accept Cell Biology as a full- 
fledged Division of The Academy. 

Auditing Committee: Dr. Robert Cooper reported the books of the 
Treasurer to be in good shape. 

Youth Activities Committee: Dr. W. J. Wayne reported that the 
Science Search dinner was well attended. 

Publications Committee: Dr. W. R. Eberly reported that the coming 
Volume of the Proceedings will contain several innovations including a 
complete list of Academy members which will improve its quality. One 
manuscript is now ready for consideration and another in preparation 
for the Special Publications Series. The Academy has agreed to support 
this program, but has not provided means of funding the project. After 
discussion by Frank A. Guthrie, William A. Daily and Paul Weatherwax, 
it was suggested that the publication of the monographs might be funded 
initially by the John S. Wright Fund subject to approval by the Re- 
search Grants Committee, and that money from the sales of the first 
monograph could be placed into a regular publications fund so that it 
could become self-sustaining eventually. Dr. Cleland suggested that W. 
R. Eberly and a member of the Library Committee meet with the Re- 


Minutes of the Executive Committee 13 

search Grants Committee to discuss the matter. Dr. Wayne agreed to 
coordinate such a meeting. 

Membership Committee: Requests have been made to Indiana Chap- 
ters of meteorology and cell biology for member lists in order that Sr. 
M. Rose may send those persons not already members of The Academy 
applications for membership. 

Scientific Areas Committee: The following resolution was adopted 

Whereas the fate of scientific and natural areas in Indiana is a 
concern of the Indiana Academy of Science, and; 

Whereas this body of men and women believes it is obliged to 
comment on policies of land use which, in its opinion, run contrary to 
the best interests, both scientific and civic, of the people of the state, 

Whereas a proposed reservoir on Big Walnut Creek in Putnam 
County, Indiana, is now planned which would permanently alter and 
partially destroy a natural valley landscape on this stream, and; 

Whereas this landscape harbors an outstanding system of plants 
and animals, including surpassing specimens of species which are relicts 
of an early post-glacial flora, species unusual in their occurrence, and; 

Whereas this landscape is deemed of greater value in its present 
natural condition, and of greater long range benefit to the people 
of the state as a nature preserve than as a reservoir as now envisioned, 

Whereas several feasible alternative sites are reported to exist by 
the Corp of Engineers, and; 

Whereas recent evidence has been submitted by the Indianapolis 
Water Company that future water supply for that metropolitan area 
is assured without said reservoir due to the newly proposed Mud Creek 
Reservoir, thus negating one of the major justifications for constructing 
the disputed reservoir, therefore 

BE IT RESOLVED by the Indiana Academy of Science by action 
of its Executive Committee this 26th day of April, 1968, to stand op- 
posed to the currently planned reservoir on Big Walnut Creek in Put- 
nam County, Indiana, as it is now proposed and to continue to oppose 
plans for any structure which would result in the inundation of any 
part of the natural valley landscape north of U. S. Highway 36. 

Science and Society Committee: Dr. Willis H. Johnson reported 
that the speaker's Bureau Brochure was published at a cost of $1000 
provided by Eli Lilly. The Progress in Government reported a willingness 
on the part of Academy members to act in governmental matters in- 
volving science. 

14 Indiana Academy of Science 

An invitation from The American Institute of Biological Science 
to The Academy to become an Affiliate member was declined unani- 

Dr. W. Wayne suggested that the fall meeting might include a 
symposium in teaching of high school science and if sufficient interest 
subsequently developed that a division could be added. Dr. J. List indi- 
cated that Indiana Biologists, a group of teachers mainly secondary, 
could act as a focal organization. Dr. Day reported that a group of 
high school teachers of chemistry was being organized. 

A motion was passed unanimously to change By-Laws, Article I. 
Dues as follows: 

Sec. 1. Initiation fees, reinstatement fees and dues for the various 
classes or types of memberships shall be determined annually by the 
Executive Committee. 

Sec. 2. Initiation fees are payable only once; reinstatement fees are 
payable each time a lapsed membership is reactivated. Annual dues are 
based upon the calendar year. 

Sec. 3. A student discount in the amount of 50% of the normal annual 
dues will be awarded to undergraduate and /or graduate students certi- 
fied to be eligible for this discount by an Academy member on the 
faculty of the student's college or university. The student discount is 
limited to a maximum of five years. 

Sec. 4. Advance payment of annual dues, beyond the current year, may 
be made at the existent rate for a period not to exceed three years, or 
in the case of students only through the calendar year in which studies 
should normally be completed, whichever occurs first. 
Sec. 5. Annual dues billings shall be made by the Treasurer prior to 
the spring meeting. Delinquent members shall be rebilled prior to the 
fall meeting. Members delinquent on December 20th shall be sent a 
third dues billing before December 31st, and if still delinquent on 
January 31st shall be dropped from the membership rolls. A former 
member, dropped for lack of dues payment, may be reinstated by pay- 
ment of both the reinstatement fee and dues for the year in which he 
wishes to resume membership. 

Sec. 6. Each member is entitled to one copy of the Proceedings of 
the Academy, and any other publications of the Academy distributed 
to the membership of the Academy, that are published in the year for 
which dues have been paid. 

Sec. 7 Former members who were dropped for non-payment of dues 
may obtain one copy of the Proceedings, or other publications, pub- 
lished in a year in which they were inactive by either of two methods, 
provided copies are still available: 

a. They may pay the reinstatement fee and dues for the year or 
years of interest. 

b. They may purchase one or more copies at the established non- 
member price. 

The meeting was adjourned at 7:00 p.m. 

Approved: October 18, 1968 James R. Gammon, Secretary 


Ball State University, October 18, 1968 

The meeting was called to order at 7:30 p.m. by Dr. William J. 
Wayne, President of the Academy, in Room 103 of the Physical Science 
and Mathematics Building. 

The minutes of the Executive Committee and the General Session 
of the Spring Meeting of the Academy held April 26-27, 1968, at the 
Honeywell Center, Wabash, Indiana, were approved. 

Treasurer — Dr. Frank A. Guthrie reported the Academy funds as 

January 1. 1968 balance $17,568.47 

Income to October 15, 1968 12,317.43 

Expended to October 15, 1968 8,456.84 

Balance, October 15, 1968 21,429.06 

Editor — Dr. William R. Eberly reported that the Proceedings will 
be printed starting next week and that it will contain a number of in- 
novations including a two-part format: Part 1: The Work of the 
Academy, and Part 2: Addresses and Contributed Papers. An entire 
member list will also be included. 

Research Grants Committee — Dr. O. K. Behrens reported that in 
conjunction with Dr. Eberly and Miss Nelle Coats, the Committee has 
worked out the details of supporting the Special Publications Series. 
It was felt that publications constituted an integral part of research. 
Therefore, $2,000.00 per annum will be set aside for the next few 
years to help the new program along. 

Six research grants totaling $2,219.00 have been approved. 

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

October 1, 1967, balance $ 625.00 

Receipts through September 30, 1968 727.00 

Disbursements through Sept. 30, 1968 300.00 

Balance, September 30, 1968 1,052.20 

In the John S. Wright Fund: 

Balance, October 10, 1967 $ 2,491.10 

Receipts to September 30, 1968 10,715.33 

Disbursements to Sept. 30, 1968 11,740.10 

Balance as of September 30, 1968 1,466.33 

Youth Activities Committee — Dr. V. Heniser discussed the methods 
of selection and financial support of the Talent Search Program and 
the general activities of the Committee. Mr. Winslow noted that 51 
clubs are now active in the Junior Academy. 

The Library Committee — Miss Nelle Coats reported that the pro- 
visions of the Lilly Endowment, Inc. grant were being completed and 
that only 372 volumes of Natural Features of Indiana remain. 


16 Indiana Academy of Science 

Relation of the Academy for the State — William A. Daily stated that 
increased support from the State was requested amounting to $5,500.00. 

Membership Committee — Sr. Mary Rose reported that efforts to 
involve more high school science teachers in the Academy continue. 

Fellows Committee — Dr. Winona Welch recommended and a motion 
was approved to accept the following members as Fellows of the 

Dr. James R. Gammon 
Dr. Helmut Kohnke 
Dr. Thomas R. Mertens 
Dr. Jerry J. Nisbet 

Emeritus Committee — Dr. Winona Welch recommended and a motion 
was approved to accept the following members as emeritus members 
of the Academy. 

Dr. Ira L. Baldwin 

Mrs. Gladys M. Friesner 

Dr. Blanch McAvoy 

Dr. John C. Roehm 

Invitations Committee — Dr. Howard Youse announced the follow- 
ing colleges as hosts for future Academy meetings: 
1969— Hanover College 
1970 — Indiana State University 
1971— Earlham College 
1972— St. Mary's College 

Biological Survey Committee — Dr. Dan Webster noted that papers 
on geographic distribution of flora and fauna are diminishing while 
ecological studies are increasing. 

Preservation of Natural Areas Committee — Dr. Robert Petty reported 
that 248 natural areas listed by counties is now available upon request 
on computer print-out. 

Committee on Science and Society — Dr. Willis H. Johnson reported 
on the current activities of the three subcommittees. 

The Speaker's Bureau Subcommittee chaired by Dr. John Christian 
is to be congratulated for their work on the brochure containing the 
names of available speakers and topics which has now been distributed 
to nearly 4,000 service clubs, women's clubs and high schools in the 
state. There is good evidence that it is already being used. A poll will 
be conducted next spring to ascertain the extent of usage and to deter- 
mine the frequency with which the speakers have been called upon. 

Dr. Day and the Subcommittee on Science and Government have 
discussed ways in which the Academy might offer its services to the 
State with Lieutenant Governor Rock and leaders of the House and 
Senate. A list of over 20 subjects of interest to the State was prepared 

Minutes of the Executive Committee 17 

and the scientists of the state were contacted about serving as con- 
sultants on one or more of these subjects. More than 160 scientists 
responded favorably. All legislators and legislative committees will be 
informed of this service and a representative of the committee will 
explain the program to the Governor and the Lieutenant Governor. Each 
member of the Academy can help further this program by contacting 
his senator and representative and explaining the available service. 

The Subcommittee on Finance reported that efforts to obtain funds 
for establishing an office with a part-time Executive Director and a 
secretary have thus far produced nothing. A new program of the NSF, 
however, has indicated an interest and a proposal will be submitted 
soon. It was suggested that support might be enlisted from businesses, 
industries and other organizations through a form of institutional mem- 
bership. Until outside funds are obtained, a continued budget of $416.42 
was requested to continue work on a limited scale. 

A motion was approved directing the Science and Society in con- 
junction with Dr. Weatherwax to prepare an amendment to the Consti- 
tution to include an "Institutional" membership to be voted upon at the 
spring meeting. 

Special financing was required for the publication of special ad- 
dresses to be distributed by the Science and Society Committee. A motion 
was approved to divert $400.00 toward their publication. 

After thorough discussion, a motion was approved to move the time 
of the Fall Meeting of the Academy from Saturday to Friday. A resolu- 
tion was passed that the Junior Academy of Science need not change 
their day of meeting unless they wished to do so. 

Dr. Wayne suggested that a short newsletter be mailed to the 
Academy membership shortly after the Spring Meeting to keep members 
informed about Academy business. He also expressed the hope that the 
Science Teaching Symposium could be continued. 

The meeting was adjourned at 9:50 p.m. 

Approved October 20, 1968. 

James R. Gammon, Secretary 


Ball State University, October 19, 1968 

The annual Fall Meeting of the Indiana Academy of Science was 
held in the Auditorium of the Teachers College Building on Saturday, 
October 19, 1968, at 11:00 a.m. Dr. William J. Wayne, President, called 
the meeting to order. Academy members were officially welcomed by 
Dr. Robert L. Carmin, Dean, College of Sciences and Humanities, Ball 
State University. 

The minutes of the Executive Committee meeting held Friday, 
October 18, 1968, were read by the secretary and approved as read. 

Fay Kenoyer Daily read a biographical sketch of each member who 
had died since the 1967 Fall meeting. These are printed under Necrology. 

Dr. Robert H. Cooper then introduced Dr. Robert E. Gordon, As- 
sociate Dean of the College of Science, University of Notre Dame, who 
presented a timely address entitled "Science, Communication and the 
Critical Mass." 

A luncheon for Junior and Senior Academy members was held in 
Cardinal Hall, Pittenger Student Center, at 12:30 p.m. 

The annual dinner meeting of the Academy was held in Cardinal 
Hall, Pettenger Student Center at 6:00 p.m., Dr. Howard R. Youse, 
President-Elect, presiding. 

Professor James E. Newman of the Resolutions Committee submit- 
ted the following resolution: "That the Academy members here assembled 
express their appreciation to Ball State University for all the courtesies 
which have been extended to the membership of the Academy during 
this meeting. We are indebted especially to Dr. James C. List and Dr. 
LeRoy A. McGrew, Co-chairmen of the Program Committee for their 
efforts in arranging facilities for this annual meeting. Further, the 
Academy is appreciative of the warm welcome extended by Dr. Robert 
L. Carmin, Dean, College of Sciences and Humanities at Ball State 
University and to Dr. Robert E. Gordon, Associate Dean, College of 
Science, University of Notre Dame, for his address to the general 
session." The resolution was approved. 

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

Dr. Alton A. Lindsey, Chairman of the Nominating Committee, 
presented the names of the divisional chairmen for 1969: Anthropology, 
Robert Pace, Indiana State University; Bacteriology, D. S. Wegener, 
Indiana University Medical School; Botany, Robert Kent, Indiana 
Central; Cell Biology, Edward J. Hinsman, Purdue University; Chem- 
istry, John W. McFarland, DePauw University; Ecology, Thomas Mc- 


Minutes of the Executive Committee 19 

Cornish, Ball State University; Entomology, Jack Munsee, Indiana State 
University; Geology and Geography, Wilton Melhorn, Purdue University; 
History of Science, B. Elwood Montgomery, Purdue University; Physics, 
Richard C. Conklin, Hanover College; Plant Taxonomy, Jack Humbles, 
Indiana University; Soil Science, James E. Newman, Purdue University; 
Zoology, James C. List, Ball State University. 

The following slate of officers and elected committees were pre- 
sented for election by Dr. Lindsey: President, Howard R. Youse, DePauw 
University; President-elect, Frank A. Guthrie, Rose Polytechnic Insti- 
tute; Treasurer, Damian Schmelz, St. Meinrad College; Director of 
Public Relations, Paul E. Klinge, Indiana University; Editor, William 
R. Eberly, Manchester College; (Secretary, James R. Gammon, DePauw 
University, continues in office for the final year of the three-year term 
of office); Bonding Committee, Robert M. Brooker, Indiana Central 
College and Howard H. Michaud, Purdue University (both 1969); Re- 
search Grants Committee, James E. Newman, Purdue University (1973). 
A motion to accept the officers and committee members was approved 

Dr. William J. Wayne presented a stimulating, illustrated speech 
which concluded the proceedings. 

The meeting was adjourned at 9:00 p.m. 

James R. Gammon, Secretary 

Approved April 25, 1969 

January 1-December 31, 1968 


A. 1968 Income: 

Item or Description Income 

Dues and Initiation Fees $ 4,003.00 

Reprint Sales to Authors (Balance of Volume 76) 450.40 

Publications Fund Income 

Sale of "Proceedings" $ 153.00 

Sale of "Natural Features of Indiana" 4 20.00 

John S. Wright Fund (Partial, Vol. 77) 3,500.00 4,073.00 

TOTAL 1968 INCOME: $ 8,526.40 

Plus interest credited to savings accounts +886.50 

Less 1968 Expenditures, below 

NET GAIN FOR 1968: $ 1,794.24 

Plus Balance, January 1, 1968 +6,925.22 

BALANCE, December 31, 1968: 

B. 1968 Expenditures: 
Item or Description 


Clerical $ 

Postage, etc. 


Postage, etc. 
Office Supplies & Expenses 
Travel Allowance & A.A.A.S. Conf. Dues 
President's Contingency Fund 
Membership Committee 

Reprints (President's Address, Necrology) 
Junior Academy of Science 
Proceedings Publication Costs 

Editorial, Vol. 77 

Printing, Vol. 77 

Mailing, Vol. 77 
Program Committee 

Chairman's & Local 

Printing & Mailing 
Science & Society Committee 
Library Binding 

From 1967 Budget 

From 1968 Budget * 1,000.00 

Reprint Costs, Vol. 77 (Authors) * 

Mailing, "Proceedings" & "Natural Features" * 

TOTAL 1968 EXPENDITURES: $ 7,618.66 $ 4,555.00 

$ 8,719.46 


re Budgeted 

$ 285.S3 

$ 250.00 

; 201.83 
































* Billings not yet received. 
** Plus Publications Income Fund Receipts, above. 
Self supporting item, not budgeted. 


Minutes of the Executive Committee 21 

ii. administered accounts 

Item or Description 

Publications Fund 
Operational Funds 

Acad. Research Fund 
Science Fair Fund 
Science Talent Search 
J.S. Wright Library Fund 
Lilly Library Fund III 
Science & Society Committee 



Bank Balances: Terre Haute First National Bank, Terre Haute, Ind. 3,882.73 
Equitable Savings & Loan Assn., Los Angeles, Calif. 5,069.26 
First Western Savings & Loan, Las Vegas, Nev. 14,584.60 


ACCOUNTS: $23,536.59 

Frank A. Guthrie, Treasurer 
December 31, 1968 

January 24, 1969 

We the undersigned have audited the Treasurer's records for the Indi- 
ana Academy of Science for the year 1968 and have found them to be 
accurate and in order. 

William G. Kessel, 
J. Lee Guernsey 

Jan. 1, 



Dec. 31, 


















































President: Dennis Waltke, Division of University Schools, Bloomington 

Vice-President: James Peterson, Brebeuf Preparatory School, Indianap- 

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


Dr. Howard Michaud, Honorary Chairman, Purdue University 

Mr. Keith Hunnings, New Haven Senior High (1965-1969) 

Mr. F. Ray Saxman, Cascade High School, Clayton (1966-1970) 

Mr. Charles Souers, Div. of University Schools, Bloomington (1966-1970) 

Miss Helen Reed, Manual High School, Indianapolis (1967-1971) 

Mr. David Blase, Arlington High School, Indianapolis (1968-1972) 


Prof. Virgil Heniser, Chairman, Indiana University, Morrison Hall 103 
Prof. Donald R. Winslow, Director, Indiana Junior Academy of Science, 
Div. of University Schools, Bloomington, Indiana 47401 


Thirty-Sixth Annual Meeting 
Saturday October 19, 1968 

8:00 A.M. 

Junior Academy Council Meeting. 

8:30-12:00 A.M. 

Registration and Election of Officers. 

8:30-10:30 A.M. 

Junior Academy Council Interviews for "Best Boy" and "Best Girl" 

9:00-10:50 A.M. 

Tours of Laboratories and Research Facilities or attend the Senior 
Divisional Meetings or Public School Science Teaching Symposium. 

11:00 A.M. 

General Session. 

Address: "Science, Communication, and the Critical Mass." Dr. 
Robert E. Gordon, Associate Dean, College of Science, University 
of Notre Dame. 


Program 23 

12:30 P.M. 

Luncheon, for Junior and Senior Academies. 

2:00 P.M. 

General Business and Presentation of Papers. Dennis Waltke, Presi- 
dent presiding. 

3:45 P.M. 

Announcements, presentation of awards. 

6:00 P.M. 

Annual Banquet. Address by retiring President of Senior Academy, 
Dr. William J. Wayne. 


1. Microbial Genetics: The Isolation, Characterization and Genetic 
Mapping of an Unknown Bacterium. 

Sam Combs, Lewis Cass Junior-Senior High School, Walton. 

2. The Characterization of East German Fossil Chlorophyll. 
Sharon Smith, Cascade High School, Clayton. 

3. The Extraction of Phytochrome from Green Plants. 
Jeanne Hagelskamp, Ladywood School, Indianapolis. 

4. A Study of the Subcellular Organization of the Leucine Specific 
Enzymes in Salmonella typhimurium CV-19. 

Marcia Stroud, Howe High School, Indianapolis. 

5. Effects of Cortisone Acetate on Hemopoiesis, the Hemopoietic 
Organs, and Growth Rate of Young Rats. 

Timothy J. O'Leary, Brebeuf Preparatory School, Indianapolis. 

6. Experiments on Pavlovian Inhibition: Inhibition VI. 
Alan Chepregi, Morton Senior High School, Hammond. 

7. The Effects of Drugs on Animal Behavior in Relation to Brain 
Serotonin and Norepinephrine. 

Vann Seawell, University Senior High School, Bloomington. 

8. In Vitro Maintenance of Rabbit Hearts on Modified Hank's Solution. 
Robert Finley McDavid 111, Schulte High School, Terre Haute. 

9. Why Do Spiders Spin Different Web Patterns ? 

Mark Watness, J. F. Kennedy Memorial High School, Indianapolis. 

10. An Observation of Effects of the Hormonal Regulators Kinetin and 
Benzyladenine on Plant Growth. 

Tammey Naab, Huntington County Community High School, Hunt- 

11. The Antiballistic Jet-Why ? 

Robert Humphreys, New Haven Senior High School, New Haven. 

12. Problems in the Construction of Lasers and Masers. 
Gary Stephen, Portland High School, Portland. 

13. Are the Chromosomal Puffs of the Chironomus tentans Influenced 
by Environmental Factors ? 

24 Indiana Academy of Science 

Martha Ziegelbauer, J. F. Kennedy Memorial High School, Indi- 

14. Quenching of Orbital Angular Momentum by Ligand Fields. 

V. Stephen Overstreet, Brebeuf Preparatory School, Indianapolis. 

15. Can the Silicone Rubber Membrane be Utilized to Separate Oxygen 
from Water? 

Gary Trowbridge, New Haven High School, New Haven. 

16. The Development Genetics of the Basidiomycetes and the Formation 
of Fruiting Bodies in Mycelium. 

Brent Kern, Howe High School, Indianapolis. 

17. A Study of Base Negative Two. 

Lucinda Lee Glentzer, Portland High School, Portland. 

18. The Role of Different Areas of the Cerebral Auditory Sensory 
System in Auditory Response. 

Dennis Waltke, University Senior High School, Bloomington. 


Members of the Portland Junior-Senior High School and Muncie 
Central High school were hosts for this meeting. Mr. Ralph Settle, 
Mr. Robert Freemyer, and Mr. William Beuoy were the sponsors. 



The thirty-sixth annual meeting of the Indiana Junior Academy of 
Science was held on Saturday, October 19, 1968 at Ball State Campus, 
Muncie, Indiana. 

One hundred and fifty-five students and nineteen sponsors repre- 
senting seventeen high schools were registered. 

President, Dennis Waltke, called the meeting to order at 2:00 p.m. 
in the University Auditorium. Previous to this session, members of the 
Junior Academy had enjoyed tours and lectures in various parts of the 

After a joint luncheon for members of both the Junior and Senior 
Academies in Cardinal Hall at Pittinger Student Center, the business 
meeting was conducted. Dennis Waltke, president, introduced the other 
officers: James Peterson, Vice-President and Rachel Koontz, Secretary. 
Dennis gave a brief welcome. The minutes of the 1967 meeting were 
read and approved. Eighteen papers were presented as listed in the 

Following the presentation of the papers, the President of the 
American Society for Microbiology, Indiana Branch, Dr. Stone, an- 
nounced the names of the winners of the Society's awards for the best 
papers in microbiology. Marcia Stroud of Howe High School received 

Minutes 25 

a certificate and a check for $25.00 for her paper entitled, "The Sub- 
cellular Organization of the Leucine Specific Enzymes in Salmonella 
typhimurium CV-19". Honorable mention included Sam Combs and 
Brent Kern. 

Dennis introduced Mr. Keith Hunnings who introduced the new 
officers and the winners of the best boy and best girl awards. Officers 
for 1969 are as follows: 

Dennis Waltke, President 
Timothy O'Leary, Vice-President 
Rachel Koontz, Secretary 

The best boy award went to Dennis Waltke, and Marcia Stroud and 
Rachel Koontz tied for the best girl award. Each received a certificate 
of recognition, a year's honorary membership in the American Associ- 
ation for the Advancement of Science and a year's subscription to Science 

Next year's meeting of the Senior Academy was set for Saturday, 
October 25, 1969, at Hanover College. However, because of its extreme 
southern location, a later announcement will confirm the time and meet- 
ing place for the Junior Academy. 

Miss Helen Reed announced that she would be distributing sum- 
maries of last year's papers directly following the meeting. The academy 
extended its thanks for her services. 

Dennis Waltke adjourned the thirty-sixth annual meeting of the 
Indiana Academy of Science at 4:25 p.m. 

Respectfully submitted by 

Rachel Koontz, Secretary 
Dennis Waltke, President 


Town Club and School 

Acton Sigma Mu Chapter of FSA, Frank- 

lin Central H. S. 

Bedford Bedford Science Problems Research 

Group, Bedford H. S. 

Bloomington National Scientific Honor Society, 
Bloomington H. S. 

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

Bloomington MSE Academy, University Junior 

Clarksville Clarksville H. S. Science Club, 
Clarksville Junior, Senior H. S. 

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

Columbus Science Club, Columbus Senior 

H. S. 

CrawfordsvilleUp-N-Atom, Crawfordsville H. S. 

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

Albertus Magnus Science Club, 
Central Catholic H. S. 

Phy-Chem Club, Elmhurst H. S. 

Springs Valley Science Club, 
Springs Valley H. S. 

Andrean Biology Club, Andrean 
H. S. 

Fort Wayne 

Fort Wayne 
French Lick 




Mu Alpha Theta, Andrean H. S. 

Biology Club, Lew Wallace H. S. 

Griffith Junior High Science Club, 
Griffith Junior H. S. 

Margaret Richwine 

Paul Hardwick 

Orville Long 

Billie Stucky 

Charles Souers 

Gerald K. Sprinkle 

Sr. Jean Marian 

L. N. Carmichael 

David Wells 
Charles Hames 

Sr. Winifred 

Ruth Wimmer 
D. L. Clark 

Sr. Marie Antoine, 

Sr. Marie Carmel, 

Sr. M. Nadine, 

Lola Lemon 

Fred Meeker 


Junior Academy of Science 27 

Griffith Griffith Senior 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 Donald G. Maines 
Preparatory School Harold J. Sommer 

Indianapolis Science Club, Howe H. S. Jerry Motley 

Indianapolis Kennedy Research Center KRC, Sr. Mary Alexandra, 
Kennedy Memorial H. S. C.S.J. 

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

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

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 0. Bennington 
Granville Wells School 

LaPorte Bi-Phi-Chem Club, La Porte H. S. Frances Mr. Gourley 

Byron Bernard 

Lebanon Junior Explorers of Science, Leb- Tom Ewing 

anon Junior H. S. 

Logansport Lewis Cass H. S. Science Club Raymond T. Kozer 

Madison Madison Science Club, Madison David Dunkerton 

Consolidated High 


Indiana Academy of Science 

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 Sen- 
ior H. S. 

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

South Bend JETS Junior Engineering 1 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 


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, 

Richard Garst 
Fredrick Calhoun 

Sr. Anna Margaret 
Sr. Aloyse 

Biological Survey Committee, J. Dan Webster Chairman 
Publications of 1967-1968 

Dealing with the Flora and Fauna of Indiana 



Vascular Plants: 





Palmer, C. M. 1969. Algal records for three Indiana 
sewage stabilization ponds. In press, Ind. Acad. Sci. 
Proc. for 1968. 

Welch, Winona H. 1969. Hookeriaceae Species and 
Distribution in North and Central America and West 
Indies. Proc. Ind. Acad. Sci. 77:351-356. 

Beesly, L. and Beesly, Adelle. 1969. Lobelias of Franklin 
County and Indiana. In press, Ind. Acad. Sci. Proc. 
for 1968. 

Craske, A. G. Jr. 1969. Ecological site preferences with- 
in two Acer saccharum - A. nigrum complexes found 
in Parke County, Indiana. In press, Ind. Acad. Sci. 
Proc. for 1968. 

Guard, A. T. 1969. Some disappearing plant species. In 
press, Ind. Acad. Sci. Proc. for 1968. 

Humbles, J. 1969. Indiana plant distribution records, 
XX, 1966-68. In press, Ind. Acad. Sci. Proc. for 1968. 

Jackson, M. T. and Allen, P. R. 1969. Detailed studies of 
old growth beech-maple upland forests in Versailles 
State Park. In press, Ind. Acad. Sci. Proc. for 1968. 

Marks, G. C. 1969. The flowering of Lemna minor and 
the establishment of Centourium pulchellum in north- 
west Indiana. In press, Ind. Acad. Sci. Proc. for 1968. 

Petty, R. O. 1969. Pattern of mesic forest succession at 
the western border. In press, Ind. Acad. Sci. Proc. 
for 1968. 

Khalil, G. M., and Cable, R. M. 1969. Germinal develop- 
ment in Philophthalmus megalurus (Cort, 1914) (Trema- 
toda: Digenea). In press, Zeit. f. Parasitenkunde 31. 

Isseroff, H. and Cable, R. M. 1968. Fine structure of 
photoreceptors in larval trematodes. A comparative study. 
Zeitschrift f. Zellforschung 86:511-534. 

Demaree, Richard S. Jr. Indiana State University. 1967. 
Ecology and external morphology of Lernaea cypri- 
nacea. Amer. Midland Natur. 78 :416-427. 

Parker, T. A. 1969. An annotated check list of the spi- 
ders of Indiana. In press, Ind. Acad. Sci. Proc. for 1968. 

Schuder, D. L. 1969. A Japanese weevil, Pseudocneor- 
hinus bifasciatus Ruelefs, discovered in Indiana. In 
press, Ind. Acad. Sci. for 1968. 

Hart, J. T. 1969. A check list of the mosquitoes of 
Indiana with a record of the occurrence of Aedes in- 
firmatus D. & K. In press, Ind. Acad. Sci. Proc. for 

Arnett, Patricia M. 1969. A study of Collembolan popu- 
lations associated with four serai stages leading to the 
beech-maple climax. In press, Ind. Acad. Sci. Proc. for 

Morgan, F. 1969. Effects of effluent on the fish popula- 
tion of Mill Creek, Rochester, Indiana. In press, Ind. 
Acad. Sci. Proc. for 19 68. 



Indiana Academy of Science 


Baker, Mrs. H. A. 1968. Breeding- bird census. Grazed, 
bushy fields and tree-bordered creek. Aud. Field 
Notes, 21:657. 

Indiana Audubon Society Members. 1968. Many titles in 
Indiana Audubon quarterly Vol. 46. 

Smith, Shelia. 1968. Breeding bird census — Suburban 
edge. Aud. Field Notes, 21:673-674. 

Webster, J. D. and West, H. C. 1968. Winter bird popu- 
lation study — Tornado-disturbed beech-maple forest. 
Aud. Field Notes 22:488-489. 


Whitaker, J. O. Jr. 1967. Habitat relationships of four 
species of field mice in Vigo County, Indiana. Ecology 

Whitaker, J. O. Jr. 1967. Hoary bat apparently hiber- 
nating in Indiana. J. Mammalogy 48:663. 

Mumford, R. E. 1969. The hoary bat in Indiana. In press, 
Ind. Acad. Sci. Proc. for 1968. 

All Animals: Gammon, J. R. 1969. The effect of inorganic sediment 

on macroinvertebrate and fish populations of a central 
Indiana stream. In press, Ind. Acad. Sci. Proc. for 

All organisms: Gerwig, F. L. and Crankshaw, W. B. 1969. Effects of 

thermal discharge on the phytoplankton and macroin- 
vertebrates of the Wabash River. In press, Ind. Acad. 
Sci. Proc. for 1968. 

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

Vascular Plants: 

Craske, A. G. Jr. 1968. Ecological site preferences and 
taxonomic differences within two Acer saccharum com- 
plexes found in Parke County, Indiana. M. A. Indiana 

Nichols, S. A. 1968. The phytocenology of selected Indi- 
ana natural areas. M. Sc. Purdue. 


Arnett, Patricia M. 1968. A study of Collembolan popu- 
lations associated with four serai stages leading to 
the beech-maple climax. M. A. Indiana State. 

McWilliams, K. 1968. A taxonomic revision of the North 
American species of the genus Thermonectus (De- 
Jean). Ph. D. Indiana U. 


Bausman, G. P. 1968. A comparative study of the habitat 
and food of sympatric populations of Plethodon dor- 
salis Cope in south central Parke County, Indiana, 
during the fall of 1967. M. A. Indiana State. 

Work in Progress, but not yet Published, Dealing with the 
Flora and Fauna of Indiana- 
Vascular Plants: Petty, R. O. Wabash. Succession in Deciduous forest 
communities — Rhodes woods, Warren Co. Bremmer 
Woods, Tippecanoe Co. Casters Woods, Montgomery 

Platyhelminthes: Cable, R. M. and students. Purdue. Studies on larval 
trematodes of Little Pine Creek and their life cycles. 

Biological Survey Committee 31 

Platyhelminthes: Abbas, M. K. Purdue. Formation of the cyst and factors 
concerning- excystation of the metaceraria of Micro- 
phallus opticus (Trematoda: Digenea). 

Pisces: Whitaker, J. O. Jr., and Wallace, I>. C. Indiana State. 

Continued studies on the fishes of Vigo County, Indi- 

Mammalia: Whitaker, J. O. Jr. Continued studies on the mammals 

of Vigo County, Indiana (parasites, food, habitat, 

Mammalia: Terrel, T. Ij. Purdue. Ecology of the swamp rabbit in 



Fay Kenoyer Daily, Butler University 

Frederick John Allen 
Elgin, Oregon Lafayette, Indiana 

November 21, 1894 June 20, 1968 

Dr. Frederick John Allen was a retired Purdue University Professor 
of Chemistry at his death June 20, 1968. He was noted for his research 
in the purification and properties of the rare gases, krypton and xenon, 
studies of cobalt and its compounds and educational methods in chem- 
istry. He was in charge of writing state tests of Indiana high school 
chemistry pupils for many years. 

Born in Elgin, Oregon, his early education was obtained in that 
state. A bachelor of science degree in agriculture was obtained from 
Oregon State in 1917. He came to Indiana then for work at Purdue 
University where he was an assistant in chemistry 1917 to 1918 and 
1920 to 1921 when he received an M.S. degree. He was an instructor at 
Oregon State College from 1919 to 1922 and at Purdue 1922 to 1929. 
He was an Assistant Professor of General Chemistry from 1930 until 
he retired in 1960. He served during World War I at the American 
University Experiment Station in Washington, D. C. 

He joined the Indiana Academy of Science in 1927 and was honored 
by becoming an Emeritus Member in 1965. During those years he gave 
a number of papers at the Chemistry Section meetings and served on 
various committees. Dr. Allen had been a member of the Academy for 
41 years at his death. 

Other societies to which he belonged were the American Association 
for the Advancement of Science and the Chemical Society. Dr. Allen is 
listed in Indiana Scientists and American Men of Science. 

Dr. Frederick John Allen gave the Indiana Academy of Science 
his loyal, active support for many years earning the respect and grati- 
tude of the society. He was honored in 1935 by election to Fellow. 

John H. Armington 
Leesburg, Indiana Indianapolis, Indiana 

December 10, 1873 June 9, 1967 

Mr. John H. Armington was known as "the dean of Hoosier 
weather forecasters" for his pioneer work in this field. Born in Leesburg, 
Indiana, in Kosciusko County, his schooling was obtained there, and 
later he taught and was principal of a school there. He studied at both 


Necrology 33 

Valparaiso and Tri-State Colleges, but did not receive a degree. His 
interest centered around English, Greek and Latin Courses in college. 
In those early days, a college degree or special course work was not 
necessary for a government job in the Weather Bureau, so in 1902 he 
began his career under Henry J. Cox, a weather forecasting pioneer 
with the Chicago Bureau. In 1914 they were co-authors of a publication 
of the University of Chicago Press called "the Weather and Climate 
of Chicago." 

Mr. Armington was transferred from the U.S. Weather Bureau in 
Chicago to the one in Indianapolis in 1914, and had completed 29 years 
of forecasting at his retirement in 1943. Mr. Armington and his wife 
celebrated their fiftieth wedding anniversary in 1947. They had a son, 
John M., and a daughter, Mary E. Mr. Armington was secretary of 
the Indianapolis Lions Club until the mid-1950's and was cited as its 
outstanding member in 1949. 

He joined the Indiana Academy of Science in 1921 and was interested 
in the Geology and Geography Sectional Meetings. He presented a 
paper on "City Smogs in Periods of General Fair Weather" in 1924 and 
served on the Library Committee for 1931 to 1932. 

Mr. Armington enjoyed a long and productive life reaching 93 years 
of age before his death at home, June 9, 1967. 

Everett Tyler Burton 
Brook, Indiana Battle Creek, Michigan 

April 21, 1893 October 21, 1967 

Everett Tyler Burton was born in the small town of Brook, Indiana, 
April 21, 1893. His education was obtained in Indiana receiving a B.A. 
and M.A. degree from Indiana University in 1920 and 1924 respectively. 
He also attended Columbia University. During 1917 and 1918, he was a 
civil engineer with the U.S. Army serving in France. He became 
a second lieutenant. He was a research engineer for the Bell Telephone 
Laboratories, New York, from 1920 to 1955. After retirement, he 
moved to Battle Creek, Michigan, where he was a registered engineer and 
consultant at the Research Institute of Michigan. 

He joined the Indiana Academy of Science in 1920 while at Indiana 
University, but moved within the year although his M.A. degree was 
completed at Indiana University in 1924. Then later he resided at 
Battle Creek, so was an out-of-state member most of the 48 years 
of membership. 

Mr. Burton also belonged to the American Association for the 
Advancement of Science, Society of Military Engineers, Institute of 
Radio Engineering, Bell Telephone Pioneers and was active in the 
Methodist Church. He is listed in Indiana Scientists and American Men 
of Science. 

34 Indiana Academy of Science 

Everett Tyler Burton was 74 years old at his death October 21, 1967, 
at Battle Creek, Michigan. 

Mable Henniger Esten 
Anderson, Indiana Indianapolis, Indiana 

October 14, 1898 February 29, 1968 

Mable Marie Esten nee Henniger was born October 14, 1898, 
near Anderson, Indiana. She attended rural public schools and graduated 
from Anderson High School in 1916. She graduated from Butler Univer- 
sity in 1922 and obtained an M.A. degree in 1932. 

She taught English and Spanish Courses at Anderson Junior High 
School from 1922 to 1924. During this period, she met and married 
Sidney R. Esten who also taught at Anderson High School. A daughter, 
Virginia, was born in 1924. Both Sidney and Virginia have also been 
Indiana Academy of Science members. Sidney, deceased, was a high 
school biology teacher and Virginia teaches in an Indinapolis High 
School now. 

Due to Sidney's influence, Mable became interested in Botany and 
majored in this subject during graduate study at Butler University. 
She had taken classwork at Indiana University while Sidney returned 
to that school for study. She also gave lectures and led nature hikes 
at Turkey Run State Park from 1927 to 1933. Sidney was naturalist 
there for the Department of Conservation. 

Following graduate work, Mable Esten became an instructor in 
the Evening Division of Butler University in 1932. She continued 
teaching both in the day and evening classes until 1953. She returned 
to teaching in 1958 in the Botany Department of DePauw University 
and taught until 1961. 

Both Sidney and Mable were enthusiastic naturalists carrying their 
professionl interests into their leisure hours. Both were energetic col- 
lectors with Mable's specialty emphasizing button collection showing 
botanical designs and mythical animals. Her activities in this field won 
much acclaim and occupied considerable time after her retirement. She 
was editor of the Button Box, a button society publication, president 
of the Indianapolis and Indiana button societies, author of several 
articles in this field and lectured both locally and nationally. After her 
husband retired, they both contributed valuable material to the dictionary 
of natural sciences published in 1966 by the Compton Co., Division of 
the Encyclopedia Britannica, Incorporated. 

Mable first joined the Indiana Academy of Science in 1931. Although 
her membership was not continuous, she belonged many years and con- 
tributed papers to the programs. She reported on the ''Beech-Maple 
Association at Turkey Run State Park" in 1931 and gave a joint paper 
with Albert G. Dannin on chlorophyll therapy in 1949. 

Necrology 35 

Mable Henniger Esten was honored by election to Phi Kappa Phi, 
national honorary society, and was a past president of the Indianapolis 
Alumni Club of Phi Kappa Phi. 

Mable died quietly in her sleep February 29, 1968, after suffering 
a heart attack, one of a series over the past several years. Despite ill 
health, she had remained active and a cheerful, gracious friend. 

Pearl C. Haslanger 
St. Joseph County, Indiana Mishawaka, Indiana 

September 9, 1906 March 21, 1967 

Pearl C. Haslanger was Mrs. Martin Haslanger of Lakeville, 
Indiana. She taught at Clay Junior High School at South Bend. She 
was born in Union Township of St. Joseph's County, Indiana, and her 
education was obtained in this state earning a degree at Indiana Univer- 
sity. Her husband, Martin, survives as well as a son, Dennis D., a 
student at Purdue University. 

She joined the Indiana Academy of Science in 1962 and was 
interested in the Geology and Geography Section, Botany Section and 
the Junior Academy of Science. 

In addition to her membership in the Indiana Academy of Science, 
Mrs. Haslanger was also a member of Detla Gamma Professional 
Sorority, Zeta Tau Alpha, American Association of University Women 
and past matron of the Lakeville Chapter, Order of the Eastern 

Stricken at 60 years of age, death came after three months of illness 
to Pearl C. Haslanger, teacher, wife, mother and academy member, 
all too soon. 

Frank E. Louraine 
Monroeville, Indiana Arlington, Virginia 

November 12, 1900 March 16, 1968 

Mr. Frank E. Louraine was born on a farm in Allen County near 
Monroeville, Indiana, November 12, 1900, and attended public school 
in Allen County where he developed an early interest in history and 
science. He received an A.B. degree in 1924 and an M.A. degree in 
1926 from Indiana University. He was assistant in the History Depart- 
ment of Indiana University from 1923 to 1924. 

He went to Saginaw Michigan High School from 1925 to 1929 to 
teach history and government. Then in 1929 he was a teaching fellow 
in American History at the George Washington University at Washing- 
ton, D.C. He joined the Library of Congress staff in 1931 where during 
his career he worked in the Stack Service, Study Room Reference Service 

36 Indiana Academy of Science 

of the former Reading Room Division and as Reference Librarian in the 
General Reference and Bibliography Division, Public Reference Section. 
He became Assistant Head of this section in 1961. He was of special serv- 
ice to researchers in American history and visiting scholars in other fields. 
He retired in 1965 and after a long illness, died March 16, 1968, at an 
Arlington Hospital. A brief account of his career was entered in the 
Library of Congress Bulletin and biographical material can be found 
in the third edition of Who's Who in Library Service, 1955. 

Mr. Louraine had been a member of the Indiana Academy of Science 
44 years at the time of his death even though he had resided elsewhere 
most of the years of membership. The field trips which he attended 
at Marengo Cave in 1924, the year he joined the society, and the next 
year at Madison, Indiana, had made a lasting impression on him. Many 
years later, he recognized Dr. Paul Weatherwax on a visit to the 
Library of Congress and recalled these meetings with considerable 
pleasure. Frank Louraine was a fine librarian and graciously accommo- 
dated scholars seeking reference material. His services were sincerely 

Roy McKee 
Jonesboro, Indiana Gas City, Indiana 

August 17, 1903 June 14, 1967 

Roy McKee was an excellent teacher at the Mississinewa High 
School in Gas City, Indiana, and was a leader in the public affairs 
of his community. He was honored at his death by resolutions from the 
Mayor and Common Council of that city praising his work as Civil 
Defense Director, chairman of the committee which organized the Gas 
City Plan Commission and Gas City Board of Zoning Appeals, and his 
service on the Plan Commission after organization. In resolutions 
adopted by the Mississinewa Board of School Trustees, his outstanding 
leadership during 37 years as teacher and community leader was recog- 
nized. Appreciation was expressed for his untiring energy, great insight, 
rare force, fine tact and love of culture and the beautiful. 

He was born at Jonesboro, Indiana, August 17, 1903, where he 
attended public school. He attended Indiana University and graduated 
from Marion College with a B.S. degree in 1931. He also attended 
Purdue and Ball State Universities. 

Roy McKee joined the Indiana Academy of Science in 1932 and 
was interested in the Chemistry, Botany and Zoology Sections. He 
was a sponsor in the Junior Academy of Science for the Gas City 
Science Club for several years after its organization in 1936. He 
served as member of the Council of the Junior Academy of Science 
from 1944 to 1948. 

Roy McKee, good teacher, good citizen and respected Academy 
member succumbed after an eight month illness June 14, 1967. 

Necrology :;7 

Millard S. Markle 
Wayne County, Indiana Richmond, Indiana 

November 26, 1883 October 31, 1968 

Dr. Millard S. Markle was almost 85 years of age when he died 
October 31, 1968. His membership in the Indiana Academy of Science 
had extended over 58 years, a record seldom equalled. He had been 
a fellow in the society for over fifty years and was quite active until 
his death. He had attended the Executive Committee Meeting on October 
18 and was a participant in the Botany field trip in the spring of 
1968, when he was as always affable and helpful. He presented an 
invited paper at the Academy on the history of Plant Taxonomy and 
Ecology in Indiana in celebration of Indiana's Sesquicentennial year, 
1966. As the editor stated, the authors selected to give papers during 
this celebration were chosen because they knew Indiana's history best 
and helped make some of it as well as write about it. Dr. Markle 
presented many papers before the Academy on a variety of subjects 
including plant abnormalities, plant ecology, microtechnique, botanical 
travel reports, teaching bacteriology, and several historical articles. He 
was President of the Indiana Academy of Science in 1945. His presi- 
dential address was entitled "Biology and the Post War World." 

On November 26, 1883, Millard S. Markle was born in Franklin, 
Wayne County, Indiana. This town, 5 miles north of Hagerstown, no 
longer exists. He later lived in Daviess County and attended school at 
Washington, Indiana. An early interest in science developed at high 
school while taking physiology. In the spring of 1904, he was allowed 
to enter Earlham College before his graduation from high school. This 
was to enable him to attend college and teach in public schools in the 
fall and winter terms. He received his B.S. degree in 1910 after being 
tutored to satisfy high school requirements. Some high school credit 
was also received for teaching at Tipton, Webster and Green Forks 
where he was superintendent of schools. He had also attended Indiana 
State Normal from 1901 to 1902. He taught a year at Earlham and 
then attended graduate school at Chicago University receiving an M.S. 
degree in 1913 and a Ph.D. degree in 1915. Dr. Markle's teaching career 
at Earlham continued during graduate school years. He was Assistant 
in Biology from 1909 to 1910, instructor from 1910 to 1912, Assistant 
Professor from 1912 to 1915, Professor of Botany from 1915 to 1918, 
Professor in the newly formed Biology Department from 1918 to 1954, 
and was Emeritus Professor from 1954 to 1968. 

Dr. Markle contributed many illustrated lectures at the Indiana 
Academy of Science meetings before the Plant Taxonomy Section. His 
beautiful color slides for projection were gleanings from interesting 
field trips and travel sequences. One source of color photographs came 
from a program for practical instruction on field trips introduced by 
Dr. David Worth Dennis at Earlham. Dr. Markle continued this policy 
with annual trips for students to the Smokey Mountains, Georgia and 
Florida. He also visited our nation's national parks, Europe and Canada. 


Indiana Academy of Science 


Necrology 39 

He had just visited Africa the summer of 1968 shortly before his 
death. He also spent much time before and after retirement on the 
preparation of microscope slides containing materials for use in teach- 
ing biology. 

Dr. Markle was honored and received positions of trust by the 
action of several other societies during his career. He was Fellow of 
the American Association for the Advancement of Science, President 
and Director of the Indiana Audubon Society, President of the Rich- 
mond Audubon and Nature Club, President of the Richmond Photo- 
graphic Society, and was Superintendent of the Sunday School at 
the West Richmond Friends Meeting. Biographical material appears 
in American Men of Science, Indiana Scientists, Who's Who in Indiana 
for 1957 and the Earlhamite. 

Memorial services at a West Richmond Friends Meeting, November 
10, 1968, gave testimony to the stature of this man. His appreciation of 
good music, good friends and his responsiveness to other's needs were 
recognized. His gentle but forceful life was an inspiration to his 
fellow men and will long be remembered by those fortunate enough 
to know him. 

Neal R. Merritt 
Pawnee City, Nebraska Wabash, Indiana 

July 18, 1899 June 9, 1968 

Neal R. Merritt was an Associate Professor of Geology and 
Geography at Manchester at his death June 9, 1968. There, he had 
attained a unique popularity by building a workshop next to his 
home in Manchester in the shape of a Dutch windmill. He covered 
it with various kinds of rocks collected on his and Mrs. Merritt's 
travels over the United States, Mexico, Canada and Europe. The 
rocks created much interest among his students and with his excellent 
teaching ability generated great popularity for his courses. 

Prof. Merritt was bom at Pawnee City, Nebraska, July 18, 1899, 
but moved to Minnesota where he graduated from high school at 
Hinckley in 1916. He specialized in social science and English in his 
undergraduate study. He attended Duluth State Teachers College in 
1937, received a B.S. degree from Bemidji State Teachers College in 
1941, B.A. in 1952. He received an M.A. from the University of Minne- 
sota in 1950 where he had attended the summers of 1946 to 1950. He 
also studied science, Spanish and the humanities at Manchester College. 
His graduate work was in political science, history and education. 

Neal R. Merritt was a rural school teacher in Pine County, Minne- 
sota, from 1917 to 1936. During this time he was married and had two 
children. He was a grade school principal of Indian School, Onigum, 
Minnesota, from 1937 to 1941; high school teacher and principal at 
Walker, Minnesota, from 1941 to 1946; history teacher and Dean of 

40 Indiana Academy of Science 

Tracy Junior College, Minnesota, 1946 to 1947; graduate assistant at 
the University of Minnesota, the summer of 1947; associate professor 
at Manchester College, Manchester, Indiana, from 1947 to 1968 when 
he died. He taught world civilization, history, geology and geography. He 
was author of several articles in educational publications. 

Prof. Merritt joined the Indiana Academy of Science in 1966. He 
was also a member of the Carpenters and Joiners Union, American 
Legion, Boy Scouts of America, Peru Y.M.C.A., Peru Rocks and Minerals 
Club, Indiana Historical Society, Indiana Veterans of World War I and 
was an ordained elder of the Presbyterian Church. 

It is regretted that we did not have the pleasure of Prof. Merritt's 
membership longer than two years. 

Frank N. Wallace 
Chicago, Illinois Washington, D.C. 

August 9, 1878 May 24, 1968 

Frank N. Wallace, former Indiana State Entomologist, died May 24, 
1968, at Washington, District of Columbia, where he had moved in 
1961. His great ability and the affection with which he was regarded 
contributed to a career unparallelled in this state. He had been state 
entomologist for 43 years under both Republican and Democratic ad- 
ministrations when he retired in 1958. He did more to publicize the 
Department of Conservation than any living person, and was a pioneer 
in developing the state park system in Indiana by working with Col. 
Richard Lieber. He was ever alert to new ways of making our Indiana 
parks more enjoyable to visitors, and publicized them by giving illustrated 
lectures on their many assets. His sharp Scottish wit and engaging 
humor were resourceful in obtaining support in the press and among 
politicians for his programs. In 1955, Frank N. Wallace was awarded 
a certificate by then Governor George N. Craig for "outstanding devo- 
tion to duty." 

Mr. Wallace was born in Chicago, Illinois, August 9, 1879. He 
came to Indianapolis with his family when he was a small child. He 
first attended school in this city where the downtown post office now 
stands on the northeast corner of Ohio and Meridian Streets. He had 
nearly finished high school when he took a bookkeeping job and con- 
tinued his education in accounting at the Y.M.C.A. It was this train- 
ing that opened the opportunity for his career in entomology. Ben 
Douglas, then state entomologist, asked Mr. Wallace to straighten out 
the accounting for his office. Under tutorship of Mr. Douglas, Frank 
Wallace became an authority on insect life. When the Indiana Depart- 
ment of Conservation was established in 1919, Mr. Wallace was made 
the Division of Entomology Director as State Entomologist. In 1956, 
Mr. Wallace was awarded an honorary doctor of laws degree by Indi- 
iana Central College. 




42 Indiana Academy of Science 

Few persons have lived a more colorful life than Mr. Wallace. 
When he was a young man, he went to northern Indiana to work 
on some trees for Gene Stratton Porter, the noted authoress. It was 
there that he met her secretary, Lorene Miller, whom he married. They 
had one son, John H. Wallace, Vice-president of the Lausche Instrument 
Company, now living in Washington, D.C. Mr. Wallace traveled widely 
in the state and country in pursuit of his profession. 

Frank Wallace joined the Indiana Academy of Science in 1920, was 
made Fellow in 1937 and Emeritus Member in 1966. He served on 
the Relation of the Academy to State Committee from 1924 to 1961, and 
he was chairman of the committee most of that time. He has the sincere 
gratitude of the society for insuring publication funds for the Proceed- 
ings during some very difficult years when the state budget was 
trimmed severely. He was President of the Indiana Academy of Science 
in 1940. His presidential address on Japanese beetle control in Indiana 
touched on a subject for which he was commended for significant con- 
trol measures in this field. His work in eradication of the beetle saved 
the state thousands of dollars. He also achieved a significant victory 
over the European corn borer which inflicted heavy losses before it 
was brought under control. 

In addition to the Indiana Academy of Science, Mr. Wallace was 
also a member of the American Association of Economic Entomologists, 
Portfolio, Masonic Lodge and the Indianapolis Press Club. He is listed 
in Indiana Scientists and is the subject of articles in the Indianapolis 
Star (Dec. 19, 1957), Outdoor Indiana (June, 1958), and the Eastern 
Indiana Farmer (November, 1958). 

New Members for 1968 

The following list contains the names and addresses of all new 
members who joined during 1968. 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 

Dr. Dorothy Adalis, Biology Dept, Ball State Univ., Muncie, Ind. 4730G 

Dr. Ernest M. Agee, Dept. Geosciences, Purdue Univ., Lafayette, Ind. 47907 

Miss Judith A. Anderson, 5510 Winston Dr., Indianapolis, Ind. 4622G 

Miss Patricia M. Arnett, 822% W. Mulberry, Kokomo, Ind. 4G901 ZD 

Mr. Frederick K. Ault, Dept. Chemistry, Ball State Univ., Muncie, Ind. 
47306 CDP 

Mr. Ned K. Bleuer, Ind. Geol. Surv., Bloomington, Ind. 47401 GSA 

Mr. Roger D. Burgess, Dept. Physics, Ball State Univ., Muncie, Ind. 47306 

Mr. N. Franklin Burnett, 11610 Crestwood Court, Indianapolis, Ind. 46239 

Mr. John M. Burns, Zoology Department, Indiana University, Bloomington, 
Ind. 47401 Z 

Dr. Robt. L. Carmin, Dean Coll. Sci. & Human, Ball State Univ., Muncie, 
Ind. 47306 G 

Mr. Eddie Cass, Dept. Geography, Ball State Univ., Muncie, Ind. 47306 GL 

Dr. Richard F. Copeland, Dept. Chemistry, Ball State Univ., Muncie, Ind. 
47306 CP 

Dr. T. J. Crovello, Biology Dept., Notre Dame University, Notre Dame, Ind. 
46556 BLT 

Miss Karen D. Curtis, Box 225, RR 1, W. Terre Haute, Ind. 47885 OZB 


44 Indiana Academy of Science 

Mr. John H. Daugherty, Dept. Anat. & Physiol., Indiana University, Bloom- 
ing-ton, Ind. 47401 ZOP 

Mr. Lyndon L. Dean, N. Ind. Public Serv., 5265 Hohman Ave., Hammond, 
Ind. 46325 G 

Mr. George F. Degler, 7939 E. Penway, Indianapolis, Ind. 46226 GLA 

Mr. Ralph M. Dinkel, Dept. Geol. & Geography, Indiana State Univ., Terre 
Haute, Ind. 47809 GSL 

Dr. Heyman C. Duecker, Dept. Chemistry, Marion Coll., Marion, Ind. 46952 

Dr. F. H. Emerson, Dept. of Horticulture, Purdue University, Lafayette, 
Ind. 47907 BO 

Mr. Larry Enochs, 2764 S. Scottland Dr., Columbus, Ind. 47201 GD 

Mr. Albert L. Esterline, 1420 Granville, Muncie, Ind. 47303 EDZ 

Dr. J. A. Gross, Dept. Life Sciences, Indiana State Univ., Terre Haute, Ind. 
47809 RBZ 

Mr. T. E. Habart, RR 3, Box 365, Knox, Ind. 46534 A 

Mr. George W. Harrison, Biology Dept., Taylor University, Upland, Ind. 
46989 ZEB 

Dr. Lester L. Hearson, Dept. Biology, Wabash Coll., Crawfordsville, Ind. 
47933 ZO 

Mr. Maurice E. Heath, Dept. Agronomy, Purdue Univ., Lafayette, Ind. 
47907 SL 

Mrs. Donna J. Howard, Muncie, Ind. 47302 DBH 

Mr. James E. Hughes, 9218 Grace PL, Highland, Ind. 46322 DH 

Lt. JG. G. S. Jones, Medical Ecology Dept., USN Med. Res. Unit 2 BX 14, 
APO Can Francisco, Calif. 96263 ZL 

Mr. Gwilym S. Jones, 337 Hilltop Ln., Wyoming, Ohio 45215 LZB 

SR. Jean G. Jones, Marion College, 3200 Cold Springs Rd., Indianapolis, 
Ind. 46222 CP 

Mr. Raymond Jones, 3465 Riverside Dr., Columbus, Ind. 47201 DG 

Miss Irene Joyce, 246 N. Rensselaer Ave., Griffith, Ind. 46303 D 

Dr. Ralph D. Joyner, Chemistry Dept., Ball State Univ., Muncie, Ind. 
47306 C 

Dr. Ronald E. Kirk, 2707 E. 66th St., Indianapolis, Ind. 46220 Z 

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

Mr. Virgil R. Knapp, RR 1, Box 119, Zionsville, Ind. 46077 E 

Dr. James M. Kortright, Rose Polytechnic Inst., Terre Haute, Ind. 47803 P 

Dr. David C. Kramer, Test Jr. High School, Richmond, Ind. 47374. ZBL 

Mr. Edward R. Lavagnino, Eli Lilly & Co., 740 S. Alabama St., Indianapolis, 

Ind. 46206. C 
Dr. Richard M. Lawrence, Dept. Chemistry, Ball State Univ., Muncie, Ind. 

47306. C 
Mr. Don T. Leonard, Biology Dept., Ball State Univ., Muncie, Ind. 4730G. 

Dr. Ralph A. Llewellyn, Rose Polytechnic Inst., Terre Haute, Ind. 47803. P 
Mr. Francis C. Lundin, Biology Dept., Ball State Univ., Muncie, Ind. 47306. 


New Members 45 

Dr. Wendell P. McBurney, Indiana University, Morrison Hall 103, Bloom- 

ington, Ind. 47401. D 
Mr. Paul T. McKelvey, Logansport High School, Logansport, Ind. 46947. 


Mr. Doren G. Martin, Dept. Anat. & Physiol., Indiana University, Bloom- 
ington, Ind. 47401. RCO 

Dr. Charles E. Mays, Dept. Zoology, Depauw Univ., Greencastle, Ind. 46135. 

Dr. Clyde R. Metz, Purdue University, Indianapolis Campus, Indianapolis, 
Ind. 46205. C 

Miss Margaret E. Meyer, Rt. 1, Corydon, Ind. 47112. BLZ 

Mr. Arthur E. Middleton, Biology Dept., St. Josephs College, Rensselaer, 
Ind. 47978. DOC 

Dr. Arthur Mirsky, Indiana University, 518 N. Delaware, Indianapolis, Ind. 
46204. GLH 

Mr. Frederic Morgan, Biology Dept., Ball State Univ., Muncie, Ind. 47306. 

Dr. William G. Nevill, Purdue University, 1201 E. 38th St., Indianapolis, 
Ind. 46207. C 

Dr. Donald J. Niederpruem, Microbiol. Dept., Indiana U. Med. Center, Indi- 
anapolis, Ind. 46202. ROB 

Mr. Billy E. Norris, 2201 Belmont, Muncie, Ind. 47304. DZ 

Dr. Sidney Ochs, I. U. Med. Center, 1100 W. Michigan, Indianapolis, Ind. 
46207. ZP 

Mr. Robert E. Pace, 24 Chickadee Dr., Terre Haute, Ind. 47803. 

Mr. Thomas A. Parker, Entomology Dept., Purdue University, Lafayette, 
Ind. 47907. ELZ 

Mr. George P. Pollock, 1314 S. Sth St., Terre Haute, Ind. 47803. Z 

Dr. H. W. Reuszer, Dept. Agronomy, Purdue Univ., Lafayette, Ind. 47907. 

Mr. David T. Rice, RR 5, Frankfort, Ind. 46041. OCY 

Mr. Rush L. Robinson, Dept. Anat. & Physiol., Indiana University, Bloom- 
ington, Ind. 47401. ZCM 

Mr. Steve Rudolph, 2800 Capitol Blvd., Evansville, Ind. 47711. COP 

Mr. Richard E. Schaffer, Dept. Anat. & Physio., Indiana Univ., Bloomington, 
Ind. 47401. ZLO 

Mr. Neil Schemehorn, 5265 Hohman, Hammond, Ind. 46325. G 

Dr. Arthur R. Schulz, Metab. Res. Lab., V. A. Hospital, Indianapolis, Ind. 
46202. CHM 

Mr. James D. Schwengel, 1557 S. Plaza Dr., Evansville, Ind. 47715. LZD 

Dr. Gregory E. Shaner, Botany & Plant Path., Purdue Univ., Lafayette, 
Ind. 47905. BLT 

Dr. James E. Shields, 7229 Wynter Way, Indianapolis, Ind. 46250. C 

Dr. Arthur C. Singer, Dept. Life Sciences, Indiana State Univ., Terre Haute, 
Ind. 47803. BZ 

Mrs. Shirley F. Smalley, 119 N. 5th, Spiceland, Ind. 47385. DOC 

Mr. Kenneth A. Smiles, Dept. Anat. & Physiol., Indiana University, Bloom- 
ington, Ind. 47401. OZC 

46 Indiana Academy of Science 

Mr. David E. Smith, 422 Bluebird Dr., Terre Haute, Ind. 47803. CBL 

Dr. Phillip J. Smith, Dept. Geosciences, Purdue Univ., Lafayette, Ind. 

47907. GPS 

Mr. Clifford M. Stamper, Wright State Campus, Colonel Glenn Highway, 
Dayton, Ohio, 45431. AG 

Dr. Larry K. Steinrauf, I. U. Med. School, Indianapolis, Ind. 46207. CPO 

Mr. Bruce N. Storuhoff, Dept. Chemistry, Ball State Univ., Muncie, Ind. 
47306. C 

Mr. Phillip L. Walker, 214 LI. Crawford St., Elkhart, Ind. 46514. AZL 

Dr. Hans W. Wendt, Psychology Dept., Valparaiso University, Valparaiso, 
Ind. 463S3. Y 

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

Mr. Woodrow W. Winstead, Rt. 3, Box 202, Newburgh, Ind. 47630. ODL 

Mr. Chas. J. Zimmerman, Jr., 1207 Ms N. Grant, Bloomington, Ind. 47401. Z 





Muncie, Indiana 
October 19, 1968 

The address, "Urban Geology — A Need and A Challenge", 
was presented by retiring president, Dr. William J. Wayne, at 
the annual dinner meeting of the Academy at the Pittenger 
Student Center on Saturday evening, October 19, 1968. It is 
an excellent statement of the necessity of including geological 
knowledge in many phases of urban planning. Dr. Wayne is 
currently a member of the Department of Geology of the Uni- 
versity of Nebraska. The address by Dr. Robert E. Gordon, 
Professor of Biology and Associate Dean of the College of 
Science at the University of Notre Dame, was given at the 
luncheon meeting on October 19 involving both the Junior 
and Senior Academy members. His subject, "Science, Com- 
munication, and the Critical Mass", deals with the public 
understanding of science as a function of the ability and 
effectiveness of the scientist's communication with the public. 


Urban Geology — A Need and A Challenge 1 
William J. Wayne- 

The Scope of Urban Geology 

Every use man makes of land is affected by the shape of the land 
and by the physical properties of the materials that lie beneath the 
surface. He depends on either the surface or the materials beneath it 
for food, water, and fuel; for building sites, building materials, and 
foundation support; for waste disposal; and for recreation. 

Geology is the study of the earth. It encompasses investigation of 
the surface, the materials beneath the surface, and all the natural 
processes that have produced those materials and landforms. Environ- 
mental geology is one of the names currently in vogue for the specific 
phase of geology that deals with the interrelationships of geologic 
processes, earth materials, and the ways in which man has met and 
used this part of his environment. 

Where man has congregated in large numbers and most extensively 
disturbed natural conditions is where most of the conflicts between man 
and his environment are likely to take place. Thus the geology of man's 
environment becomes most important in and near urban centers; the 
term urban geology is virtually synonomous with environmental geology. 
Urban geology involves the recognition and understanding of those 
geologic processes that continuously work to bring conditions on the 
earth's surface toward a state of equilibrium — the natural forces that 
operate more or less slowly but are powerfully effective in the creation 
of landscapes and the disruption of some of man's works on those land- 

Our population has increased greatly in recent decades, and with 
that population increase our intensive uses of land have also expanded 
greatly. Because of this expansion and the resulting elimination of open 
space surrounding urban centers, we find ourselves having increasingly 
less freedom to make mistakes in the development of land for uses more 
intensive than farming. 

When an error is made in developing a homesite in a rural environ- 
ment, rarely is more than a single structure and one family affected; 
an error of similar magnitude in developing homesites in an urban 
environment, however, can involve many dwellings and cause incon- 

1 Approved for publication by the State Geologist, Indiana Geological 
Survey, Department of Natural Resources. 

2 Indiana Geological Survey ; joined Department of Geology, University 
of Nebraska, Lincoln, in September 19G8. 


50 Indiana Academy of Science 

venience and unnecessary expense to many families. It is therefore in 
and near cities — the areas of large population concentrations — where it 
is of greatest importance that the men who guide our changes in land 
use recognize the ways in which natural forces act upon natural ma- 
terials and upon the works of man. The failure to recognize potentially 
destructive geologic processes can lead to unnecessary expenses in the 
urban and urbanizing areas for engineering works, costly damage to 
structures, and perhaps even the loss of human lives. 

Less spectacularly, damage may be to health, or may be limited to 
the inconvenience of wet basements or backed up drains. Nevertheless, 
as man's use of land expands, the need to have a complete and thorough 
knowledge of the geology of his environment becomes continuously more 
important. The costs of correcting mistakes increase many fold after 
available open land has been used up. 

In comprehensive planning for the future development of a com- 
munity, all the needs and interests of the area must be inventoried and 
evaluated. Ideally, potential problems should be forseen early in the 
planning process so that they can be adequately handled in the ordi- 
nances — zoning and others — that make planning effective. One basic 
phase of a comprehensive planning study — and one that is often neglected 
— is an evaluation of the geologic resources of the planning area. 

The Role of a Geologist 

Geologists are well equipped to contribute to several aspects of 
comprehensive studies in which conflicts of land use frequently arise. 
Unfortunately, geologists are sometimes asked to help explain the cause 
of a problem that might have been prevented if they had been consulted 
before the land was developed. They often find themselves cast in the 
role of trouble shooters and pessimists rather than advisors who can 
advance constructive suggestions that will help determine the optimum 
use of available land. 

Application of geologic study to urbanizing environments requires 
the ability of a generalist who is particularly well versed in geomorphol- 
ogy, engineering geology, economic geology, and hydrogeology, and who 
also has an understanding and appreciation of the principles and admin- 
istration of land-use planning. And he must be able to make the results 
of his geologic studies readily understandable to and usable by the non- 
geologically trained professional planners and citizens of the community 
to whom the decisions regarding urban development are entrusted. 

Reports on urban geology should include a brief but adequate 
review of the general geologic features of the area for which the report 
is prepared (31). The report must be directed, though, toward specific 
geologic phenomena that are likely to be of concern in planning the 
community. Features of the land about which geologists are especially 
well qualified to supply knowledge and evaluation are: (1) economic 
mineral resources and potential; (2) geologic conditions that, if un- 
recognized, could become hazards to property or health; (3) water- 

Presidential Address 


supply potential; (4) waste-disposal sites; and (5) geologic significance 
of outstanding scenic and educationally stimulating natural areas. 

Mineral Resources that Serve Urban Development 

Mineral resources that are used extensively in construction, such 
as sand, gravel, and crushed stone, must be exploited close to their 
markets (Fig. 1). These resources have a large bulk and low value per 
ton and are generally surface mined. Maintenance of a high quality 
supply at a low delivered price is important to the growth of every 
expanding urban area. To keep construction costs low haulage distances 
for aggregates must be short, because much of the final delivered price 
is the cost of hauling (9, 24, 25). 


^"""If '-' 


Figure 1. Gravel pit in suburban area of Indianapolis (13, pi 5A) where 
operation is almost completely surrounded by urban land uses. A worked- 
out part of the pit not visible in this photograph has been reclaimed for 
recreational uses. 

One of the most important pieces of information that a community 
should include in a comprehensive planning study is a map showing the 
distribution of potentially workable reserves of mineral resources. It is 
of no value to permit mineral extraction from land that has no mineral- 
resource potential, yet to restrict the industry from land that does. Only 
after the availability of the resource is known can a planner evaluate a 
particular area of a surface mineral resource and recommend a zoning 
ordinance that reflects that evaluation. Such a study was prepared for 
Marion County, Indiana, in 1958 (12) as a by-product of a county 
mapping project (13) and was used in designing zoning regulations for 
the county. 

52 Indiana Academy of Science 

Among the major objections voiced by residents of many com- 
munities to the opening or continued operation of gravel pits, clay pits 
or crushed limestone quarries are the traffic, the dust of processing, the 
noise and rocks of blasting, and the resulting wasteland they must live 
with after the resource has been worked out (26). Some mineral-resource 
operators have become increasingly aware of this criticism in recent 
years, and many of them are becoming sensitive to the desires of their 
communities that they leave the worked-out land in a readily usable 

Mineral producers should be invited to participate in the develop- 
ment of operating standards to control traffic, noise, and dust that both 
they and their neighbors can accept and of subsequent land-use plans 
for the area when they leave it (1). The concept of sequential land use 
applies particularly well to the surface-mined bulk mineral commodities 
used in construction. While the deposit is being worked the land can be 
shaped according to a predetermined design, so that it will fit well into 
a second planned use after the resource has been worked out and the 
equipment removed (21). Reclamation according to such a plan is much 
less expensive than reclamation after abandonment. In many places 
the graded and shaped abandoned pit or quarry will have great value 
to the community as a recreation site, or as building lots, or an industrial 
site; thus it can be made into a desirable and productive area rather 
than a wasteland that remains an eyesore or health hazard (3). 

Natural Hazards 

Many millenia are needed for natural processes to create landscapes. 
Landscape-producing forces work in small increments, however, and 
only small amounts of time are required for some of these to take place. 
The force of earth or rock moving across an unstable slope or of flood 
water passing down a valley is great, and where the works of man 
happen to stand in the way, they may be damaged or destroyed. 

Some natural hazards to life or property cannot be predicted well 
enough to let us avoid them entirely — the path of a particular tornado, 
for example. Many "accidents of nature" that result from failure to 
understand some fundamental geologic processes, however, can be rec- 
ognized by an alert geologist and their potential for damage forecast so 
that land uses and construction standards can be designed to reduce or 
eliminate the danger. 

From the earliest of civilization, man has used rivers for transpor- 
tation, water supply, and waste removal. Consequently he has found the 
land along the rivers desirable places to build communities in spite of 
the knowledge that high water would come regularly; he accepted this 
inconvenience for the advantages of being able to use the river the rest 
of the time. 

The flood plain, though, is the domain of the stream that built it. It is 
the relief valve of the river — the place where the excess water can spread 

Presidential Address 


out and slow down when upstream areas and tributaries deliver more 
water than the channel can carry. When man forgets this or fails to 
recognize it, he and his works can be damaged. 

We no longer use any but the largest rivers for transportation. We 
have, however, inherted the floodplain locations and have developed them 
even further. One of the results of urban expansion on floodplains has 
been an increasingly great property loss and inconvenience each time a 
heavy runoff causes the land to be inundated. Increased urbanization 
upstream in the drainage basin also increases the runoff rate (11, 19). 
Thus we have had to design and build expensive flood control works to 
protect our investments from the inevitable high water. 

Flood plains are underlain by sediments dropped by a river in 
flood. They are a normal unit on most geologic maps; therefore their de- 
lineation is one of the contributions of a geologic study to the planning 
process. Land-use regulation that restricts construction of damageable 
structures from areas of flooding is a far less expensive way of reducing 
future flood damage than is building more and bigger retaining struc- 
tures and levees. Identification of flood plain land by geologic or soils 
mapping permits it to be zoned as future open land. 

Gravity and water combine to produce downslope movement of 
masses of loose rock or soil on many hillsides. The degree of stability 
or instability of a particular slope is largely a factor of both steepness 
and moisture content and the kind of material that underlies it. For 
example, slopes of 1:1 are generally stable in the mudstones of Morgan 
County and western Brown County, Indiana; slopes of 2:1 are normally 
stable in unweathered young glacial till of central Indiana; but slopes 
of 3:1 are required for stability in the thick weathered part of the 

plant fragments 
cherl-y residuum 

thin-bedded limest-one 

Figure 2. Terain sketch and diagram of the geology along State Road 46 
in Owen County where mudflows and slump have resulted from an over- 
steepened slope and a perched water table held by a thin Pleistocene clay 


Indiana Academy of Science 

older glacial tills of southern Indiana. Where unusual conditions exist, 
such as a clay bed that serves to inhibit downward movement of moisture 
within a sequence of silty glacial sediments (Fig. 2), even more gentle 
slopes may develop. 

Slopes that seem to be stable under natural conditions may become 
unstable if moisture content, loading, or steepness should be increased 
through urban development. Such an alteration is likely to result in 
slumps, slides, mudflows (Fig. 3), and, in areas of bedrock, rock falls, 
and rock slides. At the least, such mass wasting induced by changes in 
slope equilibrium is likely to bring on expensive maintenance problems, 
such as removal of debris from the base of road cuts (Fig. 3) or re- 
building retaining walls. Where structures are built on such slopes, 
damage or destruction can be extensive. In addition to the natural 
phenomena that can take place, recent studies in Illinois (34) have 
shown that the addition of detergents, such as those found in laundry 
wastes and septic tank effluent, to unconsolidated clayey sediments will 
decrease the strength of the material and increase its tendency to move 

Most slopes that are likely to become unstable can be recognized 
in the field by a geologist who is trained in their evaluation; the local 
significance of this natural process should be reviewed in every geologic 
study for land-use planning (18, 28). 

Not all land underlain by sensitive materials is on slopes, however. 
Areas underlain by muck, peat, marl, and other soft sediments that ac- 

Figure 3. Mud flow on road cut in weathered Illinoian till along State 
Road 37 near Morgan-Monroe county line. The high clay content of the 
weathered till prevented it from remaining stable at the original cut 
dimensions, although a similar cut in unweathered till probably would 
have remained stable at that slope. 

Presidential Address 55 

cumulated under conditions of ponding but are now above water level 
are outlined on those geologic maps that show surface materials in 
detail. Such sediments are unusually common in Indiana in resort areas 
around the natural freshwater lakes, as well as in some other parts of 
the state where bodies of water have been completely filled by sedi- 
ments. These materials are not stable for foundations, and normal 
construction procedures cannot be used if they are developed intensively 
for urban uses. The high water table normally present would also 
create drainage and waste-disposal problems. Recognition and delinea- 
tion of this material in the planning process is important if the land 
is to be used without danger or damage to its occupants. 

Underground mining produces underground void space. Abandoned 
mine openings collapse and cause minor subsidence at the surface. Maps 
of underground mines on file in the offices of the Indiana Geologic 
Survey provide for Indiana the kind of information needed by planners 
who would avoid such land for construction until it has again become 

Water Supply 

An automobile in every garage and electric power for every home 
made possible the development of large residential housing additions 
far from the edges of cities that would normally supply the utility needs 
of large numbers of families. Subdivisions have been created where 
each home has a private water supply and an electric pump to deliver 
the water to the home. Not all attractive home sites in Indiana have 
enough available ground water to supply a private home, though. And 
construction of several rural schools has been well along before anyone 
realized that some water supply other than a well drilled on the school 
grounds would have to be found before the school could open for classes. 

An evaluation of the ground-water resources of the planning area 
should be included as a part of a comprehensive plan and should be 
available for the use of planning commissions in every urbanizing area. 
Hydrogeologic maps, which provide such an evaluation, can be prepared 
by a geologist, using the basic data derived from a geologic map and 
data on existing water wells. In Indiana generalized maps of this kind 
suitable for county and city planning purposes are being prepared by 
geologists in the Division of Water of the Department of Natural Re- 
sources (27). 

Not all impoundments hold enough water to become ponds or lakes. 
Artificial lakes, both large and small, have been built over materials 
that allow the water to leak out as fast as it runs in as well as in places 
that are watertight. A geologic report for planning purposes would out- 
line those places where high leakage rates could be expected and areas 
where lakes and ponds can be built successfully. Had such information 
been available and in use by the plan commissions of Indiana cities 
and counties, many investments and tax dollars of Indiana citizens could 
have been saved during the past quarter century. 

56 Indiana Academy of Science 

Waste and Refuse Disposal 

Different geologic materials and the soil profiles developed on them 
have different capabilities for absorption and transmittal of moisture. 
Although detailed planning for septic tank disposal fields probably is 
better done from a modern soils map and field studies, broad aspects of 
planning for on-site disposal of liquid household wastes can be done 
readily from the data available in a geologic study. 

Movement of liquid wastes from the soil downward to the water 
table is an aspect of sewage disposal, though, that requires the attention 
of a geologist. Surface water that seeps downward through soil and 
highly permeable materials such as sand or gravel and fractured, weath- 
ered, or cavernous rock is likely to carry with it surface contaminants. 
Where sewage and other liquid wastes are disposed of in the soil, some 
of them are likely to be flushed downward, particularly during periods 
of heavy rainfall. Runoff from livestock feed lots is another source of 
concentrated contaminants that has entered and damaged some ground- 
water aquifers in Indiana. 

In some parts of Indiana, where closely-spaced houses depend on 
both water wells and septic tanks, water supplies high in coliform 
bacteria are not unusual. Even though the bacteria may be filtered or 
have time to die before reaching a well intake, some wells have been 
seriously affected by a high nitrate content, which may cause illness, 
or by other chemicals or detergents that manage to reach an aquifer. 
A geologic report to a plan commission should outline areas where 
migration of liquid wastes is likely to damage nearby water supplies. 

Septic tank disposal fields do not all function effectively — many are 
open to the surface. Nutrients from this effluent can cause very rapid 
eutrophication of lakes downstream, as can barn lot and pasture runoff. 

When solid wastes are made part of the earth as in a dump or a 
sanitary landfill, they begin to undergo the same processes as do natural 
earth materials. Some of the rainwater that lands on the surface 
permeates the earth and passes through the soil and rocks on its way 
to a discharge area or to the water table. While it percolates through 
earth materials it dissolves any substances that may be soluble and 
carries the leachate away as part of the ground water. After it migrates 
beyond the limits of the landfill the leachate from a sanitary landfill 
can be expected to behave as would any vadose or phreatic water in 
the same geologic environment. 

Our present knowledge about leachate migration is too meagre to 
permit us to speak in authoritative terms, although landfills that are 
located in impermeable or slowly permeable materials such as shale, clay, 
or clayey till or are separated geologically from an aquifer by such 
materials probably are unlikely to cause any contamination of ground 
water. Landfills in permeable materials, though, particularly limestone 
or dolomite, are likely to yield a leachate that will migrate rather rap- 
idly and may cause considerable damage to underground water sources 

Presidential Address 57 

(5, 20, 33). Areas geologically favorable and unfavorable for solid waste 
disposal sites should be outlined in every geologic report prepared as 
part of a comprehensive planning study (16). 

Natural Areas 

Where people are, they are expected to go to school, and they want 
to recreate. Many outstanding scenic areas or unusual outcrops that 
have geologic significance have been lost to recreation or education be- 
cause they were not recognized by those who plan future land use, 
although they may have been well known to geologists, ecologists, and 
naturalists for a long time. Some of these areas, if the land is to be 
most advantageously used, probably should be considered for preserva- 
tion as natural areas or for development into park sites. The geologist 
is remiss in his responsibility if he neglects to call attention to such 
areas so that the planner can understand their significance before they 
have been overwhelmed by urbanization. Once destroyed, they cannot 
be reclaimed (15). 

Programs of Research in Urban Geology 

Less than a decade ago, few geologists and fewer planners seemed 
to be aware of ways in which geologic data could be applied to land use 
problems (29). More recently, though, the United States Geological 
Survey has recognized the need to provide geologic data for use in land 
use planning (17). Several state geological surveys, most notably those 
of California and Illinois, have also undertaken studies intended to 
supply geologic information to planners (2, 6, 7, 10). The Indiana 
Geological Survey has provided reports to a few plan commissions or 
their consultants on request during the past decade; some of the reports 
have been published (8, 12, 30), and others are available only as file 

Some of the data needed to prepare geologic reports for use in 
comprehensive land-use planning studies of rapidly urbanizing parts of 
Indiana have been acquired by Indiana Geological Survey geologists as 
part of other studies during the past 15 to 20 years. Several specific 
new research programs will be needed, though, if our geologists are 
going to be able to answer the kinds of questions that we can now 
anticipate. At this time I would like to suggest the following program 
of research in urban geologic studies for Indiana: 

1. The preparation of county or urban community geologic reports. 
— Geologic reports must be written specifically for use by a city, county, 
area, or metropolitan plan commission that is developing a compre- 
hensive study of its area of jurisdiction and must be directed to that 
audience. A standard geologic report does not provide the needed in- 
formation without interpretation. Table 1 is an outline that has served 
well in the preparation of such reports and can be adapted to most 
areas in Indiana and other midwest states. 


58 Indiana Academy of Science 

Indiana Geological Survey geologists have prepared several county 
or community reports at the request of plan commissions or their 
consultants during past years, but many requests for information were 
received too close to the planning organization's deadline to permit the 
geologist to do more than quickly draw together a report based on 
information on hand but acquired for some other purpose. Unfortunately 
such reports will be as variable in quality and in value to the user as 
the amount and quality of data on hand for the geologist to use in 
their preparation. A special research program could anticipate areas 
for study far enough in advance that most rush jobs could be avoided 
when specific requests are received. 

Geologic mapping for urban studies must be done on as large a 
scale as is practicable. At one time, maps prepared at 1 inch to 1 mile 
(1/63,360) were considered adequate and maps at 1/250,000 and 
1/125,000 were thought satisfactory for many purposes. Urban studies 
will require greater detail, however. California geologists are mapping 
some urban areas on a scale of 600 feet to the inch (2), and the flood- 
way mapping program of the U. S. Army Corps of Engineers is being 
done on an even larger scale. 

County mapping for land-use studies probably will be satisfactorily 
presented on a map with a scale of an inch to the mile. Maps for a pilot 
study of Madison County, Indiana are being prepared at an inch to the 
mile although they were compiled at a scale of 1/24,000 (32). Highly 
urban areas as well as complicated or problem areas probably should 
be presented on a larger scale, however. The study should include a 
basic map showing the distribution of surficial geologic units described 
in a non-technical style and must be supplemented by a series of 
geologic planning maps on which attention is called to specific aspects of 
geology as related to land use (Table 1). These maps would include 
waste disposal, ground-water potential, slope stability, economic ma- 
terials, and other subjects that may be appropriate (10, 16, 22). 

2. Geologic studies of ground-water contamination. — Small scale 
studies of some wells in the limestone terrane of south-central Indiana 
a few years ago led to a recommendation presented in Bulletin S. E. 15 
of the State Board of Health that much of the bacterial contamination 
of wells in such regions can be reduced or eliminated entirely by a more 
positive seal to prevent the entrance of surface water and storm water 
into the open annular space around the casing of a well (Fig. 4). Re- 
search in urban geologic problems should include additional studies of 
this kind in other geologic environments, undertaken on the university 
level or cooperatively by the State Board of Health, Division of Water, 
and the State Geological Survey. 

3. Migration of landfill leachates. — Although contaminants are 
known to have reached the water table and to have migrated away from 
a landfill site under some conditions (4), few studies have been under- 
taken to determine the significance of different geologic conditions on 
the movement of leachate from sanitary landfills. Such a study has 

Presidential Address 


been proposed on a state level in Indiana, to be done by three state 
agencies jointly. Detailed geologic and geophysical investigation of a 
study area in each of six different geological environments will be the 
responsibility of the Indiana Geological Survey. A water sampling 
program and water-quality analysis in and around each pilot site and 
the hydrology and evaluation of leachate migration and dilution will be 
undertaken by the State Board of Health and the Division of Water. 

Space filled with 
cement grout. 


Space open or 
Casing .^ fj Med with soi | 



Figure 4. Many water wells in the areas of shallow bedrock in Indiana 
have been completed without sealing the annular space around the casing 
of the well to a great enough depth to keep seepage of contaminated 
vadose water from entering the hole. If the annular space has been left 
open or has been backfilled loosely with soil or with cuttings, it remains 
a conduit through which surface water can drain downward rapidly and 
enter water-producing zones. Wells in which this space has been filled 
with cement grout or, under certain conditions, with thick drilling mud, 
and in which the sealed casing extends downward far enough to keep 
out surface waters in permeable or cavernous rock, are not likely to be 
damaged by polluted surface water. 

4. Reclamation of quarried wasteland. — Worked-out and abandoned 
land from which bulk mineral-industry commodities have been removed 
is a wasteland near many urban areas. Unconsolidated materials, such 
as gravel and sand, can be reshaped by a subsequent land owner if 
necessary, but the sand and gravel industry has started a continuing 
program of research to find better and more economical ways of pre- 
paring the land for re-use (14). Land underlain by consolidated rocks 
such as limestone are much more difficult to reclaim for subsequent 
productive use. A study of the rock properties that affect reshaping 
land as well as other possible techniques of converting worked-out land 

60 Indiana Academy of Science 

at a reasonable cost might reduce the time involved for sequential bene- 
ficial use. 

5. Damages from natural hazards of geologic origin. — Though much 
less spectacular in Indiana than in California (18) or Alaska (23), 
natural hazards exist in the state, and damages have taken place. Both 
field and laboratory studies of the geology and geometry of stable and 
unstable slopes would help in recognizing the conditions that produce 
instability of earth materials and in predicting more accurately where 
and under what conditions problems will occur in each geologic environ- 

Such a research program as this will have to be undertaken as 
soon as possible if we are to be able to apply our geologic sophistication 
to aid in the solution of these urban problems where geologic data can 
help. The longer we wait to start, the longer we will find ourselves 
forced to answer questions without sufficient information. If planners 
are to call on geologists for help, the geologists must be in a position 
to supply it. 

The Training of Geologists for Urban Studies 

As a newly-developing use of geologic information, urban geology 
requires that the geologist limit his presentation and evaluation to 
those aspects of geology that apply directly to land use problems. It 
is an applied field of geology in which the practitioner needs to under- 
stand not only the geologic aspects of the problems but must also be 
aware of the principles and techniques of the land-use planning profes- 
sion. Until 1967 few, if any, colleges and universities presented course 
material to train geologists to consider this field for employment. Some 
of the geologists who applied their knowledge to the solution of urban 
problems had picked up their background in the needs of communities 
through service on local planning commissions, and others did so 
through encountering the problems in the course of routine geologic 
studies and becoming interested in solving them. Most frequently the 
man was an engineering geologist, a ground water geologist, or an 
economic geologist. 

Within the past year, a few universities have offered courses in 
urban and /or environmental geology. At Indiana University I presented 
a series of non-credit evening lectures on the subject in the spring 
semester of 1967 and offered a 3-hour credit course in it in the spring 
semester of 1968. At the same time Dr. James Hackett left the Illinois 
State Geological Survey to set up a graduate program in environmental 
geology at Virginia Polytechnic Institute. Paul Hilpman of the Kansas 
Geological Survey began teaching a two-semester course in urban 
geology at the University of Missouri in Kansas City in 1967, and an 
evening course was offered at Oregon in the spring of 1968. The Uni- 
versity of Nebraska also has just added to its curriculum a one-semester 
course in the subject to be taught for the first time in 1969-70. 

Presidential Address Gl 

The Indiana course, as well as the one to be offered at Nebraska, 
was intended to introduce students to the ways in which geologic data 
can be used to help solve problems in land-use planning. It was designed 
to follow a course in physical geology and to present the subject to 
undergraduate majors and minors as well as provide a course in land 
use applications of geology to non-majors. At Indiana it was especially 
popular with graduate students in earth-science education. Those who 
had completed and done well in only one course in geology were able to 
complete the course successfully, but the amount of classroom partici- 
pation was directly related to the geologic background of the indi- 
vidual students. No textbook exists that is suitable for this course, but 
the use of selected readings, many of which are cited here, provided 
material that was current as well as appropriate. 

For a geology undergraduate who would like to direct his pro- 
fessional efforts into this field, the normal B. S. requirements in geology 
should be met if possible, but electives would have to include such 
courses as urban geography, land-use planning, and other courses in 
urban studies. It would be possible at Indiana University to qualify for 
a certificate in urban studies along with a strong A.B. degree in geology. 
On the graduate level a seminar in urban geologic problems coupled 
with regular advanced courses in hydrogeology, economic geology, 
limnology, and engineering techniques in geology would provide the 
required background to become a successful urban geologist. 

A non-geologist who enters any phase of land-use planning should 
anticipate the inclusion of a course in urban geology about his fourth 
year, after he has acquired an adequate background in beginning geol- 
ogy and related courses in geography and allied subjects to enable him 
to master the material presented. He would not, of course, be trained 
to make geologic studies, but should at the conclusion of a course in 
urban geology be able to read a geologic report with understanding and 
to recognize the need for a geologic study in the planning process. 


Urban land uses are rapidly converting open land into intensively 
used land. Generally, in planning those uses, a comprehensive study has 
included everything except an evaluation of the land in three dimensions. 
Geologists are eminently well equipped to supply the missing data, but 
they must also be aware of the needs of the urban community and the 
kinds of information needed by planners if they are to write reports 
usable by planners. Urban and environmental geology is a newly ex- 
panding field in the use of geologic data to help solve some urban 
problems in the planning stage. It is likely to become a part of the 
curriculum in many college geology departments within the next few 

62 Indiana Academy of Science 

Literature Cited 

1. Ahern, V. P. 1964. Land-use planning- and the sand and gravel pro- 
ducer. National Sand and Gravel Association, Silver Spring, Maryland 
30 p. 

2. Campbell, Ian, and B. W. TROXEL. 1965. Geologic hazards. California 
Div. Mines and Geology, Mineral Info. Service 18:161-163. 

3. Carnes, W. G., and others. 1966. Landscape reclamation. Landscape 
Architecture, January 1966 (9 papers on reclamation of worked-out 

4. Cartwright, Keros, and M. R. McComas. 1968. Geophysical surveys in 
the vicinity of sanitary landfills in northeastern Illinois. Groundwater 
0:22-30, 8 figs. 

5. Deutsch, Morris. 1963. Groundwater contamination and legal controls 
in Michigan. U.S. Geol. Survey Water-Supply Paper 1961. 79 p., 23 figs. 

6. Flawn, P. T. 1965. Geology and urban development. Baylor Univ. Geol. 
Studies, Bull. 8:5-7. 

7. Frye, John. 1967. Geological information for managing the environment. 
Illinois State Geol. Survey, Environmental Geology Notes 18. 12 p. 

8. Gates, G. R. 1960. Geologic considerations in urban planning for Bloom- 
ington, Indiana. Indiana Geol. Survey, Rept. Prog. 25. 21 p., 1 pi., 1 

9. Goldman, H. B. 1959. Urbanization and the mineral industry. California 
Div. Mines, Mineral Info. Service. 12(12) :l-5, 9 figs. 

10. Hackett, J. E. 1968. Geologic factors in community development at 
Illinois. Illinois State Geol. Survey, Environmental Geology Notes, 
No. 22. 16 p., 4 figs. 

11. Harris, E. E. and S. E. Rantz. 1964. Effects of urban growth on 
streamflow regime of Permanente Creek, Santa Clara County, Cali- 
fornia. U.S. Geol. Survey, Water Supply Paper 1591-B. 18 p. 

12. Harrison, Wyman. 1960. A special report on the geology of Marion 
County, Indiana. Metropolitan Planning Commission of Marion County, 
Mineral Res. Rept. 1. 53 p., 1 pi., 13 figs., 4 tables. 

13. Harrison, Wyman. 1963. Geology of Marion County, Indiana, Indiana 
Geol. Survey, Bull. 28. 78 p., 5 pis., 11 figs., 4 tables. 

14. Johnson, Craig. 1966. Practical operating procedures for progressive 
rehabilitation of sand and gravel sites. National Sand and Gravel 
Assn. Project No. 2. 75 p., 71 figs. 

15. Lindsey, A. A. 1968. Indiana's new system of scientific areas and nature 
preserves. Proc. Indiana Acad. Science, 77:75-83. 

16. McComas, Murray. 1968. Geology related to land use in the Hennepin 
region. Illinois State Geol. Survey, Circ. 422, 24 p., 10 figs., 2 tables. 

17. McGill, J. T. 1964. Growing importance of urban geology. U.S. Geol. 
Survey, Circ. 487. 4 p. 

18. Morton, D. M., and Robert Streitz. 1967. Landslides. California Div. 
of Mines and Geology, Mineral Info. Service. 20:123-129, 135-140. 

19. Savini, John, and J. C. Kammerer. 1961. Urban growth and the water 
regimen. U.S. Geol. Survey, Water-Supply Paper 1591-A. 42 p. 

20. Scheaffer, J. R., Berndt von Boehm, and J. E. Hackett. 1963. Refuse 
disposal practices in northeastern Illinois. Northeastern Illinois Metro- 
politan Area Planning Commission, Tech. Rept. 3, 72 p., 13 figs. 

21. Schellie, K. L., and D. A. Rogier. 1963. Site utilization and rehabilitation 
practices for sand and gravel operations. National Sand and Gravel 
Assn. Spec. Rept. 80 p., 47 figs., 4 tables. 

22. Schltcker, H. G., and R. J. Deacon. 1967. Engineering geology of the 
Tualatin Valley region, Oregon. Oregon Dept. Geology and Mineral 
Industries, Bull. 60. 103 p., 4 pis. 45 figs., 5 tables. 

Presidential Address G3 

23. Schmidt, R. A. M. 1964. Geology in a hurry. Geotimes 0(3):13-15. 2 figs. 

24. Sheridan, M. J. 1967. Urbanization and its impact on the mineral ag- 
gregate industry in the Denver, Colorado, area. U.S. Bur. Mines, Info. 
Circ. 8320. 53 p., 28 figs. 

25. Stephenson, R. C., and others. 1966. The interaction of urbanization 
and the mineral industries. Ohio State Univ. Nat. Res. Inst. Ann. 
Symposium 1965 (collection of 10 papers). 

26. Stollman, Israel. 1962. Land-use control in the surface extraction of 
minerals, Part 1. Am. Soc. of Planning Officials, Planning Advisory 
Service, Info. Rept. 153. 17 p. 

27. Uhl, John. 1966. Water resources of Johnson County. Indiana Dept. 
Natur. Res., Div. Water. 

2 8. Varnes, D. J. 1950. Relation of landslides to sedimentary features, p. 
229-246. In Trask, P. D., ed., Applied Sedimentation. New York, J. 
Wiley & Sons, Inc. 

29. Wayne, W. J. 1960. Geologic contributions to community planning. 
Unpublished manuscript of paper read at Am. Assn. Advancement Sci. 
meeting, Dec. 1960, 20 p. 

30. Wayne, W. J. 1968a. Geology of Morgan County (unpublished). Morgan 
Co. (Indiana) Plan Commission. 25 p., 2 pis., 3 figs. 

31. Wayne, W. J. 1968b. Urban geology as a necessity. Indiana Governor's 
Conf. on Nat. Resources, Feb. 27, 1968. p. 19-21. 

32. Wayne, W. J. in preparation. Urban geology of Madison County. Indiana 
Geol. Survey, Special Report. 

33. Weaver, Leo. 1961. Refuse disposal— its significance, p. 104-110. In 
Ground Water Contamination Symposium volume. Taft Sanitary Eng. 
Center, Tech. Rept. W 61-5. 

34. White, W. A. and S. M. Bremser. 1966. Effects of soap, a detergent, and 
a water softener on the plasticity of earth materials. Illinois State 
Geol. Survey, Environ. Geology Notes 12. 15 p. 

Table 1. Outline of a geologic report to be used as part of a 
comprehensive planning study 

1. Introduction 

a. Location and limits of area 

b. Purpose for which report is written 

c. Nature of the data available for preparation of the report (whether 
reconnaissance or detailed in nature, extent of field observations 
by author, how compiled, availability of reference material) 

2. Summary of any highly significant problems 

3. Regional geology — brief statement about geological setting of area 

4. Geology of area that may apply directly to land use studies 

a. Topographic elements (description of major terrane features, 
major drainage lines, and upland areas; origin of features may be 
mentioned but only to extent needed to aid in understanding 

b. Earth materials 

1.) Distribution, characteristics of consolidated rocks 
2.) Distribution, characteristics, thickness of unconsolidated ma- 
terials, including floodplain sediments 
(For each unit at the surface, treatment should include such 
characteristics as drainage, lithology, permeability, attitude, joint- 
ing, bedding, bearing strength, and any unusual conditions such 
as planes of weakness, zones of uneven stability, high temporary 
water zones, solutionally enlarged openings. The major character- 
istics of each geologic unit used can often be conveniently sum- 
marized in a table or chart.) 

64 Indiana Academy of Science 

c. Geological processes of importance 

1.) Weathering (type, products, depth, significance) 

2.) Mass wasting 

3.) Stream erosion and deposition 

4.) Shoreline erosion and deposition 

5.) Earthquakes 

5. Mineral resources 

a. Distribution of known and potentially exploitable mineral resources 

b. Relationship of each local resource to local and regional economy 

c. Reclamation of worked-out and abandoned surface mines 

6. Water resources 

a. Potential ground-water aquifers of area 

1.) Evaluation of each unit (note outstanding features, good and 
bad, regarding reliability, quantity, quality of water) 

b. Surface water resource potential 

1.) General statement on streamflow 
2.) Prospective impoundment sites 

7. Waste disposal 

a. Liquid wastes — septic tank method of on site disposal — evaluation 
of each geological unit and the effect on ground water and surface 
water of disposal over it. 

b. Solid wastes — geological significance of sanitary landfills; evalua- 
tion of the geologic environments in the study area and the effect 
landfills in each one may have on local water quality. 

8. Areas likely to cause unusual problems or hazards to urban develop- 

a. Flood plains 

b. Areas of potential slope failures and unstable foundation materials 

c. High water table 

d. Earthquake potential 

9. Geologic features of unusual scenic attractiveness or educational value 
10. References 


1. Basic maps 

a. Areal geology 

b. Surface geomorphic elements 

c. Bedrock topography (in glaciated areas) 

d. Bedrock geology (if different from areal geology) 
c. Bedrock structure where significant. 

2. Applied maps 

a. Natural hazard map 

1.) Floodplains, landslide or slump areas, unstable foundation areas, 
high water table areas. 

b. Thickness of unconsolidated materials 

c. Mineral resource map(s) 

1.) Areas likely to contain commercially exploitable resource, distin- 
guished from total area where that resource is at or near the 

2.) Oil and gas map, including extent of current and abandoned 
pools and fields, and possibilities for underground storage 

d. Water resource maps 

1.) Ground-water availability and potential 

2.) Surface water development areas, impoundment possibilities 

e. Areas suitable for surface waste disposal 
1.) Sanitary landfill areas 

2.) Septic tank development (Internal drainage characteristics of 
surficial units and the soils developed on them) 

f. Potential natural areas and recreation areas based on geologic 
(geomorphic) features 

Science, Communication, and the Critical Mass 

Robert E. Gordon 
University of Notre Dame 

In recent years, science has been confronted with a crisis in com- 
munication. To the average scientist this crisis has meant that the 
scientist to scientist transfer of information has bogged down. A number 
of programs to relieve this situation have been proposed and are, de- 
pending on the field, in various stages of implementation. 

I suppose that those of you who have followed the efforts of biology 
to formulate a national system, and know of my activity in this area, 
have already concluded that a biologist is about to speak to you about 
biological communication. 

Stand relieved — today I wish to talk about communication — but not 
that of scientist to scientist. 

There is an equally urgent problem which if not resolved may 
have much more deleterious effects than that posed within the scientific 

I refer to communication from the scientific community to the non- 
scientist. This problem is of equal importance to all fields of science — 
a fact which is easily demonstrable. A significant portion of the long 
range solution to this problem rests in large measure in the hands of 
that part of the scientific community concerned with education. 

Previous to World War II, research in science was the activity of 
a dedicated few — a relatively smaller number of practitioners than is 
currently the case. 

The public image of research was often revealed by caricatures in 
the popular press — in the case of biology often by a picture of an 
eccentric old man with collecting gear and butterfly net. Indeed, the 
remarks of some of my physical science peers led me to think that even 
they bought the image of a biologist, thus portrayed. 

With World War II, the public image took a radical turn. In the 
post World War decade, the public image of science and the work of 
scientists developed directly from the efforts of the scientific com- 
munity during World War II. 

And what were these efforts? The more spectacular of them dealt 
with the harnessing of nuclear energy; the development of radar and 
sonar; the screening and selecting of chemotherapeutic agents such as 
sulfur drugs and metabolic by-products of fungi as antibiotics. 

The mass media publicized the work under the label of scientific 
research. Yet it is a well known fact, at least among scientists, that the 
basic research behind these technological advances was a product of 
work carried out by the scientists of the 1930's and earlier. Further, 


66 Indiana Academy of Science 

we know that the success of the scientific community during World War 
II was not the result of directed research in the 1930's. The success re- 
mains as a clear example of how unfettered, undirected research can 
supply basic building stones on which directed development and tech- 
nology can be constructed. 

But these points were never really mentioned to the public and 
hence their image of science in the post war era was one that can be 
simply stated: given a specific problem, a relatively large amount of 
money, and an adequate concentration of investigators, the efforts of 
these scientists are direct and result in applicable solutions to the 
specific problem in a remarkably short period of time. 

Acting on this image, the public stoutly supported the growth of 
what was labelled in governmental budgets — R and D — research and 

It is difficult to pinpoint the blame for this erroneous image. Either 
few scientists spoke loud enough with corrective statements, or the mass 
media simply ignored, as non-newsworthy, disclaimers from the scientific 

But the fact that this image of capability for immediate solution 
to all problems confronting society became firmly entrenched in the 
community at large can hardly be disputed. 

With the advent of spectacular developments in space by the 
U.S.S.R., a second stimulus for public support of research and de- 
velopment was layed upon the first. I recall that academic salaries and 
departmental budgets increased dramatically — both attributable to the 
appearance in the stratosphere of a man-produced and man-projected 
satellite. At this point, the "D" of R and D began to grow at a rather 
disproportionate rate to its forerunner and counterpart — the "R". 

The scientific community began to speak out loud and clear for its 
share of funds for basic research. Perhaps — for all the wrong reasons — 
the public began to think of R and D with some degree of insight 
provided by science. 

Growth of R and D rose to a rate, in dollars spent, exceeding that 
of our gross national product. Federal expenditures in research and 
development rose over a 20 year period by about 25% each year. The 
need for scientific manpower drew national attention. From the first 
grade through the early years of college, the academic portion of the 
scientific community focused on those individuals whose potential 
predicted a career in science. 

When one examined where we were going in the funding of 
research and development, even as early as 1958, a decade ago, the 
answer was apparent. A simply extrapolation of growth curves for R 
and D and the GNP led Jerome Wisner, Science Advisor to President 
Kennedy, to inject the thought that the bubble might burst. Instead of 
a 25% per year growth, leaders now talk about a 15% per year growth 

Address 67 

— and 15% is still three times the rate of growth of the GNP. The 
problems of the growth of science and technology were apparent then a 
decade ago. 

Congress, drawing its constituents from the public whose image 
of science was that of immediate applicability, began to examine the 
results of R and D spending. This examination was and still is in terms 
of the resolution today of the multitude of problems facing the nation in 
health, transportation, agriculture, education, and urbanization. No 
consideration was or is now being given to the results of governmental 
support in terms of basic accumulation of knowledge — those building 
stones on which development and technology rest. 

The results of this intense scrutiny are familiar to you: the geol- 
ogists have lost Mohole; funding for the 200 BEV accelerator at Weston 
is slow; the chemists are frankly pragmatic in stating that the develop- 
ment of chemistry as pictured in the Westheimer Report will probably 
not be implemented; the IBP is stranded for lack of funds; and even 
NIH in the biomedical domain has some problems. It is true that the 
scrutiny and the drastic cutbacks are made more intensive by our 
involvement in Vietnam, but I submit that the bubble was due to burst 
even without Vietnam. 

And for those of you who are engaged in scholastic education, a 
reduction in the funding of science at the national level will be followed 
at the local level with little time lag. No one engaged in any facet of 
science in the United States will remain immune to the trend now begun. 

Currently, not only research but also development shares the de- 
celeration. Candidly, I think that if the Russians land the Presidium 
on the moon, we — the scientific community — may just avoid a further 
deceleration likely to become a decapitation. 

What is needed? Certainly it is not a return to the 25% per year 
increase in spending; the nation cannot afford it under present or 
projected economic conditions. The first need is a planned steady 
growth of research and development more in line with, and probably 
hinged to, the 5 or 6% annual rate of growth of the GNP. Secondly, a 
more equitable distribution of these funds between research and de- 
velopment must be attained, if the generally accepted cliche "what is 
science today is technology tomorrow" is at all correct. 

Are we likely to achieve these two needs when the war is over? 
And my answer is no, not unless we effectively communicate to a 
critical mass in the 200 million people who constitute our population. 

This communication can occur at two points — one clearly a function 
of the educators present; the other, a function of this academy and 
other institutions of organized science. 

The two points are fairly obvious: initially, during the period of 
basic education; secondly, as an input in the continuing education which 
every adult in the electronic age must carry out if he or she is to adjust 
to the rapid advances of society in this age. 

68 Indiana Academy of Science 

The mere suggestion that we look at science education conveys the 
impression that there is something amiss. What is it? The facts of 
science education as a major vehicle of communication for the past 
two decades seem to fit the following pattern. 

In the face of the most affluent period that science has witnessed 
in any civilization since its inception, in the face of a demand for 
scientific and technical personnel — we began in the elemetary school to 
single out and encourage those individuals whose talents seemed bent 
toward service of mother science. 

We instituted science fairs at the expense of the more general 
hobby fair — and we achieved positive results, plus some negative ones. 
How many students whose talents were bent in another direction were 
rebuffed by the emphasis on science ? 

We put our best teachers into advanced chemistry, advanced 
physics, advanced biology and advanced mathematics in our high 
schools. We created an elite of advanced students — and we achieved 
positive results, plus some negative ones. How many students whose 
bent was in the direction of humanities received advanced courses in 
those fields? And how many of these same students were instructed in 
basic science using examples relevant to their future role as adult 
decision-makers in an age where progress in technology is common 
place — but dependent on the progress in science ? 

We put our best teachers into major courses in institutions of 
higher learning and neglected the courses for liberal arts and business 
students; and we achieved positive results, plus some negative ones. 
How many students joined C. P. Snow's "other culture" with no under- 
standing of science and even a complete disdain for it? 

I submit that for the past two decades we have communicated in 
science education to a very select group. The total number of this group, 
although in our eyes large, simply does not represent the critical mass, 
in our population, under our system of government, necessary to insure 
support of science at a sound fiscal level. Sound in terms of the total 
national prosperity and sound in terms of permitting the type of planned 
organization and growth which characterizes science itself. 

In 20 years then, characterized by unprecedented emphasis in 
science education, we have produced one generation and begun another 
who possess a very basic misunderstanding of what science is and what 
it can do. The youth of this age — and here I speak of our current high 
school and college populations — are convinced that science does not have 
all the answers. I believe this is youthful wisdom. Not all of our 
problems — certainly not all of the socio-economic behavioral complex of 
urbanization — are susceptible to resolution by application of science. But 
I fear this glimpse of wisdom carries with it youthful immaturity. 
Immaturity that will lead — should I say is leading? — to the abandon- 
ment of science. How else do you explain the national trend of de- 
creased enrollments in science ? 

Address 69 

I call then for a very serious reconsideration of our entire approach 
to science communication in terms of basic education. I do not seek to 
materially reduce the concentration on a selected few, but I ask for 
balance and certainly more attention to the education of the non- 
scientist, particularly to his understanding of science and technology. 
This understanding should be developed in terms relevant to his role 
as an adult in an age where science can and does permeate every aspect 
of life. 

The second point of communication with the non-scientist is with 
the adult and I suggest this as a function of this academy, its sister 
institutions and other organized elements in science. 

Our aims on a long term basis would be to reinforce public under- 
standing for the sake of science and its support and to provide a 
continual updating of scientific background as a basis for decision 
making in public policy and personal problems. 

In an address to the 1966 Academy Conference of the AAAS, E. 
G. Sherburne, Jr., Director of Science Service, outlined a program for 
the academies — a program of communication to the non-scientist. I 
endorse his proposal as timely and efficient. Sherburne noted that two- 
tenths of one percent of the population in any state or city are the 
leaders. For them he suggested: 

1. that the Academy establish a science advisor to the governor 
and an advisory committee in frank imitation to the presidential 
science advisor and his committee. 

2. that the Academy conduct science seminars on relevant problems 
for members of the state legislature or even county boards. 

3. that the Academy appoint ad hoc committees to study problems 
relevant to science at the state level and to communicate the 
findings and recommendations to the decision makers. 

Through its committee on Science and Society, the Academy has moved 
to implement some of these ideas. The Committee's activity should be 
strongly endorsed by effort on the part of each member to implement 
the program. 

At the same Academy Conference, I addressed the representatives 
on the role of academies of science in the field of scientific publications. 
In my talk I reviewed the publication practices of some 48 academies of 
science. Only 2 of the 48 publish newsletters directed solely to the 
general public. Yet these two by their action recognize an important 
role of the scientist and his organizations — namely the communication 
to the attentive public of studied opinions on those matters affecting 
science, and those problems facing the state for which there is an 
answer in science. 

In conclusion, may I point out that regardless of your role as a 
scientist, you have a stake in development of a critical mass. For 
each of you in your day to day activities are communicators of science. 
Your decisions in communicating science produce both positive and nega- 
tive effects. Science, today, cannot afford negativism. If you agree with 
me, will you begin today to meet the challenge? 


Chairman : B. K. Swartz, Jr., Ball State University 

Robert E. Pace, Indiana State University, was elected 

chairman for 1969 


Stratigraphy of the White Site. Ronald L. Michael, Ball State Univer- 
sity. — This paper is a field report of archaeological excavation done in 
1968 at Mound 3, Hn-10 (IAS-BSU), located on the near southwest side 
of New Castle, Indiana. The work consisted primarily of searching for 
the base of the mound. The perimeter of the central section of the 
mound was trenched around using 5x5 excavation units. After each 
trench was excavated to a depth of about two feet below the mound 
base, soil profiles were taken of each trench. On the basis of the soil 
profiles and artifacts recovered, which included copper bracelets, pottery 
and ground stone chunkey stones, bifacially flaked chert points, and a 
unifacially flaked chert scraper, the mound was concluded to be middle 
or late Adena. 

A Preliminary Report on the Welsh-Dunlap Site, Vigo County, Indiana. 

Robert E. Pace, Stephen Coffin, and John Richardson, Indiana 
University. — Located on a sand rise along the Wabash River in northern 
Vigo County, this mixed Late Woodland village site is of especial interest 
because of its marginal position to better known traditions to the north 
and to the south. Initial excavation has recovered materials related to 
those reported from the Albee and Catlin sites, along with others 
suggesting affinities with central Illinois River valley sites. 

The Initial Excavation of the Van Nuys Site. Ben J. Morris, Ball State 
University. — The initial excavation of the Van Nuys Site, Hn-25 (IAS- 
BSU), began on June 11, 1968. By July 11, 1968, nearly 1000 square 
feet was excavated to a level averaging approximately 27 inches. The 
assemblage of artifacts recovered from the surface and during excava- 
tion, plus an extensive, multiple post hole pattern, suggests a Late 
Woodland manifestation possibly related to Hn-2, a nearby cemetery site 
with Fort Ancient affiliations. 

Mound Four New Castle Site. Terry Curren, Ball State University. — 
Mound Four is a bilobate mound located in Henry County on the 
grounds of the New Castle State Hospital. The north side of the west 
lobe was excavated during the 1968 Field School. It was originally 
thought that the area between the two lobes of the mound was filled by 
erosion. However, three secondary cremation burials were found in this 
area. A large sheet of mica and a copper band of questionable use was 
found in association with one of the burials. 

Culture Change in a Maya Community. Michael Salovesh, Purdue Uni- 
versity. — The Tzotzil-speaking Maya Indians of San Bartolome, Chiapas, 
Mexico, form a separate community within a town which includes many 


72 Indiana Academy of Science 

non-Indians. Based on research conducted between 1958 and 1962, I 
characterized this community as a strongly conservative one. I offered 
an analysis which suggested that the peculiar residence patterns followed 
by the Indians operated in ways which made these conservative tendencies 
most notable in those areas of community life which were controlled 
by men. 

In the last six years, San Bartolome's Indians have undergone a 
series of drastic shifts in cultural orientations. These have been strong- 
est in political and economic affairs, particularly in those areas which 
involve contact and interaction with the non-Indian world at the levels 
of town, state, and nation. It is remarkable that these changes have 
been most far-reaching precisely in those activities which are primarily 
controlled by men. This paper outlines the major areas of recent change, 
and seeks an explanation of those changes in a basic shift in the 
cultural ecology of the surrounding region. 

Other papers read 

An Historic Feature at the New Castle Site. Thomas Habart, Ball State 

Excavations at the Commissary Site, Henry County, Indiana: A Late 
Woodland Cemetery. Mary Lou Craig, Ball State University. 

Three Late Woodland Cemeteries in Greene County, Indiana. Curtis H. 
Tomak, Indiana University. 

Physical Types in California. Jack M. Whitehead, Ball State University. 

The Nature and Distribution of Bifrontal Occipital Cranial Deformation. 

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

Population Distance and Racial Differentiation in the American Indian. 

Georg K. Neumann, Indiana University. 

Varieties of North American Indians and Linguistic Groupings in His- 
torical Reconstruction. Norman A. Tague and Georg K. Neumann, 
Indiana University. 

Oneota Influences in the Great Lakes Area Tribes. Elizabeth J. Glenn, 
Ball State University. 

The Incidence of the Perforation of the Coronoid-olecranon 

Septum in the Middle Mississippian Population of 

Dickson Mounds, Fulton County, Illinois 

Robert L. Blakely, Randall J. Marmouze, and David D. Wynne 

Indiana University 


The humeral septum is a thin plate of bone between the coronoid and 
olecranon fossae at the distal end of the humerus. Into these fossae fit the 
coronoid and olecranon processes of the ulna and together they form the 
hinge joint of the elbow. Occasionally the septum dividing the fossae is 

During the summer of 19G8 the authors conducted a quantitative and 
qualitative study of the perforation of the coronoid-olecranon septum 
in a prehistoric Middle Mississippian American Indian population at 
Dickson Mounds, Fulton County, Illinois. Included in the study were 
determination of the incidence of bilateral and unilateral perforation, the 
relationship of the perforation with age and sex, and the possible relation- 
ship of the size of the perforation with age and sex. 

It was found that the perforation of the coronoid-olecranon septum is 
positively correlated with sex, occurring more commonly in females. The 
perforation seems to be absent in infants and children. The size of the 
perforation does not seem to increase with age. 


At the distal end of the humerus is a thin plate of compact bone 
which separates the coronoid fossa and the olecranon fossa. The bone 
between the two fossae is termed the humeral, or coronoid-olecranon, 
septum. Into these fossae fit the coronoid and olecranon processes, 
respectively, of the ulna. These hook-like processes of the ulna together 
with the rounded trochlea and fossae of the humerus form the hinge 
joint of the elbow. Extension of the arm is limited by contact between 
the olecranon process and posterior surface of the septum. Flexion at the 
elbow is restricted by contact between the coronoid process and the 
anterior surface of the septum and the intervening soft tissue on the 
anterior aspect of the upper and lower arm. 

Occasionally the coronoid-olecranon septum is perforated (Figure 1). 
Apertures in the septum may be bilateral, that is, occurring in both 
humeri of one individual, or it may be unilateral, occurring on either 
the right or left side. Hrdlicka noted the occurrence of bilaterally and 
unilaterally perforated septa in a variety of mammalian species, thus 
demonstrating that the presence of the aperture is not species-specific 
(6). Among human populations the frequency of the perforation varies 
from close to zero to almost 60% (1). In most human groups observed 
to date, perforated septa are more common among females than males, 
and when present are predominantly bilateral. The incidence of 
perforated septa is greater in the left than right humerus. 

Various theories have been put forth to explain the causes of the 
variability within and between human populations. Both genetic and 



Indiana Academy of Science 

environmental determinants have been invoked, but it remains unclear 
as to what degree the various factors contribute to the formation of the 
aperture. The mechanical hypothesis contends that the perforation is a 
result of activity which wears a hole in the septum. The aperture may 
be formed by certain types of activity or activities of long duration and 
thus might be culturally determined. 

Figure 1. Anterior view of two 
right humeri. The coronoid-ole- 
cranon septum of the humerus on 
the left is unperforated while the 
septum of the humerus on the right 
is perforated. 

In a qualitative study conducted in 1932, Hrdlicka posited a phylo- 
genetic origin for perforation of the coronoid-olecranon septum, sug- 
gesting that the potential for the perforation is inherited as one of a 
number of generalized mammalian traits (6). As evidence, he cites 
comparative data which demonstrates similar frequencies of incidence 
and comparable sexual variability in a variety of infrahuman mammals. 
Hrdlicka also pointed out that the differences in the frequency of the 
perforation among human populations sharing similar physique and 
mode of life and activity suggest that presence of the aperture is 
common to all human groups. Hrdlicka further suggests that the reali- 
zation of this genetic potential is determined by the degree of robusticity 
of the humerus. The greater the muscularity of an individual, the 
greater the bone formation and consequently the less likely that incom- 
plete ossification will result in the absence of bone at the septum. Thus 
lack of bone resorption may result in the perforated coronoid-olecranon 
septum. This theory would explain the higher incidence of the perfora- 
tion on the weaker left arm of right-handed individuals, and the greater 

Anthropology 75 

frequency of the perforation among females who are generally less 
robust than males. 

Recent studies have tended to ignore Hrdlicka's phylogenetic theory 
and have concentrated on the interpretation that the degree of robusticity 
is fundamental in determining the presence or absence of the perfora- 
tion. Benfer and McKem found a positive correlation between the size 
of the humerus and the presence of perforation of the humeral septum 
(1). Using minimum middle diameter of the humerus as a quantitative 
assessment of robusticity, the authors noted that the aperture was more 
likely to be present in humeri of smaller diameter. Glanville, in 1967, 
published evidence suggesting a positive correlation between the pres- 
ence of the perforation and longer coronoid and olecranon processes and 
increased angles of flexion and extension (5). If robusticity is positively 
correlated with the length of the ulnar processes, then Glanville's find- 
ings lend support to the robusticity theory. 


During the summer of 1968, while analyzing the skeletal material 
excavated at Dickson Mounds, the authors conducted a qualitative and 
quantitative study of the performation of the coronoid-olecranon septum. 
Dickson Mounds (F°34) is one of over fourteen hundred prehistoric 
American Indian sites in Fulton County, Illinois. The semilunar-shaped 
mound is located on a bluff overlooking the west floodplain of the Illinois 
River approximately thirty-five miles southwest of the present Peoria, 
Illinois. The artifact assemblage associated with the burials indicates 
that the site was used as a burial cemetery by a Middle Mississippian 
population (4), although recent evidence suggests that a few of the 
burials represent a Late Woodland people (3). Radiocarbon dates for 
the Eveland site, a Middle Mississippian site adjacent to Dickson 
Mounds, range from 950 to 1350 A.D. (communication from E. J. 

To date, the number of burials recovered at Dickson Mounds ap- 
proximates 1050. The majority of the burials are in an excellent state of 
preservation due to the alkalinity of the loess deposits of which the 
mound is constructed. The sample includes those burials removed during 
the 1966, 1967, and 1968 summer excavations as well as those left ex- 
posed in situ in the enclosed museum for display purposes. 

In the present study of the perforation of the humeral septum the 
authors examined only those skeletons for which age and sex had been 
previously determined (2). In addition, all Late Woodland burials were 
omitted from the analysis because it was felt that to include them in 
the study might invalidate the findings. The Late Woodland people 
may have been subjected to different cultural and physical factors than 
the Middle Mississippian population, and these factors may have con- 
tributed to the formation of the perforation as the mechanical theory 

The sample size was further reduced by selecting out for analysis 
only those burials for which at least the distal portion of both humeri 

76 Indiana Academy of Science 

were available. The purpose in doing so was to eliminate sampling error 
which would skew the relative percentages of the incidence of perfora- 
tion among males and females and the frequencies of right, left, and 
bilaterally perforated septa. As a result, the sample was pared down 
to 147 burials of which thirty-two were infants and children of indetermi- 
nant sex. These burials were excluded from the analysis so that 
an accurate sex ratio could be ascertained. Therefore 115 burials were 
utilized in the study. 


Perforation of the coronoid-olecranon septum has usually been 
treated as a qualitative trait, whose presence or absence has been cor- 
related with various properties of the humerus. The present study is 
both qualitative and quantitative in the sense that, in addition to 
observation of the occurrence and nonoccurence of the perforation, 
osteometric measurements of the aperture were taken. 

First each individual was examined for presence or absence of the 
perforation. Bilateral and unilateral perforations were noted. The fre- 
quency of unperforated septa and bilaterally and unilaterally right and 
left perforated septa were then separated out according to sex and 
graphed in Figure 2 to illustrate the sex differential. From these fre- 
quencies it was possible to derive percentages which demonstrate the 
incidence of the perforation in relation to the total sample (N) and 
male (Ni) and female (N 2 ) samples (Fig. 3). The chi-square test, 


was applied to determine the significance of the following variations: 1) 
distribution of the perforation by sex, 2) distribution of bilateral perfora- 
tion by sex, 3) distribution of unilateral right perforation by sex, and 
4) distribution of unilateral left perforation by sex. Chi-square values 
and significance, or level of confidence, (p) are given in Figure 3. In 
all cases the 0.05 level of significance was required. 

All perforations were then measured. (In four incidences, post- 
mortem damage rendered the aperture immeasurable.) The width and 
height of the perforation were measured on the anterior aspect of the 
bone using sliding calipers accurate to 0.1 mm. Width was taken as the 
maximum distance across the aperture on a line perpendicular to the 
central axis of the shaft, or diaphysis, of the humerus. The height was 
taken as the maximum distance across the aperture on a line parallel 
to the central axis of the diaphysis of the humerus. As a measure of 
the relative size of the perforation the two dimensions were summed. 
The purpose of this quantitative procedure was to provide data which 
might indicate sex and age variations in the size of the perforation. To 
this end, the mean (X) values of the width and height of the perfora- 
tions of both the right and left humerus were derived for males and 
females independently (Fig. 4). To indicate the individual variability in 
size, standard deviation, 


,.d.= yj 

S(fd a ) 







25 ■; 





Figure 2. Frequencies (vertical axis) of unperforated septa (U), bilater- 
ally perforated septa (B), and unilaterally perforated septa: left (L) and 
right (R). 

are also listed. In addition, mean values and standard deviations for the 
total sample are given. To determine the significance of the size differ- 
ence between the sexes, the chi-square test was employed and the results 

78 Indiana Academy of Science 

are presented in Fig. 4. The 0.05 level of significance was again required. 
Not shown is a table in which the size of the perforation is plotted 
against the age gradient of the total sample of individuals with apertures 
(n— 45), males displaying perforation (n=17) and females displaying 
perforation (n=28). The results are discussed below. 


Of the 115 individuals represented in the sample, forty-five, or 
39.1%, possessed either bilateral or unilateral perforations of the coro- 
noid-olecranon septum. Although the sample includes more males than 
females (59:56), apertures occur more frequently in females, twenty- 
eight to seventeen (Fig. 2). Perhaps more revealing is the fact that 
50.0% of the female population manifests bilateral or unilateral perfora- 
tions, whereas occurrence is limited to 28.8% among males. The chi- 
square test was applied to determine the significance of the differential 
incidence by sex, and although it proved insignificant at the 0.05 level 
of confidence (Fig. 3), the disparity should not be overlooked. 

Of the 115 burials, 26.1% represent individuals with bilateral per- 
forations. In both sexes bilaterally perforated septa were more common 
than unilateral perforations, occurring in thirty of the forty-five cases 
in which perforations were observed. However, bilateral perforation of 
the humeral septum is present in 37.5%, represented by twenty-one 
individuals, of the female sample (Ns), while it occurs in only 15.3% 
(nine burials) in the male population (Ni). The difference proved to be 
significant at the 0.05 level, p equalling 0.03 (Fig. 3). 

It is of interest to note that the findings are consistent with the 
data presented in previous studies and lends strong support to the 
robusticity theory. The fact that the aperture is more common among 
females suggest that the lack of robusticity observed in females is 
positively correlated with presence of the aperture. 

Unilaterally perforated septa were few, occurring in 13.0% of the 
total sample. Of the fifteen unilateral perforations, ten were noted on 
the left humerus. There is little sexual variation; both males and females 
manifesting five apertures on the left humerus and three and two per- 
forated septa, respectively, on the right humerus. The 2:1 ratio observed 
between left and right septa may reflect a lack of robusticity of the 
left humerus in right-handed individuals; however this conclusion is 
questionable since the sample size is small and whether this population 
was indeed predominantly right-handed remains unknown. The fact that 
there are twice as many bilateral perforations as unilaterial perforations 
probably indicates that the differences in robusticity between individuals 
is far greater than that between the two humeri of the same individual. 

When all burials were listed in order of ascending age at the time 
of death, it was noted that the width and height of the aperture of 
both the left and right humerus did not increase with age. If the per- 
foration is related to activity as suggested by the mechanical theory, 
one might expect the aperture to enlarge with age, at least until the 

Anthropology 79 

Sample (N) = 115 

Male (Ni) - 59, or 51.3% of N 
Female (N 2 ) = 56, or 48.7% of N 

Incidence of perforation = 45, or 39.1% of N 
Male perforation = 17, or 28.8% of Ni 
Female perforation = 28, or 50.0% of N a 

Chi-square test of significance of distribution of 
perforation by sex: X 2 = 2.68, p > 0.05 and = 0.10 

Incidence of bilateral perforation = 30, or 26.1% of N 
Male bilateral perforation = 9, or 15.3% of Ni 
Female bilateral perforation = 21, or 37.5% of N^ 

Chi-square test of significance of distribution of 
bilateral perforation by sex: X 3 = 4.80, p< 0.05 
and = 0.03 

Incidence of unilateral perforation = 15, or 13.0% of N 
Left perforation = 10, or 8.7% of N 

Male left perforation = 5, or 8.5% of Ni 
Female left perforation = 5, or 8.9% of N2 

Chi-square test of significance of distribution of 
left perforation by sex: X 3 = 0.0, p = 1.0 

Right perforation = 5, or 4.3% of N 

Male right perforation = 3, or 5.1% of Ni 
Female right perforation = 2, or 3.6% of N 2 

Chi-square test of significance of distribution of 
right perforation by sex: X 2 = 0.20, p = 0.65 

Figure 3. Frequencies and percentages of total perforations, bilateral 
perforations, and left and right unilateral perforations of the caronoid- 
olecranon septum indicating the distribution by sex and the significance 
of that distribution. 


Indiana Academy of Science 

Sample = 17 

Left Humerus 



Right Humerus 






± 1.41 


± 1.94 



Left Humerus 



Right Humerus 








Male + Female 
Sample = 45 — 

Left Humerus 



Right Humerus 



X 5.94 4.18 4.76 
s.d. ± 2.76 ± 1.78 ± 2.18 

± 1.55 

Chi-square test of significance of size differential by sex 

Chi-square Left Humerus Right Humerus 

Both Humeri 

X a 3.41 5.23 

> 0.05 < 0.05 

p — 0.06 = 0.02 

< 0.05 
= 0.04 

Figure 4. Mean values of the width and height of the perforation of the 
coronoid-olecranon septum in both humeri. Males are given at the top, 
followed by females. The third chart combines male and female values. 
Standard deviations are listed for each mean value. At the bottom are 
chi square values which indicate the significance of the difference in size 
(width -f- height) between males and females. 

third decade when a decline in activity is inevitable. The data suggests 
that the duration of the activity is not involved in the formation or 
development of the perforation. It should, however, be noted that only 
the size of the perforation at the time of death is known and not what 
the size might have been five, ten, or twenty years before death. The 

Anthropology 81 

assumption has thus been made that the age gradient for the entire 
population is an accurate reflection of the size at different ages. 

The age of the onset of the performance seems to be during the 
second decade when development of musculature is at a maximum and 
bone formation is approaching completion. Of the thirty-four individuals 
observed under age twelve years, only two evidenced perforation of 
either humerus. The two exceptions, seven and eight year old females, 
possessed large perforations which may reflect early maturation or 
genetic variability. 

The mean dimension of the width and height of perforations of 
both the right and left humerus are, in each case, larger in females than 
males (Fig. 4). When the chi-square test is employed to determine the 
significance of the defference in size between males and females, it is 
observed that the disparity is significant to the 0.05 level of confidence. 
This quantitative finding, coupled with the greater incidence of per- 
forations among females, lends credence to the robusticity theory. It 
should also be noted that the mean width and height of the aperture of 
the left humerus is greater than that of the right (Fig. 4). This fact, 
too, supports the robusticity theory. 


In summary, on the basis of the present study of the perforation of 
the coronoid-olecranon septum, the following conclusions can be stated: 

1) The incidence of perforation is greater in females than males. 

2) The incidence of bilaterally perforated septa is significantly greater 
among females. 

3) The size of the perforation in both humeri is significantly larger in 

4) Bilateral perforations are twice as common as unilaterial perfora- 

5) Perforations of the left humerus occur twice as often as perfora- 
tions of the right humerus. 

6) The size of the aperture of the left humerus is uniformly larger 
than the left. 

7) The size of the perforation does not increase with age. 

The current study provides data which may refute the mechanical 
theory. If the perforation is indeed produced by wear, one would expect 
the size of the perforation to increase with the accumulating duration 
of activity. Because the size of the perforation of the humeral septum 
does not increase with age, the authors reject the mechanical theory as 
untenable in light of the present findings. 

Both the qualitative and quantitative data collected during the 
study lend support to the robusticity theory. The authors accept 
Hrdlicka's contention that the presence of the perforation is positively 
correlated with the lack of robusticity due to incomplete bone formation 

82 Indiana Academy of Science 

of the humeral septum (6). As evidence it is possible to cite the greater 
incidence of perforation (and bilateral perforations) among females who 
are less robust than males. Quantitatively, the mean dimensions of both 
humeri are absolutely larger in females than males, suggesting a lesser 
degree of bone resorption among females. The greater frequency and 
larger size of the perforation of the left humerus also tend to substanti- 
ate the robusticity theory, if indeed the majority of the sample was 

The authors have no basis for either accepting or refuting Hrdlicka's 
proposed phylogenetic origin of the perforation of the coronoid- 
olecranon septum. However, it is suggested that perhaps it is not the 
potential for the perforation that is inherited, but rather that the poten- 
tial for degrees of robusticity are inherited and conditioned by environ- 
mental factors, the result of which is the presence or absence of the 
humeral septum. 

It seems probable that the perforation of the coronoid-olecranon 
septum is a result of more than one or two interrelated factors. Before 
it is possible to understand this complex problem, more data will be 
required. More extensive studies as well as more intensive analyses of 
existing data are necessary before the questions touched upon here can 
be fully answered. 

Literature Cited 

Benfer, R. A., and T. W. McKern. 1966. The correlation of bone robusticity 
with the perforation of the coronoid-olecranon septum in the humerus 
of man. Amer. J. Phys. Anthrop. 24:247-55. 

Blakely, R. L., and P. L. Walker. 1968. Mortality profile of the Middle 
Mississippian population of Dickson Mounds, Fulton County, Illinois. 
Indiana Acad. Science 77:102-108. 

Caldwell, J. R. 1959. The Mississippian Period. Illinois Archaeology, 
Bui. No. 1. 

Cole, F., and T. Deuel. 1937. Rediscovering- Illinois. University of Chicago 
Press, Chicago. 

Glanville, E. V. 1967. Perforation of the coronoid-olecranon septum: 
humero-ulnar relationships in Netherlands and African populations. 
Amer. J. Phys. Anthrop. 26:85-92. 

HrdliCka, A. 1932. The humerus: septal apertures. Anthropologic 
(Prague) 10:31-96. 

The Linear Growth of Long Bones in 
Late Woodland Indian Children 

Phillip L. Walker, Indiana University 


Studies of the linear growth of long bones in children have almost 
exclusively utilized indirect measurements of the living plotted against 
chronological age. Such studies are only in a very general way applicable 
to age determination in archaeological populations. The present study is 
based on direct measurements of Late Woodland Indian Children and phys- 
iological age based on the degree of the development of the dentition. By 
plotting the lengths of individual long bones against dental age, group 
growth curves were constructed which allow the investigator to determine 
the dental age of an individual when adequate dentition is not present by 
measurement of the long bone alone. To discern the applicability of growth 
curves for a specific prehistoric American Indian population to other similar 
populations a comparative study of a Middle Mississippian group was under- 
taken. Although the results of this comparison were inconclusive, it seems 
prudent to apply the growth curves for the Late Woodland population 
studied only to very similar populations until more extensive comparisons 
can be made. 

Studies of the linear growth of long bones in children have almost 
exclusively been limited to indirect measurements of the living taken 
from radiographs. Such studies introduce elements of distortion because 
of individual variability of the soft tissues and cannot be considered as 
parallel to direct measurements of dried bones (2). Usually the ages 
recorded for children in such studies are chronological rather than physio- 
logical. That is, they reflect the number of years since birth and not 
physiological maturity. In populations obtained from archaeological 
excavations only direct measurements of long bones can be made. It is 
also obvious that only the physiological age of the individual as indicated 
by dental and skeletal maturity can be obtained. Because of these dis- 
crepancies, age growth curves based on indirect measurements of the 
living are only in a very general way applicable to age determination 
from direct measurements of the skeletal remains of archaeological 

To avoid the error of applying growth curves based on indirect 
measurement and chronological age to skeletal populations, it seems 
reasonable to construct a group growth curve based on direct measure- 
ment of long bones and physiological age based on the degree of devel- 
opment of the teeth. Group growth studies of this type have previously 
been done by King B. Hunter for the Hopewellian population of the 
Klunk mound group in Calhoun County, Illinois. The present study 
follows Hunter's basic procedures. 

The use of dental development as an age determinant is based on sev- 
eral factors. Teeth are consistently well preserved in archaeological pop- 
ulations and offer a rather complete record of growth. Secondly, the 
rate of development of the dentition is much less subject to variation 
due to extraneous environmental factors than is long bone growth. 


84 Indiana Academy of Science 

Under conditions of dietary and hormonal stress the teeth seem to 
have priority for the use of deficient materials (1). For this reason 
teeth develop at a rather constant rate even in diverse populations. This 
is in sharp contrast to the rate of growth of long bones which varies 
significantly between populations. These growth factors allow the appli- 
cation of standard chronologies for tooth development to archaeological 
populations of unknown nutritional and physical makeup with a high 
degree of accuracy. In growth studies of skeletal populations therefore, 
the dental age of the individual can be treated as a relative constant 
against which the much greater variability of long bone growth can be 

Charts which plot dental age against long bone length of a popula- 
tion are of great value because they allow the investigator to determine 
the approximate dental age of a child by direct measurement of long 
bones alone. Such charts are of particular value when dealing with an 
archaeological population because demographic studies can be made 
which include individuals who do not have adequate dentition for the 
assessment of dental age but do have intact long bones. Because of the 
increased sample size obtained from such assessments the validity of 
conclusions based on resultant mortality statistics is increased. 

The skeletal material used in the present study was excavated by 
Gregory Perino during the summers of 1967 and 1968 from the Yokem 
mound group in Pike County, Illinois. The Yokem mound group consists 
of ten mounds which represent two distinct occupations of the area. Five 
of the mounds contained a bluff culture artifact assemblage and represent 
a relatively early Late Woodland occupation of the area. The remaining 
mounds represent a late occupation of the area by another group of 
Late Woodland peoples. Burial practices and artifacts from these later 
mounds indicate that this group was assimilating Middle Mississippian 
patterns. This assumption is corroborated by a relatively late 
radiocarbon date of A.D. 1208 ± 90 years. 

The date for this study was obtained from children of the later 
population of Late Woodland peoples to use the Yokem mounds. These 
were individuals excavated from mounds ll-Pk°167, ll-Pk°168, 
ll-Pk°169, ll-Pk°170, and ll-Pk°171. Forty-three children were examined 
ranging in age from zero months to 12 years. Twenty-three additional 
children were excluded from the sample, either because adequate denti- 
tion and measurable long bones were missing or because marked dental 
pathologies were present wiiich would obscure developmental patterns. 

The dental age for each individual was assessed in terms of five 
stages of dental development. These stages are: first evidence of calci- 
fication, crown completion, eruption, root completion, and root reabsorp- 
tion. All measurements of long bones were made exclusive of epiphyses. 

Figures 1, 2, and 3 represent the relationship between the long bone 
lengths and the dental ages of the Late Woodland population of the 
Yokem mounds. The dots on the graph signify the dental age plotted 
against long bone length for specific individuals. The solid lines repre- 



300 n 

3 4 5 6 7 8 

Figure 1. Length of humerus and femur plotted against dental age. Dots 
represent individual values, lines represent group growth eurves. 



9 HlO ' 11 r .12 

Figure 2. Length of fibula and ulna plotted against dental age. Dots repre- 
sent individual values, lines represent growth curves. 


Indiana Academy of Science 



Figure 3. Length of tibia and radius plotted against dental age. 
represent individual values, lines represent group growth curves. 





• ••Dickson 
— Yokem 





T T T "^ 


Figure 4. Lengths of ulnae from Dickson and Yokem mounds. Dots repre- 
sent individuals from Dickson mound, line represents group growth curve 
from Yokem mounds. 

Anthropology 87 

sent the smoothed curve formed by these values. When these curves are 
examined it will be noted that at the age of about two and one-half years 
the rate of growth consistently decreases for all long bones. This 
decrease in rate of growth represents the end of the rapid growth of 
infancy. The next marked change in the growth rate takes place at about 
nine years of age where growth accelerates. This change in rate repre- 
sents the hormonal initiation of growth in females of the population. It 
should be mentioned that no sexual identification was attempted for the 
children used in this study for two reasons. First the number of indi- 
viduals of the ages during which differential growth rates because of 
sexual differentiation is too small to produce a valid sexual separation. 
Second, sexual identification of the skeletal remains of children is often 
unreliable, particularly when the ilium is not present to permit the 
measurement of the greater sciatic notch. Because sexual identification 
was not made, the female growth spurt which occurs at about nine years 
will be partially obscured by the unchanging male growth rate. At this 
point it should be emphasized that the growth curves illustrated in 
Figures 1, 2, and 3 do not indicate the growth rate of the average or 
normal individual, but illustrate the growth of the population as a group 
(2), regardless of sex. 

To show the degree to which the growth charts constructed for the 
Yokem mound Late Woodland population can be applied to other prehis- 
toric American Indian populations, a Middle Mississippian population 
was examined and data comparable to that from Yokem mounds was 
obtained. The individuals used in this comparative study were taken 
from burials excavated at Dickson Mounds (ll-F°-34) in Fulton County, 
Illinois, during the summers of 1966 and 1967. This sample excluded the 
Late Woodland component of this site. Figure 4 represents by dots the 
dental age plotted against the ulna length of 14 children from Dickson 
Mound. There is no significant deviation in the growth rates of the two 
populations during early childhood. In later years, especially during the 
period of adolescent growth acceleration in females, the comparative 
sample shows consistently larger long bone lengths. There are several 
possible explanations for this deviation. Because of the small size of 
the comparative sample, sampling errors could easily account for this 
deviation. An alternative explanation could be that the two populations 
were nutritionally and/or physically different, and it is these differences 
that are reflected in Figure 4. Until larger comparative studies can be 
done, the growth rates of the Late Woodland population which is the 
subject of this study should be considered as strictly applicable only to 
populations which are archaeologically and physically very similar to the 
Yokem Mound population. 

Literature Cited 

1. Jenkins, G. Neil. 1954. The physiology of the mouth. Blackwell Scientific 
Publications 183. 

2. Marsh, M. M. 1955. Linear growth of long- bones of extremities from 
infancy through adolescence. J. Dis. Child. 89:725-742. 

On the Origin of the Tntelo — An Eastern Sionan Tribe 

Ralph W. Alexander, Jr., and Georg K. Neumann, Indiana University 


This paper is a contribution to the solution of the puzzling problem 
posed by the discovery of several Indian tribes in the Allegheny Piedmont 
area who spoke languages related to the Siouan tribes of the Great Plains. 
The discovery posed the question as to whether the Siouan languages 
originated in the East, the Plains area, or the intermediate area, the Obio 
Valley, that is, the direction of their spread. By correlating linguistic, 
cultural, and physical variables, the geographic and temporal relation- 
ships of these groupings are elucidated. The material that was analyzed 
is the only known sample representative of the prehistoric Eastern Siouan 
population which gave rise to the historic Tutelo Indian tribe of North 
Carolina. The physical characteristics of this group proved to be closely 
related to those of the Archaic series of crania from Indian Knoll, 
Kentucky, a fact that traces the Eastern group into the Ohio Valley. 

The Siouan tribes of North America may be divided into two main 
bodies. The larger Western group is composed of such tribes as the 
Quapaw, Osage, Kansa, Missouri, Oto, Iowa, Omaha, Ponca, Yankton, 
Santee, Yanktonai, Teton, Mandan, Hidatsa, Crow, Assiniboin, and 
Winnebago. The Eastern group, located in the region of Virginia, North 
Carolina and South Carolina, may be divided into two subgroups. The 
Northern subgroup is typically represented by the Tutelo, and the 
Southern subgroup represented by the Catawba. Although the Eastern 
tribes were known to early explorers, they were not recognized as having 
a linguistic bond with the Western group until 1870, when Horatio Hale 
suggested the existence of a Siouan dialect east of the Appalachians after 
obtaining a vocabulary from an old Tutelo man (4). Work by Hale, 
Gatschet (3), and Dorsey (2) was incorporated by James Mooney (5) 
into a classic Bureau of American Ethnology bulletin which satisfactorily 
demonstrated the Siouan connection of the Eastern Groups. The estab- 
lishment of the widespread diffusion of the Siouan-speaking tribes was 
summarized by Paul Weer in 1937 in an Indiana History Bulletin (8). 
The Northern, or Tutelo group of the Eastern Sioux are the most closely 
related linguistically to the Western Sioux. The Southern or Catawba 
group is the most aberrant of the Siouan groups linguistically. There 
is evidence, as yet not thoroughly presented, that the Siouan and 
Muskhogean linguistic families may be related, and that the Catawba 
dialect may occupy an intermediate position between the extreme 
branches of each. The problem, then, is one of the origin of the Eastern 
Sioux, whose linguistic bond with the Western group demonstrates beyond 
a reasonable doubt that the two bodies had formerly been in contact. 
We must conclude that either the tribes occupying the territory between 
had died out, that the Eastern tribes had migrated farther East or the 
Western farther West, or that both easterly and westerly movements had 
taken place. 

The ethnohistoric evidence available supports the last supposition. 
Most of the Western tribes retained until quite recently traditions of a 


Anthropology 89 

more easterly origin. The Eastern tribes, according to historic docu- 
ments, extended farther West. In the Eastern Piedmont area, many 
local migrations and tribal movements tend to obscure the earliest areas 
of occupation, but the bulk of the ethnohistoric evidence points to an 
origin of the Eastern Siouan peoples in the Ohio Valley area. Much of 
the ethnohistoric evidence bearing on the subject has been assembled by 
John R. S wanton (7). 

Archaeologically, the evidence which bears on the problem is slight, 
but convincing. JorTre Coe, who is the major contributor to our archaeo- 
logical knowledge of the Piedmont area, considers the Badin Focus cul- 
ture of the Piedmont to be directly ancestral to the historic Eastern 
Siouans (1). Further, Coe feels that the correlation between the Badin 
Focus culture and the Indian Knoll culture of Kentucky is so great that 
some direct connection must be postulated. The Badin Focus culture 
appeared in the Piedmont at the same time the shell-mound culture in 
Kentucky was replaced by the Adena people. On the basis of trait com- 
parison, the Badin-Indian Knoll relationship is closer than any other yet 

Physical evidence bearing on the varietal affiliations of the Eastern 
Siouan groups has, until now, been almost totally lacking. The senior 
author was the first to postulate that the skeletal material from the 
historic Tutelo villages was closely related to that from the Archaic 
Indian Knoll site, and used the latter as the type series, designated the 
Iswanid variety after the Catawba words iswa (river) and nie 
(people) (6). 

The solution of the problem of the origins of the Eastern Siouan 
people is only possible through the investigation of the physical anthro- 
pology of the populations of the Eastern Siouan area, comparison with 
other established varieties of American Indians, and by correlation of 
this information with the linguistic, archaeological, and ethnohistoric evi- 
dence available. This paper presents the results of the first bioanthro- 
pological examination of the only known sample of Eastern Siouan and 
specifically Tutelo skeletal material. The sample used for description, 
analysis and comparison consists of twenty-eight adult crania from 
Northern North Carolina, excavated by JorTre Coe. The original meas- 
urements and observations were taken by G. K. Neumann. The several 
sites from which the sample was obtained all date from late pre-contact 
times, and are directly ancestral to historic Tutelo villages. Since no 
other material is available, and since the Tutelo are the most repre- 
sentative of the Northern dialect group of the Eastern Siouans, the 
sample will serve to answer questions concerned not only with the 
Tutelo, but will allow inference in regard to the entire Eastern Siouan 
population. No Catawba skeletal material exists. 

Three types of comparisons were possible: (1) a comparison with 
Archaic groups, the Lenid or Iswanid varieties; (2) a comparison with 
more recent groups such as the Muskogid or Ilinid varieties; or (3) with 
a more modern variety, the Dakotid of the Plains. The Lenid variety is 

90 Indiana Academy of Science 

represented in the coastal Algonkian populations, which are unlike the 
Tutelo, and was ruled out as unlikely to bear much similarity. The 
Dakotid groups are very unlike the Eastern Siouan material morpho- 
logically, and were likewise ruled out. The Ilinid variety is geograph- 
ically less likely to be related to the Eastern Siouans than the remaining 
groups, the Muskogid and the Iswanid. There is a strong possibility 
that the Muskogid groups may be related to the Southern or Catawba 
Eastern Siouans, and this is also suggested linguistically, but there is 
no Catawba material available. The Iswanid variety bears obvious 
morphological resemblance to the Tutelo sample, and the type series for 
this variety is the Indian Knoll population responsible for the Indian 
Knoll culture which closely links the Eastern Sioux with the Ohio Valley 
area. A rigorous statistical analysis and comparison of the Indian Knoll 
Iswanid series and the Tutelo Eastern Siouan series was therefore under- 
taken, and the linguistic, archaeological, and racial traits correlated to 
find the best fit, an explanation of the origin of the Tutelo Eastern 

The statistical methods used in the complete analysis and comparison 
were the F test for homogeneity of variance of the two samples, upon 
which the t test for significant difference in means was based. A fur- 
ther test for significant difference in means was employed, the 3 
X.P.Es test, or three times the probable error of the difference between 
the means. The results were virtually identical in each case, and may be 
considered highly reliable. Morphological observations were also compiled 
in tables of frequency distribution for each observation. The computa- 
tions for the various statistical tests employed were done through the 
facilities of the Research Computing Center at Indiana University. 

The Tutelo Eastern Siouan skull may be briefly described as follows: 
the skull is medium in size, with a glabello-occipital length of 182.2 mm., 
a maximum breadth of 137.1 mm., and a basion-bregma height of 
137.4 mm. These dimensions yield a cranial index of 75.24, and a length- 
height index of 75.44. The cranial vault is therefore on the lower border 
of mesocrany and high in relation to its length. It is ovoid in form, 
with medium muscular relief, large to very large divided brow ridges, 
medium frontal height and slope, small to medium frontal breadth, a 
slight amount of sagittal cresting, medium parietal eminences, medium 
lambdoid flattening, and an occiput with medium curve and low position. 
The face as a whole is of moderate and not rugged build, and not large 
in relation to the braincase. All facial dimensions tend to be moderate, 
with a total facial height of 117.9 mm., an upper facial height of 
71.1 mm., and a total facial breadth of 133.1 mm. The indicial pro- 
portions are also moderate, with a total facial index of 88.6, 
mesoprosopic, and and upper facial index of 53.51, mesene. The 
size and height of the zygomatic bones are medium, and the 
anterior projection of the zygomatics is small to medium. The lateral pro- 
jection of the zygomatics is less moderate, and is medium to large. The 
orbit shape is most frequently square, often rhomboid, with a small to 
medium amount of inclination. The left orbital index is 80.82 if the 
breadth is taken from maxillofrontale, and 85.30 from dacryon. The first 

Anthropology 91 

index is mesoconch, the second barely falls into the hypsiconch indicial 
category. The nasal index is 48.28, or mesorrhine. Absolute diameters 
of the nasal structures are small, with a nasal breadth of 24.1 
mm., and a nasal height of 50.0 mm. Both nasal root and bridge 
dimensions are moderate; the nasal root breadth small to medium, the 
nasal bridge medium, and the nasal bridge breadth small to medium. 
Midfacial prognathism is absent to slight; aveolar and total prognathism 
are slight. The size of the mandible is medium. The most common chin 
form is bilateral, with a wide bilateral the modal subtype. Gonial eversion 
is varied, but predominantly small. In a description of this nature, the 
Tutelo Eastern Siouan skull is found to be virtually identical with the 
Indian Knoll Iswanid series. Computer analysis at the sensitivity level 
of the t test for significant difference in means between the two series 
reveals only nine significant differences out of thirty-four measurements, 
five significant differences out of twenty-three indices, and five significant 
differences out of seventy-eight morphological observations. These dif- 
ferences may be attributed to the slightly larger Tutelo skull, and repre- 
sent variation well within the limits of a local series of a variety. 

The consideration of the linguistic, ethnohistorical, and archaeologi- 
cal evidence available, and the detailed metric and morphological analysis 
of the available Eastern Siouan skeletal material leads to the following 

1. The Tutelo Eastern Siouan cranial series exhibits a high degree of 
similarity with Neumann's Iswanid variety as represented by the 
Indian Knoll series. The metric differences between the two series, such 
as the slightly larger Tutelo skull and the slightly higher Iswanid vault 
represent local differences which would be expected considering the time 
span separating the two series and, when viewed in the light of the 
great majority of measurements and indices which exhibit no statistically 
significant difference, are not sufficient to negate the conclusion that the 
Tutelo Eastern Siouan series is representative of the Iswanid variety of 
American Indian. Morphologically, the two series are also extremely 
similar, and the morphological observations substantiate the metric data 
completely. The Tutelo Eastern Siouan series may then be described as 
a local series of the Iswanid variety, with the differences between them 
and the type series being negligible on a varietal level. 

2. From the standpoint of methodology, this study has demon- 
strated that the use of the combination of metric, indicial, and morpho- 
logical characteristics as employed by G. K. Neumann (6) along with 
tests of statistically significant differences in means such as the t test and 
the 3 X.P.Es. test is sufficiently sensitive to detect minor differences and 
therefore adequate for historical reconstruction. 

3. The correlation of the linguistic picture, the archaeological and 
ethnohistorical evidence, and the physical evidence assembled in this 
paper provide a new insight into the origins of the Eastern Siouan 
peoples. The first detailed description of Eastern Siouan crania which 
can be documented as to their tribal affiliation has been provided, and 

92 Indiana Academy of Science 

the connection with the Archaic Iswanid variety makes a contribution in 
linking prehistoric archaeological manifestations with historic tribes. It 
has been shown that, of the Eastern Sioux the Tutelo are linguistically 
more closely related to the Western branch of the Siouan family than to 
the other Eastern group, and the ethnohistoric evidence available 
strongly points to an Ohio Valley origin for the Eastern Sioux. The 
archaeological evidence demonstrates the similarity between the Badin 
Focus of the Carolina Piedmont, which appears directly ancestral to the 
historic Eastern Siouan groups, and the Indian Knell culture of Ken- 
tucky, of which the Iswanid physical type is representative. Finally, the 
physical evidence presented demonstrates the varietal relationship 
between the Tutelo Eastern Siouan series and the Iswanid series of 
Indian Knoll. Therefore, from the evidence available from linguistics, 
archaeology, and physical anthropology, the conclusion is reached that 
the origin of the Eastern Siouan peoples can now be connected more 
strongly to the Ohio Valley area than was previously possible. 

Literature Cited 

1. Coe, Joffre L. 1952. The Cultural Sequence of the Carolina Piedmont. In: 
Griffin, J. B. 1952. Archeology of Eastern United States, p. 301-311. 

2. Dorsey, James O. 1893. Siouan Sociology. Bureau of American Ethnology 
15th Annual Report, 1893-1894. p. 205-244. 

3. Gatschet, A. S. A. 1894. Grammatic Sketch of the Catawba Language. 
Amer. Anthropol. 5:2. 

4. Hale, Horatio. 18S3. The Tutelo Tribe and Language. Proc. Amer. Philos. 
Soc. 21:114. 

5. Mooney, James 1894. The Siouan Tribes of the East. Bureau of American 
Ethnology Bulletin No. 22. 

6. Neumann, Georg K. 1952. Archaeology and Race in the American Indian. 
In: Griffin, James B. 1952. Archeology of Eastern United States. 

7. Swanton, John R. 1937. Siouan Tribes of the Ohio Valley. In: The 
Indianapolis Archaeological Conference, December, 1935. 

8. Weer, Paul. 1937. Preliminary Notes on the Siouan Family. Indiana Hist. 
Bull. 14(1):99-120. 

The Origin of the Shawnee Indians 

Louise M. Robbins, University of Kentucky 


Georg K. Neumann", Indiana University 

The Middle Ohio Valley has known the presence of man from pre- 
historic times to the present day. Until the coming of the White settlers, 
the aboriginal inhabitants of the valley directed most of their energies 
toward obtaining food in sufficient quantities to permit survival and 
perpetuation of their social group. Some subsistence patterns, such as 
agriculture, involved greater cultural elaboration of material items, 
whereas less diversity in the material traits generally are associated with 
earlier hunting and gathering populations. In other words, the earlier 
inhabitants of the Middle Ohio Valley concentrated on practicing their 
culture and left some of its products in passing. Although the more 
recent immigrants to the region, the White settlers, were also interested 
in populating the area and gaining a living from it, they brought 
a new interest with them. They were inquisitive about the earlier occu- 
pants whose cultural items were frequently found on the surface of the 
ground or when the soil was cultivated. As a result, considerable atten- 
tion was, and continues to be, devoted to the location, description, and 
identification of the cultures of the prehistoric inhabitants. Nor is inter- 
est in extinct cultures confined to the professionally trained investigator, 
as a perusal of early historic journals will illustrate. More frequently, 
it has been the amateur investigator who finds evidence of aboriginal 
occupations and who calls it to the attention of the trained specialist. 
Within the last one hundred years, many prehistoric sites have been 
found and subsequently given particular names in terms of location, of 
temporal placement, and of the material remains found on the site. Often 
times items of the material culture are found in association with physical 
remains of the population, and it then becomes possible to study not 
only the life ways of the people but to study the people themselves. 

An attempt is made here to reconstruct the history of the people 
who produced the Fort Ancient archaeological cultural assemblage by 
using a multidisciplinary approach toward the solution of a historic 
problem. Specifically, a test is made to ascertain to what extent the 
physical data of the Fort Ancient Aspect population support the 
conclusions that have been drawn on the basis of archaeological evidence. 

During the 1930's, Griffin (2) conducted a comprehensive investiga- 
tion of the Fort Ancient archaeological manifestation, and his data form 
the foundation for the descriptive and comparative analyses of this 
preliminary study. He establishes four foci in the Aspect — Baum, Feurt, 
Anderson, and Madisonville — on the grounds of artifactual similarities 
and differences; the same foci are accepted here as archaeological sub- 
groupings. The skeletal remains associated with these subgroupings are 


94 Indiana Academy of Science 

compared with each other, and with non-Fort Ancient groups, to denote 
the various degrees of physical homogeneity among the groups within 
the Aspect and to determine phyletic relationships with other groupings 
on a varietal level. 

The ethnohistorical identification of the earliest tribes found in the 
region is evaluated against the archaeological distribution in concordance 
with the temporal position of the Fort Ancient Aspect. While there is 
the possibility that the earliest historic tribes of the Middle Ohio Valley 
may have been derived from Siouan-, Muskogean-, or Algonquian- 
speaking stocks, it is most probable that the Fort Ancient people were 
Algonquian and therefore the ancestors of the historic Shawnee tribal 
group. Working with the assumption that the people in the area were 
affiliated with the Central Algonquian linguistic group, the cultural and 
physical correlations with other members of this division are explored to 
investigate the precise temporal and spacial placement of the population. 

When the identification of an archaeological population is being 
attempted, the investigator utilizes the morphological and metrical data 
of the skeletal remains that are available to him. The morphological 
characteristics serve as basic units since they more readily express the 
features that are typical of the group. The metrical dimensions primarily 
yield data on size rather than form, and the indicial units pertain exclu- 
sively to proportions, eliminating the size factor. If the morphological, 
metrical, and indicial traits are used together and their significant 
correlations noted, as is done in the present study, the population may 
be accurately described and identified in terms of trait combinations 
unique to it. 

The cranial material of the Fort Ancient population was collected 
by Georg K. Neumann of Indiana University during the time that 
Griffin was gathering the archaeological data for his report. The 
physical remains were so numerous that a comprehensive study of them 
was not feasible until computerized programming and facilities were 
available. A total of 732 individuals compose the Aspect sample, i.e., 
Baum Focus 40, Feurt Focus 71, Anderson Focus 118, and Madisonville 
Focus 503. Some components within each focus were excavated more 
completely than others, but the sample is considered to be more than 
adequate in providing a representation of the physical characteristics of 
the Fort Ancient people. 

In this preliminary study only the male crania of approximately 300 
individuals are examined. Since occipital cranial deformation is dis- 
played among a number of the crania, it is necessary to establish an 
undeformed and deformed category for the population of each com- 
ponent, with the exception of the components of the Anderson focus. 
The comparisons of each category are made by using Student's "t" test 
(1), a statistical means of determining significant differences between 
small populations and an important step in evaluating which traits 
would provide the most definitive information in a more comprehensive 
multivariate analysis. The assessment of population distance and infer- 
entially, the problem of local differentiation versus hybridization, is 

Anthropology 95 

expressed in terms of t-ratios and t-probabilities. An estimate of popu- 
lation distance is obtained from the ratio of significant differences found 
in the total number of variables for each measurement and index, being 
expressed as a coefficient of relatedness. Although the coefficients are 
based on a continuum, they are distributed proportionally in index 
classes to indicate near identity, close relationship, moderate relation- 
ship, and unrelatedness. This scale serves as a guide in determining 
whether crania from different components display homogeneous or 
heterogeneous physical characteristics, i.e., whether the crania represent 
similar or dissimilar physical populations (3). 

The cranial comparisons are made on three different levels — the 
intra-focus level, the inter-focus level, and the varietal level. On the 
intra-focus level, it is found that the components of the Baum Focus do 
not represent a single in-breeding population, but the series — the unde- 
formed and deformed crania — display a high degree of similarity in 
many dimensions and indices relating to traits that are commonly used 
to delineate populations. The crania from the various components of 
the Feurt Focus exhibit little physical variability, having a coefficient of 
relatedness of near identity. The series from the Anderson Focus display 
only a slight amount of physical dissimilarity and appear to be closely 
related. The crania of the components in the Madisonville Focus display 
less physical homogeneity in both undeformed and deformed groups 
than is found in the other foci. The degree of heterogeneity strongly 
suggests the presence of more than one physical variety in the focus. 

When the crania of each focus are pooled into a single "focus" 
population (maintaining undeformed and deformed categories when 
necessary), it is found that the undeformed crania of Baum, Feurt, and 
Anderson Foci appear to be closely related, displaying few significant 
differences in dimensions or indices. The deformed crania, however, 
exhibit a number of significantly differing values which may be influenced 
by the type and degree of deformation or by the presence of more than 
a single physical variety. The Madisonville crania appear to be only 
moderately related to the other foci, making it evident that more than 
one physical type was involved in the t-score comparisons. Hence, the 
crania of the Fort Ancient Aspect cannot be pooled to represent a single 
homogeneous population. 

An examination of all crania suggested that the undeformed indi- 
viduals could be sorted into an Ilinid or a Muskogid (Walcolid) category 
in accordance with the predominating physical characteristics of each 
skull in order to investigate the presence of different physical types 
in the Aspect. Smail (4) has previously shown that the Anderson popu- 
lation is closely related to the Oakwood Mound people, the "type" Ilinid 
physical variety; hence, there is strong evidence for building the Fort 
Ancient Ilinid population around the series from the Anderson Focus. 
Undeformed crania from the Baum Focus and some components of the 
Madisonville Focus also fit into the Fort Ancient Ilinid series. The 
Fort Ancient Muskogid series is composed of undeformed crania from 
the Madisonville Focus since there is little evidence of this physical 
variety in the other foci. 

96 Indiana Academy of Science 

When the Fort Ancient "derived varieties" are compared, they are 
found to be unrelated in dimensional values and only moderately related 
in indicial values. However the Fort Ancient Muskogid series display 
less of a population distance to the Fort Ancient Ilinid series than to any 
other varietal series to which it is compared. On the other hand, there is 
conclusive evidence that the Fort Ancient Ilinid series is nearly identical 
to the Ilinid physical variety and moderately related to the Muskogid 

According to various radiocarbon dates, the Fort Ancient culture 
covered a temporal span from about A.D. 1100 until historic times with 
the Baum Focus, and its Ilinid-like population, exhibiting greater 
antiquity than the other foci. The Madisonville Focus appears to be 
nearly as old as Baum and exhibits a temporal depth beyond the radio- 
carbon dates recorded for the Anderson-like components of Pleasant 
Hill and Erp. Such a time depth for Madisonville would account for the 
numerous Ilinid-like individuals in that focus and would suggest that the 
Fort Ancient Muskogid physical type was a late arrival in the Aspect, 
coming from the south. Therefore, it is proposed that the Fort Ancient 
Ilinid peoples represent the original inhabitants of the Fort Ancient 
cultural area which evolved from a Woodland base. 

The small sample of known Shawnee may not typify the entire tribe, 
but the coefficients of relatedness for the comparisons of the Shawnee 
and the Fort Ancient "type" series imply that the former represent an 
admixture of the predominant physical types in the area. Since some of 
the late prehistoric crania display similar kinds of admixture, it is 
believed that the historic Shawnee peoples are the descendants of the 
Fort Ancient archaeological population and are not recent comers to 
the Middle Ohio Valley. The fact that the Shawnee crania exhibit some 
Muskogid characteristics is considered to be merely an indication of the 
degree of admixture that is present in early historic times, but it is 
believed that the prehistoric Shawnee were more like the Fort Ancient 
Ilinid variety. The Ilinid affiliation of the Fort Ancient population, the 
early Shawnee, is substantiated by a recent study of a Fort Ancient 
series from Central Indiana that was made by Neumann. He found that 
the crania appear to be closely related to the Anderson population, or the 
Fort Ancient Ilinid variety. Thus, it appears that the Fort Ancient 
archaeological manifestation and the prehistoric Shawnee are 

Literature Cited 

1. Croxton, Frederick. 1959. Elementary Statistics with Applications in 
Medicine and the Biological Sciences. New York: Dover Publications, 

2. Griffin, James B. 1943. The Fort Ancient Aspect. University of Michigan 
Museum of Anthropology, Anthropological Papers, No. 28, Ann Arbor. 

3. Robbins, Louise M. 1968. The Identification of the Prehistoric Shawnee 
Indians — The Description of the Population of The Fort Ancient Aspect. 
Unpublished Doctoral Dissertation, Indiana University. 

4. Smail, J. Kenneth. 1965. The Uses of Female Crania in Demonstrating 
Racial Relationships. Unpublished Master's Thesis, Indiana University. 

Urban Anthropology and the Southern Mountaineer 

Ben R. Huelsman, Indiana University 


As a science Anthropology is a little over a hundred years old. In the 
nineteenth century, cultural anthropologists pioneered the concept of cul- 
ture and were primarily concerned with describing and recording the ways 
of life of so-called primitive peoples, such as the American Indian and the 
peoples of Africa and Asia. Since World "War II, many of these ethnic 
groups comprise the emergent, independent nations. Meanwhile, the twen- 
tieth century has caught up with Anthropology, and less than nine percent 
of the U.S. labor force is directly engaged in agricultural pursuits. Our 
cities are filling up with formerly rural peoples, American Indians moving 
to the city slums and away from federal reservations. Millions of blacks 
from the rural South have migrated northward to find their dreams of 
freedom frustrated in the black ghettoes of our major cities. This is a 
story of a less well-studied group of migrants, the southern mountain 
white, with whom I worked in two different slums for seven years. Since 
more than eighty percent of all Americans live in or near cities, it is 
apparent that a subfield of Urban Anthropology will emerge. 


The 1970 census will probably reveal that about eighty percent of 
U.S. citizens reside in the metropolitan areas of this nation. There seems 
to be little doubt that these urban areas now face a bewildering array of 
extremely serious problems. In an age of growing militancy of all kinds, 
the need for utilizing the best thinking of all the behavioral sciences 
toward the solution of these urban crises has never been greater. As one 
of the behavioral sciences, anthropology ought to be in a position to 
make many contributions to the solution of urgent national problems, 
especially those of our cities. In the last third of the twentieth century 
many detribalized ethnic groups will migrate to the slums of cities in the 
developing countries in ever-increasing numbers. In our own country, 
large numbers of American Indians have left their reservations to 
migrate to Chicago, Denver, and Los Angeles. It is quite predictable that 
they are met by police, employment officials and welfare workers who lack 
even the most rudimentary training in cultural anthropology. Here is at 
least one situation in some of our cities for which the professional 
anthropologist ought to be able to develop a working program of applied 
anthropology. In addition to growing numbers of American Indians who 
have migrated to our industrial centers, there are other ethnic groups 
whose presence in big city slums creates problems for both the cities and 
the migrants themselves. One of these groups is the southern moun- 
taineer. An estimated two million of them live in our midwestern indus- 
trial centers. Cultural anthropologists in the United States have studied 
the American Indian intensively for more than a century. With the 
exception of Prof. Marion Pearsall of the University of Kentucky, few 
anthropologists in this country have been concerned with studying or 
helping the southern mountain man and his family. 


98 Indiana Academy of Science 

Most of the graduate schools of anthropology in the United States 
do not offer a course or seminar in Urban Anthropology. One of the 
themes of this paper is that there should be such a course developed 
experimentally in departments of anthropology all over the United 
States. I visualize such a course as being offered in the evenings, not 
during the daytime hours, so that local government officials, policemen, 
social workers with no training in anthropology and other urban officials 
might attend. After I received my M.A. in anthropology I had the ex- 
perience of working for seven years in the cities of Cincinnati and Day- 
ton, Ohio with the southern mountaineer and his children. I often 
attended workshops and conferences sponsored by the Mayor's Friendly 
Relations Committee in Cincinnati, the purpose of which was to examine 
all aspects of the culture and urban adjustment of migrants to that city, 
both black and white. As it became generally well known that I was the 
only person in attendance with an advanced degree in anthropology, I 
became acutely aware of the image of anthropology in the United States 
which is held by such urban officials. It goes something like this. Anthro- 
pology is viewed as the study of primitive peoples, especially the Ameri- 
can Indians. The best known fields of anthropology are archeology . . . 
based on newspaper accounts of recent digs and ethnology, usually 
associated with the paperback books of Margaret Mead. To persons who 
have never studied anthropology, the field is seen as an impractical, 
romantic collection of interesting data about exotic, tribal groups. In 
other words, anthropology as they see it, cannot possibly have any 
real bearing on the kinds of practical problems such urban administrators 
and workers face on a daily basis. This is a stereotype, obviously. The 
point is that this stereotype is widely believed and acted on as though 
it were a reality by many responsible officials in major American cities. 
I might add that nothing remotely resembling a course in Urban Anthro- 
pology was ever offered at either the University of Cincinnati nor at the 
University of Dayton. It is quite true, that unlike Chicago or Denver, 
there are no American Indian migrant groups in the slums of these two 
cities, nevertheless the southern rural black and the white mountaineer 
arrived by the thousands every year in these cities. No anthropologist 
studied these groups. 

In my experience with the southern mountaineers in both Cincinnati 
and Dayton, Ohio, I noticed that their behavior varied according to the 
section of the city in which they lived. I also observed that these patterns 
of behavior differed in relationship to the length of time these people 
had resided in the city. One of the critical factors in determining the 
future adjustment of a mountain family to an urban environment is 
whether or not the male head of the family is able to find steady employ- 
ment. The folk class mountain parents I worked with tended to have a 
median level of education of about eight years of schooling in rural 
Appalachia. This is a male-dominant subculture in which the future way 
of life of the entire family is directly tied to the occupational success or 
failure of the father. Thus the success or failure of the father in finding 
steady employment in a factory or garage usually determined to some 
extent the future residence of the family in the city. Some made it to 

Anthropology 99 

home ownership in an inexpensive suburb, others, failing to find steady 
employment for some reason or other, entered the drab, barracks-like 
existence of welfare-recipients in a low-cost public housing project. By 
far the largest group of migrant mountaineers are the more recent 
arrivals to the city, usually found in an ageing slum district which is 
known as the port of entry. 

The basic premise of this paper is that there are three life styles 
to be found among the southern mountaineers in any urban community. 
Each life style consists of patterned behavior which differs from that 
observed in other parts of the city where the southern mountaineer clus- 
ters. The port of entry life style will be described first. The description 
will be as non-technical as possible to avoid cumbersome language and 
technical jargon. In Dayton, Ohio, for example, the port of entry slum 
district is the first part of that community with which the newly mi- 
grated southern mountaineer becomes familiar, while he is looking for 
a job. It is here that he hopes to break out of the intergenerational 
poverty that is choking to death so many of his kinsmen in rural Appa- 
lachia. Here is a sketch of the port of entry life style in Dayton. 

Port of Entry Life Style 

There is no deep sea harbor in Dayton and no plans to dredge one. 
There is no Ellis Island either to bring migrants from across the ocean 
in great sailing ships. But like Indianapolis, Dayton has a port of entry 
for its immigrants, and most of them speak the softly slurred speech of 
the rural South, not the tongues of Eastern Europe. There is no 
immigration service in the city to count their numbers but it is known 
they are both black and white, many of them making less than the pover- 
ty level of three thousand dollars a year. You won't find them at the local 
airport either. Most air travellers are middle class professional and busi- 
nessmen, not former sharecroppers and out of work coal miners. The 
migrant to Dayton usually gets here in a wheezing old car full of kids 
with hungry bellies and maybe a guitar. In the pocket of his faded, 
blue work shirt is a letter from a second cousin or an uncle with an 
address near Fifth and Brown or Tecumseh St. in East Dayton. This is 
in the heart of Dayton's port of entry, a strange new style of life for 
the southern white mountain migrant. He brings his wife, four, five or 
six children and about eight years of grammar schooling because he's 
heard from his kinsman that "they're hirin' on at NCR" or somewhere 
else. The southern mountain man in the port of entry is most likely to 
be from one of the mountain counties of Eastern Kentucky. His kinsman 
on Tecumseh or Brown St. has four or five young 'uns of his own, but 
he'll not turn down Dayton's newest migrant. They will all pile into one 
big flat, sleeping on couches, on the floor or in nice weather, maybe some 
of the kids will sleep out in the car. The family baggage is meagre . . . 
a few changes of old clothes, a toy or so, some snapshots, a few trinkets 
and souvenirs of life back home in the hills and hollows of Eastern 
Kentucky. The family has brought the most important baggage of all 
with them . . . the culture of the southern mountaineer, unfamiliar with 
city life. 

100 Indiana Academy of Science 

Most of the families in the port of entry hail from Eastern Ken- 
tucky, with a few from rural Tennessee and a handful from West Vir- 
ginia. The much-abused term "ghetto" hardly applies to this section of 
Dayton. Given any kind of choice, the migrants from Appalachia would 
probably decide they would rather live near each other. It's easy to 
come across friends and kin from the same county just a block or two 
away. They have the same background, share common values and the old- 
timers in the port of entry probably offer a lot of practical help to the 
just-arrived friend or kinsman from Eastern Kentucky. The newcomers 
soon learn where the flats for rent are, that you usually pay by the week 
and in advance. They also learn that it's pretty hard to get a Dayton 
landlord to rent to a family from Eastern Kentucky with a lot of kids, 
except of course in the port of entry. For a few weeks the newcomer 
and his family may live with a relative or friend while the father looks 
for a job. In the meantime his wife may be able to line up a temporary 
job as a waitress or barmaid, probably the first time she has ever been 
a wage earner. When this happens, the oldest daughter in the family 
usually takes the place of the mother, even if it means staying out of 
school. Mountain men and their wives seldom go out socially together. 
They each have their friends and acquaintances of the same sex as 
themselves. In the jargon of the social sciences, these are called refer- 
ence groups. If a man works on the assembly line at a plant, some of 
his cronies on a Friday or Saturday night are likely to be co-workers 
and others may be immediate neighbors. The fathers will probably get 
together on a regular basis at certain bars and at least one or more of 
them will be heavy drinkers. The middle class world of P.T.A.s joining 
the Junior Chamber of Commerce and involvement in community-wide 
projects is not for the southern mountain man. He is likely to be a 
union man and if he hasn't backslid, may belong to a storefront church, 
much like the ones they have down home. His religion is intensely per- 
sonal, promising immediate salvation and offering the kind of emotional 
release not to be found in a suburban church. The mountain man is 
proud and fiercely independent, but not much of a joiner, and he'd just 
as soon snoopy social workers left him and his family alone, as he 
doesn't like outside interference. 

The economics of the port of entry differs greatly from that of 
Kettering or Huber Heights in many ways. The southern white migrant 
is attuned to an existence of weekly payments. Local furniture and 
clothing stores encourage this deliberately and often charge the migrant 
family exorbitant rates of interest for their garish, poorly made prod- 
ucts. When the credit manager notices that only a few more weekly 
payments remain, the family is encouraged to add something else onto 
the bill, whether they need it or not. In an inflationary economy it is 
difficult for most people to develop budgeting skills even when the 
family size is small and the income adequate. It is even more difficult 
for the mountain family, unaccustomed to high prices, not used to han- 
dling the unexpectedly larger salary in the city and the attractive lure 
of easy credit. Slumlords know their customers well in the port of 
entry . . . all too well. They know the migrant is desperate for a 
place to stay, that most areas of Dayton will not rent to him on a 

Anthropology 101 

monthly basis if he has too many children and uncertain job prospects. 
Besides, if the migrant gives up in defeat and slips off quietly for East- 
ern Kentucky, at least the landlord will lose only a week's rent, not a 
month's rent. It is also true that coming from the grinding poverty of 
Eastern Kentucky, that the migrant is not likely to have much experi- 
ence in caring for his own home. Mountain youngsters are apt to be 
given the same kinds of unsupervised freedom they enjoyed back home, 
where there wasn't much else to do but roam free in the cool green hills 
and hollows. A boy or girl in Eastern Kentucky can do a lot of this 
without getting into trouble with the law, because there are few oppor- 
tunities for delinquent behavior in such a rural environment. The mi- 
grant newcomer to the port of entry doesn't realize he is bringing his 
children to one of the highest delinquency rate areas of Dayton. In 
East Dayton, the same casual parental supervision can lead to a police 
arrest and a referral to the Montgomery County Juvenile Court. 

There are few large cities in Appalachia, the rural areas are not 
heavily policed and a boy who gets into trouble in Eastern Kentucky is 
not likely to be arrested and hailed into a juvenile court. If a sheriff 
does arrest a boy in a mountain county he is likely to reprimand him 
on the spot or take him home where the sheriff will be personally ac- 
quainted with the boy's family. If a boy steals from a neighbor the two 
families are inclined to try to work it out in a personal, face to face 
manner, without calling in a constable or sheriff. Fights occur at roller 
rinks, outhouses get dismantled and sometimes tires or gasoline are 
stolen in rural Appalachia and some of this never gets recorded for the 
benefit of statisticians. Still, there isn't as much juvenile delinquency 
in rural Appalachia as in the port of entry slum of any large midwestern 
city. The sheer numbers and high level of efficiency of the Dayton Police 
Department ensure that a fair percentage of delinquent children in the 
port of entry will eventually be apprehended, especially the recidivists 
or repeaters. The schools, flats, streets and alleys of the port of entry 
are well known and heavily travelled by an army of police officers, social 
workers, probation officers and visiting teachers. 

When the mountain man brings his wife and children to Dayton he 
often betters himself financially by going from a chronically unem- 
ployed, hopeless way of rural poverty to a well paying, highly skilled 
factory job. But the real price he pays for pulling up his cultural roots 
in Eastern Kentucky and transplanting them to the port of entry is 
often paid by a son or daughter who someday winds up in the Boys 
Industrial School or the Girls Industrial School. In the city, justice is a 
vast, faceless bureaucracy of police, probation officers and social work- 
ers. They are all strangers and most of them uphold different cultural 
values from the southern white migrants. The person to person ways of 
handling family crises back home don't seem to work in the vast, 
impersonal sterile environment of the city. 

Welfare-Dependency Life Style 

In Dayton, Ohio there is a public housing project known as Parkside 
Homes. There are many southern mountaineers residing in this project. 

102 Indiana Academy of Science 

Their way or life is quite different from their fellow migrants in the 
port of entry. I'll call this the welfare-dependency life style. One of 
the basic requirements in most cities for welfare benefits is a residency 
requirement of not less than one year in the city. Unlike some of the 
migrants in the port of entry, the Parkside Homes mountain family has 
usually been in the city some time. The family is most often intact, con- 
sisting of both parents and anywhere from three to seven children. The 
presence of the male head of the household in the Appalachian migrant 
family makes the social structure of these families different from the 
so-called matrifocal family of the welfare-dependent black families. The 
role of the wife and mother in projects such as Parkside Homes is not 
really very different from her familiar role in rural Appalachia or in the 
port of entry. She is still the housekeeper, cook, laundress, main baby- 
sitter, nurse to younger children and all the other roles acceptable to 
mountaineer women regardless of where they live. For some of these 
mothers, the small welfare check makes possible a more stable supply 
of food than they have known since their marriage. I did not observe a 
great deal of voiced discontentment on the part of many mothers in 
Parkside Homes, even on the many occasions when I had the opportuni- 
ties to interview them alone. The children young enough to be in school 
do not seem to be essentially different in values or in behavior from 
similar children in the port of entry. The tendency to be bored with the 
sedentary school curriculum is a trait shared by most mountain youth, 
irrespective of the school attended. Parkside Homes is no stranger to 
the probation officers of the Montgomery County Juvenile Court and the 
impression is that the delinquency rate in this small housing project, if 
it could be accurately computed, is rather high. 

In my experience the member of the mountain family most dam- 
aged by the welfare-dependency life style is the father. Without a job 
or even the prospect of a job, the mountain man's life is almost without 
meaning. For this man Parkside Homes is a prison cell lined with velvet. 
The reasons why he cannot find employment in the urban labor force vary 
with each man ultimately. Sometimes it is failing health, in other cases 
functional illiteracy or alcoholism, sometimes a combination of reasons. 
The unemployed mountain man, especially if he gets a welfare check, 
serves unwittingly as the model for the city stereotype of the "lazy, 
shiftless hillbilly." For some reason the unemployed mountain man is 
not a target group for an army of civil rights organizations and Federal 
programs. His problems go unpublicized and as a group, the southern 
mountaineer is not organized. Many a good old country boy from Eastern 
Kentucky has arrived in the port of entry in Dayton, Ohio only to find 
himself in Parkside Homes a couple of years later, the empty years 
stretching endlessly ahead of him. Under such conditions and especially 
coming from dry counties in Eastern Kentucky, it is no wonder that 
so many turn up as alcoholics in the workhouse or Montgomery County 
Jail. The older teenage son of such a man always has the hope of join- 
ing some branch of military service when he drops out of school, espe- 
cially if the boy cannot find a job. Today's restrictive labor market, with 
its heavy emphasis on increased formal education, offers even less to the 

Anthropology 103 

high school dropout of both races than was true only two decades ago. 
The mountain lad who drops out of school today at age sixteen faces 
an even more dismal occupational future than ever. The situation be- 
comes even more hopeless when the armed services demand a high school 
diploma of potential enlistees. In effect this means that still one more 
traditional avenue of vertical social mobility is closing to the under- 
educated southerners of both races. The implications are that millions 
of our citizens have become technologically obsolete. 

The Assimilated Life Style 

Above Nettie Lee Roth High School on Hoover Avenue in Dayton, 
Ohio are two housing developments, namely Townview and the newer, 
brick subdivision of Western Hills. There, beginning in the 1960s, one 
will find recently arrived groups of blacks, enjoying their first taste 
of lower middle class suburban living. But for the most part the ma- 
jority of residents in these two developments are blue-collar whites and 
many of them are ultimately from rural Appalachia. A great many of 
them have spent their apprenticeships in Dayton's port of entry, have 
successfully found jobs, acquired increases in job skills and salary levels 
and bought their first homes. I lived in the Western Hills subdivision 
for two years and observed many of these people at first hand. The 
style of life of the assimilated mountaineer differs substantially from 
that of the southern mountain migrant in either the port of entry or in 
low cost public housing projects. The assimilated mountaineer is drawn 
more directly into the middle class mainstream of urban life than is 
true of the other two groups. Housekeeping standards, on the average, 
are higher than is generally observed in the decaying tenements of the 
port of entry. There is less juvenile delinquency and very little of the 
regimentation and hopelessness encountered so often in low cost housing 
projects. Except for the soft, slurred accents of the southern mountains, 
the preference for Protestant fundamentalism and country music, the 
assimilated mountaineer has lost much of his distinctive cluture. As long 
as his job lasts, the assimilated mountaineer will probably never again 
take up permanent residence in rural Appalachia, although on long 
weekends, he is likely to return to Eastern Kentucky to visit his kin 
and friends. 

In summary, I have argued that few anthropologists have taken 
the trouble to work in urban environments and even fewer yet are in- 
clined to study the southern mountaineer, either in rural Appalachia or 
as a migrant to our industrial centers. To what extent the southern 
mountain white tends to fall into three life styles in other cities can 
only be confirmed by further observation. I foresee a real and continuing 
need for the skills of anthropologists in meeting our urban problems. 
Does anyone agree ? 

A Systems Approach to the Study of Complex Society 

Melford S. Weiss 

Sacramento State College 

Early anthropological research was conducted among simple, non- 
literate, self-sufficient, hunting or quasi-agricultural communities which 
were habitually treated as cultural isolates — where contact with the 
outside-the-village world was generally regarded as inconsequential or 
at best accidental. Therefore, the society in question was invariably 
characterized as a functionally integrated and inter-related whole, a 
closed system with built-in homeostatic mechanisms which would remain, 
barring further contact, in a state of "dynamic equilibrium." 

As research interests shifted to the more complex peasant societies, 
a shift which demands new perspectives, the anthropologist, nevertheless, 
brought with him the theoretical models accumulated from tribal ven- 
tures. Robert Redfiekl (3) clearly established the peasant upon a folk- 
urban continuum; and although the peasant still retained the closeness 
of face to face primary relationships, a deep rooted and emotional at- 
tachment for the soil, and the warmth of Gemeinschaft mythology, there 
was the unavoidable added discomfort of a relationship to the greater 
outside world. Yet in spite of its extra-village contacts, the peasant 
community was generally treated as a geographically and culturally 
isolated entity whose behavioral dynamics could best be understood by 
intensive, internally-centered research activities. 

Today's complex rural and urban communities, in both the western 
and non-western world, can no longer be treated as cultural isolates 
within a closed system. Any study of changing community life must 
account for that community's relationship to the greater and invariably 
more complex outside world. Therefore, communities are open systems 
which cannot be described or analyzed in terms of static boundaries. 
The focus of outside contact and intervention cannot be ignored, for it 
is often the very nature of the relationship to the external world that 
accounts for many of the political, economic, social and ideological 
changes taking place within the village community. 

I am not disclaiming the importance of village studies, nor am I 
suggesting that the village is not, to some degree, a functionally inte- 
grated unit. However, it is time to take another look at the world's 
communities and to shift our emphasis from the internal aspects of the 
integrated village to a broader and more comprehensive understanding 
of the community and its dynamic linkage to the rest of the society. 

Because a village-centered study must now include extra-village 
structural ties, I would like to present one approach to the study of com- 
munity suggested by Professor Ishino, Donoghue, Marquis and Alchin 
(1) and Marquis (2) of the Institute for Community Development at 
Michigan State University. 


Anthropology 105 

The Systems Approach 

The systems approach provides both a method and a model for 
focusing upon a community and its connections to the greater society, 
because it views the community in terms of its interactions with out- 
side institutions. 

Following the model of the ecologists, the systems approach includes 
not only the people but also the man-made and natural components with- 
in the community; it focuses upon the interaction of the human 
components with the man-made and natural elements. 

A community "system" also includes kinship, economic, educational, 
political, religious and associational institutions which are linked not 
only to each other but to their counterparts in the greater society. 

Interaction between systems takes place through flows or movements 
of material, energy, people, and information. These flows are con- 
tinually entering and leaving the system. 

The Systems Approach and the Study of Billsburg 

In 1965 my wife and I completed a survey study of a semi-rural 
midwestern community with a population base of 2500 people. If the 
direction of our study had been internally centered and oriented in a 
restrictive and definitive temporal and spatial framework, instead of 
an open system, the linkages or channels which connect Billsburg to the 
greater outside world and the flows of material, energy, people, and 
information along these channels could have gone unnoticed, been ig- 
nored, minimized, "explained away" or more likely merely noted. It is 
because Billsburg was specifically viewed as a system constantly inter- 
acting with other systems at the local, state, national and even inter- 
national level, that the direction and nature of political, social and 
ideological change became meaningful and integral parts of the study. 

Flow Analysis as a Key Towards Understanding 
the Interaction Process Between Systems 

A flow is any movement of material, energy, people and informa- 
tion along any designated channel or pathway, either between different 
systems or within a single system. 

For example, when we first arrived in Billsburg, we spent the 
better part of the first day standing at the major highway intersection 
and observed the traffic patterns. Traffic in Billsburg is basically uni- 
directional; at 7:00 a.m. the concentration is westward toward Matson 
(a local urban industrial center), at 5:00 p.m. eastward back to Billsburg. 
The majority of the cars are newer models in the lower price field. Thus 
it was not surprising to later learn that approximately seventy percent 
of working Billsburgians are employed in the greater Matson area, that 
many of them are skilled and semi-skilled workers with average income 
of $7,000, and that Billsburg has overwhelming strong economic ties to 
the Matson area. 

106 Indiana Academy of Science 

As we began to interview, we discovered that leisure time activities 
are usually spent at movies, drive-ins, restaurants, bars, and bowling 
alleys, many of which are located outside the Billsburg area, and at the 
cultural, social, and athletic functions of Matson's State University. 

We also observed that Billsburg's all-purpose general store regu- 
larly subscribes to local, county, state and national newspapers including 
the infamous New York tabloids. 

While many major flows are inter-system oriented, internal (within 
the system) flows are also of considerable importance. My wife reported 
that women tended to do their daytime shopping at the larger shopping 
centers located outside the Billsburg city limits and near the outskirts of 
the greater Matson area, and that the downtown business area was 
poorly populated, causing the older Billsburg merchants some serious 
problems. We later found, as one might suspect, that many business 
enterprises are considering relocation closer to the Matson area. (4) 

Thus flows of material, people, and information are helping to 
establish continuing relationships between Billsburg and the greater Mat- 
son area, and, as previously mentioned, through news media, with the 
rest of the nation. Flow analysis should never be the end product of the 
research effort. It should be used as a methodological tool which is 
capable of providing much information, especially when research time- 
allocation is limited. When flow analysis is coupled with other research 
activities, it can provide immediate and valuable insights toward under- 
standing the dynamic characteristics of any community. 

The Expanding Nature of Contemporary Society 

Many problems arise within the community because the community 
itself is in a transitional (changing) stage. Many of the internal com- 
munity systems have expanded to the point where they are no longer 
subject to internal controls. This often results in organization and 
influence at higher administrative levels, and consequent conflict situa- 
tions arise which cannot be explained in terms of the village alone. 

If the Methodist Church Council in Detroit decided to send a Negro 
minister to Billsburg, (since Billsburg is still associated with the church 
hierarchy in Detroit, such a move is possible) the one hundred percent 
white Billsburg congregation would, to say the least, certainly be affected 
by such a decision. 

When we started our research Billsburg was undergoing the process 
of political upheaval. Recent studies indicate that the population base 
was changing due to a new influx of professionals, semi-professionals, 
university people, and Matson businessmen. These newcomers are of a 
more intellectual and liberal orientation than the older and more con- 
servative Billsburgians. Elections for city, township, school, agricultural 
and public safety offices are no longer controlled by the old city adminis- 
tration, for these ''newcomers" are actively participating in community 
affairs and are changing the nature of Billsburg's political and economic 

Anthropology 107 

institutions. Furthermore, these newcomers have maintained an active 
interest in state and national politics and are involved in political-social 
programs at the national and even international levels. Thus, decisions 
made in Washington, Saigon, Moscow, and London, are indirectly able 
to effect the local political ideologies and practices in Billsburg. (4) 

A Suggested Framework for Viewing the Changing Rural Community 

Scientific American (September, 1965), in discussing the alarming 
rate of urban growth in America, directs our attention to the creation 
of the Megalopolis: a concentration of linked urban centers that now 
stretches from Boston to Washington, D. C. If "megolopoly" is a poten- 
tial sign of America's future, let us not ignore the growth of the "smaller 
community ." These transitional centers are entering into dynamic rela- 
tionships with their local urban centers; and by this process, if I may 
introduce still another exotic terms, they are creating a "surbopolis": a 
concentration of linked communities about an already developed urban 


The anthropologist in his pre-occupation with a "holistic approach" 
has been primarily concerned with the social-institutional aspects of the 
community, village, or cultural system, and has neglected other possible 
research perspectives. If village studies are to expand these "limits of 
naivete," a more sophisticated approach along spatial dimensions is 
needed. While many anthropologists are clearly aware of the linkages 
and flows across and within systems and of their significance, too few 
studies have focused upon these aspects as integral parts of their in- 
vestigations. Today's communities no longer exist as "isolates"; the 
channels to other villages and cities have become the pathways to de- 
velopment, and the flows that pass along these channels are a key to 
deciphering the directions and processes of change, and a critical factor 
in understanding the complexities of twentieth century community life. 

Literature Cited 

1. Alchin, E., J. Donoghue, I. Ishino, S. Marquis. 1964. A Holistic approach 
to community development. Institute for Community Development, 
Michigan State University, East Lansing, Michigan. 

2. Marquis, S. 1962. The systems approach to community development. 
Institute for Community Development, Michigan State University, East 
Lansing, Michigan. 

3. Redfield, R. 1958. The Primitive world and its transformations. Ithaca, 
Cornell University Press. 

4. Weiss, M. S., and Weiss, P. H. Small town in transition. (In preparation 
for publication in Sacramento Anthropological Association Monographs.) 


Chairman : Hayward Campbell, Jr., Eli Lilly and Co. 
D. S. Wegener, I. U. Medical School, was elected chairman for 1969 

There was actually no meeting - of the Bacteriology Section this year. 
The paper by Petersen et al. was presented in the Cell Biology Section, 
but is printed here. Normally the Bacteriology Section of the Academy 
meets with the Indiana Branch of the American Society for Microbiology, 
because of the great overlapping of membership. This year the Indiana 
Branch met with the Ohio and Kentucky-Tennessee branches of the ASM. 
The abstracts of papers presented by the Indiana members at that meeting 
are included here. — Ed. 


Development of a Modified Antibody Plaque Technique for the Detection 
of Single Cells Making Anti-viral Antibody. B. H. Petersen, Z. Brahmi, 
J. S. Ingraham, and A. S. Levine, Indiana University Medical School. — 
Hemolytic plaque techniques detect the antibody produced by individual 
antibody-forming cells against various species of red blood cells (rbc), 
or even against haptens or proteins coupled chemically to rbc. As 
certain viruses adsorb spontaneously to rbc it was suggested that cells 
making anti-viral antibody might also be detected by a hemolytic plaque 
technique. Rabbits were given I. V. injections of Influenza virus strain 
WS-A. Suspensions of spleen cells obtained 3 days after the final injec- 
tion were examined for hemolytic plaque formation against Chicken rbc 
and Pigeon rbc coated with virus. These virus-coated rbc spontaneously 
agglutinated in our system. By use of our plaque assay in liquid medium 
(Fed. Proc. 26, 641, 1967) it was possible to observe both hemagglutina- 
tion inhibition and hemolytic plaque formation. Anti-WS-A antibody was 
demonstrated in the serums by hemagglutination inhibition. Specific 
plaques were also obtained from a rabbit immunized with Influenza strain 
Lee-B. These results demonstrate the feasibility of detecting anti-viral 
antibody produced by single cells utilizing virus-coated rbc. 

Amber Streptomycin-resistant Mutants of Escherichia coli. Karen Carl- 
son and Richard Bochrath, Indiana University Medical Center. — A 
specific genotypic defect such as the nonsense amber codon can cause 
premature termination of protein synthesis, at the site of a UAG 
codon, during translation. This defect may be mitigated by suppressor 
mutations which lead to the synthesis of specific altered transfer-RNA's. 
Streptomycin-resistant (Sm r ) mutants of E. coli WWU were isolated 
and characterized. Some Sm r mutants appeared to be amber streptomy- 
cin-resistant mutants: they were sensitive to streptomycin whenever cer- 
tain suppressor mutations were introduced. The existence of these 
mutants suggests that streptomycin resistance can be conferred by an 
incomplete protein. Experiments are in progress to determine whether 
this partial polypeptide is a ribosomal protein. 


110 Indiana Academy of Science 

Immune Response to Streptococcus faecalis in the Rat. B. PERI and M. 
Wagner, University of Notre Dame. — Rats immunized by various meth- 
ods with a formalinized suspension of Streptococcus faecalis (S.f.) strain 
ND547 were compared for agglutination titers in serum and saliva. 
Low levels of agglutinating antibody to streptococci are commonly found 
in serum and saliva of rats under the usual laboratory conditions since 
these organisms are part of the normal microflora. Response to paren- 
teral immunization was studied in rats of three categories: germfree, 
conventional and monoassociated with 5. faecalis. 

High serum titers were found in all (S.f.) monoassociated, but non- 
immunized, rats, indicating a strong immune response to gastrointestinal 
microflora alone. Immunized, monoassociated animals usually showed 
higher antibody levels than the nonimmunized monoassociates in both 
serum and saliva, but in some cases, high serum antibody level was not 
accompanied by high salivary antibody. Young rats show more antibody 
in saliva than in serum. It would seem that salivary antibody is not 
directly related to serum level. 

Germfree rats gave no immune response to immunization procedures 
which produced responses in both the conventional and monoassociated 
animals. In this case, the germfree animal, which represents removal of 
the antigenic competition of viable microorganisms, did not show the 
enhanced response anticipated. This phenomenon is being investigated 

Enzymatic Effect of Cobra Venom on Rauscher Leukemia Virus (RLV). 

A. C. Raitano and A. S. Levins, Indiana University School of Medicine. 
— The inactivation of RLV by Naja naja atra and Naja naja venoms 
was studied and an attempt made to identify the active virucidal factors 
in venom. Naja naja atra venom (150, 500, and 1000 Mg/ml sodium 
citrate) was incubated with RLV at 37° C for x k hour. The latter con- 
centration significantly decreased RLV infectivity. The RLV lipoprotein 
envelope contains 68% lecithin. Cobra venom contains a potent phospholi- 
pase A (PLA) that hydrolyzes lecithin to lysolecithin and a fatty acid. 
Therefore, several characteristics of PLA were used to identify its ac- 
tivity against RLV. PLA is heat stable at 100 °C for 10 minutes (pH 
5-6). Naja naja venom (1.5 mg/ml PBS) boiled for 10 minutes signifi- 
cantly inactivated RLV. Naja naja venom containing inactivated (boiling 
for 20 minutes) PLA did not inactivate RLV. PLA is also antigenic; 
therefore, Naja naja venom neutralization with specific antisera was 
studied. Naja naja venom (1.0 mg/ml PBS) inactivated RLV but neu- 
tralized venom (1.0 mg/ml PBS) did not. Thin layer chromatograms of 
venom on plasma phospholipids showed inhibition of PLA activity (leci- 
thin hydrolysis) by antisera. Lysolecithin (100 Mg/ml ELO) itself was 
also found to inactivate RLV while lecithin (100 Mg/ml H 2 0) did not. 
The levels of other venom enzymes were also examined. The results sug- 
gested PLA is an important factor in RLV inactivation by cobra venom. 

Suppression of Rauscher virus-induced Murine Leukemia by L-Aspara- 
ginase. W. F. Campbell and A. S. Levine, Indiana University Medical 

Bacteriology 111 

Center. — L-asparaginase from extracts of E. coli has been demonstrated 
to cause regression of several transplantable murine leukemias and 
lymphosarcomas. The leukemia cells, which apparently cannot synthesize 
L-asparagine, depend upon an exogenous supply of the amino acid. Nor- 
mal cells have high asparagine synthetase activities, and do not require 
asparagine. Apparently, asparaginase treatment depletes the plasma 
asparagine level, and deprives the leukemic cells of their exogenous 
supply of the amino acid. 

Investigations in our laboratory demonstrated that asparaginase 
treatment significantly suppressed splenomegaly in mice infected with 
Rauscher leukemia virus (RLV). The enzyme significantly increased 
survival time, but did not suppress viremia of RLV-infected mice. In 
small, multiple doses asparaginase altered the histological picture as seen 
in spleen sections, but did not completely inhibit the leukemic process. 
In apparent contradiction to the asparagine deprivation hypothesis, was 
our observation that asparagine treatment suppressed splenomegaly and 
prolonged survival. The results warrant a reappraisal of the asparaginase 
mechanism of antileukemic activity in RLV-infected mice. 

Physiological Studies of the Incorporatsion of 5-Bromouracil During 
Growth and Sporulation in Bacillus subtilis -168. PATRICIA C. MORGAN 
and Robert F. Ramaley, Indiana University. — Prior to an investigation 
of DNA synthesis and segregation during sporulation, studies were con- 
ducted on the physiological consequences of 5-bromouracil (5-BU) and 
5-bromodeoxyuridine (5-BUdR) incorporation in a thymine requiring 
strain of Bacillus subtilis-168. The results are summarized as follows: 

(1) The deoxynucleoside forms of thymine and bromouracil were in- 
corporated more rapidly and to a greater extent than were the free bases. 

(2) 5-bromouracil incorporation was stimulated by the addition of a 
small amount of thymine. This stimulation was also true for the deoxy- 
nucleosides of thymine and bromouracil. (3) Both thymidine and 5- 
bromodeoxyuridine were incorporated when they were both supplied to 
growing cells at a final ratio of BUdR/TdR + BUdR = 0.9. (4) Incor- 
poration of 5-BU or 5-BUdR for periods less than that inducing thymine- 
less death (30-60 minutes) resulted in fully viable cells. (5) Under these 
conditions the 5-BU incorporated was not selectively removed from the 
cells upon resuspension in medium containing thymine. This suggested 
that the 5-BU incorporated was not in false growing points or "unusual" 
DNA. (6) 5-BUdR incorporated during late vegetative or very early 
sporulation was included in the developing spores. Purified BU-labeled- 
spores showed the same viability as untreated spores. Thus, conditions 
have been obtained that allow the synthesis of apparently biologically 
active 5-BU DNA which is incorporated into the developing spores and 
does not decrease the viability of the completed final spores. 

Biosynthesis of Thiadiketopiperazine Antibiotics. D. R. Brannon, M. 
Gorman, B. B. Molloy, W. M. Stark, and J. Mabe, Eli Lilly and Com- 
pany, Indianapolis. — We have investigated the biosynthesis of the arano- 
tins, a new group of thiadiketopiperazines from Arachniotus aureus. The 
aranotins are structurally related to the microorganism metabolites 

112 Indiana Academy of Science 

gliotoxin and sporidesmin. Carbon-14 labeled amino acids were incubated 
with A. aureus to determine the extent of their incorporation into BDA- 
aranotin and to determine the biosynthetic relationship between the dif- 
ferent aranotins and gliotixin. Experiments were conducted to determine 
the source of sulfur in the thiadiketopiperazines and the relationship be- 
tween disulfide and thio methyl analogs. 

Mycophenolic Acid: Studies on Biological Activities. Robert H. Wil- 
liams, John C. Cline, Richard E. Holmes, and Martin J. Sweeney, 
Eli Lilly and Company, Indianapolis. The fermentation broth of a strain 
of Penicillium stolonif erum was found to possess reproducible in vitro 
antiviral activity. The active component was isolated by chloroform ex- 
traction of filtered broth at pH 3.0 followed by column chromatography 
on silica gel and crystallization. Comparison of the physical properties of 
the crystalline substance with those of mycophenolic acid led to its 
identification. Mycophenolic acid is one of the oldest known biologically 
active mold metabolites, first isolated by Gosio in 1896, and since then 
reported to have weak antimicrobial activity. The present study in- 
cludes some aspects concerning interesting new biological activities of 
this compound recently discovered in our laboratories, including antiviral 
and antitumor activity. Information concerning mechanism of action 
and the effects on modification of the basic structure to biological activity 
will also be presented. 

Genetic Evidence for Resistance of Cephalosporium to Specific Compounds. 

Paul A. Lemke, Eli Lilly & Co., Indianapolis. — One hundred compounds, 
antibiotics, antimetabolites and organic toxicants, have been surveyed 
for their toxicity to the antibiotic-producing fungus, Cephalosporium 
acremonium. This survey was designed to obtain compounds suitable as 
selective agents for resistant mutations and of potential use in the 
detection of somatic recombination through homozygosity of recessive 
mutations for resistance. 

Toxicity levels for several compounds were determined by a gradient- 
plate method, and mutations for resistance to certain compounds, princi- 
pally antimetabolites, induced. Selections for resistance were obtained 
from among survivors of 90% mortality after treatment with a chemical 
mutagen, nitrosoguanidine. Resistant cultures were selected on gradient 
plates, and mutation frequencies for resistances, spontaneous as well as 
induced, were calculated. Frequencies for induced mutations for resist- 
ance to specific compounds varied but were demonstrated to be of the 
order of 10 4 to 10 a . 

Among selections for resistance to antimetabolites no evidence for 
cross-resistance has been obtained, and markers for resistance to antime- 
tabolites have proved to be recessive by heterokaryotic tests. Data on 
selections for resistance to other compounds — specifically, actidione, acri- 
flavin, endomycin, and hydroxylamine — were less consistent. Cultures 
selected as resistant to these compounds grew poorly even in the absence 
of toxicant. 

Bacteriology 113 

In addition to the Cephalosporium organism seven other fungi 
(Penicillium chrysogenum, Emericellopsis glabra, Aspergillus nidulans, Sac- 
charomyces cervisiae, Sistotrema brinkmanni, Schizophyllum commune, 
Coprinus lagopus) and an actinomycete ( Actinoplanes utahensis) were 
compared for sensitivity to each of the hundred compounds. The 
prokaryotic nature of the actinomycete is implied by its reaction to 
specific compounds. The fungi examined were heterogeneous in response 
to the assembled compounds. 

Effect of Oxygen on the Synthesis of Nitrate Reductase in 
Bacillus stearothermophilus. JAMES H. NUNER and RONALD J. DOWNEY, 
Lobund Laboratory, University of Notre Dame. — We are investigating the 
biosynthesis of the membrane-bound electron-transfer enzyme, nitrate 
reductase (NaR). The oxygen-repressible synthesis of this enzyme in 
resting cells has been observed to begin shortly after the addition of 
nitrate. Cell division in cultures not originating in nitrate medium can 
be delayed for 10-12 hours. 

We have observed dramatic differences in the effect of imposing de- 
repressing oxygen tension (p0 2 ) on cells exposed to nitrate, with and 
without previous histories of exposure to nitrate. Cells without previous 
contact with nitrate die much more rapidly than those exposed to the 
same p0 2 in the absence of nitrate. Cells with previous contact with ni- 
trate lose viability during the first four hours and then recover. Turbidity 
of such cultures does not increase until recovery is complete. Other evi- 
dence also suggests that two populations of cells are present in the cul- 
ture: one nitrate-adapted, capable of using nitrate as a terminal oxidant, 
and the wild type, capable of using only oxygen. 

Oxygen is shown to permit growth of a nitrate-adapted culture and 
to promote the degradation of nitrate reductase to a basal level. 

The effect of different oxygen tensions on the synthesis of nitrate 
reductase in an adapted population was studied. A repressing tension of 
about 20 mm Hg was found. 

The Serum Profiles of Certain Reptile Sera and preliminary Observations 
on Antibody Formation in Snakes. Sylvia H. Kendall and S. A. Min- 
ton, Indiana University Medical Center. — Serum samples from several 
families of snakes, rock iguana (Ctenosaura) and rabbit were frac- 
tionated by zone electrophoresis (Pevikon) and gel filtration (Sephadex 
G-200). Zone electrophoresis of snake sera gave protein profiles differ- 
ing from the mammalian pattern and showed variations among the differ- 
ent snake families. No cathodic protein was detected in snake sera and 
the largest peak was not always the fast anode-migrating one, charac- 
teristic of the mammalian pattern. In addition, the electrophoretic profile 
of iguana serum differed from rabbit serum and all snake sera. No anti- 
genic relationship between iguana and snake sera could be shown by 
capillary precipitin tests. Gel filtration of rabbit serum gave a profile of 
three progressively larger peaks. Two out of three snake sera differed 
in that the first peak was largest and the third intermediate. Python 
serum gave two peaks of comparable size with a third peak suggested by 

114 Indiana Academy of Science 

a shoulder on the last peak. Peaks from fractionated snake sera, con- 
centrated and used in gel diffusion and Immunoelectrophoresis, were 
found to contain multiple components. Immunoelectrophoresis of whole 
snake sera resolved 7 to 11 components; 3 to 5 detected on the cathode 
side of the origin. Immunization of fox snakes (Elajjhe vulpina) with 
bovine serum albumin (Pentex Fraction V) in Freund's adjuvant pro- 
duced no antibody response one month after primary injection. Two 
weeks after secondary injection, snakes showed evidence of antibody 
response as measured by agglutination of erythrocytes coupled to BSA. 


Chairman: Thomas R. Mertens, Ball State University 
Robert L. Kent, Indiana Central College, was elected chairman for 1969 


A Study of the Factors Controlling Variation of Cuticular Characters. 
D. L. Dilcher and C. A. Zeck, Indiana University. — In this study we 
examined three factors which we thought might cause variation in the 
cuticular characteristics of leaves. These factors were leaf shape, leaf 
maturity, and the amount of sunlight that reached the leaf. The pur- 
pose of this study was to help resolve some of the controversy sur- 
rounding the use of cuticular analysis as a means of plant identification. 
The species investigated for differing morphology was Quercus alba, 
White Oak. Fagus grandifolia, American Beech, and Quercus rubra, 
Red Oak, were used for the study of sunlight versus shade leaves. Young 
leaves of these trees were later examined in order to understand the 
development of the epidermis. The cuticular characteristics looked for 
were the following: shape and size of the epidermal cells and stomata, 
presence and density of hairs, density of stomata, and pattern of the 
accessory cells. We found that neither the shape of the leaf nor the 
position on the leaf from which a cuticular sample was taken hindered 
the identification of the leaf cuticle. There were slight variations ob- 
served, however the variations were not severe enough to raise any 
doubts as to the species from which the cuticle had been obtained. Both 
of the species studied as sun versus shade leaves showed definite cuti- 
cular variations. There were many more stomata and the cells of the 
lower epidermis were much less lobed on the leaves that grew in the 
sunlight. The cells of the upper epidermis of the beech were more lobed 
on the shade leaves while the upper epidermis of the oak was the same 
in both sun and shade leaves. The other cuticular characters were similar 
on both sun and shade leaves. Upon studying the development of the 
epidermis in young leaves it was found that the mature situation of 
guard cells and stomata can be seen early in the development of a 
leaf while the development of the epidermal cells may follow different 
paths as the leaves mature. (Work supported by NSF GB5166) 

An Eocene Discovery of Dendropanax. G. E. Dolph and D. L. Dilcher, 
Indiana University. — Leaves of Liquidambar, Sterculia, Oreopanax, and 
Artocarpus were the only lobed forms reported from the Wilcox of 
Tennessee by Berry (1916, 1924, 1930). In addition to these forms, a 
form similar to Dendropanax (formerly placed in the form genus Aralia 
by Berry on the basis of a single specimen from Hardy Mills, Arks.) of 
the Araliaceae has been discovered. This form has the following charac- 
teristics: (1) three to five lobes; (2) shallow sinuses between the lobes; 
(3) size similarity between the main lobes; (4) entire margin; (5) 
acuminate to ovate lobes; (6) decurrent, truncate, or cordate base; (7) 
comptodrome secondary venation; and (9) papillate lower epidermis with 


116 Indiana Academy of Science 

randomly orientated anisocytic stomata. This form differs from Liquid- 
ambar which lacks an entire leaf margin, Artocarpus which lacks palm- 
ate venation, and Sterculia which lacks anisocytic stomata (fossil forms 
of Sterculia have anomocytic stomata) and reticulate tertiary venation. 
The distinction between Oreopanax and Dendropanax, both members of 
the Araliaceae, would be impossible if only the external morphological 
features of the leaves could be utilized. However a separation of these 
genera can be made by a study of the leaf cuticle. The lobed forms of 
Oreopanax have paracytic stomata while members of the genus Dendro- 
panax have anisocytic stomata. Since the fossil form also has anisocytic 
stomata in addition to the external features associated with Dendro- 
panax an assignment to the genus Dendropanax was possible. The as- 
signment of the fossil form to a living species of Dendropanax cannot be 
made due to evolution that has taken place within the genus. Although 
modern forms are similar to the fossil in external and stomatal features, 
most living species differ in their possession of sinuous upper epidermal 
cells and lack of papillae on the lower epidermis. (This work was sup) 
ported by NSF GB-5166.) 

Effects of Selenium on the Respiration of Excised Root-tip Segments of 
Maize. Raymond E. Girton, Department of Soils and Plant Nutrition, 
University of California. — This study deals with the effects of Na 2 Se0 3 
on the respiratory gas exchange of 1 cm root-tip segments cut from 3 
day germinated grains. Phosphate buffers were used to maintain a pH 
of 4.5 to 5. 

Typical Q 02 values for control (-Se) root segments were ca 5.6 
ul/hr/mg dry wt. and 5.0 for Qco 2 values. A 5xl0~ 6 M Na 2 Se0 3 con- 
centration appeared to give a 3 to 4% stimulation in 2 uptake over 
periods of 2 hours. High concentrations progressively decreased 2 up- 
take: 11% with 10- 3 M Na 2 Se0 3 and 30% with 10- M. Carbon dioxide 
output was somewhat depressed at lower selenite concentrations. High 
concentrations gave marked stimulation averaging about 30% above 
the controls. Respiratory quotient values here averaged 1.26 in contrast 
to ca 0.9 for the controls. Roots at high selenite concentrations turned 
light yellow to brown during the experiments; those at the lower con- 
centrations and the control roots remained white. 

Oxygen uptake by submerged roots in equilibrium with N 2 contain- 
ing traces of 2 was decreased to less than 1% of the controls, regard- 
less of the presence or absence of Na 2 Se0 3 . Similarly, the rate of C0 2 
production by selenite-treated roots in equilibrium with N 2 plus traces of 
2 was of the same magnitude as in the absence of selenite, viz. 81 and 
83% of the air controls. Fermentation/Respiration ratio values averaged 
0.83 in the absence of, and 0.70 in the presence of selenite and indicated a 
definite Pasteur effect. The lower (+Se) value reflects the aerobic stimu- 
lation of C0 2 output by Na 2 Se0 3 . 

Time-course studies demonstrated a gradual decrease in 2 uptake 
and C0 2 output rates for both the control and the 10 4 M selenite-treated 
roots in equilibrium with air. Roots treated with 10" M selenite indicated 

Botany 117 

a modest immediate stimulation in 2 uptake over the controls, followed 
by a steady decline to ca 50% of the initial value during the 3 hours of 
treatment. Carbon dioxide output rates of the controls and the 10* M 
Na 2 Se0 3 treated roots also declined with time. In contrast, the 10 2 M 
selenite-treated roots showed a very strong immediate stimulation in 
C0 2 output reaching an average of 153% of the initial value. This was 
followed by a rapid decline which continued to the end of the experiment. 
Respiratory quotient values ranged from a normal of ca 0.90 to a 
maximum of 1.84 for the 10 2 M Na 2 Se0 3 treated roots. This doubling of 
the R.Q. reflects both the stimulation in C0 2 output and the depression in 
2 uptake due to high concentrations of Na 2 Se0 3 . 

Some Disappearing Plant Species. Arthur T. Guard, Purdue University. 
— While in the very early history of the United States biologists were 
very much engaged in discovering new species of plants and animals, we 
may now be approaching an era in which the disappearance of species 
and conditions under which they disappear will hold the center of 
attention. Two species that seem excellent subjects for this type of study 
are Franklinia alatamaha Marsh, and Elliottia racemosa Muhl. 

These two species were first observed by John and William Bartram 
during their travels through Georgia about the year 1775. Franklinia 
alatamaha was discovered by these two botanists in the region of Fort 
Barrington, Georgia. It is no longer extant in nature, but has been 
preserved as a cultivated ornamental. The other species, Elliottia race- 
mosa, is still present, but it is found in only seven very limited areas in 
Georgia. Attempts to bring it into cultivation have been almost com- 
pletely unsuccessful. 

In view of the fact that other sympatric species observed by the 
Bartrams in this area are still abundant, the question arises as to why 
these two species have been almost completely unsuccessful. 

Weather and Corn Diseases in Indiana in 1968. A. J. Ullstrup, Purdue 
University. — Common corn smut caused by Ustilago maydis (D. C.) 
Cda. was more prevalent in 1968 than in any of the past 30 years. In 
June, 6-week-old seedlings showed large galls beneath, or at, the soil 
line. Such early symptoms haven't been reported in Indiana prior to this 
year. Corn planted in late May and early June after excessive rain in 
May showed neither the seedling galls nor the high prevalence of smut 
infection on adult plants. The unusually cold weather in May which 
slowed corn growth and thus held the meristematic tissues in a vulner- 
able state for a protracted period is believed responsible for the in- 
creased incidence of smut. 

Crazy top, a disease characterized by a condition of phyllody in the 
tassel as well as other bizarre symptoms, was also unusually prevalent in 
Indiana this year. Corn planted in late April and early May was, in 
many areas, subjected to heavy rains with consequent waterlogging of 
the soil. This condition is requisite for infection of corn by the causal 
agent Sclerophthora macrospora (Sacc.) Thirum., Shaw, & Naras. Corn 
planted in late May and early June didn't show evidence of this disease. 

118 Indiana Academy of Science 

Buggy-whip or onion leaf is a non-infectious malformation apparent- 
ly caused by excessive absorption of 2, 4-D. This condition, while ob- 
served in isolated instances almost every year, was widespread in much 
of the northern half of Indiana. In some fields the incidence reach 90%. 
The malformation is identified by the envelopment of upper portions of 
the plant in a tapering tube which often prevents tassel emergence. The 
predisposing conditions leading to this condition in corn are not fully 
understood, but the low temperatures in May are suspected. Corn show- 
ing this injury from over-absorption of 2, 4-D often had a high incidence 
of smut infection. 

Induced Resistance of Phaseolus vulgaris to Bean Anthracnose. J. E. 

Rahe and J. Kuc', Purdue University. — Etiolated hypocotyls of Phaseo- 
lus vulgaris respond hyper sensitively to Helminthosporium carbonum or 
Alternaria sp. within 24 hr. after inoculation. A similar response occurs 
at 60-72 hr. after inoculation with a non-pathogenic race of Colleto- 
trichum lindemuthianum, the causal agent of bean anthracnose. Necrotic 
lesions develop 72-96 hr. after inoculation of bean hypocotyls with a 
pathogenic race of C. lindemuthianum. Induction of hypersensitivity in 
bean hypocotyls by inoculation with H. carbonum, Alternaria sp., or a 
non-pathogenic race of C. lindemuthianum predisposes the inoculated 
tissues to resistance to a pathogenic race of C. lindemuthianum. 

Extracts of phenolics of hypocotyls inoculated with H. carbonum, 
Alternaria sp., or a non-pathogenic race of C. lindemuthianum are in- 
hibitory to both pathogenic and non-pathogenic races of anthracnose. 
This inhibitory property appears 6-24 hr. following inoculation with H. 
carbonum or Alternaria sp., and 6-24 hr. and again at 72-120 hr. after 
inoculation with a non-pathogenic but not a pathogenic race of C. 
lindemuthianum. The production of inhibitory concentrations of phenolic 
materials in response to infection by non-pathogenic races of C. 
lindemuthianum is suggested to account for the varietal resistance of 
bean plants to anthracnose. 

Lobelias of Franklin Co. and Indiana. Lloyd and Adele Beesley, Cedar 
Grove, Ind. — In the Flora of Indiana, Chas. Deam states that there are 
six species of Lobelia. In searching for Franklin County's and Indiana's 
wild Flowers, we have found all six species: Lobelia cardinalis, L. siphi- 
litica, L. puberula, L. kalmii, L. inflata and L. spicata. However, we 
have not found the white flowered form of Lobelia cardinalis f. alba 
which, according to Deam, has been reported from the Dunes by Peattie. 
Neither do we have all four well defined phases of L. spicata. 

Daily Variation in Chlorophyll Content of Corn Seedlings. Karen Kay 
Curtis and David E. Smith, Indiana State University. — Two groups of 
corn seedlings were grown for fourteen days under either normal day- 
night conditions of a greenhouse or a sixteen hours light-eight hours dark 
cycle in a growth chamber. The chlorophyll a and b concentrations of 
the first true foliage leaf were determined daily. Determination was 
based on the specific absorption coefficients of chlorophyll a and b in 
80% (v/v) acetone. Although there was a difference in the degree of 

Botany 119 

variation of the chlorophyll concentration of the plants grown under the 
two conditions, there was a striking similarity in the overall daily varia- 
tion of both chlorophyll a and b. The range of variation of total chloro- 
phyll content was from a minimum of 1.0 mg. of chlorophyll/ml. to a 
maximum of 1.8 mg. of chlorophyll /ml. Plotting either the daily varia- 
tions of chlorophyll a and b or total chlorophyll content resulted in 
graphs with maxima at 4, 8, and 12 days and minima at 3, 6, and 10 
days. The minima were shown to be significantly related statistically to 
the number of leaves unfurled at the 1% level. 

Other papers read 

The Origins of the Cultivated Peppers {Capsicum spp). Charles Heiser, 
Indiana University (by invitation). 

Pennsylvanian Coal Ball Flora of Indiana 

Robert W. Judd and Jerry J. Nisbet, Bethany Nazarene College, 
Bethany, Oklahoma, and Ball State University, Muncie, Indiana. 


This study was undertaken to further the understanding- of the flora 
of Indiana of the Pennsylvanian Period. While other workers have studied 
compression fossils of Indiana from the Pennsylvanian Period, the present 
study represents the first comprehensive work on coal ball plants. Coal 
balls are specialized calcium carbonate concretions which are sometimes 
found in coal seams. These concretions usually contain well-preserved 
petrified plant remains. 

Utilizing - the "peel technique," detailed analysis is made of selected 
coal balls which were collected in southwestern Indiana. The plants, or 
plant parts, are classified to genus, using the system proposed by An- 
drews. Five divisions are represented, including a total of 27 genera. 

The Pennsylvanian plant assemblage of Indiana, based on the present 
study, is compared to the Pennsylvanian plant assemblage of Illinois. 
Only minor differences in the represented genera appear, which suggests 
that a relatively uniform flora may have existed over portions of Indiana 
and Illinois during that Period. 

Methods and Materials 

During 1966 and 1967, collecting trips were made to the coal mining 
regions of Indiana. Most of the collecting was done during the summer 
of 1967 after analysis and sampling had revealed the most profitable 
collecting areas. 

Coal balls used in the present study were collected from eight sites 
table 1. Coal Ball Collecting Sites. 

Site 1 

T 5 S, R 8 W, Sec 10 

Warrick County 

Site 2 

T 5 S, R 8 W, Sec 14 

Warrick County 


T 1 N, R 8 W, Sec 13 

Pike County 

Site 4 

T 1 N, R 7 W, Sec 7 

Pike County 

Site 5 T 5 S, R 5 W, Sec 12 Spencer County 

Site 6 T 5 S, R 8 W, Sec 10 Warrick County 

Site 7 

T 14 N, R 8 W, Sec 27 

Vermillion County 

Site 8 

T 5 S, R 8 W, Sec 32 

Warrick County 

Site 9 

T 5 S, R 8 W, Sec 9 

Warrick County 


Botany 121 

in southern Indiana and the best specimens were then selected for 
intensive study. Table 1 shows the location of each collecting site. 

The coal balls were prepared for study by first cutting them into 
uniform slabs about % inch in thickness. The cutting was done on an 
18-inch Highland Park mechanically-fed diamond saw. Following the 
cutting process, the slabs were washed in detergent to remove the cool- 
ant. After drying, the material was prepared for study by using the 
peel technique. 

With this technique the slabs were first etched in a dilute (5-10%) 
solution of hydrochloric acid and allowed to air dry. The etched surface 
was then flooded with acetone, and a sheet of cellulose acetate film 
(.005") was immediately placed on the wet surface. By rapidly rolling 
the acetate film onto the slab, nearly all air bubbles were eliminated. 
For routine operations the slabs were then allowed to air dry for twenty 
to twenty-five minutes. When dry, the film was peeled from each slab. 
The organic material of the preserved plants which adhered to the film 
provided the thin layer of plant material necessary for microscopic 

The materials were cataloged and the acetate peels were 
examined under a 20X binocular microscope. Desired portions were cut 
from the peels, cleared in xylene, and mounted in Kleermount on micro- 
scope slides. The microscope slides prepared by this procedure were 
carefully analyzed and served as the source of data for the present study. 
The coal balls, acetate peels, and microscope slides were then stored as 
a reference collection at Ball State University. 


The classification system used in the present study is the system 
established by Andrews (1). Five divisions of plants are represented in 
the present study. 


The lycopods of the past were a rich and varied group which exhibited 
far greater dominance than do the modern forms. Whereas the existing 
lycopods are mostly creeping or herbaceous forms, many extinct members 
of this group attained tree size and developed complex reproductive 
structures approaching the seed habit. The two dominant families are 
the Lepododendraceae and Sigilliariaceae. All lycopod material in the 
present collection is placed in the Lepidodendraceae, although other 
workers might place some of the specimens in the Sigillariaceae because 
of the great similarity which exists between the organs of the two 

Probably the genus most frequently encountered by researchers is 
Lepidodendron. This genus includes many species. Even though all 
these organs have never been correlated for a single species, the genus 
Lepidodendron is very well understood. 

122 Indiana Academy of Science 

The leaves of Lepidodendron are long and grass-like and vary a 
great deal in size from one plant to the next. As the leaves fell from 
the stem they left a characteristic scar which persisted even on very old 
trunks. The cones are basically similar to those of living lycopods, but 

The anatomy of the stem of Lepidodendron is rather complex. Noth- 
ing comparable to secondary phloem was produced in the steles. Sur- 
rounding the stele is an extensive cortex that was the site of abundant 
periderm formation as the tree matured. This periderm, unlike the cork 
of modern trees, remained alive and meristematic. As it became thicker, 
splits developed in the surface and eventually the outer portions of 
periderm were sloughed off. Since the stele of the stem never attained 
great size, the large diameter of some trunks was due mainly to the large 
periderm region. 

In material from the present collection, the stem of Lepidodendron 
is represented by one nearly entire stem specimen and several stele frag- 
ments. The nearly entire stem (Figure 1) illustrates the exarch 
protostelic nature of the genus, but exhibits very little of the periderm 
region. In another specimen some traces of the periderm region are 
found in association with what is believed to be a leaf base. 

The underground portions of Lepidodendron are placed in the genus 
Stigmaria. Two closely spaced dichotomies occurred at the base of the 
trunk resulting in the formation of four main rhizomes. The small roots 
which arise from the ultimate branches of the rhizopore are protostelic. 
The small vascular trace of the root is composed of 8-10 tracheids, and is 
surrounded by three distinct regions of cortex. In the distal portion of 
the root, the middle cortex is not found, having presumably undergone 
natural disintegration. The vascular trace and small inner cortex are 
then found somewhat eccentrically suspended by small remnants of 
cortical tissue called trabeculae. 

The present collection contains many specimens of Stigmaria roots. 
Several specimens have been isolated which exhibit the three distinctive 
cortex regions and are thus believed to be sections from near the 
rhizophore. Another section (Figure 2) shows the characteristic arrange- 
ment of tissues which results when the middle cortex disintegrates and 
the vascular tissue is displaced. 

Although the size of the leaves varies from species to species, and 
even on the same plant, they remain similar in structure. The leaves are 
long and linear, with a prominent midrib projecting on the abaxial sur- 
face. The single, centrally-located vascular bundle is composed of a 
group of tracheids surrounded by thin-walled cells which probably repre- 
sent phloem tissue. The blade of the leaf contains rather extensive thick- 
walled hypodermal cells under both the upper and lower epidermis, 
between which is found a loosely arranged photosynthetic tissue. 

The previously indicated widespread occurrence of Lepidodendron in 
Indiana is further substantiated by the great number of Lepidophylloides 



Figure 3 

Figure 4 

124 Indiana Academy of Science 

found in the present collection. Some specimens show the vascular 
bundle and sheath quite well (Figure 3), but no leaves were found with 
sufficient internal preservation to illustrate phloem tissue. This specimen 
also shows the hypodermal regions and portions of the spongy mesophyll. 
All specimens found illustrate the prominent midrib on the abaxial 
surface of the leaf. 

Many species of cones have been described which are borne on 
Lepidodendron; most of these have been assigned to the genus 
Lepidostrobus. These cones vary in size over a considerable range, with 
specimens averaging 13 mm. in diameter and 15 cm. in length commonly 
found. Some exceedingly large specimens 8 cm. in diameter and over 
30 cm. long are occasionally encountered. The cones are borne terminally, 
with the sporophylls spirally arranged about a central axis. Nearly all 
the cones found have been heterosporous, with both unisexual and disexual 
cones found. The sporangia, borne on the adaxial surface of the 
sporophyll, are elongate and nearly circulate in cross section. The distal 
portion of the sporophyll, called the lamins, is upturned and overlaps 
the sporophylls above, forming a protective outer covering. 

Portions of the cone axis and sporophylls are found only occasionally 
in the present material, which would perhaps indicate that the cones 
were not so widely distributed as the other organs of this plant. The 
cross section of one cone shows the sporophylls grouped about the axis 
in an orderly arrangement. In another specimen (Figure 4) the vascular 
tissue of the axis is shown and some of the sporangia may be observed to 
be filled with microspores. 

While many cones of Lepidodendron contain a large number of 
spores in each sporangium, others contain as few as four. The genus 
Lepidocarpon, established by D. H. Scott in 1901, has only one functional 
megaspore in each megasporangium, although three aborted megaspores 
are associated with the functional one. When mature, two lateral alations 
of the sporophyll grow upward to almost completely envelop the 
sporangium, leaving only a slit-like opening at the top. The resulting 
structure is a seed in the sense that it is an integumented mega- 
sporangium with a slit-like opening in the top, analogous to the micropyle 
of a modern seed. 

Lepidocarpon "seeds" are very abundant in the present material. 
Although clusters of the "seeds" are found, they are not found in 
organic contact with each other or with any other organ. Many of the 
present specimens show the enveloping alations and slit-like opening at 
the top (Figure 5). In some longitudinal sections of the "seeds" the 
sporangial walls can be seen distinctly separate from the sporophyll 


The division Coniferophyta contains two orders, the Cordaitales and 
Coniferales. The former was a dominant group of Pennsylvanian seed 
plants from which the Coniferales are believed to have evolved. Rapid 



Figure § 


Figure T 

-Figure 8 

126 Indiana Academy of Science 

evolution and diversification of the Conife rales apparently occurred during 
the Permian Period. Some of the modern coniferous trees have changed 
little from their Permian ancestors. 

Members of the Corditales were large trees, often seventy-five to 
one hundred feet in height, with a crown of large, strap-shaped leaves. 
Although Cordaites is the name proposed first for the leaves of the tree, 
researchers include more of the plant organs within the genus Cordaites 
as the affinities of the organs become known. 

The stem of Cordaites contains a chambered or septate pith which 
is made up of closely spaced plates of parenchymatous pith separated by 
air spaces. This distinctive and characteristic pith is very similar to that 
found in the modern genus Juglans. Outside the xylem the stem pro- 
duced a narrow band of cortical tissue, encircled by a zone of periderm. 

The Cordaites leaf is superficially very similar to a modern Iris 
leaf except in size. Specimens of the leaves have been found which 
attained a length of one meter and a width of nearly 15 cm. The veins 
of the leaf are dichotomously branched, but the veins ascend so steeply 
that the venation pattern appears to be parallel. The structure of the 
individual bundles is identical to that of the vascular bundles in modern 
cycads. In some specimens of Cordaites leaves the mesophyll is undif- 
ferentiated, while in others the mesophyll is arranged into a palisade-like 
upper region and a more loosely arranged lower region. The overall 
structure suggests a tough and stress resistant leaf. 

Cordaites leaves are quite abundant in the present collection, sug- 
gesting that Cordaites must have been one of the prominent plants of 
the Pennsylvanian swamps in Indiana. Most of the leaf specimens in 
the present material display relatively poor internal preservation. Figure 
6 shows a typical Cordaites leaf in cross section. The vascular bundles 
with their hypodermal "ribs" are well illustrated in this specimen. The 
evenly spaced vascular bundles indicate the nearly parallel venation 

The reproductive structures of Cordaites are assigned to the genus 
Cordaianthus. The phrase "dwarf shoot" is accurately applied to these 
reproductive structures to distinguish them from the cones of modern 
conifers. Each shoot is 4-6 mm. long and is made up of twenty to thirty 
closely inserted scales which are spirally arranged around the central 
axis. The most distal scales of the microsporangia dwarf shoot are 
fertile, each bearing about six sporangia. In the megasporangiate shoot 
only one fertile scale is found, bearing a single ovule, which is generally 
hidden by the overlapping sterile scales. It is readily apparent that 
Cordaianthus bears little resemblance to the cones of modern conifers. 

Cordaianthus is represented sparingly in the present collection. A 
single specimen illustrates a tangential section of Cordaianthus which 
shows the sterile scales, but no fertile ones. 

Most of the isolated seeds attributed to Cordaites are assigned to the 
genus Cardiocarpus. Nearly all the seeds of the Pennsylvanian age 

Botany 127 

which exhibit bilateral symmetry in cross section are arbitrarily placed 
within this genus. 

Several good specimens of Cardiocarpus are represented in the pres- 
ent collection. Figure 7 shows a longitudinal section of Cardiocarpus 
spinatus, illustrating the projections of the sclerotesta and traces of the 


A burst of evolution in the Arthrophyta division during the Pennsyl- 
vanian Period produced some of the most unique plants that have ever 
lived. The great columnar trunks of Calamites with their whorles of 
branches, perhaps resembling a giant-size Equisetum, surely must have 
been a striking feature of the Pennsylvanian landscape. While Arthro- 
phyta is a very diverse division, certain characteristics tend to bind its 
members together as a unit. The chief characteristics of the arthrophytes 
are their ribbed and jointed stems, and the whorled arrangement of the 
leaves and sporangia-bearing organs. A rapid decline in the number and 
variety of arthrophytes apparently began during the Permian Period, 
and only a single genus, Equisetum, survives today. 

The two major orders of Arthrophyta are Sphenophyllales and 
Equisetales. The latter is made up of two families, the Equisetaceae 
containing the fossil representative. 

The family Calamitaceae contains the arborescent arthrophytes. The 
largest members of this family were trees well over fifty feet in height. 
While the isolated organs of the members of this family are assigned 
generic and specific names, the organs are believed to belong, for the 
most part, to a single, well denned plant named Calamites. 

The stem structure of Calamites is generally similar to that of 
modern Equisetum. The larger stems have a pith cavity and strongly 
developed secondary wood. Arthropitys, the most frequently encountered 
stem genus, exhibits secondary wood which is divided into sectors by 
rays only one or two cells wide. In Calamodendron, the rays are bordered 
by vertically aligned fiber cells. In the third genus, Arthroxylon, fibers 
are densely interspersed within the rays. 

One fairly complete stem specimen of Calamites is represented in 
the present collection which illustrates the pith and secondary wood in 
somewhat crushed condition. Many isolated wood fragments are char- 
acteristically found in coal ball petrifactions, and analysis of some frag- 
ments in the present collection have shown them to represent Arthro- 
pitys. Other fragmentary wood specimens from the present collection 
indicate the possibility of the presence of additional wood genera. 

The anatomy of Calamites roots, assigned to the genus Asteromyelon, 
is different from that of the stem. The roots lack the jointed feature of 
the stem, and a well preserved pith is usually present. The cortex is 
rarely preserved in the roots, but when present, large cavities similar 
to the canals found in the cortex of Equisetum are observed. 

128 Indiana Academy of Science 

Several sections from the present collection illustrate internal 
anatomy which is characteristic of Aster omyelon. The cortex is not pres- 
ent in any of the present sections, making positive identification difficult. 
The large well preserved pith and typical wood development are evident 
in several specimens. On the basis of the pith and wood features, the 
present specimens are assigned to Aster omyelon. In some of the sections 
(Figure 8) a branch root can be seen emerging from the main root. The 
relative scarcity of specimens from this genus seems to indicate that the 
roots are not widely distributed in Indiana. 

Cones which are believed to have calamitean affinities vary a great 
deal; however, all have the characteristic whorled arrangement of 
sporangiophytes. Calamostachys, the most common petrified genus, has 
suggestive whorles of sporangiophores alternating with whorles of 
sterile bracts. The sporangiophores are pelate (as in modern Equisetum) 
with the sporangia oriented toward the cone axis. 

In the present collection, specimens of Catamites cones are found in 
small numbers and are poorly preserved. The author found one speci- 
men believed to be a sporangiophore with fragments of the pelate 
sporangiophore. Based on this evidence, the specimen is assigned to 
Calamostachys. The lack of additional material suggests a rather sparse 
distribution of the cones of Catamites in Indiana. 

The order Sphenophyllales contains only one genus, Sphenophyllum, 
although several other generic names are applied to the cones of this 
plant. The growth habit has been variously interpreted as aquatic, 
creeping or climbing; and erect, terrestrial and self-supporting. The 
leaves of Sphenophyllum were borne in whorles at the nodes of a 
slender, ribbed, and jointed stem. Each leaf was somewhat wedge-shaped 
and attached to the stem at the narrow end. The broad end of the leaf 
was variously cleft or notched. The leaves were nacrophyllous, with 
from three to many vascular bundles in each leaf. While nearly all 
knowledge about the leaves of Sphenophyllum, has been derived from the 
study of coal balls. 

The stem of Sphenophyllum is protostelic, and the wood is unlike 
that found in any other plant. The center of the stem contains a tri- 
angular group of primary tracheids with small protoxylem cells located 
at the apices of the primary wood. In older stems a considerable amount 
of secondary wood is found. The tracheids of the secondary xylem 
which are located outside the apices of primary wood, are smaller than 
the remaining tracheids of the secondary wood. Some specimens have 
been found with a cortex of thin-walled cells, but the cortical tissue is 
not generally preserved. 

A fairly widespread distribution of Sphenophyllum during the Penn- 
sylvanian times is suggested by the many good preservations of the 
stele region of the stems in the present collection. Figure 9 illustrates 
the triarch protostele which characterizes the stem of Sphenophyllum. 
The structure of the secondary wood may easily be seen, and the smaller 



Figure 9 

Figure 10 


Figure 11 

Figure 12 

130 Indiana Academy of Science 

tracheids outside the protoxylem are clearly evident. This section also 
illustrates the small ray cells among the tracheids of the secondary wood. 

The roots of Sphenophyllum lack the ribbed and jointed features of 
the stem. The internal anatomy of the root is similar to that of the 
stem, except that the protostele is diarch instead of triarch. 

Several specimens of Sphenophyllum roots are represented in the 
present collection, although they are not encountered so frequently as 
specimens of the stem. The diarch protostele of the root is clearly evident 
in one specimen and the secondary wood appears similar to that of the 
stem. The cortex region is also shown in the specimen. 


The plants in this division exhibit a combination of two character- 
istics which are unique and quite interesting. The plants possess fern 
foliage and bear small nut-like seeds. The combination of these two 
characteristics gives rise to the name Pteridospermophyta or seed ferns. 
While joined by these two unifying characters, the seed ferns were in 
other ways quite diverse and were widespread in the swampy regions 
during the Pennsylvanian Period. 

Two classes of pteridosperms are recognized, the Lynginopteridaceae 
and the Medullosaceae. One member of the Lynginopteridaceae is 
Callistophyton. The stem of this plant contains a large pith region of 
very thin-walled cells. Remnants of the primary xylem tissue can often 
be observed at the periphery of the pith, followed by a broad zone of 
secondary wood. A considerable amount of secondary phloem was pro- 
duced outside the xylem, but this tissue is seldom preserved. Periderm 
was probably produced on the outside of the stem, but little positive 
evidence has been advanced to support this idea. 

The stem of Callistophyton is represented by a single specimen in 
the present collection. The large pith region is obvious (Figure 10), and 
traces of primary xylem may be observed at some points around the 
periphery of the pith. The well developed zone of secondary wood with 
its narrow rays can also be seen in this specimen. 

The class Lygihopteridaceae contains a variety of seeds. Physostoma 
is a small seed, about 6 mm. long and 2.5 mm. in diameter. The cup- 
shaped pollen chamber is very distinctive, and may contain a large 
number of pollen grains. The integument of the seed is fused with the 
nucellus except at the apex. At the apical end the integument is extended 
beyond the body of the seed into ten to twelve projections. The outer 
region of the integument is composed of brick-shaped cells arranged 
radially, while the inner region is composed of longitudinally arranged 
cells. Large unicellular hairs are sometimes found attached to the out- 
side of the integument. Conostoma is the genus of a similar, but slightly 
smaller seed than Physostoma. The structure of the integument is 
fused with the nucellus except at the apex. No projections of the 
integument are present in Conostoma. 

Botany 131 

Many seeds are represented in the present collection which are prob- 
ably assignable to the Lyginopteridaceae. Considerable difficulty was 
encountered in attempts to identify these seeds. The general literature 
does not provide adequate information to permit separation of genera of 
seeds, particularly if the entire specimen is not present. For this reason, 
the classification of the two genera below is tentative. 

Both Physostoma and Conostoma are represented in the present col- 
lection. Figure 11 shows a longitudinal section of Physostoma which illus- 
trates the arrangement of cells in the integument. The distinctive pollen 
chamber and the projections of the integument are not shown because 
a suitable section could not be made. Conostoma seen in cross section 
illustrates the typical ovoid shape. Remains of the nucellus may be 
observed in this specimen. 

The family Medullosaceae is identified chiefly by the presence of a 
polystelic vascular system in the stem. Nearly all members of this fam- 
ily are assigned to Medullosa which is evidently a genus of very broad 
limits. The foliage borne on the stem was of the Alethopteris or 
Neuropteris type and, when seen in cross section, the large petioles of 
the fronds bear a considerable resemblance to the stem of modern Zca. 
The seeds associated with Medullosa are larger than those of the 

The stem of Medullosa contains from 3-27 small oval protosteles 
surrounded by a broad region of cortical tissue. The stem itself repre- 
sented only a small part of the total size of the Medullosa trunk. The 
enveloping petiole bases account for most of the diameter of the large 

Isolated petiole bases are frequently represented in coal balls, and 
are assigned to the genus Myeloxylon. The petioles are quite large, com- 
monly 4 cm. or more in diameter where they grow free from the stem. 
Myeloxylon contains many vascular bundles scattered through a zone 
of thin-walled pith cells. The bundles, derived from repeated divisions 
of the main leaf traces consist of several xylem elements surrounded by 
a bundle sheath. Phloem tissue is not usually preserved in the bundles. 
The outer region of the petiole consists of a hypodermal zone of thick- 
walled fibrous strands, among which secretory canals may occasionally 
be observed to occur. 

Myeloxylon is represented frequently in the present collection. The 
hypodermal zone and scattered arrangement of the vascular bundles 
can be seen in several specimens. The overall resemblance of Myeloxylon 
to the stem of Zea is evident. 

The foliage of Medullosa is of the Alethopteris or Neuropteris type. 
The differences between these genera, established mainly on the basis 
of compression fossils, are mainly differences in the venation pattern and 
the mode of attachment of the leaflets to the rachis. In cross section, the 
two genera are quite similar except that the midrib of Alethopteris is 
more prominent than that of Neuropteris. 

132 Indiana Academy of Science 

As seen in cross section, Alethopteris leaves are revolute or curled 
at the margins. The parenchymatous mesophyll is palisade-like in the 
upper region and more loosely arranged in the lower portion of the leaf. 
The midrib is very prominent on the abaxial surface, and the main vas- 
cular bundle it contains shows the same structure as the bundles in the 
Myeloxylon petiole. The secondary vascular bundles exhibit the same 
tissue arrangement as the main bundle, but have some fibers associated 
with the xylem cells. 

Two genera of seeds are normally found in association with 
Alethopteris and Medullosa. Pachytesta seeds are quite large, some 
reaching a length of 6 cm. or more. The seed is more or less circular in 
median cross section and becomes somewhat triangular toward the 
micropylar end. In overall general appearance, Pachytesta resembles a 
pecan seed. 

Two specimens of Pachytesta are represented in the present collec- 
tion. One specimen, representing a small variety of seed, clearly illus- 
trates the structure of the integument. The nucellus of this specimen is 
free from the integument, and the conspicuous ribs of the integument 
are also evident. The second specimen, representing a much larger 
variety, was found partially exposed at the surface of a weathered coal 
ball. The seed was nearly complete, including the integument, and was 
5 cm. long and about 2.5 cm. in diameter. Although the external preser- 
vation of the seed was good, examination revealed little internal 

Stephanospermum is another well-known seed believed to be borne 
on Medullosa. Seeds of this genus are comparatively small, usually 
about 10 mm. long and 5 mm. in diameter. The most striking feature of 
Stephanospermum is the presence of a well-developed collar of the 
sclerotesta around the micropylar region. The apex of the pollen chamber 
extends well beyond the body of the seed. 

Several specimens of Stephanospermum are represented in the 
present collection. None of the specimens could be cut in such a way as 
to reveal the distinctive collar of the sclerotesta. In cross section, some 
of the integument structure can be observed, although the outer fleshy 
layer is not preserved in these specimens. The remains of the nucellus 
may be seen as a thin band around the inside of the seed. 

The microsporangiate organs of Medullosa are assigned to 
Dolerotheca. This is a complex organ, consisting of many tubular micro- 
sporangia imbedded in a cellular matrix. In overall appearance, 
Dolerotheca probably resembled a small wasp's nest, about 4 cm. in 
diameter and 20 mm. thick. When mature, this organ produced huge 
ovoid microspores, approximately .4 mm. long. Several fragments, 
believed to be the isolated microsporangia of Dolerotheca, are repre- 
sented in the present collection. No specimen of the entire organ was 

Botany 133 


The members of this division include both the true ferns and an 
interesting group of plants called preferns. The preferns began to 
evolve during mid-Devonian time into plants which are considered transi- 
tional between the earlier psilophytes and the later true ferns. Knowl- 
edge of preferns is based on a variety of fossil evidence from both 
compressions and petrifactions. 

Two orders of preferns exist, the Protopteridales and the 
Coenopteridales. It is unfortunate that the distinction between the two 
is highly artificial, the former known only from compression fossils and 
the latter from petrifactions. Considerable overlapping of the plants 
included in the two orders probably exists; but based on the present 
knowledge, this classification is the most expedient way to identify these 

Several families of the order Coenopteridales are represented in the 
present collection. The family Botryopteridaceae is represented by the 
genus Botryopteris. The stem of this genus is protostelic, about 12 mm. 
in diameter, and possessed a zone of cortex around the stele. Several 
petioles branched from the stem, each bearing a bundle with three 
xylem arms arranged in a characteristic W-shape. The primary petiole 
branches continued to divide and apparently terminated in slender, 
cylindrical branchlets. 

Isolated petioles of Botryopteris are represented frequently in the 
present collection. In Figure 12 the typical W-shape xylem arrangement 
of the vascular tissue, as well as the surrounding cortex region are clearly 

The family Anachoropteridaceae is represented by the genus 
Tubicaulis. The simple protostelic stem of this plant gave rise to many 
spirally arranged petioles. The unusual feature of this plant is that the 
petiole, as seen in cross section, contains a C-shape vascular trace with 
the open side of the bundle pointing toward the abaxial surface. Very 
little is known about the remaining organs of this interesting plant. 

Many isolated petioles of Tubicaulis are represented in the present 
collection. The C-shaped vascular tissue with its abaxial orientation are 
shown in several specimens. The cortex surrounding the vascular tissue 
is also quite apparent. 

The family Zygopteridaceae is represented by the petiole genus 
Etapteris. The distinguishing feature of this genus is the H-shaped 
arrangement of the vascular tissue. In recent years, Etapteris petioles 
have been definitely correlated with a more completely understood plant 
called Zygopteris. 

Only a few specimens of Etapteris are represented in the present 
collection. Figure 13 illustrates the petiole with its H-shaped xylem 
tissue. The cortex which surrounds the vascular tissue in this genus is 
similar to the cortex of the Botryopteris petiole. 


Indiana Academy of Science 

Figure 13 

Figure 14 

Figure 15 

Figure 16 

Botany 135 

Only one family of the true ferns is represented, the Marratiaceae. 
Petrified material of this family which existed during the Mississippian, 
Pennsylvanian, and Permian Periods is usually assigned to the genus 
Psaronius. This genus greatly resembled the modern tree ferns and was 
one of the dominant elements of the Pennsylvanian and Permian 
Periods. The trunk of this plant was an unbranched, gradually tapering 
column, and was covered by a dense mantle of adventitious roots. The 
mantle of roots was especially thick in the lower trunk region, resulting 
in the formation of a heavily buttressed base. The crown of the plant was 
composed of leaves which were few in number, but of great size, some 
as long as three meters. 

Two genera of sporangia exist which are attributed to Psaronius, 
with both genera apparently found on the same type of leaf. 
Cyathotrachus is a rather large synangium consisting of 7-9 sporangia, 
which are arranged around a central column for about half their length. 
Each sporangium is a distinct unit with a thin sporangial wall enclosing 
many spores. All of the sporangia are enclosed within a synangial cover, 
which apparently split at maturity to release the spores. Scolecopteris is 
the genus of a similar type of synangium which consists of six rather 
elongate sporangia joined only at their base. There is no synangial 
cover in Scolecopteris. 

Scolecopteris synangia are represented in considerable numbers in 
the present collection, suggesting a rather widespread distribution. When 
the synangia are sectioned parallel to the leaf surface, the sporangia 
appear in cross section (Figure 14), and the structure of the thin 
sporangial walls may be observed. Figure 15 shows a longitudinal section 
of Scolecopteris and illustrates the elongate nature of the sporangia and 
their attachment to the leaf. 

Another unusual feature of Psaronius is the thick mantle of roots 
which surround the stem. One specimen has been reported which had a 
stem two inches in diameter and a mantle of roots thirty inches thick. 
The broad mantle of roots in this genus is composed of two zones, an 
inner zone of small roots imbedded in a cellular matrix, and an outer 
zone of larger, free roots. Each root has a small, star-shaped protostele 
and a broad aerenchymatous cortex. The cortex is bounded by a region 
of thick-walled hypodermal cells which form the outer portion of the 
root. The presence of the aerenchymatous cortex suggests the possibility 
of a swampy habitat for the Psaronius trees. 

One of the coal balls in the present collection was found to be 
composed entirely of Psaronius roots. The roots from this specimen 
illustrate the small protostele composed of xylem tracheids (Figure 16). 
The region of hypodermal cells is also clearly evident in this section. 
Several examples of isolated Psaronius roots were found in other coal 
balls which illustrate the aerenchymatous cortex. 


Benninghoff (2) published a report of a coal ball flora from a mine 
near Petersburg, Indiana, in which he described thirteen genera of fossil 

136 Indiana Academy of Science 

plants. The genera described by Benninghoff, together with genera 
reported in selected Illinois studies, are compared to the genera in the 
present collection in Table 2. Benninghoff described three genera of 
synangia, Asterotheca, Myriotheca, and Phytocarpus, which are not rep- 
resented in the present material nor in any of the Illinois studies. All 
other genera described by Benninghoff are included in the present study. 

The degree of similarity existing between the assemblage of fossil 
plants of Indiana as represented in the present collection, and the 
assemblage of fossil plants of Illinois is summarized in Table 2. 

Darrah (3) has said that the plant assemblages found in nearly all 
American coal ball floras are basically similar. The differences lie 
mainly in the change of dominant genera from flora to flora, or the 
occasional addition or deletion of a few genera. The present study tends 
to support Darrah's statement. Considerable similarity exists between 
the plant assemblage found in Illinois and the plant assemblage of 
Indiana. The present study provides the basis for a more detailed 

table 2. Genera represented in Indiana and Illinois coal ball studies. 

o f> 

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Sphenophyllum Stem 



Sphenophyllum Root 







Botany 137 

TABLE 2. (continued) 



























, — 



















C- J 






Medullosa Stem 













: * 







Cordaites Stem 


Cordaites Leaf * 

Amyelon * 



















Psaronius Root 


* : 

|J * 


Psaronius Stem 







Cyathotrachus * 



Notoschizaea * 







The striking similarities which appear between the Illinois studies 
and the present Indiana material suggest that a relatively uniform flora 
existed over portions of Illinois and Indiana during the Pennsylvanian 

138 Indiana Academy of Science 

Literature Cited 

1. Andrews, H. N. 1961. Studies in paleobotany. John Wiley and Sons, New 

2. Bbnninghoff, W. S. 1943. Preliminary report of a coal ball flora from 
Indiana. Proc. Indiana Acad. Sci. 52:62-6S. 

3. Darrah, W. C. 1947. Studies in American coal balls. Amer. J. Sci. 239:33- 

Algal Records for Three Indiana Sewage Stabilization Ponds 

C. Mervin Palmer 

Advanced Waste Treatment Research Laboratory, FWPCA, 
Cincinnati, Ohio 


Algal identifications have been recorded from 376 samples collected from 
three Indiana sewage stabilization ponds during a period from May 1962 to 
August 196S. Although certain genera were found frequently in all three ponds, 
each pond had a distinctive algal flora. Green algae were invariably the most 
abundant of the algal groups present. However, flagellates were also prominent. 
Of a total of 64 genera of the most significant and abundant algae there were 
29 green algae, 19 flagellates, 10 blue-green algae, and 6 diatoms. Some genera 
were limited to the summr season, while others were most prominent in 
spring and fall or in the winter. The pollution-tolerant algae Euglena and 
Nitzschia were abundant and persistent in all three ponds. 

Three sewage stabilization ponds in southeastern Indiana were 
among several throughout the United States selected for biological 
studies, with particular emphasis on the algal flora. The Indiana ponds 
serve the communities of Napoleon and Sunman in Ripley County and 
St. Paul in Decatur County. These ponds range from 2.9 to 7.9 acres and 
are from 3 to 4 feet deep. The respective populations served are approxi- 
mately 300 to 800 with a pond area of about 0.01 acre per person. The 
biochemical oxygen demand (BOD) at the intakes has been ± 500 ppm 
and for the effluents less than 50 ppm. The Ripley County ponds were 
put into use in 1961 and the one in Decatur County three years later. 

Information on one or more of these ponds has been reported by 
Kuwahara (2), Palmer (5), Safferman (6), and Safferman and Morris 
(7, 8). The first virus infecting a blue-green alga was isolated from one 
of the ponds in Ripley County (6). 

Algal identifications were recorded from samples from the two 
Ripley County ponds on 43 different days during all seasons between 
May 1962 and August 1968. For the St. Paul pond, summer samples only 
have been available, these being obtained in 1966 and 1968, with collec- 
tions on eight different dates. Four samples per pond were obtained on 
each date of collection making a total of 376 samples. Each set of four 
samples represented the following sites: A — Water, near influent; B — 
Sludge or floating mat, near influent; C — Water, near effluent; D — Sludge 
or floating mat, near effluent. Identification of the algae in each sample 
was to genus and the relative abundance of each was recorded. 

Records of the microscopic analyses of samples for three dates for 
Napoleon and Sunman and two dates for St. Paul are given in Table 1. 
The data were selected to permit comparison of the algal flora in the 
three ponds and for each pond at different times of the year. Relative 
abundance of all genera is indicated using the figures from 1 through 5. 



Indiana Academy of Science 






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142 Indiana Academy of Science 

Sludge or floating mat samples (B and D) often contained a larger 
number of algal genera than did the water samples from the same 
locations (A and B). 

The number of genera of four algal groups recorded for each of the 
three ponds is listed in Table 2. In all three ponds the green and flagellate 
genera outnumbered those of blue-green algae and diatoms. 

table 2. Number of Genera by Groups in Each Indiana Pond. 

Algal Group 



St. Paul 

Green Algae 
Blue-Green Algae 
Flagellate Algae 















The highest number of genera recorded for any pond for one day 
was 21. However, the averages differed for the three ponds, being 11 for 
Sunman, 14 for Napoleon, and 18 for St. Paul. The higher the number 
of genera, the more likely that wastes in the influent have been stabilized 
as algal ingredients. A number of industrial wastes, including those 
from dairies and food canneries are generally not as readily assimilated 
by algae as is typical household sewage. Variations in loading and in 
environmental factors can also affect the algal flora. A total of 83 algal 
genera was recorded. Table 3 lists the most abundant and significant 
of these, together with the number of sampling dates they were recorded 
for each pond. Among the 64 genera listed are 24 non-filamentous green 
algae, 19 pigmented flagellates, 10 blue-green algae, 6 diatoms, and 5 
filamentous green algae. 

Ankistrodesmus, Chlamydomonas, Chlorella, Euglena, Nitzschia, and 
Oscillatoria were persistent in all three ponds. All are among the top 12 
algal genera most tolerant of pollution (4). Also fairly persistent in all 
three ponds were Anacystis, Oocystis, Pandorina, Schizothrix, and 
Scenedesmus. All but Schizothrix are among the top 35 pollution-tolerant 
genera (4). This genus would also be within the grouping if it were 
credited with the records of its synonyms as recognized by Drouet (1). 

Anabaena, Chodatella, Chroomonas, Closteriopsis, Closterinm, 
Cryptomonas, Spirulina, and Zoochlorella were present much more often 
in the Napoleon pond than in the other two ponds. Arthrospira and par- 
ticularly Pyrobotrys are the only significant ones that were distinctive at 
Sunman. At St. Paul the diatom Cyclotella was the only distinctive genus 
that appeared more than once. The interesting forms Achnanthes, 
Stigeoclonium, and the blue-green alga Johannesbaptistia were reported 
once for that pond. Navicula and other diatoms (Achnanthes, 
Gomphonema, Nitzschia) were fairly consistent in the St. Paul flora. 



TABLE 3. Relative Persistence of Algae in Indiana Sewage Ponds. 
(No. Dates Recorded per Alga) 




St. Paul 
Equivalent 1 " 










































































































































































Indiana Academy of Science 

table 3. (continued) 




St. Paul 





















































*No. of Dates recorded X5 

Napoleon and Sunman ponds were sampled during all months of 
the year, making possible a study of seasonal distribution of the algae. 
While many algae can be recorded as abundant during the summer, a 
much more restricted flora is found during the winter. Of the flagellates, 
Phacus and Chroomonas were most abundant during the spring, 
Pandorina during the summer, and Cryptomonas during the autumn and 
winter. Chalamydomonas was abundant during all seasons and Euglena, 
while present in all seasons, was less abundant during the winter. The 
diatoms Nitzschia and Navicula were present in all seasons. The former 
was much more abundant and appeared in greatest numbers in spring 
and fall. 

Most of the more common green algae tended to be primarily sum- 
mer forms. These included Actinastrum, Golenkinia, Kirchneriella, 
Oocystis, and Planktosphaeria. During the test period, Selenastrum was 
absent in the fall and most abundant in winter. Scenedesmus and 
Ankistrodesmus remained high in numbers during fall as well as sum- 
mer. Oscillatoria and Schizothrix, the two most prominent blue-green 
algae, were recorded during all seasons and were abundant from late 
spring until at least mid-autumn. 

Some algae less common in sewage ponds, but present in one or more 
of the Indiana ponds, included the diatoms Cyclotella and Hantzschia, 

Botany 145 

the blue-green algae Arthrospira, Johannesbaptistia, and Raphidiopsis, 
the green algae Actinastrum, Chodatella, Closterium, Dictyosphaerium, 
Selenastrwm, Spirogyra, and Ulothrix, and the flagellate algae 
Chlorogonium, Chroomonas, Cryptomonas, Eudorina, Massartia, 
Phacotus, Pteromonas, and Pyrobotrys. 

On each collection date two samples were taken from the influent 
area and two from the effluent area of the pond. Microscopic analyses of 
these samples indicated very little difference between the algal flora of 
the influent and effluent areas. This could be the result of rapid mixing 
of the water throughout the pond. It could be the result of rapid oxida- 
tion and stabilization of the sewage, including that in the influent area. 

At Napoleon and St. Paul the more abundant algae were generally 
present in each set of samples collected. This pattern indicates the con- 
stant quality and quantity of sewage entering the ponds. At Sunman, 
Scenedesmus was rare to absent after 1963 and Ankistrodesmus was 
rare to absent after 1965. Chlorella became more abundant in 1965 and 
the related genus Nannochloris appeared in large numbers in May 1966. 
Both persisted in abundance afterwards. Such a change in the flora has 
been considered elsewhere to be due to the development of partial 
anaerobic conditions (3). 

The algal flora of three sewage stabilization ponds in Indiana was 
composed of an average of 11 to 18 genera of algae. The most abundant 
and persistent forms were all ones previously classified as highly tolerant 
to organic enrichment. In addition to the many green algae a large 
number of flagellate algae were present, some of these being seasonal. 
Blue-green algae and diatoms, with the exception of Nitzschia, were not 

Collection of many of the samples used in this study was made by 
Dr. Robert Safferman of the Advanced Waste Treatment Research 
Laboratory; appreciation is expressed to him for his cooperation. 

Literature Cited 

1. Drouet, F. 1968. Revision of the classification of the Oscillatoriaceae. 
Monogr. 13. Acad. Natural Sciences Philadelphia (Pa.) 

2. Kuwahara, R. 1963. Report on studies in U.S.A. Ill Development of the 
lagoon in U.S.A. J. Water and Waste, Tokyo, Japan 5(4) :261-265. (In 

3. deNoyeles, P., Jr. 1967. Factors affecting phytoplankton distribution in a 
double-cell sewage lagoon. J. Phycology 3:174-181. 

4. Palmer, C. M. 1963. The effect of pollution on river algae. Ann. New 
York Acad. Sci. 108(2) :389-395. 

5. Palmer, C. M. 1967. Nutrient assimilation by algae in waste stabilization 
ponds. Proc. Indiana Acad. Sci. 76:204-209. 

6. Safferman, R. 1968. Virus disease in blue-green algae, p. 429-439. In: 
Algae, Man and Environment. Ed. D. F. Jackson, Syracuse University 
Press. Syracuse, N. Y. 

7. Safferman, R. s and M. E. Morris. 1963. Algal virus: Isolation. Science 
140(3567) :679-680. 

8. Safferman, R., and M. E. Morris. 1967. Observations on the occurrence, 

distribution, and seasonal incidence of blue-green algal viruses. Appl. 
Microbiol. 15(5) :1219-1222. 

Environmental Regulation of Experimental Leaflet Abscission 1 

D. James Morre', Bruce Rau, Richard Vieira, Tim Stanceu and Terry 
Dion, Department of Botany and Plant Pathology, Purdue University, 

Lafayette, Indiana. 

Leaf abscission is a terminal developmental sequence initiated by 
the onset of senescence (excision and deblading in experiments with 
explants) and terminated by separation of the leaf from the main axis 
of the stem (3, 4, 8, 9, 13). Studies of abscission are greatly facilitated 





20 - 

^ "^™ 

■*"' A 

Time to 50% Separation B^ / 

(23+0.5°)+ SD / / 

/ / 

— i • .1. • £Ol iflh lAAi irt lin / / 

Light W w l<iUl \d. HK / / 

Dark ■ ■ 100+ 7 HR / / 

/ / 

1 w 

1 / 

1 / 

dark/ / 

/ / 

/ A 

1 W 


/ / 

t / 

/ / 

/ / 

/ / 

/ W 

/ ^ 

~" 1 / 

1 / 

/ / 


/ / 

m,^-^L ^ 1,1,1. 

40 80 120 



Figure 1. Transition profiles of separation layer formation at 23° com- 
paring explants in light and darkness. Explants were prepared from plants 
grown in the greenhouse at high light and long photoperiod. 

1 Purdue University AES Journal Paper Number 3551. Supported in part 
by a contract with the United States Army Biological Laboratories, Fred- 
erick, Maryland and a NSF Undergraduate Research Participation Grant. 




by use of explants (excised abscission zones and adjacent tissues) which 
respond uniformly and rapidly but exhibit widely varying rates of 
abscission depending upon treatment conditions. 

The behavior of an explant population is described by a transition 
profile, a simple sigmoid defined by two parameters: the position of the 
midpoint (time to 50% separation) and the steepness of the slope at 
this point (or by an equivalent parameter, the transition width). The 
time at which abscission reaches the midpoint of the transition is a 
measure of the rapidity at which separation layers are formed. The 
transition width is a measure of the heterogeneity of the population. A 
very broad transition is indicative of a heterogeneous population whereas 
a sharp transition is indicative of a synchronous population. 

With bean, natural abscission of primary unifoliate leaves occurs 
between 30 and 40 days after planting with plants grown under green- 
house conditions (10). Explants of primary leaves reach 50% separation 
at about 100 hours following the procedures of Leopold and coworkers 
(5, 10, 13). It is desirable to utilize explants with very sharp transition 
profiles (synchronous response). For many types of abscission studies, 
it would be equally desirable to utilize explants with 50% separation 
within a day after cutting rather than the usual 100 hours. The rapid 
abscission rate would permit initiation and termination of kinetic studies 
during a single working day instead of over a period of several days or 

table 1. Abscission of Bean Explants Harvested From Plants Grown 
Under Varying Conditions of Temperature, Light Intensity and Photo- 
period. Abscission of explants was determined at three temperatures 
under continuous, low-intensity light (100 ft-c) or da/rkness. 

Conditions for Growing 
Plants Prior to 
Harvesting Explants 
Temper- Light Photo- 

ature Intensity period 

Time to 50% Separation of Ex- 
plants (Hr±SD) at Conditions 
29° 24° 22° 22° 

Light Light Light Dark 




4,800 ft-c 

8 Hr 





12 Hr 





16 Hr 







1,000 ft-c 

8 Hr 





12 Hr 





16 Hr 





20 Hr 








1,000 ft-c 

8 Hr 





12 Hr 





16 Hr 





20 Hr 






Indiana Academy of Science 

Materials and Methods 

Bean plants (Phaseolus vulgaris L., var Red Kidney) were grown 
under greenhouse or controlled environmental conditions. Unless specified 
otherwise, explants were cut from unifoliate leaves of 10 to 20 day old 
plants grown in metal flats containing potting soil overlayed with sand. 
The flats were filled to a depth of 5 cm with the soil followed by a layer 
of seeds which were then covered to a depth of about 1 cm with sand. 
Greenhouse-grown plants were supplemented with low intensity fluores- 
cent and incandescent light to provide a 16 hr. photoperiod. 

The 1 to 2 cm long debladed explants were cut to include at least 
5 mm of tissue on either side of the distal abscission zone at the pulvinus- 







a: 80 




- 1 

■ I 

' i 


1 1 1 

1 1 


22 26 30 34 



Figure 2. Acceleration of explant abscission in continuous light as a 
function of temperature. Explants prepared from greenhouse-grown plants. 
Bars show standard deviations. 



petiole juncture (10). Unless specified otherwise, approximately 20 
explants were inserted vertically (usually with the petiole end down) 
to a depth of 4 mm in 1% agar in 1 X 5 cm petri dishes. To facilitate 
inserting the pulvinus end of the explants into agar, an agar concentra- 
tion of 0.7% was used. Rate of separation of explants was independent 
of agar concentration over the range 0.7 to 2%. Results are an average 
of at least 3 experiments. 

The time of separation layer formation was estimated from the 
time when application of pressure to the pulvinus readily brought about 
separation. Details of individual experiments are given in the text. 


Transition profiles for explants prepared from greenhouse-grown 
bean plants (Fig. 1) showed a broad transition width when assays were 
conducted at 23°. In low intensity (100 ft-c) light, explants began to ab- 
scise between 60 and 70 hours after cutting but did not reach 100% abscis- 
sion until after 170 hours (50% separation at 120 hours). With green- 
house-grown plants (high light intensity and long photoperiod), 
abscission was accelerated when explant assays were conducted in 






20 40 60 80 100 

Figure 3. Transition profiles of separation layer formation comparing ex- 
plant separation in continuous light at 28 and 22.5°. 

150 Indiana Academy of Science 

darkness (50% separation at 100 hours). Except for explants prepared 
from plants grown under high light intensity and long photoperiod, ab- 
scission was generally faster at 22° when explants were incubated in 
the light (Table 1). 

Abscission is extremely sensitive to temperature (Figure 2; Table 1). 
Abscission was not detected at temperatures below 16° or above 36°. 
Increasing the temperature from 22° to 26° markedly accelerated ab- 
scission (50% separation at 120 hours at 22° vs 50% separation at 24 
hours at 26°). The temperature response curve for experimental leaflet 
abscission exhibited a broad optimum between 26 and 30°, with a decline 
in rate at 32°. Between 32 and 36°, abscission was further delayed and 
many of the explants did not abscise. Increasing temperatures decreased 
the midpoint of the transition profiles (Fig. 2) and at the same time 
increased the steepness of the slope (decreasing the transition width) 
as shown in Figure 3 for greenhouse-grown plants (compare 28°, Fig. 3 
with 22.5°, Fig. 3 and 23°, Fig. 1). The temperature at which the plants 
were grown prior to harvesting the explants had much less of an 
effect on abscission rate than the temperature during the explant bio- 
assay (Table 1). 

Effects of light on abscission were less pronounced than the effects 
of temperature. The most marked effect was the qualitative response 
to light vs. dark (Fig. 1, Table 1) which varied according to the light 
intensity and photoperiod at which the plants were grown. With all 

table 2. Abscission of Bean Explants as Influenced by Temperature, 
Light Intensity and Photoperiod. Plants grown in soil in the greenhouse 

(16 hr photoperiod) . 


Time to 


Light Intensity 


50% Separation 










































Botany 151 

plants (except those grown at high light intensity and long photoperiod), 
incubating explants in low light (100 ft-c) markedly accelerated abscis- 
sion. Increasing the light intensity from the range 700 to 1,000 ft-c to 
4,500 to 4,800 ft-c resulted in no further change in the rate of separation 
layer formation (Table 2). When explant assays were conducted at 22.5°, 
separation rate was inversely related to photoperiod with photoperiods 
of 8, 12, 16 and 20 hours. At 30°, the response to photoperiod was ir- 

table 3. Acceleration of Abscission in the Light by Inverting the 


Time to 50% Separation 
Portion of the Explant (Hr ± SD) 

Placed in Agar Dark Continuous Light 

Petiole end 109 ±9 77 ±1 

Pulvinus end 109±9 46±5 

The photoperiod at which the plants were grown affected separation 
layer formation in explants assayed under continuous light but differ- 
ences were not marked. Explants prepared from plants grown under low 
light intensity tended to abscise more rapidly in the light than those 
from plants grown at high light intensity. This response was independent 
of photoperiod and assay temperature (Table 1). Most rapid abscission 
was obtained from bean plants grown at 22.5°, 700 to 1000 ft-c and an 
8 hr photoperiod. All differences due to growth conditions of the plants 
tended to be minimized when explant assays were conducted at a near 
optimum temperature of 29° as compared with a suboptimal temperature 
of 22° (Table 1). Explants from plants grown in complete darkness 
(completely etiolated condition) did not absice when subsequently incu- 
bated either in light or darkness. 

Time to 50% separation decreased with increasing age of the plant 
particularly when assays were conducted at 23°. However, at optimum 
temperature, age of the plant did not appear to be a significant variable 
(Fig. 4). The most marked response to age was from plants grown at 
22.5°, a 12-hour photoperiod and a light intensity of 700 to 1000 ft-c. 

table 4. Influence of Stipules on the Rate of Bean Explant Abscission. 

Number of Stipules Number of Explants Time to 50% 

Per Explant Tested Separation, Hrs 

200 32 

1 100 27 

2 80 27 


Indiana Academy of Science 

Under these growing: conditions, the midpoint of abscission was shortened 
by about 5 hours comparing 12 day old and 20 day old plants. 

With assays conducted in light, acceleration of abscission was 
achieved by inverting the explants (incubation with the pulvinar end 
in agar). Inverting the explant was without effect in the dark (Table 3). 
Ethylene when applied to explants at ambient pressure and temperature 
provided a further stimulation of abscission rate. The optimum con- 
centration was near 1 ppm (Fig. 5). In some species, the stipules are 
a natural source of ethylene. With our explants, stipulate and exstipulate 
explants behaved similarly (Table 4) although abscission rate was re- 
duced slightly by removing both stipules. A shortening of the time to 
50% separation equivalent to that with ethylene was obtained by 
supplying galactose to the agar (Fig. 6). These experiments were con- 







io 10 


^ □ 


□ -D 22.5°C, 12 Hr Photo- 
period 700-1000 ft-c 

• • Average of all Conditions 







Figure 4. Rate of separation layer formation of explants excised from 
bean plants of varying- ages. All conditions are those listed in Table I. 
Assays were conducted in continuous light at 28 to 29°. 



table 5. Influence of Concentration and Placement of Sucrose and Glu- 
cose on Bean Explant Abscission at 29°. 
Sugars were supplied in 1 % Agar. 

Portion of Explant 
Placed in Agar 



Time to 50% 
Separation (Hr) 



— ■ 


















. i 





q: 20 

1 — A 



W ■ - ■ - 













1 1 

1 1 1 


Figure 5. Separation layer formation as a function of ethylene concentra- 
tion supplied to the gas phase of sealed flasks containing' the explants. Ex- 
plants were obtained from greenhouse-grown plants. Assays were con- 
ducted at 28 to 29° in continuous light with the explants inverted (pulvinus 
end down) in 0.7% agar. 


Indiana Academy of Science 

ducted with explants in the inverted position (distal end down). The 
optimum galactose concentration was 2% and explants treated with 20% 
galactose did not abscise. Under similar conditions of treatment, sucrose 
and glucose also stimulated abscission but not to the extent exhibited by 
galactose (Table 5). 

Transition profiles for inverted explants treated with ethylene or 
galactose (Fig. 7) at 28° show both a steepening of the slope and a 
marked shortening of the midpoint. When supplied together at optimal 
concentrations, ethylene and galactose were additive in their effects on 
rate of separation layer formation (Fig. 7). The studies with ethylene 
were conducted in sealed flasks with inverted explants (pulvinar end 





o 20 




h- 10 


1 1 1 1 1 

2 4 6 8 



Figure (!. Separation layer formation as a function of galactose concen- 
tration in 0.7% agar. Explants were obtained from greenhouse-grown 
plants and assays were conducted at 28 or 29° in continuous light with the 
explants inverted (pulvinus end down). 



table 6. Abscission of Bean Explants at Two Temperatures Comparing 
Plants Grown Under Widely Varying Conditions. 

Growing Conditions for Plants Prior to Harvesting Explants 

Time to 50% 

Separation of 


Temperature, Chrono- Explants 

Intensity 1 

°C logical (Hr)2 



Day Night Age, Days 22.5°C 30°C 

Silicate Soil 



32 24 18 120 60 



32 24 10 96 24 


20 7 24 120 48 

i 12 Hour Photoperiod. 

212 Hour Photoperiod, 700-1,000 ft-c. 

^Krum (Silibrico Corporation, Chicago, Illinois). 




/■ p 

^ 80 

J / 

r / 


/ $ 

/ i 

b ~« 


G /E / G 

< 60 

/ / 


/ / 


/ / 


1 t 

a 40 


/ P 


J °/ 

T / 




r / 




j \ \ \ 

8 16 24 32 40 

Figure 7. Transition profiles of separation layer formation comparing- 1 
ppm ethylene in the gas phase (E), 2.5% galacose in the agar phase (G) 
and 1 ppm ethylene plus 2.5% galactose (E + G). Explants were obtained 
from greenhouse-grown plants and assays were conducted at 28 to 29° in 
continuous light with the explants inverted (pulvinus end down). 


Indiana Academy of Science 

4 8 12 16 20 


Figure 8. Transition profiles for explants incubated in the presence of 
1 ppm ethylene plus 2.5% galactose as described for Figure 7 comparing 

plants grown under greenhouse (■— — ■) and controlled environment 

(• • — 22.5°, 700 to 1000 ft-c and 8 hr photoperiod; o — o 

— 30°, 700 to 1000 ft-c and 12 hr photoperiod) conditions. 

Botany 157 

More detailed transition profiles of rapidly abscising explants (Fig. 
8) show a transition width of approximately 2 hours and a midpoint at 
12 hours when explants were prepared from plants grown at constant 
temperature (22.5°), low light intensity (700 to 1000 ft-c) and short 
(8 hr) photoperiod. With explants from greenhouse-grown plants (vari- 
able temperature, long photoperiod, high light intensity), the midpoint 
was at 14 hours (2 hours later). When compared under less favorable 
conditions (in the absence of inducers, upright position and 22°), their 
midpoints differed by as much as 70 hrs (50 hrs to 50% separation from 
explants from plants grown at 22.5°, low light and 8 hr photoperiod vs. 
120 hr for explants from greenhouse-grown plants, Table 1, Fig. 1). The 
equalizing effect of optimum conditions for the explant assay illustrated 
in Figure 8 was evident throughout the study. In general, differences due 
to growing conditions of the plants were minimized as conditions of the 
explant assay were optimized (Table 1). Also minimized as optimum 
conditions of explant assay were approached were differences among 
individual members of each population. The net result was a steepening 
of the transition profile (Fig. 8). 


External factors such as climatic conditions have long been known 
to influence abscission of leaves from intact plants (3, 4). Low carbo- 
hydrate levels favor rapid abscission. Moderate nitrogen, low water sup- 
ply, high light intensity and conditions of carbohydrate accumulation 
reduce or retard abscission (13). Light also may have an effect through 
photoperiod. Olmsted (11) observed that constant photoperiods, re- 
gardless of length, resulted in delayed abscission and senescence. 

The effect of environment on experimental leaflet abscission (studies 
utilizing explants) has not been studied extensively. Previously, Yama- 
guchi (reviewed by Addicott (3)) found that the formation of the 
abscission layer in bean explants exhibited a temperature response 
curve with a maximum between 25 and 30° and the effects of light and 
C0 2 on explant abscission were examined by Biggs and Leopold (5). 

Being a physiological process, abscission is extremely sensitive to 
temperature particularly over the range 16 to 26°. Increasing the tem- 
perature from 22° to 24°, for example, may shorten the time of 50% 
separation of explants by nearly 50%. Between 26° and 30°, temperature 
has little effect on abscission rate with an optimum near 29°. In earlier 
abscission experiments involving explants at about 23°, it is probable that 
much of the variation was due to variations in temperature. The tem- 
peratures at which the plants were grown prior to harvesting the ex- 
plants has much less of an effect on abscission rate than the temperature 
during the explant bioassay. 

The response of explants to light is more variable and less pro- 
nounced than that due to temperature. With plants grown under low light 
intensity (conditions that tend to minimize accumulation of carbohydrate 
reserves), abscission of explants occurs more rapidly in light than in 
darkness. However, when plants are grown in high light intensity 

158 Indiana Academy of Science 

(conditions favoring accumulation of carbohydrate reserves) just the 
opposite is true, i.e. explants abscise more rapidly in the dark. 

This light-induced delay of abscission with plants grown at high 
light intensity was investigated by Biggs and Leopold (3). They found 
that plants in a C0 2 -free atmosphere under 300 ft-c of light abscised at 
the same time as those kept in the dark with or without C0 2 , while those 
kept in ambient air were delayed by about 40 hours. They concluded 
that the effect of light was due to photosynthesis since in their studies 
the effect of light was eliminated by added sucrose. 

In our studies, conducted with greenhouse-grown plants, sucrose and 
glucose had slight stimulatory effects on abscission and any regular 
correlation among light, C0 2 , photosynthesis, carbohydrate accumulation 
and abscission seemed doubtful. A further complication arose from the 
unexplained observations that explants abscised faster with the pulvinus 
end in agar (petiole end up) than when the petiole end was in agar 
(pulvinus end up) but only in light. In darkness, the explant apparently 
did not sense which end was up. 

A possible indirect involvement of photosynthesis in leaf abscission 
would be through CO- removal as it effects the response of the plant 
to ethylene. Ethylene is a natural promoter of abscission (1, 2, 6, 12) 
and carbon dioxide (CO?) is a competitive inhibitor of its action. Thus, 
a simple explanation of the effects of light and temperature on experi- 
mental leaflet abscission might be found in a consideration of the com- 
bined rates of photosynthesis and respiration as they affect C0 2 levels. 
A buildup of CO2 within the petri plates containing the explants might 
interfere with the normal acceleration of separation layer formation by 
endogenous ethylene, particularly at high temperatures. Under condi- 
tions where respiratory substrates are non-limiting, photosynthesis 
would be expected to substantially reduce the C0 2 content of the gas 
phase of explants incubated in the light with a corresponding accelerated 
rate of separation layer formation. Both ethylene and galactose appear 
to accelerate abscission through their specific action in the induction of 
a pectinase catalyzing the hydrolysis of the cementing substances be- 
tween adjacent cell layers of the abscission zone (10). 

With cotton petioles, Hall and Liverman (7) found that abscission 
was accelerated in proportion to increased light intensity up to 2,500 
ft-c but light intensities from 6,000 to 8,000 ft-c delayed abscission sig- 
nificantly. Acceleration was obtained with red, far-red, fluorescent and 
ultraviolet light but the effects of light intensity on abscission have not 
been studied extensively. Similarly, in our studies, plants grown under 
high light intensity were delayed in their abscission response as com- 
pared to plants grown under low light intensity. 

Throughout these studies, light and temperature have been most 
effective in regulating explant abscission during incubation of the ex- 
plants. Abscission of explants is only little influenced by the conditions 
under which the plants are grown except for extremes of temperature, 
nutrition and photoperiod (Table 6). Here, conditions favoring slow 

Botany 159 

growth of the plants tend to be associated with a reduced rate of separa- 
tion layer formation. 


Separation layer formation in explants of red kidney bean was 
markedly accelerated by increasing temperature between 16 and 26° C 
with an optimum between 28 and 30°. Light effects were largely qualita- 
tive. Quantitative changes in light intensity and photoperiod had little 
effect on the abscission response when explant assays were conducted at 
optimum temperatures. Light and temperature effects on explant abscis- 
sion were largely restricted to the explants themselves with minor 
modulations due to the environmental history of the plants from which 
the explants were prepared. 

By combining near optimal growing conditions of the plants prior 
to preparation of the explants, optimum temperature and continuous 
light during the explant assay in conjunction with two inducers of pec- 
tinase (galactose and ethylene) supplied under optimal conditions, we 
shortened the time of experimental leaflet abscission from 100 hours to 
12 hours. The development of an experimental system in which the time 
from excision of the explant to separation layer formation could be con- 
fined to a period of 8 hours or less would be of interest not only from 
the standpoint of practical defoliation but to expedite certain kinetic 
analyses of abscission events as well. 

Literature Cited 

1. Abeles, F. B. 19C7. Mechanism of action of abscission accelerators. 
Physiol. Plant. 20:442-454. 

2. Ajbeles, F. B. and B. Rubinstein. 1964. Regulation of ethylene evolu- 
tion and leaf abscission by auxin. Plant Physiol. 39:963-969. 

3. Addicott, F. T. 1964. Physiology of abscission. Encyclopedia of Plant 
Physiol. XV/2:1094-1126. 

4. Addicott, F. T. and R. S. Lynch. 1955. Physiology of abscission. Ann. 
Rev. Plant Physiol. 6:211-238. 

5. Biggs, R. H. and A. C. Leopold. 1957. Factors influencing abscission. 
Plant Physiol. 32:626-632. 

6. Hall, W. C. 19 5S. Physiology and biochemistry of abscission in the cotton 
plant. Texas Agricultural Experiment Station Miscellaneous Pub. 

7. Hall, W. C. and J. L. Liverman. 1956. Effect of radiation and growth 
regulators on leaf abscission in seedling cotton and bean. Plant 
Physiol. 31:471-476. 

8. Holm, R. E. and F. B. Abeles. 1967. Abscission: The role of RNA syn- 
thesis. Plant Physiol. 42:1094-1102. 

9. Morre', D. J. 1968. Leaf abscission: The chronology and control of a 
terminal developmental sequence. Proc. Western Agricultural Experi- 
ment Station Collaborator's Conference. 

10. Morre', D. J. 1968. Cell wall dissolution and enzyme secretion during 
leaf abscission. Plant Physiol. 43:1545-1559. 

160 Indiana Academy of Science 

11. Olmsted, C. E. 1951. Experiments on photoperiodism, dormancy and 
leaf age and abscission in sugar maple. Botan. Gaz. 112:365-393. 

12. Rubinstein, B. and F. B. Abeles. 1961. Relationship between ethylene 
evolution and leaf abscission. Botan. Gaz. 126:255-257. 

13. Rubinstein, B. and A. C. Leopold. 1964. The nature of leaf abscission. 
Quart. Rev. Biol. 39:356-372. 


Chairman : Ralph Jersild, Indiana University Medical Center 
Edward J. Hinsman, Purdue University, was elected chairman for 1969 


The Fine Structure of the Ventral Horn Neuron in the Calf Spinal Cord. 

E. J. Hinsman and Kathleen Moe, Purdue University. — Tissues col- 
lected from the ventral horn of the lumbar spinal cord of gluteraldehyde 
perfused calves were examined. Study was directed toward the larger 
neurons and dendritic interrelationships. The large neurons appeared 
similar in appearance to large motor neurons which have been described 
from other species. In areas of dendritic apposition interdendritic mem- 
brane modifications were found which appeared similar to desmosomes. 
These consisted of the thickening of apposed dendritic membranes, an 
increased intercellular space and electron dense cytoplasmic condensa- 
tions. Implications of their presence will be discussed. 

An Intranuclear Structure in Neurons of Human Cerebral Cortex. Itaru 
Watanabe, Sheila Donahue, and Wolfgang Zeman, Indiana Univer- 
sity Medical Center. — Examination in the electron microscope of cerebral 
cortical biopsy material from seven children with various intractable dis- 
orders of the central nervous system revealed occasional peculiar struc- 
tures in the nuclei of nerve cells. This structure is made up of filaments 
approximately 100 A in diameter. There is variation in the appearance 
of this structure which is probably due to the angle of the section. The 
filaments are seen in bundles, and forming a lattice, also as round ele- 
ments in cross section between parallel filaments bearing resemblance 
to a corncob. The whole structure is of unknown length, the longest ob- 
served measuring more than 5/a. The greatest width observed is 
1.2/4, but more often they measure only 20 to 120 m/x. They were seen in 
both sexes, ages from 2 to 9 years, in six different disorders, but 
always in nerve cells. Such structures have been observed in various 
vertebrates. Their significance is unknown. 

Chemical Composition of Membrane Fractions Isolated from Rat Liver in 
Relation to Membrane Differentiation During Secretion. Wayne Yung- 

hans and D. James Morr£, Purdue University. — The chemical composi- 
tions of Golgi apparatus, endoplasmic reticulum (ER), and plasma mem- 
brane were compared. The ratios of protein to lipid in Golgi apparatus 
and plasma membrane were similar but less than for ER. Phospholipids 
of the three membrane fractions were similar but the sterol content of 
Golgi apparatus was intermediate between that of ER and plasma mem- 
brane. Carbohydrate, RNA, and DNA were found in negligible amounts 
in the Golgi apparatus fraction indicating low contamination from 
glycogen, ribosomes, and nuclear material. 

Disc electrophoresis patterns of structural proteins from Golgi 
apparatus and ER were similar but different from those of plasma mem- 


162 Indiana Academy of Science 

hrane. Structural protein of plasma membrane separated into a character- 
istic two-band pattern with corresponding bands being present in both 
Golgi apparatus and ER. The structural protein pattern from mito- 
chondria showed several bands not present in the patterns from the 
other membrane fractions. These results together with electron micro- 
scopic evidence showed the Golgi apparatus to be a unique component 
of the endomembrane system. However, its chemical composition was 
intermediate between that of ER and plasma membrane and consistent 
with a functional role of the Golgi apparatus as a site of endomembrane 
differentiation during secretion. (Supported in part by grants from the 
NSF GB-7078 and NDEA Title IV 2599-82-11557.) 

An Electron Microscopic Study of Zinc Iodide-Osmium Staining of the 
Golgi Apparatus of Rat Intestinal Epithelial Cells. K. M. Mak and 

R. A. Jersild, Indiana University Medical Center. — A modification of the 
zinc iodide-osmium (ZIO) impregnation technique, used for the staining 
of synaptic vesicles at cholinergic junctions (Akert and Sandri, Brain 
Res., 7:286, 1968), has been applied to intestinal epithelial cells of the rat 
jejunum in an attempt to localize choline-containing phospholipids. 
Results showed that an electron dense reaction product was localized 
within the Golgi apparatus. In starved animals, one or two of the Golgi 
cisternae were ZIO-positive while associated vacuoles remained negative. 
In lecithin-fed animals, an increasing number of ZIO-positive cisternae 
were observed. In addition, reaction product was seen deposited around 
droplets, presumably of fat, accumulated in the Golgi vacuoles. Treat- 
ment of tissues in a solution containing either potassium iodide-Os0 4 
(KIO), zinc sulfate-Os04, or Os0 4 alone failed to elicit a reaction prod- 
uct within the Golgi apparatus. Thus, the reaction product is specific to 
ZIO mixture. The affinity of ZIO for the Golgi apparatus was inhibited 
by pretreatment of tissues with KIO, suggesting that KIO and ZIO were 
reacting with the same substrate. Extraction of tissue lipids with 
chloroform-methanol prevented the Golgi response to ZIO. It is sug- 
gested in this study that the Golgi apparatus is a site of accumulation 
of phospholipids. (Supported by PHS Research Grant AM 11721-01 
from National Institute of Arthritis and Metabolic Diseases.) 

The Ultrastructural Features of Intraoral Lichen Planus, Simplex. J. B. 

Whitten, Jr., Indiana University, Department of Oral Pathology, Indi- 
anapolis. — Lichen planus is a chronic, benign, dermatologic disease which 
is usually self-limiting. The skin lesions are elevated, scaly, biolaceous 
plaques which are puritic and tend to expand and coalesce forming 
larger lesions. The oral disease exists in several forms: namely (1) 
simple, (2) erosive, (3) bullous, and (4) hypertrophic. The lesions are 
elevated bluish white lines which cross forming beads. In other cases 
bluish white plaques of varying size occur without clinical identity. 

Biopsies of five patients with the simple form of the intraoral disease 
were examined with the light and electron microscopes. The tissue for 
light microscopy was fixed in 10% formalin, embedded in paraffin, sec- 
tioned at about 6/x and stained with hematoxglin and eosin. The remain- 
ing tissue for electron microscopy was fixed in 4% glutaraldehyde, post 

Cell Biology 163 

fixed in osmic tetraoxide, embedded in Epon 812, sectioned at 600 A with 
glass knives and Porter-Blum ultramicrotomes, mounted on uncoated 
200-mesh copper grids, and stained with lead citrate and uranyl acetate. 

Three phases of lichen planus were observed on ultrastructural 

Stage I — Particulate accumulations were found within some of the 
spinous cells. The particles were composed of five 
(usually) electron-dense bodies 30 A in diameter set in 
a much less electron dense amorphous material. Sur- 
rounding the dense bodies were clear zones 10-20 A in 

Stage II — The basal epithelial cell layers showed considerable 
intercellular edema with occasional interepithelial inflam- 
matory cells usually lymphocytes. The superficial lamina 
propria, beneath the anchoring fibers and lamina densa, 
was altered. The usual collagen fibers were replaced by 
an amorphous material about 3/u to 5/x in thickness. 

Stage III — The deep spinous and basal epithelial as well as the 
superficial lamina propria is degenerated and necrotic. 
In these areas aggregates of bacteria were often associ- 
ated with the most severe degeneration. This was present 
even though the surface epithelium was intact. 

Direct Studies of Nuclear Movements in Schizophyllum commune. 
Donald J. Niederpruem and Ralph A. Jersild, Indiana University Medi- 
cal Center. — Past investigations employed indirect genetic techniques to 
quantitate nuclear migration in Schizophyllum commune. The current 
study employed living hyphae of 5. commune and compared nuclear 
movements in homokaryotic mycelia, dikaryotic mycelium and an 
AxBmut homokaryon of this mushroom. Rates of nuclear movement were 
measured by phase contrast microscopy and the employment of an 
ocular micrometer, using hyphal apices and septa as reference points. 
Forward nuclear movements were observed in growing hyphal apices of 
homokaryotic mycelia and the dikaryon. Nuclear movements occurred 
within the range of hyphal growth and could account for the maintenance 
of centrally located nuclei. Opposed nuclear movements after mitosis 
greatly exceeded the rate of apical growth. Septum disruption and 
extremely rapid nuclear movements were recorded in an AxBmut 
homokaryon. Neither cytoplasmic streaming nor actively participating 
granules or filaments could account for any of these nuclear movements. 
Glutaraldehyde-fixed hyphae were examined by electron microscopy and 
revealed microtubular elements. 

Plasma Cell Antibody Against Bovine Serum Albumin in the Rabbit 
Appendix as Revealed by the Fluorescent Antibody Technique. John F. 
Schmedtje, Indiana University School of Medicine. — Evidence is accumu- 
lating that heterologous serum protein can be absorbed through the 
normal gut epithelium of adult mammals. In the present experiment, 

164 Indiana Academy of Science 

bovine serum albumin (BSA) was intubated into the appendiceal lumen of 
the adult living rabbit. Three weeks later, BSA was added to the food and 
water for one week. The appendix was removed, and portions were quick 
frozen and subsequently sectioned in a cryostat. Adjacent sections were 
stained with H and E. Purified BSA and goat anti-BSA conjugated with 
fluorescein isothiocyanate were used, according to the Coons technique, to 
identify any intracellular antibody against BSA. 

Antibody against BSA was present in plasma cells beneath the 
appendiceal luminal epithelium and beneath the outer epithelium of 
appendiceal crypts. Antibody positive cells were not present in control 
rabbits that had not received the intubation of BSA. 

The results support the hypothesis that the intubated BSA was 
absorbed and acted as a sensitizing agent. The results also support the 
hypothesis that absorption of the oral doses occurred through the gastro- 
intestinal tract, that these acted as challenge doses, and that this induced 
plasma cell antibody production in the appendix wall. 

Tumor Cell Mitotic Activity in Mice Treated with Antigenic Materials. 

William E. Stovall and Gordon L. Rosene, Ball State University. — ■ 
Current interest in tumor specific immune mechanisms has prompted this 
pilot study to investigate such mechanisms in Strong A mice afflicted 
with spontaneous mammary carcinoma. The majority of previous work, 
excluding clinical investigations, involved transplantable tumors. 

Five groups of animals were employed, consisting of one untreated 
control group and four receiving periodic injections of various antigenic 
materials. Two groups received rabbit serum. The remaining two were 
given tumor combined with Freunds adjuvant. Serum recipient groups 
received 24 injections during a 37-day period. Adjuvant recipients 
received 10 injections during the same period. Animals were sacrificed 
at 12:00 noon. Histological sections of the tumors were made and stained 
with H and E. Tumor cell mitotic indices were determined. 

Data significance was determined using the Student t test. Mean 
mitotic index values from the two rabbit sera recipient groups were 
significantly different (5% level) from the control group. Mean mitotic 
indices obtained from the two Freunds adjuvant groups were not sig- 
nificantly different from the control group. The absence of significance 
in these two groups was due to extreme data variability. 

It was anticipated that injected materials would evoke an auto- 
immune response in the recipient mouse. Literature reports suggest that 
spontaneous tumors are sometimes weakly antigenic. These procedures 
were intended to produce an enhancement of such a response. Significant 
decrease in tumor cell mitotic indices could indicate a decrease in tumor 
metabolic activity. 

Microimmunoelectrophoresis of Human Blood in Regard to the Study of 
the Gc System. Shirley Frances (Archibald) Smalley, Ball State 
University. — This study was conducted in order to determine if the Gc 
protein could be accurately and economically identified using a normal lab- 

Cell Biology 165 

oratory technique. If the test could be adapted to a routine laboratory pro- 
cedure, it would be possible to use the Gc determination in identifying 
the questionable parentage in a legal paternity suit. Hirschfeld ct al. 
and Beam and Cleve in previous experimental work have established the 
Gc inheritance line. 

Also, this study was conducted on two, three generation families. 
One family was Negro and the other family Caucasian of Anglo-Saxon 
descent. The gene frequencies were determined for three generation 

The general technique involved the collection of blood specimens 
for determination of serum protein. The serum was then treated by 
microimmunoelectrophoresis on a cellulose membrane. The Gc precipi- 
tation arcs were compared to control Gc precipitation arcs and identified 
for the correct Gc type. 

The data collected did show that the microimmunoelectrophoresis on 
cellulose acetate could be used routinely to determine the Gc component 
of the serum. This test proved to be easy to perform, accurate, repro- 
ducible, and economical. This experiment confirmed that the Gc system 
was under genetic control and could be used to help establish the 
parentage of a child. 

The verification tests by Hirschfeld could be used to eliminate any 
genetic variation due to rare Gc types. 

The data was not conclusive in determining the gene frequencies of 
the Negro and Caucasian of Anglo-Saxon descent in the midwestern 
United States region because the number of tested individuals was too 
small. In calculating the data, the gene frequencies of Beam and Cleve 
were used. 

Quantitative Measures of In Vitro Cell Mobility by Use of a Pattern 
Recognition Computer. Georges Barski, James W. Butler, and Robert 
J. Thomas, DePauw University. — The purpose of these researches is to 
find a quantitative measure of cell mobility, in vitro, which takes account 
of the changes in the shape of the cell and of the motion of the interior 
parts, while ignoring the random translation and rotation of the cell as 
a whole; and, furthermore, to use a computer to automatically obtain 
such measure. The usefulness of such a measure would be in researches 
regarding effects on mobility of such things as temperature, different 
concentrations of a chemical, different chemicals, different amounts and 
kinds of radiation, and to compare normal with abnormal cells. 

The general procedure is to take motion pictures of the cell and 
use the film as input to Chloe, the Argonne National Laboratory pattern 
recognition computer. Chloe repares a record (on magnetic tape) of 
the shapes it sees in each frame; this record of shapes is then analyzed 
by a standard general purpose computer. The program used for this 
analysis is one which is also used for automatic karyotyping of chromo- 
somes; it computes the area, centers of mass, moment of inertia, and 
other higher moments of the shapes (in combinations which are invariant 

166 Indiana Academy of Science 

under translation and rotation in the plane). Another program then 
analyzes the changes in these quantities from frame to frame and ob- 
tains a measure of the motion of the cell. It is in this latter area that 
most of the research has been and is being carried out, as there are 
many ways to combine and use the information available on the moments. 

Studies on the Mechanisms of Glutaraldehyde Stabilization 
of Cytomembranes 1 

D. James Morre' and Hilton H. Mollenhauer, Department of Botany 

and Plant Pathology, Purdue University, and Charles F. Kettering 

Research Laboratory, Yellow Springs, Ohio. 

The widespread use of glutaraldehyde for routine fixation of bio- 
logical materials (1, 9, 14, 16) and as an aid to organelle isolation (11, 
12, 13) stimulated considerable interest in the nature of its interactions 
with protoplasmic constituents (2, 3, 6). Using a combined biochemical 
and histochemical approach, Hopwood (6) demonstrated qualitatively 
that glutaraldehyde fixation introduced blockable carbonyl groups into 
tissue protein and that glutaraldehyde and formaldehyde preserved gly- 
cogen to a similar degree. However, quantitative studies of the reactions 
of glutaraldehyde with protoplasmic constituents are limited. Evidence 
that glutaraldehyde cross-links polyamines (including proteins) and poly- 
hydroxy compounds (such as polysaccharides) has come largely from 
studies conducted by the tanning industry (2, 6). 

This report presents evidence that cross-linking of free amino 
groups of proteins is critical to glutaraldehyde fixation of cytomem- 
branes, (extractability of membrane lipids is unaffected by glutaralde- 
hyde) and that glutaraldehyde reacts with small molecules such as free 
amino acids and primary amines. Most of the glutaraldehyde bound by 
tissue appears to result from its binding to small molecules and to large 
molecules other than protein. 

Materials and Methods 

Glutaraldehyde. Excised onion stem and isolated cell fractions from 
bean leaves, onion stem and mouse liver were treated with 2.5% glutaral- 
dehyde (prepared from a 50% stock solution, Fischer Scientific, Bio- 
logical Grade, to which 0.16 g/ml of activated coconut charcoal was 
added to remove impurities) at to 4°. Unless stated otherwise, glutaral- 
dehyde solutions were prepared in 0.01 M sodium phosphate buffer, pH 
7.0 to 7.3. 

Determination of protein and nitrogen. Total protein was determined 
from lyophilized fractions which were quick frozen in liquid nitrogen 
and dried to constant weight over calcium chloride. Tissue was pulverized 
to a fine powder using a mortar and pestle and a weighed aliquot was 
extracted exhaustively with hot (90°) 0.1 N sodium hydroxide. Insoluble 
material was removed by centrifugation, the supernatant was neutralized 
with trichloroacetic acid, and the protein was precipitated by addition of 
an equal volume of 1 M trichloroacetic acid (0 to 4°; 2 to 3 hr). Protein 

1 Purdue University AES Journal Paper Number 3552. C. F. Kettering 
Contribution Number 328. Supported in part by grants from the National 
Science Foundation GB-1084, -03044 and -7078 and USPHS Grant GM 15492. 


168 Indiana Academy of Science 

was estimated by the biuret method (5). Nitrogen was determined by- 
direct Nesslerization (7). 

Extraction and determination of lipids. For lipid extraction, about 
4 g fresh weight of tissue (either untreated or glutaraldehyde-fixed) was 
frozen in liquid nitrogen, lyophilized and dried as above, powdered with 
a mortar and pestle and transferred to microsoxhlet extraction thimbles. 
The chloroform extractable lipids were removed by a 3 hr extraction 
with 15 ml of chloroform. Chloroform was evaporated and the lipid resi- 
due was brought to constant weight over calcium chloride, redissolved in 
chloroform, filtered to remove insoluble materials, recovered by evapora- 
tion of solvent and brought to constant weight. Final weights were cor- 
rected for losses during filtration. To obtain the residual lipid fraction, 
the thimbles from the chloroform extraction were placed in acetylization 
flasks with 15 ml of 95% methanol and refluxed for 1 hr over a 90° 
water bath. Methanol was evaporated over a steam bath and water was 
removed by transferring the samples to a 90 to 95° oven. After drying 
to constant weight over calcium chloride, 15 ml diethyl ether were added 
to the residue followed by exhaustive soxhlet extraction (12 hr). The 
chloroform-soluble residue after decantation and evaporation of the 
ether was taken as residual lipid. 

Thin layer chromatography of phospholipids. Approximately 5 mg 
of lipid in chloroform were streaked on each 20 X 20 cm plate coated 
with Silica Gel G. Development was in chloroform :methanol:water 
(65:35:4 v/v). Regions containing lipids were detected by exposing the 
plates to iodine vapors, scraped from the plates and digested with sul- 
furic acid and hydrogen peroxide. Phosphorous content of the digests was 
determined by the method of Fiske and Subbarow (4). 

Electron microscopy. Portions of isolated pellets were post-fixed for 
1 to 24 hours in 1 per cent buffered osmium tetroxide (0.1 M sodium phos- 
phate, pH 7.2) or exposed for several days to osmium tetroxide vapors. 
Subsequent dehydration and embedding was as described previously (10). 
Sections were viewed with a Siemens Elmiskop I or Philips EM 200 
electron microscope. Magnifications are approximate. 


Glutaraldehyde stabilization of organelles during isolation. Loss of 
ultra structural detail in organelles during their isolation was prevented 
by adding 10 1 to 10" 3 M glutaraldehyde (2.5 to 0.0025%) to the homoge- 

Figure 1. Nuclei fraction from onion stem stabilized by addition of glu- 
taraldehyde to the homogenization medium. Osmium post-fixation. X 7,000. 
Figure 2. Mitochondrial fraction of onion stem stabilized by addition of 
glutaraldehyde to the homogenization medium. Osmium post fixation. 
X 13,200. 

Figure 3. Golgi apparatus fraction from onion stem containing sheets of 
rough endoplasmic reticulum (ER) stabilized by addition of glutaraldehyde 
to the homogenization medium. Dictyosomes of the Golgi apparatus are 
shown in cross section (D x and D 2 ) and sectioned tangentially (D 3 ). Osmium 
post-fixation. X 40,000. 

Cell Biology 


*>* ^s%*> 


Indiana Academy of Science 

Figure 4. Smooth membrane fraction from onion stem stabilized by addition 

of glutaraldehyde to the homogenization medium. Many of the large 

vesicular profiles are derived from fragments of the plasma membrane. 

Osmium post-fixation. X 49,000. 

Figure 5. Chloroplast fraction isolated from bean leaves in the absence of 

glutaraldehyde. Glutaraldehyde-osmium fixation. X 6,900. 

Figure 6. Chloroplasts isolated from bean leaves in the presence of 

glutaraldehyde. Osmium post-fixation. X 6,900. 

Figure 7. Glutaraldehyde-stabilized chloroplast at higher magnification. 

Cell Biology 171 

nization medium (0.5 M sucrose; 0.1 M sodium phosphate, pH 7.1; 1% 
dextran). Various fractions including nuclei (Fig. 1), mitochondria (Fig. 
2), Golgi apparatus (Fig. 3), endoplasmic reticulum (Figs. 3 and 8) 
and smooth membranes, including plasma membrane (Fig. 4), were iso- 
lated from onion stem and other plant and animal tissues in the 
presence of glutaraldehyde. These cell components were morphologically 
indistinguishable from organelles fixed in vivo. Nuclei had intact outer 
membranes and organized chromatin (Figs. 1 and 8). Mitochondria were 
dense and unswollen (Fig. 2). In the absence of glutaraldehyde, plant 
Golgi apparatus vesiculated and the stacked cisternae separated. With 
glutaraldehyde, breakdown of both Golgi apparatus (Fig. 3) and endo- 
plasmic reticulum (Figs. 3 and 8) was retarded. With chloroplasts iso- 
lated without glutaraldehyde (Fig. 5), outer membranes, osmiophilic 
granules and stroma (chloroplast matrix) were missing and the lamellae 
were distorted and swollen. With the addition of 2.5% glutaraldehyde to 
the homogenization medium (Fig. 6), the outer membranes, osmiophilic 
granules and lamellae were well preserved and clearly defined. The 
stroma was retained in the uniformly dense aspect characteristic of 
chloroplasts fixed in vivo. 

To illustrate a similar effect of glutaraldehyde on mammalian 
tissues, Figure 8 shows a pellet of an unfractionated homogenate of 
mouse liver prepared in the presence of glutaraldehyde. Organelles in- 
cluding nuclei, mitochondria, microbodies and endoplasmic reticulum 
were obtained with morphological characteristics resembling those of 
whole, fixed tissues. 

Amount of glutaraldehyde bound. With onion stem treated with 
buffer alone, the dry weight from 12 determinations was 11.3±0.3% 
of the fresh weight. With 2.5% buffered glutaraldehyde followed by four 
15 minute buffer rinses, the final dry weight was 12.4±0.5% of the fresh 
weight. The gain in dry weight due to glutaraldehyde treatment was 
1.1% of the fresh weight or about 10% of the dry weight. The increase 
in dry weight, if due to bound glutaraldehyde, would amount to 110 
/xmoles of glutaraldehyde per g fresh weight. 

Reaction of glutaraldehyde with lipids and proteins. Total lipid ex- 
tracted by chloroform-methanol-ether consisted of about 40% phospho- 
lipid and 60% neutral lipid and was unchanged by glutaraldehyde treat- 
ment of the tissue (6.1% of the dry weight) as shown by data of Table 1. 
In contrast, total protein extracted by 0.1 N sodium hydroxide declined 
from 14% of the dry weight (in untreated controls) to 3.5% of the dry 
weight after glutaraldehyde treatment. These data demonstrate that 75% 
of the protein was sufficiently cross linked to prevent extraction into 
alkali. Total nitrogen was unchanged at 3.2% of the dry weight. 

When glutaraldehyde was added to protein in solution (Fig. 9), the 
bulk of the protein was rendered insoluble with a reaction half-time of 
approximately 7 min. As with whole tissue, about 75% of the protein 
was sufficiently cross-linked to become insoluble in 0.1 N sodium hy- 
droxide following glutaraldehyde treatment. 


m. - 

Indiana Academy of Science 





Figure 8. Pellet of unfractionated mouse liver homogenized in the presence 
of glutaraldehyde. A nucleus (N), segments of rough endoplasmic reticulum 
(RER), a microbody (^B) and numerous mitochondria (M) are shown. 
Osmium post-fixation. X 16,500. 

Cell Biology 


Although lipid extractability was unchanged by glutaraldehyde 
treatment (Table 1), it was of interest to consider the possibility that 
certain lipids might react with glutaraldehyde and yet remain lipid solu- 
ble and be extracted into the chloroform-soluble fraction. To test this 
possibility, the phospholipid fraction was separated by thin layer chroma- 
tography into low Rf phospholipids (glyco- and sulfolipids), lecithin 
(phosphatidylcholines), ethanolamine phospholipids (phosphatidylethanol- 
amines) and two phospholipid fractions containing serine (phosphatidyl- 
serines). As shown by data of Table 2, lecithin accounted for about 30% 
of the total lipid phosphorous and was unchanged by glutaraldehyde 
treatment. In contrast, the amino phospholipids (those containing serine 
and ethanolamine) were greatly reduced in amount at positions corre- 
sponding to their characteristic R f 's on the thin layer plates. Recovery 
of serine phospholipids was 35% and recovery of ethanolamine phospho- 
lipids was 66% in contrast to 95% recovery for lecithin. An apparent 
increase in low R f phospholipids following glutaraldehyde treatment 
approximately compensated for the decrease in ethanolamine and serine 
phospholipids. Total recovery of lipid phosphorus was 94% of that of 
unfixed control tissue. 

Reaction of glutaraldehyde with small molecules. Glutaraldehyde 
reacted rapidly with the primary amino groups of all twenty protein 
amino acids tested (Table 3) as evidenced by loss of ninhydrin reactivity. 
Paralleling the loss of chemical reactivity of the a-amino group, was the 

12 3 4 


Figure 9. Solubility of bovine serum albumin in 1.0 N sodium hydroxide as 
a function of time in a 2.5% glutaraldehyde solution at to 4°. 

174 Indiana Academy of Science 

table 1. Composition of Onion Stem Determined in the Presence or 
Absence of Glutar aldehyde (GA). 

% of Dry Weight 

Constituent — GA +GA 

Total Lipids 6.1 6.1 

40% Phospholipids 

60% Neutral Lipids 

Alkali Extractable Protein 14.0 3.5 

Total Nitrogen 3.2 3.2 

TABLE 2. Phospholipid Composition of Onion Stem Determined in the 
Presence or Absence of Glutar aldehyde (GA). 

Lipid Phosphorous 

'> (v-glg 

dry wei 







— GA 

+GA - 


+ GA 

— GA +GA 

Low Rf Phospholipids 














Serine Phospholipids A 







Ethanolamine Phospholipids 







Serine Phospholipids B 







Sterols and Neutral Lipids 














appearance of a yellow chromogen with an absorption maximum near 
440 m/x and suggestive of Schiff base formation. Cysteine, in addition to 
forming the yellow coloration given by all amino acids, rapidly pro- 
duced an insoluble precipitate not given by cysteine. The imino acids, 
proline and hydroxyproline, retained ninhydrin reactivity following glu- 
taraldehyde treatment. Proline did not react. 

Of the nucleotide bases tested, adenine and guanine were most re- 
active with the formation of an ultraviolet light-absorbing adduct and 
the yellow coloration. Neither cytidine nor uridine produced a visible 
reaction at room temperature but upon heating reacted in a manner 
similar to adenine. 

Role of protein cross-Unking in cytomembrane stabilization. To 
test the relative contributions of protein and lipid in maintaining the 
form of cytomembranes, a cell fraction containing dictyosomes and a 
few fragments of endoplasmic reticulum (Fig. 3) was prepared by glu- 
taraldehyde stabilization and lyophilized. After drying to constant weight 

Cell Biology 





-* ■$* 












' ' 




Golgi apparatus fraction from onion stem stabilized by addition 
of glutaraldehyde to the homogenization medium, freeze dried, stained with 
osmium vapor and embedded in epon-araldite. The field was chosen to 
contain a fragment of endoplasmic reticulum (ER) as well as a dictyosome 
showing cisternae sectioned both tangentially and in cross section. 
X 120,000. 

Figure 11. As in Figure 10 except lipids removed by chloroform- 
methanol-ether extraction prior to exposure to osmium. X 120,000. 

176 Indiana Academy of Science 

over calcium chloride, the lipids were extracted with chloroform- 
methanol-ether from one half of each pellet as described for whole tissue. 
The other pellet half served as an unextracted control. Prior to examina- 
tion in the electron microscope, both pellets were stained by exposure to 
osmium tetroxide vapor and embedded directly in an epon-araldite mix- 
ture (10). Figure 10 shows a portion of a control pellet with a single 
dictyosome and a segment of endoplasmic reticulum. With both cell com- 
ponents, the osmium was localized largely in the lumina of dictyosome 
cistemae and with both the lumina and ribosomes of the endoplasmic 

In the extracted pellet (Fig. 11), the appearance of the membrane 
was similar to that of the unextracted controls except that the dictyo- 
somes and endoplasmic reticulum were more clearly denned. It is clear 
that chloroform extraction of lipids did not alter the form of glutaralde- 
hyde-stabilized dictyosomes and endoplasmic reticulum, and that the 
reactions of glutaraldehyde with protein of the membrane were appar- 
ently sufficient to stabilize the form of the organelle as viewed in the 
electron microscope. 

Binding of small molecules to membranes in homogenates. Since 
glutaraldehyde is a dialdehyde, it was possible that reactive small mole- 
cules were bound to cytomembranes through reaction of the second alde- 
hyde to reactive groups on large molecules. To test this possibility, radio- 
active metabolites were thoroughly mixed with homogenized onion stem 

table 3. Reactivity of Amino Acids, Imino Acids and Nucleotide Bases 
With Glutaraldehyde. 

Amino Acids* 




+ + 


+ + 


+ + 

Aspartic Acid 

+ + 








+ + 



Glutamic Acid 


Imino Acids 3 - 








+ + 




+ + 

Nucleotide Bases 


+ + 




+ + 







a Determined from loss of ninhydrin reactivity following- paper chromatog- 
raphy and from increase in absorbance at 440 m^ during reaction in 

b In addition to the yellow chromogen given also for cystine, cysteine gave 
a copious precipitate not given by cystine suggesting a reaction of glu- 
taraldehyde with the cysteine -SH. 

c Determined from altered chromatographic properties and absorbance at 

Cell Biology 


Glutaraldehyde Cone, % 

Figure 12. Inactivation of the enzyme CDP-choline-cytidyl transferase of 
homogenates of onion stem stabilized with varying- concentrations of 

(same medium as for organelle isolation) after which glutaraldehyde was 
added to a final concentration of 2.5%. After 30 minutes at to 4°, 
the homogenates were pelleted at 12,000 g for 30 min and washed with 
distilled water through several resuspension-centrifugation cycles until 
constant specific activity was achieved. With C 14 -acetate, a molecule not 
expected to react with glutaraldehyde, the level of radioactivity in the 
homogenate was 1.8 m/anoles/^mole/mg N. Choline gave a similar value 
which suggested little or no binding. The value for C 14 -uridine, although 
obtained from mung bean stem homogenates rather than onion, was 
sufficiently low to suggest little or no binding. Radioactivity from C 14 - 
glucose bound to the 12,000 g pellet was about 3 times that of acetate 
showing a small but significant reactivity of glutaraldehyde with a sugar. 
Of the compounds tested, leucine was most reactive, binding at 5 times 
the amount of glucose. In competition experiments, it was found that the 
level of radioactive leucine bound to components of the homogenates was 
little influenced by a 100-fold excess of unlabeled leucine. 

Loss of enzymatic activity. Typical denaturation kinetics are shown 
in Figure 12 for the enzyme CDP-choline-cytidyl transferase of onion 
stem. This enzyme catalyzes the formation of CDP-choline, an inter- 
mediate in the biosynthesis of lecithin, from phosphorylcholine (P- 
choline) and cytidine triphosphate (CTP). 

178 Indiana Academy of Science 


Although glutaraldehyde provides superior structural preservation 
for electron microscopy (1, 9, 14, 16), its use in organelle stabilization 
or in metabolic studies may modify enzymatic activity. Loss of enzymatic 
activity is generally associated with protein cross-linking by glutaralde- 
hyde (3). Examples of losses of enzymatic activity are given by Hop- 
wood (6), Sabatini et al. (14), Barrnett (1) and in Figure 12. Not all 
enzymatic activities decline following treatment with glutaraldehyde. 
Some activities appear to be unchanged (15), and De Jong et al. (3) 
have reported activation by glutaraldehyde of an acid phosphatase of 
nuclei in cultured plant cells. Preliminary results indicate an activation 
of a CTPase of dictyosomes isolated from onion stem following treat- 
ment of homogenates with glutaraldehyde (unpublished observations). 

For a fixative to be of general use in preparing tissues or organelles 
for chemical analysis, the chemical moiety under study must remain un- 
modified. With onion stem, lipids are recovered quantitatively after glu- 
taraldehyde stabilization and the phospholipid lecithin appears to be un- 
changed. The lipid content after treatment appears to reflect the lipid 
content before fixation. Glutaraldehyde-stabilized dictyosome prepara- 
tions from onion stem contain 50 to 60% protein, 20 to 30% phospholi- 
pids (rich in lecithin) with the remainder consisting mostly of sterols 
and neutral lipids (12). 

Studies of Park et al. (13) suggest that quantum conversion and 
electron transport in photosynthesis takes place within the rigid frame- 
work of glutaraldehyde-stabilized plastids and that these processes do 
not require conformational changes in proteins. They showed that chloro- 
plasts isolated from leaves previously fixed in 6% glutaraldehyde retain 
optical rotatory dispersion activity and absorption spectra of unfixed 
chloroplasts and the capability of limited photochemical activity. Ferri- 
cyanide-Hill activity measured by 2 evolution or indophenol reduction 
was 25% of that of unfixed chloroplasts. The oxygen evolution was light 
dependent and sensitive to various photosynthetic inhibitors. 

That the morphological integrity of glutaraldehyde-treated organ- 
elles is maintained after lipid extraction suggests that the cross-linked 
protein framework of the organelle is the critical feature of glutaralde- 
hyde stabilization. Golgi apparatus fractions containing small fragments 
of endoplasmic reticulum when isolated by glutaraldehyde stabilization 
showed little or no morphological response to changes in osmotic environ- 
ment. They maintained their in vivo shapes even after freeze drying 
(Fig. 10) or freeze diying followed by extraction of total lipids using 
chloroform-methanol-ether (Fig. 11). Except for slight increases in 
osmium deposition after lipid extraction such preparations were indis- 
tinguishable from each other. 

Associated with the reaction of glutaraldehyde with tissue, tissue 
homogenates or isolated cell fractions is a yellow coloration which in- 
creases in intensity for about 1 hour after addition of glutaraldehyde at 
to 4°C. Various protein, amino acid or amine solutions treated with 

Cell Biology 179 

glutaraldehyde assume a similar color. The color is probably due to the 
formation of Schiff bases, which in proteins would be with basic amino 
acids. Cystine which consists of two cysteine residues linked by a di- 
sulfide bond did not precipitate with glutaraldehyde as did cysteine. This 
is evidence for a reaction of glutaraldehyde with the free sulfhydryl 
group of cysteine. The imino acids proline and hydroxyproline contain an 
amino group in the ring which is apparently unreactive with glutaralde- 
hyde. Hydroxyproline appeared to react slowly with glutaraldehyde pos- 
sibly through an involvement of the hydroxyl group. 

Being a dialdehyde, glutaraldehyde is capable of forming cross- 
linking bridges in proteins involving both of the aldehyde groups, or 
under certain circumstances only one of the aldehyde groups may react 
with protein. Hopwood (6) demonstrated qualitatively that free carbonyl 
groups are introduced into proteins following reaction with glutaralde- 
hyde. The free aldehyde could then react with other cell components in- 
cluding free amino acids. That this type of reaction occurs is shown by 
data of Table 4. Radioactive leucine added to total homogenates of 
onion stem cross-links to organelles sedimenting at 12,000 g. This bind- 
ing is apparently nonspecific and in competition with a variety of re- 
active molecules since addition of excess unlabeled leucine had little effect 
on the amount of labeled leucine bound by the organelles. Other metab- 
olites including choline, acetate and uridine are not bound significantly. 
Glucose is bound to the organelle fraction, presumably via cross-linking 
reactions involving the free hydroxyls, at a level 1/10 that of leucine. 

These results show that glutaraldehyde fixation of tissues or or- 
ganelle fractions incubated with radioactive metabolites may lead to non- 
specific binding of the metabolites via the cross-linking reactions. This 
possibility must be taken into account in autoradiographic or biochemical 
studies involving radioactive metabolites and especially with amino 
acids when supplied in conjunction with glutaraldehyde stabilization. 

table 4. Binding by Glutaraldehyde of C H -Labeled Metabolites to the 
12,000 o Particulate Fraction of Onion Stem*. 

Metabolite m^Moles/VMole/Mg Nitrogen 

Acetate-2-C 14 1.8 

Choline-2-C 14 1.8 

Uridine-U-C lib 0.7 

Glucose-U-C 14 5.5 

Leucine-U-C 14 50.4 

a Homogenates were mixed with isotope (1.5 to 30 ^moles/5 g fresh weight 
of tissue in 6 ml of total homogenate) followed by glutaraldehyde (final 
concentration of 2.5%) and equilibrated 30 minutes. After clearing the 
mixture of cellar debris, the 12,000 g pellet was washed with buffer to 
constant specific activity on a total nitrogen basis. Values are ± 30%. 

b Mung bean stem. 

180 Indiana Academy of Science 

To summarize our findings, calculations of the distribution of po- 
tential glutaraldehyde binding sites among various cell constituents is 
provided in Table 5. From the dry weight changes associated with re- 
action with glutaraldehyde, approximately 110 /mioles of glutaraldehyde 
are bound per g fresh weight of onion stem. Of this 110 ^moles, less 
than 15 might be expected to bind with protein on the basis of lysine 
and cysteine content. Other basic or hydroxy amino acids may react with 
glutaraldehyde but evidence presented here and elsewhere (2) suggests 
the a-amino group of lysine and the j3-SH group of cysteine as poten- 
tially the most reactive groups in the interiors of protein chains (pre- 
sumably the N-terminal residue would also be available for reaction with 
glutaraldehyde). The potential contribution of nucleic acid to the amount 
of glutaraldehyde bound in onion stem is probably less than 2 ^moles due 
to the small amounts of nucleic acid present. Adenine and guanine (A + 
G) are considerably more reactive in solution as the free base than 
either uridine or cytidine (U + C). Amino lipids react with glutaralde- 
hyde but represent only 25% of the lipid phosphorus and therefore ac- 
count for less than 1 ^mole of the total glutaraldehyde bound. The largest 
proportion of glutaraldehyde bound to amino groups would be to free 
amino acids and other small molecules containing primary amines. As- 
suming that 80% of the nonprotein nitrogen consists of amino nitrogen 
(8), this class of molecules could account for up to 55 ^moles of glutaral- 
dehyde per g fresh weight or approximately 50% of the total glutaral- 
dehyde bound by the tissue. 

We have no estimate of the amount of glutaraldehyde bound to cell 
wall, storage polysaccharides and small molecules such as sugars or 
alcohols. Assuming maximal binding to amines, this fraction would repre- 
sent about 37 /rnioles of the 110 ^moles total. However, it appears that 
under usual conditions of fixation no more than 75% of the available 

table 5. Relative Proportions of Potential Glutaraldehyde Binding Sites 

of Onion Stem. 

(i Moles Glutaraldehyde Bound 
Constituent per G Fresh Weight 

Total (By Weight) -110 

Protein (Calculated on the basis of 6.5 g lysine <15 

N and 1 g cysteine N per 100 g protein N) (8) 
Nucleic Acid (Calculated on the basis of 0.45% of 
the dry weight; 50% A + G) (8) <2 

Free Amino Acids and Primary Amines (Calcu- 
lated on the basis of 6.6% of the dry weight; 
average M. W. 120) -55 

Amino Lipids (Calculated on the basis of 25% of 
the total lipid P; average M. W. 750) <1 

Unaccounted For (Cell wall, storage polysaccha- 
rides, small molecules other than primary amines) >37 

Cell Biology 181 

sites react with glutaraldehyde (Fig. 9, Table 2 and estimates from re- 
action of amino acids and nucleic acids in solution). If 75% is used as the 
extent of reaction, less than half the bound glutaraldehyde is accounted 
for by reaction with amino and sulfhydryl groups (both as polymers or 
small molecules). Thus, binding of glutaraldehyde to cell wall and other 
polyhydroxycompounds might be more extensive than the 37 /mioles/g 
fresh weight estimated and, in plant tissues, might account for binding 
equal in magnitude to that by primary amines. 


Chemical evidence suggests that glutaraldehyde stabilization of cyto- 
membranes occurs primarily by cross-linking free amino groups of 
adjacent polypeptide chains. Hydroxy 1 and sulfhydryl groups may also 
contribute to the cross linking. However, 90% or more of the total 
lipid of glutaraldehyde treated tissue is recovered by solvent extraction. 
The bulk of the glutaraldehyde bound by tissue appears to be the result 
of its binding to small molecules and to large molecules other than pro- 
tein. Cross-linking of protein, however, appears critical to glutaralde- 
hyde fixation of cytomembranes. 

Literature Cited 

1. Barrnett, R. J. 1964. Localization of enzymatic activity at the fine 
structural level. J. Royal Micros. Soc. 83:143-151. 

2. Bowes, J. H., and C. W. Cater. 1966. The reaction of glutaraldehyde with 
proteins and other biological materials. J. Royal Micros. Soc. 85:193-200. 

3. DeJong, D. W., A. C. Olson, and E. F. Jansen. 1967. Glutaraldehyde acti- 
vation of nuclear acid phosphatase in cultured plant cells. Science 

4. Fiske, C. H., and Y. Subbarow. 1925. The colorimetric determination of 
phosphorus. J. Biol. Chem. 66:375-400. 

5. Gornall, A. G., C. J. Bardawill, and M. M. David. 1949. Determination of 
serum proteins by means of the biuret reaction. J. Biol. Chem. 

6. Hopwood, D. 1967. Some aspects of fixation with glutaraldehyde. J. 
Anatomy 101:83-92. 

7. Koch, F. C, and T. L. McMeekin. 1924. A new direct Nesslerization 
micro-Kjeldahl method and a modification of the Nessler-Folin reagent 
for ammonia. J. Am. Chem. Soc. 46:2066-2069. 

8. Long, C. 1961. Biochemists Handbook. D. Van Nostrand, New York. 

9. Maunsbach, A. B. 1966. The influence of different fixatives and fixation 
methods on the ultrastructure of rat kidney proximal tubule cells. II. 
Effects of varying osmolarity, ionic strength, buffer system and fixative 
concentration of glutaraldehyde solutions. J. Ultrastructure Res. 

10. Mollenhauer, H. H. 1964. Plastic embedding mixtures for use in elec- 
tron microscopy. Stain Technology 39:111-114. 

11. Moore', D. J., H. H. Mollenhauer, and J. E. Chambers. 1965. Glutaralde- 
hyde stabilization as an aid to Golgi apparatus isolation. Experimental 
Cell Res. 38:672-675. 

12. Morre*, D. J., H. H. Mollenhauer, and W.P.Cunningham. 1965. Chemical 
and structural analysis of dialdehyde stabilized Golgi apparatus. Plant 
Physiol. 40 (Suppl.):39. 

182 Indiana Academy of Science 

13. Park, R. B., J. Kelly, S. Drurt, and K. Saurer. 1966. The Hill reaction of 
chloroplasts isolated from glutaraldehyde-fixed spinach leaves. Proc. 
Nat. Acad. Sci. 55:1056-1062. 

14. Sabatini, D. D., K. Bensch, and R. J. Barrnett. 1963. Cytochemistry and 
electron microscopy. The preservation of cellular ultrastructure and 
enzyme activity by aldehyde fixation. J. Cell Biol. 17:19-58. 

15. Sommer, J. R., and J. J. Blum. 1965. Cytochemical localization of acid 
phosphatases in Euglena gracilis. J. Cell Biol. 24:235-251. 

16. Trump, B. F., and R. E. N. Bulger. 1966. New ultrastructural character- 
istics of cells fixed in a glutaraldehyde-osmium tetroxide mixture. Lab. 
Invest. 15:368-379. 

Adenosine Mono-, Di- and Trinucleotidase Activities 
of Rat Liver Cytomembranes 1 

Arthur E. Middleton, Ronald Cheetam, Donald Gerber and D. J. 
Morre', Department of Botany and Plant Pathology Purdue University. 

The liver continuously secretes triglycerides and cholesterol 
into the circulation in the form of very low density lipoproteins (6). The 
successful isolation of the various components of the secretory system 
(endoplasmic reticulum, Golgi apparatus and plasma membrane) from 
rat liver (7) provides new approaches to the identification and characteri- 
zation of these structures. This report describes the procedure for the 
isolation of plasma membrane from rat liver adopted from those of 
Neville (8) and Emmelot et al. (4). The rates at which these plasma 
membrane fractions hydrolyze the mono-, di- and triphosphates of adeno- 
sine are compared with those of other components of the rat liver secre- 
tory system. 

Materials and Methods 

Isolation of plasma membrane. Male rates, 200 to 300 g (50 days 
old), purchased from the Holtzman Company, 421 Holtzman Road, Madi- 
son, Wisconsin, were anesthetized by intraperitoneal injections of 0.5 to 
1 ml pentobarbital solution (Abbott Laboratories Nembutal, 20 mg/ml). 
Each liver was drained of blood, minced finely and homogenized in 5 
to 10 ml 1 mM sodium bicarbonate using a very loose fitting Potter- 
Elvejheim all-glass tissue homogenizer (30 sec at 1,000 rpm). The 
homogenate was mixed with 300 ml of 1 mM sodium bicarbonate and 
stirred for 4 min, after which the suspension was filtered through a single 
layer of premoistened (with bicarbonate) cheesecloth followed by a second 
filtration through two layers of premoistened cheesecloth. The filtered 
homogenate was then centrifuged for 10 min at 3,500 rpm (about 1,500 g) 
using 50 ml centrifuge tubes and an angle rotor (Servall SS-34). The 
supernatant was discarded and the pellet was resuspended in 80 ml 
1 mM bicarbonate. The 1,500 g centrifugation-resuspension cycles were 
continued until a clear supernatant was obtained (usually 3 to 4 times). 

The final washed pellets were combined in 1 mM bicarbonate (total 
volume of 3 ml) and 9 ml of sucrose (density 1.3) was added slowly (by 
drops) with constant stirring. The following gradient was layered over 
the plasma membrane suspension: 5 ml sucrose density 1.20; 8 ml su- 
crose density 1.18; 5 ml sucrose density 1.16; 4 ml sucrose density 1.14. 
The gradient was then centrifuged for 75 min at 24,000 rpm (SW 25 
rotor, Spinco Model L Ultracentrifuge). Plasma membrane was collected 
from the 1.16-1.18 density interface. 

1 Purdue University AES Journal Paper No. 3553. Work supported in 
part by grants from the NSF GB7078, GY4122, and GY2683. 



Indiana Academy of Science 

Methods for obtaining Golgi apparatus and endoplasmic reticulum 
fractions will be described in detail elsewhere. These fractions were rela- 
tively free of plasma membrane and mitochondrial contamination (7). 

Enzyme assays. Assays for 5'-mononucleodidase (EC with 
AMP as substrate and ADPase (EC were performed with the 
nucleotides (5 mM) as sodium salts in a medium containing 100 mM 
KC1, 5 mM MgCl 2 and 50 mM Tris (pH 7.4) and a total volume of 2 
ml with an incubation time of 15 min at 37°. For assays of the Na — 
K — Mg 2 — ATPase, the potassium salt of ATP was used in a medium 

-it ' 

, . 4-.. 




::■■ ,:■ 





n 4 


%. i 


>rl %^ x ' 

f ; - % 




" ■<%• 



f V 

Figure 1. Thin section of plasma membrane pellet. Two junctional com- 
plexes are shown at arrows. Osmium fixation. X 33,300. 

Cell Biology 185 

containing 66 mM NaCl, 33 mM KC1, 5 mM MgCl 2 and 25 mM Tris 
(pH 7.4). The corresponding medium for the Mg 3 — ATPase (EC contained 100 mM KC1 and no NaCl according to the procedure 
of Emmelot and Bos (3). Protein was determined by the Lowry procedure 
(5). The complete reaction mixtures contained approximately 0.1 mg 

Electron microscopy. Samples of isolated fractions were prepared 
for electron microscopy by fixation in 2% osmium tetroxide buffered to 
pH 7.1 with sodium phosphate (12 hrs, 4°) followed by dehydration and 
embedding in an Epon-Araldite plastic mixture. Negative staining of the 
unfixed plasma membrane fractions was done on carbon-stabilized, 
collodion-coated grids by first resuspending a portion of the pellet in 
water followed by mixing the suspension with an equal volume of 2% 
phosphotungstate (PTA) neutralized with potassium hydroxide to pH 
6.8. Specimens were observed and photographed using a Philips EM/ 200. 

Results and Discussion 

Yield and purity of membranes. The yield of plasma membrane was 
1 to 4 mg protein from 10 g fresh weight of liver. A 20-fold enrichment 
of 5'-nucleotidase and a 50-fold enrichment of Na + -activated ATPase 
relative to the total homogenate was obtained (Tables 1 and 2). The 
final pellets consisted of a pinkish-white layer of plasma membrane over 
a small layer of mitochondria. Based on estimates of succinic dehydro- 
genase (9), the activity of the plasma membrane fraction was 25% that 
of purified mitochondria. Glucose-6-phosphatase activity was 25% that 
of purified endoplasmic reticulum. Perhaps no more than 50% of the 
final pellet was plasma membrane. 

Electron microscopy. Samples of preparations taken for electron 
microscopy contained vesicles of many sizes (Fig. 1). Occasionally a 
desmosome, a feature of the plasma membrane, was observed. Granules 
suggestive of ribosomes were not observed but mitochondria and 
mitochondrial fragments were prevalent. 

table 1. Specific Activities of the ATPase and (Na + -K+) — ATPase of 
the Plasma Membrane-Rich Cell Fraction and of the Total Homogenate 

from Rat Liver. 

Specific Activity 


(^Moles iP/Hour/Mg Protein) 

Cell Fraction 





Total Homogenate 

Mg+ + ,K + 




Plasma Membrane 

Mg++,K + ,Na+ 







+ 0.10 

Mg+ + ,K + 




Mg++,K + ,Na + 




ANa + 




186 Indiana Academy of Science 

table 2. SjJecific Activities of AMPase and ADPase of Rat Liver Cell 


Specific Activity 
(,uMoles iP/Hour/Mg Protein) 
Cell Fraction AMP ± SD ADP ± SD ATP ± SD 

Total Homogenate 

1.2 ± 0.4 

1.2 ± 0.3 

0.7 ± 0.4 

Plasma Membrane 

23.0 ± 3.0 

3.6 ± 2.6 

9.6 ± 6.1 

Golgi Apparatus 

5.8 ± 0.3 

1.7 ± 0.4 

2.5 ± 0.9 

Endoplasmic Reticulum 

0.1 ± 0.8 

0.1 ± 0.1 

2.8 ± 1.3 


4.7 ± 3.1 



1.2 ± 0.3 



* Single determination 

After staining with phosphotungstic acid (PTA), the collapsed mem- 
branes were embedded in a thin amorphous film of stain. Most of the 
membranes examined showed a fine granular structure in surface view 
with smooth edges. Occasionally, membrane margins exhibited globular 
knobs with an approximate diameter of 50 or 60 A (Fig. 2). These knobs 
have been described as characteristic of certain regions of the plasma 
membrane (1, 2), probably the microvilli extruding into the bile spaces 
(Benedetti and Emmelot, unpublished). In addition, a hexagonal array 
of subunits was observed in certain membrane sheets (Fig. 3) as pre- 
viously reported by Benedetti and Emmelot (1). The hexagonal subunit 
patterns were prevalent in preparations containing tight junctions (2). 

Sodium-potassium activated ATPase. The specific activities of the 
Mg a +-ATPase and (Na + -K + )-ATPase of freshly prepared plasma mem- 
brane (Table 1) showed considerable variation from preparation to 
preparation. This is consistent with reports for other membrane ATPase 
systems (3). The (Na+-K+) -ATPase is the Na + -mediated increase of 
inorganic phosphate released from ATP over that observed in the absence 
of Na+ with adjustment of [K+] to maintain osmolarity (Table 1). 
Sodium ions increased the inorganic phosphate released from ATP in the 
presence of potassium ions by most plasma membrane preparations to 
about 170% that obtained in the absence of sodium ions. Results with 
Golgi apparatus and endoplasmic reticulum fractions were variable and 

5' '-Nucleotidase. As summarized in Table 2, the 5'nucleotidase 
(AMPase) activities of the plasma membrane fractions were significantly 
higher than either endoplasmic reticulum or Golgi apparatus. This 
activity appears to be localized as an intrinsic constituent of the liver 
plasma membrane and serves as a convenient measure of plasma mem- 
brane contamination of these cell fractions. 

ADPase. In contrast to previous reports of Emmelot and Bos (3), 
plasma membrane fractions obtained by our procedures showed a low 
level of ADPase activity (Table 2). It has not been verified that this 
increase in inorganic phosphorus is due to a genuine ADPase. 

Cell Biology 


Figure 2. Liver plasma membrane negatively stained with phosphotungstic 
acid. Globular knobs are found at the edge of some membrane sheets 
(arrows). X 300,000. 

Figure 3. Liver plasma membrane negatively stained with phosphotungstic 
acid. The surface is dotted with small particles in hexagonal array. 
X 90,000. 

188 Indiana Academy of Science 


A plasma membrane-rich cell fraction isolated from rat liver was 
shown to contain both a Mg' J+ — ATPase and a sodium-potassium-activated 
ATPase. The fraction was characterized by a high 5'-nucleotidase activ- 
ity with AMP as substrate. This activity was low or absent in isolated 
Golgi apparatus and endoplasmic reticulum fractions. The absence of 
AMPase in other cytomembrane fractions and the ease and unambiguity 
of its estimation makes AMPase a useful marker enzyme for plasma 

Literature Cited 

1. Benedetti, E. L.., and P. Emmklot. 1965. Electron microscopic observations 
on negatively stained plasma membranes isolated from rat liver. J. Cell 
Biol. 26:299-305. 

2. Benedetti, E. L.., and P. Emmelot. 1968. Hexagonal array of subunits in 
tight junctions separated from isolated rat liver plasma membranes. 
J. Cell Biol. 38:15-24. 

3. Emmelot, P., and C. J. Bos. 1966. Studies on plasma membranes. III. 
Mg a + — ATPase, (Na+' — K+ — Mg 2 +) — ATPase and 5'-nucleotidase activ- 
ity of plasma membranes isolated from rat liver. Biochim. Biophys. Acta 


4. Emmelot, P., C. J. Bos, E. L. Benedetti, and P. Rumke. 1964. Studies on 
plasma membranes. I. Chemical composition and enzyme content of 
plasma membranes isolated from rat liver. Biochim. Biophys. Acta 

5. Lowry, O. H., N. J. Rosebrough, A. L. Farr, and R. J. Randall. 1951. 
Protein measurement with the Folin phenol reagent. J. Biol. Chem. 


6. Middleton, A. E., D. J. Morre', L. M. Alves, R. L. Hamilton, and R. 
Mahley. 1968. Immunochemical identification of very low density serum 
lipoproteins in Golgi apparatus from rat liver. Proc. Indiana Acad. Sci. 

7. Morre', D. J., R. Cheetham, and W. Yunghans. 1968. Biochemical char- 
acterization of a Golgi apparatus-rich cell fraction isolated from rat 
liver. J. Cell Biol. 39:96a. 

8. Neville, D. M. 1960. The isolation of a cell membrane fraction from rat 
liver. J. Biophys. Biochem. Cytol. 8:413-422. 

9. Pennington, R. J. 1961. Biochemistry of dystrophic muscle. Mitochondrial 
succinate-tetrazolium reductase and adenosine triphosphatase. Biochem. 
J. 80:649-654. 

Membranifibrils on Cristae and Grana Membranes 1 

J. D. Hall, J. W. Stiles, Y. Awasthi, and F. L. Crane 
Purdue University 


Mitochondria cristae show a linear arrangement of the 90A headpieces 
on the membrane surface when observed by negative staining. Treatment 
with Triton releases fibrous structures of 70A diameter to which the 90A 
headpieces are attached. (J. W. Stiles, Ph.D. thesis, Purdue University, 
1969) These fibers have been purified and contain 15% lipid and very little 
cytochrome. Extraction of the membranifibril with acetone leaves a 30A 
diameter fibrous, insoluble protein which may be the basis for the fibrous 

Similar linear arrangements of particles variously referred to as 
quantosomes or chloroplast Fi have been observed on chloroplast grana 
membranes. Extraction of the grana with triton releases fibrous structures 
of 70A diameter to which 100A particles are attached. We propose that the 
basis of linear organization of particles on the surface of cristae and grana 
membranes is based on their attachment to a fibrillar structure in the 
membrane called the membranifibril. 


Globular particles have been demonstrated on the surfaces of various 
biological membranes. The membranes of mitochondrial cristae and of 
chloroplast grana are two such examples. Spherical 90a particles are 
seen projecting on stalks from the surface of inner mitochondrial mem- 
branes when observed by negative staining (3). Oligomycin-insensitive 
ATPase activity has been shown to be associated with these particles 
(15). Numerous particles, 70-140A in diameter, are found associated with 
the membranes of chloroplast lamellae when viewed by negative stain- 
ing. Some of these particles have been identified as carboxydismutase 
(8, 21). Others have been variously associated with chloroplast fraction I 
protein, ca + +-dependent ATPase, and quantasomes (1, 9, 14). 

The regular orientation of particles on the surfaces of their respec- 
tive membranes suggests that a discrete substructure may be responsible 
for holding particles in place. Evidence for linear substructure in mito- 
chondrial cristae (18, 19) and chloroplast grana (1, 14, 16) has been 
previously reported. Evidence has also been presented for fibers in 
plasma membrane (5). Linear elements have been observed in cholate 
extracts of mitochondria, but have not been interpreted as fibers (10, 13). 
A fibrillar structure which may be the basis of linear organization of 
particles in chloroplast grana and mitochondrial cristae will be described 
in this paper. 

1 Supported under a grant AM04663 from the National Institutes of 
Arthritis and Metabolic Diseases, and training grants GMOH95 (J. W. S.) 
and GM01392 (J. D. H.) and career grant K6-21,S39 (F. E. C.) from the 
National Institutes of General Medical Science. 


190 Indiana Academy of Science 

Materials and Methods 

Beef heart mitochondria prepared according to the method of Low 
and Vallin (12) were homogenized in 0.25 M sucrose and stored at — 20° C 
for at least 4 days prior to preparation of electron transport particles 
(ETP). Following thawing, the mitochondria were homogenized in 0.001 
M Tris-HCl, pH 7.4, and centrifuged at 11,700 x g. The pellet was 
washed 4 times in this manner then resuspended in the buffer. The 
suspension was sonicated for two 5-min. periods in a brine bath using a 
Branson probe sonifier at maximal output. Heavy beef heart mitochondria 
were removed by centrifugation, at 11,700 x g. The supernatant was 
then centrifuged at 78,000 x g for 30 min. The pellet was washed and 
resuspended in 0.001 M Tris-HCl, pH 7.4. If the ETP were to be stored, 
the preserving medium of Hansen and Smith (7) was used. 

To ETP suspended in buffer at a protein concentration of 30 mg/ml 
was added aqueous 10% Triton X-114 to give a final concentration of 
Triton of 3%. Following mixing, the solution was cooled in ice for 15 
min. then solid KC1 was added to a final concentration of 0.2 M. The mix- 
ture was incubated for 30 min. with stirring. Centrifugation at 78,000 
x g for 30 min. yielded a yellow supernatant which was further purified 
by dilution with an equal volume of 0.001 M Tris-HCl, pH 7.4, which was 
0.2 M in KC1, followed by centrifugation. Ammonium sulfate fractiona- 
tion of the supernatant was achieved by the addition of the solid salt to 
25% saturation. Centrifugation at 27,000 x g for 30 min. resulted in 
a sinking precipitate, a floating precipitate, and a straw-colored 

Chloroplasts prepared from spinach leaves were obtained by the 
method of Crane (2). After 2 washings with 0.05 M Tricine, pH 7.4, 
followed by centrifugation at 8,720 x g, the pellet was resuspended in 
buffer which was 4% in Triton X-100 and incubated for 30 min. with 
occasional stirring. The mixture was then centrifuged at 144,000 x g 
for 1 hr. 

table 1. Chemical Composition of Mitochondrial Membranifibril 





Cl b 



Lipid 2 




0.028 0.020 



1 ^moles/g protein. 

2 Determined as % lipid 


mg lipid N 

X 100 

mg- lipid + nig pr 

Cell Biology 


Figure 1. Mitochondrial membrane fragments showing- 90A particles and 
linear arrangement of these particles on the surface of the membrane. 
Phosphotungstate. X 164,000. Marker 500A. 

Figure 2. Membraniflbrils purified by ammonium sulfate fractionation from 
Triton-treated FTP. Uranyl acetate. X 184,600. Marker 500A. 

192 Indiana Academy of Science 

Protein was assayed by the modified biuret procedure of Yonetani 
(25). Phospholipid was extracted using the standard Folch procedure (4) 
with aqueous 0.74% KC1 as the salt. The organic layer was evaporated 
to dryness and the phospholipid taken up in 10 ml of chloroform-methanol, 
2:1. One ml of this was combined with an equal amount of concentrated 
sulfuric acid and heated over a flame until the solution blackened. Then 
2-3 drops of concentrated hydrogen peroxide were added and the mixture 
boiled again. The clarified solution was then tested for phosphate by the 
method of Lindberg and Ernster (11). Cytochromes were determined 
from difference spectra recorded with a Unicam SP 800 recording spec- 
trophotometer by the method of Williams (24). The extinction coefficients 
of Vanneste (23) were used. 

Samples were negatively stained with either 1% phosphotungstate 
at pH 6.9 or saturated uranyl acetate. Microscopy was done on a Philips 
EM 200 using Kodak Electron Image Plates. 

Sonication to release membranifibrils was carried out with a high- 
power sonicator kindly supplied by Quigley-Rochester Scientific Inc. 

Results and Discussion 

Examination of untreated electron transport particles by negative 
staining reveals flattened vesicles from the surface of which project 90A 
particles (Fig. 1). These particles are visible on the membrane surface 
as well as at the edges of the vesicles. In this latter profile view, particles 
can frequently be seen attaching to the membrane edge by a stalk. Closer 
examination of Figure 1 suggests that the white rim to which the 
particles attach at the edge of the vesicle might be a discrete structure 
rather than just a rolled edge. At arrow A this white rim is seen to 
extend out from the membrane surface as a linear structure. The paral- 
lel array of lines on the membrane surface (at arrow B) also suggests 
a linear substructure associated with these membranes. 

Treatment of ETP with Triton X-114 releases linear structures to 
which the 90a particles are attached. Following ammonium sulfate 
fractionation, a purified fraction of these structures was obtained (Fig. 2). 
The central strands have an average diameter of 70 A. The 90a particles 
are arranged opposite one another in a regular fashion along the central 
strand. This linear structure has been called the membranifibril (17). 
Treatment of ETP with deoxycholate and KC1, according to the method of 
Tzagoloff et al. (22), releases linear elements which resemble the mem- 
branifibril. Similar structures have also been observed following 
sonication of whole beef heart mitochondria (Fig. 3). 

Preliminary chemical analysis of the purified Triton-prepared mem- 
branifibrils revealed that they are composed mostly of protein. Hence 
the name fibril is applicable. Several bands were observed on gel electro- 
phoresis. The membranifibrils also contain a small amount of lipid, about 
15%, and very little cytochrome. The preparation shows ATPase activity. 
Initial flavin analysis indicates that there may be an acid-extractable 

Cell Biology 


Figure 3. Sonicated mitochondria showing released membranifibrils. Phos- 
photungstate. X 123,000. Marker 500A. 

Figure 4. Chloroform-methanol extracted membranifibrils. Phospho- 
tungstate. X 213,000. Marker 500A. 


Indiana Academy of Science 


Figure 5. Chloroplast grana membrane showing linear arrangement of 
100A particles on the surface of the membrane. Phosphotungstate. X 136,600. 
Marker 500A. 

Figure 6. Membranifibrils isolated from chloroplasls by Triton treatment. 
Phosphotungstate. X 184,600. Marker 500A. 

Cell Biology 195 

flavin associated with the membraninbril fraction. The chemical 
composition of the mitochondrial membranifibrils is found in Table 1. 

Kopaczyk et al. (10) have isolated a rutamycin-sensitive ATPase 
fraction (P 2 ) by cholate and ammonium sulfate treatment of inner mito- 
chondrial membrane which resembles our membraninbril fraction both 
chemically and structurally. Their P 2 fraction showed no cytochrome aa 3 
and low levels of the b and c cytochromes. The lipid content of their frac- 
tion was reported as 10.7%. The fraction showed minimal rutamycin- 
sensitive ATPase activity in the absence of added phospholipid. Maximal 
activity was achieved by the addition of 1 mg of phospholipid per mg of 
P 2 preparation to the assay medium. 

Membranifibrils are clearly evident in their electron micrographs of 
the rutamycin-sensitive ATPase (P 2 ) fraction. However, they interpret 
these structures as headpiece-stalk sectors projecting exteriorly from 
phospholipid bilayers or coiled phospholipid micelles. A lipid-based struc- 
ture hardly seems likely considering the low lipid content of the P 2 
fraction and of our membranifibril preparation. 

Extraction of the membranifibril with acetone or chloroform-methanol 
leaves a 30a diameter fibrous, insoluble protein (Fig. 4). This fibrous 
protein may be the basis of membranifibril structure. 

When untreated washed and lysed chloroplasts are observed by nega- 
tive staining, the grana membranes are seen to have a particulate surface 
(Fig. 5). These particles have an average diameter of 100a and appear 
to be aligned in rows (indicated by arrow). Treatment of the chloroplast 
fragments with Triton releases fibrous structures very similar to the 
mitochondrial membranifibrils (Fig. 6). These structures are collected in 
the 144,000 x g supernatant and have been called chloroplast membrani- 
fibrils (20). These structures consist of 70a diameter strands to which 
the 100a particles are attached. Chemical composition and enzymatic 
activity of the chloroplast membranifibrils is currently under investiga- 
tion. Structures resembling chloroplast membranifibrils have been seen 
following sonication of intact chloroplasts. 

It is proposed that the membranifibril is the basis of the linear 
arrangement of particles on the surface of mitochondrial cristae and 
chloroplast grana membranes by virtue of the attachment of these 
particles to the fibrillar structure. In mitochondria, such a structure 
would correspond to the basepieces which Green has proposed as the site 
of the electron transfer chain (6). However, since the membranifibril 
contains very little cytochrome it is unlikely that cytochrome dependent 
electron transport function is associated with the site of attachment of 
the 90a particles. 


A new structural element, the membranifibril, was isolated from the 
membranes of mitochondria and chloroplast grana by Triton treatment. 
Preliminary investigation of the chemical composition and the enzymatic 

196 Indiana Academy of Science 

activity of these fibrils was conducted. It was proposed that the mem- 
branifibrils are the basis of the linear organization of the particles on 
these membranes. 

Literature Cited 

1. Branton, D., and R. B. Park. 19 67. Subunits in chloroplast lamellae. 
J. Ultrastruct. Res. 19:283-303. 

2. Crane, F. L. 1959. Internal distribution of coenzyme Q in higher plants. 
Plant Physiol. 34:128-131. 

3. FernAndez-MorAn, H. 1962. Cell membrane ultrastructure. Low temper- 
ature electron microscopy and x-ray diffraction studies of lipoprotein 
components in lamellar systems. Circulation 26:1039-1065. 

4. Folch, J., M. Lees, and G. H. S. Stanley. 1957. A simple method for the 
isolation and purification of total lipids from animal tissues. J. Biol. 
Chem. 226:497-509. 

5. Glaeser, R. M., T. Hayes, H. Mel, and C. Tobias. 1966. Membrane struc- 
ture of Os04-fixed erythrocytes viewed "face on" by electron microscope 
techniques. Exp. Cell Res. 42:467-477. 

6. Green, D. E., and A. Tzagoloff. 1966. The mitochondrial electron transfer 
chain. Arch. Biochem. Biophys. 116:293-304. 

7. Hansen, M., and A. L. Smith. 1964. Studies on the mechanism of oxi- 
dative phosphorylation. Biochim. Biophys. Acta 81:214-222. 

8. Haselkorn, R., FernAndez-MorAn, F. Kieras, and E. van Brugger. 1965. 
Electron microscopic and biochemical characterization of fraction I 
protein. Science 150:1598-1601. 

9. Howell, S. H., and E. N. Moudrianakis. 1967. Function of the quanta- 
some in photosynthesis. Proc. Nat. Acad. Sci. 58:1261-1268. 

10. Kopaczyk, K., J. Assai, D. W. Allman, T. Oda, and D. E. Green. 1968. 
Resolution of the repeating' unit of the inner mitochondrial membrane. 
Arch. Biochem. Biophys. 123:602-621. 

11. Lindberg, O., and L. Ernster. 1956. Determination of organic phos- 
phorous compounds by phosphate analysis, p. 1-22. In: D. Glick, [ed.], 
Methods of Biochemical Analysis. Vol. III. Interscience Publishers, 
New York. 

12. Low, H., and I. Vallin. 1963. Succinate-linked diphosphopyridine nucleo- 
tide reduction in submitochondrial particles. Biochem. Biophys. Acta 

13. Oda, T., and S. Seki. 1966. Molecular organization of the energy trans- 
ducing system in the mitochondrial membrane, p. 369-370. In: R. Uyeda, 
[ed.], Sixth International Congress for Electron Microscopy 1966. Vol. 
II. Maruzen Co., Ltd., Tokyo. 

14. Park, R. B., and N. G. Pon. 1963. Chemical composition and the substruc- 
ture of lamellae isolated from Spinacea oleracea chloroplasts. J. Mol. 
Biol. 6:105-114. 

15. Racker, E., D. D. Tyler, R. W. Estabrook, T. E. Conover, D. F. Parsons, 
and B. Chance. 1965. Correlations between electron transport activity 
ATPase and morphology of submitochondrial particles, p. 1077-1101. In: 
T. E. King, H. S. Mason, and M. Morrison, [ed.l, Oxidases and Related 
Redox Systems. John Wiley and Sons, New York. 

16. Staeiiklin, L. A. 1967. Chloroplast fibrils linking the photosynthetic 
lamellae. Nature 214:1158. 

Cell Biology 197 

17. Stiles, J. D. 1969. Linear substructure in the membranes of mitochondria 
and chloroplasts. Ph.D. Thesis. Purdue University. 104 p. 

18. Stiles, J. W., W. P. Cunningham, F. F. Sun, and F. L. Crane. 19G6. The 
relation of structure to function in the mitochondrial electron transport 
system. J. Cell Biol. 31:113A. 

19. Stiles, J. W., F. F. Sun, and F. L. Crane. 1967. Fibrous structures from 
mitochondrial cristae. J. Cell Biol. 35:129A-130A. 

20. Stiles, J. W., J. T. Wilson, and F. L. Crane. 1968. Membranifibrils in 
cristae and grana. Biochim. Biophys. Acta 162:631-634. 

21. Trown, P. W. 1965. An improved method for the isolation of carboxy- 
dismutase. Biochemistry 4:908-918. 

22. Tzagolofp, A., K. IT. Byington, and D. H. MacLennan. 1968. Studies of 
the mitochondrial adenosine triphosphate system. J. Biol. Chem. 


23. Vanneste, W. H. 1966. Molecular proportion of the fixed cytochrome 
components of the respiratory chain of the Keilin-Hartree particles and 
beef heart mitochondria. Biochim. Biophys. Acta 133:175-178. 

24. Williams, J. N. 1964. A method for the simultaneous quantitative esti- 
mation of cytochromes a, b, ci and c in mitochondria. Arch. Biochem. 
Biophys. 107:537-543. 

25. Yonetani, T. 1961. Studies on cytochrome oxidase. J. Biol. Chem. 


Chairman: L. A. McGrew, Ball State University 
John W. McFarland, DePauw University, was elected chairman for 1969 


X-Ray Diffraction Study of Aqueous Thallium (III) Chloride. Richard 

M. Lawrence, Ball State University. — The radial distribution function 
of 2.5 M aqueous thallium (III) chloride having a mole ratio of chloride 
ion to thallium ion of 4.3 was calculated from x-ray diffraction data. The 
radial distribution function is consistent with the existence of the tetra- 
chlorothallate (III) ion as the dominate ionic species in the solution. 
There is evidence that the shape of the TlCL-ion is that of a distorted 
tetrahedron with each chloride ion being at about 95° from two others 
and about 127° from a third. The distortion in the complex ion is quali- 
tatively attributed to solvent stabilization effects. There is no evidence 
for thallium-ion — water-molecule contact in the proposed model of the 
tetrachlorothallate (III) ion. 

Studies of a Cyanamide Complex of Iron (II). F. J. Holler and R. D. 
Joyner, Ball State University. — A number of possible ways that the 
cyanamide unit could coordinate with a metal are suggested. These pos- 
sibilities include simple ionic and molecular complexes as well as bridge 
structures. A pentacyano-cynamide complex of iron (II) has been 
referred to in analytical studies of cyanamide, but no previous studies 
have been concerned with its actual preparation. Optimum conditions for 
the formation of this complex have now been determined. Additional 
studies have been made concerning the isolation and purification of the 

Phthalocyaninogermanium(II) and Other Phthalocyaninogermanium Com- 
pounds. R. L. Stover and R. D. Joyner, Ball State University. — Reac- 
tions of germanium compounds of the form RiGeCk-n (R = phenyl, 
n -butyl, hydrogen; n = 0, 1, 2, 3, 4) with metal free phthalocyanine, 
sodium phthalocyanine, and diiminoisoindoline are described. Apparently 
phthalocyaninogermanium compounds are not formed from compounds of 
the type R4GeCl4-.11 where n>l. The reaction of phenyl germanium 
hydride with metal-free phthalocyanine to apparently give phthalocy- 
aninogermanium (II) is described. 

A Kinetic Study of the Reaction of Phenyl Isocyanate with Water. 

LeRoy A. McGrew and Stephen C. Robling, Ball State University. — 
The rate of the reaction of phenyl isocyanate with water in acetone solu- 
tion was followed by measuring the rate of carbon dioxide evolution. In 
the presence of excess water the reaction was found to be first order with 
respect to isocyanate concentration. The pseudo-first order rate constant 
was found to be 5.5 X 10 3 sec.- 1 . The reaction was found to be catalyzed 
by tertiary amine bases. Primary amines suppressed the normal decom- 
position by reacting with the isocyanate in a fast step to produce 


200 Indiana Academy of Science 

N,N-diphenylurea. This urea was also observed to be the only product of 
the normal reaction with water. The experimentally determined rate law 
was shown to be consistent with a proposed mechanism. 

Organonitrile Complexes of Rhenium (I) and Manganese (I). Bruce N. 
Storhoff, Ball State University, and J. R. Doyle, the University of 
Iowa. — Bromopentacarbonylrhenium(I) reacts with acetonitrile, propioni- 
trile, acrylonitrile and benzonitrile to yield complexes of the stoichiometry 
Re(CO) 3 (RCN) 2 Br. Bromopentacarbonylmanganese(I) reacts, in an anal- 
agous manner, with acetonitrile and benzonitrile to yield the tricarbonyl 
complexes Mn(CQ) 3 (RCN) 2 Br. The infrared spectra of the complexes 
suggest octahedral structures with the two nitriles cis to each other and 
cis to the bromine. 

Bromopentacarbonylmanganese(I) reacts with acrylonitrile and 
propionitrile to yield complexes of the stoichiometry Mn(CO) 3 (RCN)Br 
which are suggested to be halogen bridged dimers on the basis of molecu- 
lar weight and infrared studies. J9i-/x-bromooctacarbonyldirhenium(I) 
reacts with acetonitrile to yield cis-Re(CO) 4 (CH 3 CN)Br. 

Infrared studies indicate that for each complex the organonitriles 
are attached to the metals through the pair of electrons associated with 
the nitrogen. Molecular weight studies and nuclear magnetic resonance 
studies of the tricarbonyl complexes, M(CO) 3 (CN) 2 Br, indicate that they 
dissociate in chloroform solution but not in dimethylsulfoxide solution. 

Replacement reactions indicate that the nitriles of all the manga- 
nese (I) and rhenium (I) complexes can be replaced by pyridine and sug- 
gest that olefins do not replace the nitriles. 

Use of Computers in Undergraduate Physical Chemistry. Richard F. 
Copeland, Ball State University. — Computers can be very valuable tools 
for the student of physical chemistry, but for most students the junior or 
senior year course in undergraduate physical chemistry is not the time to 
begin to learn computer programming. The best results should be 
obtained by a program integrating the use of computers into chemistry 
courses beginning with the freshman course in general chemistry. 
Instruction at the freshman level may be in either a special course, or, 
preferably, incorporated into the general chemistry laboratory sequence. 
Analysis of data using library programs is a very useful asset, but 
emphasis should be on the student's use of programming to solve his 
individual problems. Simulation of complex chemical systems involving 
large numbers of variables can be a valuable tool in instruction. 

Synthesis and Properties of [2.2.2] Bicyclooctyl Enamines. A. G. Cook 
and T. A. Hecht, Valparaiso University. — Some [2.2.2]bicyclooctyl 
enamines have been synthesized and some of their properties investi- 
gated. The ultraviolet spectra of some of them seems to be indicative 
of some homoconjugation. 

Reduction of Selected Enamines with Lithium Aluminum Hydride. A. G. 

Cook and D. J. Schultz, Valparaiso University. — It has been observed 
that certain enamines possess structural features which make them sus- 
ceptible to reduction by lithium aluminum hydride. These features will 
be discussed. 


Chairman: William B. Crankshaw, Ball State University 
Thomas McComish, Ball State University, was elected chairman for 



Temperature and Moisture Relationships of Green County, Indiana Strip 
Mine Areas. R. Michael Dinkel and Fred Rothwell, Indiana State 
University. — Temperature and moisture are physical factors which must 
be taken into consideration when one considers the reclamation of strip- 
mined land by means of forestation. During the spring and summer of 
1968, temperature and moisture readings were made on and in the 
vicinity of the Green County spoil bank location. These were measured 
periodically and data was compiled in order to determine the significance 
of varying temperature and moisture relationships on the viability and 
growth of tree seedlings introduced on the spoil bank and coal-mine 

Ecological Site Preference and Taxonomic Differences within Two Acer 
saccharum-Acer nigrum Complexes Found in Parke County, Indiana. 

Alfred G. Craske, Jr., Indiana State University. — A study was con- 
ducted on the ecological site preferences and taxonomic differences 
within two Acer saccharum-Acer nigrum complexes found in Parke 
County, Indiana. A suitable means of identification was determined for 
the two species based on leaf characteristics. A significant difference was 
found between flood plain and slope and flood plain and upland popula- 
tions based on leaf morphology. The flood plain was found to have sig- 
nificantly higher pH and phosphorus values and significantly less canopy 
cover than those values for slope or upland. No significant differences 
were found between slope and upland for the ecological parameters. 
Sugar maple, black maple and the intermediates were found to represent 
three populations. The intermediate population represented the central 
portion of a continuum of leaf characters running from black maple to 
sugar maple. Acer nigrum Michx. f. showed a definite site preference 
for the flood plain. Acer saccharum Marsh, showed a definite site prefer- 
ence for the slope and upland. It would thus appear that the character 
of site preference would be valuable for identification and should be 
incorporated into discriptions of the two species. 

Effects of Thermal Discharge on the Phytoplankton and Macroinverte- 
brates of the Wabash River. Jerry L. Gerwig and Wm. B. Crankshaw, 
Ball State University. — The effects of thermal discharge on the phyto- 
plankton and macroinvertebrates were studied on the Wabash River in 
Sullivan County, Indiana. The temperatures recorded in the normal river 
water ranged between 72 and 78 degrees F. Temperatures recorded in 
the heated-water zone varied from 87 to 74 degrees F. 


202 Indiana Academy of Science 

Nineteen genera and nine families of green algae (Chlorophyceae), 
blue-green algae (Myxophyceae), and diatoms (Bacillariophyceae) con- 
stituted the phytoplankton population. Green algae represented 95 to 97 
percent of the algal population, blue-green algae represented 2 to 3 per- 
cent, and diatoms represented 1 to 2 percent regardless of the station or 
the average number of organisms per liter of water. The invertebrates 
collected were mainly various insect larvae and nymphs. 

Effects of Effluent on the Fish Population of Mill Creek, Rochester, Indi- 
ana. Frederic Morgan, Ball State University. — This is a report of pre- 
liminary data showing the effect of city and light industrial effluent on 
fish life. Mill Creek is located in North-Central Indiana. It begins as 
the overflow of Lake Manitou and runs approximately five miles to the 
Tippecanoe River, passing through the city of Rochester. 

Fish were collected from eleven stations: four sites above the major 
sources of effluent, three sites scattered through them, and four sites 
below them. The fish were collected by blocking off a measured segment 
(60 m) of the stream with quarter-inch knotted minnow seines and 
making two sweeps with an electric seine. At the eleven sites, production 
of fish varied from a high of 41.1 g/m 2 at station number four above the 
sources of effluent to zero at station number seven. Downstream in the 
last four stations the fish population slowly started to recover: .03 
g/m 2 , .05 g/m 2 , .8 g/m 2 , and 3.1 g/m 2 . From all sites we removed 1166 
fish representing 31 different species. The number of species per site 
varied from nineteen at station four above the sources of effluent to zero 
at station seven. A recovery was noted in the four downstream sites: 
3, 5, 7 and 11. 

During the past twelve months, water samples were taken at four 
sites and analyzed for D.O., temperature, hydrogen sulfide, hardness, 
turbidity, anionic detergent, total phosphate, nitrite, nitrate, ammonia 
and pH. 

A Study of Selected Physiochemical Properties of Two Indiana Main- 
Stream Reservoirs. Paul T. McKelvey and Charles E. Smith, Jr., Ball 
State University. — Selected physiochemical properties of Lake Shafer and 
Lake Freeman, two shallow, narrow main-stream reservoirs near Monti- 
cello, Indiana, were determined from November, 1967, through June, 
1968. Bi-monthly sub-surface samples from eight stations, four on each 
reservoir, were analyzed. 

These reservoirs were found to have certain properties resembling 
those of rivers, e.g., width, depth, carbon dioxide concentration, and the 
absence of distinct thermal stratification. Other properties, typical of 
natural lakes and/or large reservoirs were, e.g., total alkalinity, dissolved 
oxygen, nitrate, pH, and sulfate. Phosphate, nitrite, and turbidity values 
were intermediate between those of rivers and natural lakes and/or large 
reservoirs. Total alkalinity, nitrate, and phosphate cycles were also 

Ecology 203 

Lake Shafer and Lake Freeman were considered eutrophic based on 
the high nutrient levels, especially nitrate and phosphate, present in 
their waters. 

Pattern of Mesic Forest Succession at the Western Border. R. O. Petty, 

Wabash College. — Data and analysis are presented for forest stands 
which depict the successional trends currently underway at the western 
extension of the mesophytic forest. Emphasis is given to the performance 
of several significant species, especially Quercus velutina, Acer saccharum 
and Fagus grandifolia. A model of successional pattern is offered to 
explain the several cliseral shifts which have occurred during the post- 
glacial period, as seen in palynology and in the phytosociology of west- 
central Indiana. 

The Effect of Inorganic Sediment on Macroinvertebrate and Fish Popula- 
tions of a Central Indiana Stream. James R. Gammon, DePauw Univer- 
sity. — The composition and density of populations of macroinvertebrates 
and fish were studied during 1967 and 1968 in a 1.5 kilometer section of 
Deer Creek which received wastewater from a crushed limestone quarry. 
Heavy deposits of sediment which accumulated during 1966 were swept 
downstream by floods during late 1966 and early 1967. Little sediment 
accumulated in pools below the quarry during 1967 because of unusually 
high levels of stream flow throughout the year and because the quarry 
settling ponds were dredged regularly. Nevertheless, 35 tons of 
sediment entered the stream from the quarry during 1967. 

The standing crop of fish in three pools downstream from the quarry 
averaged only 55% of those in two pools upstream in June 1967 (224 
Kg. /ha. compared to 417 Kg. /ha.). Carpsuckers and gizzard shad were 
almost completely lacking downstream, while carp, redhorse and suckers 
were reduced in numbers. By early August many small individuals had 
migrated into the pools below the quarry and the standing crops rose to 
an average of 255 Kg./ha. compared to 362 Kg. /ha. in the upstream pools. 

The quarry settling ponds received relatively little attention follow- 
ing a thorough dredging in early August 1967 and became completely 
filled and, therefore, inoperative by late fall of 1967. Heavy concentra- 
tions of stonedust entered Deer Creek throughout 1968 as a result. By 
August 2, 1968 over 3300 tons of sediment were contributed to Deer 
Creek. The standing crop of fish in June 1968 in the pools below averaged 
only 25% of those above (100 Kg./ha. compared to 400 Kg./ha.). 

The macroinvertebrate populations in several riffles above and 
below the point of pollution were sampled monthly during months of 
stable water levels. The density in the riffles below averaged less than 
50% of that above. All orders, families, genera and species seemed to be 
reduced equally in abundance. 

The Effect of Ground Cover on the Soil Moisture 
Regime in a Mixed Mesophytic Woods 

William B. Crankshaw, Ball State University 


A 25-acre mixed mesophytic woods in southwestern Delaware County 
was used to study the effect on soil moisture regime by presence or absence 
of ground cover. Four sites were selected within the woods, a Brookston 
silty clay loam with cover, a Brookston silty clay loam without cover, a 
Crosby silty clay loam with cover and a Crosby silty clay loam without 
cover. Bouyoucos soil moisture blocks were permanently installed at 
5, 10, 25, and 50 cm. depths at each site. Available soil moisture was moni- 
tored daily for a period of thirteen months. Significant differences were 
found in rate and amount of loss and gain of available moisture content at 
the different depths for both soil types and for both cover situations. 


A 25-acre mixed mesophytic woods in southwestern Delaware County 
was selected to study the effect of presence or absence of ground cover 
on soil moisture. Stewart Woods, the area chosen, met several predeter- 
mined requisites. These requisites included: a woods bisected by a fence, 
one side undisturbed, the other side lightly grazed and also at least two 
soil types, the latter oriented perpendicular to the fence line. These 
simple specifications quickly excluded most of the wooded areas in east 
central Indiana. 


Within the woods, four study sites were established. These sites 
were located on a Brookston silty clay loam with ground cover, a 
Brookston silty clay loam without cover, a Crosby silty clay loam with 
cover and a Crosby without cover. Soil moisture and soil temperature 
probes were permanently installed, with a minimum of disturbance to 
the soil structure and profile, at 5, 10, 25, and 50 cm. depths at each of 
the four sites. Taylor maximum-minimum thermometers and rain gages 
were mounted on posts one meter above the surface at each site. Air and 
soil temperatures, precipitation and available soil moisture were 
monitored daily for thirteen months. 

A complete census was made of the 24.25-acre woods by species and 
diameter. The phytosociological parameters of relative and absolute 
density and basal area were determined as well as importance value 
(Table 1). This was done to compare the grazed and ungrazed areas on 
the basis of relative importance of represented species and to evaluate 
the difference exerted by a 12-year period of light grazing (one calf per 4 

Sampling of ground cover was accomplished by establishing four 
one-meter square plots near each site. These plots were located 10 meters 





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Ecology 207 

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208 Indiana Academy of Science 

in each cardinal direction from the instrument post. All shrubs, tree 
reproduction and herbaceous vegetation were tallied by species. 

Results and Discussion 

Little significant difference was noted in a comparison of importance 
values between the grazed and ungrazed area. The grazed portion of the 
woods appeared to be more xeric with combined importance value for 
oaks and hickories of 53.1% as opposed to 37.8% for the ungrazed. Note 
that the importance for the ungrazed portion is distributed fairly evenly 
between five species, all rather mesic in their requirements. 

Ground cover was very sparse on the grazed side even with the 
minimal grazing. No shrubs were tallied and very few forbs; 82.5% 
of all individuals in the grazed area were sedges and grasses. 

An evaluation of the soil moisture data indicated that the two sites 
with relatively little ground cover reached field capacity earlier than 
sites with cover (Table 2). This difference increased with depth, i.e., the 
50 cm. depth showed the greatest difference in moisture content between 
cover and non-cover. The 25 and 50 cm. depths, once they achieved field 
capacity, remained consistently higher in available moisture content than 
the shallower depths not only during the growing season, but all year. 
Fluctuations in available moisture at the 5 and 10 cm. depths were very 
wide during the summer as an effect of convectional rain and higher 
evaporation rates. The moisture regime did not actually stabilize at any 
depth until mid-March; at this time field capacity was constant and 
remained so until June and July. 

Differences in soil moisture existed between the two soil types as 
well as in the presence or absence of cover. Available moisture content 
began decreasing sooner in the Crosby than in the Brookston and 
dropped to 15% available moisture content at an earlier date. The differ- 
ence in time required to reach 15% available moisture was less marked 
between the Crosby and Brookstone as the soil depth increased. Fall 
rains caused a sharp and steady increase in available moisture content 
at the 50 cm. level. This occurred about two weeks earlier in the Crosby 
than in the Brookstone. Other depths in both soils evidenced an increase 
but with the increase occurring at the same time at each depth and at 
a less steady rate than 50 cm. Though the penetration was better on the 
Crosby, the Brookston had in general a higher moisture holding capacity 
through the year. 

In summary, soil moisture reached field capacity earlier in the 
grazed portion of the woods on the Crosby soil, though penetration of 
moisture was greater in the ungrazed. Moisture retention was greater 
at all four depths on the grazed side, particularly on the Brookston soil. 
Apparently, grazing, if minimal to avoid compaction, has a favorable 
effect on the soil moisture regime. 

Ecology 209 

Literature Cited 

1. Crankshaw, Wm. B., Qadir, S. A., and A. A. Lindsey. 1965. Edaphic Con- 
trols of Tree Species in Presettlement Indiana. Ecology 46(5) :688-698. 

2. Dambach, C. A. 1944. A ten-year ecological study of adjoined grazed and 
ungrazed woodlands in NE Ohio. Ecol. Mon. 14:253-270. 

3. Diller, O. D. 1937. Soil moisture content during critical periods in the 
regeneration of previously grazed farm woodlands. J. Forestry 35:299- 

4. Geiger, R. 1966. The Climate Near the Ground. Harvard University 
Press, Cambridge, Mass. 

5. Jeffrey, W. W. 1963 Soil temperature measurements in forests of north- 
western Canada. Ecology 44:151. 

6. Olson, D. F., and M. D. Hoover. 1954. Methods of soil moisture determina- 
tion under field conditions. U.S.F.S., S.E. For. Exp. Sta. Paper 38. 

Detailed Studies of Old-Growth Forests in Versailles State 
Park, Indiana 

M. T. Jackson and P. R. Allen, Indiana State University, Terre Haute, 
and Columbus High School, Columbus, Indiana 


Full censuses were taken for 17.46 and 6.44 acre segments of two 
old-growth upland forests located in Versailles State Park, Ripley County, 
Indiana. The stands have been designated as Jackson's and Potzger's 
Woods, respectively. Transects were also taken in the slope forests of 
Jackson's Woods and the forest composition of limestone sinks was 

Jackson's Woods had 28 species over 4 inches and 33 species over 2 
inches dbh. Density per acre of stems over 4 inches was 109 with basal 
area per acre of 112 ft. 2 Beech and sugar maple ranked first and second 
in importance value with 32 and 25%, respectively. Tulip-poplar followed 
with 14%, and dogwood was fourth at 6.5%. In the sinkholes, beech and 
sugar maple decreased in importance, and walnut, elm and tulip-poplar 
increased in importance in larger and better drained sinks. The W-facing 
slope forest is mixed mesophytic; oak and hickory increases on exces- 
sively-drained NW and N-facing slopes; the NE and E-facing slopes are 
watered by subsurface seeps and have forests similar to floodplains. 

Potzger's Woods had 24 and 27 species over 4 and 2 inches, respec- 
tively. The density per acre was 121 with a basal area per acre of 117 ft. 2 
Sugar maple and beech ranked first and second in importance with 39 and 
31%, respectively; walnut was third at 5% followed by white ash and 
black gum. Sugar maple was exceedingly dominant in the smaller size 
classes in both stands. The present full tally for Potzger's Woods indicates 
that sugar maple has increased in stand importance since 1956, most 
likely at the expense of beech which has declined slightly in importance. 

The stands appear to be transitional between the mixed mesophytic 
forests of extreme southeastern Indiana and the beech-maple types of 
the flat uplands. As the physiography matures the woods will probably 
shift to a more mixed mesophytic composition. 


Plant ecologists have been interested in the forests of southeastern 
Indiana for a number of years. The Versailles State Park and Laughery 
Creek Valley areas of Ripley County have been of particular interest 
because the diverse topography of that region induces striking differ- 
ences in forest types within very short distances. According to Potzger 
(9), the well-drained uplands and moderate slopes support a climax of 
mixed mesophytic forest dominated by beech (Fagus grandifolia) and 
sugar maple {Acer saccharwn) , and which includes twenty or more addi- 
tional species of lesser importance. Beech is also an important member 
of the forests on the poorly-drained flat uplands, but sugar maple is 
replaced as a co-dominant on these wet sites by red maple {Acer rubrum) 
and sweet gum (Liquidambar styraciflua) . On the excessively-drained 
slopes beech is absent; however, sugar maple continues as a dominant 
species, along with mesophytic oaks, ash and hickories (9). 


Ecology 211 

The faculty and students of the Conservation Education Summer 
Camp, held at Versailles State Park for several years, focused their at- 
tention on this wide range of habitats and forest composition in the vicin- 
ity of the Laughery Creek Valley. Potzger (9) and Stearns (15) studied 
the ecological relationships of the forest communities within the State 
Park area. In addition, Potzger and Potzger (13) and Potzger and Lim- 
ing (12) worked out secondary succession patterns on the flat uplands 
of Ripley County. The distribution patterns and associated species of 
beech were examined by Potzger and Chandler (10) in ten stands in 
the general Laughery Valley area. In a later study, Potzger and Chan- 
dler (11) described four oak-dominated forests of the same region. 

The aims of this study were: 1) to intensively study one of the best 
remaining old-growth upland and adjacent slope forests of the area that 
somehow had escaped earlier study; 2) to compare the above stand to 
a frequently studied old-growth stand located in the southeast part of 
the Park; and 3) to supplement the current search for and description of 
areas in Indiana that are suitable for preservation under the Natural 
Areas Act. 

Location and Description of the Stands 

Versailles State Park lies in the Illinoian Till Plain section of Indiana. 
Upland areas are flat to gently rolling, but Laughery Creek and its 
tributaries have cut deep valleys and ravines into the upland plateaus. 
There is about 300 ft. of relief in the immediate area with Laughery 
Creek dropping to about 700 ft above mean sea level and the highest flat 
uplands reaching slightly above 1,000 ft elevation. Laughery Creek cut 
a deep, broad steep-walled valley into the underlying Ordovician lime- 
stone as it carried Pleistocene meltwater. The region is very scenic with 
broad panoramas. 

Laughery Creek is the approximate dividing line between the Dear- 
born Upland Physiographic Provine characterized by Ordovician Age 
limestone and the Muscatatuck Regional Slope underlain by younger 
Silurian Limestone. One of the study sites is situated in each physio- 
graphic province. Both stands occupy well-drained Cincinnati silt loam 
soils. The study areas are located in the infrequently visited section of 
the Park that lies south of U. S. Highway 50. 

The first stand, referred to as Jackson Woods, is located west of 
Laughery Creek on a high trapezoid-shaped upland promentory. (The 
stand names used in this paper follow the nomenclature used by Lindsey 
[6]). It is situated in the northwest corner of section 18, Twp. 7N, Range 
12 E. There is about 100 acres of forest in the immediate area, but the 
old-growth upland stand covers less than 25 acres. The west, north, 
northeast, and east-facing slopes of the upland ridge are also clothed in 
good mixed forest. The upland area slopes gently toward the northeast, 
but drainage is largely internal into numerous sinkholes which then 
feed springs and seeps along the valley walls. The slopes that are 
watered by sub-surface seeps support a forest that is strikingly floodplain 
in character, although it occurs on a 75-80% slope. 

212 Indiana Academy of Science 

The second stand is referred to as Potzger Woods in honor of the 
late, outstanding plant ecologist from Butler University. It is located 
on a gentle northeast-facing upland slope, which is drained by small 
ravines in the upper reaches of Turkey Creek, a tributary of Laughery 
Creek. The stand is situated east of Laughery Creek in the northeast 
quarter of Section 20, Twp. 7 N, Range 12 E. The least disturbed part of 
the stand covers less than 10 acres. 

Both areas suffered some disturbance as a consequence of private 
ownership prior to park acquisition. Jackson Woods was lightly grazed 
by cattle for a number of years, and about 30 trees were cut in 1936. 
For about seven years prior to 1936, a few beech trees were cut each 
year for firewood. Little information exists concerning the history of 
Potzger Woods, but apparently only limited cutting occurred in the 
1920's and 30's. The presence of abundant high grade walnut in both 
stands indicates that neither area was seriously damaged by logging. 


A full census of all trees greater than 2.0 inches dbh (diameter 
breast high) was taken for the best sections of each stand. A total of 
17.46 acres was tallied in Jackson Woods and 6.44 acres in Potzger 
Woods. Stand margins were excluded from the full tallies to avoid com- 
munity transitions. All trees over 4.0 inches were measured to the near- 
est 0.1 inch with diameter tapes, and trees 2.0 to 3.9 inches were counted 
and recorded by species. Smaller trees, tree seedlings, shrubs and herbs 
were stratum ranked according to the method advanced by Lindsey, 
et al. (5) to establish the importance of each species within each stratum. 
Stratum ranking is a subjective assignment of numerical values to each 
species according to its contribution to the density and cover in its re- 
spective stratum. Values range from stratum rank 9 (a pure stand of 
a single species) to stratum rank 1 (a species with a single individual 

In addition, the changes in forest composition on contrasting slope 
aspects of Jackson Woods were examined by laying a continuous one- 
half mile horizontal belt transect along the contour at mid-slope. Two 
600-ft vertical belt transects were run from the upland to the flood- 
plain to determine the effect of topographic position on forest composi- 
tion. The transects were divided into sections 43.56 ft by 100 ft so that 
a 1/10 acre sample was obtained per 100-ft horizontal distance. 

Six of the upland sinkholes in Jackson Woods were full tallied for 
trees to compare: 1) small sinks (ca. 0.15 acre each) that are subject 
to flooding during heavy rains; 2) small sinks (ca. 0.15 acre each) that 
apparently seldom or never contain standing water; and 3) large sinks 
(ca. 0.4 acre each) that apparently do not flood. The occurrence of 
ponding in small sinks was established by examination of the silt coat- 
ings on the leaf litter and by the absence of herbs, shrubs and tree re- 
production in the flooding portion. 

Ecology 213 

The soil reaction of upland and slope samples was determined with 
a Beckman pH meter; soil moisture values were expressed as percentage 
oven dry weight. Canopy heights were measured with an Abney level. 

The field work was done in early July and mid-September. Full tally 
plot corners were marked with permanent iron rods and exact locations 
are available from the authors. 

Stand attributes according to Lindsey (4) are as follows: D 2 = 
density per acre; D 3 = relative density; B 2 = basal area per acre; B 3 = 
relative basal area; and V 3 = importance value = 

D 3 + B 3 . 

Species nomenclature follows Little (7) and Fernald (3) for trees and 
other species, respectively. 

Results and Discussion 

Jackson Woods 

Upland Stand Description. The primary stand attributes of the 
full-tally section of this stand are shown in Table 1. Twenty-eight species 
greater than 4 inches dbh represent a stand density of 109.03 stems 
per acre and a stand basal area of 111.67 square ft per acre. Although 
these totals are slightly less impressive than those for the best old- 
growth stands in the state, they indicate a relatively undisturbed condi- 
tion. The stand is of mixed composition, but heavily dominated by beech 
and sugar maple, which total 57.5% of stand importance value. 
Beech represents nearly a third of stand importance, largely because of 
its large size (over half of the stand basal area); whereas, sugar maple 
attains its one-fourth of the stand importance because of its high density 
(nearly 40% of the stand total), particularly in the small size classes. 

Examination of the size-class distribution in Table 2 gives the im- 
pression that sugar maple is rapidly invading the stand, in large measure 
at the expense of beech. This is particularly noteworthy when the 
ratios of the number of stems less than 12 inches dbh to the number 
greater than 12 inches are examined for the two species. These ratios 
are 1675/40 or 41.88 and 26/242 or 0.11 for sugar maple and beech, 
respectively, when stems greater than 2 inches dbh are considered, and 
699/40 or 17.48 and 8/242 or 0.03, respectively, for stems greater than 
4 inches dbh. These ratios obviously are useful only for determining 
possible trends in stand composition since many beech-maple dominated 
stands exhibit a similar preponderance of young maples. However, beech 
usually maintains its position as a co-dominant and frequently increases 
its importance value as the stand matures. This natural process of selec- 
tion toward beech dominance as successional maturity is reached is 
frequently enhanced by selective logging in which higher grade species 
such as oak, walnut and tulip-poplar are removed, leaving the lower 
quality beech. The differential rate of sugar maple reproduction when 

214 Indiana Academy of Science 

compared to beech is largely offset by differential survival of beech 
with respect to sugar maple. Most undisturbed stands seem to have 
much greater mortality of small maples than of small beech. Long-term 
detailed studies of a number of beech-maple dominated stands should 
help resolve this question. Notwithstanding these comments, the ab- 
sence of any sugar maple stems greater than 24 inches in this stand 
indicates that sugar maple will undoubtedly increase in the larger size 
classes in the future, assuming the continued absence of disturbance. 
The projected increase in sugar maple may be partially at the expense 
of beech, but a replacement of less tolerant species such as tulip-poplar 
(Liriodendron tulipif era) , walnut (Juglans nigra), sassafras (Sassafras 
albidum) and wild cherry (Primus serotina) is more probable. 

Tulip-poplar is the sub-dominant species with an importance value 
of 14.1%. Moderate selective cutting 30 to 40 years ago has apparently 
favored this relatively intolerant species. Although most of the 30 trees 
cut in 1936 (just prior to park acquisition) were tulip-poplar, the 
crown openings created by this cutting undoubtedly favored release, 
rapid growth and regeneration of tulip-poplar, serving to increase 
rather than decrease tulip importance. Nearly half (96 of 204) of the 
tulip-poplar stems over 4 inches dbh fall in the 12 to 24 inch size 
classes. Expected growth increments of thrifty tulip-poplars in canopy 
openings would be on the order of one-fourth to one-half inch diameter 
increase per year on the average. This growth rate would date the estab- 
lishment of many of the middle-sized tulip-poplars as canopy trees at 
about 30 to 40 years ago. 

Other species worthy of comment include flowering dogwood (Cornus 
florida) which represents 6.5% of the stand (Table 1), largely by virtue 
of its high density (13 stems per acre larger than 4 inches dbh), and 
walnut at 4.6% importance. Walnut is very well represented in the 12 
to 24 inch sizes (49 of 66 total stems), indicating that it was also favored 
by the canopy openings created by cutting. Walnut is frequently clumped 
in the larger sinkholes (see Table 3) of the area. This contagious dis- 
persal is favored by increased light, gravity seed dispersal and accumula- 
tion of fertile soil in the sink bottoms. Elm, wild cherry and sassafras 
were probably favored to a lesser extent by the earlier cutting. Six 
species of oak and hickory represent only 1.17% of the stand importance 
value. The absence of these species on this well-drained site is a matter 
of conjecture, since there is no record of recent cutting of either oak or 
hickory. Apparently, even the moderately tolerant white oak is unable 
to successfully compete in mature upland stands in this area. Canopy 
opening would not have favored oak and hickory, because of their slower 
growth rate as compared to tulip-poplar. Large chinquapin oak 
(Quercus muehlenbergii) is rather common along the drier bluffs and 
slopes, where, as Reynolds and Potzger (14) pointed out, it even sur- 
passes white oak in abundance. On the uplands it is an inconsequential 
species, however. 

Although there are no trees greater than 40 inches dbh, there are 
128 stems in the 28-inch or greater size classes for an average of nearly 



TABLE. 1. Stand Attributes for Jackson Woods, based on trees U" dbh and greater 

17.4-6 acres — full tally. 



D 3 

B 2 

B 3 


B A /Tree 

Fagus grandifolia 







Acer saccharum 







Lirio dendron t u lipifera 







Comus florida 







Ulmus rubra 







Juglans nigra 







Fraxinus americana 







P?'unus serotina, 







Nyssa sylvatica 







Ulmus americana 







Sassafras albidum 







Celtis occidentalis 







Quercus alba 







Quercus rubra 







Ostrya virginiana 







Gary a glabra 







Vitis spp. 







Juglans cinerea 







Acer negundo 







Fraxinus quadrangulata, 







Tilia americana 







Carya ovata 







Robinia pseudoacacia 







Carpinus caroliniana 







Quercus muehlenhergii 







Carya cordifoi mis 







Asimina triloba 







Aesculus glabra 










8 large trees per acre. All but 15 (88.3%) of these large trees are 
beech which contributes to the impression that this stand is almost en- 
tirely beech. Potzger and Chandler (10) recorded only 17 trees greater 
than 30 inches (based on a total sample of about 5 acres) in the entire 
ten stands that they studied. The largest individuals of noteworthy 
species in the immediate vicinity of this old-growth stand include: beech 
39.7, chinquapin oak 39. 1, tulip-poplar 36.7, red oak (Quercus rubra) 35.2, 
blue ash (Fraxinus quadrangulata) 29.8, black gum (Nyssa sylvatica) 
29.7, walnut 26.7, pignut hickory (Carya glabra) 24.0, Kentucky coffee- 
tree (Gymnocladus dioica) 24.0, box-elder (Acer negundo) 20.4, butter- 
nut (Juglans cinerea) 17.1, ironwood (Ostrya virginiana) 13.9, and 
dogwood 9.0. Many of the larger forest-grown tulip trees, beech, walnut 


Indiana Academy of Science 


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218 Indiana Academy of Science 

and ash have clear boles of 30 to 60 ft with very little taper. The general 
canopy height averages about 115 to 120 ft in the upland stand, with 
tulip-poplars reaching the greatest overall height. 

The tree reproduction data are summarized in Table 2, so stratum 
rank values are not listed for small trees. In all, 27 species contribute 
1,569 stems in the 2.0 to 4.0 inch dbh size class. 

The shrub and vine stratum is dominated by hinder a Benzoin (stra- 
tum rank 8), particularly in the larger sinks and other very mesic 
situations. Asimina triloba is stratum rank 5; Rhus radicans and Smilax 
rotundifolia, 4; Parthenocissus quinquefolia and Rites cynosbati, 3; 
Sambucus canadensis and Viburnum acerifolium are minor shrubs. 

Twenty-two species of summer herbs were stratum-ranked as follows : 
Galium concinnum and Podophyllum peltatum, 7; Arisacma triphyllum 
and Galium circaezans, 5; Actaea alba, Boehmeria cylindrica, Circaea 
latifolia, Viola sp., 4; Impatiens pallida, Polygonatum pubescens, and 
Sanguinaria canadensis, 3; Cimicifuga racemosa, Dryopteris hexagonop- 
tera, Hydrastis canadensis, Jeffersonia diphylla, Menispermum canadense, 
Panax quinquef olius , Phytolacca americana, Trillium gleasoni, 2; 
Osmorhiza claytoni and Hydrophyllum appendiculatum, 1. The only 
herbs present with marked affinities to southern forests are Cimicifuga 
and Jeffersonia. The fern flora is very depauperate for a rich, mesic 

Sinkhole Comparison. The upland stand has Karst topography that 
typically develops as a result of surface collapse into solution chambers 
in the underlying limestone. There are about two dozen sinks of various 
sizes in the 17% -acre stand. Most range from 8 to 15 feet deep, and vary 
in bottom configuration and subsurface drainage. They may be grouped 
into three broad categories as defined in the Methods Section. Some of 
the larger sinks have almost impenetrable tangles of grapevine, Virginia 
creeper and spice bush thoroughly interlaced with a rank growth of 
jewel weed that commonly grows shoulder high. 

The total area tallied in the six sinks was lVz acres or about 8% of 
the entire stand. The importance values for species present in the three 
types of sinks are summarized in Table 3. The six species present in 
the small sinks subject to ponding were confined to the rims above the 
level of flooding, except for an occasional large beech in the bottom. It 
is of interest that tulip-poplar, walnut and slippery elm (Ulmus rubra) 
are absent from the small, ponding sinks, although they represent nearly 
V± of the importance value of the total stand (Table 1). The periodic 
ponding may prevent seed germination near the sink bottoms, particularly 
the largely gravity dispersed walnut. Collective importance values for 
beech and sugar maple are reduced along the sinkhole gradient from 
79.0% to 68.8% to 58.5% (Table 3), although they are clearly the domi- 
nant species in all three cases. This decrease is compensated for by an 
increase in walnut, slippery elm, American elm {Uhnus americana) and 
tulip-poplar. The fertile soil and mesic conditions of the large sink 
bottoms provide a situation similar to high floodplains, where these 

Ecology 219 

species typically reach their best development. Tree reproduction, shrub 
and herb strata are well represented in all of the non-ponding sinks, 
but are almost totally excluded from the ponded sections. Consideration 
of sinkhole microenvironments is important because they add variety 
to an otherwise uniform area, and correspondingly enrich the plant 
communities present. 

table 3. Comparison of Average Importance Values for Woody Species 
Located in Three Contrasting Types of Sinkholes. 












Ostrya virginiana 


Prunus serotina 



Cornus florida 




Fraxinus americana 




Acer saccharum 




Fagus grandifolia 




Juglans nigra 



Ulmus rubra 



Ulmus americana 



Liriodendron tulipifera 




Celtis occidentalis 




Slope Forests. The west, northwest and northeast-facing slope 
aspects represent 600 feet or 0.6 acre each of the 2,600-foot belt transect, 
and the north and east-facing slope aspects represent 400 feet or 0.4 
acre each. No south slope was present in that immediate area. Data for 
basal area per acre and importance values are summarized in Table 4. 
The slope both lengthens (vertically) and steepens from the west side 
around to the east. The richness of the woody flora decreases and soil 
moisture increases from west to east. The soil pH remains moderately 
acid until the steeper northeast and east-facing slopes with numerous 
limestone outcroppings are encountered, then it becomes slightly basic. 
On the latter slopes, there are abundant mid-slope seepage outlets from 
the internally-drained upland sinks. Since soil moisture readings were 
taken in July following a long dry period, none of the areas had high 
moisture values. 

Species shifts follow, in general, the drouth susceptibility of the 
slopes (Table 4). Trees more typical of more xeric slopes, such as black 
locust {Robinia pseudoacacia) (probably an escape), Eastern red cedar 
(Juniperus virginiana), blue beech (Carpinus caroliniana) , redbud 
{Cercis canadensis) and dogwood are restricted to the west and north- 
west-facing slopes. Oaks, hickories and beech are absent from the mesic 
northeast and east-facing slopes, except for one bitternut hickory (Carya 


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222 Indiana Academy of Science 

cordif ormis) . Both basswood (Tilia americana) and walnut increase pro- 
gressively from drier to mesic slopes. Sugar maple is well represented 
on all slopes although fluctuating widely in both basal area per acre and 
importance value. Also of interest is the addition of species more typical 
of floodplains (American elm, hackberry (Celtis occidentalis) , green 
ash (Fraxinus pennsylvanica) and box-elder) as the mesic slopes with 
subsurface seeps were reached. Kentucky coffeetree, sycamore (Platanus 
occidentalis) and honey locust (Gleditsia triacanthos) were present on 
the east slope, although they did not fall within the transect. 

The species-substrate relationships described above serve to illus- 
trate quantitatively how community types of the Laughery Valley change 
progressively with the compass in this diverse topography. The richest 
flora occurs on the west slope community, which, according to Braun (1), 
is more typically mixed mesophytic than is either the upland or the moist 
east and northeast-facing slopes. The latter are undoubtedly late serai 
stages, rather than mature stands, held there in this case by physio- 
graphic instability and excess subsurface moisture. 

The results from two vertical belt transects down the east slope 
from the upland to the beginning of the Laughery floodplain are com- 
bined and summarized in Table 5. The linear 100-foot increments repre- 
sent horizontal distances since slope corrections were made. The combined 
transect represents 0.2 acre per 100 feet increment. Sections 1 and 2 
are on the relatively level upland; section 3 represents the abrupt break 
at the bluff; sections 4 and 5 the steep (75-80°) mid-portion of the slope, 
and section 6 the less steep beginning of the floodplain. (Slope percent- 
ages are given for only sections 4 and 5 because the slope percentages 
change continually in the other sections.) 

The stand attributes of the upland sections were not appreciably 
different from those of the upland stand in general (Table 1), except 
that the relatively small transect sample missed many of the rarer 
species of the upland stand. Sugar maple was represented in all slope 
sections, although the basal area and importance values fluctuated rather 
widely. Beech stopped at the bluff; whereas, oak, hickory and blue ash 
were restricted to the bluff edge and dropped out as the mesic lower 
slopes were reached. The vertical distribution of these species is similar 
to the horizontal distribution shown in Table 4, with respect to moisture 
conditions. Basswood, walnut and slippery elm increased in response to 
increasing mesic conditions as they did in the slope aspect comparison. 
Species typical of wet sites, such as box-elder, sycamore and honey-locust 
were encountered as the floodplain was neared. 

Potzger Woods 

Stand Description. Species attributes are listed in Table 6 for the 
24 species greater than 4 inches dbh. Both the stand density of 121.43 
stems per acre and the stand basal area of 116.58 square feet per acre are 
slightly higher than the corresponding values for the preceding stand. 
This stand is heavily dominated by sugar maple and beech with the two 
species accounting for nearly 70% of stand importance. Also of interest 



table 6. Stand Attributes for Potzger Woods, based on trees U" dbh and 

6.1+U acres — full tally. 



D 2 

D 3 

B 2 

B 3 


B A/ Tree 

Acer saccharum 







Fag lis grandi folia 







Juglans nigra 







Fraxinus americana 







Nyssa sylvatica 







Ulmus rubra 







Liriodendron tiilipifera 







Prunus serotina 







Cornus florida 







Ostrya virginiana 







Ulmus americana 







Tilia americana 







Celtis occidentalis 







Carya cordiformis 







Quercus rubra 







Fraxinus pennsylvanica 







Fraxinus quadrangulata 







Quercus alba 







Sassafras albidum 







Aesculus glabra 







Carya ovata 







Carya glabra 







Quercus muehlenbergii 







Acer rubrum 










is the 5.05% importance value for walnut, which represents one of the 
highest upland stand values for that species in the state. 

When the size-class data are examined (Table 7) the preponderance 
of small sugar maple is immediately apparent. Again, ratios were calcu- 
lated for the proportion of sugar maple and beech below and above 12 
inches. For sugar maple greater than 2 inches the ratio of small to 
large trees is 1002/26 or 38.54, and the corresponding figure for greater 
than 4 inches is 416/26 or 16.00. The ratios for beech are 10/84 or 0.12 
and 8/84 or 0.10, respectively. A comparison with previous studies of 
Potzger Woods indicates that sugar maple is gaining in stand impor- 
tance, possibly at the expense of beech. This is discussed in more detail 
in the section comparing the two stands. 

There are 52 stems greater than 28 inches, or an average of 8 large 
trees per acre. Forty-four (84.4%) of the large trees are beech. The 
largest individuals of species in the full tally plot include: beech 45.8, 
tulip-poplar 35.2, sugar maple 32.4, sassafras 29.0 (44 feet clear bole), 


Indiana Academy of Science 



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Ecology 225 

black gum 28.7, hackberry 28.1, basswood 25.4 and walnut 24.5. The 
general canopy height of this stand is 110 to 115 feet with a few taller 
trees reaching 120 feet. Increment cores taken from the outer portion 
of large tulip-poplar and beech trees (34.8 and 34.1 inches dbh, respec- 
tively) indicate that they are still steadily growing: a 3-inch increase in 
radius in 24 years or 0.25 inch diameter average increase per year for 
tulip-poplar, and a 3-inch radial increase in 33 years or 0.18 inch 
average diameter increase per year for beech. A complete coring of a 
22.5-inch walnut showed the tree to be 66 years old, or an average 
annual diameter growth increase of slightly over Vz inch. 

Stratum rank values for 10 species of shrubs and vines are: hinder a 
benzoin and Smilax rotundi folia, 5; Viburnum aceri folium and Vitis sp., 
4; Parthenocissus quinquefolia and Ribes cynosbati, 3; Asimina triloba, 
Rhus radicans, Viburnum rufidulum and Sambucus canadensis, 2. 

Twenty-four species of summer herbs and ferns had the following 
stratum rank values: Impatiens pallida, 7; Galium concinnum and 
Urtica procera, 6; Galium circaezans and Podophyllum peltatum, 5; 
Adiantum pedatum, Arisaema triphyllum, Athyrium asplenioides, Circaea 
latifolia, Osmorhiza claytoni, and Polygonatum pubescens, 4; Cimicifuga 
racemosa, Dryopteris hexagonoptera, Menispermum canadense, Ranunculus 
septentrionalis, Smilacina racemosa, Solidago caesia and Viola sp., 3; 
Actaea alba, Anemone quinquefolia Boehmeria cylindrica, Polystichum 
acrostichoides and Sanguinaria canadensis, 2; and Panax quinquef olius , 
1. The fern flora is much richer than in Jackson Woods. 

Comparison with Previous Studies. Potzger Woods is one of the 
few stands in the state to have four forest ecological studies during a 
20-year period. Unfortunately, the exact plot and sample point locations 
were not permanently marked so that exact comparisons could be made. 
Potzger (9) plot sampled the woods using 33 ten-meter square quadrats, 
for a sample totalling about 12.5% of the nearly 6V2 acres full tallied in 
this study. In 1955, Petty (8) full-tallied 5 acres of the least-disturbed 
part of the stand; however, it is unknown if the present full-tally com- 
pletely included the earlier 5-acre census. Stearns (15) intensively 
sampled the area by the random pairs method by establishing 140 points 
and recording data for 280 trees over 4 inches dbh. His sample would 
be equivalent to nearly 36% of the 782 trees included in this tally. 

A comparison of these studies clearly indicates the value of taking 
full tallies of forest remnants whenever possible. Although there have 
undoubtedly been some compositional changes in the stand during the 
past 18 years, many of the differences in stand attributes obtained in the 
four studies are due to differences in sampling adequacy and data 

This is particularly apparent when the rarer species are considered. 
In both full tallies, 24 species were recorded, although only 21 were com. 
mon to the two studies. Differences in the species lists result largely 
from the plots not being exactly the same size and in the same location, 
since only one individual was involved for five of the six species in 

226 Indiana Academy of Science 

question. The other case is a question of how consistently white ash 
(Fraxinus americana) and green ash (F. peyinsylvanica) can be separated 
in the field. There should, obviously, be no problem in obtaining com- 
parable species lists for a stand when full tallies include the identical 
stand segment. By comparison, Potzger (9) recorded only 16 of the 24 
species listed in this study with his 12.5% sample and Stearns (15) 
encountered only 15 species in a 36% sample, the latter being a much 
more intensive forest sample than is usually taken. Coefficients of com- 
munity, based on relative density, were calculated for the four studies of 
the single stand. Relative density is used in this comparison since impor- 
tance values were not determined in the two earlier studies. The coeffi- 
cients of community ranged from 61.8 to 82.3% similarity with the two 
full tallies giving the greatest similarity. The similarity between the two 
full tallies probably would have been even closer except that density for 
the former study was based on 3-inch and larger trees; whereas, this 
study was based on trees greater than 4 inches dbh. 

Substantial composition changes could also occur in an 18-year 
period. The main changes in density during the 13-year period between 
Petty 's (8) study and this study include an increase in sugar maple 
from 42.4% to 56.5% and a decrease in American elm from 5.0% to 
1.3%. All other species have remarkably constant values between the 
two studies. The latter change is largely due to mortality of American 
elm due to disease. Sugar maple most likely is increasing substantially 
in density, particularly in the smaller size classes (Table 7). When 
the present sugar maple density values are compared to Potzger's 1950 
study, the change is even greater: 39.9% to 56.5%. 

Stearns (15) found beech and sugar maple importance values of 
43.8% and 24.5%, respectively, as compared to 30.7% and 38.5%, respec- 
tively, in this study. Comparable shifts in relative density and relative 
basal area are as follows: sugar maple increased from 33.9% to 56.5% 
density, and from 15.0% to 20.5% for basal area; whereas, beech 
decreased from 27.5% to 11.8% density and from 60.1% to 49.6% basal 
area. Some differences would occur due to sampling differences, as men- 
tioned previously, but these data indicate a definite trend toward increased 
maple importance largely at the expense of beech. 

This comparison is not intended to be a criticism of any previous 
study. Each study was well conceived and properly executed. The prob- 
lem is one of sampling intensity and adequacy, particularly with respect 
to low frequency species. Since full tallies can be made for most 
accessible stands at the rate of about one acre per hour for all field labor, 
it is hardly justifiable when working with small stands, to compromise 
the adequate stand data obtained from full tallies for the slight gain 
in field efficiency obtained by low intensity sampling. Furthermore, it is 
questionable whether Potzger Woods could have been sampled by taking 
140 sets of random pairs of trees in less time than the six hours re- 
quired in this study to full tally the stand. 

Ecology 227 

Comparison of Stands 

There is remarkable vegetational and floristic similarity between the 
two stands (Tables 1 and 6). Twenty-one of the 24 species of woody 
plants greater than 4 inches dbh occurring in Potzger Woods are also 
found in Jackson Woods. Beech has almost identical absolute values for 
density and basal area in the two stands, but sugar maple represents over 
13% greater importance in Potzger Woods by virtue of a very high 
density per acre of 68.6 and nearly double (23.9 to 12.7) its former basal 
area value. The only other substantial shifts are decreases in importance 
value for tulip-poplar from 14.1% to 2.8% and dogwood from 6.5% 
to 1.5%. The combined importance value for beech and sugar maple is 
11.7% greater in Potzger Woods than in Jackson Woods, compensating, in 
part, for the decreased importance of tulip-poplar in the former stand. 

When the two stands were compared by coefficients of community 
based on tree importance values, the stands were 77.8% similar. This 
compares well with the highest coefficient found (82.3%) when the four 
studies of Potzger Woods were compared. 

The two stands are also very similar with respect to shrub and 
herb composition. All nine of the shrub and vine species present in 
Jackson Woods were present in Potzger Woods. The only additional 
shrub in Potzger Woods was Viburnum ruficluhim, the southern black 
haw. Coefficients of community based on stratum rank values for shrubs 
and vines yielded a similarity percentage of 80.0. Sixteen of the 22 and 
24 herb species found in Jackson and Potzger Woods, respectively, were 
in common to both woods. The coefficient of community based on herb 
species was 61.5% similarity. 

With such close similarity between the local climates, physiography, 
drainage and soils of the two sites, lack of vegetation similarity would be 
anomalous. However, the degree of similarity described here is much 
closer than for most individual stands within a forest association. 

Apparently even the time and amount of past disturbance has been 
similar. When the density was plotted (on the ordinate) per size class 
(on the abscissa) on semi-log paper, each of these stands exhibited the 
same departure from the straight line relationship considered typical of 
old-growth stands. Both stands have similar pronounced plateaus in the 
18- to 30-inch size classes, which indicate substantial disturbance during 
the same time period. The disturbance largely reflects the moderate 
cutting about 30 to 40 years ago, as previously mentioned. This selective 
cutting favored seedling establishment and release of established indi- 
viduals of relatively intolerant species such as tulip-poplar, walnut, ash, 
slippery elm, wild cherry and sassafras. These relationships become clear 
when Tables 2 and 7 are examined. 

Phytogeographic Position of the Stands 

In her discussion of the Area of Illinoian Glaciation in southeastern 
Indiana and adjacent southwestern Ohio, Braun (1) states: "In transi- 
tional bands between the flats and the slopes of the area, where drainage 

228 Indiana Academy of Science 

and aeration are better and yet dissection is not apparent, sugar maple 
appears with the beech. This beech-maple forest is a serai community 
which is replaced by mixed mesophytic forest whenever dissection 
becomes more evident." She goes on to add concerning the dissected 
areas of that region: "In most of the mixed mesophytic forest communi- 
ties of this section of the Western Mesophytic Forest region, beech 
forms approximately 50 per cent of the canopy. In the mixed mesophytic 
communities, tulip tree, sugar maple, basswood, walnut and white ash 
are almost always present together with some six or eight species of 
lesser frequency." 

The authors feel that the upland stands described in this paper fit 
Braun's description almost exactly. Both sites are well-drained, but not 
excessively dissected. Although the importance values (Tables 1 and 6) 
give the impression that maple is co-dominant with beech, inspection of 
the basal area columns (the best single attribute for expressing domi- 
nance) places sugar maple in more nearly its actual position. In both of 
these stands, beech represents about 50% of the stand basal area, while 
10 and 8 additional species (including all of those Braun mentioned) have 
about 1% or greater relative basal area in Jackson and Potzger Woods, 
respectively. Presently, these upland stands should be classified as 
beech and sugar maple dominated mixed mesophytic forests. 

As previously mentioned, many old-growth stands have prodigious 
sugar maple reproduction and very few young beech, but differential 
survival rates compensate to keep beech persisting or even increasing as 
a co-dominant. This will no doubt be the short-term effect as the stands 
continue to recover during the next 50 years or so from the moderate 
past disturbance. It would appear, however, that over long periods beech 
may lose importance as the physiography and stands mature. Should this 
occur, the stands would be expected to become more mixed in total com- 
position with sugar maple gaining somewhat in importance value. The 
latter situation is exemplified in the west-facing slope data in Table 4. 
Excluding black locust (one individual) and Eastern red cedar (which 
was confined to one small limestone outcrop), 14 species each represent 
greater than 1% of the west-facing slope importance value, indeed a 
very mixed stand with no species clearly dominant. The west-facing slope 
represents mature but not over-dissected topography and a slightly drier 
than average aspect. It is expected that the upland stand composition 
will approach that currently occurring on the west-facing slope, as the 
physiography of the upland matures. 

Although these mixed forests are not the true Mixed Mesophytic 
forests typical of the Cumberland Mountains, they were derived from 
that type in response to Post-Pleistocene northward migration and the 
attendant compositional changes as the forest segregated in the complex 
of habitats in the Laughery Valley (1). Neither yellow buckeye nor 
white basswood, considered by Braun (1) as indicators of the Mixed 
Mesophytic type, were found in the vicinity of the study area. Both are 
reported in Clifty Falls State Park, located about 25 miles south, and 
Deam (2) reports white basswood for Ripley County, but a persistent 
search in these stands failed to produce either species. 

Ecology 229 

Future work on the forests of the Laughery Valley should center on 
long-term changes within the best described old-growth stands and relat- 
ing these stands to the forest complex of the region as a whole. 


1. Two old-growth upland forests were studied by taking full tallies of 
all trees over 2.0 inches dbh for segments of 17.46 acres and 6.44 acres, 
respectively, in Jackson and Potzger Woods. 

2. Both stands are beech and sugar maple-dominated mixed mesophytic 
forests with the two dominant species totaling 57.5% and 69.2% of 
the importance value, respectively. Ten and eight other species each 
represent about 1% or greater importance values in the two stands, 

3. Sugar maple dominates the smaller size classes, indicating that it is 
likely increasing in stand importance, probably in response to limited 
disturbance about 30 to 40 years ago. 

4. Intolerant species such as tulip-poplar, walnut, elm and wild cherry 
have been favored by past cutting which created rather extensive 
canopy openings. 

5. The forest composition of sinkholes changes in response to ponding or 
improved internal drainage. Beech and sugar maple decrease in 
importance, but walnut, elm and tulip-poplar increase with increases 
in internal drainage and increases in sinkhole size. 

6. The effect of slope aspect on forest composition was determined by 
transect sampling. The west-facing slope is a mixed-mesophytic 
forest with many species sharing dominance. Oak and hickory species 
increase in importance value on the excessively-drained northwest 
and north-facing slopes. The steep northeast and east-facing slopes 
are very moist due to subsurface seeps; this results in a species com- 
position similar to floodplains. Sugar maple occurs on all slopes, but 
beech is absent from the northeast and east slopes. 

7. A comparison of four studies of Potzger Woods based on different 
sampling methods indicates that rarer species were encountered only 
in full tallies. Since 1956, sugar maple has increased in stand im- 
portance, possibly at the expense of beech, which has decreased. 

8. Changes in stand composition over the next several decades will likely 
show additional recovery from disturbance and a gain in sugar im- 
portance at the expense of the more intolerant species. Beech will 
likely hold its position as the stand co-dominant. 

9. As the physiography matures and becomes more dissected over a 
great number of years, the stand should become more mixed meso- 
phytic in composition rather than the present beech and maple 
dominated mixed composition. 

230 Indiana Academy of Science 

Literature Cited 

1. Braun, E. Lucy. 1950. Deciduous forests of Eastern North America. 
Blakiston Co., New York. 596 p. 

2. Deam, C. C. 1940. Flora of Indiana. Indiana Dept. of Cons., Indianapolis. 
1236 p. 

3. Pernald, M. L. 1950. Gray's Manual of Botany. American Book Co., 
New York, N.Y. 1632 p. 

4. Lindsey, A. A. 1956. Sampling methods and community attributes in 
forest ecology. For. Sci. 2:287-296. 

5. Lindsey, A. A., R. O. Petty, D. K. Sterling, and W. Van Asdall. 1961. 
Vegetation and environment along the Wabash and Tippecanoe Rivers. 
Ecol. Monogr. 31:105-156. 

6. Lindsey, A. A., D. V. Schmelz, and S. A. Nichols. 1969. Natural Areas 
in Indiana and their Preservation. Indiana Natural Areas Survey, 
Dept. Biological Sciences, Purdue Univ., Lafayette. 594 p. 

7. Little, E. L., Jr. 1953. Check list of native and naturalized trees of the 
United States. Agric. Handbook No. 41, U.S. Forest Service, Washington, 
D.C. 472 p. 

8. Petty, R. O. 1955. Tabulation of woody species on five acres at Ver- 
sailles State Park. Unpublished Undergraduate Thesis, Butler University. 

9. Potzger, J. E. 1950. Forest types in the Versailles State Park Area, 
Indiana. Amer. Midi. Natur. 43:729-742. 

10. Potzger, J. E., and Leland Chandler. 1950. Beech in the forests about 
Laughery Creek Valley. Proc. Indiana Acad. Sci. 59:82-94. 

11. Potzger, J. E., and Leland Chandler. 1952. Oak forests in the Laughery 
Creek Valley, Indiana. Proc. Indiana Acad. Sci. 62:129-135. 

12. Potzger, J. E., and A. N. Liming. 1953. Secondary succession in stands of 
red maple-sweet gum-beech forests in Ripley County, Indiana. Butler 
Univ. Bot. Stud. 11:50-59. 

13. Potzger, J. E., and Esther Potzger. 1950. Secondary succession in an 
Illinoian Tillplain habitat. Proc. Indiana Acad. Sci. 59:95-101. 

14. Reynolds, Wm., and J. E. Potzger. 1950. Distribution of Quercus muehlen- 
bergii in Indiana. Butler Univ. Bot. Stud. 10:71-79. 

15. Stearns, Forest. 1956. Forest communities in Versailles State Park, 
Indiana. Butler Univ. Bot. Stud. 13:85-94. 

A Study of Collembolan Populations Associated with Four 
Serai Stages Leading to the Beech-Maple Climax 

Patricia M. Arnett, Indiana State University 


Ninety-six litter samples of one square decimeter each were taken in 
April through July from an old field, oak and maple-oak dominated serai 
stages, and a beech-maple climax in Parke County, Indiana. The objective 
was to determine the relationship between Collembola and the serai stages. 

From the 1821 Collembolans collected through modified Tullgren fun- 
nels, 59 species were found. The maple-oak had more species than any 
other area. This difference was highly significant. The oak area had a 
higher number of individuals per 100 cm 3 of litter than the other areas. 
This difference was highly significant. The oak area had the most indi- 
viduals, the largest volume of litter, the highest total of prominence values 
of all species, and the greatest weekly average of individuals. In all attri- 
butes the field had the lowest value. Isotobryoides ochracius Maynard and 
Onychiurus armatus Tullberg were more frequent in the maple-oak area 
than in the other areas. The difference was highly significant. A three- 
dimensional ordination showed the similarity between the Collembola and 
the tree composition increased with each successive serai stage. The 
ordination indicated the collembolan populations of the wooded areas were 
more closely related to each other than to the Collembola in the field. 


Collembola are common inhabitants of the humus and upper soil 
layers in many ecological situations. The principal objective of this study 
was to determine the relationship of collembolan populations to plant 
serai stages. Leaf litter was collected from an old field, and the oak, 
maple-oak, and climax beech-maple areas of Allee Memorial Woods, 
which is located one and one-half miles northwest of Annapolis, Parke 
County, Indiana. The Collembola extricated from the litter were counted 
and identified. 

Description of study area 

Allee Woods is in the Tipton Till Plain close to the southern 
boundary of the Wisconsin glaciation (8). Soils are melanized podzolics 
with low fertility (7). The woods are severely dissected with 170 feet of 
relief. Three deep gorges are present along with a high bluff over Sugar 
Creek and numerous sloping ravines. All study sites were on the upland. 
Allee Woods was chosen for study because of the presence of numerous 
serai stages and a remnant of beech-maple dominated climax forest. 

The beech-maple area represented the climax vegetation in the 
woods. There has been no cutting although windthrow has removed a 
few mature Acer saccharum Marsh, and Fagus grandifolia Ehrh. Beech 
and sugar maple are co-dominants with Liriodendron tulipifera L. an 
important associate. The understory consists primarily of sugar maple 
and beech saplings and Cornus florida L. The soil is silt loam and is 


232 Indiana Academy of Science 

slightly acid. The litter depth averaged 3.83 centimeters. A slight 
southwestern slope was present. 

The maple-oak probably represented an intermediate stage between 
the beech-maple climax and the oak stage as evidenced by the large 
number of sugar maple saplings. It was selectively cut about the turn of 
the century when some white oaks were removed. Quercus alba L., 
Quercus rubra L., and sugar maple are dominant species. The under- 
story is mainly beech, sugar maple, and dogwood. The soil is slightly 
acid silt loam. The average litter depth is 4.41 centimeters. The letter is 
more uniformly spread over the ground in this area than in the others. 
The area slopes gently toward the west. 

The oak area is predominantly white oak and red oak. Sugar maple, 
Asimina triloba (L.) Dunal., and Ostrya virginiana (Mill) K. Koch form 
the understory. The soil varies from acid to slightly acid loam. This 
area was completely cut over about fifty years ago and has remained 
undisturbed. The present evenly aged trees represent reproduction and 
sprouting stumps. This area had the highest average litter depth, 4.58 
centimeters, consisting primarily of oak leaves. The area slopes slightly 
toward the south. 

Tulip-poplar, Acer rubrum L., and Sassafras albidum (Nutt.) Nees 
are the trees most abundant in the old field. The soil is silt loam and is 
slightly acid. The field has a slightly western slope. This area had the 
least average amount of litter, 3.21 centimeters. The field was previously 
pastured and cultivated but has been abandoned for about thirty years. 
Several successful stages are present with parts of the field covered 
mainly by grasses while other sections have low shrubby vegetation or 
small to medium-sized trees. 

Methods and Materials 

A 100- by 200-foot tract, presumably representative of the vegeta- 
tion in each of the four areas, was chosen. Within each tract, sample 
plots one decimeter square were selected at random before the actual 
sampling began. Each week for the 12 weeks from April 20 to July 5, 
1968, two samples of leaf litter from each tract were taken by means of 
a trowel. The depth of the litter was measured to the nearest 

To prevent condensation on the sides of the funnel and the consequent 
loss of organisms, the litter was inverted and gently placed in a 
Tullgren funnel so that numerous air passages existed between the 
litter and funnel wall. A 15-watt bulb was suspended 3-4 inches above 
the sample for 24 hours. Insects were collected in 95 per cent ethanol to 
preserve coloration. The nomenclature followed that of Maynard (6). A 
species list is located in Table 1. Voucher specimens are on file in the 
Indiana State University entomological research collection. 

Ecology 233 

table 1. A list of collembolan species of Allee Woods. 


Entomobrya assuta Folsom 

Entomobrya atrocincta f. pseudoperpulchra Mills 

Entomobrya marginata Tullberg 

Entomobrya multif asciata Tullberg 

Entomobrya sp. A 

Entomobrya sp. B 

Isotobryoides ochracius Maynard 

Lepidocyrtus curvicollis Bourlet 

Lepidocyrtus unifasciatus James 

Lepidocyrtus sp. A 

Lepidocyrtus sp. B 

Lepidocyrtus sp. C 

Orchesella ainsliei Folsom 

Tomocerus elongatus Maynard 

T omocerus flavescens Tullberg 

Tomocerus minor Lubbock 

Tomocerus vulgaris Tullberg 

T omocerus sp. A 

Willowsia sp. A 

Willowsia sp. B 


Folsomia fimentaria L. 

Folsomia quadrioculata Tullberg 

Folsomia sp. A 

Isotoma eunotabilis Folsom 

Isotoma olivacea Tullberg 

Isotoma viridis Bourlet 

Isotoma sp. A 

Isotomu?*us palustris Muller 

Proisotoma immersa Folsom 

Proisotoma minuta Tullberg 


Onychiurus armatus Tullberg 


Anurida sp. A 

Hypogastrura tigrina Harvey 
Neanura barberi Handschin 
Pseudachorutes simplex Maynard 
Xenylla welchi Folsom 
Poduridae sp. A 
Poduridae sp. B 
Poduridae sp. C 

234 Indiana Academy of Science 

Table 1 — Continued 


Arrhopalites binoculatus Borner 

Denisiella sp. A Folsom and Mills 

Dicyrtomina variabilis Maynard 

Katiannina macgillivrayi Banks 

Neelus albus Maynard 

Neelus maculosus Maynard 

Neelus sp. A 

Neelus sp. B 

Phenothrix sp. A 

Sminthurides lepus Mills 

Sminthurides sp. A 

Sminthurides sp. B 

Sminthurides sp. C 

Sminthurinns radiculus Maynard 

Sminthurinus radiculus f. pictus Maynard 

Sminthurinus sp. A 

Sminthurinus sp. B 

Sminthuridae sp. A 

Sminthuridae sp. B 

Sminthuridae sp. C 

Analysis of data 

The ordination procedure (3, 1) is based on prominence values per 
species per community. From the prominence values, which are equal 
to the density times the square root of the frequency, a coefficient of 
community can be calculated. The inverse of the coefficient of two 
stands can be equated with linear distance and transformed into a three- 
axis spatial pattern which will show a significant correlation between 
the actual, measured distances and the original coefficient of community 
between the two given stands. Collembola were ordinated in three dimen- 
sions by this method. The trees were ordinated in a slightly different 
manner. A full tally of all trees whose diameter, breast height, was 
greater than 2 inches was made for each area. The average of relative 
density and relative basal area for each species with dbh equal to 4.0 
inches or more is called the importance value (5). Importance values 
were used instead of prominence values in the computations. Chi- 
square tests were used to test conclusions concerning relative density and 
frequency of species in each area. 

Results and discussion 

A total of 1821 individuals of Collembola was collected, including 59 
species. The total amount of litter collected was 38,500 cubic centimeters. 
For all four tracts combined there was an average of 4.72 individuals per 
unit volume. A unit volume is defined as one-tenth of a cubic decimeter. 





















































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236 Indiana Academy of Science 

There was an average depth of 4.02 centimeters of litter per sample. The 
lowest number of Collembola ever collected from a sample was 0.00 in 
one old field plot. The highest number was 163 from a plot in the oak 

The oak area was unique in many ways (Table 2). It had (a) the 
highest average number of individuals (density) per unit volume; (b) 
the greatest number of individuals; (c) the highest average litter depth; 
(d) the highest total of prominence values of all species; and (e) the 
largest average number of individuals per week. In every attribute the 
old field had the lowest value. The beech-maple area was intermediate in 
value for every characteristic. Only once did the maple-oak area have 
the highest value; it had the greatest total number of species present 
per area. 

The number of individuals per unit volume was greatest in the oak 
area followed by decreases in relative abundance in the beech-maple, 
maple-oak, and old field areas. This difference was highly significant 
(X 2 — 213.96, 3df). The maple-oak area had more species present in the 
litter (44) than the other areas. The difference was significant 
(X 2 = 9.60, 3df). A possible explanation is that the maple-oak area com- 
bines the features and hence Collembola of the oak and beech-maple 
areas. Although the old field had an average depth of 3.21 centimeters, 
some of this depth is a result of the cushioning and buoyant effect of 
the grasses in the litter. Hence the value cited is probably high. 

Isotobryoides ochracius Maynard appeared in 71 per cent of all 
samples in the maple-oak area but in only eight per cent of the samples in 
the old field. This difference was highly significant (X 2 = 15.32, 3df). 
Onychiurus armatus Tullberg was present in 83 per cent of the maple- 
oak samples but in only 12 per cent of the old field samples. The differ- 
ence was highly significant (X 2 = 10.48, 3df). Table 3 lists the density 
per unit volume and prominence values for the ten most common species. 

Some species could not be identified because of the absence of 
suitable taxonomic keys. More samples over a longer period of time might 
allow us to identify an increased number of the more difficult species. 
Some of the immature stages cannot be identified beyond family (2). 
Sminthuridae sp. A, which has occurred three times but only in the old 
field, may be an indicator species. Similarly, Sminthuridae sp. C has 
occurred only in the beech-maple area and could be an indicator species. 

The most important species in terms of density, frequency, and 
prominence value is Tomocerus minor Lubbock. The second most impor- 
tant is Onychiurus armatus. The former was cosmopolitan in distribu- 
tion while the latter is prevalent in the maple-oak area. The most 
important genus in terms of density, frequency, and prominence value is 
Entomobrya followed by Tomocerus and Onychiurus. The most impor- 
tant family by the same criteria is the Entomobryidae. However, both 
the Entomobryidae and Sminthuridae have 20 species represented. 

Because trees are one of the primary constituents of a community 
and their leaves form the bulk of the litter, an ordination was made on 



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Indiana Academy of Science 

a full tally of the trees and on the Collembola of each area to determine 
if there is a relationship between the two populations. The data were 
plotted on the same set of axes (Figure 1). The interpoint distances 
were measured graphically from the trees of one stand to the Collembola 
of the same stand to determine the degree of similarity between the two 
populations. In lowest terms the distances are as follows: old field, 131 
units; oak, 121 units; maple-oak, 42 units; and beech-maple, 34 units. 
There was a definite straight line relationship showing that the distance 
between the collembolan population and the tree composition, i.e., the 

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populations of four serai stages. 

Ecology 239 

dissimilarity between the two as measured by interpoint distance, 
decreases as the climax area is approached and stability is reached. In 
the more advanced serai stages, the maple-oak and beech-maple, the ordi- 
nation results indicated equilibrium is approached between the woody 
vegetation and the collembolan population. 

When the Collembola and trees were ordinated in the third dimen- 
sion, the Collembola occurred in two main groups with respect to each 
other. The Collembola of the wooded areas were closely grouped together. 
Interpoint distances were beech-maple (BM) to maple-oak (MO) = 36.2; 
BM to oak (0) = 40.8; and MO to = 48.0. When each wooded area was 
compared to the old field, another grouping occurred. Interpoint distances 
were BM to old field (OF) = 134.2; MO to OF = 146.2; and to 
OF = 151.7. The obvious conclusion is that the collembolan population 
of the wooded areas are closely related to each other while the collembolan 
population of the old field is a separate group. 


1. The largest numbers of Collembola per unit volume existed in the oak 
area and the smallest numbers in the old field. 

2. The maple-oak area had the largest number of species (44) while the 
old field had the smallest (20). 

3. Isotobryoides ochracius and Onychiurus armatus were present in the 
highest frequency in the maple-oak area. 

4. The most important species in terms of density, frequency, and 
prominence value was Tomocerus minor. Second was Onychiurus 
armatus. The most important genera are Entomobrya, Tomocerus, 
and Onychiurus. The most important family was Entomobryidae. 

5. Ordination showed that the similarity between the collembolan popu- 
lation and the tree composition in each area increased with each 
succeeding serai stage; the greatest similarity was found in the 
climax beech-maple area. 

6. The Collembola in wooded areas were more similar to each other than 
to those in the old field. 

Literature cited 

1. Beals, E. 1960. Forest bird communities in the Apostle Islands of 
Wisconsin. Wilson Bull. 72:156-181. 

2. Bellinger, P. F. 1954. Studies of soil fauna with special reference to the 
Collembola. Conn. Agr. Exp. Station, New Haven, Conn. Bull. 583. 67 p. 

3. Bray, J. R., and J. T. Curtis. 1957. An ordination of the upland forest 
communities of Southern Wisconsin. Ecol. Monog. 27:325-49. 

4. Folsom, J. W. 1937. Nearctic Collembola or spring-tails, of the family 
Isotomidae. Bull. U. S. Nat. Mus. 168. 145 p. 

240 Indiana Academy of Science 

5. Lindset, A. A. 1956. Sampling methods and community attributes in 
forest ecology. Forest Sci. 2:287-296. 

6. Maynard, E. C. 1951. The Collembola of New York State. Comstock Pub- 
lishing- Company, Inc., Ithaca, New York. 388 p. 

7. Petty, R. O., E. C. Williams, and R. A. Laubengayer. 1961. Ecologic stud- 
ies of a ridge forest and adjacent flood plain, Parke County, Indiana. 
Wabash College, Crawfordsville, Ind. 197 p. 

8. Schneider, A. P. 1966. Physiography, pp. 41-56. In A. A. Lindsey (ed.), 
Natural features of Indiana. Indiana Academy of Science, Indianapolis. 
600 p. 

Woodcock Singing Ground Descriptions for Two Indiana Sites 

William B. Crankshaw, Jerry A. Smith, and Ralph D. Kirkpatrick, 

Ball State University 


The American Woodcock, Philohela minor, has a statewide breeding - dis- 
tribution in Indiana. While conducting- a survey of woodcock activity on 
singing grounds in the spring of 1968 we found that singing ground cover 
requirements were more comprehensive than anticipated. Two contrasting 
areas were selected which were known singing grounds, one in Randolph 
Co. and one in Delaware Co. The Delaware Co. site is situated in cultivated 
fields with subdivisions in close proximity. The Randolph Co. site is also 
in a cultivated area but due to poor drainage the site proper has minimal 
disturbance. No subdivisions are located near the Randolph Co. site. A 
description was made of each site on the basis of vegetation, soils, drain- 
age, land use and the more common fauna. 


The American woodcock {Philohela minor) has a statewide breed- 
ing distribution in Indiana. While conducting a survey of woodcock ac- 
tivity on singing grounds during April and May 1968, we noted that 
singing ground cover requirements appeared to be quite broad. A search 
of the literature revealed no descriptions of singing grounds in Indiana. 

Study areas in Delaware and Randolph Counties were chosen after 
woodcock were known to be using them as singing grounds. Field work 
was conducted during June. 

Delaware County Study Area 

The site chosen for the following description was considered typical 
of several woodcock singing areas located north of Muncie in central 
Delaware county. The site is located at T21N, R9E, Sec 25, NE 1 ^. The 
general area is one of rolling cultivated land, with occasional small wood- 
lots and scattered subdivisions. The woodlots are generally oak-hickory 
stands with a brushy understory indicating a history of intensive timber 
removal. The subdivisions are composed of one story homes usually on 
half acre lots with virtually no trees. 

The site surveyed was one of cultivated fields through which a small 
dredged stream, Jake's Creek, flows in a southwesterly direction. A small 
stand of trees is located approximately three hundred yards to the south 
of the stream. The singing birds had been heard in the cultivated fields 
north of and adjacent to the stream. North of the surveyed area is a com- 
plex of old farm buildings used only for implement storage. North of the 
barns and across a road, is a small housing development. There is no 
evidence of livestock pastured in the singing area. The land use for the 
past few years has been that of cash grain crops. 

Jake's Creek is located about eight feet below the general land 
level due to dredging. It averages about six feet in width, is relatively 


242 Indiana Academy of Science 

deep for its width, has a low turbidity and is a perennial stream. A 
strip of natural vegetation 35 feet on each side parallels the creek 

The soils represented in the singing area are of the Morley-Blount- 
Pewamo series. 

The cultivated fields on both sides of the stream have been under 
apparently continuous crop cover for some years with soybeans, wheat, 
oats, and corn evident this current year. The strip along the stream 
was composed of small trees, shrubs, and herbaceous plants. The dom- 
inant plant of the herbaceous species was Redtop {Agrostis vulgaria); 
dominance was shared in the woody plants by several species. The fol- 
lowing is a list of the woody plants present on the surveyed area: Black 
walnut (Juglans nigra), Pignut hickory (Carya glabra), Shagbark 
hickory (Carya ovata), Hackberry (Celtis occidentalis) , White ash Haw- 
thorn (Crataegus spp.) , Red-osier dogwood (Cornus stolo?iifera) , Honey- 
locust (Gleditsia triacanthos) , Black willow (Salix nigra), Boxelder 
(Acer negundo), Bur oak (Quercus macrocarpa) , and Smooth sumac 
(Rhus glabra). The herbaceous plants were: Redtop (Agrostis alba), 
Wild parsnip (pastinaca sativa) , Wild carrot (Daucus carota) , Common 
ragweed (Ambrosia trifida), Red clover (Trifolium erectum) , Dandelion 
(Taraxacum officinale), Chicory (Cichorium intybus) , Common milkweed 
(Asclepias syriaca) , Wild rose (Rosa virginiana), Burdock (Arctium 
lappa), Horseweed (Erigeron canadense) , Common thistle (Cirsium 
lanceolatum) , Canada thistle (Cirsium arvense) , Teasel (Dipsacus syl- 
vertria), Wild morning glory (Ipomoea spp.), Sweet clover (Melilotus 
alba and M. officinalis), Poison ivy (Toxicodendron radicans) , Wild 
lettuce (Lactuca canadensis), Dock (Rumex spp.), Green foxtail (Seta- 
ria viridis), Timothy (Phleum pratense) , Moth mullein (Verbascum blat- 
taria) , and Great mullein (Verbascum thapsus) . 

The birds listed were actually seen while some of the animals were 
included on the basis of various indications of their presence. The birds 
observed were: mourning dove (Zenaidura macroura) , purple martin 
(Progne subis subis), barn swallow (Hirundo rustica ery throg aster) , 
bank swallow (Riparia riparia riparia) , goldfinch (Spinus triatis tria- 
tis) , redwing (Agelaius phoeniceus) , sparrow (Passer domesticus do- 
mesticus) , bronzed grackle (Quiscalus versicolor) , and bobwhite (Colinus 
virginianus) . The mammals observed or which gave evidence of their 
activities were: deer mouse (Peroynyscus leucopus) , meadow mouse 
(Microtus pennsylvanicus) , muskrat (Ondatra zibethica) , woodchuck 
(Marmota monax) , and raccoon (Procyon lotor) . 

Randolph County Study Area 

Cabin Creek Bog was the location of a woodcock singing ground 
in Randolph county. The bog is located about 500 feet south of Cabin 
Creek on Indiana State Hwy. #1. The topography of the region is rolling 
with several smaller bogs nearby. The Cabin Creek Bog was selected due 
to the contrast it presents to the site described in Delaware county. The 

Ecology 243 

bog is elevated above the general terrain and perennial springs flow 
from its highest points. The area surrounding the bog is farmland of 
moderate quality mainly in the U. S. Soil Conservation Service Land 
Capability Classes of III and IV. Woodcock have been observed singing 
and nesting in the bog proper for many years. 

The soil is peaty, probably Rifle peat, and very unstable to walk 
upon. The soil in the bog stays wet throughout the summer as evi- 
denced by the many springs flowing from the bog. 

The more dominant plants in the bog are given here. On the west- 
ern side, above a marshy border of Carex spp., Juncus spp., and Skunk 
cabbage (Symplocarpus foetida) the dominants are mainly prairie plants 
and occur in the open. This area is covered with small hummocks formed 
by plant roots and peat, the most prevalent plant appears to be Prairie 
dock (Silphium terebinthinaceum) and several grasses, Big and Little 
Bluestem (Andropogon gerardii and A. scoparins) , Indian grass (Sor- 
ghastrum nutans), and Panic Grass (Panicum implicatum) . The center 
of the bog, used by nesting woodcocks, is the location of groves of small 
trees and shrubs that have formed an overstory. The main plants of 
this area are Ninebark (Physocarpus opulfolius) and Quaking aspen 
(Populus tremuloides) . A large variety of other shrubs and trees occupy 
this area in association with the aspen and Ninebark. There are: Dog- 
wood (Cornus stolonifera and C. obliqua) , Sumac (Rhus vernix) , Buck- 
thorn (Rhamus laceolata) , Viburum (V. lantago) , Willow (Salix spp.) 
and Hazelnut (Corylus americana) . East of the bog center is another 
open area dominated by the Shrubby cinquefoil (Potentilla fructicosa) . 

The mammals of the area include the meadow jumping mouse 
(Zapus hudsonius), shorttail shrew (Blarina brevicanda) , eastern mole 
(Scalopus aquaticus), and the red fox (Vulpes fulva). Birds observed 
include bronzed grackle and flicker (Colaptes auratus) . 

Discussion and Conclusions 

Delaware County singing ground surveyed was selected because it 
was very typical of the other singing grounds in east-central Indiana. 
It had relatively little cover and was located in the proximity of houses. 
Some of the other singing grounds in the country were situated between 
subdivisions with houses located within two hundred feet in either direc- 
tion. There was some evidence of a correlation between soil types and 
woodcock breeding areas. Most of the breeding grounds were located on 
Pewamo silty clay loams. This soil type has a high moisture holding 
capacity; in Delaware county it is generally artificially drained. 

The Randolph County singing ground is relatively undisturbed by 
agricultural activities. There is open space as is required for singing 
grounds (2, 3) as well as a soft soil suitable for probing for food. Scat- 
tered shrubs are present as Marshall (1) found on Minnesota singing 

244 Indiana Academy of Science 

The American woodcock is apparently not uncommon in east-central 
Indiana. Its singing grounds are located in cover types that are very 
extensive as well as in some unusual less extensive types. 

Literature Cited 

1. Marshall, William H. 19 5S. Woodcock singing grounds at the Cloquet 
Experimental Forest, 1947-1956. Trans. N. American Wildl. Conf. p. 

2. Mendall, Howard L,., and Clarence M. Aldons. 1943. The ecology and 
management of the American Woodcock. Maine Cooperative Wildlife 
Research Unit, Orono. 202 p. 

3. Studholme, Alan T., John D. Buele and Russell T. Norris. 1940. A Study 
of Pennsylvania Woodcocks. Pa. Game News 11(11) :6-7, 23, 30. 

Hemmer Woods : An Outstanding Old-Growth Lowland Forest 
Remnant in Gibson County, Indiana 

M. T. Jackson, Indiana State University 

Interest in old-growth forest stands and other little-disturbed 
remnants of original Indiana vegetation has been heightened by the 
current Indiana Natural Areas Survey. More commonly than not, areas 
that have escaped the major causes of disturbance are founa by ecologists 
to be either extremely inaccessible or on marginal, rugged land that 
has precluded clearing. Hemmer Woods is a welcome exception. The 
Hemmer family has preserved this nearly level lowland forest remnant 
because of their love for large trees and quiet woods. The central pur- 
pose of this paper is to provide a detailed description of this remarkable 
old-growth forest stand before it is ' 'developed" into one of the all-too- 
common land use patterns of the state. 

Location and Description of the Area 

Hemmer Woods is located about two miles southeast of the village 
of Mackey in extreme southeastern Gibson County. The area is in the 
Wabash Lowland Physiographic Province that was subjected to glacia- 
tion during the Illinoian Period. Smith Fork of Pigeon Creek, which is, 
in turn, tributary to the Ohio River, meanders through the entire length 
of the area. The least disturbed section of the woods comprises about 20 
acres located principally in the North Half of the Northwest Quarter of 
the Southwest Quarter of Section 24, Township 3 South, Range 9 West. 
Another nearby wooded tract of about 70 acres is also owned by the 
Hemmers. This large stand has had slight disturbance by limited cuting 
several years ago. Although a description of the latter stand is not in- 
cluded in this paper, it should be the site of future ecological studies 
because it represents a gradual transition from the floodplain to the dry 

The study area is part of an alluvial deposit known locally as the 
"Buckskin Bottoms." The area has less than 10 ft of relief with the 
entire 20-acre stand lying between 410 and 420 ft above mean sea level. 
The soil type was mapped in the 1922 Gibson County Soil Survey Report 
as Waverly silt loam. It has a light gray surface horizon that is medium 
in organic content. Soil samples taken from six locations ranged from 
pH 6.0 to 6.2. The area was naturally poorly drained, owing to its flat 
surface, low position and gentle stream gradient. A deeply-dredged ditch 
located a few rods from the stand has lowered the water table until the 
section of the creek that is located within the stand remains very low 
or dry during most of the year. According to Mr. L. H. Hemmer, the 
ditch was dug in 1918, and prior to that, the creek retained a good flow 
of water all year. 

No living trees have been removed, but dead individuals have been 
cut following lightening strikes or other natural mortality. A small 


246 Indiana Academy of Science 

tornado struck the southern edge of the stand in 1916 and several wind- 
thrown and damaged trees were salvaged shortly thereafter. Mr. Hem- 
mer states that there has been no fire in the stand during this century 
and grazing has been excluded since World War I or earlier. There is 
no present evidence of disturbance except for a very few stumps that 
remain from past cutting of dead trees. The area has been designated 
as a classified forest for about 20 years by the Indiana Division of 


A full census was made by the usual full tally method for 11.33 acres 
centered in the main part of the stand. All stand edges were excluded. 
The major study section (600 ft square) is located in the main east-west 
part of the tract. A smaller contiguous section (roughly a 30°-60° tri- 
angle with a 400 ft base) was tallied in the northeastern part of the 
stand. Temporary markers will be replaced by permanent stakes at the 
plot corners. All trees over 4 inches dbh (diameter breast high) were 
measured to the nearest one tenth inch with diameter tapes and those 
2.0 to 3.9 inches were counted and recorded by species. Tree heights were 
taken with an Abney level. 

Since the area has quite uniform topography, the section tallied is 
treated as a single stand. The tally for one small segment of the stand 
which is located on slightly higher ground was recorded separately and 
is discussed later. Data are summarized in Tables 1 and 2. Stand attri- 
butes are those of Lindsey (2); species nomenclature for trees follows 
Little (5); and shrub and herb nomenclature follows Fernald (1). Com- 
mon names of trees are used in the text for convenience in reading. 

Stand Description 

A total of 1,259 trees were tallied in the size classes greater than 
4.0 inches. The 2.0 to 3.9-inch size class contained 789 individuals, in- 
cluding 28 vines which are omitted from the tables. As indicated in 
Table 1, the stand is of mixed lowland composition with several species 
sharing dominance. Thirty-two species were represented by individuals 
larger than 4 inches dbh, and eleven species had importance values of 
nearly 5% or greater. It is best characterized as a sweet gum — tulip- 
poplar — red maple — elm stand. The presence of tulip-poplar as one of 
the dominant species in this lowland stand is understandable when one 
considers that Ridgway (6) found the species growing to great pro- 
portions in the original forests of the fertile high bottoms of the southern 
Wabash River floodplain. The low gradient stream of this stand has 
probably never flooded extensively for lengthy periods, conditions similar 
to those on the Wabash high bottoms. Lindsey, ct al. (4), failed to find 
tulip-poplar present in the Wabash floodplains, since complete clearing 
for agriculture had removed most of the suitable sites. Increased fre- 
quency and duration of flooding in recent decades undoubtedly has ac- 
celerated the loss of species such as tulip-poplar, which are relatively 
intolerant to inundation. Hemmer Woods has increased value as a 

Ecology 247 

natural area since it is the only known example of floodplain forest in 
the state that includes such large tulip-poplar trees. 

The size of individual trees and the great number of large trees 
are the most striking features of the stand (Table 2). Eleven species 
have a total of 74 individuals in the 30-inch or larger size classes, for 
an average of over six 30-inch plus stems per acre. Eighteen stems 
greater than 40 inches dbh fell within the study plot. At least three trees 
greater than 40 inches fell outside the plot, including a 48.3 inch syca- 
more. Three tulip-poplars exceeded 50 inches dbh; the largest, at 54.6 
inches, may be the largest individual of that species in the state. Twelve 
species had representatives larger than 30 inches with sycamore 48.6 
inches, white oak 44.0, sweet gum 40.7, green ash 37.4, red oak 36.5, 
American elm 35.9, red maple 34,4, hackberry 33.5, river birch 30.8 and 
swamp white oak 30.5. The stand is also noteworthy for large sassafras 
(nine over 20 inches) with the largest at 31.9. A great number of the 
larger trees have clear boles of 40 to 60 feet. The largest tulip-poplar 
measured 151 feet tall and 53 feet to the first limb. The larger tulip 
trees stand 20 to 30 feet taller than the general canopy level of about 
120 feet. Another interesting feature is the large grove of very large 
paw paw trees. Twenty paw paw stems exceeded 4 inches dbh with two 
reaching 6.2 inches. These are among the largest forest grown paw paw 
trees that the author has measured. 

The shrub stratum is dominated by Lindera benzoin, Staphylea tri- 
folia, Euonymns afj'opurpureus, and Smilax rotundifolia. Urtica pro- 
cera abounds in the herb layer, which could aid in the preservation of 
the area! Mr. Hemmer stated that the spring wildflower display is 
very showy, including large colonies of Trillium gleasoni and Mertensia 

Effect of Drainage on Stand Composition 

Although there is little relief within the area, there is remarkable 
ecological separation of species that are usually restricted to either up- 
land or lowland sites. Along such a low gradient stream, three or four 
vertical feet alter the drainage and soil moisture patterns appreciably. 
Since several of the tally strips crossed the area at right angles to the 
stream, it was easy to note the gradual but definite species shift to 
species less tolerant of high water tables as the gentle topographic rise 
was topped. This helps explain the presence of shagbark and mockernut 
hickory, and black oak in the same stand with swamp white oak, syca- 
more, river birch, sweet gum, hackberry and shellbark hickory. 

Another interesting moisture relationship is illustrated by a com- 
parison between the species importance values of a small segment (ap- 
proximately 1 acre) of the stand located on the highest small rise with 
importance values of the main stand. This small segment, on a gentle 
southwest-facing slope, had a combined importance value for the more 
typically upland oaks (white, red and black) and upland hickories (shag- 
bark and mockernut) of 30.9% as compared to 11.6% for the same 
species in the remainder of the stand. A change in importance value of 


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250 Indiana Academy of Science 

nearly 20% for these species indicates much lower soil moisture levels 
on the slope. In addition, the only individual of the most xeric of the 
oak species present (black oak) was found on the same slope. 

Other species with substantial shifts in importance in the slope 
segment include: a) increases on slope; slippery elm 6.3% to 12.3%, 
white ash 2.8% to 8.1%, wild cherry 1.4% to 6.0% and sassafras 5.9% 
to 10.4%) ; b) decreases on slope; sweet gum 14.0% to 3.8%, red maple 
8.7% to 0.6%, American elm 7.9% to 2.4%,, hackberry 7.6% to 3.8%, 
sycamore 6.0% to 0.0%, shellbark hickory 2.2% to 0.0% and river birch 
1.5% to. 0.0%. It is of interest that all species with increases in im- 
portance are more typical of upland stands, while all species with de- 
creases in importance are typical of depressional or floodplain situations. 

Of the five sugar maples larger than 4.0 inches, three were in the 
slope segment. Four of the sugar maples were in the 6-inch size class 
with the lone exception at 13.3 inches. One 4.9 inch sugar maple was 
found on a very slight rise along the creek and another at 6.6 inches 
occurred on higher ground near the south edge of the stand. Both of 
the 2-4 inch maples occurred on higher sites. Only a single red maple 
was found on the slope, as compared to an average of 12.7 red maples per 
acre in the remainder of the stand. 

Future Changes in Composition 

Perhaps the shift from red maple to sugar maple on the drier sites 
is a definitive change that portends a future trend within the stand 
toward species less typical of floodplains. This could indicate the begin- 
ning of a species shift in response to less hydric conditions following the 
lowering of the water table 50 years ago. 

Inspection of the size class data in Table 2 reveals that several 
species seem to be shifting significantly in importance value. Both river 
birch and sycamore are restricted to the immediate vicinity of the creek 
bank and neither is reproducing. Although tulip-poplar has more general 
distribution throughout the stand, apparently it is not reproducing. One 
could assume that it will be maintained in canopy openings resulting from 
windthrow or death, since abundant seed sources are at hand. There are 
few tuilp trees in the area disturbed by the tornado, however. Instead, 
other intolerant species such as sassafras, wild cherry, slippery elm, 
American elm and hackberry represent a larger percentage of the 
medium-sized stems in that area. 

When ratios of the number of individuals smaller than 12 inches to 
the number of individuals greater than 12 inches are compared for 
several species, projected compositional changes become clearer. The 
three members of the Ulmaceae plus red maple have ratios greater than 
3:1 and seem to be invading the stand rapidly. Wild cherry, ash, black 
gum, bitternut and shagbark hickory have ratios decreasing from 2:1 to 
%:1 in the order listed. These species seem to be maintaining their 
position in the stand. River birch, sweet gum, tulip-poplar, white oak, 
shellbark hickory and sycamore have ratios less than 0.5:1 and appear 



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252 Indiana Academy of Science 

to be losing their share of stand dominance. The size distribution ratios 
are, at best, only indications of trends, since they do not completely ac- 
count for differential survival rates among young trees of different 

Since the number of standing dead trees per species gives some in- 
formation about survival rates, all identifiable dead trees were measured 
and recorded. Overall, 4.4% of the total standing stems were dead. This 
appears to be a relatively low mortality rate until one considers that 
most small trees are largely sapwood which decays to a condition that 
precludes species identification in a very few years. Dead trees repre- 
sented a substantial percentage of total standing stems for the follow- 
ing species: sassafras 12.1, black cherry 9.7, American elm 8.6, black 
gum 6.0, sweet gum 4.2 and slippery elm 3.0. American elm seems to be 
maintaining its position in the stand, since widespread mortality from 
elm diseases is not in evidence. 

The stand does not appear to be at climax although largely un- 
disturbed by man. Perhaps an equilibrium had been reached prior to 
drainage. Presently the composition shift appears to be favoring species 
typical of situations of medium soil moisture. Those species favoring 
very moist situations, such as river birch, sweet gum, shellbark hickory 
and sycamore, are apparently decreasing. Windthrow has probably helped 
retain intolerant species such as wild cherry, sassafras and tulip-poplar. 
It is of interest that no beech trees were found in any size class, al- 
though beech occurs quite commonly in surrounding stands. Perhaps the 
relatively recent invasion of sugar maple is the initial step toward a 
beech-maple dominated stand, and that beech invasion will follow suit 
in the future. Long term studies would be of interest here. 

Comparison with Other Stands 

Bottomland and depressional forests frequently have higher basal 
area per acre than most undisturbed upland stands which partially 
explains the moderately high (134 sq ft per acre) basal area in this 
stand. It is surpassed in basal area by only four of the old-growth stands 
reported by Schmelz and Lindsey (7). It has both higher basal area and 
lower density per acre than either Donaldson's Woods in Lawrence 
County or Cox Woods, Orange County, which are often considered to be 
the most impressive forests in the state (7). Beckville Woods in Mont- 
gomery County, which occupies a depressional Brookston silt loam soil, 
was found by Petty (personal communication) to have 135 sq ft per 
acre. A full tally of 21 acres of Kramer Woods along the Ohio River 
in Spencer County was found by Schmelz (personal communication) to 
have 128.5 sq ft per acre, but the stand density was much lower at 78.9 
trees per acre. The mean diameter of trees in Hemmer Woods at 12.2 
inches is about average for old-growth stands in the state, but Kramer 
Woods at 14.6 and Beall Woods at 14.2 (7) indicate a greater percentage 
of large trees. 

When the numbers per size class were plotted on semi-log paper 
according to the method used by Schmelz and Lindsey (7) to analyze 

Ecology 253 

disturbance in stands, the relationship approaches a straight line more 
closely than did any of the 19 stands that they studied. Slight plateaus in 
the 18 to 26-inch size classes and 42 to 50-inch size classes indicate only 
mild disturbance. The former plateau probably reflects the windthrow of 
52 years ago. 

Hemmer Woods has 18 of its 32 species in common with the Beall 
Woods bottomland stand which is located on the larger floodplain of the 
lower Wabash River (3), and 24 of its 32 species in common with Kramer 
Woods. Most important Beall Woods species absent from Hemmer 
Woods are bur oak, silver maple, and pin oak. Shumard's oak repre- 
sented 13% of Beall Woods, but had only 0.11% importance in Hemmer 
Woods. Although pin oak is absent from the stand reported herein, 
it is abundant in medium size classes in the larger Hemmer Woods 
stand. Of the important species in Kramer Woods, only southern red 
oak, swamp chestnut oak and beech are absent here. Again, Shumard's 
oak is very important (23%) in Kramer Woods. 

Coefficients of community, based on importance percentages, were 
calculated to compare the degree of similarity among the three bottom- 
land stands. Hemmer Woods was 39.6% and 32.8% similar to Beall and 
Kramer Woods, respectively. Beall and Kramer Woods were 48.1% 
similar to one another. These values indicate that stands located on flood- 
plains of larger rivers, within a given area, tend to be more similar struc- 
turally than are stands which are located on floodplains of large and 
small streams, respectively. Differences in the frequency and duration 
of flooding between large and small streams is undoubtedly a major 
factor influencing stand composition. 

The only species of southern affinity in this stand are persimmon and 
Shumard's oak, although floodplains normally offer excellent migration 

The extremely large trees, excellent size class distribution, rich 
species composition, high basal area and relatively low density combine 
to make Hemmer Woods one of the finest forests in Indiana. The area 
is certainly of the quality that should be preserved as part of the 
Natural Areas System of Indiana. 

The author wishes to thank Miss Amanda Hemmer and Messrs. 
E. J. and L. H. Hemmer for permission to do the study. 

Literature Cited 

Fernald, M. L. 1950. Gray's Manual of Botany. 8th Ed., American Book 
Company, New York, New York. 1632 p. 

Lindsey, A. A. 1956. Sampling - methods and community attributes in 
forest ecology. Forest Science 2:287-296. 

. 1962. Analysis of an original forest of the Lower Wabash 

Floodplain and Upland. Proc. IndianaAcad.Sci. 72: 

254 Indiana Academy of Science 

R. O. Petty, D. K. Sterling and W. Van Asdall. 1961. Vege- 

tation and environment along- the Wabash and Tippecanoe Rivers. Ecol. 
Monogr. 31:105-156. 

Little, E. H. 1953. Cheek list of native and naturalized trees of the 
United States. Agriculture Handbook No. 41, U. S. Forest Service, Wash- 
ington, D. C. 472 p. 

Ridgewat, R. 1876. Our native trees. The tulip tr,e — Liriodendron tulip- 
ifera. Field and Forest 1:49-53. 

Schmelz, D. V. and A. A. Lindsey. 1965. Size-class structure of old- 
growth forests in Indiana. Forest Science 11:258-264. 


Chairman : Leland Chandler, Purdue University 
Jack R. Munsee, Indiana State University, was elected chairman for 1969 


A Japanese Weevil Discovered in Indiana. 

Donald L. Schuder, Purdue University. — An infestation of adult Jap- 
anese weevils, Pseudocneorhinus bifasciatus Roelefs, was reported 1 by 
a homeowner in Vincennes on June 26, 1968. (The owner had noticed the 
insect's damage in 1967.) Investigation revealed adults to be common in 
a V2 city block area and causing foliar damage to mock orange, privet, 
rose, spirea and weigela and other shrubs. 

This weevil, native to Japan, was first reported in the U. S. in 
1923 in Connecticut. It is now known from Delaware, Georgia, Maryland, 
New Jersey, New York, North Carolina, Pennsylvania, South Carolina, 
Virginia and the District of Columbia, on the eastern coast of the United 
States. The Indiana record is the first report from the midwest. 

The host list of this omnivorous pest is extensive and includes the 
following trees, shrubs, flowers, fruit and vegetables: Ash (white), citrus 
sp., elm, hackberry, hemlock, mimosa, oak, azalea, barberry (Japanese), 
deutzia, camelia, clematis, firethorn, forsythia, honeysuckle, ivy (Eng- 
lish), lilac, mahania, mock orange, mountain laurel, privet (California 
and Regels), rhododendron, rose, rose of Sharon, spirea (Anthony Water- 
er, vanhoutte), weigela, ageratum, chrysanthemum, fern, geranium, hi- 
biscus, lilly-of-the-valley, veronica, lima beans and strawberry. The insect 
occurs in large numbers and cuts broad, rounded sections from the 
margins of the leaves resulting in a crenulated appearance. Injury can 
be extensive and death of hemlocks and rose bushes has been reported. 
The insect feeds in the daytime. 

The weevil is parthenogenetic and the wing covers are fused so 
that it cannot fly. It is robust, grey in color and the elytra are crossed 
by two blackbands. Adults are present from early June until freezing 
weather. Eggs are laid in partially eaten, curled, dried leaves. The 
immature stages are unknown. 

Because of its omnivorous habits this insect constitutes a serious 
threat to the ornamental plantings in homeyards, cemeteries, golf 
courses, strawberry plantings and commercial nurseries. It can be con- 
trolled with early applications of 2.5 percent aldrin, dieldrin, heptachlor, 
malathion or parathion dusts according to Smith (J. Econ. Ent. 48: 628). 

1 The infestation was reported to Robert Dolphin of the U.S.D.A. stationed 
at Vincennes. Identification was made by Rose Ella Warner of the U. S. National 


256 Indiana Academy of Science 

Other papers read 

Insects and Other Arthropods of Economic Importance in Indiana during 
1968. Roger T. Huber and John V. Osmun, Purdue University. 

Notes on the Biology of Aedes flavescens (Muller) in Indiana. R. E. 

Siverly, Ball State University. 

The Artificial Introduction of Microorganisms into an Insect's Alimentary 
Canal. Lois Herzog and Harold L. Zimmack, Ball State University. 

A Histological Study of the Bacteria Escherichia coli and Serratia marc- 
escens in the Larval Stages of the European Corn Borer. Albert 
Esterline, Ball State University. 

Indiana Odonata — 1968. B. Elwood Montgomery, Purdue University. 

A Checklist of the Mosquitoes of Indiana with a Record of the 
Occurrence of Aedes infirmatus D&K 

John W. Hart, Richmond, Indiana 

The first record of a mosquito from Indiana was published in 1827 
when Thomas Say (8) described Culex musicus [=Psorophora ferox 
Humb.]. There were apparently no further writings listing mosquitoes 
for 95 years. Dyar's 1922 publication, The Mosquitoes of the United 
States, (4) listed another four species. 

In 1942 the author prepared a manuscript entitled "Preliminary 
Studies of the Mosquitoes of Indiana" in partial fulfillment of the re- 
quirements for a degree at Purdue University. In July and August of the 
same year, the author worked in Indiana in the Malaria Control in War 
Areas (MCWA) Program of the U. S. Public Health Service. Mosquito 
records gained as a result of both of these studies were reported in a 
preliminary list of Indiana mosquitoes (5). This 1944 paper noted 24 
species known to occur in the state. 

Christensen and Harmston also published a preliminary list of the 
mosquitoes of Indiana in 1944 (3). They reported species collected in the 
1941 and 1942 MCWA studies and included five not in the author's list. 
Carpenter reporting on collection of mosquitoes at military installations 
in Indiana during 1944 and 1945 added two to the list (2), and Brooks 
reported another in 1947 (1). 

The next 13 records were added by Siverly (9 thru 16), Newhouse 
and Siverly (7), and Siverly and Burhardt (17). Two additional mos- 
quito findings were reported in 1968, one by Truman and Craig (18), 
and one by the author (6). 

This paper reports Aedes infirmatus Dyar and Knab not previ- 
ously known from the state. The Purdue University collection contains 
an excellent female specimen taken as part of the 1958 studies at Hovey 
Lake in Posey County. Collection date was August 20. 

Forty-eight species of mosquitos have been taken in Indiana. 
Anopheles barberi Coq. — Christensen and Harmston (3) 
A. crucians crucians Wied. — Christensen and Harmston (3) 
A. punctipennis (Say) — Hart (5) 

A. quadrimaculatus Say — Dyar (4) Collected by E. B. Williamson 
A. walkeri Theo. — Christensen and Harmston (3) 
Toxorhynchites rutilus septentrionalis (D&K) — Hart (6) 
Wyeomyia smithii (Coq.) — Siverly (13) 
Uranotaenia sapphirina (O. S.) — Hart (5) 
Culiseta inornata (Will.) — Hart (5) 
C. silvestris minnesotae Barr — Siverly (15) 
C.morsitans (Theo.) — Christensen and Harmston (3) 
C. melanura (Coq.) — Siverly (9) 
Orthopodomyia alba Baker — Brooks (1) 


258 Indiana Academy of Science 

O. signifera (Coq.) — Hart (5) 

Mansonia perturb an s (Walk.)— Dyar (4) Collected by E. B. Williamson 

Psorophora ciliata (Fab.) — Dyar (4) Collected by E. B. Williamson 

P. howardii Coq. — Hart (5) 

P. cyanescens (Coq.) — Hart (5) 

P.ferox (Humb.)— Say (8) 

P.horrida (D&K)— Hart (5) 

P. varipes (Coq.) — Hart (5) 

P. confinnis (L.A.) — Hart (5) 

P. discolor (Coq.) — Siverly and Burkhardt (17) 

Aedes abserratus (F&Y) — Siverly (16) 

A. aurifer (Coq.) — Siverly (10) 

A. canadensis canadensis (Theo.) — Hart (5) 

A. excrucians (Walk.) — Siverly (12) 

A.fitchii (F&Y)— Siverly (14) 

A. flavescens (Mull) — Siverly (14) 

A. grossbecki D&K — Siverly (10) 

A. infirmatus D&K — Hart — Purdue University Collection 

A. sollicitans (Walk.) — Christensen and Harmston (3) 

A. sticticus (Meig.) — Hart (5) 

A. stimulans (Walk.) — Siverly (14) 

A. thibaulti D&K— Siverly (11) 

A. trivittatus (Coq.) — Dyar (4) Collected by J. J. Davis 

A. hendersoni Cock. — Truman and Craig (18) 

A. triseriatns (Say) — Hart (5) 

A. aegypt (L.) — Hart (5) 

A. vexans (Meig.) — Hart (5) 

A. cinereus Meig. — Carpenter (2) 

Culex pipiens pipiens L. — Hart (5) 

C. pipiens quinquefasciatus Say — Newhouse and Siverly (7) 

C. restuans Theo. — Hart (5) 

C. salinarius Coq. — Carpenter (2) 

C. tarsalis Coq. — Hart (5) 

C. erraticus (D&K) — Hart (5) 

C. territans Walk.— Hart (5) 

Orthopodomyia signifera, (Coq.) and O. alba Baker are indistin- 
guishable in the adult stages. Records of signifera prior to Brooks (1) 
were based on adult females. Though they were probably correctly called 
signifera, a positive identification can not be claimed. The author studied 
the larval material collected by Brooks and concurred in the identifica- 
tion of both species. 

The occurrence of alba in Indiana is to be expected. It is found in 
reliable lists of the mosquitoes of both Ohio and Illinois. 

Entomology 259 

Literature Cited 

Brooks, I. C. 1947. Tree-hole mosquitoes in Tippecanoe County, Indiana. 
Proc. Indiana Acad. Sci. 56:154-156. 

Carpenter, S. J. 1952. Notes on mosquitoes in North America: II, Collec- 
tions at military installations in Indiana during 1944 and 1945. Mosquito 
News. 12(4):252-253. 

Christensen, G. R., and F. C. Harmston. 1944. A preliminary list of 
the mosquitoes of Indiana. J. Econ. Ent. 37(1) :110-111. 

Dyar, H. G. 1922. The mosquitoes of the United States. Proc. U. S. 
Nat. Mus. 62:1-119. 

Hart, J. W. 1944. A preliminary list of the mosquitoes of Indiana. Amer. 
Mid. Natur. 31(2) :414-416. 

. 1968. Occurrence of Toxorhynchites rutilus septentrionalis 

(Dyar and Knab) in Indiana. Mosquito News. 28(1) :118. 

7. Newhouse, V. P., and R. E. Siverly. 1965. The Culex pipiens, complex in 
southern Indiana. Mosquito News. 25(4) :489-490. 

8. Say, Thomas. 182 7. (described Culex musicus=Psorophora ferox from 
Indiana) J. Acad. Nat. Sci. Phila. 6:149. 

9. Siverly, R. E. 1958. Occurrence of Guliseta melanura (Coquillett) in Indi- 
ana. Proc. Indiana Acad. Sci. 67:137. 

10. . 1959. Occurrence of Aedes grossbecki Dyar and Knab and 

Aedes aurifer (Coquillett) in Indiana. Proc. Indiana Acad. Sci. 68:149. 

11. . 1961. Occurrence of Aedes thibaulti Dyar and Knab in Indi- 
ana. Proc. Indiana Acad. Sci. 70:137. 

12. . 1963. Occurrence of Aedes excrucians (Walker) in Indiana. 

Proc. Indiana Acad. Sci. 72:140. 

13. . 1964. Occurrence of Wyeomyia smithii in Indiana. Proc. Indi- 
ana Acad. Sci. 73:144-145. 

14. . 1966. Mosquitoes of Delaware County, Indiana. Mosquito 

News. 26(2):221-229. 

15. . 1966. Occurrence of Culiseta minnesotae Barr in Indiana. Proc. 

Indiana Acad. Sci. 75:108. 

16. . 1967. Occurrence of Aedes abserratus (Felt and Young) and 

Culiseta morsitans (Theobald) in Indiana. Mosquito News. 27(1) :1 16. 

17. Siverly, R. E., and R. W. Burkhardt, Jr. 1965. Occurrence of Psorophora 
discolor (Coquillett) in Indiana. Proc. Indiana Acad. Sci. 74:195. 

18. Truman, J. W., and G. B. Craig, Jr. 1968. Hybridization between Aedes 
hendersoni and Aedes triseriatus. An. Ent. Soc. Amer. 61(4) :1020-1025. 

Crosses of Tropisternus from Central America with Other Color 

Forms of the Tropisternus collar is Complex 

(Colcoptera: HYDROPHILIDAE) 

Frank N. Young, Indiana University 

In several previous papers (1, 3, 5) different color forms of the 
Tropisternus collaris (Fabricus) Complex have been shown to be cap- 
able of interbreeding. North and South American forms readily inter- 
breed and produce viable offspring, but these offspring are highly in- 
fertile. The principal work to date has been carried on with a melanic 
form from Lago de Ayapel, Colombia, and the three North American 
subspecies (striolatus, mexicanus, and viridis). A non-melanic form from 
Puerto Rico is now available, and some of the previous crosses have been 
repeated with it. 

The melanic form from Colombia resembles typical collaris from 
Brazil and Venezuela in the basic arrangement of the elytral pattern, 
but has the elytral dark lines expanded so that in some individuals the 
intermediate areas are eliminated and the elytra appear solid green. The 
pronotal pattern is greatly expanded and the green metallic area becomes 
broader than long in contrast to collaris or mexicanus. The head pattern 
is also extended, but not to the degree seen in viridis from Florida, 
Georgia, and South Carolina. 

In contrast the usual form of Tropisternus collaris found in Panama 
{mexicanus Castelnau) and to the north has six nearly complete dark 
lines on the elytra separated by areas in which yellow, fluorescent pig- 
ments are developed. The pronotal pattern is similar to typical collaris, 
and the dark head pattern is usually reduced, barely or not extending 
anterior to the Y-shaped epicranial suture. 

In the British Museum are four speciments collected by Champion 
in Panama and marked "T. collaris var." apparently by Sharp. These 
were apparently not included in the Biologia Americana material (2), 
unless they are the basis of the citation of Panama under collaris (2, p. 
55). T. collaris is cited from Mexico (collection Saunders) on the 
basis of a very typical specimen which is almost certainly mislabelled 
and probably came from Venezuela. The four Panama specimens col- 
lected by Champion have the elytral lines expanded and the yellow 
pigments largely eliminated, but the head and pronotal markings are 
much as in typical mexicanus. I believe that these specimens are natural 
hybrids between the melanic form of collai^is from northern Colombia 
and the subspecies mexicanus which extends throughout Central America 
and northward into the United States. Other specimens which I have 
seen from Panama are similar to typical mexicanus. 

There are, however, indications in several series from Honduras, 
Guatemala, Costa Rica, San Salvador, and Panama (collected by Kenneth 
Mc Williams and Borys Malkin) that there may be genes from the 


Entomology 261 

melanic form which have flowed northward. Similar lateral extension of 
the dark pronotal blotch, however, is completely lacking in all the 
northern Mexican specimens I have seen and in the United States until 
the zone of intergradation between mexicanns and striolatus is reached 
in eastern Texas, Oklahoma, Missouri, and Ilinois. 

It was hoped that the probem of the taxonomic status of the North 
and South American forms could be solved by laboratory experiments with 
the Central American forms and the laboratory stock from Colombia. In 
the summer of 1967, Mr. (now Dr.) Kenneth McWilliams collected speci- 
mens for me in several countries of Central America. Unfortunately, 
specimens from Panama died in transit, but specimens from Honduras 
and Guatemala arrived alive and six crosses were made with the Colom- 
bian form (AYA), the melanic form viridis from Florida, and between 
the two (Guatemala female x Honduran male). Several of these crosses 
proved successful and 174 adults were reared from 376 larvae (over 

The accompanying tables summarize the crosses, backcrosses, and 
outcrosses made to test the fertility of the parents and hybrids. None of 
the Fi x Fi crosses nor backcrosses of the Honduran x Colombian froms 
proved fertile. However, when Fi hybrids were outcrosses with T. c. 
viridis from South Carolina some fertility was shown. As shown in the 
table, this was low and many deaths occurred in the larvae before the 
first molt. This may have been due to the fact that all of the males 
available for use were laboratory reared from originally natural hybrids 
between T. c. viridis and striolatus. 

The result of these crosses indicate that there is still a very high 
degree of infertility in the F x of hybrids of T. c. mexicanus and the 
melanic form from Colombia even when the populations are closer to- 
gether. The distance, however, is still considerable (over 900 miles), 
and the parent populations may have been separated for a considerable 
time. It still seems that study of the population from southern Panama 
will be necessary before it is possible to declare that T. collaris and T. 
mexicanus (along with its color forms striolatus and viridis) are specifi- 
cally distinct. 

The color patterns in these crosses have not yet been analyzed in 
detail. However, they follow in general the pattern previously noted in 
crosses between the melanic Colombian form and the North American 
forms, that is, in general the darker patterns are recessive to the lighter 
patterns, but there are differences in penetrance. For example, the head 
patterns tend to be slightly more extended than in the lighter parent, but 
not as extended as in the darker parent. Also, there is a persistent 
tendency for the partial expression of the recessive gene in the presence 
of the full expression of the dominant gene. For example, in the cross of 
Honduras x T. c. viridis (Table 1, 5X) all offspring showed darker elytra 
and 14 of 16 showed a more extended head pattern than the lighter 
female parent. (Unfortunately, all of these proved very feeble and 
further crosses could not be made.) 


Indiana Academy of Science 




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Entomology 265 

In the outcross (Table 4, 24X, 25X, 26X) of the Honduras x 
Colombia hybrid, the color patterns of head, pronotum, and elyta show 
great variation. The elytral pattern varies from lighter than the lighter 
parent to darker than the darker and the head and pronotum show a 
similar range. Some of the effects in this cross are doubtless due to 
interacting genes. Some individuals from this cross are still living, but 
to date have proven highly infertile (Tabl 5, 31X, 32X). I wish to express 
my thanks to Christina Fenn (now Mrs. Ronald W. Smith) for her 
assistance in this research. 

Sunmmary and Conclusions 

The low degree of fertility in the Fi offsprings of Tropisternus col- 
laris mexicanus from Honduras with a melanic form of T. collaris from 
Colombia suggests that these populations have been separated for a 
considerable length of time. The lowered fertility is about of the degree 
that is found in crosses of the other North American forms with the 
South American or West Indian forms. Only one adult was obtained 
from Fi x Fi cross and none from backcrosses with the Colombian 
parent. Outcrosses of hybrids with the melanic T. c. viridis from South 
Carolina indicates, however, that the hybrids are not sterile but that 
their failure to produce viable embryos is probably due to lethal genes 
and possibly translocations which cause death early (or later) in 

Literature Cited 

1. Dancis, Barry M. 1967. Experimental hybridization of an insular form 
of Tropisternus collaris (Fabricus) with mainland subspecies (Coleoptera: 
Hydrophilidae). Proc. Indiana Acad. Sci. 76:279-283. 

2. Sharp, David. 1882. Biologia centrali-americana. Insects. Coleptera 1 (Part 
2), pp. xv + 140, illus. 

3. Young, Frank N. 1965. Hybridization between North and South American 
Tropisternus (Coleoptera: Hydrophilidae). Proc. XII. Intern. Congress 
of Entomology, London, p. 246 

4. Young, Frank N. 1966. The genetic basis of color patterns of aquatic 
beetles of the Tropistermis collaris complex (Coleoptera: Hydrophilidae). 
Proc. North Central Branch, Entomol. Soc. Amer. 20:87-92. 

5. Young, Frank N. 1967. Studies on the color patterns in crosses of Tropis,- 
ternus from western Mexico with other color forms of the Tropisternus col- 
laris complex (Coleoptera: Hydrophilidae). Proc. Indiana Acad. Sci. 76:272- 

An Annotated List of the Spiders of Indiana 1 

Thomas A. Parker, Purdue University 


The history of Indiana araneology began in 1847 when Nicholas 
Marcellus Hentz, the Father of American Araneology, described A galena 
naevia from Indianapolis (29). Since that time many have contributed to 
the list of spiders of Indiana. Notable among these contributions are the 
works of Fox (25), Banks (2), and Elliott (20, 21, 22). 

Dr. William H. Fox published the first list of Indiana spiders (25), 
which was based on a collection of a chemistry professor at Purdue Uni- 
versity, Mr. Frederick C. Test. This list included 79 species of spiders in 
what would today be 16 families, although family names were not used in 
this list. 

Nathan Banks published a preliminary list of Indiana spiders in 
1907 (2). This work included keys to the families and genera of spiders 
and, in addition, a list of species belonging to other arachnid orders. The 
list of spiders included 148 species found in 16 families, two species of 
which were described as new to science. The list was based on spiders 
collected by W. S. Blatchley, Mel T. Cook of Greencastle, W. J. Moenkhaus 
of Huntingburg, and A. M. Banta who collected spiders in Indiana caves. 

In 1930 the late Dr. Frank R. Elliott published on spiders of a beech- 
maple forest near Richmond, Indiana (20). Combining the records of 
Banks with newly collected material and his beech-maple study, Elliott 
published a list of Indiana spiders in 1932 (21). This list contained 218 
species of spiders found in 20 families. 

Elliott again published a list of Indiana spiders in 1953 (22). It was 
based on records in the literature and newly collected material. The work 
of Lowrie (38, 39) on the spiders of the Chicago area dunes greatly con- 
tributed to the records of Indiana spiders. Using these records and 
others, Elliott assembled a list containing 303 species of spiders found in 
21 families. 

No major studies of the spiders of Indiana have been published since 
Elliott's 1953 list. This paper, then, will enlarge, correct, and update the 
list of Indiana spiders. 

The List 

The present list of Indiana spiders contains 378 species of spiders 
found in 29 families. The families are arranged following the system used 
by Kaston (32). The genera are alphabetically arranged in the families 
and likewise the species in the genera. Nomenclature generally follows 

1 Journal Paper No. 3771, Purdue University Agricultural Experiment 


Entomology 267 

that used by Kaston. Synonyms are included beneath the accepted scien- 
tific name in cases where name changes have occurred within the last 
twenty years. The species are numbered consecutively. 

Collection records are arranged by county with the oldest record pre- 
ceding all others. The counties are in bold-faced type. Usually the local- 
ity is given in parenthesis, followed by citations in which the record was 
found. In instances where no locality was given in the literature or where 
recent collectors gave no specific locality on their labels, just the citation 
or date collected and collector's initials are included in the parenthesis. 
If the record is a recent one, the collection dates and collector's initials 
are included. Names of collectors may be found in Appendix I. 

Many of the localities recorded in the older publications no longer 
exist, having been incorporated into neighboring cities. Other names 
appearing in the literature were local names and were not formally recog- 
nized. Appendix II enables the reader to locate such places more accu- 
rately. Also included in this appendix are a few localities visited by the 
author or his colleagues for which a more specific geographical description 
was deemed necessary. 

The writer's spider collection is housed in the Entomology Depart- 
ment at Purdue University. All identifications of recent collections were 
made by the writer. Some verifications were obtained from araneologists 
in the United States and Canada. Such determinations are credited to 
these men individually in the text. This list is based on records from the 
literature and the collection of the writer. This collection represents spi- 
ders collected during a three-year study of spiders along the Wabash and 
Tippecanoe Rivers, spiders sent for determination through the Cooperative 
Extension Service, and specimens collected by students and staff of the 
Entomology Department at Purdue University. 

ORDER Araneida 
Suborder Orthognatha 

family Atypidae 

1. Atypus milberti (Walckenaer), 1837 

Crawford (Wyandotte Cave, Sept. 8, 2, 21, 22); Posey (New 
Harmony, 2, 21, 22); INDIANA (49); Warren (South Pine 
Creek, May 5, 1968, GRF) 

family Ctenizidae 

2. Aptostichus flavipes Petrunkevitch, 1925 

Lawrence (Mitchell, 50, 51, 22) 
Petrunkevitch (51) reported this species in his Catalogue of American 
Spiders, Part 1, not in his synonymic index (49), as Elliott (22) has 

3. Pachylomerides audouini (Lucas), 1836 

Floyd (Floyd's Knobs, 45); Harrison (Corydon, May 12, 1968, 

268 Indiana Academy of Science 

The specimen from Harrison County was sent to the writer by Jack 
Washburn, County Agent. It was found in the driveway at the home of 
Cova Bates. 

Suborder Labidognatha 

family Dysderidae 

4. Dysdera crocata C. L. Koch, 1839 

Whitley (Columbia City, Aug. 28, 1967, RLG); Cass (Logans- 
port, Mar. 30,1967, TH) 

family Segestriidae 

5. Ariadna bicolor (Hentz), 1842 

Tippecanoe (Lafayette, 25, 22); Crawford (Wyandotte, Apr. 17, 
2, 21, 22); Lawrence (Mitchell, Apr. 2, 2, 21, 22); Parke (Mec- 
ca, Apn\ 27, 2, 21, 22); INDIANA (49); Wayne (Richmond, 20, 
21, 22); Steuben (Crooked Lake, 21, 22); Porter (Valparaiso, 
21, 22) 
Elliott (22) listed this species from Wyandotte Cave. The original 
record of Banks (2) was from the town of Wyandotte, not the cave. 

family Scytodidae 

6. Scytodes thoracica (Latrielle), 1804 

Wayne (Richmond, 20, 21, 22); White (Brookston, Apr. 22, 
1968, TAP); Tippecanoe (West Lafayette, May 16, 1968, TAP) 

family Loxoscelidae 

7. Loxosceles reclusa Gertsch & Mulaik, 1940 

Gibson (Princeton, 46); Harrison (Bradford, Oct. U, 1967, 
TAP) ; Posey (Mount Vernon, Jan. 19, 1968, TAP; Poseyville, 
May 23, 1968, TAP); Clark (Jeffersonville, Mar. 28, 1968, 
TAP); Knox (Bruceville, Mar. 18, 1968, RED); Vanderburgh 
(Evansville, May 28, June 3, 1968, TAP); Dubois (Jasper, June 
26,1968, DJM) 

family Pholcidae 

8. Pholcus jihalangioides (Fuesslin) , 1775 

Wayne (Richmond, 20, 21, 22); Steuben (Crooked Lake, 21, 
22); Tippecanoe (West Lafayette, June 20, 1966, TAP) 

9. Spermophora meridionalis Hentz, 1841 

Tippecanoe (Ross Biological Reserve, 5) 

family Theridiidae 

10. Asagena americana Emerton, 1882 

Tippecanoe (Lafayette, 25, 22; Ross Biological Reserve, 5; 
Merritt's Pine Plantation, July 11, 1967, RWM); Crawford 
(Wyandotte, Apr. 17, 2, 21, 22); INDIANA (49); Wayne 
(Richmond, 20, 21, 22) 

11. Conopistha rufa (Walckenaer), 1841 
Argyrodes trigonum (Hentz), 1850 

Monroe (Mayfield's Cave, 4, 22; Twin Cave, 4, 21) 

Entomology 269 

The Twin Cave record of Banks is from a specimen sent to him by 
Banta and probably is Banta's "Lower Twin Cave at Mitchell" which is 
in Lawrence County. 

12. Crustulina altera Gertsch & Archer, 1942 

C. guttata: (Emerton), 1882 (nee Theridion guttatum Wider, 1834) 
Wayne (Richmond, 20, 21, 22, 39); Steuben (Crooked Lake, 21, 
22); Porter (Valparaiso, 21, 22; Ogden Dunes, 39, 22); Tippe- 
canoe (Ross Biological Reserve, 5) 

Elliott (21, 22) listed Crustulina guttata (Wider). Willis J. Gertsch 
considers our species separate from the European guttata (Wider), hence 
the name altera. Kaston (32), page 75, says, "In general appearance and 
structure of the genitalia this species [C. altera] closely resembles gut- 
tata (Wider), with which, in fact, it may be identical, although Dr. 
Gertsch thinks not." 

13. Ctenium pumilus (Emerton), 1909 

Wayne (Richmond, 20, 21, 22) 

14. Ctenium riparius (Keyserling), 1886 

Knox (May 26, 2, 21, 22); INDIANA (49); Wayne (Richmond, 
21, 22); Porter (Ogden Dunes, 39, 22) 

15. Enoplognatha marmorata (Hentz), 1850 

Marion (Indianapolis, 30, 22); Tippecanoe (Lafayette, 25) 

16. Euryopis argentea Emerton, 1882 

Porter (Ogden Dunes, 39, 22; Ind. Dunes State Park, 22); 
Tippecanoe (Ross Biological Reserve, 5) 

17. Euryopis limbata (Walckenaer), 1841 
E. funebris (Hentz), 1850 

Tippecanoe (Lafayette, 25, 22); Posey (New Harmony, June 
4, 2, 21, 22); Wayne (Richmond, 20, 21, 22, 39); LaPorte 
(Smith, 39, 22); Fountain (Attica, Sept. 7, 1967, TAP) 

18. Latrodectus mactans (Fabricius), 1775 

Tippecanoe (Lafayette, 25, 22); Crawford (Wyandotte, Sept. 9, 
2, 21, 22, 35); Lawrence (Mitchell, July 15, 2, 22); Dubois 
( Huntingburg, 2, 21, 22); Jennings (North Vernon, Sept. 14, 
2, 21, 22); Martin (Shoals, Apr. 22, 2, 21, 22, 35); Floyd (New 
Albany, May 5, 2, 21, 22); Clark (Charlestown, HS, 35) 
Banks (2) collected this species at Wyandotte, not in Wyandotte 
Cave, as Elliott (22) has indicated. 

19. Latrodectus variolus Walckenaer, 1837 

Putnam (Greencastle, 2, 21, 22, 42); Porter (Dunes Acres, 38, 
39, 42; Ogden Dunes, 39, 42); Brown (Aug. 10, 1967, May 1, 
1968, JJF) 

Levi (35) synonymized Latrodectus variolus with L. 7nactans. Actu- 
ally what is now known as L. variolus was included in Levi's records of 
L. curacaviensis. McCrone and Levi (42) showed that L. variolus is a 
valid species based on coloration and ecology. Therefore the records 

270 Indiana Academy of Science 

listed for Latrodectus curacavie?isis for Indiana in Levi's paper are 
actually L. variolus and are herein listed as such. 

20. Lithyphantes albomaculatus (DeGeer), 1778 

Porter (Dunes Acres, 38, 39; Ogden Dunes, 39) 

21. Steatoda borealis (Hentz), 1850 

Tippecanoe (Lafayette, 25, 22; Americus, Aug. 1, 1967, TAP) ; 
Putnam (Greencastle, 2, 21, 22); Wayne (Richmond, 20, 21, 22, 
39); Porter (Valparaiso, 21, 22; Ogden Dunes, 39, 22); LaPorte 
(Smith, 39, 22); Benton (Otterbein, Apr. 29, 1968, JOS); 
Pulaski (Winamac, June 16, 1966, Aug. 22, 1967, TAP); Ver- 
million (Clinton, July 1U, 1966, TAP); Fulton (Rochester, Aug. 
29, 1967, TAP) 

22. Teutana grossa (C. L. Koch), 1838 

Tippecanoe (Ross Biological Reserve, 5) 

23. Teutana triangulosa (Walckenaer), 1802 

Tippecanoe (Lafayette, 25, 22; Ross Biological Reserve, 5); 
Marion (Indianapolis, Oct. 15, 1967, TAP) 

24. Theridion alabamense Gertsch & Archer, 1942 

T. cinereum Emerton, 1913, (cinereum preoc. by Thorell, 1875) 

Lake (Miller, 53; Pine, 53); Porter (Ind. Dunes State Park, 53); 
LaPorte (Smith, 39, 22) 

25. Theridion differens Emerton, 1882 

Marshall (Arlington, June 10, 2, 21, 22); Starke (Bass Lake, 
June 17, 2, 21, 22); Wayne (Richmond, 20, 21, 22, 39); Steuben 
(Crooked Lake, 21, 22); Brown (Nashville, 21, 22); Porter 
(Valparaiso, 21, 22; Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 
22; Ind. Dunes State Park, 22); Tippecanoe (Americus, June 23, 

1966, TAP) 

26. Theridion frondeum Hentz, 1850 

Knox (Vincennes, July 10, 2, 21, 22); Posey (Grand Chain, 
June 3, 2, 21, 22; Mount Vernon, Aug. 17, 18, 1966, June 8, 1967, 
July 27, 1967, TAP), INDIANA (49), Lake (Hessville, 53; 
Liverpool, 53); LaPorte (Otis, 53; Smith, 39, 22; LaPorte, June 
19, 1968, TAP); Porter (Woodville, 53; Valparaiso, 21, 22; 
Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22; Ind. Dunes State 
Park, 22); Wayne (Richmond, 20, 21, 22, 39); Tippecanoe (Ross 
Biological Reserve, 5; Americus, Aug. 1, 1967, TAP); Carroll 
(Delphi, July 6, 1966, TAP); Huntington (Huntington, June 
29, 1966, TAP); Fountain (Attica, July 25, 28, 1967, TAP); 
Cass (Logansport, Aug. 3, 1967, TAP); Vermillion (Clinton, 
July 13, Ik, 1966, July 20, 1967, TAP); Sullivan (Merom Sta- 
tion, July 20, 21, 1966, TAP); Gibson (Mount Carmel, July 18, 

1967, TAP) 

27. Theridion glaucescens Becker, 1879 
T. spirale Emerton, 1882 

Lake (Pine, May 25, 2, 53, 21, 22); Porter (Ind. Dunes State 
Park, 53); Tippecanoe (Ross Biological Reserve, 5; Americus, 

Entomology 271 

June 23, 1966, Aug. 1, 1967, TAP); Huntington (Huntington, 
June 29, 1966, Aug. 15, 1967, TAP); Fulton (Rochester, June 
6, 7, 1966, Aug. 29, 1967, TAP) ; Pulaski (Winamac, June 15, 
1966, Aug. 22, 1967, TAP) ; Cass (Logansport, July 6, 1966, 
Aug. 3, 1967, TAP); Vermillion (Clinton, July 20, 1967, TAP); 
Gibson (Mount Carmel, July 18, 1967, TAP) 

28. Theridion lyricum Walckenaer, 1841 
T. lyra Hentz, 1850 

T. kentuckyense Emerton, 1909 

Monroe (Mayfield's Cave, 2, 4, 21, 22; Twin Cave, 2, 21, 22; 
Truett's Cave, 4, 22) ; INDIANA (49) 

29. Theridion murarium Emerton, 1882 

Tippecanoe (Lafayette, 25, 22; West Lafayette, June 18, 1968, 

30. Theridion pennsylvanicum Emerton, 1913 

Wayne (Richmond, 20, 21, 22); Steuben (Crooked Lake, 21, 22) 

31. Theridion porteri Banks, 1897 

Owen (Porter's Cave, 1, 2, 3, 6, 21, 22); Monroe (Truett's Cave, 

1, 2, 6, 4, 21; Mayfield's Cave, 4, 44, 21, 22); INDIANA (49) 

Blatchley (6) receives credit for a Truett's Cave record because he 
states, on page 204, "This species was also noted in the first room of 
Truett's Cave— W.S.B." 

The date of the description of this species should be 1897, not 1896, 
as Banks (2, 3) has indicated. Banks described Theridion porteri in the 
same publication as the description of Tegenaria cavicola. The reader 
is referred to a discussion of the date change under Tegenaria cavicola. 

32. Theridion puncto spar sum Emerton, 1882 

Pulaski (Winamac, June 16, 1966, TAP) 

33. Theridion rupicola Emerton, 1882 

LaPorte (Smith, 39, 22) 

34. Theridion unimaculatum Emerton, 1882 

LaPorte (Smith, 39, 22) 

Elliott (22) copied Lowrie's record incorrectly. The record should 
have been Theridion unimaculatum, not T. albomaculatum as Elliott has 
indicated. T. albomaculatum occurs from Georgia, Texas, and Florida, 
south to Brazil. 

35. Theridion tepidariorum C. L. Koch, 1841 

Marion (Indianapolis, 30, 22); Tippecanoe (Lafayette, 25, 22; 
Americus, June 23, 1966, TAP; West Lafayette, June 20, Aug. 
10, 1966, TAP); Kosciusko (2, 21, 22); Putnam (Greencastle, 

2, 21, 22); Wayne (Richmond, 21, 22); Porter (Valparaiso, 21, 
22); LaPorte (Smith, 39, 22); Brown (Brown Co. State Park, 
Sept. 13, 1966, TAP); Fulton (Rochester, July 27, 1967, TAP) 

36. Theridion zelotypum Emerton, 1882 

Fountain (Attica, Aug. 11,25, 1967, TAP) 

272 Indiana Academy of Science 

37. Theridula opulenta (Walckenaer), 1841 

Wayne (Richmond, 20, 21, 22) 

38. Ulesanis americana Emerton, 1882 

LaPorte (Smith, 39) 

family Nesticidae 

39. Nesticus carteri Emerton, 1875 

Harrison (Bradford Cave?, 23, 47, 48, 2, 21, 22); Owen (Por- 
ter Cave, 6, 2, 21, 22); Monroe (Coon Cave, 6, 2, 21, 22; May- 
field's Cave, 4, 22; Truett's Cave, 4, 22); Crawford (Marengo 
Cave, 6, 2, 21, 22); INDIANA (49) 
The Bradford Cave record for this species will always remain du- 
bious. Emerton (23) says, "A cocoon collected by Mr. Packard, from 
Bradford Cave, Ind., contains young, which had passed their second 
moult, probably of this species." 

Packard (47) states, ". . . while in the Bradford Cave occurred a 

Nesticus thought by Mr. Emerton to be identical with Nesticus Carteri." 

Packard (48) does not accept the Bradford Cave record of Nesticus 

carteri Emerton as a valid one, for on page 83 he lists, "Nesticus carteri 

Emerton, Amer. Naturalist, ix, May, 1875. Carter Caves, Kentucky." 

family Linyphiidae 

40. Bathyphantes canadensis (Emerton), 1882 

Wayne (Richmond, 21, 22) 

41. Bathyphantes concolor (Wider), 1834 

Fulton (Rochester, July 27, Sept. 16, 1967, TAP) 

42. Bathyphantes nigrinus (Westring), 1861 

Tippecanoe (Lafayette, 25, 22); Monroe (Mayfield's Cave, 4, 
22); Wayne (Richmond, 20, 21, 22, 39); Porter (Dunes Acres, 

43. Bathyphantes pallida (Banks), 1892 

Diplostyla nigrina Emerton, 1881 (nee Linyphia nigrina Westring) 
Tippecanoe (Americus, Sept. 7, 1967, July U, 1968, TAP); 
Fountain (Attica, Aug. 11, 1967, TAP); Fulton (Rochester, 
Sept. 16,1967, TAP) 

44. Bathyphantoides brevis (Emerton), 1911 

Fountain (Attica, Aug. 11, 1967, TAP); Fulton (Rochester, 
Sept. 16, 1967, TAP) 

45. Centromerus cornupalpis (O.P.-Cambridge), 1875 

Wayne (Richmond, 20, 21, 22) 

46. Centromerus latidens (Emerton), 1882 

Spencer (Santa Claus, Apr. -4, 1966, RWM) 

47. Florinda coccinea (Hentz), 1850 

Sullivan (Merom Station, July 20, 1966, TAP) 

48. Frontinella pyramitela (Walckenaer), 1841 

LaPorte (Smith, 39, 22); Tippecanoe (Ross Biological Reserve, 

Entomology 273 

5); Brown (Brown Co. State Park, Sept. 13, 1966, TAP); Pu- 
laski (Winamac, Aug. 22, 1967, TAP); Carroll (Pyrmont, July 
28, 1968, RWM) 

49. Lepthyphantes nebulosa (Sundevall), 1830 

Tippecanoe (Americus, Aug. 10, 1967, TAP) 

50. Lepthyphantes zebra (Emerton), 1882 

Wayne (Richmond, 20, 21, 22) 

51. Linyphia clathrata Sundevall, 1830 

Wayne (Richmond, 20, 21, 22, 39); Porter (Ogden Dunes, 39, 
22); LaPorte (Smith, 39, 22) 

52. Linyphia maculata Emerton, 1909 

LaPorte (Smith, 39, 22); Cass (Logansport, July 6, 1966, TAP) 

53. Linyphia marginata C. L. Koch, 1834 

Tippecanoe (Lafayette, 25, 22; Sept. 2, 1966, TAP); Marshall 
(Arlington, June 10, 2, 21, 22); Crawford (Wyondotte, Apr. 17, 
2, 21, 22?); Putnam (Greencastle, 2, 21, 22); Monroe (May- 
field's Cave, 4, 22); INDIANA (49); Wayne (Richmond, 20, 21, 
22, 39); Steuben (Crooked Lake, 21, 22); Porter (Valparaiso, 
21, 22); LaPorte (Smith, 39, 22); Sullivan (Merom Station, 
July 21, 1966, TAP) 
Banks (2) collected this species at the town of Wyandotte, not in 
Wyandotte Cave as Eliott (22) has indicated. 

54. Linyphia weyeri Emerton, 1875 

Crawford (Marengo Cave, 44, 21, 22). 
Emerton (23) and Packard (47, 48) list Linyphia subterranea Emer. 
from Wyandotte Cave, not L. weyeri, as Banks (3) and Elliott (22) have 
indicated. Linyphia weyeri Emer. was described from specimens taken 
in Weyer's Cave, in Augusta County, Virginia. 

55. Meioneta fabr a (Keyserling), 1886 

Spencer (Santa Claus, June 12, 1966, RWM); Tippecanoe 
(Americus, Aug. 2U, 1967, July U, 1968, TAP) 

56. Meioneta micaria (Emerton), 1882 

Wayne (Richmond, 20, 21, 22); Tippecanoe (Americus, Aug. 10, 
24, 1967, TAP) ; Fountain (Attica, Aug. 25, 1967, TAP) ; 
Vanderburgh (Apr. 1, 1966, RWM) 

57. Meioneta unimaculata (Banks), 1892 

Porter (Valparaiso, 21, 22); Tippecanoe (Americus, Aug. 10, 
2U, 1967, TAP); Carroll (Prymont, July 28, 1968, RWM) 

58. Pityohyphantes costatus (Hentz), 1850 

Linyphia phrygiana: Emerton, 1882 (nee L. phrygiana C. Koch, 

Marshall (Culver, June 29, 2, 21, 22); Lake (Hessville, 53; 
Liverpool, 53); Wayne (Richmond, 20, 21, 22, 39); Steuben 
(Crooked Lake, 21, 22); Brown (Nashville, 21, 22); Porter 
(Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22); LaPorte 
(Smith, 39, 22); Putnam (Bainbridge, Apr. 23, 1967, TAP) 

274 Indiana Academy of Science 

Chamberlin and Ivie (11) are convinced that P. costatus is different 
from the European Linyphia phrygiana. They state, on page 28, "This 
common species of the eastern United States is distinct from the Euro- 
pean phrygiana, where it has been consistently placed by American au- 
thors. The above name [P. costatus (Hentz)], therefore, should be used." 

59. Pusillia mandibulata Emerton, 1882 
Linyphia pusilla (of most authors) 

Tippecanoe (Lafayette, 25, 22; Americus, July 27, 1967, TAP); 
Wayne (Richmond, 21, 22); Porter (Dunes Acres, 38, 39, 22); 
Fountain (Attica, Aug. 3, 1966, July 25, 1967, TAP); Hunting- 
ton (Huntington, Aug. 15, 1967, TAP) 

Chamberlin and Ivie (11) say that Linyphia pusilla Sundevall, 1830 
is European and does not occur in North America. They consider their 
Pusillia pusilla distinct from Pusillia mandibulata Emerton, 1882. Ap- 
parently what was formerly known as Linyphia pusilla is now Pusillia 
mandibulata in the East and their new Pusillia bonita in the West. 

GO. Stemonyphantes lineatus (Linnaeus), 1758 

Tippecanoe (Lafayette, 25, 22); Putnam (Greencastle, 2, 21, 
22); Wayne (Richmond, 21, 22) 

61. Tennesseellum formicum (Emerton), 1882 

Porter (Dunes Acres, 38, 39, 22); Fountain (Attica, Aug. 11, 
1967, July Jt, 1968, TAP) 

G2. Troglohyphantes cavernicolus (Keyserling), 1886 

Lawrence (Donnelson's Cave, 4, Nov. 9, 2, 21, 22; Twin Caves 
at Mitchell, 4, 22; Shawnee Cave, 43, 44, 21, 22); INDIANA 

Emerton (23) did not record this species from Wyandotte Cave as 
Elliott (22) has indicated. Mclndoo (44), on page 183, says, "This species 
[Linyphia Weyeri Emer.] in this cave [Marengo Cave] is as abundant as 
Troglohypha?ites ( = Willibaldia) cavernicola Keys, is in the Shawnee 

Elliott (21) lists Mayfield's Cave and Hamer's Cave for this species. 
Elliott (22) credits the Mayfield's Cave record to Banta and drops the 
Hamer's Cave record. Banta (4) did not collect this species in Mayfield's 
Cave, and the writer is certain that a Hamer's Cave collection never 

Elliott (22) should not have included Troglohyphantes incertus Emer. 
in the list of Indiana spiders. Emerton described this spider in 1875 from 
specimens taken in caves in Virginia and Kentucky. The last paragraph 
of the description, which is also included in Packard (48), pages 275-276, 
says, "Fountain Cave, Virginia, among stalactites, in company with 
Nesticus ijallidus (Packard); also in Bat Cave, Carter County, Kentucky 
(Shaler and Packard)." 

family Micryphantidae 

63. Anthrobia sp. 

Crawford (Wyandotte Cave, 18, 22) 

Entomology 275 

The writer is hesitant to include this spider as a record for Indiana. 
Elliott (22) included this spider as A. mammouthia Tellkampf. Cope (18) 
listed specimens from Wyandotte Cave as Anthrobia. He gave these 
specimens no specific name, believing them different from A. mam- 
mouthia, which he had listed from Mammouth Cave, Kentucky. Cope 
gives this account of his collections in Wyandotte Cave: "Two species of 
Arachnidans were observed, one a true spider, the other related to the 
"long-legs" of the woods. A species similar to the former is found in the 
Mammouth Cave [A. mammouthia], and others in other caves, but in 
every instance where I have obtained them, they have been lost by the 
dissolution of their delicate tissues in the impure alcohol." 

Emerton (23) had A. mammouthia listed from Mammouth, Proctor's 
and Diamond Caves, all of which occur in Kentucky. Emerton made no 
mention of collecting A. mammouthia from Wyandotte Cave as Elliott 
(22) has indicated. 

64. Catabrithorax oxypaederotipus (Crosby), 1905 

Wayne (Richmond, 20, 21, 22); Steuben (Crooked Lake, 21, 22); 
Porter (Valparaiso, 21, 22) 

65. Catabrithorax plumosus (Emerton), 1882 

Tippecanoe (Americus, Aug. 24, 1967, TAP) 
This specimen was determined by Wilton Ivie of the American 
Museum of Natural History. 

66. Catabrithorax probatus (Cambridge), 1874 

Wayne (Richmond, 20, 21, 22); Steuben (Crooked Lake, 21, 
22); Porter (Valparaiso, 21, 22) 

67. Ceraticelus emertoni (O.P.-Cambridge), 1874 

Wayne (Richmond, 21, 22); Porter (Dunes Acres, 38, 39, 22; 
Ogden Dunes, 39, 22); LaPorte (Smith, 39, 22); Tippecanoe 
(Ross Biological Reserve, 5) 

68. Ceraticelus fissiceps (O.P.-Cambridge, 1874 

Porter (Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22; Ind. 
Dunes State Park, 22) 

69. Ceraticelus laetus (O.P.-Cambridge), 1874 

Fountain (Attica, July 4, 1968, TAP) 

70. Ceraticelus limnologicus Crosby & Bishop, 1925 

Porter (Dunes Acres, 39, 22; Ogden Dunes, 39, 22) 
Lowrie (39) reported C. limnologicus, not C. homologicus as Elliott 
(22) has indicated. 

71. Ceraticelus minutus (Emerton), 1882 

Wayne (Richmond, 20, 21, 22, 39); Steuben (Crooked Lake, 21, 
22); Porter (Valparaiso, 21, 22; Ogden Dunes, 39, 22) 

72. Ceratinella brunnea Emerton, 1882 

Wayne (Richmond, 20, 21, 22); Steuben (Crooked Lake, 21, 22); 
Porter (Valparaiso, 21, 22) 

73. Ceratinops rugosa (Emerton), 1909 

Tippecanoe (Ross Biological Reserve, 5) 

276 Indiana Academy of Science 

74. Ceratinopsis anglicana (Hentz), 1850 

Porter (Dunes Acres, 38, 39, 22; Ind. Dunes State Park, 22) 

75. Ceratinopsis interpres (O.P.-Cambridge) , 1874 

LaPorte (Otis, 53); Porter (Woodville, 53; Valparaiso, 21, 22); 
Wayne (Richmond, 20, 21, 22); Steuben (Crooked Lake, 21, 22); 
Brown (Nashville, 21, 22) 

76. Cornicularia minuta Emerton, 1882 

Wayne (Richmond, 20, 21, 22) 

77. Eperigone contorta (Emerton), 1882 

Porter (Dunes Acres, 38) 

78. Eperigone indicabilis Crosby & Bishop, 1928 

Tippecanoe (Americus, July 27, 1967, TAP) 
This specimen was determined by Wilton Ivie of the American 
Museum of Natural History. 

79. Eperigone tridentata (Emerton), 1882 

Owen (Porter's Cave, 6, 2, 21, 22); INDIANA (49) 
Nathan Banks determined the specimen sent to him by Blatchley 
from Porter's Cave as Tmeticus tridentatus Emerton. Blatchley (6) 
reported this species in his paper. Banks (2) reported the same species 
from Porter's Cave, putting it in the genus Gonglydmm. Elliott (22) 
should have recorded Oedothorax tridentatus Emer., not bidentatus, as he 
has for number 126. Elliott (21) recorded O. bidentatus synonymizing 
this species with Gonglydium tridentatus Emer. This synonymy is 

80. Eridayites erigonoides (Emerton), 1882 

Spencer (Santa Claus, May 22, 1966, RWM) 

81. Erigone aletris Crosby & Bishop, 1928 

Fountain (Attica, Aug. 16, Sept. 6, 1968, TAP) 

82. Erigone atra Blackwall, 1833 
?E. longipalpis Emer., 1882 

Marshall (Arlington, June 10, 2, 21, 22); INDIANA (49); 
Fountain (Attica, Aug. 11, 1967, TAP) 

83. Erigone autumnalis Emerton, 1882 

Porter (Dunes Acres, 38, 39, 22); Tippecanoe {Aug. 1, 1967, 
RWM); Harrison (May 11, 1966, RWM); Switzerland (May 11, 
23, 1966, RWM); Spencer (Santa Claus, Aug. 21, 1966, RWM); 
Vanderburgh (May 21, June 1, July 1, 21, RWM) 

84. Erigone blaesa Crosby & Bishop, 1928 

Fountain (Attica, July 4, 1968, TAP) 

85. Erigone dentigera O.P.-Cambridge, 1874 
?E. longipalpis Emer., 1882 

Porter (Dunes Acres, 38, 39, 22); Cass (Logansport, Aug. 3, 
1967, TAP) ; Fountain (Attica, July 28, 1967, TAP) 

86. Erigone infernalis Keyserling, 1886 

Monroe (Mayfield's Cave, 4, 43, 44, 21, 22); Lawrence (Twin 
Cave at Mitchell, 4, 2, 21, 22) ; INDIANA (49) 

Entomology 277 

87. Goneatara platyrhiuus (Crosby & Bishop), 1927 

Wayne (Richmond, 20, 21, 22) 

88. Gra?)imonota inomata Emerton, 1882 

Tippecanoe (Lafayette, 25, 22); Wayne (Richmond, 21, 22) 

89. Hypselistes florens (O.P.-Cambridge), 1875 

Porter (Dunes Acres, 38, 39, 22); LaPorte (Smith, 39, 22) 

90. Islandiana longisetosus (Emerton), 1882 

Fountain (Attica, July U, 1968, TAP) 

91. Mythoplastoides exiguus (Banks), 1892 

Wayne (Richmond, 20, 21, 22) 

92. Oedothorax montiferus (Emerton), 1882 

Wayne (Richmond, 20, 21, 22) 

93. Origanates rostratus (Emerton), 1882 

Wayne (Richmond, 20, 21, 22); Steuben (Crooked Lake, 21, 22); 
Warren (South Pine Creek, Apr. 13, 1968, GRF) 

94. Pelecopsis bishopi Kaston, 1945 

Tippecanoe (Ross Biological Reserve, 5) 

95. Phanetta subterranea (Emerton), 1875 

Crawford (Wyandotte Cave, 23, 47, 48, 6, 2, 21, 22; Little 

Wyandotte Cave, 43, 22; Saltpetre Cave, 43, 21, 22); Owen 

(Spring Cave, 43, 22); Monroe (Truett's Cave, 4, 2, 21, 22; 

Mayfield's Cave, 4); INDIANA (49)' 

Mclndoo (43) recorded this species from Spring Cave, not from 

Marengo Cave as Elliott (21) has indicated. Spring Cave is situated % 

mile west of Marengo Cave. 

Emerton (23) recorded this species "under stones in Carter and 
Wyandotte Caves (Packard)", not in Bradford Cave as Elliott (22) had 
indicated. Carter Cave is in Kentucky. 

96. Scylaceus pallidus (Emerton), 1882 

Wayne (Richmond, 20, 21, 22); Steuben (Crooked Lake, 21, 22) 

97. Walckenaera vigilax (Blackwall), 1853 

Tippecanoe (Americus, Aug. 2U, 1967, July 18, 1968, TAP) 

family Araneidae 

98. Acacesia hamata (Hentz), 1847 

Posey (Grand Chain, July 8, 2, 21, 22); Putnam (Greencastle, 2, 
21, 22); INDIANA (49); Wayne (Richmond, 20, 21, 22) 

99. Acanthepeira stellata (Walckenaer), 1841 

Tippecanoe (Lafayette, 25, 22; West Lafayette, Oct. 1U, 1967, 
JEW); Putnam (Greencastle, 2, 21, 22); Knox (Vincennes, July 
10, 2, 21, 22, Apr- 1U, 1968, TAP); Posey (Grand Chain, July 
8, 2, 21, 22; Mount Vernon, Aug. 17, 1966, TAP); Starke (Bass 
Lake, June 20, 2, 21, 22) ; Jennings (North Vernon, Sept. 14, 
2, 21, 22); Porter (Dunes Acres, 38, 39, 22; Ind. Dunes State 
Park, 22); Parke (Aug. 21, 1966, RWM) ; Benton (Otterbein, 
Apr. 25, 1968, JOS) 

278 Indiana Academy of Science 

100. Allepeira lemniscata (Walckenaer), 1841 
Linyphia conferta Hentz, 1850 
Hentzia basilica: McCook, 1878 

Wayne (Richmond, 20, 21, 22); Steuben (Crooked Lake, 21, 22); 
Porter (Valparaiso, 21, 22) 

101. Aranea cavatica (Keyserling), 1881 

Wayne (Richmond, 21, 22) 

102. Aranea miniata (Walckenaer), 1841 

LaPorte (Smith, 39, 22); Wayne (Dalton, Aug. 2, 1967, GLW) 

103. Aranea nordmanni (Thorell), 1870 

Tippecanoe {Aug. 30, 1966, ES) 

104. Aranea solitaria (Emerton), 1884 
A. angulata Clerck, 1757 

Putnam (Greencastle, 2, 21, 22); Wayne (Richmond, 20, 21, 22, 
39); LaPorte (Smith, 39, 22) 

105. Araneus nigripes (Keyserling), 1883 

Posey (Grand Chain, May 12, 2, 21, 22); INDIANA (49) 

106. Araniella displicata (Hentz), 1847 

Tippecanoe (Lafayette, 25); Kosciusko (Tippecanoe Lake, June 
6, 2, 21, 22; Vawter Park, June 1, 2, 21, 22); Posey (Grand 
Chain, June 3, 2, 21, 22); Martin (Shoals, May 16, 2, 21, 22); 
Fountain (Attica, June 20, 2, 21, 22); Porter (Dunes Acres, 38, 
39); LaPorte (Smith, 39); Warren (Pine Village, June 1, 
1968, TH) 

107. Argiope aurantia Lucas, 1833 

Tippecanoe (Lafayette, 25, 22); Whitley? (Round and Shriner 
Lake, 55); Putnam (Greencastle, 2, 21, 22); Marshall (Lake 
Maxinkaukee, Aug. 7, 2, 21, 22); Wayne (Richmond, 21, 22); 
Brown (Nashville, 21, 22); Steuben (Crooked Lake, 21, 22); 
Porter (Valparaiso, 21, 22; Dunes Acres, 38, 39, 22; Ind. Dunes 
State Park, 22); Pulaski (Winamac, Aug. 22, 1967, TAP) 

E. B. Williamson (55) was probably referring to this species when 
he stated on page 155, "In the webs of a species of large black and yellow 
spider ( A rgiope) I have found the remains [of dragonfl ies] ." 

108. Argiope trifasciata (Forkstal), 1775 

Tippecanoe (Lafayette, 25, 22); Putnam (Greencastle, 2, 22); 
Lake (Clark, 53; Miller, 53); Porter (Ind. Dunes State Park, 
53, 22; Valparaiso, 21, 22; Dunes Acres, 38, 39, 22); Wayne 
(Richmond, 21, 22); Brown (Nashville, 21, 22); Fountain 
(Attica, Sept. 7, 1967, TAP) 

109. Cyclosa turbinata (Walckenaer), 1841 
C. caudata (Hentz), 1850 

Tippecanoe (Lafayette, 25, 22); Putnam (Greencastle, 2, 22); 
Porter (Dunes Acres, 38, 39, 22; Ind. Dunes State Park, 22) 

110. Epeira dumetorwm (Villers), 1789 

Tippecanoe (Ross Biological Reserve, 5); Posey (Mount Vernon, 

Entomology 279 

Aug. 18, 1966, TAP); Sullivan (Merom Station, July 21, 1966; 
TAP); Gibson (Mount Carmel, July 27, 28, 1966, July 12, 1967, 
TAP); LaPorte (LaPorte, June 19, 1968, TAP); Vermillion 
(Clinton, July 20, 1967, TAP); Fulton (Rochester, Aug. 29, 

1967, TAP) 

111. Epeira foliata (Fourcroy), 1785 

Tippecanoe (Lafayette, 25, 22; June 4, 1967, TAP); Kosciusko 
(Tippecanoe Lake, Aug. 17, 2, 21, 22; Vawter Park, June 1, 2, 
21, 22); Knox (Vincennes, Nov. 26, 2, 21, 22; July 2, 2, 21, 22); 
Marshall (Arlington, June 10, 2, 21, 22; Culver, June 29, 2, 21, 
22); Posey (Grand Chain, July 8, 2, 21, 22; Mount Vernon, June 
8, 1967, TAP); Putnam (Greencastle, 2, 21, 22); Clark (Jeffer- 
sonville, June 23, 2, 21, 22); Wayne (Richmond, 21, 22); Porter 
(Dunes Acres, 38, 39, 22); Huntington (Huntington, June 28, 
29, 1966, Aug. 15, 1967, TAP); Fulton (Rochester, June 7, 1966, 
TAP); Gibson (Mount Carmel, July 27, 28, 1966, Sept. 29, 1967, 
TAP); Sullivan (Merom Station, July 20, 21, 1966, TAP); 
Brown (Brown Co. State Park, Sept. 13, 1966, TAP); Franklin 
(Brookville, Aug. 30, 1966, TAP); Benton (Otterbein, Apr. 22, 

1968, JOS); LaPorte (LaPorte, June 19, 1968, TAP); Pulaski 
(Winamac, Aug. 22, 1967, TAP) 

112. Epeira pegnia Walckenaer, 1841 

Tippecanoe (Lafayette, 25, 22); Putnam (Greencastle, 2, 21, 22) 

113. Epeira raji (Scopoli), 1763 

Tippecanoe (Lafayette, 25, 22); Knox (Vincennes, Aug. 23, 2, 
21, 22); Lawrence (Mitchell, July 13, 2, 21, 22); Parke (Monte- 
zuma, Aug. 12, 2, 21, 22); Marion (May 30, 2, 21, 22); Putnam 
(Greencastle, 2, 21, 22); Wayne (Richmond, 20, 21, 22, 39); 
Steuben (Crooked Lake, 21, 22); Porter (Woodville, 53; Val- 
paraiso, 21, 22; Ogden Dunes, 39, 22); Brown (Nashville, 21, 
22; Brown Co. State Park, Sept. 13, 1966, TAP); LaPorte (Otis, 
53; Smith, 39, 22); Lake (Hessville, 53; Liverpool, 53) 

114. Epeira thaddeus Hentz, 1847 

Putnam (Greencastle, 2, 21, 22); Porter (Dunes Acres, 38, 39, 
22; Ind. Dunes State Park 22); LaPorte (Smith, 39, 22) 

115. Epeira trifolium Hentz, 1847 

Tippecanoe (Lafayette, 25, 22); Marshall (Culver, Aug. 22, 2, 
21, 22); Putnam (Greencastle, 2, 21, 22); Wayne (Richmond, 
21, 22); Steuben (Crooked Lake, 21, 22); Porter (Valparaiso, 
21, 22; Dunes Acres, 38, 39, 22; Ind. Dunes State Park, 22); 
LaPorte (Smith, 39, 22) 

116. Epeira undata (Olivier), 1787 

Tippecanoe (Lafayette, 25, 22); Putnam (Greencastle, 2, 21, 
22) ; Cass (Logansport, Jidy 15, 1968, TH) 

117. Eustala anastera (Walckenaer), 1841 

Tippecanoe (Lafayette, 25, 22; Americus, Aug. 1, 1967, TAP); 
Kosciusko (Tippecanoe Lake, June 6, 2, 22); Vawter Park, June 

280 Indiana Academy of Science 

1, 2, 22) ; Lake (Hammond, June 16, 2, 22; Pine, June 29, 2, 22; 
Miller, 39); Knox (Vincennes, July 10, Aug. 23, 2, 22; July 2, 

2, 22); Marshall (Arlington, June 10, 2, 22; Culver, June 29, 
2, 22); Starke (Bass Lake, June 20, 22, 2, 22); Crawford 
(Wyandotte, June 25, 2, 22); Posey (Grand Chain, July 8, 
June 3, 2, 22; Mount Vernon, Aug. 18, 1966, TAP); Fountain 
(Attica, June 20, 2, 22, Aug. 3, 1966, July 25; Aug. 11, 1967, 
TAP); Wayne (Richmond, 20, 21, 22, 39); Porter (Valparaiso, 
21, 22; Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22; Ind. Dunes 
State Park, 22); LaPorte (Smith, 39, 22); Sullivan (Merom 
Station, July 20, 21, 1966, July 18, 1967, TAP); Gibson (Mount 
Carmel, July 27, 28, 1966, TAP); Vermillion (Clinton, July lh, 

1966, July 20, 1967, TAP) ; Fulton (Rochester, Aug. 29, 1967, 

118. Eustala cepina (Walckenaer), 1841 

Carroll (Delphi, Aug. 3, 1967, TAP), Sullivan (Merom Station, 
July 20, 1966, July 18, 1967, TAP), Gibson (Mount Carmel, 
July 27, 1966, TAP), Fountain (Attica, Aug. 3, 1966, July 25, 

1967, TAP); Posey (Mount Vernon, Aug. 18, 1966, TAP) 

These specimens were determined by Wilton Ivie of the American 
Museum of Natural History. 

119. Mangora gibberosa (Hentz), 1847 

Wayne (Richmond, 20, 21, 22, 39); Brown (Nashville, 21, 22); 
Porter (Valparaiso, 21, 22; Dunes Acres, 38, 39, 22; Ogden 
Dunes, 39, 22; Ind. Dunes State Park, 22), LaPorte (Smith, 39, 
22); Tippecanoe (Ross Biological Reserve, 5; Ent. Research 
Area, July 2U, 1968, MKP) 

120. Mangora ornata (Walckenaer), 1841 

M. maculata (Keyserling), 1865 

Lake (Hessville, 53; Liverpool, 53); Porter (Dunes Acres, 38, 
39, 22; Ogden Dunes, 39, 22); LaPorte (Smith, 39, 22); Tippe- 
canoe (Ross Biological Reserve, 5); Carroll (Pyrmont, July 28, 

1968, RWM) 

121. Mangora placida (Hentz), 1847 

Crawford (Wyandotte, June 26, 2, 21, 22); Posey (New Har- 
mony, May 6, 2, 21, 22); Sullivan (Merom Station, July 18, 
1967, TAP) 

122. Mastophora bisaccata (Emerton), 1885 

Wayne (Richmond, 20, 21, 22); Tippecanoe (Lafayette, Sept. 
29, 1967, GL) 

123. Mastophora cornigera (Hentz), 1850 

Tippecanoe (Lafayette, 25, 22), Putnam (Greencastle, 2, 21, 22) 

124. Met a menardi (Latrielle), 1804 

Tippecanoe (Lafayette, 25, 22; Ross Biological Reserve, 5); 
Monroe (Mayfield's Cave, 6, 2, 4, 21, 22; Strong's Cave, 6, 2, 
21, 22; Saltpetre Cave, 6, 2, 44; Coon Cave, Sept. 14, 1967, 
TAP); Lawrence (Donnehue's Cave, 6, 2, 22; Donnelson's Cave, 

Entomology 281 

6, 4, 2, 21, 22; Mitchell, «/*% 15, 2, 21, 22; Lower Twin Cave 
at Mitchell, 2, 4, 22; Shawnee Cave, 43, 22); Bartholomew 
(Clifty Cave, 6, 2, 21, 22); Crawford (Wyandotte Cave, 6, 2, 21, 
22; Saltpetre Cave, 6, 2, 21, 22); Putnam (Greencastle, 2, 21, 
22); Owen (Spring Cave, 44, 22); Porter (Dunes Acres, 38) 

Elliott (22) states on page 312 that Emerton (23) collected Met a 
menardi from Bradford and Wyandotte Caves. This inclusion is erroneous. 
Emerton (23), page 278, says, 'Two species were found only about the 
mouths of caves. These are Theridion vulgare Hentz, a spider found 
all over the country in shady places, and a large species of Meta, which 
has been found in similar situations in Massachusetts and New Hamp- 
shire, and resembles Epeira fusca Blackwall." 

Emerton did not say near which caves these spiders were found, nor 
did he positively identify these spiders as Meta menardi (Latrielle), 1804. 

Banta (4) did not report Meta menardi from Strong's Cave, as 
Banks (2) and Elliott (22) have indicated. 

125. Metepeira lahyrinthea (Hentz), 1847 

Putnam (Greencastle, 2, 21, 22); LaPorte (Smith, 39, 22) 

126. Micrathena gracilis (Walckenaer), 1841 

Marion (Indianapolis, 30, 22); INDIANA (40); Tippecanoe 
(Lafayette, 25, 22; Ross Biological Reserve, 5); Crawford 
(Wyandotte, Sept. 9, 2, 21); Putnam (Greencastle, 2, 21); Lake 
(Hessville, 53; Liverpool, 53); Wayne (Richmond, 20, 21, 22); 
Porter (Valparaiso, 21, 22) 

Lindsey, et al. (37) state that Micrathena agilis Walckenaer is found 
on islands in the Wabash River. The name agilis has never been used 
for a North American member of this genus. Apparently M. gracilis 
was collected, not M. agilis. 

127. Micrathena mitrata (Hentz), 1850 

Putnam (Greencastle, 2, 22); Brown (Brown Co. State Park, 
Sept. 13, 1966, TAP) 

128. Micrathena sagittate (Walckenaer), 1841 

Marion (Indianapolis, 30, 22); Tippecanoe (Lafayette, 25, 22); 
Crawford (Wyandotte, June 26, July 25, 2, 21, 22); Lawrence 
(Mitchell, July 15, 2, 21, 22); Putnam (Greencastle, 2, 21, 22); 
Knox (Vincennes, July 10, 2, 21); Lake (Hessville, 53; Liver- 
pool, 53); Wayne (Richmond, 20, 21, 22, 39); Porter (Valpa- 
raiso, 21, 22; Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22; 
Ind. Dunes State Park, 22); LaPorte (Smith, 39, 22) 

129. Neoscona arabesca (Walckenaer), 1841 

Tippecanoe (Lafayette, 25, 22, Sept. 24, 1966, ES); Kosciusko 
(Tippecanoe Lake, June 6, 8, 2, 21, 22); Lake (Hammond, May 
29, July 30, 2, 21, 22); Knox (Vincennes, Aug. 23, 2, 21, 22); 
Posey (Arlington, June 10, 2, 21, 22; Mount Vernon, Aug. 30, 
1966, TAP); Crawford (Wyandotte, Sept. 9, 2, 21, 22); Starke 
(Bass Lake, June 17, 20, 22, 2, 21, 22); Marshall (Culver, June 
29, 2, 21, 22); Putnam (Greencastle, 2, 21, 22); LaPorte 

282 Indiana Academy of Science 

(Wilders, July 25, 2, 21, 22; Smith, 39, 22); Wayne (Rich- 
mond, 20, 21, 22, 39); Steuben (Crooked Lake, 21, 22); Porter 
(Valparaiso, 21, 22; Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 
22; Ind. Dunes State Park, 22); Brown (Brown Co. State Park, 
Sept. 13, 1966, TAP); Carroll (Delphi, Sept. 25, 1966, TAP); 
Benton (Otterbein, Apr. 25, 1968, JOS); Fulton (Rochester, 
June 6, 1966, Aug. 29, 1967, TAP); Fountain (Attica, July 25, 
1967, TAP) 

130. Neoscona benjamina (Walckenaer), 1841 

Tippecanoe (Lafayette, 25; Sept. 26, 1966, JDH); Posey (Grand 
Chain, July 8, 2, 21; Mount Vernon, Aug. 18, 1967, TAP); 
Dubois (Huntingburg, 2, 21); Putnam (Greencastle, 2, 21); 
Lake (Miller, 53, Pine 53; Hessville, 53; Liverpool, 53); Porter 
(Woodville, 53; Ind. Dunes State Park, 53; Dunes Acres, 38, 
39); LaPorte (Otis, 53: Smith, 39); Brown (Brown Co. State 
Park, Sept. 13, 1966, TAP); Pulaski (Winamac, Aug. 22, 1967, 
TAP); Benton (Otterbein, Apr. 29, 1968, JOS) 

131. N eoscona pratensis (Hentz),1847 

Porter (Dunes Acres, 38, 39, 22; Ind. Dunes State Park, 22) 

132. Nephila clavipes (Linnaeus), 1758 

Wayne (Richmond, 21, 22) 

133. Shiga hentzi Banks, 1907 

Perry (Cannelton, May 31, 2, 3, 21, 22); INDIANA (49) 

134. Singa variabilis Emerton, 1884 

Kosciusko (Tippecanoe Lake, 2, 21, 22) 

135. Verrucosa arenata (Walckenaer), 1841 

Tippecanoe (Lafayette, 25, 22); Knox (July 2, 2, 21, 22); Vin- 
cennes, July 10, 2, 21, 22); Parke (Montezuma, Aug. 12, 2, 21, 
22); Crawford (Wyandotte, Sept. 9, 2, 21, 22); Putnam (Green- 
castle, 2, 21, 22); Wayne (Richmond, 20, 21, 22); Steuben 
(Crooked Lake, 21,22) 

136. Wixia anaglyphe (Walckenaer), 1841 
W. ectypa (Walckenaer), 1841 

Posey (Grand Chain, June 3, 2, 21, 22); Wayne (Richmond, 20, 
21, 22) 
Walckenaer's description of W. anaglyphe came one page before 
W. ectypa of the same publication. 

family Theridiosomatidae 

137. Theridiosoma gemmosum (L. Koch), 1877 

Monroe (Mayfield's Cave, 2, 4, 21, 22); LaPorte (Smith, 39, 

family Tetragnathidae 

138. Leucage venusta (Walckenaer), 1841 
L. hortorum (Hentz), 1847 

Tippecanoe (Lafayette, 25, 22; Ross Biological Reserve, 5); 
Posey (Arlington, June 10, 2, 21, 22; Grand Chain, May 12, 

Entomology 283 

2, 21, 22); Putnam (Greencastle, 2, 21 ?, 22; Bainbridge, Apr. 
23, 1967, TAP); Wayne (Richmond, 20, 21, 22, 39); Steuben 
(Crooked Lake, 21, 22); Porter (Valparaiso, 21, 22; Dunes 
Acres, 38, 39, 22; Ogden Dunes, 39, 22; Ind. Dunes State Park, 
22); LaPorte (Smith, 39, 22) 
Banks (2) collected this species at Greencastle, not New Castle, as 
Elliott (21) has indicated. 

139. Mimognatha foxi (McCook), 1894 

Posey (Mount Vernon, June 8, 1967, TAP); Fountain (Attica, 
Aug. 11, 25, 1967, TAP); Tippecanoe (Lafayette, Oct. 23, 1966, 
Wilton Ivie of The American Museum of Natural History deter- 
mined the Mount Vernon collection. 

140. Pachygnatha brevis Keyserling, 1883 

Cass (Logansport, Apr. 6, 1967, TAP) 
Pachygnatha tristriata and P. brevis are not synonymous, as Elliott 
(22) has indicated. 

141. Pachygnatha tristriata C. L. Koch, 1845 

Tippecanoe (Lafayette, 25, 22); Posey (New Harmony, June U, 
2, 21, 22); INDIANA (49); Wayne (Richmond, 21, 22); Knox 
(Oct. 31, 1966, RWM) 

142. Pachygnatha xanthostoma C. Koch, 1845 

Wayne (Richmond, 21, 22) 

143. Tetragnatha banksi McCook, 1894 
T. seneca Seeley, 1928 

Fulton (Rochester, June 7, 1966, Aug. 29, 1967, TAP); Sullivan 
(Merom Station, July 20, 21, 1966, TAP); Posey (Mount Ver- 
non, Aug. 18, 1966, TAP); Pulaski (Winamac, Aug. 22, 1967, 
TAP); Sullivan (Clinton, July 20, 1967, TAP); Gibson (Mount 
Carmel, July 27, 1966, July 12, 1967, TAP) 

After a thorough examination of paratype material of Tetragnatha 
guatemalensis 0. P. Cambridge, 1889, the writer is convinced that T. 
banksi and T. guatemalensis are not synonymous, as Chickering (13, 
14), has concluded. The writer reviewed McCook (41), Levi and Field 
(36), 0. P. Cambridge (8), F. P. Cambridge (7), Seeley (52), and 
Chickering (13, 14, 15) in reaching the above conclusions. The writer 
feels certain that T. seneca Seeley is a synonym of T. banksi McCook, 

Both T. banksi and T. guatemalensis occur in the United States. 
Tetragnatha guatemalensis reaches a few southern states (as e.g., 
Chamberlin and Ivie, [10] and possibly McCook's specimens from Flor- 
ida). Tetragnatha banksi is more northern in distribution (as e.g., 
Seeley's collections, Levi and Field's collections, the writer's collections 
and McCook's specimen from Wisconsin). 

The "large tooth" on the male chelicera of T. guatemalensis is al- 
ways missing in males of T. banksi. Chickering (13) says the "large 
tooth" of T. guatemalensis males is not well developed and that of T. 

284 Indiana Academy of Science 

banksi is a reduced condition. The writer has found and D. J. Clark 
of the British Museum reaffirms that the "large tooth" is quite strong 
in the male of T. guatemalensis. 

Since Tetragnatha guatemalensis is found in the United States, a 
series of "large tooth" reductions should be found in males as one travels 
from south to north if one wishes to synonymize this species with T. 
banksi. The writer can find no evidence of such a cline. 

144. Tetragnatha elongata Walckenaer, 1841 
T. grallator Hentz, 1847 

Tippecanoe (Lafayette, 25, 22; Ross Biological Reserve, 5; 
Americus, June 23, 1966, Aug. 1, 1967, TAP); Kosciusko (2, 
21, 22; Tippecanoe Lake, June 6, 8, 2, 21, 22; Winona Lake, 33, 
22); Posey (Arlington, June 10, 2, 21, 22; Mount Vernon, Aug. 
18, 1966, June 8, 1967, TAP); Dubois (Huntingburg 2, 21, 22); 
Wayne (Richmond, 21, 22); Steuben (Crooked Lake, 21, 22); 
Porter (Valparaiso, 21, 22); Brown (Brown Co. State Park, 
Sept. 13, 1966, TAP); Fulton (Rochester, June 6, 7, 1966, Aug. 
29, 1967, TAP); Cass (Logansport, July 6, 7, 1966, Aug. 3, 
1967, TAP); Carroll (Delphi, June 23, 1966, TAP); Vermillion 
(Clinton, July 13, U, 1966, July 20, 1967, TAP); Gibson (Mt. 
Carmel, July 27, 28, 1966, July 12, 1967, TAP); Pulaski (Wina- 
mac, June 15, 16, 1966, Aug. 22, 1967, TAP); Sullivan (Merom 
Station, July 20, 21, 1966, July 18, 1967, TAP); Huntington 
(Huntington June 28, 29, 1966, Aug. 15, 1967, TAP); Fountain 
(Attica, Aug. 3, 1966, July 25, Aug. 25, 1967, TAP) 

145. Tetragnatha extensa (Linnaeus), 1758 
T. rusticana Chickering, 1959 

Kosciusko (Tippecanoe Lake, June 6, 8, 2, 21, 22; Winona Lake, 
33); Knox (July 2, 2, 21, 22; Vincennes, July 10, 2, 21, 22); 
Posey (Grand Chain, July 8, 2, 21, 22); Marshall (Culver, 
June 29, 2, 21, 22) 

Dr. A. M. Chickering has informed this writer that his T. rusticana 
is generally considered a synonym of T. extensa, a good description of 
which may be found in Chickering (14). 

146. Tetragnatha laboriosa Hentz, 1850 

Tippecanoe (Lafayette, 25, 22; Ross Biological Reserve, 5; 
Americus, June 23, 1966, TAP); Kosciusko (Tippecanoe Lake, 
June 6, 2, 21, 22; Vawter Park, June 1, 2, 21, 22; Winona Lake, 
33, 22); Lake (Hammond, May 29, June 16, July 30, 2, 21, 22; 
Pine, May 25, 2, 21, 22); Posey (Arlington, June 10, 2, 21, 22; 
Grand Chain, June 3, 2, 21, 22; Mount Vernon, June 8, 1967, 
TAP); Crawford (Wyandotte, June 25, 2, 21, 22); Lawrence 
(Mitchell, Apr. 2, 2, 21, 22; Twin Caves, 4, 22; Donnelson's 
Cave, 4, 22); Putnam (Greencastle, 2, 21, 22); Fountain (Attica, 
June 20, 2, 21, 22, Aug. 3, 1966, July 25, Aug. 11, 25, Sept. 7, 
1967, TAP); LaPorte (Wilders, July 25, 2, 21, 22; Smith, 39, 
22); Brown (Nashville, 21, 22); Steuben (Crooked Lake, 21, 
22); Wayne (Richmond, 20, 21, 22, 39); Porter (Dunes Acres, 

Entomology 285 

38, 39, 22; Ogden Dunes, 39, 22; Ind. Dunes State Park 22); 
Sullivan (Merom Station, July 20, 21, 1966, July 18, 1967, 
TAP); Pulaski (Winamac, Aug. 22, 1967, TAP); Carroll (Del- 
phi, Aug. 3, 1967, TAP); Vermillion (Clinton, July 13, 14, 
1966, July 20, 1967, TAP); Cass (Logansport, July 6, 1966, 
TAP) ; Gibson (Mount Carmel, July 27, 28, 1966, TAP) ; 
Fulton (Rochester, June 6, 7, 1966, TAP); Huntington (Hunt- 
ington, Aug. 15, 1967, TAP) 
Banks (2) did not report a separate collection from Lake County 
as Elliott (22) has indicated. 

147. Tetragnatha pallescens F.O.P.-Cambridge, 1903 

Kosciusko (Winona Lake, 33, 22); Porter (Dunes Acres, 38, 39, 
22); Posey (Mount Vernon, Aug. 18, 1966, June 8, 1967, TAP); 
Sullivan (Merom Station, July 20, 21, 1966, TAP); Fountain 
(Attica, Aug. 3, 1966, July 25, 1967, TAP); Gibson (Mount 
Carmel, July 28, 1966, TAP); Cass (Logansport, Aug. 3, 1967, 
TAP) ; Vermillion (Clinton, July 20, 1967, TAP) 

148. Tetragnatha straminea Emerton, 1884 

Kosciusko (Tippecanoe Lake, June 6, 2, 21, 22; Winona Lake, 
33, 22); INDIANA (49); Porter (Dunes Acres, 38, 39, 22; Ind. 
Dunes State Park, 22); LaPorte (Smith, 39, 22); Tippecanoe 
(Ross Biological Reserve, 5) 

149. Tetragnatha vermiformis Emerton, 1884 

Kosciusko (Winona Lake, 33, 22); Tippecanoe (Ross Biological 
Reserve, 5) 

150. Tetragnatha veriscolor Walckenaer, 1841 

Tippecanoe (Ross Biological Reserve, 5; Americus, June 23, 
1966, TAP); Sullivan (Merom Station, July 20, 21, 1966, TAP); 
Huntington (Huntington, June 28, 29, 1966, Aug. 15, 1967, 
TAP); Cass (Logansport, July 6, 1966, July 25, Aug. 3, 1967, 
TAP); Fulton (Rochester, June 6, 7, 1966, TAP); Fountain 
(Attica, Aug. 3, 1966, TAP) ; Pulaski (Winamac, June 16, 1966, 
TAP); Vermillion (Clinton, July 20, 1967, TAP); Posey 
(Mount Vernon, June 7, 8, 1967, TAP) 

family Ctenidae 

151. Anahita animosa (Walckenaer), 1837 

Crawford (Wyandotte, Apr. 17, Sept. 8, 2, 21, 22), INDIANA 

family Mimetidae 

152. Ero furcata (Villers), 1789 

Tippecanoe (Lafayette, 25, 22); Wayne (Richmond, 20, 21, 22, 
39) ; Porter (Ogden Dunes, 39, 22) 

153. Mimetus dissimulatus (Walckenaer), 1841 
M. interfector Hentz, 1850 

Martin (Shoals, May 16, 2, 21, 22); Porter (Dunes Acres, 38, 
39; Ogden Dunes, 39; Ind. Dunes State Park, 22); LaPorte 
(Smith, 39) 

286 Indiana Academy of Science 

154. Mimetus puritanus Chamberlin, 1923 

Sullivan (Merom Station, July 20, 1966, TAP) 
Wilton Ivie of The American Museum of Natural History determined 
the writer's specimen. 

family Agelenidae 

155. Agelenopsis naevia (Walckenaer), 1841 

Marion (Indianapolis, 29, May 30, 2, 21, 22); Tippecanoe (La- 
fayette, 25, 22); Kosciusko (Tippecanoe Lake, Aug. 17, 2, 21, 
22); Knox (Vincennes, Aug. 31, 2, 21, 22); Jennings (North 
Vernon, Sept. H, 2, 21, 22); Floyd (New Albany, Sept. 11, 2, 
21, 22); Steuben (Clear Lake, Aug. 13, 2, 21, 22; Crooked Lake, 

21, 22); Putnam (Greencastle, 2, 21, 22); Dubois (Huntingburg, 
2, 21, 22, Aug. 17, 18, 1967, ES); Wayne (Richmond, 20, 21, 
22); Brown (Nashville, 21, 22; Brown Co. State Park, Sept. 13, 
1966 TAP); Porter (Valparaiso, 21, 22); Fulton (Rochester, 
Sept. 16, 1967, TAP) 

Agelenopsis pennsylvanica is not a synonym of A. naevia as Elliott 
(22) has indicated. 

156. Agelenopsis pennsylvanica (C. L. Koch), 1843 

Porter (Dunes Acres, 38, 39); LaPorte (Smith, 39); Tippe- 
canoe (Lafayette, Sept. 15, 1966, TAP; West Lafayette, Oct. 

22, 1966, TAP) 

Lowrie (39) reported that Elliott (20) had collected A. pennsylvanica 
from Richmond. Elliott (20, 21) collected A. naevia, not A. pennsyl- 
vanica from Richmond. 

157. Agelenopsis utahana (Chamberlin & Ivie), 1933 

Porter (Ogden Dunes, 39, 22); LaPorte (Smith, 39, 22) 

158. Auchicybaeus ovalis Gertsch, 1933 

Crawford (Marengo Cave, 26) 
Gertsch (26), page 12, states, ''TYPE LOCALITY— Female holotype 
from Marengo Spring Cave, Crawford County, Indiana, October 20, 1911, 
taken in the 'area of total darkness' by Arthur W. Henn." 

159. Circurina arcuata Keyserling, 1887 

Tippecanoe (Lafayette, 25, 22; Merritt's Pine Plantation, July 
26, 1968, RWM) ; Starke (Bass Lake, Apr. 9, 2, 21, 22) ; Wayne 
(Richmond, 20, 21, 22); Porter (Valparaiso, 21; Ind. Dunes 
State Park, 21, 22); Warren (S. Pine Creek, Apr. 13, May 5, 
1968, GRF); Adams {Apr. 16, 1968, TAP) 

A very closely related species, C. robusta Simon, 1886, occurs in the 
Rocky Mountains. Cicurina arcuata has chevrons on its abdomen, where- 
as C. robusta has an immaculate abdomen. 

160. Cicurina brevis (Emerton), 1890 

Starke (Bass Lake, Apr. 9, 2, 21, 22); INDIANA (49); Wayne 
(Richmond, 20, 21, 22, 39); Porter (Dunes Acres, 39, 22; Ind. 
Dunes State Park, 22); LaPorte (Smith, 39, 22) 

Entomology 287 

161. Circurina pallida Keyserling, 1887 

Starke (Bass Lake, Apr. 19, 2, 21, 22); Putnam (Apr. 22, 2, 
21, 22; Bainbridge, Apr. 23, 1967, TAP); Crawford (Wyandotte, 
Aug. 17, 2, 21, 22?) ; Monroe (Mayfield's Cave, 4, 22; Truett's 
Cave, 4, 22); INDIANA (49); Wayne (Richmond, 20, 21, 22, 
39) ; LaPorte (Smith, 39, 22) 
Banks (2) collected this species at Wyandotte, not in Wyandotte 
Cave, as Elliott (22) has indicated. 

162. Coras juvenilis (Keyserling) , 1881 

LaPorte (Smith, 39, 22); Putnam (Bainbridge, Apr. 23, 1967, 

163. Coras lamellosus (Keyserling), 1887 

Porter (Ogden Dunes, 39, 22); LaPorte (Smith, 39, 22); Put- 
nam (Bainbridge, Apr. 23, 1967, TAP); Tippecanoe (Merritt's 
Pine Planation, Apr. 10, 1968, RWM) 

164. Coras medicinalis (Hentz), 1821 

Tippecanoe (Lafayette, 25, 22); Putnam {Apr. 22, 2, 21, 22); 
Wayne (Richmond, 20, 21, 22) 

165. Coras mont anus (Emerton), 1890 
Coelotes montanus Emerton, 1890 

Wayne (Richmond, 20, 21, 22) 

166. Tegenaria cavicola Banks, 1897 

Crawford (Saltpetre Cave, 1, 2, 3, 6, 21, 22); INDIANA (49) 
Mclndoo (43, 44) reported Willibaldia (== Troglohyphantes) cav- 
ernicola Keyserling from Shawnee Cave, not Tegenaria cavicola Banks, 
as Elliot (22) has indicated. 

Banks (2, 3) lists the date of his description of Tegenaria cavicola 
as 1896. The date should be 1897, for the 21st Annual Report of the 
Indiana Dept. of Geology and Natural Resources for 1896 was not 
published until 1897. The title page states, "Indianapolis: 1897." 

167. Tegenaria derhami (Scopoli), 1763 

Tippecanoe (Lafayette, 25, 22); Crawford (Wyandotte, Sept. 9, 
2, 21, 22); Monroe (Mayfield's Cave, 4, 22; Twin Cave?, 2, 
21, 22), Putnam (Greencastle, 2, 21, 22); Wayne (Richmond, 
21, 22); Steuben (Crooked Lake, 21, 22); Porter (Valparaiso, 
Banks (2) collected this species at Wyandotte, not in Wyandotte 
Cave, as Elliott (22) has indicated. 

Banks (2) and Elliott (21, 22) report that Banta (4) collected this 
species from Twin Cave. Banta says nothing about a Twin Cave collec- 
tion. Banks probably means Banta's Lower Twin Cave at Mitchell, 
which is in Lawrence County. Apparently Banta had done some collect- 
ing around Mitchell, Indiana, and did not mention many of these collec- 
tions when he reported on Mayfield's Cave in Monroe County. 

168. Wadotes calcaratus (Keyserling), 1887 

Tippecanoe (Lafayette, 25, 22); Posey (New Harmony, Sept. 3, 
2, 21, 22), INDIANA (49); Wayne (Richmond, 20, 21, 22); La- 

288 Indiana Academy of Science 

Porte (Smith, 39, 22); Putnam (Bainbridge, Apr. 23, 1967, 
TAP); Warren (S. Pine Creek, May 5, 1968, GRF) 

169. Wadotes hydridus (Emerton) , 1890 

Wayne (Richmond, 20, 21, 22); Putnam (Bainbridge, Apr. 23, 
1967, TAP) 

family Hahniidae 

170. Hahnia cinerea Emerton, 1890 

Wayne (Richmond, 20, 21, 22, 39); Steuben (Crooked Lake, 21, 
22); Porter (Ogden Dunes, 39, 22); Spencer (Santa Claus, 

May 8, 15, 22, July 17, 1966, RWM) 

171. Neoantistea agilis (Keyserling), 1887 

Wayne (Richmond, 20, 21, 22, 39); Steuben (Crooked Lake, 21, 
22); Tippecanoe (Ross Biological Reserve, 5; Americus, Aug. 
2 A, Sept. 7, 1967, TAP) 

172. Neoantistea radula (Emerton), 1890 
Hahnia riparia Keyserling, 1887 
Hahnia radula Emerton, 1890 

N. riparia race radula Gertsch, 1934 

Wayne (Richmond, 20, 21, 22, 39); LaPorte (Smith, 39, 22) 

family Pisauridae 

173. Dolomedes scriptus Hentz, 1845 

Monroe (Mayneld's Cave, 4, 22); Porter (Dunes Acres, 38, 39, 
22) ; Tippecanoe (Americus, June 23, 1966, TAP) ; Vermillion 
(Clinton, July 1U, 1966, TAP); Fountain (Attica, Aug. 3, 1966, 
TAP); Fulton (Rochester, Aug. 29, 1967, TAP); Sullivan 
(Merom Station, July 18, 1967, TAP) 

174. Dolomedes sexpunctatus Hentz, 1845 

Tippecanoe (Lafayette, 25, 22); Kosciusko (Tippecanoe Lake, 
June 6, 2, 21, 22); Putnam (Greencastle, 2, 21, 22); INDIANA 
(49); Porter (Dunes Acres, 38, 39, 22); Fulton (Rochester, 
Sept. 16, 1967, TAP); Posey (Mount Vernon, June 21, 1968, 

175. Dolomedes striatus Giebel, 18G9 

Porter (Dunes Acres, 38, 39) 

176. Dolomedes tenebrosus Hentz, 1844 

Tippecanoe (Lafayette, 25, 22; Americus, July 27, 1967, TAP); 
Crawford (Wyandotte, July 25, 2, 21, 22); Starke (Bass Lake, 
Oct. 10, 2, 21, 22) ; Lawrence (Mitchell, July 15, 2, 21, 22) ; 
Jackson (Medora, Sept. 2h, 2, 21, 22) ; Knox (May 26, 2, 21, 22; 
Cypress Swamp, Sept. 25, 2, 21, 22); Marshall (Lake Maxin- 
kuckee, Aug. 7, 2, 21, 22) ; Gibson (Sept. 1, 2, 21, 22) ; Put- 
nam (Greencastle, 2, 21, 22); LaPorte (Wilders, July 25, 2, 21, 
22; Smith, 39, 22); INDIANA (49); Wayne (Richmond, 20, 
21, 22, 39); Steuben (Crooked Lake, 21, 22); Brown (Nashville, 
21, 22); Porter (Valparaiso, 21, 22; Dunes Acres, 38, 39, 22; 
Ind. Dunes State Park, 22), Carroll (Delphi, June 22, 1966, 

Entomology 289 

DEW); Harrison (Mauckport, May 6, 1968, TAP); Monroe 
(Bloomington, July 3, 1968, TAP) 

Banks (2) collected this species from Wyandotte, not in Wyandotte 
Cave, as Elliott (22) has indicated. Banta (4) collected Dolomedes scrip- 
tus from Mayfield's Cave, not D. tenebrosus, as Elliott (22) has indi- 
cated. Dolomedes striatus, D. scriptus, and D. tenebrosus are not synony- 
mous to one another, as Elliott (22) has indicated. 

177. Dolomedes urinator Hentz, 1845 

Lawrence (Donnelson's Cave, 6, 2, 21, 22; Mitchell, Apr. 2, 2, 
21,22); INDIANA (49) 

178. Pisaurina brewipes (Emerton), 1911 

LaPorte (Smith, 39, 22) 

179. Pisaurina mir a (Walckenaer) , 1837 

Marshall (Arlington, June 10, 2, 21, 22; Culver, June 29, 2, 
21, 22); Starke (Bass Lake, June 20, 2, 21, 22); Martin (Shoals, 
July 13, 2, 21, 22); Putnam (Greencastle, 2, 21, 22); Dubois 
(Huntingburg, 2, 21, 22); Wayne (Richmond, 20, 21, 22, 39); 
Steuben (Crooked Lake, 21, 22); Brown (Nashville, 21, 22); 
Porter (Valparaiso, 21, 22; Dunes Acres, 38, 39, 22; Ogden 
Dunes, 39, 22; Ind. Dunes State Park, 22); LaPorte (Smith, 
39, 22) ; Tippecanoe (Ross Biological Reserve, 5) 

180. Pisaurina mira var. subinflata (Hentz), 1850 

Posey (New Harmony, Sept. 3, 2, 21, 22); Jennings (North 
Vernon, Apr. U, 2, 21, 22); INDIANA (49) 

family Lycosidae 

181. Arctosa emertoni Gertsch, 1934 

Tippecanoe (West Lafayette, Sept. 30, 1966, TAP) ; Fountain 
(Attica, July 28, 1967, TAP) 

182. Arctosa funerea (Hentz), 1844 

Knox (July 2, 2, 21, 22) ; INDIANA (49); Wayne (Richmond, 
20, 21, 22) ; Tippecanoe (Ross Biological Reserve, 5) 

183. Arctosa littoralis (Hentz), 1844 

INDIANA (40), Lake (Hammond, June 16, 2, 21, 22; Pine, 
May 25, 2, 53, 21, 22; Miller, 53, 39, 22); Posey (Grand 
Chain, Sept. 5, 2, 21, 22; Mount Vernon, Aug. 17, 1966, June 
7, 1967, June 8, 1968, TAP); Putnam (Greencastle, 2, 21, 22); 
Porter (Ind. Dunes State Park, 53; Dunes Acres, 38, 39, 22; 
Ogden Dunes, 39, 22) 

184. Arctosa noctuabunda (Montgomery), 1904 

Gibson (Mount Carmel, July 12, 1967, TAP) 

This specimen was determined by Dr. Willis J. Gertsch of The 
American Museum of Natural History. It has been retained at the 

185. Arctosa rubicunda (Keyserling), 1876 

Tippecanoe (Lafayette, 25, 22); INDIANA (49) 

290 Indiana Academy of Science 

186. Geolycosa fatifera (Hentz), 1842 

Crawford (Wyandotte, May 27, Sept. 8, 2, 21, 22); Posey 
(New Harmony, Sept. 3, 2, 21, 22) ; Marshall (Culver, Aug. 22, 
2, 21, 22); Starke (Bass Lake, Oct. 10, 2, 21); Marion (May 30, 
2,21,22); INDIANA (49) 

187. Geolycosa missouriensis (Banks), 1895 

Lake (Buffington, 54); Pulaski (Oct. 6, 1930, JJD, 54) ; Porter 
(Dunes Acres, 38, 39, 22) 

Wallace (54) states, "Geolycosa fatifera (Hentz) is obviously closely 
related to missouriensis (Banks). Superficially they resemble each other 
closely, but their genitalia are distinct. It may be that, when males have 
been taken from a wider area, this species will be found to intergrade 
with missouriensis. ," 

188. Geolycosa pikei (Marx), 1881 

Lake? (Miller, 53; Pine, 53; Clark, 53); Porter? (Ind. Dunes 
State Park, 53) 

Comstock (16, 17) states, 'The range of this species includes the 
Eastern and Middle States and extends to the District of Columbia and 
to Indiana." Comstock apparently included the "Middle States" and 
"Indiana" on the basis of Shelford's records. The writer feels certain 
that Shelford's records were misidentifications. Shelford's G. pikei was 
most likely G. wrightii which is indigenous to the dunes area and was 
described by Emerton in 1912, the same year Shelford published his 

Wallace (54) says that Geolycosa pikei is "the sandy-beach-inhabit- 
ing form of the New England seaboard." 

189. Geolycosa wrightii Emerton, 1912 

Lake (Buffington, 24?, 54; Miller, 54); Porter (Tremont, 54; 
Ind. Dunes State Park, 54, 22; Ogden Dunes, 54, 39, 22; Dunes 
Acres, 54, 38, 39, 22) 

Although Emerton (24) designated no types, Wallace (54) states 
that the probable female type was collected at Chicago, Illinois, by 
C. B. Davenport in April, 1904. The probable male type was collected 
at Buffington, Indiana, on Sept. 6, 1909 by W. H. Wright. 

190. Lycosa aspersa Hentz, 1844 

Tippecanoe Lafayette, 25, 22); INDIANA (49); Porter (Dunes 
Acres, 38, 39, 22; Ogden Dunes, 39, 22; Ind. Dunes State Park, 
22; LaPorte (Smith, 39; Michigan City, Sept. 10, 1967, TAP); 
Carroll (Delphi, June 23, 1966, TAP) 

Lowrie (39) collected this species at Smith, not Miller, as Elliott 
(22) has indicated. 

191. Lycosa avara (Keyserling), 1876 

Tippecanoe (Lafayette, 25, 22, Oct. 10, 1966, TAP); Jennings 
(North Vernon, May 7, 2, 21, 22); INDIANA (49); Porter 
(Dunes Acres, 38, 39, 22); Spencer (Santa Claus, May 22, 1966, 

Entomology 291 

192. Lycosa avida Walckenaer, 1837 

Tippecanoe (Lafayette, 25, 22, Oct. 10, 1966, TAP) ; Lake 
(Hammond, May 7, 2, 21, 22); Wayne (Richmond, 21, 22); 
Porter (Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22; Ind. 
Dunes State Park, 22; Lake (Miller, 39, 22); Fountain (Attica, 
July 28, Aug. 11, 1967, TAP); Monroe (Bloomington, July 3, 
1968, TAP) 

193. ' Lycosa baltimoriana (Keyserling), 1876 
Porter (Dunes Acres, 38, 39, 22) 

194. Lycosa carolinensis Walckenaer, 1837 

Tippecanoe (Lafayette, 25, 22, Sept. 22, 1966, TAP); Crawford 
(Wyandotte, July 25, 2, 21, 22?); Jennings (North Vernon, 
Sept. 1U, 2, 21, 22); LaPorte (Aug. 26, 2, 21, 22); Boone 
(Sept. 20, 1966, TRH) ; Benton (Otterbein, Apr. 22, 1968, 

Banks (2) collected this species at Wyandotte, not in Wyandotte Cave 
as Elliott (22) has indicated. 

195. Lycosa frondicola Emerton, 1885 

Tippecanoe (Lafayette, 25, 22); Lawrence (Mitchell, Aug. 28, 
2, 21, 22); Jennings (North Vernon, Apr. 1U, 2, 21, 22); Dubois 
(Huntingburg, 2, 21, 22); Floyd (New Albany, May h, 2, 21, 
22); Porter (Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22; Ind. 
Dunes State Park, 22); Lake (Miller, 39, 22); Warren (S. Pine 
Creek, May 5, 1968, GRF); Benton (Otterbein, May 2, 1968, 

196. Lycosa gulosa Walckenaer, 1837 

Kosciusko (Tippecanoe Lake, Aug. 17,2,21,22) ; Dubois (Hunt- 
ingburg, 2, 22); Starke (Bass Lake, Oct. 10, 2, 21, 22); Jackson 
(Medora, Sept. 21*, 2, 21, 22) ; Posey (Grand Chain, Sept. 5, 2, 
21, 22); Crawford (Wyandotte, SejH. 8, 9, 2, 21); INDIANA 
(49); Wayne (Richmond, 20, 21, 22, 39); LaPorte (Smith, 39, 
22); Tippecanoe (Merritt's Pine Plantation, Apr. 7, 1968, RWM; 
Lafayette, Apr. 15, 1968, TAP) 

Although the records of Banks (2) are Lycosa kochii Keyserling, 
1876, he probably had L. kochii Emerton which is synonymous to Lycosa 
gulosa Walckenaer. All other records are of L. gulosa. 

197. Lycosa helluo Walckenaer, 1837 

INDIANA (40); Tippecanoe (Lafayette, 25, 22; West Lafayette, 
Oct. 18, 1966, TAP; Americus, July 27, Aug. 15, Sept. 7, 1967, 
TAP); Marshall (Arlington, June 10, 2, 21, 22); Kosciusko 
(Tippecanoe Lake, June 6, 2, 21, 22); Starke (Bass Lake, Apr. 
9, 2, 21, 22); Lawrence (Mitchell, Aug. 28, 2, 21, 22); Posey 
(Grand Chain, May 12, June 3, 2, 21, 22); Jackson (Medora, 
Sept. 2U, 2, 21, 22); Parke (Mecca, Apr. 27, 2, 21, 22); Clark 
(Jeffersonville, June 23, 2, 21, 22); Marion (May 30, 2, 21, 22); 
Dubois (Huntingburg, 2, 21, 22); Porter (Dunes Acres, 38, 39; 
Ogden Dunes, 39; Ind. Dunes State Park, 22); Allen (Fort 

292 Indiana Academy of Science 

Wayne, Sept. 21, 1966, TAP); Fulton (Rochester, July 27, 1967, 
TAP); Fountain (Attica, July 28, 1967, July 18, 1968, TAP) 
Marx (40) first recorded this species from Indiana, not Emerton, as 
Elliott (22) has indicated. 

198. Lycosa permunda Chamberlin, 1904 

LaPorte (Smith, 39, 22) 

199. Lycosa punctulata Hentz, 1844 

Tippecanoe (Lafayette, 25, Sept. 26, 1966, JDH); Jennings 
(North Vernon, Sept. IU, 2, 21, 22); Putnam (Greencastle, 2, 

200. Lycosa rahida Walckenaer, 1837 

Lawrence (Mitchell, Aug. 28, 2, 21, 22); Putnam (Greencastle, 
2, 21, 22); Dubois ( Huntingburg, 2, 21, 22); Wayne (Richmond, 
20, 21, 22, 39); Porter (Dunes Acres, 38, 39, 22; Ogden Dunes, 
39, 22; Ind. Dunes State Park, 22); LaPorte (Smith, 39, 22); 
Spencer (Santa Claus, July 1U, 1966, RWM); Tippecanoe 
(Americus, July 27, 1967, TAP) 

201. Pardosa lapidicina Emerton, 1885 

Kosciusko (2, 21); Tippecanoe (Ross Biological Reserve, 5) 
This species was not recorded by Banks (2) from Bass Lake, Jeffer- 
sonville, and Huntingburg as Elliott (22) has indicated. 

202. Pardosa milvina (Hentz), 1844 

Tippecanoe (Lafayette, 25, 22); Starke (Bass Lake, Apr. 10, 
2, 22); Clark (Jeffersonville, June 23, 2, 21); Dubois (Hunting- 
burg, 2, 22); Wayne (Richmond, 21, 22); Porter (Dunes Acres, 
38, 39, 22; Ogden Dunes, 39, 22; Ind. Dunes State Park, 22); 
Lake (Miller, 39, 22); LaPorte (Smith, 39, 22); Carroll (Delphi, 
June 23, 1966, TAP); Huntington (Huntington, June 28, 1966, 
TAP); Vermillion (Clinton, July 13, 1966, July 20, 1967, TAP); 
Sullivan (Merom Station, July 20, 21, 1966, July 18, 1967, 
TAP); Gibson (Mount Carmel, July 27, 1966, July 12, 1967, 
TAP); Fountain (Attica, Aug. 3, 1966, July 28, Aug. 11, 25, 
Sept. 7, 1967, TAP) ; Fulton (Rochester, July 27, Sept. 16, 
1967, TAP) ; Brown (Brown Co. State Park, Sept. 13, 1966, 

Pardosa flavipes is a synonym of Pardosa milvina (Hentz), not of 
Pardosa flavipalpis F. Cambridge, as Elliott (21, 22) has indicated. 

203. Pardosa modica (Blackwall), 1846 
P. fuscula (Thorell), 1875 

Porter (Dunes Acres, 38, 39, 22) 

204. Pardosa moesta Banks, 1892 

Porter (Dunes Acres, 38, 39, 22) 

205. Pardosa xerampelina (Keyserling), 1876 

Tippecanoe (Ross Biological Reserve, 5) 

206. Pirata insidaris Emerton, 1885 

Lake (Hammond, May 7, 2, 21, 22); INDIANA (49); Porter 

Entomology 293 

(Valparaiso, 21, 22; Ogden Dunes, 39, 22); LaPorte (Smith, 
39, 22) 

207. Pirata macidatus Emerton, 1909 

Carroll (Delphi, July 7, 1966, TAP); Fountain (Attica, July 25, 
1967, TAP) ; Tippecanoe (Americus, July 27, Aug. 10, 1967, 
TAP); Fulton (Rochester, July 27, 1967, TAP) 

208. Pirata marxi Stone, 1890 

Wayne (Richmond, 21, 22); LaPorte (Smith, 39, 22); Porter 
(Ind. Dunes State Park, 22); Tippecanoe (Ross Biological Re- 
serve, 5) 

209. Pirata minutus Emerton, 1885 

Tippecanoe (Lafayette, 25, 22; Americus, July 27, 1967, TAP); 
Wayne (Richmond, 21, 22) 

210. Pirata montana Emerton, 1885 

Tippecanoe (Lafayette, 25) 

211. Pirata piratica (Olivier) , 1789 

Steuben (Clear Lake, Aug. 13, 2, 21, 22); Knox {May 26, 2, 
21, 22); INDIANA (49); Wayne (Richmond, 20, 21, 22, 39); 
Porter (Dunes Acres, 38, 39; Ogden Dunes, 39); LaPorte 
(Smith, 39); Tippecanoe (Americus, June 23, 1966, July 27, 
1967, TAP) 

212. Schizocosa hilineata (Emerton), 1885 

Wayne (Richmond, 21, 22); Porter (Valparaiso, 21, 22); Tippe- 
canoe (Merritt's Pine Plantation, June 10, 1968, RWM) 

213. Schizocosa crassijyalpis (Emerton) , 1909 

Warren (S. Pine Creek, May 5, 1968, GRF) 

214. Schizocosa crassipes (Walckenaer), 1837 

Tippecanoe (Lafayette, 25, 22; Americus, July 27, Aug. 24, 
1967, TAP); Kosciusko (Tippecanoe Lake, June 8, Aug. 17, 2, 
21, 22; Vawter Park, June 1, 2, 21, 22); Lawrence (Michell, 
July 15, 2, 21, 22) ; LaPorte (Aug. 26, 2, 21, 22; Smith, 39, 22); 
Wayne (Richmond, 20, 39); Porter (Valparaiso, 21, 22; Dunes 
Acres, 38, 39, 22); Fulton (Rochester, July 27, Aug. 2, Sept. 16, 
1967, TAP) ; Huntington (Huntington, Sept. 16, 1967, TAP) 

215. Schizocosa saltatrix (Hentz),1844 

Tippecanoe (Lafayette, 25, 22; Ross Biological Reserve, 5); 
Crawford (Wyandotte, Apr. 17, 2, 21, 22?); Porter (Dunes 
Acres, 38, 39, 22; Ogden Dunes, 39; Ind. Dunes State Park, 22); 
Carroll (Delphi, June 23, 1966, TAP) ; Posey (Mount Vernon, 
June 7, 1967, TAP); Benton (Otterbein, May 2, 1968, JOS); 
Spencer (Santa Claus, May 8, 1966, RWM) 
Banks (2) recorded this species from Wyandotte, not Wyandotte 

Cave as Elliott (22) has indicated. Lowrie (39) recorded this species from 

Ogden Dunes, not Smith as Elliott (22) has indicated. 

216. Trochosa pratensis (Emerton), 1885 

Knox (Vincennes, Apr. 25, 2, 21, 22); Crawford (Wyandotte, 

294 Indiana Academy of Science 

Aug. 17, 2, 21, 22?); Starke (Bass Lake, Apr. 10, 2, 21, 22); 
Jackson (Medora, Sept. 24, 2, 21, 22); Lake (Pine, Oct. 29, 2, 

21,22); INDIANA (49) 
Banks (2) recorded this species from Wyandotte, not Wyandotte 
Cave as Elliott (22) has indicated. 

family Oxyopidae 

217. Oxyopes salticus Hentz, 1845 

Wayne (Richmond, 21, 22); Porter (Dunes Acres, 38, 39, 22); 
Tippecanoe (Ross Biological Reserve, 5; Ent. Research Area, 
July 24, 1968, MKP); Carroll (July 7, 1966, TAP); Fountain 
(Attica, Aug. 25, 1967, TAP) 

family Gnaphosidae 

218. Callilepis imbecilla (Keyserling), 1887 

Porter (Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22); Tippe- 
canoe (Ross Biological Reserve, 5); Spencer (Santa Claus, June 
5, 1966, RWM) 
The writer has a Callilepis in his collection, but it is not imbecilla. 

219. Cesonia bilineata (Hentz), 1847 


220. Drassodes neglectus (Keyserling), 1887 

Porter (Dunes Acres, 38, 39, 22; Ind. Dunes State Park, 22) 

221. Drassyllus creolus Chamberlin & Gertsch, 1940 

Spencer (Santa Claus, July 3, 11, 1966, RWM) 

222. Drassyllus depressus (Emerton), 1890 

Tippecanoe (Lafayette, 25, 22; Merritt's Pine Plantation, July 
11, 1967, June 15, 1968, RWM) 

223. Drassyllus rufulus (Banks), 1892 

LaPorte (Smith, 39, 22) 

224. Drassyllus virginianus Chamberlin, 1922 

Tippecanoe (4-H Leadership Camp, July 12, 1968, TAP) 

225. Gnaphosa sericata L. Koch, 1866 

G. bicolov (Hentz), 1847 (bicolor preoc. by Hahn, 1831) 

INDIANA (9); Porter (Ogden Dunes, 39, 22; Ind. Dunes State 
Park, 22); Tippecanoe (Merritt's Pine Plantation, July 11, 
1967, RWM; Warren (S. Pine Creek, May 5, 1968, GRF) 

226. Haplodrassus bicornis (Emerton), 1909 

LaPorte (Smith, 39, 22) 

227. Haplodrassus signifer (C. L. Koch), 1839 
Drassus robustus Emerton, 1890 

Tippecanoe (Lafayette, 25, 22); Porter (Dunes Acres, 38, 39, 
22); Lake (Miller, 39, 22; Smith, 39, 22) 

228. Herpyllus ecclesiasticiis Hentz, 1832 

H. vasifer (Walckenaer) , 1837 

Tippecanoe (Lafayette, 21, 22, June 20, 1968, TAP); Putnam 

Entomology 295 

(Greencastle, 2, 21, 22; Bainbridge, Apr. 23, 1967, TAP); 
Wayne (Richmond, 20, 21, 22, 39); Porter (Valparaiso, 21, 22; 
Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22); Lake (Munster, 
May 2, 1968, TAP); Vanderburgh (Evansville, May 10, 1968, 
TAP); Fulton (Rochester, Aug. 29, 1967, TAP); Gibson 
(Mount Carmel, July 27, 1966, TAP) 

The writer can see no reason why many have chosen to use 
Walckenaer's Drassus vasifer, rather than Hentz's Herpyllus ecclesiasti- 
cus. Hentz (28) gives an accurate description of the species, gives it a 
valid name, and tells of its habits. He states, on page 102, ". . . ; and 
a blackish one with a white band on the cephalothorax, a band on the 
abdomen, beginning at base and reaching the middle, and a spot near the 
apex white. This one attains a great size and is found in houses, 
under stones, planks, the bark of decaying trees, etc. I call it 
H. ecclesiasticus, . . ." 

Elliott (22) lists Nodocion sp. for number 208. The specimen was 
taken at Richmond, Wayne Co., Indiana. Levi (34) described Nodocion 
melanie from Wisconsin. Elliott's specimen may be this species, but the 
writer has not had the opportunity to examine the specimen. Three other 
species of Nodocion are found in the West. 

229. Sergiolus capulatus (Walckenaer), 1837 
5. variegatus (Hentz), 1847 

Kosciusko (Tippecanoe Lake, June 6, 2, 21, 22); INDIANA 
49, 9); Wayne (Richmond, 20, 21, 22, 39); Porter (Valparaiso, 
21, 22); Tippecanoe (Ross Biological Reserve, 5) 

230. Sergiolus montanus (Emerton) , 1890 
(nee montanus Emerton, 1909) 

Sullivan (Merom Station, July 21, 1966, TAP) 
Wilton Ivie of The American Museum of Natural History determined 
this specimen. 

231. Sosticus insularis (Banks), 1895 

Porter (Dunes Acres, 38, 39, 22; Ind. Dunes State Park, 22); 
Tippecanoe (Americus, July 27, 1967, TAP) 

232. Zelotes duplex Chamberlin, 1922 

Tippecanoe (Ross Biological Reserve, 5); Posey (Mount Ver- 
non, Aug. 18, 1966, TAP); Spencer (Santa Claus, May 22, July 
2h, 1966, RWM) 

233. Zelotes hentzi Barrows, 1945 

Fountain (Attica, Aug. 25, 1967, TAP); Warren (S. Pine Creek, 
Apr- 13, 1968, GRF) 
The writer's specimen was determined by Wilton Ivie of The Ameri- 
can Museum of Natural History. 

This species, which occurs in the West, may be synonymous to 
Zelotes melancholica (Thorell), 1877, according to Kaston (32). 

234. Zelotes laccus (Barrows), 1919 

Tippecanoe (Ross Biological Reserve, 5; Merritt's Pine Planta- 
tion, July 20, 1968, RWM) 

296 Indiana Academy of Science 

235. Zelotes subterraneus (C. L. Koch), 1839 
Z. ater (Hentz), 1832 

Crawford (Wyandotte, Apr. 17, 2, 21, 22) ; Wayne (Richmond, 
20, 21, 22, 39); Steuben (Crooked Lake, 21, 22); Porter (Val- 
paraiso, 21, 22) ; Fulton (Rochester, Sejrt. 16, 1967, TAP) 
The writer has arbitrarily assigned the record of Banks (2) and 
Elliott (20, 21, 22) to Zelotes subterraneus. Hentz (28) probably had two 
species when he described Herpyllus ater. The writer refers the reader to 
Kaston (32), pages 356 and 357, for a discussion of this taxonomic 

family Clubionidae 

236. Agroeca minuta Banks, 1895 

Wayne (Richmond, 21, 22); Steuben (Crooked Lake, 21, 22); 
Porter (Ogden Dunes, 39, 22) 

237. Agroeca pratensis Emerton, 1890 

Marshall (Arlington, 2, 21, 22) ; INDIANA (49) 

238. Castianeira cingulata (C. L. Koch), 1842 

Wayne (Richmond, 20, 21, 22, 39); Steuben (Crooked Lake, 21, 
22); Porter (Valparaiso, 21, 22; Dunes Acres, 38, 39, 22; Ogden 
Dunes, 39, 22; Ind. Dunes State Park, 22); LaPorte (Smith, 39, 
22); Tippecanoe (Ross Biological Reserve, 5; Americus, Aug. 
10, 1967, TAP) 

239. Castianeira descripta (Hentz), 1847 

Tippecanoe (Lafayette, 25, 22); Putnam (Greencastle, 2, 21, 
22); Fountain (Attica, Aug. 25, Sept. 7, 1967, TAP); Hunt- 
ington (Huntington, July 27, 1967, TAP) 

240. Castianeira longipalpus (Hentz), 1847 

Kosciusko (Tippecanoe Lake, Aug. 17, 2, 21, 22); Knox (July 2, 
2, 21, 22); Wayne (Richmond, 20, 21, 22, 39); LaPorte (Smith, 
39, 22) 

241. Castianeira trilineata (Hentz), 1847 

INDIANA (40, 49); Porter (Dunes Acres, 38, 39, 22) 

Marx (40) recorded this species for Indiana, not Hentz, as Elliott 

(22) has indicated. Elliott (22) lists Fox as having recorded this species 

from Lafayette. Actually Fox (25) obtained his Lafayette record from 

Marx's catalogue, as he states in a footnote at the bottom of page 268. 

242. Chiracanthium inclusum (Hentz), 1847 

Knox (Vincennes, Aug. 23, 2, 21, 22) 

243. Clubiona abbotii L. Koch, 1866 

Tippecanoe (Lafayette, 25, 22, Apr. 22, 196S, TAP; Americus, 
June 23, 1966, TAP); Marshall (Arlington, June 10, 2, 21, 
22); INDIANA (49); Huntington (Huntington, June 29, 1966, 
TAP); Sullivan (Merom Station, July 20, 1966, July 18, 1967, 
TAP); Fountain (Attica, Aug. 3, 1966, Sept. 7, 1967, July U, 
1968, TAP) 

244. Clubiona elizabethae Kaston, 1945 

Tippecanoe (Ross Biological Reserve, 5) 

Entomology 297 

245. Clubiona obesa Hentz, 1847 

Wayne (Richmond, 20, 21, 22); Porter (Valparaiso, 21, 22; 
Ogden Dunes, 39, 22); LaPorte (Smith, 39, 22); Huntington 
(Huntington, June 28, 1966, TAP); Posey (Mount Vernon, June 
7,1967, TAP) 

246. Clubiona pallens Hentz, 1847 

Wayne (Richmond, 20, 21, 22, 39); Porter (Valparaiso, 21, 22; 
Dunes Acres, 38, 39, 22); LaPorte (Smith, 39, 22); Tippecanoe 
(Ross Biological Reserve, 5) 

247. Clubiona plumbi Gertsch, 1941 

Gibson (Mt. Carmel, July 27, 1966, TAP); Tippecanoe (Ameri- 
cus, Aug. 1, 1967, TAP) 

248. Clubiona pygmaea Banks, 1892 

Tippecanoe (Americus, June 23, 1966, TAP) 

249. Clubiona riparia L. Koch, 1866 

Cass (Logansport, Aug. 3, 1967, TAP) 

250. Clubiona tibialis Emerton, 1890 

Cass (Logansport, July 6, 1966, TAP); Gibson (Mt. Carmel, 
July 27, 1966; TAP); Fountain (Attica, Aug. 3, 1966, TAP); 
Posey (Mt. Vernon, Aug. 18, 1966, June 7, 1967, TAP); Carroll 
(Delphi, Aug. 3, 1967, TAP); Fulton (Rochester, July 27, 1967, 

251. Clubiona tribola Banks, 1907 

Crawford (Wyandotte, 2, 3 49, 21, 22) 

This species was described as new by Banks (2) on the basis of one 
female. It was taken at Wyandotte, not in Wyandotte Cave, as Petrunke- 
vitch (49) has indicated. 

252. Meriola decepta Banks, 1895 

Tippecanoe (Ross Biological Reserve, 5) 

253. Micaria aurata (Hentz), 1847 

Porter (Dunes Acres, 39, 22; Ind. Dunes State Park, 22) 

254. Micaria montana Emerton, 1890 

Porter (Dunes Acres, 38) 

255. Phrurolithus similis Banks, 1895 

Wayne (Richmond, 20, 21, 22) 

256. Phrurotimpus alarms (Hentz), 1847 
Phrurolithus palustris Banks, 1892 

Posey (Grand Chain, May 12, 2, 21, 22); Wayne (Richmond, 
20, 21, 22, 39); Steuben (Crooked Lake, 21, 22); Porter (Ogden 
Dunes, 39, 22); Tippecanoe (Ross Biological Reserve, 5); 
Spencer (Santa Claus, May 8, 15, 22, 29, June 5, 12, 26, July 11, 
1U, Aug. 15, 1966, RWM) 

257. Phrurotimpus borealis (Emerton), 1911 

Spencer (Santa Claus, June 5, 1966, RWM); Gibson (Mt. Car- 
mel, Sept. 23, 1967, TAP) 

298 Indiana Academy of Science 

258. Phrurotimpus minutus (Banks), 1892 

Spencer (Santa Claus, Sept. 4, 1966, RWM) 

259. Tracheitis tranquillus (Hentz), 1847 

Tippecanoe (Lafayette, 25, 22, Sept. 26, 1966, TAP; West 
Lafayette, Sept. SO, 1966, JWB); Kosciusko (Tippecanoe Lake, 
June 8, 2, 21, 22); Putnam (Greencastle, 2, 21, 22); Dubois 
(Huntingburg, 2, 21, 22); Wayne (Richmond, 20, 21, 22, 39); 
Porter (Valparaiso, 21, 22; Dunes Acres, 38, 39, 22; Ind. Dunes 
State Park, 22); Bartholomew (Columbus, Oct. 25, 1967, TAP) 

260. Zora pumila (Hentz), 1850 

Tippecanoe (Ross Biological Reserve, 5) 

family Anyphaenidae 

261. Anyphaena celer (Hentz), 1847 

LaPorte (Smith, 39, 22); Porter (Ogden Dunes, 39, 22; Ind. 
Dunes State Park, 22); Tippecanoe (Ross Biological Reserve, 5) 

262. Anyphaena fraterna (Banks), 1893 

Tippecanoe (Ross Biological Reserve, 5); Warren (South Pine 
Creek, May 5, 1968, GRF) 

263. Anyphaena pectorosa L. Koch, 1866 

Wayne (Richmond, 20, 21, 22, 39); LaPorte (Smith, 39, 22) 

264. Anyphaenella saltabunda (Hentz), 1847 

Wayne (Richmond, 20, 21, 22); Porter (Valparaiso, 21, 22); 
Brown (Nashville, 21, 22); Tippecanoe (Ross Biological Re- 
serve, 5; Ent. Research Area, July 2U, 1968, MKP) 

265. Aysha gracilis (Hentz), 1847 
Anyphaena rubra Emerton, 1890 

Tippecanoe (Lafayette, 25, 22; Ross Biological Reserve, 5); 
Lake (Hammond, June 16, 2, 21, 22) ; Fountain (Veedersburg, 
May 12, 2, 21, 22); INDIANA (49); Porter (Dunes Acres, 38, 
39, 22; Ogden Dunes, 39, 22); LaPorte (Smith, 39, 22) 
This species is not synonymous with Microthena gracilis Walck. 

(sic), Acrosona gracilis Walck. (sic), and Epeira rugosa as Elliott (22) 

has indicated. 

266. Sillus conspersa (Keyserling), 1887 
Anyphaena conspersa Keyserling, 1887 

Lake (Hessville, 53; Liverpool, 53) 
F.O.P. — Cambridge erected the genus Sillus in 1900, not in 1896, as 
Petrunkevitch (49) has indicated. 

family Thomisidae 

267. Coriarachne lenta (Walckenaer), 1837 
Xysticus versicolor (Keyserling), 1880 

Crawford (Wyandotte, Apr. 17, 2, 21, 22) ; Posey (Grand Chain, 
Apr. 19, 2, 21) ; Parke (Mecca, Apr. 27, 2, 21, 22) ; Floyd (New 
Albany, May 4, 2, 21, 22); Putnam (Greencastle, 2, 21, 22); 
Porter (Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22); LaPorte 
(Smith, 39, 22) 

Entomology 299 

Banks (2) did not record this species from Vincennes as Elliott (22) 
has indicated. 

268. Ebo latithorax Keyserling, 1883 

Porter (Dunes Acres, 38, 39, 22; Ind. Dunes State Park, 22); 
Tippecanoe (Ross Biological Reserve, 5) 

269. Ebo pepinensis Gertsch, 1933 

Porter (Ogden Dunes, 39, 22) 

270. Misumena calycina (Linnaeus), 1758 

Tippecanoe (Ross Biological Reserve, 5); Huntington (Hunt- 
ington, June 29, 1966, TAP) 

721. Misumenoides formosipes (Walckenaer), 1837 
M. aleatorius (Hentz), 1847 

Steuben (Lake James, May 8, 2, 21, 22); Lake (Hammond, 
July 30, 2, 21, 22); Posey (Grand Chain, Sejjt. 5, 2, 21); Put- 
nam (Greencastle, 2, 21, 22); Porter (Dunes Acres, 38, 39, 22; 
Ogden Dunes, 39, 22; Ind. Dunes State Park, 22); LaPorte 
(Smith, 39, 22); Tippecanoe (Ross Biological Reserve, 5); Sulli- 
van (Merom Station, July 20, 1966, TAP); Cass (Logansport, 
July 6, 1966, TAP); Pulaski (Winamac, Aug. 22, 1967, TAP) 

Banks (2) did not record this species from Vincennes as Elliott (22) 
has indicated. 

272. Misumenops americanus (Keyserling), 1880 

Starke (Bass Lake, June 17, 2, 21, 22); Posey (Grand Chain, 
June 3, 2, 21, 22) ; Fountain (Attica, June 20, 2, 21, 22) ; Mar- 
shall (Culver, June 29, 2, 21, 22) ; Knox (July 2, 2, 21, 22) ; 
Putnam (Greencastle, 2, 21, 22); Porter (Valparaiso, 21, 22) 

273. Misumenops asperatus (Hentz), 1847 

Crawford (Wyandotte, June 23, Sept. 20, Apr. 17, 2, 21, 22); 
Posey (New Harmony, May 6, 2, 21, 22); Lake (Pine, May 25, 
2, 53, 21, 22; Miller, 53; Clark, 53; Hessville, 53; Liverpool, 53; 
Dubois (Huntingburg, 2, 21, 22); LaPorte (Otis, 53; Smith, 39, 
22); Porter (Woodville, 53; Ind. Dunes State Park, 53, 22; 
Valparaiso, 21, 22; Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 
22); Wayne (Richmond, 20, 21, 22, 39); Tippecanoe (Ross Bio- 
logical Reseive, 5) 

274. Misumenops oblongus (Keyserling), 1880 

Fountain (Veedersburg, May 12, 2, 21, 22; Attica, Aug. 3, 1966, 
TAP), Kosciusko (Vawter Park, June 1, 2, 21, 22); Putnam 
(Greencastle, 2, 21, 22); INDIANA (49); Wayne (Richmond, 
20, 21, 22, 39); Steuben (Crooked Lake, 21, 22); Porter (Valpa- 
raiso, 21, 22; Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22); 
LaPorte (Smith, 39, 22); Posey (Mount Vernon, June 8, 1967, 
TAP); Gibson (Mount Carmel, July 27, 1966, TAP) 

275. Oxyptila americana Banks, 1895 

LaPorte (Smith, 39,22) 

300 Indiana Academy of Science 

276. Oxyptila barrow si Gertsch, 1939 

Lake (Pine, Oct. 29, NB, 27); Fulton (Rochester, July 27, 1967, 
TAP); Tippecanoe (Americus, July 27, 1967, TAP) 
The male holotype of this species was taken at Pine, Indiana, by Na- 
than Banks in 1907. He recorded it as 0. conspurcata Thorell, 1877. 
Gertsch described O. barrowsi in 1939, using Banks' Pine, Ind., record 
as the holotype. Dr. Gertsch determined both of the writer's male speci- 

277. Oxyptila conspurcata Thorell, 1877 

Knox (May 26, 2, 21, 22; Cypress Swamp, Sept. 25, 2, 21, 22); 
INDIANA (49); Tippecanoe (Ross Biological Reserve, 5) 
Gertsch (27) found that Banks' Pine, Ind., collection was Oxyptila 
barrowsi n.sp. 

278. Oxyptila monroensis Keyserling, 1883 

Wayne (Richmond, 20, 21, 22) 

279. Philodromus abbotii Walckenaer, 1837 
P. marxii Keyserling, 1889 

P. ornatus Banks, 1892 

Posey (Arlington June 10, 2, 21, 22) ;LaPorte (Wilders, July 25, 2, 

21, 22; Smith, 39, 22); INDIANA (49) 

280. Philodromus alasciensis Keyserling, 1883 

Lake (Miller, 53; Pine, 53); Porter (Ind. Dunes State Park, 53, 
22; Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22) 
The preferred spelling of this species is as above, not P. alaskensis, 
as Shelf ord (53) has indicated. 

281. Philodromus cespiticolis Walckenaer, 1802 

P. aureolus (of most authors) 

Lake (Hammond, June 16, 2, 21, 22, 19) ; Knox July 2, 2, 21, 22, 
19; Vincennes, Aug. 23, 2, 21, 22); INDIANA (49); Porter 
(Dunes Acres, 38, 39, 22, 19; Ogden Dunes, 39, 22, 19); Tippe- 
canoe (Americus, June 23, 1966, TAP); Carroll (Delphi, July 

7, 1966, TAP) ; Posey (Mount Vernon, 19, Aug. 18, 1966, June 

8, 1967, TAP) ;Gibson (Mount Carmel, July 27, 1966, TAP) ; 
Sullivan (Merom Station, July 20, 21, 1966, TAP); Vermillion 
(Clinton, July 1J+, 1966, TAP); Pulaski (Winamac, June 16, 
1966, Aug. 22, 1967, TAP); Fountain (Attica, July 25, 1967, 
TAP); Fulton (Rochester, July 27, 1966, TAP) 

282. Philodromus imbecillus Keyserling, 1880 

Porter (Dunes Acres, 38) 

283. Philodromus keyserlingi Marx, 1890 
P. washita Banks, 1932 

Posey (Arlington, June 10, NB, 19); Huntington (Huntington, 

June 28, 1966, TAP), Vermillion (Clinton, July 20, 1967, TAP) 

Dondale (19) considers P. washita a synonym of P. keyserlingi Marx, 

1890, whereas Kaston (32) considers them distinct species. The writer 

considers them synonymous. Banks (2) originally recorded this species 

as Philodromus placidus Banks, 1892. 

Entomology 301 

284. Philodromus pernix Blackwall, 1846 

Kosciusko (Tippecanoe Lake June 17, 2, 21, 22) ; Posey (Grand 
Chain, June 3, July 8, 2, 21, 22); Lake (Pine, Oct. 29, 2, 21, 22; 
Miller, 53; Clark, 53); Knox (July 2, 2, 21, 22); Putnam (Green- 
castle, 2, 21, 22); Porter (Ind. Dunes State Park, 53, 22; Dunes 
Acres, 38, 39, 22; Ogden Dunes, 39, 22); Wayne (Richmond, 
21, 22); Pulaski (Winamac, June 15, 16, 1966, TAP); Tippe- 
canoe (West Lafayette, Apr. 17, 1968, TAP); Benton (Otter- 
bein, Apr. 29, 1968, JOS); LaPorte (LaPorte, June 19, 1968, 

Banks (2) did not record this species from Vincennes, as Elliott 
(22) has indicated. 

285. Philodromus placidus Banks, 1892 

Knox (Vincennes July 10, 2, 21, 22) ; Posey (Grand Chain, 
May 12, July 8, 2, 21, 22); Lake (Pine, June 29, 2, 21, 22); 
Marshall (Culver, June 29, 2, 21, 22); Pulaski (Winamac, June 
16, 1966, TAP) 

286. Philodromus praelustris Keyserling, 1880 
Starke (Bass Lake, Apr. 9, NB, 19) 

Banks (2) originally recorded this collection as Philodromus vulgaris 
Hentz, which is a synonym of Philodromus pernix. Dondale (19) found 
that Banks' Bass Lake record was actually Philodromus praelustris. 

287. Philodromus rufus Walckenaer, 1825 

Kosciusko (Tippecanoe Lake, June 6, 2, 21, 22); Posey (Arling- 
ton, June 10, 2, 21, 22); Porter (Dunes Acres, 38, 39, 22); 
Tippecanoe (Ross Biological Reserve, 5); Fulton (Rochester, 
June 7, 1966, TAP) ; Pulaski (Winamac, June 16, 1966, TAP) ; 
Parke (June 26, 1967, TAP) 

288. Philodromus satullus Keyserling, 1880 

Wayne (Richmond, 20, 21, 22); Tippecanoe (Ross Biological 
Reserve, 5) 

289. Philodromus thorelli Marx, 1889 

Porter (Dunes Acres, 38, 39, 22) 

290. Synema varians (Walckenaer), 1837 
S. parvulum (Hentz), 1847 

Posey (Arlington, June 10, 2, 21, 22; Grand Chain, May 12, 2, 
21, 22); Fountain (Veedersburg, May 12, 2, 21, 22); Marshall 
(Culver, June 29, 2, 21, 22); INDIANA (49) 

291. Thanatus formicinus (Olivier) , 1789 

Warren (Pine Village, June 1, 1968, TH) 

292. Tibellus duttoni (Hentz), 1847 

Lake (Pine, Oct. 29, 2, 21, 22) ; Kosciusko (2, 21, 22) 
Tibellus duttoni (Hentz) is not synonymous with Tibellus oblongus 
(Walckenaer), as Elliott (21, 22) has indicated. 

293. Tibellus maritimus (Menge), 1874 

Porter (Dunes Acres, 38, 39, 22; Ind. Dunes State Park, 22) 

302 Indiana Academy of Science 

294. Tibellus oblongus (Walckenaer), 1802 

LaPorte (Wilders, July 25; 2, 21; Smith, 39, 22); Putnam 
(Greencastle, 2, 21, 22); Wayne (Richmond, 21, 22); Porter 
(Valparaiso, 21, 22; Dunes Acres, 38, 39, 22); Brown (Nashville, 
21, 22); Tippecanoe {Sept. 28, 1966, JDH); Cass (Logansport, 
July 6, 1966, TAP) ; Fulton (Rochester, July 27, 1967, TAP) 

295. Tmarus angulatus (Walckenaer), 1837 

Wayne (Richmond, 21, 22); Steuben (Crooked Lake, 21, 22); 
Porter (Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22; Ind. 
Dunes State Park, 22); LaPorte (Smith, 39, 22) 

296. Tmarus rubromaculatus Keyserling, 1880 

Tippecanoe (Ross Biological Reserve, 5) 

297. Xysticus autificus Keyserling, 1880 

Pulaski (Winamac, June 20, 1966, TAP); Tippecanoe (Ameri- 
cus, July 4, 1968, TAP) 
Drs. Redner and Turnbull of the Research Institute at Belleville, 
Ontario, Canada determined the specimen taken at Winamac. 

298. Xysticus banksi Bryant, 1933 

LaPorte (Smith, 39, 22) 

299. Xysticus bicuspis Keyserling, 1887 

Tippecanoe (Merritt's Pine Plantation, Apr. 29, May 1, 8, 1968, 

300. Xysticus discursans Keyserling, 1880. 

Sullivan (Sullivan, July 11, 1966, RWM) 

301. Xysticus elegans Keyserling, 1880 

Lake (Miller, 39); LaPorte (Smith, 39, 22) 

302. Xysticus fraternus Banks, 1895 

Wayne (Richmond, 20, 21, 22, 39); Steuben (Crooked Lake, 21, 
(Greencastle, 2, 22); Porter (Dunes Acres, 38, 39, 22; Ogden 
Dunes, 39, 22); LaPorte (Smith, 39, 22) 

303. Xysticus gulosus Keyserling, 1880 

LaPorte (Wilders, July 25, 2, 21 22; Smith, 39, 22); Putnam 
(Greencastle, 2, 22); Porter (Dunes Acres, 38, 39, 22; Ogden 
Dunes, 39, 22; Ind. Dunes State Park, 22); Tippecanoe (West 
Lafayette, Oct. 19, 1966, TAP) 

304. Xysticus luctans (C. L. Koch), 1845 

Kosciusko (2, 21,22) 

305. Xysticus punctatus Keyserling, 1880 
X. formosus Banks, 1892 

Lake (Pine, 53), Porter (Ind. Dunes State Park, 53) 

306. Xysticus transversatus (Walckenaer), 1837 
X. ferox (Hentz), 1847 

INDIANA (40); Knox (Vincennes, Aug. 23, 2, 21, 22); Posey 
(Arlington, June 10, 2, 21, 22, June 7, 1967, TAP); Crawford 
(Wyandotte, July 25, 2, 21, 22); Starke (Bass Lake, June 22, 

Entomology 303 

2, 21, 22); Marshall (Culver, June 29, 2, 21, 22); Putnam 
(Greencastle, 2, 21, 22); Wayne (Richmond, 20, 21, 22, 39); 
Porter (Valparaiso, 21, 22); LaPorte (Smith, 39, 22); Spencer 
(Santa Claus, May 8, 22, 29, June 12, 1966, RWM); Vander- 
burgh (Evansville, June 25, 1968, TAP) 

307. Xysticus transversus Banks, 1892 

Posey (New Harmony, Feb. 23, 2, 21, 22) ; INDIANA (49) 
The writer does not consider this species a valid one. The description 
was based on a single, immature specimen. He must report it, however, 
since it was taken in Indiana. 

308. Xysticus triguttatus Keyserling, 1880 

Lake (Hammond, July 30, 2, 21, 22); LaPorte (Wilders, July 
25, 2, 22), INDIANA (49); Wayne (Richmond, 21, 22); Porter 
(Valparaiso, 21, 22; Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 
22; Ind. Dunes State Park, 22) 

309. Xysticus tumef actus (Walckenaer), 1837 
X. funestus Keyserling, 1880 

X. nervosus Banks, 1892 

Kosciusko (Tippecanoe Lake, June 6, 2, 21, 22); Marshall (Cul- 
ver, June 29, 2, 21, 22); Putnam (Greencastle, 2, 21, 22); 
INDIANA (49); Wayne (Richmond, 20, 21, 22); Porter (Val- 
paraiso, 21, 22); Harrison (Corydon, Oct. 29, 1967, TAP); 
Tippecanoe (West Lafayette, Oct. 4, 1967, TAP; Merritt's Pine 
Plantation, Oct. 30, 1967, RWM) 

family Salticidae 

310. Admestina tibialis (C. L. Koch), 1848 

Porter (Ogden Dunes, 39, 22; Ind, Dunes State Park, 22); 
LaPorte (Smith, 39, 22) 

311. Agassa cerulea (Walckenaer), 1837 
A. cyanea (Hentz), 1845 

Lake (Hammond, June 16, 2, 21, 22); INDIANA (49) 

312. Attus varus Hentz, 1846 

Putnam (Greencastle, 2, 21, 22) 
Petrunkevitch (49), page 730, says that the identification of this 
spider is impossible because of the meager description given by Hentz. 

313. Attus rufus Hentz, 1846 

Tippecanoe (Lafayette, 25, 22); Knox (Vincennes, July 10, 2, 
21, 22); Crawford (Wyandotte, Sept. 9, 2, 21, 22); Starke 
(Bass Lake, June 17, 2, 21, 22) ; Posey (Grand Chain, July 8, 
2, 21, 22); LaPorte (Wilders, July 25, 2, 21, 22); Dubois 
(Huntingburg, 2, 21, 22) 

The status of Attus rufus Hentz, 1846 remains in question. Banks 
(3) synonymized this species with Phidippus whitmani Emerton, 1909. 
Kaston (32) following Petrunkevitch (49) states that Attus rufus Hentz 
is unknown to modern araneologists since the descriptions are too meager 
to place this spider in a proper genus. 

304 Indiana Academy of Science 

Elliott (21, 22) synonymized A. rufus with P. whitmanii Peckham, 
1909. The writer is convinced that Attus rufus Hentz must remain as is, 
and that Phidippus rufus Peckham, 1888 and Phidippus whitmanii Em- 
erton, 1909 are synonyms of Phidippus whitmanii Peckham, 1909. 

314. Evarcha hoyi (Peckham), 1883 

Porter (Dunes Acres 38, 39, 22); Tippecanoe (Ross Biological 
Reserve, 5) 

315. Habrocestum morosum (Peckham) , 1888 

Tippecanoe (Lafayette, 25) 

316. Habrocestum pulex (Hentz), 1846 

Tippecanoe (Lafayette, 25, 22; West Lafayette, June 20, 1966, 
TAP); Crawford (Wyandotte, SejJt. 8, 2, 21, 22); INDIANA 
(49); Wayne (Richmond, 20, 21, 22, 39); Porter (Valparaiso, 21, 
22; Ogden Dunes, 39, 22); LaPorte (Smith, 39, 22); Fulton 
(Rochester, July 27, 1967, TAP); Carroll, (Delphi, June 23, 
1966, TAP); Vermillion (Clinton, July 20, 1967, TAP) 

317. Habronattus agilis (Banks), 1893 

Porter (Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22; Ind. 
Dunes State Park, 22) 

318. Habronattus borealis (Banks), 1895 

Steuben (Crooked Lake, 21, 22); Porter (Dunes Acres, 38, 39, 
22; Ogden Dunes, 39, 22; Ind. Dunes State Park, 22) 

319. Habronattus calcaratus (Banks), 1904 

Porter (Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22; Ind. 
Dunes State Park, 22) 

320. Habronattus decorus (Blackwall), 1846 
Pellenes roseus (Hentz), 1846 

Steuben (Crooked Lake, 21, 22); Porter (Valparaiso, 21, 22); 

Huntington (Huntington, July 27, 1967, TAP) 
Hentz's description of Attus roseus was published in June of 1846, 
whereas Blackwall published his description of Salticus decorus in Janu- 
ary of 1846. 

321. Hasarius adansonii (Audouin), 1827 
Sidusa borealis Banks, 1904 

Porter (Dunes Acres, 39, 22; Ind. Dunes State Park, 22) 

322. Hentzia ambigua (Walckenaer), 1837 
W ala palm-arum (Hentz), 1845 

Lake (Hammond, May 29, June 16, July 30, 2, 21, 22; Pine, 
May 25, 2, 21, 22); Posey (Arlington, June 10, 2, 21, 22); IN- 
DIANA (49); Porter (Dunes Acres, 38, 39, 22; Ogden Dunes, 
39, 22); Huntington (Huntington, Aug. 15, 1967, TAP) 

323. Hentzia mitrata (Hentz), 1845 

Tippecanoe (Lafayette, 25); Posey (Arlington, June 10, 21, 22; 
Grand Chain, May 12, June 3, July 8, 2, 21, 22) ; Martin (Shoals, 
May 16, 2, 21, 22); Fountain (Veedersburg, May 12, 2, 21, 22); 
Kosciusko (Vawter Park, June 1, 2, 21, 22); INDIANA (49); 

Entomology 305 

LaPorte (Otis, 53; Smith, 39, 22); Porter (Woodville, 53; 
Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22) 

324. Hyctia Una (Hentz), 1845 

Posey (Mount Vernon, June 8, 1967, TAP) 
This specimen was determined by Wilton Ivie of The American Mu- 
seum of Natural History. He considers Icius formosus Banks, 1892 a 
synonym of Hyctia bina. 

325. Hyctia pikei Peckham, 1888 

Wayne (Richmond, 21, 22) 

326. Icius elegans (Hentz), 1845 

Tippecanoe (Lafayette, 25, 22); Lake (Hammond, May 29, 2, 

21, 22); Posey (Arlington, June 10, 2, 21, 22) ; Knox (July 2, 
2, 21, 22); Starke (Bass Lake, June 22, 2, 21, 22); Martin 
(Shoals, May 16, 2, 21, 22) ; Marshall (Culver, June 29, 2, 21, 
22); Porter (Dunes Acres, 38, 22); LaPorte (Smith, 39, 22) 

327. Icius hartii Emerton, 1891 

Wayne (Richmond, 20, 21, 22, 39); Brown (Nashville, 21, 22); 
LaPorte (Smith, 39) 

328. Icius similis Banks, 1895 

Tippecanoe (West Lafayette, June 20, 1966, TAP) 

329. Maevia inclemens (Walckenaer), 1837 
M. vittata (Hentz), 1845 

M. niger (Hentz), 1845 

Tippecanoe (Lafayette, 25; Ross Biological Reserve, 5); Posey 
(Arlington, June 10, 2, 21, 22); Floyd (New Albany, 2, 21, 22); 
Lake (Hessville, 53; Liverpool, 53); Wayne (Richmond, 20, 21, 

22, 39); Steuben (Crooked Lake, 21, 22); Brown (Nashville, 21, 
22); Porter (Valparaiso, 21, 22; Dunes Acres, 38, 39, 22; Ogden 
Dunes, 39, 22; Ind. Dunes State Park, 22); LaPorte (Smith, 38, 
22); Huntington (Huntington, June 29, 1966, TAP) 

330. Marpissa undata (DeGreer), 1778 

Marptusa familiaris Emerton, 1891 

Tippecanoe (Lafayette, 25; Americus, June 23, 1966, TAP); 
Putnam (Greencastle, 2, 21, 22); Dubois (Huntingburg, 2, 21, 
22); LaPorte (Smith, 39, 22; LaPorte, May 15, 1968, TAP) 

331. Metaphidippus canadensis (Banks), 1897 

Tippecanoe (Ross Biological Reserve, 5) 

332. Metaphidippus flavipedes (Peckham), 1888 

Tippecanoe (Ross Biological Reserve, 5) 

333. Metaphidippus flavus (Peckham) , 1888 

Porter (Dunes Acres, 38, 39, 22) 

334. Metaphidippus galathea (Walckenaer), 1837 

Porter (Ogden Dunes, 39); Tippecanoe (Ross Biological Re- 
serve, 5); Carroll (Delphi, July 7, 1966, TAP) 

306 Indiana Academy of Science 

335. Metaphidippus protervus (Walckenaer), 1837 
Dendryphantes capitatus (Hentz), 1845 

D. octavus (Hentz), 1845 

Tippecanoe (Lafayette, 25, 22; Ross Biological Reserve, 5); 
Lake (Hammond, May 29, June 16, July 30, 2, 21, 22; Pine, 
June 29, 2, 53, 21, 22); Marshall (Arlington, June 10, 2, 21, 22; 
Culver, June 29, 2, 21, 22) ; Posey (Grand Chain, May 12, June 
3, 2, 21, 22); Crawford (Wyandotte, June 25, 26, 2, 21, 22); 
Starke (Bass Lake, June 22, 2, 21, 22); Martin (Shoals, 
May 16, 2, 21, 22); Fountain (Attica, June 20, 2, 21, 22; 
Veedersburg, May 12, 2, 21, 22); Floyd (New Albany, 2, 
21, 22); Kosciusko (Vawter Park, June 1, 2, 21, 22); Porter 
(Ind. Dunes State Park, 53; Valparaiso, 21, 22; Dunes Acres, 
38); Wayne (Richmond, 20, 21, 22); Steuben (Crooked Lake, 
21, 22); Brown (Nashville, 21, 22); LaPorte (Smith, 39, 22) 

336. Neon nellii Peckham, 1888 

Wayne (Richmond, 20, 21, 22); Steuben (Crooked Lake, 21, 22); 
Brown (Nashville, 21, 22); Porter (Valparaiso, 21, 22) 

337. Onondaga lineata (C. L. Koch), 1848 

Tippecanoe (Ross Biological Reserve, 5) 

338. Paraphidippus aurantius (Lucas), 1833 

Attus multicolor Hentz, 1846 

Putnam (Greencastle, 2, 21, 22); Knox (Vincennes, July 10, 2, 
21, 22); Martin (Shoals, July 13, 2, 21, 22), Crawford (Wyan- 
dotte, June 25, 26, 2, 21, 22) ; Posey (Grand Chanin, July 8, 2, 

339. Paraphidippus marginatus (Walckenaer), 1837 
Dendryphantes militaris (Hentz), 1845 

Tippecanoe (Lafayette, 25; Ross Biologoical Reserve, 5); Craw- 
ford (Wyandotte, June 25, 2, 21, 22); Lawrence (Mitchell, July 
15, 2, 21, 22); Porter (Dunes Acres, 38, 39, 22; Ogden Dunes, 
39, 22); LaPorte (Smith, 39, 22); Fulton (Rochester, June 7, 
1966, TAP) 

340. Paraphidip]Jus pineus Kaston, 1945 

Tippecanoe (Ross Biological Reserve, 5); Vermillion (Clinton, 
June 13, 1966, TAP) 

341. Pellenes arizonensis Banks, 1904 

Porter (Ogden Dunes, 39, 22) 

342. Phidippus audax (Hentz), 1844 

Marion (Indianapolis, 31, 22); Tippecanoe (Lafayette, 25, 22, 
Oct. 23, 1966, RWM; West Lafayette, Sept. 20, 1966, AF, Sept. 
31, 1966, TAP, May 15, 1968, KHK) ; Posey (Arlington, June 
10, 2, 21, 22; New Harmony, May 6, Sept. 3, 2, 21, 22; Grand 
Chain, May 12, June 3, 2, 21, 22); Starke (Bass Lake, June 22, 
2, 21, 22); Fountain (Attica, June 20, 2, 21, 22); Jennings 
(North Vernon, May 7, 2, 21, 22); Marshall (Culver, June 29, 2, 
21, 22); Putnam (Greencastle, 2, 21, 22); Dubois (Huntingburg, 

Entomology 307 

2, 21, 22); Wayne (Richmond, 20, 21, 22, 39); Steuben (Crooked 
Lake, 21, 22); Porter (Valparaiso, 21, 22; Dunes Acres, 38, 39, 
22; Ogden Dunes, 39, 22; Ind. Dunes State Park, 22); LaPorte 
(Smith, 39, 22); Benton (Otterbein, Apr. 25, 1968, JOS) 

343. Phidippus cardinalis (Hentz), 1845 

Jennings (North Vernon, Sept. U, 2, 21, 22?) ; INDIANA (49) 
Banks (2) recorded this species from North Vernon, not Mount 
Vernon, as Elliott (22) has indicated. 

344. Phidippus fraudulentus (Walckenaer), 1837 
Dendryphantes insignarius (C. Koch), 1846 

Lake (Hammond, June 16, 2, 21, 22); Posey (Arlington, June 
10, 2, 21, 22); INDIANA (49); Porter (Dunes Acres, 38, 39, 22; 
Ogden Dunes, 39, 22); LaPorte (Smith, 39, 22) 

345. Phidippus mccookii (Peckham), 1883 

Porter (Dunes Acres, 38, 39, 22) 

346. Phidippus princeps (Peckham), 1883 
P. brunneus Emerton, 1891 

Porter (Dunes Acres, 38, 39, 22); LaPorte (Smith, 39, 22) 
Lowrie (38, 39) reported P. brunneus Emerton, not P. brunneus 
F. Cambridge, as Elliott (22) has indicated. Emerton's Phidippus brun- 
neus is a synonym of Phidippus princeps (Peckham), 1883. 

347. Phidippus purpuratus Keyserling, 1884 

Porter (Ogden Dunes, 39, 22) 

348. Phidippus putnami (Peckham), 1883 

Knox (July 2, 21, 22); Posey (Grand Chain, 2, 21, 22); 

349. Phidippus rimatov (Walckenaer) , 1837 
P. clarus Keyserling, 1884 

Putnam (Greencastle, 2, 21, 22); Wayne (Richmond, 20, 21, 22, 
39); Brown (Nashville, 21, 22); Porter (Dunes Acres, 38, 39, 
22; Ogden Dunes, 39, 22); LaPorte (Smith, 39, 22); Tippecanoe 
(July 26, 1967, TAP; Ross Biological Reserve, 5); Carroll 
(Delphi, Aug. 3, 1967, TAP) 
Banks (2) recorded Philaeus rimator Peckham from Greencastle. As 
far as the writer can determine, Philaeus rimator is a synonym of Phidip- 
pus rimator (Walckenaer), 1837 and not of Dendryphantes formosus 
(Peckham), 1883, as Elliott (21, 22) has indicated. The writer bases his 
opinions on Petrunkevitch (49), pages 772 and 630. 

350. Phidippus variegatus (Lucas), 1833 
P. morsitans (Walckenaer), 1837 

Tippecanoe (Lafayette, 25) 

351. Phidippus whitmanii Peckham, 1909 

Tippecanoe (Ross Biological Reserve, 5) 

352. Phlegra fasciata (Hahn), 1831 
P. leopardus (Hentz), 1846 

308 Indiana Academy of Science 

Tippecanoe (Lafayette, 25); Posey (New Harmony Sept. 3, 2, 
21,22); INDIANA (49) 

353. Salticus scenicus (Linnaeus), 1758 

Tippecanoe (Lafayette, 25; West Lafayette, May 20, 1968, 
RWM); Wayne (Richmond, 21, 22); Posey (Mount Vernon, 
Aug. 18, 1966, TAP) 

354. Sitticus palustris (Peckham), 1883 

Porter (Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22; Ind. 
Dunes State Park, 22) 

355. Synemosyna lunata (Walckenaer), 1837 
S. formica Hentz, 1845 

Posey (Arlington, June 10, 2, 21, 22) ; Marshall, (Culver, June 
29, 2, 21, 22); Wayne (Richmond, 20, 21, 22); Tippecanoe 
(Americus, Aug. 10, 1967, TAP); Franklin (Brookville, Sept. 
1, 1966, RWM) 

356. Talavera minuta (Banks), 1895 

Porter (Ogden Dunes, 39, 22); Tippecanoe (Ross Biological 
Reserve, 5) 

357. Thiodina iniquies (Walckenaer), 1837 
T. sylvana (Hentz), 1845 

T. retarius (Hentz), 1850 

Crawford (Wyandotte, June 25, July 25, 2, 21, 22); Posey 
Grand Chain, June 3, 2, 21, 22) ; Dubois ( Huntingburg, 2, 21, 

358. Zygoballus bettini Peckham, 1888 

Knox (July 2, 2, 21, 22); INDIANA (49); Porter (Dunes 
Acres, 38, 39, 22; Ind. Dunes State Park, 22); LaPorte 
(Smith, 39) 
Lowrie (39) collected this species at Smith, not Ogden Dunes, as 
Elliott (22) has indicated. 

369. Zygoballus nervosus (Peckham), 1888 

Porter (Dunes Acres, 39; Ogden Dunes, 39; Ind. Dunes State 
Park, 22) 

FAMILY Dictynidae 

360. Dictyna angulata Emerton, 1915 

Tippecanoe (Ross Biological Reserve, 5) 

361. Dictyna arundinaceoides Keyserling, 1883 

Tippecanoe (Lafayette, 25, 22) 

362. Dictyna bicornis Emerton, 1915 

Porter (Ogden Dunes, 39, 22) 

363. Dictyna bostoniensis Emerton, 1880 

Porter (Dunes Acres, 38, 39, 22; Ogden Dunes, 39, 22) 

364. Dictyna foliacea (Hentz), 1850 

Lake (Miller, 53; Hessville, 53; Liverpool, 53; Clark, 53); 
LaPorte (Otis, 53; Smith, 39, 22); Porter (Woodville, 53; Ind. 

Entomology 309 

Dunes State Park, 53, 22; Valparaiso, 21, 22; Dunes Acres, 38, 
39, 22; Ogden Dunes, 39, 22); Wayne (Richmond, 20, 21, 22, 
39); Steuben (Crooked Lake, 21, 22) 

365. Dictyna formidolosa Gertsch & Ivie, 1936 

D. armata Banks, 1911 (armata preoc. by Thorell, 1875) 
Tippecanoe (Ross Biological Reserve, 5) 

366. Dictyna frondea Emerton, 1888 

Lake (Hammond, May 29, 2, 21, 22); Marshall (Arlington, June 
10, 2, 21, 22); INDIANA (49); Porter (Dunes Acres, 38, 39, 
22; Ogden Dunes, 39, 22; Ind. Dunes State Park, 22) 
Chamberlin and Ivie (12) consider Dictyna foliacea and Dictyna 

frondea separate species, whereas Kaston (32) considers them 


367. Dictyna maxima Banks, 1892 

Tippecanoe (Ross Biological Reserve, 5) 

368. Dictyna minuta Emerton, 1888 

Wayne (Richmond, 20, 21, 22); Steuben (Crooked Lake, 21, 
22); Porter (Valparaiso, 21, 22) 

369. Dictyna sublata (Hentz),1850 

Kosciusko (Tippecanoe Lake, June 8, 2, 21, 22; Vawter Park, 
June 1, 2, 21, 22); Posey (Grand Chain, June 3, 2, 21, 22; Mount 
Vernon, Aug. 18, 1966, June 8, 1967, TAP); Wayne (Richmond, 

20, 21, 22); Steuben (Crooked Lake, 21, 22); Porter (Valpa- 
raiso, 21, 22); Tippecanoe (Ross Biological Reserve, 5; Ameri- 
cus, June 23, 1966, TAP); Fulton (Rochester, June 7, 1966, 
Sept. 16, 1967, TAP) ; Pulaski (Winamac, June 15, 16, 1966, 
TAP); Huntington (Huntington, June 29, 1966, TAP); Cass 
(Logansport, July 6, 1966, Aug. 3, 1967, TAP); Gibson (Mount 
Carmel, July 27, 1966, TAP); Sullivan (Merom Station, July 

21, 1966, July 18, 1967, TAP) ; LaPorte (LaPorte, June 19, 
1968, TAP) 

370. Dictyna volucripes Keyserling, 1881 

Porter (Dunes Acres, 38, 39, 22); LaPorte (Smith, 39, 22) 

371. Lathys foxii (Marx), 1891 

Wayne (Richmond, 20, 21, 22) 

372. Scotolathys maculatus (Banks), 1900 

Tippecanoe (Ross Biological Reserve, 5) 

373. Scotolathys pallidus (Marx), 1891 
Lathys pallida (Emerton), 1894 

Wayne (Richmond, 20, 21, 22); Tippecanoe (Ross Biological 
Reserve, 5) 

family Uloboridae 

374. Hyptiotes cavatus (Hentz), 1847 

Tippecanoe (Lafayette, 25, 22); Wayne (Richmond, 20, 21, 22, 
39); Steuben (Crooked Lake, 21, 22); Porter (Ogden Dunes, 
38, 22); LaPorte (Smith, 39, 22) 

310 Indiana Academy of Science 

375. Uloborus glomosus (Walckenaer), 1841 
U. americanus (of most authors) 

Wayne (Richmond, 20, 21, 22, 39); Steuben (Crooked Lake, 21, 
22); Porter (Valparaiso, 21, 22; Ogden Dunes, 39, 22); LaPorte 
(Smith, 39, 22) 

Chamberlin and Ivie (12) page 139, say that Uloborus americanus 
Walckenaer, 1841 is a synonym of Thanatidius americanus (Walckenaer), 
1841. T. americanus is in the family Pisauridae. 

The U. americanus of most authors is actually U. glo?nosus 
(Walckenaer), 1841. Kaston (32), page 513, states, "According to 
Chamberlin and Ivie (1944) americanus is another species and ours is 
really glomosus Walckenaer 1841." 

family Amaurobiidae 

376. Amaurobius bennetti (Blackwall), 1846 

Monroe (Mayfield's Cave, 4, 22; Truett's Cave?, 2, 21, 22); 
Wayne (Richmond, 20, 21, 22, 39); Steuben (Crooked Lake, 21, 
22); LaPorte (Smith, 39, 22); Putnam (Bainbridge, Apr. 23, 

1967, TAP) 

Banta (4), page 59, collected two specimens of this species in 
Mayfield's Cave. He makes no mention of a Truett's Cave collection. 
Banks (2) determined Banta's specimens and reported them from Truett's 
Cave. Banks was in error; he should have reported this species from 
Mayfield's Cave, not from Truett's Cave. 

377. Amaurobius fer ox (Walckenaer), 1837 

Tippecanoe (Lafayette, 25, 22); White (Brookston, June 12, 

1968, TH) 

378. Titanoeca americana Emerton, 1888 

Wayne (Richmond, 20, 21, 22, 39); Porter (Dunes Acres, 38, 39, 
22; Ind. Dunes State Park, 22); Tippecanoe (Merritt's Pine 
Plantation, May 12, 1967, RWM) 

Appendix I 

NB— Banks, Nathan KHK— Knauer, Kenneth H. 

CB — Bates, Cova GL — LaRocca, George 

JWB— Brewer, J. Wayne RWM— Meyer, Robert W. 

JJD— Davis, John J. DJM— Miller, Dale J. 

RED— Dolphin, Robert E. MKP— Parker, Martha K. 

JJF— Favinger, John J. TAP— Parker, Thomas A. 

GRF— Finni, Gary R. ES— Saugstad, Ed 

AF — Frishman, Austin JOS — Sillings, John 0. 

RLG— Giese, Ronald L. HS— Speith, H. 

JDH — Hacker, Jan D. JEW — Wappes, James E. 

TH— Henry, Thomas GLW — Ward, Gertrude L. 

TRH— Hintz, Thomas R. DEW— Weaver, Dix E. 

Entomology 311 

Appendix II 
Obscure Collecting Localities 

Americus, Tippecanoe County — The collections were made along the 
Tippecanoe River, approximately one mile upstream from the junction of 
the Tippecanoe and Wabash Rivers. 

Arlington, Marshall County — Banks (2) states that this town was in 
Marshall County, not the Arlington in Rush County. The writer cannot 
determine its exact location in the county. 

Buffington, Lake County — This town was formerly called Edgemoor 
and was located in section 26 of Calumet Township. It was later taken 
into Gary. 

Clark, Lake County — This town was located on the eastern edge of 
North Township which is now Hammond. 

Cypress Swamp, Knox County — This area, also known as Great 
Cypress Swamp, is approximately two miles northeast of the confluence 
of the Wabash and White Rivers. 

Entomological Research Area, Tippecanoe County — This collecting 
area is part of the Horticultural Farm which lies approximately two 
miles west of the Purdue Campus on Sharon Chapel Road. 

Grand Chain, Posey County — This area is approximately eight miles 
south of New Harmony along the Wabash River. 

Hessville, Lake County — This town, which was in North Township, 
is now part of Hammond. 

Liverpool, Lake County — This town is located at the western edge of 
Hobart Township. 

Merritt's Pine Plantation, Tippecanoe County — This privately-owned 
Christmas tree plantation is located approximately % mile east of the 
Wildcat Creek bridge on the south side of State Road 25. 

Miller, Lake County — This town was located in section 6 of Hobart 
Township and is now part of Gary. 

Mount Carmel, Gibson County — Although Mount Carmel is actually 
situated on the west bank of the Wabash River (Illinois), all collections 
were made on the east bank of the river (Indiana). 

Mount Vernon, Posey County — All collections were made along the 
east bank of the Wabash River which is approximately eight miles due 
west of Mount Vernon. The Mount Vernon record for Loxosceles reclusa 
was taken in the town, not along the Wabash. 

Otis, LaPorte County — This town is in New Durham Township. 

Pine, Lake County — This town was situated along Lake Michigan 
and is now the north end of Clark Street in Gary. 

Ross Biological Reserve, Tippecanoe County — This preserved area 
lies along the Wabash River adjacent to the staff recreation area, "The 
Hills." These areas are located approximately ten miles southwest of 
West Lafayette. 

312 Indiana Academy of Science 

Smith, LaPorte County — This small town is located at the fork of 
Fail Road where it turns into 175 East Road and 125 East Road, 1.2 miles 
east of Tee Lake in Galena Township. The town is approximately 15 
miles due east of Michigan City. 

Woodville, Porter County — This town is in Liberty Township which 
is north of Valparaiso. 

Literature Cited 

Banks, N. 1897. Indiana caves and their fauna, p. 202-204. In: Indiana 
Dept. Geol. Natur. Resources, 21st Annual Report for 1896. Indianapolis. 

. 1907. A preliminary list of the Arachnida of Indiana with 

keys to the families and genera of spiders, p. 715-747. In: Indiana Dept. 
Geol. Natur. Resources, 31st Annual Report for 1906. Indianapolis. 

. 1910. Catalogue of Nearctic spiders. Bull. U. S. Nat. Mus. 


4. Banta, A. M. 1907. The fauna of Mayfield's Cave. Carnegie Inst. Wash. 

5. Becker, H. 1951. A quantitative study of the vertical distribution of spiders 
of the Ross Biological Reserve. Unpublished M.S. Thesis, Purdue University. 
63 p. 

6. Blatchley, W. S. 1897. Indiana caves and their fauna, p. 121-212. In: Indi- 

ana Dept. Geol. Natur. Resources, 21st Annual Report for 1896. Indianapolis. 

7. Cambridge, F. O. P. 1897-1905. Arachnida, Araneida. Vol. II, in Biologia 
Centrali- Americana. R. H. Porter, London. 610 p. 

S. Cambridge, O. P. 1889-1902. Arachnida, Araneida. Vol. I, in Biologia Cen- 
trali-Americana. R. H. Porter, London. 317 p. 

9. Chamberlin, R. V. 1922. The North American spiders of the family Gnapho- 
sidae. Proc. Biol. Soc. Wash. 35: 145-17 2. 

10. Chamberlain, R. V., and W. Ivie. 1942. A hundred new species of Ameri- 
can spiders. Bull Univ. Utah 32(13) ; Biol. Series7(l) : 1-1 17. 

11. — — . 1943. New genera and species of North American linyphiid 

spiders. Bull. Univ. Utah 33(10) ; Biol. Series 7(6) :l-39. 

12. . 1944. Spiders of the Georgia region of North America. Bull. 

Univ. Utah 35(9) ; Biol. Series 8(5) :l-267. 

13. Chickering, A. M. 1957. The genus Tetragnatha (Araneae, Argiopidae) in 
Jamaica, B.W.I., and other neighboring islands. Brevoria, Mus. Comp. Zool. 

14. — . 1959. The genus Tetragnatha (Araneae, Argiopidae) in 

Michigan. Bull. Mus. Comp. Zool. 119(9) :475-499. 

15. . 1962. The genus Tetragnatha (Araneae, Argiopidae) in Ja- 
maica, W. I. Bull. Mus. Comp. Zool. 127(8) :423-450. 

16. Comstock, J. H. 1912. The spider book. Garden City, N. Y. 725 p. 

17. . 1940. The spider book. Revised and edited by W. J. Gertsch. 

Doubleday, Doran and Co., New York. 729 p. 

18. Cope, E. D. 1872. On the Wyandotte Cave and its fauna. Amer. Natur. 

19. Dondale, C. D. 1961. Revision of the aureolas group of the genus Philodro- 
mus (Araneae: Thomisidae) in North America. Can. Entomol. 93(3) :199- 


20. Elliott,, F. R. 1930. An ecological study of the spiders of the beech-maple 
forest. Ohio J. Sci. 30:1-22. 

Entomology 313 

21. . 1932. Revisions and additions to the list of Araneae (spiders) 

of Indiana. Proc. Indiana Acad. Sci. 41 :419-430. 

22. . 1953. The araneology of Indiana. Proc. Indiana Acad. Sci. 


23. Emerton, J. H. 1875. Notes on spiders from caves in Kentucky, Virginia, 
and Indiana, Amer. Natur. 9:278-281. 

24. . 1912. Four burrowing- Lycosa (Geohjcosa Montg. Scaptocosa 

Banks) including one new species. Psyche 19:25-36. 

25. Fox, W. H. 1892. A list of spiders from Indiana. Proc. Entomol. Soc. Wash., 
D. C. 2(2) :267-269. 

2 6. Gertsch, W. J. 1933. Diagnosis of new American spiders. Amer. Mus. Novi- 

tates 637 :1-14. 

27. . 1939. A revision of the typical crab-spiders (Misumeninae) of 

America north of Mexico. Bull. Amer. Mus. Natur. Hist. 76:277-442. 

28. Hbntz, N. M. 1832. On North American spiders. Amer. J. Sci. Arts 21(1) :99- 

29. . 1847. Descriptions and figures of the Araneides of the United 

States. J. Boston Soc. Natur. Hist. 5:443-478. 

30. . 1850. Descriptions and figures of the Araneides of the United 

States. J. Boston Soc. Natur. Hist. 6:18-35, 271-295. 

31. Hentz, N. M., ed. S. Scudder. 1S67. Supplement to the descriptions and fig- 
ures of the Araneides of the United States. Proc. Boston Soc. Natur. Hist. 

32. Kaston, B. J. 194S. Spiders of Connecticut. Conn. State Geol. and Natur. 
Hist. Survey Bull. 70:1-874. 

33. Kinter, E. 1935. Some spiders of the genus Tetragnatha. Proc. Indiana 
Acad. Sci. 44:207-209. 

3 4. Levi, H. W. 1951. New and rare spiders from Wisconsin and adjacent 

states. Amer. Mus. Novitates 1501:1-41. 

3 5. . 1959. The spider genus Latrodectus (Araneae, Theridiidae). 

Trans. Amer. Microscopical Soc. 78(1) :7-43. 

36. Levi, H. W., and H. M. Field. 1954. The spiders of Wisconsin. Amer. Midi. 
Natur. 51:440-467. 

37. Lindset, A. A., R. O. Petty, W. VanAsdel, and D. K. Sterling. 1960. 
Vegetation and environment along the Wabash and Tippecanoe Rivers. Ecol. 
Mono. 31 :105-156. 

38. Lowrie, D. C. 1942. The ecology of the spiders of the xeric dunelands in 
the Chicago area. Bull. Chicago Acad. Sci. 6(9) :161-189. 

39. . 1948. The ecological succession of spiders of the Chicago 

area dunes. Ecology 29(3) :334-351. 

40. Marx, G. 1890. Catalogue of the described Araneae of temperate North 
America. Proc. U.S. Nat. Mus. 12:497-594. 

41. McCook, H. C. 1894. American spiders and their spinningwork. Vol. III. 
Acad. Natur. Sci. Philadelphia. 40 6. p. 

42. McCrone, J. D., and H. W. Levi. 1964. North American widow spiders of 
the Latrodectus curacaviensis group (Araneae: Theridiidae). Psyche 
71(1) :12-27. 

43. McIndoo, N. E. 1910. Biology of the Shawnee Cave spiders. Biol. Bull. 
Woods Hole, Mass. 19(6) :303-323. 

44. ■ — — . 1911. Notes on some arachnids from Ohio Valley caves. Biol. 

Bull. Woods Hole, Mass. 20(3) :1S3-1S6. 

45. Minton, S. A. 1956. The trap-door spider Pachylomerides adouinii, in 
southern Indiana. Proc. Indiana Acad. Sci. 64:255. 

314 Indiana Academy of Science 

46. Minton, S. A., and C. Olson. 1964. A case of spider bite with severe hemo- 
lytic reaction. Pediatrics 33(2) :283-284. 

47. Packard, A. S. 1875. The invertebrate cave fauna of Kentucky and adjoin- 
ing states. Amer. Natur. 9:274-278. 

48. . 1888. The cave fauna of North America, with remarks on 

the anatomy of the brain and origin of the blind species. Nat. Acad. Sci. 
Memoir 4:3-156. 

49. Petrunkevitch, A. 1911. A synonymic index-catalogue of spiders of North, 
Central, and South America with all adjacent islands. Bull. Amer. Mus. 
Natur. Hist. 29:1-809. 

50. . 192 5. Descriptions of new or inadequately-known American 

spiders. Ann. Entomol. Soc. Amer. 18:313-322. 

51. . 1939. Catalogue of American spiders. Part I. Conn. Acad. 

Arts Sci. 33:133-338. 

52. Seeley, R. M. 192S. Revision of the spider genus Tetragnatha. Bull. New 
York State Mus. 278:99-147. 

53. Shelford, V. E. 1912. Ecological succession. IV. Vegetation and the control 
of land animal communities. Biol. Bull. 23:59-99. 

54. Wallace, H. K. 1942. A revision of the burrowing spiders of the genus 

Geolycosa. Amer. Midi. Natur. 27:1-62. 

55. Williamson, E. B. 1900. Biological conditions of Round and Shriner Lakes, 
Whitley County, Ind. Proc. Indiana Acad. Sci. 9:151-155. 


Chairman: Lowell I. Dillon, Ball State University. 
Wilton N. Melhorn, Purdue University, was elected chairman for 1969. 


Gypsum Resources of the Midwestern United States. John H. Cleveland 
and Carol F. Tiefel, Indiana State University. — Gypsum (CaS04 # 2H 2 0) 
as an economic mineral is primarily utilized in the construction industry 
for prefabricated plasterboard. Although the value of raw gypsum is less 
than $5.00 per ton, its production is economically significant because the 
fabricated products are normally produced at plants adjacent to the mine 
and constitute a ten-fold increase in value. Commercial gypsum deposits 
must have large reserves, minable thickness, moderate depth, and a loca- 
tion near major population centers. Present regional production is con- 
fined to six districts in four states (Indiana, Iowa, Michigan, and Ohio) 
and is obtained from rocks of four different geologic periods (Silurian, 
Devonian, Mississippian, and Permian?). All of the region's known 
commercial gypsum deposits are sedimentary in origin and genetically 
related to evaporite basins, but are the final product of recent rehydration 
of anhydrite at shallow depths. Contrary to prevailing mining industry 
trends, quarry operations are being replaced by underground mines with 
future development likely in the form of multipurpose underground mines 
in close proximity to major populations. Recent published investigations 
of newly recognized gypsum reserves at LaPorte, Indiana and Albia, 
Iowa suggest they may fit this pattern. 

A Significant Exposure of Pleistocene Drift in South-Central Indiana.* 

Allan F. Schneider, Indiana Geological Survey. — Unconsolidated 
deposits of the Pleistocene Series are exposed in a railroad cut at the 
glacial boundary in northwestern Brown County. The section is signifi- 
cant because it exhibits a more complete sequence of Pleistocene sedi- 
ments than other exposures in the area and because it corroborates W. J. 
Wayne's conclusion that in northwestern Brown County glacier ice of the 
Kansan Age advanced somewhat farther than ice of the Illionian Age. 

Siltstone of the Borden Group (Mississippian System) at the base of 
the cut is overlain unconformably by 3% feet of unoxidized and oxidized 
till interpreted as Kansan in age. The till is overlain by about 30 feet of 
fine gravel and pebbly sand, much of which is strongly oxidized. In its 
upper part this deposit is altered to a sticky reddish-brown sandy mate- 
rial that clearly represents a significant period of weathering. The 
gravel and sand is regarded as Illinoian outwash, and the weathering pro- 
file is interpreted as having formed during the interglacial Sangamonian 

The section is capped by about 6 feet of silt, which is probably of 
eolian rather than lacustrine origin. In its lower part the loess is 

1 Publication authorized by the State Geologist, Department of Natural Re- 
sources, Geological Survey. 


316 Indiana Academy of Science 

clayey and similar in color to the underlying weathered outwash but 
grades upward into more typical noncalcareous loess. The reddish-brown 
silt is tentatively considered to be Illinoian and Sangamonian and the 
upper part Wisconsinan in age. 

Unconsolidated Deposits on the Mitchell Plain of Indiana. i Richard L. 
Powell, Indiana Geologicol Survey. — The Mitchell Plain of south-central 
Indiana is a limestone plateau partly mantled with unconsolidated ma- 
terials that consist mostly of loess, gravels, and clays that attain 60 feet 
in thickness. Most of the material is clay in layers of red, brown, or 
olive and yellow. In places the clay overlies coarser sedimnts. A persis- 
tent bed of cherty gravel, which in places contains a few allochthonous 
geode fragments and lenses of sandstone pebbles, overlies the clays 
where they have not been eroded. The cherty gravel bed lies at an alti- 
tude similar to that of some Lafayette gravel deposits along entrenched 
meandering drainage routes. The present entrenched drainage pattern 
evolved from drainage on a westward-sloping erosion and deposition 
surface that had developed on the Mitchell Plain and the Norman Upland 
by the time that the cherty gravel bed was deposited. Entrenchment of 
the major streams during late Tertiary or early Pleistocene time and 
during mid-Pleistocene time accelerated cavern and sinkhole develop- 
ment on the Mitchell Plain and was accompanied by erosion of the clays 
and gravels and partial redeposition in sinkholes and caverns. Loess of 
late Pleistocene age blanketed the region prior to recent erosion. 

1 Published with permission of the State Geologist, Indiana Geological 

Analysis of Surficial Landform Properties: The Regionalization 
of Indiana into Units of Morphometric Similarity 1 

Laurence A. Lewis, Temple University 


This study offers a more rigorous alternative to the traditional reli- 
ance on qualitative data and the subjective approach in the regionalization 
of geomorphic units. Using analytical data and techniques, regional geo- 
morphic units are developed for Indiana. The results of a principal com- 
ponets analysis and a cluster analysis indicate that Indiana can be region- 
alized using three primary morphometric components: a vertical componet, 
an areal magnitude factor and a stream network development factor. In 
addition, a lacustrine factor is found to be of secondary importance within 

The areal patterns of these morphomeric components indicate that the 
previous physiographic or regional geomorphic units developed for Indiana 
fail in delimiting the State into regions of form similarity. The previous 
investigations, by concentrating on geologic structure and/or process in- 
stead of the actual topographic expression, most likely account for this 


In previous physiographic or regional geomorphic studies, the cri- 
teria utilized in delimiting "natural" regions have been highly subjective 
and somewhat inconsistent. Traditionally, regional geomorphic studies 
have relied heavily on qualitative descriptions of bedrock structure, 
general landform characteristics, geologic history and/or climatic charac- 
teristics as the basis for forming different regions. Likewise, criteria 
which are considered crucial for the location of regional boundaries vary 
from area to area in the regional studies of Fenneman (1), Thornbury 
(4) and Hunt (3). Hammond has reported on the need to use the iden- 
tical criteria throughout a single study and has attempted to minimize 
subjective data by substituting measurable landform characteristics as 
the standard for delimiting landform regions (2). In his regionalization 
of the United States, Hammond utilized four variables: slope inclina- 
tion, vertical dimension, general profile character and some aspects of 
surface material. While Hammond's work represented a start, it did 
not satisfactorily eliminate the problem of subjective data or completely 
meet the needs of geomorphology. First, only two of the variables used 
could be measured — slope inclination and vertical dimension; the two 
remaining variables are subjective in nature. Second, the purpose of 
the study was to develop regions that would indicate areas of similar 
land-use. From a geomorphic point of view, this is not a particularly 
useful purpose. Clearly, the need still exists to use analytical criteria in 
the construction of regions that meet the needs of geomorphology. 

A landform assemblage results from the interaction of three basic 
factors: process, time and geologic structure. Only the result of these 

1 The author wishes to acknowledge the assistance of Johnson Akin- 
bola and Daniel Knuth in the collection of data and for their valuable 



Indiana Academy of Science 

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Geology and Geography 


Sample Points 
GUI -1.556 to -0.582 
□ -0.548 to 0.540 
0.556 to 1.538 
(tactor scores) 

Figure I: Vertical Component 

320 Indiana Academy of Science 

three factors, namely topographic expression, should be the prime con- 
cern when delimiting geomorphic regions. Therefore, the criteria used in 
the formation of geomorphic regions should be limited to surficial form 
elements. If one of the primary factors strongly affects a given area, 
the morphometric properties of the surface will reflect the causal in- 

This study is the initial report of an attempt to delimit geomorphic 
regions solely using quantitatively measurable data. By limiting the 
criteria to variables that can be measured, analytical techniques can be 
used to develop regions that will minimize subjectivity and permit re- 
liable comparisons between features found in different areas. These re- 
gions, where the form elements are evaluated numerically, in turn may 
be used to evaluate the controls of the environment in individual process 
oriented geomorphic studies. Specifically, this study regionalizes the 
State of Indiana into areas of morphometric similarity. 


Thirteen variables (Table 1), all of which have been utilized in previ- 
ous geomorphic studies, form the basis for the regionalization of Indiana 
into similar morphometric areas. This set of variables measures the ma- 
jor surficial properties of landforms. The data for these variables are 
obtained from U.S.G.S. topographic maps. 

Within a general systems framework, the stream basin can be con- 
sidered the basic or prevalent geomorphic system; furthermore, within 
the United States, all land areas are affected by streams and their re- 
lated processes. For these reasons, stream basins are chosen as the basic 
unit of measurement in this study. With the exception of one variable 
(percent of water), all of the variables in this study are calculated from 
4th order stream basins. 

A random sample of sixty quadrangles was drawn from all the 
U.S.G.S. topographic maps (scale 1:24,000) that cover Indiana. The 
sample size insured ample areal coverage of the whole State (Figure 1). 
On each quadrangle drawn from the sample, the center of the map 
served as the base from which the nearest fourth order basin was lo- 
cated. As an indication of basin size, a fourth order basin might extend 
over six quadrangles. For the variable, percentage of water, only the 
original quadrangle drawn in the sample was utilized. 


The multiplicity of variables used in this study requires simplifica- 
tion in order to remove redundancy among the variables and to discover 
the contrasting morphometric elements. This can be achieved by utiliz- 
ing the technique of principal components analysis. The purpose of using 
this technique is to investigate how much of the total variance within 
the sixty stream basins, exhibited in the thirteen variables, can be ac- 
counted for by a smaller number of new "principal components." These 

Geology and Geography 









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Indiana Academy of Science 

principal components will form the basis for regionalizing Indiana into 
morphometric areas. 

Transformation of ten of the variables into their logarithmic equiva- 
lent was required to satisfy the assumptions of normality and linearity 
for the data (Table 2). After the transformations, the correlation matrix 
was subjected to a principal axis solution in which five of the eigen- 
vectors (those having values greater than 1.00) were rotated to a normal 
varimax position. These five orthogonal factors account for approxi- 
mately eighty-three percent of the total variance in the original data 
matrix (Table 3). From examination of the varimax solution, these five 
factors designated the vertical or ''steepness" factor, the horizontal or 
areal magnitude factor, level of stream network development, basin 
shape and the lacustrine factor respectively. 

With the isolation of the five principal components, the next step in 
the regionalization is to aggregate the sixty individual factor scores into 
groups of similarity. The degree of similarity among the factor scores 
is determined by the distance between each observation in factor space. 
Close points are considered similar to each other; distant points or ag- 
gregates of points distant from each other are considered dissimilar. 

The Vertical Component. The vertical component, which accounts 
for 36.3 percent of the total variance (Table 3), is identified primarily 
with relief, the slope of the land, and stream gradient (Table 2). Since 
this component accounts for the largest proportion of the total variance, 
it appears to be the most important surficial element of variation within 

TABLE 3. Percent of Communality Over the Factors and Eigenvalues 

Percent of Communality Over: 

All 13 

5 Rotated 

Factor Number 




































































-1.556 to 












Geology and Geography 


ED -1.165 to -0.539 
[ | -0.439 to 0.384 
[Ml 0.416 to 1.205 
(factor scores) 

Figure 2: Areal Magnitude 


Indiana Academy of Science 

□ -0.695 to 622 
HI 0717 to 2.240 
(factor scores) 

Figure 3: Stream Network Development 

Geology and Geography 325 

Indiana. Inspection of the plot of the factor scores for the first com- 
ponent indicated that the vertical factor aggregates into three basic 

Figure 1 illustrates the areal expression of the vertical component for 
the three factor groupings. 

The general pattern of vertical land expression increases toward the 
south. The three regions, as delimited on the map, show no simple correla- 
tion to the previous more genetic oriented regions. While it is true that 
the area of maximum vertical expression is primarily located in areas not 
glaciated during the Wisconsin Period, it is not limited to the non- 
glaciated portion of the State. Likewise, physiographic regions such as 
the Crawford upland and Mitchell plain appear not to be valid divisions 
in regard to morphometric elements. This is substantiated further by the 
second and third components. 

Areal Magnitude Factor. The highest factor loadings on the second 
component are essentially measures of stream basin extensiveness. Simi- 
lar to the first component, the plot of the factor scores for the magnitude 
factor indicate three grouping: 

group one — 1.165 to — 0.439 

group two —0.539 to 0.384 

group three 0.416 to 1.205 

The areal expression of these three factor groupings are delimited on 
Figure 2. As with the first component, no strong correlations between 
the previous physiographic units of glaciated and non-glaciated areas 
and various glacial phenomena are reflected by this component. 

Stream Network Development Factor. This component represents 
the last major dimension contributing to morphometric variation within 
the State (Table 3). The remaining two components account for only 
16 percent of the total variability. The plot of the factor scores for this 
component indicates the following groupings: 

group one —1.870 to —0.796 

group two —0.695 to 0.622 

group three 0.717 to 2.240 

The areal expression of these groupings is given on Figure 3. The areas 
of maximum development follow the major stream systems within the 
State regardless of the underlying materials. This would seem to indicate 
that if physiographic units use genetic criteria as the basis for division, 
more emphasis should be placed on current-day processes than past 

The Shape Factor. It is suggested that the shape component, which 
is heavily weighted only by one variable (Table 2), be disregarded as a 
criterion for regionalization. First, inspection of the factor score plot 
showed no apparent groupings. The values appeared as a continuum. 
Second, inspection of the areal expression of the absolute values of the 
factor scores indicated a random pattern throughout the State. The be- 
havior of this component could be interpreted as indicating that stream 


Indiana Academy of Science 






[MO Less than' 0.514 
HH Greater than 1.080 
(factor scores) 

Figure 4: Lacustrine Factor 

Geology and Geography 327 

basin shape is purely a result of random processes. However, it probably 
reflects the difficulty of developing any quantitative measure of shape 
without the use of vectors. 

The Lacustrine Factor. With the fifth component accounting for 
only 7.9 percent of the total variability, it is considered to be of second- 
ary importance when delimiting regions within Indiana. The plot of the 
factor scores indicated a dichotomous arrangement : 

group one less than 0.514 

group two greater than 1.080 

Figure 4 shows that the land areas in the immediate vicinity of Lake 
Michigan and the northeastern portion of Indiana are different from 
the remaining portion with regard to this component. 


This study offers a more rigorous alternative to the traditional re- 
liance on qualitative data and the subjective approach in the regional- 
ization of geomorphic units. Using analytical data and techniques, re- 
gional geomorphic units are developed for Indiana. Since the individual 
form elements within the regions can be evaluated numerically, reliable 
comparisons between different areas are possible in contrast to the sub- 
jective regional comparisons that must result when using the traditional 

This study establishes that three basic factors determine the vari- 
ance of morphometric properties throughout Indiana; namely, the verti- 
cal expression of the land, the horizontal expression of the land and the 
development of stream networks. While this study successfully isolates 
areas having similar morphometric properties, no attempt is made to 
combine the three primary components into a composite regionalization. 
The need still exists to investigate other areas in order to develop a 
better understanding of the exact values of the factor scores and to deter- 
mine if the limits of the factor scores used in this study can be applied 
universally before a composite classification is developed. 

From the inspection of the areal pattern of the three components, it 
appears that previous physiographic studies do not delimit Indiana into 
regions of form similarity. Most likely, this results from their over 
emphasis on geologic structure and geologic history as the basic criteria 
for regionalization. By concentrating on the actual topographic expres- 
sion and using analytical techniques offered in this study, the undue in- 
fluence of a single contributing factor, such as geologic structure, is 
prevented. If a purpose of regional geomorphic studies is to complement 
process-oriented geomorphic studies, regionalization using analytical pro- 
cedures will be more fruitful than the traditional approach. 

328 Indiana Academy of Science 

Literature Cited 

1. Fenneman, Nevin M. 1938. Physiography of Eastern United States. McGraw- 
Hill Book Company, Inc. New York. 

2. Hammond, Edwin H. 1964. Analysis of Properties in Land Form Geography: 
An Application to Broad-Scale Land Form Mapping. Ann. Assoc. Amer. 
Geographers 54 (1) 11-18. 

3. Hunt, Charles B. 1967. Physiography of the United States. W. H. Freeman 
and Company, San Francisco. 

4. Thornbury, William D. 1965. Regional Geomorphology of the United States. 
John Wiley & Sons, Inc., New York. 

Earth Science Teaching in the Secondary Schools of Indiana 

Rogek F. Boneham, Indiana University at Kokomo 

An earth science course in the secondary schools of Indiana is a 
relatively new innovation. The purpose of this study was to provide in- 
formation about earth science teachers and to learn what are some of 
the problems they are faced with in conducting this new course. 

This secondary school instruction in earth science will have a direct 
bearing on the enrollments in those college departments which deal with 
some phase of earth science. Some students with an inclination toward 
a scientific career may become interested in earth science after taking 
such a course and decide to major in one of its numerous branches in 

The responsibility for maintaining viable earth science courses in 
secondary schools rests, of course, with the teachers and supervisors of 
the respective school systems. However, we teachers at the college level 
cannot divorce ourselves completly from the secondary school system. 
Many of its students ultimately become our students. If we can help 
them become better prepared for college study then it is beneficial to 
both the students and ourselves. The Earth Science Curriculum Project 
was just such an acknowledgment of our responsibility toward better 
secondary school education. 

The Indiana University School of Science has a list of secondary 
school earth science teachers in Indiana. A three-page questionnaire was 
sent to the 165 teachers on this list. Seventy-three questionnaires were 
returned, sixty-eight from earth science teachers and five from teachers 
who did not teach earth science. It is likely that a number of teachers 
who did not return the questionnaire do not teach earth science. 

The following list contains the counties in which there are earth 
science teachers: 

County with 9 teachers: Marion 

Counties with 6 teachers : Allen, Lake 

County with 5 teachers: Bartholomew 

County with 4 teachers : Delaware 

County with 3 teachers: Porter 

Counties with 2 teachers: Fulton, Hamilton, Howard, Jackson, Madison, 
Montgomery, Tipton, Vigo 

Counties with 1 teacher: Boone, DeKalb, Dubois, Elkhart, Harrison, Hen- 
dricks, Henry, Jefferson, Knox, Kosciusko, LaPorte, Noble, Orange, 
Parke, Posey, Randolph, Spencer, Sullivan, Vermillion, Wayne 

A few of the respondents taught seventh or eighth grade courses in 
earth science. These classes were not included in this survey, since I was 
concerned with senior high school. However, the teachers were included 
with the senior high school teachers of this survey. 


330 Indiana Academy of Science 

The large majority of school systems offered a two-semester course 
rather than a one-semester course. The median number of weeks devoted 
to each phase of earth science in the two-semester courses were: 

Introduction 2 weeks 

Geology 16 weeks 

Astronomy 6 weeks 

Meteorology 6 weeks 

Oceanography 4 weeks 

In the senior high school for 1967-68, there were 3924 students in 142 
sections for a mean of 27 students per section. The approximate enroll- 
ment for 1966-67 was 3566 students. This shows an increase of approxi- 
mately 350 students during 1967-68 over 1966-67 or approximately 13 
sections of 27 students. 

Earth science teaching in Indiana really began in the last decade as 
the following list shows: 

Number of schools offering earth science for the first time 

1928 1958 1960 1962 1963 1964 1965 1966 1967 

1212335 18 5 

Earth science is taught in various schools from grade 7 through 
grade 12. The schools were divided into 32 teaching the course to a 
single grade and 29 teaching multiple grade sections. The majority 
taught earth science in grades 8-10. Two schools offered it to seventh 
graders and 21 offered it to grades 11 and 12. 

The teachers were requested to supply information about the aca- 
demic ability of their students. The following list is the median 
percentages of earth science students: 

College caliber 30% 

Terminal high school 60% 

Potential drop-outs 10% 

There was approximately a 3:1 favorable response to the question, 
"Should the earth science curriculum be expanded?" Those who favored 
curriculum expansion usually listed more than one of the following: 
more laboratory equipment (34 teaechers); more laboratory space (24 
teachers); more library volumes (20 teachers); more teachers (6 teach- 

The greatest problems of earth science teachers were mainly: low 
ability of students (21 teachers); lack of equipment (19 teachers); lack 
of background courses of respondent (16 teachers); lack of space (12 
teachers); lack of library volumes (10 teachers); lack of student interest 
(9 teachers); lack of administration interest (4 teachers). 

Approximately 30% of the teachers used the text prepared under 
the supervision of the Earth Science Curriculum Project. In view of the 
fact that the text has been available for a short period of time, it is 
being used by a large number of students in Indiana. 

Geology and Geography 331 

The two training areas which the teachers found most beneficial to 
their teaching were: courses attended on a degree program and N.S.F. 
Summer Institutes. To a lesser extent were: in-service-training and 
field work. 

Slightly more than half of the teachers had never attended an N.S.F. 
Summer Institute. The great majority of those who had attended an 
institute replied that they had attended either one or two. 

In nearly every case, those who took an N.S.F. Summer Institute 
course in earth science listed the Institute as their most helpful training 
for teaching earth science. 

The median number of years as a teacher listed by the respondents 
was eight. The median number of years as an earth science teacher was 
two. This tends to indicate that the majority of the earth science teachers 
were not originally trained as such. This point is further proven later 
in the report. 

Approximately 65% of the teachers had a master's degree. Those 
with a bachelor's degree only had a median of 6-10 hours beyond the 
bachelor's degree. Those with a master's degree had a median of 20+ 
hours beyond the master's degree. 

The science background of the teachers was examined. The following 
list gives the particular science and the median number of semester hours 
credits of the teachers. This question was arranged so that each 
respondent could check hours, 1-9 hours, 10-20 hours, or 20+ hours. 

Median semester hours credit of earth science teachers 

Earth Science 




















The courses which the majority of teachers thought would be most 
beneficial to their teaching were: Oceanography (76%), Astronomy 
(61%), Geology (60%), and Meteorology (38%). Judging from the 
median semester hours of the teachers' background, it is obvious that 
they recognize their definciencies. What is surprising though is that so 
few of them felt a course in Meteorology would be beneficial since more 
than 50% had never had such a course. 

Approximately 10% of the teachers listed earth science or geology 
as their major for the bachelor's degree, while approximately 40% listed 
biology as their undergraduate major. The remaining teachers mainly 
listed various fields of the natural or biological sciences as their 
undergraduate major. 

Those teachers with a master's degree mainly listed the following 
majors: education (35%), earth science (25%), and biology (15%). 
Their graduate degree minors were mainly education (40%) and biology 

332 Indiana Academy of Science 


The typical earth science teacher in Indiana has the following char- 
acteristics. He — the large majority are men — has a master's degree in 
education or earth science and a bachelor's degree in biology. He has 
more than twenty hours of course work beyond the master's degree, 
which means that he has probably spent a number of his summers in 
school since he has only been teaching for eight years. 

He has taught earth science for the last two years, so he must have 
taught another science, probably biology, general science, or possibly 
chemistry during most of his career. 

There is a 50% likelihood that he initiated the earth science course 
at the school in which he now teaches. The school is located in an urban 
area and his students are in the ninth or tenth grade. His pupils are in 
the upper half of their class and a substantial number will continue their 
education after high school graduation. 

There is a 50% likelihood that he attended at least one N.S.F. Sum- 
mer Institute in earth science. If he did attend such an institute he feels 
this was his best training for instructing his earth science classes. 

He wishes to expand the earth science course at his school with the 
addition of more laboratory equipment. He would also like to take 
additional courses in oceanography, astronomy, and geology in order 
to improve his teaching ability. 

This study illustrates that earth science teachers in Indiana generally 
did not major in earth science at college. However, there is a sizeable 
minority, nearly 20%, who do have a master's degree in earth science. 
Those teachers whose major field was other than earth science have been 
greatly helped by having attended an N.S.F. Summer Institute in earth 

Earth science teaching in Indiana has really come about within the 
last ten years and as the number of pupils continue to expand, there is 
little doubt that their instructors will become better qualified to teach 
the subject. 

Stratigraphy and Correlation of Middle Devonian Strata in the 
Logansport Sag, North-central Indiana 

R. William Orr, Ball State University 


The name Traverse Formation of northern Indiana and the Michigan Basin 
is extended into the Logansport Sag to include approximately 25 feet of strata 
previously separated into Miami Bend Formation, Logansport Limsetone, and 
Little Rock Creek Limestone. Traverse rocks of the Sag and those of the Michi- 
gan Basin of equivalent late Middle Devonian age display similar lithologies, 
have similar depositional histories, and contain nearly identical conodont 
faunas. The Traverse rocks of the Sag lie within two Middle Devonian 
conodont zones high in the Givetian Stage and correlate with strata in 
New York ranging from the Levanna Shale Member of the Skaneateles 
Shale (Hamilton Group) through the Tully Formation. The lower zone 
comprises the body of strata containing Icriodus latericrescens latericrescens 
below the lowest position of Polygnathus varcus; the upper zone is the 
P. varcus Zone. 


Upper Silurian and Devonian rocks are exposed in the valley of the 
Wabash River between Delphi, Carroll County, and Lewisburg, Cass 
County, Indiana (Fig. 1). The outcrop area is situated near the crest of 
the northwest extension of the Cincinnati Arch in the structural depres- 
sion that Cumings and Shrock (10) named the Logansport Sag. 

Approximately 25 feet of Middle Devonian carbonate rocks lie 
between Silurian strata and the New Albany Shale (Upper Devonian). 
Although the Middle Devonian part of the section is quite thin and 
exposures are relatively few, several units assigned the rank of forma- 
tion have been proposed for rocks that crop out in the Sag. These are 
Miami Bend Formation (6), Logansport Limestone (7), and Little Rock 
Creek Limestone (5). These three units nowhere are known to be present 
in a single outcrop, and their lateral and vertical relationships as a 
result are unclear. With few exceptions, among them the work of Cooper 
and associates and that of Campbell (3), these names rarely have been 
used. They have not been adopted for use by the Indiana Geological 

Middle Devonian rocks of the Logansport Sag lithologically more 
closely resemble equivalent rocks of the Michigan Basin than those of the 
Illinois Basin. As pointed out by Pinsak and Shaver (15), rocks equiva- 
lent to part of the Traverse Group of Michigan are recognized as far 
south as Carroll, Cass, Howard, and Miami Counties, Indiana. During 
Middle Devonian time basin type sedimentation was taking place on 
the northeast side of the Cincinnati Arch and a southward-thinning wedge 
of sediments was deposited in northern Indiana (15). The Miami Bend, 
Logansport, and Little Rock Creek represent the southern extension of 
this wedge into the Logansport Sag. 

Field work was conducted during the summer of 1967 when the 
writer was associated with the Indiana Geological Survey. Charles 


Indiana Academy of Science 

R R L L 

Figure 1. Map of Carroll and Cass Counties showing collecting localities of 
Middle Devonian limestone units in the Logansport Sag. 

A. Pollock, Department of Geology, Indiana University, accompanied the 
writer in the field and assisted in the collecting of samples. Charles 
Pollock, Carl B. Rexroad and William J. Wayne (both of the Indiana 
Geological Survey), and Robert S. Nicoll, Department of Geology, The 
University of Iowa, discussed the stratigraphy of the area on several 
occasions. Carl Rexroad and Robert Nicoll critically read the manuscript. 

Development of Stratigraphic Nomenclature 

Middle Devonian rocks of the Logansport area have been studied by 
numerous geologists for nearly 100 years. Collett (4) made a brief 
survey of Silurian and Devonian rocks in Carroll and Cass Counties 
among others in 1872. He recognized two divisions of the Devonian part 
of the section, a lower limestone unit (unnamed by Collett) 20 to 22 feet 
thick and an upper black shale unit 60 feet thick (Fig. 2). Collett used 
the name "Louisville-Delphi black slate" for the latter unit in two col- 

Geology and Geography 





















































Figure 2. Chart showing development of nomenclature applicable to Middle 
Devonian rocks in the Logansport Sag. 

umnar sections. As pointed out by Wilmarth (23), the "Louisville-Delphi 
black slate" probably was named for its surface distribution from Louis- 
ville, Kentucky, to Delphi, Carroll County, Indiana. Brown (2) used the 
name "Delphi black slate" to designate the same unit in Marion County, 
Indiana. Both names have since been replaced in common usage by the 
older and more widely accepted name New Albany Shale (1). 

Kindle (13) distinguished two divisions of Devonian carbonate rocks 
in the Wabash Valley near Logansport, a lower, gray, crystalline, thin- 
to thick-bedded limestone and an upper limestone that varies from bluish 
drab and conchoidally fracturing to dark colored and arenaceous. He 
correlated the lower unit with the Jeffersonville Limestone (an untenable 
correlation on the basis of conodont succession) of southern Indiana and 
applied the southern Indiana term Sellersburg beds (12) to the upper 
limestone and noted its Hamilton fauna. He provisionally assigned to the 
Silurian the stromatoporoid-bearing limestone that he observed to be 
10 to 13 feet thick below the lower unit (Fig. 2). 

Cooper and Warthin (7) proposed the name Logansport Limestone 
for "light-colored granular limestone" in which "corals and other fossils 
are abundant particularly in the upper 6 feet" for Middle Devonian lime- 
stone that rests on the Silurian at many places in the Logansport area. 
They designated as type section the exposure at Pipe Creek Falls (Local- 
ity 7) where Middle Devonian rocks consist of a lower, gray, fine-grained, 
stromatoporoid-bearing unit (assigned to the Silurian by Kindle but 

336 Indiana Academy of Science 

more properly Middle Devonian) and an upper, pink, granular, abundantly 
fossiliferous unit (= the lower of Kindle's two Devonian units). 

Cooper (5) introduced the name Little Rock Creek Limestone (= the 
upper of Kindle's two Devonian units) for 7 feet of "gray, brittle, con- 
choidally fracturing limestone above the Logansport" that is exposed 
along the bed and banks of Little Rock Creek upstream from the bridge 
located 1 mile southeast of Lockport, Carroll County (Locality 2). 

Cooper and Phelan (6) proposed the name Miami Bend Formation 
for the gray, stromatoporoid-bearing limestone that formerly was 
included as the lower part of the Logansport Limestone (7). They desig- 
nated the west side of the France Stone Company quarry (Locality 5) as 
type section. Here the unit is a biostrome about 15 feet thick and 
consists of gray, fine-grained, indistinctly bedded, stromatoporoid- and 
coral-bearing limestone that disconformably overlies the Kokomo Lime- 
stone Member of the Salina Formation (Silurian). 

All three Middle Devonian limestone units may be observed resting 
disconformably on Silurian strata along the Wabash River. The Ken- 
neth Limestone Member (probably Silurian in age) of the Salina 
Formation is not present at any of the studied localities. 

Conodont Faunas 

The Miami Bend, Logansport, and Little Rock Creek units all con- 
tain characteristic upper Middle Devonian (Givetian) conodont species. 
The Miami Bend (sampled at Localities 5 to 7) is a stromatoporoid-coral 
biostrome that yields small numbers of conodonts. Platform species 
include only Icriodus latericrescens latericrescens Branson and Mehl and 
/. expansus Branson and Mehl. At these three localities the Miami Bend 
is overlain along a sharp contact by the granular Logansport Limestone, 
the lower part of which contains a conodont fauna identical to that of the 
Miami Bend. At Locality 6, the upper part of the Logansport contains 
Polygnathus varcus Stauffer in addition to the above two taxa. At Local- 
ities 2 to 4-, I. latericrescens latericrescens, I. expansus, I. cymbiformis 
Branson and Mehl, P. varcus, and P. linguiformis linguiformis Hinde are 
irregularly distributed throughout the Logansport. The conodont fauna 
of the Little Rock Creek (sampled at Localities 1 and 2) is identical to 
that of the Logansport at Localities 2 to 4- listed above. 


The above faunas show that two conodont zones are present in 
Middle Devonian strata of the Logansport Sag. The lower is character- 
ized by the presence of Icriodus latericrescens latericrescens below the 
lowest position of Polygnathus varcus. The upper is the P. varcus Zone 
(25) that is characterized by the association of these two taxa. 

In Germany, Icriodus latericrescens latericrescens is confined to the 
Polygnathus varcus Zone high in the Givetian Stage (24). In North 
America, /. latericrescens latericrescens is widely distributed below the 

Geology and Geography 337 

P. varcus Zone and in New York ranges from the Levanna Shale Member 
of the Skaneateles Shale (Hamilton Group) into the Tully Formation. 
Polygnathus varcus ranges from the Centerfield Limestone Member of 
the Ludlowville Shale (Hamilton Group) also into the Tully (14). See 
Orr and Klapper (18) for a discussion of the correlation of this interval. 

In the Logansport Sag, the Miami Bend and the lower part of the 
Logansport lie within the zone of Icriodus later icrescens later icrescens 
below the lowest position of Polygnathus varcus. In the Michigan Basin, 
the lower part of the Traverse Formation (20) of northern Indiana and 
that part of the Traverse Group of Michigan below and including the 
lower part of the Alpena Limestone lie within this conodont zone (17). 
The highest part of the Silver Creek Member of the North Vernon 
Limestone of southern Indiana also lies within this zone (19). 

The upper part of the Logansport and the Little Rock Creek lie 

within the Polygnathus varcus Zone. In the Michigan Basin, the upper 
part of the Traverse Formation of northern Indiana and that part of the 
Traverse Group of Michigan above and including the middle part of the 
Alpena Limestone to the base of the Squaw Bay Limestone lie within this 
conodont zone (17). In southern Indiana, the Beechwood Member of the 
North Vernon Limestone lies within the P. varcus Zone (16). 

Cooper and others (8) correlated the Logansport with the Beechwood 
Member of southern Indiana and with the Four Mile Dam Limestone of 
Michigan. They considered the Little Rock Creek to be younger than the 
Beechwood and correlated it with the Potter Farm Formation and 
Thunder Bay Limestone (both of the Traverse Group) of Michigan. 
Campbell (3) also correlated the Devonian rocks exposed at Pipe Creek 
Falls with the Beechwood on the basis of megafaunal similarities. 

Galloway and St. Jean (11) studied the stromatoporoid faunas of 
both the Logansport and Little Rock Creek units. They indicated the 
Hamilton age of the Logansport fauna and pointed out the correlation 
with Frasnian and Faminian faunas from the Dinant Basin, Belgium. 
They considered the Little Rock Creek to be of Tully age and correlated 
its stromatoporoid fauna with that of the Potter Farm Formation of 

In 1966 Cooper and Phelan (6) reported for the first time the 
brachiopod String ocephalus in Indiana. They obtained their specimens 
from the Miami Bend unit at several localities just east of Logansport 
including the upper 18 inches of the unit in the west wall of the France 
Stone Company quarry (Locality 5). On the basis of brachiopod faunas, 
Cooper and Phelan correlated the Miami Bend with the Rogers City 
Limestone of the Michigan Basin and with an interval in New York 
between the Marcellus and Skaneateles Shales. In terms of conodont 
zones, the Miami Bend lies within the same zone as the Skaneateles 
Shale of New York. 

Cooper and Phelan's (6) correlation of the Miami Bend with the 
Beauvais Sandstone of Ste. Genevieve County, Missouri, is herein rejected 
as untenable because of the presence in the Beauvais of Icriodus angus- 

338 Indiana Academy of Science 

tus Stewart and Sweet. This important species is found also in the 
Dundee Limestone of northwest Ohio (14), Delaware Limestone of cen- 
tral Ohio (21), and lower part of the intertonguing Speed and Silver 
Creek Members of the North Vernon Limestone of southern Indiana (16). 
The correlation of these units with the Marcellus Shale of New York is 
well established. 

Nomenclatural Proposal 

Thornbury and Deane (22) and Pinsak and Shaver (15) previously 
have pointed out the lithologic affinities of the Middle Devonian rocks of 
the Logansport Sag to Traverse rocks of the Michigan Basin. The 
Miami Bend, Logansport, and Little Rock Creek units represent a south- 
ern extension of southward-thinning Traverse strata into the Sag. The 
Four Mile Dam Limestone of Michigan and the Miami Bend unit repre- 
sent similar biostromal developments of similar Late Givetian age. 
The conodont faunas of the Middle Devonian rocks of the Sag are essen- 
tially identical to those of Traverse strata of the Michigan Basin. 

Because of lithologic similarities, similar depositional histories, and 
equivalent conodont faunas, it is here proposed that the name Traverse 
Formation (20) be used for those rocks in the Sag that previously have 
been designated as Miami Bend Formation, Logansport Limestone, and 
Little Rock Creek Limestone. The extension of the name Traverse into 
the Logansport Sag is consistent with Middle Devonian paleogeographic 
interpretations of Pinsak and Shaver (15) and eliminates needless nomen- 
clature applicable to outcrops in a few-county area. In contrast to the 
Miami Bend, Logansport, and Little Rock Creek, the Traverse is a map- 
pable unit that can be traced in the subsurface of northern Indiana and 
the Michigan Basin. 

Because the Miami Bend, Logansport, and Little Rock Creek units 
are quite thin, have unclear lateral relationships, and have been recog- 
nized only in outcrop in a few-county area, but do possess identifying 
lithologic characteristics, it is here suggested that when these names are 
used they be assigned the rank of lithofacies rather than formation. 

Collecting Localities 

All cited topographic maps are from the U. S. Geological Survey 
7.5 minute series (1:24000). 

Locality 1 : Delphi Limestone Company quarry on north side of 
U. S. Highway 421 northwest of Delphi, SW% SW% sec. 19, T. 25 N., 
R. 2 W., Carroll County, Indiana (Delphi quadrangle). Location cited by 
Cumings and Shrock (9). 

Fauna from Little Rock Creek Lithofacies (7 feet 9 inches to 9 feet 
2 inches below top) contains Icriodus latericrescens latericrescens, 
Polygnathus varcus, and P. linguiformis linguiformis. Fauna from 
Little Rock Creek (4 feet 6 inches to 6 feet 6 inches below top) contains 
the same fauna but without /. latericrescens latericrescens. Fauna from 
Little Rock Creek (3 feet 2 inches to 4 feet 6 inches below top) contains 

Geology and Geography 339 

only P. linguiformis linguiformis. Total thickness of Little Rock Creek is 
9 feet 2 inches. The limestone rests on the Huntington Lithofacies of the 
Wabash Formation (Silurian) and is unconformably overlain by the New 
Albany Shale (Upper Devonian). A conodont fauna recovered from a 
limestone band low in the shale contains Palmatolepis gigas Miller and 
Youngquist and P. suhrecta Miller and Youngquist. 

Locality 2: Little Rock Creek, bed and banks of stream from the 
bridge near its mouth to about 300 yards upstream, SE^i SW 1 ^ sec. 17, 
T. 26 N., R. 1 W., Carroll County, Indiana (Burrows quadrangle). This is 
the type locality of the Little Rock Creek Lithofacies as designated by 
Cooper (5); location cited by Cooper and others (8). 

Fauna from Logansport Lithofacies includes Icriodus latericrescens 
later icrescens, I. expansus, Polygnathus varcus, and P. linguiformis 
linguiformis. Fauna from Little Rock Creek Lithofacies includes 1. 
latericrescens latericrescens, I. expansus, I. cymbiformis, and P. varcus. 
Approximately 12 to 15 feet of strata are poorly exposed at intervals in 
the bed and along the banks of the stream. Silurian rocks crop out in 
the creek bed below the bridge, but the contact with Devonian strata is 
not exposed. 

Locality 3: Low bluff with small natural bridge on north side of 
South River Road along south side of Wabash River across from George- 
town, SW*4 SW 1 ^ sec. 35, T. 27 N., R. 1 W., Cass County, Indiana (Bur- 
rows quadrangle). 

Fauna from Logansport Lithofacies (0 to 1 foot 7 inches above base) 
contains Icriodus latericrescens latericrescens. Fauna from Logansport 
(4 feet 3 inches to 6 feet 6 inches above base) includes I. latericrescens 
latericrescens, Polygnathus varcus, and P. linguiformis linguiformis. 
Exposed thickness of Logansport is 6 feet 6 inches of granular fossilifer- 
ous limestone that overlies the Kokomo Limestone Member of the Salina 
Formation (Silurian). 

Locality 4: West bluff of Grant's Run on east side of South River 
Road 1 mile east of Georgetown, NE 1 ^ SE^ sec. 35, T. 27 N., R. 1 W., 
Cass County, Indiana (Burrows quadrangle). 

Fauna from Logansport Lithofacies (4 feet 9 inches to 11 feet 3 
inches below top) includes Icriodus latericrescens latericrescens, I. 
expansus, and Polygnathus linguiformis linguiformis. Fauna from 
Logansport (0 to 4 feet 9 inches below top) includes these species but in 
addition P. varcus and I. cymbiformis. Exposed thickness of Logansport 
is 11 feet 3 inches. A covered interval of approximately 20 feet separates 
the Logansport from the highest exposure of Silurian strata situated in 
the stream bed at the base of the bluff. 

Locality 5: West wall of France Stone Company quarry on north 
side of U. S. Highway 24, 2 miles east of Logansport city limits, SWM 
NE% sec. 27, T. 27 N., R. 2 E., Cass County, Indiana (Logansport 
quadrangle). This is the type section of the Miami Bend Lithofacies as 
designated by Cooper and Phelan (G). Cooper and Phelan recovered 

340 Indiana Academy of Science 

specimens of String ocephalus from the upper 18 inches of the Miami 
Bend exposed in the abandoned west wall of the quarry (U. S. National 
Museum Locality 391a). 

Fauna from Miami Bend Lithofacies (0 to 1 foot 4 inches, 1 foot 4 
inches to 3 feet, 3 feet to 5 feet above base) contains Icriodus expansus. 
Fauna from Miami Bend (5 feet to 7 feet 6 inches above base, 12 feet 
8 inches to 14 feet 8 inches above base) contains /. latericrescens lateri- 
crescens. Fauna from Miami Bend (7 feet 6 inches to 10 feet 2 inches, 
10 feet 2 inches to 12 feet 8 inches above base) contains both of these 
species. Total thickness of Miami Bend is 14 feet 8 inches. It rests on 
the Kokomo Limestone Member of the Salina Formation (Silurian) and is 
overlain by isolated patches of the Logansport Lithofacies. 

Locality 6: South wall of France Stone Company quarry near gravel 
pile, SW 1 ^ NW& sec. 26, T. 27 N., R. 2 E., Cass County, Indiana (Logans- 
port quadrangle). Location given by Cooper and Phelan (6). 

Fauna from Logansport Lithofacies (0 to 1 foot 10 inches above 
base) includes Icriodus latericrescens latericrescens and /. expans%is. 
Fauna from Logansport (12 feet 7 inches to 14 feet 3 inches above base) 
contains same association but in addition Polygnathus varcus. Exposed 
thickness of Logansport is 14 feet 3 inches. The underlying Miami Bend 
Lithofacies is 6 feet 2 inches thick and rests on the Kokomo Limestone 
Member of the Salina Formation (Silurian). 

Locality 7: Pipe Creek Falls, east bluff of Pipe Creek below the 
dam, 4000 feet north and 1200 feet east of southwest corner of grant 1, 
T. 26 N., R. 3 E., Cass County, Indiana (Onward quadrangle). This is 
the type section of the Logansport Lithofacies as designated by Coopei 
and Warthin (7); section described by Cumings and Shrock (9); location 
cited by Cooper and Warthin (7), Campbell (3), and Cooper and Phelan 

Fauna from Miami Bend (0 to 2 feet 6 inches below base of over- 
lying Logansport) contains Icriodus expansus. Fauna from Logansport 
(0 to 3 feet above base) contains /. expansus. The Miami Bend is 6 feet 
6 inches thick and rests on the Kokomo Limestone Member of the 
Salina Formation (Silurian). Exposed thickness of Logansport is 7 
feet 10 inches. 

Literature Cited 

1. Borden, W. W. 1874. Report of a geological survey of Clark and Floyd 
Counties, Indiana* Indiana Geol. Survey Ann. Rept. 5:133-189. 

2. Brown, R. T. 1883. Report of a geological and topographical survey of 
Marion County, Indiana. Indiana Dept. Geology and Nat. History Ann. 
Rept. 12 :79-99. 

3. Campbell, Guy. 1942. Middle Devonian stratigraphy of Indiana. Geol. Soc. 
Araer. Bull. 53:1055-1071. 

4. Collett, John. 1872. Geological reconnaissance of Jasper, White, Car- 
roll, Cass, Miami, Wabash, and Howard Counties. Indiana Geol. Survey 
Ann. Repts. 3 and 4:289-337. 

Geology and Geography 341 

5. Cooper, G. A. 1941. New Devonian stratigraphic units. J. Washington 
Acad. Sci. 31:179-181. 

6. Cooper, G. A. and Thomas Phelan. 1966. String ocephalus in the Devonian 
of Indiana. Smithsonian Misc. Coll 151(1). 20 p. 

7. Cooper, G. A. and A. S. Warthin. 1941. New Middle Devonian strati- 
graphic names. J. Washington Acad. Sci. 31:259-260. 

8. Cooper, G. A. and others. 1942. Correlation of the Devonian sedimentary- 
formations of North America. Geol. Soc. America Bull. 53:1729-1794. 

9. Cumings, E. R. and R. R. Shrock. 1928a. The geology of the Silurian 
rocks of northern Indiana. Indiana Dept. Conservation Pub. 75. 226 p. 

10. Cumings, E. R. and R. R. Shrock. 1928b. Niagaran coral reefs of Indi- 
ana and adjacent states and their stratigraphic relations. Geol. Soc. 
Amer. Bull. 39:579-620. 

11. Galloway, J. J. and Joseph St. Jean, Jr. 1955. Middle Devonian Stroma- 
toporoidea from Indiana (abs.). Geol. Soc. America Bull. 60:1562-1563. 

12. Kindle, E. M. 1899. The Devonian and Lower Carboniferous faunas of 
southern Indiana and central Kentucky. Bull. Amer. Paleontology 
3(12). Ill p. 

13. Kindle, E. M. 1901. The Devonian fossils and stratigraphy of Indiana. 
Indiana Dept. Geology Nat. Resources Ann. Rept. 25:529-758, 773-775. 

14. Klapper, Gilbert and Willi Ziegler. 1967. Evolutionary development of 
the Icriodus latericrescens group (Conodonta) in the Devonian of 
Europe and North America. Palaeontographica no. 127:68-83. 

15. Pinsak, A. P. and R. H. Shaver. 1964. The Silurian formations of northern 
Indiana. Indiana Geol. Survey Bull. 32. 87 p. 

16. Orr, R. W. 1964. Biostratigraphic zonation and correlations based on 
conodonts of Middle Devonian strata of southern Illinois and adjacent 
states. Unpublished M. A. thesis, The University of Texas, Austin. 

17. Orr, R. W. 1967. Conodonts from Middle Devonian strata of the Michi- 
gan Basin. Unpublished Ph.D. thesis, Indiana University, Bloomington. 

18. Orr, R. W. and Gilbert Klapper. 1968. Two new conodont species from 
Middle-Upper Devonian boundary beds of Indiana and New York. J. 
Paleontology 42:1066-1075. 

19. Orr, R. W. and C. A. Pollock. 1968. Reference sections and correlation 
of Beechwood Member (North Vernon Limestone, Middle Devon- 
ian) of southern Indiana and northern Kentucky. Amer. Assoc. Petrol- 
eum Geologists Bull. 52:2257-2262. 

20. Schneider, A. F. and S. J. Keller. In preparation. Geologic Map of the 
Chicago 1° X 2° Quadrangle, Indiana, Illinois, and Michigan, showing 
bedrock and unconsolidated deposits. Indiana Geol. Survey Regional 
Geol. Map 4. 

21. Stewart, G. A. and W. C. Sweet. 1956. Conodonts from the Middle De- 
vonian bone beds of central and west-central Ohio. J. Paleontology 

22. Thornbury, W. D. and H. L. Deane. The geology of Miami County, 
Indiana. Indiana Geol. Survey Bull. 8. 49 p. 

23. Wilmarth, M. G. 1938. Lexicon of geologic names of the United States 
(including Alaska). U. S. Geol. Survey Bull. 896. 396 p. 

24. Wittekindt, Hanspeter. 1966. Zur Conodontenchronologie des Mittelde- 
vons. Fortschr. Geol. Rheinland u. Westfalen no. 9:621-646. 

25. Ziegler, Willi. 1962. Taxionomie und Phylogenie Oberdevonischer Cono- 
donten und ihre stratigraphische Bedeutung. Hess. Landesamt. Bodenf. Abh. 
no. 38. 166 p. 

Planning for and Utilization of the Web Pattern of 
Physical Urban Development in Cities 

Thomas Frank Barton, Indiana University 

In two earlier papers, (1, 2) the writer identified and described a 
web pattern of physically urban development consisting of: 1. business- 
industrial-circulation corridors with residential meshes located in urban 
areas and 2. a web of corridors of urban development in the countryside 
enclosing- farm and forest lands. 

This paper recommends that rather than permitting a web of busi- 
ness-industrial-circulation corridors to develop haphazardly, American 
city planning departments should design and zone a web of these corri- 
dors. There are seven primary advantages in taking this progressive 
and decisive planning and zoning action: 

1. Aids industry by (a) providing for wider and more even distribution 
of business and industry throughout the urban area and thereby 
helping rlatively to relieve congestion in the downtown; (b) permit- 
ting concentration, yet dispersal, of industry; (c) increasing the 
opportunity for the number of and greatly increasing the foot- 
frontage for strategic industrial sites fronting on (or nearby) major 
transportation lines and primary utility lines; and (d) making possi- 
ble more efficient location of industrial districts. 

2. Provides for nodal business development. 

3. Permits better location of trucking terminals and warehouses. 

4. Makes possible better siting of high rise apartments. 

5. Enables utilities to provide more efficient, economical service. 

6. Helps concentrate populations and travel destinations thereby sup- 
porting existing or potential mass transit. 

7. Protects and helps make more viable and liveable neighborhood and 
community residential areas. 

Aid Industry 

Better city-wide distribution. Today in many American cities too 
much industry is concentrated near the business-industrial core of the 
city or has migrated to peripheral and /or suburb locations. If too high 
a fraction of the city's employment opportunities is concentrated in the 
downtown, traffic congestions at morning rush hours will be caused 
primarily by too many people converging from various directions of the 
compass to a relatively smaller and smaller area. In contrast if relatively 
too many businesses and industries are located on the periphery of a 
large city in political suburbs, the lower income bracket group of em- 
ployees finds it impossible, difficult or expensive to make their journeys 
to work. Moreover the central city is hard pressed to maintain a suffi- 
ciently high tax base to support city services. The zoning of a web of 


Geology and Geography 343 

business-industrial-circulation corridors not only encourages the city- 
wide distribution of business and industry instead of having them con- 
centrated in large islands or on one or more edges of the downtown but 
such zoning will also reduce if not prevent future scatteration of these 
establishments on city peripheries where they may be subsequently en- 
gulfed by residential subdivisions. In contrast, industries and businesses 
built in these corridors are protected from encroachment and enclosure 
by residential sub-division promoters. Owners of industries now scattered 
throughout the residential areas should be instructed not to expand or 
make improvements in their present plants. Moreover any industries 
wishing to locate, expand or relocate should be helped to secure new 
sites in the corridors. Present owners of residences in the newly desig- 
nated corridors should be advised not to improve their property and 
to be encouraged to find new locations. Within a few decades if not 
earlier the scatteration of industries in residential areas and of resi- 
dences in the business-industrial corridors could be materially reduced 
and primarily eliminated. 

Permits concentration yet dispersal. The web pattern with its corri- 
dors will provide an alternate choice to the management of new or ex- 
panding businesses and industries that do not wish to choose between 
two extremes. These extremes are: 1. staying in the downtown area 
or 2. locating on the edge of the city. The web pattern of corridors pro- 
vides a large number of locational opportunities between the center of 
the city and its edges and in relation to other major considerations such 
as an airport or a university. 

When business and industry are widely dispersed over the city in 
corridors, there is a greater opportunity for more people to live in the 
residential meshes of the web pattern and be closer to their places of 
employment. In this way those employed in a nearby corridor may save 
hours each day and have transportation costs reduced in their journeys 
to work and shop and to secure services and recreation. This will also 
relieve at least relatively downtown congestion. 

Increases strategic industrial sites. Efficient, quick, low cost trans- 
portation is vital to most businesses and industries. The amount of land 
adjacent to nodal areas, produced by the junctions and crossings of 
several types of transportation such as highways, railways, water- 
ways and airways is limited. However in business-industrial corridors 
where railroad lines, limited access expressways and subways are built 
roughly parallel to each other for miles and miles, the land fronting 
on these lines (or located within a block or two) is increased enormously. 
At the present time in many cities railroad lines and limited-access ex- 
pressways parallel to each other or perhaps one or two miles apart 
extend for long distances through residential areas where their presence 
is considered a nuisance if not an economic and social liability. With the 
development of corridors: 1. transportation line right-of-ways are re- 
stricted to compact areas, 2. the transportation facilities can be co- 
ordinated to give more efficient service, 3. frontage sites for industry 
and to a lesser extent for business are increased by linear development 

344 Indiana Academy of Science 

and 4. single-dwelling residential areas may be better shielded with 
parkways, high-rise apartments and semi-public buildings. In this manner 
residential land adjacent to but not fronting on transportation lines has 
its value increased rather than decreased. 

Of course each corridor will not have a railroad, subway and ele- 
vated line but each corridor will be served by a major limited access ex- 
pressway consisting of 4 to 12 lanes or more. 

Efficient location of industrial districts. The zoning of business- 
industrial corridors and the provision of adequate transportation and 
utility facilities reduces the competition, pressure and scramble for 
industrial sites and encourages the sorting of industries into roughly 
homogeneous groups. Some industries have clean facilities, use clean raw 
materials and are on well-landscaped grounds. These industries desire 
prestigious environments and wish to be in a cluster of similar indus- 
tries rather than adjacent to plants with extensive truck activity and 
open-lot storage of raw materials which at best give a cluttered, un- 
sightly appearance. In the corridors tertiary industries using raw ma- 
terials delivered to the plant by truck and airplane would be located in 
one section and secondary industries using railroad cars and ship or 
barge loads of coal, ores, lumber and other materials could more con- 
genially and efficiently occupy a different one. The former could be 
adjacent to an airport and the latter an ocean, lake and/or river port. 

Encourages Nodal Business Development 

Although nearly all large cities with populations of about 250 thou- 
sand or more (as well as many cities of smaller size) have extensive com- 
mercial strips or ribbon development occupying net patterns, most of 
the city planning departments are attempting to discourage and/or block 
this development. Long-range plans shown on land use maps indicating 
future city goals show a hierarchy of business nodal areas ranging in 
size from the neighborhood through the community to the regional 
shopping and service areas with the apex in the downtown. 

If land were zoned and developed as business-industrial corridors, 
business could be more easily and justifiably restricted to corridor junc- 
tions with the regional centers at the major junctions and the community 
centers at the secondary junctions. The same transportation and utility 
systems would serve both urban land-use functions — business and in- 
dustry. Ribbon development should be reduced and discouraged and in 
time primarily eliminated. Zoning would prevent industry from occupying 
nodal corridor junctions which provide strategic sites for retail, service 
and recreational establishments. Moreover zoning would prevent retail 
establishments from occupying scattered sites that become dispersed in 
a nuisance fashion among the industries. 

Better Location of Truck Terminals and Warehouses 

Without business-industrial corridors strategic sites for trucking 
terminals and warehouses wax and wane. Moreover in some cities the 

Geology and Geography 345 

zoning departments are attempting to concentrate these facilities in 
one large area on one side of the downtown. However different types of 
trucking terminal and warehouses should be grouped in districts adjacent 
to the types of businesses and industries they service. Warehouses could 
be located in the corridors fronting on or within a few blocks of limited 
access expressways, railroad lines and water transport or airfields. 

Siting High-Rise Apartment Districts 

With the businesses, industries, and transportation concentrated in 
corridors which enclose the residential meshes, at least two general types 
of high rise apartment sites become available. These may be found adja- 
cent to and flanking the corridors or near the center of the communities 
enclosed in the meshes. The writer would like to see many of the high 
rise apartments occupying an open-space greenbelt of parkways which 
would help to screen the corridors. Some of the more expensive high- 
rise apartments could overlook the better landscaped industrial districts. 
Apartments for the less affluent and perhaps hand and semi-skilled labor- 
ers could be located adjacent to the secondary industries where their 
employment is located. No attempt would be made to segregate income 
groups. The primary criterion would be to locate types of apartment 
house suitable for the non-management employees as near potential 
places of employment as possible so as to reduce the time spent in 
going to and from work. 

Bicycling and walking to work which is so commonplace in Europe 
would then become potentially possible for many here in the United 
States. If this type of travel were adopted, automobile traffic could be 
relatively reduced and perhaps the worker's health improved. 

More Efficient Utility Service 

If retail, wholesale, service and industrial establishments were lo- 
cated in corridors with high-rise apartments flanking them, utility man- 
agements might become believers in the philosophy that a "little bit of 
heaven or paradise" can exist on earth. These corridors then would be- 
come the sites of the primary arterials of water, sewage, gas, electricity, 
telephone and other services. Tributary lines would connect the single- 
house, duplex and townhouse apartment residential areas in the meshes 
with the primary lines in the corridors. This would relatively reduce both 
construction and maintenance costs and provide better service. For 
example, better pressures could be maintained in water systems and 
periods of interrupted service reduced. The present-day maze of utility 
requirements found in many cities hinders efficient management. Busi- 
ness-industrial corridor development would help bring order out of chaos. 

Mass Transit 

Billions of words have been written to support the theme that gaso- 
line-engine traffic is choking and doing almost irreparable damage to not 
only the downtowns but entire large cities. And many urban leaders ask 
why mass transit is not economical and feasible? Still others in thought- 

346 Indiana Academy of Science 

less exasperation recommend that mass transit be subsidized by govern- 
mental funds. Such people apparently forget that mass transit lines are 
going out of business each decade because most people prefer to use their 
own car rather than mass transit. Such transit is only feasible when 
large numbers of people wish to use such transportation to reach a con- 
centration of mass destinations. It is interesting to note that one of the 
largest cities in North America, where mass transit has been relatively 
successful, has adopted a long-range plan for developing through zoning 
business-industrial corridors. 

Permits Community Development 

By designing the location of the corridors, the size of the resi- 
dential areas enclosed by such may be controlled to the extent that 
each mesh would have large enough space for at least several (four to 
five) neighborhoods and one community. Some residential meshes may 
have several communities. But the areas for these neighborhoods and 
communities should be ample to accommodate single-house, duplex and 
town apartments, as well as neighborhood and community schools, play 
grounds and shopping centers. 

Unplanned yet rapidly developing web patterns of business-indus- 
trial corridors have enclosed or are slicing through residential areas 
dividing and subdividing neighborhoods and communities. These un- 
planned and non-designed corridors are yearly destroying billions of 
dollars in undeterminable residential values and destroying or helping 
to block community development. 


The writer's paper entitled "The Web Hypothesis of Physical Urban 
Growth" (2) was written primarily to call attention to and to identify a 
new hypothesis or generalization concerning the physical pattern of 
city growth, and to show how the urban web pattern with its business- 
industrial threads or corridors encloses, and as growth continues, sub- 
divides residential areas and produces residential islands of various sizes. 
The partially-enclosed residential areas both within and on the periphery 
of the web pattern indicate that additional subdivisions of residential 
areas will occur. 

Although written in November, 1966, that paper contained the fol- 
lowing two paragraphs: 

"Even in its initial step (of formation), this hypothesis may 
aid city planning by: 1. helping to justify the planning and con- 
structing of both city and private services in the new primary arte- 
ries on fringes of the geographic city before the construction of 
stores, offices and factories begins, 2. supplying a defense for the 
zoning of land along the primary arterials for business and industry 
and 3. zoning the location of business-industrial corridors to regu- 
late the size of residential communities so that these may be more 
economical and viable. 

Geology and Geography 347 

In fact, the development of multiple distribution systems — of 
streets and transit lines; water, storm water and gas lines; below- 
surface telephone and electrical wires and other forms of transporta- 
tion, utilities and communication both public and private — may pro- 
vide the means of helping control patterns of urban development." 

And after these preceding two paragraphs were written the writer 
obtained a copy of The Comprehensive Plan of Chicago in which "Corri- 
dors of High Accessibility" with associated "Industrial Areas" are 
shown (3). 

This paper has stressed seven major advantages of zoning a city- 
wide pattern of business-industrial-circulation corridors for: 1. long- 
established cities, 2. potential urban areas on the perpheries of cities, 3. 
megalopolises and 4. potential large new cities (over 100,000 popula- 

Literature Cited 

1. Barton, Thomas Frank. 1967. Notes on a New Pattern and Process of 
Physical City Development: The Web Theory. Proc. Indiana Acad. 
Science 76:339-346. 

. 1967. The Web Hypothesis (Theory) of Physical Urban 

Growth, (abstr.) Annals Assoc. Amer. Geographers 57:781-782. 

Duba, John G. 1966. The Comprehensive Plan of Chicago. Chicago Plan- 
ning Commission, Chicago, Illinois. 

Transportation of Mineral Aggregates in Indiana 1 

Robert R. French, U. S. Bureau of Mines 


Mineral aggregates, such as crushed stone, sand, and gravel, are essen- 
tial to the expansion and renewal of urban areas and to the improvement 
of highway systems. Because of the mineral producer's ability and willing- 
ness to improve production techniques and to absorb increasing costs, the 
price of mineral aggregates has remained remarkably stable. The stability 
of f.o.b. prices, depletion, and the exclusion by zoning of some well-located 
deposits has caused transportation costs to become an increasingly 
important part of the overall price of mineral aggregates in Indiana. 

Published and unpublished rate schedules show that trucking is the 
most economical method of transporting aggregates up to a distance of 
about 35 miles. Between 35 and 230 miles railroad rates appear to be the 
most economical, although substantial variations exist between companies 
and between northern and southern Indiana. Barge transportation is most 
economical for distances of more than 230 miles. Average charges are 
approximately 2.5 to 5 cents per ton mile plus 25 cents per ton base charge 
for road transport, 0.82 cents per ton mile plus $1.40 per ton base charge 
for rail transport, and 0.45 cents per ton mile plus $2.25 per ton base charge 
for water transport. 


Most mineral aggregates, such as crushed rock, sand, and gravel, 
are low-value, high-volume commodities that are essential to the ex- 
pansion and renewal of housing units, transportation systems, and work 
facilities for urban populations. Under normal conditions, the con- 
struction commodities can bear only nominal transportation charges and 
must be produced as near the market area as is economically, socially, 
and geologically feasible. 

Numerous economic and social problems develop and become more 
serious as rapid urbanization increases the demand for construction ma- 
terials. Producing companies that have served a growing market for any 
length of time may ultimately be faced with increasing operating costs 
because of thicker overburden, higher lift or longer quarry haul, higher 
incidence of equipment repair, increased pumping, and anti-pollution re- 
quirements. These problems, however, are mostly overshadowed by the 
proximity of the production site to the market area. The economic ad- 
vantage of good site location can, and commonly does, result in an 
increased number of complaints about vibration, traffic congestion, noise, 
physical danger, and air and water pollution, by the local residents. The 
social incompatibility of producer and consumer creates a tendency for 
new production sites to be selected or relocated in less densely populated 
areas. However, most production sites are selected in areas served by 

1 The data used in this paper were compiled in 19G6-1967 while the 
author was employed by the Indiana Geological Survey, and are used with 
the permission of Dr. John B. Patton, State Geologist, Bloomington, 


Geology and Geography 349 

good highways and if unplanned 'strip' urbanization occurs along these 
transportation routes, then the social conflicts are repeated. 

Geologic and geographic distribution of commercial grade deposits 
is another major factor to be considered in the production and subsequent 
marketing of mineral aggregates in an urban area. Geologic conditions in 
Indiana are such that large areas in the northern, southwestern, and 
southeastern parts of the State are covered with thick glacial drift and 
fine-sized outwash, or are underlain by sandstone, shale, or thin lime- 
stone. Aggregates, especially the coarser sizes (plus l 1 /^ inch), must be 
transported into these areas by road or rail, and the cost is considerable. 
For example, coarse aggregates that are normally priced about $1.50 per 
ton f.o.b., commonly retail for $3.10 per ton in Vincennes, and $2.30 to 
$3.15 per ton at stockpiles in Indianapolis, and for more than $4.00 per 
ton in South Bend. 

The net effect of the economic, social, and geologic factors, and the 
low capital requirements (compared to other mining operations^) of 
mineral aggregate production, has been to create a highly competitive 
industry in Indiana. Average prices of crushed rock, sand, and gravel 
have remained quite low (the average price of sand and gravel was $0.94 
and crushed rock $1.27 per ton in 1966 as compared to $0.72 and $1.20 
per ton, respectively, in 1947), and the number of producing quarries 
has remained reasonably constant (80 to 89) during the past 20 years. 
Company data from the sand and gravel industry are not readily avail- 
able, but an analysis of crushed stone operations in Indiana (2) has 
shown that most of the 35 independent company failures that occurred 
between 1947 and 1965 were in areas where production facilities were 
relatively concentrated (5 to 10 quarries within 20 miles). Most of the 
failures in these highly competitive areas are believed to be the result 
of both the market size and the minimization of the protective effect of 
transportation costs. 

Highway Transport 

During the first quarter of the twentieth century, most mineral ag- 
gregates were shipped by rail or used in the immediate vicinity of the 
quarry or pit. Producing companies began trucking their own aggregate 
about 1925, and later, in the 1940's, started using owner-operated ve- 
hicles. This was a workable arrangement for small single-quarry com- 
panies but mergers, purchases, and normal growth brought larger inte- 
grated companies and a definite trend toward the use of contract haulers. 
Today more than three-fourths of the aggregate produced in Indiana is 
transported to the consumer by trucks, most of which are owned by 
independent trucking companies. 

Contract haulers are required to file their current freight rates 
with the Public Service Commission, State Office Building, Indianapolis. 
Typical published rates for three independent trucking companies are 
shown in Table 1. The wide variations in charges for hauling crushed 
rock, sand, and gravel on a cents per ton-mile basis suggest that per- 
haps some "ghost" rates or other factors may be included in these data. 


Indiana Academy op Science 

Haulage rates are, in fact, commonly bid on a single job basis and de- 
pend largely on the size of the job and traffic and road conditions be- 
tween the production site and the construction project. Some of the 
actual 1967 rates charged to one rural producer of crushed rock are 
shown in Figure 1. The basic charges are approximately 5 cents per ton- 
mile for the first 6 miles, 3 cents per ton-mile for the next 18 miles, and 
2y 2 cents per ton-mile thereafter. A base charge of 25 cents per ton was 
also applicable. These charges are similar, but are not necessarily identi- 
cal to those reported to be in effect in other areas of the country (1, 4, 5, 
7) and Canada (6, 8). 


I | 1 

| i | i | y\ 

' ^ 

y ^s^ 

X ^s^ 









k 100 






5 75 








i 1 i 

1 1 1 1 1 1 



20 30 




Figure 1. Two unpublished tariffs for highway transport of mineral ag- 
gregates in a rural area. 

table 1. Some iMiblished rates for highway trans}Jortation of mineral 
aggregates in Indiana 

Cost per 

Cost per 

Cost per 


ton (cents) 


ton (cents) 


ton (cents) 









































For 30 miles or more, cost is 3.5 cents per ton-mile. 

Geology and Geography 


The average f.o.b. price of crushed rock in Indiana is approximately 
$1.27 per ton, thus a haul of about 31 miles in some rural areas will 
effectively double the price of the aggregate to the consumer. In urban 
and some suburban areas, the transfer distance required to double the 
f.o.b. price may be as short as 9 or 10 miles, depending mostly on time 
and traffic conditions. The cost of sand and gravel, which normally sells 
for less than $1.00 per ton, can be doubled by a transfer distance of only 
6 to 8 miles in some densely populated areas. 

Rail Transport 

Rail transportation is used to haul crushed rock and minor amounts 
of sand and gravel into northern and western Indiana, and into the 
neighboring states. A substantial part of the demand for crushed rock 
in Indianapolis is also satisfied via rail transport of material produced 
in the surrounding counties (3). Freight rates for rail transportation of 
mineral aggregates vary widely and, in effect, between areas of the 
State. Average freight rates discussed here are for point-to-point hauls 
as filed with the Public Service Commission. Loading and unloading of 
freight cars is normally the responsibility of the shipper and the cost 
varies from about 20 to 40 cents per ton, depending upon the type of 
rolling stock and equipment available. 

Variations in published point-to-point rates range from about 10 
cents per ton-mile to less than one cent per ton-mile with minimum 
charges ranging from $0.99 to $1.84 per ton. Reasons for these variations 
may include complex routes requiring one or more transfers, bulk rates 


— Northern Indiana 

— Central Indiana 

— Southern Indiana 


100 150 




Figure 2. Some published point-to-point tariffs for rail transport of 
mineral aggregates. 

352 Indiana Academy of Science 

for high annual tonnages or large shipments, or other market condi- 

The regression lines shown in Figure 2 were derived from estimated 
track mileage and published rates for point-to-point hauls. Most of the 
lower rates were in effect in southern Indiana where the average charge 
was .61 cents per ton-mile plus 99 cents per ton minimum charge. In 
northern Indiana, where quarries are few and moderately long hauls are 
common, the charges for rail transportation were approximately .92 
cents per ton-mile plus $1.84 per ton minimum. The average rate for 
all data considered in Figure 2 is .82 cents per ton-mile plus $1.40 per 
ton minimum. 

Water Transport 

A few of Indiana's crushed rock and sand and gravel operations are 
strategically located along the bluffs and flood-plains of the Ohio River, 
and are able to take advantage of barge transportation for long hauls. 
Company-owned or leased barges of 500 to 1,200 tons capacity are used 
to ship aggregates to major markets in southwestern Indiana, Illinois, 
Kentucky, Ohio, and West Virginia. Contract haulers rates are not 
regulated by the Interstate Commerce Commission, but are commonly 
quoted by the individual companies according to market conditions and 
are subject to change. In late 1966, one barge company established some 
point-to-point transportation rates of .3 to 1.02 cents per ton-mile, f.o.b. 
in barge, with a minimum load of 500 tons per barge. Another company 
quoted an average rate of .81 cents per ton-mile, and a third company 
quoted .4 to .65 cents per ton-mile for minimum barge loads of 1,200 tons 
(plus 50-100 cents per ton for loading and 100-200 cents per ton for un- 
loading.) The data shown in Figure 3 are published point-to-point rates 
from docks in Indiana and Kentucky. Relatively high (compared to the 



Z 6 

« 5 


"• 4 

3 3 


Q 2 

50 100 150 200 250 300 350 


Figure 3. Some published point-to-point tariffs for water (barge) trans- 
port of mineral aggregates. 
















__i— *— -"■ 



• • 

• • 











Geology and Geography 


cost of aggregate) loading and unloading charges normally prohibit the 
use of barge transport for all but the longer hauls, although direct load- 
ing via conveyor belt from processing plant to barge is possible at some 
sites and permits reduced handling charges and greater distance of 
economic haul. The use of flat-deck barges can further reduce the cost 
of loading and unloading to about 25-50 cents and 50-100 cents per ton, 

Summary and Conclusions 

Trucking will probably continue to be the dominant mode of mineral 
aggregates transportation because of the ease of loading and the ability 
to deliver directly to the point of consumption. Factors of production and 
transportation generally restrict deliveries to within about 35 miles of 
the production site, and it is within this radius that trucking appears to 
be the most economic (Figure 4). Geologic conditions do not permit 
large scale open pit production of mineral aggregates in northern and 
southwestern Indiana and it is to these areas, which are mostly more 
than 50 miles from production, that railroads provide the most economic 
means of transportation. Lack of suitable crushed rock aggregate in some 
parts of the neighboring states has allowed some Indiana deposits that 
are strategically located on the bluffs of the Ohio River to be exploited 
and the aggregate shipped by barge to Illinois, Ohio, Kentucky, and West 
Virginia. Water transport appears to be the most economic method for 
distances of more than 230 miles, but direct loading facilities from plant 
to barge may extend the competitive range. 

1 ! ~ 

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Figure 4. Comparison of average tariffs for highway, rail, and water 
transport of mineral aggregates. 

354 Indiana Academy of Science 

Social conditions and the concentration of major markets on the 
fringe of expanding metropolitan areas, in 'satellite' cities, and in high- 
way construction projects, create a tendency for aggregates producers to 
locate or relocate their open pit operations in the more sparsely popu- 
lated areas if geologic conditions and transportation facilities are amen- 
able. Thus, continued urbanization and the depletion of some of the cen- 
trally-located deposits may require longer transportation hauls than 
those previously considered feasible and competitive. The cost of longer 
transportation must ultimately be reflected in higher costs of construc- 
tion, assuming no major changes in technology. Greater truck or rail- 
road car capacity, better highways, unit trains, and some economies of 
scale in larger and more remote production facilities, will counterbalance 
the increased transportation costs to some degree. Crushed rock and 
stone sand from underground mines located within some urban areas 
offer an alternative to increasing transport cost, but the industry has 
not yet developed this possibility to any great extent. Land values, and 
the demand for underground shelters, reservoirs, waste disposal areas, 
or constant temperature storage space may require that we examine the 
open space left within some urban areas for the possibility of obtaining 
mineral aggregates and creating usable underground space. 

Literature Cited 

1. Dunn, J. R. 1966. Mineral resources and minerals conservation. Speech 
prepared at Rensselaer Polytechnic Institute. 

2. French, R. R. 1967. Indiana's crushed stone industry, 1947-1965. Indiana 
Business Review 42 :7-ll. 

3. French, R. R. and D. D. Carr. 1967. Geologic factors affecting the explo- 
ration for mineral aggregates in the Indianapolis area. Purdue Engi- 
neering Bull. 127:86-103. 

4. Goldman, H. B. 1959. Urbanization and the mineral industry. California 
Div. Mines, I. S. 12. 

5. Gray, J. J., N. S. Peterson and G. A. Kingston. 1968. Mineral transporta- 
tion costs in the Pacific Northwest. U. S. Bureau of Mines, I. C. 8381. 

6. Hewitt, D. F. 1962. Urban expansion and the mineral industries in the 
Toronto-Hamilton area. Ontario Dept. Mines, I. M. Rept. 8. 

7. Sheridan, M. J. 1967. Urbanization and its impact on the mineral aggre- 
gate industry in the Denver, Colo. area. U. S. Bureau of Mines, I. C. 

8. Tyler, P. M. 1964. Cost of acquiring and operating mineral properties, 
p. 167-223. In: Economics of the mineral industries. Amer. Inst. Mining 
Metall. Engineers. 

9. Wimpfer, S. P. and N. Severinghaus. 1968. Industrial minerals and rocks, 
p. 849-874. In: Surface mining. Amer. Inst Mining Metall. Engineers. 

An Economic Appraisal of Reclamation Practices on a Strip Coal 
Mine Site in Greene County, Indiana 

R. Michael Dinkel and Lee Guernsey, Indiana State University 


The main purpose of this study was to ascertain the impact of very 
adverse physical conditions on reclamation practices and to determine the 
costs involved in revegetating the toxic strip mine site. A number of 
experiments were carried out in order to obtain a comparison of selected 
factors of reclamation. 

Four expenditures were analyzed in order to determine the total costs 
of reclaiming- land which had developed a very harsh environment from 
strip coal mining- operations. These were the costs of utilities for prepar- 
ing media, the transportation expenses of delivering the media for field 
application, the labor costs of preparing the test sites and the planting of 
trees, and the costs of materials which included the media used, the plants, 
and the seed. 

The total cost of reclaiming an acre of land to trees at the project site 
was 2.4tf per ton of coal mined or about $150 per acre. In contrast, the cost 
of reclaiming the test site to grass averaged 8.4tf per ton of coal mined or 
about $435 per acre. However, if the test site had more favorable 
characteristics, the costs would have been greatly reduced. 

Throughout the Eastern Interior Coal fields of the United States, 
a significant percentage of productive land is undergoing strip coal min- 
ing operations. In this process, massive machinery is used to remove the 
overburden of soil and rocks from above the various seams of coal. The 
condition in which the areas are left after the mineral has been ex- 
tracted is in need of reclamation. 

This study was conducted to ascertain the significance of the physi- 
cal and economic factors involved in the revegetation of a strip mined 
area was internal and runoff flows into strip mined ponds of the area and 
two spoil bank ridges at the Greene County site. One of these ridges 
was composed entirely of sandstone, whereas the second ridge was made 
up primarily of shaly material. 

The slopes of the project area were not too steep for planted vegeta- 
tion, but all the plantings were made upon the struck off surface on top 
of the two spoil bank ridges. The ridge tops had a width of nearly twenty 
feet between the shoulders of the banks. Most of the drainage in the 
area was internal and runoff flows into strip mined ponds of the area and 
remains stagnant after reaching these water bodies. This fact has been 
instrumental in the ponds having a high iron content and a very reddish 

Another physical characteristic of primary importance is the soil 
texture of the project area. The character of materials making up the 
spoil surfaces has a direct effect on plant growth and ultimate soil de- 
velopment. Since the shale spoil bank had a much finer texture than the 
sandstone spoil bank, more compaction occured on this spoil material and 
resulted in an impervious hardpan developing on the shale ridge. 


356 Indiana Academy of Science 

The sandstone ridge in the project area had a pH range of from 
3.0 to 4.9 and was a marginal spoil bank. This is in contrast to the shale 
ridge, which has an average pH of 2.7 and was classified as being toxic. 
The non-toxic portions of the marginal spoils were plantable, but were 
usually small in size and randomly scattered. The pH readings were 
taken at various depths in the spoil banks. This was done since leaching 
processes often cause the surface spoil to be less acidic than that found 
at a depth of a few inches. Qualitative chemical tests were made on the 
spoil from both strip mine ridges within the experimental area. Both 
the shale and the sandstone ridge were found to be very rich in phos- 
phorus. Traces of potassium, magnesium, and calcium were also found 
in the area. Iron was present in samples from both ridges, but was 
more abundant from the sandstone area. Both ridges had a very high 
sulfate content, which reduced the viability of the various plants in this 
reclamation project. 

The adjacent farm land has been severely damaged by the toxic ma- 
terial which has been washed down from the spoil banks. Loose material 
has been carried down onto the field which will undoubtedly decrease the 
productive capabilities of the land. This problem might be alleviated if 
drainage ditches were maintained near the boundary of these two 
adjacent areas. The simple construction of ditches would not eliminate 
the problem because these ditches would fill with sediment very rapidly. 

One of the initial steps in planning this study was to determine a 
site layout for the experimental plots. Preliminary field work was con- 
ducted during March, 1968, to determine the specific areas in which the 
experiments would be conducted upon the various woody species to be 
used. Two herbaceous species were planted in October of 1967, and simi- 
lar preliminary work was done prior to that time. Upon evaluating the 
slopes of the two ridges, considering the exposure of the different species 
to the sunlight and the drainage of the plots, it was decided that the top 
of the two ridges would be used for the plantings of both trees and 
grasses. Since the sides were not equal in degree of slope, it was felt 
that the flattened area on the ridge tops should be utilized, thus giving a 
more accurate means of equating the actual potential of each of the two 
ridges. These tops were struck off in 1966, six years after the strip 
mining had taken place. 

Definite plots were then staked out on the ridges. A total of twelve 
separate plots were arranged, with six being located on each ridge, and 
trees were planted at six foot intervals within these plots (Figures 1 
and 2). It was decided to use six different species of trees and to divide 
these various species into three groupings. Two of the groups were 
treated with different fungal culture filtrates, an organic matter to be 
utilized by the plants in their growth, and the third was a control group. 
In the site layout of experimental plots for the herbaceous species, less 
subdividing of the plots was necessary. In this phase of the experiment, 
only the sandstone ridge was utilized. Prior to the actual sowing of the 
grasses, the region was staked out into three separate plots (Figure 
3). In comparing the general physical background of the two spoil bank 

Geology and Geography 



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A- White Oak 
B - Red Oak 
C - Yellow Poplar 
D- Sycamore 
E- Black Locust 
F- Virginia Pine 

I - Control 

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3- Aspergillus fiavus Addition 

X- Soil Moisture Test 

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ridges, it was apparent that the greatest differences were in the acidity 
and the chemical composition of the ridge itself. Most of the other physi- 
cal properties were quite similar. 

There was a great difference in the natural vegetation found on 
each of the two spoil bank ridges within the experimental area. Numerous 


Indiana Academy of Science 


A- White Oak 
B- Red Oak . 
C - Yellow Poplar 
D - Sycamore 
E- Black Locust 
F- Virginia Pine 

1- Control 

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X- Soil Moisture Test 

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Cottonwood trees and trumpet vines were located on the sides of the 
sandstone ridge, near the site where the experimental trees were planted. 
These cottonwood trees, as well as a number of pine trees, were planted 
by the Indiana Coal Producers and appear to be growing very slowly. 
The trumpet vines and other weeds were not planted here, but have 

Geology and Geography 



Plot I 

Plot 2 , 

Plot 3 


RF |:|92 

Fig. 3 

grown on a volunteer basis. But there was absolutely no natural vegeta- 
tion to be found surrounding the test plots on the shale ridge. 

Three separate expenditures were analyzed in order to determine the 
total costs of reclaiming land that had undergone strip mining opera- 
tions. These were the costs of utilities, labor, and materials which were 
utilized in the various aspects of the reclamation experiment. The shake 

360 Indiana Academy of Science 

machine, the device in which the fungal culture nitrates were formed, 
cost one-half cent per hour to operate. It operated for 72 continuous 
hours, which was the total time required for the run to be completed. A 
total of twelve runs were made with the shake machine for an accumu- 
lated cost of $4.32. On a larger scale, the utilities would run $43.20 to 
treat one acre of land in a similar manner. The utility costs for the 
operation of the 100 gallon fermenter, with 96 hours of operation time 
per batch, amounted to $9.85. The utility cost would be $197.20 for 
one acre. The labor cost of planting and treating one acre of such land 
with trees would be approximately $71, based on the present minimum 
wage of $1.60 per hour. 

TABLE 1. Cost of Reclaiming one Acre of Strip Mined Land 

Trees Grass 

Raw Material for Media ___$19.20 Raw Materal for Media ___$ 40.00 

Seedlings 16.00 Seed 170.00 

Labor 70.95 Labor 

Utilities 43.20 Utilities 197.20 

TOTAL __.. _____ $149.35 TOTAL $407.20 

(plus labor) 

The three key materials required for this research project were 
media used in treating the experimental groups, the plants, and the 
seed. The costs of these materials were considered in order to determine 
a detailed cost analysis (Table 1). The cost of the raw materials for 
media used in the tree planting experiment was $1.92. From these costs, 
it can be extrapolated that it would cost approximately $19.20 to treat 
an entire acre of strip mine land with tree plantings in a similar manner. 
It was necessary to acquire the tree seedlings for this study in multiples 
of one hundred. Although only part of the tree seedlings were planted, 
one hundred of each of the six different species of trees were acquired 
and only the more healthy specimens were used in the actual planting 
on the spoil banks. 

By planting a total of only 180 trees on the two strip mine sites, 
the actual cost of plants used in the study was about $1.60. These were 
planted in an area which was approximately 4,000 square feet in size. In 
extrapolating this to a scale of acreage, the cost of plants necessary to 
reclaim an acre of similar land would be nearly $16. Two grasses were 
used in this study, Kentucky 31 Fescue and perennial rye. A seed mix- 
ture containing 2% lb. of each was sown on the experimental plots. The 
cost of seeding the plots was $3 and the sum of all grass seed utilized in 
the project totaled $9. The cost of seeding an acre of strip mined land in 
a similar manner would cost about $170. Since the experiments were con- 
ducted on a comparatively small area, the total cost of reclaiming this 
type of land would be much less if the practices were conducted on a 

Geology and Geography 361 

larger scale, as most certainly would be the case in state-wide reclama- 
tion endeavors. 

Another way of expressing the cost analysis of reclamation for a 
larger area in this vicinity is on a tonage basis. This figure was arrived 
at by using the average depth and thickness of the number III coal seam, 
which is presently being mined in this portion of Greene County. This 
seam, which averages two to three feet in thickness, is approximately 
sixty feet beneath the surface of the earth. If these figures remain uni- 
form throughout the project area, a total of nearly 6,000 tons of coal 
would be mined from one acre of ground. In 1968, the average cost of 
recovering this coal was about $4 per ton. It would cost an additional 
$.017 per ton of coal mined to reclaim the stripped land in trees. To 
this, one would need to add the cost of labor, which would amount to 
approximately $.007 per ton. Therefore, the total cost of reclaiming one 
acre of land to trees at the project site would amount to $149.35, which 
would average 2.4c per ton of coal mined. 

The cost of reclaiming the test site to grass land is more expensive. 
The materials and utilities needed in this operation would add an addi- 
tional 8.4c to the cost of extracting one ton of coal in the stripping meth- 
ods now in use. Labor costs would vary significantly, even on the local 
scale and, therefore, would be almost impossible to estimate with any 
degree of accuracy. However, without the labor costs being taken into 
consideration, it would cost $407 to reclaim an area of 1.0 acres to grass- 


Although the survival rates of the experimental plantings in this 
reclamation study were not outstanding, a great deal of information was 
acquired concerning the importance of the physical environment on 
various herbaceous and woody species. This strip mine site was un- 
doubtedly one of the most adverse areas in this coal mining region. Very 
few spoil banks in Greene County, Indiana had such a low pH reading 
and such a high sulphur content. By simple observing the project area, 
one was able to note the dramatic contrast between the experimental 
planting site and the other nearby stripped land, which had been natural- 
ly revegetated. Before any major steps are taken in the reclamation of 
a stripped area, considerable time should be spent to thoroughly analyze 
the physical characteristics of the region. Although some of the physi- 
cal characteristics may be favorable, one poor characteristic may cause 
the project to be unsuccessful. 

In addition, economic factors must also be considered. The overall 
cost of reclaiming spoil banks has been a major concern of the private 
coal companies. Many of these organizations feel that the initial expense 
of reclaiming the stripped land far outweighs any possible economic 
return in the near future. This concept has been diminishing rapidly, 
however, due to many successful operations in the past few years. 

On most strip mined areas in this portion of Indiana, favorable re- 
sults would be achieved by utilizing the methods practiced in this study. 

362 Indiana Academy of Science 

As previously shown in this study, it would be possible to reforest an 
acre of land for less than $150. This figure would include the cost of all 
plants, culture filtrate, and labor for the project. The economic returns 
from this endeavor would be a definite asset to the entire region. From 
the cost figures arrived at in this study, it may be too expensive to 
plant grasses on this particular strip mine site in Greene County. How- 
ever, if the area possessed more favorable physical characteristics, this 
operation would not be uneconomic, since income from the grass lands 
would eventually exceed the initial investment. 

Popcorn Production in Indiana 1 

Lowell I. Dillon, Ball State University 


Commercial popcorn production, important only since 1890, thoug-h 
showing- much fluctuation, has increased greatly since the early 1940's. 
National harvested totals in the last decade have averaged approximately 
180,000 acres and 416 million pounds. The crop's value has been as high as 
15.4 million dollars. Most production is in the Corn Belt with Indiana the 
leader in most recent years. Production by various counties has been quite 
variable but the chief areas has recently been in northeastern Indiana and 
particularly Huntington County. Optimum physical conditions and methods 
of cultivation are similar to those for field corn but more care is necessary 
in harvesting and storage. The chief reasons for the importance of popcorn 
production in Indiana are the government feed grains allotment program 
and the location here of established buyers and processors. The state 
should continue as one of the major United States producers. 

Popcorn was probably grown by the Indians of both North and 
South America. It has been important commercially, however, only 
since about 1890. Production, though showing a great deal of fluctuation 
from year to year, has increased greatly since the early 1940's (1). This 
coincided approximately with the development of popcorn hybrids, but 
growing demand would probably have caused an increase anyhow. In 
the last decade the national harvested area has varied from about 109,- 
000 to over 240,000 acres with an average of approximately 180,000 
acres. Pounds produced have varied from 272 million to 518 million, 
averaging about 416 million (Table 1), and value of the harvest has 
ranged from about 6.2 million to 15.4 million dollars (2) and (3). 

As shown on Table 1, the commercial crop is chiefly produced in the 
Corn Belt states of Ohio, Indiana, Illinois, Iowa, Missouri, and Nebraska, 
although the bordering states of Michigan and Kentucky produce signifi- 
cant enough amounts to be listed in the annual reports of the Department 
of Agriculture. Minor amounts are at times reported from states as far 
apart as Maryland, Texas, and Idaho. Indiana or Iowa is usually the 
leading state with Illinois third (2). The exact ranking of states is some- 
times questionable, as statistics from the two major sources, the Census 
of Agriculture and Agricultural Statistics do not always agree. Based on 
measurements of the last ten years by the latter source, Indiana is the 
leader in both harvested acreage and in total pounds produced. 

Within Indiana the leading producing counties are scattered widely 
over the state. Huntington County has been the leading producer in the 
two most recent agricultural census years. Other leading counties include 
Kosciusko, LaPorte, Vermillion, Harrison, Wells, and Noble (Table 2). 
As has been true nationally, there has been considerable fluctuation of 

1 Appreciation is expressed for the assistance of Mr. Max G. Miller, 
County Extension Agent — Agricultural, Huntington County ; Mr. Jack Wade, 
U.S. Soil Conservation Service, Huntington County; and Mr. Eugene 
Kiracofe, Fieldman, Weaver Popcorn Co., Van Buren, Indiana, in gathering 
information for this paper. 


364 Indiana Academy of Science 

production from county to county within the state. At various census 
periods LaGrange, Parke, Tippecanoe, and Vigo Counties have ranked 
high (4). In the post-war period the most stable areas of important pro- 
duction have been in northeastern Indiana in Huntington, Wells, Noble, 
and Kosciusko Counties. Even here, however, stability is only relative 
and there have been significant changes from year to year in both acreage 
and production. 

Here the question arises as to why Indiana is a leading popcorn pro- 
ducer and why popcorn production is of major importance in certain 
counties and not in others. 

The usual methods of commercial popcorn growing are very similar 
to those used for field corn (1). Physical conditions of climate and soil 
which encourage the growth of one also encourage the other. Some 
growers claim that a somewhat slower ripening period is better for pop- 
corn. This would favor the more northern Corn Belt areas. This surely 
is only a minor factor. It has not significantly discouraged growers in 
such areas as southern Indiana and Kentucky. 

Methods of planting and cultivation are nearly identical and fertiliza- 
tion varies only slightly. There are major differences, however, in har- 
vesting and curing the crop. While a mechanical com picker can be 
easily adapted to harvest popcorn, this crop is always harvested as ear 
corn. Most field corn is both picked and shelled in the field. Further, 
damage to the standing crop by wind, rain, or hail may have a more 
serious effect on the harvesting of popcorn than on that of field corn. 
Popcorn is more susceptible to picking damage than field corn since 
cracked or broken kernels are useless for popping. 

Storage and curing practices are also much more important in the 
production of popcorn. Since popcorn is grown for human consumption, 
standards are higher than for field corn and rodent or insect damage 
may make the crop completely unacceptable to the processor. In this case 
the farmer can only use his popcorn for animal feed, a use for which 
field corn is far more suitable. 

Moisture content is also of paramount importance. The crop should, 
if possible, be completely mature before the first killing frost, and slow 
natural drying in the field or crib is recommended. Artificial drying, how- 
ever, is common but must be done carefully. Too rapid loss of moisture 
may reduce popping expansion resulting in a lower price or loss of 

Such differences in harvesting or curing, however, are not the major 
factors involved in the decision of whether or not to grow popcorn. The 
greatest differences in field corn and popcorn production and the most 
important determinants in the farmer's choice of crops are governmental 
and/or economic. 

The government feed grains program is of major importance. Under 
this program the acreage allotted to a farm for the growing of corn, 
wheat, sorghums, and other feed grains has been calculated on the 

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Geology and Geography 367 

basis of the acreage planted in these crops in the 1958-1959 base year. 
This has affected popcorn production in two ways. In some parts of 
Indiana farmers were at that time already growing popcorn fairly exten- 
sively. Their acreage allotment for feed grains was thus somewhat low 
when compared to their total farm acreage. The farmer thus is encour- 
aged to continue to use his unallotted land for popcorn or, for the only 
other real cash crop choice, soybeans. This has tended to cause permanent 
popcorn production by the same farmers over a longer period of time. 

Acreage allotments have also, since they restrict the planting of 
feed grains, left surplus land available for crops not covered by the 
federal programs and have encouraged some farmers who had not grown 
popcorn previously to enter into this type of agriculture. Again, as men- 
tioned above, the only real cash crop choice is soybeans. The crop planted 
will depend upon the farmer's opinion as to which will be the more 
profitable. In the case of popcorn he will have a more exact idea of 
his profit since in most cases, a definite price has been promised him and 
the only variables are yield and quality. Also, he may get a bonus if 
quality is especially good. 

Although all the above factors no doubt affect the farmer's decision 
and play a part in the concentration of popcorn production in certain 
areas, the most important determinant is the location of an established 
processor. The processor has the necessary equipment for drying, shell- 
ing, storage, and packaging; has wholesale and retail markets available; 
has transportation facilities; and keeps a close eye on national demand 
and holdover supplies. Based on this knowledge he makes his acreage 
contracts for the year. He makes a firm promise as the price per pound 
harvested, granted acceptable quality, and for all practical purposes, 
determines the annual acreage planted. Very few farmers, unless they 
are also processors, are prepared to make the investment in time and 
money required and to take a chance on the future market. 

No one can be definite as to the future trends in popcorn production 
generally. Factors involved include the general economic prosperity of 
the country, competition from the myriads of other "snack foods" now on 
the market or to be developed in the future, and possible expansion of 
foreign markets. Even now popcorn is being exported to Japan and to 
western Europe in small amounts. Future growth of these markets 
depends on successful educational and promotional campaigns. There 
may also be some other possible markets as yet untapped in relatively 
prosperous areas of the world, but the vast, underdeveloped regions offer 
little encouragement. 

Indiana will probably continue as a major producer. The present 
markets built up by Indiana processors and lower shipping costs to the 
populous eastern states should insure our state a continued important 
share of total national production. 

368 Indiana Academy of Science 

Literature Cited 

1. Brunson, Arthur M. and Dbwatne L. Richardson. 1958. Popcorn. Farmer's 
Bulletin No. 1679. U.S.D.A., Washington. 17p. 

2. U.S. Department of Agriculture. 1957-1967. Agricultural Statistics. Gov- 
ernment Printing Office, Washington. 

3. U.S. Department of Agriculture, Purdue University Agricultural Experi- 
ment Station. 1968. Popcorn Acreage and Production Forecast. 

4. U.S. Department of Commerce, Bureau of the Census, 1954, 1959, 1964. 
Census of Agriculture, Government Printing Office, Washington. 


Chairman: Robert H. Cooper, Ball State University 
B. Elwood Montgomery, Purdue University, was elected chairman 

for 1969 

Other papers read 

Some Fragments of Indiana Entomological History. 

B. Elwood Montgomery, Purdue University 

Academic Origins of Members of the Genetics Society of America 

Donna Howard and Thomas R. Mertens, Ball State University 


The academic and geographic origins of current members of the 
Genetics Society of America who received their undergraduate degrees from 
U. S. institutions and who are listed in the 1965 edition of American Men of 
Science were investigated. The 1019 geneticists included in this study 
received their baccalaureate degrees from 281 different institutions of 
higher education. The 999 who have earned doctorates were awarded these 
degrees by 93 different institutions. Land-grant and state colleges and 
universities were productive of geneticists at both the baccalaureate and 
doctoral levels; e.g., eight of the ten leading institutions granting bacca- 
laureates to the geneticists were state institutions, and over 60% of the 
geneticists' doctorates were awarded by such schools. Over 45% of the 
geneticists were born in eight states in the Middle Atlantic and East 
North Central United States. Of these eight states, Indiana was the least 
productive, only 20 of the 460 geneticists from these states having been 
born in Indiana. Currently 36 of the 1019 individuals in this study are 
employed in the state of Indiana. The states in Southeastern United States 
failed to produce as many geneticists as they employ. Only one state, 
Mississippi, failed to produce any of the 1019 geneticists. 


A study of members of the Genetics Society of America was con- 
ducted to reveal the geneticists' academic and geographical backgrounds, 
which institutions graduate the greatest numbers of geneticists, and the 
types of institutions in which geneticists find employment. Only the 1019 
members of the Genetics Society of America who earned baccalaureate 
degrees at U.S. institutions and who were listed in the eleventh edition of 
American Men of Science (1) were included in this study. Using this pro- 
cedure may be expected to result in the inclusion of active researchers 
and contributors to the discipline of genetics, and it eliminated the need 
for sending questionnaires to the individuals who are the subjects of this 
investigation, since all pertinent data were directly obtained from 
American Men of Science. 

The growing need for highly trained scientists has been especially 
recognized since the early 1950's. Since that time there have been several 
studies of the academic backgrounds of scientists in general, the most 
extensive having been conducted by Lindsey R. Harmon, Director of 
Research for the Office of Scientific Personnel of the National Academy 
of Sciences. In his 1965 study, Harmon stressed the importance of 
research dealing with the academic origins of scientists: 

Advancement of the public welfare and development of all aspects of a 
modern technological society are intimately bound up with the educa- 
tion of an adequate number of the society's members to the highest 
levels of wbich they are capable. . . . Comparatively little is known, 
of a quantitative nature, regarding the career patterns of the most 
highly-trained segment of the populace, and but little more is known 
about the backgrounds from which it comes. (5) 

Lyon studied the origins of American botanists, and also stressed the 
importance of researching the academic origins of scientists: 


History of Science 371 

Scientists should learn more about themselves than is possible through 
personal observations. As a basis for maintaining the proper supply of 
trained men in each of the special fields, we should have accurate 
information about the number, ages, and professional preparation of 
the workers in each area. (8) 

The research most closely related to the present investigation is 
Chiscon's "The Academic Origin of Drosophila Workers in the United 
States" (2). In his study, Chiscon determined the academic origins of 
471 Drosophila workers. As sources of data, he utilized the volumes of 
American Men of Science and Drosophilia Information Service. 

The present study of 1019 members of the Genetics Society of Amer- 
ica identifies those institutions which have provided instruction in the 
field of genetics, the genticists* areas of research specialization, the types 
of academic backgrounds the geneticists have, the types of institutions 
in which the geneticists find employment, and the regions of the U.S. 
which have been the most productive of geneticists. 

Materials and Methods 

The names and current addresses of the geneticists were taken from 
the 1967 membership list of the Genetics Society of America (3). The 
1965 edition of American Men of Science was used as a source for the 
following additional information which was recorded on a 5" x 7" card for 
each geneticist: age, sex, date and place of birth, degrees earned, institu- 
tions where degrees were earned, years in which degrees were earned, 
age when degrees were earned, post-doctoral experience, current institu- 
tion of employment, and field of specialization. 

The geneticists' ages were determined from the birth date and were 
calculated as of May, 1968 — the anticipated completion date of this inves- 
tigation. The World Almanac (10) was used to determine whether the 
institutions which granted the degrees were state, land-grant, denomi- 
national, private, etc. These data were then analyzed in order to obtain 
generalizations about geneticists in the United States. 

Data and Discussion 

Academic origins. The baccalaureate origins of the geneticists were 
more varied than the doctoral origins. There were three times as many 
baccalaureate institutions as there were doctorate institutions. The 1019 
baccalaureates earned by the geneticists were awarded by 281 different 
undergraduate institutions. One hundred forty institutions awarded only 
one baccalaureate, while 53 each awarded two such degrees. The remain- 
ing 88 institutions granted the balance of 773 baccalaureate degrees. 

Both state and private institutions were productive of geneticists at 
the baccalaureate level. Land-grant institutions awarded 35.7% of the 
baccalaureate degrees in the present investigation. In Chiscon's study of 
the academic origin of Drosophila workers, 23% of the baccalaureate 
degrees were awarded by land-grant institutions (2). In the present 
investigation, land-grant and state colleges and universities together 

372 Indiana Academy of Science 

granted 595 baccalaureates, 58.4% of the total, while private and denomi- 
national institutions awarded 36.8% of the total, or 375 baccalaureates. 
Kiefler observed that the nation's liberal arts colleges have played an 
important role in producing scientists and have produced more than half 
the science doctorates in the past (7). In the present study, however, 
only 36.8% of the baccalaureates were awarded by liberal arts colleges. 

In the top-ten institutions granting baccalaureate degrees were eight 
state and two private universities (Table 1). Concerning the productivity 
of institutions in relation to the natural sciences, Thistlewaite stated: 

Natural science productivity is associated with large freshman enroll- 
ments, graduate programs offering the Ph.D., public support, and 
absence of religious affiliation. These characteristics are typical of the 
state university, which . . . tends to be outstandingly effective in 
stimulating achievement in the natural sciences. (9) 

The present study supports Thistlewaite's generalization in two 
ways: 1) land-grant and state colleges and universities awarded 58.4% 
of the baccalaureates and 2) eight of the ten leading institutions grant- 
ing baccalaureates were state universities. 

Comparing the ten leading baccalaureate institutions in the present 
investigation with the ten leading institutions in Chiscon's study of 
Drosophila workers and with Lyon's study of botanists revealed that 
many of the institutions which appeared in the top-ten in this study were 
also in the top-ten in Chiscon's and Lyon's studies. California and Illinois 
were among the ten leading universities in all three studies. Harvard 
and Texas were among the ten leading universities granting baccalaure- 
ates to the members of the Genetics Society of America and also to 
the Drosophila workers in Chiscon's study (2). Cornell University and 
the universities of Minnesota, Nebraska, and Wisconsin were among the 
ten leading institutions granting baccalaureates to the members of the 

table 1. Top Ten Institutions Granting Baccalaureate Degrees to 
Members of the Genetics Society of America 

Number of 
University Degrees Granted 

Cornell 38 

California (Berkeley) 36 

Illinois 35 

Iowa State (Ames) 31 

Minnesota 29 

Nebraska 24 

Wisconsin 21 

Harvard 20 

Chicago 18 

Texas 18 

History of Science 373 

Genetics Society of America and also to the botanists in Lyon's 
study (8). 

In the present study, 999 of the 1019 geneticists completed doctoral 
degrees; 954 earned the Ph.D. degree. Ninety- three institutions awarded 
these doctorates. Among the ten leading institutions granting doctorate 
degrees were six state and four private universities (Table 2). In 
general, the top-ten baccalaureate institutions were also the top-ten doc- 
torate institutions. However, Columbia and Yale, which were among the 
ten leading doctorate universities, replaced the universities of Nebraska 
and Illinois, which were among the ten leading baccalaureate institutions. 

table 2. Top Ten Institutions Granting Doctoral Degrees to 
Members of the Genetics Society of America 

Number of 
University Degrees Awarded 

California (Berkeley) 83 

Columbia 79 

Wisconsin 68 

Harvard 56 

Texas 53 

Iowa State (Ames) 51 

Cornell 50 

Yale 43 

Minnesota 38 

Chicago 35 

Land-grant institutions awarded 38.7% of the geneticists' doctoral 
degrees. Land-grant and state institutions together granted 605 doctor- 
ates or 60.5% of the total, while private and denominational institutions 
awarded 381 doctorates, or 38.1% of the total. 

Geographic Origins. To reveal which areas of the U.S. were the 
most productive of geneticists, the geneticists were divided according to 
the nine regions of the U.S. in which they were born (Table 3). These 
nine regions were used in Harmon's 1961 study (4). The total number 
of geneticists produced by all the regions is slightly less than the 
number employed, because the places of birth for six of the geneticists 
were not listed in American Men of Science. 

Together the Middle Atlantic and East North Central states pro- 
duced 45.5% of the geneticists. Indiana was the least productive of the 
five states included in the latter region (Table 3). Only 20 of the geneti- 
cists included in this investigation were born in Indiana and of these six 
retain their Indiana residences. Thirteen of the 20 obtained baccalaureate 
degrees from institutions in the state, while only six earned their doc- 
torates within the state. A total of 27 geneticists (Table 4) earned 

374 Indiana Academy of Science 

table 3. Productivity of Regions of the United States: Comparison 

Between Number of Members of the Genetics Society of America 

Born and Employed in Each Region 

Number Born 

Number Employed 


in Region 

in Region 

New England 



Middle Atlantic 



East North Central 


















West North Central 



East South Central 



South Atlantic 



West South Central 















baccalaureate degrees from Indiana schools. Indiana and Purdue Univer- 
sities ranked thirteenth and fourteenth, respectively, in the nation in 
granting doctoral degrees to geneticists in this study, Indiana having 
granted 27 and Purdue 23. 

Harmon and Soldz' 1960 study (6) showed that the regions producing 
the greatest number of science doctorates are the Middle Atlantic, East 
North Central, and Pacific. The latter, however, has not been highly pro- 
ductive of geneticists (Table 3). Among the states New York was the 

table 4. Indiana Institutions Granting Baccalaureate Degrees to 
Members of Genetics Society of America 

Number of 
Names of Institutions Degrees 

Indiana University 13 

Purdue University 9 

Wabash College 3 

Ball State University 1 

Indiana Central College 1 

Total 27 

History of Science 375 

most productive (173 geneticists) and Mississippi the least productive 
(no geneticists). Mississippi is included in the East South Central states, 
which generally have been low producers of scientists. 

Comparisons of Data Concerning Male and Female Geyieticists 

In Harmon's study of Ph.D.'s in the sciences, slightly over 10% of 
the doctorate holders were women (5). In the present investigation of 
1019 geneticists, 12.5% or 127 were women. 

Thirty-eight percent of the men but only fifteen percent of the 
women earned baccalaureates at land-grant colleges and universities. The 
women, however, were more frequently graduated from private colleges 
and universities than were the men. Land-grant institutions awarded 
30.6% of the women's doctoral degrees and 39.8% of the men's doctorates. 
Private colleges and universities awarded 45.4% of the women's doctoral 
degrees and 36.6% of the men's doctorates. Institutions outside the 
United States granted 1.2% of the men's doctoral degrees, but none of 
the women earned doctorates from foreign universities. 

A higher percentage of women are employed by private colleges 
and universities, while a higher percent of the men find employment in 
state colleges and universities. Colleges and universities employ 77.2% 
of the women and 78.6% of the men. In Harmon's 1965 study (5) a 
higher percentage of men than women found employment in business 
and industry. Among the geneticists, however, no difference between the 
sexes is apparent with 6.0% of the men and 6.3% of the women finding 
employment in business and industry. 

In general, the men completed the doctorate in fewer years following 
completion of the baccalaureate than did the women. Five years after 
the baccalaureate, 47.2% of the men had earned the doctorate; but only 
26.5% of the women had completed the doctoral requirement. The aver- 
age time beyond the baccalaureate for the male to complete the doctorate 
was 6.7 years, while the 121 women completing that degree required 

table 5. Areas of Research Specialization of Members of the 
Genetics Society of America 

No. of 


No. of 


Area of Genetics 











































376 Indiana Academy of Science 

an average of just under 9 years. Only 9.7% of the men but 20.7% 
of the women earned doctorates after age 34. None of the 121 women 
earned a Sc.D. degree, but 1.8% of the men earned that degree. The 
M.D. degree was earned by 3.0% of the men and almost 3.0% of the 
women. The majority of both sexes earned the Ph.D. degree. Certain 
areas of research were more attractive to men than to women (Table 5). 
Men more frequently specialized in plant genetics, statistics, and bio- 
physics than did the women; whereas, women more frequently entered 
the fields of microbial and biochemical genetics than did the men. About 
50% of both men and women have specialized in some aspect of animal 


The data obtained in this study justify the following conclusions: 

1. The baccalaureate origins of the geneticists were more varied 
than their doctoral origins, the baccalaureate degrees having been 
awarded by 281 different institutions and the doctorates having been 
granted by 93 institutions. 

2. Private colleges and universities produced over one-third of the 
geneticists, while state and land-grant institutions granted over 50% 
of their baccalaureates and over 60% of their doctorate degrees. Eight 
of the ten leading institutions granting baccalaureates were state institu- 
tions; six of the ten leading doctorate institutions were state-supported. 

3. Many of the ten leading institutions granting baccalaureates to 
the geneticists also were included in the leading institutions granting 
baccalaureates to botanists and Drosophila workers. This may be indica- 
tive that certain institutions are generally productive of life scientists. 

4. The Middle Atlantic and East North Central states constituted 
regions in the U.S. that were most productive of geneticists. Indiana pro- 
duced only 20 of the 1019 geneticists in this study. 

5. At both the baccalaureate and doctorate level, women geneticists 
were more frequently graduated from private colleges and universities 
than were men. The men more frequently earned degrees from land- 
grant colleges and universities. Geneticists tended to find employment 
in the kinds of institutions from which they earned their degrees; a 
higher percentage of women were employed by private colleges and 
universities, whereas a higher percent of the men found employment 
in state colleges and universities. 

Literature Cited 

1. American Men of Science. The Physical and Biological Sciences. Edition 
11. 1965. R. R. Bowker Company, New York. 

2. Chiscon, J. Alfred. 1956. The academic origin of Drosophila workers in 
the United States. J. Heredity 47:292-295. 

3. Genetics Society of America Supplement: Directory of Members of the 
Genetics Society. 1967. Genetics 56 (3-2): s27-s62. 

History of Science 377 

Harmon, Lindsey R. 1961. High school backgrounds of science doctor- 
ates. Science 133:679-688. 

-. 1965. Profiles of Ph.D.'s in the Sciences. National Academy 

of Sciences, Washington, D.C. 

6. Harmon, Lindsey R. and Herbert Soldz. 1960. The Science Doctorates of 
1958 and 1959. National Science Foundation, Washington, D.C. 

7. Kiefler, William F. 1967. Editorially speaking. Chem. Education 44:119. 

8. Lyon, Charles J. 1957. Origins and status of American botanists. Science 

9. Thtstlewaite, Donald. 1959. College environments and the development 
of talent. Science 130 :73. 

10. The World Almanac. 1968. Newspaper Enterprise Association Inc., 

Original Science Apparatus Preserved in 
Science Museums and Universities in Free Europe 

N. G. Sprague, Ball State University 

Original science apparatus is where you find it: Florence, Leyden, 
Glasgow, or Cambridge. A search during the spring of 1968 for original 
apparatus behind present-day physics and astronomy led the author 
through many interesting by-ways of free Europe. 

Using World of Learning and American Ephemeris and Nautical 
Almanac as sources for addresses, one hundred scientists and museum 
directors and /or curators were contacted in fifty observatories and 
museums. All persons were cordial in their response concerning a visit. 
Original pieces of science apparatus were located in twenty museums in 
fourteen countries. The number of items ranged from one or two to an 
entire room dedicated to a single scientist. It was observed that there 
was twice as much stored away as there was on display. This probably 
accounted for the overcrowding observed in most museums. Some pieces 
of apparatus were displayed in a regular museum, some in the seminar 
room of a university department, and others along corridors. 

In 1657, Cardinal Leopoldo de 'Medici together with his brother, the 
Granduke Ferdinando II — both followers of Galileo — founded in Flor- 
ence the Accademia del Cimento, the first modern scientific institute in 
Europe. In the Museum of the History of Science, situated in the medi- 
eval Castellani Palace in Piazza dei Guidici near the Arao River, many 
of the most interesting and valuable relics of Galileo and his pupils of 
the Accademia del Cimento are preserved. 

Many scientific objects in the outer rooms tend to divert the visitor, 
however one pushes forward to the blue velvet lined room of Galileo 
Galilei. There are two of the telescopes with which he studied the skies, 
discovering the valleys of the moon, the phases of Venus, and four of 
the satellites of Jupiter; his campassec; pendulum diagrams and weights; 
air thermometer; astrolab; and publications. Some of these were re- 
lated to his "swinging lamp" and "leaning tower" experimentation in 
Pisa, about fifty miles to the west. 

Instruments of one of his famous pupils, Evangelista Torricelli, are 
there: a metal case telescope and his 1643 mercury tube barometer. 
Several rooms are filled with later dated telescopes of unusual designs 
and mountings. Others contain case after case of astrolabs and measur- 
ing devices. In the center of one room was a large 16th century armillary 
sphere made by the Florentine instrument maker, Volpaja. Elsewhere in 
the museum were located a lens grinder, ruling machine, and spectro- 

Three weeks later a large collection was photographed in the Rijks- 
museum voor de Geshiedenis der Natuurwelenschappen at Leyden, Hol- 
land. The name brings to mind windmills and capacitors. They had both. 


History of Science 379 

However, it was also the home territory of Huygen, Leeuwenhoek, and 

There, in 1656, Christian Huygens made the important invention of 
the pendulum clock, which had been conceived independently by Galileo 
fifteen years earlier but had never been expanded to its basic capa- 
bilities. Huygens also applied the clock principal to a plate-type plane- 

Antony van Leewenhoek in the 1670's made the first study of the 
miniature water-world with single lens microscopes. He taught himself 
to grind lenses and made microscopes capable of magnifying up to 270 
times. An entire room is given over to his microscopes and later im- 
provements in the field of microscopy. 

From a much later date there is the apparatus of Heike Kamerlingh 
Onnes, the 1913 Nobel prize for physics winner. With it he discovered a 
method of liquefying helium. 

Across the English channel science history was also made, and many 
pieces of apparatus are still preserved. A very fine collection from Lord 
Kelvin and James Joule was found in the new Physics Building of the 
University of Glasgow, Scotland. 

Early in life Lord Kelvin became fascinated with the possibilities 
of electrical currents for signalling over long distances and soon was 
personally involved in the company that was trying to lay the first 
Atlantic cables. In one display case were many pieces of apparatus 
from that association: a high voltage electrostatic voltmeter; the "cable" 
galvanometer used to receive the first message, "Europe and America 
are united by telegraphic communication! Glory to God in the Highest 
and on earth peace and good will to all men," which was sent by cable 
under the Atlantic Ocean; a Centi-ampere Balance with which Lord 
Kelvin originated the principle of "weighing" electric current by balanc- 
ing the attraction between sets of coils against weights; and a Mouse 
Cage Electric Motor. 

Kelvin was also noted for some of his teaching devices: the French 
horn which he himself played in the classroom to illustrate problems in 
acoustics; a ballistic momentum-impulse firearm; pitch glacier started 
1890; and Call Box with its three compartments nick-named by his 
students as "purgatory, heaven, and not passed." 

James Prescott Joule carried out much scientific research in his own 
private laboratory. However, some of his discoveries were made with 
Lord Kelvin. It is fitting that several pieces of his apparatus are also 
included in the collection. A Joule Magnetic Engine, an electric motor 
made by Joule in 1840-3, is one of the earliest motors in existence. It was 
undoubtedly used in his studies on eletrical energy. A Joule Calorimeter 
made of brass was used by Joule in the final determination of the me- 
chanical equivalent of heat. 

In England one would expect to find science "originals" at Oxford 
and Cambridge. Henry Gwyn Jeffres Moseley, while a lecturer at Ox- 

380 Indiana Academy of Science 

ford, proved the existence of a simple relationship between the X-ray 
spectrum of an element and its atomic number, thereby showing that 
the properties of an element are determined, not by its atomic weight, 
but by its atomic number. He thus provided a new method of chemical 
analysis which has since aided immeasurably in solving various out- 
standing problems of atomic structure. Several of his X-ray spectrom- 
eters are located in the Museum of the History of Science. 

At Cavendish Laboratory, just off of Free School Lane in Cambridge, 
are located several items used in their research by many famous scien- 
tists of the late 19th century and the beginning of the 20th. Found there 
on the third floor in cases along a hall are J. J. Thomson's positive ray 
apparatus (complete parabolae were first recorded with this model); 
Aston's original mass spectrograph (without the magnet); and C. T. 
R. Wilson's cloud chamber, which was used without any major altera- 
tions for all the photographic work he did. 

The thinking which was behind these originals is very impressive. 
It is hoped that all students, in future classes where various pieces of 
apparatus will be used, will recognize and be inspired by our scientific 

A Half Century of Science at Ball State University 

Robert H. Cooper, Ball State University 

In October, 1952, 0. B. Christy and Robert H. Cooper co-authored 
a paper, "The First Thirty-four Years of Science at Ball State Teachers 
College," which was presented at the Fall meeting of the Indiana 
Academy of Science at Valparaiso University. The paper reviewed a 
few of the professors who contributed much to the development of 
the institution and a number of graduates who have done much in the 
field of science at various locations in the world and in numerous pro- 
fessions. The present paper reviews in some detail the administrative 
and curricular science development of Ball State and brings the in- 
formation up-to-date. 

The Administration Building, where science was first taught on the 
Ball State University campus, was built in 1898. The institution went 
by a number of names up to June 17, 1918, when it became known as the 
Indiana State Normal School — Muncie Branch. For some years the 
president of the Indiana State Normal — Terre Haute Branch also served 
in that capacity for both schools. 

The professors who stand out as setting up the initial high standard 
of training in the fields of science included Frederick J. Breeze, Pro- 
fessor of Geography and Geology; Dr. Otto B. Christy, Professor of 
Botany and Agriculture; Richard A. Gantz, Professor of Physiology and 
Zoology; Frank V. Graham, Professor of Chemistry and Physics; Harry 
H. Howick, Assistant Professor of Mathematics, Physics, and Chemistry. 
These five instructors were outstanding teachers and the author of this 
paper had the privilege of having classes with each of them. Professor 
Graham in Chemistry remained in the institution until his death in 1944; 
Professor Howick in Physics remained in the institution until his retire- 
ment in 1956; and Dr. 0. B. Christy in Biology remained with the insti- 
tution until his retirement in 1950. Dr. Christy is still living and resides, 
along with Mrs. Christy, at the Teachers Retirement Home (Greenwood 
Village), Greenwood, Indiana. 

In February 1921 the Indiana General Assembly appropriated $125,- 
000 to start the work on the Science Hall. In 1923 the State provided an 
additional $128,000 to complete and also to equip the Science Building. In 
the Fall of 1923 some classes were held in Science Hall before its entire 
completion. The building was not in full operation until the summer of 
1924. At that time it had located within it the Department of Business 
Administration, as well as English, Mathematics, and Science. 

During the Spring of 1924 a change too place in the college and in 
the science program which helped many of the teachers who were work- 
ing in eight-month schools. Announcement was made of a six-week Mid- 
Spring Term from May 5 to June 13, 1924. The terminology Summer 
Quarter was now changed to First Summer Term and Second Summer 


382 Indiana Academy of Science 

Term. This made it possible for a student in an eight-month school to 
enter Ball State at the Mid-Spring Term and continue with the First 
and Second Summer Terms and thus get in a half year of credit. 

On September 9, 1929, Burris Laboratory School was opened. This 
was to be a very important part of Ball State for the remainder of the 
half century and more. Dr. Earl Johnson started as principal and be- 
came Professor of Education and finally Dean of the College of Edu- 
cation when Ball State became a university in 1965. 

In 1930-31 the course numbering changed to 100's, 200's, 300's and 
400's. The second digit in the listings under the different divisions was 
the key to that division — thus 2 became the number indicating the Divi- 
sion of Biology, so Science 120 was the freshman General Biology; 3 
indicated the Division of Agriculture; 4 indicated the Division of Chemis- 
try; 5 indicated the Division of Geography and Geology; 6 indicated the 
Division of Physics; and 7 indicated the Division of Human Physiology 
and Hygiene. 

Dr. Willis S. Blatchley, famous author-naturalist of Indiana, visited 
Ball State Teachers College, and lectured during the early 1930's. Dr. 
Alfred Kinsey came to Ball State and lectured and conducted field study 
with children in Christy Woods during the 1930's. Both men were very 
skillful in field work and were very knowledgeable. 

Mr. Frank Wallace, State Entomologist, was a guest speaker and 
adviser in the Science Department many times through the years. His 
excellent photography, his wholesome Hoosier philosophy, and his 
sage advice were of much value. His death in May 1968 was a real loss 
to Indiana. 

Dr. Charles Deam, one of Indiana's most important plant taxono- 
mists, aided with advice concerning Christy Woods and other work within 
the Science Department. The numerous visits by the flora classes to his 
well-labeled arboretum at Bluffton were of inestimable value. During 
these visits he gave much valuable information to the Ball State stu- 
dents. His death occurred in 1953. 

For a number of years the curriculum in Agriculture was an inte- 
gral part of the work at Ball State. The institution was fortunate to 
have a 17-acre woods and arboretum developed by Dr. Christy, known 
later as Christy Woods. More recently the institution purchased 16 acres 
about two miles from the campus which became known as Ball State 
University Wildlife Preserve, and is an area for study and for collect- 
ing both fauna and flora. Christy Woods has become an outstanding 
research and teaching laboratory with a sizeable greenhouse and prepa- 
ration facility. 

In more recent years Ball State University has offered curricula for 
the Nursing Degree, the Degree in Medical Technology, the Pre-medical 
Program, the Pre-dental Program, the Pre-engineering Program, the 
Pre-veterinary Program, and the Pre-pharmacy Program. 

Dr. Donald Crooks joined the Biology staff in 1929 and later became 
a research worker in the U. S. Department of Agriculture, specializing 

History op Science 383 

in the research on the virus of tobacco. He retired December 15, 1967, 
with honors. Dr. Floy Hurlbut, a geographer, joined the staff in 1931 
and was a very outstanding teacher in the field of Geography. She re- 
tired from Ball State in 1954. Dr. Donald E. Miller joined the staff as a 
Professor of Biology in 1936 and has done a fine piece of work with the 
Indiana Academy of Science, with the counseling of Pre-medic stu- 
dents, with his own teaching, and with Sigma Zeta, national science 
honorary organization. Dr. Robert H. Cooper, Professor of Biology, joined 
the staff in 1936 and became Head of the Department of Science after 
Dr. Christy's retirement in 1950. He continued in this capacity until 
1965 when he became Coordinator of Sciences and Mathematics until 
his retirement in 1968. 

In 1947 Dr. P. A. Wiseman, with special training in Organic Chemis- 
try, joined the staff and is at the present time Chairman of the De- 
partment of Chemistry in the new Physical Science Building. Dr. Jerry 
J. Nisbet and Dr. George W. Welker joined the Science Department in 
1950. With the organization of separate departments Dr. Nisbet became 
Chairman of the Department of Biology. Dr. Welker has counseled in 
Medical Technology and aided students in many ways. Malcom E. 
Hults joined the Ball State staff in 1953 and has given much time to 
research and to the development of the Department of Physics, of 
which he is now Head. Dr. George F. Beatty, a graduate of Ball State, 
returned to his Alma Mater in 1958 as Professor of Geography and Geol- 
ogy and is presently Head of the Department of Geography and Geology 
and Professor of Geography and is located in the new Physical Science 
Building with the other members of his staff. Dr. Warren E. Schaller 
came to Ball State University in 1959 and has co-authored a college 
health text and has done much toward organizing and reorganizing the 
certification of teachers of health in the State of Indiana. He is cur- 
rently Chairman of the Department of Physiology and Health Science. 

Through the years Ball State University has moved from offering 
the Two-year Certificate for Teaching to the Four-year Bachelor's De- 
gree in the many fields of teaching and research. The Master of Arts in 
Education, the Master of Arts and the Master of Science Degrees are 
offered as the beginning of the graduate work. In recent years more ad- 
vanced degrees have been offered and at the present time the Doctor 
of Education Degree with specialization in Science Education and speci- 
fically a major in the Department of Biology is being offered. Minors 
for the doctoral degrees are available in the other fields of science. 

There was a Physical Science Club devoted to the consideration of 
physics and chemistry problems listed in the catalog in 1937-38. Later 
there was a Science Club to represent all of the sciences. In 1938 the 
Sigma Zeta math and science honorary was organized and has served 
as a fine stimulus for science research. 

In September of 1939, under the authorship of Dr. 0. B. Christy, a 
bulletin was put out entitled An Outdoor Laboratory at Ball State 
Teachers College. This was a bulletin concerning the history and use of 
Christy Woods and was the first of its kind. It was revised in 1960. 

384 Indiana Academy of Science 

Listed in the catalog of 1939-40 is the Shrawder Collection. Through 
contributions of the George A. and Frances Ball Foundation, the William 
H. Shrawder Collection of rocks and minerals was added to the science 
geology collection and is still carefully maintained as one of the finest 
in this part of the United States. Mr. Shrawder, a resident of Indiana, 
was a teacher of geology in Schenley High School, Pittsburgh, Pennsyl- 

With the publication of the 1951-53 catalog, an offering was listed 
as Science 400, Science of Distant Areas, four or eight quarter hours 
credit. This listing finally led to rather extensive field study in various 
parts of the United States and in other countries. The purpose of the 
course was to help the student become acquainted within the five weeks' 
time, with the flora and fauna, the agriculture and conservation, and 
any unique science peculiar to the area. Beginning with the summer of 
1956 this became a reality in that a class was organized. Since that 
time, under Dr. Robert Cooper, five summer studies have been offered 
in the Northwestern Rocky Mountains areas of the United States and 
the Southwestern Rockies of Canada; four summers of study have been 
spent in the State of Alaska, including the Point Barrow and Pribilof 
Island areas, two summers in the State of Hawaii, including field work 
in six of the islands; and one summer in the study of countries around 
the world, including Norway, Switzerland, Africa, India, Australia, New 
Zealand, and Fijii. Also, field study, under Dr. Forrest Stevenson, has 
been done in Jamaica in two different summers. 

In 1954 Ball State Teachers College, along with Earlham College, 
started to sponsor the Eastern Indiana Regional Science Fair and this 
has been continued by Ball State up to the present time. Earlham de- 
cided some years ago to go into a different type of activity to inspire 
young high school students to go into the field of science. The Fair has 
been sponsored by Dr. Gerald E. Doeden and Dr. George W. Welker of 
Ball State, along with other members of the staff, including Dr. Robert 
L. Shelley, Dr. Homer D. Paschall and Dr. Leon Reynolds. 

By 1959 the requirement for the training of elementary teachers 
had been balanced in such a way that there were two courses required in 
the life sciences, two in the physical sciences, two in the earth sciences, 
two in the health sciences, and one methods course taught by persons 
in the Science Department who had had experience in the elementary 
school. This type of program is still in existence and has proved very 
successful for the training of students going out to do elementary 
work and including science in their teaching. 

In 1960 Dr. Robert H. Cooper was awarded the first annual Dr. 
James A. McClintock Award. The $500 was applied toward the publica- 
tion of a revision of Christy Woods — Outdoor Laboratories. 

In 1961 the Science Lecture and Discussion Series was held the 
first week of February and has been conducted annually up to and in- 
cluding the current year, 1968. At least five outstanding scientists have 
appeared each time on this two-day or one-day lecture series. Prominent 

History of Science 385 

persons who have appeared include Nobel prize winners and others from 
the National Severe Storms Forecast Center, the Bell Telephone Labora- 
tories of New Jersey, and the National Aeronautics and Space Adminis- 
tration of Washington, D. C. and Huntsville, Alabama. Speakers from 
medical colleges, from research departments of various colleges, and 
from industry have been included in these series over the years. This 
has proved very stimulating for students, staff, and the community. 

In 1962 A Sigma Xi Club was organized by staff members in the 
Departments of Science, Mathematics, Psychology, and Social Science 
(Anthropology). This club has been continuing its meetings and will be 
applying for Sigma Xi Society chapter status in 1968. This has been a 
stimulating addition, so far as faculty activity is concerned. 

In 1965 a complete reorganization of the departments occurred with 
the Science Department being organized into a Department of Biology, 
Department of Chemistry, Department of Geography and Geology, De- 
partment of Physics, and Department of Physiology and Health Science. 
Each department has its own department head and offers graduate de- 
grees and curricula of various types. Physics, Chemistry, Mathematics, 
and Geology moved into the new Physical Science Building in the summer 
of 1967. This five-story building was completed at a cost of $4,200,000. 

At the present time construction is progressing on the new Life 
Science Building, which will house the Departments of Biology, Physiol- 
ogy and Health Science, and Geography. The completion of this five- 
story building is anticipated late in the school year of 1969 or early 
1970 at a cost of $5,500,000. 

As of 1967 additional staff members in the different departments of 
science have made worthwhile contributions to the development of each 
science in the fields of research and teaching. Included are: 

Biology: Dr. Jerry J. Nisbet, Chairman of the Department, Dr. 
Alice H. Bennett, Dr. Frank Bernhardt, Dr. William B. Crankshaw, Dr. 
Arthur L. Eiser, Dr. Clyde W. Hibbs, Dr. Kalph D. Kirkpatrick, Dr. 
James C. List, Dr. Margaret McElhinney, Dr. Thomas R. Mertens, Dr. 
D. E. Miller, Dr. Jeanette C. Oliver, Dr. Charles E. Smith, Dr. Forrest 

F. Stevenson, Dr. George W. Welker, Dr. Charles D. Wise, Dr. Harold 
L. Zimmack. 

Chemistry: Dr. P. A. Wiseman, Head of Department, Gerald L. 
Alexander, Dr. William H. Bowman, Dr. Gerald E. Doeden, Dr. Ralph 
D. Joyner, Dr. Richard M. Lawrence, Dr. LeRoy McGrew, Dr. Robert L. 

Geography-Geology: Dr. George F. Beatty, Chairman of the De- 
partment, Dr. Lowell I. Dillon, Dr. Henry E. Kane, Edward E. Lyon, 
Dr. Harlan H. Roepke, William H. Stevenson. 

Physics: Dr. Malcom E. Hults, Head of Department, Roger D. Bur- 
gess, Dr. Edwin C. Craig, Ben Hurd, Dr. Leon M. Reynolds, Dr. Newton 

G. Sprague. 

386 Indiana Academy of Science 

Physiology and Health Science: Dr. Warren E. Schaller, Head of 
Department, Dr. William Bock, Dr. Raymond E. Henzlik, Dr. Homer D. 
Paschall, Dr. Gordon L. Rosene, Dr. Russell E. Siverly, Dr. Robert J. 

Ball State, with its 675 acres, approximately 50 buildings, about 
15,000 students, and a faculty of over 600, is considered "one of the 
emerging universities." It developed from a private school to an institute, 
to a normal school, to a teachers college, and to a university. From an 
institution with 230 students and 30 faculty members it has developed 
into a university with five colleges — Architecture and Planning, Busi- 
ness, Fine and Applied Arts, Teachers College, and College of Science 
and Humanities. The science instructional staffs now include not only 
full-time professors but also local physicians, local research workers 
and laboratory technicians and registered nurses who may serve on a 
part-time basis. 


Chairman: Edwin C. Craig, Ball State University 
Richard Conklin, Hanover College, was elected chairman for 1969. 


Preliminary Results of a Muon Energy Study. Robert Callis and Edwin 
Craig, Ball State University. — This study was conducted in order to 
determine the energy of muon particles observable through liquid scin- 
tillation detection when muon travel was limited to a short distance. 
Knowing the time delay necessary for each path of electronic signal to 
achieve coincidence, and the distance of muon flight between scintillation 
tanks, the muon velocity can be calculated. Knowing the muon velocity, 
and the mass as 206 m e , the energy of the muon can be determined. 

The coincidence system used in this study included time to height 
conversion and had a resolving time of 20 nanoseconds. With the cali- 
brated nanosecond delay module it was possible to introduce delays in 
increments of one nanosecond. The average muon velocity was calcu- 
lated to be 2.18 x 10 8 m/sec. The energy corresponding to this velocity 
was 40.5 MeV. The probability of chance coincidence occurring was 2.76 x 
10~s based upon a probability certainty of 1.00. The average number of 
chance coincidences occurring in one second was found to be 9.94 x 10- 5 

Determination of Absolute D* for Photovoltaic, Infra-red Detectors. John 
F. Houlihan, Shenango Valley Campus, Penn State University. — A 
practical and accurate method of determining an absolute value of D :: 
for photovoltaic, infra-red detectors is discussed. The D* performance 
parameter is defined and is expressed in physically measurable quantities. 
The experimental set-up is shown and discussed briefly. Also the correc- 
tion factors which are necessary due to the experimental techniques 
used, are considered in some detail and practical methods of determining 
them are given. A Fourier analysis of a square-wave pulse-train is pre- 
sented in an appendix and the affect of such a wave train on an rms 
meter considered in detail. Finally, several excellent reference sources 
for infra-red detector work are listed. 

Possible Methods for Observing Shadow Bands at the Next Solar Eclipse 
in North America. Roger D. Burgess and Malcom Hults, Ball State 
University. — Immediately preceding and following an eclipse of the sun, 
light and dark bands are often seen moving across the surface of the 
earth. A great deal more study of these so-called Shadow Bands, both 
theoretical and experimental, is needed. This paper reviews the history of 
observations of the bands and the attempts to explain them. A report 
of an observation of the bands in Rio Grande do Sul, Brazil, November 
12, 1966, is given. More advanced methods of visually observing the 
bands are suggested and attempts to detect and measure shadow bands 
electronically are reported. The presentation emphasizes the importance 


388 Indiana Academy of Science 

of both visual observation and electronic detection of shadow bands at 
the next eclipse March 7, 1970, which is readily accessable to physicists 
and astronomers in North America. 

Nuclear Electric Quadrupole Resonance Analysis of Chemical Bonds in 
CI 35 - Containing, Straight-Chain Hydrocarbons. David E. Koltenbah, 
Ball State University. — The nuclear electric quadrupole moment of nuclei 
of spin greater than or equal to unity interacts with the electric field 
gradient arising from the molecular or crystalline electric field in the 
environs of the given nucleus. The frequency of this interaction is sensi- 
tive to changes in the surrounding electron distribution arising from 
an altered chemical structure. Consequently, the nuclear quadrupole 
resonance (NQR) frequency is related to the structure of the chemical 
bond, and the study of the shift of this frequency from one member to 
another in a homologous group of compounds has been employed as a 
means of studying chemical bonds. Extensive investigations are reported 
in the literature in which this effect has been studied in halogenated ring 
and halogenated saturated straight-chain compounds. This paper re- 
viewed a systematic study of NQR frequencies of CI 35 in several chloro- 
alkenes, of which virtually no previous study had been made. The in- 
ductive effect of -CI, -CCIH2, and -CH 3 substitutents upon the electron 
density of the C-Cl bonds was determined. The effects of hybridization 
and 7r-bonding characteristics were estimated by the Townes-Dailey 
rules, and the C-Cl bond ionicities were calculated and compared with 
the ionicities of more abundantly studied chloroalkanes. 

Low Energy Elastic Scattering of K - Mesons off Protons. Gerald P. 
Thomas, Ball State University. — Forty-four K~ -p low momentum elastic 
scattering events between and 250 MeV/c have been found by following 
265.12 meters of Krmeson tracks, backward from their capture points in 
nuclear emulsion. The total scattering cross-section is plotted as a func- 
tion of kaon momentum and this is fitted with the s-wave zero-effective- 
range theory with the scattering lengths A = — 1.1 + 0.55i, Ai = — 0.20 + 
0.44 i for the isotopic spin T = O and T = 1 channel respectively. Fur- 
ther evidence for s-wave interaction is discussed through the angular 
distribution in the center-of-mass system for the elastic events. 

The Design and Construction of a System for 
Direct Measurement of Atomic Lifetimes 

Theodore V. Blanc, i Ake G. Danemar,- and David E. Koltenbah 
Department of Physics, Ball State University 


The project reported in this paper involved the design and construction 
of a system to directly measure atomic lifetimes, and follows the technique 
employed by Jules Klose of the National Bureau of Standards. A low energy 
10 to 3 eV, 20 nsec-duration, 10 kHz square wave electron excitation pulse 
was used to excite a gas at a pressure of 1 X 10-* Torr. The visible emission 
spectrum was observed by a photomultiplier through a grating spectrometer 
which permitted the selection of the desired spectral line or energy level 
transition. Employing delayed coincidence techniques familiar in nuclear 
physics, the square excitation pulse and the ensuing photomultiplier pulse 
were fed into a time-to-pulse height analyzer whose output would display 
the decay of the emission intensity as a function of the time for relaxation 
of the atomic systems. The project was completed to the construction of 
the time-to-pulse height converter and promises to be useful in obtaining 
data on atomic lifetimes. 


This project was the first study by the Department of Physics at 
Ball State University in the measurement of atomic lifetimes. For this 
initial work the project was divided into two segments. The theoretical 
computations and calculations comprised the M.S. thesis work by Mr. 
Danemar (4), while the actual design and construction of the system 
comprised the M.S. thesis work of Mr. Blanc (2). This paper is essentially 
a synopsis of the latter thesis. 

The purpose of the project was to design and construct a system cap- 
able of measuring the rate of decay of excited atomic energy levels in 
neutral atoms. The atomic excitation was achieved by bombarding a gas 
with an electron beam of sufficiently low energy so as to obtain excita- 
tion of neutral atoms rather than ionization. A short square wave pulse 
of low energy electrons was used as the primary beam. The measurement 
of the decay time of the subsequent atomic relaxation after the pulse was 
made by observation of the decay time of the intensity of the invisible 
radiation emitted by the gas as detected by a grating spectrometer. The 
desired spectral line was selected from the spectrometer and the emitted 
photons received by a photomultiplier. By analysis of the delayed coinci- 
dence between the detected photomultiplier pulses and the primary elec- 
tron pulses, the decay of the excited state could be determined. 


The method of delayed coincidence in photomultiplier scintillation 
counting in nuclear physics was first applied to direct measurement of 

1 Present address: Physics Department, University of Kentucky, Lexing- 
ton, Kentucky. 

2 Present address: Physics Department, Princeton University, Princeton, 
New Jersey. 



Indiana Academy of Science 

atomic lifetimes by Heron, McWhirter, and Rhoderick (6) about 1954 at 
the University of Glasgow. They sought to develop an accurate and 
direct method of measuring atomic lifetimes capable of determining 
whether or not the decay was of an exponential form. The technique was 
further developed by Bennett, Javan, and Ballik (1) in 1960 at the Bell 
Telephone Laboratories by using a multichannel pulse-height analyzer 
in conjunction with a pulse time-to-height converter. This method proved 
to be both enormously faster and stabler than the single-channel tech- 
nique employed by Heron, et al (6). Later this technique was employed in 
1964 by Holzberlein (7, 5) at the University of Oklahoma, as well as 
by Pendleton and Hughes (10) at the University of Arkansas, to measure 
the lifetime of helium. The same technique was employed by Klose (8, 9) 
at the National Bureau of Standards to study neon in 1965 and argon 
in 1967. Wolff and Davis (13) at the University of California used a laser 
excitation system to study cesium and sodium in 1967. 

Project Rationale 

The project was appealing to the authors for a number of reasons. 
First, up to the time of the initiation of the project there had been only 
five elements known to have been in some part studied — helium, neon, 
argon, cesium, and sodium. Thus there existed a need to extend knowl- 
edge to other elements and to attempt to expand the state of the art. 
Second, the field was relatively new and unpopulated. With the develop- 
ment of current electronics a successful technique had been acquired, 
but insofar as could be determined by the authors it had been success- 


Grid Power ■ 
• Supplies 








Figure 1. Block Diagram of Apparatus. 



fully employed by only four other research groups. Lastly, the equipment 
that would be required for the project for the most part consisted of 
multi-purpose equipment of relatively moderate cost available at most 
smaller universities. 

Experimental Method and Apparatus 

It is essentially after the method and technique of Dr. Jules Klose 
of the National Bureau of Standards that this project was patterned. 
A block diagram of the system used by the authors is given in Figure 1. 
The source of excitation was a specially designed low pressure gas excita- 
tion tube which employed a low energy electron gun. A square wave sig- 
nal from a pulse generator was applied to the control grid of the electron 
gun so as to produce a square electron excitation pulse in the gas under 
study in the tube. The spectral emission of the excited gas was separated 
into its components by a grating spectroscope. The particular energy level 
transition or spectral line to be studied was then isolated, detected by 
a photomultiplier, and subsequently amplified. The same square wave 
signal from the generator was also used to "start" a time-to-pulse-height 
converter. The pulse from the photomultiplier was used to "stop" the 
time-to-pulse-height converter. The function of the converter was to 
feed a voltage pulse proportional to the time interval between the 
"start" and "stop" into a multichannel pulse height analyzer. Thus the 
pulse height analyzer displayed the emission intensity on the ordinate 
and the emission life time on the abscissa. 

The entire system can be broken down into four basic sections: 1) 
the vacuum or low pressure gas system, 2) the spectrograph, 3) the elec- 
tronics, and 4) the excitation tube. 




I I I 

• I I Excitation Tube 
I i I 




-( mh}- 

1 (mA 


SQ. Wave 

D.C. Power 



D.C. Power 

( vtvm) 

Figure 2. Block Diagram of Excitation Tube Electronics 


Indiana Academy of Science 

The vacuum or low pressure gas system was constructed entirely of 
glass, including a Delmar DS-7050-1 glass mercury diffusion pump which 
was fused directly into the system via a liquid nitrogen cold trap. The 
gas pressure in the excitation tube was operated at approximately 
1 X 10"* Torr or O.lfx. Since the system was originally designed with a 
McLeod pressure gauge fused directly into the system and since the 
gauge was found to be a large source of mercury contamination, it was 
decided to study first the atomic lifetimes of mercury. 

The spectograph system consisted merely of a slightly modified large 
Cenco Model 87102 grating spectograph with a dispersion of 16 A /mm. 

The electronics system consisted of an HP-214A square wave pulse 
generator, an RCA-6655A photomultiplier and Victoreen PVS-C pre- 
amplifier, a time-to-pulse height converter (3), a Victoreen PIP-400A 
pulse-height analyzer, and an excitation tube control system. The last 
system, which is depicted diagrammatically in Figure 2, consisted of a 
series of meters and power supplies to enable the operator to have 
complete control and monitoring capabilities over the excitation tube. 

Lastly, the excitation tube — the very heart of the project — proved to 
be one of the most difficult parts to design and construct. The excitation 
technique employed by Klose (8, 9) was basically designed by Simpson and 
Kuyatt (12) at the National Bureau of Standards in which, by using a 
multistage device, electrons were first drawn from the cathode by a high 
potential of the order 300 V and then decelerated to obtain a beam of the 
desired energy range in the order of 10 to 30 eV. This design was suc- 
cessfully used by Klose in his studies but was found by the authors to be 
impractical to construct due to the lack of sufficiently sophisticated 
machining facilities. Thus it was decided to adapt a presently existing 
production TV electron gun to the above technique. A diagram of the 
authors' electron gun appears in Figure 3. The electron guns used in tele- 

I.Jcm Collector 

[ P.P. 0.9cm / 

•Figure 3. Diagram of Excitation Tube. 

Physics 393 

vision picture tubes are designed to operate at potentials of the order of 
500 to 5000V. Use was made of an RCA 19GWP22 color television gun 
(11) in which the operating potentials on each of the gun elements were 
proportionately scaled down to produce an electron beam energy in the 
10 to 30 eV range. This gun system had two advantages — it was designed 
to operate at comparatively low potentials and, since it was a color picture 
tube gun system, it actually consisted of three guns. Due to the fact that 
the gun was to be operated at relatively low potentials, the high potential 
grid 4 and magnetic shield were removed from the gun assembly leaving 
a three-grid electron gun. This had an advantage because the three guns 
could be operated singly, so that if one gun burned out it was possible to 
change to another gun by external switching without opening the tube. 
The collector was constructed out of stainless steel and formed into a 
"cup" shape to minimize the mechanical recoil of the low energy electrons 
back into the excitation viewing area. 

The excitation tube functioned sufficiently well for obtaining some 
initial spectrograms, but was not as stable as desired because of the 
existence of a space charge in the viewing area arising from recoil elec- 
trons from the anode. A fine mesh suppressor grid at the anode potential 
should eliminate this annoyance and allow the tube to be used for atomic 
lifetime measurements. 

Literature Cited 

1. Bennett, W. Jr., A. Javan, and E. A. Ballik. 19G0. Measurement of radia- 
tive lifetimes. Bull. Amer. Phys. Soc. 5:496. 

2. Blanc, T. V. 1968. The design and construction of a system for direct 
measurement of atomic lifetimes. Unpublished M.S. Thesis, Department 
of Physics, Ball State University. 

3. Culligan, G., and N. H. Lipman. 1960. Fast transistorized time-to-pulse 
height converter. Rev. Sci. Instr. 31:1209. 1960. 

4. Danemar, A. G. 1968. Theory of measurements of atomic lifetimes. 
Unpublished M.S. Thesis, Department of Physics, Ball State University. 

5. Fowler, R. G., T. M. Holzberlein, C. H. Jacobson, and S. J. B. Corrigan. 
1964. Direct measurements of lifetimes of excited states of neutral 
helium. Proc. Phys. Soc. 84:539. 

6. Heron, S., R. W. P. McWhirter, and E. H. Rhoderick. 195 6. Measurements 
of lifetimes of excited states of helium atoms. Proc. Roy. Soc. (London) 

7. Holzberlein, T. M 1964. Direct measurement of atomic lifetimes: 
helium. Rev. Sci. Instr. 35:1041. 

8. Klose, J. Z. 1966. Atomic lifetimes of neon I. Phys. Rev. 141:181. 

9. Klose, J. Z. 1967. Lifetimes of some 4p levels in argon I. J. Opt. Soc. 
Amer. 57:1242. 

10. Pendleton, W. R., and R. H. Hughes. 1965. Radiative lifetimes of excita- 
tion mechanics in helium. Phys. Rev. 13S:A683. 

11. RCA design information data booklet 19GWP22, 2-67. Radio Corporation 
of America, Electronic Components and Devices, Harrison, N. J. 

12. Simpson, J. E. and C. S. Kuyatt. 1963. Design of low voltage electron 
guns. Rev. Sci. Instr. 34:265. 

13. Wolff, R. J., and S. P. Davis. 1968. Direct measurement of atomic life- 
times of cesium and sodium. J. Opt. Soc. Amer. 58:490. 


Chairman: Carrolle A. Markle, Earlham College 
Jack E. Humbles, Indiana University, was elected chairman for 1969 

Other papers read 

Additional Records of Flora of Wayne County, Indiana. 

Carrolle A. Markle, Earlham College. 

Hookeriaceae Species and Distribution in South America 

Winona H. Welch, DePauw University 


This paper is the second in the series on the distribution of the 
Hookeriaceae in the world. The first and second studies pertain, especially, 
to North, Central, and South America, and to the West Indies. Apparently 
the center of distribution is in northern South America, particularly in the 
NW portion of the continent. The greatest endemism occurs in northern 
South America. North America and southern South America have the 
smallest number of species and endemics, the former having the least. 
There is evidence that the Hookeriaceae is chiefly a tropical family, in the 
Americas, although species occur from Alaska to the southern tip of South 
America. East-West as well as North-South distribution are shown by 
species of the Hookeriaceae. Hookeria acutifolia occurs in Hawaii, Canada, 
United States, Mexico, Central America, South America, West Indies, and 
Asia. Other species occur in South America, Australia, Tasmania, New 
Zealand, and on islands near South Africa. 

This paper is the second in the series on the distribution of the 
Hookeriaceae, based largely upon epithets and distribution areas cited in 
Index Muscorum. It is assumed that the introductory pages of the first 
paper (9) will be reviewed before reading this report. 

The following islands have been included with South America: 
Hermite, Falkland, Galapagos, Trinidad, and Tobago. The area is that of 
the Index Muscorum. 

For reference, the areas in the first paper are repeated: America 1. 
North America (Canada, United States, Mexico), Greenland, Aleutian 
Islands, Bermudas; America 2. Central America and Cocos Island; Amer- 
ica 3. West Indian Islands (Greater and Lesser Antilles, Bahamas) except 
Trinidad and Tobago. The three distribution divisions of South America 
numerically follow those of North and Central America: 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. 

As assumed in the beginning of this project, as monographic work 
progresses in the family and as collectors publish their papers on new 
records of species, often extending the distribution range, changes are 
necessary to keep the information up-to-date. Differences since the first 
paper (9) follow. 

Actinodontiiim portoricense Crum & Steere, Am 3, 4, not Am 3 
endemic; A. sprucei (Mitt.) Jaeg., Am 2, 4. Callicostella grossiretis Bartr., 
Am 2 endemic; C. subpallida Ren. & Card., Am 3 endemic. Crossomitrium 
herminieri (Besch.) Jaeg., Am 3 endemic instead of Am 2; C. oerstedi- 
anum C. Muell., Am 2, 4, not Am 2 endemic; C. orbiculatum C. Muell., Am 
3, 5, not Am 3 endemic. Hookeriopsis diffusa (Mitt.) Jaeg., Am 2, 4, not 
Am 2 endemic; H. guadalupensis (Brid.) Jaeg., Am 2, 3, not Am 3 
endemic; H. websteri Crum & Bartr., Am 3 endemic. Hypnella cymbifolia 
(Hampe) Jaeg., Am 3, 4, 5, not Am 3 endemic; H. jamesii H. Robins., 


Plant Taxonomy 397 

Am 2 endemic. Lepidopilum apollinairei Broth. & Paris, Am 2, 4, not 
Am 2 endemic; L. tortifolium Mitt., Am 1, 2, 3, 4. Thamniopsis pendula 
(Hook.) Fleisch., Am 2, 4. Species, varieties, and forms reduced to 
synonymy: Crossomitrium heterodontium Ren. & Card., Am 2 = C. 
patrisiae (Brid.) C. Muell.; Cyclodictyon blandum (Lor.) Kuntze, Am 3 = 
C. varians (Sull.) Kuntze; Lepidopilum cubense (Sull.) Mitt. f. integri- 
folia Ther., f. latifolia Ther., and f. robusta Ther., Am 3 = L. cubense 
(Sull.) Mitt.; L. genuflexum C. Muell., Am 2 = Rhynchostegiopsis flexuosa 
(Sull.) C. Muell.; L. polytrichoides (Hedw.) Brid. var. pellucens Besch. 
(not var. pallescens) = L. polytrichoides. 

Three genera, Callicostellopsis, Helicoblepharum, and Stenodesmus, 
occur in South America, only. Callicostellopsis is a genus composed of 
one species, meridiensis, which has been recorded only in NW South 
America. The genus, Helicoblepharum, consists of four species, all occur- 
ring in northern South America, one in the NE and three in the NW. 
Two species, latifolius and tenuicuspis, comprise the genus, Stenodesmus, 
and are known only from NW South America. 

Two genera, Amblytropis and Philophyllum, are chiefly, but not 
totally, South American genera. One of the five species of Amblytropis, 
A. denticulata, is endemic in the West Indies. The other four species, 
A. gemmacea, A. hispidula, A. ovata, and A. setosa, have been reported 
only from NW South America. The genus, Philophyllum, is composed of 
four species, all of which occur in NE South America. One species, 
P. tenuifolium, has a more general distribution, occurring in Central 
America and northern South America. 

In the lists of Hookeriaceae in Am 4, 5, 6, the asterisk indicates the 
endemic species, varieties, and forms. These plants presently are not 
known to occur in geographical areas other than the ones cited. The 
species of additional areas follow. 

Steere (4) stated that many species of mosses of the eastern slopes 
of the Andes and the headwaters of the Amazon extend from eastern 
Peru to the West Indies, and that the Andean highlands of the sierra are 
much wetter in the north and are most closely related to the wet montane 
and alpine forests of Colombia and Central America. Steere also points 
to the relationship between the genera of Ecuador, Peru, and Bolivia, and 
those of Mexico. 

In the distribution patterns of the Hookeriaceae species, the following 
have been noted by the author. 

Callicostella pallida has been recorded from northern South America, 
Central America, Mexico, southern United States, and the West Indies. 

Lepidopilum polytrichoides occurs throughout South America, in 
Central America, Mexico, Florida, and the West Indies. 

Lepidopilum scabrisetum has been reported from the three distribu- 
tion areas of South America, from Central America, Mexico, and the 
West Indies. 

398 Indiana Academy of Science 

Adelothecium bogotense, Crossomitrium patrisiae, Cyclodictyon 
albicans, Hookeriopsis cruegeriana, Isodrepanium lentulum, and 
Lepidopilum radicale have been recorded from northern South America, 
Central America, Mexico, and the West Indies. 

Cyclodictyon roridum, Daltonia longifolia, and Lepidopilum torti- 
folium occur in NW South America, Central America, Mexico, and the 
West Indies. 

Daltonia gracilis is known from three distribution areas of South 
America, Central America, and from Mexico. 

The distribution ranges of Cyclodictyon rubrisetum, Lepidopilum 
brevipes, and L. carneum follow the Cordilleran pattern, in NW South 
America, Central America, and Mexico. 

Callicostella scabriseta and Hookeriopsis incurva may be collected 
throughout South America, in Central America, and in the West Indies. 

The following species have been cited from northern South America, 
Central America, and the West Indies: Cyclodictyon cuspidatum, 
Daltonia stenophylla, Hemiragis aurea, and Hookeriopsis undata. 

Hookeriopsis falcata and Leskeodon andicola occur in NW South 
America, Central America, and the West Indies. 

The distribution range for Callicostella depressa, Hookeriopsis 
acicularis, and Leskeodon cubensis is NE South America, West Indies, 
and Central America. 

Hookeriopsis crispa, H. variabilis, Hypnella pilifera, Neohypnella 
chrysophyllopodia, Philophyllum tenuifolium, and Thamniopsis pendula 
have been recorded in northern South America and in Central America. 

Crossomitrium oerstedianum, C. wallisii, Daltonia lindigiana, D. 
tenuifolia, Hookeriopsis diffusa, H. subfalcata, Lepidopilum apollinairei, 
L. semi-laeve, Leskeodon pusillus, and Thamniopsis pendula are species 
of NW South America and Central America. 

Hypnella cymbi folia, H. diversifolia, H. leptorrhyncha, and 
Lepidopilum intermedium are species in the distribution area of northern 
South America and the West Indies. 

The following species have been collected in NW South America and 
the West Indies: Actinodontium portoricense, Callicostella colombica, C. 
rivularis, C. subpallida, Cyclodictyon denticulatum, C. lindigianum, C. 
ulophyllum, Lepidopilum aureo-fulvum, L. mitellcri, and L. robustum. 

Species reported from NE South America and the West Indies are: 
Crossomitrium cruegeri, C. orbiculatum, Cyclodictyon olfersianum, Dis- 
tichophyllum cubense, Lepidopilidium portoricense, Lepidopilum subauri- 
folium, Leskeodon auratus, and L. cubensis. 

The following species are known, presently, from northern South 
America only: Callicostella aspera, C. martiana, C. merkelii, C. micro- 
carpa, C. paulensis, C. rufescens, Cyclodictyon limbatum, C. regnellii, 
Hookeria viridula, Hookeriopsis asprella, H. hypnacea, H. parkeriana, 
Hypnella pallescens, Lepidopilidium divaricatum, Lepidopilum affine, L. 

Plant Taxonomy 399 

ambiguum, L. biductulosum, L. nanothecium, L. sub fie xi folium, and 
Thamniopsis killipii. 

Three South American species are known only from distribution areas 
four and six: Callicostella scabriuscula, Daltonia trachyodonta, Hookeria 
lorentzii, H. uliginosa, Lepidopilum erectiusculum, and Pterygophyllum 

Lepidopilum plebejum has been collected in S and NE South 

Hookeria acutifolia has been recorded from islands of the Pacific or 
in Oceania (Hawaii), in Canada (British Columbia), the United States 
(from Washington to Alabama, Georgia, and Connecticut), in Mexico, 
Central America (Guatemala, Costa Rica), South America (Ecuador, 
Bolivia, Brazil), West Indies (Cuba, Jamaica, Haiti, Porto Rico, Guade- 
loupe), and in Asia (Japan, India, Nepal, Ceylon, North and Central 
Vietnam, Sumatra, Java). It is evident that this species shows an exten- 
sive north and south as well as east and west distribution range. 
However, H. acutifolia may not be regarded as a cosmopolitan species. 

The following species have east-west or lateral distribution instead of 
north-south or vertical. The method of distribution may be clarified as 
additional localities are discovered. Distichophyllum assimile and 
Pterygophyllum obscurum have been collected in southern South America, 
in Australia, and Tasmania. 

Distichophyllum rotundifolium, Eriopus apiculatus, Pterygophyllum 
dentatum, Sauloma tenella, and S. tenella f. propagulifera have been 
reported from southern South America, Australia, Tasmania, and New 

Eriopus flexicollis has been collected in southern South America and 
New Zealand. 

Eriopus cristatus is known from southern South America, Oceania, 
Australia, Tasmania, New Zealand, Madagascar, Maritius, and Reunion. 

It would seem that this east-west distribution has been due to conti- 
nental drift, using Wegener's explanation (6); that is "to the process of 
relative movements among continents." Wegener (5) assumed that all 
the continents had been united, comprising adjacent parts of a large 
supercontinent. The latter broke up into the present continents which 
drifted apart. Since fossil remains of identical animals and plants have 
been found in the widely separate continents of Africa and South 
America, it seems plausible that there can be like species of mosses living 
today on these distant continents. 1 

Creer (2) proposed that the southern hemisphere supercontinent 
started to break up about 150-200 million years ago, to form the conti- 
nents of South America, Africa, Australia, and Antarctica; and the 
subcontinents of India and Madagascar. 

1 The author is indebted to Dr. James A. Madison, Professor of Geology, 
DePauw University, for the helpful geological references. 

400 Indiana Academy of Science 

Dietz (3) states that "most paleontologists claim that it is easier to 
explain the occurrence of similar species on continents widely separated 
by the ocean on the basis of transfer of living creatures by rafting on 
flotsam, by easy moves along ancient island chains, or by long subsided 
land bridges, rather than by continental drift." The paleontologists also 
explain that a "supposedly simpler method than continental drift, would 
be dispersion of living creatures over isthmian links (like the Isthmus of 
Panama today), which included narrow land bridges between Africa and 
South America, for instance." 

Axelrod (1) reviewed the paleontological evidence for the late Paleo- 
zoic and Mesozoic, and concluded that the fossil floras suggested stable, 
not drifting continents. One of his conclusions was that the vegetation- 
climatic zones display a symmetrical arrangement from northern to 
southern hemispheres consistent with continental stability. 

Whichever theory one accepts, the evidence of these species of the 
Hookeriaceae on such distant land bodies is a challenge to a bryologist. 

America 4: Venezuela, Colombia, Peru, Bolivia, Ecuador, 
Galapagos Islands 

Actinodontium portoricense Crum & Steere, A. sprucei (Mitt.) Jaeg. 
Adelothecium bogotense (Hampe) Mitt. *Amblytropis gemmacea (Mitt.) 
Broth., *A. hispidula (Mitt.) Broth., *A. ovata (Mitt.) Broth., *A. setosa 
(Mitt.) Broth. *Callicostella acutifolia Ther., C. aspera (Mitt.) Jaeg., 
C. colombica Williams, *C. galipanoana (C. Muell.) Broth., *C integri- 
folia (C. Muell.) Broth., C. martiana (Hornsch.) Jaeg., C. merkelii 
(Hornsch.) Jaeg., C. microcarpa Aongstr., C. pallida (Hornsch.) Aongstr., 
C. paulensis Broth., *C. plicatula Ther., C. rivularis (Mitt.) Jaeg., 
C. rufescens (Mitt.) Jaeg., *C. saxatilis (Mitt.) Jaeg., *C. scabripes (C. 
Muell.) Broth., C. scabriseta (Hook.) Jaeg., C. scabriuscula (C. Muell.) 
Jaeg., C. strumulosa (Hampe & Lor.) Jaeg., C. subpallida Ren. & Card. 
*Callicostellopsis meridensis (C. Muell.) Broth. *Crossomitrium epiphyl- 
lum (Mitt.) C. Muell., *C. goebelii C. Muell., C. oerstedianum C. Muell., 
C. patrisiae (Brid.) C. Muell., *C. phragmidiaceum C. Muell., *C. 
rotundi folium Herz., *C. saprophyllwni Broth., *C splendens Broth., *C. 
spruceanum C. Muell., *C. tenellum C. Muell., C. wallisii C. Muell. 
*Cyclodictyon aeruginosum (Mitt.) Kuntze, C. albicans (Hedw.) Kuntze, 
*C allionii Broth., *C. amnigenum (C. Muell.) Broth., *C. angustirete 
Herz., *C. bakeri (Britt.) Par., *C benoistii Ther., *C. bombonasicum 
(Mitt.) Kuntze, *C. breve Herz., *C. caespitosum (Mitt.) Kuntze, *C. 
capillatum (Mitt.) Kuntze, *C. castaneum (Mitt.) Kuntze, * C. 
chimborazense (Mitt.) Kuntze, C. cuspidatum Kuntze, C. denticulatum 
Kuntze, *C. fendleri (C. Muell.) Broth., *C. flexieuspis Broth., *C humile 
(Mitt.) Kuntze, *C. jagianum (C. Muell.) Kuntze, *C. hraiiseanum 
(Hampe & Lor.) Kuntze, *C latifolium Kuntze, C. limbatiim (Hampe) 
Kuntze, C. lindigianum (Hampe) Kuntze, *C. mittenii (Jaeg.) Kuntze, 
*C. nivale (C. Muell.) Kuntze, *C. ob s cur i folium (Mitt.) Kuntze, *C. ob- 
scurum Herz., *C. pandurif olium (Mitt.) Kuntze, *C. plicatulum (C. 
Muell.) Broth., *C.pusillum Herz., C.regnellii (Aongstr.) Kuntze, C. rori- 

Plant Taxonomy 401 

dum (Hampe) Kuntze, C. rubrisetum (Mitt.) Kuntze, *C. rugulosum 

(Mitt.) Kuntze, *C. shillicaiense (Mitt.) Kuntze, *C. stephanii Herz., 

*C. tocoraniense Herz., C. ulophyllum (Besch.) Broth. *Daltonia bilim- 

bata Hampe, *D. brevinervis Bartr., *Z>. cucullata Hampe, D. gracilis 

Mitt., *D. jamesonii Tayl., *D. jamesonii var. laevis Herz., *D. latolim- 

bata Broth, m Herz., D. lindigiana Hampe, D. longifolia Tayl., *Z>. 

macrotheca Mitt., *Z>. ovah's Tayl., *D. pellucida Herz., *£>. peruviana 

Mitt., *D. pulvinata Mitt., £>. stenophylla Mitt., D. tenuifolia Mitt., 

D. trachydonta Mitt. *Distichophyllum elongatum Mitt. * Eriopus 

deflexus C. MuelL, *2£. mniadelphus Spruce, *£ r . nutans (Hampe) Mitt., 

*£'. papillatus Herz. *Helicoblepharu?n daltoniaceum (Hampe) Broth., 

*i?. fuscidulum (Mitt.) Broth., *#. venustum (Mitt.) Broth. Hemiragis 

aurea (Brid.) Ren. & Card. Hookeria acutifolia Hook. & Grev., H. 

lorentzii C. Muell., *H. orbignyana Mont., H. uliginosa C. MuelL, H. 

viridula Mitt. *Hookeriopsis acuminata (Mitt.) Jaeg., *H. adunca 

(Mitt.) Jaeg., *H. armata Broth., H. asprella (Hampe) Broth., *H. 

brunneophylla (C. Muell.) Fleisch., *H. cavifolia (Mitt.) Jaeg., H. crispa 

(C. Muell.) Jaeg., H. cruegeriana (C. Muell.) Jaeg., *H. curvifolia 

(Mitt.) Jaeg., *H. cuspidata Jaeg., *H. cuspidatissima (Hampe) Broth., 

H. diffusa (Mitt.) Jaeg., *H. exigua (Mitt.) Jaeg., H. falcata (Hook.) 

Jaeg., *H. glandulifera (Hampe) Jaeg., *H. gracilis (Mitt.) Jaeg., 

H. hypnacea (C. Muell.) Jaeg., H. incurva (Hornsch.) Broth., *H. 

lepidopiloides Herz., *H. longiseta Williams, *H. pachydictyon Herz., 

*H. papillidioides (C. Muell.) Broth., H. parkeriana (Hook. & Grev.) 

Jaeg., *H. pernutans (C. Muell.) Broth., *H. plumicaulis (C. Muell.) 

Broth., *H. ptari-tepuiensis Bartr., *H. purpureophylla (Britt.) Broth., 

*H. scabrella (Mitt.) Jaeg., *if. sinuata (Mitt.) Jaeg., *H. steyermarkii 

Bartr., H. sub falcata (Hampe) Jaeg., *H. subscabrella Fleisch. ex 

Broth., * H. subsecunda (Mitt.) Jaeg., *H. taylorii (C. Muell.) Wijk & 

Marg., *H. tenuis (Mitt.) Jaeg., *H. terrestris (Mitt.) Jaeg., H. 

undata (Hedw.) Jaeg., *H. undatula (C. Muell.) Broth., *H. vaga 

(Mitt.) Jaeg., H. variabilis (Mitt.) Jaeg., *H. velutina (Hampe) Jaeg., 

*H. viridissima (Mitt.) Jaeg., *H. williamsii Herz. *Hypnella brotheri 

Herz., H. cymbifolia (Hampe) Jaeg., H. diversifolia (Mitt.) Jaeg., H. 

pallescens (Hook.) Jaeg., *H. philonotula (C. Muell.) Kindb., H. pilifera 

(Hook. & Wils.) Jaeg., *H. recurvula (C. Muell.) Broth., *H. 

sigmatelloides (C. Muell.) Broth. Isodrepanium lentulum (Wils.) Broth. 

Lepidopilidium divaricatum (Doz. & Molk.) Broth., *L. purpurissatum 

(C. Muell.) Broth., *L. synoicum Herz., *Lepidopilum acutmn Mitt., 

L. affine C. Muell., *L. allionii Broth., L. ambiguiim Broth., *L. anceps 

Mitt., ;i: L. angustifrons Hampe, L. apollinairei Broth. & Par., *L. 

arcuatum Mitt., *L. argutidens Broth., *L. armatum Mitt., *L. aubertii 

Ther., L. aureofulvum C. Muell., *L. awriculatum Herz., *L. aurifolium 

Mitt., *L. ballivianii Herz., L. biductulosum (P. Beauv.) Wijk & Marg., 

*L. br achy phy Hum Broth, in Herz., *L. brevifolium Mitt., L. brevipes 

Mitt., :!: L. calvum Mitt., L. carneum Bartr, *L. caudatum C. Muell., *L. 

caviuscidum Mitt., *L. chloroneuron (Tayl.) Hampe & Lor. *L. coruvallium 

(Brid.) Mitt., *L. crispum Herz. *L. curvifolium Mitt., *L. curvirameum 

(C. Muell.) Par., *L. cuspidans Mitt., L. erectiusculum (Tayl.) Mitt., L. 

erubescens C Muell., *L. excelsum C. Muell., *L. filosum Herz., *L. 

402 Indiana Academy of Science 

frondosum Mitt., *L. gertrudis Herz., *L. goniothecium C. Muell., *L. 
gracile Mitt., *L. herzogii Broth, in Herz., *L. huallagense Broth., *L. 
imbricatif olium Mitt., *L. inflexum Mitt., *L. integerrimum Mitt., L. 
intermedium (C. Muell.) Mitt., *L. krauseanum C. Muell., *L. 
leiomitrium C. Muell., *L. leucomioides Broth., *L. longifolium Hampe, 
:|: L. maculatum C. Muell., *L. malachiticum Herz. *L. mniaceum C. 
Muell., *L. mnioides C. Muell., L. muelleri (Hampe) Spruce, L. 
nanothecium C. Muell., *L. nudum Mitt., *L. ovatifolium Herz., *L. 
pallido-nitens (C. Muell.) Paris, *L. pectination Mitt., *L. per gracile C. 
Muell., *L. perlaxum Ther., *L. permarginatum Williams, *L. phyllophilum 
Broth., L. poly tricho ides (Hedw.) Brid., *L. pumilum Mitt., L. radicale 
Mitt., L. robustum Mitt., L. scabrisetum (Schwaegr.) Steere, L. semi-la eve 
Mitt., *L. spendens Broth., *L. steyermarkii Bartr., *L. stillicidiorum 
Mitt., L. sub fie xi folium C. Muell., *L. subgracile Broth., *L. subpoly- 
trichoides C. Muell., *L. tenuifolium Mitt., *L. tenuissimum Herz., L. 
tortifolium Mitt., *L. wallisii C. Muell. Leskeodon andicola Spruce ex 
Mitt., *L. palmarum (Mitt.) Broth., *L. pungens (Mitt.) Broth., L. 
pusillus (Mitt.) Broth., *L. wallisii (C. Muell.) Broth. Neohynella 
chrysophyllopodia (C. Muell.) Bartr. Philophyllum tenuifolium (Mitt.) 
Broth. Pterygophyllum rigidum (Schwaegr.) Brid. *Rhynchostegiopsis 
complanata C. Muell., *R. tunguraguana (Mitt.) Broth. *Stenodesmus 
latifolius Bartr. & Herz., *S. tenuicuspis (Mitt.) Jaeg. *Stenodictyon 
nitidum (Mitt.) Jaeg., *S. saxicola Williams. Thamniopsis killipii (Wil- 
liams) Williams, T. pendula (Hook.) Fleisch. 

America 5 : Brazil, Paraguay, Guiana, Trinidad, Tobago. 

Adelothecium bogotense (Hampe) Mitt. *Callicostella apophysata 
(Hampe) Jaeg., C. aspera (Mitt.) Jaeg., *C. circinata (Broth.) Broth., 
*C. cruegeri (C. Muell.) Broth., *C. daltoniaecarpa (C. Muell.) Broth., 
C. depressa (Hedw.) Jaeg., *C. diatomophila (C. Muell.) Fleisch., *C. 
glabrata Broth., *C. irrorata (C. Muell.) Broth., *C. jungermannioides 
Herz., *C. juruensis Broth., *C liynosa (Broth.) Broth., *C. lorifolia 
(Hampe) Jaeg., C. martiana (Hornsch.) Jaeg., C. merkelii (Hornsch.) 
Jaeg., C. microcarpa Aongstr., *C mollis (Wils.) Jaeg., *C mono f aria 
(Geh. & Hampe) Broth., *C. mosenii (Broth.) Broth., C. pallida 
(Hornsch.) Aongstr., *C. paludicola Broth., C. paulensis Broth., *C. 
pellucida (Mitt.) Jaeg., *C. perpallida (Broth.) Broth., *C. pilotrichidio- 
ides Broth., C. rufescens (Mitt.) Jaeg., *C. rufescens var. demerarae 
Richs., *C. scaberrima Broth., C. scabriseta (Hook.) Jaeg., *C sellowiana 
(Hampe) Jaeg., *C. spur io -pallida (Broth.) Broth., *C. subdeprcssa 
(Besch.) Broth., *C. submicrocarpa (Geh. & Hampe) Broth., *C. S2t6- 
monofaria Broth., *C. torrentium (Broth.) Broth., *Chaetephora perrinii 
(Spreng.) Brid., Crossomitrium cruegeri C. Muell., C. patrisiae (Brid.) 
C. Muell., *C. paulense Broth. & Sebille, *C. radulae forme C. Muell., *C. 
ramulicola C. Muell., *C. sellowii C. Muell., *C. split gerberi (Mont.) C. 
Muell., *C. ^Zei C. Muell. *Cyclodictyon aciculif olium (C. Muell.) Broth., 
*C. albatum (C. Muell.) Kuntze, C. albicans (Hedw.) Kuntze, *C, 
chloroleucum (Broth.) Broth., C. cuspidatum Kuntze, *C. glareosum 
(Broth.) Broth., *C glaucif olium (C. Muell.) Broth., *C iporangeanum 

Plant Taxonomy 403 

(Geh. & Hampe) Broth., *C. laxifolium Herz., *C leucomitrium (C. 
Muell.) Broth., C. limbatum, (Hampe) Kuntze, *C. longifrons (Broth.) 
Broth., *C. marginatum (Hook. & Wils.) Kuntze, *C. minarum 
(Aongstr.) Kuntze, *C minor (Aongstr.) Kuntze, ::: C. molliculum 
(Broth.) Broth., C. olfersianum (Hornsch.) Kuntze, *C. pallens (Mitt.) 
Kuntze, *C. pergracile Broth., *C. regnellianum (C. Muell.) Fleisch., 
C regnellii (Aongstr.) Kuntze, *C. rivale (C. Muell.) Broth., *C. 
submarginatum (Aongstr.) Kuntze, *C. viridissimum Kuntze. *Daltonia 
androgyna Geh. & Hampe, *D. aristata Geh. & Hampe, *D. brasiliensis 
Mitt., D. gracilis Mitt., D. stenophylla Mitt. *Distichophyllum densirete 
Broth., *D. gracile Aongstr., *D. minutum C. Muell., *D. minutum var. 
perlimbatum C. Muell. *Eriopus albescens (Hampe) Jaeg., *Z£. flexicaulis 
(Hampe) Paris, *E. lorifolius (Hampe) Paris, *E. monilidontius 
(Hampe) Paris, *E. setigerus Mitt. *Helicoblepharum brasiliense 
Herz. Hemiragis aurea (Brid.) Ren. & Card. Hookeria acutifolia 
Hook. & Grev., *H. com?nutata Paris, *H. janeirensis Paris, H. 
virdula Mitt. Hookeriopsis acicularis (Mitt.) Jaeg., H. asprella 
(Hampe) Broth., *H. beyrichiana (Hampe) Broth., *//. brachypelma 
(C. Muell.) Broth., *H. caldensis (Aongstr.) Broth., *H. cirrhosa (Hampe) 
Jaeg., *H. corcovadensis (Reichdt.) Jaeg., *H. crispa (C. Muell.) 
Jaeg., H. cruegeriana (C. Muell.) Jaeg., *H. cruegeriana var. dimorpha 
(C. Muell.) Jaeg., *H. drepanophylla (Geh. & Hampe) Broth., *//. 
exesa (C. Muell.) Broth., *H. fliiminensis (Geh. & Hampe) Broth., *H 
glaziovii (Hampe) Jaeg., *H. hornschuehiana (Jaeg.) Broth., *H. 
hydrophila (C. Muell.) Broth., H. hypnacea (C. Muell.) Jaeg., H. in- 
curva (Hornsch.) Broth., *H. latifrondea (C. Muell.) Broth., *H. leu- 
comioides (Broth.) Broth., *H. lonchopelma (C. Muell.) Broth., *H. 
luteo-viridis (Besch.) Broth., *H. minutiretis (C. Muell.) Broth., *H. 
negrensis Broth, ex Ther., H. parkeriana (Hook. & Grev.) Jaeg., *H. 
perfulva (C. Muell.) Fleisch., *H. planiuscida (Hampe) Jaeg., *H. 
puiggarii (Geh. & Hampe) Broth., *H. rhynchostegioides (Broth.) Broth., 
*H. rubens (C. Muell.) Broth., *H. saprophila (Broth.) Broth., *H. 
saprophila var. major C. Muell., *H. schiffneri Broth., *H. serrata 
(Aongstr.) Jaeg., *iJ. subaurescens (Geh. & Hampe) Broth., *H. tenera 
(Hampe) Jaeg., H. undata (Hedw.) Jaeg., H. variabilis (Mitt.) Jaeg., 
*H. vesicularia (C. Muell.) Broth. Hypnella cymbifolia (Hampe) Jaeg., 
H. diversifolia (Mitt.) Jaeg., H. leptorrhyncha (Hook. & Grev.) Jaeg., 
H. pallescens (Hook.) Jaeg., H. pilifera (Hook. & Wils.) Jaeg., *H. 
punctata Broth., *H. verrucosa (Hampe) Jaeg. Isodrepanium lentulum 
(Wils.) Britt. *Lepidopilidium aureo-purpure?n (Geh. & Hampe) Broth., 
*L. brevisetmn (Hampe) Broth., *L. brevisetum var. purpurascens 
Broth., *L. caudicaule (C. Muell.) Broth., L. divarication (Doz. & 
Molk.) Broth., *L. entodontella (Broth.) Broth., *L. gracilifrons (C. 
Muell.) Broth., *L. laevisetum (Hampe) Broth., *L. nitens (Hornsch.) 
Broth., *L. nitens var. latior Geh. & Hampe, L. portoricense (C. Muell.) 
Crum & Steere, *L. rupestre (C. Muell.) Broth., *L. tenwisetum (C. 
Muell.) Broth., *L wainioi (Broth.) Broth. Lepidopihim affine C. Muell., 
L. ambiguum Broth., L. biductidosum (P. Beauv.) Wijk & Marg., *L. 
flavescens Geh. & Hampe, *L. glaziovii Hampe, L. intermedium (C. 
Muell.) Mitt., *L. latifolium (C. Muell.) Mitt., *L. laxirete C. Muell., 

404 Indiana Academy of Science 

*L. leptoloma Broth., *L. michelianum Broth. & Par., *L. mittenii C. 
Muell., *L. mosenii Broth., L. nanothecium C. Muell., *L. oblongifolium 
Mitt., *L. obtusulum C. Muell., *L. ovalifolium (Dub.) Broth., *L. plebe- 
jum C. Muell., L. poly tricho ides (Hedw.) Brid., *L. pycnodictyum C. 
Muell., L. radicale Mitt., *L. rupestre Broth., L. scabrisetum (Schwaegr.) 
Steere, L. subaurifolium Geh. & Hampe, L. subflexifolium C. Muell., *L. 
subfuscum Mitt., *L. subobtusulum Broth., *L. subsubulatum Geh. & 
Hampe, *L. subulatum Mitt., *L. surinamense C. Muell. *Leskeodon art- 
status (Geh. & Hampe) Broth., *L. aristatus var. tenuilimbatus Broth., 
L. auratus (C. Muell.) Broth., L. cubensis (Mitt.) Ther., *L. densiretis 
(Broth.) Broth., *L. longicaulis Broth., *L. minusculus (C. Muell.) 
Fleisch. Neohypnella chrysophyllopodia (C. Muell.) Bartr. *Philophyllum 
bromeliophilum C. Muell., P. tenuifolium (Mitt.) Broth. *Rhynchostegi- 
opsis brasiliensis Broth. Thamniopsis killipii (Williams) Bartr., T. 
pendula (Hook.) Fleisch. 

America 6: Chile, Argentina, Uruguay, Falkland Islands, Hermite Island. 

Callicostella scabriseta (Hook.) Jaeg., C. scabriuscula (C. Muell.) 
Jaeg. Crossomitrium patrisiae (Brid.) C. Muell. *Cyclodictyon sublim- 
batum (C. Muell.) Kuntze. Daltonia gracilis Mitt., D. trachyodonta 
Mitt. Distichophyllum assimile Broth, in Skottsb., *D. cavifolium (Card.) 
Card., *Z>. dicksonii (Hook. & Grev.) Mitt., *D. ellipticum Herz., *D. 
eremitae (Jaeg.) Paris, *D. fernandezianum Broth, in Skottsb., *Z). flac- 
cidum (Hook. f. & Wils.) Mitt., *D. nanospathulatum Herz., *D. nigri- 
cans Besch., *Z). patagonicum Besch., D. rotundif olium (Hook. f. & 
Wils.) C. Muell. & Broth., *D. subelimbatum Broth, in Skottsb. Eriopus 
apiculatus (Hook. f. & Wils.) Mitt., *£". apiculatus var. platyloma Card. 
& Broth., E. cristatus (Hedw.) Brid., E. flexicollis (Mitt.) Jaeg., *E. 
grandiretis Broth, in Skottsb., *E. leptoloma Broth, in Skottsb. Hookeria 
lorentzii C. Muell., *H. magellanica (P. Beauv.) Arnott, H. uliginosa C. 
Muell. Hookeriopsis incurva (Hornsch.) Broth. *Lamprophyllum splen- 
dissimum (Mont.) Broth. *Lepidopilum aurescens C. Muell., L. erecti- 
usculum (Tayl.) Mitt., L. plebejum C. Muell., L. polytrichoides (Hedw.) 
Brid., L. scabrisetum (Schwaegr.) Steere. *Pterygophyllum anomalum 
(Schwaegr.) Mitt., P. anomalum var. pallidum Card. & Broth., *P. chono- 
ticum Mitt., P. dentatum (Hook. f. & Wils.) Dix., *P. fragile Mitt., *F. 
magellanicum Besch., P. obscurum Mitt., *F. obscurum f. thermalis Herz., 
P. rigidum (Schwaegr.) Brid., *P. tenuinerve Broth, in Skottsb. Sauloma 
tenella (Hook. f. & Wils.) Mitt., S. tenella f. propagulifera Sainsb. 

A summary of the Hookeriaceae species, varieties, and forms, and 
the endemics, based upon data presently known, in North, Central, and 
South America follows: Am 1: 36 species, 8 of which are endemic; Am 
2: 96 species, 39 of which are endemic; Am 3: 107 species, 54 of which 
are endemic; Am 4: 255 species, 173 of which are endemic; Am 5: 192 
species, 137 of which are endemic; Am 6: 47 species, 23 of which are 
endemic. The greatest endemism occurs in northern South America as 
shown by a total of 310 endemics. The largest number of species and 
endemics occurs in Am 4 or northwestern portion of South America and 
decreases to the east, north, and south. North America and southern 

Plant Taxonomy 405 

South America have the smallest number of species and endemics, the 
former having the least. It is also evident that the family Hookeriaceae 
is chiefly a family of tropical mosses in the Americas, although species 
occur from Alaska to the southern tip of South America, Hermite Island, 
and the Falkland Islands. 

Literature Cited 

1. Axelrod, D. I. 1963. Fossil Floras suggest stable, not drifting Continents. 
J. Geophys. Res. 68:3257-3264. 

2. Creer, Kenneth M. 1966. Continents on the Move. Sea Frontiers 12(3): 

3. Dietz, Robert S. 1967. More about Continental Drift. Sea Frontiers 13(2): 

4. Steere, William C. 1948. Mosses of Ecuador I. The Bryologist 51(3): 

5. Wegener, A. 1912. Die Entstehung der Kontinente. Geol. Rundsch. 3(4): 


6. Wegener, A. 1924. The Origin of Continents and Oceans. (English trans- 
lation by J. G. A. Skerl. Methuen, London.) 212 p. 

7. Welch, Winona H. 1962. The Hookeriaceae of the United States and 
Canada. The Bryologist 65(l):l-24. 


-. 1966. The Hookeriaceae of Mexico. The Bryologist 60(1): 

9. - — — ■. 1968. Hookeriaceae Species and Distribution in North and 

Central America and West Indies. Proc. Indiana Acad. Sci. 77:351-356. 

10. . 1969. The Hookeriaceae of Cuba. The Bryologist 72(2). In 


11. Wijk, R. van der, W. D. Margadant, and P. A. Florschutz. 1959, 1962, 
1964, 1967. Index Muscorum. Vol. 1-4. Utrecht, The Netherlands. 

Some Late Glacial Charophytes Compared to Modern Species 

Fay Kenoyer Daily, Butler University 


Some Date Glacial Charophytes Compared to Modern Species. Fay 
Kenoyer Daily, Butler University. — Fossil fructifications from New York pro- 
vided by Norton G. Miller were referred to the moden taxa, Chara sejuncta 
and Tolypella glowierata. This Tolypella displayed a triply-segmented basal 
plate on the oospore while a Tolypella (undoubtedly T. prolifera) reported 
in an earlier study had an undivided basal plate. In view of these findings 
and the conflicting literature on this subject, some selected modern speci- 
mens were studied. It was found that Tolypellas of the Section Conoideae to 
which T. prolifera belongs had an undivided basal plate similar to that in 
the Chareae. Members of the Section Allantoideae represented by T. glom- 
erata had doubly or triply-segmented basal plates as in Nitella. For the 
first time in a Tolypella, some well-developed, hard limeshells were dis- 
covered in a collection of Tolypella prolifera from Indiana. Calcium de- 
posits inside the spiral cells were also found in oogonia of other members of 
the Conoideae. No similar deposits were found in the Allantoideae or in 
Nitella. Descriptions and illustrations as well as a consideration of the 
evolutionary significance of these findings are given. 


In 1961, some fossil charophytes were reported from New York (5). 
Among them were charophytes referred to the recent species, Chara se- 
juncta and Tolypella (probably T. prolifera) . 

The Tolypella oospore (Text-figure 1) was damaged, but its un- 
mistakable characteristics caused no hesitation about referral to that 
genus. However, it was perplexing that the specimen had an undivided 

Figure 1. Oospore of Tolypella prolifera, fossil from Erie Co., New York 
(Table 1, No. 13): 1. lateral view; 2. basal view; 3. apical view, ca. 80 X. 

basal plate (Text-figure 1, no, 2). Grambast (6) described the basal 
plate of a fossil Tolypella from the Tertiary of the Paris Basin as having 
three segments similar to that in Nitella. Horn af Rantzien (7) in 1959 
questioned the persistence of sterile oogonial cells in Nitella and 
Tolypella to produce the three-parted plate. He reported not having 
seen any although some of his photographs suggest their presence. In 
1962, Grambast (9) again discussed the triply-segmented basal plate of 

1 The abundant charophytic remains provided for this study by Norton 
G. Miller, Michigan State University, East Dansing, Mich., made this con- 
tribution possible. 


Plant Taxonomy 407 

charophyte fructifications from the Tertiary of the Paris basin. He 
explained that Horn af Rantzien had examined them and "has seen 
that there is considerable similarity between the triply-segmented basal 
plate of these fructifications and the three oogonial cells demonstrated 
to occur in Nitella." Since both genera belong to the Nitelloideae, they 
were expected to be similar in this respect. 

Recently, more fossil charophyte specimens collected in New York 
were made available for study. Mr. Norton G. Milleri sent four vials 
which contained oospores from the basal marl at Houghton Bog, several 
miles north of Springville, Erie Co., New York. It is 11,880± 730 years 
B. P. (Norton G. Miller communication). The oospores had been treated 
with hydrochloric acid. Some untreated material from this site was 
also obtained. This had been collected with a piston sampler. Oospores 
and lime-shells of Chara sejuncta A. Br. were obtained from that site. 
(For a description and illustration of this species see Daily, 5) 

Another vial contained oospores picked from untreated organic 
debris collected ca. ten feet beneath a gravel deposit near Lockport, 
N. Y. This material was deposited near the shore of Lake Iroquois and 
has been dated at 12,000± 400 years B. P. (1). Chara sejuncta A. 
Br. and Tolypella glomerate Desv. in Lois, were found in that deposit. 
(A description and illustrations can be found in the Systematic Section 
for T. glomerata). Pollen found in the sediments imply that a spruce 
forest was dominant during the time the charophytes were being de- 
posited. (Norton G. Miller communication). 

One of the interesting things about the T. glomerata oospores was 
that the basal plates were triply-segmented instead of undivided as in 
the T. prolifera oospores from the New York deposits. (See also Plate 
1, figs. 8-10, T. intricata, for an undivided basal plate.) It was then 
recognized that these two species represented the two sections of Toly- 
pella. T. prolifera belongs to the Section Conoideae with short, conical 
and acute ultimate cells of the branchlet rays, spiral cells not swelling 
at the apex and persistent coronula. T. glomerta belongs to the Allan- 
toideae with ultimate cells of the branchlet rays elongate, spiral cells 
swelling at the apex and coronula deciduous. 

Therefore, it was decided to examine some herbarium specimens of 
extant species representing the two sections of Tolypella to see if the 
basal plate in the Allantoideae was consistently segmented and in the 
Conoideae undivided. The basal plates were best demonstrated by 
transmitted light and high power. The integuments were removed from 
the basal portion of the oospore which had been hand sectioned and 
flattened on the slide. 

Results obtained with selected specimens are given in Table 1. They 
show that species of Tolypella belonging to the Conoideae do have 
oospores with a consistently undivided basal plate (Plate 1, Fig. 10). 
Members of the Allantoideae may have oospores with a two- or three- 
segmented basal plate (Plate 1, Figs. 12, 14, 15). Furthermore, hard 
lime-shells were found in Tolypella prolifera (Plate 1, Figs. 1-4), T. 


Indiana Academy of Science 

Plant Taxonomy 409 

intricata (Plate 1, Figs. 5-7), and T. fimbriata (Table 1, nos. 12, 9, & 
11 respectively), which belong to the Conoideae. However, none of the 
specimens examined belonging to the Allantoideae produced lime-shells. 
Hard lime-shells in the Conoideae have apparently gone unnoticed before. 
Various authors have described the accretion of crystals on the ex- 
terior of Tolypella oogonia or internal discrete crystals on the oospores 
beneath the integuments in damaged specimens, but hard lime-shells in 
undamaged specimens were thought not to exist. 

A description and illustration of the lime-shell of T. prolifera from 
Indiana is included in the systematic part of this paper. Calcification 
developes inside undamaged spiral cells of the oogonium beginning as 
discrete crystals on the adaxial wall, but finally consolidating and 
filling the slightly convex spirals producing an opaque, smooth, hard 
lime-shell without concentric lamination. A suggestion of the original 
discrete crystalline consistency remains even after the crystals consoli- 
date because of a difference in opacity between the crystals and their 


While studying morphological development in extant Characeae, 
Sawa (10) found two- and three-parted basal plates in Nitella, but ob- 
served only undivided basal plates in Tolypella. However, he stated, 
"there is a separate group of Tolypella-like plants with Nitella-like 
oogonia and oospores some of which might have been identified as vari- 
ous species of Tolypella in the past." That these were stated to be 
laterally compressed is a misprint (personal communication). They 
were not laterally compressed or in other words, they were terete. They 
also had oospores with a three-parted base similar to the Paris Basin 
Tolypella reported by Grambast. They are undoubtedly allied to the 
Allantoideae. The suggestion by Sawa to remove Tolypellas with an un- 
divided basal plate of the oospore from the Nitelleae seems unwarranted. 
Many other morphological characteristics in common between the genera 
Nitella and Tolypella seem sufficient to place them in the same category. 
The possession of an undivided basal plate in the Conoideae, however, 
indicated that this Section of Tolypella is intermediate in this respect 
between the Allantoideae and the Chareae and evolutionarily nearer. 
As far as lime-shells are concerned, none have been found in the genus 
Nitella, none in the Allantoideae, developed in the Conoideae, may or 
may not be developed in the Chareae. 

With the finding of lime-shells in Tolypella, a basis is provided 
for returning several fossil species of charophytes to that genus. The 

Figs. 1-4. Tolypella ■prolifera (Table 1, no. 12). Limeshell : 1. axial section, 
X 100 ; 3-4. apical, lateral and basal views respectively, X 62. Figs. 5-7. T. 
intricata (Table 1, no. 9). Limeshell in lateral, basal and apical views respective- 
ly, X 62. Figs. 8-10. T. intricata (Table 1, no. 10). Oospore in apical, lateral, and 
basal views respectively, X 85. Figs. 11-13. T. glomerata (Table 1, no. S). 
Oospore in lateral, basal and apical views respectively, X 100. Fig - . 14. 
T. glomerata (Table 1, no. 6). Basal plate of the oospore X 10 0. Fig. 15 
T. nidifica (Table 1, no. 1). Basal plate of the oospore, X 270. 

410 Indiana Academy of Science 

lime-shells of extant Tolypella and the fossil genus Sphaerochara (illus- 
trated by Horn af Rantzien, 7, Plate X) are similar. Horn af Rantzien's 
excellent photographs of Sphaerochara (Maedlerisphaera) ulmensis 
could very well be used to illustrate the lime-shell of Tolypella prolifera 
which differs chiefly in size. It is similar in shape, number of convolu- 
tions of the spiral cells, apical configuration with a groove, base, texture 
of limeshell and basal plug. The apex in lime-shells of T. intricata 
(Table 1, no. 9) was more like that of the fossil Sphaerochara heado- 
nensis without the apical groove. Horn af Rantzien (8) removed several 
species formerly assigned to Tolypella to Sphaerochara because: ''Toly- 
pella fructifications do not develope lime-shells (Horn af Rantzien, 8, 
p. 210) and gyrogonites cannot accordingly be lime-shells of fossil 
Tolypella species." However, with the finding of lime-shells in Tolypella, 
that genus is undoubtedly synonymous with Sphaerochara. 

Summary and Conclusions 

The presence of an undivided basal plate of the oospore and pro- 
duction of lime-shells were found to be consistent characteristics in the 
Section Conoideae of the genus Tolypella while divided basal plates and 
lack of lime-shells are consistent in the Allantoideae. In these respects, 
evolutionary development apparently followed a trend from segmented 
basal plates and lack of lime-shells in the genus Nitella and the Allan- 
toideae of the genus Tolypella to unsegmented basal plates and presence 
of lime-shells found in the Conoideae of the genus Tolypella and unseg- 
mented basal plates associated with or without lime-shells in the 
Chareae. The Conoideae are thus an intermediate group with reference 
to these characteristics. However, the two sections of Tolypella form a 
natural group based upon many other characteristics held in common 
and show the greatest affinity to each other. 

The finding of firm lime-shells without lamination in Tolypella 
removes the objection to placing fossil specimens assigned to Sphaero- 
chara in this genus. 

Systematic Section 

Tolypella glomerata (Desv. in Lois.) Leonh. 1863 Lotus 13:129. 

The following is a description of fossil oospores found at the Lock- 
port, N. Y. site. (For full citation see Table 1, no. 8. Illustrated in 
Plate 1, Figs. 11-13). 

Oospore ca 0.312 mm. long and 0.260 mm. wide with 7-9 ridges in 
lateral view. Spirals slightly narrower at apical periphery than at 
equator, then widen slightly and again narrow meeting at a point at 
the apex, narrower at the base meeting around a triply or doubly 
segmented basal plate. Outer colored membrane light brown, decorated 
with granules in linear arrangement with the bases of the granules 
forming an indistinct reticulate pattern. 

Tolypella prolifera (Ziz ex A. Br.) Leonh. 1863. Lotus 13:57. 

Description of lime-shells from extant specimens collected in Poka- 
gon State Park, Indiana. (Full citation given in Table 1, no. 12. Il- 
lustrated in Plate 1, Figs. 1-4). 

Plant Taxonomy 





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412 Indiana Academy of Science 

Lime-shell without utricle and formed by 5 spirals, 0.465 mm. long, 
0.417 mm. broad, spherical to ellipsoid with rounded to protruding apex 
and rounded base, with 11 striae. Spirals flat to convex, ca. 0.062 mm. 
thick and wide at equator, thinner forming a groove around the apical 
periphery, then the cell ends slightly protruding and distinct at the 
apex, at the base meeting around a pentagonal pore completely filled 
with the basal plug. Plug 0.052 mm. broad and about as thick at ma- 
turity. Texture of limeshell crystalline, with occasional fractures and 
without lamination or zonation. 

Literature Cited 

Buckley, J. D., M. A. Trautman, and E. H„ Willis. 19GS. Amer. J. Sci. 
Suppl. 10:246-294. 

Daily, Fay Kenoyer. 1946. Species of Tolypella in Nebraska. Butler Univ. 
Bot. Stud. 8:113-117. 

. 1950. Tolypella prolifera Leonh. in Indiana. Butler Univ. 

Bot. Stud. 9:273-27* 

1954. A rare Tolypella new to the United States of America. 

Butler Univ. Bot. Stud. 11:144-148. 

5. . 1961. Glacial and post-glacial eharophytes from New York 

and Indiana. Butler Univ. Bot. Stud. 14(1) :39-72. 

6. Grambast, Louis. 1956. La plaque basale des Characees. C. R. Acad. Sci. 
(Paris) 242:2585-2588. text-figs. 

7. Horn af Rantzien, Henning. 1959. Recent charophyte fructifications 
and their relations to fossil gyrogonites. Kungl. Svenska Vet. Ak. 
Arkiv. f. Bot. Ser. 2. 4(7) :165-332. 

8. . and H. P. Ulrich. 1965. Fragipans — what are they? Res. Prog- 

charophyte fructifications. Acta Univ. Stockholm, Stockholm Contrib. 
in Geol. 4(2):45-197. 21 pis. 

9. and Louis Grambast. 1962. Some questions concerning re- 
cent and fossil charophyte morphology and nomenclature. Acta Univ. 
Stockholm, Stockholm Contrib. in Geol. J)(3) :135-144. 

10. Sawa, T. 1968. Abstract. Oogonia and fructifications of recent eharo- 
phytes concerning the taxonomic position for Tolypella. J.Phycol. Suppl. 

11. Wood, R. D. 1965. Monograph of the Characeae. In: R. D. Wood and K. 
Imahori, A revision of the Characeae. Vol. 1. J. Cramer, W^einheim. 

Indiana Plant Distribution Records, XX. 1966-68 
Jack Humbles, Indiana University 

Genera are listed in the order of their appearance in Deam's Flora 
of Indiana. Species within each genus are in alphabetical order, and 
are followed by the name of the county in which they were collected. 
Nomenclature is in accord with that used in Gray's Manual of Botany, 
8 ed., 1950. 

Specimens have been collected by John Bard, Mark Fraker, Mary 
Ann Hart, C. Bixler Heiser, III., Jack Humbles, C. Eugene Jones, Dirk 
Walters, and Jim Whitis. 

Voucher specimens for all the new records are in the herbarium of 
Indiana University. 

Taxonomic Entities 

Setaria faberii, Bartholomew, Boone, Brown, Carroll, Cass, Clinton, 
Dearborn, Decatur, Fayette, Franklin, Grant, Hamilton, Hancock, How- 
ard, Jefferson, Jennings, Johnson, Madison, Marion, Miami, Morgan, 
Ohio, Rush, Shelby, Tipton, Wayne. Symplocarpus foetidus, Madison. 
Spiranthes gracilis, Wayne. Saururus cernuus, Orange, Washington. 
Silene nivea, Monroe. Dianthus armeria, Harrison. Spiraea tomentosa, 

Melilotus officinalis, Jackson. Robinia pseudo-acacia, Harrison. Coro- 
nilla varia, Monroe. Desmodium cuspddatum, Ohio. Ailanthus altissima, 
Clark. Acalypha rhomboidea, Ohio. Impatiens caj^ensis, Jackson. Sida 
spinosa, Ohio. Asclepias incarnata, Washington. Verbena urticifolia, 
Scott. Mentha piperita, Ohio. Mentha spicata, Ohio, Scott. 

Datura stramonium, Ohio. Verbascum thapsus, Scott. Chaenorrhinum 
minus, Monroe. Campsis radicans, Jackson. Cephalanthus occidentalis, 
Scott. Lobelia siphilitica, Ohio. Vernonia altissima, Ohio. Eupatorium 
coelestinum, Ohio. Eupatorium rug o sum, Ohio. Silphium perfoliatum, 
Jackson. Ambrosia artemisiifolia var. elatior, Ohio, Switzerland. Xan- 
thium italicum, Ohio. Heliopsis helianthoides, Ohio. Rudbeckia triloba, 
Ohio. Actinomeris alternifolia, Switzerland. Bidens polylepis, Monroe. 
Artemisia annua, Ohio. Senecio glabellus, Morgan. Cirsium arvense, 
Scott. Cirsium vulgare, Scott. Sonchus arvensis var. glabrescens, Monroe. 


The Flowering of Lemna minor and the Establishment of 
Centaurium pulchellum in Northwestern Indiana 

Gayton C. Marks, Valparaiso University 

The flowering of Lemna minor is considered to be an infrequent 
occurrence (4, 5, 6). This rarity of flowering has imposed serious re- 
strictions on studies which contribute to an understanding of the life 
cycle of the Lemnaceae (7). It has been recommended that, when such 
phenomena occur, flowering material be collected for critical study (6). 

Such an occasion did present itself this summer while a routine 
microscopic examination was made of a Schererville pond. The first 
indication of a serendipitous find was an almost transparent bubble- 
like structure adjacent to several thalli of Lemna minor. A more critical 
observation revealed this to be the cup-shaped stigma attached to an 
equally semi-transparent style protruding from the reproductive pouch 
of a parent duckweed. 

Although in no way related to the ferns, the term frond is com- 
monly employed in describing the thallus of the duckweeds. While there 
seems to be considerable controversy in interpreting the inflorescence 
of this group, little disagreement exists in the belief that the Lem- 
naceae is a degenerate offshoot of the Araceae. 

Maheshwari and Kapil have described the frond of Lemna pauci- 
costata as being composed of three indistinct regions: 1. The distal end 
which is primarily photosynthetic in function; 2. A nodal sector flanked 
by the reproductive pouches, and from which arise the root, the daugh- 
ter fronds and the flowers; 3. A basal axial region from which arises 
a pedicel and is partly modified to form the pouches. The frond is asym- 
metric, largely undifferentiated and somewhat broader at the distal end 

With the exception of the pedicel, this description may be cautiously 
applied to Lemna minor. Environmental conditions often lead to such 
variation that species are not readily distinguished (3, 8). 

The primordium of the foot arises from a subepidermal layer which 
is overarched by a protective sheath of epidermis. This sheath is soon 
pierced by a root cap which becomes winged in L. paucicostata but not 
L. minor. The epidermis is without root hairs. The anatomy of the 
root reveals four layers of cells. While the outer two are chlorophyllous, 
the inner two are devoid of pigment and may be compared to the en- 
dodermis and the xylem respectively. The vascular strand in the frond 
is simple. In L. paucicostata, it is composed of a narrow strip of elon- 
gated cells. In Lemna minor, these cells may become lignified (7). 

The question of the floral assemblage is more difficult to assess. It 
commonly has been interpreted as several flowers (usually one female, 
two males) in a common membranous spathe (1, 4, 5). Pool describes 
the family as bearing extremely simple flowers consisting of one stamen 
and one pistil (9). In L. paucicostata the pistil is lateral with respect 


Plant Taxonomy 415 

to the stamens, but in L. minor the carpel lies between two stamens. The 
latter arrangement lends support to the view that what is usually con- 
sidered an infloresence is in reality a flower (7). 

If this assumption is true, it may be appropriate to consider this 
species as near polygamomonecious since what seem to be perfect flowers 
and staminate flowers may be borne on the same individual. It has been 
the practice to regard this entire group as monecious (1, 10). 

As earlier stated, the first indication of the flowering process was 
the appearance of style and stigma. Closer scrutiny revealed that the 
pistil was flanked on either side by immature stamens. This discovery 
prompted a quantitative study. Of six hundred plants examined on 
June 19, 95 or 15.8% bore this type of "flower." At this point it sud- 
denly became apparent that a second type of "flower" was present, this 
being entirely staminate, composed of two stamens only. The next four 
hundred plants examined on this same day produced 113 "flowers" of 
either the perfect or staminate type. This represented about 28% of 
the total population. 

On June 23 a duckweed sampling was taken from a small lake near 
St. John. Only five, or less than 2% of three hundred plants examined, 
yielded flowering individuals. On the same day the original Schererville 
pond yielded 28 flowering plants of one hundred studied. On June 25, a 
ditch connected to the Kankakee River near Schneider was explored 
for flowering specimens. None of the one hundred plants examined 
exhibited "flowers." The Schererville pond produced 117 flowering plants 
of two hundred examined on July 9. This represented approximately 
58% of the population. By August 29, this percentage dropped to 31%, 
and by September 17 it was estimated that this percentage had declined 
to about 15%. 

Statistical errors are easily introduced because of the sampling 
techniques used. Chief among these is properly identifying one plant. 
As in other duckweeds, L. minor reproduces asexually by buds or daugh- 
ter fronds in the same reproductive pouches where flowering occurs. In 
most cases the daughter, granddaughter and great granddaughter fronds 
remain attached to the parent frond forming a chain of thalli. Even 
gentle manipulation produces fragmentation so that the pure definition 
of a plant is at best obscure. A lesser obstacle was the immaturity of 
the stamens. Few seemed to reach anthesis. Many remained subepidermal 
and had to be gently pressed out with a teasing needle to prove their 
existence. Of the hundreds of flowers observed, only a few anthers ex- 
hibited dehiscence. The anthers are tetralocular, but all four micro- 
sporangia do not develop synchronously. One cell may be metaphase 
while others in early prophase (7). 

The stigma and style appear to be but one cell thick. There is but 
one ovule in an ovary and this may be removed with little difficulty. The 
membranous sac (spathe) which invests the "perfect flowers" is curiously 
but sparingly pigmented with red spots. 

With a great amount of disappointment, no fruiting has been ob- 

416 Indiana Academy of Science 

In the Flora of Indiana, Deam listed Centaurium pulchellum as an 
excluded species (2). He had received a report of a specimen from the 
Dunes area without a specific locality and had seen a plant collected in 
South Chicago. He cautiously preferred to "wait and see" if this species 
would establish itself in Indiana. 

Centaurium pulchellum is a member of the Gentianaceae. It produces 
a handsome flower with a salverform corolla and is pleasingly pink. 
Three or four specimens were found cringing from periodic onslaughts 
of a power mower in a Schererville lawn in midsummer of 1968. Since 
that time abundant stands have been found near the hangars and on run- 
ways of the Griffith airport at the eastern edge of that community. These 
plants seemed stunted and judging from other plants, this could be 
explained on the basis of soil. A smaller but more luxuriant population 
was found on the extreme southern edge of Griffith in late summer. 

Literature Cited 

1. Benson, Lyman. 1957. Plant Classification. D. C. Heath and Co. 

2. Deam, Charles C. 1940. Flora of Indiana. Department of Conservation, 
State of Indiana. 

3. Fassett, Norman C. 1957. A manual of Aquatic Plants. University of 
Wisconsin Press. 

4. Fernald, Merritt L. 1950. Gray's Manual of Botany, 8th Ed. American 
Book Co. 

5. Gleason, Henry A. and Cronquist, Arthur. 1963. Manual of Vascular 
Plants of Northeastern United States and Adjacent Canada. D. Van 
Nostrand and Co. 

6. Lawrence, George. 1964. Taxonomy of Vascular Plants. The Macmillan 

7. Maheshwari, Satish and Kapil, R. 1963. Morpholological and Embryo- 
logical Studies on the Lemnaceae. I. The Floral Structure and Gameto- 
pytes of Lemna paucicostata. Amer. J. Bot. 50:677-686. 

8. McClure, Jerry and Alston, Ralph. 1966. A Chemotaxonomic Study of 
Lemnaceae. Amer. J. Bot. 53:849-860. 

9. Pool, William H. 1941. Flowers and Flowering Plants. McGraw Hill. 
10. Porter, C. L. 1967. Taxonomy of Flowering- Plants. W. H. Freeman Co. 


Chairman : M. L. Baumgardner, Purdue University 
James E. Newman, Purdue University, was elected chairman for 1969 


Fluctuations of Bacteriological Numbers in Farm Ponds. L. E. Hughes 
and H. W. Reuszer, Purdue University. — Studies of the fluctuations of 
bacterial numbers in farm ponds was conducted for sixteen months. These 
studies were conducted in three ponds, Ponds A, B, and C. Samples were 
taken approximately once each month from both the surface water and 
water near the bottom of each pond. 

Bacterial numbers in Pond B were about equal in the bottom water 
and the surface water throughout the year, ranging from about 5,000 
per ml in August, 1967, to somewhat greater than 40,000 per ml in 
May, 1967. Larger numbers and greater fluctuations of bacterial num- 
bers were found in Ponds A and C, with generally higher bacterial num- 
bers in the bottom water of Pond A than in the surface water, while 
generally higher numbers occurred in the surface water of Pond C than 
in the bottom water. 

Some factors influencing these bacterial numbers, including total 
carbon, have also been studied. Pond B had consistently lower total 
carbon values in both the surface water and bottom water than did Ponds 
A or C with values ranging from 7.0 mg per liter to 11.6 mg per liter. 
Pond A had values as high as 16 mg per liter while Pond C had even 
higher values, ranging up to 22 mg per liter. 

Earth Mound in Eastern Indiana. James M. Smith, University of Day- 
ton, and Bruce Miller, Miami University. — This paper concerns an 
earth mound about 110 feet in diameter located S. W. X A, Sec. 35, T-ll- 
N, R-2-W, Union County, Indiana. The area is described as a soils prob- 
lem, stressing beginning of B horizon development. Numerous chert 
objects — in all liklihood hearth stones of Adena Indian culture — were 
found in the mound. 


Tillage Techniques on Indiana Prairie Soil 1 

Helmut Kohnke and S. A. Barber, Purdue University 


Several variations of tillage methods and of residue management on 
a fertile prairie soil during six years have resulted in only minor effects 
on corn yields. Even the no tillage plots produced corn within 2.5 percent 
of the conventionally treated plots. However, distinct changes in soil 
properties were observed. These point to eventual major effects on the 
productivity of the soil. 


The introduction of increasingly more powerful tractors has made it 
possible to combine several tillage operations that formerly have been 
performed by horse-drawn equipment. Recognition of the danger of 
erosion has pointed to the need of keeping the soil surface receptive for 
water in order to avoid excessive runoff. Considerably increased fertiliza- 
tion, especially with nitrogen compounds, and use of herbicides have 
caused many farmers to change from crop rotations that included grass 
and legume meadows to continuous cropping with corn and soybeans. 

Historically one reason for tillage has been to stimulate the de- 
composition of the mineral and organic components of the soil in order 
to provide an adequate nutrient supply for the succeeding crop. With the 
plentiful and relatively inexpensive fertilizers available this reason has 
ceased to exist. 

All these facts have caused the introduction of new systems of tillage 
and of crop residue management during the last two decades. How do 
crop yields and soil quality react to this change? In an attempt to find 
quantitative answers an experiment has been conducted on the Purdue 
University Agronomy Farm comparing several tillage treatments for 
corn. It is the purpose of the research reported in this paper to determine 
what effects continuous cropping with corn has on soil conditions and 
corn yields under various types of tillage. 

Experimental Procedure 

An experiment has been designed in which four systems of tillage, 
and four levels of residues were used. Corn was planted annually on six 
of the treatments, while two treatments remained without any crop. 
Every year fertilizer is uniformly broadcast over the entire area at 
rates designed to result in high yields. Herbicides are used and the plots 
with corn are cultivated twice during the season. 

The treatment are given in Table 1. 

The design of this experiment permits the comparison of the effects 
of corn versus no corn, and of the relative effects of four tillage methods. 

1. Journal Paper No. 3514, Purdue University Agricultural Experiment 


Soil Science 


The items that were studied include yield, stalk diameter, and root growth 
of corn and organic matter content, pore space, and aggregation of 
the soil. 

The soil of the experimental area is a mollisol, a tall grass prairie 
soil, grading from a Raub silt loam to a Chalmers silt loan. Table 2 pre- 
sents some of the properties of these soils. 

Table 1. Design of the tillage experiment. 

















Stover Removed 







Field Cultivator 






















* Conventional tillag-e consists of fall or spring- plowing- followed by 

TABLE! 2. Properties of the soils of the experiment area. 


silt loam 

("2" profile) 


















Silt loam 




Silt loam 



Silt loam 






Silty clay loam 






Clay loam 



4 .,8 








Chalmers silt loam ("8" profile) 


















Silt loam 




Silt loam 









Silty clay loam 






Clay loam 






Clay loam 






Indiana Academy of Science 

Except for the variables studied all other treatments were identical 
for the entire experimental area. The same corn hybrid was used. Fertili- 
zation consisted of an annual application of 250 lbs N, 100 lbs P 2 5 , and 
100 lbs. K 2 per acre. Atrazine at a rate of 4 lbs /acre was used to con- 
trol grassy weeds. In the fall the stover on all plots was cut with a stalk 

Conventional tillage (treatments 1, 2, 3, 7) consisted of plowing to 
20 cm and disking three times. Whenever the soil moisture conditions 
permitted plowing was done in the fall. All other operations were done 
immediately preceding corn planting. The field cultivator (Graham- 
Hoeme type) was used twice to a depth of 20 cm. The rototiller loosened 
the entire surface soil to about 8 cm depth. Both these treatments included 
two cultivations. The soil of these treatments (4 and 5) were disked 



S no 



Figure 1 

Cultivator (4) 

^Conventional (I) 
xNo Tillage (6) 



62 63 64 65 66 67 

Comparison. of the corn yields of the residue management 
plots with those in Tippecanoe County and in Indiana 

Soil Science 


twice. In the case of the "no tillage" treatment the corn planter was 
used twice in succession in order to get the seed deep enough into the 
ground. The soil treatments 1, 2, 3, 4, 5, and 7 were cultivated twice. Since 
treatments 6 and 8 were not cultivated they received an application of 
2, 4-D in addition to the atrazine. 

While treatment 7 (no corn) was managed in exactly the same 
manner as treatments 1, 2, and 3, treatment 8 (no corn) consisted ex- 
clusively of application of fertilizers and herbicides. 

Results and Discussion 
Corn Yields 

The average coin yields of the four plots of each of the treatments 
during the first six years of the experiment are given in Table 3. Figure 
1 compares the yields of three of the treatments with the average yields 
for the State of Indiana and Tippecanoe County during the same years. 
The effect of the changing weather from year to year is quite evident. The 
yields from the field cultivator plots have been consistently the highest. 
The yields from the three "Conventional tillage" treatments and from 
the rototiller treatment did not vary statistically from each other. Prob- 

Figure 2 















2 3 4 5 6 

62 63 64 65 66 67 

Comparison of the effects of treatments with those of 
weather on corn yields 


Indiana Academy of Science 
Figure 3 









' .1 .2 .3 .4 .5 .6 .7 


Corn root distribution with depth 

ably the most astonishing result is that the average yield of the no-tillage 
treatment was only five percent lower than that of conventional treat- 
ment. This difference becomes still smaller when the first two years are 
excluded from the comparison (2.5 percent difference). In the first two 
years the no-tillage treatment suffered from inadequate stand because 

Soil Science 423 

of the difficulty to get the seed into the ground. This small difference in 
yield indicates that the soil of the experimental area is very favorable for 
corn growth even without any tillage treatment. 

It is interesting to note (Figure 2) that the differences in yield due 
to differences in treatment were smaller than due to the variations due to 
weather from year to year. 

The root distribution of corn in one of the "conventional tillage" 
plots was compared with that in a "no tillage" plot (Figure 3). At a 
location 6 inches from the plant "conventional" corn had three times the 
weight of roots as "no tillage" corn. At a location 20 inches from the 
plant (midway between the rows) this ratio was two to one. In spite of 
this great difference in root growth yields were quite similar. 

TABLE 3. Cora yields, bushels per acre. 





































































In the seventh year of the experiment (1968) all plots were planted 
to corn, otherwise the treatments remained the same. A measurement of 
the diameters of the cornstalks (Figure 4) indicates the cumulative effects 
of six years of differential treatments. The three conventional plow 
treatments and the field cultivator treatment gave about the same re- 
sults. The biggest diameters occurred in the roto-till plots. There was a 
definite decrease of diameters in the plots that had been fallow for six 
years as well as in the no-tillage plots. These latter, as has been men- 
tioned before, had a substantially smaller amount of roots than the 
conventionally treated plots. 

Organic matter content of the soil 

The organic matter content of the soil of the various treatments 
was determined anually since 1965. Because the organic matter content 
of the surface soil has the greatest effect on the moisture regime most 
of the determinations were made on soil of the 0-4 inch depth. While there 
have been variations from year to year the relations of organic matter 


















Indiana Academy of Science 
Figure 4 

3 4 5 6 7 



Diameters of corn stalks as affected by residue treatment 
and tillage August 1968 

TABLE 4. Organic matter content of experimental plots of the 0-4 inch 
depth of the soils of the treatments of the residue management 


Sampled May 1968 



Matter Percent 


Conventional, normal residue 



Conventional, residue removed 



Conventional, double residue 



Field Cultivator 






No tillage 



No corn — tillage 



No corn — no tillage 


Soil Science 


content from treatment to treatment have essentially remained the 
same. Table 4 shows the organic matter situation in the spring of 1968. 

It is interesting to note that the highest organic matter content in 
the surface soil occurs in the roto-tiller plots where the residues are 
incorporated in the upper four inches only. Leaving the residues on top 
of the ground as done in the no-tillage plots evidently causes them to 
decompose more quickly. The addition of extra residues has resulted in 
a substantial increase in organic matter compared to leaving the normal 
amount of residues in the field. As was to be expected removing the 
residues reduced the organic matter content considerably. Leaving the 
soil bare of any vegetation (treatments 7 and 8) resulted in a decline of 
the organic matter content even greater than where only the corn roots 
were left in the ground (treatment 2). 

After six years of conventional cropping with corn the soil had 
0.4% more organic matter than the soil that remained fallow for the 






32 r 


t 24 



2 20 


: : 




h- 12 




October 1967 

April 1968/ 

3 4 5 6 



Effect of treatment and season upon aeration porosity 
0-3 inch depth 


Indiana Academy of Science 

same period. This represents an average annual loss of 1300 lbs per 
acre of organic matter (dry basis) from the soils that had no vegetation. 
Surely an outstanding testimony for the value of corn as a soil conserving 
crop — under conditions of high fertilization and no erosion! 

In all the treatments in which the soil was tilled down to 20 cm, the 
organic matter content was rather uniform to this depth. In the rototill 
and the no-till plots, however, there was a substantial decrease of 
organic matter below 10 cm. 


A comparison of the aeration porosities of the soil of all the treat- 
ments in the fall of 1967 and in the following spring (Figure 5) shows 

Figure 6 













3 4 5 6 7 



Aggregation index of the surface soil (3-5 inch depth) as 
affected by residue treatment and tillage 

Soil Science 427 

very little lasting influence of the treatments. Only the roto-tilled plots 
(treatment 5) had a substantially higher aeration porosity in the spring 
than the other treatments. The data for October 1967 are a direct reflec- 
tion of the mechanical treatments. Most of the soils have maintained their 
loose structure throughout the growing season. The greatest decrease 
of porosity of the "conventionally" treated plots occurred in treatment 
7 where no vegetation existed that could have protected the soil from 
the impact of the rain. 

Previous experience has shown that 10 percent aeration porosity in 
the upper root zone is sufficient for satisfactory corn growth. In most 
of the treatments the actual aeration porosity in the spring before culti- 
vation was somewhat lower than this value. This indicates a need for 
loosening of the soil and the reason for a decrease in yield of corn on the 
no-tillage treatment. 

The water-stable aggregation of the soil of the various treatments 
was determined by the wet-sieving method (Figure 6). Most of the soils 
showed rather satisfactory values between 0.25 and 0.35 mm aggregation 
index. The two treatments with accumulation of organic residues in the 
surface (Nos. 5 and 6) had considerably greater aggregation indices. The 
no-corn, conventional tillage treatment (No. 7) had the lowest value. The 
fact that no organic matter was added and that the cultivation caused 
oxidation of the most active part of the humus may be the reason for this. 

Temperature fluctuations 

A temperature study conducted in the summer of 1966 showed 
interesting differences in daily temperature fluctuations between the 
treatments. These data are shown in Table 5. 

The fact that temperature fluctuations in the soil increase with 
increasing thermal conductivity is clearly brought out by these data. The 
corn in treatments 1 and 2 reduced insolation compared to treatments 
7 and 8. The corn residue plowed into the soil of treatment 1 reduced the 
thermal conductivity compared to treatment 2, from which the residues 
were removed. The much denser structure of the no-tillage soil (treat- 

TABLE 5. Daily temperature fluctuations at 10cm depth in the soil and 

in the air July 1966. 

Daily temperature 
Treatment degrees F. 

1 Corn, conventional tillage 7.8 

2 Corn, conventional tillage, residues removed 10.7 

7 Fallow, conventional tillage 12.3 

8 Fallow, no tillage 16.9 
Air, 30cm above ground 27.7 

428 Indiana Academy of Science 

ment 8) had a higher thermal conductivity than the soil of treatment 7 
which has been plowed every year. 

The greater the daily soil temperature fluctuations are, the greater 
are the vapor pressure fluctuations and consequently losses of soil mois- 
ture in the vapor phase. These data indicate clearly that protection from 
insolation and incorporation of crop residues into the soil assist in mois- 
ture conservation. 


During the six years this experiment has run the effects of tillage and 
crop residue management on corn yields have not been great. However, 
the changes in the physical conditions of the soil have been pronounced. 
As compared with corn produced with conventional tillage, some treat- 
ments caused a decrease in soil physical conditions, whereas other treat- 
ments improved them. Where the practices used cause a marked reduc- 
tion in one or more of the physical conditions this portends a warning 
that these changes could eventually cause yield decreases. If this is true 
on the fundamentally very productive mollisols, it would be much more 
so on soils of lower quality. The farmer must stay forever vigilant to use 
the best management practices for his soil. 

Fitting Plants to Fragipan Soils in Southern Indiana 

Maurice E. Heath, Purdue Agricultural Experiment Station 

This paper deals with recent research and observations of fragipan 
soils and plant relations found in the southern Indiana sandstone shale 
soil region (Figure 1). These soils comprise 2,348,000 acres — nearly a 
tenth of the state. The sandstone shale soil region extends into all or 
parts of 19 counties. It is an unglaciated area with deeply cut valleys and 
rolling to steep topography. Approximately 40 percent of the area is 
wooded — too steep for the use of farm equipment. The native pH of the 
loess covered hills ranges from 5-3 to 5.5. 


O >^j; 

Figure 1. The sandstone shale soil region (in black) extends into all or 
parts of the 19 southern Indiana counties. Soil fragipans are commonly 
found throughout the area. 

The Environmental Model 

Soil fragipans restrict moisture movement as well as root depth 
development. Fragipans are formed under low pH in humid areas (8). 
Plant roots will not penetrate the fragipan unless through a cleavage or 
crevice. Thus, for many of these soils, the shallow root zone depth will 
commonly range from 18 to 34 inches (7). The root zone water holding 



Indiana Academy of Science 

capacity may range from less than 4 to 4% inches (5). The fragipan 
itself may range from 10 to 15 inches in thickness. 

Rainfall averages 40 to 46 inches annually. Soils are often water- 
logged in winter and very early spring. Nearly half the rainfall is lost as 
run-off. There is little, if any, lasting snow cover during winter with 
diurnal temperatures frequently fluctuating above and below freezing. 
Tap rooted alfalfa is a high risk crop. The total vertical winter soil motion 
during 1965-66 was shown to range from 22.03 inches at the surface to 
0.39 inches at the 12-inch soil depth (3). It was considerably greater 
during the winter of 1966-67. Droughts are common in mid-summer. 
Moisturewise the root zone is an environment of extremes — from wet 
to dry. 

Since 1953, when research first started on the Southern Indiana 
Forage Research Farm (Dubois County), several major environmental 
differences have been observed between the Brookston-Crosby soil area 
(Lafayette) and the sandstone shale area (Forage Farm). The observed 
differences are as follows: 

Environmental Factors 

Root zone 

Soil fragipans 

Water-holding capac- 
ity in the root zone 

Water logging in 


as deep as needed 


8-12 inches 

seldom when tiled 

Sandstone Shale 

shallow 18"-34" 


less than 4" to AW 


Winter heaving 

little except ponded 



Annual rainfall 



Annual run-off 



Summer drought 


30 days or more, 80 
percent of the years 


level to undulating 

rolling to very steep 

Percent of land in 

93 (Tippecanoe 


54 (Lincoln Hills — in- 

U.S.-SCS classes 

cludes Crawford, 


Harrison, Spencer and 
Perry Counties) 

Erosion hazards 

little to none 


Cation exchange 




Organic matter 

low to medium 

extremely low 

Examples of agronomic crops considered low risk for the Brookston- 
Crosby soil region are corn, soybeans, winter wheat, alfalfa, and brome- 
grass. For the sandstone shale soil region examples of low risk agronomic 
crops are the sorghums, winter wheat and rye, red clover, Korean 
lespedeza and tall fescue. 

Fitting Crop Plants and Practices to the Environmental Model 

To protect many of these hillside soils from the ravages of erosion 
it would appear, in general, that either the natural vegetation of wood- 

Soil Science 


land or a close growing adapted forage grass and/or crop plant must be 
employed. Several grasses, legumes and agronomic practices of promise 
will be discussed. 

Tall fescue (Festuca arundinacca Schreb), a perennial forage grass, 
is a comparatively late comer to Indiana. Kentucky 31 tall fescue was dis- 
covered in 1931 on Mr. William Suiter's farm in Menefee County, Ken- 
tucky. It was not publicized greatly until about 1945. Since that time 
thousands of acres of tall fescue have been seeded on hill land fragipan 
soils in Southern Indiana. It has particular adaptation to this environ- 
ment of extremes from wet to dry and greatly insulates the soil surface 
reducing freezing and thawing during winter. Presently it is by far the 
most adapted grass for these conditions. Much research is now underway 
to learn how to use this grass more profitably in a livestock system on 
these kind of soils. Excellent winter pastures of tall fescue have been 
produced for beef cows using the practice of the round bale and aftermath 
growth (Figure 2) rationed with an electric fence (6). 

Figure 2. This hill land field was reclaimed from abandonment in 1957 and 
seeded to tall fescue. With the round bale and ration grazing technique 
and a practical level of fertilization, as much as 180 beef cow pasture days 
per acre in mid-winter has been achieved. 

432 Indiana Academy of Science 

Reed canarygrass (Phalaris arundinacea L.) performance results 
elsewhere would strongly indicate that it fits this environmental model of 
extremes — from wet to dry. It is now being evaluated as a possible 
drought pasture for beef cows on the overflow valley soils with slow per- 
meability. Here again, the rationed round bale-aftermath pasture looks 
promising to bridge the summer drought. 

Several perennial lateral rooted legumes appear promising for use 
on these kinds of soils. Crownvetch (Coronilla varia L.) when managed 
as a hay plant has shown very good response (2). In 1954 roadside plots 
10 feet in width were seeded. Ten years later the crownvetch had spread 
75 feet much of the spreading was through a good tall fescue sod. In fact, 
crownvetch allowed to reach full bloom will shade out tall fescue. The 
composition of crownvetch is reported by several workers to be similar to 
that of alfalfa. Crownvetch research is continuing as to its place in 
mixtures for hay production, beef cattle acceptability, fertility and 
management requirements for intensified production. 

Zigzag clover (Trifolium medium L.) planted in 1955 has spread a 
distance of 25 feet through a dense tall fescue sod. Zigzag clover in 
bloom from a distance resembles red clover. Baled zigzag clover hay has 
been consumed as readily by beef cows as other common hays. Currently 
it is necessary to propagate vegetatively due to a lack of a good seed pro- 
ducing type. A small root nursery, approximately 10' x 20' in size, has 
been established on a selected farm in each of 15 counties of the sand- 
stone shale soil region. As these nurseries develop it is planned to use 
the roots for propagation. With the aid of a mechanical planter the 
4- to 6-inch root cuttings will be planted in hillside pasture and meadow 
sods. Field observations will be made under actual farm conditions while 
at the same time small plot management trials will be expanded at the 
Forage (research) Farm. 

In 1954 several big trefoil accessions from the Pacific Northwest 
were tested on the Forage Farm but did not prove winter hardy. How- 
ever, in 1964, I identified big trefoil (Lotus uliginosus Schkuhr) growing 
on Mr. Clarence Kaiser's farm in Crawford county. Mr. Kaiser had 
observed this legume growing and spreading in eight different areas in a 
well fertilized tall fescue meadow. He thought it to be birdsfoot trefoil 
(L. corniculatus L.). The field was managed as hay in the spring and 
then grazed the rest of the season. Three of the eight ecotypes obtained 
from Mr. Kaiser's tall fescue field have shown excellent forage and seed 
production characteristics in small plots on the Forage Farm. Presently 
seed is being increased of the three ecotypes and will be further tested 
for hay and pasture use. They look very promising to grow with fescue 
on these soils. Big trefoil has tolerance to wet soils and low pH. 

Lateral rooted alfalfa has been reported to be much more tolerant 
to heaving than the commonly used tap rooted alfalfas (4). In an attempt 
to tailor make a lateral rooted alfalfa that would be happy in this environ- 
ment, Dr. R. L. Davis made many crosses between the lateral rooted and 
tap rooted types. These were grown on the Forage Farm f ragipan soils for 
several years and their growth characteristics observed. The best growing 

Soil Science 433 

and spreading crosses were selected for further screening in the green- 
house. From these, the best genotypes were combined into an experi- 
mental synthetic (1). Currently the experimental is being "on farm" 
tested in eight counties in the sandstone shale area. Management research 
is also being conducted on the Forage Farm. The goal is an alfalfa that 
will give persistence of stand along with acceptable productivity for 
pasture and hay. 

Crambe, a new industrial oil seed crop of promise, fits this environ- 
mental model when planted in mid-April. It can be harvested approxi- 
mately 85 days after planting and prior to the mid-summer drought. It 
has excellent seedling vigor and can be grown in close drills. It will fur- 
nish much soil protection from raindrop energy in a short period of time 
after seeding. It is visualized that the crambe crop could be followed by 
a summer seeding of grasses and legumes in August or by a wheat or rye 
crop in the fall. 

Tall fescue sod planting on hill land looks promising. The idealized 
model would provide sod dormancy for approximately two months while 
the summer annual, such as sorghum, became sufficiently developed to 
shade the fescue. After the sorghum matured the fescue would again 
grow to furnish fall pasture along with the sorghum residue and provide 
an improved pasture or hay meadow the following year. Thus far we 
have not found a satisfactory growth regulator to obtain consistent sod 
dormancy. Presently, it is necessary to use cover crops and reseed 
following sod planting on sloping land. 


1. The environmental model of the unglaciated sandstone shale 
fragipan soils in Southern Indiana is a contrast of extremes when com- 
pared to the well drained corn soils of Central Indiana, thus requiring 
plants with special adaptation. 

2. Presently, tall fescue is the most adapted perennial grass to 
furnish hillside protection and stabilization. 

3. Lateral rooted legumes of promise are being tested as associates 
to grow with tall fescue and other grasses to further improve the forage 
quality and productivity on these hill land soils. Crambe, a promising 
industrial crop, also appears to fit the environmental model. 

4. Special cultural and management practices such as sod planting 
and the rationed round bale technique for winter and summer drought 
pastures appear very promising on these kinds of soils. 

Literature Cited 

1. Busch, Robert and R. L. Davis. 1966. Creeping rooted alfalfa research on 
the Southern Indiana Forage Farm. Res. Progress Report 233; also 
R. P. R. 282 (1967). Purdue (Indiana) Agr. Exp. Sta. 

2. Heath, Maurice E. 1968. Crownvetch (Coronilla varia L.) trials on Southern 
Indiana fragipan soils. Second crownvetch Symposium. Penna. State Univ. 

434 Indiana Academy of Science 

3. Newman, James E. 1967. Total soil heaving motions at the Southern 
Indiana Forage Farm during the 1965-66 winter season. Res. Progress 
Report 297. Purdue (Indiana) Agr. Exp. Sta. 

4. Southworth, W. 1921. A study of the influence of the root system in 
promoting hardiness in alfalfa. Sci. Agr. I, 5-9. 

5. Wiersma, Daniel. 1962. Soil moisture at the Forage Farm during the 
1961 season. Res. Progress Report 33. Also see R. P. R's 66 (1963), 132 
(1964), 185 (1965), and 246 (1966). Purdue (Indiana) Agr. Exp. Sta. 

6. Wilson, Lowell, R. C. Peterson, M. E. Heath and R. E. Erb. 1965. Re- 
stricted versus unrestricted winter grazing of round fescue bales and 
aftermath for the beef cow herd on the Forage Farm. Res. Progress 
Report 189, Purdue (Indiana) Agr. Exp. Sta. 

7. Zachary, A. L., et al. 1964. Soils of the Forage Farm. Res. Progress Re- 
port 131. Purdue (Indiana) Agr. Exp. Sta. 

8. . and H. P. Ulrich. 1965. Fragipans — what are they? Res. Prog- 
ress Report 184. Purdue (Indiana) Agr. Exp. Sta. 

Fertilizer Experiments with Corn on Several Soils in 
Indiana, 1963-1965 1 

Russell K. Stivers, Purdue University 2 

The primary purpose of this research was to study the response of 
continuous com (Zea mays L.) to phosphorus and potassium fertilization 
on several different soils in Indiana. Barber and Humbert (1), after 
reviewing advances in the knowledge of potassium fertilization from 
1951 to 1961, concluded that more data are needed on the relationship 
between crop response to added potassium and the level of available 
potassium in the soil. On a Runnymede loan testing medium in potassium, 
Stivers and Griffith (7) found that it took four years of cropping with 
continuous high yielding corn before a significant yield increase with 
potassium fertilizer was obtained. It was thought that ths soil might 
have fixed potassium since the increase in available soil potassium, after 
four years of applying 100 pounds per acre of potassium annually, was 
small. Barber and Humbert (1) state that soil type may have to be 
considered more in the future because of the relationship between soil 
type and potassium fixation and potassium release. 

Barber and Stivers (2) have reported that broadcast and plow under 
phosphorus fertilizer is more important than row phosphorus fertilizer 
for corn when more than 17 pounds per acre of phosphorus is applied. 
At low soil test levels, corn yields were increased by row applications of 
phosphorus, but not a medium and high levels. 

Methods and Procedures 

Different rates and placements of commercial fertilizers were applied 
to soils on field plots on which corn was grown. Height, yield of grain 
and chemical composition of the ear leaf were determined. 

These experiments were conducted on three different farms, all of 
which were in Tippecanoe County (Table 1). The phosphorus fertility and 
the potassium fertility experiments on the Gwin Farm were adjacent to 
each other at the same location. The Byers and Brown farms experiments 
were initiated in 1963; the two experiments on the Gwin Farm were 
initiated in 1964. 

One of the three replications on the Byers Farm was on Fincastle 
soil, and two replications were on Russell soil (Table 1). On the Gwin 
Farm, no clear demarcation between Martinsville and Russell soils was 
made. The two soils are very similar except that Martinsville has coarser 
parent material in the D horizon and is slightly more droughty than 

1. Journal Paper No. 3519, Purdue University Agr. Exp. Sta. 

2. The author acknowledges the help of S. R. Miles, Ethel Tudor, N. 
T. Houghton, PT. K. Kesler, Enola Ruff, R. P. Shaw, and Paul Crane in 
conducting this research. 



Indiana Academy of Science 



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Soil Science 437 

On the Crosby soil location and on the Fincastle and Russell soils 
location, three replications were used. Experimental design was an unbal- 
anced incomplete block design with 28 treatments. A randomized complete 
block design with four replications was used for the Martinsville and 
Russell soils experiments. All plots were 65 feet long and four 42-inch 
corn rows wide. The center two rows minus five feet for border on each 
end were harvested. 

Soil samples were taken prior to the start of each experiment from 
every plot. Fifteen or more cores were taken seven inches deep in each 
plot and composited for one sample. They were then air dried, screened 
and mixed, and subsamples were tested by the Purdue Soil Testing 
Laboratory using methods of Spain and White (5, 6). 

On the Fincastle and Russell soils having a pH of 5.6, two and 
one-half tons of agricultural limestone were applied before plowing, and 
one and one-half tons of limestone were applied after plowing in 1963. 
On Crosby soil agricultural limestone was applied at the rates of one and 
one-half tons per acre before plowing and three-fourths ton per acre after 
plowing in 1963. In 1964, three and three-quarters tons per acre of agri- 
cultural limestone were applied to the same soil and plowed under for the 
phosphorus and potassium experiments. 

Fertilizer materials used were 33.5 percent nitrogen from ammonium 
nitrate, 24 percent phosphorus from superphosphate in 1963 and 1965 
and 9 percent phosphorus from superphosphate in 1964, and 50 percent 
potassium from muriate of potash. 

Two hundred pounds per acre of nitrogen was applied on all experi- 
ments each year prior to plowing. An additional ten pounds per acre of 
nitrogen was applied beside two of the four corn rows of all treatments in 
each plot in 1963 on both the Crosby soil location and on the Fincastle 
and Russell soils location. Rates of broadcast phosphorus and potassium 
fertilizer were applied in only one year, 1963, on the Crosby soil location. 
All broadcast phosphorus and potassium fertilizers were applied in the 
spring, half before plowing and half after. On the Martinsville and 
Russell phosphorus fertility experiment, 598 pounds per acre of potassium 
was broadcast. On the potassium fertility experiment on the same loca- 
tion, 120 pounds per acre of phosphorus was broadcast and 40 pounds 
per acre of P was used in the row. In all experiments where row fertilizer 
was used, it was applied approximately two inches horizontally away 
from the seed at planting. 

Aldrin granules for controlling soil insects were broadcast on plowed 
ground and disked in immediately prior to planting on all experiments 
conducted in 1964. Broadcast atrazine spray after planting to control 
weeds was applied to both Martinsville and Russell soils experiments in 
1964 and to the Crosby experiment in 1965. Conventional cultivation and 
hand hoeing were used on all experiments as needed to control weeds. 

Funks G-96 and Indiana 678 corn hybrids were used for seed on all 
experiments in 1963 and in 1964 respectively. Pfisters Associated Growers 
Sx29 was used on the Crosby location in 1965. 

438 Indiana Academy of Science 

Stands per acre counted at harvest were approximately 16,200 on 
the Crosby location in 1963, 19,800 in 1964, and 16,300 in 1965. Final 
stands on the Martinsville and Russell experiments in 1964 were approxi- 
mately 16,000 plants per acre. On the Fincastle and Russell location final 
stands were approximately 16,500 plants per acre. 

Dates of planting on the Crosby location were June 4 in 1963, May 
22 in 1964, and May 13 in 1965. The Martinsville and Russell soils location 
was planted May 29, 1964. The Fincastle and Russell soils location was 
planted May 24, 1963. 

Height measurements were made on the tallest extension of leaves 
of each of 10 or more plants within the harvest area of each plot of all 

Composite corn ear leaf samples of 15 or more leaves per treatment 
taken at the green silk stage, were ground and then sent to the Depart- 
ment of Agronomy, Ohio Agricultural Research and Development Center, 
Wooster, Ohio, for analyses. 

For yield determination, ear corn was hand harvested. Weights and 
moisture percentages in the grain were determined, and yields were 
calculated using Remmenga's tables (3). 

Results and Discussion 

These experimental sites were selected because they had low or very 
low soil test values for the fertilizer nutrient being studied, and they 
represented important soil types in Indiana (Table 1). 


Relation of height of corn 31 days after planting to rates of broadcast 
phosphorus and potassium fertilizer treatment. Fincastle and Russell 
soils, H. Byers Farm, Lafayette, Indiana, 1963. 

Rate of P in Lbs. per A. 

Rates of K 

in Lbs. per A. 

52 105 



— — 



19.1 20.7 



18.7 18.1 

Lsd at the 20% level = 

: 1.7 inches 

Lsd at the 5% level = 

: 2.6 inches 

Lsd at the 1% level = 

3.5 inches 

Height of corn increased 3.3 to 7.9 inches as phosphorus rates 
increased on the Fincastle and Russell soils location in 1963 (Table 2). 
There was no significant differences in height between the Fincastle soil 
(Replication I) and the Russell soil (Replications II and III). 

Soil Science 439 

Yields on the Fincastle replication were 11 to 12 bushels per acre 
higher than the yields on the two Russell replications. The probability 
was more than 99 to 1 that this difference was real and was not due to 
chance alone. Yields of corn grain were significantly increased 4.5 to 
15.1 bushels per acre as rates of phosphorus fertilization increased in the 
same experiment (Table 3). There was also a trend toward higher phos- 


Relation of yields of corn grain with 15.5 percent moisture to rates of 
broadcast phosphorus and potassium fertilizer treatment, Fincastle and 
Russell soils, H. Byers Farm, Lafayette, Indiana, 1963. 

Rate of P in Lbs. 



Rate of K 

in Lbs. per A. 



Bu. per A. 


. — , 

— . 









Lsd at the 20% level = 


bu. per A. 

Lsd at the 5% level = 

13.8 bu. per A. 

Lsd at the 1% level = 

21.7 but. per A. 

phorus content of the ear leaf of corn taken at silking as rates of phos- 
phorus increased (Table 4). Where no phosphorus was applied, percent 


Relation of chemical composition of the ear leaf of corn at silking to 
fertilizer treatment, Fincastle and Russell soils, H. Byers Farm, Lafay- 
ette, Indiana, 1963. 



A. of 

Percent Composition 

Parts per Mill 










B Cn 









18 17 









18 13 







A l 


17 13 










16 12 











2-3 2-5 


phosphorus in the ear leaf was in the low range as denned by the Purdue 
Plant and Soil Anailysis Laboratory (4). The response to phosphorus was 
large for the first increment of fertilizer as might have been expected 
from the very low soil test value for phosphorus. 

In the same experiment there was no increase in height of corn in 
relation to rates of potassium fertilization (Table 2), but there was a 

440 Indiana Academy of Science 

significant increase in yield (Table 3). Where phosphorus was applied 
and potassium was not, corn ear leaf content of potassium was in the 
low range (Table 4). Evidently, soil potassium available to corn was not 
low enough to influence height 31 days after planting, but it did influence 
the plants later in the season. 

After 1963, it was impossible to obtain additional data from this 
Fincastle and Russell soils experiment. After the death of the owner, 
H. Byers, the new owner would not lease the land for agricultural 
research purposes. 

On the Martinsville and Russell soils location height of corn 46 days 
after planting was significantly increased as rates of both row and 
broadcast phosphorus increased (Table 5). Twenty pounds per acre of 


Relation of height of corn 46 days after planting to rates of row and 
broadcast phosphorus fertilizer treatment, Martinsville and Russell 
soils, C. Gwin Farm, Lafayette, Indiana, 1964. 

Broadcast P R° w Application of P in Lbs. per A. 

Lbs. per A. 

5 10 20 30 40 

44 52 55 56 57 56 

60 49 56 58 58 — — 

120 56 — — — — — 

300 — — — 60 

Lsd at the 20% level = 3 inches 
Lsd at the 5% level = 5 inches 
Lsd at the 1% level = 7 inches 

row phosphorus resulted in the same height of corn as 120 pounds per 
acre of broadcast P, or a 1 to 6 efficiency ratio. Drouth was so severe 
at this location in July and August, 1964, that yields ranged from 2 to 87 
bushels per acre, and they were not significantly related to treatment. 
This great range in yields reflected a large difference in available soil 

Height of corn 48 days after planting was significantly reduced at the 
high rates of broadcast potassium on the Martinsville and Russell soils 
location (Table 6). It is thought that high soluble salts in the soil and 
relatively dry soils reduced rate of growth in this drouthy season. Yields 
of corn grain were very low, ranging from 2 to 27 bushels per acre, and 
they were not significantly related to potassium treatment. 

On both of these experiments on the Martinsville and Russell soils 
there was great variation in initial soil test values from individual plots. 
As a result of this variation and the yield variation, particularly in the 
phosphorus experiment, the location was discontinued. 

Soil Science 441 


Relation of height of corn 46 days after planting to rates of row and 
broadcast potassium fertilizer treatment, Martinsville and Russell soils, 
C. Gwin Farm, Lafayette, Indiana, 196 U> 

Row Applications of K in Lbs. per A. 

Broadcast K 

Lbs. per A. 25 50 75 100 125 

67 66 69 64 68 66 

149 69 69 65 64 

598 62 — — — — 59 

Lsd at the 20% level = 3 inches 
Lsd at the 5% level = 5 inches 
Lsd at the 1% level = 6 inches 

On Crosby soil in a three-year period (1963-1965) height and yields 
of corn were significantly increased as phosphorus fertilization increased, 
but height and yields were not meaningfully related to potassium fertiliza- 
tion even though initial soil test levels for phosphorus and potassium were 
very low and low respectively (Tables 7 and 8). Analyses of the ear leaf 


Relation of height of corn six weeks after planting to rates of broadcast 
phosphorus and potassium fertilizer treatment, Crosby soil, E. R. Brown 
Farm, Lafayette, Indiana, 1963-1965 inclusive. 

Rate of K 

Applied in 1963 

Lbs. per A. 

Rate of P Applied in 1963 
Lbs. per A. 

66 131 

31 — — 

149 28 37 39 

299 31 36 41 

Lsd at the 20% level = 3 inches 
Lsd at the 5% level = 4 inches 
Lsd at the 1% level = 5 inches 

of the 1963 crop indicated that phosphorus was low when none was 
applied, and potassium was deficient (4) when none was applied (Table 
9). Higher yield levels might have resulted in yield differences related 
to potassium fertilization. However, on this Crosby soil in this 1963-1965 
period soil test levels did predict response to phosphorus fertilizer, but 
they did not predict response to potassium fertilizer. 

442 Indiana Academy of Science 


Relation of yields of corn grain with 15.5 percent moisture to rates of 
broadcast phosphorus and potassium fertilizer treatment, Crosby soil, 
E. R. Brown Farm, Lafayette, Indiana, 1963-1965 inclusive. 

Rate of K 

Applied in 1963 

Lbs. per A. 

Rate of P Applied in 1963 
Lbs. per A. 

0' 66 131 

Bu. per A. 
44.0 — — 

149 38.9 70.1 78.4 

299 47.2 78.5 94.0 

Lsd at the 20% level = 17.2 bu. per A. 
Lsdatthe 5% level = 26.3 bu. per A. 
Lsd at the 1% level = 35.0 bu. per A. 


Relation of chemical composition of the ear leaf of corn at silking to 
fertilizer ti-eatment, Crosby soil, E. R. Brown Farm, Lafayette, Indiana, 



L<bs. per A. of 


t Composition 

Parts per 



P K 




























5 2 










3 9 

135 299 










LiOav range 










Deficiency range 

less than — 











Average yields of corn on one replication of a Fincastle soil were 11 
more bushels per acre higher than those on two replications of a Russell 
soil of a fertilizer experiment. On this experiment and also on a similar 
experiment on Crosby soil, height of corn, yield of grain and phosphorus 
composition of corn ear leaves were increased as rates of phosphorus fer- 
tilization increased. On a third location on Russell and Martinsville soils, 
height of corn was increased, but yields were not increased in a 

Soil Science 443 

drouthy season, as rates of phosphorus fertilization increased. The initial 
phosphorus soil test levels of all three locations were low or very low. 

Potassium fertilization increased yield and potassium composition 
of ear leaf of corn on the Fincastle and Russell soils location testing low 
in potassium. High rates of broadcast potassium fertilizer on a Martins- 
ville and Russell soils location reduced corn height in a very dry year. 
On a Crosby soil there was no height or yield response of corn to 
potassium fertilization over a three-year period even though the initial 
soil test level for potassium was low. 

Literature Ctied 

1. Barber, Stanley A. and Roger P. Humbert. 1963. Advances in the knowl- 
edge of potassium fertilization. Fertilizer Technology and Usage. Soil 
Science Society of America, Madison, Wisconsin. 

2. and Russell K. Stivers. 1963. Phosphorus fertilization of 

of field crops in Indiana — research prior to 1963. Purdue University Re- 
search Bulletin No. 759. 

3. Brunson, Arthur M. 1959. Prior to harvest estimating corn yields. Pur- 
due University Extension Circular 472. 

4. Hood, E. L. 1968. Interpretation of corn leaf analysis as used by the 
Purdue F'ant and Soil Analysis Laboratory, 1968. Mimeograph. 

5. Spain, J. M. and J. L. White. 1960. pH and lime requirement determina- 
tion. Purdue University soil testing mimeograph. 

6. and . (no date). Procedure for the determination of 

phosphorus and potassium. Purdue soil testing mimeograph. 

7. Stivers, R. K. and D. R. Griffith. 1966. Nitrogen and potassium rates 
for continuous corn on Runnymede loam, 1962-1965. Purdue University 
Research Progress Report 248. 


Chairman: John 0. Whitaker, Jr., Indiana University 
James C. List, Ball State University, was elected chairman for 1969 


Temperature Preferences in the Eastern Garter Snake 
(Thamnophis sirtalis sirtalis). David C. Kramer, Ball State University. — 
Nine Eastern Garter Snakes were maintained in a thermal gradient box 
with a controlled photoperiod for 54 days. The snakes were observed 
twice daily and the environmental temperature selected by each was 
recorded. After 20 days' exposure to a daily photoperiod of 14 hours, the 
snakes were exposed to a photoperiod of 10 hours for 34 additional 
days, but no change in temperature preferences was noted. The average 
temperature selected by all of the snakes for the entire test period 
was 22.9 degrees C. However, there was a consistent tendency for 
all snakes to select a slightly warmer temperature in the afternoon than 
in the morning. The average difference between the preferred morning 
and afternoon temperatures was approximately 2.0 degrees C. 

Relationship between Emergence Rhythm and Metabolic Rhythm in 
Drosophila melanogaster. William J. Brett, Indiana State University. — 
A population of wild type Drosophila melanogaster was maintained in a 
twelve-hour-light — twelve-hour-dark cycle with light onset at 9:00 a.m. 
Females were allowed to oviposit over a 24-hour period to provide com- 
parably aged samples. The oxygen consumption for a number of pupae 
obtained from each sample was determined for a one-hour period at 
three-hour intervals over a 48-72 hours period. The remaining pupae in 
each sample were permitted to emerge and number of adults determined 
at three-hour intervals. The peak for emergence occurred at 12:00 
noon. Oxygen consumption exhibited a major peak at 10:00 p.m. and 
a minor peak at 1:00 p.m. With the exception of the minor peak the 
two curves were almost mirror images of one another. These results 
suggest that emergence may actually occur during a metabolic-low. 

The Effects of Chicken Luteinizing Hormone on the Pullet Ovary. John 
M. Burns, Indiana University. — Mammalian luteinizing hormone (LH) 
has very little effect on the immature chicken ovary and, therefore, a 
study was initiated to determine the response of the pullet ovary to 
chicken LH. As no purified samples of chicken LH were available it was 
necessary to develop a method for obtaining LH activity from chicken 
pituitary glands without interference from the pituitary follicle stimulat- 
ing hormone (FSH). This was done by treating chicken pituitary glands 
with neuraminidase which inactivated the FSH. The administration of 
chicken LH to immature chickens resulted in: 

(1) a significant 32 P uptake by the ovary in 15 minutes. 

(2) an increase in 32 P incorporation into the RNA fraction of the 


446 Indiana Academy of Science 

(3) a marked increase in the levels of ovarian 17-B estradiol, 
estrone, and estriol. 

Tracheal Mucous Velocities in the Rabbit, Dog, and Rat. Andrew W. 
Gruenholz and Henry Tamar, Indiana State University. — The tracheas 
of 24 Dutch rabbits, 16 dogs, and 6 Charles River albino rats were excised, 
opened, and immersed in oxygenated Tyrode's solution. The transit time 
of a 1 mm. platinum disc over 5 mm. of pseudo-stratified ciliated epi- 
thelium was determined at 39° C for the rabbits and dogs, and at 
37° C for the rats. The insulated test chamber was leveled, extraneous 
heat was eliminated, and a 15-minute adaptation period was allowed. 
Transit times for the rabbit and dog remained constant for one hour. 
The mean mucous-flow velocity was 0.292 mm. /second for the rabbit, 
0.585 mm. /second for the dog, and 0.167 mm. /second for the rat. The 
difference between the rabbit and dog values was significant at the 0.1 
level. Solutions of 0.2M and 0.25M NaCl depressed ciliary activity in 
the dog respectively 1.9 and 1.68 times more than the equivalent concen- 
trations of KC1. In the rabbit as well 0.2M NaCl was more depressant 
than 0.2M KC1. The trachea of the rabbit exhibited rhythmic contractions 
of its blood vessels following immersion in 0.2M KC1. 

Studies on the Growth Rate of the Juvenile Pilot Black Snake 

(Elaphe obsoleta). Robert E. Geyer, Jr. and William B. Hopp, Indiana 
State University. — Fifteen pilot black snakes were hatched from eggs in 
the laboratory. These snakes were individually housed and divided into 
five groups of three snakes per group. Each snake in group I received 
1 cc of food per week, group II, 1.5 cc per week, group III, 2.0 cc per 
week, group IV, 2.5 cc per week, and group V, 3.0 cc per week. The 
snakes were force-fed a mixture of strained baby foods consisting of beef, 
beef liver, and veal by the syringe-catheter technique. Although several 
measurements were taken, weight gain was found to be the most reliable 
basis for determining growth. Weekly recordings of weight were made 
during the 12-week period this experiment was conducted. 

At the end of 12 weeks the snakes in group I had gained an average 
of 1.93 grams, group II, 2.80 grams, group III, 3.33 grams, group IV, 
4.77 grams, and group V, 6.25 grams. The average gain in weight per 
week for each group was 0.16, 0.23, 0.28, 0.40, and 0.52, respectively. 

Rabies in Indiana Bats 

John 0. Whitaker, Jr., Walter A. Miller, and William L. Boyko 

Indiana State University 


Indiana State Board of Health 


A total of 626 bats routinely submitted to the Indiana State Board of 
Health from 1965 to 1968 was examined for rabies using- the fluorescent 
antibody test and sometimes the mouse test. Of these 42, or 6.7%, were 
rabid. Twenty-four of 364 big brown bats, Eptesicus fuscus, were rabid as 
were 11 of 152 red bats, Lasiurus, borealis. Bat rabies was more common in 
the southern than in the northern portions of the state, and was more 
common in the summer and fall than at other times. There were two apparent 
outbreaks of rabies in Indiana in bats in 1967, one in Eptesicus fuscus in 
Jefferson County and one in Lasiurus borealis in Vanderburgh County. Bat 
rabies has been reported in a total of 17 Indiana counties. There appears 
to be no positive correlation in Indiana between rabies in bats and rabies in 
other species. A total of 133 big brown bats was collected in "normal wild" 
populations of Eptesicus in Jefferson County at the height of the 1967 rabies 
outbreak, but all proved negative for rabies. 


Rabies was first found in a United States bat in 1953 when a Florida 
Yellow bat was killed while attacking a seven-year-old boy (7). Rabies 
has now been reported in all of the 48 contiguous states; Rhode Island 
became the last on June 12, 1967 (1). The disease has now been found 
in at least 20 species of bats. 

Some summarized incidence figures for bat species in which large 
numbers of bats have been examined in Florida (5, 7), Texas (6), south- 
western United States (2), southern New England (3) and Illinois 
(4, 8) are: 

examined Number rabid Percent rabid 

Lasiurus borealis 




Eptesicus fuscus 




Tadarida brasiliensis 




Lasiurus floridanus 




Lasiurus seminola 




Pipistrellus subflavus 




Myotis velifer 




My otis grisesce?is 




Myotis lucifugus 




Myotis austroriparius 





448 Indiana Academy of Science 

Illinois is the only midwestern state in which there have been exten- 
sive surveys for bat rabies. A total of 652 bats was examined (8, 4) and 
none were found to be rabid. Unfortunately only 12 were red bats, the 
species in which the highest incidence of rabies generally occurs. Of the 
remainder, 504 were Myotis, the genus which generally has the lowest 
incidence of rabies. Fifty-five were Eptesicus, 65 were Pipistrellus, 8 
were Plecotus and 8 were Lasionycteris noctivagans. 

Before the present study there were four confirmd cases of bat 
rabies in Indiana. A red bat, Lasiurus borealis, taken in Tippecanoe 
County, September 5, 1960, was rabid, as was a silver-haired bat, 
Lasionycterius noctivagans, taken September 12-13, 1960, in Montgomery 
County. Thirty-three additional bats, unidentified, taken in 1960, were 
not rabid. None were examined from 1961 through 1963, but in 1964, 37 
unidentified bats were found to be negative for rabies. Two rabid speci- 
mens were found in 1965 among 70 unidentified bats. One of the rabid 
bats was from Delaware County and one from Monroe County. 

The present study was initiated in 1965 to determine: 

a. the distribution and incidence of rabies in Indiana bats 

b. the relationship between rabies in bats and rabies in other 
species of mammals in Indiana 

c. the relation between incidence of rabies in bats turned in by 
Indiana citizens to the Indiana State Board of Health and 
rabies in wild-taken bats. 

Materials and Methods 

The brains of all bats submitted to the Indiana State Board of 
Health were examined directly for Negri bodies and by the fluorescent 
antibody method. Brain material from many was injected into white mice 
as a further test. These bats came from throughout the state. Starting 
in 1966, all bats submitted to the Indiana State Board of Health were 
identified by the senior author. 

Many bats came from a small number of counties. In two counties, 
Jefferson and Vanderburgh, there were widely publicized bat rabies 
scares in 1967; these are the counties from which we have examined the 
largest numbers of specimens. Three other counties from which large 
numbers of bats have been examined are Marion, Johnson, and St. Joseph, 
all of which have yielded rabid bats. Bat rabies in Vanderburgh County 
was chiefly in red bats, Lasiurus borealis, while in Jefferson County it 
was primarily in big brown bats, Eptesicus fuscus. Efforts were made to 
obtain large numbers of bats from "normal wild colonies" of Eptesicus in 
Jefferson County on July 21, 1967, at the peak of the outbreak. A total 
of 97 bats was collected that day by the senior author and Robert Kerr, 
the Jefferson County Sanitarian. Later samples by Mr. Kerr increased 
the total of bats from the "wild" populations to 133. 

Information on rabies in species other than bats was acquired from 
the files of the Indiana State Board of Health. This information was 
compared with the bat rabies information. 



A total of 759 bats was examined during this study, 626 of them 
submitted to the Indiana State Board of Health by citizens, physicians, 
veterinarians, and law or health officials for routine rabies examination, 
and 133 from the Jefferson County "wild sample." 

Species of bats infected 

Two of the species collected in small numbers, Pipistrellus subflavus 
and Lasiurus cinereus, had the highest incidence of rabies of any of the 
bats taken during this study (Table 1), but this may be chance. Only 16 
individuals of each species were taken. 

table 1. Rabies in bats collected by citizens of Indiana, summarized 

by species, 1965-1968. 

No. examined 

No. rabid 

% rabid 

Eptesicus fuscus 




Lasiurus borealis 




My otis lucifugus 




(incl. 2 probably this 


My otis sodalis 


M. keeni 


Lasionycteris noctivagans 


Nycticeius humeralis 


Pipistrellus subflavus 




Lasiurus cinereus 







The two most common bats of Indiana appear to be Lasiurus borealis, 
the red bat, and Eptesicus fuscus, the big brown bat. Lasiurus borealis 
is a solitary species, but is quite common, especially in southern Indiana. 
This species would appear to be the most commonly infected of the United 
States bats that have been studied. In Indiana, 7.2% of the red bats 
submitted by citizens to the State Board of Health were rabid (Table 1). 
Eptesicus fuscus is most often directly associated with human beings, 
often forming large colonies in barns, houses and other buildings. For 
this reason, and since large numbers of Eptesicus have not been examined 
for rabies, data on this species are particularly desirable. The rate of 
rabies infection in Eptesicus, 6.6%, was slightly lower than in red bats. 

In Indiana, the major cave bats are species of Myotis, of which 6Q 
specimens were examined, one (1.5%) being rabid. 

Overall, 42 of 626, or 6.7% of the bats submitted to the Indiana 
State Board of Health were rabid. 


Indiana Academy of Science 

Geographic distribution of bat rabies in Indiana 

During the present study (Table 2), rabies was found in bats from 
fourteen counties: Daviess, Dearborn, Fayette, Gibson, Greene, Hendricks, 
Jefferson, Jennings, Johnson, LaPorte, Marion, Monroe, St. Joseph, and 
Vanderburgh, bringing to 17 the total number of counties from which 
rabid bats have been taken (including Delaware, Montgomery, and 
Tippecanoe, counties in which bat rabies was found before the initiation 
of this study). 

table 2. Rabies in bats collected by citizens of Indiana, summarized 

by county, 1965-1968. 

No. examined 

No. rabid 

% rabid 































































































table 2. Rabies in bats collected by citizens of Indiana, summarized 
by county, 1965-1968. — (Continued) 

No. examined 

No. rabid 

% rabid 


* Marion 

* Montgomery- 


St. Joseph 






































* Bat rabies found before initiation of present study. 

During this study, most of the rabid bats were from southern Indi- 
ana. The state was divided roughly into thirds, from north to south. 
Three of 97, or 3.0% of the bats from the northern third of the state were 
rabid, 9 of 221, or 4.1% from the central third, and 30 of 308 or 9.7%, 
from the southern third of the state were rabid. This difference was 
significant (Chi-square = 9.68**, 2 df). However, the two major bat 
rabies counties, Jefferson and Vanderburgh, are in the southern third. 
Excluding information from those two counties, seven of 81 bats, or 
8.6%, were rabid from the southern third of the state, thus still consti- 
tuting a greater percentage infection than in the north or central 
counties, but this difference was not significant (Chi-square = 3.10, 1 df). 


Indiana Academy of Science 

Time of year of infection 

Rabies infections in bats were summarized by time of year of occur- 
rence (Table 3). Most rabid bats were taken in the summer, but these 
were also the months in which most of the bats were taken. Of 429 bats 
taken in the summer (June- August), 32, or, 7.5% were rabid, while in 

table 3. Time of year of rabies infection of bats caught by 

Indiana citizens. 

(No. rabid per year is in parentheses.)