Skip to main content

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

See other formats 


Digitized  by  the  Internet  Archive 

in  2012  with  funding  from 

LYRASIS  Members  and  Sloan  Foundation 


http://archive.org/details/proceedingsofin791969indi 


PROCEEDINGS 

of  the 

Indiana  Academy 
of  Science 

Founded  December  29, 1885 


Volume  79 
1969 


Marion  T.  Jackson,  Editor 

Indiana  State  University 

Terre  Haute,  Indiana  47809 


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

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

Published  at  Indianapolis,  Indiana 
1970 


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

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

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

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

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

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

6.  The  Constitution  and  By-Laws  reprinted  from  v.  74  are  available  to  members 
upon  application  to  the  Secretary.  Necrologies  reprinted  from  the  various  volumes  can 
be   supplied  to   relatives   and   friends   of   deceased   members    by   the   Secretary. 

7.  Officers  whose  names  and  addresses  are  not  known  to  correspondents  may  be 
addressed  care  of  the  State  Library.  Papers  published  in  the  Proceedings  of  the  Academy 
of  Science  are  abstracted  or  indexed  in  appropriate  services   listed  here : 

Annotated  Bibliography  of  Economic  Geology 

Bibliography  of  Agriculture 

Bibliography  of  North  American  Geology 

Biological  Abstracts 

Chemical  Abstracts 

Chemisches  Zentralblatt 

Current  Geographical  Publications 

Geological  Abstracts 

Metallurgical  Abstracts 

Pesticides  Documentation  Bulletin 

Psychological  Abstracts 

Review  of  Applied  Entomology 

The  Torrey  Bulletin 

Zoological  Record 


TABLE  OF  CONTENTS 

Part  1 

THE  WORK  OF  THE  ACADEMY 

Page 

Officers  and   Committees  for  1969    3 

Minutes  of  the  Spring  Meeting 6 

Minutes  of  the  Fall  Meeting  (Executive  Committee)    9 

Minutes  of  the  Fall  Meeting  (General  Session)    12 

Annual  Financial  Statement    14 

Annual  Report,  Junior  Academy  of  Science 16 

Biological  Survey  Committee  Report  23 

Necrology     27 

New  Members  for  1969   39 

Part  2 

ADDRESSES  AND  CONTRIBUTED  PAPERS 

Presidential   Address    45 

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

Anthropology 

E.    Dolan — Use    of    Historical    and    Archaeological    Evidence    to 

Reconstruct   the    Cherokee    Past*     57 

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

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

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

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

A  Probable  Late  Woodland  Occupation*   58 

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

Western  Origin  of  the  Delaware  Indians   60 

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

of  the  Illinois  Hopewell  Indians   62 

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

Extinctions    65 

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

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

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

Indiana     75 


: Abstract  or  Note  only 


iv  Indiana  Academy  of  Science 

Botany  Page 

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

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

of  Cyanidium  caldariwm*   83 

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

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

in    Indiana*     83 

L.  Ford — Chromosome  Associations  in  Corn  Monoploids*  84 

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

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

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

Gramineae     85 


Cell  Biology 

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

to  the  Administration  of  Thiouracil*   91 

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

Skeletal  Muscle  in  Vitamin  E-Deficient  Rabbits*    91 

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

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

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

Human   Enteric   Epithelial   Cells*    92 

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

on  the  Morphology  of  Heart  Mitochondria*   93 

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

Brain*    93 

S.    Yumoto   and    S.   Ishikawa — Ultrastructural   and    Enzymological 

Observations  of  Isolated  Kidney  Microvilli*   93 

J.     B.     Whitten — Ultrastructural     Features     of     the     Calcifying 

Epithelial   Odontogenic   Tumor*    94 

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

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

Activities  of   Rat  Liver   Cytomembranes    107 

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

Membrane:  Cytochrome  Oxidase   110 


*Abstract  or  Note  only 


Table  of  Contents  v 

Chemistry  Page 

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

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

Coordination  Compounds*    121 

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

on  the  Flavin  Enzyme  L-Amino  Acid  Oxidase*    121 

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

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

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

Dependence  of  Eo  for  the  Daniell  Cell   123 

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

Ecology 

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

of   the    Bluegill*    135 

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

Growth    of    Bluegill*    135 

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

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

on    Growth    of    Bluegill    with    Varied    Feeding    Frequency    and 

Constant   Ration*    136 

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

Growth  and   Social  Hierarchy  in  the  Green  Sunfish*    136 

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

County,    Indiana*    137 

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

Factors  on  the  Concentration  of  Hydrocyanic  Acid  in  Manihot 

esculenta*      137 

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

Expressions  for  Tree   Species  in   Foley  Woods,  Edgar  County, 

Illinois*    137 

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

Old-growth  Forests*  138 

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

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

Indiana    Woodcocks     162 

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

noveboracensis,  from  Pike  County,  Indiana   172 


*Abstract  or  Note  only 


vi  Indiana  Academy  of  Science 

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

and  Behavior  of  Indiana  Ruffed  Grouse    177 

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

through   1968    187 

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

Ecotypes  Discovered  in  Crawford  County,  Indiana    193 

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

and    a    New    Method    of    Classifying    Midwestern    Hardwood 

Forests     198 

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

the  Commercially  Valuable  Mussels  of  the  Wabash  and  White 

Rivers   of   Indiana    205 

Entomology 

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

Potential  Screening  Technique  for  Insect  Pathogens*    227 

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

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

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

Citations      229 

Gertrude    L.    Ward — The    occurrence    of    Chalybion    zimmermanni 

Dahlbom    (Sphecidae)    in   Indiana    231 

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

Serai  Stages  Leading  to  the  Beech-Maple  Climax   234 

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

Mosquito   Populations   in    Indiana    238 

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

Geology  and  Geography 

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

Miocene*    253 

A.  H.  Siddiqi — Urbanization  in  Asia*    253 

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

Shale   of   Southern   Indiana    254 

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

Indiana     263 

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

Hadley  Lake  Depression,  Tippecanoe  County,  Indiana   270 


*  Abstract  or  Note  only 


Table  of  Contents  vii 

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

Karst  Groundwater  Zones  in  South-Central  Indiana 281 

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

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

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

of    the    Neighborhood    Grocers    in    Terre    Haute,    Indiana:    A 

Geographic   Analysis    309 

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

Indiana 318 

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

of  Central  Indiana  to  the  Walter  Christaller  Model   325 

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

Microbiology  and  Molecular  Biology 

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

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

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

Dictyostelium  discoideum   Spores*    345 

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

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

Rate  Equations  for  Enzyme-catalyzed  Reactions*    346 

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

Materials  in  Nerve  Fibers*    346 

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

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

Physics 

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

Lacunarity   Technique*    357 

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

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

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

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


'Abstract  or  Note  only 


viii  Indiana  Academy  of  Science 

Page 

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

Horse   Shoe   Lake*    359 

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

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

C.   C.    Sartain — Semiconductors   Produced  by   Doping  Oxideglasses 

with  Ir,  Pd,  Rh  or  Ru    361 

Plant  Taxonomy 

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

Fossil   Fungal   Spores*    375 

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

Winona    H.    Welch — Hookeriaceae    Species    and    Distribution    in 

Africa,  Europe,  Asia,  Australia  and  Oceania   377 

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

Ferns    388 

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

onomic  Notes  on  Genus  Polygonum,  Section  Polygonum   396 

Soil  Science 

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

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

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

on  Water  Infiltration  into  Soils    407 

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

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

Populations  in  Indiana  Farm  Pond  Waters   423 

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

Insecticides  on  Pond  Sediments  and  Watershed  Soils    432 

Zoology 

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

ing  Aminoglutethimide    Treatment*    439 

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


♦Abstract  or  Note  only 


Table  of  Contents  ix 

Page 

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

in   Vigo   County,   Indiana    441 

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

of  the  House  Mouse,  Mus  musculus  449 

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

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

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

with  the  Big  Brown  Bat   (Eptesicus  fuscus)    466 

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

Indiana     470 

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

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

Instructions  for  Contributors 484 

Index    486 


*  Abstract  or  Note  only 


PART  1 

THE  WORK 

OF  THE 
ACADEMY 

1969 


Howard  R.  Youse,  President 


Officers  and  Committees  for  1969 

OFFICERS 


President Howard   R.   Youse,   DePauw   University 

President-elect Frank  A.  Guthrie,  Rose  Polytechnic  Institute 

Secretary James  R.  Gammon,  DePauw  University 

Treasurer Damian  Schmelz,  St.  Meinrad  College 

Editor William   R.   Eberly,   Manchester   College 

Director  of  Public  Relations Paul  E.  Klinge,  Indiana  University 

DIVISIONAL  CHAIRMEN 

Anthropology Robert  E.  Pace,  Indiana  State  University 

Botany Robert  L.  Kent,  Indiana  Central  College 

Cell  Biology Edward  J.  Hinsman,  Purdue  University 

Chemistry James  E.  George,  DePauw  University 

Ecology Thomas  S.  McComish,  Ball  State  University 

Entomology Jack  R.  Munsee,  Indiana  State  University 

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

History  of  Science B.  Elwood  Montgomery,  Purdue  University 

Microbiology  and  Molecular  Biology D.  S.  Wegener, 

Indiana  University  Medical  Center 

Physics Richard  L.  Conklin,  Hanover  College 

Plant  Taxonomy Jack  E.  Humbles,  Indiana  University 

Soil  Science   James  E.  Newman,  Purdue  University 

Zoology James  C.  List,  Ball  State  University 


EXECUTIVE  COMMITTEE 

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


*Baldinger,  L.  H. 

Behrens,  0.  K, 

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

Coats,  N. 

Conklin,  R.  L. 

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

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

Gammon,  J.  R. 

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

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


Heniser,  V. 

Hinsman,  E.  J. 

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

Kaufman,  K.  L. 

Kent,  R.  L. 

Kessel,  W.  G. 

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

List,  J.  C. 

McComish,  T. 
*Markle,  C.  A. 

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

Montgomery,  B. 


"Morgan,  W.  P. 

Moulton,  B. 

Munsee,  J.  R. 

Newman,  J.  E. 

Nisbet,  J. 

Pace,  R.  E. 

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

Schmelz,  D. 

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

Webster,  J.  D. 

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

Winslow,  D. 

Youse,  H.  R. 


4  Indiana  Academy  of  Science 

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

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

COMMITTEES  APPOINTED  BY  THE  PRESIDENT 

(President  an  ex  officio  member  of  all  committees) 

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

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

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

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

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

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

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

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

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

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

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


Officers  and  Committees  5 

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

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

Necrologist:  F.  K.  Daily. 

Parliamentarian:  P.  Weatherwax. 

SPECIAL  COMMITTEES  APPOINTED  BY  THE  PRESIDENT 

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

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

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

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


SPRING    MEETING 

Hanover  College,  Hanover,  Indiana 

MINUTES  OF  THE  EXECUTIVE  COMMITTEE  MEETING 

April  25,  1969 

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

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

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

Academy  Accounts 

Balance  as  of  January  1,  1969 $  8,719.46 

Receipts  through  April  22,  1969 4,147.40 

Expenditures  through  April  22,  1969 2,855.02 

Balance  as  of  April  22,  1969 10,011.84 

Administered  Accounts 

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

Receipts  through  April  22,  1969 12,125.00 

Expenditures  through  April  22,  1969   10,441.66 

Balance  as  of  April  22,  1969 16,500.47 

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

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

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

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

Johnson,  our  representative,  attended  the  1968  meeting. 

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

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

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

6 


Minutes  of  the  Executive  Committee  7 

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

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

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

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

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

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

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

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

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

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

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


8  Indiana  Academy  of  Science 

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

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

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

Adjournment  at  5:35  pm. 

Approved  October  23,  1969. 

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


FALL    MEETING 

MINUTES  OF  THE  EXECUTIVE  COMMITTEE  MEETING 

Hanover  College,  Hanover  Indiana 

October  23,  1969 

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

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

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

January  1,  1969  balance    $23,536.59 

Income  to  October  17,  1969  26,126.28 

Expended  to  October  17,  1969   17,738.06 

Balance,  October  17,  1969 31,924.81 

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

October  1,  1968  balance $  1,050.20 

Receipts  through  September  30,  1969 725.32 

Disbursements  through  Sept.  30,  1969   1,302.46 

Balance,  September  30,  1969 473.06 

In  the  John  S.  Wright  Fund: 

Balance,  October  1,  1968 $  1,466.33 

Receipts  to  September  30,  1969 11,351.42 

Disbursements  to  Sept.  30,  1969   12,605.72 

Balance  as  of  September  30,  1969 212.03 

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

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

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

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

9 


10  Indiana  Academy  of  Science 

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

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

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


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

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

Robert  I.  Fletcher 

John  A.  Leighty 

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

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

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

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

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

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

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


Minutes  of  the  Executive  Committee  11 

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

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

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

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

The  meeting  was  adjourned  at  9:20  pm. 

Approved  October  24,  1969. 

James  R.  Gammon,  Secretary 


MINUTES  OF  THE  GENERAL  SESSION 

October  24,  1969 

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

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

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

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

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

Article  II,  Section  8  to  be  added: 

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

This  amendment  was  approved. 

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

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

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

12 


Minutes  of  the  General  Session  13 

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

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

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

The  meeting  was  adjourned  at  9:00  pm. 

Approved  May,  1970. 

James  R.  Gammon,  Secretary 


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

I.    ACADEMY  ACCOUNTS 

Income  Expenditure       Budgeted 

Dues    $  3,906.00 

Reprints   1,847.10  $   1,828.73         $       150.00 

Publication    476.17 

Editor    $      400.00  400.00 

Mailing 76.17  100.00 

Secretary    360.55  300.00 

Clerical 354.28 

Postage,   etc.   6.27 

Treasurer     222.23  225.00 

Clerical    61.25 

Postage,   etc.    160.98 

Office    120.56  175.00 

Travel,   Dues   170.10  180.00 

President's   Fund   100.00 

Membership    Committee    18.00  75.00 

Junior     Academy     22.18  150.00 

Program    Committee    472.43                 650.00 

Chairman   66.08 

Printing    212.09 

Mailing 194.26 

Transfer  to  Administered 

Accounts:     2,100.00 

Science  and   Soc.    600.00  600.00 

Library    Binding    1,000.00  1,000.00 

Publication     500.00  500.00 

Transfer  from  Administered 

Accounts:    22.03 

Interest    on    Savings    1,070.70 

Miscellaneous      43.05  43.05 

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


14 


Financial  Report 


15 


II.     ADMINISTERED  ACCOUNTS 


Jan.  1 
Balance 

Sciene    Talent    $  1,995.80 

Science   Fair   7,515.99 

Science  and  Society 466.42 

Research     307.67 

J.   S.   Wright  Library 134.28 

Lilly  III  Library 4,374.94 

Publication     

Proceedings    

Natural    Features    

Academy    Budget    

Library  Binding  

NSF  Funded  Project: 

Science  to   People 

Transfer  to  Academy 

Account    from    Miscellaneous    22.03 


1969 

1969 

Expen- 

Dec. 31 

Income 

diture 

Bal. 

$  1,860.45 

$       135.35 

$10,730.00 

5,545.99 

12,700.00 

600.00* 

554.53 

511.89 

6,659.35 

6,533.50 

433.52 
134.28 

34.88** 

1,487.00 

2,922.82 
1,029.50 

217.00* 

312.50* 

500.00* 

1,000.00* 

1,000.00 

11,663.33 

4,492.61 

7,170.72 

22.03 


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


$26,038.08 


♦Transfer  from  Academy  Accounts 
** Payment  cancelled  on  check 


III.     SUMMARY 

Academy  Administered  Total 

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

1969    Expenditure    5,834.00  20,496.11  26,330.11 

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

1968     Balance     8,719.46  14,817.13  23,536.59 

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

BANK  BALANCES 

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

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

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

Total    Assets    in    All    Accounts    $35,812.42 

Rev.  Damian  Schmelz,  Treasurer 
December  31,  1969 


January  30,  1970 

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

W.  G.  Kessel 
J.  L.  Guernsey 


INDIANA  JUNIOR  ACADEMY  OF  SCIENCE 
OFFICERS 

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

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

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

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

STATE  DIRECTOR 

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


PROGRAM 

Thirty-Seventh  Annual  Meeting 
Saturday,  November  1, 1969 

8:00  am-10:00  am 

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

9:30  am 

Tours  of  several  academic  facilities.  Check  registration  area. 

9:30  am 

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

11:30  am-1:00  pm 

Lunch — Purdue  Union  Cafeterias. 

1:00  pm 

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

16 


Junior  Academy  of  Science  17 

1:30  pm 

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

3:00  PM 

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


PROGRAM  OF  PAPERS 

1.  The  Origin  of  the  Universe. 

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

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

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

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

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

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

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

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

6.  The  Characterization  of  RE  Virus. 

Mark  Coates,  Madison  Consolidated  High  School,  Madison. 

7.  Comparative  Analysis  of  Commercial  Dentrifices  Containing  Fluoride 
Compounds. 

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

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

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

9.  Drugs  Cause  Damage. 

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

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


18  Indiana  Academy  of  Science 


ACKNOWLEDGMENT 


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


MINUTES  OF  THE  THIRTY-SEVENTH  ANNUAL 

MEETING  OF  THE 

INDIANA  JUNIOR  ACADEMY  OF  SCIENCE 

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

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

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

Dennis  Waltke    President 

Timothy  O'Leary    Vice-President 

Rachel  Koontz    Secretary 

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

Rachel  Koontz  President 

Mark   Coates    Vice-President 

John  Farrell    Secretary 

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


Junior  Academy  of  Science  19 

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

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

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

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

Respectfully  submitted, 

Rachel  Koontz,  Secretary 
Dennis  Waltke,  President 


INDIANA  JUNIOR  ACADEMY  OF  SCIENCE 
1969-70 

Town  Club  and  School  Sponsor 

Acton  Sigma  Mu  Chapter  of  FSA,  Frank-       Margaret  Richwine 

lin  Central  H.   S. 

Bedford  Bedford  Science  Problems  Research      Paul  Hardwick 

Group,  Bedford  H.  S. 

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

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

Bloomington     MSE   Academy,   University   Junior       Charles  Souers 
High 

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

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

Columbus  Science  Club,  Columbus  Senior  L.  N.  Carmichael 

H.  S. 

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

ville 

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

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

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

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

Springs  Valley  H.  S. 

Gary  Andrean     Biology     Club,     Andrean       Sr.  Marie  Antoine 

H.  S.  SS.C.M. 

Sr.  Marie  Carmel 
SS.C.M. 

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

SS.C.M. 

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

20 


Junior  Academy  of  Science  21 

Town  Club  and  School  Sponsor 

Griffith  Griffith   Junior   High   Science   Club       Fred  Meeker 

Griffith  Junior  H.  S. 

Griffith  Griffith  Junior  High  Science  Club,       Geraldine  R.  Sherfey 

Griffith  Senior  H.  S. 

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

H.  S. 

Hartford  City  Hartford  City  H.   S.   Science  Club,      

Hartford  City  H.   S. 

Highland  Science   Club,  Highland   H.    S.  Jon  Hendrix 

Hobart  Hobart  Senior  High   Science   Club,       Stanley  J.  Senderak 

Hobart  Senior  H.  S. 

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

Huntington       Science,  Huntington  H.  S.  Robert  Diffenbaugh 

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

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

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

Indianapolis      Science  Club,  Howe  H.  S.  Jerry  Motley 

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

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

Indianapolis      Roncalli  H.  S.  Sr.  Mary  Alexandra 

C.S.J. 

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

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

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

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

Byron  Bernard 

Lebanon  Junior  Explorers  of  Science,  Leba-       Tom  Evving 

non  Junior  H.  S. 


22 


Indiana  Academy  of  Science 


Town  Club  and  School 

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

Madison  Madison     Science     Club,     Madison 

Consolidated  High 

Muncie  Muncie  Central  Science  Club,  Mun- 

cie  Central  H.  S. 

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

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

Portland  Science  Club,  Portland- Wayne 

Township  Junior  H.  S. 

Portland  Portland  Senior  H.  S.  Science  and 

Mathematics    Club,   Portland   Sr. 
H.  S. 

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

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

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

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

South  Bend       LaSalle  High  School 

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


Tipton  Tipton  H.  S.   Science  Club,  Tipton 

H.  S. 

Trafalgar  Indian  Creek  School 

Vincennes  Sigma  Tau  Science  Club,  St.  Rose 

Academy 


Sponsor 
Raymond  T.  Kozer 

David  Dunkerton 

William  Beuoy 

Roger  Moody 


Keith  Hunnings 
E.  H.  Sanders 

Mary  Zehner 


Ralph  Settle 
Robert  Freemyer 

Ernest  Litweiler 


John  V.  Davis 
John  Marker 


Sr.  Marie  Barbara 
S.P. 

Richard  Garst 
Frederick  Calhoun 


Sr.  Anna  Margaret 
Sr.  Aloyse 


BIOLOGICAL  SURVEY  COMMITTEE 

J.  Dan  Webster,  Chairman,  Hanover  College 

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


Virales : 


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


Tracheophyta 


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

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

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

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

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

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

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


Protozoa : 


Worms  and  Arthropods 


Crustacea : 


Insecta  : 


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

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

448. 

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

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

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

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

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

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

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

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

23 


24  Indiana  Academy  of  Science 

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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


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

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

system.  Ph.D.  Purdue. 

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

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


Biological  Survey  Committee 


•if. 


Pisces 


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

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

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

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


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


Algae : 


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


Reptilia  and 
Amphibia  : 
Mammalia  : 


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

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

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

(Geomys  bursarius)    in  Indiana. 


NECROLOGY 

Fay  Kenoyer  Daily,  Butler  University 

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

March  6,  1912  November  7,  1969 

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

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

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

March  5,  1896  June  30,  1969 

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

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

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

27 


28  Indiana  Academy  of  Science 

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

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

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

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

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

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


Necrology  29 

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


Leslie  Willard  Freeman 
Escanaba,  Michigan  Indianapolis,  Indiana 

August  17,  1915  July  7,  1969 

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

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

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

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

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


30  Indiana  Academy  of  Science 

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

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

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

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


Vaclav  Hlavaty 
Louny,  Czechoslovakia  Bloomington,  Indiana 

January  27,  1894  January  11,  1969 

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

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


NECROLOGY  31 

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

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

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

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

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

Ruth  Jordan 
Danville,  Indiana  Lafayette,  Indiana 

July  4,  1891  July  14,  1968 

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

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


32  Indiana  Academy  of  Science 

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

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

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

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


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

January  13,  1924  October  6,  1969 

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

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


Necrology  33 

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

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

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

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


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

December  9,  1903  January  26,  1969 

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

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

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

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


34  Indiana  Academy  of  Science 

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

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

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


William  Lowell  Toms 
Hancock  County,  Indiana  Greenfield,  Indiana 

December  8,  1896  August  20,  1969 

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

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


Necrology  35 

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

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

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

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

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


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

October  23,  1883  December  28,  1968 

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


36  Indiana  Academy  of  Science 

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

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

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

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

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

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

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


Necrology  37 

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


Howard  Ford  Wright 
Wharton,  Ohio  Indianapolis,  Indiana 

December  27,  1904  October  25,  1969 

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

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

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

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


38  Indiana  Academy  of  Science 

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

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

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


NEW  MEMBERS  FOR  1969 

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

A — Anthropology 

B — Botany 

C — Chemistry 

E — Entomology 

G — Geology  and /or  Geography 

H — History  of  Science 

L — Ecology 

M — Mathematics 

O— Cell  Biology 

P — Physics 

R — Bacteriology 

S — Soil  Science 

T— Plant  Taxonomy 

Y — Psychology 

Z — Zoology 

D — Secondary  Science  Teaching 

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

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

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

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

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

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

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

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

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

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

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

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

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

39 


40  Indiana  Academy  of  Science 

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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


New  Members  41 

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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


PART  2 

ADDRESSES 

AND 

CONTRIBUTED 

PAPERS 


Hanover,  Indiana 
October  24,  1969 


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


PRESIDENTIAL  ADDRESS 


The  World  of  the  Honey  Bee 

Howard  R.  Youse 

DePauw  University,  Greencastle,  Indiana 

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

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

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

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

45 


46  Indiana  Academy  of  Science 

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

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

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

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

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


Presidential  Address  47 

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

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

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

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


48  Indiana  Academy  of  Science 

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

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

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


Literature  Cited 

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

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


To  Ruine  a  World 

John  B.  Patton 

Chairman,  Department  of  Geology,  Indiana  University, 

and 

State  Geologist,  Indiana  Department  of  Natural  Resources 

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

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

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

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

49 


50  Indiana  Academy  of  Science 

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

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

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

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

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

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

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


Address  51 

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

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

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


52  Indiana  Academy  of  Science 

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

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

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

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

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

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


Address  53 

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

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

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

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

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


54  Indiana  Academy  of  Science 

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

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

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

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

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

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


Address  55 

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

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

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

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

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

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


56  Indiana  Academy  of  Science 

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

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

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

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


ANTHROPOLOGY 

Chairman,  Robert  E.  Pace,  Indiana  State  University 

Edward  V.  McMichael, 

Indiana  State  University,  was  elected  Chairman  for  1970 

ABSTRACTS 

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

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

NOTES 

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

57 


58  Indiana  Academy  of  Science 

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

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

One  hundred-fifty  body  sherds  were  excavated  in  1968,  and  one  rim 
sherd  identified  as  Late  Woodland  and  similar  to  Albee  Mound  material. 
There  were  but  15  fragmented  sherds  recovered  in  1969.  The  1969  exca- 
vation of  the  perimeter  of  the  previous  profile  did  not  yield  a  single  post 
hole  between  the  22-inch  to  30-inch  level.  The  1968  excavation  yielded 
approximately  500  post  holes  that  ranged  in  size  from  1  inch  to  91/£ 
inches  in  diameter.  The  post  holes  were  found  between  the  22-  and  24-inch 
level  and  had  a  5-degree  tilt  in  an  easterly  direction.  The  holes  that  were 
recorded  in  1969  were  found  at  the  10-  to  20-inch  level.  They  were  2  to  2V2 
inches  in  diameter  with  no  tilt. 

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

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


Anthropology  59 

OTHER  PAPERS  READ 

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

Elizabeth  J.  Glenn,  Ball  State  University. 

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


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

Georg  K.  Neumann,  Indiana  University 


Abstract 

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

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

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

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

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

60 


Anthropology  61 

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

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


Literature  Cited 

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

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


Origins  and  Racial  Affiliations  of  the  Illinois  Hopewell  Indians 

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

Abstract 

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

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

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

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

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

62 


Anthropology  63 

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

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

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

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

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


64  Indiana  Academy  of  Science 

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

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

Conclusions 

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

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

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

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

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

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

Literature  Cited 

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

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

Springfield.    271  p. 

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

Chicago.    597  p. 

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

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

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

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

Acad,  of  Sci.  60:66-68. 


Some  Possible  Causes  for  Late-Pleistocene  Faunal  Extinctions 

William  H.  Adams,  Indiana  University 

Abstract 

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

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

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

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

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

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

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

65 


66  Indiana  Academy  of  Science 

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

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

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

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

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


Anthropology  67 

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

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

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

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

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


68  Indiana  Academy  of  Science 

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


Conclusions 

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


Literature  Cited 

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

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

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


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

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

Abstract 

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

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

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

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

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

69 


70  Indiana  Academy  of  Science 

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

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

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

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


Anthropology 

Table  1.    Cranial  measurements — means  for  8  males. 


71 


Measurement 


Abbreviation 


Average 


Cranial  Vault 

Cranial  module 

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

Face 

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

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

Orbit 

Left  orbital  height 

Left  orbital  breadth   (mf ) 

Left  orbital  breadth  (d) 

Dental  arch  and  profile 

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

Angles 

Facial  profile  angle 
Midfacial  profile  angle 
Alveolar  profile  angle 
Gonial  angle 

Mandible 

Length  of  mandible 

Bicondylar  breadth 

Height  of  mandibular  symphysis 

Biangular  breadth 

Minimum  ramus  length 


CM 

TP 

L 

B 

MF 

FC 

H 

PAH 

BPH 

LB 


TFH 

UFH 

TFB 

MFB 

IOB 

SIOB 

BOB 

AIM 


NB 

Nil 
DC 
DS 

MN 
SMN 
BNB 
HNB 


LOH 

LOBM 

LOBD 


MB 
ML 
FL 
FLA 


FP< 
MP< 
AP< 

G< 


LM 
BCB 
SH 
HA 

HI, 


140.2 

5.4 

191.5 

136.5 

93.1 
116.4 
141.0 
118.1 

26.1 
107.3 


125.1 
73.9 

137.5 
97.5 
97.6 
20.6 
99.3 
19.4 


25.5 
52.6 
22.2 
13.8 
9.3 
5.0 
59.8 
24.8 


32.8 

4  2.4 


63.6 

55.6 

101.0 

99.0 


84.2° 

88.5° 

69.8° 

124.8° 


108.7 
124.9 

37.9 
107.3 

34.9 


72 


Indiana  Academy  of  Science 

Table  2.    Cranial  indices — means  for  8  males. 


Indices 


Abbreviation 


Average 


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

Face 

Total  facial 

Upper  facial 

Midfacial 

Trans,  cranio-facial 

Zygo-frontal 

Fronto-mandibular 

Zygo-mandibular 

Facial  flatness 

Ant.  interorbital 

Nasal 

Nasal 

Nasal  root  height 
Nasal  bone  height 
Nasal  bridge  height 

Orbit 

Left  orbital   (mf ) 
Left  orbital   (d) 

Dental  Arch 

Maxillo-alveolar 

Mandible 

Mandibular 


B/L 

71.39 

H/L 

73.35 

H/B 

102.76 

H/(L  +  B/2) 

85.48 

PAH/L 

61.79 

BPH/H 

18.56 

MF/B 

68.24 

MF/M'F 

80.75 

TFH/TFB 

91.12 

UFH/TFB 

53.77 

UFH/MFB 

76.96 

TFB/B 

100.82 

MF/TFB 

70.62 

BA/MF 

114.32 

BA/TFB 

77.51 

SIOB/IOB 

21.45 

AIB/BOB 

19.27 

NB/NH 

49.19 

DS/DC 

62.28 

SMN/MN 

55.98 

HNB/BNB 

41.76 

LOH/LOB 

77.43 

LOH/LOBD 

82.31 

MB/ML 


LM/BCB 


114.41 


86.42 


Anthropology 

Table  3.    Morphological  observations — percentages  and  frequencies 


sm* 

m 

1 

vl 

Muscularity 

0    (0) 
It 

0    (0) 
m 

87    (7) 
h 

13    (1) 

Weight 

0    (0) 

sm 

71    (5) 
m 

29    (2) 

1 

Face  size 

0    (0) 

62    (5) 

38    (3) 

med 

div 

con 

tor 

Brow  ridge  shape 

0    (0) 

100    (8) 

0    (0) 

0    (0) 

tr 

sm 

m 

1 

vl 

Brow    ridge    size 

0    (0) 
sm 

12    (1) 
m 

25    (2) 

1 

50    (4) 
vl 

13    (1) 

Glabella 

12    (1) 

38    (3) 

38    (3) 

12    (1) 

vlo 

lo 

m 

hi 

vhi 

Frontal  height 

0    (0) 

13    (1) 

75    (6) 

12    (1) 

0    (0) 

n 

bul 

si 

m 

P 

vp 

Frontal    slope 

0    (0) 
sm 

0    (0) 
m 

13    (1) 

1 

87    (7) 

0    (0) 

0    (0) 

Post-orb.   constr. 

75    (6) 
sm 

25    (2) 
m 

0    (0) 

1 

Frontal  eminences 

75    (6) 
n 

25    (2) 
sm 

0    (0) 
m 

1 

Med.    front,    crest 

75    (6) 

25    (2) 

0    (0) 

0    (0) 

n 

sm 

m 

1 

vl 

Sagittal  elev. 

37    (3) 

sm 

50    (4| 
sm 

37    (1) 

m 

0    (0) 

1 

0    (0) 

Parietal  eminences 

0    (0) 
fl 

100    (8) 
m 

0    (0) 

1 

Temp,  fullness 

25    (2) 

sm 

38    (3) 
m 

37    (3) 

1 

0    (0) 
vl 

Mastoids,  size 

0    (0) 

50    (4) 

25    (2) 

25    (2) 

n 

sm 

m 

P 

vp 

Occip.  curve 

0    (0) 
hi 

0    (0) 
m 

25    (2) 

1 

62    (5) 

13    (1) 

Occip.   position 

87    (7) 

13    (1) 

0    (0) 

bun 

na 

m 

wi 

Occip.  breadth 

13    (1) 

87    (7) 

0    (0) 

0    (0) 

n 

srn 

m 

P 

Lamb,  flattening 

0    (0) 
sm 

13    (1) 
m 

50    (4) 
1 

37    (3) 
vl 

Elev.   occ.   condyles 

0    (0) 

14    (1) 

57    (4) 

29    (2) 

lo 

m 

hi 

vhi 

Basion 

0    (0) 
n 

71    (5) 

s 

15    (1) 
m 

14    (1) 

1 

Platybasia 

100    (8) 
vsm 

0    (0) 
sm 

0    (0) 
m 

0    (0) 

1 

Styloids 

25    (2) 
sm 

50    (4) 
m 

13    (1) 
1 

12    (1) 

Lacerate   foramen 

17    (1) 

sm 

17    (1) 
m 

66    (4) 
1 

Glenoid  fossa 

0    (0) 
sm 

25    (2) 
m 

75    (6) 

1 

Post-glen,  process 

0    (0) 

75    (6) 

25    (2) 

th 

m 

tk 

vtk 

Tympanic  plate 

87    (7) 
s 

13    (1) 
m 

0    (0) 
1 

0    (01 

Petrous  depr. 

0    (0) 

50    (4) 

50    (4) 

obi 

rhm 

sq 

ell 

id 

74 


Indiana  Academy  of  Science 


Table  3.      (Continued) 


sm* 

m 

1 

vl 

Orbit  shape 

38    (3) 

62    (5) 

0    (0) 

0    (0) 

0    (0) 

n 

sm 

m 

P 

vp 

Orbit   inclin. 

25    (2) 

38    (3) 

37    (3) 

0    (0) 

0    (0) 

i) 

si 

m 

dp 

Suborbit.  fossa 

0    (0) 

63    (5) 

37    (3) 

0    (0) 

sm 

m 

1 

vl 

Zygomatic  size 

25    (2) 
sm 

63    (5) 
m 

12    (1) 

1 

0    (0) 

Zygo.   lat.  proj. 

0    (0) 
sm 

50    (4) 
m 

50    (4) 

1 

Zygo.  ant.  proj. 

0    (0) 

87    (7) 

13    (1) 

lo 

m 

hi 

Zygo.  bone  ht. 

0    (0) 

100    (8) 

0    (0) 

sm 

m 

1 

Size  of  nasals 

0    (0) 

100    (8) 

0    (0) 

lo 

m 

hi 

vhi 

Nasal   root  height 

13    (1) 

50    (4) 

37    (3) 

0    (0) 

srn 

m 

1 

vl 

Nasal  root  breadth 

37    (3) 

38    (3) 

25    (2) 

0    (0) 

lo 

m 

hi 

vhi 

Nasal  bridge  height 

17    (1) 
sm 

17    (1) 
m 

66    (4) 

1 

0    (0) 

Nasal   bridge   breadth 

50    (3) 

50    (3) 

0    (0) 

str 

cone 

ccv 

cvx 

Nasal  profile 

0    (0) 

0    (0) 

100    (5) 

0    (0) 

n 

sm 

m 

dp 

Nasion    depr. 

13    (1) 

62    (5) 

25    (2) 

0    (0) 

par 

hyp 

ell 

sU 

III 

Palate  shape 

50    (4) 

0    (0) 

13    (1) 

37    (3) 

0    (0) 

lo 

m 

hi 

vhi 

Palate  height 

0    (0) 

0    (0) 

38    (3) 

62    (5) 

n 

sm 

m 

1 

Inframax.  notch 

33    (2) 

33    (2) 

17    (1) 

17    (1) 

r 

sm 

m 

1 

vl 

Mandib.   size 

0    (0) 

50    (4) 

50    (4) 

0    (0) 

nb 

wb 

int 

med 

Chin  form 

25    (2) 

75    (6) 

0    (0) 

0    (0) 

neg 

neu 

sm 

m 

1 

Chin  proj. 

0    (0) 

87    (7) 

0    (0) 

13    (1) 

0    (0) 

ti 

sm 

m 

1 

Gonial  eversion 

50    (4) 

37    (3) 

13    (1) 

0    (0) 

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


Literature  Cited 


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


Preliminary  Report  on  the  Excavation  of  the 

"Great  Mound"  at  Mounds  State  Park  in 

Madison  County,  Indiana 

Kent  D.  Vickery,  Indiana  University 


Abstract 

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


Introduction 

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

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

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


Mound  Structure 

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


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

75 


76  Indiana  Academy  of  Science 

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

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

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

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

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

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


Anthropology 


77 


Post  Hole  Patterns 

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


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


78  Indiana  Academy  of  Science 

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

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

Features 

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

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


Anthropology  79 


Burials 


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

Artifacts 

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

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

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

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

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

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


80  Indiana  Academy  of  Science 

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

Test  Trench 

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

Conclusions 

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

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

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

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


Anthropology  81 

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

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

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


Literature  Cited 

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

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

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

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


82  Indiana  Academy  of  Science 

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

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


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

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


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

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

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


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

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


BOTANY 

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

ABSTRACTS 

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

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

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

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

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

83 


84  Indiana  Academy  of  Science 

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

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

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

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

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

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


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


Abstract 

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

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

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

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

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

85 


86 


Indiana  Academy  of  Science 


^STO^, 


% 


3         4 


/: 


i%Jllr' 


.* 


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

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

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

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

Figure  10.  Clustered  corms  of  Arrhenatherum. 

Figure  11.    Corm  of  Cinna. 

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

Figure  13.    Section  of  stem  of  Hymenachne  pseudointenupta. 

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


Botany  87 

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

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

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

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

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

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


88  Indiana  Academy  of  Science 

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

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

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

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

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


Botany  89 

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

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

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

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

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


90  Indiana  Academy  of  Science 

Literature  Cited 

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

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

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

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

13  :207-8. 

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

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


CELL  BIOLOGY 

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

ABSTRACTS 

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

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

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

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

91 


92  Indiana  Academy  of  Science 

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

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

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

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

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

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

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


Cell  Biology  93 

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

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

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

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


94  Indiana  Academy  of  Science 

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

NOTE 

Ultrastructural  Features  of  the  Calcifying  Epithelial  Odontogenic  Tumor. 

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

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

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


Cell  Biology  95 

OTHER  PAPER  READ 

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


Comparisons  of  Isolated  Plasma  Membranes  from 
Plant  Stems  and  Rat  Liver1 

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

Abstract 

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

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

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

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

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


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

96 


Cell  Biology  97 

Materials  and  Methods 

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

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

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

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

The  following  homogenization  media  were  employed: 

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

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

(3)  Medium  II  minus  MgCl2  (for  estimation  of  phosphatide  acid 
phosphatase  which  was  inhibited  by  Mg+  + ) ; 

(4)  1  mM  sodium  bicarbonate  (22). 

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

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


98  Indiana  Academy  of  Science 

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

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

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

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

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

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

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

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

Results 

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


"*\ 


W- 


PM 


y 


*. 


w 


I  - 

0 


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

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


100  Indiana  Academy  of  Science 


i 

1  t 


xw     t 


© 


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


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

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


Cell  Biology  loi 


GA 


f 


% 


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


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

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


102 


Indiana  Academy  of  Science 


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


0.3 


0.2  - 


—  0.1 


h- 

v  O 

0.0 

> 

—  CD 

o* 
<£ 

y^ 

1.0 

u_  ._ 

—   CL 

O 

CO      . 

0.8 

-    A.   ONION    STEM 

-   B.   RAT    LIVER 

\    A\      Total 

x'X^x        Total 
/         \       Particulate 

1  J     \     Homogenate 

1  /        V 

\/ 

-          /            x\ 

x                      \ 

/                       \ 

~~xv                 Total    Particulate 

'                                                                             —  y 

'                                 •                          TOtQl 

^ — ^•^^•^Homogenate 

x      x~x                 x 

V   ^x 

s            ^*^ 

1     1     1     1     x     1     1 

Ir^l           1           1           1           1           1      - 

o 

^  0.6 
0.4 


0.2  - 


0,0 

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

CELL  FRACTION 

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

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

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

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

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

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


Cell  Biology  103 

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

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

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

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

Discussion 

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

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

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


104 


Indiana  Academy  of  Science 


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


Constituent 


Specific  Activity  or  Amount 
Rat  Liver  Onion  Stem 


5'-Nucleotidase 

Mg++ -adenosine 

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

triphosphatase 
Phosphatidic  acid 

phosphatase 
Sphingomyelin 

Sialic  acid 
Sterols 
Buoyant  density 


50  /xmoles/hr/mg 

protein 
45  ^moles/hr/mg 

protein 

10  /umoles/hr/mg 

protein 

1  //mole/hr/mg 

protein 

18%  of  total  lipid 

phosphorous 

32  mum/mg  protein 

5-6%  of  dry  weight 

1.16-1.18 


not  detected 

not  detected 

not  detected 

0.1  /xmoles/hr/mg 

protein 

not  detected 
not  detected 

present 

est.  1.4* 


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


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

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

Our  results  support  the  contention  of  Emmelot  and  Benedetti  (2) 
that  the  plasma  membrane  is  a  highly  differentiated  membrane  system 
with  its  characteristics  and  composition  varying  according  to  species, 
cell  type  and  perhaps  even  the  metabolic  state  of  the  cell.  Our  com- 
parisons of  rat  liver  vs.  onion  stem  show  the  plasma  membranes  to  be 
different  in  many  ways  and  that  species  differences  are  reflected  in  the 
characteristics  of  the  surface  membrane. 


Cell  Biology  105 

Literature  Cited 

1.  Benedetti,  E.  L.,  and  P,  Emmelot.  1967.  Studies  on  plasma  membranes.  IV.  The 
ultrastructural  localization  and  content  of  sialic  acid  in  plasma  membranes  isolated 
from  rat  liver  and  hepatoma.  J.  Cell  Sci.  2  :499-512. 

2.  Benedetti,  E.  L.,  and  P.  Emmelot.  1968.  Structure  and  function  of  plasma  membranes 
isolated  from  liver,  p.  33-120.  In  A.  Dalton  and  F.  Haguenau  [eds.  1  The  Membranes. 
Academic  Press,  New  York. 

3.  Cheetham,  R.  D.,  D.  J.  MORBE,  and  W.  N.  Yunghans.  1970.  Isolation  of  a  Golgi 
apparatus-rich  fraction  from  rat  liver.  II.  Enzymatic  characterization  and  com- 
parison with  other  cell  fractions.  J.  Cell  Biol.  44  :492-500. 

4.  Clowes,  F.  A.  L.,  and  B.  E.  Juniper.  1968.  Plant  Cells.  Blackwell  Scientific  Publica- 
tions, Oxford  and  Edinburgh.  546  p. 

5.  Edleman,  J.,  and  M.  A.  Hall.  1965.  Enzyme  formation  in  higher-plant  tissues. 
Development  of  invertase  and  ascorbate-oxidase  activities  in  mature  storage  tissue  of 
Helianthus  tuberosus  L.  Biochem.  J.  95:403-410. 

6.  Eisinger,  W.  R.,  and  D.  J.  MORRE.  1968.  The  effect  of  sulfhydral  inhibitors  on  plant 
cell  elongation.  Proc.  Indiana  Acad.  Sci.  77:136-143. 

7.  Emmelot,  P.,  C.  J.  Bos,  E.  L.  Benedetti  and  Ph.  Rumke.  1964.  Studies  on  plasma 
membranes.  I.  Chemical  composition  and  enzyme  content  of  plasma  membranes 
isolated  from  rat  liver.  Biochim.  Biophys.  Acta  90:126-145. 

8.  Epstein,  E.  1960.  Spaces,  barriers,  and  ion  carriers :  Ion  absorption  by  plants.  Amer. 
J.  Bot.  47  :393-399. 

9.  FlSKE,  C.  H.,  and  Y.  Subbarow.  1925.  The  colorimetric  determination  of  phosphorus. 
J.  Biol.  Chem.  66  :375-400. 

10.  Frey-Wyssling,  A.,  and  K.  Muhlethaler.  1965.  Ultrastructural  Plant  Cytology. 
Elsevier  Press,  Amsterdam,  The  Netherlands. 

11.  Jorgensen,  K.  H.,  and  H.  Dam.  1957.  An  ultra  micromethod  for  the  determination  of 
total  cholesterol  in  bile,  based  on  the  Tschugaeff  color  reaction.   Acta  Chemica  Scand. 

11:1201-1208. 

12.  Keenan,  T.  W.,  and  D.  J.  Morre.  1970.  Phospholipid  class  and  fatty  acid  composi- 
tion of  Golgi  apparatus  isolated  from  rat  liver  and  comparison  with  other  cell 
fractions.  Biochemistry  9  :  19-25. 

13.  Lowry,  O.  H.,  N.  J.  Rosenbrough,  A.  L.  Farr  and  R.  J.  Randall.  1951.  Protein 
measurement    with    the    Folin    phenol    reagent.    J.     Biol.    Chem.    193  :265-275. 

14.  Middleton,  A.  E.,  R.  Cheetham,  D.  Gerber  and  D.  J.  Morre.  1969.  Adenosine 
mono-,  di-  and  trinucleotidase  activities  of  rat  liver  eytornembranes.  Proc.  Indiana 
Acad.  Sci.  78:183-188. 

15.  Morre,  D.  J.  Cell  wall  dissolution  and  enzyme  secretion  during  leaf  abscission. 
Plant  Physiol.  43:1545-1559. 

16.  Morre,  D.  J.,  and  W.  R.  Eisinger.  1968.  Cell  wall  extensibility:  Its  control  by  auxin 
and  relationship  to  cell  elongation,  p.  625-645.  In  F.  Wightman  and  G.  Setterfield  [eds.] 
Biochemistry  and  Physiology  of  Plant  Growth  Substances.  Runge  Press,  Ottawa, 
Canada. 

17.  Morre,  D.  J.,  T.  W.  Keenan  and  H.  H.  Mollenhauer.  In  press.  Golgi  apparatus 
function  in  membrane  transformations  and  product  compartmentalization  :  Studies 
with  cell  fractions  from  rat  liver.  Proc.  1st  Intern.  Sym.  Cell  Biol.  Cytopharm., 
Venice,  Italy,  1969. 

18.  MORRE,  D.  J.,  and  J.  L.  Key.  1967.  Auxins;  pp.  575-593.  In  F.  H.  Wilt  and  N.  K. 
Wessels   [eds.]  Methods  in  Developmental  Biology.  Thomas  Y.  Crowell  Co.,  New  York. 

19.  Morre,  D.  J.,  R.  L.  Hamilton,  H.  H.  Mollenhauer,  R.  W.  Mahley,  W.  P.  Cun- 
ningham, and  V.  S.  Lequire.  1970.  Isolation  of  a  Golgi  apparatus-rich  fraction 
from  rat  liver.  I.  Method  and  morphology.  J.  Cell  Biol.  44  :484-491. 

20.  Morre,  D.  J.,  and  H.  H.  Mollenhauer.  1964.  Isolation  of  the  Golgi  apparatus  from 
plant  cells.  J.  Cell  Biol.  23  :295-305. 


106  Indiana  Academy  of  Science 

21.  Morrk,  D.  J.,  H.  H.  Mollenhauer  and  J.  E.  Chambers.  1965.  Glutaraldehyde  stabili- 
zation   as    an    aid   to    Golgi    apparatus    isolation.    Exp.    Cell    Res.    38 :672-675. 

22.  Neville,  D.  M.  I960.  The  isolation  of  a  cell  membrane  fraction  from  rat  liver.  J. 
Biophys.  Biochem.  Cytol.  8:413-422. 

23.  Ordin,  L.,  and  M.  A.  Hall.  1967.  Studies  on  cellulose  synthesis  by  a  cell-free  oat 
coleoptile  enzyme  system  :  Inactivation  by  airborne  oxidants.   Plant  Physiol.  42  :205-212. 

24.  Parsons,  J.  G.,  and  S.  Patton.  1967.  Two  dimensional  thin-layer  chromatography  of 
polar  lipids  from  milk  and  mammary  tissue.  J.  Lipid  Res.  8  :696-699. 

25.  Reynolds,  E.  S.  1963.  The  use  of  lead  citrate  at  high  pH  as  an  electron-opaque  stain 
in  electron  microscopy.  J.  Cell  Biol.  17:208-212. 

26.  Roland,  J.  C.  1969.  Mise  en  evidence  sur  coupes  ultrafines  de  formations  poly- 
saecharidiques  directement  associees  au  plasmalemme.  C.  R.  Acad.  Sc.  Paris  269D  :939- 
942. 

27.  Rohser,  G.,  A.  N.  Siakotos  and  S.  Fleischer.  1966.  Quantitative  analysis  of  phospho- 
lipids by  thin-layer  chromatography  and  phosphorous  analysis  of  spots.   Lipids  1  :85-86. 

28.  Smith,  S.  W.,  S.  B.  Weiss  and  E.  P.  Kennedy.  1957.  The  enzymatic  dephosphoryla- 
tion   of   phosphatidic   acids.    J.    Biol.    Chem.    228 :915-922. 

29.  Poux,  N.  1967.  Localisation  d'activites  enzymatiques  dans  les  cellules  du  meristeme 
radiculaire  de  Cuctimis  sativus  L.  I.  Activites  phosphatasiques  neutres  dans  les  cellules 
du  protoderme.  J.  Microscopie  6:1043-1058. 

30.  Straus,  J.  1962.  Invertase  in  cell  walls  of  plant  tissue  cultures.  Plant  Physiol. 
37:342-348. 

31.  Villemez,  C.  L.,  J.  M.  McNab  and  P.  Albersheim.  1968.  Formation  of  plant  cell 
wall  polysaccharides.  Nature  218:878-880. 

32.  Widnell,  C.  C,  and  J.  C.  Unkeless.  1968.  Partial  purification  of  a  lipo-protein  with 
5'nucleotidase  activity  from  membranes  of  rat  liver  cells.  Proc.  Nat.  Acad.  Sci. 
61:1050-1057. 


Di-  and  Tri-nucleotidase  Activities  of  Rat  Liver  Cytomembranes1 

R.  D.  Cheetham  and  D.  J.  Morre,  Purdue  University 

Abstract 

Enzymatically  active  fractions  of  endoplasmic  reticulum,  Golgi  apparatus  and  plasma 
membrane  were  isolated  from  rat  liver.  Without  exception,  the  di-  and  trinucleotide  phos- 
phatase activities  of  the  Golgi  apparatus  fraction  were  intermediate  between  those  of  the 
endoplasmic  reticulum  and  plasma  membrane.  These  data  support  the  proposal  that  the 
Golgi  apparatus  serves  as  a  site  of  endomembrane  differentiation  from  endoplasmic 
reticulum-like  to  plasma  membrane-like. 

Morphological  evidence  has  provided  the  basis  for  a  proposal  that 
Golgi  apparatus  function  as  sites  of  cytomembrane  differentiation  in 
the  formation  of  membranes  which  are  plasma  membrane-like,  beginning 
with  an  input  of  membrane  constituents  from  endoplasmic  reti2ulum 
and  implying  a  progressive  change  in  the  composition  or  arrangement 
of  constituents  within  the  membrane  (5,  10).  An  understanding  of  the 
functional  significance  of  the  transitional  nature  of  the  Golgi  apparatus 
requires  information  on  the  chemical  and  enzymatic  composition  of 
Golgi  apparatus  relative  to  membranes  of  the  endoplasmic  reticulum 
and  plasma  membrane. 

In  this  report,  we  compare  nucleoside  di-  and  triphosphatase  ac- 
tivities of  endoplasmic  reticulum,  Golgi  apparatus  and  plasma  mem- 
brane. With  every  nucleotide  tested,  the  activity  of  the  Golgi  apparatus 
is  intermediate  between  that  of  the  endoplasmic  reticulum  and  plasma 
membrane. 

Materials  and  Methods 

Methods  for  isolation  (2,  9,  10),  yield  and  purity  (8)  of  the  endo- 
plasmic reticulum,  Golgi  apparatus  and  plasma  membrane  fractions  have 
been  described.  Enzyme  assays  were  at  37°  C  under  conditions  where 
activity  was  proportional  to  time  of  incubation  and  protein  concentration. 
Protein  was  determined  by  the  method  of  Lowry  et  al.  (7)  and  inorganic- 
phosphate  by  the  method  of  Fiske  and  Subbarow  (4).  All  assays  were 
with  nucleotides  as  sodium  salts  in  medium  A-l  of  Emmelot  et  al.  (3), 
pH  7.4. 

Substrates  were  of  the  highest  purity  obtainable  from  the  suppliers 
indicated:  inosine-5'-diphosphate  (IDP);  thy  midine-5'-diphosphate 
(TDP);  thymidine-5'-triphosphate  (TTP)  and  guanosine-5'-diphosphate 
(GDP)  [Calbiochem];  adenosine-5'-diphosphate  (ADP),  cytidine-5'-di- 
phcsphate  (CDP),  uridine-5'-diphosphate  (UDP),  inosine-5'-triphosphate 
(ITP),  uridine-5'-triphosphate  (UTP)  [Sigma];  adenosine-5'-triphosphate 
(ATP),  guanosine-5'-triphosphate  (GTP)   [Nutritional  Biochemicals]. 


1  Purdue  University  AES  Journal   Paper  No.    3910.    Work  supported   in   part  by  grants 
from  the  NSF  GB  1084  and  707S. 

107 


108 


Indiana  Academy  of  Science 


Results  and  Discussion 

The  relative  nucleoside  di-  and  triphosphatase  activities  of  the 
isolated  fractions  are  shown  in  Figure  1.  The  specific  activity  (/xmoles 
Pi/hr/mg  protein)  of  each  fraction  was  divided  by  the  specific  activity 
of  the  corresponding  total  homogenate.  This  corrects  for  fluctuations  in 
feeding  and  age  of  the  animals  and  for  variations  in  the  isolation  pro- 
cedures. The  activity  of  the  Golgi  apparatus-rich  fraction  is  intermediate 
between  that  of  endoplasmic  reticulum-  and  plasma  membrane-rich  frac- 
tions even  though  the  activity  is  increasing,  as  with  uridine  triphos- 
phatase; decreasing,  as  with  guanosine  diphosphatase;  or  unchanging,  as 
with  inosine  diphosphatase;  in  going  from  endoplasmic  reticulum  to 
plasma  membrane. 

Morphological  evidence  for  a  transitional  nature  for  the  Golgi  ap- 
paratus has  come  from  studies  with  the  fungus  Pythium   ultimum    (5) 


12 


UJ 

|s 

LU 
CD 

o 
o 

X 

_l 

<    4 
o 


go 


I 


1 


m 


1 


I 


^ 


i 

I 


H_i 


II 


m  m 


I 


5 


I 


s 


o 


4- 


ERGAPM 
ATPase 


ER  GAPM 

UTPase 


ER  GA  PM 

GTPase 


ER  GA  PM 

ITPase 


ER  GAPM 

CTPase 


i 


IZZl 


m 


m 


m 


m. 


11 


L. 


I 


ERGAPM 

TTPase 


ERGAPM 
ADPase 


ER  GA  PM 
UDPase 


ER  GA  PM 

GDPase 


ER  GA  PM 

IDPase 


ER  GA  PM 

CDPase 


Figure  1.  Relative  specific  activities  of  hydrolysis  of  nucleoside  di-  and  tri- 
at  pH  7.-4  by  endoplasmic  reticulum  (ER)-,  Golgi  apparatus  (GA)-  and  plasma 
(PM)-rich  cell  fractions  from  rat  liver.  Relative  specific  activity  is  the  ratio 
activity  (umo/es  iP/hr/my  protein)  of  each  fraction  to  that  of  the  total  h 
(Abbreviations  for  substrates  are  explained  in  the  text.) 


ER  GA  PM 

TDPase 

phosphates 

membrane 
of  specific 
omoociiatc. 


Cell  Biology  109 

and  rat  liver  (10).  These  studies  show  a  progressive  change  in  both 
membrane  staining  and  membrane  thickness  across  stacked  cisternae 
from  a  morphology  which  resembles  that  of  the  endoplasmic  reticulum 
to  a  morphology  which  is  plasma  membrane-like.  The  transitional  nature 
of  the  Golgi  apparatus  is  also  reflected  in  the  lipid  composition  of  these 
fractions  where  the  Golgi  apparatus  is  again  intermediate  between  the 
endoplasmic  reticulum  and  the  plasma  membrane  (10).  The  enzyme  ac- 
tivities presented  here  lend  further  support  to  our  proposal  that  the 
Golgi  apparatus  functions  in  the  transformation  of  membranes  from 
endoplasmic  reticulum-like  to  plasma  membrane-like. 

Golgi  apparatus  are  not  known  to  be  sites  of  protein  synthesis  (6) 
but  enzyme  proteins  might  be  added  to  or  removed  from  the  membranes 
during  transformation.  If  transfer  of  proteins  from  endoplasmic  reticu- 
lum to  Golgi  apparatus  occurs  in  the  formation  of  plasma  membrane- 
like  secretory  vesicles,  then  it  appears  that  the  process  must  involve  a 
selective  transfer  so  that  certain  proteins  become  concentrated  in  the 
secretory  vesicle  membranes  whereas  others  are  not  incorporated  (1). 
As  an  alternative,  certain  activities  of  enzyme  proteins  derived  from 
endoplasmic  reticulum  might  be  progressively  activated  or  inhibited  con- 
current with  the  changes  in  lipid  and  carbohydrate  composition  of  the 
membranes  (1).  Hopefully,  pulse  labeling  studies  involving  one  or  more 
of  the  protein  constituents  of  plasma  membrane  can  be  used  to  provide 
a  direct  test  of  the  concept  of  membrane  flow  in  rat  liver. 

Literature  Cited 

1.  Cheetham,  R.  D.,  T.  W.  Keenan,  S.  Nyquist,  and  D.  J.  Morre.  1969.  Biochemical 
comparisons  of  endoplasmic  reticulum-,  Golgi  apparatus-,  and  plasma  membrane-rich 
cell  fractions  from  rat  liver  in  relation  to  cytomembrane  differentiation.  J.  Cell  Biol. 
43:21a. 

2.  Cheetham,  R.  D.,  D.  J.  Morre,  and  W.  N.  Yunghans.  1970.  Isolation  of  a  Golgi 
apparatus-rich  fraction  from  rat  liver.  II.  Enzymatic  characterization  and  comparison 
with  other  cell  fractions.  J.  Cell  Biol.  44  :492-500. 

3.  Emmelot,  P.,  C.  J.  Bos,  E.  L.  Benedetti,  and  P.  H.  Rumke.  1964.  Studies  on  plasma 
membranes.  I.  Chemical  composition  and  enzyme  content  of  plasma  membranes  isolated 
from  rat  liver.  Biochim.  Biophys.  Acta  90:126-145. 

4.  Fiske,  C.  H.,  and  Y.  Subbarow.  1925.  The  colorimetric  determination  of  phosphorus. 
J.  Biol.  Chem.  66:375-400. 

5.  Grove,  S.  N.,  C.  E.  Bracker,  and  D.  J.  Morre.  1968.  Cytomembrane  differentiation 
in   the  endoplasmic   reticulum-Golgi   apparatus-vesicle   complex.    Science   161:171-173. 

6.  LeBlond,  C.  P.  1965.  General  conclusions,  p.  321.  In:  C.  P.  LeBlond  and  K.  B. 
Warren  [eds.  ]  The  Use  of  Radioautography  in  Investigating  Protein  Synthesis. 
Academic  Press,  New  York. 

7.  Lowry,  O.  H.,  N.  J.  Rosenbrough,  A.  L.  Farr,  and  R.  J.  Randall.  1951.  Protein 
measurement   with    the    Folin    phenol    reagent.    J.    Biol.    Chem.    193:265-275. 

8.  Middleton,  A.  E.,  R.  D.  Cheetham,  D.  Gerber,  and  D.  J.  Morre.  1969.  Adenosine 
mono-,  di-  and  trinucleotidase  activities  of  rat  liver  cytomembranes.  Proc.  Indiana 
Acad.  Sci.  78:183-188. 

9.  Morre,  D.  J.,  R.  L.  Hamilton,  H.  H.  Mollenhauer,  R.  W.  Mahley,  W.  P.  Cun- 
ningham, R.  D.  Cheetham,  and  V.  S.  Lequire.  1970.  A  Golgi  apparatus-rich  cell 
fraction    isolated  from   rat   liver.    I.    Method  and  morphology.    J.    Cell    Biol.    44:484-491. 

10.  Morre,  D.  J.,  T.  W.  Keenan,  and  H.  H.  Mollenhauer.  In  press.  Golgi  apparatus 
function  in  membrane  transformations  and  product  compartmentalization :  Studies 
with  cell  fractions  isolated  from  rat  liver.  Proc.  1st  Intern.  Symp.  Cell  Biol,  and 
Cytopharmacol.,  Venice,  Italy,  7-11  July  1969. 


A  Model  Mosaic  Membrane:  Cytochrome  Oxidase1 

T.  F.  Chuang,  Y.  C.  Awasthi  and  F.  L.  Crane, 
Purdue  University 


Abstract 

During  purification  of  a  lipid-free  cytochrome  oxidase,  beef  heart  mitochondrial  mem- 
branous structure  was  broken  down  through  use  of  the  nonionic  detergent  Triton, 
together  with  KC1.  After  removal  of  lipid,  the  purified  cytochrome  oxidase  appears  as 
90A  diameter  globules  or  as  assemblies  of  rod-like  structure  with  the  same  thickness. 
Upon  addition  of  mitochondrial  phospholipid,  structural  transformation  of  the  enzyme 
occurs  and  enzyme  activity  is  enhanced.  Three  phases  of  transformation  in  structure 
depending  on  the  amount  of  phospholipid  added  are  observed:  1)  at  0-20  g  Atom  P  of 
phospholipid/mole  cytochrome  oxidase  there  is  transformation  from  swollen  particles  or 
rod-like  elements  (200A)  to  sheets  made  up  of  50A  globules;  2)  at  20-65  g  Atom  P/mole, 
the  50A  globules  are  evenly  dispersed  to  form  mosaic  membrane  vesicles;  3)  at  higher  than 
65  g  Atom  P/mole,  there  is  excess  phospholipid  around  the  membrane  vesicle,  and  the 
subunits  appear  to  be  30-50A  in  diameter.  During  this  transformation  there  is  a  regular 
increase  in  activity  which  attains  maximum  at  65  g  Atom  P/mole  in  oxidase  and  this 
activity  remains  unchanged  even  at  higher  phospholipid  concentration  up  to  235  g  Atom 
P/mole.  We  conclude  that  the  oxidase  protein  globules  form  a  mosaic  membrane  with 
phospholipid  interspersed  between  the  globules. 

Introduction 

Purified  cytochrome  oxidase  has  been  shown  to  be  able  to  form 
membranes.  Deoxycholate  will  disperse  the  membrane  structure  and  after 
removal  of  deoxycholate  the  solubilized  cytochrome  oxidase  is  organized 
to  form  membrane  vesicles  (8).  In  the  study  of  the  relationship  between 
membrane  formation  and  ionic  strength,  we  observed  that  the  membrane 
formation  could  be  accomplished  in  the  presence  of  high  concentration 
of  nonionic  detergent  (10).  By  employing  nonionic  detergents,  Tritons, 
together  with  KC1,  a  lipid-free  cytochrome  oxidase  has  been  prepared 
(2,  10)  and  it  appears  as  90 A  globules.  In  the  investigation  of  phospho- 
lipid function  in  the  cytochrome  oxidase  reaction  system  (2)  we  found 
that  this  lipid-free  preparation  is  rather  inert  to  its  substrate,  cytochrome 
c.  Maximal  activity  can  not  be  obtained  using  even  lipid-cytochrome  c 
(5)  as  its  substrate  if  phospholipid  is  absent  on  the  enzyme  side  (2). 
Enzyme  activity  toward  either  cytochrome  c  or  lipid-cytochrome  c  in- 
creases on  addition  of  phospholipid  micelles  to  the  enzyme.  In  the  pres- 
ent study  we  discuss  the  relationship  between  the  enzyme  activity  and 
amount  of  phospholipid  present  as  well  as  the  structural  transforma- 
tion of  the  enzyme  on  addition  of  mitochondrial  phospholipids. 

Materials  and  Methods 

Mitochondria  were  isolated  by  the  method  of  Low  and  Vallin  (7)  and 
were  stored  in  concentrated  suspension  at  — 20°  C  before  use.  Enzyme 
and  phospholipid  were  prepared  from  mitochondria  which  were  less  than 


1  Supported  under  grant  AM04663  from  the  National  Institutes  of  Arthritis  and 
Metabolic  Diseases.  F.  L.  Crane  is  supported  by  Career  Grant  K6-21.839  from  the  National 
Institutes  of  General  Medical  Science. 

110 


Cell  Biology  111 

two  weeks  old.  Lipid-free  cytochrome  oxidase  was  prepared  as  described 
by  Sun  et  al.  (10  with  some  modification  (2).  One  mM  succinate  was 
added  before  sonication  and  0.02  m  Tris  pH  7.4  was  used  instead  of 
phosphate  buffer.  This  enzyme  contained  only  2-3  g  Atom  P/mole  cyto- 
chrome oxidase  or  2-3%  phospholipid  (w/w)  and  it  has  heme  a  concen- 
tration of  8-9  rn.fi  moles /mg  of  protein  as  determined  by  its  differential 
spectrum  using  a  millimolar  extinction  coefficient  AA  at  605-630  m/x  at 
13.1  (12). 

Phospholipid  micelles  and  the  lipid-free  cytochrome  oxidase  were 
soluble  and  could  not  be  centrifuged  out  at  108,000  x  g  for  1  hr.  To 
study  the  amount  of  phospholipid  bound  to  cytochrome  oxidase  during 
membrane  formation,  different  amounts  of  phospholipid  were  added  to 
40  mg  cf  lipid-free  cytochrome  oxidase  and  the  mixture  was  diluted 
with  0.25  m  sucrose,  0.02  M  Tris,  pH  7.4  to  3  mg  of  protein  per  ml.  The 
mixture  was  sonicated  at  maximal  output  with  a  Branson's  Sonifer  for 
five  30  sec  intervals  and  centrifuged  at  108,000  x  g  for  30  min.  The  pellet 
was  resuspended  in  sucrose-Tris  buffer  and  centrifuged  at  the  same 
speed.  After  three  washes  all  free  lipid  and  free  cytochrome  oxidase 
which  were  not  in  the  membrane  were  washed  out. 

Protein  was  determined  according  to  Yonetani  (13).  Molecular 
weight  of  cytochrome  oxidase  was  based  on  the  value  of  72,000  (3). 
Phospholipid  micelles  were  prepared  by  the  sonication  method  of  Flei- 
scher and  Fleischer  (6).  Cytochrome  oxidase  activities  were  assayed 
polarographically  in  the  following  mixture  at  pH  6.5:  16  mM  potassium 
phosphate;  10  mM  potassium  citrate;  0.80  mM  EDTA;  13  mM  potassium 
ascorbate;  1.11  mM  TMPD;  15  ^M  cytochrome  c  in  a  total  volume  of  1.8 
ml  using  a  Gilson  oxygraph  at  37°  and  the  enzyme  in  the  range  of 
5-20  Mg. 

Tritons  and  cytochrome  c  Type  III  were  purchased  from  Sigma 
Chemical  Company.  Other  chemicals  were  reagent  grade. 

Samples  negatively  stained  with  phosphotungstate,  pH  6.8  were 
prepared  for  electron  microscopy  according  to  the  procedure  of  Cunning- 
ham and  Crane   (4)   and  observed  with  a  Philips  EM  300. 

Results 

Membranes  of  mitochondrial  electron  transport  particles  (ETP) 
(Fig.  1)  were  split  into  two  membranous  fractions  by  using  Triton  X  114 
and  KC1  as  described  by  Prezbindowski  et  al.  (9).  Further  purification 
of  the  green  fraction  results  in  a  lipid-free  cytochrome  oxidase  which 
appears  as  90a  diameter  globules  or  assemblies  of  rod-like  structure 
with  the  same  thickness  (Fig.  2).  This  lipid-free  enzyme  has  low  activity 
which  can  be  recovered  upon  addition  of  phospholipid.  Figure  3  shows 
the  relationship  between  the  enzyme  activity  and  the  amount  of  phos- 
pholipid added.  Three  phases  of  the  enzyme  activity  and  enzyme  struc- 
ture related  to  the  amount  of  phospholipid  added  can  be  observed.  At 
low  amount  of  phospholipid  (0-20  g  Atom  P/mole  cytochrome  oxidase), 
the  90a   globules  and  rod-like   elements  began   to   swell   to  the   size   of 


112 


Indiana  Academy  of  Science 


'■,: 


MM^Sm.  mam 

photung state.    X    103,700.    Marker 


WmmmBBMW 

FIGURE     1.      Mitochondrial     membrane    fragments.     Phi 
1000.1. 

Figure  2.  Purified  lipid-frec  cytochrome  oxidase  showing  90  A  globules  and  rod-like  struc- 
ture made  of  90.1  subunits.  Phosphotung state.  X  103,700.  Marker  1000.1.  Arrow  A  indicates 
90 A  enzyme  globules;  arrow  B,  a  rod-like  element  with  90 A  subunits. 


Cell  Biology 


113 


200a  in  diameter  and  gradually  to  transform  to  a  small  sheet  of  mem- 
brane made  up  to  50a  globules  (Fig.  4).  Addition  of  phospholipid 
rapidly  changed  the  activity  of  the  enzyme  (Fig.  3).  With  addition  of 
more  phospholipid  to  the  enzyme,  (20-65  g  Atom  P/mole  cytochrome 
oxidase),  the  lipoprotein  globules  became  organized  to  form  a  larger 
sheet  of  membrane  with  the  same  size  (50a)  globules  visible  in  surface 
views  (Fig.  5). 


0  50  100  150  200  250 

PHOSPHOLIPID    ADDED    (g    ATOM   P/MOLE    CYTOCHROME    OXIDASE) 

Figure  3.  Enzyme  activity  of  a  purified  lipid-free  cytochrome  oxidase  preparation  and 
the  amount  of  phospholipid  added.  Phospholipid  was  added  and  incubated  at  cold  for  5 
minutes  before  dilution  for  assay.  This  enzyme  contained  1.5  mg  of  Triton  x  100  per  mg 
of  protein. 


At  phospholipid  concentrations  higher  than  65  g  Atom  P/mole  cyto- 
chrome oxidase,  excess  of  phospholipid  was  observed  around  the  mem- 
brane vesicles  and  the  summit  was  reduced  to  about  30-50A  diameter 
(Fig.  6).  Activity  at  this  stage  remains  at  the  maximum  and  addition  of 
more  phospholipid  (up  to  235  g  Atom  P/mole  cytochrome  oxidase)  did 
not  increase  activity. 

When  the  lipid-free  enzyme  was  sonicated  with  increasing  amounts 
of  phospholipid  micelles  (up  to  235  g  Atom  P/mole  cytochrome  oxidase) 
membrane  obtained  by  centrifugation  followed  by  several  washes  to  re- 
move unbound  phospholipid  and  cytochrome  oxidase  showed  regular  in- 
crease in  its  phospholipid  content  up  to  a  constant  value  of  65  g  Atom 
P/mole  cytochrome  oxidase  which  corresponds  to  about  40%  of  lipid 
in  the  membrane  (Fig.  8).  Any  excess  phospholipid  was  washed  out  and 
in  no  experiment  did  the  value  of  bound  phospholipid  in  the  washed  mem- 
brane increase  beyond  this  constant  value.  No  phospholipid  micelle  struc- 


114 


Indiana  Academy  of  Science 


Figure  4.  Cytochrome  oxidase  with  10  g  Atom  P  of  phospholipid  per  mole  of  enzyme 
(or  10%  phospholipid,  w/w)  showing  swollen  particles,  rod-like  elements  and  small  mem- 
brane with  50  A  subunits.  Phosphotung state.  X  103,700.  Marker  1000. 1.  Arrow  points  on 
a  swollen  rod-like  element  with  50 J  subunits. 

Figure  5.  Cytochrome  oxidase  with  .11  g  Atom  P  of  phospholipid  per  mole  of  enzyme 
(or  25%  phospholipid,  w/w)  showing  vesicle  membrane  with  50A  subunits  phosphotung- 
slate.  X  103,700.  Marker  1000A. 


Cell  Biclogy 


115 


Pil 


■-i*.,? 


Figure  6.  Cytochrome  oxidase  with  235  g  Atom  P  of  phospholipid  per  mole  of  enzyme 
(70%  phospholipid,  w/w)  showing  excess  of  phospholipid  micelles  around  membrane 
vesicles.  Phosphotwng state.  X  103,700.  Marker  1000.1. 

Figure  7.  Membraneous  cytochrome  oxidase  after  washing  away  the  free  phospholipid 
micelles.  Excess  of  phospholipid  (235  g  Atom  P  of  phospholipid  per  mole  of  cytochrome 
oxidase)  was  added  to  lipid-free  cytochrome  oxidase  as  described  in  the  text  and  centri- 
fuged  at  108,000  x  g  for  30  minutes  and  3  washes  to  remove  unbound  phospholipid.  Phos- 
pholipid content  decreased  to  65  g  Atom  P  of  phospholipid  }><  r  mole  of  cytochrome  oxi- 
dase. No  micelle  is  seen  around  membrane  vesicles.  Note  50.1  subunits.  Phosphotung state. 
X  103,700.  Marker  1000A. 


116 


Indiana  Academy  of  Science 


uj 

>-    ^ 

N     uJ 
2     CO 

UJ      < 

o 

2  x 
H   o 

UJ 

o 

X 

o 

o 

h- 
>- 
o 


I < 

O      cp 


80 


60 


40 


20 


0  50  100  150  200  250 

PHOSPHOLIPID   ADDED  (g.  ATOM  P/M0LE   CYTOCHROME  OXIDASE) 

Figure  8.     Amount  of  phospholipid  bound  to  cytochrome  oxidase. 


ture  was  observed  around  the  edge  of  the  membrane  vesicles  after  wash- 
ing (Fig.  7).  Subunits  of  the  membrane  consistently  showed  a  diameter 
of  50a  and  completely  filled  the  sheet  surfaces. 

Enzyme  with  phospholipid  added  (vesicle  membraneous  form)  was 
found  to  be  much  more  stable  to  heat  treatment  than  the  lipid-free  en- 
zyme. Figure  9  shows  the  activity  change  of  both  lipid-free  and  lipid- 
added  enzymes  after  incubation  at  different  temperatures.  After  incuba- 
tion at  the  temperature  and  time  indicated,  lipid-free  enzyme  was  incu- 
bated with  phospholipid  at  4°  C  for  10  minutes  before  enzyme  assay. 
Lipid  added  cytochrome  oxidase  has  90%,  81%  and  19%  of  the  original 
activity  after  20  minutes  incubation  at  30°  C,  38°  C  and  50°  C  respec- 
tively while  the  lipid-free  enzyme  had  only  75%,  40%  and  0%  of  original 
activity.  The  cytochrome  oxidase  of  washed  beef  heart  mitochondria  was 
also  found  to  be  more  stable  to  heat  treatment  (Fig.  10). 


Discussion 

The  capability  of  cytochrome  oxidase  to  form  membrane  was  demon- 
strated earlier  by  McConnell  et  al.  (8)  and  our  laboratory  (10).  Mem- 
brane formation  demonstrated  in  the  present  study  requires  the  presence 
of  both  protein  and  phospholipid.  Phospholipid  content  is  also  a  critical 
factor  for  enzyme  activity.  Deficiency  in  lipid  content  causes  the  enzyme 
molecules  to  cluster  together  and  may  prevent  access  of  substrate  to  the 
active  site.  As  can  be  seen  in  Figures  2  and  4,  on  addition  of  phospholipid 


Cell  Biology 


117 


100 


TIME  OF  INCUBATION   (MIN) 

Figure  9.  Stability  of  Upid-free  and  lipid-added  cytochrome  oxidase  activity.  Enzyme 
was  incubated  at  temperature  indicated  at  protein  concentration  of  1  mg/ml.  120  g  Atom 
P  of  phospholipid/mole  enzyme  was  added  before  or  after  incubation.  Solid  lines  are  the 
lipid-added  cytochrome  oxidase  activity;  dotted  lines  the  Upid-free  cytochrome  oxidase 
activity.  Control  activity  as  100%  was  it  1.2  ninoles  Q-i  uptake /min/mg. 


micelles  the  enzyme  swells  and  50a  lipoprotein  globules  are  loosely 
packed  into  a  sheet  to  form  membrane  vesicles.  In  this  sheet  form  the 
substrate  has  ready  access  to  active  sites  on  the  enzyme  as  shown  by 
the  high  cytochrome  oxidase  activity. 

Further  addition  of  phospholipid  micelles  (20-65  g  Atom  P/mole 
enzyme)  allows  a  more  even  dispersion  of  the  enzyme  in  the  form  of 
mosaic  membrane  vesicles  with  50a  globules  evenly  dispersed  as  shown 
in  Figure  5.  Thin  sectioning  of  this  membrane  shows  unit  membrane 
structure  which  is  50-55A  thick.  Once  the  phospholipid  requirement  for 
the  membrane  formation  and  maximal  activity  is  reached,  further  addi- 
tion of  phospholipid  does  not  further  increase  the  activity.  The  negative 
staining  data  show  excess  phospholipid  micelles  around  the  edge  of 
membrane  vesicles.  The  globule  size  on  the  membrane  is  somewhat  re- 
duced to  30a.  This  reduction  of  apparent  globule  size  may  indicate  an 
excess  of  phospholipid  among  the  lipoprotein  globules. 

Sonification  facilitates  the  interaction  of  cytochrome  oxidase  pro- 
tein and  phospholipid  to  form  tightly  bound  membrane.  The  unbound 
lipid  is  still  in  the  state  of  micelles  and  can  be  easily  removed  by  wash- 


Hi 


Indiana  Academy  of  Science 


38° 

inn  * 

n 

IUU 

o 

o 

^ 

A__^ 

V 

-^^^45° 

h-    80 

—  y 

> 

h- 

o 

< 

-1   ,-~ 

O    60 

— 

(T 

h- 

Z 

o 

o 

fe    40 

- 

____^       50° 

h- 

z 

LJ 

u 

cr 

IH    20 

— 

Q_ 

n 

1                      1 

i 

1 

1                       1 

10  20  30  40 

TIME   OF  INCUBATION    (MIN) 


50 


60 


Figure  10.  Stability  of  cytochrome  oxidase  in  a  beef  heart  mitochondrial  preparation. 
Mitochondria  were  adjusted  to  protein  concentration  of  U  mg/ml  and  incubated  at  indi- 
cated temperature.  Control  activity  as  100%  was  5:20   ^motcs  02  uptake  /  min  j  mg  protein. 


ing.  A  consistent  value  of  60-70  g  Atom  phosphorus  of  phospholipid  per 
mole  of  cytochrome  oxidase  was  obtained  after  washing  when  an  excess 
of  phospholipid  was  added  (Fig.  8).  The  same  saturated  value  of  phos- 
pholipid was  obtained  for  the  maximal  activity  of  cytochrome  oxidase 
(Fig.  3).  The  electron  micrograph  (Fig.  7^  showed  50a  globules  on  the 
washed  membrane  surface  and  no  phospholipid  micelles  were   observed. 

The  90a  particles  characteristic  of  the  lipid-free  enzyme  are  equiva- 
lent to  a  molecular  weight  of  290,000  daltons  if  the  protein  density  is 
assumed  as  1.25.  Molecular  weight  of  monomer  cytochrome  oxidase  is 
72,000  (3).  Thus,  our  lipid-free  cytochrome  oxidase  may  exist  as  a 
tetramer.  Ball  et  al.  (1)  suggested  that  cytochrome  oxidase  might  be  a 
tetrapolymer  consisting  of  four  identical  hemoprotein  submolecules. 
Takemori  et  al.  (11)  also  suggested  that  their  Emasoln.to  solubilized 
cytochrome  a  as  a  pentamer  based  on  sedimentation  experiments.  After 
cytochrome  oxidase  is  reorganized  into  membrane  the  globule  size  ap- 
pears to  be  50-60A  which  is  equivalent  to  the  molecular  weight  of 
50,000-85,000.  This  value  agrees  with  the  molecular  weight  of  72,000 
suggested  by  Criddle  and  Bock  (3).  Takemori  et  al.  (11)  suggested  that 
cytochrome    oxidase   existed   as   a   monomer   with    only   one   heme   group 


Cell  Biology 


119 


per  molecule  in  mitochondrial  particles,  it  tended  to  polymerize  after 
extraction  from  mitochondria  with  bile  salt  and  purification  with  am- 
monium sulfate.  Similar  polymerization  probably  also  occurs  in  our 
preparation  and  depolymerization  occurs  again  during  reconstitution. 


50  A  ~j~~ 


Q  PROTEIN    GLOBULE 
T    PHOSPHOLIPID 


Figure  11.     A  model  mosaic  membrane:  cytochrome  oxidase. 


In  a  study  of  the  heat  stability  of  the  enzyme  we  found  that  the 
membraneous  form  of  cytochrome  oxidase  (made  by  addition  of  phos- 
pholipid to  lipid-free  cytochrome  oxidase)  was  more  stable  than  the 
lipid-free  enzyme.  Figure  9  shows  the  cytochrome  oxidase  activity  after 
incubation  at  a  series  of  temperatures  for  enzyme  to  which  phospholipid 
was  added  before  or  after  incubation.  The  membraneous  cytochrome 
oxidase  has  90%,  81%  and  19%  of  the  control  activity  while  the  incu- 
bated lipid-free  enzyme  has  only  75%,  40%  and  0%  of  the  control  ac- 
tivity after  incubation  for  20  minutes  at  30°  C,  38°  C  and  50°  C  re- 
spectively. Stability  of  cytochrome  oxidase  activity  in  mitochondria  is 
much  greater  than  in  the  purified  lipid-free  state  (Figs.  9  and  10). 

Wallach  and  Gordon  (14)  proposed  that  in  membrane  formation  the 
various  subunits  were  assembled  to  form  a  protein  lattice  penetrated  by 
cylinders  of  lipid.  We  propose  that  a  model  membrane  made  from  inter- 
action of  purified  membrane  protein,  cytochrome  oxidase,  and  phospho- 
lipid may  simulate  in  part  the  membrane  structure  in  mitochondrial 
cristae  as  in  Figure  11.  Upon  addition  of  phospholipid  to  lipid-free  cyto- 
chrome oxidase  a  complex  forms  between  these  two  components,  and 
structural    transformation    occurs    parallel    to    and    in    as    pronounced    a 


120  Indiana  Academy  of  Science 

fashion  as  the  enhancement  of  enzyme  activity.  Protein  molecules,  as 
membrane  subunits,  are  assembled  to  form  mosaic  structure  with  phos- 
pholipid. After  formation  of  protein-lipid  complex,  the  enzyme  is  much 
more  insensitive  to  heat  treatment  than  the  lipid-free  enzyme.  The  mem- 
brane of  cytochrome  oxidase  may  represent  a  model  for  some  biological 
membranes  in  vivo.  Further  comparison  between  this  membrane  model 
and  the  mitochondrial  cristae  membrane,  with  respect  to  sensitivity  to 
enzyme  attack  (e.g.,  phospholipases  and  proteases)  as  well  as  com- 
parison of  molecular  conformations  will  be  necessary  before  we  can  tell 
how  closely  the  model  membrane  reflects  the  structure  in  cristae. 


Literature  Cited 

1.  Ball,  E.  G.,  C.  F.  Strittmatter,  and  O.  Cooper.  1951.  The  reaction  of  cytochrome 
oxidase  with  carbon  monoxide.  J.  Biol.  Chem.  193  :635-647. 

2.  Chuang,  T.  F.,  F.  F.  Sun,  and  F.  L.  Crane.  1969.  Proteolipids  VI.  The  dual  role  of 
phospholipid  in  cytochrome  oxidase  reaction.  J.  Bioenergetics. 

3.  Criddle,  R.  S.,  and  R.  M.  Bock.  1959.  On  the  physical-chemical  properties  of  water- 
soluble  cytochrome  oxidase.  Biochem.  Biophys.  Res.  Commun.  1:138-142. 

4.  Cunningham,  W.  P.,  and  F.  L.  Crane.  1965.  Identification  of  isolated  cellular  mem- 
branes by  electron  microscopy.  Plant  Physiol.  40:1041-1044. 

5.  Das,  M.  L.,  and  F.  L.  Crane.  1964.  Proteolipids  I.  Formation  of  phospholipid- 
cytochrome  c  complexes.  Biochemistry  3  :696-700. 

6.  Fleischer,  S.,  and  B.  Fleischer.  1967.  Removal  and  binding  of  polar  lipids  in 
mitochondria  and  other  membrane  systems,  p.  406-433.  In  R.  W  Estabrook  and  M.  Z. 
Pullman    [eds.  1   Methods  in   Enzymology,  Vol.   X.   Academic   Press,   New   York. 

7.  Low,  H.,  and  I.  Vallin.  1963.  Succinate-linked  diphosphopyridine  nucleotide  reduc- 
tion in  submitochondrial  particles.   Biochim.   Biophys.   Acta  69:861-374. 

8.  McConnell,  D.  G.,  A.  Tzagoloff,  D.  H.  MacLennan,  and  D.  E.  Green.  1966.  Studies 
on  the  electron  transfer  system  LXV.  Formation  of  membranes  by  purified  cytochrome 
oxidase.  J.  Biol.  Chem.  241 :2373-2382. 

9.  Prezbindowski,  K.  S.,  F.  J.  Ruzicka,  F.  F.  Sun,  and  F.  L.  Crane.  1968.  A  double 
layer  of  protein  in  mitochondrial  cristae.  Biochem.   Biophys.  Res.  Commun.  31:164-169. 

10.  Sun,  F.  F.,  K.  S.  Prezbindowski,  F.  L.,  Crane,  and  E.  E.  Jacobs.  1968.  Physical 
state  of  cytochrome  oxidase,  relationship  between  membrane  formation  and  ionic 
strength.  Biochim.  Biophys.  Acta  153:804-818. 

11.  Takemori,  S.,  I.  Sekuzu,  and  K.  Okunuki.  1961.  Studies  on  cytochrome  a.  Physico- 
chemical  properties  of  purified  cytochrome  a.  Biochim.  Biophys.  Acta  51  :464-472. 

12.  Vanneste,  W.  H.  1966.  Molecular  proportion  of  the  fixed  cytochrome  components  of 
the  respiratory  chain  of  Keilin-Hartree  particles  and  beef  heart  mitochondria.  Biochim. 
Biophys.  Acta  113:175-178. 

13.  Yonetani,  T.  1961.  Studies  on  cytochrome  oxidase  III.  Improved  preparation  and 
some  properties.  J.  Biol.  Chem.  236:1680-1688. 

14.  Wallach,  D.  F.  W.,  and  A.  Gordon.  1968.  Lipid  protein  interactions  in  cellular  mem- 
branes. Fed.  Proc.  27:1263-1268. 


CHEMISTRY 

Chairman:  James  E.  George,  DePauw  University 
Joseph  R.  Siefker,  Indiana  State  University,  was  elected  Chairman  for 

1970 

ABSTRACTS 

Determination  of  the  Hydrolysis  Constant  of  the  Stannous  Ion  by  an 
Electromotive  Force  Method.  Lusan  G.  Ong  and  Eugene  Schwartz, 
DePauw  University. — The  formal  hydrolysis  constant 

K=  [SnOH  +  ]  [H  +  ] 
Sn2  + 
for  the  hydrolysis  reaction 

Sn^+  +  H20  =  SnOH+  +  H  + 

was  obtained  at  25°  C  and  at  an  ionic  strength  of  unity  by  an  electro- 
motive force  method  employing  the  concentration  cell 

/    HC1O4(1.00m)  /   HC104(XF)  / 

Sn(Hg)     /  /    Sn(ClO4)2(0.020F)     /     (Hg)Sn 

/      Sn(ClO4)2(0.020M)    /     NaClO4(1.0-XM)        / 

in  which  x  was  varied  from  0.04  to  0.96.  The  liquid  junction  potentials 
for  the  above  cell  were  obtained  from  voltages  of  the  cell 

Ag,  AgCl/HCl  (1.0m)       /     HCI(Xm)  /        AgCl,  Ag 


/     HCI(Xm)  / 

/     NaCl(l.O-XM)        / 


The  hydrolysis  constant  of  the  stannous  ion  was  found  to  be  1.5±  0.5  X 
10  2. 

The  Infrared  Spectra  of  Coordination  Compounds.  James  Nowak,  Robert 
Williams  and  James  George,  DePauw  University. — The  infrared  spec- 
tra of  [Co(NH3)-Cl]Cl2,  [Co(NDH)nCl]Cl2,  [Cr(NH;s)5Cl]Cl2  and  a 
series  of  related  compounds  containing  ions  of  the  form  [M(NHa)5X]p  + 
(where  M  =  Co(III)  or  Cr(III)  and  X  =  H20,  F",  Cr,  Br  N02",  SCN"  and 
C032)  have  been  obtained.  The  spectra  of  these  compounds  are  very 
similar  to  those  of  other  metal-ammonia  systems  which  have  been  more 
extensively  studied.  The  absorptions  arising  from  the  vibrations  of  the 
coordinated  ammonia  molecules  occur  in  the  regions:  31,500  cm1;  1,600- 
1,650-1;  1,300-1,350  cmi;  and  800-850  cnri.  The  metal-ammonia  stretch- 
ing frequencies  are  near  500  cm*1.  The  coordinated  polyatomic  ligands 
could,  in  all  cases  studied,  be  distinguished  from  their  ionic  counterparts. 

pH  Dependent  Isotope  Effects  on  the  Flavin  Enzyme  L-Amino  Acid  Oxi- 
dase. Robert  L.  VanEtten  and  David  S.  Page,  Purdue  University. — 
The  effects  of  changes  in  pH  and  of  deuterium  substitution  upon  the 
kinetic  parameters  of  the  reaction  of  L-amino  acid  oxidase  with  L-leucine 
have  been  examined.  The  Michaelis-Menten  parameter  KIH  was  measured 
for  the  L-amino  acid  oxidase — L-leucine  system  as  a  function  of  pH  in 
H20  at  25,  30  and  35°  C  in  the  pH  range  5.5  to  9.5.  Treatment  of  the 

121 


122  Indiana  Academy  of  Science 

data  according-  to  Dixon's  rules  yields  pK  values  for  a  group  situated  in 
the  active  center  (i.e.,  in  the  ES  complex)  and  for  a  group  in  the  free 
enzyme.  The  temperature  dependence  of  these  pK  values  allows  the 
calculation  of  an  enthalpy  of  ionization  of  7-8  kcal/mole  for  this  ioniz- 
able  group.  This  value  together  with  the  observed  pK  values  lead  to  the 
conclusion  that  a  histidyl  group  is  involved  in  the  catalytic  interaction 
between  enzyme  and  substrate.  Studies  of  the  pK  dependence  of  the 
system  in  D20  at  25°  C  reveals  substantial  D20  solvent  isotope  effect, 
particularly  at  lower  pD  values.  A  major  part  of  this  D20  effect  is  at- 
tributable to  effects  upon  the  ionization  constant  of  one  or  more  cata- 
lytically  active  groups.  Such  an  interpretation  is  consistent  with  the  ob- 
served shift  of  0.7  units  in  the  pK  value  of  the  catalytically  important 
group  described  previously,  as  well  as  the  changed  pKa  of  free  imida- 
zole in  D20.  When  DL-[a--H]leucine  is  employed  as  a  substrate  there  is 
observed  a  strikingly  pH  dependent  kinetic  isotope  effect,  with  a  value 
of  kn/kD  of  4.0  being  observed  at  pH  values  less  than  6.5,  but  approaching 
1.0  at  pH  values  above  8.5.  Stopped-flow  experiments  have  been  con- 
ducted to  locate  the  particular  steps  in  the  action  mechanism  which  are 
affected  by  deuterium  substitution. 

A  Kinetic  Study  of  the  Decarboxylation  of  Duroic  Acid  in  Sulfuric  Acid 
Solutions.  John  T.  Snow  and  Gerald  R.  Barker,  Earlham  College. 
— The  decarboxylation  of  diortho-substituted  benzoic  acids  in  strong 
mineral  acids  has  long  been  known  and  is  of  some  synthetic  use,  but  the 
mechanism  of  the  reaction  is  not  well  understood.  Kinetic  data  are  pre- 
sented on  the  decarboxylation  of  duroic  acid  in  various  concentrations  of 
sulfuric  acid  and  at  several  different  temperatures.  The  reaction  is  first 
order  in  duroic  acid,  but  the  rate  dependence  on  sulfuric  acid  concen- 
tration allows  no  simple  interpretation.  Several  possible  mechanisms 
are  presented. 

Kinetic  data  were  obtained  by  measuring  evolved  carbon  dioxide. 
The  development  of  the  gasometric  technique  is  discussed  and  a  com- 
parison is  made  of  the  gasometric  method  and  the  infrared  and  ultra- 
violet spectrophotometric  methods  of  analysis. 

Molecular  Complexes  of  Bromine  and  Various  Substituted  Carbostyrils 
and  their  Hydrolysis  Products.  Donald  J.  Cook,  DePauw  University. — 
A  number  of  1:1  molecular  compounds  between  bromine  and  various 
carbostyrils  have  been  prepared.  These  compounds  are  fairly  stable  at 
room  temperature  and  are  inert  in  nonpolar  solvents.  However,  in  the 
presence  of  water,  hydroxide  ion,  or  pyridine,  the  molecular  compound 
is  destroyed  resulting  in  the  substitution  of  a  bromine  on  the  three  or 
six  position  of  the  carbostyril.  Identification  of  the  substituted  bromine 
carbostyrils  was  made  by  infrared  studies  and  by  independent  synthesis 
of  the  compound  by  known  methods.  The  position  of  the  bonded  bromine 
molecule  on  the  carbostyril  is  not  known  but  some  studies  with  the 
nuclear  magnetic  resonance  have  been  initiated  to  determine  the  struc- 
ture. 

The  molecular  compounds  have  also  been  shown  to  be  brominating 
agents  for  alkenes  and  ketones. 


Temperature  Dependence  of  E°  for  the  Daniel]  Cell 

Sister  Barbara  Buckbeei,  Ronald  E.  Surdzial^  and  Clyde  R.  Metz-% 

Indiana  University 

Abstract 

EMF  measurements  for  the  cell  Zn/ZnSOj  (1m)//CuS04  (lM)/Cu  were  made  at  various 
temperatures  over  the  range  of  0-50 °C.  The  equation  describing  the  voltage  as  a  function 
of  temperature  is 

E°(volt)  =  (1.1028  ±  0.0026)  —  (0.641  ±  0.425)10-3  t  +  (0.72  ±  0.87)10-5  t2 
where  t  is  the  Celsius  temperature.  This  result  compares  favorably  with  electrochemical 
measurements  reported  in  the  literature  for  similar  cells,  but  the  derived  values  of  ^G°, 
AS°  and  /\H°  differ  considerably  with  accepted  thermodynamic  values.  This  disagreement 
probably  results  from  the  presence  of  a  residual  voltage  and  corresponding  temperature 
coefficient  inherent  in  the  cell  from  incomplete  elimination  of  the  liquid  junction  potential. 

The  conventional  Daniell  cell,  Zn/Zn-  +  //Cu2  + /Cu,  is  often  used  in 
general  chemistry  courses  to  demonstrate  the  calculation  of  overall  cell 
voltage  by  combining  half-cell  potentials.  Using  the  data  commonly 
found  in  textbook  tables  (3)  for  the  Zn/Zn-+  and  Cu/Cu2+  couples, 
-0.763  v  and  0.337  v,  respectively,  the  predicted  value  of  E°  at  25 °C  is 
1.100  v.  Upon  careful  construction  of  the  cell  using  0.5m  solutions  of 
the  nitrates  or  sulfates  of  the  metals,  the  observed  voltage  is  somewhat 
lower  than  the  predicted  value  (11). 

For  a  cell  containing  the  sulfates  of  the  metals,  the  overall  cell 
potential  E(exp)  is  given  by 

RT 

E(exp)=E°  — lnQ  +  E(LJ)  [1] 

nF 

where  Q,  the  thermodynamic  activity  quotient,  is  denned  as 

Q  =:  aZn-+  aCu/aCu2+  aZn  =    a2ZnS0^    aCu/a2CllS0^    aZn    [2] 

and  E(LJ)  is  the  liquid  junction  potential  arising  from  the  ZnS04-KCl- 
CuS04  interfaces.  The  activities  of  the  metals  in  Equation  [2]  are  unity 
by  convention  and  the  product  of  concentration  and  mean  activity  co- 
efficient gives  the  mean  activities  for  the  salts.  Using  the  values  given 
by  Robinson  and  Stokes  (8)  for  the  activity  coefficients,  the  emf  contri- 
bution in  Equation  [2]  for  the  activity  term  is  negligible  for  the  1m 
concentrations. 

As  written  above,  the  cell  contains  a  salt  bridge  to  minimize  the 
liquid  junction  potential.  Maclnnes  (5)  states  that  E(LJ)  is  negligible 
provided  the  ionic  mobilities  of  the  cation  and  anion  in  the  bridge  are 
equal.  If  a  saturated  KC1  bridge  is  used,  this  equality  is  nearly  achieved 
at  25°  C.  Thus  any  observed  voltage  should  represent  E°  for  the 
chemical  reaction  of  interest:  Zn  +  Cu-+  =  Zn-  +  -f  Cu. 


1  Current  address  :  St.  Joseph  High  School,  Brooklyn,  N.Y. 

2  Current  address:  Highland  High  School,  Highland,  Indiana. 

3  Current  address  :  Indiana  University-Purdue  University  at  Indianapolis. 

123 


124  Indiana  Academy  of  Science 

Experimental 

The  half  cells  consisted  of  strips  of  Zn  and  Cu  metal  (10x1x0.1  cm) 
dipping  into  1M  solutions  of  ZnS04  and  CuS04,  respectively.  To  reduce 
heat  transfer  from  the  solutions  to  the  air,  the  electrodes  were  thermally 
insulated  using  1-inch  thicknesses  of  plastic  foam.  The  half  cells  were 
joined  by  a  salt  bridge  consisting  of  a  saturated  KC1  solution  suspended 
on  a  strip  of  chromatographic  filter  paper.  To  reduce  possible  interaction 
between  the  metals  and  KC1,  the  electrodes  and  a  small  quantity  of 
solution  were  isolated  from  the  bulk  solution  by  enclosing  them  within 
glass  tubing  which  was  drawn  to  a  capillary  tip.  At  each  temperature 
fresh  solutions  and  a  newly-constructed  salt  bridge  were  used.  All  chemi- 
cals were  of  reagent  grade  quality  and  standard  quantitative  procedures 
were  used  in  preparation  of  the  solutions. 

The  complete  Daniell  cell  was  placed  in  a  constant  temperature  bath 
and  the  temperatures  of  the  half  cells  were  monitored  by  separate 
thermometers.  Once  temperature  equilibrium  was  reached,  the  tempera- 
ture was  recorded  to  the  nearest  0.05°  C  (corrected  for  stem  immersion) 
and  the  open-cell  voltage  was  read  from  a  Honeywell  Potentiometric 
Voltmeter,  Model  852,  to  the  nearest  0.1  mv.  At  the  sensitivity  used,  the 
input  impedance  was  10  Mil,  so  negligible  current  was  drawn  from  the 
cell. 

Results 

The  data  appear  in  Figure  1.  Using  the  method  of  Bennett  and  Frank- 
lin (1),  the  regression  coefficients  for  a  linear  and  a  quadratic  dependence 
on  temperature  were  determined.  An  analysis  of  variance  indicated  the 
quadratic    term    to    be    significant    and    the    corresponding    equation    is 

E°  (volt)  =  (1.1028  ±  0.0026)  —  (0.641  ±  0.425)10 3  t+ 
(0.72  ±  0.87)10-5  t2  [3] 

where  t  is  the  Celsius  temperature  and  varies  from  0°  to  50°  C.  The 
standard  deviation  is  1.36  mv  and  the  estimated  errors  in  the  regression 
coefficients  are  calculated  on  a  95%  probability  limit  basis. 

The  maximum  random  error  in  the  voltage  resulting  from  tempera- 
ture measurement  is  negligible,  ±  0.02  mv.  The  error  resulting  from 
temperature  gradients  is  estimated  as  ±  0.5  mv  based  on  observations 
made  with  nonisothermal  cell  conditions. 

The  important  thermodynamic  properties  AG0,  AS°  and  AH0  can  be 
derived  from  Equation  [3].  These  are 

AG°(kcal/mole)  =  -nFE°  =  -50.863  +  0.0296  t  —  0.33x10 -3  t-'       [4] 

AS°   (gibbs/mole)  =  -d(AG°)/dT  =  -29.6  +  0.66  t  [5] 

AH°(kcal/mole)  =  AG°  +  TAS°  =  -58.948  +  0.1813  t  + 

0.33x10-3  t2.  [6] 

Table  1  contains  values  of  these  properties  and  accepted  thermodynamic 
values  (10).  Although  the  quadratic  equation  for  AG°  allows  the  esti- 
mation of  AC°p,  the  confidence  limits  in  the  regression  coefficients  are 
too  large  to  provide  reliable  values. 


Chemistry 


125 


%L 


10    20     30    40   50 
TEMPERATURE    (°C) 


Figure    1.     Plot   of   experimental   emfs   for   the   cell   Zn/ZnSO',(  1m)//CuSOj,( lu) /Cu.    The 
curve  is  the  least  squares  equation. 


Table   1.     Thermodynamic   values  for   the   Daniell  cell  calculated  from 

Equations  [3] -[6]. 

t  (°C)  E°  (volt)    AG°  (kcal/mole)       AS°  (gibbs/mole)     AH°  (kcal/mole) 


0 

1.1028 

—50.863 

—29.6 

—59.948 

10 

1.0971 

—50.600 

—23.0 

—57.168 

20 

1.0929 

—50.403 

—16.4 

—55.354 

25 

1.0913 

— 50.329  (- 

-50.71)* 

— 13.1(- 

-3.73)* 

— 54.621(- 

-51.82)* 

30 

1.0901 

—50.272 

—9.8 

—53.806 

40 

1.0887 

—50.207 

—3.2 

—52.224 

*  The  values  given   in   parentheses  are  accepted   thermodynamic   values 
(10). 


Discussion 

The  calculated  value  of  E°  at  25°  C  from  Equation  [3]  of  1.0913  ± 
0.0026  v  is  lower  than  the  predicted  value  of  1.100  v  by  roughly  9  mv. 
Reported  values  for  E°  at  15°  C  of  1.09337  by  Cohen,  Chattaway  and 
Tombrock  (2)  for  a  cell  consisting  of  amalgamated  electrodes  in  satu- 
rated solutions  and  1.0962  v  (average)  by  Jahn  (4)  give  similar  differ- 
ences between  experimental  and  calculated  values. 


126  Indiana  Academy  of  Science 

The  value  of  dE°/dT  at  25°  C  calculated  from  Equation  [3]  is 
— 0.289  mv/deg.  This  value  compares  favorably  to  — 0.429  mv/deg 
reported  by  Cohen,  Chattaway  and  Tombrock  (2),  to  — 0.2  mv/deg  re- 
ported by  Rosset  (9),  and  to  — 0.182  mv/deg  listed  by  deBethune  and 
Loud  (3)  as  an  experimental  determination.  These  results  are  consider- 
ably more  negative  than  — 0.083  mv/deg  as  predicted  by  combining  half 
cell  values  of  dE°/dT  based  on  thermodynamic  values  given  by  deBethune 
and  Loud  (3).  The  thermodynamic  value  for  AS°  in  Table  1  corresponds 
to  — 0.0809  mv/deg  (10).  Considering  the  similarities  of  the  species  in- 
volved in  the  cell,  the  smaller  thermodynamic  values  appear  to  better 
express  AS0  for  the  reaction  than  do  the  electrochemical  values. 

The  source  of  the  discrepancies,  9  mv  and  0.2  mv/deg,  probably 
results  from  the  last  two  terms  in  the  expressions  for  the  overall  cell 
potential,  Equation  [1],  and  the  corresponding  temperature  coefficient 

dE(exp)        dE°        R  d  In  Q  dE(LJ) 

-dT-  =  ^-^(lnQ  +  T-l^)+"^  [7] 

which  are  present  because  of  experimental  conditions. 

As  mentioned  earlier,  the  emf  contribution  of  the  second  term  in 
Equation  [1]  is  negligible  at  25°  C  and  so  the  9  mv  must  be  the  result 
of  the  two  liquid  junction  potentials  for  the  ZnS04-KCl  and  KCl-CuS04 
interfaces.  Maclnnes  (5)  gives  the  following  general  equation 

soln  II 

soln  I 

which  must  be  applied  at  each  interface.  Unfortunately  Equation  [8] 
cannot  be  integrated  directly,  but  Maclnnes  (5)  describes  two  approxi- 
mate methods  for  obtaining  values  of  E(LJ). 

The  first  approximation  is  based  on  the  assumptions  that  ionic 
mobilities  are  independent  of  concentration  and  that  activities  and 
concentrations  are  equal.  This  results  in 


E(LJ)=f±l^fli/z'i>^i-^>      In    f-^f-  [9] 


RTn  Wz,)  (Ci  —  Ci)  nC./ii 

n  Mi  (Ci"— c!)  nC     ^i 

where  /m  is  the  ionic  mobility.  Using  data  from  Milazzo  (7)  for  Equation 
[9],  one  obtains 

E(LJ)  =  E(LJ)  +  E(LJ)  =  —1.3  +  1.6  =  0.3  mv 

ZnSO,  —  KC1  KC1— CuSO^ 

which  is  considerably  less  than  the  9  mv  and  has  the  incorrect  sign. 

The  second  technique  for  obtaining  E(LJ)  values  from  Equation  [8] 
is  by  graphical  integration  of  values  of  t-f/Cf  plotted  against  Cf  between 
the  appropriate  concentrations.  Following  the  procedure  given  by 
Maclnnes   (5)  and  assuming  that  the  activities  of  the  sulfate  ion  in  1m 


Chemistry  127 

ZnS04  and  in  lM  CuS04  are  equal  and  that  the  changes  in  the  activities 
of  Cl~,  SOj'-  and  K2S04  passing  from  the  salt  bridge  to  1m  ZnS04  and 
lM  CuS04  to  be  identical,  the  total  liquid  junction  potential  is  given  by 

sat  KC1 

— RT    r       O  'Zn~  +  d(C  f  ) 

E(LJ)   =       ~-  I  Vj      q~       ~t~  ZnS04     ZnSO, 


/  V        ZnS04     ZnS04 

"   1m  ZnS04 

sat  KC1 

S{Cu^+  d(C 

C  f 

CuS04     CuS04 
1m  CuS04  Llt'J 


■    •    >7 

CuS04     CuS04     / 


Using  available  data  (5)-(8)  and  making  the  approximation  that  the 
activity  coefficients  for  the  components  of  mixtures  to  be  equal  to  the 
activity  coefficients  for  the  pure  components  at  the  concentration  of  the 
component,  the  graphical  integration  of  Equation  [10]  gives 

E(LJ)  =  —38.7  +  38.3  =  —0.4  mv. 

Although  the  magnitude  is  low,  the  sign  of  E(LJ)  is  correct  and  if 
proper  data  were  available,  Equation  [10]  should  predict  values  more  in 
line  with  the  9  mv.  It  is  felt,  therefore,  that  the  9  mv  difference  can  be 
accounted  for  by  the  value  of  E(LJ). 

From  data  reflecting  the  temperature  dependence  of  the  limiting 
ionic  conductances  (7),  Equation  [9]  can  be  used  to  estimate  dE(LJ)/dT. 
For  a  2%  increase  in  the  ionic  mobilities  for  an  increase  in  temperature 
of  one  degree,  the  value  of  E(LJ)  increases  by  0.1  mv.  Thus  a  significant 
portion  of  the  0.20  mv/deg  discrepancy  is  accounted  for  by  the  dE(LJ)  /dT 
term  of  Equation  [7].  No  attempt  was  made  to  estimate  dE(LJ)/dT 
from  Equation  [10]  because  of  the  uncertainty  in  the  assumptions  regard- 
ing the  choice  of  data. 

In  conclusion,  the  values  of  E°,  AG°,  AS°  and  AH0  given  in  Table  1 
are  valid  for  the  experimental  Daniell  cell  which  includes,  by  necessity, 
a  liquid  junction. 


Acknowledgment 

Two  of  the  authors  (SBB  and  RES)  were  participants  in  the  Institute 
for  High  School  Chemistry  Teachers  sponsored  by  the  National  Science 
Foundation  at  Indiana  University. 


128  Indiana  Academy  of  Science 

Literature  Cited 

1.  Bennett,   C.    A.,   and   N.   L.    Franklin.    1954.    Statistical   Analysis   in   Chemistry   and 
the  Chemical  Industry.  John  Wiley  and  Sons,  New  York.  724  p. 

2.  Cohen,    E.,    T.    D.    Chattaway,    and    W.    Tombrock.    1907.    Zur    thermodynamik    der 
normalelemente.  Z.  Physik.  Chem.  60:706-727. 

3.  deBethune,  A.   J.,   and  N.   A.   Swendeman  Loud.   1964.   Standard  Aqueous   Electrode 
Potentials  and  Temperature  Coefficients  at  25°C.  Clifford  A.  Hampel,  Skokie,  111.   19  p. 

4.  Jahn,     H.     1886.     Uber    die     Beziehung    von    chemischer    Energie    und    Stromenergie 
galvanischer  Elemente.  Wied.  Ann.  28  :21-28. 

5.  MacInnes,    D.    A.    1961.   The   Principles   of   Electrochemistry.    2nd.    ed.    Dover    Publica- 
tions, New  York.  478  p. 

6.  Maron,    S.    H.,    and    C.    R.    Prutton.    1958.    Principles    of    Physical    Chemistry.    The 
Macmillan  Company,  New  York.  789  p. 

7.  Milazzo,   G.    1963.   Electrochemistry.   Elsevier   Publishing  Company,   New  York.   708   p. 

8.  Robinson,  R.  A.,  and  R.  H.  Stokes.  1949.  Tables  of  Osmotic  and  Activity  Coefficients 
of  Electrolytes  in  Aqueous  Solution  at  25°C.  Trans.   Faraday  Soc.   45  :612-624. 

9.  Rosset,   G.    1904.   Daniell  Cell  as   Standard  Cell   for  Technical   Purposes.   Electrochem. 
Ind.  2:246. 

10.  Rossini,  F.  D.,  D.  D.  Wagman,  W.  H.  Evans,  S.  Levine,  and  D.  Jaffe.  1952.  Selected 
Values  of  Chemical  Thermodynamic   Properties.  Nat.   Bur.   Stand.   Circ.   500.   1268  p. 

11.  Young,  J.  A.,  and  J.  G.   Malik.   1969.   Chemical  Queries.  J.   Chem.   Educ.   46:227-228. 


The  Study  of  Complexes  of  Di-n-butyloxamidine  with 
Transition  Metals 

Warren  E.  Hoffman,  Maurice  Jacobs,  Gerhard  Kennepohl,  Dennis 

W.    Parrot,    Phillip    Reed,    Theodore    R.    Stout    and    James    Sundy, 

Indiana  Institute  of  Technology. 

Abstract 

The  determination  of  solution  complexes  of  di-n-butyloxamidine  with  Cr(III),  Mn(II), 
Fe(III),  Co  (III),  Ni(Il),  and  Cu(II)   was  made  using  Job's  method  of  continuous  variation. 

Upon  investigation  of  the  chemical  reactions  of  aliphatic  oxamidines 
(3,  4),  we  found  that  Cu(II)  and  Ni(II)  salts  formed  solid  metal  com- 
plexes of  the  general  formula, 

(  R-NH-C-C-NH-R  )  •  MCI,  •  2H,0,  whereas  the  Mn(II)  salt  formed 

II   II 
HNNH 

(  R-NH-C-C-NH-R  )  •  MCI,  •  4H,0.  Attempts  at  that  time  to  obtain 

II   II 
HNNH 

solid  complexes  with  Co(II),  Co(III),  Fe(II),  Fe(III),  and  Cr(III)  failed. 
However  the  color  changes  with  these  salts  pointed  to  the  formation  of 
solution  complexes.  Subsequently,  a  brown  Fe(III)  solid  complex  was 
isolated  using  benzene  as  solvent  (1). 

Recently,  we  have  re-examined  the  solutions  involving  the  di-n- 
butyloxamidine  with  Cr(III),  Mn(II),  Fe(III),  Co(II),  Ni(II)  and  Cu(II). 
Applying  Job's  method  of  continuous  variation  (2),  we  have  attempted  to 
establish  the  ligand  to  metal  ion  ratio  for  any  complexes  that  might  be 
formed  in  these  solutions.  All  the  aforementioned  metal  (II)  ions  form 
complexes  in  solution  with  the  ligand  to  metal  ion  ratio  of  2:1.  The 
Cr(III)  and  Fe(III)  ions  appear  to  form  more  than  one  complex  in 
solution.  Thus,  Job's   (2)   method  is  not  applicable. 

Assuming  a  reaction  in  solution  to  be  pM  -f  qN  =  MpNq  and  deter- 
mining p  and  q,  Job's  (2)  method  shows  a  plot  of  change  in  the  absorb- 
ance  versus  mole  fraction  to  yield  a  maximum  or  a  minimum  at  the 
stoichiometric  composition  of  the  complex.  If  the  measurements  are  made 
at  several  wavelengths,  all  should  yield  the  same  result.  The  method  of 
continuous  variation  can  yield  reliable  results  only  when  the  following 
conditions  are  observed: 

1)  Only  a  single  chemical  reaction  occurs  in  the  solution.  There  is 
no  association,  protolysis,  solvolysis,  etc.,  of  either  the  reactants  or  the 
products. 

2)  The  law  of  mass  action  is  applicable  in  terms  of  concentration. 

3)  The  reactants  form  only  one  complex. 

129 


130  Indiana  Academy  of  Science 

Methods 

1)  A  stock  solution  of  di-n-butyloxamidine  of  0.01  M  (0.001  for 
Fe(III))  was  made  in  anhydrous  alcohol,  either  methanol  or  ethanol.  The 
choice  of  alcohol  made  little  or  no  difference  in  the  results.  In  the  case 
of  Mn(II),  a  water  solution  of  about  0.1  m  was  prepared.  Due  to  the 
ready  facility  of  hydrolysis  of  the  oxamidines,  water  solutions  should 
he  avoided  except  where  they  can  be  used  immediately. 

2)  The  metal  salts  were  dissolved  in  the  same  solvent  and  made  up 
to  the  same  strength  as  the  corresponding  oxamidine  solution.  The 
Cr(III),  Co  (II)  and  Ni(II)  salts  were  all  hydrated.  The  Mn(II)  salt  was 
dissolved  in  water. 

3)  The  spectra  of  solvent,  metal  salt  solution,  and  oxamidine  solution 
were  determined  using  a  Beckman  DB  spectrophotometer.  Comparison  of 
these  spectra  with  one  of  the  solution  containing  both  metal  ion  and 
oxamidine  led  to  a  selection  of  wavelengths  at  which  Job's  (2)  method 
would  be  applied.  The  absorbance  of  the  mixtures  as  the  mole  fraction  of 
oxamidine  varied  from  0.00  to  1.00  was  read  using  a  Beckman  DU 
spectrophotometer. 

4)  The  room  in  which  the  experiments  were  performed  and  the  in- 
struments housed  was  held  at  68  °F  and  under  30%  relative  humidity. 

5)  The  wavelengths  at  which  Job's  method  for  each  of  the  salts  was 
applied  are  listed  in  Table  1. 

6)  The  plot  of  A  versus  wavelength  for  the  Ni( II) -oxamidine  system 
using  methanol  as  solvent  is  shown  in  Figure  1. 

7)  The  plot  of  A  versus  mole  fraction  of  oxamidine  is  shown  in 
Figure  2. 

Figures  1  and  2  are  representative  of  the  types  of  curves  obtained 
for  all  the  salts  with  the  exception  that  for  Cr(III)  and  Fe(III),  Figure 
2  had  more  than  one  maximum  or  minimum  or  both. 

Table  1.    Wavelengths  for  application  of  Job's  Method  to  salts  of 
transition  metals. 


Salt 

Wavelengths  (mu) 

Cr(III) 

370,  380,  540,  550 

Mn(II) 

370,  380,  400 

Fe(III) 

355,  365,  375 

Co(II) 

390,  400,  410 

Ni(II) 

460,  480,  500 

Cu(II) 

530,  560,  590 

Summary 

Figure  2,  which  is  typical  of  the  data  obtained  with  the  M-+  ions, 
indicates  a  solution  complex  of  oxamidine /M-+  of  2:1.  Work  is  continu- 
ing using  other  methods  to  determine  the  nature  of  the  M;5  +  complexes 
formed  in  solution. 


Chemistry 


131 


2.4r 

2.2 


2.0- 


ujl.4 

I'2 
$  1.0- 

< 

0.8 
06- 
0.4- 
02- 


NiCI2  Solution 
■Oxamidine  Solution 
■Complex  Solution 


200 


280  3  20  3  60  400  440 

Wavelength,  mu 


480 


520 


560 


600 


Figure  1.    Absorbance  vs.  wavelength  for  NiCk  system. 


132 


Indiana  Academy  of  Science 


uj 
o 


1.3 
1.21- 
U 
1.0 
0.9- 

0&' 
0.7  • 


4$0mu 

-480  mu 
•500mu 


0.1     02     Q3   0.4    05    Q6   07      08     0.9 

Mole  Fraction  Oxamidine 


Figure  2.    Ni-\-2-oxamidine  complex. 


Chemistry  133 

Literature  Cited 

1.  Jay,  Theodore.  1964.  The  Reaction  of  Ferric  Chloride  with  Di-n-Butyloxamidine.  M.S. 
Thesis.  Indiana  Institute  of  Technology,  Ft.  Wayne. 

2.  Job,  P.  1928.  Method  of  Continuous  Variation,  Ann.  de  Chimie  9:113(10). 

3.  Woodburn,  H.  M.,  and  W.  E.  Hoffman.  1958.  The  Chemistry  of  Oxamidines  I.  J.  Org. 
Chem.  23  :262-268. 

4.  Woodburn,  H.  M.,  R.  R.  Salvesen,  J.  R.  Fisher,  W.  E.  Hoffman,  E.  L.  Graminski, 
and  R.  L.  Van  Deusen.  1967.  Metal  Complexes  of  Cyanoformamidines,  Oxamidines  and 
Oxalimidates.  J.  Chem.  and  Eng.  Data  12  :615-617. 


ECOLOGY 

Chairman:  Thomas  S.  McComish,  Ball  State  University 
James  R.  Gammon,  DePauw  University,  was  elected  Chairman  for  1970 

ABSTRACTS 

The  Annual  Growth  Cycle  of  the  Bluegill.  Thomas  S.  McComish,  Ball 
State  University,  and  Richard  0.  Anderson,  University  of  Missouri. — 
Continuous  growth  experiments  were  conducted  from  fall  1965  through 
1967  with  bluegill,  Lepomis  macrochirus,  under  similar  natural  seasonal 
photo-periodicity  and  temperature  fluctuations  to  those  occurring  in  ponds 
near  Columbia,  Missouri.  Each  fish  was  isolated  in  an  aquarium  and  fed 
frozen  chironomid  larvae  in  excess.  Growth  in  length  and  weight;  food 
consumed;  conversion  efficiency  of  live,  dry,  and  protein  weights;  and 
calories  energy  were  measured  monthly. 

Growth  of  yearling  bluegill  increased  steadily  from  March  to  a  peak 
in  June  and  July  followed  by  a  steady  decline  to  a  low  in  December. 
Food  consumed  followed  a  similar  pattern.  Growth  of  two-year-old 
sexually  mature  bluegill  started  in  March,  peaked  in  May,  decreased 
steadily  to  a  low  in  July,  followed  by  a  steady  increase  to  a  second  peak 
in  September,  and  decreased  to  a  low  point  in  November.  Food  con- 
sumed and  conversion  efficiencies  followed  similar  patterns. 

The  mid-summer  slump  in  growth  for  two-year-old  bluegill  was 
correlated  with  temperature.  The  May  and  September  growth  peaks 
occurred  at  20  to  22°  C  and  the  mid-summer  low  at  about  27°  C.  Meta- 
bolic rate  was  implicated  in  the  growth  cycle  as  a  function  of  tempera- 
ture, season,  size,  and  perhaps  sexual  maturation  in  preparation  for 
spawning. 

The  Effect  of  Photoperiod  on  Growth  of  Bluegill.  Paul  G.  Davidson  and 
Thomas  S.  McComish,  Ball  State  University. — Growth  experiments 
were  conducted  with  bluegill,  Lepomis  macrochirus,  for  102  days  under 
3  different  photoperiods  at  26°  C.  One  photoperiod  increased  from  15.50 
to  19.78  hours  daily,  another  decreased  from  15.50  to  12.25  hours  daily, 
and  a  third  was  held  constant  at  15.50  hours  daily.  Growth,  food  con- 
sumption (mealworms,  Tenebrio  molitor) ,  and  food  conversion  efficiency 
were  evaluated  for  bluegill  in  each  set  of  conditions. 

Under  the  conditions  used  in  this  experiment  there  was  no  apparent 
effect  of  photoperiod  on  the  growth  of  bluegill.  This  was  true  for  all 
measurements  of  growth  in  length,  growth  in  weight,  food  consumption 
and  food  conversion  efficiency.  It  was  also  true  when  males  and  females 
were  compared  for  each  of  these  measurements. 

These  results  were  not  expected.  A  possible  explanation  is  that  the 
relatively  high  temperature  of  26°  C  increased  metabolism  enough  to 
mask  the  effects  of  photoperiod. 

135 


136  Indiana  Academy  of  Science 

The  Response  of  Fish  to  Heated  Effluents.  James  R.  Gammon,  DePauw 
University. — The  specter  of  temperature  elevation  of  surface  waters  due 
to  electric  generating-  stations  has  caused  great  concern  to  aquatic 
ecologists  because  of  projected  construction  estimates.  The  distribution 
and  abundance  of  adult  fish  in  two  three-mile  segments  of  the  Wabash 
River  have  been  studied  since  1967.  One  site  is  bisected  by  an  860  mega- 
watt station,  while  the  other  will  receive  heated  effluents  beginning  in 
1970. 

During  normal  summer  flows  the  temperature  of  the  effluent  was 
about  7°  C  higher  than  the  river  water.  Complete  mixing  was  achieved 
about  %  mile  below  the  effluent  jetty  at  which  point  the  temperature  was 
about  4°  C  higher.  Some  species  populations  were  estimated  by  mark- 
and-recapture  techniques  using  hoop  nets  and  electrofishing  apparatus, 
but  for  most  species  only  relative  indices  of  abundance  based  on  catch 
data  could  be  obtained. 

Consistent  differences  in  the  distribution  patterns  were  noted  for 
some  species  which  seemed  related  to  temperature.  Species  which  tended 
to  avoid  the  heated  water  included  northern  river  carpsucker  (Carpiodes 
carpio  carpio),  golden  redhorse  (Moxostoma  erythrurum) ,  shorthead 
redhorse  (M.  breviceps),  spotted  bass  (Micropterus  punctulatus) ,  long- 
ear  sunfish  (Lepomis  megalotis  megalotis),  and  sauger  (Stizostedion 
canadense).  Species  which  were  significantly  more  abundant  in  the 
heated  water  included  carp  (Cyprinus  carpio),  buffalo  fish  (mostly 
Ictiobus  babalus),  gar  (Lepisosteus  osseus  and  L.  plato stomas ) ,  channel 
catfish  (Ictalurus  punctatus)  and  flathead  catfish  (Pilodictis  olivanis) . 

Preliminary  Experiments  on  Growth  of  Bluegill  with  Varied  Feeding 
Frequency  and  Constant  Ration.  Thomas  E.  Mangum,  III  and  Thomas 
S.  McComish,  Ball  State  University. — Growth  of  bluegill,  Lepomis 
macrochirus,  in  length  and  weight,  and  food  conversion  were  compared 
between  groups  of  fish  fed  once  and  three  times  daily.  Daily  ration  and 
evironmental  conditions  were  constant  between  groups.  Groups  fed 
three  times  daily  showed  greater  growth  and  conversion  efficiencies  in 
most  cases.  Differences,  however,  were  small  and  not  significant. 

The  Relationship  Between  Growth  and  Social  Hierarchy  in  the  Green 
Sunfish.  Ruth  A.  Wilsey  and  Thomas  S.  McComish,  Ball  State  Uni- 
versity.— Growth  in  weight  and  length,  and  food  conversion  efficiencies 
were  compared  between  similar  sized  control  and  paired  female  green 
sunfish  {Lepomis  cyanellus)  in  aquaria.  Growth  was  related  to  hier- 
archy. Prior  residence  and  size  were  major  factors  in  hierarchy  estab- 
lishment. Interaction  between  paired  fish  stimulated  growth  and  food 
consumption  to  an  optimal  level  beyond  which  a  decline  was  observed. 
Control  and  paired  fish  grew  more  than  subordinate  fish.  Control  fish 
converted  food  more  efficiently  than  paired  fish.  Social  interaction  re- 
sulted in  the  death  of  four  subordinate  fish.  This  was  related,  at  least 
in  part,  to  elevated  maintenance  levels  and  food  consumption  reductions. 
Growth  and  conversion  efficiency  of  dominant  fish  after  the  death  of 
subordinates  improved  in  direct  relation  to  the  length  of  the  recovery 
period. 


Ecology  137 

A    Taxonomic    Survey    of   the    Ostracods  of   Delaware    County,    Indiana. 

Daniel  R.  Goins,  Ball  State  University. — A  survey  of  ostracods  (Class 
Crustacea,  Order  Ostracoda)  was  taken  throughout  Delaware  County, 
during  May  and  June,  1968.  One  hundred  random  samples  were  taken 
from  creeks,  drainage  ditches,  farm  ponds,  temporary  streams  and  ponds, 
and  rivers  in  this  county.  A  bottom  dredge  net  was  used  to  sample  bodies 
of  water,   and  collections  were  preserved  in   50%    alcohol   in   pint  jars. 

Sixty-two  %  of  the  samples  contained  ostracods.  Many  of  these 
ostracods  were  dissected  in  the  laboratory  and  mounted  in  permanent 
media  on  microscope  slides.  The  ostracods  were  then  identified  and  re- 
corded. Comprehensive  surveys  of  ostracods  have  been  made  in  Ohio 
and  Illinois,  but  only  one  collection  previously  had  been  reported  from 
Indiana.  This  study  increased  the  known  Indiana  genera  from  3  to  19 
and  revealed  several  new  species.  Names  and  descriptions  of  these  new 
genera  and  species  will  be  published  in  the  near  future. 

The  Influence  of  Environmental  Factors  on  the  Concentration  of  Hydro- 
cyanic Acid  in  Manihot  esculenta.  Robert  D.  Hart  and  Wm.  B.  Crank- 
shaw,  Ball  State  University. — Manioc  or  yuca  (Manihot  esculenta),  a 
common  cultigen  of  milpa  agriculture  in  the  tropics,  varies  from  site  to 
site  with  respect  to  the  concentration  of  hydrocyanic  acid  in  the  tubers. 

A  short  term  study  was  conducted  to  determine  the  possibility  of  a 
correlation  between  acid  concentration  and  microclimatic  factors.  A 
site  was  selected  on  the  edge  of  the  Amazon  Basin  in  the  rain  forest 
of  eastern  Ecuador  to  conduct  the  study. 

Microclimatic  factors  were  measured  at  7  stations  along  a  60  m 
transect  through  a  milpa  of  manioc.  These  factors  included  insolation, 
precipitation,  air  temperature,  soil  temperatures,  and  soil  moisture.  At 
the  termination  of  the  field  study,  the  manioc  plant  closest  to  each 
station  was  removed  and  assayed  for  hydrocyanic  acid  content.  Soil 
samples  were  also  taken  at  each  station  and  analyzed  for  percent  nitro- 
gen, phosphate  and  potassium. 

Regression  analysis  indicated  that  the  highest  correlation  existed 
between  insolation  and  temperatures  and  acid  content.  Enzyme  action 
converting  cyanogenetic  glucoside  to  hydrocyanic  acid  seems  to  be  in- 
hibited by  the  stress  situation  of  high  air  and  soil  temperatures. 

A  Comparison  of  Dominance  Expressions  for  Tree  Species  in  Foley 
Woods,  Edgar  County,  Illinois.  M.  T.  Jackson,  Indiana  State  University, 
and  R.  0.  Petty,  Wabash  College. — Foley  Woods  is  a  120-acre  stand 
occupying  a  till  plain  depression  just  north  of  the  Wisconsin  terminal 
moraine.  The  diverse  stand  (39  tree  species)  is  notable  as  a  western 
edge  outlier  of  American  beech  near  the  prairie  border.  Sugar  maple, 
red  oak,  bur  oak,  white  oak,  shellbark  hickory,  white  and  green  ash,  and 
basswood  lead  in  importance. 

Canopy  openings  created  by  selective  cutting  of  large  trees  have 
allowed  sugar  maple  and  elm  to  increase  greatly  in  density  in  the  smaller 
size   classes.    High   densities    of   small   trees   give   these   species    a   two- 


138  Indiana  Academy  of  Science 

factor  importance  value  index  (the  average  of  relative  density  and 
relative  basal  area)  disproportionately  higher  than  their  actual  contribu- 
tion to  stand  dominance. 

To  evaluate  this  disparity,  a  10%  sample  of  the  south  80  acres  was 
tallied  by  taking  forty  1/5-acre  line  strips  (200  X  43.56  feet).  All  trees 
over  2  inches  dbh  were  recorded  and  line-intercept  crown  cover  was 
determined  for  each  plot.  Density,  frequency,  basal  area,  cover,  average 
diameter  and  average  basal  area  were  determined  for  each  species. 
Various  two-,  three-  and  four-attribute  importance  value  indices,  and 
other  synthetic  indices  were  examined.  The  indices  were  weighed  against 
relative  cover  to  determine  which  integrating  expressions  best  reflect 
the  contribution  of  each  species  to  total  stand  dominance.  A  Density- 
Double  Dominance  Index  in  which  relative  density  plus  relative  basal 
area  weighted  twice  were  averaged 


/  D3  +  B3  +  B3  \ 


gave  the  best  approximation  of  importance.  Relative  basal  area  alone 
was  the  next  most  accurate  expression,  followed  by  the  four  factor 
index  (the  average  of  the  relative  values  of  density,  basal  area,  fre- 
quency and  cover). 

Testing  the  Quarter  Method  against  Full  Tallies  in  Old-growth  Forests. 

Damian  Schmelz,  St.  Meinrad  College. — The  quarter  method,  although 
used  with  good  results  by  workers  interested  in  the  general  characteris- 
tics of  many  stands,  has  been  criticized  for  its  bias  for  species  tending 
toward  clumped  dispersal,  for  the  amount  of  work  involved  in  office 
computations  of  vegetational  attributes,  for  the  difficulty  in  computing 
standard  error,  and  for  its  relative  field-inefficiency  when  compared 
with  other  rapid  sampling  methods.  A  further  criticism  of  the  accuracy 
of  the  quarter  method  results  from  a  comparison  of  data  obtained  by 
this  method  with  that  obtained  by  full  tallies  in  seven  old-growth  stands 
in  Indiana.  The  stands  ranged  in  size  from  4  to  21  acres.  Two  or  three 
quarter  point  samples  were  obtained  for  each  stand,  giving  a  total  of 
17  samples  for  comparison.  The  number  of  points  in  each  sample  aver- 
aged two  per  acre.  The  quarter  method  consistently  overestimated  the 
stand  basal  area,  from  1.0%  to  99%,  averaging  39%.  Stand  density  was 
overestimated  in  most  samples,  by  as  much  as  60%.  Of  the  total  number 
of  species  in  each  stand,  from  43%  to  73%  were  recorded  by  the  samples. 
One  of  the  five  most  important  species  was  missed  in  five  samples,  and 
the  order  among  them  was  different  in  ten  samples.  In  one  sample  the 
most  important  species  was  different,  and  in  six  samples  the  second  most 
important  species  was  different.  The  quarter  method  is  not  judged  use- 
ful for  any  detailed  analysis  of  hardwood  forests. 


Bluegill  Predation  by  Three  Fish  Species 

William  J.  Gulish,  Indiana  Department  of  Natural  Resources 


Abstract 

Seven  ponds  at  Driftwood  Experimental  Station  were  used  to  evaluate  the  ability  of 
three  species  of  predaceous  fish  to  control  bluegill  populations.  These  were  the  largemouth 
bass,  northern  pike  and  white  catfish  (Ictalurus  catus) .  The  ponds  were  stocked  in  April, 
1967,  with  100  predators  and  1,000  bluegill  per  acre.  Two  control  ponds  were  stocked 
only  with  bluegill  at  1,000  per  acre.  Four  ponds  were  drained  in  October,  1967,  and  the 
populations  evaluated.  Three  of  the  ponds  were  refilled  and  the  fish  returned.  All  ponds 
were  drained  in  August,  1968,  and  the  populations  evaluated. 

The  largemouth  bass  was  the  most  effective  bluegill  predator  followed  by  the  northern 
pike  and  white  catfish.  The  effects  of  intraspecific  competition  on  the  dynamics  of  the 
bluegill  populations  and  its  relation  to  the  predator  population  is  discussed. 


Introduction 

The  tendency  of  the  bluegill  to  overpopulate  has  long  been  a  major 
problem  for  the  fisheries  manager.  The  problem  is  particularly  acute  in 
farm  ponds,  as  the  stocking  of  bluegill  in  these  bodies  of  water  is  wide- 
spread. This  study  was  conducted  to  evaluate  the  ability  of  three  species 
of  predaceous  fish  to  control  bluegill  populations. 


Methods 

Seven  ponds  at  the  Driftwood  Experimental  Station  were  used  in 
the  experiments.  The  ponds  range  from  0.62  to  2.05  acres  in  size,  have 
maximum  depths  of  5  feet  and  are  supplied  with  water  from  Starve 
Hollow  Lake.  Screens  placed  over  the  inlet  valves  to  prevent  the  en- 
trance of  lake  fish  were  not  completely  effective.  While  contamination 
did  occur,  it  was  not  serious  in  most  of  the  ponds. 

Three  species  of  predaceous  fish,  white  catfish,  northern  pike  and 
largemouth  bass  were  used.  Two  ponds  each  of  white  catfish-bluegill  and 
northern  pike-bluegill  were  stocked  while  one  pond  with  largemouth 
bass-bluegill  and  two  ponds  with  only  bluegill  were  used  as  controls.  The 
ponds  were  stocked  during  April  and  May,  1967.  A  rate  of  100  preda- 
tors to  1,000  bluegill  per  acre  was  used  in  all  the  ponds.  Stocking  is 
summarized  in  Table  1. 

Ponds  2,  4,  6  and  7  were  drained  during  October,  1967.  Pond  7  con- 
tained a  large  bass  (1.2  lbs.)  and  a  large  number  of  green  sunfish  and 
crappies  so  the  data  were  discarded.  All  predaceous  fish  and  bluegills 
from  the  three  remaining  ponds  were  counted,  weighed  and  measured. 
Both  the  white  catfish  and  largemouth  bass  had  spawned,  but  the  young 
were  easily  separated  from  the  original  stock  by  size.  The  same  was 
true  for  bluegill  in  all  ponds. 

All  bluegill  were  separated  by  year  class.  The  survivors  of  the 
original  bluegill  stock   (age   1)    were  counted  and  lengths   and  weights 

139 


140 


Indiana  Academy  of  Science 


Table  1.     Stocking  of  seven  experimental  ponds  at  Driftwood  Experi- 
mental Station,  April  19-May  11,  1967. 


Pond 

Area 

Bluegill 

Northern 

White 

Largemouth 

(Acres) 

Pike 

Catfish 

Bass 

1 

0.62 

No. 
Size 

620 
1.0-2.0" 

2 

1.07 

No. 
Size 

1070 
1.0-2.0" 

107 
2.5-7.0" 

3 

1.07 

No. 
Size 

1070 
1.0-2.0" 

107 
2.5-5.0" 

4 

1.23 

No. 
Size 

1230 
1.0-2.0" 

123 

1.5-4.0" 

5 

2.05 

No. 
Size 

2050 
1.0-2.0" 

205 
1.5-4.0" 

6 

1.37 

No. 
Size 

1370 
1.0-2.0" 

137 
6.0-7.0" 

7 

1.4(1 

No. 
Size 

1400 
1.0-2.0" 

were   recorded.    Young-of-the-year  bluegill   were   bulk   weighed   and   the 
total  number  calculated  from  counts  of  weighed  random  samples. 

After  processing,  all  fish  (except  for  Pond  7)  were  returned  to  the 
refilled  ponds.  As  many  as  possible  of  the  contaminating  species,  such 
as  green  sunfish,  were  sorted  out  and  discarded.  Handling  mortality 
among  the  large  fish  was  quite  low.  A  high  percentage  of  the  young- 
of-the-year  bluegill  were  lost,  however. 


Table  2.     Species  compositioyi  and  standing  crop  of  fishes  in  six  ponds 
at  Driftwood  Experimental  Station,  August,  1968. 


Pond  1 

Pond  2 

Pond  3 

Pond  4 

Pond  5 

Pond  6 

White   Catfish 

No./A. 
Lbs./A. 

117 
31.4 

7 
1.7 

Northern    Pike 

No./A. 
Lbs./A. 

10 
14.0 

10 

15.4 

Largemouth   Bass 

No./A. 
Lbs./A. 

274 
87.1 

Bluegill 

No./A. 

72,684 

29,147 

8,923 

13,429 

17,914 

8,054 

Lbs./A. 

368.2 

109.6 

131.1 

330.6 

177.5 

52.3 

Other  Sunfish 

No./A. 

5 

2 

1,785 

2,776 

6,183 

2 

Lbs./A. 

1.1 

0.3 

17.4 

14.5 

54.0 

0.7 

Crappies 

No./A. 
Lbs. /A. 

1 

186 
14.9 

4 
1.1 

Bullheads 

No./A. 
Lbs./A. 

2 
0.6 

1 
0.9 

Total 

No./A. 

72,691 

29,267 

10,901 

16,215 

24,112 

8,330 

Lbs./A. 

369.9 

141.3 

165.1 

359.1 

248.9 

140.1 

Ecology  141 

In  August,  1968,  all  ponds  were  drained.  All  bluegill  of  4  inches 
total  length  or  longer  were  sorted  into  1-inch  groups,  counted  and 
weighed.  A  minimum  of  100  fish  from  each  1-inch  group  were  measured 
to  the  nearest  0.1  inch.  The  relative  abundance  of  bluegills  below  4 
inches  was  calculated  from  random  samples  because  of  the  high  num- 
bers of  these  fish.  A  minimum  of  100  fish  from  the  2  and  3-inch  groups 
were  measured  to  the  nearest  0.1  inch.  The  remaining  bluegills  were 
then  bulk  weighed.  No  measurements  were  taken  from  fish  in  the  1-inch 
group,  which  included  all  fish  less  than  2.0  inches  long.  However,  aver- 
age weights  of  the  1-inch  group  fish  were  determined  by  counting  and 
weighing  a  minimum  of  500  fish.  All  predaceous  fish  were  sorted  into 
inch  groups,  weighed  and  measured. 

It  was  possible  to  separate  the  different  year  classes  by  their  length 
distributions.  Scale  samples  taken  as  cross  checks  showed  this  method 
to  be  sufficiently  accurate. 


Results 
Bluegill  Mortalities 

Mortality  rates  for  the  original  stock  of  bluegill  in  the  study  ponds 
were  calculated  from  the  tables  in  Ricker  (4)  and  are  given  in  Table  3. 
These  rates  were  similar  in  both  white  catfish  ponds.  The  same  was  true 
in  both  northern  pike  ponds. 

Table  3.     Survival,  seasonal  mortality  rate  (a)  and  instantaneous  rate 

of  natural  mortality  (i)  for  the  original  bluegill  stock  in  six  ponds  at 

Driftwood  Experimental  Station,  1967-68. 


Predator 

None 
Pondl 

White  Catfish 

Northern  Pike 

Bass 

Pond  2 

Pond  3 

Pond  4 

Pond  5 

Pond  6 

No./A.  stocked 

1000 

1000 

1000 

1000 

1000 

1000 

No./A.  10-67 

936 

751 

140 

a  (4-67  to  10-67) 

0.064 

0.249 

0.860 

i  (4-67  to  10-67) 

0.07 

0.29 

1.97 

Days  (4-67  to  10-67) 

L83 

174 

1S4 

No./A.  8-68 

SS4 

86] 

951 

347 

444 

104 

a  (4-67  to  8-68) 

0.116 

0.139 

0.049 

0.653 

0.556 

0.896 

i   (4-67  to  8-68) 

0.12 

0.15 

0.05 

1.06 

0.81 

2.26 

Days  (4-67  to  8-68) 

475 

470 

467 

454 

453 

490 

a  (10-67  to  8-68) 

0.080 

0.538 

0.257 

i   (10-67  to  8-68) 

0.08 

0.77 

0.30 

Days  (10-67  to  8-68) 

287 

280 

306 

Table  3  indicates  that  there  was  little  predation  by  the  white  cat- 
fish on  the  original  bluegill  stock.  Change  in  bluegill  mortality  rate  in 
Pond  2  between  drainings  was  negligible,  indicating  a  uniform  predation 
rate  over  the  study  period. 


142  Indiana  Academy  of  Science 

A  Saprolegnia  infection  in  March,  1968,  drastically  reduced  the  white 
catfish  population  in  Pond  3,  since  41  were  found  dead  from  March  19 
to  23.  When  drained  in  August,  1968,  only  seven  white  catfish  were 
found  in  the  pond.  Several  white  catfish  also  died  of  the  infection  in 
Pond  2  along  with  a  few  bluegills  in  both  ponds.  The  fungus  infection 
probably  caused  the  decrease  in  the  6-inch  bluegill  group  in  Pond  2 
(Table  4). 

The  lack  of  Age  I  fish  in  Pond  2  (Table  5)  is  a  result  of  the  1967 
draining  which  almost  completely  eliminated  the  1967  bluegill  year  class. 

Table  4.     Length  distribution  of  white  catfish  and  bluegill  from  Ponds 

2  and  3,  1967-68. 


Pond 

2 

Pond 

3 

10-19-67 

8-2-68 

7-30-68 

Inch   Group 

Catfish               Bluegill 

Catfish 

Bluegill 

Catfish 

Bluegill 

1 

6,500 

33 

30,260 

7,541 

2 

7 

6 

17 

3 

1 

12!) 

6 

379 

4 

768 

1 

500 

1,162 

5 

6 

r,4 

402 

354 

6 

9 

50 

10 

18 

1 

92 

7 

14 

5 

1 

1 

3 

s 

17 

15 

3 

9 

19 

20 

1 

Ki 

17 

17 

1 

11 

6 

5 

12 

X 

Id 

13 

•>> 

Table  5.     Number  per  acre,   average   length   and  percent   total  weight 

of  bluegill  year  classes  from  six  ponds  at  Driftwood  Experimental 

Station,  1968. 


Age  II 

Age   I 

Age  0 

Pond 

No./A. 

Ave.  L. 

%  wt. 

No./A. 

Ave.  L. 

%  Wt. 

No./A. 

Ave.  L. 

%  wt. 

1 

SS4 

5.3 

24.7 

* 

* 

* 

* 

* 

* 

2 

861 

5.0 

73.7 

6 

3.2 

o.l 

28,280 

1.3 

26.2 

3 

95 1 

5.1 

68.3 

925 

4.1 

29.9 

7,048 

1.3 

1.8 

4 

347 

5.9 

16.4 

10,938 

3.4 

82.3 

2,143 

1.3 

1.8 

r> 

444 

5.0 

21.1 

12,817 

2.7 

74.4 

1,325 

1.3 

4.5 

6 

104 

7.1 

56.9 

102 

5.4 

28.3 

7,853 

1.3 

14.S 

*  Not  separable  with  available  data. 

Total  bluegill  production  in  the  white  catfish  ponds  (Table  2)  was 
the  lowest  in  any  of  the  ponds  except  Pond  6,  which  was  overpopulated 
with  bass. 


Ecology  143 

Despite  a  high  level  of  mortality,  the  northern  pike  significantly 
reduced  the  original  bluegill  stock.  This  stock  declined  from  55  to  65% 
over  the  study  period.  However,  data  from  Pond  4  indicate  that  predation 
did  not  occur  at  a  uniform  rate  (Table  3).  Pike  predation  on  the  original 
bluegill  stock  shows  a  marked  increase  with  time.  Early  in  the  study,  the 
small  size  of  the  pike  made  the  bluegills  relatively  invulnerable  to  them. 
However,  the  pike  grew  faster  than  the  bluegill,  increasing  the  latter's 
vulnerability  accordingly.  Beyerle  and  Williams  (1)  reported  that  north- 
ern pike  fed  in  aquaria  showed  a  preference  for  the  smaller  size  groups 
of  centrarchids.  Results  from  Pond  4  indicate  the  opposite.  Pike  in  this 
pond  appeared  to  feed  selectively  on  the  larger  sizes  of  bluegill,  despite 
the  fact  that  the  pond  contained  a  huge  quantity  of  smaller  size  bluegill 
(Table  6).  Total  weight  of  the  original  bluegill  stock  decreased  from  75 
to  68  lb.  in  the  time  between  the  first  and  second  drainings.  Average 
increment  in  length  of  the  original  stock  in  this  same  time  period  was 
only  0.9  inch. 

Despite  impressive  predation  on  the  original  bluegill  stock,  northern 
pike  are  poorly  adapted  for  controlling  bluegill  populations.  This  is 
particularly  true  in  ponds,  where  their  spawning  requirements  are  un- 
likely to  be  met.  A  major  problem  in  the  pike-bluegill  combination  is  the 
difference  in  hatching  dates  between  the  two  species.  The  pike  is  an 
early  spawner.  The  young-of-the-year  become  piscivorous  (in  this  area) 
in  late  April  or  early  May. 

There  are  seldom  young-of-the-year  bluegill  available  for  forage 
before    June.    During   this    interval    the    young   pike    must    feed    on    the 


Table  6.     Length  distribution  of  northern  pike  and  bluegill  from  Ponds 
U  and  5  at  Driftwood  Experimental  Station,  1967-68. 


oup 

Pond 

Jt 

Pond 

5 

10-31 

-67 

8-7-68 

8-5-68 

Inch  Gi 

Pike 

BlueKill 

Pike 

BlueKill 

Pike 

Bluegill 

1 

22,218 

2,636 

9,540 

•1 

1,515 

22,980 

3 

31 

11,603 

3,294 

5 

537 

274 

365 

6 

L6 

150 

39 

7 

3 

3 

8 

1 

13 

5 

14 

2 

15 

5 

1 

16 

4 

1 

17 

2 

18 

4 

4 

1!) 

2 

10 

20 

1 

3 

-n 

1 

144 


Indiana  Academy  of  Science 


preceding  year  class  (age  I)  of  bluegill.  As  is  indicated  by  Table  3, 
vulnerability  of  this  size  bluegill  to  the  pike  is  low.  As  a  result,  where 
other  forage  species  are  not  available,  the  size  of  a  given  year  class  of 
pike  may  well  be  determined  by  the  abundance  and  size  of  the  pre- 
ceeding  year  class  of  bluegill.  Where  other  species  are  present,  pike  seem 
to  feed  on  them  in  preference  to  the  bluegill  (5). 


Table  7.     Length  distribution  of  largemouth  bass  and  bluegill  in  Pond 

6,  1967-68. 


10-25-67 


8-27-68 


Inch  Group        Bass 


Bluegill 


Bluegill 


60,463 


3 

87 
1 02 


112 

1 


18 
121 
23 
22 
73 


10,718 
31 

10 
25 
107 
09 
84 


In  terms  of  both  bluegill  control  and  size  of  fish  produced,  the  large- 
mouth  bass-bluegill  combination  provided  the  best  results  (Table  7). 
Because  of  the  relatively  large  size  of  the  bass  in  relation  to  the  blue- 
gill at  stocking  (Table  1),  the  vulnerability  of  the  bluegill  was  high. 
The  result  was  a  high  mortality  rate  among  the  bluegill  and  good  growth 
by  both  species.  Total  bluegill  production  in  this  pond  was  low  (Table 
2)  as  a  result  of  a  large  bass  overpopulation  (274  bass/acre)  coupled 
with  the  destruction  of  a  large  portion  of  the  1967  bluegill  year  class 
during  the  fall  draining  in  1967. 

Bluegill  Dynamics 

In  most  cases,  the  bluegill  carrying  capacity  of  a  body  of  water  is 
determined  by  the  food  supply.  The  degree  of  intraspecific  competition 
within  the  bluegill  population  is  determined  by  the  difference  between 
the  standing  crop  and  the  carrying  capacity   (2). 

The  amount  of  recruitment  into  a  bluegill  population  is  a  function 
of  the  degree  of  food  competition.  If  carrying  capacities  were  uniform, 
many  problems  could  be  solved  easily.  Unfortunately,  in  addition  to 
the  differences  in  basic  fertility  among  bodies  of  water,  carrying  capacity 
for  bluegill  may  be  influenced  by  relationships  within  the  population. 
Swingle  (6)  reported  an  inverse  relationship  between  the  percentage 
of  large  fish  in  largemouth  bass-bluegill  populations  and  the  standing 
crop. 


Ecology  145 

Gerking  (3)  has  shown  that  small  bluegill  are  more  efficient  in 
using  protein  for  growth  than  are  large  bluegill.  This  gives  the  smaller 
bluegill  an  advantage  over  the  larger  one  in  food  competition.  It  also 
suggests  that  the  potential  carrying  capacity  is  greater  for  small  blue- 
gill than  larger  ones,  increasing  the  likelihood  of  overpopulation. 

Evaluation  of  Results 

Pond  1  is  a  good  example  of  what  happens  in  the  absence  of  preda- 
tion  and  all  bluegill  control  is  intraspecific.  No  length  frequency  dis- 
tribution or  scale  samples  were  taken  from  the  1-inch  group  bluegill, 
making  positive  year  class  assignment  impossible.  However,  the  length 
distribution  curve  from  the  2-inch  group,  if  extended,  reaches  a  peak 
between  1.5  and  2.0  inches.  The  large  average  individual  size  of  the 
1-inch  group  bluegill  (321 /lb.)  indicates  that  most  of  them  were  members 
of  the  1967  year  class.  Intraspecific  competition  appeared  to  severely 
limit  bluegill  reproduction  in  Pond  1  in  1968. 

The  difference  between  the  standing  crop  and  the  carrying  capacity 
in  Pond  1  was  originally  very  large,  resulting  in  the  production  of  a 
large  1967  year  class.  This  year  class  expanded  to  fill  the  carrying 
capacity,  thereby  limiting  further  reproduction  or  growth. 

The  results  from  Ponds  2  and  3  indicate  that  the  white  catfish 
is  more  of  a  bluegill  competitor  than  a  predator.  Both  catfish  ponds 
remained  turbid  throughout  the  growing  season.  This  probably  re- 
duced the  carrying  capacity  for  bluegill.  If  white  catfish  do,  in  fact, 
reduce  the  carrying  capacity  for  bluegill,  the  difference  between  the 
standing  crop  (original  stock)  and  carrying  capacity  was  much  lower 
than  in  Pond  1  resulting  in  less  recruitment. 

The  northern  pike  mortality  in  Ponds  4  and  5  was  so  high  that  they 
never  achieved  a  population  level  capable  of  controlling  the  bluegill.  As 
in  Pond  1,  the  difference  between  the  standing  crop  and  carrying  ca- 
pacity was  large,  resulting  in  high  bluegill  recruitment.  The  apparent 
selectivity  of  the  pike  for  larger  bluegill  tended  to  aggravate  the  prob- 
lem. 

Since  northern  pike  rarely  reproduce  in  ponds,  their  applicability 
for  bluegill  control  in  ponds  seems  very  limited.  With  sufficient  effort, 
a  pike  population  might  be  maintained  at  a  level  that  would  control  the 
bluegill  population.  However,  this  can  be  said  of  almost  any  predator. 
At  present,  it  appears  that  the  largemouth  bass  is  the  best  adapted 
species  for  controlling  bluegill  population  in  ponds. 

Pond  6  developed  an  overpopulation  of  bass.  Bass  predation  kept 
the  bluegill  population  far  below  carrying  capacity.  The  standing  crop 
of  bluegill  actually  decreased  slightly  between  1967  and  1968.  While 
the  production  of  bluegill  recruits  under  these  conditions  had  to  be 
high,  there  were  sufficient  bass  present  to  limit  their  survival  to  a  very 
low  level. 


146  Indiana  Academy  of  Science 

Discussion 

Since  recruitment  into  the  bluegill  population  is  such  an  important 
component  of  population  dynamics,  the  effect  of  various  management 
procedures  on  it  should  be  kept  in  mind.  Preoccupation  with  growth 
rate  often  results  in  failure  to  consider  the  effect  of  various  procedures 
on  the  entire  system. 

A  case  in  point  is  the  stocking  rates  used  for  farm  ponds.  A  com- 
monly used  rate  is  100  fingerling  largemouth  bass  and  100  fingerling 
bluegill  per  acre.  In  ponds  of  average  fertility,  the  original  bluegill 
stock  will  rapidly  grow  to  a  large  size.  They  will  spawn  at  age  I,  pro- 
ducing a  tremendous  number  of  young.  This  very  high  level  of  bluegill 
recruitment  results  in  good  growth  of  the  original  bass  population. 
However,  it  exceeds  the  ability  of  the  bass  to  reduce  the  number  of 
bluegill  recruits  enough  to  maintain  good  bluegill  growth.  If  the  bass 
are  unable  to  reduce  the  numbers  of  the  first  bluegill  year  class  suf- 
ficiently, this  single  year  class  will  tie  up  most  of  the  ponds  carrying 
capacity  causing  low  bluegill  recruitment  the  following  year.  The  young 
bass  produced  at  this  time  find  very  few  bluegill  of  a  vulnerable  size 
and  a  high  degree  of  competition  with  the  bluegill  for  other  food  organ- 
isms. Since  the  original  bass  stock  will  have  reached  desirable  size  at 
this  stage,  they  are  subject  to  fishing  harvest,  reducing  the  number  of 
effective  predators  on  the  oversize  bluegill  year  class.  While  the  original 
stock  of  bluegill  has  also  reached  desirable  size  and  are  being  harvested, 
they  make  up  a  relatively  small  portion  of  the  total  standing  crop  and 
their  removal  does  not  appreciably  increase  the  growth  of  the  remaining 
bluegill. 

This  stocking  rate  (100  to  100)  results  in  good  growth  of  the 
original  fish  but  makes  a  bluegill  overpopulation  likely  in  short  order. 
Since  a  population  that  will  maintain  a  high  equilibrium  yield  is  desir- 
able, stocking  rates  must  be  adjusted  so  that  the  bulk  of  the  standing  crop 
will  be  composed  of  harvestable  fish.  If  a  situation  producing  a  high  level 
of  recruitment  is  set  up,  there  should  be  enough  predators  stocked  to 
control  it. 

It  is  nearly  impossible  to  recommend  a  good  stocking  rate  without 
some  knowledge  of  the  carrying  capacity  of  the  body  of  water  involved. 
However,  if  a  single  fingerling  stocking  ratio  is  to  be  used,  it  would 
be  advantageous  to  increase  the  number  of  bluegill  stocked.  This  would 
slow  the  growth  rate  somewhat,  and  reduce  recruitment  to  a  more 
manageable  level. 

An  alternative  is  to  decrease  the  survival  of  bluegill  recruits  by 
increasing  the  number  of  bass  stocked.  Which  method  would  produce 
the  highest  fishing  yield  over  an  extended  time  period  cannot  be  pre- 
dicted. 

Another  area  where  factors  affecting  recruitment  are  important  is 
that  of  partial  kills.  Nothing  seems  more  ridiculous  to  me  than  attempt- 
ing to  destroy  nests  or  young- of -the-y ear  in  a  stunted  bluegill  popula- 
tion.  Since  recruitment  into  a  stunted  population  is  low  to  begin  with, 


Ecology  147 

the  food  made  available  by  their  destruction  will  not  appreciably  alter 
the  growth  of  the  remaining  fish.  Besides,  this  reduces  the  supply  (low 
anyway)  of  vulnerable  fish  to  the  young-of-the-year  predators. 

A  more  common  practice  is  the  large  scale  reduction  in  the  bluegill 
population  concentrated  on  the  smaller  size  groups.  It  should  be  remem- 
bered here  that  the  same  factors  that  increase  growth  also  increase 
recruitment.  Where  a  large  reduction  in  standing  crop  is  made,  precau- 
tions against  the  recruitment  of  a  massive  bluegill  year  class  should  be 
taken.  This,  again,  could  take  different  forms.  One  way  would  be  to 
increase  the  predator  population  to  accommodate  the  higher  level  of 
bluegill  recruitment.  The  other  would  be  to  replace  the  destroyed  year 
class  (es)  with  a  number  of  bluegill  more  suited  to  the  carrying  capacity. 


Literature  Cited 

1.  Beyerle,   George   B.,   and  John   E.    Williams.    1968.    Some   observations   of  food   selec- 
tivity  by  northern   pike   in   aquaria.   Trans.   Amer.   Fisheries   Soc.   97(1):28-31. 

2.  Carlander,    Kenneth    D.    1966.    Relationship    of    limnological    features    to    growth    of 
fishes  in  lakes.  Verh.  Internat.  Verein.  Limnol.  16:1172-1175. 

3.  Gerking,   Shelby  D.   1962.    Production   and  utilization   in   a  population   of  bluegill   sun- 
fish.  Ecol.  Monogr.  32:31-78. 

4.  Ricker,    W.    E.    1958.    Handbook   of   computations    for   biological    computations    of   fish 
populations.  Fisheries  Res.  Board  Canada  Bull.  119. 

5.  Seaburg,    Keith    G.,    and   John    B.    Moyle.    1964.    Feeding   habits,    digestive    rates    and 
growth  of  some  Minnesota  warm  water  fishes.   Trans.   Amer.   Fisheries   Soc.   93(3)  :269- 


6.     Swingle,   H.    S.    1961.    Some   relationships   within    fish   populations   causing   fluctuations 
in  productions.  Proc.  Pacific  Sci.  Cong.  10  :43-45. 


Practicality  of  Endrin  as  a  Fish  Toxicant1 

H.  E.  McReynolds,  U.  S.  Forest  Service 

Abstract 

When  fisheries  bioassays  documented  the  lethality  of  endrin  to  fish  at  almost  ineoiv- 
ceivably  low  concentrations,  the  question  of  the  suitability  of  this  pesticide  as  a  replacement 
for  rotenone  naturally  followed.  Fisheries  workers  immediately  noted  two  characteristics  of 
this  chemical  that  would  make  it  superior  to  rotenone  as  a  piscicide :  (1)  it  was  cheaper; 
and  (2)  it  was  more  persistent,  thereby  more  likely  to  effect  a  complete  kill. 

However,  many  other  characteristics  and  effects  of  endrin  as  a  fish  toxicant  were 
unknown  or  poorly  understood.  Experiments  were  set  up  in  selected  southern  Indiana  ponds 
to  test  this  chlorinated  hydrocarbon.  Laboratory  bioassays  with  test  fish  established  an 
LCno  (Lethal  Concentration,  50r/f )  or  TLm  (Median  Tolerance  Limit)  of  about  1.3  parts 
per  billion.  Field  tests  indicated  highly  disparate  Lc's.  After  testing  in  a  diversity  of 
aquatic  situations,  it  was  postulated  that  the  amount  of  suspended  particulate  matter 
exerted  a  considerable  effect  on  the  lethality  of  the  chemical  in  field  tests.  It  was  surmised 
that  the  mechanism  of  lowered  toxicity  in  more  tui'bid  situations  was  the  adsorption 
of  the  chemical  to  organic  (and  possibly  inorganic)  suspensoids.  Verification  of  these 
postulations  was  not  pursued,  since  other  endrin  experiments  of  a  public  health  nature 
were  beginning  to  point  out  the  potential  dangers  of  endrin  in  potable  water  supplies. 
In  addition,  the  duration  of  toxicity  of  the  pesticide  showed  evidence  of  extending  beyond 
that  desired  in  the  ideal  fish  toxicant,  especially  in  the  sediments.  Later,  more  thorough 
experiments  have  generally  tended  to  establish  the  validity  of  these  surmisals.  It  was 
concluded  that  despite  the  economy  and  kill  efficiency  of  endrin,  other  undesirable  and 
dangerous  characteristics  outweighed  the  kill-cost  attributes. 


Introduction 

At  the  time  of  these  experiments,  fisheries  biologists  were — and 
still  are — looking  for  a  cheaper,  more  effective  replacement  for  rotenone 
as  a  fish  toxicant.  The  use  of  piscicides  for  eliminating  undesirable  fish 
populations  has  become  an  important  tool  in  fish  management.  While 
rotenone,  a  ketone  of  botanical  origin,  has  certain  advantages  of  low 
toxicity  to  mammals  and  low-level  of  danger  to  the  applicator,  it  also 
has  a  number  of  limitations  that  fisheries  technicians  have  found  frus- 
trating (difficulty  in  killing  resistant  species  such  as  bullhead  catfish; 
the  problem  of  getting  effective  vertical  dispersal  through  metalimnetic 
barriers;  relatively  quick  oxidation  to  non-toxic  levels;  high  toxicity  for 
many  fish  food  organisms;  and  relatively  high  cost). 

When  biologists  began  to  notice  that  organochlorine  pesticides  were 
displaying  piscicidal  potencies  many  times  greater  than  rotenone,  first 
reactions  were  dismay  and  apprehension.  After  the  first  shock  waves 
had  passed,  however,  some  biologists  began  to  wonder  if  this  chemical 
lightning  might  be  harnessed  for  fish  management  use.  Toxaphene,  a 
chlorinated  camphene,  was  the  first  to  receive  extensive  testing  as  a 
fish  toxicant,  and  was  even  used — somewhat  surreptitously — in  a  com- 
mercial fish  toxicant  product.  Unfortunately,  a  toxicity  of  extended  du- 
ration   (up   to   8   years   in    Michigan   lakes)    made    its   use   unfeasible    in 


1  This  study  was  supported  in  part  by  Federal  Aid  to  Fish  Restoration  funds    (Project 
F-4-R-8,  Indiana). 

148 


Ecology  149 

most  situations.  Interest  turned  to  other  members  of  the  generically 
similar  chlorinated  hydrocarbon  pesticides.  While  Canadian  biologists 
began  testing  thiodan,  some  American  fisheries  people  became  intrigued 
with  endrin,  an  insecticide  and  rodenticide  with  toxicity  thresholds  for 
fish  measured  in  low  or  fractioned  parts  per  billion!  This  paper  reports 
one  of  these  earlier  experiments  with  endrin. 

Laboratory  Aquaria  Tests 

Methods 

The  preliminary  tests  were  conducted  in  seven  15-gallon  metal  frame 
glass  aquaria  (Fig.  1)  having  aeration  and  filtration  systems.  Diluent 
waters  were  obtained  from  the  two  ponds  on  the  Crosley  State  Fish  and 
Game  Area  which  were  to  be  used  for  the  field  tests.  These  are  desig- 
nated merely  as  Pond  1  and  Pond  2  in  this  paper.  Two  species  of  cen- 
trarchids  were  used  as  test  animals:  the  bluegill  sunfish  (Lepomis  m. 
macrochirtis  Rafinesque),  and  the  redear  sunfish  (Lepomis  microlophus 
[Gunther]).  Fish  were  obtained  by  seining  and  were  held  in  the  aquaria 
for  10  days  before  testing  began.  The  fish  ranged  in  size  from  approxi- 
mately 1  inch  to  3  inches.  Although  this  range  does  not  meet  size- 
uniformity  recommendations  (12)  no  grading  was  done  since  these  pre- 
liminaries were  merely  exploratory  in  nature.  Actually,  the  larger  and 
smaller  fish  were  purposely  combined  in  each  aquarium  to  observe  any 
difference  in  size-susceptibility.  Two  bluegills  and  2  redears  were  placed 
in  each  of  6  aquaria  and  10  fish,  a  mixture  of  both  species,  were  held  in 
the  7th  aquarium  as  controls. 

The  endrin  was  obtained  as  a  75%  wettable  powder  and  weighed  on 
an  analytical  balance  in  the  following  amounts:  0.19,  0.38,  0.75,  and  1.50 
g.  These  weights  are  for  the  active  ingredient  and  assume  that  the  manu- 
facturer's assay  (75%)  is  correct.  The  validity  of  this  assumption  was 
not  investigated. 

The  minute  concentrations  at  which  these  tests  must  be  conducted 
present  a  problem  in  dilution.  It  was  decided  that  10  gal  of  water  would 
be  used  in  the  aquaria,  and  calculations  of  dosage  began  from  there.  To 
get  the  lowest  concentration,  0.26  parts  per  billion  (ppb),  it  was  neces- 
sary to  mix  0.19  g  active  ingredient  of  the  endrin  powder  with  5  gal  of 
water,  take  1  g  of  this  solution  and  introduce  this  amount  into  the  10 
gal  of  aquarium  water.  Addition  of  another  0.19  g  of  endrin  to  the 
original  solution  gave  the  0.52  ppb  dosage,  and  so  on  up  the  scale.  The 
test  solution  was  drawn  off  by  pipette  from  the  center  of  the  5  gal 
bottle  after  extended  agitation.  A  shell  vial  was  weighed  on  a  triple 
beam  balance  having  a  sensitivity  of  0.01  g,  and  the  weight  recorded. 
The  balance  was  set  up  one  more  gram  and  the  test  solution  was  pipetted 
into  the  vial  until  the  correct  weight  was  reached.  The  contents  of  the 
vial  were  then  dripped  into  the  aquarium  and  the  aquarium  water 
stirred  with  a  clean  glass  rod.  Hourly  checks  were  made  by  the  study 
leader  or  biologist  aides,  and  notes  were  made  on  the  reactions  of  the 
fish.   Observations  were  made   from   8:00   AM   until   4:30   pm    (the   work 


150 


Indiana  Academy  of  Science 


4-1 

x 

CNJ 

«1 

4J 
W 

i  mi 

4-1  ~ 

x 

CM 

en 

<U 

5  tin 

Hi 

«1 

IH1 

M     • 
•H    !-i 

O 


4J  <U 

r-i  5q 

3  CO 

in  en 

Q)  O 

P4  T3 


> 

m 

CO 

CO 

0J 

o 

> 

> 

> 

4-J 

5 

> 

•H 

CU 

>-i 

> 

> 

> 

r-l 

<U 

3 

U 

U 

rJ 

ex 

M 

tn 

CO 

3 

en 

o 

3 

6 

CO 

c 

en 

c 

C) 

to 

o 

CO 

o 

0 

o 

o 

•H 

en 

o 

•r-l 

0 

o 

T3 

4-J 

CN 

r. 

c 

C 

■r-l 

•r-l 

CD 

r— I 

Tl 

.  n 

Tl 

.~ 

.  r. 

en 

CO 

TJ 

X 

X 

Tl 

X 

X 

o 

C 

cc 

ex 

CX 

CO 

a, 

a 

tj 

o 

ex 

ex 

CX 

ft 

•r-l 

PS     r-l     r-l     [Z; 


P 

T) 

0 

'i. 

p 

•r-l 

=\ 

CO 

4-1 
CO 

U 

T- 

CI) 

CO 

4_) 

4J 

c 

C 

= 

CI) 

CI' 

O 

1— 1 

M 

CJ 

er 

a 

c 

- 

h 

r4 

0 

0 

c 

a 

a 

P 

P 

p 

o 

0 

CJ 

n 

o 

0 

o 

0 

o 

CJ 

~ 

•r-l 

•H 

•H 

C4-I 

r-l 

4J 

4-1 

4-1 

4-1 

4-J 

n 

CO 

C) 

CJ 

CJ 

CJ 

CJ 

CJ 

cu 

CO 

c 

- 

CO 

ctl 

CO 

CO 

CO 

CO 

en 

> 

B 

•r-l 

~ 

cu 

0) 

CU 

cu 

CU 

(!) 

O 

P 

W) 

n 

)-l 

H 

rJ 

r-l 

U 

T3 

> 

r-l 

CZ) 

u 

i. 

0 

o 

o 

O 

0 

0 

O 

3 

o 

^ 

(5 

y 

13 

y 

JS 

r3 

y 
o 

co 

CD 

B 

•H 
4-J 

en 

■rl 

X 

C4-I 

o 

P 

o 

4-J 

CO 

c 
•- 

rJ 

>. 

4J 

Cu 

4-1 

4-1 

P 

•r-l 

4-J 

CO 

Cr 

X) 

X 

X 

X 

X) 

X 

o 

r— 1 

CO 

CO 

B 

x 

a 

ex 

CU 

Cu 

Cu 

Cu 

CJ 

CO 

Cu 

ex 

a, 

Cu 

CX 

CX 

v~' 

4J 

5-i 

c 

O 

TJ 

0 

e 

<o 

CN 

O 

x> 

r-j 

o 

o 

O 

•r-l 

O 

? 

CN 

m 

o 

CN 

m 

o 

p 

>: 

4-1 

c 

"53 

o 

o 

r-l 

o 

o 

H 

0 

y 

ii 

CO 

rJ 

60 

c 

"5 

rH  cn  cn  <j-  in  MD  r- 


=  2 


0 

o 

0 

0 

o 

o 

o 

o 

cc 

•^ 

"S 

y 

y 

y 

r3 

IS 

12 

y 

4-J 

O 

y 

s 

Tl 

T) 

nd 

Tl 

Tl 

Tl 

Tl 

c 

r. 

— 

p 

p 

c 

p 

p 

p 

p 

<D 

5 

o 

o 

0 

o 

o 

0 

o 

•> 

C/l 

s 

p-i 

(X 

PU| 

Pm 

Ph 

Pu 

Pu 

X) 

CD 

CO 

CU 
X) 

rJ 

ex 

i~ 

> 

Hfl 

^ 

CO 

(3 

r- 

-s 

> 

o 

C5 

rJ 

4-1 

•rl 

P 

P 

cn 

3 

o 

CO 
J-i 

B 

X 

- 
- 

o 

II 

& 

P 

0) 

1 

o    o     fe 


Ecology  151 

day)  each  day  from  Monday  through  Saturday.  Water  temperatures 
were  not  controlled  during  these  tests  and  ranged  between  63°  F  and 
75°  F. 

Results  i 

Concentrations  of  0.26  ppb  and  0.52  ppb  were  run  concurrently  for 
a  period  of  96  hours,  and  at  neither  level  were  there  any  mortalities. 
There  were  no  indications  of  even  slight  distress  in  any  fish.  These  fish 
were  observed  for  several  weeks  after  completion  of  the  96-hour  test, 
and  no  reaction  to  the  chemical  was  observed.  The  next  dosage  tested 
was  1.0  ppb.  During  a  48-hour  test  no  mortalities  occurred  and  the  fish 
showed  no  observable  evidence  of  discomfort.  At  the  termination  of 
the  48-hour  test,  another  1.0  ppb  was  added  to  Aquarium  3,  and  the 
concentration  in  Aquarium  6  was  augmented  by  2.0  ppb.  Assuming  that 
the  original  1.0  ppb  dosage  had  not  been  appreciably  dissipated  during 
the  previous  48  hours,  concentrations  at  this  time  were  upwards  to  2.0 
ppb  in  Aquarium  3  and  3.0  ppb  in  Aquarium  6.  After  18  hours,  the  first 
indication  of  distress  was  noted.  A  small  bluegill  in  Aquarium  6  showed 
the  initial  signs  of  disquietude.  Symptoms  of  distress  in  this  fish  became 
more  intense  at  each  30-min  observation  period,  and  90  min  after  the 
first  manifestations  the  fish  was  dead.  Soon  afterward,  other  fish  in 
both  aquaria  began  to  show  reactions  to  the  test  solution,  and  42  hours 
after  increasing  the  concentration  all  fish  in  both  aquaria  were  dead. 

The  physiological  reactions  were  similar  to  those  described  by 
Public  Health  Service  biologists  (14).  An  increase  in  respiration  and  fin 
movements  was  first  noted.  Later,  fish  exhibited  annoyed  wanderings  to 
the  surface  or  into  bottom  corners  of  aquaria.  As  distress  became  more 
intense,  darting  movements  were  noted,  with  fish  occasionally  breaking 
the  surface  and  in  one  instance  jumping  clear  of  the  aquarium.  Later,  a 
loss  of  hydrostatic  equilibrium  was  evidenced  by  fish  swimming  abruptly 
to  the  surface  and  settling  slowly  to  the  bottom.  At  this  time,  the  test 
animals  were  oriented  at  an  angle  to  the  horizontal.  The  body  axis  was 
a  line  diverging  from  the  horizontal  by  20° -35°,  caudal  end  down.  Fish 
near  expiration  showed  little  control  of  their  movements;  some  were 
attempting  feeble  swimming  efforts  while  on  their  sides  or  backs. 

These  results  indicated  that  the  toxicity  threshold  of  endrin  lay 
between  1.0  ppb  and  1.52  ppb  in  these  experimental  waters  and  with 
these  test  animals.  However,  since  the  first  dosage  of  1.0  ppb  was  fol- 
lowed for  only  48  hours,  there  was  a  possibility  that  a  more  extended 
test  period  would  have  resulted  in  some  mortality.  Another  test  at  this 
level  (1.0  ppb)  was  run  for  72  hours  but  no  mortalities  resulted.  Again, 
1.0  ppb  was  added  to  each  aquarium  and  once  more  no  fish  survived  an 
additional  72-hour  test.  From  these  exploratory  tests,  it  was  assumed 
that  the  toxicity  threshold  was  a  value  between  1.0  ppb  and  2.0  ppb, 
indicating  that  the  field  tests  should  begin  with  a  concentration  of  1.0 
ppb  (apparently  a  sublethal  dose)  and  continue  upward  until  concentra- 
tion at  which  all  test  fish  were  killed  was  reached  (Lethal  Concentration, 
100%,  or  simply  LC100).    It  should  be  noted  here  that  where  a  chemical's 


152  Indiana  Academy  of  Science 

use  as  a  piscicide  is  concerned,  the  LC100  becomes  a  more  important  figure 
than  the  more  commonly  used  LC50  (also  commonly  termed  LD50  Lethal 
Dosage,  50%  or  tlih).  This  stems  from  the  fact  that  many  chemicals 
show  a  non-linear  regression  from  the  concentration  producing  total 
extermination  to  those  of  sublethal  levels  (LC10ft  to  LC0).  Thus,  where  the 
economic  feasibility  of  a  product  is  one  of  the  important  criteria,  it  be- 
comes of  greater  concern  to  establish  this  level  by  direct  testing  rather 
than  to  estimate  it  by  projection  from  intermediate  levels  (lc50). 

Later  checks  of  the  2.0  ppb  solutions  were  made  to  determine  if  they 
were  still  toxic.  Two  fish  were  placed  in  each  of  2  aquaria  15  days  after 
the  solutions  were  made.  All  fish  failed  to  survive  until  the  19th  day  (96- 
hour  test)  indicating  that  dissipation  had  not  progressed  to  any  great 
degree. 

An  interesting  sidelight  which  supports  results  of  a  Japanese  study 
(19)  was  the  presence  of  cladocerans  and  planarians  in  the  3.0  ppb 
solution.  Two  days  after  all  fish  had  been  killed,  these  organisms  were 
noted  in  large  numbers  on  the  sides  of  the  aquarium. 

Crosley  Pond  Tests 

During  1960,  the  endrin  study  suffered  from  a  low  priority,  and  there 
was  no  time  available  for  this  project  until  mid-October.  On  October  13, 
a  small  pond  on  the  Crosley  State  Fish  and  Game  Area  was  treated 
with  endrin  calculated  to  produce  a  concentration  of  1.0  ppb.  After  120 
hours  without  any  mortality  of  test  fish,  another  1.0  ppb  of  endrin  was 
added.  This  presumably  raised  the  endrin  concentration  in  the  pond  to 
2.0  ppb,  a  level  that  had  killed  all  test  fish  in  laboratory  experiments. 

After  144  hours  of  testing  at  this  level,  there  were  no  mortalities, 
nor  even  discomfort  among  test  fish.  It  was  theorized  that  this  puzzling 
situation  resulted  from  lower  temperatures  and /or  higher  turbidities  than 
had  been  present  in  laboratory  tests.  This  study  was  not  continued  dur- 
ing 1961. 

Driftwood  Pond  Tests 

In  1962  the  study  was  given  a  higher  priority  and  the  testing  site 
was  moved  to  the  Driftwood  Farm  Pond  Experiment  Station  where  four 
ponds  were  available.  It  was  felt  that  the  common  water  source,  depth, 
basin  morphometry,  and  soil  types,  and  similar  thermal  gradients  would 
eliminate  some  of  the  variables  encountered  in  the  heterolimnetic  en- 
virons of  dissimilar  farm  ponds. 

Three  ponds  were  used  for  testing  and  the  fourth  served  as  a  con- 
trol. The  study  here  was  designed  to  bracket  the  minimum  LC100  with  a 
concentration  below  the  LC100  (at  1  ppb)  and  one  calculated  to  give  a 
total  kill  (4  ppb).  These  figures  were  based  on  the  preliminary  lab  tests 
which  showed  LC50  of  1  ppb  or  more,  and  no  mortality  in  the  Crosley 
pond  at  an  assumed  2  ppb  level.  It  was  reasoned  that  4  ppb  in  waters 
of  higher  temperature  would  effect  a  complete  kill.  A  third  pond  was 
treated  with  a  tremendous   dose  to   observe   the   duration   of  toxicity  in 


Ecology  153 

cases  of  miscalculated  dosages.  Treatment  in  this  pond  was  at  46  ppb. 
No  treatment  was  made  on  the  control  pond.  Ponds  ranged  in  size  from 
0.62  acres  to  1.37  acres.  Actual  testing  began  in  mid-September  when 
water  temperatures  were  near  16°  C. 

Pond  1 

Pond  1  (0.62  acres)  received  the  initial  treatment.  This  pond  was 
sprayed  with  a  solution  calculated  to  give  a  4  ppb  concentration  of 
endrin  (active  ingredient).  Twenty-four  hours  previous  to  the  spraying 
of  the  toxicant,  2  test  cages  each  containing  10  bluegills  were  placed  in 
the  pond.  One  cage  was  placed  in  water  of  18-inch  depth,  the  other  in 
5  feet  of  water.  Test  cages  were  approximately  36"  X  36"  X  22".  Since 
bluegills  used  as  test  fish  were  from  adjacent  ponds,  the  24-hour  acclima- 
tion period  seemed  sufficient. 


Il|lllijililt« 


&00*       ' 


v.      ~a      . 


ill 


mm 


Figure  2.    One  of  the  endrin  test  ponds  at  the  Driftwood,  Station  showing  deepwatt 
cage  raised  for  examination.  Dead  test  fish  are  visible  in  cage. 


Test  cages  were  checked  periodically  to  observe  fish  reactions,  but 
for  the  purpose  of  recording,  data  were  considered  only  on  a  24-hour 
interval  basis  (i.e.,  fish  were  recorded  as  having  died  only  at  the  termi- 
nation of  a  24-hour  test  period).  Although  the  last  fish  in  a  test  group 
may  actually  have  expired  at  42  hours,  the  LC100  was  recorded  as  having 
occurred  at  the  48-hour  level.  (There  seems  to  be  little  justification  for 
carrying  the  recording  of  results  to  a  precision  beyond  the  sophistication 
of  the  basic  technique  itself.) 

After  24  hours,  2  fish  in  the  deep  water  cage  had  died,  but  there  were 
no  mortalities  in  the  shallow  water  cage  although  2  bluegills  were  in 
extreme  distress.  At  the  48-hour  check,  all  fish  in  the  deep  cage  were 
dead  and  there  was  but  one  survivor  in  the  shallow  cage.  The  following 
day  (72-hour  test)  the  last  fish  had  expired. 


154  Indiana  Academy  of  Science 


Pond  2 


The  same  procedure  was  followed  in  Pond  2  testings  as  had  been 
used  in  Pond  1  (a  deep  and  a  shallow  cage  each  with  10  test  fish).  This 
pond  (1.07  acres)  was  sprayed  with  a  dosage  calculated  to  give  a 
concentration  of  1  ppb  of  endrin  (active  ingredient).  After  introduction 
of  the  toxicant,  no  mortalities  were  recorded  during  the  first  4  days  (the 
24-,  48-,  72-,  and  96-hour  tests),  nor  was  any  distress  noted  in  any  of 
the  test  fish  during  this  time.  In  view  of  subsequent  results,  the  signifi- 
cance of  this  lack  of  mortality  during  the  96-hour  period  will  be  discussed 
in  some  detail  later. 

On  the  fifth  day,  one  bluegill  showed  the  first  signs  of  agitation 
and  symptoms  of  distress  became  more  intense.  Before  the  end  of  the 
120-hour  test,  this  fish  had  become  the  first  mortality  in  Pond  2.  The 
following  day  another  fish  died  (144-hour  test)  and  sporadic  mortalities 
continued  until  the  last  fish  expired  3  weeks  after  introduction  of  the 
toxicant  (504-hour  test).  These  events,  although  instructive,  were  some- 
what disconcerting  since  this  level  (projected  from  aquaria  and  Crosley 
Pond  tests)  was  intended  to  be  a  sublethal  concentration.  It  seemed  un- 
tenable to  suspect  disease,  parasitism,  debility  from  feeding  inter- 
ference, etc.  as  the  causative  agent  of  the  mortalities.  Observations 
showed  no  external  parasites  or  disease  and  body  forms  of  the  dead 
fish  displayed  no  noticeable  emaciation.  In  addition,  during  this  period, 
the  control  fish  in  an  adjacent  pond  had  suffered  no  mortalities  under 
very  similar,  if  not  identical,  conditions — with  the  exception  of  the  test 
toxicant,  of  course.  I  was  forced  to  the  conclusion  that  the  LC100  concen- 
tration in  these  Driftwood  waters  was  somewhat  lower  than  it  had  been 
in  the  Crosley  pond  waters.  The  major  differences  in  the  ponds  were 
size,  shading,  temperature,  and  transparency.  The  Driftwood  ponds  were 
larger,  unshaded,  warmer,  and  had  greater  clarity.  All  of  these  physical 
factors  have  been  suspect  in  influencing  the  toxicity  of  various  chlori- 
nated hydrocarbons  in  solution.  These  will  be  discussed  later  at  some 
length. 

Pond  4 

Pond  4  was  a  1.23-acre  pond  used  for  testing  the  residual  effect 
of  an  extremely  high  dosage.  This  pond  was  sprayed  with  a  quantity  of 
endrin  calulated  to  give  a  concentration  of  46  ppb.  The  same  procedure 
(two  cages  with  10  bluegills  each)  that  had  been  used  in  Ponds  1  and  2 
was  used  in  Pond  4. 

At  the  termination  of  the  24-hour  test  (8:00  AM,  September  18, 
1961)  all  fish  were  dead.  Until  the  pond  froze  over,  test  fish  (usually 
at  2-week  intervals)  were  placed  in  the  cages,  and  each  time  died  within 
the  24-hour  period.  After  ice  cover  sealed  the  pond,  no  tests  were  made 
until  a  warmer  period  opened  it  in  January.  At  this  time,  test  fish  again 
died  within  a  24-hour  period.  It  was  decided  to  replace  the  water  and 
see  if  the  residual  endrin  in  the  sediments  would  produce  a  toxicity  in 
previously-uncontaminated  water. 

The  pond  was  drained  February  1-3,  1962,  and  allowed  to  stand  idle 
for    about    3    weeks.    Then    it   was    refilled    with    uncontaminated    water 


Ecology  155 

(March  1)  and  allowed  to  stand  until  March  13.  On  this  date,  a  new 
group  of  10  bluegills  was  placed  in  a  single  cage  in  the  pond.  Two  of 
these  subsequently  escaped  from  encagement  and  one  was  later  found 
dead  along  the  pond  edge.  Of  the  eight  fish  remaining  in  the  test  cage, 
the  first  mortalities  occurred  on  March  26.  At  this  time,  four  fish  were 
found  dead.  One  more  bluegill  was  dead  on  March  28,  another  on  March 
29,  and  the  final  two  expired  on  April  1. 

The  pond  was  then  drained  a  second  time,  and  refilled  on  April  13. 
Nine  test  fish  were  put  in  the  cage  on  April  25.  The  first  mortality  was 
recorded  on  May  2  and  by  May  9  all  remaining  test  fish  were  dead.  The 
pond  was  immediately  drained  again  and  had  been  refilled  on  May  15. 
Ten  new  fish  were  placed  in  the  pond  on  May  20,  and  on  June  2  the 
first  death  was  recorded.  Another  died  on  June  5,  and  two  more  on 
June  7.  Although  five  fish  were  still  alive,  it  was  obvious  that  a  long- 
term  lethal  toxicity  still  existed. 

Once  more  the  pond  was  drained  and  refilled,  and  was  ready  for 
the  continuation  of  testing  by  June  15.  Fish  again  were  put  in  the  cage, 
and  these  fish  showed  no  mortality  in  a  subsequent  month  of  observa- 
tions. Tests  were  suspended  at  this  time  since  the  pond  appeared  to 
have  become  non-toxic  and  was  needed  for  another  study.  Apparently, 
the  dissipation  of  the  toxicant  was  finally  complete  since  the  other  fish 
stocked  in  this  test  pond  were  not  affected. 

It  should  be  stressed  that  the  water  source  for  these  ponds  was 
piped  by  gravity  flow  from  adjacent  Starve  Hollow  Lake  which  is  lo- 
cated upstream  from  the  station.  Ponds  drained  into  a  small  cresk 
which  also  drained  the  lake.  The  source  water  was  entirely  uncontami- 
nated  by  endrin  and  no  other  insecticides  were  known  to  have  been  used 
in  the  lake  itself.  In  addition,  no  fish  kills  were  noted  in  the  lake.  Thus 
it  appears  that  the  toxicity  was  of  an  autochthonous  nature,  and  is  as- 
sumed to  have  been  recycled  or  brought  into  re-solution  from  the  con- 
taminated sediments. 


Discussion 

Only  in  1962  were  the  endrin  tests  given  precedence  over  other 
duties.  By  1962  it  was  becoming  apparent  from  personal  experiences  and 
those  of  other  investigators  that  this  chemical  was  too  dangerous  for 
use  as  a  piscicide  except  in  extremely  isolated  instances. 

As  data  accumulated  from  various  studies  of  endrin,  certain  prob- 
lems became  apparent.  It  has  been  noted  that  other  chlorinated  hydro- 
carbons were  present  in  fish  flesh  at  higher  levels  than  were  present  in 
solution  (6,  7,  8,  18,  20).  This  accumulative  effect  is  indicated  for 
endrin  also  in  Bridges'  (3)  study  of  a  Colorado  fish  kill,  and  by  implica- 
tion (22)  in  aquaria  tests.  The  concentrating  potential  of  lower  forms 
in  the  food  chain  is  well  documented  for  hydrocarbons:  moss  (15); 
Potamogeton  (8);  vegetation  (3,  4);  earthworms  (2);  and  plankton, 
frogs  and  fish  (18).  It  is  generally  assumed  that  these  accumulations  in 


156  Indiana  Academy  of  Science 

fish  flesh  are  dietary  translocations  of  residues  through  ingestion  of 
contaminated  food  items.  The  role  of  direct  absorption  of  the  materials 
from  the  aquatic  medium  through  integumentary  tissues  has  not  been 
as  convincingly  demonstrated,  although  there  are  data  (22)  that  imply 
direct  absorption  of  endrin.  Hoss  (17)  has  effectively  shown  that  both 
pathways  (absorption  and  ingestion)  may  be  important  in  his  studies 
of  zinc65  accumulations  in  the  flounder,  and  Williams  and  Pickering  (28) 
have  shown  that  bluegills  accumulate  cesium1-^  and  strontium^  by  both 
ingestion  and  absorption.  Though  not  conclusive,  these  data  are  insinua- 
tive. 

Regardless  of  whether  the  accumulative  deposition  in  fish  flesh  re- 
sults from  ingestion  or  from  absorption,  it  forces  us  to  re-evaluate  our 
reliance  on  short  term  bioassays  (24-  to  96-hour  tests)  where  chronic 
exposures  to  chlorinated  hydrocarbons  are  involved.  One  of  the  two 
major  points  arising  from  this  present  study  is  that  short  term  bio- 
assays are  insecure  evidence  of  non-toxic  levels  in  situations  of  chronic 
or  long-persisting  contamination  and/or  with  materials  accumulatively 
stored  by  metabolic  processes.  The  need  for  studies  of  the  effects  of 
long  term  exposure  has  been  pointed  out  (14),  but  there  is  a  rather 
dramatic  example  in  the  study  of  Driftwood  Pond  2.  Had  testing  in 
this  pond  been  suspended  after  a  common  48-hour,  72-hour,  or  even 
96-hour  period,  the  assumption  of  a  sublethal  level  would  have  ap- 
peared valid.  However,  the  first  mortality  occurred  on  the  5th  day  and 
all  test  fish  had  expired  within  3  weeks  from  time  of  exposure.  There 
were  no  mortalities  among  the  control  fish  during  this  period.  There- 
fore, one  must  carefully  examine  over  extended  periods  those  agents 
of  an  accumulative  or  an  insidious  nature  to  ascertain  that  seeming  sub- 
lethality  is  not  actually  slow  response.  This  is  particularly  true  as 
testing  approaches  the  toxicity  threshold. 

The  second  point  of  interest  deduced  from  this  and  other  studies 
involves  the  variance  in  toxic  limits  for  fish  established  by  several 
workers  (Table  1).  Katz  and  Chadwick  (22)  obtained  a  96-hour  LC50 
(their  TLm)  of  0.27  ppb  for  coho  salmon,  and  0.60  ppb  for  bluegills. 
Henderson  et  al.  (14)  established  a  96-hour  LC5o  (their  TLm)  of  0.44  ppb 
for  bluegills  in  hard  water  using  the  emulsible  concentrate.  In  the  present 
study,  I  failed  to  get  any  kill  on  test  fish  in  the  Crosley  test  at  an 
assumed  2  ppb.  On  the  other  hand,  there  was  a  complete  kill  of  test  fish 
over  a  3-week  period  in  one  Driftwood  pond  at  a  1  ppb  concentration.  A 
resume  of  Michigan  lake  and  stream  rehabilitations  (16)  notes  a  sur- 
vival of  some  fish  (Fundulus)  at  8  ppb.  Bridges  (3)  points  to  a  partial 
survival  of  largemouth  bass,  bluegills,  pumpkinseed  sunfish,  and  black 
crappies  in  a  Colorado  pond  adjacent  to  a  beet  field  which  had  been 
sprayed  with  an  emulsible  formulation  of  endrin.  At  the  time  of  investi- 
gation (4  days  after  the  spraying),  water  analyses  indicated  a  concen- 
tration of  40  ppb  of  endrin.  Fish  were  not  dying  at  this  time.  Bridges, 
noting  the  disparity  of  his  results  with  those  of  the  USPHS  team,  points 
a  suspicious  finger  at  the  9.1  pH  of  the  Colorado  pond  and  the  fact  that 
the  highest  pH  tested  by  Henderson's  group  (14)  was  8.2. 


Ecology 


157 


Temperature  has  been  shown  to  exert  a  tremendous  effect  on  the 
toxicity  of  endrin.  The  studies  of  a  Japanese  group  (19)  observed  that 
endrin  toxicity  to  fish  increased  at  higher  temperatures,  and  Katz  and 
Chadwick  (22)  found  96-hour  LC50  levels  for  bluegills  of  8.25  ppb  at 
1.0°-4.5°C  and  0.33  ppb  at  25 °C.  The  data  of  the  latter  study  indicate  a 
25-fold  increase  in  toxicity  with  an  increase  of  20.5°-24.0°C. 

There  are  wide  differences  between  the  toxic  thresholds  of  endrin, 
DDT  and  dieldrin  in  diverse  animal  groups  (1,  5,  9,  10,  13,  14).  An 
instance  of  this  is  obvious  in  Table  1.  Species  susceptibility  to  endrin  has 
been  pointed  out  by  numerous  investigators  (14,  21,  22,  25,  27).  Even 
within  a  particular  species  there  is  evidence  that  susceptibility  to  endrin 
changes  with  age,  with  embryonic  stages  being  more  resistant  than  larval 
stages  (19). 


Table  1.      The     varying     toxicity     levels     of     three     chlorinated     hydrocarbons     in     their 

effect     on     several     classes     of     organisms     (measured     in     mg/kg     of     body     weight     or 

in    ppb    of    active    ingredient) . 


Organism 

DDT 

Dieldi 

in 

Endrin 

Estimated  fatal  dose  for  a 

30   g    (5,    13) 

(  =    an 

5  g    (13)    (  = 

an  esti- 

? 

man  weighing  70  kg 

estimated  428 

mg/kg) 

mated  71  mg 

/kg) 

Pheasants  (LD50) 

300  mg/kg 

50  mg/kg 

14  mg/kg 

(9,  10) 

9,  10) 

(9,  10) 

Bluegills  (LC50  for 

8.8  ppb  (14) 

9.1  ppb  (14) 

0.44  ppb 

emulsible  concentrate) 

(14) 

Daphnia   (50-hour 

1.4  ppb  (1) 

330  ppb  (1) 

352  ppb 

Immobilization  test) 

(1) 

It  has  been  pointed  out  that  the  tolerance  of  some  species  is  different 
in  different  volumes  of  water  even  though  the  concentration  of  the  active 
ingredient  is  the  same.  This  so-called  volume  effect  has  been  noted  for 
DDT  (24)  and  for  endrin  (22).  This  increase  in  toxicity  in  greater  vol- 
umes of  the  same  concentration  as  smaller  volumes  can  be  logically  cor- 
related with  the  previously  discussed  accumulative  nature  of  this  hydro- 
carbon. In  addition  to  fish  as  accumulators  of  pesticidal  hydrocarbons, 
studies  have  indicated  the  importance  of  macrophytes  as  concentrators 
of  these  chemically  related  insecticides  (4,  8).  Bridges  and  his  co-workers 
(4),  by  use  of  isotope-labeled  DDT  (Cu-DDT),  have  found  that  storage 
of  this  relative  of  endrin  is  about  Vz  external  (i.e.,  adsorption  to  external 
portions  of  the  plant)  and  %  internal  (absorption  into  the  tissues).  This 
could  be  presumably  true  for  endrin  also.  If  such  a  presumption  has  any 
basis  in  actuality,  it  is  quite  conceivable  that  in  aquatic  ecosystems, 
where  macrophytes  form  an  important  part  of  the  biocoenosis,  with- 
drawal of  hydrocarbons  by  these  plants  would  have  a  depressing  effect  on 
the  toxicity.  The  result  would  be  a  balancing  or  a  suppression  of  the 
volume    effect,    if    one    assumes    the    plants'    retention    of    endrin    until 


158  Indiana  Academy  of  Science 

degradation  of  the  chemical.  The  volume  effect  has  been  reported  for 
aquaria  studies,  and  seldom,  if  ever,  in  natural  waters.  Similarly,  lab- 
oratory studies  have  generally  established  lower  toxicity  levels  than  field 
studies,  sometimes  with  great  disparities.  My  experience  with  1  ppb  in 
Driftwood  Pond  2  is  a  notable  exception,  but  as  previously  pointed  out 
this  would  have  been  considered  an  lc„  at  the  96-hour  level. 

Fish  bioassayists  have  noted  different  toxicity  levels  produced  by 
various  formulations  of  the  same  chemical  compound.  These  are  most 
graphically  depicted  by  Henderson,  et  al.  (14).  They  compared  96-hour 
LCsos  for  bluegills  when  tested  in  2  formulations  (acetone  solution  and 
emulsible  concentrate)  of  a  number  of  insecticides.  DDT  was  considerably 
more  toxic  in  emulsible  concentrate,  dieldrin  was  less  toxic  in  this  form, 
and  endrin  showed  little  difference.  From  other  experiments,  they  con- 
clude that  the  toxicity  of  wettable  powders  (used  in  the  present  study) 
was  similar  to  that  of  the  acetone  solution.  Other  chemical  and  physical 
factors  such  as  the  effects  of  light,  dissolved  mineral  content,  suspended 
particulate  matter,  etc.,  are  not  well-understood  and  could  have  an 
important  or  contributing  influence. 

In  spite  of  all  these  possible  contributory  influences,  the  distribution 
of  variously-established  toxic  thresholds  seems  implausibly  disparate. 
From  these  doubts  is  born  a  second — or  at  least,  an  accessory — explana- 
tion. This  is  simply  that  a  lack  of  standardization  of  analytical  techniques 
exists.  Extraction  and  analysis  of  almost  infinitesimal  hydrocarbon  resi- 
dues in  tissues  has  been  a  tedious,  painstaking  job  with  interfering  sub- 
stances having  unknown  effects  on  the  results.  It  appears  that  the  more 
exacting  gas  chromatography-infra-red  spectrophotometric  method  now 
in  use  will  narrow  the  divergence  of  results.  However,  even  this  advance 
did  not  produce  incontrovertible  evidence  in  the  lower  Mississippi  River 
incidents. 

I  suspect  that  higher  temperatures  and  water  transparency  were 
responsible  for  the  total  kill  in  Driftwood  Pond  2  at  a  level  which,  in 
laboratory  tests,  was  sublethal.  Conversely,  the  failure  to  get  a  kill  in 
the  Crosley  Pond  at  a  high  dosage  (2  ppb)  is  attributed  to  the  high 
turbidity  in  this  pond,  with  a  possible  assist  from  the  decreasing  water 
temperatures. 

Different  investigators  have  used  various  designations  for  their 
toxicity  measurements  (ld,  tl,  and  lc).  Lethal  dosage  (ld)  has  been 
used  for  some  time  as  the  index  of  relative  toxicity  to  terrestrial  or 
avian  animals,  and  the  method  generally  has  used  milligrams  of  a  sub- 
stance per  kilogram  of  body  weight  as  its  comparative  measurement.  The 
Subcommittee  on  Toxicity,  of  the  Federation  of  Sewage  and  Industrial 
Wastes  Associations  (12)  recommended  the  term  Median  Tolerance  Limit 
(tl,,,)  to  express  the  concentration  at  which  just  50%  of  the  test  animals 
die.  This  designation  was  aimed  at  measurements  of  chemicals  in  a 
liquid  medium  with  fish  as  the  test  animals.  It  is  usually  derived  by  a 
straight  line  graphical  interpolation  from  tests  producing  higher  and 
lower  percentages  of  mortality.  More  recently,  several  investigators  have 


Ecology  159 

shown  an  increasing  preference  for  Lethal  Concentration  (lc).  Usually  LD 
and  lc  tests,  like  tl„,,  have  attempted  to  establish  the  50%  level   (ld51„ 

lc5o). 

For  a  piscicide,  as  briefly  mentioned  previously,  the  minimum  con- 
centration which  produces  100%  mortality  of  the  fish  population  (LC100) 
constitutes  a  more  important  figure  than  the  LCr.o.  Desirably,  for  field 
use  as  a  fish  toxicant,  a  material  should  be  effective  in  eradicating  the 
population  within  a  specific,  preferably  short,  time  (LC100  during  x  time 
interval).  It  is  obvious  then  that  the  LC100  is  not  a  precise  unity  but 
rather  a  fluctuating  value  between  time  parameters  (e.g.,  the  24-hour 
LC100  may  be  4  ppb  whereas  the  LC100  for  a  96-hour  test  may  be  2  ppb). 

Lennon  and  Walker  (23)  have  used  the  term  Effective  Concentra- 
tion (EC100)  in  their  delineative  screening  program.  If  we  define  an  effec- 
tive concentration  (EC)  as  the  minimum  concentration  that  produces  a 
100%  mortality  to  problem  species,  this  can  be  written  without  the  mor- 
tality-percentage subscript  numeral.  This  would  allow  contraction  of  the 
abbreviate  symbols  by  inclusion  of  the  time  interval  as  a  superscript 
numeral  as  has  Dorris  et  al.  (11)  in  refinery  waste  bioassays  (tl„,  48  = 
tl,„  for  48-hour  test).  This  would  avoid  complicating  the  formula 
with  both  a  subscript  and  a  superscript  (LC10048),  and  it  would  be  simpler 
to  use.  While  EC  (or  LC100)  is  a  varying  quantity  with  time-influenced 
parameters,  the  EC34,  the  EC4S,  etc.,  are  precise  figures  for  particular  test 
situations. 


Recommendations 

Among  a  number  of  desirable  characteristics  of  a  fish  toxicant  (low 
cost,  easy  application,  homogeneous  dispersal,  relatively  persistent  tox- 
icity, etc.),  the  mandatory  nature  of  one  trait  makes  it  the  primary  con- 
sideration. This  is  its  effect  on  human  health.  It  must  first  meet  regula- 
tory standards  set  up  to  guard  the  public  (against  itself?).  At  the  time 
these  tests  were  initiated,  endrin  was  merely  the  latest  and  most  potent 
of  an  exponentially-increasing  number  of  chlorinated  hydrocarbon  in- 
secticides. Its  entry  into  the  commercial  market  was  accompanied  by  the 
usual — at  that  time — incomplete  testing  of  its  biological  field  effects. 
(Erstwhile  tests  by  chemical  manufacturers  might  be  paraphrased  as  the 
development  of  broad  spectrum  pesticides  with  narrow  spectrum 
evaluations.) 

It  soon  became  apparent  that  endrin  (for  fish,  at  least)  was  the 
most  lethiferous  substance  to  come  out  of  the  chemists'  cauldron.  When 
certain  doubts  concerning  its  effect  on  human  health  arose,  its  status 
became  immediately  suspect,  in  spite  of  its  use  in  harvesting  food  fish 
in  Malaysia  (26).  All  the  ramifications  of  its  use  are  still  not  under- 
stood, and  it  appears  possible  that  authorities  may  belatedly  prohibit  its 
sale.  In  view  of  its  possible  health  hazards  and  the  probability  of  its 
imminent  withdrawal  from  the  commercial  market,  it  seems  that  health 
considerations  and  unavailability  have  negated  whatever  potential  endrin 
may  have  had  as  a  fish  toxicant.  One  cannot  with  any  integrity  abhor 


160  Indiana  Academy  of  Science 

the  fallout  from  atomic  testings,  deplore  the  dispersal  of  heptachlor  to 
kill  some  insignificant  bug,  and  then  put  on  the  Hydean  mask  to  espouse 
the  widespread  dissimination  of  endrin  in  our  waters. 


Literature  Cited 

1.  Anderson,  Bertil  G.  1960.  The  toxicity  of  organic  insecticides  to  Daphnia.  Robert  A. 
Taft  Sanitary  Eng.  Center,  U.  S.  Public  Health  Serv.  Tech.  Rep.  W60-3.  p.  94-95. 

2.  Barker,  Roy  J.  1958.  Notes  on  some  ecological  effects  of  DDT  sprayed  on  elms. 
J.  Wildl.  Mgt.  22  :269-274. 

3.  Bridges,  W.  R.  1961.  Disappearance  of  endrin  from  fish  and  other  materials  of  a 
pond  environment.  Trans.  Amer.  Fish.  Soc.  90  :332-335. 

4.  Bridges,  W.  R.,  B.  J.  Kallman,  and  A.  K.  Andrews.  1963.  Persistence  of  DDT  and 
its  metabolites  in  a  farm  pond.  Trans.  Amer.  Fish.  Soc.  92:421-427. 

5.  Brown,  A.  W.  A.  1951.  Insect  control  by  chemicals.  John  Wiley  and  Sons,  Inc., 
New  York.  817  p. 

6.  Burdick,  G.  E.,  E.  J.  Harris,  H.  J.  Dean,  I.  M.  Walker,  Jack  Skea,  and  David 
Colby.  1964.  The  accumulation  of  DDT  in  lake  trout  and  the  effect  on  reproduction. 
Trans.  Amer.  Fish.  Soc.  93:127-136. 

7.  Cope,  Oliver  B.  1960.  The  retention  of  DDT  by  trout  and  whitefish.  Robert  A.  Taft 
Sanitary  Eng.  Center,  U.  S.  Public  Health  Serv.  Tech.  Rept.  W60-3.  p.  72-75. 

8.  Cope,  Oliver  B.  1961.  Effects  of  DDT  spraying  for  spruce  budworm  on  fish  in  the 
Yellowstone  River  system.  Trans.  Amer.  Fish.  Soc.  90  :239-251. 

9.  DeWitt,  J.  B.,  C.  M.  Menzie,  V.  A.  Adormaitis,  and  W.  L.  Reichel.  1960.  Pesticidal 
residues  in  animal  tissues.  Trans.  25th  N.  Amer.  Wildl.  and  Natur.  Resources  Conf. 
p.  277-285. 

10.  DeWitt,  J.  B.,  and  J.  L.  George.  1960.  Bureau  of  Sport  Fisheries  and  Wildlife 
Pesticide-Wildlife  Review:  1959.  Fish  and  Wildlife  Circular  No.  84,  Revised.  U.  S. 
Dept.  Interior,  Washington,  D.  C. 

11.  Dorris,  Troy  C,  William  Gould,  and  Charles  R.  Jenkins.  1960.  Toxicity  bioassay 
of  oil  refinery  effluents  in  Oklahoma.  In  Biological  Problems  in  Water  Pollution. 
Trans,  of  the  1959  Seminar  Eng.  Center,  Cincinnati,  Ohio.  p.  276-285. 

12.  Doudoroff,  P.,  B.  G.  Anderson,  G.  E.  Buddick,  P.  S.  Galtsoff,  W.  B.  Hart, 
R.  Patrick,  E.  R.  Strong,  E.  W.  Surber,  and  W.  M.  Van  Horn.  1951.  Bioassay 
methods  for  the  evaluation  of  acute  toxicity  of  industrial  wastes  to  fish.  Sewage  and 
Ind.  Wastes  23:1380-1397. 

13.  DuBois,  K.  P.  1958.  Insecticides,  rodenticides,  herbicides,  household  hazards.  Post- 
grad. Med.  24  :278. 

14.  Henderson,  C,  Q.  H.  Pickering,  and  C.  M.  Tarzwell.  1959.  Relative  toxicity  of  ten 
chlorinated  hydrocarbon  insecticides  to  four  species  of  fish.  Trans.  Amer.  Fish.  Soc. 
88  :23-32. 

15.  Hoffman,  C.  H.,  and  A.  T.  Drooz.  1953.  Effects  of  a  C-47  airplane  application  of 
DDT  on  fish-pond  organisms  in  two  Pennsylvania  watersheds.  Amer.  Midland  Natur. 
50:172. 

16.  Hooper,  Frank  F.,  John  E.  Williams,  Mercer  Patriarche,  Fred  Kent,  and  James 
C.  Schneider.  1964.  Status  of  lake  and  stream  rehabilitation  in  the  United  States  and 
Canada  with  recommendations  for  Michigan  waters.  Mich.  Inst,  for  Fish.  Res.  Rep. 
1688.  56  p. 


Ecology  161 

17.  Hoss,  Donald  E.  1964.  Accumulation  of  zinc-65  by  flounder  of  the  genus  Paralichthys. 
Trans.  Amer.  Fish.  Soc.  93  :364-368. 

18.  Hunt,  Eldridge  G.,  and  Arthur  I.  Bischoff.  1960.  Inimical  effects  on  wildlife  of 
periodic  DDD  applications  to  Clear  Lake.  Calif.  Fish  and  Game  46  :91-106. 

19.  Iyotomi,  Kisabu,  Tomotsu  Tamura,  Yasno  Itazawa,  Isao  Hanyu,  and  Syotoro 
Sugiura.   1958.   Toxicity  of   Endrin  to   Fish.   Prog.   Fish   Cult.   20:155-162. 

20.  Kallman,  B.  J.,  O.  B.  Cope,  and  R.  J.  Navarre.  1962.  Distribution  and  detoxication 
of  toxaphene  in  Clayton  Lake,  New  Mexico.  Trans.  Amer.  Fish.  Soc.  91:14-22. 

21.  Katz,  Max.  1961.  Acute  toxicity  of  some  organic  insecticides  to  three  species  of 
salmoids  and  to  the  three-spine  stickleback.  Trans.  Amer.  Fish.  Soc.  90  :264-268. 

22.  Katz,  Max,  and  George  G.  Chadwick.  1961.  Toxicity  of  endrin  to  some  Pacific 
Northwest  fishes.  Trans.  Amer.  Fish.  Soc.  90  :394-397. 

23.  Lennon,  Robert  E.,  and  Charles  R.  Walker.  1964.  Investigations  in  fish  control : 
(1)  Laboratories  and  methods  for  screening  fish-control  chemicals.  U.  S.  Dept.  of  the 
Interior,  Bureau  of  Sport  Fisheries  and  Wildlife,  Circular  185.  15  p. 

24.  Prevost,  G.,  C.  Lannouette,  and  F.  Grenier.  1948.  Effect  of  volume  on  the  determi- 
nation of  DDT  or  rotonone  toxicity  on  fish.  J.  Wildl.  Mgt.  12  :241-250. 

25.  Rudd,  R.  L.,  and  R.  E.  Genelly.  1956.  Pesticides:  their  use  and  toxicity  in  relation 
to  wildlife.  Calif.  Dept.  Fish  and  Game,  Game  Bull.  No.  7.  209  p. 

26.  Soong,  MlN  Kong.  1960.  Shell  "Endrex"  used  as  a  fish  toxicant.  Prog.  Fish-Cult. 
22:93. 

27.  Screenivasan,  A.,  and  M.  V.  Natarajan.  1962.  Use  of  endrin  in  fishery  manage- 
ment. Prog.  Fish-Cult.  24:181. 

28.  Williams,  Louis  G.,  and  Quentin  Pickering.  1961.  Direct  and  food-chain  uptake  of 
cesium137   and   strontium*   in    bluegill    fingerlings.    Ecology   42  :205-206. 


Courtship  and  Territorial  Behavior  of  Some  Indiana  Woodcocks1 

Harmon  P.  Weeks,  Jr.,  Purdue  University 

Abstract 

Woodcocks  were  observed  during  evening  flight  performances  from  their  beginning 
on  March  15  until  conclusion  on  May  24  at  Shidler  Forest  in  western  Tippecanoe  County. 
Aspects  of  flight  performances  were  noted,  timed  when  possible,  and  recorded.  Territorial 
observations  were  recorded.  Three  birds  were  mist-netted  for  banding  and  identification 
purposes.  Taped  woodcock  calls  were  used  to  attract  males  for  netting  and  for  testing 
territorial  extent. 

The  greatest  number  of  performing  males  was  observed  during  the  last  half  of 
March.  This  was  probably  the  peak  migration  period.  A  few  birds  remained  on  the  study 
area  throughout  the  breeding  season.  Light  intensity  controls  the  onset  and  cessation 
of  performance  events.  Low  temperatures  and  precipitation  had  little  observed  effect  on 
performances.  Longer  flight  times  and  greater  ground  area  coverage  by  flights  were 
noted  in  this  study  than   in  previous  studies.  These  two  parameters  are  interrelated. 

With  the  aid  of  the  taped  woodcock  call,  a  woodcock  was  found  to  defend  an  area 
of  at  least  4.38  acres.  Most  territorial  defense  consisted  of  threats  and  retreats  with  no 
physical  contact  observed  between  combatants.  When  attracted  by  taped  calls,  one  bird 
did  attack  the  author  and  struck  him  several  times.  One  singing  site  appeared  most 
attractive  because  of  the  sequential  appearance  there  of  three  males  coinciding  exactly 
with  disappearances  of  males  from  other  sites. 


Introduction 

The  American  woodcock  (Philohela  minor)  is  an  uncommon  summer 
resident  in  Indiana,  but  because  they  commonly  nest  farther  to  the  north 
and  winter  to  the  south,  large  numbers  may  be  seen  here  during  fall  and 
spring  migrations  (10).  By  the  time  woodcocks  reach  this  area  in  the 
spring,  they  have  already  begun  their  courtship  and  this  performance 
may  be  seen  almost  anywhere  in  the  state  if  one  searches  the  correct 
habitats. 

The  courtship  performance  consists  of  a  rather  spectacular  spiraling 
flight  by  the  male  and  a  harsh,  nasal  call,  generally  described  as  a 
"peent"  (8),  which  is  given  from  the  ground  between  flights.  A  true  vocal 
song  is  given  in  the  flight  during  the  first  part  of  a  rather  direct  descent. 
The  performing  area  is  defended  against  other  males  by  the  territorial 
bird,  and  females  move  into  this  area  to  be  mated. 

Although  this  behavior  has  been  described  by  many  observers  (1,  8, 
9),  the  territorial  and  courtship  behavior  of  the  woodcock  are  poorly 
described  and  little  work  has  been  done  on  this  species  in  Indiana.  The 
objectives  of  this  study  were  to  attempt  to  answer  some  of  these 
questions  and  to  establish  some  performance  parameters  for  Indiana 
woodcocks. 

The  study  area  was  a  200-acre  tract  known  as  Shidler  Forest,  owned 
by  the  Department  of  Forestry  and  Conservation,  Purdue  University,  and 
located   10   miles   west  of  Lafayette,   Indiana,   in   Tippecanoe   County.    It 


Journal    Paper   No.    3876   from    Purdue   University   Agricultural    Experiment    Station. 

162 


Ecology 


163 


consists  of  bottomlands  along  Indian  Creek,  ridges,  and  ridgetops.  There 
are  openings  in  the  bottomlands  and  on  the  ridgetops  which  are 
used  as  woodcock  singing  fields.  The  area  has  a  small  resident  woodcock 

population. 

Methods  and  Materials 

Beginning  in  early  March,  the  open  fields  of  the  study  area  were 
checked  periodically  for  performing  birds.  When  birds  began  to  perform, 


LEGEND 
Field 

Woodland 
Stream 
Census  Route 

0  500 


1000 


Scale  (feet) 

Figure   1.     A    major    portion    of    Shidler    Forest    showing    performing    areas    which    were 
occupied  by  singing  woodcocks  at  times  during  1969. 


164  Indiana  Academy  of  Science 

a  route  was  established  to  include  visits  to  all  occupied  and  other  likely 
singing  sites  on  the  area.  These  sites  were  numbered  for  identification, 
incorporating  woodcock  territorial  numbers  of  previous  studies.  This 
accounts  for  the  non-sequential  numbering  (Fig.  1). 

This  route  was  walked  several  evenings  a  week,  and  a  record  was 
kept  of  sites  being  used  by  singing  woodcocks.  No  morning  censuses  were 
made.  When  a  site  was  in  use  on  two  consecutive  observations,  it  was 
assumed  to  have  been  used  during  the  period  between  those  visits. 
Records  were  kept  of  numbers  of  birds  on  and  off  territories,  behavioral 
observations,  and  timing  of  performance  events. 

On  some  days  all  field  time  was  spent  observing  complete  perform- 
ances of  individual  territorial  males.  The  chronological  order  and  duration 
of  performance  events,  the  number  of  flights  and  peents,  and  behavioral 
observations  were  recorded. 

In  an  attempt  to  identify  individual  birds  performing  on  an  area, 
three  males  were  mist-netted  and  banded  with  U.S.  Fish  and  Wildlife 
Service  bands.  In  the  first  netting  attempts  the  net  was  simply  set  in  the 
usual  path  of  flight  of  the  bird.  Later  a  portable  cassette  tape  recorder 
was  used  to  play  a  tape  recording  of  the  peenting  of  a  territorial 
woodcock  to  lure  the  territorial  bird  into  the  net. 

The  size  of  the  territory  (defended  area)  of  resident  birds  was 
tested  with  the  taped  peenting  call.  This  tape  was  played  at  various 
distances  from  the  landing  site  of  a  territorial  male  and  a  positive 
response  was  considered  to  be  an  overflight  with  the  issuance  of  a 
cackling  threat  note.  The  tape  was  played  at  the  same  volume  at  which  it 
was  recorded.  This  technique  also  allowed  the  observation  of  various 
reactions  of  territorial  birds  to  intruding  males. 

Weather  conditions  were  recorded  each  day  on  the  area.  Temperature 
was  measured  with  a  mercury  thermometer.  Wind  speed  was  estimated. 
In  the  consideration  of  light  intensity  effects,  only  completely  clear  or 
completely  cloudy  days  were  used. 

Data  were  tested  for  significance  when  necessary  with  Students'-t 
test.  Differences  were  considered  significant  at  the  95%  confidence  level 
and  highly  significant  at  the  99%  confidence  level. 

Results  and  Discussion 

Occupancy  of  Singing  Grounds 

The  first  woodcock  was  observed  performing  on  Area  1  on  March  15, 
1969.  The  number  of  birds  performing  and  the  total  number  of  birds 
observed  on  the  area  increased  from  that  date  until  about  March  25  and 
then  decreased,  more  or  less  stabilizing  during  the  first  week  in  April 
(Fig.  2).  Individual  singing  fields  were  occupied  for  periods  of  from  1  day 
to  2  months.  The  last  half  of  March  would  seem  to  be  the  time  of  the  main 
migratory  flight  in  this  area.  Migratory  males  obviously  occupied  per- 
forming areas  during  short  pauses  in  their  northward  movement. 


Ecology 


1(55 


15 
14 
13 
12 
Number      11 


of 


10 


Woodcocks        | 

6 

5 
4 
3 
2 

1 


^-  Total    birds  performing 
|-  Total    birds  observed 


J 


Milan 


15    16    17  18  21    24   25  26  28  29    3     6     8    10  14   16   17  20  22  23  25   26  29    4      7     9     14  21 
March  April  May 

Figure  2.     Number   of   performing    woodcocks    and   total   woodcocks    recorded   during    cen- 
suses at  Shidler  Forest,  1969. 

The  requirements  for  a  singing  field  are  fairly  rigid,  for  even 
transient  birds  performing  along  their  migratory  routes  prefer  certain 
areas  (11).  A  few  conditions  seemed  to  be  necessary  on  the  study  area. 
A  site  was  always  an  area  with  no  tall  herbaceous  vegetation,  either  an 
area  of  bare  ground,  of  matted  vegetation,  or  of  short  grasses  (mowed 
or  otherwise  checked),  and  small  rather  widely  scattered  woody  plants. 
Acceptable  heights  of  woody  vegetation  ranged  from  2-15  feet. 

Temperatures  did  not  seem  to  affect  the  performance  level  even 
when  coupled  with  precipitation;  birds  performed  at  normal  levels  on 
March  29  when  the  temperature  was  at  27  °F  and  on  March  25  when  the 


8 

7 
Number       0 

of  Birds        5 
4 

".  3 
2 

1 


Performing 


H  -  Temperature 

P~  No.  of  Birds  Performing 


60 


50 

°F 

at 

Time 

of 

Census 

40 

30 

20 

17        18       21        24        25       26       28       29 

Clear  Clear  Clear    Rain     Snow  Snow  Cdy.  Clear 

Date    (March) 

Weather 
Figure  3.     Comparison    of    temperature    and    the    number    of    woodcocks    performing     in 
late  March,   1969,   at   Shidler  Forest   (one  inch  of  snow  on  ground,   drifts  to  four  inches). 


166  Indiana  Academy  of  Science 

temperature  was  at  31  °F  and  a  light  snow  was  falling  (Fig.  3).  On 
March  26,  however,  with  the  temperature  at  28  °F,  a  light  snow  falling 
and  1  inch  of  snow  on  the  ground,  only  a  few  very  irregular  flights 
occurred  (Fig.  3).  Although  other  studies  showed  that  performance  levels 
decreased  as  temperatures  reached  35°  to  41°F  (2,  5,  7),  no  decrease  was 
evident  in  this  study  at  these  temperatures.  Snow  cover  was  apparently 
the  influencing  factor.  Mendall  and  Aldous  (7)  observed  that  birds  hesi- 
tated to  land  on  snow  and  Pettingill  (8)  reported  that  a  snow  cover  cur- 
tailed performances  and  that  those  that  did  occur  were  irregular. 

Fairly  strong  winds  also  failed  to  alter  performance  levels.  On  March 
28,  a  10-20  miles  per  hour  (mph)  wind,  with  gusts  to  30  mph,  blew  without 
inhibiting  performances  (Fig.  2).  Light  rain  had  no  depressing  effect,  but 
during  periods  of  heavy  rain  no  flights  were  made,  although  the  birds 
continued  peenting  from  the  ground.  Song  flights  resumed  when  intensity 
of  the  rain  decreased.  Sheldon  (11)  found  that  high  winds  with  or  with- 
out rain  and  heavy  downpours  without  wind  curtailed  breeding  activity. 
It  seems  that  the  drive  to  perform  courtship  flights  is  so  great  that  only 
the  most  extreme  environmental  conditions  would  cause  elimination  of  a 
performance. 

Display  Behavior 

Territorial  birds  usually  began  peenting  from  their  diurnal  areas 
5-15  min  before  moving  onto  their  singing  grounds.  Some  individuals 
gave  no  preperformance  peenting.  Mendall  and  Aldous  (7)  found  it 
unusual  for  distances  from  the  diurnal  area  to  the  singing  ground  to 
be  greater  than  100  yards.  Although  the  same  was  found  in  this  study, 
one  bird  had  its  diurnal  area  in  the  bottomland,  about  0.4  miles  from 
its  singing  ground  on  the  ridgetop,  Area  4-5.  This  bird,  however,  after 
11  days  shifted  to  a  performing  territory  within  75  yards  of  its  diurnal 
area,  when  that  territory,  Area  1,  was  abandoned  by  its  performing  bird 
(Fig.  1). 

On  clear  days  woodcocks  arrived  on  their  singing  grounds  an 
average  of  16  min  after  sunset  and  began  the  first  flight  an  average  of 
21  min  after  sunset.  Cloudy  conditions  caused  the  birds  to  occasionally 
move  onto  the  areas  well  before  sunset  and  greatly  increased  variability 
in  time  of  first  peent  and  of  first  flight  relative  to  sunset  (Table  1). 
Light  intensity  is  the  major  influencing  factor  on  time  of  first  on-area 
peent  and  first  flight.  The  greater  variability  in  time  of  first  peent  than 
in  time  of  first  flight  on  clear  days  is  undoubtedly  because  of  the 
variable  light  intensities  on  the  individual  diurnal  areas  (Table  1).  All  of 
singing  grounds  have  very  similar  light  conditions,  for  none  have  over- 
head cover.  This  first  flight  begins  when  the  light  intensity  reaches  about 
2  foot-candles  (4,  7).  The  greater  variability  evident  on  cloudy  days  in 
the  times  between  sunset  and  the  first  peent  and  flight  comes  from  the 
fact  that  even  a  complete  cloud  cover  can  let  through  extremely  variable 
amounts  of  light  (Table  1). 

A  total  of  89  flights  in  9  complete  performances  were  timed.  There 
was  much  variation  within  and  among  performances.  The  average  flight 


Ecology  167 

Table  1.  Time  after  official  sunset  of  first  peent  from  performing  area 
and  of  first  flight  of  male  woodcocks  at  Shidler  Forest,  1969. 


First  Peent 

First  Flight 

Measurement 

Clear  Sky 

Cloudy  Sky 

Clear  Sky 

Cloudy  Sky 

Mean  Length  (min) 
Range  (min) 
Standard  Deviation 
No.  of  Observations 

16.29 
7  to  24 
4.37 
24 

—0.08 
—  17  to  20 
12,03 
13 

21.18 

16  to  27 
2.28 
22 

8.00 
—13  to  22 

12.44 
11 

length  was  61.5  sec  with  a  range  of  52-76  sec.  The  time  intervals  between 
flights  were  also  very  variable. 

Three  component  parts  of  a  total  of  29  flights  from  3  separate 
performances  were  timed — the  ascent  (period  before  vocal  song),  the 
song,  and  the  silent  descent.  By  far  the  most  variable  component  among 
the  three  was  the  ascent  with  the  other  two  being  relatively  invariable. 
The  mean  length  of  time  for  the  ascent  was  45.2  sec  with  a  standard 
deviation  of  3.30;  of  the  song  9.7  sec  with  a  standard  deviation  of  0.81; 
and  of  the  silent  descent  7.0  sec  with  a  standard  deviation  of  0.63.  Almost 
all  of  the  variability  in  the  flights  was  due  to  the  variation  in  ascent 
times.  Because  the  song  and  silent  period  occurred  during  fairly  direct, 
though  somewhat  zigzagged,  groundward  plunges,  the  altitude  attained 
during  such  flights  must  have  been  fairly  uniform.  Estimates  of  heights 
attained  vary  from  200-300  feet  (7,  8);  Sheldon  (11)  measured  the 
altitude  of  3  flights  of  the  same  bird  and  found  all  to  be  275  feet.  If  this 
constancy  of  song  and  silent  descent  length  hold  for  woodcocks  elsewhere, 
then  variation  evident  in  flight  length  can  be  attributed  to  variation  in 
ascent  times.  Brewster  (3)  reported  the  length  of  2  songs  as  11  and  12 
sec.  The  average  and  the  range  of  flight  times  in  this  study  were  longer 
than  those  of  other  studies,  and  the  area  covered  in  one  flight  was 
also  larger,  averaging  over  2  acres.  Pitelka  (9)  reported  average  flight 
coverage  of  only  V3  acre  and  flight  times  of  29  to  60  sec  with  an  average 
of  43  sec  (calculated  by  author).  The  majority  of  the  flights  timed  by 
Mendall  lasted  50  to  55  sec  with  a  range  from  44  to  63  sec  (7).  The 
author  suggests  that  the  time  of  an  ascent,  and  thus  a  flight,  is  deter- 
mined by  the  time  required  to  reach  a  certain  altitude,  and  that  this  time 
is  directly  proportional  to  the  amount  of  area  encompassed  in  a  flight. 
This  area  may  in  turn  be  determined  by  several  factors  including  size  of 
singing  field,  height  of  surrounding  vegetation,  and  juxtaposition  of  other 
territorial  males. 

The  number  of  peents  per  minute  varied  greatly,  ranging  from  7  to 
28  per  minute.  The  woodcocks  occasionally  walked  or  ran  short  distances 
between  peents  but  most  often  remained  in  one  spot,  usually  pivoting  a 
little  after  each  peent.  Some  birds  turned  360°  in  one  direction  while 
others  pivoted  90-180°  in  one  direction  and  then  reversed  directions.  This 
rotation   caused    the    volume    of   the    peenting    to    seem    to    vary    to    an 


168  Indiana  Academy  of  Science 

observer  and  probably  served  to  make  territorial  birds  conspicuous  for 
the  maximum  distance  possible  in  all  directions. 

The  average  length  (from  first  peent  on  the  area  to  the  last)  of  26 
performances  not  influenced  by  moonlight  was  34.2  min,  ranging  from 
16  to  53  min.  On  a  moonlit  night,  April  29,  a  performance  lasted  74  min. 
It  was  found  that  performances  were  highly  significantly  longer  on  cloudy 
than  clear  days,  41.3  and  31.5  min,  respectively.  The  lengths  of  per- 
formances were  also  more  variable  on  cloudy  than  on  clear  days.  Total 
performance  length  was  greatly  influenced  by  light  conditions.  Cloudiness 
extended  performances  because  of  earlier  starting  times  and  fairly  com- 
parable stopping  times;  the  greater  variability  in  lengths  of  perform- 
ances on  cloudy  days  was  undoubtedly  caused  by  differential  light  pene- 
tration through  a  full  cloud  cover.  A  long  performance  on  a  moonlit  night 
as  observed  in  the  study  has  been  reported  by  many  investigators.  It  is  a 
good  example  of  the  degree  to  which  various  aspects  of  the  performance 
are  light  intensity  dependent  (7,  8,  11). 

The  manner  in  which  performances  ended  differed  among  individuals. 
A  bird  left  an  area  after  the  last  flight  either  immediately,  after  a  long 
period  of  silence,  or  most  commonly  after  several  minutes  of  decelerating 
peenting. 

Territorial  Behavior 

Territorial  birds  tolerated  no  territorial  intrusions  by  other  males. 
Cackling  was  the  most  common  threat  exhibited.  When  an  intruder 
peented  within  an  occupied  territory,  the  territorial  bird  usually  would 
fly  immediately  toward  the  source  of  the  strange  peenting  and  cackle. 
This  generally  discouraged  the  intruder  which  hid  and  became  silent  or 
departed.  The  peent  was  the  major  communication  in  woodcock  relation- 
ships and  served  the  dual  purpose  of  advertising  occupancy  and  of  dis- 
closing the  presence  of  intruders. 

Several  chases  were  observed  and  in  all  cases  both  birds  seemed  to 
flush  from  nearly  the  identical  spot.  When  first  sighted,  the  birds  were 
usually  10  to  15  feet  above  the  ground  with  one  about  5  feet  ahead  of  the 
other.  This  interval  was  kept  throughout  the  chase  as  the  birds  climbed 
at  a  45  to  90°  angle  from  the  horizontal.  Usually  the  only  sound  emitted 
during  these  chases  was  wing-twitter  although  cackling  occasionally 
occurred.  About  half  of  these  chases  ended  with  the  bird  giving  chase 
veering  off  and  beginning  a  flight  song.  At  other  times  the  chase  con- 
tinued upward  until  both  birds  were  lost  from  view  and  several  minutes 
elapsed  before  the  territorial  bird  returned  to  the  area.  No  physical  con- 
tact between  birds  was  ever  observed.  Similar  chases  were  observed  by 
Pitelka  (9)  who  attributed  them  to  two  males  simultaneously  starting 
their  flights.  From  the  observations  in  this  study,  this  does  not  seem 
likely.  It  is  also  unlikely  that  this  is  a  courtship  flight  including  male  and 
female,  because  in  none  of  the  11  chases  observed  did  the  bird  being 
chased  ever  return  to  the  field.  It  seems  most  probable  that  it  is  a  very 
ritualized  form  of  chase  which  has  developed  evolutionary  to  maximize 


Ecology  169 

survival  by  preventing  injury  to  combatants  and  still  maintaining  terri- 
torial integrity. 

An  attempt  was  made  to  determine  the  area  defended  by  a  terri- 
torial bird.  On  May  5  and  6,  as  the  peenting  tape  was  played  from 
various  locations  on  the  area,  the  territorial  male  on  Area  1  reacted  by 
flying  toward  it  and  cackling  over  a  total  area  of  4.38  acres.  The 
greatest  distance  from  which  a  response  was  drawn  was  330  feet.  It 
had  been  thought  previously  that  the  birds  defended  at  most  the  area 
covered  by  their  flights,  estimates  of  which  varied  from  V3  to  2  acres  in 
different  studies  (8,  9).  Because  this  bird  reacted  at  all  points  tested,  4.38 
acres  is  the  minimum  estimate  of  the  territorial  size.  If  the  greatest  dis- 
tance from  which  a  response  was  drawn,  330  feet,  is  assumed  to  be  the 
territorial  limit  and  a  circle  circumscribed  about  the  landing  site  with 
this  as  the  radius,  the  area  included  is  7.85  acres,  a  considerably  larger 
territory   than   anyone   had   previously   suspected. 

Several  aggressive  reactions  of  territorial  birds  to  the  presence  of 
the  investigator  were  encountered.  On  May  1,  I  was  lying  about  100  feet 
from  the  usual  landing  site  of  the  bird  on  Area  1.  The  taped  peenting 
was  played  to  test  the  bird's  reaction  to  it.  The  territorial  bird  cackled, 
flew  toward  the  sound  and  lit  within  20  feet  of  the  recorder,  which  was 
on  the  ground  about  3  feet  from  my  head.  After  flying  closer  to  the 
recorder  and  peenting  several  times,  it  flushed  and  returned  to  its  usual 
landing  site.  The  complete  approach  process  was  repeated  twice  more. 
On  the  third  approach,  the  bird  landed  5  feet  from  the  recorder  and 
approached  it  on  foot.  It  stood  silently  next  to  it  for  a  few  seconds  and 
then  began  searching  through  the  grass  around  the  recorder.  Suddenly  it 
moved  through  the  weeds  and  appeared  about  a  foot  from  my  face. 
Almost  immediately  the  woodcock  lunged  toward  me  and  struck  my  eye 
with  its  bill  and  then  grabbed  a  tuft  of  hair  in  its  bill  and  yanked  vigor- 
ously several  times.  The  bird  then  released  its  hold,  jumped  back  about  a 
foot,  and  stood  watching  me.  I  remained  as  still  as  possible.  The  whole 
attack  was  then  repeated  with  the  bird  first  striking  my  eye  and  then 
yanking  my  hair.  After  this  attack  an  attempt  was  made  to  catch  the 
bird  by  hand,  but  its  reactions  were  much  too  quick  and  it  flushed.  How- 
ever, it  immediately  responded  again  to  the  peenting  tape  and  attacked 
my  hand  several  times  as  I  shuffled  it  in  the  grass.  These  attacks  indicate 
how  strong  a  role  peenting  plays  in  the  territorial  behavior  of  the  wood- 
cock. This  woodcock  and  others  were  invariably  attracted  by  the  peenting 
from  the  tape  even  seconds  after  unsuccessful  attempts  to  capture  them. 
After  a  woodcock  is  attracted  to  an  area  by  alien  peenting,  any  movement 
is  evidently  attributed  to  the  intruder  and  is  attacked.  In  both  face-to-face 
encounters  the  woodcock  first  struck  my  eye,  the  only  place  in  which 
movement  occurred.  It  also  attacked  my  hand  only  when  it  was  shuffled 
in  the  grass.  This  would  be  the  expected  reaction  sequence  in  a  species 
which  defends  territory  only  during  periods  of  marginal  light  intensity 
and  against  somber  colored  intruders. 

In  an  attempt  to  band  birds  so  that  homing  could  be  tested  in  subse- 
quent years,  the  bird  on  Area  1  was  mist-netted  and  banded  on  April  25. 


170 


Indiana  Academy  of  Science 


Soon  after  this  a  different  behavior  pattern  for  the  bird  on  this  area 
was  noted  (i.e.,  changes  in  landing  site,  flight  pattern,  diurnal  area, 
reaction  to  taped  peenting,  and  various  other  performance  parameters) 
and  a  male  caught  on  May  7  proved  to  be  a  different  bird.  The  subse- 
quent appearance  soon  afterward  of  a  third  behavior  pattern  lead  to  the 
capture  and  banding  of  a  third  male  from  Area  1  on  May  16.  The  appear- 
ance of  the  second  bird  on  Area  1  was  closely  linked  with  the  discontinu- 
ance of  the  performances  of  the  bird  on  Area  4-5,  and  the  appearance  of 
the  third  bird  with  the  cessation  of  the  performances  of  the  bird  on  Area 
10  (Fig.  4).  Woodcocks  from  adjacent  areas  undoubtedly  replaced  previ- 
ous territorial  birds  as  they  disappeared  from  Area  1.  Both  birds  which 
were  replaced  on  Area  1  disappeared  soon  after  netting  and  banding.  It 
is  impossible  to  determine  what  caused  this  disappearance,  but  the 
trauma  of  the  netting  and  banding  operation  may  have  caused  the  birds 
to  move  off  the  study  area.  However,  the  subnormal  post-banding  per- 
formance by  bird  "A"  illustrated  that  banded  birds  may  have  ceased 
performing  while  remaining  on  the  study  area  (Fig.  4).  Because  of  this 
replacement  pattern,  Area  1  was  interpreted  as  being  the  prime  per- 
formance site  on  the  study  area,  and  thus,  the  one  most  competed  for. 
The  fact  that  it  had  at  least  three  males  on  it  with  not  a  day  of  non- 
occupancy  is  indicative  of  its  attractiveness. 

Sheldon   (11)   suggested  that  competition  for  open  spaces  was  very 
important  in  the  evolutionary  shaping  of  the  territorial  behavior  of  the 


B 


March 
15         20        25 

I 


30  1 


April 
10        15        20 


May 


25       30 

H«,i— 


H  '  h 


First  performing  bird  netted   J 

Subnormal  performance   - 

Study  area  not  checked   - 

New  pattern  noted  - 

Bird  on  area   4-5  no  longer  performing   - 

Performing  bird  "B"  netted  - 
Normal  performance  of  "B"  pattern 
New  pattern   noted 
Bird  on  area  JO    no  longer   performing 


15        20     24 


Performing  bird  "C"  netted 

Normal    performance   of   "C"  pattern 

Subnormal  "C"  performance 

No    flights— only  peenting 

No  bird  on  area 


FIGURE  4.      Chronological    sequence    of    events    on    Area    1    in    woodcock    study    at    Sh idler 
Forest,  1969. 


Ecology  171 

woodcock,  for  in  primeval  times  very  few  openings  existed  in  the  eastern 
United  States  and  Canada.  It  is  probable  that  even  now  strong  competi- 
tion for  preferred  sites  exists.  This  competition  and  territorial  defense 
appears  to  be  principally  a  ritualistic  type  which  involves  little  physical 
contact.  Lack  (6)  said  that  avoidance  by  others  of  occupied  areas  was  the 
primary  factor  in  maintaining  territorial  integrity  in  birds.  Tinbergen 
(12)  said  that  his  previous  view  that  hostility  was  the  primary  factor  as 
well  as  Lack's  that  avoidance  was  primary  were  both  one-sided  and  that 
the  two  factors  operate  in  tandem.  The  cofunctioning  of  these  two  factors 
may  offer  a  partial  explanation  for  the  tenacious  drive  to  perform  which 
has  evolved  in  the  woodcock.  When  a  bird  is  on  territory,  it  is  at  an 
advantage  since  its  defense  is  nearly  always  successful  because  of  the 
natural  avoidance  of  occupied  areas  by  unattached  males.  However,  if  a 
bird  fails  to  occupy  its  territory  for  a  day  or  more  due  to  the  weather 
or  some  other  factor  and  another  bird  occupies  the  area  in  the  interim, 
the  former  territorial  bird  is  then  at  a  disadvantage.  Those  birds  which 
establish  a  territory  in  a  favorable  location  and  perform  there  daily, 
regardless  of  conditions,  will  be  selected  for  evolutionarily,  for  they  have 
a  greater  chance  of  being  on  territory  throughout  the  breeding  season  and 
thus  more  chances  to  procreate. 


Literature  Cited 

1.  Bent,  A.  C.   1927.  Life  histories  of  North  American  shorebirds,  part  1.  U.S.   Nat.   Mus. 
Bull.  142.  420  p. 

2.  Blankenship,    L.    H.    1957.    Investigations    of    the    American    woodcock    in    Michigan. 
Mich.  Conserv.  Dep.,  Game  Div.,  Rep.  No.  2123.  217  p. 

3.  Brewster,  William.    1894.  Notes  and  song  flight  of  the  woodcock    {Philohela  minor). 
Auk  11:291-298. 

4.  Duke,    Gary    E.    1966.    Reliability    of   censuses    of   singing   male    woodcocks.    J.    Wildl. 
Mgt.  30  :697-707. 

5.  Goudy,    William    H.    1960.    Factors   affecting   woodcock    spring    population    indexes    in 
southern  Michigan.  Mich.  Conserv.  Dep.,  Game  Div.,  Rep.  2281.  44  p. 

6.  Lack,    David.    1954.    The    natural    regulation    of   animal    numbers.    Oxford    University 
Press,  London.  343  p. 

7.  Mendall,  Howard  L.,  and  Clarence  M.  Aldous.   1943.  The  ecology  and  management 
of  the  American  woodcock.  Maine  Coop.  Wildl.  Res.  Unit,  Orono.  201  p. 

8.  Pettingill,     Olin     Sewall,     Jr.     1936.     The     American     woodcock,     Philohela     minor 
(Gmelin).  Mem.  Boston  Soc.  Natur.  Hist.  9(2)  :167-391. 

9.  Pitelka,    Frank    A.    1943.    Territoriality,    display,    and   certain   ecological    relations    of 
the  American  woodcock.  Wilson  Bull.  55(2)  :88-114. 

10.  Robbins,   Chandler   S.,   Bertel   Bruun,   and   Herbert   S.    Zim.    1966.    Birds   of   North 
America.  Golden  Press,  New  York.  340  p. 

11.  Sheldon,  William   G.   1967.  The  book  of  the  American  woodcock.  The  University  of 
Mass.  Press,  Amherst.  227  p. 

12.  Tinbergen,  N.  1957.  The  function  of  territory.  Bird  Banding  VIII  il>  :l-8. 


Foods  of  the  White-footed  Mouse,  Peromyscus  leucopus 
noveboracensis,  from  Pike  County,  Indiana1 

Gwilym  S.  Jones,  Purdue  University 

Abstract 

During  a  fauna]  study  of  a  strip-mined  area  in  Pike  County,  Indiana,  the  stomachs 
of  489  white-footed  mice  (Peromyscus  leucopus  noveboracensis)  were  examined  to  de- 
termine food  preferences.  The  stomach  contents  were  identified  by  comparison  with 
seeds,  arthropods,  and  other  materials  collected  on  the  study  plots  where  the  mice  were 
trapped.  The  average  %  volume  of  each  food  was  estimated  for  each  stomach.  Of  the 
68  different  materials  identified,  undifferentiated  starchy  substance  and  invertebrates  were 
the  most  common.  Wild  cherry  (Prunus  sp.),  blackberry  (Rubus  sp.)  and  wood  son-el 
(Oxalis  sp.)  were  the  most  abundant  identifiable  plant  foods.  The  average  %  volumes  of 
food  types  (i.e.  plant,  animal,  and  "other")  were  computed  for  each  season.  The  results 
showed  that  plant  foods  are  dominant  throughout  the  year,  but  increase  noticeably  during 
the  fall  and  winter. 

Introduction 

As  part  of  a  faunal  study  of  a  strip-mined  area  (6),  stomachs  of  all 
white-footed  mice  captured  during  a  trapping-  program  were  examined 
to  determine  food  preferences  of  the  species.  This  paper  summarizes  the 
findings  of  the  study. 

Study  Area 

The  study  area  covers  approximately  IV2  square  miles  in  Monroe 
and  Lockhart  Townships  (R7W  and  R8W,  T3S),  Pike  County,  Indiana. 
About  V2  of  the  area  has  been  strip-mined  for  coal.  The  topography 
resulting  from  the  strip-mining  is  either  pyramidal,  parallel  ridges  rang- 
ing in  height  to  75  feet,  or  extensive  flat  areas  with  intermittent  small 
hills.  The  stripping  began  in  1921  and  ended  in  1961.  The  unstripped  half 
of  the  area  has  rolling  topography  with  intermittent  low,  wet  areas. 

The  vegetation  was  divided  into  nine  distinct  cover-types  based  on 
dominant  species.  Pines,  mixed  pine-hardwoods,  pine-hardwood  saplings, 
and  black  locust  occurred  only  on  the  stripped  portions.  Hardwoods,  brush 
and  weeds  occurred  on  both  stripped  and  unstripped  land,  and  bottom- 
land hardwoods  and  crops  occurred  only  on  the  unstripped  land.  Except 
for  brush  and  weeds,  all  cover  types  on  the  stripped  land  had  been 
planted. 

Methods 

The  food  habits  study  was  based  on  mice  that  were  captured  during 
a  population  study  which  employed  100  plots  randomly  selected  from  a 
gridded  map  of  the  area.  The  plots  were  125  feet  square  and  contained 
18  trapping  sites  of  3  traps  each.  Each  plot  was  set  for  three  nights.  The 
stomachs  of  the  captured  mice  were  removed  as  soon  as  possible  and 
preserved  in  70%  alcohol.  During  trapping  all  plants  found  on  each  plot 
were  recorded  and  samples  of  their  seeds  and  fruits  collected. 


1  Work  was  supported  by  the  Indiana  Dept.  of  Natural  Resources,  Midwest  Coal 
Producers,  Central  States  Forest  Experiment  Station  and  Purdue  Agricultural  Experiment 
Station. 

172 


Ecology  173 

After  the  trapping  program  was  completed,  the  stomachs  were  dis- 
sected and  the  contents  washed  into  a  dish  of  alcohol.  The  contents  were 
then  identified  by  comparison  with  the  sample  collections  of  seeds  and 
animal  matter.  The  volume  of  each  food  was  estimated  according  to  the 
methods  of  Hamilton  (4)  and  Whitaker  (9).  Every  effort  was  made  to 
identify  all  stomach  materials.  However,  some  unidentifiable  contents 
were  listed  as  unidentified  plant  or  animal  materials,  or  unknown,  depend- 
ing upon  their  nature.  The  data  were  programmed  and  analyzed  by 
computer.  Because  the  study  emphasized  plant  materials,  arthropod 
remains  were  identified  only  to  general  classifications. 

Results 

The  most  prevalent  year-round  food  eaten  by  the  white-footed  mouse 
was  undifferentiated,  starchy  material.  Insects  and  Lepidoptera  or 
Hymenoptera  larvae  were  other  preferred  foods.  Unidentified  seeds  ranked 
below  starchy  material  and  insects.  Of  the  identifiable  seeds,  those  with 
the  highest  volumes  were  wild  cherry  (Prunus  serotina) ,  blackberry 
(Rubus  sp.),  and  yellow  wood  sorrel  (Oxalis  sp.).  All  stomach  items 
except  those  in  the  next  paragraph  are  listed  in  Table  1. 

The  following  foods  were  found  in  quantities  of  less  than  0.1% 
volume:  Compositae  seeds,  0.8%  frequency;  crowfoot  (Ranunculus) 
seeds,  Annelida,  and  touch-me-not  (Impatiens)  seeds,  0.6 %  frequency; 
pupae  cases,  pokeberry  (Phytolacca)  seeds,  tick  trefoil  (Desmodium) 
seeds,  and  metal,  0.4%  frequency;  trumpet  creeper  (Campsis)  seeds, 
Coleoptera  larvae,  flower  petals,  bones,  redbud  (Cercis)  seeds,  oxeye 
daisy  (Chrysanthemum)  seeds,  avens  (Geum)  seeds,  sedge  (Scirpus) 
seeds,  daisy  fleabane  (Erigeron)  seeds,  lettuce  (Lactuca)  seeds,  bluets 
(Houstonia)  seeds,  St.  John's-wort  (Hypericum)  seeds,  and  plantain 
(Plantago)  seeds,  0.2%  frequency. 

The  seeds  of  planted  trees  consumed  by  the  white-footed  mouse  were 
maple  (Acer),  pine  (Pirius) ,  ash  (Fraxiyius),  elm  (Ulmus),  and  black 
locust  (Robinia).  Of  the  planted  species,  maple  had  the  highest  average 
volume  and  %  frequency,  but  these  values  were  only  1.2%  volume  and 
8.3%  frequency. 

A  consideration  of  the  seasonal  preferences  indicates  that  plant  foods 
are  dominant  throughout  the  year  but  increase  noticeably  during  the 
fall  and  winter,  after  the  plants  have  dropped  their  fruits  and  arthropods 
have  become  scarce. 


174  Indiana  Academy  of  Science 

Table  1.  Average  percent  volume  and  percent  frequency  of  foods  found  in  489  white- 
footed  mice  (Peromyscus  leucopus)  stomachs  from  September,  1965,  to  August,  1966,  in 
Pike  County,  Indiana. 

Food  Volume         Frequency 

Starchy  material 

Insects 

Lepidoptera  or  Hymenoptera  larvae 

Unidentified   seeds 

Wild    cherry     (Prunus)     seeds 

Rubus    (sp. )    seeds 

Yellow    wood    sorrel     (Oxalis)    seeds 

Unidentified    plant 

Hair 

Chilopoda 

Green   vegetation 

Indian    hemp    (Apocynum)    seeds 

Cranesbill     {Geranium)     seeds 

Mollusca 

Maple    (Acer)    seeds 

Unknown 

Arachnida 

Flesh 

Pine    (Pinus)    seeds 

Endogone 

Ash    {Fraxinus)    seeds 

Sumac     (Rhus)     seeds 

Elm    iUlmus)    seeds 

Bush  clover  (Lespedcza)   seeds 

Rose   (Rosa)    seeds 

Rumex     (sp. )     seeds 

Bittersweet     (Celastrus)     seeds 

Cottonwood   (Populus)   seeds 

Poison    ivy    (Rttus)    seeds 

Sassafras    (Sassafras)    seeds 

Pebbles 

Honeysuckle    (Loniccra)    seeds 

Unidentified  grass  seeds 

Panic  grass    (Panicum)    seeds 

Dogwood    (Cornus)    seeds 

Unidentified    animal 

Everlasting  pea    (Lathyrus)    seeds 

Unidentified   fungus 

Grape     (Vitis)    seeds 

Greenbriar     (Smilax)     seeds 

Sedge    (Car  ex)    seeds 

Feathers 

Peppergrass     (Lcpidium)    seeds 

Foxtail    (Setaria)    seeds 

Black  locust    (Robinia)    seeds 

Sweet  clover    {Melilotus)    seeds 

Apple   (Pyrus)   seeds 


30.1 

77.3 

14.5 

7S.fi 

9.2 

42.2 

7.5 

41.0 

5.:-! 

21.4 

3.7 

10.0 

3.6 

9.4 

•A. -A 

23.3 

2.2 

41.2 

1.9 

18.7 

1.8 

12.7 

l.fi 

7.0 

1.5 

5.0 

1.2 

8.9 

1.2 

8.3 

0.!) 

8.3 

0.8 

9.8 

0.8 

4.8 

0.8 

4.8 

0.7 

6.0 

0.7 

4.4 

<u; 

2.3 

0.5 

A.A 

0.5 

A  A 

0.5 

2.0 

0.4 

3.7 

0.4 

2.0 

0.4 

2.5 

0.4 

2.5 

0.4 

1.5 

0.3 

7.:-! 

0.3 

1.5 

0.2 

3.3 

0.2 

2.7 

0.2 

2.1 

0.2 

1.5 

0.2 

0.6 

0.1 

3.5 

0.1 

1.5 

0.1 

1.2 

0.1 

1.2 

0.1 

1.0 

0.1 

1.0 

0.1 

0.6 

0.1 

o.i; 

0.1 

o.t; 

0.1 

0.2 

Ecology  175 


Discussion 


I  agree  with  Hamilton  (4)  that  the  starchy  material,  called  mast  by 
Whitaker  (9),  is  composed  of  many  different  materials  (such  as  mast, 
root  tips,  and  seeds)  which  cannot  be  separated  by  observation.  Hamilton 
(4)  found  stomachs  containing  nothing  but  starchy  material.  Such  was 
the  case  in  this  study.  Undoubtedly  some  of  the  starchy  material  was 
composed  of  acorns  (Quercus) ,  hickory  nuts  (Carya) ,  and  other  such 
fruits.  Whitaker  (personal  communication)  indicates  that  he  has  found 
portions  of  the  exocarp  of  such  fruits  still  clinging  to  the  material.  Such 
was  not  the  case  in  this  study. 

Green  vegetation  was  separated  from  unidentifiable  plants  because 
this  material  is  composed  of  leaves,  buds,  or  other  vegetative  plant  parts. 
This  is  supported  by  Hamilton's  (4)  contention  that  the  remains  of  buds, 
and  Jameson's  (5)  conclusion  that  cotyledons  of  sprouting  seeds,  when 
eaten  by  mice,  retain  their  green  color.  By  my  methods  it  was  impossible 
to  determine  the  origin  of  the  plants  or  the  parts.  The  unidentified  plant 
material  was  thought  to  be  fragments  of  xylem  or  phloem  (2). 

The  fungus  Endogone  was  a  common  item  in  the  stomachs  of  many 
small  mammals  by  a  number  of  investigators  (1,  2,  3,  4,  7,  and  11). 
However,  it  was  not  a  common  food  of  the  white-footed  mouse  in  this 
study,  ranking  20th  in  volume  with  a  frequency  of  0.7%. 

Although  insects  were  a  major  food,  they  never  surpassed  plant  food 
in  volume  or  frequency.  Similar  observations  were  reported  by  Hamilton 
(4),  Jameson  (5),  Williams  (10),  and  Whitaker  (8,  9). 

Hair,  found  in  41.5%  of  the  stomachs,  was  often  in  the  form  of  balls, 
suggesting  that  it  resulted  from  the  grooming  habits  of  the  mice.  Some 
hair  undoubtedly  accompanied  ingested  flesh  which  was  found  in  4.8%  of 
the  stomachs. 

Stomach  analyses  admittedly  provide  only  an  indication  of  food 
preferences.  However,  they  provide  some  information  concerning  the 
ecology  of  the  mammal  and  indicate  what  plants  it  eats.  Although  some 
believe  that  enough  food  habits  work  has  been  done  for  Peromyscus,  of 
the  68  foods  identified,  seeds  of  27  plant  species  were  new  foods  for  the 
species.  To  the  author's  knowledge  none  has  been  reported  before.  The 
most  common  of  these  were  Indian  hemp  (Apocynum)  and  cranesbill 
{Geranium)   with  frequencies  of  7.9%    and  5.0%,  respectively. 


Acknowledgements 

Thanks  are  extended  to  Dr.  R.  E.  Mumford,  Purdue  University, 
Dr.  J.  O.  Whitaker,  Jr.,  Indiana  State  University;  and  William  Crawford, 
Enos  Coal  Mining  Company. 


176  Indiana  Academy  of  Science 

Literature  Cited 

1.  Bakerspigel,    A.    1958.    The   spores    of   Endogone    and   Melanogaster   in    the    digestive 
tracts  of  rodents.  Mycologia  50  :440-442. 

2.  Calhoun,    J.    B.    1941.    Distribution   and   food   habits   of   mammals   in    the   vicinity    of 
the  Reelfoot  Biological  Station.  J.  Tenn.  Acad.  Sci.  6:177-225. 

3.  Dowding,   E.    S.    1955.   Endogone  in   Canadian   rodents.    Mycologia   47 :51-57. 

4.  Hamilton,    W.    J.,    Jr.    1941.    The   food   of   small   forest    mammals    in    eastern    United 
States.  J.  Mammal.  21  :250-263. 

5.  Jameson,    E.    W.,    Jr.    1952.    The    food    of    deer    mice,    Peromyscus    maniculatus    and 
P.  boylei,  in  the  northern    Sierra  Nevada  California.   J.   Mammal.   33  :50-60. 

6.  Jones,    G.    S.    1967.    Vertebrate   ecology    of    a    strip-mined    area    in    southern    Indiana. 
M.  S.  Thesis,  Purdue  Univ.  158  p. 

7.  Whitaker,    J.    O.,    Jr.    1962.    Endogone,    Hymenog  aster,    and    Melanogaster    as    small 
mammal  foods.  Amer.  Midland  Natur.  67:152-156. 

8.    .     1963.    Food    of    120    Peromyscus    leucopus    from    Ithaca,    New    York.    J. 

Mammal.  44  :418-419. 

9.  -  — .    1966.    Food   of   Mus   musculus,    Peromyscus   maniculatus,    and   Peromyscus 
leucopus  in  Vigo  County,  Indiana.  J.  Mammal.  47:473-486. 

10.  Williams,  O.  1959.  Food  habits  of  the  deer  mouse.  J.  Mammal.  40:415-419. 

11.  Williams,    O.    and    B.    A.     Finney.    1964.    Endogone — food    for    mice.    J.    Mammal. 
45:265-271. 


Observations  on  Ecology  and  Behavior  of  Indiana  Ruffed  Grouse1 

John  P.  Muehrcke  and  Charles  M.  Kirkpatrick,  Purdue  University 


Abstract 

On  an  840-acre  study  area  in  Monroe  County,  vegetation  types  consisted  of  second- 
growth  hardwoods  including  a  19-year-old  burn,  old  fields,  pine  plantations,  and  wildlife 
openings.  Flush  surveys  showed  that  grouse  used  hardwoods  most  consistently  (especially 
the  burn),  pines  for  winter  and  hunting  season  protection,  and  other  types  very  little. 
Age  ratios  of  summer-flushed  and  summer-trapped  grouse  were  essentially  even  in  1968. 
Eight  grouse  broods  found  in  summer  of  1968  averaged  3.1  young,  less  than  noted  in 
1965.  Brood  hens  showed  protective  behavior  until  young  were  10-12  weeks  old.  Extent 
of  brood  separation  movements  varies  among  individual  young.  Fifteen  drumming  males 
were  found  in  1969  compared  with  12  in  1968.  In  1969,  drumming  began  February  28, 
the  number  of  drumming  males  increasing  until  April  10  when  all  15  males  drummed, 
and  decreasing  afterward  until  only  2  were  heard  April  23.  The  daily  period  from 
about  %  hour  before  sunrise  until  y2  hour  after  sunrise  marked  the  most  intense  drum- 
ming activity.  The  intervals  between  drumming  performances  were  shortest  just  before 
and  just  after  sunrise.  The  drumming  act  averaged  5+  seconds.  Males  occasionally  roosted 
on  drumming  logs,  but  rarely  drummed  at  night.  Besides  their  main  drumming  logs,  all 
males  had  one  or  more  alternate  logs. 

A  century  ago  ruffed  grouse  (Bonasa  umbellus)  were  probably  found 
throughout  Indiana  woodlands,  being  reported  in  58  counties  (13),  but 
the  effect  of  man  has  limited  this  species  to  the  less  intensively  used 
hills  of  south-central  Indiana.  Evidence  mounts  that  the  bird  is  holding 
its  own  if  not  actually  thriving  in  the  managed  public  forest  lands. 
Thurman  (25)  started  a  study  on  ruffed  grouse  ecology  in  Monroe  County 
to  which  a  report  by  Muehrcke  (19)  represents  a  continuation.  The  work 
of  these  students,  supervised  by  the  junior  author,  proposes  to  accumulate 
data  for  the  better  understanding  of  Indiana  ruffed  grouse  ecology  to 
aid  in  management  practices  that  will  benefit  this  fine  bird. 

The  study  concentrated  on  an  840-acre  part  of  the  Hoosier  na- 
tional Forest  in  southeastern  Monroe  County  locally  known  as  Geiger 
Ridge.  It  lies  within  the  western  mesophytic  and  oak-hickory  climax 
vegetation  (17)  in  which  man  has  wrought  many  changes  to  the  forest. 
Most  of  the  wider  valley  bottoms  and  ridge  tops  were  cleared  and 
farmed  and  remaining  timber  was  extensively  logged.  Now  the  most 
common  species  in  the  overstory  include  chestnut  oak  (Quercus  spp.), 
black  oak  (Quercus  velutina  Lam.),  red  oak  (Quercus  spp.),  white  oak 
(Quercus  alba  L.) ,  and  hickory  (Carya  spp.).  Species  nomenclature  fol- 
lows Deam  (5).  In  1935  the  U.  S.  Forest  Service  began  buying  the 
worked-out  farms,  allowing  some  cleared  fields  to  revert  to  forest  suc- 
cessional  stages  and  planting  some  sites  with  pines.  This  history  of 
land  use  has  resulted  in  some  distinct  vegetation  types  consisting  of 
second-growth  hardwoods  including  a   19-year-old  burn,   old  fields,  pine 


1  Journal  Paper  No.  3856  from  Purdue  University  Agricultural  Experiment  Station, 
based  on  a  master's  thesis  by  John  P.  Muehrcke  with  research  support  from  the  Indiana 
Division  of  Fish  and  Game  and  cooperation  from  U.S.  Forest  Service  personnel,  Hoosier 
National  Forest. 

177 


178  Indiana  Academy  of  Science 

plantations,   and   wildlife   openings.    Thurman    (25)    describes   these   and 
other  physical  and  historical  features  of  the  area  in  detail. 

Methods 

To  locate  grouse  broods,  the  observer  made  intensive  searches  in 
all  cover  types  in  early  summer.  Brood  locations  later  became  focal 
points  for  trapping  with  cloverleaf  traps  (4).  Trapping  was  done  pri- 
marily to  gather  information  about  age  ratios,  grouse  movements,  and 
to  provide  a  basis  for  population  estimation.  Sex  and  age  were  deter- 
mined for  each  trapped  bird,  which  was  then  color-banded  and  released. 

Throughout  the  study,  major  effort  was  devoted  to  observing  be- 
havior of  grouse  in  their  natural  environment.  Hens  with  broods  and 
drumming  males  provided  most  such  contacts.  Whenever  a  brood  was 
discovered,  usually  by  flushing,  the  observer  concealed  himself  to  watch 
the  behavior  of  hen  and  young  as  they  regrouped  from  hiding.  Ten  1/10- 
acre  vegetation  samples  were  taken  to  describe  the  plant  communities 
where  broods  were  flushed.  Muehrcke  (19)  used  a  line  transect  method 
as  the  sampling  technique. 

The  observer  located  all  drumming  males  on  the  area  by  walking 
along  ridge  tops  from  xk  hour  before  sunrise  to  Vz  hour  after  sunrise. 
Each  ridge  top  was  covered  at  least  twice  during  the  clear  calm  days  of 
March  and  April.  The  sound  of  drumming  revealed  drumming  log  loca- 
tions and  offered  chances  to  study  the  performing  males.  Certain  drum- 
ming males  were  watched  intensively  for  recording  behavior.  Drumming 
logs  of  all  cocks  were  periodically  examined  for  signs  of  use  as  a  clue 
to  drumming  male  population.  Cover  analyses  were  made  of  10  drumming 
log  sites  by  the  method  of  Lindsey  et  al.  (16). 

Results  and  Discussion 

The  study  area  divided  itself  roughly  into  the  part  burned  over  in 
1951  and  the  unburned  hardwoods.  The  burn  is  an  interspersion  of  pines, 
field  edges,  and  open  canopy  hardwoods  with  a  great  deal  of  brush,  sap- 
lings, and  pole  timber.  The  unburned  area  is  predominantly  oak-hickory 
with  little  understory.  The  burned  hardwoods  in  the  southern  half  of 
the  area  obviously  held  more  grouse  than  the  unburned  portion.  Even 
though  grouse  were  not  uniformly  distributed  in  the  burn,  most  field 
work  was  concentrated  there.  In  general,  most  grouse  appeared  where 
brushy  vegetation  was  densest  and  along  ridge  tops  in  the  burn  and 
near  pine  plantations.  Ridge  tops  with  older  unburned  timber  were  not 
used  extensively  except  by  drumming  males. 

From  June  1968  to  April  1969,  250  grouse  were  flushed  in  the 
study  area.  Since  the  1968  estimated  fall  population  based  on  marked 
birds  was  42,  obviously  many  birds  were  flushed  more  than  once.  Al- 
though grouse  appeared  to  be  most  abundant  in  the  burned  section,  when 
Thurman  (25)  more  completely  searched  the  whole  area  by  randomized 
routes,  his  recorded  flushes  in  unburned  hardwoods  led  in  all  seasons 
except  fall.  In  the  present  study,  of  all  broods  flushed  in  summer,  86% 


Ecology  179 

came  from  burned  over  hardwoods,  the  rest  from  pines.  Slopes  and 
valley  bottoms,  as  opposed  to  ridge  tops,  produced  70%  of  brood  flushes. 
Temperature  and  humidity  data  from  valley  bottom  (670-foot  contour) 
and  ridge  top  (850-foot  contour)  showed  that  the  valley  had  a  wider 
range  of  temperature  and  was  more  humid  than  the  ridge.  The  ap- 
parent preference  of  grouse  for  the  lower  levels  in  summer  may  be  re- 
lated to  temperature  and  moisture  conditions  more  comfortable  to  the 
birds,  especially  as  the  same  conditions  associate  with  denser  vegetation 
affording  protection  and  food  supplies.  Crop  contents  from  four  summer 
juveniles  were  strong  in  occurrence  of  fungi  and  touch-me-not  (hnpatiens 
spp.),  both  abundantly  found  in  bottoms. 

In  September,  the  first  noticeable  difference  occurred  in  grouse  loca- 
tions with  birds  increasingly  found  on  ridge  tops.  During  the  period  of 
most  movement  in  the  first  week  of  October,  grouse  often  appeared  on 
roads  and  in  areas  where  previously  unseen.  The  October  movement  was 
most  frequently  away  from  dense  hardwood  thickets  of  slopes  and  bot- 
toms, the  high  use  area  of  summer,  to  edges  around  pine  plantations.  In 
October,  hunting  pressure  was  associated  with  increased  grouse  use  of 
pine  plantations,  the  densest  protective  cover  on  the  area.  Thurman  (25) 
also  noted  increased  use  of  pines  in  fall  and  winter.  In  midwinter,  grouse 
use  in  and  around  pines  remained  high,  but  they  also  spread  back  into 
dense  hardwood  vegetation,  often  feeding  there  on  the  abundant  crop  of 
sumac  (Rhus  spp.)  fruit.  Groups  of  grouse  were  more  commonly  seen  in 
winter  than  in  fall,  indicating  a  tendency  to  concentrate  at  feeding  sites. 

During  early  spring,  grouse  again  used  slopes  and  ridge  tops,  males 
moving  upward  to  more  mature  timber  to  establish  drumming  territories. 
Males  flushed  from  drumming  logs  showed  no  unusual  behavior.  Even 
those  flushed  from  logs  on  moonlit  nights  flew  normally.  They  had  no 
special  escape  routes  but  usually  flew  in  a  direction  away  from  the  ob- 
server. 

In  three  summer  months  of  1968,  a  total  of  80  flushed  grouse  gave 
nearly  an  even  adult:  juvenile  age  ratio  (38  juveniles,  37  adults,  5  un- 
known). Of  26  different  grouse  captured  from  July  to  October  inclusive, 
the  age  ratio  was  even.  This  is  unusual  as  a  healthy  post-breeding  popu- 
lation of  small  game  birds  should  include  60-80%  young  of  the  year  (1). 
In  1968,  broods  accompanying  hens  averaged  3.1,  substantially  less  than 
the  6-8  found  by  Thurman  (25)  on  the  same  area  in  1965.  Bump  et  al.  (3) 
reported  6-8  young  in  other  states.  Obviously,  1968  was  a  poor  year  for 
recruitment  of  ruffed  grouse  on  the  study  area.  A  few  reasons  of  circum- 
stantial nature  suggest  themselves. 

Rainfall  amounting  to  9.4  inches  occurred  in  14  days  in  May,  more 
than  twice  as  much  as  a  previous  100-year  average  for  the  period.  Ex- 
treme wet  conditions  at  the  time  when  ruffed  grouse  are  hatching  may 
cause  high  chick  mortality  (7).  To  underline  the  possibility  of  wet 
weather  mortality,  54%  of  summer  trapped  grouse  were  males.  There- 
fore, by  interpolation,  17  of  37  adult  grouse  flushed  in  summer  1968  may 
have  been  hens.  There  were  actually  eight  known  females  with  broods, 


180 


Indiana  Academy  of  Science 


leaving  a  possible  nine  broodless  hens.  Torrential  rains  could  easily 
destroy  whole  broods  of  young  chicks,  clutches  near  hatching,  or  decimate 
either  to  account  for  small  brood  size  and  broodless  hens  in  1968.  (A 
better  brood  year  probably  occurred  in  1969.  In  a  survey  for  cover 
mapping  on  a  nearby  area,  John  D.  Vanada  found  10  broods  during  the 
period  June  11-August  22.  They  averaged  6.9  young  per  brood  and  no 
broods  had  less  than  5  chicks). 

On  the  basis  of  recaptured  banded  grouse,  the  1968  fall  population 
of  the  study  area  was  estimated  at  42±14.8  birds  at  the  95%  confidence 
interval  (18).  Meager  data  make  this  only  an  approximation,  but  other 
methods  of  crude  estimating  give  about  the  same  number. 

Composition  and  arrangement  of  forest  ground  cover  and  overstory 
determine  ruffed  grouse  distribution  and  influence  productivity  (7).  Va- 
riety among  food  producing  plants  is  essential  to  counteract  irregulari- 
ties in  food  production  because  of  weather  or  alternate  seeding  years. 
Because  of  the  sedentary  nature  of  ruffed  grouse,  good  habitat  meets 
food  and  cover  requirements  within  a  relatively  small  area  (24).  Sum- 
mer brood  range  poses  the  most  critical  requirements.  Broods  must  have 
a  complex  understory  of  woody  and  herbaceous  plants  of  proper  height 


Table  1.  Vegetation  analysis  of  canopy  and  shrub  strata  for  ten  l/lO-acre  samples  at 
brood  locations  (1968)  and  ten  l/25-aere  samples  at  drumming  log  sites  (1969),  Geiger 
Ridge  Study  area.    (Only  the  10  highest  ranking  species  for  importance  value  are  shown 

in  each  case.) 


Relative 

Trees 

Stem  den- 

stem 

Basal 

Relative 

Relative 

Impor- 

and 

sity  pei- 

density 

area 

basal  area 

Fre- 

fre- 

tance 

shrubs 

acre 

per  acre 

per  acre 

per  acre 

quency 

quency 

value 

Brood 

Locations 

Hickory  spp. 

341 

9.1 

5.9 

22.6 

100 

3.5 

11.7 

Shortleaf    pine 

83 

2.1 

7.S 

30.3 

20 

0.7 

11.0 

Sumac  spp. 

783 

20.7 

100 

3.5 

8.6 

Black   oak 

i7<; 

4.6 

4.S 

IS.  5 

SO 

2.S 

8.6 

White   oak 

74 

1.9 

2.S 

L0.8 

90 

3.2 

5.3 

Sassafras 

450 

11.9 

0.08 

0.3 

100 

3.5 

5.2 

Ironwood     (Ostrya) 

203 

5.3 

90 

3.2 

4.2 

Sugar    maple 

101 

2.6 

1.5 

0.0 

100 

3.5 

4.0 

Dogwood 

269 

7.1 

0.08 

0.3 

100 

3.5 

3.6 

Hazelnut     ( Corylus ) 

L73 

4.5 

70 

2.5 

3.5 

Drumming  Log  Sites 

Black   oak 

272 

5.4 

20.4 

37.2 

100 

6.4 

16.3 

Sumac  spp. 

592 

11.9 

100 

6.4 

9.1 

Red    oak 

32 

0.5 

9.4 

17.1 

30 

1.9 

S.5 

Sassafras 

627 

12.6 

3.4 

6.2 

100 

6.4 

S.4 

Hickory  supp. 

462 

9.2 

3.1 

5.S 

100 

6.4 

7.1 

Dogwood 

460 

9.2 

L.5 

2.S 

90 

5.S 

5.9 

Grape    (Vitis) 

11)7 

3.9 

100 

6.4 

5.1 

Blue   beech    (Carpinus) 

i         345 

6.9 

50 

3.2 

5.0 

Ironwood 

215 

4.3 

90 

5.S 

5.0 

Tulip  tree 

( Liriodendron ) 

27 

0.5 

6.1 

11.1 

50 

3.2 

4.8 

Ecology  181 

and  density  to  allow  easy  movement  of  chicks  as  they  forage  (24).  The 
fact  that  grouse  have  persisted  on  the  study  area  without  any  special 
attention  indicates  that  the  range  is  meeting  at  least  the  minimum 
requirements  for  their  survival.  It  is  likely  that  extensive  interspersion 
of  woodland  successional  stages  on  Geiger  Ridge  similarily  provide  the 
mixture  of  types  found  necessary  for  ruffed  grouse  in  Missouri  (14). 

The  study  area  supported  a  minimum  of  8  broods,  usually  found 
in  relatively  open  canopy  situations  with  dense  underbrush.  In  a  com- 
parison of  basal  areas  of  different  species,  hickory,  black  and  white 
oaks,  sugar  maple  (Acer  saccharum  Marsh.),  and  shortleaf  pine  (Pinus 
echinata  Mill.)  dominated  the  overstory  of  brood  locations.  In  stems 
per  acre,  sumac,  sassafras  (Sassafras  albidiim),  hickory,  and  dogwood 
(Cornus  spp.)  led  in  that  order  of  understory  dominants  (Table  1). 
Ground  cover  consisting  of  grasses,  greenbrier  (Smilax  spp.)  tickclover 
(Desmodium  spp.),  cinquefoil  (Potentilla  spp.),  goldenrods  (Solidago 
spp.),  and  other  species  was  thinner  where  broods  were  found  than  in 
other  spots. 

As  noted  by  others  (2,  23),  the  grouse  brood  is  held  together  by  the 
hen's  vocalizations.  When  a  brood  is  flushed,  the  hen  typically  feigns 
injury,  attempting  to  decoy  the  intruder  away  from  the  hiding  young. 
After  a  more  or  less  intensive  display,  the  hen  flushes  to  rejoin  the 
brood,  responding  to  their  shrill  peeping  whistle  with  her  own  rasping 
peep  or  clucking.  The  intensity  of  the  bond  between  hen  and  chicks  ap- 
pears strongest  when  chicks  are  less  than  10  weeks  old  (up  to  July  30) 
and  gradually  weakens  as  they  mature.  This  was  shown  by  timing  the 
return  of  flushed  hens  to  their  broods  (Fig.  1).  By  September,  hens  did 
not  try  to  hold  broods  together,  and  at  this  time  break-up  of  broods  and 
wandering  of  some  individuals  occur. 

A  relatively  small  number,  26,  of  different  grouse  were  trapped  and 
marked  in  this  study.  Marked  grouse  were  recaptured  or  identified  by 
sighting  their  color  bands  11  times.  The  recovery  data  are  insufficient  to 
suggest  more  than  considerable  variation  in  the  tendency  to  move.  One 
juvenile  captured  in  July  was  recovered  in  August,  1,800  feet  away  in 
the  same  valley;  and  was  recovered  a  second  time  in  October,  1,600  feet 
from  the  last  site,  this  time  in  another  valley.  This  bird  was  with  a  brood 
on  the  first  two  occasions,  but  was  alone  the  last  time.  Another  mem- 
ber of  the  same  brood  was  recaptured  the  same  day  in  October  as  the 
first  juvenile  mentioned,  but  in  the  same  trap  where  it  was  originally 
caught.  These  records  from  two  individuals  in  the  same  brood  suggest 
that  young  grouse  move  up  and  down  a  valley  system  and  sometimes 
wander  into  other  valleys  as  brood  separation  occurs.  In  18  resightings 
of  marked  grouse  frcm  the  Geiger  Ridge  area,  Thurman  (25)  found  9 
juveniles  less  than  300  yards  between  points  of  observation,  although  a 
10th  was  recovered  2%  miles  away  and  the  11th,  5  Ms  miles  away. 
Adults  on  the  average  were  even  more  sedentary  than  the  shorter- 
ranging  juveniles.  Although  our  data,  and  those  of  Thurman  (25)  as 
well,  are  slight  by  comparison,  they  fit  the  general  conclusion  of  Hale 
and  Dorney  (11)  that  juvenile  ruffed  grouse  are  more  mobile  than  adults. 


182  Indiana  Academy  of  Science 

50n 


40  - 


c/>  30 
Id 

Z> 

i 


20- 


10 


Figure  1. 
broods. 


i   n      i      i      i      i      I      i     i      i 

16      23       30       7        14       21       28       4        II         18 
JUNE  JULY  AUGUST 

Time    required   for   ruffed   grouse    hens    flushed   with    broods    to    return    to    the 


In  this  respect,  Indiana  grouse  resemble  northern  birds.  The  phenomenon 
of  juvenile  mobility  implies  a  sexual  difference  as  discussed  by  Gullion 
(10)  on  the  basis  of  his  extensive  experience  with  Minnesota  grouse. 
Young  cocks  evidently  prefer  to  claim  territories  within  the  familiar 
brood  range,  whereas  young  hens  disperse  more  erratically  to  new 
habitats. 

Drumming  Males  and  Breeding  Populations 

During  the  last  week  of  March  and  the  first  week  of  April  1968, 
drumming  males  found  on  the  study  area  totaled  12  or  1  per  70  acres. 
In  1969,  the  count  was  15  or  1  per  56  acres.  Thurman's  (25)  count  for 
this  area  in  1966  was  25  males  or  1  per  32  acres.  Wise  (personal  com- 
munication) found  13  drumming  logs  on  an  835-acre  area  6  miles  from 
Geiger  Ridge.  Lewis  et  al.  (15)  estimated  2  grouse  per  100  acres  in 
Missouri,  half  of  them  males. 

The  number  of  drumming  males  on  an  area  from  one  year  to  the 
next  is  an  indicator  of  breeding  male  population  trends  although  not  a 


Ecology  183 

precise  one  (6).  If  a  total  census  of  drumming  males  can  be  obtained, 
it  is  often  assumed  that  a  sex  ratio  of  1:1  exists,  hence  the  total  spring 
population  is  at  least  twice  the  number  of  drumming  males.  This  is  a 
conservative  estimate  since  nondrumming  males  are  overlooked  and  since 
selective  predation  is  heavier  on  males  than  on  females  in  late  winter 
and  early  spring  (8).  In  spite  of  caution  recommended  in  estimating 
breeding  populations  or  trends  by  this  method,  the  difference  between 
Thurman's  25  drumming  males  and  Muehrcke's  15  appears  significant. 
As  discussed  above,  1968  apparently  was  a  poor  production  year. 

Further  evidence  of  a  reduced  1969  population  is  seen  in  the  dis- 
tribution of  drumming  males.  Thurman  found  13  drumming  territories 
in  the  unburned  hardwoods.  Muehrcke  found  only  2  there,  but  found  13 
territories  in  the  burned  section,  which  is  believed  to  be  the  better  grouse 
habitat.  In  1969  the  population  was  small  enough  to  allow  most  terri- 
torial males  to  establish  themselves  in  the  better  habitat.  In  view  of  the 
wet  weather  catastrophe  in  1968,  and  the  attachment  of  sexually  ma- 
turing males  to  their  brood  territories,  the  situation  also  suggests  better 
survival  of  young  in  the  burn. 

Drumming  Log  Sites 

Biologists  (9,  20)  have  observed  that  grouse  use  certain  logs  year 
after  year.  Gullion  (9)  called  them  perennial  logs.  In  the  present  study, 
Muehrcke  found  3  of  the  25  drumming  logs  identified  by  Thurman  in 
1966  still  used  in  1968.  Eight  of  12  logs  used  in  1968  were  also  used  in 
1969.  In  their  analyses  of  drumming  log  relationships,  neither  Palmer 
(20)  nor  Gullion  (9)  found  a  single  site  quality  factor  or  other  good 
explanation  for  the  phenomenon  of  traditional  use. 

Detailed  data  were  collected  for  the  physical  features  of  drumming 
logs  in  1969.  On  6  of  15  territories,  alternate  logs  were  also  used.  De- 
composition of  logs  varied  from  moderate  to  very  much.  Their  diameters 
ranged  9-16  inches,  averaging  12.8.  Total  length  ranged  8-38  feet,  averag- 
ing 23.2.  Distance  the  log  lay  from  ridge  top  usually  ranged  from  less 
than  10  to  more  than  600  feet,  although  4  logs  were  fairly  well  down 
slope.  Distance  from  field  edge  was  less  than  100  feet  except  those  on 
slopes  which  were  210  to  more  than  600  feet  away.  Hardy  (12)  located 
18  drumming  sites  on  his  eastern  Kentucky  study  area,  all  on  or  near 
ridge  tops;  and  Thurman  (25)  tabulated  the  exposure  of  25  sites  on  the 
Geiger  Ridge  area,  noting  that  some  were  on  ridge  tops  and  others  on 
slopes.  The  distance  of  the  latter  from  ridge  tops  is  not  on  record  but 
CMK  remembers  that  many  were  near  the  tops.  The  higher  levels  for 
display  sites  may  offer  some  survival  value  in  permitting  flushed  males 
to  fly  downward  quickly  into  denser  escape  cover.  Otherwise,  a  ridge 
top  location  may  play  a  part  in  natural  selection  to  the  extent  that 
sound  may  travel  farther  from  a  higher  than  a  lower  level,  and  hence 
attract  more  distant  hens. 

A  vegetation  analysis  of  drumming  log  and  brood  location  sites  is 
summarized  in  Table  1.  According  to  basal  areas  of  hardwood  species, 


184  Indiana  Academy  of  Science 

drumming  male  grouse  use  more  mature  timber  than  broods.  Otherwise, 
stems  per  acre  and  importance  values  show  the  essential  nature  of  young 
growth  and  shrub  species  in  both  habitats.  This  coincides  with  Palmer's 
(20)  observation  that  woody  vegetation  over  8  feet  high  is  more  dense 
near  drumming  sites  than  in  surrounding  cover.  We  believe  that  our 
grouse  generally  inhabit  dense  vegetation  in  the  valleys  during  inclement 
weather  of  spring,  and  that  males  move  upward  to  drumming  logs  in 
brushy  areas  dominated  by  older  trees. 

1969   Drumming  Season 

On  January  15  at  least  two  grouse  visited  drumming  logs  as  shown 
by  tracks  in  snow  and  droppings  around  logs.  Snow  cover  shortly  dis- 
appeared and  no  further  activity  around  logs  was  apparent  until 
February  24.  Droppings  showed  a  number  of  logs  visited,  but  undisturbed 
leaves  below  the  stages  (spots  on  the  logs  where  the  grouse  habitually 
stand  to  drum)  hinted  that  drumming  had  not  yet  begun.  First  drum- 
ming was  heard  February  28  after  a  week  of  mild  weather  when  tempera- 
tures ranged  from  a  high  of  41°  F  to  a  low  of  28°  F.  Drumming  activity 
then  increased  in  the  following  days  with  4  males  drumming  on  March  7, 
and  droppings  showing  that  other  males  were  visiting  their  logs.  From 
March  8  to  17  all  males  were  silent  during  which  time  temperatures 
ranged  from  10°  F  to  21°  F.  After  that  the  number  of  different  males 
drumming  increased  until  April  10  when  all  15  males  drummed. 

The  peak  of  drumming  intensity,  as  measured  by  the  largest  num- 
ber of  different  active  males,  extended  from  April  7  to  12  inclusive.  No 
less  than  8  and  as  many  as  15  of  the  15  different  males  performed  daily. 
Thurman  (25)  gives  the  first  week  in  April  for  this  intense  activity 
period  in  1966.  In  1969,  drumming  intensity  tapered  off  until  only  2  males 
were  heard  on  April  23.  No  observations  were  made  after  that. 

As  also  noted  by  Petraborg  et  al.  in  Minnesota  (22),  Muehrcke  (19) 
found  the  daily  duration  of  drumming  was  longest  in  the  period  of  peak 
activity,  with  drumming  beginning  earlier  before  sunrise  and  ending 
later  after  sunrise  as  the  peak  period  approached.  On  April  8  one  grouse 
started  drumming  at  0515  hours  EST,  the  earliest  time  noted,  and 
stopped  at  0614.  On  April  6,  grouse  drummed  sporadically  on  the  study 
area  until  1710  hours  exept  for  a  silent  period  from  1200  to  1410  hours. 
Usually  the  early  morning  drumming  period  ended  about  xk  hour  after 
sunrise,  but  some  males  returned  to  drum  intermittently  for  another  2 
hours.  Bent    (2)    also  reported  this  behavior. 

During  the  peak  period  of  drumming  activity,  Muehrcke,  from  a 
hide,  observed  one  cock  intensively  to  time  the  frequency  of  successive 
drumming  performances.  His  raw  data  are  less  extensive  and  precise  than 
those  of  Palmer  (21)  but  do  show  that  the  time  interval  between  drums  is 
shortest  just  before  and  just  after  sunrise.  The  raw  data  further  show 
that  the  intervals  were  shorter  when  a  second  grouse  is  present.  It  is  to 
be  expected  that  the  presence  of  a  hen  stimulates  an  increased  rate  of 
drumming    (8).   The   data   are   summarized   in  Table   2.   They   show  that 


Ecology  185 

drumming"  behavior  was  more  erratic  when  a  second  grouse  was  present; 
drumming  continued  longer  in  the  morning;  the  intervals  between  drums 
were  extended  as  the  cock  left  his  log  frequently  to  react  with  the 
visitor;  and  length  of  drums  ranged  up  to  3  sec  longer  than  when  the 
cock  was  alone. 

One  aberrant  cock  often  drummed  between  1500  and  1800  hours  in 
the  same  area  where  a  grouse  drummed  in  evenings  in  1988.  Gullion  (8) 
observed  that  some  males  are  predominantly  afternoon  or  evening  drum- 
mers. Night  drumming  apparently  occurs  commonly  (2).  On  calm,  moon- 
lit nights,  grouse  drummed  at  2200  hours  on  2  occasions,  and  at  0345 
on  3  occasions  on  Geiger  Ridge.  Thurman  (25)  also  heard  night  drum- 
ming on  the  area. 

Table  2.    Time  data  for  ruffed  grouse  drumming  performances  on  Geiger 
Ridge  study  area,  1969. 


Drumming  male 

Second  grouse 

was  alone 

was  present 

Date 

March  23 

April  4 

First  drumming 

0555  EST 

0535 

Last  drumming 

0655 

0727 

Number  of  drums 

24 

39 

Interval  between  drums 

Range  (minutes) 

1-5 

1-10 

Average 

2.3 

2.9 

Length  of  drums 

Range  (seconds) 

4-6 

3-9 

Average 

5.3 

5.3 

Only  certain  males  roosted  on  their  drumming  logs  and  even  then 
uncommonly  except  for  the  week  of  most  intensive  activity.  During  that 
7-day  period,  6  logs  were  used  1-3  nights  and  4  logs  were  used  4  nights, 
Muehrcke  actually  flushing  grouse  from  these  logs.  One  grouse  flushed 
from  a  drumming  log  at  2100  on  September  19.  More  intensive  field  work 
in  the  fall  would  probably  show  that  this  is  not  unusual.  Other  studies  (9) 
have  shown  that  many  drumming  logs  are  closely  attended  in  summer  as 
well  as  in  fall.  Young  males,  seeking  to  become  established  on  logs  in  fall, 
probably  cause  increased  activity  in  older,  established  males. 

Male  grouse  are  especially  wary  when  drumming  and  react  to 
unfamiliar  sounds  or  movement  by  remaining  silent  or  taking  flight. 
Grouse  leave  their  logs  by  walking  if  undisturbed,  or  by  running  or 
flushing  apparently  according  to  their  level  of  fright.  All  drumming  males 
habitually  performed  from  a  main  log  but  had  one  or  two  alternative  logs. 
At  least  7  males  drummed  on  alternate  logs.  If  a  bird  ran  from  its  main 
log  before  sunrise  during  the  peak  drumming  period,  it  invariably 
drummed  on  an  alternate  log  within  the  half  hour.  Males  flushed  from 
their  main  logs  would  not  use  alternate  logs  the  same  day  but  the  follow- 
ing day. 


186  Indiana  Academy  of  Science 

Literature  Cited 

1.  Allen,   D.   L.    1962.    Our  wildlife  legacy.    Funk  and   Wagnalls    Co.,   New   York.    421   p. 

2.  Bent,  A.  C.  1932.  Life  histories  of  North  American  gallinaceous  birds.  U.  S.  National 
Mus.  Bull.  162,  Washington,  D.  C.  490  p. 

3.  Bump,  Gardiner,  R.  W.  Darrow,  F.  C.  Edminster,  and  W.  F.  Crissey.  1947.  The 
ruffed  grouse — life  history — propagation — management.  New  York  State  Cons.  Dept. 
Albany.  915  p. 

4.  Chambers,  R.  E.,  and  P.  F.  English.  1958.  Modifications  of  ruffed  grouse  traps. 
J.  Wild!  Mgmt.  22:200-202. 

5.  Deam,    C.    C.    1940.    Flora   of    Indiana.    Dept.    of    Cons.,    Indianapolis.    1236    p. 

6.  Dorney,  R.  S.,  D.  R.  Thompson,  J.  B.  Hale,  and  R.  F.  Wendt.  1958.  An  evaluation 
of  ruffed  grouse  drumming  counts.  J.  Wildl.  Mgmt.  22  :35-40. 

7.  Edminster,  F.  C.  1954.  American  game  birds  of  field  and  forest.  Chas.  Scribner's 
Sons,  New  York.  490  p. 

8.  Gullion,  G.  W.  1966.  The  use  of  drumming  behavior  in  ruffed  grouse  population 
studies.  J.  Wildl.  Mgmt.  30:717-729. 

9.  Gullion,    G.    W.    1967.    Selection   and   use   of   drumming   sites   by   male   ruffed   grouse. 

Auk  84  :87-112. 

10.  Gullion,  G.  W.  1969.  The  ruffed  grouse  in  northern  Minnesota.  Processed  report. 
Forest   Research   Center,   U.   Minn.,   Cloquet.   20   p.  +  18   append. 

11.  Hale,  J.  B.,  and  R.  S.  Dorney.  1963.  Seasonal  movements  of  ruffed  grouse  in  Wis- 
consin. J.  Wildl.  Mgmt.  27:648-656. 

12.  Hardy,  F.  C.  1950.  Ruffed  grouse  studies  in  eastern  Kentucky.  Ky.  Div.  Fish  and 
Game,  Frankfort.  26  p. 

13.  Haymond,  Rufus.  1856.  Birds  of  southeastern  Indiana.  Proc.  Acad.  Natur.  Sci. 
VIII  :286-298. 

14.  Korschgen,  L.  J.  1966.  Foods  and  nutrition  of  ruffed  grouse  in  Missouri.  J.  Wildl. 
Mgmt.  30:86-100. 

15.  Lewis,  J.  B.,  J.  D.  McGowan,  and  T.  S.  Baskett.  1968.  Evaluating  ruffed  grouse 
reintroduction    in    Missouri.    J.    Wildl.    Mgmt.    32:17-28. 

16.  Lindsey,  A.  A.,  R.  O.  Petty,  D.  K.  Sterling,  and  Willard  Van  Asdall.  1961.  Vege- 
tation and  environment  along  the  Wabash  and  Tippecanoe  Rivers.  Ecol.  Monogr. 
31  :105-156. 

17.  Lindsey,  A.  A.,  W.  B.  Crankshaw,  and  S.  A.  Qadtr.  1965.  Soil  relations  and  dis- 
tribution   map   of  the  vegetation   of  presettlement   Indiana.    Bot.    Gaz.    126:155-163. 

18.  Mosby,  H.  S.  (editor).  1963.  Wildlife  investigational  techniques.  The  Wildlife  Society, 
Washington,  D.  C.  419  p. 

19.  Muehrcke,  J.  P.  1969.  Observations  on  ruffed  grouse  ecology  and  behavior  in  south- 
eastern Monroe  County,  Indiana.  Unpub.  master's  thesis,  Purdue  University,  Lafay- 
ette, Indiana.  98  p. 

20.  Palmer,  W.  L.  1963.  Ruffed  grouse  drumming  sites  in  northern  Michigan.  J.  Wildl. 
Mgmt.  27:656-663. 

21.  Palmer,  W.  L.  1969.  Time  frequency  between  successive  drumming  performances  of 
ruffed  grouse.  Wilson  Bull.  81  :97-99. 

22.  Petraborg,  W.  H.,  E.  G.  Wellein,  and  V.  E.  Gunvalson.  1953.  Roadside  drumming 
counts  a  spring  census  method  for  ruffed  grouse.   J.   Wildl.   Mgmt.   17:292-295. 

23.  Sawyer,  E.  J.  1923.  The  ruffed  grouse  with  special  reference  to  its  drumming. 
Roosevelt  Wildl.  Bull.  1  :355-386. 

24.  Sharp,  W.  M.  1963.  The  effects  of  habitat  manipulation  and  forest  succession  on 
ruffed  grouse.  J.  Wildl.  Mgmt.  27:664-671. 

25.  Thurman,  J.  R.  1966.  Ruffed  grouse  ecology  in  southeastern  Monroe  County,  Indiana. 
Unpub.  master's  thesis,  Purdue  University,  Lafayette,  Indiana.  101  p. 


Fox  Bounty  in  Indiana  During  the  Years  1961  through  1968 

Ralph  D.  Kirkpatrick,  Ball  State  University 


Abstract 

Fox  bounty  records  for  1961  through  1968  were  obtained  from  each  county  auditor 
in  Indiana.  The  number  of  Indiana  counties  paying  fox  bounties  declined  from  63  in 
1961  to  51  in  1968.  Numbers  of  fox  bountied  ranged  from  20,066  in  1963  to  23,863  in  1966. 
Fox  bounty  records  may  reflect  gross  fox  population  fluctuations.  Estimates  of  relative 
abundance  of  foxes  in  Indiana,  derived  from  the  annual  Sportsman's  Questionnaire  and 
the  annual  Furbuyer's  Report  are  compared  to  data  from  fox  bounty  records.  During  this 
8-year  period,  the  number  of  foxes  bountied  peaked  at  1.036  foxes  per  square  mile  in 
1966.  The  number  of  fox  pelts  purchased  by  Indiana  fur  buyers  also  peaked  in  1966  at 
0.258  pelts  per  square  mile.  Numbers  of  fox  pelts  purchased  annually  indicated  a  greater 
correlation  with  total  numbers  bountied  than  with  economic  value  of  pelts.  A  lesser 
degree  of  correlation  was  obtained  between  numbers  of  foxes  bountied  and  numbers  of 
hunter-killed  foxes  as  determined  by  the  annual  Sportsman's  Questionnaire. 

Laporte  County  paid  fox  bounties  from  1875  through  1877.  Fox 
bounties  were  not  paid  in  Indiana  again  until  1911.  One  or  more  counties 
have  expended  funds  for  fox  bounties  each  year  since  that  date  (2). 

Early  Indiana  fox  bounty  records  through  1948  are  given  and  dis- 
cussed by  Haller  (2).  Fox  bounty  records  for  the  years  1949  through 
1960  are  listed  and  discussed  by  Brooks  (1).  No  Indiana  state  agency 
compiles  fox  bounty  records  by  county  or  by  year.  Both  Haller  and 
Brooks  comment  on  the  difficulty  of  compiling  an  accurate  and  complete 
fox  bounty  record.  County  records,  including  fcx  bounty  information, 
tend  to  be  stored  in  courthouse  attics  or  basements  where  they  are 
occasionally  damaged  or  lost. 

Fox  bounty  records  for  each  Indiana  county  for  the  years  of  1961 
through  1968  were  secured  in  the  present  study.  Numbers  of  foxes 
bountied  were  compared  with  numbers  cf  fox  pelts  purchased  by  Indiana 
furbuyers  and  with  numbers  of  foxes  supposedly  killed  by  Indiana 
hunters. 


Methods 

Fox  bounty  records  for  the  years  1961  through  1968  were  secured 
by  a  personal  contact  with  every  County  Auditor's  Office  in  northeastern 
Indiana.  A  questionnaire  was  mailed  to  all  other  County  Auditors.  County 
Auditors  that  did  not  return  to  questionnaire  were  contacted  in  person 
or  by  telephone. 

Records  of  the  number  of  fox  pelts  purchased  by  Indiana  furbuyers 
and  average  fur  prices  were  secured  from  the  Annual  Furbuyers  Reports 
filed  with  the  Indiana  Department  of  Natural  Resources  (William  E.  Ginn, 
personal  communication). 

Estimates  of  annual  hunter  kill  of  fox  were  obtained  from  the 
Annual  Sportsman's  Questionnaire  compiled  by  the  Indiana  Department  of 
Natural  Resources  (William  E.  Ginn,  personal  communication). 

187 


188 


Indiana  Academy  of  Science 


Results  and  Conclusions 

The  number  of  counties  paying  fox  bounty  has  declined  from  63  in 
1961  to  51  in  1968,  with  the  majority  of  bounty-paying  counties  being 
located  in  northern  Indiana  (Table  1,  Fig.  1).  States  north  and  east  of 
Indiana  have  also  experienced  a  decline  in  the  number  of  counties  paying 


Table  1.    Number  of  foxes  bountied  in  Indiana  1961  through  1968. 


County 

1961 

1962 

1963 

1964 

1965 

1966 

1967 

1968 

Total 

Adams 

270 

17ti 

1  95 

218 

206 

242 

203 

252 

1,762 

Allen 

408 

808 

241 

444 

297 

8S1 

525 

390 

2,989 

Benton 

407 

318 

2S1 

310 

233 

306 

240 

346 

2,441 

Blackford 

224 

IBS 

174 

207 

205 

223 

184 

141 

1,526 

Boone 

584 

496 

546 

299 

347 

425 

00 

00 

2,647 

Carroll 

458 

467 

482 

42S 

377 

519 

442 

236 

3,354 

Cass 

308 

8X1 

500 

400 

49S 

500 

500 

500 

3,587 

Clinton 

62S 

421 

8S7 

36S 

321 

627 

407 

332 

3,491 

Decatur 

75 

89 

XS 

94 

1  05 

00 

00 

00 

451 

DeKalb 

454 

308 

810 

320 

321 

366 

352 

307 

2,741 

Delaware 

450 

405 

880 

271 

80S 

435 

306 

309 

2,814 

Dubois 

769 

767 

00 

00 

00 

00 

00 

00 

1,536 

Elkhart 

1S1 

228 

203 

2S3 

3S7 

422 

3S3 

366 

2,448 

Fayette 

133 

4X9 

250 

1S9 

192 

240 

207 

234 

1,934 

Fountain 

842 

8  IS 

227 

213 

1S3 

325 

390 

312 

2,310 

Fulton 

491 

8S0 

872 

416 

500 

S2S 

46S 

442 

3,897 

Grant 

850 

870 

876 

319 

351 

435 

271 

267 

2,739 

Hamilton 

400 

897 

824 

309 

301 

311 

2S4 

262 

2,588 

Hancock 

405 

842 

321 

236 

211 

361 

303 

299 

2,478 

Hendricks 

701 

500 

472 

391 

429 

579 

500 

493 

4,065 

Henry 

487 

857 

800 

196 

274 

404 

462 

425 

2,905 

Howard 

500 

4S0 

460 

430 

890 

3S0 

400 

223 

3,263 

Huntington 

844 

857 

2S8 

297 

273 

3S7 

270 

279 

2,490 

Jasper 

762 

658 

655 

599 

519 

672 

774 

543 

5,182 

.)  a  y 

281 

895 

812 

335 

221 

2S0 

243 

200 

2,217 

Jefferson 

820 

800 

801 

345 

00 

00 

00 

00 

1,266 

Jennings 

167 

100 

816 

263 

00 

00 

00 

00 

846 

Johnson 

00 

00 

00 

00 

226 

252 

2S3 

36S 

1,129 

Knox 

00 

00 

00 

170 

200 

220 

00 

00 

590 

Kosciusko 

494 

550 

5  OS 

677 

6S7 

S33 

594 

565 

4,908 

LaGrange 

248 

2S0 

2S9 

266 

264 

3S9 

413 

331 

2,480 

Lake 

479 

548 

400 

465 

313 

315 

292 

330 

3,137 

LaPorte 

5  t;t; 

624 

665 

696 

667 

7S3 

929 

732 

5,662 

Madison 

408 

488 

293 

323 

320 

389 

414 

307 

2,882 

Marion 

248 

287 

143 

22S 

97 

123 

171 

00 

1,247 

Marshalll 

47X 

857 

33S 

552 

471 

6S9 

54S 

517 

3,950 

Miami 

871 

821 

362 

347 

863 

42S 

3S0 

347 

2,919 

Monroe 

401 

2S0 

314 

395 

325 

500 

590 

520 

3,325 

Montgomery 

441 

504 

455 

350 

440 

00 

00 

00 

2,190 

Morgan 

425 

850 

242 

2S5 

266 

333 

330 

571 

2,802 

Newton 

682 

874 

43S 

46S 

353 

524 

371 

421 

3,631 

Noble 

490 

801 

4S6 

533 

672 

6S6 

471 

376 

4,015 

Ohio 

81 

87 

45 

65 

00 

00 

00 

00 

178 

Owen 

00 

00 

402 

399 

269 

00 

00 

00 

1,070 

Parke 

888 

81 S 

27S 

196 

213 

342 

331 

332 

2,343 

Porter 

00 

00 

00 

495 

500 

500 

400 

446 

2,341 

Posey 

840 

840 

00 

00 

00 

00 

00 

00 

680 

Pulaski 

600 

560 

496 

664 

551 

691 

612 

5 1  5 

4,689 

Ecology 

189 

Table 

.    Numb 

er  of  foxes  boun 

ied  in  Indiana  1961  thron 

gh  1968- 

-Contim 

ed. 

County 

1961 

1962 

1963 

1964 

1965 

1966 

1967 

1968 

Total 

Futnam 

425 

380 

380 

325 

396 

466 

466 

466 

3,304 

Randolph 

00 

00 

00 

00 

00 

00 

537 

442 

979 

Ripley 

460 

357 

329 

277 

00 

00 

00 

00 

1,423 

Rush 

391 

331 

295 

273 

298 

442 

333 

00 

2,363 

St.    Joseph 

200 

228 

232 

392 

300 

366 

352 

405 

2,475 

Scott 

L67 

174 

157 

85 

83 

17 

17 

00 

700 

Shelby 

442 

387 

328 

293 

268 

366 

442 

499 

3,025 

Starke 

2f>r> 

230 

215 

228 

298 

348 

246 

228 

2.048 

Steuben 

365 

222 

290 

374 

499 

500 

500 

655 

3,405 

Sullivan 

00 

00 

00 

00 

250 

300 

250 

00 

800 

Switzerland 

85 

77 

116 

00 

111 

00 

00 

00 

389 

Tippecanoe 

47(1 

301 

263 

36,7 

321 

398 

403 

332 

2,855 

Tipton 

369 

300 

274 

328 

300 

278 

268 

210 

2,327 

Union 

229 

250 

249 

223 

218 

245 

273 

00 

1,687 

Vermillion 

190 

174 

163 

195 

178 

206 

267 

266 

1,639 

Vigo 

284 

333 

358 

335 

313 

384 

332 

00 

2,339 

Wabash 

253 

377 

292 

391 

362 

400 

400 

325 

2,800 

Warren 

400 

299 

276 

176 

169 

194 

211 

194 

1,919 

Wells 

269 

336 

236 

302 

342 

405 

324 

306 

2,520 

White 

576 

472 

477 

423 

503 

428 

467 

405 

3,751 

Whitley 

1X9 

366 

326 

420 

385 

475 

359 

273 

2.793 

Total 

23,843 

22,168 

20,066 

20,751 

20,243 

23,863 

21,690 

18,842 

171.676 

Counties 

Paying 

63 

63 

62 

63 

60 

56 

54 

51 

69 

Bounty 

Mean  No. 

Bountied/ 

378.5 

348.7 

323.6 

335.9 

337.4 

426.1 

401.7 

369.5 

2,488.1 

County 

fox  bounty.  The  number  of  Ohio  counties  paying  fox  bounty  declined  from 
66  in  1961  to  52  in  1968  (5)  and  Michigan  stopped  paying  fox  bounty 
entirely  in  1965  (3), 

The  custom  of  fox  chasing  persists  in  southern  Indiana  but  has  prac- 
tically died  out  in  northern  Indiana  where  it  was  practiced  in  Grant 
County  as  recently  as  1931  (4).  Fox  chasers  have  discouraged  the  expendi- 
ture of  funds  to  pay  fox  bounty  in  southern  Indiana  counties.  A  reason 
for  the  discontinuance  of  bounty  payments  in  the  few  southern  counties 
that  appropriated  bounty  funds  is  the  import  of  dead  foxes  from  nearby 
counties. 

The  mean  number  of  foxes  bountied  per  Indiana  county  peaked  in 
1966  when  an  average  of  426.1  foxes  (or  1.036  foxes  per  square  mile) 
were  bountied  per  county  (Fig.  2). 

Brooks  (1)  reported  peak  Indiana  fox  bounty  payments  in  1926,  1934, 
1946,  1954,  and  1960.  It  appears  that,  if  bountied  foxes  actually  are  an 
index  to  total  fox  numbers,  the  Indiana  fox  population  fluctuates  with 
high  populations  occurring  each  6  to  8  years. 


190 


Indiana  Academy  of  Science 


•'H-H  I  I  I  I  M° 


PAID  BOUNTIES 
IN   1968 


•   •  ceased  paying 

o*#  BOUNTIES 

PRIOR  TO   1968 


Figure  1.     Map    of    Indiana    showing    counties    reporting    bounty    payments    during    the 
years  1961  through  1968. 


Fox  pelts  sold  may  reflect  fur  price  to  a  greater  extent  than  it 
reflects  fox  numbers,  but  Figure  3  indicates  that  this  is  not  true  during 
the  last  8  years  in  Indiana. 


Ecology  191 


1.5 

1.4 

1.5 

1.2 

1.1 

4) 
rH 

1.0 

a 

u 

CO 

cr 

CO 

0.9 
0.8 

U 
ft 

0.7 

CD 

X 

o 

0.6 

O 

0.5 

u 

1 

S25 

0.4 
0.5 

0.2 

0.1 

0 

\____ 


1961  1962  1965  1964  1965  1966  1967  1968 

••••fox  kill  by  hunters* 

11  ■  foxes  bountied 

—  —  fox  pelts  sold  to  furbuyers 

*  fox  kill  data  for  1968  is  not  available;  validity  of  kill 
data  for  I966  is  questionable 

Figure  2.     Comparison    of    three    measures    of    fox    population    levels    in    Indiana   for    the 
years  1961  through  1968. 


Bounty  amount  paid  per  fox  during  the  years  of  this  study  has  been 
$3.00  per  animal.  Brooks  (1)  points  out  that  the  amount  paid  per  fox  does 
not  directly  affect  numbers  of  foxes  bountied  per  year.  Fluctuations  in 
numbers  of  foxes  bountied  during  the  years  of  1961  through  1968 
parallel  fox  kill  data  supplied  by  the  Indiana  Sportsman's  Questionnaire 
(Fig.  2).    These  data  may  be  an  index  of  gross  populations  fluctuations. 


192 


Indiana  Academy  of  Science 


u 

CO 

C 
00 

1.10 
1.05 

u 

1.00 

a, 

CO 

0.95 

0.90 

T3 
■•-> 

c 
o 

0.85 
0.80 

<3 

0.75 

,0 

0.70 

53 

O.65 

6.5c 

6.00 
5.50 
5.00 
4.50 
4.00 
3.50 
3.00 
2.50 
2.00 
1.50 
1.00 

0.50 

0.00 


1961    1962    1963   1964    1965    1966    1967    1968 


foxes  bountied 
•  •  ••  price  paid  per  red  fox  pelt 
—  — price  paid  per  gray  fox  pelt 

Figure  3.     Comparison  of  average  prices  paid  for  fox  pelts  by   Indiana  fur  buyers  with 
numbers   of  foxes  bountied  per  square  mile  for  the  years   1961    through    1968. 


Literature  Cited 

1.  Brooks,   D.   M.    1962.   A   study  of  the  fox  population.    Ind.   Dept.    of   Cons.,   Wildl.    Res. 
Rep.  23(2)  :68-118. 

2.  Haller,   F.    D.    1950.    The  bounty  system   in   Indiana.    Ind.   Dept.   of   Cons.,   Wildl.   Res. 
Rep.  11(2)  : 93-125. 

3.  Laun,    C,   ed.    1968.    68   bounty  survey    (United   States).    Bounty   News,   2(1)  :n. p. 

4.  Leopold,    A.    1931.    Report   on    a   game   survey   of    the    North    Central    States.    Sporting 
Arms   and    Ammunition    Manufacturers'    Institute,    Madison,    Wisconsin.    299    p. 

5.  Russell,    K.    R.    1968.    Fox    bounty    payments    in    Ohio,    1944-1967.    Wildlife    In-Service 
Note  No.   97,   Ohio   Dept.   of  Natural   Resources,  Division  of   Wildlife.    10  p. 


Naturalized  Bi*j;  Trefoil   (Lotus  pedunculatus  Cav. )   Ecotypes 
Discovered  in  Crawford  County,  Indiana 

Maurice  E.  Heath,  Purdue  Agricultural  Experiment  Station 


A bstract 

In  1964,  eight  naturalized  ecotypes  of  big  trefoil,  (Lotus  pedunculatus  Cav.  ;  for- 
merly L.  ugilinosis  Schkuhr.,  L.  major  Sm.)  were  discovered  growing  in  a  tall  fescue 
(Festuca  arundinacea  Schreb. )  meadow  on  the  Clarence  E.  Kaiser  farm  in  Crawford 
County.  Subsequent  evaluations  show  three  of  the  ecotypes  superior  in  forage  and  vegeta- 
tive characteristics  for  use  in  tall  fescue  pastures  on  fragipan  soils  in  southern  Indiana. 
The  environmental  model  under  which  the  big  trefoil  ecotypes  have  become  naturalized 
is  described. 

Big  trefoil  (Lotus  pedunculatus  Cav.),  where  adapted,  is  a  long 
lived  perennial  forage  legume.  It  is  of  European  origin  and  is  a  recog- 
nized forage  plant  in  France,  Italy,  Denmark,  Germany  and  other 
countries. 

It  is  not  definitely  known  when  big  trefoil  was  introduced  into  the 
United  States.  However,  it  has  been  grown  in  this  country  for  over  95 
years  (5).  It  has  become  naturalized  in  western  Oregon,  western  Wash- 
ington and  northwest  California  where  it  is  used  for  pasture  and  hay.  It 
has  shown  good  adaptation  on  wet  soils  in  Georgia,  North  Carolina  and 
Florida  (3,4). 

In  1947,  Roland  McKee  (personal  communication),  Senior  Agrono- 
mist for  the  Bureau  of  Plant  Industry,  U.S.  Department  of  Agriculture, 
reported,  ".  .  .  some  work  with  Lotus  uliginosus  for  the  southern  part  of 
the  Plains  Corn  Belt  region  would  be  justified  as  I  am  inclined  to  believe 
that  by  selection,  a  sufficiently  hardy  strain  can  be  obtained  for  use  in 
combination  with  some  of  the  grasses."  Several  accessions  of  big  trefoil 
were  tested  on  the  Southern  Indiana  Forage  Farm,  Dubois  County,  Indi- 
ana, in  1954-55  but  failed  to  survive  the  winter. 


Ecotypes  Discovered 

On  August  13,  1964,  Mr.  Clarence  E.  Kaiser  showed  me  big  trefoil 
growing  in  eight  different  areas  in  a  vigorous  tall  fescue  (Festuca 
arundinacea  Schreb.)  field  on  his  farm  (Fig.  1).  Mr.  Kaiser  thought  the 
plants  to  be  a  particularly  adapted  form  of  birdsfoot  trefoil  (L.  cornicu- 
latus  L.)  since  the  areas  were  gradually  increasing  in  size.  Upon  exami- 
nation of  the  rhizomatous  and  stoloniferous  type  of  growth  and  spread- 
ing, I  immediately  recognized  these  plants  to  be  big  trefoil.  Mr.  Kaiser's 
farm  is  in  Crawford  County,  Indiana,  located  1  mile  east  and  1  mile 
south  of  the  junction  of  Indiana  Highways  164  and  145. 

193 


194 


Indiana  Academy  of  Science 


^IbsSPl 


-  -      -     #..*" 


Figure  1.  Naturalized  big  trefoil  (Lotus  pedunculatus  Cav.)  discovered  August 
growing  on  fragiyan  soils  on  Mr.  Clarence  E.  Kaiser's  farm  in  Crawford  County, 
Mr.  Kaiser  is  second  from  the  right. 


IS,   1964, 

Indiana. 


Mr.  Kaiser  believes  that  the  big  trefoil  was  a  contaminant  in  a 
grass-legume  seed  mixture  he  frost  seeded  over  the  field  in  the  late  winter 
of  1939.  The  seed  mixture  included: 

Rate  per  acre 
Species  in   pounds 

Birdsfoot  trefoil  (L.  corniculatus  L.)  1 

Ladino  clover  (Trifoliwm  repens  L.)  1 

Alsike  clover  (T.  hybridwm  L.)  1 

Timothy  (Phleum  pratense  L.)  4 

Smooth  bromegrass   (Bromus  inermis  Leyss.)  5 

Redtop  (Agrostis  alba  L.)  2 

In  1941,  five  pounds  per  acre  each  of  tall  fescue  and  orchardgrass 
(Dactylis  glomerata  L.)  seed  were  drilled  into  the  meadow.  Because  of 
the  excellent  adaptation  of  tall  fescue  the  field  is  now  a  heavy  fescue  sod. 


How  Field  Was  Managed 

Each  year  since  the  1939  seeding,  the  spring  growth  of  grasses  and 
legumes  has  been  harvested  for  hay.  The  summer  and  fall  growth  has 
been  used  for  beef  cow  pasture  from  late  September  following  calf 
weaning  until  late  January. 

From  1939  to  1951,  a  mid-summer  Ladino  clover  seed  harvest  was 
made  annually.  In  1964,  red  clover  (T.  pratense  L.)  was  frost  seeded  in 
late  winter.  The  red  clover  has  been  allowed  to  produce  and  shatter  seed 


Ecology 


195 


Figure  2.     Primary    area    of    adptation    of    tall    fescue     (Festuca    arundiacsa    Schreb.)     in 
the  east  humid  area  of  the   United  States  including   the  southern    third  of  Indiana    (1). 


on  the  ground  every  other  year.  Thus,  the  red  clover  has  behaved  as  a 
reseeding  biennial  legume  growing  in  the  tall  fescue  sod.  The  meadow 
has  been  fertilized  every  other  year  with  phosphorus  and  potassium 
according  to  soil  test  and  limed  as  needed. 

The  Environmental  Model  (2) 

The  hill  soils  on  Mr.  Kaiser's  farm  belong  to  the  Zanesville-Wellston 
soil  association  commonly  found  throughout  the  unglaciated  sandstone 
shale  region.  The  soil  fragipan  common  to  many  of  these  soils  restricts 
water  movement  and  root  penetration.  The  shallow  root  zone  depth  may 
range  from  18  to  34  inches.  Although  rainfall  averages  46  inches 
annually,  20  inches  are  lost  as  run-off,  and  droughts  are  common  in 
mid-summer.  The  soils  are  often  water-logged  in  winter  and  early 
spring.  Severe  winter-heaving  is  a  common  occurrence.  The  soil  moisture 
is  one  of  extremes  from  wet  to  dry.  Tap  rooted  alfalfa  is  considered  a 
high  risk  crop  on  these  soils  when  compared  to  tall  fescue  and  biennial 
red  clover.  The  native  pH  of  these  soils  ranges  from  5.3  to  5.5. 


Tall  Fescue  Adaptation 

The  southern  third  of  Indiana  is  in  the  primary  area  of  adaptation 
of  tall  fescue  (Fig.  2).  Since  1950,  tall  fescue  has  been  increasingly  grown 
on  the  fragipan  soils  of  southern  Indiana  where  it  currently  excells  all 
other  perennial  grasses  in  adaptation. 

It  is  believed  that  big  trefoil  may  have  its  greatest  promise  for  use 
in  southern  Indiana  as  a  perennial  pasture  legume  in  combination  with 


l  ;m; 


Indiana  Academy  of  Science 


1    ^v^iO.-.  '*      •  ., 


I'x.'- 


/.- 


FIGURE  3.     One  of  the  eight  ecotypes  of  big  trefoil  observed  to  be  associating  compatabily 
with    a   heavy  stand  of   tall  fescue  on   the  Kaiser  farm    (Photographed  June   17,    1966) 


Figure  4.  One  of  many  big  trefoil  nursery  plants  growing  on  fragipan  soils  on  the 
Southern  Indiana  Purdue  Agricultural  Center,  Dubois  County,  Indiana.  It  has  spread 
three  and  a  half  feet  through  a  good  stand  of  tall  fescue  during  a  period  of  three  growing 
seasons  without  evidence  of  any  heaving.  Tall  fescue  was  seeded  when  the  big  trefoil 
seedlings  were  transplanted  from  the  greenhouse  April  15,  1967  (Photographed  October 
3,  1969). 


Ecology  197 

tall  fescue  on  fragipan  soils  (Fig.  3).  Big  trefoil  appears  to  be  a  very 
efficient  supplier  of  readily  available  nitrogen.  Tall  fescue  has  a  dark 
green  vigorous  growth  during  the  spring  season  when  growing  with  big 
trefoil. 

Agronomic  Progress 
Approximately  15  two-inch  plugs  were  obtained  from  each  of  the 
eight  ecotypes  in  the  fall  of  1965  for  propagation  and  evaluation  pur- 
poses. These  were  planted  on  the  Southern  Indiana  Forage  Farm  in  the 
spring  of  1966.  Two  years  of  observations  showed  three  of  the  ecotypes 
to  be  superior  in  vigor  of  growth  and  seed  production.  A  total  of  10.3 
pounds  of  seed  of  the  3  ecotypes  were  produced  in  1968  for  experimental 
purposes  by  special  seed  production  facilities  of  the  Minnesota  Agricul- 
tural Experiment  Station  at  Rosemont,  Minnesota.  Seed  germination 
ranged  from  85  to  90%.  Excellent  stands  from  seeding  resulted  in  field 
plots  at  the  Southern  Indiana  Purdue  Agricultural  Center  (known  as  the 
Southern  Indiana  Forage  Farm  prior  to  1969)  in  the  spring  of  1969. 
Seed  produced  from  the  three  superior  ecotypes  (#1015,  #1016  and 
#1019)  equally  blended  together  will  be  known  as  'Kaiser'  trefoil  in  honor 
of  Mr.  Clarence  E.  Kaiser  who  introduced,  observed  and  nurtured  these 
plants  for  many  years  on  his  farm. 

Significance  of  Discovery  and  Summary 

1.  The  naturalized  ecotypes  of  big  trefoil  from  the  Clarence  E.  Kaiser 
farm  in  Crawford  County,  Indiana,  are  strongly  perennial  and  appear 
to  fit  the  environmental  model  of  the  hill  land  fragipan  soils  of  the 
unglaciated  sandstone  shale  soil  region  of  southern  Indiana. 

2.  Five  years  of  observation  have  not  shown  any  disease  or  heaving 
tendencies  among  the  eight  big  trefoil  ecotypes  discovered. 

3.  Several  of  the  big  trefoil  ecotypes  compete  aggressively  and  com- 
patibly with  tall  fescue  (Fig.  4)  which  is  the  dominant  perennial  grass 
of  the  unglaciated  sandstone  shale  soil  region. 

4.  The  big  trefoil  ecotypes  not  only  produce  seed  but  they  can  also  be 
readily  established  from  seed. 

5.  It  is  hypothesized  that  the  ruminant  response  from  the  tall  fescue-big 
trefoil  mixture  will  be  superior  to  that  of  the  ruminant  response  from 
tall  fescue  alone. 

6.  The  seed  produced  from  the  three  agronomically  superior  big  trefoil 
ecotypes  when  equally  blended  together  will  be  known  as  "Kaiser" 
trefoil  in  honor  of  Mr.  Clarence  E.  Kaiser  who  introduced,  observed 
and  nurtured  these  plants  for  many  years. 

Literature  Cited 

1.  Cowan,   J.   Ritchie.    1962.   Forages.   Iowa  State  University   Press,   Ames,   Iowa.    300  p. 

2.  Heath,  Maurice  E.  1969.  Fitting  plants  to  fragipan  soils  in  southern  Indiana.  Proc. 
Indiana  Acad.  Sci.  78  :429-434. 

3.  Henson,  Paul  R.,  and  H.  A.  Schoth.  1962.  The  trefoils — adaptation  and  culture. 
ITSDA-ARS  Agric.  Handbook  223. 

4.  McKee,  Roland  and  H.  A.  Schoth.  1941.  Birdsfoot  trefoil  and  big  trefoil.  USDA  Cir. 
625.  13  p. 

5.  McKee,  Roland.  1948.  Big  trefoil  proved  valuable  as  a  forage  crop  for  the  West  and 
South.  USDA-ARS  Res.  Achiev.  Sheet  101. 


The  Forest  Types  of  Indiana  and  a  New  Method  of  Classifying 
Midwestern  Hardwood  Forests 

Alton  A.  Lindsey  and  Damian  V.  Schmelz,i  Purdue  University 


Abstract 

Evidence  from  the  plotting  of  modified  importance  sums  for  3  species-groups  of 
dominant  forest  trees  on  a  3  axis,  triangular  graph  supported  the  classification  of  58 
outstanding  old-growth,  relatively  undisturbed  stands  in  Indiana  into  four  major  forest 
types :  Oak-hickory,  Beech-maple,  Lowland-depressional  and  Mixedwoods.  Two  clearly 
recognizable  subtypes  of  the  latter  were  well  drained  mixedwoods  versus  poorly  drained 
mixedwoods. 

Other  lines  of  evidence,  such  as  the  species  association  gradient  presented  herein, 
and  a  3  dimensional  ordinational  model  published  elsewhere,  corroborate  this  interpre- 
tation. 

This  classification  appears  applicable  to  the  hardwood  forests  of  all  nearby  mid- 
western  states  where  the  forest  stands  also  seem  to  be  dominated  in  different  sites  by 
essentially    the   same    3    general    species-groups,    and    intermediate   mixtures. 

The  Indiana  Natural  Areas  Survey  of  1967-9  provided  an  opportunity 
for  extensive  field  work  on  the  old  growth  forest  stands  of  the  state. 
Detailed  descriptions  of  many  such  stands  were  presented  in  the  book  on 
the  survey  results  (5).  The  best  36  forest  stands  (i.e.,  those  least  dis- 
turbed and  mest  nearly  in  equilibrium  with  their  specific  environment) 
were  used  for  ordination  and  an  intensive  comparison  of  forest  types. 
After  completion  of  this  work,  the  authors  jointly  developed  several  sim- 
pler, more  readily  applicable  approaches  for  the  classification  and  com- 
parison of  midwestern  hardwood  forests,  using  58  relatively  undisturbed 
stands,  including  the  36  stands  of  the  Schmelz  and  Lindsey  analysis  (7). 
The  best  one  of  these  approaches  appears  worthy  of  recommendation 
for  use  in  all  midwestern  states  where  3  essentially  similar  groups  of 
hardwood  tree  species  attain  high  importance  values;  it  therefore  forms 
the  basis  of  the  present  brief  paper.  The  method  depends  on  importance 
percentage  sums  of  the  respective  species-groups,  plotted  on  a  special 
triangular  graph.  It  does  not  require  tediovs  computations,  ordination 
or  the  construction  of  three-dimensional  models. 

The  potential  usefulness  of  the  present  method  is  not  restricted  to 
application  by  professional  forest  ecologists.  Because  of  current  and 
probable  continuing  interest  in  preservation  of  natural  areas  by  citizens' 
groups,  state  conservation  departments,  federal  agencies  and  others,  an 
accurate  but  relatively  simple  approach  to  comparison  and  classification 
of  forest  stands  should  aid  in  determining  priorities  for  official  protection. 

Methods  and  Materials 

The  58  forest  stands  we  used  (Table  1)  were  either  subjected  to  full 
tallies  of  substantial  portions  (up  to  23  acres  complete  census)  or  were 
intensively  sampled  by  a  number  of  Vs  acre  strips  or  %  acre  strips,  each 


Present  address:    Biology   Department,    St.   Meinrad    College,    St.    Meinrad,    Indiana. 

198 


Ecology  199 

400  feet  in  length.  The  unusually  thorough  gathering  of  basic  data  was 
prompted  by  the  demonstration  by  Lindsey,  Barton  and  Miles  (3)  that 
a  satisfactory  level  of  adequacy  depends  on  quite  high  intensity  of 
sampling. 

In  each  stand,  the  following  attributes  were  computed  for  each 
species  represented  by  individual  stems  exceeding  4  inches  dbh:  density 
per  acre  (D-),  relative  density  (D.f),  basal  area  per  acre  (BL.),  relative 
basal  area  (B.{)  and  importance  (V3).  The  latter  was  obtained  by  aver- 
aging relative  density  and  relative  basal  area  figures.  The  V3  for  separate 
selected  spe:i:s  within  a  given  stand  were  summed,  to  obtain  the  total 
importance  for  each  of  three  basic  species-groups  which  dominated  and 
characterized  stands  representing  particular  forest  types. 

The  oak-hickory  species-group  included  the  species  of  upland  Quercus 
and  Carya  characterizing  rather  xeric  forest  sites  in  Indiana,  i.e.,  white 
(Q.  alba),  chestnut  (Q.  prinus)  and  black  (Q.  velutina)  oaks,  and 
pignut  (C.  glabra)  and  shagbark  (C.  ovata)  hickories.  The  lowland- 
depressional  species-group  included  13  species  on  flood  plains  or  poorly 
drained  depressions — red  and  silver  maples  (Acer  rubrum,  A.  saccha- 
rinum) ,  big  shellbark  hickory  (C.  laciniosa) ,  hackberry  (Celt/is  occi- 
dentalis),  green  ash  (Fraxinus  pennsylvanica) ,  sweet  gum  (Liquidam- 
bar  styraciflua) ,  black  gum  (Nyssa  sylvatica) ,  sycamore  (Platanus 
occidentalis) ,  swamp  white  oak  (Q.  bicolor) ,  bur  oak  (Q.  macrocarpa) ,  pin 
oak  (Q.  palustris) ,  Shumard  oak  (Q.  Shiimardii)  and  American  elm 
(Ulmus  americana) . 

The  beech-maple  group  typical  of  upland  mesic  sites  consisted  of 
Fagus  grandifolia  and  Acer  saccharum. 

The  use  of  a  graph  in  the  form  of  an  equilateral  triangle  made  it 
possible  to  plot,  by  one  dot  for  each  stand,  the  selected  population  impor- 
tance sum  for  each  of  the  3  species-groups  in  a  single  plane.  This  graph, 
reproduced  as  Figure  1,  shows  values  for  the  beech-maple  species-group 
increasing  upward,  i.e.,  the  base  of  the  triangle  stands  for  zero  and  apex 
of  the  triangle  would  represent  100%  beech  and /or  sugar  maple.  How- 
ever, the  plotted  figures  cannot  be  simply  the  straight  importance  per- 
centage sums.  The  total  importance  for  a  stand  is  actually  1007r,  but 
we  disregard  some  species,  namely,  those  not  included  in  any  of  the 
three  species-group  lists  above.  Because  the  use  of  the  three-axis  graph 
depends  on  the  3  values  for  each  plotted  point  adding  up  to  100,  we  con- 
verted each  species-group  figure  to  a  new  per  cent  value,  on  the  basis  of 
a  full  100 %  for  the  sum  of  only  the  3  species-group  importance  figures. 
These   adjusted   values,   then,   were   plotted   for   Figure    1. 

Such  figures  for  the  oak-hickory  species-group  were  plotted  on  the 
axis  that  has  values  increasing  from  the  entire  right-hand  side  (zero  for 
OH)  to  the  lower  left  point  (100%  for  OH).  Conversely,  all  along  the 
left  side  is  the  zero  value  for  the  lowland-depressional  species  group 
component  of  a  given  stand,  while  this  component  increases  rightward 
and  downward  to  100%   LD  species  at  the  lower  right-hand  point.  Upon 


200 


Indiana  Academy  of  Science 


OAK-H  ICKO  RY 


An       BB  IS.LC.PI 
PO.SI.TI 


Figure  1.  Location  of  58  hardwood  forest  stands  on  a  three-axis  graph,  plotting  sums 
of  the  8  species-group  importance  percentages  per  stand  (see  text)  on  the  triordinates 
to  determine  the  stand  type.  The  60  per  cent  importance  level  (heavy  line)  delimits  the 
3  corners  where  beech-maple,  oak-hickory  and  lowland  depressional  stands  occur,  while 
the  stands  falling  centrally  represent  the  mixedwoods  type.  (Abbreviated  stand  names 
are  explained  in  Table  1.  In  several  cases,  one  dot  represents  more  than  one  closely 
similar  stand). 


this  3-axis  graph,  then,  we  plotted  the  3  major  components  of  each  of  the 
58  stands  selected  for  minimal  history  of  disturbance  by  human  activity. 

For  example,  the  Bear  Creek  Valley  slope  forest  (BV  in  Table  1) 
contained  no  lowland-depressional  species,  hence  its  dot  (BV)  fell  on  the 
zero  LD  line  that  forms  the  left  side  of  the  Figure  1  graph.  The  figures 
for  the  other  axes  were  approximately  20%  beech-maple  and  80 %  oak- 
hickory,  as  the  BV  dot  in  Figure  1  indicates. 


Results  and  Discussion 

Since  the  abovementioned  position  of  stand  BV  on  the  graph  closely 
approaches  the  oak-hickory  (lower  left)  point,  this  stand  doubtless  rep- 
resents the  oak-hickory  forest  type.  It  would  be  classed  as  an  oak- 
hickory  stand  by  the  criteria  of  Crankshaw  et  «/.  (2)  or  on  any  other 
basis  of  which  we  have  heard.  But  how  should  stand  BU  (with  the  same 
[20%]  beech-maple  percentage  but  only  63%   oak-hickory,  and  with  17% 


Ecology 


201 


Table   1.     Names,   symbols,   forest   types   and  locations   of   the   58   stands   considered   herein. 


County 

U.S.G.S.  quad 

Town 

Range 

Section 

BEECH-MAPLE 

Allee   (Al) 

Parke 

Montezuma 

16N 

xw 

3 

Bendix  (Bx) 

St.  Joseph 

Lydick 

37N 

1W 

11 

Caster  (Ca) 

Montgomery 

Shannondale 

19N 

3W 

34 

Cring  (Cr) 

Jay 

Portland 

23N 

14E 

10 

Hayes  (Ha) 

Wayne 

New  Paris 

14N 

1W 

35 

Hoot  (Ho) 

Owen 

Patricksburg 

9N 

4W 

6 

Jackson   (Jk) 

Ripley 

Milan 

7N 

12E 

18 

Logansport  (Lo) 

Cass 

Anoka 

27N 

2E 

33 

Manlove  (Mu) 

Fayette 

Connersville 

15N 

12E 

29 

Meltzer  (Crosby)    (MC) 

Shelby 

Rays  Crossing 

12N 

8E 

7 

Nature  Conservancy   (NC) 

Montgomery 

Alamo 

17N 

6W 

2 

Officer's  North  (ON) 

Jefferson 

Volga 

4N 

9E 

22 

Pine  Hills   (  PH  ) 

Montgomery 

Alamo 

17N 

6W 

1 

Pioneer  Mothers   (PM) 

Orange 

Paoli 

IN 

IE 

7 

Potzger  (Po) 

Ripley 

Milan 

7N 

12E 

20 

Rocky  Hollow   (RH) 

Parke 

Wallace 

17N 

7W 

27 

Rosbrugh  (Ro) 

Kosciusko 

Leesburg 

33N 

BE 

30 

Rush   (Ru) 

Montgomery 

Alamo 

18N 

6W 

27 

S.  LaPorte   (SL) 

LaPorte 

LaPorte  East 

36N 

3W 

2 

Spurgeon   (Sp) 

Noble 

Ligonier 

35N 

9E 

18 

Warren   (Wr) 

Berrien   (Mich) 

Three  Oaks 

7S 

1W 

27 

Weaver  (Wv) 

Fayette 

Connersville 

15N 

12E 

30 

Wygant   (Wy) 

Huntington 

Majenica 

27N 

10E 

3 

OAK-HICKORY 

Beall   (Upland)    (BU) 

Wabash  (111.) 

Mt.  Carmel 

2S 

13W 

11 

Bear  Creek  Plateau   (BP) 

Fountain 

Stonebluff 

21N 

8W 

33 

Bear  Creek  Valley   ( BV ) 

Fountain 

Stonebluff 

20N 

8W 

4 

Dunes  (Xeric)    (DX) 

Porter 

Dune  Acres 

37N 

<;w 

L3 

Fox  Island  (FI) 

Allen 

Fort  Wayne  W. 

30N 

he 

25 

Johnson   (Jo) 

Posey 

Wabash  Is. 

17S 

14W 

32 

Lilly-Dickey   (Ly) 

Brown 

Nashville 

9N 

3E 

8 

Ross  Reserve   (RR) 

Tippecanoe 

Otterbein 

23N 

tiW 

26 

Wing  Haven   (Wi) 

Steuben 

Angola  E 

38  N 

13E 

35 

LOWLAND-DEPRESSIONAL 

Andrus   (An) 

Knox 

E.  Mt.  Carmel 

IS 

12W 

11 

Beall   (Bottom)    (BB) 

Wabash  (111.) 

Mt.  Carmel 

2S 

13  W 

11 

Beckville  (Bk) 

Montgomery 

Shannondale 

18N 

3W 

11 

Davis  1   (Dv) 

Randolph 

Redkey 

21N 

12E 

23 

Giants  (WG) 

Vermillion 

Perrysville 

18,   19N 

9W 

3,  34 

Hemmer  (Hm) 

Gibson 

Lynnville 

3S 

9W 

24 

Independence   (IB) 

Tippecanoe 

Otterbein 

22N 

i;w 

3 

Kramer  (Kr) 

Spencer 

Owensboro  W. 

SS 

7W 

12 

Little  Cypress  (LC) 

Knox 

E.  Mt.  Carmel 

IS 

12W 

14 

Meltzer  (Brookston)    (MB) 

Shelby 

Ray's  Crossing 

12N 

8E 

7 

Paramecium  Is.   (PI) 

White 

Buffalo 

28N 

3W 

1 

Pin  Oak   (PO) 

Gibson 

E.  Mt.  Carmel 

IS 

12W 

34 

Sigmoid  Is.   (SI) 

Carroll 

Brookston 

25N 

3W 

15,   16 

Terrace  Is.   (TI) 

Tippecanoe 

Brookston 

24N 

3W 

17 

Tippecanoe   (Upper)    (TU) 

Pulaski 

Bass  Lake 

31N 

1W 

18,   19 

Wesselman   (Ws) 

Vanderburgh 

Evansville 

6S 

10W 

22,  23 

MIXED  WOODS 

Big  Walnut  (BW) 

Putnam 

Roachdale 

15N 

3W 

29,  31,  32 

Botany  Glen   (BG) 

Grant 

Gas  City 

23N 

8E 

11,   14 

Bradford  (Br) 

Morgan 

Mooresville  W 

13N 

IE 

r> 

Clifty   (CI) 

Montgomery 

Alamo 

17N 

<;w 

l 

Conboy  (Cy) 

Jennings 

Vernon 

6N 

9E 

10 

Donaldson's  (Dn) 

Lawrence 

Mitchell 

3N 

IE 

4 

Dunes  (Mesic)    (DM) 

Porter 

Dunes  Acres 

37  N 

6W 

13 

McCormicks  Cove  (McC) 

Owen 

Gosport 

ION 

3W 

22 

Officer's  (South)    (OS) 

Jefferson 

Volga 

4N 

9E 

22 

Tippecanoe  (Lower)    (TL) 

Pulaski 

Bass  Lake, 
Winamac 

31N 

1W 

18,   19 

202  Indiana  Academy  of  Science 

lowland-depressional),  be  typed?  An  inspection  of  the  distribution  of  all 
the  stands  on  the  graph  aids  in  establishing  improved  criteria  for  stand 
classification. 

The  58  original  or  very  old  growth  stands  utilized  do  not  appear  as 
a  continuum  throughout  Figure  1  since  intergradation  between  the  two 
sides  (OH  versus  LD)  appears  lacking  not  only  along  the  zero  BM  (base) 
line  but  throughout  a  large  central  region  and  high  up  toward  the  BM 
apex.  The  intergradations  can  be  traced  instead  from  LD  with  high 
available  moisture  supply,  through  BM  with  median  moisture  and  drain- 
age (mesic  sites),  to  OH  with  limited  moisture  and  excessive  drainage. 
Although  these  old  stands  must  at  present  be  relatively  stabilized  and  in 
balance  with  their  sites  and  overall  environments  in  accord  with  current 
polyclimax  interpretation,  reading  the  graph  upward  from  either  the  OH 
or  LD  corner  parallels  the  successional  trend  in  Clementsian  theory  from 
moisture  extremes  to  optimum  conditions  for  the  beech-maple  type  on 
mesic  sites.  This  conforms  with  the  results  shown  by  the  much  less  direct 
approach  (through  3-axis  ordination  technique)  of  Schmelz  and  Lindsey 
(7)  based  on  a  more  stringently  selected  36  stands  of  these  58. 

An  area  of  some  selected  size  subtending  each  point  of  the  triangle 
will  serve  to  delimit  or  define  the  major  forest  types  for  Indiana  in  a 
revised  type  classification.  Any  criteria  adopted  are  somewhat  arbitrary, 
as  were  those  of  Crankshaw  et  al.  (2)  and  Lindsey  et  al.  (4)  in  earlier 
papers.  The  present  authors  consider  that  the  information  in  Figure  1 
supports  recognition  of  4  major  hardwood  forest  types,  when  each  "point" 
is  delimited  at  the  60%  cross  line  of  the  graph,  as  shown  by  heavier 
lines.  The  3  forest  types  occupying  the  extreme  site  conditions,  near  the 
points  of  the  triangle  (Lowland-depressional,  Beech-maple  and  Oak- 
hickory)  are  defined  by  having  at  least  a  60  (adjusted)  importance  per- 
centage sum  for  the  one  dominant  species-group,  hence  40%  or  less  for 
the  other  two  species-group  sums  taken  together.  Stand  BU,  having  63% 
upland  OH  species,  is  therefore  placed  in  the  oak-hickory  type.  Obviously, 
the  three  importance  sums  are  sufficient  to  designate  forest  types  (with- 
out plotting  stand  positions  on  a  graph)  once  a  quantitative  criterion  is 
accepted.  The  graph  facilitates  broad  comparisons. 

Stands  that  are  not  conspicuously  dominated  by  a  single  species- 
group  fall  within  the  remaining  central  hexagonal  figure.  We  term  this 
hardwood  type  as  Mixedwoods.  This  fourth  type  is  subdivided  rather 
clearly  (within  the  stands  examined,  at  least)  into  well  drained  and 
poorly  drained  subtypes,  at  the  left-center  and  right-center  respectively, 
and  without  intergradation  between  them  within  the  Mixedwoods  Type. 
The  latter  includes,  basically,  both  the  Mixed  Woods  and  Western  Meso- 
phytic  Types  of  Schmelz  and  Lindsey  (7)  (both  these  types  representing 
the  well-drained  Mixedwoods  subtype)  plus  the  poorly-drained  mixed- 
woods  subtype  represented  here  by  TL,  OS  and  Cy  at  the  right.  The 
latter  show  very  low  oak-hickory  representation,  because  low-ground  oaks 
like  pin  and  swamp  white  oaks  are  not  included  in  the  (upland)  oak- 
hickory  species-group.  However,  these  3  stands  have  moderate  amounts 
of  beech  and /or  sugar-black  maple;   this  raises  them  above  the  lowland 


Ecology  203 

depressional  type,  into  the  Mixedwoods  Type,  as  the  subtype  of  the  latter 
which  lies  intermediate  between  Beech-maple  and  Lowland-depressional 
rather  than  between  Beech-maple  and  Oak-hickory. 

Since  about  87%  of  Indiana  was  covered  by  forest  in  presettlement 
times  (6),  all  substrate  conditions  paralleling  the  forest  type  triangle  in 
Figure  1  probably  supported  forests  in  this  state  originally.  The  large 
central  blank  space  appearing  on  the  graph  represents,  by  and  large, 
the  vegetation  of  sites  without  extremes  of  ponding  or  droughtiness, 
steep  slope  or  excessive  internal  drainage.  Since  these  median  sites  have 
been  chiefly  cleared  for  agricultural  utilization,  fine  present-day  forests 
tend  to  be  restricted  to  sites  less  favorable  for  farming.  It  seems  reason- 
able that  mixedwoods  were  somewhat  more  prevalent  in  the  original 
vegetation  of  Indiana  than  in  that  of  today,  not  so  much  as  an  extensive 
mappable  type  as  on  some  isolated  median  sites  within  the  regions 
mapped  (4,  5,  6)  as  beech-maple  or  oak-hickory.  The  portions  of  southern 
Indiana  mapped  previously  (4,  5,  6)  as  "western  mesophytic"  we  now 
prefer  to  consider  Mixedwoods,  in  part  because  Aescuhis  octandra  and 
Tilia  heterophylla  which  Braun  (1)  considered  mesophytic  indicators  do 
not  extend  very  far  northward  in  Indiana. 

Data  on  soil  requirements-tolerances  of  certain  Indiana  tree  species 
were  published  by  Crankshaw,  Qadir  and  Lindsey  in  1965  (2).  We  have 
arranged  those  species  according  to  the  ecological  affinities  shown  in 
Table  2  into  a  gradient  from  black  oak  at  the  xerophytic  end  through 
black  ash  at  the  hydrophytic  extreme.  They  fall  clearly  into  the  3 
groups  roughly  corresponding  with  the  3  apical  regions  of  our  Figure  1 
graph  and  with  the  3  extreme  positions  on  the  ordination  model  (of  36 
stands)  published  py  Schmelz  and  Lindsey  (7).  The  upland  oak-hickory 
type  (upper  third  of  Table  2)  prefers  high  per  cent  sand,  whereas 
high  silt  content  characterizes  the  sites  of  the  dominant  tree  species  of 
beech-maple  forests,  and  high  per  cent  clay  favors  lowland-depressional 
stands.  Highly  leached  soils,  low  pH  and  low  available  nitrogen  are 
compatible  with  oak-hickory.  The  lowland  depressional  species  are 
favored  by  high  available  water,  neutrality  of  reaction,  high  clay  con- 
tent and  at  least  4  of  them  by  high  per  cent  nitrogen  in  the  soil. 


204 


Indiana  Academy  of  Science 


Table  2.  Gradient  of  influence  of  the  most  important  soil  factors  on  tree 
species  in  the  presettlement  Indiana  forest.  Plus  sign  indicates  high  value 
favors  species  under  natural  competition.  Minus  sig?i  indicates  that  low 
value  is  favorable  or  well  tolerated.  A  reworking  of  certain  data  from 
Crankshaw,  Qadir  and  Lindsey  (2). 


Avail. 

Leach- 

% 

% 

% 

'/< 

H,0 

ing 

pH 

N 

Sand 

Silt 

Clay 

Black  Oak 



+ 





+ 

Chinquapin  Oak 

— 

+ 

— 

+ 

White  Oak 

6.1 

— 

+ 

— 

W.  B.  Cherry 

— 

6.1 

+ 

— 

Shingle  Oak 

— 

+ 

Post  Oak 

+ 

— 

— 

— 

Upl.  Hickories 

+ 

— 

— 

+ 

Red  Oak 

+ 

Tulip  Tree 

+ 

+ 

Sugar  Maple 

+ 

+ 

+ 

Basswood 

+ 

— 

+ 

+ 

White  Ash 

— 

< » 

+ 

+ 

Beech 

< > 

— 

+ 

Bur  Oak 

+ 

+ 

Sweet  Gum 

+ 

— 

+ 

Amer,  Elm 

7 

— 

+ 

Bl.  Walnut 

+ 

7 

+ 

Buckeye 

+ 

+ 

+ 

• 

+ 

Hackberry 

+ 

0 

7 

+ 

— 

Cottonwood 

+ 

0 

7 

+ 

Honey  Locust 

+ 

< — > 

7 

< > 

Sycamore 

+ 

7 

+ 

Black  Ash 

+ 

— 

7 

+ 

+ 

Literature  Cited 

Braun,  E.  Lucy.   1950.  Deciduous  forests  of  Eastern  North  America.   Hafner  Pub.   Co., 

New  York.  610  p. 

Crankshaw,   W.  B.,   S.   A.   Qadir,  and  A.   A.   Lindsey.    1965.   Edaphic  controls  of  tree 

species  in  presettlement  Indiana.  Ecology  46  :688-698. 

Lindsey,  A.   A.,  J.  D.   Barton,  Jr.,  and  S.    R.   Miles.   1958.   Field  efficiencies  of  forest 

sampling  methods.  Ecology  39  :428-444. 

— ,  W.  B.  Crankshaw,  and  S.  A.  Qadir.  1965.  Soil  relations  and  distribution 
map  of  the  vegetation  of  presettlement  Indiana.   Bot.  Gaz.  126:155-168. 

— ,  D.  V.  SCHMELZ,  and  S.  A.  Nichols.  1969.  Natural  Areas  in  Indiana  and 
their  Preservation.  Indiana  Natural  Areas  Survey,  Purdue  University,  Lafayette. 
606  p. 

Petty,  R.  O.,  and  M.  T.  Jackson.  1966.  Plant  communities,  p.  264-296.  /»  A.  A. 
Lindsey  |ed.  1  Natural  Features  of  Indiana,  Indiana  Sesquicentennial  Volume.  Indiana 
Acad.  Sci.,  Indianapolis.  630  p. 

Schmelz,  D.  V.,  and  A.  A.  Lindsey.  1970.  Relationships  among  the  forest  types  of 
Indiana.  Ecology    (in  press). 


A  Survey  of  the  Commercially  Valuable  Mussels  of  the 
Wabash  and  White  Rivers  of  Indiana1 

Louis  A.  Krumholz,  Roy  L.  Bingham,-'  and  Edward  R.  Meyers 
University  of  Louisville 

Abstract 

The  shells  of  several  unionid  mussels  of  the  Wabash  River  system  in  Indiana  are  of 
considerable  value  in  the  manufacture  of  nuclei  for  cultured  pearls  in  Japan.  Because  of 
increased  exploitation  of  that  fauna  for  commercial  purposes,  the  Indiana  Department  of 
Natural  Resources  sponsored  a  survey  to  determine  the  status  of  the  mussel  populations 
over  about  500  stream  miles  of  the  Wabash,  White,  and  East  Fork  of  the  White  rivers, 
and  to  gather  data  upon  which  to  base  recommendations  for  the  preservation  of  that 
resource. 

The  survey,  conducted  by  the  University  of  Louisville,  extended  through  1966  and 
1967  and  included  99  collections  made  either  with  a  crowfoot  bar  or  by  diving  and  hand- 
picking  with  or  without  auxiliary  air  supply.  In  those  collections,  30  species  of  unionids 
were  represented,  but  only  10  were  considered  important  in  the  commercial  market; 
representatives  of  those  10  species  made  up  77.1%  of  the  total  catch  of  the  survey.  The 
most  abundant  mussel  in  the  commercial  market  as  well  as  in  the  survey  collections  was 
the  mapleleaf,  Quadrula  quadrula  (Rafinesque).  Data  from  the  survey  as  well  as  from  the 
commercial  market  indicated  that  the  mussel  stocks  in  the  rivers  were  depleted  seriously 
in  1966,  mostly  through  the  very  efficient  efforts  of  divers  with  auxiliary  air  supply. 
Those  data  led  to  the  passage  of  Discretionary  Order  No.  136  by  the  Indiana  Department 
of  Natural  Resources  prohibiting  the  use  of  such  gear  in  commercial  musseling. 

Mussels  of  commercial  value  grow  very  slowly ;  it  requires  from  4  to  5  years  for  a 
mapleleaf  mussel  to  reach  the  legal  size  of  2.5  inches.  The  serious  depletion  of  breeding 
stocks  in  1966  probably  will  result  in  a  very  low  yield  of  marketable  shells  at  least  until 
1970  and  perhaps  later. 

Introduction 

Freshwater  bivalve  mussels  of  the  family  Unionidae  have  been  in 
existence  since  early  Mesozoic  times  in  the  lakes  and  rivers  of  the  north- 
ern temperate  and  subtropical  areas  of  the  world  (29).  At  present, 
there  are  between  500  and  600  living  species  of  unionids  within  the  con- 
tinental United  States  (10),  and  of  these,  several  are  commercially  valu- 
able because  of  the  quality  of  the  material  in  their  shells.  Although  the 
soft  parts  of  these  mussels  are  edible,  no  commercial  market  has  ever 
been  established  for  freshwater  clams  comparable  to  that  for  marine 
clams. 

The  shells  of  the  freshwater  mussels  have  become  valuable  at  two 
different  times  in  recent  history  of  the  United  States,  for  two  quite  dif- 
ferent reasons.  During  the  last  decade  of  the  nineteenth  century,  the 
pearl  button  industry  was  established  at  Muscatine,  Iowa,  by  J.  P. 
Boepple,  a  German  button  cutter  (8).  That  industry  flourished  for  about 


1  Contribution  No.  126  (New  Series)  from  the  Department  of  Biology,  University  of 
Louisville,  Louisville,  Kentucky  40208. 

J  Present  Address:  Department  of  Oceanography,  Texas  A&M  University,  College 
Station,  Texas  77843. 

3  Present  Address  :  Department  of  Zoology,  Arizona  State  University,  Tempe,  Arizona 
85281. 

205 


206  Indiana  Academy  of  Science 

50  years,  but  by  the  beginning  of  the  twentieth  century,  the  mussel  beds 
in  the  Mississippi  River  began  to  show  signs  of  depletion   (28). 

Even  though  the  use  of  mussel  shells  in  the  manufacture  of  pearl 
buttons  essentially  disappeared  in  1953,  new  demands  were  put  upon  the 
mussel  populations  by  the  cultured  pearl  industry. 

Freshwater  mussel  shells  from  rivers  in  the  United  States  are  used 
in  the  manufacture  of  the  spherical  nucleus  that  is  implanted  in  the 
oyster  by  the  Japanese.  Shells  from  the  Wabash  River  and  Tennessee 
River  systems  in  the  United  States  have  an  extraordinary  translucence 
that  makes  them  particularly  valuable  as  nuclei.  The  diameter  of  the 
spherical  nucleus  prepared  from  mussel  shells  usually  ranges  from  Vs  to  hi 
inch,  but  may  be  even  larger  on  occasion.  In  preparing  the  nuclei,  blemish- 
free  mussel  shells  are  cut  into  strips,  then  into  squares,  and  then  into 
cubes.  The  cubes  are  placed  between  rotating  discs  similar  in  shape  to 
the  stone  of  a  grist  mill,  together  with  a  slurry  of  abrasive  material.  The 
discs  are  rotated  and  the  nuclei  are  worn  down  until  they  become  spheri- 
cal. At  this  point  the  spheres  are  selected  on  the  basis  of  their  perfection 
and  are  placed  within  the  visceral  mass  of  the  living  oyster.  The  yield 
of  cultured  pearls  of  superior  quality  using  such  nuclei  is  very  high.  The 
time  required  for  a  cultured  pearl  to  be  formed  ranges  from  1  to  5  years, 
depending  on  the  desired  thickness  of  the  nacreous  covering  on  the 
nucleus  (27). 

Persons  associated  with  the  mussel  industry  realized  that  the  mussel 
supply  is  exhaustible  and  efforts  were  made  to  conserve  this  valuable  nat- 
ural resource.  In  April  1967,  the  Indiana  Department  of  Natural  Resources 
passed  Discretionary  Order  No.  136  which  restricted  the  methods  for 
taking  mussels  for  commercial  purposes  to  handpicking,  short  forks, 
tongs,  or  brail  (crowfoot)  bar.  The  use  of  mechanical  dredges  and  diving 
with  auxiliary  air  supplies  was  prohibited. 

Because  of  the  very  great  demand  for  mussel  shells  of  high  quality 
in  the  cultured  pearl  industry,  the  beds  of  commercially  valuable  mussels 
in  the  Tennessee  River  have  been  virtually  wiped  out.  Similarly,  the 
mussel  beds  of  the  Wabash  River  system  are  in  danger  of  serious  deple- 
tion. In  many  instances,  populations  of  desirable  species  of  shells  have 
been  extirpated.  However,  it  should  be  pointed  out  that  the  cultured  pearl 
industry  and  their  demands  on  mussel  shells  of  the  Wabash  River  is  not 
the  sole  factor  operating  in  the  depletion  of  those  shells.  Pollution, 
whether  it  be  industrial,  domestic,  or  agricultural,  has  played  a  leading 
part  in  the  decline  of  mussel  populations  in  most  Indiana  streams.  Also, 
the  invasion  of  the  Asiatic  clam  (Corbicula  fluminea  Muller)  has  become 
widespread  in  Indiana  streams.  Since  the  Asiatic  clam  does  not  form 
glochidia,  and  hence  needs  no  fish  host  in  its  life  history,  it  has  a  decided 
competitive  advantage  over  other  mussels.  By  late  1989,  many  sections 
of  the  Wabash  and  White  rivers  were  heavily  infested  with  Asiatic  clams. 

Materials  and  Methods 

All  together,  99  collections  were  made  either  by  crowfoot  bar  or 
by  diving  and  handpicking  at  63  sites  in  the  Wabash  River,  the  White 
River,  and  the  East  Fork  of  the  White  River. 


Ecology 


207 


Figure  1.  Sections  of  the  Wabash  and  White  rivers  included  in  the  study  area.  Collections 
were  made  at  approximate  10-mile  intervals  from  Delphi,  Indiana,  to  the  mouth  of  the 
Wabash  River  and  from  Tunnelton,  Indiana,  on  the  East  Fork  of  the  White  River  to  the 
mouth  of  the  White  River. 


208  Indiana  Academy  of  Science 

Sampling  Stations 

The  area  under  investigation  extended  from  Delphi,  at  Wabash  River 
Mile  331,  to  the  Ohio  River,  and  in  the  East  Fork  of  the  White  River 
from  Tunnelton,  at  White  River  Mile  158,  to  its  mouth  at  Mt.  Carmel, 
Illinois,  a  total  of  about  500  miles  of  stream  (Fig.  1).  In  sampling  the 
study  area  for  distribution  and  relative  abundance  of  commercially  im- 
portant mussels,  the  river  was  divided  into  10-mile  stretches,  and  1  mile 
of  each  stretch  was  marked  off  and  sampled  intensively  using  crowfoot 
bars  either  alone  or  in  conjunction  with  diving  and  handpicking.  The 
location  of  each  sampling  station  by  river  mile  was  obtained  from  flood- 
plain  charts  published  by  the  U.  S.  Army  Corps  of  Engineers.  Rather 
than  sampling  every  tenth  mile,  a  1-mile  section  out  of  each  10  miles,  suit- 
able for  operation  of  the  crowfoot  bar,  was  selected.  If  no  such  section 
was  available  in  any  10-mile  stretch,  a  1-mile  section  either  immediately 
above  or  below  that  stretch  was  sampled.  All  together,  63  such  sampling 
sites  were  selected  and  the  crowfoot  bar  operated  over  those  63  miles  of 
stream. 

Many  of  the  sites  were  visited  more  than  once  to  determine  whether 
the  population  had  changed  either  because  of  changes  brought  about  by 
the  activities  of  the  mussel  fishermen,  or  by  some  other  change  such  as 
increased  pollution. 

The  Crowfoot  Bar 

The  crowfoot  bar  or  brail  bar  (Fig.  2)  has  been  used  for  collecting 
mussels  since  1897  in  the  upper  Mississippi  River  and  its  tributaries  (8, 
10).  Crowfoot  bars  are  fabricated  from  sections  of  iron  pipe  0.75  to  1.0 
inch  in  diameter  and  10-20  feet  in  length.  At  intervals  of  1  to  3  inches 
over  the  length  of  the  bar,  3-foot  lengths  of  seine  cord  or  trot  line  are 
attached  by  loose  half  hitches.  Three  crowfoot  hooks  are  attached  at 
equal  intervals  to  each  strand  of  cord,  and  one  end  of  a  loose  wire  or  rope 
bridle  is  attached  to  each  end  of  the  bar.  When  in  use,  the  bar  is  towed 
downstream  by  a  length  of  rope  attached  by  a  snap  hook  to  an  iron  ring 
mounted  in  the  center  of  the  bridle.  Crowfoot  hooks  are  constructed  from 
two  10-  or  12-inch  lengths  of  No.  9  or  No.  11  steel  wire  bent  into  loops, 
then  twisted  together  in  such  a  manner  that  the  ends  of  the  loops  may 
be  bent  at  equal  intervals  into  4  adjacent  curved  hooks  (Fig.  2).  The 
crowfoot  bar  in  use  today  is  constructed  essentially  as  it  was  70  years  ago. 

Crowfoot  bars  are  carried  in  pairs  in  johnboats,  each  bar  carried  on 
two  iron  stanchions,  mounted  one  on  either  end  of  each  side  of  the  boat. 
When  in  use,  a  0.5-1.0  inch  manila  rope  is  affixed  to  the  bridle  ring  and 
the  whole  bar  comes  to  lie  on  the  river  bottom,  perpendicular  to  the 
shoreline  with  the  strands  of  hooks  spread  out  behind  the  bar  on  the 
bottom.  The  connecting  line  is  allowed  to  assume  an  angle  of  approxi- 
mately 45°  to  the  plane  of  the  river  bottom.  Then  it  is  attached  to  a 
cleat  mounted  on  the  bow  of  the  boat. 

The  boat  usually  is  carried  downstream  by  the  current  with  the  aid 
of  a  sea  anchor,  commonly  referred  to  as  a  mule,  attached  by  cords  to  the 


Ecology 


209 


Figure  2.     A.   Johnboat    with    crowfoot   bar  on   stanchions.    B.    Detailed   view   of   crowfoot 
hooks. 


210  Indiana  Academy  of  Science 

stern  of  the  boat.  Sea  anchors  are  made  of  canvas,  wood,  cr  metal  about 
3  feet  deep  and  5  feet  long.  The  boat  may  be  steered  by  adjusting  the 
cords  that  attach  the  mule  to  the  boat.  Whenever  there  is  insufficient  cur- 
rent for  the  mule  to  propel  the  boat,  an  outboard  motor  is  used. 

In  the  stream  bottom,  mussels  orient  themselves  with  the  open 
posterior  portion  of  the  valves  facing  upstream.  As  the  crowfoot  bar 
hooks  trail  down  the  river  bottom,  they  may  become  lodged  between  the 
open  valves  of  the  mussel.  The  presence  of  a  hook  between  its  valves  irri- 
tates the  mussel  and  it  immediately  closes  its  shell,  thereby  effectively 
attaching  itself  to  the  hook. 

Mussel  fishermen  using  crowfoot  bars  choose  an  area  of  clean  river 
bottom  known  to  support  large  populations  of  mussels.  A  bar  is  lowered, 
pulled  along  for  an  appropriate  time,  and  then  returned  to  the  boat.  As  it 
is  returned  to  the  boat,  its  counterpart  is  lowered  to  the  river  bottom 
so  that  fishing  is  continuous.  The  speed  of  such  dragging  is  slow,  usually 
not  more  than  0.5  mph.  While  one  bar  is  fishing,  the  other  bar  is  cleaned 
of  its  clams  and  any  debris  and  made  ready  for  fishing  again.  As  fishing 
proceeds,  the  two  bars  continue  to  be  alternated  over  the  entire  stretch 
of  stream. 

In  the  present  study,  we  used  a  14-foot  johnboat  carrying  2  crowfoot 
bars,  each  10  feet  long,  and  each  fitted  with  150  crowfoot  hooks,  affixed 
in  groups  of  3  to  each  of  50  three-foot  lengths  of  seine  cord  (Fig.  2).  To 
move  about  at  will,  the  boat  was  powered  with  an  18-horsepower  out- 
board motor. 

Crowfoot  bars  are  most  effective  in  water  more  than  6  feet  deep  and 
on  bottoms  of  clay  or  sand  with  little  large  gravel.  Crowfoot  bars  were  of 
little  use  in  water  less  than  3  feet  deep  or  in  areas  having  bottoms 
covered  with  dead  twigs,  mats  of  dead  leaves,  or  other  such  debris.  In 
such  situations,  the  hcoks  became  entangled  shortly  after  they  touched 
the  river  bottom. 

Each  collection  made  during  this  study  routinely  used  a  pair  of  crow- 
foot bars,  and  on  some  occasions  auxiliary  samples  were  taken  with 
SCUBA  (self-contained  underwater  breathing  apparatus)  gear.  At  each 
station,  one  crowfoot  bar  was  dragged  downstream  for  about  20  min  and 
then  replaced  by  the  second  bar  and  the  process  repeated  over  the  1-mile 
section. 

Diving  and  Handpicking 

Diving  was  performed  with  SCUBA  gear  similar  to  that  used  by 
sportsmen  and  by  commercial  mussel  fishermen.  A  tank  of  compressed 
air  carried  on  the  back  enables  the  diver  to  stay  under  water  for  30-45 
min.  In  some  instances,  a  portable  compressor  is  fixed  to  a  small  raft,  and 
two  or  more  divers  are  supplied  with  air  continuously. 

Mussels  are  placed  in  a  bucket  and  taken  to  the  surface  for  unloading. 
SCUBA  gear  is  a  very  effective  technique  for  sampling  mussel  populations 
since  it  can  be  used  any  time  except  during  flood  conditions,  and  hence 


Ecology  211 

virtually  no  area  of  the  Wabash  or  White  river  systems  of  Indiana  is 
impregnable  to  such  diving  methods.  On  several  occasions,  SCUBA  gear 
was  used  to  collect  in  an  area  that  had  been  collected  previously  with 
a  crowfoot  bar  to  determine  the  relative  efficiencies  of  the  two  methods. 

In  addition  to  diving,  many  samples  were  collected  by  handpicking 
mussels  in  shallow  water.  In  some  areas,  this  was  the  most  productive 
method  of  collecting. 

Reproduction 

The  cultured  pearl  industry  of  today  is  just  as  dependent  upon  the 
continuous  production  of  freshwater  mussels  as  the  pearl  button  industry 
was  a  half  century  ago.  Effective  practices  for  the  conservation  of  any 
resource  necessitates  a  knowledge  of  the  complete  biology  of  the  organ- 
ism concerned. 

For  this  part  of  the  study,  the  gonads  of  369  individual  mussels, 
representing  15  species,  were  examined  (Table  1).  The  reproductive 
processes  of  Quadrula  quadrula  (Rafinesque)  were  studied  intensively, 
and  the  extents  of  the  breeding  seasons  were  determined  tentatively  for 
that  species  and  for  Quadrula  metanevra  (Rafinesque),  Quadirula  pustu- 
loses (Lea),  Obovaria,  olivaria  (Rafinesque),  Tritogonia  verrucosa 
(Barnes),  Megalonaias  gigantea  (Barnes),  Amblema  costata  Rafinesque, 
and  Actinonaias  carinata  (Barnes).  Principal  emphasis  was  placed  on 
those  8  commercially  valuable  species,  and  limited  data  were  available  on 
6  others  as  follows:  Lampsilis  anodontoides  (Lea),  2  males,  2  females; 
Lampsilis  ovata  ventricosa  (Barnes),  2  males,  1  female;  Obliquaria 
reflexa  Rafinesque,  1  male,  2  females;  Elliptio  crassidens   (Lamarck),  2 

Table  1.     Numbers,   kinds,  and  sexes  of  freshwater   mussels  from   the 

Wabash  and  White  rivers,  Indiana,  used  in  studies  of  reproduction.  Only 

those  species  for  which  at  least  15  specimens  were  available  for  study 

are  listed.  Data  for  the  other  species  are  given  in  the  text. 

Number  of  Specimens 
Species 

Actinonaias  carinata 
Amblema  costata 
Megalonaias   gigantea 
Obovaria    olivaria 
Quadrula  metanevra 
Quadrula  pustulosa 
Quadrula  quadrula 
Tritogonia  verrucosa 
Others 

Totals  185  184  369 


Male 

Female 

Total 

9 

14 

23 

11 

9 

20 

11 

6 

17 

18 

21 

39 

7 

8 

15 

24 

14 

38 

78 

89 

167 

13 

7 

20 

14 

16 

30 

212  Indiana  Academy  of  Science 

males,  2  females;  Fusconaia  ebenus  (Lea),  1  male,  3  females;  Fusconaia 
undata  (Barnes),  5  males,  5  females;  Plethobasus  cyphyus 
(Rafinesque),  1  male,  1  female. 

It  is  not  practicable  to  attempt  identification  of  the  sex  of  fresh- 
water mussels  without  aid  of  a  compound  microscope.  In  this  study,  sex 
was  determined  by  inserting  a  hypodermic  needle  through  the  incurrent 
siphon  and  into  the  gonad,  withdrawing  a  small  amount  of  gonadal  tissue, 
and  examining  it  under  a  compound  microscope.  The  differences  between 
the  male  and  female  tissues  were  obvious,  and  it  is  believed  that  unionid 
mussels  can  be  sexed  in  this  manner  with  complete  confidence.  So  far  as 
could  be  determined,  this  method  for  sex  determination  had  no  deleterious 
effects  on  any  of  the  species  studied. 

Sexually  mature  males  and  females  of  Q.  quadrula,  collected  in 
March,  were  studied  histologically  to  determine  whether  there  were  con- 
current areas  of  active  and  inactive  gametogenesis  within  a  single  individ- 
ual. The  complete  foot,  along  with  the  inner  and  outer  gills  were  excised, 
cut  midsagittally,  and  the  central  and  peripheral  areas  compared  by 
studying  stained  histological  sections  from  the  various  midsagittal  areas. 
The  gills  were  fixed  in  Bouin's  fluid  (15)  and  preserved  in  70%  ethanol. 
Serial  sections,  6  microns  thick,  were  prepared  and  stained  with  Heiden- 
hain's  iron-haematoxylin  and  orange  G. 

Breeding  seasons  were  determined  by  correlating  the  degree  of 
gametogenic  development  in  the  gonads  of  both  sexes  with  the  presence 
or  absence  of  eggs  in  the  water  tubes  or  marsupia  of  the  females.  A 
female  was  considered  gravid  if  the  gills  had  become  modified  into  a 
marsupium  and  contained  eggs.  In  determining  the  extents  of  the  breed- 
ing seasons,  the  condition  of  the  development  of  the  ova  was  considered 
the  best  indicator,  and  the  next  best  indicator  was  the  condition  of  the 
male  gonad.  In  some  species,  such  as  O.  olivaria  and  L.  ovata  ventricosa, 
the  marsupium  is  very  pronounced  whereas  in  Q.  quadrula  the  appear- 
ance of  the  marsupium  is  not  typical  even  though  it  may  contain  eggs. 
In  that  species,  the  eggs  are  in  the  inner  and  outer  gills,  and  the  water 
tubes  had  to  be  examined  at  various  times  of  the  year  to  determine  the 
extent  of  the  breeding  season. 

Relative  Abundance  and  Distribution  of  Unionid  Mussels 

Although  no  attempt  was  made  toward  a  revision  of  unionid  mussels, 
it  is  important  here  to  provide  a  brief  account  of  the  various  attempts 
to  delineate  the  systematics  of  those  animals.  The  first  attempt  was  that 
of  Stein  (26)  and  was  followed  by  several  preliminary  lists  by  Call  (4,  5, 
6),  who  presented  generalized  geographic  distributions  for  the  aquatic 
forms  known  to  occur  within  the  boundaries  of  Indiana.  In  addition  to 
these  preliminary  studies  of  Call,  a  rather  superficial  list  of  endemic 
species  was  published  by  Simpson  (25). 

The  first  detailed  study  of  the  unionid  fauna  of  Indiana  was  Call's 
Descriptive  Illustrated  Catalogue  of  the  Molltisca  of  Indiana  (7),  which 
included    detailed    descriptions    and    excellent    drawings    of    each    species 


Ecology 


213 


known  to  exist  in  the  state  at  that  time.  Blatchley  and  Daniels  (3)  pro- 
vided a  supplement  to  Call's  catalogue  adding  a  number  of  species,  mostly 
gastropods.  Daniels  (11,  12)  provided  up-to-date  checklists  of  the  mol- 
luscan  fauna  of  Indiana. 

The  most  recent  comprehensive  study  of  the  unionid  fauna  of  Indiana 
is  Goodrich  and  van  der  Schalie's  (14)  Revision  of  the  Mollusca  of  Indi- 
ana. The  latter  paper  was  used  in  this  study  for  species  identification. 


Table  2.  Distribution  and  abundance  of  unionid  mussels  m  the 
Wabash,  White,  and  East  Fork  of  the  White  rivers  of  Indiana  based  on 
99  collections  in  1966  and  1967.  R,  rare;  — ,  not  present;  C,  common; 
A,  abundant.  Upper  Wabash  River:  Delphi  to  Terre  Haute,  Indiana; 
Middle  Wabash  River:  Terre  Haute  to  Mount  Carmel,  Illinois;  Lower 
Wabash  River:  Mount  Carmel  to  Ohio  River  (22). 


Wabash    River 

White 

River 

Main 
Stream 

East 

Species 

Upper 

Middle 

Lower 

Fork 

Subfamily  Anodontinae 

Alasmidonta  marginata 

R 

— 

— 

— 

— 

Anodonta  grandia 

— 

— 

R 

— 

— 

Anodontoides  j '  eruaaacianua 

R 

— 

— 

— 

— 

Laamigona  complanata 

C 

C 

C 

C 

C 

Laamigona  compressa 

R 

— 

— 

— 

— 

Laamigona  coatata 

K 

K 

— 

— 

— 

Strophitua  rugoaua 

C 

— 

— 

— 

— 

Subfamily  Lampsilinae 

Actinonaiaa  carinata* 

A 

A 

c 

c 

c 

Cyprogenia  irrorata 

— 

R 

— 

— 

— 

Lampailia  anodotitoidea 

C 

C 

V 

— 

— 

Lampailia  ovata  ventricoaa 

c 

C 

c 

c 

(' 

Leptodea  fragilia 

c 

c 

C 

V 

(' 

Leptodea  laeviaaima 

— 

— 

R 

— 

— 

Obliquaria  reftexa 

K 

R 

R 

R 

c 

Obovaria  olivaria* 

A 

A 

A 

C 

c 

Obovaria  aub  rotunda 

K 

— 

— 

— 

H 

Proptera  alata 

c 

C 

C 

C 

c 

Truncilla  truncata 

R 

U 

R 

R 

R 

Subfamily  Unioninae 

Amblema  coatata* 

C 

c 

C 

C 

A 

Cyclonaiaa  tuberculata 

— 

— 

— 

— 

R 

Elliptio  craaaidena 

— 

— 

— 

— 

(.' 

Fuaconaia  ebenua* 

R 

R 

R 

c 

C 

Fuaconaia  undata* 

R 

R 

— 

R 

C 

Megalonaiaa  gigantea* 

R 

C 

— 

R 

(' 

Plethobaaua  cyphyua 

R 

— 

— 

— 

K 

Pleurobema  cordatum 

— 

— 

— 

— 

R 

Quadrtda  metanevra* 

R 

R 

R 

K 

R 

Quadrula  puatuloaa* 

A 

A 

A 

A 

A 

Quadrula  quadrula* 

A 

A 

A 

A 

A 

Tritogonia  verrucoaa* 

C 

C 

C 

— 

— 

The  10  species  of  greatest  commercial  value. 


214  Indiana  Academy  of  Science 

The  distribution,  relative  abundance,  and  locations  of  collections  are 
shown  in  Table  2  (22).  The  species  are  arranged  in  alphabetic  order  under 
three  subfamilies.  Many  species  taken  were  of  little  or  no  commercial 
value,  but  their  occurrence  is  listed.  Particular  emphasis  is  placed  on  10 
species  of  known  commercial  value  (Table  2). 

All  species  collected  in  this  study  were  listed  by  Goodrich  and  van  der 
Schalie  (14)  and  most  were  described  and  illustrated  by  Call  (7).  How- 
ever, some  rare  and  less  abundant  species  listed  by  those  authors,  such  as 
Amblema  peruviana  (Lamarck),  were  not  encountered  here,  and  some, 
once  common  in  Indiana  rivers,  such  as  Elliptio  dilatatus  (Rafinesque) 
and  Quadrula  cylindrica  (Say),  were  in  evidence  only  through  isolated 
dead  shells.  Those  two  species  apparently  have  been  extirpated  from  the 
study  area  within  the  last  two  decades.  Other  species,  such  as  Pleurobema 
cordatum  (Rafinesque),  Cyclonaias  tuberculata  (Rafinesque),  Cyprogenia 
irrorata  (Lea),  and  Obovaria  sub7'otunda  (Rafinesque)  are  much  less 
abundant  and  have  much  more  restricted  patterns  of  distribution  than 
those  recorded  by  Goodrich  and  van  der  Schalie  (14).  No  species 
encountered  in  this  study  exhibited  wider  ranges  of  distribution  since 
the  earlier  studies,  and  most  species  appeared  much  less  abundant.  Only 
two  species,  Q.  pustulosa  and  Q.  quadrula,  were  as  abundant  or  more 
abundant  than  indicated  by  earlier  authors. 

Such  information  indicates  a  definite  trend  toward  restriction  in 
range  and  reduction  in  absolute  abundance  of  most  unionid  mussels  in  the 
Wabash  and  White  rivers.  Although  some  species  may  have  been  extir- 
pated from  the  areas  under  investigation  there  may  still  be  limited  pop- 
ulations in  other  parts  of  the  drainage  system.  If  there  are,  hopefully 
the  populations  in  the  mainstreams  of  the  Wabash  and  White  rivers  may 
be  re-established  when  conditions  permit. 

This  pattern  of  reduction  and  elimination  of  unionid  mussel  popula- 
tions may  be  attributed  to  two  principal  factors:  1)  the  disturbance  and 
destruction  of  mussel  beds  through  overexploitation  for  commercial  pur- 
poses; and  2)  deterioration  of  the  environment  as  suitable  habitat  for 
mussels  through  increasing  burdens  of  pollution. 

The  devastating  effects  of  industrial  and  organic  pollutants  on  unionid 
mussels  were  first  pointed  out  by  Ortmann  (23)  in  his  report  on  the  effects 
of  such  pollution  in  the  Allegheny,  Monongahela,  and  Ohio  rivers  in 
western  Pennsylvania.  Ten  years  later,  Forbes  and  Richardson  (13)  noted 
a  direct  correlation  between  increasing  levels  of  pollution  and  decreasing 
ranges  and  numbers  of  mollusks  in  the  Illinois  River.  More  appropriate 
to  the  present  work  was  the  study  by  Baker  (1)  in  which  he  reported  in 
detail  the  elimination  of  all  mussels  from  certain  areas  of  the  Big 
Vermillion  River,  a  tributary  to  the  Wabash  River,  by  heavy  loads  of 
municipal  organic  wastes.  More  recently,  Wurtz  (30)  stated  that  unionid 
mussels  were  quite  intolerant  to  pollution  of  any  kind  and  reported 
unequivocally  that  freshwater  mussels  disappear  quickly  from  streams 
carrying  moderately  heavy  burdens  of  pollutants. 

Polluted  streams  exhibit  remarkable  patterns  of  recovery  down- 
stream from  sources  of  pollution  or  following  the  abatement  of  pollution 


Ecology  215 

(1,  16,  18,  19).  Perhaps  the  first  detailed  description  of  the  biological 
recovery  of  a  river  system  downstream  from  a  source  of  pollution  was 
that  of  Kolkwitz  and  Marsson  (17)  who  reported  on  the  changes  in  the 
species  composition  of  aquatic  communities,  and  pointed  out  that  the 
abundance  and  diversity  of  organisms  increased  as  the  effects  of  pollution 
diminished. 

Disruption  of  the  stream  bottom  by  mechanical  means  may  be  just 
as  devastating  to  a  mussel  population  as  pollution.  One  of  the  methods 
for  collecting  mussels  that  is  most  destructive  of  the  streambed  is 
mechanical  or  hydraulic  dredging.  In  the  operation  of  such  a  dredge,  about 
the  uppermost  2  feet  of  stream  bottom  is  lifted  out  of  place,  carried  to 
a  barge  where  the  mussels  are  removed,  and  then  dumped  back  into 
the  river.  In  this  operation,  the  stream  bottom,  with  all  its  biotic  com- 
munities, is  completely  discommoded.  Any  benthic  organisms  that  survive 
must  become  re-established  rather  quickly  or  those  segments  of  the 
stream  ecosystem  that  depend  on  benthos  for  their  livelihood  will  not 
survive. 

Tongs,  crowfoot  bars,  and  other  such  equipment  disturb  the  stream- 
bed  to  a  limited  extent,  but  they  usually  are  used  only  in  localities  where 
they  are  known  to  be  effective.  At  most,  those  tools  disrupt  small  areas 
of  bottom  to  a  depth  of  a  few  inches,  instead  of  upheaval  of  the  entire 
streambed. 

Perhaps  the  method  least  destructive  to  the  stream  bottom  is  hand- 
picking.  Here,  the  musseler  simply  walks  along  the  stream  bottom  or 
swims  over  it  and  picks  up  whichever  mussels  he  desires. 

Numbers  and  Kinds  of  Mussels  Collected 

Data  for  the  20  species  of  mussels  taken  in  at  least  6  different  col- 
lections are  arranged  according  to  method  of  collection  in  Table  3. 

From  the  data  in  Table  3,  it  is  obvious  that  about  %  (34.0%)  of  the 
total  catch  was  made  up  of  mapleleaf  (Quadrula  quadrula) ,  one  of  the 
most  highly  sought-after  commercial  mussels  for  the  cultured  pearl 
industry.  The  mapleleaf  was  followed  in  order  by  the  sandshell  (Obovaria 
olivaria)  which  contributed  10.2%  and  the  pimpleback  (Q.  pustulosa) 
with  9.0%.  Thus,  these  three  species  made  up  53.2%  of  the  total  catch. 
The  remaining  7  of  the  10  commercially  important  species  listed  earlier 
comprised  another  23.9%  of  the  catch,  and  all  together  the  10  species 
made  up  more  than  %   (77.1%)  of  all  mussels  taken  during  the  study. 

Of  the  20  species  listed  in  Table  3,  several  have  shells  that  are  too 
thin  and  fragile  or  are  too  highly  colored  to  be  of  value  as  nuclei  for 
cultured  pearls.  Others  were  taken  in  such  small  numbers  that  they  could 
not  be  considered  important  in  the  commercial  market.  Only  15  of  the  20 
species  were  represented  by  more  than  25  specimens. 

The  white  heelsplitter,  Lasmigona  complanata  (Barnes),  is  used 
sparingly  in  the  cultured  pearl  industry,  but  has  some  commercial  value 


216  Indiana  Academy  of  Science 

Table  3.  Numbers  of  individuals  of  the  20  species  of  the  unionid  mus- 
sels represented  in  six  or  more  collections  taken  by  crowfoot  bar  and  by 
handpickifig,  together  with  the  numbers  of  collections  in  which  they 
occurred,  from  the  Wabash  and  White  rivers  during  1966  and  1967.  The 
averages  are  for  the  numbers  of  individuals  per  collection. 


All 

Collections 

Crowfoot  Bar 

Handpicking 

Total 

Species 

No. 

Coll. 

No. 

Coll. 

No. 

Coll. 

Lasmigona  complanata 

24 

5 

40 

11 

04 

10 

Lasmigona   costata 

1 

1 

5 

5 

0 

0 

Others* 

12 

11 

3 

2 

15 

13 

Actinonaias  carinata 

XX 

21 

21 

6 

109 

27 

Lampsilis   anodontoides 

36 

0 

1 

1 

37 

7 

Lampsilis  ovata  ventricosa 

34 

20 

54 

15 

XX 

35 

Leptodea  fragilis 

54 

22 

3  5 

15 

X9 

37 

Obliquaria  reflexa 

10 

0 

3 

2 

13 

8 

Obovaria  olivaria 

162 

35 

32 

10 

194 

45 

Obovaria  subrotunda 

9 

4 

2 

2 

11 

6 

Propter  a  alata 

9 

7 

14 

6 

23 

13 

Truncilla  truncata 

4 

3 

0 

6 

10 

9 

Others* 

— 

— 

2 

2 

2 

2 

Amblema  costata 

41 

If) 

08 

10 

104 

25 

Elliptio  crassidens 

35 

3 

22 

3 

57 

6 

Fusconaia  ebenus 

4 

4 

51 

8 

55 

12 

Fusconaia  undata 

S 

6 

13 

7 

21 

13 

Megalonaias  gigantea 

30 

X 

08 

it 

93 

17 

Quadrula  metanevra 

35 

12 

12 

0 

47 

18 

Quadrula  pustulosa 

XX 

25 

84 

14 

172 

39 

Quadrula  quadrula 

359 

39 

288 

17 

047 

56 

Tritogonia  verrucosa 

27 

12 

— 

— 

27 

12 

Others* 

6 

2 

10 

12 

22 

14 

Totals 

1076 

02 

830 

37 

1906 

99 

Average 

17.4 

22.4 

19.3 

*  The  total  number  of  specimens  for  each  of  the  species  included  in  the  category  of 
"Others"  are:  Alasmidonta  marginata  (Say),  2  specimens;  Anodonta  grandis  Say,  2; 
Anodontoides  ferussacianus  (Lea),  2;  Lasmigona  compressa  (Lea),  4;  Strophitus  rugosus 
(Swainson),  5;  Cyprogenia  irrorata,  1;  Leptodea  laevissima  (Lea),  1;  Cyclonaias 
tubcrculata,  3 ;  Plethobasus  cyphyus,  8 ;  Pleurobema  cordatum,  4 ;  and  7  unidentified 
individuals. 


because  of  its  relative  abundance.  The  shell  is  fairly  heavy  and  the 
nacre  is  creamy  white.  The  64  specimens  taken  made  up  3.4%  of  the  total 
catch;  however,  most  of  those  shells  were  collected  by  hand  in  the  East 
Fork  of  the  White  River  in  water  less  than  6  feet  deep. 

Lasmigona  costata  (Rafinesque),  the  fluted  shell,  was  rare  in  the 
collections  and  is  of  very  little  commercial  value. 

The  mucket,  Actinonaias  carinata,  was  fourth  in  numerical  abundance 
in  our  collections  and  made  up  5.7%  of  the  total  catch.  The  shell  of  this 
species  is  commercially  important  and  is  of  high  quality  in  the  manufac- 
ture of  nuclei  for  cultured  pearls. 


Ecology  217 

The  yellow  sandshell,  Lampsilis  anodontoides,  is  not  of  commercial 
value  because  usually  it  does  not  have  enough  thick  portion  of  the  shell 
to  warrant  the  expense  of  cutting  it  for  nuclei. 

Although  the  pocketbook,  Lampsilis  ovata  ventricosa,  and  the 
floater,  Leptodea  fragilis  (Rafinesque),  comprised  4.5  and  4.7%,  respec- 
tively, of  the  total  catch,  neither  is  important  because  of  the  quality  of 
the  shell.  The  pocketbook  usually  is  pale  green  to  brown  in  color  and  the 
shell  of  the  floater  is  so  fragile  that  it  can  easily  be  crushed  between  the 
fingers  of  one  hand. 

With  the  exception  of  Obovaria  olivaria,  all  other  members  of  the 
Lampsilinae  were  taken  in  such  small  numbers  that  they  are  not  of 
commercial  importance.  The  sandshell  is  one  of  the  most  highly  sought- 
after  shells  and  is  relatively  abundant,  particularly  in  the  Wabash  River, 
where  it  consistently  makes  up  a  significant  part  of  the  commercial 
market. 

The  threeridge,  Amblema  costata,  ranked  fifth  in  the  number  of 
specimens  collected  in  this  study  and  made  up  5.5%  of  the  total  catch. 
This  mussel  was  common  throughout  the  study  area  and  was  particularly 
abundant  in  the  upper  reaches  of  the  East  Fork  of  the  White  River.  In 
that  area,  in  contributes  consistently  to  the  commercial  market. 

Elliptio  crassidens,  the  elephant's  ear,  is  of  no  commercial  value  in 
the  cultured  pearl  industry  because  of  the  purple  color  of  its  nacre. 

Although  the  two  representatives  of  the  genus  Fusconaia,  F.  ebenus 
and  the  pigtoe,  F.  undata,  contributed  only  2.9  and  1.1%,  respectively,  to 
the  total  catch  in  this  study,  they  are  highly  desirable  as  shells  in  the 
cultured  pearl  industry.  Fusconaia  ebenus  is  particularly  desirable  and 
actually  makes  up  a  significant  portion  of  the  commercial  market. 

The  largest  freshwater  mussel  taken  in  the  study  was  the  wash- 
board, Megalonaias  gigantea.  In  this  study  it  contributed  4.9%  of  the 
total  catch  and  was  particularly  important  in  the  East  Fork  of  the  White 
River.  The  washboard  has  a  relatively  high  commercial  value  because  of 
its  size  and  thickness  of  the  shell,  but  the  quality  of  the  nacre  is  not  as 
desirable  as  that  of  some  other  shells. 

The  most  important  genus  of  freshwater  unionids,  so  far  as  the 
contribution  of  its  representatives  to  the  cultured  pearl  industry  is  con- 
cerned, is  Quadrula.  In  the  Wabash  River  system,  three  members  of  the 
genus,  the  monkeyface,  Q.  metanevra,  the  pimpleback,  Q.  pustulosa,  and 
the  mapleleaf,  Q.  quadrula,  contributed  45.4%  to  the  total  number  of 
mussels  taken  during  this  study.  These  3  shells,  together  with  Obovaria 
olivaria  and  Fiisconia  ebenus  are  the  5  shells  of  greatest  commercial  value 
for  pearl  nuclei  and  they  made  up  56.7%  of  the  total  catch  in  this  study. 
When  these  5  shells  are  considered  together  with  the  other  5  of  the  10 
most  important  commercial  species  listed  earlier,  the  contribution  to  the 
total  number  collected  in  this  study  was  77.1%.  Thus,  the  10  most  sought 
after  species  are  in  aggregate  the  10  most  abundant  species. 


218  Indiana  Academy  of  Science 

The  only  other  species  of  commercial  importance  is  the  pistolgrip, 
Tritogonia  verrucosa,  which  has  a  distribution  limited  almost  entirely  to 
the  Wabash  River  upstream  from  Vincennes,  Indiana.  Although  this  shell 
contributed  only  1.47c  to  the  overall  catch,  in  the  area  above  Vincennes  in 
1966,  it  contributed  3.7%. 

Comparisons  of  Catches  in  Different  Localities 

In  the  present  study,  it  was  assumed  that  the  sites  for  collections  by 
crowfoot  bar  and  by  handpicking  were  distributed  randomly  over  the 
river  systems  investigated  and  that  those  methods  of  collection  were 
non-selective  in  the  kinds  of  mussels  taken.  Thus,  the  number  and  kinds 
of  mussels  in  the  collections  should  be  a  good  indication  of  their  distribu- 
tion throughout  the  study  area  (Table  3).  As  is  true  of  most  animals  in 
the  world,  the  distribution  of  unionid  mussels  in  the  Wabash  and  White 
river  systems  was  not  uniform.  Rather,  some  of  the  species  were  not 
taken  in  the  Wabash  River  and  others  were  not  taken  in  the  mainstream 
or  the  East  Fork  of  the  White  River   (Tables  2  and  3). 

Among  the  20  species  mentioned  in  Table  3,  only  one,  Elliptio  crassi- 
dens,  was  not  taken  in  the  Wabash  River  at  any  time,  and  another, 
Obliquuria  reflexa,  was  represented  by  a  single  female  specimen  taken 
on  a  crowfoot  bar  near  Cayuga,  Indiana,  24  August  1967.  Conversely, 
E.  crassidens  was  represented  by  57  specimens  collected  by  crowfoot  bar 
and  handpicking  from  the  East  Fork  of  the  White  River,  and  12  speci- 
mens of  O.  reflexa  were  collected  from  the  mainstream  and  East  Fork 
of  the  White  River  in  1967.  In  that  part  of  the  White  River  system 
under  study,  5  of  the  20  species  were  not  present  in  the  collections  from 
the  main  stem  and  3  were  not  taken  in  the  East  Fork  of  the  White 
River.  However,  of  all  of  those  species,  only  Tritogonia  verrucosa  is  of 
commercial  value. 

It  is  also  obvious  from  the  data  in  Table  3  that  some  species  are 
relatively  much  more  abundant  in  one  area  than  in  another,  whereas 
others  may  be  fairly  evenly  distributed.  Several  species,  Actinonaias 
carinata,  Obovaria  olivaria,  Quadrula  metanevra,  Q.  quadrula,  and  Tri- 
togonia verrucosa  apparently  are  much  more  common  in  the  Wabash 
River  than  in  the  White  River.  By  the  same  token,  Amblcma  costata, 
Fusconaia  ebenus,  and  Megalonaias  gigantea  appear  to  be  more  common 
in  the  White  River.  Only  one  of  the  10  most  commercially  valuable  spe- 
cies, Quadrula  pustulosa,  seems  fairly  evenly  distributed  over  the  study 
area. 

Another  datum  that  may  give  an  indication  of  the  distribution  of 
mussels  in  the  rivers  investigated  is  the  number  of  collections  in  which 
each  kind  of  mussel  occurred.  Only  four  species,  Lampsilis  ovata  ventri- 
cosa,  Obovaria  olivaria,  Quadrula  metanevra,  and  Q.  quadrula,  were 
taken  by  crowfoot  bar  and  by  handpicking  from  the  Wabash  River  in 
both  1966  and  1967,  and  from  the  main  stem  and  East  Fork  of  the 
White  River  (Table  3).  One  other  species,  Propter®  alata  (Say),  was 
taken  by   handpicking   from   each   area   and   by   crowfoot  bar   from   the 


Ecology  219 

main  stem  and  East  Fork  of  the  White  River  and  from  the  Wabash 
River  in  1966,  but  not  in  1967.  Similarily,  Truncilla  truncata  Rafinesque, 
although  rare,  was  collected  in  each  of  the  areas. 

For  the  entire  study,  19  of  the  20  species  were  represented  in  the 
61  collections  from  the  Wabash  River,  17  were  represented  in  the  27 
collections  from  the  East  Fork  of  the  White  River,  and  15  were  taken 
in  the  11  collections  from  the  White  River  proper.  Here,  it  should  be 
pointed  out  that  the  61  collections  from  the  Wabash  River  were  taken 
over  a  distance  of  about  330  miles,  the  27  from  the  East  Fork  from 
about  110  miles  of  stream,  and  the  11  from  the  main  stem  of  the  White 
River  from  about  50  miles  of  stream,  so  that  the  intensity  of  sampling 
was  about  the  same  in  each  area. 

In  considering  the  occurrence  of  only  the  10  species  of  greatest 
commercial  value  in  all  collections  made  during  this  study,  only  1, 
Q.  quadrula,  was  represented  in  more  than  half  those  collections.  Four 
others,  O.  olivaria,  Q.  pustulosa,  A.  carinata,  and  A.  costata,  were  repre- 
sented in  more  than  %,  and  each  of  the  remaining  5  was  represented 
in  from  10  to  20%  of  all  collections.  In  the  Wabash  River  alone,  2 
species,  Q.  quadrula  and  O.  olivaria  were  represented  in  more  than  half 
the  61  collections,  5  were  represented  in  from  20  to  35%,  and  3  were 
present  in  less  than  10%  of  the  collections.  In  the  main  stem  of  the 
White  River,  where  only  11  collections  were  made,  no  species  was  taken 
in  more  than  four  collections,  and  one,  T.  verrucosa,  was  not  repre- 
sented. In  the  East  Fork  of  the  White  River,  2  species  were  present  in 
more  than  half  the  27  collections,  2  others  were  represented  in  more 
than  %,  3  others  in  more  than  hi,  2  others  in  from  15  to  20%,  and  1, 
T.  verrucosa,  was  not  represented.  From  these  data  it  is  apparent  that 
Q.  quadrula  and  O.  olivaria  are  ubiquitous  and  that  the  other  eight 
species,  with  the  exception  of  T.  verrucosa,  are  fairly  widespread 
throughout  the  Wabash  River  system. 

Comparisons  of  Catches  in  1966  and  1967 

During  the  course  of  this  study,  10  one-mile  sections  of  the  Wabash 
River  between  Delphi  (Mile  331)  and  Terre  Haute  (Mile  220),  Indiana 
were  sampled  with  a  crowfoot  bar  during  June  or  July  1966,  and  again 
in  August  1967.  All  collections  were  made  with  the  same  equipment 
operated  by  the  same  personnel,  and  the  data  are  believed  comparable. 

In  1966,  297  specimens  referable  to  21  species  were  taken  from  the 
10  collecting  sites  (Table  4),  whereas  in  1967,  only  56  mussels  referable 
to  11  species  were  collected.  This  amounts  to  a  reduction  in  numbers 
of  specimens  of  81%  and  a  reduction  in  the  diversity  of  species  of 
48%.  The  only  instance  in  which  more  individuals  were  collected  in 
a  single  station  in  1967  than  in  1966,  was  at  Wabash  River  Mile  253-252, 
where  24  individuals  were  taken  the  second  year  as  compared  with  19 
the  first.  Mussels  referable  to  9  species  were  taken  during  the  2-year 
period,  of  which  7  were  collected  in  1966  and  6  in  1967.  The  numbers 
of  individuals  of  each  species  taken  in  1966  were:  Lasmigona  complanata, 
8;   Lampsilis  anodontoides,  2;   Proptera  alata,  2;    Obovaria  olivaria,   2; 


220  Indiana  Academy  of  Science 

Table  4.     Collections  of  mussels  from  each  of  10  1-mile  sectio?is  taken 
with  a  crowfoot  bar  in  1966  and  1967,  showing   the  numbers  of  mus- 
sels a?id  numbers  of  species  taken  in  each  collection. 

1966  1967 


50 

12 

22 

9 

42 

7 

2 

1 

45 

8 

1 

1 

19 

5 

2 

1 

7 

2 

■ — 

— 

27 

9 

2 

2 

39 

7 

— 

— 

19 

7 

24 

(5 

3 

2 

— 

— 

46 

10 

3 

2 

No.  of         No.  of  No.  of         No.  of 

River  Mile  Specimens     Species         Specimens     Species 

320-319 
311-310 
294-293 
285-284 
281-280 
271-270 
259-258 
253-252 
239-238 
230-229 

Totals  297  21  56  11 


Quadrula  pustulosa,  1;  Q.  quadrula,  3;  and  Tritogonia  verrucosa,  1.  In 
1967,  the  numbers  were:  Actinonaias  carinata,  4;  Obliquaria  reflexa,  1; 

0.  olivaria,    6;    Q.    pustulosa,    1;    Q.    quadrula,    11;    and    T.    verrucosa, 

1.  An  analysis  of  these  catches  indicates  that  the  mussels  taken 
in  1967  had  a  much  higher  commercial  value  than  those  taken  in  1966; 
of  the  10  commercially  important  mussels  listed  earlier,  there  were  23 
taken  in  1967  and  only  7  taken  in  1966.  The  reasons  for  these  differences 
in  the  locality  under  discussion  are  not  readily  apparent.  In  the  light 
of  the  data  from  the  other  nine  localities  where  collections  were  made 
both  years,  it  seems  most  likely  that  the  chance  of  a  crowfoot  hook 
coming  in  contact  with  a  particular  mussel  was  very  important  in  the 
makeup  of  that  part  of  the  study.  Still,  the  consistently  low  catches 
in  1967  in  the  other  9  localities  indicates  that  the  overall  populations 
of  mussels  in  the  upper  Wabash  River  had  declined  rather  spectacularly. 

The  numbers  of  individuals  of  each  species  taken  each  year  in  all 
10  collections  are  listed  in  Table  5.  Of  the  22  species  listed,  21  were 
taken  in  1966  and  11  were  taken  in  1967.  A  single  specimen  represented 
the  total  catch  for  each  of  8  species  in  1966,  and  there  were  5  species 
represented  by  a  single  individual  in  1967.  Although  9  of  the  10  com- 
mercially desirable  species  listed  earlier  were  taken  in  1966  com- 
pared with  6  such  species  in  1967,  the  percentages  of  the  total  catch 
made  up  by  those  species  for  the  2  years  was  about  the  same.  However, 
the  actual  drop  in  the  numbers  of  such  highly  desirable  species  as  A. 
carinata  and  Q.  quadrula  cannot  be  regarded  lightly.  This  is  even  more 
impressive  when  it  is  considered  that  all  of  the  specimens  of  Q.  quadrula 
collected  in  1967  were  taken  in  the  single  collection  at  Mile  253-252. 


Ecology 


221 


Table  5.     Numbers  of  each  species  of  mussels  collected  by  crowfoot  bar 

from  the  same  10  one-mile  sections  of  the   Wabash  River  in  1966  and 

1967  (see  Table  U  for  location  of  collecting  sites). 


Species 


No.  Taken 

No.  Taken 

in  1966 

in  1967 

1 



1 

__ 

8 

2 

1 

1 

5 

— 

43 

9 

2 

— 

9 

4 

16 

1 

— 

1 

44 

15 

3 

— 

5 

— 

1 

— 

1 

1 

1 

— 

1 

— 

1 

— 

15 

_ 

24 

4 

110 

17 

5 

1 

Alasmidonta  marginata 
Anodontoides  terussacianus 
Lasmigona  complanata 
Lasmigona   compressa 
Strophitus  rugosus 
Actinonaias  carinata 
Lampsilis    anodontoides 
Lampsilis    ovata    ventricosa 
Leptodea  f?'agilis 
Obliquaria  reflexa 
Obovaria  olivaria 
Obovaria   subrotunda 
Proptera  alata 
Truncilla  truncata 
Amblema  costata 
Fusconaia  ebenus 
Fusconaia  undata 
Plethobasus  cyphyus 
Quadrula  metanevra 
Quadrula  pustulosa 
Quadrula   quadrula 
Tritogonia  verrucosa 


Totals 


297 


5(5 


Only  two  species,  O.  olivaria  and  Q.  quadrula,  were  represented  in 
each  of  the  10  collections  in  1966,  whereas  O.  olivaria  was  represented 
in  5  of  the  1967  collections  and  Q.  quadrula  was  represented  in  4.  Seven 
other  species  were  represented  in  4  or  more  collections  in  1966,  but  only 
2  species  other  than  those  mentioned  above  were  represented  in  more 
than  a  single  sample.  The  lack  of  representation  in  more  than  a  single 
collection  in  1967  lends  credence  to  the  decline  in  species  diversity  as 
well  as  actual  abundance  of  mussels  during  the  period  under  considera- 
tion. 

In  comparing  the  catches  in  1966  with  those  in  1967,  it  is  important 
to  compare  them  with  changes  in  the  numbers  of  shells  sold  for  use  in 
the  cultured  pearl  industry.  In  1965,  2000  tons  of  Indiana  shells  were  sold 
for  use  in  Japan;  in  1966,  that  number  was  4200  tons;  in  1967  it  was 
1080  tons;  and  in  1968  the  total  number  probably  did  not  exceed  250 
tons.    Using  these   data   for   comparison,   the   percentage    drop   between 


222  Indiana  Academy  of  Science 

1966  and  1967  was  essentially  the  same  as  for  our  samples  taken  in 
the  survey  and  there  was  another  substantial  drop  between  1967  and 
1968  for  which  we  have  no  data. 

It  is  important  here  to  note,  however,  that  in  1969  the  com- 
mercial musselers  have  reported  very  large  numbers  of  small  mussels 
in  the  Wabash  River,  and  from  the  estimated  size  of  these  young  ani- 
mals they  must  be  in  their  second  or  third  year  of  life.  On  examination, 
it  was  found  that  most  of  these  small  mussels  were  Asiatic  clams. 

Efficiency  of  Different  Kinds  of  Gear 

Only  two  methods  for  collecting  mussels  were  used  in  this  study, 
the  crowfoot  bar  and  handpicking  with  or  without  SCUBA  gear.  When 
a  crowfoot  bar  is  dragged  over  an  area,  only  those  mussels  at  the 
surface  of  the  stream  bottom  with  their  valves  open  are  capable  of 
being  caught.  With  diving  or  handpicking,  each  mussel  seen  or  touched 
by  the  musseler  is  capable  of  being  captured.  Thus,  it  is  obvious  that 
the  opportunities  for  collecting  mussels  by  handpicking,  including  div- 
ing with  or  without  auxiliary  air  supply,  are  much  greater  than  with  a 
crowfoot  bar. 

On  13  September  1966,  in  the  Wabash  River  near  Americus,  Indiana, 
we  roped  off  an  area  of  stream  bottom  10  feet  wide  and  170  feet  long. 
That  area  was  then  dragged  with  a  crowfoot  bar,  and  a  total  of  8 
mussels  representing  3  species  and  weighing  8.4  lbs.  was  collected. 
Following  the  study  with  the  crowfoot  bar,  2  divers  equipped  with 
SCUBA  gear  covered  the  same  marked-off  area  and  picked  up  220 
mussels  of  legal  size  including  201  specimens  of  commercial  value  as 
follows:  8  Actinonaias  carinata,  3  Obovaria  olivaria,  13  Amblema  costata, 
7  Quadrula  metayievra ,  6  Q.  pustulosa,  and  164  Q.  quadrula  that  weighed 
a  total  of  138.8  lbs.  The  other  19  specimens  represented  4  species  of 
no  commercial  value  and  weighed  8.7  lbs. 

On  the  basis  of  the  data  in  Table  3,  where  the  average  catch  per 
drag  of  the  crowfoot  bar  over  a  1-mile  section  of  river  yielded  17.4 
mussels  and  the  average  catch  per  collection  by  handpicking  was  22.4 
mussels,  the  difference  does  not  seem  great.  It  should  be  pointed  out 
here  that  each  collection  with  a  crowfoot  bar  required  2  men  more  than 
2  hours  of  work  whereas  the  average  time  for  the  collections  by  hand- 
picking  was  2  men  for  less  than  30  min.  Thus,  it  is  apparent  that  the 
latter  method  is  more  than  four  times  as  efficient  on  the  basis  of  time 
alone. 

Handpicking  with  the  aid  of  an  auxiliary  air  supply  is  so  efficient 
that  an  entire  mussel  bed  can  be  wiped  out  in  a  few  hours.  We  know 
of  1  instance  in  which  3  men  with  auxiliary  air  supply  collected  more 
than  3000  lbs.  of  commercially  valuable  mussels  from  a  single  bed  in 
less  than  2  days.  Of  course,  they  collected  only  mussels  of  legal  size 
and  left  the  smaller  individuals  behind.  That  act  did  assure  the  survival 
of  the  population,  but  the  growth  rate  of  most  thick-shelled  mussels  is 
slow;  it  requires  about  4  or  5  years  for  a  mapleleaf  or  threeridge  mussel 


Ecology  223 

to  attain  the  legal  size  of  2.5  inches.  Still,  by  removing  all  the  larger 
individuals,  the  reproductive  capacity  of  the  population  is  seriously  im- 
paired, and  it  may  take  another  4  or  5  years  or  even  longer  for  the 
population  to  recover  its  reproductive  potential. 

Reproduction 

The  cell  lineage  in  unionid  mussels  has  been  adequately  described 
and  illustrated  by  Lillie  (20),  and  atypical  spermatogenesis  in  mollusks 
has  been  reported  by  Coe  and  Turner  (9)  and  Loosanoff  (21).  No  at- 
tempt has  been  made  in  the  present  study  to  describe  the  details  of 
meiosis  in  either  the  male  or  female  mussel,  but  atypical  spermatogenesis 
was  demonstrated  in  Quadrula  quadrula  (2). 

Atypical  sperm  cells  develop  from  the  same  type  of  spermatogenic 
cells  as  normal  sperm  cells,  but  the  atypical  cells  undergo  several  mitot- 
ic divisions  while  still  surrounded  by  the  original  cytoplasm  within  the 
original  cell  membrane.  The  number  of  cells  undergoing  atypical  sperma- 
togenesis usually  are  much  less  numerous  than  those  undergoing  normal 
spermatogenesis,  but  there  are  occasions  when  the  atypical  cells  out- 
number the  typical  ones. 

Spermatogenesis  is  a  continuous  process  in  Q.  quadrula,  the  most 
active  period  being  in  the  seasons  immediately  before  and  during  spawn- 
ing. In  March  and  April,  before  the  breeding  season  commences, 
spermatogenesis  is  active  and  continuous  until  late  June. 

In  the  ovary  of  Q.  quadrula,  the  primordial  wall  becomes  differen- 
tiated and  the  ovum  is  formed  following  the  usual  maturation  divisions. 
The  ovum  moves  from  the  lumen  of  the  ovary  through  ducts  to  the 
suprabranchial  chamber  and  down  into  the  water  tubes  of  the  inner 
and  outer  gills.  The  spermatozoa  move  through  ducts  from  the  lumina 
of  the  acini  to  the  suprabranchial  chambers  and  cloacal  chamber  and  out 
the  excurrent  siphon  into  the  water.  The  sperm  are  drawn  into  the  in- 
current  siphon  of  the  female  by  respiratory  action,  and  are  carried 
through  the  mantle  cavity  to  the  gills  where  they  enter  the  water  tubes 
through  the  ostia. 

Syngamy  and  cleavage  take  place  in  the  water  tubes  and  the  gills 
become  modified  into  a  marsupium  to  accommodate  the  developing  em- 
bryos. Q.  quadrula  is  larviparous,  the  homolecithal  ova  are  incubated 
within  the  interlamellar  spaces  of  the  modified  gills.  In  about  2  to  4 
weeks,  glochidia  are  formed  and  discharged  from  the  water  tubes  into 
the  suprabranchial  chambers,  through  the  cloacal  chamber,  and  out  the 
excurrent  siphon.  Since  glochidia  are  obligate  temporary  parasites  of 
fishes,  if  they  do  not  find  a  host  fish  within  a  few  days  they  perish. 

If  there  is  a  large  or  dense  adult  population  of  mussels  or  if 
there  is  a  sparse  population  of  suitable  fishes  to  serve  as  hosts  for 
the  developing  glochidia,  the  survival  of  young  individuals  is  low.  If, 
however,  there  is  an  adequate  breeding  population  of  mussels  but  they 
are  scattered  rather  sparsely  over  a  large  area,  the  survival  of  glochidia 


224  Indiana  Academy  of  Science 

will  be  high  if  enough  suitable  host  fishes  are  present.  Thus,  if  the  legal- 
sized  mussels  are  harvested  after  they  have  reproduced,  any  glochidia 
that  survive  the  host-parasite  relationship  with  fishes,  will  have  a  much 
better  rate  of  survival  but  they  will  not  be  able  to  add  to  the  harvest- 
able  crop  for  about  5  years. 

According  to  Pennak  (24),  all  members  of  the  Subfamily  Unioninae 
are  short-term  breeders  and  are  gravid  sometime  between  April  and 
August;  whereas,  all  members  of  the  Subfamilies  Anodontinae  and 
Lampsilinae  are  long-term  breeders  among  which  fertilization  takes 
place  in  mid  or  late  summer  and  the  embryos  are  carried  until  the 
next  spring. 

Among  the  species  reported  here  (Table  1),  only  Actinonaias  car- 
inata  and  Obovaria  olivaria  belong  to  subfamilies  other  than  the  Union- 
inae. Histological  studies  of  the  gonads  of  21  specimens  of  Actinonaias 
carinata  indicated  that  the  ovary  was  not  in  breeding  condition  until 
late  July  and  continued  in  that  condition  until  mid-October.  Spermato- 
genesis became  active  during  the  late  spring,  but  no  mature  sperm  were 
encountered  until  late  July.  Thus,  based  on  the  relatively  few  specimens 
studied,  it  is  apparent  that  fertilization  could  not  have  taken  place 
much  before  August  and  could  continue  on  through  early  October.  No 
observations  were  made  on  the  time  of  release  of  glochidia  in  this  spe- 
cies, but  the  evidence  at  hand  indicates  that  it  is  a  long-term  breeder. 

In  O.  olivaria,  the  gonads  of  39  specimens  were  examined  in  detail. 
Uncleaved  but  mature  ova  were  observed  in  the  ovaries  of  specimens 
sacrificed  in  August,  and  at  that  time  the  ovary  appeared  to  be  in 
typical  breeding  condition.  Mature  spermatozoa  were  present  in  the 
lumina  in  late  July,  indicating  that  active  spermatogenesis  was  in 
progress.  These  conditions  in  the  gonads  of  each  sex  persisted  through 
September,  typical  for  the  long-term  breeder.  No  observations  were 
made  on  the  release  of  glochidia. 

All  other  species  for  which  there  were  adequate  samples  for  de- 
tailed study  belong  to  the  Unioninae  and  would  be  expected  to  be  typi- 
cally short-term  breeders.  In  Amblema  costata,  the  breeding  seasons 
extend  from  early  May  to  early  July.  The  breeding  season  for  Mega- 
lonaias  gigantea  extends  from  early  June  through  August.  The  breed- 
ing seasons  for  the  three  species  of  Quadrula  studied  here  extend 
through  May,  June,  and  July. 

Of  the  19  specimens  of  Tritogonia  verrucosa  reported  here,  there 
were  7  females  and  12  males.  Examination  of  the  testes  from  specimens 
collected  in  July  indicated  that  they  were  in  prebreeding  condition  and 
specimens  collected  in  November  and  early  May  contained  mature  sperm. 
The  ovary  of  a  specimen  taken  in  November  was  in  typical  breeding 
condition.  The  walls  of  the  primordium  were  thin,  the  vitelline  mem- 
branes were  well  developed,  amphinuclei  were  present,  and  the  ova  were 
crowded  in  the  lumen.  Ovaries  from  females  collected  in  June  and 
July  were  characteristically  in  prebreeding  condition.  Thus,  it  appears 
that  T.  verrucosa,  although  a  member  of  the  Unioninae,  is  a  long-term 


Ecology  225 

breeder.  However,  more  data  will  have  to  be  gathered  in  order  to  estab- 
lish the  precise  limits  of  the  breeding  season,  and  observations  must  be 
made  on  the  time  of  release  of  glochidia. 

The  only  evidence  of  sexual  dimorphism  among  the  mussels  studied 
was  in  individuals  of  T.  verrucosa.  In  that  species,  the  valves  of  the 
females  have  long  posterior  extensions  not  present  in  the  valves  of 
the  males.  There  is  no  evidence,  based  on  this  study,  of  any  relation- 
ship between  degree   of  obesity  and  maleness   or  femaleness. 

Acknowledgements 

This  study  was  supported  by  the  Indiana  Department  of  Natural 
Resources,  Division  of  Fish  and  Game,  under  Sub-Project  No.  4-10-R,  in 
cooperation  with  the  U.  S.  Department  of  the  Interior,  Bureau  of  Com- 
mercial Fisheries.  We  are  grateful  to  Messrs.  Woodrow  M.  Fleming  and 
R.  Eugene  Bass  of  the  Indiana  Division  of  Fish  and  Game;  to  Darrell 
Christensen  and  William  W.  Oakes  for  field  assistance;  to  Joe  K.  Neel 
for  initiating  the  project;  and  to  E.  Nelson  Cohen,  Terre  Haute,  Indiana, 
for  providing  crowfoot  bars  and  information  on  commercial  aspects  of 
mussel  shells  in  the  Japanese  cultured  pearl  industry. 

Literature  Cited 

1.  Baker,  F.  C.  1922.  The  molluscan  fauna  of  the  Big  Vermillion  River,  Illinois,  with 
special  reference  to  its  modification  as  a  result  of  pollution  by  sewage  and  manu- 
facturing wastes.  111.  Biol.  Monogr.  7:105-224. 

2.  Bingham,  R.  L.  1968.  Reproductive  seasons  of  eight  freshwater  mussels  from  the 
Wabash,  White,  and  East  Fork  of  the  White  rivers  of  Indiana.  Unpublished  M.S. 
Thesis.  Univ.  Louisville,  Louisville,  Ky.  102  p. 

3.  Blatchley,  W.  S.,  and  L.  E.  Daniels.  1902.  On  some  mussels  known  to  occur  in 
Indiana.  Annu.  Rep.  Ind.  Geol.  Surv.  26  :557-628. 

4.  Call,  R.  E.  1892.  A  contribution  to  a  knowledge  of  Indiana  Mollusca.  Proc.  Indiana 
Acad.  Sci.  3  :140-160. 

5.    .    1896a.    Second    contribution    to    a    knowledge    of    Indiana    Mollusca.    Proc. 

Indiana  Acad.  Sci.  6:135-146. 


1896b.    The   hydrographic   basins   of   Indiana   and   their   molluscan    fauna. 


Proc.  Indiana  Acad.  Sci.  6  :247-258. 


7. .    1900.    A    descriptive    illustrated    catalogue    of    the    Mollusca    of    Indiana. 

Annu.  Rept.  Geol.  Surv.  Ind.  24  :335-535. 

8.  Carlander,  H.   B.   1954.   A  history  of  fish  and  fishing  in   the  upper  Mississippi  River. 
Upper  Miss.  R.  Conserv.  Comm.  96  p. 

9.  Coe,  W.  R.,  and  H.  J.  Turner,   Jr.  1938.  Development  of  the  gonad  and  gametes  in 
the  soft-shell  clam,   (Mya  arenaria) .  J.  Morph.  62:91-111. 

10.  Coker,   R.   E.    1921.   Freshwater  mussels   and   mussel   industries   of   the   United   States. 
Bull.  U.  S.  Bur.  Fish.  36:13-89. 

11.  Daniels,    L.    E.    1903.    A    check-list   of   the    Indiana    Mollusca,    with   localities.    Annu. 
Rep.  Ind.  Geol.  Surv.  26  :629-652. 

12.    .    1914.   A   supplemental   check-list  of   Indiana  Mollusca,   with   localities    and 

notes.  Annu.  Rep.  Ind.  Dept.  Geol.  Natur.  Resources  39:318-326. 


226  Indiana  Academy  of  Science 

13.  Forbes,   S.   A.,   and   R.   E.   Richardson.   1919.   Some  recent  changes   in  Illinois  biology. 
Bull.  111.  State  Lab.  Natur.  Hist.  13:137-156. 

14.  Goodrich,   C,  and  H.   van  der  Schalie.   1944.   A   revision  of  the  Mollusca  of   Indiana. 
Amer.  Midland  Natur.  32(2)  :257-326. 

15.  Guyer,    M.    F.    1953.    Animal    micrology.    5th    ed.    The    University    of    Chicago    Press, 
Chicago.  327  p. 

16.  Hynes,   H.   B.   N.    1960.   The  biology  of  polluted  waters.   Liverpool  Univ.   Press,   Liver- 
pool, England.  202  p. 

17.  Kolkwitz,  R.,  and  M.   Marsson.    1909.   okologie  der  tierische  Saprobien.   Beitrage   zur 
Lehre    von    der    biologische    Gewasserbeurteilung.    Int.    Rev.    ges.    Hydrobiol.    2  :  126-152. 

18.  Krumholz,  L.  A.   1946.  Repopulation  of  the  West  Fork.   Outdoor  Indiana  13  ((3)  :12. 


19.  -  — ,  and  W.  L.  Minckley.  1964.  Changes  in  the  fish  population  in  the  upper 
Ohio    River    following    temporary    pollution    abatement.    Trans.    Amer.     Fish.    Soc.    93 

(l):l-5. 

20.  LlLLlE,  F.  R.   1895.  The  embryology  of  the  Unionidae.  J.  Morph.  10(1)  :  1-100. 

21.  Loosanoff,  V.   L.   Reproductive  cycle  in  Cyprina  islandica.  Biol.   Bull.   104(2)  :146-155. 

22.  Meyer,  E.  R.  1968.  The  distribution  and  abundance  of  freshwater  mussels  of  the 
family  Unionidae  (Pelecypoda)  of  the  Wabash,  White,  and  East  Fork  of  the  White 
rivers  of  Indiana.   Unpublished  M.S.  Thesis.  Univ.  Louisville,  Louisville,  Ky.  68  p. 

23.  Ortmann,  A.  E.  1909.  The  destruction  of  the  freshwater  fauna  in  western  Pennsyl- 
vania. Proc.  Amer.   Phil.  Soc.  48:90-110. 

24.  Pennak,    R.   W.    1953.    Freshwater   invertebrates   of  the   United   States.    Ronald    Press. 

New  York.  769  p. 

25.  Simpson,   C.   T.   1900.   Unionidae  of   Indiana.   Nautilus   14  :95-96. 

26.  Stein,  F.  1881.  The  molluscous  fauna  of  Indiana.  Annu.  Rep.  Ind.  Geol.  Surv.  16  :451- 
467. 

27.  Stephens,   W.   M.   1963.  Man-made  pearls.   Sea  Frontiers  9  (5)  :299-308. 

28.  van  der  Schalie,  H.  1938.  Hitch-hiking  mussels  and  pearl  buttons.  Mich.  Conserv. 
7:4-5,  11. 

29.  Walker,  B.  1917.  The  method  of  evolution  in  the  Unionidae.  Occas.  Pap.  Univ.  Mich. 
Mus.  Zool.  45:1-10. 

30.  Wurtz,  C.   B.   1956.  Freshwater  mollusks  and  stream  pollution.  Nautilus  69  (3)  :97-102. 


ENTOMOLOGY 

Chairman:  Jack  R.  Munsee,  Indiana  State  University 
John  W.  Hart,  Earlham  College,  was  elected  Chairman  for  1970 

ABSTRACTS 
The  Use  cf  Heartbeat  as  a  Potential  Screening  Technique  for  Insect 
Pathogens.  Mildred  G.  Ware  and  Harold  L.  Zimmack,  Ball  State 
University. — The  purposes  of  this  study  were:  1)  to  develop  a  rapid 
and  accurate  screening  technique  which  will  determine,  by  observing 
changes  in  heartbeat,  whether  or  not  a  specific  species  of  bacteria  is 
pathogenic  to  a  specific  insect  and  2)  to  determine  criteria  which  would 
make  it  possible  to  apply  this  technique  to  other  insects. 

On  the  basis  of  these  data,  it  is  possible  that  the  heartbeat  of  the 
European  corn  borer,  Ostrinia  nubilalis  (Hubner),  can  be  used  to  detect 
bacterial  pathogens. 

Results  obtained  indicate  that  the  non-pathogenic  species  of  bac- 
teria, Escherichia  coli  (Migulo),  increased  the  mean  post-ingestion  heart 
rate  4.36  heartbeats/minute;  whereas  mean  post-ingestion  heart  rates  in 
the  larvae  ingesting  the  pathogenic  species  of  bacteria,  Bacillus  thurin- 
giensis  Berliner,  Serratia  marcescens  Bizi  and  Bacillus  subtilis  Cohn, 
decreased  19.44  to  24.21  heartbeats/minute. 

The  investigators  believe  that  age  had  a  definite  influence  on  the 
heart  rates  and  the  susceptibility  of  the  larvae  to  pathogenicity.  In  the 
test  group  using  B.  thuringiensis,  reported  to  be  the  most  vigorous  of 
the  experimental  pathogens,  all  late  fifth  instar  larvae  were  used.  The 
mean  pre-  and  post-ingestion  heart  rates  for  this  group  were  69.68 
heartbeats /minute  and  50.24  heartbeats  /minute,  respectively.  These  fig- 
ures are  significantly  lower  than  the  mean  pre-  and  post-ingestion  heart 
rates  of  early  fifth  instar  larvae  used  to  test  the  effects  of  the  moderate 
pathogenic  species,  S.  marcescens,  and  the  controversial  pathogen,  B. 
subtilis.  The  mean  figures  in  the  S.  marcescens  test  group  were  96.39 
heartbeat/minute  and  72.18  heartbeats/minute  and  in  the  B.  subtilis 
test  group  the  mean  pre-  and  post-ingestion  heart  rates  were  96.77 
and   74.78   heartbeats/minute. 

Further  Studies  on  the  Interbreeding  of  an  Insular  Form  of  Tropis- 
ternus  collaris  (Castelnau)  with  Mainland  Forms.  Frank  N.  Young, 
Indiana  University. — Experimental  crosses  of  a  form  of  Tropisternus 
collaris  from  Puerto  Rico  indicate  that  the  color  pattern  shows  partial 
or  complete  dominance  over  that  of  forms  from  Florida,  Indiana,  and 
Mexico.  Such  crosses  are,  however,  highly  sterile  or  infertile.  Crosses 
with  a  melanic  form  from  Colombia  (Ayapel)  show  dominance  of  color 
pattern  but  nearly  complete  fertility  of  progeny.  Backcrosses  have 
produced  interesting  examples  of  particulation  of  the  elements  of  the 
color  pattern. 

OTHER  PAPER  READ 
On  the  Nature  of  Communication  of  Bees.  Harold  Esch,  University  of 
Notre  Dame  (by  invitation). 

227 


The  Second  Record  of  Coelioxys  obtusiventris 
Crawford  (Hymenoptera,  Megachilidae)1 

Leland  Chandler,  Purdue  University 

Abstract 

The  parasitic  bee,  Coelioxys  obtusiventris  Crawford,  was  described  in  1914  from 
Florida  and  has  been  known  only  from  the  unique  female  holotype.  A  second  specimen  has 
now  been  collected  at  Lafayette,  Indiana,  and  is  recorded. 

The  parasitic  megachilid,  Coelioxys  obtusiventris  Crawford,  was 
described  on  the  basis  of  a  single  female  located  in  the  C.  F.  Baker 
Collection  (1).  The  specimen  bore  a  label  "Florida;  Palm."  without 
additional  data.  The  holotype  is  Type  Specimen  Catalog  Number  18217, 
U.S.  National  Museum  (1,  2).  Mitchell  (2)  redescribed  the  species  and 
indicated  a  possible  relationship  with  C.  modesta  Smith. 

Females  of  C.  modesta  and  C.  obtusiventris  characteristically  have 
the  apex  of  the  sixth  metasomal  tergite  upturned  abruptly;  however, 
a  number  of  individuals  of  C.  modesta  do  not  exhibit  this  condition.  In 
either  case,  the  apical  third  of  this  tergite  (in  C.  modesta)  bears  nu- 
merous long,  erect,  black  hairs,  but  these  are  not  sufficiently  dense  to 
obscure  the  upturned  portion  when  present.  Comparatively,  this  por- 
tion of  the  apical  tergite  of  C.  obtusiventris  is  covered  so  extensively 
with  brownish-black  hairs  that  the  upturned  portion  is  nearly  hidden. 

The  most  distinguishing  structure  of  C.  obtusiventris  is  the  sixth 
metasomal  sternite,  described  by  Mitchell  (2)  as  ".  .  .;  sternum  6 
slightly  flared  apically,  with  a  long  apical  spine,  apical  margin  with  a 
prominent  fringe  of  long  brownish  hairs,  nearly  equally  the  spine  in 
length."  This  feature  is  so  striking  and  differs  so  greatly  from  the 
apical  sternite  of  other  species  that  its  diagnostic  value  has  probably 
been  under-emphasized  within  longer  descriptions. 

The  second  specimen  of  C.  obtusiventris  was  identified  among  ma- 
terial in  the  Purdue  Entomological  Research  Collection.  The  specimen, 
a  female,  bears  the  label  "Lafayette,  Ind.  VI  •  16  •  59."  No  other 
information  is  available  nor  was  the  bee  associated  with  other  speci- 
mens of  Coelioxys  or  of  the  host  genus  Megachile. 

The  status  of  species  based  upon  unique  specimens  usually  creates 
a  puzzling  situation,  especially  when  localities  are  indefinite.  A  second 
record  from  a  remote  region  does  not  reduce  the  puzzle,  but  it  does 
confirm  a  continued  existence. 

Literature  Cited 

1.  Crawford,   J.   C.    1914.    Some  species  of  the  bee  genus  Coelioxys.   Ann.   Entomol.    Soc. 
America  7(2)  :148-159. 

2.  Mitchell,  T.  B.  1962.  Bees  of  the  eastern  United  States.  Vol.  II.  North  Carolina  Agric. 
Exp.  Sta.  Tech.  Bui.  No.  152.  557  p.   (esp.  214-215). 


1  Journal  Paper  No.   2963  of  the   Purdue  University  Agricultural  Experiment  Station. 

228 


Indiana  vs.  Indian  Territory :  Misinterpreted  Locality  Citations 

Leland  Chandler,  Purdue  University 


Abstract 

A  number  of  insect  species  have  been  erroneously  recorded  from  Indiana,  whereas  the 
localities  are  actually  in  Oklahoma.  An  examination  of  locality  labels  on  some  of  these 
specimens  show  the  abbreviation  [Ind.].  Whether  some  designation  of  Territory  was 
included  originally  and  removed,  or  perhaps  not  printed  originally,  is  not  known.  Several 
zoogeographic  interpretations  are  based  on  these  resulting  in  erroneous  conclusions. 

Modern  systematic  treatments  include  data  from  numerous  disci- 
plines. Thence,  by  analysis  of  these  data,  certain  zoogeographic,  eco- 
logical, and  phylogenetic  concepts  are  derived.  A  basic  source  of  these 
data  is  the  locality  label. 

This  paper  is  devoted,  in  part,  to  a  clarification  of  one  series  of 
locality  labels  which  has  been  misinterpreted.  Fortunately,  reinterpre- 
tation  does  not  result  in  major  changes  for  most  of  the  species  involved. 

Mitchell  (4)  recorded  the  leaf-cutter  bee,  Megachile  p.  parallela 
Smith,  from  South  McAlister,  the  site  listed  as  an  Indiana  locality. 
Since  this  bee  does  occur  in  the  state,  no  significance  was  attached  to 
this  specific  place.  Stephen  (6)  recorded  the  silk  bee,  Colletes  mandi- 
bularis  Smith,  from  Macalester,  also  credited  to  Indiana.  This  species, 
likewise,  occurs  throughout  the  state.  La  Berge  (3)  recorded  three  spe- 
cies of  bees  from  Indiana  as  follows:  Svastra  (as  Melissodes)  o.  obliqua 
(Say)  from  McAllister;  Svastra  (as  Melissodes)  p.  petulca  (Cresson) 
from  South  McAlester;  and  Melissodes  c.  communis  Cresson  from  South 
McAllester.  Of  the  three  species,  S.  petulca  is  the  only  one  not  known 
to  occur,  being  of  more  southern  and  western  distribution.  Deleting  the 
Indiana  record  from  the  distributional  map  [(3),  fig.  9,  pg.  1010]  re- 
sults in  a  somewhat  different  configuration.  On  this  basis,  it  would  be 
predicted  that  this  species  is,  however,  likely  to  occur  within  the  state  in 
the  Lower  Wabash  Valley  or  "Pocket"  Biotic  Unit  (2). 

As  the  number  of  references  to  this  locality  increased,  in  addition 
to  the  various  spellings,  it  became  important  to  discover  the  exact 
location  and  the  reasons  for  the  interpretations.  With  but  meagre  facts 
to  substantiate  the  conclusions,  it  was  reasonable  to  accept  the  idea 
that  all  records  should  be  attributed  to  McAlester,  Oklahoma.  This 
would  mean  that  labels  read  as  "Indiana"  were  in  fact  "Indian  Territory." 

Confirmation  that  such  an  idea  was  correct  has  been  found  in  the 
recent  publications  by  Campbell  (1)  and  Quate  and  Thompson  (5). 
Campbell  recorded  Lobopoda  yiigrans  (Melsheimer)  from  South  McAles- 
ter and  Atoka,  both  localities  given  as  being  in  Indiana.  The  inclusion 
of  Atoka,  also  an  Oklahoma  locality,  further  represents  a  label  mis- 
interpretation. 

The  reference  by  Quate  and  Thompson  strangely  does  not  have 
reference  to  Indiana,  but  to  Arkansas.  It  does,  however,  give  complete 

229 


230  Indiana  Academy  of  Science 

label  information  as  follows:  "Vinita,  Ind.  T.,  June  7-8,  '99,  Wickham." 
The  species  referred  to  is  Melanotus  lanei  Quate  and  the  locality  is 
cited  as  being-  in  Arkansas. 

It  has  become  apparent  that  one  or  more  collections  of  beetles  and 
of  bees  contains  materials  with  labels  that  can  be  misread.  Assuredly,  all 
references  to  Atoka,  McAlester  (and  its  derivative  spellings),  and  Vinita 
are  Oklahoma  localities,  not  Indiana  or  Arkansas. 


Literature  Cited 

1.  Campbell,  John  M.  1966.  A  revision  of  the  genus  Lobopoda  (Coleoptera  :Alleculidae) 
in  North  America  and  the  West  Indies.  Illinois  Biol.  Monog.  37,  Univ.  Illinois  Press, 
Urban  a. 

2.  Chandler,  Leland.  1966.  The  origin  and  composition  of  the  insect  fauna,  p.  345-361. 
In  A.  A.  Lindsey  [ed.]  Natural  Features  of  Indiana.  Indiana  Acad.  Sci.  Sesquicenten- 
nial  Volume,  Indianapolis.  600  p. 

3.  La  Berge,  Wallace  E.  1956.  A  revision  of  the  bees  of  the  genus  Mclissodcs  in  North 
and  Central  America.  Part  1  (Hymenoptera,  Apidae).  Univ.  Kansas  Sci.  Bull.  37,  Pt.  2 
(18)  :911-1194. 

4.  Mitchell,  T.  B.  1937.  A  revision  of  the  genus  Megachile  in  the  Nearctic  Region.  Part 
Vi.  Taxonomy  of  subgenera  Argyropile,  Leptorachis,  Pseudocentron,  Acentron  and 
Melanosarus.   (Hymenoptera  :Megachilidae).  Trans.  American  Entomol.  Soc.  63:45-83. 

5.  Quate,  Laurence  W.,  and  Sarah  E.  Thompson.  1967.  Revision  of  click  beetles  of 
genus  Melanotus  in  America  north   of  Mexico.   Proc.   U.   S.   Nat.   Mus.    121    (3568)  :l-83. 

6.  Stephen,  W.  P.  1954.  A  revision  of  the  bee  genus  Colletes  in  America  north  of 
Mexico.  Univ.   Kansas  Sci.  Bull.  36,  Pt.  1   (6)  :149-527. 


The  Occurrence  of  Chalybion  zimmermanni  Dahlbom  (Sphecidae) 

in  Indiana 

Gertrude  L.  Ward,  Earlham  College 

Abstract 

Chalybion  zimmermanni  Dahlbom  (Hymenoptera,  Sphecidae)  is  added  to  the  list  of 
insects  of  Indiana.  The  northern  extent  of  this  insect's  range  was  shown  by  Bohart  in  1963 
as  Tennessee.  It  was  found  nesting  in  holes  in  wood  in  Wayne  County,  Indiana,  in  1968, 
and  in  1969  evidence  of  nesting  was  found  in  six  other  counties.  Nests  are  provisioned  with 
small  spiders,  generally  Argiopidae  or  Theridiidae.  An  unusual  two-layered  plug  is  made 
of  mud  and  uric  acid.  The  white  uric  acid  contrasts  distinctly  with  the  old  wood  around 
the  nest  hole.  A  small  number  of  nests  were  located  in  deserted  nests  of  the  yellow-legged 
mud  dauber,  Sceliphron  caementarium. 

During  a  recent  study  of  three  mud-using  wasps,  Sceliphron  cae- 
mentarium (Drury),  Trypargilum  politum  (Say),  and  Chalybion  cali- 
fornicum  (Saussure),  another  large  mud-using  wasp  appeared  in  my 
study  area  near  Centerville,  Wayne  County,  Indiana.  This  was  Chaly- 
bion zimmermanni  Dahlbom.  On  1  August  1968,  an  assistant,  Jay 
Myers,  called  my  attention  to  the  metallic  blue  wasp  which  was  plugging 
a  hole  in  the  wooden  plate  of  an  old  tool  shed.  The  original  borer  was 
probably  a  beetle. 

We  watched  this  wasp  clean  the  debris  out  of  a  second  hole,  gather 
mud  from  old  Sceliphron  caementarium  nests  and  form  a  barricade 
deep  in  the  hole.  She  collected  small  spiders,  laid  an  egg  on  the  abdomen 
of  one  of  the  first  spiders,  and,  on  two  occasions,  built  a  wall  about 
halfway  down  the  hole.  The  remaining  space  was  used  for  a  second  cell. 
When  this  was  filled,  the  female  formed  a  countersunk  plug  of  dark 
mud.  This  sometimes  required  three  or  four  loads  of  mud,  apparently 
all  of  it  gathered  from  old  nests. 

After  the  dark  mud  was  dry,  she  made  a  level  or  slightly  concave 
seal  of  white  material.  We  collected  some  of  this  white  plaster  from  one 
cell,  and  analysis  by  infra-red  spectrophotometry  showed  that  it  was 
mainly  uric  acid.  The  wasp  disappeared  after  completing  six  holes. 

In  May,  1969,  six  small  cylindrical  traps  made  of  fiberglass  screen- 
ing were  fastened  over  the  white-plugged  holes.  Emergence  of  wasps 
from  these  cells  was  noticed  first  on  28  June.  Nothing  emerged  from  one 
cell.  Two  males  were  collected  and  are  in  the  Joseph  Moore  Museum  of 
Earlham  College.  Five  wasps,  including  both  males  and  females,  were 
released.  A  summary  of  the  adults  which  emerged  is  shown  in  Table  1. 

Several  of  the  wasps  which  emerged  were  chilled  in  a  glass  jar  in  a 
food  freezer  for  about  4  minutes,  and  then  were  measured  before  their 
release.  The  clypeal  teeth  were  examined  to  determine  the  sex.  The  male 
has  a  median  tooth  which  is  longer  than  either  of  the  side  teeth,  and  the 
female  has  three  small  flaps  with  the  middle  one  not  shorter  than  the 
others,  as  it  is  in  Chalybion  calif  or  nicum. 

231 


232  Indiana  Academy  of  Science 

Table  1.     Adults  emerging  from  cells  of  Chalybion  zimmermanni  Dahl- 

bom,  Indiana,  1969. 


Number 

Body 

Cell 

emerging 

length  (mm) 

Sex 

A 

1 

18 

F  (?) 

B 

2 

19,  15.5 

F,  M 

C 

0 

— ■ 

— 

D 

1 

16 

M 

E 

2 

18,  16 

F  (?),  M 

F 

1 

large 

F   (?) 

Two  females  started  to  make  nests  in  old  holes  in  the  wooden  plate 
on  16  July  1969.  Eight  holes  were  filled,  four  of  which  had  been  occupied 
in  1968. 

An  attempt  was  made  in  1969  to  determine  the  extent  of  C.  zimmer- 
manni penetration  into  Indiana.  In  1963,  Bohart  and  Menke  (1)  reported 
the  northern  extent  of  the  range  as  Tennessee.  Although  no  other 
specimens  were  collected,  the  distinctive  two-layered,  two-colored  plugs 
were  found  in  the  following  counties:  Clark,  Crawford,  Dearborn,  Ohio, 
Ripley,  and  Switzerland.  An  equal  amount  of  collecting  yielded  negative 
results  in  Decatur,  Jackson,  Lawrence,  Rush,  and  Washington  counties. 

Most  of  the  cells  were  in  the  old  timbers  of  barns  and  sheds,  but  in 
Clark  County  four  of  these  plugs  were  found  in  a  mass  of  deserted 
Sceliphron  caementarium  cells.  Rau  (5)  reported  that  in  Mexico  C. 
zimmermanni  uses  the  old  cells  of  Sceliphron  assimilis  (Dahlbom). 

A  total  of  40  spiders  was  examined  from  C.  zimmermanni  nests. 
The  majority,  60%,  were  in  the  family  Araneidae,  and  the  remaining 
40%  were  in  the  Theridiidae.  These  are  shown  in  Table  2. 

Table  2.     Prey  of  Chalybion  zimmermanni  Dahlbom  in  Indiana,  1969. 


No. 

% 

Total  % 

Araneidae 

Araneus  spp. 

VI 

30.0 

Argiope  aurantia  Lucas 

9 

22.5 

Argiope  trifasciata  (Forskal) 

2 

5.0 

Cyclosa  conica  (Pallas) 

1 

2.5 

60.0 

Theridiidae 

Theridion  frondeum  Hentz 

15 

37.5 

Asagena  americana  Emerton 

1 

2.5 

40.0 

40 

100.0 

Entomology  233 

Whether  or  not  the  uric  acid  serves  as  a  deterrent  to  parasites 
which  might  invade  the  completed  cell  through  the  plug  has  not  been 
determined.  The  use  of  two  colors  of  material  in  the  final  plug  was 
reported  by  Williams  (6)  for  Chalybion  violaceum  (Fabricius)  in  the 
Philippines.  He  said  that  this  wasp  ".  .  .  stores  her  small  spiders  in 
some  convenient  hollow,  as  a  rung  socket,  penholder  base,  old  mud  nest, 
etc.,  and  simply  plugs  up  the  aperture,  first  with  mud  or  moist  earth,  and 
finishes  this  off  with  a  mixture  of  the  excreta  of  geckos  (lizards),  giving 
the  plug  a  whitish  or  plaster-like  appearance." 

Williams  (6)  also  reported  that  d'Herculais  in  1882,  observing 
Chalybion  chalybeiis  (Smith)  at  Port  Natal,  Africa,  noticed  "this  curious 
habit"  of  using  a  light-color  final  plug.  The  material  used  at  Port  Natal 
was  bird  excrement. 

Iwata  (2)  reported  that  he  saw  Sceliphron  (Chalybion)  inflexum 
Sickmann  on  Taiwan  plaster  the  mud  seal  of  her  nest  with  white 
material,  and  also  (3)  reported  this  in  Thailand.  Yamamoto  (7)  said 
that  he  saw  this  wasp  in  Japan  collecting  bird  droppings  which  were 
still  damp,  and  using  them  for  plastering. 

In  India,  Jayaker  and  Spurway  (4)  observed  Chalybion  bengaleyise 
Dahlbom  make  a  plug  of  brown  mud  and  then  cover  it  with  a  plug  of 
white.  One  wasp  collected  the  white  material  from  the  feces  of  a  pet 
tortoise  and  another  wasp  used  bird  feces. 

Summary  and  Conclusions 

Chalybion  zimmewnanni,  having  the  northern  edge  of  its  range 
reported  in  1963  as  Tennessee,  appears  to  be  moving  northward  in 
Indiana.  The  food  stored  for  the  young  is  small  spiders  from  the 
families  Araneidae  and  Theridiidae.  It  most  often  nests  in  borings  in  old 
timbers,  but  has  been  seen  to  use  the  deserted  mud  nests  of  Sceliphron 
caementarium. 

Acknowledgments 

The  author  wishes  to  express  appreciation  to  Alan  Rushton  for 
spectrophotometric  analysis,  to  National  Science  Foundation  for  an 
undergraduate  assistant  under  Grant  GY  4495,  and  to  Research  Corpora- 
tion for  a  portion  of  the  institutional  grant  to  Earlham  College. 

Literature  Cited 

1.  BOHART,  R.  M.,  and  A.  S.  Menke.  1963.  A  reclassification  of  the  Sphecinae  with  a 
revision  of  the  nearctic  species  of  the  tribes  Sceliphronini  and  Sphecinae.  Univ.  Calif. 
Berkeley.  Pub.  Entomol.  30:19-182. 

2.  Iwata,  K.  1939.  Habits  of  some  solitary  wasps  in  Formosa  (IV).  Trans.  Natur.  Hist. 
Soc.  Formosa  29:161-178. 

3.  Iwata,  K.  1964.  Bionomics  of  non-social  wasps  in  Thailand.  Nature  and  Life  in 
Southeast  Asia  3  :323-383. 

4.  Jayaker,  S.  D.,  and  H.  Spurway.  1963.  Use  of  vertebrate  faeces  by  the  sphecoid  wasp 
Chalybion  bengalense  Dahlb.  J.  Bombay  Natur.  Hist.  Soc.  60  :748-749. 

5.  Rau,  P.  1943.  The  nesting  habits  of  certain  Sphecid  wasps  of  Mexico,  with  notes  on 
their  parasites.  Ann.  Entomol.  Soc.  Amer.  36  :647-653. 

6.  Williams,  F.  X.  1919.  Philippine  wasp  studies.  Bull.  Expt.  Sta.  Hawaiian  Sugar 
Planters'  Assoc,  Entomol.  Ser.  No.  14  :19-181. 

7.  Yamamoto,  D.  1942.  Habits  of  Sceliphron  (Chalybion)  inflexum  Sickmann.  Kontyu 
16  :69-75. 


A  Taxonomic  Key  to  the  Collembola 
in  Four  Serai  Stages  Leading  to  the  Beech-Maple  Climax 

Patricia  M.  Arnetti,  Indiana  State  University 


Abstract 

From  April  through  July,  1968,  96  leaf  litter  samples  were  taken  from  an  old  field, 
oak  and  maple-oak  dominated  serai  stages,  and  a  beech-maple  climax  in  Parke  County, 
Indiana.  Collembola  were  extricated  by  a  modified  Tullgren  funnel  apparatus,  collected, 
and  identified.  A  key  was  based  on  morphology  and  color  for  32  species,  which  represent 
20  genera  and  5  families.  A  table  of  the  distribution  of  each  species  by  serai  stage  was 
included. 

Introduction 

From  April  through  July,  1968,  96  leaf  litter  samples  of  1.0  dm^ 
each  were  taken  from  an  old  field,  oak  and  maple-oak  dominated  serai 
stages,  and  a  beech-maple  climax  in  Allee  Woods,  Parke  County, 
Indiana.  Collembola  were  extricated  by  a  modified  Tullgren  apparatus 
and  identified  (2,  3).  With  current  keys,  not  all  individuals  could  be 
identified  to  species.  The  key  represents  1,533  individuals,  32  species, 
20  genera  and  5  families.  The  purpose  of  this  paper  was  to  present  a 
simplified  key  to  the  species  in  the  four  serai  stages  leading  to  the 
beech-maple  climax.  The  changes  in  Collembola  populations  as  influenced 
by  plant  successional  patterns  was  previously  described  (1). 

Method 

After  all  species  were  identified,  the  most  obvious  external  char- 
acteristics were  selected  for  these  32  species  and  a  key  was  constructed. 
The  key  was  based  on  color  and  morphology.  The  primary  morphological 
characteristics  were:  1)  length  and  shape  of  the  body;  2)  degree  of 
fusion  and  length  of  abdominal  segments;  3)  nature  of  prothorax; 
4)  presence  of  scales,  body  hair,  and  setae;  5)  number  of  eyespots;  and 
6)  number  of  segments  and  length  of  antennae.  The  distribution  of  each 
species  by  serai  stage  (Table  1)  shows  the  relative  abundance  of  each 
species  per  ecological  area  and  hence  could  be  helpful  in  confirming  an 
identification  of  an  individual  from  a  comparable  sere. 

Taxonomic  Key 

1.  Body  elongate;  abdominal  segments  distinct  although   IV,  V,  and  VI  or  V  and  VI 
may    be    ankylosed    2 

suborder  Arthropleona  Borner 

1'.   Body   globular ;    abdominal   segments   not   distinct ;   the   first   four   abdominal    seg- 
ments fused  with  thorax 26 

suborder  Symphypleona  Borner 
family  Sminthuridae 

2.  Prothorax  reduced  and  membraneous 3 

superfamily  Entomobryoidea  Womersley 

2'.   Prothorax   similar  to  other  segments   22 

superfamily  Poduroidea  Womersley 


Present  address:  Northwestern  High  School,  Kokomo,  Indiana  46901. 

234 


Entomology  235 

3.  Scales  absent ;  body  segments  equal  to  subequal  in  length ;  antennae  with  4 
simple  segments  ;   last  abdominal  segments  may  be  ankylosed 4 

family  Isotomidae 

3'.  Scales  and/or  brush-like  setae  present ;  third  or  fourth  body  segment  elongate ; 
antennae  with  4-6  segments,  the  third  and  fourth  sometimes  annulated  ;  abdominal 

segments   always  distinct 11 

family  Entomobryidae 

4.  Fourth,   fifth,  and  sixth  abdominal  segments  ankylosed 5 

Folsomia 

4'.   Fourth,  fifth,  and  sixth  abdominal  segments  not  ankylosed 6 

5.  E>es   absent;   pigment   absent   

Folsomia  fimentaria  L. 

5'.   Eyes  2  and  2  ;  pigment  gray  to  black   

Folsomia  quadrioculata  Tullberg 

6.  Body  with   bothriotrichia    (long  sensory  body  hairs)    

Isotomurus  palustris  Muller 
6'.   Body    without    bothriotrichia    7 

7.  Manubrium  (single  part  of  furcula ;  is  attached  to  abdomen)  much  shorter  than 
dentes  (middle  part  of  furcula;  is  forked),  with  many  ventral  setae;  Abd.  IV 
usually  shorter  than   Abd.   Ill   8 

Isotoma 
7'.   Manubrium    often    longer    than    dentes    with    few    or    no   ventral    setae ;    Abd.    IV 

usually   longer   than    Abd.    Ill    10 

Proisotoma 

8.  Eyes  4  and  4  on  round  patches  connected  by  an   inverted  V-shaped  mark 

* Isotoma  eunotabilis  Folsom 
8'.   Eyes  8  and  8  on  elongate  patches  without  an  inverted   V-shaped  mark 9 

9.  Length    0.6   mm    

Isotoma  viridis  Bourlet 

9'.   Length    1.5    mm    

*Isotoma  olivacea  Tullberg 

10.  Dentes   shorter   than   manubrium    

Proisotoma  minuta  Tullberg 

10'.   Dentes   longer  than   manubrium   

*Proisotoma  immersa  Folsom 

11.  Abd.   Ill  longer  than  Abd.   IV;  mucrones    (tip  of  furcula)   hairy;  Ant.   Ill  longest 
segment   and   annulated;   antennae   4-segmented   12 

subfamily  Tomocerinae 

Tomocerus 

11'.   Abd.  Ill  shorter  than  Abd.  IV;  mucrones  not  hairy;  antennae  4-  to  6-segmented  __     15 

subfamily  Entomobryinae 

12.  Maxilla     bearded     13 

12'.  Maxilla  not  bearded   14 

13.  Antennae  longer  than   body 

*Tomocerus  elongatus  Maynard 

13'.  Antennae  shorter  than  body 

*  Tomocerus  flavescens  Tullberg 

14.  Dental  spines  tridentate;  Th.   II  overlapping  but  not  obscuring  Th.   I  dorsally 

*Tomocerus  minor  Lubbock 

14'.  Dental  spines  simple;  Th.   II  obscuring  Th.   I  dorsally 

*Tomocerus  vulgaris  Tullberg 

15.  Antennae   with   6   segments    

Orchesella  ainsliei  Folsom 
15'.  Antennae  with  4  segments 16 

16.  Body   without  scales   17 


236  Indiana  Academy  of  Science 

16'.   Body  with  scales   21 

Lepidocyrtus 

17.  Eyes  not  on  daik  patches 

*Isotobryoides  ochracius  Maynard 

17'.  Eyes   on   dark  patches   18 

Entomobrya 

18.  Body  unicolorous  without  crossbands   of  contrasting  color 19 

18'.   Body    with    dark    dorsal    and    lateral    spots    or    bands    or    both    on    light    ground 

color 20 

19.  Color  gray  to  olive  green  to  bluish  purple 

Entomobrya  marginata  Tullberg 

19'.   Color   yellow   to    yellow-orange   

^Entomobrya  atrocincta  f. 
pseudopcrpulchra  Mills 

20.  Transverse  bands  on  every  segment 

*  Entomobrya  multifasciata  Tullberg 

20'.  Transverse  bands  on  most  segments  ;  Abd.   I  with  2  dark  dorsal  spots 

Entomobrya  assuta  Folsom 

21.  Purple  pigment  on  Abd.   IV 

*  Lepidocyrtus  unifasciatus  James 

21'.   Purple  pigment  on   antennae  and  legs.   

Lepidocyrtus  curvicollis  Bourlet 

22.  Eyes  absent 

family  Onychiuridae 
*Onychiuru8  armatus  Tullberg 

22'.  Eyes  present 23 

family  Poduridae 

23.  Pigment    present    24 

23'.   Pigment   absent   

*Neanura  barberi  Handschin 

24.  Furcula    well    developed    25 

24'.   Furcula  reduced 

*Xenylla  welchi  Folsom 

25.  Color  brown  and  yellow  mottled ;  abdomen  not  considerably  distended 

*Hypogastrura  tigrina  Harvey 

25'.   Color  dark  gray  to  black  ;  abdomen  considerably  distended 

*Pseudachorutes  simplex  Maynard 

26.  Antennae    shorter    than   head;    eyes   absent    27 

Neelus 
26'.  Antennae  longer  than  head  ;  eyes  present 28 

27.  Color   white   

*Neelus  albus  Maynard 

27'  Color  yellow  with  red  speckles 

*Neelus  maculosus  Maynard 

28.  Color  purplish  red 

*Arrhopalites  binoculatus  Borner 
28'.   Color  not  purplish  red 29 

29.  Black   pigment   spots    present   on    abdomen    30 

29'.   Black  pigment  spots  absent;  eyes  orange 

Katiannina  maegillivrayi  Banks 

30.  Antennae    pale    basally    

*Dicyrtomina  variabilis  Maynard 

30'.   Antennae    dark    basally    31 

Sminthtirinus 


Entomology 


237 


31.    Body   with   much   dark   purplish   black   pigment ;    antennae   and   legs   banded   with 

dark  pigment 

*Sminthurinus  radiculus  Maynard 

31'.   Body  with   black   pigment  reduced   in   form   of  lateral   polygons;   buff   and   orange 

spots    present    

*Sminthurinus  radiculus  f.  pictus  Maynard 


Species  first  reported  for  Indiana. 

Table  1.    Species  distribution  by  serai  stage. 


Beech- 

Maple- 

Old 

Species 

maple 

<  »ak 

Oak 

field 

Total 

Arrhopalites  binoculatus 

2 

1 

3 

1 

7 

Dicyrtominia  variabilis 

0 

1 

0 

o 

1 

Entomobrya  assuta 

42 

7 

it; 

s 

73 

Entomobrya  atrocincta 

f.   pseudoperpulchra 

L6 

8 

41 

2 

67 

Entomobrya  marginata 

13 

8 

10 

11 

42 

Entomobrya  multij asciata 

44 

69 

121 

19 

253 

Folsomia  fimentaria 

10 

31 

is 

4 

63 

Folsomia  quadrioculata 

6 

1 

0 

0 

7 

Hypogastrura  tigrina 

o 

0 

1 

1 

2 

Isotobryoides  ochracius 

35 

49 

42 

2 

L28 

Isotoma  eunotabilis 

1 

2 

3 

o 

6 

Isotoma  olivacea 

8 

ir, 

46 

16 

85 

Isotoma  viridis 

0 

1 

0 

0 

1 

Isotomurus  palustris 

2 

2 

3 

0 

7 

Katiannina  macgillivrayi 

2 

1 

2 

1 

6 

Lepidocyrtus  curvicollis 

5 

3 

1 

0 

9 

Lepidocyrtus  unifasciatus 

0 

2 

0 

0 

2 

Neanura  barberi 

0 

0 

2 

0 

2 

Neelus  albus 

1 

3 

2 

0 

6 

Neelus  maculosus 

0 

o 

1 

0 

1 

Onychiurus  armatus 

94 

124 

43 

4 

265 

Orchesella  ainsliei 

(i 

0 

o 

I 

1 

Proisotoma  immersa 

2 

6 

14 

1 

23 

Proisotoma  minuta 

(l 

0 

5 

0 

5 

Pseudachomtes  simplex 

1 

1 

\ 

o 

<■. 

Sminthurinus  radiculus 

2 

1 

0 

0 

3 

Sminthurinus  radiculus 

f.  pictus 

5 

3 

17 

0 

25 

Tomocerus  elongatus 

1 

1 

1 

0 

3 

Tomocerus  flavescens 

2 

2 

2 

0 

<; 

Tomocerus  minor 

i:\ 

74 

L15 

36 

298 

Tomocerus  vulgaris 

1 

3 

38 

5 

47 

Xenylla  welchi 

20 

10 

33 

20 

S3 

Literature  Cited 

1.  Arnett,   Patricia  M.    1969.   A  Study  of  Collembolan  populations   associated  with   four 
serai   stages   leading   to  the   beech-maple   climax.    Proc.    Indiana   Acad.    Sci.    78 :231-240. 

2.  Folsom,    J.    W.    1937.    Nearctic    Collembola    or    springtails,    of    the    family    Isotomidae. 
Bull.  U.  S.  Natur.  Mus.  168.  145  p. 

3.  Maynard,    E.    C.    1951.    The    Collembola    of    New    York    State.    Comstock    Publishing 
Company,  Inc.,  Ithaca,  New  York.  388  p. 


Factors  Influencing  the  Species  Composition  of 
Mosquito  Populations  in  Indiana1 

R.  E.  Siverly,  Ball  State  University 


Abstract 

Climate,  and  natural  features  such  as  forests,  bodies  of  water,  soils,  vegetation  and 
shade  are  location-dependent  factors  which  influence  species  composition  of  mosquito 
populations  in  Indiana.  Other  factors  include:  urbanization,  suburban  development,  de- 
forestation, drainage,  and  tillage  of  soil. 

The  mosquito  fauna  of  Indiana  has  both  northern  and  southern  climatic  elements. 
Indiana  is  the  northern  range  limit  for  certain  southern  species  (e.g.  Psorophora  cyanc- 
scens,  P.  howardii),  and  the  southern  limit  for  certain  northern  species  (e.g.,  Aedcs 
abserratus,  A.  excrucians). 

Indiana's  mosquito  fauna  is  characteristically  sylvan,  plains  species  being  few  and 
limited  in  distribution.  Aedes  stimulans,  a  northern  forest  mosquito,  is  well  established 
north  of  the  Wisconsin  glacial  boundary.  South  of  this  boundary  it  is  largely  limited  to 
beech-maple  tracts  in  the  Southwestern  Till  Plain. 

About  12  of  the  50  species  of  Indiana  mosquitoes  are  produced  in  permanent  water. 
Included  are  species  of  Mansonia,  Anopheles,  Culex,  and  Uranotaenia.  Temporary  pools 
in  depressions  produce  both  univoltine  and  multivoltine  Aedes  and  Psorophora.  Recurrent 
rains  during  summer  disposes  continual  production  of  multivoltine  members  of  these  two 
genera. 

Some  species  are  directly  dependent  on  certain  plants  either  for  production  or  as 
sources  of  carbohydrate.  Indirect  effects  of  vegetation  include  shade  and  humidity. 

Soils  underlying  depressions  used  by  early  spring  mosquitoes  contained  clay  loam  and 
were  poorly  drained.  In  one  instance,  soil  disturbance  appeared  to  enhance  mosquito  pro- 
duction. The  need  for  further  studies  of  relationships  between  soil  characteristics  and 
mosquito  production  is  stressed. 

Urbanization  tends  to  substitute  one  set  of  mosquito  problems  for  another,  rather 
than  eliminate  all  problems.  Species  diversity  decreases,  with  attendant  increase  in 
density  of  adaptive  species.  Domestic  species  (e.g.,  Culex  pipiens,  the  house  mosquito)  can 
attain  dominance  as  a  result  of  improper  liquid  waste  and  solid  waste  disposal  practices. 
Aedes  triseriatus,  A.  vexans,  A.  sticticus,  A.  trivittatus  and  Psorophora  confinnis  are 
some  of  the  para-domestic  mosquitos  which  annoy  suburban  dwellers. 

Because  of  diverse  conditions  in  Indiana,  one  set  of  recommendations  for  mosquito 
abatement  will  not  suffice  for  all  localities,  and  control  recommendations  for  a  given 
community  will  require  periodic  revision. 


Introduction 

The  species  composition  of  mosquito  populations  varies  from  one 
part  of  Indiana  to  another.  Presently,  50  species  are  listed  for  Indiana. 
Twenty-seven  species  were  reported  from  Delaware  County  (8)  and 
30  species  are  listed  for  Wayne  County  (J.  H.  Hart,  personal  communi- 
cation). 

Some  species  are  limited  to  one  or  more  of  the  northernmost  tier 
of  counties.  Other  species  are  known  to  occur  only  in  the  extreme 
southern  parts  of  Indiana.  Species  composition  even  varies  between 
adjacent  counties.  Evidently,  not  one  factor   (e.g.,  climate),  but  a  com- 

1  This   investiyiration   was  supported   in    part  by   a  Riant   from   the   Indiana    State   Board 
of  Health,  funded  by  PL  89-749,  Section  314(d). 

238 


Entomology  239 

plex  of  factors  influences  these  patterns  of  distribution.  This  paper 
identifies  some  of  the  factors  responsible  for  variations  in  species 
composition  of  mosquito  populations  in  Indiana,  and  makes  some  pre- 
dictions, based  upon  these  observations,  regarding  species  composition 
of  mosquito  populations  in  future  years.  Factors  are  considered  as 
location-dependent  and  location-independent. 

Location — Dependent  Factors 
Climate 

Members  of  the  genus  Psorophora  generally  are  considered  as 
southern  mosquitoes  which  develop  rapidly  in  temporary  bodies  of  water 
following  summer  rains.  In  years  with  cool  or  dry  summers,  some  of 
the  eight  species  which  represent  this  genus  in  Indiana  may  be  absent, 
at  least  in  the  northern  part  of  the  state.  Psorophoi^a  ferox,  P.  varipes 
and  P.  horrida  were  not  found  in  Delaware  County  during  an  intensive 
survey  in  1964  (8).  These  species  were  taken  in  biting  collections  during 
the  summer  of  1969  at  one  of  the  sampling  sites  used  in  the  1964  survey. 
Thus,  climatic  variations  from  year  to  year  will  influence  the  species 
make-up  of  mosquito  populations  in  a  given  area. 

Neither  P.  cyanescens  nor  P.  howardii  has  been  reported  north  of 
Bartholomew  County.  From  state  distribution  records  (1,  2)  it  appears 
that  both  of  these  species  are  rare  or  absent  north  of  40°  latitude. 

Conversely,  some  northern  species  attain  the  southern  limit  of 
their  ranges  in  northern  Indiana.  Aedes  abserratus,  one  of  the  black- 
legged,  univoltine,  cold-hardy  mosquitoes,  had  not  been  reported  as 
occurring  south  of  LaPorte,  Lagrange,  and  Steuben  Counties.  Nationwide, 
it  has  not  been  reported  south  of  40°  latitude.  One  of  the  band-legged 
Aedes — A.  excrucians — is  found  in  kettles  and  bogs  in  northern  Indiana, 
but  it  fades  out  in  numbers,  and  its  occurrence  is  infrequent  or  rare  in 
central  Indiana.  As  in  the  case  of  A.  abserratus,  Illinois  and  Indiana 
appear  to  be  the  southern  limit  of  its  range. 

The  necessity  for  water  in  mosquito  development  is  axiomatic.  Rise 
and  decline  in  number  of  some  species  is  directly  dependent  upon  patterns 
of  precipitation  in  summer.  Production  of  Aedes  and  Psorophora  species 
from  forest  tracts  during  August  often  is  prevented — not  because  of 
insufficient  warmth — but  because  of  insufficient  rainfall.  Rainfall  of 
three  inches  or  more  within  a  24-hour  period  may  be  required  to  inundate 
soil  depressions  containing  eggs,  and  to  provide  standing  water  for  a 
week  or  more  thereafter.  With  other  species,  numbers  produced  in  a  given 
year  remains  about  the  same,  regardless  of  whether  summers  are  wet 
or  dry.  This  applies  both  in  Indiana  and  Wisconsin.  (8,  9). 

Although  other  factors  undoubtedly  play  a  role,  climate  probably  is 
the  overriding  influence  in  the  distribution  of  sub-tropical  Psorophora 
and  certain  Nearctic  Aedes  in  Indiana.  Mean  daily  minimum  tempera- 
tures and  mean  daily  maximum  temperatures  vary  10°  F  or  more  between 
northern  and  southern  Indiana  (6).  This  suggests  sufficient  gradient  for 
differences  in  species  composition  in  the  two  areas;  such  is  true  with 
mosquitoes. 


240  Indiana  Academy  of  Science 

Natural  Features 

Natural  features  which  influence  the  species  make-up  of  local  mos- 
quito populations  include  forests,  bodies  of  water,  soils,  and  vegetation. 

Forests.  In  northern,  and  in  certain  parts  of  central  Indiana,  the 
immature  stages  of  one  forest  mosquito,  Aedes  stimulans,  occur  in 
cattail  ponds  and  roadside  ditches,  as  well  as  in  woodland  pools.  With 
some  exceptions,  the  most  recent  Wisconsin  glacial  boundary  (10)  is  the 
southern  limit  of  its  range. 

A.  stimulans  was  collected  south  of  this  boundary  in  southeastern 
Indiana  in  the  spring  of  1969,  but  only  in  relict  populations  in  forested 
tracts  over  wet  soils.  On  the  other  hand,  this  species  was  absent  in  wet- 
soil  areas,  which  formerly  were  forested,  just  north  of  the  Wisconsin 
glacial  line.  In  such  habitats  it  apparently  is  displaced  with  competitive 
species  such  as  A.  canadensis  or  A.  vexans,  which  tolerate  exposure  in 
unshaded  areas.  The  distribution  of  A.  stimulans  is  discussed  further  in 
connection  with  soils  and  vegetation. 

Aedes  triseriatus  is  another  forest  mosquito,  widely  distributed  in 
Indiana.  It  is  commonly  known  as  the  treehole  mosquito  since  it  utilizes 
rot  holes  containing  water  in  stumps,  as  well  as  cavities  which  develop 
on  trunks  and  limbs  and  hold  water  as  a  result  of  natural  processes. 

Stumps  need  to  be  in  the  right  stage  of  decay  to  hold  water.  With 
extensive  decay,  the  stump  becomes  entirely  hollow  and  no  water  is  re- 
tained. This  suggests  a  time  span  for  a  stump  hole  as  a  larval  habitat. 
Over  a  10-year  period,  a  stump  hole  in  Delaware  County  yielded  Ano- 
pheles barberi  and  Cnlex  restuans  as  well  as  Aedes  triseriatus.  After  10 
years  the  cavity  rotted  through,  and  no  longer  held  water. 

Aedes  triseriatus  lately  has  come  into  prominence  because  of  its 
implication  as  a  potential  vector  of  California  encephalitis.  Small  forest 
mammals,  such  as  squirrels,  are  believed  to  act  as  a  reservoir  for  the 
causative  arbovirus.  This  vector  is  not  limited  to  forest  tracts,  however, 
since  immature  stages  also  are  found  in  artificial  containers,  such  as  old 
tires.  The  negative  association  between  California  encephalitis  and  urban 
environment  may  result  from  limited  numbers  of  reservoir  hosts  instead 
of  limited  numbers  of  suitable  vectors  in  towns  and  cities. 

Bodies  of  Water.  The  simplest  classification  of  bodies  of  water  is 
permanent  or  temporary.  Permanent  water  describes  a  site  where  water 
stands  all  year  during  most  years.  Temporary  bodies  of  water  contain 
water  for  shorter  periods. 

Contrary  to  popular  belief,  the  size  of  the  mosquito  crop  is  not 
directly  proportional  to  the  expanse  of  water.  Only  about  a  dozen  of 
Indiana  mosquitoes  can  utilize  permanent  water  as  production  sites.  Such 
species  have  adaptations  which  permit  them  to  complete  development 
in  a  habitat  which  is  alive  with  predators.  Larvae  and  pupae  of  Mansonia 
perturbans  attach  anal  siphons  to  the  subterranean  tracheal  systems  of 
such  aquatic  plants  as  cattails  and  sedges,  and  acquire  oxygen  in  this 
way.  They  never  need  to  surface  except  for  the  brief  period  required 
for  emergence  of  the  adult. 


Entomology  241 

Steuben  County  has  about  7%  of  its  area  in  wetlands,  much  of  it 
in  dry  marshes,  open  water,  shallow  and  deep  shrub  swamps,  all  in  the 
permanent  water  category  (4).  Adult  trap  collections  made  in  this 
county  in  1969  contained  relatively  high  percentages  of  Mansonia  per- 
turbans  (42%)  and  Anopheles  species  (8%). 

Larvae  of  Anopheles  mosquitoes  are  commonly  found  in  emergent 
vegetation  and  in  mats  of  floating  vegetation  where  fish  and  other 
predators  do  not  easily  gain  access.  The  resemblance  of  these  larvae 
to  floating  sticks  may  have  some  protective  value.  The  same  general 
habitat  used  by  Anopheles  is  also  used  by  Culex  species. 

Larvae  of  Culex  territans,  a  rather  ubiquitous  mosquito  which  feeds 
on  cold  blooded  animals,  also  may  be  found  along  with  Uranotaenia 
sapphirina,  in  permanent  bodies  of  water. 

Farm  ponds  seldom  serve  as  production  sites  for  mosquitoes  when 
clean  shore  lines  are  maintained,  where  there  is  little  if  any  emergent 
vegetation,  and  when  fish  and  other  predators  are  present. 

Many  kinds  of  depressions — some  man-made — hold  temporary  pools 
of  water  which  serve  as  production  sites  for  mosquitoes  after  spring 
thaw,  snow  melt  or  rains.  There  may  be  a  predator  problem  for  those 
which  develop  slowly.  For  the  most  part,  mosquitoes  which  develop  in 
pools  and  puddles  either  have  a  lower  temperature  threshold  for  develop- 
ment than  their  predators,  or  they  simply  are  geared  for  more  rapid 
development.  The  univoltine  Aedes  which  develop  during  early  spring 
while  the  water  is  40-50°F  include:  A.  abserratus,  A.  aurifer,  A.  cana- 
densis, A.  cinereus,  A.  excrucians,  A.  fitchii,  A.  flavescens,  A.  grossbecki, 
A.  stimulans,  and  A.  thibaulti.  These  are  off  the  water  and  on  the  wing 
before  crayfish,  predaceous  beetles,  dragonfly  nymphs  and  other  preda- 
tors become  established  in  the  temporary  pools  and  puddles. 

Multivoltine  Aedes  and  Psorophora  which  undergo  rapid  develop- 
ment in  temporary  water  impoundments  following  spring  and  summer 
rains  include:  A.  trivittatus,  A.  sticticus,  A.  vexans  and  all  of  the 
Psorophora.  A  week  to  10  days  is  sufficient  for  these  mosquitoes  to  com- 
plete development,  assuming  normal  summer  temperatures.  They  may 
utilize  the  same  habitats  as  the  univoltine  Aedes  which  developed  earlier 
in  the  year. 

The  importance  of  temporary  bodies  of  water  should  not  be  under- 
rated. One  woodlot  pool  in  Delaware  County  was  estimated  to  produce 
more  than  600,000  Aedes  stimulans  (8).  Flood  plains,  such  as  the  Wabash 
in  Vigo  County,  are  notorious  for  mosquito  production.  Aedes  vexans  is  a 
prime  contributor  to  this  kind  of  mosquito  problem. 

Soils.  Probably  no  one  factor  is  more  important  in  mosquito  produc- 
tion than  soil,  yet  no  factor  is  more  poorly  understood.  Soil  character- 
istics and  numbers  and  kinds  of  immature  mosquitoes  present  in  the 
water  overlaying  these  soils  were  studied  in  the  spring  of  1969.  Soil 
samples  were  taken  with  a  soil  auger  at  the  approximate  center  of  the 
depressions  where  mosquito  collections  were  made.  Soil  cores  approxi- 
mately IV2,  inches  long  and  1%  inches  in  diameter  were  transported  in 
plastic  bags  to  the  Soil  Conservation  Office  in  Muncie  for  soil  descriptions. 


242 


Indiana  Academy  of  Science 


•3  8 


^  -qj  ^ 

M    »    N 


05  5>  os  C 

.  .  .  O 

^  ^  ^  Z 

t~  ©  rH 


.2    »    «: 


"t     ^  ^     ^  "^     "t     "t 


Irt  rH 


"■*  00     O 


£  o 


P  CQ 
ft 

~c 
..   c 


— '    ° 

3  6 


c  5 


jo 
o    u 


S3 

CD      tS 


T3    +-> 


?    +* 

ft   ^ 


>,   a*    >>   5 


C     0     cJ 

O      O      *H 
U     ft   13 


I  t;    c 


,C    >—  ^3    *— 


.3    ft 

u   £ 


0)    .J3 

"S   £ 


Si  - 
2 

v  £ 

o   43  o 


£  1 
0     J 


"3   2 


S3     O 


£  b 


£   S 


I  -a  1  |  §  ||  1 

>>     .2      >>  °  - 


C     4J 


Entomology  243 

Soil  results  are  summarized  in  Table  1.  One  characteristic  shared  by 
all  soil  samples  was  poor  drainage.  With  the  exception  of  the  samples 
taken  in  Steuben  County,  all  soils  had  a  clay  component.  Amounts  of 
organic  matter  in  the  samples  were  quite  variable.  In  cases  where 
development  had  passed  the  larval  stage,  pupae  or  adults  were  collected. 
With  the  exception  of  Steuben  County  samples,  all  soil  samples  were 
taken  from  forest  soils. 

Samples  collected  in  Franklin  County  are  noteworthy.  One  soil  sample 
was  taken  in  a  water-filled  depression  where  no  mosquitoes  were  present; 
another  soil  sample  was  taken  in  a  ditch  nearby,  where  larvae  were 
present.  According  to  the  Soil  Conservation  Service,  the  ditch  sample 
revealed  more  evidence  of  disturbance  than  the  soil  taken  from  the 
depression.  Conceivably  more  minerals  and  nutrients  were  available  in 
the  ditch,  having  been  brought  to  the  soil  surface  as  a  result  of  excava- 
tion. This  same  phenomenon  was  observed  in  northern  Wisconsin  where 
greater  density  of  larvae  was  observed  in  a  borrow  pit  adjoining  a 
logging  road  than  in  an  adjacent  swamp. 

Aedes  fiavescens  is  rare  in  Indiana  (8).  Although  peat  overlaying 
marl  probably  is  not  prerequisite  for  production  of  this  mosquito,  the 
unusual  density  of  both  immatures  and  adults  near  Orland,  in  Steuben 
County,  suggests  that  A.  fiavescens  may  be  produced  over  Edwards  or 
Warner  Muck  in  other  areas. 

Less  definite  is  the  association  between  A.  stimulans  and  Brookston 
soils.  These  are  timber  soils  formed  from  fine  textured  till  from  recent 
(Wisconsin)  glaciation.  South  of  the  Wisconsin  glacial  boundary  A. 
stimulans  was  found  in  minority  numbers  in  Clark  and  Ripley  Counties. 
The  associated  soil  series  were  Avonburg  and  Clermont,  respectively. 
These  are  timber  soils  formed  on  old  (Illinoian)  glacial  till. 

From  Table  1,  one  could  generalize  that  mosquitoes  are  likely  to  be 
produced  in  depressions  in  poorly  drained  forest  soils  which  have  clay 
loam  components.  Undoubtedly,  some  species  are  tolerant  of  a  wide 
variety  of  soils,  others  less  so.  More  precise  information  regarding  soil 
characteristics  of  habitats  will  have  to  be  obtained  before  predictions  can 
be  made  relative  to  species  occurrence.  This  subject  warrants  further 
investigation. 

Vegetation.  The  association  of  Wyeomyia  smithii  with  the  pitcher 
plant,  Sarracenia  purpurea,  is  an  example  of  direct  influence.  W. 
smithii  is  not  known  to  occur  in  the  absence  of  this  pitcher  plant.  W. 
smithii  was  reported  in  LaPorte  County  in  1964  (8)  and  subsequently 
collected  in  Steuben  County.  The  direct  dependence  of  Mansonia  per- 
turbans  on  plants  such  as  cattail,  water  lily  and  sedge  has  been 
mentioned. 

Plants  may  be  of  direct  benefit  to  adult  mosquitoes  as  a  source  of 
nectar  for  food.  Preliminary  observations  in  Delaware  County  indicate 
that  Aedes  stimulans  feeds  on  nectar  of  Phlox  divaricata  (wild  phlox). 
The  occurrence  of  nectar-yielding  plants  near  its  sources  of  production 
may  influence  its  distribution. 


244  Indiana  Academy  of  Science 

Aedes  stimulans  is  associated  with  both  oak-hickory  and  beech- 
maple  climax  forests  in  the  northern  part  of  the  state  but  is  limited  in 
its  occurrence  to  beech-maple  tracts  south  of  the  Wisconsin  glacial  line. 
Aedes  canadensis  and  A.  vexans  usually  are  dominant  in  oak-hickory 
forests  on  Illinoian  drift. 

Some  of  the  most  influential  effects  of  vegetation  on  species  composi- 
tion are  the  indirect  effects  of  shade  and  humidity.  Some  mosquitoes  are 
more  tolerant  of  open,  well-lighted  conditions  than  others.  A  given  species 
may  seek  shade  in  the  southern  part  of  its  range,  but  tolerate  unshaded 
conditions  in  the  northern  part  of  its  range.  This  has  been  observed  with 
Aedes  aurifer.  Larvae  were  found  under  dense  shade  along  the  Muscata- 
tuck  Flats  in  Washington  County,  but  also  in  a  peat  bog,  near  button 
bush,  in  fairly  open  water  in  LaPorte  County.  Aedes  aurifer  were  also 
collected  from  very  open,  unshaded  pools  in  a  bog  near  Tomahawk,  Wis- 
consin. 

Culiseta  melanura  and  Culex  territans  larvae  were  collected  from  the 
same  pools  in  Delaware  County.  Culex  territans  larvae  occupied  the 
open  portions  of  the  pools.  Culiseta  melanura  larvae  were  found  in  the 
darker  recesses,  close  by  hollow  rotting  bases  of  trees  or  in  densely 
shaded  root  holes  in  the  forest  floor.  However,  in  northern  Wisconsin, 
C.  melanura  larvae  were  collected  in  sphagnum-leatherleaf  bogs,  quite 
similar  in  appearance  to  habitats  utilized  by  A.  aurifer. 

During  warm  weather  the  tolerance  for  exposure  to  light  is  an 
indication  of  capacity  for  rapid  development,  since  lighted  pools  are  also 
exposed  to  the  warmth  of  the  sun's  rays.  Several  Psorophora  species  and 
some  Aedes  (especially  A.  trivittatus) ,  which  develop  in  open  shallow 
sunlit  pools,  are  able  to  complete  development  before  depressions 
become  dry  through  drainage  and  evaporation. 

Because  of  their  small  size  and  high  surface: volume  ratio,  adult 
mosquitoes  are  particularly  subject  to  desiccation.  The  association  of 
adult  mosquitoes  and  shrubbery  is  well  known.  Here  again,  the  effect  of 
vegetation  for  mosquitoes  is  an  indirect  one,  through  providing  more 
humid  conditions  for  harborage  than  would  be  encountered  in  unshaded 
areas.  In  general,  Indiana's  mosquito  fauna  reflects  its  history  of  a  pre- 
dominant forest  vegetation.  Plains  mosquitoes  are  infrequent  or  absent 
in  the  state. 

Location-Independent  Factors 

Urbanization 

In  a  strict  sense,  whether  towns  eventually  become  cities  may 
depend  upon  location,  but  cities  of  50,000  or  more  occur  in  all  regions  of 
the  state.  Regardless  of  location,  the  most  significant  feature  of  urban 
mosquito  problems  is  their  striking  similarity.  Mosquito  problems  in 
these  urban  communities  arise  mainly  from  solid  waste  and  liquid  waste. 

Litter  is  solid  waste,  including  artificial  receptacles  which  hold  water 
such  as  cans,  buckets,  and  old  tires.  Old  tires  hold  water  for  surprisingly 


Entomology  245 

long  periods,  and  are  utilized  as  production  sources  by  Culex  pipiens,  C. 
restuans,  Aedes  triseriatus,  and  other  species  of  mosquitoes.  Used  tires, 
stockpiled  in  the  open  at  junk  yards  and  service  stations,  often  serve  as 
production  foci  for  domestic  mosquitoes.  Urbanization  and  heavy  traffic 
do  not  appear  to  be  a  deterrent,  especially  for  C.  pipiens.  The  implication 
of  this  species,  the  northern  house  mosquito,  in  the  transmission  of 
St.  Louis  Encephalitis  will  be  discussed  later. 

The  yellow  fever  mosquito,  Aedes  aegypti,  was  reported  from 
Charlestown,  in  Clark  County,  in  1941  (3).  A  visit  to  Clark  County  in  the 
spring  of  1969  revealed  no  trace  of  A.  aegypti.  There  was  no  litter  along 
the  creek  where  early  collections  were  made.  Other  production  sites  were 
plentiful  in  adjacent  areas,  and  tires  were  positive  for  house  mosquito 
larvae.  Later  in  1969,  tires  at  an  open  dump  in  Posey  County  were 
checked  for  A.  aegypti,  but  none  were  collected.  Other  mosquitoes, 
chiefly  Culex  pipiens,  were  present.  The  availability  of  artificial  con- 
tainers, particularly  in  areas  within  its  former  range,  poses  a  threat  of 
recurrence  of  A.  aegypti  within  the  state. 

Not  all  artificial  water-holding  receptacles  are  litter.  Bird  baths,  eave 
troughs  and  garden  pools  also  may  serve  as  production  sources  for 
domestic  mosquitoes.  These  tend  to  be  less  important  than  litter  recep- 
tacles, and  more  residential  than  commercial. 

The  discharge  of  household  sewage  into  streams  is  a  common  prac- 
tice in  towns  and  cities  in  the  midwest.  Often,  the  water  in  these  streams 
moves  slowly  or  tends  to  stagnate.  Organic  enrichment  and  stagnation 
provide  conditions  which  are  ideal  for  production  of  Culex  pipiens. 
Heavy  house  mosquito  production,  in  proximity  to  concentrations  of  both 
human  and  bird  populations  are  the  basic  ingredients  for  epidemics  of 
St.  Louis  encephalitis.  The  isolation  of  St.  Louis  encephalitis  virus 
from  Culex  pipiens  in  Evansville  and  Boonville  in  1964  was  reported  by 
Newhouse  and  Siverly  (5). 

Water  polluted  by  human  sewage  is  not  the  only  medium  created 
by  urbanization  and  used  by  domestic  mosquitoes.  Liquid  waste  from 
meat  packing  plants  may  be  a  potent  source  of  mosquitoes,  whether  run 
into  standing  ponds,  lagoons,  or  other  bodies  of  water. 

Obviously,  the  solution  to  these  mosquito  problems  created  by 
urbanization  is  through  sanitation  programs  and  adequate  waste  disposal 
systems.  My  purpose  is  not  to  discuss  how  this  can  be  done,  but  to 
show  that  species  composition  of  mosquito  populations  does  change  with 
industrialization,  urbanization,  and  concentrations  of  human  population. 

Suburban  development,  and  the  tendency  for  families  to  live  in  or 
near  wooded  tracts  has  intensified  certain  mosquito  problems.  Such 
species  as  Aedes  vexans,  A.  trivittatus,  A.  sticticus,  Psorophora  confinnis 
and  Anopheles  punctipennis — once  regarded  as  country  mosquitoes — 
now  may  also  be  regarded  as  para-domestic,  coming  onto  lawns,  porches, 
and  even  inside  dwellings  to  obtain  blood  meals.  Residents  inadvertently 
aggravate  this  situation  by  watering  lawns  and  providing  favorable  condi- 


246  Indiana  Academy  of  Science 

tions  for  mosquito  harborage  in  their  lush,  ornamental  shrubs.  Residents 
who  are  subjected  to  this  kind  of  mosquito  annoyance  often  strongly 
assert  there  is  no  standing  water  in  their  entire  neighborhood.  Usually 
this  is  not  the  case.  On  the  other  hand,  there  is  insufficient  information  on 
flight  range  and  dispersal  habits  of  many  para-domestic  species.  Until 
such  evidence  is  obtained,  one  should  not  rule  out  the  possibility  that 
such  mosquitoes  might  have  travelled  several  miles  from  production 
sites. 


Agricultural  Practices 

Agricultural  practices,  also,  can  alter  species  composition.  Draining 
and  deforestation  are  examples. 

Decatur  County  offers  good  examples  of  the  effects  of  drainage  on 
mosquito  production.  Much  of  the  land  in  this  county  is  wet  soil,  and 
conceivably  was  well  infested  with  mosquitoes  when  it  was  in  native 
forest.  Now,  only  remnants  of  forest  remain,  mostly  as  drained  woodlots. 
In  some  cases,  the  drainage  leads  into  farm  ponds.  Farms  are  well  tiled 
and  crop  production  is  high.  The  few  depressions  found  positive  for  mos- 
quito larvae  in  these  wooded  tracts  in  1969  were  producing  Aedes  vexans 
and  A.  canadensis.  These  mosquitoes  develop  more  rapidly,  and  in  more 
temporary  type  pools  than  A.  stimulants,  which  probably  was  present,  if 
not  dominant,  in  the  native  forests  of  Decatur  County. 

Shelby  County,  also,  shows  effects  of  drainage  and  deforestation. 
Much  of  the  land  is  in  cultivation  and  few  forest  tracts  remain.  Aedes 
stimulans  was  collected  in  Meltzer  Woods  in  association  with  A.  vexans 
and  A.  canadensis,  and  as  a  single  species  in  a  woodlot  in  the  southern 
part  of  the  county.  Mosquitoes  in  the  rural  parts  of  this  county  appear 
to  have  been  largely  eliminated  through  deforestation  and  draining. 
Psorophora  and  summer  Aedes  may  be  produced  in  sites  where 
drainage  is  blocked.  Thus,  both  Decatur  and  Shelby  Counties  afford 
examples  of  alteration  of  species  composition  through  agricultural 
practices. 

Discussion 

It  is  apparent  that  species  composition  of  mosquito  populations  in 
any  given  locality  in  Indiana  is  influenced  by  a  complex  of  many 
factors.  Superimposed  upon  such  location-dependent  factors  as  climate, 
natural  features  and  soil  are  the  effects  of  urbanization  and  agricultural 
practices. 

A  characteristic  pattern  of  species  succession  in  rural  areas  is  as 
follows:  a  rather  explosive  peak  of  univoltine  Aedes  occurs  in  late  April 
or  early  May.  This  is  shortly  followed  by  a  succession  of  overlapping 
peaks  of  multivoltine  Aedes  and  Psorophora,  the  occurrences  of  these 
peaks  varying  from  year  to  year  depending  upon  patterns  of  precipita- 
tion. These  peaks  are  concurrent  with  steady  but  sustained  populations 
of  such  permanent  water  breeders  as  Mansonia  and  Anopheles. 


Entomology  247 

This  pattern  is  likely  to  change  quite  markedly  with  urbanization. 
The  univoltine  Aedes  are  drastically  reduced  or  entirely  eliminated.  Some 
of  the  multivoltine,  para-domestic  species  are  able  to  survive,  especially 
those  which  are  versatile  in  utilizing  such  sites  as  blocked  drainage  ditches 
or  artificial  containers.  There  is  likely  to  be  a  domestic  mosquito  problem 
unless  municipal  ordinances  requiring  solid  waste  and  liquid  waste  dis- 
posal keep  apace  with  the  increase  in  urban  populations. 

Thus,  a  result  of  severe  environmental  disturbance  associated  with 
construction  of  streets,  highways,  shopping  centers,  schools,  housing 
developments,  factories  and  other  trappings  of  urbanization  is  reduction 
in  the  number  of  species  and  increase  in  the  numbers  of  members  of  a 
few  species.  The  effect  is  similar  to  that  observed  in  streams.  As  pollu- 
tion increases,  diversity  of  species  decreases  until  through  selection  all 
that  remain  are  those  which  tolerate  pollution.  Representatives  of  these 
few  species  are  in  much  greater  abundance  than  previously. 

Can  any  predictions  be  made  regarding  the  picture  as  a  whole  for 
Indiana?  If  current  trends  continue  toward  movement  of  human  popula- 
tions from  rural  to  urban  areas,  and  increased  use  of  land  for  crop 
production,  one  might  predict  that,  for  mosquitoes:  1)  the  number  of 
species  will  tend  to  decrease;  2)  that  reduction  will  most  likely  occur  in 
numbers,  if  not  in  species  of  sylvan  mosquitoes  and  permanent  water 
breeders;  3)  the  numbers  of  specimens  of  domestic  mosquitoes  likely 
will  increase,  with  attendant  likelihood  of  transmission  of  diseases  com- 
monly associated  with  domestic  mosquitoes;  4)  that  practices  which 
preserve  or  maintain  the  undisturbed  environment  will  tend  to  dampen 
the  foregoing  effects. 

The  day  is  rapidly  approaching  when  communities  in  Indiana  will 
want  organized  mosquito  control.  Oddly  enough,  the  recognition  of  this 
need  is  contemporary  with  increasing  sensitivity  to  the  use  of  pesticides. 
It  should  be  recognized  that  any  method  of  control — especially  such  a 
sophisticated  method  as  radio-sterilized  male  techniques,  or  chromosomal 
incompatibility,  or  biological  control — can  only  succeed  in  a  given  locality 
if  species  composition  in  that  particular  locality  is  known  and  at  least 
partly  understood.  The  factors  mentioned,  and  undoubtedly  others,  are 
worthy  of  studies  in  depth. 

Acknowledgments 

Thanks  are  extended  to  A.  A.  Lindsey,  H.  P.  Ulrich  and  A.  L. 
Zachary  of  Purdue  University  for  information  received.  Also,  apprecia- 
tion is  expressed  to  Keith  Huffman  and  John  Hart  of  the  Soil  Conserva- 
tion Service  for  assistance  and  suggestions. 

Literature  Cited 

1.  Carpenter,    Stanley    J.    1968.    Review    of   recent   literature    on    mosquitoes    of    North 
America.  Calif.  Vector  Views  15(8)  -.11-97. 

2.  Carpenter,     Stanley    J.,     and    Walter    J.     LaCasse.     1955.     Mosquitoes    of     North 
America  (north  of  Mexico).  University  of  California,  Berkeley  and  Los  Angeles.  360  p. 


248  Indiana  Academy  of  Science 

3.  Christensen,  G.  R.,  and  F.  C.  Harmston.  1944.  A  preliminary  list  of  the  mos- 
quitoes of  Indiana.  J.  Econ.  Entomol.  37:110-111. 

4.  Hamilton,  Max.  1962.  Wetlands  of  Steuben  County.  Indiana  Department  of 
Conservation,  Indianapolis. 

5.  Newhouse,  V.  F.,  and  R.  E.  Siverly.  1966.  St.  Louis  encephalitis  virus  from  mos- 
quitoes   in    southwestern    Indiana,    1964.    J.    of    Med.    Entomol.    3(3-4)  :340-42. 

6.  Schall,  Lawrence  A.  1966.  Climate,  p.  156-170.  In  A.  A.  Lindsey  [ed.]  Natural 
Features  of  Indiana.  Indiana  Acad,  of  Sci.  Sesquicentennial  Volume,  Indianapolis. 
600  p. 

7.  Siverly,  R.  E.  1964.  Occurrence  of  Wyeomyia  smithii  in  Indiana.  Proc.  Indiana 
Acad,  of  Sci.  73:144-145. 

8.  Siverly,  R.  E.  1966.  Mosquitoes  of  Delaware  County,  Indiana.  Mosquito  News. 
26(2)  :221-229. 

9.  Siverly,  R.  E.,  and  G.  R.  DeFoliart.  1968.  Mosquito  studies  in  northern  Wisconsin 
II.  Light  trapping  studies.  Mosquito  News.  28(2)  :  162-167. 

10.  Wayne,  William  J.  1966.  Ice  and  Land,  p.  21-39.  In  A.  A.  Lindsey  [ed.]  Natural 
Features  of  Indiana.  Indiana  Acad,  of  Sci.  Sesquicentennial  Volume,  Indianapolis. 
600  p. 


A  Check  List  of  Indiana  Collembola 
John  W.  Hart,  Earlham  College 


Abstract 

Several   entomologists   have   studied   Indiana   Collembola.    Most   of   their    findings    have 
not  been  published.   This  checklist  includes  all  species  known   from  the  state  at  this  time. 

Hoosier  entomologists  have  shown  little  interest  in  the  Collembolan 
insects.  Eight  species  were  known  from  the  state  when  Wilkey  (15) 
reported  21  new  records  in  his  taxonomic  study  of  a  small  area  in  central 
western  Indiana.  Pedigo  (8,  9,  10,  11,  12)  contributed  substantially  to 
both  the  bionomic  and  taxonomic  knowledge  of  the  Collembolan  fauna. 

Springtails  are  flightless  insects  and  restricted  for  the  most  part  to 
a  life  on  or  in  damp  soil,  under  bark,  or  on  water  surfaces.  In  a  few 
instances  they  are  found  on  growing  plants.  Only  a  few  different  kinds  of 
habitats,  involving  a  limited  number  of  soil  types  and  a  small  geographi- 
cal area,  were  investigated  in  finding  the  69  species  and  forms  and  40 
genera  reported  herein.  Far  more  study  is  indicated  if  we  are  to  know 
the  true  distribution  of  this  order  in  Indiana. 

Considering  the  multitude  of  possibilities  for  introduction  of  Col- 
lembola from  other  states  (or  even  other  nations)  one  may  conclude  that 
given  satisfactory  growing  conditions,  any  species  might  be  found  in 
almost  any  place.  In  fact,  introduction  is  often  effected  in  a  package  of 
"home  soil"  making  survival  quite  probable. 

Scott  (14),  in  his  study  of  springtails  of  New  Mexico,  reported  28 
of  the  species  and  forms  noted  in  this  check  list.  Of  the  28  reported  by 
Scott  (14),  26  were  also  found  in  New  York  by  Maynard  (5).  When 
Maynard  (5)  made  his  study,  63  of  the  species  listed  in  this  paper  had 
already  been  described.  He  found  47  of  them  in  New  York.  Mills  (6) 
studied  Iowa  springtails  at  a  time  when  only  59  of  the  69  Collembola 
we  now  know  from  Indiana  were  recognized.  He  identified  49  of  them  as 
occurring  in  Iowa. 

There  is  a  close  relationship  between  some  species  of  springtails 
and  the  soil.  These  have  interesting  possibilities  as  indicator  species. 
Environmental  disturbance  by  pollution  with  wastes  or  chemicals  (includ- 
ing insecticides)  could  be  reflected  by  variations  in  densities  of  one  or 
more  species  or  by  variation  in  composition  of  Collembolan  populations. 
There  appears  to  be  much  of  both  importance  and  interest  yet  to  be 
studied. 


249 


Checklist  of  Indiana  Collembola 


Mesaphorura  clavata  (Mills 
granulata    (Mills), 


System  of  classification,  Salmon,  1964 

Order  Collembola 

Suborder  Arthropleona 

Superfamily  Hypogastruroidea 

Family  Onychiuridae 

Subfamily  Tullbergiinae 

3),  1934 

1934 


Hart,  1969 
Hart,   1969 


Subfamily  Onychiurinae 
Hymenaphorura  subtenuis    (Folsom),    1917 

Paronychiurus    ramosus     (Folsom),    1917 

Protaphorura  encarpata    (Denis),    1931 

Family  Hypogastruridae 
Xenylla    humicola     (Fabricius),    1780 

Hypogastrura  armata    (Nicolet),   1841 
lucifuga     (Packard),    1888 
matura    (Folsom),    1916 
packardi    (Folsom),    1902 

Suborder  Neoarthropleona 
Family  Brachystomellidae 
Brachystomella  stachi  Mills,   1934 

Family  Anuridae 
Aphoromma  granaria    (Nicolet),    1847 

Friesea  claviseta  Axelson,   1900 

Family  Neanuridae 
Pscudachorutes  aureo-fasciatus   (Harvey),  1898 
saxatilis   Macnamera,    1920 

Neanura    barberi    (Handschin),    1928 

Superfamily  Entomobryoidea 
Family  Tomoceridae 
Maynardia    elongata    (Maynard),    1951 

Pogonognathellua   flavescens    (Tullberg),    1871 

Family  Isotomidae 
Subfamily  Proisotominae 
Folsomia  fimetaria    (Linne),   1758 

quadrioculata  (Tullberg),  1871 

Proi80toma    minuta     (Tullberg),     1871 

Subfamily  Isotominae 
Isotomurus  palustris    (Miiller),   1776 
palustroides  Folsom,   1937 

Pscudisotoma  sensibilis    (Tullberg),   1871 

Isototniella   minor    (Schaffer),    1896 

Isotomina   constricta    (Folsom),    1937 

Isotoma    trispinata    MacGillivray,    1896 

violacea  form   caerideatra    (Guthrie),    1903 
viridis   form    catena    (Guthrie),    1903 

Heteroisotoma  andrci    (Mills),   1934 

Vertagopus    arborea    (De    Geer),    1740 
cincrca    (Nicolet),    1842 


Hart,  1969 

Wilkey,   1950 

Hart,   1969 

Pedigo,  1969b 

Wilkey,  1950 

Packard,  1888 

Mills,   1934 

Wilkey,  1950 


Pedigo,   1969b 

Hart,   1969 
Hart,  1969 

Pedigo,   1969a 
Pedigo,  1969b 

Hart,  1969 


Pedigo,   1969a 
Folsom,  1913 


Wilkey,   1950 
Hart,   1969 

Pedigo,   1969b 


Mills,  1934 

Wilkey,  1950 

Wilkey,  1950 

Hart,  1969 

Hart,  1969 

Pedigo,  1969a 

Wilkey,  1950 

Folsom,  1937 

Mills,  1969 

Wilkey,  1950 

Wilkey,  1950 


250 


Entomology 


251 


Family  Entomobryidae 
Subfamily  Entomobryinae 
Orchesella   ainsliei   Folsom,    1924 

hexfasciata    (Harvey),    1895 
zebra  Guthrie,    1903 

Parasinella  cavernarum    (Packard),   1888 

Entomobrya    assuta    Folsom,    1924 

marginata    (Tullberg),    1871 
nivalis    (Linne),   1758 

Entomobryoides    purpurascens     (Packard),    1873 

Pseudosinella    petterseni    Borner,    1901 
rolfsi    Mills,    1932 
violenta    (Folsom),    1924 

Lepidocyrtus    curvicollis    Bourlet,    1839 
cyaneus  Tullberg,   1871 
cyaneus   form    cinercus    Folsom,    1924 
lanuginosus    (Gmelin),    1788 

(near)     jmllidus  Reuter,   1890 
paradoxus    Uzel,    1890 

Salina   banksii  MacGillivray,    1894 


Subfamily  Paronellinae 


Suborder  Metaxypleona 
Family  Poduridae 


Podura    aquatica   Linne,    1758 


Suborder  Symphypleona 
Family  Sminthuridae 
Subfamily  Sminthuridinae 
Sminthurides   hyogramme    Pedigo,    1966 
lepus    Mills,     1934 
malmgreni    (Tullberg),    1876 
pseudassimilis   Stach,   1956 

Sphaeridia   pumilis    (Krausbauer) ,    1898 

Subfamily  Sminthurinae 
Tribe  Katiannini 
Sminthurnius   aureus    (Lubbock),    1862 
elegans   (Fitch),   1863 
similitortus  Maynard,  1951 


Tribe  Sminthui 


Sminthurus    mcdialis   Mills,    1934 
trilineatus    Banks,    1903 


Sphyrotheca    minnesotensis    (Guthrie),    1903 

Tribe  Bourletiellini 
Katiannina  macgillivray    (Banks),   1897 

Bourletiella    hortensis     (Fitch),    1863 
millsi    Pedigo,    1968 

Pseudobourletiella  chandleri  Pedigo,   1968 

Deuterosminthurus  yumanensis  Wray,  1967 

Subfamily  Dicyrtominae 
Ptenothrix  marmorata    (Packard),   1873 
unicolor     (Harvey),    1893 


Wilkey,  1950 
Wilkey,  1950 
Wilkey,   1950 

Packard,   1888 

Wilkey,  1950 
Wilkey,  1950 
Wilkey,   1950 

Wilkey,   1950 

Pedigo,   1969a 

Wilkey,  1950 
Wilkey,   1950 

Pedigo,   1969a 

Pedigo,  1969a 

Pedigo,   1967 

Pedigo,   1969a 

Pedigo,  1969b 
Pedigo,   1969b 


Pedigo,   1968 


Folsom,   1916 


Pedigo,  1966 
Pedigo,  1969a 
Pedigo,  1969b 
Pedigo,   1969a 

Pedigo,   1969a 


Wilkey,  1950 
Pedigo,  1969a 
Pedigo,  1969a 

Pedigo,   1969a 
Pedigo,   1966 

Pedigo,   1969a 

Wilkey,   1950 

Gould,   1945 
Pedigo,   1968 

Pedigo,  1968 

Wray,   1967 

Pedigo,   1969a 
Wilkey,  1950 


252  Indiana  Academy  of  Science 


Literature  Cited 

1.  Folsom,    J.    W.    1913.    North    American    springtails    of    the    subfamily    Tomocerinae. 
Proc.  U.  S.  Nat.  Mus.  46  :460. 

2.  — .    1916.    North    American    Collembolous    insects    of    the    subfamilies    Achoru- 


tinae,   Neanurinae,   and    Podurinae.    Proc.   U.    S.    Nat.    Mus.    50:515. 

3.  -  — .  1937.  Nearctic  Collembola  or  springtails  of  the  family  Isotomidae.  Bull. 
U.  S.  Nat.  Mus.  168:1-472. 

4.  Could,  G.  E.  1945.  Insect  pests  of  cucurbit  crops  in  Indiana.  Proc.  Indiana  Acad. 
Sci.  53:169-170. 

5.  Maynard,  E.  A.  1951.  A  Monograph  of  the  Collembola  or  Springtail  Insects  of 
New   York   State.    Comstock   Publishing    Co.,    Ithaca,    N.Y.    388    p. 

6.  Mills,  H.  B.  1934.  A  Monograph  of  the  Collembola  of  Iowa.  Collegiate  Press, 
Ames,  Iowa. 

7.  Packard,  A.  S.  1888.  The  cave  fauna  of  North  America.  Mem.  Nat.  Acad.  Sci. 
4:1-156. 

8.  Pedigo,  L.  P.  1966.  A  new  Sminthurid  from  northwestern  Indiana  with  a  rede- 
scription  of  Sminthurus  trilrneatus  Banks.  (Collembola:  Sminthuridae) .  J.  Kansas 
Entomol.  Soc.  39(1)  :90-98. 

9.    .   1967.   Selected  life  history  phenomena  of  Lcpidocyrtus  cyancus  f.  cincrcus 

Folsom    with    reference    to    grooming    and    the    role    of    the    collophore     (Collembola: 
Entomobryidae)   Entomol.  News  78  (10)  :263-267. 

10.    .     1968.    Pond    shore    Collembola:     a    redescription    of    Salina    bankaii    Mac- 


Cillivray      (Entomobryidae)      and     new     Sminthuridae.     J.      Kansas      Entomol.      Soc. 
41(4)  :548-556. 

11.  -  .    1969a.    Activity    and    local    distribution    of    surface-active    Collembola     (In- 

secta)  :     I.     Woodland    populations.    Amer.    Midland    Natur.    83 :107-118. 

12.  -  — .    1970.    Activity    and    local    distribution    of    surface-active    Collembola    (In- 
secta)  :    II.    Pond-shore    populations.    Annals    Entomol.    Soc.    Amer.    63:753-760. 

13.  Salmon,   John   T.    1964.   An   index   to   the   Collembola.    Bull.    7,   Royal    Soc.    New   Zeal. 
651  p. 

14.  Scott,   H.    G.    1960-1965.    The   Collembola   of   New   Mexico,    I-III,    V-XII   and    XIV-XV. 
Entomol.  News  71  through  76. 

15.  Wilkey,    R.    F.    1950.    Collembola    of    Tippecanoe    and    surrounding    counties.    Unpub- 
lished  B.S.    Thesis,    Purdue   University. 

16.  WRAY,      D.      L.      1967.      Some     new     North      American      Collembola.      Entomol.      News 
78(3)  :53-62. 


GEOLOGY  AND  GEOGRAPHY 

Chairman:  W.  N.  Melhorn,  Purdue  University 
Robert  Drummond,  Indiana  State  University,  was  elected  Chairman 

for  1970 


ABSTRACTS 

Faunal  Assemblage  Study  of  the  Maryland  Miocene.  C.  Tom  Statton, 
Hanover  College. — The  location  is  approximately  eight  miles  north  of  the 
mouth  of  the  Patuxent  River  on  the  shore  of  the  Chesapeake  Bay.  The 
area  is  known  as  the  coastal  plain,  and  the  study  was  that  of  the  faunal 
assemblages  of  the  Miocene  deposits.  Emergence  and  submergence  of  the 
coastal  plain  eastward  of  the  fall  line  are  recorded  by  subsequent 
advances  and  retreating  of  the  Miocene  seas.  The  three  Miocene  seas 
under  study  resulted  in  the  Calvert,  Choptank,  and  St.  Mary's  Forma- 
tions. Included  in  this  study  are  faunal  lists,  block  counts,  and  measured 
sections  of  these  sediments.  Determination  of  the  boundaries  of  the 
Miocene  deposits  are  either  shown  in  measured  section  or  described  as  to 
the  nature  of  the  boundary,  and  should  be  readily  apparent.  The  zonation 
of  the  Miocene  sediments  is  rather  arbitrary,  based  upon  interpretive 
analysis  of  the  earlier  zonation  set  forth  by  Shattuck  in  1904.  Following 
the  presentation  of  field  exploration,  some  interpretive  data  dealing  with 
ecology  and  climatology  are  presented.  This  is  an  attempt  at  conclusion 
analysis  of  data. 

Urbanization  in  Asia.  Akhtar  Husain  Siddiqi,  Indiana  State  University. 
— The  study  brings  out  the  comparative  degree  of  urbanization  among  the 
Asian  countries.  Briefly  reviewing  the  historical  background  of  urbaniza- 
tion in  Asia,  in  the  light  of  the  economic  growth  of  the  Asian  countries, 
the  current  (1950-60)  structure  of  the  urban  population  was  discussed. 
Finally,  with  the  help  of  the  component  analysis,  an  attempt  was  made 
to  measure  the  levels  of  urbanization  in  Asia. 


OTHER  PAPER  READ 

Fact   and    Fancy.    Mary    E.    Cedars,    Roosevelt   High    School,    Kokomo, 
Indiana. 


25:? 


Acritarchs  (Leiosphaeridia)  in  the 
New  Albany  Shale  of  Southern  Indiana 

Roger  F.  Boneham,  Indiana  University  at  Kokomo 

Abstract 

The  Devonian-Mississippian  New  Albany  Shale  of  southern  Indiana  is  one  of  the 
numerous  black  shale  facies  in  central  and  eastern  North  America.  The  shale  is 
divided  into  five  lithologically  distinct  members.  These  members  contain  varying  per- 
centages of  two  acritarchs  Leiosphaeridia  plicata  Felix  and  L.  linebacki  sp.  n.  The  mean 
percentages  of  these  two  species  are  sufficiently  different  in  the  uppermost  Clegg  Creek 
and  Camp  Run  Members  to  separate  them  from  the  lower  members.  Leiosphaeridia  may 
be   a   useful   genus   for   stratigraphic   separation   of   other   black   shales. 

This  study  was  made  to  determine  whether  the  green  alga  Tasmanites 
and  the  acritarch  Leiosphaeridia  (of  unknown  biological  affinity)  might 
be  used  to  differentiate  the  various  members  of  the  New  Albany  Shale  in 
Indiana. 

The  New  Albany  Shale  was  originally  named  by  Borden  (2)  from 
exposures  along  the  Ohio  River  in  Floyd  County.  The  lithology  is  charac- 
teristic of  the  Devonian-Mississippian  dark  shales  which  outcrop  in  many 
parts  of  the  eastern  United  States  and  Ontario. 

The  New  Albany  Shale  contains  intermittent  layers  of  dolomite  and 
occasional  silty  layers  high  in  quartz.  The  formation  contains  pyrite 
scattered  throughout  its  layers  although  the  pyrite,  for  the  most  part,  is 
minute  crystals  seldom  being  larger  than  a  few  millimeters.  The  shale  has 
two  aspects:  a  brownish-black  to  black  carbon-rich  facies  and  a  greenish- 
gray  to  gray  carbon-poor  facies.  In  the  area  of  southern  Indiana 
from  which  the  samples  of  this  report  came,  the  brownish-black  to  black 
carbon-rich  facies  is  the  predominant  one. 

The  New  Albany  Shale  is  continuous  over  much  of  north-central 
United  States.  Its  equivalents  in  Ohio  are  the  Huron,  Ohio,  Cleveland, 
Bedford  and  Sunbury  shales  and  in  Michigan  the  Antrim  and  Ellsworth 
shales  (4). 

Campbell  (3)  divided  the  New  Albany  Shale  into  a  number  of  forma- 
tions and  members.  A  recent  study  by  Lineback  (15)  has  subdivided  the 
New  Albany  Shale  into  five  members.  At  the  top  of  the  uppermost  mem- 
ber (Clegg  Creek),  Lineback  has  four  beds  which  are  only  recognizable 
locally. 

The  New  Albany  Shale  contains  Tasmanites  and  Leiosphaeridia 
specimens  scattered  throughout  its  various  members  (3,  15).  The  genus 
Tasmanites  has  been  enigmatic  for  many  years  being  classified  as  worm 
eggs,  trilobite  eggs,  land  plant  spores,  hystrichospheres,  algal  spores,  and 
green  algae.  I  believe  that  Wall  (19)  has  offered  convincing  arguments 
for  placing  Tasmanites  in  the  class  Chlorophyceae  as  single  cell  algae, 
The  same  diverse  origins  have  been  proposed  for  Leiosphaeridia  as  for 
Tasmanites.  Indeed,  Leiosphaeridia  was  not  recognized  as  a  distinct  genus 

254 


Geology  and  Geography 


255 


New  Providence  Shale 

Lower  Mississippian 

Rockford  Limestone 

Clegg  Creek  Member 

Upper  Devonian 

Camp  Run  Member 

OS 

m 

Morgan  Trail  Member 

< 

Selmier  Member 

Blocher  Member 

£ 

Middle  Devonian 

North  Vernon  Limestone 

Figure    1.     Column    showing    stratigraphic    position    of    the    Neiv    Albany    Shale    and    its 
members  in  southern  Indiana. 


until  fairly  recently  (8).  Eisenack  (8)  placed  Leiosphaeridia  with  the 
hystrichospheres  while  others  (6,  7,  10,  16)  have  placed  the  genus  in  the 
group  Acritarcha  (a  group  of  unknown  and  possible  diverse  biological 
affinities). 

The  Devonian-Mississippian  boundary  is  apparently  present  near 
the  top  of  the  New  Albany  Shale.  Huddle  (12)  thought  the  uppermost 
strata  might  be  Mississippian  from  his  study  of  the  conodont  faunas. 
Read  and  Campbell  (17)  placed  the  whole  formation  within  the 
Devonian.  However,  Campbell  later  (3)  decided  that  the  uppermost  strata 
contained  a  fauna  of  Kinderhook  age.  Cross  and  Hoskins  (5)  also  placed 
the  Devonian-Mississippian  boundary  at  the  uppermost  strata  of  the  New 
Albany  Shale. 

Excellent  discussions  on  the  earlier  reports  of  Tasmanites  (20)  and 
Leiosphaeridia  (6)  are  readily  available  and  need  not  be  repeated  here. 
There  are  two  published  studies  of  attempts  to  use  Tasmanites  for  corre- 
lation of  rock  units.  The  first  study  (13)  was  done  on  subsurface  samples 
from  the  Williston  Basin  of  southwestern  Manitoba.  I  did  the  second  study 
(1).  My  attempt  was  unsuccessful  mainly  because  the  correlation  was 
attempted  over  too  large  an  area  using  too  many  formations. 

This  paper  is  the  first  attempt  to  use  Leiosphaeridia,  as  a  strati- 
graphic  marker.  The  New  Albany  Shale  contains  both  Tasmayiites  and 
Leiosphaeridia  but  the  number  of  Leiosphaeridia  specimens  is  far  greater 
than  the  Tasmaiiites  specimens.  In  this  study  I  found  that  the  most  sig- 
nificant results  were  obtained  by  concentrating  on  the  two  most  common 
species,  Leiosphaeridia  plicata  and  L.  linebacki,  and  excluding  all  other 
specimens.  Since  the  other  specimens  in  all  cases  totaled  less  than  2%  of 


256 


Indiana  Academy  of  Science 


the  entire  count,  I  do  not  believe  their  inclusion  in  this  paper  would  have 
altered  my  final  conclusions. 

In  this  study,  I  used  only  one  formation  in  a  relatively  small  area 
(Fig.  2).  I  was  fortunate  to  have  the  outcrop  localities  of  Lineback  who 
has  spent  much  time  in  the  field  studying  and  mapping  the  New  Albany 
Shale  (14,  15). 


I  oVernon 

2 


JEFFERSON 


k       6       Han 


?     CLARK 

Henry-  ° 

ville  £ 


9  oSellersburg    <J 


FLOYD 

New 

Albany  y    £ 

II  ^L    ? 

•Jefferson- 

Ville       N 


J L 


25 


MILES 


Figure  2.     Map  of  southern  Indiana  showing  collection  localities. 


Geology  and  Geography  257 

Section  Localities 

Section  1  (Lineback,  (14)  sec.  23).  Stream  bank,  Six-Mile  Creek,  T6N, 
R7E,  sec.  11,  NW  *4,  Jennings  Co.  Morgan  Trail  Mbr.  (top)  3  m  brownish- 
black,  fissile  shale.  Selmier  Mbr.  2  m  olive-gray  to  greenish-gray,  blocky 
shale  containing  dolomitic  septarian  concretions  up  to  1  m  in  diameter. 

Section  2  (Lineback  (14)  sec.  18).  Road  cut,  State  Highway  3,  1  mile 
south  of  Vernon,  T6N,  R8E,  sec.  11,  SE1^,  NW1/*,  Jennings  Co.  Blocher 
Mbr.  2  m  brownish-black,  fissile  shale  interbedded  with  thin,  grayish 
dolomitic  layers. 

Section  3  (Lineback  (14)  sec.  19).  Road  cut,  State  Highway  3,  iy2 
miles  south  of  Vernon,  T6N,  R8E,  sec.  14,  NE  % ,  NW  %,  Jennings  Co. 
Selmier  Mbr.  2  m  greenish-gray,  blocky  shale. 

Section  4  (Lineback  (14)  sec.  20).  Stream  bank  and  road  cut  along 
Quick  Creek,  T4N,  R7E,  sec.  15,  NE  hi,  NE  %,  Scott  Co.  Selmier  Mbr. 
(top)  1  m  dark  greenish-gray,  blocky  shale.  Blocher  Mbr.  2  m  brownish- 
black,  fissile  shale. 

Section  5  (Lineback  (14)  sec.  1).  Along  State  Highway  56  beginning 
at  bridge  in  T3N,  R8E,  sec.  9,  SW  XA,  SW  V±,  thence  along  north  line  sec. 
16,  NW  *4,  Jefferson  and  Scott  Cos.  Camp  Run  Mbr.  3  m  brownish-black 
to  grayish-black,  fissile  to  blocky  shale. 

Section  6  (Lineback  (14)  sec.  11).  Standard  Materials  Quarry,  4  miles 
west  of  Hanover,  T3N,  R9E,  sec.  16,  SW  &,  NE  %,  Jefferson  Co.  Morgan 
Trail  Mbr.  (top)  1  m  brownish-black,  fissile  shale.  Selmier  Mbr.  1  m 
greenish-gray,  blocky  shale.  Blocher  Mbr.  1  m  brownish-black  to  greenish- 
gray,  fissile  shale. 

Section  7  (Lineback  (14)  sec.  9).  Road  cut  State  Highway  203,  1  mile 
northwest  of  Lexington,  T3N,  R8E,  sec.  33,  NW  %,  NE  %,  Scott  Co. 
Morgan  Trail  Mbr.  2  m  brownish-black,  thinly  bedded  shale. 

Section  8  (Lineback  (14)  sec.  7).  Road  cut  State  Highway  160, 
3  miles  southeast  of  Henryville,  Clark  Grant,  lot  223,  N  %,  Clark  Co. 
Clegg  Creek  Mbr.  5  m  brownish-black,  fissile  shale. 

Section  9  (Lineback  (14)  sec.  3).  Road  cut  State  Highway  31-W,  just 
west  of  Sellersburg,  Clark  Grant,  lot  110,  center  southwest  line,  Clark  Co. 
Clegg  Creek  Mbr.  6  m  brownish-black  to  greenish-gray,  fissile  shale. 
Lineback  (14)  considers  the  lower  4  m  to  be  the  Camp  Run  Mbr.  but  the 
Leiosphaeridla  linebacki/L.  plicata  ratio  indicates  the  lower  part  is  Clegg 
Creek  (Table  1). 

Section  10  (Lineback  (14)  sec.  13).  Slate  Run  just  west  of  Blackiston 
Mill  at  north  edge  of  New  Albany,  Clark  Grant,  lot  63,  Floyd  Co.  Camp 
Run  Mbr.  SV2  m  brownish-black  to  olive-gray,  fissile  shale. 

Section  11  (Lineback  (14)  sec.  14).  Falling  Run,  just  below  bridge  to 
Silver  Hills,  west  side  of  New  Albany,  T3S,  R6E,  Sec.  3,  NW  %,  SE  %, 
Floyd  Co.  New  Providence  Shale  (top)  1  m  greenish-gray,  blocky  shale. 


258  Indiana  Academy  of  Science 


Chapel  Shale  io   m  greenish-gray,  glauconitic  shale.  Clegg  Creek,  Mbr. 
1  m  brownish-black  to  brownish-gray,  fissile  shale. 

Section  12  (Lineback  (14)  sec.  15),  Atkins  Quarry,  north  of  Jefferson- 
ville,  Clark  Grant,  lot  10,  center  southwest  line,  Clark  Co.  Morgan  Trail 
Mbr.  (top)  1  m  brownish-gray,  soft  shale.  Selmier  Mbr.  1/20  m  greenish- 
gray,  blocky  shale.  Blocher  Mbr.  3  m  brownish-black  to  brownish-gray, 
soft  to  fissile  shale. 

Section  13  (Lineback  (14)  sec.  8).  Road  cut,  State  Highway  160, 
V2  mile  east  of  Henryville,  Clark  Grant,  lot  255,  West  y2,  Clark  Co.  Rock- 
ford  Ls.  (top)  8/10  m  light  brownish-gray,  fossiliferous  limestone.  Jacobs 
Chapel  Bed  2/10  m  greenish-gray,  calcareous  shale.  Clegg  Creek  Mbr. 
3  m  brownish-black,  fissile  shale. 

Field  and  Laboratory  Methods 

Each  outcrop  was  sampled  vertically  at  1  m  intervals.  The  sample 
spacing  was  closer  if  a  particular  shale  member  was  less  than  1  m  thick. 

The  samples  were  crushed  in  the  laboratory  with  a  rotary  crusher. 
It  was  necessary  not  to  crush  the  material  too  finely  since  the  leiospheres 
are  rather  large  and  might  be  damaged.  A  working  sample  of  approxi- 
mately 10  g  was  obtained  by  repeatedly  quartering  the  crushed  field 
sample. 

This  laboratory  sample  was  boiled  in  concentrated  hydrofluoric  acid 
(52%)  for  approximately  30  minutes.  The  residue  was  centrifuged  and 
washed.  Then  it  was  placed  in  a  2%  solution  of  sodium  hypochlorite  over- 
night. The  residue  was  again  centrifuged  and  washed,  then  stained  in  a 
1%  aqueous  solution  of  methyl  green.  The  Tasmanites  and  leiospheres 
usually  did  not  take  the  stain.  However,  the  amorphous,  organic  matter 
which  is  universally  present  in  the  shales  did  take  the  stain.  The  color 
contrast  between  the  green-stained  amorphous  material  and  the  light 
yellow  to  dark  orange  color  of  the  Tasmanites  and  leiospheres  made  it 
much  easier  to  conduct  specimen  counts  under  the  microscope. 

The  stained  material  was  placed  in  glycerine  jelly  and  from  there 
mounted  on  glass  slides.  A  minimum  of  200  grains  were  counted  for 
each  sample  which  contained  fossils.  The  results  of  these  counts  are 
summarized  in  Table  1. 

Stratigraphic  Separations 

Table  1  illustrates  the  fact  that  Leiosphaeridia  linebachi  and  L. 
plicata  are  present  in  differing  quantities  in  the  various  members  of  the 
New  Albany  Shale.  Note  that  at  any  outcrop  the  L.  linebacki/L.  plicata 
ratio  may  vary,  in  some  cases,  over  rather  wide  limits.  However,  when 
the  specimen  counts  of  all  the  exposures  of  a  given  member  are  averaged 
we  arrive  at  a  meaningful  conclusion.  Namely,  that  the  Blocher,  Selmier 
and  Morgan  Trail  Members  contain  similar  mean  values  of  L.  linebachi 
and  L.  plicata  and  it  follows  that  the  L.  linebacki/L.  plicata  ratios  fall 


Geology  and  Geography 


259 


o  ^ 


«    S 


COc^OC^iHtHCOi-I'-ICO 


tji    N    M    W    O    M    O 
>    t-    H    «    «     O) 


co   eo  10   th   o   co  to 
in   n  oi   •*   ^   «   N 


«  oo  n   t- 

M    H    M    !C 


C~     CO     O     ,H     T}l 

oj   co   c-  c<i   co 


1  § 
§  .1 


,-H     r-t     cq     ,_|      ,-H 


<J5     ^f     00     O     00 
OS     O     -<*     t-     <£) 


CO    <-!    i-l    «5    O    us    O 


t    t-    -tf    <M 


oo   oo   a   oi   ffl   oo   oo 


WOfflOOOltCttlQ 


C<1    O    lO    K5    lO 


q  a 


NWiftiO<ON»N 


ft  ft       ^ 

O    H    N    M     o    W    CO 


ft 
H 


o  u 


ft 


ft  ft 

O    -I    <M     O 


O    <N    CO 


260  Indiana  Academy  of  Science 

within  similar  ranges.  The  Camp  Run  and  Clegg  Creek  Members  can 
easily  be  separated  from  each  other  and  from  the  underlying  members 
either  by  their  difference  in  means  or  the  L.  linebacki/L.  plicata  ratios. 

Conclusions 

The  significance  of  the  differences  in  the  means  and  L.  linebacki/L. 
plicata  ratios  of  some  members  of  the  New  Albany  Shale  is  that  they  are 
most  likely  a  reflection  of  a  portion  of  the  changing  biota  of  the  floating 
algal  mat  which  covered  areas  of  the  sea  during  the  deposition  of  the 
New  Albany  Shale.  Some  species  of  Leiosphaeridia  do  have  at  least 
limited  stratigraphic  value  within  the  New  Albany  Shale  and  similar 
studies  of  other  black  shales  may  prove  the  value  of  these  fossils  in  help- 
ing stratigraphers  with  some  of  their  problems  in  correlating  various 
black  shale  exposures. 

Systematic  Descriptions 

Group  Acritarcha  Evitt  1963 

Subgroup  Sphaeromorphitae 

Downie,  Evitt,  and  Sarjeant  1963 

Genus  Leiosphaeridia 

Eisenack  1958 

Leiosphaeridia  plicata  Felix  1965 
Figure  3,  Illustrations  4  and  5 

Diagnosis:  Spherical  to  subspherical.  Diameter  100-(110-225)-245  fi 
(50  specimens  measured)  96%  of  measured  specimens  are  in  the  size 
range  110-225  fi.  Wall  thickness  3-7  fi.  Surface  laevigate  with  no  pylome 
or  pores.  Wall  often,  but  not  always,  folded.  Lunate  folds  common  on 
thinner  walled  specimens.  Straight  or  slightly  sinuous  folds  common  on 
thicker  walled  specimens. 

Remarks:  Felix  (11)  found  L.  plicata  in  Neogene  age  sediments  of 
southern  Louisiana.  This  would  indicate  an  extremely  long  time  span  for 
the  species.  There  is  a  possibility  that  Felix  (11)  was  working  with 
reworked  specimens.  Another  possibility  is  that  the  New  Albany  L. 
plicata  are  not  the  same  species  as  the  Louisiana  L.  plicata  but  resemble 
each  other  due  to  convergence.  Speciation  within  the  leiospheres  is  difficult 
since  they  have  so  few  diagnostic  characters. 

Felix  did  note  that  of  the  hundreds  of  specimens  he  observed  only 
one  was  larger  than  200  /*.  It  was  245  fi.  Felix  gives  a  size  range  of 
120-200  fi  for  the  Louisiana  specimens.  The  Indiana  material  extends 
this  range  slightly.  However,  I  do  not  feel  the  great  age  range  or  slightly 
different  size  range  is  sufficient  to  believe  the  Indiana  specimens  are  a 
different  species  than  L.  plicata.  Leiosphaeridia  plicata  is  similar  in 
many  respects  to  Tasmanites  plicatilis  Boneham  1967  which  I  have  found 
in  the  Upper  Devonian  black  shales  of  Michigan,  Ohio,  and  Ontario  (1). 
The  size  range  and  lunate-shaped  wall  folds  are  similar.   However,   T. 


Geology  and  Geography 


2(\\ 


plicatilis  has  wall  pores,  the  diagnostic  feature  which  separates  the  two 
genera  Tas?nanites  and  Leiosphaeridia. 

Leiosphaeridia  linebacki  sp.  n. 
Figure  3,  Illustrations  1-3 

Diagnosis:  Spherical  to  subspherical.  Diameter  35-(40-90)-95  /* 
(50  specimens  measured)  96%  of  measured  specimens  are  within  the 
size  range  40-90  t*.  Wall  thickness  ca.  1  /*.  Surface  laevigate  with  no 
pylome  or  pores.  Wall  often  but  not  always,  folded.  Folds  straight  or 
slightly  sinuous. 

Remarks:  Most  of  the  specimens  of  L.  linebacki  are  easily  distin- 
guished from  L.  plicata  by  their  size  difference.  However,  there  is  some 
suggestion  of  an  intergradational  population  since  the  largest  specimens 
of  L.  linebacki  outwardly  resemble  the  smallest  specimens  of  L.  plicata 
if  the  latter  have  sinuous  rather  than  lunate  folds.  However,  there  is  a 
distinct  size  difference  between  the  two  species.  L.  linebacki  is  distin- 
guished from  L.  ralla  Felix  mainly  by  a  size  difference.  L.  valla,  has  a 
diameter  of  87-100  (i  and  a  wall  thickness  of  1-3  fi.  L.  linebacki  is  dis- 
tinguished from  L.  tenuissima  Eisenack  by  the  diameter,  the  latter  has 
a  diameter  of  ca.  100  p.  Eisenack  (9)  shows  figures  of  L.  tenuissima 
that  are  somewhat  larger  than  100  /u.  Also  L.  tenuissima  has  lunate  folds 


FIGURE  3.  1)  Leiosphaeridia  linebacki,  holotype,  x900,  IU  12129G40/2;  2)  L.  linebacki, 
x900,  IU  12129J26/0;  3)  L.  linebacki,  x200,  IU  12129J26/0;  4)  L.  plicata,  x 
200,  IU  12130J26/3;  5)  L.  plicata,  x200,  IU  12131Y29/3.  (All  slides  are  deposited  toith 
the  Department  of  Geology,  Indiana  University,  Bloomington.  Individual  figures  are 
located  on  a  given  slide  using  an  England  Finder  Slide.) 


262  Indiana  Academy  of  Science 

which  is  not  the  case  with  L.  linebacki.  The  size  of  L.  linebacki  is 
similar  to  Tasmanites  decorus  Boneham  from  the  Devonian  black  shales 
of  Michigan,  Ohio  and  Ontario.  However,  T.  decorus  has  wall  pores. 
L.  linebacki  bears  a  close  resemblance  to  Leiosphaeridia  (Protoleios- 
phaeridium)  major  (Staplin)  Downie  and  Sarjeant  1963.  L.  major 
(Staplin)  (18)  has  a  more  restricted  size  range  (55-85  /jl)  and  is  rela- 
tively thick  walled.  These  two  differences  lead  me  to  believe  that 
L.  major  and  L.  linebacki  are  two  distinct  species. 


Literature  Cited 

1.  Boneham,  R.  F.  1967.  Devonian  Tasmanites  from  Michigan,  Ontario,  and  northern 
Ohio.  Papers  Michigan  Acad.  Sci.  52:163-173. 

2.  Borden,  W.  W.,  1874.  Report  of  a  geological  survey  of  Clark  and  Floyd  Counties. 
Indiana  Dept.   Geol.  and  Natur.   Resources  Ann.   Rept.   5:133-189. 

3.  Campbell,  Guy.   1946.   New  Albany  Shale.   Geol.   Soc.   Amer.  Bull.   57:829-903. 

4.  Collinson,  Charles.  1968.  Devonian  of  the  north-central  region,  United  States. 
p.    933-971.    In    International    Symposium    on    the    Devonian    System. 

5.  CROSS,  A.  T.  and  J.  H.  Hoskins.  1951.  The  Devonian-Mississippian  transition  flora 
of  east-central  United  States.  C.R.  3  erne  Congres  Strat.  et  Geol.  du  Carbon., 
Heerlen:  113-122. 

6.  Downie,  Charles  and  W.  A.  S.  Sarjeant.  1963.  On  the  interpretation  and  status 
of  some  hystrichosphere  genera.  Palaeontol.  6(1)  :83-96. 

7.    .    1964.    Bibliography    and    index    of    fossil    dino-flagellates    and    acritarchs. 

Geol.  Soc.  Amer.  Mem.  94:180  p. 

8.  Eisenack,  Alfred.  1958a.  Tasmanites  Newton  und  Leiosphaeridia  n.g.  als  Gattungen 
der    Hystrichosphaeridea.    Palaeontographica.     (A)     110:1-19. 

9.    .    1958b.    Mikrofossilien   aus    dem    Ordovizium    des    Baltikums.    Senckenberg. 

leth.  39  :389-405. 

10.  Evitt,  W.  R.  1963.  a  discussion  and  proposals  concerning  fossil  dinoflagellates, 
hystrichospheres,    and    acritarchs,    II.    Proc.    Nat.    Acad.    Sci.    49 :298-302. 

11.  Felix,  C.  J.  1965.  Neogene  Tasmanites  and  leiospheres  from  southern  Louisiana, 
U.S.A.  Palaeontol.  8(1)  :  16-26. 

12.  Huddle,  J.  W.  1934.  Conodonts  from  the  New  Albany  Shale  of  Indiana:  Bull. 
Amer.  Paleontol.  21  :  136  p. 

13.  Jodry,  R.  L.  and  D.  E.  Campau.  1961.  Small  pseudochitinous  and  resinous  micro- 
fossils:  new  tools  for  the  subsurface  geologist.  Amer.  Assn.  Petrol.  Geol.  Bull. 
45(8)  :1378-1391. 

14.  Lineback,  J.  A.  1964.  Stratigraphy  and  depositional  environment  of  the  New 
Albany  Shale  (Upper  Devonian  and  Lower  Mississippian)  in  Indiana.  Unpublished 
Ph.D.    Thesis.    Indiana   University,    Bloomington.    136   p. 

15.    .    1968.    Subdivisions   and   depositional   environments   of  New   Albany    Shale 

(Devonian-Mississippian)   in  Indiana.  Amer.  Assn.  Petrol.  Geol.  Bull.  52:1291-1303. 

16.  NORRIS,  G.  and  W.  A.  S.  Sarjeant.  1965.  A  descriptive  index  of  genera  of  fossil 
Dinophyceae   and    Acritarcha.    New    Zeal.    Geol.    Surv.    Paleontol.    Bull.    40 :    72    p. 

17.  Read,  C.  B.  and  Guy  Campbell.  1939.  Preliminary  account  of  the  New  Albany  Shale 
flora.  Amer.  Midland  Natur.  21  :435-453. 

18.  Staplin,  F.  L.  1961.  Reef-controlled  distribution  of  Devonian  microplankton  in 
Alberta.  Palaeontol.  4(3)  :392-424. 

19.  Wall,  David.  1962.  Evidence  from  recent  plankton  regarding  the  biological  affini- 
ties of  Tasmanites  Newton  1875  and  Leiosphaeridia  Eisenack,  1958.  Geol.  Mag. 
94  :353-363. 

20.  Winslow,  M.  R.  1962.  Plant  spores  and  other  microfossils  from  Upper  Devonian 
and   Lower   Mississippian   rocks   of   Ohio.   U.S.    Geol.    Surv.    Prof.    Paper   364 :    93   p. 


Factors  Affecting  Coal  Roof  Rock  in  Sullivan  County,  Indiana 

Charles  E.  Wier,  Indiana  Geological  Survey 


Abstract 

Maintaining-  a  safe  and  stable  roof  is  both  an  economic  and  geologic  aspect  of  an 
underground  mine  operation.  Roof  falls  occur  when  the  rock  is  too  weak  to  support  the 
overlying  pressure  across  the  open  span  where  coal  has  been  removed.  Roof  falls  are 
related  to  characteristics  of  the  rocks.  They  may  be  described  as  dust,  lenticular,  con- 
cretion, slate,  clay  squeeze  and  massive  falls.  Lithology  and  thickness  of  beds,  jointing, 
strength  of  bedding  plane  bond,  and  the  effect  of  moisture  are  important  considera- 
tions.  No  single  criterion   seems   to  be   adequate  for   practical   roof  evaluation. 

Introduction 

Probably  no  single  aspect  of  underground  coal  mining  is  less  under- 
stood than  the  evaluation  of  roof  strength  before  actual  mining  begins. 
Certain  areas  of  good  roof  and  certain  areas  of  poor  roof  may  be  recog- 
nized but  the  roof  in  most  of  the  area  is  likely  to  be  of  questionable 
strength.  Before  mining  begins  the  geologist  can,  by  using  cores  of  the 
roof  rock,  run  physical  and  chemical  tests  that  characterize  the  roof 
rock.  During  the  mining  operation  the  engineer  can  devise  a  system  of 
mining  that  leaves  approximately  50%  of  the  coal  as  supporting  pillars 
and  a  system  of  roof  support  using  roof  bolts,  supplemented  by  posts, 
rails,  or  bars.  If  the  roof  collapses,  the  engineer  can  design  a  method 
of  cribbing  to  stabilize  the  roof  in  the  roof  fall  area.  Coal  mines,  how- 
ever, should  operate  at  a  profit  and  if  too  much  material  and  too  many 
man  hours  are  required  for  roof  control  this  cost  is  more  than  the 
margin  of  profit  and  the  mine  has  a  deficit  operation  regardless  of  the 
efficiency  of  actual  coal  removal.  The  mine  superintendent  must  see 
that  the  roof  is  supported  well  enough  that  it  will  not  endanger  the 
miners  nor  collapse  in  rooms  or  entries  and  restrict  the  flow  of  fresh  air 
through  the  passages,  or  the  flow  of  coal  out  on  conveyor  belts.  On 
the  other  hand,  his  job  is  to  keep  costs  down  and  put  the  minimum 
amount  of  money  into  roof  control. 

In  an  attempt  to  characterize  good  versus  poor  roof  condition,  in- 
formation has  been  accumulated  intermittently  during  the  past  15  years. 
Data  for  this  report  was  gathered  from  drill  holes  and  from  the  under- 
ground workings  of  the  Minnehaha,  R.  S.  and  K.,  and  Thunderbird  Mines, 
all  in  Sullivan  County. 

Kinds  of  Roof  Falls 

The  roof  in  an  underground  mine  collapses  when  the  rock  is  too  weak 
to  support  the  overlying  pressure  across  the  area  where  the  coal  has 
been  removed.  The  kinds  of  rocks  are  most  important  in  an  evaluation 
of  the  roof.  Thick  homogeneous  beds  are  more  competent  than  thin 
heterogeneous  ones.  In  general,  thick  massive  sandstone  or  limestone 
provides  the  best  roof,  and  shale  or  mixed  sand,  silt,  and  shale  beds  are 
the  poorest.  However,  a  fairly  homogeneous  gray  shale  may  be  a  satis- 

263 


264  Indiana  Academy  of  Science 

factory  roof.  Vertical  joints  across  a  bed  and  abundant  mica  and  carbon 
films  in  a  bedding  plane  allow  slippage  and  decrease  the  competence  of 
the  rocks.  Expandable  clays  in  the  rock  swell  in  the  presence  of  moisture 
and  cause  disruptive  pressures.  Water  in  the  rock  also  adds  weight  to 
the  roof,  tends  to  weaken  some  rocks,  and  lubricates  the  moving  surface. 
A  combination  of  the  above  parameters  may  produce  a  variant  of  the 
six  different  kinds  of  roof  falls  described  below. 

A  dust  roof  fall  (Fig.  1)  is  related  to  the  thin  soft  dark  gray  shale 
that  overlies  the  coal  in  some  places.  This  shale  contains  finely  dissimi- 
nated  pyrite,  calcareous  shells,  and  carbon  films.  As  soon  as  the  coal  is 
removed  and  moist  air  comes  in  contact  with  the  pyrite  in  the  shale, 
the  iron  sulfide  (pyrite)  changes  to  an  iron  sulfate  and  swells,  and  the 
shale  literally  falls  apart.  Alternate  moist  and  dry  air  that  circulates 
through  the  mine  may  hasten  the  dissociation  of  the  shale.  This  soft 
shale  crumbles  into  dust  and  falls  out  between  the  roof  bolts  even  if 
large  metal  or  woods  plates  are  used  at  the  bottom  of  the  bolt.  Not  much 
can  be  done  about  holding  this  part  of  the  roof,  but  such  falls  are  not 
a  serious  problem  because  this  shale  commonly  ranges  only  a  few  inches 
to  a  foot  in  thickness  and  the  dust  that  falls  to  the  floor  creates  only 
a  minor  nusiance. 

The  lenticular  roof  fall  is  related  to  a  sandstone  roll  (Fig.  2).  In 
this  situation,  the  bottom  surface  of  the  sandstone  is  quite  irregular 
and  the  upward  concave  areas  are  filled  with  dark  gray  to  black  shale 
that  at  that  place  forms  the  roof  of  the  coal.  This  shale  is  fairly  soft 
and  is  not  well  cemented  to  the  sandstone.  Thus  when  the  coal  is  re- 
moved, this  lenticular  unit  of  shale  will  come  down. 

Coal  beds,  such  as  the  Springfield  coal  (V),  that  are  overlain  by 
black  fissile  shale  (called  slate  by  the  miners)  that  contains  ironstone 
concretions  have  two  special  kinds  of  roof  falls — the  concretion  fall 
(Fig.  3)  and  the  "slate"  fall  (Fig.  4).  In  areas  where  the  ironstone  con- 
cretions are  developed  they  are  from  an  inch  to  four  feet  in  diameter 
and  commonly  occur  at  the  top  of  the  coal  and  the  base  of  the  overlying 
black  shale.  When  coal  is  removed  these  concretions  may  hang  down 
from  the  roof  into  the  passage  (Fig.  3).  They  are  called  pots  or  kettles 
by  the  miners  who  generally  pry  them  out  of  the  roof  before  they  fall. 

In  areas  where  the  black  slaty  shale  does  not  contain  concretions  it 
commonly  makes  a  good  roof.  If  a  break  occurs,  a  large  slab  of  rock  may 
pull  partly  loose  from  the  roof  (Fig.  4)  and  hang  there  for  days  before 
falling  unexpectedly. 

The  fifth  type  of  fall  is  related  to  weakness  through  jointing  or 
fracturing,  in  the  roof  rock  and  in  the  coal.  This  weakness  is  most 
obvious  when  it  is  also  a  clay  squeeze  (Fig.  5).  An  underclay  squeeze 
occurs  where  the  clay  beneath  the  coal  flows,  probably  by  a  combination 
of  shear  and  plastic  flowing,  from  beneath  the  coal  into  a  vertical 
crack  or  joint  in  the  coal.  In  some  cases  the  clay  moves  through  the  coal 
and  several  feet  above  the  coal  into  a  joint  in  the  roof  rock.  During 
mining  the  clay  may  flow  in  mined-out  entries  or  rooms.  Although  clays 


Geology  and  Geography 


265 


1              — 

— 

—            = 

Blnrk    Shale  ^^ 

H         r^       H 

3 

Figure  1.     Dust  roof  fall. 
Figure  2.     Lenticular  roof  fall. 
Figure  3.     Concretion  roof  fall. 

Figure  4.     Slate  roof  fall. 

Figure  5.     Clay  squeeze  and  fall. 

Figure  6.     Massive  roof  fall. 


266 


Indiana  Academy  of  Science 


that  readily  squeeze  have  somewhat  different  physical  properties  than 
those  that  do  not,  the  trigger  that  starts  the  process  is  differential 
weight  on  different  parts  of  the  coal  bed  and  thus  on  the  underlying 
clay.  Thus  if  a  clay  squeeze  occurs  across  a  mined-out  area,  commonly 
the  clay  moves  out  from  under  the  coal  on  one  side  of  the  joint  lowering 
the  coal  and  the  overlying  roof  rock  relative  to  the  mostly  undisturbed 
coal  and  roof  rock  on  the  other  side  of  the  joint.  The  roof  then  is  con- 
siderably weakened  and  the  probability  of  a  roof  fall  is  increased  tre- 
mendously. 

Fractures  or  joints  in  the  roof  rock  either  in  bedding  planes  or 
across  bedding  planes  may  be  present  and  not  easily  recognized.  It  is 
difficult  to  predict  the  probability  of  a  roof  fall  in  this  case.  The  roof 
may  look  the  same  as  in  Figure  1,  but  a  massive  roof  fall  occurs  un- 
expectedly filling  the  entry  and  extending  20  or  30  feet  upward,  mostly 
through  thin-bedded  and  interlaminated  shale,  sandstone,  and  siltstone. 
This  material  is  full  of  weak  bedding  planes  that  consist  mostly  of 
carbonaceous  films  and  mica.  When  the  coal  is  removed,  the  weight  of 
the  overlying  material  begins  to  pull  these  bedding  planes  apart;  the 
individual  beds  start  breaking  and  form  nearly  vertical  cracks.  If 
water  seeps  in  through  the  cracks,  it  weakens  the  shale  further,  acts 
as  a  lubricant  to  bedding  plane  movement,  and  adds  weight  making  the 
chances  of  roof  fall  much  greater.  This  interbedded  material  ordinarily 
does  not  contain  water  in  the  natural  state.  The  shale  beds  effectively 
seal  off  the  sand  beds  and  lenses  from  each  other  and  make  the  whole 
material  more  or  less  impermeable,  that  is,  before  cracking  takes  place. 

Two  conditions  are  favorable  for  the  accumulation  of  water:  1)  a 
thick-bedded  porous  sandstone  on  top  of  the  coal  or  close  enough  to  be 
reached  by  the  4-  to  8-foot  long  roof  bolt  holes,  and  2)  swags  or  struc- 


000  feet 


Figure  7.     Cross   section   near   the   east   side   of    Thundcrbird  Mine   showing   variations  in 
dip  of  the  coal  and  in  the  distribution  of  kinds  of  roof  rock. 


Geology  and  Geography  267 

tural  lows  in  the  coal.  These  two  conditions  are  well  illustrated  at  the 
east  end  on  Main  East  in  the  now  abandoned  Coal  VI  workings  of  the 
Thunderbird  Mine  (Fig.  7).  Where  a  porous  sandstone  rests  on  top  of 
the  coal  (and  may  cut  into  the  coal)  it  likely  will  cause  a  water  problem 
even  without  being*  in  a  structural  low.  On  the  other  hand  a  structural 
low  that  has  a  less  porous  laminated  sandstone  may  allow  the  roof  to 
be  saturated  with  water  so  that  it  becomes  quite  heavy.  Water  may 
weaken  the  bonding  ability  of  the  clay  and  increase  the  probability  of 
collapse.  Another  minor  factor  in  some  roof  falls  may  be  gas  pressure. 
At  places  where  mining  progresses  up  hill,  gas  in  the  top  of  the  coal 
and  in  the  roof  may  exert  some  lateral  pressure  on  the  roof  where  it  is 
slightly  downhill  and  where  the  coal  has  just  been  mined. 

Evaluation  of  the  Rocks 

In  an  attempt  to  try  to  predict  areas  of  massive  roof  falls  in  the 
Hymera  coal  (VI)  in  the  Thunderbird  Mine,  43  cores  were  studied  in 
detail.  The  common  sequence  for  the  20  feet  of  rocks  above  Coal  VI  is, 
from  the  top  down:  1)  thick-bedded  sandstone,  2)  laminated  sandstone, 
3)  laminated  shale,  4)  dark  gray  non-laminated  shale,  and  5)  dark 
gray  shale  containing  plant  fossils  and  pyrite. 

1)  Sandstone:  light-gray,  medium-grained,  thick-bedded;  locally  may 

be    replaced    by    siltstone,    laminated    sandstone,    or 
laminated  shale;  locally  cuts  down  into  coal. 

This  medium-grain,  thick-bedded  sandstone,  should  theoretically  be 
an  excellent  roof  where  it  is  close  to  the  coal,  because  it  is  structurally 
strong.  But  in  some  areas  it  carries  much  water  and  completely  saturates 
the  underlying  laminated  sandstone  and  shale  such  that  they  fall  from 
as  high  as  the  base  of  the  sandstone. 

2)  Sandstone:  light-    to    medium-gray,   laminated   with    10    to    50% 

shale;  contains  chlorite,  mica,  and  carbonaceous  ma- 
terial in  bedding  planes. 

3)  Shale:    dark-  to   medium-gray,  laminated  with   10   to   50%    fine- 

grained sandstone;   contains  mica  and  carbonaceous  ma- 
terial in  bedding  planes. 

The  laminated  shale  and  laminated  sandstone  differ  mostly  in 
amount  of  sandstone  versus  the  amount  of  shale.  Both  units  contain  weak 
bedding  planes  composed  of  lineated  mica  flakes  and  carbonaceous  films 
(from  fossil  plants).  The  sandstone  has  a  greater  capacity  to  contain 
water  (more  porous)  and  a  greater  capacity  to  absorb  water  (more 
permeable)  through  the  rock  if  drill  holes  connect  it  to  water-bearing 
rock.  This  water  not  only  makes  the  roof  heavier  but  tends  to  react 
electrochemically  with  some  of  the  clays  and  chlorite  (in  shale  laminae) 
in  some  bedding  planes.  The  sandstone  laminae  act  as  a  conduit  for  the 
water  but  remain  solid,  but  shale  laminae  desintegrate.  Montmorillinite 
clays  that  expand  when  wet  were  not  found  in  the  samples  tested.  Sawed 


208 


Indiana  Academy  of  Science 


samples  of  the  more  clayey  bands  of  these  two  units  readily  break  into 
small  pieces  when  immersed  in  water  for  several  hours. 

4)  Shale:    dark-gray,   not   laminated   with   standstone;    locally    con- 

tains   in   lower   part   calcite   or   pyrite    brachiopod   shells 
and  crinoid  columnals. 

This  shale,  although  not  particularly  strong,  is  homogeneous  and 
seems  to  hold  fairly  well  where  it  is  2  feet  or  more  thick.  The  strength 
of  this  shale  is  adversely  affected  by  water,  but  it  has  a  low  enough 
permeability  that  ordinarily,  water  is  not  a  problem. 

5)  Shale:    dark-gray;    contains    abundant    plant    remains,    some    of 

which  contain  pyrite  and  some  carbon   (both  vitrain  and 
fusain). 

The  shale  is  commonly  less  than  a  foot  in  thickness  and  is  absent 
in  many  areas.  Where  present  it  commonly  falls  down  on  to  the  floor  of 
the  entries  as  small  fragments  or  as  dust. 

Physical  Tests 

In  the  sequence  of  rocks  discussed  the  key  to  predicting  good  versus 
poor  roof  conditions  seems  to  be  hidden  in  the  physical  and  chemical 
variations  of  units  2  and  3,  the  laminated  sandstone  and  laminated  shale. 
Several  tests  were  tried.  A  few  look  promising. 


IOOO  2000 

Modulus   of    Rupture 


3000 


4000 


Figure  8.  Graph  showing  vertical  variation  in  modulus  of  rupture  for  samples  taken 
from  two  cores.  Solid  line  represents  core  from  Thundcrbird  Mine  area;  dashed  line 
from  R.  S.  &  K.  Mine  area. 


Geology  and  Geography  269 

Modulus  of  rupture  was  determined  for  blocks  cut  both  from  hand 
samples  and  from  cores.  In  general  roof  rock  known  to  be  strong  had 
high  shearing  strength  and  poor  roof  rock  had  low  shearing  strength. 
Where  the  first  20  feet  of  rock  immediately  above  the  coal  was  laminated 
sandstone  and  shale  the  readings  were  irregular.  Cores  were  obtained 
from  an  area  of  poor  roof  rock  in  the  Thunderbird  Mine  and  from  an 
area  of  satisfactory  roof  rock  in  the  R.  S.  &  K.  Mine  (Fig.  8).  Modulus 
of  rupture  tests  do  not  show  significant  difference. 

Clays  were  separated  from  the  shale  laminae.  Illite  is  the  most 
abundant  clay  mineral.  X-ray  patterns  indicate  that  the  001  peak  for 
illite  is  usually  symmetrical  in  the  area  of  good  roof  and  asymmetrical 
in  the  area  of  poor  roof.  The  asymmetry  indicates  a  degraded  illite  that 
will  take  water  readily  and  will  swell. 

Viscosity  tests  were  run  on  clays  by  mixing  50  g  of  clay  with  100  ml 
of  water  and  measuring  the  resistance  to  a  rotating  paddle.  The  viscosity 
of  the  clay  slurry  is  related  to  internal  friction  and  cohesion  of  particles. 
In  general  the  good  roof  rocks  had  the  highest  viscosity,  but  viscosity 
varies  vertically  over  short  distances. 

Conclusions 

In  evaluating  roof  rock  conditions  the  geologist  must  look  at  the 
rocks  from  many  viewpoints.  No  single  physical  or  chemical  parameter 
tells  the  whole  story.  Not  only  are  lateral  variations  in  the  rocks  im- 
portant but  vertical  variations  are  also.  Not  only  may  the  rock  be 
significantly  different  from  one  foot  to  the  next  but,  in  some  cases,  from 
one  centimeter  to  the  next. 


Proposed  Origins  for  the  Hadley  Lake  Depression, 
Tippecanoe  County,  Indiana 

Nils  I.  Johansen  and  Wilton  N.  Melhorn,  Purdue  University 


Abstract 

This  paper  concerns  a  study  of  an  unusual  linear  topographic  depression  in  north- 
western Tippecanoe  County,  Indiana.  This  depression  contains  Indian  Creek,  Hadley 
Lake,  a  branch  of  Burnett  Creek,  and  a  reach  of  the  Wabash  River. 

Four  hypotheses  are  presented  for  the  origin  and  development  of  the  topographic 
depression  : 

1.  A  partly  abandoned  glacial  sluiceway. 

2.  An  esker  trough  or  rinnentaler. 

3.  A  segment  of  the  "original"  Wabash  River  channel. 

4.  Stream  capture. 

After  evaluating  available  evidence,  it  is  concluded  that  hypotheses  1  and  4  may 
be  combined  and  appear  to  provide  a  reasonable  solution  to  the  problem.  Hypothesis  2 
is  an  exciting  possibility  and  hypothesis  3  is  not  totally  implausible. 

Additional  fieldwork,  particularly  shallow  exploration  geophysical  methods  to  de- 
termine bedrock  configuration  and  altitude,  is  needed  to  verify  one  of  the  hypotheses,  a 
combination  of  more  than  one  of  them,  or  provide  evidence  that  none  are  reasonable 
explanations  of  the  observed  phenomenon. 


Introduction 

The  most  striking"  topographic  feature  in  Tippecanoe  County  is  the 
Wabash  River  and  its  tributary  valleys.  The  Wabash  enters  the  county 
near  the  northeastern  corner  and  leaves  near  the  middle  of  the  western 
boundary.  All  other  streams  drain  to  the  Wabash  River  making  it  the 
local  base  level  of  erosion.  Changes  in  gradient  or  level  of  the  Wabash 
thus  will  change  the  erosive  and  transportive  power  of  streams  entering 
it. 

The  Wabash  flows,  for  the  most  part,  in  a  wide,  alluviated  valley 
bounded  by  relatively  steep,  terraced  valley  walls  (Fig.  1).  The  modern 
stream  may  be  classed  as  "underfit"  in  the  sense  of  volume  of  flow  and 
channel  width  and  configuration  compared  with  total  valley  width.  Pres- 
ent-day flow  did  not  carve  this  wide  valley;  thus,  proglacial  and  immediate 
postglacial  flow  were  much  greater  than  now.  Two  distinct  levels  of 
paired  terraces  along  the  valley  sides,  the  Mississinewa  and  Maumee 
terraces  of  Thornbury  (8)  and  an  indistinct  third  terrace  level,  whose 
place  in  the  late  glacial  history  of  the  valley  is  uncertain,  suggest  the 
importance  of  the  Wabash  valley  as  a  spillway  for  meltwater  draining 
away  from  the  retreating  Wisconsinan  glacier. 

But  did  late-glacial  Wabash  River,  in  the  Lafayette  vicinity,  always 
follow  its  present-day  course  ?  The  reason  for  posing  this  question  lies 
in  examining  the  topographic  maps  of  Tippecanoe  County  and  neighbor- 
ing areas  to  the  northeast  and  southwest.  Overall,  the  Wabash  flows  from 
northeast  to  southwest;  the  river  enters  the  county  flowing  southwest- 
ward,  gradually  changes  course  to  south,  but  at  Lafayette  makes  a 
nearly  right-angle   bend   and   flows   in   a   nearly   due   west   direction  for 

270 


Geology  and  Geography 


271 


Figure   1.     Oblique  topographic  model  of  the  Lafayette  area,    Tippecanoe  County,   Indiana 
(After  A.K.F.  Turner). 


several  miles.  It  then  resumes  a  southwestward  course  along  nearly  the 
same  vector  followed  when  entering  the  county  from  the  northeast 
(Fig.  2). 

The  near  right-angle  bend  at  Lafayette  is  also  marked  by  a  sig- 
nificant increase  in  width  of  an  already  wide  valley.  This  has  been  at- 
tributed to  the  fact  that  at  this  location  the  present  river  intersects  the 
glacially-filled  valleys  of  the  preglacial  Teays  River  and  Anderson  Valley, 
the  latter  a  tributary  of  the  ancient  Teays  which  entered  it  from  the 
east  along  the  general  line  of  present-day  Wildcat  Creek  (9).  Thus  we 
have  an  apparently  adequate  explanation  for  the  great  bend  of  the 
Wabash,  but  another  problem  arises.  It  is  generally  accepted  that  the 
present  drainage  of  the  glaciated  portion  of  Indiana  is,  except  for 
reaches  of  the  upper  middle  Wabash  drainage,  independent  of  lines  of 
preglacial  drainage.  Why,  then,  should  the  modern  Wabash  intersect 
the  buried  Teays  Valley  at  this  particular  location? 

The  key  to  the  question  may  lie  in  an  explanation  of  an  unusual 
topographic  lineament,  just  northwest  of  West  Lafayette,  where  a  small 
natural  lake,  Hadley  Lake,  and  several  other  closed,  undrained  depres- 


272 


Indiana  Academy  of  Science 


sions,  lie  athwart  the  crest  of  the  present  drainage  divide.  Two  creeks 
drain  away  from  Hadley  Lake,  but  the  lake  has  no  outlet.  These  creeks 
are  Indian  Creek  flowing:  southwestward  and  the  northeastward  flow- 
ing* Burnett  Creek.  These  creeks  also  appear  "underfit"  in  terms  of  cross- 
section  profile  versus  flow,  indicating  that  both  valleys  carried  a  greater 
flow  of  water  in  the  past. 

The    question   now   becomes,   "Is   the   apparent   alinement   of   Indian 
Creek,  Hadley  Lake,  Burnett  Creek,  and  the  Wabash   River  significant, 


Geology  and  Geography  273 

and  if  so  what  is  its  meaning  in  respect  to  development  of  the  modern 
Waabsh?"  There  may  be  several  acceptable  answers,  considering  the 
paucity  of  currently  available  data,  but  before  elaborating  the  problem 
a  brief  review  is  required  of  the  late  glacial  history  of  the  area  as  now 
interpreted. 

Late  Glacial  History 

By  late  Tertiary  time,  bedrock  subcrop  data,  especially  abundant 
from  Illinois,  indicates  regionally  mature  topography  (in  the  Davisian 
sense).  Lithologies  of  the  region  are  Paleozoic  sedimentary  rocks — sand- 
stones, shales,  and  limestones — on  which  polycyclic  episodes  of  regional 
planation  during  the  Tertiary  produced  smoothly  rounded  uplands,  broad 
valleys  of  meandering  streams,  and  local  rock  benches  or  straths  mark- 
ing the  record  of  the  polycyclic  events.  The  master  drainage  system 
was  the  Teays  River  and  its  tributaries.  The  flow  direction  of  the  Teays 
was  westward  to  join  the  preglacial  Mississippi  River  in  central  Illinois. 
Preglacial  Wabash  River  was  tributary  to  preglacial  Ohio  River,  as 
today,  but  the  Wabash  drainage  basin  was  much  smaller  than  now,  and 
separated  from  Teays  drainage  by  a  watershed  divide  located  some- 
where south  of  Lafayette,  probably  along  a  line  intersecting  the  modern 
Wabash  bedrock  gorge  somewhere  between  Attica  and  Covington.  The 
drainage  of  Tippecanoe  County  was  entirely  within  the  Teays  basin. 

Multiple  Pleistocene  glaciations  effected  many  topographic  changes 
and  drainage  diversions  in  the  northern  %  of  Indiana.  The  principal 
effects  of  ice  invasion  were  to  smooth  terrain  by  filling  of  valleys  in 
existing  topography  with  glacial  till  or  outwash.  During  interglacial 
stages,  progressive  erosion  occurred,  with  many  streams  re-establishing 
themselves  in  their  original  valleys  partly  filled  with  glacial  debris.  The 
present  topography  of  the  Wabash  bend  area,  however,  is  principally  a 
function  of  late  Wisconsinan  deposition  and  subsequent  erosion  after 
the  ice  departed  perhaps  12,000±  years  ago.  Minor  deposition  of  loess 
on  uplands  since  glacial  retreat  has  effected  small  topographic  modifica- 
tions but  these  are  irrelevant  to  the  problem  at  hand. 

Retreat  and  melting  of  ice  produced  great  volumes  of  flood  waters 
to  be  drained,  but  drainage  was  complicated  by  obstructions  of  various 
kinds — local  ice  lobation,  ice-damming  to  create  lakes,  different  erosional 
properties  of  surficial  materials,  and  existing  surface  topography.  Glacial 
drainage  lines,  or  sluiceways,  are  commonly  occupied  by  modern  streams; 
however,  these  valleys  are  too  large  for  the  present  flow  volume,  and 
are  in  this  sense  "underfit."  This  is  a  key  factor  in  attempting  to  re- 
establish drainage  systems  as  they  existed  during  retreat  and  melting 
of  the  ice.  Indeed,  underfit  streams  (in  the  sense  used  in  this  paper)  are 
common  in  Tippecanoe  and  adjacent  counties.  Essentially  all  the  larger 
streams  fit  into  this  category. 

Glaciation  also  made  broad-scale  changes  in  drainage-basin  organi- 
zation in  the  Midwest.  Teays  River  disappeared  and  the  upper  Wabash 
River  occupied  in  part,  and  by  sequential  events  not  yet  understood,  the 


274  Indiana  Academy  of  Science 

old  Teays   drainage  basin  in  Indiana.  Thus   Tippecanoe   County  now  is 
totally  incorporated  within  the  Wabash  River  drainage  system. 

In  the  northeastern  part  of  the  state,  drainage  is  toward  the  lower 
Great  Lakes,  or  more  specifically,  is  part  of  the  St.  Lawrence  River 
drainage  basin.  During  later  stages  of  Wisconsinan  glaciation,  there  is 
evidence  that  part  of  the  St.  Lawrence  drainage  was  reversed  for  some 
time.  The  eastern  glacial  Great  Lakes  used  the  upper  Wabash  River  as 
an  outlet,  and  a  significant  flow  of  water  (Maumee  flood)  was  added  to 
the  Wabash  Valley  sluiceway. 

In  summation,  the  picture  of  what  happened  to  drainage  systems 
during  ice  melting  and  shortly  after  the  glaciers  disappeared  from  the 
region  is  by  no  means  totally  clear.  Climatic  and  vegetation  conditions 
are  unknown  or  only  inferred.  These  two  factors  have  great  significance 
in  determining  type  and  rate  of  erosion  and  landscape  modification.  Ero- 
sion and/or  change  in  climate  and  vegetation  may  obscure  or  destroy  the 
evidence  of  former  location  of  a  stream,  and  we  must  accept  the  fact 
that  a  stream  is  where  it  is  for  no  good  reason  that  we  can  ascertain. 
The  stream  is  where  it  is  because  it  started  out  in  this  location  for 
reasons  not  now  apparent.  These  unknowns  probably  explain  why  many 
areas  in  Indiana  look  peculiar  to  the  geomorphologist  or  airphoto  inter- 
preter; they  can  not  be  fully  explained,  and  look  more  like  a  "geo- 
morphic  happening"  than  something  that  developed  by  rational  ordering. 
But  the  reasons  are  there.  We  have  not  yet  found  them. 

Previous  Work 

Tippecanoe  County  and  the  great  bend  of  the  Wabash  appears  to 
have  been  a  popular  area  for  geographic  field  investigations  in  the 
early  part  of  this  century.  McBeth  (5)  investigated  many  features  re- 
sulting from  glaciation,  as  well  as  development  of  the  Wabash  drainage 
system,  but  his  papers  make  no  reference  to  a  Hadley  Lake-Wabash  River 
lineament;  very  probably  it  escaped  his  notice  because  it  is  not  a  topo- 
graphically prominent  feature  as  compared  to  eskers  and  terraces,  and 
of  course  he  lacked  topographic  maps  or  air  photos  on  which  the  linea- 
ment immediately  attracts  attention.  F.  J.  Breeze  (1)  apparently  recog- 
nized the  peculiar  relations  of  Burnett  Creek  and  Indian  Creek  in  a 
paper  presented  before  the  Indiana  Academy  in  1916  but  no  abstract  or 
paper  was  published  and  thus  details  of  his  observations  or  interpreta- 
tions are  lost.  The  early  report  on  Tippecanoe  County  by  Gorby  (3) 
likewise  fails  to  mention  the  Hadley  Lake  depression.  Gorby's  report  is 
not  particularly  admirable  even  by  geologic  standards  of  his  time,  though 
his  statement  that  ".  .  .  The  assertion  may  be  ventured  that  no  county 
in  the  Union  affords  a  more  varied  exhibit  of  this  puzzling  deposit  (i.e., 
the  Glacial  drift)  than  Tippecanoe  .  .  ."  might  draw  plaudits  from 
modern  workers.  Leverett's  classic  work  (4)  also  lacks  mention  of  the 
Hadley  Lake  lake  and  trench. 

No  further  geomorphic  work  was  done  in  the  Tippecanoe  region 
until  recent  years.  Schneider  and  others  (6)  describe  three  sets  of  linear 


Geology  and  Geography  275 

features  in  western  Indiana,  which  consist  of  straight,  shallow  troughs 
and  discontinuous  low  ridges  or  elongated  knobs  of  sand  and  gravel.  They 
relate  these  linear  features  (one  of  which  is  Hadley  Lake  trough)  to  sub- 
parallel  northeast-southwest  drainage  developed  in  accord  with  ice  move- 
ment from  the  northeast.  West  and  Barr  (11)  describe  groundwater  prob- 
lems in  the  area  immediately  west  of  Hadley  Lake  and  infer  a  substantial 
thickening  of  glacial  drift  eastward  into  the  depression.  Schneider 
and  Wayne  (7)  suggest  that  the  depression  is  an  esker-trough  that  is 
a  downstream  continuation  of  that  segment  of  the  Wabash  River 
extending  from  near  Lafayette  upstream  to  Georgetown  in  Cass  County. 

We  are  now  in  a  position  to  again  ask  the  question,  "Is  the  aline- 
ment  of  Burnett  Creek,  Indian  Creek,  and  Hadley  Lake  significant,  and 
in  what  relation  to  the  modern  course  of  the  Wabash  River?"  Let  us  look 
at  four  preliminary  hypotheses  in  an  attempt  to  answer  this  question. 

Hypotheses  of  Origin 

We  suggest  four  possible  hypotheses  for  development  of  this  peculiar 
alinement  of  land  forms. 

1.  An  abandoned  glacial  sluiceway 

2.  An  esker  trough,  or  perhaps  a  rinnentaler 

3.  The  "original"  post-glacial  Wabash  river  channel 

4.  Stream  capture 

All  hypotheses  deal  with  fluvial  processes  as  an  erosional  factor. 
What  is  not  taken  into  consideration  are  bedrock  topography,  joint 
control,  faulting  (if  any),  and  general  structural  geology  of  the  area. 
As  the  buried  extension  of  the  Knobstone  cuesta  must  pass  almost  di- 
rectly beneath  the  linear  surface  depression,  bedrock  cannot  be  elimi- 
nated as  a  determinant  in  developing  a  satisfactory  explanation  for 
surface  drainage  adjustments  despite  a  rather  thick  covering  blanket  of 
glacial  till  or  outwash.  Too  few  bedrock  data  are  available,  however,  to 
assess  its  importance  at  this  time,  although  Wayne  (10)  produced  a 
preliminary  bedrock  physiography  map  of  the  area  based  on  limited 
information. 

Let  us  look  at  each  hypothesis  in  some  detail. 

Glacial  Sluiceway 

This  is  the  most  popular  local  explanation  of  the  landforms  by 
geomorphologists  and  air  photo  interpreters.  The  assumptions  behind 
this  interpretation  follow: 

1)  The  Wabash  River  has  not  changed  its  course. 

2)  The  sluiceway  received  large  volumes  of  water  from  a 
stagnant  glacier  or  glacier  receding  primarily  by  vertical 
ablation. 

3)  The  glacial  source  of  meltwater  was  nearby,  perhaps  as  close 
as  the  northern  border  of  Tippecanoe   County   because   the 


276 


Indiana  Academy  of  Science 


sluiceway  is  not  traced  farther  northeast;  if  it  did  extend 
farther,  post-glacial  erosion  or  colluviation  has  destroyed 
evidence  of  former  existence. 

4)  The  time  of  origin  is  thus  placed  as  late-glacial,  when  the 
glaciers  left  the  area  about  12,000 ±  years  ago,  or  about  the 
time  of  formation  of  the  Iroquois   (  ? )   moraine. 

5)  The  sluiceway  carried  water  for  only  a  short  time,  and  later 
creeks  occupied  the  depression  and  began  to  cut  new  valleys. 


NW 


SE 


Typical    (7) 


N.W.-S.E.    Sec. 9,    T.23N.,    R.5W. 


Near  headwater© 


N.W.-S.E.   Sec.  3,   T.  23  N.,  R.5W. 


SE 


Near  the  Wabosh 
River  @ 


N.W.-S.E.  Sec. 24,  T.23N.,   R.6W. 


Ficuke  3.    Cross-section  of  present  Indian  Creek  (See  Fig.  2  for  locations). 


Geology  and  Geography  277 

Indian  Creek  and  Burnett  Creek  thus  formed.  Hadley  Lake 
was  left  as  a  ponded  local  segment  of  the  trench  because  the 
bedrock  high  southwest  of  the  lake  prevented  free  drainage 
in  that  direction. 

The  conclusion  is  thus  reached  that  the  trench  is  essentially  a 
temporary  overflow  distributary  of  the  Wabash.  With  further  retreat 
of  the  ice,  the  source  of  water  was  gone  and  the  sluiceway  was  occupied 
by  small,  newly-formed  streams. 

Cross-sections  of  Indian  Creek  valley  (Fig.  3)  show  clearly  at  least 
two  cycles  of  erosion,  with  a  wide  valley  representing  the  sluiceway  stage, 
and  a  smaller  valley  of  the  modern  creek.  Thus  the  sluiceway  hypothesis 
appears  adequate  to  explain  the  landform  feature  of  the  trench;  how- 
ever, it  explains  neither  the  reason  for  location  or  alinement.  The 
hypothesis  fails  to  take  into  account  either  bedrock  topography  or 
correlation  between  this,  the  present  Wabash  Valley,  and  the  preglacial 
Teays  River  Valley.  The  glacial  sluiceway  interpretation  suggests  that 
the  landforms  are  independent  of  these  latter  factors  and  dependent  only 
on  an  abundant  source  of  melt-water. 

Esker  Trough  or  Rinnentaler 

This  hypothesis  implies  subglacial  or  englacial  scour  creating  a 
bedrock  trench  of  linear  section.  Leverett  used  the  term  esker  trough  for 
similar  linear  features  in  Montgomery  County,  near  Muncie,  and  else- 
where in  the  state,  limiting  the  term  to  linear  depressions  now  contain- 
ing only  relicit  patches  or  knobs  of  sand  and  gravel,  and  as  noted  previ- 
ously, Schneider  has  continued  usage  of  this  term  in  describing  three 
sets  of  linear  features   in  west-central  Indiana. 

As  no  relict  patches  remain  in  the  Hadley  Lake  trench  there  is  no 
proof  that  an  esker  was  ever  present.  The  suggestion  is  tenable  that 
the  trench  is  a  rinnentaler  (channel-valley).  These  valleys  may  be  cut 
uphill  beneath  the  ice  because  of  pressure;  terminal  base-level  of 
streams  may  raise  and  streams  flow  englacially  or  even  superglacially 
over  collapsed  ice  filling  the  rinnentaler  (2) 

Acceptance  of  this  hypothesis  requires  formation  in  a  glacial  rather 
than  proglacial  environment  as  suggested  by  the  glacial  sluiceway  theory. 
The  fact  that  the  Hadley  Lake  trench  cuts  uphill  and  across  a  bedrock 
divide  appears  to  favor  the  rinnentaler  over  the  esker  trough  hypothesis 
of  origin. 

"Original"  Post-glacial  Wabash  River  Channel 

This  is  a  bold  hypothesis  but  is  worth   consideration. 

1)  The  hypothesis  is  not  dependent  on  a  particular  source  of 
meltwater,  but  rather  a  large  flow  for  a  longer  period  of 
time,  as  could  be  expected  during  the  time  of  glacial  melt- 
ing. 

2)  The  hypothesis  assumes  a  correlation  between  alinement  of 
the  landform,  bedrock  topography,  the  Teays  valley,  and  the 
modern  Wabash  River. 


278 


Indiana  Academy  of  Science 


Let  us  project  a  sequence  of  events  which  could  produce  the  aline- 
ment  of  the  Indian  Creek-Hadley  Lake-Burnett  Creek  system  and  the 
Wabash  River  (Fig.  4). 

When  the  ice  retreated  and  the  land  surface  was  re-exposed,  pre- 
existing topography  was  buried  by  glacial  drift.  Preglacial  drainage 
channels  were  buried  and  running  water  commenced  re-dissecting  the 
new  surface.  The  ample  water  available  gave  birth  to  a  Wabash  River 


L  Wabash  encounters  obstruction. 


2.  Downcutting  ceases,  possible  ponding. 

Tributary  eroding  in  unconsolidated 
debris  partly  filling  the  buried  Teays 
channel. 


3.   Tributary  reaches  the  Wabash. 
Overflow  with  rapid  downcutting. 


4.   Reversal  of  drainage  in  the 
future  Burnett  Creek  segment. 


5.    Modern  distribution  of   streams. 


Figure  4.    Projected  sequence  of  events  creating  the  Hadley  Lake  trough. 


Geology  and  Geography  279 

flowing  in  almost  a  straight  southwestern  direction.  The  new  stream  was 
unaffected  by  buried  topography;  soon,  however,  downcutting  encountered 
the  bedrock  "high"  located  southwest  of  present  Hadley  Lake.  This 
produced  a  temporary  base  level;  the  rate  of  downcutting  could  have 
slowed  upstream  from  the  obstruction  and  ponding  possibly  took  place. 
Downstream,  Indian  Creek  continued  its  work  of  grading  this  segment 
to  the  next  lower  temporary  base  level  control  at  some  unknown  point 
further  down  the  Wabash  valley. 

In  time,  overflow  from  the  upstream  segment  may  have  occurred 
approximately  where  Tippecanoe  River  enters  the  modern  Wabash.  Over- 
flow may  have  combined  with  headward  erosion  of  tributaries  from  the 
east  and  south;  these  tributaries  in  part  were  re-occupying  an  only  partly 
filled  segment  of  the  old  Teays  Valley.  Once  diversion  occurred,  upstream 
erosion  was  accelerated,  for  the  stream  was  now  completely  cutting  in 
glacial  drift  filling  the  old  Teays  and  Anderson  valleys  and  no  longer  had 
to  contend  with  the  bedrock  high  athwart  its  previous  course.  As  the 
softer  fill  was  exposed  to  less  overall  consolidation  by  glaciation,  it 
provided  no  obstruction  to  lateral  corrasion.  In  time,  a  marked  widening 
of  the  Wabash  Valley  occurred  southward,  in  the  upstream  direction  of 
the  old  Anderson  Valley. 

The  "original  Wabash"  valley  was  thus  abandoned.  The  small 
streams  in  this  valley  continued  slow  development.  Hadley  Lake  was 
left  behind  as  a  ponded  depression  blocked  by  the  bedrock  divide.  Some 
factors  needing  verification  if  this  hypothesis  is  to  be  supported  are: 

1)  The  presence  of  lacustrine  sediments  in  the  Hadley  Lake 
basin,   provided   the   original   lake   existed   long  enough. 

2)  Geophysical  profiles  or  drill  holes  along  the  axis  of  the 
trough  to  determine  depth  to  bedrock,  gradient  of  the  postu- 
lated river,  and  evidence,  if  any,  of  former  terraces  along 
this  postulated  course. 

3)  Composition  of  soil  materials  along  the  postulated  course 
to  determine  if  they  were  deposited  by  running  water  or 
if  the  till  has  been  reworked  by  fluvial  action. 

There  can  be,  admittedly,  substantial  objections  to  this  hypothesis. 
Nevertheless,  it  deserves  consideration  until  negative  proof  is  forth- 
coming. 

Stream  Capture 

This  hypothesis  describes  a  chain  of  events  to  explain  the  linear 
depression.  Some  fundamental  assumptions  used  in  suggesting  a  simple 
case  of  stream  capture  are: 

1)  The  Wabash  River  did  not  change  its  course,  but  did  have  a 
much  greater  flow  volume. 

2)  Notation  of  the  peculiar  alinement  of  Burnett  Creek,  with 
a  northeastward,  or  opposite,  direction  of  flow. 


280  Indiana  Academy  of  Science 

3)  The  presence  of  a  bedrock  high  or  divide  southwest  of  Hadley 
Lake. 
The  sequence  of  events  may  have  been  as  follows.  Wabash  River 
and  its  tributaries  were  carrying-  a  tremendous  volume  of  meltwater, 
whereas  the  north  fork  of  Burnett  Creek  followed  the  Hadley  Lake- 
Indian  Creek  channel  southwestward.  As  flow  was  obstructed  by  the 
bedrock  high,  the  upper  reaches  of  the  system  (Burnett  Creek)  lost 
most  of  its  erosive  power.  Wabash  River  and  the  Indian  Creek  segment 
continued  downcutting.  Meanwhile  a  small  tributary  was  eroding  north- 
ward from  the  Wabash  River  in  section  22,  T.  24  N.,  R.  4  W.  This 
tributary  eventually  reached  Burnett  Creek  in  the  E  V2  section  21,  T.  24 
N.,  R.  4  W.,  and  at  this  time  a  more  favorable  outlet  to  the  Wabash 
River  was  provided.  Consequently,  Burnett  Creek  reversed  its  flow  di- 
rection and  Hadley  Lake  was  left  as  a  local  depression  behind  the 
bedrock  dam.  Indian  Creek  and  Burnett  Creek  became  two  independent 
drainage  systems,  and  a  wide  abandoned  valley,  or  col,  was  left  across 
the  bedrock  divide.  Proof  of  this  hypothesis  of  drainage  reversal  awaits 
identification  of  terrace  deposits  on  Burnett  Creek  and  determination  of 
whether  the  gradient  of  the  terraces  correspond  to  a  supposed  flow  in 
the  opposite  direction  from  that  of  the  modern  stream. 

Conclusions 

Four  hypotheses  have  been  presented  to  explain  the  origin  of  the 
Hadley  Lake  depression  and  related  fluvially  carved  landforms.  Two  of 
these  (abandoned  glacial  sluiceway  and  stream  capture)  may  be  com- 
bined and  appear  to  offer  a  reasonable  explanation.  All  hypotheses, 
however,  must  remain  in  consideration  until  sufficient  evidence  or  proof 
is  provided  to  explain  the  observed  phenomena. 

Literature  Cited 

1.  Breeze,  F.  J.  1916.  Diversion  of  drainage,  Indian  Creek  to  Burnett  Creek.  Proc. 
Indiana  Acad.  Sci.  for  1916:49. 

2.  Embleton,  C,  and  C.  A.  M.  King.  1968.  Glacial  and  Periglacial  Geomorphology. 
Edward  Arnold,  Ltd.,  London.  608  p. 

3.  Gorby,  S.  S.  1886.  Geology  of  Tippecanoe  County.  Indiana  Dept.  Geol.  and  Natur. 
Resources  15:61-69. 

4.  Leverett,  F.,  and  F.  D.  Taylor.  1915.  The  Pleistocene  of  Indiana  and  Michigan 
and  the  history  of  the  Great  Lakes.  U.  S.  Geol.  Surv.  Mongr.  53  :529  p. 

5.  McBeth,  W.  A.  1901.  Wabash  River  terraces  in  Tippecanoe  County,  Indiana.  Proc. 
Indiana  Acad.  Sci.  for  1901  :237-243. 

6.  Schneider,  A.  F.,  G.  H.  Johnson,  and  W.  J.  Wayne.  1963.  Some  linear  glacial 
features  in  west-central  Indiana.  Proc.  Indiana  Acad.  Sci.  72:172-173. 

7.  Schneider,  A.  F.,  and  W.  J.  Wayne.  1968.  Segmentation  of  the  Upper  Wabash 
River  across  Indiana.  Second  Ann.  Meeting  North-Central  Section,  Geol.  Soc. 
Amer.  Iowa  City,  Iowa.  Program  Abstr.  :36-37. 

8.  Thornbury,  W.  D.  1958.  The  geomorphic  history  of  the  upper  Wabash  Valley. 
Amer.  J.  Sci.  256  :449-469. 

9.  Wayne,  W.  J.  1952.  Pleistocene  evolution  of  the  Ohio  and  Wabash  valleys.  J.  Geol. 
60:575-585. 

10.  Wayne,  W.  J.  1956.  Thickness  of  drift  and  bedrock  physiography  north  of  the 
Wisconsin  glacial  boundary.   Indiana  Geol.   Surv.   Rep.    Prog.    7:70  p. 

11.  West,  T.  R.,  and  D.  J.  Barr.  1965.  Economic  groundwater  problems  encountered 
in  the  development  of  a  housing  area  near  West  Lafayette,  Indiana.  Proc.  Indiana 
Acad.  Sci.  74  :259-267. 


Base  Level,  Lithologic  and  Climatic  Controls  of  Karst  Groundwater 
Zones  in  South-Central  Indiana 

Richard  L.  Powell,  Indiana  Geological  Survey- 
Abstract 

Nearly  all  groundwater  movement  with  the  carbonate  bedrock  of  south-central 
Indiana  has  been  within  a  karst  groundwater  zone  that  occupies  the  lower  part  of  the 
vadose  groundwater  zone  above  the  zone  of  permanent  saturation  (phreatic  zone)  or 
local   base   level   and  that   is   characterized  by   a  highly   fluctuating   water   table. 

Variations  of  precipitation  within  climatic  cycles  and  the  local  relief  above  base 
level  control  the  vertical  range  of  the  water  table  and  subsequent  solution  within  the 
karst  groundwater  zone  during  that  time  period  corresponding  to  each  subaerial  erosion 
level.  Variations  in  carbonate  solubility,  thickness  of  beds,  and  intensity  of  jointing 
determine  the  texture  and  shape  of  the  integrated  subterranean  conduits.  The  volume  of 
water  transmitted  through  the  karst  groundwater  zone  and  its  acidity  determine  the 
size. 

Limestone  is  relatively  soluble  compared  with  dolomite  and  clastic  sediments,  but 
it  is  less  permeable  than  some  dolomites  or  sandstones.  Initial  permeability  in  limestone  is 
along  joints  and  bedding  planes  and  varies  greatly  from  bed  to  bed.  Shales  and  silty 
carbonate  units  may  form  perched  water  bodies  that  may  be  breached  by  joints  or 
erosion.  Sandstone,  dolomite,  and  intensely  jointed  limestone  may  form  perched  water 
bodies  or  aquifers  within  relatively  impermeable  limestone  strata  and  release  the  water 
to  less  permeable,  subjacent  limestone. 

Introduction 

Solutionally-enlarged  openings  and  caverns  in  dynamic  karst  ground- 
water zones  carry  the  greatest  amount  of  groundwater  in  the  non-glaci- 
ated portion  of  south-central  Indiana  (Fig.  1).  Base  level,  lithology  and 
climate  are  the  three  major  controls  of  karst  groundwater  zones  in 
carbonate  rocks  of  Mississippian  age  that  underlie  the  Mitchell  Plain 
and  Crawford  Upland  physiographic  units  of  south-central  Indiana. 

Karst  Groundwater  Zone 

The  karst  groundwater  zone  is  characterized  by  a  highly  fluctuating 
water  table  within  the  confines  of  any  integrated  openings,  such  as  joints 
and  bedding  planes,  in  carbonate  bedrock  (13)  (Fig.  2).  It  is  above 
the  zone  of  permanent  saturation  (phreatic  groundwater  zone)  and  there- 
fore includes  the  lower  part  of  the  zone  of  aeration  (vadose  ground- 
water zone).  This  zone  is  somewhat  the  same  as  the  fluctuating  water 
table  zone  described  by  numerous  hydrologists,  such  as  Finch  (4)  and 
Meinzer  (8),  and  defined  by  Swinnerton  (14)  as  the  zone  within  which 
limestone  caverns  are  developed. 

Water  table  fluctuations  within  karst  terrains  have  been  ignored 
by  most  theorists  concerned  with  formulating  a  universal  theory  of 
origin  of  limestone  caverns,  even  though  the  most  intense  and  frequent 
water  table  fluctuations  documented  have  been  within  karst  or  cavern 
areas.  A  few  modern  hydrologists  argue  that  a  true  water  table  does 
not  exist  within  the  widely  spaced  openings  in  carbonate  rocks.  The  fact 
that  the  relative  grain  size  between  the  interstices,  usually  open  joints 

281 


282 


Indiana  Academy  of  Science 


Outcrop  Area 
of  Middle  j 
Mississippian 
Limestones 


1 000 -i 


CRAWFORD  UPLAND 
Stephensport   and 


MITCHELL 
PLAIN 


600- 


200 


FIGURE  1.  Generalized  map  and  cross  section  of  south-central  Indiana  showing  the 
location  of  the  Mitchell  Plain  and  Crawford  Upland  and  their  relationship  to  rocks  of 
M iss iss ippian  ag e . 


Geology  and  Geography 


28.'] 


Figure  2.     Idealized  diagram    showing   the  range  of  fluctuations  of  a  normal  toater   table 
in  a  homogeneously  permeable  medium. 


and  bedding  planes,  is  measured  in  feet  rather  than  in  microns  does  not 
alter  the  fact  that  groundwater  level  fluctuates  within  the  integrated 
openings.  The  fluctuations  are  commonly  measurable  in  feet  and  tens 
of  feet  in  Indiana,  but  rises  of  the  water  table  up  to  a  few  hundred  feet 
are  known  elsewhere. 

A  fluctuating  water  table  exists  in  all  permeable  strata.  The  fluctua- 
tions of  the  water  table  are  caused  by  the  difference  in  frequency  and 
amounts  of  precipitation,  infiltration  rates,  transmission  rates,  storage 
capacity  of  the  bedrock,  and  discharge  rates  from  the  permeable  strata 
or  bedrock.  Precipitation  is  variable  seasonally,  annually,  or  within  cli- 
matic cycles  and  geologic  episodes,  causing  corresponding  fluctuations 
of  the  water  table.  The  rate  of  infiltration  is  dependant  upon  the  presence 
of  exposed  surface  openings  and  is  affected  greatly  by  the  type  and 
density  of  vegetation.  There  may  be  no  surface  run-off  from  a  forested 
area  while  from  a  denuded  area  it  may  be  total.  The  transmission  of 
karst  groundwater  is  dependent  upon  the  size,  shape  and  sinuosity  of 
integrated  openings  between  an  intake  area  and  discharge  point.  The 
larger  the  openings,  the  greater  the  potential  groundwater  movement. 
The  amount  of  groundwater  movement  and  discharge  is  controlled  by 
size  and  abundance  of  integrated  openings,  and  by  storage  capacity  within 
the  fluctuating  zone.  The  rate  of  groundwater  discharge  to  surface  out- 
lets is  controlled  by  the  size  of  the  integrated  openings  and  the  hydro- 
static head  that  may  be  developed  within  the  subterranean  tributaries  to 
the  outlets. 

The  greatest  volume  of  groundwater  movement  within  carbonate 
bedrock  of  a  karst  terrain  occurs  within  the  zone  that  is  most  affected 
by  infiltration  of  acidic  meteoric  waters  and  discharge  of  the  accumulated 
karst  groundwaters.  The  shallow  zone  just  above  the  zone  of  permanent 
saturation  within  the  normal  range  of  water  table  fluctuation  that  is 
affected    by    even    slight    infiltration    transmits    the    greatest    volume    of 


284  Indiana  Academy  of  Science 

fresh  karst  groundwater  (Fig.  2).  Therefore,  it  is  the  zone  of  the 
greatest  amount  of  solution.  The  amount  of  karst  groundwater  trans- 
mitted above  or  below  the  normal  range  of  the  water  table  decreases 
proportionally  to  distance  from  the  normal  zone.  A  decrease  in  infiltra- 
tion results  in  a  decrease  in  the  amount  of  acidic  karst  groundwater, 
therefore,  a  significant  decrease  in  solutional  enlargement  and  a  lower- 
ing of  the  water  table.  An  increase  of  precipitation  causing  a  rise  of 
the  water  table  above  the  zone  of  normal  fluctuations  would  increase  the 
hydrostatic  head,  rate  of  flow  and  transmitted  acidic  karst  groundwater 
in  pre-existing  openings  developed  within  the  underlying  zone  of  normal 
fluctuations.  The  normal  range  of  water  table  fluctuations  should  de- 
crease to  shallower  limits  during  a  climatic  or  geologic  episode  as  open- 
ings within  the  karst  groundwater  zone  enlarged  by  solution  to  cavernous 
routes  of  proportions  capable  of  containing  normal  amounts  of  infiltration. 

Heavy  precipitation  or  snow  melt  may  flood  integrated  openings 
causing  a  rise  in  the  water  table,  an  increase  in  hydrostatic  head,  and  a 
high  velocity  within  the  integrated  openings  with  a  corresponding  in- 
crease in  solution  rates.  The  greater  the  velocity  of  a  solvent,  the 
greater  the  rate  of  solution  and  the  greater  the  amount  of  solvent,  the 
greater  the  amount  of  solution.  Thus  a  large  volume  of  solvent  becomes 
partly  saturated  much  more  rapidly  than  a  small  volume  becomes  com- 
pletely saturated  (6).  Swinnerton  (14)  stated  that,  "A  solution  of  low 
concentration  dissolves  more  rapidly  than  a  highly  concentrated  solution. 
Four  volumes  of  water  become  one-fourth  saturated  with  CaCO;>(  or  any 
similar  solute  under  ordinary  conditions  in  far  less  time  than  one  volume 
becomes  completely  saturated." 

Recent  papers  indicate  that  groundwater  in  carbonate  rocks  which 
is  nearly  saturated  with  calcium  carbonate  is  incapable  of  a  significant 
amount  of  solution  unless  it  changes  temperature  or  velocity,  or  is 
mixed  with  other  saturated  groundwater  of  a  different  temperature  (1, 
15).  They  have  also  based  nearly  all  of  their  cave  stream  analysis  on 
samples  taken  during  normal  flow  conditions  and  none  during  flood  stages, 
yet  the  latter  condition  fills  the  cavern  and  produces  ceiling  solution 
features  that  are  cited  as  evidence  for  development  below  the  water 
table.  Analysis  of  groundwater  taken  at  spring  in  Indiana  that  are 
discharging  from  caverns  indicate  that  from  about  150  to  300  ppm  of 
carbonate  and  sulphate  ions  are  removed  by  solution  during  normal  to 
low  flow  stages  (9).  The  concentration  of  the  solution  decreases  during 
flood  stages,  but  the  great  increase  of  volume  of  groundwater  transmitted 
results  in  a  net  increase  in  solution  of  the  carbonate  bedrock.  Nearly 
saturated  groundwater  has  previously  accomplished  the  significant  amount 
of  solution  of  which  it  was  capable. 

Most  theorists  have  noted  that  many  caverns  contain  evidence  of 
complete  water  filling,  at  least  in  early  stages  of  cavern  development, 
such  as  the  presence  of  solution  features  on  the  upper  walls  and  the 
ceiling  of  cavern  passages  (2).  Consequently,  they  have  based  their 
theories  on  the  presumption  that  the  caves  were  formed  at  a  time  when 
the  water  table  was  stable  and  permanently  above  the  zone  of  solution 


Geology  and  Geography  285 

and  cavern  formation  (15).  Solution  features  on  the  roofs  of  cavern 
passages  are  not  proof  of  permanent  saturation  nor  stable  water  table 
conditions.  They  prove  only  that  the  passages  were  filled  when  these 
features  were  developed.  Stable  water  table  conditions  are  probably 
non-existent.  Climates  are  not  stable,  and  if  the  water  table  were  to 
stabilize,  it  would  indicate  that  there  is  a  lack  of  imbalance  between 
intake  and  discharge  within  the  carbonate  bedrock  which  would  suggest 
a  lack  of  significant  groundwater  movement.  Thus,  solution  features  on 
cavern  ceilings  were  most  likely  developed  when  the  passages  were 
water  filled  with  moving  groundwater  during  a  temporary  high  position 
of  the  water  table. 

Base  Level  Control 

Base  level  has  been  cited  classically  as  a  control  of  surface  geomor- 
phic  erosion  cycles,  as  well  as  a  control  of  levels  at  which  caverns  may 
develop.  Ultimate  base  level  is  commonly  considered  as  a  landward 
extension  of  sea  level,  while  regional  or  local  base  level  is  defined  in 
terms  of  strata  or  materials  resistant  to  downcutting.  Base  level  in 
regard  to  cavern  development  has  been  designated  as  the  base  level 
of  solution,  generally  referring  to  non-soluble  rocks  below  the  carbonate 
host. 

Base  level  is  here  defined  at  the  top  of  the  zone  of  permanent  satu- 
ration (phreatic  groundwater  zone)  or  the  base  of  the  karst  groundwater 
zone  whenever  the  zone  of  permanent  saturation  was  relatively  stable 
during  any  definable  episode  of  cavern  development  (Fig.  2).  Such 
episodes  are  usually  contemporary  with  surficial  geomorphic  episodes. 
Thus  base  level  is  not  a  flat  planar  surface,  but  rather  is  a  gently  sloping- 
surface  graded  in  a  downstream  direction.  Surface  erosion  or  downcut- 
ting controls  the  position  of  the  somewhat  permanent  water  table.  Sub- 
terranean tributaries  do  not  develop  significantly  below  this  permanent 
water  table,  or  base  level  of  significant  groundwater  movement,  or  base 
level  of  significant  solution.  Caverns  that  develop  at  a  designated  base 
level  may  exhibit  irregularities  caused  by  differential  solution  of  various 
carbonate  or  non-carbonate  lithologies,  but  generally  they  reflect  the 
position  of  the  karst  groundwater  zone  that  existed  when  they  formed. 

Perched  water  bodies  caused  by  lithologic  differences  are  not  re- 
garded as  base  levels.  However,  a  karst  groundwater  zone  may  become 
perched  following  surface  rejuvenation  if  the  surface  stream  cuts  through 
an  impervious  layer  or  aquitard. 

Lithologic  Control 

A  few  theoretical  papers  have  considered  groundwater  flow  within 
a  carbonate  bedrock  as  transmission  of  phreatic  groundwater  within  a 
homogenously  permeable  medium  hundreds  of  feet  thick  (2,  3).  Stratified 
rocks,  including  the  carbonates,  are  seldom  of  uniform  lithology,  thick- 
ness, porosity  or  permeability  for  more  than  a  few  feet  vertically  and 
a  few  hundred  feet  laterally.  Thick  sequences  of  carbonate  strata  are 
usually   composed   of  various   thick,  medium,   and   thin-bedded   types   of 


286  Indiana  Academy  of  Science 

limestone,  shaly  and  sandy  limestones,  calcareous  shales  and  sandstones 
and  dolomites,  and  perhaps  including  non-calcareous  (non-soluble) 
strata,  which  are  all  of  different  solubility  rates.  The  intensity  of 
jointing-  within  each  type  of  lithology  also  varies  greatly  from  bed  to 
bed  and  according  to  the  location  in  relation  to  local  structural  features. 
Thus,  any  general  theory  which  assumes  a  homogeneously  permeable 
carbonate  bedrock  for  groundwater  movement  or  cavern  development,  for 
other  than  one  bed  in  a  very  small  area,  is  ignoring  the  most  important 
geologic  problem  concerned. 

Most  limestones  are  relatively  impermeable  rock  types,  although 
most  are  very  soluble  and  some  have  a  high  porosity.  Most  limestones 
owe  their  high  permeability  to  integrated  openings  along  bedding  planes 
and  joints  and  subsequent  solutional  enlargement  of  these  openings. 
Coarse-grained  and  porous  limestones  appear  to  be  more  readily  soluble 
than  fine-grained  or  compact  limestones.  The  more  dense,  compact,  thin 
limestone  beds  of  the  lower  part  of  the  Sanders  Group  and  the  litho- 
graphic limestone  beds  of  the  upper  part  of  the  Blue  River  Group  and 
some  of  the  limestone  units  of  the  West  Baden  and  Stephensport  Groups 
(Table  1)  commonly  project  as  resistant  ledges  into  the  larger  cave 
passages,  or  have  a  comparatively  narrow  slot  dissolved  through  them 
if  they  are  thick-bedded.  Conversely,  large  cavern  passages  formed  by 
solution  are  more  common  in  thick-bedded,  coarse-grained  or  porous 
limestones,  as  in  the  Salem  Limestone  Member.  Most  coarse-grained  and 
porous  limestones,  even  if  thin-bedded,  have  been  more  extensively  dis- 
solved and  form  recesses  in  the  cave  passages  formed  in  strata  of  the 
upper  part  of  the  Blue  River  Group  and  the  lower  part  of  the  Sanders 
Group.  The  compact,  denser  limestones  offer  less  surface  exposure  and 
allow  less  penetration  of  solvents  than  more  porous  and  coarse-grained 
limestones;  therefore,  the  latter  rocks  are  more  easily  removed  by  solu- 
tion. Rocks  low  in  carbonate  content  are  generally  less  soluble  than 
those  with  a  higher  carbonate  content.  Purity  of  the  limestones  in 
respect  to  high  calcium  carbonate  content  is  of  questionable  importance. 
Calcite  veins,  pure  calcium  carbonate  deposited  as  crevice  fillings,  com- 
monly protrude  as  resistant  ledges  and  dikes  in  cave  passages.  Caverns 
within  the  St.  Louis  Limestone  Formation,  which  is  generally  a  hetero- 
genous unit  of  argillaceous  limestones,  are  generally  unknown,  except 
for  small  solution  tubes.  Care  should  be  taken  to  distinguish  between 
large  solutionally-formed  cavern  passages  and  large  passages  which 
have  resulted  from  collapse  enlargement  of  a  small  solution  passage 
and  solutional  removal  of  the  debris  at  stream  level. 

Dolomites  are  less  readily  soluble  than  many  types  of  limestone,  but 
they  are  an  important  zone  of  groundwater  movement  within  the  Ste. 
Genevieve  Limestone  because  of  their  high  inherent  permeability.  Sig- 
nificant amounts  of  solution  occurs  in  some  dolomite  beds  simply  because 
the  dolomite  is  the  groundwater  transporting  medium  which  comes  into 
contact  with  the  acidic  groundwater.  Cavern  passages  developed  in 
association  with  dolomite  beds  are  known  in  several  Indiana  caverns, 
notably    in    Wyandotte    Cave,    where    solution    has    occurred    within    or 


Geology  and  Geography  287 

Table  1.  Sequence  of  important  carbonate  stratigraphic  units  of 
Mississippian  age  in  south-central  Indiana. 

Group  Formation 

Glen  Dean  Limestone 
Hardinsburg  Formation 
Stephensport  Golconda  Limestone 

Big  Clifty  Formation 
Beech  Creek  Limestone 

Elwren  Formation 
Reelsville  Limestone 
West  Baden  Sample  Formation 

Beaver  Bend  Limestone 
Bethel  Formation 


Paoli  Limestone 
Blue  River  Ste.  Genevieve  Limestone 


St.  Louis  Limestone 


Salem  Limestone 
Sanders  Harrodsburg  Limestone 


through  dolomite  beds  as  well  as  the  underlying  limestone  beds  owing 
to  discharge  of  water  from  the  dolomite  beds  into  the  underlying  lime- 
stone (12).  Very  permeable  porous  and  vuggy  dolomites  may  project 
as  ledges  because  the  groundwaters  pass  through  them  easily,  while 
some  very  fine-grained,  loosely-cemented  dolomites  are  easily  removed, 
perhaps  partly  by  mass  wasting  and  mechanical  erosion.  Medium  beds 
of  dolomite  in  several  Indiana  caverns  have  been  widened  within  cave 
passages  more  easily  than  adjacent  limestone  beds.  Some  thick  beds  of 
dolomite,  in  now  abandoned  passages,  have  apparently  caused  passage 
enlargement  by  exfoliation  or  disintegration  by  weathering. 

Calcareous  shales  that  are  particularly  susceptible  to  solution  occur 
stratigraphically  within  the  upper  part  of  the  Blue  River  Group.  These 
shales  are  very  soluble  and  passages  within  several  caves  have  developed 
partly  by  solutional  removal  of  the  shale,  perhaps  aided  by  some  me- 
chanical erosion.  Shales,  including  calcareous  and  non-calcareous  varieties, 
are  generally  compact  and  appear  to  lack  significant  open  joints;  thus, 
they  are  commonly  barriers  to  vertically  moving  groundwater.  Perched 
water  bodies  occur  on  the  top  of  the  shale  within  its  unbreached  limits. 
However,  once  breached  the  perched  water  is  discharged  into  the  under- 
lying carbonate  strata.  If  the  relative  relief  between  the  perched  water 
body  on  the  shale  and  the  normal  water  level  of  the  karst  groundwater 
in  the  underlying  carbonate  strata  is  sufficient  and  if  the  descending 
water  is  transmitted  through  a  vertical  joint  or  set  of  joints  that  pene- 
trates several  beds,  a  vertical  shaft  or  pit  may  result  from  water  running 
down  the  walls.  Progressive  upstream  migration  of  the  breached  edge 


288  Indiana  Academy  of  Science 

of  the  shale  may  result  in  a  laterally  elongated  shaft,  or  successive 
breaching  at  different  points  may  cause  successive  abandonment  of  the 
former  horizontal  cave  passage  and  each  preceding  shaft. 

Vertical  solution  shafts  or  pits  are  formed  below  any  lithology  such 
as  shale,  sandstone,  limestone,  or  dolomite  which  is  relatively  resistant 
to  solution  in  comparison  to  carbonate  strata  below,  and  which  is  over- 
lain with  relatively  permeable  strata  or  material.  The  overlying  perme- 
able unit  serves  as  a  perched  water  body  reservoir  for  groundwater  to  be 
discharged  through  an  opening  in  the  resistant  bed.  An  opening,  such 
as  a  joint,  through  the  resistant  bed  allows  somewhat  regulated  discharge 
into  the  underlying  carbonate  rocks.  The  character  of  the  reservoir  rock 
and  the  opening  in  the  resistant  bed  determines  the  rate  of  groundwater 
discharge  into  the  underlying  strata.  Vertical  shafts  or  pits  are  enlarged 
primarily  by  regulated  flows  or  seepage  down  the  walls  of  the  vertical 
opening  rather  than  by  complete  flooding  and  filling  which  is  probably 
possible  only  in  the  initial  small  openings.  Some  vertical  solution  shafts 
are  now  fed  directly  by  diverted  surface  drainage  and  they  may  have 
originated  by  this  means. 

Thin  sandstone  beds,  particularly  the  Popcorn  Sandstone  Bed  at  the 
base  of  the  Paoli  Limestone  Member,  serve  as  resistant  beds  to  form 
perched  water  bodies  within  overlying  limestones,  discharging  water 
through  restricted  outlets  into  underlying  strata,  commonly  forming 
vertical  solution  shafts,  or  laterally  elongated  shafts.  Perched  ground- 
water within  thick  sandstone  units,  such  as  those  within  the  West  Baden 
and  Stephensport  Groups,  is  the  prime  source  for  the  groundwater  which 
develops  caverns  in  underlying  limestones  (11).  Groundwater  movement 
from  a  perched  water  body  in  a  permeable  sandstone  into  a  limestone 
as  a  means  of  cavern  development  is  probably  the  major  exception  to  the 
concept  that  most  caverns  are  formed  by  groundwater  solution  within 
a  karst  groundwater  zone. 

Groundwater  movement,  and  subsequent  cavern  enlargement,  in 
south-central  Indiana  is  primarily  along  the  joints  and  bedding  planes 
in  the  carbonate  strata.  Indiana  caverns  commonly  have  developed 
parallel  to  the  dip  of  the  strata  following  sets  of  joints  that  allow  the 
steepest  gradient  to  the  nearest  or  lowest  surface  outlet.  Most  Indiana 
cave  passages  trend  with  the  regional  structure  which  has  average  dips 
of  about  25  feet  per  mile  to  the  southwest.  There  are  local  structures, 
however,  which  deviate  from  this  pattern,  and  caverns  situated  on  these 
structures  follow  the  local  dip.  Some  caverns  follow  the  strike  of  the 
structure  where  the  karst  groundwater  zone  within  the  particular  cavern- 
ous strata  has  no  outlets  in  a  downdip  direction  along  streams  which 
dissected  the  cavernous  strata  or  karst  groundwater  zone. 

Each  bed,  depending  upon  its  lithologic  type  and  thickness,  has  a 
unique  pattern  of  joints  or  joint  sets.  Generally  any  particular  set  of 
joints  does  not  extend  into  overlying  or  underlying  strata,  but  some  joints 
are  common  to  several  strata,  and  some  master  joints  apparently  extend 
through    many    beds    of    dissimilar    lithologies.    Thin-bedded    strata    are 


Geology  and  Geography  289 

usually  more  intensely  jointed  than  thiek-bedded  units.  In  general  it 
seems  that  there  is  a  joint  spacing-  of  a  particular  bed  that  is  related 
to  the  unit  thickness  and  type  of  lithology.  Although  the  joints  from  bed 
to  bed  may  not  be  superposed,  they  do  cross  and  this  allows  vertical 
groundwater  movement  without  necessitating  significant  flowage  along 
bedding  planes. 

Multi-level  cavern  passages  that  have  developed  in  progressive  stages 
as  tributaries  related  to  adjacent  surface  drainage  levels  or  geomorphic 
episodes  commonly  reflect  the  effect  of  joint  patterns  shifting  from  bed 
to  bed.  Passages  formed  along  a  master  joint  which  dissects  numerous 
beds,  usually  have  a  canyon-like  appearance  with  wide  places  repre- 
senting temporary  halts  in  solutional  deepening  or  more  soluble  strata. 
In  addition,  passages  in  one  stratum  may  form  an  orientation  pattern 
related  to  somewhat  different  intake  and  discharge  areas  than  passages 
at  a  different  level  within  another  stratum.  Goss  Cave,  in  Washington 
and  Harrison  Counties,  is  an  excellent  example.  The  upper  levels  have 
developed  within  one  set  of  joints  in  the  upper  part  of  the  Salem  lime- 
stone owing  to  a  general  infiltration  of  meteoric  water,  while  the  lower 
level  within  the  lower  part  of  the  Salem  Limestone  has  developed  pri- 
marily by  surface  run-off  diverted  into  sinkholes  in  addition  to  water 
diverted  from  the  upper  levels.  Similar  subterranean  diversions  in 
Wyandotte  Cave  have  been  described  (12). 

An  important  cause  of  the  down  dip  development  of  caverns  and 
groundwater  movement  is  the  dissection  of  permeable  strata  by  en- 
trenched surface  drainage  in  places  down  dip  from  intake  or  recharge 
areas.  Gardner  (5)  based  his  "static  water  zone"  theory  of  cavern  de- 
velopment on  the  concept  that  dissection  of  permeable  strata  was  re- 
sponsible for  initiating  groundwater  movement  and  solution  of  caverns. 
Although  much  of  his  theory  is  correct,  he  places  too  much  importance 
on  structural  control  of  cavern  development.  Other  aspects  which  he 
mentions,  such  as  climate,  are  equally  important. 

Climatic  Control 

Numerous  authors  have  either  suggested  that  the  misfit  streams 
in  modern  caverns  are  not  large  enough  to  have  dissolved  the  cavern, 
and  that  therefore  they  were  either  formed  in  the  past  when  the  passages 
were  water  filled  below  a  permanently  higher  water  table  (3),  or  they 
have  been  formed  by  diversion  of  surface  streams  to  subsurface  routes 
at  flood  stages  (7)  or  by  subterranean  stream  piracy  (16). 

Although  the  streams  in  most  Indiana  caverns  are  now  misfit,  it  is 
obvious  that  each  level  was  essentially  filled  with  water  at  times  during 
its  period  of  solutional  enlargement.  The  thick  and  extensive  cave  sedi- 
ments present  in  Indiana  caverns  are  further  evidence  of  drastic  changes 
in  cave  stream  regimen  that  have  taken  place,  particularly  where  the 
deposits  are  within  abandoned  upper  levels  which  no  longer  contain  a 
stream.  Some  of  the  passages  which  contain  misfit  streams  now  flood 
with   even   light   precipitation,    but   most   are   known   to    flood    only   in- 


290  Indiana  Academy  of  Science 

frequently  during  exceptionally  heavy  periods  of  precipitation.  Some 
water  filled  caverns  now  exist  only  because  they  are  flooded  owing  to 
alluvial  sediments  which  have  dammed  their  outlets  (10). 

Three  factors,  misfit  cave  streams,  abandoned  cave  levels  and  ex- 
tensive cave  sediments,  are  proof  that  the  carbonate  bedrock  area  of 
south-central  Indiana  has  been  subjected  to  multiple  cycles  of  wetter 
and  drier  conditions.  Further  proof  of  significant  climatic  change  is 
available  in  the  interpretation  of  the  Pleistocene  history  of  the  immedi- 
ately adjacent  areas.  At  least  three  ice  sheets  of  the  Kansan,  Illinoian 
and  Wisconsinan  glaciations  were  close  enough  to  the  area  to  effect 
moister  conditions  than  during  the  interglacial  stages.  These  wetter 
epochs  undoubtedly  caused  more  frequent  temporary  filling  of  the  solu- 
tional  openings  within  the  karst  groundwater  zones  existing  during 
each  glacial  episode,  with  solutional  enlargement  taking  place  at  a 
corresponding  rate.  Waning  of  the  glacial  climatic  conditions  would 
result  in  a  decrease  of  precipitation,  a  change  to  drier  conditions  and 
the  subsequent  misfit  stream  conditions. 

Ancient  climatic  changes  would  also  have  caused  changes  in  the 
amount  and  type  of  vegetation  in  south-central  Indiana.  A  dense  wood- 
land cover  during  a  wet  climatic  episode  would  have  caused  a  lack  of 
surface  run-off  with  a  corresponding  increase  in  groundwater  infiltration 
and  an  increase  in  groundwater  acidity,  whereas  a  drier  climatic  episode 
would  cause  a  decrease  in  vegetal  cover,  an  increase  in  surface  run-off 
and  perhaps  in  sedimentation  in  surface  and  subsurface  drainage  routes. 

Although  some  evidence  is  available  to  correlate  certain  cave  levels, 
cave  sediments  or  karst  groundwater  conditions  with  specific  geologic 
events,  data  are  generally  lacking  to  devise  an  exact  correlation  of  all 
cavern  development  episodes  with  known  geologic  events.  Data  available 
at  this  time  suggest  that  episodes  of  cavern  solution  and  deposition  per- 
haps are  the  only  evidence  of  geologic  or  climatic  episodes  which  are  yet 
to  be  detected  as  surface  geomorphic  features.  That  is,  evidences  of 
past  climates  and  geomorphic  events  may  perhaps  be  best  interpreted 
from  evidence  obtained  in  caverns  where  the  evidence  has  not  been  de- 
stroyed by  subsequent  events.  Multiple  cavern  levels  in  south-central 
Indiana  and  the  cave  sediments  deposited  within  them  appear  to  record 
a  sequence  of  events  ranging  in  age  from  late  Tertiary  to  Recent  times. 
Each  of  the  cavern  levels  was  obviously  initially  formed  by  solution  within 
a  karst  groundwater  zone  and  perhaps  later  filled  with  cave  sediments 
as  each  succeeding  cave  level  or  karst  groundwater  zone  became  re- 
established at  another  level,  higher  or  lower  than  the  preceding  level, 
owing  to  changes  in  base  level  and  climate. 


Geology  and  Geography  291 

Literature  Cited 

1.  BOGLI,   A.    1965.   The  role  of   corrosion  by   mixed   water   in   cave  forming.    Problems   of 
the  Speleological  Research.  Prague,  p.  125-131. 

2.  Bretz,    J.    H.     1942.    Vadose    and    phreatic    features    of    limestone    caverns.    J.    Geol. 
50:675-811. 

3.  Davis,  Wm.  M.  1930.  Origin  of  limestone  caverns.  Bull.  Geol.  Soc.  Amer.  41  :475-628. 

4.  Finch,    J.    W.    1904.    The   circulation    of   underground    aqueous    solutions.    Proc.    Colo. 
Sci.  Soc.  7:193-252. 

5.  Gardner,   J.    H.    1935.    Origin   and   development  of   limestone  caverns.    Bull.    Geol.    Soc. 
Amer.  46:1255-1274. 

6.  Kaye,    C.    A.     1957.    The    effect    of    solvent    motion    on    limestone    solution.    J.    Geol. 
65:35-46. 

7.  Malott,   C.    A.    1938.    Invasion   theory   of   cavern    development    (Abstract)     p.    323.    In 
Geol.  Soc.  Amer.  Proc.  for  1937. 

8.  Meinzer,    O.    E.     1923.    Outline    of    groundwater    hydrology,    with    definitions.    U.     S. 
Geol.  Surv.  Water-Supply  Paper  494.  71  p. 

9.  Powell,    R.    L.    1961.    A    geography    of    the    springs    of    Indiana.    Unpublished    M.A. 
Thesis.  Indiana  University. 

10.    .    1963.    Alluviated    cave    springs    in    south-central    Indiana.    Proc.    Indiana 

Acad.  Sci.  72:182-189. 

11.    .    1966.    Groundwater    movement    and    cavern    development    in    the    Chester 


Series  of  Indiana.  Proc.  Indiana  Acad.  Sci.  75:210-215. 

12.    .     1968.    The    geology    and    geomorphology    of    Wyandotte    Cave,    Crawford 

County,  Indiana.  Proc.  Indiana  Acad.  Sci.  77  :236-244. 


13.    .     1969.     Cavern    development    in    a    karst    groundwater    zone     (Abstract) 

p.   38.   In  Geol.   Soc.   Amer.   Absti-acts  with  programs  for   1969.   Part   6. 

14.  Swinnerton,     A.     C.     1932.     Origin    of    limestone    caverns.     Bull.     Geol.     Soc.     Amer. 
43  :663-694. 

15.  Thrailkill,    J.     W.     1968.     Chemical    and    hydrologic    factors    in    the    excavation    of 
limestone  caverns.  Bull.  Geol.  Soc.  Amer.  79:19-46. 

16.  Woodward,    H.    P.    1961.    A   stream    piracy    theory    of    cave    formation.    Nat.    Speleol. 
Soc.  Bull.  23:39-58. 


Cooling  Degree  Days  in  Indiana 

Lawrence  A.  Schaal,1  Purdue  University 

Introduction 

Abstract 

Cooling  degree  days  are  used  by  the  air-conditioning  industry  to  estimate  power 
requirements  for  cooling  buildings  as  temperatures  vary  from  day  to  day,  seasonally  and 
geographically.  Normal  cooling  degree  days  are  calculated  for  various  climatic  stations 
in  Indiana  and  the  appropriateness  of  the  base  temperature  used  will  be  related  to 
daily  data  recorded  from  a  typical  Indiana  home. 

Cooling  degree  days  is  a  meteorological  statistic  derived  from 
accumulating  the  excess  of  the  daily  mean  outdoor  temperature  above  a 
chosen  base  temperature.  They  closely  parallel  the  energy  necessary  to 
cool  for  human  comfort  the  interior  of  buildings  exposed  to  a  hot  exterior 
environment.  The  chosen  base  temperature  has  ranged  from  60  to  75  °F. 
This  study  will  hopefully  add  to  the  apparent  meager  work  done  on  this 
question  for  ordinary  homes,  or  small  buildings.  Normal  cooling  degree 
days  are  calculated  for  some  weather  stations  in  Indiana  from  monthly 
mean  temperatures  and  a  chart  for  the  state  is  constructed  from  the 
data. 

One  cooling  degree  day  is  a  day  when  the  mean  temperature  exceeds 
the  base  temperature  one  degree.  Lower  mean  temperatures  are  given  a 
value  of  zero.  The  resulting  sum  for  a  day,  a  week,  or  season  enables  com- 
parisons from  one  area  to  another,  within  various  time  periods,  and  in  a 
locale  to  assess  cooling  energy  requirements  of  structures.  Heating 
degree  days  which  are  calculated  from  mean  temperatures  below  65  °F 
have  a  wide  use  in  the  heating  industry,  a  fine  example  of  profitable  use 
of  climatology. 

Interest  in  this  study  started  as  the  author  considered  the  appropri- 
ateness of  65  °F  as  the  standard  base  temperature  for  cooling  degree 
days.  The  Glossary  of  Meteorology  (3)  gives  the  common  base  of  75 °F. 
In  selected  Climatic  Maj)s  of  the  United  States  (2)  the  base  temperature 
used  is  65 °F.  The  Climatological  Handbook,  Columbia  Basin  States  (4) 
suggests  60  °F.  In  Indiana,  a  daily  mean  temperature  of  65  °F  results 
from  minimum  temperature  for  the  day  of  about  54 °F  and  a  maxium 
temperature  of  about  76  °F.  An  air  temperature  of  76  °F  is  still  in  the 
comfortable  range  for  non-working  humans  under  common  humidity, 
wind  and  solar  conditions.  But  as  described  in  this  paper,  the  solar 
radiation  load  on  a  small  building  seems  to  require  a  lower  base 
temperature  at  least  when  the  maximum  interior  temperature  is  set 
at  78  °F. 

The  correlation  of  cooling  requirements  for  several  types  of  com- 
mercial buildings  was  reported  by  Thorn    (7).   Correlations  in  a   15-day 


1  Agronomy   Department  and   Environmental    Science   Services   Administration,    United 
States  Department  of  Commerce. 

292 


Geology  and  Geography 


293 


study  in  Washington,  D.  C.  ranged  from  0.405   (office  building)   to  0.918 
(hotel).  The  base  temperature  was  65 °F. 

Methods 

In  this  low  budget  study  the  author  compared  the  minutes  the  air 
conditioner  ran  in  his  own  home  to  the  measurements  of  air  temperature 
and  solar  radiation  at  the  Purdue  University  Agronomy  Farm  6  miles 
west-northwest  of  the  house.  Readings,  taken  for  42  days  at  about  8  AM, 
consisted  of  the  total  minutes  the  home  thermostat  set  at  78  °F  caused 
the  coolant  compressor  to  run  and  cool  the  house  (Fig.  1). 


58 


One-tenth  of  day's  solar 
radiation  given  at  points, 


Line  equation: 


31.75  T  -  1888.9 


200 


300       400      500 
COOLING  MINUTES 


600 


700 


800 


Figure  1.  Daily  observations  of  outdoor  mean  temperature  are  related  to  minutes 
of  cooling  to  maintain  a  temperature  of  78° F  or  lower  in  a  typical  dwelling.  The 
day's  solar  radiation  in  tens  of  calories  per  square  centimeter  is  given  at  the  point. 
Cooling  minutes  accumulated  when  the  daily  mean  temperature  exceeded  59° F  generally. 


The  circulating  fan  of  the  furnace  ran  continuously  and  the  ther- 
mostat was  set  to  turn  on  the  coolant  compressor  when  78  °F  was 
exceeded  during  the  42  days.  A  simple  electric  clock  was  connected  to 
the  circuit  on  the  furnace  which  actuated  the  coolant  compressor  and  the 
coolant  fan  outside  the  house.  The  room  thermostat  closed  the  circuit  for 
running  the  cooling  equipment. 

Cooling  minutes  used  in  this  paper  could  be  converted  directly  to 
electrical  energy  by  using  the  electrical  specifications  of  the  electric 
motors. 


294  Indiana  Academy  of  Science 

Of  some  interest  was  the  frequency  and  time  periods  when  the 
cooling-  system  ran.  These  were  monitored  by  placing  a  hygrothermo- 
graph  beside  the  outlet  of  the  furnace.  Thus  the  rise  and  fall  of  outlet 
air  temperature  and  humidity  were  charted.  These  data  made  it  quite 
obvious  how  the  radiation  load  on  the  house  had  to  be  removed  by  the 
air  conditioner.  Cooling  began  on  many  days  at  about  noon  and  con- 
tinued until  evening  or  midnight. 

Results  and  Discussion 

Linear  regression  was  calculated  for  the  42  daily  observations  using 
C  =  aT  +  k  where:  C  was  cooling  minutes;  T  equalled  the  mean  tem- 
perature for  the  24-hour  period  (the  average  of  the  maximum  and 
minimum  temperature);  a  was  the  regression  coefficient  of  T;  and  k 
was  the  constant.  The  equation  of  the  regression  line  was: 
C  =  31.75  T—  1888.9 

1888  9 

when  C  =  0  then  T  = —  =  59.5 

31.75 

The  intercept  on  T  when  cooling  minutes  were  zero  was  59.5  °F.  This 
data  suggests  a  base  temperature  of  about  60  °F.  The  simple  correlation 
coefficient  was  0.880. 

During  the  period  of  observation  it  was  obvious  that  solar  radiation 
had  an  effect  on  the  minutes  of  cooling.  For  this  reason  solar  radiation 
observed  at  the  Agronomy  Farm  became  the  second  factor  in  a  multiple 
regression  problem.  The  effect  of  showers  was  not  studied.  There  were 
very  few  showers  during  the  42  days  but  during  one  evening  a  shower 
cooled  the  house  and  environs  reducing  the  cooling  minutes  compared  to 
other  days  at  the  same  temperature.  Showers  favor  shifting  the  base 
temperature  higher  since  cooling  minutes  are  less.  A  sunny  day  favors 
a  lower  base  temperature  because  the  interior  temperature  of  the  house 
increased  by  solar  radiation  greatly  exceeds  exterior  air  temperature.  The 
multiple  correlation  equation  using  both  temperature  and  solar  radiation 
was  C  =  aT  +  bS  +  k  where:  C  =  cooling  minutes;  T  =  mean  tempera- 
ture of  day;  S  =  solar  radiation  (daily  total  gram  calories  on  a  square 
centimeter  of  horizontal  surface).  The  coefficients  calculated  were: 
a  =  30.15;  b  =  0.288;  resulting  in  C  =  30.15  T  +  0.288  S  —  1939.24.  If 
the  first  equation  for  C  (obtained  when  the  solar  radiation  term  was  not 
included)  is  placed  equal  to  the  above  equation  including  the  solar  radia- 
tion term,  we  obtain:  31.75  T  —  1888.87  =  30.15  T  +  0.288  S  —  1939.24; 
1.6  T  =  0.288  S  —  50.37;  T  =  0.18  S  —  31.48.  This  shows  a  relation  of 
1°F  temperature  change  equal  to  0.18  Langley  where  the  two  linear 
equations  intersect.  In  the  above  equation  the  constant  1939.24  is  a  little 
greater  than  the  1888.9  of  the  first  equation  which  did  not  have  the 
solar  radiation  term.  The  intercept  of  T  with  zero  minutes  of  cooling 
turns  out  to  be   61.1  °F. 

The  coefficient  of  T  and  S  may  also  be  normalized  to  show  a  relative 
relationship  for  one  standard  deviation  of  C  by  calculating  Beta  Coeffi- 
cients   (1).    This   is    done    by    dividing    the    coefficients    a    and   b    by    the 


Geology  and  Geography  295 

standard  deviation  of  C  which  was  calculated  to  be  172.48.  They  were 
next  multiplied  by  their  respective  standard  deviations  as  follows:  Cal- 
culated  standard   deviations:   for   C   172.48;   for   T   4.781;    for   S.    155.38. 

30.15    X    4.781 
Beta  for  T  = —  =  0.8357; 

172.48 

0.2884   X    155.38 
Beta  for  S  = =  0.2598. 

172.48 

Therefore  one  standard  deviation  of  C  relates  to  0.84  standard  deviation 
of  T  and  0.26  standard  deviation  of  S. 

The  significance  of  solar  radiation  to  cooling  minutes  was  calculated 
as  follows  (N  =  42): 

Source  of  Variation  Degrees  of  Freedom  Sum  of  Squares 
C  on  T                                                                    40  275,116 

C  on  T  and  S  39  195,118 

Reduction  due  to  adding  S  1  79,928 

79  928            79  928 
For  the  well  known  F  test, =  — ■ =  15.97 

195,188/39  5005 

From  F-table  values  (5),  an  F  of  8.87  or  greater  indicates  that  the 
chance  is  less  than  1  in  200  that  this  set  of  observations  accidentally 
showed  such  a  contribution  from  solar  radiation  increasing  cooling 
minutes  after  the  effect  of  temperature. 

The  correlation  coefficients  based  on  additional  tests  and  computa- 
tions are  listed  in  Table  1. 


Table  1.  Correlation  coefficients. 

Daily  mean  temperature  and  cooling  minutes  0.880 

Daily  highest  temperature  and  cooling  minutes  0.881 

Daily  mean  temperature  and  solar  radiation  with  cooling  minutes  0.917 

Daily  highest  temperature  and  solar  radiation  with  cooling  minutes  0.886 


Linear  correlation  of  the  maximum  temperature  with  cooling  min- 
utes was  about  the  same  as  with  the  mean  temperature  and  did  not 
improve  much  with  the  addition  of  the  solar  radiation  term.  This  seems 
to  indicate  that  the  maximum  temperature  reflected  the  effect  of  solar 
radiation  with  very  little  contribution  by  the  minimum  temperature  or 
night  temperature. 

The  importance  of  interaction  between  temperature  and  solar  radia- 
tion (product  of  T  and  S)  to  cooling  was  computed.  The  addition  of  this 
third  term  to  the  equation  became  significant  at  the  5%  level  but  not  at 
the  1%  level  using  the  F  test. 


296 


Indiana  Academy  of  Science 


The  one-story  house  of  about  1200  square  feet  plus  a  full  basement 
was  insulated  by  4-inch-thick  bats  of  fiber  glass  insulation  in  ceilings 
and  walls  and  is  believed  fairly  typical  of  the  contemporary  3-bedroom 
home.  The  roof  was  grey  with  a  slope  of  3  to  12.  The  window  area  was 
more  than  the  average  house,  and  storm  windows  were  in  place.  Con- 
siderable attic  ventilation  came  from  openings  under  the  eaves.  It 
seems  that  the  solar  radiation  load  was  proportionately  larger  than  for 
a  large  apartment  or  office  building  having  large  interior  areas  away 
from  exterior  walls  and  roof.  Afternoon  relative  humidity  varied  little 
during  the  experiment.  Daily  lows  ranged  from  39  to  86  at  the 
Agronomy  Farm  and  averaged  53%  relative  humidity. 


It                    r\        j-\                        "                                                                                                                         "' 

900      A 

i— L- 1 

1000 

•  900     I 

1 
1 

*100C 

! 

V1 

1100 

— X_ 

— — 

r 

1200 

1       ^ 

13( 

)0> 

^w^ 

^y 

S T 

SJ 

44-noo 

1— J*»1200 

COOLING 
DEGREE 
DAYS 
NORMAL 

ANNUAL       . 

14 

i"" 

00 

kh 

1/1300 

D<>140C 

) 

BASE   65° 

Figure   2.     Normal  annual  cooling   degree  days  calculated  from   daily  mean  temperatures 
above  C5°F. 


Geology  and  Geography  297 

I  conclude  that  in  a  sunny  climate  and  for  small  houses  the  base 
temperature  for  cooling  is  closer  to  60 °F  than  65  °F,  but  perhaps  for 
large  multistoried  buildings  the  base  temperature  may  be  more  appro- 
priately 65 °F.  Since  65 °F  is  becoming  the  most  frequently  used  base 
temperature,  a  normal  cooling  degree  day  map  for  Indiana  has  been 
drawn  (Fig.  2).  Monthly  mean  temperatures  for  the  period  of  1931-1960 
were  used  since  the  World  Meteorological  Organization  and  Environ- 
mental Science  Services  Administration,  Environmental  Data  Service 
use  this  average  as  the  normal. 

The  monthly  mean  temperature  of  the  mid-summer  months  readily 
convert  to  cooling  degree  days  with  practically  no  error  by  subtracting 
65  from  the  mean  and  multiplying  by  the  number  of  days  in  the  month. 
However,  for  the  peripheral  months  of  the  season  when  the  mean  is 
near  65  we  used  the  standard  deviation  of  the  monthly  mean  tempera- 
ture to  estimate  the  additional  degree  days  needed.  Thorn's  unpublished 
nomogram   (6)   was  used  to  estimate  the  increment. 

For  an  example  in  the  use  of  Figure  2,  consider  a  location  in  Indiana 
where  the  normal  of  cooling  degree  days  is  1200  for  a  season.  A  sum- 
mer has  just  passed  when  the  cost  of  cooling  a  new  building  was  $300 
and  the  cooling  degree  days  totalled  1500  as  calculated  from  daily  tem- 
peratures nearby.  What  is  the  normal  expected  expense  ?  Since  1500 
cooling  degree  days  is  300  more  than  the  normal  1200  and  300  is  20% 
of  1500,  then  20%  of  $300,  or  $60,  is  the  above  normal  cost  of  cooling  the 
building.  A  normal  or  average  summer  would  result  in  cooling  costs  $60 
less  than  $300,  or  $240.  Supposing  the  next  summer  averaged  near  normal 
in  cooling  degree  days  and  costs  were  appreciably  different  from  $240.  It 
would  then  be  evident  that  changes  in  the  efficiency  of  cooling  had 
occurred. 

The  Indiana  map  relates  to  the  standard  exposure  of  official  ther- 
mometers generally  found  in  rural  or  village  areas.  These  localities  are 
generally  cooler  than  the  brick  and  concrete  areas  of  the  cities.  There- 
fore city  islands  of  heat  are  not  shown  unless  included  unknowingly  in 
some  substation  data  used. 

Conclusions 

In  conclusion,  this  experiment  favored  a  base  temperature  nearer  60 
than  65  °F  for  the  computation  of  cooling  degree  days  as  they  relate  to 
home  cooling.  The  test  period,  however,  did  cover  a  sunny  period  and  a 
humidity  much  higher  than  found  in  the  western  plains,  both  of  which 
should  be  inversely  related  to  the  base  temperature. 

Acknowledgments 

The  author  is  indebted  to  James  E.  Martin  and  Oscar  Luetkemeier 
for  Agronomy  Farm  observations,  to  Walter  L.  Stirm,  Advisory  Agricul- 
tural Meteorologist  for  instrumentation,  and  Robert  May  for  computer 
work. 


298  Indiana  Academy  of  Science 

Literature  Cited 

1.  Arkin,  Herbert,  and  Raymond  R.  Colton.  1946.  An  Outline  of  Statistical  Methods, 
4th  Edition.  Barnes  and  Noble  Inc.,  N.  Y.  224  p. 

2.  Data  Information.  1968.  Selected  Climatic  Maps  of  the  United  States.  Environ- 
mental   Science    Services    Administration,    U.    S.    Department   of    Commerce.    32    p. 

3.  Huschke,  Ralph  E.  1959.  Glossary  of  Meterology.  American  Meteorological  Society. 
638  p. 

4.  Meteorology  Committee,  Pacific  Northwest  River  Basins  Commission.  1969.  Climato- 
logical  Handbook,  Columbia  Basin  States,  Temperature,  Vol.  1,  Part  B.  540  p. 

5.  Snedecor,  George  W.  1965.  Statistical  Methods,  Fifth  Edition.  Iowa  State  University 
Press,  Ames,  la.  534  p. 

6.  Thom,  H.  C.  S.  1956.  Monthly  Degree  Days  derived  from  Monthly  Mean  Tempera- 
ture, nomogram.  Environmental  Data  Service,  Environmental  Science  Services  Ad- 
ministration, U.  S.  Department  of  Commerce. 

7.  Thom,  H.  C.  S.  1966.  Cooling  Degree  Days  and  Energy  Consumption.  Air  Condition- 
ing, Heating  and  Ventilation.  63  :53-54. 


The  Climatology  of  Indiana  Tornadoes 

Ernest  M.  Agee,  Purdue  University 

Abstract 

Various  tornado  statistics  for  Indiana  have  been  determined  based  upon  climato- 
logical  records  from  1916  through  1968.  Yearly,  monthly,  and  diurnal  variations  of 
tornado  frequencies,  injuries,  and  deaths  are  examined  on  a  state-wide  and  county 
basis.  An  attempt  has  been  made  to  remove  the  problem  of  the  population  bias  in 
tornado  reporting.  Also,  the  effect  of  terrain  on  the  areal  distribution  of  tornadoes  is 
noted.    Further   study    into   the   matter   of   orographic    effects    is   suggested. 

Introduction 

A  national  summary  of  tornado  statistics  by  Flora  (1)  has  provided 
the  scientific  community  with  considerable  information  on  tornado  fre- 
quencies and  occurrence  patterns.  It  is  recognized,  however,  that  the 
scope  of  study  in  a  national  summary  does  not  provide  the  detailed 
information  that  is  often  desired  on  a  state-wide  basis.  Precedence  for 
the  value  of  state  summaries  of  tornado  statistics  and  related  climato- 
logical  inferences  is  evident  in  the  work  by  Darkow  (unpublished  data) 
on  Missouri  tornadoes.  In  fact,  the  study  by  Darkow  provided  consider- 
able incentive  to  do  a  similar  study  for  Indiana,  especially  since  it  is 
well  above  the  national  average  as  a  tornado-producing  state.  Indiana 
has  been  involved  in  some  of  the  most  dramatic  tornado  outbreaks  of 
this  century.  Most  notable  are  the  tri-state  tornadoes  of  1925  and  the 
Palm  Sunday  tornadoes  of  1965. 

The  primary  objectives  of  this  investigation  were  to  determine 
yearly,  monthly,  and  diurnal  variations  of  tornado  frequencies,  injuries, 
and  deaths  for  Indiana.  In  addition  to  obtaining  an  areal  distribution  of 
tornadoes,  a  limited  effort  was  made  to  examine  the  effects  of  population 
and  topography.  Tornado  data  for  the  United  States  (2)  do  show  a 
tendency  for  decreasing  tornado  frequencies  toward  the  southeast  corner 
of  Indiana,  which  could  be  attributed  to  population  and /or  terrain 
effects.  It  was  one  of  the  intentions  of  this  study  to  substantiate  or 
dismiss  this  observation  by  examining  the  data  in  greater  detail  on  a 
state-wide  basis.  Finally,  it  was  hoped  that  this  work  would  also  provide 
a  suitable  format  and  method  for  examining  the  climatology  of  tor- 
nadoes for  other  states.  A  generalized  computer  program  has  been 
compiled  and  is  available   to   other  investigators. 

Methods 

The  period  of  this  study  dates  from  1916,  the  beginning  of  official 
tornado  records,  through  1968.  Data  assembled  from  United  States 
Weather  Bureau  records  (3,  4,  5,  6,  7)  were  checked  against  the  records 
maintained  by  Indiana's   State   Climatologist,   Mr.   Lawrence  A.   Schaal. 

Data  extraction  required  that  certain  decisions  be  made  in  making 
the  tornado  tallies.  Funnel  clouds  aloft  were  not  counted  but  water- 
spouts over  Lake  Michigan  were.  Also,  tornadic  winds  were  counted 
when    accompanied    by    path    damage    with    tornado    characteristics.    In 

299 


300  Indiana  Academy  of  Science 

nearly  all   cases,   however,   the   storm  was  logged   in   the  records   as   a 
tornado. 

The  format  of  the  data  reduction  was  prescribed  by  the  computer 
program  developed  simultaneously  to  handle  the  data  analysis.  Infor- 
mation recorded  included  the  date,  time  of  day,  and  place  of  origin  of 
tornado  by  county  (in  some  cases  out  of  state).  Also,  counties  affected, 
width  and  length  of  damage  path,  direction  of  movement,  injuries  and 
deaths,  and  estimated  property  damage  were  recorded.  All  available 
information  was  placed  on  one  computer  card  per  tornado.  Problems  in 
handling  the  data  included  such  matters  as  broken  storm  paths,  chang- 
ing directions,  large  time  intervals,  varying  damage  paths,  and  different 
translational  speeds. 

Other  data  collected  for  the  study  include  rural  population  figures 
per  Indiana  county  (9)  for  1920,  1940,  and  1960.  These  three  sets  of 
values  were  averaged  to  yield  a  mean  rural  population  per  county  and 
for  the  state  during  the  period  of  the  study.  Also  obtained  were  the 
number  of  square  miles  in  each  county  and  the  state. 

Results  and  Discussion 

During  the  period  from  1916  through  1968  a  total  of  551  different 
tornadoes  were  observed  and  recorded  in  climatological  data.  Of  this 
total,  20  came  from  Illinois,  2  from  Kentucky,  and  529  originated  in 
Indiana.  Some  of  those  tornadoes  only  touched  down  briefly  in  a  single 
county  while  others  had  continuous  paths  on  the  ground  through  several 
counties  and  into  the  adjacent  states  of  Michigan  and  Ohio. 

Figure  1  shows  the  number  of  tornadoes  affecting  each  Indiana 
county  from  1916  through  1968.  Twenty-four  tornadoes  were  observed 
in  Porter  County,  more  than  any  other.  Close  behind  are  Elkhart  with 
22,  Lake  with  20,  and  Marion  and  St.  Joseph  with  19  each.  These  coun- 
ties are  highly  populated  and  certainly  reflect  the  effect  of  population 
density  on  tornado  reporting.  No  tornadoes  have  been  officially  recorded 
for  Ohio  and  Crawford  counties.  Also  shown  are  the  number  of  tor- 
nadoes originating  in  each  county.  A  maximum  of  20  tornadoes  origi- 
nated in  Lake  County.  On  the  other  hand  none  have  originated  in 
Crawford,  Dearborn,  Ohio  and  Union  counties.  Of  course  counties  are 
political  boundaries  and  do  not  represent  equal  geometrical  areas.  But, 
a  county-by-county  analysis  has  been  made  for  the  sake  of  convenience. 
Figure  1  also  gives  the  number  of  tornado  days  per  county.  Porter 
County  has  the  highest  value  with  21. 

In  Figure  2  the  effects  of  population  and  unequal  county  areas 
have  been  considered.  To  reduce  the  number  of  tornadoes  affecting  each 
county  to  a  more  meaningful  statistic  a  tornado  index  has  been  formed. 
This  dimensionless  quantity  is  defined  as, 


Tornado  Index  = 
f  Tornadoes /Unit  Area"!  f      Rural  Population 


I      Rural  Population      J  I  Tornadoes/Unit  Area 

county  state 


(1) 


Geology  and  Geography 


30] 


ELKHART 

16 


4©2 


STEUQEN 

4©5 


Figure  1.     Tornadoes     affecting     Indiana     counties     (circled),     tornadoes     originating     in 
county  (underlined) ,  and  the  number  of  tornado  days  per  county  from  1916  through  1968. 


302 


Indiana  Academy  of  Science 


Figure  2.      Tornado    indices    per    Indiana    county    for    1916    through    1968. 


Geology  and  Geography  303 

The  tornado  index  for  each  county  is  therefore  based  on  a  common 
denominator  of  1.00,  the  state  average.  Counties  with  indices  greater 
than  1.00  are  above  the  state  average  and  those  less  than  1.00  are  below 
the  state  average.  The  use  of  rural  population  was  observed  to  more 
aptly  handle  the  problem  of  population  bias  than  total  population.  Fur- 
ther, it  seems  logical  that  a  given  area  should  only  be  inhabited  to  a 
certain  extent,  beyond  which  additional  population  would  not  improve 
the  chances  of  tornado  detection.  Rural  population  figures  were  also 
used  by  Darkow  in  his  Missouri  study,  supporting  the  approach  in  this 
investigation.  The  largest  tornado  index,  2.60,  was  computed  for  Porter 
County.  Other  high  values  include  Jasper  with  2.09  and  Pike  with  2.06. 
The  northwest  quadrant  and  the  southwest  corner  of  the  state  are  well 
above  the  state  average.  As  was  expected  the  southeast  portion  of  the 
state  is  considerably  below  the  state  average.  It  was  not  the  objective 
of  this  study  to  examine  in  detail  the  effect  of  topography  on  tornado 
distribution  but  these  data  are  indicative  of  terrain  effects  and  do 
suggest  further  study  into  the  problem.  It  should  be  mentioned,  how- 
ever, that  the  method  used  to  handle  the  population  bias  in  tornado 
reporting  may  not  be  the  most  accurate  and  the  tornado  indices  could 
be  slightly  misrepresentative. 

Figure  3  shows  the  yearly  distribution  of  tornadoes  and  tornadoes 
causing  injury  or  death.  Most  notable  is  the  increase  in  tornado  occur- 
ences since  1954.  This  can  be  partially  attributed  to  the  development  of 
the  severe  local  storm  (SELS)  forecasting  and  warning  program  by  the 
U.  S.  Weather  Bureau.  This  program  also  improved  and  updated  the 
efforts  in  the  tornado  recording  network  which  should  account  for  much 
of  this  increase.  Figure  3  is  also  indicative  of  more  tornadoes  causing 
injury  and  death  in  recent  years.  The  most  tornadoes  for  any  single 
year  was  the  52  recorded  in  1965.  Nineteen  of  these  caused  injury  or 
death.  However,  the  Palm  Sunday  tornado  outbreak  in  1965  did  make 
this  an  exceptionally  high  tornado  year  for  Indiana.  Also  shown  is  the 
yearly  distribution  of  tornado  days  and  tornado  death  days  for  Indiana. 
Again,  the  initiation  of  the  program  by  the  U.  S.  Weather  Bureau  is 
reflected  in  the  data. 

Figure  4  shows  the  monthly  distribution  of  tornadoes.  More  tor- 
nadoes occurred  in  April  than  any  other  month  and  78%  of  all  tornadoes 
occurred  during  the  period  March  through  July.  The  maximum  number  of 
tornado  days  occurred  in  June  but  tornado  days  in  early  spring  produced 
more  tornadoes.  For  instance,  45  tornado  days  produced  63  tornadoes 
in  July,  but  30  tornado  days  produced  71  tornadoes  in  March.  Another 
interesting  feature  is  that  a  larger  portion  of  the  tornadoes  occurring 
in  late  winter  and  early  spring  caused  injury  and /or  death.  Notice  that 
March  had  71  tornadoes,  30  of  which  caused  injury  and /or  death.  But, 
April  had  124  tornadoes  and  only  29  caused  injury  and/or  death.  This 
seems  to  indicate  that  the  tornado  season  must  be  in  progress  for  awhile 
before  people  begin  to  take  it  seriously.  The  data  also  suggest  a  small 
second  maximum  of  tornado  occurrences  in  early  fall.  This  is  associated 
with  increased  frontal  activity  which  is  not  as  violent  as  that  associated 
with  the  storms  that  develop  in  the  spring. 


304 


Indiana  Academy  of  Science 


Geology  and  Geography 


305 


130 

120 

110 

100 

90 

80 

70 

60 

• 

50 

40 

30 

20 

10 

•78% 


□   TORNADOES 
BS3   TORNADO  DAYS 
■i   TORNADOES  CAUSING 
INJURY  AND/OR   DEATH 


JFMAMJ      J      ASOND 


FiGURi:  4.     Indiana  tornado  occurrences  by  month  from  1916  through  1968. 


306 


Indiana  Academy  of  Science 


Table  1.  Indiana  tornado  deaths  and  tornado  death  days  by  month 
from  1916  through  1968. 

MJJASOND 
3   2   10   0   10   0 


J 

F 

M 

A 

Tornado 

Death  Days 

1 

0 

10 

4 

Tornado 

Deaths 

1 

0 

207 

157 

0   0 


0   0 


Table  1  gives  the  number  of  Indiana  tornado  deaths  and  tornado 
death  days  by  month  for  the  period.  More  deaths  have  occurred  in 
March  which  interestingly  enough  is  not  the  month  of  maximum  tornado 
activity. 


00     02     04     06    08      10     12      14      16      18     20     22     24 

Figure  5.     Diurnal    variation    of    Indiana    tornado    activity   from    1916    through    1968. 


The  diurnal  variation  of  tornado  activity  is  given  in  Figure  5. 
Tornadoes  can  occur  any  hour  of  the  day  just  as  they  could  any  day  of 
the  year,  but  late  afternoon  is  the  preferred  time.  More  tornadoes 
occurred  between  5:00  and  6:00  pm  than  any  other  hour  of  the  day.  A 
second  maximum  appears  between  midnight  and  1:00  am  which  is  prob- 
ably attributed  to  the  midwest's  nocturnal  thunderstorm  activity.  Also, 
tornadoes  causing  injuries  appear  to  be  less  in  proportion  to  the  num- 
ber of  tornadoes  for  nighttime  compared  to  daytime.  If  such  is  true  this 
could  be  related  to  the  fact  that  more  people  are  at  rest  or  asleep  in 
their  homes  and  are  therefore  not  quite  as  vulnerable  to  injury. 


Geology  and  Geography  307 

Table  2.  Diurnal  variations  of  Indiana  tornado  deaths  from 
1916  through  1968. 


00-01 

01-02 

02-03 

03-04 

04-05 

05-06 

06-07 

07-08 

08-09 

09-10 

10-11 

11-12 

0 

0 

3 

2 

0 

0 

1 

0 

0 

0 

0 

0 

12-13   13-14   14-15   15-16   16-17   17-18   18-19   19-20   20-21   21-22   22-23   23-24 
0      2     24     66     118    174     40     73     30      5      0      0 


Table  2  shows  the  diurnal  variation  of  tornado  deaths.  The  maxi- 
mum is  associated  with  peak  tornado  activity.  Again  it  is  evident  that 
nighttime  deaths  are  much  less  than  for  daytime  in  proportion  to  the 
number  of  tornadoes  occurring.  Since  death  producing  tornadoes  did 
occur  at  the  beginning  and  end  of  adjacent  hourly  periods,  the  tabular 
data  indicate  more  deaths  than  actually  happened  (Table  1). 

The  direction  of  movement  in  percentages  for  Indiana  tornadoes 
has  also  been  determined.  Over  81%  of  the  tornadoes  traveled  in  an 
east  through  northeast  direction  with  47%  moving  toward  the  north- 
east. Tornadoes  have  been  observed  to  move  in  all  directions  but  from 
southwest  to  northeast  is  the  general  tendency. 

Summary 

Yearly,  monthly,  and  diurnal  variations  of  tornado  frequencies, 
injuries,  and  deaths  for  Indiana  have  been  presented  based  upon  data 
assembled  for  the  period  1916  through  1968.  The  data  are  summarized 
in  Figures  1-5  and  Tables  1  and  2.  An  attempt  has  been  made  to  remove 
the  problem  of  population  bias  in  tornado  reporting.  Also,  the  effect  of 
topography  on  the  areal  distribution  of  Indiana  tornadoes  has  been 
touched  upon  and  further  study  is  intended  along  these  lines.  A  gen- 
eralized computer  program  has  been  developed,  and  is  available,  to  do 
similar  climatological  analyses  for  other  states. 

Acknowledgments 

I  am  grateful  to  Dr.  P.  J.  Smith  for  his  review  of  the  manuscript 
and  to  Mrs.  Carol  Gilliom  for  her  assistance  in  the  data  analysis  and 
computer  programming.  This  work  was  partially  supported  by  NASA 
Engineering  System  Design  Traineeship  Program  Project  No.  NGR  15- 
005-069. 

Literature  Cited 

1.  Flora,    Snowden    D.    1954.    Tornadoes   of   the   United    States.     University   of    Oklahoma 
Press,  Norman,  Okla.  194  p. 

2.  U.   S.   W.   B.   Technical  Note  No.   20,   1960:   Tornado  Occurrences   in   the  United   States. 
United  States  Weather  Bureau,  Washington,  D.  C. 

3.  Report   of   the    Chief   of   the   U.    S.    W.    B.,    1916-1931.    U.    S.    Weather    Bureau,    Wash- 
ington, D.  C. 

4.  U.    S.    Meteorological    Yearbook,    1931-1949.    U.    S.    Weather    Bureau,    Washington,    D.    C. 


308  Indiana  Academy  of  Science 

5.  Monthly    Weather    Review,     1921-1949.    United    States    Weather    Bureau,    Washington, 
D.  C. 

6.  National    Climatological    Data,    1950-1961.    U.    S.    Department    of    Commerce,    Weather 
Bureau,  Washington,  D.  C. 

7.  Indiana    Climatological    Data,    1950-1961.    U.    S.    Department    of    Commerce,    Weather 
Bureau,  Washington,  D.  C. 

8.  Storm  Data,   1959-1968.  U.  S.  Department  of  Commerce,  Weather  Bureau,  Washington, 
D.  C. 

9.  U.  S.  Bureau  of  the  Census,  1920,  1940,  1960.  Census  of  Population,  Washington,  U.   S. 
Government  Printing  Office. 


The  Changing  Location  Patterns  of  the  Neighborhood  Grocers  in 
Terre  Haute,  Indiana :  A  Geographic  Analysis 

R.  Michael  Dinkel  and  A.  J.  Cantin,  Indiana  State  University 


Abstract 

In  today's  corporate  society,  the  small  businessman  is  becoming  less  and  less 
common.  A  prime  example  of  this  presently  can  be  found  in  the  food  marketing 
industry.  Since  the  year  1900,  there  have  been  many  significant  changes  in  the  grocery 
business  throughout  the  United  States.  This  study  specifically  deals  with  the  causes 
and  effects  of  changing  location  patterns  of  the  small  grocer  in   Terre  Haute,   Indiana. 

The  location  of  the  small  grocer  with  respect  to  major  thoroughfares,  does  appear 
to  be  instrumental  in  the  survival  of  the  small  grocer  against  the  onslaught  of  various 
supermarket  chains.  Therefore  the  geographic  location  seems  to  be  a  major  influence  on 
the  survival  of  this  particular  type  of  small  business  establishment. 


Introduction 

The  changes  in  the  location  and  distribution  of  retail  grocers  within 
the  corporate  limits  of  Terre  Haute,  Indiana,  is  an  example  of  the 
changing  economic  environment  of  man.  This  study  was  undertaken 
during  the  spring  of  1969  when  both  authors,  in  personal  discussion, 
observed  the  rather  rapid  rate  at  which  certain  neighborhood  grocers 
were  being  eliminated,  due  perhaps  to  strong  competition  of  the 
numerous  supermarket  chains. 

We  questioned  whether  the  elimination  of  the  neighborhood  grocers 
was  due  to  the  location  of  supermarket  chain  stores  in  shopping  centers. 
Since  in  Terre  Haute  there  are  only  a  relative  few  such  centers,  it  could 
not,  by  itself,  account  for  the  change.  We,  therefore,  decided  to  test  his- 
torically, the  hypothesis  of  location  with  respect  to  major  thorough- 
fares. 

It  might  be  assumed  that  the  retail  grocer  would  tend  to  polarize 
around  the  major  thoroughfares  within  the  city.  This  was  found  to  be 
true  in  the  early  1900's  and  in  the  past  two  decades.  However,  in  the 
intervening  years,  a  negative  trend  was  apparent  with  regard  to  locat- 
ing on  major  thoroughfares. 

Data  for  this  study  were  gathered  from  the  Terre  Haute  City  Di- 
rectory for  the  years  1900  to  1967.  Recognizing  that  time  would  bring 
considerable  change  in  the  location  pattern,  we  decided  that  a  sample 
of  one  year  in  a  decade  would  demonstrate  such  change.  The  years 
selected  were  drawn  randomly,  and  if  data  for  that  year  were  lacking, 
the  next  year  was  arbitrarily  selected.  These  data  were  then  plotted 
on  a  city  map  (using  the  city  limits  for  the  year  selected).  Stores  out- 
side of  the  city  limits  were  not  included  in  the  study  and  were  not 
plotted.  For  purposes  of  finding  trends  in  recent  years,  data  for  both 
1962  and  1967  were  included  (Table  1). 

309 


310 


Indiana  Academy  of  Science 


Otiwwt* 


Fig 


URE  1.    The  location  of  retail  grocery  stores  in   Tcrre  Haute,  Indiana,  in  1! 


Geology  and  Geography  311 

Table  1.     Retail  grocers  on  major  traffic  arteries 


1900 

1908 

1918 

1927 

1937 

1944 

1954 

1962 

1967 

Wabash  Ave. 

27 

22 

28 

31 

12 

12 

11 

5 

3 

Lafayette    Ave. 

16 

14 

16 

34 

21 

17 

13 

8 

5 

Poplar   St. 

16 

18 

10 

12 

12 

9 

7 

6 

6 

Thirteenth    St. 

20 

20 

25 

27 

22 

17 

18 

10 

7 

Third  and 

Fourth   Sts. 

29 

27 

39 

56 

39 

26 

27 

13 

13 

Twenty-Fifth   St 

9 

7 

6 

Harding  Ave. 

21 

13 

14 

15 

10 

12 

— 

— 

— 

Analysis  of  Patterns,  1900-1967 

1900  Map 

In  1900,  location  with  respect  to  major  thoroughfares  was  ex- 
tremely important  (Fig.  1).  Four  major  concentrations  were  observable: 
along  Lafayette,  Wabash,  Poplar,  and  Third  and  Fourth  Streets.  Concen- 
trations were  present  on  Thirteenth  Street  and  on  Harding  Avenue  (Sec- 
ond Street),  which  formerly  was  the  major  north-south  route  in  the 
city.  The  heavy  concentration  on  Fourth  Street  was  an  enigma  until 
it  was  learned  of  a  Farmers  Market  in  the  Court  House  area.  Along 
Fourth  Street,  a  huge  "hay  rack"  had  been  installed  by  farmers  to 
mass  feed  their  horses  when  they  brought  in  their  produce  for  sale 
and  grocers  located  with  respect  to  this  potential  market. 

Located  along  all  streets  mentioned,  with  a  heavy  emphasis  in 
the  Central  Business  District  (CBD),  were  129  of  191  grocers  in  Terre 
Haute,  nearly  68  %  of  the  total.  Concentration  along  major  thorough- 
fares in  1900  was  extremely  important.  Note,  by  contrast,  the  sparse 
pattern  in  the  NE  and  SE  neighborhoods  of  Terre  Haute.  One  can  sum- 
marize the  city  for  1900  as  having  very  few  neighborhood  grocers. 

1908  Sample 

Vast  changes  occurred  between  1900  and  1908.  The  number  of 
neighborhood  grocers  increased,  particularly  in  the  NE  and  SE  sec- 
tions of  the  city.  Now  only  114  of  224  (or  about  50%  of  the  total)  were 
located  on  the  6  main  arteries  of  the  city — a  drop  of  18%  from  1900. 

As  the  city  grew  toward  the  NE  and  SE  and  construction  of  many 
homes  of  the  working  class  were  completed,  neighborhood  grocers  ap- 
peared within  these  areas. 

However,  the  major  distribution  of  stores  was  still  dominated  by 
location  with  respect  to  points  of  major  traffic  flow.  A  major  redistribu- 
tion was  taking  place.  The  heavy  concentration  near  the  "hay  rack" 
had  diminished,  and  the  Third  and  Fourth  Street  concentration  had 
spread  toward  the  city  limits.  These  two  streets  had  but  one  additional 
store   in    1908  as  compared   to   1900,   but  those   stores   pre-dating    1908 


312  Indiana  Academy  of  Science 

were  no  longer  concentrated  near  the  CBD  to  the  degree  they  were  in 
1900. 

Between  1900  and  1908  a  decline,  to  remain  unchecked  for  nearly 
half  a  century,  began  for  stores  located  along  major  thoroughfares, 
with  consequent  growth  in  importance  of  neighborhood  stores. 

1918  Sample 

By  1918  the  trend  toward  greater  numbers  of  neighborhood  grocers 
continued  with  a  drop  of  nearly  5%  of  the  thoroughfare-oriented  stores 
from  1908.  Note  the  relatively  dense  pattern  of  neighborhood  stores  in 
NE  and  SE  Terre  Haute. 

The  traffic-flow  locations  are  realigned,  with  growing  importance 
for  Thirteenth,  Wabash,  and  Third  and  Fourth  Streets,  plus  a  50%  loss 
of  the  grocers  located  on  Poplar.  Reappearing  is  a  very  heavy  con- 
centration on  Third  and  Fourth  Streets  between  Ohio  and  Chestnut 
Streets.  These  stores  were  largely  small  stores  owned  by  newly  arrived 
Syrian  immigrants  in  a  low-income  area  of  the  city. 

Between  College  and  Hulman  and  Twelfth  and  Fourteenth  Streets, 
a  Negro  district  referred  to  as  "Baghdad,"  had  nearly  a  250%  increase 
from  7  to  18  stores.  Again,  as  in  the  Third  and  Fourth  Street  complex, 
this  illustrates  a  growing  tendency  for  large  numbers  of  small  stores 
to  locate  in  low-income  areas. 

1927  Map 

This  was  the  peak  year  for  grocers  in  Terre  Haute,  as  there  were 
489  stores  located  in  the  city,  up  from  295  in  1918  (Fig.  2).  A  large  per- 
centage of  this  was  due  to  the  36-store  Oakley  Chain  that  was  opened 
during  this  period,  and  years  later,  sold  out  to  Kroger  Supermarkets,  Inc. 
These  stores  were  spaced  evenly  throughout  the  entire  city  and  were 
supplied,  in  part,  through  Oakley's  farm. 

This  locational  trend  continued.  The  major  traffic  arteries  accounted 
for  only  40%  of  the  stores,  despite  a  noticeable  growth  of  stores  in  the 
Third  and  Fourth  Street  area  and  Lafayette  Avenue  which  increased 
from  16  stores  to  34. 

Major  traffic  arteries  had  a  higher  number  of  stores  than  ever 
before,  but  a  smaller  percentage  of  the  total.  Store  coverage  was  most 
uniform  in  the  68  year  period.  More  stores  were  located  in  neighbor- 
hoods, except  in  the  "Baghdad"  area  where  the  number  of  stores  fell 
from  18  to  7,  the  number  originally  found  in  1908. 

1937  Sample 

The  effect  of  the  depression  is  demonstrated  by  the  rapid  decline  in 
grocer  numbers,  from  439  stores  to  338,  or  a  23^  decrease.  In  1937 
only  116  of  338  stores  (or  34%)  were  located  on  major  traffic  routes, 
down  6%  from  the  previous  decade. 


Geology  and  Geography 


313 


Figure  2.    The  location  of  retail  grocery  stores  in  Terre  Haute,  Indiana,  in  1927. 


The  remaining  stores  had  even  distribution,  with  only  a  few  cluster- 
ings. A  new  cluster  in  the  Twelve  Points  area,  which  began  in  1927, 
continued  through  1937,  due  perhaps  to  the  rebuilding  of  Garfield  High 
School  and  a  resultant  increase  of  residences  in  north  and  northeast 
Terre  Haute. 


314 


Indiana  Academy  of  Science 


FIGURE  3.    The  location  of  retail  grocery  stores  in  Terre  Haute,  Indiana,  in  195U- 


Of  the  101  stores  that  failed  from  1927  to  1937,  59  of  these  were 
located  on  major  thoroughfares,  indicating  the  declining  importance 
of  this  location  factor. 


Geology  and  Geography 


315 


Figure  4.    The  location  of  retail  grocery  stores  in  Terre  Haute,  Indiana,  in  1967. 


1944  Sample 

The  number  of  grocers  declined  from  338  to  276,  a  drop  of  62 
stores.  However,  an  important  trend  change  occurred  at  this  time.  Al- 
though only  93  stores  of  the  276  were  located  on  traffic-flow  routes    the 


316  Indiana  Academy  of  Science 

percentage  of  stores  so  located  remained  at  347c,  the  same  figure  as 
for  1937.  This  marks  the  first  time  the  decrease  was  stayed.  Stores  lo- 
cated on  traffic-flow  had  fewer  failures  than  neighborhood  stores,  indi- 
cating a  trend  away  from  the  neighborhood  stores  for  the  first  time. 
No  particular  section  of  the  city  had  a  disproportional  share  of  grocer 
failure. 

1954  Map 

By  this  date  new  thoroughfare  alignments  had  been  made  in  Terre 
Haute  (Fig.  3).  Harding  Avenue  (Second  Street)  was  no  longer  a  major 
north-south  route,  but  was  replaced  by  Seventh,  Eighth,  and  Ninth 
Streets.    Also,    Twenty-fifth    Street    had    grown    to    major    status. 

The  percentage  of  stores  on  major  traffic  arteries  had  grown  to 
41%,  up  considerably  from  1944.  This  marks  the  first  time  in  50  years 
that  the  pattern  had  definitely  reversed,  the  gain  for  major  traffic 
locations  came  from  a  net  loss  for  neighborhood   stores. 

1962  Sample 

The  rate  of  decline  was  now  accelerating  as  there  were  but  161 
grocers  left  in  Terre  Haute,  a  drop  of  nearly  100  stores  in  an  8-year 
period.  Of  those  remaining,  68  or  43%  were  on  major  traffic  locations, 
up  2%  since  1954.  This  indicates  the  weakening  position  of  the  neigh- 
borhood grocer  and  that  traffic-flow  and  shopping  centers  (as  a  location 
factor)  were  making  inroads. 

1967  Map 

All  trends  continued,  but  now  at  an  accelerated  rate  (Fig.  4).  Of  the 
107  remaining  stores,  53  (or  about  50%)  are  located  with  respect  to 
major  traffic,  up  7%  in  5  years.  In  addition,  at  least  six  major  super- 
markets were  established  in  shopping  centers,  leaving  relatively  few 
neighborhood  grocers  in  Terre  Haute. 

The  net  number  of  stores  in  the  city  is  declining  rapidly.  Obviously 
the  supermarket  era  which  began  after  World  War  II  is  now  establish- 
ing itself,  and  larger  stores  have  spread  farther  apart  to  areas  of  easy 
automobile  access.  The  decline  of  the  neighborhood  grocer  is  apparent 
both  in  relative  importance  and  in  total  numbers. 

Conclusion 

It  is  now  apparent  that  there  are  many  reasons  for  small  grocers 
locating  where  they  do  in  Terre  Haute,  but  location  within  a  neighbor- 
hood is  no  longer  significant.  Location  with  respect  to  shopping  centers 
is  a  growing  factor;  nonetheless,  the  most  important  factor  at  present 
is  major  avenues  of  transportation. 

The  future  of  the  neighborhood  grocer  in  the  city  appears  to  be 
rather  precarious.  Although  the  total  volume  of  each  retail  store  was 
unattainable,   that   of  the    small    neighborhood   grocer   doubtless    is   very 


Geology  and  Geography  317 

low.  The  authors  estimated,  by  using  a  small  sample  of  shoppers,  that 
the  major  chain  stores  plus  allied  independent  grocers  (i.e.,  IGA)  con- 
trol in  excess  of  90  (/<  of  the  total  retail  volume. 

It  would  appear  that  the  average  consumer,  due  to  the  higher  prices 
charged  by  the  neighborhood  grocer,  will  most  often  patronize  the 
larger  chain  for  purely  economic  reasons.  The  convenience  of  a  neigh- 
borhood store  appears,  therefore,  to  remain  secondary  to  the  savings 
effected  by  shopping  at  a  larger  volume   establishment. 


Literature  Cited 

1.  Dinga,   Carl  F.    1968.   Analysis  of  retail  site   locations   in   Terre   Haute,    Indiana.    Proc. 
Indiana  Acad.  Sci.  77:321-25. 

2.  Guernsey,    Lee.    1961.    Characteristics    of    the    Terre    Haute    Central    Business    District. 
Proc.  Indiana  Acad.  Sci.  71  :203-09. 

3.  Terre  Haute,  Indiana  City  Directories,  1900-1967.  R.  L.  Polk  Publishing  Company. 


Population  and  Settlement  Decline  in  Southwestern  Indiana 

Thomas  Frank  Barton,  Indiana  University 

Abstract 

The  largest  contiguous  and  historically  continuous  area  of  population  and  settlement 
decline  in  Indiana  is  found  in  the  southwestern  part  of  the  state.  This  mobile  and 
declining  population  influences  many  facets  of  the  economy  and  life  style.  Reasons  for, 
and  potentialities  of,  a  Southwestern  Indiana  Regional  Planning  Commission  are  pre- 
sented. Population  shifts  are  not  a  disaster  if  proper  short-  and  long-ranged  plans  are 
adopted  to  readjust  to  the  situation. 

Introduction 

The  largest  continuous  area  of  population  and  settlement  decline 
in  Indiana  is  found  in  the  southwestern  part  of  the  state.  Except  for 
Vermillion  County,  there  are  17  counties  located  in  the  crotch  of  the 
Wabash  and  Ohio  Rivers  in  which  populations  declined  between  1950- 
1960,  and  4  counties  which  gained  population  only  because  births  ex- 
ceeded the  combined  number  of  deaths  and  emigration.  The  counties 
designated  in  this  paper  as  southwestern  Indiana  are  listed  in  Table  1. 
Population  mobility  is  characteristic  of  this  area  with  some  people 
moving  from:  1)  one  county  in  the  area  into  another  in  the  area,  2) 
the  countryside  into  the  settlements,  3)  smaller  settlements  to  the 
largest  cities,  and  4)  from  the  area.  This  moving  and  declining  popu- 
lation influences  many  facets  of  the  society,  such  as:  taxation  and 
debts  on  formerly  fully-used  public  and  private  facilities;  property 
values;  need  and  support  for  commercial  establishments  and  services; 
land-use  requirements;  and  future  economic  development.  Population 
shifts  in  a  state  are  not  a  disaster  if  proper  short-ranged  and  long- 
ranged  plans  are  adopted  to  re-adjust   to  the  situation. 

County  Population  Decline 

During  1950-1960,  21  Indiana  counties  decreased  in  population  and 
17,  or  over  4/5  of  these,  are  located  in  southwestern  Indiana.  The  four 
counties  outside  of  this  region  that  declined  in  population  are  Warren, 
adjacent  to  the  Indiana-Illinois  boundary  and  adjacent  to  this  region 
on  the  north,  and  Union,  Ohio  and  Switzerland  counties,  adjacent  to 
the    Indiana-Ohio    boundary    in    southeastern   Indiana. 

Not  only  did  counties  in  this  region  decline  in  population  between 
1950-1960  but  also  the  number  of  counties  with  such  declines  steadily  in- 
creased in  the  past  30  years,  1930-1960  (Table  1).  During  the  1930-1940 
decade,  14  counties  increased  in  population  and  only  7  decreased;  the 
highest  percentage  loss  was  6.2  in  Vermillion  County.  However,  dur- 
ing   the     decade     of     1940-1950     when     employment     opportunities     and 


Statistics  in  this  paper  preceding  the  centerhead  "Some  Chain  Reactions"  are  taken 
from  the  1930,  1940,  1950  and  1960  Census  of  Population  of  the  United  States  or  secured 
by  using  the  data  from  these  volumes. 

318 


Geology  and  Geography 


319 


Table  1.     Population    percentage     changes    in    Southwestern    Indiana. 


County- 


Estimated 
1930-1940     1940-1950     1950-1960     1966-1985 


Clay 

Crawford 

Daviess 

Dubois 

Gibson 

Greene 

Knox 

Lawrence 

Martin 

Orange 

Owen 

Parke 

Perry 

Pike 

Posey 

Spencer 

Sullivan 

Vanderburgh 

Vermillion 

Vigo 

Warrick 


. — ■ 

4.2 

— 

5.7 

+ 

1.2 

—     4.1 

+ 

0.1 

— 

8.7 

— 

9.8 

—  25.0 

+ 

L.3 

+ 

2.3 

— 

0.5 

—  15.3 

+ 

9.9 

+ 

5.3 

+ 

15.5 

+    20.6 

+ 

5.2 

+ 

* 

— 

2.5 

—  10.3 

— 

0.5 

— 

11.0 

— 

5.6 

—  25.9 

+ 

0.4 

■ — 

1.3 

— 

4.3 

—  21.4 

— 

1.5 

— 

2.0 

— 

6.5 

—     5.2 

r 

1.9 

+ 

3.7 

— 

0.7 

—     9.0 

— 

0.8 

— 

2.5 

— 

* 

—     6.2 

\ 

6.5 

— 

2.7 

— 

3.1 

—  16.6 

+ 

4.8 

— 

9.7 

■ — 

5.6 

—  14.2 

+ 

6.9 

— 

2.3 

— 

0.8 

-  16.6 

f- 

4.2 

— 

12.0 

— 

14.7 

—  33.3 

+ 

7,1 

+ 

3.3 

— 

3.0 

—  23.8 

— 

3.0 

— 

0.2 

— 

0.6 

—  17.6 

— 

4.0 

— 

12.4 

— 

8.2 

—  23.8 

f 

15.4 

+ 

22.7 

+ 

3.3 

+    29.0 

— 

6.2 

— 

9.5 

— 

10.3 

—  18.7 

+ 

0.9 

+ 

5.5 

+ 

3.1 

+      3.6 

+ 

6.6 

+ 

10.8 

+ 

9.5 

+    12.0 

Change  too  small  to  compute  percentage. 


employment  surpassed  the  previous  decade,  13  counties  declined  in 
population,  7  gained  and  Orange  County  had  a  small  net  loss.  During 
1950-1960  only  5  counties  gained  in  population  and  3  of  these  only 
slightly— Clay  1.2%,  Vigo  3.1%  and  Vanderburgh  3.0%. 

Although  11  counties  in  this  region  had  both  gains  and  declines 
during  the  30-year  period  from  1930-1960,  10  counties  experienced 
consistent  declines  or  gains.  Four  counties  (Dubois,  Vanderburgh,  Vigo 
and  Warrick)  registered  continuous  gains  and  six  counties  (Greene, 
Lawrence,  Orange,  Spencer,  Sullivan  and  Vermillion)  had  continuous 
losses.  The  six  declining  counties  are  located  adjacent  to  the  four 
gaining  counties.  Two  of  the  four  counties  with  continuous  population 
growth,  Vanderburgh  and  Vigo,  contain  the  two  largest  cities  in  this 
area,  Evansville  and  Terre  Haute,  respectively.  Three  counties  with 
continuous  gains  (Vanderburgh,  Warrick  and  Dubois)  are  adjoining, 
but  the  three-county  cluster  is  surrounded  by  Indiana  counties  where 
one  or  two  decades  of  population  decline  has  taken  place. 

The  severity  of  population  decline  was  greater  during  the  more 
prosperous  decades  of  1940-1960  than  during  depression  years  of  the 
1930's.    The   population   declines   during   the    1930-1940   period   in   the    7 


320  Indiana  Academy  of  Science 

counties  ranged  between  1.5  and  6.2%.  In  contrast,  the  population  de- 
clines during  the  decade  of  1940-1950  in  13  counties  ranged  between 
1.3  and  12.0 %  with  6  counties  having  declines  greater  than  6.2%.  Dur- 
ing the  1950-1960  decade  the  decline  in  16  counties  ranged  between  0.7 
and  14.7%. 

Population  Growth  in  the  Region 

The  21  counties  in  this  region  have  had  a  small  population  gain 
during  1940-1960.  The  gain  was  2.8%  between  1940-1950  and  dropped 
to  about  0.5%  during  the  1950's. 

This  region  is  characterized  by  emigration.  None  of  the  21  counties 
escaped  emigration.  Any  gain  in  population  was  chiefly  due  to  births 
outnumbering  the  total  deaths  and  emigration.  Not  one  of  these  coun- 
ties registered  an  increase  of  18.5%,  which  was  both  the  state  and  na- 
tional growth  rate  during  the  1950-1960  decade.  In  contrast,  22  counties 
in  Indiana  had  a  population  growth  greater  than  18.5%  with  increases 
ranging  between  19.7%  in  Tippecanoe  and  66.3%    in  Hendricks  counties. 

Growth  in  Settlements 

Settlements  with  over  1,000  inhabitants  increased  in  15  counties 
and  decreased  in  4  during  1930-1960.  The  increase  ranges  between 
3.3%  in  Spencer  County  to  39.7%  in  Parke  County,  and  the  rate  of 
decrease  between  3.0%  in  Knox  County  and  23.9%  in  Vermillion  County. 
Crawford  County  does  not  have  a  settlement  with  1,000  inhabitants,  and 
in  Sullivan  County,  settlements  with  over  1,000  decreased  by  7  persons. 

During  this  30-year  period  the  number  of  settlements  with  popu- 
lations over  1,000  increased  from  41  to  46.  Six  of  these  46  did  not  have 
a  population  of  1,000  in  1930.  And,  only  one  settlement  with  1,000  or 
more   inhabitants   in   1930   dropped  below  that  level   before   1960. 

While  it  is  true  that  the  population  in  settlements  of  over  1,000 
has  increased  in  15  counties  during  1930-1960,  18  of  these  were  smaller 
in  1960  than  they  were  in  1950  (Table  2).  Moreover,  12  of  these  18  were 
cities  with  a  minimum  population  of  2,000  and  the  largest,  Vincennes, 
had  18,046  in  1960.  The  18  settlements  with  population  declines  totaled 
78,312  in  1960,  as  compared  with  83,605  in  1950,  or  a  loss  of  5,293. 
This  loss  is  relatively  small  when  compared  with  the  gain  of  12,907 
persons  by  Evansville  in  the  same  decade.  At  the  same  time,  3  of  the 
small  cities  lost  over  10%  of  their  inhabitants:  Oakland  City,  14.8%, 
Jasonville  17.0%  and  Bicknell  37.0%.  During  the  preceding  decade, 
1940-1950,  Oakland  City's  population  gained  while  Bieknell's  declined 
by  10.5%,  and  Jasonville  lost  14.1%   (2). 

More  settlements  of  1,000  or  more  declined  in  population  during 
1940-1950  than  during  the  following  decade.  There  were  41  settlements 
of  this  size  in  1950;  20  of  these  had  a  population  decline  during  the 
1940's. 

The  cities  with  populations  of  over  5,000  registered  a  smaller  de- 
cline during  1950-1960  than  cities  of  less  than  5,000.   Of  the   12  cities 


Geology  and  Geography 


321 


Table  2.   Population  trends  in  county  seats  and /or  largest  cities. 


County 


County  Seat  and/or 
Largest  Cities 


1930- 
1940 


1940-   1950-  Population  greater  in  1960 
1950  1960    than  any  other  decade 


Clay 

Brazil 

Crawford 

English 

Daviess 

Washington 

Dubois 

Jasper 

Gibson 

Princeton 

Greene 

Bloomfield* 

Linton 

Knox 

Vincennes 

Lawrence 

Bedford 

Martin 

Loogootee* 

Shoals 

Orange 

Paoli 

Owen 

Spencer 

Parke 

Rockville 

Perry 

Cannelton 

Tell  City* 

Pike 

Petersburg 

Posey 

Mount  Vernon 

Spencer 

Rockport 

Sullivan 

Sullivan 

Vanderburgh 

Evansville 

Vermillion 

Clinton* 

Newport 

Vigo 

Terre  Haute 

Warrick 

Boonville 

+ 
+  + 
+ 
+ 
+ 


+ 

+ 
+      + 


1 


*  Largest  city  in  county  but  not  the  county  seat. 

with  over  5,000,  7  had  a  larger  population  in  1960  than  in  1950,  and  5 
had  less.  The  two  largest  cities,  Evansville  and  Terre  Haute,  with  over 
70,000,  gained  in  population  during  the  1950's  while  the  next  2  largest 
cities,  Vincennes  and  Bedford,  lost. 


Summary  of  Population  Decline 

Counties,  towns  and  cities  have  lost  population  in  Southwestern 
Indiana.  Some  of  these  declines  existed  during  only  1  decade,  some 
during  2  and  in  others,  declines  were  continuous  over  30  years,  between 
1930-1960.  Projections  of  population  growth  between  1966  and  1985  pre- 
dict that  17  of  these  21  counties  will  decline  by  approximately  57,000 
and  4  counties,  Dubois,  Vanderburgh,  Vigo  and  Warrick,  will  gain  about 
61,000  inhabitants  during  the  same  period  (4).  The  total  gain  would  be 
only  4,000,  a  very  small  gain  (less  than  1%).  Should  the  projection  be 
correct,  since  the  total  population  gain  was  only  0.5%  during  1950- 
1960,  it  is  obvious  that  this  region  has  problems  with  population 
declines,  almost  stagnant  growth  and  emigration.  In  contrast,  other  coun- 
ties in  Indiana  will  rapidly  gain  in  population  due  to  births  and  immi- 
gration. For  example,  according  to  John  M.  Huie,  Monroe  County's 
population  increased  by  35.5%  between  April  1,  1960,  and  July  1,  1966, 
and  immigration  of  more  than  14,000  was  the  highest  of  any  county  in 
the  state.  During  the  same  period  Vanderburgh's  emigration  reached 
11,800. 


322  Indiana  Academy  of  Science 

Some  Chain  Reactions 

As  the  population  of  an  area  declines  over  a  period  of  several 
decades  a  chain  of  economic,  social  and  political  reactions  bring  about 
complications.  Some  of  these  interrelated  reactions  are  briefly  described 
in  the  following  paragraphs. 

1)  Local  employment  opportunities  either  stagnate  or  decline. 
In  Southwestern  Indiana  employment  in  mining,  agriculture,  com- 
merce and  services  has  declined  in  recent  decades.  Technological  change 
and  new  techniques  have  influenced  need  for  employment  in  all  these 
occupations.  New  machinery  and  techniques  in  coal  mining  have  dras- 
tically cut  the  need  for  a  large  number  of  miners.  Machinery,  farm 
consolidation  and  corporate  farming  have  resulted  in  rapid  declines  in 
population  and  farm  employment  opportunities.  With  decline  of  mining 
and  farming  employment,  fewer  people  are  needed  to  supply  goods  and 
services  resulting  in  some  becoming  temporarily  or  permanently  un- 
employed. As  work  opportunities  appeared  in  adjacent  counties  or  in 
nearby  cities  people  were  forced  either  to  commute  to  work  or  to  move. 

2)  Areas  with  declining  populations  generally  have  a  larger  num- 
ber of  their  people  in  the  upper  age  brackets.  Emigration  generally  con- 
sists of  the  younger  workers  and  those  more  economically  "foot  loose." 
And  the  older  people  who  are:  unable  to  liquidate  their  property;  will- 
ing to  accept  less  affluent  living  conditions;  and /or  incapable  of  leaving 
life-long  friends,  remain  behind. 

3)  Employees  who  remain  in  the  area  spend  more  time  and  money 
in  commuting  to  work  in  cities  such  as  Evansville,  Terre  Haute,  and 
Vincennes  and  to  factories  in  Dubois  County  or  in  counties  outside  the 
area  such  as  Monroe. 

4)  As  the  population  declines  in  townships,  counties,  incorporated 
villages  and  small  cities,  school  districts  and  special  districts,  it  often 
becomes  more  expensive  per  person  or  family  to  support  the  existing 
services  and  to  provide  new  facilities.  Moreover,  as  commercial  and/or 
small  industrial  establishments  go  out  of  business  or  move  from  the 
area,  heavier  taxes  fall  upon  residential  and  agricultural  land  and 
structures. 

5)  In  attempting  to  reduce  costs,  new  services  (typical  of  more 
prosperous  counties  and  cities)  are  often  postponed  and  the  quality  of 
the  service  already  established  may  decline.  For  example,  the  size  of 
police  and  fireman  forces  may  be  cut,  new  equipment  may  not  be  pur- 
chased and  that  on  hand  may  not  be  adequately  repaired. 

6)  With  the  resulting  population  decline,  public  services  retarded 
and /or  restricted,  and  tax  rates  increased,  it  becomes  very  difficult  for 
settlements  to  attract  manufacturing  and  service  industries  which  could 
give  the  community  a  more  varied  employment  mix.  Industrial  manage- 
ment makes  careful  investigations  of  the  quality  and  quantity  of  serv- 
ices available  before  moving  into  a  city. 


Geology  and  Geography  323 

7)  Residential  property  values  may  decline  in  villages  and  small 
cities  thus  putting  owners  with  large  debts  in  a  financial  squeeze  and 
preventing  people,  if  they  must  sell,  from  receiving  a  reasonable  price 
for  their  land  and  buildings. 

8)  The  number  and  quality  of  commercial  and  service  establish- 
ments in  small  cities  may  decline  which  may:  reduce  customer  selection; 
raise  prices;  and /or  influence  customers  to  shop  in  other  cities. 

9)  The  number,  kinds  and  quality  of  professional  services  may 
also  decline  and  some  may  disappear.  Some  villages  and  smaller  cities 
which  formerly  had  several  doctors,  dentists,  lawyers  and  bankers  now 
have  only  one  or  none  of  each  profession. 

10)  As  cities  become  dormitory  settlements  where  most  of  the 
taxes  are  levied  on  residences,  the  taxing  unit  cannot  afford  to  provide 
the  services  considered  necessities  in  a  twentieth  century  community 
or,  if  it  does,  it  becomes  a  burden. 

11)  Vacant  urban  structures  and  land  provide  a  poor  economic 
image  to  potential  developers. 

12)  Commuters  who  live  outside  the  corporate  limits  may  be  sub- 
ject to  payroll  taxes.  Mayor  Frank  F.  McDonald,  Evansville,  estimates 
that  15,000  persons  who  work  in  Evansville  live  outside  the  political 
city  (1). 

As  these  and  other  actors  interact  the  economic  situation  spirals 
downward  aggravating  forces  already  in  operation,  sometimes  intro- 
ducing other  degrading  forces,  and  stimulating  additional  emigrations 
to  more  prosperous  areas  in  Indiana  and  other  states.  According  to 
Huie  (4),  Indiana's  employment  grew  by  approximately  206,000  jobs 
between  1950-1960  but,  according  to  the  United  States  Department  of 
Commerce,  if  Indiana's  employment  had  grown  at  the  national  rate  there 
would  have  been  an  additional  29,000  jobs. 

Some  Suggested  Action 

What  has  been  done  in  the  last  decade  or  two  to  help  the  county 
leaders  and  people  to  meet  the  problems  associated  with  declining 
populations?  Have  conferences  been  held,  study  groups  organized  and/ 
or  a  program  of  investigations  been  launched?  Is  there  literature  on 
these  problems  that  could  be  made  available  to  the  high  schools  and 
universities  in  this  area?  Does  the  Legislature  or  the  State  have  study 
groups  for  research  and  to  offer  suggestions  and  alternate  courses  for 
action?  ,..;.._.,.,_ 

It  seems  that  not  too  much  has  been  written  either  about  the  21 
counties  under  consideration  or  the  problems  associated  with  declining 
populations  and  settlements.  And  one  of  the  best  publications  a  propos 
to  this  area,  entitled  Regional  Development  and  the  Wabash  Basin  (3), 
either  has  not  been  read  or  understood  or,  if  so,  has  not  stimulated  much 
action  or  reaction  in  Southwestern  Indiana. 


324  Indiana  Academy  of  Science 

It  seems  that  most  of  the  American  literature  on  settlements,  urban- 
ism  and  urban  problems  and  solutions  to  problems  is  about  cities  with 
populations  over  100,000.  Moreover,  the  literature  on  problems  created 
by  rapid  population  and  city  growth  far  exceeds  that  on  problems  and 
alternate  solutions  for  areas  of  population  and  settlement  decline.  There 
is  need  for  material  written  on  smaller  settlements.  Southwestern 
Indiana  is  a  region  of  small  settlements  with  only  5  of  the  46  having 
populations  of  10,000  or  more.  Twenty  of  the  46  have  populations  in  the 
1,000  to  2,000  bracket.  And  in  Indiana  where  a  settlement  qualifies  as 
a  city  if  it  has  a  population  of  2,000,  there  are  26  cities  with  14  in  the 
2,000-5,000  bracket  and  7  in  the  5,000-10,000  bracket. 

The  government  provides  information  on  surveys  and  studies  deal- 
ing with  the  Wabash  and  Ohio  River  watersheds  and  has  made  or  will 
make  suggestions  for  economic  and  social  development.  But  most  of 
these  studies  focus  on  areas  of  both  rapidly-expanding  and  declining 
counties. 

Is  it  possible  for  the  21  counties  in  this  area  and  Warren  (adjacent 
and  to  the  north)  to  create  a  regional  planning  organization  which  could 
focus  directly  on  the  problems,  potentialities  and  beneficial  adjustments 
for  this  area  ?  Such  an  organization  could  provide  a  forum  wherein  the 
problems  of  the  area  could  be  aired,  freely  discussed  and  courses  of 
action  agreed  upon  in  a  non-partisan  atmosphere.  Conferences  could  be 
held  for  high  school  students  and  their  parents,  university  audiences, 
and  in  settlements  throughout  the  region  to  discuss  economic  and  so- 
cial problems  and  alternative  solutions.  A  Southwestern  Indiana  Re- 
gional Planning  Commission  could  outline  and  start  a  program  of 
study  investigations.  It  would  seem  that  such  a  program  would  be 
better  than  the  present  situation.  This  is  serious  and  action  is  needed 
now. 

Literature  Cited 

1.  Anonymous.    1969.    Politics    in    perspective:    eminent    mayors    favor    payroll    tax.    The 
Indianapolis  Star,  Section  2,  September  28,  p.  8. 

2.  Barton,    Thomas    Frank.     1952.    Cities    with    a    population    decline    in    southwestern 
Indiana,  1940-1950.  Proc.  Indiana  Acad.  Sci.  62  :250-255. 

3.  Boyce,    Ronald  R.    (Ed.)    1964.   Regional   development   and  the   Wabash   basin.   Univer- 
sity of  Illinois  Press,  Urbana. 

4.  Huie,    John   M.    1968.    Population    changes    pressure    local    governments.    Economic    and 
Marketing  Information  for  Indiana  Farmers.  December  31.  p.  2-4. 


A  Comparison  of  the  Central  Place  Hierarchy  Pattern  of 
Central  Indiana  to  the  Walter  Christaller  Model 

Neil  V.  Weber,  Indiana  State  University 


Abstract 

Walter  Christaller's  theory  on  the  hierarchy  of  central  places  was  used  as  a  model 
for  the  analysis  of  the  distribution  of  agglomerated  centers  in  central  Indiana.  The  132 
study  centers  were  analyzed  for  potential  hierarchy  breaks  relating  to  the  degree  of 
centrality  of  the  individual  centers.  The  t  test  method  was  used  to  check  whether  or  not 
significant  differences  existed  between  the  spacing  of  Indiana  centers  and  the  theoretical 
Christaller  pattern.  Finally,  the  Near-Neighbor  Analysis  was  used  to  test  whether  or  not 
the  distributional  pattern  of  the  study  centers  conforms  to  the  maximum  dispersal  of  the 
hexagonal  framework.   Conclusions  drawn  from  the  study  affirm  the  hypotheses  examined. 

The  purpose  of  this  study  was  to  empirically  analyze  the  basic 
principles  of  the  central  place  theory.  The  problem  is  focused  on  the 
interpretation  of  the  spatial  relationships  which  are  found  among  the 
central  places  on  the  Tipton  Till  Plain  of  central  Indiana — namely  the 
size,  spacing  and  distribution  of  central  places  within  the  study  area. 

The  method  of  analysis  was  concerned  with  testing  two  primary 
hypotheses.  The  first  being  that  certain  ordering  principles  of  settle- 
ment exist  in  nature;  second,  that  there  is  a  distinct  relationship  be- 
tween the  organization  and  distribution  of  agglomerated  centers  in 
central  Indiana  and  the  Walter  Christaller  pattern  for  southern  Ger- 
many. 

Delimitation  of  Study  Area 

The  study  area  consists  of  the  52-county,  20,362  square  mile  central 
portion  of  Indiana.  These  counties  reflect  homogeneous  unity  in  that 
they  are  all  directly  associated  with  the  physiographic  sub-province  of 
Indiana  known  as  the  Tipton  Till  Plain.  All  counties  either  border  on  or 
lie  within  this  physiographic  unit. 

The  Tipton  Till  Plain  is  a  nearly  flat  to  gently  rolling  glaciated 
surface  (6)  with  virtually  featureless  topographic  expression  (relief 
seldom  greater  than  50-100  feet).  The  monotony  of  the  flat-lying  land- 
scape is  evident  throughout  the  central  portion  of  the  state  with  only 
the  extreme  southeastern  section  (the  gradational  boundary  zone)  re- 
flecting any  topography  related  to  the  underlying  bedrock   (8). 

The  Tipton  Till  Plain  exemplifies  a  homogeneity  of  economic  ac- 
tivity as  well  as  topography.  It  is  an  agricultural  region  (4)  contain- 
ing three  predominant  types  of  farming:  cash-grain  in  the  west,  grain- 
livestock   in   the   central   portion,   and   general   farming  in   the   east. 

Some  portions  of  the  study  area  deviate  from  the  normal  pattern 
described  in  this  section.  These  particular  areas  will  be  discussed  in 
detail  later  in  the  study.  The  vast  majority  of  central  Indiana,  how- 
ever, is  a  flat-lying  agricultural  region. 

325 


32(5 


Indiana  Academy  of  Science 


This  study  is  concerned  with  the  distributional  pattern  of  132  ag- 
glomerated settlements  within  the  study  area.  These  settlements  repre- 
sent all  centers  within  the  region  having  populations  of  1,000  or  more. 
The  cut-off  limit  was  set  at  1,000  due  to  the  problem  of  acquiring 
comprehensive  data  for  centers  below  this  figure. 

Evaluation  of  the  Hierarchy  Pattern  for  Indiana  Centers 

Population  is  a  common  measurement  of  the  importance  of  place 
(3).  It  may,  therefore,  serve  as  a  major  criterion  for  determining  the 


,000,000- 


First  order  canter 


Second    order   centers 


Third   order   centers 


Fourth    order  centers 


*•*. 


'•000t — i 1 1 1 1 1 1 1 1 1 1 r 

0  10        20         30        40         50         60         70        80         90         100        110        120       130        140 

RANK 


Figure   1.     Rank-size  relationship  of  urban  centers  in  Central  Indiana. 


Geology  and  Geography  327 

functional  grade  of  trade  centers.  Population  gaps  in  the  progression 
from  small  to  large  centers  would  reflect  major  distinctions  in  the  size 
groups  (1).  Theoretically,  a  hierarchy  of  centers  would  develop  re- 
lating to  the  degree  of  centrality  of  the  groupings. 

By  plotting  the  132  centers  on  a  semi-logarithmic  graph  according 
to  their  rank-size  value,  distinct  groupings  within  the  series  are  ob- 
served (Fig.  1).  The  progression  from  largest  to  smallest  is  geo- 
metrically constant  from  80,000  to  30,000;  23,000  to  11,000;  and  9,600 
to  1,000.  Distinct  gaps  in  the  trend  line  are  evident  between  400,000  to 
80,000;  30,000  to  23,000;  and  11,000  to  9,600.  This  pattern  would  imply 
four  well  developed  hierarchy  groupings  for  the  central  places  dis- 
tributed within  the  study  area. 

The  first  order  or  primary  group  descends  to  approximately  the 
100,000  level.  This  would  include  only  one  center,  that  of  Indianapolis. 
The  secondary  grouping  ranges  from  80,000  to  30,000.  This  grouping 
contains  8  cities.  There  are  13  tertiary  centers  ranging  from  23,000  to 
11,000;  and  110  centers  of  quaternary  level  with  a  population  of  9,000 
to  1,000. 

Description  of  Areas  Excluded  from  Study 

There  are  two  factors  which  tend  to  upset  the  normal  pattern  of 
homogeneity  within  the  study  area.  Both  have  had  a  noticeable  effect 
on  the  development  and  distribution  of  central  urbanized  centers  within 
the  region. 

In  the  south-central  portion  of  the  study  area,  there  is  a  hilly, 
unglaciated  surface  located  within  the  Norman  Upland  physio- 
graphic region  (5).  This  is  a  rugged  area  with  the  greatest  local  relief 
in  the  state.  The  natural  agglomerating  principles  on  which  this  study 
is  based  are  restricted  to  a  great  extent  in  topographic  regions  as  com- 
plex as  this  upland  section.  Second,  in  the  late  19th  century,  natural 
gas  was  discovered  in  the  Muncie- Anderson  area  (7).  The  locations 
within  this  region  never  developed  in  such  a  way  that  they  can  be 
classified  trade  centers  within  the  criteria  set  forth  in  this  study. 
These  centers  developed  as  a  result  of  proximity  to  a  natural  resource 
rather  than  in  the  context  of  having  a  functional  service  location.  Since 
these  two  elements  are  not  suitable  to  space  and  distance  calculations, 
they  will  henceforth  be  segregated  from  the  rest  of  the  study  area  by 
suitable  boundaries  of  their  own. 

Model  Pattern  for  Indiana  Centers 

Having  once  established  a  distinct  hierarchy  pattern  for  the  study 
area,  the  next  step  is  to  analyze  the  spacing  of  the  centers  in  light  of 
the  various  groupings. 

Christaller  ordered  the  pattern  of  development  in  the  form  of 
hexagons.  Six  centers  within  each  net  are  located  at  equal  distances 
from  one  another  as  well  as  from  their  focal  place.  Each  center  within 


328  Indiana  Academy  of  Science 

the  group   represents   the   same  magnitude  of  centrality.    The   distance 
between  centers  of  each  successive  group  increases  by  the  \/  3. 

The  model  from  the  Tipton  Till  Plain  region  is  composed  of  46  type 
cast  centers  from  the  4  hierarchy  groupings.  The  centers  were  selected 
on  the  basis  of  the  best-fit  rule.  The  theoretical  distribution  based  on 
the  Christaller  scheme  portrays  the  centers  symmetrically  placed  about 
the  landscape. 

The  primary  (first  order)  city  is  Indianapolis.  It  is  the  most  cen- 
tralized of  the  major  cities  in  the  state.  Indianapolis  was  designated 
the  state  capitol  in  1825  largely  because  of  the  central  location  and  easy 
accessibility.  It  continues  to  function  as  the  major  focus  of  all  adminis- 
trative and  consumer  activity  within  the  study  area. 

There  are  six  secondary  centers  (excluding  those  in  the  indus- 
trialized area)  fairly  evenly  spaced  around  Indianapolis.  These  are 
Terre  Haute-West  Terre  Haute,  Bloomington,  Lafayette-West  Lafayette, 
Marion,  Richmond  and  Columbus-East  Columbus.  Each  occupies  a  focal 
point  on  an  apex  of  the  hexagon. 

Between  the  primary  city  and  the  six  secondary  centers  are 
grouped  six  tertiary  centers:  Crawfordsville,  Frankfort,  New  Castle, 
Shelbyville,  Franklin  and  Greencastle.  Although  Greencastle  and  Frank- 
lin were  previously  classified  fourth  order  centers,  their  strategic  loca- 
tion with  respect  to  serviceability  warrants  a  third  order  trade  dis- 
tinction. 

Around  each  of  the  first,  second  and  third  order  centers  are  quater- 
nary centers  in  clusters  of  6  (33  centers  in  all).  These  centers  range  in 
population  from  1,000  to  a  little  over  10,000.  Although  centers  such  as 
Bedford  were  previously  ranked  as  third  order  places,  their  true  trade 
status  is  more  comparable  to  the  quaternary  level.  Therefore,  a  few 
such  centers  have  been  adjusted  to  fit  the  model  more  correctly. 

Comparison  of  Spacing  Patterns 

In  actuality  the  centers  are  distributed  about  the  landscape  in  a 
more  random  fashion  than  the  model  prescribes.  However,  a  statistical 
analysis  concerning  the  spacing  of  the  trade  centers  gives  impressive 
support  to  the  hexagonal   theorem. 

The  theory  states  that  the  mean  distance  between  centers  within 
the  four  groupings  is  directly  proportional  with  the  distance  to  func- 
tional groups  of  the  next  higher  order.  The  mean  distance  between 
secondary  centers  is  38.6  miles,  while  the  mean  distance  to  the  primary 
city  is  58.0  miles.  The  tertiary  cities  have  a  mean  distance  of  35.5  miles; 
the  distance  to  the  secondary  cities  averages  35.1  miles.  A  mean  dis- 
tance of  20.7  miles  separates  quaternary  centers  with  the  nearest  places 
of  higher  order  approximately  20.9  miles  away. 

Christaller  further  projected  that  each  successive  hexagonal  net 
would  increase   by   the   radical   of  three.   This   would   postulate   that  by 


Geology  and  Geography  329 

ascribing  a  theoretical  value  to  the  secondary  centers,  the  lower  group- 
ings   would    approximate    the    \/3    rule    in    descending    order. 

Based  on  the  mean  distance  figure,  the  distance  between  secondary 
centers  is  ordered  at  58.3  miles.  This  places  the  tertiary  centers  at  a 
model  distance  of  33.7  miles;  the  actual  computed  distance  is  35.1  miles. 
Quaternary  centers  should  average  a  distance  of  20.4  miles;  in  reality 
they   are   20.9   miles. 

When  observed  distances  and  theoretical  distances  are  analyzed 
quantitatively,  the  results  reflect  a  minimal  aberration  among  the  units 
tested.  The  average  variation  is  computed  to  be  3.18%  with  a  correla- 
tion coefficient  of  0.9984  (Table  1). 

Table  1.     Comparison  of  spacing  patterns. 


Hierarchy  Groupings 


2nd  to  1st  58.0  58.3  —.52    (.0032) 

2nd  to  2nd  58.6  58.3  +.51 

3rd  to  2nd  35.1  33.7  +4.15 

3rd  to  3rd  35.5  33.7  +5.34 


Mean 

Theoretical 

Variation 

Measured 

Distance 

from 

Distance 

( Miles — Based 

Theoretical 

(Miles)* 

on   V3> 

(Percentage) 

4th  to  3rd 

22.6 

20.4 

+  10.78 

4th  to  2nd 

20.5 

20.4 

+  .49 

(nearest) 

4th  to  1st 

19.6 

20.4 

—3.92 

(around     3rd) 

4th  to  4th 

20.7 

20.4 

+  1.47 

(around    2nd) 

4th  to  4th 

20.7 

20.4 

+  1.47 

*  The  distance  measurements  were  taken  from  a  1968  Indiana  Chamber  of  Commerce 
road  map.  Calculations  were  based  on  straight  line  distances.  It  was  found  that  straight 
line  distances  between  centers  averaged  from  77,-19%  less  than  road  distances.  Therefore 
a  mean  correction  factor  of  13%  should  be  applied  to  the  straight  line  distances  to  obtain 
the  approximate  read  distances  between  centers. 


Therefore  one  may  conclude  that  the  centers  of  this  region  tend 
to  lie  in  a  uniform  pattern  about  the  landscape,  evenly  spaced  within 
their  own  unique  groupings  as  well  as  externally  between  groupings. 
The  distance  between  the  various  hierarchies  conforms  directly  with 
Christaller's  observed  tendency  toward   W3  as  the  set  norm  of  increase. 

Analysis  of  Deviations  from  Theoretical  Locations 

The  theoretical  hexagon  system  was  rotated  to  a  position  of  best 
fit  with  reference  to  the  observed  location  of  the  model  centers  (Fig.  2). 
Best  fit  was  determined  by  calculating  the  position  which  warranted  the 
lowest  sum  departure  of  the  observed  secondary  centers  from  their 
theoretical  locations.  After  the  theoretical  pattern  had  once  been  fixed 


330 


Indiana  Academy  of  Science 


in  position,  departures  were  calculated  for  each  of  the  model  centers. 
These  figures  were  then  analyzed  to  determine  to  what  degree  each 
hierarchy    grouping    correlated    with    the    Christaller    framework. 

A  "t"  test  was  used  to  test  the  significance  of  the  difference  be- 
tween the  observed  and  the  theoretical  distances  to  adjacent  centers. 
The  test  was  computed  for  each  of  the  three  groups  in  an  attempt  to 
determine  the  amount  of  uniformity  within  units.  The  result  from  the 
"t"  test  showed  that  in  all  3  groupings  the  observed  distances  and  the 
theoretical  distances  were  not  significantly  different  at  the  0.05  level. 


LEGEND 

Primary     center 

Secondary    canters 

Tertiary   centere 
Quaternary    centers 


FIGURE  2.      Comparison,  of  theoretical  pattern  and  model  centers. 


Geology  and  Geography  331 

Therefore,  one  may  conclude  that  even  though  the  centers  do  devi- 
ate to  a  certain  degree  from  their  theoretical  location,  the  spatial 
patterns   are   similar  enough  to   be   considered   statistically   related. 

Near-Neighbor  Analysis 

To  test  if  the  centers  were  located  in  a  pattern  of  maximum  dis- 
persal (which  is  in  essence  what  the  hexagonal  theorem  prescribes), 
the  spacing  of  each  unit  with  respect  to  its  adjacent  settlements  must 
be  analyzed. 

Examination  of  this  pattern  can  be  accomplished  by  the  use  of 
the  Near-Neighbor  Analysis.  This  technique  indicates  the  degree  to 
which  any  observed  distribution  of  points  deviates  from  what  might  be 
expected  if  the  points  were  distributed  in  a  random  manner  within  the 
same  area  (2).  The  theory  states  that  there  are  three  distinct  patterns 
of  settlement:  Aggregate  (clustering),  Random  and  Uniform. 

The  mathematical  test  for  the  Near-Neighbor  pattern  assigns  an 
"R"  value  for  the  observed  density  of  points  in  the  area  under  con- 
sideration. An  "R"  value  of  less  than  1.00  represents  a  tendency  toward 
clustering  (0.00  equals  a  perfect  aggregate  pattern).  An  "R"  value  of 
greater  than  1.00  reflects  a  tendency  toward  uniform  spacing  (2.15 
equals  maximum   dispersal   or  the  perfect  hexagonal   pattern). 

The  following  results  were  obtained  by  use  of  Near-Neighbor  analy- 
sis. The  "R"  for  the  second  order  grouping  is  2.011.  Deviation  is  0.139 
(6.4%)  from  the  perfect  hexagonal  framework.  The  tertiary  centers 
have  an  "R"  of  2.126.  This  pattern's  deviation  is  less  than  2%  (0.024) 
from  the  maximum  hexagonal  dispersal.  The  "R"  for  the  fourth  order 
centers  is  2.060  with  a  departure  from  uniformity  of  only  0.090  (4.1%). 

Therefore,  one  may  conclude  on  the  basis  of  the  aforementioned 
data,  that  there  is  a  distinct  distributional  pattern  of  settlement  for 
this  region.  The  model  centers  do  in  effect  reflect  a  near  perfect  rela- 
tionship with  the  uniform  spacing  (2.15)  tendency  of  the  Near-Neighbor 
equation. 

Conclusion 

The  results  of  this  study  clearly  support  the  two  hypotheses  tested. 
Nature  dictates  a  certain  degree  of  order  to  the  development  of  central 
places.  The  central  places  of  central  Indiana  have  a  tendency  to  conform 
to  such  an  ordering  principle;  and  the  pattern  for  this  area  is  very 
much  similar  to  the  K-3  network  of  Walter  Christaller. 


Literature  Cited 

1.  Berry,    B.    J.   L.,    and   W.    L.    Garrison.    1958.    Functional   bases    of   the    central    place 
hierarchy.  Econ.  Geogr.  34:145-154. 

2.  Berry,   B.   J.  L.,  and  D.   F.   Marble.    1968.   Spatial   Analysis :   A   Reader   in   Statistical 
Geography.  Prentice-Hall,  Inc.,  Englewood  Cliffs,  N.J.  512  p. 


332  Indiana  Academy  of  Science 

3.  BRUSH,  J.  E.  1953.  The  hierarchy  of  central  places  in  southwestern  Wisconsin.  Geogr. 
Rev.  43  :380-402. 

4.  Hart,  J.  F.  1968.  Field  patterns  in  Indiana.  Geogr.  Rev.  63:450-471. 

5.  Kingsbury,  R.  C.  1966.  An  Atlas  of  Southern  Indiana.  Occasional  Pub.  No.  3.  Dept.  of 
Geogr.  Indiana  Univ.,  Bloomington.  60  p. 

6.  Schneider,  A.  F.  1966.  Physiography,  p.  40-56.  In  A.  A.  Lindsey  [ed.]  Natural  Fea- 
tures of  Indiana.  Indiana  Acad.  Sci.  Sesquicentennial  Volume.  Indianapolis.  600  p. 

7.  Visher,  S.   S.   1923.  Economic  Geography  of  Indiana.  D.  Appleton  and  Co.,  N.Y.   511  p. 

8.  Wayne,  W.  J.  1956.  Thickness  of  drift  and  bedrock  physiography  of  Indiana  north  of 
the  Wisconsin  glacial  boundary.  Progress  Rep.  No.  7.  Indiana  Geol.  Surv.,  Bloomington, 
Ind. 


Parameter  Measurement  in  Fluvial  Morphology 

Daniel  M.  Coffman,  Purdue  University 


Abstract 

Because  of  the  quantitative  significance  of  stream  order,  an  accurate,  ordered  drain- 
age map  has  become  a  base  for  much  research  in  fluvial  morphology.  These  maps  are 
often  prepared  from  U.  S.  Geological  Survey  1:24,000  topographic  maps  and  doubt  is 
created  about  their  usefulness  as  sources  of  primary  data.  This  study  indicates  that 
blue  lines  of  such  topographic  maps  represent  third-  and  fourth-order  streams.  Even 
interpretation  of  topographic  maps  by  Strahler's  method  of  "V's"  does  not  necessarily 
result  in  an  accurate  drainage  map.  Relief,  age,  and  geology  appear  to  control  the 
quality  of  drainage  maps  which  are  produced.  The  relative  relief  of  apparent  first-order 
streams  may  be  used  to  predict  the  quality  of  a  drainage  map  which  will  be  produced 
from  a  topographic  quadrangle  and  permits  classifications  of  these  quadrangles  as 
excellent,  marginal,  and  unacceptable. 


Introduction 

For  nearly  25  years  attempts  to  quantitatively  define  river  systems 
and  landforms  derived  from  their  development  have  stemmed  from  Hor- 
ton's  concept  of  an  ordered  stream  system  (3).  The  advantage  of  stream 
order,  in  a  given  physiographic  region,  is  its  relation  to  the  number, 
average  lengths,  gradients,  drainage  area,  and  perhaps  discharge  of 
stream  segments  of  varying  size.  A  modification  of  Horton's  work 
whereby  the  smallest  stream  tributaries  are  assigned  the  lowest  order 
was  introduced  by  Strahler  (8)  and  resulted  in  the  system  of  stream 
order  which  is  widely  used  today.  An  example  of  a  stream  ordered  ac- 
cording to  Strahler's  system  is  shown  in  Figure  1.  Because  of  stream 
order's  quantitative  significance,  an  accurate,  ordered  drainage  map 
has  become  a  base  from  which  research  into  fluvial  processes  normally 
begins. 

The  results  of  any  research  are  no  better  than  the  quality  of 
original  input  data,  and  this  is  especially  true  in  the  study  of  stream 
patterns,  geometry,  and  mechanics.  Here  the  raw  data  generally  in- 
volve measurement  of  the  number,  length,  and  gradient  of  stream 
segments;  width,  depth,  and  velocity  of  water  in  the  channel;  and 
area,  main  stream  length,  and  shape  of  the  drainage  basin.  If  these 
parameters  are  measured  and  mapped  in  the  field  the  quality  of  the 
data  is  generally  excellent.  Unfortunately,  few  researchers  have  time 
or  money  to  devote  to  this  procedurally  difficult  type  of  data  collection, 
resulting  in  the  use  of  topographic  maps  as  a  major  source  of  original 
data  for  studies  in  fluvial  morphology. 

The  purpose  of  this  paper  is  to  examine  the  problems  involved  in 
obtaining  an  accurately  ordered  drainage  map  from  commonly  available 
U.S.G.S.  topographic  maps.  This  study  was  sponsored  by  the  Purdue 
University  Water  Resources  Research  Center,  Office  of  Water  Resources 
Research,  but  this  paper  has  not  had  the  benefit  of  its  review.  The 
author   also    acknowledges    the    assistance    of    Dr.    W.    N.    Melhorn    and 

333 


334  Indiana  Academy  of  Science 

Prof.    R.    D.    Miles    in    discussions    relating    to    aspects    of   fluvial    mor- 
phology. 

Measurement  from  Maps 

In  the  field,  first-order  stream  channels  (defined  as  the  smallest 
unbranched  tributaries  of  a  river  system)  are  easily  recognized  result- 
ing in  the  production  of  a  very  accurate  drainage  map.  The  U.S.G.S. 
topographic  maps  on  which  many  studies  rely,  however,  do  not  gen- 
erally show  all  stream  channels.  Apparent  first-order  stream  segments 
identified  from  these  maps  thus  become  functions  of  map  scale  and  the 
type  of  map  interpretation  employed,  and  in  the  published  literature 
may  not  represent  from  article  to  article,  the  same  size  stream  channel. 
As  a  result,  parameters  such  as  average  length  may  be  correlated  with 
stream  orders  which   are  incorrectly  numbered. 

Because  of  this  common  practice  of  compiling  stream  data  directly 
from  topographic  maps,  much  effort  has  been  devoted  to  determining 
the  validity  of  ordering  streams  from  the  various  types  of  maps  com- 
monly available.  Morisawa  (5)  concluded  that  U.S.G.S.  topographic 
maps  scaled  1:62,500  are  not  reliable  for  measuring  drainage  basin 
characteristics  other  than  area.  She  also  stated  that  data  on  numbers 
and  lengths  of  stream  segments  show  great  variation  when  taken  from 
maps.  Coates  (2),  in  working  with  streams  in  southern  Indiana,  found 
that  on  U.S.G.S.  maps  scaled  1:24,000  first-  and  second-order  streams 
were  rarely  shown  and  that  most  channels  interpreted  as  first-order 
were  actually  third-order.  Strahler  (9)  proposed  an  improvement  in 
interpreting  topographic  maps,  by  adding  to  those  stream  channels 
shown  in  blue,  segments  where  "V's"  in  contour  lines  indicate  that 
valleys  are  present.  This  method  greatly  aids  in  adding  otherwise  over- 
looked stream  segments  to  the  drainage  net,  but  produces  an  un- 
certainty as  to  the  size  of  the  smallest  segment  now  shown. 

Because  of  the  ever  increasing  need  for  basic  data,  some  research- 
ers have  ignored  published  warnings  about  the  questionable  validity 
of  a  drainage  map  prepared  from  topographic  maps.  One  example  is 
provided  by  Stall  and  Fok  (7)  who  ordered  drainage  basins  from 
U.S.G.S.  topographic  maps  scaled  1:62,500  without  interpretation  by  the 
method  of  "V's."  Their  correlation  of  discharge,  drainage  basin  area, 
length,  and  slope  which  they  ascribe  to  low  order  segments  must  con- 
servatively belong  to  fourth-  and  fifth-order  streams.  Although  some 
authors  merely  bypass  the  question  of  their  data's  validity,  Scheidegger 
(6)  suggested  what  the  techniques  of  other  investigators  appear  im- 
plicitly to  assume;  that  if,  for  instance,  second-order  streams  are 
treated  as  first-order  streams  because  of  map  inaccuracy  the  results 
obtained  in  a  study  are  proportional  to  the  parameters  of  the  actual 
stream  system.  If  this  is  true,  after  a  field  check  the  values  of  the 
actual  system  may  be  obtained  from  the  study  by  using  Horton's  law 
of  drainage  composition  or  some  other  mathematical  tool  of  extrapola- 
tion. 


Geology  and  Geography  335 

Objectives  of  this  Study 

Publication  of  the  Atlas  of  County  Drainage  Maps  for  Indiana  in 
July,  1959,  permits  large  areal  studies  of  stream  ordering-  to  be  con- 
ducted while  maintaining  a  high  level  of  confidence  in  the  data  (1). 
These  maps  were  prepared  by  the  staff  of  the  airphoto  laboratory  of 
the  Engineering  Experiment  Station  at  Purdue  University  from  1940 
A. A. A.  aerial  photographs.  Depending  on  the  time  of  year  that  photo- 
graphs were  taken  these  maps  are  estimated  to  show  an  average  of 
90%  of  all  existing  stream  channels  (R.  D.  Miles,  personal  communica- 
tion). A  field  check  in  the  vicinity  of  Tippecanoe  County  demonstrated 
that  these  maps  show  almost  all  first-order  segments.  Those  stream 
channels  not  shown  appeared  young,  probably  having  developed  since 
the  date  of  photography. 

Using  these  maps  as  a  base,  a  study  was  conducted  to  answer  the 
following  questions  about  drainage  maps  prepared  from  U.S.G.S.  topo- 
graphic maps  scaled   1:24,000  with  10-foot  contour  interval: 

1)  What  portion  of  a  drainage  basin  and  what  order  stream  seg- 
ments are  actually  shown  by  the  blue  lines  of  these  maps  ? 

2)  What  increase  in  accuracy  is  obtained  by  interpreting  these 
maps  using  Strahler's  method  of  "V's"  ? 

3)  Are  drainage  maps  produced  by  analysis  of  data  in  the  above 
two  cases  proportional  to  the  actual  stream  net,  and  if  not,  what 
effect  do  lost  segments  have  on  ordering  the  basin? 

4)  Does  accuracy  of  these  maps  vary  with  location  in  the  state  of 
Indiana  ? 


Procedure 

Drainage  basins  up  to  sixth-order  in  size  were  selected  randomly 
from  the  different  physiographic  provinces  of  the  state  (4).  Topographic 
maps  for  these  basins  were  obtained  and  drainage  maps  were  produced, 
initially  by  tracing  only  those  streams  shown  by  solid  or  dashed  blue 
lines,  and  secondly  by  adding  all  stream  segments  which  could  be 
recognized  using  the  method  of  "V's."  Experiments  with  different  oper- 
ators indicated  that  more  than  a  general  knowledge  of  topographic 
maps  was  required  to  apply  the  method  of  "V's"  and  that  some  practice 
was  necessary  before  any  operator  made  the  best,  consistent  use  of  this 
method.  Once  familiar  with  the  method,  however,  different  operators 
could  produce  reasonably  identical  maps  of  the  same  drainage  basin. 

Once  the  two  tracings  were  prepared  they  were  ordered  and  com- 
pared with  an  ordered  base  map  taken  from  the  Atlas  of  County  Dram- 
age  Maps.  The  data  for  this  segment-to-segment  comparison  was 
punched  on  IBM  cards  and  analyzed  by  a  CDC  6500  computer. 


336 


Indiana  Academy  of  Science 


■8-2 

*  * 

Q  P 

-■ 

o 

E 


<U     ft 


<J       EC 


83    2 

< 


C     c  to   « 

S    I  B  W 

Sis  S» 

*  m  P 


U     ft 

Ph    ft 


ft 

E 


O    M 

0) 


ft   X 

ft  H 
<! 


w  P 


+>  GO    °   g 

Slew 
«   S   ai  a 

££  ££ 


15 

Sg. 

CO    £    ft 


O     O    (N    O 


O    lO    ^     lO 


o   o   o   o 


©  o  o  © 


©    O    ©    "* 


©  w  as   © 

rH     ■>*     CO     © 


Geology  and  Geography 


337 


Table  2.    Accuracy  of  U.S.G.S.  maps  according  to  physiographic  province. 


Physiographic  Province 


Range  of 

General 

Relief 


Geology 


Percent  of  Various 

SeK 

men 

ts  Shown 

on 

Map 

1 

2 

3 

4 

its 

LOO 

LOO 

100 

97 

LOO 

LOO 

100 

97 

LOO 

LOO 

100 

89 

96 

100 

100 

79 

90 

100 

100 

(57 

90 

99 

100 

19 

4X 

91 

100 

15 

45 

100 

100 

Crawford   Upland 
Norman    Upland 
Dearborn   Upland 
Scottsburg    Lowland 
Muscatatuck   Slope 
Wabash    Lowland 
Tipton    Till    Plain 
Lake    and    Moraine 
Mitchell     Plain 
Indiana 


220-350  Mississippian     SS,     LS,     SH 

170-260  Mississippian     SS,     SH 

120-250  Ordovician    LS 

100-150  Devonian   &   Mississippian   SH 

100-150  Silurian     LS 

100-150  Pennsylvanian    SH,   SS 

100-150  Glacial  Till 

80-120  Fluvial    Glacial 

Underground  Drainage 
All  Provinces  Using  Only  the  Blue  Lines 


Results  of  Computer  Analysis 

The  percentage  of  each  order  of  stream  segments  shown  by  blue 
lines  on  the  topographic  maps  was  found  to  be  about  the  same  for  all 
physiographic  provinces  of  the  state  (see  "Indiana"  in  Table  2).  Table  1 
shows  results  for  basins  from  the  Tipton  Till  Plain.  These  findings 
agree  well  with  those  of  Coates  (2)  as  almost  no  first-  and  second-order 
streams  are  shown.  From  one-third  to  one-half  of  the  third-order  seg- 
ments appear  and  almost  all  fourth-order  segments,  are  shown.  As  the 
breakdown  into  apparent  order  indicates,  nearly  all  third-order  streams 
and  one-half  of  the  fourth-order  streams  would  be  classed  as  first-order 
if  only  blue  lines  were  used  to  construct  a  drainage  map.  Of  4,498  seg- 
ments from  various  basins  used  in  this  analysis,  only  153  or  about  4% 
of  all  segments  were  shown  in  blue  on  the  1:24,000  U.S.G.S.  topographic 
maps. 

An  increase  in  the  number  of  stream  segments  located  using  the 
1:24,000  U.S.G.S.  topographic  maps  was  found  to  be  significant  in  every 
physiographic  province  if  interpretation  by  Strahler's  (9)  method  of 
"V's"  was  employed.  Table  1  shows  that  in  the  Tipton  Till  Plain  about 
Vs  of  the  first-order,  V2  of  the  second-order,  and  almost  all  of  the  third- 
order  channels  are  shown.  The  proportion  of  total  segments  shown  in- 
creased from  4<7r  to  28%  in  this  province,  whereas  in  other  physiographic 
provinces  of  the  state  the  method  of  "V's"  resulted  in  almost  100%  re- 
covery of  all  stream  segments.  Even  in  areas  where  this  method  proved 
most  successful,  however,  it  still  contained  two  major  uncertainties.  First, 
the  correct  length  of  an  added  segment  was  always  in  doubt,  commonly 
by  a  factor  many  times  the  contour  interval.  Second,  many  segments 
were  recognized  whose  point  of  junction  with  other  channels  was 
uncertain,  which  ultimately  could  result  in  incorrect  ordering  of  the 
stream  net. 

In  provinces  where  almost  all  stream  segments  in  a  basin  are 
obtained   using  the   method   of  "V's,"   only  the   uncertainty   of  junction 


338 


Indiana  Academy  of  Science 


Us 

N 

L 

Us 

N 

L 

1 

236 

.2 

4 

--. 

9 

1.6 

2 

81 

.4 

5 

-== 

3 

2.6 

3 



27 

.8 

6 

1 — - 

1 

- 

R  =  3 


Figure  1.    Hypothetical  stream. 


locations  will  cause  lack  of  proportionality  between  actual  and  traced 
stream  nets.  However,  if  a  portion  of  segments  is  missed,  either 
because  of  the  map  or  inaccurate  procedures  used  in  tracing  a  stream 
net,  the  ordered  stream  system  which  results  will  be  in  no  way  propor- 
tional to  the  real  stream  system.  To  emphasize  this  point,  consider  the 
stream  system  shown  in  Figure  1.  It  has  a  perfect  bifurcation  ratio  (R) 
of  three,  and  the  numbers  (N)  and  average  lengths  (L)  for  the  various 
orders  are  indicated.  Figure  2  is  the  same  drainage  basin  without  first- 
and  second-order  segments.  This  remains  a  reasonable  representation 
of  the  original  basin  and  provides  an  example  of  the  stream  net  which 
some  researchers  assume  is  obtained  by  ordering  stream  systems  from 
large-scale    topographic    maps.    The    bifurcation    ratio    is    unaltered    and 


Geology  and  Geography 


Mi) 


Us 

N 

L 

Us 

N 

L 

1 

27 

.8 

3 



3 

2.6 

2 

9 

1.6 

4 

. 

1 

- 

Figure  2.    Hypothetical  stream  without  1st  and  2nd  orders. 


after  a  field  check  the  laws  of  drainage  composition  would  allow  calcula- 
tion of  the  numbers  and  average  lengths  of  stream  orders  not  shown  by 
this  map. 

Figure  3,  however,  shows  what  actually  happens  when  a  stream  net 
is  ordered  directly  from  the  blue  lines  of  a  U.S.G.S.  topographic  map 
scaled  1:24,000.  Since  approximately  only  V3  of  the  third-order  stream 
segments  are  shown,  the  ordered  basin  is  no  longer  proportional  to  the 
real  stream  net.  The  bifurcation  ratio  is  altered  and  the  average  seg- 
ment lengths  become  distorted.  It  is  impossible  to  extrapolate  the  prop- 
erties of  the  original  stream  net  from  this  ordered  basin.  Using  the 
data  for  the  Tipton  Till  Plain,  Figure  4  shows  a  theoretical  drainage 
map  produced  from  a  U.S.G.S.  topographic  map  using  the  method  of 
"Vs."  It  shows  more  segments  than  the  stream  net  in  Figure  3,  but  the 


340 


Indiana  Academy  of  Science 


Us 

N 

L 

Us 

N 

L 

1  , 

10 

2.0 

3 



1 

6.0 

2 

4 

.9 

4 



0 

- 

Figure  3.    Stream  as  it  might  appear  on  USGS  map. 


bifurcation  ratio  is  still  altered  and  the  measurement  of  average  lengths 
is  meaningless. 

The  ability  of  interpretation  using  the  method  of  "V's"  to  add  seg- 
ments to  the  blue  lines  of  topographic  maps  was  found  to  vary  consid- 
erably depending  on  location  in  the  state.  This  variability  appears  to  be 
a  function  of  1)  basin  relief,  2)  age  of  development  of  topography 
and  3)  geology.  Basin  relief  (the  difference  in  elevation  from  the 
drainage  divide  to  the  mouth  of  the  stream)  is  an  indication  of  slope. 
The  greater  the  slope  the  more  contour  lines  a  given  length  of  stream 
crosses  and  thus  the  easier  it  is  to  map.  As  the  length  of  time  during 
which  the  topography  has  been  evolving  increases,  low  order  streams 
develop  more  distinct  channels,  commonly  rectangular  in  shape,  result- 
ing in  more  distinct  "V's"  in  the  contour  lines.  Geology  effects  resistance 


Geology  and  Geography 


341 


Us 

N 

L 

Us 

N 

L 

1 

43 

.7 

3 



4 

2.1 

2 



12 

1.5 

4 

•m^» 

1 

— 

Figure  4.    Stream  as  it  might  be  mapped  using  method  of  V 


to  erosion  and  thus  controls  the  rate  of  gully  development.  Some  geologi- 
cal materials  are  characterized  by  shallow  channels  whereas  others 
possess  steep,  deep  channels  more  likely  to  produce  pronounced  "V's"  in 
contour  lines. 

Because  physiographic  divisions  of  the  state  reflect  differences  in 
relief,  age,  and  geology  some  generalizations  about  the  topographic 
maps  covering  these  areas  may  be  made.  Table  2  lists  the  divisions  and 
indicates  the  percent  of  various  order  streams  which  would  likely  be 
obtained  by  interpreting  a  1:24,000  U.S.G.S.  topographic  map  using 
Strahler's  (9)  method  of  "Vs."  This  table  intends  to  show  only  average 
conditions  and  because  of  variations  within  any  province  it  should  not 
be  used  as  an  accurate  guide  for  all  topographic  quadrangles  within  this 
physiographic    division.    No    data    were    taken    for    the    Mitchell    Plain 


342 


Indiana  Academy  of  Science 


because  drainage  in  that  province  is  essentially  underground.  Data  for 
the  Lake  and  Moraine  Province  are  poor  and  inconclusive  because  of 
extensive  development  of  ditch  systems  over  most  of  the  area,  which  are 
mapped  in  detail  as  cultural  features  on  U.S.G.S.  quadrangles. 

Estimation  of  Map  Accuracy 

As  most  first-  and  second-order  streams  are  tributary  to  third-order 
segments,  and  because  they  account  for  the  majority  of  those  segments 
not  shown  on  the  topographic  maps,  a  method  to  estimate  their  presence 
or  absence  was  sought.  It  has  been  demonstrated  that  the  smallest 
dashed,  blue  lines  on  a  U.S.G.S.  quadrangle  represent  third-  and 
fourth-order  channels  and  that  relief  is  of  prime  importance  in  the 
ability  to  interpret  a  map  using  the  method  of  "Vs."  Thus,  the  average 
relative  relief  of  apparent  first-order  streams  was  compared  with  the 
quality  of  interpretation  using  the  method  of  "Vs."  Ten  apparent  first- 
order  segments  were  chosen  from  a  basin  (or  quadrangle  if  the  basin 
was    small)    and    their    relative    relief    calculated.    Highest    and    lowest 


100 


KEY 

• 

LAKE  AND 

MORAINE 

PROVINCE 

O 

TIPTON 

TILL 

a 

< 
2 

PLAIN 

A 

(0 

WABASH 

© 

LOWLAND 

u> 

Z> 

X 

z 

o 

CRAWFORD 

z 

UPLAND 

o 

I 

D 

v> 

NORMAN 

t- 

UPLAND 

z 

kJ 

2 

© 

Ui 

SCOTTSBURG 

V) 

LOWLAND 

z 

< 

UI 

A 

a: 

l- 

MUSCATATUCK 

V) 

SLOPE 

u. 
o 

*- 

■ 

z 

ID 

DEARBORN 

o 
a: 

UPLAND 

Ui 

a. 

0  50  100  150  0  50  100  0 

AVERAGE      RELATIVE     RELIEF     OF     APPARENT     FIRST     ORDER     STREAMS 

Figure  5.    Plot  of  average  relative  relief  versus  percent  of  streams. 


Geology  and  Geography  343 

values  were  discarded  and  the  average  relative  relief  of  the  eight 
remaining  segments  was  plotted  against  the  percent  of  first-,  second-, 
and  third-order  segments  shown  on  the  map.  Results  as  shown  in 
Figure  5  indicate  that  the  average  relative  relief  provides  a  good 
estimate  of  the  accuracy  of  a  single  U.S.G.S.  quadrangle  map. 

Average  relative  relief  of  apparent  first-order  streams  has  been 
used  to  divide  U.S.G.S.  quadrangles  into  three  separate  classes;  excel- 
lent, marginal,  and  unacceptable.  Quadrangles  with  an  average  relative 
relief  of  150  feet  or  more  are  excellent,  showing  all  stream  segments  of 
all  orders,  and  may  be  used  with  confidence  to  produce  an  accurate 
drainage  map.  Quadrangles  with  an  average  relative  relief  less  than  80 
feet  are  unacceptable  because  they  fail  to  show  first-,  second-,  and  some 
third-order  segments,  making  any  drainage  map  produced  from  them 
unproportional  and  unreliable. 

If  average  relative  relief  is  between  80  and  150  feet  the  topographic 
map  is  marginal  as  a  base  for  production  of  an  accurate  drainage  map. 
Assuming  random  failure  to  detect  "lost"  stream  segments  the  mini- 
mum acceptable  omission  of  first-order  streams  may  be  calculated  using 
the  formula 

2 

(1 )  •  (100)  where  R  =  bifurcation  ratio 

R 

and  is  compared  to  the  probable  percentage  of  first-order  streams  as 
obtained  from  Figure  5  after  calculating  the  average  relative  relief.  If 
enough  first-order  segments  are  shown  the  quadrangle  should  produce 
a  drainage  map  proportional  with  the  real  system.  If  R  is  not  known  it 
is  recommended  that  three  be  used  because  it  is  the  worst  possible  value. 

Conclusions 

Unless  drainage  maps  are  produced  by  field  mapping  or  from 
interpretation  of  large-scale  aerial  photographs  their  reliability  as 
accurate  models  of  the  real  stream  net  must  be  initially  questioned.  No 
acceptable  drainage  map  can  be  made  from  a  U.S.G.S.  topographic  map 
scaled  1:62,500  without  considerable  field  verification.  If  drainage  maps 
are  traced  from  1:24,000  U.S.G.S.  topographic  maps,  using  only  the 
streams  shown  by  blue  lines,  the  apparent  first-order  segments  actually 
will  represent  third-  and  fourth-order  streams.  If  these  quadrangles 
are  interpreted  using  the  method  of  "V's"  uncertainty  about  order, 
length,  and  junction  location  of  added  segments  is  introduced. 

The  quality  of  interpretation  by  the  method  of  "V's"  varies  as  a 
function  of  1)  basin  relief,  2)  age  of  topography,  and  3)  geology.  An 
approximate  guide  to  the  quality  of  a  drainage  map  produced  from  a 
given  topographic  map  is  thus  the  physiographic  province  from  which  it 
comes.  It  has  been  demonstrated  that  the  average  relative  relief  of 
apparent  first-order  stream  segments  shown  by  blue  lines  can  be  used 
to  predict  the  quality  of  a  topographic  map  for  use  in  preparing  an 
accurate    drainage    map.    Based    on    this    average    relative    relief    the 


344  Indiana  Academy  of  Science 

1:24,000    U.S.G.S.    topographic    maps    for    Indiana    may    be    classed    as 
excellent,  marginal,  and  unacceptable. 

Literature  Cited 

1.  Atlas  of  County  Drainage  Maps  of  Indiana.  1959.  Purdue  Univ.  and  State  Highway 
Dept.  Ext.  Ser.  Rep.  No.  97. 

2.  Coates,  D.  R.  1958.  Quantitative  geomorphology  of  small  drainage  basins  of  southern 
Indiana.  Columbia  Univ.  Dept.  Geol.  Tech.  Rep.  10  :  67  p. 

3.  Horton,  R.  E.  1945.  Erosional  development  of  streams  and  their  drainage  basins ; 
hydrophysical  approach  to  quantitative  morphology.  Geol.  Soc.  Amer.  Bull.  56 :275- 
370. 

4.  Malott,  C.  A.  1922.  The  physiography  of  Indiana,  p.  57-257.  In  W.  N.  Logan  and 
others.  Handbook  of  Indiana  geology.  Pub.  21  Indiana  Dept.  Cons.,  Indianapolis. 

5.  Morisawa,  M.  E.  1959.  Relation  of  quantitative  geomorphology  to  stream  flow  in 
representative  watersheds  of  the  Appalachian  Plateau  Province.  Columbia  Univ.  Dept. 
Geol.  Tech.  Rep.  20  :  94  p. 

6  Scheidegger,  N.  A.  1965.  The  algebra  of  stream-order  numbers.  U.  S.  Geol.  Surv.  Prof. 
Paper  525-B  :  B187-B189. 

7.  Stall,  J.  B.,  and  Yu-Sl  Fok.  1968.  Hydraulic  geometry  of  Illinois  streams.  Univ.  of 
111.  Water  Resources  Center  Rep.  15  :  47  p. 

8.  Strahler,  A.  N.  1952.  Hysometric  (area-altitude)  analysis  of  erosional  topography. 
Geol.  Soc.  Amer.  Bull.  63:1117-1142. 

9.  Strahler,  A.  N.  1957.  Quantitative  analysis  of  watershed  geomorphology.  Amer. 
Geophys.  Union  Trans.  38  :913-920. 


MICROBIOLOGY  AND  MOLECULAR  BIOLOGY 

Chairman:  Warner  S.  Wegener,  Indiana  University  Medical  Center 
Larry  Day,  Eli  Lilly  &  Co.,  was  elected  Chairman  for  1970 

ABSTRACTS 

Stream  Pollution  from  Coal  Mine  Waste  Piles:  Effect  of  Sulfur  and  Iron 
Oxidizing  Bacteria.  Robert  Ramaley  and  Richard  Kindig,  Indiana 
University. — Analysis  of  100  samples  of  stream  water  taken  during 
hydrological  studies  of  the  Patoka  (Pike  County,  Ind.)  and  the 
Busseron  (Sullivan  County,  Ind.)  watersheds  showed  a  correlation 
between  the  acidity  of  the  samples  and  the  number  of  sulfur  or  iron 
oxidizing  bacteria  (Thiobacillus-F 'errobacillus) . 

Data  obtained  during  local  washouts  of  surface  mined  areas  and 
experiments  with  sterilized  mine  waste  material  are  consistent  with  the 
production  of  the  acid  pollution  and  perhaps  some  of  the  oxidized  iron 
by  the  action  of  sulfur  and  iron  oxidizing  bacteria  on  sulfur  bearing 
mine  wastes  and  a  subsequent  washing  out  of  both  the  acid  and  some 
of  the  bacteria  into  the  streams  following  rainfall. 

Core  samples  taken  of  land  reclaimed  in  accordance  with  the  1967 
Indiana  Mining  Act  showed  that  such  reclamation  was  effective  in 
reducing  the  number  of  sulfur  and  iron  oxidizing  bacteria  in  the  covered 
mine  maste  material  and  thereby  decreasing  the  acid,  production  to 
negligible  amounts. 

Fine  Structure  Changes  during  Germination  of  Dictyostelium  dis- 
coideum  Spores.  David  A.  Cotter.  Indiana  University  Medical  Center. 
—The  earliest  developmental  stage  in  the  life  cycle  of  the  cellular 
slime  mold,  Dictyostelium  discoideum,  is  spore  germination.  Spores  of 
this  cellular  slime  mold  can  be  induced  to  germinate  by  exposure  to  a 
mild  heat  shock  of  45°  C  for  30  minutes.  The  spore  germination  process 
occurs  in  four  well-defined  stages:  1)  dormant  spore  activation,  2) 
post-activation  lag,   3)    spore   swelling,   and   4)    myxamoeba   emergence. 

Electron  microscopy  revealed  significant  changes  in  the  fine  struc- 
ture of  germinating  spores  during  stages  three  and  four.  The  mito- 
chondria progressively  became  less  dense,  lost  their  peripherally 
attached  ribosomes,  and  revealed  more  pronounced  tubuli  as  germina- 
tion proceeded.  During  spore  swelling,  the  three-layered  spore  wall 
broke  down  in  two  stages:  1)  the  outer  and  middle  layers  were  rup- 
tured as  a  unit;  and  2)  the  inner  wall  was  breached.  Crystals  and  dark 
(lipid)  bodies  seemed  to  disappear  shortly  before  or  during  emergence 
of  the  myxamoebae.  Autophagic  vacuoles  were  found  in  dormant  spores 
and  throughout  the  entire  germination  process. 

The  addition  of  cycloheximide  to  germinating  spores  inhibited  the 
loss  of  the  crystals  and  dark  (lipid)  bodies.  In  addition,  the  drug 
inhibited  the  breakdown  of  the  inner  wall  layer.  Cycloheximide  did  not 

345 


346  Indiana  Academy  of  Science 

prevent  the  formation  of  the  water  expulsion  vesicle  or  the  apparent 
function  of  the  autophagic  vacuoles.  A  hypothesis  is  offered  to  explain 
the  inhibition  of  myxamoebae  emergence  by  this  drug. 

Thermal  Induction  of  Bacteriophage  PL  S.  S.  Lee,  Indiana  University 
Medical  Center. — The  induction  of  a  triply  auxotrophic  (thy  ,  ura",  met) 
Escherichia  coli  lysogenic  for  PI  was  studied  under  different  nutritional 
and  temperature  conditions.  At  37°  C,  as  reported  by  others,  induction 
occurs  only  under  thymineless  conditions.  At  45°  C,  induction  occurs 
under  thymineless  conditions,  but  also  under  conditions  of  amino  acid 
deprivation.  Although  thermal  induction  appears  to  be  a  different 
process  than  thymineless  induction,  the  fact  that  the  two  interact 
synergistically  suggests  that  both  involve  repressor  inactivation. 
Thermal  induction  is  generally  less  complete  than  thymineless  induction 
at  37°  C.  The  fact  that  pre-treatment  at  37°  C  under  condition  of 
amino  acid  deprivation  lowers  subsequent  thermal  induction  under  con- 
ditions where  thymine,  uracil  and  methionine  are  all  withheld  suggests 
that  cells  that  have  just  completed  one  round  of  DNA  replication  and 
have  not  initiated  another  round  are  relatively  insensitive  to  thermal 
induction.  Finally,  non-induced  cells  undergo  little,  if  any,  thymineless 
death  at  45°  C,  although  induced  cells  slowly  lose  plaque-forming 
ability  at  this  temperature. 

Computer-based  Derivation  of  Rate  Equations  for  Enzyme-catalyzed 
Reactions.  Arthur  R.  Schulz  and  Donald  D.  Fisher,  Indiana  Univer- 
sity Medical  Center. — An  algorithmic  process  has  been  developed  which 
provides  for  derivation  of  rate  equations  for  enzyme-catalyzed  reactions 
by  a  digital  computer.  An  explicit  notation  system  is  employed  which 
is  adaptable  to  computer  processing  techniques.  The  sequence  of  an 
enzymic  reaction  is  represented  by  two  matrices.  A  connection  matrix 
is  used  to  determine  the  valid  paths  which  connect  the  enzyme  species, 
and  a  matrix  of  substrate  and  product  names  is  used  to  append 
reactant  symbols  to  the  proper  path  vectors. 

The  denominator  of  a  rate  equation  is  given  by  the  sum  of  the 
valid  paths  which  connect  the  enzyme  species.  The  algorithm  provides 
for  derivation  of  the  numerator  of  the  rate  equation  and  for  alphabetical 
sorting  of  the  numerator  and  denominator  terms.  Reformulation  of  the 
rate  equation  from  the  "coefficient"  form  to  the  more  useful  "kinetic" 
form  is  accomplished  by  expressing  each  denominator  term  as  a  vector 
of  reactant  concentration  exponents.  This  provides  for  computer-based 
definition  of  the  Michaelis  constants  and  for  all  possible  inhibition  con- 
stants, and  for  the  reformulation  of  the  equation  using  the  definitions 
selected  by  the  investigator. 

Axoplasmic  Transport  of  Materials  in  Nerve  Fibers.  S.  Ochs,  M.  I. 
Sabri,  and  N.  Ranish,  Indiana  University  Medical  Center.— Axoplasmic 
flow  is  present  in  nerve  fibers.  Our  recent  studies  have  shown  that  after 
lumbar  seventh  ganglion  injection  with  HMeucine,  a  crest  of  labelled 
activity   is   present   in   the   sciatic   nerve   with   the   position   of  the   crest 


Microbiology  and  Molecular  Biology  347 

moving  outward  at  a  rate  of  400  mm  per  day.  Evidence  that  the  labelled 
activity  is  intra-axonic  was  gained  by  freeze  block.  Extraction  of  labeled 
material  showed  it  to  be  present  in  particulate  and  soluble  form,  as  high 
molecular  weight  protein  (450,000  and  65,000)  and  polypeptide 
(4-13,000).  The  precursor  P:i2-orthophosphate  is  incorporated  into  a 
slower  moving  material.  The  possible  molecular  basis  of  the  fast  trans- 
port system  and  other  studies  regarding  the  mechanism  underlying 
intra-axonic  transport  were  discussed. 


Development  of   a    Clear,    Photopolymerizable   Acrylamide    Gel    and    Its 
Use  In  Immobilizing  and  Staining  Nucleic  Acids. 

Thomas  A.  Cole  and  Wayne  F.  Middendorf,  Wabash  College 

Abstract 

A  clear,  photopolymerizable  gel  of  a  poly  acrylamide  has  been  developed.  The  com- 
ponents will  form  a  gel  in  presence  of  high  concentrations  of  cesium  chloride.  DNA 
immobilized  in  such  a  gel  can  be  stained  and  the  blank  gel  subsequently  destained.  The 
application  of  this  technique  to  isopycnic  density  gradient  centrif ligation  of  cesium 
chloride   solutions    was    discussed. 

Introduction 

Polymerization  of  acrylamide  and  N,N'-methylene-bis-acrylamide 
has  been  widely  used  in  the  preparative,  routine  and  analytical  versions 
of  disc  electrophoresis  (1).  Additionally,  photopolymerization  of  these 
compounds  have  been  used  in  conjunction  with  sucrose  density  gradient 
centrifugation  for  immobilization  of  components  separated  by  centri- 
fugation  (2,  3,  4).  This  report  extends  the  immobilization  technique  to 
dense  solutions  of  cesium  chloride  and  details  the  development  of  a 
clear  photopolymerizable  gel.  The  techniques  of  staining  and  destaining 
the  immobilized   gel  are  reported  also. 

Methods  and  Materials 

Preparation  of  Cesium    Chloride-Containing  Gels 

Cesium  Chloride  (SC11352)  was  purchased  from  the  Sargent-Welch 
Company.  Cesium  chloride  was  dissolved  in  water  to  give  a  solution 
with  a  density  of  around  1.875  gm/ml.  Such  a  solution  requires  about 
1.200  gms  CsCl/ml  solution  at  20 °C.  Three  parts  of  the  cesium  chloride 
solution  were  mixed  with  one  part  of  polymerizable  solution. 

Polymerizable  solution  is  made  of  three  parts,  Solutions  B  and  C  and 
riboflavin.  Solution  B  consists  of  11.4  gm  Tris  buffer,  1.2  ml  N,N,N',N'- 
tetramethylethylenediamine  (TEMED,  practical  grade,  Matheson  Cole- 
man and  Bell  8563)  and  distilled  water  to  a  final  volume  of  33  ml.  The 
pH  is  adjusted  to  6.9  with  85  percent  phosphoric  acid.  Solution  C  is 
24  gm  of  acrylamide,  0.735  gm  N,N'-methylene-bisacrylamide  (Bis,  East- 
man 8383)  and  distilled  water  to  give  a  final  volume  of  50  ml.  Approxi- 
mately 0.5  mg  riboflavin/5  ml  aliquot  is  dissolved  in  the  polymerizable 
solution.  Solutions  B  and  C  are  stable  and  may  be  stored  at  room  tem- 
perature. After  addition  of  riboflavin  the  polymerizable  solution  must  be 
protected  from  light. 

Layers  of  gel  were  successively  polymerized  in  a  cellulose  nitrate 
centrifuge  tube.  Alternate  layers  contained  DNA.  Each  layer  was 
polymerized  from  0.4  ml  of  solution  (0.1  ml  of  Cesium  chloride  solution, 
0.3  ml  of  polymerizable  solution)  with  a  thin  (2  mm)  overlay  of  water. 
Those  layers  which  contained  DNA  were  made  by  diluting  stock  DNA 

348 


Microbiology  and  Molecular  Biology  349 

solution  with  cesium  chloride-polymerizable  solution.  Polymerization  was 
completed  for  each  layer  in  10  to  20  minutes  at  a  distance  of  4  inches 
from  a  15  watt  fluorescent  bulb. 

Staining  and  Destaining 

After  polymerization  the  gels  were  removed  from  the  cellulose 
nitrate  tubes  by  shaking  or  rimming  with  a  microspatula  and  were 
stained  by  one  of  a  variety  of  procedures.  Three  procedures  gave  the  best 
results  and  they  are  reported  here.  A  methyl  green  procedure  (5,  6)  gives 
a  green  stain,  pyronin  B  (5)  gives  a  lavender  stain  and  methyl  green  and 
pyronin  B  together  (7)  give  a  blue-green  stain.  The  gels  were  immersed 
in  the  staining  solution  for  1  hour  or  more.  The  excess  stain  was  re- 
moved by  repeated  washing  in  0.2n  acetate  buffer  of  an  appropriate  pH 
(4.0  for  methyl  green,  4.5  for  pyronin  B  and  4.25  for  methyl  green-py- 
ronin  B).  Electrophoretic  destaining  may  be  done  in  the  same  buffers. 
A  suitable  destaining  apparatus  may  be  constructed  from  lucite  (8).  The 
washing  procedure  takes  about  2  days  with  occasional  changes  of  buffer. 
The  electrophoretic  procedure  requires  about  24  hours  at  3-4  ma  of 
current  per  tube. 

DNA  Samples 

Calf  thymus  DNA  (Sodium  Salt,  Type  I,  lot  115B-1690)  was  pur- 
chased from  Sigma  Chemical  Company,  St.  Louis,  Missouri.  A  stock 
solution  of  DNA  was  prepared  by  dissolving  DNA  in  dilute  NaCl-citrate 
(0.015m  NaCl,  0.0015m  sodium  citrate,  pH  7). 

Results 

The  results  show  that  within  the  range  of  approximately  100  to  250 
/xg  calf  thymus  DNA/ml  good  contrast  bands  were  obtained.  At 
500fig/ml  the  DNA  partially  inhibited  the  polymerization  process. 
Consequently,  the  500^g/ml  band  was  small  due  to  loss  of  material. 

Discussion 

The  photopolymerizable  gel  of  disc  electrophoresis  (1)  and  sucrose 
density  gradient  techniques  (2,  3,  4)  is  translucent.  The  transparent  gel 
reported  here  is  the  result  of  lowering  the  Bis/acrylamide  ratio.  This 
means  that  the  transparent,  photopolymerizable  gel  is  not  as  highly 
cross-linked  as  the  translucent  one. 

The  results  reported  here  show  that  DNA  can  be  immobilized, 
stained  and  destained  in  a  gel  containing  cesium  chloride  at  concen- 
trations comparable  to  those  used  in  isopycnic  density  gradient  centri- 
fugation.  Thus,  cesium  chloride  does  not  interfere  with  the  polymeriza- 
tion process. 

The  methyl  green  stain  has  been  used  successfully  for  staining 
native  DNA  in  acrylamide  gels  (5);  since  these  gels  are  much  thicker 
than  those  of  disc  electrophoresis  the  primary  exposure  time  to  the 
stain  was  1%  more  hours  rather  than  1  hour  as  reported  by  Boyd  and 


350  Indiana  Academy  of  Science 

Mitchell.  The  pyronin  B  method  has  been  used  for  denatured  DNA  (5). 
Presumably,  our  DNA  samples  were  not  denatured  and  the  pyronin  B 
method  gave  the  least  satisfactory  results.  The  methyl  green-pyronin 
B  method  gave  the  best  contrast  but  the  methyl  green  alone  could  be 
completely  removed  from  non-DNA  containing  areas  of  the  gel. 

With  the  techniques  developed  here,  it  should  be  possible  to  carry 
out  isopycnic  cesium  chloride  density  gradient  centrifugation  with  DNA 
and  stain  the  samples  after  polymerization.  With  disc  electrophoresis 
many  staining  procedures  have  been  quantitated  with  a  microdensi- 
tometer  scanning  of  the  gel  (1).  This  instrument  costs  $3900.  Recently, 
an  inexpensive  high-resolution  densitometer  for  disc  electrophoresis 
has  been  developed  (9).  The  total  cost  for  materials  for  this  instrument 
are  reported  not  to  exceed  $300.  Thus,  a  guantitative  and  analytical 
data-acquisition  system  for  density  gradient  centrifugation  should  be 
possible  without  the  use  of  an  analytical  ultracentrifuge  (Model  E, 
Spinco  Division,  Beckman  Instruments,  Inc.,  Palo  Alto,  California  94304) 
or  drop  collecting  techniques.  The  requirements  for  such  a  system  are 
a  preparative  ultracentrifuge,  the  immobilization  and  staining  pro- 
cedure reported  here,  and  a  microdensitometer  for  gel  scanning. 


Acknowledgements 

This   work   was   supported   by   Grant   NIH-GM-11860,    U.    S.    Public 
Health    Service,   and    an   NSF-COSIP    grant   to    Wabash    College. 


Literture  Cited 

1.  Davis,  Baruch  J.  1964.  Polyacrylamide  Gel  Electrophoresis.  Ann.  New  York  Acad, 
of  Sci.  121  :404-427. 

2.  Jolley,  Weldon  B..  H.  W.  Allen,  and  O.  M.  Griffith.  1967.  Ultracentrifugation 
Using    Acrylamide    Gel.     Anal.     Biochem.     21:454-461. 

3.  Prins,  H.  K.,  and  D.  D.  Smink.  1965.  Zone-Centrifugation.  Bibl.  Haematol.  23:1186. 
(Proc.    10th    Congr.    Intern.    Soc.    Blood    Transfusion,    Stockholm,    1964.) 

4.  Cole,  Thomas  A.,  and  Thomas  W.  Brooks,  Jr.,  1968.  Density  Gradient  Centrifu- 
gation:   Fixation    of    Bands    by    Photopolymerization    of   Acrylamide.    Science:    161:386. 

5.  Boyd,  James  B.,  and  H.  K.  Mitchell.  1965.  Identification  of  Deoxyribonucleases  in 
Polyacrylamide  Gel  Following  Their  Separation  by  Disc  Electrophoresis.  Anal. 
Biochem.  13:28-42. 

6.  Kurniok,  N.  B.  1950.  The  Quantitative  Estimation  of  Desoxyribosenucleic  Acid 
Based   on    Methyl    Green    Staining.    Exp.    Cell    Res.    1:151-158. 

7.  Jensen,  W.  A.  1962.  Botanical  Histochemistry.  W.  H.  Freeman  and  Company,  San 
Francisco.  251  p. 

8.  Maurer,  H.  R.  1966.  Einfacher  Entfarbeappararat  fur  die  Disk-Electrophorese.  Z. 
Klin.  Chem.  4:85-86. 

9.  Petrakis,  Peter  L.  1969.  An  Inexpensive  High-Resolution  Densitometer  for  Disc 
Electrophoresis.  Anal.  Biochem.  28:416-427. 


The  Effect  of  Avidin  on  the  Biosynthesis  of  Fatty  Acids  in 

Aspergillus  niger  and  Aspergillus  flavus 

K.  Schwenk  and  A.  S.  Bennett,  Ball  State  University 

Abstract 

Submerged  cultures  of  Aspergillus  flavus  and  Aspergillus  niger  were  grown  in  a 
medium  containing  avidin,  a  substance  which  inhibits  the  conversion  of  acetate  to 
malonate.  Control  cultures  were  grown  without  avidin. 

Mycelium  samples  were  taken  at  time  points  over  a  70-hour  incubation  period, 
harvested  by  centrifugation,  washed  with  distilled  water  and  re-suspended  in  absolute 
methanol.  After  saponification,  the  fatty  acids  were  extracted  with  hexane,  methylated, 
isolated  by  thin  layer  chromatography,  and  separated  and  identified  by  gas  liquid 
chromatography. 

An  increase  in  Cm  fatty  acids  and  a  decrease  in  Cis  fatty  acids  by  cultures  grown 
with  avidin  suggest  that  malonate  plays  an  important  role  in  the  elongation  of  long- 
chain  fatty  acids  in  these  organisms  during  the  time  interval  from  5  to  15  hours. 

In  control  cultures,  the  initially  high  percentage  of  stearic  acid  decreased  while  the 
percentage  of  oleic  acid  and  linoleic  acid  increased,  further  indicating  the  conversion  of 
stearate  to  oleate. 

Several  monoenoic  acids  are  present  in  these  organisms,  but  the  only  dienoic  acid 
found  is  linoleic  acid.  This  suggests  that  the  conversion  of  oleate  to  linoleate  involves  a 
highly  specific  desaturase. 

The  first  step  in  the  biosynthesis  of  long  chain  fatty  acids  by  way 
of  the  malonate  pathway  involves  the  enzyme,  acetyl-CoA-carboxylase. 
This  enzyme  is  one  of  the  biotin-containing  carboxylases  and  catalyzes 
the  overall  reaction. 

Acetyl-CoA  +  HCCV  +  ATP  ?±  Malonyl-CoA  +  ADP  +  P, 
This  reaction  is  followed  by  the  successive  addition  of  2-carbon  units  in 
the  form  of  malonyl-CoA  or  malonyl-ACP  to  acetyl-CoA    (4).  The  end 
product   of   this   sequence   of   reactions    is   dependent   upon   the    type    of 
organism  and  is  either  palmitate  (9)  or  stearate  (3). 

Although  the  malonate  pathway  is  thought  to  be  the  predominant 
pathway  for  the  biosynthesis  of  fatty  acids,  evidence  for  the  existence 
of  alternate  pathways  has  been  presented  (4).  Mattoo  et  al.  (5)  reported 
that  the  biosynthesis  of  fatty  acids  in  intact  mycelium  of  Aspergillus 
flavus  was  only  partially  inhibited  after  increasing  amounts  of  avidin, 
a  biotin  inhibitor,  had  been  added  to  the  medium.  Their  results  sug- 
gested that  the  formation  of  malonate  was  not  essential  for  the  synthesis 
of  fatty  acids  in  this  organism.  The  fatty  acid  distribution  of  the  prod- 
ucts recovered  from  the  inhibited  cultures  was  not  determined. 

In  the  present  experiments  avidin  was  used  to  study  the  effect  of 
the  inhibition  of  malonate  formation  on  the  types  of  fatty  acids  synthe- 
sized by  A.  niger  and  A.  flavus. 

Materials  and  Methods 

Aspergillus  niger  and  Aspergillus  flavus,  obtained  from  Dr.  K.  B. 
Raper,   University  of  Wisconsin,  were  maintained  on  slants   of  Czapek 

351 


352  Indiana  Academy  of  Science 

medium  and  stored  at  4°  C.  Spores  were  lightly  scraped  and  washed 
from  the  slants  with  sterile  medium.  Five  ml  of  spore  suspension 
(A  =  0.1  at  525  m^)  was  added  to  25  ml  of  sterile  (autoclaved;  15  psi, 
15  min)  culture  medium  (glucose,  40  g;  NfLNO.,,  1  g;  MgS04«7H20, 
0.3  g;  KH,P04,  0.3  g;  H,0,  1  liter)  in  250  ml  Erlenmeyer  flasks.  The 
inoculated  cultures  were  incubated  for  24  hours  at  28  °C  on  a  recipro- 
cating shaker.  At  the  end  of  this  period,  stir  cultures  were  prepared  by 
adding  the  contents  of  the  flask  to  420  ml  of  culture  medium  in  a 
2-liter  Erlenmeyer  flask,  aerating  (500  ml/min)  and  stirring  for  60 
hours.  Forty-five  units  of  avidin  (1  U=amount  capable  of  inactivating 
1  A  biotin)  had  been  added  to  the  experimental  culture  medium;  no 
avidin  was  added  to  control  cultures. 

Aliquots  of  the  mycelial  suspension  were  removed  from  the  stir 
culture  at  various  time  points.  After  centrifugation,  the  mycelial  mat 
was  washed  with  distilled  water,  placed  in  15  ml  absolute  methanol,  and 
sonified  for  3  min  (J-17A  Sonifier,  Branson  Sonic  Co.  Power).  The 
sonified  material  was  diluted  with  an  equal  volume  of  15 %  KOH  in 
methanol,  refluxed  for  2  hours,  acidified  with  concentrated  HC1,  and 
extracted  with  hexane.  After  washing  the  extract  with  distilled  water, 
drying  over  anhydrous  sodium  sulfate  and  removing  the  solvent  on  a 
rotary  evaporator,  the  resulting  fatty  acids  were  methylated  with 
diazomethane  (7).  The  methyl  esters  were  isolated  on  silica  gel  G  thin 
layer  plates  and  separated  and  identified  by  gas  liquid  chromatography 
using  a  10-foot  glass  column  packed  with  5%  poly(diethylene-glycol 
adipate)  on  60/80  mesh  Chrom  GA/W;  column  temperature,  190°  C; 
gas  flow,  70  ml/min;  Varian  Aerograph  90-P  instrument.  For  further 
identification  of  unsaturated  fatty  acids,  methyl  esters  were  separated 
by  thin  layer  chromatography  on  10%  AgN03-impregnated  silica  gel  G 
with  chloroform :acetone  (99.5:0.5)  as  the  developing  solvent  prior  to 
GLC  analysis. 

Results  and  Discussion 

Members  of  the  class  Ascomycetes  have  a  fatty  acid  composition 
similar  to  that  of  plant  seeds  (2)  and  like  the  growing  leaves  and  seeds 
of  plants  produce  the  stearate  to  linolenate  series  of  unsaturated  fatty 
acids.  Shaw  (8)  reported  that  the  only  Cis  fatty  acids  produced  by  the 
ascomycetes  are  the  A9-unsaturated  fatty  acids.  Analysis  of  the  products 
in  the  present  experiments  revealed  that  linoleic  acid  was  the  only 
dienoic  acid  synthesized  by  A.  niger  and  A.  fla/vus,  although  the 
monoenoic  form  of  C9  through  Cis  acids  were  identified,  suggesting  that 
the  desaturase  which  is  responsible  for  the  conversion  of  oleate  to 
linoleate  is  highly  specific.  This  finding  makes  members  of  the  genus 
Aspergillus  particularly  well-suited  as  experimental  organisms  for 
studying  the  biosynthesis  of  unsaturated  fatty  acids  in  plant-like 
organisms. 

Cultures  of  Aspergillus  -niger  grown  in  a  medium  containing 
inhibitory  amounts  of  avidin  did  not  differ  from  those  grown  in  the 
absence  of  avidin  as  to  the  relative  rate  of  palmitate  synthesis  (Fig.  1). 


Microbiology  and  Molecular  Biology 


353 


25     h 


20 


1  5 


1  0 


1  0 


20  30  40  50 

Age   of   Culture    (hours) 


6  0 


Figure  1.     Percent  distribution  of  palmitic  and  stearic  acids   in  mycelium   of  Aspergillus 
niger  grown  in  media  with  and  without  avidin. 


a)  r) 

PS  -H 


■P  -P 

G  -P 


50    |- 


4  0 


A/ith  Avidin 


18:2 


2  0  3  0  4  0  5  0 

Age   of  Culture   (hours) 

Figure  2.     Percent  distribution  of  fatty  acids  in  mycelium  of  Aspergillus  niger  grown  in 
media  with  and  without  avidin. 


354 


Indiana  Academy  of  Science 


However,  the  relative  amount  of  stearic  acid  produced  was  substantially 
lower  in  the  mycelium  grown  in  the  avidin  containing  medium.  In  the 
control,  the  relative  amount  of  stearic  acid  recovered  increased  from 
12%  in  the  inoculum  to  22%  after  a  5-hour  incubation  period;  with 
avidin,  stearic  acid  decreased  from  12%  to  7%.  This  difference  in 
stearate  concentration  is  evident  during  the  first  30  hours  of  the  incu- 
bation period. 

At  the  5-hour  time  point  only  23%  of  the  fatty  acids  recovered 
from  the  mycelium  grown  in  the  avidin-containing  medium  was  of  the 
Cis  series,  compared  to  33%  in  the  control  (Fig.  2). 

These  data  suggest  that  malonate  plays  an  important  role  in  the 
elongation  of  palmitate  to  stearate.  Although  Mattoo  (5)  reported  a 
partial  overall  inhibition  of  fatty  acid  synthesis  upon  the  addition  of 
avidin  to  the  medium,  our  data  show  that  all  steps  in  the  biosynthetic 
pathway  were  not  affected  to  the  same  degree.  In  both  A.  niger  and 
A.  flavus,  the  addition  of  avidin  to  the  culture  medium  inhibited  the 
formation  of  long  chain  fatty  acids  to  a  greater  degree  than  short 
chain.  The  importance  of  malonate  in  the  elongation  of  palmitate  to 
stearate  in  Aspergillus  is  in  agreement  with  the  report  of  Nagai  and 
Bloch  (6)  who  found  that  elongation  to  stearate  in  bacterial  and  plant 
extracts  was  dependent  upon  the  presence  of  malonate.  Barron  (1) 
found  that  malonate  was  used  for  the  elongation  of  preformed  fatty 
acids  by  the  soluble  fraction  of  rat  liver  homogenates,  while  acetate  was 
used  by  the  mitochondrial  fraction. 


-P    -U 

O  tu 

U 

Q) 


1  0 


20  30  40  50 

Age  of  Culture  (hours) 


Figure    3.     Percent    distribution   of   fatty   acids    in    the   mycelium    of    Aspergillus    niger    at 
various  times  in  the  incubation  period. 


Microbiology  and  Molecular  Biology  355 

Bennett  and  Quackenbush  (2)  previously  reported  studies  on  the 
biosynthesis  of  fatty  acids  in  Penicillium  chrysogenum  which  indi- 
cated that  endogenous  palmitate  was  rapidly  elongated  to  stearate  and 
desaturated  by  the  sequential  pattern  of  oleate  to  linoleate  to  linolenate. 
In  the  present  experiments,  relatively  large  amounts  of  palmitate  and 
stearate  were  synthesized  at  early  time  points  in  the  growth  period 
(22%  and  12%,  respectively,  at  5  hours),  followed  by  increases  in  oleate 
and  linoleate  concentration  at  45  hours  (Fig.  3),  to  give  additional 
support  for  this  pattern  of  elongation,  then  desaturation. 


Literature  Cited 

1.  Barron,  E.  J.  1966.  The  Mitochondial  Fatty  Acid  Synthesizing  System :  General 
Properties  and  Acetate  Incorporation  into  Monoenoic  Acids.  Biochim.  Biophys.  Acta. 
116:425-435. 

2.  Bennett,  A.  S.,  and  F.  W.  Quackenbush.  1969.  Synthesis  of  Unsaturated  Fatty 
Acids  by  Penicillium  chrysogenum.  Arch.  Biochem.  and  Biophys.  130  :567-572. 

3.  Lynen,  F.  1961.  Biosynthesis  of  Saturated  Fatty  Acids.  Fed  Proc.  20:941-951. 

4.  Majerus,  W.  P.,  and  P.  R.  Vagelos.  1967.  Fatty  Acid  Biosynthesis  and  the  Role  of 
the  Acyl  Carrier  Protein.  Adv.  Lipid  Res.  5  :2-33. 

5.  Mattoo,  A.  K.,  V.  V.  Modi,  and  R.  N.  Patel.  1966.  Biotin  and  Fatty  Acid  Biogenesis 
in  Aspergillus  flavus.  Experimentia.  22  :436-437. 

6.  Nagai,  J.,  and  K.  Bloch.  1966.  Elongation  of  Acyl  Carrier  Protein  Derivatives  by 
Bacterial  and  Plant  Extracts.  J.  Biol.  Chem.  242  :357-365. 

7.  Schlenk,  H.,  and  J.  L.  Gellerman.  1960.  Esterification  of  Fatty  Acids  with 
Diazomethane  on  a  Small  Scale.  Anal.  Chem.  32:1412-1413. 

8.  Shaw,  R.  1965.  Occurrence  of  Linolenic  Acid  in  Fungi.  Biochim.  Biophys.  Acta. 
98  :230-236. 

9.  Wakil,  S.  J.  1961.  Mechanism  of  Fatty  Acid  Snythesis.  J.  Lipid  Res.  2:1-24. 


PHYSICS 

Chairman:  Richard  L.  Conklin,  Hanover  College 
Ralph  A.  Llewellyn,  Rose  Polytechnic  Institute,  was  elected  Chairman 

for  1970 

ABSTRACTS 

Measurement  of  Ionization  in  Nuclear  Emulsion  by  Lacunarity  Tech- 
nique. W.  David  Mueller,  Ball  State  University. — Lacunarity  was 
studied  as  a  technique  of  measuring  ionization  and  energy  loss  of  a 
charged  particle  passing  through  nuclear  emulsion.  Lacunarity  is  a 
measure  of  track  density  and  is  defined  as  the  linear  fraction  of  a 
track  segment  that  consists  of  gaps.  Measurements  were  made  of  blob 
density  and  lacunarity  of  segments  of  tracks  of  particles  emanating 
from  nuclear  disintegration  in  nuclear  emulsion.  Both  blob  density  and 
grain  density  as  determined  by  lacunarity  were  plotted  versus  residual 
range  of  track  segments  to  compare  the  two  methods  of  measuring 
ionization. 

Except  for  tracks  of  very  heavy  ionization  and  tracks  of  light 
ionization,  grain  density  as  determined  by  lacunarity  is  a  better  measure 
of  ionization  than  blob  density.  Both  theoretical  and  experimental  re- 
sults indicate  that  over  a  considerable  range  of  ionization,  blob  density 
does  not  vary  greatly  with  the  degree  of  ionization.  Also,  particles 
could  be  distinguished  by  lacunarity  which  could  not  be  distinguished  by 
blob  density. 

Operation  and  Flux  Determination  of  a  Neutron  Generator.  Martin  A. 
Burkle  and  Leon  M.  Reynolds,  Ball  State  University. — A  neutron  gene- 
rator-accelerator consisting  of  a  150  kv  Cockroft-Walton  accelerator  which 
makes  use  of  the  3H(d,  n)4He  reaction  was  placed  in  operation  by 
the  Department  of  Physics,  Ball  State  University.  Experiences  as- 
sociated with  the  installation  and  startup  of  the  equipment  were  de- 
scribed along  with  projected  uses.  Preliminary  results  for  neutron  flux 
determinations  using  2.8  Mev  neutrons  produced  by  the  2H(d,  n)3He 
reaction  were  given.  The  usefulness  of  the  device  for  experimentation 
in  the  advanced  undergraduate  or  beginning  graduate  laboratory  was 
considered. 

Molecular    Structure    of   Thyroxine    by    X-Ray    Crystallography.    L.    K. 

Steinrauf,  0.  Seely,  J.  A.  Hamilton  and  J.  M.  H.  Pinkerton,  Indiana 
University  Medical  Center. — The  molecular  structure  in  the  crystal 
form  of  the  thyroid  hormone  thyroxine  was  determined  by  single  crystal 
x-ray  diffractometry  using  the  Supper-Pace  Autodiffractometer.  The 
structure  was  found  to  consist  of  planar  layers  of  the  hormone  sepa- 
rated by  layers  of  water  molecules,  all  connected  by  a  highly  complicated 
network  of  hydrogen  bonds.  A  possible  charge  transfer  bond  exists  be- 
tween  molecules    of   thyroxine.    The    hydrogen    bonds    and    the    charge 

357 


358  Indiana  Academy  of  Science 

transfer  bond  provide  the  means  for  speculation  on  the  manner  in  which 
thyroxine  can  bond  to  blood  serum  proteins. 

Electron  Paramagnetic  Resonance  Studies  on  The  Magneli  Phases  of  the 
Titanium-Oxygen  System.  1  John  F.  Houlihan  and  L.  N.  Mulay, 
Pennsylvania  State  University. — Several  transition  metal  oxides  exhibit 
a  number  of  stable  phases  with  variable  stoichiometry,  different  from 
that  predicted  by  simple  valence  considerations  of  the  cation.  These 
oxides  have  well-defined  crystal  structures  and  are  known  as  Magneli 
phases. 

Magnetic  susceptibility  studies  by  Mulay  and  others  on  the  phases 
of  the  titanium-oxygen  system  described  by  Tin02n-i  and  recent  elec- 
trical conductivity  data  by  Bartholomew  of  the  Materials  Research 
Laboratory  have  revealed  several  interesting  semi-conductor  to  metal 
transitions. 

In  this  paper,  typical  exploratory  electron  paramagnetic  resonance 
spectra  studied  as  a  function  of  temperature  are  presented  for  the  fol- 
lowing oxides:  TLO5,  T^Ot,  TioOu,  and  TiTOw.  The  electron  paramag- 
netic resonance  data,  in  general,  have  confirmed  the  transitions  previously 
observed  by  other  means. 

Proposed  lines  of  interpretation  correlating  electron  paramagnetic 
resonance  parameters  (such  as  g  values,  asymmetry  of  line  shapes,  etc.) 
with  the  magnetic  data  and  the  observed  transitions  were  presented. 

An  Evaluation  of  Relativistic  Thermodynamics.  Darryl  L.  Steinert, 
Hanover  College. — The  problem  of  the  accuracy  of  the  formulations  of 
relativistic  thermodynamics  proposed  by  Planck,  Eckart,  Ott,  and  Lands- 
berg  was  examined.  Due  to  the  lack  of  experimental  data  on  relativistic 
thermodynamic  systems,  it  is  not  possible  to  compare  predictions  made 
by  the  various  formulations  with  experimental  data.  But,  using  the 
process  of  evaporation  as  a  model,  I  found  that  it  is  possible  to  study 
the  consistency  between  the  transformation  laws  for  temperature  and 
for  mechanical  energy.  The  result  obtained  was  that  only  Ott's  formula- 
tion is  free  of  contradiction. 

Ott's  proposed  transformation  laws  were  further  evaluated  in 
terms  of  their  compatibility  with  relativistic  formulations  of  fluid  dy- 
namics and  statistical  mechanics.  Compatibility  is  to  be  expected  be- 
cause thermodynamics,  fluid  dynamics,  and  statistical  mechanics  are 
compatible  in  their  non-relativistic  formulations. 

The  lack  of  contradiction  between  Ott's  formulation  and  the  trans- 
formation law  for  mechanical  energy,  and  its  compatibility  with  formu- 
lations of  relativistic  fluid  dynamics  and  statistical  mechanics,  gave 
support  to  a  conclusion  that  Ott's  formulation  of  relativistic  thermo- 
dynamics is  correct. 


1  We  acknowledge  Prof.  W.  B.  White  for  providing  samples  and  Mr.  W.   J.   Danley  for 
assistance.  This  work  was  initially  sponsored  by  AEC  contract  AT(30-1  )-2581. 


Physics  359 

Physical  Oceanography  in  Indiana:  A  Study  of  Horse  Shoe  Lake.  Ralph 
A.  Llewellyn,  Rose  Polytechnic  Institute. — During  the  spring  of  1969, 
a  thorough  experimental  oceanographic  study  was  conducted  of  a  large 
artificial  lake  in  west-central  Indiana.  Initiated  as  an  educational 
exercise  for  a  group  of  67  science  and  engineering  students,  the  study 
developed  into  an  integrated  recording  and  analysis  of  7  parameters  of 
the   lake  over  a  several  week  period. 

The  results  of  the  study  included  a  computer-drawn  contour  map 
of  the  bottom,  the  discovery  of  an  unexpected  region  of  "dead"  water, 
evidence  for  a  sub-surface  current,  and  the  formulation  of  plans  to  moor 
continuous  recording  gear  in  the  lake.  This  concentrated  study  could 
well  serve  as  a  prototype  for  such  investigations  by  other  colleges. 


NOTES 

Polarized  "He+  Ion  Source  for  the  Indiana   University  Cyclotron.  J.   H. 

Hettmer,  Indiana  University. — An  optically  pumped  polarized  Heli- 
um-3  ion  source,  similar  to  that  designed  and  constructed  at  Rice  Uni- 
versity (1),  is  being  built  for  use  with  the  cyclotron  now  under  construc- 
tion at  Indiana  University  (2,  3,  4).  This  cyclotron  will  be  particularly 
well-suited  to  this  purpose  due  to  its  unique  property  of  accepting 
low-energy  positive  ions  at  ground  potential  for  acceleration  to  high 
energies.  This  is  of  interest  because  experimental  data  concerning  the 
scattering  of,  and  nuclear  reactions  induced  by,  these  particles  are 
extremely  difficult  to  obtain  by  any  other  technique. 

Previous  work  on  this  type  of  ion  source,  together  with  adaptations 
and   improvements   associated    with   this   application   was    described. 


Literature  Cited 

1.  Findley,  D.  O.,  S.  D.  Baker,  E.  B.  Carter,  and  N.  D.  Stockwell.  1969.  A  Polarized 
3He+   Ion  Source.  Nucl.  Instr.  and  Method.  71  :125-132. 

2.  Sampson,  M.  B.,  M.  E.  Rickey,  B.  M.  Bardin,  and  D.  W.  Miller.  1967.  Planned  200- 
MeV  Indiana  University  Cyclotron:  Properties  and  Unique  Features.  Proc.  Indiana 
Acad.  Sci.  77:347-348. 

3.  Rickey,  M.  E.,  M.  B.  Sampson,  B.  M.  Bardin,  and  D.  W.  Miller.  1966.  Proposed 
Indiana  University  200-MeV  Multi-Particle  Variable  Energy  Cyclotron  Facility. 
I.E.E.E.  Trans.  Nucl.  Sci.  NS-13  :464-468. 

4.  Rickey,  M.  E.,  M.  B.  Sampson,  and  B.  M.  Bardin.  1969.  General  Design  Features  of 
the   Indiana   University   200-MeV   Cyclotron.    I.E.E.E.    Trans.    Nucl.    Sci.    NS-16  :397-414. 


360  Indiana  Academy  of  Science 

Non-local  Contributions  to  the  Cyclotron  Absorption  Spectrum  for  a 
Single  Valley  Semiconductor  Model.  Uwe  J.  Hansen,  Indiana  State 
University,  and  James  L.  Hazelton,  Oklahoma  State  University. — In 
a  material  like  PbTe  for  which  at  35  Ghz  the  skin  depth  is  of  the  same 
order  of  magnitude  as  the  radius  of  carrier  orbits  at  cyclotron  reso- 
nance, semiclassical  calculations  of  the  cyclotron  absorption  coefficient 
indicate  that  absorption  maxima  should  be  observed  for  dielectric  anom- 
aly, rather  than  the  expected  cyclotron  resonance  conditions  (1). 
Experiments  indicate,  nevertheless,  that  some  structure  is  observed  at 
the  cyclotron  frequency  (1).  A  non-local  calculation,  adapted  from 
Hebel's  calculations  for  Bismuth  (2),  for  a  single  ellipsoidal  Fermi 
Surface  with  PbTe  parameters  was  carried  out  for  the  limited  case  of 
the  magnetic  field  orientation  along  the  major  symmetry  axis  and  the 
microwave  electric  field  parallel  to  the  magnetic  field.  This  calculation 
indicated  an  absorption  maximum  at  the  cyclotron  frequency  (3).  More 
extensive  calculations  are  in  progress. 


Literature  Cited 

1.  Hansen,  U.  J.,  J.  H.  Gardner,  and  K.  F.  Cuff.  1966.  Analysis  of  cyclotron  absorption 
in  PbTe.  Bull.  Amer.  Phys.  Soc.  11(5)  :755. 

2.  Hebel,    L.    C.    1965.    Cyclotron    resonance    in    Bi    with    slightly    anomalous    skin    effect. 
Phys.  Rev.  138(6A)  :1641-1649. 

3.  Hazelton,     J.     L.     1969.     Semiclassical    calculation     of    cyclotron     absorption     in     lead 
telluride.  Unpublished  M.S.  Thesis,  Indiana  State  Univ.,  Terre  Haute. 


Semiconductors  Produced  by  Doping  Oxide-glasses 
With  Ir,  Pd,  Rh  or  Ru 

C.  C.  Sartain,  Indiana  State  University- 


Abstract 

Semiconductors  were  produced  by  diffusion  doping  oxide  glasses  with  more  than 
1  wt  %  of  Ir,  Pd,  Rh  or  Ru  and  by  implanting  40  kilovolt  Ir  ions  into  several  oxide 
glasses.  Hall  mobility  at  300°  K  and  78°  K  was  less  than  0.005  cm2/volt  sec.  Mobility 
from  impurity  concentration  and  conductivity  was  0.001  cm2/volt  sec.  Conductivity  was 
not  ionic.  Enough  direct  current  was  passed  through  one  sample  to  have  plated  out  a 
million  times  as  many  Ir  ions  as  were  in  the  sample  with  no  change  in  conductivity. 
X-ray  small  angle  scattering  indicates  that  conductivity  was  not  due  to  electron  hopping 
between  conducting  islands.  Conductivity  was  ohmic  at  300°  K  and  was  field  dependent 
below  4°  K.  Thermoelectric  power  of  12  to  25  microvolts  per  degree  relative  to  copper 
indicated  hole  conduction.  The  material  absorbed  throughout  the  visible  region.  For  con- 
stant firing  temperature  and  time  at  a  constant  oxygen  pressure  conductivity  increased 
as  impurity  concentration  increased  and  varied  from  10-3  to  10J  reciprocal  ohm-cm. 
For  a  constant  concentration  of  platinum-metal  ions  and  for  equilibrium  firing,  the 
conductivity  was  directly  proportional  to  the  oxygen  pressure.  Firing  in  hydrogen  to 
remove  oxygen  reduced  conductivity  at  the  rate  of  two  carriers  per  oxygen  atom 
removed.  These  data  indicate  that  the  frozen-in  valence  states  of  the  platinum-metal  ions 
serve  as  the  source  of  the  carriers. 


Introduction 

Oxide  glasses  were  caused  to  become  amorphous  semiconductors 
by  heavily  doping  them  with  iridium,  ruthenium,  rhodium  or  palladium. 
The  conductivity  is  between  10  ~'5  and  102  ohms-1  cnr1. 

Most  metals  will  oxidize,  dissolve  into  and  become  a  part  of  molten 
oxide  glass.  They  are  so  chemically  active  with  oxygen  that  all  of  their 
electrons  form  binding  orbitals.  They  are  unable  to  retain  charges  which 
can  act  as  donors  or  acceptors.  Thus,  no  one  has  reported  doping  oxide 
glasses  with  ordinary  dopants  such  as  boron,  phosphorous,  etc. 

Certain  members  of  the  platinum  metal  family,  while  not  inert,  are 
relatively  inactive  chemically.  These  metals  have  more  than  one  oxi- 
dation state  and  they  can  retain  charges  which  contribute  to  the  elec- 
trical conductivity. 


Conductivity  vs.  Temperature 

Sample  resistance  was  measured  as  a  function  of  temperature 
from  1.48°K  to  1150°K.  For  several  hundred  samples,  the  d-c  resistance 
was  measured  by  the  volt-amp  method  for  about  20  different  tempera- 
tures between  the  triple  point  of  nitrogen  and  200  °C. 

Typical  data  are  shown  in  Figure  1.  There  exists  a  striking  simi- 
larity between  the  shapes  of  these  graphs  and  those  shown  in  Pearson 
and  Bardeen's  Figure  4A,  "Resistivity  of  Silicon-Boron  Alloys  as  a 
Function  of  Inverse  Absolute  Temperature''  (7). 

361 


362 


Indiana  Academy  of  Science 


10  Meg 


Figure  1.  Logarithm  of  resistance  vs.  reciprocal  absolute  temperature  for  various  con- 
centrations of  iridium  in  an  oxide  glass.  These  curves  are  typical  examples  of  platinum 
metals  doped  into  any  one  of  many  kinds  of  oxide  glasses. 


At  high  temperatures  the  intrinsic  resistance  of  one  sample  of 
oxide  glass  was  shown.  For  low  doping  and  low  temperature  the  resist- 
ance is  given  by  R  =  A  exp  (W/kt).  For  high  doping  the  carriers  seem 
to  be  degenerate.  At  low  temperature  as  the  doping  increases,  the  slopes 
of  the  curves  decrease  monotonically  as  far  into  the  "degenerate"  re- 
gion as  it  was  carried. 

It  is  obvious  from  the  shape  of  these  curves  that  at  low  doping,  the 
conduction  cannot  be  metallic. 

The  conduction  cannot  be  ionic  because  there  was  no  transport  of 
ions.  Ionic  conduction  in  glass  usually  leads  to  a  depletion  of  ions  from 


Physics 


363 


some  volume  and  a  change  in  resistance  as  current  is  passed  through 
the  unit.  Tests  of  many  hours  duration  on  many  samples  showed  that 
the  resistance  did  not  change  as  much  as  one  part  in  10r>  after  enough 
charge  had  passed  through  the  samples  to  have  plated  out  a  million 
times  as  many  iridium  atoms  as  were  in  the  sample. 

There  was  no  electrolysis  of  metals  nor  of  oxygen.  In  some  cases 
the  only  element  common  to  different  samples  was  oxygen.  For  example 
a  lead-borate-oxide  glass  doped  with  iridium  conducted,  yet  so  did  an 
alkali-silicate-oxide  glass  doped  with  ruthenium.  In  every  case  at  least 
half  of  the  atoms  in  the  system  were  oxygen.  In  no  case  was  there  any 
evolution  of  oxygen  at  the  terminals. 

The  resistance  at  300 °K  was  ohmic  over  more  than  6  orders  of 
magnitude  between  100  /*v  per  cm  and  500  v  per  cm.  Conductivity  did 
depend  on  the  field  below  4°K.  The  data  here  are  too  meager  to  estab- 
lish the  relationship  between  conductivity  and  electric  field  with  accu- 
racy. 


5.080 


5.070 


5.060 


5.050 


5.040 


5.030 


5.020 


1000/T 

Figure  2.  Logarithm  of  resistance  vs.  reciprocal  absolute  temperature  for  sample 
I  -  1  -  C  -  D  of  Figure  1.  Note  how  the  experimental  points  fit  a  smooth  curve  even  at 
greatly  expanded  vertical  scale. 


364 


Indiana  Academy  of  Science 


The  conduction  was  not  due  to  electron  hopping  between  conduct- 
ing islands  with  dimensions  of  the  order  of  100  A  as  described  by  Neu- 
gebauer  and  Webb  (6).  Many  X-ray  small  angle  scattering  experiments 
were  performed  in  search  of  this  effect.  In  some  cases  particles  of  this 
size  were  found,  but  there  was  no  correlation  between  particle  size 
and  resistivity  of  the  sample.  Based  on  their  model,  Neugebauer  and 
Webb  (6)  predicted  the  magnitude  of  the  resistivity  and  its  temperature 


0  Meg. 


1  Meg. 


100K 


1.80  x 

10-3   ev 

a 

0-3  ev. 

.3 

1  /  T 


Figure  3.  Logarithm  of  resistance  vs.  reciprocal  absolute  temperature  for  a  sample  with 
a  room  temperature  resistance  of  100  kilo  ohms.  The  slope  of  this  curve  in  the  liquid 
nitrogen  is  2.73  X  lO-3  ev.  The  slope  in  the  liquid  helium  range  appears  to  be 
1.80  XJ0-3  ev.  The  point  at  148°  K  is  below  the  extrapolated  straight  line.  This  prob- 
ably indicates  that  the  curve  is  not  truly  straight  anywhere,  but  drops  off  continuously 
as  predicted  by  Mott  (5).  See  text. 


Physics  365 

dependence.  The  resistivity  in  this  study  was  several  orders  of  magni- 
tude too  low  to  have  been  due  to  this  effect  and  the  temperature  depend- 
ence differed  from  that  predicted. 

Ir  02  and  Ru  02  single  crystals  are  metallic  conductors  (8),  but  they 
play  no  part  in  the  conduction  in  these  Cermets. 

The  conduction  was  due  to  the  presence  of  metals  belonging  to  the 
platinum  family.  The  glasses  did  not  conduct  (that  is,  the  conductivity 
was  many  orders  of  magnitude  less)   unless  these  metals  were  present. 

There  were  no  compensating  metals  present. 

Figure  2  is  the  curve  which  appeared  flat  in  Figure  1.  The  points 
fit  a  smooth  curve  and  at  low  temperature  the  line  is  straight  with  a 
slope  of  1.90  x  10"8  ev. 

Figure  3  shows  how  one  sample  behaves  at  liquid  helium  tempera- 
tures. The  resistance  of  this  sample  was  105  ohms  at  300  °K.  In  the 
liquid  nitrogen  range  the  slope  was  2.73  milli-electron  volts.  The  curve 
as   drawn   shows   a  slope   of   1.80   milli-electron  volts   below   4°K. 

However,  the  point  at  1.48  °K  is  definitely  below  the  curve. 

This  curve  exhibits  the  fall-off  in  slope  of  the  In  p  vs  1/T  at  low 
temperature  as  predicted  by  Mott  (5).  Probably  at  no  point  is  the 
curve  truly  a  straight  line.  It  appears  to  fit  a  straight  line  over  limited 
temperature  ranges,  but  its  slope  probably  decreases  continuously.  This 
change  in  slope  may  occur  for  all  of  our  samples. 


Hall  Effect 

In  no  sample  could  a  Hall  voltage  as  large  as  10'9  volts  be  detected 
at  any  temperature.  For  some  samples,  10~9  volts  would  represent  a 
mobility  of  about  3  x  10"8  cm2  per  volt  sec.  The  carrier  mobility  in  all  of 
our  samples  was  less  than  this  amount. 


Seebeck  Effect 

Thermoelectric  emf  was  measured  in  two  ways.  In  the  first  method, 
one  end  of  a  long,  slender  sample  was  fixed  in  an  ice  bath  while  the 
other  end  was  heated  to  various  temperatures.  In  a  variation  of  this 
method,  the  cold  end  was  kept  in  a  liquid  nitrogen  bath. 

In  the  second  method,  pt-pt  10  rh  thermocouples  were  welded  to 
each  end  of  a  Cermet  sample  about  5  cm  long.  The  sample  was  inserted 
to  a  series  of  positions  of  increasing  temperature  in  a  tube  furnace  so 
that  the  ends  of  the  sample  were  at  different  temperatures.  The  tem- 
perature of  each  end  could  be  measured.  The  Seebeck  voltage  could  be 
measured  between  the  two  platinum  wires. 

In  every  case,  the  Seebeck  Voltage  indicated  hole  conduction. 


366 


Indiana  Academy  of  Science 


Conductivity  vs.  Metal  Concentration 

The  conductivity  as  a  function  of  concentration  of  ruthenium  in  one 
kind  of  glass  fired  at  one  temperature  for  a  fixed  time  in  air  is  shown 
in  Figure  4.  Each  point  on  the  curve  represents  five  samples.  The  bars 
cover   the    maximum    and    the   minimum    resistance    of   the    group.    The 


300  - 


LU 

o 

< 
co 

CO 
UJ 


ID 

o 


100  - 


2  5  10 

Wt  %    Ru    in     Glass     "  A 


FIGURE  4.  Logarithm  of  conductivity  at  :i00°  K  vs.  logarithm  of  weight  percent 
ruthenium  in  glass  A.  The  Cermets  were  fired  in  air  through  a  tunnel  kiln  so  that  each 
sample  was  exposed  to  the  same  temperature-time  profile.  For  these  samples 
a  =   CW*.44  up  to  about  6  wt%  where  devitrification  occurs. 


Physics 


367 


curve  is  fitted  by  a  power  law  up  to  the  point  where  devitrification 
begins.  The  data  will  not  fit  an  exponential  dependence  of  conductivity 
on  concentration. 

For  Glass  "A"  containing  lead,  silicon,  Cadmium  and  aluminum 
oxides,  devitrification  occurs  at  about  6  wt  %  Ru.  Devitrified  samples 
look  like  black  emery  paper. 


Log   CONCENTRATION      GLASS    B 


100 


Figure  5.  Logarithm  of  conductivity  at  300°  K  vs.  logarithm  of  concentration  of 
ruthenium  in  glass  B.  The  samples  were  fired  in  air  for  constant  times,  but  for 
different    temperatures.    The    exponents    of    the    different    curves    vary    from    2.6    to    IS. 3. 


368  Indiana  Academy  of  Science 

For  a  different  glass,  labeled  B,  the  effect  of  varying  the  firing 
temperature  is  shown  in  Figure  5.  Firing  time  was  a  constant.  Devitri- 
fication occurred  at  higher  Ru  concentration  for  this  glass. 

Resistance  vs  Firing  Time  in  Air 

If  the  number  of  metal  atoms,  the  external  oxygen  concentration 
and  the  firing  temperature  were  held  constant  and  a  number  of  similar 
samples  are  fired  for  different  times,  the  resistance  appeared  to  decrease 
exponentially  with  time  to  approach  a  constant  value. 


Oxygen  Concentration 

Figure  6  shows  the  conductivity  of  a  series  of  similar  samples  fired 
for  the  same  time  and  temperature  at  different  pressures  of  oxygen.  The 
arbitrary  line  with  a  slope  of  one  drawn  on  the  graph  indicates  that  the 


100 


0.3 


1.0 

OXYGEN 


3.0 
PRESSURE  , 


10.0 
INCHES  Hg 


300 


Figure  6.  Logarithm  of  conductivity  at  300°  K  vs.  logarithm  of  oxygen  pressure.  The 
samples  were  fired  under  a  controlled  atmosphere  of  commercial  oxygen  for  a  constant 
temperature  and  time.  The  arbitrary  straight  line  has  a  sloj)e  of  1.  Each  bar  on  the 
graph  covers  the  measured  range  of  10  samples  which  were  not  precisely  uniform  in 
thickness.  Later  techniques  improved  the  reproducibility  of  the  sample  dimension,  but 
this  experiment  was  not  repeated. 


Physics 


M) 


conductivity  varied  directly   as   the  number  of  oxygen  atoms   diffusing 
into  the  sample  and  reacting  with  the  ruthenium. 

Firing  in  Hydrogen 

An  especially  large  sample  was  made  for  thermogravimetric  analy- 
sis. A  Cermet  mix  containing  3.60  wt  %  Ru  was  screened  and  fired  on 
a  substrate.  The  measured  mass  of  the  sample  was  0.303  g  and  its  re- 
sistance at  300 °K  was  23  kilo  ohms.  The  sample  was  fired  in  hydrogen 
for  30  minutes  at  500  °C. 

The  sample  lost  a  mass  of  760  jxg  and  its  resistance  at  300  °C 
increased  to  704  kilo  ohms. 

A  previous  test  showed  that  the  hydrogen  firing  at  500°  C  did  not 
affect  the  glass. 

At  500  °C,  apparently  hydrogen  converted  Ru  02  to  Ru-O:..  The  water 
formed  evaporated  from  the  sample.  Each  Ru  O2  presumably  contributes 
one  hole  for  conduction.  Conversion  of  each  Ru  (X  to  V2  (Ru20;f)  or 
removing  one-half  of  an  oxygen  atom  from  each  Ru  atom  removes  one 


Conductance 

Micromhos 

47.8 


Measured  43.5 


Oxygen  Wt. 


No.  of 
Holes  at 


Measured  1.4 


ru  07       Micrograms     300  K 
864  — 6.47xl019 


5.91x10 


19 


5.72xl019 
Holes 


0.19x10 


19 


Ru203 


0   —     0 


Figure    7.      Schematic    diagram    showing    how    removal    of    oxygen    from    one    sample    of 
Cermet  removes  holes  and  correspondingly  reduces  conductance. 


370  Indiana  Academy  of  Science 

hole  thus  removing  one  charged  carrier.  The  Ru  in  the  sample  was  3.6% 
of  0.303  gm  or  10.9  mg,  or  6.47  X  1019  atoms.  The  loss  of  760  fig  of 
oxygen  represented  a  loss  of  2.86  X  1019  atoms  of  oxygen,  or  5.72  X  1019 
holes  (two  holes  per  oxygen  atom  removed).  The  conductance  became 
1/704  kilo  ohms  or  1.4  micromhos.  The  loss  in  conductance  was  43.5  — 
1.4  or  42.1  micromhos. 

In  Figure  7,  the  quantity  of  oxygen  required  to  convert  Ru2  03  to 
Ru  02  is  plotted  schematically  on  the  vertical  scale.  For  this  sample, 
which  contained  10.9  mg  of  Ru,  864  fig  of  oxygen  was  required  for  the 
conversion. 

On  the  left,  conductance  of  the  sample  in  micromhos  was  plotted, 
and  on  the  far  right  the  number  of  holes  in  the  sample  at  300  °K. 

When  the  sample  was  originally  fired  in  air,  the  ruthenium  was 
oxidized  to  the  level  such  that  its  measured  conductance  was  43.5  mi- 
cromhos. The  sample  was  then  fired  in  hydrogen.  This  firing  removed 
a  measured  quantity  of  760  fig  of  oxygen  or  5.72  X  1019  holes  and  re- 
duced the  conductance  to  the  measured  value  of  1.4  micromhos.  The 
assumption  that  removing  5.72  X  1019  holes  removed  42.5  micromhos 
of  conductance  gave  the  scale  of  the  graph.  For  this  sample  one  mi- 
cromho  was  equivalent  to  0.1345  X  10lfi  holes,  or  10'"  holes  was  equiva- 
lent to  7.44  micromhos.  Now  the  measured  level  of  1.4  micromhos  above 
zero  was  set  to  be  0.19  X  1010  holes. 

Firing  this  sample  in  air  at  the  temperature  and  time  did  not 
oxidize  all  of  the  Ru  to  Ru  Oz.  Only  5.91  to  1019  atoms  of  Ru  out  of  a 
possible  6.47  X  1019  or  91.3%  were  combined  as  Ru  O2.  The  remainder 
was  oxidized  only  to  the  Ru2  On  level. 

For  this  sample  the  carrier  concentration  was  11.2  X  1020  Ru  atoms 
per  cm3  times  91.3%-  or  10.2  X  1030  holes  per  cnr.  <r  =  0.204  fiJ  cm1.  The 
mobility  fi  =  a/ne  —  1.25  X  10~3  cmVvolt  sec. 


Other  Physical  Properties 

No  magneto-resistance  was  observed  up  to  14  kilogauss.  No  photo- 
conductivity was  observed  at  liquid  nitrogen  temperatures  or  above. 
A  very  small  increase  in  conductivity  was  noted  when  light  from  an 
incandescent  bulb  (microscope  illuminator)  was  focused  on  a  sample 
bathed  in  liquid  helium.  This  was  attributed  to  the  heating  of  the  sample, 
not  to  photoconductivity  of  the  sample.  The  samples  absorbed  energy 
throughout  the  visible  region. 

The  kind  of  glass  does  affect  the  conductivity,  suggesting  that  the 
conductivity  might  be  proportional  to  the  square  of  the  dielectric  con- 
stant. However,  in  these  experiments,  the  effect  of  the  dielectric  con- 
stant was  not  isolated  from  other  effects,  particularly  that  due  to  the 
temperature  of  firing,  i.e.,  that  of  quenching  the  oxide  into  a  particular 
frozen-in  valance  state.  Obviously  this  latter  effect  depends  on  the 
softening  point  of  the  glass. 


Physics  371 

No  effects  such  as  rectification  were  observed  which  could  be  inter- 
preted as  due  to  the  contacts  or  terminations. 

Noise  was  proportional  to  1 /frequency  for  these  samples  at  room 
temperature.  The  noise  was  not  measured  at  any  other  temperature. 

An  unsuccessful  attempt  was  made  to  measure  mobility  directly  (2, 
3)  by  injecting  carriers  and  measuring  the  time  to  travel  a  microscopic 
distance.  This  experiment  indicated  the  lifetime  was  very  short.  If  it  is 
assumed  that  /*  r=  €  r/m*  with  p,  =  10-3  and  m*  =  m,  T  is  indeed  very 
small.  Smaller  values  of  m*  require  even  smaller  values  of  T. 

Conductivity  in  alternating  current  below  4°K  is  independent  of 
frequency  up  to  10  kc/sec. 

It  is  interesting  to  compare  Sb-doped  Ge  (1)  with  Ir-doped  glass. 

I  noted  that  at  the  doping  level  just  high  enough  to  make  Fritzsche's 
Sb-doped  Ge  become  degenerate,  N  =  2.7  X  1017  atoms  per  cm3,  p  300  = 
0.015  ohm  cm  and  fi  =  1500  cnrVvolt  sec.  At  the  doping  level  required  to 
make  Ir  in  glass  "degenerate,"  N  =  1.2  X  10"',  p  =  2.1  and  ix  —  1.5  X 
lO-3.  The  ratio  of  N  is  about  5000,  that  of  p  is  about  200  and  that  of  y, 
is  about  10-6. 

Ion  Implantation1 

Forty  kilovolt  ions  of  Ir-193  were  implanted  into  fused  Si02  and  five 
other  glasses.  The  doping  levels  were  1014,  10]5  and  10*16  ions  per  cma. 
The  Lindhard  depth  of  penetration  and  the  deviation  of  the  range  for 
40  kv  Ir-193  into  SiO-  were  computed  by  Gibbons  and  Johnson  (4)  to  be 
222a  with  a  spread  of  ~  22a.  The  electrical  properties  of  the  ion  im- 
planted material  is  about  the  same  as  that  of  the  diffused  material.  If  it 
is  assumed  that  half  of  the  Ir-193  ions  do  stop  in  a  layer  only  45a 
thick,  if  the  quantum  mechanical  effect  of  such  a  thin  conductor  is 
neglected,  and  if  the  conductivity  based  on  the  behavior  of  the  same 
number  of  ions  diffused  into  the  glass  is  estimated,  the  measured  con- 
ductivity is  obtained.  This  is  good  evidence  that  the  holes  do  not  reflect 
or  scatter  from  the  surface  of  the  conductor  and  therefore  they  hop 
short  distances  in  short  lifetimes,  Also,  the  conductivity  of  the  various 
glasses  were  crudely  proportional  to  the  square  of  the  dielectric  constant. 

It  appears  that  when  the  40  kv  Ir  ions  stop  in  oxide  glass,  local 
heating  and  other  effects  permit  the  ions  to  form  chemical  bonds  with 
neighboring  oxygen  atoms  and  that  Ir  introduced  by  ion  bombardment 
forms  the  same  bonds  as  Ir  diffused  into  the  glass  at  high  temperature. 

Discussion 

These  experiments  indicated  that  each  tetravalent  atom  of  one  of 
the  platinum  metals  in  an  oxide  glass  can  contribute  a  hole  which 
participates  in  the  conduction  mechanism.  The  trivalent  atoms   seemed 


1  We  are  indebted  to  Mr.   G.  Alton  and  the  Stable  Isotopes  Division   of  the  Oak  Ridge 
National  Laboratory  for  implanting  the  ions. 


372  Indiana  Academy  of  Science 

to  take  no  part  in  the  conduction.  The  valence  state  was  fixed  at  high 
temperature  by  adding  or  removing  oxygen  from  the  platinum  metal 
ion.  When  the  sample  was  cooled  this  valence  state  was  frozen-in.  Since 
the  rate  of  cooling  was  rapid,  this  may  be  considered  a  quenching 
effect.  The  number  of  platinum  metal  ions  frozen  in  the  tetravalent  state 
determined  the  number  of  holes  available  to  carry  the  current,  thus  fix- 
ing the  resistance  (or  conductance)  of  the  sample.  The  number  of  car- 
riers was  easily  changed  by  changing  the  number  of  atoms  in  the 
tetravalent  state.  This  could  be  done  only  at  high  temperatures. 

The  resistance  could  be  changed  by  changing  the  total  number  of 
platinum  metal  atoms  in  a  sample.  However,  in  a  sample  already  manu- 
factured which  contains  a  fixed  number  of  platinum  metal  atoms,  the 
number  of  tetravalent  atoms  could  be  changed  by  increasing  or  de- 
creasing the  oxygen  content  of  the  sample.  This  could  be  done  by  firing 
for  a  different  time  at  a  given  oxygen  pressure  and  temperature,  by 
firing  in  an  oxidizing  or  reducing  atmosphere  or  even  by  firing  at  a 
different  temperature. 

At  high  temperatures  the  oxides  of  the  platinum  metals  were  vola- 
tile, so  only  a  limited  temperature  range  was  available  for  work. 

Under  certain  conditions  semi-conducting  glass  could  be  made 
from  osmium.  However,  osmium  oxide  was  very  volatile  and  poisonous, 
and  it  is  difficult  to  process  the  glass  without  losing  osmium. 

In  these  experiments  the  conductivity  apparently  depended  on 
the  state  of  one  of  the  electrons  belonging  to  the  d-shell  of  the  platinum- 
metal  atom.  If  this  electron  was  bound  to  an  oxygen  atom,  there  existed 
a  hole  in  the  d-shell  (band?)  which  may  act  as  a  trap  or  otherwise  par- 
ticipate in  conduction.  If  there  was  no  oxygen  atom  nearby  to  attract 
the  electron,  it  occupied  its  ground  state  in  the  d-shell  and  no  hole 
was  available. 

The  conductivity  was  a  discontinuous  function  of  concentration.  If 
the  concentration  was  less  than  about  1  wt  %  or  about  3  X  1030  Pt-metal 
atoms  per  cm3  the  sample  conductivity  decreased  by  many  orders  of 
magnitude  to  that  of  the  intrinsic  conductivity  of  the  glass.  At  this  con- 
centration the  average  volume  occupied  by  only  one  ion  was  about  3 
X  10 -*  cm3.  Thus,  at  the  average  distance  between  ions  of  about  15a,  the 
samples  change  from  the  conducting  to  the  non-conducting  state. 


Physics  373 

Literature  Cited 

1.  Fkitzsche,   H.    1958.   Resistivity   and  Hall   coefficient  of  antimony-doped  germanium   at 
low  temperatures.  J.  Phys.  and  Chem.  of  Solids.  6  :69-80. 

2.  Haynes,    J.    R.,    and    W.    Shockley.    1951.    The    mobility    and    life    injected    holes    and 
electrons  in  germanium.  Phys.  Rev.  81  :835-843. 

3.  Haynes,   J.   R.,  and  W.   C.   Westphal.   1952.   The  drift  mobility  of  electrons  in  silicon. 
Phys.  Rev.  85  :680. 

4.  Johnson,    W.    S.,   and   J.    F.    Gibbons.    1966.    Statistical   range    distribution    of   ions    in 
single  and  multiple  element  substrates.  Applied  Phys.  Letters  9  :321-322. 

5.  Mott,    N.    F.    1968.    Conduction    in    glasses    containing    transition    metal    ions.    J.    Non- 
Crystalline  Solids.  1  :36-52. 

6.  Neugebauer,   C.   A.,  and  M.   B.   Webb.   1962.   Electrical  conduction   mechanism   in  ultra- 
thin,  evaporated  metal  films.  J.  Appl.  Phys.  33  :74-82. 

7.  Pearson,  G.  L.,  and  J.  Bardeen.  1949.  Electrical  properties  of  pui-e  silicon  and  silicon 
alloys  containing  boron  and  phosphorus.  Phys.  Rev.  75  :865-883. 

8.  Ryden,   W.   D.,   A.   W.   Lawson,   and  C.   C.   Sartain.   1968.   Temperature   dependence   of 
the    resistivity    of    RuO-    and    Ir02.    Phys.    Letter    26A  :209-210. 


PLANT  TAXONOMY 

Chairman:  Jack  E.  Humbles,  Indiana  University 
Jeanette  C.  Oliver,  Ball  State  University,  was  elected  Chairman 

for  1970 

ABSTRACTS 

Eocene  Euphorbiaceous  Fruits.  Neal  E.  Lambert  and  David  L.  Dil- 
cher,  Indiana  University. — A  recent  study  of  a  population  of  20  well- 
preserved  fossil  fruits  has  shown  that  these  fruits  have  probable  af- 
finities with  the  Euphorbiaceae.  These  fruits  were  collected  from  Eocene 
clay  deposits  in  Henry  County,  Tennessee.  In  1922,  E.  W.  Berry  de- 
scribed similar  fruits  collected  from  Eocene  deposits  in  western  Ten- 
nessee and  assigned  these  to  the  genus  Monocarpellites.  He  tentatively 
referred  the  genus  to  the  Malvaceae  but  expressed  uncertainty  concern- 
ing its  botanical  affinities.  M.  E.  J.  Chandler  described  similar  fruits 
from  Lower  Tertiary  deposits  of  Egypt,  Isle  of  Wight,  Sussex  of 
England.  Chandler  referred  her  material  to  two  genera,  Wetherellia  and 
Palaeow ether ellia,  and  placed  the  genera  in  the  Euphorbiaceae.  The 
fruits  from  western  Tennessee  most  resemble  the  genus  Palaeowether- 
ellia;  however,  there  are  some  consistent  differences.  Both  the  external 
features  and  internal  anatomical  structure  of  the  fossil  material  indi- 
cate an  affinity  with  some  of  the  large  capsules  found  in  extant  woody 
genera  of  the  Euphorbiaceae.  The  fossil  fruits  are  septicidal  capsules, 
24-39  mm  in  diameter  with  7-10  locules  which  dehisce  radially  exposing 
the  seeds.  Sections  showing  internal  cellular  detail  have  been  prepared 
and  will  be  discussed  in  addition  to  the  overall  aspects  of  the  fruits. 

Morphology  and  Taxonomy  of  Fossil  Fungal  Spores.  M.  V.  Sheffy  and 
D.  L.  Dilcher,  Indiana  University. — The  Eocene  clay  deposits  of  west- 
ern Tennessee  and  Kentucky  contain  large  numbers  of  preserved  dis- 
persed fungal  spores.  Extensive  research  has  been  done  on  the  fossil 
leaves  associated  with  these  clays  and  one  study  by  Dilcher  (1965)  re- 
ports numerous  epiphyllous  fungi  found  on  these  fossil  leaves.  A  large 
assemblage  of  dispersed  fungal  spores  was  isolated  from  the  sediments 
by  zinc  bromide  flotation  and  mounted  in  glycerine  jelly.  Camera  lucida 
line  drawings  and  photographs  were  made  of  each  spore  type  to  ac- 
company the  spore  descriptions.  Distinct  morphological  characters  such 
as  shape,  size,  sculpture,  and  number  of  cells  and  pores  were  used  to 
delimit  14  genera  and  81  species.  They  were  classified  according  to  an 
artificial  system  of  nomenclature  followed  by  von  der  Hammen,  Rouse, 
Clarke  and  Elsik.  This  preliminary  taxonomic  work  is  necessary  for 
any  further  work  with  fungal  spores  as  part  of  a  complete  organic 
assemblage  or  as  marker  fossils  in  stratigraphic  correlations.  Many  of 
the  spores  found  are  similar  to  dispersed  spores  recorded  from  Cre- 
taceous-Recent sediments  of  North  America,  Europe  and  Africa.  Al- 
though some  spores  have  been  tentatively  assigned  to  modern  taxa  by 

375 


376  Indiana  Academy  of  Science 

Bradley,  Dilcher,  Wolf  and  Graham,  this  continues  to  be  a  difficult  task 
until  extensive  modern  reference  collections  are  required. 

The  Cultivated  Solanaceae  of  Ecuador.  Charles  B.  Heiser,  Jr.,  Indiana 
University. — The  family  Solanaceae  is  well  represented  among  the  plants 
cultivated  in  Ecuador.  Several  members  of  the  family  are  important 
food  plants,  including  such  well  known  ones  as  the  potato  (Solanum 
tuberosum)  and  peppers  {Capsicum  spp.) ,  as  well  as  a  number  of  lesser 
known  ones,  including  the  naranjilla  (S.  quitoense),  the  pepino  (S. 
muricatum)  and  the  tree  tomato  {Cyphomandra  crassi  folia) .  Most  of  the 
ornamentals  of  the  family  grown  in  the  temperate  zone  are  also  culti- 
vated there,  and  in  addition  a  number  of  shrubs  are  grown  for  their 
ornamental  value  (Solanum  spp.,  Iochroma  fuchsioides,  Streptosolen 
jamesonii,  and  Datura  spp.).  Various  species  of  the  genus  Datura  are 
also  employed  for  use  as  narcotics.  Chromosome  counts  were  obtained 
for  the  following:  Datura  arborea,  n=12;  D.  aurea  n=12;  D.  Can- 
dida, n=12;  Iochroma  fuchsioides,  n=ca.24;  and  Streptosolen  jame- 
sonii,  n=ll. 


Hookeriaceae  Species  and  Distribution  in  Africa,  Europe,  Asia, 
Australia  and  Oceania 

Winona  H.  Welch,  DePauw  University 

Abstract 

This  is  the  third  and  last  in  the  series  of  papers  on  the  distribution  of  the 
Hookeriaceae.  The  first  (4)  pertained  to  North  and  Central  America  and  West  Indies, 
and  the  second  (6)  to  South  America. 

No  Antarctic  or  cosmopolitan  species  of  Hookeriaceae  have  been  noted.  Endemism 
occurs  frequently  in  the  family.  An  observation  of  the  world  distribution  of  genera 
shows  that  nine  genera  are  known  only  from  one  area  and  twelve  genera  occur  in  more 
than  one  area  but  have  a  hemisphere  or  a  continental  distribution,  such  as  in  the 
Americas,  in  the  West  Indies  and  the  Americas,  or  in  Asia. 

In  the  progress  of  the  monographic  studies  in  the  Hookeriaceae  and 
due  to  the  publication  of  Volume  5  of  Index  Muscorum  (7),  the  follow- 
ing changes  and  additions  in  synonymy,  authors  of  epithets,  and  distri- 
bution have  been  made  since  publications  (4)  and  (6)'  Callicostella 
depressa  var.  rubella  (Besch.)  Wijk  &  Marg.  =  C.  depressa  (Hedw.) 
Jaeg. — Am  2-5;  *C.  filescens  (Besch.)  Jaeg. — Am  3;  C.  leonii  Ther.  = 
Pilotrichidium  leonii  (Ther.)  Crum  &  Bartr. — Am  3;  *C.  merkelii 
(Hornsch.)  Aongstr. — Am  5;  C.  strumulosa  (Hampe  &  Lor.)  Jaeg. — 
Am  4,  5;  *C.  subdepressa  (Besch.)  Kindb. — Am  5;  C.  wallisii  Fleisch. 
r=  nom.  nud.  — Am  2.  Crossomitrium  crugeri  C.  Mull.  =  C.  patrisiae 
(Brid.)  C.  Mull. — Am  1-6;  C.  herminieri  (Besch.)  Jaeg. — Am  2,  3. 
Cyclodictyon  blandum  (Lor.)  Kuntze  =  C.  varians  (Sull.)  Kuntze — Am 
1,  3,  5;  C.  pattens  (Mitt.)  Kuntze  =  C.  varians  (Sull.)  Kuntze — Am  1, 
3,  5;  *C.  regnellianum  (C.  Mull.)  Broth,  ex  Par. — Am  5;  C.  varians 
(Sull.)  Kuntze — Am  1,  3,  5.  Distichophyllum  densirete  Broth.  —  *Les- 
keodon  densiretis  (Broth.)  Broth. — Am  5;  *D.  fasciculatum  Mitt. — 
Afr  4,  Am  6,  Oc.  *Eriopus  haitensis  Crum  &  Steere — Am  3.  Hemiragis 
anrea  (Brid.)  Kindb. — Am  2-5.  Hookeria  viridula  Mitt.  =  Cyclodictyon 
viridulum  (Mitt.)  Kuntze — Am  4,  5;  H.  wallisii  C.  Mull.  =  nom.  nud. 
— Am  2.  Hookeriopsis  diffusa  (Wils.)  Jaeg. — Am  2,  4;  *H.  fissideyitoides 
(Hook.  &  Wils.)  Jaeg. — Am  3;  *H.  heteroica  Card,  has  recently  been 
collected  in  Mississippi,  previously  known  from  Mexico — Am  1 ;  *H, 
luteo-viridis  (Besch.)  Kindb. — Am  5;  *H.  perfulva  (C.  Mull.)  Broth. 
ex  Par. — Am  5;  *H.  websteri  Crum  &  Bartr. — Am  3.  Hypnella  diversi- 
folia  (Mitt.)  Jaeg.  =  Neohypnella  diversifola  (Mitt.)  Welch  &  Crum 
— Am  3-5.  Isodrepanium  lentulum  (Wils.)  Britt. — 1-5.  Lepidopilidium 
divaricatum  (Doz.  &  Molk.)  Broth. — Am  2-5.  Lepidopilum  calomicron 
Broth.  =  * Crossomitrium  calomicron  (Broth.)  Welch — Am  3;  L.  dia- 
phanum  (Hedw.)  Mitt. — Am  2-4;  L.  pumilum  Mitt. — Am  2,  4;  L.  sub- 
tortifolium  Bartr. — Am  1-2.  *Leskeodon  parvifolius  Bartr. — Am  4;  *L. 
wallisii  (C.  Mull.)   Broth,  ex  Par. — Am  4.  Neohypnella  chrysophyllopodia 


1  It  is  assumed  that  the  introductory  pages  of  the  first  (4)  and  second  (6)  papers  will 
be  reviewed  before  reading  this  treatise. 

*  The  asterisk  before  the  epithet  in  the  following  lists  indicates  the  species,  varieties, 
and  forms  which  are  not  known  to  occur  in  geographical  areas  other  than   the  one  cited. 

377 


378  Indiana  Academy  of  Science 

(C.  Mull.)  Bartr.  =  N.  diversifolia  (Mitt.)  Welch  &  Crum— Am  2,  4, 
5;  N.  mucronifolia  Bartr.  =  N.  diversifolia.  Pseudohypnella  mucroni- 
folia  Bartr.  ex  Welch  &  Crum  in  error  for  Neohypnella  mucronifolia. 
Thamniopsis  killipii   (Williams)    Bartr. — Am  4,  5. 

Africa,  America,  Asia,  Australia,  Europe,  and  Oceania  are  abbre- 
viated, respectively,  as  Afr,  Am,  As,  Austr,  Eur,  and  Oc.  The  numbers 
following  the  abbreviations  refer  to  specific  areas  in  these  countries,  as 
used  in  Index  Muscorum  and  by  previous  papers  on  the  distribution  of 
Hookeriaceae. 

An  observation  of  the  world  distribution  of  genera  shows  that  the 
following  are  known  from  one  area  or  one  continent  only:  Archboldiella 
and  Leskeodontopsis,  As  4;  Orontobryiim,  As  3;  Bellia,  Austr  2;  Cal- 
licostellopsis,  Stenodesmus,  and  Thamniopsis,  Am  4;  Lamprophyllnm, 
Am  6;  and  Tetrastichium,  Afr  1. 

Further  observations  show  that  additional  genera  occur  in  more 
than  one  area  and  have  a  hemispheric  or  a  continental  distribution: 
Adelothecium,  Am  1-5;  Amblytropis,  Am  3-4;  ChaetomitHopsis,  As  2-4; 
Crossomitrium,  Am  1-6;  Dimorphocladon,  As  3-4;  Helicoblepharum,  Am 
4-5;  Hemiragis,  Am  2-5;  Isodrepanium,  Am  1-5;  Neohypnella,  Am  2, 
3,  5;  Philophyllum,  Am  2,  4,  5;  Rhynchostegiopsis,  Am  2-5;  and  Steno- 
dictyon,  Am  2-4. 

The  areas  of  distribution  in  the  earlier  publications  (4,  6)  are  re- 
peated for  reference.  America  1:  North  America  (Canada,  United 
States,  Mexico),  Greenland,  Aleutian  Islands,  Bermudas;  America  2: 
Central  America  and  Cocos  Island ;  America  3 :  West  Indian  Islands 
(Greater  and  Lesser  Antilles,  Bahamas)  except  Trinidad  and  Tobago; 
America  4:  Venezuela,  Colombia,  Peru,  Bolivia,  Ecuador,  Galapagos 
Islands:  America  5:  Brazil,  Paraguay,  Guiana,  Trinidad,  Tobago;  and 
America  6:  Chile,  Argentina,  Uruguay,  Falkland  Islands,  and  Hermite 
Island. 

The  following  islands  have  been  included  with  Africa:  the  Azores, 
Canary  Islands,  Madeira  Islands,  Saint  Helena  Island,  Madagascar 
Island,  Mauritius  Island  (formerly  Isle  de  France),  Reunion  Island 
(formerly  Bourbon)  and  Kerguelen  (Desolation  Island). 

If  tropical  Africa  is  considered  as  a  distribution  center,  we  find 
species  which  occur  in  Africa  also  occurring  in  Europe  and  Asia,  as  well 
as  Australia  to  the  east  and  the  Americas  to  the  west,  particularly 
South  America.  Cyclodictyon  laete-virens  has  been  reported  from 
throughout  Africa  and  in  Europe.  Daltonia  angustifolia  occurs  in 
Africa,  Asia,  and  New  Zealand,  and  D.  strictifolia  in  Africa  and  Asia. 
D.  splachnoides  has  been  recorded  in  America,  Africa,  Europe,  and 
Australia.  Collections  of  Eriopus  cristatus  have  been  reported  from 
Africa,  South  America,  Australia,  New  Zealand,  and  Oceania.  Ap- 
parently Hookeria  lucens  grows  in  cooler  areas  than  the  great  majority 
of  the  species  since  it  occurs  in  northern  Africa  and  adjacent  Asia, 
Europe,   and   northern   North   America. 


Plant  Taxonomy  379 

Endemism  in  African  Hookeriaceae  is  greatest  in  Afr  2,  consist- 
ing of  67  species  and  varieties.  Afr  3  has  35  species,  and  Afr  1  and 
Afr  4,  two  and  six,  respectively.  The  Hookeriaceae  are  represented  in 
Afr  1  by  four  species;  Afr  2  by  89  species,  varieties,  and  forms;  Afr 
3  with  56  species  and  varieties;  and  Afr  4  by  14. 

The  European  species  of  Hookeriaceae  have  extensive  distribution 
records.  Cyclodictyon  laete-virens  occurs  also  in  Afr  1-4;  Daltonia 
splachnoides  in  Afr  2,  Am  2,  Austr  1,  2;  Distichophyllum  carinatum  in 
As  2;  Eriopus  apiculatus  (1)  (introduced  and  established  in  England), 
in  Austr  1,  2,  Am  6;  and  Hookeria  lucens  in  Am  1,  Afr  1,  and  As  5. 

One  hundred  and  sixty-eight  species,  subspecies,  and  varieties  have 
been  reported  from  As  4,  57  from  As  3,  27  from  As  2,  one  from  As  5, 
and  one  from  As  1.  It  is  evident  that  the  center  of  distribution  of 
Asiatic  species  is  As  4.  Ten  endemic  species,  subspecies,  and  varieties 
of  Hookeriaceae  are  known  in  As  2,  28  in  As  3,  and  126  in  As  4. 

Distichophyllum  carinatum  occurs  in  Eur  as  well  as  in  As  2.  As  2 
has  in  common  with  Oc:  Callicostella  papiUata,  Cyclodictyon  blume- 
anum,  Distichophyllum  mittenii,  and  Hookeria  acutifolia  which  also 
occurs  in  Am  1-5  and  As  3-4.  In  addition  to  As  2-4,  Daltonia  angusti- 
folia  also  occurs  in  Afr  2-3  and  Austr  2,  and  var.  strictifolia  in  Afr  3. 
Twenty  Asian  species  occur  in  Oc,  six  in  Afr  3,  four  in  Afr  2,  three  in 
Austr  1,  and  three  in  Austr  2.  Hookeria  lucens  has  been  reported  from 
As  5,  Eur,  Afr  1,  and  Am  1.  H.  acuti  folia  has  been  recorded  in  As  2-4, 
Am  1-5,  and  Oc.  The  following  species  in  As  3  also  occur  in  Oc :  Calli- 
costella papiUata,  C.  prabaktiana,  Cyclodictyon  blumeanum,  Daltonia 
contort  a,  Distichophyllum  'mittenii,  and  Hookeria  acutifolia  (also  in 
As  2,  4,  and  Am  1-5).  Hookeriopsis  pallidifolia  has  been  reported  from 
Afr  2  as  well  as  As  3.  In  addition  to  As  4,  Callicostella  kaernbachii, 
Chaetomitrium  tahitense,  var.  deplanchei,  and  Cyclodictyon  lepidum 
occur  in  Austr  1 ;  Callicostella  papiUata,  C.  prabaktiana,  Chaetomitrium 
tahitense,  var.  deplanchei,  Cyclodictyon  blumeanum,  var.  vescoanum,  Dal- 
hitense,  var.  deplanchei,  Cyclodictyon  blumeanum,  var.  vescoanum,  Dal- 
tonia contorta,  Distichophyllum  mittenii,  D.  undulatum,  Eriopus  re- 
motif  olia,  Hookeria  acutifolia  (also  As  2,  3,  Am  1-5),  and  Leskeodon 
acuminatus  occur  in  Oc;  Daltonia,  angustifolia  in  Austr  2  and  Afr  2,  3, 
and  its  var.  strictifolia  in  Afr  3.  One  species,  Hookeria  lucens  (Hedw.) 
Sm.,  is  the  only  species  known  from  As  5.  This  species  also  grows  in 
Eur,  Afr  1,  and  Am  1,  a  northern  distribution  in  comparison  with  the 
subtropical  and  tropical   habitats  of  the  great  majority  of  species. 

Australia  and  Tasmania  have  34  species  of  Hookeriaceae,  19  of 
which  are  endemic;  and  New  Zealand,  etc.,  26  species,  varieties,  and 
forms,  15  of  which  are  endemic.  In  the  distribution  of  species,  Australia 
and  Tasmania  have  9  species  in  common  with  New  Zealand,  and  7  in 
common  with  Am  6,  3  each  with  As  4,  Afr  3,  and  Oc,  and  1  each  with 
Eur,  Afr  2,  and  Am  2.  New  Zealand  has  9  species  which  also  occur  in 
Australia  and  Tasmania,  and  6  in  common  with  Am  6,  2  each  with 
Afr  2  and  3,  and  1  each  with  As  3  and  4,  Eur,  Am  2,  and  Oc.  It  is  ap- 


380  Indiana  Academy  of  Science 

parent  that  Australia,  Tasmania,  and  New  Zealand  have  more  species 
of  Hookeriaceae  in  common  with  each  other  and  with  Am  6  than  they 
have  with  other  countries. 

Pterygophyllum  balantii  Broth.  (Hepaticina  balantii  C.  Mull,  in 
Hedwigia  41:128.  1902)  was  described  in  his  Symbolae  ad  Bryologicam 
Australiae  III,  without  citation  of  locality  as  Australia,  Tasmania,  or 
New  Zealand. 

Austr  1  has  19  endemic  species  in  the  Hookeriaceae.  Austr  1  and  2 
have  in  common  nine  species.  Distichophyllum  rotundi folium,  Eriopus 
apiculatus,  Pterygophyllum  dentatum,  and  Sauloma  tenella  occur  in 
Austr  1,  2,  and  Am  6;  Distichophyllum  assimile  and  Pterygophyllum 
obscurum  have  been  reported  from  Austr  1  and  Am  6;  Callicostella 
kaernbachii,  Chaetomitrium  tahitense,  and  var.  deplanchei  have  habitats 
in  Austr  1  and  As  4.  Austr  1  and  Oc  have  in  common :  Chaetomitrium 
tahitense,  var.  deplanchei,  and  Eriopus  cristatus.  Species  with  a  very 
extensive  range  of  distribution  are  Daltonia  splachnoides  in  Austr  1, 
2,  Afr  2,  Eur,  Am  2,  and  Eriopus  cristatus  in  Austr  1,  2,  Afr  3,  Am  6, 
and  Oc. 

Austr  2  has  seven  endemic  species,  six  varieties,  and  two  forms  in 
26  representatives  of  the  Hookeriaceae.  Nine  species  occur  in  both 
Austr  1  and  2,  and  six  of  those  in  Austr  2  also  grow  in  Am  6;  Disti- 
chophyllum rotundi  folium,  Eriopus  apiculatus,  E.  cristatus,  E.  flexi- 
collis,  Pterygophyllum  dentatum,  and  Sauloma  tenella.  Three  species  of 
Austr  2  have  an  extensive  range  of  distribution.  Daltonia  angustifolia 
has  been  reported  from  As  2-4  and  Afr  2,3;  D.  splachnoides  from  Austr 

2,  Afr  2,  Am  2,  and  Eur;  and  Eriopus  cristatus  from  Oc,  Austr  1,  Afr 

3,  and  Am  6. 

Endemism  in  the  Hookeriaceae  of  Oceania  consists  of  45  species,  six 
varieties,  and  one  form.  Hookeria  acutifolia  occurs  in  As  2-4  and 
Am  1-5,  as  well  as  in  Oc.  Eriopus  cristatus  has  been  reported  from  Oc, 
and  also  from  Austr  1,  2,  Afr  3,  and  Am  6.  The  distribution  range  of 
four  species  from  Oceania  extends  into  As  2:  Callicostella  papillata, 
Cyclodictyon  blumeanum,  Distichophyllum  mittenii,  and  Hookei'ia 
acutifolia;  six  into  As  3;  Callicostella  papillata,  C.  prabaktiana,  Cyclo- 
dictyon blumeanum,  Daltonia  contorta,  Distichophyllum  mittenii,  and 
Hookeria  acutifolia;  and  12  into  As  4:  Callicostella  papillata,  C. 
prabaktiana,  Chaetomitrium  tahitense,  var.  deplanchei,  Cyclodictyon 
blumeanum,  var.  vescoanum,  Daltonia  contorta,  Distichophyllum  mit- 
tenii, D.  undulatum,  Eriopus  re  motif  olius,  Hookeria  acutifolia,  and 
Leskeodon  acuminatus.  Austr  1  and  Oc  have  two  species  and  a  variety 
in  common:  Chaetomitrium  tahitense,  var.  deplanchei,  and  Eriopus 
cristatus;  and  Austr  2  and  Oc,  1:  E.  cristatus. 

Hookeriacae  species  of  Oceania  have  a  distributional  relationship  of 
11  species  with  As  4,  six  with  As  3,  four  with  As  2,  two  with  Austr  2,  and 
one  each  with  Austr  2,  Afr  3,  and  Am  1-6. 


Plant  Taxonomy  381 

Africa  1:  North  Africa,  Madeira,  Azores,  Canary  Islands. 

Cyclodictyon  laete-virens  (Hook.  &  Tayl.)  Mitt.;  Hookeria  lucens 
(Hedw.)  Sm. ;  *Lepidopilum  virens  Card.;  '•  Tetrastich  ium  fontanum 
(Mitt.)  Card. 

Africa  2:  Central  Africa,  Saint  Helena  Island,  Fernando  Po. 

*Actinodontium  dusenii  Broth.;  *A.  streptogoyieum  Broth.;  Cal- 
licostella  africana  Mitt.;  *C.  ascenionis  (C.  Mull.)  Kindb.;  *C.  attenuata 
(C.  Mull.)  Kindb.;  C.  brevipes  (Broth.)  Broth.;  :::C.  chevalieri  Broth. 
in  Corb.;  *C  chinophylla  (C.  Miill.)  Kindb.;  C.  constricta  (C.  Mull.) 
Kindb.;  *C.  emarginatula  Broth,  in  Corb.;  C.  eroso-truncata  Card.; 
C.  fissidentella  (Besch.)  Kindb.;  *C.  gabonesis  Broth.  &  P.  Varde; 
C.  lacerans  (C.  Miill.)  Jaeg. ;  C.  leptocladula  (Broth.)  Broth.;  *C. 
maclaudii  (Par.  &  Broth.)  Broth.;  *C.  papillosula  Broth.  &  P.  Varde; 
*C.  perpapillata  Broth.  &  P.  Varde;  C.  pusilla  Broth,  ex  Demar.  & 
P.  Varde,  C.  salaziae  (Besch.)  Kindb.;  C.  seychellensis  (Besch.)  Kindb.; 
*C.  sub  emarginatiila  Broth.  &  P.  Varde;  C.  tristis  (C.  Miill.)  Broth.; 
:!:C.  usambarica  (Broth.)  Broth.;  *Chaetomitrium  dusenii  C.  Miill.  ex 
Broth.;  *C.  dusenii  var.  brevinerve  (P.  Varde)  P.  Varde;  ^Cyclodictyon 
bidentatum  Demar.;  C.  borbonicum  (Besch.)  Broth.;  *C.  brevifolmm 
Broth,  in  Mildbr. ;  :;:C.  crassicaule  Broth,  in  Mildbr.;  *C.  delicatum  P. 
Varde;  *C.  dixonianum  Demar.;  *C.  filicuspis  P.  Varde;  C.  hildebrandtii 
(C.  Miill.)  Kuntze;  :|:C.  immersum  Broth.  &  P.  Varde;  *C  krebedjense 
Broth.;  *C  krebedjense  var.  argutidens  P.  Varde;  C.  laete-virens 
(Hook,  and  Tayl.)  Mitt:;  *C.  lebrunii  Demar.  &  P.  Varde;  *C.  per- 
limbatum  Broth.;  *C.  preussii  (Broth.)  Broth.;  :':C.  recognitum  De- 
mar. ■&  P.  Varde;  :|:C.  spectabile  Broth,  in  Mildbr.;  *C.  subbrevi- 
folium  Broth.;  :|:C.  subobtusifolium  Dix.  ex  Demar.  &  P.  Varde;  C. 
vallis-gratiae  (C.  Miill.)  Kuntze;  C.  vallis-gratiae  f.  breutelianivm 
(Hampe)  Demar.  &  P.  Varde;  C.  vesiculosum  (Brid.)  Kuntze; 
Daltonia  angustifolia  Doz.  &  Molk.;  :D.  constricta  P.  Varde; 
*D.  dusenii  C.  Miill.  ex  Broth.;  *Z).  euryloma  P.  Varde;  *D. 
longinervis  Mitt.;  *Z).  mildbraedii  Broth,  in  Mildbr.;  *D.  mildbraedii 
var.  laevis  Demar.  &  Leroy;  *D.  minor  Besch.;  *Z).  minuta  Ther. ;  *D. 
plicata  P.  Varde;  D.  splachnoides  (Sm.)  Hook.  &  Tayl.;  *D.  tortifolia 
Demar.  &  P.  Varde;  *Distichophyllidium  africanum  Demar.  &  P.  Varde; 
*Distichophyllum  procumbens  Mitt.;  *D.  rigidicaule  (Dus.)  Broth.;  *D. 
rigidicaule  var.  gabonense  (P.  Varde)  P.  Varde;  *  Hookeria  contracta 
Gepp  in  Hiern.;  *Hookeriopsis  ambigna  P.  Varde;  *H.  angolensis 
(Welw.  &  Dub.)  Broth,  ex  Par.;  *H.  cheiloneura  (Broth.)  Broth.;  *H. 
gabonensis  Broth.  &  P.  Varde;  *H.  mittenii  P.  Varde;  H.  pallidifolia 
(Mitt.)  Geh.  &  Herz.;  *H.  papillosula  Broth.  &  P.  Varde;  H.  pappeana 
(Hampe)  Jaeg. ;  *H.  staudtii  (Broth.)  Broth.;  *Hypnella  abrupta 
(Mitt.)  Jaeg.,  *H.  guineensis  Broth.  &  Par.,  ■'Lepidopilidium  cyrtoste- 
gium  (Ren.  &  Card.)  Card,  in  Grand.;  *L.  devexum  (Mitt.)  Broth.; 
*L.  hanningtonii  (Mitt.)  Broth.;  *L.  subdevexum  (Broth.)  Broth.; 
*L.  subdevexum  var.  intermedium  P.  Varde;  *L.  theriotii  Nav.  in  Dix. 
&  Ther.;    *Lepidopilum   callochlorum    (C.   Miill.)    ex   Broth.;    L.   dusenii 


382  Indiana  Academy  of  Science 

C.  Mull,  ex  Broth.;  *L.  hirsutum  (Besch.)  Broth,  var.  tuberculatum 
Ther.;  *L.  lastii  Mitt.;  L.  niveum  (C.  Mull.)  Kindb.;  *Sauloma  afri- 
cana  Dix.  &  Ther.;  *S.  tisserantii  P.  Varde. 

Africa  3:  Madagascar,  Mauritius,  Reunion  Islands. 

* Actinodontium  hirsutum  Besch.  var.  ramosum  Besch.;  Callicostella 
af vicuna  Mitt.;  C.  brevipes  (Broth.)  Broth.;  C.  constricta  (C.  Mull.) 
Kindb.;  C.  erosotruncata  Card.;  C.  fissidentella  (Besch.)  Kindb.;  C. 
lacerans  (C.  Mull.)  Jaeg. ;  *C.  laeviuscula  Mitt.;  C.  leptocladula 
(Broth.)  Broth.;  C.  papillata  (Mont.)  Mitt.;  C.  papillata  var.  brevifolia 
Fleisch.;  *C.  parvocellulata  Demar.  &  P.  Varde;  *C  perrotii  (Par.) 
Broth.;  C.  pusilla  Broth,  ex  Demar.  &  P.  Varde;  C.  salaziae  (Besch.) 
Kindb.;  C.  seychellensis  (Besch.)  Kindb.;  * Chaetomitrium  borbonicum 
Besch.;  *C.  comorense  Hampe  ex  C.  Mull.;  *Cyclodictyon  aubertii  (P. 
Beauv.)  Kuntze;  C.  borbonicum  (Besch.)  Broth.;  C.  hildebrandtii  (C. 
Mull.)  Kuntze;  C.  laete-virens  (Hook.  &  Tayl.)  Mitt.;  *C.  perrottetii 
Demar.  &  P.  Varde;  C.  vallis-gratiae  (C.  Mull.)  Kuntze;  C.  vallis-gratiae 
f.  breutelianum  (Hampe)  Demar.  &  P.  Varde;  C.  vesiculosum  (Brid.) 
Kuntze;  Daltonia  angustifolia  Doz.  &  Molk.;  D.  angustifolia  var. 
strictifolia  (Mitt.)  Fleisch.;  *D.  forsythii  Broth,  ex  Card,  in  Grand.; 
:|:Z).  latimar ginata  Besch.;  *D.  latimarginata  var.  madagassa  Ren.; 
*Z>.  stenoloma  Besch.;  *Distichophyllum  mascarenicum  Besch.;  *Eriopus 
asplenioides  (Brid.)  Besch.;  E.  cristatus  (Hedw.)  Brid.  in  C.  Mull. ; 
*E.  fragilis  C.  Mull.;  *Hookeria  splachnif olia  (Brid.)  Arnott;  *Hooker- 
iopsis  darntyi  (Besch.)  Broth,  in  Card,  in  Grand.;  *H.  diversifolia 
(Ren.  &  Card.)  Broth,  ex  Card,  in  Grand.;  *Hypnella  semi-scabra  Ren. 
&  Card.;  *H.  viridis  Ren.  &  Card.;  '■Lepidopilidium  attenuatum  Ther.; 
*L.  branneoleum  (C.  Mull.)  Broth.;  L.  cespitosum  (Besch.)  Broth.; 
L.  chenagonii  (Ren.  &  Card.)  Card,  in  Grand.;  *L.  corbieri  (Ren.  & 
Card.)  Card,  in  Grand.;  *L.  flexuosum  (Besch.)  Broth,  ex  Par.,  *L.  is- 
leanum  (Besch.)  Broth.;  :|:L.  parvulum  Card,  in  Grand.;  *L.  subrevolutum 
(Ren.  <&  Card.)  Card,  in  Grand.;  *Lepidopilum  carrougeauanum  Ther. 
&  P.  Varde;  L.  hirsutum  (Besch.)  Broth.;  'L.  hirsutum  var.  ramosum 
Besch.;  :|:L.  hirsutum  var.  tuberculatum  Ther.;  :;:L.  humblotii  Ren.  & 
Card.;  *L.  verrucipes  Card,  in  Grand. 

Africa  4:  South  Africa,  Kerguelen  Island,  Tristan  da  Cuhna. 

^Callicostella  applanata  Broth.  &  Bryhn;  C.  fissidentella  (Besch.) 
Kindb.;  C.  salaziae  (Besch.)  Kindb.;  C.  tristis  (C.  Miill.)  Broth.; 
Cyclodictyon  laete-virens  (Hook,  and  Tayl.)  Mitt.;  C.  vallis-gratiae 
(C.  Mull.)  Kuntze;  C.  vallis-gratiae  f.  breutelianum  (Hampe)  Demar.  & 
P.  Varde;  *  Daltonia  tristaniensis  Dix.  in  Christ.;  -Distichophyllum 
fasciculatum  Mitt.;  D.  imbricatum  Mitt.;  W.  nniiifoliuni  (Hornsch.) 
Sim;  *Z>.  taylorii  Sim;  *Eriopus  mniaceum  (C.  Miill.)  Broth.; 
Hookeriopsis  papeana  (Hampe)  Jaeg*. 

Europe,  Iceland,  the  Caucasus. 

Cyclodictyon  laete-virens  (Hook.  &  Tayl.)  Mitt.;  Daltonia  splach- 
noides  (Sm.)  Hook.  &  Tayl.;  Distichophyllum  cariiiatum  Dix.  &  Nich.; 
Eriopus  apiculatus  (Hook.  f.  &  Wils.)  Mitt,  (introduced  and  established 
in  England)  ;  Hooker ia  lucens  (Hedw.)  Sm. 


Plant  Taxonomy  383 

Asia  1 :  North  Asia  including  Sakhalin  Island. 

Hookeria  lucens  (Hedw.)  Sm. 

Asia  2:  China,  Mongolia,  Japan,  Korea,  Taiwan  (Formosa). 

Callicostella  papillata  (Mont.)  Mitt.;  Chaetomitriopsis  glaucocarpa 
(Schwaegr.)  Fleisch.;  Cyclodictyon  blumeanum  (C.  Mull.)  Kuntze; 
Daltonia  angustifolia  Doz.  &  Molk. ;  D.  angustifolia  var.  strictifolia 
(Mitt.)  Fleisch.;  D.  aristifolia  Ren.  &  Card.;  *Distichophylhim  breviro- 
stratum  Ther.;  D.  carinatwm  Dix.  &  Nich.;  *D.  cavaleriei  Ther.;  *D. 
collenchymatosum  Card.;  D.  cuspidatum  (Doz.  &  Molk.)  Doz.  &  Molk.;  D. 
gracilicaule  Fleisch.;  D.  jungermannioides  (C.  Mull.)  Bosch  &  Lac;  ":D. 
maibarae  Besch.;  D.  mittenii  Bosch  &  Lac;  D.  montagneanum  (C.  Mull.) 
Bosch  &  Lac;  D.  nigricaule  Mitt,  ex  Bosch  &  Lac;  *D.  obtusif olium 
Ther.;  *D.  osterwaldii  Fleisch.;  *D,  stillicidiorum  Broth.;  *Eriopus  ja- 
ponicus  Card.  &  Ther. ;  E.  parviretis  Fleisch. ;  *E.  spinosus  Nog. ;  Hook- 
eria acutifolia  Hook.  &  Grev.;  Hookeriopsis  pappeana  (Hampe)  Jaeg. ; 
H.  utacamundiana   (Mont.)    Broth.;    *H.  yakushimensis  Toyama. 

Asia  3:  India,  Pakistan,  Ceylon,  Burma,  Thailand  (Siam),  Vietnam 

(Indochina). 

Actinodontium  ascendens  Schwaegr.;  A.  rhaphidostegum  (C.  Mull.) 
Bosch  &  Lac;  Callicostella  papillata  (Mont.)  Mitt,;  C.  prabaktiana  (C. 
Mull.)  Bosch  &  Lac;  Chaetomitriopsis  glaucocarpa  (Schwaegr.)  Fleisch.; 
Chaetomitrium  ciliatum  Doz.  &  Molk.  ex  Bosch  &  Lac;  *C.  confertum 
Thwait.  &  Mitt.;  C.  cucullatwm  Dix.;  C.  leptopoma  (Schwaegr.)  Bosch  & 
Lac;  *C.  nervosum  Dix.;  C.  papillif olium  Bosch  &  Lac;  C.  philippinense 
(Mont.)  Bosch  &  Lac;  C.  torquescens  Bosch  &  Lac;  *C.  volutum  Mitt.; 
Cyclodictyon  blumeanum  (C.  Mull.)  Kuntze;  Daltonia  angustifolia  Doz.  & 
Molk.;  D.  angustifolia  var.  strictifolia  (Mitt.)  Fleisch.;  *Z).  apiculata 
Mitt.;  *D.  aristifolia  Ren.  &  Card.  ssp.  leptophylla  Fleisch.;  *D. 
brevipedunculata  Mitt.;  D.  contorta  C.  Mull.;  D.  contorta  ssp.  inac- 
gregorii  (Broth.)  Fleisch.;  *D.  flexifolia  Mitt.;  *D.  gemmipara  Dix.;  *D. 
marginata  Griff.;  *D.  perlaxiretis  Dix.;  *D.  reticulata.  C.  Mull.;  *D. 
semitorta  Mitt.;  *D.  sub  angustifolia  Ren.  &  Card.;  Dimorphocladon 
borneense  Dix.;  *Distichophyllum  ceylanicum  (Mitt.)  Par.;  D.  cuspi- 
datum  (Doz.  &  Molk.)  Doz.  &  Molk.;  *£>.  griffithii  (Mitt.)  Par.;  *D. 
heterophyllum  (Mitt.)  Par.;  *D.  humifusum  (Mitt.)  Par.;  *Z>.  limpidum 
Thwait.  &  Mitt.;  *D.  madurense  Ther.  &  P.  Varde;  D.  mittenii  Bosch  & 
Lac;  D.  montagneanum,  (C.  Miill.)  Bosch  &  Lac;  D.  nigricaule  Mitt,  ex 
Bosch  &  Lac;  *D.  obovatum  (Griff.)  Par.;  D.  schmidtii  Broth.;  D. 
sinuosulum  Dix.;  'D.  succulentum  (Mitt.)  Broth.;  *Eriopus  bonianus 
Besch.;  *£\  lucidus  Thwait.  &  Mitt.;  E.  parviretis  Fleisch.;  E.  remoti- 
folius  C.  Miill.;  Hookeria,  acutifolia  Hook.  &  Grev.;  Hookeriopsis  pallidi- 
folia  (Mitt.)  Geh.  &  Herz.;  *H.  purpuvata  (Mitt.)  Broth.;  *H.  secunda 
(Griff.)  Broth.;  'H.  thwaitesiana,  (Mitt.)  Broth.;  H.  utacamundiana, 
(Mont.)  Broth.;  *Lepidopilidium  furcatum  (Thwait.  &  Mitt.)  Broth.; 
*Orontobryum  hookeri  (Mitt.)  Fleisch.;  Pseudohypnella  verrucosa  (Doz. 
&  Molk.)  Fleisch. 


384  Indiana  Academy  of  Science 

Asia  4:  Indonesia,  Malaya,  Philippine  Islands,  New  Guinea. 

*Actinodontium  ascendens  Schwaegr. ;  A.  rhaphidostegum  (C.  Mull.) 
Bosch  &  Lac;  *Archboldieila  pilifera  Bartr.;  *Callicostella  aiomensis 
Bartr. ;  *C.  armata  Herz.;  *C.  beccariana  (Hampe)  Jaeg\ ;  *C.  chloneura 
C.  Mull.;  *C.  eberhardtiana  Broth.  &  Par.;  C.  kaernbachii  Broth.;  *C. 
loriae  Fleisch. ;  C.  papillata  (Mont.)  Mitt.;  *C.  papillata  var.  brevifolia 
Fleisch.;  *C  papillata  var.  viridissima  Dix.;  :|:C.  paupera  (C.  Mull.) 
Kindb.;  C.  prabaktiana  (C.  Mull.)  Bosch  &  Lac;  *C.  prabaktiana  var. 
acuminata  Baumg. ;  *C.  pterygophylloides  (Broth.)  Broth.;  Chaetomitri- 
opsis  glaucocarpa  (Schwaegr.)  Fleisch.;  *C  diversifolia  Zant. ;  *Chaeto- 
mitrium  acanthocarpum  Bosch  &  Lac;  *C  auriculatum  Dix.  &  Herz.; 
*C.  beccarii  Dix.;  *C.  bornense  Mitt.;  *C.  brassii  Bartr.;  C.  ciliatmn 
Doz.  &  Molk.  e:r  Bosch  &  Lac;  *C.  crispifolium  Bartr.;  *C.  ctenidioides 
Broth.;  C.  cucullatum  Dix.;  *C.  divergens  Dix.;  *C.  elegans  Geh.;  *C. 
elmeri  Broth.;  *C.  elongatum  (Doz.  &  Molk.)  Doz.  &  Molk.;  *C. 
everettii  Mitt,  e:r  Dix.;  :::C.  fimbriatum  (Doz.  &  Molk.)  Bosch  &  Lac; 
*C.  finisterrae  Dix.  &  Herz.;  *C.  horridulum  Bosch  &  Lac;  *C.  integri- 
folium  Bartr.;  *C  laevifolium  Dix.;  :::C.  laevisetum  Dix.;  :;:C.  lanceolatum 
Bosch  &  Lac;  *C.  lancifolium  Mitt.;  :;:C.  lauterbachii  Broth.;  C.  lep- 
topoma  (Schwaegr.)  Bosch  &  Lac;  *C.  leptopoma  var.  massartii  Ren.  & 
Card.;  :|:C.  longisetulum  Bartr.;  *C.  macrohystrix  C.  Mull. ;  :::C.  madan- 
gense  Bartr.;  *C.  nanohystrix  C.  Miill.;  C.  nematosum  Broth.;  C. 
orthorrhynchum  (Doz.  &  Molk.)  Bosch  &  Lac;  *C.  paleatum  (Hampe) 
Fleisch.;  C.  papillifolium  Bosch  &  Lac;  *C.  papuanum  Bartr.;  *C. 
parcesetulosum  Bartr.;  *C.  perakense  Broth.;  :::C.  perarmatum  Broth.; 
:i:C.  perlaeve  Dix.;  C.  philippinense  (Mont.)  Bosch  &  Lac;  *C.  plicatum 
Bartr.;  :::C.  poecilophyllum  Dix.;  :::C.  pseudo-eiongatum  Broth.;  *C. 
pseudo- papillifolium  Bartr.;  :::C.  recurvifolimn  Fleisch.;  *C.  rigidulum 
Broth.;  *C.  robbinsii  Bartr.;  :|:C.  roemeri  Fleisch.;  *C.  seriatum  Broth.; 
:!:C.  setosum  Broth.;  :;:C.  spinosum  Bartr.;  :::C.  sublaevisetum  Dix.;  :::C. 
subplicatum  Bartr.;  C.  tahitense  (Sull.)  Mitt.;  C.  tahitense  var. 
deplanchei  (Besch.)  Wijk  &  Marg. ;  C.  torquescens  Bosch  &  Lac;  *C. 
torquescens  var.  barbatum  Dix.;  *C  vrieseanum  Bosch  &  Lac;  *C. 
warburgii  Broth.;  C.  weberi  Broth.;  *C.  werneri  Herz.;  *C.  wildei  Zant.; 
Cyclodictyon  blumeanum  (C.  Miill.)  Kuntze;  *C  blumeanum  var. 
morokae  Fleisch.;  C.  blumeanum  var.  vescoanum  (Besch.)  Fleisch.;  C. 
lepidum  (Mitt.)  Broth.  &  Watts;  Daltonia  angustifolia  Doz.  &  Molk.; 
*D.  angustifolia  var.  gemmiphylla  Fleisch.;  *D.  angustifolia  var.  longi- 
pedunculata  (C.  Miill.)  Fleisch.;  *D.  angustifolia  var.  rcvoluta  (Broth.) 
Bartr.;  D.  angustifolia  var.  strictifolia  (Mitt.)  Fleisch.;  D.  aristifolia 
Ren.  &  Card.;  *D.  armata  Bartr.;  *Z).  baumgartneri  Froehl.;  D.  contorta 
C.  Miill.;  D.  contorta  ssp.  mac-gregorii  (Broth.)  Fleisch.;  :|:Z).  contorta 
var.  humilis  Fleisch.;  *D.  himalayeusis  Dix.  &  Herz.;  D.  pseudosteno- 
phylla  Bartr.;  *D.  schiffneri  Froehl.;  :D.  tuberculosa  Bartr.;  Dimorpho- 
cladou  borneense  Dix.;  *Distichophyllidium  antarense  Zant.;  *D.  junger- 
manniaceum  Fleisch.;  -:D.  nymanianum  Fleisch.;  :]:D.  rhizophorum 
Fleisch.;  *Distichophyllum  aciphyllum  Dix.;  *D.  angustifolium  Dix.;  *D. 
angustissimum  Dix.;  *D.  borneense  Broth.;  :|:Z).  brevicuspidatum  Bartr.; 
*D.  brevicuspis  Fleisch.;  *D.  catinifolium  Froehl.;    'D.  cucullatum  Bartr.; 


Plant  Taxonomy  385 

D.  cuspidatum  (Doz.  &  Molk.)  Doz.  &  Molk. ;  *D.  denticulatum  Dix.;  *D. 
dixonii  Herz.;  *D.  elmeri  Broth.;  *D.  evanidolimbatum  Fleisch.;  D.  graci- 
licaule  Fleisch.;  D.  jungermannioides  (C.  Mull.)  Bosch  &  Lac;  -D.  leio- 
pogo7i  Dix.;  *Z>.  lixii  (Broth.)  Ther.;  -D.  longipes  Broth.;  *D.  longobasis 
Fleisch.;  *D.  lorianum  Fleisch.;  *D.  macropodium  Dix.;  D.  mittenii  Bosch 
&  Lac;  D.  montagneanum  (C.  Mull.)  Bosch  &  Lac;  *D.  nidulans  Herz.;  D. 
nigricaule  Mitt,  ex  Bosch  &  Lac;  *Z>.  nigricaule  var.  cirratum  (Ren.  & 
Card.)  Fleisch.;  *D.  nigricaule  var.  complanatum  Fleisch.;  *D.  perundu- 
latum  Dix.;  *D.  pullei  Dix.;  *D.  santosii  Bartr. ;  D.  schmidtii  Broth.; 
D.  sinuosulum  Dix.;  *Z>.  spathulatum  (Doz.  &  Molk.)  Doz.  &  Molk.;  *D. 
stipitati folium  C.  Mull.;  *Z).  submucronatum  Fleisch.;  *D.  subnigricaiile 
Broth.;  *D.  tortile  Doz.  &  Molk.;  *D.  turgidum  Bartr.;  D.  undulatum 
Doz.  &  Molk.;  *Eriopus  cristatus  (Hedw.)  Brid.;  *E.  flaccidus  Broth.; 
■fE.  microblastus  Broth.;  E.  parviretis  Fleisch.;  *E.  perlimbatiis  Dix.; 
*£'.  ramosus  Fleisch.;  E.  remotifolius  C.  Mull. ;  E.  subremotifolius  Broth.; 
Hookeria  acutifolia  Hook.  &  Grev.;  *Hookeriopsis  gemmidens  Broth.: 
*H.  ?nacropus  (Bosch  &  Lac)  Broth.;  *//.  majurei  Bartr.;  H.  utacamun- 
diana  (Mont.)  Broth.;  :H.  wichurae  Fleisch.;  Leskeodon  acuminatus 
(Bosch  &  Lac)  Fleisch.;  *L.  acuminatus  var.  laeviseta  Zant. ;  *L.  pan- 
durifolius  Fleisch.;  *L.  philippinensis  Broth.;  *L.  robbinsii  Bartr.; 
*L.  rotundifolius  Bartr.;  *Leskeodontopsis  pustulata  Zant.;  Pseudo- 
hypnella  verrucosa  (Doz.  &  Molk.)  Fleisch.;  *Pterygophyllum  javense 
Dix.;   *P,  novae-gaineae  Bartr.;   *Sauloma  tenuis  C.  Mull. 

Asia  5:  Asiatic  part  of  the  Middle-East,  including  Cyprus. 

Hookeria  lucens  (Hedw.)  Sm. 

Australia  1 :  Australia  and  Tasmania. 

■fCallicosteUa  bailey i  (Broth.)  Kindb.;  C.  kaernbachii  Broth.;  *C. 
rugiseta  Dix.;  *Chaetomitrium  entodontoides  Broth.  &  Watts;  C.  tahi- 
tense  (Sull.)  Mitt.;  C.  tahitense  var.  deplanchei  (Besch.)  Wijk  &  Marg. ; 
Cyclodictyon  lepidum  (Mitt.)  Broth.  &  Watts;  *Daltonia  pusilla  Hook.  f. 
&  Wils.;  D.  splachnoides  (Sm.)  Hook.  &  Tayl.;  DistichophyUum  assimile 
Broth.;  *D.  baileyanum  C.  Mull. ;  *D.  beccarii  (Harape  &  Geh.)  Par.; 
*D.  complanatum  (Hampe)  Mitt.;  D.  crispulum  (Hook.  f.  &  Wils.)  Mitt.; 
*jD.  levieri  (Geh.)  Broth.;  *D.  longicuspis  Broth.;  D.  microcarpum 
(Hedw.)  Mitt.;  *D.  minutif olium  C.  Mull.;  D.  pulchellum  (Hampe) 
Mitt.;  D.  rotundif olium  (Hook.  f.  &  Wils.)  C.  Mull.  &  Broth.;  *£>. 
sub  minutif  olium  (Broth.  &  Geh.)  Fleisch.;  *D.  whiteleggeanum  C.  Mull.; 
Eriopus  apiculatus  (Hook.  f.  &  Wils.)  Mitt.;  *E.  brassii  Bartr.;  E. 
cristatus  (Hedw.)  Brid.  in  C.  Mull.;  *E.  tasmanicus  Broth.;  *Pterygo- 
phyllum  bryoides  Broth.;  P.  dentatum  (Hook.  f.  &  Wils.)  Dix.; 
*P.  flaccidissimum  Broth.;  P.  obscurum  Mitt.;  *P.  subrotundum  (Hampe) 
Jaeg. ;  *P.  wattsii  Broth.;  Sauloma  tenella  (Hook.  f.  &  Wils.)  Mitt.; 
*S.  zetterstedtii  (C.  Mull.)  Jaeg. 

Australia  2:  New  Zealand,  etc. 

*Bellia  nervosa  (Hook.  f.  &  Wils.)  Broth.;  Daltonia  angustifolia 
Doz.  &  Molk.;  D.  splachnoides  (Sm.)  Hook.  &  Tayl.;  DistichophyUum 
crispulum  (Hook.  f.  &  Wils.)   Mitt.;  *Z).  crispulum  var.  adnatum   (Hook. 


386  Indiana  Academy  of  Science 

f.  &  Wils.)  Dix.;  D.  microcarpum  (Hedw.)  Mitt.;  fD.  microcarpum  var. 
homodictyon  Sainsb.;  *D.  microcladum  (Col.)  Broth.;  D.  pulchellum 
(Hampe)  Mitt.;  *D.  pulchellum,  var.  elliptic? folium  Sainsb.;  *D.  pul- 
chellum  var.    parvirete    Sainsb.;    D.    rotundifolium    (Hook.    f.    &    Wils.) 

C.  Mull.  &  Broth.;  Eriopus  apiculatus  (Hook.  f.  &  Wils.)  Mitt.;  *E. 
brownii  Dix.;  E.  cristatus  (Hedw.)  Brid.  in  C.  Mull.;  E.  flexicollis 
(Mitt.)  Jaeg.;  *Hookeria  flava  Col.;  Pterygophyllum  dentatum  (Hook.  f. 
&  Wils.)  Dix.;  *P.  dentatum  var.  latifolium  (C.  Miill.)  Wijk  &  Marg.; 
*P.  dentatum  var.  robustum  (Hook.  f.  &  Wils.)  Dix.;  *P.  distichophyl- 
loides  Broth.  &  Dix.;  *P.  quadrif avium  (Sm.)  Brid.;  *P.  quadvif  avium  f. 
mavginata  Sainsb.;  *Sauloma  macvospova  Sainsb.;  S.  tenella  (Hook.  f.  & 
Wils.)  Mitt.;  *S.  tenella  f.  pvopagulifeva  Sainsb. 

Oceania:  Pacific  Islands. 

*Callicostella  bisexualis  (Besch.)  Jaeg.;  :;:C.  caledonica  Ther.;  *C. 
coMipbelliana  (Hampe)  Jaeg. ;  *C.  chlovina  (Besch.)  Broth.;  :|:C.  fvatevi 
Broth.  &  Watts;  *C.  melanotheca  (Besch.)  Jaeg.;  *C.  melanotheca  var. 
scabviseta  Ther.;  :i:C.  nukahivensis  (Besch.)  Broth.;  *C.  oblongifolia 
(Sull.)  Jaeg.;  C.  papillata  (Mont.)  Mitt.;  C.  prabaktiana  (C.  Miill.) 
Bosch  &  Lac;  *C.  vesiculata  (C.  Miill.)  Jaeg.;  *Chaetomitvium-  aneitense 
Broth.  &  Watts;  *C.  callichvoum  Besch.;  *C.  densum  Dix.  ex  Bartr. ; 
*C.  depvessum  Mitt.;  :!:C.  frondosum  Mitt.;  :;:C.  hebridense  Dix.;  C. 
ovthovvhynchum  (Doz.  &  Molk.)  Bosch  &  Lac.  var.  vitense  Bartr.; 
*C.  vugifolium  (Sull.)  Mitt.;  :!:C.  smithii  Bartr.;  *C.  speciosum  (Sull.) 
Mitt.;  C.  tahitense  (Sull.)  Mitt.;  C.  tahitense  var.  deplanchei  (Besch.) 
Wijk  &  Marg.;  C.  weberi  Broth.;  *C.  wheeleri  Hampe;  *Cyclodictyon 
bescherellei  (Par.)  Broth.;  C.  blumeanum  (C.  Miill.)  Kuntze;  *C.  blumea- 
nwm  f.  gvaeffeana  (C.  Miill.)  Fleisch.;  C.  blumeanum  var.  vescoanum 
(Besch.)     Fleisch.;    *Daltonia    baldwinii    Broth.;    D.    contorta    C.    Miill.; 

D.  pseudostenophylla  Bartr.;  *D.  vufescens  Broth.;  *D.  sphaevica  Besch.; 
*Distichophyllidium  maticiim  Broth.  &  Par.;  *Distichophyllum  apicidi- 
gevum  Broth.  &  Par.;  *D.  capillatum  Mitt.;  D.  cuspidatum  (Doz.  & 
Molk.)  Doz.  &  Molk.;  D.  fasciculatum  Mitt.;  *D.  flavescens  (Mitt.)  Mitt. 
in  Seem.;  :]D.  fossombvonioides  Ther.;  *Z).  fvancii  Ther.;  *Z>.  fveycinetii 
(Schwaegr.)  Mitt.;  *D.  fveycinetii  var.  crassetuvgescens  C.  Mull.;  *D. 
graeffeanum  (C.  Miill.)  Broth.;  D.  imbvicatum  Mitt.;  *D.  koghiense 
Ther.;  *D.  limbatulum  (C.  Miill.)  Par.;  'D.  lingulatum  Bartr.;  D.  mit- 
tenii  Bosch  &  Lac;  D.  montagneanum  (C.  Miill.)  Bosch  &  Lac;  :]:D. 
nadeaudii  Besch. ;  VD.  pavadoxum  (Mont.)  Mitt.;  *D.  samoanum  Fleisch.; 
■•'D.  samoanum  var.  bvevipes  Bartr.;  ':D.  semimavginatum  Ther.;  *D. 
tahitense  Besch.;   *D.  tovquati folium  Dix.;  D.  undulation   Doz.  &  Molk.; 

D.  vitianum  (Sull.)  Mitt.;  Eriopus  cristatus  (Hedw.)  Brid.  in  C.  Mull.; 
*E.  marginatus  Ther.;  *E.  pacificus  (Besch.)  Fleisch..;  E.  remotifolius  C. 
Miill.;  *E.  subvemotifolius  Broth.;  Hookevia  acutifolia  Hook.  &  Grev. ; 
*H.  sandvicensis  Reichdt. ;  *Hookeviopsis  purpurea  (C.  Miill.)  Broth.; 
*H.  purpurea  var.  acuminatula,  (C.  Miill.)  Bartr.;  *H.  purpurea  var. 
ligulacea  (C.  Miill.)  Bartr.;  Leskeodon  acuminatus  (Bosch  &  Lac.) 
Fleisch. 


Plant  Taxonomy  ::x7 

Literature  Cited 

Paton,    Jean    A.    1968.    Eriopus    apiculatus    (Hook.    f.    &    Wils. )     Mitt,    established    on 
Tresco.  Trans.  Brit.  Bryol.  Soc.  5(3)  :460-462. 

Welch,    Winona    H.    1962.    The   Hookeriaceae   of   the   United   States    and    Canada.    The 
Bryologist  65(1)  :l-24. 

— .  1966.  The  Hookeriaceae  of  Mexico.  The  Bryologist  69(1)  :l-68. 


— .     1968.     Hookeriaceae     Species     and     Distribution     in     North     and     Central 
America  and  West  Indies.  Proc.  Indiana  Acad.  Sci.  77  :351-356. 

.  1969.  The  Hookeriaceae  of  Cuba.  The  Bryologist  72(2)  :93-136. 

— .     1969.     Hookeriaceae    Species    and    Distribution    in     South    America.     Proc. 


Indiana  Acad.  Sci.  78  :396-405. 

7.      Wijk,   R.  van  der,  W.   D.   Margadant,  and  P.  A.   Florschmtz.     1959,   1962,    1964,    1967, 
1969.  Index  Muscorum.  Vol.  1-5.  Utrecht,  The  Netherlands. 


Biosystematic  Studies  of  the  Beech  and  Marsh  Ferns1 

Jeanette  C.  Oliver,  Ball  State  University 


Abstract 

The  taxonomic  status  of  five  species  of  ferns  common  to  the  Northeastern  United 
States  has  been  in  a  state  of  confusion  for  many  years.  The  broad  beech  fern,  the  long 
beech  fern,  the  marsh  fern,  the  New  York  fern,  and  the  bog  fern  have  been  placed  in 
the  genus  Thelypteris  by  some  authors.  Others  consider  them  members  of  the  large 
cosmopolitan  genus  Dryopteris.  In  other  interpretations  the  beech  ferns  are  separated 
into  the  genus  Phegopteris. 

Evidences  from  morphological,  cytological,  and  biochemical  studies  do  not  warrant  the 
inclusion  of  these  species  in  the  genus  Dryopteris.  Significant  differences  were  noted 
between  the  beech  ferns  and  the  marsh  ferns  which  would  appear  to  justify  their 
separation  into  different  genera. 

This  study  was  undertaken  in  an  attempt  to  clarify  the  confusion 
regarding  the  taxonomic  status  of  five  species  of  ferns  common  to  the 
northeastern  United  States,  These  ferns,  and  their  counterparts  from 
other  regions,  have  been  variously  classified  generically. 

Some  have  considered  the  broad  beech  fern,  the  long  beech  fern, 
the  bog  fern,  the  marsh  fern,  and  the  New  York  fern  as  being  of  com- 
mon generic  rank.  These  ferns  were  treated  as  members  of  the  large 
cosmopolitan  genus  Dryopteris  by  Christensen  in  his  monograph  (4,  5). 
This  interpretation  was  followed  by  Fernald  in  his  Gray's  Manual  of 
Botany  (9)  and  by  Deam  in  his  Flora  of  Indiana   (8)    (Table  1). 

Christensen,  in  a  later  work,  set  the  genus  Thelypteris,  with  the 
marsh  fern,  Thelypteris  palustris  as  its  type,  apart  from  the  genus 
Dryopteris  (6).  Morton  (12)  in  his  contributions  to  Britton  and  Brown's 
Illustrated  Flora  (10)  recognized  the  differences  between  the  "thelyp- 
teroid"  and  the  "dryopteroid"  ferns    (Table  1). 

On  the  basis  that  Thelypteris  Schmidel  was  not  validly  published, 
Copeland  (7)  in  his  Genera  Filicum  used  the  name  Lastrea  Bory.  The 
Nomenclature  Committee  at  the  International  Botanical  Congress  in 
Stockholm  decided,  however,  that  the  name  Thelypteris  is  correct  and 
voted  down  a  proposal  to  conserve  Lastrea  (13,  14). 

Cytologically  the  beech  ferns  differ  from  both  the  dryopteroids  and 
the  thelypteroids;  further,  their  sori  lack  indusia  in  contrast  with  the 
latter.  Therefore,  some  feel  that  these  ferns  should  be  placed  in  a 
separate  genus,  Phegopteris    (11,  21)    (Table  1). 

Methods  and  Materials 

Extensive  collections  of  the  five  species  were  made  throughout 
Indiana  and  Ohio.  A  total  of  422  herbarium  specimens  from  the  Field 
Museum  of  Natural  History  and  432  specimens  from  the  Missouri 
Botanical  Garden  was  examined. 


1  This    work    was   supported   by   grants   from    the    Indiana   Academy    of   Science,    Sigma 
Xi,  and  Ball  State  University. 

388 


Plant  Taxonomy 


M) 


a. 

o 


s  §  e 

©     <»  *» 

.■«  <  p* 

8^  ■§ 


6^ 

t.      CO 


«    © 


fe^ 


6 

2 

x 

■« 

.OS 

^ 

a. 

ft 

© 

2 

53 

o 

<» 

•CI 

.1 

~ 

90     o3 

*«K 

e 

© 

•2  ^ 

ft 

•  2 

a 

o 

w 

m 

•  2 

,2! 

s 

,C0       fe 

'£ 

£ 

*£   a 

3^ 

ft 

3> 

% 

as 
ft 

a 

CD 

> 

05     (L> 

ft  2 

» a 
^  s 

« 

<*> 

OS 

o3 

<»   . 

-^ 

J 

-=; 

*s 

Q 

tVl     w 

^H 

&H 

in 

&H 

Q  S 


ft 

o 

©} 


2 

c 

8 

© 

© 

c- 

CO 

i 

a 

CD 

*© 

g 

co 

c 

•  2 

.2  1* 

- 

j-   a 

<s 

» 

ft 

CD        <D 

•^ 

"*w 

a 

ft 

+j 

a 

=5  ^ 

5* 

-P 

==3 

1 — . 

o 

*■»*  .—. 

Q» 

» 

<»  -^- 

>s£ 

-=: 

Q 

^  j 

tV, 

W 

&H 

&H    w 

a 

ft 

o 
53} 

05 

Oh 

«3 

C 

CD 

Q 

« 

02 

( . 

'£ 

"fi 

X 

X 

rC 

a. 

u 

CJ 

o 

od 


ft 
o 


5v 

S     ft 

a  a 

O      <D 
^3     > 

O  Q 


'** 

a 

cu 

u 

a 

fe 

<v 

fe 

0) 

X 

a 

fe 

CD 

a 

4«J 

o 

O 

PQ 
a 

03 

^ 

o 

o 

CD 

pq 

J 

^ 

CQ 

Z 

390  Indiana  Academy  of  Science 

The  external  morphology  was  carefully  studied.  Drawings  and  photo- 
graphs were  made  to  record  distinguishing  characteristics. 

Rhizomes  and  stipes  were  cross  sectioned  at  10>  and  stained  with 
safranin-fast  green. 

Spore  samples  were  obtained  from  both  living  and  moribund  speci- 
mens. The  spores  were  mounted  in  Hoyer's  medium  for  clearing  and 
preservation. 

For  cytological  study,  meiotic  materials  were  fixed  in  3  parts  ethyl 
alcohol  :  1  part  glacial  acetic  acid.  Squashes  were  made  in  aceto-orcein. 

Biochemical  comparisons  were  made  by  means  of  paper  chromatog- 
raphy. Extracts  were  prepared  by  powdering  dried  fronds  and  soaking 
the  materials  in  80%  methanol  for  48  hours.  Fifty  microliters  of  each 
sample  were  applied  to  Whatman  #1  paper  using  the  spot  method.  Both 
1  and  2-dimensional  chromatograms  were  run  descendingly  using  bu- 
tanolracetic  acidiwater  (12:3:5)  as  the  solvent. 

Dried  chromatograms  were  examined  in  the  presence  of  ultra- 
violet light.  The  chromatograms  were  sprayed  sequentially  with  ninhy- 
drin  and  Ehrlich  reagent  for  the  detection  of  free-amino  acids  and  re- 
lated substances.  Some  chromatograms  were  sprayed  with  Pauly's 
reagent  or  alkaline  silver  nitrate  for  the  detection  of  phenolic  com- 
pounds (16). 

Morphological  Observations 

The  frond  of  the  beech  ferns  consists  of  a  deltoid  blade  borne  by 
a  pilose  stipe.  The  lanceolate  apex  of  the  blade  is  pinnatifid. 

The  blade  of  the  long  beech  fern  has  10-12  pairs  of  separate  pinnati- 
fid pinnae  below  the  apex.  The  length  of  the  blade  exceeds  its  breadth  at 
the  widest  point.  Nine  to  18  pairs  of  surcurrent  pinnae  are  noted  in  the 
broad  beech  fern.  The  lower  3-4  pairs  are  bipinnatifid  and  the  upper  ones 
are  pinnatifid  (Fig.  1,  2). 

Both  species  bear  rounded,  supramedial  sori  which  lack  indusia. 

The  blades  of  the  New  York,  bog,  and  marsh  ferns  are  lanceolate  in 
outline  and  are  composed  of  pinnatifid  segments  below  a  pinnatifid  apex. 
The  dark  stipe  is  glabrous  in  the  marsh  fern  and  scaly  in  the  other  two 
species. 

The  blade  of  the  New  York  fern  is  composed  of  18-40  pairs  of 
oblong-lanceolate  pinnae.  The  lower  4-5  pairs  are  greatly  reduced  in 
size  (Fig.  3).  The  sori  are  located  submarginally  and  have  a  kidney- 
shaped  indusium. 

The  marsh  fern  possesses  13-25  pairs  of  linear-lanceolate  pinnae 
which  bear  medial  indusiate  sori  (Fig.  4). 

The  blade  of  the  bog  fern  is  composed  of  10-38  pairs  of  oblong- 
lanceolate  pinnae.  The  lowermost  are  basioscopic  (Fig.  5).  The  indusiate 
sori  are  borne  submarginally. 


Plant  Taxonomy 


8i)l 


I  0 


/m    T  ,    t     PwtTaphS     °f     herba™™      specimens.      1)      Phegopteris      hexagonoptera 
(Mich*.)    Feet    broad   beech  fern;   2)    Phegopteris   polypodioides   Fee>    long    beech   fcrn. 

Thelypteris  noveboracensis   (L.)   Nieuwl,  New   York  fern;  4)    Thelypteris  palustris  Schott 
marsh  fern;   5)    Thelypteris   simulata    (Davenp.)    Nieuwl.,    bog   fern 

Figures   6-10.     Fern   spores   X   675.    6)    Broad   beech   fern;    7)    Long    beech   fern;   8)    Bog 
fern;  9)  Marsh  fern,  10)  New  York  fern. 


392  Indiana  Academy  of  Science 

Spore  Morphology 

The  spores  of  all  five  species  are  bilateral  in  shape  and  bear  an 
adherent  perispore.  Spores  of  the  beech  ferns  are  light  greenish-brown 
in  color.  Their  perispore  does  not  form  conspicuous  ridges  or  wings. 
The  spores  of  the  broad  beech  fern  are  20-27  x  30-50/a.  The  surface  is 
tuberculate  in  texture.  The  perispore  is  not  winged  (Fig.  6).  Spores  of 
the  long  beech  fern  are  28-30  x  55-60/a.  The  spore  surface  is  tuberculate. 
The  perispore  varies,  some  spores  being  wingless  and  ridgeless  and 
others  bearing  short  broken  ridges  and  narrow  discontinuous  wings 
(Fig.  7). 

The  remaining  species  are  yellow-brown  in  color  and  possess  a 
tuberculate  winged  and  ridged  perispore.  The  spores  of  the  marsh  fern 
are  34-45  x  53-63/u.  Ridges  are  few  and  broken  (Fig.  9).  The  New  York 
fern  spores  are  highly  sculptured  with  wide  apical  wings.  The  spores  are 
25-30  x  33-45/*  (Fig.  10).  The  spores  of  the  bog  fern  are  sculptured 
similarly  to  those  of  the  New  York  fern,  but  approach  those  of  the  marsh 
fern  in  size  (Fig.  8). 

Cytological  Observations 

It  is  apparent  from  this  and  other  studies  that  different  chromosome 
numbers  exist  among  the  thelypteroid  ferns. 

Broad  beech  fern  materials  collected  in  Indiana  showed  a  chromosome 
number  of  n  =  30  (Voucher  Specimen  #0159  Oliver).  Counts  made  of 
Virginia  specimens  by  Wagner  (18)  and  by  Britton  (1,  2,  3)  of  materials 
from  Ontario  confirm  this  number. 

Studies  of  the  long  beech  fern  have  been  made  by  Manton  (11)  in 
Great  Britain  and  by  Britton  (1,  2,  3).  Both  have  reported  numbers  of 
n  =  90  and  In  =  90.  The  long  beech  fern  is  apogamous.  A  doubling  of 
the  diploid  number  of  90  chromosomes  occurs  just  prior  to  meiosis;  thus 
the  sporophytic  and  the  gametophytic  numbers  are  the  same. 

Counts  of  the  marsh  fern  (Voucher  #0172  Oliver)  and  the  New  York 
fern  (Voucher  #0195  Oliver)  yielded  numbers  of  n  =  35  and  n  =  27, 
respectively.  These  numbers  are  in  agreement  with  those  reported  by 
Wagner  (17,  18).  Britton  reported  a  number  of  n  =  35  for  the  marsh 
fern;  however,  his  tentative  studies  of  the  New  York  fern  indicated 
n  =  29  (1). 

Meiotic  material  of  the  bog  fern  was  unavailable  for  this  study. 
Wagner  reported  a  number  of  n  =  64  for  specimens  collected  in 
Maryland  (17). 

Biochemical  Observations 

Chromatograms  were  prepared  from  dried  specimens  of  ferns  col- 
lected throughout  the  northeastern  United  States.  Materials  had  been 
preserved  for  periods  of  1  month  to  25  years.  Remarkable  consistency 
was  obtained  from  materials  of  different  ages  and  localities. 


Plant  Taxonomy  393 

Examination  of  chromatograms  in  the  presence  of  ultraviolet  light 
revealed  one  compound  with  a  rf  value  of  0.67  common  to  all  species. 
Another  substance,  rf  0.80,  was  common  to  the  marsh,  the  bog,  and  the 
New  York  fern.  Several  additional  spots  appeared  to  be  species  specific. 

Subsequent  spraying  revealed  all  ninhydrin  reacting  compounds  to  be 
species  specific.  The  compound  common  to  all  and  that  common  to  the 
three  species  were  indicated  by  their  positive  reactions  with  Pauly's 
reagent  and  alkaline  silver  nitrate  to  be  phenolic  (Fig.  11). 

Negative  results  were  obtained  with  Ehrlich's  reagent  in  all  cases. 


0 


*     / 


On  ©» 

KJ"  ©n 

On 

0„ 


>_'  n 


a 


0 

.»Q,     0P      o;     ©p       e, 

.*o0p  Op 


•; 


am 


I  11 

L  .    _  i 

Figure  11.  One-dimensional  descending  chromatograms.  A.  Marsh  fern,  B.  Bog  fern, 
C.  Long  beech  fern,  D.  Broad  beech  fern,  E.  Netv  York  fern  (n  =z  ninhydrin  postive 
substance  p  =  phenolic  compound) . 


Discussion  and  Summary 

All  aspects  of  this  study  indicate  exclusion  of  the  beech  and  marsh 
ferns  from  the  genus  Dryopteris. 

The  blades  of  these  ferns  are  light  green,  thin-membranous,  and  non- 
evergreen;  whereas,  those  of  true  Dryopteris  are  deep  green,  subcori- 
aceous  and  evergreen.  The  vascular  bundles  reach  the  margins  of  the 
pinnae  in  contrast  to  those  of  the  dryopteroids  which  terminate  in 
hydathodes. 


394  Indiana  Academy  of  Science 

The  thelypteroids  bear  slender  rhizomes;  the  stipes  contain  two 
vascular  bundles  which  unite  at  the  base  of  the  blade.  Massive  rhizomes 
and  stipes  bearing  3-7  bundles  are  characteristic  of  the  dryopteroids. 

The  spores  are  distinguished  with  some  difficulty.  Basically,  the 
spores  of  the  thelypteroids  have  a  tuberculate  surface;  perispore  ridges, 
if  present,  are  quite  discontinuous.  Dryopteroid  spores  tend  to  be  cristate 
with  closed,  anastomosing  ridges  (15). 

Extensive  cytological  studies  have  been  made  of  the  American 
species  of  Dryopteris  sensu  stricto  (19,  20).  A  base  chromosome  number 
of  n  =  41,  accompanied  by  various  levels  of  ploidy,  was  noted.  This 
number   has   not  been   seen  in   any  thelypteroid  fern. 

There  appears  to  be  much  basis  for  maintaining  the  beech  ferns  as 
a  separate  genus  Phegopteris.  Some  special  characters  common  to  the 
beech  ferns  as  opposed  to  the  marsh  ferns  are  as  follow:  deltoid  blades; 
exindusiate  sori;  spores  wingless  or  bearing  much  reduced  broken  wings; 
chromosomes  extremely  small.  Chromatographically,  fewer  common 
phenolic  substances  were  detected  in  the  beech  than  in  the  marsh  ferns 
(Fig.  11). 

In  conclusion,  the  results  of  this  study  indicate  the  exclusion  of  the 
thelypteroid  ferns  from  Dryopteris;  further  basic  differences  appear  to 
justify  the  delimitation  of  the  beech  ferns  as  a  distinct  genus.  A  number 
of  outstanding  problems  are  inherent  in  these  ferns.  The  relationship  of 
the  oak-fern  to  the  beech  ferns  is  one  such  problem.  Another  is  the  rela- 
tionships of  the  thelypteroids  of  the  western  United  States  to  those  of 
the  east.  Extensive  studies  of  Western  North  American,  Asian,  and 
European  counterparts  are  needed  before  the  taxonomy  of  the  group  can 
be  understood  fully. 

Literature  Cited 

1.  Britton,  D.  M.  1953.  Chromosome  studies  on  ferns.  Amer.  J.  Bot.  40  :575-583. 

2.  Britton,  D.  M.  1961.  The  problems  of  variation  in  North  American  Dryopteris. 
Amer.  Fern.  J.  51  :23-30. 

3.  Britton,  D.  M.  1965.  The  cytology  and  distribution  of  Dryopteris  species  in 
Ontario.  Can.  J.  Bot.  44  :63-78. 

4.  Christensen,  Carl.  1913.  A  monograph  of  the  genus  Dryopteris  Part  I.  The  tropical 
American   pinnatifid-bipinnatifid  species.   Kgl.   Dansk.   Vid.   Selsk.   Skr.,   VII.    10  :55-582. 

5.  Christensen,  Carl.  1920.  A  monograph  of  the  genus  Dryopteris  Part  II.  The 
tropical  American  bipinnate-deeompound  species.  Kgl.  Dansk.  Vid.  Selsk.  Skr., 
VIII.  6:1-123. 

6.     Christensen,    Carl.    1938.    Manual    of    Pteridology.    Chronica    Botanica    Press.    550    p. 

7.     Copeland,    E.     B.     1947.     Genera    Filicum.     Chronica    Botanica    Press.    247    p. 

8.     Beam.     C.     C.     1940.     Flora    of     Indiana.     Department    of     Conservation,     Indianapolis. 
1236  p. 

9.  Fernald,  M.  L.  1950.  Gray's  Manual  of  Botany.  American  Book  Co.,  New  York. 
1632  p. 


Plant  Taxonomy  395 

10.  Gleason,     H.    A.     1952.     Illustrated    Flora    of    the    Northeastern     United    States    and 
Adjacent    Canada.    New    York    Botanical    Garden,    New    York.    Vol.    1.    482    p. 

11.  Manton,   I.   1950.   Problems  of  cytology  and  evolution  in  the  Pteridophyta.   University 
Press,  Cambridge,  Eng.  316  p. 

12.  Morton,    C.    V.    1950.    Notes   on   the   ferns   of   the   eastern    United   States.    Amer.    Fern 
J.  40:213-225. 

13.  Morton,     C.     V.     1958.     The     Californian     species     of     Thelypteris.     Amer.     Fern     J. 
48:136-162. 

14.  Morton,    C.    V.    1963.    The  classification  of   Thelypteris.    Amer.    Fern   J.    53:149-154. 

15.  Oliver,   J.    C.    1968.   A   study   of  spore  characteristics   of  the   ferns   of   Indiana.    Amer. 
Fern  J.  58  :5-12. 

16.  Smith,    I.    1958.    Chromatographic    techniques:     Clinical    and    biochemical    application. 
Heinemann  Medical  Books,  Ltd.  London.  419  p. 

17.  Wagner,    W.     H.    Jr.     1963.    A    biosystematic    survey    of    United    States    ferns — pre- 
liminary abstract.  Amer.  Fern  J.  53  :1-16. 

18.  Wagner,    W.    H.    Jr.    1966.    Pteridophytes    of    the    Mountain    Lake    area,    Giles    Co., 
Virginia:  Biosystematic  studies,  1964-5.  Castanea  31:121-140. 

19.  Walker,     S.     1961.     Cytogenetic    studies    in    the    Dryopteris    spinulosa    complex     II. 
Amer.  J.  Bot.  48  :607-614. 

20.  Walker,  S.   1962.  Further  studies  in  the  genus  Dryopteris;  the  origin  of  D.   clintoni- 
ana,  D.  celsa  and  related  taxa.  Amer.  J.  Bot.  49  :497-503. 

21.  Wherry,    E.    T.    1937.    Guide   to   eastern    ferns.    Science    Press,    Lancaster,    Pa.    252    p. 


Cytotaxonomic  Notes  on  Genus  Polygonum,  Section  Polygonum1 

Donald  N.  Moore,  Thomas  R.  Mertens  and  Joyce  E.  Highwood, 
Ball  State  University 


Abstract 

The  most  accurate  identification  of  species  in  genus  Polygonum,  section  Polygonum 
is  based  on  fruit  and  perianth  characteristics.  Preliminary  investigations  suggested  that 
these  morphological  features  could  be  effectively  correlated  with  specific  chromosome 
numbers.  The  achenes  of  five  species  of  Polygonum  collected  in  Indiana,  Nova  Scotia  and 
New  Brunswick  are  briefly  described  and  chromosome  numbers  for  these  species  are 
reported  as  follows:  P.  aviculare  L.  sensu  stricto  2n  =  60;  P.  buxijorme  Small  In  = 
60;  P.  arenastrum  Bor.  In  =  40;  P.  fowleri  Robinson  2n  =  ca.  40;  and  P.  erectum  L. 
2«  =   40. 

Although  numerous  taxonomic  investigations  of  genus  Polygonum, 
section  Polygonum  (Avicularia)  have  been  conducted,  there  is  little 
agreement  as  to  species  identity  in  North  America  or  elsewhere.  Much  of 
the  taxonomic  confusion  comes  from  the  tremendous  morphological 
variation  within  the  individual  species  of  the  section.  At  present  the 
most  accurate  identification  of  Polygonum  specimens  is  based  on  fruit 
and  perianth  characteristics.  The  purpose  of  this  investigation  was  to 
attempt  to  gain  evidence  to  support  this  basis  of  identification  by  showing 
that  morphological  features  can  be  correlated  with  chromosome  numbers. 
Plants  identified  as  a  given  species  on  the  basis  of  morphological  charac- 
ters may  be  expected  to  have  identical  chromosome  numbers. 

The  fruit  of  Polygonum  is  a  small,  dry,  indehiscent  achene  with  a 
relatively  thin  wall.  According  to  Styles  (5),  Mertens  and  Raven  (2), 
Savage  and  Mertens  (4),  and  Mertens  (3),  fruit  and  perianth  character- 
istics are  the  most  consistent  morphological  features  used  in  the 
identification  of  species  within  section  Polygonum.  The  criteria  employed 
for  identification  in  this  investigation  included:  achene  color,  texture, 
shape,  and  size;  the  depth  of  perianth  sinuses;  and  the  position  of 
inflorescences  on  the  stem.  Inflorescences  are  located  either  at  the  apices 
of  branches  or  in  the  axils  of  the  leaves  along  the  stem,  depending  on  the 
species. 

Cytological  investigation  by  Love  and  Love  (1)  has  revealed  that  the 
chromosomes  of  species  within  section  Polygonum  are  uniformly  small 
in  size  with  centrally  located  centromeres.  The  present  investigation  sup- 
ports the  same  conclusion.  Love  and  Love  claim  that  the  basic  chromo- 
some number  of  the  section  is  2n  =  20  with  tetraploid  and  hexaploid 
plants  being  quite  common.  Mitotic  chromosome  counts  of  40  and  60 
for  species  in  section  Polygonum  have  been  frequently  reported  in  the 
literature  (2,  5,  6). 

This  report  consists  of  a  brief  discussion  of  achene  characteristics 
and  chromosome  numbers  for  several  species  of  section  Polygonum 
collected  in  Indiana,  Nova  Scotia  and  New  Brunswick. 


1  This   study   was   supported   by   grants   to   Dr.    Mertens   from   the    Indiana   Academy   of 
Science  and  The  Society  of  Sigma  Xi. 

396 


Plant  Taxonomy 


897 


# 


**A 


I 

a 


w    •*%%> 


B 


1     *t 


f 


f 


t 


' 


0 


Figure  1.     Typical  achenes.    A.  Polygonum  aviculare  L.,  B.  P.  buxifoime  Small,  and  C.   P. 
fowleri  Robinson. 


398  Indiana  Academy  of  Science 

Polygonum  aviculare  L.  sensu  stricto,  has  achenes  (Fig.  1A)  which 
are  dull  and  heavily  striated  (5).  Their  color  ranges  from  dark  brown  to 
black.  The  achene  has  two  more-or-less  equal  concave  sides  and  a  third 
flat  side,  and  is  completely  enclosed  in  the  perianth,  which  is  divided  for 
three-fourths  or  more  of  its  length.  The  achenes  of  P.  aviculare  from 
Nova  Scotia  and  New  Brunswick  varied  from  3.58  to  4.08  mm  in  length 
(mean:  3.93  mm)  and  from  2.16  to  2.92  mm  in  width  (mean:  2.54  mm). 

Chromosome  counts  of  2n  —  60  and  2??  —  ca.  60  were  obtained  for 
P.  aviculare  from  specimens  collected  in  Nova  Scotia  and  New  Brunswick. 
Counts  of  2n  =  60  for  this  species  are  well  documented  in  the  literature 
(2,5,6): 

1)  P.  aviculare  L.,  rocky  beach  on  Sand  Beach  south  of  Yarmouth, 
Yarmouth  Co.,  Nova  Scotia,  August  14,  1968,  T.  R.  Mertens  NS-2. 
(BSU).  2n  =  60. 

2)  P.  aviculare  L.,  Yarmouth  Co.,  Nova  Scotia,  August  16,  1968, 
T.  R.  Mertens  NS-16.  (BSU).  2m  =  ca.  60. 

3)  P.  aviculare  L.,  St.  John  Co.,  New  Brunswick,  August  19,  1968, 
T.  R.  Mertens  NB-35.  (BSU).  2w  =  ca.  60. 

4)  P.  aviculare  L.,  Deer  Island,  Charlotte  Co.,  New  Brunswick, 
August  19,  1968,  T.  R.  Mertens  NB-42.  (BSU).  2n  =  ca.  60. 

The  achenes  (Fig.  IB)  of  Polygonum  buxiforme  Small  are  small  in 
size  in  comparison  to  those  of  the  closely  related  P.  aviculare.  Specimens 
from  Nova  Scotia  and  New  Brunswick  measured  2.58  to  3.16  mm  in 
length  (mean:  2.87  mm)  and  1.58  to  2.93  mm  in  width  (mean:  2.25  mm). 
The  achene  is  dull  to  mildly  shiny,  dark  brown  in  color  and  heart-shaped 
with  two  equal,  concave  sides  and  one  flat  side.  The  achene  is  enclosed 
in  the  perianth,  which  has  a  distinctive  flanged  edge  (Fig.  IB).  The 
perianth  is  divided  for  %  to  %  of  its  length. 

Chromosome  numbers  of  2n  =z  60  and  2n  —  ca.  60  were  obtained  for 
the  following  specimens  of  P.  buxiforme: 

1)  P.  buxiforme  Small,  Dearborn  Co.,  Indiana,  July  9,  1966,  A.  D. 
Savage  17-2.  (BSU).  2n  =  60. 

2)  P.  buxiforme  Small,  Dearborn  Co.,  Indiana,  July  9,  1966,  A.  D. 
Savage  17-3.  (BSU).  2n  =  60. 

3)  P.  buxiforme  Small,  Halifax  Co.,  Nova  Scotia,  August  18,  1968, 
T.  R.  Mertens  NS-26.  (BSU).  2n  =  60. 

4)  P.  buxiforme  Small,  Queens  Co.,  Nova  Scotia,  August  17,  1968, 
T.  R.  Mertens  NS-22.  (BSU).  2n  =  ca.  60. 

5)  P.  buxiforme  Small,  Queens  Co.,  Nova  Scotia,  August  17,  1968, 
T.  R.  Mertens  NS-23.  (BSU).  2n  =  ca.  60. 

6)  P.  buxiforme  Small,  Lunenburg  Co.,  Nova  Scotia,  August  17, 
1968,  T.  R.  Mertens  NS-24.  (BSU).  2n  =  ca.  60. 


Plant  Taxonomy  399 

Polygonum  arenastrum  Bor.,  is  characterized  by  achenes  which  are 
dark  brown  with  two  more-or-less  equal  convex  and  one  narrow  concave 
side.  Savage  and  Mertens  (4)  report  that  the  achenes  of  this  species 
range  from  1.58  to  2.50  mm  in  length  and  from  1.00  to  1.75  mm  in 
width.  The  achene  surface  is  dull  but  shiny  along  the  edges. 

A  chromosome  count  of  2n  —  40  was  obtained  from  the  following 
specimen  collected  in  Indiana: 

1)  P.  arenastrum  Bor.,  Delaware  Co.,  Indiana,  September  18,  1968, 
Joyce  Highwood  1-2.  (BSU).  2n  —  40. 

Another  specimen,  identified  on  the  basis  of  morphological  features 
as  P.  buxiforme,  was  found  to  have  a  diploid  chromosome  number  of  40 
thus  suggesting  that  it  was,  in  fact,  P.  arenastrum. 

2)  P.  arenastrum  Bor.,  (?)  Warrick  Co.,  Indiana,  September  17, 
1966,  A.  D.  Savage  62-1.  (BSU).  2n  =  40. 

Styles    (5)    and    Mertens    and    Raven    (2)     also    report    2n    =    40    for 
P.  arenastrum. 

Polygonum  fowleri  Robinson  (=  P.  allocarpum  Blake)  produces 
highly  distinctive  achenes  having  a  granular  texture  and  a  "beak-like" 
apex  (Fig.  1C).  A  high  incidence  of  biconvex  fruits  are  encountered  in 
this  species  (2).  The  length  of  the  achenes  determined  from  the  study  of 
specimens  from  New  Brunswick,  Canada,  ranged  from  2.92  to  3.67  mm 
with  an  average  of  3.34  mm.  The  width  of  these  achenes  ranged  from 
1.63  to  2.08  mm  with  the  average  of  1.85  mm. 

The  color  of  the  granular  achenes  varied  from  a  light  to  a  medium 
brown.  Although  normally  three  sided  with  two  sides  convex  and  one 
side  concave,  some  achenes  have  only  two  sides  (Fig.  1C). 

The  light  green  perianth  completely  encloses  the  achene  of  P. 
fowleri.  The  perianth  is  divided  for  %  to  %  of  its  length,  varying  with 
individual  specimens.  The  inflorescences  appear  in  the  axils  of  the 
leaves  along  the  stem. 

Taylor  and  Mulligan  (6)  report  n  =  20  for  P.  fowleri  specimens 
from  Graham  Island  in  the  Queen  Charlotte  Islands,  British  Columbia, 
Canada.  Supporting  their  data  are  the  following  chromosome  counts  of 
2n  r=  ca.  40  obtained  from  specimens  of  P.  fowleri  collected  in  New 
Brunswick: 

1)  P.  fowleri  Robinson,  Lord's  Cove,  Deer  Island,  Charlotte  Co., 
New  Brunswick,  August  19,  1968,  T.  R.  Mertens  NB-36.  (BSU). 
2n  =  ca.  40. 

2)  P.  fowleri  Robinson,  Lord's  Cove,  Deer  Island,  Charlotte  Co., 
New  Brunswick,  August  19,  1968,  T.  R.  Mertens  NB-38.  (BSU). 
2n  =  ca.  40. 

The  achenes  of  Polygonum  erectum  L.  are  light  brown  to  tan  in 
contrast  to  the   much  darker  fruits   exhibited  by   the   other   species   of 


400  Indiana  Academy  of  Science 

section  Polygonum.  The  P.  erectum  achene  is  dull  and  granular  with 
two  convex  and  one  concave  side.  It  has  been  reported  that  the  achenes 
range  from  2.33  to  2.92  mm  in  length  and  1.58  to  2.17  mm  in  width  (4). 
The  perianth  is  characteristically  bottle-shaped  and  divided  for  less 
than  one-half  of  its  length. 

The  chromosome  number  of  2n  —  40  was  established  for  the 
following  specimens  from   Indiana: 

1)  P.    erectum,    L.,    Porter    Co.,    Indiana,    August    29,    1966,    A.    D. 
Savage  58-1.  (BSU).  2m  =  40. 

2)  P.    erectum    L,,    Porter    Co.,    Indiana,    August    29,    1966,    A.    D. 
Savage  58-2.  (BSU).  2n  -  40. 

Voucher  specimens  for  the  plants  for  which  chromosome  numbers 
are  reported  in  this  paper  are  housed  at  Ball  State  University. 

This  investigation  supports  the  importance  of  fruit  and  perianth 
characteristics  and  chromosome  number  in  the  identification  of  species 
within  Polygonum,  section  Polygonum.  Members  of  a  given  species, 
identified  on  the  basis  of  morphological  features,  may  be  expected  to 
have  identical  chromosome  numbers.  This  was  generally  found  to  be 
the  case  in  the  present  investigation.  Chromosome  numbers  reported 
herein  agree  with  those  reported  in  the  literature  in  those  cases  where 
such  reports  are  known.  On  the  other  hand,  the  fact  that  morphologically 
distinct  species  {e.g.,  P.  arenastrum,  P.  fowleri,  and  P.  erectum)  have 
identical  chromosome  numbers  {i.e.,  2n  =  40)  indicates  that  chromo- 
some number  alone  cannot  be  used  in  identifying  species  in  section 
Polygonum. 


Literature  Cited 

1.  Love,  A.,  and  D.  Love.  1956.  Chromosomes  and  taxonomy  of  eastern  North  American 
Polygonum.  Can.  J.  of  Bot.  34:501-521. 

2.  Mertens,  T.  R.,  and  P.  H.  Raven.  1965.  Taxonomy  of  Polygonum,  section  Polygonum 
(Avicularia)   in  North  America.  Madrono  18(3):85-92. 

3.  Mertens,  T.  R.  1968.  Polygonum — A  knotty  problem  in  plant  taxonomy.  Amer.  Biol. 
Teacher  30(10)  :832-840. 

4.  Savage,  A.  D.,  and  T.  R.  Mertens.  1967.  A  taxonomic  study  of  genus  Polygonum, 
section  Polygonum  (Avicularia)  in  Indiana  and  Wisconsin.  Proc.  Indiana  Acad.  Sci. 
7  7:357-369. 

5.  Styles,  B.  T.  1962.  The  taxonomy  of  Polygonum  aviculare  and  its  allies  in  Britain. 
Watsonia.  5:177-214. 

6.  Taylor,  R.  L.,  and  G.  A.  Mulligan.  1968.  Flora  of  the  Queen  Charlotte  Islands  Part 
2.  Cytological  aspects  of  the  vascular  plants.  Research  Branch,  Canada  Department  of 
Agriculture.    Monogr.    No.    4    Part    2.    Ottawa,    Canada.     148    p. 


SOIL  SCIENCE 

Chairman:  James  E.  Newman,  Purdue  University 
Clyde  W.  Hibbs,  Ball  State  University,  was  elected  Chairman  for  1970 

Distribution  of  Corn  (Zea  mays  L .)  Roots  in  Two  Soils  in 
Relation  to  Depth  of  Sampling  and  Type  of  Sampler1 

Russell  K.  Stivers,  Purdue  University13 

Abstract 

A  high  yielding  field  containing  both  Crosby  silt  loam  and  Ragsdale  silty  clay  loam 
was  selected  for  this  study.  The  roots  of  the  crop  were  sampled  while  the  corn  was  in 
the  maturation  or  ear  filling  stage.  Soils  cores  in  15  cm  (6  in.)  vertical  samples  were 
taken  in  each  soil  type  with  a  bucket  auger  and  with  a  Pit-O-Matic  core  sampler  to  a 
depth  of  152  cm  (60  in)  where  possible.  Gravel  in  the  Crosby  profiles  made  it  im- 
possible to  sample  deeper  than  107  cm  (42  in.)  in  4  of  the  5  subsampling  areas.  Roots 
and  soil  were  separated.  Dry  weights  of  roots  decreased  highly  significantly  with  depth. 
The  0-30  cm  (0-12  in.)  depths  contained  from  75  to  82%  of  the  total  weight  of  roots  in 
the  sampled  profiles.  Less  than  2'/(  of  the  roots  were  below  107  cm  (42  in.)  The  bucket 
auger  took  more  soil  and  with  it  more  roots  per  15  cm  vertical  sample  than  did  the 
Pit-O-Matic  core  sampler.  Sample  holding  diameters  of  these  two  tools  were  the  same, 
but  sample  cuttings  diameters  were  different. 

Distribution  of  corn  roots  in  soil  is  important  because  corn  plants 
obtain  water  and  nutrients  from  soil.  Fehrenbacher  (2,  3)  has  shown 
that  physical  properties  of  soil  and  their  genetic  horizons  are  related 
to  available  water-holding  capacity  and  to  root  penetration  and  total 
weight.  The  purpose  of  this  study  was  to  compare  vertical  distribution 
of  corn  roots  by  using  two  different  soil  sampling  tools  on  two  different 
adjoining  high-yielding  soils. 

Methods  and  Procedures 

Hand  soil  sampling  tools  were  used  in  this  study  because  it  was  not 
possible  to  use  a  Kelley  (1)  or  similar  type  mounted  soil  core  sampler 
while  the  corn  was  growing.  The  hand  type  soil  sampling  tools  used  were 
a  standard  bucket  auger'  and  a  Pit-O-Matic  core  sampler4  of  approxi- 
mately the  same  size,  5.5  cm  (2i%  in)  inside  diameter.  The  cutting  edge 
diameter  of  the  bucket  auger  was  7.0  cm  (2%  in),  and  that  of  the  core 
sampler  was  5.2  cm  (2i16  in). 

A  Crosby  silt  loam  (Aerie  Ochraqualff)  and  a  Ragsdale  silty  clay 
loam  (Typic  Argiaquoll)  adjacent  to  each  other  in  the  same  field  of 
William  Franklin,  Jamestown,  Boone  County,  Indiana,  were  used  for 
this  study.  The  soils  were  described  by  D.  P.  Franzmeier  and  P.  W. 
Harlan    (personal  communication).    Some   properties   of  these    soils   are 


1  Journal   Paper  No.   3875,   Purdue  University  Agr.   Exp.   Sta. 

J  The   author   acknowledges   the   help   of   C.    D.    Raper,    Richard   Fletcher,    Ethel   Tudor, 
and  Enola  Ruff  in  conducting  this  research. 

:!  Manufactured  by  Art's  Machine  Shop,  American  Falls,  Idaho. 

4  Manufactured  by  the  Soil  Testing  Company,  Smithville,  Tennessee. 

401 


402 


Indiana  Academy  of  Science 


**>  s  **  5 

J  o  o  o 

3  £  £  £ 


TJ  T!  73  -C 
O  V  9>  II 


F  o  o  o  o 

M   £  £  £  £ 


>>  a  a 

*  a  ■ 

>>  °  °  °     _*      5 

1   II   II  II  II   II   II   II   II 


J  X 


X>    T3  .O 


jjK^^JJ   SK^J 


H  > 


£  o  o  o  ta 

M  00  00  ,o  o 


O  W  lO  LO  o 
rH  Jft  OJ  <X>  N 
M      O]      rH      T-H      H 


<j  w  w  w  w  w 
jj  >  j>  >'  >'  > 

-  j  ti  X  X 
X  >  >  >  > 

> 

^     »     (C     !fi     »     » 
J    M    M    CO     Q3     CO 

lO    t-    N    ffl    (O 
00                      CO    CO 

■o    II 
c 

V 

J  > 

II  II 


J  § 


II  II 

X 
X  > 


N    H    »    O 


C-    t-    C-    00 


.-h   o   o   o   o 


S3    S3    S3    S3 


S3    fl    S3    S3 


£    £ 


£  !S 


£  «  s 


H      O 


lO     t- 

lO    ™    CO    LO 
N    in 

'      C>]     +j     +j 

c  MM 

bo 


CM     N     H 


»       M,    CM     Ol     „     CM      ryj      H    „     N     CM 

^<|  P3  ffl  o  u    o  «;  pq  pq  pq 
«  6 


Soil  Science  403 

listed  in  Table  1.  Ragsdale  is  higher  in  organic  matter,  more  alkaline  in 
subsurface  horizons  to  parent  material,  and  has  a  higher  strong  acid 
phosphorus  test  value  in  the  B  horizon  to  56  cm  (22  in)  than  does  the 
Crosby  soil.  Ragsdale  is  found  in  lower  areas  to  which  runoff  comes 
from  the  higher  Crosby  soil. 

Soil  and  root  samples  were  taken  between  August  9  and  August  14, 
1968,  after  midsilk  date  of  this  corn  crop,  while  the  ears  were  still 
filling  and  while  the  brace  roots  were  still  growing.  The  variety  was 
Northrup  King  610  planted  April  27.  For  the  soil  and  root  sampling  and 
for  yield  determinations,  a  nested  design  suggested  by  W.  E.  Nyquist 
(personal  communication)  was  used.  Soils  were  main  treatments.  Five 
replications  or  subsampling  areas  approximately  15.3  m  (50  ft)  apart, 
located  in  the  center  and  on  the  points  of  the  compass,  were  used  in  each 
soil  type.  Within  each  sublocation,  depths  were  sampled  to  152  cm  (60 
in)  where  possible,  in  15  cm  (6  in)  vertical  increments  starting  at  the 
surface.  At  each  sublocation  two  samplers,  the  standard  bucket  auger 
and  the  Pit-O-Matic  core  sampler,  were  used,  one  on  each  side  of  a 
corn  plant  located  18  cm  (7.1  in)  from  another  corn  plant  in  the  row  and 
96.5  cm  (38  in)  from  plants  in  adjoining  rows.  Surface  increments  in 
the  row  touched  the  stalk. 

Soil  and  root  subsamples  for  each  15  cm  increment  were  put  in 
paper  bags  and  air  dried  at  room  temperature.  Prior  to  separating  roots 
from  soil,  the  samples  were  dried  at  38°  C  (100°  F)  for  24  hours  and 
weighed.  After  the  dried  sample  was  broken  up  into  smaller  pieces  with 
a  mortar  and  pestle,  the  roots  were  separated  from  the  soil  by  the 
method  of  Raper  (5).  This  consisted  of  a  wet  screening  procedure  with 
oscillation  in  a  saturated  calgonite  solution.  Tweezers  and  flotation  in 
water  were  used  where  soil  particles  containing  roots  failed  to  break 
down.  Roots  were  placed  in  steel  cans  and  dried  at  38°  C  for  24  hours, 
weighed,  and  placed  in  labeled  plastic  bags  for  reference. 

Corn  grain  yields  were  taken  in  1968  in  the  same  five  sub-locations 
on  each  soil  where  the  soil  and  root  samples  were  taken.  Yields  are 
reported  with  15.5%  moisture  in  the  grain. 

Weights  of  roots,  soil  plus  roots  (roots  were  less  than  1%  of  the 
total  weight),  and  yields  were  subjected  to  analyses  of  variance  and  F 
tests  of  significance. 

Results  and  Discussion 

Weight  of  roots  per  15  cm  depth  of  soil  sampled  decreased  highly 
significantly  from  the  0-15  cm  depth  to  the  15-30  cm  depth.  The  average 
root  weight  of  the  0-15  cm  depth  was  1.33  grams  or  67%  of  the  total 
root  weight  to  152  cm  (60  in)  depth  and  that  of  the  15-30  cm  depth  was 
0.22  grams  or  11%  of  the  total  root  weight  to  152  cm  (60  in)  depth. 
Even  though  average  root  weights  declined  as  soil  depth  increased 
below  30  cm  depth,  these  differences  in  root  weights  were  not  significant 
at  19  to  1  or  greater  odds.  These  averages  are  shown  in  Table  2  for  all 
depths   sampled,  both   soils,   and   both   soil   sampling  tools.   Part  of  the 


404 


Indiana  Academy  of  Science 


=  S 


K     73 


5.  c1 
.2   e 


5  I 

o 


V     ft 


»  ft  02 

♦i  £  £ 

*  g  a 

.  .£>  t^ 

>  s 

0  » 


.£     3     60 
"to 

g    fi    £ 


a  a  » 

.j  £  £ 

*  g  2 

.  -O  fed 


©©t-oooot-;©©©© 

lO    O    »    00    ■*    N    ri 


iM    O    <M     O    Tji     i-l 


"*     CM     C5     CT5     O     O     CO 


t-     00     CM     SO     O     O    00 

»   •*   *   t-   o>  w  o 


lO    O    O    lO    N    M    M 


<S    O    CM    ©    «C    fr- 


ill       i        i       i        i      O     O     O 

oo©cmcm©©©;7;2;£ 

t-   to   to   ©   io   .-h   O 
CM    ^H    o    o    o    o    © 


I- 

S 

■t 

© 

X 

~ 

o 

1- 

o? 

-+ 

00 

a 

\- 

» 

tfi 

co 

— 

-M 

IN 

CM 

© 

"-1 

H 

— 

o 

© 

O] 

CO 

". 

■f 

© 

rH 

1- 

,_i 

CO 

S 

~ 

t 

M 

o 

N 

X 

CM 

: 

' 

— 

>'. 

- 

© 

O 

~ 

2 

© 

c 

C: 

© 

IO 

to 

to 

lO 

IO 

to 

CO 

- 

o 

© 

H 

O 

,. 

o 

o 

CM 

o 

© 

o 

1- 

■--■ 

~. 

°\ 

•"< 

© 

© 

o 

IC 

to 

to 

to 

to 

to 

to 

to 

to 

IC 

X 

© 

© 

■en 

Ol 

o 

© 
© 

to 

V- 

CO 

o 

© 

o 

CM 

5C 

c 

IC 

— 

© 
© 

rH 

iMO)00»NOi»H 
N'fXO^OOH 


£  a 


ire   ©   -*   ©   © 


Ooohmwhnoo 

■*  CO  M1  <fi  N  t-  O0 
©  fr-  CO  CO  C5  "*  i-H 

O  ■*  «   H  H  O   O 


CO  IO  IO  CO 


HOOCDCSMOOIIOM 


CO     ■**     CO     ©     © 


lO    O    00    00     © 

t-     H     N      H      O     O 

©      r-i      ©      ©      ©      © 


o    o    o 
B  fc  £ 


■t 

IM 

t- 

-t 

H 

CO 

"1 

>  1 

'I 

© 

© 

© 

© 

© 

©. 

J, 

^ 

f 

J 

CC 

■t 

1- 

■"* 

-r 

«■• 

•■;-! 

IC 

7 

© 

X 

CM 

-/ 

i- 

,,, 

SC 

iC 

r 

. 

© 

^ 

■  c 

© 

© 

© 

© 

© 

© 

© 

© 

-. 

IC 

to 

IC 

iC 

IC 

to 

CM 

rH 

.-H 

5 

© 

© 

© 

>c 

© 

CO 

© 

^ 

© 

© 

fr- 

-/ 

■■'.• 

CI 

IC 

© 

CM 

ee 

T. 

— 

3i 

:  i 

_ 

-_■ 

t- 

0-1 

'I 

^ 

© 

© 

© 

_ 

© 

IC 

iC 

IC 

IC 

IC 

ire 

IC 

to 

iC 

to 

co 

iC 

© 

H 

IC 

© 

CO 

r 

-f 

CM 

.■x 

— 

CM 

© 

•■i 

' 

•:"- 

© 

/ 

fr- 

-t 

© 

■■"! 

^ 

-- 

© 

© 

© 

© 

© 

© 

t-H     rH     CM     CO 
©     CO     CM     00     Tf 


©     CM     00     Tf     © 

co    Tf    tt    ic    © 

©     ©     CM     00     tJ< 


©    ire    ©   © 


©     fr-     Oi     ©     CM     CO 


©    ire    ©    © 


©     fr-     OS     ©     CM 


Soil  Science  405 

wide  ranges  in  weight  per  subsample  in  the  0-15  cm  and  15-30  cm 
samples  can  be  attributed  to  presence  of  brace  roots.  There  were  brace 
roots  in  some  samples  and  not  in  others.  However,  after  drying  it  was 
impossible  to  determine  rapidly  what  was  a  brace  root  and  what  was 
not.  Foth  (4)  found  that  brace  root  growth  increased  total  root  weight 
of  corn  nearly  50%  between  the  67th  and  80th  days  after  planting. 
Harvests  in  this  experiment  were  made  between  the  104th  and  109th 
days  after  planting — probably  before  total  brace  root  growth  was 
completed. 

Another  possible  explanation  in  the  wide  range  in  root  weights  is 
related  to  the  method  of  sampling.  Samples  of  soil  and  roots  were  taken 
out  of  the  hole  after  each  successive  15  cm  depth  was  reached.  This 
allowed  extra  soil  and  roots  which  fell  into  the  hole  to  be  brought  up. 
Coefficients  of  variation  of  the  dry  weights  of  soil  samples  (with  very 
small  weights  of  roots  in  them)  varied  from  14%  up  to  18%.  However, 
coefficients  of  variation  of  corn  root  weights  found  in  these  soil  samples 
ranged  from  a  low  of  62 %  to  a  high  of  125%.  These  are  extremely  high 
coefficients  of  variation.  When  variation  in  weights  of  soil  samples  and 
root  samples  is  compared,  it  is  found  that  root  weights  were  four  to 
seven  times  as  variable  as  weights  of  soil  samples.  This  indicates  that 
corn  roots  were  not  uniformly  distributed  in  the  soils  sampled  (Table  2). 

Root  weights  from  the  bucket  auger  averaged  27%  heavier  than 
those  from  the  core  sampler  in  samples  taken  to  91  cm  (36  in).  This 
difference  was  significant  at  4  to  1  odds  and  was  expected  because  of 
the  wider  cutting  edge  of  the  bucket  auger.  The  cutting  diameter  of  the 
bucket  auger  was  35%  wider  than  that  of  the  core  sampler.  However, 
the  average  weight  of  soil  and  roots  per  15  cm  increment  taken  by  the 
bucket  auger  was  699  g.  This  compared  to  465  g  for  the  core  sampler 
increment  or  50%  more.  The  difference  of  234  g  was  highly  significant. 
Root  samples  taken  by  the  bucket  auger  were  macerated  and  harder  to 
separate  than  those  taken  with  the  core  sampler.  However,  the  bucket 
auger  was  able  to  by-pass  stones  and  sample  to  a  greater  depth  in  one 
sublocation  than  was  the  core  sampler  in  Crosby  soil.  Gravel  in  pockets 
probably  laid  down  by  water  prevented  sampling  any  deeper  than  122  cm 
(48  in)  with  the  core  sampler  in  Crosby  soil.  In  the  one  sublocation 
where  it  was  possible  to  sample  to  the  desired  depth  (152  cm  or  60  in) 
on  the  Crosby  soil,  root  weights  were  less  than  the  averages  found  in 
Ragsdale  soil  at  the  same  depths.  However,  it  was  possible  to  sample 
all  depths  to  152  cm  at  all  sublocations  of  the  Ragsdale  soil.  There  were 
no  significant  differences  between  the  two  soils  in  root  weights  found  to 
91  cm.  Below  this,  other  sampling  procedures  are  needed  to  determine 
whether  or  not  Ragsdale  has  more  roots  at  greater  depths  than  has 
Crosby.  The  methods  used  in  this  study  to  sample  corn  roots  in  Crosby 
soil  below  91  cm  were  not  satisfactory. 

Yields  of  corn  grain  averaged  10,057  kg  per  ha  (160  bu  per  A)  on 
the  Ragsdale  soil  in  1968.  On  the  Crosby  soil,  yields  were  8809  kg  per 
ha  (140  bu  per  A).  The  difference  in  yields  of  1248  kg  per  ha  (20  bu 
per  A)   was  significant  at  19  to  1  odds.  Since  the  rainfall  of  2.77  cm 


406  Indiana  Academy  of  Science 

(1.09  in)  in  July  and  7.29  cm  (2.87  in)  in  August  was  deficient,  the  less 
gravelly  Ragsdale  soil  must  have  supplied  more  water  to  the  crop — ■ 
probably  because  of  deeper  root  penetration. 

Summary 

Roots  were  sampled  from  five  sublocations  of  both  a  Crosby  silt 
loam  and  a  Ragsdale  silty  clay  loam  in  the  same  field.  Two  types  of 
sampling  tools  were  used  to  obtain  vertical  soil  cores.  Subsamples  were 
taken  by  15  cm  (6  in)  increments  from  the  surface  down  to  152  cm 
(60  in)  where  possible.  The  roots  were  separated  from  the  soil.  Highly 
significant  differences  were  found  between  root  weights  from  the  0-15  cm 
(0-6  in)  depth  subsamples  and  root  weights  from  subsamples  taken 
below  that.  About  67r/<-  of  the  total  weight  of  roots  to  a  depth  of  152  cm 
was  found  in  the  0-15  cm  subsamples.  Weight  of  roots  rapidly  declined 
from  the  0-15  cm  depth  to  the  137-152  cm  (54-60  in)  depth.  Gravel  in 
the  Crosby  profiles  made  it  impossible  to  sample  deeper  than  107  cm 
(42  in)  in  4  of  the  5  sublocations.  The  standard  bucket  auger  with 
wider  effective  cutting  diameter  took  more  soil  and  with  it,  27%  more 
roots  than  did  the  Pit-O-Matic  core  sampler.  The  bucket  auger  macerated 
the  roots  while  the  core  sampler  did  not.  The  very  wide  range  in  root 
weights  taken  in  subsamples  to  30  cm  depth  (12  in)  was  partially 
explained  by  the  presence  of  brace  roots  in  some  subsamples  and  not  in 
others.  It  was  shown  that  weight  of  roots  in  soil  subsamples  was  four  to 
seven  times  as  variable  as  the  weights  of  the  soil  from  which  they 
came.  Grain  yields  of  corn  in  1968  of  10057  kg  per  ha  (160  bu  per  A) 
on  the  Ragsdale  soil  were  significantly  greater  than  the  8809  kg  per  ha 
(140  bu  per  A)  on  the  Crosby  soil.  Since  July  and  August  were  relatively 
dry,  the  less  gravelly  Ragsdale  soil  apparently  supplied  more  available 
water — probably  because  of  deeper  root  penetration. 


Literature  Cited 

1.  Fbhrenbacher,  J.  B.,  and  J.  D.  Alexander.  1955.  A  method  of  studying  corn  root 
distribution  using  a  soil-sampling  machine  and  a  shaker-type  washer.  Agron.  J. 
47:468-472. 

2.  Fehrenbacher,  J.  B.,  P.  R.  Johnson,  R.  T.  Odell,  and  P.  E.  Johnson.  1960.  Root 
penetration  and  development  of  some  farm  crops  as  related  to  soil  physical  and 
chemical  properties.  Transactions  of  the  7th  Inter.  Congress  of  Soil  Science,  Madison, 
Wise,  U.S.A.,  1960.  Vol.  111:243-252. 

3.  Fehrenbacher,  J.  B.,  B.  W.  Ray,  and  J.  D.  Alexander.  1967.  Root  development  of 
corn,  soybeans,  wheat,  and  meadow  in  some  contrasting  Illinois  soils.  Illinois  Re- 
search,   University    of    Illinois    Agricultural    Experiment    Station,    Spring    1967:3-5. 

4.  Foth,   H.   D.   1962.   Root  and  Top   Growth  of  Corn.    Agron.   J.   54  :49-52. 

5.  Raper,  Charles  David,  Jr.  1970.  Significance  of  Root  Characteristics  in  Nutrient 
Uptake  and   Growth   of   Soybeans.    Ph.D.    Thesis,   Purdue  University. 


The  Effect  of  Rainfall  Energy  on  Water  Infiltration  into  Soils1 

J.  V.  Mannering  and  D.  Wiersma,  Purdue  University 


Abstract 

Simulated  rain  was  applied  in  the  field  to  seven  Indiana  soils  ranging  in  texture 
from  a  sand  to  a  silty  clay.  Each  site  was  divided  so  that  one  half  of  the  plot  was  bare 
fallow  while  the  other  half  was  protected  by  a  layer  of  screenwire  suspended  10  cm  above 
the  surface.  A  comparison  of  the  infiltration  characteristics  of  these  soils  under  these  two 
conditions  when  exposed  to  high  energy  rain  showed  rainfall  energy  to  be  the  principal 
causative  factor  in  surface  sealing.  The  magniture  of  these  differences  was  greatly 
influenced  by  soil  texture,  however,  medium  textured  soils  were  affected  most  severely. 
After  30  minutes  of  rainfall,  infiltration  rates  on  medium  textured,  bare  soils  were  20  to 
30%  of  those  on  protected  soils. 

The  formation  of  a  layer  at  the  soil  surface  that  reduces  water 
intake  has  been  recognized  for  over  a  century  (1).  The  principal  factor 
responsible  for  the  formation  of  this  surface  seal  has  been  shown  to  be 
the  impact  of  raindrops  on  the  soil  surface  (5).  Duley  and  Kelly  (2), 
for  example,  showed  that  when  the  surface  was  protected  by  cover,  a 
broad  range  of  infiltration  rates  occurred  between  different  soil  types. 
The  differences  diminished,  however,  when  these  same  soils  were  bare 
and  subjected  to  rain.  Other  workers  (3,7)  have  reported  permeabilities 
for  surface  seals  of  1/5  to  1/2000  of  those  of  underlying  materials.  In 
the  solution  of  a  mathematical  model,  Swartzendruber  (6)  showed  that 
the  conductivity  of  the  least  permeable  layer  does  not  of  itself  control 
flow  but  is  dependent  on  the  physical  characteristics  of  both  the  least 
permeable  layer  and  the  other  material  in  the  system. 

Although  much  effort  has  been  directed  to  understanding  water 
intake  into  sealed  soil  surfaces,  the  magnitude  of  changes  in  infiltration 
rates  of  Indiana  soils  brought  about  by  rainfall  energy  remains 
unsolved. 

It  is  the  purpose  of  this  study  to  show  the  accumulative  influence 
of  rainfall  energy  on  infiltration  rates  for  a  wide  range  of  soil 
textures. 

Methods  and  Procedures 

Infiltration  rates  were  determined  by  using  the  rainfall  simulator 
described  by  Meyer  and  McCune  (4).  Two  storms  of  1-hour  duration  at 
an  intensity  of  7.0  ±  0.5  cm  per  hr  were  applied  on  successive  days  to 
soils  that  had  been  maintained  in  a  fallow  condition  for  several  months. 
Two  treatments  or  conditions  were  used  to  measure  the  influence  of 
high  energy  rainfall  on  infiltration  rates  of  fallow  (bare)  soils:  1) 
rainfall  of  high  kinetic  energy  was  applied  to  a  bare,  unprotected  soil, 

1  Contribution  from  Purdue  University  Agronomy  Department,  Lafayette,  Indiana. 
Published  with  the  approval  of  the  Director  of  the  Purdue  University  Agricultural  Experi- 
ment Station  as  Paper  No.  3894.  The  authors  wish  to  acknowledge  the  Soil  and  Water 
Conservation  Research  Division,  Agricultural  Research  Service,  USDA  for  their  con- 
tribution in  this  study. 

407 


408  Indiana  Academy  of  Science 

and  2)  rainfall  with  low  kinetic  energy  was  applied  to  this  same  soil. 
This  was  accomplished  in  the  following  manner.  Prior  to  the  application 
of  simulated  rainfall,  one  plot  (12  x  35  feet  in  size)  was  covered  with 
a  double-layer  of  18  x  14  mesh  screenwire.  This  screenwire  was  sus- 
pended 10  cm  above  the  soil  surface  to  reduce  drop  size  and  velocity 
(kinetic  energy)  of  the  rain  without  obstructing  overland  flow  or 
runoff.  A  second  adjacent  plot  was  left  bare  and  unprotected. 

This  study  was  conducted  on  seven  Indiana  soils  (Table  1)  ranging 
in  texture  from  a  sand  to  a  silty  clay.  Some  of  the  relevant  soil 
properties  are  shown  in  Table  1. 

Table  1.     Properties  of  soils  tested. 


Organic 

Aggre- 

Sand 

Silt 

Clay 

Matter 

gation 

(%) 

(%) 

(%) 

(%) 

Index 

Oakville  sand 

94 

4 

2 

0.5 

0.065 

Fox  gravelly  sand  loam 

80 

12 

8 

1.1 

0.181 

Warsaw  sandy  loam 

62 

25 

13 

3.3 

0.408 

Fox   silt  loam 

22 

57 

21 

1.3 

0.494 

Zanesville  silt  loam 

0 

72 

10 

1.3 

0.160 

Cincinnati   silt  loam 

0 

72 

19 

1.3 

0.205 

Markland  silty  clay 

4 

55 

41 

3.0 

0.986 

The  kinetic  energy  of  the  rainfall  from  the  simulator  is  approxi- 
mately 800  ft  tons  per  acre  inch  (4).  The  kinetic  energy  of  the  same 
intensity  rainfall  falling  through  a  double-layer  of  screenwire  is  not 
known.  However,  it  has  been  observed  by  means  of  high-speed  pho- 
tography that  the  screenwire  effectively  disperses  large  drops.  As  a 
consequence,  the  rainfall  energy  of  single  drops  is  expended  over  a 
much  larger  area  and  the  velocity  of  a  sizable  portion  of  the  falling 
drops  is  reduced.  Both  of  these  factors  would  greatly  reduce  the 
kinetic  energy  of  the  falling  rain  and,  therefore,  its  dispersive  power  in 
the  formation  of  surface  seals. 

Results  and  Discussion 

The  effects  of  the  rainfall  energy  on  infiltration  rates  of  the  various 
soils  over  a  2-hour  period  are  shown  in  Figures  1  through  7. 

The  protective  screenwire  is  shown  to  have  very  little  influence  on 
the  infiltration  on  the  Oakville  sand  during  the  first  60-minute  storm 
(Fig.  1)  since  on  both  the  protected  and  unprotected  plots  essentially 
all  of  the  rain  applied  entered  the  soil.  This  was  true  for  the  protected 
plot  throughout  the  2-hour  storm.  However,  the  infiltration  on  the  bare 
plot  was  reduced  to  70 %  of  that  on  the  protected  plot  at  the  end  of 
2  hours. 


Soil  Science 


401) 


More  evidence  of  surface  sealing-  occurred  on  the  Fox  gravelly 
sandy  loam  (Fig.  2)  when  the  protective  cover  permitted  essentially  all 
water  applied  during  the  2-hour  storm  to  enter  the  soil.  On  the  other 
hand,  infiltration  rates  on  the  unprotected  plots  were  only  55%  and  32%, 
respectively,  of  that  on  the  protected  plots  at  the  end  of  1  and  2  hours. 
The  formation  of  the  surface  seal  on  the  bare  plot  had  largely  occurred 


id      in      ^t      io 
(jy/wo)  3ivH  NOUVUmJNI 


r--       co       u~>      <3-       ro      cm       —       o 
(Ji|/wo)  3iyy  NOIlVdlHIdNI 


[     I 

- 

1             o 

ID- 

Q             f 

CD 

-z. 
< 

J 

c. 

CO                  J 

p 

a 

LU 

/ 

O 

_1                  J 

A 

to  _ 

_l 

r/ 

c 

>                  * 

o 

i 

CO 

5 

a> 

0) 

c 

c  " 

0> 

(D 

i-      c 

~c 

O       a) 
CO      a) 

o  " 

o     b 

o 

2     CO 

_c  _ 

II 

C\J 

CON-tOiO^t-rOOJ       —       o 

(jq/wo)  3ivH  NOIlVdllldNI 


h-cDLfi^rroco       —       o~" 
(jil/Luo)  3ivd  NOIlVdXllHNI 


410  Indiana  Academy  of  Science 

after  only  20  minutes  of  rain  as  evidenced  by  the  rapid  drop  in 
infiltration  rate. 

Figure  3  shows  the  effect  of  rainfall  energy  on  the  infiltration  rate 
of  a  Warsaw  sandy  loam.  Again,  the  infiltration  rate  remained  high  on 
the  protected  plot  throughout  the  2-hour  test  while  the  infiltration  rate 
on  the  sealed  plot  had  decreased  48%  and  24%,  respectively,  at  the  end 
of  1  and  2  hours.  Although  the  soils  studied  in  Figures  2  and  3  are  both 
sandy  loam  in  texture,  the  Warsaw  soil  had  approximately  twice  the 
silt  +  clay  content  of  the  Fox  soil  which  probably  accounted  for  the 
appreciably  lower  infiltration  rate  at  the  end  of  the  2-hour  storm. 

After  only  10  minutes  of  rain,  a  seal  had  formed  on  the  Fox  silt 
loam  soil  and  little  further  reduction  in  infiltration  rate  occurred  after 
30  minutes  of  rain  during  the  first  1-hour  storm  (Fig.  4).  Infiltration 
rates  on  the  bare  plots  were  35%  and  80%,  respectively,  of  those  on  the 
protected  plots  after  1  and  2  hours.  The  swelling  of  the  clay  present  is 
thought  to  be  responsible  for  the  rapid  decrease  in  infiltration  on  the 
protected  plot  during  the  second-day  test. 

Figures  5  and  6  illustrate  the  influence  of  rainfall  energy  on  the 
infiltration  rates  of  soil  high  in  silt.  When  not  protected  these  two  soils, 
Zanesville  and  Cincinnati,  containing  72%  silt,  both  exhibited  a  marked 
reduction  in  infiltration  after  only  15  minutes  of  rain.  Further  reduc- 
tions in  infiltration  rate  after  30  minutes  of  rain  were  relatively  small. 
Infiltration  rates  on  the  bare  plots  after  the  first  hour  of  rain  were 
approximately  35  %  of  those  on  protected  plots  for  both  soils.  After  2 
hours,  the  differences  between  rates  on  the  bare  and  protected  plots 
were  greater  on  the  Zanesville  (61%)  than  on  the  Cincinnati  (34%). 
The  reduced  infiltration  with  time  on  protected  plots  indicate  that 
either  internal  soil  properties  are  controlling  infiltration  rate  or  that 
the  rain  falling  through  the  screenwire  contained  sufficient  energy  to 
produce  seals  on  these  particular  soils. 

Rainfall  energy  did  not  result  in  a  major  reduction  in  infiltration 
on  the  Markland  silty  clay  (Fig.  7).  After  60  minutes  of  rain,  the  rate 
on  the  bare  plot  was  still  60%  of  that  on  the  protected  plot  and  there 
was  essentially  no  difference  between  treatments  at  the  end  of  2  hours. 
The  high  level  of  water  stable  aggregation  resulting  from  high  clay 
content  and  relatively  high  organic  matter  levels  prevented  the  forma- 
tion of  a  severe  surface  seal  on  this  soil. 

These  results  show  that  when  exposed  to  high  energy  rainfall,  all 
of  the  soils  tested  are  subject  to  surface  sealing,  thus  reduced  water 
infiltration.  The  magnitudes  of  these  differences  vary  greatly  among 
soils  and  are  to  a  great  extent  controlled  by  soil  texture  and  structure. 
The  soils  most  seriously  affected  are  the  medium  textured  soils  where 
infiltration  rates  into  exposed  soils  can  drop  to  as  low  as  Ys  those  of 
protected  soils  after  only  a  few  minutes  of  rain.  Organic  matter  and 
clay  contents  when  sufficiently  high  to  produce  stable  structure  greatly 
reduce  the  soil  sealing  effect  of  rainfall  energy. 

The  realization  that  the  rate  of  water  infiltration  into  soils  can  be 
reduced  as  much  as  four  or  five-fold  because  of  surface  sealing,  is 
essential  in  many  forms  of  hydrological  planning. 


Soil  Science 


411 


N        (D         ifi         ^t        IO        N         —         O 

(A|/LUO)  3ivd  NOIlVdllldNI 


O 

CO 

o 

O 

m 

CD 

TJ 

O 

c     - 
o 

00 

O 

CD    - 

c 

c 

CD 

E 

<v 

O  w 

3    ' 

CD  UJ 

"5 

1 

XI 

I— 

"o 

> 

a> 

O 

c 

c 

^r 

O) 

Ol 

c 

c 

CO 

CD 

D 

(juywo)  3iv&  NOIlVdllldNI 


s      w      m      <t      «      w      -      o 
(JLj/ujo)  3ivd  NOIlVdllldNI 


412  Indiana  Academy  of  Science 

Literature  Cited 

1.  Baver,  L.  D.   1956.  Soil  Physics,  Third  Edition,  John  Wiley  and  Sons,  Inc.,  New  York. 

2.  Duley,  F.  L.,  and  L.  L.  Kelly.   1939.  Effect  of  soil  type,  slope  and  surface  conditions 
on  intake  of  water.  Nebr.  A.E.S.  Res.  Bull.  112:1-16. 

3.  McIntyre,   D.    S.    1958.    Permeability   measurements   of   soil   crusts   formed  by   raindrop 
impact.  Soil  Sci.  85:185-189. 

4.  Meyer,  L.  Donald,  and  D.  L.  McCune.   1958.  Rainfall  simulator  for  runoff  plots.  Agr. 
Eng.  39  :644-648. 

5.  Neal,    J.    H.,    and   L.    D.    Baver.    1937.    Measuring   the   impact   of   raindrops.    J.    Amer. 
Soc.  Agron.  29  :708-709. 

6.  Swartzendruber,   D.    1960.   "Water  flow   through   a  soil  profile  as   affected   by  the   least 
permeable  layer.  J.  Geophys.  Res.  65(12)  :4037-4042. 

7.  Tackett,  J.   L.,  and  R.   W.  Pearson.   1964.   Some  characteristics  of  soil  crusts  formed 
by  simulated  rainfall.  Soil  Sci.  99  :407-413. 


Effects  of  Organic  Matter  on  the  Multispectral  Properties  of  Soils1 

M.  F.  Baumgardner,  S.  Kristof,  C.  J.  Johannsen,  and  A.  Zachary, 

Purdue  University 


Abstract 

The  use  of  data  obtained  from  an  airborne  optical-mehanical  scanner  in  determining 
the  organic  matter  content  of  surface  soils  is  presented.  Multispectral  data  covering  the 
electromagnetic  spectrum  from  0.32  to  14  microns  was  recorded  on  magnetic  tape  in 
analog  form.  After  conversion  to  digital  form,  the  data  were  analyzed  by  computer  to 
provide  pictorial  printouts  of  soils  areas.  The  computer  was  trained  to  recognize  the 
spectral  responses  of  different  levels  of  organic  matter  of  surface  soils  and  produce  maps 
showing  the  locations  of  organic  matter  content.  The  computer  maps  were  compared 
with  results  from  approximately  200  surface  soil  samples  analyzed  for  organic  matter. 
These  samples  were  collected  in  a  25-hectare  field  in  the  center  of  a  flight  line  in  Tippe- 
canoe   County,    Indiana. 


Introduction 

For  many  years  the  soil  scientist  has  been  using"  a  color  designa- 
tion as  part  of  his  description  of  the  various  horizons  of  the  soil  profile. 
The  color  designation  commonly  used  in  many  countries  in  the  world  is 
the  Munsell  notation  system  (5). 

From  a  certain  color  designation  the  soil  scientist  can  make  a 
number  of  inferences  about  other  properties  of  the  soil.  He  relates 
variability  of  the  soil  color  to:  1)  organic  matter  content,  2)  the 
presence  or  absence  of  oxidized  or  reduced  iron  compounds,  3)  the 
internal  drainage  characteristics  of  the  soil,  and  even  to  a  certain 
extent  4)  the  potential  productivity  of  the  soil. 

Attempts  have  been  made  to  quantify  the  soil  color  measurements 
and  to  relate  these  measurements  quantitatively  with  other  soil 
properties.  Many  of  these  attempts  have  met  with  little  success  because 
of  the  difficulties  encountered  in  making  quantitative  color  measure- 
ments. Variations  of  soil  color  are  found  when  measuring  undisturbed 
soil  samples  versus  disturbed  samples.  Surface  roughness  and  variations 
in  soil  moisture  content  also  affect  soil  color   (2). 

During  the  past  4  years  research  has  been  conducted  at  the  Labora- 
tory for  Applications  of  Remote  Sensing  (LARS)  at  Purdue  Uni- 
versity to  develop  techniques  and  instrumentation  for  measuring*  and 
characterizing  earth  surface  features  with  remote  sensing  devices  from 
aerospace  platforms.  The  use  of  an  airborne  optical-mechanical  scanner 
for  obtaining  earth  resources  data  and  the  use  of  computer  techniques 
for  reducing  the  data  have  been  described  in  numerous  papers  (2,  3, 
6,7). 


1  Agricultural  Experiment  Station  Journal  No.  3939.  This  research  was  supported 
jointly  by  the  U.  S.  Department  of  Agriculture  and  the  National  Aeronautics  and  Space 
Administration,    and    conducted   at   the    Laboratory    for    Applications    of    Remote    Sensing. 

413 


414 


Indiana  Academy  of  Science 


Kristof  (4)  and  Baumgardner  et  al.  (2)  describe  the  use  of  these 
techniques  in  the  automatic  identification,  separation,  and  mapping'  of 
soil  categories  based  on  the  differences  of  spectral  properties  of  the 
surface  soils. 

The  initial  results  of  a  study  of  the  effects  of  soil  organic  matter 
on  the  spectral  properties  of  soils  are  reported  in  this  paper.  Data 
essential  to  this  study  were  obtained  with  an  airplane  containing  an 
optical-mechanical  scanner  flown  by  the  University  of  Michigan.  The 
data  were  analyzed  by  computer  techniques  at  LARS. 


Materials  and  Methods 

General   soils   studies   have   been   conducted   on    data    obtained   from 
flight    lines    flown    in    Tippecanoe    County,    Indiana,    during    May,    1969. 

TIPPECANOE    COUNTY,    INDIANA 
21       22  23  24 


25  ttrfcR 


Figure  1.  FUghtlincs  fioivn  for  scanner  data  collection  and  location  of  Soil  Area  D. 


Soil  Science 


415 


Sample   Locations  -  Dieterle  Farm 


it 


Figure  2.    Soil  sampling  locations  on  Test  Site  D. 


f"  C°U"t?'  heS  m  a  transitional  zone  between  prairie  and  forest 
so  s.  In  the  northwestern  part  of  the  County,  soils  were  formed  under 
tall  prarne  grasses.  Soils  of  the  southeastern  portion  were  formed 
under  deciduous  hardwood  forest.  M 

A  25-heetare  field  was  selected  for  more  intensive  soil  studies    This 
field    shown  as  test  site  D  in  Figure  1,  is  located  in  the  native  fore 
yegetat.on  area  and  has  surface  soil  patterns  typical  of  the  Hapludalfs 
(gray-brown  podzols).  The  field  is  a  portion  of  the  SEV4,  Sec.  6    T21N 
R3W,  Tippecanoe  County,  Indiana. 


416  Indiana  Academy  of  Science 

Scanner  and  photographic  data  were  obtained  on  the  flight  lines 
shown  in  Figure  1  on  May  26,  1969,  between  1100  and  1300  hours. 
Ground  truth  data  showed  that  test  site  D  had  been  plowed  and  disked 
in  preparation  for  planting  corn  and  soybeans.  Since  there  was  no  sur- 
face vegetative  cover,  the  soil  patterns  were  clearly  defined  and  easily 
discernible  by  visual  means. 

Within  a  few  days  after  the  scanner  flight,  a  sampling  grid  plan 
was  drawn  (Fig.  2)  and  1-kg  surface  soil  samples  were  obtained  at 
32m  intervals.  A  total  of  197  samples  was  taken  from  the  surface  to  a 
maximum  depth  of  2  cm. 

One  of  the  objectives  of  this  research  was  to  study  the  relation- 
ship between  the  organic  matter  content  of  surface  soils  and  the  spec- 
tral response  patterns  in  the  spectral  range  from  0.40ft  to  2.6/x.  Ac- 
cordingly, organic  matter  determinations  were  made  on  each  of  the 
197  samples  by  the  Walkley  and  Black  method  (8). 

It  was  theorized  that  if  the  correlation  between  these  two  variables 
was  good,  then  the  scanner  data  could  be  used  in  the  rapid  preparation 
of  computer  printouts  or  maps  of  fields  showing  different  levels  of 
organic  matter  content  in  the  surface  soils. 

The  scanner  data  used  in  this  study  was  obtained  at  an  altitude  of 
approximately  1300  m  (4000  feet).  The  scanner  data  were  digitized  at 
an  interval  such  that  each  sampling  point  on  one  scan  line  would  repre- 
sent the  average  energy  of  a  specific  area  on  the  ground.  At  the  altitude 
of  1300  m,  each  sampling  point  or  resolution  element  represented  ap- 
proximately 16  to  24  m3  (150  to  200  ft2). 

To  study  the  correlation  between  organic  matter  content  and  spec- 
tral response,  it  was  necessary  to  obtain  a  quantitative  spectral  value 
in  each  wavelength  channel  which  was  representative  of  the  area  from 

Table  1.     Wavelength    ranges    in    12    channels    of    the    University    of 
Michigan  optical-mechanical  scanner. 

Channel  Number  Wave  length  Range  (microns) 


1  0.40-0.44 

2  0.46-0.48 

3  0.52-0.55 

4  0.55-0.58 

5  0.58-0.62 

6  0.62-0.66 

7  0.66-0.72 

8  0.72-0.80 

9  0.80-1.00 

10  1.00-1.40 

11  1.50-1.80 

12  2.00-2.60 


Soil  Science  417 

which  each  surface  sample  of  soil  was  obtained.  On  a  gray-scale  com- 
puter printout  of  the  entire  field,  four  resolution  elements  were  selected 
to  represent  each  soil  sample  location.  The  average  radiance  values  of 
the  4  resolution  elements  for  each  surface  sample  in  each  of  12  wave- 
length channels  were  used  to  plot  against  the  organic  matter  content. 
Table  1  shows  the  wavelength  frequencies  measured  in  each  of  the  12 
channels  used  in  this  study. 

Alexander  (1)  developed  a  color  chart  for  estimating  organic 
matter  in  mineral  soils  in  Illinois.  Correlations  were  made  of  soil 
colors  (Munsell  notations)  with  laboratory  analyses  for  organic  matter 
determined  from  a  large  number  of  Illinois  soils. 

Table  2  indicates  the  five  levels  of  organic  matter  which  Alexander 
uses  in  his  color  chart.  Soils  of  the  test  site,  which  were  used  for 
analysis  with  the  multispectral  scanner  data,  contained  similar  amounts 
of  organic  matter. 

Table  2.     Levels  of  soil  organic  matter  content  determined  in  Illinois  (1). 

Average   (%)  Range  (%) 

5  3.5-7.0 

3.5  2.5-4.0 

2.5  2.0-3.0 

2  1.5-2.5 

1.5  1.0-2.0 


In  this  study  the  desired  task  was  to  obtain  a  computer  printout 
(or  map)  of  the  test  site  showing  the  locations  of  soils  having  five 
levels  of  organic  matter  similar  to  the  Illinois  study  (1).  The  organic 
matter  content  and  corresponding  spectral  responses  of  12  wavelength 
bands  for  each  soil  sample  location  were  used  to  train  the  computer.  The 
samples  were  divided  into  five  levels  of  organic  matter  content  (Table 
3).  The  computer  classified  each  resolution  element  of  each  scan  line 
in  the  test  site  using  a  pattern  recognition  technique  which  used  the 
data  provided  by  the  training  samples.  Less  than  V±  of  the  area  in  the 
test  site  was  used  to  train  the  computer. 

Table  3.     Number   of   samples   in    each   of   five    levels    of   soil    organic 
matter  used  for  training  the  computer. 


Percent 

Number  of 

Organic  Matter 

Samples 

>3.5 

46 

2.5-3.5 

37 

2.0-2.5 

18 

1.5-2.0 

63 

0-1.5 

46 

418 


Indiana  Academy  of  Science 


Results  and  Discussion 

To  provide  a  laboratory  model  of  the  soil  patterns  on  the  test  site 
area  a  soil  mosaic  was  constructed  with  the  use  of  portions  of  each  of 
the  197  surface  soil  samples.  Figure  3  contains  a  photograph  of  this 
mosaic  and  provides  a  good  general  representation  of  the  soils  pattern 


M0msmm 

■■<:■■    ?■  '     ^^^M^W0^^^^W^^^^^^^^:  ^  ■■■:■    'v 

:■'■■"-    ■:■ .  ■        ■    •'         •■**'  <■-,  - 


.mmj 


.«f 

lf>; 

si 

.....?.,■'».: 

ftr;.|  ;^'4 

::      ■  r 

.*E 

"1. 

Jil^ij^ 

^ii;> '      -  ■> 

.Hf' 

;    '* 

*Jpi 

;.|S|! 

Ji,,  ::; 

-vfcS-:;'*;i 

*            -  ;; 

. 

II1  ..;^ 

'. 

Figure  3.    General  soil  patterns  of  Test  Site  D. 


Soil  Science 


419 


mmsmm 
««lilimi^ 

^filillliligi.,, 

/■iiiiiSilti. 

■;  f%%j .itlfllJIS  i 


mmmmmi 

...........,,...■.......'......... 


f^sx  x  a 


•    3LLLf  f 

-  '   •'  .    .    /  /   ft--   :/—/  , 

**////// 


iflsifi 


I//T/7 

a  a  *  ig  $g 

s«;^a«~~«~«..^  —  «. 
X  « '38  <~  —  —  ->  ~.  «« -» ». 
a**—  *~~«a/*aa«™,a~. 

|^r~«*~ib  s  »»-«.*«*/// ^ 

>»-«»*«*««»////// 

•/-*«**«««////£/ 

s*  —««»*«/////// 

s»  a  »*«**'*  *  s*  «»«/ 

s *a a® »»*» *«*«/// 
K«a»**aa»//«// 


,     .<rv*  **.*.; 

*.  ->'  A  #  r  t  t  v  -,  v  . ... 

<  <  X  /  *  «•   *■  *  <   i  «  *  .<  *  r  ,• 

777/ /  -    •       '        , 

-«*//// 

.'—  ■ 

sass*///** 

/ 


*      ,.     *    *     V    * 

..,■.*  ■>■  •    :iy*:*^  *'■*  ;■■  **  v 

♦Illy  /fli;ii§§!.  / 
'      ■  •  •  •  '    --  /// 

Tiiii^afcii 

/lii 


'//  / 


*3h 


M 


****** 


***; 


!t#**i 


Ifr 


X  k  ^  * 


•V 


jiiiiiiti 


// 


"ssi 


"    .      <     W     .      .      .    ,  ,.  ,  ,      , 

Figure  4.     Photograph    of    computer   printout    of    general    soil    patterns    at    Test    Site    D. 


420 


Indiana  Academy  of  Science 


of  the  field.  A  photograph  of  a  computer  printout  (Fig.  4b)  showing 
five  different  levels  of  organic  matter  in  the  field  is  shown  for  compari- 
son with  the  soil  mosaic.  This  organic  matter  map  compares  very  well 
with  the  color  patterns  observed  in  the  soil  mosaic.  The  depressional 
soils  which  have  the  greatest  amount  of  organic  matter  lie  near  the 
broad  drainage  ditch.  In  the  lower  right  hand  area  of  the  illustration  a 
high  organic  matter  content  is  also  indicated.  This  is  a  slight  depression 
and  natural  drainageway. 


190 


170 


150 


130 


I  10 


90 


70 


00 


1 


000     00 
0     00000 

0           0   0 

0 

0        00   0 

0   0 

0           0  0    0 

0 

0  0    0  00 

00     0 

0 

00                 00 

0 

0 

0 

0 

0 

.    0 
Oo      0 

0 

o              r.-o.74 

0 

0 

0 

0 

0 

0 

0 

0      0 

0      0 

0 

0 

0      0 

8°o 

00                 0 

00      0 

0    0 

00 

DO    0      0 

00  00 

0       0 

0    000 

00              o         o 

00o 

0     0    00  0 
0        0      0 

000 

00            0            0    0  0            0           0 
DO            00000                       0 
000         0            00000 

0    00      0    0        00    0   00    00 

0 

oo                          oo 

,0             0                         o   00 

0.0      0.5       1.0       1.5       2.1      2.6      3.1       3.6     4.1       4.6      5.2 
Organic   Matter   Content     (Percent) 


5.7      6.2 


P^igure  5.     Correlation    between    average    radiance    level    (relative    response)    and    organic 
matter  content  for  197  samples  from  Test  Site  D. 


The  test  site  contains  undulating  land.  The  lighter  areas  to  the 
left  and  right  of  the  center  of  the  illustration  represent  sloping  topog- 
raphy. In  some  cases,  these  slopes  are  as  much  as  3  to  4  m  higher  in 
elevation  than  the  high  organic  matter  soils  along  the  drainageways. 
It  was  observed  that  with  more  detailed  examination  of  the  organic 
matter  classes,  severely  eroded  and  moderately  eroded  spots  in  the 
field  could  be  identified  on  the  computer  printouts.  In  these  instances, 
erosion  has  been  sufficiently  severe  that  subsoil  was  exposed. 

Average  radiance  levels  of  the  12  wavelength  channels  were 
plotted  against  soil  organic  matter  content  of  each  sample  location. 
Most  of  the  plottings  or  correlations  were  quite  similar.  Figure  5  pre- 
sents data  for  channel   6    (0.62-0.66*0 .   Linear  regression   analysis   gave 


Soil  Science  421 

an  r  value  of  — 0.74.  However,  the  plotted  data  seem  to  indicate  that 
perhaps  a  linear  relationship  may  not  be  valid  over  the  range  of  organic 
matter  used.  It  appears  that  above  2.0  or  2.5%  organic  matter,  there 
may  be  a  linear  correlation  with  a  much  higher  r  value.  Below  2.0% 
organic  matter,  the  curve  becomes  much  steeper. 

It  appears  that  organic  matter  plays  a  dominant  role  in  bestowing 
spectral  properties  upon  soils  when  the  organic  matter  content  exceeds 
2.0%.  As  the  organic  matter  drops  below  2%  it  becomes  less  effective 
in  masking  out  the  effects  of  other  soil  constitutents  such  as  iron  or 
manganese  on  spectral  response  of  soils. 

Training  samples  which  are  used  for  the  test  site  were  also  used 
for  obtaining  a  5-level  organic  matter  map  of  the  entire  north-south 
flight  line  across  Tippecanoe  County.  The  area  covered  was  approxi- 
mately 24  miles  long  and  1  mile  wide.  The  results  look  very  promising. 
More  research  is  necessary,  however,  to  check  the  accuracy  of  the 
results  and  to  determine  the  proper  methods  for  determining  organic 
matter  levels  with  the  use  of  the  computer.  Size  of  the  training  samples 
is  also  being  evaluated. 

Summary  and  Conclusions 

Results  of  this  research  have  well  established  the  feasibility  of 
using  multispectral  scanner  data  and  computer  analysis  to  prepare 
maps  which  outline  surface  soils  with  varying  levels  of  organic  matter. 
Surface  soil  samples  were  obtained  from  a  test  site  area  and  organic 
matter  content  was  determined.  Linear  regression  of  organic  matter 
content  and  wavelength  response   gave   a  r  value   of  — 0.74. 

More  research  is  needed  to  define  more  precisely  the  effects  of 
and  interactions  between  organic  matter,  iron,  and  aluminum  com- 
pounds, surface  roughness,  and  moisture  as  these  factors  affect  the 
spectral  properties  of  soils. 

An  operational  capability  to  map  five  or  more  levels  of  soil  organic 
matter  in  a  rapid  manner  would  provide  a  very  useful  tool  to  the  soil 
surveyor,  the  urban  and  regional  planner,  the  agricultural  chemical 
industry,  the  drainage  engineer,  the  farm  manager,  and  many  others. 


Literature  Cited 

1.  Alexander,  J.  D.   1968.  Color  chart  for  estimating  organic  matter  in  mineral  soils  in 

Illinois.  University  of  Illinois,  Urbana. 

2.  Baumgardner,  M.  F.,  C.  J.  Johannsen,  and  S.  J.  Kristof.  In  press.  Multispectral 
studies  of  soils  and  clay  minerals.  Proc.  German  Soil  Sci.  Soc.  Meetings.  Hannover, 
W.  Germany.  September,  1969. 

3.  Johannsen,  C.  J.,  and  M.  F.  Baumgardner.  1968.  Remote  Sensing  for  planning 
resource  conservation.  Proc.  Soil  Conserv.  Soc.  of  Amer.  Athen,  Ga.,  August,  1968,  p. 
149-155. 

4.  Kristof,  S.  J.  In  press.  Preliminary  Multispectral  Studies  of  Soils.  J.  Soil  and  Water 
Conserv. 


422  Indiana  Academy  of  Science 

5.  Munsell,  A.  H.  1947.  A  color  notation.  Ed.  10.  Munsell  Color  Company,  Baltimore. 

6.  Laboratory  for  Agricultural  Remote  Sensing.  Remote  Multispectral  Sensing  in  Agricul- 
ture. 1967.  Vol.  2    (Annual  Report).  Res.  Bui.  No.  832,  Purdue  University. 

7.  Laboratory  for  Agricultural  Remote  Sensing.  Remote  Multispectral  Sensing  in  Agricul- 
ture. 1968.  Vol.  3   (Annual  Report).  Res.  Bui.  No.  844,  Purdue  University. 

8.  Walkley,  A.,  and  I.  A.  Black.  1934.  An  examination  of  the  Degtjareff  method  for 
determining  soil  organic  matter  and  a  proposed  modification  of  the  chromic  acid  titra- 
tion method.  Soil  Sci.  37  :29-38. 


A  Two- Year  Study  of  Bacterial  Populations  in  Indiana 
Farm  Pond  Waters1 

L.  B.  Hughes  and  H.  W.  Reuszer,  Purdue  University 


Abstract 

A  two-year  study  of  bacterial  populations  in  farm  pond  waters  was  conducted  on 
three  farm  ponds  on  the  Southern  Indiana  Purdue  Agricultural  Center.  Samples  of  water 
from  the  surface  and  near  the  bottom  were  taken  approximately  once  per  month  from 
each  pond.  Bacterial  numbers  were  determined  by  colony  counts.  Great  differences  in 
bacterial  numbers  were  found  between  ponds  and  large  seasonal  variations  in  bacterial 
numbers  were  found  within  ponds.  Maximum  bacterial  numbers  occurred  in  late  spring 
or  early  summer  in  all  three  ponds.  Consistently  higher  numbers  of  bacteria  were 
present  in  Pond  C  and  lowest  numbers  were  present  in  Pond  B.  Water  temperatures 
were  quite  similar  in  all  three  ponds,  although  a  significant  correlation  between  water 
temperature  and  bacterial  numbers  was  found  only  in  Pond  C.  Organic  carbon  content 
was  consistently  highest  in  Pond  C  and  lowest  in  Pond  B.  Significant  positive  correlation 
coefficients  between  organic  carbon  and  baterial  numbers  were  found  in  Ponds  A  and  B, 
but  a  significant  negative  correlation  was  found  in  Pond  C. 


Introduction 

Farm  ponds  have  many  uses  in  our  day  including  serving  as  sources 
of  recreation,  sport  and  commercial  fishing,  livestock  water,  and  even 
water  for  human  consumption  and  general  household  use.  Consequently, 
the  chemical  and  biological  nature  of  farm  pond  waters  is  of  extreme 
importance.  A  thorough  search  of  the  literature  revealed  few  micro- 
biological studies  in  farm  pond  waters.  The  purpose  of  this  study  was 
to  determine  the  changes  in  numbers  of  bacteria  in  farm  pond  waters 
over  a  2-year  period  and  to  investigate  some  of  the  factors  possibly 
influencing  the  bacterial  numbers. 

Literature 

Many  workers  have  reported  large  fluctuations  in  bacterial  num- 
bers in  bodies  of  water.  Fred  et  al.  (1)  and  Snow  and  Fred  (4)  found 
large  fluctuations  of  bacterial  numbers  in  Lake  Mendota,  Wisconsin, 
in  studies  covering  4  years  and  extreme  fluctuations  could  not  always 
be  explained  (1).  In  a  brief  study  of  Flathead  Lake,  Montana,  Graham 
and  Young  (2)  found  maximum  bacterial  numbers  below  8,000  per  ml. 
Lower  numbers  of  bacteria  were  found  in  the  surface  water  than  in 
water  below  the  surface.  Stark  and  McCoy  (5)  reported  striking  dif- 
ferences in  bacterial  numbers  in  the  surface  water  at  different  loca- 
tions on  the  same  lake. 

Irwin  and  Claffey  (3)  found  wide  variations  in  numbers  of  bac- 
teria in  the  waters  of  20  ponds  in  Oklahoma  with  fewer  bacteria  found 
in    ponds    with    higher    turbidity.    Wilson    et    al.    (7)    reported    5,000    to 


1  Journal  Paper  No.  3889,  Purdue  University  Agricultural  Experiment  Station, 
Department  of  Agronomy,  Lafayette,  Indiana.  This  study  was  supported  in  part  by  the 
Office  of  Water  Research,  U.  S.  Department  of  the  Interior. 

423 


424  Indiana  Academy  of  Science 

30,000  bacteria  per  ml  in  6  West  Virginia  ponds.  Little  differences  in 
numbers  of  bacteria  were  detected  at  different  depths.  Water  tempera- 
ture and  pH  showed  no  correlation  to  the  bacterial  numbers. 

Procedures 

The  three  farm  ponds  studied  were  constructed  in  1953  or  1954 
on  the  Southern  Indiana  Purdue  Agricultural  Center.  The  ponds  were 
arbitrarily  labeled  A,  B,  and  C.  Pond  A  was  the  largest  with  a  surface 
area  of  0.96  acre,  a  maximum  depth  of  12  feet,  and  a  watershed  area 
of  7  acres.  Substantial  aquatic  plants  in  the  edge  of  Pond  A  con- 
tributed organic  matter  to  the  water.  Pond  B  had  a  surface  area  of 
0.66  acre,  a  maximum  depth  of  12  feet  and  a  watershed  area  of  2 
acres.  Very  little  aquatic  plant  growth  occurred  in  this  pond.  Pond  C 
had  a  surface  area  of  0.30  acre,  a  maximum  depth  of  15  feet  and  a 
watershed  area  of  3  acres.  Dense  growth  of  aquatic  plants  extended 
several  feet  into  the  edge  of  the  water.  Substantial  algal  growth  was 
noted  in  Ponds  A  and  C  in  late  spring  and  early  summer,  but  little 
algal  growth  was  noticed  in  Pond  B.  A  mixture  of  alfalfa  and  orchard 
grass  was  the  common  forage  crop  growing  on  the  three  watersheds  and 
provided  good  protection  against  erosion  of  the  soil. 

Samples  of  water  were  taken  from  a  raft  at  the  surface  and  6 
inches  from  the  bottom  at  4  different  locations  on  each  pond  approxi- 
mately once  per  month  over  a  2-year  period  (April,  1967  to  March, 
1969).  Samples  of  water  were  immediately  placed  in  ice  and  trans- 
ported back  to  the  laboratory  (4  hours  travel  time).  The  water  samples 
were  then  stored  overnight  at  5°  C  and  plated  the  following  day  on  an 
agar  medium  containing  1.0  gm  glucose,  1.0  gm  peptone,  0.5  gm  yeast 
extract,  0.25  gm  K2HPO,,  12.0  gm  agar,  and  1,000  ml  deionized  water. 
Bacterial  numbers  were  determined  by  colony  counts  made  after  14 
days  incubation  in  the  dark. 

Water  temperatures  were  obtained  four  times  daily  by  E.  J.  Monke 
and  P.  R.  Goodrich  of  the  Purdue  University  Agricultural  Engineering 
Department  using  thermocouples  at  various  depths.  Data  used  in  this 
paper  are  the  means  of  the  four  temperatures  recorded  on  the  day  of 
sampling.  Water  temperatures  were  available  only  from  April  to  De- 
cember, 1967.  Aliquots  of  water  were  evaporated  and  organic  carbon 
was  determined  using  the  manometric  procedure  described  by  Van  Slyke 
and  Folch  (6).  Data  on  pH,  nitrate  concentration,  and  water  turbidity 
were  obtained  from  the  Indiana  State  Board  of  Health. 

Results  and  Discussion 

The  factors  possibly  influencing  the  bacterial  numbers  that  were 
studied  include  organic  matter  content,  water  temperature  and  to  a 
lesser  extent,   pH,  nitrate   concentration,   and   turbidity   of  the   water. 

Nitrate  concentration,  pH,  and  water  turbidity  of  the  three  ponds 
are  shown  in  Table  1.  The  highest  average  pH  found  in  Pond  B  would 
seem    to    be    farther    from    the    optimum    pH    for    maximum    bacterial 


Soil  Science  425 

growth  than  the  pH  in  either  Ponds  A  or  C.  Water  turbidity  was  also 
highest  in  Pond  B.  The  small  variations  in  nitrate  concentrations  would 
not  be  expected  to  have  significant  effect  on  bacterial  numbers. 


Table  1.     Turbidity,  pH  and  nitrate  concentration  of  the  pond  water, 

at  different  times." 


April 

August 

December 

April 

July 

Avg. 

1967 

1967 

1967 

1968 

1968 

pH 

Pond   A 

7.4 

6.8 

8.0 

7.1 

7.0 

7.3 

Pond    B 

7.4 

8.0 

7.4 

7.7 

7.8 

7.7 

Pond   C 

7.3 

7.1 

7.4 

7.6 

7.3 

7.3 

Nitrate  Concentration  (ppm) 

Pond   A 

0.1 

0.1 

0.3 

0.1 

0.1 

0.14 

Pond   B 

0.1 

0.2 

0.2 

0.1 

0.1 

0.14 

Pond   C 

0.1 

0.2 

0.3 
Turbidity 

0.1 

0.2 

0.18 

Pond  A 

0.3 

0.1 

10 

2 

0.3 

2.5 

Pond  B 

15 

0.1 

3 

15 

3 

7.2 

Pond   C 

0.7 

0.2 

3 

1 

0.0 

1.1 

2  The   authors   express   gratitude   to    the    Indiana    State    Board   of    Health   for    use   of 
these  data. 


Water  temperatures  of  the  3  ponds  for  a  9-month  period  are  shown 
in  Figure  1.  Surface  water  temperatures  were  quite  similar  in  all  3 
ponds,  reaching  a  maximum  near  80° F  in  June,  and  a  minimum  near 
40° F  in  December.  The  maximum  bottom  temperatures  in  Ponds  A  and 
B  were  above  70  °F  in  August.  Cooling  began  one  month  earlier  in  Pond 
C.  Minimum  bottom  water  temperatures  occurred  in  December  and 
were  about  equal  to  surface  temperatures  at  that  time. 

Seasonal  variations  in  organic  carbon  content  are  shown  in  Figure 
2.  The  highest  organic  carbon  content  generally  occurred  in  late  spring 
or  early  summer  in  both  years.  Pond  B  had  the  least  seasonal  variation 
of  organic  matter  content,  with  a  range  of  7.1  to  14.8  mg  of  organic 
carbon  per  liter  of  water  with  similar  quantities  of  organic  carbon  in 
the  surface  water  and  in  the  water  near  the  bottom.  Pond  A  showed 
larger  seasonal  variation  of  organic  matter  at  both  depths  with  organic 
carbon  quantities  ranging  from  8.0  to  24.0  mg  per  liter.  Higher  organic 
matter  content  generally  was  prevalent  near  the  bottom  than  at  the 
surface  in  Pond  A.  Both  depths  of  Pond  C  contained  the  highest  or- 
ganic matter  content  and  the  largest  seasonal  variation,  ranging  from 
11.1  to  29.2  mg  of  organic  carbon  per  liter  of  water. 


426 


Indiana  Academy  of  Science 


80 

POND  C 
• "Surface  Water 

/°^ 

~~\    °-—o  Water  6"  From  Bottom 

/U 

/          / 

/ 

-o-- 

• — <\\ 

\  \ 
\  \ 
\  \ 

60 

x" 

X 

50f 

^.  N. 

/in 

,         , 

1 

1 

>5> 

1                    1                     1                   T 

M 


Ll_ 
o 


POND  B 

• •  Surf  acen  Water 

o — o  Water  6"  From  Bottom 


POND  A 
—•Surface  Water 
o—  -o  water  6"  From  Bottom 


'A         M 
1967 


J  J         A         S         0 

TIME  OF  SAMPLING 


Figure    1.     Seasonal    variations    in    water    temperature    for    a    nine-month    period.     (The 
authors   express   gratitude    to   P.    R.    Goodrich   and   E.    J.    Monke   for   use   of    these    data.) 


Bacterial  numbers  are  given  in  Table  2.  Seasonal  variations  in 
bacterial  numbers  are  shown  in  Figure  3.  The  highest  numbers  of 
bacteria  generally  were  present  in  late  spring  or  early  summer  in  both 
years  of  the  study.  High  numbers  of  bacteria  seemed  associated  with 
the  active  growth  period  of  aquatic  vascular  plants  rather  than  the 
autumn  period  of  plant  death  and  decay.  The  role  of  algae  with  respect 
to  numbers  of  bacteria  was  not  clear.  Pond  A  had  bacterial  maxima  at 
the  same  time  each  year.  The  surface  water  usually  had  fewer  bacteria 
than  the  bottom  water.  Bacteria  in  Pond  A  ranged  from  4,400  to  102,700 


Soil  Science 


427 


POND  C 

• 'Surface  Water 

0---0  Water  6"  From 
Bottom 


AMJJASONDJFMAMJJASONDJFM 

POND  B 

• •  Surface  Water 

0---0  water  6"  From 
Bottom 


AMJJASONDJFMAMJJASONDJFM 
1967  1968  1969 

TIME  OF  SAMPLING 

Figure  2.     Seasonal    variations    in    organic    carbon    content    of    the    pond    waters    for    a 
two-year  period. 


per  ml  in  the  surface  water  and  from  5,100  to  118,100  in  the  bottom 
water.  Pond  B  had  only  small  variations  in  bacterial  numbers  with 
similar  numbers  of  bacteria  at  both  depths  throughout  the  2-year 
period.  Bacteria  in  Pond  B  ranged  from  4,200  to  48,000  per  ml  in  the 
surface  water  and  from  3,400  to  28,300  per  ml  in  the  water  near  the 
bottom.  Pond  C  had  the  highest  numbers  of  bacteria  and  the  largest 
seasonal  variations  of  bacterial  numbers.  In  Pond  C,  the  bacteria  ranged 


428 


Indiana  Academy  of  Science 


Table  2.     Seasonal  variations  of  bacterial  number 
for  a  two-year  period. 


in  thousands  per  ml 


Pond  A 

Pond  B 

PondC 

Month 

Upper 

Lower 

Upper 

Lower 

Upper 

Lower 

April  1967 

20.7 

28.3 

14.2 

10.4 

7.7 

3.2 

May 

94.1 

110.6 

48.0 

10.8 

17.0 

18.2 

June 

70.6 

40.1 

11.8 

12.5 

160.0 

___ 

July 

11.1 

22.4 

6.2 

11.6 

9.9 

17.6 

August 

4.8 

9.7 

5.3 

10.2 

13.3 

19.2 

September 

4.4 

7.8 

5.0 

4.2 

10.9 

24.5 

October 

10.4 

12.2 

9.6 

6.9 

15.0 

3.6 

December 

15.1 

16.3 

5.7 

6.4 

25.2 

31.3 

February   1968 

15.9 

20.1 

15.9 

15.4 

77.8 

75.1 

April 

6.8 

20.1 

16.9 

20.6 

64.6 

75.6 

June 

102.7 

118.1 

10.6 

28.3 

109.3 

48.4 

July 

46.7 

42.8 

25.2 

12.0 

102.5 

37.3 

August 

16.6 

32.7 

25.1 

24.1 

68.4 

56.4 

September 

11.6 

14.3 

7.1 

27.7 

18.8 

26.7 

November 

5.3 

15.5 

10.2 

9.6 

13.1 

41.3 

December 

4.5 

5.1 

4.2 

3.4 

10.4 

8.8 

January 

45.2 

47.2 

5.8 

6.5 

111.2 

121.0 

March 

42.7 

44.1 

9.2 

8.5 

25.7 

45.8 

from  7,700  to  160,000  per  ml  in  the  surface  water  and  from   3,200   to 
121,000  per  ml  in  the  bottom  water. 

Many  chromogenic  bacterial  colonies  appeared  on  the  plates  in- 
cluding red,  pink,  shades  of  yellow  and  orange,  white  and  cream  colored. 

A  statistical  analysis  of  variance  of  the  bacterial  numbers  is  given 
in  Table  3.  In  each  of  the  ponds,  the  numbers  of  bacteria  varied  sig- 
nificantly  with   time   of   sampling,   location   of   sampling,   and   depth   of 


Table  3.     Analysis  of  variance  of  bacterial  numbers. 


Source  of 

F  Values 

Variation 

Pond  A 

Pond  B 

Pond  C 

Time   of   Sampling 

859.41** 

276.68** 

681.51** 

Location  of  Sampling 

55.58** 

49.72** 

23.15** 

Depth  of  Sampling 

74.58** 

3.49** 

20.73** 

Time  x  Location 

40.72** 

55.07** 

19.76** 

Time  x  Depth 

24.61** 

186.74** 

100.26** 

Location  x  Depth 

24.87** 

47.92** 

6.76** 

Time  x  Depth  x  Location 

18.67** 

61.62** 

14.79** 

**  Significant  at  the  1%  level. 


Soil  Science 


42'.) 


POND  C 
•  Surface  Water 
°  Water  6"  From 
^ov  Bottom 


r=      AMJJASONDJFMAMJJASONDJFM 


en 


POND  B 
-•  Surfacei  Water 
-o  Water  6"  From 
Bottom 


A  M  J 
1967 


JASONDJFMAMJJASONDJFM 
1968  1969 

TIME  OF  SAMPLING 


Figure  3.    Seasonal  variations  in  bacterial  numbers  for  a  two-year  period. 


sampling.  Table  4  shows  correlation  coefficients  for  organic  carbon  and 
water  temperature  with  bacterial  numbers  in  each  of  the  ponds.  Corre- 
lation coefficients  between  water  temperature  and  bacterial  numbers  were 
not  significant  in  Ponds  A  and  B  but  were  highly  significant  in  Pond  C. 
The  correlation  coefficients  between  organic  carbon  and  bacterial  num- 
bers were  highly  significant  in  both  Ponds  A  and  B  while  a  negative 
correlation  was  significant  at  the  57c  level  in  Pond  C. 


430 


Indiana  Academy  of  Science 
Table  4.     Correlation  coefficients 


Organic 

Watsr 

Bacterial 

Carbon 

Temperature 

Numbers 

Pond  A 

Organic    Carbon 

1.000 

.445 

.440** 

(580) 

(260) 

(580) 

Water  Temperature 

.445 

1.000 

.042 

(260) 

(320) 

(320) 

Bacterial  Numbers 

.440** 

.042 

1.000 

(580) 

(320) 

(720) 

Pond  B 

Organic  Carbon 

1.000 

.122 

.299** 

(580) 

(260) 

(580) 

Water  Temperature 

.122 

1.000 

—.005 

(260) 

(320) 

(320) 

Bacterial  Numbers 

.299** 

—.005 

1.000 

(580) 

(320) 

(720) 

Pond  C 

Organic  Carbon 

1.000 

.355 

—.115* 

(560) 

(260) 

(560) 

Water  Temperature 

.355** 

1.000 

.325** 

(240) 

(300) 

(300) 

Bacterial   Numbers 

—.115* 

.325** 

1.000 

(560) 

(300) 

(700) 

*  Significant  at  5%  level. 

**  Significant  at  1%  level. 

Values  in  parentheses  indicate  number  of  samples. 


Summary 

Seasonal  variations  of  bacterial  numbers  were  found  in  a  2-year 
study  of  three  Indiana  farm  pond  waters.  Statistical  analysis  showed 
significant  variation  of  numbers  of  bacteria  in  each  pond  with  time 
of  sampling,  location  of  sampling,  and  depth  of  sampling.  In  addition, 
correlation  coefficients  for  water  temperature  with  bacterial  numbers 
were  significant  in  Pond  C  and  not  significant  in  Ponds  A  and  B.  Cor- 
relation coefficients  for  organic  matter  content  with  bacterial  numbers 
were  significant  in  Ponds  A  and  B,  but  a  significant  negative  correlation 
co-efficient  was  found  in  Pond  C. 


Soil  Science  431 

Literature  Cited 

1.  Fred,  E.  B.,  F.  C.  Wilson,  and  A.  Davenport.  1924.  The  distribution  and  significance 
of  bacteria  in  Lake  Mendota.  Ecology  5  :322-339. 

2.  Graham,  V.  E.,  and  R.  T.  Young.  1934.  A  bacteriological  study  of  Flathead  Lake, 
Montana.  Ecology  15:101-109. 

3.  Irwin,  W.  H.,  and  J.  Claffey.  1968.  Soil  turbidity,  light  penetration  and  plankton 
populations  in  Oklahoma  ponds  and  lakes.  Proc.  Okla.  Acad.  Sci.  47  :72-81. 

4.  Snow,  Laetitia  M.,  and  E.  B.  Fred.  1926.  Some  characteristics  of  the  bacteria  of 
Lake  Mendota.  Wis.  Acad.  Sci.,  Arts  and  Letters.  22:143-154. 

5.  Stark,  W.  H.,  and  Elizabeth  McCoy.  1938.  Distribution  of  bacteria  in  certain  lakes 
of  northern  Wisconsin.  Zentrbl.  Bakt.  Abt.  II.  98  :201-209. 

6.  Van  Slyke,  D.  D.,  and  J.  Folch.  1940.  Manometric  carbon  determination.  J.  Biol. 
Chem.  136:509-551. 

7.  Wilson,  H.  A.,  T.  Miller,  and  Rosa  Thomas.  1966.  Some  microbiological,  chemical 
and  physical  investigations  of  farm  ponds.  West  Virginia  Agriculture  Experiment  Sta- 
tion Bull.  522T.  p.  1-17. 


Adsorption  of  Insecticides  on  Pond  Sediments  and  Watershed  Soils1 
N.  L.  Meyers,  J.  L.  Ahlrichs  and  J.  L.  White,  Purdue  University 

Abstract 

Adsorption  of  malathion,  phorate,  and  carbaryl  was  studied  on  pond  sediments  and 
watershed  soils.  Mineralogical  characterization  showed  the  clay  fractions  of  both  the 
sediment  and  soil  to  contain  kaolinite,  micaeous  minerals  and  vermiculite.  Adsorption 
studies  with  the  soils  revealed  malathion  was  adsorbed  to  the  greatest  extent  followed 
by  carbaryl  and  then  phorate.  Since  adsorption  tends  to  retain  pesticides  in  soil,  the 
probability  of  contamination  of  pond  water  following  application  of  these  insecticides  is 
slight. 

Introduction 

The  pollution  of  water  resources  by  pesticides  and  other  organo- 
toxicants  has  received  much  attention  since  contamination  of  the  Ten- 
nessee River  by  toxaphene  in  1951  as  reported  by  Young  and  Nicholson 
(7).  Entrance  of  a  pollutant  into  water  from  use  on  agricultural  land  is 
regulated  by  the  factors  controlling  the  fate  of  pesticides  in  the  soil. 
Removal  of  the  pesticide  from  the  soil  might  occur  through  leaching, 
volatization,  or  runoff  while  adsorption  of  pesticides  tends  to  retard 
or  prevent  removal.  In  addition,  pesticides  may  undergo  alterations  in 
the  soil  as  the  result  of  chemical,  biological,  or  photochemical  processes. 
The  ultimate  fate  of  a  pesticide  depends  on  a  combination  of  these 
parameters.  However,  adsorption  on  colloidal  surfaces  of  the  soil  ap- 
pears to  determine  to  a  greater  extent  than  any  other  single  factor 
the  ultimate  fate  of  pesticides.  The  nature  and  extent  of  adsorption 
has  been  discussed  by  Bailey  and  White   (2)   and  Meyers   (4). 

This  study  deals  with  the  adsorption  of  three  insecticides  on  water- 
shed soils  and  their  corresponding  pond  sediments.  Generally,  adsorp- 
tion of  a  pesticide  to  a  significant  degree  will  greatly  reduce,  if  not 
eliminate,  movement  into  surface  waters.  To  facilitate  the  study,  three 
small  farm  ponds  were  selected  on  the  Purdue  University  Southern 
Indiana  Forage  Farm  in  Dubois  County.  The  watershed  soil  types  were 
Zanesville  (6-18%  slope)  and  Welston  (12-18%  slope)  silt  loams  which 
have  developed  from  sandstone  and  shale.  The  soils  are  similar  except 
for  the  fragipan  formation  in  the  Zanesville  soil.  Each  of  the  water- 
sheds was  devoted  to  alfalfa  production.  Prior  to  the  application  of  in- 
secticides, soil  samples  were  collected  from  the  watersheds  and  sedi- 
ment samples  were  collected  from  the  pond  bottoms.  The  mineralogical 
characterizations  and  the  adsorption  studies  described  below  were  con- 
ducted using  these  materials.  Since  the  mineralogical  and  physical 
properties  of  the  three  ponds  were  similar,  only  results  from  one  of 
the  ponds  are  reported. 


1  Contribution  from  Purdue  University  Agronomy  Department,  Lafayette,  Indiana. 
Published  with  the  approval  of  the  Director  of  the  Purdue  University  Agricultural 
Experiment  Station  as  Paper  No.  3888.  This  study  was  supported  by  the  Office  of  Water 
Resources  Research,  Department  of  Interior. 

432 


Soil  Science 


4)5.", 


Results 
Mineralogical  Characterization 

The  mineralogical  composition  of  the  clay  fraction  of  pond  sedi- 
ment in  relation  to  its  watershed  soil  has  not  previously  been  reported 
in  Indiana.  Therefore,  the  mineralogical  composition  of  both  soil  and 
sediment   was    determined   to    provide    information    on    this    relationship 


SOIL  CLAY 


POND  I5A 
<  2.0  u 


SEDIMENT  CLAY 


Mg-SATURATED 


GLYCEROL  SOLVATED        I 


K- SATURATED 


I      i        /       K- SATURATED 
HEATED  300°C 


K-SATURATED 
HEATED  55CTC 


Figure  1.    X-ray  diffractogram  of  the  soil  and  sediment  clay. 


434 


Indiana  Academy  of  Science 


as  well  as  to  provide  a  basis  for  meaningful  interpretation  of  the 
adsorption  studies.  The  mineralogical  composition  was  determined  by 
x-ray  diffraction  and  infrared  techniques.  The  x-ray  diffractograms  of 
the  soil  and  sediment  clay  fraction  (<2.0/x)  from  the  pond  are  shown 
in  Figure  1.  The  positions  of  intensities  of  the  peaks  show  the  min- 
eralogical composition  of  the  soil  and  sediment  to  be  essentially  identi- 
cal. The  14  A  peak  present  on  Mg  saturation,  glycerol  solvation,  and 
mild  heating  was  interpreted  to  indicate  the  presence  of  vermiculite. 
The  10  A  peak  is  characteristic  of  micaeous  minerals  and  is  greatly 
enhanced  by  collapse  of  the  14  A  material  on  strong  heating  (550°  C). 
The  presence  of  kaolinite  is  confirmed  by  the  7  A  peak  remaining  on 
K  saturation  and  mild  heating  and  by  the  disappearance   of  the   7   A 


SOIL  CLAY 
POND  I5A 


SEDIMENT 
CLAY 


SOIL  CLAY 
POND  I5B 

SEDIMENT 
CLAY 


SOIL  CLAY 

POND  I3B 

SEDIMENT 
CLAY 


1020 


780 


3800 


COO 


OOO 


800 


FREQUENCY  (CM'') 


FIGURE  2.    Infrared  absorption  spectra  for  soil  and  sediment  clays. 


Soil  Science 


435 


peak  on  heating  to  550°  C  which  destroys  the  kaolinite  structure.  De- 
tailed procedures  for  the  identification  of  clay  minerals  are  given  by 
Whittig  (6). 

The  infrared  patterns  of  the  clay  fractions  are  shown  in  Figure  2. 
The  presence  of  kaolinite  is  confirmed  by  the  weak  band  at  3690  cm  " 
and  a  strong  band  at  3620  cm'.  Montmorillonite  or  vermiculite  would 
also  exhibit  a  strong  band  at  3620  cmJ  in  addition  to  a  broad  band  in 
the  3400  cmJ  region.  Details  concerning  the  application  of  infrared 
spectroscopy  to  clay  mineral  systems  has  been  given  by  Ahlrichs 
etal.  (1). 

Although  the  mineralogical  composition  of  the  clays  appears  to  be 
identical,  the  quantity  of  clay  is  much  higher  in  the  sediment  than  in 
the  soil  (Table  1).  This  is  in  agreement  with  the  work  of  Kohnke  (3) 
and  is  in  the  order  expected.  Thus,  clay  and  silt  are  preferentially 
eroded  into  the  pond  at  the  expense  of  sand  but  no  differential  erosion 
of  clay  types  occurs. 


Table  1.     Texture  of  the  watershed 

soil 

and  pond  sediment. 

%  Sand            %  Silt            %  Clay 

Coarse  clay 

(/c  of  total 

Fine  clay 
%  of  total 

Soil 
Sedin 

15.6                 68.4                 16.0 
lent        4.6                 70.4                 25.0 

63.1 

58.7 

36.9 
41.3 

600 

i               i 1 1 1 1 1 1 

- 

ADSORBENT 

O              O 
O              O 

- 

E 

v. 
Q 

MALATHION 

- 

w    300 

CD 

QL 

O 

O) 

§    200 

y' 

- 

E 

/                                                                                      CARBARYL 

- 

100 

/    ^^^                                                                     PHORATE 

(r            i               i               i               i               i               i               i               i 

10  20  30  40  50  60  70 

EQUILIBRIUM  CONCENTRATION  figm/ml 

Figure  3.  Adsorption  isotherms  for  the  three  insecticides. 


80 


436  Indiana  Academy  of  Science 

Adsorption  Studies 

Adsorption  of  malathion,  phorate,  and  carbaryl  was  studied  using 
watershed  soils  and  the  pond  sediments  as  adsorbents.  Selection  of 
these  materials  was  based  largely  on  their  current  use  on  alfalfa  for 
weevil  and  spittlebug  control.  Adsorption  of  the  insecticides  is  repre- 
sented as  Fruendlich  isotherms  in  Figure  3.  It  can  be  noted  that  mala- 
thion is  strongly  adsorbed  followed  by  carbaryl,  with  phorate  showing 
limited  adsorption. 

Comparison  of  adsorption  on  soil  with  adsorption  on  sediment 
showed  little  difference  for  malathion  and  phorate  while  carbaryl  ad- 
sorption was  greater  on  the  sediment.  Examination  of  Table  1  would 
lead  one  to  expect  greater  adsorption  on  the  sediment  due  to  the  in- 
creased clay  content.  However,  the  expected  increase  occurred  only 
with  carbaryl. 

Absolute  interpretation  of  adsorption  data  is  of  limited  value  but 
comparison  of  the  relative  extent  of  adsorption  is  worthwhile.  If  a 
pesticide  is  adsorbed  it  would  be  less  likely  to  enter  a  pond  as  com- 
pared to  a  non-adsorbed  counterpart.  We  might  expect  then  that  the 
degree  of  pollution  for  these  insecticides  following  application  to  the 
soil  would  be  in  the  order  phorate  >  carbaryl  >>  malathion.  In  addi- 
tion, adsorption  in  the  soil  under  field  conditions  should  be  more  com- 
plete since  concentrations  used  in  laboratory  studies  represent  an  appli- 
cation of  10-300  times  the  normal  rates  of  application. 

Conclusions 

Little  difference  was  found  between  the  clay  mineralogy  of  a  pond 
sediment  and  the  soil  from  which  it  is  derived.  The  soils  and  sediments 
used  in  this  study  contain  kaolinite,  micaeous  minerals,  and  vermiculite. 

Adsorption  studies  showed  that  malathion  is  adsorbed  to  the 
largest  extent  followed  by  carbaryl  and  then  phorate  with  significant 
adsorption  by  the  soil  of  all  three.  If  a  pesticide  is  adsorbed,  it  should 
not  be  subject  to  movement  into  a  pond  by  leaching  or  runoff;  thus,  we 
would  conclude  that  contamination  of  a  farm  pond  probably  would  not 
occur  following  application  of  malathion,  carbaryl  or  phorate  to  the 
watershed,  and  that  phorate  would  probably  be  the  first  to  reach  the 
pond  if  contamination  did  occur.  These  conclusions  are  based  on  the 
assumption  that  pesticide  applications  are  made  according  to  the  manu- 
facturers suggestion  and  applied  at  the  recommended  rate. 

Further  support  of  these  observations  is  given  by  the  work  of 
other  cooperators  on  the  project  (5).  Continuous  monitoring  of  the 
pond  water  for  8  months  following  application  of  phorate  and  carbaryl 
at  levels  4  times  their  recommended  dosage  showed  no  trace  of  carbaryl 
at  anytime  in  the  water  and  only  a  slight  temporary  trace  of  phorate. 
When  recommended  rates  were  applied,  no  trace  of  either  insecticide 
was  found  in  the  pond. 


Soil  Science  437 

Literature  Cited 

1.  AHLRICHS,  J.  L.,  J.  R.  Russell,  R.  D.  Harter,  and  R.  A.  Weismiller.  1965.  Infrared 
Spectroscopy  of  Clay  Mineral  Systems.  Proc.  Indiana  Acad,  of  Sci.  75  :247-255. 

2.  Bailey,  G.  W.,  and  J.  L.  White.  1964.  Review  of  adsorption  and  desorption  of  organic 
pesticides  by  soil  colloids,  with  implication  conerning  bioactivity.  J.  Agr.  Food  Chem. 
12:324-332. 

3.  Kohnke,  H.   1950.  The  reclamation  of  coal  mine  spoils.   Advances  in  Agron.   2:318-349. 

4.  Meyers,  N.  L.  1968.  Adsorption  of  organic  insecticides  on  well  characterized  watershed 
soils  and  their  corresponding  pond  sediments.  M.S.  Thesis.  Purdue  University. 

5.  Office  of  Water  Resources  Research.  1969.  Effect  of  pesticide  residues  and  other 
organic-toxicants  on  the  quality  of  surface  and  ground  water  resources.  Annual 
Report.  Purdue  University,  Lafayette,  Indiana. 

6.  Whittig,  L.  D.  1965.  X-ray  diffraction  techniques  for  mineral  identification  and 
mineralogical  composition.  In  Methods  of  Soil  Analysis.  Part  1.  Ch.  49.  Number  9  in 
the  series  Agronomy.  Amer.  Soc.  of  Agron.  Inc.,  Madison,  Wisconsin. 

7.  Young,  L.  A.,  and  H.  D.  Nicholson.  1951.  Stream  pollution  resulting  from  the  use 
of  organic  insecticides.  Prog.  Fish  Cult.  13:193-198. 


ZOOLOGY 

Chairman:  James  C.  List,  Ball  State  University 
Ralph    D.   Kirkpatrick,   Ball   State   University,  was   elected    Chairman 

for  1970 


ABSTRACTS 

Some  Modifications  in  Rat  Ovaries  and  Uteri  Following  Aminoglute- 
thimide  Treatment.  Egerton  Whittle,  Indiana  State  University. — 
Immature  female  albino  rats  (Charles  River  Strain)  were  injected 
subcutaneously  with  aminoglutethimide  phosphate  (AGP)  for  a  15-day 
period  beginning  on  day  25  post  partnm.  The  experimental  animals  were 
given  dosages  of  either  50,  100,  or  200  mg/kg  body  weight  per  day.  At 
the  end  of  the  period  of  treatment  the  ovaries  and  uteri  were  excised 
and  the  weights  compared  with  the  weights  of  ovaries  and  uteri  of 
control  animals  to  determine  whether  AGP  produced  a  definite  effect  on 
the  size  and  weight  of  these  organs  and  the  general  stature  of  the 
animals. 

The  overall  body  weight  of  the  treated  groups  displayed  an  in- 
creasing retardation  with  increasing  dose  levels  of  AGP.  The  amount 
of  weight  gain  decreased  almost  linearly  with  increased  amounts  of 
AGP  injected. 

The  effect  of  AGP  on  ovarian  weight  in  the  treated  animals  was 
that  of  an  apparent  intial  stimulation  to  growth  at  the  lower  dose  level 
but  an  abrupt  reversal  of  this  trend  when  higher  dose  levels  were  ad- 
ministered. It  would  seem  likely  that  AGP  effects  involve  more  than 
a  single  mechanism  of  action. 

The  growth  of  uteri  was  inhibited  increasingly  with  increasing 
dosage  of  AGP.  At  the  highest  dose  level  the  uterine  weight  was  only 
about  Vz  that  observed  in  control  animals. 

The  evidence  from  this  study  indicates  some  relationship  between 
presence  of  AGP  in  a  system  and  the  amount  of  estrogen  present  in  the 
system,  as  circulating  estrogen  is  the  main  governing  factor  for  de- 
velopment and  growth  of  the  organs  observed. 

Big  Brown  Bat  Eptesicus  fuscus  Movement  in  Tunnel  Cave,  Cliffy 
Falls  State  Park,  Indiana.  James  B.  Cope  and  Richard  Mills,  Earl- 
ham  College. — The  senior  author  and  students  have  studied  the  bat 
population  in  Tunnel  Cave  for  the  last  five  winters.  During  this  time, 
the  disturbance  by  banding  and  the  subsequent  reading  of  bands  caused 
unnatural  movements  in  the  cave.  Color  banding  techniques  were  tried 
which  reduced  the  disturbance  considerably  after  the  initial  banding. 
Movement  was  greater  than  expected  and  further  refinement  of  tech- 
nique was  employed. 

439 


440  Indiana  Academy  of  Science 

Every  6  hours  (  6  AM,  12  Noon,  6  pm,  and  12  Midnight)  for  a  5-day 
period  during  the  last  week  in  January,  the  cave  was  monitored  for 
banded  bats.  Flashlights  were  used  and  only  color  bands  that  were 
visible  were  recorded.  No  bats  were  disturbed  even  if  the  band  could 
not  be  read.  There  is  overwhelming  evidence  that  some  bats  come  out 
of  torpor  and  fly  during  this  period  even  when  the  outside  temperature 
isO°F. 


Parasites  of  Feral  Housemice,  Mus  musculus,  in 
Vigo  County,  Indiana 

John  0.  Whitaker,  Jr.,  Indiana  State  University 

Abstract 

The  major  external  parasites  of  the  housemouse,  Mus  musculus,  in  Vigo  County, 
Indiana,  are  Myobia  musculi,  Radfordia  affinis,  Myocoptes  musculimis,  Ornithonyssus 
bacoti,  Androlaelaps  fahrenholzi,  and  Dermacarus  heptneri  (all  are  mites),  in  approxi- 
mate order  of  decreasing  abundance.  Among  the  internal  parasites,  Heligmosomoides 
polygyrus  is  the  most  abundant,  followed  by  cestodes  (presently  unidentified  and  listed  as 
a  group),  Protospirura  sp.,  and  Syphacia  sp.  Males  and  females  harbored  similar  parasite 
infestations,  but  there  was  a  definite  increase  in  internal  parasite  load  with  increased 
age  of  the  mouse.  The  same  trend  was  apparent  in  one  species  of  mite,  Radfordia 
affinis,  but  was  not  evident  in  the  rest  of  the  species  of  ectoparasites.  Heligmosomoides 
is  primarily  a  spring  and  winter  form  and  occurred  at  its  greatest  abundance  in  the 
single  habitat  present  only  at  that  time,  winter  wheat,  while  cestodes  were  most  common 
during  the  summer  and  fall.  The  data  are  scanty,  but  suggest  Syphacia  to  be  a  spring 
and  summer  form.  Myocoptes  was  primarily  a  fall  form.  Most  external  parasites  reached 
their  greatest  abundance  in  the  fall  and  summer.  Myobia  was  most  abundant  in  winter 
wheat  and  corn,  Radfordia  in  soybeans  and  corn,  Myocoptes  in  corn  and  Ornithonyssus 
bacoti  in  sorghum.  No  relation  was  found  between  internal  and  external  parasites ;  their 
distributions  seemed  independent  of  each  other. 

Introduction 

A  number  of  housemice,  Mus  musculus,  were  taken  during  studies 
of  the  mammals  of  Vigo  County,  Indiana  (2).  Of  these,  470  from 
randomly  selected  plots  were  examined  for  external  parasites,  and  503 
for  internal  parasites.  The  present  study  was  initiated  to  determine 
if  there  were  seasonal,  sex,  age  or  habitat  differences  in  parasite  in- 
festations. 

Some  of  the  information  has  been  presented  previously,  in  a  paper 
on  the  fleas  of  the  mammals  of  Vigo  County  (3),  in  a  paper  on  the  mites 
of  the  small  mammals  of  Vigo  County  (4),  and  in  a  paper  on  the  Labi- 
dophorine  mites  of  North  America  (1). 

Donald  Norris  was  kind  enough  to  make  the  nematode  identifica- 
tions. 

Methods 

External  parasites  were  obtained  by  searching  the  fur  with  dissect- 
ing needles  and  a  10  to  60X  zoom  binocular  dissecting  microscope.  Mites 
were  cleared  and  stained  overnight  in  cold  Nesbitt's  Solution,  mounted 
in  Hoyer's  Solution  and  ringed  with  asphaltum.  Fleas  were  run  through 
the  alcohols  and  mounted  in  permount.  Internal  parasites  were  preserved 
in  70 %  alcohol  or  formal-acetic  acid  (FAA). 

Data  concerning  incidence  of  parasitism  were  presented  in  terms 
of  percentage  of  mice  infested,  with  Chi-square  being  used  to  test  for 
significant  differences  using  the  actual  numbers  of  parasitized  mice  in 
the  different  categories.  Abundance  information  was  presented  in  terms 
of  average  number  of  parasites  per  mouse,  but  this  number  often  seemed 

441 


442 


Indiana  Academy  of  Science 


of  less  value  than  the  incidence  values  because  of  the  great  amount 
of  variation  in  the  numbers  of  parasites  infesting  individual  mice,  with 
a  few  individuals  harboring  large  numbers.  For  this  reason  "t"  tests 
were  not  run. 

External  parasites 

Other  than  some  of  the  species  of  mites,  there  were  few  external 
parasites  on  the  housemouse.  One  mouse  yielded  about  15  larval  ticks,  2 
mice  each  had  1  flea,  1  had  2  lice,  while  124  mice  had  mites  (Table  1). 

Table  1.    Comparison  of  parasites  of  male  and  female  Mus  musculus. 


270  Males 

219  Females 

Infestation 

No. /Mouse 

Infestation 

No./M 
Total 

ouse 

Parasites 

No. 

% 

Total 

Avg. 

No. 

% 

Avg. 

Internal  Parasites 

Heligmosom  oides 

polygyrus 

35 

L3.0 

482 

1.70 

21 

9.5 

171 

0.78 

Cestodes 

21 

7.8 

93 

0.34 

22 

9.6 

04 

0.43 

Syphacia 

5 

1.9 

27 

0.10 

3 

1.4 

40 

0.22 

Protospirura 

5 

Lit 

6 

0.02 

0 

2.7 

0 

0.04 

Cuterebra  larvae 

1 

0.4 

1 

0.004 

0 

0.0 

0 

0.0 

Ascarid  larvae 

0 

0.0 

0 

0.0 

1 

0.5 

1 

0.005 

Heterakis 

0 

0.0 

0 

0.0 

1 

0.5 

1 

0.005 

253  Males 

214  Females 

External  Parasites 

Myobia    musculi 

2  4 

9.5 

132 

0.52 

12 

5.6 

29 

0.14 

Radfordia    affinis 

23 

9.1 

51 

0.20 

12 

5.6 

22 

0.10 

Ornithonyssus     bacoti 

16 

6.3 

35 

0.14 

2 

0.0 

3 

0.01 

Androlaelaps  fahrenholzi 

11 

4.3 

25 

0.10 

3 

1.4 

3 

0.01 

Myocoptes    musculinus 

7 

2.8 

10 

0.04 

0 

4.2 

27 

0.13 

Dermacarus     heptneri 

2 

o.s 

57 

0.22 

1 

0.5 

1 

0.005 

Dermacarus     hypudaei 

2 

o.s 

2 

0.01 

0 

0.0 

0 

0.00 

Hirstionyssus      talpae 

1 

0.4 

3 

0.01 

l 

0.5 

1 

0.005 

Eulaelaps    stabularis 

1 

0.4 

1 

0.004 

0 

0.0 

0 

0.0 

Larval  ticks 

1 

0.4 

ir> 

0.06 

0 

0.0 

0 

0.0 

Hoplopleura  cajjtiosa 

1 

0.4 

2 

0.01 

0 

0.0 

0 

0.0 

Misc.  Mites 

13 

5.1 

IS 

0.07 

11 

5.1 

15 

0.07 

Listrophorus  leuclcarti 

0 

0.0 

0 

0.00 

1 

0.5 

1 

0.005 

Orchopcas    leucopus 

0 

0.0 

0 

0.00 

1 

0.5 

1 

0.005 

Ctenojnhalmus    pseudagyr. 

tea    0 

0.0 

0 

0.00 

1 

0.5 

1 

0.005 

Androlaelaps    morlani? 

0 

0.0 

0 

0.00 

1 

0.5 

1 

0.005 

Mites,  then,  were  the  dominant  form  of  external  parasites,  with  11 
species  being  found  in  all.  Seven  of  these,  Androlaelaps  morlani  (?), 
Dermacarus  hyjmdaei  (not  previously  reported),  Eulaelaps  stabularis, 
Haemogamasus  longitarsus  and  Listrophorus  leuckarti  were  repre- 
sented by  only  one  or  two  specimens  each,  and  Hirstionyssus  talpae  by 
only  four,  hence  these  species  were  not  considered  as  important  para- 
sites of  the  housemouse  in  the  area  under  consideration. 


Zoology  443 

The  hypopial  form  of  the  mite,  Dermacarus  heptneri,  was  found  on 
3  housemice,  totaling;  58  individuals.  This  species  is  very  tiny  and  can  be 
easily  overlooked,  hence  it  may  be  more  abundant  than  indicated.  Mites 
of  this  type  cling  tenaciously  to  the  individual  hairs;  one  must  separate 
the  hairs  with  dissecting  pins  and  examine  their  bases  to  find  the 
hypopi.  D.  heptneri  was  found  on  no  other  species  of  Vigo  County 
mammal,  but  the  hypopi  of  another  species  of  Dermacarus,  D.  hypudaei, 
were  found  on  two  housemice.  D.  hypudaei  is  primarily  a  species  of 
Zapus,  but  is  also  found  on  other  species,  especially  Microtus  ochro- 
gaster  and  M.  pennsylvanicus. 

Androlaelaps  fahrenholzi,  totaling  29  individuals  on  14  mice,  was  a 
parasite  of  the  housemouse,  as  was  the  case  with  most  species  of  small 
mammals  examined.  It  occurred  at  a  lower  rate,  at  0.07  individuals  per 
housemouse,  than  on  most  of  the  other  species   (4). 

Fifteen  housemice  yielded  56  specimens  of  the  tiny  Listrophorid 
mite,  Mycoptes  musculinus.  This  species  of  mite  was  not  restricted  to 
the  housemouse,  but  all  except  seven  specimens  taken  were  from  that 
host. 

Ornithonyssus  bacoti  was  an  important  mite  on  Mus  musculus,  34 
specimens  being  taken  from  18  mice.  This  mite  occurred  at  a  similar 
rate  on  Peromyscus  maniculatus  bairdi,  and  three  specimens  were 
taken  from  P.  leucopus. 

Radfordia  affinis  was  taken  almost  entirely  on  Mus,  with  65  individ- 
uals being  taken  from  a  total  of  36  different  mice.  Two  specimens  were 
taken  from  Peromyscus  maniculatus  in  the  area  under  consideration  (4). 

Myobia  musculi,  a  tiny  white  form  similar  in  general  appearance  to 
Radfordia  affinis,  and  like  that  species  a  myobiid  mite,  was  the  most 
common  species  of  external  parasite  on  Mus  musculus  in  Vigo  County. 
It  was  taken  on  9.1%  of  the  housemice,  but  was  not  found  on  any  of  the 
other  species  of  small  mammals  examined  (4). 

The  housemouse,  in  Indiana,  appears  to  be  relatively  flealess,  only  2 
fleas  being  taken  on  the  470  housemice  examined  during  the  present 
study,  1  each  of  Orchopeas  leucopus  and  Ctenopthalmus  pseudagyrtes. 
A  third  species,  Epitedia  wenmanni,  again  one  specimen,  was  pre- 
viously reported  from  a  Vigo  County  housemouse  (3).  Likewise  Wilson 
(5)  took  only  two  fleas  from  Indiana  Mus,  one  an  Epitedia  wenmanni, 
the  other  a  specimen  of  Orchopeas  leucopus. 

One  of  the  housemice  examined  yielded  two  lice,  Hoplopleura 
captiosa,  the  only  louse  recorded  from  Mus  in  Indiana.  Wilson  (5) 
took  four  specimens  from  the  housemouse  from  Carroll  and  Tippecanoe 
Counties. 

One  housemouse  yielded  about  15  larval  ticks,  which  were  pre- 
served, but  subsequently  lost.  Wilson  (5)  reported  three  different 
individuals  of  the  tick,  Dermacentor  variabilis,  from  Mus  from  three 
Indiana  counties. 


444  Indiana  Academy  of  Science 

Internal  Parasites 

No  trematodes  or  acanthocephalans  were  taken  from  any  of  the  503 
housemice  examined  for  internal  parasites,  while  both  nematodes  and 
cestodes  were  found  to  be  relatively  common. 

Among  the  internal  parasites  the  most  abundantly  represented 
group  overall  was  the  Nematoda,  especially  Heligmosomoides  polygyrus. 
This  species  was  generally  tightly  coiled  when  seen  among  materials 
from  the  intestinal  tract,  and  was  red.  Fifty-six  of  the  mice  examined, 
or  11.1%  contained  from  one  to  over  100  individuals  of  this  species, 
totaling  653  worms,  and  averaging  1.30  worms  per  mouse.  Eight  of  the 
mice  yielded  over  25  worms  per  host.  This  species  was  found  mostly  in 
Mus,  but  a  few  individuals,  apparently  H.  polygyrus  were  found  in 
Peromyscus  maniculatus  bairdi. 

Eleven  mice  had  15  nematodes  of  the  genus  Protospirura,  nearly 
all  of  which  were  in  the  stomach  rather  than  in  the  intestine.  Female 
oxyurids,  Syphacia,  were  found  in  nine  mice,  totaling  96  specimens. 

Cestodes  were  important  as  parasites  of  Mus  musculus,  but  unfor- 
tunately these  have  not  yet  been  identified  and  are  treated  here  as  a 
group.  Forty-three  housemice  yielded  187  cestodes. 

Cuterebra,  sp.,  a  botfly  larva,  was  found  in  one  mouse;  one  mouse 
yielded  a  nematode  of  the  genus  Heterakis;  one  harbored  five  objects 
from  under  the  skin  of  the  head  which  appeared  to  be  larval  cestodes; 
and  one  yielded  a  larval  Ascarid  nematode. 

Parasites  in  Relation  to  Sex  of  Mice 

A  total  of  270  male  housemice  were  examined  for  internal  parasites, 
of  which  62,  or  22.9%,  yielded  593  parasites,  or  2.20  per  mouse.  Of  the 
210  females  examined,  47,  or  22.3%,  yielded  325  parasites,  or  1.55  per 
mouse.  Thus,  males  and  females  were  about  equally  infested  in  terms 
of  incidence,  but  the  rate  of  infestation  in  terms  of  average  number  of 
parasites  per  mouse  was  higher  in  males  than  in  females  in  this 
sample. 

The  two  main  types  of  internal  parasites  were  nematodes, 
Heligmosomoides  polygyrus,  and  cestodes.  Of  the  270  male  housemice 
examined,  35,  or  13.0%,  harbored  482  nematodes,  Heligmosomoides, 
averaging  1.79  worms  per  mouse  for  all  mice,  or  13.77  worms  per  mouse 
for  those  parasitized.  Twenty-one  of  the  219  females,  or  9.5%,  yielded 
171  Heligmosomoides ,  averaging  0.78  worms  per  mouse  overall,  and  8.1 
in  those  infested.  The  higher  average  number  in  the  males  was  pri- 
marily because  of  three  males  with  particularly  large  infestations  of 
40,  58  and  100  worms. 

A  total  of  93  cestodes  were  taken  from  21,  or  7.7%  of  the  male  Mus 
examined,  averaging  0.34  cestodes  per  mouse  overall,  or  4.43  per  para- 
sitized mouse.  In  the  females,  22  of  219,  or  10.0%,  harbored  94  cestodes, 
for  an   average   of  0.43   per  mouse,  or  4.27   in  the   mice   infested.    Thus 


Zoology  445 

males  and  females  harbored  relatively  similar  internal  parasite  loads. 
(Comparisons  for  the  remainder  of  the  individual  kinds  of  parasites  can 
be  found  in  Table  1.) 

Parasite  Infestation  and  Age  of  Animal 

To  determine  the  relationship  between  age  of  the  animal  and  para- 
site infestation,  the  mice  were  divided  into  4  groups  based  on  weight, 
those  under  10  g,  10.0  to  14.9  g,  15.0  to  19.9  g,  and  those  over  20  g. 

There  was  a  significant  increase  (Chi-square  =  28.75**,  3  df)  in  the 
incidence  of  animals  with  internal  parasites  with  increased  age  of  the 
animals,  going  from  9.1  through  16.5,  23.5  and  47.8%  of  the  mice  being 
parasitized  in  the  four  size  groups  (Table  2).  This  trend  was  apparent  in 
both  the  most  important  parasite  groups,  the  nematode  H  elig  mosomoides , 
and  in  the  cestodes.  For  Heligmosomoides,  the  largest  class  again  har- 
bored the  most  worms  per  mouse,  on  the  average,  while  the  three 
smallest  classes  were  about  the  same,  but  for  cestodes  there  was  an 
increased   mean   number   of  individuals   per   mouse   with   increased   age. 


Table  2.    Major  parasites  of  four  size  classes  of  housemice,  Mus  musculus. 


Tinder  10  g 

10.0-14.9  g 

15.0-19.9  g 

20  g 

and  over 

% 

Avg.#/ 

% 

Avg.#/ 

% 

Avg.#/ 

% 

Avg.#/ 

Parasites 

Infest . 

Mouse 

Infest. 

Mouse 

Infest. 

Mouse 

Infest. 

Mouse 

Interna]  Parasites 

Number  examined 

(66) 

(206) 

(162) 

(69) 

(503) 

All    internal    parasites 

9.1 

1.32 

16.5 

1.32 

23.5 

1.32 

47.8 

5.58 

Heligomosomoides   polygyrus 

4.5 

1.02 

9.2 

0.86 

11.7 

0.90 

21.7 

3.86 

Cestodes 

0.0 

0.00 

4.9 

0.17 

8.6 

0.35 

26.1 

1.44 

Syphacia    sp. 

3.0 

0.32 

i.r. 

(1.24 

1.2 

0.03 

2.9 

0.30 

Protospirura 

1.5 

0.02 

0.9 

1.94 

3.7 

0.05 

2.9 

0.03 

External  Parasites 

Number  examined 

(61) 

(194) 

(153) 

(62) 

All  external  parasites 

26.2 

0.51 

21.6 

0.79 

31.5 

1.14 

29.0 

1.10 

Myobia    musculi 

3.2 

0.03 

5.2 

0.16 

11.7 

0.66 

9.7 

0.44 

Radfordia  affinis 

3.2 

0.0c 

4.6 

0.11 

7.S 

0.1S 

1(1.1 

0.25 

Myocoptes  musculinus 

4.9 

0.K 

2.6 

0.04 

3.3 

0.10 

3.2 

0.08 

Ornithonyssus    bacoti 

0.0 

0.0( 

4.1 

0.07 

4.6 

0.10 

4.8 

0.10 

Androlaelaps   fahrenholzi 

4.9 

0.2« 

2.1 

0.04 

2.6 

0.03 

4.S 

0.06 

Among  external  parasites  as  a  group  there  was  no  such  apparent 
trend  in  incidence  (Table  2).  Respective  values  for  the  4  size  classes  are 
26.2,  21.6,  31.5  and  29.0%.  In  most  cases  there  was  no  definite  relation- 
ship between  parasites  and  age,  or  else  the  data  were  too  scanty  to  draw 
conclusions.  One  species,  however,  Radfordia  affinis,  did  show  the  same 
tendencies  as  the  internal  parasites.  It  is  more  apt  to  be  taken  in  the 
older  mice    (Chi-square  r=   10.10,  2  df). 


440 


Indiana  Academy  of  Science 


Parasite  Infestation  and  Season 

Data  were  summarized  (Table  3)  on  a  seasonal  basis  as  Spring 
(Mar.  through  May),  Summer  (June  through  Aug.),  Fall  (Sept.  through 
Nov.),  and  Winter  (Dec.  through  Feb.). 

Table  3.    Seasonal  distribution  of  major  parasites  of  Mus  musculus. 


Spring 

Summer 

Fall 

Winter 

% 

Avg.#/ 

% 

Avg.  #/ 

% 

Avg.#/ 

%      Avg.#/ 

Parasites 

Infest. 

Mouse 

Infest. 

Mouse 

Infest. 

Mouse 

Infest.    Mouse 

Internal  Parasites 

Number  examined 

(79) 

(88) 

(194) 

(142) 

All    internal    parasites 

40.5 

19.3 

18.0 

19.0 

Heligmosomoides 

34.2 

4.48 

5.7 

0.89 

5.2 

0.24 

9.9       1.23 

Cestodes 

1.3 

0.06 

11.4 

0.65 

12. !) 

0.57 

4.2       0.10 

Syphacia 

2.5 

0.51 

4.5 

0.35 

0 

0 

2.1        0.18 

Protospirura 

5.1 

0.09 

2.2 

0.02 

1.5 

0.02 

1.4       0.02 

External  Parasites 

Number  examined 

(77) 

(85) 

(168) 

(140) 

All  external  parasites 

19.3 

34.1 

42.3 

12.9 

My  obi  a 

3.8 

0.43 

8.2 

0.46 

13.1 

0.41 

2.9       0.15 

Radfordia 

2.6 

0.17 

10.6 

0.27 

12.5 

0.1S 

1.4       0.01 

Myocoptes 

0 

0 

2.4 

0.02 

7.7 

0.32 

0           0 

Omithonyssus 

1.3 

0.04 

11.8 

0.24 

3.0 

0.07 

1.4       0.02 

A.    fahrenholzi 

1.3 

0.01 

1.2 

0.01 

5.4 

0.14 

2.1       0.02 

The  greatest  percentage  of  housemice  harbored  internal  parasites 
during  the  spring,  at  40.5%,  while  just  under  20%  of  those  taken  were 
parasitized  during  the  other  3  seasons.  This  difference  was  significant 
(Chi-square  =  14.45,  1  df).  The  parasite  load  is  heaviest  during  the 
spring  because  the  principle  internal  parasite,  Heligmosomoides 
polygyrus,  is  primarily  a  spring  parasite,  reaching  by  far  its  greatest 
percentage  infection  of  mice,  and  its  greatest  average  number  of  worms 
per  mouse  at  that  time.  This  nematode  had  its  second  greatest  occurrence 
during  the  winter.  Cestodes,  as  a  group  the  second  most  important  of 
the  internal  parasites,  were  least  common  during  the  spring  and  winter, 
and  most  common  during  the  summer  and  fall.  For  Syphacia,  the  data 
are  scanty,  but  this  form  would  appear  to  be  a  spring  and  summer 
form.  It  was  not  taken  at  all  in  the  fall  sample  of  mice,  even  though 
this  was  the  largest  sample,  at  194.  Protospirura  seemed  to  be  most 
abundant  in  the  spring,  but  data  are  too  few  concerning  this  species  to 
be  reliable. 

External  parasites,  as  a  group,  were  most  abundant  in  the  fall  and 
summer,  and  least  abundant  in  the  winter  (Chi-square  =  29.47,  3  df). 
Myobia,  musculi,  Radfordia  ajfinis  and  Myocoptes  mus&ulinus  were  fall 
and  winter  mites,  with  Myocoptes  occurring  only  during  that  season. 
Omithonyssus  bacoti  was  taken  at  its  greatest  rate  in  the  summer,  and 
Androlaelaps  fahrenholzi  was  taken  at  its  greatest  rate  in  the  fall.  With 
the  exception  of  A.  fahrenholzi,  all  species  occurred  at  their  lowest 
abundance  in  the  winter. 


Zoology 


447 


Parasites  of  Mus  as  Associated  with  Habitat 

There  were  enough  data  concerning:  Mus  parasites  in  eight  habitats 
to  make  a  meaningful  presentation  (Table  4).  Late  stage  winter  wheat 
(over  6  inches  high)  was  the  one  habitat  available  only  during  the 
spring.  As  seen  previously,  Heligmosomoides  was  primarily  a  spring 
parasite,  and  as  one  might  expect,  winter  wheat  was  the  habitat  in 
which  the  greatest  incidence  and  abundance  of  this  nematode  occurred. 
The  second  greatest  abundance  occurred  in  cut  corn,  primarily  a  winter 
habitat,  although  some  cut  corn  areas  were  available  in  the  fall,  and 
some  were  still  present  in  the  spring.  The  major  habitats  for  cestodes 
were  corn  and  cut  wheat.  Syphacia  was  most  abundant  in  the  winter 
wheat  over  6  inches,  and  in  winter  wheat  which  had  been  cut.  This  latter 
habitat  was  available  in  late  spring  and  early  summer. 


Table   4. 


Relationship   of   Mus   parasites    to   habitat.    (Numbers    in   parentheses    are    the 
numbers  of  plots  in  the  habitats.) 


Internal  Parasites 

Helimosomoides 

Cestodes 

Syphacia 
Avg.#/ 

Prot 

ospirura 

Avg.#/ 

Avg.#/ 

Avg.#/ 

Habitat 

% 

Mouse 

% 

Mouse 

% 

Mouse 

% 

Mouse 

Weedy    field    (58) 

3.4 

0.05 

5.2 

0.17 

0 

0 

Grassy   field    (69) 

5.8 

0.72 

4.3 

0.35 

0 

2.9 

0.06 

Soybeans    (25) 

12.0 

0.12 

4.0 

0.04 

4.0 

0.04 

4.0 

0.08 

Winter  wheat 

over    6"    (37) 

29.7 

5.22 

5.4 

0.27 

5.4 

1.08 

5.4 

0.08 

Corn     (118) 

3.8 

0.18 

15.4 

0.65 

0 

2.3 

0.02 

Sorghum    (31) 

0.7 

0.77 

0 

3.2 

0.03 

0 

Wheat,    cut    (79) 

10.1 

0.71 

12.7 

0.65 

3.8 

0.62 

2.5 

0.03 

Corn,  cut   (69) 

27.5 

4.35 

4.3 

0.07 

2.0 

0.07 

1.4 

0.01 

External  Parasites 

Myobia 

Radfordia 

A.  fahrenholzi 

Avg.#/ 

Myocoptes 

Avg.#/ 

0. 

bacoti 

Avg.#/ 

Avg.#/ 

Avg.#/ 

Habitat 

% 

Mouse 

% 

Mouse 

% 

Mouse 

% 

Mouse 

% 

Mouse 

Weedy   field    (54) 

5.6 

0.37 

0 

3.7 

0.04 

0 

3.7 

0.07 

Grassy   field    (65) 

4.6 

0.05 

7.7 

0.12 

6.1 

0.06 

1.5 

0.05 

t.5 

0.02 

Soybeans    (23) 

4.3 

0.04 

17.4 

0.39 

0 

4.3 

0.04 

0 

Winter  wheat 

6"    (37) 

8.1 

0.89 

2.7 

0.03 

0 

0 

0 

Corn    (118) 

14.4 

0.64 

16.1 

0.26 

5. 1 

0.17 

10.2 

0.42 

2.5 

0.08 

Sorghum  (25) 

8.0 

0.12 

0 

0 

0 

20.0 

0.24 

Wheat,    cut    (78) 

7.7 

0.28 

3.8 

0.10 

1.3 

0.01 

1.3 

0.04 

7.7 

0.21 

Corn,   cut    (67) 

1.5 

0.04 

3.0 

0.19 

1.5 

0.02 

o 

1.5 

0.03 

Myobia  was  most  abundant  in  the  winter  wheat  and  in  the  corn 
while  Radfordia  was  taken  at  its  greatest  rates  in  soybeans  and  corn. 
Myocoptes  was  quite  definitely  a  form  of  the  cornfields  while 
Ornithonyssus  bacoti  was  found  at  its  greatest  rate  in  sorghum. 


448  Indiana  Academy  of  Science 

Discussion 

Season,  habitat  and  age  of  the  host  all  seemed  to  influence  the  para- 
site population  of  the  housemouse,  Mus  muscuhis,  while  the  fourth 
factor  under  consideration,  sex  of  the  animal,  seemed  to  have  little  or  no 
effect.  Some  factors  appeared  interrelated  in  such  a  way  that  it  was 
difficult  to  determine  which  of  two  factors  was  affecting-  the  mice.  For 
example,  the  nematode  parasite,  Heligmosomoides  polygyrus,  was  most 
abundant  in  the  spring  in  winter  wheat  fields  in  which  the  wheat  was 
at  least  6  inches  high.  Hence,  one  can  conclude  that  the  best  situation 
in  which  to  look  for  this  species  is  in  winter  wheat  fields  during  the 
spring,  but  I  was  unable  to  evaluate  the  relative  effects  of  the 
particular  season   as   opposed  to  those  of  the  habitat. 

The  relationship  of  internal  and  external  parasitism  was  assessed 
in  an  attempt  to  determine  whether  animals  became  parasitized  because 
of  their  generally  poor  physical  condition,  or  if  the  animals  simply 
happened  to  be  in  the  right  place  at  the  right  time  to  become  para- 
sitized. If  animals  were  infected  because  of  a  general  overall  poor 
physical  condition,  then  the  same  animals  that  were  parasitized  inter- 
nally should  also  tend  to  be  parasitized  externally.  On  the  other  hand, 
if  parasitism  was  strictly  a  chance  happening,  then  one  should  be  able 
to  compute  the  approximate  number  of  mice  expected  to  have  both 
internal  and  external  parasites  by  multiplying  the  percentage  of  mice 
with  external  parasites  times  the  percentage  with  internal  parasites. 
For  this  calculation,  only  the  470  mice  examined  for  both  internal  and 
external  parasites  were  used.  Of  these,  130,  or  0.277  were  found  to  have 
external  parasites,  and  102,  or  0.217  were  found  to  harbor  internal  para- 
sites. One  would  expect,  by  chance,  that  0.277  X  0.217  =  0.060  of  the 
470,  or  28.20  mice  would  have  both  internal  and  external  parasites.  The 
actual  number  of  mice  with  both  internal  and  external  parasites  was  30, 
hence  we  can  conclude  that  the  relationship  between  internal  and 
external  parasites  of  the  housemouse  in  Vigo  County  is  strictly  a 
chance   one. 


Literature  Cited 

1.  Rupes,  V.,  and  J.  O.  Whitaker,  Jr.  1968.  Mites  of  the  subfamily  Labidophorinae 
(Acaridae,  Acarina)  in  North  America.  Acarologia  10  :493-499. 

2.  Whitaker,  J.  O.,  JR.  1967.  Habitat  and  reproduction  of  some  of  the  small  mammals 
of  Vigo  County.  Indiana,  with  a  list  of  mammals  known  to  occur  there.  Occas.  Papers 
Adams  Ctr.  Ecol.  Studies.  16  :l-24. 

3.  Whitaker,  J.  O.,  Jr.,  and  K  W.  Corthum,  Jr.  1967.  Fleas  of  Vigo  County,  Indiana. 
Proc.  Indiana  Acad.  Sci.  76  :431-440. 

4.  Whitaker,  J.  O.,  Jr.,  and  N.  Wilson.  1968.  Mites  of  small  mammals  of  Vigo  County, 
Indiana.  Amer.  Midland  Natur.  80  :537-542. 

5.  Wilson,  N.  1961.  The  ectoparasites  (Ixodides,  Anoplura  and  Siphonaptera)  of 
Indiana  mammals.  Unpublished  thesis.  Purdue  University.  527  p. 


Molt  in  Two  Populations  of  the  House  Mouse,  Mus  musculus 

Ronald  E.  Brechner  and  Ralph  D.  Kirkpatrick, 
Ball  State  University 

Abstract 

This  study  correlated  molt  in  two  population  samples  of  the  house  mouse,  Mus 
musculus,  with  sex,  age,  geographic  origin  and  time  of  year.  Sample  A  consisted  of 
273  house  mice  trapped  in  1964  and  1965  on  Sand  Island,  Johnston  Atoll,  central 
Pacific  Ocean.  The  272  mice  of  Sample  B  were  caught  over  a  6-month  period  in  Delaware 
County,  Indiana,  in  1968  and  1969. 

Incidence  of  pigmented  areas  (molt)  on  the  fleshside  of  the  pelts  of  both  samples 
was  recorded  and  statistically  analyzed  to  find  patterns  and  percent  molt.  There  was 
no  positive  correlation  between  incidence  of  molting  and  origin  and  age  of  sample  mice, 
nor  between   incidence  of  molting  and  sex  or  month   of  collection. 

Introduction 

The  present  study  correlated  molt  (as  indicated  by  pigmentation  of 
the  fleshside  of  pelts)  in  two  feral  populations  of  the  house  mouse,  Mus 
musculus,  with  sex,  age,  geographic  location  and  time  of  year.  The  term 
"feral"  as  used  here  implies  mice  which  are  either  living  out-of-doors  or 
in  unheated  buildings. 

Sample  A  represented  an  insular  population  from  the  central 
Pacific  Ocean.  Sample  B  was  taken  from  Delaware  County,  Indiana. 
Percentage  molt  and  areas  of  molt  were  recorded  and  each  specimen 
was  assigned  to  an  arbitrary  age  class. 

Related  Literature 

Osgood  (11)  noted  three  different  phases  of  pelage  color  in 
Peromyscus  which  corresponded  to  age — juvenile,  adolescent,  and  adult. 
Allen  (1)  described  molt  in  this  same  genus  as  generally  beginning  in 
the  feet  and  around  the  nose  and  extending  dorsally  and  also  proceed- 
ing anteriorly  from  the  base  of  the  tail.  Collins  (3)  agreed  that  this  is 
the  general  pattern  of  spring  molt  in  Peromyscus  maniculatus,  but  that 
it  is  subject  to  much  irregularity.  He  found  that  the  juvenile  pelage  was 
complete  at  4  to  5  weeks  and  transition  from  juvenile  to  post  juvenile 
pelage  occurred  between  the  ages  of  6  and  8  weeks.  Molt  on  the  ventral 
surface  was  usually  completed  before  the  dorsal  surface  and  usually 
proceeded  from  anterior  to  posterior.  Collins  (3)  checked  for  molt  by 
parting  the  hair  and  checking  for  the  presence  of  new  hair.  But  as 
pointed  out  by  Golley  et  al.  (5),  the  molt  is  more  readily  determined 
from  the  underside  of  the  skin  and  may  be  well  underway  before  it  is 
detected  on  the  surface. 

Gollschang  (6)  stated  that  some  Peromyscus  leucopus,  both  captive 
and  wild,  molt  over  an  indefinite  period  of  time,  and  that  body  weight 
had  no  direct  relationship  to  onset  of  pelage  change. 

Most  mammals  have  similar  pigmented  areas  as  illustrated  by 
Kopenen   (9)   who  examined  the  sequence  of  pelages  of  the  Norwegian 

449 


450  INDIANA   ACADEMY    OF    SCIENCE 

lemming,  Lemmus  lemmus,  by  using"  the  pigment  patterns  of  the 
fleshside  of  the  skin  and  Skoczen  (12),  who  studied  seasonal  changes  of 
pelage  of  the  mole,  Talpa  europaea,  as  measured  by  the  planimetric 
measuring  of  pigmented  areas  on  the  underside  of  the  skin. 

Golley  et  al.  (5)  studied  skins  of  Peromyscus  polionotus  by  pinning 
the  skins  out  to  dry  on  cardboard  for  6  months  with  the  furside  down 
and  then  using  a  planimeter  to  measure  total  area  exhibiting  molt.  He 
found  that  molt  was  influenced  by  increasing  age;  however,  he  also  noted 
it  may  be  influenced  by  sex,  body  size,  trauma,  and  other  environ- 
mental factors.  The  percentage  of  the  pelt  involved  in  the  molt  process 
progressively  declined  with  age.  He  found  80  to  90%  of  total  pelage 
involved  in  molt  in  juveniles,  but  it  never  exceeded  45%  at  later  stages 
of  development.  There  was  a  35%  maximum  in  post-juvenile  molt. 
Golley  et  al.  (5)  also  noted  areas  of  gray  pigmentation  preceding  or 
following  the  areas  of  most  active  molt. 

The  pigmentation  of  flat-skin  mounts  of  33  specimens  of  known  age 
Microtus  calif ornicus  used  in  an  experiment  by  Ecke  and  Kinney  (4) 
revealed  a  close  age-molt  correlation  from  17  to  60  days  of  age.  Animals 
could  be  aged  by  degree  of  molt  to  within  4  days  of  their  actual  age. 
The  method  was  88%  accurate  with  laboratory-reared  mice.  Evidence 
from  skins  of  old  animals  indicated  that  all  adult  molts  are  irregular 
after  the  molting  of  post-juvenile  pelage.  These  occurred  as  mottled 
patterns  of  dark  and  light  areas  on  the  underside  of  the  skin  with  no 
apparent  consistency  of  design.  No  significant  molt  variation  was 
found  between  the  two  sexes. 

Methods 

Sample  A  was  collected  on  Sand  Island,  Johnston  Atoll,  central 
Pacific  Ocean,  by  the  junior  author  while  working  with  the  Pacific 
Ocean  Biological  Survey  Program  in  1964  and  1965.  The  atoll  covers  less 
than  800  acres  and  this  insular  population  has  developed  since  1923  when 
the  U.S.S.  Tanager  Expedition  biologists  found  no  mammals  on  the 
island   (8). 

Sample  B  was  taken  from  Delaware  County,  Indiana.  A  minimum 
of  30  mice  per  month  was  collected  during  a  6-month  period  from 
September  27,  1968,  through  March  22,  1969.  The  mice  came  from 
2  county  locations:  1)  Delaware  County  Fairgrounds,  Muncie,  Indiana; 
2)  Earl  Southworth's  farm — 1  mile  west  of  Tillotson  Avenue  on  Bethel 
Pike,  Muncie,  Indiana. 

All  but  6  of  the  272  mice  in  Sample  B  were  assigned  to  arbitrary 
age  classes  based  on  molar  wear  after  Lidicker  (10).  The  remaining  6 
were  not  aged  because  the  skulls  were  destroyed  during  trapping. 
Data  collected  from  the  266  usable  specimens  were  compared  with  data 
from  the  273  mice  in  Sample  A. 

Mice  were  collected  using  live-traps  and  snap-traps.  The  pelt  was 
removed  from  the  carcass,  tagged,  and  pinned  with  the  furside  down  to 
dry.  Facia  and  excess  fat  were  removed  as  suggested  by  Clark  (2), 
to  prevent  their  masking  of  molt  pattern. 


Zoology 


151 


Percentage  of  molt  was  estimated.  A  grid  system  composed  of 
Vs-inch  squares  was  imposed  on  a  plastic  transparency.  This  was  then 
placed  over  the  skinside  of  each  pelt,  and  the  percentage  of  molt  was 
calculated  by  comparing  the  total  number  of  squares  covered  by  the 
pelt  to  the  number  of  squares  covered  by  pigmented  areas. 

We  decided  that  a  modified  version  of  the  pelt  pattern  used  by 
Hendricks  (7)  would  best  demonstrate  the  possible  patterns  (Fig.  1). 
A  map  with  14  areas  was  prepared  for  each  pelt.  To  designate  an  area 
as  molting,  a  minimum  of  Vs  square  inch  of  that  area  must  be  molting 
as  demonstrated  by  the  presence  of  pigmentation.  This  indicator  was 
arbitrarily  chosen. 

The  areas  of  molt  on  each  pelt  were  marked  on  the  map  of  the 
pelt,   along  with   other   pertinent   information   available   for  that   speci- 


Figure  1.    Pelt  map  of  house  mouse  illustrating  areas  of  molt. 


452 


Indiana  Academy  of  Science 


men.   A  combination  of  Chi-square   and   co-factor   analysis   was   run  to 
find  the  most  frequently  occurring  patterns. 

After  these  patterns  were  determined,  each  pelt  map  was  again 
examined  and  classified  as  representing  one  of  the  main  patterns,  as 
irregular,  or  as  displaying  no  molt. 

A  pelt  was  recorded  as  having  a  certain  molt  pattern  if  it  dis- 
played molt  in  50%  or  more  of  the  areas  comprising  that  pattern.  It 
was  possible  for  more  than  one  pattern  to  be  represented  on  the  same 
pelt.  This  information  was  analyzed  for  correlation  of  molt  pattern  with 
age,  sex,  geographic  origin,  and  time  of  year. 

Results  and  Discussion 

Population  Samples  A  and  B  were  analyzed  separately  for  repre- 
sentative molt  patterns.  Five  patterns  were  found  in  Sample  A  and  four 
patterns  in  Sample  B  (Fig.  2).  The  molt  patterns  were  lettered  A 
through  I;  J  was  used  to  indicate  an  irregular  unclassified  molt  pattern; 
and  K  represented  no  molt. 


SAMPLE  A 

3A5A1! 


2,  12,13 


SAMPLE 


Figure  2.    Predominant  molt  pattern  in  samples  from  two  house  mouse  populations. 


Each  pelt  was  again  checked  to  determine  which  pattern  or  patterns 
it  represented.  It  was  possible  for  several  patterns  to  be  represented  on 
one  pelt.  When  this  occurred,  the  composite  of  patterns  was  considered 
as  one  pattern.  For  example,  many  of  the  pelts  represented  one  clear-cut 
pattern,  such  as  I  or  H;  but,  some  of  the  pelts  had  patterns  F,  G,  H,  I 
represented  on  the  same  pelt.  Such  pelts  were  considered  as  representing 
one  pattern,  the  pattern  of  FGHI. 

Males  and  females  within  the  same  sample  were  considered  jointly, 
since  they  were  characterized  by  the  same  distribution  and  frequency 
of  molt  patterns.  The  molt  patterns  were  randomly  distributed  through- 
out  the   age   classes   and   the   months   of   collection,   and    did   not   show 


Zoology  453 

positive  correlation  with  either  of  these.  The  samples  were  represented 
by  different  patterns,  with  the  exception  of  C  and  G,  which  were  identical 
in  both  samples. 

Ecke  and  Kinney  (4)  were  able  to  age  Microtus  calif ornicus  up  to 
60  days  of  age  by  using  a  combination  of  weight  and  molt  pattern. 
They  found  that  all  older  mice  had  irregular  molt.  Our  findings  indi- 
cated that  in  the  house  mouse,  regular  patterns  indicative  of  the 
populations  are  present.  These  are  randomly  distributed  throughout  the 
age  classes  and  months  in  both  sexes. 

There  was  no  significant  difference  between  the  percentage  molt 
of  the  sexes  (P  =  0.05).  In  Age  Classes  3  and  7,  the  females  nearly 
doubled  the  percentage  of  molt  displayed  by  the  males.  However,  most 

Table  1.    Mean  percent  molt  by  sex  and  age  class  in  samples  from  two 
house  mouse  populations. 


Age  Class                3 

4 

5 

6 

7 

Mean  % 

Sand  Island,  Johnston  Atoll 

Male                        15.6 

20.0 

16.0 

13.2 

10.0 

14.9 

Female                     33.6 

24.8 

20.0 

21.5 

19.1 

23.8 

Combined                24.6 

22.4 

18.0 

17.3 

14.5 

19.3 

Delaware  County,  Indiana 

Male                        10.1 

11.1 

4.2 

5.2 

1.7 

6.4 

Female                     20.5 

10.0 

3.6 

5.9 

8.1 

9.6 

Combined                15.3 

10.5 

3.9 

5.5 

4.9 

8.0 

Table    2.     Percent   molt    by   month   for   male    and   female    house   mice 
collected  from  two  populations. 


Sand  Island,  Johnston  Atoll 
Male          Female          Sum 

Delaware  County, 

Indiana 

Male 

Female 

Sum 

Sept. 

19 

30 

49 

3 

2 

5 

Oct. 

30 

37 

67 

15 

11 

26 

Nov. 

14 

31 

45 

9 

9 

18 

Dec. 

21 

15 

36 

5 

3 

8 

Jan. 

14 

19 

33 

7 

15 

22 

Feb. 

14 

9 

23 

6 

5 

11 

Mar. 

— 

— 

— 

5 

20 

25 

Apr. 

10 

12 

22 

— 

— 

— 

May 

25 

10 

35 

— 

— 

- — 

June 

16 

25 

41 

— 

— 

— 

July 

18 

29 

47 

— 

— 

— 

Aug. 

10 

19 

29 

— 

— 

— 

454  Indiana  Academy  of  Science 

of  the  sample  representatives  are  in  Age  Classes  4,  5,  and  6,  with  only 
a  small  number  in  Age  Classes  3  and  7  (Table  1). 

The  amount  of  molt  was  inversely  proportional  to  age.  The  greatest 
percentage  molt  occurred  in  Age  Class  3  (the  youngest  mice),  and  the 
least  percentage  of  molt  in  Age  Class  7  (the  oldest  mice). 

Sample  A  showed  the  greatest  percentage  of  molt  during  June, 
July,  September,  October  and  November.  The  least  molt  was  evidenced 
in  February  and  April  (Table  2).  There  were  no  March  specimens  used 
in  this  study. 

Month  of  collection  did  not  correlate  significantly  with  percentage 
of  molt  in  Sample  B.  September  and  December  had  the  smallest  percent 
molt;  whereas,  October  and  March  had  the  greatest  percent  (Table  2). 
However,  this  sampling  period  was  but  6  months  in  length. 

Geographic  origin  of  sample  mice  was  related  to  mean  percent  molt 
and  was  significant  at  the  0.05  level.  Table  1  reveals  that  in  each 
age  class,  percentage  of  molt  in  Sample  A  was  at  least  IV2  times  that 
of  Sample  B. 

Acknowledgments 

Thanks  are  extended  to  Mr.  Tom  Harris  and  Dr.  Terry  Schurr, 
Ball  State  Office  of  Research;  William  B.  Wilson,  Ronald  D.  Toombs, 
and  Larry  G.  Scherich  for  collection  assistance;  and  Phillip  Lehner  and 
Dennis  Stadel,  Pacific  Ocean  Biological  Survey  Program. 

Literature  Cited 

1.  Allen,   Glover   M.    1914.   Pattern   development   in   mammals   and   birds.   Amer.    Natur. 
48:385-412,  467-484,  550-566. 

2.  Clark,   James   L.    1937.   The  preservation  of  mammal  skins  in  the  field.   J.   Mammal. 
18:89-92. 

3.  Collins,    H.    H.    1918.    Studies    of    normal    molt   and   of    artificially   induced    regenera- 
tion of  pelage  in  Peromyscus.  J.  Exp.  Zool.  27  :73-99. 

4.  Ecke,    Dean    H.,    and    Alva    R.    Kinney.    1956.    Aging    meadow    mice,    Microtus    cali- 
fomicus,    by   observation   of    molt   progression.    J.    Mammal.    37:249-254. 

5.  Golley,    Frank   B.,    Eric   L.   Morgan,   and   James   L.    Carmon.    1965.    Progression    of 
molt   in    Peromyscus   polionotus.    J.    Mammal.    47:145-149. 

6.  Gollschang,    Jack    L.    1956.    Juvenile   molt   in   Peromyscus    leucopus    novcboraccnsis. 
J.  Mammal.  37:516-520. 

7.  Hendricks,   Donovan   E.   1967.   The  ectoparasites  and  other  arthropods  associates   of 
the  13-Lined  Ground   Squirrel.   Purdue  University  Research  Bulletin  No.   817.   14   p. 

8.  Kirkpatrick,    Ralph   D.    1966.   Mammals   of   Johnston   Atoll.    J.    Mammal.    47:728-729. 

9.  KOPENEN,     TERTTU.     1965.     The     sequence     of     pelages     in     the     Norwegian     lemming, 
Lemmus    lemmus    L.    Arch.    Soc.    Zool.    Bot.    Fennial    Venamo.    18 :200-278. 

10.  Lidicker,  William  Z.,  Jr.  1965.  Ecological  observations  on  a  feral  house  mouse  popu- 
lation declining  to  extinction.  Ecol.  Monogr.  36  :27-50. 

11.  Osgood,   Wilfred   H.   1909.   Revision   of  the  mice  of  the   American  genus   Peromyscus. 
Washington:   Bur.  of  Biol.  Surv.,  N.  Amer.  Fauna.  No.  28:1-285. 

12.  Skocqen,     Stanislaw.     1966.     Seasonal    changes    of    the    pelage    in    the    mole,     Talpa 
curopaea  L.,  1758.  Acta  Theriologica.  11:537-549. 


Some  Effects  of  Aminoglutethimide  on  Water  and  Electrolyte 
Metabolism  in  the  Female  Rat 

R.  E.  Zimmerman  and  W.  J.  Eversole,  Indiana  State  University 

Abstract 

Aminoglutethimide  phosphate  (AG)  was  injected  subcutaneously  at  a  dosage  level  of 
100  mg/kg  body  wt/day  into  Charles  River  strain  female  rats  of  approximately  160  g. 
Control  rats  were  treated  under  the  same  conditions,  but  were  injected  with  distilled 
water.  Comparisons  were  made  at  1  and  3  days. 

Urine  volumes  were  compared  between  water-fasted  controls  and  treated  animals. 
Water  consumption  was  compared  between  nephrectomized  controls  and  treated  animals. 
Water  consumption  and  urine  output  were  compared  between  intact  controls  and  treated 
animals  allowed  to  drink  ad  libitum. 

Urine,  tissue,  and  plasma  sodium  and  potassium  levels  were  compared  in  treated  and 
control  animals,  both  intact  and  adrenalectomized. 

Chronic  administration  of  AG  to  intact  rats  induced  a  diabetes-insipidus-like  state 
and  many  animals  doubled  their  24-hour  water  exchange.  Aminoglutethimide  phosphate 
elicited  polydipsia  in  the  absence  of  the  adrenals  or  the  kidneys,  suggesting  that  polyuria 
was  secondary  to  increased  water  intake  in  the  complete  absence  of  adrenal  cortical 
hormones. 

Sodium  and  potassium  concentrations  in  urine,  tissue  and  plasma  of  intact  animals 
treated  with  AG  resembled  those  in  adrenalectomized  non-treated  animals,  while  treat- 
ment in  adrenalectomized  animals  appeared  to  produce  an  additive  effect.  Such  results 
may  be  interpreted  as  indicating  that  the  effects  of  AG  are  mediated  only  in  part  by  the 
adrenal  cortex. 

Introduction 

Aminoglutethimide  phosphate  (AG)  [a-(p-aminophenyl)a-ethylglu- 
tarimide]  has  been  used  clinically,  and  appears  to  have  value  in  the 
treatment  of  generalized  seizure  when  administered  in  dosages  of  250- 
500  mg  three  times  a  day  (1).  Kahnt  and  Neher  (4)  in  a  continuing 
study  of  the  effect  of  large  numbers  of  drugs  on  beef  adrenal  homog- 
enates  in  vitro  found  that  aminoglutethimide  (AG),  among  others, 
inhibited  steroid  synthesis.  It  is  now  generally  agreed  that  the  primary 
activity  of  AG  is  to  inhibit  the  synthesis  of  corticoids  at  some  step(s) 
between  the  conversion  of  cholesterol  to  pregnenolone,  the  latter  being 
a  precursor  of  all  steroid  hormones  (2,  3).  It  has  been  shown,  in  rats, 
that  AG  markedly  decreased  the  output  of  corticosterone  (3). 

The  purpose  of  this  investigation  was  to  determine  the  effects  of 
aminoglutethimide  on  water  exchange  and  sodium  and  potassium  levels 
in  body  fluids  and  tissues  of  intact  and  adrenalectomized  animals.  We 
expected  this  study  to  answer  the  question  of  whether  AG  induces 
physiological  changes  characteristic  of  adrenal  dysfunction  and  whether 
or  not  its  effects  on  water  and  electrolyte  metabolism  are  brought  about 
by  direct  action  or  by  mediation  through  changes  induced  in  adrenal 
cortical   function. 

Materials  and  Methods 

Three  groups  of  12  rats  each  were  used  for  the  first  phase  of  the 
experiment.  Half  of  the  animals  were  given  daily  subcutaneous  injections 

455 


456  Indiana  Academy  of  Science 

of  AG  (100  mg/kg  body  wt).  Controls  were  treated  the  same,  but  were 
given  injections  of  distilled  water. 

The  first  group  was  fasted  for  24  hours  then  placed  in  metabolism 
cages  without  water  or  food.  Urine  volumes  were  collected  in  graduate 
cylinders  for  48  hours.  The  urine  output  of  controls  and  treated  animals 
was   compared. 

The  second  group  was  placed  in  metabolism  cages  and  allowed  to 
drink  ad  libitum.  The  amount  of  water  consumed,  the  amount  of  urine 
excreted,  and  the  amount  of  weight  gained  were  measured  daily  for 
72  hours. 

The  ureters  were  clamped  off  in  the  third  group.  They  were  then 
put  in  cages  and  allowed  to  drink  ad  libitum.  Water  consumption  was 
measured  at  24  hours. 

In  the  second  phase  of  the  work,  1-  and  3-day  electrolyte  levels  were 
checked  both  in  intact  and  adrenalectomized  controls,  and  in  treated 
animals  receiving  daily  subcutaneous  injections  of  AG  (100  mg/kg  body 
wt).  Serum,  urine,  and  tissue  electrolytes  were  done  by  flame  photometry 
using  procedures  outlined  in  the  Coleman  Flame  Photometry  Manual. 
Tissue  electrolytes  were  done  using  the  procedure  for  urinary  electro- 
lytes, and  employing  1  g  samples  of  gastrocnemius  muscle  homogenized 
in  10  ml  of  distilled  water. 

The  data  were  statistically  evaluated  using  the  Student's  "t"  test. 
A  "P"  value  of  less  than  0.01  was  considered  significant. 

Results 

Aminoglutethimide  had  no  effect  on  the  24-  and  48-hour  urine 
volume   of  water-fasted   rats.    In   animals    drinking   ad   libitum,    it    did, 

Table  1.     Effects  of  aminoglutethimide  on  urine  volume  in  water  fasted 

rats,  and   those   drinking   ad  libitum    (numbers   in   parenthesis   indicate 

number  of  cases  and  results  are  expressed  as  mean  ±  SE). 


Treatment 
lOOrng/ 

Milliliters 

Urine 

Mean 

Body 

Weight  in 

Water  Fasted 

Drinking 
ad  libitum 

Grams 

kg/ day 

24  hr 

48  hr 

24  hr 

48  hr 

170 

AG** 

4.0±.4 

6.1±.7 

25±6 

41±11 

(12) 

(12) 

(12) 

(12) 

P>.05 

P>.05 

P<.01 

P<.01 

165 

none 

4.0±.6 

5.6±.9 

11±1 

20+2 

(12) 

(12) 

(12) 

(12) 

'Aminoglutethimide. 


Zoology 


457 


Table  2.     The   effects   of  amino g lute thimide   on   water  consumption   in 
intact  and  nephrectomized  animals  drinking  ad  libitum. 


Mean 

Treatment 
lOOmg/ 
kg7day 

Milliliters  Water  Consumed 

Body 
Weight 

Intact 

Nephrectomized 

in  Grams 

24  hr 

48  hr 

24  hr 

150 
150 

AG 
none 

43±4 

(12) 
P<.001 
20±2 

(12) 

73  ±20 

(12) 
P<.001 

37±5 
(12) 

30±2.4 

(12) 

P<.01 

20±1.5 

(12) 

however,  cause  significant  increases  in  urine  volume  at  both  24  and  48 
hours    (Table   1). 

Nephrectomized  and  intact  animals  treated  with  AG  exhibited  sig- 
nificant increases  in  24-hour  water  intake  compared  to  controls  (Table 
2). 

Aminoglutethimide  significantly  reduced  the  concentration  of  plasma 
sodium,  in  intact  animals,  at  1  day.  Intact  or  adrenalectomized  controls 
showed  no  changes  in  plasma  sodium,  whereas  the  intact  and  adrenalec- 
tomized treated  animals  exhibited  about  the  same  sodium  decrease  at 
1  day   (Table  3). 

Intact  animals  treated  for  3  days  showed  a  decrease  in  sodium 
equal  to  that  of  the  intact  animals  treated  for  1  day,  and  also  equal  to 
the    3-day    adrenalectomized    non-treated    rats.     The     adrenalectomized 


Table  3.     The  effects  of  aminoglutethimide  on  plasma  sodium  and  po- 
tassium levels  in  the  intact  and  adrenalectomized  animals. 


Mean 

Treatment 

Milliequivalents/ Liter 

Body 

Intact 

Adrenalectomized 

Weight 

lOOmg/ 

in  Grams 

kg/day 

Na 

K 

Na 

K 

175 

AG 

128±.8 

8.4±.2 

126±1 

8.0  ±.5 

(lday) 

(12) 

(12) 

(12) 

(12) 

P<.01 

P<.01 

P<.01 

P<.01 

177 

none 

143±.6 

6.9  ±.2 

141±.9 

4.5±.3 

(lday) 

(12) 

(12) 

(12) 

165 

AG 

(3  days) 

128±1 

8.4±.2 

115±6 

9.5±.7 

P<.01 

P<.01 

P<.01 

P<.01 

165 

none 

144±.7 

6.5±.3 

126±.8 

8.4±.4 

(3  days) 

(12) 

(12) 

(12) 

(12) 

458 


Indiana  Academy  of  Science 


treated  animals,  however,  showed  an  even  greater  decrease  at  3   days 
(Table  3). 

The  plasma  potassium  of  intact  and  adrenalectomized  animals 
treated  with  AG  was  elevated  at  1  day,  and  the  potassium  level  of  the 
adrenalectomized  non-treated  animals  was  somewhat  lower  than  in  in- 
tact controls,  but  not  significantly  so.  The  potassium  levels  of  the  intact 
animals  treated  for  3  days  was  the  same  as  in  the  1-day  intact  treated 
animals.  The  3-day  adrenalectomized  non-treated  animals  showed  an 
elevation  of  plasma  potassium.  The  plasma  potassium  of  adrenalectom- 
ized treated  animals  was  elevated   (Table  3). 

Total  urinary  sodium  was  markedly  reduced  in  animals  treated  with 
AG  for  1  day.  One-day  adrenalectomized  controls  also  showed  a  reduc- 
tion in  urine  sodium.  At  3  days  the  values  for  the  intact  treated  animals 
were  slightly  elevated.  The  adrenalectomized  animals,  treated  and  con- 
trols, showed  a  reduction  in  sodium  (Table  4). 


Table  4.     The  effects  of  amino  glut  ethimide  on  urine  output  of  sodium 
and  potassium  in  the  intact  and  adrenalectomized  animals. 


Mean 

Milliequivalents/day 

Body 

Treatment 

Intact 

Adrenalectomized 

Weight 

lOOmg/ 

in  Grams 

kg/day 

Na 

K 

Na 

K 

177 

AG 

.17±.02 

.2±.04 

.17±.02 

.18±.02 

(lday) 

(12) 

(12) 

(12) 

(12) 

P<.01 

P<.02 

P<.02 

P<.03 

175 

none 

.46±.02 

.14±.02 

.195±.01 

.19±.01 

(lday) 

(12) 

(12) 

(12) 

(12) 

165 

AG 

,46±.05 

.23±.03 

.15±.01 

.21±.01 

(3  days) 

(12) 

(12) 

(12) 

(12) 

P>.()5 

P>.05 

P<.01 

P<.02 

165 

none 

.47±.01 

.21±.02 

.1±.01 

.23±.01 

(3  days) 

(12) 

(12) 

(12) 

(12) 

Urinary  potassium  in  intact  animals  treated  for  1  day  with  AG 
was  elevated.  The  potassium  levels  in  adrenalectomized  treated  and 
control  animals  were  equal  to  that  in  the  intact  treated  animals  (Table  4). 

At  3  days  the  potassium  output  in  the  intact  treated  and  non- 
treated  animals  was  the  same.  The  potassium  level  in  the  3-day 
adrenalectomized  treated  was  elevated  (Table  4). 

The  sodium  concentration  in  muscle  tissue  of  intact  treated  animals 
was  elevated  at  1  day.  The  1-day  adrenalectomized  treated  animals 
showed  a  level  approximately  equal  to  that  in  the   1-day  intact  treated 


Zoology 


45!) 


Table  5.     The  effects  of  amino glutethimide  on  tissue  sodium  and  potas- 
sium in  the  intact  and  adrenalectomized  animals. 


Mean 

Treatment 

Milliequivalents/lOOg 

Body 

Intact 

Adrenalectomized 

Weight 

lOOmg/ 

in  Grams 

kg/ day 

Na 

K 

Na 

K 

175 

AG 

4.4±.5 

44  ±10 

5.0±.4 

,'}()  •  2 

(1  day) 

(12) 

(12) 

(12) 

(12) 

P<.01 

P<.01 

P<.02 

P<.05 

177 

none 

2.7±.2 

61±1 

3.9±.2 

36±10 

(1  day) 

(12) 

(12) 

(12) 

(12) 

165 

AG 

2.3±.4 

74±10 

4.9±.04 

21±1 

(3  days) 

(12) 

(12) 

(12) 

(12) 

P<.05 

P<.05 

P<.01 

P<.01 

165 

none 

2.4±.3 

73±50 

2.9±.3 

38±5 

(3  days) 

(12) 

(12) 

(12) 

(12) 

animals.  At  3  days  the  intact  non-treated,  intact  treated,  and  the  adren- 
alectomized non-treated  rats  exhibited  equal  tissue  sodium  levels.  The 
3-day  adrenalectomized  treated  animals  demonstrated  an  increase  in 
tissue  sodium  (Table  5).  Intact  or  adrenalectomized  animals  treated 
with  AG  for  1  day  showed  a  reduction  in  tissue  potassium  which,  at  3 
days,  remained  low  in  adrenalectomized  rats,  but  was  near  the  control 
value  in  intact  rats   (Table  5). 

Discussion 

The  data  show  that  aminoglutethimide  has  a  significant  effect  on 
water  intake.  Treated  rats  allowed  to  drink  ad  libitum,  consumed  sig- 
nificantly more  water,  excreted  more  urine,  and,  over  short  periods, 
retained  more  water  than  did  their  controls.  Treatment  induced  water 
retention  during  the  first  24-hour  period  after  injection,  but  subsequent 
injections  failed  to  exacerbate  this  condition.  Also,  when  treated  rats 
were  deprived  of  water  the  urine  volume  was  not  increased  over  the 
control  value,  thus  indicating  that  a  primary  effect  of  the  drug  was  on 
thirst  rather  than  on  excretory  mechanisms.  These  findings  are  inter- 
preted as  indicating  that  increase  in  thirst  was  caused  by  a  change  in 
internal  osmotic  pressure  which  was  then  corrected  for  by  drinking 
and  establishing  a  new  high  internal  fluid  level;  this  level  when  reached 
was  maintained  but  not  increased. 

Treated  animals  which  had  been  nephrectomized  showed  an  increase 
in  fluid  intake.  This  demonstrated  that  the  increase  in  urine  output 
found  in  treated  animals,  allowed  to  drink  ad  libitum,  was  a  product  of 
the  polydipsic  effect  and  not  a  cause  for  it. 

Aminoglutethimide  treatment  caused  an  increase  in  plasma  potas- 
sium and  a  concomitant  decrease  in  plasma  sodium.  The  adrenalectomized 
controls  did  not  show  a  reduction  in  plasma  sodium  at  1  day,  but  did 
show  this  reduction  at  3  days.  The  reason  for  the  difference  in  plasma 


460  Indiana  Academy  of  Science 

sodium  at  3  days  compared  to  1  day  after  adrenalectomy  is  probably  be- 
cause the  levels  of  adrenal  steroids  in  the  circulation  1  day  after  adrenal- 
ectomy had  not  been  sufficiently  depleted  to  permit  a  lowering  of  plasma 
sodium. 

In  the  intact  treated  animals,  regardless  of  the  length  of  treatment, 
there  was  a  drop  in  plasma  sodium  equal  to  the  levels  in  adrenalec- 
tomized treated  animals. 

The  plasma  potassium  levels  in  1-  or  3-day  treated,  intact  or 
adrenalectomized,  animals  were  inversely  related  to  the  sodium  levels. 
The  potassium  levels  in  the  adrenalectomized  controls  were  unchanged 
at  1  day.  The  potassium  level  in  the  intact  and  adrenalectomized  treated 
animals  at  1  day  was  increased. 

As  was  expected,  the  plasma  of  3-day  adrenalectomized  non-treated 
animals  was  low  in  sodium.  The  plasma  sodium  in  the  intact  animal 
treated  for  3  days  was  reduced  to  a  level  equal  to  that  in  the  adrenalec- 
tomized controls.  The  adrenalectomized  animals  treated  for  3  days 
showed  a  greater  reduction  m  plasma  sodium  than  did  any  of  the  other 
animals  regardless  of  operative  and  treatment  procedures.  This  is 
indicative  that  AG  has  direct  effects  on  blood  electrolyte  levels  inde- 
pendently of  any  action  mediated  via  the  adrenal  cortex. 

The  urine  sodium  output  was  depressed  in  intact  animals  treated 
for  1  day  and  in  1-day  adrenalectomized  non-treated  animals.  These 
results  could  be  interpreted  as  indicating  that  at  1  day  neither  the 
adrenalectomized  non-treated  nor  the  intact  treated  animals  had  com- 
plete adrenal  insufficiency.  At  3  days  the  adrenalectomized  rats  did  show 
the  expected  urinary  loss  of  sodium,  whereas  adrenalectomized  treated 
animals  at  either  1  or  3  days  showed  a  marked  decrease  in  sodium 
output.  This  is  further  evidence  of  an  effect  of  AG  which  is  independent 
of  adrenal  function. 

The  variations  and  errors  in  urine  potassium  determinations  were 
such  that  possible  correlation  with  adrenal  involvement  could  not  be 
made.  However,  the  drug  did  influence  potassium  excretion  in  that  it 
increased  it  in  the  intact  animals  at  1  day  and  in  the  adrenalectomized 
animals  at  1  day. 

The  tissue  sodium  levels  in  the  intact  treated  and  adrenalectomized 
treated  rats  were  increased  at  1  day.  Such  increases  in  tissue  sodium 
lend  credence  to  the  concept  that  rapid  osmotic  shifts  occur  in  the 
drug-induced  polydipsia.  The  action  of  aminoglutethimide  could  cause 
a  shift  of  sodium  into  the  tissue  and  this  in  turn  could  set  up  an 
osmotic  gradient  between  the  plasma  and  the  tissue.  The  increase  in 
thirst  could  be  concomitant  with  shifts  in  plasma  and  tissue  sodium  since 
hypothalamic  thirst  centers  are  regulated  by  the  osmotic  balance 
between  the  tissue  and  plams  fluids  (5).  The  lowering  of  plasma  sodium 
levels,  along  with  increase  in  tissue  sodium  levels  would  tend  to  cause 
water  to  shift  into  tissues  thus  causing  increase  in  tissue  water  and 
decrease  in  plasma  water.  A  concomitant  decrease  in  urinary  sodium 
levels,  and  an  increase  in  water  intake  would  be  expected. 

The  extra-adrenal  effects  of  AG  on  urinary  and  tissue  sodium  and 
potassium,  in  the  intact  animals  were  not  clearly  evident  at  3  days. 
Following  treatment  for  3  days  the  intact  rats  tended  to  exhibit  more 


Zoology  461 

clearly  the  traditional  symptoms  of  adrenal  insufficiency.  This  would 
be  expected  since  AG  does  not  block  the  effects  of  ACTH  on  adrenal 
cell  function,  morphology,  and  growth  (4),  and  in  intact  animals  the 
inhibition  of  glucocorticoid  synthesis  allows  compensatory  hypersecre- 
tion of  ACTH  and  leads  to  adrenal  hypertrophy  (3).  The  increased  pro- 
duction of  ACTH  would  tend  to  cause  a  compensatory  reaction  and  a 
change  in  the  urine  and  tissue  sodium  and  potassium. 

The  way  in  which  aminoglutethimide  affects  molecular  processes  in 
adrenal  steroid  snythesis  is  rapidly  being  clarified.  However,  its  effects, 
such  as  its  influences  on  extra-adrenal  action  on  water  and  electrolyte 
metabolism  and  on  sex  organs  and  thyroid  function,  are  as  yet  poorly 
understood.  Whether  it  has  one  site  of  action  which  is  common  to  all 
these  organs  is  not  known,  but  it  is  possible  that  the  drug  acts  at  one 
locus  necessary  to  the  function  of  several  organs. 

Summary 

1.  A  distinct  relationship  exists  between  the  effects  of  amino- 
glutethimide and  water  consumption. 

2.  Aminoglutethimide  produced  a  polydipsic  effect  which  is  then 
followed  by  a  polyuria. 

3.  Injections  of  aminoglutethimide  caused  adrenal  insufficiency- 
like effects. 

4.  Aminoglutethimide  appeared  to  cause  shifts  between  blood  and 
muscle  in  sodium  and  potassium,  independently  of  the  adrenals,  setting 
up  an  osomotic  gradient  which  caused  alterations  in  urine  output  of 
sodium  and  potassium. 

5.  The  presence  of  the  adrenals  appeared  to  cause  a  compensation 
which  decreaed  the  effects  of  aminoglutethimide  on  water  and  electro- 
lyte metabolism. 

Literature  Cited 

1.  Bauer,   R.,   and   J.   S.   Meyer.   1964.   Clinical   Evaluation   of  Elipten.   J.   Mich.   Med.    Soc. 
59:1829-1832. 

2.  Cash,   Ralph,   David   Schteingart,   and   Jerome   Conn.    1966.    Aminoglutethimide   and 
Metastatic  Adrenal  Cancer.  J.  Amer.  Med.  Ass.  198:1007-1010. 

3.  Dexter,   R.    N.,   L.   M.    Fishman,   A.    C.    Black,   Jr.   and    R.    L.    Ney.    1967.    Clin.    Res. 
14:61. 

4.  Kahnt,    F.    W.,   and    R.    Neher.    1966.    Uberdie   Adrenal    Steroid    Biosynthesis   in   vitro. 
Helv.  Chim.  Acta.  49:1457. 

5.  Turner,   C.   D.     1966.     General   Endocrinology,    Fourth   edition.     W.    B.    Saunders   Com- 
pany, Philadelphia,  Pennsylvania. 


The  Effect  of  Steroids  on  the  Follicle  Stimulating  Hormone  (FSH) 
Content  of  Chicken  Anterior  Pituitary  Glands 

Frank  J.  Zeller  and  Walter  R.  Rathkamp,  Indiana  University1 

Abstract 
A  variety  of  steroids  were  administered  to  cockerels  and  capons  to  note  their  effect 
on  anterior  pituitary  gland  weight  and  FSH  content.  Short-term  experiments  with 
capons  indicated  that  FSH  content  was  increased  by  testosterone  propionate  (TP), 
decreased  by  progesterone,  while  androstenedione,  dehydroisoandrosterone,  dexa- 
methasone,  pregnenolone,  androstenediol,  and  small  amounts  of  estradiol  had  little  if  any 
effect.  Birds  injected  with  TP  for  a  40-day  period  after  caponization  had  pituitary 
weights  similar  to  cockerel  controls  but  the  FSH  content  was  still  high  above  normal. 
Results  indicate  that  testosterone  alone  does  not  control  FSH  pituitary  content  in  the 
male  bird. 

The  negative  feedback  relationship  between  the  gonads  and  the 
anterior  pituitary  gland  is  still  incompletely  understood  despite  many 
years  of  experimentation.  This  is  particularly  true  in  the  case  of  the 
male.  In  addition,  the  majority  of  studies  have  been  done  with  mam- 
mals; therefore,  even  less  is  known  of  the  system  in  the  non-mammalian 
vertebrates.  The  purpose  of  this  paper  is  to  attempt  to  answer  the  ques- 
tion whether  testosterone  alone  is  responsible  for  the  control  of 
pituitary  FSH  content  and  weight  in  the  male  bird. 

Materials  and  Methods 

The  single  comb  White  Leghorn  chickens  used  in  these  experiments 
were  obtained  when  1  day  old  from  the  Indianapolis,  Indiana,  Farm 
Bureau  Co-op.  The  female  rats  used  to  assay  pituitary  FSH  content 
were  obtained  when  21  days  old  from  the  Holtzman  Company,  Madison, 
Wisconsin.  The  experimental  procedure  was  essentially  as  follows. 
Caponization  was  performed  before  the  birds  were  2  weeks  old  and 
the  capons  and  intact  controls  were  then  grown  to  the  desired  age  for 
an  experiment.  The  birds  were  then  injected  with  various  steroids. 
Testosterone  propionate  was  a  gift  from  the  Schering  Corporation,  es- 
tradiol dipropionate  a  gift  from  CIBA  Pharmaceutical  Corporation,  and 
the  dexamethasone  a  gift  from  the  Merck  Corporation.  The  other  ster- 
oids were  purchased  from  Nutritional  Biochemicals.  All  of  the  steroids 
were  carried  in  sesame  oil  and  injected  subcutaneously  in  0.1  cc 
amounts.  At  the  end  of  the  injection  period  the  animals  were  killed  and 
the  pituitary  glands  quickly  removed,  weighed,  and  then  homogenized 
in  cold  distilled  water.  The  homogenate  was  centrifuged  at  4°C,  washed, 
recentrifuged  and  the  supernatant  frozen  until  assayed  for  its  FSH 
activity.  The  FSH  activity  of  the  chicken  pituitary  glands  was  deter- 
mined by  means  of  the  human  chorionic  gonadotropin  (HCG)  augmen- 
tation assay  (4).  The  HCG  was  generously  supplied  by  Dr.  J.  B.  Jewell 
of  the  Ayerst  Laboratory.  That  chicken  pituitaries  can  be  assayed  by 
this  method  for  their  FSH  activity  was  shown  in  an  earlier  report  (5). 
At  least   10  mg  equivalent  fresh   pituitary  material  was   given  to   each 


1  Contribution   No.    840   from   the   Zoology  Department,   Indiana  University.    Supported 
by  NSF  grant  6957. 

462 


Zoology  463 

assay  rat.  Usually,  20  to  30  mg  was  administered.  In  any  one  experi- 
ment, however,  all  rats  received  the  same  amount.  The  end-point  of  the 
FSH  assay  was  ovarian  weight,  i.e.,  the  more  FSH  the  higher  the 
weight.  Appropriate  known  FSH  controls  were  run  with  each  assay 
but  only  the  data  from  the  chicken  pituitaries  are  presented  in  the 
tables.  The  FSH  was  a  gift  from  the  NIH  Endocrinology  Study  Section. 

Results 

The  response  of  cockerels  and  capons  of  several  age  groups  to 
testosterone  propionate  (TP)  is  shown  in  Table  1.  It  can  be  seen  that 
short  term  treatment  in  the  cockerel  resulted  in  no  change  in  pituitary 
weight  but  a  decrease  in  FSH  content  as  noted  by  the  lighter  weight 
ovaries  of  the  assay  animals.  In  the  capon  there  was  some  decrease  in 
pituitary  weight  but  increased  FSH  content.  Table  2  presents  data  show- 
ing the  effect  of  TP  treatment  and  withdrawal  on  FSH  content  over  a 
2-week  period.  Within  2  days  of  TP  treatment  there  was  an  increase 
in  pituitary  FSH.  This  continued  to  a  peak  at  day  4  and  was  main- 
tained through  day  14.  Withdrawal  of  TP  quickly  resulted  in  a  de- 
crease in  pituitary  FSH  and  within  7  days  the  level  was  back  to  that 
of  the  original  untreated  capons.  There  were  no  significant  differences 
in  pituitary  weights  between  the  groups.  These  results  are  similar  to 
those  obtained  in  the  male  rat  (2). 

Table   1.     The  effect  of  testosterone  propionate  and  testosterone  precursors   on   cock   and 
capon  comb  weight  and  anterior  pituitary  weight  and  FSH  content. 


Comb 

Pituitary 

Rat  FSH  Assay 

Treatment 

N 

g 

mg 

N 

Ovary  mg  ±  SE 

Testosterone  propionate  (TP) 

82-day-old  birds. 

Cock  controls 

46 

8.7 

6 

L02 

± 

lit 

Cock    +    400  Mg  TP  x  5 

53 

S.2 

5 

68 

+ 

■I" 

Capon  controls 

22 

13.8 

6 

83 

H 

5 

Capon   +   400  ^g  TP  x  5 

39 

11.9 

6 

190 

-+-_ 

18** 

105-day-old  birds. 

Cock  controls 

18 

3.23 

9.3 

9 

59 

± 

3 

Capon  controls 

15 

0.60 

15.3 

9 

tilt 

± 

5 

Capon    +   200  ^g  TP  x  8 

11 

3.41 

12.8 

7 

US 

t- 

5** 

115-day-old  birds. 

Cock  controls 

15 

9.9 

7 

56 

± 

■1 

Cock   +   200  Mg  TP  x  8 

14 

9.8 

7 

43 

+ 

■1 

Capon  controls 

8 

18.1 

s 

39 

■t- 

4 

Capon    +   200  ^g  TP  x  4 

7 

14.8 

■7 

56 

+ 

4** 

Capon    +   200  ^g  TP  x  8 

s 

L2.8 

8 

79 

-t 

2** 

Testosterone  precursors.  55-day-old  bii 

ds. 

Cock  controls  46  5.40  ±  .47  6.5  ±  .1  8  68  ±  4 

Capon  controls  26  0.56  ±  .05  9.8  ±  .3**  6  144  ±  13* 

Capons   +    500   wg  steroid  x   10  days. 

Androstenedione  27  0.93  ±  .03**  9.5  ±  .3  8  122  ±  8 

Dehydroisoandrosterone  28  0.75  ±  .06**  10.2  ±  A  8  111  ±  5* 

Androstenediol  31  0.64  ±  .04  12.1  ±  .4**  9  121  ±  3 

Pregnenolone  29  0.59  ±  .04  8.7  ±  .3**  8  147  ±  8 

Significance  levels :  *  5%  ;  **  1%. 


464  Indiana  Academy  of  Science 

Because  testosterone  alone  did  not  appear  to  return  capon  FSH 
pituitary  levels  to  that  seen  in  the  intact  chicken,  the  possibility  that 
other  steroids  might  affect  FSH  content  was  studied.  This  included  the 
female  sex  hormones  and  some  metabolic  precursors  of  testosterone 
such  as  pregnenolone,  androstenediol,  androstenedione,  and  dehydroiso- 
androsterone.  The  latter  two  are  also  secretory  products  of  the  testis 
(1).  In  Table  1  it  can  be  seen  that  only  these  two  compounds  had  any 
androgenic  effect  as  noted  by  the  comb  response.  Only  dehydroiso- 
androsterone  had  any  effect  in  reducing  FSH  content  toward  the  normal 
cockerel  level. 

Table  2.     The  effect  of  testosterone  propionate  on  the  pituitary  FSH  activity   of   109-123 

day  old  capons. 

Testosterone 
Treatment,  days 

Capon  controls 
200  ^gx2 
200  ^g  x  4 
200  ^g  x   7 
200  ^g  x   14 
200  ^g  x   7,   off  2 
200  ^g  x  7,   off  4 
200  ^g  x   7,   off  7 

The  effects  of  estradiol  and  progesterone  on  the  pituitary  gland  are 
presented  in  Table  3.  In  large  amounts,  estradiol  dipropionate  (ED) 
decreased  pituitary  weight  and  FSH  content  when  given  alone  or  in 
combination  with  TP.  Progesterone  decreased  FSH  content,  a  situation 
opposite  to  that  reported  in  the  mammal  (3). 

It  is  possible  that  the  long  time  interval  between  caponization  and 
the   experimental   period   results   in   a    system    that   is    less    sensitive    to 

Table  3.     The  effect  of  estradiol,  progesterone,   and  estradiol  plus  testosterone  on  chicken 
pituitary  weight  and  FSH  content. 


Pituitary 

Rat  FSH  Assay 

% 

N 

niK 

N 

Ovary  mg  +  se 

Increj 

5 

16.8 

6 

40   +  2 

_ 

5 

16.1 

5 

58   +  3 

48 

5 

L6.4 

6 

97  ±  7 

148 

5 

15.6 

5 

81  +  4 

104 

5 

17.6 

6 

74  +  8 

88 

5 

17.1 

6 

52  +  5 

32 

5 

14.5 

6 

54   +  4 

37 

5 

16.2 

5 

45   +   1 

13 

Total  7-day 

Pituitary 

Rat  FSH  Assay 

Treatment                                                         N 

mg 

N 

Ovary  mg  +  SE 

Estradiol  dipropionate   (ED)   or  progesterone, 

62-day-old  bii 

ds. 

Cock  controls                                                37 

6.1 

7 

78 

± 

5 

Capon  controls                                                16 

S.l 

6 

128 

+ 

6** 

Capon  +     50  Atg  ED                                  19 

7.S 

5 

115 

± 

14 

Capon    +    100  ^g  ED                                19 

8.0 

(i 

106 

± 

8 

Capon    +    200  ^g  ED                                20 

7.0 

6 

61 

± 

5** 

Capon    +    3.5  mg 

progesterone                                              18 

10.0 

5 

84 

± 

^** 

Estradiol  dipropionate  +   testosterone  propionate 

(TP),  68-day- 

old  capons. 

Controls                                                           13 

11.6 

6 

78 

± 

9 

1.4  mg  TP                                                        13 

it.  2 

4 

135 

± 

g** 

1.4  mg  TP  +   105  ^g  ED                         15 

9.7 

6 

75 

+ 

5** 

1.4   mg  TP  +  210  ^g  ED                         16 

9.5 

6 

78 

+ 

5** 

**  1%  significance. 


Zoology  465 

hormone  action  than  is  found  in  normal  birds.  Therefore,  an  experi- 
ment was  performed  in  which  birds  were  caponized  on  day  14,  then 
injected  with  50  /mg  TP  every  other  day  for  30  days.  These  birds  then 
received  30  /ng  TP  alone  or  in  combination  with  some  other  steriod  for 
10  days.  Thus,  the  total  experimental  period  was  40  days.  The  object 
was  to  try  and  create  as  normal  an  environment  as  possible  as  far  as 
TP  was  concerned  to  see  if  it  alone  would  maintain  normal  pituitary 
weight  and  FSH  content,  or  if  other  steroids  were  also  needed.  The 
results  in  Table  4  indicate  that  the  long  term  treatment  with  TP, 
although  apparently  below  physiologic  level  as  noted  by  comb  weight, 
did  keep  the  pituitary  weight  down,  but  did  not  maintain  a  normal  FSH 
concentration.  Of  the  other  steriods,  only  progesterone  had  a  depressing 
effect  on  pituitary  FSH  levels.  Dexamethasone  appeared  to  synergise 
the  action  of  TP  on  the  comb. 

Table  4.  The  effect  on  the  comb  and  pituitary  gland  of  birds  caponized  on  day  H, 
injected  every  other  day  for  30  days  with  50  nil  testosterone  propionate  (TP),  then 
injected  daily  for  10  days  with  30  ng  TP  alone  or  in  combination  with  200  ^g  TP  5  ng 
estradiol  dipropionate  (ED),  500  ng  progesterone  (prog),  or  20  ng  dexamethasone  (dexa). 


Comb  Pituitary  Rat  FSH  Assay 

Treatment                                                     N                  g  mg  N             Ovary  mg  +  SE 

Untreated  controls. 

Cock  controls                                               46                5.46  6.5  8                  68   ±     4 

Capon  controls                                            26                0.56  9.8  6                144  ±  13** 

Capons   -j-   50    ng  TP   every   other   day  for   30   days   followed   by   10-day   treatment   with   30 

ug  TP  alone  or  in  combination  with  other  steroids. 


30  ^g  TP  control 

38 

2.56 

5.5 

7 

140  ±     3 

+    200  ^g  TP 

37 

5.03 

5.0 

5 

147  +   12 

+    5  Mg  ED 

36 

2. OS 

5.3 

6 

145  ±     6 

+    500  ^g  prog. 

33 

2.38 

5.9 

6 

96  ±     8** 

+    20  ^g  dexa. 

36 

3.26 

6.7 

6 

126  ±   11 

+    ED  and  prog. 

38 

2.06 

5.3 

6 

109  +     8** 

+   TP,  ED,  prog., 

and  dexa. 

36 

6.91 

5.7 

6 

111  +     7** 

**  1%  significance  level. 

Conclusion 

The  results  of  the  experiments  presented  in  this  paper  suggest 
that  testosterone  alone  did  not  control  FSH  levels  of  the  male  chicken 
pituitary  gland. 

Literature  Cited 

1.  Connell,  G.  M.,  C.  J.  Connell,  and  K.  B.  Eik-Nes.  1966.  Testosterone  synthesis  by 
the  two-day-old  chick  testis  in  vitro.  Gen.  Comp.  Endrocrinol.  7:158-165. 

2.  Gay,  V.  L.,  and  E.  M.  Bogdanove.  1969.  Plasma  and  pituitary  LH  and  FSH  in  the 
castrated  rat  following  short-term  steroid  treatment.  Endrocrinology  84:1132-1142. 

3.  Schwartz,  N.  B.  1968.  Newer  concepts  of  gonadotropin  and  steroid  feedback  control 
mechanism,  p.  33-50.  In  J.  Gold  (ed. )  Textbook  of  gynecologic  endrocrinology. 
Hoeber,  N.Y. 

4.  Steelman,  S.  L,  and  F.  M.  Pohley.  1953.  Assay  of  FSH  based  on  the  augmentation 
with  human  chorionic  gonadotropin.  Endrocrinology  53  :604-616. 

5.  Zeller,  F.  J.  1965.  The  follicle  stimulating  hormone  content  of  chicken  anterior 
pituitary  glands.  Proc.  Indiana  Acad.  Sci.  74  :351. 


Radiotelemetry  with  the  Big  Brown  Bat  (Eptesicus  fuscus)1 

James  B.  Cope,  Donald  R.  Hendricks  and  William  B.  Telfair, 

Earlham  College 

Abstract 
A  frequency  modulated  transmitter   was   designed  specifically  for  the   big   brown   bat, 
Eptesicus  fuscus,   to  aid  in   studying   movement   in   a  hibernaculum.    The   signal   from   the 
1.2    g   transmitter   was   recorded   on    a   tape   for    1    second   in    every    6   minutes,    providing 
recordings  from  a  period  of  7  days  and  6  hours  on  a  30-minute  tape. 

The  research  worker  dealing  with  live  animals  in  the  natural  en- 
vironment is  plagued  with  the  possibility  of  disturbing  the  subject 
sufficiently  to  make  his  data  unreliable.  We  have  been  concerned  about 
the  disturbance  factor  in  bat  hibernacula.  To  aid  in  studying  bat  move- 
ment, we  designed  a  transmitter  which  was  light  enough  so  it  did  not 
appear  to  hinder  the  bats  in  flight  nor  alter  their  behavior. 

Methods 

Cochran  et  al.  (1)  used  a  2.5  g  transmitter  to  study  thrushes  and 
found  no  hindrance  in  flight  or  change  in  behavior  with  transmitters  up 
to  15%  of  the  body  weight.  Because  the  weight  of  Eptesicus  fuscus 
ranges  from  14  to  24  g  (3),  the  transmitter  had  to  weigh  less  than  2.1  g 
to  allow  freedom  in  flight  and  movement. 

A  transmitter  designed  by  Skutt  et  al.  (2)  was  modified  by  con- 
structing the  body  out  of  plexiglas  0.562  inch  in  diameter  and  0.141 
inch  in  thickness.  The  completed  transmitter  weighed  1.1  to  1.2  g  as 
opposed  to  the  1.5  g  transmitter  built  by  Skutt  et  al.  (2).  Three  turns  of 
No.  28  magnet  wire  in  the  threads  on  the  outside  of  the  form  (tapped 
at  one  turn)  served  as  the  inductor  in  an  LC  (Hartley)  oscillator  and 
also   as   the   transmitting  antenna    (Figs.    1,   2,   3). 

The  signal  is  an  unmodulated  carrier  which  produced  quieting  in  a 
normal  FM  radio.  To  produce  an  audio  output  from  the  receiver  when 
the  functioning  transmitter  was  within  range,  the  oscillation  of  the 
local  oscillator  was  frequency  modulated  at  an  audio  rate.  Later,  a 
1000  ohm  resistor  was  dropped  from  the  construction  to  increase  the 
efficiency  of  the  transmitter  and  lengthen  the  life  of  the  battery  (Figs. 
1,2). 

The  original  transmitter  drew  approximately  140  n  amp.  When 
one  resistor  was  dropped,  it  drew  65  /x  amp  which  more  than  doubled 
the  life  of  the  battery.  This  caused  the  oscillator  to  "motor  boat"  at 
an  audio  rate.  The  expected  battery  life  was  approximately  14  days. 

A  gating  circuit  was  also  designed  to  turn  on  both  the  receiver  and 
tape  recorder  for  1  second  every  6  minutes.  This  was  an  astable  oscil- 
lator, followed  by  a  phase  inverter  (a  mono-stable  oscillator),  then  a 
current  amplifier  and  power  transistor  switch  (Fig.  4).  This  allowed  a 
30-minute  tape  to  record  for  7  days  and  6  hours. 


1  Funds  were  provided  by  National   Science   Foundation   Undergradute   Research   Par- 
ticipation  grant  No.    GY5991   and   COSIP   grant  No.    GY4707. 

466 


Zoology 


407 


SIDE     VIEW 

note:    threads  are    .009"    deep   with    26   to  32  turns    per  inch. 

Figure  1.    Modified  Skutt  transmitter. 


A  mock  transmitter  was  built  by  using  a  soft  aluminum  shell 
measuring  0.562  inch  in  diameter  and  0.125  inch  in  thickness.  It  was 
filled  with  lead  shot  to  give  it  a  weight  of  1.5  g.  The  mock  transmitter 
was  attached  to  a  bat  with  adhesives  to  test:  1)  irritation  from  glue 
and,  2)  irritation  from  placement.  Surgical  cement  (Vi-drape)  was  used 
but  proved  unreliable  because  it  caused  loss  of  hair  and  skin  irritation. 
A  benzoin  tincture  first  applied  to  the  hair,  then  contact  cement  (Weld- 
wood)  to  both  the  hair  and  transmitter  surface  gave  the  best  bond 
and  least  irritation.  When  the  transmitter  was  attached  to  the  neck 
region  or  the  scapular  region,  the  bat  refused  to  fly,  and  bit  and 
scratched  at  the  transmitter  until  it  was  pulled  off.  When  attached 
just  above  the  caudal  region,  it  caused  no   irritation  and  no   noticeable 


4(58 


Indiana  Academy  of  Science 


IOOK-  470K 


01  J L  ispf 


T jif-L..* 


I00K-470K 

7pf 

*                .01- 

.l/.f    = 

LI,.,  j- 

<  'K 

Figure  2.    Circuitry  of  transmitter. 


Figure  3.    Modified  transmitter. 


7^7 
Figure  4.    Gating  circuit. 


change  in  flight  pattern  or  feeding  behavior  (Fig.  5).  A  sample  of  8 
bats,  4  males  and  4  females,  with  an  average  weight  of  18  g  was  used 
for  testing. 

The    bats    and   all    equipment   were    taken    to    the   hibernaculum.    A 
transmitter  was  attached  to  a  bat  and,  hand-held,  the  bat  was  moved 


i 1 

1  e*. 


Figure  5.    Eptesicus  fuscus  with  transmitter  attached. 


Zoology  469 

to  different  possible  locations  in  the  site  to  determine  effectiveness  of 
equipment  and  antennae.  Best  reception  was  with  a  half-loop  antenna 
at  the  range  0-15  m.  The  signal  could  be  picked  up  within  this  range 
despite  placement  of  the  bat  behind  jagged  walls  or  in  holes  in  the 
ceiling.  Because  the  study  area  is  a  tunnel,  no  interference  was  found 
inside  the  center  half  of  the  tunnel  from  FM   stations. 

All   equipment  was   tested  for  temperature   stabilization  at  — 8°C; 
all  proved  effective. 

Acknowledgement 
We  thank   Dr.  Richard  W.   Stow  of  Ohio   State   University  for  as- 
sistance and  use  of  his  laboratory,  Margaret  Brown  for  art  work,  and 
Richard  Otis  for  a  photograph. 

Literature  Cited 

1.  Cochrane,  W.  W.,  G.  G.  Montgomery,  and  R.  R.  Graber.  1967.  Migratory  flights 
of  Hylocichla  thrushes  in  spring :  a  radiotelemetry  study.  In  The  Living  Bird,  6th 
Ann.   Lab.   of  Ornithology,   Cornell  Univ.,   Ithaca,  N.Y.   249  p. 

2.  Skutt,  H.  R.,  R.  G.  Beschle,  D.  G.  Moulton,  and  W.  P.  Loella.  1967.  New  sub- 
miniature  amplifier-transmitters  for  telemetering  biopotentials.  Electroeneephal.  and 
Clin.  Neurophysiol.  22  :275-277. 

3.  Walker,  E.  P.,  and  associates.  1964.  Mammals  of  the  World,  v.  1.  The  Johns 
Hopkins  Press,  Baltimore.  644  p. 


Status  of  My otis  lucifugus  in  Indiana1 

James  B.  Cope  and  Donald  R.  Hendricks,  Earlham  College 

Abstract 
Twelve  nursery  colonies  of  the  little  brown  bat,  Myotis  lucifugus,  were  observed  in 
Indiana  during  June-August  1969.  Seven  colonies  declined  in  population  while  5  other 
colonies  seemed  stable  when  compared  to  previous  data.  Samples  were  taken  from  each 
colony  for  banding  and  the  numbers  of  remaining  bats  were  estimated,  giving  an 
estimated  total  population. 

In  the  past  few  years  much  has  been  written  about  the  decline  of 
animal  populations.  Bats  in  particular  have  been  included  in  this  de- 
cline (1,  2).  The  senior  author  suspected  this  decline  but  had  no  docu- 
mented evidence.  The  junior  author  devoted  the  past  summer  to  a  study 
of  12  well-established  nursery  colonies  of  Myotis  lucifugus.  The  results 
of  this  study  are  in  Table  1.  Seven  colonies  declined  in  population  while 
five  colonies  seemed  to  have  a  stable  population. 

Table  1.    Population  decline  in  7  nursery  colonies  of  Myotis  lucifugus  in  Indiana. 


Colony 

No. 

First 

No. 

Last 

No. 

% 

Date 

Bats 

Date 

Bats 

Banded 

Decrease 

Reelsville 

1 

8 

Aug    59 

600 

12 

Aug    69 

0 

3197 

100% 

Brookville 

2 

23 

Aug    60 

650 

20 

Jul     69 

0 

647 

100% 

Newbern 

3 

22 

Aug   60 

850 

L3 

Aug    69 

0 

2751 

100% 

Shoals 

4 

28 

Aug   58 

1133 

L3 

Aug    69 

275 

unknown 

76% 

Milroy 

5 

27 

Jul     62 

800 

2 

Jul     69 

125 

1586 

84% 

New  Castle 

6 

12 

Aug   58 

1000 

15 

Aug    69 

35 

1239 

96% 

Pennville 

7 

20 

Jul     65 

1060* 

22 

Jul     69 

581* 

1719 

48% 

*  Evening  count. 

The  breakdown  of  the  seven  colonies  showing  a  declining  popula- 
tion is  as  follows: 

Three  colonies  were  completely  gone  from  the  nursery  sites.  One 
of  these  was  built  out;  one  was  exterminated  with  50%  DDT  dust;  the 
third  colony  left  because  $10.00  worth  of  mothballs  was  tacked  up  in 
their  roosting  areas.  Over  the  past  10  years,  6,595  bats  were  banded  in 
these  colonies. 

The  fourth  colony  decreased  approximately  76%  over  the  past  11 
years,  probably  due  to  exterminators  using  DDT  dust  in  the  roost  and 
to  attempts  to  asphyxiate  the  bats  with  automobile  exhaust  fumes.  Fly- 
ing bats  and  dead  bats  were  seen;  it  is  unknown  how  these  bats  re- 
mained alive.  The  strength  of  the  DDT  is  unknown. 

The  fifth  colony  decreased  84%  over  the  past  6  years.  The  owner 
informed  us  that  physiologists  from  a  university  had  been  taking  banded 
and  unbanded  bats  each  year  for  their  experiments.  They  also  took 
females  from  the  nursery  colony  before  the  young  had  dropped.  The 
authors  are  convinced  that  there  needs  to  be  better  cooperation  between 


1  Funds   were   provided   by   National   Science   Foundation    Research    Participation    Grant 
No.  GY  5991. 

470 


Zoology  471 

researchers    so    that    one    set   of   research    efforts    is    not    destroyed    by 
another.  A  total  of  1,586  bats  have  been  banded  in  this  colony. 

The  sixth  colony  declined  96%,  reason  unknown.  The  seventh  colony 
had  a  48%  reduction,  reason  unknown,  in  spite  of  the  fact  that  this 
population  has  been  examined  (by  evening  counts)  more  critically  than 
any  other  in  the  state. 

Literature  Cited 

1.  Humphrey,    S.    R.    1964.    Extermination    at    Indiana    Myotis    lucifugua    nurseries.    Bat. 
Res.  News  5(4)  :34. 

2.  Mumford,    R.   E.,   and   J.    B.   Cope.    1964.   Distribution   and   status   of   the   Chiroptera   of 
Indiana.  Amer.  Midland  Natur.  72(2)  :473-489. 


Vertebrate  Remains  from  an  Indiana  Cave 
Ronald  L.  Richards,  Indiana  University 

Abstract 

Excavation  of  a  shallow  sandstone  capped  cave  in  Monroe  County,  Indiana,  pro- 
duced skeletal  remains  of  13  vertebrate  species.  These  were  opossum,  bat,  black  bear, 
raccoon,  gray  fox,  chipmunk,  cottontail,  deer,  bison,  vulture,  frog  or  toad,  snake,  and 
tortise.  The  bear  and  bison  may  be  of  some  antiquity,  125  years  Before  Present  at 
minimum.  The  remainder  are  more  recent. 

The  remains  differ  from  those  reported  for  other  Indiana  caves  in  a  higher  fre- 
quency of  opossum,  a  greater  variety  of  sub-mammalian  forms,  and  possibly  in  the  lack 
of  mustelid  carnivores. 

Only  a  few  surface  investigations  of  vertebrate  material  from 
more  "typical"  Indiana  caves  have  been  made  (2,  3,  5,  6).  One  paper 
reports  minor  excavation  of  Pleistocene  material  (7).  The  present 
excavation  is  the  first  extensive  study  of  the  deposits  from  any  Indiana 
cave. 

Thundermug  Bone  Cave  is  located  6.6  miles  west-southwest  of 
Bloomington,  Indiana  (NE&  of  the  NWV4  of  the  SW&,  sec.  17,  T8N, 
R2W,  Whitehall  Quadrangle,  Monroe  Co.,  Indiana).  A  ridge  top  sand- 
stone formation  forms  the  cave  ceiling.  The  cave  consists  of  three  main 
chambers.  The  entrance  room  was  9  feet  long,  6  feet  wide,  and  6  feet 
high;  the  side  room  was  10  feet  long,  8  feet  wide,  and  4  feet  high;  and 
the  lower  room  an  average  diameter  of  about  6  feet  with  a  10-foot 
dimension  and  15  feet  high.  Excavation  thereafter  increased  all  the 
room  depths. 

The  fill  consisted  of  sandstone  breakdown  and  limestone  solution 
fragments,  well  interspersed  in  a  great  quantity  of  dirt.  The  sequence 
of  events  within  the  cave  thus  seems  to  have  been  a  gradual  dissolving 
free  of  the  irregularities  of  the  limestone  walls  and  crumbling  and 
collapse  of  the  sandstone  ceiling,  accompanied  by  a  gradual  filling  of 
the  cavern  with  dirt  from  the  cave  entrance. 

Bones  were  found  throughout  most  of  the  stratigraphic  sequence. 
The  deposition  of  bear  bones  in  the  entrance  room  was  mostly  second- 
ary, being  derived  from  near  the  cave  entrance.  All  bones  in  the  lower 
room,  mostly  of  deer  and  bear,  were  in  secondary  deposits  derived  from 
the  entrance  room.  The  bones  in  the  side  room  were  apparently  at  the 
original  locus  of  deposition,  although  much  disarrayed. 

Excavations  extended  from  November,  1964  to  July,  1967.  The  en- 
trance room  was  excavated  to  bedrock  at  5.5  feet  posteriority,  and  7.5 
feet  to  a  base  of  sandstone  breakdown  and  thick  clay  deposits  near  the 
entrance  slope.  The  lower  room  was  dug  to  bedrock  at  1  foot  and  the 
side  room  excavated  to  3  feet,  a  level  where  bones  were  absent. 

Only  partial  skeletons  were  recovered.  This  is  due  to  bone  loss 
through  solution  fissures  in  the  floors  of  the  lower  and  side  rooms,  as 
well  as  to  possibilities  of  only  partial  skeletons  having  eroded  into  the 
cave  (bison?),  and  partial  animals  having  been  dragged  in  by  predators 
and  scavengers. 

The  shallow  chambers  of  Thundermug  are  nearer  the  surface  than 
those  of  the  more  typical  limestone  caves.  Consequently  the  temperature 

472 


Zoology 


473 


.2  § 

CO    «* 

*  5 

a   O 

OX 
O  +> 


3 


"Ml 

r2      ^^ 

""51 J^ 


m    m    2  V 


to   co   ;-   r   >-. 

r  *  g  S  1  si 


IS 
,11 


■S-S  It  «  5  3 


X 


££ 


i  e 


b.  :>  M 

1)    0)    a> 


b  ^  u 


co  co 

a  a 

as  a! 

5h  Jh 


*  s  S 


HOONOH" 
CO        oi 


C 
3 


0)  ■» 


0 

II            CO  ff- 

ii        bt>  co 

1  1— ' 

i     i        p;  <u 
i     i      .5  cj 

£  2 

i    i       sjx 

c 

bJD 

HNHMH 


•2 

1 1 


3  J. 


^2 


^d    ~' 


2    3    CO  *«i 


j    «    S 


o 
•2  .£> 


5e  °C 


^  g  e  e  | 


CO    C 


CO     J 

|S  5  co  co 

co    C3    ~ 


8   ^ 


5*  .2    ft-W     S 


g  s  »  a)  ° 


<o  e  s> 


CS     O  r— i 


^      h^^O^Oh 


474 


Indiana  Academy  of  Science 


FIGURE   1.      Skull  of  one  of  the  black  bears,  Urus  americanus,  recovered. 


of  the  cave  is  modified  more  by  the  above-ground  temperature  than  in 
a  typical  cavern  in  which  the  temperature  of  the  limestone  bedrock 
exerts  a  dominant  control.  Thus  in  Thundermug  summer  temperatures 
were  observed  to  be  higher  and  winter  temperatures  lower  than  the 
approximate  54°F  of  the  average  Indiana  cavern  (1).  In  addition,  the 
cave  is  often  dry  and  somewhat  south-facing,  and  manages  to  filter 
some  dim  light  into  its  shallower  portions,  thus  the  shallower  portions 
of  the  cavern  probably  were  attractive  to  animals  which  would  not 
ordinarily  use  the  typical  cave  (e.g.  vultures).  The  lower  winter  tem- 
peratures may  have  contributed  to  death  of  some  of  the  animals. 

The  forms  recovered  in  the  excavation  are  listed  within  Table  1. 
(All  material  collected  is  on  file  with  the  author).  Three  of  the  six 
opossums  found  had  not  yet  completed  the  third  lower  premolar  erup- 
tion. Vultures  often  nest  in  shallow  cavities.  The  Thundermug  remains 
may  have  been  a  nesting  group,  since  an  adult  and  two  young  birds 
were  closely  associated. 

Indications  of  scavengers  and  predators,  in  the  form  of  crushed 
or  tooth-punctured  bone  seem  to  be  present  among  the  opossum,  rac- 
coon, and  cottontail  remains.  Gnawings  on  many  of  the  deer  and  bear 
bones  indicate  the  presence  of  small  rodents,  possibly  Peromyscus 
leucopus  (Table  1). 

Bear  seem  to  have  been  exterminated  from  south-central  Indiana 
about  1837,  and  the  bison  about  1778  (8).  The  related  deposits  presum- 
ably are  at  minimum  125  years  old.  The  bears  were  at  the  2.5  and  3 
feet  levels,  and  the  bison  at  nearly  3  feet,  indicating  some  antiquity, 
although  no  conclusion  can  be  drawn  because  the  accumulation  and 
erosional  history  of  the  cave  are  not  known. 


Zoology  475 

The  black  bear  material  is  noteworthy.  Although  historical  records 
are  abundant,  as  is  evidence  of  their  former  presence  such  as  wallows 
and  scratch  marks  on  trees  and  cave  walls,  nothing  remains  of  the  bears 
themselves  except  for  occasional  finds  of  skeletal  material  (4,  8).  The 
Thundermugs  ursids  were  compared  with  measurements  from  other 
speciments  of  Ursus  americanas,  and  are  clearly  of  the  U.  americanus 
type  (7,  9).  Figure  1  illustrates  one  of  the  skulls. 

In  comparing  Indiana  cavern  studies,  the  scope  of  the  investigations 
should  be  taken  into  account.  Both  Banta  (2)  and  Blatchley  (3)  were 
concerned  with  the  observation  of  living  animals  and  evidence  of  their 
presence.  Banta's  study  was  of  one  cave;  Blatchley's  of  several  caves. 
Hahn's  study  (6)  and  the  present  excavation  both  concerned  past 
vertebrate  faunas.  Hahn's  (6)  collection,  from  only  one  cave,  seems  to 
be  of  more  recent  material  than  that  in  Thundermug. 

The  results  from  the  various  cavern  studies  for  Indiana  are  com- 
pared in  Table  1.  Thundermug  included  an  unusual  number  of  opossums 
at  different  stratigraphic  levels.  This  may  indicate  use  of  the  cave  over 
a  long  period  of  time.  Bats  and  carnivores  were  found  in  all  four  studies, 
although  evidence  of  mustelids  was  found  in  both  the  "observation" 
studies,  and  none  found  in  either  of  the  osteological  studies.  Several 
of  the  carnivore  types  presumably  inhabited  the  cave.  In  contrast  to  the 
possible  inhabitants  are  the  accidently  introduced  forms.  These  might 
include  bison,  deer,  and  chipmunk. 

In  summary,  the  Thundermug  material  differs  from  that  of  other 
Indiana  caverns  studied  in  the  larger  number  of  opossums,  greater 
variety  of  sub-mammalian  forms,  and  possibly  in  the  lack  of  mustelid 
carnivores.  These  differences  may  be  related  in  part  to  the  ''attractive- 
ness" of  the  cavern. 

Acknowledgements 

I  thank  A.  Gregory  James,  Mark  A.  Wright,  and  Steven  Rand  Red- 
meier  for  help  with  the  excavation.  J.  L.  Paradiso,  U.  S.  National 
Museum,  kindly  identified  the  bison  fragments,  and  William  H.  Adams, 
Indiana  University,  aided  me  with  comparative  ethnozoological  material. 

Literature  Cited 

1.  Addington,  A.  R.  1926.  A  preliminary  report  upon  the  survey  of  Indiana  caves 
with  special  reference  to  Marengo  Cave.  Indiana  Yearbook  for  1926,  p.  303-313. 

2.  Banta,  Arthur  M.  1907.  The  fauna  of  Mayfields  cave.  Carnegie  Inst,  of  Wash., 
Pub.  No.  67,  114  p. 

3.  Blatchlfy,  W.  S.  1897.  Indiana  caves  and  their  fauna.  21st  Annu.  Rep.  Dep.  Geol. 
Natur.  Resources  Indiana  for  1896,  p.  121-212. 

4.  Collett,  John.  1883.  List  of  fossils  found  at  Porter's  Quarry,  one  and  one  fourth 
miles  west  of  Rensselaer  on  the  south  side  of  the  Iroquois  River.  12th  Annu.  Rep. 
Indiana  Dep.  Geol.  Natur.  Hist  for  1882,  1883,  p.  73. 

5.  Cope,  E.  D.  1873.  Report  on  the  Wyandotte  Cave  and  its  fauna.  Third  and  Fourth 
Annu.  Repts.  Geol.  Surv.  Indiana  for  1871  and  1872,  p.  157-182. 

6.  Hahn,  Walter  L.  1906.  The  mammalian  remains  of  the  Donaldson  Cave  (3  miles 
southeast   of   Mitchell).    Proc.    Indiana   Acad.    Sei.    22:142-144. 

7.  Hay,  Oliver  P.  1911.  The  Pleistocene  period  and  its  vertebrata.  36th  Annu.  Rep. 
Dep.  Natur.  Resources  Indiana,  p.  541-784,  1912. 

8.  Lyon,  Marcus  Ward.  1936.  Mammals  of  Indiana.  Univ.  Press,  Notre  Dame,  Indiana, 
384  p. 

9.  Merriam,  C.  Hart.  1896.  Preliminary  synopsis  of  the  American  bears.  Proc.  Biol. 
Soc.  Wash.  10  :65-83. 


Comparisons  of  Rewarming  from  Natural  Torpidity  and  Induced 

Hypothermia  in  Chipmunks  (Tamias  striatus)  with  Reference  to 

Heart  Rate  and  Temperature  Relationships 

Richard  E.  Schafferj  and  Robert  W.  Bullard,  Indiana  University 

Abstract 

Chipmunks  live-trapped  in  the  Bloomington  area  during  the  summer  and  early  fall 
of  1968  were  housed  under  four  environmental  conditions  to  study  their  effects  on 
natural  torpidity.  Beginning  in  late  December  and  continuing  into  March,  torpid  animals 
were  observed  in  three  of  the  four  groups.  Depending  on  the  group,  30  to  50%  of  the 
animals  experienced  forms  of  torpidity  ranging  from  deep  to  very  shallow.  In  a 
number  of  animals  provoked  to  arouse,  heart  rates  and  thoracic  and  colonic  temperatures 
were  recorded  during  the  rewarming  process.  Since  a  majority  of  torpid  states  were  so 
shallow  that  arousal  was  provoked  before  the  animals  could  be  fitted  with  recording 
leads,  individuals  from  the  different  groups  were  subjected  to  an  induced  hypothermia  by 
"jar  cooling,"  after  which  their  heart  rates  and  body  temperatures  were  recorded  dur- 
ing rewarming. 

Animals  provoked  to  arouse  from  natural  torpidity  demonstrated  intensive  shivering, 
while  rewarming  with  extreme  rapidity,  generating  steep  heart  rate  and  thoracic 
temperature  curves  with  time,  which  contrasted  to  lagging  colonic  temperatures  as 
much  as  18.5°C  below  thoracic  temperatures.  Rewarming  of  "jar  cooled"  animals,  at- 
tended by  varying  degrees  of  shivering,  was  considerably  slower,  resulting  in  heart  rate 
and  both  temperature  curves,  with  time,  of  similar  sigmoid  shape  and  smaller  temperature 
gradients.    The    significance    of    these    relationships    is    discussed. 

The  eastern  chipmunk,  Tamias  striatus,  is  a  member  of  the  tribe 
Marmotini,  (9).  This  tribe  also  includes  the  woodchuck  {Marmota)  and 
the  ground  squirrels  (Citellus).  While  physiological  investigations  of 
hibernation  experienced  by  the  Marmotini  have  mostly  concentrated  on 
Citellus  and  Marmota,  typical  deep  hibernating  forms,  only  a  few 
preliminary  investigations  have  been  conducted  on  T.  striatus  (4,  7, 
8,  10),  and  their  results  appear  to  indicate  this  animal  is  not  a 
typical  marmotine  hibernator. 

Although  the  depths  of  torpidity  have  been  described  by  rectal 
temperature  measurements  (7,  8,  10),  temperature-heart  rate  relation- 
ships of  isolated  hearts  have  been  reported  (4),  and  oxygen  con- 
sumption measurements  of  torpid  and  arousing  animals  have  been 
made  (7),  one  important  physiological  aspect  of  the  torpor  experienced 
by  T.  striatus  is  missing.  No  data  concerning  measurements  of  heart 
rates  and  temperatures  during  arousals,  as  exist  for  the  other  Marmo- 
tine hibernators  (3,  5,  6),  have  been  offered.  Because  of  this,  it  was 
decided  that  the  first  object  of  this  investigation  should  be  a  study  of 
heart  rates   and   temperatures    during  provoked   arousals. 

Second,  since  the  degree  of  torpor  experienced  by  T.  striatus  was 
generally  brief  and  shallow,  possibly  suggesting  either  an  intolerance 
to  deep  hypothermia  or  insufficient  rewarming  abilities,  from  which 
most  animals  aroused  before  they  could  be  fitted  with  recording  leads, 
the  decision  was  made  to  attempt  to  assess  their  tolerance  to  hypo- 
thermia and  rewarming  capabilities  by  subjecting  individuals  to  an 
induced  hypothermia   (1)   by  means  of  "jar  cooling." 


1  Trainee    of    Public    Health    Service,    National    Institutes    of    Health,    Grant    TOl    ES 
00075-03S1. 

476 


Zoology  477 

Materials  and  Methods 

Chipmunks  were  live-trapped  in  the  vicinity  of  the  Indiana  Uni- 
versity campus  from  May  through  October,  1968.  All  animals  were 
caged  individually  in  two  animal  rooms  with  different  photo-periods. 
One  room  was  windowless  in  which  a  12-hour  photoperiod  of  artificial 
light  and  a  temperature  of  19°  to  21  °C  were  maintained.  The  other 
room  contained  a  window  which  admitted  natural  light  and  had  a 
temperature  which  fluctuated  between  17°  and  23  °C.  All  chipmunks 
were  maintained  on  a  diet  of  Wayne  Lab-Blox  Shorts  (Allied  Mills,  Inc., 
Chicago,  111.),  periodically  supplemented  with  sunflower  seeds  and 
pieces  of  fresh  lettuce,  carrots  and  apples,  and  water  ad  libitum. 

Early  in  October,  1968,  six  chipmunks  were  placed  outdoors  on  a 
roof  in  individual  cages  provided  with  hinged-topped  nest  boxes.  These 
animals  were  then  supplied  with  nesting  materials  until  additional  ma- 
terials were  refused.  By  mid-October,  the  chipmunks  housed  indoors 
had  also  been  provided  with  the  same  type  of  accomodations. 

In  mid-December,  three  chipmunks  were  placed  in  a  refrigerator. 
For  the  first  week,  the  temperature  was  gradually  decreased  to  9°C.  At 
the  beginning  of  the  second  week,  the  temperature  was  dropped  to  6° 
to  3°C.  The  refrigerator  was  unlighted  except  for  a  small  amount  of 
12-hour  light  entering  a  3V2-inch  diameter  ventilation  hole  near  the  top 
of  one  side  of  the  refrigerator.  Humidity  was  always  high,  as  evidenced 
by  water  condensation  on  the  inside  of  the  refrigerator. 

All  animals  were  provoked  to  arouse  at  room  temperature  (20°  to 
24°C)  by  removing  them  from  their  nests,  fitting  them  with  EKG  and 
temperature  recording  leads,  and  placing  them  in  a  rectangular  acrylic- 
plastic  chamber  provided  with  ventilation  ports.  Total  time  elapsed  for 
these  manipulations  was  3  to  5  minutes. 

Electrocardiogram  leads  were  fabricated  from  highly  polished 
safety  pins  connected  to  a  Sanborn  High  Gain  Preamplifier  (model 
150-2700)  and  Recorder  (model  154-100B).  The  ground  was  connected  to 
copper  screening  sandwiched  between  the  walls  of  the  chamber.  Prior 
to  subdermal  insertion  in  a  Lead  II  orientation,  the  tips  of  the  safety 
pins  were  wiped  with  70%  alcohol  and  dipped  in  1%  Novocain  (Win- 
throp  Laboratories,  New  York,  N.  Y.).  The  EKG  was  recorded  every  1 
or  2  minutes  and  the  heart  rate  calculated  by  counting  from  this  record. 
The  EMG  was  recorded  as  interference  on  the  EKG  record. 

Temperatures  were  measured  with  a  YSI  Tele-Thermometer,  Model 
41TS,  and  400  and  500  Series  thermistor  probes  (Yellow  Springs  Instru- 
ment Co.,  Inc.,  Yellow  Springs,  Ohio).  Thoracic  temperatures  were 
obtained  using  a  511  probe  inserted  into  the  chest  cavity  through  the 
barrel  of  an  18G  sterile  needle  which  had  penetrated  between  the  third 
and  second  last  ribs.  Previously,  the  probe  had  been  wiped  with  70% 
alcohol  and  the  needle  dipped  in  1%  Novocain.  After  the  probe  had  been 
pushed  anteriorly,  to  loop  the  tip  near  the  base  of  the  heart,  the  needle 
was  withdrawn  and  the  lead  secured  to  the  animal's  back  with  adhesive 
tape.  Deep  colonic  temperatures  were  obtained  with  a  401  probe,  wiped 
with  70%  alcohol  and  dipped  in  glycerine,  then  inserted  3-5  cm  into  the 
colon  and  secured  to  the  base  of  the  tail  with  adhesive  tape.  All  probes 
had  been  previously  calibrated  ±  0.2  °C  in  a  well-stirred  water  bath. 


478 


Indiana  Academy  of  Science 


Hypothermia  was  induced  by  placing  a  weighed  animal  in  a  glass 
vessel  which  had  been  pre-cooled  to  0°  to  2°C  by  surrounding  it  with 
crushed  ice  in  a  styrofoam  chest.  The  vessel  was  then  sealed  air-tight 
with  the  animal  rebreathing  the  chilled  air  as  it  slowly  cooled  and  lost 
consciousness.  This  hypothermia  was  judged  complete  when  the  animal 
1)  exhibited  a  shallow,  abdominal  respiration  rate  of  30-50  per  minute; 
and  2)  was  unable  to  right  itself.  Approximtely  2%  hours  of  cooling 
were  required  to  meet  these  conditions,  after  which  the  animal  was 
removed  from  the  vessel  and  prepared  for  recording  of  rewarming,  ac- 
cording to  the  procedure  for  provoked  arousals. 

Results 

It  was  the  end  of  December  before  any  torpor  was  observed  in  the 
chipmunks  and  this  was  observed  in  an  outdoor  animal.  From  then  on, 
more  torpid  animals  were  observed.  The  greatest  frequency  of  torpidity 
occurred  from  mid-January  to  mid-March,  after  which  there  was  an 
abrupt  end.  Of  six  outdoor  chipmunks,  three  were  observed  in,  and 
provoked  from  torpor  one  or  more  times  during  this  period.  Animals  in 
the  naturally  lighted  room  began  showing  signs  of  lethargy  in  mid- 
January  with  4  of  12  animals  eventually  being  observed  in  states  of 
light  torpor.  While  a  few  of  the  12-hour  photoperiod  animals  appeared 
to  show  some  lethargy  during  this  time,  none  was  observed  in  any  state 
of  torpor.  One  of  the  three  refrigerator  animals  was  observed  twice  in 
deep  torpor,  the  first  occurring  late  in  February. 

Figure  1  shows  the  course  of  the  second  arousal  provoked  in  a 
female   chipmunk,    weighing    132. 3g,   which    had    become    torpid    in    the 


40 

36 

32 

w     28 

UJ 

CC 

cr 

UJ 

o_ 

u!     20 

H 

16 
12 


3/12/69, NATURAL   F-IG/15/68  ,  R  EF  (2) 


600 


520 


440  2 


-  360  y 


200 


120 


40 


I  0    20 


50   60    70 


30   40 

TIME  (MIN  ) 

Figure  1.     Progression     of     thoracic     and     colonic     temperatures,     and     heart     rate     in     a 

refrigerator    housed,    female    chipmunk     (132.3g)     while    rewarming    from    natural    torpor 

during  a  provoked  arousal. 


Zoology 


479 


560 


480 


?    400 

2 


<     320 

LU 

CD 


240 


80 


3/12/69, NATURAL   F   10/15/68 ,  REF   (2) 


0  4  8  12         16        20        24        28        32        36  40 

TEMPERATURE  CO 

Figure  2.     Heart   rate    and   thoracic    temperature    (°C)    relationships,    during  acceleration 

of  the  heart  rate  to  maximum,  for  the  female  chipmunk  shown  arousing   in  Figure   1. 


refrigerator.  From  the  first  recorded  heart  rate,  thoracic  and  colonic 
temperatures  of  68  beats  per  minute,  11.7°C  and  10.8°C,  respectively, 
this  animal  completely  rewarmed  in  70  minutes.  Figure  1  also  shows 
how  closely  heart  rates  and  thoracic  temperatures  increase  together  such 
that  by  27  minutes  into  recording  time  a  maximum  heart  rate  of  535 
beats  per  minute  was  associated  with  a  thoracic  temperature  of  35.4 °C. 
The  rapid  increase  in  heart  rate  and  thoracic  temperature  was  attended 
by  intense  shivering.  Shivering  was  recorded  as  interference  on  the  EKG 
from  the  onset  of  recording,  increasing  in  intensity,  finally  ceasing  at  28 
minutes,  when  a  heart  rate  of  530  beats  per  minute  and  a  thoracic 
temperature  of  35.7  °C  were  observed.  After  shivering  ceased,  thoracic 
temperatures  increased  only  slightly  over  the  remaining  time  and  heart 
rate  fluctuated  greatly.  Colonic  temperatures,  on  the  other  hand,  showed 
an  entirely  different  progression.  While  thoracic  temperature  increased 
an  average  of  0.86° C  per  minute  during  the  period  of  shivering,  colonic 
temperatures  lagged  considerably,  only  increasing  an  average  of  0.31  °C 
per  minute.  It  was  during  this  period  (at  25  minutes)  that  a  maximum 
gradient  of  18.4 °C  existed  between  the  two  temperature  regions.  From 
this  point  colonic  temperatures  increased  rapidly,  and  during  the  next 
12  minutes  averaged  almost  1°C  per  minute.  Thereafter  the  rate  of 
colonic  temperature  increase  markedly  declined.  The  colonic  temperature 
curve  also  shows  a  "dip"  or  "plateau"  just  prior  to  the  development  of 
the  maximum  gradient.  This  corresponded  to  the  few  minutes  after  the 
animal  had  arisen  to  its  feet  (at  21  minutes)  and  during  which 
shivering  was  most  intense. 

Results  were  similar  for  other  chipmunks  provoked  to  arouse. 
Rapid  increases  in  heart  rates  and  thoracic  temperatures  were  attended 
by  intensive  shivering.  Two  other  animals,  one  male  and  one  female, 
for  which  complete  arousals  were  recorded,  showed  thoracic  temperature 


480 


Indiana  Academy  of  Science 


increases    of   0.75 °C    per    minute    and    0.78°C    per    minute,    respectively, 
during-  the  rapid  phase. 

Figure  2  shows  the  relationship  of  heart  rate  to  thoracic  tempera- 
ture of  the  same  animal.  The  relationship  is  approximately  linear  in  the 
upper  portion  of  the  curve  (36°  to  24 °C).  However,  at  24°  to  22 °C 
there  is  a  turn  toward  curvilinear  which  becomes  more  evident  below 
22 °C.  Extrapolation  of  this  curve  indicates  that  the  heart  should  cease 
to  beat  at  approximately  2°C,  much  lower  than  would  be  predicted 
(approximately  14 °C)  for  an  extrapolation  of  the  upper  part  of  the 
curve.  Such  curves  for  the  other  provoked  chipmunks  indicated  similar 
properties.  Partial  confirmation  of  these  extrapolations  exists  in  the 
results  of  a  hypothermic  cardiac  arrest  induced  in  this  particular  chip- 
munk. Atrio-ventricular  dissociation  occurred  at  1.3°C  with  atrial 
activity  ceasing*  at  0.2°C. 

Figure  3  is  an  example  of  the  results  obtained  during  the  rewarming 
of  "jar  cooled"  animals.  The  general  pattern  is  quite  similar  to  that  of 
provoked  animals.  The  major  exceptions  are  a  more  uniform  rate  of 
rewarming  and  a  longer  time  necessary  for  rewarming  to  a  given  level. 
Thoracic  temperatures  and  heart  rates  increased  less  rapidly  than,  and 
colonic  temperatures  increased  more  rapidly  than,  their  counterparts  in 
animals  arousing  from  natural  torpor.  The  result  was  the  generation  of 
similar  sigmoid  temperature  curves  and  a  smaller  temperature  gradient. 
These  conditions  developed  in  spite  of  shivering  which,  although  not 
beginning  immediately,  was  almost  as  intense  as  and  persisted  longer 
than  that  demonstrated  by  animals  provoked  to  arouse.  Again  a  slight 


40 


3b  - 


32 


28 


24  - 


Q. 

J?     20 


16 


12 


•  ••  THORACIC 
HEART   RATE 

•  ••COLONIC 


2/26/6  9,  INDUCEOM-8/  7/68,  N  R  C  (I) 


440  2 


-360  u 


280  < 


600 


520 


120 


40 


I  0 


20 


50 


60 


70 


30        40 
TIME  (MIN  ) 
Figure  3.     Thoracic    and    colonic    temperature    and    heart    rate    progressions    during    re- 
warming   from    an    induced    hypothermia    in    a    male    chipmunk    (126.8g)    of    the    12-hour 
photoperiod  group. 


Zoology 


481 


560 


z 

400 

5 

to 

< 

VO 

111 

cu 

UJ 

< 

240 

ce 

H 

IX. 
< 

160 

2/26/69, INDUCED    M -  8/ 7/6 8 , NRC  (  I  ) 


5  2 


3  6 


40 


0  4  8  12         16         20        24         I 

TEMPERATURE  (°C) 
Figure  4.     Heart    rate-thoracic    temperature    (°C)    relationships   during    heart    rate    accele- 
ration to  maximum,  for  the  male  chipmunk  shown  rewarming  in  Figure  3. 


"plateau"  was  present  in  the  colonic  temperature  curve  when  the  animal 
rose  to  its  feet. 

A  plot  of  the  heart  rate-thoracic  temperature  relationships  for  this 
animal  (Fig".  4)  indicates  further  differences  between  the  values  obtained 
from  an  animal  provoked  to  arouse  from  natural  torpor  and  those 
obtained  from  an  animal  rewarming  from  an  induced  hypothermia.  The 
reduced  slope  for  the  induced  animal  is  a  reflection  of  comparative  heart 
rates  occurring"  at  higher  temperatures  (approximately  2°C).  However, 
such  differences  may  be  due  to  the  different  hypothermic  states. 

While  the  only  example  of  rewarming  from  induced  hypothermia 
offered  here  is  that  of  a  12-hour  photoperiod  animal,  results  from  fur- 
ther induced  hypothermias  performed  on  both  male  and  female  chip- 
munks housed  under  this  and  two  other  conditions  indicate  a  possible 
difference  among  the  groups.  These  differences,  again,  are  mainly 
reflected  in  the  increased  times  necessary  to  achieve  rewarming  to  a 
given  heart  rate  level,  in  this  case  the  generation  of  maximum  heart 
rates  and  the  thoracic  temperatures  corresponding  to  those  heart  rates. 
Table  1  summarizes  these  data. 

An  evaluation  of  Table  1  reveals  the  animals  under  the  12-hour 
photoperiod  conditions  appear  to  outperform  the  other  groups  of  animals 
in  their  ability  to  rewarm  after  an  induced  hypothermia,  in  both  the 
achievement  of  a  maximum  heart  rate  and  speed  with  which  it  is 
attained.  On  the  other  hand,  chipmunks  housed  in  the  room  with  natural 
illumination  exhibited  the  poorest  performance. 

Discussion 

Confirming  the  previous  reports  about  T.  striatus  (7,  8,  10)  and  its 
western  counterpart,  Eutaynias  (2),  this  investigation  has  also  encoun- 
tered the  unpredictable,  brief,  and  generally  shallow  torpor  experienced 


482  Indiana  Academy  of  Science 

Table   1.     Comparison  of  mean  heart  rates  and  their  corresponding   temperatures   in  the 

different    groups    of    animals.    The    mean    time    required    to    achieve    these    conditions    was 

measured  from   a  mean  heart   rate   and  mean   thoracic   temperature   of    126    and   15.35° C, 

and  108  and  16.9°C,  respectively,  in  provoked  and  induced  chipmunks. 


Provoked  from  _ 

In 

duced  Hypotherm 

ia 

Group 

Natural  Torpor 

12-hr  Light 

Natural  Light 

Roof 

Number 

3 

5 

4 

4 

Mean  max  H.R.  -t-S.D. 

498+44 

521+23 

478+21 

495+25 

Mean  thoracic  temp  +S.D. 

31.0+4.9 

35.8+1.0 

36.9+0.9 

37.0+0.7 

Mean  time  +S.D. 

19.3+6.8 

42.2+7.7 

65.0±13.3 

51.0+16.4 

by  T.  striatus  during  the  winter  months.  It  has  also  shown  that  chip- 
munks do  experience  relatively  deep  torpor.  In  addition,  the  explosive 
nature  of  this  arousal,  as  indicated  by  oxygen-consumption  experiments 
(7),  has  for  the  first  time  been  complemented  by  measurements  of  heart 
rates,  thoracic  and  deep  colonic  temperatures  during  provoked  arousals. 
The  results  for  3  chipmunks,  in  which  complete  measurements  during 
provoked  arousal  have  been  obtained,  indicate  the  ability  to  rapidly 
increase  heart  rates  and  thoracic  temperatures,  reaching  maximum 
heart  rates,  in  less  than  30  minutes.  During  this  period  of  rapid  accelera- 
tion, thoracic  temperatures  increased  at  an  average  rate  of  0.8  °C  per 
minute,  while  colonic  temperatures  lagged  those  of  the  thoracic-heart 
region  by  as  much  as  18.4° C.  However,  thoracic  and  colonic  temperatures 
were  only  as  low  as  11.7°  to  18.7 °C.  The  average  rate  of  thoracic  tem- 
perature acceleration  is  considerably  larger  than  the  0.5  °C  per  minute 
rate  in  rectal  temperatures  observed  by  Cade  (2)  in  a  fasted,  refrig- 
erator-housed Eutamias  amoenus.  The  development  of  large  temperature 
gradients  between  thoracic  and  colonic  regions  indicates  the  physiologi- 
cal ability  of  circulatory  shunting  of  heat  to  critical  organs  (3),  a 
method  of  differential  rewarming  demonstrated  by  all  hibernators 
which  have  been  studied.  The  apparent  "plateaus,"  followed  by  an 
increased  rate  of  increase,  in  the  colonic  temperatures  curves  may 
indicate  a  rapid  vasoconstriction,  followed  by  vasodilation,  of  the  vas- 
culature to  the  posterior  portion  of  the  animal. 

Because  the  majority  of  torpid  animals  experienced  a  hypothermia 
of  a  very  brief  and  shallow  nature,  it  appears  that  a  physiological 
limitation  may  govern  the  depth  of  torpor.  Evidence  for  this  has  been 
presented  by  Lyman  and  Blinks  (4)  who  have  demonstrated  that  the 
ventricles  of  the  isolated  chipmunk  heart  cease  activity  between  7°  and 
5°C,  with  the  atria  stopping  between  3.3°  and  0.7  °C.  These  results  were 
compared  to  those  of  Citellus  trideceml meatus  hearts  which  continued 
beating  as  low  as  — 1°C.  Their  isolated  chipmunk  hearts  also  showed 
a  linear  temperature-heart  rate  relationship  from  30°  to  12 °C,  with  an 
abrupt  curvilinear  relation  below  this  point,  such  that  the  hearts  con- 
tinued to  function  at  a  lower  temperature  than  would  be  predicted  from 
an  extrapolation  of  the  linear  portion  of  the  curve.  The  heart  rate- 
temperature  relationships  presented  here,  while  not  as  linear  as  those 
for  the  isolated  hearts    (4)    do  indicate  that  the  chipmunk  heart  in  the 


Zoology  483 

whole,  intact  animal  also  continues  to  function  at  a  lower  temperature 
than  predicted.  The  finding  here  that  one  heart  ceased  normal  activity  in 
the  whole  animal  at  1.3  °C  is  offered  in  support. 

It  was  hoped  that  further  insight  into  possible  physiological  limita- 
tions, either  in  the  form  of  reduced  hypothermic  tolerance  or  rewarm- 
ing  capabilities,  could  be  obtained  by  subjecting  individuals  to  a  suitable 
hypothermic  stress.  This  was  accomplished  by  "jar  cooling"  in  which 
the  animals  experienced  a  state  of  "artificial  hibernation"  (1).  At  this 
time  it  should  be  emphatically  pointed  out  that  natural  hibernation  and 
induced  hypothermia  are  distinctly  different  (1,  3,  5).  The  most  impor- 
tant factor  characterizing  natural  hibernation  is  the  hibernator's  capa- 
bility of  rewarming  from  this  state  without  the  aid  of  external  heat 
(1,  3,  5).  Animals  under  induced  hypothermia  usually  show  only 
vestiges  of  rewarming  capabilities  (1).  Accordingly,  then,  the  chipmunks 
in  this  investigation  performed  rather  well.  They  showed  both  the 
ability  to  withstand  and  rewarm  from  a  hypothermia  which  would 
have  produced  dire  consequences  to  most  homotherms,  unless  artificially 
rewarmed.  The  fact  that  these  animals  were  capable  of  rewarming 
from  body  temperatures  of  11°  to  17°C,  in  a  relatively  rapid  period  of 
time  and  without  artificial  rewarming  is  significant.  The  differences 
in  rewarming  capabilities  between  the  different  groups  of  animals  seems 
to  suggest  environmental  and  possible  seasonal  influences,  for  which 
studies  are  now  in  progress. 

Literature  Cited 

1.  Adolph,  E.  F.,  and  J.  Richmond.  1955.  Rewarming  from  natural  hibernation  and 
from  artificial  cooling.  J.  Appl.  Physiol.  8  :48-58. 

2.  Cade,  T.  J.  1963.  Observations  on  torpidity  in  captive  chipmunks  of  the  genus 
Eutamias.  Ecology.  44  :255-261. 

3.  Lyman,  C.  P.  1965.  Circulation  in  mammalian  hibernation.  In  W.  F.  Hamilton 
[ed.]  Handbook  of  Physiology,  Section  2,  Circulation  III.  American  Physiological 
Society,  Washington,  D.  C. 

4.  Lyman,  C.  P.,  and  D.  C.  Blinks.  1959.  The  effect  of  temperatures  on  the  isolated 
hearts  of  closely  related  hibernators  and  non-hibernators.  J.  Cell.  Comp.  Physiol. 
54:53-64. 

5.  Lyman,  C.  P.,  and  P.  O.  Chatfield.  1955.  Physiology  of  hibernation  in  mammals. 
Physiol.  Rev.  35  :403-425. 

6.  Lyman,  C.  P.,  and  R.  C.  O'Brien.  1963.  Autonomic  control  of  circulation  during 
the    hibernating    cycle    in    ground    squirrels.    J.    Physiol.    168 :477-499 

7.  Neumann,  R.  L.  1967.  Metabolism  in  the  eastern  chipmunk  (Tamias  striatus)  and 
the  southern  flying  squirrel  (Glaucomys  volans)  during  the  winter  and  summer. 
In  K.  C.  Fisher,  A.  R.  Dase,  C.  P.  Lyman,  E.  Schonbaum,  F.  E.  South,  Jr.  [eds.l 
Mammalian  Hibernation  III.  American  Elsevier  Publishing  Co,  Inc.,  New  York, 
N.  Y. 

8.  Panuska,  J.  A.  1959.  Weight  patterns  and  hibernation  in  Tamias  striatus.  J. 
Mammal.  40  :554-556. 

9.  Simpson,  G.  G.  1945.  The  principles  of  classification  and  a  classification  of  the 
mammals.  Bull.  Amer.  Mus.  Natur.  Hist.  85:1-350. 

10.  Woodward,  A.  E.,  and  J.  M.  Condrin.  1945.  Physiological  studies  on  hibernation  in 
the  chipmunk.  Physiol.  Zool.  18:162-167. 


INSTRUCTIONS  FOR  CONTRIBUTORS 
Eligibility 

Indiana  Academy  of  Science  members  in  good  standing  are  eligible  to  submit 
papers  for  publication  in  the  Proceedings.  When  a  paper  is  signed  by  more  than  one 
author,  at  least  one  must  be  a  member  of  the  Academy.  Preferably,  eligibility  should  be 
established  before  submitting  the  paper,  as  such  papers  are  given  priority.  In  any  case, 
all  authors  must  be  certified  by  the  treasurer  for  payment  of  dues  and  old  reprint  bills 
at  the  time  of  the  deadline  (see  below).  Invited  papers  may  be  considered  for  publication 
regardless   of  the  membership  status   of  the   author. 

All  papers  submitted  for  publication  in  full  will  be  reviewed  by  qualified  reviewers, 
selected  by  the  Publication  Committee.  Papers  read  by  title  only  at  the  Fall  Meeting 
may  also  be  considered  for  publication.  Among  papers  of  primarily  regional  interest, 
e.g.,  in  certain  aspects  of  botany,  zoology,  geology,  geography,  and  anthropology,  those 
dealing  with  Indiana  material  will  be  accorded  preference.  The  selection  of  papers  for 
the  Proceedings  is  the  responsibility  of  the   Publication   Committee. 

Abstracts 

Three  copies  of  an  abstract  should  be  submitted  to  the  Divisional  Chairman  at  the 
time  the  title  of  a  paper  is  submitted  for  the  Fall  program.  All  abstracts  will  be  pub- 
lished in  the  Proceedings,  either  separately  or  with  papers  that  are  published  in  full. 
Two  copies  of  the  abstract  should  be  marked  "for  the  editor."  The  third  copy  of  the 
abstract  should  be  marked  "for  the  divisional  chairman"  and  may  include  information 
about  time,  projection  facilities  needed,  etc.  The  abstract  should  be  prepared  according 
to  the  form  currently  used  in  the  Proceedings  (see  the  latest  copy  of  the  Proceedings). 
The  abstract  should  be  complete  and  clear  in  itself  not  over  5%  of  the  length  of  the 
paper.  Normally  abstracts  should  not  exceed  200  words  in  length.  Abstracts  are  not 
reprinted    (except  for  those  which  are  included  at  the  head  of  a  paper  published   in  full). 

Deadline  at  the  Editorial  Office 

When  sent  via  the  Divisional  Chairman  as  prescribed,  or  directly,  all  material  to  be 
considered  for  publication  in  the  Proceedings  must  reach  the  editor  within  20  days 
following  the  Fall  Meeting. 

Preparation  of  Manuscripts 

A.  Refer  to  the  latest  copy  of  the  Proceedings  for  the  accepted  style  of  abstracts  and 
papers,  and  follow  this,  especially  in  literature  citations,  headings,  footnotes,  table 
and  figure  construction. 

B.  Type  on  11  x  8%  inch  bond  paper  with  a  new  ribbon,  leaving  some  margin.  Double 
space  everything,  including  title,  author's  name,  department  and  institution,  foot- 
notes, quotations,  legends  and  literature  list.  Manuscripts  must  be  submitted  in 
duplicate.  The  original  will  become  the  printer's  copy ;  if  it  must  be  retyped,  it  will 
be  sent  back  to  the  author  for  this. 


(', 


Footnotes  are  to  be  kept  to  a  minimum.  Necessary  footnotes  are  numbered  consecu- 
tively   throughout,    and   referred   to   in    the   text   as   superscripts,    without    parentheses. 

Literature  citations  are  listed  alphabetically  at  the  end  of  the  paper,  headed  Litera- 
ture Cited.  List  complete  literature  citations,  i.e.,  author,  date,  title,  journal  (or 
publisher  and  city),  volume  and  total  pages.  The  highly  abbreviated  form  used  in 
some    journals    has    not    been    adopted    for    the    Proceedings.     Follow    these    models : 

7.  Doe,  J.  B.  and  R.  C.  Roe.  1949.  New  light  from  old  radioactive  carbon.  J.  Amer. 
Biol.  Soc.  34  :273-305. 

8.  Milazzo,  G.  1963.  Electrochemistry.  Elsevier  Publ.  Co.,  New  York,  N.Y.  708  p. 
References  cited  should  be  numbered  consecutively  (in  the  alphabetized  list)  and 
should  be  referred  to  in  the  text  by  number  in  parentheses  on  the  line  of  type  and 
before  the  period  if  at  the  end  of  a  sentence. 

484 


Instructions  for  Contributors  485 

E.  Do  not  underline  anything  except  scientific  names,  words  to  be  italicized,  and  titles 
of    books    when    they    appear    in    the    text    only,    not    in    literature    list. 

F.  All   literature   listed,   tables   and   illustrations   should   be   referred   to   in   the   text. 

G.  Tables,  which  are  costly  to  print,  should  be  reduced  to  a  minimum.  Avoid  small 
tables  scattered  through  the  text.  Each  table  (including  heading)  should  be  typed 
on  a  separate  letter-size  sheet  and  placed  at  the  end  of  the  paper.  Outsize  tables 
cannot  be  accepted. 

H.  Photographs  should  be  printed  on  glossy  paper  and  have  good  contrast.  It  is  best 
to  mount  them  trimmed  to  fit  tightly  together  at  the  edges  in  groups,  on  stiff  card- 
board with  rubber  cement.  Proportion  the  group  for  a  full  page  of  the  Proceedings, 
or  use  the  full  width  of  the  paper  (4%")  and  any  part  of  the  page's  height.  Do 
not  mix  line  drawings  and  photographs  in  the  same  group.  All  figure  captions 
should  be  on  a  single  letter-size  sheet,  numbered  to  correspond  and  placed  at  end 
of  paper. 

I.  The  originals  for  line  drawings  need  be  no  more  than  twice  the  size  desired  for  the 
printed  figure.  They  should  be  proportioned  and  arranged  to  fit  the  page  size  of  the 
Proceedings.  All  line  drawings  must  be  drawn  in  India  ink,  lettered  with  a  lettering 
set,  and  of  suitable  size  to  allow  for  necessary  reduction.  Do  not  submit  printed 
maps  when  the  necessary  reduction  will  efface  the  narrower  lines  or  render  some  of 
the  lettering  hardly  legible ;  such  maps  should  be  redrawn  and  lettered  in  adequate 
size  letters,  omitting  unnecessary  details.  All  illustrations  requiring  a  size  scale 
must   portray   the   scale    in    a   manner   that   permits   size    reduction. 

J.  Major  professors  are  urged  to  review  all  papers  by  their  graduate  students,  for  both 
form  and  content,  before  they  are  sent  in  for  publication.  Of  those  based  on  uni- 
versity theses,  manuscripts  carrying  the  approval  by  the  professor  will  be  given 
preference  over  those  without  such  approval.  New  authors,  especially,  are  reminded 
that  a  scientific  paper  should  summarize  the  work,  not  recapitulate  it.  It  must  be 
much  more  concise  than  a  university  thesis,  avoiding  all  extraneous  material, 
especially  long  tables  and  lists  of  little  interest  except  to  the  author.  All  manuscripts 
should  be  as  concisely  written  as  possible. 

K.  Reprints  of  papers  are  paid  for  by  authors,  at  cost.  Directions  for  ordering  re- 
prints accompany  the  galley  proof  and  the  orders  are  placed  at  the  time  the  author 
returns  the  corrected  galley  proof  to  the  editor.  The  order  form  supplied  by  the 
editor  must  be  completed  and  returned.  If  you  have  any  special  institutional  forms 
regarding  payment  for  the  reprints,  these  should  be  sent  directly  to  the  Treasurer 
of  the  Indiana  Academy  of  Science  at  the  time  the  reprints  are  paid  for.  Abstracts 
are  not  reprinted. 

L.  The  editor  needs,  at  the  time  he  mails  out  galley,  current  addresses  for  all  authors 
and  coauthors  of  all  abstracts  and  papers.  Many  former  graduate  students  lose  the 
opportunity  to  order  reprints  when  there  are  faulty  forwarding  addresses.  It  is 
suggested  that  the  student's  permanent  home  address  be  written  on  the  reverse  side 
of  the  abstract  copy  marked  "for  the  editor." 
Revised  July  14,  1970. 


INDEX 


Acritarchs  in  New  Albany  Shale,  254 
Acrylamide     gel,     use     in    staining     nucleic 

acids,  348 
Adams,  W.  H.,  65 
Aedes  and  soils,  238 
Aedes,  distribution  of,  Indiana,  238 
Agee,  E.  M.,  299 
Ahlrichs,  J.  L.,  432 
Allee  Woods,  Collembola  in,  234 
Aminoglutethimide,  effects  of,  455 
Aminogluthethimide,    effects   on    rat   ovaries 

and  uteri,  439 
Anderson,  R.  O.,  135 
Archaeology,  Mounds  State  Park,  75 
Archaeology,    Oxendine    Site,    Vigo    County, 

57 
Archaeology,  State  of  Delaware,  69 
Arnett,  Patricia  M.,  234 
Asia,  urbanization,  253 
Aspergillus     niger,     biosynthesis     of     fatty 

acids,  351 
Avidin,    effect    on    fatty    acid    biosynthesis, 

351 
Awasthi,  Y.  C,  110 


Bacteria  in  farm  pond  waters,  423 

Bacteria,  sulfur  and  iron  oxidizing,  and 
coal  mine  stream  pollution,   345 

Bacterial  ingestion,  European  corn  borer, 
effect  on  heartbeat,  227 

Bakker,  G.  R.,  122 

Barton,  T.  F.,  318 

Baumgardner,  M.  F.,  413 

Beech  ferns,  taxonomy  of,  388 

Beesley,  L.  and  Adele,  83 

Behavior,  of  Indiana  ruffed  grouse,  177 

Bennett,  A.  S.,  351 

Big  Brown  Bat  (Eptesicus  fuscus),  move- 
ments, 439 

Big  trefoil  and  tall  fescue,  193 

Big  trefoil,  ecotypes  in  southern  Indiana, 
193 

Bingham,  R.  L.,  205 

Single,  G.,  92 

Bioassays,  need  for  longer  term  tests,  148 

Biological  Survey  Committee  Report,  23 

Bison,  bones  from  Indiana  cave,  472 

Blair,  P.  V.,  93 

Blair,  B.  O.,  83 

Bloom,  W.  W.,  83 

Bluegill  growth,  effect  of  photoperiod,  135 

Bluegill,  response  to  predation,  139 

Bluegill,  seasonal  growth,  135 

Boneham,  R.  F.,  254 

Bounty,  fox  in  Indiana,  187 

Brechner,  R.  E.,  449 

Buckbee,  Sister  Barbara,  123 


Bullard,  R.  W.,  476 
Burkle,  M.  A.,  357 

Cantin,  A.  J.,  309 

Carlton,  W.  W.,  91 

Carotenoid,    mutant    of    Cyanidium    caldari- 

um,  83 
Carr,  Merrill  T.,  memorial,  27 
Cathartes,  bones  from  Indiana  cave,  472 
Cave,    Thundermug    Bone,    Monroe    County, 

Indiana,  472 
Cavern  development,  281 
Central  places,  hierarchy  of,  325 
Chalybion  zimmermanni  in  Indiana,  231 
Chandler,  L.  228,  229 
Charophytes,  pleistocene,  84 
Check  list,  Indiana  Collembola,  249 
Cheetham,  R.  D.,  107 
Cherokee  past,  evidence  for,  57 
Christaller,  W.,  325 
Chromosome  associations,  84 
Chromosome  numbers  in  Polygonus,  396 
Chuang,  T.  F.,  110 
Cities,  near-neighbor  analysis,  325 
Coal  mining,  263 
Coal,    stream    pollution    from    mine    waste, 

345 
Coelioxys      obtusiventris      Crawford,      from 

Tippecanoe  County,  Indiana,  228 
Coffing,  S.  J.,  57 
Coffman,  D.  M.,  333 
Cole,  T.  A.,  348 

Collembola,  Indiana  records,  249 
Collembola,  Indiana  species,  249 
Collembola,  key  to,  234 
Cook,  D.  J.,  122 

Cooling  degree  days  in  Indiana,  292 
Coordination     compounds,     infrared    spectra 

of,  121 
Cope,  J.  B.,  439,  466,  470 
Corn    monoploids,    chromosome    associations 

in,  84 
Corn  roots,  distribution  in  soils,  401 
Cotter,  D.  A.,  345 
Crane,  F.  L.,  110 
Crankshaw,  W.  B.,  137 
Cyanidium  caldarium,   a  carotenoid  mutant, 

83 
Cycloheximide,     blockage     of     Dictyostelium 

discoidcum  myxamoeba  release,  345 
Cyclotron       resonance,       theory,       non-local 

terms,  360 
Cytochrome  oxidase,  membrane  form,  110 

Daily,  Fay  K.,  27,  84 

Daniell     cell,     temperature     dependence     of 
voltage,  123 


487 


488 


Index 


Daughtery-Monroe  Site,  excavations  at,  57 

Davidson,  P.  G.,  135 

Deay,  Howard  O.,  memorial,  27 

Degree  days,  cooling,  in  Indiana,  492 

Delaware  Indian  tribe,  60,  69 

Di-n-butyloxamidine,  complexes  of,  129 

Dilcher,  D.,  375 

Dinkel,  R.  M.,  309 

DNA,    microspectrophotometric   analysis   of, 

84 
Dolan,  E.,  57 

Drainage  maps  from  topographic  maps,  333 
Duroic     acid,     decarboxylation     mechanism, 

122 

Ecology,  of  Indiana  ruffed  grouse,  177 
Ecuador,  cultivated  Solanaceae,  376 
Education     and    undergraduate     oceonogra- 

phy,  359 
Endoplasmic  reticulum,  nucleotide  phospha- 
tase activities,  107 
Endothelial  cell  nuclei,  brain,  93 
Endrin,  use  as  a  fish  toxicant,  148 
Enteric  epithelium,  human,  92 
Environment,  preservation  of,  49 
Enzyme-catalyzed  reactions,  346 
Enzmes,  L-amino  acid  oxidase,  121 
Epithelial  tumor,  94 
Euphorbiaceous  fruits,  Eocene,  375 
European  corn  borer  control,  heartbeat  and 

bacterial  pathogens,  227 
Eversole,  W.  J.,  455 

Extinctions,  megafauna  of  late-Pleistocene, 
65 

Farm  ponds,  423 

Fatty  acids,  avidin  effects  and  biosynthesis, 
351 

Fauna,  Maryland  Miocene,  253 

Feeding  frequency,  effect  on  bluegill 
growth,  136 

Ferguson,  R.  J.,  58 

Fisher,  D.  D.,  346 

Fluvial  morphology,  parameter  measure- 
ment, 333 

Foley  Woods,  Edgar  County,  Illinois,  137 

Follicle  stimulating  hormone,  effect  of  ster- 
oids on  chicken  pituitary  content,   462 

Foods,  Peromyscus  leucopus,  172 

Foods,  white-footed  mouse,  172 

Ford,  L.,  84 

Forest  dominance  expressions,  137 

Forest  types,  Indiana,  198 

Fossil,  fruits  of  Eocene  age,  375 

Fox,  bounty  in  Indiana,  187 

Franklin  County,  Indiana,  Trilliums  of,  83 

Fruits,  Euphorbiaceous  of  Eocene  age,  375 

Fungi,   dispersed  fossil   spoies,   375 

Freeman,  Leslie  Willard,  memorial,  29 

C.amma-A  globulin,  92 
Gammon,  J.  R.,  136 


Geology,  coal  roof  rock,  263 

George,  J.,  121 

Germination,  in  Dictyostelium  discoideum, 
345 

Glasses,  oxide,  produced  by  Ir,  Pd,  Rh  and 
Ru,  361 

Coins,  D.  R.,  137 

Golgi  apparatus,  mucleotide  phosphatase 
activities,  107 

Gramineae,  stem  structure  in,  85 

Grasses,  stem  structure  in,  85 

"Great  Mound,"  excavation  of,  75 

Green  sunfish,  growth  and  effect  of  hier- 
archy, 136 

Grocer  locations,  Terre  Haute,  Indiana,  309 

Groundwater  in  limestones,  281 

Growth,  bluegill,  effect  of  photoperiod,  135 

Growth,  bluegill,  effected  by  feeding  fre- 
quency, 136 

Growth,  bluegill,  seasonal,  135 

Gulish,  W.  J.,  139 


Hadley  Lake  depiession,  Tippecanoe  Coun- 
ty, 270 

Hall,  B.  V.,  91 

Hamilton,  J.  A.,  357 

Hansen,  U.  J.,  360 

Hart,  J.  W.,  249 

Hart,  R.  D.,  137 

Hazelton,  J.  L.,  360 

Heated  effluents,  fish  response  to,  136 

Heath,  M.  E.,  193 

Heiser,  C.  B.,  Jr.,  376 

Hendricks,  D.  R.,  466,  470 

Hettmer,  J.  H.,  359 

Hierarchy,  effect  on  green  sunfish  growth, 
136 

Highwood,  Joyce  E.,  396 

Hlavaty,  Vaclav,  memorial,  30 

Hoffman,  W.  E.,  129 

Hookeriaceae,  species  and  distribution  in 
Africa,  Europe,  Asia,  Australia  and 
Oceania,  377 

Hopewell  Indians  of  Illinois,  62 

Houlihan,  J.  F.,  358 

Hughes,  L.  B.,  423 

Hunter,  K.  B.,  62 

Hydrocyanic  acid,  content  in  Manioc,  137 

Hypothermia  and  Tamias  striatus,  476 


Indian   territory,  misinterpreted  as  Indiana, 

229 
Indiana,  early  man,  65 
Insect    heartbeat    and    bacterial    pathogens, 

227 
Insect  pathology  and  heartbeat,  227 
Insecticide  adsorption  on  soils,  432 
Ion    source,    polarized    helium-3,    at    Indiana 

University,  359 
Iron,     oxidizing     bacteria,     and     coal     mine 

stream  pollution,  345 


Index 


48!) 


Ishikawa,  S.,  93 

Isotope  effects,  deuterium,  121 

Jackson,  M.  T.,  137 
Jacobs,  M.,  129 

Job's    method    and    transition    metal    com- 
plexes, 129 
Johannsen,  C.  J.,  413 
Johansen,  N.  I.,  270 
Jones,  G.  S.,  172 
Jordan,  Ruth,  memorial,  31 
Junior  Academy  of  Science,  16 

'Kaiser'     trefoil,     ecotypes     from     Crawford 

County,  Indiana,  193 
Karst  groundwater  zone,  281 
Kennepohl,  G.,  129 
Kidney   microvilli,   ultrastructure   and  enzy- 

mology,  93 
Kindig,  R.,  345 
Kirkpatrick,  C.  M.,  177 
Kirkpatrick,  R.  D.,  187,  449 
Kristof,  S.  J.,  413 
Krumholz,  L.  A.,  205 

Lambert,  N.,  375 

La  Motte  Culture,  site  of,  57 

Lee,  S.  S.,  346 

Lciosphacridia  in  New  Albany  shale,  254 

Lembi,  C.  A.,  96 

Limnology,  experimental,  in  a  western  In- 
diana lake,  359 

Lindsey,  A.  A.,  198 

Lipid  in  membrane  structure,  110 

Liquid  junction  potentials,  Daniell  cell,  123 

Llewellyn,  R.  A.,  359 

Lotus  pcdunculatus  Cav.,  formerly  L.  uli- 
ffinosus  Schkuhr.,  L.  major  Sm.,  193 

McComish,  T.  S.,  135,  136 

McMichael,  E.  V.,  57 

McReynolds,  H.  E.,  148 

Magneli  phases  and  electron  spin  reso- 
nance, 358 

Mangum,  T.  E.,  Ill,  136 

Manihot  esculenta,  influence  of  microcli- 
mates, 137 

Manioc,  influence  of  microclimates,  137 

Mannering,  J.  V.,  407 

Marsh   ferns,   taxonomy  of,    388 

Masilea,  rhizoid  formation  by,  83 

Melhorn,  W.  N.,  270 

Membrane,  mosaic  structure,  110 

Mertens,  T.  R.,  396 

Metal  oxides  and  electron  spin  resonance, 
358 

Metz,  C.  R.,  123 

Meyer,  E.  R.,  205 

Meyers,  N.  L.,  432 

Microclimate,  effect  on  acid  content  of 
Manioc,  137 


Middendorf,  W.  F.,  348 

Mills,  R.,  439 

Mitochondria,   ADP   induced   changes   in,    93 

Molecular  complexes,  bromine  and  substi- 
tuted carbostyrils,  122 

Molt  in  Mus,  449 

Monroe  County,  Indiana,  ruffed  grouse  ecol- 
ogy and  behavior  in,  177 

Moore,  D.  N.,  396 

Morre,  D.  J.,  96,  107 

Mosquitoes,  Indiana  distribution,  238 

Mounds  State  Park,  excavations,  75 

Muehrcke,  J.  P.,  177 

Mueller,  W.  D.,  357 

Mulay,  L.  N.(  358 

Murad,  T.  A.,  69 

Mus,  molt  in  two  populations,  449 

Mus  musculus,  parasites  in,  441 

Mussels,  valuable,  Wabash  and  White  Riv- 
ers, 205 

Myelination,  deficient,  quaking  mouse,  92 

Myotis  lucifugus,  status  in  Indiana,  470 

Natural  resources,  outlook,  49 
Nerve  fibers,  axoplasmic  transport  in,  346 
Neumann,  G.  K.,  60,  62,  69 
Neutron  generator,  flux  determination,  357 
New  Albany  Shale,  acritarchs,  254 
Nichols,  K.  E.,  83 
Nowak,  J.,  121 

Nuclear  emulsion,  ionization  in,  357 
Nucleic    acids,    acrylamide    gel    staining    of, 
348 

Ochs,  S.,  346 
Oliver,  Jeanette  C,  388 
Ong,  L.  G.,  121 

Organic  carbon  in  farm  pond  waters,  423 
Ostracods,   survey   in   Delaware   County,   In- 
diana, 137 

Pace,  R.  E.,  57 

Page,  D.  S.,  121 

Paleo-Indian,  cause  for  megafauna  extinc- 
tions, 65 

Paleozoic  rocks,  southern  Indiana,  254 

Parasites,  Mus  musculus,  441 

Parrot,  D.  W.,  129 

Patton,  J.  B.,  49 

Pcromyscus  leucopus.  Pike  County,  Indi- 
ana, 172 

Pesticide,  toxicity  of  endrin  to  fish,  148 

Petty,  R.  O.,  137 

pH,  effects  on  enzyme  kinetics,  121 

Phenology  in  Indiana,  83 

Photoperiod,  effect  on  bluegill  growth,  135 

Pinkerton,  J.  M.  H.,  357 

Plasma  membrane,  nucleotide  phosphatase 
activities,  107 

Plasma  membranes,  plant  stems  and  rat 
liver,  96 


490 


Index 


Pleistocene,  faunal  extinction  causes,  65 
Pollution,  and  mosquitoes,  Indiana,  238 
Pollution,  avoidance,  abatement,  49 
Pollution,  stream,  from  coal  mines,  345 
Polygonum,  cytotaxonomic  notes  on,  396 
Pond  sediment  mineralogy,  432 
Population    decline,    Southwestern    Indiana, 

318 
Population,  fix  in  Indiana,  187 
Powell,  R.  L.,  281 
Predation,  effect  on  bluegill,  139 
Proglacial    drainage,    Hadley    Lake    depres- 
sion, 270 

Quaking  mouse,  deficient  myelination,  92 
Quarter  method,  testing,  138 


Racial   history,   Indians  of  eastern  U.S.,   60, 

62,  69 
Radiotelemetry  and   Big   Brown   Bat    (Epte- 

sicus  fuscus),  466 
Ramaley,  R.,  345 
Ranish,  N.,  346 
Rathkamp,  W.  R.,  462 
Reed,  P.,  129 

Relativistic  thermodynamics,  358 
Reuszer,  H.  W.,  423 
Reynolds,  L.  M.,  357 

Rhizoids,  auxin  effects  on  formation  of,  83 
Richards,  R.  L.,  472 
Rogers,  Richard  M.,  memorial,  32 
Roland,  J.  C,  96 
Ruffed     grouse,     ecology     and     behavior,     in 

Monroe  County,  Indiana,  177 

Sabri,  M.  I.,  346 

Salvage  of  waste,  decline  in,  49 

Sartain,  C.  C,  361 

Schaal,  L.  A.,  292 

Schaffer,  R.  E.,  476 

Schmedtje,  J.  F.,  92 

Schmelz,  D.  V.,  138 

Schulz,  A.  R.,  346 

Schwartz,  E.,  121 

Schwenk,  K.,  351 

Seely,  O.,  357 

Semiconductors  produced  by  Ir,   Pd,  Rh  and 

Ru,  361 
Settlement    decline,    Southwestern     Indiana, 

population,  318 
Sheffy,  M.  V.,  375 
Shelley,  R.  L(avere),  memorial,  33 
Siakotos,  A.  N.,  93 
Siddiqi,  A.  H.,  253 
Simon,  J.,  91 
Siverly,  R.  E.,  238 

Skeletal  muscle,  regeneration  in  rabbits,  91 
Snow,  J.  T.,   122 

Soil,  corn  root  distribution  in,  401 
Soil,  multispectral  properties,  413 
Soil,  organic  matter,  413 


Soil  sampling  depth  and  corn  root  distribu- 
tion, 401 

Soils  and  trees,  198 

Soils,  effect  of  rainfall  energy  on,  407 

Soils,  surface  sealing  by  rain,  407 

Soils,  water  infiltration  into,  407 

Solanaceae,  cultivated  of  Ecuador,  376 

Solar  radiation  and  cooling  degree  days, 
292 

Southwestern  Indiana,  population  and  set- 
tlement decline,  318 

Spores,  fungal,  in  Eocene  deposits,  375 

Statton,  C.  T.,  253 

Steinert,  D.  L.,  358 

Steinrauf,  L.  K.,  357 

Stem  structure  in  grasses,  85 

Steroids,  effect  on  chicken  pituitary  FSH 
content,  462 

Stivers,  R.  K.,  401 

Stout,  T.  R.,  129 

Streams,  accuracy  of  ordering  procedures, 
333 

Sulfur,  oxidizing  bacteria  and  coal  mine 
stream  pollution,  345 

Sundy,  J.,  129 

Surdzial,  R.  E.,  123 


Tall  fescue  and  big  trefoil,  193 

Tamias    striatus,    torpor    and    hypothermia, 

476 
Telfair,  W.  B.,  466 

Temperature  and  cooling  degree  days,  292 
Thermal  induction,  bacteriaphage  PI,  346 
Thermodynamic  properties,  Daniell  cell,   123 
Thermodynamics,  relativistic,  358 
Thiouracil,  thyroid  response  to,  91 
Thyroxine,  molecular  structure  of,  357 
Tin,  hydrolysis  constant,  121 
Tippecanoe    County,    Hadley    Lake    depres- 
sion, 270 
Titanium-oxygen    system   and   electron    spin 

resonance,  358 
Toms,   William  Lowell,  memorial,  34 
Topographic  maps  and  stream  nets,  333 
Tornadoes,     Indiana    climatology    and    sta- 
tistics, 299 
Torpor,    rewarming    from    in    Tamias    sti-ia- 

tus,  476 
Transition  metals,  di-n-butyloxamidine  com- 
plexes, 129 
Transitions,     in     transistion     metal     oxides, 

358 
Trees  and  soils,  198 

Trilliums,  Franklin  County,  Indiana,  83 
Tropistcrneis     collorus     (Castelnau),     inter- 
breeding of,  227 


Urbanization    and    Indiana    mosquito    popu- 
lations, 238 
Ursus,  bones  from  Indiana  cave,  472 


Index 


491 


VanEtten,  R.  L.,  121 

Van  Nuys  Site,  continued  excavation,  58 

Van  Vleet,  J.  F.,  91 

Vickery,  K.  D.,  75 

Vitamin  E  deficiency,  rabbits,  91 


Wabash  River,  proglacial  drainage,  270 

Ware,  Mildred  G.,  227 

Ward,  Gertrude  L.,  231 

Warner  Glen  W(ones),  memorial,  35 

Watanabe,  I.,  92 

Water,  ethical  implications,  49 

Watershed  soil  mineralogy,  432 

Weatherwax,  P.,  85 

Weber,  N.  E.,  93 

Weber,  N.  V.,  325 

Weeks,  H.  P.,  Jr.,  162 

Welch,  Winona  H.,  377 

Welser,  J.  R.,  91 

Whitaker,  J.  O.,  Jr.,  441 

White,  J.  L.,  432 


White-footed  mouse,    Pike   County,    Indians 

172 
Whitten,  J.  B.,  Jr.,  94 
Whittle,  E.,  439 
Wier,  C.  E.,  263 
Wiersma,  D.,  407 
Williams,  R.,  121 
Wilsey,  Ruth  A.,  136 
Woodcock,  territorial  behavior,  162 
Woodcocks,  courtship  behavior,  162 
Wright,  Howard  F.,  memorial,  37 

Yemma,  J.,  84 

Young,  F.  N.,  227 

Youse,  H.  R.,  presidential  address,  45 

Yumoto,  S.,  93 

Zachary,  A.,  413 
Zeller,  F.  J.,  462 
Zimmack,  H.  L.,  227 
Zimmerman,  R.  E.,  455