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PROCEEDINGS
of the
Indiana Academy
of Science
Founded December 29, 1885
Volume 78
1968
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
1969
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
Academy.
6. The Constitution and By-Laws reprinted from v. 74 are available to
members upon application to the Secretary. Necrologies reprinted from the
various volumes can be supplied 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
TABLE OF CONTENTS
Part 1
THE WORK OF THE ACADEMY
Page
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
ADDRESSES AND CONTRIBUTED PAPERS
Presidential Address 49
"Urban Geology — A Need and A Challenge", William J. Wayne
"Science, Communication, and the Critical Mass" 65
Robert E. Gordon
Anthropology
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
Bacteriology
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 colvf 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
Botany
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
Chemistry
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
Ecology
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
Entomology
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
Physics
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
Zoology
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
PARTI
THE WORK
OF THE
ACADEMY
1968
William J. Wayne, President
Officers and Committees for 1968
OFFICERS
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
DIVISIONAL CHAIRMEN
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
EXECUTIVE COMMITTEE
(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
BUDGET COMMITTEE
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.
COMMITTEES ELECTED BY THE ACADEMY
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.
COMMITTEES APPOINTED BY THE PRESIDENT
(President an ex officio member of all committees)
Academy Representative on the Council of A.A.A.S.: W. H. Johnson.
Auditing Committee: 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.
Smith.
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.
SPECIAL COMMITTEES APPOINTED BY THE PRESIDENT
Biological Survey Committee: J. D. Webster, chairman; L. Chandler; C.
B. Heiser; G. C. Marks; R. Mumford; W. H. Welch; F. Young.
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.
SPRING MEETING
The Honeywell Center, Wabash, Indiana
MINUTES OF THE EXECUTIVE COMMITTEE MEETING
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
Center.
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-
lows:
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-
12
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
unanimously:
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,
and;
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,
and;
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-
mously.
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
FALL MEETING
MINUTES OF THE EXECUTIVE COMMITTEE MEETING
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
follows:
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.
15
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
Academy:
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
MINUTES OF THE GENERAL SESSION
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-
bers.
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-
18
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
unanimously.
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
FINANCIAL REPORT OF THE INDIANA ACADEMY OF SCIENCE
January 1-December 31, 1968
I. ACADEMY ACCOUNTS
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
TOTAL 1968 INCOME & CREDITS:
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
Secretary
Clerical $
Postage, etc.
Treasurer
Clerical
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
Expenditu
re Budgeted
$ 285.S3
$ 250.00
; 201.83
84.00
235.00
225.00
130.00
105.00
243.91
175.00
ues
172.16
180.00
100.00
12.00
75.00
3gy)
*
150.00
55.22
150.00
4,339.86
400.00
400.00
3,939.S6
500.00**
*
100.00
700.23
650.00
155.73
544.50
600.00
600.00
974.45
* Billings not yet received.
** Plus Publications Income Fund Receipts, above.
Self supporting item, not budgeted.
20
Minutes of the Executive Committee 21
ii. administered accounts
Item or Description
Publications Fund
Operational Funds
ACADEMY ACCOUNTS:
Acad. Research Fund
Science Fair Fund
Science Talent Search
J.S. Wright Library Fund
Lilly Library Fund III
Science & Society Committee
Miscellaneous
STATE ACCOUNTS: 1
TOTAL IN ALL ACCOUNTS 17,568.47
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
TOTAL ASSETS IN BANKS AND SAVINGS
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,
1968
1968
Dec. 31,
Balance
Receipts
Expenditures
Balance
724.00
4,073.00
4,339.86
457.14
6,201.22
5,339.90
3,278.80
8,262.32
6,925.22
9,412.90
7,618.66
8,719.46
678.58
790.00
1,160.91
307.67
1,938.76
6,375.00
797.77
7,515.99
2,319.95
1,000.00
1,324.15
1,995.80
134.28
134.28
5,565.68
1,190.74
4,374.94
1,650.00
1,183.58
466.42
6.00
16.03
22.03
0,643.25
9,831.03
5,657.15
14,817.13
7,568.47
19,243.93
13,275.81
23,536.59
INDIANA JUNIOR ACADEMY OF SCIENCE
OFFICERS
President: Dennis Waltke, Division of University Schools, Bloomington
Vice-President: James Peterson, Brebeuf Preparatory School, Indianap-
olis
Secretary: Rachel Koontz, New Haven Senior High, New Haven
JUNIOR ACADEMY COUNCIL
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)
YOUTH ACTIVITIES COMMITTEE
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
PROGRAM
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"
Awards.
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.
22
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.
PROGRAM OF PAPERS
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-
ington.
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-
anapolis.
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.
Hosts
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.
MINUTES OF THE THIRTY-SIXTH ANNUAL
MEETING OF THE
INDIANA JUNIOR ACADEMY OF SCIENCE
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
campus.
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
program.
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
magazine.
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
INDIANA JUNIOR ACADEMY OF SCIENCE
1967-1968
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
High
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
Gary
Gary
Gary
Griffith
Mu Alpha Theta, Andrean H. S.
Biology Club, Lew Wallace H. S.
Griffith Junior High Science Club,
Griffith Junior H. S.
Sponsor
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,
SS.C.M.
Sr. Marie Carmel,
SS.C.M.
Sr. M. Nadine,
SS.C.M.
Lola Lemon
Fred Meeker
26
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
28
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 Engineering1 Technical
Society, Central H. S.
South Bend Second Year Biology Class, Clay
H. S.
South Bend IONS Club, J. W. Riley H. S.
South Bend LaSalle High School
Terre Haute Pius X Science Teens, Schulte H. S.
Tipton Tipton H. S. Science Club, Tipton
H. S.
Trafalgar Indian Creek School
Vincennes Sigma Tau Science Club, St. Rose
Academy
William Beuoy
Roger Moody
Keith Hunnings
E. H. Sanders
Mary Zehner
Ralph Settle
Robert Freemyer
Ernest Litweiler
John V. Davis
John Marker
Sr. Marie Barbara,
S.P.
Richard Garst
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
Algae:
Bryophyta:
Vascular Plants:
Platyhelminthes:
Crustacea:
Arachnoidea:
Insecta:
Pisces:
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
1968.
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
1968.
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.
29
:m
Indiana Academy of Science
Aves:
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.
Mammalia:
Whitaker, J. O. Jr. 1967. Habitat relationships of four
species of field mice in Vigo County, Indiana. Ecology
48:867-872.
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
1968.
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
State.
Nichols, S. A. 1968. The phytocenology of selected Indi-
ana natural areas. M. Sc. Purdue.
Insecta:
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.
Amphibia:
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
Co.
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-
ana.
Mammalia: Whitaker, J. O. Jr. Continued studies on the mammals
of Vigo County, Indiana (parasites, food, habitat,
reproduction).
Mammalia: Terrel, T. Ij. Purdue. Ecology of the swamp rabbit in
Indiana.
Necrology
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
32
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
Star.
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
appreciated.
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.
38
Indiana Academy of Science
MILLARD S. MARKIJF
1883-1968
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.
Necrology
41
FRANK N. WALLACE
1878-1968
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
ZRL
Dr. Ernest M. Agee, Dept. Geosciences, Purdue Univ., Lafayette, Ind. 47907
PSG
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
PHM
Mr. N. Franklin Burnett, 11610 Crestwood Court, Indianapolis, Ind. 46239
DLZ
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
43
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
C
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.
DLZ
Dr. Ralph A. Llewellyn, Rose Polytechnic Inst., Terre Haute, Ind. 47803. P
Mr. Francis C. Lundin, Biology Dept., Ball State Univ., Muncie, Ind. 47306.
Z
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.
CLZ
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.
OZC
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.
LZB
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.
RS
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
PART 2
ADDRESSES
AND
CONTRIBUTED
PAPERS
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.
PRESIDENTIAL ADDRESS
Urban Geology — A Need and A Challenge1
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-
scapes.
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.
49
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
51
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).
"2|L,
^"""If '-'
m
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
condition.
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
53
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
bed.
54
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
downslope.
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
stable.
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
reports.
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.
I
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
r»«)
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.
trwimTTiipTTnTTrrmrt
Space open or
Casing .^ fjMed with soi|
Casing,
pTTmrnrrnTiTTTTiTTTTm
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-
ment.
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.
Summary
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
years.
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
land).
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
table.
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
description.)
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-
ment
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
Illustrations
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
surface
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
community.
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,
65
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
development.
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
community.
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?
ANTHROPOLOGY
Chairman : B. K. Swartz, Jr., Ball State University
Robert E. Pace, Indiana State University, was elected
chairman for 1969
ABSTRACTS
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
71
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
University.
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
Abstract
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
perforated.
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.
Introduction
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
73
71
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.
Sample
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.
Blasingham).
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
suggests.
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.
Methods
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 (N2) samples (Fig. 3). The chi-square test,
e
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,
*=2
,.d.= yj
S(fda)
N
Anthropology
?Y
ko
35
30
25 ■;
,'>'
IS
10
0,
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.
Results
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 (N2) = 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 Na
Chi-square test of significance of distribution of
perforation by sex: X2 = 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: X3 = 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: X3 = 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 N2
Chi-square test of significance of distribution of
right perforation by sex: X2 = 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.
80
Indiana Academy of Science
Male
Sample = 17
Left Humerus
Width
(mm)
Height
(mm)
Right Humerus
Width
(mm)
Height
(mm)
X
s.d.
5.33
3.26
3.76
± 1.41
4.76
2.22
3.76
± 1.94
Female
Sample
28
Left Humerus
Width
(mm)
Height
(mm)
Right Humerus
Width
(mm)
Height
(mm)
X
s.d.
6.25
2.99
4.39
1.91
4.77
2.14
3.82
1.26
Male + Female
Sample = 45 —
Left Humerus
Width
(mm)
Height
(mm)
Right Humerus
Width
(mm)
Height
(mm)
X 5.94 4.18 4.76
s.d. ± 2.76 ± 1.78 ± 2.18
3.79
± 1.55
Chi-square test of significance of size differential by sex
Chi-square Left Humerus Right Humerus
Both Humeri
Xa 3.41 5.23
> 0.05 < 0.05
p — 0.06 = 0.02
4.32
< 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.
Conclusions
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
females.
4) Bilateral perforations are twice as common as unilaterial perfora-
tions.
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
right-handed.
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
Abstract
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
populations.
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.
83
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
plotted.
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-
Anthropology
Ho
300n
3 4 5 6 7 8
AGE IN YEARS
Figure 1. Length of humerus and femur plotted against dental age. Dots
represent individual values, lines represent group growth eurves.
300-.
AGE IN YEARS
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.
86
Indiana Academy of Science
3WH
AGE IN YEARS
Figure 3. Length of tibia and radius plotted against dental age.
represent individual values, lines represent group growth curves.
Dots
30&-,
250-
5<H
ULNA
• ••Dickson
— Yokem
T^7
T^T
T~T
AGE IN YEARS
TTT"^
»2
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
Abstract
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
88
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
reported.
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
Sioux.
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
conclusions:
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
and
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
93
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
variety.
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
coterminous.
Literature Cited
1. Croxton, Frederick. 1959. Elementary Statistics with Applications in
Medicine and the Biological Sciences. New York: Dover Publications,
Inc.
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
Abstract
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.
Introduction
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.
97
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.
104
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.
(4)
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
complex.
Conclusion
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.)
BACTERIOLOGY
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.
ABSTRACTS
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 (Smr) mutants of E. coli WWU were isolated
and characterized. Some Smr 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.
109
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
further.
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 xk 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 H20) 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 104 to 10a.
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 (p02) 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 p02 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.
BOTANY
Chairman: Thomas R. Mertens, Ball State University
Robert L. Kent, Indiana Central College, was elected chairman for 1969
ABSTRACTS
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
115
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 Na2Se03
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 Q02 values for control (-Se) root segments were ca 5.6
ul/hr/mg dry wt. and 5.0 for Qco2 values. A 5xl0~6 M Na2Se03 con-
centration appeared to give a 3 to 4% stimulation in 02 uptake over
periods of 2 hours. High concentrations progressively decreased 02 up-
take: 11% with 10-3 M Na2Se03 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 N2 contain-
ing traces of 02 was decreased to less than 1% of the controls, regard-
less of the presence or absence of Na2Se03. Similarly, the rate of C02
production by selenite-treated roots in equilibrium with N2 plus traces of
02 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 C02 output by Na2Se03.
Time-course studies demonstrated a gradual decrease in 02 uptake
and C02 output rates for both the control and the 104 M selenite-treated
roots in equilibrium with air. Roots treated with 10" M selenite indicated
Botany 117
a modest immediate stimulation in 02 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
Na2Se03 treated roots also declined with time. In contrast, the 102 M
selenite-treated roots showed a very strong immediate stimulation in
C02 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 102 M Na2Se03 treated roots. This doubling of
the R.Q. reflects both the stimulation in C02 output and the depression in
02 uptake due to high concentrations of Na2Se03.
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.
Abstract
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
Site3
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
120
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
examination.
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.
Discussion
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.
Lycopodophyta
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
families.
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
larger.
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
Botany
123
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
alations.
Coniferophyta
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
Botany
v>r>
Figure §
Figure 0
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
pattern.
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
nucellus.
Arthrophyta
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
Botany
129
Figure 9
Figure 10
St..
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.
Pteridospermophyta
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
Lyginopteridaceae.
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
trunk.
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
preservation.
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
found.
Botany 133
Pterophyta
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
plants.
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
evident.
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.
134
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.
Interpretation
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
correlation.
table 2. Genera represented in Indiana and Illinois coal ball studies.
o f>
Genus *j g g
US* d) 4> (0 O O
O o ft tf tf W W
Stigmaria
*
*
*
Lepidodendron
*
*
*
*
Lepidocarpon
*
*
N
*
Lepidophylloides
*
Sigillaria
*
Lepidostrobus
*
*
*
Calamites
*
*
*
Asteromyelon
*
*
Calamostachys
*
Paleostachys
*
Asterophyllites
*
Annularia
*
Sphenophyllum Stem
*
*
Sphenophyllum Root
*
Callistophyton
*
Physostoma
*
Conostoma
Botany 137
TABLE 2. (continued)
Genus
fft
CO
CO
CI
CI
Z'
<Tj
C".
CTj
c-
CI
«3
CO
O)
£
s
C
C5
CTj
■7.
m
Pi
H
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rt
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Heterangium
*
Medullosa Stem
*
Myeloxylon
*
*
•'!■
Neuropteris
*
Alethopteris
*
Pachytesta
*
H
: *
Stephanospermum
*
*
Telangium
:
'?
Cordaites Stem
*
Cordaites Leaf *
Amyelon *
Cordaianthus
*
Cardiocarpus
*
*
*
Botryopteris
*
*
•'-
Pecopteris
*
*
*
Tubicaulis
*
Etapteris
*
Psaronius Root
:|:
* :
|J *
*
Psaronius Stem
*
*
*
Scolecopteris
*
*
Cyathotrachus *
Anachoropteris
*
Notoschizaea *
Asterotheca
*
Myriotheca
*
Phytocarpus
*
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
Period.
138 Indiana Academy of Science
Literature Cited
1. Andrews, H. N. 1961. Studies in paleobotany. John Wiley and Sons, New
York.
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-
53.
Algal Records for Three Indiana Sewage Stabilization Ponds
C. Mervin Palmer
Advanced Waste Treatment Research Laboratory, FWPCA,
Cincinnati, Ohio
Abstract
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.
139
140
Indiana Academy of Science
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Botany
141
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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
Napoleon
Sunman
St. Paul
Green Algae
Blue-Green Algae
Flagellate Algae
Diatoms
29
11
23
4
23
7
19
4
17
7
12
5
TOTAL
07
53
41
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.
Botany
143
TABLE 3. Relative Persistence of Algae in Indiana Sewage Ponds.
(No. Dates Recorded per Alga)
ALGAL GENUS
Napoleon
Sunman
St. Paul
Equivalent1"
Achnanthes
Actinastrum
3
5
5
Anabaena
9
3
5
Anacystis
10
18
20
Ankistrodesmus
■11
33
40
Aphanizomenon
1
-
5
Arthrospira
-
3
-
Carteria
3
1
-
Characium
1
2
-
Chlamydomonas
39
30
40
Chlorella
35
40
40
Chlorococcum
3
5
10
Chlorogonium
16
6
25
Chodatella
L5
-
-
Chromulina
8
2
5
Chroomonas
13
1
-
Cladophora
1
1
-
Closteriopsis
5
-
-
Closterium
5
-
-
Coelastrum
7
10
10
Cosmarium
-
1
-
Cryptomonas
22
1
10
Cyclotella
-
-
20
Dictyosphaerium
8
1
5
Eudorina
2
5
5
Euglena
41
42
40
Francea
1
-
-
Golenkinia
8
7
15
Gomphonema
3
4
10
Gonium
1
2
-
Hantzschia
1
-
-
Johannesbaptistia
-
-
5
Kirchneriella
7
9
-
Lepocinclis
3
1
5
Lobomonas
4
1
-
Massartia
2
-
5
Micractinium
14
7
20
Nannochloris
-
13
15
Navicula
14
15
35
Nitzschia
41
36
40
Oedogonium
-
2
-
Oocystis
11
22
30
Oscillatoria
37
35
40
144
Indiana Academy of Science
table 3. (continued)
ALGAL GENUS
Napoleon
Sunman
St. Paul
Equivalent*
Palmella
Pandorina
Phacotus
Phacus
Phormidium
Planktosphaeria
Pleodorina
Pteromonas
Pyrobotrys
Raphidiopsis
Scenedesmus
Schizothrix
Selenastrum
Sphaerocystis
Spirogyra
Spirulina
Stigeoclonium
Tetraedron
Trachelomonas
Ulothrix
Zoochlorella
3
16
2
11
1
14
1
5
4
39
26
1
2
2
5
16
30
7
15
2
-
3
20
2
-
-
15
11
-
4
10
15
35
19
35
8
5
*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
common.
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
Japanese).
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 Abscission1
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
100
80-
60-
.o40
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
1 /LIGHT
/ /
t /
/ /
/ /
/ /
/ W
/ ^
~" 1 /
1 /
/ /
■/
/ /
m,^-^L ^ 1,1,1.
40 80 120
TIME AFTER EXCISION, HR.
160
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.
146
Botany
147
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
weeks.
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
Indicated
29° 24° 22° 22°
Light Light Light Dark
22.5°
4,500
to
4,800 ft-c
8 Hr
26±4
58±8
73±10
110±20
12 Hr
30±1
69±11
104±12
90±12
16 Hr
32±4
69±11
105±6
100±10
700
to
1,000 ft-c
8 Hr
21±1
40±6
55±10
101±12
12 Hr
23±3
52±8
60±9
150±10
16 Hr
28±5
64±10
73±20
133±20
20 Hr
29±2
50±13
73±20
127±11
30°
700
to
1,000 ft-c
8 Hr
24±3
40±7
42±1
154±19
12 Hr
27±3
34±6
44±5
118±6
16 Hr
26±3
40±1
60±20
155±13
20 Hr
27±3
35±6
68±20
103±20
148
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:
x
0s
O
e
UJ
120-
O
a: 80
2
UJ
40-
0
- 1
■ I
' i
i
1 1 1
1 1
18
22 26 30 34
38
TEMPERATURE, °C
Figure 2. Acceleration of explant abscission in continuous light as a
function of temperature. Explants prepared from greenhouse-grown plants.
Bars show standard deviations.
Botany
149
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.
Results
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
100-
o
<
<
a.
LU
CO
0 20 40 60 80 100
TIME AFTER EXCISION, HR
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) .
Photo-
Time to
Temperature
Light Intensity
period
50% Separation
(C°)
(Ft-C)
(Hr)
(Hr)
22.5
4,500-4,800
8
88
12
98
16
112
700-1,000
8
68
12
98
16
108
20
112
30
700-1,000
8
44
12
24
16
56
20
24
15
700-1,000
8
>144
12
>144
16
>144
20
>144
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-
regular.
table 3. Acceleration of Abscission in the Light by Inverting the
Explants.
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
0 200 32
1 100 27
2 80 27
152
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-
30
«r
20
<
en
UJ
CO
6^-
O
io 10
UJ
^ □
•□■
□ -D 22.5°C, 12 Hr Photo-
period 700-1000 ft-c
• • Average of all Conditions
10
12
16
18
20
AGE OF BEAN PLANTS, DAYS
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°.
Botany
153
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
Sugar
Concentration
(mM)
Time to 50%
Separation (Hr)
Petiole
None
— ■
Sucrose
30
Glucose
3
30
Pulvinus
None
—
Sucrose
3
30
Glucose
3
:m
42
42
28
44
44
32
26
24
27
29
30
en
i
. i
>
2
O
h-
q: 20
1 — A
A
<c
W ■ - ■ -
W
q:
lu
</)
$
o
IT)
10
O
h-
LlI
5
1 1
1 1 1
ETHYLENE CONCENTRATION, ^l/l
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.
154
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
down).
/
40
X
SEPARATION,
o
o20
-
o
LU
h- 10
n
1 1 1 1 1
2 4 6 8
GALACTOSE CONC, % IN AGAR
10
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).
Botany
155
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
Light
Temperature, Chrono- Explants
Intensity1
°C logical (Hr)2
Substrate
(Ft-C)
Day Night Age, Days 22.5°C 30°C
Silicate Soil
Conditioners
5,500-5,800
32 24 18 120 60
Soil-Sand
5,500-5,800
32 24 10 96 24
100
20 7 24 120 48
i 12 Hour Photoperiod.
212 Hour Photoperiod, 700-1,000 ft-c.
^Krum (Silibrico Corporation, Chicago, Illinois).
CO
£100
>
<
0 /■ p
^ 80
J /
r /
o
/ $
/ i
b ~«
/E+
G /E / G
< 60
/ /
or
/ /
<
/ /
o_
1 t
a 40
1-
/ P
z
J °/
T /
Sao
—
(T.
r /
LU
/
°- 0
j \ \ \
8 16 24 32 40
HOURS AFTER EXCISION
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).
156
Indiana Academy of Science
4 8 12 16 20
TIME AFTER EXCISION, HR
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).
Discussion
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
C02 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 C02-free atmosphere under 300 ft-c of light abscised at
the same time as those kept in the dark with or without C02, 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, C02, 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 C02 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 C02 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.
Summary
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.
285:1-23.
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.
CELL BIOLOGY
Chairman : Ralph Jersild, Indiana University Medical Center
Edward J. Hinsman, Purdue University, was elected chairman for 1969
ABSTRACTS
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-
161
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-Os04
(KIO), zinc sulfate-Os04, or Os04 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
examination.
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
thickness.
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
families.
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 Cytomembranes1
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 0 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.
167
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.
Results
Glutaraldehyde stabilization of organelles during isolation. Loss of
ultra structural detail in organelles during their isolation was prevented
by adding 101 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 (Dx and D2) and sectioned tangentially (D3). Osmium
post-fixation. X 40,000.
Cell Biology
icj
*>* ^s%*>
170
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.
172
m. -
Indiana Academy of Science
;#
it
m
«
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
173
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 Rf'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 Rf 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
0 12 3 4
HOURS IN 2.5% GLUTARALDEHYDE
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 0 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
ght)
CHC1.3-
■Soluble
Residual
Total
Constituent
— GA
+GA -
-GA
+ GA
— GA +GA
Low Rf Phospholipids
292
346
57
65
349
411
Lecithins
258
244
16
14
274
258
Serine Phospholipids A
78
34
2
T
80
34
Ethanolamine Phospholipids
54
27
7
12
61
39
Serine Phospholipids B
78
20
T
2
78
22
Sterols and Neutral Lipids
27
41
7
14
34
55
Total
787
712
80
107
876
819
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
175
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t
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at
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i%
51l
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
reticulum.
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*
Alanine
++
Phenylalanine
+ +
Arginine
+ +
Threonine
+ +
Aspartic Acid
+ +
Tryptophan
+
Cysteine
++b
Tyrosine
+
Cystine
+ +
Valine
+
Glutamic Acid
+
Imino Acids3-
Glycine
+
Histidine
++
Proline
—
Isoleucine
+ +
Hydroxyproline
■±
Leucine
+ +
Nucleotide Bases0
Lysine
+ +
Adenine
+
Methionine
+ +
Guanine
+
Cytosine
—
Uracil
—
a Determined from loss of ninhydrin reactivity following- paper chromatog-
raphy and from increase in absorbance at 440 m^ during reaction in
solution.
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
177
Glutaraldehyde Cone, %
Figure 12. Inactivation of the enzyme CDP-choline-cytidyl transferase of
homogenates of onion stem stabilized with varying- concentrations of
glutaraldehyde.
(same medium as for organelle isolation) after which glutaraldehyde was
added to a final concentration of 2.5%. After 30 minutes at 0 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 C14-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 C14-uridine, although
obtained from mung bean stem homogenates rather than onion, was
sufficiently low to suggest little or no binding. Radioactivity from C14-
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
Discussion
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 02 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
0 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 CH-Labeled Metabolites to the
12,000 o Particulate Fraction of Onion Stem*.
Metabolite m^Moles/VMole/Mg Nitrogen
Acetate-2-C14 1.8
Choline-2-C14 1.8
Uridine-U-Clib 0.7
Glucose-U-C14 5.5
Leucine-U-C14 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.
Summary
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
155:1672-1674.
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.
177:751-766.
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.
15:283-309.
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 Cytomembranes1
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.
183
184
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 3.1.3.5) with
AMP as substrate and ADPase (EC 3.6.1.6) were performed with the
nucleotides (5 mM) as sodium salts in a medium containing 100 mM
KC1, 5 mM MgCl2 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 — Mg2 — ATPase, the potassium salt of ATP was used in a medium
-it '
, . 4-..
¥
%-V
w
::■■ ,:■
M
V
m
%i:h
n 4
'&.
%. i
*§V
>rl %^x'
f;- %
n%*~
Y
*«IJNi
" ■<%•
®&
$&&*■%
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 MgCl2 and 25 mM Tris
(pH 7.4). The corresponding medium for the Mg3 — ATPase (EC
3.6.1.4) 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
protein.
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
Cations
(^Moles iP/Hour/Mg Protein)
Cell Fraction
Added
I
II
III
Total Homogenate
Mg+ + ,K +
0.39
1.05
0.89
Plasma Membrane
Mg++,K + ,Na+
0.45
1.30
0.99
ANa+
+0.06
+0.25
+ 0.10
Mg+ + ,K +
2.9
15.2
10.8
Mg++,K + ,Na +
5.0
23.2
19.6
ANa +
+2.1
+8.0
+8.8
186 Indiana Academy of Science
table 2. SjJecific Activities of AMPase and ADPase of Rat Liver Cell
Fractions.
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
Mitochondria
4.7 ± 3.1
0.9*
Supernatant
1.2 ± 0.3
1.0*
0.65*
* 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
Mga+-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
inconclusive.
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
187
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
Summary
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
membrane.
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.
Mga+ — ATPase, (Na+' — K+ — Mg2+) — ATPase and 5'-nucleotidase activ-
ity of plasma membranes isolated from rat liver. Biochim. Biophys. Acta
120:369-382.
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
90:126-145.
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.
193:265-275.
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.
77:159-163.
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 Membranes1
J. D. Hall, J. W. Stiles, Y. Awasthi, and F. L. Crane
Purdue University
Abstract
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
structure.
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.
Introduction
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.
189
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
supernatant.
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
Fractions.
Fraction
c
Cytochromes^
Cl b
a+a^
%
Lipid2
Triton-prepared
membranifibrils
DOC-prepared
membranifibrils
0.058
0
0.028 0.020
0 0
0.078
0
14.4
9.6
1 ^moles/g protein.
2 Determined as % lipid
/
mg lipid N
X 100
mg- lipid + nig pr
Cell Biology
191
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
193
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.
194
Indiana Academy of Science
iflffji
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 (P2) by cholate and ammonium sulfate treatment of inner mito-
chondrial membrane which resembles our membraninbril fraction both
chemically and structurally. Their P2 fraction showed no cytochrome aa3
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
P2 preparation to the assay medium.
Membranifibrils are clearly evident in their electron micrographs of
the rutamycin-sensitive ATPase (P2) 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 P2
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.
Summary
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
69:361-374.
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.
243:2405-2412.
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.
236:1680-1688.
CHEMISTRY
Chairman: L. A. McGrew, Ball State University
John W. McFarland, DePauw University, was elected chairman for 1969
ABSTRACTS
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
complex.
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 103 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
199
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)2Br. Bromopentacarbonylmanganese(I) reacts, in an anal-
agous manner, with acetonitrile and benzonitrile to yield the tricarbonyl
complexes Mn(CQ)3(RCN)2Br. 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(CH3CN)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)2Br, 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.
ECOLOGY
Chairman: William B. Crankshaw, Ball State University
Thomas McComish, Ball State University, was elected chairman for
1969
ABSTRACTS
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
areas.
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.
201
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/m2 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/m2, .05 g/m2, .8 g/m2, and 3.1 g/m2. 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
noted.
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
Abstract
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.
Introduction
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.
Methods
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
acres).
Sampling of ground cover was accomplished by establishing four
one-meter square plots near each site. These plots were located 10 meters
204
Ecology
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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-
402.
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
Abstract
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
studied.
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.
Introduction
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).
210
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.
Methods
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
present).
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: D2 =
density per acre; D3 = relative density; B2 = basal area per acre; B3 =
relative basal area; and V3 = importance value =
D3 + B3.
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
Ecology
215
TABLE. 1. Stand Attributes for Jackson Woods, based on trees U" dbh and greater
17.4-6 acres — full tally.
SPECIES
Da
D3
B2
B3
V,
B A /Tree
Fagus grandifolia
14.32
13.13
57.66
51.63
32.38
4.03
Acer saccharum
42.33
38.82
12.67
11.34
25.08
.30
Lirio dendron t u lipifera
11.68
10.71
19.53
17.49
14.10
1.67
Comus florida
13.23
11.08
2.18
1.95
6.52
.16
Ulmus rubra
8.30
7.61
2.67
2.39
5.00
.32
Juglans nigra
3.78
3.47
6.46
5.78
4.63
1.71
Fraxinus americana
3.95
3.62
2.63
2.35
2.99
.67
P?'unus serotina,
3.38
3.10
1.19
1.07
2.09
.35
Nyssa sylvatica
1.72
1.58
1.61
1.44
1.51
.94
Ulmus americana
1.37
1.26
1.26
1.13
1.20
.92
Sassafras albidum
1.55
1.42
.79
.71
1.07
.51
Celtis occidentalis
.86
.79
.67
.60
.70
.80
Quercus alba
.23
.21
.52
.47
.34
2.26
Quercus rubra
.17
.16
.57
.51
.34
3.35
Ostrya virginiana
.57
.52
.1!
.10
.31
.19
Gary a glabra
.23
.21
.44
.39
.30
1.91
Vitis spp.
.34
.31
.04
.04
.18
.12
Juglans cinerea
.17
16
.14
.13
.15
.82
Acer negundo
.11
.10
.16
.14
.12
1.45
Fraxinus quadrangulata,
MU
.06
.10
.09
,08
1.67
Tilia americana
A\
.10
.03
.03
.07
.27
Carya ovata
.0(1
.06
.09
.08
.07
1.50
Robinia pseudoacacia
.06
.06
.09
.08
.07
1.50
Carpinus caroliniana
.11
.10
.02
m
.06
.18
Quercus muehlenhergii
.11
.10
.02
.02
.06
.18
Carya cordifoi mis
II
.10
.02
.02
.06
.18
Asimina triloba
.06
.06
.01
.01
.04
.17
Aesculus glabra
.06
.06
.01
.01
.04
.17
Total
109.03
111.69
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
216
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
woods.
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.
Small
Temporarily-
Small
Large
Ponding
Non-Ponding
Non-Ponding
Species
Sinks
Sinks
Sinks
Ostrya virginiana
3.4
Prunus serotina
3.8
1.7
Cornus florida
6.5
3.0
1.7
Fraxinus americana
7.4
6.6
2,2
Acer saccharum
24.4
17.0
9.6
Fagus grandifolia
54.6
51.8
37.0
Juglans nigra
8.4
21.5
Ulmus rubra
11.6
12.0
Ulmus americana
___
12.0
Liriodendron tulipifera
___
___
2.1
Celtis occidentalis
—
—
L.7
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
220
Indiana Academy of Science
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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
Ecology
223
table 6. Stand Attributes for Potzger Woods, based on trees U" dbh and
6.1+U acres — full tally.
-eater.
SPECIES
D2
D3
B2
B3
V,.
B A/ Tree
Acer saccharum
68.63
56.52
23.93
20.53
38.53
.35
Fag lis grandi folia
14.29
11.77
57.84
49.61
30.69
4.05
Juglans nigra
4.50
3.71
7.44
6.38
5.05
1.65
Fraxinus americana
6.52
5.37
4.51
3.87
4.62
.69
Nyssa sylvatica
2.64
2.17
5.24
4.49
3.33
1.98
Ulmus rubra
3.88
3.20
3.12
2.68
2.94
.80
Liriodendron tiilipifera
2.48
2.04
4.13
3.54
2.79
1.66
Prunus serotina
3.57
2.94
2.44
2.09
2.52
.68
Cornus florida
3.11
2.56
.42
.36
1.46
.14
Ostrya virginiana
2.48
2.04
.45
.39
1.22
.18
Ulmus americana
1.55
1.28
1.00
,86
1.07
.65
Tilia americana
1.24
1.02
.90
.77
.90
.73
Celtis occidentalis
.78
.64
.98
84
.74
1.26
Carya cordiformis
.93
.77
.74
.63
.70
.80
Quercus rubra
.93
.77
.60
.51
.64
.65
Fraxinus pennsylvanica
.78
.64
.66
.57
.61
.85
Fraxinus quadrangulata
.93
.77
.19
.16
.47
.20
Quercus alba
.62
.51
.40
.34
,43
.65
Sassafras albidum
.16
.13
.71
.61
.37
4.43
Aesculus glabra
.62
.51
.24
.21
.36
.39
Carya ovata
.31
.26
.36
.31
.29
1.16
Carya glabra
.16
.13
.19
.16
.15
1.19
Quercus muehlenbergii
.16
.13
.05
.04
.09
.31
Acer rubrum
.16
.13
.04
.03
.08
,25
Total
121.43
116.58
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),
224
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
interpretation.
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.
Summary
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,
respectively.
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
Abstract
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 cm3 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.
Introduction
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
231
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
centimeter.
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.
Entomobryidae
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
Isotomidae
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
Onychiuridae
Onychiurus armatus Tullberg
Poduridae
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
Sminthuridae
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.
Ecology
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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
area.
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
(X2 — 213.96, 3df). The maple-oak area had more species present in the
litter (44) than the other areas. The difference was significant
(X2 = 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 (X2 = 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 (X2 = 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
Ecology
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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
100 — <»
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beech-maple
old field
maple-oak
oak
Collembola
trees
Figure 1. A two-dimensional ordination of the collembolan and tree
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 0 = 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 0 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.
Summary
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
Abstract
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.
Introduction
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, NE1^. 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
241
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
channel.
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
grounds.
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.
296-305.
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
upland.
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
245
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
Forestry.
Procedure
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
virginica.
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
248
Indiana Academy of Science
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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
Ecology
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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
species.
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
corridors.
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.
ENTOMOLOGY
Chairman : Leland Chandler, Purdue University
Jack R. Munsee, Indiana State University, was elected chairman for 1969
NOTES
A Japanese Weevil Discovered in Indiana.
Donald L. Schuder, Purdue University. — An infestation of adult Jap-
anese weevils, Pseudocneorhinus bifasciatus Roelefs, was reported1 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
Museum.
255
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)
257
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
260
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
47%).
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 Fx 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.)
262
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
development.
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-
278.
An Annotated List of the Spiders of Indiana1
Thomas A. Parker, Purdue University
Introduction
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
Station.
266
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
indicated.
3. Pachylomerides audouini (Lucas), 1836
Floyd (Floyd's Knobs, 45); Harrison (Corydon, May 12, 1968,
CB)
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,
TAP); INDIANA (49)
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,
38,39,22)
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,
1836)
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
(49)
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
Cave."
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
existed.
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
incorrect.
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,
TAP)
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,
22)
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,
RWM)
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,
1894.
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
(49)
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,
TAP)
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,
21,22)
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,
TAP)
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
Museum.
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
study.
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,
RWM)
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,
JOS)
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,
JOS)
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,
21,22)
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,
TAP)
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
INDIANA (9)
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
problem.
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,
TAP)
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-
mens.
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,
TAP)
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,
RWM)
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,
21,22)
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);
INDIANA (49)
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,
22)
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
synonymous.
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.
72:1-80.
4. Banta, A. M. 1907. The fauna of Mayfield's Cave. Carnegie Inst. Wash.
67:1-114.
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.
68:1-15.
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.
6:406-422.
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-
222.
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.
62:299-317.
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-
109.
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.
11:103-111.
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.
GEOLOGY AND GEOGRAPHY
Chairman: Lowell I. Dillon, Ball State University.
Wilton N. Melhorn, Purdue University, was elected chairman for 1969.
ABSTRACTS
Gypsum Resources of the Midwestern United States. John H. Cleveland
and Carol F. Tiefel, Indiana State University. — Gypsum (CaS04#2H20)
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
Age.
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.
315
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
Survey.
Analysis of Surficial Landform Properties: The Regionalization
of Indiana into Units of Morphometric Similarity1
Laurence A. Lewis, Temple University
Abstract
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
Indiana.
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
failure.
Introduction
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
suggestions.
317
318
Indiana Academy of Science
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Geology and Geography
319
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-
fluence.
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.
Variables
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.
Analysis
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
321
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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
Eigenvalue
Factors
Factors
1
4.720
36.3
36.3
43.7
43.7
2
2.408
18.5
54.8
22.3
66.0
3
1.601
12.3
67.1
14.8
80.8
4
1.056
8.1
75.3
9.8
90.5
5
1.021
7.1)
83.1
9.5
100.0
C>
0.721
5.5
88.7
7
0.526
4.0
92.7
8
0.359
2.8
95.5
9
0.303
2.3
97.8
10
0.143
1.1
98.9
11
0.112
0.9
99.8
12
0.030
0.2
100.0
13
0.001
0.0
100.0
group
one
-1.556 to
-0.582
group
two
—0.548
to
0.540
group
three
0.556
to
1.538
Geology and Geography
!23
ED -1.165 to -0.539
[ | -0.439 to 0.384
[Ml 0.416 to 1.205
(factor scores)
Figure 2: Areal Magnitude
324
Indiana Academy of Science
□ -0.695 to 0 622
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
groups:
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
phenomena.
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
32G
Indiana Academy of Science
r"
1
<r
r^
,!I£L_^
[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.
Conclusions
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
approach.
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
college.
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.
329
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-
ers).
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 0 hours, 1-9 hours, 10-20 hours, or 20+ hours.
Median semester hours credit of earth science teachers
Earth Science
(1-9)
Geology
(1-9)
Astronomy
(1-9)
Meteorology
(0)
Oceanography
(0)
Chemistry
(10-20)
Physics
(1-9)
Biology
(20+)
Mathematics
(1-9)
Geography
(1-9)
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
(25%).
332 Indiana Academy of Science
Summary
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
science.
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
Abstract
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.
Introduction
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
Survey.
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
333
Indiana Academy of Science
R R 0 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
335
COLLETT
KINDLE
COOPER
ft OTHERS
GALLOWAY
a ST. JEAN
COOPER
a PHELAN
THIS
PAPER
1872
1901
1942
1955
1966
LOUISVILLE- DELPHI
NEW ALBANY
DELPHI
NOT CONSIDERED
DELPHI
NEW ALBANY
BLACK SLATE
SHALE
SHALE
SHALE
SHALE
SELLERSBURG
LITTLE
LITTLE
LITTLE
ROCK CREEK
ROCK CREEK
ROCK CREEK
BEDS
LIMESTONE
LIMESTONE
LIMESTONE?
DEVONIAN
JEFFERSONVILLE
LOGANSPORT
TRAVERSE
LIMESTONE
LIMESTONE
LOGANSPORT
LIMESTONE
LOGANSPORT
LIMESTONE
LIMESTONE
FORMATION
SILURIAN
MIAMI BEND
FORMATION
SILURIAN
SILURIAN
SILURIAN
SILURIAN
SILURIAN
SILURIAN
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.
Correlation
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
Michigan.
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 SW1^ 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 SW1^ 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, NE1^ 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, SW1^ 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
(6).
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
30:261-273.
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
342
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.
Conclusion
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-
tion).
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 Indiana1
Robert R. French, U. S. Bureau of Mines
Abstract
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.
Introduction
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,
Indiana.
348
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 l1/^ 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.
350
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
2y2 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).
1/3
I | 1
| i | i | y\
' ^
y ^s^
X ^s^
150
125
—
s£S^
—
z
o
H-
k 100
—
—
Ul
Q.
</>
5 75
._
OJ
O
ySy
50
—
25
i 1 i
1 1 1 1 1 1
-
10
20 30
MILES
40
50
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
Miles
ton (cents)
Miles
ton (cents)
Miles
ton (cents)
1-5
25
1-10
200
0-5
55
5-10
45
11-30
270
5-10
(55
10-15
70
31-50
290
10-15
80
15-20
95
51-80
350
15-20
100
20-25
120
81-110
390
20-25
120
25-30
145
111-150
430
*See
footnote.
30-35
170
151-200
530
For 30 miles or more, cost is 3.5 cents per ton-mile.
Geology and Geography
351
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
KEY
— Northern Indiana
— Central Indiana
— Southern Indiana
50
100 150
MILES
200
250
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-
tions.
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
8
1
Z 6
O
« 5
Ul
"• 4
or.
3 3
-J
O
Q 2
50 100 150 200 250 300 350
MILES
Figure 3. Some published point-to-point tariffs for water (barge) trans-
port of mineral aggregates.
1
1
1
1
1
1
-
~
-
-
-
•
0
—
—
-
__i— *— -"■
*
••
• •
• •
-
°^~"
-
__L
1
1
•
1
1
1
—
Geology and Geography
353
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,
respectively.
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 ! ~
' A ' ' ' 1
/
s ^> —
- i
l
—
-/
—
/
I ... i ,
i i l i i ■
3 -
2 -
100
200
MILES
300
400
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.
8320.
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
Abstract
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
color.
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.
355
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
357
H
A3 |
C-2
Legend
;i
& <
A- White Oak
B - Red Oak
C - Yellow Poplar
D- Sycamore
E- Black Locust
F- Virginia Pine
I - Control
2- Aspergillus niger Addition
3- Aspergillus fiavus Addition
X- Soil Moisture Test
RF 1-384
A-l
J
L
N
F3
i D-2
TREE PLOTS
SANDSTONE RIDGE
Fig. I
B 3 i
A2
! CI
I
I
I
Dl |
E-3
I F2
F, J
IV;i
Llj*
•— x-1
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
358
Indiana Academy of Science
Legend
A- White Oak
B- Red Oak .
C - Yellow Poplar
D - Sycamore
E- Black Locust
F- Virginia Pine
1- Control
2- Aspergillus niger Addition .
3- Aspergillus flavus Addition
X- Soil Moisture Test
RF h384
TREE PLOTS
SHALE RIDGE
Fig. 2
r.
Di '
X— i
PI
E2
F;3
! D-2
f i
A-3,
C-2
!•■
I
I
T"
B-2
B3
A-2
! CI
•— X-
L
n
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
359
L
Plot I
Plot 2 ,
Plot 3
GRASS PLOTS
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-
lands.
Conclusions
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 Indiana1
Lowell I. Dillon, Ball State University
Abstract
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.
363
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
market.
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
Geology and Geography
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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.
HISTORY OF SCIENCE
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
Abstract
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.
Introduction
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:
370
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
Baccalaureate
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
Doctoral
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
Regions
in Region
in Region
New England
76
95
Middle Atlantic
256
156
East North Central
204
197
Ohio
52
33
Indiana
20
36
Illinois
75
60
Michigan
31
45
Wisconsin
26
23
West North Central
148
86
East South Central
13
51
South Atlantic
73
149
West South Central
70
61
Mountain
49
44
Pacific
69
159
Foreign
55
21
Totals
1013
1019
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
%of
No. of
%of
Area of Genetics
Women
Women
Men
Men
Plant
18
14.2%
258
28.9%
Animal
64
50.4
428
48.0
Microbial
31
24.4
136
15.2
Radiation
3
2.3
10
1.1
Biochemical
11
8.7
46
5.2
Statistics
0
0,0
11
1.2
Biophysics
0
127
0.0
3
892
0.4
Totals
100.0%
100.0%
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
genetics.
Conclusions
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
125:1071-1074.
9. Thtstlewaite, Donald. 1959. College environments and the development
of talent. Science 130 :73.
10. The World Almanac. 1968. Newspaper Enterprise Association Inc.,
Cleveland.
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-
scope.
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.
378
History of Science 379
However, it was also the home territory of Huygen, Leeuwenhoek, and
Onnes.
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-
tarium.
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
heritage.
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
381
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.
Shelley.
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.
Synovtiz
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.
PHYSICS
Chairman: Edwin C. Craig, Ball State University
Richard Conklin, Hanover College, was elected chairman for 1969.
ABSTRACTS
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 me, 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 108m/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
counts/sec.
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
387
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
CI35- 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 CI35 in several chloro-
alkenes, of which virtually no previous study had been made. The in-
ductive effect of -CI, -CCIH2, and -CH3 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 0 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 A0= — 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
Abstract
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 0 eV, 20 0 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.
Introduction
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.
History
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.
389
390
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-
Acceleration
Grid Power ■
• Supplies
Vacuum
System
Photomultiplier
Time-to-Puls
Height
Converter
Preamplifier
Pulse-Height
Figure 1. Block Diagram of Apparatus.
Physics
391
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.
<7>
0-12VDC
Power
Supply
Variable
Transformer
I I I
• I I Excitation Tube
I i I
E>
Impedance
Matcher
G3
-( mh}-
1 (mA
^-Negative^r^
SQ. Wave
Pulse
Generator
D.C. Power
Supply
a
Voltage
Divider
D.C. Power
Supply
( vtvm)
Figure 2. Block Diagram of Excitation Tube Electronics
392
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)
A234:565.
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.
PLANT TAXONOMY
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
Abstract
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.,
396
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
rigidum.
Lepidopilum plebejum has been collected in S and NE South
America.
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
Zealand.
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):
148-151.
3. Dietz, Robert S. 1967. More about Continental Drift. Sea Frontiers 13(2):
66-82.
4. Steere, William C. 1948. Mosses of Ecuador I. The Bryologist 51(3):
65-167.
5. Wegener, A. 1912. Die Entstehung der Kontinente. Geol. Rundsch. 3(4):
276-292.
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.
1-68.
-. 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
press.
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
Abstract
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.
Introduction
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.
406
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.
408
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
juncture.
Discussion
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
411
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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.
4:4-5.
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,
Clark.
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.
413
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
(V).
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
414
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-
served.
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
Co.
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.
SOIL SCIENCE
Chairman : M. L. Baumgardner, Purdue University
James E. Newman, Purdue University, was elected chairman for 1969
ABSTRACTS
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. XA, 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.
417
Tillage Techniques on Indiana Prairie Soil1
Helmut Kohnke and S. A. Barber, Purdue University
Abstract
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.
Introduction
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
Station.
418
Soil Science
419
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.
Treatment
Tillage
Depth
Crop
Residue
No.
Cm
1
Conventional*
20
Corn
Normal
2
Conventional
20
Corn
Stover Removed
3
Conventional
20
Corn
Doubled
4
Field Cultivator
20
Corn
Normal
5
Rototiller
8
Com
Normal
6
None
0
Com
Normal
7
Conventional
20
None
None
8
None
0
None
None
* Conventional tillag-e consists of fall or spring- plowing- followed by
disking.
TABLE! 2. Properties of the soils of the experiment area.
Raub
silt loam
("2" profile)
Bulk
Total
Aeration
Organic
Depth
Density
Porosity
Porosity
Matter
cm.
Texture
g/cc
percent
percent
pH
percent
0-6
Silt loam
6.6
2.7
10-14
Silt loam
6.3
36-40
Silt loam
1.27
52
9.0
(5.4
54-58
Silty clay loam
1.33
50
6.6
6.4
75-79
Clay loam
1.54
42
4 .,8
7.0
96-100
Loam
1.81
31
5.7
8,2
Chalmers silt loam ("8" profile)
Bulk
Total
Aeration
Organic
Depth
Density
Porosity
Porosity
Mattei-
cm.
Texture
g/cc
percent
percent
pH
percent
0-6
Silt loam
6.0
3.0
10-14
Silt loam
6.0
32-36
Loam
1.33
50
8.3
5,7
55-59
Silty clay loam
1.28
52
5.4
5.8
75-79
Clay loam
1.37
48
4.5
6.5
96-100
Clay loam
1.54
42
4.1
7.0
420
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 P205, and
100 lbs. K20 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
cutter.
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
m
160
150
140
130
120
S no
100
90
80
70
Figure 1
Field
Cultivator (4)
^Conventional (I)
xNo Tillage (6)
Indiana
Tippecanoe
County
62 63 64 65 66 67
YEARS
Comparison. of the corn yields of the residue management
plots with those in Tippecanoe County and in Indiana
Soil Science
421
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
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a
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O
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150
140-
130
120
2 3 4 5 6
TREATMENTS
62 63 64 65 66 67
YEARS
Comparison of the effects of treatments with those of
weather on corn yields
422
Indiana Academy of Science
Figure 3
CO
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o
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a.
Q
o
20-
0 ' .1 .2 .3 .4 .5 .6 .7
GRAMS OF DRY ROOT MATERIAL IN 1000 cc OF SOIL
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.
Roots
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.
Treat-
Six
ment
Year
No.
1962
1963
1964
1965
1966
1967
Average
1
146.1
149.7
134.6
140.7
127.7
137.6
139.7
2
147.4
146.7
136.2
147.2
125.6
128.2
138.5
3
146.4
144.1
119.1
152.5
123.1
130.5
136.0
1
143.1
155.0
138.0
153.4
138.5
143.9
143.6
5
145.3
138.2
138.2
156.4
125.6
125.6
138.2
6
133.1
134.3
135.5
143.4
123.8
124.7
132.5
Annual
Average
143.6
144.7
133.6
148.9
124.9
133.1
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
424
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32
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30
29
28
27
or
0 26
25-
24
Indiana Academy of Science
Figure 4
3 4 5 6 7
NUMBER OF TREATMENTS
8
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
experiment.
Sampled May 1968
Treatment
Organic
Matter Percent
1
Conventional, normal residue
2.6
2
Conventional, residue removed
2.4
3
Conventional, double residue
3.0
4
Field Cultivator
2.8
5
Rototiller
3.2
G
No tillage
3.0
7
No corn — tillage
2.2
8
No corn — no tillage
2.2
Soil Science
4L.T>
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
LlJ
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28
t 24
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2 20
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lb
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October 1967
April 1968/
3 4 5 6
NUMBER OF TREATMENTS
8
Effect of treatment and season upon aeration porosity
0-3 inch depth
426
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.
Structure
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
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x
LU
Q
2
Q
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.5
.2-
3 4 5 6 7
NUMBER OF TREATMENTS
8
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.
Average
Daily temperature
fluctuation
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.
Conclusion
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.
><-n
0 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
429
430
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
winter
Brookston-Crosby
as deep as needed
none
8-12 inches
seldom when tiled
Sandstone Shale
shallow 18"-34"
common
less than 4" to AW
frequent
Winter heaving
little except ponded
much
areas
Annual rainfall
36"
46"
Annual run-off
10"
20"
Summer drought
seldom
30 days or more, 80
percent of the years
Topography
level to undulating
rolling to very steep
Percent of land in
93 (Tippecanoe
County)
54 (Lincoln Hills — in-
U.S.-SCS classes
cludes Crawford,
I-IV
Harrison, Spencer and
Perry Counties)
Erosion hazards
little to none
severe
Cation exchange
24-Brookston
12-Zanesville
capacity
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
431
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.
Summary
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, 1963-19651
Russell K. Stivers, Purdue University2
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
Russell.
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.
435
436
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
replications.
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).
TABLE 2.
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.
0
52 105
Inches
0
13.7
— —
149
12.8
19.1 20.7
299
14.8
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-
TABLE 3.
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.
per
A.
Rate of K
in Lbs. per A.
0
52
105
Bu. per A.
0
91.0
. — ,
— .
149
113.3
128.4
127.3
299
119.2
123.8
123.7
Lsd at the 20% level =
9.8
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
TABLE 4.
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.
Lbs.
per
A. of
Percent Composition
Parts per Mill
Kill
P
K
N
P
K
Ca
Mg
Mn
B Cn
Zll
(i
0
2.68
0.20
1.86
0.67
0.35
158
18 17
no
0
299
2.85
.22
2.43
.56
.20
L37
18 13
52
109
0
2.66
.24
1.44
.87
A l
172
17 13
39
109
299
2.91
.26
2.48
.62
.20
191
16 12
45
Low
range
2.46-2.75
.16-.24
1.26-1.
70
.11-.20
.11-20
16-19
2-3 2-5
11-20
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
TABLE 5.
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.
0 5 10 20 30 40
Inches
0 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
moisture.
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
TABLE 6.
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. 0 25 50 75 100 125
Inches
0 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
TABLE 7.
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.
0 66 131
Inches
0 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
TABLE 8.
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.
0 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.
TABLE 9.
Relation of chemical composition of the ear leaf of corn at silking to
fertilizer ti-eatment, Crosby soil, E. R. Brown Farm, Lafayette, Indiana,
1963.
Fertilizer
Treatment
L<bs. per A. of
Percen
t Composition
Parts per
Mill
on
P K
N
P
K
Ca
Mg
Mn
B
Cn
Zn
0 0
2.64
.20
1.09
0.69
0.61
89
15
10
46
0 299
2.58
.18
1.83
0.56
0.34
118
17
9
5 2
135 0
2.68
.29
0.65
1.18
0.90
175
16
12
3 9
135 299
2.72
.27
1.90
0.68
0.39
132
11
9
34
LiOav range
2.46-2.75
.16-.24
1.26-1.70
.11-.20
.11-20
16-19
2-:;
2-5
11-20
Deficiency range
less than —
2.45
.15
1.25
0.10
0.10
15
2
2
10
Summary
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.
ZOOLOGY
Chairman: John 0. Whitaker, Jr., Indiana University
James C. List, Ball State University, was elected chairman for 1969
ABSTRACTS
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 32P uptake by the ovary in 15 minutes.
(2) an increase in 32P incorporation into the RNA fraction of the
ovary.
445
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
and
Indiana State Board of Health
Abstract
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.
Introduction
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:
Number
examined Number rabid Percent rabid
Lasiurus borealis
191
12
6.28
Eptesicus fuscus
177
5
2.82
Tadarida brasiliensis
963
27
2.80
Lasiurus floridanus
717
20
2.79
Lasiurus seminola
846
6
0.71
Pipistrellus subflavus
406
2
0.49
Myotis velifer
273
1
0.37
My otis grisesce?is
281
1
0.36
Myotis lucifugus
984
3
0.30
Myotis austroriparius
1998
1
0.05
447
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.
Zoology
449
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
364
24
6.6
Lasiurus borealis
152
11
7.2
My otis lucifugus
50
1
2.0
(incl. 2 probably this
species)
My otis sodalis
12
0
0
M. keeni
4
0
0
Lasionycteris noctivagans
3
0
0
Nycticeius humeralis
9
0
0
Pipistrellus subflavus
16
2
12.5
Lasiurus cinereus
16
4
25.0
626
•12
6.7
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.
450
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
Allen
13
0
0
Bartholomew
6
0
0
Boone
4
0
0
Brown
2
0
0
Cass
2
0
0
Clark
3
0
0
Clinton
2
0
0
Daviess
1
1
100.0
Dearborn
6
1
16.7
Decatur
1
1)
0
*Delaware
5
0
0
Elkhart
3
0
0
Fayette
1
1
100.0
Floyd
1
0
0
Fountain
2
0
0
Franklin
1
(1
0
Fulton
2
0
0
Gibson
4
1
25.0
Greene
4
2
50.0
Hamilton
1
0
0
Hancock
4
0
0
Hendricks
8
2
25.0
Henry
4
0
0
Howard
6
0
0
Huntington
1
0
0
Jackson
8
0
0
Jefferson
132
13
9.9
Jennings
2
1
50.0
Johnson
51
3
5.9
Knox
2
0
0
Kosciusko
3
0
0
LaGrange
3
0
0
Lake
11
0
0
Laporte
2
1
50.0
Lawrence
8
0
0
Zoology
451
table 2. Rabies in bats collected by citizens of Indiana, summarized
by county, 1965-1968. — (Continued)
No. examined
No. rabid
% rabid
Madison
* Marion
Marshall
Miami
Monroe
* Montgomery-
Morgan
Noble
Pike
Posey-
Putnam
Randolph
Ripley
Rush
St. Joseph
Scott
Spencer
Steuben
Sullivan
Switzerland
*Tippecanoe
Vanderburgh
Vigo
Wabash
Warren
Wayne
Wells
White
7
69
3
2
16
9
5
1
1
4
2
1
2
6
47
3
2
1
4
1
10
95
7
1
1
15
1
1
626
0
0
3
4.3
0
0
0
0
1
6.3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
4.3
0
0
0
0
0
0
o
0
0
0
0
0
10
10.5
0
0
0
0
o
0
0
0
0
0
0
0
i:j;
6.7
* 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).
452
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.)
Total
1966
1967
1968
Total
Rabid
%
January
0
7
1
11
0
0
February
4
2
2
8
0
0
March
1
6
9
16
0
0
April
6
(1)
5
15
26
1
3.8
May
14
13
15
(2)
42
2
4.8
June
12
(2)
30
(3)
32
74
5
6.8
July
10
(1)
80
(12)
60
(3)
150
16
10.7
August
42
(2)
12
(7)
39
(2)
205
11
5.4
September
11
(3)
40
(1)
7
58
5
8.6
October
4
(1)
5
(1)
9
2
22.2
November
5
8
13
0
0
December
6
8
14
626
0
42
0
115
(10)
328
(24)
183
(8)
6.7
%
8.6%
7.3%
4.4%
the fall (September to November) 7 of 80, or 8.8% were rabid, in the
winter (December to February), 0 of 33, and in the spring (March to
May), 3 of 84, or 3.6% were rabid. Thus, bat rabies in Indiana has been
occurring in the summer and fall.
Change in rate of occurrence of disease through years of study
The overall rate of infection in 1966 was 8.6% (10 of 135 infected);
in 1967 it was 7.3% (24 of 328), and in 1968 it was 4.4% or 8 of 183.
Thus it would not appear that the disease has been increasing in Indiana
bats over the past three years. These data would suggest a decrease
over the period, but the observed difference was not significant (Chi-
square = 2.33, 2 df).
Marion, Johnson, St. Joseph, Vanderburgh, and Jefferson Counties
Five counties were studied intensively (Table 4). There were rabies
"outbreaks" in 1967 in big brown bats in Jefferson County and in red
bats in Vanderburgh County. Both outbreaks were publicized, hence
Zoology
4 5:
numerous bats were submitted to health authorities, and we have rather
large citizen samples from these counties. Also we have a rather large
citizen sample from Marion, Johnson, and St. Joseph Counties. These are
counties in which rabies is present but in which there was no major
publicity.
Jefferson County is in southeastern Indiana. In 1967, 11 of 93 bats,
or 11.8% were found to be rabid. In 1968, 2 of 36, or 5.5% were rabid.
All but one of the rabid bats from this county were big brown bats, and
all were from the city of Madison. None of the 16 red bats, Lasiurus
borealis, taken there were rabid.
In Vanderburgh County, seven of the ten rabid bats were red bats,
Lasiurus borealis, but only thirteen big brown bats, Eptesicus, were
table 4. Rabies in bats in Vanderburgh, Jefferson, Marion, Johnson, and
St. Joseph Counties.
Marion,
Johnson,
Vanderburgh
Jefferson
St.
Joseph
No.
Rabid %
No.,
Rabid %
No. Rabid %
L. borealis
61
1
11.5
L6
0
0
27
2
7.4
L. cinereus
9
1
11.1
0
0
0
4
1
25.0
E. fuscus
13
1
7.7
102
12
11.8
96
4
4.2
Other
12
I
8,3
14
1
7.1
33
1
3.0
95
10
10.5
132
13
9.8
160
8
5.0
1966
2
0
0
3
0
0
42
4
9. ,5
1967
67
9
13.4
93
11
11.8
69
2
2.9
1968
26
1
3.8
36
2
5.6
49
2
4.1
95
10
10.5
132
18
9.8
169
8
5.0
January
0
0
0
0
0
0
4
0
0
February
0
0
0
0
0
0
1
0
0
March
1
0
0
3
0
0
6
0
0
April
1
0
0
1
0
0
9
0
0
May
3
0
0
4
0
0
13
1
7.7
June
6
0
0
18
1
5.6
L5
1
6.7
July
:m
4
11.8
5!)
10
16.9
25
1
4.0
August
43
5
11.6
34
2
5.9
55
3
5.5
September
7
1
14.3
14
0
0
20
2
10.0
October
0
0
0
1
0
0
3
0
0
November
0
0
0
0
0
0
0
0
0
December
0
0
0
0
0
0
3
0
0
95
10
10.5
132
13
9.8
160
8
5.0
454 Indiana Academy of Science
taken, of which one was rabid. The total number of bats examined from
Vanderburgh County was 95.
Thus the bat rabies situation was much different in Jefferson and
Vanderburgh Counties, with red bats being the main species infected in
Vanderburgh County, and big brown bats being the main species infected
in Jefferson County. Hopefully more red bats from Jefferson County and
more big brown bats from Vanderburgh County will be taken in the
future.
Marion, Johnson, and St. Joseph Counties were counties in which bat
rabies has been found, but where this fact apparently was not widely
publicized, thus there was no major effort by citizens to turn in bats.
However, large numbers of bats were submitted from these counties, and
data from them, taken collectively, was considered as a control, for com-
parison with data from the two counties with publicized rabies. In the
three counties, eight of 160 bats, or 5.00% were rabid, as compared to 23
of 227 (10.1%) in Jefferson and Vanderburgh Counties. This difference
was not significant (Chi-square = 3.35, 1 df).
Relation between Rabies in the "Citizen Sample" and the "Wild Sample"
It was hypothesized that if the bats were being found by the citizens
because they were "sick" from rabies, then the incidence of rabies should
be higher in the citizen sample than in the wild simple. On the other
hand, if the bats were not affected by the virus, then rabies should occur
in the wild sample and the citizen sample at about the same rate. (In
this latter case, they could be acting as carriers of rabies in that they
harbor the virus, yet do not get the disease themselves, as has been
suggested. If this were the case then bats might constitute a natural
source of infection from which other species might contract the disease.)
For comparison with the citizen sample, 133 bats were collected from
seven colonies during the height of the 1967 rabies outbreak in Jefferson
County (97 in six colonies in the city of Madison, some in the same
buildings from which rabid bats had originated, and 36 bats from a
locality 15 miles southwest of Madison.) None of these bats was rabid.
The above hypothesis was not supported. However, it is possible that
rabies is found only in certain colonies, and that the wrong colonies were
sampled. The possibility that bats leave the colonies when they
become rabid, and that they are more likely to come into contact with
humans, when rabid, seems a probable explanation of the data at this
time.
Hopefully, future laboratory studies can be undertaken to determine
whether individual bats become sick or die when they contract the disease,
and if so, how long this takes.
Relation of Rabies in Bats to Rabies in Other Species in Indiana
If there is a relationship between rabies in bats and rabies in other
species of mammals as has been suggested, then it would seem that
there should be a positive association between areas with bat rabies and
Zoology 455
those with rabies in other spec-ies. If there is no such relationship, then
it would seem that incidence of rabies in other species would be about
the same in counties with bat rabies and counties without bat rabies.
Since the greatest number of cases of bat rabies occurred in 1967, and
since the Indiana State Board of Health has summarized information
concerning rabies in all species in 1967, data from that year were
examined for relationship between bat rabies and rabies in other species.
In the six counties with bat rabies in 1967, a total of 949 individuals
other than bats were examined. Five of these, or 0.5% were found to be
rabid. From all other counties, a total of 2369 individuals was examined,
of which 58, or 2.45% were rabid.
This would lead one to believe that, in Indiana, there is no positive
correlation between rabies in bats and rabies in other species. In fact,
one could draw the reverse conclusion, that there is a negative correla-
tion between bat rabies and rabies in other species, since significantly
more rabid animals were found in other species in those counties not
reporting bat rabies than in those with bat rabies in 1967 (Chi-
square — 13.18, 1 df). Unfortunately, the counties in which numbers of
bats have been examined are not always the same counties in which
adequate numbers of other species have been examined.
Information from some of the critical counties is as follows: in
Jefferson County rabies was not found in the 27 other mammals examined
in 1967; in Vanderburgh County, three of 193 other mammals examined
were rabid (1.55%). In Marion County, in which 3 of 67 bats over the
past three years have proved rabid, no rabies cases were reported in 537
mammals examined in 1967. Other than Vanderburgh, the major rabies
counties for other species in 1967 were Daviess, Knox, Lake, and Shelby,
each having at least three rabid animals other than bats in 1967. Thus,
it would be instructive to examine large numbers of bats from these
counties, but a total of only five was received in 1967.
Twelve of 168 bats (7.1%) v/ere rabid from the 44 counties with
rabies in other species in 1967, while in the counties not reporting rabies
in other species, 22 of 160 bats (13.8%) were rabid, but this difference
was not significant (Chi-square = 3.44, 1 df).
As a further test, the data from the counties adjoining the two major
bat rabies counties, Jefferson and Vanderburgh, were summarized and
compared to the data from the remainder of southern Indiana. In
Vanderburgh, Posey, Gibson, Pike, and Warrick Counties, 243 individuals
other than bats were examined, of which 3 or 1.23% were rabid. In Jef-
ferson, Scott, Clark, Jennings, Ripley, Dearborn, Ohio, and Switzerland
Counties, a total of 188 individuals of other species were taken, of which
8, or 4.2% were rabid. In the remaining southern Indiana counties, 379
individuals other than bats were examined, of which 24, or 6.3% were
rabid, thus again indicating lack of association or even negative associa-
tion between bat rabies and rabies in other species.
456 Indiana Academy of Science
Although important data are lacking, one can only conclude that
there is no positive correlation between bat rabies and rabies in other
species in Indiana.
Thanks to Mr. Robert Kerr, the Jefferson County Sanitarian, we
have relatively complete information concerning the status of many of
the Jefferson County bats when collected. Of 109 bats collected 17, or
15.6% were dead when taken. Among the 96 non-rabid bats, 14 or 14.6%
were dead, while among the 13 rabid bats, 3 or 23% were dead.
It is recognized that there are certain problems inherent in data of
the kind that have been collected during this study. For example, in the
comparison between bat rabies and rabies in other species, most of the
bats were from cities, while many of the specimens of wild species may
be from rural areas, but there is no way to check this. We have no actual
or relative information concerning the size of the bat populations in the
various counties. Data of this sort would be very valuable. Most
important, none of the data were collected in a truly random fashion.
Literature Cited
1. Anonymous. 1968. Bat Research News. 9(1) :8.
2. Burns, K. F., C. F. Farinocci, T. G. Murnone, and D. F. Shelton. 1956.
Insectivorous bats naturally infected with rabies in southwestern United
States. Amr. J. Public Health 46:1089-1097.
3. Girard, K. F., H. B. Hitchcock, G. Edsell, and R. A. MacCreadt. 1965.
Rabies in bats in southern New England. New England J. Med. 272:75-80.
4. Pearson, E. W., and T. R. B. Barr. 1962. Absence of rabies in some bats
and shrews from southern Illinois. Trans. 111. State Acad. Sci. 55:35-37.
5. Schneider, N. J., J. E. Scatterdat, A. L. Lewis, W. L. Jennings, H. D.
Venters, and A. V. Hardy. 1957. Rabies in bats in Florida. Amr. J. Public
Health 47:983-989.
6. Sullivan, T. D., J. E. Grimes, R. B. Eads, G. C. Menzies, and J. F. Irons.
1954. Recovery of rabies virus from colonial bats in Texas. Public Health
Reports 60:766-768.
7. Venters, H. D., W. R. Hoffert, J. E. Scatterdat, and A. V. Hardy. 1954.
Rabies in bats in Florida. Amer. J. Public Health 44:182-185.
8. Verts, B. J., and T. R. B. Barr. An effort to identify rabies in bats from
northwestern Illinois. Cornell Vet. 51:384-387.
Preference for Toxic and Non-toxic
Artificial Sweeteners in Rodents1
Mahlon W. Wagners and W. C. Gunther
Valparaiso University
Abstract
Four species of deer mice (Peromyscus) were given choices of glucose
and three artificial sweeteners: sodium saccharin and sodium or calcium
cyclamate. Preferences for glucose vs saccharin were based on relative
sweetness, with the sweeter of a pair being- chosen. However, Ss preferred
glucose to the cyclamates regardless of relative sweetness. Evidence is
presented to suggest that aversion to the cyclamates may be based either
on noxious stimuli associated with ingestion (according to recent toxicity
studies of cyclamates) or on taste qualities in addition to sweetness that
may be present in the cyclamates. Therefore, learning and physiological
factors as well as taste are implicated in food preferences.
Introduction
Artificial sweeteners have been available as sugar substitutes for
diabetics and dieters for many years. While saccharin has a bitter after-
taste to humans, it has been used extensively as an effective reward in
many animal learning studies since 1933 (5). Squirrel monkeys, however,
do not prefer saccharin (2). New artificial sweeteners, the cyclamate
salts, were introduced in 1950 and rapidly surpassed saccharin in popu-
larity, accounting for sales of $25 million in 1961. Perhaps because of the
newness of the cyclamates, few studies with animals have been reported:
Murray et al. (8) found rats to prefer water to all concentrations of Na
cyclamate, but Smith and Ross (15) found C57 black mice to prefer
maximally a 1% Na cyclamate solution to water.
The basis for sweet preference is usually ascribed to innate and /or
learning factors. Glucose is sweet and directly utilizable for energy by
the cells. These sweet preferences can be "unlearned" when paired with
noxious stimuli (12). Nees and Derse (11) and Japanese researchers
recently reported that cyclamates fed to rats and mice resulted in infer-
tility, laxative effects and fetal deaths. It may be hypothesized that these
toxic effects would result in a preference change similar to that reported
above when sweet compounds were paired with noxious stimuli, although
the chemical need not be ingested in sufficient quantities to produce toxic
effects, since Nachman (9, 10) has shown laboratory rats to avoid LiCl
once having experienced ill effects and Fregly (3) has found adult rats
to avoid LiCl although never having experienced toxic effects. Gunther
(1), on the other hand, finds that lactose fed to early posthatching chicks
resulted in many deaths and that apparently there was no preference
change to the less noxious stimuli available. Kare (6), however, reports
that chicks and adult chickens are indifferent to lactose, suggesting an
inability to discriminate a lactose solution from water.
1 This study was supported by a National Science Foundation Institu-
tional grant and a Valparaiso University research grant to the first author.
2 Now at State University College, Oswego, N. Y. 13126
457
458 Indiana Academy of Science
It was the purpose of this study to examine the preferences of sev-
eral rodent species for artificial sweeteners either known to be non-toxic
or suspected of having toxic effects.
Materials and Methods
Four species of deer mice — males and females of Peromyscus
leucopus (N=7 and 4, mean weight 23.5 and 21.6 gm.), Peromyscus
eremicus (N=4 and 3, mean weight 18.5 and 17.9 gm.), Peromyscus
maniculatus bairdii (N=2 and 3, mean weight 20.4 and 15.5 gm.), and
Peromyscus floridanus (N=3, mean weight 33.0 gm.) — were supplied by
John A. King at Michigan State University.
Sweet substances used were anhydrous glucose, sodium saccharin,-**
and sodium and calcium cyclamate.4
All &s were housed individually with standard (Big Red) laboratory
chow always present and water available except during testing. Using a
paired-comparison procedure, each S was presented with pairs consisting
of a 4, 8, or 16% glucose solution and either a 0.23% saccharin, 0.91% Na
cyclamate or a 1.81% Ca cyclamate solution. These particular concentra-
tions were chosen since Stellar (16) reported rats to prefer maximally a
0.23% Na saccharin solution, and Smith and Ross (15) found mice to pre-
fer a 1% Na cyclamate solution. Saccharin-glucose pairs were presented
between July 3 and 24, 1967. Glucose — 0.45% Na cyclamate pairs were
presented for 11 days from July 25 to August 5 under the mistaken
assumption that saccharin might be only twice as sweet as cyclamate.
(However, Schutz and Pilgrim (14) report that saccharin is ten times
sweeter than Ca cyclamate to humans, when both are paired with
sucrose.) The Na cyclamate concentration was doubled to 0.91%,
approximating that of Smith and Ross, and paired with glucose for 15
days from August 7 to 22. For three days 0.91% Ca cyclamate was paired
with each of the three glucose solutions. Because of the continuing
rejection of the artificial sweetener the concentration was again doubled
and the 1.81% Ca cyclamate-glucose pairs were presented for 15 days
from August 26 to September 9, 1967. Due to the rather preliminary
nature of the use of 0.45% Na cyclamate and 0.91% Ca cyclamate solu-
tions, the intake for these pairs is not included in the data to be reported.
Each glucose-artificial sweetener pair was repeated five times on a
randomly cycled schedule. Pairs of solutions were presented for two
hours per day. Solution positions were altered randomly to inhibit devel-
opment of drinking patterns based on position rather than on sweetness.
Analysis of variance was computed on intake corrected for body weight
since there were wide variations in weight across species. The data are
presented graphically as percent glucose preference, which is a method
that not only takes body weight into account but also, by examining their
relative preference rather than their absolute intake differences, tends to
equate preferences for animals who drink small or large amounts.
3 The sodium saccharin was provided by the Monsanto Chemical
Company.
* The cyclamate salts were provided by Abbott Laboratories.
Zoology
459
Results
An analysis of intake uncorrected for body weight showed that
Peromyscus floridanus drank more and showed a greater absolute degree
of preference and that P.m. bairdii males generally drank the least and
showed indifference to the choices. (The greater intake by P. floridanus
is probably linked to their greater size.) The fact that these generaliza-
tions are maintained in the percent-preference data attests to the stabil-
ity of the correction factor used. All further data presented and analyzed
are on the basis of the intake corrected for body weight rather than on
the basis of raw intake.
The preference data are presented differently in Figure 1 from
Figures 2 and 3 because for saccharin there are clear differences in
preference depending on the glucose concentration. This glucose depend-
ence is not seen for the cyclamates since in all cases glucose was chosen
to a very great extent over the cyclamates regardless of concentration
(the exception in all cases being P.m. bairdii males).
Na Saccharin
Glucose vs. .23% Saccharin
—4% glucose — 8% glucose — 16% glucose
ioo-,P leucopus-male - Pleucopus-female .. P floridanus
80
60-
40-
20
/ A--.
1 2345 I 2345
12 3 4 5
)o-jPm. bairdii- male
/ \ / \
/
-i — — i 1 r
2 3 4 5
Pm.bairdii-female
/
/
i
v — *>
p
sremicus
-male
o-
. 0 o"*'
"X
DT-
.0.
*">.a o
Q-*
o —
—° -o
T 1 1 1
12345 12345
TRIALS
Peremicus- female
t 1 i r— r
12 3 4 5
Figure 1. Relative Intake of Glucose and Sodium Saccharin in Peromyscus.
460 Indiana Academy of Science
Only for the saccharin-glucose choice did most of the species prefer
the artificial sweetener (See Figure 1). When given the 4% glucose vs.
0.23% saccharin all subjects preferred saccharin (F=15.15; df=l/6;
p<.001), with P. fioridanus showing the greatest preference and P.m.
bairdii males being indifferent. P.m. bairdii females showed preferences
similar to P. eremicus. P. eremicus showed a greater corrected intake
than P. leucopus (F=17.92; df=l/19; p<.005) and a greater saccharin
preference (F=81.0; df=l/9; p<.001). No sex differences were found for
these last two species but sex differences were significant for P.m. bairdii
(F=16.76; df=l/4; p<.025).
When 8% glucose was paired with saccharin, P. fioridanus developed
a marked glucose preference, P. eremicus and P.m. bairdii females
showed a saccharin preference, and P. leucopus and P.m. bairdii males
showed no consistent preferences (F=5.58; df=6/24; p<.001). Again,
no sex differences were found for P. leucopus or P. eremicus although by-
examining these two species separately preference differences were found
(F=5.43; df=l/9; p<.05). For P.m. bairdii sex was significant
(F=17.76; df=l/4; p<.025).
All Ss preferred 16% glucose to saccharin, and it is obvious from
graphical inspection that there are no significant species differences.
Na Cyclamate
The various glucose concentrations were preferred to cyclamate
except by P.m. bairdii males, who were inconsistent in their preferences
as is seen in Figure 2. Generally, P. fioridanus showed a greater degree
of preference, and when these data are coupled with the P.m. bairdii
male drinking pattern, there is produced a significant species X sweet-
ener interaction (F=31.78; df=6/24; p<.001) for 16% glucose vs. Na
cyclamate. It is also seen in Figure 2 that the 8% glucose combination
produced a similar difference in species preference. P. eremicus and
P. leucopus exhibited no species or sex-based differences in preference
for glucose. P.m. bairdii sex differences were found for the 8%
glucose combination (F— 29.24; df— 1/4; p<.01) and for the 16% glucose
combination (F=30.37; df=l/4; p<.01).
In spite of the pronounced glucose preference, the mice still man-
aged a total solid intake of from 0.04 gm Na cyclamate for the P.m.
bairdii to 0.07 gm for P. eremicus females, to 0.08 to 0.10 gms for the
remaining mice.
Ca Cyclamate
As is seen in Figure 3, for all pairs of solutions, P. fioridanus drank
more and showed the greatest glucose preference, P.m. bairdii males
were inconsistent or preferred Ca Cyclamate, and the remaining animals
were intermediate (for 4% glucose, F=85.0; df=6/24; p<.001, for 8%
glucose, F=63.0; df=6/24; p<.001, and for 16% glucose, F=6.37;
df=6/24; p<.001). The only sex-based intake differences in preference
for P. eremicus and P. leucopus were found when the choice was a 4%
glucose solution or Ca cyclamate. P. eremicus females maintained a
Zoology
461
consistent glucose preference while the other Ss showed an initial glucose
preference changing to indifference (F=3.28; df=4/4; p<.05). P.m.
bairdii showed consistent sex differences for all three pairs of solutions
(for 4%, F=14.91; df=l/4; p<.025, for 8%, F=122.1; df=l/4;
p<.001, and for 16%, F=118.6; df— 1/4; p<.001).
At the conclusion of the exposure to the three artificial sweeteners
and glucose, the mice had a total solid cyclamate consumption of from
0.15 gms for P.m. bairdii to 0.21 gms for P. leucopus males.
If possible toxic effects would cause an increased aversion to cycla-
mates with extended experience, such aversion should be seen as the
significant effect of "days" or "days by sweetener." Significant "day"
and "day by sweetener" effects were found for 16% glucose vs. Na
cyclamate and 4% glucose vs. Ca cyclamate, and "days" were significant
for 8% glucose vs. Ca cyclamate. However, the significances were not
caused by a consistent decline in cyclamate preference, but rather by
marked day-to-day variation in sweetener preference by the several
species. In addition, the already low initial preferences for cyclamates
Glucose vs .91% Na Cyclamate
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Figure 2. Relative Intake of Glucose and Sodium Cyclamate in Peromyscus.
462
Indiana Academy of Science
makes it difficult for them to decrease to any appreciable extent. Conse-
quently, an unequivocal answer is not provided to the hypothetical effects
of toxicity on preference.
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Zoology 463
Discussion
The species differences found here present further evidence of the
more general phenomena referred to in the introduction (for species
other than deer mice) and also reported for glucose preferences in deer
mice (21). At present these differences cannot be explained in terms of
the differing natural habitats. P. floridanus and P. eremicus live in xeric
(dry) areas and P. leucopus and P.m. bairdii live in mesic (moist) areas,
but the sweet preferences of these species living in similar areas are not
consistently similar. In fact, there often seem to be similar preferences
between species of quite different natural habitats. This phenomenon was
also found for glucose-glucose preferences by Wagner and Rowntree (21).
The pronounced sex differences in preferences for glucose vs. sac-
charin in laboratory rats recently noted by Valenstein (18, 19) were not
found for P. leucopus or P. eremicus, although P.m. bairdii also showed
consistent sex differences in preferences for all three artificial sweeteners.
However, where Valenstein found males to switch to glucose and females
to continue preferring saccharin, here the male P.m. bairdii remained
indifferent to or preferred the artificial sweetener while the females
switched to a glucose preference. It is difficult to evaluate the species
differences in preference for Na cyclamate in the other studies (8, 15)
since the rats and mice were given a choice of cylamate and water and
the rats were partially food deprived. As Stellar and McCleary (17) and
Wagner and Rowntree (20) have emphasized, the type of choice available
determines the preference (as most obviously does the presence of
hunger).
The strong glucose preferences over the cyclamates are very inter-
esting, especially when it is realized that the 1.81% Ca cyclamate is
approximately as sweet as a 16% glucose solution (to these writers).
This means that preferences (or aversions) for cyclamates are probably
not based on sweetness per se. There is also literature suggesting that
sugar of lead (lead acetate), while sweet, is avoided by rats due to its
toxicity. However, this research is somewhat suspect in suggesting that
some sweets are avoided; Mason and Safford (7) have reported that
although this compound may be sweet it also has strong taste components
of bitter and astringent or metallic which probably produce the aversion.
While these taste components have not been reported for the cyclamates,
many persons do report anecdotally a different, sort of "thin" or
"metallic" character in the taste of artificially sweetened beverages.
It is not possible to show definitively that the cyclamate aversion in
deer mice is based on toxicity or physiological discomfort. Nees and
Derse (11) fed rats for one year on diets with 5% and 10% Ca
cyclamate added. This amounted to between 312 gms and 780 gms of
total solid cyclamate ingested or from slightly less than one gram to
slightly more than two grams per day for females and males respectively.
(These animals, it should be noted, were not given a choice of an alter-
native unadulterated diet.) With animals averaging over 400 gms this is
equivalent to an intake of less than 0.1 gms in a 20 gm mouse. Addi-
tional recent findings by Tanaka in Japan (personal communication and
464 Indiana Academy of Science
translation from Nees) suggest that an acute LD50 for 20 gm mice is
.10-.20 gm Na cyclamate and, further, that a more sensitive measure of
fetal deaths, FLD50, is 0.0036 gm! On the basis of these two studies it is
quite possible that the deer mice could have taken sufficient cyclamate to
experience toxic effects and could have developed an aversion to these
solutions.
However, it is not even necessary to show that the animals are
experiencing marked physiological discomfort in learning an avoidance
since Fregly (3) found adult rats to avoid all concentrations of poisonous
LiCl, even those too dilute to produce discomfort; Rozin (13) also found
half -wild and domestic rats to subsequently avoid a Lithium-poisoned diet
for a long time after having experienced the ill effects only once.
This study points up the need for more research to separate prefer-
ences based on taste or other receptors. Toxic sweet substances such as
xylose, ribose, and dulcin may give clues to the basis for preferences or
aversions. In addition, the study of various species and sex differences is
needed to understand further the basic aspects of taste.
Literature Cited
1. Eichelkraut, Nan, and W. C. Gunther. 1968. Sugar preference among
heat-stressed chicks. Proc. Indiana Acad. Sci. 77:413.
2. Fisher, G. L., C. Pfaffmann, and E. Brown. 1965. Dulcin and saccharin
taste in squirrel monkeys, rats and men. Science 150:506-507.
3. Fregly, M. J. 1958. Specificity of the sodium chloride appetite of
adrenalectomized rats: Substitution of lithium chloride for sodium
chloride. Amer. J. Physiol. 195:645-653.
4. Garcia, J., D. J. Kimeldorf, and R. A. Koelling. 1955. Conditional aver-
sion to saccharin resulting from exposure to radiation. Science
123:157-158.
5. Hausmann, M. F. 1933. The behavior of albino rats in choosing foods:
II. Differentiation between sugar and saccharine. J. Comp. Psychol.
15:419-428.
6. Kare, M. R. 1961. Comparative aspects of the sense of taste, pp. 6-15.
In : The Physiological and Behavioral Aspects of Taste. Univer. Chicago
Press.
7. Mason, D. J., and H. R. Safford. 1965. Palatability of sugar of lead.
J. Comp. Physiol. Psychol. 59:94-97.
8. Murray, E. J., H. Wells, M. Kohn, and N. E. Miller. 1953. Sodium
sucaryl: a substance which tastes sweet to human subjects but is
avoided by rats. J. Comp. Physiol. Psychol. 46:134-137.
9. Nachman, M. 1962. Taste preference for sodium salts by adrenalec-
tomized rats. J. Comp. Physiol. Psychol. 55:1124-1129.
10. . 1963. Learned aversion to the taste of lithium chloride and
generalization to other salts. J. Comp. Physiol. Psychol. 56:343-349.
11. Nees, P. O., and P. H. Derse. 1967. Effects of feeding calcium cyclamate
to rats. Nature 213:1191-1195.
12. Revusky, S. H., and E. W. Bedarf. 1967. Association of illness with prior
ingestion of novel foods. Science 155:219-220.
Zoology 465
13. Rozin, P. 1968. Specific aversions and neophobia resulting from vitamin
deficiency or poisoning- in half-wild and domestic rats. J. Comp. Physiol.
Psychol. 66:82-88.
14. Schutz, H. G., and F. J. Pilgrim. 1957. Sweetness of various compounds
and its measurement. Food Research 23:206-213.
15. Smith, M. P., and S. Ross. 1960. Acceptance of sodium sucaryl by C57
black mice. J. Genet. Psychol. 96:101-104.
16. Stellar, E. 1967. Hunger in man : Comparative and physiological studies.
Amer. Psychologist 22:105-117.
17. Stellar, E., and R. A. McCleary. 1952. Food preference as a function of
the method of measurement. Amer. Psychologist. 7:256.
18. Valenstein, E. S., V. C. Cox, and J. W. Kakolewski. 1967. Further
studies of sex differences in taste preferences with sweet solutions.
Psychol. Reports 20:1231-1234.
19. Valenstein, E. S., J. W. Kakolewski, and V. C. Cox. 1967. Sex differences
in taste preference for glucose and saccharin solutions. Science
156:942-943.
20. Wagner, M. W., and J. T. Rowntree. 1966a. Methodology of relative sugar
preference in laboratory rats and Deer mice. J. Psychol. 64:151-158.
21. . 1966b. Comparative sugar preference in various rodents.
Amer. Zoologist 6:536.
Melanoma in Helodernia suspectum Cope
Robert H. Cooper, Ball State University
In August of 1945 two gila monsters were received from Arizona
expressed in a wooden box 24" x 16" x 12". The animals have been
living in this same container continuously since that time, except for the
last few months, when one of them was removed because of surgery on
the other. The animal under discussion measures 497 mm from tip to
tip and 335 mm from tip of nose to vent. This is almost the exact size
that it was 23 years ago.
Approximately once a week the specimen has received one raw egg
or some newly hatched English sparrows or some newly born mice and
rats.
In 1963 a projection began to show on the dorsal surface at the
base of the tail. By 1968 this tumor was approximately 51 mm in width,
35 mm in length from front to back, and 20 mm in depth. In order to
try to prevent further migration of the growth, surgery was performed
on the morning of March 9, 1968, by a competent Muncie surgeon. Since
the growth was of the type that had migrated through the tissue, the
entire dorsal surface was removed, including the fascia. The specimen
was immobilized for the surgery.
The tumor was taken to the Ball Memorial Hospital Pathology Lab-
oratories. Macroscopic and microscopic studies were made of the tissues
and cells. Sections were made and stained in order to make cellular stud-
ies, and malignancy was determined. Kodachrome 35 mm slides were
made of the gross structure and microscopic structure. Kodachrome
views of the incision were made from time to time.
Dr. Montgomery and Dr. Branam point out that "The tumor has no
particular arrangement and occurs simply in multiple nodules separated
from each other by bands of connective tissue. Here and there, there are
hyalinized fibrous areas intimately associated with some tumor cells.
The individual cells are eliptical-oval or spindle shaped. They have a
relatively large amount of brownish pigment in their cytoplasm, and
inspection by oil power magnification revealed that many contain very
large nuclei as compared to the nuclei of the normal pigment-bearing
cells in the epidermis and dermis, and revealed that these nuclei contain
increased amounts of chromatin material and exhibit an irregular bead-
ing of the inner surface of the nuclear membrane, and considerable vari-
ation in their configuration. These are features of malignant cells,
usually. On the other hand careful inspection of many fields shows no
mitotic figures.
"This particular lesion was called a melanoma in the first analysis
because it is a neoplastic growth of nevus cells producing pigment, that
exhibits abnormal cells with abnormal nuclei, and demonstrates invasion.
Since the lesion was large it was impossible to embed all of the material,
466
Zoology 467
and it was assumed that there must be a junctional component some-
where along the line. However, this has not proven to be the case, and
for this reason the lesion might best be called a cellular blue nevus, with
cytologic features suggesting malignancy. " The tumor and microscopic
sections are on file at the Ball Memorial Hospital laboratory.
Contact had been made with Dr. Herbert L. Stahnke, Director of the
Poisonous Animals Research Laboratory at Arizona State University at
Tempe, Arizona, who is a specialist in working with gila monsters and
scorpions, to see whether there was any information he could give as to
anesthesia and post-operative treatment. Dr. Stahnke had used barbitu-
ates, injecting approximately % cc. every 15 to 20 minutes. He had also
used fluothane in small amounts, watching the animal very carefully
and removing it from the gas chamber upon loss of equilibrium.
Dr. Sherman Minton, Professor of Bacteriology at Indiana Univer-
sity Medical School and an authority on reptiles, was also interviewed and
mentioned the use of chilling at 5° centigrade for one or two hours which
he had used on cobras, and then being able to work five or ten minutes
before the specimen would become active. Dr. Minton mentioned that
chloroform had not been satisfactory with the reptiles, but that he is
trying fluothane with success.
Two days after surgery the animal was offered a raw egg and ate it.
From that time on a raw egg was broken into a culture dish and fed
to the specimen approximately once a week. No healing seemed to take
place for some weeks.
During the last part of April it was suggested by Dr. Minton that
heat be added to the extent of keeping the temperature pretty close to
80°. This has been done and a scab has formed and come off. On May
22 three drops of "mycolog" ointment were smeared over the incision.
This was at the recommendation of a local veterinarian and has served
as a control for fungus and bacterial growth.
Feeding and "mycolog" treatment are continuing and new tissue is
forming. Shedding has taken place over the entire animal. The surgeon
believes this increased activity to be due to the stimulus from the
replacement of tissue over the incision.
Acknowledgment is due to Ralph 0. Butz, Jr., M.D., for the skilled
surgery, and to Lall Montgomery, M.D., and George E. Branam, M.D.,
pathologists, for the Kodachrome slides of the gross and microscopic
study of the tumor.
A Comparative Study of Some Effects
of Aminoglutethimide Phosphate
on Serum Potassium and Sodium
George P. Pollock, Indiana State University
Abstract
A comparative study of the effects of 2-(p-aminophenyl)-2-ethyl
glutethimide phosphate on serum potassium and sodium concentrations was
conducted. Rattus norvegicus, Coturnix japonicum, Rana pipiens and Chrysemys
picta were used. A significant increase in serum potassium and a significant
decrease in serum sodium were observed in the homeothermic species. No
significant changes in the two electrolytes were observed in the
poikilothermic species.
Introduction
The compound 2-i-(p-aminophenyl)-2-ethyl glutethimide phosphate
(AGP) has been used clinically as an anticonvulsant in the treatment of
generalized seizure and petit mal and temporal lobe epilepsy (1). Admin-
istration of the drug has resulted in some undesirable side effects.
Changes in endocrine function similar to those produced with amphenone
B were reported by Hertz et al (4). Pittman and Brown (5) observed
hypertrophy of thyroid, ovarian, and adrenal tissues. Dexter (2) indi-
cated a steroidogenic block with AGP prior to A-5-pregnenolone. The
specificity for the site of this block was indicated since addition of the
latter compound allowed normal synthesis to occur. The experiments of
Eversole and Zimmerman (3) suggested that abnormal mineralocorticoid
synthesis occurred in rats following AGP treatment, since they reported
deviations in urinary and blood electrolyte values in these animals. The
present investigation was conducted to determine if similar effects of
AGP are found in the serum electrolytes of other classes of vertebrates.
Representative mammals, birds, amphibians, and reptiles were used in
this study.
Materials and Methods
Female Rattus norvegicus, white rats of the Charles-River strain,
Coturnix coturnix japonicum, Japanese quail, Rana pipiens, grass frogs,
and Chrysemys picta, painted pond turtles, were used in this experiment.
All animals used were fasted 24 hours prior to treatment but were
allowed water ad libitum. The AGP solution administered was prepared
in a concentration of 100 mg of AGP/ml of distilled water. Each animal
was given two injections of AGP in a dosage of 100 mg/kg body weight
at 12-hour intervals. The injections for the rats, quail, and turtles
were given subcutaneously in the neck region. The frogs were injected
in the dorsal lymph sacs. Each control animal received an injection of
distilled water comparable to the volume of AGP that it would have
received as an experimental animal. Twenty-four hours after the first
injection, the animals were sacrificed by guillotining. The blood was col-
lected, and the serum was separated by centrifugation. Serum electro-
468
Zoology 469
lytes were measured by flame photometry using the "Coleman Flame
Photometry Manual" method. This procedure required serum dilutions
of 1:50 for potassium and 1:200 for sodium samples. Since small quanti-
ties of serum were obtained from the frogs and quail, all sodium samples
were prepared from the previously tested potassium samples to maintain
consistency in all tests.
The standard "t" test was used for the statistical evaluation of the
Results
The AGP injections caused extreme inactivity in the rats and quail
with many specimens unresponsive to external stimuli. This condition
was more severe and prolonged in the quail than in the rats. Only
initial irritation, shown by considerable kicking, was evident in the frogs
and turtles. AGP caused significant differences in serum potassium and
sodium concentrations in both the rats and quail. These differences were
expressed as an increase in potassium and a decrease in sodium. No
variations in these electrolytes were observed in the frogs and turtles.
These results are cited in Table 1.
table 1. Serum Sodium and Potassium Concentrations.
Na-
f
AGP
Control
A>
Species
Control
AGP
R. norvegicus
147.0 ± 2.2
104.6 ± 3.1*
6.2 ± .3
8.1 ± .3*
(12)
C. japonicum
149.2 ± 3.4
124.2 ± 3.6*
5.5 ± .3
8.1 ± .3*
(10)
R. pipiens
114.8 ± 2.0
114.2 ± 1.5
2.6 ± .04
2.7 ± .06
(24)
C. picta
118.5 ± 1.0
119.3 ± 1.2
2.4 ± .08
2.5 ± .04
(24)
P < .01.
* Indicates significant difference.
All serum concentrations are in mEq/1 of solution.
Discussion
Some individuals question the use of guillotined animals for electro-
lyte determinations because of simultaneous collection of body fluids.
The results of this experiment are all relative values since all samples
were obtained in the same manner. It is to be noted that the control
values correspond with those reported by Prosser (6) for the rats, frogs,
and turtles.
470 Indiana Academy of Science
The results obtained in the experiments by Eversole and Zimmerman
(3) in rats were similar but less profound than those found here in both
rats and quail. These differences might be expected in the experiments
of those authors since they gave one injection per 24 hours as compared
with two injections per 24-hour period for the present experiment. The
increased response might then be attributed to i. a pharmacological
booster effect of the AGP titer in the bloodstream, or 2. an increased
response due to more AGP being present per 24-hour period.
The evidence that AGP produces a steroidogenic block prior to
A-5-pregnenolone, prepares one to assume a marked reduction or absence
of mineralocorticoid secretion after treatment with the drug. The
changes in the electrolytes of the rats and quail found here would not be
unexpected. Furthermore one might expect identical changes in frogs and
turtles since identical hormones have been isolated from these vertebrates.
The question then arises as to why there were no changes in the electro-
lyte balance of these animals following treatment with AGP.
Several possible answers to this question exist, but they are highly
speculative. This is due in part to lack of a method for qualitative
determination of AGP in the bloodstream of these animals. If one
assumes that AGP is absorbed and acts in cold-blooded animals in the
same way as in warm-blooded animals, then one cannot overlook the idea
of the existence of an alternate pathway of synthesis for corticosteroids
unaffected by AGP treatment. There are tissues of unknown function
which are suspected of steroidogenic action in both amphibians and
reptiles (7).
A second area that needs investigation is that of absorption of the
compound by these animals. The degree of vascularity in the subcu-
taneous neck region is not as extensive in the turtle as in the rats and
quail. The results obtained would indicate that absorption of the com-
pound was negligible for both the frogs and the turtles. When the mass
of the turtle shell is considered as part of the total mass, and the
injections were given on a body weight basis, the dose given to the
turtles was the highest received by any of the animals. The frog injec-
tions were given in the lymph sacs and should have reached the general
circulation more rapidly than in any of the other animals. This method
of injection almost insures the presence of AGP in the bloodstream, and
since no response was illicited, it then seems logical to assume the drug
does not block steroidogenesis in the frog. Metabolic alteration of AGP
by the frog and turtle exists as a possible cause for their lack of response.
If the compound is altered to some intermediate before it can cause its
block, then no change could occur. To be able to support this hypothesis,
some means of qualitative determination of AGP must be developed.
Literature Cited
1. Bauer, R., and J. S. Oleyer. 1960. Clinical evaluation of Elipten, J. Mich.
Med. Soc, 59:1829-1832.
2. Dexter, R. N., L. M. Fisher, A. C. Black, and R. L. Ney. 1966. Inhibition
of adrenal corticoid synthesis by aminoglutethimide. Clinical Res. 14:16
(abstr.)
Zoology 471
3. Eversole, W. J., and R. E. Zimmerman. 1968. Effects of aminoglutethimide
on water and electrolyte metabolism. Fed. Proc. 27:691.
4. Hertz, R., W. W. Tullner, J. A. Schricker, P. G. Dhtse, and L. P. Hall-
man. 1955. Recent Prog-. Hormone Res. 11:119.
5. Pittman, J. A., and R. W. Brown. 1964. Antithyroid and antiadrenocortical
activity of aminoglutethimide. J. Clin. Endo. and Metal. 26:1014-1016.
6. Prosser, C. L. 1966. Comparative Animal Physiology, ed. 3. W. B.
Saunders Co., Philadelphia.
7. Turner, C. D. 1966. General Endocrinology, ed. 4. W. B. Saunders Co.,
Aspects of Water Loss Physiology in Certain
Plethodontid Salamanders
Barbara L. Pickard and Albert E. Reynolds, Hudson Valley Community
College (Troy, N. Y.) and DePauw University (Greencastle)
Abstract
Weight reductions due to evaporative loss of water under varying- con-
ditions of temperature and relative humidity are presented for Plethodon
jordani jordani, Plethodon glutinosus, and Desmognathus ochrophaeus carolinen-
sis, the data for the latter two being somewhat less conclusive. The experi-
mental conditions employed were those presumed to approximate conditions
encountered by animals in nature.
Introduction
It is a commonplace of even casual observation that the various
species of amphibians do not all live under the same environmental condi-
tions, but that, rather, each species occurs in its own reasonably distinc-
tive situation. These situations are, however, characterized by one rela-
tively constant feature in that all amphibian organisms require some
access to water, but even in this respect there is a diversity of habitats
varying from those that are definitely aquatic to those that are almost
completely terrestrial. With respect to the capacity to conserve body
water and to resist, or survive, the desiccating influence of atmospheric
air, the members of the Class Amphibia thus present a whole spectrum of
adaptations the achievement of which must have been very crucial in the
history of vertebrate evolution toward complete terrestrialism.
Among studies on the reactions of caudate amphibians to the
evaporating power of air, an early one was that of Shelf ord (9) which
afforded salamanders the opportunity to sample a humidity gradient.
Both Plethodon cinereus and Plethodon glutinosus spent relatively little
time in the dry and medium parts of the gradient, and spent most of the
time in the moist portion. The general results enabled Shelford to state
that "P. glutinosus is clearly more sensitive to dryness than is P.
cinereus" Hall (3) found that Amby stoma punctatum could withstand
evaporational loss of 11% to 47% of body weight and still survive. Little-
ford et al. (4) found that P. cinereus survived greater loss of water
than two of its ecologically more aquatic plethodontid relatives, Eurycea
bislineata bislineata and Desmognathus fuscus fuscus.
In a study of two plethodontids, Aneides lugubris and Ensatina
eschscholtzii xanthoptica and a salamandrid, Triturus torosus, Cohen (1)
found a differential in the time required for the specimens to be
desiccated to a 20% loss of initial body weight, as follows: Triturus
averaging 13.9 gm, a mean of 257 minutes, Aneides averaging 8.47 gm,
a mean of 261 minutes, and Ensatina averaging 5.66 gm, a mean of 185
minutes. About 30% larger than Ensatina, the Aneides also required
about 30% more time to lose 20% of the body weight. The Triturus was
the largest of the three by a considerable margain, yet it lost 20% of its
472
Zoology 473
weight in approximately the same time as did the smaller Aneides.
Within any one species, Cohen was unable to demonstrate a correlation
between body size and rate of water loss.
In a study that encompassed eight species of Western plethodontids,
Ray (8) stressed the specimens by desiccation to loss of a righting reac-
tion, or to lethality. Ray found a definite correlation between size and
rate of water loss, with the larger individuals in each species showing
the slower rates of loss, a relationship which he claimed to be an
expected one on the basis of a difference in surface-to-mass ratio.
The experiments reported in the present paper were undertaken to
obtain information under conditions deemed to be somewhat similar to
those encountered by salamanders in nature, but with the a priori under-
standing that any experimental procedure would necessarily be some-
what unnatural. Some of these conditions were: 1. exposure to a degree
of evaporative loss that would be essentially non-stressful; 2. an air
movement rate of a quite gentle intensity or magnitude; 3. an
experimental run of moderate length.
Materials and Methods
Routine and regular maintenance of salamanders of the type used in
this study was accomplished in a constant-temperature room at 14-16°C.
The animals were housed, in varying numbers, in either glass or plastic
containers of various sizes, marked as to place and date of capture. Adult
Drosophila and small or medium-sized mealworms (Tenebrio larvae)
were fed to the animals periodically. In addition to some provision for
air interchange between the room and the interior of the container, each
container was supplied with sphagnum moss or wadded towel paper,
partly to supply the animals with a means of concealment, but also to
provide a moisture-holder that would incure a saturated air or one of
quite high relative humidity within the container.
For the experiments reported herein, plastic containers 6 inches in
diameter were equipped in the manner just described, and each sala-
mander selected was placed in its own individual container, appropriately
marked, and thereafter not fed until after the experiments were con-
cluded. In order that its weight might be stable, no salamander was sub-
jected to its first experimental desiccation run until after adequate
acclimation to the lack of food. For the experimental runs each sala-
mander was transferred to and confined in a small plastic rectangular
"weighing box" perforated by as many small holes as the box structure
would permit. The weight of these boxes was frequently checked, and in
such boxes each specimen was weighed immediately before and immedi-
ately after each desiccation run. Weighing was accomplished on a
magnetically-damped chainomatic balance located in the constant-
temperature room. For transport between the constant-temperature room
and the environment-control chamber, the salamander in its box was
placed in a small plastic bag.
The experiments were conducted in the exposure chamber of an
environment-control device known as a Vapor-Temp (Model VP-200At,
474 Indiana Academy of Science
Blue M Manufacturing Company) which has built-in equipment giving it
the capability of automatically maintaining relative humidities from 34%
to 98% and dry bulb temperatures from 4°C to 77 °C. Air-flow control
can vary from 0 to 30 complete changes of exposure chamber volume per
minute. The Vapor-Temp is furthermore provided with built-in features
to record both wet-bulb and dry-bulb temperatures on a circular record
sheet. This was supplemented by the installation within the exposure
chamber of standard centigrade thermometers, one of which was wet-
bulb, the other dry. By coating the interior of the Pyrex glass exposure
chamber with an antifog solution, complete visibility within the chamber
was achieved.
In preparation for an experimental run, the Vapor-Temp was turned
on and regulated to the desired temperature-humidity environment within
the chamber. All runs were conducted at minimal air flow. When the
weighed salamander was introduced, a short wait ensued until the desired
environmental conditions were re-established in the chamber, and the
run was then continued for one hour. Following this sixty minutes of
exposure, the animal was re-weighed, and then returned to its own
individual container. Each animal was then periodically re-placed in a
"weighing-box" and its weight recorded until it had re-gained its original
weight. In so far as was feasible, each animal was used in all of the runs
included in the program.
The animals studied in this program included 19 specimens of
Plethodon jordani jordani and four specimens of Desmognathus
ochrophaens carolinensis taken at Indian Gap, Sevier County, Tennessee.
Appreciation is hereby extended to the Naturalist Service of the Great
Smoky Mountains National Park for cooperation with this field work.
One included specimen of Plethodon glutinosus glutinosus was collected
west and north of Dillard, Georgia, approximately 2.5 miles north of the
Georgia-North Carolina border along the Betty Creek Road, in Macon
County, North Carolina. All 24 specimens just mentioned were collected
in a program aided by a National Science Foundation Grant adminis-
tered by the Highlands Biological Station, Highlands, North Carolina, at
which one of the authors was a resident investigator. The two remaining
specimens of P. glutinosus were both Indiana specimens, one collected in
the Morgan-Monroe State Forest, the other at Friends Camp in Putnam
County.
Weight changes experienced by the animals during the runs are
regarded as all being due to water loss: no defecation occurred during
any of the runs. In reporting these changes, the raw data of weight
change in grams have not been recorded; instead, the changes of weight
are expressed as a proportion of the original weight.
Results
1. Plethodon jordani
In all cases the environmental "starting point" for the salamanders
used was water-saturated air at 14-16° C. A glance at Table 1 will reveal
Zoology
475
table 1. Weight lost by specimens of Plethodon jordani in one hour
under various eyivironmental conditions.
Speci-
men
Initial
Percentages of
initial weight lost at indicated
No.
Wt. (gm)
relative humidity (in %) and temperature.
36%
48%
70%
85%
92.5%
11°C
17°C
15C°
26°C
29°C
A14
1.5736
13.371
10.089
11.021
6.217
2.127
A15
1.7692
5.658
11.053
10.768
3.754
2.526
Al
1.8680
3.81
24.38
10.72
9.112
A 5
1.9740
10.088
17.509
gain
4.600
gain
A7
1.9740
14.453
9.192
3.608
4.200
1.396
A8
2.1479
13.855
13.177
10.472
5.055
3.060
A10
2.3206
15.340
9.404
8.954
3.942
1.921
All
2.4084
11.879
11.725
8.873
2.254
5.870
A3
2.5147
10.22
8.866
6.000
4.37
0.070
A13
2.7094
12.453
10.973
9.238
3.271
4.622
A9
2.7546
11.685
7.666
6.625
3.416
1.591
A16
2.8924
13.383
13.009
9.648
3.572
2.334
A18
2.9209
9.268
9.197
6.646
2.881
3.514
A19
3.0263
13.132
17.596
2.624
4.655
0.400
A17
3.4884
8.497
8.143
7.399
4.759
3.306
A6
3.7439
9.917
11.075
9.930
4.251
7.885
A12
3.9314
7.707
9.921
9.559
2.215
2.651
A4
4.0264
7.319
8.676
4.587
3.316
3.685
A2
4.8146
36.220
■
5.198
4.22
13.10
Mean
2.7820
12.013
11.758
7.881
4.213
3.532
at once that the animals lost weight under all conditions to which they
were experimentally subjected, with two exceptions. At the bottom of
Table 1 the means recorded show a general trend for the animals to lose
a greater proportion of body weight on exposure to progressively less
humid air. With respect to this trend, a column-by-column inspection of
Table 1 supports the summary which follows.
Lost more weight at 36%/ll°C than at 48%/17°C = 11 specimens:
A14, A7, A8, A10, All, A3, A9, A16, A18, A17, A13;
Lost more weight at 48%/17°C than at 70%/15°C = 17 specimens:
A15, Al, A5, A7, A8, A10, All, A3, A13, A9, A16, A18, A19, A17, A6,
A12, A4;
Lost more weight at 70%;/15°C than at 85%/26°C = 16 specimens:
A14, A15, Al, A8, A10, All, A3, A13, A9, A16, A18, A17, A6, A12,
A4, A2;
476 Indiana Academy of Science
Lost more weight at 85%/26°C than at 92.5% /29°C = 11 specimens:
A14, A15, A5, A7, A8, A10, A3, A9, A16, A19, A17.
Thus under all of the conditions imposed, from 11 to 17 of the 19
animals showed less water loss per hour with increasing relative
humidity, or, conversely, greater water loss per hour as the relative
humidity of the surrounding air decreased. In the array of experimental
circumstances, perhaps the most "natural" temperatures for the sala-
manders were the 15° and 17° exposures; in these exposures differing by
only 2°C, the relative humidity difference was of 22%, and 17 out of the
19 animals showed a difference in greater water loss in the less humid
environment. Six animals showed a particularly consistent behavior
under all conditions in that they lost progressively less water with
increasing relative humidity, as follows: A8, A10, A3, A9, A16, A17.
The general trend of the relationships just reviewed is supportive
on the thesis that P. j. jordani exhibits an hourly water loss rate
inversely proportional to the ambient relative humidity in its environment.
In Table 1 the specimens are listed, from top to bottom, in order of
increasing size as reflected by body weight, an arrangement that was
deliberately adopted in view of a possibility that water loss rate might
be size-related. Since evaporative water loss is skin-related, generally-
accepted principles concerning surface-area-to-mass ratios might make
it logical to expect that the smaller salamanders would sustain greater
loss than the larger ones. If such occurred, inspection of the vertical
columns in Table 1 should make it apparent. Such an inspection of
Table 1, column by column, conveys strongly the idea that the sala-
manders were extremely variable in their response to desiccation. Within
this variability, however, weak trends may be discerned at 48%/17°C,
70%/15°C, and 85%/26°C; under these conditions, the larger animals
show a tendency to lose a smaller proportion of the body weight than
do the smaller ones.
table 2. Plethodon jordani: Mean weight loss by size groups.
Mean Weight 36%
48%
70%
85%.
92.5%
Specimens
(gms)
11°C
17°C
15°C
26°C
29°C
A14, A15,A1, A5
1.7962
8.231
15.757
10.836
5.920
2.326
A7,A8,A10,A11
2.2127
13.881
10.874
7.976
3.862
3.061
A3,A13,A9,A16
2.7177
11.935
10.128
7.877
3.657
2.154
A18, A19, A17, A6
3.2948
10.203
11.502
6.649
4.136
3.776
A12, A4, A2
4.2574
17.082
9.298
6.448
3.250
6.478
For further investigation of this tendency, Table 2 has been con-
structed by arbitrarily dividing the 19 specimens into five groups, and
presenting the loss rates as means for the resulting size groups. Such
a consolidation has the virtue of affording slightly clearer evidence of
Zoology 477
a weak trend for the smaller salamanders to lose more weight at
48%/17°C, 70%/15°C, and 85%/20°C than do the larger ones.
2. Other Salamanders
Two other species of Plethodontid salamanders were studied, and
these were subjected to an additional experimental run at 81% relative
humidity and 16° C. The number of specimens available was small, and
some of the animals failed to regain original weight and were not used
in some of the experimental runs (Table 3).
table 3. Weight lost by individuals of two salamander species in one
hour under various environmental conditions.
Speci- Initial
men Weight
No. (gm)
Percentages of initial
relative humidity (in
weight lost at indicated
% ) and temperature.
36% 48%
ire i7°c
70%
15°C
81%
16°C
85%
26°C
92.5%
29°C
A. Desmognathus ochrophaeus carolinensis
E3 0.6995
E4 1.3226
El 1.3326
E2 1.3884
21.806 21.668
17.753 18.937
12.457 11.754
18.475
16.909
15.359
12.034
9.768
10.769
5.657
0.072
0.516
0.034
2.991
Means 1.1857
17.6227 17.4530
14.7673
9.7680
5.4993
1.1803
B. Plethodon glutinosus glutinosus
B2 2.0423
B3 2.6068
Bl 6.5051
17.808 11.663
14.021 19.319
10.253 9.430
8.517
9.919
18.050
9.1927
10.745
4.958
4.028
2.906
1.9792
5.554
3.818
Means 3.7180
14.0273 13.4706
12.1620
8.2985
3.4670
3.7837
Reference to Part A of Table 3 will show that the largest specimen
of D. o. carolinensis was still smaller in terms of body weight than the
smallest specimen of P. j. jordani listed in Table 1. The mean percentages
of weight loss of D. o. carolinensis shown in Table 3 were consistently
greater than that of P. j. jordani exposed to comparable experimental
conditions, a result logically to be expected on the basis of surface-mass
ratio relationships. Also consistent with such a relationship was the
fact that (except at 92.5% /29°C) the smallest carolinensis exhibited
a higher hourly loss rate than did the three somewhat larger specimens,
all three of which were about the same size. With respect to hourly
478 Indiana Academy of Science
rate of water loss in correlation with environmental condition, the
mean values recorded in Part A of Table 3 show a progressive decrease
as the relative humidity increased. Inspection of the separate values for
each specimen indicate the same as a general trend, but with an
occasional exception.
The three specimens of P. g. ghitinosus used included two specimens
near the same size plus one quite large animal (Table 3, Part B) which
was in fact the largest of the total of 26 animals involved in these experi-
ments. The two other ghitinosus specimens were somewhat larger than
the D. o. carolinensis and near the mean weight of the P. j. jordani used.
This largest ghitinosus exhibited the least weight loss in all ghitinosus
runs except those at 70%/15°C and 92.5% /29°C. For comparable environ-
mental exposures, mean weight loss of glutinosus was less than of D. o.
carolinensis in all runs except that of 92.5% /29°C, but greater than that
of P. j. jordani in all runs except at 85%/26°C. Although not consistent
for the 85%/26°C and 92.5% /29°C runs the mean percentage of weight
loss in glutinosus generally increased progressively with reduced relative
humidity (Table 3, Part B).
Discussion
One of the most striking features of the results just presented is the
marked amount of variation observed. One specimen of P. jordani lost
as much as 13% of body weight in relatively humid air at 92.5% rela-
tive humidity, while one other specimen lost a rather insignificant 0.07%
of body weight; these represent the extremes of responses to conditions
that evoked an average loss of 3.53% of body weight. In the 36%/ll°C
runs there occurred the largest "spread" between greatest and least
weight loss, a difference of 32.41% of body weight under conditions
where the mean weight loss was 12.0%; yet in this "driest" air of these
36%/ll°C runs, one animal lost only 3.81% of body weight. The smallest
spread in response was seen in the runs at 85%/26°C where the difference
was 6.9% under conditions where the average loss was 4.2%. The more
scant data on the other salamanders, D. o. carolinensis and P. glutinosus,
exhibited differences more intermediate in magnitude, but nevertheless
show variation.
In spite of the very considerable variation manifested, salamanders
of all three taxa exhibited a trend, as has been indicated in the previous
section, to lose body weight in direct proportion to the dryness of the
surrounding air. This response of the animals to the evaporative power
of air is in thorough accord with logical expectation.
In view of the generally-held view that small organisms have a
higher ratio of surface area to mass than do larger organisms, another
logical expectation was that the smaller salamanders would lose more
weight in percentage of initial weight, during the 60-minute runs than
would the larger salamanders. This expectation was borne out only to
the extent that the data presented above give evidence of a weak trend.
Inspection of the Tables will show several deviations from this expecta-
Zoology 479
tion, and one gross violation of it was exhibited by P. jordani No. A2
which was the largest of the jordani series, yet it sustained the greatest
weight loss under two sets of conditions, 36%/ll°C and 92.5% /29°C.
These experiments thus fail to confirm unequivocally a definite correla-
tion between size and rate of water loss found by Ray (8) yet indicate
more of a trend in this direction than was found by Cohen (1), who
found no correlation.
MacMahon (5) studied water loss rate in P. jordani, P. glutinosus,
and also Pleihodon yonahlosse at 20°C, air flow of 2000 cc/minute, and
0% relative humidity, and concluded that the living salamander was
unable to exert any effective physiological control over loss of water but
suffered a rate of loss determined entirely by body composition and body
proportion (surface-volume ratio). MacMahon paralleled his studies of
living salamanders by the use of inanimate "model salamanders" made
up of 80% water and 20% agar by weight, and cast in molds that were
exact replicas of typical individuals of the three species, and found that
these lost weight at the same rate as did their living counterparts. In
another report (MacMahon, 6) based on 573 specimens of the same three
species, air flow rates of 0, 1000, 2000, and 3000 cc/minute were used, and
runs were made a 0% and 100% relative humidity, all at 20°C. That
water loss was least in P. jordani and greatest in P. yonahlosse was
ascribed by MacMahon to the fact that the former had the greatest, the
latter the least, surface-volume ratio. Hourly weight checks during the
experimental runs enabled MacMahon to determine a critical activity
point (CAP) at which the salamander concerned failed to accomplish a
righting reaction. Although all three species were variable, in general
both P. jordani and P. yonahlosse had lesser CAP values than P.
glutinosus. In addition P. glutinosus from montane areas had smaller
CAP values than specimens from non-montane localities. These latter
considerations constrained MacMahon to propose a genetically-controlled
CAP in these salamanders.
In the present report, in all runs except that in the most humid air,
the greatest mean water loss rates were experienced by specimens of
D. o. carolinensis, the smallest organism used, a result in conformity
with MacMahon's (5) emphasis on the role of the surface-volume ratio.
It may also be pertinent, however, to point out that D. o. carolinensis
lives in natural situations that are typically somewhat more moist than
those harboring either P. jordani or P. glutinosus, so the faster rate of
water loss could be due to intrinsic morphologic as well as physiologic
factors that are genetically determined and which are part and parcel of
the total adaptive make-up of the species.
With respect to P. glutinosus, the mean values in Table 3 indicate
a slower rate of water loss than was characteristic of D. o. carolinensis,
a result consonant with the surface-volume relationship since glutinosus
is a larger animal. However, glutinosus also lost more water per hour
than did P. jordani in all runs except that at 85%/26°C. The mean
weight in Table 3 for P. glutinosus is skewed upward because of one
quite large animal. To make an additional comparison, the two most
480 Indiana Academy of Science
similar glutinosus (B2, B3) and the four most comparable jordani (A8,
A10, All, A3) were singled out and separate sub-means calculated for
each group; the mean initial weight for both groups was slightly greater
than 2.3 grams. In terms of these sub-means, P. glutinosus still lost
more water in an hour than did P. jordani, and under all of the experi-
mental conditions. These P. glutinosus vs. P. jordani comparative
responses to evaporative water loss are not in strict accord with the
thesis that surface-volume relationships control water loss. The possi-
bility exists that the capacity of P. glutinosus to lose water at a slightly
faster rate could be one facet of its organization making possible the
higher CAP value found by MacMahon (6). Certain it is that P.
glutinosus as a total population must be so constituted as to be reason-
ably well adapted to quite a variety of conditions since it has a very
extensive range as compared to either P. jordani or D. o. carolinensis, a
range that includes much more non-montane territory than the high-
rainfall southern Appalachian mountainous area where it overlaps both
of the other two forms (2). MacMahon (6) pointed out that because of
higher temperature and lower rainfall animals in non-montane areas
may have had to withstand greater moisture-loss stress, and may have
been the objects of selective action resulting in the higher CAP.
Throughout this paper attention has been focused on the relative
humidity of the atmosphere surrounding the experimental animals, and
no analysis or special study has been made of temperature relationships.
In general, and with one exception, higher experimental humidities have
been accompanied by higher temperatures where evaporation rates could
logically be expected to be greater. Furthermore, full recognition is given
to the small sample sizes, particularly as applied to D. o. carolinensis and
P. glutinosus, and to the consequent fact that the results do not support
any high degree of conclusiveness. This investigation was undertaken
in the spirit of an exploratory study utilizing a basic experimental design
that differed from several other studies of a like nature chiefly in that
less severe physiological stresses were imposed on the experimental
animals. It was felt desirable that the actual results, as set forth in
the Tables, be made a matter of record, along with the logical inferences
that may be derived from them in terms of indicated trends.
The material presented in this paper formed a portion of a some-
what broader program used in another connection (7). This broader
program took note of the recovery of the experimental animals following
evaporative water loss. The time required for the recovery of 99%
to 100% of initial weight was variable, but occurred as follows:
P. jordani — 3 hrs. 35 min. to 38 hrs. 35 min.
P. glutinosus — 2 hrs. 20 min. to 23 hrs. 11 min.
D. o. carolinensis — 4 hrs. 30 min. to 22 hrs. 56 min.
Summary
In this preliminary study nineteen specimens of Plethodon jordani
were subjected to relative humidity and temperature conditions as fol-
lows: 36%/ll°C, 48%/17°C, 70%/15°C, 85%/26°C, and 92.5%/29°C. To
Zoology 481
supplement this, four specimens of Desmognathas ochrophaeus carolinen-
sis and three specimens of Plethodon glutinosus were exposed to the
same conditions plus an additional experimental run at 81%/16°C. All
experimental runs were one hour in length, and weight losses were all
considered to be due to evaporative loss of water. Weight losses were
presented in tabular form as percentages of initial weight. Weight loss
was quite variable, the extremes extending from as little as 0.07% to
as much as 36.22% of initial body weight. The most distinct trend evinced
by the results was weight loss, or loss of water, in direct proportion
to atmospheric dryness. The data gave weak support to the thesis
that water loss is very largely controlled by the ratio of surface-to-mass
(or volume). Other inferences from the data suggested a taxonmic,
genetically controlled, difference in response to desiccation. After a
desiccation run, the animals recovered initial weight in periods varying
generally from 3 to 23 hours.
Literature Cited
1. Cohen, Nathan A. 1952. Comparative Rates of Dehydration and Hydra-
tion in Some California Salamanders. Ecology 33:462-478.
2. Conant, Roger. 1958. A Field Guide to Reptiles and Amphibians. Hough-
ton Mifflin Company, Boston.
3. Hall, F. E. 1922. The Vital Limits of Exsiccation of Certain Animals.
Biol. Bull. 42:31-51.
4. Littleford, R. A., W. F. Keller, and N. E. Phillips. 1947. Studies on the
Vital Limits of Water Loss in the Plethodont Salamanders. Ecology
28:440-447.
5. MacMahon, James A. 1964. Factors Influencing the Rate of Water Loss in
Salamanders. Amer. Zool. 4(3):144.
6. . 1965. Water Loss in Three Species of the Salamanders of the
Genus Plethodon. Amer. Zool. 5(2):116.
Pickard, Barbara L. Christopher. 1965. Comparative Physiology of Water
Loss in Plethodontid Salamanders. Master's thesis, Roy O. West Library,
DePauw University.
Ray, Carleton. 1958. Vital Limits and Rates of Desiccation in Sala-
manders. Ecology 39:75-83.
Shelford, Victor E. 1918. The Reactions of Certain Animals to Gradients
of Evaporating Power of Air. Biol. Bull. 25:79-120.
Rapid Approximations for some Chi Square and Derived
Correlational Statistics Used in the Social and
Biological Sciences.1
Hans W. Wendt, Valparaiso University"
Abstract
Short-cut expressions are given which approximate basic forms of Chi
Square first and second order interactions, the fourfold point coefficient of
correlation, and the contingency coefficient. This strictly empirical approach
assumes that cell frequencies in tabled data distributions are roughly sym-
metrical. Under these conditions, differences among marginal totals of
2 by k tables are small and little error is introduced by substituting, in
effect, an average value for pairs of unequal diagonal cell entries. The esti-
mates are compared with conventional solutions for 20 examples, with
two thirds of these falling within approximately ± 10% of the value of the
statistic in question as ordinarily computed. The short cuts approach the
more precise solutions as the cell entries involved become more fully sym-
metrical. Criteria are offered for predicting the direction of the error
resulting from the simplified substitutions.
Introduction
Some time ago a short cut method was pointed out for dealing with
special cases of the Chi Square situation (6), intended to complement
a number of standard discussions of nonparametric data treatments (1,
2, 3, 5, 7). Extensions of the original procedure have since been used in
experimental work and in teaching students to run rapid over-all checks
on conventional calculations. We have found them helpful in brief treat-
ments of the significance and correlation topics in courses for non-spe-
cialists who needed some basic statistical criteria that could be used in
field settings and would permit quick decisions without incurring exces-
sive error.
Summaries of the modified procedures are given below. Analytically
or empirically, criteria might be derived by means of which some equiva-
lent of confidence intervals would be available for these estimates. How-
ever, this complication could easily defeat their basic purpose. Instead,
we are presenting a number of practical examples (not systematically
selected except for the constraints discussed below) where standard
computational procedures and short cuts have been used side by side, to
show the type or error that can be expected. The investigator will de-
cide for himself in which situations the rapid estimates should merely
complement the more rigorous solutions (as in preliminary screening
of data from field work and gross checks on more elaborate work); or
where circumstances suggest that they might be used independently.
1 The author is indebted to Waldemar C. Gunther for suggesting possi-
ble applications of this methodology, originally based on behavioral studies,
to biological and zoological research.
2 Now at Macalester College.
482
Zoology 483
The 2 by 2 table
Experimental and survey data are often cast in the form of a
fourfold table of classes of events or subjects, with observed frequencies
labelled
A
B
C
D
The formula customarily used for determining interaction is
abbreviated from the definitional expression for Chi Square, or (without
correction for continuity),
N (BC— AD)3
«?
X"~" (A+B) (C+D) (A+C) (B+D)
While simpler than another (basic) expression, computation for any N
over approximately 50 magnitude becomes laborious. Furthermore, deci-
mal errors are often made by students who are not thoroughly practised
with contingency tables.
If possible, the 2 by 2 table is set up such that all four marginal
totals are equal. This design assures the most powerful test of the null
hypothesis and minimizes the problem of small expected frequencies.
Where data from continuous distributions are to be collapsed into two
categories on each variable this means dichotomizing around the re-
spective medians. Given the special situation of successful median splits,
however, the conventional formula reduces to
(|A — C| + |B— D|)a
2 ■
r~~ N
Moreover, this expression is still satisfactorily close to the long solution
where only the general criterion of "ordinal symmetry" is satisfied, that
is, where observed cell frequencies are arranged as
A>B
V A
C<D
Given such a basic pattern (or its mirror image),
2_^ (|A — C| ' |B — D|) Approximation for Chi Square
^ N in 2 by 2 table3
In effect, we simply square the sum of the left and right column (abso-
lute) differences and divide by total cases. The procedure can be equally
well phrased, of course, in terms of row differences as well as differences
between diagonal sums. We have found the formulation above the most
practical and least confusing to the student, and it is consistent with
the approximation proposed for special 2 by 2k tables.
3 It was pointed out to the author by G. Lienert that the analytical
basis for the validity of the above assumption has been worked out earlier
by M. H. Quenouille {Rapid statistical calculations, London, Griffin, 1959),
who arrived at an equivalent formula for this special 2 by 2 case.
484
Indiana Academy of Science
If ordinal symmetry in the table deviates greatly from complete
symmetry in metric terms, that is, where any two diagonal cells are
greatly different although the basic condition stated earlier is satisfied,
X2 will be systematically over- or under-estimated depending on whether
the larger or the smaller diagonal is involved (see later).
The examples in Table 1, some taken from the texts quoted, compare
the rapid approximation with the standard method and show the conse-
quences for accepting or rejecting the null hypothesis. The last two
examples illustrate the type of error which results from gross imbalances
in a diagonal.
TABLE 1.
Basic 2 by 2
table
X2, standard
procedure
P
X2, estimate
P
Error, % of
true value
J 6
11
8
25
7.59
<.01
8.1
<.01
+7
17
13
12
24
3.62
<.10
3.9
<.05
+8
37
52
66
49
5.02
<.05
5,0
<.05
0
16
30
28
25
3.24
<.10
2.9
<.10
—11
75
98
113
89
5.93
<.02
5.9
<.02
—1
24
20
17
22
0.99
>.30
1.0
>.30
+ 1
34
94
37
35
12.40
<.001
19.2
<.001
+55a
19
1
12
18
15.40
<.001
11.5
<.001
— 25a
aNote asymmetry in diagonal cells.
The 2 by 2k table
In the standard treatment, individual terms of the type
(O— E);
E
are summed after expected (E) frequencies in each cell are computed
from the marginal sub-totals of the observed (O) frequencies. There is
a special case where (a) row totals are equal, and (b) where observed
cell frequencies assume only two different values. Here the procedure
can be much simplified. For example, if a contingency table
A
B
C
1)
E
F
G
H
Zoology
consists of the following hypothetical cell frequencies
485
15
25
15
25
25
15
25
15
an analogy exists to the 2 by 2 case, and
(|A— E| + |B— F| + |C — G| + |D— H)
X2=
N
ii)
= =10.0 (df=3, P < .02).
160
This result is identical with the conventional long solution. If the
stated equalities are satisfied only approximately, the identity no longer
holds in the general case. However, it can often be written, without
excessive error,
(|A — E| + |B — F| + |C — G| + |D — H|)2 Approximation for
N
Chi Square
in 2 by U table
The solution extends to 2 x 2k tables. (With constraints stated, the
number of columns is necessarily even.) There we add all observed
column differences, square the total, and divide by N.
Table 2 compares some solutions for cases of moderate deviations
from the desired pattern.
TABLE 2.
X
Basic 2 by 4 table
a, standard
procedure
P
X3, estimate
P
Error, % of
true value
7 15
17 8
18
6
6
23
21.76
<.001
21.2
<.001
—3
13 8
6 14
12
7
L0
20
8.69
<.05
8,7
<.05
(»
The 2 by 2 by 2 and Higher Order Interactions
One standard treatment, after M. S. Bartlett, for the second order
(2 by 2 by 2) interaction consists in solving a cubic equation (4) and
is rarely given in textbooks. The reasons may be the forbidding amount
of labor required where no computer is available and a relatively low
efficiency of this statistic. Consequently, other measures have come to
be preferred. As the Ns in research often do not exceed the magnitude
of 100 it can be argued that there is mainly one situation where a gross
significance estimate for an interaction may still be desired. Consider
486
Indiana Academy of Science
the case where two 2 by 2 tables show suggestive and opposite trends
in the observed cell entries, as in the example,
9
40
38
5
X
44
3
7
51
Here the condition is satisfied that
A < B A' > B'
A V V A
C> D C < D'
where A'+B'+C'+D'=N'
A+B+C+D=N,
and N~N'
Given this type of situation (and only where opposite trends are evi-
dent from inspection) a workable estimate of the interaction, with df=l,
is possible as
,i
(|A-C| + |B— D| + | A'— C'| + |B'— D'|)3 Approximation
N + N'
for second order
interaction
Chi Square
The amount of time saved over the precise long method tends to be
substantial.
The examples in Table 3 compare solutions by way of the standard
method and the approximation, respectively. It will be noted that the
conditions stated above are violated to some extent in both cases. How-
ever, the basic opposition of diagonal trends is still visible.
TABLE 3.
Basic 2 by 2 by 2
table
X2, standarc
procedure
i
P
X3, estimate
Error, % of
P true value
63 70
70 55
43
35
19 5 51
31 5-51
<.02
4.0
<.05 —17
61 77
101 82
61
42
31
51 n-89
<.001
13.9
<.001 +17
(Speculatively it would seem that the approximation given for the
2 by 2 by 2 case could be generalized to interactions of third or higher
order. Precise computation of this type of statistic by way of solving
the appropriate equations is, to our knowledge, not usually attempted.
Occasionally, however, a significance estimate might be sought by an
investigator if a high degree of symmetry is obvious among the trends
of all sub-tables. For example, in the following hypothetical example an
Zoology
487
interaction may be suspected among such variables as sex, birth order,
affiliation need, and resulting anxiety:
9
12
15
10
15
8
11
20
X
17
10
7
16
15
12
12
15
Extending the earlier reasoning by analogy, the resulting 2 by 2 by 2
by 2 interaction might be approximated from the respective eight column
differences in the four tables as being of the order of
4 V
205
—8.2
We can offer no empirical check on this estimate precisely because no
"standard solution" seems to be readily available.)
Correcting for Continuity
The Yates correction is usually required for both 2 by 2 and 2 by
2 by 2 tables where small expected frequencies would inflate the %2 esti-
mate. It can be applied for the approximations discussed above. For
the regular 2 by 2 table, a constant of 2 is subtracted from the sum of
the column differences before squaring the total. For the 2 by 2 by
2 table, a constant of 4 should first be subtracted in the numerator.
The Fourfold Point Coefficient of Correlation
The definitional formula for rP suggests that computation can be
simplified wherever the cell entries are symmetrical in nature, that is,
result from median splits along both variables. The standard formula,
rp=
BC— AD
V
(A+B) (C+D) (A+C) (B+D)
in the case of equality among marginal sub-totals reduces to
| A— C| + |B— D|
*p=
N
The expression disregards the sign of the coefficient which is easily
determined from the actual cell distribution. If symmetry is not com-
488
Indiana Academy of Science
pletely achieved by reason of tied scores in the original data, or because
a larger number of cells cannot be conveniently combined,
rp~
[A — C| + |B— D|
N
Approximation for fourfold point
coefficient of correlation
In other words, we add the absolute values of the two column differ-
ences and divide the sum by N. This version of a correlation coefficient
is readily computed without as much as a slide rule wherever N is man-
ageable (e.g., an even number and under 50). The ease would seem to
make it unique among a number of other shortcuts since it is faster
than the cosine pi approximation of the tetrachoric, or the Chown-Moran
and Mosteller approximations to the product-moment measure (cf. 5).
The method may be particularly helpful for preliminary item analyses
and gives good precision with test data where near-median splits can be
achieved. It also serves as a general check where Tp or possibly r are
computed by means of the standard procedures.
Table 4 illustrates the adequacy of the simplified formula in some
empirical cases.
TABLE 4.
Basic fourfold
table
rP, standard
procedure
tp, estimate
Error, % of
true value
16
51
64
49
0.319
0.28
—12
20
41
70
18
0.470
0.49
+4
23
37
67
33
0.280
0.30
+7
2
13
10
5
0.544
0.53
—3
7
44
64
19
0.624
0.61
—2
22
73
95
0
0.790
0.77
—3
The Contingency Coefficient for Four Cells
The standard formula is usually written as
-V
N+xa
and, in the special case of the fourfold table,
c=
J
Zoology 489
N(BC— AD)3
(A+B) (C+D) (A+C) B+D)
N+ N(BC— AD):
(A+B) (C+D) (A+C) (B+D)
Where all observed cell frequencies are reasonably symmetrical the
original expression reduces to
C<*-> _ Approximation for four-cell
V
contingency coefficient
(|A-C| + |B— D|)2 + N2
The two examples of Table 5 illustrate the adequacy of the estimate.
TABLE 5.
Basic fourfold
table
C, standard
procedure
C, estimate
Error, % of
true value
3 23
17 2
0.612
0.61
0
11 32
14 13
0.258
0.30
+16
Direction and Size of Error
In the 20 illustrations presented above approximately two thirds
fall within — 12 and +8% of the statistic in question, and one fourth
within — 1 and +1%. This evaluation is based on a small sample of
five somewhat heterogeneous situations. It is also open to the criticism
that the cell frequencies were selected so as to satisfy, primarily, the
constraints stated at the outset. In place of the gross summary above,
however, the short cuts can also be examined in terms of the direction
and amount of error to be expected a priori.
It can be shown, and is demonstrated by inspection of the sixteen
examples in the 2 by 2 category, that the approximation tends to over-
estimate the exact value of the statistic whenever the deviation from true
symmetry is (proportionately) greater within the larger of the two pairs
of diagonal cells. The exact value is underestimated whenever dis-
crepancies are greater between the two smaller cells. At the same time,
asymmetry in the smaller pair of diagonals generally carries less weight
in the overall statistic than a comparable degree of asymmetry involving
the larger pair. Furthermore, it may be argued that the composition of
diagonals in 2 by 2 tables ("symmetrical" in the sense used above) is
distributed as the result of sampling from fixed diagonal sums. Intuitive-
ly, then, larger degrees of asymmetry (in terms of per cent discrepancy)
would normally occur within the lesser cell diagonal because of the
490 Indiana Academy of Science
small numerical values, which readily form extreme proportions as they
approach zero. This would not apply equally to the cell frequencies ob-
served within the other, larger diagonal. If this analysis is correst we
would expect positively skewed error distributions in large scale appli-
cations of the short cut methods. That is, (a) underestimates of x2, rP
and C will be more common, and their amount will be small — the approxi-
mations tend to be conservative; (b) overestimates will be less common
but will be more extreme when they occur.
Literature Cited
1. Lienert, G. A. 1962. Verteilungsfreie Methoden in der Biostatistik. Wein-
heim, Germany: Beltz.
2. McNemar, Q. 1955, 1962. Psychological statistics. New York, Wiley.
3. Siegel, S. 1956. Nonparametric statistics for the behavioral sciences.
New York, McGraw-Hill.
4. Snedecor, G. W. 1946. Statistical methods, 4th edition. Ames, Iowa, Iowa
State College Press.
5. Tate, M. W., and R. C. Clelland. 1957. Nonparametric and shortcut sta-
tistics in the social, biological, and medical sciences. Danville, Illinois,
Interstate Printers.
6. Wendt, H. W. 1958. Naeherungsloesungen fuer Chi-Quadrat-Werte:
Wechselwirkungen erster unci zweiter Ordnung in Vierfeld-Tafeln.
Psychologie und Praxis 2:39-44.
7. Wilcoxon, F. 1949. Some rapid approximate statistical procedures. New
York, American Cyanamid Co.
The Food of Rana catesbeiana in Three Habitats in
Owen County, Indiana
By F. Don Fulk and John 0. Whitaker, Jr.
Abstract
A total of 442 bullfrogs, Rana catesbeiana, was collected, of which 3 67 had
food in the stomachs. Frogs were collected from the White River, strip-pit
ponds and farm ponds by gigging. The most important foods in stomachs
of frogs from the river were scarabaeid beetles, crayfish, lucanid beetles,
terrestrial snails, earthworms, and carabid beetles. Corresponding foods
of frogs from the strip-pit ponds were crayfish, Lepidoptera, spiders, vege-
tation, Dytiscidae and Libellulidae, and from farm ponds they were Rana
tadpoles, crayfish, Libellulidae, Lepidoptera, young Rana sp., and Aeschnidae.
Introduction
Among the papers concerning bullfrog food habits are: Korschgen
and Moyle (7) 455 bullfrog stomachs examined from Missouri; Korschgen
and Baskett (6) 408 Missouri stomachs; Cohen and Howard (3) 300 Cali-
fornia stomachs examined, Brooks (1) 138 Virginia stomachs examined,
Perez (9) 50 Puerto Rico stomachs examined, Surface (10) and Frost
(4) 29 and 25 Pennsylvinia stomachs examined, and Bush (2) 18 Ken-
tucky stomachs examined.
The present study was initiated to determine and compare the food
habits of Rana catesbeiana in three different habitats, river, farm ponds
and strip pit ponds, in central Indiana.
Materials and Methods
The study area was located in Owen County and consisted of approxi-
mately twenty miles of the White River from Gosport to Freedom, eleven
strip pits and fourteen farm ponds. Bullfrogs were collected in June and
July of 1966, 1967, and 1968, by using gigs and lights from a boat in the
river, and from shore in the ponds.
Stomachs were removed soon after collection of the frogs and were
examined in the laboratory using a dissecting microscope.
Description of Study Areas
Around the strip pit ponds were characteristic rows of spill piles in
various serai stages of succession but much Rubus was present. The
ponds were clear and contained Typha latifolia and algae. The immedi-
ate shore of most of the strip pits was essentially void of vegetation.
The farm ponds were in pasture situations usually with relatively
little ground cover along the shore. Typha latifolia in the shallow end
and algae were predominant forms of vegetation.
The White River meanders slowly through flat bottom land and
rolling hill country characteristic of Owen County. Dominant plants
491
492 Indiana Academy of Science
along its banks are Populus deltoides (cottonwood), Betida nigra (river
birch), Salix interior (sandbar willow), and Acer negundo (box elder).
Some important reptiles and amphibians present in the study area
were: Rana pipiens (leopard frog), R. clamitans (green frog), Bufo
woodhousei (Fowler's toad), Hyla crucifer (spring peeper), and Natrix
sipedon (common water snake).
Discussion
During the three years 442 frogs were taken. The stomachs of 367
contained food: 178 from the river, 111 from the strip pits, and 78 from
farm ponds (Table 1.)
table 1. Stomach contents of 367 bullfrogs, Rana catesbeiana, from the
White River, strip-pits, and farm ponds in Owen County, Indiana.
WHITE RIVER
STRIP-PITS
FARM
PONDS
178 stomachs
111 stomachs
78 stomachs
%
%
%
%
%
%
FOOD ITEMS
vol.
freq.
vol.
freq.
vol.
freq.
Scarabaeidae
14.2
23.6
2.3
4.5
3.5
5.1
Crayfish
12.3
18.0
21.3
31.5
14.8
16.7
Lucanidae
9.6
11.2
— -
1.3
1.3
Snails (terrestrial)
8.2
13.5
1.3
6.3
0.4
1.3
Earthworms
7,1
14.6
0.1
0.9
Carabidae
6.8
14.5
1.5
9.0
0.7
2.6
Snails (aquatic)
6.5
11.8
Spiders
5.3
12.4
7.7
21.6
2.5
5.1
Unidentified minnows
4.8
11.2
0.7
0.9
1.3
1.3
Diplopoda
3.6
8.4
4.9
7,0
Elateridae
2.0
3.9
1.5
2.6
Vegetation
1.8
11.0
7.0
33.0
2.8
10.3
Lepoinis machrochirus
1.5
1.7
0.8
0.9
Notropis atherinoides
1.3
1.7
N. chrysocephalus
1.2
1.1
Sialid larvae
1.1
1.7
Rana sp.
1.1
1.1
3.9
6.3
3.9
3.8
Blattidae
1.1
1.1
Libellulidae
1.0
1.7
5.4
10.9
10.4
11.5
Formicidae
0.9
3.9
0.2
2.7
3.3
5.1
Chilopoda
0.8
2.8
Hyla versicolor
0.6
0.6
2.6
2.6
Ambystoma texanum
0.6
1.1
Hydrophilidae
0.6
0.6
0.6
1.8
1.3
1.3
Omophronidae
0.6
0.6
- — -
Natrix sipedon
0.5
1.1
— ■
Graptemys pseudo-
geographica
0,1
0.6
Zoology
493
TABLE
1 (Continued)
WHITE RIVER
STRIP-PITS
FARM
PONDS
178 stomachs
111 stomachs
78 stomachs
%
%
%
%
%
%
FOOD ITEMS
vol. freq.
vol.
freq.
vol.
freq.
Unidentified insects
0.4
2.2
0.4
3.6
Tenebrionidae
0.4
1.1
0,2
1.8
—
Bombidae
0.4
0.6
0.2
01)
0.8
1.3
Plecoptera naiads
0.4
0.6
Oniscidae (sowbugs)
0.3
2.2
0.9
L.8
Silphidae
0.3
0.6
Chrysomelidae
0.3
0.6
0.2
1.8
Unidentified Coleoptera 0.3
1,1
0.7
4.5
1.3
L3
Gryllidae
0.3
1.7
0,7
2.7
Phalangida
(Harvestmen)
0.2
1,7
Notropis spilopteris
0.2
0.6
Lampyridae
0.2
1.1
0.8
1.3
Vespidae
0.2
0.6
1.5
3.6
Lepidopterous larvae
0.2
0.6
3.6
9.9
1 .5
2.0
Cantharidae
0.1
1.1
1 .7
3.8
Pentatomidae
0.1
1.1
0.1
0.9
Apidae
trace
o.o
0.1
0.9
L.9
3.8
Notonectidae
trace
0.6
Agrionidae
trace
0.6
0.4
0.9
0.4
1 .3
Adult Lepidoptera
10.5
21.6
4.7
5.1
Dytiscidae
5.8
14.4
Aeschnidae
2,7
6.3
3.9
3.8
Odonata naiads
1.9
5.4
2,/;
2.0
Trichoptera
1.8
2.7
—
Acrididae
1.6
3.6
—
Tipulidae
1 .3
2.7
—
Stratiomyidae
1.3
4.5
—
Ephemerida
1.:;
2.7
—
Gryllotalpidae
0.9
0.9
—
Curculionidae
0.8
7.0
1.4
2.6
Buprestidae
0.4
0/.)
■
Phalacridae
0.1
0.9
Reduviidae
— -
0,1
1.8
Coreidae
0.3
0.9
1.3
1.3
Belostomatidae
0.2
0.9
0.7
1.3
Cicadellidae
0.2
0.6
Mantidae
0.2
0.0
Mussels
0.1
0.0
Haliplidae
0.1
0.0
Anobiidae
0.1
0.0
Rhysodidae
0.1
0.0
Cerambycidae
0.1
0.9
Coleopterous larvae
0.1
0.0
494
Indiana Academy of Science
table 1 (Continued)
WHITE RIVER
STRIP-PITS
FARM
PONDS
178
stomachs
111 stomachs
78 stomachs
%
%
%
%
%
%
FOOD ITEMS
vol.
freq.
vol.
freq.
vol.
freq.
Unidentified Diptera
0.1
1.8
1.6
3.8
Nepidae
0.1
0.9
Aradidae
0.1
0.9
Gerridae
0.1
1.8
1.3
1.3
Unidentified Hemiptera
0.1
0.9
Membracidae
0.1
0.9
Cercopidae
0.1
0.9
Rana tadpoles
—
20.0
21.8
Plecoptera
___
1.3
1.3
Culicidae
1.3
1.3
Corixidae
0.8
1.3
Chrysopidae
0.3
2.6
Staphylinidae
—
0.2
1.3
Coccinelidae
0.1
1.3
Bombyliidae
—
0.1
1.3
Chrysididae
0.1
1.3
A large variety of animal foods was represented, undoubtedly
because the diet of Rana catesbeiana is greatly influenced by availability.
Almost anything that moves and is of appropriate size is probably taken.
Forty-four food items were recorded from stomachs from the river
frogs, 38 from the farm ponds, and 57 from the strip ponds.
The six most important foods of bullfrogs in the river listed in order
of decreasing utilization were Scarabaeid beetles, crayfish, lucanid
beetles, terrestrial snails, earthworms, and carabid beetles. In the strip-
pit ponds corresponding foods were crayfish, adult Lepidoptera, spiders,
vegetation, Dytiscidae and Libellulidae. In the farm ponds the most
important foods were Rana tadpoles, crayfish, Libellulidae, adult
Lepidoptera, young Rana sp., and Aeschnidae.
Of the six most important foods in each of the habitats, only one,
crayfish, was listed in all three. In the river habitat, various beetles were
important, comprising a total of 35.5% of the total volume of food, and
with three families of beetles included among the top six. Crayfish was
the top food in strip-pits, and second in importance in the other two
habitats. It would appear that crayfish are often a major food of the
bullfrog. Crayfish was the only food in the top six in the river which was
also in the top six in the other habitats. The similarity between stripped
ponds and farm ponds was greater, with three major shared foods,
crayfish, adult Lepidoptera, and Libellulidae. The top food in the farm
ponds, however, was tadpoles, forming 20% of the food by volume. Tad-
Zoology
495
table 2. Ten ?nost importa?it foods (highest volumes) of bullfrogs from
three Indiana, habitats.
RIVER
STRIP-PITS
i
FARM PONDS
% vol.
% vol.
% vol.
1.
Scarabaeids
14.2
Crayfish
21.3
Rana tadpoles
20.0
2,
Crayfish
12.3
Lepidoptera
10.5
Crayfish
14.8
3.
Lucanids
9.6
Spiders
7.7
Libellulidae
10.4
4.
Terrestrial
snails
8.2
Vegetation
7.0
Lepidoptera
4.7
5.
Earthworms
7.1
Dytiscidae
5.8
Yg. Rana sp.
3.9
6.
Carabids
6,8
Libellulidae
5.4
Aeschuidae
3.9
7.
Aquatic snails
6.5
Diplopoda
4.9
Scarabaeidae
3.5
8,
Spiders
5.3
Rana sp.
3.9
Formicidae
3.3
9.
Unidentified
minnows
4.8
Lepid larvae
3.6
Hyla versicolor
2.6
10.
Diplopoda
3.6
Aeschnidae
2.7
Odonata naiads
2.6
poles were very abundant in the farmponds, but not in the river or
stripped ponds.
It would appear that the bullfrog feeds both underwater and on land.
Several of the important foods, the tadpoles, fish, aquatic snails, and pre-
sumably many of the crayfish, are probably or definitely gotten from the
water. However, it is possible that the frogs catch the aquatic prey while
sitting on the shore or in shallow water. Much of the food must be taken
on land, where the frogs would appear to wait on the shore for prey to
appear, probably at night, as indicated by the Scarabaeids, Lucanids, and
the numbers of moths in stomachs.
Somewhat surprising were the numbers of dragonflies since it
would appear difficult for bullfrogs to catch them. The frogs might get
them during the day or at dusk by remaining motionless and capturing
them as they light, or else they might capture them from among the
vegetation at night. The former would appear to be the case. Bullfrogs
were seen on several occasions among cattails in the ponds during the
day, apparently waiting for prey items.
Frost (4) stated that crayfish, frogs, and mice were principal foods
of larger specimens. Percent volumes of crayfish from the White River
were 12%, with 21.3% from strip pits and 14.8% in the farm ponds.
Crayfish formed the highest volume of food from the pits and ponds, but
ranked second to Scarabaeidae, at 14.2% in the river.
Kirn (5) reports a young bullfrog catching and eating another frog,
presumed to be a bullfrog. In California, Cohen and Howard (3) found
17 frogs present in 300 stomachs examined. Young frogs, Rana, were
found to be a relatively minor food item in all three habitats during the
present study. Hyla versicolor specimens were taken from stomachs from
farm ponds and from the White River.
496 Indiana Academy of Science
Tadpoles of Rana (probably R. catesbeiana, R. clamitans and R.
pipiens) were an important item in the diet of the farm pond frogs.
Minton (8) found a 17-inch coral snake in a bullfrog stomach. Two
frogs from the White River had eaten 10- and 11-inch snakes, Natrix
sipedon.
Minnows were found in stomachs in all three areas. Three species of
Notropis were taken from frogs from the river. Lepomis macrochirus
was taken in both river and strip-pit areas. A turtle, Graptemys psendo-
geographica, was found in a stomach from the river.
Among the insects, beetles were the most important order in
stomachs from all three habitats.
Vegetable matter from the three areas occurred frequently, but
presumably was ingested accidentally.
The average length of strip-pit frogs was 103.5 mm; farm pond
frogs averaged 107.2 mm; and the White River frogs averaged 174.8 mm.
Average weights were 158.8 for the strip-pit frogs; 153.2 for the farm
pond frogs and 373.7 for the river frogs. Thus the river frogs were much
larger, but part of the weight difference could be attributed to the heavy
volume of eggs since 78% of females taken in June contained eggs, con-
trasting to 31.3% in July and 14.3% in August. All frogs from the river
were taken in June.
It is likely that the river frogs are larger because of the relative
lack of fishing pressure in the more remote parts of the river. The strip-
pits and farm ponds are heavily fished, thus the larger frogs are probably
rapidly removed by fishermen.
Literature Cited
1. Brooks, G. R., Jr. 1964. An analysis of the food habits of the bullfrog,
Rana catesbeiana, by body size, sex, month, and habitat. Virginia J.
Science (New series) 15:173-186
2. Bush, F. M. 1959. Foods of some Kentucky herptiles. Herpetologica
15:73-77.
3. Cohen, M. W., and W. E. Howard. 1958. Bullfrog Food and Growth at the
San Joaquin Experimental Range, California. Copeia 1958:223-225.
4. Frost, S. W. 1935. The food of Rana catesbeiana. Copeia 1935 :15-1S.
5. Kirn, A. J. 1949. Cannibalism among Rana pipiens Berlandieri and pos-
sibly by Rana catesbeiana, near Somerset, Texas. Herpetologica 5:84.
6. Korschgen, L. J., and T. S. Baskett. 1963. Foods of impoundment and
stream-dwelling bullfrogs in Missouri. Herpetologica 19:89-99.
7. Korschgen, L. J. and D. L. Moyle. 1955. Food habits of the bullfrogs in
central Missouri farm ponds. Amer. Midi. Natur. 54:332-341.
8. Minton, J. E. 1949. Coral Snake Preyed upon by Bullfrogs. Copeia 1949:
288.
9. Perez, M. E. 1951. The Food of Rana catesbeiana in Puerto Rico.
Herpetologica 7:102-104.
10. Surface, H. A. 1913. First report on the economic feature of the
amphibia of Pennsylvania. Zool. Bull. Pa. Dept. Agric.
The Hoary Bat in Indiana
Russell E. Mumford, Purdue University
Abstract
The hoary bat (Lasiums cinereus) has been recorded from 34 counties in
Indiana. All but one record of occurrence fall between April 12 and October
5 ; one specimen was taken January 31. The species probably breeds throughout
the state. Most young are evidentliy born in June. Adult males appear to make
up an extremely small percentage of the Indiana population.
First reported from Indiana in 1844 by Plummer (11) the hoary bat
(Lasiums cinereus) was long considered rare in the state. Lyon (4)
knew of extant specimens from only Monroe and St. Joseph Counties,
although other authors had published records (either without specimens
or specimens since lost) from other localities. For a century after
Plummer caught a hoary bat in a building in Wayne County, little was
learned about this showy species in Indiana. Some Purdue University
students became interested in bats in the late 1940's and in the spring
of 1948 shot eight Lasiums cinereus at West Lafayette (12). James B.
Cope and I initiated a bat banding program in Indiana in 1951 and have
been assisted by Earlham College students and others to the present.
Nixon A. Wilson began banding later. Norman C. Negus did the first
bat banding in Indiana, but worked with non-lasiurine species. John O.
Whitaker, Jr., working in cooperation with the State Board of Health,
has added numerous records through the identification of bats submitted
to the Board for rabies tests.
Distribution and Seasonal Occurrence. The hoary bat probably
occurs throughout Indiana and specimens are now in museums from 25
counties (7, 13, and hitherto unpublished records). Reliable sight records
of lost or discarded specimens are at hand from 9 additional counties.
Findley and Jones (1) summarized the seasonal distribution of the
hoary bat throughout its contiguous range, but their data for Indiana are
incomplete. It seems worthwhile, therefore, to summarize Indiana rec-
ords in more detail herein (Table 1). Except for the specimen (J. O.
Whitaker 4003) taken January 31, 1967, at Terre Haute, Vigo County
(14), all records for which dates of collection are known fall between
April 12 and October 5. Most of the reports are for the months of May
to August, probably the period when hoary bats are most abundant in
Indiana. The single January record suggests that a few may winter.
It is generally felt that over the northern part of the range of the hoary
bat (which includes Indiana) the species migrates southward in the fall.
Banding to date has failed to confirm such movement, but sightings and
other circumstantial evidence indicate that migration does occur. The
hoary bat is hardy and wide-ranging, capable of long flights. Its
tolerance to northern winters requires investigation.
Population data are not available, thus the seasonal abundance of
this species in Indiana is unknown. On the evening of May 29, 1948, I
497
498
Indiana Academy of Science
table 1. Number of Records of Lasiurus cinereus in Indiayia
by County, by Month.
County
Jan. Apr. May
June
July
Aug-.
Sept.
Oct.
Month
Un-
known
Blackford
1
Clay
5
2
Franklin
1
Gibson
1
Grant
1
Harrison
1
Howard
1
Jackson
1
Jefferson
1
Jennings
1
Johnson
1
Knox
1
LaGrange
2
Lake
5
Lawrence
3
Madison
1
Marion
1
1
1
Marshall
1
Miami
2
Monroe
1
1
Montgomery 1
Morgan
1
Newton 1
Parke
1
Pike 2
Pulaski 2
St. Joseph
2
1
Zoology 499
table 1 (Continued)
County
Jan.
Apr.
May
June
July
Aug-.
Sept.
Oct.
Month
Un-
known
Tippecanoe
5
4
1
4
Vanderburgh
1
2
3
2
1
Vermillion
1
Vigo
1
1
Wayne
2
2
5
7
1
Wells
1
White
1
Totals
1
3
12
29
23
12
2
1
10
shot two of at least three hoary bats feeding together along the border
of a pastured woodlot near West Lafayette. Thomas W. Hoekstra and
J. Scott Grundy shot five hoary bats at dusk on June 27, 1962, as the bats
flew over a small pond near Munster (Lake County). Hoekstra told me
that he and Grundy stopped shooting at the numerous bats when the five
collected all proved to be Lasiurus cinereus; all bats seen that evening
appeared to be this species. James B. Cope and his students netted and
banded three hoary bats the night of July 12, 1962, near Richmond
(Wayne County). More field work is necessary before the true status
of this bat can be determined.
Habitat. Lasiurus cinereus is thought to be essentially a tree-
inhabiting bat, roosting by day among the foliage. Such known roosting
sites in Indiana include a pasture, residential areas, a wooded lake shore,
a wooded campus, and an open area of mixed woodland, brush, and old
fields. One individual was taken in a building; another was found cling-
ing to the side of a building. There are a few records of skulls and other
remains of hoary bats from Indiana caves (2, 6), but neither a live nor
an intact animal has been discovered in caverns in this state. We would
like to know at what season and under what conditions this bat enters
caves. Most Indiana specimens have been shot or netted about woodlot
borders, along small streams through pasture or cropland, or over ponds
(mostly in non-wooded areas). The eight females taken at West
Lafayette in 1948 were shot around the borders of a pastured, deciduous
woodlot surrounded on three sides by pasture and on the fourth by a
cultivated field. Other species of bats shot at this location include
Eptesicus fuscus, Myotis sodalis, Pipistrellus subflavus, Lasionycteris
noctivagans, Lasiurus borealis, and Nycticeius humeralis. Judging from
the frequency with which hoary bats are submitted to the State Board
of Health for examination, residential areas may be utilized to a
considerable extent.
500 Indiana Academy of Science
Breeding. The first breeding record for Indiana appears to be that
of McAtee (5), who reported a female with two young at Bloomington
(Monroe County) on July 12, 1905. Hahn (3) mentions this same report,
but states that the specimens (now lost) were taken "in June." Addi-
tional breeding data have accumulated slowly, but the presence of hoary
bats in most parts of Indiana in summer suggests that the species breeds
throughout the state. On July 18, 1952, I shot a subadult female (U. S.
National Museum 296436) in Jackson County (8). This bat flew weakly
and evidently had been reared locally. A female (R. E. Mumford 2361)
and two attached young were captured June 15, 1960, at Lafayette.
Another female (Joseph Moore Museum 2009) and two attached young
were taken in Wayne County on June 14, 1963. Single, non-flying young
(JMM 2667, 2686) were taken in Wayne County on June 30 and July 6,
1966. Emery Jenkins found two young (REM 3996, 3997) at Brazil
(Clay County) on June 12, 1964. An adult female (JOW 2613) captured
at Terre Haute on May 28, 1964, gave birth to two young the same day
(13). It is possible that the latter record represents a premature birth
occasioned by the confinement of the female, for none of the eight gravid
females taken at West Lafayette between May 19 and June 8, 1948, had
given birth. An immature (JOW 4836) taken in Vanderburgh County on
June 18, 1968, was probably not yet able to fly (forearm length 46 mm)
and no doubt constitutes a local breeding record.
Lasiurus cinereus normally produces two young per litter; nine
gravid females and three females with attached young from Indiana each
had this number. In contrast, the smaller red bat (Lasiurus borealis)
usually produces three, and sometimes four, young per litter. Parturition
in hoary bats in Indiana probably occurs mostly in late May and the first
half of June. The largest embryos examined, both from a female taken
June 1 in Tippecanoe County, each had a forearm length of 16.5 mm.
Two young, born to a confined female in Vigo County on May 28, had
forearm lengths of 15 and 17 mm. At birth, young probably have fore-
arms measuring from 16 to 20 mm (9, 10). Single flying young taken
July 18 (Jackson County) and July 19 (Howard County) each had
forearm lengths of 53 mm.
From available data, adult male hoary bats are quite rare in Indiana,
especially from April through June. One (JOW 4946) was captured in
Jennings County on May 31, 1968. For the April to June period there
are records of 40 adult females (for a few sight observations the sex is
not known). Likewise, most July and August records for adults are of
females. Some of the males taken in July and August are of questionable
age, but most are clearly young of the year.
Literature Cited
1. Findley, James S., and Clyde Jones. 1964. Seasonal distribution of the
hoary bat. J, Mammalogy, 45:461-470.
2. Hahn, Walter L. 1907. The mammalian remains of the Donaldson Cave.
Proc. Indiana Acad. Sci. 15:142-144.
3. . 1909. The mammals of Indiana. 33rd Ann. Report Ind. Dept.
Geol. Nat. Resources, pp. 419-654.
Zoology 501
4. Lyon, Marcus W., Jr. 1936. Mammals of Indiana. Amer. Midi. Natur.
17:1-384.
5. McAtbe, Waldo L». 1907. A list of the mammals, reptiles and batrachians
of Monroe County, Indiana. Proc. Indiana Acad. Sci. 20:1-16.
6. Mumford, Russell E. 1953. Hoary bat skull in an Indiana cave. J. Mam-
malogy 34:121.
7. Mumford, Russell E., and James B. Cope. 1964. Distribution and status
of the chiroptera of Indiana. Amer. Midi. Natur. 72:473-489.
8. Mumford, Russell E., and Charles O. Handley, Jr. 1956. Notes on the
mammals of Jackson County, Indiana. J. Mammalogy 37:407-412.
9. Munyer, Edward A. 1967. A parturition date for the hoary bat, Lasiurus
c. cinereus, in Illinois and notes on the newborn young. Trans. 111. Acad.
Sci. ©0:95-97.
10. Nero, Robert W. 1958. Hoary bat parturition date. The Blue Jay
16:130-131.
11. Plummer, John T. 1844. Scraps in natural history (Quadrupeds). Amer.
J. Sci. Arts 46:236-249.
12. Provost, Ernest E., and Charles M. Kirkpatrick. 1952. Observations on
the hoary bat in Indiana and Illinois. J. Mammalogy 33:110-113.
13. Whitaker, John O., Jr. 1967. Habitat and erproduction of some of the
small mammals of Vigo County, Indiana, with a list of mammals known
to occur there. Occ. Pap. Adams Center Ecol. Studies, No. 16:24 pp.
14. . 1967. Hoary bat apparently hibernating in Indiana. J.
Mammalogy 48:663.
INSTRUCTIONS FOR CONTRIBUTORS
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Indiana Academy of Science members in good standing- are eligible to
submit papers for publication in the Proceedings. When a paper is signed
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502
Contributors 503
7. Doe, J. B. and R. C. Roe. 1949. New light from old radioactive
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Revised April 28, 1967
INDEX
Abscission of bean leaf, 14 6
Academic origins of geneticists, 370
Aedes infirmatus D&K, new Indiana
record, 257
Alexander, R. W., Jr., 88
Algae, sewage ponds, 139
Allee Woods, Collembola, 231
Allen, Frederick John, memorial, 32
Allen, P. R., 210
Aminoglutethimide, effects on serum
electrolytes, 468
Antibiotics, Thiadiketopiperazine, 111
Antibody plaque technique, 109
Antibody studies, 163
Appalachia, culture of, 97
Araneae of Indiana, an annotated list,
266
Archaeology of Eastern Siouan Tutelo,
88
Armington, John H., memorial, 32
Arnett, P. M., 231
Artificial sweeteners, rodents' prefer-
ence for, 457
Atomic lifetimes, measurement of, 3S9
Awasthi, Y., 189
Bacillus subtilis growth and sporula-
tion, 111
Bacteria in farm ponds, 417
Ball State University science depart-
ment, 381
Barber, S. A., 418
Barski, G., 165
Barton, T. F., 342
Bats, incidence of rabies in, 447
Beech-maple climax, Collembola pres-
ent in, 231
Beesley, L. & A., 118
Beetles (Hydrophilidae), 260
Big Walnut Creek Reservoir, resolu-
tion, 13
Blakely, R. L., 73
Blanc, T. V., 389
Blood, Gc system, 164
Bochrath, R., 109
Bone growth, prehistoric Indians, 83
Boneham, R. F., 329
Boyko, W. L., 447
Brahmi, Z., 109
Brannon, D. R., Ill
Brett, W. J., 44 5
Bryophyta, 396
Burgess, R. D., 387
Burns, J. M., 445
Burton, Everett Tyler, memorial, 33
Butler, J. W., 165
By-laws, revision, 14
Callis, R., 387
Campbell, W. F., 110
Carlson, K., 109
Cell membrane composition, 161
Cell mobility, 165
Centaurium pulchellum, 414
Cephalosporium genetics, 112
Cerebral cortex, neuron structure, 161
Chara, fossils, 406
Cheetham R., 183
Chi square, statistical approximations,
482
Chiroptera, incidence of rabies in, 44 7
Chiroptera, the Hoary Bat, 497
Chlorophyll variation in corn, 118
City planning and web pattern, 342
Cleveland, J. H., 315
Cline, J. C, 112
Coal balls, 120
Coal mine reclamation costs, 355
Cobra venom action on virus, 110
Coffin, S., 71
Coincidence techniques in measure-
ment of atomic lifetimes, 389
Collembola in beech-maple climax
forest, 231
Communication, scientist to non-scien-
tist, 65
Community development, systems ap-
proach, 104
Computers in education, 200
Cook, A. G., 200
Cooper, R. H., 381, 466
Copeland, R. F., 200
Corn diseases in Indiana, 117
Corn, response to phosphorus and po-
tassium fertilization, 435
Correlation coefficients, statistical ap-
proximations, 482
Craig, E., 387
Craig, M. L., 72
Crane, F. L., 189
Crankshaw, W. B., 201, 204, 241
Craske, Al G., Jr., 2 01
Crop plants on fragipan soils, 429
Crushed stone, transportation costs,
348
Curren, T., 71
Curtis, K. K., 118
Cuticle variability in plants, 115
Cyanamide complex of iron, 199
D* absolute value, 387
Daily, F. K., 32, 406
Danemar, A. G., 38 9
Data analysis, life sciences, statistical
shortcuts, 482
Dendropanax in the Eocene, 115
505
506
Index
Desiccation, response to, in salaman-
ders, 472
Desmognathus ochrophaeus carolinen-
sis, water loss physiology in, 47 2
Devonian rocks, 333
Dickson Mounds, Fulton County, Illi-
nois, 73
Dilcher, D. L., 115
Dillon, L. I., 363
Dinkel, R. M., 201, 355
Dolph, G. E., 115
Donahue, S., 161
Downey, R. J., 113
Doyle, J. R., 200
Drosophila periodicity and rhythm,
445
Earth mound, 417
Earth science teaching in Indiana, 320
Eclipse, solar, 387
Education, the non-science major, 65
Elaphe obsoleta, 446
Electrolytes, potassium and sodium,
468
Enamine synthesis and properties, 200
Endoplasmic reticulum, 183
Escherichia coli, mutant, 109
Esten, Mable Henniger, memorial, 34
Esterline, A., 256
Fertilizer experiments with corn, 435
Fixation, glutaraldehyde, 167
Food habits, of Rana catesbeiana, 491
Forest analysis, 245
Forest ecology, 2 01, 204
Forests, old-growth, 210
Forest succession, 2 03
Fort Ancient, archaeological popula-
tion, 93
Fossils, charophytes, 406
Fragipan soils and suitable crops, 429
French, R. R., 348
Fulk, F. D., 491
Galileo's telescopes, air thermometer,
and astrolab, 378
Gammon, J. R., 203
Garter snake, 445
Genetics Society of America, academic
origins of members, 370
Gentianaceae, 414
Geologic reports for land-use planning
studies, 49
Geomorphology of Indiana, 317
Gerber D., 183
Gerwig, J. D., 201
Geyer, R. E., Jr., 416
Girton, R. E., 116
Glenn, E. J., 72
Glutaraldehyde, reactions with cyto-
membranes, 167
Golgi apparatus, 162, 1S3
Gordon, R. E., 65
Gorman, M., Ill
Greene County, Indiana, 355
Gruenholz, A. W., 446
Guard, A. T., 117
Guernsey, L., 3 55
Gunther, W. C, 457
Gypsum resources in U.S., 315
Habart, T., 72
Hall, J. D., 189
Hart, J. W., 257
Haslanger, Pearl C, memorial, 35
Heath, M. E., 429
Hecht, T. A., 200
Heiser, C, 119
Heloderma svspcctum, tumor in, 466
Hemmer Woods, Gibson County, In-
diana, 2 45
Herzog, L., 256
Hinsman, E. J., 161
Holler, F. J., 199
Holmes, R. E., 112
Hookeriaceae species in South Amer-
ica, 3 96
Hopp, W. B., 446
Hormones in chickens, 445
Houlihan, J. F., 3 87
Howard, D., 370
Huber, R. T., 2 56
Huelsman, B. R., 97
Hughes, L. E., 417
Hults, M., 387
Humbles, J., 413
Humeral septum, performation of, 73
Hunter, K. B., 72
Huygens' pendulum clock and plane-
tarium, 3 7S
Indiana Academy of Science, bylaws,
changes, 14
Indiana fossil plants, 120
Indiana plant distribution records, 413
Indiana spiders, annotated list, 266
Indians, Shawnee, origin, 93
Ingraham, J. S., 109
Jackson, M. T., 210, 2 45
Japanese Weevil in Indiana, 255
Jersild, R. A., 161, 163
Joyner, R. D., 199
Judd, R. W., 120
Junior Academy of Science, 22
Kendall, S. H., 113
Kirkpatrick, R. D., 241
Kohnke, H., 418
Koltenbah, D. E., 38 8, 389
Kramer, D. C, 445
Kuc', J., 118
Index
507
Land use planning', 49
Landform properties, 317
Lasiurus cinereus, in Indiana, 497
Late Woodland American Indians, S3
Lawrence, R. M., 199
Leaf abcission, environmental regula-
tion, 146
Leewenhoek's microscopes, 37 8
Lemke, P. A., 112
Letnna minor, the flowering of, 414
Lemnaceae, 414
Leukemia suppression by L-Aspara-
ginase, 110
Levine, A. S., 109, 110
Lewis, L. A., 317
Lichen planus, 162
Lobelias of Indiana, 118
Logansport Sag, 333
Louraine, Frank E., memorial, 3.")
Mabe, J., Ill
Mak, K. M., 161
Markle, C. A., 395
Markle, Millard S., memorial, 37
Marks, G. C, 414
Marmouze, R. J., 73
Maya Indians, 71
McGrew, L. A., 199
McKee, Roy, memorial, 36
McKelvey, P. T., 202
Melanoma in Helodermu suspectum,
466
Members (1968), 43
Membranifibrils, 189
Membranes, fibrils in mitochondria
and chloroplasts, 189
Merritt, Neal R., memorial, 39
Mertens, T. R., 370
Meson scattering, 38 8
Michael, R. L., 71
Middle Mississippian American In-
dians, 73
Middleton, A. E., 183
Miller, B., 417
Miller, W. A., 44 7
Mineral aggregates, transportation
costs, 348
Minton, S. A., 113
Mitchell Plain deposits, 316
Moe, K., 16
Mollenhauer, H. H., 167
Molloy, B. B., Ill
Montgomery, B. E., 2 56, 3 69
Morgan, F., 202
Morgan, P. C, 111
Morre', D. J., 146, 161, 167, 183
Morris, B. J., 71
Mosquitoes, Indiana check list, 25 7
Mosses, 396
Mound (White Site) excavation, 71
Mucous velocities in trachea, 446
Mumford, R. E., 497
Muon energy study, 387
Mycophenolic acid, 112
Neumann, G. K., 72, 88, 93
Neuron, fine structure, 161
New Castle Site, excavation, 71
Niederpruem, D. J., 163
Nisbet, J. J., 120
Nitrate reductase synthesis, 113
Nuclear resonance analysis, 3SS
Nuner, J. H., 113
Organonitrile complexes, 200
Orr, R. W., 333
Osmun, J. V., 256
Pace, R. E., 71
Paleobotany, 120
Palmer, C. M., 139
Parker, T. A., 266
Pennsylvanian Mora, 12 0
Peri, B., 110
Petersen, B. H., 109
Petty, R. O., 203
Phaseolus resistance to anthracnose,
US
Phaseolus vulgaris, leaf abscission,
146
Phenyl isocyanate reaction with water,
199
Phosphatases, nucleotide, 183
Phthalocyaninogermanium compounds,
199
Pickard, B. L., 472
Pilot Black Snake growth, 446
Plant distribution records, 413
Plant species disappearing, 117
Plasma membrane, 183
Pleistocene drift in Indiana, 315
I'lethodon glutinosus, water loss physi-
ology in, 472
I'lethodon jordani jordani, water loss
physiology in, 472
Pollock, G. P., 468
Pollution effect on fish, 2 02
Pollution effect on organisms, 203
Pollution-tolerant algae, 139
Popcorn production in Indiana, 363
Powell, R. L., 316
Prairie soils, tillage techniques, 418
Pseudocneorhinus. bifasciatus, 255
Rabies in Indiana bats, 447
Rahe, J. E., US
Raitano, A. C, 110
Ramaley, R. F., Ill
Rana catesbeiana, food habits, 491
Reclamation practices, strip coal mine,
355
Reptile serum profiles, 113
Reservoir limnological studies, 202
508
Index
Reuszer, H. W., 417
Reynolds, A. E., 472
Richardson, J., 71
Robbins, L. M., 93
Robling, S. C, 199
Rodents' preference for artificial
sweeteners, 457
Rosene, G. L., 164
Rothwell, F., 201
Salovesh, M., 71
Schizophyllum commune, nucleus, 163
Schmedtje, J. F., 163
Schneider, A. F., 315
Schuder, D. L., 255
Schultz, D. J., 200
Science and Society Committee re-
ports, 13, 16
Science apparatus originals ; Florence,
Leyden, Glasgow, and Cambridge,
378
Science at Ball State University, 381
Science history ; astronomy and
physics, 378
Secondary education, earth science, 3 29
Selenium effect on respiration, 116
Sewage stabilization ponds, algae, 139
Shawnee Indians, 93
Siverly, R. E., 256
Smalley, S. F., 164
Smith, C. E., Jr., 202
Smith, D. E., 118
Smith, J. A., 241
Smith, J. M., 417
Soil moisture, forest, 204
Soils, fertilizer response of corn, 43 5
Soils (fragipan) and suitable plants,
429
Southern mountaineers, migration of,
97
Spectroscopy, atomic, in the measure-
ment of atomic lifetimes, 3 89
Spiders of Indiana, 2 66
Sprague, N. G., 378
Stark, W. M., Ill
Statistical approximations, Chi Square,
correlation coefficients, 4 82
Stiles, J. W. 1S9
Stivers, R. K., 43 5
Storhoff, B. N., 2 00
Stovall, W. E., 164
Stover, R. L., 199
Stratigraphy, Devonian rocks, 333
Streptococcus faecalis. immunity in rat,
110
Strip coal mine reclamation, 355
Strip mine areas, temperature and
moisture, 201
Sweeney, M. J., 112
Sweeteners, toxic and non-toxic, as
preferred by rodents, 457
Tague, N. A., 72
Tamar, H., 446
Teaching earth science in Indiana, 329
Thallium chloride, 199
Thamnophis sirtalis, 445
Thermal discharge, in river, 201
Thomas, G. P., 388
Thomas, R. J., 165
Tiefel, C. F., 315
Tillage techniques, 4 IS
Tolypella, fossils compared to modern,
405
Tomak, C. H., 72
Transportation costs, mineral aggre-
gates in Indiana, 3 48
Trees, site preference, 201
Tropistemus collaris and supspecies,
genetics, 260
Tumor cells, mitotic activity, 164
Tutelo Indian tribe of North Carolina,
88
Ullstrup, A. J., 117
Urban development and planning, 342
Urban geology, research programs, 49
Urban structure analysis, 104
Van Nuys Site, excavation, 71
Versailles State Park, Indiana, 210
Wagner, M., 110
Wagner, M. W., 457
Walker, P. L., 83
Wallace, Frank N., memorial, 40
Watanabe, I., 161
Wayne, W. J., 49
Web pattern in urban development,
342
Weiss, M. S., 104
Welch, W. H., 396
Welsh-Dunlap Site, excavation, 71
Wendt, H. W., 4S2
Whitaker, J. O. Jr., 447, 491
Whitehead, J. M., 72
Whitten, J. B., Jr., 161
Wildlife management, 241
Williams, R. H., ]12
Woodcock singing ground descriptions,
241
Wynne, D. D., 73
Young, F. N., 260
Yunghans, W., 161
Zea maris L.. response to fertilization,
435
Zeck, C. A., 115
Zeman, W., 161
Zimmack, H. L., 2 56
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