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DNA POLYMERASE

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Volume XXVIII, Nos. 1-4

March, 1977

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SECTION I

MATHEMATICAL SCIENCES

Mathematics, Statistics,
Operations Research

SECTION IV
BIOLOGICAL SCIENCES
Agriculture, Botany,
Medical Science,
Zoology

SECTION XI
EORENSIC SCIENCES

SECTION II
PHYSICS

SECTION X
AQUATIC SCIENCES

SECTION III
EARTH SCIENCES
Geography
Geology

SECTION IX
COMPUTER SCIENCES

SECTION V
SOCIAL SCIENCES
Anthropology, Education,
Economics, History,
Psychology, Sociology

SECTION VI
ENVIRONMENTAL
SCIENCES

SECTION VIII
SCIENCE EDUCATION

SECTION VII
CHEMISTRY

AFFILIATED ORGANIZATIONS
Texas Section, American Association of Physics Teachers
Texas Section, Mathematical Association of America
Texas Section, National Association of Geology Teachers

GENERAL INFORMATION

MEMBERSHIP. Any person engaged in scientific work or interested in the promotion of
science is eligible for membership in The Texas Academy of Science. Dues for annual
members are $12.00; student members, $5.00; sustaining members, at least $10.00 in
addition to annual dues; life members, at least $240.00 in one payment; patrons, at least
$500.00 in one payment; corporation members, $250.00 annually; corporation life
members, $2000.00 in one payment. Annual subscription rate is $45.00. Dues should be
sent to the Secretary -Treasurer. Subscription payments should be sent to the Managing
Editor.

Texas Journal of Science. The Journal is a quarterly publication of The Texas Acad¬
emy of Science and is sent to all members and subscribers. Single copies may be pur¬
chased from the Managing Editor.

Manuscripts submitted for publication in the Journal should be sent to the
Manuscript Editor, P.O.Box 13066, North Texas State University, Denton, Texas 76203.

Published quarterly by The Talley Press, San Angelo, Texas, U.S.A. (Second Class
Postage paid at Post Office, San Angelo, Texas 76901.) Please send 3579 and returned
copies to the Editor (P.O.Box 10979, Angelo State University, San Angelo, Texas 76901.)

Volume XXVIII, Nos. 1-4

March, 1977

CONTENTS

Editorially Speaking . . . . 3

Instructions to Authors . . . . 3

Invited Paper

Excision Repair Following Ultraviolet Irradiation in Toluene-Treated Escherichia

coli. By John W. Dor son and Robb E. Moses . 5

Markov- Renewal Programming Under Incomplete Information -I. By J.S. Prasad

and A. G. Walvekar . . . . . . 19

Markov-Renewal Programming Under Incomplete Information - II. By A. G. Walvekar

and J. S. Prasad . . . . . . 33

Some Comments Concerning the Impact Ball Apparatus. By C. T. Haywood ...... 45

Petrography of Pleistocene Volcanic Ash from Palo Duro Canyon, Texas. By

Peter Jeschof nig . . . 51

Application of Huygens’ Principle to the Reflection of Seismic Waves at a Free Surface.

By Ethel Ward McLemore and Ira L. Wright . . . . 61

The Effect of Lysosomal Enzymes onChloroplasts in vitro. By J. Aune, Barbara Jean

Smith, and M. M. Aboul-Ela . . . . 85

A Simple, Inexpensive Freeze-Etch Device for Use in Electron Microscope Labora¬
tories. By G. D. Cagle and G. R. Vela . . . . 91

The Flower Gardens Coral Reefs. By M. Samuel Cannon and Edmond S. Alexander . . 99

A Reevaluation of Two New Species of Fossil Bats from Inner Space Caverns. By

Sara L. Dorsey . . . 103

Seasonal Distribution of Mites Associated with Popilius disjunctus (Illiger)(Coleop-

tera;Passalidae) in Hardin County , Texas. By George E. Gibson, Jr . . 109

Crown Positions Within Unthinned Loblolly Pine Plantation Canopies. By J. David

Lenhart, Chi-Yun Ho, and Dwight R. Hicks . . . . . . . 113

Production of Amino Acids by Soil Microorganisms. By F. R uiz -Berraquero and

A. Ramos- Cormenzana . . . . . . 1T9

The Bats of East Texas. By David J. Schmidly, Kenneth T. Wilkins, Rodney L.

Honeycutt, and Bernard C. Weynand . . . 127

A Key to the Lice of Man and Domestic Animals. By Donald W. Tuff . 145

An Index to the Genera of Hosts in YamagutVs SystemaHelminthum. By D. W. Tuff

and D. G. Huffman . . ....161

Effects of Prometryne on RNA and Protein Metabolism in Bean Leaf Discs. By

David C. Whitenberg and James H. Flippen . . . 193

Results of a Magnetometer Survey at Hueco Tanks, A Prehistoric Mogollon Village

in Western Texas. By J. Barto Arnold III and George B. Kegley III . . . 201

A Sodium Iodide Centrifugation Technique for Isolation of Newly -Replica ted DNA

from Rat Liver. By Roger F. Brown . . . 209

Barriers to Internal Rotation in 2-Bromopropene and 2-Iodopropene. By G. A. Crowder

and Royce W. Waltrip . . . 219

The Polarographic Reduction of Metal Ions in Acetone and Acetophenone in the
Presence of Lithium Perchlorate and Tetraethylammonium Perchlorate. By
Ivory V. Nelson . . . . . 223

Effects of Simple Sugars on Certain Serum Constituents and Erythrocyte Osmotic

Fragility in Rats. By M. E. Purdom, M. Bayer, S. J. Norton, and P. T. Sullivan ... 231

Rayleigh-Schroedinger Perturbation Calculations for Beryllium. By G. P. Saxon

and D. L. Hardcastle . . . 239

Synthesis of a Radiolabeled Carrier to Precipitate Anti-Hapten Antibodies. By

A. C. Schram and C. P. Christenson . . . . 253

Corrosion Studies in a Water Flood Oil Field System. By Larry G. Spears . 259

The Salinity and Temperature Tolerance and the Growth of Macrobrachium ohione

(Smith) 1874 Reared in Laboratory Tanks. By Kyung-Suk Chung . . 271

The Ecology of a B-ranchiobdellid (Annelida : Oligochaeta) from a Texas Pond. By

Stephen J. Koepp and Edgar A. Schlueter . . . . . 285

Limnological Theses and Dissertations Concerning Texas Waters, 1897-1976. By

John M. Pettitt . . . . . 295

DIALECTIC

Is Canopy Interception an Accurate Measure of Loss from the Hydrologic Budget?

By Mingteh Chang . . . . . 339

Critical Levels ofPhosphorus and Nitrogen in Texas Impoundments. By G. Fred Lee . . . 347

Wolves, Coyotes, Ducks, and Hybridism. By Bob H. Slaughter . . 351

NOTES SECTION

Procoelus Versus Opisthocoelus Vertebrae. By William D. Gosnold and Bob H. Slaughter . . 355

A Noteworthy Record of the Silver-Haired Bat in Southeast Texas. By Chester O. Martin ... 35 6

Occurrence of Striped Bass ( Morone saxatilis ) in Coastal Waters of Texas. By R. L.

Benefield, A. W. Moffett, and L. W. McEachron . 357

A Survey of Ectoparasites of Hares and Rabbits in Grant County, New Mexico. By

Paul H. Rodriguez . . 358

A Kiam Incised Pot from Louisiana. By Russell J. Long . 358

Resolution of 2-(o-Chlorophenyl)-2-(p-Chlorophenyl)Acetic Acid. By J. Guilford,

E. Hickman, and D. Ghosh . . . . . 360

Observations on the Ecology of Micropterus treculi in the Guadalupe River. By R. L.

Boyer, G. W. Luker, and R. J. Tafanelli . . . . 361

An Ambicolorate Windowpane ( Scophthalmus aquosus), with Notes on Other

Anomalous f latfish. By Andre M. Landry , Jr., and Kenneth W. Johnson ...... 362

Analysis of Stump Vegetation in an East Texas Pond. By E. S. Nixon, W. F. Trotty,

B. L. Bates, and D. L. Wilkinson . 366

INDEX, Volume XXVIII - 1977 ............................... 368

INSTRUCTIONS TO AUTHORS

Papers intended for publication in The Texas Journal of Science are to be sub¬
mitted to Dr. Roland Vela, Editor, P. O. Box 13066, North Texas State University,
Denton, Texas 76203.

The manuscript submitted is not to have been published elsewhere. Triplicate
typewritten copies (the original and 2 reproduced copies) MUST be submitted.
Typing of both text and references should be DOUBLE-SPACED with 2-3 cm
margins on STANDARD SYl X 11 typing paper. The title of the article should be
followed by the name and business or institutional address of the author(s). BE
SURE TO INCLUDE ZIP CODE with the address. If the paper has been
presented at a meeting, a footnote giving the name of the society, date , and occasion
should be included but should not be numbered. Include a brief abstract at the
beginning of the text (abstracting services pick this up directly) followed by an
introduction (understandable by any scientist) and then whatever paragraph
headings are desired. The usual editorial customs, as exemplified in the most
recent issues of the Journal, are' to be followed as closely as possible.

In the text, cite all references by author and date in a chronological order, i.e.,
Jones (1971); Jones (1971 , 1972); (Jones, 1971); (Jones, 1971 , 1972); Jones and
Smith (1971); (Jones and Smith, 1971); (Jones, 1971; Smith, 1972; and Beacon,
1973). If there are more than 2 authors, use: Jones, et al. (1971); (Jones, et al,
1971). References are then to be assembled, arranged ALPHABETICALLY, and
placed at the end of the article under the heading LITERATURE CITED. For a
PERIODICAL ARTICLE use: Jones, A. P., and R. J. Wilson, 1971-Effects of
chlorinated hydrocarbons./. Comp. Phys., 37:116. (Only the 1st page number
of the article is to be used.) For a PAPER PRESENTED at a symposium, etc., use
the form: Jones, A. P., 1971— Effects of chlorinated hydrocarbons. WMO Sym¬
posium on Organic Chemistry, New York,N.Y. For a PRINTED PAPER use: Jones,
A.P., 1971— Effects of chlorinated hydrocarbons. Univ. of Tex., Dallas, or Jones,
A. P., 1971— Effects of chlorinated hydrocarbons. Univ. of Tex. Paper No. 14,46 pp.
A MASTERS OR Ph.D THESIS should appear as: Jones, A. P., 1971— Effects of
chlorinated hydrocarbons. M.S. Thesis, Tex. A&M Univ., College Station. For a
BOOK, NO EDITORS, use: Jones, A. P., \91 \ — Effects of Chlorinated Hydrocarbons.
Academic Press, New York, N.Y., pp. 13-29. Fora CHAPTER IN A BOOK WITH
EDITORS: Jones, A. R, 1971— Structure of chlorinated hydrocarbons. In A. P. Jones
and T. S. Gibbs (Eds.), Effects of Chlorinated Hydrocarbons. Academic Press,
New York, N.Y.,pp. 3-12. For an IN PRESS PERIODICAL reference, use: Jones, A.P.,
1971— Effects of chlorinated hydrocarbons./, of Org. Chem. , In Press. For an
IN PRESS BOOK reference, use: Jones, A. P., 197 1— . Effects of Chlorinated Hydro¬
carbons. Academic Press, New York, N.Y. In Press.

All tables are to be typed with a carbon ribbon, free of error, without hand¬
written notations, and be prepared for photographic reproduction. Tables should
be placed on separate sheets with a marginal notation on the manuscript to indicate
preferred locations. Tables should have a text reference, i.e., Table 2 shows ... or
(see Table 2).

4

THE TEXAS JOURNAL OF SCIENCE

Figures are to be original inked drawings or glossy photographs NO LARGER
than 6V2X4V2 inches and mounted on standard 8V2X 11 paper. Legends for figures
are to be typed separately and lettering within' the figure kept to a minimum.

All photographs, line drawings, and tables are to be provided with self-
explanatory titles or legends. Each illustration should be marked on the back
with the name of the principle author, the figure number, and the title of the
article to which it refers.

Galley proof of each article will be submitted to the author. This proof must
be carefully corrected and returned within 3 days to the Managing Editor’s Office
(Dr. Mike Carlo, Managing Editor, P. 0. Box 10979— ASU Station, San Angelo,
Texas 76901). Page proof will not be submitted. Page charge ($35/page) and
reprint costs MUST accompany the return of the corrected galley of the manu¬
script (Check or Purchase Voucher). A delay in the printing of the manuscript
will occur if payment is not submitted with the return of the galley.

Reprint price list and page charge information will accompany galley proofs.
Reprints are delivered approximately 6 to 8 weeks after articles appear.

NOTICE: IF YOUR ADDRESS OR TELEPHONE NUMBER CHANGES, NOTIFY US
IMMEDIATELY SO WE CAN SEND YOUR GALLEY PROOF TO YOU
WITHOUT LOSS OR DELAY.

BIOGRAPHICAL SKETCH

5

ROBB E. MOSES, M.D.
BIOGRAPHICAL SKETCH

Robb E. Moses received his M.D. degree from the Johns Hopkins University
Medical School. While there, he was a Fellow in the laboratory of Dan Nathans.
Following an internship in Internal Medicine at Hopkins, he joined Maxine Singer
at the National Institutes of Health as a Staff Associate. Subsequently, he was
appointed a USPHS Special Fellow with Charles Richardson at Harvard. He has
been on the faculty at Baylor College of Medicine in Houston since 1971.

EXCISION REPAIR FOLLOWING ULTRAVIOLET IRRADIATION
IN TOLUENE-TREATED ESCHERICHIA COLI

by JOHN W . DORSON and ROBB E . MOSES

Mans McLean Department of Biochemistry
Baylor College of Medicine, Houston 77030

ABSTRACT

Repair of UV-rad&tion photoproducts has been examined in bacteria made permeable
by exposure to tolj/ehe. In toluene- treated cells this repair is due to the excision repair path¬
way. An active participation in this process by DNA polymerase I is demonstrated by two
different experimental approaches. In the first, irradiated cells are allowed to accumulate in¬
cisions in the absence of repair synthesis and then repair synthesis is allowed. In wild-type
cells this results in a rapid reformation of high molecular weight DNA. The other approach
is more direct as the polymerization by DNA polymerase I is directly stimulated. This stimu¬
lation is caused by the addition of nicotinamide mononucleotide which blocks polynucleotide
ligase activity. This addition causes a large increase in the amount of repair synthesis after
UV which is specific for DNA polymerase I. These experiments lead to the conclusion that
the repair synthesis catalyzed by DNA polymerase I is an efficient process in which the repair
synthesis is easily terminated by polynucleotide ligase and that few nucleotides are inserted
into the damage site.

INTRODUCTION

UV radiation causes the formation of dimers between adjacent pyrimidines
on the same DNA strand (Setlow, 1966). There are at least 3 mechanisms for the
repair of these dimers (Hanawalt and Setlow, 1975). Photo-reactivation is the
process of in situ reversal of the dimers. This enzymatic process requires visible
irradiation. Postreplication or recombinational repair requires DNA replication
and the rec gene products. This process repairs gaps in the daughter strands in
regions opposite dimers .Excision repair is a cut and patch process wherein the
dimers are removed from the DNA and replaced via repair synthesis. It is this final
process which we will discuss in this report.

In the simplest terms, excision repair consists of individual steps of incision,
excision, repair synthesis and ligation (Grossman, et al., 1970). Incision is the
introduction of a break into the phosphodiester backbone of the DNA near the
dimer and is under the control of the uvr gene products. Excision is the step in
which the dimer is actually removed from the DNA. Repair synthesis then fills
in the damaged region with correct nucleotides using the opposite strand as tem¬
plate. These nucleotides are joined to the undamaged DNA strand by polynucleo-
Accepted for publication: November 1, 1976.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

8

THE TEXAS JOURNAL OF SCIENCE

tide ligase to complete the repair process and return both sequence integrity and
intactness to the DNA.

Toluene -treated Escherichia coli are permeable to small molecules and present
an in vitro system for a more thorough examination of the repair processes than
in vivo (Moses and Richardson, 1970). We have used this system to dissect the
repair process into its individual steps and then examine these steps. For example,
incision is difficult to study in vivo because the nicks that are made are rapidly
repaired so that at any time during repair there are actually few nicks present in
the DNA. In toluene- treated cells by not supplying the deoxyribonucleoside tri¬
phosphates (dNTPs) repair synthesis is blocked and incisions cannot be repaired,
thus accumulating. In this way, we have been able to study incision as a discrete
step. We have shown that incision requires the presence of ATP and proceeds for
about 10 to 15 min at 37 C (Dorson, et al. , 1975). In this report we examine the
contribution of the DNA polymerases to the repair synthesis.

MATERIALS AND METHODS

Radioactive materials were obtained from Schwarz/Mann. NMN was obtained
from Sigma.

Cells were grown in L-broth (10 g tryptone, 5 g yeast extract and 5 gNaCl)
supplemented with 10 Mg/ml thymine. To label DNA, the cells were grown in the
same media with the addition of 5 pc/m\ tritiated thymidine.

Cells were toluene -treated as previously described (Moses and Richardson, 1970).
Cells were irradiated at a dose of 6 J/m2 min following toluene treatment and
immediately placed into reaction mixtures.

Sucrose Gradient Sedimentation

Labeled toluene -treated cells were incubated after irradiation in reaction mix¬
tures which contained no radioactive nucleotides. Following incubation the cells
were lysed directly on the top of alkaline sucrose gradients. The gradients were
centrifuged for 2 hrs at 25,000 rpm at 20 C. Collection was from the bottom direct¬
ly onto Whatman Number 3 paper strips which were then washed with TCA and
cut into scintillation vials and counted. The data is expressed as % of the total
counts recovered from the gradient versus the % of the gradient migrated. Molec¬
ular weight determinations were by comparison to 0X174 RF1 DNA.

Incorporation Determinations

To measure DNA synthesis unlabeled cells were incubated in reaction mixtures
containing deoxyribonucleoside 5'- triphosphates one of which was radioactive .
The incubations were stopped by the addition of TCA and acid-insoluble material
collected by filtration. The filters were washed with TCA, placed in scintillation
vials and counted. Based on specific radioactivity of the precursor, the incorpora¬
tion was converted to picomoles.

EXCISION REPAIR

9

RESULTS

Sedimentation Analysis

The complete process of excision repair is comprised of many steps or partial
reactions. These steps are tightly coordinated. As excision repair requires the intro¬
duction of breaks into DNA it seems that these interruptions are removed as rapidly
as possible. The result is that in live cells after UV- irradiation the DNA is remark¬
ably intact at any instant. That this is also the case in vitro is shown in Figure 1.

% GRADIENT

400 150 25 0

M X 10-6

Figure 1 . Maintenance of high molecular weight DNA during excision repair when all the
requirements are available. Toluene- treated wild-type cells (W3110) were ir¬
radiated (30 J/m2) and incubated in reaction mixtures including ATP and dNTPs
and stopped at various times. A = 2 min, O = 20 min, • = unirradiated,
20 min.

10

THE TEXAS JOURNAL OF SCIENCE

After irradiation, toluene- treated wild- type cells incubated in complete reaction
mixtures containing the appropriate substrates for DNA synthesis including ATP
demonstrate little breakdown in the DNA. This analysis is performed on alkaline
sucrose gradients which denature the DNA and is therefore a sensitive indicator
of the intactness of the DNA strands.

The rapidity of the complete excision process requires jthat a block be intro¬
duced in order to examine any of the individual steps. By blocking the repair syn¬
thesis step any breaks that have been introduced into the DNA may be observed.
The arrow (Figure 1) near the top of the gradient indicates the peak position of
the counts when repair synthesis is blocked by withholding the deoxyribonucleoside
5 '-triphosphates (dNTPs) which are the substrates for the repair synthesis. We
previously demonstrated that UV-specific incision requires the presence of ATP
(Dorson, et al., 1975). Incisions accumulate in the absence of repair synthesis
which is blocked by omitting the dNTPs from the reaction mixtures. This experi¬
ment illustrates the efficiency of excision repair in maintaining the intactness of
the DNA even though there are breaks being introduced into the DNA.

One way for this process to be so efficient would be for the repair and sealing
of the DNA to be very rapid compared to the introduction of the initial incision.
Incisions accumulate when irradiated toluene-treated cells are incubated in the
presence of ATP but in the absence of dNTPs. The closed circles (Figure 2) are
the distribution of the DNA after such an incubation for 10 min. The cell’s ability
to repair and reseal these breaks is examined by the subsequent addition of dNTPs
to allow repair synthesis and the rejoining of the repaired region to the DNA. This
rejoining requires the correct configuration in the DNA at the juncture of the
break. The results (Figure 2) indicate that as quickly as the first analysis is made
(2 min) that the majority of the breaks in the DNA have been repaired and sealed.
Even though these breaks required about 10 min to accumulate most are sealed
much more quickly indicating the speed of the repair process after the incision
has occurred. Further incubation (20 min, open squares) results in a distribution
that is very similar to that seen for DNA from unirradiated cells taken through
the same protocol (closed squares).

When a similar experiment was performed with polA~ cells (DNA polymerase I
deficient) there was only a small increase in the molecular weight distribution
when the dNTPs were added. Thus, the rapid complete reformation seen in wild
type is the result of participation by DNA polymerase I.

Incorporation Studies

The activity of the DNA polymerases can be followed by directly measuring
the synthesis of DNA during the postirradiation incubation. This incorporation,
however, is complicated, due to replicative synthesis. The data in Figure 3 is pre¬
sented to distinguish the patterns of incorporation due to repair and replication.
In the left panel all the incorporation is due to replication as the strain is uvrA"
preventing excision repair (Moses and Moody, 1975). Replicative incorporation

EXCISION REPAIR

11

Figure 2. Reformation of high molecular weight DNA subsequent to incision accumula¬
tion. Toluene- treated wild-type cells (W3110) were irradiated (30 J/m2) and
incubated 5 min in reaction mixtures containing ATP but no dNTPs (A). The
dNTPs are then added and incubated for additional time (• = 1 min and 0=4 min).

falls as the UV dose is increased indicating the inhibition to replication (Bowersock
and Moses, 1973). Repair synthesis is, of course, low until the cells are exposed
to UV. This is seen in the center panel when replicative synthesis is blocked in a
mutant with a conditional defect in replication, dnaB ts. As the UV dose increases
the repair synthesis also increases until the system appears to saturate and levels
off. The result of combining both types of synthesis in the same cell is illustrated

12

THE TEXAS JOURNAL OF SCIENCE

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for a p#M~ strain in the right panel. The initial high level of replicative synthesis
falls as the UV dose is increased just as before. However, there is now a residual
synthesis which is constant at the higher UV exposures. This level of incorporation
is seen to be nearly the same as in the center panel in which only repair synthesis
occurs. This residual level of synthesis is about thirty percent of the replicative
level.

EXCISION REPAIR

13

All of the panels in Figure 3 were in polA~ strains which lack DNA polymerase I.
When a comparable experiment is performed in a toluene-treated wild- type strain,
the results in Figure 4 are obtained. The level of replicative synthesis is not influenced

Figure 4. DNA synthesis in toluene- treated wild- type cells (W3110) following irradiation.
Experiment performed as described in Figure 3. •= complete, O = -ATP.

by the absence or presence of DNA polymerase I. Once again, the level of incor¬
poration due to replication falls as the UV dose is increased. In a DNA polymerase I-
containing strain, however, the level of synthesis that is observed after high UV
exposures is very low, in this case near to the background incorporation seen in
the absence of ATP.

14

THE TEXAS JOURNAL OF SCIENCE

How can such low levels of repair incorporation be accounted for and is there
repair synthesis at all? It was demonstrated in Figure 1 that there is a very efficient
repair in wild type. If the number of nucleotides being inserted at each damage
site were few enough then the amount of synthesis could be so small as to escape
detection. One way for this to occur is by an active participation by polynucleotide
ligase. Ligase could seal the nick in the DNA quickly after the dimer had been re¬
moved, thus limiting the number of nucleotides inserted. In that event, blocking
ligase should lead to a more extensive repair synthesis.

Nicotinamide mononucleotide (NMN) is a product in the ligase activation reac¬
tion (Little, et al, 1967). NMN addition reverses the reaction producing an un¬
charged ligase which is incapable of sealing nicks in DNA. The addition of NMN
to toluene-treated cells, blocking ligase activity, results in a large stimulation of
repair incorporation (open circles. Figure 5). The addition of NMN is seen to have
little effect on the replicative level of synthesis. Increasing the UV exposure First
results in a drop in the level of incorporation followed by increasing incorporation
at the higher doses. The initial drop is due to the UV being more inhibitory to
replication than in stimulating the repair synthesis. At the higher doses the incor¬
poration is almost exclusively repair synthesis.

The NMN-stimulated repair synthesis is ATP dependent as shown by the level
of synthesis in its absence (open squares, Figure 5). The ATP independent back¬
ground level of synthesis is slightly increased by NMN addition but does not vary
with changes in the UV exposure. The ATP requirement for repair synthesis can
be at least partially explained by the ATP requirement in incision as incision pro¬
ceeds repair synthesis (Dorson, et al, 1975).

DISCUSSION

Data presented in this and other publications from our laboratory are summa¬
rized in Figure 6. Excision repair is dependent upon the uvrA, B gene products
which control ATP-dependent incision. The requirements for this initial event
are consistent with the observation that other processes which follow incision
such as excision and repair synthesis are all ATP and uvrA, B dependent (Moses
and Moody, 1975; Bowersock and Moses, 1973; Deutsch, et al., 1976). Once
incision has occurred, however, the pathway may take 2 possible branches. This
branching depends on the presence of DNA polymerase I. Purified DNA polymer¬
ase I is capable of binding to incisions and removing dimers by a mechanism of
nick translation wherein the associated 5'-K3' exonuclease removes nucleotides
as the polymerase inserts undamaged ones in their place (Kelly ,et al, 1969). In
vivo nick translation is efficiently terminated by polynucleotide ligase once the
dimer region is passed. This accounts for the very rapid reformation observed
following incision accumulation in toluene-treated polA+ cells. That there are
few nucleotides inserted into each patch site explains why we are unable to ob¬
serve repair synthesis in wild-type cells. A dose of 30 J/m2 produces about 1500
dimers/genome (Deutsch, et al, 1976). At 8 x 108 cells/reaction this corresponds

pmoles dTMP Incorporated

EXCISION REPAIR

15

Figure 5. Effect of NMN on DNA synthesis in toluene-treated wild-type cells (W3110)
following irradiation. Experiment performed as described in Figure 3. • = com¬
plete; O = complete, +NMN; ■ = -ATP; and □ = -ATP, +NMN.

16

THE TEXAS JOURNAL OF SCIENCE

u i
if)
<

2

O

CL

<

Z

Q

TT

TT

—I - L,

|uv

| uvrA.B.(C)

a

yj

(/)

<

E

UJ

2

O

CL

<

Z

Q

-

Figure 6. Scheme for excision repair in toluene-treated cells.

to 1.2 x 10 12 dimers/reaction. If approximately 10 nucleotides are inserted at each
damage site in vitro as in the case in vivo (Cooper and Hanawalt, 1972) then the
upper limit for the incorporation in polA+ cells would be 20 pmoles. This would
occur if all the dimers were repaired. As about 30% of the dimers are actually re¬
moved in vitro (Deutsch, et al., 1976) this results in an upper limit of 6 to 7 pmoles
total incorporation which is too low to distinguish from the ATP-in dependent
background incorporation.

The addition of NMN to toluene -treated cells greatly stimulates the amount
of repair synthesis due to DNA polymerase I. NMN does not interact directly
with DNA polymerase I nor is it a necessary cofactor or activator. NMN acts by
blocking polynucleotide ligase and it is the absence of ligase activity which pro¬
motes DNA polymerase I synthesis. The inactivated ligase cannot seal nicks in
the DNA during repair and there is nothing to impede DNA polymerase I in exten¬
sive nicly translation. Conversely, the stimulation by blocking ligase is a direct
indication that the DNA contains sealable nicks during the normal repair mediated

EXCISION REPAIR

17

by DNA polymerase I. This efficiency in sealing the DNA polymerase I repair is
independently shown by the rapid and nearly complete reformation of the DNA
in Figure 2.

The 2 branches for DNA polymerase I involvement in excision repair in toluene-
treated cells are consistent with in vivo situations. In polA+ cells the repair patch
is short being only about 10 nucleotides whereas the patch in polA~ is about 1 ,000
nucleotides or a long patch (Cooper and Hanawalt, 1972). Therefore, the level of
repair synthesis in vitro parallels this same pattern, being more extensive in the
polAT strains.

The branch in the polAT strains is referred to as the DNA polymerase III path¬
way (Figure 6). The data supporting this contention is presented elsewhere
(Bowersock and Moses, 1973). However, the mechanism for this branch is dis¬
tinctly different than the nick translation mechanism in polA+ cells. There is no
stimulation by the addition of NMN to polA~ strains (Dorson and Moses, 1975)
and the breaks are not as readily sealed. This had led us to suggest that the prob¬
able DNA structure during DNA polymerase III repair is one containing a gap
with several to a few hundred nucleotides actually removed before the polymerase
begins acting. This most likely is the result of an independent exonuclease being
responsible for excision and removal of nucleotides past the damage. The physical
disconnection of the excising nuclease from the polymerase supposedly would
not allow the tight coupling or concerted action. The final step in either branch
of the pathway is the rejoining of the DNA break by polynucleotide ligase.

ACKNOWLEDGEMENTS

This work was supported by NIH Grant No. GM-19122, Robert A. Welch
Foundation Grant No. Q-543 and the American Cancer Society Grant No. NP-153B.
J. W. D. is supported by NIH Grant No. GM-0008 1-02. R. E. M. is recipient of
P. H. S. Career Program Award GM-70314.

LITERATURE CITED

Bowersock, D., and R. E. Moses, 197 3 -Ultraviolet irradiation inhibition of replicative Deoxy¬
ribonucleic Acid synthesis in toluene-treated Escherichia coli.J. Biol. Chem, 248:7449.

Cooper, P. K., and P. C. Hanawalt, 1972 -Heterogeneity of patch size in repair replicated
DNA in Escherichia coli. J. Mol. Biol, 67:1.

Deutsch, W. A., J. W. Dorson, and R. E. Moses, 1976-Excision of pyrimidine dimers in
toluene-treated Escherichia coli. J. Bacteriol., 125:220.

Dorson, J. W., W. A. Deutsch, and R.E. Moses, 1975 -DNA repair in toluene-treated Escheri¬
chia coli cells. Tex. J. of Sci., In press.

- , and R. E. Moses, 1975 -DNA polymerase I involvement in repair in toluene-treated

Escherichia coli. In M. Goulian and P. Hanawalt (Eds.), DNA Synthesis and Its Regulation.
Benjamin, Reading, pp. 8 15-821 .

18

THE TEXAS JOURNAL OF SCIENCE

Grossman, L., A. Braun, R. Feldberg , and I. Mahler, 1975— Enzymatic repair of DNA. Ann.
Rev. Biochem., 44:19.

Hanawalt, P. C., and R. B. Setlow (Eds.), 191 5 -Molecular Mechanisms for Repair of DNA.
Plenum, New York.

Kelly, R. B., M. R. Atkinson, J. A.Huberman, and A. Kornberg, 1969-Excision by Thymine
dimers and other mismatched sequences by DNA polymerase of Escherichia coli. Nature
(London), 244:495.

Little, J. W., S. B. Zimmerman, C. K. Oshinsky, and M. Gellert, 1967 -Enzymatic joining of
DNA strands. Proc. Nat. Acad. ofSci., U. S. A., 5 8:2004.

Moses, R. E., and C. C. Richardson, 1970 -Replication and repair of DNA in cells of Escherichia
coli treated with toluene . Proc. Nat. Acad. ofSci., U. S. A., 67:67 4.

- — — , and E. E. M. Moody, 1975 -DNA repair synthesis dependent on the uvrA, B gene

products in toluene-treated cells.,/. Biol. Chem., 250:8055.

Setlow, R. B., 1966— Cyclobutane-type pyrimidine dimers in polynucleotides. Science, 153:379.

MARKOV- RENEWAL PROGRAMMING UNDER INCOMPLETE
INFORMATION - 1 .

by J.S. PRASAD

Consolidated Reactive Metals, Mamaroneck, New York 10543
and A. G. WALVEKAR

P.O. Box 4230, Department of Industrial Engineering,

New Mexico State University, Las Cruces 88003

INTRODUCTION

Multistage decision processes have been the subject of active research during
the last decade. The scope of sequential decision making has grown with the
development of new mathematical models to represent practical situations. This
paper deal0 with the optimization of the output of a system whose behavior is
represented by a Markov- renewal process— while the structure of this mathematical
model is know, the decision maker is assumed to be operating with incomplete
information about the parameters of the mode.

Statement of the Problem

Consider a Markov- renewal process with rewards described by the quadruplet
P, F, R, and Z where

p={pfj}

F = {Fy (t) }

R= {Ry(t|x(i,j))}

is the matrix of rewards due to the
transition from i to j in the clock
time t,0 <_■ t < t (i,j), since the begin¬
ning of the transition,

is the conditional transition proba¬
bility matrix (i,j = 1,2,... ,N),

is the matrix of transition interval
distribution functions,

is the set of alternatives available to
the decision maker when the system
is in state i, i = 1 , 2, . . . , N.

Accepted for publication: September 12, 1974.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

20

THE TEXAS JOURNAL OF SCIENCE

A Markov- renewal decision problem is denoted by MRD in the sequel. If an
MRD involves incomplete information about P and/or F, they are treated as ran¬
dom variables and are designated by a tilde (~) placed above those parameters.
The lack of information about the parameters of a Markov- renewal process leads
to 3 distinct MRD’s, namely:

1. MRD: F, R, Z }

2. MRD: {P.F.R.Z}

3. MRD: {F,F, R, Z}

Optimization Under Incomplete Information About Transition Probabilities
Consider an MRD: {F, F, R, Z} Let the prior distribution of? be G(P|m)
with the parameter p and where G belongs to the family of distributions T which
is closed under consecutive sampling rule. Additionally, assume that the 1st and
2nd moments of the transition interval distributions are finite and nontrivial.

The expected discounted return due to the transition from state i to state j
under the zth alternative in i is

pu(a>V°°dF5(T);/V“x-dxRu(x!T) (1)

where a continuous discounting, e"ax with a>0, is used.

The average 1-step discounted return when starting from state i and with the
knowledge ofF* being indexed by p is

P?(a,*0= s pfj(ot) •/ pfj(ju)*dG(P|M)

j=l J SZN

= 2 p?j(a)*p?:(M) (2)

j = 1 1J 1J

Here, pz.(ju) is the expected value of the probability of making a transition from

1J f-L

state i to state j under the z alternative in i.

Let Vj(ju) be the maximum expected discounted reward over an infinite period
when the system starts from the generalized state (i, p). Then, each v^p) must
satisfy the following simultaneous functional equations:

VjO) = Zmez) {Q-(a’0+ .2i^j(Ju)-/dF^(t).e-«,vj(T|(M))}

for i = 1 , 2, . . . , N, pe M; and a >0.

Now, define the functions v^n, p) recursively as follows:

vi(n.M) = Z™2X. {pj<a,M)'+ ,^Py(p)- gy(a) '^(n-l, Tj-O)) }

(3)

for i = 1,2, . . . ,N; peM; n = 1, 2, . . . ; a >0

(4)

MARKOV- RENEWAL PROGRAMMING -I

21

where {v^O, £t) }= (V^/i) }is a set of bounded terminal rewards, and

^(a) = Aat'dFy(t) (5)

=0/°6"at *fy(t)dt (6)

if f?. (t) exists. It will be shown that {v.(n,ju) }forms successive approximations to
{v.(ju) }. It should be noted that 0<. 3^ (a)< 1 for all (i, j, z) whenever a>0, for, if

By (a) = /‘2 e'atdF?.(t), 0< t,<t2<»

‘i

then, by the mean value theorem, there is a point t* such that tl <t*<t2 , for which
Sy(a) = e'ai*f2 dF?j(t) = e‘at* < 1 .

tl

Lemma 1

Let V and R* be the bounds defined as follows:

| VjCju) I < v,

R* =™zX { I pf ( Ct , ju) | } for i = 1, 2, . . . , N; zeZj
Then, the functions v^n, /j ) are bounded:

k(n,M)| < gnV+i^R* (7)

1- P

where B - {^(o).} (8)

Proof: The hypothesis of the lemma is trivially true for n = 0. Assume the
lemma for n and let

vj(n+l , u) = P* (04 m) + . Py (11) • By (a) *Vj(n, T| (ji))

Then,

|Vj(n+l,/j) | < |pj*(a, m) l+B .^PyO^)* lvj(n> Ty (a0) |

< R*+ B{B"V + i^R*} 2 p*(u)

1-B j = l 1J

= (R* - gR* + gR*_ gn+lR*)+ gn+ly

1- 3

= 3n+1y + lr.3n+1 r*

1-3

22

THE TEXAS JOURNAL OF SCIENCE

That is,

|v.(n+l,At) I < gn+1V + l-.e"!1 R*.

>' n : ■' 1-3

Hence, by induction, the lemma is true for all n>0.

Lemma 2

If V, R* 3 and the set of functions {v^n, n) }are defined as in earlier equations,
then,

KCn, M) - Vj(m, M)| < (Bn+ Bm)V + R* (9)

for all m<n, i = 1 , 2, . . . , N.

Proof: From Lemma 1 we have that
|v.(n, //) - v.(0, n)\< (1 + gn)V + R*.

Now, assume the hypothesis of this lemma to be true for m. Then,
|vi(n,(u)-Vj(m+l,jU)| < 6nV+^R*-R*

1 r> m

+ £5{BmV - If^R*}
l- p

= (en + 6m+1)v+ i^R*

1-3

P*rl-B+ B- 6m + 1 1

1 l-B

Therefore,

| v,(n , ») - Vi(m+ 1 , ai) | < ( Bn + Bm + 1 )V + R* •

Hence, by induction, the lemma is true for all m<n.

Theorem 1

The limit of the sequence {v^n ,/jl) \ as n tends to infinity, exists.

Proof: From Lemma 2, it can be seen that
|vi(n,iu)-vi(n-l)/i)|.;< (Bn+Bn'1)V+

MARKOV- RENEWAL PROGRAMMING -I

23

Note that

lim {(on + gn-l)V+ gn-1-gn R*}=0.

n->oo^M P / 1_ g J

That is, for any e > 0, there exists an n* such that for all n>n*

|(en + + gn-1- b"r* | < e.

1- P

This implies that, for all n>n*

| Vj(n , //) - v.(n-l,/i) | < e,i = 1,2, . . . ,N
Satisfying the cauchy criterion for the convergence of the sequence {v.(n , /i) }.

Theorem 2

Let

v* = min {Vi(M) }. v* = max{Vi(M)

r* = ™‘z {pf («. P) ■>?■ R* = ?,? {pf (a, At) };

and

min rflz/ \-i
i,jj2{gij(“)}>

0" _ max rfl z / \ -1
3 -i i ztSyU)}.

If r* + 3'v* _> V*, then {v^n, p) }increases monotonically to {v. (/*)}; and if
R* + $"V* < v* then {v.(n, fi) decreases monotonically to fyOu) } as n increases.

Proof: Case l,r* + 3'v*> V*
It is clear that

- Vj(0,M) =

> r*+ g'v*-V* > 0 i = 1,2, . , , , N.

Now, assume the theorem for n and let
vj(n+l , n) = p?(a,/i)+
and

'/i(n,u)= pJ’(a,M)+ .2 p^(p)-e^(a).vj(n-l,Tb(p)).

24

THE TEXAS JOURNAL OF SCIENCE

Then,

Vj(n+1 , n) - Vj(n,/z) = (p? (a, p) - p?’ (a,u))

+ j Py (M) • By (a) • Vj(n , Ty (p))

- . ^ pj] (m) • By (a) -v/n-l , T?] (p))

“ j ?, Pij ^ ‘ ey (“) ' (vj(n > T!j O')) - Vj(n-1 , Ty (u)) }
>0 i = 1,2,. ...N.

Hence, by induction, the theorem is true for all n>0.

Case 2, R* + 3"V* <_ v*
It is seen that

Vj(l, u) - vj(0, M) = {ff (a, m) + s p?j(*i) • By (a)*Vj(0, Ty Qt)) }- V(M)
<_ R* + 3” V* - v* <. 0

The remainder of the proof follows along similar lines as for Case 1 .

Now, letting n approach infinity, it is seen that v^n , /u) converges monotonically
to v^ju) for all i = 1 , 2, . . . , N, thus proving the theorem.

In the preceeding 2 lemmas and 2 theorems, the existance of an optimal solu¬
tion to the MRD with incomplete information about the transition probabilities
has been established. Theorem 4 also establishes a set of sufficient conditions
under which the functions (v^n,//)} converge monotonically to {v-(m) }•

The recursive equations (4) can be used to compute {v^n, n)} iteratively for
n = 1, 2, . . . when the terminal functions (Vf/x)} are specified. In using this suc¬
cessive approximation technique, it is necessary to develop a reliable rule for
stopping the iterations. The sequence {v^n, ju)} has been shown to converge to
{VjO-t)}, but only sufficient conditions were developed for its monotonicity. A
more acceptable stopping rule is established in Theorem 3 which follows Lemma 3
which is needed first.

Lemma 3

Let r*, R*, and 3" be defined as before. Then

<

Vj(Ai) <

R*

1-3"

(10)

i=l,2,...,N.

MARKOV-RENEWAL PROGRAMMING -I

25

Proof: The mean reward per transition must satisfy
r* < p?(a ,M) ^ R*

1 =1,2, . . . , N z£Z.

If the system is run over an infinite period, the bounds on are obtained by
summing the bounds on p? (a, ju) with the appropriate discounting. Thus,

oo oo

2 r*(3')n < v.(/i) < 2 R*(3")n

n = 0 1 n - 0

or

Vj (n) <

R *

1-3"

i = 1,2,. . . ,N.

Theorem 3

Let the terminal functions {V.(ju)}be constants and let e.(n, /i) be the error of

th 1 1

the n approximant. That is, let

ejOi.ju) = Vj(n) - v^n,//) i=l,2,...,N. (11)

Then

jejCn.M) j < (B")n {maxC-^p - v*. V* - yyy)} • (12)

Proof: When n = 0,

ej(0 , At) = Vj(m) - V..

If w.^jd) > Vj, then

|ej(0> m) I i - V*.

On the other hand, if v^ju) < V., then

|ej(0, n) | = V.-v.( ji)-< v*- Tf^r
Hence, |e.(0,p) | 1 max{jr|Fr - v*,V* - yyy } •

Having established the theorem for n = 0, assume that it holds n and let
VjGO = P^(a,M) + j|1^(M)-^(a)-vj(T|(M»

26

THE TEXAS JOURNAL OF SCIENCE

and

Vj(n = 1,m) = pf (a, M) = . pjj (m) • By (a) ’Vj(n, Ty (u))

Thus,

. 2 p]> (M) • By (a) [Vj(Ty GO) - Vj(n, Ty 00)]'

< e;(n+l,M)

< $ Py (At) • ^(^[YjCTjO*) - vj(n> TyC/t))]

, N

Let 2 Ph (m) ' Bh (ct) [v-(T? (u)) - vi(n, Tj?(/t))] be the larger of the absolute
1 j = I y y j y j y 1

values of the 2 bounds. Then,

|ej(n+l » At) j' - . Sj pS O) • By (a) • | vj (T y (m) - ^(n, TJ (At)) |

:l B''[(B'')n{max(I^fr - v*. V* - ]

That is,

|ej(n+l , m) | < (e")n+1{niax('T^r-v*,V*-;-^r)}.:

Hence, by induction, the theorem is true for all n >. 0.

Note that the sequence

(B’TWj^r - v*, V* - )

converges monotonically to zero as n approaches infinity even though {|ej(n,/i)|j
need not be a monotonic sequence. Thus, if e> 0 is an error acceptable to the
decision maker, it is possible to compute the smallest n which satisfies

( b" )n {max( - v*. V* - — }<E. (13)

Then so computed gives the maximum number of iterations that need to be done
on {v^n, ju) } to guarantee an e-optimal estimate of (v^q) } . This can therefore
be used effectively as a stopping rule.

In dealing with an MRD: { P , F, R, Z } , one way of estimating this unknown
quantity is on the basis of frequency counts. Consider an MRD from which a
sample Xn = {x0 , Xj , x2 x } of n transitions have been observed under con¬
secutive sampling rule. Here, x0 is the initial state known before sampling. Let

MARKOV- RENEWAL PROGRAMMING -I

27

vz- be the number of transition in X from i to i under the zth alternative in i.
ij _ n J

Let v = {vy } be the Z x N matrix of transition counts. Then, the likelihood of
observing Xn given that F = P is

N Z * z

n n1 (pp^ij. (14)

i=l z —1 1J

j=l

It is seen that v is a sufficient statistic for this data generating process. The natural
conjugate distribution is matrix beta distribution. That is, P has the joint density
function with parameter /!,

where

f(p|M) = k(M) n n> (pf.fij"1

i=l z=l 1J

j=l

n z. rou?)
k(m) = n n1 ^ -

1=1 Z=1 n r(pz)

j=i 1]

and ii - {jUy} is a Z x N matrix such that

N

(15)

(16)

(17)

(18)

Theorem 4

Let P have matrix beta distribution with parameter //. Suppose that a sample
with transition counts vis observed under consecutive sampling rule. Then the
posterior distribution of F is also matrix beta with parameter

m" = T + v

(19)

Proof: By Bayes’ theorem the posterior density function , f(P | (i , v), is proportion¬
al to the product of the kernel of the likelihood function and the kernel of the
prior density function. Therefore,

f(P l/A v) -

n n; (p^r1

1=1 z=l J

j=l

(20)

28

THE TEXAS JOURNAL OF SCIENCE

The right-hand side of (20) is the kernel of a matrix beta density function
with parameter p! + v. Hence the theorem.

The fact that the family of matrix beta distributions is closed under consecutive
sampling rule follows directly from this theorem.

If? follows a matrix beta distribution with parameter p = then

Example of an MRD: {?, F, R, Z)

Consider an MRD with 2 states and 2 alternatives in each state. Let the transi¬
tion intervals be random variables with the following density functions:

with

ff.(t) = Gf-e"6^

e = (efj } =

5

4

6

10

3
2

4

5

Assume that the transition probability matrix P is not known completely but
is known to follow matrix beta distribution with parameter p,

When the system is in state i and is being operated under the zth alternative,
let the reward accumulated by r.z units per unit of time,

R = {rf } -

9

6

-3

-6

The problem is to maximize the total expected discounted reward when the
system is operated for an indefinitely long time. Continuous discounting with
a = 40 is to be used. Now,

P-j(a) = / dF£(tJ./V
J 0 J 0

ax

1XRU (x|t>

a + 0:

and,

MARKOV- RENEWAL PROGRAMMING -I

Assume

Hence,

fSy(a) = / e“atdFy (t)

a + 0

m = {pfj(p)}

{pfj(a)}

{6y(a)|

N

pf(a,^) = S
1 j = l

(p?(a,//)}

v(0,m)

” 0.5

0.5“

0.8

0.2

0.4

0.6

_ 0.8

0.2^

~ 0.20

0.2 f

0.14

0.14

-0.07

-0.07

-°.i2

-0.13_

0.11

0.0?"

0.09

0.05

0.13

0.09

0.20

0.11

M) Py (a)

" 0.205“

0.140

-0.070

-0.120

v,(0,ju)

0

v2(0, n)

0

vi(l,M) = 2£Zj {p? (a, m) + ^ PyO) • By (a) -VjCO, Ty (m)) }

_ max r Z( ,ai
” zeZ. fPi

v(l ,jti)

0.205

-0.070

30

THE TEXAS JOURNAL OF SCIENCE

The policy corresponding to this is £ (1 , ju) -

N

. Now,

vj(2 , m) = ™z. {p^(a,M) +.?1Py(M)-By(a)'Vj(l,TS(M))}
i, j = 1 , 2; z = 1,2.

Hence,

From this, {p? (a, T} ^ju))} is computed to be

0.20

{p-CcnT} ,(/*))}

0.14

0.07

0.12

which yields

The following can be computed using a similar procedure:
v( 1 , Tj j(/i))

r 0.20"

[-0.07 _

v(i, t}2(m))

v(1,T^(M))

v(1,T120)) =

= r °-2ii -

L-0.07J ’

= r °-2fi ■

L-0.07J ’

= r °-2ii •

L-0.07J ’

0.21]

■0.07] ’

v(1,T?2(m)) =

0.21]

■0.07J

v( 1 , Tj ,(/J))

v(l,T222(M))

= r°-

L-0J

= r°-

L-o.

0.21

0.07

0.21

0.07

This can be used in computing v(2, ju) as follows:

Pi (a, m) + pj 1 (m) • i(a) *v2 (1 , Tj 2(/z))
+ pi2(M)-e!2(c0-v2(i,T|2(M))
p?(a,/u) + p?1(p)*eu(a)-v1(l,Tf1(M))

+ Pu(aO • 612(01) -v2(l , T?2(m))

v1(2,Ju) =max

MARKOV- RENEWAL PROGRAMMING -I

31

max

0.214

0.154

0.214 .

Similarly,

Thus,

v2(2,m) = max

-0.063

-0.088

0.063.

r°-

L-o,

214

.063

v(2 , u)

The policy corresponding to the 2nd approximation is
S(2,M) = |

By following a similar procedure, the 3rd approximants are computed.lt should
be noted that to calculate v.(3, ju), it is necessary to compute v^, T y(M)) which
further involves computing v^l , Tj^CIy (/*))). It can be shown that

and

v(3,m)

0.214

-0.063

C(3,M) =

RESULTS

The results are summarized in Table 1 . The error bound shown in the last column
is computed using the inequality (13). Note that .

v* — y* = o; R* = 0.205, r* = -0.120,

3' - 0.05, 3" = 0.20

_ . v* = 0.256; V* - -E,- = 0.126.

1-8 1-8

Therefore, the upper bound on the error of the nth approximant is
(e'Tt-r^rr - v*} = (0.2)n(0.256) .

1- P

Thus, v(ju) can be estimated with a 3rd decimal accuracy by performing 3 iterations.

32

THE TEXAS JOURNAL OF SCIENCE

TABLE 1

Summary of Computations for the Example MRD : F, R, Z } .

Error

n

v(n, j U)

£(n,M)

Bound

0

0

0

-

0.256

1

0.205

-0.070

1

1

0.051

2

0.214

1

0.010

-0.063

1

3

0.214

-0.063

1

1

0.002

LITERATURE CITED

Astrom, K. J., 1965 -Optimal control of Markov Processes with incomplete state information.
/. Math. Anal. & Appl., 10:174.

DeCani, J. S., 1964-A dynamic programming algorithm for embedded Markov chains when
the planning horizon is at infinity. Mgmt. Sci., 10:716.

Ferguson, T. S., 1967 -Mathematical Statistics, A Decision Theoretic Approach. Academic
Press, New York.

Howard, R. A., 19 60 -Dynamic Programming and Markov Processes. MIT Press, Cambridge, Mass.

Jewell, W. S., 1963 -Markov- renewal Programming I: Formulation, finite return models. Oper.
Res., 11:938.

- , 1963— Markov-renewal Programming II: Infinite return models, example. Oper.

Res., 11:949.

Martin, J. J., 1967 -Bayesian Decision Problems and Markov Chains. John Wiley and Sons,
New York.

MARKOV- RENEWAL PROGRAMMING UNDER INCOMPLETE
INFORMATION -II

by A. G. WALVEKAR

P.O. Box 4230, Department of Industrial Engineering,

New Mexico State University, Las Cruces 88003

and J. S. PRASAD

Consolidated Reactive Metals, Mararoneck, NY 10543
INTRODUCTION

The Markov- Renewal decision problem was introduced in the preceeding paper,
and the case of MRD: {P, F, R, Z} was discussed. The other 2 problems, namely
MRD: {P, F , R, Z} and { P , F, R, Z} are presented here. The solution procedure
is illustrated by numerical examples.

Optimization Under Incomplete Information About Transition Interval Distributions
Consider an MRD: {P, F, R, Z } . Let the transition interval distributions,
F = {F?j(t/\p)} indexed on \p, belong to a family $ which is closed under consecutive
sampling rule.

The expected discounted reward that is accumulated when the system makes
a transition from i to j under the zth alternative in i is

OO f

Py(a,ip) = J dFy(t/i|j) vf eax *dxRy(x/t). (1)

Again, continuous discounting, eaX with a > 0 is used.

The expected discounted reward when starting from state i and using zth alterna¬
tive is,

pf(a, \p) = 2^ Py • Py(ot, \p). (2)

When the system is operated for an indefinitely long time, let v^ip) be the
maximum total expected discounted reward accumulated when starting from
State (i, \p). Then Vj( ip) must satisfy the following set of simultaneous functional
equations:

Accepted for publication: September 12, 1974.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

34

THE TEXAS JOURNAL OF SCIENCE

v# = zmezx; {pjM + .sp:

N

j=l

/■dFjjCt/n,)

•eat.

Vj(T*z(^)} (3)

for i = 1 , 2, . . . , N; a > 0.

Here, Tjz is the mapping of i/jinto ipdue to the transition from i under the policy
z in an interval of time t.

A sequence {v^n, ip) } can be defined recursively as follows:

Vj(n, ip)

max
z eZ.

N

{pf (ex, ip) + . 2 p?j / ^ Py(t/iJ))'ea *• V: (n -1 , T jz ( f ) } (4)

j=l

1,2,. ...N.

Let the range of the transition interval be O^tj < t2 <°°- Then, by mean value
theorem, there is a value s such that t^ < s< t2, and

That is,

/2dFy(t/i|))*eat-vj(n,TtIZ(iJ;)) = eas .v.(n, T‘z(p)). (5)

tl

It should be remembered that s is generally a function of i, j , and z.

The equations (4) can now be written as,

Vj(n, 4)) = <pf(ot, i|>) + Pye“svj(n-l>T‘z(t(J))} (6)

i = 1,2, . . . ,N.

Note that

0<es< 1 (7)

for i,j = 1,2, . . . ,N; zeZ^ and a<0.

The following 2 lemmas and 2 theorems establish the existance of an optimal
solution to the MRD under consideration. They also show that the sequence of
approximants, {v^n, \]j)} converges to the optimal solution.

Lemma 1
If

V > |V.(*)| (8)

MARKOV- RENEWAL PROGRAMMING -II

35

R* = ™azx{|pz(a^)|} (9)

and

B = max{eas}; (10)

1 j J ? ^

then

|Vi(n,*)|i B”V+^R* (11)

for all n > 0, i = 1 , 2, . . . , N.

Lemma 2

Let V, R*, and 3 be defined as for the previous lemma. Then, for all m <n
|vj(n,*)-vi(m>H,)| < (gn+ em)V+ (12)

.i- 1,2 . N; n>0.

Theorem 1

Limit of the Sequence {v.(n, ip) } , as n

approaches infinity, exists.

Theorem 2

Let

V* = ““{VjOj,)} ,

V* = mf{V^)}

r* = ™z (Pf(«> <l>)} - '

R* = ““{pfO*.*)}

B* = ,

B" = {?“*}.

’ J’

If r* + 3#v* > V*, then {v.(n, ip) } increases monotonically to (v.(i|j)} , and if
R* + 3"V* <. v* then {v.(n, \jj)} decreases monotonically to {Vj( rp) } as n increases
(i = 1, 2, . . . , N).

The equations (4) can be used to arrive at the optimal solution by iterating on n.
A stopping rule that guarantees an e-optimal solution in a finite number of steps
is based on the fact that the sequence

(e")n {max(rSI7 _ v*, V* - yf^)}

converges monotonically to zero.

If the transition interval can take only discrete values as in the example pre¬
sented in the next section, then the frequency count can be used to estimate the

36

THE TEXAS JOURNAL OF SCIENCE

corresponding probability mass function. Let sample In = {i0 , ix , i2 , . . . , in } of
n transitions of the system be obtained by the consecutive sampling rule. Here ^ is
the initial state of the system known prior to the sampling. Let Xn= {Xj,x2, ...,xn}
be the transition intervals corresponding to In. When starting from State i and using
the zth alternative, let v!2 be the observed number of transitions of interval x. The
matrix of transition counts, v= {v,z } is a ZxT matrix where Z = Z- and T is

X j ” |

the number of discrete values that the transition interval can assume.

Given that the prior probability mass function the transition interval is (p(t/ip)
= U‘z}, the conditional probability of observing Xn is

N

n

i= 1

if

z= 1

n (tiz)v

x=l*x '

iz

x

(13)

If the stopping rule that determined the sample size n is noninformative, then
equation (13) is the likelihood of Xn . Hence <p(t/ip) = {(fdz } has a matrix beta
distribution with parameter ip; ip - { } isaZxT matrix, and

ip'xZ = VZ + 1 (14)

i= 1,2, . . . ,N; z= 1,2, . . . ,Z.; x= 1,2, . . . ,T.

It can be shown that if a sample Xn is obtained from a population that has
matrix beta prior distribution with parameter ip , then the posterior distribution
is also matrix beta with parameter + v where v is the matrix of frequency counts
resulting from XR.

Example of an MRD: {P, P, R, Z }

Consider an MRD: (P, F, R, Z } with 2 states and 2 alternatives in each state.
The transition intervals can be either 1 or 2 units of time, the probability mass
function governing it is now known. The matrix of frequency counts is

v = {vj2} =

from which ip is obtained to be

ip = {^tiz}= (v;z + i} =

6 4

1 12
2 5

5 2

7 5

2 13

3 6

6 3

The transition probabilities are:

MARKOV” RENEWAL PROGRAMMING -II

37

p= {Pp =

Now.

= pH:

0.5

0.5

0.8

0.2

0.4

0.6

0.8

0.2

Hence

let the reward accumulated be r? units per unit of time.

0.29

0.21

0.29

0.21

0.11

0.69

0.03

0.17

0.13

0.27

0.20

0.40

0.53

0.27

0.13

0.07

being operated under the

zth alt

R =

{rf }

9

6

-3

”6

The problem is to maximize the total expected discounted reward when the
system is operated for an indefinitely long time . Continuous discounting, eat with
a = 2, is to be used.

Now,

p?j(a,if)) = 2 * /*eax *r? «d

1J t = 1 J o

Thus,

2.057 2.057

2.318 0.579

-0.566 -0.849

-2.171 -0.544 _

and

2.057 “

1.970
-0.736 ‘

-1.846

{p?j(a, ip)} -

{pjZ(a, ifj) } -

38

THE TEXAS JOURNAL OF SCIENCE

Assume a starting policy ^(0

Then,

[-»:

057

736

v(0, ip)

If alternative 1 is used when starting from State 1 , then,

vl(l,» = pKa.^+pl.rfjRl/^^-v^O.Tj1^))

+ f{1(2/i|J).e2a.v1(0,T^1(i()))}

+ pl2f;2(l/^)-ea-v2(0>T!1(^))

+ fJ2(2/*)-e2a.v2(0,T‘1(^))}.

Similarly Vj(l , ip), ( 1 , ip), and v2(l , ip) are computed. Then

Vi(l,i|>) = max(v[(l,^); v*(l,iji))

and

v2(l, ip) = max(v2(l, iji); vf (1,-rp)).

Next,

t!1^)

5

13

6
3

which yields

v(0,Tj‘4)) =

Similarly, v(0, T2
is computed:

"«)= [

2.046

-0.736

2.085

0.736

, etc. Using these values, the next approximant

F 2.

L-°-

085

733

v(l, ip)

Repeating the iteration once more
v(2, ip)

The results are presented in Table 1.

2.086

-0.732

c(l. 'I') -

?(2, <Jj)

MARKOV- RENEWAL PROGRAMMING -II

39

TABLE 1

Summary of Computations for the Example MRD:

{P,F,R,Z}.

n

v(n, \p)

.C(n, i|d

0

2.057

1

-0.736

1

1

2.085

1

-0.733

1

2.086

1

2

-0.732

1

Optimization Under Incomplete Information About Both P and F

Consider and MRD: {¥*, V, R, Z}. Let the transition probability matrix,
P = {jp -}, have a distribution function G(P/^) with parameter p. Let G belong
to a family T of distributions closed under consecutive sampling rule. Similarly,
let the transition interval distribution, V - (P^(t/ijj) } with parameter ip, belong
to a family $ which is closed under consecutive sampling rule.

When the system starts from the state (i, p, ip), the expected discounted reward
due to the transition from i to j under the zth alternative in i is,

OO f

Py(a, <|0 =J dF?j(t /<(<)• / e“x .dxR*(x/t). (15)

The expected 1-step reward is,

P?(ct,/J, Tp) = f|0u)*Py(oi, <10- (16)

Let VjOu, ip) be the maximum expected discounted reward that is accumulated
if the system is operated over an indefinitely long period of time starting from
State (i, p, \p). Then v^p, \p) must satisfy the following simultaneous functional
equations:

N

Vj (MS = zrnea^.{p^(a,M,<|<)+ ZpgGO;/ dF^.e^.v^T^^)) }

i - 1 , 2, . . . , N; a >0; pe M; ipe^.

(17)

Here, again, T^t(jU,ip) is the mapping of Min to M and T into T due to the transition
from i to j under the zth alternative in i in an interval of t units.

Define a sequence, {y.(n , p, \p) }, recursively as follows:

V|(n,At,i/j) = 2 py(M) • / dF?(t/<|0 •

eat.

Vj(n-1

>T^,<K))}

(18)

i - 1, 2, . . . , N; a> 0

40

THE TEXAS JOURNAL OF SCIENCE

Let the range of the transition intervals be 01 tx <tj < ». By mean value theorem,
there is a value s such that tj < s < ^ , and

/t2dFg(t/i|,).e“t.vj(n,T^0i, <|t)) = (m, *))• /W*(t/*).

That is,

/^d FS (t/ •eat>Vj(n,Tjjt(M, 0) = (19)

Thus, equations (18) can be rewritten as.

N

i = 1 , 2 . N.

Lemma 3
If

then

V > |VjOi.*)

R* = ™aX{|pf(a,f<, <l')|)>
6 =imjaL«“s>’

i-e

(21)

(22)

(23)

(24)

|v:(n, /!,*)}< BnV+ -^R*

for all n -1 0, i = 1,2, ... ,N.

Lemma 4

If V, R*, and £ are defined as for the previous lemma, then for all m < n,

(3m _ |3n

|v.(n, n, i|)) - Vj(m, m, i|^) | - ( 3n + 3m )V + — — — R* (25)

for all n > 0, i = 1 , 2, . . . , N.

Theorem 3

The limit of the sequence (v^n , ii, rp ) } , as n approaches infinity, exists.

Theorem 4
Let

V* = mjn{Vj(M,i|;)}, v* = max(Vi(M, *)},

MARKOV- RENEWAL PROGRAMMING - II

41

r* = ™£{Pz(a,/i^)},
3' =iJ^gaS>’

R* = ™azx{Pz(a,M, if;)},

3" = ^{e-}.

If r* + 3'v* > V*, then (v^n, ju, \p) } increases monotonically to {v^ju, i />)}, and
if R* + 3"V* 5 v*, then {v^n, ju, \Jj)} decreases monotonically to {v-(p, ^)} as
n increases (i = 1 , 2 , . . . , N).

The preceding 2 lemmas and the 2 theorems establish that the sequence of
approximants, {Vj(n , ju, \p)j, converges to the optimal solution {v^/i, ip)}. Theorem 4
also establishes a set of sufficient conditions for the convergence of that sequence
to be monotonic.

The equations (18) can be used iteratively to arrive at the optimal or an accep¬
table solution. The number of iterations, n, required to guarantee a prescribed
e-optimality may be computed from

(6")nmax-^r -v*; V*--l-}<s (26)

1“ P I" P

Example of an MRD: {P,V,R, Z}

Consider an MRD: {P, F, R, Z) with 2 states and 2 alternatives in each state.
The transitions can take place in either one or two units of time with unknown
probability. A matrix of frequency counts from such a system is given below:

State 1

Trans. Interval
1 unit 2 units

State 2

Trans. Interval
1 unit 2 units

State 1
v =

State 2

II

N

7 3

5 5

z = 2

1 21

1 3

ii

N

1 4

2 7

N

II

K)

13 6

3 1

From this matrix, P(/i) and {f^(t/i|j)} are calculated to be

m =

0.50

0.80

0.39

0.78

0.33

0.07

0.11

0.52

0.50

0.20

0.61

0.22

0.17

0.73

0.28

0.26

0.25

0.07

0.17

0.15

0.25

0.13

0.44

0.07

{fy(t/«K>} =

42

THE TEXAS JOURNAL OF SCIENCE

When the system is in State i and is being operated under the zth alternative,
let the reward be r? units per unit time, with,

9 ~

6

-3 •

-6 _

The problem is to determine the maximum expected discounted reward that
is accumulated from operating the system for an indefinitely long period. Con¬
tinuous discounting with a = 2 is to be used.

Now,

fy(a, til) = o/dF'-(t/i|;) * /eax * if * dx

= ^fij(1/^)(1-ea) + ^(2/^)(l-e2“).

Therefore,

2.078
0.565
0.869
0.595

and,

N _

pf(a = ^p?-Qj)*pZ(a-, i/j),

hence

{pzj(a, t/j)} -

2.036

2.332

-0.555

-2.115

Assume

Therefore,

{pz(a, M, ip)} ~

2.057

1.979

-0.738

-0.811

tp) =

v(0,At. if»)

2.057

-0.738

Next,

MARKOV-RENEWAL PROGRAMMING -II

43

= ™X2{pf(a,M,if>) + PiZi(M) [f iZi(1/i^) -e01 - v, (0, T,^ x (m, i/i))
+ f1z1(2/^).e2“.v1(0>T1z12(M,^))]

+ PiZ2(M)[fiZ2(l/ip)-eol'V2(0,T1z21(M, *))

+ fz(2/^.e2“.v2(0,Tz2(M^))]}.

From this, Vj(l , /i, i/j) was found to be 2.092. Similarly, v2(l , p, was found to
be -0.736. The calculations were performed for the next successive approximants
and the results are presented in Table 2.

TABLE 2

Summary of Computations for the Example MRD: {l\ F, R, Z }.

n

v(n ,11, ifj)

£(n,j U, ifj)

0

2.057

1

-0.738

1

1

2.092

1

I

-0.736

1

9

2.092

1

z

-0.736

1

LITERATURE CITED

Astrom, K. J., 1965— Optimal control of Markov Processes with incomplete state information.
J. Math. Anal and Appl., 10 : 174.

DeCani, J. S., 1964-A dynamic programming algorithm for embedded Markov chains when
the planning horizon is at infinity. Mgmt. Sci., 10:716.

Ferguson, T. S., 1967 -Mathematical Statistics, A Decision Theoretic Approach. Academic
Press, New York.

Howard, R. A., 1960 -Dynamic Progamming and Markov Processes. MIT Press, Cambridge,
Mass.

Jewell, W. S., 1963 -Markov- renewal Programming I: Formulation, finite return models. Oper.
Res., 11:938.

- , 1963-Markov-renewal Programming II: Infinite return models, example. Oper.

Res., 11:949.

Martin, J. J., 1961 -Bayesian Decision Problems and Markov Chains. John Wiley and Sons,
New York.

SOME COMMENTS CONCERNING THE IMPACT BALL APPARATUS

by C.T. HAYWOOD

Department of Physics, Midwestern State University
Wichita Falls 76308

A typical impact ball apparatus consisting of an arbitrary number of balls in
contact is illustrated in Figure 1 . The usual demonstration is to pull one or more
balls aside, maintaining contact between them, and then to allow these to collide
with the remaining spheres. Despite previous discussions in the literature (Chapman,
1960; Lemon, 1935; Schilling and Yeagley, 1947; Walkiewicz and Newby, 1972),
this apparatus is still misunderstood by many teachers and authors. It is the pur¬
pose of this paper to analyze the simplest case so that physics teachers can give
their students a simple, yet accurate, interpretation of the results.

Quite often the statement is made that a simple application of the principles
of conservation of momentum and conservation of energy are sufficient to define
the motion immediately after the collision. This statement is incorrect. Further¬
more, when the motion is allowed to continue for a few collisions in a “symmetric”
fashion, the observation that things get “messed-up” is usually blamed on the
suspension system. This explanation, at best, is misleading. These explanations
fail to touch on the central point necessary to understand the demonstration.

The case to be examined is shown in Figure 2. The spheres are designated 1,2,3,
left to right. The positive direction is to the right. Sphere 1 has an initial positive
speed v^ just before impact. As shall be discussed, the principles of conservation
of energy and momentum do not guarantee that the speed of Sphere 1 will be zero
after the impact. Its speed after the collision will be designated vlf . Other speeds
are similarly designated. The initial conditions are v2i = v3i = 0. Application of
the conservation conditions and elimination of v1A immediately leads to

Vif =

<v2fHv3f>
v2f+v3f ’

(1)

From this relation, the usual observation is made, i.e., if vif = 0 then either v2f
or v3f must be zero. The constraint that one ball not pass through the other then
dictates that v2f = 0. The result that v3f = v^ then follows. However, this is not
the only solution since positive and negative values of Vff should be considered.
Without loss of generality, the magnitude of vif can be taken to be unity. Then

v2f

v2f =

'3f

1 + v

3f

'3f

1 - v

3f

Case A, vlf = +1

(2)

Case B, vlf = -1

(3)

Accepted for publication: January 14, 1976.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

46

THE TEXAS JOURNAL OF SCIENCE

Figure 2. A head-on perfectly elastic collision in which Ball 1 impacts Ball 2. Balls 2
and 3 are at rest and in contact before the collision.

The graphs for these hyperbolic functions are shown in Figures 3 and 4. The dashed
lines indicate regions of solutions to the equations which do not represent per-
missable motion. In Case A, none of the solutions are permissable since, if vlf is
positive, both v2f and v3f must be positive or Ball 1 would pass through the other
two. In Case B, the regions indicated are not permitted for the following reasons:

48

THE TEXAS JOURNAL OF SCIENCE

Figure 3. Graph of Equation 2, corresponding to vlf = +1. This motion cannot occur.

Quadrant I - Since Ball 2 cannot pass through Ball 3, we require v3f 2L v2f.

Quadrant II - Since vlf is negative, v3f cannot also be negative since this
would mean that all 3 balls would be traveling in the negative
direction after the collision.

Quadrant III - Since vlf is negative, v2f can only be negative if v3f is positive
and v3f > v2f such that the net momentum will be positive
after the collision, no point on the curve satisfies this condition.

It is reasonable to ask if one should have anticipated that vlf could not be positive.
The answer can probably be deduced quite simply by considering the head-on
collision of a ball of mass m with another at rest whose mass is m'. The conser¬
vation principles show that the ball of mass m will rebound if m' > m. In the three-
ball case considered here, one would expect m2 to behave as if its mass were
slightly greater than mq due to the influence of m3 . Thus, rebound of m: is to
be expected.

IMPACT BALL APPARATUS

49

Figure 4. Graph of Equation 3, corresponding to vlf = -1. Only the solid-line branch
represents permitted motion.

The motion observed when the demonstration is performed is that both vlf
and v2f are small. The experiments and calculations of Chapman (1960) show
that the only cases which seem to occur are those for which vlf is small and negative.
Chapman modeled the problem by imagining the spheres to be separated by mass¬
less springs, thus invoking a Hooke’s Law interaction force. The results were
calculated to be vlf = -0. 1 303 vli? v2f = +0.15 03 vli5 and v2f = +0.9800 v^. Chapman
also calculated results using a 3/2 power interaction. The results were qualitatively
the same as for Hooke’s Law. In an experiment involving steel balls, the results
were close to the calculated values.

The application of the principles of conservation of energy and momentum to
the problem of a perfectly elastic collision as indicated in Figure 2 is thus seen to
permit a variety of motions after the collision. These motions are limited to vlf < 0,
v2f>0, and v3f >0. These solutions are, of course, in addition to the single case

50

THE TEXAS JOURNAL OF SCIENCE

vlf = v2f = 0, v3f = vii- In order to uniquely predict the motion, a specific force
law must be assumed. The only case observed to occur, however, is in the asymptotic
region of Figure 4 corresponding to v3f being large.

Since this problem is often misunderstood, it can be instructive to ask students
to analyze it on their own. This suggestion is also made by Walkiewicz and Newby
(1972), but the method of analysis they use, though of a more general nature,
requires more insight on the part of the student. A simple method for giving the
assignment would be to have them derive Equation 1 and then discuss the physically
acceptable motion consistent with the equation.

LITERATURE CITED

Chapman, S., 1960-4ra. J. Phys., 28:705 .

Lemon, H., 1935 -Am. J. Phys., 3:36.

Schilling, Harold K., and Henry Yeagley, 1947 -Am. J. Phys., 15:60.
Walkiewicz, T. A., and N. D. Newby, \912-Am. J. Phys., 40:133.

PETROGRAPHY OF PLEISTOCENE VOLCANIC ASH FROM
PALO DURO CANYON, TEXAS

by PETER JESCHOFNIG

Department of Anthropology , Southern Methodist University,

Dallas 75272

ABSTRACT

Samples from several Palo Duro Canyon volcanic ash outcrops were examined petro-
graphically and particle size distributions were determined through dry-sieving and pipet¬
ting. The examined properties of the glass and the heavy minerals indicate that these ash
lenses correlate with the Pearlette-like ash type O as defined by Izett,ef al.

INTRODUCTION

It was the purpose of this study to identify, petrographically, several Palo
Duro Canyon volcanic ashes and to correlate them with better known ash
beds. On the basis of stratigraphic position, megascopic resemblance, under¬
lying mollusk fauna (Johnston and Savage, 1955) and micro-vertebrate fauna
(Schult, personal communication, 1970), the local ash lenses have been con¬
sidered to be Pearlette ash. To determine the validity of this assumption, a
petrographic examination of a Pearlette ash sample from the Borchers locality
in Meade County, Kansas, was included in this study. To complement the pe¬
trographic findings, grain size distributions were determined for each ash
sample through dry sieving and pipetting.

ASH LOCATIONS

Bull Draw Ash Locality

The main exposure is approximately 50 ft long, 3 Vi ft thick, and is located
on the west side of Bull Draw, Vi mi above its junction with North Cita Creek.
Sample 68L38 was collected from the lower 6 in of the main exposure, while
sample 68L39 was collected from the upper 6 in of the outcrop.

Woody Draw Ash Locality

The ash is exposed in several small canyons near the head of Woody Draw,
a tributary of North Cita Creek. Sample 68L42 was collected from the lower
6 in of the 2 Vi ft thick ash bed.

Accepted for publication: Date unknown.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

52

THE TEXAS JOURNAL OF SCIENCE

Sunday Creek Ash Locality

The main outcrop is 8V2 ft thick and is located near the head of a north
flowing tributary of Sunday Creek. Sample 68L43 was collected from the
lower 6 in of the bed.

Sample 68L44 was collected about 750 ft WNW of locality 68L43 on the
other side of the interfluve, 2-3 ft above the base of the ash bed which is
about 10 ft thick at this locality.

Figure 1. Ash sample locations in Palo Duro Canyon, Texas.

PETROGRAPHY OF VOLCANIC ASH

53

Little Sunday Creek Ash Locality

The ash is exposed near the head of an east flowing tributary of Little Sun¬
day Creek. The sample site is on the east side of a north flowing arroyo which
drains into the tributary. Sample 68L45 was collected 2 ft above the base of
the 2-5 ft thick bed.

All of the above ash locations are in Randall County, Texas, and are shown
in Figure 1.

LABORATORY PROCEDURE

Approximately 400 g of each ash sample were carefully desegregated and
scrubbed ultrasonically in water for 5 min and decanted 5 times after settling
periods of 60 sec. The decanted materials were filtered, dried and studied.
The remaining slurries were poured into large evaporating dishes and dried
in a heat chamber for 2 or more hours.

Before heavy liquid separation, the magnetic fraction was removed by
magnet. The non-magnetic fraction was separated into glass and 2.42+ SG
fractions using a Bromoform-Acetone mixture with a specific gravity of 2.42.
The samples were stirred for 3 min in separatory funnels containing the 2.42
SG Bromoform-Acetone mixture and the ash. After settling periods of 10 min
the heavier fractions were tapped off. This stirring, settling, and tapping pro¬
cess was repeated 3-5 times per separation. Both, the glass and the 2.42+ SG
heavies were rinsed with acetone and dried.

The 2.42+ fraction was again separated into a 2.42-2.8 fraction and a 2.8+
fraction using pure Bromoform with a specific gravity of 2.8. These separat¬
ing processes were the same as described above.

The indices of refraction of the glass were determined by Becke line and
oblique illumination methods. Due to the inavailability of high dispersion
liquids, the more accurate focal masking technique was not used. Glass shards
were examined in form of immersion mounts using microscope slides, oil of
known index and no. 0 cover glasses. All determinations were made with
white light and repeated with a sodium filter (594 nm) placed over the light
source. A thermometer was attached to the microscope stage and readings
were taken before and after each determination in order to compensate for
the refractive index change of the oils due to tempaerature changes.

The shards were studied by traversing the slide on the mechanical stage,
identifying and counting each grain that passed under the crosshair. A mini¬
mum of 300 grains was counted for each slide. Each shard was identified as to
shard type and index of refraction. -The indices were recorded relative to the
oil: nglass <noil, nglass = noil, or nglass >noil.

In smaller grains, it was often difficult to make rapid distinctions between
glass shards and quartz or feldspar, but between crossed nicols and with the
gypsum plate inserted, the separation was made easy by the red color of the

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isotropic shards which contrasted sharply with the yellow and blue inter¬
ference colors of quartz and feldspar. This method also made more prominent
the fine-grained alteration products that coat the glass shards.

Using a Bausch and Lomb binocular microscope, the heavy minerals were
examined and prepared for spindle stage study. The Wilcox spindle stage was
used to identify the minerals and to determine their indices. For minerals
with variable indices like Amphiboles and Pyroxenes, nx was recorded, other¬
wise the minerals were recorded by name only.

Heavy minerals were selected for spindle stage study only if they had glass
adhering to them and if there was no doubt that they were of volcanic, and
not of detrital origin. The index of the adhering glass was also determined to
make sure it matched the index of the other glass shards. The petrographic
work was done with both Leitz and Zeiss polarizing research microscopes.

For the grain size analysis, fresh ash samples were used. About 50-100 g
of each ash sample were desegregated in an ultrasonic waterbath and oven
dried: Each sample was dry- sieved through lA 0 mesh sieves and a Rotap
mechanical shaker. The 44 p fraction was further subdivided through pipetting
into Vi 0 fractions. (Galehouse, 1971).

GLASS

Megascopically, all glass fractions are silver-gray in color, while micro¬
scopically they appear to be somewhat pink under plain light. Few shards
show any alterations, and in some cases, what appears to be alterations may
be clay adhering to the shard. Under the microscope several kinds of glass
shards can be distinguished (Figure 2). The most common ones are plain
plates and plates with one or two ridges, which represent bubble walls and
bubble junctions. Quite common are pumice fragments with elongate vesi¬
cles, while pumice fragments with spherical vesicles and spherical vesicle frag¬
ments are not very common in the Palo Duro Canyon ash. The indices of re¬
fraction range from 1.499 to 1.501 with the majority of the shardshaving an
index of 1.500 (Table 1).

The decanted material consists mainly of clay, very small shards, and un¬
broken glass bubbles, with refractive indices matching those of the shards
(Table 2). The bubbles float and are normally decanted during the clean¬
ing cycle. It is apparently for this reason, that they have not been reported
before.

HEAVY MINERALS

The ash samples 68L38, 68L42, 68L43, and 68L44 contain clinopyroxene
(ferroaugite, nx - 1.730 ± .005) and hornblende (nx = 1.692 ± .002) as dia¬
gnostic minerals, which have not been found in the Borcher’s ash 67L16.

PETROGRAPHY OF VOLCANIC ASH

55

1 . Spherical vesicles and vesicle fragments.

2a. Plain plates.

2b. Plates with one or two ridges (connecting walls).

3a. Pumice fragments with spherical vesicles.

3b. Pumice fragments, fibrous or with elongate vesicles.

Figure 2. Major glass shard types. Modified from Pirsson, 1915.

Chevkinite, allanite, quartz, plagioclase, sanidine, and colorless zircon have
been found in all samples, while pink zircon was found only in the Borcher’s
ash and the Palo Duro ash 68L45. No hornblende was found in sample 68L45.

The most common detrital minerals, which are more or less abundant in
all samples, are epidote, quartz, plagioclase, alkali feldspar, almandine garnet,
hematite, metamorphic hornblende, and augite.

PARTICLE SIZE DISTRIBUTION

As no fresh ash sample was available for the particle size analysis of the
Borchers ash, the size analysis of this ash was performed on decanted remains

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TABLE 1

Indices of Refraction and Shard Types of Glass.

Shard types-%

Plates Pumice fragments

Sample

No.

Index of
refraction

plain

1 or 2
ridges

fibrous or
elongate vesicles

spherical

vesicles

68L38

1.500 ± .001

50

38

12

<.l

68L39

1.500 ± .001

48

40

12

<.l

68L42

1.500 ± .001

47

40

13

<.l

68L43

1.500 ± .001

47

36

17

<.l

68L44

1.500 ±.001

45

40

15

<.l

68L45

1.500 ± .001

47

38

15

<.l

67L16

1.500 ± .001

48

42

10

<.l

TABLE 2
Heavy Minerals.

Sample

No.

Clino-
pyroxene
nY = 1. 730
±.005

Horn¬
blende
nx - 1.692
±.002

Zircon
color- pink
less

Allan-

ite

Chev-

kinite

Plagio-

clase

Quartz

Sani-

dine

68L38

X

X

X

X

X

X

X

X

X

68L39

X

X

X

X

X

X

X

X

o8L42

X

X

X

X

X

X

X

X

68L43

X

X

X

X

X

X

X

X

68L44

X

X

X

X

X

X

X

X

68L45

X

X

X

X

X

X

X

X

67L16

X

X

X

X

X

X

X

x = present

from earlier experiments. Consequently, the very fine fractions of this sample
are missing and the percentages are skewed toward the larger grain sizes. Con¬
sidering the absence of the fine portion of this sample (67L16), the particle
size distribution curve is in fairly good agreement with those of the Palo Duro
ashes.

Table 3 shows the distribtuion of particle sizes by percentage, while
Figure 3 shows the relationship of the various size distribution curves to each
other. The loss in sieving was reasonably small (less than IV2 g per sample) and
was not included in the percentages.

PETROGRAPHY OF VOLCANIC ASH

57

TABLE 3

Particle Size Distribution by % of Weight.

Size

r

68L38

(%-wt)

68L39

(%-wt)

68L42

(%-wt)

68L43

(%-wt)

68L44

(%-wt)

68L45

(%-wt)

67L16

(%-wt)

420

1. 43

1. 28

. 17

. 10

. 07

.41

2. 40

354

. 33

. 17

. 15

. 20

. 07

. 15

2. 07

297

. 33

. 27

. 09

. 28

. 20

.31

3. 35

250

. 53

. 34

. 48

. 10

. 76

. 39

5. 59

210

1. 37

. 84

. 83

. 18

2. 18

. 72

8. 50

177

. 20

. 57

. 63

. 10

3. 34

.75

9. 50

149

. 82

1. 42

2. 17

. 26

7. 29

1. 47

12. 74

125

. 86

1. 42

. 39

. 42

3. 80

. 83

9. 78

105

4. 81

5. 12

4. 74

2. 72

10. 45

2. 89

14. 20

88

7. 66

7. 62

7. 61

5. 69

10. 71

4. 85

9. 50

74

1 1. 85

12. 17

12. 26

9. 82

14. 25

10. 78

9. 33

63

5. 71

6. 47

5. 20

5. 00

6. 02

5. 93

3. 75

53

10. 61

12. 00

10. 33

9. 40

10. 45

12. 74

4. 92

44

14. 93

16. 22

14. 96

14. 31

1 1. 24

18. 29

4. 36

31

14. 13

13. 86

15. 91

9. 76

13. 64

16. 28

-

22

14. 66

15. 17

17. 59

15. 87

2. 77

16. 77

-

16

5. 86

3. 37

6. 48

12. 90

2. 77

6. 45

-

16

3. 91

1. 69

-

12. 90

-

-

-

100. 00

100. 00

99. 99

100. 01

100. 01

100. 01

99. 99

CONCLUSIONS

On the basis of petrographic and chemical data, Izett, et al. , (1970, 1971)
informally classified Pearlette and Pearlette-like ash into types B, S, and 0.

99

95

90

80

70

60

50

40

30

20

10

5

2

1

. 5

. 2

. 1

gurf

typ

j

Di

1.

THE TEXAS JOURNAL OF SCIENCE

1 2 3 4 5 6

Particle Size, ^

3. Relationship of the various size distribution cures to each other.

e O ash is characterized by the presence of hornblende (nx= 1.688-
Lve. 1.690) and clinopyroxene (nx= 1.725-1.734; avg. 1.730). The
ro ashes also contain hornblende (nx= 1 .692) and clinopyroxene
730), and the petrographic properties of the glass fractions are also

PETROGRAPHY OF VOLCANIC ASH

59

identical to those of the type O ash. Therefore, the Palo Duro ashes correlate
quite well with Izett’s type O ash. The Verdos ash of Colorado (Scott, 1963),
Harpole Mesa ash, Upper Onion Creek ash of Utah (Richmond, 1962), and
Cudahy ash of Kansas (Frye, Swineford, and Leonard; 1948) are among the
ash deposits that have been classified as type O (Lambert, personal communi¬
cation, 1970).

The Palo Duro ashes differ greatly from the Borchers ash of Kansas.
While all Palo Duro ashes contain clinopyroxene, and all except sample
68L45 contain hornblende, the Borchers ash contains neither clinopyroxene
nor hornblende, Izett and Wilcox (Izett, personal communication, 1970) have
also examined Bor chefs ash samples and noted the absence of clinopyroxene
and hornblende. Therefore, it seems very unlikely that additional ash separa¬
tions and heavy mineral studies will reveal any of these critical minerals in the
Borchers ash.

The Palo Duro ashes are also very different from the Bishop ash and
from type S ash. The Bishop ash’s most diagnostic features are its chalky
white color. The predominance of minutely pumiceous glass shards with in¬
dices ranging from 1.494-1.496, and the presence of biotite. In comparison,
the Palo Duro ashes are silver-gray in color with bubble walls and junctions as
major glass shards and with higher indices (1 .499-1 .501) than those of the Bis¬
hop ash. The Palo Duro ashes do not contain any biotite, and even the glass
chemistry comparisons show that they are different from the Bishop ash.

The type S ash is distinguishable from the Palo Duro ashes by comparison
of the refractive indices of hornblende, (nx = 1.680 for type S and nx = 1.692
for Palo Duro). The indices of clinopyroxene differ also (nx = 1.720 for
type S, nx = 1.730 for Palo Duro), but more work is needed with clinopy-
roxenes before they can be used conclusively. The chemistry of the glass is also
a good criterion for distinguishing Palo Duro ash from type S ash (Izett, per¬
sonal communication, 1970).

The absence of hornblende in the sample 68L45 is most likely due to an
insufficient size of sample, and it is expected that hornblende will be found in
additional separations.

The Palo Duro ash has not yet been dated, but K-Ar dates on sanidine of
several type O ashes indicate an age of about 0.6 million years.

ACKNOWLEDGMENTS

I wish to thank Frank W. Daugherty, Wayne P. Lambert, and Robert L.
Laury for their suggestions and encouragement in the course of this project.
This research was begun as part of an Lienors’ thesis in geology at West Texas
State University, Canyon, Texas and supported by a NSF undergraduate re¬
search grant.

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LITERATURE CITED

Frye, J. C., A. Swineford, and A. B. Leonard, 194 8 -Correlation of Pleistocene deposits
of the Central Great Plains. J. GeoL, 56(6) :50 1.

Galehouse, J. S., 1971 -Sedimentation analysis. In R. E. Carver (Ed.), Procedu res in
Sedimentary Petrology, pp. 67-94.

Izett, G. A., R. E. Wilcox, H. A. Powers, and G. A. Desborough, 1970-The Bishop Ash
Bed, a Pleistocene marker bed in the western United States. Quaternary Res., 1:121,

- , - , J. D. Obradovich, and R. L. Reynolds, 1971 -Evidence for two

Pearlette-like ash beds in Nebraska and adjoining areas. Paper presented at G.S.A.
meeting, Washington, D.C.

Johnston, C. S. and D. E. Savage, 1955-A survey of various late Cenozoic vertebrate
faunas of the Panhandle of Texas, Part 1, Univ. Calif. Pub. Geol. Sci., 3 1(2) :46 .

Naeser, C. W., G. A. Izett, and R. E. Wilcox, 1971-Zircon fission track ages of Pearlette-
like volcanic ash beds in the Great Plains. Paper presented at G.S.A. Annual Meeting,
Washington, D.C.

Pirsson, L. V., 19 15 -The microscopical characters of volcanic tuffs-a study for stu¬
dents, Amer. J. Sci., 4th series, 40(236): 191.

Richmond, G. M., 1962-Quaternary stratigraphy of the La Sal Mountains, Utah, U.S.
Geol. Sur. Prof. Pap., 324.

Scott, G. R., 1963-Quaternary geology and geomorphic history of the Kassler quadran¬
gle, Colorado, U.S. Geol. Sur. Prof Pap., 412A.

APPLICATION OF HUYGENS’ PRINCIPLE TO THE REFLECTION
OF SEISMIC WAVES AT A FREE SURFACE

by ETHEL WARD McLEMORE

Geophysicist, 11625 Wander Lane, Dallas 75230

and IRA L. WRIGHT

Consulting Systems Representative, IBM, Dallas 75230

ABSTRACT

This paper deals with the problem of a time-domain solution of a reflected spherical wave
from a free surface. It is a study of the properties of the coefficients of the wave functions
of the reflected compressional and shear waves in Kirchhoff’s surface integral, developed by
application of Huygens’ Principle to a plane wave approximation of the velocity potentials.
A technique for location of the secondary source in the form of equal-time rings on the
reflecting surface is developed. The values of the coefficients of the first derivatives in the
integral are insignificant as compared with those of the second derivatives, for both waves.
The larger magnitudes occur immediately after reflection, and the largest when the observa¬
tion point and the source point are on the same vertical. In this case the time rings are con¬
centric circles with center and reflection point at the origin. When the observation point is
near the reflecting surface and the origin, the magnitudes of the compressional wave coeffi¬
cients are greater than those of the shear wave. As the observation point is moved away from
the origin and the vertical, the magnitudes of the shear wave coefficients are greater than
those of the compressional wave.

INTRODUCTION

The purpose of this study has been the application of Huygens’ Principle by
use of Kirchhoff’s integral to a plane wave approximation of compressional wave
on a stress free surface.

In general terms Huygens’ Principle states that if a source 0 is surrounded by
a closed surface S, the various points of the surface S, when reached by the wave
of the disturbance, become the origin of secondary waves, and the disturbance
observed beyond the surface S results from the super-position of these secondary
waves (Rossi, 1959). The Principle states that the disturbance at some later time,
t = t0 + At, at a given observation point, can be obtained by considering the effect
produced by each point on a wave surface at a given time, t = t0 , acting as a sec¬
ondary source (Officer, 1958).

Accepted for publication: May 19, 1976.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

62

THE TEXAS JOURNAL OF SCIENCE

Huygens published his geometrical theory of wave propagation in optics, based
on an analogy between the propagation of sound waves of small amplitude and
the propagation of light, in 1690. Erroneously, he treated a light wave as a scalar
phenomenon. Much of the work in his Treatise was vague, provoking a controversy
which has lasted for may years and resulted in a wealth of published work on
wave propagation.

A solution of the problem of transient elastic wave propagation through use
of the Laplace transform was developed by Cagniard (1930). The transform
methods give an answer, but not in the case of non-horizontal layers, nor do they
give an understanding of the mechanism involved. Use of Kirchhoffs formula in
the application of Huygens’ Principle gives one a greater grasp of the physics of
the problem.

Burridge (1963) made an exhaustive study of the reflection of a pulse in a solid
sphere, using Kirchhoffs formula, and defined the pulse shape.

The application of Huygens’ Principle to problems of elastic diffraction, using
Kirchhoffs formula, has been investigated recently by Trorey (1970). He treated
only the compressional wave.

This work is composed of 3 parts:

1. Development of the coefficients of the wave functions through substitution
of given boundary conditions in Kirchhoffs surface integral (Baker and Copson,
1939).

2. Location of the secondary source reflecting surface, S, in the form of equal¬
time rings, for any given observation point within the medium bounded by S,
but not on S.

3. Mapping of the secondary source rings and the values of the coefficients of
the wave functions as time passes, for any given observation point, depth of
source point, and velocity ratio; with numerical examples for varying positions
of the observation point.

COEFFICIENTS OF THE WAVE FUNCTIONS IN
KIRCHHOFF’S SURFACE INTEGRAL

Let us assume a seismic pulse generated by a sudden application of unit pres¬
sure, such as the explosion of dynamite in a hole in the ground; in a spherical
cavity of such relatively small radius that we treat the source hole as a point source
under a plane boundary S. At any time t after the explosion an observer, situated
at a point Q within the medium bounded by S will “see” the reflected image of
rays diverging from the simple source at a point O' (Figure 1); their intersection
with the surface S taking the form of a closed curve. From this section of the sur¬
face plane, acting as a secondary source, rays converge at Q, which is the “eye”
of the “observer.”

We use rectilinear coordinates with the surface location of the source point as
origin and the point Q in the xz-plane with the point 0. We take the z-axis of
our configuration as positive downward, the inward normal to S.

APPLICATION OF HUYGENS’ PRINCIPLE

63

IMAGE OF POINT SOURCE

o1

Figure 1. A sketch illustrating the point source image concept.

Kirchh off’s formula which follows (Baker and Copson, 1939) states a general
theorem concerning solutions of the scalar wave equation with constant coefficients:
Let u(x,y, z, t) be a solution of the equation

V2u = (1/V2 )62u/6t2 (V2 is Laplace’s operator)

whose partial derivatives of the first and second orders are continuous within and
on a closed surface S, and let (xj , yq , zx ) be a point within S.

Then,

uOi,yi>zi,t) =

(l/4Tr)//{|u|6(l/r)/6n -(l/Vr)6r/6n|6u/6t|- (l/r)|6u/6n|}dS, (1)

where r is the distance from (xj , yl , zx) to a typical point of S, 6/6 n denotes
differentiation along the inward normal to S, square brackets ( | | ) indirate retarded
values, and V denotes wave velocity.

If, however, the point (xx iyl,zl) lies outside S, the value of the integral is zero.

64

THE TEXAS JOURNAL OF SCIENCE

We define the retarded value of a function u at a variable point P relative to a
fixed point Q, when the distance QP is r, as

| u | = u(x, y, z, t-r/V). (2)

We assume a general form of the incident dilatational wave generated by the
explosion at the source 0(0, 0, d) and set the scalar potential Aj for the incident
wave equals fj(Vct - r0)/r0, where fj is some dimensionless function of the argu¬
ment (Vct -r0 ), and r0 is the distance from the point 0(0, 0, d) to a point P(x,y, z«e),
in the direction of the propagation of the incident wave, on the surface S (Figures 2-1
and 2-2). We use the very small figure e to indicate the proximity of Kirchhoff’s
surface S to our xy-plane of the earth’s surface. Kirchhoff’s S is a mathematical
surface and the integral is discontinuous at the boundary. In setting z = 0 after
differentiation, we are approximating the values of the coefficients in the integral.

Figure 2-1. Geometry of the data used in the 3 numerical examples for the compressional
wave equal- time rings.

APPLICATION OF HUYGENS’ PRINCIPLE

65

100

100

Figure 2-2. Geometry of the incident compressional wave originating at point source
0(0, 0, d) and reflected from surface S at a different point P(x, 0, 0) with the
reflected shear wave arriving at point Q(Xp 0, zp.

Let us approximate the spherical wave at P(x, y, z« e) by a plane wave, and
in differentiating in the direction of the normal require that r0 be constant for a
given P, but allow both the amplitude and direction to vary for different points on S.

Now, consider the reflection of a plane compressional wave incident on a free
surface which bounds an isotropic, homogeneous, perfectly elastic medium occupy¬
ing a half space. We consider only the energies of the 2 independent reflected waves,
the compressional and the shear. To develop the scalar potential, Ar(xl5 yl5 zl5 1),
for the reflected compressional wave at Q(xj , yx , zx) we substitute H(I)Aj for the
u of Kirchhoff’s formula, z for n, rt for r, and Vc for V (Figure 2-1). To develop
the vector potential, R(xj , y x , zl , t), for the reflected shear wave, we substitute
G(I)Aj for u, z for n, r/ forr, and Vs for V (Figure 2-2). The functions H(I) and
G(I) are the compressional and shear wave reflection coefficients for incident
dilatational waves, respectively, for a free surface (McLemore and Wright, 1976).

66

THE TEXAS JOURNAL OE SCIENCE

Then ,

Ar(x, > Yi , ?i . t) - (l/41i)//{|H(I)Ai|6(l/i1)/6z

- (l/Vcri)6r,/6z | 6(H(I)Ai)/6t |

- (1/r, ) 1 6(H(I) A,)/ 6z | } dS; (3)

and

R(xljy;,Zl,t) = (1/4tt)//{ |G(I)Aj | 6(l/i'[)/ 6z

- (■l/Vfr1')6r17«i|6(G(I)Ai)/6t|

- (1/r/) | 6(G(I)Ai)/6z | }dS. (4)

In making the substitutions of these reflection coefficients it should be under¬
stood that Kirchhoff ’s formula applies so long as the path structure is large compared
with the wave length.

Performing the indicated differentiation in (3) and (4), see Appendix A, and
substituting our definition of retarded values in (2), we get

Ar(xi>yi>zi>t) - (l/4Tt)//(l/r0r,){(H(I)(CosI/r0 +Cos 0/r, )

+ (dH(I)/dI)(Sin I/r0)) .fj(Vct - (r0 + r, ))

+ (H(I)(Cos I + Cos 0) *fj(Vct - (r0 + r, ))} dS; (5)
and

R(Xi ,yi , Z] , t) a (l/4Tr)//(l/r0r,'){(G(I)(CosI/r0 +Cos B/r/ )

+ (dG(I)/dl)(Sin I/r0)) .fj(Vct - (r0 + ar/ ))

+ G(I)(Cos I + aCos 6)-f/(Vct-(r0 +ar,'))}dS, (6)

where a = Vc/Vs, the ratio of the reflected compressional wave velocity to the
reflected shear wave velocity.

From (5) we get the coefficient of the wave function f^V^t - (rQ + y{ )) for the
reflected compressional wave as

FH = (l/r0 r, ){(H(I)(Cos I/r0 + Cos 0/r, ) + (dH(I)/dI)Sin I/r0) } , (7)

and the coefficient of the wave function fV(V t - (rQ + r} )) as

FPH - (l/r0r1 ){H(I)(Cos I + Cos 0)} . (8)

From (6) we get the coefficient of the wave function f.(V t - (rQ + ar^)) for
the reflected shear wave as

FG - (l/r0r;){(G(I)(CosI/r0 +Cos 3/r/) + (dG(I)/dI)Sin I/rQ) } ,

(9)

APPLICATION OF HUYGENS’ PRINCIPLE

67

and the coefficient of the wave function f/(Vct - (rQ + ar^)) as

FPG = (l/r0r/){G(I)(Cos I + aCos 3)} . (10)

As can be seen from the graphs of the numerical examples which follow
(Figures 4-1 and 5-1), the values of the coefficients FH and FG are insignificant
as compared with the values of FPH and FPG. For a First approximation we write
for (5) and (6):

A/x^y^Zj, t) - (l/4'n)//{(l/r0r1 )H(I)(CosI + Cos 0)*F(\^t -(rQ+rj^JdS; (5.1)
and

=(l/4Tf)//{(l/r0r;)G(I)(CosI+aCos eH/(Vj-(r0+ar/))}dS; (6.1)

where

t t/t \ _ 4Sin33Cos 3Cos I - SinICos* I 22 3
H(l) — — -

4Sin3BCosgCos I + SinICos22B

(5.2)

and

G(I)

4Sm 3Cos 2 3Cos

4Sin33Cos 30osl + Sin ICos22 3

dH(I)/dI =

(8Sin33Cos23) ("2Cos23Cos2I(l + 2Sin23)-Cos2 3(Sin23Cos2I+Sin2ICos23)
Cos 3 ( (4Sin33Cos3CosI + SinICos223)2

and

dG(l)/dl =

4 Sin 2 3 ( 4Sin33Cos3I - Sin I Cos2 2 3Cos 3(Cos 21 + 2Sin23)

(6.2)

■(5.3)

SinIC

os3 (

(4Sin33Cos3CosI + Sin ICos223)2

; (6.3)

where

I is the angle between and rQ ,

0 is the angle between and ,

3 is the angle between zx and r
and

a is the ratio of the reflected compressional wave velocity to the reflected shear
wave velocity.

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THE TEXAS JOURNAL OF SCIENCE

LOCI OF EQUAL-TIME RINGS ON THE SURFACE S

We are interested in the location of the secondary source points on the reflecting
surface S of rays arriving at Q in phase. The phase, (V t-(rQ + rl )) or (V t-(rQ + ar^)),
of a ray arriving at Q depends on time, position of the source, position of Q,
and retardation — making the problem essentially one of geometrical analysis. For
any time t these two expressions are constants, and, as the velocities are constants,
for any time t the expressions (rQ + r1 ) and (rQ + ar^) are other constants.

Equal- time Rings for the Compressional Wave

The expression Vt -(rQ +r1) = 0 represents a family of confocal prolate spheroids,
or ellipsoids of re volution, with foci at 0 and Q. The intersection of this 3 -dimen¬
sional system of curves with the earth‘s surface — our S as the plane z = 0 — is a
family of eccentric ellipses, symmetrical with respect to the x-axis (Figure 3).
These curves define the loci on the reflecting surface S corresponding to equal¬
time rings for the reflected compressional waves (Rayleigh, 1877). Their eccentric¬
ity depends on the ratio of the depth of the point Q to the depth of the point 0,
and on the xx coordinate.

Figure 3. XZ-Cross- section of the ellipsoids with perspective view of the corresponding
compressional wave equal-time rings.

APPLICATION OF HUYGENS’ PRINCIPLE

69

The larger the ellipsoids the later the arrival. The earliest arrival corresponds
to the ellipsoid just tangent to the surface. This point of tangency is the ordinary
reflection point, as the tangent plane to an ellipsoid makes equal angles with the
focal radii at the point of tangency, and the angle of incidence is equal to the
angle of reflection of the compressional wave at this point. The rings are concen¬
tric when the fociO and Q are at equal distances from the reflecting surface, with
center at the reflection point equal to l/2xr For vct» OQ, the ellipses become
concentric circles, with center equal to l/2xr If the Points 0 and Q are on the
same vertical, the rings are concentric circles with the reflection point and center
at the origin.

If we let (rQ + q ) = k, and substitute the coordinate values of rQ and q, rationali¬
zation yields (Appendix B, Figure 2-1)

(x>l/2XlK/(k2-x^))2 + (k2/(k2-Xj))y2 = (k2/4(k2-x2)2)(K2^4d2(k2-x2)), (11)

where

K = ((k2 -x2 ) + (d2 -z2 )) .> d2.

Equation (1 1) defines a system of ellipses with center at
((l/2x,K)/(k2-x?),0,0),
major semi- axis equal to

(k/2(k2 - x2 ))(K2 - 4d2 (k2 - x2 ))1/2 ,
and minor semi- axis equal to

( l/2(k2 - x2 ) 1/2 )(K2 - 4d2 (k2 - x2 )) 1/2 .

The value of the parameter k for the reflection point, which we denote by kQ is
k0 - (x2 + (z1 + d)2)1/2, (12)

and the reflection point is at the value of x for which
K2 = 4d2(k2 - x2),

or,

x = (d/(d + Zl))xr (13)

If we let Ak represent a constant difference between succeeding values of k,
then a unit change of Ak in k is equal to a unit change in distance corresponding
to a unit change in retardation of time. For consistency in comparing potential
values at various observation points, Q(xx , 0, zx ), we can set

Ak = k0 - OQ

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THE TEXAS JOURNAL OF SCIENCE

and

k = kA + n Ak, where n =1 , 2, 3, . . . .

However, as most of the energy represented by the functions occurs immediately
after reflection, we can locate the maximum effect by subdividing the first interval
(n 1), making the unit change of time as small as our computation errors permit.
Since kQ corresponds to the time of reflection, we begin the computations with
kQ + 1 as representing the first ring.

Equal- time Rings for the Shear Wave

The expression V t - (rQ + ar^ ) = 0 represents a system of surfaces of revolution,
or space quartics, with foci at the Points 0 and Q. The intersection of these 3-
dimensional surfaces with the reflecting surface S is a system of 2 quadratics which
do not touch, or a biquadratic, symmetrical with respect to the x-axis.

If we let rQ + ar^ = k' and substitute the coordinate values of rQ and r^ for the
shear wave, rationalization yields (Appendix B, Figure 2-2)

x4 + 2Dx3 + (D2 - E)x2 + Fx + G + y 4 + (2x2 + 2Dx - E)y 2 = 0, (1 4)

where

D = -wa2x1, E = wK' + w2k'2, F = -wK'D,

G = (w2K'2/4) - w2k'2d, w = 2/(1 -a2),

K' = ((k'2 -a2x2) + (d2 -a2z2)) > d2 ,
and, by definition,

a = v/v.

c s

We solve this quartic for y in terms of given values of x. Setting the expression
x4 + 2Dx3 + (D2 - E)x2 + Fx + G = A,
and the expression

2x2 + 2Dx - E = B,
equation (14) becomes

y4 + By2 + A = 0, (15)

a biquadratic system for the parameters k' and a, containing the 2 quadratics
y2 = (-B - (B2 - 4 A) 1/2 )/2
and

y2 = (-B + (B2 - 4 A) 1/2 )/2.

APPLICATION OF HUYGENS’ PRINCIPLE

71

If B2 - 4A >. 0, the roots are real and these 2 quadratics, or one of them, represent
the equal time rings for the shear wave on the surface S.

In our computations we found that the values of x and y which satisfy the
quadratic

y2 = (-B + (B2 - 4 A) 1/2 )/2

for real roots do not satisfy the original equation rQ + ar^ = k', for the values of k'
used. We found that the values of x and y which satisfy the quadratic

y2 = (-B -(B2 - 4 A) 1/2 )/2

for real roots do satisfy the original equation for the values of k' used. We con¬
cluded that, for our problem, the first quadratic has no physical meaning, and
that the equal-time rings which we seek for the reflected shear wave are represented
by the equation

y2 = (-B - (B2 -4A)1/2)/2, (16)

a family of closed curves symmetrical with respect to the x-axis, which appear
on the graph as slightly distorted ellipses. When the Points 0 and Q are on the same
vertical, the rings are concentric circles with the center and reflection point at
the origin.

The shear rings are larger than the compressional rings for the same point Q(x1,0,z1 )
because the value k' is greater than the value k for the compressional waves, which
means that the shear waves arrive at Q at a later time.

The reflection point P(x, 0, 0) for the shear wave at Q(xL ,0,2^ is

x = (d/(d + z1))x2, (17)

where x2 is a coordinate of a point Q'^ , 0, ) for the compressional wave cor¬

responding to the Point Q(Xj , 0, z2 ) for the reflected shear wave. We do not know
the value of x2 corresponding to the value x: for the shear wave. We cannot use
Equation (17) to find the reflection point.

To locate the reflection point P(x, 0, 0) for the shear wave at Q(x2 , 0, z2 ) we
use Snell’s Law which states that

Sin 0/V = Sin 3/V (In this case 0 is the angle between the

normal n and Q'P.)

At the reflection point, Sin 0 = Sin I = x/r and Sin 3 = (Xj - x)/q (Figure 2-2).
Substituting, we get x/arQ = (xx - x)/^', where a = V /V .

Rationalization (Appendix B) yields

x4 - 2Xj x3 + (C(z2 - a2 d2 ) + x2 )x2 + 2Ca2 d2 x: x - Ca2 d2 x2 = 0, (18)

where

c = 1/(1 - a2).

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The reflection point is that value of x, of this quartic (18), which is less than
x and greater than (d/(d + z1))xr

The value of k' at the reflection point when x has the value of (17) is

= (x2 + d2)1/2 + a((Xj - x)2 + z2)1/2. (19)

We set

Ak' = k; - OQ ,
and

kn = k^ + n Ak', where n = 1,2,3,....

Using equations (7), (8), (1 1), (12), and (13) for the reflected compressional
wave and equations (9), (10), (14), (15), (16), (17), (18), and (19) for the reflected
shear wave, we wrote a general FORTRAN IV program for computation of a plane
wave approximation of the velocity potentials created by the reflection of a
spherical incident compressional wave on a stress free surface.

Figures 4, 4-1, 5, and 5-1 show 2 numerical examples of the reflecting surface
S and the resulting values of the coefficients of the wave functions, at a given point
within the medium through which the sound waves travel as time passes. The
coefficients assume the greatest magnitude when the source point is on the same
vertical with the observation point. When the observation point is near the reflecting
surface and the origin, the compressional wave coefficients are greater than the
shear wave coefficients. As the observation point is moved away from the origin
and the reflecting surface, the shear wave coefficients are greater than those of
the compressional wave. Most of the effect occurs immediately after reflection.

Figure 4. (See page 73.) Equal-time rings on surface S for the 2 positions of the obser¬
vation point Q(10, 0, 10) and Q(200, 0, 200), with d = 100 feet and velocity
ratio a = (3)1/2. Note the compressional wave.

Figure 4-1. (See page 74.) Compressional wave coefficients corresponding to rings of
Figure 4.

Figure 5. (See page 75.) Equal- time rings on surface S for the 2 positions of observation
point Q(10, 0, 10) and Q (200, 0, 200), with d = 100 feet and velocity ratio a = (3)1/2.
For the shear wave.

Figure 5-1 . (See page 76.) Shear wave coefficients corresponding to rings of Figure 5.

APPLICATION OF HUYGENS’ PRINCIPLE

73

x

REFLECTION PD I NT x I ' REFLECTION POINT

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THE TEXAS JOURNAL OF SCIENCE

- FPH

For Position of Observation Point Q(200, 0, 200)

X

INI

o

cno

UJ CM
>

cr ii

r* — <

x

cr ii
UJUJ

x o
cn

Figure 5. (See legend, page 72.)

REFLECTION PD I NT x ' REFLECTION POINT

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THE TEXAS JOURNAL OF SCIENCE

APPLICATION OF HUYGENS’ PRINCIPLE

77

APPENDIX A

The Differentiation in (3), Ar(Xj ,yl,zl, t):

rj - ((x-x1)2+y2+(z~z1)2)1/2.

Aj = f(Vc-r0)/r0.

<5(l/r,)/6z = (— 1/r j )6r t / 6z-

6r,/Sz = 6((x - Xj)2 + (z -Zj)2)1/2/6z

= l/2((x - Xj)2 + y2 +(z-z1)2)'1,2-2(z-z1)

= (z-z1)/r1

= -Cos 0 (when z - 0).

6(l/r,)/6z = (l/r^ )Cos 8.

H(I) Aj |6(l/r,)/6z | = (l/r0r,) •H(I)(Cos Bh1 • |fi(Vct - r„) |.
r0 = (x2 +y2 +(z-d)2)1/2
6(H(I)Aj)/6t = H(I)6Aj/6t + Aj6H(I)/6t.

6Aj/6t = 6((l/r0)fi(Vct-r0))/6t
= (l/r0)6(fi(Vct-i0))/6t
= (l/r0)-Vc.fi'(Vct-r0),

where fj is the derivative of fj with respect to the argument.

6H(I)/6t = 0 (H(I) contains no t).

6(H(I)Aj)/6t = (l/ro)-H(I)-Vc-fi'(Vct-r0).

-(l/Vcr1)6r1/6z|6(H(I)Ai)/6t| = (l/^i^HflJ-Cos e>fj(Vct - r0).

— ( 1/r , ) |6(H(I)Ai)/6z| = — ( 1 /r j ) |Ai6H(I)/6z + H(I)6Ai/6z |.

6(H(I))/6z + (dH(I)/dI/dz) = -(Sin I/r0)dH(I)/dl.

Aj6H(I)/6z = (-(SinI/r0)dH(I)/dI)-fi(Vct-r0)/r0.

H(I)6Aj/6z = (H(I))6((l/r0)fj(Vct-r0»-6z

= (H(I))fi(Vct - r0)6(l/r0)/6z + (H(I))6fi(Vct - r0)/6z -( l/rQ).
6(l/r0)/6z = -(l/r2)6r0/6z

= -(l/r2)*l/2(x2 + y2 + (z - d)2)"I/2.2(z -d)

= -d/r2)*(z-d)/r0
= -(l/r2)CosI (when z = 0).

H(I)fj(\;t - r0)6(l/rQ)/6z = -(l/r0)<H(I).(CosI/r0).fi(Vct-r0).

6fi(Vcl - r0)/6z = 6(Vct - r0)/6z -f/ (Vct - r0)

= -CosI.f;(Vct-r0).

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THE TEXAS JOURNAL OF SCIENCE

Appendix A (Continued)

H(I)6fj(Vct-r0)/6z*(l/r0) = -(l/r0)H(I) -Cos I .f/(Vct - rQ).

H(I)6Aj/& = -(W0)-H(I)-(CosI/r0)-fi(Vct-r0)-(l/r0)-H(I)*CosI-fi'(Vct-r0).
— (1/r j ) |6(H(I)/\j)/6z| = (l/r0rI)((dH(I)/dI)(Sin I/rQ)* |f;(Vct - r0) |

+ (l/^r^HOKCos I/r0). |fj<Vct - r0) |

+ (l/r0r1)H(I)CosI- |f/(Vct-i0) |.

Substituting these values for the three terms in (3) and substituting our value of retarded
time (2) where t = t - ^l/'Vc, we get Equation (5), page 66;

|fi<Vct-r„)| = fi(Vct-(t0 +ri)),

where

jfi'(Vct - r0) f = f/(Vc -(!„+!,)).

— >

The Differentiation in (4) R(x1 , y 2 , t):

rj = (x -x^2 + y2 + (z -Zl)2)l/2.

6(l/rJ)/6z = (-l/rj2)6rj/6z.

6rJ/6z = (z — Zj )/r j

= -Cos 3 (when z = 0).

6(l/rj)/6z = (l/rJ2)Cos 3.

|G(I) Aj 1 6(l/r |)/6z | = (l/r0rj)*G(I)(Cos 3/r')- |fj(Vct - rQ) j.

6(G(I)Ai)/6t = G(I)6Aj/6t + Aj6G(I)/6t.

6Aj/6t = (l/r0)*Vc-f/(Vct-r0).

6G(I)/6t = 0 (G(I) contains no t).

6(G(I)Aj)/6t = (l/r0)*G(I)*Vc.fj'(Vct-r0).

-(l/\^r')6r'/5z |6(G(I)Ai)/%| = (l/r0r;)*G(IKVc/Vs)-Cos |f/(Vct - rQ) |.

-<l/r') |6(G(I) Ai)/6z | = -(1/r j) | Aj6G(I)/6z + G(l)6Aj/6z |.

6(G(I))/6z = (dG(I)/dI)(dI/dz)

= -(Sin I/r0)dG(I)/dI.

Aj6G(I)/6z = (-(SinI/r0)dG(I)/dI)-fi(Vct-r0)/r0.

G(I)6A,/6z = (G(I))6((l/i0)fi(Vct-r0)>/Sz

= (G(I))fj(Vct - r0)6(l/r0)/6z + (G(I))6fi(Vct - rQ)/6z-( 1 /rQ) .
6(l/r0)/6z = -(l/r„)CosI (when z = 0).

G(I)fi(Vct-r0)6(l/r0)/6z = -( l/r0)-G(I> -(Cos I/r0)-fj(Vct - r0).

6fj(Vct - r0)/6z = -CosI»f/(Vct-r0).

APPLICATION OF HUYGENS’ PRINCIPLE

79

Appendix A (Continued)

G(I)6fi(Vct-r0)/6z-(l/r0) = -(l/r0)G(I)-Cos - rQ).

G(I)6Ai/6z = -(l/r0)*G(IHCos I/r0Hj(Vct - - (l/r0KKD*CosI*f/(Vct-r0).

-d/r') 1 6(G(I) Aj)/6z | = (l/r0rJ)((dG(I)/dI)(Sin I/r„)- |fj(Vct-i0) |

+ (l/i0rJ)G(I)(Cos I/r0)- |fj(Vct -r0) |

+ d/r0rJ)G(I)Cos I* |f/(Vct -r0) |.

Substituting these values for the 3 terms in (4) and substituting our value of retarded time (2)
where t = (t - r^), we get Equation (6), page 66;

|fi(Vct-r0)| = fi(Vct-(x0+ar;)),

where

|fi'(Vct - r0) I - fi'(Vct - (r0 + arj)),

and

a = Vc/Vs.

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THE TEXAS JOURNAL OF SCIENCE

APPENDIX B
THE RATIONALIZATION

Compressional Wave

r0 = (x2 + y2 + d2)y2
rj = ((x-Xl)2 +y2 +z2)1/2
r0 + ri = k

ri = k“ro

r2 = k2 - 2kr0 + r2

x2 - 2xj x + x2 + y2 + z2 - x2 - y2 - d2 - k2 = -2kr0
-2xj x + x2 + z2 - d2 - k2 = -2kr0
Set K = ((k2 - x2 ) + (d2 - z2 ))

2X]lx + K = 2k(x2 + y2 +d2)1/2

4x2x2 +4Kx1x + K2 = 4k2 x2 +4k2y2 +4k2d2

(k2 -x2)x2 - Kxxx + k2y2 = K2/4 -k2d2

x2 - (KXl/(k2- x2 ))x + (k2/(k2 - x2 ))y2 = K2/4(k2 - x2 ) - k2d2/(k2 - x2 )
x2 - 2(KXl/2(k2 - x2 ))x + K2x2/4(k2 - x2 )2 + (k2/(k2 - x2))y2
= K2/4(k2 -x2) -k2d2/(k2 - x2 ) + K2x2/4(k2 -x2)2.

(x - Kxj/2(k2 - x2 ))2 + (k2/(k2 - x2 ))y2

= (K2 - 4k2 d2 )/4(k2 -■ x2 ) + K2x2 /4(k2 - x2 )2 .

(x-Kxj/2(k2 -x2))2 +(k2/(k2 - x2 ))y2

= (k2/4(k2 - x2 )2 ) (K2 - 4d2(k2 - x2 )). (11)

Shear Wave

r0 = (x2 +y2 +d2)1/2
rj = ((x-x1)2+y2+z2)1/2

t /

r0 + arx = k
arj = k'-r0
a2r2 = k'2 - 2k r0 + rj

a2x2 - 2a2Xj x + a2x2 + a2y2 + a2z2 - k'2 - x2 -y2 - d2 = ~2k,r0.

(a2 - l)x2 + (a2 - l)y2 - 2a2XjX + a2x2 + a2z2 - k'2 - d2 = -2k rQ.

Set K# = ((k^2 - a2x2 ) + (d2 - a2z2 ))

(a2 - l)x2 + (a2 - l)y2 - 2a2x2x - K* = -2kf(x2 + y2 + d2 )1/2.

Set w = 2/(a 2 - 1)

(x2 + y2 ) - wa2XjX - wK//2 = -wk^x2 + y2 + d2 )1/2

APPLICATION OF HUYGENS’ PRINCIPLE

81

Appendix B (Continued)

(x2 +y2)2 + 2(x2 + y2)(-wa2x1x -wK72) + (-wa2x1x - wK'/2)2
= wV2x2+w2k'2y2+w2k'2d2.

x4 - 2wa2x1x3 +(w2a4x2 - wK* - wV2 )x2 + w2a2XjKx + (wV2/4) - wV2d2
+ (2x2 -2wa2x1x-wK' -w2k'2)y2 + y4 = 0.

Set D = -wa2Xj ,

E = wK' + wV2,

F = -wK'D,

G = (w2k'2/4) - w2k'2d2.

Then

x4 + 2Dx3 + (D2 - E)x2 + Fx + G + (2x2 + 2Dx - E)y2 + y4 =0. (14)

The Reflection Point for the Shear Wave
r0 - (x2+d2)1/2
r; = «x-Xl)2- z])112
Sin 3 = (l/a)Sin 0
= (l/a)Sin I
= (Xj -x)/rj
x/ar0 = (xL - x)/r|

x2/a2(x2 + d2 ) = (x2 - 2xjX + x2)/(x2 - 2xjX + x2.+ z2)

(1 -a2)x4 - 2X]l(l -a2)x3+(z2-a2d2 + (l-a2)x2)x2 + 2a2d2x1x-a2d2x2 = 0.
SetC = 1/(1 - a2)

x4 -2Xlx3 +(C(z2 -a2d2) + x2)x2 + 2Ca2d2Xlx -Ca2d2x2 = 0. (18)

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NOTATION

S is a plane surface taken as the xy-plane. For the approximation we regard the sur¬

face S as coinciding with the earth’s surface.

0(0, 0,d) is the source point of the explosion or disturbance and d is depth of source. The
dimensions of the source are small compared with other distances involved.

Q(x1? 0, Zj) is an observation point within the medium bounded by S.

P(x, y, 0) is any point on the surface S.

r0 is the distance OP.

rt is the distance QP for the reflected compressional wave,

rj is the distance QP for the reflected shear wave,

n is the normal into the medium.

I is the angle between n and r0.

0 is the angle between n and rr

3 is the angle between n and r{.

is the compressional wave velocity in ft/sec.
is the shear wave velocity in ft/sec.
a is Poisson’s ratio,

a is the ratio \/\

H(I) is the reflection coefficient of the compressional wave at the free surface, and is

a function of Ws, and I.

G(I) is the reflection coefficient of the shear wave at the free surface, and is a function

ofVc,Vs, and I.

Aj is the scalar potential for the incident wave.

Ar is the scalar potential for the reflected compressional wave.

— ^

R is the vector potential for the reflected shear wave,

fj is some dimensionless function of the given argument,

fj is the derivative of f| with respect to the given argument.

FH is the coefficient of the wave function fj(\^t - (r0 + rj)).

FPH is the coefficient of the wave function fj(\^t - (r0 + r^).

FG is the coefficient of the wave function fj(\^t - (r0 + ar/)).

FPG is the coefficient of the wave function fj(Vct - (r0 + arj )).

FH2 is the value of FH for a given time.

FPH2 is the value of FPH for a given time.

FG2 is the value of FG for a given time.

FPG2 is the value of FPG for a given time.

APPLICATION OF HUYGENS’ PRINCIPLE

83

LITERATURE CITED

Baker, Beran B., and E. T. Copson, 1939 -The Mathematical Theory of Huygens’ Principle.
The Claredon Press, London.

Burridge, R., 1963-The reflection of a pulse in a solid sphere. Proc. of the Royal Society, A,
276:367.

Cagniard, L., (1930), 19 62 -Translation by Edward A. Flinn and C. Hewitt Dix, Reflection
and Refraction of Progressive Seismic Waves. McGraw-Hill, New York, N.Y.

McLemore, Ethel Ward, and Ira L. Wright, 1976-Reflection coefficients and their derivatives.
Tex. J. Sci., 27(3) : 3 3 1 .

Officer, C. B., 1958 -Introduction to the Theory of Sound Transmission. McGraw-Hill,
New York, N.Y., pp. 269-274.

Raleigh, J.W. S., \%11 -Theory of Sound. Dover 1945 Edition, Vol. 2, New York , N.Y. , pp. 119-126.
Rossi, Bruno, 1959 -Optics. Addison-Wesley Pub. Co., Reading (2nd printing), p. 19.
Trorey, A. W., 1970-A simple theory for seismic diffractions. Geophys. , 35:762.

■

THE EFFECT OF LYSOSOMAL ENZYMES ON CHLOROPLASTS
IN VITRO

by J. AUNE, BARBARA JEAN SMITH, and M. M. ABOUL-ELA

Texas Woman ’s University, Denton 76204

ABSTRACT

Isolated chloroplasts from spinach leaves were incubated with rat liver cells in monolayer
culture. The chloroplasts were phagocy tized by the cells and remained visibly intact for a per¬
iod of 3 hrs. The chloroplasts were incubated with the lysosomal extract from the liver cells for
15 min. There was no evidence of a release of chlorophyll or protein from the chloroplasts.

INTRODUCTION

There are numerous examples of chloroplasts existing in an endosymbiotic
relationship within the cytoplasm of certain animal cells (Taylor, 1968; Trench
and Smith, 1970; Trench, et al., 1972). Although these animal cells contain a
variety of lysosomal enzymes capable of degrading the major categories of biologi¬
cally important molecules, the chloroplasts do not appear to be digested by these
enzymes. Chloroplasts phagocy tized by fibroblasts in culture have retained their
ultrastructural integrity for 5 cell generations (Nass, 1969). When introduced into
the white of hens’ eggs, isolated chloroplasts have undergone repeated division
and maintained their metabolic activity as indicated by C02 fixation and Hill
activity (Giles, 1972).

These observations of the cell’s inability to dispose of internalized chloroplasts
have prompted this further examination of the effect of lysosomal enzymes on
chloroplasts.

MATERIALS AND METHODS

After it was confirmed in our laboratory that chloroplasts were phagocytized
but not degraded by rat liver cells in vitro, we investigated the effect of lysosomal
enzymes on intact chloroplasts. Isolated chloroplasts were incubated with lyso¬
somal enzyme extract and chloroplast degradation determined by microscopic
examination and by quantitation of free chlorophyll and soluble protein.

Phagocytosis of Chloroplasts by Liver Cells

Liver cells were isolated by a modification of the Dulbecco and Vogt method
(Dulbecco and Vogt, 1954), suspended in TC-199 (Difco) supplemented with 20%

Accepted for publication: April 14, 1976.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

86

THE TEXAS JOURNAL OF SCIENCE

calf serum, and seeded in Leighton tubes at a density of 3 - 4x 10s cells/cm2.
Chloroplasts were diluted with the same medium and added to the monolayer in
a ratio of approximately 5:1. After 1 hr of incubation in the dark the glass cover-
slip was removed from the Leighton tube, inverted onto a glass slide and the cells
examined by phase microscopy at 37 C (Reichert Zetopan, 1250x).

Chloroplasts were isolated from fresh spinach (Nass, 1969), and the resulting
chloroplast pellet was resuspended in a modified Honda medium (Honda, et al,
1966; Nass, 1969). Each preparation was examined by phase microscopy to de¬
termine the quality of the chloroplasts then stored at 4 C for no longer than 48
hrs prior to use.

Isolation of Lysosmal Enzymes

Three male Sprague Dawley rats (200-250 gm) were sacrificed by decapitation
after fasting for 22-24 hrs. The livers were excised, weighed and minced. Five vol¬
umes of 0.25 M sucrose chilled to 4 C were added per gram of original liver tissue
(5 ml/gm). The liver was then homogenized using 1 stroke of a motor driven (1000
RPM) Potter-Elvehjem homogenizer. The liver homogenate was centrifuged using
a differential centrifugation procedure as described by DeDuve, et al (1955). The
resulting lysosomal pellet was made into 0.2% Triton X-100, refrigerated 15 min
and assayed for 3-glucouronidase.

Enzyme Assay

3-Glucouronidase was selected as the marker enzyme (Stahl andTouster, 1971)
in the lysosomal extract and assayed as follows. To 0.25 ml 0.2 M sodium phenol-
phthalein glucouronide and 0.025 ml lysosomal extract previously made 0.2% in
Triton X-100, plus sufficient water to make the total volume of the mixture to
1 .0 ml. The mixture was incubated at 37 C for 15 min. The reaction was stopped
with 3 ml stopping reagent (consisting of 0.13 M glycine, 0.67 M NaCl and 0.083 M
Na2C03 per liter after adjusting the pH to 10.7 with 1 N NaOH) and absorbance
determined at 552 nm. The absorbance values indicated an average release of
0.013 [x moles of phenophthalein which is equivalent to 0.05 enzyme unit activity.

Chlorophyll and Protein Determination

To each of three 0.25 ml aliquots of chloroplast suspension the following were
added: 0.25 ml sodium acetate buffer, lysosomal enzyme fraction (0.025, 0.05,
and 0.1 ml), and sufficient water to make the entire incubation mixture total 1 ml.
The mixtures were incubated for 15 min at 37 C with constant gentle mixing.
Enzyme activity was stopped by adding 3 ml stopping reagent (described above),
and the incubation mixtures were then centrifuged at 2000 x g for 7 min to remove
chloroplasts and debris. The absorbance of an aliquot of the supernatant in acetone
was measured at 663 and 664 nm to determine the amounts of chlorophylls a and b,
and total chlorophyll in mg/g of tissue was calculated according to Arnon (1949)
and Koski (1950). The values obtained were then converted into % of the total
chlorophyll extracted from a sample of 0.25 ml chloroplast suspension made into
10 ml in acetone.

LYSOSOMAL ENZYMES

87

A modification of the Lowry method was used to determine the total protein
in each sample (Campbell and Sargent, 1967; Lowry, et al, 1951). The values ob¬
tained were converted into % of the amount of protein in 0.1 ml enzyme preparation.

RESULTS AND DISCUSSION

Microscopic examination of the rat liver monolayer after 1 hr of incubation
demonstrated that although the cells did phagocytize between 5 and 6 chloro-
plasts/cell, continuous observations for 3 hrs showed no evidence of chloroplast
degradation occurring within the cytoplasm of these cells.

The amount of chlorophyll present in the supernatant of chloroplast suspensions
varied with the preparation; therefore, chlorophyll values were expressed as a %
of the total chlorophyll extracted from the chloroplast preparation. The mean
values of free chlorophyll in the supernatants of the chloroplast control and the
chloroplast-enzyme mixtures ranged from 5% to 7% (Figure 1). As a result of
statistical analyses it could be inferred that the amount of chlorophyll leached
from the chloroplasts into the supernatant was not influenced by lysosomal enzyme
activity.

The amount of soluble protein released from the chloroplasts did not change
significantly (at 0.05 probability) after incubation with the enzyme extract. It is
seen in Figure 2 that the curve representing chloroplast-enzyme and the curve
representing enzyme alone are parallel. The difference between the 2 curves rep¬
resents the amount of soluble protein in the supernatant of 0.25 ml chloroplast
suspension. As the aliquots of enzyme protein increased from 0.025 to 0.1 ml,
there was a proportionate increase in the total protein of the chloroplast-enzyme
supernatant. The loss of protein from chloroplasts has been previously reported
by Kahn and Wettstein (1961) who found that chloroplasts isolated in an aqueous
medium such as Honda medium lose much of the stromal protein within 30 min
after isolation even though they appear to be structurally intact.

Earlier unpublished studies in our laboratory using neutral red as a lysosomal
marker showed that human neutrophils readily phagocytize chloroplasts, but that
the lysosomes do not coalesce with internalized chloroplasts. These observations
of chloroplast-lysosome interaction support the work reported by other investi¬
gations (Giles, 1972; Nass, 1969; Taylor, 1968; Trench and Smith, 1970; Trench,
etal, 1972).

From this study there is evidence that neither chlorophyll nor protein are re¬
leased from chloroplasts which have been incubated with lysosomal enzyme extract.
The lysosomal enzymes are unable to degrade the chloroplast membrane under the
conditions specified. Of course the possibility cannot be ruled out that the chloro¬
plasts released an inhibitor which deactivated the lysosomal extract. These results
are in agreement with the possible symbiotic origin of the chloroplast-bearing
eukaryotic cells.

88

THE TEXAS JOURNAL OF SCIENCE

0.0 0.025 0.05 0.10

ML LYSOSOMAL ENZYME EXTRACT

Figure 1 . Chlorophyll present in supernatant of chloroplast-enzyme incubation mixtures.

All samples contain 0.25 ml chloroplast suspension. Mean chlorophyll values
(expressed as % of total chlorophyll extracted) with their standard error of the
mean represent 2 experiments with 3 replicates each. Total chlorophyll in chloro¬
plast suspension was assayed as 650jUg/ml.

SUMMARY

Rat liver cells in monolayer culture phagocytized isolated spinach chloroplasts;
however, these internalized organelles were not visibly degraded after 3 hrs of
continuous observation. Chloroplasts were incubated with lysosomal enzyme ex¬
tract prepared by homogenization and differential centrifugation of rat liver, and
the supernatants of chloroplast-enzyme incubation mixtures were analyzed to
determine chloroplast degradation. The amount of free chlorophyll and soluble
protein present in the supernatant of chlorophyll-enzyme incubation mixtures
were not influenced by lysosomal enzyme activity.

SOLUBLE PROTEIN AS % OF AMOUNT OF 0.1 ML ENZYME

LYSOSOMAL ENZYMES

89

ML LYSOSOMAL ENZYME EXTRACT

Figure 2. Amount of soluble protein from 0.25 ml chloroplast suspension incubated with

lysosomal enzyme, o - o, and soluble protein in enzyme preparation alone,

expressed as % of amount in 0.1 ml. Each value with its standard error
of the mean represents 2 experiments with 3 replicates each.

ACKNOWLEDGEMENTS

The authors wish to express their thanks to Dr. Phillip Stahl for his technical
advice and critical discussions.

LITERATURE CITED

Arnon, D. I., 1949 -Plant Physiol., 24:1.

Campbell, P. M., and F. R. Sargent, 1967 —Techniques in Protein Synthesis. Academic Press,
New York, N.Y., 429 pp.

DeDuve, C.,B. C. Pressman, R. Granetto, R. Waltioux, and F. Applemans, 19 55-Biochem. /.,
60:604.

Dulbecco, R., and M. Vogt, 1954-/. Exp. Med., 99:167.

Giles, K. L., 191 2 -Nat. New Biol, 236:56.

90

THE TEXAS JOURNAL OF SCIENCE

Honda, S. J T. Hongladarom, and G. G. Laties, 1966-7. Exp. Bot., 17:460.

Kohn, A., and D. Wettstein, 1961-7. Ultrastruct. Res., 5:557 .

Koski, V., 1950 -Arch. Biochem. Biophys., 29:339.

Lowry, O. H., N. J . Rosebrough, A. L. Farr, and R. S. Randell, 195 1-7. Biol. Chem., 193:265.
Nass, M. M., 1969— Sci., 165:1128.

Stahl, P. D., and O. Touster, 197 1 -7. Biol. Chem., 246:5 398.

Taylor, D. L., 1968-7. Mar. Biol. Assoc. U. K., 48:1.

Trench, R. K„ and D. C. Smith, \910-Nat., 227:196.

- , M. E. Trench, and L. Muscatine, \912-Biol. Bull., 142:335.

A SIMPLE, INEXPENSIVE FREEZE-ETCH DEVICE FOR USE IN
ELECTRON MICROSCOPE LABORATORIES

by G.D. CAGLE

New Product Research (Microbiology)

Alcon Laboratories, Inc., Fort Worth 76134

and G. R. VELA

Department of Biological Sciences,

North Texas State University, Denton 76203

ABSTRACT

A simple and inexpensive freeze-etch device for use in procedural electron microscopy
courses and laboratories is described. Frozen cells are cleaved beneath the surface of liquid
nitrogen, and the cleaved cell surface etched by a method which allows differential, although
unregulated, heating of the holding block and the colder lid, to which ice sublimes.

INTRODUCTION

A replica of the frozen-etched surface of biological material may be prepared
by: i) rapidly freezing; ii) cleaving; iii) etching; and iv) replicating the specimen.
Adhering cellular material is removed and the replica observed directly with the
electron microscope. The technique of freeze-etching has been reviewed by Kohler
(1968) and Bullivant (1973) who cite the advantages of freeze -etching as: i) fixation,
dehydration, and embedding are not necessary; ii) cells differ from the living state
only in that they are frozen, and iii) fracture proceeds along a structurally weak
path, following the plasma membrane. Although previous attempts had been
made to freeze-etch biological material (Steere, 1957; Haggis, 1961), Moor, et al.
(1961), were the first to obtain consistent results with a cryoultramicrotome freeze-
etch device (Balzers’ apparatus). Subsequently, other workers (Koehler, 1966;
Steere, 1966) developed freeze-etch devices similar to the Balzers’ apparatus which
allow cleaving and etching of the specimen to be performed in an evacuated cham¬
ber. In contrast to the cryoultramicrotome freeze-etch devices of Moor, et al.
(1961), simple freeze-cleave devices have been developed (Bullivant and Ames, 1966;
Winklemann and Meyer, 1968). Types of each of the simple freeze-cleave devices
are commercially available, and are moderately expensive ($2,000 or more), al¬
though considerably less expensive than the Balzers’ freeze-etch device. A simple,

Accepted for publication: November 4 , 1976.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

92

THE TEXAS JOURNAL OF SCIENCE

inexpensive freeze-etch apparatus which lacks precise temperature controls can
be constructed for use in electron microscope laboratories equipped with a rotary-
stage vacuum evaporator.

MATERIALS AND METHODS

The principle components of the freeze-etch device, illustrated in Figure 1,
include an insulating cup (IC), holding block (HB), and lid (L), machined from
brass. The insulating cup (IC) is 3.2 cm high, 4.7 cm in diameter, with a 0.3 cm
cone centered in the bottom to support the holding block. The holding block (HB),
2.6 cm high and 3.8 cm in diameter, has a small opening (arrow) in the center of

Figure 1. Photograph of specimen-holding device. Cells are placed in the hole (arrows)
in the center of the holding block (HB), cleaved, covered with the lid (L), and
placed in the insulating cup (IC) containing liquid nitrogen.

the block surface 0.25 cm in diameter and 0.6 cm deep. This opening holds the
cells during freezing, cleavage, and replication. Damage and contamination of the
cleaved cell surface is reduced by the cover (L) which is flanged to fit closely to
the top of the holding block (HB). Other parts of the device, depicted on the
stage of the vacuum evaporator, are an insulating platform (IP), an arm-guide (A-G)
and a threaded shaft (TS) on which the arm, attached to the lid (L) by a hook (H), is
raised (Figure 2).

Bacteria were prepared for freeze-etching by removing them from the surface
of a solid medium and placing them in the opening in the holding block. In some
cases, cells were pretreated with 28% glycerol for 30 min before freezing. The
holding block was placed in the insulating cup and the cells rapidly frozen by
immersing the apparatus in liquid nitrogen (-196 C). The cells were cleaved beneath
the surface of the liquid nitrogen with a razor blade cooled to liquid nitrogen
temperature. The lid was submerged in liquid nitrogen, placed over the cleaved

INEXPENSIVE FREEZE-ETCH DEVICE

93

Figure 2. Photograph of the apparatus as it appears on the insulating platform (IP) of
the vacuum evaporator. The lid (L) covering the cleaved cell surface is attached
to the arm-guide (A-G) by a hook (H), and the lid is raised by rotation of the
threaded shaft (TS). The Pt-C and C electrodes are also apparent.

cell surface, and the entire apparatus quickly transferred to the insulating platform
inside a Mikros VE 10 vacuum evaporator (Mikros, Inc., Portland, OR). During
evacuation, frost forms on the top of the lid and on the sides of the insulating
cup, but not on the surface of the cells in the holding block. The cleaved surface
was etched by withdrawing a thin block of insulating material, separating the in¬
sulating platform and insulating cup, with a length of fine cord, and then adjusting
the lid to a position 0.1 -0.2 cm above the cleaved cell surface. In this way the
temperature of the block is increased gradually , and the cleaved cells surface etched
by the slightly raised, and colder, lid. The method is unsophisticated, the tempera¬
ture of the block and lid are not monitored, and etching times must be empirically
determined, with periods between 3-12 min routinely employed.

After the cells were etched, they were shadowed with Pt-C (open arrows, Figure 3)
at an angle of approximately 45°, and backed at 90° with C (solid arrow, Figure 3)
for support. The chamber was pressurized and the holding block allowed the

94

THE TEXAS JOURNAL OF SCIENCE

Figure 3. Photograph of the apparatus as it appears during etching and replication, bell
jar removed. After a vacuum of 10 5 Torr is achieved, the lid is raised 0.1-0.2 cm,
the cell surface etched, and the lid raised approximately 8.5 cm to permit un¬
obstructed deposition of Pt-C (open arrow) and the C (solid arrow) by evapor¬
ation.

equilibrate to ambient temperature. Upon immersion in distilled water, the replica
invariably broke apart. Replicas could be collected with less contamination by
placing a thin plastic mask over the holding block, exposing only the cells which
were frozen cleaved, and etched. The replica was treated for 1 hr in H2S04, rinsed
several times in distilled H20, and treated for another hour in approximately 5%
sodium hypochlorite (commercial Chlorox) to remove adhering cellular material.
The replica was rinsed again in distilled H20 and collected on a collodion-coated
300-mesh copper grid. All replicas were examined with an RCA-EMU-3G electron
microscope at 50 kV. Photographic negatives were taken at initial magnifications
of 7, 700-1 1,500.

An aquatic Bacillus subtilis and a Pseudomonas sp. were cultured on trypticase
soy agar for 18-24 hr at 30 C prior to freeze-etching.

INEXPENSIVE FREEZE-ETCH DEVICE

95

RESULTS AND DISCUSSION

Replicas of bacterial cells obtained with this technique (Figures 4 and 5) are
similar to those obtained with other freeze-etch procedures (Nanninga, 1969;
Cagle, et al, 1972). Figure 4 is a replica of B. subtilis cleaved obliquely along the
cell wall (CW) and plasma membrane (PM), and through the cytoplasm (Cy).

Figure 4. Frozen-etched preparation of B. subtilis. A small portion of the ordered arrange¬
ment of the cell wall (CW) subunits is apparent (solid arrows), with areas of
plasma membrane (PM) and cytoplasm (Cy) exposed. Strand-like extracellular
material (EM) which may be polymer or artifact is observed. In some cases,
deep etching also exposes a second layer of cells (open arrows). Marker = 0.5 ^m
(x 8 1,000).

96

THE TEXAS JOURNAL OF SCIENCE

Several flagella (F) adhere to the cell wall of the organism and numerous strands
of extracellular material (EM) are also apparent. A small portion of the ordered
structure which comprises the cell wall is evident on the exterior of one cell (solid
arrows), which has been previously reported as a surface component of numerous
Bacillus species (Holt and Leadbetter, 1969). The plasma membrane (PM) contains
particles, characteristic of this layer in frozen-etched preparations. The absence
of temperature control results in deep etching, and some replicas reveal a second
layer of cells (open arrows) (Figure 4).

A similar cleavage pattern is also observed with the aquatic pseudomonad, frac¬
tured to reveal a large portion of the cell wall (CW), and a small portion of the
granular plasma membrane (PM) and cytoplasm (Cy) (Figure 5). The structure of
the bacterial cell wall is partially observed (open arrows), although much of the
wall is masked by eutectic material (solid arrows). The presence of this material
indicates that the temperature of the holding block increased during etching, with
melting of some low melting-point, salt-containing matrices.

Figure 5 . Freeze-etching of Pseudomonas sp. Eutectic material (solid arrows) masks most
of the cell wall (CW) substructure (open arrows). Portions of the cell wall have
been removed exposing the granular plasma membrane (PM) and the deeply
etched cytoplasm (Cy). Marker = 0.5 /dm (x5 3,000).

The freeze-etch device and procedures employed (Figures 1-3) combine the
etching method introduced by Hall (1950) and Steere (1957) with the simple
freeze-cleave apparatus of Bullivant and Ames (1966). The replicas in Figures 4

INEXPENSIVE FREEZE-ETCH DEVICE

97

and 5 are representative of those which may be routinely obtained, although better
results may be attained with practice. Cells cleaved through the cytoplasm appear
to be adversely affected by the relatively long etching times, and the best replicas
obtained were those cleaved along the plasma membrane, or tangent to the cells
which are subsequently exposed by etching. Although replicas produced with this
device do not always possess the fine structure which may be achieved with sophis¬
ticated freeze-etch devices, as the Balzers device (nor is it designed to replace the
Moor device as a research tool), it does provide a means for introducing the students
to the methodology of freeze -etching, as well as to some techniques of cryo-biology.
The advantages of this freeze-etch device are its simplicity and low cost (less than
$45.00); however, experience with the apparatus has shown that this method is
highly empirical, and that successful replicas are not always initially obtained.

LITERATURE CITED

Bullivant, S., 197 3 -Freeze-etching and freeze-fracturing. Advanced Techniques in Biological
Electron Microscopy. Springer-Verlag, Berlin, pp. 67-112.

- , and A. Ames, 1966-A simple freeze-fracture replication method for electron

microscopy./. Cell Biol, 29:435.

Cagle, G. D., G. R. Vela, and R. M. Pfister, 1972 -Freeze-etching of Azotobacter vinelandii :
examination of wall, exine, and vesicles./. Bacteriol., 109:1 191.

Haggis, C. E., 1961 -Electron microscope replicas from the surface of a fracture through
frozen cells./. Biophys. Biochem. Cytol., 9:841.

Hall, C. E., 1950-A low temperature replica method for electron microscopy./. Appl. Phys.,
21:61.

Holt, S. C., and E. R. Leadbetter, 1969— Comparative ultrastructure of selected aerobic spore¬
forming bacteria: a freeze-etching study. Bacteriol. Rev., 33:346.

Koehler, J. K., 1966-Fine structure observations in frozen-etched bovine spermatozoa./ Ultra-
struc. Res., 16:359.

- , 1968 -The technique and application of freeze-etching in ultrastructure research.

Ad vane. Biol. Med. Phys., 12:1.

Moor, H., K. Muhlethaler, H. Waldner, and A. Frey-Wyssling, 1961 -A new freezing ultra¬
microtome./. Biophys. Biochem. Cytol., 10:1.

Nanninga, N., 1969 -Preservation of the ultrastructure of Bacillus subtilis by chemical fixation
as verified by freeze-etching./. Cell Biol., 42:733.

Steere, R. L., 195 7 -Electron microscopy of structural detail in frozen biological specimens.
/. Biophys. Biochem. Cytol., 3:45.

- , 1966-Development and operation of a simplified freeze-etching unit. /. Appl.

Phys., 37:3939.

Winklemann, H., and H. W. Meyer, 1968-A routine freeze-etching technique of high effectivity
by simple means. Exp. Path., 2:277 .

THE FLOWER GARDENS CORAL REEFS

by M. SAMUEL CANNON

Department of Human Anatomy, College of Medicine,

Texas A &M University , College Station 77843

and EDMOND S. ALEXANDER

Department of Medical Illustration , University of Texas Health
Science Center, Dallas 75235

ABSTRACT

The Flower Gardens Coral Reefs, located 110 nautical miles SSE of Galveston Island,
Texas are the northernmost living coral reefs in the Gulf of Mexico. Today, these reefs are
an area of intense exploration, partly because of their ecological isolation and preservation
of plant and animal life. Approximately 360 marine species, many previously unreported
in the Gulf, have been identified within them. Also, marine invertebrates available at the
reefs are animals ideally suited for comparative neurobiological studies. In addition, the
West Flower Garden reef possess the most fully-developed reefal biotic communities along
the Texas-Louisiana coast.

INTRODUCTION

The Flower Gardens Coral Reefs (East and West), located approximately
110 nautical miles SSE of Galveston, Texas on the outer edge of the conti¬
nental shelf were virtually unexplored, until recently. Today, partly because
of their ecological isolation and preservation of marine plant and animal life,
the reefs are the focus of intense exploration, much of it by scientists from the
Marine Biomedical Institute (MBI) of the University of Texas Medical Branch
at Galveston, Texas A&M University (TAMU), the Universities of Houston
and Southwestern Louisiana and the National Museum of Natural History.

WORK UNDERTAKEN

The “Miss Freeport5’ is a 135 -foot research vessel (under operational control
of MBI) which can support up to 21 diver-scientists, who usually work in depths
from 70 to 85 ft. Occasionally they dived to 190 ft along the edge of the reef.
Near the Great Escarpment, the 2-man submersible, “Nekton Gamma,” (General
Oceanographies, Inc., Newport Beach, California) carried scientists to depths
exceeding 450 ft. The excursion defined the reefs diverse biological and geo-
Accepted for publication: December 13, 1976
The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

100

THE TEXAS JOURNAL OF SCIENCE

logical forms, and provided evidence refuting the earlier conclusion that the
Flower Gardens coral reef was dead. It’s alive, thriving and much deeper (more
than 450 ft) and older than imagined. Moreover, it is the northernmost living
coral reef in the Gulf of Mexico and to date some 360 marine species, many
previously unreported in the Gulf, have been identified within it. Under the
sponsorship of MBI, Drs. Thomas Bright and Linda Peguegnat of TAMU and
their collaborators published a comprehensive survey of living forms on the
reef; the book is entitled, “Biota of the West Flower Garden Bank” (Gulf
Publ. Co., Houston, Texas). In addition, geologists and engineers edited a de¬
tailed bottom map. Scaled at one in/1 00 ft, the map’s resolution exceeds those
of the lunar surface compiled by NASA prior to the Apollo Missions. To quote
Robert Alderdice, operation’s officer in charge of the Flower Gardens Cruises,
“Incredibly, man has more detailed information of another planetary body
than its own coastal waters, which have been navigated for centuries.”

The continental shelves of the United States possess a tremendous potential
as new geographic support areas near the coastal zone and an ultimate goal is
to make the shelves inhabitable. But, how to successfully integrate them into
the nation’s economic geography is still another problem. The solution re¬
quires, among other things, detailed maps and complete geological knowledge
of the shelves. The Flower Gardens reefs, on the edge of the Texas Continental
shelf probably formed originally atop sedimentary rock forced upward by a
salt dome. Whether the coral is solid and continuous or only a cap over a sub¬
strate of rock is unknown, but core sampling may provide the answer.

Moreover, the Flower Gardens are a unique biological and chemical com¬
munity, and in many respects it is an ecologically closed environment. The
water is remarkably clear (to around 200 ft), always between 21 C to 22 C
(70 F to 72 F) and much of the indigenous flora and fauna are found nowhere
else in the Gulf. Furthermore, the West Flower Garden coral reef has the most
fully-developed series of reefal biotic communities along the Texas-Louisiana
coast and studies here may furnish answers applicable to other biotic structured
communities. The crest of the reef itself is approximately 75 ft beneath the
water’s surface. The depth across most of the reef is 70 to 85 ft, while sur¬
rounding waters are 350 to 460 ft deep. Approximately 40% of the lower
80 acres of reef atop the West Flower Garden is covered by viable building
corals, primarily species of Diploria, Montastrea and Pontes ; the massive coral
heads approach 10 ft in diameter. These spectacular coral growths, plus com¬
munities of sessile and mobile organisms attract divers and undersea photo¬
graphers during spring and summer. Myriad species of fish and hundreds of
invertebrate varieties are encountered. Commercial snappers ( Lutjanus sp .)
and groupers (Family Serranidae) occupy the deeper flanks of the Flower
Gardens reefs which are frequented by fishing boats. Utilizing submersibles,
underwater television and other appropriate techniques, investigators hope
to observe migration, schooling reproductive behavior, habitat preference and
feeding characteristics of the snappers and other commercial and sport fishes.

FLOWER GARDENS

101

Other organisms found at the Flower Gardens include foraminifers, sponges
(Porifera), marine worms (Polychaeta), brachipods, numerous varieties of
gastropods, mollusks, crustaceans and echinoderms. There are over 100 species
of tropical fish. Particularly fascinating on the living reef above 150 ft are the
diverse polychaetous worms. Some worms ( Eunice sp.) are exceptionally large
with one specimen of E. aphroditois nearly 2 ft long. Echinoderms occupy
all depths of the West Flower Garden bank, while a sizeable crustacean pop¬
ulation occurs between 150 and 280 ft. Among the bottom fish, Chromis
enchrysurus is most numerous. The grouping of exclusively deep reef fishes
is intriguing and contains the most important commercial food fish in the Gulf
of Mexico. The Red snapper ( Lutjanus campechanus) and several relatives, in¬
cluding the Vermillion snapper (Rhomboplites aurorubens ). Catches of Red snapper
are better during fall and winter, but in June, 1972, considerably more Red
than Vermillion snapper were observed over the drowned reef and additional
investigation of the feeding behavior of these 2 species appears warranted.
Also significant were the occasional sightings of the rather rare serranid, the
Spanish flagfish, ( Gonioplectrus hispanus) and the butterfly fish, ( Chaetodon
aya), the latter previously reported absent. Further descriptive and quanti¬
tative data regarding the biota of deeper portions of the West Flower Garden
bank and other reefs should clarify the complex inter and intracommunity
relationships.

Virtually all exploration of the Flower Gardens reefs involved putting marine
scientists into the sea, they “went after” their subjects of study. This is quite
remarkable as “traditional oceanography” has been a science of remote control
using dredges, pumps and elaborate instrumentation usually manipulated from
the deck of a ship. The direct approach of “diver oceanography” is more de¬
pendable; it produces extremely accurate scientific data and is less harsh to
the ecosystem. O

Environmental studies acquire high-priority from marine scientists. The
Gulf of Mexico receives nearly 2/3 of the sediment and industrial waste from
the continental United States and over 3,000 offshore oil wells and miles of
subsea pipelines dot the coastal waters of Texas, Louisiana and Mexico. The
concentration of chlorinated hydrocarbons and heavy metals in organisms
from the Flower Gardens is somewhat higher than from other Gulf areas. By
performing environmental quality baseline studies, scientists hope to create
a continuous program of biological sampling and monitoring.

In addition, medicine can learn much from the sea. And, comparative neuro¬
biology offers infinite possibilities for medicine. The elaborate neuronal organ
system of mammals, including man, is fashioned upon structures and principles
evolved over millions of years, primarily in the sea. The chemical transmitters,
hormones and enzymes in man also occur in ancient, less complicated marine
invertebrates. These chemical substances vary between species, but their fund¬
amental molecular structure is identical in almost every respect. Similarly,
specialized tissues such as neurons are nearly identical in marine organisms and
man. Medicine demands an understanding of how human beings adapt to their
environment, and the brain, with its neurons and trillions of circuits is the chief

102

THE TEXAS JOURNAL OF SCIENCE

regulator of adaptability. Study of simpler nerve networks and untangling the
complexities of neural integration and transmission in organisms such as the
marine mollusk, Aplysia brasiliana, a shell -less sea snail whose brain is con¬
veniently located in its belly and contains approximately 1,500 neurons, is a
first step in penetrating the circuitry of man’s brain. Thus, tapping the sea’s
resources will harvest new knowledge prerequisite to solving many of today’s
environmental and medical problems. Indeed, the sea is man’s final frontier
on earth.

A REEVALUATION OF TWO NEW SPECIES OF FOSSIL BATS
FROM INNER SPACE CAVERNS

by SARA L. DORSEY

Shuler Museum of Paleontology ,

Southern Methodist University, Dallas 75275

ABSTRACT

Choate and Hall (1967) proposed 2 new species of My otis on the basis of a small collection
of Pleistocene fossils recovered from Inner Space Caverns, Georgetown, Texas. Since that
time a large number of topotypes have been recovered and as a result a reevaluation of the 2
extinct species was undertaken. One,M magnamolaris is considered con specific with the living
M. velifer and the other, M. rectidentis is considered valid.

INTRODUCTION

Pleistocene bat remains from Inner Space Caverns (formerly Laubach Cave) in
Georgetown, Texas, were first reported by Slaughter (1966) and referred to as
“My otis sp.” This small collection was later studied and reported on by Choate
and Hall (1967) and 2 new species were proposed, Myotis magnamolaris and
My otis rectidentis.

The cave assemblage, is believed to have accumulated during the last major
interstadial, probably between 25,000 and 40,000 years before the present. A
radiocarbon date on bones of the extinct peccarry \Platygonus, produced an age
of 13,900 ± 400. Considering the stage of bone dating at that time, this is con¬
sidered to be a minimum date and Slaughter believes the bones to be in excess of
20,000 B.P.

Choate and Hall’s study (1967) was based on 23 partial and complete mandibles
and 2 left maxilli. Since that time the cavern has been revisited and an additional
79 mandibles and 6 maxilli were recovered. These 2 collections are combined for
a reevaluation of the 2 species. Forty -seven specimens of modern Myotis velifer
incautus were also included in the study for comparative purposes. Measurements
included in the table and discussions are in mm and were taken by means of an
ocular micrometer in a binocular microscope.

Myotis rectidentis Choate and Hall (1967)

Referred Specimens

Holotype (SMU-SMP 61780); 10 mandibles including the paratype and new
topotypes. All specimens are from Inner Space Caverns, Georgetown, Travis
County, Texas.

Accepted for publication: November 3, 1975.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

104

THE TEXAS JOURNAL OF SCIENCE

Comparison With Other Species

Considerations for comparative species were based on the modern geographic
ranges of the various species.

Af. re ctidentis can be distinguished from Af. yumanensis by the smaller size of
the latter.

Af. lucifugus occultus is about the size of M. rectidentis, but the premolars are
more crowded.

Finally, Af. keenii apache has a thinner mandible with a straight shape in con¬
trast to the curving shape of the mandible of Af. rectidentis .

Discussion

Morphologically, Af. rectidentis seems most like modern Af. velifer but scatter
diagrams showing the length-width ratios of ml , m2, and m3*.place the holotype
of Af. rectidentis and 9 other specimens in a distinct cloud from both Af. magna-
molaris and modern Af velifer. As the results of the molar diagrams are identical,
only the one for m2 is presented (Figure 1). Only 2 lower canines are present in
the collection so they were not plotted. With canine lengths of 1.25 and 1.5,
they were smaller than those of Af. velifer. No significant differences other than
size were found between Af. rectidentis and Af. velifer. Nevertheless, the size is so

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Af. velifer, squares represent fossils of Af. rectidentis, and circles represent fossils
of M. velifer from Inner Space Caverns, the holotypes of Af. rectidentis and Af.
magnamolaris are designated H, paratypes P, and topotypes are the unlettered
symbols.

FOSSIL BATS

105

consistent, it is felt that the species is valid, and it is agreed that M. rectiden tis is
a bat “similar to M. velifer but smaller.” The M. velifer sample included almost
an equal number of males and females, eliminating the possibility that the 2 clouds
on the scatter diagram represent sexual dimorphism.

Measurements

For means and extremes of lower teeth see Table 1 . There are 4 specimens
containing upper molars that fit in with the smaller M. rectidentis species. One
such specimen was described by Choate and Hall (1967) while the other 3 were
more recent finds at the cave. The new maxilli have crown lengths of Ml ranging
from 1 .37 - 1 .62, of M2 ranging from 1 .37- 1 .56, and of M3 ranging from 0.75 -
0.81 . The greatest width of the molars varies from 1 .31 - 1 .93 in Ml , from 1 .37 -
1.56 in M2, and the lone measurement of M3 is 1.37. These measurements are
smaller than similar measurements of M. velifer.

TABLE 1

Lower molar
tooth row

l

Length

rn 1

Width

m2

Length Width

m3

Length Width

Modern velifer

4.31-4.68

1.43-1.62

1.00-1.18

1.43-1.56

0.93-1.18

1.31-1.56

0.87-0.93

Means

4.47

1.5 3

1.08

1.51

1.05

1.41

0.90

SMU Fossils

4.25-5.06

1.50-1.7 5

1.06-1.37

1.43-1.68

1.00-125

1.06-1.62

0.68-1.06

Means

4.64

1.60

1.16

1.5 7

1.07

1.46

0.95

M. rectidentis

3.12-3.68

1.12-1.31

0.75-0.87

1.00-125

0.62-0.93

0.87-1.12

0.50-0.75

Means

3.42

1.20

0.83

1.15

0.80

1.04

0.65

My otis velifer (J. A. Allen)

My otis magnamo laris Choate and Hall (1967)

Referred Material

Sixty-eight mandibles, 4 maxilli, and 1 complete palate from Inner Space
Caverns, Georgetown, Travis County, Texas. This includes the holotype (SMU-SMP
61772) and paratypes of M. magnamolaris.

Discussion

Choate and Hall’s (1967) diagnosis of M. magnamolaris placed greatest signifi¬
cance on the large size of the fossil form. Length -width ratios were plotted on
scatter diagrams for ml, m2, and m3, for the holotype, paratypes, for the greatly
expanded collection of topotypes, and finally for the specimens of modern M.
velifer (Figure 1). As the results were exactly the same for each molar, only the
diagram of m2 is presented. The fossils from Inner Space Caverns and the modern
sample form a continuous cline, although the holotype of M. magnamolaris is
the largest of the collection.

106

THE TEXAS JOURNAL OF SCIENCE

Choate and Hall’s (1967) diagnosis also characterizes^. magnamo laris as having
a lower canine that is massive and long. Only one canine (the holotype’s) however,
was preserved in the original collection (see Figure 2). We now have several others
and the size of the canine seems to be a matter of allometric growth, considering
the greater size of the holotype. No other morphological differences were found
in the fossils that cannot be duplicated in modern M. velifer.

B

Figure 2. View of the tooth row of the holotypes of (A )M. magnamolaris and (B )M. rectidentis.

It seems possible that the Inner Space bats represent a rather basic group for
the species and may actually be ancestral to one or more modern subspecies.
Choate and Hall (1967) suggested that the fossils are “possibly the direct ancestor
of M. velifer .” It seems equally possible, however, that the fossils represent an
extinct geographic subspecies that on an average was a bit larger than the modern
myotids. It is known that many extant species were larger in size during the late
Pleistocene. In any case , only maximum sized specimens of M. magnamolaris could
be distinguished from M. velifer.

If, of course, additional material (i.e., skulls) should provide viable characters
allowing separation of the form from the modern form, it would be of no more
than subspecific rank. Whether it proved to be an extinct geographic race, a dead¬
end temporal race, or a race ancestral to one or more modern subspecies, it would
be proper to be designated as M. velifer magnamolaris Choate and Hall (1967).

FOSSIL BATS

107

Comparison With Other Species

M. volans has anterior premolars that are crowded out of the anterior-posterior
tooth row as in the fossil M. velifer, but the protoconid basin of P4 is constricted
both anterior and posterior to form a barbell -shaped basin. Also the hypocone is
not as prominent a cusp as in the fossil forms.

M. thysanodes differs from the fossil M. velifer in several ways. First, there is
no crowding of the anterior premolars, also there is a weaker development of the
crest that extends from the protocone-hypocone to the lingual face of the meta¬
cone. Finally, the protocone extends farther lingually than the hypocone , while
in the fossil forms the protocone and hypocone are subequal, giving it a more
quadrangular appearance.

Measurements

For means and extremes of lower teeth, see Table 1 . In the way of maxillary
measurements there are 4 specimens including a complete palate in addition to
the previously described maxillary from 1967. The crown -length range of Ml is
from 1.88-1 .75, in M2 from 1.62-1.68, and in M3 from 0.93-1 .12. The greatest
breadth of Ml is from 1.62-1.8, of M2 from 1.75-1. 93, and of M3 from 1.37-1.81.
These fall well within the range of modern velifer.

SUMMARY

The collection of bat fossils from Inner Space Caverns (formerly Laubach Cave)
has been greatly expanded. A reevaluation of the 2 extinct species proposed by
Choate and Hall (1967) was undertaken as a result. The following conclusions
have been reached.

M. rectidentis Choate and Hall (1967) cannot be distinguished fromM velifer
on any morphological basis other than by its smaller size. Scatter diagrams of the
length -width ratios of the molars demonstrate that the size range does not over¬
lap with either modern Texas myotids or those from Inner Space previously de¬
scribed as M. magnamolaris. I therefore feel that this species is valid.

The holotype and paratypes of M. magnamolaris form a continuous cline,
with respect to size ratios of the teeth with the modern species of M. velifer and
the newly collected topotypes from Inner Space Caverns. Myo tis magnamolaris is
therefore considered conspecific withM. velifer. The fossil population may average
slightly larger than the modern population, but 95% of the specimens could not
be distinguished from the modern form. The fossils could represent an extinct
geographic race, or a temporal race ancestral to one or more modern subspecies.

ACKNOWLEDGEMENTS

Acknowledgements are due Bob H. Slaughter, Shuler Museum of Pale ontology,
Southern Methodist University, for suggestions and reading of the manuscript.
Thanks also are due Dr. Walter Dalquest, Midwestern University, for the loan of

108

THE TEXAS JOURNAL OF SCIENCE

the recent Myotis velifer collection and Dr. Henry Setzer, U.S. Natural History
Museum, for allowing my study of the collections under his care.

LITERATURE CITED

Choate, J. R., and E. R.Hall, 1967-Twonew species of bats, Genus Myotis, from a Pleistocene
deposit in Texas. Am, Mid. Nat., 78(2) :5 31.

Slaughter, B. H., 1966-Platygonus compressus and associated fauna from the Laubach Cave
of Texas. Am. Mid. Nat., 75 (2) :475 .

SEASONAL DISTRIBUTION OF MITES ASSOCIATED WITH
POPILIUS DISJUNCTUS (ILLIGER) (COLEOPTERA; PASSALIDAE)
IN HARDIN COUNTY, TEXAS

by GEORGE E. GIBSON, JR.1

Department of Biology, Lamar University, Beaumont 77710

ABSTRACT

Seasonal distribution of mites associated with Popilius disjunctus was studied by collecting
2 sets of beetles, 1 in the winter and 1 in the summer of 1974, from a beech-magnolia forest
in Hardin County, Texas. Some correlation between climatic conditions and mite population/
beetle was noted. The region being temperate-subtropic, the seasonal variation in temperature
was minimal and thus had an effect on mite population density and kinds present.

INTRODUCTION

Popilius disjunctus (Illiger), as described in many works, is a semisocial beetle
found in rotting oak, hickory, gum, beech, and magnolia logs. It has been utilized
by several researchers for both laboratory and field experiments and observations
(Borror and DeLong, 1954; Pearse, et al, 1936; Gray, 1946; Hunter and Mollin,
1964; Harrel, 1967). Work on the seasonal distribution of the commensal and
parasitic mites associated with P. disjunctus has been limited to relatively few re¬
searchers. Hunter and Mollin (1964) and Harrel and Mathis (1965) have dealt
specifically with seasonal influences on mite populations on P. disjunctus in North
Carolina and Oklahoma. No work is known for the southeastern portion of Texas.

To further investigate this symbiotic relationship under seasonal influence and
originate a study in southeast Texas, 2 collections of beetles were made in a post
climax beech -magnolia forest in Hardin County, Texas, during the months of
August and December, 1974.

METHODS

On August 30, 1974, and December 22, 1974, 25 beetles were collected in a
beech-magnolia forest in Hardin County, Texas, just off Highway 96. The mites
were removed from 10 randomly chosen beetles and placed in 80% isopropyl alcohol.

When all mites on the exterior had been found and identified, the elytra and
hind wings were removed to search for Zercon passalorum and Heterochelytus

Present address: La Marque High School, 300 Vauthier Road, La Marque 77568.

Accepted for publication: November 11, 1976.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-A, March, 1977.

no

THE TEXAS JOURNAL OF SCIENCE

fusiformis. These 2 species of mites are believed to be the only 2 parasitic species
found on the beetle (Pear se, etal., 1936). In addition, numerous Hypopi (nymphs)
(Family: Tyroglyphidae) were encountered in clusters under the elytra.

Identification was made on all mites, except Cosmolaelaps passali and Oppelia sp.,
by comparison with specimens identified by Dr. Preston Hunter. C. passali was
identified utilizing a work by Hunter and Mollin (1964). Oppelia sp. was identified
through personal communication with Dr. G.W. Wharton at Ohio State University
in March, 1975.

When necessary, slides were made of each of the species found, particularly of
the 3 species of the genus Uroobovella. The Uroobovella sp. were not separately
identified for correlative purposes by either Pearse, et al. (1936), or Harrel and
Mathis (1965).

RESULTS

Results are presented in Table 1. The average number of each species, where
applicable, is compared to corresponding month averages obtained by Pearse, et al
(1936), indicated by an asterisk (*), and to the March averages of Harrel and Mathis
(1965), identified by a double asterisk (**).

A total of 20 beetles were examined. The total number of mites found in August,
1974, was 1858. In December, 1974, 3173 mites were found. Average mites/beetle
was 185.8 and 3 17.3 respectively. A total of 14 mite genera were found, although
Oppelia sp., C passali , and Caebnopsis latus were found only in the August collection.

DISCUSSION AND CONCLUSIONS

As indicated by Pearse, et al (1936), Hunter and Mollin (1964), and Harrel
and Mathis (1965), the species averages are highly variable. However, win ter aver¬
ages are somewhat higher than summer averages. This is especially true for Zercon
passalorum and Heterochelytus fusiformis . Variations in results when compared
to previous individual patent leather beetle works may be explained by many
factors. Environmental conditions of Southeast Texas are unique. The 2 other
studies used here for comparative purposes were done in North Carolina and
Oklahoma. Those areas, while considered temperate in climatic conditions, are
subjected to much more severe winters than the temperate-subtropic climatic
conditions of Southeast Texas.

During the study the difference in the August and December temperature was
not extreme. This held true of the whole winter, but overall climatic conditions
for the area were not atypical to normal seasonal change. It is concluded that the
increase in the number of mites found in the second collection (December) was
due to the somewhat cooler climatic conditions and availability of habitat provided
to mites when the beetles become more colonial during the winter months.

SEASONAL DISTRIBUTION OF MITES

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The forest composition did not seem to have any correlative significance to the
study, but further study is indicated to determine if tree specificity is relative to
mite density and genera present.

Further research is also necessary to determine external body surface and subely-
tral subwing mite populations during a noticeable extreme of seasonal temperature
in Southeast Texas.

LITERATURE CITED

Borrow, D. J., and D. M. DeLong, 195 4 -An Introduction to the Study of Insects. Holt, Rinehart
and Winston, p. 382.

112

THE TEXAS JOURNAL OF SCIENCE

Gray, I. E., 1946— Observation on the life history of the horned Passalus. Amer. Mid . Nat.,
35:729.

Harrel, Richard C., 1967 -Popilius disjunctus (Illiger) (Coleoptera: Passalidae) as a laboratory
animal. Turtox News, 45 (11) :270.

- , and Billy J. Mathis, 1965 -Mites associated with Popilius disjunctus (Illiger)

Coleoptera: Passalidae) in McCurtain County, Oklahoma. Proc. Okla. Acad. Sci., 45 :66.

Hunter, E. P., and Karl Mollin, 1964-Mites associated with Passalus beetles. I. Life stages and
seasonal abundance of Cosmolaelaps passali: n. sp. (A carina: Laelaplidae) . Acarologia,
6(2):247.

Pearse, A. S., M.T. Patterson, J. S. Rankin, and G.W. Wharton, 1936 -The ecology of Passalus
cornu tus. Fabricius. Ecol. Monog., 6(4):456.

CROWN POSITIONS WITHIN UNTHINNED LOBLOLLY PINE
PLANTATION CANOPIES

by J. DAVID LENHART1, CHI-YUN HO2’3, and DWIGHT R. HICKS2’4

School of Forestry, Stephen F. Austin State University

Nacogdoches 75961

ABSTRACT

Crown class percentages are not affected by age or trees/acre. The percentage of dominant
and suppressed trees is affected by land productivity.

INTRODUCTION

Thinning practices employed to grow trees for selected products are usually
defined by designating proportions of trees to be removed from each of the 4
main recognized crown classes. There is little available published information,
however, regarding the distribution of tree crowns within the canopy prior to
thinning.

Information collected from 219 unthinned old- field loblolly pine (Pinus taeda L.)
plantations in the Interior West Gulf Coastal Plain during a mensurational study
provided an opportunity to describe the effects of age, site index, and trees/acre
on crown classes (Lenhart, 1972).

Plantation Measurements

Plantations selected for sampling were unthinned, at least 9 yrs old, and un¬
damaged by fire, insects, or disease. Within each plantation mensurational infor¬
mation, including age, site index (base age = 25), and trees/acre, were collected
from a sample plot. In addition, the trees (10 to 16) on a sub -sample within the
plot were each classified into one of 4 crown positions within the canopy— domi¬
nant, co-dominant, intermediate, or suppressed. These crown positions were
converted to percentages.

Crown Class Percentages

For all sample plots, the average % of trees in each crown class was as follows:

1 Associate Professor and Assistant to the Dean.

2 Former Graduate Research Assistants.

Presently employed by Ministry of Communications in Taiwan.
Presently employed by Drummond Coal Company.

Accepted for publication: October 14, 1975.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

114

THE TEXAS JOURNAL OF SCIENCE

Crown Class Percentage

Dominant 31

Co-dominant 28

Intermediate 31

Suppressed 10

The paucity of suppressed trees could suggest that many trees of this class have
already died and disappeared. More probably, however, the spacing of these
plantations— mostly nominally 6X8 feet— has been sufficient to support stands
in which few trees have not yet been crowded into the suppressed condition.

Crown Class Percentages by Age Classes

Since the sampled plantations ranged in age from 9 to 30 yrs, we were able to
describe crown positions in relation to 5-yr age classes (Figure 1).

AGE

Figure 1. Percentage of trees in the 4 crown classes by age classes. Number of sample
plots for each age class is shown in parentheses.

LOBLOLLY PINE PLANTATION CANOPIES

115

In age classes 10, 15, and 20, which included 95% of the sample plots, the %
of crowns in each class is fairly constant. In these age classes, less than 6 %-points
separate the upper 3 crown classes. Across all age classes about 10% of the trees
had suppressed crowns. There was no indication of much shifting of trees between
crown classes as they grow older.

Crown Class Percentages by Trees-Per-Acre Classes

The relationships between relative tree frequency in the 4 crown positions
and trees/acre are shown in Figure 2. The sampled plantations ranged from 200
to 1700 trees/acre with 1 sample each in the 1400 and 1700 classes. About 92%
of the sample plots were in the 400- to 1000-trees/acre classes.

200 300 400 500 600 700 800 900 1000 1100 1200

TREES PER ACRE

Figure 2. Percentage of trees in the 4 crown classes by trees/acre classes. Numbers in paren¬
theses represent the sample size for each trees-per-acre class.

Trends in Figure 2 resemble those in Figure 1; each of the upper 3 crown classes
includes about 30% of the trees, while the suppressed class represents about 10%.
Between 600 and 1100 trees/acre about 4 to 7 %-points represents the spread
between the dominant, co-dominant, and intermediate crown classes. No trends
toward change in crown class with density are apparent.

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Crown Class Percentages by Site Index Classes

Figure 3 depicts crown positions in relation to 5 -ft site index classes. About
91% of the sampled plantations had site index values between 45 and 65 ft, inclusive.

SITE INDEX

Figure 3. Percentage of trees in the 4 crown classes by site index classes. Numbers in
parentheses represent the sample size for each site index class.

The proportions of trees in the top and bottom crown positions, definitely
tend to vary with site index. As site index increases, the dominant % increases and
the suppressed % decreases. Also, intermediate % appears to decrease slowly with
increasing site index, while co-dominant % shows no definite trend.

On the poorer sites— 35, 40, and 45 ft, about 40% of the trees are either domi¬
nant or suppressed. On the better sites-65 , 70, and 75 ft, about 40% of the trees

LOBLOLLY PINE PLANTATION CANOPIES

117

are dominant, and 1 in 10 or less is suppressed. Across all site index classes, except
75 ft, the majority of the tree crowns are in either the co-dominant or intermediate
classes.

The data suggest that in un thinned plantations within the ranges observed,
the forester cannot expect any change in canopy composition as age or trees/acre
increase or decrease. It may require some type of thinning regime to increase, for
example, the % of dominant trees in a plantation. On the other hand, the % of
dominant trees is larger and the % of suppressed trees is smaller on more productive
than on less productive land. On the better sites about 40% of the trees are dominant.

ACKNOWLEDGEMENTS

Research for this project was supported in part by funds made available under
the Mclntire-Stennis Act.

This paper is based in part on a masters thesis by Mr. Ho.

LITERATURE CITED

Lenhart, J. D., 1972-Cubic-foot yields for unthinned old-field loblolly pine plantations in

the Interior West Gulf Coastal Plain. Texas Forestry Paper No. 14, 46 pp.

PRODUCTION OF AMINO ACIDS BY SOIL MICROORGANISMS

by F. RUIZ-BERRAQUERO and A. RAMOS-CORMENZANA

Department of Microbiology

University of Granada, Spain

INTRODUCTION

At the present time there is a revival in the use of microorganisms as agents
for agricultural soil fertilization (Voznyakouskaya, 1963; Azcon, et al., 1973).
The basis for this practice resides in the knowledge that many soil microorganisms
excrete amino acids, auxins, vitamins, etc., and that these substances enhance plant
growth, either singly or in combinations showing synergistic effects (Vagnerova
and Vancura, 1962; Brown, 1974). It is assumed that some microorganisms enhance
plant growth better than others and that these can be identified even in the soil
milieu.

This is a report on the production of specific amino acids by soil bacteria. The
research objective was to determine the numbers of microorganisms in the soil
which secrete amino acids and to assess the relative quantities produced. It is
assumed that the data presented will serve to identify those organisms that can
be used most effectively in commercial agricultural applications and perhaps even
in the commercial production of amino acids.

MATERIALS AND METHODS
Microorganisms

The production of amino acids was studied in the laboratory using 17 1 differ¬
ent organisms isolated from soils from the provinces of Barcelona and Alicante
in Spain. Assays of amino acids produced were performed using Leuconostoc
mesenteroides ATCC 8042 auxotrophic for arginine, aspartic acid, glutamic acid,
leucine and lysine while the production of alanine was monitored with Pediococcus
cerevisiae ATCC 8081 .

Media

The soil bacteria studied were isolated and maintained on medium consisting
of glucose , 20 g; NH4 Cl, 7 g; MgS04 • 1W2 0, 0.5 g; KH2 P04 , 2 g; agar, 1 1 g; and

Accepted for publication: July 1, 1975.

The Texas J ournal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

120

THE TEXAS JOURNAL OF SCIENCE

enough water to make 1 1. the pH was adjusted to 7 before sterilization. Stock
cultures of the auxotrophic organisms were maintained on Lactobacillus Broth
AOAC (Difco) supplemented with the required amino acid. Production of amino
acids was measured on specific amino acid assay media (Difco).

Production of Amino Acids

The assay for amino acids was based on the method used by Udaka (1960) for
the isolation of glutamic acid-producing bacteria. This method was originally in¬
troduced by Steele, et al, (1949) and more recently employed by Guinea (1970).

For this study, the auxotrophic organisms were spread on specific amino acid-
deficient media and then soil isolates were streaked onto these seeded plates.
Replicates were used to insure isolated and well-separated colonies. The plates
were incubated at room temperature for 5 days and then evaluated as shown
below.

Numerical value 0:

No growth of the auxotrophic organisms was observed macro- or microscop¬
ically.

Numerical value OM:

No growth of the auxotrophic organisms was observed macroscopically and
only scant growth was seen microscopically. Growth was limited to the
periphery of the colony of the amino acid-producing organism.

Numerical value 1 :

Growth visible macroscopically; limited (1 to 2 mm) to the periphery of
the colony of the amino acid-producing organism.

Numerical value 2 :

A definite halo of growth surrounding the colonies of the amino acid-pro-
ducing organisms. Growth extended some 3 to 4 mm from the colony and
the auxotroph colonies were well developed. (Figures 1 and 2)

Numerical value 3:

Abundant, confluent growth surrounding the colonies and extending more
than 4 mm from the colony.

While these criteria do not yield exact measurements, the growth patterns ob¬
served (Figures 1 and 2) are easy to assess for assignment of numerical values. Since
several parameters are used, i.e., number of auxotroph colonies, density of growth,
and extent of zone which supports growth, it is assumed that the numerical value
is a measure superior to any single criterion.

Evaluation of Amino Acid Production

Each of the 171 soil isolates as assayed for production of alanine, arginine,
aspartic acid, glutamic acid, leucine, and lysine.

The “index” of production of amino acids (P) by any given microorganisms
can be determined by summing the numerical values; 0, 1,2,3 and dividing this
sum by the number (n) of amino acids measured.

PRODUCTION OF AMINO ACIDS

121

Figure 1. Growth of auxotrophic organism associated with amino acids excreted by soil
bacteria.

In like manner, the “index” of production of amino acids by a group of micro¬
organisms (e.g., Gram positive heterotrophs) can be obtained by dividing the sum
of indices for each organism in the group by the number of microorganisms.

Statistical Treatment

In order to evaluate the significance of the differences (percentages) of organ¬
isms capable of producing a given amino acid, the following formula was employed.

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THE TEXAS JOURNAL OF SCIENCE

ki

and: k

then : d

where: Pi

p2

n

Pi; k2
ni n2

Pi +?2 + a * * P;

ni + n2 + • • • n.

/ k(l-

k) (

T>

the index of production of amino acids by a given organism
the index of production of amino acids by a second organism
number of organisms tested

Figure 2. Microcolonies of auxotrophic organism surrounding colonies of amino acid-
producing bacteria.

A *

PRODUCTION OF AMINO ACIDS

123

The calculated value of d was assumed to be derived from a population of values
distributed normally about the mean and the value of 1 .96 (a = 0.05 ; 95% confi¬
dence level) was used as a measure of statistical significance. It was assumed that
all values of d greater than 1.96 show statistically significant differences in the
populations of soil bacteria capable of excreting specific amino acids.

In like manner, differences in indices of production of amino acids by these
microorganisms or groups of microorganisms can be evaluated by the following
equation:

where: X, Y = means of indices

2 2

Sx, Sy = variance of the measured values
n = number of values obtained

The value of S2 was determined from the equation

S2 = ^ [2(x2)-i(Sx)*]

The number Zis also assumed to be derived from a normally distributed population;
for a = 0.05 the value 1 .96 is used.

RESULTS

The data shown in Table 1 indicate that numbers of organisms nable of ex¬
creting amino acids and the extent of excretion.

TABLE 1

Percentage of Bacteria Isolated from Soil
Which Secrete Amino Acids Into the Growth Medium

Numerical value
of production

Amino acids

Ala

Arg

Asp

Glu

Leu

Lys

OM, 1,2, 3

91

87

88

79

94

69

1,2,3

83

47

76

42

74

43

2,3

45

18

42

20

36

27

1

38

29

34

22

38

16

0

9

13

12

14

6

31

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THE TEXAS JOURNAL OF SCIENCE

The data in Table 2 show the index of production of amino acids by 2 major
groups of bacteria isolated from the soil. The data indicate different levels of pro¬
duction of each of the 6 amino acids studied.

TABLE 2

Index of Production of Amino Acids
by Groups of Soil Microorganisms.

Index

Amino acids

Group of

microorganisms

Ala

Arg

Asp

Glu

Leu

Lys

Gram positive

1.63

0.79

1.50

0.71

1.33

0.78

Gram negative

1.24

0.78

1.15

0.77

1.35

0.95

DISCUSSION

Analyses of Amino Acid Production

The quantities of amino acids secreted by bacteria isolated from different soils
vary over a wide range. For example, 94% of the organisms tested (161/171)
showed some production of leucine. Seventy- four % showed production at levels
1,2, and 3; 36% at levels 2,3; and 18% at level 3 (Table 1). We infer from these
data that although the majority of bacteria isolated from the soil secrete leucine,
only a small number (31, 17 1 ; 18%) secrete sufficient quantities to support exten¬
sive growth of other microorganisms. Analysis of the differences observed indicated
that these differences are statistically significant at the 95% level of confidence
(Table 3).

The data in Table 2 also show that alanine, aspartic acid, and leucine are pro¬
duced in larger quantities than arginine, glutamic acid, and lysine. The difference
between these 2 groups is statistically significant but the differences seen within
each group (i.e., alanine- aspartic acid, alanine-leucine, aspartic acid-leucine, etc.)
are not. The values for Z shown in Table 4 confirm this observation at the 95%
confidence level.

When the values obtained were analyzed with respect to Gram reaction of the
organisms studied, it was seen (Table 2) that both groups, Gram positive and Gram
negative, secreted greater amounts of alanine, aspartic acid, and leucine than of
arginine, glutamic acid, and lysine. This difference was statistically significant but
the differences observed within each group were not. From this we infer that the
type and quantity of amino acids secreted by Gram positive bacteria is not differ¬
ent from that secreted by Gram negative bacteria and that certain amino acids
are secreted in relatively large quantities by both groups.

PRODUCTION OF AMINO ACIDS

125

TABLE 3

Values of d for Comparison of Percentages of Bacteria
Capable of Excreting Amino Acids

Amino

acid

d for difference*

0, 1,2,3

vs.

1,2,3

1,2

vs.

2,3

2,3

vs.

3

Alanine

1.949

7.372

5.242

Arginine

8.105

2.698

0.901

Aspartic Acid

2.740

6.379

6.241

Glutamic Acid

6.302

4.315

3.176

Leucine

4.662

6.864

3.336

Lysine

2.732

2.709

2.915

♦Values of d greater than 1.96 represent statistically significant differences at the 95% con¬
fidence level.

TABLE 4

Values for Z for Index of Amino Acid Production
by all the Organisms Tested*

Ala

Arg

Asp

Glu

Leu

Lys

Ala

__

6.548

1.088

6.651

1.220

4.869

Arg

6.548

-

4.802

0.400

4.513

0.707

Asp

1.088

4.802

-

5.226

0.161

3.672

Glu

6.651

0.400

5.226

-

4.882

1.065

Leu

1.220

4.513

1.161

4.882

-

3.435

Lys

4.869

0.707

3.672

1.065

3.435

-

♦Values of Z greater than 1.96 represent statistically significant differences at the 95% con¬
fidence level.

SUMMARY

A simple technique was used to study the secretion of alanine, arginine, aspar¬
tic acid, glutamic acid, leucine, and lysine by 171 different bacteria isolated from
soils in the provinces of Barcelona and Alicante in Spain. Statistical analyses of
the data obtained showed that a large portion of soil isolates excrete amino acids
into the growth media but that only a much lower percentage secrete sufficient
quantities to support abundant growth of auxotrophic bacteria.

LITERATURE CITED

Azcon, R J . M . Barea, and V. Callao, 197 3 — Inoculacio n conjunta de microorganismos movil-
izadores de fosforo y Rhizobium en cultivos enarenados de judia. Microbiol. Expanola,
26:31.

126

THE TEXAS JOURNAL OF SCIENCE

Brown, M. E., 1974-Seed and root bacterization. Ann. Rev. Phytopathol., 12:181.

Guinea, J., 1970-Analisis colonial de la produccion de A. glutamico por Citrobacter inter¬
medium C3. Microbiol. Espanola, 23:13.

Steele, B. F., H. E. Sauberlich, M. S. Reynolds, and C. A. Baumann, 1949-Media for Leuco-
nostoc mesenteroides P-60 and Leuconostoc citrovorum 8081./. Biol. Chem., 177:5 33.

Udaka, S., 1960-Screening method for microorganisms accumulating metabolites and its
use in the isolation of Micrococcus glutamicus. J. Bacteriol., 79:754.

Vagnerova, K., and V. Vancura, 1962 -Production and utilization of amino acids by various
species of rhizosphere bacteria. Fol. Microbiol., 7:55.

Voznyakouskaya, Y. M., 1963-Choice of microorganisms for use in the composition of bac¬
terial fertilizers. Mikrobiologiya, 32:168.

Yamada, K., S. Kinoshita,T. Tsunoda, and K. Aida, 197 2 -The Microbial Production of Amino
Acids. Haldsted Press, New York.

THE BATS OF EAST TEXAS

by DAVID J. SCHMIDLY, KENNETH T. WILKINS,

RODNEY L. HONEYCUTT, and BERNARD C. WEYNAND

Department of Wildlife and Fisheries Sciences,

Texas A&M University, College Station 77843

ABSTRACT

Information concerning the distribution of bats in East Texas is scanty at best. Bailey
(1905), in his Biological Survey of Texas, recorded only 4 bats from east Texas. McCarley
(1959) presented data on 84 specimens of 8 species from 12 counties but recent changes in
the taxonomy of bats make the paper outdated and difficult to use. Davis (1974) mapped
county locality records, gave a recent taxonomic arrangement of the species, and summarized
much of the natural history information for bats in the area. His book does not give exact
collecting localities within counties, institutions where specimens are deposited, or literature
references.

In an attempt to document all records of bats collected in East Texas, we have examined
749 specimens from 29 counties. Also, for the past 2 years, one of us (Weynand) studied the
distribution of bats on a selected area in Newton County in extreme eastern Texas. Information
is given herein for 12 species of bats, including collecting localities and the institutions where
specimens are deposited. In addition, pertinent natural history information on bats from
Newton County is presented.

INTRODUCTION
Region Studied

The region herein defined as “East Texas” is essentially the same as that de¬
limited by McCarley (1959). The eastern and southern boundaries are the Texas-
Louisiana state line and the Gulf of Mexico, respectively (Figure 1). The Brazos
River bounds the southern half of the western limits of the area. The remaining
boundary is a line running from the Brazos River in Robertson County north¬
eastward to the Red River in Lamar County and thence to the Texas-Louisiana
state line.

East Texas contains a portion of 2 biotic provinces— the western extremities
of the Austroriparian and the eastern half of the Texan (Blair, 1950). The Austro-
riparian Province includes 4 of Tharp’s (1926) vegetational regions: oak-hickory,
long-leaf pine, coastal prairie, and pine-oak forest (Figure 2). Only 2 bats (Pipistrel-
lus subflavus and Lasiurus borealis) were among the 47 species of mammals listed
for the Austroriparian Province by Blair (1950). The portion of the Texas Province
in East Texas included parts of Tharp’s (1926) coastal prairie, oak-hickory, and
pine-oak regions. Of the 49 species of mammals occurring in this province, 41
also occur in the Austroriparian (Blair, 1950). No bats are listed as members of
the Texan fauna by Blair (1950).

Accepted for publication: March 31, 1976.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

128

THE TEXAS JOURNAL OF SCIENCE

Figure 1. The East Texas region (dark area of the Texas map; counties included are to
the right of the heavy line).

BATS OF EAST TEXAS

129

Figure 2. Map of East Texas illustrating biotic provinces and vegetational regions within
the area. The heavy broken line separates the Texan (left of the line) from the
Austroriparian (right of the line) Biotic Province. Solid lines separate the 4
vegetational regions: (A) oak-hickory; (B) pine- oak forest; (C) long-leaf pine;
and (D) coastal prairie.

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THE TEXAS JOURNAL OF SCIENCE

Newton County is situated in extreme eastern Texas on the Texas- Louisiana
border (Figure 1). The mammal fauna on the proposed site of a nuclear power
plant near Burkeville was inventoried during 1972-1974 (Inglis, et al, 1974).
Using conventional collecting techniques (including use of guns and mist nets),
bats were sampled on the power plant site in and over essentially 4 distinct situ¬
ations: streams in bottomlands, permanent man-made ponds in forested areas,
flyways in forested areas, and man-made structures such as old wells, cisterns,
and abandoned buildings.

Accounts of Species

Specimens examined in the account below are deposited in the following col¬
lections (museum abbreviations given in parentheses): Texas Cooperative Wildlife
Collection, Texas A&M University (TCWC); Stephen F. Austin State University
Vertebrate Collection (SFAVC); University of Texas at Arlington Collection of
Vertebrates (UTAVC); The Museum, Texas Tech University (TTU); Museum of
Zoology, Louisiana State University (LSUMZ); Bird and Mammal Laboratories,
U.S. Bureau of Sport Fisheries and Wildlife (BS); The University of Kansas, Museum
of Natural History (KU); and University of California, Museum of Vertebrate
Zoology, Berkeley (MVZ). The use of common names and species accounts follow
the sequence of species given in Jones, et al., (1975). Counties are listed from
west to east and north to south. Unless otherwise indicated, “additional records”
are from the literature.

Myotis lucifugus (Le Conte). Little Brown Bat.

This species has not been previously recorded from East Texas. The only other
record from Texas is that of a single specimen collected prior to 1900 at Fort Hancock
in Hudspeth County. This specimen was referred to the subspecies M. I occultus
by Findley and Jones (1967). A Newton County specimen is clearly referable to
M. 1. lucifugus on both geographical and morphological grounds. This specimen
was taken along Mill Creek under ecological conditions described below for
M. austroriparius. The nearest locality from which this subspecies has been recorded
is in central Arkansas, approximately 200 mi northeast of Newton County (Sealander
and Young, 1955).

Specimens examined: total, 1. Newton Co.: 12 mi N Burkeville (TCWC).

Myotis austroriparius (Rhoads). Southeastern Myotis.

The Southeastern Myotis, which ranges from the Ohio River Valley into the
southeastern states, reaches its western limits in extreme eastern Texas. Although
this species is abundant in Louisiana (Lowery, 1974), previous collecting records
suggest it is rare in East Texas. Packard (1966) reported the first Texas specimen
from Bowie County; Mi chael,ef a/., (1970) reported a single specimen from Panola
County. We follow LaVal (1970) in considering M. austroriparius asmonotypic;
thus, no subspecific assignment has been made for Texas specimens.

BATS OF EAST TEXAS

131

The frequency with which this bat was taken in Newton County suggests it
may be more common in East Texas than previously suspected. Specimens were
obtained from Mill and Indian creeks, 12 and 8.5 mi N Burkeville, respectively.
Both creeks are slow-moving narrow waterways surrounded by buttonbush and
blackberry bushes. The adjacent woods consist of large upland areas of loblolly
and shortleaf pines, narrow beach magnolia bottoms, and stands of hardwood
trees (white oak, red oak, and hickory). The crown closure above the creeks
ranged from 50 to 100% and the trees ranged in height from 50 to 100 ft.

Specimens examined: total, 9. Panola Co.: 8mi SWGary, 1 (TCWC). Newton Co.:
12 mi N Burkeville, 5 (TCWC): 8.5 mi N Burkeville, 2 (TCWC). Bowie Co.:
New Boston, 1 (TTU).

Lasionycteris noctivagans { Le Conte). Silver-Haired Bat.

The only record of the silver-haired bat in East Texas is that of a female found
alive hanging from a fence in a residential section of Galveston on 24 September 1975
(Martin , In Press). The nearest place in Texas where this species has been recorded
is from Bandera County which is over 300 airline mi west of Galveston (Blair,
1952). Since this species’ migratory habits are well-known, the Galveston speci¬
men probably represents a migrant. Silver-haired bats have been observed migrating
at sea along the Main ^oast (Lowery, 1974).

Specimens examined: total, 1. Galveston Co.: Galveston (TCWC).

Pipistrellus subflavus (F. Cuvier). Eastern Pipistrelle.

This species has been recorded from 10 East Texas counties. It has been collected
in every month of the year and is apparently a permanent resident of the area. In
Newton County, it was commonly collected along streams in bottomlands and in
flyways in forests. Specimens from East Texas are referable to the subspecies
P. s. subflavus on both morphological and geographical grounds.

Specimens examined: total, 42. Rusk Co.: 4 mi SE Gary,4(SFAVC). Anderson Co.-
Long Lake, 3 (BS). Panola Co.: Lake Murvaul, 1 (SFAVC). Shelby Co.: 15 mi N Center,

I (TTU); 14 mi N Center, 4 (SFAVC); cave between Timpson and Gary, 1 (SFAVC);
no specific locality, 6 (LSUMZ). Nacogdoches Co.: cave 4 mi SW Garrison, 7 (SFAVC),
15 mi E Nacogdoches-Garrison Hwy, 1 (SFAVC); 1 mi S Cushing, 1 (SFAVC);
Hwy 204, 10 mi E Cushing, 1 (SFAVC); Nacogdoches, 1 (SFAVC). Walker Co.:

II mi NW Waverly, 1 (TMM). Polk Co.: 4 mi E Livingston, 2 (TCWC); 3 mi S
Livingston, 1 (TCWC). Newton Co.: 9.3 mi N Burkeville, 1 (TCWC); 12 mi N
Burkeville, 2 (TCWC); 11.5 mi NBurkeville, 1 (TCWC); 8.5 mi N Burkeville,
1 (TCWC). Harris Co.: Huffman, 1 (TCWC). Galveston Co.: Clear Creek, 1 (BS).

Eptesicus fuscus (Palisot de Beauvois). Big Brown Bat.

The big brown bat has been recorded from only 7 counties in East Texas and
all recorded specimens are referable to the subspecies f fuscus. All collecting
records are within the pine-oak forest and long-leaf pine vegetational regions.

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THE TEXAS JOURNAL OF SCIENCE

Available evidence suggests the favored roosts of this species in East Texas are
man-made structures. In July 1975, we located a colony of approximately 35 big
brown bats in a crack between a brick chimney and an old house within the city
limits of Gilmer, Upshur County. Specimens have been collected from East Texas
in all months except November and December.

Specimens examined: total, 27. Upshur Co.: Gilmer, 13 (TCWC). Marion Co.:
Jefferson, 1 (BS). Nacogdoches Co.: SFA Campus, 4 (SFAVC); 1 (TTU); Nacogdoches,

1 (SFAVC). Trinity Co.: Trinity, 1 (TCWC). Walker Co.: 2 mi SW Huntsville,

2 (TCWC). San Jacinto Co.: 5 mi NW Cleveland, 2 (TTU). Hardin Co.: Sour Lake,
1 (BS), Grady, 1 (BS).

Lasiurus borealis (Muller). Red Bat.

This species is one of the most common bats in East Texas. It has been recorded
from 23 counties and in all major vegetation regions. In Newton County, red bats
were commonly collected along streams in bottomlands, in fly ways in forests,
and around man-made ponds. They have been collected at all seasons of the year
and appear to be permanent residents of the area. East Texas specimens are referable
to the subspecies L. b. borealis.

Specimens examined: total, 137. Lamar Co.: Arthur City, 3 (BS); Paris, 3 (BS).
Red River Co.: Clarksville, 1 (BS). Bowie Co.: 8 mi N New Boston, 1 (TCWC).
Van Zandt Co.: 1 mi SW Grand Saline, 1 (TTU). Smith Co.: Tyler, 1 (TCWC).
Rusk Co.: 1.6 mi NE New London, 2 (SFAVC). Marion Co.: Jefferson, 2 (BS).
Panola Co.: 6 mi E Mt. Enterprise, 1 (SFAVC); 7.5 mi ENE Carthage, 4 (TCWC).
Cherokee Co.: Maydelle, 1 (SFAVC); 3 mi W Forrest, 1 (SFAVC), 1 (TTU).
Shelby Co.: Choice, 1 (LSUMZ). Nacogdoches Co.: Martinsville, 1 (SFAVC);
SFA Campus, 5 (SFAVC); SFA Experimental Forest, 2 (TTU); Sam Rayburn
Reservoir, 1 (SFAVC); Nacogdoches, 1 (SFAVC), 5 (TTU). Angelina Co.: 23 mi
S Nacogdoches, 1 (TTU); 1 mi S Lufkin, 1 (SFAVC); no specific locality, 1 (SFAVC).
Trinity Co.: 1.3 mi E Trinity, 1 (TCWC); Trinity, 1 (TCWC). Walker Co.: 16mi SW
Huntsville, 1 (TCWC); 2 mi NE Huntsville, 2 (TCWC); Huntsville, 1 (TCWC).
San Jacinto Co.: San Jacinto River, Farm Rd 995,8 (TTU); 5 mi NW Cleveland,
13 (TTU); San Jacinto River, Farm Rd 945, 13 (TTU). Jasper Co.: Bouton Lake,

1 (UTAVC). Newton Co.: 7.5 mi N Burke ville, 5 (TCWC); 1 1.5 mi N Burke ville,

2 (TCWC), 7 mi N Burkeville, 1 (TCWC); Newton, 1 (TTU). Hardin Co.: Sour Lake,
1 (BS). Brazos Co.: Bryan, 3 (TCWC); College Station, 11 (TCWC); 6.5 mi SE
College Station, 4(TCWC); 6.5 mi SW College Station, 1 (TCWC). Montgomery Co.:
20 mi SW Huntsville, 4 (TMM). Liberty Co.: 12 mi N Dayton, 3 (TTU); 20 mi
NW Liberty, 1 (BS). Harris Co.: Houston, 4 (TCWC); no specific locality, 1 (SFAVC).
Jefferson Co.: Port Arthur, 6 (SFAVC), 6 (TTU). Galveston Co.: Bolivar Peninsula,
7.5 mi NE Ferry landing, 1 (TMM).

Lasiurus seminolus (Rhoads). Seminole Bat.

The seminole bat is abundant throughout the pine-oak and long-leaf pine
forest regions of East Texas; it is less common in the oak -hickory region. Since
it has been taken in East Texas from February through November, it probably

BATS OF EAST TEXAS

133

is a year-long resident in the area. In Newton County, this bat was collected along
Mill and Indian creeks under ecological conditions as described in the account of
M. austroriparius. This species is monotypic.

Specimens examined: total, 71. Panola Co.: 6 mi EMt. Enterprise, 1 (SFAVC);
7.5 mi ENE Carthage, 1 (TCWC). Cherokee Co.: 3 mi W Forrest, 1 (SFAVC).
Shelby Co.: 4 mi S Joaquin, 1 (SFAVC); 2.3 mi SE Patroon, 1 (SFAVC). Nacogdoches
Co.: 9.5 mi SW Nacogdoches, 1 (TMM); Nacogdoches, 6 (SFAVC); SFA Campus,
2 (SFAVC), 1 (TTU). Trinity Co.: Trinity, 1 (TCWC). Sabine Co.: 5 mi N Geneva,
1 (TCWC); no specific locality, 1 (SFAVC). San Jacinto Co.: 5 mi NW Cleveland,
15 (TTU), Farm Rd 945 San Jacinto River, 10 (TTU). Polk Co.: 3 mi S Livingston,
1 (TCWC); 4 mi E Livingston, 1 (TCWC); no specific locality, 2 (SFAVC). Newton
Co.: 8.5 miN Burkeville, l(TCWC); 11.5 mi NBurkeville, 6 (TCWC). Montgomery
Co.: 5 mi S Richards, 1 (TCWC). Harris Co.: Houston, 10 (LSUMZ); no specific
locality, 4 (TCWC). Liberty Co.: 12 mi N Dayton, 1 (TTU). Jefferson Co.: 1 1.7 mi
W Sabine Pass, Hwy 87, 1 (TCWC).

Additional records: Rusk Co. (McCarley, 1959); Harris Co. (Olsen, 1966).

Lasiurus cine reus (Palisot de Beauvois). Hoary Bat

This species has not been reported previously from East Texas, although it is
relatively common in other parts of Texas and throughout Louisiana. The East
Texas specimen, which is referable to the subspecies L. c. cinereus, was taken
near Cleveland in San Jacinto County on 13 January 1974. The nearest recorded
Texas specimen is from McLennan County, 140 miles northwest of Cleveland.

Specimens examined: total, 1 . San Jacinto Co.: 5 mi NW Cleveland, 1 (TTU).

Lasiurus intermedius (H. Allen). Northern Yellow Bat.

Collecting records indicate this species is one of the rarest of the tree bats in
East Texas. Until recently, it was known only from Harris County. However, on
24 April 1975, one of us (Wilkins) netted an adult female over a narrow pool of
water in Brazos County. The uterus of this female contained four embryos which
were each 5 mm in crown -rump length. According to Hall and Jones (1961),
East Texas specimens are referable to the subspecies/,. i.floridanus (Miller) rather
than to L. i. intermedius (H. Allen) which occurs in South Texas.

Specimens examined: total, 8. Brazos Co.: 8 mi SW College Station, 1 (TCWC).
Harris Co.: Houston, 1 (KU), 2 (MVZ), 2 (TCWC); 4 mi N Huffman, 1 (TCWC).
Fort Bend Co.: Brawner Ranch, 2 mi S Dewalt, 1 (TTU).

Nycticeius humeralis (Rafinesque). Evening Bat.

This species is widely distributed throughout East Texas and is probably the
most common bat in the area. It occurs in all major vegetation regions and is
represented in collections by more than 150 specimens collected from 16 counties.
In Newton County we collected specimens along streams in bottomlands and
around man-made ponds. In Nacogdoches County, evening bats have been collected

134

THE TEXAS JOURNAL OF SCIENCE

in all months of the year, although records indicate they are more abundant in
summer and fall than in the winter (Table 1). Specimens from East Texas are
referable to the subspecies N. h. humeralis on both geographical and morphological
grounds.

TABLE 1

Monthly Collecting Records of Nycticeius humeralis from Nacogdoches County, Texas.

Month

Males

Females

Sex Unknown*

Totals

J

1

0

1

2

F

2

0

1

3

M

0

4

0

4

A

1

1

3

5

M

0

1

0

1

J

1

3

5

9

J

1

9

9

19

A

4

3

0

7

S

6

4

1

11

0

6

5

2

13

N

5

2

1

8

D

4

0

0

4

TOTALS

31

32

23

86

*Sex not recorded on specimen labels.

Non -geographic variation has not been previously investigated in Nycticeius
humeralis. We utilized the large sample of evening bats from Nacogdoches County
to determine the extent of individual and sexual variation in this species. Of the
10 external and cranial characters we examined, all but 2 (interorbital constriction
and width across the upper molars) exhibited significant secondary sexual dimorph¬
ism (see Table 2). In all characters, females averaged larger than males. Coeffi¬
cients of variation (CV’s) of external measurements averaged larger than those of
cranial features. Of the cranial measurements, length of palate had the highest CV.
With the exception of cranial breadth and width across the upper molars, CV’s
of males averaged larger than those of females.

Specimens examined: total, 160. Lamar Co.: Paris, 3 (BS); Arthur City, 1 (BS).
Bowie Co.: 8 mi N New Boston, 1 (TCWC); Texarkana, 3 (BS). Marion Co.: Jefferson,
1 (BS). Rusk Co.: 1.6 mi NE New London, 6 (SFAVC). Anderson Co.: Palestine,
Gus Engling Wildlife Management Area, 1 (TCWC). Cherokee Co.: Bates Farm,
3 mi W Forrest, 3 (SFAVC); Cherokee Co., 1 (SFAVC). Nacogdoches Co.: Nacogdoches,
33 (TTU), 56 (SFAVC); SFA Campus, 12 (SFAVC); 1/2 mi E Nacogdoches, 1
(SFAVC); 1 mi E Nacogdoches, 1 (SFAVC); 11 mi SW Nacogdoches, 1 (TTU).

BATS OF EAST TEXAS

135

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Shelby Co.: Shelbyville, 2 (SFAVC). Sabine Co.: Pineland, 1(TCWC). Trinity Co.:
Trinity, 5 (TCWC). San Jacinto Co.: 5 mi NW Cleveland, 7 (TTU); Jet Farm Rd 945,
San Jacinto River, 1 (TTU). Jasper Co.: Jasper, 3 (BS). Newton Co.: 12 mi N
Burkeville, 1 (TCWC); 11.5 mi N Burkeville, 2 (TCWC); 9.2 mi N Burkeville, 3 (TCWC);
Brazos Co.: 3 mi S Bryan, 1 (TCWC); 2 mi SE Bryan, 1 (TCWC); College Station,

136

THE TEXAS JOURNAL OF SCIENCE

5 (TCWC); 8 mi SE College Station, 1 (TCWC). Montgomery Co.: 5 mi S Richards,
1 (TCWC). Harris Co.: Houston, 1 (BS). Fort Bend Co.: Brawner Ranch, 3 mi S
Dew alt, 1 (LSUMZ).

Additional records, Harris Co.: Houston (Olsen, 1966).

Plecotus rafinesquii (Lesson). Rafinesque’s Big-eared Bat.

This is another species that reaches the western limits of its range in East Texas.
Michael and Birch (1967) recorded the first Texas specimens from Polk County.
This bat was relatively common in Newton County where it seemed to prefer
wells and cisterns as roosting sites. Of the 5 bats collected there, 4 were taken in
wells and cisterns; 1 was obtained in a mist net strung amongjunipers and loblolly
pines. Specimens were obtained in November, December, and June. Our obser¬
vations support the conclusions of Jones and Suttkus (1975) that P. rafinesquii
is found in small numbers at scattered localities. This species is monotypic.

Specimens examined: total, 36. Marion Co.: 27 mi W Jefferson, 1 (SFAVC).
Nacogdoches Co.: Nacogdoches, 14 (SFAVC); 10 mi SE Nacogdoches, 5 (TCWC).
Sabine Co.: 3 mi NE Milam, 3 (SFAVC). Polk Co.: 1 mi S Neches River, 3 mi E
Hwy 59, 2 (TCWC); no specific locality, 5 (SFAVC). Newton Co.: 6 mi N Burke-
ville, 1 (TCWC); 7 mi N Burke ville, 1 (TCWC); 10 mi N Burke ville, 2 (TCWC);
9.5 mi N Burkeville, 1 (TCWC). Hardin Co.: 7 mi SE Silsbee, 1 (TCWC).

Tadarida brasiliensis (Saussure). Brazilian Free-tailed Bat.

Two kinds of Brazilian Free-tailed Bats have been reported from the southern
United States and both are known from East Texas. T. b. mexicana (Saussure) is
a migratory, cave -dwelling (sometimes house -dwelling) bat of the southwestern
United States which reaches the western limits of its range in East Texas. These
bats do not hibernate, and during June and July the females give birth to their
young in caves where females congregate in extremely large numbers. The other
form, T. b. cynocephala (LeConte), is a sedentary, tree- or house-dwelling bat of
the southeastern United States which reaches the western limits of its range in
East Texas. This bat hibernates during the winter and gives birth to its young during
June and July.

The taxonomic status ofT. b. mexicana and T. b. cynocephala has been debated
for years. Carter (1962) and Davis (1974) consider them as distinct species, but
certain recent authors (for example, Barbour and Davis, 1969; Lowery, 1974)
have preferred to consider them both as subspecies of T. brasiliensis as was pro¬
posed by Schwartz (1955).

The primary distinctions between cynocephala and mexicana are ethological,
although some morphological dissimilarity is evident between the two (Carter,
1962). Carter (1962) has suggested that cynocephala and mexicana are seasonally
sympatric in East Texas because during early March, when both forms are pre¬
sumably copulating, some mexicana pass through the western edge of the geo¬
graphic range of cynocephala. If intergradation is possible, it should occur in an

BATS OF EAST TEXAS

137

area between the forested region of East Texas and the Balcones Escarpment, and
the resulting populations should be morphologically and ethologically intermediate
between T. cynocephala and T. mexicana (Carter, 1962).

In order to allocate and define populations of free-tailed bats inhabiting East
Texas, we found it necessary to study geographic variation among populations
from Texas and the southeastern United States. A total of 359 specimens was
examined from the following places (sample sizes given in parenthesis): 1. Florida
(21 females); 2. Georgia (14 males, 29 females); 3. Alabama (6 males; 18 females);
4. Sabine Co., Texas (58 males, 51 females); 5 . Nacogdoches Co., Texas (14 males,
10 females); 6. Anderson Co., Texas (10 males); 7. Harris Co., Texas (6 males,
15 females); 8. Grimes Co., Texas (5 males, 5 females); 9. Brazos Co., Texas
(10 males, 8 females); 10. Kerr Co., Texas (16 females); 1 1. Comal Co., Texas
(13 males, 23 females); 12. Val Verde Co., Texas (16 males, 7 females). For each
specimen, we recorded (to the nearest 0.01 mm using dial calipers) the following
cranial measurements (numbers refer to characters in Table 2): (1) greatest length
of skull, (2) zygomatic breadth; (3) greatest width across the braincase; (4) inter¬
orbital constriction; (5) length of maxillary toothrow (C-M3); (6) width across
molars (M3-M3 ); (7) length of palate measured from front of incisors to posterior-
most point near midline of palate; (8) greatest length of mandible; and (9) length
of mandibular toothrow (C-M3 ).

Since earlier studies have shown that significant sexual dimorphism exists in
populations of T. brasiliensis, we treated the sexes separately in analyzing geo¬
graphic variation. To assess the degree of divergence among samples, we used a
multivariate analysis of variance (MANOVA) and a canonical analysis program in
the Statistical Analysis System (SAS) designed and implemented by Barr and
Goodnight (Service, 1972). Canonical analysis of the data aimed at providing
weighted combinations of the measurements which maximize the distinction
between the groups (for a detailed discussion of the methodology, see Schmidly
and Hen dricks , In Press) . Sample means and individual specimens were plotted on
those canonical variates that account for the greatest fractions of total variation.
The relative importance of each original variable to a particular canonical variate
was computed by multiplying the vector variable coefficient by the median value
of the dependent variable, summing all variable values for a particular vector,
and then computing the percent of relative importance of each variable per vector.

The first 3 canonical variates were computed from the variance -covariance
matrix among the 9 cranial characters for all samples of males and females separately.
For males, the first canonical variate expresses 60.6% of the phenetic variation;
the second, 14.9; and the third, 10.1. These same values for females are 59.2,
22.8 and 7.1%, respectively. Two-dimensional plots of the first 2 canonical variates
(including the mean and 2 standard errors either side of the mean for each sample)
for males and females are shown in Figure 3.

Examination of Figure 3 (males) reveals that the primary separation of samples
currently defined as cynocephala and mexicana occurs along Vector I. Samples
of cynocephala (2, 3, 4, 5, and 7) cluster on Side A of the graph and include 3

138

THE TEXAS JOURNAL OF SCIENCE

I

Figure 3. Projections of the first 2 canonical vectors illustrating the phenetic position of
male (top) and female (bottom) samples of Tadarida brasiliensis. Numbers are
positioned at the mean value for each sample in the character space; the ellipse
surrounding each number represents 2 standard errors around the mean. See
text for key to samples.

samples (4, 5, and 7) from Sabine, Nacogdoches, and Harris Counties in East Texas.
Samples of mexicana (6, 9, 1 1, and 12) cluster on Side B of the graph and include two
samples (6 and 9) from Brazos and Anderson Counties in East Texas. Sample 8 from

BATS OF EAST TEXAS

139

Grimes County in East Texas is intermediate and overlaps both the cynocephala and
mexicana clusters. Those characters which have the greatest influence on Vector I
and hence which contribute most to the separation of cynocephala and mexicana
are greatest length of the skull, zygomatic breadth, and breadth of the cranium
(Table 3). In all of these features, specimens of cynocephala average larger than
specimens of mexicana (see Table 4).

TABLE 3

Variable Coefficients for Canonical Variates I and II with
an Estimate of the % Influence of Each Variable on Each
Vector for Samples of Male and Female Tadarida brasiliensis.

MALES

Vector I Vector II

Variable

Percent

Variable

Percent

Character

Coefficient

Influence

Coefficient

Influence

1

0.162

31.49

-0.026

4.32

2

0.227

25.76

-0.007

0.68

3

0.134

14.24

-0.091

8.29

4

0.167

7.52

0.221

8.51

5

0.008

0.55

-0.096

5.63

6

-0.083

6.50

-0.022

1.48

7

0.038

2.49

-0.183

10.25

8

0.043

5.91

0.388

45.71

9

0.047

5.53

-0.236

15.11

FEMALES

1

0.067

15.90

0.034

14.56

2

0.211

29.38

0.008

2.02

3

0.102

13.37

-0.007

1.66

4

0.101

10.10

0.185

18.45

5

0.026

2.60

0.077

11.74

6

0.036

3.49

-0.019

3.33

7

-0.129

12.90

0.191

27.38

8

-0.038

3.80

0.035

10.60

9

0.147

13.46

-0.062

10.25

Examination of Figure 3 (females) reveals the same general pattern as that
observed for males. Again the major separation of cynocephala and mexicana is
along Vector I. Samples of cynocephala (1,2, 3, 4, 5, and 7) congregate on Side A
of the graph and samples of mexicana (9,10,11, and 12) on Side B. Once again
Sample 8 from Grimes County in East Texas is intermediate and overlaps both
the cynocephala and mexicana clusters. Characters which have the highest percent

140

THE TEXAS JOURNAL OF SCIENCE

TABLE 4

Selected Cranial Measurements (Mean ± 1 Standard Devia¬
tion) of Samples of T. brasiliensis from the Southeastern
United States.

Sex

N

Greatest length
of skull

Zygomatic

breadth

Greatest width
across braincase

Length of

mandibular toothrow

Georgia (cynocephala)

Male

14

17.42 ±0.17

10.19 ±0.14

9.61 ±0.12

6.67 ±0.16

Female

29

17.02 ±0.28

10.08 ±0.17

9.46 ±0.16

6.61 ±0.18

Sabine Co., Texas (< cynocephala )

Male

58

17.50 ±0.23

10.30 ± 0.18

9.60 ±0.20

6.78 ±0.20

Female

51

17.10 ±0.25

10.06 ±0.15

9.49 ±0.20

6.57 ±0.23

Grimes Co., Texas (intergrades)

Male

5

17.29 ±0.24

9.85 ±0.40

9.30 ±0.22

6.60 ±0.28

Female

5

17.06 ±0.11

10.00 ±0.18

9.30 ±0.12

6.50 ±0.12

Brazos Co., Texas ( mexicana )

Male

10

16.65 ±0.07

9.88 ±0.11

9.05 ±0.07

6.50 ±0.07

Female

8

16.71 ±0.20

9.74 ±0.19

9.15 ±0.20

6.32 ±0.12

Comal Co., Texas (mexicana)

Male

13

17.01 ±0.25

9.89 ±0.26

9.23 ±0.15

6.45 ±0.17

Female

23

16.69 ±0.26

9.74 ±0.18

9.20 ±0.12

6.38 ±0.21

influence on Vector I for females include zygomatic breadth, greatest length of
skull, length of the lower mandibular toothrow, and breadth of cranium (Table 3).
In all of these features, specimens of cynocephala average larger than specimens
of mexicana (Table 4).

We interpret the morphological evidence as indicating that although samples
of cynocephala and mexicana are somewhat morphologically distinct, there is
evidence of morphological intergradation where their ranges come in contact in
East Texas. The sample from Navasota in Grimes County is morphologically
intermediate and overlaps both the sample of mexicana from College Station in
Brazos County and samples of cynocephala from eastern Texas. On the basis of
the morphological analysis, we find it difficult to justify recognizing cynocephala
and mexicana as separate species. However, this interpretation does not eliminate
the possibility that ethological isolation, which would completely prevent them
from interbreeding, might exist between the two forms. Supposedly, copulation
in the migratory mexicana occurs in Mexico, whereas copulation in the nonmigratory
cynocephala takes place within its range in the United States (Barbour and Davis,

BATS OF EAST TEXAS

141

1969). Spenrath and LaVal (1974) studied the population dynamics of the colony
of T. b. mexicana at College Station and found it to be somewhat atypical in
comparison to other colonies of this bat in that males remain in the colony through¬
out the year. We have studied this colony since 1971 and our data support Spenrath
and LaVal ’s (1974) conclusions. The year-round presence of males of T. b. mexicana
at College Station, a locality only 100 mi from colonies of T. b. cynocephala in
eastern Texas, plus the fact that a population of bats morphologically intermediate
between mexicana and cynocephala is located at Navasota in Grimes County
strongly suggests that gene flow exists between populations of these 2 taxa. Ob¬
viously the behavior of the bats in Grimes County needs to be investigated to
support or refute our hypothesis. But until these studies are conducted, we feel
the data now available best support the conclusion that cynocephala and mexicana
are conspecific. We follow Schwartz (1955) in treating the 2 taxa as subspecies
of the wide-ranging species T. brasiliensis.

Specimens examined: total, 256. T. b. cynocephala . Cherokee Co.: 1 (SFAVC).
Shelby Co.: 20 mi S Center, 1 (SFAVC). Nacogdoches Co.: Nacogdoches, 59 (SFAVC),
13 (TTU); 1 1 mi SW Nacogdoches, 1 (TTU). Angelina Co.: Lufkin, 2 (SFAVC).
Sabine Co.: Pineland, 107 (TCWC); 11 mi N Hemphill, 1 (SFAVC). Harris Co.:
Houston, 20 (TCWC). T. b. mexicana. Anderson Co.: Palestine, 12 (TCWC).
Brazos Co.: College Station, 17 (TCWC). Grimes Co.: Navasota, 22 (TCWC).

Distributional Patterns

Distributional patterns of East Texas bats by biotic province and vegetational
regions are presented in Table 5. Of the 2 biotic provinces in East Texas, the
Austro riparian supports the most diverse chiropteran fauna. All 12 East Texas

TABLE 5

Distributions of East Texas Bats by Biotic Provinces and Vegetation Regions.

Species

Biotic Province

Vegetational Region

Austro-

Oak-

Pine-

Long-leaf

Coastal

riparian

Texan

Hickory

Oak

Pine

Prairie

Pipistrellus subflavus

X

X

X

X

X

X

Eptesicus fuscus

X

X

X

X

Lasiurus borealis

X

X

X

X

X

X

Lasiurus seminolus

X

X

X

X

Lasiurus cinereus

X

X

Lasiurus intermedius

X

X

X

X

Nycticeius humeralis

X

X

X

X

X

X

My otis austroriparius

X

X

X

My otis lucifugus

X

X

Lasiony cteris noctivagans

X

X

Plecotus rafinesquii

X

X

X

Tadarida brasiliensis

X

X

X

X

X

X

TOTALS

12

5

5

9

9

8

142

THE TEXAS JOURNAL OF SCIENCE

bats occur in this province whereas only 5 occur in the Texan Province. Seven
species occur only in the Austroriparian Province, whereas none are restricted to
the Texan Province. Among the 4 vegetational regions in East Texas, the western¬
most (oak-hickory) region supports the fewest species (5), whereas the pine-oak
and long-leaf pine regions (each with 9 species) support the most diverse fauna.
Eight species have been recorded from the coastal prairie region. Four species
( Pipistrellus subflavus, Lasiurus borealis , Nycticeius humeralis, and Tadarida
brasiliensis) are found throughout East Texas. Collecting records indicate that Myotis
lucifugus, Lasionycteris noctivagans, and Lasiurus cinereus are rare in the area
with only 1 county record for each.

ACKNOWLEDGEMENTS

We wish to thank the following curators who kindly allowed us to examine
specimens under their care: J. L. Darling, University of Texas at Arlington; Charles D.
Fisher, Stephen F. Austin University; Hugh H. Genoways, Texas Tech University;
Robert S. Hoffman, University of Kansas; George W. Lowery, Louisiana State
University; Robert F. Martin, Texas Memorial Museum, Austin; James L. Patton,
University of California, Berkeley; Don E. Wilson, Bird and Mammal Laboratories,
U. S. National Museum, Washington, D. C. Work on the Blue Hills site in Newton
County was supported by Bechtel Power Corporation and Gulf State Utilities
through the Texas Agricultural Experiment Station, Project 6024.

LITERATURE CITED

Bailey, V., 1905 -Biological survey of Texas. A. Amer. Fauna, 25:1.

Barbour, R. W., and W. H. Davis, 1969 -Bats of America. Univ. Press Kentucky Lexington,

286 pp.

Blair, W. F., 1950 -The biotic provinces of Texas. Tex. J. Sci., 2:93.

- , 1952-Bats of the Edwards Plateau in Central Texas. Tex. J. Sci. , 4:95.

Carter, D. C., 1962 -The systematic status of the bat Tadarida brasiliensis (I. Geoffrey) and
its related mainland forms. Unpublished Ph.D. dissertation, Texas A&M Univ.

Davis, W. B., 1914-Mammals of Texas. Texas Parks and Wildlife Dept., Austin, 294 pp.

Findley, J. S., and C. Jones, 1967 -Taxonomic relationships of bats of the species Myotis
fortidens, M. lucifugus, and M. occultus. J. Mamm., 48:429.

Hall, E. R., and J. K. Jones, 1961-North American yellow bats “ Dasypterus and a list of
the named kinds of the genus Lasiurus Gray. Publ. Univ. Kansas, Mus. Nat. Hist., 14:73.

Inglis, J. M., W. J. Clark, H. D. Irby, and D. M. Moehring, 1974 -Preconstruction ecological
and biological studies. Blue Hills Nuclear Power Plant Environmental Study. Texas A&M
Univ., 1300 pp.

Jones, J. K., D. C. Carter, and H. H. Genoways, 1975 -Revised Checklist of North American
Mammals North of Mexico. Occ. Papers, The Museum, Texas Tech Univ., 28:1.

BATS OF EAST TEXAS

143

Jones, C., and R. D. Suttkus, 1975-Notes on the natural history of Plecotus rafinesquii.
Occ. Papers Mus. Zool., La. State Univ., 47:1.

LaVal, R. K., 1970 -Infraspecific relationships of bats of the species Myotis austroriparius.
J. Mamm., 5 1:542.

Lowery, G. H., 191 4 -The Mammals of Louisiana and Its Adjacent Waters. La. State Univ.
Press, 565 pp.

Martin, C. O., 1976 -A noteworthy record of the silver-haired bat in Texas. Tex.JSci., In Press.

McCarley, H., 195 9 -The mammals of eastern Texas. Tex. J. Sci., 11:385.

Michael, E. D., and J. B. Birch, 1967-First Texas record of Plecotus rafinesquii. J. Mamm.,
48:672.

- , R. L. Whisennand , and G. Anderson , 1 970 - A recent record of Myotis austroriparius

from Texas. /. Mamm., 51:620.

Olsen, R. E., 19 66 -A heretofore unnoted collection of Texas mammals. Tex. J. Sci., 18:225.

Packard, R. L., 19 66 -Myotis austroriparius in Texas./. Mamm., 47:128.

Schmidly, D. J., and F. S. Hendricks, 1976-Systematics of the southern races of Dipodomys
ordii. Bull. S. Calif. Acad. Sci., In Press.

Schwartz, A., 1955 -The status of the species of the brasiliensis group of the genus Tadarida.
J. Mamm., 36:106.

Selander, J. A., and H. Young, 195 5 -Preliminary observations on the cave bats of Arkansas.
Proc. Arkansas Acad. Sci., 7:21.

Service, J., 1912-A User’s Guide to the Statistical Analysis System. Institute of Statistics,
Raleigh Division, North Carolina State University.

Spenrath, C. A., and R. K. LaVal, 1974— An ecological study of a resident population of
Tadarida brasiliensis in eastern Texas. Occ. Papers, The Museum, Texas Tech Univ., 21:1.

Tharp, B. C., 1926-Structure of Texas vegetation east of the 90th meridian. Univ. Texas,
Austin, Bull., 2606:1.

■

■

■b.

A KEY TO THE LICE OF MAN AND DOMESTIC ANIMALS

by DONALD W. TUFF

Department of Biology,

Southwest Texas State University,

San Marcos 78666

ABSTRACT

Dichotomous keys are given to the species of biting and sucking lice normally found on
man and domestic animals. Illustrations of diagnostic characters used in the keys are included.

INTRODUCTION

The following keys are designed primarily for the non -specialist to facilitate
the identification of the lice parasitizing man and domestic animals. The terms
used in most keys, particularly those unaccompanied by illustrations and often
poorly defined, are difficult to interpret.

It is hoped that the keys will fulfill two basic needs: first, make the identifi¬
cation of the most frequently encountered species of lice as simple as possible by
elimination of the seldom encountered species; and second, explain the more
important terms used in the keys by the use of an adequate number of illustrations.
Throughout the keys, I have avoided using characters of the genitalia to separate
species. More often than not, a person will have a specimen of the opposite sex
than that referred to in the key. Also, specimens must be in an exceptionally
good condition and properly mounted so that the characters of the genitalia can
be seen.

It is essential to remember that although the wording in a key is fixed, the
organisms are not and some variation within species’ populations may be encountered.

The keys will give an individual practice in identifying specimens of lice from
domestic animals when the specific host is unknown. Generally, specimens can be
identified simply by knowing the host, then confirmation of the determination
by comparing the specimen with descriptions and illustrations in the laboratory
is an easy matter. For the diagnosis of louse infestations, a host list, and photo¬
graphs of several of the more common species found on domestic animals, the
reader is referred to the works of La Page (1968) and Sloss (1973).

KEY TO THE ORDERS OF LICE

1 Anterior margin of head acute, rostrum present, mouthparts adapted

for piercing and sucking; opposing mandibles absent; tarsal claws
single, often large . . . Parti, ANOPLURA

Accepted for publication: November 30, 1976.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

146

THE TEXAS JOURNAL OF SCIENCE

1 Anterior margin of head generally broadly rounded, rostrum ab¬

sent, mouthparts adapted for chewing; opposing mandibles well
developed, on ventral surface; tarsal claws small, either single or
paired . Part II, MALLOPHAGA

Parti

KEY TO THE FAMILIES, GENERA AND SPECIES OF
ANOPLURA OF DOMESTIC ANIMALS

1 Eyes well developed, with distinct lens (Figure 1) .... Pediculidae, 3

l' Eyes vestigial or absent . . . 2

2( 1 ;) Paratergal plates of abdominal segments heavily sclerotized forming lateral

lobes (Figure 2); all legs of equal size (Figure 3) . . . Haematopinidae, 4

2 Paratergal plates of abdominal segments absent; if present, greatly re¬

duced and weakly sclerotized; first pair of legs smaller than second
and third pairs (Figure 4) . . . Linognathidae, 6

Family Pediculidae

3(1) All legs equal in size and shape, tarsal claws slender (Figure 5); abdominal
segments normal, with sclerotized paratergal plates, each plate bearing a spi-
acle (Figure 6) . . . (the head louse and body louse) Pediculus humanus

3' First pair of legs reduced (Figure 5), second and third pairs greatly

enlarged with robust tarsal claws (Figure 7); abdomen with 4 pairs of
prominent lateral tubercles (Figure 8); a group of 3 distinct spiracles
located dorsally on each side of the abdomen behind the third pair
of legs . (the crab louse) Pthirus pubis

Family Haematopinidae

4(2) Head 2 to 3 times as long as wide (Figure 9) . . 5

4f Head about as long as wide (Figure 10); paratergal plates tuberculated

(Figure 11) . . . (the shortnosed cattle louse) Haematopinus eurysternus

Another louse is recognized here,//, quadripertusus, the cattle tail louse. This

louse is ecologically isolated from H. erysternus, being found on the tail of cattle,

but is morphologically identical with H. erysternus.

5(4) Paratergal plates rounded (Figure 2); tarsal claws toothed (Figure 12)
. (the hog louse) Haematopinus suis

5' Paratergal plates tuberculate (Figure 11); tarsal claws simple, not

toothed (Figure 13) . . . . (the horse sucking louse) Haematopinus asini

Family Linognathidae

6(2;) Abdominal segments with only one transverse row of setae; abdominal

spiracles borne on small tubercles (Figure 14) ...... . .

. (the capillate cattle louse) Solenopotes capillatus

6' Abdominal segments with more than one transverse row of setae; ab¬
dominal spiracles not borne on tubercles (Figure 15) . 7

LICE

147

7(6/) Postantennal region laterally produced (Figure 16) .......... .

. . (on sheep and goats) Linognathus africanus

l' Postantennal region not laterally produced . . . 8

8(7') Head twice as long as broad; preantennal region as long as broad

(Figures 17, 18) . . . 9

8f Head about as long as broad or slightly longer; preantennal region

much broader than long (Figures 19, 20) . . . 10

9(8) Preantennal region elongate, apically acute (Figure 17a); lateral mar¬
gins of postantennal regions straight, appearing rectangular (Fig¬
ure 17b) ....... . (the long-nosed cattle louse) Linognathus vituli

9' Preantennal region acute (Figure 18a); lateral margins of postantennal regions

slightly convex (Figure 18b) . (the goat sucking louse) Linognathus stenopsis

10(8 ) Head as long as broad, preantennal region very short (Figure 19a), lateral

margins of postantennal region slightly convex (Figure 19b) .

. . . . (the sheep foot louse) Linognathus pedalis

lO* Head slightly longer than broad, preantennal region well developed, with

lateral margins straight and apex blunt, (Figure 20 a), lateral margins of post¬
antennal region parallel (Figure 20b) . . . .

. . . (the dog sucking louse) Linognathus setosus

Part II

KEY TO THE SUBORDERS, FAMILIES, GENERA AND SPECIES
OF MALLOPHAGA OF DOMESTIC ANIMALS

1 Antennae distinctly clubbed or capitate, concealed in a groove on un¬
derside of head (Figure 21) . . AMBLYCERA 2

1 Antennae not clubbed or capitate, project from lateral margins of head

(Figure 22) . . ISCHNOCERA 6

KEY TO THE FAMILIES OF THE SUBORDER AMBLYCERA

2(1) Underside of head with two long stout backward projecting spine¬
like processes; antennae clubbed (Figures 23, 24) ...... Boopidae

2' Underside of head with spine-like processes reduced or absent; antennae

not strongly clubbed (Figures 25 , 26) ......... Menoponidae 3

Family Boopidae

There is one species of Boopidae, Heterodoxus spininger, found in the United

States and occurs on dogs. H. spininger is easily recognized by the shape of the

head and the larger backward projecting spines. The body is covered with large

and numerous setae.

Family Menoponidae

3(2') Underside of head with 2 short spine-like processes (Figure 25) ... 4

3 Underside of head without spine-like processes . .

............. (the shaft louse [on chickens] ) Menopon gallinae

148

THE TEXAS JOURNAL OF SCIENCE

4(3)

4'

5(4')

5'

Abdominal tergites III-VTI each with one transverse row of setae
(Figure 27) . . (on chickens) Menacanthus pallidulus

Abdominal tergites III-VII each with two transverse rows of setae
(Figure 28) . . . 5

Dorsal portion of meso and metathorax with numerous short setae
(Figure 29) . (the chicken body louse) Menacanthus stramineus

Dorsal portion of meso and metathorax with a few short setae on the
lateral margins (Figure 30) .... (on chickens) Menacanthus comutus

KEY TO THE FAMILIES OF THE SUBORDER ISCHNOCERA

Tarsi each with two claws (Figure 31) ......... Philopteridae 7

Tarsi each with one claw (Figure 32) . Trichodectidae 13

Family Philopteridae

Temporal lobes angulate (Figures 33, 34) . . 8

Temporal lobes rounded (Figure 35) . . 11

Temporal lobes extending backward and terminating in long stylet-like
processes (Figure 36) . . (the large turkey louse) Chelopistes meleagridis

Temporal lobes not produced into stylet-like processes . 9

Lateral margins of temporal lobes parallel; two long setae at posterior

margin of each lobe (Figure 33) . . . . . .

. (the fluff louse [on chickens]) Goniocotes gallinae

Lateral margins of temporal lobes distinctly angulate, not parallel; two
or three long setae at posterior margin of each lobe, a short spine at
angular process (Figure 34) . . . . 10

10(9*) Two long setae at posterior margin of each temporal lobe .

. (the brown chicken louse) Goniodes dissimilis

10* Three long setae at posterior margin of each temporal lobe (Figure 34)
. (the large chicken louse) Goniodes gigas

11(7 ) Anterior portion of head with a series of backward projecting chitinized
processes (Figure 37) . . (the slender turkey louse) Oxylipeurus poly trapezius

11 Anterior portion of head without such chitinized processes .... 12

12(11 ) Dorsal posterior margin of pterothorax with four groups of long setae
(Figure 38) .... (the chicken head louse) Cuclotogaster heterographus

12 Dorsal posterior margin of pterothorax without groups of setae; pos¬

terior lateral margins of pterothorax each with a patch of long setae
(Figure 39) . (the wing louse [on chickens]) Lipeurus caponis

6(l')

6'

7(6)

7'

8(7)

8'

9(8')

9'

Family Trichodectidae

1 3(6 f) Lateral margins of preantennal region straight, strongly convergent;

apex of head with distinct narrow emargination (Figure 40) ......

. (the cat louse) Felicola subrostrata

13' Lateral margins of preantennal region rounded, arched; apex of head

blunt, broadly emarginate or rounded (Figure 41) .......... 14

LICE

149

14(13')

14'

15(14)

15'

16(15)

16'

17(16')

17'

18(17)

18'

B. equi and B. ovis are closely related species, their separation is quite diffi¬
cult, generally the characters mentioned in the key will be useful.

19(17') Anterior margin of head slightly flattened or concave in female (Fig¬
ure 47), not so in the male; second abdominal tergite in male with a

concave posterior border (Figure 48) . . .

. . (the angora goat biting louse) Bovicola limbata

19' Anterior margin of head slightly flattened or concave in male (Figure 47),

not so in female; second abdominal tergite in male with a straight pos¬
terior border (Figure 49) .... . (the goat biting louse) Bovicola caprae

B. limbata and B. caprae have been considered distinct species by some authors;
others consider them the same. B. limbata is more common on angora goats whereas
B. caprae is more common on short-haired goats. The separation of the females
of these species is practically impossible since only minor variation exists. The
separation of the males is an easy matter because the genitalia of each species is
characteristic for that species.

Figure 1. Illustration of diagnostic characters used in this key.

Median transverse bands on abdominal tergites present; antennae nor¬
mal (Figure 42) . . . 15

Median transverse bands on abdominal tergites absent; antennae stout
. . (the dog biting louse) Trichodectes canis

Body sparsely covered with short setae . . . 16

Body densely covered with long setae. . . (on angora goats) Bovicola crassipes

Head narrowing anteriorly, anterior margin sharply rounded or blunt
(Figure 43) . . . (the cattle biting louse) Bovicola bovis

Head having anterior margin broadly rounded or flattened (Figure 41) . . 17

Anterior margin of head rounded, preantennal angle not markedly ex¬
panded; antennae long, basal segment of male antenna robust (Fig¬

ure 44) .................. . ...... 18

Anterior margin of head flattened or slightly concave; preantennal angles
expanded, pointed, antennae same in both sexes (Figure 45) ..... 19

Width of temporal lobes about equal to that of preantennal region
(Figure 44) . . . . (the horse biting louse) Bovicola equi

Width of temporal lobes less than that of preantennal region (Fig¬
ure 46) ............... (the sheep biting louse) Bovicola ovis

150

THE TEXAS JOURNAL OF SCIENCE

6

Figures 2-6. Illustrations of diagnostic characters used in this key.

LICE

151

Figures 7- 10. Illustrations of diagnostic characters used in this key.

152

THE

TEXAS JOURNAL OF SCIENCE

irmii

n i in

14

Figures 11-15. Illustrations of diagnostic characters used in this key.

LICE

153

154

THE TEXAS JOURNAL OF SCIENCE

fmfflf m

27

rriTH inriiTf? r r?n'\

mm

immnn p v? t

ffwnw

Ur u Tprrr t pttu

■ffffiffttf

Figures 2 3- 28. Illustrations of diagnostic characters used in this key.

LICE

155

Figures 29-34. Illustrations of diagnostic characters used in this key.

156

THE TEXAS JOURNAL OF SCIENCE

Figures 35-37. Illustrations of diagnostic characters used in this key.

LICE

157

Figures 38-42. Illustrations of diagnostic characters used in this key.

158

THE TEXAS JOURNAL OF SCIENCE

Figures 43-49. Illustrations of diagnostic characters used in this key.

LICE

159

LITERATURE CITED

Anderson, D. M., 1975 -Common Names of Insects, 1975 Revision. Entomological Society
of America. Spec. Publ. 75-1, 37 pp.

Emerson, K. C., 1956-Mallophaga (chewing lice) occurring on the domestic chicken. J Kansas
Entomol. Soc 29(2) :63.

Ewing, H. E., 1929-/n Charles C. Thomas (Ed.), A Manual of External Parasites. Baltimore,
225 pp.

Ferris, G. F., 1951 -The sucking lice. Memoirs of the Pacific Coast Entomological Society ,
Vol L 320 pp.

La Page, G., 19 62 -Monnig’s Veterinary Helminthology and Entomology. Williams and Wilkins
Co., Baltimore (5th ed.), 567 pp.

- , 1968 -Veterinary Parasitology. Oliver and Boyd, London (2nd ed.), 1182 pp.

Metcalf, C. L., W. P. Flint, and R. L. Metcalf, 195 1 -Destructive and Useful Insects. McGraw-
Hill Book Co., Inc., New York, N.Y., 1071 pp.

Morse, M., 1903-Synopses of North American invertebrates. XIX. The Trichodectidae.Amer.
Nat., 37 (44 1) : 609 „

Scanlon, J. E., 1960-The Anoplura and Mallophaga of the mammals of New York. Wildlife
Disease, No. 5. 121 pp. printed on 3 microcards.

Seguy, E., 1944-Insectes ectoparasites (Mallophages, Anoploures, Siphonap teres). Faune de
France, No. 43. P. Lechevalier, Paris, 684 pp.

Sloss, M. W., 1973 -Veterinary Clinical Parasitology. Iowa State University Press (4th ed.), 250 pp.

Werneck, F. L., 1950 -Os Malofagos de Mamiferos. Parte 2. Edicao do Instituto Oswaldo
Cruz, Rio de Janeiro, 207 pp.

Wiseman, J. S., 195 9 -The genera of Mallophaga of North America north of Mexico with
special reference to Texas species. Ph.D. Thesis, A&M College of Texas, College Station,
Texas, 339 pp.

AN INDEX TO THE GENERA OF HOSTS IN YAMAGUTI S
SYSTEMA HELMINTHUM

by D. W. TUFF and D. G. HUFFMAN

Southwest Texas State University,

San Marcos 78666

INTRODUCTION

The first volume of Yamaguti’s Sy sterna Helminthum was published in 1958
and was immediately hailed by many parasitologists as a monumental first step
toward systematically cataloging the helminths of the world. Unfortunately, none
of the now completed five volumes contains an index to the hosts. Therefore,
Systema Helminthum is most useful when used in conjunction with the host index
of some other reference, such as the Index Catalogue of Medical and Veterinary
Zoology or Helminthological Abstracts. However, both of these references are in
serial form and their use under most laboratory conditions presents many logistical
problems. Consequently, we have compiled this compact host index to compliment
the use of Systema Helminthum in the laboratory.

This host index is divided into five sections corresponding to the five volumes
of Systema Helminthum The sections are arranged according to the usual taxonomic
sequence, rather than in order of the publisehd volume number. Within each sec¬
tion, generic names of hosts are listed alphabetically, followed by page number
in the appropriate volume.

Numerous synonyms and misspellings ofhost genera were detected in Yamaguti’s
text and were corrected in the index, but undoubtedly many more were trans¬
cribed undetected. When synonymies were suspected, a number of modern taxo¬
nomic references were consulted to determine the currently recommended name.
We omitted the parasites of “dog,” “cat,” “man,” etc., because page references
to the well-known fauna of these hosts would be so extensive as to be virtually
useless. The reader may encounter similar difficulties with such large and abundant
genera as Anas, Larus, and Rana.

We wish to emphasize that Yamaguti did not attempt to include complete
host lists in Systema Helminthum. Consequently, the reader is cautioned not to
draw conclusions regarding new host records based on the exclusive use of this
index.

ACKNOWLEDGEMENTS

We wish to thank Interscience Publishers for permission to condense and publish
this information. Southwest Texas State University is to be acknowledged for
Accepted for publication: January 11, 1977.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

162

THE TEXAS JOURNAL OF SCIENCE

financial support of the project and for providing unlimited access to their DEC
System 10 computer which was used for sorting and condensing the material. And
finally, we wish to express appreciation for the encouragement of certain of our
colleagues, without whose moral support this project would have been shelved
long before the many weeks of typing and editing had really begun.

INDEX: SYSTEMA HELMINTHUM

163

MONOGENE A and ASP/DOCOTYLEA (vol IV)

( Ablennes-257 ,268) ( Abramis-11 , 12, 13, 15, 16, 17, 24, 27, 30, 35, 36, 37, 38, 40, A 1 ,44,233,316) ( Acanthlas-144.304)
(Acanthocybium-276) ( Acanthogobius-246 ) ( Aoanthorhinus-304 ) ( Acanthorhodeus-25 , 43 , 91 ,233) ( Ac anthostrac ion-50 )
( Acanthurus-65 ) (Acerina-1 1 , 14, 16, 17,25,28) ( Ac lpenser- 1 5 . 1 33 , 233 , 300 . 30 1 ) ( Acrocheilus - 3 1 )
(Aetobatus-153, 163) ( Agonostomus-240,248) ( Albatrossla- 1 95 ) (Albula-232) ( Alburnoides- 1 90 . 1 98)
(Aiburnus-12, 17,27,29,30,31,34,36,37,38,40,41) (Aleblon-165) ( Alestes-233 ) ( AIloteuthis-20) (Allotis-68,75)
(Alopias-162) (Alosa-212,213,215) (Alytes-288) ( Ambassls-64 ) ( Amblopl ites-62 . 63 ) (Ameiur us- 13,51 ,62,63,83)
( Ammodytes- 1 1 ) ( Amyda-297 . 315 . 31 7 , 3 1 9 ) ( Ancyrocephalus-67 ) ( Angel Ichthys- 128 . 24 1 ) ( Anguilla- 1 1 , 34 , 35 )
(Ani3Qtremus-65.244) ( Anodonta-3 1 9 ) ( Anodontostonia-222) ( An thias- 1 23 . 262 ) ( Anyperodon-50 ) ( Aphredoder us-25 )
( Apl ites-76 ) ( Aplodlnotus-242 , 245 . 265 , 321 ) (Apogon-25) ( Apomotis-53 , 62 , 75 ) ( Archopl ites-68 )
(Archosargus-154,241 ) ( Areliscus-236) (Argulus-18) ( Argyreiosus-192) ( Ar lus-49 . 50 , 92 , 93 ) ( Armochelys-297)
(Arripis-241 ) ( Arvis-71 ) (Aspius-30.44.233.234.316) ( Atheresthes-130, 185) ( Atherina- 1 1 , 49 ) (Auxis-204)

( Bagar ius-60 ) (Bagre-70) ( Bairdiella- 1 04 , 1 87 , 207 ) ( Balistes-65 . 66 , 1 92 ) ( Barbus- 1 1 . 1 3 . 15 , 26 , 28 ,
29,34,35,36,37,39,43,233,234,315,316) ( Bathystoma- 1 82 ) ( Batrachocottus-1 1 ) (Belone-255 , 257) (Blepsias-14)
( 31 icca- 16, 26. 27. 30, 34. 35. 36, 37. 40, 44. 233, 31 6) (Bopyrus-179) ( Boreogadus-1 1 ) (Bothus-88) (Box-100, 179,242,244)
(Brachirus-70) (Brachygenys-182) ( Brachymystax- 14,86) (Brama-131 ,202,280) ( Br anchiostegus-24 1 )
(Brevoortia-214,218,220,221 ) ( Bufo-288 , 29 1 )

(Caesio-66 ) (Calamus-131 .244,316,320) { Cal igus- 1 64 ) ( Ca llichrous-74 ) ( Ca 1 lorhynchus- 1 70 . 17 1 , 305 )
(Campostoma-34,35) (Cantharus-134,247,248,320) ( Capoetobrama-26 ) ( Caranx- 1 39 , 227 , 23 1 , 249 , 252 , 263 , 269 ,
271 ,272,274,277,284) ( Carasslus-1 1 ,13,15,25,26,27,28,31,35,36,233) ( Carcharias- 1 4 1 , 144,162,305,306)
(Carcharinus-141 , 307) ( Caretta-296 ) (Caspialosa-212) (Catla-12,26,45) ( Catonotus-76 ) (Catostomus-16 ,
72,91,198) (Cau?olatilus-132, 182) ( Centrarchus-84 ) (Centronotus-242) (Centropomus-103, 165) ( Centropr istes-24 1 )
(Cepola-241 ) ( Cestracion-306 ) ( Chaenobrytt us-25 ,53,54,61,68,75,83) ( Chaenogobius-66 ) ( Chaetod i pter us- 6 1 ,128)
( Chaetodon-64 , 244 ) ( Chae topt er us-79 ) ( Chalc alburn us-26 , 44 ) ( Che il ichthys- 199) (Che lid on Ichthys- 135)
(Chelodina-296) (Chelydra-295.297,319) ( Chilogobio- 1 3 . 30 , 39 , 40 , 49 ) (Chimaera-158, 170,322)
(Chirocentr us-229, 267) (Chironemus-131 ) (Chi or oscombr us-275) ( Choerops- 1 83 ) ( Chondro stoma-26 ,30,40,50,233)
(Chorlnemus-191 ,201,250) ( Chrysemys-295 . 296 ) ( Chr ysophr ys-96 , 1 32 , 1 82 , 24 1 , 277 , 320 ) ( Cir rhina- 12 , 26 , 233 )
( Citula- 1 92 ) (Clarias-21 ) (Clemmys-295.296.297) ( Clinostomus-36 ) ( Clupea-14 , 18, 21 2. 21 3) ( Clupeonella-212)
( Cl upisoma-80 ) ( Cobit is- 11 , 12 , 1 3 , 1 4 , 15 . 16 . 50 ) ( Coelorhynchus- 1 77 ) ( Comephor us- 1 2 ) (Conger-129) ( Corbicul a-3 17 )
(Coregonus-14,86,107.190) ( Corv ina-92 . 96 , 97 ) ( Cor yphaena-1 18. 123.276) ( Cottocomephor us- 1 2)
(Cottus-1 1 , 13,14,134,137.233) (Couesius-12,29, 35) (Cratinus-129) (Crenilabrus-132) ( Ctenobrycon-234 )
( Ctenophar yngodon-26 .39.51) (Culter-36,37.39.40,41 ,233) ( Curlmatus-324 ) ( Cybium-1 19 .21 4 , 280, 282 . 283)
( Cyclanorbls-3 1 9) ( Cyclopterus-1 1.12) ( Cymothoa-179 . 182) (Cynoglossus-96,237) (Cy nose ion -15, 96, 185,247)
(Cyprinus-11, 12. 13 . 15.24 .25.26 .27 .28. 30. 31 . 34 . 36 . 37 . 40 . 41 .49.91 .233.315.316) ( Cypsel ur us-255 , 256 )

(Daicocus-76 ) (Dasyatis-126, 141 . 152 . 153 . 154 . 157 . 160 . 163 . 303 ) (Dasybatis-153) (Diodon-116)
(Diplophysa-15, 16,91) (Diptychus-26 . 30) ( Discoglossus-288 ) (Ditrema-73 , 24 1 ,245) (Donax-316)
(Dorosoma-217,220,221 .222) (Drepane-49,51) ( Dussumler ia-2 1 3 )

(Echeneis-139) (Echinorhinus-305) (Elagatis-280.281 ) (Eleglnus-1 1,13) (Eleutheronema-97 )
(Eloplchthys-34.38) (Emys-296) (Enedrias-242) (Enophrys-14) (Epinephelus-80,96,97, 103, 123, 124, 127,
132,135,137,243,244,245) (Ericymba-36.43) (Erythrocul ter-34 . 35. 36 . 37 . 38 . 39 . 40 . 41 . 42 .50 . 66 . 233) (Esox-11 .
19, 17,83, 107,233,255) (Etmopterus-306) (Eucalia-13,27) (Eucinostomus-102) (Eugaleus-305)
(Eupomotis-54, 56, 61 .63.68.75.76.83) (Eurycea-310) (Euthynnus-1 18.204.205.260) ( Eutropi ichthys-69 , 74 , 80 )
(Exocoetus-255 ) (Exoglossum-42)

(Formio-268) (Fundulus-1 3 , 16 , 17 , 83 , 84 )

(Gadopsis-59 ) (Gad us- 12. 15. 164. 173. 174. 175. 177. 178.213.220) (Galeichthys-66) (Galeus-305) (Gambusia-83 )
(Gasterosteus-1 1.12.15.16) (Geomyda-295 , 296 ) (Germo-1 19) (Gerres-50 , 242) (Ginglymostoma-305 ) (Girella-98. 128)
(Glossogobius-27 .28) (Gnathopogon-27.36.37.44.51 .233) (Gobio-1 3 , 15 . 30 . 36 . 38 . 233) (Gobius- 1 1 , 15 . 28)
(Gonialosa-213.221 .222) (Gonlistius-1 35) (Goniobasis-321 ) (Graptemys-295 , 31 9) (Gymnelis-14)
(Gymnocorymb us-234 ) (Gymnosarda-1 18)

(Hadropterus-83 ) (Haemulon-93 , 182. 184,262) (Halichelys-296) (Hapalogenys-262) ( Har podon-49 ) (Haustor-83)
(Hel lastes-1 3 1 ) (Hel icolenus- 1 34 , 1 37 . 245 ) (Helioperca-52,61 ,75) (Helotes-242) (Hemibarbus- 1 3 , 38 , 42 , 43 , 50 , 51 )
(Hemichromis-6 1 ) (Hemicul ter-34 . 35 . 36 . 37 . 39 . 40 . 41 . 42. 43 . 233 ) (Hemiramphus-82, 196, 197,255,256)
(Heptranchias-305 ) (Hexagrammos-251 ) (Hexanchus-305 ) (Hiatula-242) (Hippoglossina-126)
(Hippoglossus-121 .126.164) (Hippopotamus-294 ) (Histiophorus-1 18,119) (Holacanthus-240) (Hoplognathus- 1 27 )
(Hucho-86 ) (Hur o-5 3 ,54, 68, 91 ) (Hybognathus-28.29.35) (Hybopsis-31 , 34) (Hyborhynchus-25 , 30 . 42 ) (Hydrocyon-38)
(Hy la-288, 289. 292) (Hypentellum-34.72.91 ) (Hypophthalmlchth-30. 38 . 39 ) (Hypopr ion-307 )

(Ictalur us-62, 63) (Ilisha-94) ( Indonaia-321 ) (Iniistius-183)

164

THE TEXAS JOURNAL OF SCIENCE

(Julls-97)

MONOGENEA and ASPIDOCOTYLEA , cont.

(Kachuga-295) (Katsuwonus-118,204.259,260) (Kinosternon-297 . 298 )

(Labeo-12,25,26,28,35,45,316) (Labrax-96, 182,22 4.242) (Labrus-165.240) (Lactar ius-201 ) (Lactophr ys-50)
(Lagodon-79 ) (Lamellidens-321 ) (Lampris-204 ) (Lampsil is-321 ) (Lateolabrax-33 , 87 , 245 ) (Lates-96)
(Latris-136,138) ( Lebistes- 1 2 ) (Lefua- 1 2 , 1 4 , 15 , 233 ) (Leiocassis-76 ) (Leiognathus-53 , 242) (Leiostomus-1 82 , 262)
(Lepeoph their us- 165) (Lepibema-83 ) (Lepidopus-202 ) (Lepidorhinus-1 49)
(Lepomis-1 1,25,52,53,56,61,62,63,64,68,69,76,83,84) (Lethr inus-50 , 99 . 124 ) (Leucaspis-233) (Leucichthys- 1 90 )
(Leuciscus-1 1,12, 13, 15, 17, 25, 26, 27, 28, 30, 31, 34, 36, 38, 40, 42, 44, 233. 3 15, 31 6) (Leuresthes-269) (Lichia-201 ,242)
(Ligumia-319) (Limanda-12,77) (Llmnoeottus-1 1 ,26) (Liocassis-58 . 59 , 85 ) (Liopsetta-13, 17) (Lissemys-321 )
(Loligo-3,20) (Longirostrum-249) (Lota-11,14,165.233) (Lucioperca-26 . 49 , 316 ) (Lutianus-50.66.81 , 131 )

(Hacrochelys-317) (Hacropodus-85) (Macrurus-178, 194) (Maena-182,247) (Makaira-1 19) (Malacoclemmys-319)
(Mallotus-17, 18) (Margarlscus-62) (Mar gar it if era-317) (Megalaspis-262) (Megalobrama-39,41 ,43,233)
( Megalops-98 ) (Heinertia-179 . 182) (Melanogrammus-15) (Melo-317) (Menticirrhus-104 . 182) (Hentula-315)
(Herlangus-165, 173) (Herluccius-178, 193) (Merlucius-165) ( Micropogon-15 , 104 , 207 , 203 , 316 )
( Micropter us-15,53. 62, 63, 69,76, 91 ) (Misgurnus-12, 14, 15. 16,233) ( Mogurnda-66 ) ( Mola- 1 1 6 , 1 1 7 , 1 1 8) (Molva-88)
•(Morone-83, 187.306) (Horrhua-174 ) ( Moxostoma-37 , 44 ,72 ,91 ) (Mugil-49, 52. 61 ,67, 123.243,244,250) (Muraena-149)
(Muraenesox-49,73,99) (Mustelus- 136, 144, 158,303,304,305,306,322,323) (Mycteroperca-128)
(Myliobatis-130, 151,166) (Mylocheilus-29) (Myoxocephalus-12, 14) ( Mystus-30 . 60 , 73 )

(Narcine-56) (Naucrates-124) (Necturus-310) (Nemaehilus-1 1 .12,14,15.16,19.29.91,233) (Nematalosa-221 )
(Neomaenis-184,249) (Neothunnus-1 17 ) (Nephrops-324 ) (Notemigonus-25,29, 198) (Notopterus’-58 )
(Notothenia-111,130) ( Notropl s-25 , 26 , 29 , 34 , 35 , 36 . 37 , 38 , 41 , 42 . 44 , 198 )

(Ocadia-295 , 296 ) (Odax-243) (Qligoplites-201 ,262) (0ncorhynchus-18, 107, 131 ) (Onos-15)
(Onychodactylus-298) (Ophiocephalus- 1 5 , 18) (Ophlodon- 1 65 , 245 ) (Opisthocentrus- 1 4 ) (Opisthonema-221 )
(Opsariichthys-39,41 ) (Orioleueiscus-27) (Orthoprist is- 182, 209) (Osteogeneosus-50,93) (Otolithes-97 , 199)
(Otr ynter-243 ) (Oxygaster-234 )

(Pagellus-131 , 182,241 ,242,243) (Pagrosomus- 100, 123, 132, 182,243,247) (Pagrus-96, 100, 166, 182) (Pama-243)
(Pangasius-69 ) (Parabramis-29, 32,41 .233) (Paralabrax-96) (Paraleucogobio-43) (Paralichthys-77, 126, 185)
(Paranthias-199) (Parapercis-180,241 ) (Parapetalus-1 17) ( Par a si lur us-57, 58, 59) (Parastromateus-261 ,268)
(Parathunnus-1 18,204,205) (Paratractus-271 ) ( Parexoglossum-42) ( Par upene us-66 , 67 ) (Pastinachus-152,153.163)
(Pelamys-1 1 9 , 204 ) (Pelecus-43 , 233) (Pellona-182) (Pelobates-288) (Pelteobagr us-78) (Pentapodus-243)
(Peprilus-243) ( Perea- 1 1 ,14,31,43,83.233.316) (Percina-83) ( Percottus-50 ) (Peristedion-223) (Phenacobius-41 )
(Pholis-16) (Phoxinus-1 1,1 3, 14, 15. 16. 17. 24, 34. 35, 36, 38, 39. 4 1,233) (Phycis-174, 178) ( Plagiognathops-42 . 43 . 44 )
(Platycephalus-49.50.51 ,52.241 ) (Plecoglossus-14 ) (Plectorhynchus-1 32) (Plectroplites-100)
(Pleuronectes-1 1.13,15.16.17.121) (Pneumatophorus-219) ( Pogonias-246 ) (Poll ac hi us -16, 174) (Pollasina-16)
(Polydactylus-97) ( Pol ynemus-24 3 , 249 ) (Polyodon-300 , 31 9 ) (Polypterus-21 ) (Pomacanthus-128,244)
( Pomad as y s-51 , 132,242) ( Pomatomus-24 1 ,244) ( Pomolobus-220 ) ( Pomox is-62 , 63 ) ( Poronotus-244 )
(Priacanthus-51 .244) (Prionotus-183) (Pristiophorus-159) (Pristipoina-278) (Pristis-155) (Pristiurus-145)
(Promicrops-96 ) (Prosopium-1 90 ) (Pseudaspius-29) (Pseudemys-295,296) (Pseudobagrus-58,59,85)
( Pseudogob io-1 3 ,37,38.39.42) ( Pseudorasbora- 13.41,43,51) (Pseudorhombus-51 ) (Pseudosciaena-243)
(Pseudupeneus-66) ( Ptychocheilus-26 . 29 . 31 ) ( Pungi t ius- 1 1 , 1 2 ) ( Puntius-26 . 28 . 29 . 30 , 31 . 38) (Pygosteus-16)

(Rachycentron-139) (Raimas-25) (Raja-55,110,111 ,112,113,144,158,159,161,162,163,307,308,309,324)
(R ana-2 88, 289, 292, 293) (Remora-139) (Rhacophorus-289 ) (Rhatndia-62,63) (Rhina-158) ( Rhinichthys-16 , 29 , 30, 35)
(Rhinobatus-1 53. 156,309.323) (Rhlnoptera-125, 323) ( Rhodeus-34 , 233 ) (Rhombus-137) (Rhynchobatus-155)
(Rhynchobedella-49,69) (Richardson ius- 12, 16,30,35) (Rita-30) (Roccus-85, 96, 193,243,280) (Rohtee-36)
(Rostrogobio- 13,43) (Rutilus- 16,17,25,30,31,35,38,40,42,43,233,316)

(Salmo-16, 18, 107, 190) (Salvelinus-86. 107) (Sarcocheil ichthy- 1 3.40) (Sarda-1 17,204,205) (Sargus-96,
99,100,195,244) (Saurogobio-37 ■ 38 , 42) (Sawara-280) (Scaphiopus-291 ,294) (Scard inius-27 , 30 , 36 , 37 , 233 , 31 6)
( Scatophagus-7 1 ) (Schizo thorax- 1 5 , 1 7 , 26 , 28 . 29 , 32 , 233 ) (Sciaena-90 , 94 , 97 , 99 , 122 , 165 , 229 , 244 )
(Scoliodon-141 ,323) (Scomber-33,214,216,219,274.282) ( Scomberomor us-244 , 277 , 280 , 283 ) (Scombrops-242)
(Scorpaena-1 32) (Scorpis-262) (Scyllium- 1 45 , 303 ) (Scymnodon-148) (Scymnorhinus-305) (Scymnus-304 )
(Sebastes-241 .245) (Sebastichthys-245 ) ( Sebastiscus-245 , 257 ) (Sebastodes- 1 23 , 126 , 1 30 , 137,241,245) (Selene-192)
(Semicossyphus-183) (Semotllus-25,26 , 28 , 29 . 31 . 35 , 42, 62) (Seriol a- 123, 204. 242, 255. 261 ,263,264) (Serranus-97 )
(Setipinna-216) (Siebenrockiella-318) ( Siganus-78 , 79 , 80 , 240 , 24 3 , 245 ) ( Si llaginoides-243 , 247 )
(Sillaglnopsis-97) (Sillago-97) (Sllondia-29.80) (Si lur us-30. 57. 58, 59, 316) (Siniperca-66) (Smaris-247)
(Solea-126) (Somniosus-304 , 306 ) (Sparus-53 . 54 . 72 , 99 , 100 , 132 , 1 47 , 246 ) (Spheroides-126, 128, 164, 184, 199)
(Sphyraena-196 ,275) (Sphyrna- 120, 142, 162,305,306) (Spinax-306) (Squalalburnus-35)
(Squaliobarbus-26,35.37.39.43) (Squalius-37,50) (Squalus-56 . 1 1 9, 144 , 153 , 158 , 304 , 306 ) (Squatina- 1 38 , 149)
(Stenotomus-245 ) ( St er no ther us-297 ) ( St izos ted ion-82) (Stromateus-266) (Strongylura-50 , 51 ) (Synagris-99, 123)

(Tandanus-59) (Tautoga-242 ) (Temnodon-245 ) (Terrapene-297 ) (Testudo-296 ) (Tetranarce-55 , 56 )
(Tetraodon-81 ,117.184) (Tetrapterus-118, 119) (Tetrathyra-319) (Tetrodon-178) (Teuthis-50 , 51 ,246)

INDEX: SYSTEMA HELMINTHUM

165

MONOGENEA and ASPIDOCOTYLEA , cont.

(Thalassoehelys-317) (Theragra-15) (Therapon-64 , 65 , 97 , 101 ,105) (Thrlssocles-216,275) (Thywallus-107, 168, 190)
(Thynnus-1 17, 118, 119, 120,204,205,215,282) (Thyrsltes- 165,202) (Thysanophrys-51 ) (Tllapla-61)

(Tinea-1 1,15,17,25,27,30,39,40) (Torpedo-55,56,305,306) (Trachlnotus-193.285,316) (Trachinus-241 ,246)

(Trachurops-182,241 ) (Traehurus-179 ) (Treblus-166 ) (Triacanthus-51 ,59) (Trichiurus-225) (Trlgla-134 ,
135,1611,165,22*1,256) (Trlonyx-295,297,317,319) (Tr istrawella-61 ) (Trygon-153. 156, 163) (Trygonorrhlna-158)
(Tylosur us-51 ,243,255,257,258,270)

(Umbra-14) (Umbrlna-12 , 92 , 96 , 97 , 104,122,243) (Unio-315) (Upeneoides-67) (Upeneus-66,67) (Uranoscopus-108)
(UroIophus-154 ) (Urophycis-178)

(Varlcorhlnus-15,29,31) (Vimba-16 , 17 , 30 , 44 , 31 6) (Vlvipara-315, 321 )

(Wallago-84 ) (Wallagonla-69,72,74,81 )

(Xenentodon-69) (Xenoeypr ls-39 , 42 , 43 , 44 ) (Xenopus-19.290) (Xenoti3-52.62.75) (Xiphias-1 18, 120, 121 )

(Xurel-227,272)

( Zoarees- 15,16)

D! GENE A (vol 0

(Abramldopsis-65) ( Abr amls-8 , 65 , 65 , 86 , 87 , 88 , 10 1 , 570 , 58 1 , 582 , 588 , 59 1 ,702,703,855,863,869,882,942)
(Abudefduf-57, 157,205,269) ( Acanthias-90) ( Acanthls-616) ( Acanthocepola-312) ( Acanthocottus-121 ,
145,263,271,708) ( Acanthocyblum-252 , 327 ) ( Acanthogobio-722 ) ( Acanthogobius-1 8, 20 , 43 , 52, 1 17 , 160 ,212,

297.712.867.873.877.880.885) ( Acanthosaura-475 ) ( Acanthur us-145, 146, 151 , 178,205,292,298) ( Acartia-264 ,
271 ,296) ( Acciplter-563 , 565 , 567 , 573 ,585 , 586 ,587 ,615,694,755) ( Acer lna-86, 88. 117. 128. 160, 195,268,570,591 ,
703,869) ( Acestrorhamphus-17 , 48 ,285) (Acheilognathus-180,224,230,599.644,687,706,707,716,872,873,882,883,898)
( Aohlrus-1 45 ) ( Acilius-391 ) ( Aclnonyx-950) ( Acipenser-65, 129, 132, 209 , 210,21 1 , 304 , 305, 588) (Acrldotheres-665)
(Aerocephalus-616,682) (Acropoma-57.293) ( Ac tltis-6 16, 671 ,681 ,682,727,734,764) ( Adenota-954 , 957 )
(Aegialltis-732,793) ( Aegypius-563 ) ( Aepyceros-957 ) ( Aequidens-101 ) ( Aeschna-391 . 392 , 393) ( Aetobatis-83 . 84 )
( Agama-401 , 479) (Agamla-588,589,590) ( Agelaius-614,616) ( Agkistrodon-443 . 444 , 448 , 449 , 450, 452)
( Agnostomus-43 , 676 . 677 , 909 ) ( Agriolimax-829,912) (Agrlon-380,392) (Aldemosyne-633) ( Ailurus-926)
(Alx-569.792.793.923) (Ajaia-563,659) (Akodon-836) (Alaeops-53) (Alauda-681 ,683,761 ,763) (Albula-56)
(Alburnus-8, 64, 86, 87. 101 .366.570.859) (Alca-569. 570. 571 ,648.649,664,694,708,874,899)
(Alcedo-596.597.602.661 .693) ( Aleelaphus-958) (Alces-844) ( Alectorls-755.763.764) ( Alepidosaurus-190)
(Aleposomus-309) (Alligator-482,489.536.543.546) (Allotis-198) (Alopex-886) ( Alopochen-6 1 2)
(Alosa-166.263.296.296.301) ( Alphestes-142) ( Alsophis-449) (Aluco-585) (Alutera-133,213,318)
( Alytes-397 . 402,41 1.451 .485) ( Ambassis-722 , 868 . 877 , 878 . 880) ( Amberiza-607 ) ( Amblopl ites-9 ,
12,131 ,132,195,243,246,483,589,595,622,638,735) (Amblyrhynchus-498,499,501 .502) ( Amblystoma-86 , 243 , 376 ,
379,380,381,382,399,402,409,413,478,485) ( Amelurus-86 , 87 . 88 . 1 1 2. 131 . 152. 1 83 . 195 . 210, 21 8, 221 . 243 . 245 , 247 . 483 ,
582,589,638,890,929) (Amerla-429,776) (Amerianna-626,637,892) (Amla- 167, 183, 195, 197,243,483,589,870)
(Amlatus-195) (Ammodytes-1 45, 166,214.263.264,305) ( Ammomanes-789 ) ( Ammospi za-681 ,729,737,757)
(Amnlcola-I 13,138, 195.223.397.485.644,666.693.703.727.735.856.860.870) (Amphacanthus-345,868) (Amphithoe-120,
166,729) (Amphluma-376.396.413.459) ( Amphotlstius-84 ) (Ampullarla-893.932.938) (Amyda-424,436,455,
481,509,529,534) (Anabas-707.873.877.880) ( Anampses-69 ) (Anarrhlchas-27, 51 ,52,55,58. 114, 117,270)
(Anas-567. 569, 570, 570. 57 1.572. 599. 600. 609. 61 0.61 1.61 2. 61 3. 61 4. 6 16. 626 .627. 629. 629. 63 1.632 .633. 634. 637. 638. 645.
648,660,661 ,663,664,665,666,667,669,679,681 ,692,693,694,695,708,720,722,727,732,733,768,770,774,781 , 783, 783,

784.786.791.792.793.795.797.874.885) ( Anaspides-159) (Anastomus-629.642.685) ( Anatopynla-615) (Anax-667 .732)
(Anchovia-266) (Anchoviel la-266) ( Anelstrodon-431 .450.558) ( Ancylochellis-641 ) ( Ancylopsetta-1 69 ,290,308)
(Angelichthys-149.319.341 ) (Anguilla-8.72. 108, 109, 117. 118, 120,125,130,131,159,166,175,195,210,
218,232,241,263,271,281,282,286,304,306,308) ( Anguispira-754 , 91 1 ) (Anhlnga-572,575,599,600,608,623,638,
646,649,692) ( Anlsogammarus-806 ) ( Anlsotremus-62 , 69 . 107 . 108 , 120 . 273 . 299 ) ( Anlsus-375 , 893 , 959 ) (Anobol la-101 )
(Anodonta-8,9) (Anolls-420.425.518.838.916) (Anopheles-392.824) (Anous-606) (Anser-569,
571,600,627,634,667,708,71 1,720,773,778,780,783,784,792,793,794,795,797,874) ( Anseranas-783,784)
(Anslotremus-212) ( Anthura-727) (Anthus-616.681 ,759,763) (Antigone-627,637,645,692,775) (Aoria-9, 10)
(Apeltes-120. 165,296) ( Aphanius-723 ) ( Aphareus-225) ( Aphredoder us-88 . 132) (Aplnephel us-208) ( Aplltes-12,595)
(Aplodlnot us-87 .113.138) (Aplysla-145) ( Apodemus-832 . 908 ) (Apodlchthys-120) (Apogon-42, 43. 167.265)
( Apogoniehthys-1 17.166) (Apomotls-87. 198.595) (Apus-615.753.755.763) (Aqulla-563.567.587.653,692,695,710,763)
(Aramldes-629.734.760) (Aramus-639.659) (Arbacia-51) ( Archlbuteo-591 .695.882) ( Archlplanus-761 )
( Archoplltes-1 16) (Archosargus-1 14, 145) (Ardea-79 , 565 , 566 , 567 ,572,578,
582,588,589,591,609,618,622,629,632,637,638,644,645,646,654,665,687,688,689,690,692,693,703,704,705, 706, 708,
711,727,773,775,781,799,870,871,872,874,882) (Ardeola-566 , 588 ,635,644, 667 , 687 , 690) (Ardetta-656,704,705,720)
(Arelise us-105. 150.213.264.293) (Arenar la-606, 631 .651 ,672.674.727.728.729.730.732) ( Arenicola-651 )
(Argentina-28.213.272.284.294) (Argia-383) (Arion-908) ( Arius-12 , 1 8 , 157 , 257 , 259 ,292, 307 > ( Armadillidlum-757)
(Arnoglossus-13.307) ( Aromoehelys-458 ) (Arquatella-774) (Arripis-13,277) ( Artamus-742,743)

166

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DIGENEA, cont.

(ArtedieUus-21 4,26*1)
(Asellus-117, 39 1,727, 735)
(Aspidophoroides-63 ,109)
(Astacus-8 19,93*1)

( Ateleopus - 1 7 0 )

(Artibeus-811 ,831 ) (Arvlcola-805.896.902.923.926)

( Asio-563 ,585,599,703,727,869, 882,950) ( Aspicottus-273 )

( As pi us- 8 ,65,87.89,588.591 . 9*12) (Aspro-129, 591 .703.869)

( Astatotilapia-548,877,884)

( Ateles-833 )

( Ascalobotes-475 )
( Aspidonectes-481 )
(Assiminea-932,933)
( Astraea-1 07 ) ( Astur-586 , 587 , 692 ) ( Astyanax- 17, 34 , 228)

(Atelodactinls-758) (Athene-563,586,661,670)

(Atherina-1 4, 56, 119, 120,221 ,222,266,277,343.648,708,868,874)
(Auchenoglanls-366) ( Aulopus-1 70 ) ( Austr alorbis-938 )
(Axlnurus-687,689) (Aythya-739) ( Azuma- 1 04 , 11 2)

(Atherinopsis-64,879) ( AttUa-752 ,760)

(Auxis-1 1 ,42,325,330,331 , 3*10 ) (Avis-783)

(Bagarius-21 ,78,79) (Bagrus-44.88.233.235.246) ( Balrdlella-43 . 117, 138. 161 ) ( Balaena-848)
(Balaenoptera-848,925) ( Balantiopteryx-823 , 824 ) ( Balanus-732 ) (Balistes-36,91 ,108,111,124,148,151,158,188)

( Barbus-60, 65, 87, 89, 101 , 102, 129,260,355,588,591 ,688,705,855,867,869,872,878,883,897,942) ( Bascanlon-443 )
(Baslllchthys-58,591 ,597) ( Basil lscus-427 ) ( Batagur-557 ) (Bathygoblus-160) ( Bathystoma-70) (Batlllaria-607)
(Baza-585) ( Bel lator- 1 42 , 1 69 ) ( Belone- 1 5 , 1 9 , 20 , 21 , 43 , 56 . 87 , 292 , 297 . 71 2) ( Belonopter us-742 ) ( Bernbic ium-673 )
( Beroe-32 1 ) ( Beryx-20 , 266 ) (Bibos-972) ( Biomphalar ia-394 , 938 , 955 )

( Bithynia-64, 65, 160, 175,388,392,571 ,599.600,614,618,626,654,661 ,662,694,792,856,891 ) (Bittium-210,232,730)
(Blarina-908.909.910,912) (Blastocerus-955) (Blennicottus-120, 120, 121 ) ( Blennius-29, 37 , 51 , 53 , 62 , 103 , 145)
(Blepsias-292) (Blicca-8, 65, 66, 87, 101 ,702,703,855,856,869,882,942) (Boa-419,457) (Bodianus-145)

( Boleophthalmus-868 ,878,885, 885) ( Boleosoma-87 , 10 1 ,130,131,195,226,589) ( Bombinator-374 , 379 , 382 , 41 0)

( Bombycilla-681 ) ( Bonasa-666 . 676 , 682 . 762 . 946 ) ( Boreogadus-264 ) ( Bos-797 , 829 , 835 , 836 ,
841 ,842,936,938,940,954,955,956,957,959,964,968,969,972,973,974,975,979) ( Boselaphus-954 ) ( Botaurus-565 ,
566,567,575,588,589,590,591,627,642,644,646,654,656,665,687,688,703,704,798) ( Bothrocara-27 . 142)'
( Bo t hr ops- 4 38 ,449,450,474,476) ( Bothus-277 , 293) (Box-297 , 3*t7 ) ( Brachiodontes-38 ) ( Brachygenys-53 ,70)
( Br achygobius-305 ) ( Brae hyp tenus-585 , 586 ) ( Brachyrhaphis-28) ( Br adybaena-835 ) ( Br ama-334 , 336)
(Branchiostegus- 117 ,150,171,177,284) (Branta-572, 608, 629, 768, 791 ,794,795,925) ( Brodenia-833 ) (Brotula-39,
225,271,290) (Brycon-88) ( Br yttosus- 1 60 ) ( Bubal is-972 , 974 ) ( Bubalus-940 . 954 , 955 , 956 , 957 , 959 , 960 , 973 . 975 )
(Bubo-563,585,586.587,604,755.799) ( Bubulcus-565 . 566, 589. 6 13, 635, 649, 654, 689, 694, 707, 860, 873, 878)
(Buccinum-1 14,212) (Bucco-627) ( Bucephala-570 , 57 1 , 572 , 609 , 632 , 644 , 665 , 67 1 , 727 , 773 , 79 1 ,795)
(Buffelus-835, 841 ,973,974) ( Bufo-64 , 88 , 374 , 375 , 376 , 380 , 381 . 382, 383 , 388 , 390 , 391 , 393 , 399 , 400 , 401 , 402, 403 , 404 ,
408,409,410,411,412,413,479,480,566,688,707,873,942,946,950) (Bulimnea-445,570) (Bullmulus-758,762)
( Bui imus-65, 390, 391 .599,644.694.791 .857.860.898) ( Bui inus-402, 4 19. 566, 628, 895, 936, 938, 940, 954, 971 )
(Burhinus-564,765) ( Butastur-564 , 567 , 585 ) ( Buteo-563 , 564 , 567 , 574 , 578 , 581 , 585 , 586 , 587 , 59 1 , 601 , 632, 695 ,703 ,
714,726,752,754,759,869,882) (Butorides-565.567.589,644.687.704,705.708.720,874)

( Cacatua-759 ) (Caccabis-755,759,764) (Cacicus-755.757,758,763) (Caesio-151 ,298, 307) (Caiman-468,
4 .2,487,489,536,537,541,543,544,545,546) ( Cair ina-572 , 628, 694 , 695 .780 ,784 ,797 ) (Calamus-33,37,38,47,110,
115,117,124,125,140,145,147,214,265,294,308,368) ( Cal idris-617 , 651 , 673 , 674 , 728,729 ,730,732,777 ,796 )
(Callichrous-44,86, 122) ( Cal 1 ichthys-565 ) ( Call inectes-730 ) ( Call ionymus- 1 3 . 1 8 , 52 , 1 04 , 1 1 7 , 1 1 8 , 270 , 29 1 ,

293.371) (Callorhinus-861 ) ( Callyodon-1 05 . 159 . 201 , 292 , 353) ( Calopter yx-394 , 408) (Calospiza-755)
(Calotes-4 31,472,474,475) ( Calotomus-1 05 ,117) (£ajLurom^s-8311 ) (Cambaroides-932) (Cambarus-129.243,
379,396,483,553,932) ( Camelus-835 , 937 ) (Campeloma-406,602,679,890.929) ( Campostoma-1 18) ( Campyloramphus-760)
( Canachites-682 ) (Cancer-729) (Cancroma-687.702,869) (Canls-565,597,705,809,
835,857,859,863,867,869,871 ,874, 878, 885, 897, 899, 93*1, 943, 945, 946, 947) ( Cantharus-62, 145) (Cantherines-36,

145.151.170.293.364.371) (Capella-570, 607. 626, 627, 644, 647. 652. 727. 729, 732. 742, 765, 773, 774, 776, 787. 788, 893)
(Capltella-727) ( Capoetobrama-8 , 588 ) (Capra-835,841,957,959,972,973) (Capreolus-926) (Caprimulgus-615.
741,743,748,750,762) ( Caranx-9 , 10 . 12 , 1 8 , 1 9 , 20 , 33 . 40 , 42 , 43 , 7 1 , 74 , 80 , 86 , 88 , 154 , 1 57 , 169 , 207 , 21 2 , 21 3 , 21 4 , 21 5 ,
277,278,279,293,297,307,308,338) (Carapus-162) ( Carassiops-579 ) (Carassius-64,65,86,87,224,230,249,
366,579,588,591 ,599,638,687,707,714,716,859,873,877,878,880,882,883,898,942,943) (Carbo-583,586,648,651 ,704)
(Carcinides-727) ( Carcinus-729 ) (Cardium- 19, 65, 142,739) ( Carduel is-681 ) (Caretta-462,465,487,501 ,
506,517,518,530) (Cariama-565 ,752) (Caridina-160,732) (Carinogammarus-120, 166) ( Carphibis-563 , 626 )
( Car piodes-66 ) (Carpodacus-681 ,682) (Casarca-571 ,572,614,615,638,795) (Casmerodius-565,566,622,653)
(Caspialosa-40,263) ( Cassiculus-627 , 629) ( Cassid ix-754 ,757 . 758) (Castor-904.906,968) ( Cataphractus-358 .
362,363) (Cathaica-835) (Catharista-637) ( Cathar tes-564 , 565 . 57 1 . 637 ,705) (Catoptrophorus-671 ,727)
(Catostomus-66 ,67.87,88.118.131,569.579 .589 .703.860.870.890) (Caularchus-1 10, 120) (Caulolatilus-148, 188,
189,308) (Caulopsetta-167 ) (Causus-436) (Cebidichthys-300) (Cebus-820,831 ) ( Celestus-479 ) ( Celeus-755)
( Centrarchus-88 ) (Centriscops-160, 170) ( Centroc ere us-676 ) (Centrolophus-149,280,302) (Centropristis-109, 142)
(Centropus-628,678) ( Centroscymnus- 1 86 , 196) ( Cepaea-829 , 908 ) ( Cephalophol is- 1 57 ) ( Cephalophus-957 )
(Cepo la-13, 40, 277, 284, 289, 29 8) (Cepphus-621 ,708) (Cerastes-485) (Ceratacanthus-125, 142) (Cerataspis-276)
(Ceratophora-474) (Ceratophrys-404) ( Cerchneis-632 , 757 , 760 ) ( Cercibis-629 ) ( Cercocebus-829 , 832 )
(Cercopithecus-829.936.977) (Cerdocyon-831 ) ( Cerithidea-671 ,715,723,740,794,879) ( Cerithium-205 , 34 1 , 342)
( Cer ivoula-834 ) ( Certhia-681 ) ( Cervicapra-974 ) ( Cervus-797 , 830 , 842 , 964 )
(Ceryle-592, 593, 594, 595. 596, 597. 620, 640, 646. 665. 745) ( Chaenobryttus-1 3 1 . 1 38 , 245 , 360 , 687 ) (Chaenogobius-65,
130,160,224,260,286,867,873,882) (Chaetodon-33,42,49,69,75, 114,352) (Chaetoclipterus-1 14) (Chaetogaster-66)
(Chaetopteryx-101 ) ( Chaetur lchthys-37 1 ) ( Chalcalburnus-65 . 588 , 59 1 ) ( Chalcides-479 ) ( Chalcinus- 1 52 , 248 )
( Chal inura- 1 6 1 ) ( Chamaeleo-392 , 40 1 , 470 , 47 1 , 477 , 479 , 838 ) ( Champse-537 , 54 1 , 546 ) ( Channa-707 , 87 3 ) (Chanos-12)
( Chaobor us-6 15) ( Char adr ius-569 , 630 , 68 1 ,727 ,728 ,729 ,733 ,756 ,780 ,782 ) (Charax-101 ) (Chascanopsetta-266 )
(Chasmagnathus-887,932) ( Chatoessus-30 ) ( Chaulelasmus-778 ) ( Chaunax-265 , 289 , 294 )
(Cheilichthys- 134, 188, 319,368) (Cheilinus-167) (Cheilodactylus-289) (Chelidon-615,616,744,761 ,763)
(Chelidonias-580,648,789,899) (Chelidonlchthys-58, 109, 1 17, 118,212,281 ) (Chelidoperca-170)

INDEX: SYSTEMA HELMINTHUM

167

PIGENEA, cont.

(Chelonia-432, 462, 465, 479, 487, 495, 496, 497, 498, 499, 500, 501, 503, 504, 505, 506, 507, 51 1,51 2, 51 3, 51 4, 515, 51 8, 524, 526,
528,552,557,558) (Chelopu3-457.458.459.521 ) (Chelydra-445.457.458.459.463,483,507.514,516,521 ,522,529,
544,553) (Chelys-554) (Chen-791 .792,795) (Chenolopex-628) (Chenopsis-572.780,791 ) (Cherox-806) ( Chettusia-787 )
( Chllomycterus- 113,349) (Chllonyeteris-817) (Chlloscyllium-83,84, 115) (Chlltonia-806) (Chlmaera-196)
(Chlonls-791 ,795) (Chirocentr us-207, 288) (Chlroleptes-41 D (Chironectes-856 ) (Chlronlus-438.440.476)
(Chironomus-391 ,615,806,821 ) (Chlroxiphla-760) (Chltra-424) (Chlamydoselachus-196)
(Chlidonlas-570,580,632.648.688,711) ( Chlor lchthys-109) (Chlorophthalmus-170,266) ( Choerodon- 1 17 . 1 1 8)
(Chondrostoma-102,588,882) (Chorlnemus-17) (Chrionema-13) (Chrolcocephal us-787) (Chromls-601 ,941)
(Chrysemys-243, 446, 457, 458, 459, 460, 483, 507, 521 ,522,557) ( Chryslchthys-687 ) ( Chrysolophus-779 )
(Chrysophrys-12,37,62, 102,121 ,235,280) (Clchlasoma-138)
(Cleon la-2 12, 565. 591 ,602,618,642,644,649.687,703.742,869,882) (Cinclus-738,74 3) (Cinlxys-479) (Clnnyrls-645)
(Clnosternum-544 ) (Clonella-829) (Clpangopaludlna-894) (Clrcaetus-563,587.646,759.867) (Circus-563,
565,566,567,578,585,586,587,591 ,602,644,646,665,670,670,692,695,754) (Cirrhitus-220,227) (Cistudo-459)
( Citellus-806 ) (Cltharlchthys-14.304) ( Clangula-588 , 599 . 658.664 ,727 .739 .778. 783 .791 ,795.796) (Clarlas-157.
172,182,191,245,690,873,878,880) ( Clemmys-457 , 458 , 459 . 459 . 460 . 520 . 555 ) (Cleopatra-601,867,878,941,971)
(Clethrionomys-832) (Cllnocottus-1 10, 157,272) (Cloelia-457 ) (Clupanodon-149.266.277) (Clupea-9.11,
34, 166,254,263,264,266,271,276,280,296,297,298,300,301 ,305) (Clupisoma-172) (Cobltis-8, 65, 102.224,230,
588,694) (Cobus-938,954,957,974,979) (Coceothraustes-676,683,788) ( Cochlearius-565 ) (Coelopeltis-479)
(Coelorhynchus-14.39.142.161 . 170,270.290) (Coenagrlon-614) (Colobus-829) (Coloeus-614,665,676,755,789)
(Cololabis-371 ) (Colossoma-356 , 361 , 363) (Coluber-430,434.438,444,449,450,452,453,457,467,485,488,540)
(Columba-633, 634, 638, 660, 676, 678, 703, 708, 720, 788, 869, 874) (Columblgallina-676) (Colymbus-569 . 570, 57 1 , 573,
578,579,581,632,635,638,644,647,648,649,694,701,702,708,708,717,720,768,786,798,868,869,873,874,883, 884, 899)
(Complecta-703) (Condylura-908) (Conger-17,18,108,166,214,264,281,294,304,307,308) (Connochaetes-938)
(Constrictor-44 4,475) (Contia-478) (Coracias-564) ( Coracina-583 ) ( Coragyps-564 ) (Corbicula-89,893)
(Corcorax-681 ) (Cordulla-393) ( Coregonus-8 . 53 . 86 , 130. 131 . 132.591 ) ( Coreperca-883 ) (Corethra-404 )
( Cor idodax-368 ) ( Corone-757 .761 ,763) (Corvina-200,235,238) (Corvus-564 , 599 . 600 , 61 1,614,627,628,
629,630,645,646,647,649,662,665,666,669,670,675,676,677,681,692,693,696,749,754,755,758,763,787,788, 789, 893)
(Corydalis-406) (Coryphaena-213.252.253.255.276.318) (Coryphaenoides-142.284) (Cosmetira-149)
(Cottocomephorus-130) ( Cottunculus- 1 02 , 103,271,290,370) ( Cottus-17 , 27 , 86 , 87 , 88 , 89 , 102, 109,114,117,118,
119,129,130,131,132,159,212,214.264,270,271,286,297,305,547,601,709,711 ) (Cotur nix-6 15. 669, 676, 681 .703,751 .
754,763,764) (Cotyle-6l5.754.755.762) (Craterocephalus-579) ( Cratlnus-271 ) (Crax-763) (Creagrutus-228)
(Creel sc us-628, 629) (Creisson-880 ) (Crenicichla-102,228) ( Crenllabrus- 17 , 37 ,53, 86 , 1 17 , 307 ) ( Crepidula-202 , 651 )
(Crex-681 ,682,763) (Cricetus-807,937.948) (Crimata-93) (Crlnia-632) (Cristivonier-131 )
(Crocethia-730,733.734,799) (Crocldura-809.830.907) (Crocod 11 us-468. 482, 491 .537.538.543.546) ( Crocothemls-390)
(Crotophaga-586.627.628.630.646.744.758.759) (Cryptacanthodes-214.271 ) (Cryptobranchus-384 , 41 3)
(Cryptoglaux-563) (Cryptotomus-205 ) ( Cr ypturellus-789 ) (Ctenolabrus-108) (Ctenopharyngodon-878)
(Cue ulus-6 14, 6 16, 665, 754) (Culex-615) (Culicoldes-615) ( Cumingla-62, 651 ) ( Cuncuma-799 ) (Cuon-963)
(Curimatus-228) (Cyanocltta-681 ,754.755.761 .762) (Cyanocorax-758,760,789) (Cyblum-20)
(Cyelagras-425.438.446.473) (Cyclas-379.633.651 ) (Cyclemys-458) ( Cyclops-263 . 264 . 287 . 395)
(Cyclopterus-121 .264.286.294) (Cygnus-569 . 57 1 , 609 , 61 1 , 629 . 658 .661 , 664 , 665 , 694 ,791 ,793,794,795)
(Cymatogaster-37.71 ,76) ( Cymblum-9 ) (Cynchramus-616) (Cynopolius-773) (Cynoscion-21 ,118,134,145,
166,200,213,282,310,368) (Cypr inus-64 . 65 . 65 . 87 . 101 . 131 . 175 ■ 249 . 293 . 366 . 586 . 588 .707 .71 4 .71 6 . 855 . 868 . 870. 873 .
877,878,882,883,885,942,943) (Cypselurus-333) (Cypselus-615.753) (Cystignathus-403.404) (Cyttus-118. 170.281 )

(Dacelo-564 .580) (Dacnis-755) (Dactylopagrus-1 17.272.368) (Dactylopter us-280) (Pactylosargus-349)
(Dafila-570, 609, 61 1, 664, 727, 733, 778, 791, 792, 793, 794, 923) ( Daicocus - 1 5 8 ) (Dampleria-148, 168) (Daphlla-734 )
(Daphnia-806) (Dascyllus-292) (Dasyatls-84 ,85,90) (Dasycottus-214,271 .290) (Dasylophus-681 ) (Pasymys-938)
(Pasyprocta-937 ) (Pasypter us-823. 824) (Pasypus-837) ( Pa syur us-903. 908) (Pecapterus-166) (Pecodon-57 ,117,140)
(Pelrochelys-457) (Pel lchon-744 ) (Pelma-475) (Pelphinapterus-850) (Pelphlnus-846 . 848 , 851 . 852, 856 , 859 . 876 , 95 1 )
(Pendroealaptes-677 ) ( Pend rocoelum-64 , 392) (Pendrocopus-616,681 ,763.789) (Pendrocygna-628.637,638)
(Pendrodryas-374) (Pendrolca-681 .684) (Pendronias-681 ) (Peniscus-429 ) (Pentex-235 , 304 )
(Permatemys-464 .515.550.555. 557 ) (Permophis-459) (Peroceras-754. 757.762,909. 910, 91 2) (Pesmana-905 , 945 )
(Pesmognathus-376, 381 ,383,399,413) (Piabrotica-754 ) (Piacope-107,264) (Piagramma-69 . 115,120) (Pibranchus-289 )
(Plcamptodon-383.386.876) (Picerus-97 1 ) (Pichoceros-585) (Plcholophus-565 ) (Plcotyles-968 ) (Picrostonyx-908)
( Pi crur us-6 16. 64 8. 667. 74 3) (Pldelphis-734,805.830,838,856,886,891 ,
901,908,909,933,934,937,944,947,948,949,950) (Plemictylus-86.399.400,405) (Piodon-1 13,316,349,371 )
(Piomedea-705 ,871 ) (Plplacanthopoma-30.289) (Piplectrum-305) (Plplodus-1 17) (Piscoglossus-485 , 856 )
(Piscognathus-869. 883) (Plssoura-660 . 652) (Pistelra-488 ) (Pistichodus-239) ( Pi trema- 104, 159. 167,213)
(Doderlelnia-57 ) (Ponax- 19. 53. 71 .222.739) (Poras-358. 363) (Porcelaphus-797 ) (Porosoma-1 18,297 .701 ) (Praco-476)
(Preissenla-86 , 87 ) (Prepanopsetta-27 ) (Dromas-650 ) ( Dr ymar chon- 450. 450, 47 4) (Prytnobius-438,539)
(Dryobates-665.677 .681 .682) (Pryophylax-438) (Pugong-920,923.979) (Dumetella-779) (Puymaeria-37,297)
(Puymar la-37 ) (Dyromys-81 1 .838) (Pytiscus-388, 391 . 392)

(Eehlnogatmnarus-159) (Echlnorhinus-1 96 ) (Egretta-565 . 566, 582. 588, 591 .636,644.654,687.706,715.716,873,
882,883) (Elagatis-213) (Elanoides-563) (Elaphe-426, 431 .449,479.541 .558.950) (Elaphis-450) (Elaps-446)
( E leg ino ps-63. 74. 270) (Eleotr is-950) (Elephas-841 .844,965.967.971 ,976) (Elestris-269) (Eleutheronema-880 )
(Eliomys-907,908) (Elliptio-12) (Elolsa-41 1 ) (Elosia-381) (Emballonura-821 ) (Emberiza-615,681 ,
748,754,755,767,788,789) (Embiotoca-51 .71 ) (Emerlta-727) (Emoia-476) (Emozamla-673 ) (Empidonomus-762 )
(Emyda-4 36 , 480 ) ( Emydur a-422 , 437 ) ( Emys-243 , 424 . 459 . 460 . 464 . 483 . 516 , 521 , 522 . 523 ) (Enallagma-380 . 383 )

168

THE TEXAS JOURNAL OF SCIENCE

DIGENEA, cont.

(EneheXyopu 3-208 ) (Endophrys-120) (Engraul ls-30 . 1 49 . 264 . 267 . 292 . 297 . 298 ) (Engyophrys-142) (Enhydra-851 .
861,870,886) (Enhydr is-431 .5*17,950) (Enneacanthus-88 .595) (Enophr ys-281 ) (Ensls-651 ) (Eophona-676,754)
(Ephemera- 101 ,131 ,667,821 ) (Ephippiorhynchus-629) (Epicordula-666 ) (Epigonus- 1 42 )
(Eplnephelus- 16. 17. 18.37,89. 107.108. 109. 120. 127, 142. 149. 157. 159. 174.213.214.278.281 .282.294.305.329. 333, 3*12,
867) (Epltheca-379 , 380) (Epomops-837 ) (Epteslcus-805. 806. 807, 81 6. 817.822.823.824.826. 91 5) (Eques-163, 169)
(Equula-69 ) (Equus-937.971 .976) (Eremophila-756) (Eretmochelys-462. 465, 496. 497. 498. 499. 500, 501 ,505,526,530)
(Eriehthoni us-222) (Er lgnathus-855 ) (Erimyzon-66) (Erinaceus-832.897.908.908.936) (Eriocheir-932)
(Erolia-563, 634. 65 1.673. 727. 729. 732. 734. 799) (Erythrolamprus-449) (Erythropus-563)
(Esox-8, 9, 12, 17. 44, 86. 88. 10 1.129. 131. 160. 183. 195. 24 3. 483. 570. 581. 59 1.592. 890. 942) (Eteone-727) (Etheostoma-87)
(Etrumeus-52) (Eucalia-129.578.592) (Eucinostomus-72. 138, 165.265) (Eucynopotamus-270) (Eudryas-474 )
(Eudynamis-586) (Euhadra-400 , 678 ) (Eulota-678) (Eumeces-479.480.950) (Eunectes-425, 426 . 446 , 457 . 473 , 61 5)
(Euomphal la-829. 908) (Euparypha-677 ) (Euphagus-757 ) (Euphractus-937 ) (Eupomacentrus-1 10)
(Eupomotis-12 , 87 , 131 . 1 38 , 195 , 360,589.595 , 622 . 687 . 688 ) (Eurota-908) (Eurycea-382. 399) (Eurynia-9)
(Eurynorhynchus-751 ) (Eurypyga-741 .752) (Eurystomus-694 ,766 , 860) (Eutaenla-447) (Eutamias-806)
(Euthynnus-1 2. 42. 276, 305. 308, 31 4. 325. 326. 327. 328 .334. 335. 339) (Eutropilchthys-20,-21 .89. 106.229,233)
(Euxenura-619.642.646.742) (Exocoetus-57 )

(Falco-563. 564, 573. 585. 586. 587. 591 .615.628.666.670.679.684.692.695.751 ,758,759) (Farancia-396.441 ,445,
457 , 488 ) (Fellchthys-308) (Felis-597. 837. 838. 855. 856. 857. 859. 863. 869. 871 .874.877.878.897.934.940.945.950)
(Fennecus-597,945) (Fiber-793,807,892,895,923,924,925,946,948,969) ( Fistular ia-57 . 143 . 21 3. 304 ) (Florida-566,
704,733) ( Fluminlcola- 1 18, 812, 81 4 ) (Fluta-132) (Fontogammarus-160.658) (Formica-829) (Formicarlus-760,760)
(Fossar la-459. 570. 841 .842.957.968) (Francolinus-763.781 ) (Fratercula-708.729.738) ( Fregata-575 .

601.603.711.868) (Fringilla-6 15. 665. 681 .683.755.763.767.789) (Fulica-570, 591 ,599,612,626,627,628,629,630,
632,634,662,663,665,667,681,682,746,748,770,773,774,781 ,782,788,789,792,795,799) (Fullgula-609.633.660.66l .
662,727,778,780,783,786,902) (Fulix-692) (Fulmarus-769) (Fundulus- 12.70. 131 .138,152,165.184,589,592,
715,723,799,871,879,881) ( Furcimanus-27 , 142) (Furnarius-760)

(Gadus-9, 19,59. 120. 121 , 142 , 165 , 205 , 212, 214 , 263,263 , 264 , 271 , 286 , 290 , 292 , 297 , 304 , 708) (Gagata-79)
(Galax las-159, 160, 24 1,579) (Galba-841 .842) (Galbula-744 ) (Galeichthys-28.241 . 308) (Galeorhinus-256)
(Galictis-565) (Galllcrex-694 ) (Gall inago-569 .574 . 594 . 606 . 629 .727 .773 .774 .782 . 797 ) (Gal 1 inula-569 . 594 ,
627,628,629,630,636,661,677,681,682,683,746,752,773,774,778,780,781 ,792,794,799) (Gallus-564 , 570 , 600 , 616 , 627 ,
633,634,646,665,666,669,670,675,678 681,694,694,708,723,749,779,788,792,793,794,797,874) (Gambusia- 1 84 . 37 1 ,
579,601,638,645,714,723,798,867,878,941,942) (Gammarus-101 ,117,120,159,391,392,412,733)
(Garr ulus-564, 665. 676. 681 ,742,746,755,755,761 ,763,767,788,789) (Garzetta-635,690) (Gasterosteus-1 03 , 121 , 305)
(Gastrodonta-908.909.912.913) (Gastroidea-754 ) (Gasterosteus-1 19) (Gavia-638.647,649.662,697,698,703,708,874)
(Gavialls-469.491.541 ) (Gavllan-583 ) (Gecko-474) (Gelochel idon-673 ) (Gemma-740) (Genetta-950)
(Gen ypter uS-304 , 306) (Geoclemmys-457,458) (Geoemyda-457,458,459.479,516) (Geothelphusa-931 .932)
(Geothlypis-684,743) (Gephyrochar ax- 1 82 , 228 ) (Gerbillus-675,908) (Germo-335) (Geronticus-628.629.647.648)
(Gerres- 138. 190.207.722.868.877.878.880) (Gerris-722. 868 . 877 . 878) (Gibbonsia-1 1 0. 120.170) (Gillichthys-723)
(Ginglymostoma-84) (Giraffa-841 ) (Glrella-1 48. 149. 157.205.292) (Glanidium-1 19) (Glareola-74 1 .743.748)
(Glaucidlum-563.758) (Glauclonetta-570.649.693.708.728.729.735.778.792.795.874) (G1 irulus-824 ) (Glis-908)
(Globlocephalus-848.849.853) (Glossogobius-1 8, 157, 195,225,226,269,277,645,707,720,873,877,878,880,884)
(Glottis-7 7 3. 774. 77 6. 779) (Glyptanisus-960 ) (Glyptocephalus-212) (Glyptosternum-172) (Gnathonemus-689 )
(Gnathopogon-65, 118,224,230,260,707,716,873,883) (Goblo-87, 117.588.860.869) (Gobiomorphus-125, 159,241)
(Goblus-10. 17.53.65.80. 102. 108,109,160,166,167,286,293,304,307,708,709,874) (Gomphus-390 , 666 )
(Goniistius-105. 107. 115. 151 . 157,293) (Goniobasis- 1 18, 121 , 198,376,506,507,749,870,874,888)
(Gorsakius-644.716.883) (Goura-651 ) (Gralllna-564 ) (Graptemys-483.507.516.520.521 .533) (Grayia-491 .558)
(Gr ison-838 ) (Gr us-628, 754. 770. 775. 779. 783 .893) (Guara-704) (Gulra-630.744,752,788) (Gulo-855 . 863)
(Guttera-774) (Gymnacanthus-27 . 106 . 1 2 1 . 142.264) (Gymnarchus-78 . 87 . 235 ) (Gymnodactylus-475) (Gymnosarda-253)
(Gymnothorax-1 1,17. 304. 307. 308. 309) (Gyraulus-375. 406, 409, 582. 61 4. 626, 891 .893.894,897.925)

(Hadropterus-87 ) (Haematopus-627 , 651 .658.728.732.733) (Haemopis-569 . 57 1 ) (Haemulon-33.47.57.62.69,
70,71,107,108,109,162,163,214,273,294,332,601) (Hagedashia-612) (Halcyon-573.595.596,661,754,765.773)
(Hal iaetus-563. 564, 571 .585.586.587.591 .695.705.710.882) (Hal iastur-564 . 587 . 603 ) (Hal ichoeres-104 , 107,110,
160,205,297) (Halichoerus-855.859.863) (Halicore-919.920.922.978) (Halieutichthys-142) ( Haminoea- 6 1 4 )
(Hapalogenys-167) (Haplochromls-178) (Hardella-420.506.525) (Harelda-658 ) (Harengula-149,222,266,

277.298.701.868) (Harpagus-758) (Har pi prion-646,741 ,752) (Harpodon-280,351 ) (Hedimela-742) (Hedymela-762 )
(Hedymeles-757 .761 ) (Hel ice-735 . 887 . 932) (Helicella-676.779.829.908.829) (Hel icigona-908) (Helicodiscus-91 3)
(Hel lcogena-829) (Helicolenus-109 . 165 . 169 . 170 . 277 ■ 281 . 283 ) (Hel icoperca-87 ) (Helicopsis-676)
(Helioperca-1 38. 198.589) (Hel isoma-66 . 183 , 184 , 245 . 263 . 371 . 379 . 395 , 405 , 406 , 458 , 459 , 463,
521,522,553,571,572,592,595,622,638,688,703,784,797,799,892,909,946,979) (Helix-676.681 .908.908,909)
(Helizella-914) (Helodrias-633) (Helodromas-6l6.632.776.794) (Helostoma-358 ) (Hemibarbus-65) (Hemichromis-942)
(Hemiclepsis-89 ) (Hemiculter-880) (Hemidactylus-472 . 474 . 475 ) (Hemigrapsus-727 . 887 ) (Hemipipa-375)
(Hemiramphus-56. 311 .722.868.880) (Hemitripterus-121 ,157.212.214,271,282) (Heosemys-556 )
(Hepsetia-342.722.868.880) (Herod ias-565. 566. 582. 588. 628. 635. 64 8) (Her pailurus-838) (Herpestes-943)
(Herpetodryas-448.551 ) (Her petotheres-564 ) (Herpobdella-64.569.571 .572) (Hesperiphona-754 ) (Heterandria-184)
(He ter odon-4 49. 450. 450. 452. 61 7) (Heterodontus-90) (Heteropneustes-44 , 45 . 164, 172, 191 ,192) (Heteropygia-696 )
(Heterotis-490) (Hexagenia-1 31 . 152) (Hexagrammos-60 . 1 1 5 . 157 . 272 ) (Hexanchus-196. 197) (Hierac idea-563. 646, 692)
(Hleraetus-581 .584) (Hierococcyx-567 )

INDEX: SYSTEM A HELMINTH UM

169

DIGENEA, eonfc.

(Himantopus.-649, 671 ,702,727,729,741,752,773.777,778,781 ,782.784,787,789.794,797,798,869) (Hlodon-1 *1 . 1 31 )
(Hippeutis-897) (Hippocampus-212, 215) (Hippoglosgoides-53,55, 106, 165,212.271 )
(Hlppoglosaua-30.55.212.237.263.270.271.272.290) (Hippopotamus-956, 958. 960. 961 .962.963.97*1) (Hlppotragus-918.
938,958,97*1) (Hirundo-6 15, 61 6, 665. 742, 79*1.766. 768) (Hiatrlonicu3-737.792) (Holacanthus-34 . 360)
(Holconotus-162) (Holoe«intru8-126, 134 .169.170) (Homalonyx-682) (Hoplerythrinua-689) (Hopllas-244,689) .
(Hoplichthya-13) (HopIognathus-28, 29,57 . 1 15, 127 ) (Hucho-195) (Huro-87 . 131 .223 , 366 . 547 . 589 . 601 ) (Huso-211)
(Hyalella-121) (Hyborhynohus-n8.582.589.856) (Hydraspis-549.554) (Hydrobia-65 ,268,694)
(Hydrochelldon-578.607.615.616.632.649.681.768.787.788) (Hydrochoer us-929. 928. 968) (Hydrocoloeus-606.607.613)
(Hydromy s-808.88S,94 9) (Hydroprogne-573 , 578 ) (Hydropsalis-627 ) (Hydroscopus-457 ) (Hyelaphus-963)
(Hyla- 14, 64, 307, 374, 375. 376. 379. 380. 38 1.388. 39 1.392. 399. 400. *101 .403. 408. 4 1 1 . **29. 4*45. 4*19. 454. 478. 479. 548, 565,
571,629,632) (Hylophylu3-759) (Hynnodus-199.265) (Hynoblus-892 . 893) (Hyodon-118) (Hypentel ium-66 . 11 8)
(Hyperoodon-851 ) (Hypleurochilus-1 14) (Hypoclydonla-12) (Hypodyfces-109) (Hypolais-789 ) (Hypomesus-260)
(Hypomor phus-586 ) (Hyporhamphus-34 , 205 , 297 . 371 ) (Hypotaenid la-645, 682. 699, 742)
(Hypotriorchls-563.567.815.616.754.755) (Hypsoblennius-n4)

qbls-600.614.548.848.687.742.765) (Icelua-118) (Ichthyoborus-647) (Ietalurus-1 12. 152. 184. 185.226.
243,245,483,547,601,870) (Ieteria-745) (Icterus-564, 682, 758, 789) (Ictlobua-66. 199.337) (Ictis-946)
(Iduna-615.789) (Idus-1 75. 581 .855.856.882.898.942) (Iguana-465.492) ( Iher lngichthys-1 32. 366 )
(Ilyanassa-51 .727) ( Ilybius-404 ) (Indoplanorbls-895.937.938.973.976) (Inlmlcus- 18. 157.270.281 .304)
(Ionornls-795) (Iridlo-109) ( Ischnura-383 ) (Istlophorus-267 ) (Ixobrychus-566.573) ( Ixoreus-747)

(Jablru-591.742) (Jaeana-639,752.773) ( Jenynsia-591 ) (Johnlus-212.278) (Jordanella-184 ) (Julis-37)
(Junee-780)

(Kachuga-424 , 425 , 426 , 427 , 458 , 459 ,480,501, 506 , 508 , 525) (Kantlmurla-816,824) (Katayawa-932,937)
(Kathetostoma-9 , 10 ) (Katsuwonus-330) (Ketupa-579.714) (Klnlxys-950) (Klnosternon-457.458.459.5l6.529)
(KlrkaIdla-390) (Klttaclnela-588) (Kyphosus- 135. 136.205.250.300.369)

(Labeo-358) (Labeobarbus-60 ) (Labes-337) (Labldesthes-152) (Labrax-9 . 14 . 103 .235 . 304) (Labr isomus- 1 88 )
(Labrus-37 ,51,53.103.108.109.111.117.270.709) (Laecotrephes-390 ) (Lacerta-419.475) (Laehesis-450 . 475)
(Laehnolaemus-213) (Laehnolalmus-32, 109. 141 , 145.213) (Lactophrys-91 . 186,202,203) (Laemonema-106 , 142,214,169)
(Lagoeephalus-1 34 ) (Lagodon-29, 112, 145, 166) (Lagopu3-665.676.68l ) (Lahlllella-93) (Lampr is-295 . 326 , 337)
(Lawpropeltls-447.449.450.451 .452) (Lampsllls-12) (Lanlus-564. 61 5. 627. 665. 676. 681 .742.744.754.759,763)
(Lapemis-485) (Lar us-134. 564. 566. 569, 570. 570. 571 .572.573.578.579.583.591 .599.600.606.607.613.614.615.616.621 .
622,626,632,633,644,647,648,649,650,651,652,658,665,669,670,671,673,687,688,694,696,701 ,702,703,708, 711, 712,
715,716,722,724,727, 728, 729, 732, 733, 738, 755, 756, 768, 787, 868, 869, 871, 874, 877, 884, 885, 899, 904)
( Las Ion ye ter 1 s-91 5. 91 6) (Laslurus-823 , 824 . 830, 915 ) (Lateolabrax-222)
(La ter all us-758, 760) (Later idopsls-1 58. 289) (Laternula-72) (Lates-158.213.231 ) (Laticauda-486 , 49 1 )
(Latllus-179) (Latr idopsls-1 23 .281 ) (Leander-87 . 109. 160) (Lebistes- 1 2 . 1 1 8. 228. 582. 638) (Leclthoconcha-683)
(Leiwadophls-474 ) (Leiognathus-1 54 . 208 . 279 . 297 . 37 1 ) (Leiolopisma-399 . 478 . 479 ) (Leiostomus-7 1 . 76 . 1 38 . 166 . 21 3)
(Lepadogaster-108.289) (Lepibema-227) (Lepidaplois-710) (Leplsosteus-I 2 , 183 . 184 , 37 1 . 870)
(Lepomis-9 .12. 87 .88. 121. 129. 131 , 195,223,589,622,638,687,856) (Lepor inus-48 , 93. 152.270) (Leptagonus-103)
(Lepfcoeella-87) (Leptoeephalus-1 3. 17. 18.54. 120. 142. 157.281.304) (Leptocottus-121 .271 .272)
(Leptodactylus-374 .375.381. 382 .402.403. 404 . 409 . 488 . 565 . 582) (Leptodira-449) (Leptonychotes-925)
(Leptoptllos-618.641 ,694) (Leptoscarus-1 17,159) (Leptosynapta-651 ) (Lepus-835.914.941 ) (Lernaeoeera-271 )
(Lestes-408.614) (Lefchr inus-1 8.53 . 69 . 106. 107. 108 . 1 15 . 1 17 , 118, 120. 121 .124.176) (Leucasplus-600)
(Leuelehthys-131) (Leuclscus-8.9.65.66.86,87. 101 , 175, 176, 195,260,582,588,591 , 702, 855, 856, 859, 869, 882 ,9**2)
(Leueogobio-224 ) (Leuconoe-807.821 ) (Leucophoyx-653 ) (Leucorr hi n la-666 ) (Leucosareia-677 ) (Leuresthes-7 1 1 )
(Llbellula-383. 391 .392.393.395.666) (Lichia-1 1,277,292,297,304,316,705.867,872,883) (Ligurlnus-676)
(Limanda-166.213.271 ) (Umax-681) (Limnodromus-733) (Llmnodynastes-408. 41 1 . 375 . 380. 429) (Llmnopardal is-795 )
(Limnophilus-412.812) (Limnothlypus-788 ) (Limnotragus-974 ) (Limosa-569.574.626.632.633.727.773.782)
(Llmulus-885 ) (Liocassis-88) (Liophis-375.428.438.439.448.472.474.540.551) (Liopsetta-157) (Liparis-17, 121 )
(LisseMys-396.420.424.48l .532) (Llssotr lton-374 ) (Llthofaloo-615) (Littorina-120.606.708.709.728.729.796)
(Liza-867.873) (Lo-351) (Lobivanellus-628.777) (Lobodon-925 ) (Lobotes-20 , 297 )
(Lophlus-1 9. 20. 142.212.213.225.264.271.277.281 .290.304.307.308) (Lophodytes-57 1 . 588 . 703 ) (Lophol ati lus-264 )
(Lophopsetta-1 42. 166.213.708) (Lophura-475) (Lophuromys-938) (Loricaria-122,270) (Loris-820) (Lorlus-763)
(Lota-8.86,88,130.131.132.195) (Lotella-127 ,284 . 308) (Lovettia-266 ) (Loxodon-965 ) (Loxodonta-939 )
(Lueioperea-8, 9, 88. 88, 128, 195.293.591 .703 .869) (Luclopimelodus-270) (Lumbriculus-391 ) (Lumpenus-62 , 121 )
(Luseinia-681 .699.759.763) (Lutlanus-107 . 112, 157,213,225,227,278,280)
(Lutra-705 .807.863. 872 . 895 . 896 .897.933.946.947.948) (Lutreola-863.875.893.897.929.946.946) (Luvarus-318)
(Luxllu3-870) (Lyeodes-27.106.109.114) (Lyeodontis-274 . 308 . 31 1 . 37 1 ) (Lygosoma-401 )
(Lymnaea-64. 366, 379. 389. 391 .396.404.411 .412. 433. 445. 459 .569 .570 .572 .578 .579 .611 .61 2. 61 5 .61 8 .626 .632. 633, 634,
637,648,651,682,687,773,784,791 .793,806,821 .841 .842.891 .892.893.895.897.936.937.938.939.940.941 .974) (Lynx-939)
(Lyrua-885)

(Habuia-401 ,479) (Habuya-475) (Maeaea-932) (Maeacus-835 . 878 . 936 ; 966 . 977 ) (Machetes-627 , 665 ) (Hacoma-739)
(Maerobrachium-87. 160.886) (Maorochlamys-91 4 ) ( Maerones-9 . 20 , 87 , 89 . 164 , 173 , 233 , 260)
(Maorophthalmus-729.734 .886) (Hacropodus-431 .707.715.873.880) (Hacrorhamphosus-41 ) (Hacrotus-815)
(Hacr our us-59. 283) (Hacruronus-281 .290) (Hactra-19, 19) (Haena-145.277.297) (Halacoclemmys-457)

170

THE TEXAS JOURNAL OF SCIENCE

DIGENEA, cont.

(Malaeoctenus-160,521 ,162) (Malapterurus-88.239) (Manta-315) (Mareea-570, 571 .612,629.632.639.693,791 ,
792,795,797) (Mari la-570. 606. 630. 658. 797) (Martes-837.932) (Mastacembalus-88. 167. 195.286) ( Mastacemberus- 1 32)

(Megalaspis-280 )
(Melanerpes-761 )
(Melanogrammus-271 )
( Me lano taenia-579 )
(Mellttophagus-71l3)

( Mastomys-938 ) ( Mazama-955 ) (Mediolaria-9) (Megaeeryle-596) (Megaderma-822)

(Megalobatrachus-86.375.913) (Megalornls-665.773.779) (Megarhynchus-762)

(Melania-288,719,823,857,867,878,932) (Melanltta-571 ,599.630.729.737.739.783)

(Melanoides-193) (Melanonyx-792) (Melanopsis-598) (Melanosuchus-536.537.599)
(Meleagris-569.633.678.708.788.797.879) (Meles-897 ,908,993) ( Mel ichthys- 195, 170.355)

(Melospiza-68 1.682, 760. 789) (Menidia- 13. 19.21 ,70. 199.219,282.297.708)

(Mentieirrhus-97.71.118.138. 195,161,166,213,219,282,708) (Mephitis-933. 996. 996. 997. 998) (Mergellus-571 ,
633,778) ( Mergus-570. 571 .579.580.588.599.609.626.627.632.633.639.695.699.669.699.699.695.703.708.722.791 .

793.869.879) (Meriones-908) (Merlangus-628) (Merlinus-166 .282) ( Mer luce lus-269. 266, 281 .282.290.292,308)

( Merluc i us-269 ,292) ( Merops-793 . 755 .76 1 ) (Merula-699.681 .761 ) (Mesodon-759 ,837.909) (Mesogobius-708.879)
( Mesomphlx-909 ) ( Hesothemis-667 ) (Metachirops-805) (Metachirus-950) (Microscelis-753) (Micraeca-632)

(Micrastur-586.758.759) (Microcarbo-582,636) (Mlcrogadus-121 .708) (Mlcrohyla-950) (Mlcrometrus-7 1 )
(Micronycteris-809) ( Mi cr opal ama-792) (Mlcroperca-226) (Micropogon-195,213)

(Micropterus-1 2 . 87 . 88 . 129.131, 152, 195, 197, 218, 223, 226, 293, 293, 366, 983, 597, 589, 595, 602, 622, 735)
(Microsarcop3-782) (Microtus-805.806.856,900.908.929,925.926,991 ) (Miliaria-789) (Milvago-752,758)
(Milvus-563, 579,580, 589, 585, 586, 587, 599, 600, 601 ,628,69M,695,695,696,669,692,699,701 ,706,719,715,716, 720, 753,
755,791,860,867,871 ,872,877,878,882,889,898,991) (Mimus-757 ,762) (Miniopter us-807, 809, 822, 823, 886)

(Mlsgurnus- 160, 183.229,230,931 ,591 .695.697.707.807.873.877.878,880.883.892) (Mitrella-52, 165)

(Mniotilta-682,683) (Modiolaria-17) ( Modiolus -20, 65 1 ) ( Mogurnda-65 . 87, 88, 160,230,260,286,572,579)

( Mola-290 ,319,321) (Molge-399 .902.912) (Molllenesla-189 ,798) (Molossldis-829 ) (Molossus-8 15. 821 .821 ,822.915)
( Mo lothr us-682 , 763 ) (Molpastes-619) (Molva-269) ( Monacanthus- 36 , 129 , 1 39 , 1 95 , 197 , 1 98 , 202 , 287 , 291 , 363 , 369 )
(Monatus-966) (Monedula-619,676) (Monocentr ls-58 ) (Monolene-39) (Monostoma-325) (Montifr ingil la-759 ,789)
( Morenia-508 , 557 ) (Moroco-65) (Morone-1 18, 131 ,138,195,219,226) (Horrhua-192) ( Moschus-830)

(Motacilla-6 19, 61 5. 61 6. 681 .729.767.773.789) ( Motella-208 . 21 9 ) (Moxostoma-1 16) (Mugil-12,

92,93,99,95,209,229,298,280,292,298,299,581 ,701 ,709,705,717,720,722,867,868,869,872,873,876,877,878, 880, 883,
889,885,885) (Mugllogobius-579) (Mulimulus-759 ) (Mulloides-157 ) (Hulloidlchthys-165) (Mullus-68,

197.165.166.168.292.293.708.708.879) (Munia-633) (Muraena- 17 .77 . 309 . 307 . 308) (Muraenesox-17.258.281 .310)

(Mus-705,797,806,871 ) (Musclcapa-6 16. 681 ,683.759,766.789) (Musculium-86 , 87 , 131 , 152,379,892)

(Mustela-9 13, 807, 81 9, 861 ,875,876,882,890,895,896,929,932,939,993,996,996,997,998) (Mustelus-90,256) (Mya-651 )

( Mycothera-677 ) (Mycter ia-569 , 591 . 687 . 688 ,689 ) (Mycteroperca- 17, 18, 107, 195, 179,213,219,278) (Myiospiza-763)
(Myiozetetes-758) (Myleus-356.361 ) ( Myl iobatls-90 ) (Mylossoma-356,361 ,363) (Myocastor-967 ) (Myopotamus-929 )
( My ospiza-7 88 ) ( Myotis-806 , 807 . 81 2 , 81 3 . 815 . 81 6 . 81 9 . 820 , 82 1 , 822 , 823 . 829 , 839 , 9 1 5 )

(Myoxocephalus-28, 119,269,271 .297.621) (Mystus-78) ( My tilus-9, 17.92,651 .739.790) (Myxus-161 )

(Naja-920, 933, 994, 459, 989, 991 ) (Nandus-689) ( Nannoperca-579 . 637 ) (Nannugo-907) (Narcine-21 3 )

( Naracion-257 ) (Nassa-51 ,5E, 195.214,651 .739,735) (Nasua-997) (Natalus-809,815.819.822) (Natrix-293,

395,396,930,931,933,939,935,994,445,947,948,999,950,951,952,953,955,958,959,983,988,597,599,558,573, 601 , 995,
950) (Necturus-1 31 ,226,382,398,913) (Nemachilus-101 , 102) (Nematoscells-319) (Nemichthys-170) (Nemorhaedus-829)
(Neobythites-70.299) (Neocar id ina-87 , 391 . 732 . 807 . 887 ) (Neofiber-925) (Neogobius-65 , 66 ) (Neomeris-853)

(Neomaenis-107, 110.213.273) (Neomys-809.859.860.885.900.916.930) (Neopercis-1 05 . 1 17 . 1 1 8, 167,299)

(Neophocaena-896.899.853) (Neophron-563 . 587 . 669 ) (Neosebastes-109, 392) (Neotis-799) (Neotragus-957)
( Nephel is-175 ) (Nephron-569.637) (Neptunea-212) (Nereis-51.52,210,727) (Nerithina- 1 60 ) (Neritina-1 17)

(Nerophis-121 ) ( Netta-629 . 630 . 639 . 692 ) (Nett ion-570. 609. 61 0,61 3. 628. 690. 661 ,791 .792.799.797) (Nettopus-791 )

(N inox-569, 585. 75 9) (Nocomis-88 , 1 1 8) ( Noc til io-9 1 5 ) (Notechis-929 ) (Notemigonus-180,592) (Notharchus-794)
(Nothura-676) (Notophoyx-566 ) ( No topter us-206, 688) (No to then la- 122, 27 0,299, 399)

(Notropis-88. 118, 129.588.589.602.638.693.856.890) (Noturus-152,295) (Nucifraga-759)

(Numenius-569. 575. 608. 626. 630. 633. 650. 651 .655.665.666.670.729.736.765.779.775.777.781 ,797) (Numida-696,678,

720,751) (Nutria-967) (Nyc talus-806 , 807 . 81 1 , 81 9 , 821 ) (Nyctanassa-582.601 ,671 ,679,

687,739,736,763) (Nyctea-563.585.695) (Nyctereutes-863 . 995 ) (Nycter is-823,824 ) (Nyctibius-563.791 )

(Nycticebus-820) (Nycticeius-447 . 81 1,821,822,823,825,830,838,915,916) (Nyc ticor ax-565 . 566 , 582 , 588 , 589 ,
591,618,628,629,649,695,696,699,650,673,686,687,688,689,690,692,703,709,706,707,708,719,715,720,869, 871 , 872,
873,879,877,878,898) (Nyctlnomus-807 . 81 1,822,823,829) (Nyroca-569. 570. 571 ,572,599,607,609,61 1,612,626,

629,630,632,633,639,660,664,665,667,670,693,708,727,728,729,735,768,778,783,789,791 ,792,795,797,879)
(Nystalus-758 )

(Obelia-149) (Oblata-53 , 62 , 101 ) (0cadla-955.506.532) (Ochthodromus-630,797) (Ocyurus-57 , 107 , 1 10,
195,213,232,259,292) (Odobaenus-899 , 851 ) (Odocoileus-955 ) (Odontobutis-88 , 89 , 229 , 697 ) (Odontogadus-219,269)
(Odontophorus-763) (Odontoscion-163) (Odostomia-296) (Oecetis-87 ) (Oedicnemus-569.761 ) (Oenanthe-760)
(Oenops-637 ) (Oidemia-599. 632, 650, 658, 699, 708. 729. 730. 738, 739. 770, 773, 778, 783, 79 1,795, 879) (Oligocottus-121 )
(Oligoplites-215.279) (Ollvella-727) (Olor-791 ,795) (Oncoides-838) (Oncomelania-732 . 937 )
(Oncorhynchus- 102, 103. 119. 124.132.269.286.297.305.319) (Ondatra-622,812,856,860,871,876,896,899,902,
909,915,916,923,925,968) (Oniscus-757 ) (0nos-9 ,121,271.289,306,974) (Ooeidozyga-375,931 ,950)
(Oph ice phal us-688 ) (Ophichthus-1 16) (Qphidium-109,309) (Ophiocara-645) (Ophiocephalus-65,77 ,101,
172, 195,260,286,687,689,690,707,873,877,878,880) (0phiodon-1 1 ,17,215,264,271,272,304) (Ophis-438)
(Ophisaurus-399 ) (Qpisthognathus-160, 162) (Opisthonema-29,266) (Oporornls-677 . 684 ) (Opsan us-21 3, 227, 232, 282)
(Opsarlichthys-65. 101 .707.873) (Orchestia-886 ) (Oreinus-689 ) (Orioclncla-716) (Oriolus-563.799.789)
(Ornlthorhynchus-663.903.917) (Orophls-474) (Orthagor iscus-21 5 , 31 8, 31 9 , 320, 322 , 325 , 337 ) (Orthetr um-390 )

INDEX: SYSTEMA HELMINTHUM

171

DIGENEA, oont .

( Or bhopr 1st 19-29,70, 71 ,76, 21 3, 232, 29*0 (Qrthostoechus-220) (Ortygometra-627 )

(Or yzlas-SSM, 836,845,71 5, 880) (Osmerus-8, 263, 271 .305.570,708) (Osteolaemus-537 )
(08traclon-31.32.117) (Ostrea- 18, 19,38,739) (Otis-865) (Otogyps-597 )

(Ovls-797, 835, 841 ,937,957,958,972,973) (Oxyehilus-677 ) (Oxyechus-782) (Oxyloma-882 )

(Oxyphilus-676) (Oxyura-626, 829, 632,791 ) (Oy am la-81 6)

(Ortygonax-758 )
( Os tlno ps-564 , 682)
(Ot us-585, 6 15, 709)
(Oxymycterus-837)

(Pachydlplax-667) (Paehynathus-9 1 ) (Pachyuromys-936) (Pachyurus-132) (Pagellus-102, 304 )
(Pagrosomus-37,38,51 , 105,124,214,225,280,281,305,333) (Pagrus-108) (Paguma-943) (Pagurus-479) (Palaemon-86)
(Palaemonetes-727 , 886 ) (Pallnurlchthys-145, 166) (Palonla-972.974) (Paludestrina-729,733,739)
(Paludlna-129,626,651 ,891 ) (Paludoaus-288,936) (Pend lon-563, 573, 574. 587, 604, 648, 768, 769) (Pangaslus-20, 357)
( Pan ur us-882, 787, 789) (Paplo-833,977) (Parabramis-880) (Parabuteo-585) (Paracalllope-159, 159)
(Paracentrotus-51 ) ( Par ache ilognathu-224) ( Par adoxur us-933) ( Pa r a fossar ulus-897 ,943)
(Paralabrax-157.165.271.308) (Parallchthys-20, 28 . 55 , 123. 165 , 166 , 21 3. 232.266 , 282, 290 , 307 , 308 , 371 )
(Paranthlas-149, 157,278,308) (Parapercis-29, 1 18. 139, 166,290) (Parapr istipoma-1 19 , 127 , 157 , 167 . 342)
(Parasllur us-20, 86, 88, 224, 230, 260, 571 .580,707,873) (Parathelphusa-932) (Paratraetus-213) (Parexocoetus-56,57)
(Paroarla-788) (Parophrys-142) (Parra-629,795) (Parupeneus-75, 165,710,717) (Parus-6 15, 681 ,683,755,756,767)
(Passer-586,614,615,665,666,667.670,676,681 ,682,742,743,748.753,761 ,762,767,770,788,789) (Pastor-759,789)
(Pavo-665) (Pavoncella-776) (Pecten-651) (Pedetalthyla-638.644) (Pediocetes-752) (Pelagia-318)
(Pelawys-253.276.337) (Pelates-722,868,880) (Pelecanus-239, 581 ,582,599,603,613,645,653,
692,701,702,711,714,716,718,720,724,768,769,770,867,869,877,882,884) (Pelecus-87 . 366 , 588 , 859 , 882)
(Pelidna-732,788) (Pelobates-54 ) (Pelomedusa-557) (Pelomys-938) (Pelophylax-374 ) (Pelotretus-28)
(Pelteobagrus-160,286) (Pelusios-436,463,481 ) (Peprllus-145,276) (Perameles-908) (Perca-8 , 9 , 86 , 88 ,
109,128,129,130,130,131,195,218,243,256,268,366,483,570,579,581,588,589,591 ,592,622,638,687,702,703, 735, 869,
870,886,890) (Percalates-579 ) (Perclna-87 , 88 . 101 . 132) (Percopsls-88, 132,218,579) (Perdlx-676 , 677 , 678)
(Pericroeotus-752) (Perlngla-708) (Perlsoreus-682) ( Per isted ion-39. 57, 169.266.283) (Perlthewis-667)
(Perla-821) (Pernls-563,644,692) (Peromyscus-838.876.908.910.9n ,913) (Peropteryx-822) (Petrochelidon-614)
(Petromy2on-591 ) (Phacellodomus-760) (Phacochoerus-971 ) (Phalacrocor ax-571 ,582,
583,597,600,603,608,619,621,628,631,632,633,636,637,638,644,646,648,649,688,689,692,702,706,708,711 , 715, 716,

720.722.769.869.872.873.882.883) (Phalaropus-617,787,791 ) (Phalloeeros-638.705) (Phasianus-666,678,
694,762,779,789,860,895) (Phllander-820.838,856) (Phllodryas-438.439.475.476) (Philohela-574)
(Phll0Hiachu3-569.6l7.628,648,68l ,791 ,795,899) (Philomyeus-678 ) (Philypnodon-241 .579,637) (Phlmosus-628)
(Phoca-708, 85 1.86 1,863. 869. 870. 892. 903) (Phoeaena-846.855.862) (Phocaenoides-846,850,853)
(Phoenlcopter us-626 ,630.631 ,774.792,794) (Pholas-222) (Pholis-708) (Phoxlnus-101 ,183.570,580,588) (Phoyx-688)
(Phragmitlcola-6 1 5 ) (Phr yganea-388 . 399 . 41 2 . 822) (Phyeis-121 ,214) (Phyllopezus-752) (Phylloscopus-767 )
(Phyllostomus-9 1 5 ) (Phyrne-374) (Physa-64 . 379 , 380 , 395 , 402 , 403 . 41 9 , 445 , 449 , 450 , 452 ,
454,459,534,570,572,589,611,612,614,618,626,628,629,632,633,651,793,841,891,892,895,923,936,938,948, 949, 954)
(Physella-405, 447, 458.61 1 .784) ( Physeter-847 ) (Physlculus-214 , 266 , 270, 281 , 304 ) (Physopsis-841 ,936,937)
(Piaraetus-345.361) (Piaya-586,744) (Pica-587,614,616,665,676,677,695,755,756,761,763,768,779,788,789)
(Pictada-9 ) (Picus-681 ,755,761 ) (Plla-893) (Pilherodius-646) (Plmelodella-228) (Plmelodus-132.228,270,356)
(Pimelometopon-140, 145) (Pimelopterus-1 11 . 135) (Pimephales-592.799) (Pipile-646) (Pipilo-682,754,755,760)
(Pipistrel lus-807, 811 .821.823.824.825.834) (Piranga-682,758) ( Pirenella-723 . 867 )
(Pisidium- 130. 131 .379.380.382.383) (Plsobla-727,730) (Pisodontophis-72) (Pitangus-585 , 645) (Pithecophaga-705)
(Plagyod us-264) ( Planar la-64 . 66 . 39 1 . 626 . 89 1 . 892 ) (Planestlcus-677 ) (Planorbls-194,
199,374,375,394,396,402,419,451 ,553,566,572,582,591,597,618,628,629,632,633,634,635,637,682,688,784, 791 , 792,
829,843,892,893,936,937,938,945,954,955,956,979) (Planorbula-409.566,946,946) (Platalea-591 , 648 , 659 , 666 ,
706,770,771,873) (Platanista-857 . 907 ) (Platax-10) ( Platessa-30 ) (Plathemis-390 , 409 ) (Platichthys- 19,267)
(Platycephalu8-18.21.108. 150.283.325) (Platyclchla-757.761 ) (Platyrhynoidis-90) (Platysomatichthy-263,271 )
(Plecoglos3us-120.224.230.647.715.882) ( Pleeostomus-94 , 1 46 . 287 , 359 ) (Plecotus-805 , 807 , 821 )
(Plectorhlnchus-69.70. 104,110,122.127) (Plectropoma-18) (Plectropterus-780.784 ) (Plegadls-648.776.787,789)
(Plethodon-24 3,399) (Pleurobrachla-149) (Pleurocera- 11 2, 198,547,602)
(Pleuronectes-27.29.30.52. 119. 121 ,212,213,270,271,293,305,370,708,873) (Pleuronlchthys-18) (Plotiopsis-645)
(Plotosus-58. 117. 167.297) (Plotus-608,622,628.629,634.687.693) (Pluvlalis-627) (Pneumatophorus-145,280)
(Podarcis-419.475) (Podargus-564 ,628) (Podiceps-571 ,590,591,614,637,638,645,646,648,649,689,694,

694.716.742.768.776.786.860.883) (Podiclper-646) (Podilymbus-638.644.697.784) (Podioecetes-676)
(Podoenemis-457. 458. 461 .462.477.492.554.556) (Poecllla-582) (Poecilonetta-605 , 660 , 693 ,797 ) (Pollocephalus-599)
(Pollaohlus-263.264 .271.708) (Polydactylus-15) (Polygordlus-264 ) (Polygyr a-459 ,754 . 907 , 908 , 909 , 91 1 )
(Polymix la- 120. 170. 305) ( Polynemus- 1 7 . 165 . 1 66 . 258 . 308 ) (Polyodon-285) (Polypedates-382 . 385. 389 , 400 ,750 , 950)
(Polyprion-127,281 ) (Polypterus-46) (Polypylis-614) (Polystlcta-572) ( Pomacanthus- 1 07 , 25 1 , 269 , 34 1 , 344 , 354 )
(Pomacea-639) (Pomacentr us-205. 292) (Pomadasys-70, 127,304) (Pomalobus-166) (Pomatiopsis-928,932,937)
(Pomatomus-145.166.213.708) (Pomatopsis-876.932) (Pomatostomus-681 ) (Poaolobus-166,232,294) (Pomotis-687)
(Pomoxls-131 .223.360) (Pongo-839) (Pontinus-169) (Poppella-268) (Poronotus-145, 166,280,708)
(Porphyr io-626 , 627 .628.694 .752) (Porphyr iola-628) (Por zana-627 . 639. 646. 681 .682.683,763,781 ,788,792,793)
(Potamoblus-1 18) (Potamon-807 , 865. 889 . 931 .932) (Potamopyrgus-125, 159,241 ) (Praomys-938) (Praticollela-754 )
(Prlacanthus-9. 43. 57. 169. 170.305.308) (Prlonodes-142.305) (Prlonotus-169) (Pristiophorus-196)
(Pr istipoma-1 57, 162. 195. 225. 226. 333) (Prlstis-214) (Procerodes-145) (Prochllodus-93. 366 ) (Procnias-760)
(Procto pus-644 ) (Proeyon-677.679.837.870.874.875.939.948.949) (Progne-758 ,762) (Promenetus-946 )
(Premier ops- 18. 2 14, 308. 329) (Promops-915) (Pronotogrammus-170, 309) (Prosopium-1 31 ) (Proteus-1 18)
(Protopsetta-20. 30) (Protopterus-35) (Prunella-684 .763) (Psammechinus-51 ) (Psectrotanypus-615)
(Psenopsls-280.302) (Psettodes-15) (Pseud acr 13-376,399,445, 449, 478, 632, 948) (Pseud ammlnicola-733)

172

THE TEXAS JOURNAL OF SCIENCE

(Pseudaphritls-160.579)
(Pseudeut.ropiu3-20.21 ,233)
1 18, 224, 647, 687, 699, 882)
(Pseudopitnelod us-48, 230)
(Pseudopriacanthu-169)

DIGENEA, cont.

(Pseudaspi us-88) (Pseudemys-452. 457. 458. 459. 460, 480. 506. 507, 51 6, 521 .533.553)

( Pseud is-375. 382. 404. 409) (Pseudobagrus-86 . 88 . 172, 260, 57 1 ) (Pseudogobio-66,
(Pseudogyps-567.799) (Pseudolabrus-105, 117.141,167) (Pseudoperil am pus- 599)
( Pseudoplaty stoma-81 ) ( Pseud opl euronect-28 . 138,145,166,212,213,708)

(Pseudorasbora-1 1 , 12,65, 180,224,230,599,600,644,647,694,706,714,720,872,878,882,884,

898) (Pseudorhombus-17. 145. 170.271 .282) (Pseudosalamandra-400 . 401 ) ( Pseudoscar us-89 . 205 . 251 , 345)
(Pseudosuccinea-379. 380. 406. 408, 459. 626. 841 .842.892.909) (Pseudotantalus-628) (Pseudothelphusa-932)
(Pseudotr i ton- 379 . 381,383) ( Pseud upeneus- 165 . 305) ( Pseudur la-768 ) (Pslttacula-661 ) (Psophia-752)
(Pteronura-874) (Pteropteryx-814) (Ptyas-400 . 425 , 430 . 431 , 444 , 479 . 488 ) ( Ptychocheil us- 1 17 , 588 )
(Puffinus-607, 628, 638, 711 ,712,716,722,768,883,884) (Pungtungia-224,647) (Puntius-65,707,715,873,880)
(Purpura-673) (Pu tori us-859. 861 .863) ( Pygosteus-8 ) (Pyramldula-91 1 ) (Pyrgophysa-566,936) (Pyroderus-564,583)
(Pyrogophysa-566,936) (Pyrreroides-707.873) (Pyrrhocorax-677 .789) (Pyrrhula-754) (Pyrrula-676,681 )
(Python-419, 429. 444)

(Querquedula-570.611.612.627.634.638.666.667.682.693.696.786.791 ,792,793,795,797)
(Qui seal us-579, 754, 757. 758, 788)

(Quickel la-683 )

(Rachycentron-213) (Radix-65,615) (Raja-196.256) (Rallina-677) (Rallus-612.627.629.676.681 .682,
699,734,773,780,781 ,787,792,793) (Ramphocelus-758.789) ( Rana-64 , 131 , 184 , 210 , 243 , 285 , 374 , 375 , 376 , 377 , 378 ,
379 ,380,381, 382 ,383,387, 388 , 389 ,390,391, 392 ,393,394, 395 , 396 , 397 , 399 ,400,401,403, 404 ,405,407,408,409, 410, 411,
412,413,419,430,431 ,445,447,449,451 ,453,454,463,479,485,541 ,548,553,566,573,583,591 ,618,632,633,644, 687, 689,
707,856,873,874,875,876,892,893,894,897,942,945,946,946,948,949,950) (Ranzania-321 ) (Rappia-409.446)
(Rattus-841 .871 .874.892.893.894.895.896.897.898.938.948.971 ) (Recurvirostra-627.649.670) ( Redunca-938 .
954,957,974,979) (Remora-264,304) (Retinella-909.910) (Retropinna-579.637) (Rhacophorus-400)
(Rhamdia-235.270.688) (Rhamphastus-587.759.788) (Rhamphocoelus-757.763) (Rhea-564) ( Rhinemys-554 )
(Rhinich thy s-589) ( Rhinobatos-90 ) (Rhinoceros-965) ( Rhinoderma-381 ) (Rhinodoras-132,219) (Rhinogobius-17,288)
( Rhinolophus-806 , 807 .809,81 1,821,822,823,824,867,907) (Rhinoplagusia-293) (Rhinopoma-821 ,822,829)
(Rhinoptera-84) (Rhodeus-88.224.714.872.878.882,950) (Rhombus-264,271,304) (Rhyacophilus-616.630,774)
(Rhyacosiredon-383) (Rhynchobatus-102) (Rhynchops-573.589,648,712,734) (Rhynchopsis-792)
( Richard sonius-260, 589) (Rlchmondena-761 ) (Riparla-615) (Rissa-571 ,578,649,708,874)
(Rita-78.80,82,102.172.233) (Rivulogammarus-160,658) (Roccus-214,215) ( Roeboides-285 ) (Roncador-47 .71 )
( Rostratula-644 ,732) (Rostrhamus-639,775,792) (Rucervus-955) (Rumina-909) (Rupicola-758,759)
(Rupornis-758,759) (Rusticola-688) ( Rutilus-8 . 65 . 86 . 87 ) (Rypticus-17. 18)

(Saccobranchus-687 ) (Sagitta-149,263,270,271 ) (Salamandra-399 , 411 . 412) (Salangichthys-716,883)
(Salminus-17 , 48,79. 285 , 356 ) (Salmo-60, 86 . 88. 1 17 . 1 18, 121 . 129.130.131,132,159.160,195,263.264,271,297,305,344,
355,356,361,362,363,589,703,870) (Salvelinus-86 . 87 , 88 . 131 , 132. 195,256,264,271,297,703,870) (Sander-591)
(Sarcld iornls-783 , 797 ) (Sarcocheil ichthy-1 1 8, 224 , 260 , 340 . 599 . 694 , 882) (Sarcogrammus-590,593) (Sarcogyps-564 ,
574,584,586,600,692) (Sarcorhamphus-564 . 587 . 637 ) (Sarda-1 1 , 19 , 166 , 293 , 307 , 308 , 329, 338) (Sardinella-266 )
(Sardinia-267) (Sargus-53. 62. 101 . 147.267) (Saur ida-265 . 281 . 290 . 304 . 308 . 371 ) (Saurophaga-756) (Saurothera-743 )
(Saxlcola-767) (Sayonara-170) (Scaphirhynchus-132,209.210) (Scardinius-8 , 64 , 87 ,
132,366,581,588,591 ,702,703,856,863,869,898,942) (Scarus-105,251 ,341 ) (Scatophagus-9.249,258)
( Schilbeodes- 1 3 1 ,221 ,245) (Schistosomophora-937 ) (Schizothor ax-87. 102, 131 .689) (Sciaena-20,80,200,207,
225,258,280,282,293,337) (Sciurus-161 .949) (Scollodon-368) (Scolopax-569 . 570 . 586 . 593 . 594 . 616 . 627 , 633 . 773 .79 1 )
(Scomber-1 9. 42. 43. 104. 110. 121. 149. 166.252.276.277.280.337.708) (Scomberoides-277 ) (Scomberomorus-1 1 . 12 . 19 . 327)
(Scombrops-21 . 157.170) (Scoplmera-732) (Scops-587) (Scopus-574 ,629) (Scorbicularia-651 )
(Scorpaena-9, 109. 120. 164. 170.200.270.271 .281 ,290,293) (Scorpaenichthys-18,272,282) (Scotophilus-822,823,838)
(Scyllium-53.90) (Scymnodon-196) (Sebastes-121 .264.271 ) (Sebastichthys-157) (Sebastiscus-1 17. 128,271 )
(Sebastodes-28.57.58. 108. 117. 120. 142. 149. 157. 158. 165.213.264.271 ■ 308) (Sebastopyr-1 8)
(Segmentina-375.614.843.892.893.894.897.950) (Segnitlla-955) (Seiur us-677 .683.755.788) (Selar-71 . 167,169)
(Semlcossyphus-37 . 140) (Semlnatrix-449) (Semisulcospira-230,287,645,647,706,707,
715,716,792,824,872,873,882,883,898,931 ) (Semnopithecus-829) (Semotilus- 1 2. 66 . 88 . 101 , 102,589,592)
(Seriola-9. 18. 20. 208. 21 3. 267. 276. 282. 308 .325. 327. 334. 335. 342. 367) (Ser iolella-280 ) (Seriolichthys-302)
(Serranus-10. 15. 17. 18.89. 107. 108. 121 .282.304.342) (Sesarma-932,933) (Setarches-290 ) (Sewertzovia-914)
(Sialia-762) (Slbynomorphus-480) (Siganus-159 . 201 . 251 . 292 . 294 . 345 . 351 ) (Sika-926) (Sillago-57, 104,
105,110,118,722) (Silundia-20.233.357.362) (Silur us-8.88. 160, 172.356.361.363,588) (Simlimnea-572,632,841 )
(Slphonostoma-121 ) (Slphostoma-1 34 ) (Sir edon-4 1 2 ) (Sirembo-71 , 170) (Siren-413) (Si strur us-450 ) (Sitta-742)
(Smaris-62.277.293) (Solariella-30) (Solea-168) (Somateria-569. 570, 571 ,572,633,645,708,
728,729,739,768,778,783,791,795) (Sorex-809.830.876.907.908,913.916.931 ) (Spar lsoma-17 , 107,205,251 )
(Spar us-29. 37. 52. 80. 105 , 1 1 8, 120 , 159 , 159 , 281 , 346 , 347 ) (Spatula-570.61 1,634,667,768,783,785,791,792,795)
(Speotyto-752.760.789) (Sphaerechinus-51 ) (Sphaerium-101 .131 .132.379.380.381.382) (Sphaeroides-36 , 59 .
73,88,91,112,113,134,149,166,188,214,293,316,319,348,349,367,370,658) (Sphargis-496 ) (Spheniscus-573 . 694 )
(Sphyraena-9 .11. 12.18.19.20.213.282.307.308.325) (Sphyrapicus-681 .682) (Spilogale-947) (Spi lornis-564 )
(Spilotes-444,467) (Spinachia- 121 ) (Splnaperca- 11 ) (Spinus-614) (Spio-145) (Spiral ina-565 . 629 )
(Spizaet us-564 ,565) ( Spod iopsar-6 1 4 ) (Spondyl iosoma-56 ) (Squalius-.IOI , 175,588) (Squal us-90 ,196,256,314,315)
(Squatarola-630.636.727.729.734.741 .781 ) (Stagnicola-445,569,570,578,579,61 1,639,793,806,807,
842,896,923,941,955) (Stegostoma-84 ) (Stenella-95 1 ) (Stenodus-195) (Stenopharyngodon-714,943)
(Stenopsyche-806 . 824 ) (Stenostoma-91 1 ) (Stenothyra-224 ) (Stenotomus- 145,166) (Stenotrema-754 ,912)
( St er cor ari us-571 .578.578.708) (Stereolepis-774 ) (Sterna-570 . 570 . 57 1 . 573 . 578 , 579 , 580 , 599 , 600 , 602 , 603 , 606 ,

INDEX: SYSTEM A HELMINTH UM

173

DIGENES , cont.

607, 61 3, 61 5, 617, 61 8, 632, 648, 61)9,652,673 ,688, 696, 698, 703 ,708, 71 1,722, 723 ,724, 729, 734, 755, 768, 787, 789, 869, 874,
899) ( St er no ther us-4 14, 457, 459, 5 16, 52 9) (Stizosted ion-21 . 88 . 195 , 21 8, 366 , 589 , 595 ,702) (Storer la-478 . 479 )
(Streblospio-727) (Strepera-755) (Strepsllas-674 ,727) (Streptopelia-563,629.676,681 .755)
(Strix-563 , 585 , 586 , 587 , 627 , 629 , 660,675,676,755,755,878) (Stromateoides-280 , 333) (Stromateus-219,280)
(Strongylura-12, 13,21 ,56,205,297) (Sturnella-754,757) (Sturnla-563.614) (Sturnopastor-599 ,761 )
(St urn us-6 14, 6 15, 665, 676, 681 ,742,762,763,767) (Subulina-678,687,788,838) (Subzebrlnus-914) (Succlnea-681 ,
682,683,908,909,910,914) (Sufflamen-1 88 ) (Sula-603,712,719.722,769.871 ) (Suncus-806 , 950 ) (Sus-835,
905,968,971 ) (Sylvaemus-838) (Sylvia-616,681,767) (Sylvilagus-9 1 4 ) (Sympetrum-409 ) (Symphaemia-774 )
(Symphurus-76 ) (Synagrls-1 18, 188) (Synagrops-12, 170) (Synaphobranchus-266 , 304 ) (Synaptura-17,40)
(Syndoamya-19) (Syngnathus-1 19, 121 ,162,165,167,709) (Synodontis-44 , 46) (Synodus-15,70. 118. 166,167,279,
282,290,305,308,310,342) (Syrietes-166) (Syr igma-635) (Syrmatieus-677 , 682) (Syrnlum-585 , 587 , 627 , 629 , 757 )

(Taehyeres-793 ) (Tachyphonus-682 ) (Tadar lda-807, 81 5, 821 ,822,824,830) (Tadorna-570, 727, 791 ,794,795)
( Ta lore best ia-7 34 ) (Tal pa-808, 905, 91 2, 930) (Tatnias-91 1 , 91 3) (Tandanus-259 ,260,637) (Tantalus-567,591,618,689)
(Tapes-1 9, 53, 145,221 ,222,650,739) (Taphozous-822) (Taplrus-966 ) (Tarpon-308) (Tartaruga-502)
(Tautoga-52, 138, 166,708,709) (Tautogolabrus- 1 45 , 166,708,709) (Tax idea-947 ) (Taxydromus-431 ) (Tayassus-968 )
(Teleseoplas-12) (Telmatias-594 ) (Temenuchus-744,761 ) (Terathropius-563,799) (Terekia-633 ) (Terrapene-399.458)
(Te3tudo-457.459.482.492) (Tetragoneur la-666 , 667 ) (Tetragonopterus-153) (Tetrao-676,764) (Tetraodon- 112, 200 )
(Tetraogallus-678,755.760.764) (Tetrapterus-332) (Tetrastes-683.754.764) (Tetrathyra-480 )
(Tetrodon-18,36,48,201,245) (Teuthis-37 , 67 . 107 . 251 . 273 , 292 . 296 . 298 , 345 , 351 , 868 , 877 . 878 . 880 )
(Thalasseus-57 1,607) (Thalassochelys-432 ,451,453,462, 465 , 479 , 496 , 497 ,500,505,518, 529 , 552 )
(fhalassoma-58, 108,119) (Thalassornis-612) (Thamnomys-938)
(Thamnophis-243.445,447,449,450.452.453,454,458,460,555,946) (Thelotornls-436) (Theodoxus-65 ) (Theragra-165,
264) ( The rapon- 157, 227, 29 3, 64 5, 707, 722, 87 3. 877 ,878, 880) (Thereiceryx-779 ) (Ther 1st icus-563. 630, 646, 648)
(Thlara-706) (Thraupls-757.760.789) (Thymallus-86 , 88 . 129, 130. 131 . 132, 195) (Thynnus-12,21 ,252,
253,304,325,326,327,328,329,334,335,337,338) (Thyrlnops-166) (Thyrsltes-307.314) (Thysanoessa-314) (Tiara-872)
(Tlgrl soma-649 ,688) (Tllapla-601 , 867 , 87 1 , 878 , 883. 94 1 ) (Tillqua-401 ,479) (Tlnamus-629 , 676 )
(Tinea-65,87, 101 , 175,366,582,591,855.856,863,942) (Tomichla-628) (Tomodon-438) (Torpedo-196,197)
(Torqullla-829) (Totanus-579, 648, 649, 674, 682. 708. 727, 732. 773. 774. 776. 777. 778, 781 ,787,874)
(Toxostoma-596,754 ,762) (Toxotes-160) (Trachlnocephal us-281 ,304) (Tr achlnotus- 1 1 .43,142.149,166.167.

287.293.319.708) (Trachlnus- 1 0 , 1 09 , 237 , 27 1 , 292 , 304 ) (Traehurops-43 . 154, 166, 169,277) (Traehurus-42 , 43 ,

63.154.157.169.170.277.278.293.297.304.708) (Trachynotus-10) (Trachyphonus-763 ) (Trachypterus-277 )
(Traehysaurus-476,481 ) (Tragelaphus-938. 974) (Tragulus-97 1 ) (Trematomus-102, 121 , 142,267,293) (Triacanthus-43)
(Trlchechus-919,966) (Tr lehiur us-1 9, 21 , 145 , 195 , 304 . 308) (Trichogaster-689) (Tr identiger- 160, 180,224.882)
(Trlgla-1 1,1 8, 21, 108. 109, 21 5. 264. 270. 301) (Tr iglops-109) (Trigonorrhlna-90) (Tr imeresurus-438 . 444 )
(Tr lnga-565,569. 578. 609, 61 5. 61 6, 629. 632. 633. 649. 650. 651 .665.673.681 .682.696.726.727.729.730.733.737. 768, 773,
774,775,776,777,782,787,788) (Tr lonyx-396 . 424 , 480 , 481 , 484 , 509 . 532) (Tripterygion-167)
(Trlsotropis-1 8) (Triton-375,399,411,451) (Tritur us-376 .377,381, 382 . 383 . 399 ■ 405 , 406 , 454 ) (Trochalopteron-744 )
(Troglodytes-758.788) (Trogonurus-758 ) (Troplcorbls-938) (Tropldonotus-396 ,
402,419,420,424,425,430,431,433,433,434, 444, 445, 451, 452, 454, 458, 479, 482, 485, 541, 548, 549, 558, 563)
(Tropldurus-476) (Trutta-102, 121 . 131 , 195) (Trygon-84) (Trypanoeor ax-564 .676,763) (Tuplnambis-475 )
(Turacus-675) (Turdus-675. 676. 681 .683.684.699.742.754.757.761 ,762,763,764,774,789,789) (Turnix-682)
(Turrls-1 49 ) (Tur siops-951 ) (Turtur-676.682.703.869.882) (Tuxtepec-615) (Tylosurus-21 . 204 ) (Tylototr lton-400)
(Tympanotonus-720.867.884) (Tyrannus-615.762) (Tyto-567.587.759)

(Oea-734) (Umbra-86.89.890) (Umbrina-47 .71 . 102.170.212.225) (Unlo-8) (Upeneoides-70. 157. 158)
(Upeneus-57 , 142. 157 . 158) (Upupa-616.648.732.754.765) (Uraeotyphlus-381 ) (Uranoseopus-10.214.236.237.238.240)
(Ur la-570. 570. 571 .579.647.708.737.768.769) (Urlnator-647.649) (Urobatis-90) (Uroeissa-764.773.774)
(Urocyon-946) (Uroleuea-761 . 789 ) (Uromastix-454 . 466 . 474 ) (Urophye is-39 . 121 . 142 . 166 . 214 . 264 . 265 . 266 . 290 , 293)
(Urosalplnx-673) (Urotrlchus-910) (Urubitinga-585 .586)

(Valvata-626.633.891 ) (Vanellus-569. 570. 580. 626. 632. 633. 636. 665. 681 .727.729.756.768.773.777.781 .782.791 ,
792) (Var anus-420, 44 3. 454. 473. 484) (Var leorhlnus-869 ) (Ventridens-909,910.912) (Venus-740) (Vertigo-914)
(Vespertillo- ! 30 , 805 , 80? , 81 7 , 82 1 , 82 1 , 823 , 824 , 825 , 826 , 829 , 907 , 924 ) ( Vesper ugo-81 1 . 81 8 . 821 . 824 . 825 ■ 826 )
(Vipera-433,436,479.485) (Vireo-683.684.743) (Vlreosylva-682 ) (Viverra-836.933)
(Vlvlparus-1 32. 602. 61 8. 628 .635. 683. 893 .894. 897 .898 .899 .943) (Volatlnia-758) ( Vul panser-739 )
(Vulpes-597. 703. 705. 708. 837. 859. 861 .869.870.871 .872. 874 .875 .882 .94 3 .945 .946) (Vultur-564)

(Wallago-43.78.260) (Wallagonia-82) (Wllsonla-755)

(Xanthoeephalus-614) (Xanthornus-758) (Xenistius- 158) (Xenodermichthys-309) (Xenodon-438,449)
(Xenomystax-1 18) (Xenopus-375 . 402) (Xenorhynchus-618.642) (Xenotis-589 ) (Xerophlla-829 . 909 )
(Xesur us-1 49. 160.173.292.351 .352) (Xiphlas-253 . 327 ) (Xiphocolaptes-757 .760) (Xiphostoma-48 ) (Xystrias-304)

(Zacco-101 . 119.224.599.647.706.714.872.878.882)
(Zamenis-431 ,444,449,450,454,479,485) (Zanelus-371 )
(Zenop8is-170) (Zeugopterus-108) (Zeus-108.212,307)
757,910,912) (Zonotrlch la-755. 758 .760) ( Zonurus-473 )

(Zaloey s-149,278) ( Zalophus-648 . 847 , 870 , 899 )
(Zapus-925.926) (Zebrias-150,293) (Zebrina-829)
( Zoarees-58 ) ( Zoniehthys-208)

(Zonltoides-677,

174

THE TEXAS JOURNAL OF SCIENCE

CESTODA (vo/ll)

(Abramls-20, 21, 2*1,58, 187) (Acanthlas-65.80,86,92. 113, 117. 135) ( Acanthocottus-45 ) ( Acanthodactylus-179 ,
415) ( Acartia-126) ( Acciplter-235, 345) (Acerlna-38,58. 141 , 1*42) ( Acheilognathus-44 ) ( Achipterla-385)
( Aclnonyx-4 37 ,**38) ( Aclpenser-12, 35 . 38 . 40 . 143) ( Acomys-406 , 447 ) (Acridotheres-231 .236,252.290,293.308,
322,323,324) (Acrls-162, 165) ( Acrocephalu3-235.273) ( Actitis-218,260,289) ( Actophilus-346 ) (Adenota-381 )
(Aegialitis-226,252,253.260,261 ,299,300,31 1,313,337,342) ( Aegithalus-253.271 .272) ( Aegollus-269.270)
(Aepyceros-380,381 ,398,399.400) ( Aethecfiinus-394 ) ( Aethiopsar-309) ( Aethya-310)
( Aetobatis-62,69,73,74,76,80,83,84,89,96,98,99, 100, 103, 113, 134) (Aetomylaeus-103) ( Afrotis-225) (Agama-179,
180) ( Agonoderus-219) ( Agr iol imax-202 , 208 , 21 8) (Agrion-332) (Ailurophis-414,415) ( Alx-289 , 292 ) (Akis-418)
(Alauda-1 91 ,260,262,271 ,273,325) ( Alburnus-20 , 58 , 143, 187) (Alca-188, 193.235.238) ( Alcelaphus-381 )
( Alces-380,381 ) ( Alector ls-215 . 216 . 21 8 . 220 . 308 . 325) (Alepisaurus-78) (Alepocephalus-137) (Alestes-27.29)
(A1 log alumna- 385) (Allotis-140) ( Alopeclas-HO) ( Alopex-360 . 364 , 437 . 442) (Alopias-66,69.75)
( Alouatta-395 , 404 , 405 ) (Alphitoblus-219) ( Alphitophagus-260 , 41 9) (Alutera-59. 115) ( Amara-218.219.221 )
( Amber i za-271 ) ( Amblopl ites-4 5 , 58 , 1 40 . 142 ) ( Ambystoma-1 62 ) (Amelur us-51 .58.140, 142, 144,249) ( Ameiva- 1 79 , 1 80)
(Am la-51 ,58,140, 142) ( Ammocoetes-1 87 ) ( Ammodytes-46 , 135) ( Ammospl za-260 ) ( Ammotragus-399 ) (Amphlbolurus-174)
( Amphiodon-45) (Amphlsbaena-181 ) ( Amphluma-162) ( Amyda-176) ( Anaferonla- 219) ( Anarhlcas- 4 7 )
(Anas- 187, 188, 200, 21 0,21 3. 21 6. 21 8. 220, 222, 236. 257, 284, 285, 287. 288, 289. 290. 291 .292,293,295,297,298,300,301 ,304,
305,306,308,309,310,311,312,316,317,318,319,320,324,325,326,328,332,334,348) ( Anastomus-222)
( Aneistrodon-174 , 175 ) ( Ancylochllus-226.238) ( Aneldes-165) (Anguilla-44,
46,47.58,113,126,141) (Anhinga-243,325) ( Anlsodactylus-219) (Anisolabis-418,419) ( Annlella-179) (Anoa-381)
(Anolis-179) ( Anorthura-231 ) ( Anser-1 91 . 289 . 291 . 295 , 297 . 304 , 310 . 31 8, 321 , 322, 325) ( Anseranus-320)
(Anthicus-219,419) (Anthus-231 .235.244 .262.271 . 309) ( Antilocapra-380 , 397 ) (Ant.llope-380.381 ) ( Antrozous-394 )
( Apel tes-45 ) (Apes-136) ( Aphod lus-21 8 , 21 9 . 260 . 323 . 4 1 9 ) ( Aphredoderus-51 .140) (Aphrodite-134) (Aplites-140, 142)
( Aplodlnotus-140, 142) ( Aplopel ia-21 8 , 31 9) ( Apocellus-260)
(Apodemus-345, 385. 404. 41 8, 421 .427,429.432,441 ,447.448) (Apogon-45) (Aprion-128) (Aptenodytes-195.254)
( Apteryx -2 37, 25 1 ) (Apus-270) (Aqulla-191 .264,274) (Ara-215) ( Arantia-218) ( Archibuteo-344 , 345)
( Archiptera-372) (Arctocephalu3-68,353) ( Arctomys-426) ( Arc tone tta-246, 288, 291 ,292) (Ardea-185, 187,
188,231 ,232,233,235,237,242,244,246,316,335,354) (Ardeola-231 ,232,233,244,254) (Arenaria-260,297,298)
(Argentina-113.134) ( Arlon-208,235) (Arius-93. 132) ( Arnoglossus-44 , 45 . 47 , 126 ) ( Arquatell a-289 , 299 . 307 , 31 8)
(Artediellus-34) ( Arthrenus-394 ) ( Arthroleptls-165) (Arvicanthis-389,406,407,418,432)
(Arvicola-384,385,418,419.426,431 .433.434) ( Asio-269 . 345 ) (Aspldltes-175) (Aspius-21 , 143,187) ( Astacus-31 1 )
(Astimastlllas-273) (Astur-270.345) (Asturina-274) ( Ataenius-260) (Athene-261,263) ( Atherls-175)
( Atractosteus-141 ) ( Attagenus-394 ) ( Attlcora-238) ( Auchenoglanis-30) ( Auxis-1 15)
( Ay thya-290 ,292,304,305,318,319,334)

(Bagarius-12. 143) (Bagre-1 15) ( Balaenoptera-356 . 366 . 368) (Ballstes-128, 129.134,136) (Balopogan-315)
( Bar bus-20, 21 .38.41 ) ( Bar tramia-237 ) ( Bassar lscus-364 ) (Baslllchthys-141 ) (Batrachoseps-165) ( Belone-54 )
(Belonopter us-237. 260. 262. 333. 342) (Beryx-46) (Betta-56) (Bison-381) (Bitis-174, 175)
(Blarina-395, 42 1,425, 426, 427) ( Blastocercus-380 . 397) (Blatta-405) (Blicca-21 , 187) (Boa-170, 173, 182)
( Boiga-1 79 ) ( Boleosoma-45 . 47 ■ 140) ( Bonasa-208,209 , 260, 298 , 330) (Boodon-175) ( Bos-380 , 381 , 397 , 398 , 399, 400)
( Bo t a ur us- 188, 242, 24 3, 24 4) ( Bothrops- 173 . 174 . 175) (Bothus-44) ( Brachycellus-219) (Brachyteles-395)
( Br achy plat ystoma-69 . 147.148.151.155.156) ( Brachyplatytoma-155 ) ( Brachypter us-221 ) ( Br achyteles-381 )
( Br adytus-2 1 9 ) ( Brama-45 , 1 1 0 , 1 1 5 , 122 , 1 35 , 1 36 , 1 37 ) ( Branchiostegus-45 ) ( Branta-295 . 305 . 334 ) ( Brevoortia-1 31 )
(Brotogerys-209.21 1 ) (Brotula-45) ( Bubal is-400 ) (Bubalus-399 ) (Bubo-269,270) (Bubulcus-308) (Bucco-279)
( Bucephala-246 . 289 . 305 . 322) ( Buceros-2 1 4 ) ( Buchanga-222 , 323 ) (Bucorax-203,224,227) (Bucorvus-223.224,227,270)
(Buffelus-398,399) (Bufo-162, 164, 165, 166,413,414) (Bulimulus-208) (Burhinops-252,265)
(Burhinus-232, 235, 25 1.252, 261 .342) (Busarellus-232) ( Butaster-223 , 257 ) (Buteo- 191 ,222,223,345)
(Butorldes-215.233.249.294.346) (Bycanistes-203.213) (Bythotrephes-140)

(Cacatua-201 ,213.215,216) (Caccabis-210, 215 . 21 8,219 . 222.260, 275.278, 303, 325 ) (Cactua-215)
(Cairina-279.287.288,319) (Calathrus-219) (Calathus-218,260) (Caleonyx-180)
( Cal idrls-226, 238, 256. 260, 261 . 289 .298.299 . 300 . 301 . 31 1 .325.327) (Callianassa-1 18) (Callithrix-395,419)
(Callorhinus-353.357) (Callorhynchus- 1 7 ) (Caloenas-216) (Calotes-179, 180) ( Calyptocephalus-162)
(Calyptomena-213) ( Camelus-380 , 398 . 399 . 400) ( Campethera-2 1 4 ,216) (Cancer-118) ( Cancroma-246 , 255 ) ( Candona-289 )
(Canis-360. 361 .364,365.410.411 .414.437.438.439.442.443.444) (Cannochaetus-400) (Capella-231 ,235,239,252,
253,256,258,289,291,296,297,300,301 ,311,327,344,382) (Capoeta-20) ( Capra-380 , 381 , 398, 399 , 439)
(Capreolus-380. 381 ,397.398) (Caprimulgus-2 18. 231 .248.251 .252,253,273.323.346) (Capromys-376,382,405)
(Caracina-346) (Caranx-1 1 5 . 1 34 , 1 37 ) (Carassius-20,29, 186. 187) (Carcharhinus-67.69.72,89, 113, 115, 128,
129,131,134,135) (Carchar ias-62. 66 . 67 . 68 , 69 .71 .72 . 83 . 87 . 88 . 89 . 102 , 11 3 , 1 15 , 1 1 8 . 128 , 1 30, 134 , 1 35 , 154 , 157 , 158 )
( Carcharodon-67 .71 .102,128.154) (Carcinus-1 17) ( Cardinal is-278 ) (Cariacus-380) (Cariama-224) (Carina-246)
( Car inogammar us-9 ) ( Car piodes-23 . 25 . 26 ) (Carpophaga-200.213,214,215) (Casarca-295,297) (Cassicus-222,271 ,279)
(Casuarlus-222) (Casus-440) ( Cathar acta- 1 94 ) (Catonotus-140) (Catostomus-21 ,22,23,58, 187) (Causus-176)
(Cavia-432) ( Cebus-375 . 381 . 395 ) (Celeucides-309) (Celeus-215,221 ) (Celia-221) (Centrarchus-140, 142)
(Centrocercus-218.278.330) (Centrologhus-38 , 42 , 50 , 357 ) (Centrophor us-66, 67, 110, 126, 132) (Centroprlstis-129)
(Centropus-213.214.215.218.220) (Centroscymnus-126, 127, 133) (Centurus-220) (Cephalophus-380 , 381 , 398 , 399 )
( Cephaloscyll ium-76 ) (Cepola-45, 133. 135) (Cepphus-237.238) ( Cerastes- 179 . 180 . 41 3 . 41 4 ) (Ceratacanthus-1 15)

INDEX: SYSTEM A HELMINTHUM

175

CESTODA, cont.

(Cerato£h£llus-418,419,432) (Ceratozetes-385) (Cerchneis-191 .236.237 , 345 ) (Cercopithecus-390.408.418,431 .444)
( Cereus-4 39 ) ( Cerorchinea-238 ) ( Certhia-326 ) ( Cervus-380 , 397 , 437 ) (Cestrac ion-67 , 75 . 83 , 88 . 1 1 3, 1 1 4 , 154 )
(Cetopsis-151 ) (Cetorhinus-71 ,75,80) (Chaenobryttus-45, 190) (Chaenogobius-105) (Chaetura-236)
(Chaeturlchthys-131 ) (Chalcaburnus-58) (Chalcides-179) (Chalcidus-915) ( Chal cococcyx- 323 )
(Chalcophaps-210,216,292) (Chalococeyx-323) (Chamaeleo-180, 181 )
( Char ad ri us-226, 23 3, 239 ,235, 236, 238, 252, 253, 257, 260, 261, 262, 269, 289, 299, 312, 323, 333, 337, 338, 392)
( Chare harhlnus-88 ) ( Chaulelasmus-292 , 293, 297 , 31 1 . 31 8, 31 9, 339 ) (Chelidon-238) (Chelidonaria-236,238)
(Cheliodonichthys-129) (Chelodlna-168, 182) (Chenolopex-285) (Chenopis-295.398) ( Chenoplox-296 )
(Chenopsis-293,286,293) (Cherubineho-131 ) (Chilonycteris-931 ) ( Chi loseyll ium-9 , 66 , 70 , 89 , 87 , 91 ,157)
(Chimaera-16, 17, 18,66) (Chlonis-251 ,289) ( ChionomyA-373 ) (Chiroeentrus-132) ( Chironemus- 1 35 ) (Chirostoma-187)
(Chlamydera-251 ) (Chlamydoselachus-76, 138) (Chlamydotis-272,318) (Chlorls-309. 326) ( Chlorophis-9 1 3 . 9 1 9 )
(Chlorophthalmus-55) ( Chlorura-269 ) ( Choeridium- 219) ( Chondrostoma-20 , 21 ,187) ( Chord eiles-290 , 253 )
(Chorinemus-115.132.136) ( Chor iotis-229 , 225 , 395) (Chorthippus-275) (Chrysichthys-25, 191 , 193)
(Chrysochlorls-232,908,929,933) (Chrysococcyx-217) (Chrysocolaptes-295) (Chrysolampis-319)
(Chrysomitris-271 ,278) (Chrysopelea-179) (Chrysopelia-180) (Chrysophrys-1 16) (Chrysotis-203.215.381 )
(Cichla-191 , 192) (Ciehlosoma-96) (Ciconia-1 87 , 188 , 261 ) (Clnclus-236.238. 396) (Cinnyrls-227,273)
(Clrcaetus-191 .279,395) (Circus-191,252,319,399,395) (Citellus-379 , 376, 385 , 399 , 912 , 9 1 8, 91 9, 937 , 942, 445)
(Citharichthys-120) ( Civettictis- 9 13,919) (Clangula-1 87 , 246 , 251 . 258,290, 293 , 300, 31 0, 31 1 , 312, 329, 334)
(Clangulamareca-309) (Clareola-312) (Clar ias-20, 21 .27,28.29,31 .55,56, 191 , 150) (Clarotes-193)
(Clethrionomys-373.385.919,992.997.998) (Clevelandia-125) (Clibanarius-1 18) (Clivicola-239,236)
( Cl upea-38, 39. 99, 126.130. 132) (Cnemidophorus-179) ( Coassus-380 ) (Cobltis-20, 193. 187) (Cobus-399)
(Coccystes-270) ( Coccyzus-270 ) (Cochlearius-259 ) (Cochliocopa-912) (Coelopeltis-179.913.919)
(Coelorhynchus-133) ( Coendou-909 ) (Coephloeus-221 ) (Colaptes-213,219,222) ( Collnus-212 . 21 3 . 215 . 218, 221 )
( Col i us-209 ,217) ( Collocal ia-253 . 256 . 266 . 276 ) (Colobus-375) (Coloeus-231 .236, 309. 315) (Coluber-175. 176, 180,
913,915) (Columba-202. 209, 208, 209. 210, 21 1,21 3. 21 9, 21 5. 21 6. 21 7. 21 8, 260, 263. 292, 305, 3 19, 329, 330)
(Colymbus- 185, 186, 187, 188, 192. 193. 199, 196, 286, 296, 310. 31 1.31 9. 325. 332, 391) (Conger-126, 133) (Conopophlla-220)
(Constrictor- 170. 173) (Conurus-215) (Copsychus-315) ( Coraclas-270 . 273 . 308. 315 . 325) (Cordeiles-298)
(Cordeilus-323) (Coregonus-33,39,38.39,97.58. 190, 191,192,193.187,359) (Coris-116) (Corone-220, 231 .232,323)
(Coronella-175) (Corophium-9 ) (Corvus- 189, 188,210,216,220,222,
227,231 ,232,235,236,238,296,253,260,261 ,270, 308,309,310,315,322,323,332) ( Cor yl 1 is-203 . 209 ) (Coryphaena-109.
111,112,128,136) (Corythaeola-213.217) (Coscoroba-288,296) (Cossyphus-136) (Cottus-38,44,46,58, 140,
192,187,188) (Coturnix-217 ,218.219, 220,260,278, 298 , 325) ( Cratacanthus-260 ) (Crathacanthus-219) (Craticus-261 )
(Crax-215) (Crematogaster-212) (Crex-238) (Cricetomys-389 . 390) ( Cr icetulus-379 . 399 , 4 1 8. 926 , 928)
(Cricetus-426,427,432,437) (Crlniger-316) ( Cr isti vomer-33 . 38 . 39 . 58 , 140, 14 1 , 142, 143) (Cristovomer-33.58. 141 )
(Crocethia-289 , 297 ,312, 325, 337,342) ( Crocidura-350 , 393 . 41 2, 415 . 421 . 422. 425 . 426 , 427 , 429 . 430 . 432, 435 , 439, 448)
(Crocopus-213,214,217) (Crocothemls-332) (Crocuta-438) (Crossophthalmus-202) (Crossopus-439) (Crotalus-179)
(Crotophaga-235,237) (Crotophopeltis-175) ( Cr ymophllus-300 ) (Cryptobranchus-162) ( Cr yptoglaux-269 )
(Cryptomys-432) (Cr ypturus-21 3 . 305 ) (Crystallias-32) (Ctenocephal us-41 0,4 18. 432) ( Ctenopharyngodon-46 )
(Ctenopsyllus-419) (Ctenosaura-181 ) (Cuculus-222 , 235 , 308 ) (Curacus-271 ) (Cursorius-237.251 ,262)
(Cyanocitta-313) (Cyanops-203) (Cyblum-128, 132,136) (Cyclocypr is- 326)
(Cyclops-38. 51, 58. 140. 14 1,142, 143. 144. 16 1.170 .174. 176. 185 .186. 187 .188. 289. 294. 295. 304. 309. 31 0.31 1.322. 326, 328.
354,358,359,360) (Cyclopsittacus-203. 21 3.290) (Cyclopterus-38,45) ( Cygnopsis-304 )
(Cygnus-227.288.289.295.297.318.325.326.348.349) (Cylindrosteus-140) (Cymogaster-125) (Cynias-66,68,82.86)
( Cynlctis-4 14 ) ( Cynomys-344 ) (Cynosclon-1 1 3 . 1 15. 1 30. 1 31 ) (Cypridepsis-293) (Cyprlnus-20, 21 .29.46. 187)
(Cypris-289. 305. 31 0.322. 326) (Cypseloides-236.238) (Cypselus-231 .236.270) ( Cystophora-358 )

(Dacelo-267 ) (Dacnis-273) (Dactylopterus-44 ) (Dafila-288.297.319) (Dallia-185. 353) (Daption-197)
(Dasyatis-65.66.69.73.77.80.82.83.84.85.86.89.95.96.98. 118. 131 . 132. 133.134. 135. 136. 137) (Dasycottus-126)
( Dasyproc ta-4 42 ) (Dasypus-395 ) (Dasyurus-441 ) (Del phlnus-352 . 353 , 366 , 368 ) (Demansia- 175 )
(Dendrocitta-227. 231 .232.271 .292.316) (Dendrocoptus-278 ) (Dendrocopus-214) (Dendrocygna-162,246,292,
294,319,334) (Dendrolaphis-413) (Dendromus-427 . 444 ) (Dendropus-180) (Dendrornls-254 ) (Denisonia-173)
(Dermestes-394 ,419) (Desmognathus-162, 165) (Diaeum-220) (Diagramma-13, 16, 129) (Diaptomus-58 . 142,
143,185,186,294,297,304,310,322,325,326,328,346,354) (Dicheros-214) (Dicholophus-286 ) (Picotyles-382)
(Dicromorpha-260) (Dlerostonyx-373 ) (Didelphis-353 . 360. 365 . 392 . 394 . 395 ) (Dlkerogammarus-9 ) (Dinotopterus-141 )
(Dlomedea-1 93 .194.195.197) (Dlpodomys-394 . 447 ) (Dipsadomorphus-69 . 173 ) (Dipus-394) (Discoderus-219)
(Disphelldus-180) (Pol icholophus-224 ) (Dorus-1 31 ) (Drepane- 1 32 ) ( Promaeus-209 ) (Dromaius-21 3)
(Dryobates-214.236.261 ) (Dr yoscopus-273 ) (Ducula-215.221 ) (Pytes-296)

(Echeneis- 113) (Echidna-392) (Echinocotyle-325) (Echiostoma-44) (Eclectus-215.216.222) (Ectopistes-202)
(Egernia-1 81 ) (Elanoides-313) (Elaphe- 1 75 . 179 . 182 . 363 ■ 364 ) (Elaps-175) (Elapsoidea-175) (Elephantulus-394 )
(Elephas-372) (Eliomys-432) ( Ember lza-2 16. 235. 237. 251 .271 .273) (Enhydra-358) (Enneoc tonus-235) (Enophrys-44 )
(Entomyza-220) (Ephestia-394.419) (Epimys-390.404) (Epinephelus-126, 128) (Epischura-142)
(Epteslcus-429. 431 .432.433) (Equula-131) (Equus-372. 381 ) (Erethizon-382 . 383 , 438 . 445) (Er ignathus-353)
(Erlmyzon-22 . 140) (Er inaceus-359 . 360 . 394 . 395 . 407 . 427 . 44 1 ) (Erionetta-293.299.323) (Erlsmatura-292.304)
(Er lthacus-252 ) (Erolia-232 , 235 ,238,239,244, 257 ,2 60,261, 262 , 289 ,290,297,299,301,312) (Erythr ina-322 )
(Erythrinus-131) (Erythropus-237) (Eryx-179. 181 ) (Esox-33 . 38. 45 , 57 , 58 . 140 , 14 1 , 142, 1 87 , 354 ) (Estr ilda-308)
(Etmopterus-121 ) (Eucichla-232) (Eucitharus-44 ) (Eucyclops-58. 140, 143,294) (Eudromius-236) (Eudyptes-193. 196)
(Eulopa-4 12) (Eumeces- 179 . 180) (Eumetopias-353 . 357 . 358 , 367 ) (Eunectes-173,437) (Eupagur us-73 . 1 17)
(Eupetomena-272. 320) (Euplocamus-220) (Eupodotls-214,224,225) (Eupomotls-45 . 51 . 140) (Euponera-221 )

176

THE TEXAS JOURNAL OF SCIENCE

(Eurocephal us-271 ) ( Eurycercus-

(Evotomys-385,418,426,440)

CESTODA, cont.

( Eurystomus-326 ) (Eurytemora-44 ) (Eutropiichthys-145) (Euxerus~447)

(Faloo-187 .189, 191,223,237,274,344,345) (Farancia-175. 179) ( Fel ls-355 . 360, 361 , 410 . 41 2. 41 3 . 41 4 . 415 , 437 .
439,440,442,4*13,4410 ( Fennecus-4 1 A ) ( Fiber-408, 4A9) (Fistularia-74) (Florida-232,233)
(Francolinus-212,213,215,216,232.238,260,310) (Fratercula-193.263) (Fregata- 193. 194. 195.268) (Fridericia-300)
(Fringilla-231, 262,27 1,278,301,315) (Fulica-247 , 291 . 292, 293.297 , 310. 31 1 . 326 , 329, 332)
(Fuligula-288, 289, 290, 292, 297, 300, 302, 305, 31 0,311 ,312,318,327,334) (Fulmarus-193. 194,299) (Fundulus-55)
( Furnar ius-24 1 )

(Gaeko-1 80 ) (Gadus-21 . 38 , 40 , 41 . 44 . 45 . 115 , 126 . 135 . 1 36 . 143 . 358) (Galaehr ysia-300 . 31 3) (Galeichthys-1 15.
122) (Ga^eocerdo-65,66,67,77,87,88,90, 101 , 113, 128, 131 , 135) (Galeopithecus-375,405) (Galeorhinus-68,80. 127)
(Galerella-394) (Galerita-214,217,231 ,262,271,273,309) (Galeus-65 , 68 ,69 , 77 , 80 , 82 , 85 , 86 , 1 1 3, 1 15 , 1 33)
(Galictis-444) (Gall inago-233 . 236 , 238 , 245 . 25 1 , 252 , 253 . 260 , 26 1 , 289 , 291 . 298 , 299 , 301 . 31 3 , 344 )
(Gal linula-238. 247, 26 1.292. 293) (Gallirex-215.217) (Gallus-1 98 , 208 , 209 ,210, 21 1 . 212, 21 3 . 21 4 , 21 6 , 217 , 21 8.
219,220,221,222,233,234,260,275,292,298,303,320,325,326) (Galumna-365 , 372 . 376 , 380, 381 , 385 , 400) (Gammarus-9 .
33,34,61 ,246,300,309,310,311,322,421,432) (Gar rul ax-21 9 , 237 , 25 1 , 316) (Garr ulus-23 1 . 236 . 263, 301 , 309 , 315 , 322)
(Garzetta-222,233) (Gasterodonta-208) (Gasterosteus-45 , 46 , 47 . 58 , 140, 142, 185, 186, 187, 188) (Gasteroteus-140)
(Gatto-365) (Gavia-1 86 , 253 , 258 , 286 , 296 , 31 1 ) (Gazella-380 . 398 , 399 ) (Gecinus-209, 21 3, 21 4 , 221 , 222, 261 , 308)
(Genetta-396,413.414,415,439,440,443,444) (Genypterus-59 ) (Geoclchla-242) (Geomys-374 . 376 , 382. 385 )
(Geopelia-209 ,216) (Georhynchus-439) (Geosclurus-447) (Geotr upes-260 , 309 ) (Geranospizias-274)
(Gerbillus-388,444,445,447) (Ger rosaur us- 1 80 , 1 81 ) (Gila-23,142) (Ginglymostoma-67 . 87 . 1 1 3 , 1 1 8 , 1 35 . 1 36 , 137)
(Giraffa-380 ) (Gladiunculus-45 ) (Glanidium-69 ) (Glareola-237 . 261 . 325 , 326. 327 ) (Glaridichthys-144 )
(Glaucomas- 37 4) (Glis-420) (Globicephal us-353 . 355, 369) (Globlcera-213) (Glomer is-429 ) (Glottis-260)
(Glyciphagus-447) (G 1 yptocephal us- 1 26 ) (Gnathocerus-394 ) (Gnathopogon-44 , 142.250) (Gobio-20 . 23 . 58 . 1 87 )
(Gobius-44, 105) (Golunda-389 ) (Gongylus-4 1 3 . 41 4 ) (Gonlodiscus-412) (Gorilla-372) (Gorsakius-254 )
(Goura-214,215,216) (Gracula-273) (Gracupica-322) (Grammomys-4 1 8 . 444 ) (Guira-237) (Guttera-216,221 ,277)
(Gymnodactylus-4 1 4 ) (Gymnostinops-238, 315) (Gymnura-85) (Gy paetus-237 , 345 ) (Gypagus-247)
(Gyps-201,205,237,263)

(Haematopus-226,253,289,300,312,322,329,337,346) (Haemulon-132) (Halcyon-262) (Hal iaetus-222 , 244 ,
250,345,346) (Hallchelys-1 12) ( Ha loporphyr us-38 ) (Hamadryas-375 ) (Hapale-395 ) (Har eld a- 199, 246, 251 )
(Harpactes-291 ) (Harpalus-219) (Harpipr ion-231 .250) (Harpodon-1 32) (Hel icops- 176 ) (Hel ioperca-58 . 140)
(Helix-212.214) (Helodromas-238 , 252 ) (Hemidactylus-179 . 180 . 41 3, 41 4 , 415 ) (Hemigaleus-67,91 , 132)
(Hemignathus-315) (Hemigrapsus-85,118) (Hemitripterus-44) ( Heptane hus- 66 , 126,127,134)
(Herodias-231 .233,235,240,243) (Her pestes-210, 216 , 361 , 394 , 404 ) (Herpobdella-252, 306) (Heterobranchus-56, 150)

(Heterodontus-66 , 84 )
(Hieraetus-345)
(Hippoglossus-55 ,137)
(Hirundapus-236)
(Histrionicus- 311)

( He terohyr ax-372, 389 ,390) (Hexagraminos-44 ) (Hexanchus-66 , 67 , 69 , 86 , 112, 126, 127)

(Hlmantopus-201 .209,226.231 ,262,265,288,290,291,333,335,342,343) (Hiodon-45 , 47 )

(Hippolyte-62 ) (Hippopotainus-380 , 381 ) (Hipposideros-438) (Hippotragus-381 , 398,399)
(Hirundella-271 ) (Hir undo-234, 236, 237. 238, 253, 270. 271 ,308,346) (Histiophorus-46 )

(Holopterus-252 ) (Holorhinus-73 ,84,85) (Homalopsis-175)

(Hoplopterus-233.237.240.252.337.342) (Hoplox ypterus-226 . 232 . 342 ) (Houbara-223,225.272) (Huro-45 , 142)

(Hyaena-438) (Hyas-1 17) (Hybomys-427 .428) (Hyborhynchus-24 ) (Hydrarga-68 ) (Hyd roc he lid on- 193. 237, 24 6)

(Hydrochoerus-382 ) (Hydrolagus-17) (Hydromedusa-175) (Hydroprogne-290 , 300) (Hydropsalis-248) (Hydrurga-355)
(Hyla-1 62, 164 , 165 . 166 . 360,401 ) (Hylocichla-309) (Hymenophysa-45 ) (Hypentelium-22) (Hype rood on- 350, 368)

(Hyplophus-67) (Hypochera-297 ) (Hypopr ion-66 . 69 , 89 . 1 15 , 1 16) (Hypotr iorchls-237 ) (Hyrax-404)

(Iartes-365) (Ibex-380) (Ibis-250,251.310) (Ichneumia-414) (Ictalur us-85, 140, 144,146) (Icterus-271 )
(Ictinia-274) ( Ictiobus-21 , 22 , 23 , 25 , 26 , 140) (Idus-143) (Iguana-181 ) (Ilisha-132) (Inachus-1 17)
(Iridomyrmex-212) (Isur us-67, 69 ,86. 110, 122) (Ixobrychus-221 .243) (Ixoreus-300)

(Jaculus-445) ( Julus-4 12) (Jj

-261 )

(Kerivoula-432) (Ketupa-325) (Kogia-68) (Kryx-398)

(Labeo-145) (Labidethes-140) (Labrax-46) (Labrus-44 , 47 ) (Lacerta- 179 , 180. 1 91 . 21 8, 4 1 0, 41 3 , 415 )
(Laches is- 173 ■ 174 ) (Laemargus-67.76) (Lagenorhynchus-368 , 437 . 44 9) (Lag id ium-375 . 376 . 384 , 386 )
(Lagopus-208 .212.216.218.222.278.279, 300. 325 .330) (Lagorchestes-386 , 401 ) (Lagurus-447 ) (Lakoeta-20)
(Lamna-65, 68.69.71 .86. 103. 109. 114) (Lampris-78) (Lanius-270.273,274,278.279.309) (Larus-184 , 185, 186,
187,188, 192, 193, 194,226,232,233,235,237,246,251 ,252 , 253 , 263 , 289 , 290 , 294 , 295 , 297 , 299 , 300 , 301 , 304 , 31 0, 319, 324,
325,353) (Lasionycteris-433) (Lasiurus-393 ) (Lateolabrax-44 , 46) (Latus-354) (Lebia-260) (Lebistes-51 )
(Lechesls-176) (Leggada-427) (Leiobatus-62 , 83 ) (Le iolopisma-1 65 ) (Leiopoa-215) (Lemmus-373 , 4 1 9 , 438 , 439 )
(Lemniscomys-444) (Lemur-391 . 445 ) (Leo-437) (Lepibema-58 , 140, 142) (Lepidopus- 1 1 0 , 1 22 ) ( Lepidorhinus- 1 26)
(Lepisosteus- 140,141 .142.143) (Lepomis-45 . 58 , 140,142,143) (Leptail ur us-4 1 5 ) (Leptocephalus- 1 35 )
(Leptocottus-44 . 46 ) ( Leptocyc lops- 360 ) (Leptodactylus-1 62 . 165, 359) (Leptodira-175) (Leptodora-140)
(Leptonychotes-354.355.357) (Leptops-144) (Leptopsylla-419) (Lepus-345 . 373 , 376 , 383, 385 , 390 , 406 , 445)
(Lerwa-213) (Lethr inus-128) (Leucichthys-33 . 34 . 38 . 45 . 58 . 141 , 1 43 . 185 , 353 ) (Leuciscus-20. 21 ,141,143,187)
(Llacarus-372 ) (Lialis-176) (Lichia-135) (Lichtenstelniplc-220) (Limanda-44 , 45 , 126) (Limax-208 , 21 8 , 235 )
(Llmicola-233) (Limnocryptes-235.252.261 . 300. 301 ) (Limnodrilus-20,22,23) (Limnodromus-344 )

INDEX: SYSTEM A HELMINTH UM

111

CESTODA, cont.

(Umonites-232,238,260,261 ,289 , 299 , 312, 3<t5) (yjmosa-226,227,232,265,289,297,3'*2) (Liolepis-1 80)
(Liophis-173, 176) (Liparis-32) (Lissotls-223.22tt.225) (Lobipes-300.326) (Loblpluv ia-231* . 245 . 26 1 . 265 . 337 )
( Lob i van ell us-252, 262, 266, 342) (Lobodon-355 ) (Lobotes-131 ) (Locustella-262 . 315) (Loligo-80) (Lonchura-308)
(Lophaethyla-296) (Lophaetus-345 ) (Lophius-44, 46 , 55 , 1 1 1 , 1 1 3 , 1 26 , 1 27 , 1 31* . 1 35 , 1 36 , 1 37 ) (Lophoceros-227.269,270)
(Lophodytes- 188,291 ) (Lophophor us-210) (Lophopsetta-44 , 131 ) (Lophorus-200) (Lophotibis-226)
(Lophotis-22tt,272,Htl9) (Lorlus-203.215.216) (Lota-33 , 38 , 58 . 85 , 126 , 1 36 , 1 »t 1 , 192, 14 3, 359) (Lotella-126)
(Loxla-271 ) (Loxops-271 ) (Lucioperoa-tl5,57 . 187) (Lullula-278) (Lumbrlculus-238. 300) (Lutnbricus-233)
(Lurocal ls-252 ) (Lusclnla-233. 323) (Lut janus-129, 132, 136) (Lutra-351 .361 ) (Lutreola-36H ) (Lybius-317)
(Lycaon-360,438,439,442) (Lycodes-38) (Lycodon-180) (Lygosotna-1 80) (Lymnocryptes-2 38 , 253 . 299 . 327 ) (Lynx-360.
36*1 ,365, 438, 939, 440,444) (Lyrur us-21 6, 21 8, 21 9. 222. 260, 278. 279, 330)

(Habuya- 179) ( Macacus-360 , 445 ) (Machetes-235 , 238 , 260 , 261 , 300 ) ( Hacrocorax-221 ) ( Macrocyclops- 1 HO , 1 6 1 )
( Macronectes- 1 95 ) (Macrones-12,20, 145) ( Macropter yx-254 , 269 , 270 ) (Maeropus-361 . 378. 379, 386,
396,401 ,406,408,438) ( Macrorhlnus-356 ) (Macroscelldes-432) (Haena-137) (Hajaqueus-193) ( Halapterus-1 44 )
(Halapterurus-135,145) (Mallotus-39) (Halopterus-144) ( Mai polon- 1 79 , 1 80 . 4 1 3 ) (Malurus-263 )
(Manis-401 ,403,406,427) ( Manucod ia-221 , 273 ) (Mareca-288 , 289 , 290 , 292 , 304 , 31 8) (Margaritifera-99)
(Marila-289, 291 ,293,304) ( Marmosa-392 , 394 , 395 ) (Marmota-376 , 377 . 385 . 447 ) ( Martes-364 , 433 , 440)
(Mastoeembalus-56 ) (Mastomys-405.427.428,448) (Mazama-380) (Megalalma-203) (Megaloperdix-303) (Megalotis-414)
(Megapodius-232.248) (Melanerpes-213,222,308) (Melanitta-309 , 310, 31 1 , 328 . 329 ) (Melanobucco-217 , 220)
(Melanogrammus-40) (Melanoplus-260.275) ( Meleagr is-209 . 21 4 , 217 . 21 8. 220, 221 , 260 . 275 , 290 . 298 , 320 , 326)
(Meles-364 , 393 , 440 , 446 ) (Helierax-270) (Heliornls-262) (Melitophagus-273) (Melivora-364) (Melizophilus-279)
(Melophus-268,316) ( Memtnus-385 ) (Menachilus-143) (Menticirrus-44 ) (Mephitis-364 , 365 . 396 , 437 )
(Merganser-311.318) ( Mergus- 1 84 , 185 . 186 , 187 , 1 88 , 226 , 284 , 287 , 289 , 290 , 303 . 304 , 305 , 31 1 , 312)
(Mer iones-394 , 395 , 406 ,418,419, 427 ,434,443,445.447,448) ( Merlangus- 1 36 . 142) (Merluccius-40,
41,44,55,126,127,134) ( Merops-246 , 273 ) (Merula-309) (Mesembrinibis-231 ) (Mesenehytraeus-300)
(Mesocrleetus-394,418,434,447) ( Mesocyclops- 1 6 1 ,163,174,360) (Mesoplodon-366) ( Metachirops-392 )
( Me t ac hi r us-392, 394) (Metamysis-9 ) (Metopriana-288 ) (Microcarbo-232 . 250 . 31 0 . 324 ) (Hicrocyclops-174 )
(Mlcrogad us-40, 135) (Mlcrohyla-359 , 360) (Micrometrus-125) (Microinys-432) (Micropogon-122, 131,132)
(Micropter us-45, 58, 140, 141 , 142, 143. 187) ( Micropus-234 , 249 . 276 ) ( Microsarcops-252 ) ( Microtus-345 . 360 .

373.374.383.384.385.412.418.419.426.427.432.433.437.440.441.442.447) (Migapodus-251 ) (Milax-203)
(Milvus-1 91 ,210,222,223.224.237.238.250,252.253.257.262.313.345) (Miniopterus-429.430) (Mino-221 )
( Mi sg urn us-20, 23) (Mogurnda-105, 142) (Mola-46 , 66 , 1 16 , 122, 1 35 ) (Molossus-431 ) (Molothrus-273)
(Molpastea-203 ,221) (Molybdophanes-324) (Monticola-231 ) ( Morel ia- 170 ) ( Mormyr us-27 ) ( Motac il 1 a-235 ,
251 ,262,271 ,272,273,346) ( Motella-38 , 40 , 46 , 47 ) ( Moxostoma-21 ) ( Mozama-397 ) (Mugil-128. 136) (Mullus-44 , 47 , 1 36 )
(Huraena-44,45, 135) (Muraenesox-48, 1 15) (Mus-345 . 361 , 364 , 389 , 390 , 394 , 395 , 404 , 405 ,

406.418.419.420.421.426.427.432.433.437.438.440.441.447) ( Musca-2 1 2 . 21 8 . 260 ) ( Muscicapa-235 , 238 . 27 1 . 27 3 .
308,309,315) ( Mustela-364 , 437 , 438 , 439 . 440 . 443 ) (Mustelus-17 , 62 , 65 , 66 , 67 , 68 , 69 ,77 , 83 , 85 , 86 , 89 , 90 , 1 1 3 , 1 16 ,
117,120,125,127,131,135,138) (Hycetus-375) (Mycteroperca-128. 137) (Myiagra-310) (Myliobatis-69.73.
74,76,98,103,137) (Mylobdophanes-295) ( Myonax- 1 91 . 394 ) ( Myopotamus-427 . 444 ) (Myothera-252)
(Myotis-424, 429. 431 .432.434) (Myoxocephalus-33 . 34 . 38 . 44 ) ( Myoxus-420 , 432 ) ( Myrmecophaga-395 ) (Myzomela-221 )

(Naja-1 70. 17 1,175. 176. 180. 221) (Namls-407) (Narcine-69 , 84 . 157 ) (Narke-84 , 103 ) (Nassa-61 ,62)
(Nasua- 375, 393,437) (Natr lx-174 . 175 . 1 80 . 360. 415) (Necomis-22) (Nectarinia-271 ) (Necturus- 1 62 )
( N eg apr ion-65 , 84 , 115) (Nemachilus-20, 187) (Nemorhaedus-380 ) (Neobythites- 1 1 3 ) (Neofiber-376,385)
(Neomenis-353 ) (Neowys-4 12 . 4 1 9. 420 . 421 . 423 . 429. 430 . 432 ) (Neophoca-353 ) (Neophron-199) (Neotis-224,226)
(Neotoma-374) (Nerophls-44 , 58 ) (Neseatus-218) (Nesocla-215,404.406,440) (Nestor-204) (Nethium-246 )
(Netta-288, 291 .293.322.334) (Nettapus-304 ) (Nettion-287 , 292, 300, 304 , 31 8, 325 , 334 ) (Nettium-246 )
(Nomadacrls-277 ) (Notaspis-374) (Notechls-173) (Notemlgonus-24 ) (Nothura-218) (Notiopsar-27 1 ) (Notophoyx-240 )
(Notorhynchus-127. 129) (Notropis-21 ,58. 140, 142, 144.187) ( Noturus- 1 4 0 ) ( Nuc i fr aga-23 1 , 309)
(Numenius-226, 232. 237. 251 .252.253.262.299.327) (Numida-206.209.210.212,213.214,216,
217,218,220,221,257,265,275,277,303,320,404) (Nyctalus-429) (Nyctea-269) (Nyctibius-233.252)
(Nycticebus-394 .407) (Nycticeius-433) (Nycticorax-193. 194.240.244.249,255.258) (Nyctiprogne-323)
(Nyctophil us-395 ) (Nymenius-327) (Nyroca-236 . 246 . 284 . 288 . 289 , 290 . 291 . 292 . 293 . 295 . 297 . 300. 304 , 305 , 309 , 310 .
311,312,317,319,324,334) (Nythotrephes-140)

(Ochotona-387) (Ochthodromus-338) (Oenerodrilus-233 ) (Odobaenus-353) (Qdocoileus-380, 381 ,397,444)
(Odontaspis-66. 135) (0ediceros-6l ) (Oed icnemus-232 , 247 , 252 , 263 . 265 . 31 0, 337 ) (Qena-202, 315, 319, 323)
(Oenopopelia-202,216) (Oestrelata-192. 193) (Ogmorhinus-354 . 355 ) (Oidemia-246 , 290 , 291 , 292 , 309 ,
310,311,322,326,334) (Oligocottus-125) (01 igopl ites-44 ) (Ommastrephes-66 .113) (Ommatophoca-355 . 357 )
(Qmmatostrephes-71 ) (Oncoides-439) (Oncorhynchus-38 . 39 . 40 . 58 . 66 . 67 , 140 , 145 , 184,354,355)
(Ondatra-374. 423. 427 .440. 44 3) (Onthophagus-212) (Onychogale-379 . 386 ) (Ophiocephal us-56 . 1 45 ) (Ophiodon- 11 3 )
(0phisurus-1 33 ) (Qpisthocomus-251 ) (Qpladelus-140) (Opsariichthys-46) (Orca-350) (Qrchestia-304 ) (Ore inus-437 )
(Oreotragus-398 ) (Or ibatula-380 ) (Or iolus-214 , 220, 221 . 23 1 . 236 . 262, 272. 273 . 275 . 279 . 309 . 315 . 322) (Qrnis-202)
(Orthagoriscus-59 . 122) (Qrthophagus-320) (Or tygometr a-333 ) (Ortyx-278) (Or yctolagus-373 . 376 , 383 , 385 , 430)
(Oryx-398) (Osmerus-38 , 39 . 46 . 58 . 141 . 142, 185) (Ostinops-279. 315) (Otar ia-68 . 355 )
(0tis-206, 223, 224, 225, 237, 260, 270, 302, 303, 309, 325, 326) (Otocorys-253 ) (Qtocyon-360 . 41 1 ) (Otomela-270)
(Otomys-385. 389.444.445) (Otus-263 . 269 . 449 ) (Ourebia-398 ) (Ovibos-380)
(Ovis-380, 381. 382. 397. 398. 399. 400. 401. 439) (Oxyechus-338. 342) (Oxyrhina-1 13) (Oxytelus-323 ) (Oxyura-288 , 298)

178

THE TEXAS JOURNAL OF SCIENCE

CESTODA, cont.

(Pachycepha la-272, 279) (Pachygrapsus- 1 18) (Pagellus-122) (Pagodroma- 1 93 ) ( Pagol la-392 ) ( Pagrosomus- 1 26)
(Palaeornls-209) (Palinurlchthys-9M) ( Paloma-2 1 6 ) ( Palombus-21 8 ) ( Palorus- 419) ( Pandion-250 . 395)
(Panthera-938) (Pa£io-938) (Paraealanus-126) (Paraeynictis-909) (Par ad isea-273 )
( Par adoxur us- 170, 909, 9 15, 939,990) (Paralichthys-99 , 131 , 135) (Parameles-392) (Parasllurus-192, 199,195)
(Pariparus-315) (Parisomius-262) (Paroxya-275) (Par us-253 . 262, 271 , 272. 278. 301 . 315 , 326 ) (Passer-219,
216,218,231,236,251,253,261,262,271,308,315) (Passerella-253,300) (Pavo-21 0,21 1 .212, 21 3, 219 , 217 . 222 , 320)
( Pa voncel la-260, 322) (Pedioecetes-260.278) (Pedlotragus-381 ) (Pelamys-1 1 1 , 126, 128) (Pelecanus-189,

187.199.196.250.286.313.325) (Pelecus-20) (Pel idna-226, 236, 238, 252, 289, 298, 299, 300) (Pelorlbates-380)
(Penelope-216) (Penthetriopsls-279) ( Pepr ilus- 128) ( Perameles- 909,933) (Peramys-392) ( Perea- 3 3 ,
38,95,57,58,190,191,1112,187,359) (Percina-95, 193) (Percopsls-95,58) (Percottus-105) (Perdlx-208,

209.219.215.218.219.220.222.260.278.308.320.325) ( Per lops-9 1 3 ) (Periparus-271 ) (Perlplaneta-905) (Pernls-395)
(Perognathus-399,997) (Peromyscus-269 , 369 , 365 . 9 12 . 992 . 999 , Hil7) (Petaur ista-375) (Petroclnela-315)
(Petroclidurus-313) (Petrodromus-928) (Petrogale-901 ) ( Petromyzon-38 , 58) (Phacoehoerus-382 . 389 )
(Phaeochroa-199) ( Phalacrocorax- 1 85 , 1 87 . 1 88 , 250 , 309 . 31 0 . 31 3 . 329 ) (Phalanger-375) (Phalanglsta-375)
(Phalaropsls-235) (Phalaropus-252\298,326) (Phascolarctos-937) (Phascolarctus-375) (Phascolomys-381 ,386)
(Phasianus-198.219,215,216.218,260.278,308.320) (Pheidole-212,219) (Phenacomys-373) (Phllaete-295.297)
(Philander-399) (Philemon-220) (Philohela-238,239) (Philomachus-260, 261 ,297,298,300,312)
(Phoca-353, 357 , 358 , 366 , 367 , 369 ) (Phocaenoides-366) (Phoebetria-199, 195. 197) (Phoeniconaias-291 )
(Phoeni copter us-2 85 ,298 , 299 , 309 , 325, 331,338,339) (Phoenicurus-262,273) (Phoxinus-187) (Phraetocephalus-199,
155) (Phrynorhombus-9H ) ( Phrynosoma- 178) (Phrynosomatis-178) (Phycis-136) (Phyllorhlza-17)
(Phyl Iosco pus-2 35 , 273 ) (Physa-208) (Physeter-350, 367) (Pica-221,231,236,263,309,315,322,359) (Plcolaptes-259)
(Picus-2 19, 216, 221 ,233,236,297,253,261 ,308) (Pllherodlas-293) (Pilumnus-1 18) (Pimelodus-191 . 153,156)
(Pimeloldus-151) (Plnicola-267 ) (Pinopslttacus-215) (Pionus-215) (Pipistrellus-929 . 933) (Piraeeba-155)
(Pirarara-155) (Pirlnampus-198. 153) (Pisobia-299 , 300) (Pitta-265) (Pltuophis-180) (Pitymys-939 , 997)
(Pityophis-179) (Planar la-190) (Planesticus-261 ,290,323) (Platalea-233 . 292, 250 , 310, 325) (Platibis-325)
(Platycephalus-1 15) (Platycercus-209.210.21 1 .215,231) (Platydactylus-177) (Platysqualus-129) (Platystoma-152)
(Platystomatichth-1 51 .152,156) (Plecoglossus-192) (Plecotus-929,939) (Plegadls-292 , 27 1 . 390) (Plethodon-162)
(Pleuronectes-39, 35. 99.97.111, 11 3. 126. 137) (Ploceus-268 . 316) (Plodia-399) (Pluvialis-297 ) (Podabrus-219)
(Podager-220,253.323) (Podargus-255) (Podica-292) (Pod jeeps- 1 86 . 187 , 1 88 . 192 , 296 . 286 , 296 , 316 , 332 , 333 , 390, 39 1 )
( Pod ilymbus-307, 332.333) ( Pod iotr agus-398 ) ( Poec ilogale-937 ) (Poecilonetta-319,326) (Poephagus-381 )
(Pogonias-1 30) (Pollachius-90,99) (Pollasiomys-9M7) ( Polyborus-257 ) (Polygyra-208) (Polyodon-93)
(Polypedates-165) (Polypterus-55. 192, 193) (Polysticta-293 , 301 ) (Polytells-210) (Pomatomus-55, 115, 129, 131 )
(Pomatorhynchus-273) (Pomolobus-99 , 131 ) (Pomoxis-190, 192) (Pontogammarus-33) (Pontoporeia-33 ) (Porogynia-277)
(Poronotus-99, 128) (Portunus-1 17) (Porzana-326, 333) (Pratincola-262) (Prenolepls-212) (Prinace- 159 )
(Priocella-193) (Pr iodontes-395 ) ( Priofinus- 1 97 ) (Pr ionace-67 , 69 ,78 , 87 , 89 . 109 , 1 12 , 1 1 3 , 159 ) (Pr ionailurus-369 )
(Prionodon-130) (Pr lonotus-1 35 ) (Prlstipoma-1 32) (Prlstls-69,87,95.97, 129, 132) (Pristiurus- 1 39 )
(Procavia-372 , 389 , 390) (Procell aria- 192, 193,199,195,197) (Procyon-359 , 355 . 369 , 393) (Progne-271 ) (Promomys-927)
(Promoxls-190) (Prosopium-58, 191 ) (Protemnodon-386) (Protoschelobates-380 , 383) (Prunella-251 ,262)
(Psammodromus-179) (Psammodynastes-176) (Psammophis-179 , 180,181,919) (Psenopsis-50,52,53) (Pseudacris- 1 65 )
( Pseud ageniosus-69) (Pseudaspius-175 ) (Pseudechis-173) (Pseudeudotropius-197) (Pseudocal anus- 126)
(Pseudochirus-375 , 389 ) (Pseud odoras-191 ) (Pseudogenlosus-198) (Pseud ogobio-23) (Pseudogyps-237)
(Pseudopimelodus-155) (Pseudoplatystoma-199, 151 , 156) (Pseudopleuronect-39 ,35.95) (Pseudorhombus-1 13)
(Pseudotantalus-292) (Pseudothemis-332) (Pseudotr iacis-1 10, 126) (Pseudotriton-162) (Pslttacula-210,21 1,216)
(Psittacus-205,215.217.218) (Psophia-232) (Pternistes-21 3) (Pteroaetus-395)
(Pterocles-210, 213, 215, 216. 22 1.313) (Pteroclidurus-213,218) ( Pterocnemla-203 ) (Pterodroma- 1 92 )
(Pteroglossus-252) ( Pteroplatea-69 . 89 , 85 . 95 . 96 . 97 . 1 32) (Pteropus-939 ) (Pterostichus-219) (Ptilorhis-273)
(Ptllotis-261 ,315) (Ptistes-210) (Ptyas-165) (Ptychochellus-39, 191 , 192) (Ptyodaetylus-919)
(Puffinus- 192, 193, 199,233) (Pulex-910.it18.932) (Pulvianus-337) (Punctoribates-369) (Putor ius-938. 939, 990, H93 )
(Pycnonotus-209 , 323) ( Pycnotus-220 ) (Pygoscel is- 193, 199, 195,259) (Pygosteus-191 ,188) (Pyralis-918)
(Pyromelana-220,279,297) ( Pyr rhur a-1 99 ) (Python-170.180) (Pytilia-213)

(Querquedula-260, 288, 290 ,291 ,293.297.301 .317.319.326.339.399) (Quinquarius-39) (Qulscalus-272)

(Raja-99,61,62,65,66,67,68,69,70,72,73, 79, 79, 80, 82, 83, 89, 85, 86, 88, 103, 112, 113, 117, 139, 137)
(Rallus-238.306) (Ramenis-9 1 9 ) (Rana-162 , 163 , 169 , 165. 166 . 179 , 175 , 360) (Rangifer-376 , 380, 381 , 398) (Raniceps-96)
(Rappia-165) (Rattus-375 , 399 , 909 , 905 , 906 , 907 , 91 2, 91 8, 91 9, 920, 926 , 927 , 931 , 932, 933 , 939 , 990, 99« , 9H7 , 998)
( Recur virostra-200. 209, 233. 290. 291 .305.310.335.338) (Redunca-381 ,382) (Remora-115,137) (Rhabdomys-906,9H8)
(Rhabdo phis- 175. 176) (Rhagadus-219) ( Rhamdia-99 , 1 46 ) ( Rhamphastus-209 ) (Rhea-200.211) ( Rhina-87 , 1 1 8)
(Rhinobatus-73,83.89,85. 97,98, 157.158) (Rhinoceros-372) (Rhinochetus-238) (Rhinolophus-932) (Rhinoplagusia-79)
( Rh i no pt era-62, 72,73 , 79 ,98,103.139) (Rhodeus-20) (Rhomboidichthys-9it,97) (Rhombomys-919,99 3,999,9118)
(Rhombus-90, HU. H7) ( Rhyac lphilus-289 ) (Rhyacophilus-238,260) (Rhynchaea-252, 310, 392) (Rhynchobatis-69,70)
(Rhynchobat us-62. 66 .78,87.97.98.100.103.118. 129,133,135,136,137,157,158) (Rhyncholus-220) (Rhynchops-253)
(Rhynchotus-218) (Rlchardsonius-192) (Riparia-239 ,238) (Rissa-185, 186 , 1 87 , 192 , 193,237 , 251 , 253, 300, 301 )
(Rita-192) ( R 1 v ulogammar us-33 ) ( Rostr atula-1 99 , 23 1 . 232 . 252 . 306 , 31 1.315.327) ( Rowettia-238 ) ( Runambulus- 9 1 9 )
(Rupicapra-380) ( Rupicola-227 , 323 ) (Rupornis-279) (Rutlcllla-238,262, 396) (Rutllus-20,21 ,187)

(Salamandra-169) (Salmo-33 . 39 . 38 . 39 . 90 , 9U , 95 , 97 . 57 . 139 . 136 . 137 , 190 , 19 1 , 192,193,176,189,187,359)
(Salpinctes-270) (Salvelinus-33 , 39 , 38 . 39 . 95 , 98 . 58 , 190,192,193,189,187,188) (Sarcidiornls-296)
(Sarclophorus-239) (Sarcogrammus-266 ) (Sarcophilus-991 ) (Sarcorhamphus-193) (Sardinella-89 )

INDEX: SYSTEMA HELMINTHUM

179

CESTODA, cont.

(Saurlda-45.46,47.48) (Saxicola-238.262,263,273,326) (Scalopus-419,429) (Scapanus-427) (Scaphiopus- 1 65 . 166)
(Seardlnius-20, 187) (Scaurus-4 1 8) (Seelophorus-180) (Schelor ibates-364 . 365 . 372 . 374 , 380 . 381 , 385 . 400)
(Schilbeodes-140) (Schizopygopsis-187) (Schizothorax-20) (Sciaena-47 . 131,137) (Scincus-414 )
(Sclur us-350, 35 1,373. 383, *109, 91 2, 920, 937, 990, 947, 948) (Sclerotis-140) (Scol iodon-68 , 69 . 88 , 102, 112, 128,
130,132) (Scolopax-233 . 234,235, 237 , 251 ,252 , 253 , 258 , 260 , 26 1 , 263 , 289 , 299 , 300 , 301 , 308 , 326 , 327 , 329 , 346)
(Scomber-111 . 112, 115, 126, 135. 136, 137) (Scomberoides-1 15) (Scomberomorus- 1 31 ,135) (Scophthalmus-44 ) (Scops-269)
(Scorpaena-44,136) (Scutovertex-379 , 376) (Scyliiorhlnus-67,82) (Scyl 1 lum-69 ,76 .77 , 83 , 86 , 1 1 3)
(Scymnorhinus-1 10, 126) (Scymnus-66 , 67 ,76 , 1 10 . 1 16, 126,134,135,137) (Sebastes-45 , 46 ) (Sebastodes-46)
(Selache-70) (Selenophor us-219) (Seleucldes-275) (Semnlosus-76 ) (Semotilus-45) (Senicebus-395 ) (Sepedon-180)
(Sepiella-113) (Seps-413,415) (Ser iola-44 , 1 15) (Serranus-129, 132, 136) (Setonix-386 )
(Slgmodon-374 ,382,404,406,418) (Silur us-38 , 1 44 , 145 . 1 46 , 151 ,155) (Silvicapra-380, 398) (Simotes-175)
(Siniperca-56) (Sltta-272,278, 326) (Smarts- 127, 137) (Solea-35 , 44 . 47 . 11 3 ) (Solen-61 ) (Solenodon-433)
(Solenophorus-218,221 ) (Solenopsis-2 12 ) (Somater ia-192 . 246 . 288 . 290 , 301 , 309 . 31 0. 329) (Somniosus-67 , 76 )
(Sorex-350, 408, 409, 4 12, 423, 424. 425, 426, 427, 428, 429, 430, 433, 434. 435, 439, 440, 442) (Spalax-445) ( Sparus- 1 1 5 , 1 31 )
(Spatula-284,288,292,297,318.319.325,334) (Speotyto-263 . 269 ) (Spermaspiza-297 ) (Spermestes-297 )
(Spermophila-271 ) (Sphaeniscus-194) (Sphecotheris-222) (Spheniscus- 1 96 ) (Sphenocerc us-21 0)
(Sphenorhynchus-262) (Sphyraena- 1 33 ) (Sphyrna-69 , 89 ,90 , 1 1 4 , 128, 136 , 137 ) (Spilogale-365 , 396 ) (Spinax-65 , 66 , 194)
(Splzaetus-274) (Spreo-260) (Squalius-38, 187)
(Squalus-97, 55, 66. 67, 69, 80. 83. 85. 109. 112, 113.120, 125, 133. 134, 135. 136, 154) (Squatarola-238. 300, 338)
( Squat ina-65 ,66,67,68,69.83,86,99,113.118,127.136) (Stallio-415) (Stegoblum-4 1 9 ) (Stegostoma-101 .124)
(Stellio-364.413.414) (Steno-366 , 368 , 449 ) (Stenocellus-260) (Stenodus-58 ) (Stenolophus-219,260)
(Stenorhynchus-1 18) (Stercorar ius-193 . 237 , 263) (Sterna- 185, 186, 187, 188,192, 193, 194,237,246,251 ,252,253,

260.299.354) (Stigmatopel ia-202 . 210, 21 3 . 215. 31 9) ( St i zosted lon-38 . 45 . 58 , 1 40 . 1 4 1 ,143.354) (Stomoxys-298)
(Streps ilas-244, 261 ) (Streptopel la-202, 210,215 , 216 . 217 .21 8 . 292 , 31 9 . 327 ) (Stringops-200) (Strix-269,270,410)
(Strongylura-54) ( Str uthlo-1 83 , 21 1 ) (Sturnla-322) (Sturnopastor-323 ) (Sturnus-231 ,236,
251,253,262,263,300,309,322) (Stylar ia-2 1 ) (Succinea-412) (Sula-1 93 , 1 94 , 1 95 ) (Suricatta-427) (Surnia-269)
(Surnic ulus- 320) (Sus-372 , 384 . 387 . 400) (Sylvia-262,270 , 273 . 279 . 315 , 323) (Sy 1 vilagus-376 . 383 . 406, 444 )
(Syn allaxis-27 3. 275) (Synaphobranchus-45 , 110,113) (Syngnathus-46 , 47 ) (Synodontis-24,25. 143) (Syphonostoma-44)
(Syrnlum-269) (Syrrhaptes-317)

(Taehyglossus-376,392) (Tachyoryctes-445 ) (Tachyphonus-273 ) (Tachys-219) ( Tad or n a- 2 46 , 25 1 , 291 , 293 , 31 8)
(Taenlopt era-27 3) (Taenlura-136) (Tal pa-4 12, 4 19. 429, 433, 4 37) (Tamandua-395 ) (Tamiasciurus-373,419)
(Tand anus-22, 28, 30) (Tandorna-320) (Tantalus-231 ) (Tapirus-377 . 385) (Tnrentola-164 , 177 , 4 1 0, 41 3 , 4 1 4 , 415)
(Tarpon-46) (Tatera-410) (Tater ia-374 ) (Taterona-448) (Tatusla-395 ) (Taurotragus-380 . 38 1 , 397 . 398 , 400)
(Taxidea-364,440,443) (Tchagra-323 ) (Telephonus-346 ) (Temora-126) (Tenebr io-219 . 394 . 41 8 , 427 . 432, 447 )
(Tenebrloldes-219,394) (Tenezio-432) (Terrapene-181 ) (Tetraceros-381 ) (Tetramoriuni-212,214)
(Tetrao-209,216,218.220,222.260.278,279,330) (Tetraogallus-216.222.278, 303) (Tetrapterus-46 )
(Tetrarhynchus-1 33) (Tetrastes-21 8, 220 . 260 , 263 . 278 . 303 . 308 ) (Tetrax-223 , 224 , 303 ) (Tetronarce-84 )
(Thalasseus-233 ) (Thalassoeca-194 ) (Thalassogeron-193. 194. 195) (Thalassoica- 1 95 ) (Thamnophilus-273)
(Thamnophis-174 ) (Theragra-40 . 113, 126) (Thereiceryx-222) (Therlsticus-226,232) (Thomomys-374 . 376 , 379 , 41 9 . 447 )
(Thos-364) (Thous-438, 440,444) (Thryonomys-378 . 405 , 406 , 407 ) (Thrysites-1 34 ) (Thylacis-433)
(Thymallus-33. 38. 39.58. 141. 143, 184.354) (Thynnus-78 . 1 1 1 .115.137.138) (Tichodroma-327 ) (Tigriopus-84 , 125)
(Tigrlsoma-231 ) (Tilapla-141 ) (Tinamus-218,220) (Tinca-20 , 21 . 22 , 23 , 29 , 1 87 , 233 . 244 ) (Tityra-273) (Tockus-270)
(Todaropsis-71 ) (Torgos-205 ) (Torpedo-66.67.69.82.83.84.85.126,138)
(Totanus- 187, 226, 232, 238, 253, 256, 260, 261 ,264,267,290,298) (Trachinotus-132) (Tr achinus-40, 44 . 116 )
(Trachurops-44 ) (Trachurus-73 ) (Trachypterus-46 . 47 ) (Trachysaurus-179, 180) (Tragelaphus-380 . 399 , 400)
(Tragopan-221) (Trapelus-415) (Treron-213) (Tretanorhinus-174 ) (Tr iacis-67 , 68 . 85 ) (Tr iakis-67 . 125)
(Tr ibol ium-21 9 . 260 .394.419.432) (Trichiurus-1 15. 132) (Trichodestes-410) (Tr lchoglossus-203 . 21 3, 216)
(Trlchomycterus-109) (Trichoplossus-203) (Trichopteron-315) (Trie horlbates- 364) (Trichosurus-439)
(Trigla-44.46.47.113.H6.126.135.138) (Trimeresurus-175. 176)
(Tringa-191 . 209, 226, 232, 233, 235, 237, 238, 239, 240, 244, 247, 252, 253, 256, 260, 261, 262, 263,
264,289,291,293,297,298,299,300,301 ,307,312,322,337,342) (Tringoides-238.260) (Trionyx-172)
(Triplecturus-218.219) (Trlturus-161 ) (Troglodytes-261 , 321 , 374 ) (Trogoderma-394 ) (Trogon-236)
(Tropidonotus-175. 176.364.413.414.415) (Tropidurus-179 ) (Tropocyclops-143) (Tr utta-38 , 47 . 136. 14 1 , 142,

185.186.354) (Tr ygon-61 . 62 . 65 , 66 , 67 , 68 .70 .72. 73 .74 . 80 . 82 . 83 ■ 84 . 85 ■ 91 . 95 . 97 . 98 . 99 . 101 . 102. 103 . 1 1 3 ■ 11 8. 1 1 9.
126,128,131,132,133,134,135,136,137,138) (Trypanocorax-231 .309.315) (Tubifex-20) (Tupinambis-174 )
(Turacus-215.217) (Turdoldes-207.308.315) (Turdus- 189, 21 8. 231 .232,233.235.236,237.239.240.247,263,267.
273,300,315,322,323,332,344,345,346,349) (Turtoides-271 ) (Turtur-202.210.213,215,218,319) (Turturoena-202,
213) (Tylosurus-54 . 128) (Tympanuchus-220) ( Ty phi otri ton-161 ) (Tyto-269 )

(Umbra-174) (Uncia-439) (Upogebia-1 19) (Upupa-263 .276 . 309 . 324 . 327) (Uranidea- 1 88 ) (Uranoscopus-1 33)
(Ur ia-1 88. 192. 193.235.236.237.238) (Urinator- 185, 187,296) (Urobatis-69 ,72 .73 . 74 , 84 , 85 . 1 1 8 . 1 1 9) (Urocissa-274 )
(Urocyon-364 . 365 . 396 .439) (Urogymn us-67 .69.70.84,99,103) (Urolophus-84 ) (Uromastix-179) (Uronoscopus- 1 38)
(Urophycls-40.44.55. 135) (Ur sus-355 . 425 . 440 ) (Urubutinga-238)

(Vallonia-208) (Vanellus-234.237.238.239.261 .262. 31 1 . 313. 342) (Vanga-270) (Varanus- 1 70 , 171 , 172, 173,
177,179,181,182,415) (Varicella-208) (Vastres-14) ( Vesper til io-432. 439 . 440) (Vesperugo-429 ) (Vidua-297)
(Vlnago-213.214.215.217) (Vipera-176) (Viscacia-383) (Vitrea-412) (Vitr ina-4 12)
(Viverra-361.363.411.413.437.443) (Vulpecula-79.88. 123. 125. 134) (Vulpes-361 . 364 , 365 , 366 . 4 1 4 , 437 , 439 , 440 , 442)

180

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CESTODA, cont.

(Wallago-145)

(Xantholaema-221 ) (Xema-1 87 , 1 92 . 300 , 301 ) (Xenopeltis-173) (Xenopsylla-418.419.432) (Xenopus-161 )
(Xenorhynchus-241 ) (Xenotis- 1 40 ) (Xerus-374 , 447 ) (Xiphias-60, 111 , 113. 123,126,129.134.135.138) (Xystrlas-1 1 3)

(Zamenis-180,363,364,413,414,415) ( Zanthopygia-27 2) (Zaocys-175) ( Zebethai lur us-355 . 4 1 0) (Zenaidura-202)
( Zenopsi s-60 ) (Zeus-60,122) ( Zibethail ur us-4 1 5 ) (Zoarces-58) ( Zonifer-253 ) ( Zonitoides-208 . 4 1 2 )

( Zonotr icha-27 1 ) (Zonurus-181 ) ( Zosterops-236 , 27 1 . 279 , 31 6 . 323 ) ( Zygaena-68 , 69 , 11 4 , 1 28 . 1 54 ) ( Zygonectes-44 )

( Zygor ibatula-381 )

NEMATODA fro/ M)

( Ablabophis- 162) ( Abramis-30 . 35 . 36 . 42 , 77 ) ( Acanthias-57 ) ( Acanthochaera-260 ) ( Acanthodaetylus- 1 40 . 1 43 )
( Acanthogenys-314) ( Ac an thogobi us-620 ) ( Acanthophi s- 1 14,177) (Accentor-203,204) ( Acc ipi ter-201 ,
202,241,242,248,250,263,267,271 ,277,280,292,306) ( Acer ina-35 , 42 ) ( Acestorhamphus- 1 0 ) (Achirus-46)
(Acinonyx-345, 362,577) ( Acipenser- 1 0, 1 1 , 27 . 30 . 43 , 48 , 49 . 50 . 61 , 64 ,71 .72,76) ( Acomys-343 . 6 11 ) ( Acontias- 1 42 , 1 43 )
( Ac r ido there s-266 ,311,313,314) ( Acris-85 , 89 , 180 ) ( Acrocephalus-209,242, 31 1 ) ( Acrochordus-182)
( Acr yl 1 ium-223 , 225,247) ( Actitis-270 , 277 ) ( Addax-451 ) ( Adenota-395 , 644 , 645 , 646 ) ( Aedes- 1 1 6 , 1 1 7 , 645 ,
650,656,659,661,663,666) ( Aegialitis-201 ) ( Aegol ius-24 3 . 277 ) ( Aegotheles-247 ) ( Aegypius-243 ) ( AeIopus-267 )
( Aelurus-362 , 578 ) ( Aenomys-478 ) ( Aepyceros-368 , 4 1 6 , 4 1 7 , 4 1 9 , 422 , 423 . 45 1 , 480 , 502 , 646 ) ( Aethechinus-340 . 594 . 629 )
(Aethia-224) ( Aethomys-343 ) ( Aetobatls-57 ) ( Afrocichla-243.) ( Afrotis-254 )
(Agama-89, 125, 142, 143, 146, 152, 153,176,177,178,179,180,188,189,567) ( Agelaius-205 . 31 4 ) ( Agkistrodon-121 ,
122, 128, 129, 166, 178, 179,621 ) ( Agonostomus-4 1 .218) (Agonus-59) (Agouti-463.476,480.481,567,628)
(Agriocharis-234) ( Agr iol imax-4 95 . 498 , 499 . 501 , 504 , 508 . 509 . 51 0 . 51 1.524,527.528) (Agrion-42) ( Agrodroma-31 3 )
(Ailuropoda-575) ( Ailur us-37 3 ) ( Aiolopus-267 ) ( A j a ja-205 . 300 ) ( Akodon-343 . 482 ) ( Alactaga-600)
(Alauda-200, 203, 242, 263, 31 1 ,313,314) ( Albida-495 , 499 , 527 ) (Alburnus-30. 35,42,66,73.77) (Alca-12,218,
219,237,264,270) ( Alcedo-262 , 274, 284 . 300 . 308 , 620) ( Alcelaphus-363) (Alces-339,340,416,434,435,440,441 ,
451,459,460,512,527,641,643) ( Alcichthys-66 ) ( Alector is-2 1 0 , 223 , 229 . 234 . 27 1 , 290 , 291 , 327 ) ( Alectur a-253 )
( Alestes-67) (Alle-201 ) ( Alligator- 1 69 , 1 78 , 21 9 , 621 ) ( Allolobophora-200 , 201 , 342) ( Alopex-342 . 51 0) ( Alosa-26 , 27 )
(Alouatta-482,541) ( Alphi tobius-229 ) (Aluco-243) (Alytes-99) ( Atnazona-233 . 235 ) ( Amblopl ites-9 , 43 , 51 , 66 ,73 ,76 )
( Amblyrhamphus-290) ( Ambystoma-86 , 96 , 106,112,157,158) ( Ameiur us-36 , 5 1 ,61,73, 174,219,238,533)
(Aweiva-91,139,141,154,178) ( Amia-62 ,73,238) ( Ammocoetes-47 ,81) ( Ammocr ypta-43 ) ( Ammodytes-27 )
( Ammophor us-229) ( Ammospermophil us-540) ( Ammospi za-260 , 292 ) ( Ammotr agus-397 , 45 1 ) ( Am pel is-300 )
(Amphibolurus-152, 177, 189) ( Amphisbaena- 1 25 . 144 , 178 , 179) ( Amphiuma-1 06 , 109 . 1 17 . 162. 21 9) (Amyda-158, 174)
(Anabas-42,43,57) ( Anabates-300 . 3 1 2 ) ( Anadorhynchus-255 ) ( Anas- 1 2 , 200 , 201 , 204 , 209 ,
210,215,217,218,219,222,223,224,229,234,235,238,239,241 ,242,251 ,262,264,271 ,272, 289, 290, 292, 312, 329, 348)
( Anax-60 1 ) ( Anchoviella-36 ) (Anguilla-24 ,26,34,35,36,38,41,42,48,50,56,57,62,66,70,73,76,80,178,620)
( Anguis-84 ,88,99 ,100,122,126,146) ( Anhinga-203 . 21 8 . 21 9 . 238 , 239 . 240 , 24 1 , 267 . 309 . 329) (Anisolabis-613)
(Anniella-142) (Anolis-139) ( Anomalurus-557 . 559 ) (Anopheles-645, 650. 659. 661 .663.666,670) (Anous-239)
(Anser-200, 209, 21 0,21 5. 21 6, 223. 233. 234. 271 .291 .326) ( Antechinus-508) (Anthropithecus-343.395.541 ,595)
( Anthropopithecus-335 ,371 ,395) ( Anthus-202, 203 , 242 , 296 , 31 3 ) ( Ant idorcas-339. 363.394.416,419.423,436,441 ,
451,458) (Antigone-234.290) ( Antilocapra-335 . 342 . 344 . 416 . 430 . 434 , 438 . 457 . 458 . 459 . 670) (Antilope-339.
365,394,417,451) (Aonyx-586) ( Aotus-541 ) ( Aphanopus-590 ) ( Aphareus-54 ) ( Aphelocephala-298)
(Aphelocoma-313,314) ( Aphodius-599 . 604 , 61 4 ) ( Aphredoderus-238) ( Apl ites-27 , 43 , 73 ) ( Aplodinotus-43,
66,69,73,76,238) ( Apodemus-332 , 342 , 343 , 345 , 453 , 454 , 482 , 484 , 544 , 553 , 575 , 580 , 599 , 61 3 , 625 , 629 , 630)
( Apolichthys-29) ( Aptenod ytes-239) ( Aptychotrema- 1 0 . 57 ) ( Aquil a-24 1 , 242 , 256 , 262 , 263 , 275 , 277 , 292) (Ara-233)
(Aramides-200,201 ,291 ) ( Ar apaima-34 , 43 . 77 , 1 70 . 62 1 ) ( Aratinga-233 ) (Arbacia-55) ( Arbicol a-566 )
( Arbor ico la-225 , 327 ) ( Ar chibuteo-205 . 24 1 ) ( Arctictis-631 ) ( Arc tocephal us-587 , 590 )
( Arctogale-335 ,344,345, 524 ,528) (Arctomys-538,540,544,575) ( Ardea-3 1 , 21 8, 21 9, 233 , 236 , 237 , 239 , 240 , 24 1 , 242 ,
243,258,272,277,278,289,292,300,307,309,318,319,323,329) ( Ardeola-218 . 239 . 259 , 278 . 31 8, 31 9 , 329) ( Arenaria-281 )
(Argas-650) (Argentina-35) (Argusianus-226,227) ( Ar ion-498 , 51 0 . 51 1 . 527 ) ( Arius-27 , 47 ) ( Armadillidium-271 )
( Armigeres-645 , 659 ) ( Aromochelys- 1 83 . 184) ( Artibeus-343.462,469,655) ( Artogale-431 )
(Arvicanthis-482.492.599.623.627.631 ) ( Arvicola-341 ,346,453,484) (Asellus-42) ( Ashmunell a-96 ) ( Asida-247 , 250 )
(Asio-202, 204, 209, 242, 243, 247, 277, 299, 305, 307, 608) ( Aspid ites- 1 62 , 177) ( Aspis-176) ( Aspius-30 , 77 )
(Asthenes-312) ( Astr inula-278) ( Astrophis-162) ( Astur-3 1 , 204 , 238 , 240 , 242 , 246 , 252 , 256 , 267 , 269 ,
270,271 ,277,280,292,297,309,317) ( Astyanax-44 , 46 ) ( Astylosternus- 1 04 . 109 , 1 44 ) ( Ataenius-608) ( Ateleopus-37 )
(Atelerix-340,624,625) ( Ateles-336 , 343 . 539 . 54 1 , 542 , 6 1 4 , 626 . 649 , 666 ) ( Ateuchus-6 1 3 )
(Athene-209,229,243,263,297,326) (Atherinopsis-45) (Atilax-586) ( Atractomor pha-267 ) ( Attila-202) (Auxis-42)
( Ay thya-2 1 5.291) (Azuma-10)

(Bagarius-23.44) ( Bagre-39 ) ( Bagrus-53 ) ( Bakksinel la-659 ) ( Balaena-676 ) ( Balaenopt era-585 ,586,590,676)
(Balanosphyra-314) ( Bal antiopter yx-63 1 ) ( Balear ica-234 ) ( Bal istes-27 , 48 , 49 , 55 ) ( Bainbusicola-277 . 319)
(Barbus- 15. 20. 30. 42. 43. 48. 50. 66. 67. 69. 71 ,72,73,218,219,278) (Bascanion-178) ( Bassar iscus-367 )
(Bathyergus-428,481 .566) ( Batrachemys- 1 34 ) ( Batrachoseps-92 ) ( Batrachus-52) ( Batracocottus-9 )
( Bdellonyssus-654 ) ( Bdellophis-84 , 1 0 1 ) ( Bdellostoroa-81 ) ( Beaufortia-66 ) (Beluga-522) ( Bernlcla-223 , 264 )

INDEX: SYSTEM A HELMINTHUM

181

NEMATODA, cont.

(Bhringa-306) (Bison-342,416,422,435,440,451 ,457,458,512,645) (Bitis-121. 128. 130.162, 163,164,180,599,621)
(Biziura-215.289) (Bizura-215) ( Blabera-569 ) ( Blaps-229 , 23 1 , 594 , 6 1 2 , 6 1 3 ) (Blarina-342.482.505.589.625)
(Blatta -594,599,613) ( B1 attel la-252 , 253 , 289 . 290 , 292 , 599 , 61 4 , 625 , 627 , 628 ) (Blennius-27,75) ( Blicca-9 , 30 , 35)
(Boa-128,130,178) (Boaedon- 162, 164,605) ( Bombinator-84 , 86 , 88 , 9*1 , 99 , 100,146) (Bombyna-608)
( Bonasa-1 99 ,200.,223, 231 , 233 , 23*1 , 260,268, 271 , 322) ( Bos-339 , 3*10 , 3^2 , 363, 365 ,
416,421,422,434,435,451,457,458,531,621,635,644,645,668,669) (Boselaphus-397,418) (Botaurus-219,238.239,
264,267,277,278,319,323) (Bothrops-128, 129, 164. 178) (Brachylophus-146) (Braehyotls-305) ( Brachypter us-278 , 292)
(Brachyteles-449) (Bradypus-449,469,606,607,609,616,653,662) (Branchiostegus-47.52)
(Branta-200,209,210,215,216,223,272) ( Brosmius-35 ) (Brycon-66) ( Bryttosus-4 1 ) ( Bubal is-428 , 435 , 45 1 , 646 )
( Bubalus-669 ) (Bubo-202,205,242,243,247,260,277,292,306) ( Bubulcus-21 7 , 21 8 . 247 . 26 1 , 278 , 289 , 293 )
(Bucco-231 ,292) (Bucephala-203,204,264) ( Buceros-3 1 3 ) ( Buchanga-267 ) ( Budytes-266 . 267 ) ( Buf felus-436 , 45 1 , 635 )
(Bufo-76, 84, 85, 86, 88, 89,90,93,94,95,96,97,99, 100,101,102,104,105,106,108,109,110,113,114,116,118,146,154, 158,
167,601,620) rfeulinnnus-495) (Bungarus-129, 130, 163,164,184) ( Bur linus-254 )
(Butastur-240) (Buteo-201 ,202,209,238,241 ,242,247,248,256,261 ,263,277,278,296,297,307,608) ( Butor ides- 1 99 .
239,240,259,289,309)

(Cabassus-424,429,44 3,475,560,575.581 ) ( Cacatua-233 ) ( Caccabis-223 , 225 , 229 , 230 , 234 , 235 , 249 , 252 , 254 ,
267,271,290,317,327) (Cacopus-109) (Caiman-168, 169, 190, 195) (Cairlna-200, 223, 234, 241 ,264,329)
(Calamanthus-311 ) ( Cal idr is-242 , 269 , 276 ) ( Call ichrous-23 , 44 , 46 , 48 ) ( Call ionymus-9 , 27 ) (Calliope-200)
( Call ipepla-230, 232, 234) (Callisaurus-136) ( Call ithrix-524, 541 ,569,624) ( Callorhinus-372 , 590 )
(Callosciurus-476 ) ( Callospermophi 1 u-540 ) ( Cal lotar ia-590 ) (Caloenas-212) ( Calopterocles-223 )
(Calotes-142, 143, 152, 164, 169, 173.176,178,192) (Calotis-152) ( Caluromys-569) ( Cambarincola-534 ) ( Cambarus-620 )
(Camelopardalis-339) (Camelus-335 , 339 , 34 1 , 397 , 416 . 417 , 420, 422 , 430 , 434 , 435 , 436 , 45 1 , 457 , 458 ,
460,480,512,617,653,668) (Gampethera-306) ( Campostoma-66 ) ( Cancroma-205 ,238,289,290) (Canis-339.
343,345,348,372,401,505,510,543,577,599,600,601,624,626,627,629,630,631,635,640,662) ( Cantharus-49 )
(Canther ines-9 ) (Canthon-608) (Capella-209,262-,269,286) ( Capoetobrama-66 ) (Capra-336,339.340,341,
34 4,383,397,416,417,420,421 ,430,4311,1135,441 ,1)51 ,457,458,459,496,499,501 ,502,504,508,527,531 ,538,556, 614, 644)
(Capreol us-339, 340, 4 16, 4 17, 421 ,434, 435, 439, 440, 441, 451, 457, 458, 499, 512, 527, 538, 609, 643, 644, 672)
(Caprimulgus-201 ,229,230.231 ,233,255,256,263.283,297,302.330) ( Caprolagus-433 ) ( Capromys-482 , 487 , 555 )
(Capros-27) ( Car anx-26 , 28 , 42 . 48 . 49 , 60 ) ( Carassiops-39 , 21 9) (Carassius-30 ,76 ,77 , 608) ( Carcarhinus-9 , 10)
(Carcharias-37 ) (Carcinus-57) (Cariama-229) ( Caridina-73 ) (Carine-205,230,270) (Carollia-655)
( Carpiodes-42 , 66 ) ( Carpophaga-257 ) ( Car pophilus-267 ) ( Cascara-209 ) ( Casmerod ius-329 ) ( Cassiculus-260 )
(Cassicus-300.314) (Cassidix-313) (Castor-339,343,353,441.575,651) ( Casuarius-209 , 247 ) (Cathaica-499)
( Cathar tes-289 ) ( Catheturus-223 , 233 ) ( Catolynx-506 ) ( Catorrhac tes-270 ) ( Catostomus-9 , 36 , 49 , 66 , 67 , 77 , 238 )
(Caularchus-76,77) ( Causus- 1 09 , 1 30 , 163 , 1 64 ) ( Cavia-264 , 339 , 343 , 41 8 , 483 , 49 1 , 567 , 589 , 6 1 3 , 620 , 652 )
(Cav lei la-54 5, 555, 652) ( Cebus-335, 343 , 394 , 41 6 , 431 ,492,525,541,612,613,614,624,630,640,649,650,651)
( Centetes-624 ) (Centrocercus-249,266) (Centro no tus-26) ( Centrophi 1 us-627 ) (Centro pus-229,230,231 ,
233,236,260,282,298,304,319) ( Cepaea-495 , 498 , 499 . 510 . 527 ) ( Cephalophus-340 , 396 , 434 , 45 1 , 599 , 609 , 640 , 644 ,
645,646,662,670) (Cephalopter us-249. 300) ( Cephaloscyll ium-37 ) (Cerastes-140,143,164,178,179,180)
( Cer atodus-1 6 ) ( Cer atophora- 1 51 . 307 ) ( Ceratophr ys-88 , 90 , 94 . 95) ( Ceratophyl 1 us-600 ) ( Cerchneis-248 ,
270,271,277) ( Cercoeebus-395 . 662 ) (Cer come la-205, 297) ( Cercomys-478 , 483 ) ( Cercopithecus-336 , 359 ,
395,396,398,569,587,611,613,624,626,630,636,640,662,665) (Cerdocyon-624) (Ceriornis-223) (Certhis-321 )
(Cervicapra-644,645) ( Cervus-338. 339 . 340. 365 . 383 . 395 . 421 ,440,44 1,451,457,458,492,499,500,503,504,512,531 ,
609,614,640,642,644,670,679) ( Cer yle-252 . 269 . 308 ) ( Cetorhinus-29 ) ( Ceuthophi 1 us-630 ) ( Chaenichthys-52 . 586 )
(Chaenobryttus-43.73.238) (Chaenocephal us-52, 590) (Chaenogobius-42) ( Chaetophractus-361 ,616) (Chaetura-322)
( Chalc alburn us- 30 ) (Chalc ides- 142, 143,144) ( Chalcophaps-2 1 2 ) (Chalinura-59)
(Chamaeleo-109, 144. 152. 154.164.176.177.178.179,189) ( Chamaepel ia-235 ) ( Champsa- 1 78 , 1 90 )
( Champsocephalus-52 .60,586) ( Char ad r ius-201 , 21 8 , 21 9 , 242 , 243 , 264 , 276 , 277 , 287 , 290 , 292 , 297 ) ( Chaul el asmus-2 1 5 ,

271.289) (Chauna-216) (Chaunax-27) (Chelidon-268.330) (Chelidoptera-231 ) (Chelone-156, 167, 170)
(Chel^dra-123, 158, 159, 174, 183, 184, 194) (Chen-209,210,216,224) ( Chenonetta-223 ) ( Chenopsis-202 , 21 5 , 222 , 289 )
(Chersus-134) (Chettusia-313) ( Chibia-256 , 306 ) ( Chiloscyll ium- 1 9 . 37 , 56 ) ( Chimaera-35 ) ( Ch ionomys- 34 3 . 54 4 )
(Chlrocentrus-36 ) (Chironitis-613) (Chitra-174) (Chlamydosaurus-152. 189) (Chlidonias-202,269) (Chloea-36)
( Chloephaga-223 ) ( Chlorophoneus-3 1 2 ) (Choloepus-627,662,664) (Chondrula-506) (Chordeiles-322)
(Choriotis-230. 231 .254.268.282) ( Chor tophaga-252 . 290 ) ( Chr istivomer-9 ) ( Chrosomus-66 )
(Chrysemys-157. 158. 174 . 183. 184 . 185.219) (Chryserpes-313) (Chrysoconops-650)
( Chr ysolophus-200, 201 .203.204.221 .223) ( Chr ysoperca-238 ) ( Chr ysophr ys-39 , 48 , 52 , 53 ) (Chrysops-650,665)
(Chrysotis-201 ) ( Ciconia-1 76 . 209 , 21 9 ■ 225 . 239 . 246 . 27 1 . 272 . 277 . 278 . 292 . 305 . 330 ) ( Cinclosoma-242 ) (Cinclus-201 ,
276) (Cinixys-134, 136, 154) ( Cinnyr is-307 ) ( Cinosternum- 1 83 , 1 84 ) ( Circaetus-242 , 263 ) (Circus-202,
209, 234, 238, 241, 242, 243, 247, 248, 277, 278, 300, 313AS) ( Cissa-3 1 6 , 327 ) ( Cistudo- 1 58 , 179 ) (Citellus-339.
340,342,343,416,417,446,451,458,540,553,575,594,600,605,612,613,624,625,627,630) (Clangula-203,218,219,264,

272.290) (Claravis-212) (Clarias-42,43.44,45.46.76,619) ( Clarotes-48 ) (Clemmys-158, 174, 183, 185)
(Clethrionomys-332.345.415.453.455.484.544.553.599.631) ( Cloel ia-178 ) ( Clupea-26 , 28 , 29 . 35 )
( Cnemidophorus- 1 40 .141. 178) ( Cnidoglanis-52 ) (Cobitis-36) (Cobra-158) (Cobus-365,422,429.451,645,646)
(Cochlearius-241 ) ( Cochl icel la-495 . 499 ) ( Cochl icopa-504 . 528) ( Coelopel t is- 1 64 , 1 76 )
(Coelorhynchus-10. 30.33.59.70) ( Coeluromys-630 ) (Coendu-371 ,478,480,482,558,655,656) (Coilia-27)
(Colaptes-203,247,311 ) ( Coleonyx- 1 40 , 14 1 ) ( Col inus- 1 99 . 200 . 209 . 21 0 , 21 2 , 222 , 223 , 229 , 23 1 , 232 , 234 , 252 , 260 ,
268,271 ,289,290,322) ( Colobus-353 . 61 1.662,663.666) ( Coloeus-201 , 204 ) (Colonocus-345 . 373 , 431 , 51 0. 524 ,
535,575,627) (Colossoma-15) (Coluber-106.121. 122,128,129,130,143,158,162,163,167,176,177,178,179,190)
(Columba-201 .203,212.233.235.271 .289.290.292.300.314) ( Columbigal 1 ina-2 1 2 , 303 ) (Columbina-212) (Columella-496)

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NEMATODA, cont.

(Coljrmbus-12,218,219,237,240,243,264,269,272,274) (Comephorus-63.68) (Conepatus-362.625,639)
(Conger-27. 28, *48, *49. 76) (Connochoetes-451 ) (Conolophus-146, 148) (Constructor- 12 8, 187) (Cophotis-151 )
(Copris-253.608) (Copsychus-260 , 266 , 301 , 323 , 327 ) ( Cor aclas-230, 231 .2*17,2*18,2*4 9.250.252.
266,271,286,302,31 1,327) (Coracina-202) ( Cor agyps-250 , 289 ) ( Coregonus-9 , 31 . *42 , 68 . 69 . 73 ,78) (Coreopsis-223)
(Coronella-128, 163) (Corucia-151 ) (Corvus-200, 201 , 202 , 20*4 , 209 , 223 , 230 , 2*42 , 2*4 3 , 256 , 260,
263,266,267,271 ,284 , 291 , 292 , 296 , 297 , 300 , 301 , 309 , 31 1 , 31 3 , 31 *4 , 321 , 32*4 , 329) (Corynorhinus-565) (Coryphaena-62,
69) (Coscoroba-209) (Cottocomephorus-61 ,62,63.68.587) (Cottus-26, 27. 35, 42. 59, 61 ,62.63.66,590)
(Coturnix-201 ,203,204,222,223.229.230.231 .234,252,266,267) (Cotyle-268) ( Cracticus - 3 1 2 ) (Cranorhinus-250)
(Craterocephalus-219) ( Cr atogeomys-482 ) ( Cr ax-224 , 255 , 26 1 ) ( Cren ilabr us-49 ) ( Cr icetomys- 339 . 34 3 .
428,483,489,566,599) (Cricetulus-343.613) ( Cr icetus-344 , 348 , 453 . 486 . 544 , 599 ) (Criniger-311 )
(Cristivomer-49,69,79) ( Crocidura-332 , 335, 343 , 344 , 346 , 481 ,483,595,614,625,628) (Crocodilus-123, 167,168,
169,170) (Crocopus-233) (Crossodactylus-86) ( Crossoptilon-224 ) (Crotalus-123 , 128, 129, 1 38 , 163 , 164 , 165 , 187 , 196)
( Crotaphopeltis- 1 62 ) (Crotophaga-230.252,260,271 ,282) ( Cr yptobranchus- 1 06 ,107.114,118) ( Cr yptoglaux-202 )
(Crypturellus-212,225,256) ( Cryptur us-202 , 223 . 225 , 230 . 231 ) (Crysolophus-224) (Ctenocephal ides-651 , 66 1 )
(Ctenodactyl us-458, 558, 627, 633) ( Ctenodon-1 54 . 178) ( Ctenomys-567 ) (Ctenosaura-1 36) (Cuckoo-236)
(Cue ulus-228, 229, 23 1,234, 282, 330) (Culex-1 16, 117, 189.645.650.659,661 ) (Cul iciomyia-659) (Culicivora-300)
(Culicoides-326, 650, 651 ,660,668) ( Cumana-235) (Cupidonia-223) (Cupidonica-222) (Curimatus- 17, 19,45)
( Cursor ius-263 ) (Curucujus-322) ( Cyanerpes-287 ) (Cyanocephal us-339) (Cyanocitta-266,293)
(Cyanocorax-230,267,311 ,314) ( Cyanoplca-202 , 31 4 ) (Cyclemys-158) (Cyclodus-177)
(Cyclops-42,45,76,77.174,194.195,329,619.620.621.678) (Cyclopterus-28, 35) (Cycloturus-490)
(Cyclura-134.138.146) (Cygnopsis-223) ( Cygnus-202 . 204 . 215 , 216 . 217 . 21 8, 223 . 237 . 27 1 , 289 , 327 , 330)
( Cyn aelur us-36 1 ) ( Cynict is-4 31 , 569 ) ( Cynolebias-4 1 ) ( Cynomolgus-349 , 394 . 395 . 398 . 569 ) (Cynomys-343.
417,624,625) ( Cynonycter is-579 ) ( Cynopi thee us-569 ) (Cynoselon-27,30) ( Cyornis-299 ) ( Cypr inodon-27 )
(Cyprinotus-278) ( Cypr inus-1 0 , 30 , 42 , 48 , 66 ,77 . 620) (Cypselus-330) (Cysticola-314) (Cystignathus-88, 1 16)
(Cystophora-587 , 590 )

(Dacelo-230,256,294) ( Dactylophora-53 ) (Dactylosternum-267 ) (Daf i la- 1 99 . 20 1 , 24 1 . 289 , 290) (Dama-397.
421,422,440,457,458) ( Damal iscus-335 . 339 . 340 , 363 . 4 1 7 , 4 1 9 . 422 , 434 . 436 , 45 1 . 480 . 5 1 6 , 645 , 646 ) (Damonia- 1 84 )
(Daphnia-272,290) (Daption-274 ) (Par yprocta- 339 ) (Pasyatis-34 , 59 ) (Pasycottus-78) (Pasymys-431 )
(Pasypeltis-162) (Pasyprocta-335 . 37 1 . 383,477,480,488,555,566,608,636) (Pasypus-361 ,362,418,423,424,429,
430,438,487,551,560,561,581,649,653) (Pasyurus-601 ,632,649.650) (Pecticus-267 ) (Pel ichon-266 . 31 3 , 31 4 )
(Pelphinapter us-520 , 522 , 585 ) (Pelphinoptera-676 ) ( Pel phinus-5 1 8 , 51 9 , 520 , 52 1 , 585 , 586 , 587 ) (Pemansia-1 14.
163,177) (Pendroaspis-1 30 ) (Pendrocitta-266 , 268 , 327 ) ( Pendrocolaptes-300 , 31 3 , 330 ) (Pendrohyrax-427 .
573,583,645) (Pendrolagus-651 ) (Pendromus-599 ) (Pendropicos-261 ) (Penisonia-1 14,177) (Pentex-47)
(Permatemys-1 84 ) ( Permestes-229 . 267 ) (Permophis-1 00 ) (Peroceras-96 , 498 ) (Pesmiegretta-259) ( Pesmodus - 4 6 8 )
(Pesmognathus-86,96,98, 101,115) (Piagramma-53 ) ( Pi aptomus-45 , 77 ) (Pichocercus-292) (Picbolophus-235, 260)
(Picotyles-370 , 383 , 607 , 609) (Picrodon-180) (Picrostonyx-453,553) (Picrur us-286 . 306 . 307 . 31 2) (Pictychus-72)
(Pidelphis-340, 341 ,342,343.420,429.442,475.477,482,516.526.555.560.563,564,566,570.576,596,614, 615, 620, 621 ,
623,628,658,678) (Piemyctylus-85 , 86 , 112) (Pikerogammarus-1 1 ) (Pinodon-129)
(Pinomys-557 ) (Piodon-37 , 1 30 , 620 ) (Piomedea-238, 239 , 240 , 274 , 285 , 289 , 298 ) (Piomedeadaption-274 )
(Pioptrornis-261 ) (Piplodus-38.50) (Piplophysa-72) (Pipodomys-339,558,612,613,630) (Piporiphora-152)
(Pipsadomorphus-128,163,178) (Pipsosaurus-146) (Piscoglossus-278) (Pispharagus-278) (Pispholidus-109, 166)
(Pissemurus-321 ) (Pissoura-319) (Pistiehodus-17) (Piuca-312) (Pives-31 1 ) (Polieholophus-229)
(Polichotis-339,449) ( Pomicella-233 ) (Poras-15) (Porcelaphus-451 ) (Porcopsis-405) (Porosoma-29) (Promaeus-212)
(Prymarchon-129, 163) (Prymobius-106, 122.123.125.158,163.171) (Pryobates-204 . 248 ) (Pryobatus-248)
(Pryocopus-268) (Pryolimnas-281 ) (Pryomys-629) (Pr yophis- 1 28 , 1 29 , 179 ) (Pryoscopus-312)

(Echidna-55,432) (Echinus-76) (Echymipera-626) ( Egocerus-422 , 646 ) ( Egretta-21 9 , 239 , 24 3 , 259 , 277 ,
278,309,329) ( Eisenia-200, 201 .208.514.515.531 ) (El anus-247 , 248 . 260) (Elaphe-121 ,128,129,130,163,182,
189,278,608,620) (Elaphomorphus-192) (Elassoma-238) (Eleginops-52) ( Elephan tulus-570 , 594 . 65 1 ) (Elephantus-617)
(Elephas-354, 367 ,386,532.579.617) (Eliomys-453.633) (Ellobius-608) (Elosia-86 , 95 , 106, 158) (Elseya-158)
(Emberiza-202.204,270.312) ( Emissola- 1 3 . 37 ) (Empidonax-294 ,296) (Emyda-174) (Emys- 157, 158, 159, 167, 174, 183, 184)
(Ena-499 ) (Enhydra-590) (Enneacanthus-219,238) (Enneoctonus-260 , 267 ) (Enygrus-129) (Eophona-262)
(Ephippiorhynchus-319) (Epigeichthys-76) (Epigrainmophora-501 ) (Eplmys-336,554,599,613,624)
(Epinephel us-28, 36 ,76 , 77 ) ( Epiphragmophora-498 .510,528) (Eptesicus-346,470,565)
(Equus-336, 354, 356, 357. 358. 376, 377. 378, 379. 380. 389. 390 .391 ,416.538.597.612.640.644.645.651 ,678) (Eremias-175)
( Erethizon-492 . 555 .558,639.649,656.663) (Er iculus-624 ) (Ericymba-43) (Erignathus-587.590) (Erimyzon-41 ,238)
(Erinaceus- 176, 342, 343. 346. 348, 51 0.529 .60 1,61 3. 61 4, 623. 624. 625. 626. 627, 631 .637) (Eriocheir-620)
(Erithacus-201 ,297) (Erolia-243 , 277 ) (Erythrocebus-61 1 ) ( Er ythrocul tur-76 ) ( Er ythrol amprus- 1 60 , 1 63 , 178 )
(Erythrourus-624) (Esox-9 , 12 , 27 , 35 , 36 , 4 1 , 42 , 43 , 73 . 76 , 77 , 21 8 , 21 9 ) (Etheostoma-43) (Eualus-59)
(Euarctos-575 , 662 ) (Euborellia-229) (Eucalia-66) (Euchiloglanis-66) (Euconulus-504 ) ( Eucyc lops-6 19 , 620 , 62 1 )
( Eudromias-242 ) ( Eudyptes-239 . 285 ) (Eudyptula-239) (Eulota-511) (Eumeces-96, 126, 158, 180)
(Eumetopias-372.528,586,587.590) (Eumops-630) (Euneacanthus- 1 2 ) (Eunectes-123 . 196 , 372) (Euomphalis-498)
(Eupagurus-57 ) (Euparypha-499 ) ( Eupemphix- 11 6 ) ( Euper ypha-495 ) ( Euphr ac tus-36 1 , 423 , 424 , 429 , 4 38 , 44 3 )
(Euplocamus-222.224,247) (Eupodotis-247 ) (Eupomoti s-9 . 10 , 36 , 43 , 73 , 238 ) (Euprepis-178) (Euprocyon-361 )
(Europaeus-613) (Eurycea-69.88.101.115) (Eurypyga-281 ) (Eurystomus-230) (Eusimulium-326,669)
(Eutamias-446.548.553.594.630) (Eutropichthys-73) (Euxestus-267 ) (Evotomys-342 , 344 , 453 , 455 )

(Excalfactoria-231 )

INDEX: SYSTEMA HELMINTHUM

183

NEMATODA, cont.

( Falco-202, 209, 2^1, 2A2, 2»t 3, 2A8, 2^19,256, 262 ,263, 266, 270 ,27 1,277, 278, 290, 297, 300 ,309, 31 0,312,31*1, 321, 330)
( Fel ls-335. 339, 3*1 1, 3*13, 3*1**, 3*15. 3*<8, 36 1,362, 363. 367, 372, 43 1 , **6 1 ,*<98, 506, 51 6, 529, 53 1,569, 576, 577, 579, 599, 605, 606
619,620,623,624,625,626,627,636,637,662,663,678) (Fennecus-579,605) (Fiber-340,343,345,416) (Florida-238,300)
( FI uv ldraco-36 , 46) (Forcipomyla-1 17) (Forficula-625) (Francollnus-223 ,229, 230, 234 ,252 , 263 , 267 , 31 8, 31 9, 323)
( Fraterula-264 ) ( Fregata-239) (Frlngilla-200,202,203.270, 31 1 ) (Frutlcicola-495,498,499,504,527) (Fuadias-310)
(Fullea-1 99, 21 4, 21 5. 21 6, 21 8, 289, 290, 291 .300,301) (Fuligula-214.215.264) (Fulmarus-285) (Funambulus-553.568)
(Fundulu3-30.36.53.219) (Funisciur us-345. 483. 599) (Furnarlus-31 1 )

(Gadopsls-218) (Gadus-26,27,28,35,47,59,.70,72,590) (Gaidropsarus-47 ) (Galago-362, 569, 623. 625)
(Galaxlas-41 ,218) (Galba-500) (Galea-415.449,483,491,548,652) (Galeichthys-50) (Galeocerdo-26 , 37 . 57 )
(Galeopi thee us-547 ) (Galeopterus-546 ) (Galeorhlnus-50) (Galeus-57) (Gal 1 inago-289 ) (Gallinula-242,262)
(Galloperdix-230.325) (Gallus-1 99, 200. 201 ,203,204,205,210,212,222,223,224,225,229,
231,233,234,235,253,254,261,264,267,271,282,289,291,348,625) (Gambelia-137) (Gambusia-43,278) (Gammarus-1 1 ,
68,264,289,290) (Gampsoelels-267 ) (Garrulax-324 ) (Garrulus-200 , 202, 204 , 242, 243, 260, 270. 301 , 31 2, 31 4 ,
318,321,324,326,330) (Gasterosteus-36,42.68) (Gastrophr yne-96 , 106 ) (Gavia-209.238) (Gav ial is-39 .
165,168,170,194) (Gazella-339, 340 , 34 1 . 397 . 416 . 417 . 420 . 426 . 451 , 458 . 459 . 496 . 556 , 670 ) (Gecinus-268) (Gecko-139,
140, 142, 14*1,151,196, 620) (Gehyra-140) (Genetta-431 ,599,605,626,659,662) (Gennaeus-223 , 224 )
(Geoelemys-158, 183,620) (Geomyda-1 34 ) (Geomys-599) (Geopelia-235) (Geophagus-17) (Georychus-339)
(Geosciur us-6 1 1 . 624 ) (Geotrupes-601 ,608.613) (Geotrygon-235) (Geranospiza-300)
(Gerbillus-453. 483. 569. 611 .612,626.630.631) (Gila-9,66,238) (Glnglymostoina-37 ) (Giraffe-339) (Girasia-503)
(Girella-48 , 59 ) (Glanidium-66 ) (Glareola-281 ) (Glaucidiuin-205,223.230,256) (Glaucionetta-239 , 289 . 329)
(Glaucomys-342,446.541 .554) (Glis-453,482,483,629) (Globiocephal us-520. 522. 585. 586) (Glossogobius-45.67,619)
(Glossophaga-655 ) (Glyptothorax-67 ) (Gnathopogon-36 ) (Gobio-66) (Gobiomorus-27 ) (Goblus-26 , 21 8) (Golunda-343 )
(Gongylo phis- 129) (Gongylus- 1 40 , 152 ) (Gonocephal um-229 . 267 ) (Gonyocephal us- 152 . 153 , 177 ) (Gopherus-1 34 , 149)
(Gorgon-363) (Gorilla-398. 428, 541 ,551 .595,666) (Gorsakius-620) (Grallina-202) (Grampidelphis-676 )
(Grampus-522) (Graphlurus-176 ) (Graphophaslanus-223,252) (Graptemys-158, 159. 174, 183) (Graucalus-324 )
(Grayla-1 80) (Grison-344) (Grossiptodon-222,223) (Grossospiculagia-434 )
(Gr us-203, 209. 235, 241 ,242,243,280.290.330) (Gryllus-627) (Guevei-645) (Guira-260)
(Gulo-345, 431 ,510.535,575.627) (Guttera-235) (Gymnocranius-59 ) (Gymnodactylus- 1 40 , 177 )
(Gymnopleurus-601 ,604,613) (Gymnorhina-202, 311 ) (Gymnothorax-48) (Gypaetus-242,248) (Gyps-242.243,277)
(Gypsophoca-587 ) (Gyrlnophllus-1 15)

(Hadropterus-43 ) (Haematopoda-665 ) (Haematopus-280 ) (Haemulon-36 ) (Hagedashia-226,239)
(Halcyon-246,250.256.269,276,277.283,307.313.316.326) (Hal iaetus-239 . 256 ) (Hal iastur-239 ) (Hal ichoerus-586 ,
590) (Hal lcore-588 ) (Halieutaea-33) (Hal iotis-55 ) (Halmaturus-651 ) (Hardella-159. 185) (Harelda-200,218,219)
(Harpagifer-52.59) (Harpallus-625) (Hegeter-605 ) (Heleioporus-H8) (Helicel la-495, 497, 499, 506, 527)
( He 1 ichoer us-587 ) (Hel icops-1 63 . 182) (Helicosciurus-599) (Helioperca-9.43.73.238) (Heliosciurus-152,554)
(Helix-495.496,498.499.501.502.510.527.528) (Helminthoglypta-498 ) (Heloderma- 1 91 ) (Helodrilus-201 .514,513.515)
(Helosla-94) (Helotarsus-24 1 ) (Hemidactylus-140. 1*12, 14 3, 178, 180.340) (Hemlergls-142) (Hemilepidotus-590 )
(Heosemys-126, 159. 183) (Hepsetia-36) (Heptagenea-73) (Herodias-219,239.330)
(Her pestes-367. 503. 522. 526. 579. 594. 631 .636.637) (Herpetodryas-164) (Hesperomys-655 ) (Hetairus-59 )
(Heterobranchus-44 .46) (Heterocephalus-558 ) (Heterodon-96, 122, 123. 128.163.178.179) (Heterodontus-48,55.56,585)
(Heterohyrax-392.583) ( He ter opneust us-46) (Heterotetrax-254 ) (Hexagenia-61 ,73) (Hexanchus-57 )
(Hierac idea-256, 263, 267. 293) (Hieremys-159) (Himantolophus-31 ) (Himantopus- 199 ,201 .215.242,269.290)
(Hiodon-4 3,66) (Hlppocentrum-665 ) (Hippoglossus-27 . 28 , 59 ) (Hippopotamus-348.427.532,563.575,650)
(Hippotragus-395.396.4l6.417.422.435.451 .645.646) (Hipsirhina-185) (Hir undo-202 . 203 . 266 . 268 . 31 3. 31 4 . 325 )
(Histiophoca-590 ) ( Hi stiophor us-28) (Hodotermes-254 ) (Holacanthus-1 8) (Holbrookia-1 36) (Holocentrus-50)
(Holochilus-479. 481 .482.489.654) (Holoquiscalus-31 1 ) (Homo-343,395,416,417,451,624) (Homopus-196)
(Hopllas-10,46) (Hoplobrotula-33) (Houbara-223.254 ) (Huro-43, 51 ,73,76,77,238) (Hyaena-624.663)
(Hyalella-72 , 289 ) (Hyalinia-495.527) (Hybomys-599,623) (Hyborhynchus-66) (Hydrochel idon-203 , 273 , 286 )
(Hydrochoer us-335. 343 .491 .543.597) (Hydromedusa-112) (Hydromys-61 8) (Hydropo tes-340 , 344 , 397 , 4 1 5 , 4 16 , 435 , 457 )
(Hydroprogne-417) (Hydrosaurus-157. 187) (Hydrurga-528 . 586 , 587 . 588) (Hydrus-169)
(Hydurga-588 ) (HYla-85 . 88 . 89 , 91 . 92. 93 . 94 . 95 . 96 , 99 . 100 , 106 . 110, 112 . 1 1 4 . 1 1 8, 158,177,620)
(Hylobates-334. 335. 361. 395. 396. 540. 551 .608.611 .650.662.678) (Hylochoerus-381 . 395) (Hylodes-90,93,97)
(Hylomotus-268) (Hylophylax-274 ) (Hylotomus-248) (Hynobius-86 , 620) (Hyperoodon-518,585,676)
(Hyphantornis-267 . 324 ) (Hypomesus-28) (Hypophthalmiehth-42) (Hy potr iorchis-248, 309 , 323)
(Hyrax-538.573.583.641) (Hystrix-340.554.558.566.639.655.679)

(Ibis-204 ^217. 237. 272. 284) ( Ichnotropi s- 1 80 ) (Iehthyophaga-239) ( I ctalur us-9.43, 51 ,53,73,238)
(Ieter us-259 . 260. 314 ) (Ictiobus-238) (Ietonyx-639.663) (Idotea-26) (Idus-10.66) (Iguana-133.138,
139,146,148,149,191,652) (Ilisha-28) (Iltis-528) ( Inimicus-76 ) (Irrisor-250) (Isoodon-424,507.650)
(Ithagenes-234) ( Ithaglnls-224 )

(Jaculus-544) ( Japalura-1 52 . 165) (Johnius-36) (Junco-199,314)

(Kaehuga-158. 159. 183) (Kaloula-88) (Kannabateomys-483 , 490 . 656) (Kaupifalco-263) (Ketupa-270)
(Kinosternon-158. 174.184) (Klttacincla-261 ) (Kogia-589.676)

(Labldesthes-43) (Labrax-38.77) (Labrus-27 , 49 )

(Lacerta-88 ,91 ,124,126,139,140,143,154,175,176,180,

184

THE TEXAS JOURNAL OF SCIENCE

NEMATODA, cont.

189,196) (Lachesls-130.162.178) (Laemargus-35) (Lagenorhynchus-519.585.586) (Lagidlum-650)
(Lago£us-201 ,203,210,222,223,231 ,233,234,21)8,311 ,326,330) (Lagostomus-449) (Lagothrix-539.62H,625.6i)9)
(Lama-3110) (Lamna-81) (Lampr is-590) (Lampronessa-223) (Lampropeltis-85. 121 . 128.130.163.176,179)
(Lamprotornis-312) (Lan lus-243 . 254 , 260 , 262 , 263 . 268 , 286 . 305 . 307 . 31 1 . 31 *) . 608 ) (Larus-201 .202.203.209.237.
238,239,240,242,264,269,270,272,273,274,281 ,291 ,297,302,324,330) (Lasiorhlnus-384 . 390 ) (Laslurus-565 . 604 )
(Lateolabrax-76) (Lates-48,53) (Laticauda-129) (Latridopsis-10.53) (Lebius-590) (Leggada-479.482.599)
(Leimadophis-123, 158) (Leiocassis-39 ) (Leioloplsma-96, 126, 178, 179.180) (Leiostomus-53 ) (Leipoa-232)
(Lemmus-632,679) ( Lemni scorn ys-4 82 , 599 ) (Lemur-541 ,569.673) (Leon toe ebus-569 , 650 , 656 ) (Lephuromys-569 )
(Lepibema-43) (Lepldocephalicht-43 ) (Lepidogrammus-327) (Lepidopsetta-76 , 77 ) (Lepilemur-460,650)
(Lepisosteus-69,238) (Lepomis-27 ,51 ,73.238) (Lepor inus-1 0, 44 ,45,46,49) (Leptocephal us-50) (Leptocottus-29)
(Leptod actyl us-85 ,86,88,89,90,94,95,102, 106,110,112,116,117,158) (Leptodira-1 30. 164)
(Leptonychotes-587 , 588 ,590.625) (Leptopel ls-95 ) (Leptoptlla-211 ,212,213) (Leptoptilus-218.272.290)
(Lepus-332, 335, 340, 341 ,343.348,415.416,417.433,447.457.458,459.483.496,497,543.548,549.550,558. 571 , 613, 640,
663,672,673) (Leucichthys-9 . 69 .73) (Leuciscus-9 . 10 . 30 . 35 . 36 . 42 . 48 . 66 . 72 , 77 . 21 8)
(Leucopareia-215) (Leucophaea-261 ) (Leucophoyx-278 , 290 ) (Leucosarcia-223,234) (Leucosomus-66 ) (Lichanotus-460)
(Llmanda-76) (Llmax-4 95 , 499 , 506 , 509 , 51 0 . 525 . 52885 . 89 ) (Limnodynastes-91 . 93 . 1 1 4 , 1 17 , 9~3 ) (Liobagrus-67 )
(Llolaemus-143) (Llopeltis-121 . 176) (Liophi s- 1 23 . 163 ) (Liparis-27,59) (Liponyssus-654) (Lissemys-174)
(Llssotr 1 ton-86 , 88 ) (Litargus-267 ) (Lnphius-27) (Lo-45) (Lobivanel 1 us-286 ) (Lobodon-587 , 588 ) (Loncheres-655 )
(Lophlus-1 5 , 27 , 28 , 29 , 33 , 37 , 48 , 49 , 81 ) (Lophoaetus-236) (Lophoceros-250 ,263) (Lopholatilus-49 )
(Lophophor us-222, 223) (Lophopsetta-53 ) (Lophortyx-203 . 230. 231 , 325 ) (Lophorus-224 ) (Lophura-223 . 224 )

(Lophuromys-630) (Lor is-569 )

(Loxodonta-387. 391 .395.581 ,617)

( Lumbricus-242 , 287 , 342 , 344 , 345 , 51 4 , 533 )

(Lutra-345 , 357 , 358 , 528 , 586 , 587, 629.637)

(Lycaon-524 , 577 ) (Lycod lchthys-59 )

(Lymnaea-174 , 500) (Lynx-516,577,579,598,601,627,636,662,663)

(Lyrur us-200, 201 ,203,204,205,210,223.234,242,248,266,326) (Lystrophi s- 1 63 )

(Lota-9 ,27,35,50,63,66,70,73,218,219) (Loxodon-650 )

(Lucilia-209) (Lucioperea-21 .69.218) (Lullula-31 4 )

(Lumpenus-26 ) (Luscinia-242, 260,296 ,311) (Lutianus-36 )

(Lutreola-335, 344, 345, 431 ,510,524,528,620,621 ,639) (Lutzia-659)

(Lycodon-1 80 ) (Lycodontis-50) (Lygophis-178) (Lygosoma-181 )

(Lyr iocephalus- 151)

(Habuya-140, 142, 151 ,153,173,180,196) (Macaca-650 , 663 ) ( Hacacus-335 . 336 . 340 . 349 , 359 . 361 , 394 , 395 , 398 ,
416,461,601,611,613,614,628) (Macquaria-29) (Macrelaps-129) ( Macroehelys- 1 58 ) ( Hacrochl amys-496 , 503 , 504 )
(Macrocyclops-195) (Macrodipteryx-230) ( Macrohectopus-587 ) ( Macronectes-239 , 274 , 285 ) ( Macrones - 2 3 , 24 )
(Macro pod us-4 44) ( Macropus-359 , 369 , 382 , 385 , 388 , 392 , 399 , 400 , 40 1 ,403,404,405,406,407,403,
409,410,419,424,444,640,649,651,652,664,674) ( Macropygia-233 ) ( Macrorhinus-586 ) ( Macrorhynchus-590 )
( Mac rose el ides-565 ,614) (Mac rose incus- 149) (Macrotus-471 ,655) ( Macrozoarces-29 ) (Macrurus-34 )
(Malacoclemmys-183, 184) (Malacomys-478,483,599) (Malacoptila-231 ) (Malapterus-9 , 31 ) (Malocoptila-231 )
( Mai ur us-3 12 ) ( Manatus-582 ) (Mandrillus-665) ( Manis-37 1 . 492 , 597 ) (Mansonia-650,659) (Mansonioides-650,659)
(Mar eca-204, 215.289) ( Margar iscus-66 ) ( Marmosa-340 , 560 , 563 , 626 , 628 , 658 ) (Marmota-343,416,433,
446,540,543,576,605) ( Mar tes-335 . 340 . 342 . 344 . 345 . 373 . 43 1 . 51 0 . 524 , 528 . 530 . 535 , 575 . 579 . 599 . 627 , 63 1 , 639 , 678 )
(Mastacembelus-43.58.73.74) ( Mastomys-481 .554,599.605.630) (Mayleus-15) ( Mazama-383 , 44 1 , 451 , 459 , 491 , 492, 607)
(Mccullochella-10,29.39) (Megaceryle-240, 308) ( Megaderma-631 ) ( Megadyptes-239 ) ( Megalobatr achus- 1 10)
(Megaloperdix-224) (Megalopterus-239) (Megalot is- 361 ,571,624) ( Megaptera-676 ) ( Megascops-243 ) ( Megophr ys- 1 05 )
(Melanerpes-292 .313) (Melanitta-203) (Melanogrammus-29) ( Me 1 ano pi us-268 , 290 , 292 )
(Meleagr is- 199 .200. 201 .203.204.210.222.223.224.229.234.235.252.261 .289) (Meles-342,
345,348,372,373,498,510,538,575,579,628,636,639,643) (Mellchthys-29,49) ( Mel ierax-263 . 297 ) (Mellivora-373,
639) (Melomys-623) (Melopelia-234 ) (Melursus-362 , 578 ) (Menidia-36,238) (Menura-242) (Mephitis-335,

362.367.443.506.510.528.575.596.614.625.626.628.639.678) (Merganser-200.219) (Mergellus-219) (Mergus-12,
203,217,218,219,237,241,264,270,289) ( Mer iones-483 , 61 3 . 625 , 649 , 61 1 ) (Merlangus-72) ( Merluccius - 1 0 , 27 ,
35,47,590) (Merops-252 , 280 ,282.313. 327 ) (Merula-202, 242,287) (Mesocr icetus-340 , 453 , 605 , 608 , 61 3)
(Mesocyclops-43.619.620.621 ) (Mesomys-371 ,482) ( Mesopus-28 , 29 , 69 ) ( Metachirops- 342 . 34 3 , 344 , 358 , 475 , 560 , 6 1 4 )
(Metachir us-340. 442, 475. 560. 563. 626. 628) (Metopldius-271 ) ( Microcarbo-2 1 9 , 238 , 264 ) (Microcebus-630 , 637 )
(Microdactylus-231 ) ( Microhyla-85 , 95 , 97 ) (Micropogon-46 , 49 . 50 , 203 , 263 ) (Micropterus-9 , 27 , 43 , 51 , 66 , 73 , 238)
(Micropus-202 ) ( Microsarcops-28 1 ) ( Microscel is-261 ) (Microtus-340,
343,344,453,454,455,482,484,544,553,568,599,613,630,631) (Micrurus-96. 128) (Midas-621) ( Milax-5 1 0 , 527 , 528 )
( Mil vus-202, 240, 241 .242.248,263.620) ( Min iopter us-344 , 346 . 467 . 470 . 47 1 , 654 ) ( Mirounga-372 , 586 , 587 , 588 )
(Misgurnus-239.608.619) (Mitu-256) ( Mogera-332 . 450 . 485 ) ( Mogurnda- 1 0, 39 , 42,72 , 73 ,77 ) ( Molge-88 , 89 )
(Molossus-342) (Molva-35.47.73.76.590) (Momotus-259) (Monacha-495 , 499 , 506 , 525 , 527 ) ( Monachus-586 , 587 , 588 , 590)
(Monasa-231 ,306) (Monax-576) (Monodon-520.585.590) (Monopterus-619) (Monotaxis-45) ( Mont icol a-202 , 260 , 296 , 312)
(Morellia-635) ( Morenia - 1 5 8 , 159) (Mor ica-594 . 624 ) ( Morone-38 , 51 , 52 , 73 ) (Morphnus-300) (Moschus-502, 645)
(Motacilla-209.242.260.266.292.311.312.313) ( Moxostoma-67 ) ( Mugil-30 . 35 . 38 ) ( Mul lus-27 . 29 , 49 ) Mungos-256 , 635 )
( Mur aena-48 ) (Muraenesox-27 . 28 . 33 . 56 ) (Murenophis-49 ,70)
(Mus-342.343.344.345.346.348.361 ,455,461,479,483,484,486,540,544,548,549,553,566,569,575,599,600,612,613, 623,
625,626,627,629,630,631,654,662,678,679) (Musca-209.597.635,636) ( Muscard inus-629 ) (Muscicapa-260,
267,330) (Musicapa-300) (Muscisaxicola-313) (Mustela-335 , 343 , 344 , 345 , 348 , 43 1 , 528, 535 , 625 , 639)
(Mustelus-26.35.37.56.57) (Mycetes-362,525,615) (Mycteria-200) ( My cteroperca-36 , 50 ) ( Myd aus-625 , 626 )
( My lagra-262 ) ( My iozetetes-308 ) (Myletes-15) ( My 1 iobatis-55 ) ( Myocastor-335 ,340,445,433,569,640,656)
( Myomys-479 ) ( Myonax-524 , 602 ) ( Myophonus-3 1 2 ) (Myopot-amus-335 . 340,343) (Myoprocta-643)
(Myotis-202, 343. 344. 346. 347, 437. 466, 470. 565. 626. 630. 633. 653 .655) (Myoxus-344 ,453) (Myrmecophaga-469,490,

625.626.653.678) (Mystus-22 . 24 . 46 . 47 . 74 ) ( My ther a-300 ) ( Myxocephal us- 1 0 , 30 , 61 ) (Hyzantha-301 ) (Myzomela-261 )

INDEX: SYSTEMA HELMINTHUM

185

NEMATODA, cont.

(Naja-1 21, 129, 130, 162, 163, 164, 182, 184) (Nandlnla-630) (Nannoperca-39.219) (Nasalis-397) (Nasilio-626)
(Nasua-335, 361. *13 1,53*1.579. 625. 627, 662, 663) (Natalus-344 , 472 , 633) (Natrix-100, 109, 121 , 122, 128,129,130,
162,163,166,167,177,178,182,18*1,195,196,620,621 ) (Necrosyrtes-256) (Nectarinia-287) (Nectomys-554 , 62*4 , 65** )
(Nectophrynoides-99 ) (Necturus-88,219) (Nemachilus-30.66.72) (Nematonurus-33 ) (Nematophila- 1 3*1 )
(Nemorhaedus-339,340,342.397,416,417,434,435,436,440,451 .458,495) (Neobythites-4 1 ) (Neocellia-659)
(Neomaenis-50 , 53 ) (Neomerls-519,521 .676) (Neomys-344 , 3*15 , 505 , 589 , 61 8) (Neophoca-587) (Neophron-249)
(Neotls-231 ) (Neotoma-339, 340, 4*15, 458. 459. 483, 545) (Neotomys-625) (Nerophius-27 ) (Nesocia-343 ) (Nettapus-217 )
(Nettlon-201 ,214,271 ,289) (Nicoria-106, 158) (Ninox-205 , 278 , 294 , 307 ) (Nisaetus-242, 278) (Nisus-277)
(Noctilio-472) (Noctura-230) (Nonnula-231 ) (Notechis-1 1 4 , 163, 177 ) (Notemigonus-9)
(Nothura-203,233.250,257,315) (Notipsar-162) (Notophoyx-238) (Notophthalmus-1 12) (Notopoeon-30,43)
(Notopterus-61 ,73.74) (Notothenia-29.52 , 59 . 586 , 590) (Notropis-36 , 43 , 66 ) ( Noturus- 4 3 , 7 3 )
(Nucifraga-204,209,219,242,314) (Numenius-217,218,219,280,281 ,284) (Numida-200, 201 ,204,209,210,
222,223,228,229,2.31,232,233,234,235,241,248,264,271) (Nutrla-335) (Nyctaea-243) (Nyctala-204 )
(Nyctalu3-345.346.470) (Nyetea-292) (Nyctereutes-335 . 34 1 . 342, 362. 371 . 510 ,577 , 580 . 635) (Nyctiardea-239)
(Nyctibeus-231 ) (Nycticebus-359 , 362 . 54 1 , 63 1 , 652 , 659 ) (Nycticeius-345.468,654) (Nycticorax-200,218,219.
225,236,238,239,240,242,243,256,262,277,278,289,290,304) (Nyctidromus-260 , 26 1 , 322) (Nyctinomus-345,437)
( Nyroca-20 1 , 203 , 204 , 264 , 274 , 289 , 290 )

(Oceanites-285 ) (Oceanodroma-275 ) (Ochotona-4*l8,486,545,548,550) (Ocnera-231 ) (Octolasium-242)
(Ocyurus-36) (Odobenus-587 . 590) (Odocoileus-340 , 383 , 397 , 416 , 430 , 434 , 435 , 436 , 440 , 451 , 457 , 458 , 496 , 500 , 501 ,
556,614,644,670) (Odontobut i s-6 1 9 ) (Odontophorus-2Q5.223.231 ) (Oedaleus-267 ) (Oed icnemus-231 , 242 , 254 )
(Qedipomidas-541 ) (Oedura- 140) (Oenanthe-314) (Ogcocephal us-29) (Ogmorhinus-587 ) (Qidemia-200 ,
203,214,215,264,272,284,289) (Okapia-339 , 356 , 396 , 422, 435 , 451 , 480 , 532 , 61 4 , 617 , 650) (Ommatophoca-587,588,590)
(Qnchidium-509 ) (0ncorhynehus-27 ,28,29,30,39,41,61,67,69,76,79,238) (Ondatra-343. 344,416,455,481,
482,509,624,625,631,661) (Onitis-613) (Onos-27 , 39 , 59 , 590) (Onthophagus-594 . 604 , 608 , 61 3 . 61 4 )
(Onychodactyl us-86 ) (Onychogale-404.650,651 .674) (Onychomys-599 , 631 ) (Ophiocephalus-42,43.44,45,46,64,
73,76,619,620) (Ophiodes-178) (Ophiodon-29,48) (Ophis-178) (Ophisaurus-179, 195) (Opladelus-238) (Opsanus-28)
(Opsarlichthys-36,42,77) (Orchestia-267) (Oreamnos-430,435,497,556) (Orectolobus- 10,37) (Oreinus-67)
(Oreotragus-645) (0restias-4 1 ) (Or iocincla-242) (Oriolus-250, 260, 266 , 267 , 293 , 301 , 31 3, 323 , 327)
(Ornithodorus-649 , 650) (Ornithonyssus-655) (Or thagorisc us-49 ) (Ortyx-222 , 234 , 247 , 31 4 ) (Or yc ter opus- 362 , 639 )
(Oryctolagus-335.340,343.415,416,433.447.458.497.543.613) (Oryx-416) (Oryzomys-63 1 ) (Osmerus-26 , 34 , 68 )
(Qssifraga-201 ) (Otar ia-372 . 586 , 587 . 588 ) (Qtis-201 ,210,212,222,
223,224,229,230,254,260,261 ,282,296,297,299,306,330) (Otocampsa-31 1 ) (Otocolobus-598 ) (Otolicnus-569)
(Qtoinela-260, 307 , 321 ) (Otomys-373.492.623) (Otopterus-346 ) (Otospermophilus-543) (Otus-242.243,256,
261,270,277,302) (Ourebia-340,422,435,436) (Ovibos-435 , 451 , 459 ) (Ovis-336 , 339 , 340 , 34 1 , 34 2, 34 4 , 397 , 4 16 ,
417,419,420,421 ,422,423,430,432,434,435,436,438,439,440,441 ,451 ,457,458,459,480,495,496,497,501 ,502, 512, 556)
(Oxanna-646 ) (Oxya-26?) (Oxydema-267 ) (Oxyrhopus-122, 178, 196)

(Pachydactylus-142) (Pachypti la-274 ,285) (Pagellus-76) (Pagrosomus-29.52,53.55) (Paguma-366.631.663)
(Pagurus-59 ) (Palamedea-210) (Pal inur ichthys-38 ) (Palomas-234 ) (Palorus-267 ) (Pan-541 ,651 ,666) (Pandalus-59 )
(Pandion-240, 241 .275.300.620) (Panopl ites-650) (Panthera-531 ,601 ) ( Papio-335, 395 , 397 , 538 , 54 1 , 61 1 , 626 , 666 )
(Parachaenichthys-590) (Parachondrula-499 ) ( Farad i sea- 3 1 2 ) (Paradoxurus-366 ,431,631)
(Paral ichthys-27 ,29,52,53) (Parameles-626 , 632 ) (Parapr isti poma-76 ) (Parascalops-589 )
(Parasilurus-39.42,45,77.78.620) ( Par axer us-554, 569. 599. 623) (Parcoblatta-630) (Paroaria-330) (Parus-291 ,
311,314) (Passalus-604,608) ( Passarel la-202 ) (Passer-200,201 . 268 , 270 , 27 1 , 31 4 . 324 , 608 ) ( Passer ina-294 )
(Pastor-272,313) (Paulicea-49 , 50 ) ( Pavo-222 , 223 , 224 , 227 , 230 , 234 , 235 , 252 , 261 , 268 , 27 1 , 290 ) (Pavoncella-201 )
(Pecari-437.607) (Pedetes-396 , 549 , 557 ) (Pedioecetes-223 , 224 , 231 , 234 , 252 , 253 , 260, 268) (Ped iotragus-451 )
(Pelagodroma-274) (Pelamys-42) (Pelea-4 34 , 45 1 ) ( Pelecanoides-285 ) ( Pelecanus- 1 2 , 202 , 21 8 , 237 , 238 ,
239,240,262,269,270,271,278,290,293,303) (Pelecus-10, 30, 43 , 66 ) ( Pelobates-84 , 88 , 94 , 99 , 100, 146) (Pelomys-431 )
(Pelteobagrus-73) (Penelope-235,256) (Peragale-570,626.627) ( Per ameles-341 .570.626,632) (Perca-9,
26,30,35,42,43,51 ,52,70,72,73,76,218,219) ( Percalates-1 0, 29 , 39 , 73 .77 ) (Percina-43) (Perdix-200,201 ,
203,204,205,210,222,223,224,225,229,231 ,234,252,271,290,300,326) (Per iplaneta-594 . 599 , 601 , 61 3 ) (Pernis-241 ,
242,263,291) (Perodicticus-432) ( Perognathus-34 1 ,558,599,600) (Peromyscus-34 1 , 342, 455 , 482, 486 , 544 , 599 ,
545,554,613,630,631) ( Peroryc tes-626 ) (Petaurista-483.657) (Petauroides-538,539,543) (Petrodromus-572,673)
(Petroeca-296) (Petrogale-385 . 399 . 400 . 401 .406.409) (Petromys-540,559) ( Petromyscus-559 ) (Petromyzon-50,81 )
(Petrosaurus-137) ( Phaeochoer us-386 . 387 . 389 , 394 . 395 , 396 . 397 . 552 . 575 . 625 . 645 ) (Phainopepla-322)
(Phalacrocorax-45. 200. 201 .202. 204. 209 .21 8. 21 9. 237. 238 .239. 240. 241 .242 .258. 259 .264. 270 .275. 276, 290, 317, 321)
( Phanaeus-253 , 608 ) ( Phaneroptera-267 ) ( Phaps-234 ) ( Phascogal a-603 ) ( Phased omys- 389 , 395)
(Phasianus-200, 201*203, 204, 205, 21 0,222, 223, 224, 225, 231, 234, 252, 260, 271) ( Pheniscus-209 ) (Pheretima-5 1 4 )
(Phi lander-340. 429. 464. 560. 563. 569. 570. 620. 628 .658) ( Philantomba-646 ) (Philemon-261 ) ( Phi lodryas- 1 22 , 1 30 , 187)
(Philomachus-209.272) (Philypnodon-39.218) (Phimosus-217) (Phlogoenas-233) (Phoca-513,
528,585,586,587,588,590,657) (Phocaena-519.520.522.585.586) (Phocaenoides-519,520) (Phoebetria-239 , 274 )
(Phoenicopter us-242. 243 .272. 289. 303) ( Phol is-27 . 76 ) ( Phox inus-29 . 36 , 66 , 72 ) ( Phrynobatrachus-94 )
(Phrynoeephalus-176 . 177) (Phrynosoma-178) (Phyeis-47 .73) (Phylan-247,250) (Phyllodactylus-140)
(Phyllomedusa-88 ) (Phyllosoma-678) (Phyllostomus-465) ( Physeter-585 , 586 , 677 ) (Physiculus-10)
(Physignathus-177. 178. 193) (Piaractus-15.20) (Piaya-229,230,252) (Pica-202,204,243,260,266,267,292,
301,314,315,318,321) (Picus-204 , 248. 256 . 268 . 300. 306 . 307 , 31 3 ) (Pimelia-594 ,605) (Pimelodella-66,73,74)
(Pimelodus-15.49.76) (Pimephales-36) (Pionus-233) (Pipile-256) (Pipilo-271 ) (Pipistrellus-345.633) (Pipra-330)
(Piranga-204) (Pitangus-294 ) (Pithecia-557 ) (Pithecus-336, 54 1 , 569 , 61 1 ) (Pitta-256) (Pituophis-128, 176)

186

THE TEXAS JOURNAL OF SCIENCE

NEMATODA, cont.

(Pitymys-453) (Planesticus-209.242.260,271 ) (Platalea-218.270.289) (Platichthys-76.77)
(Platycephalus-28,55.59) (Platyclchla-293) (Plecoglossus-36) (Plecotus-470.624)
(Plectroplites-10,29.39.59.73.77.218) ( Pleetropterus-291 ) (Plestiodon-140) (Plethodon-85.86,89.96, 101)
(Pleuronectes-26 ,28 .30.31.35,47.48.52.76. 590) (Pleuronichthys-52) (Plocepasser-254) (Plotus-230.237,
239,240,241) (Podager-231 ) (Podarcls-196) (Podargus-229) ( Pod iceps- 12. 203. 204, 21 8, 21 9. 237, 240, 24 1 .
262,264,269,270,272,273,289,301 ,305,619) (Podinema-178, 196) (Podoenemi3-134. 183, 185) (Poeelle-312)
(Poecilictis-637) (Pogonomyrmex-178) ( Pol loaetus-248 ) ( Pol iocephal us-240) (Polyborus-261 ) (Polygyra-96 )
(Polymltarcys-6 1 ) (Polyodon-238) (Polypedates-85 . 91 ) (Polyplectron-222, 224 .225, 247 .255 . 327) (Polyteles-234 )
(Pomatostomus-204,294) (Pomox Is- 1 2 , 43 . 73 . 21 9 . 238) (Pongo-438,541 .663) (Porcell lo-270, 27 1 ) (Poronotus-38)
(Porphyr io-263 ) (Potamochoer us-646) ( Potamogale-678-) (Potamon-620) (Potomophls- 1 22 ) (Potorous-404 , 444 )
( Potos-4 3 1 ,510) (Praomys-345 ,478.483.554.569. 599 ) (Prionailur us-605) (Pr lonoeypr is-278) (Pr ionops-322)
(Pr istls-37 ) (Prlstiurus-26) (Probrevleeps-109, 167) (Procapra-430) (Proeavia-392.573.583.627.645)
(Procellar la-275) (Prochilodus-72) (Procyon-348 . 361 . 367 . 371 ■ 372. 431 .
510,575,578,618,621,625,627,628,650,663,678) (Prodelphinus-586 ) (Proechimys-475.477.483.488)
( Promicro ps-28 , 3D (Promops-468 ) (Propithecus-663) (Prosopium-49.73.79) (Protogeris-235) (Protopsetta-76)
(Protoxerus-479) (Psammodromus-180) (Psammophis-1 30 . 162,164,176,178) (Psammosaurus-187) (Psephurus-39)
(Pseudacris-85.89.96.99. 106) (Pseudaphritls-39.67 .218) (Pseudaspls-164) (Pseudaspius-77) (Pseudaxis-398.418)
(Pseudechis-1 14,163.177) ( Pseud emys- 158 .174.184) (Pseudobagrus-45) ( Pseudoboa- 1 61 ) (Pseudocordylus- 143)
(Pseudogyps-248) ( Pseudois- 434,436) (Pseudolabr us-55 ) (Pseudomy2omyia-659) (Pseudophls-106, 158, 178)
(Pseudoplmelod us-50) (Pseudopl at 1 stoma-49) (Pseudopsittacus-324) (Pseudopus-176) (Pseudorhombus-46.52)
(Pseudoscaphirhyn-67) (Pseudotantalus-240) (Pseudotriton-100) (Pseudotropius-73) (Psl ttacula-235)
(Psittacus-233.235.257.300.330) (Psophia-223,224) (Pternistes-223. 225. 231 .254.267.319) (Pterocles-221 )
(Pteroclur us-221 ) ( Pterogal e-65 1 ) ( Pteroglossus-205 ) (Pteromylaeus-34) (Pteromys-558) ( Pteronura- 587,663)
(Pterophyllum-10) (Pteropus-580.679) (Ptilopachys-268) (Ptiloscelis-297) (Ptiocichla-311 ) (Ptyas-184, 123)
(Ptychocheil us-67 ) (Ptychodactylus-91 ,139,140) (Ptychozoon-183) (Puffinus-238.270.274) (Pulex-650,651 )
(Puntius-42) (Pupilla-496) (Putor ius-335 . 342 . 344 . 345 . 348 . 372 . 431 . 51 0 . 524 , 528 . 535 , 575 , 627 , 639)
(Pycnonotus-31 3, 324 ) (Pygocentrus-45, 46) (Pygodactylus-178) (Pygoscells-238 , 285 ) (Pyromel ana-229 .313)
(Pyrrhocor ax-266, 312) (Pyrrhura-233,297) ( Pyrrhurus-322)
( Python- 1 14, 123, 125,129, 162, 163, 164, 177, 182, 187, 195,406)

(Querquedula-200.215.216.217.271 .272.289.290) (Quiscalus-204 , 256 , 267 , 268 , 27 1 )

(Rachycentron-28) ( Racovitzia-60 ) (Raja-29.32.35.37.57.81 ) (Rallus-281 .284) ( Ramphastos-256 . 257 .
260,270,300,325) ( Rana-1 2 , 84 , 85 , 86 , 88 , 89 , 90 , 91 , 93 , 94 , 95 , 96 , 97 , 99 , 100, 102 , 103 , 104 , 105 , 106 , 107 , 108 , 109 , 1 1 1 ,
112,113,114,115,116,117,118,144,158,162,167,169,174,184,219,619,620,621) (Rangifer-340.434.435.436.440.451 .
459,460,512,556,644,646) (Raphicerus-436.480.556.645.646) (Rasbora-42,43) ( Ratelus-624 ) (Rattus-336,
342, 343, 344, 345, 348, 479, 483, 484, i486, 507, 508, 544, 545, 553, 554, 566, 580, 599, 611, 620, 625, 626, 628, 63 1 )
(Recurvirostra-201 .272,289.290) (Redunca-421 ,422,614,645) ( Regalecus-27 ) (Reighardia-302)
( Reithrodontomys-554 .613) ( Retinella-96 , 506 ) (Retropinna-39.41 .218) (Rhabdomys-479.482.605)
(Rhacophor us-85. 88. 106. 115) (Rhadinaea-1 12. 122) ( Rhamdia-30 , 67 , 168) (Rhamphocenus-300) (Rhea-203,
207,235,249,250,306) (Rheocrypta-43) (Rhinaster-402) (Rhinichthys-9.43,66,238) (Rhinobatus-57)
(Rhinoceros-369. 371 .381.384.386.387.391 .401 .402.538.562.597.607.617) (Rhinoderma-93.94) (Rhlnogoblus-42)
(Rhlnolophus-343.345.346.467.470.624.634.653.654) (Rhinoptynx-307) (Rhlpicephalus-649.650.661 ) (Rhipidura-298)
(Rhizothera-224) (Rhodestethla-238) ( Rhodothraupls-294 ) (Rhodymenichthys-77) ( Rhombomys-34 1 . 545 . 549 , 625 . 654 )
( Rhombosolea-4 1,47) (Rhombus-27) ( Rhyacosiredon- 1 06 ) ( Rhynchotus-230 . 235 . 257 , 4 1 7 ) ( Rhyparobia-599 )
(Richard son! us-9. 66. 238) (Ripar ia-266 , 268 , 31 3) ( Rissa-264 , 281 , 302 , 324 ) (Rita-45,74) (Roccus-38)
(Roll ulus-224 ,227.230.267) (Romerolag us-340, 416, 482, 550) ( Rupicapr a-339 .
417,430,434,435,440,441,451,457,458,495,497,501,527,556) (Ruticllla-201 ) (Rutilus-35, 66,77)

(Sacrophllus-432) (Saiga-420.642) (Saimir i-431 . 482 . 524 . 542 . 61 4 . 650) (Salamandra-88,94,95,99, 100, 146)
(Salmlnus-10. 45. 48.73.76) (Salmo-9 . 14 .26.27 . 31 . 33. 35. 38. 39 . 41 . 42. 43.49 .50.61 .66 .67.69 .70.73.79 . 218,238,590)
(Salvel inus-9 . 10. 12,29,30,35,36,49,60,61,66,67,68,69,76,79,219,238) (Sarcidiornis-209) (Sarcochell ichthy-36)
(Sarcogyps-238 .256) (Sarcophilus-627 ) (Sardlnella-27) (Sardlnius-10) (Sargus-53) (Sarkidlornis-272)
(Saurothera-311) (Sax lcola-268 . 311.312) (Sax icololdes-267 ) (Scalopus-589.625.631 ) (Scaphiodon-20)
(Scaphlopus-85 .89.95.96,99) (Scaphirhynchus-81 ) (Scar abac us-601 . 608 .613) (Scardafella-212,235)
( Sc ard ini us-30. 35. 76. 77) (Scelocnemus-178) (Sceloporus-136, 137. 140, 141 , 152,178,179,190) (Schilbeodes-238)
(Schizothorax-30.67.72) (Sciaena-28,77 ,590) (Sciaenops-27.53) (Scincus-140, 142, 143, 144, 154, 180, 185)
(Sciurotamias-554 ) (Sciur us-335. 336. 340. 342, 362, 4 15, 4 16, 445,
446,476,478,482,538,540,541,554,559,568,569,571,575,599,613,614,625,626,627,630,631,640,654) (Sclerostoma-209)
(Sclerotis-238) (Scolecomorphus-101 ) (Scoliodon-37 ) (Scolopax-218 . 242. 262, 276 , 284 , 291 ) (Scolopendra-209)
(Scomber-12,26,27,29,30,35,42) (Scomberomorus-28.30.77) (Scops-247) (Scopus-229.231 ) (Scorpaena-48 , 81 )
(Scorpaenichthys-61 ) (Scutiger-95) (Scylllna-289) (Scyllium-26.55.57) (Scythrops-229) (Sebastes-27 , 35 , 48 )
(Sebasti sc us-77 ) (Sebastodes-10.27.60.76.77) (Selurus-315) (Selenidera-318. 326) (Semnopithecus-397 .541 .662)
(Semotilus-66) (Sepedon-121 ) (Sephoena-231 ) (Ser lnus-324 ) (Ser lola-28 . 30 .79 ) (Serranellus-77 )
(Ser ran us-4 6. 49. 2 18. 590) (Setonix-444) (Setopagis-260 ) (Sialia-268,313) (Slaphos-126) (Siganus-44 .45,49)
(Sigmodon-343. 37 1.4 17. 481 .486.566.575.599.600.607.624.625.626.631.654) (Sika-358 . 373 . 440,500)
(Silenus-359.613) (Sillaginodes-55) (Silur us-30. 35. 38.43,72,219) (Silvicapra-480) (Silvilagus-417)
(Simia-4 38. 492. 541 .612.624) (Simul lum-326 . 64 4 . 650. 668 . 669 ) (Siredon-106, 112, 157) (Siren-99. 106,169)
(Sitagra-324) (Sitophilus-267 ) (Sltta-261 ,315) (Sminthurus-209) (Solenodon-439)

INDEX: SYSTEMA HELMINTH UM

187

NEMATODA, cont.

(Soniaterla-201 ,203,21A,218,219,26A,271 ,272,290,293) (Sorex-2A2, 332. 337 . 339. 3A3 . 3AA , 3A5 . 3A6 . 3A8 , A5A , A55 . A81 ,
A82.A83, 505, 510, 535, 589, 618, 625, 628) (Sotalia-519,585) (Spalax-566.575.613.61A) (Sparus-60) (Spatula-20A ,
218,289,290) (Speotyto-297 , 306 ) (Sperinophilopsls-263.559,605) (Spermospiza-313) ( Sphenomorphus- 178)
(Spheroides-9,A8) (Sphyraena-36,590) (Sphyrna-26 . 37 , 61 , 69 ) (Spllogale-510.528.575,625) (Spllopel la-23A )
(Spilotes-178) (Spinlperea-30) (Spirontocaris-59) (Spizaetus-256.263) (Splzella-20A ) (Spreo-312)
(Squalius-30,67) (Squalus-10, 35,50) (Squatarola-2A2,2A3,280) (Squatina-3A ) (Steatomys-569.599) (Stegostoma-37 )
(Stellio-1A2, 152) (Stenopelwatus-625) (Stenops-637 ) (Stenorhynehus-587 ) (Stenosaura- 1 38 . 191 )
(Stercorarius-237.270) (Sterna-201 , 203,239, 2A2, 270 , 273 , 280 , 281 , 28A , 305 ) (Sternotherus-105, 17A)
(Stietoenas-235) (Stlzostedlon-9, A3, 51 ,73,76) (Stotnoxys-597 ,598) (Storeria-96, 122)
(Strepera-2A3.260.268.3H ,318) (Strepsleeros-363 , 397 . A22, A51 .669) ( Streptonoura-7 3 )
(Streptopella-212,23A,326,328) (Streptosaurus-IAO, 152) (Sir lx-20A , 223 . 228 , 230 , 2A2 , 2A3 , 2A8 , 260 , 263 , 26A ,
277,278,289,300,307,330) (Stromateoides-76) (Stromateus-27) (Str uthlo-206, 21 0,235, 306, A27)
(Sturnel la-260, 267, 268) (Sturnla-202 ) (Sturnira-3A5 ) (Sturnopaster-313) (Sturnus-201 .202.203.2A2.
267, 272, 287, 311, 313, 31A) (Stylopyga-613) (Subzebr inus-A96 , 50A ) (Succinea-A95.A96.A98. 500,509,510,511 ,528)
(Sudls-39) (Sula-239) (Suncus-3AA , 3A6, 628 . 651 ) (Sur lcata-571 ,575.598.651 ) (Surnia-223.2A3)
(Sus-335, 336 , 3A6 , 3A8 , 358 , 373 , 395 ,A17,A26.A51.A61 ,512,605 , 607,609 . 61 A , 620,621 , 6A0 , 6A5 , 6A6 , 6A7 ) (Sycorax-1 17 )
(Sylvaemu3-A53 ,627) (Sylvia-205,2A2,260,266,268,312,31A) (Sylvicapra-61A,6A5) (Sylvilagus-3AO.3A1.3A3,
A16,A33,AA7,A57,A58,A59,A83,A91 ,A96,A97,5A3,5A9,558,599,663,66A) (Sylvlsorex-61 A) (Symbranehus-81 .218)
(Synaelum-A8) (Synalpheus-579 ) (Syncerus-669 ) (Syngnathus-26 , 30) (Synodontis-AA , A3 ) (Syrmatle us-200, 225)
(Syrnium-20A )

(Tachydromus-620) (Tachyglossus-A 32 ) (Tachyphonus-292) (Tachyptes-239) (Tadarida-3A5.A37.A68)
(Tadorna-201 ,209,217,271 ) (Taeniorhynchus-650 , 656 ) (Taenlura-3A ) (Taius-29 . 30) (Talpa-2A2, 337. 3AA ,
3A6,3A8,A85,A89,535,589,59A) (Talpacota-23A ) (Tamandua-A2A . A25 . AA8 , AA9 . A69 . A81 , A8A , A87 , A88 , A90 , A92)
(Tamias-3AO,3A6) (Tamlasclurus-AA6,A78.55A) (Tanagra-300 . 31 1 ) (Tandanus-10, 39.63) (Tantalus-2 A0)
(Taphozus-A65) (Tapldurus-1A3) (Tapirus-368 , 371 , 387 , A02, 608, 609 ) (Tarentola-91 . 139.1AO. 1A1, 1A2, 1A3, 183. 193)
(Tarsius-569) (Tartaruga-158) (Tatera-505) (Taterillus-55A,631 ) (Taterona-A82 . 55A ) (Tatus-551 ,560,561 ,56A ,651 )
(Tatusia-AA3) (Taurotr agus-396 , 397 , A 17 . A2 1 , A22 , A5 1 . A80 , A85 ) (Tax idea-A 3 1 , 506 . 575 . 625 . 628 , 639 )
(Tayassus-383.A26.A58, 61 2, 662) (Tayra-A31 ,505) (Tchitrea-287 ) (Telmatobius- 1 12) (Temenuchus-261 ,292)
(Tenebrio-292,613) (Tenebroldes-267 ) (Tentyr la-2A7 ) (Terekla-273 . 276 , 280 , 287 )
(Ter rapene-96 , 126, 1 3A , 158, 160, 17A , 179 , 191 ) (Testudo-1 32. 1 3 A , 136, 1A1 , 1A2, 1A3, 1 A A , 1A6, 1A8, 1A9, 155, 156, 157, 159,
167,608) (Tetragonopterus-10.66) (Tetrao-200 , 201 , 203 , 20A , 205 , 212, 222, 223, 231 , 23A , 235, 2A7 , 2A8 , 2A9 , 260, 326 , 330)
(Tetraodon-53,77) (Tetraogallus-223 ,22A , 230. 231 , 235. 303 ) (Tetrapterus-28) (Tetrapter yx-290 )
(Tetrastes-205 , 22A , 23A , 235. 2A8 , 260 , 276) (Tetrax-25A) (Tetrix-267) (Tetusia-551 ) (Teuthls-A6) (Textor-297 , 31 A )
(Thalassarehe-201 ) (Thai assochelys- 156 , 170, 185) (Thalassoma-285) (Thamnophilus-31 1 )
(Thamnophis-85 .121 , 122, 123, 128, 130, 17 A, 176, 179, 195,510) (Thaumalea-223 , 22A ) (Theba-A95.A99.506.525,527)
(Thecadactylus-IAO) (Therapon-10,39,218) (Theropithecus-628) (Thomoniys-339. 3A3 , 356 . A8A ) (Thos-361 .362,629)
(Threpterlus-28,53.59) (Thrlponax-256 , 257 ) (Thryonomys-3A 1 , 358 , 393 , A28 , A83 . A91 , 623) (Thryothorus-271 )
(Thylogale-392.A00.A01 .A03.A0A.A09. AAA. 651) ( Thymal lus-35 . A2 . 60 . 61 . 66 . 67 , 68 . 78 ) (Thynnus-27 . 28 . A2 , 81 )
(Thyrsltes-585,586) (Tilapia-23) (Til iqua-1 39. 1A0. 1A1 . 15A. 177. 181 ) (Tlnamus-20A ,
212, 213, 21A, 223, 225, 231, 235, 256, 266, 268, 31A, 315) (Tinca-30. A2.66) (Tinnunculus-2A 1 ,268,278) (Tipula-209)
(Tityra-260) (Tockus-270 . 31 A ) (Todorna-223 . 26A ) (Tolypeutes-A83.560.561 ,56A) (Totanus-200 , 205 . 26 1 ,
270, 280, 28A, 285) (Trachelotis-250,283) (Trachinus-27 . 39 .76) (Trachops-A72 ) (Trachurus-60)
(Trachysaurus-1A2, 1 A3, 150) (Tragelaphus-3A0.A18.A35.A51.61A.6AA.6A5.6A6. 669) (Traqopan-20A . 223 . 22A . 225)
(Tragulus-397.61A.6A3.6A5) (Trematomus-52.60) (Tr iatoma-661 ) (Tribolium-267.627) (Trichcchus-583)
(Tr lchida-500) (Tr lchiurus-30. 36 . 37 ) (Trichogaster-A3) (Tr ichosurus-A 19 .55A . 599 ■ 651 . 652) (Tridentiger-67)
(Tr lgla-26 , 27.59) (Trimeresurus-123. 128. 129) (Trimerorhinus-1 30) (Tringa-200,209,
215, 220, 2A2, 270, 27A, 276, 277, 280, 286, 609) (Tr ingoides-270) (Trionyx-1 35. 17 A, 185. 195) (Tr iothorus-31 1 )
(Triton-86.91 ,9A. 105. 110. 112. 118. 139) (Triturus-73.86.91 .93.9A. 101 , 112. 113. 139. 17A.620) (Trochalopteron-31A)
(Troehilus-28A ) (Troglodytes-595 ) (Trogodon-300) (Trogon-229, 231 ,252,256) (Tropidolepidura- 178 )
(Tropldonotus-99. 100. 121 , 123, 125, 129, 1A6, 162, 16A , 17A , 176, 182, 195, 621 ) (Tropldurus-123, 1 A 3, 152, 176, 178, 180)
(Trox-608 ) (Trutta-68) (Trygon-3A . 5A .55 ) (Tr ygonorrhlna-57 ) (Trypanocorax-201 .20A.276)
(Tuplnambis-91 , 127, 139, 160, 166, 178, 563, 56A) (Turdus-201 ,202, 203 , 20A , 205 , 209 , 235 , 2A2, 2A3 , 260 , 27 1 , 287 , 293 , 300 ,
311, 312, 313, 31A, 316.32A, 325) (Turnix-231 ,260) (Tursiops-519.522.586.676) (Turtur-229,23A,235) (Tylotriton-96)
(Tympanuchus-223.23A.252.260) (Typhaea-267 ) (Typhl ias-67 ) (Typhlops-130. 16A) (Tyrannus-268. 299)
(Tyto-2A3.2A8.26l.277)

(Uca-67) (Umbra-9 , 1 7 A) (Umbrlna-38) (Upenelchthys-28,60) (Upeneoldes-A8 ) (Upupa-2A7 ,250,286)
(Uraeotyphlus-85 ) (Uraeus-16A, 196) (Uranoscopus-28. 29. 32.75) (Urax-300) (Uria-218, 21 9, 237 , 2A0, 270, 285 , 32A )
(Uroaetus-256) (Urocissa-266 . 31 A . 321 ) (Uroeyon-3A2 . 361 . 510, 577 . 623. 627) (Urogymnus-55 ) (Urolepsis-3A )
(Urolophus-33. 55.57) (Uromastix-92. 1A2. 1A3. 1A8. 177.188) (Ursus-209.373.A51 .575. 578. 6 12. 61 A, 66 A) (Uta-152)

(Vallonia-A96.50A) (Vanellus-201 , 203 , 205 , 2A2 , 2A3 , 26A , 27A , 280 , 286 , 290 ) ( Varanus- 1 25 . 129, 130, 15A, 162,
163,167,170,176,177,178,179,182,187) (Var icorhlnus-67 ) (Vertigo-A96 ) ( Vesper til io-3A6.A70.62A, 630. 63 1 ,
633 , 653 , 65A ) (Vlnago-23A,235) (Vipera-126, 129, 130. 163, 16A. 177) ( Viscacia - A 9 1 ) (Vitr ina-50A )
(Viverra-A98. 579.630) (Viverricula-362.366.630) (Vormela-A31 .636) (Vulpes-336 , 339, 3A2, 3A5 , 3A8 , 372, A61 ,
510,535,570,571,577,579,580,601,605,606,611,620,62A, 625, 627, 629, 63 1,636, 637, 677) (Vultur-2A2.2A9)

(Wallabla-A00.A03.A05.A09. 587) (Wallago-A5 .73) (Wallagonia-37 )

188

THE TEXAS JOURNAL OF SCIENCE

NEMATODA, cont.

(Xanthopygus-441 ) (Xanthornus-323) (Xantusia-1 42 , 1 M3 , 1 4 8 ) (Xenodon-162, 163, 178) (Xenopsylla-600,61 3)
(Xenopus-45, 113) (Xer US-5‘19 . 550 . 554 , 571 . 59*1 , 599 . 62*0 (Xiphias-28) (Xiphocolaptes-296)

(Zacco-42.43.67.77) ( Zacholus- 1 96 ) ( Zalophus-528 . 586 . 587 . 66 1 .662) (Zamenis-123, 128. 129, 143. 163. 164)
(Zanclorhynchus-60) (Zaocys-121 . 129, 130. 179) ( Zapus-570 . 602 ) (Zebra-382) (Zebrias-37) (Zebrina-495.499.527)

(Zebu-4 17.434) ( Zenaidur a-2 1 2 . 243 ) ( Zenobiel la-528 ) ( Zenopsis-30 ) ( Zeugopterus-27 ) (Zeus-26,27,28,29,30,36)
(Ziphius-586,676) ( Zoarces-26 ,27,59.61) ( Zonitoides-504 .511) (Zonotrichia-314) (Zonurus-180) ( Zygonectes-233)

ACANTHOCEPHALA (vol.V)

(Abramis-17.40.42.44.94) ( Acanthurus - 3 3 , 51 ) (Accipiter-1 16, 120, 121 .122) ( Acer ina- 1 7 . 40 , 44 . 46 . 83 . 94 . 96 )
( Ac i pen ser- 3 9, 40, 42, 44, 58, 94, 96) ( Acridotheres-126) ( Adesmia- 1 1 8 ) ( Aeg ial i tes- 1 24 ) ( Aethechinus- 1 37 . 1 39 )
( Agelaius- 1 17,118) ( Agrobates- 1 52 ) (Ajaja-77) ( Alaeops-56) ( Alauda-1 1 6 . 1 17. 152) (Albula-108)
(Alburnus-17.44.94) (Alcedo-156) ( Alectura-1 15) ( Alestes- 1 05 ) ( Alosa- 1 00 . 108 ) ( Amblopl ites- 1 7 . 1 8 . 42 , 58 )
( Ambystoma-48) (Amelurus-18,58,95) ( Ameiva-1 27 ) ( Amia-18,58) ( Ammodytes-39 ) ( Amphacanthus-62)
(Amphimallon-141 ) ( Amphithoe-39) ( Amyda-35) ( Anarchichas-39 ) ( Anas-64 ,71 . 72 , 73 , 74 , 84 , 151 )
( Anguilla- 17. 18. 27. 35. 40. 42. 43. 44. 46. 58. 71 .80.94.97.108) (Anhinga-73) ( An isopl ia- 1 4 1 ) ( Anomal a- 1 4 1 )
(Anorthura-123) (Anser-64,73) ( Anthus-76 , 117) ( Aoria-33) ( Aphelocephala-1 39) ( Aphodius-143) ( Aphriza-75)
(Aplodinotus-25.45.63) ( Apodemus- 1 30 . 1 3 1 . 141 ) (Aquidens-27) ( Aquil a-73 . 123 ) ( Aramides-77 ) ( Ar apaima- 1 09 . 1 1 0 )
(Arctocephalus-83) ( Ardea-72 . 73 . 74 , 75 . 76 . 77 . 11 7 . 1 2 1 . 125.156) ( Ardetta-74 . 152) ( Arenar ia-76 )
( Armadillidium-150. 151 ) (Armides-65) (Artibeus-144) ( Asellus-45 . 46 . 48 . 49 . 64 ) (Asio-122, 125) ( Aspius-44 . 94 )
(Aspro-94) (Astacus-156) (Astur-156) (Asturinula-123) ( Astyanax-31 ) (Athene-123) ( Ather ina-7 1 . 77 . 1 08 )
(Atherinlchthys-47.60) (Attila-127) (Aulopus-87) ( Automol us-90 ) ( Auxis-101 )

(Bagrus-105) (Balaena-86) (Balaenoptera-86,87) ( Bal istes- 1 57 ) ( Barbus- 1 7 . 33 . 46 , 94 ) (Batara-90, 127)
(Baza-121,154) (Belone-100, 108, 157) (Biziura-72) ( Blabera- 1 43 ) ( B1 aps- 1 30 . 1 32 ) ( B1 ar in a- 1 22 )
(Blatella-131 . 143,144) ( Blennius-96 , 156 ) ( B1 icca-17 . 42 . 44 . 80 . 94 ) (Boa-135,136) (Bodianus-103) (Boedon-125)
(Bombinator-48) (Botaurus-72.74.75.76, 152) ( Bothrocara-39) (Box-100) ( Brachymystax-4 1 ) ( Br achyspi za- 1 17 )
(Brama-87. 100) (Branta-71 ) (Brosme-39) (Brycon-40) (Bubo-118, 121 ) ( Bucephal a-7 3 . 74 ) ( Buceros- 1 39 )
(Buf o-4 8, 49, 90, 120, 127,130.132,146.155.156) ( Burhinus-1 17 , 124 , 125) (Busarell'us-135) ( Butastur-1 24 )
(Buteo- 120, 122, 123, 124, 125, 135) ( Butor ides-77 )

(Cacicus-14, 117) (Caiman-110) ( Calamus-1 03 ) ( Cal idr is-7 1 , 77 , 78 , 89 ) ( Cal 1 ionymus-4 3 . 56 ) ( Cal 1 iopi us-39 )
(Callithrix-143, 144, 145) ( Cal lorhinus-80 . 83 . 86 , 87 ) (Caluromys-1 14) ( Campostoma- 1 8)
(Canis-80, 130, 138, 139, 141 , 146, 147, 148) ( Cantharus-76) (Capella-78, 127, 151 ) ( Caranx-87 . 1 00 , 101,110,158)
(Car ass i us-1 7 , 29 . 44 , 47 ,76 , 94 , 100) ( Carchar ias- 1 8) ( Cariama- 1 35 ) (Carina-84) ( Car piodes- 1 8 , 20 , 58 , 95 )
(Caspialosa-108) ( Cassid ix- 1 25 ) ( Castor-73 ) (Catantops-128) (Cathartes-134) (Catla-28)
(Catostomus-T7, 18.20.22.41 .42.46) ( Cebus- 1 4 3 , 1 44 , 1 45 ) ( Centetes- 1 32 ) ( Centonia- 1 4 1 ) ( Centropr istes- 1 1 1 )
(Centro pus- 154 ,155) ( Cepphus-7 1 , 84 ) ( Cerchneis- 1 24 ) (Cercopithecus-1 36) (Certhia-150) (Ceryle-77)
(Chaenichthys-61 ) (Chaenobrytt us-58) ( Chaenocephalus-81 ) ( Chaenogobius-46 , 47 ) (Chaetodon-43)
(Chalamydotis-1 18) (Chanodichthys-67 ) ( Char adr ius-7 1 , 73 , 89 , 90 , 118,121,124,135) (Chaulelasmus-7 1 )
(Cheirogaleus-143) (Chelydra-21 . 156) (Cherax-72) ( Chi idon ias-7 1 ) ( Chond rostoma- 1 7 . 94 ) ( Chor ioti s- 1 1 7 . 1 1 8)
(Chromis-54) (Chrysemys-17.21 ) (Chrysichthys-77) (Ciconia-76) (Cinclosoma-1 39) (Cinclus-72,73) ( Circaetus- 1 27 )
(Circus-120,122,123,124,125,126,130,135) (Cisor-45) ( Ci tellus- 1 30 . 1 3 1 , 1 4 1 ) ( Clangul a-64 , 65 , 7 1 , 72, 73 , 80 , 84 , 85 )
( Clemmys-2 1 ) ( Cl imac ter is- 1 39 ) (Clupea-40,42.47.80,83. 100. Ill ) ( Clupeonella- 1 08 ) ( Cobitis- 1 7 , 3*1 , 95 )
(Coccothraustes-1 17) (Coccygus-127) ( Coelopel tis- 1 23 ) (Coilia-18) (Colaptes-1 18, 135, 151 ) (Colinus-1 17)
(Colisa-35) (Coloeus-121 . 126.127.151) (Cololabis-101 ) (Colonocus-1 41 ) (Coluber- 128, 135, 140)
( Colymbus-64 ,72,74.80.83) ( Compsothl ypi s- 1 4 ) (Conepatus-138, 144, 145, 148) ( Conger-39, 87,158) ( Conopophaga- 1 5 1 )
( Cor ac ias- 126, 132. 135. 151 ) ( Coragypus-1 34 ) (Corcorax-1 17) ( Coregonus-17 , 20, 39 , 40 , 42 , 44 . 45 , 46 , 47 , 80 , 94 )
(Coris-43) ( Corn itermes- 138) (Coronella-122) ( Corophium-95 ) (Corvina-43)
(Corvus-71 , 117, 121 ,.122,124,126,127,135,151,152,157) ( Cor yphaena- 100 , 1 1 1 )
(Cottus-1 7 ,30,39,40,42,58,80,83,94,95,96) (Cratinus-102) ( Crenic ichla-27 , 31 ) (Cricetus-130) (Cristovomer-42)
(Crocethia-65 , 89 ) (Crocidura-122, 128, 129) ( Crotophaga- 1 27 ) ( Ctenophar yngodon- 1 7 ) ( Ctenosaur a- 1 30 , 132)
(Cuculus-89. 124, 127) ( Cur imata- 20) ( Cyclops- 1 14) (Cyclopterus-71 ) ( Cygnus-64 ,71 , 74 ) (Cynoglossus-19,20,43)
(Cy nose ion-96, 109.11 1) (Cyphocaris-39) (Cyprinodon-24) ( Cyprinotus-21 )
(Cyprinus-17,32,40,44,45,46,47,58,94,95,96) (Cypris-21) ( Cystophor a-80 , 83 )

(Dafila-72, 84 ) (Dasylophus-154) (Dasypus- 1 47 ) ( De 1 ph inapt er us- 80 , 81 ) (Delphinus-81 ,87)
(Dendrocopus- 1 18. 156) (Dentex-43) (Desmognathus-48 ) (Diaphoropterus-1 19) ( Dicotyl es- 1 4 1 ) (Didelphis- 147,149)
(Diemictylus-48 , 49 ) (Pi loboderus-1 4 1 ) ( Dinopi urn- 1 21 ) (Diplophysa-32) ( Pi psadomor phus- 1 35 , 136)
(Piptychus-1 9 . 32 . 95 ) (Pitrema-1 10) (Polichonyx-1 19) (Porosoma-25 , 27 ) (Prepanidopsetta-47 )
(Prymobius-126, 127, 135. 136) (Dryobates-150. 151 ) ( Pumetella- 1 5 1 )

(Egretta-72 ) (Elacate-1 1 1 ) (Elanoides- 1 26 ) ( Elaphe-76 . 79 ) ( El aphis- 1 22 ) (Eleginops-56 ) ( El iomys- 1 30 )

(Elopichthys-47) (Elops-19) ( Ember iza-73 , 76 . 1 1 7 , 1 27 ) (Emerita-65) (Emys-21 ,48) (Engraul is- 1 08 )

INDEX: SYSTEM A HELMINTHUM

189

ACANTHOCEPHALA, eont.

(Enhydra-80 ,81,83) (Bpjcometis-I 41 ) (Eplnephelus-103) (Erlgnathus-80.83) (Erlmy2on-18)

(Erinaceus-130, 132, 137, 139, 141 ,143,144,152) (Erionetta-72) (Erisinatura-73,84) (Erithacus-123, 151 , 152)
(Erolia-75,76) (Er ythroeulter-45 , 47,53) (Erythrolamprus-135. 136) (Esox-1 7 ,

18,20,40,42,44,45,46,47,58,80,94,95) (Eucalla-58) (Eueinostomus-77 ) ( Eudromias- 1 24 ) (Eumetopias-80 ,83,86)

(Euphagus-1 17 ) (Euphasia-156) (Eupodotis-1 15) (Eupomotis-58) (Eur ycea-49 ) (Eurynorhynchus-78) (Eutamlas-141 )
(Eutermes-1 38 ) (Euthynnus-104) (Excalfactoria-140) (Exocoetus-100, 108) (Exosphaeroma-56 )

(Falco-116, 120, 122,123,124,126) ( Fel Is- 1 29 , 1 46 , 1 47 . 1 48 , 155 , 157 ) (Florida-78) (Formicivora-151 )

(Francolinus-146) (Fringilla-1 17) (Fulica-64 , 72 ,73 , 74 ) (Fuligula-64 ,71 ) ( Fulmarus-7 1 ) (Fundulus-19,23,75)

(Gad us-39, 40, 42, 43, 58, 84 ,87. 94, 96, 100) (Galelchthys-102) (Galictuis-1 45) (Gallinula-64 ,71 , 151 )
(Gallus-7 1,115,117,143,151) (Gammarus-39 , 40 , 42 . 46 , 49 , 71 ,73,74,95,156) (Garrulus-1 17, 127) (Gasterosteus-17,
39,40,42,46,95,108) (Gempylus-103) (Geothlypis-14 , 1 1 8) (Gephyroeharax-31 ) (Gephyroglanis-77 ) (Geranospiza-1 35)
(Germo-87 ) (Glareola-7 1 . 150) (Glaucldlum-123) (Glaucomys-1 31 ) (Gleoides-100) (Globicephalus-86 )
(Glossogobius-33) (Gnathonemus-1 00 ) (Gnathopogon-47 ) (Gobio-17 , 40 , 46 , 86 , 94 , 95 ) (Goblus-43 , 44 , 47 , 7 1 , 94 , 1 08 , 158)
(Gorsakius-1 53 ) (Graptemys-2 1 ) (Gromphas-141 ) (Gulra-127) ( Gym nocr an i us-96 ) (Gymnopleurus-141 ) (Gypsophoca-84 )

(Haematopus-65 , 89 , 90 . 151 ) (Haemulon-1 03 ) (Halcyon-92,123) (Hal iaetus-74 , 120, 122) (Haliastur-124)

(Halichoeres-69) (Hal iehoerus-80 , 81 , 82 , 96 ) (Harelda-7 1 ) (Harpalus-141 ) (Harpyaliaetus-135) (Heleodytes-1 17)
(Hel io perea-4 5 , 58 ) (Hemerobius-17) (Hemlbarbus-47 , 53 ) (Hemiechinus- 1 37 ) (Hemlgnafchus-1 4 ) (Hemlrhamphus-67 )
(Hemitrlpterus-84) (Herodlas-74 ,77 , 126) (Herpestes-146, 155) (Her petodryas-90 , 127) (Heterospi2ias-123, 127, 135)
(Heterotis-27) (Hexagrammos-46 ) (Hieracidea- 121) (Hllsa-33) (Hlmantopus- 1 24 ) (Hiodon-58 )

(Hippoglossoides-39,41 ) (Hippoglossus-39,82) (Hister-143) (Hoplerythr inus-31 ) (Hoplias-31 ) (Hoplognathus-96)
(Hoplopterus-122, 124, 125) (Houbara- 1 1 5 ) (Huro-42,45) (Hyaena-141 ) (Hyalella-58) (Hyas-72) (Hydrocyon-105)

( Hydromys- 1 55 ) (Hydrurga-81 ) (Hy la-4 8, 90, 120, 123, 127, 128, 154) (Hylacola-139) (Hylocichla-151 ) (Hymenophysa-4? )
(Hypentelium-58) (Hyperoodon-86 ) (Hypnomorphus-1 35) (Hypoedalius-90) ( Hypoedalus- 151) (Hypophthalmichth-17)
(Hyporhamphus-67 ,102) (Hypostomus-26 ) (Hypotaenidia-91 ,140,151)

(Ictalurus-45,58) (Icteria-118) (Ictiobus- 18, 19,20,25,27,58.95) (Inegocla-50) (Isoodon-1 32) (Ixoreus-126)

( Johnius-19) (Jordanella-19)

(Kyphosus-52 )

(Labeo-28 , 33 ) (Labrax-43 , 108) (Labrus-156, 158) (Lacerta-122, 127) (Lachesis- 127,135,136)
(Lachnolaimus-103) (Lachnosterna-141 ) (Lagodon-1 1 1 ) (Lam pro torn is- 139) (Lanlus-72) (Larus-64 , 65 ,
71,72,73,75,76,77,78,80,152,157) (Lateolabrax-103, 104) (Lates-105) (Leiostomus-108) (Lemur-143,144)
(Leontocebus-1 43) ( Leon to pi thee us- 1 45 ) ( Lepidolepr us-87 , 158) (Lepidopsetta-4 1 ,80) (Lepidopus- 1 00 )
(Lepldosteus-58 ) (Lepomis-18,25.58.63) (Leptagonus-39 ) (Leptodactylus-90, 123,127) (Leptodon-123, 135)
(Leptogaster-40 ) (Leptonychotes-81 ,82) (Leptoptila-147) (Lepus-156) (Lerimus-109) (Leuclchthys-20,41 )
(Leue i sc us- 1 7 , 38 . 40 , 42 , 44 , 46 , 47 , 57 , 68 . 76 , 80 , 94 ) (Leucophoyx-77 ) (Leucopternis-127) (Licengraul is- 1 9 )
(Ligyrus-142) (Limanda-39) (Limnobaenus-151 ) (Limnocottus-42 ) (Limnodynastes-123, 154) (Limnopar alls- 152)
(Llmosa-71 ,72.73. 124) (Liocassis-45 ) (Liopsetta-42 , 58 ) (Lipidopus-87 ) (Lobodon-81 .82) (Lobotes-108)
(Lophius-39 , 43 , 87 ) (Lophodytes-72) (Lota-17,18,39.40,41 .42,44,46.80.94,96) (Lotella-40, 41 )
(Lucloperca-40 ,42,44,80) (Lullula-117) (Luscinia- 1 52 ) (Luterola- 1 29 ) (Lutlanus-97 , 111 ) (Lycichthys-39)
(Lycodon-1 28 , 154) (Lynx-141 , 148) (Lystrophus- 135,136)

(Habuia-128) (Hacacus-143) (Hacrorhinus-81 ) ( Halacopterus- 106 ) (Manls-136. 137) (Marila-64,71 )
(Hartes-141 ) (Hedialuna-106) (Hegalobatrachus-48) (Megalurus-150) ( Hegaptera-86 ) ( Melanitta-64 )
(Helanoglaea-4 1 ) ( Hel anogrammus-39 ) (Heleagris-1 15) (Heles-141 ,143) ( Helolontha- 1 4 1 ) ( Helospi za-1 16,118)
( Hen tic ir rhus- 107. 108) (Henura-90) (Mephitis-138,142) (Merganser-83) (Hergus-64 ,71 ,73,75,76,80,84,120,122)
(Mer tones- 130. 132) (Herlangus-94 ) (Merluccius-61 .83.87) (Merula-150, 153) (Hesoplodon-87 ) (Hetachirus-1 14)
(Metopiana-84 ) ( Mi cr acanthus-52) (Micrastur-126) ( Microcarbo-84 ) ( Mi cromesist us-39) ( Mlcropalama-7 3 )
(Micropogon-1 08 .111) ( Micropterus-1 8 . 20. 42. 46 . 58) (Microtus- 1 30 , 1 31 ) (Midas-143) (Milvus-120, 122, 123,124,125)
(Mirounga-8 1 ) ( Misgurnus-47 , 97 ) (Mogurnda-46,47 ,76) (Holge-48) (Molothrus- 1 17) ( Monachus-83 )
(Monochlr us-26 . 43 ) (Monopterus-35) (Montieola-76 . 1 1 9 , 152) ( Morgurnda-76 ) (Horone-58)
(Motaeilla-71. 116. 117.121 ,127) ( Moxostoma-1 8 , 20 ) ( Mugil- 1 8 , 1 9 , 24 , 62 , 156) ( Hus-64 , 1 30 , 1 32 ) ( Muse icapa- 1 50 , 15 1 )
(Hustela-80.82. 129. 139. 141 . 142) ( Mustel us-87 ) (Mycterla-72) (Mycteroperca-82, 103) (Myiochanes-116)
(Myletes-1 8) (Myllobatis-158) (Mylosoma-42) (Myoxocephal us-82) (Myoxus-130. 133) ( Hyrmecophaga- 114,132)
(Mystax-143. 145) (Mystus-35) (Myxocephalus-39,80.82,94)

(Naja-128, 138) (Nandus-35) (Nasua-144) (Natrix-80, 122, 128) (Naucrates-1 1 1 ) (Nemacheilus-17,44,55)
(Heroaehil us-94 . 95 ) (Nematalosa-1 9) (Neobythites-69) (Neogobius-108) (Neomoenis-103) (Neomys-122) (Neophoca-81 )
(Heophron-1 5 ) (Nesokia-1 32) (Netta-73) (Nettion-84 , 1 45 ) (Ninox-121 ) (Nothura-1 18) (Notothenia-58 , 61 , 81 ,82,92)
(Humenlus-71 .75.76.78) (Numida-1 15, 117. 118) (Hyctianassa-65.77.78) (Nyctlcorax-64 ,75 ,77 ,78 , 83 , 125, 152,153)
(Nyroca-64.71 ,72,73,74,83,84,151)

(Qcyurus-103) (0dobaenus-80 , 83 ) (0edicnemus-1 15. 118, 122,124) (Oedipomldas-143, 145) (Oenanthe-76, 118, 152)
(Ogmorhinus-82.83) (Oidemia-64, 71 ,73,84) (0mmatophoea-82) (Oncorhynchus-17 , 39 . 42 , 45 , 46 , 86 ) (Ondatra-73,141)

190

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ACANTHOCEPHALA, cont.

( Ontophagus- 1 4 3 ) ( Onychomys- 131) (Ophiocephalus-39 , 35 . <17 , 127 ) (Opsanus-18) (Opsariichthys-97) (Orca-87 )
(Oreinua-1 9 ) (Oreochichla-1 39) ( Orestias- 157) (Oriolus-121 ) (Orthopristis-1 1 1 )
(Osmer us-90, 92, 58, 80, 82, 83. 85. 156) (Ostlnops-19 , 1 17) (Qtaria-80.81 .83) (Otis-118, 158)
(Qtus-120 . 122. 129 . 127 . 153 ) (Oxyrhophus-1 35 . 1 36 ) (Ox yura-72.73)

(Paehycara-39) (Pachycephala-139) ( Pacu-90 ) (Pagophoca-80) ( Pagrosomus-96 ) (Pagurus-72) ( Palaemon-76 )
(Paludicola-98) ( Pand lon-75 ) ( Pangasius- 103) (Papio-193) (Parabramls-17.95,97,69) (Parachaeniehthys-6 1 ,81 , 82 )
(Paralabrax-102) (Paralichthys-39,50.75 ,111) (Paralinurichthys-109) (Parameles-132 ) (Parapr istipoma-96)
(Parascalops-131 . 192) ( Par asilur us-39 . 35, 91 .95,96.97.97) (Parus-157) (Passer-116,118) (Passerella-90. 118)
( Ped ionomus- 1 39 ) ( Pelamys- 1 06 ) (Pelecanus-80) (Pelecus-9 1,99,99) (Pelteobagr us-90) (Pel torhamph us-55 )
(Penelopldes-1 18) (Perea- 17, 18,90.92.99,96.58.80.99.95) (Perclna-58) (Percottus-99.95)
(Periplaneta-130. 131. 132. 193) (Petromyzon-91 .92) (Phalaerocorax-73 .77 . 80 . 81 . 83 . 89 . 85 ) (Phalaropus-71 .89)
(Phaneus-191) (Phasgale-132) (Phllodryas-135. 136) (Philomachus-73 , 129 ) (Phoca-80.82.83.87) (Phocaena-80,81 .83)
(Phoenieothraupes-151 ) (Phoxinus- 1 7 , 97 , 99 , 95 ) (Phycis-87) (Phyllodromia-131 ,132,191)- (Phyllophaga-191 . 192)
(Phyl lose opus-123) ( Physeter-82 . 86 , 87 ) (Physocorax-92. 125) ( Piaraetus-9 1 ) (Pica-121 , 122, 125 , 126 , 127 , 152)
(Picus-150) (Piplio-1 18,151) (Pitangus-125) (Pitta-151) (Pitymys-131 ) (Platessa-71 .76.77,99,96)
(Platlchthys-9 1 .80) ( P 1 at ycephal us-89 ,111) (Platyeichla-91 . 151 ) (Platydactylus-122) ( Plecoglossus-95 , 97 )
(Plestlodon-193) (Plethodon-98) (Pleuronectes-26,39.90,92,93.80.83.99. 157) (Pleuronichthys-50)
(Plotosus-60,97, 103) (Plotus-71) (Pluvialis-152) (Podargus-123, 129. 159) (Podiceps-69,76) (Poecilonetta-89 )
( Pol lachi us-39 .89 ) ( Pol yboro ides- 129 ) (Polydaetylus-106 ) ( Polynemus-20 , 109) (Polypedates-98) (Polyphylla-19 1 )
(Pomadasys-92. 96 ) ( Pomatomus- 108,111) (Powatostomus-1 39) ( Pomox 13-25 ,58,95) (Pontoporeia-39.92) (Poocetes-1 16)
(Porichthys-97 ) (Porzana-69 ,90,116,152) ( Potamobius-7 1 ) (Potamochoerus-136) (Potos-199 ) (Pratincola-123)
( Prlonotus- 39,111) ( Pristlpoma- 1 00 ) (Prochilodus-18) ( Procyon- 192, 195, 152) ( Psammodynastes- 128)
( Psettodes- 111) ( Pseud aspius-97 .69 ) ( Pseudechls-93 ) (Pseud emys-21 ) ( Pseudobagrus-97 ) (Pseudochloris-1 17)
(Pseudogyps-122) (Pseudopleuronect-39.91 .83) (Pseudorasbora-97) ( Pseudorca-86 ) (Pseudotadorna-7 1,73)
(Pterocles-132) (Pteroglossus-1 19) (Ptyas-129 , 127 , 128) (Pungitius-17, 18,95,58)
(Putorius-80,83. 129. 130.139.191) ( Pygosteus-92 ) (Pyrrholaemus-1 39)

(Querquedula-69.71 .72.89) (Quiscalus-1 17. 151 )

(Rachycentron-1 1 1 ) (Raja-93) (Rallus-71) (Rana- 17, 21 ,98,99,76,77,83,93,95, 120, 127, 192, 153, 155, 156)
( Rasbora-39 ) (Rattus-122. 130.131) (Recurvirostra-71 ) ( Regalecus-87 ) ( Rhad inea- 135) (Rhamphastos-1 19)
(Rhamphocoelus-1 17) ( Rhea-152) ( Rhinogobius-76 ) (Rhinoplagusia-69) ( Rhombus- 9 2 , 80,111) (Rhynchobatus-1 1 1 )
(Rhynchobdel la-39 ) ( Rhyparobla- 193) (Rissa-89 ) (Roccus- 18,39,58,96) (Roncador-108) (Ruplcola-1 19)
(Rupornis-123. 127) (Rutieilla-123) ( Rutilus-90 , 99 , 99 ) ( Ruvettus-87 )

(Sagittarius-123) (Saimiris-193. 195) (Salamandra-98 . 99 ) (Salmo-17 . 18. 39 , 90. 91 . 92 , 99 , 95 , 96 , 87 , 99 , 95 , 96 )
( Saltator- 150) (Sal v el in us-17 . 39.91.92,59.58.80) (Sarcorhamphus-1 39 ) ( Saurogobio- 1 7 ) (Saurothera- 1 27 )

(Sax icola-1 17,119) (Sayornis-1 16) (Scalopus-1 31 ) (Scarabaeus-130, 191 ) (Scardinius- 17 , 92. 99 . 96 , 99 )
(Scarturus-130) (Scatophagus-52.97 ) (Schilbe-105 ) (Schizopygopsis-39 . 96 ) (Sehizothorax-17, 19,20,32,91,95)
(Sciaena-93.80.82. 157) (Sciurus-130. 131 . 191 ) (Scolopax-150) (Scomber-87, 100, 101 , 105) (Sco»hthalmus-9 1 )
(Scorpaena-93 . 157 ) (Sebastes-90 ) (Sebastlchthys-96) (Sebastiseus-69 ) ( Semblls- 1 7 ) (Semotilus- 17,20,23)

(Seriola-100, 105,106) (Serranus-51 .87. 108) (Setlpinna-33) (Sialis-17) (Sillago-96) (Si lurus-90, 92. 99 , 96 , 99 )
(Siniperca-30.39.97) (Solea-93 , 99 , 157 , 158 ) (Solenodon-136) (Somater ia-69 ,7 1 , 72 ,79 , 89 , 157 ) (Sorex-122, 129)

(Spar us-90. 92. 96. 87. 96. 97) (Spatula-71 ,72) (Spermophllus-133, 139) (Spheroides-59 .77 ) (Sphyraena-93)
(Spilogale-129, 138. 196) (Spllornis-129 ) (Squallus-17.95.99,95) (Squatarola-71 , 129) (Steno-86)

(Stenotomus-39. Ill) (Stercorar lus-9 1 ) (Sterna-7 1,77,89) (Stlzostedion-18.20,58) (Strix-120, 122,125,126,157)
(Sturnella-117) (Sturnus-71 .116.117,118.126.151.152) (Sus-191 ) (Sylvia-117.123) (Symbranchus-31 )

(Synagris-100) (Synodontis-22 ) (Synodus-96 ,111)

(Tachytriorchis-135) (Tadorna-69 .71 ) (Talpa-138) (Tamandua-1 19 ) (Tamias-131 . 191 ) (Tanagra-158)

(Tatus-197.199) (Tayra-199) (Tejus-127) (Temenuchus- 1 21 ) (Tenebrio-193) (Tereckia-7 1 ) (Tetrao-69)

(Teuthls-96 , 53 ) (Thamnophil us-90) (Thamnophis-122, 126) (Theragra-39) (Therapon- 1 9 ) (Thymallus-90 ,
92,99,96,156) (Thynnus-87 . 100) (Tilapia-39 ) (Tinea- 17 . 1 8. 99 . 96 , 99 . 95 ) (Tolypeutes-199) (Toxostoma-1 18)
(Trachinus-9 3.71 .80) (Trachurus-90.96.53.96, 105.108) (Trachypter us-87 ) (Trematomus-90 , 58,61,81,82)

(Tr ichaelurus-19 1 ) (Tr ichogaster-39 ) (Trigla-93. 157) (Tr iglops-39 ) (Tringa-71 ,77,90, 116, 150, 151 )
(Tr iportheus-32 . 91 ) (Triton-98,157) (Tr iturus-97 , 98 ) (Tropidonotus-98 , 99 , 120 . 127 , 128, 193) (Tropidurus-127)

(Trutta-99) (Turdus-76 , 89 . 91 .118,129,126,127,150.151.152) (Turnlx-1 15) (Tursiops-81 )

(Tylosurus-18.20.77. 100. 109. 108) (Tympanuchus- 1 16) (Tyto-123 , 127 )

(Umbrina-93,76,82, 108) (Upenichthys-107 ) (Upupa-135) (Uria-71 ,86) (Urocissa-125) (Urocyon-138. 192)

(Vampyrus-131 ) (Vanellus-71 .73.90. 129, 151 , 152) (Varanus-128) ( Var icorhinus-99 . 95 ) ( Verasper-9 1 )

(Vimba-92,99 ) (Vipera-80, 122, 125, 126, 129) (Vul pes- 1 30 , 19 1 , 193 ) (Vultur-118)

(Wilsonia-19)

(Xenarthra-1 38) ( Xenocypr is- 9 7 ) (Xenodon- 135,136) ( Xenopeltis- 136) (Xesurus-53)

(Zacco-19, 21 .97.66.67) ( Zalophus-80 . 82 ) ( Zamenis-122) (Zanclorhynchus-92) (Zebrias-50) (Zebrlus-96)

(Zeus-158) (Zoarces-39.92.99.99.96) ( Zosterops-1 19)

INDEX: SYSTEMA HELMINTHUM

191

LITERATURE CITED

Yamaguti, Satyu, 1958 -Sy sterna Helminthum. Vol. /. The Digenetic Trematodes of Verte¬
brates. Parts 1 and 2. Interscience Publishers, Inc., New York, N.Y., 1575 pp.

- - , \9S9-Sy sterna Helminthum. Vol. II. The Cestodes of Vertebrates. Interscience

Publishers, Inc., New York, N.Y., 860 pp.

- - , 1961 -Systema Helminthum. Vol. III. The Nematodes of Vertebrates. Parts 1 and 2.

Interscience Publishers, Inc., New York, N.Y., 679 and 1261 pp.

— - , 1963 -Systema Helminthum. Vol. IV. Monogenea and Aspidocotylea. Interscience

Publishers, Inc., New York, N.Y., 699 pp.

- , 1963 -Systema Helminthum. Vol V. A canthocephala. Interscience Publishers, Inc.,

New York, N.Y., 423 pp.

EFFECTS OF PROMETRYNE ON RNA AND PROTEIN
METABOLISM IN BEAN LEAF DISCS

by DAVID C. WHITENBERG and JAMES H. FLIPPEN1

Biology Department, Southwest Texas
State University, San Marcos 78666

ABSTRACT

The effects of 2,4-bis (isopropylamino)-6-methylmercapto-s--triazme (prometryne)
on RNA and protein metabolism in bean leaf discs were studied over a period of 7 days.
Prometryne was found to have no effect on the RNA content of the discs or RNA syn¬
thesis as measured by 14C-uracil incorporation during the 7 day treatment period. In
contrast, the protein content of the discs declined significantly by the 5 th day of treat¬
ment, and a significant reduction in protein synthesis as measured by 14C-alanine incor¬
poration was observed at the same time. The data did not reveal a specific site(s) of action
of prometryne on protein synthesis, but did suggest an increased rate of senescence of
the discs, probably due to blockage of the photosynthetic mechanism.

INTRODUCTION

The primary effect of 2, 4-bis (isopropylamino)-6-methyl-mercapto-s-
triazine (prometryne) and other substituted s-triazine herbicides is generally
believed to be on photosynthetic reactions (Gysin and Knusli, 1960). Reduc¬
tion of C02 fixation in plants treated with s-triazineshas been shown, (Ashton,
et al., 1960; Sikka and Davis, 1969; Smith and Bucholtz, 1962; Zweig and
Ashton , 1962) and the effect of these herbicides on the Hill reaction of iso¬
lated chloroplasts has been well documented (Gysin and Knusli, 1960; Good,
1961; Hilton, et al, 1963; Moreland, et al, 1959; Moreland and Hill, 1962).
Although s-triazines unquestionably affect photosynthetic reactions it is pos¬
sible that such effects are secondary in the intact plant, and a close relation¬
ship between protein metabolism and photosynthetic activities has been sug¬
gested (Rhodes and Yemm, 1963). Prometryne has been found partially to re¬
place uracil in E. coli RNA (Moreland, 1967), Singh and West (1967) have
reported that 2-chloro-4, 6-bis (e thylamino)-s- tri azine (simazine) inhibits
RNA and protein synthesis in oat chloroplasts and Beruter and Temperli
(1970) found that prometryne inhibits RNA synthesis before C02 fixation

1 Microbiology Department, University of Texas, Health Science Center, 7703 Floyd Curl
Drive, San Antonio, Texas 78284

Accepted for publication: September 10, 1975

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977

194

THE TEXAS JOURNAL OF SCIENCE

is affected in spinach. Ashton, et al, (1963) reported that 2-chloro-ethyl-
amino-6-isopropylamino-s-triazine (atrazine) severely altered the structure
and morphology of kidney bean chloroplasts, even to the extent of their com¬
plete destruction. McDaniel and Frans (1969) have found that prometryne
inhibits electron transfer and oxidative phosphorylation in soybean mito¬
chondria in a manner similar to oligomycin. However, Mann et al, (1965)
found no effect of atrazine on protein synthesis in barley and Sesbania seed¬
lings, Moreland et al, (1969) reported that atrazine did not affect RNA and
protein synthesis in excised com and soybean tissues, and Truelove et al,
(1973) obtained questionable results concerning an effect of prometryne on
protein synthesis in cucumber leaf discs. Thus the available information is
somewhat contradictory concerning the effects of s-triazines on RNA and
protein metabolism in plant tissues.

This paper reports the effects of prometryne on RNA and protein meta¬
bolism in bean leaf discs.

MATERIALS AND METHODS
Plant Materials

Kidney beans ( Phaseolus vulgaris L.) were planted in flats of sterilized
vermiculite, watered as needed with either tap water or Hoagland’s so¬
lution, and grown in the greenhouse for 14 days. The primary leaves were
excised, washed in deionized water, and discs 1 cm in diameter were punched
from the inverveinal areas of the leaves. Groups of 10 discs (average total weight
of 0.1 25g) were floated on 15 ml of the appropriate aqueous solution contained
in petri dishes. The dishes were placed in a vacuum desiccator, subjected to a
negative pressure of 10 mm Hg for 20 min to infiltrate the tissue, and trans¬
ferred to the laboratory bench under a 24 hr photoperiod of approximately
1,000 lux from cool-white fluorescent lamps at a temperature of 26 C. Five
sets of 10 discs each were frozen for zero time analysis, and 5 replications of
each treatment were removed at 1,2, 3, 5 and 7 days and frozen until analyzed.

Treatments

The basal solution consisted only of water that contained 20,000 units/
liter of penicillin. Solutions used to determine the effect of prometryne
on RNA and protein content of the leaf discs consisted of the basal solution
made to 16.6 //M with prometryne. Synthesis of RNA was studied by follow¬
ing the incorporation of 14C-uracil in discs floated on solutions composed of
basal solution, 16.6 /M prometryne and 1/jlc 14C-uracil (Amersham/Searle,
sa 50 mc/m mole). Solutions used to follow protein synthesis contained 1
H c 14C-alanine (Amersham/Searle, sa 105 mc/m mole). Control solutions
contained no prometryne, but were otherwise identical. Kinetin, which is
frequently incorporated in solutions used in studies of this type, was not in¬
cluded because Goldthwaite and Laetsch (1967) have reported it to be relatively
ineffective in retarding senescence in bean leaf discs, whereas white light was
very effective.

EFFECTS OF PROMETRYNE ON RNA

195

Analytical Procedures

Protein and RNA were extracted by a modification of the techniques
of Barker and Hollingshead (1964) and Osborne (1962). The replicates
of 10 discs were homogenized in a minimum volume of 80% ethanol and
centrifuged at 2000 xg for 5 min. The residue was washed once with 1 ml
of 95% ethanol saturated with sodium acetate, and then suspended in 5
ml of ice cold 0.2N perchloric acid for 10 min. The precipitate was recov¬
ered by centrifugation as before, washed twice with 1 ml cold 0.2N per¬
chloric acid, once with 1.5 ml 95% ethanol saturated with sodium acetate, and
twice with 3 ml of a mixture of 95% ethanol saturated with sodium acetate
and ether (3:1 v/v). The residue was suspended in 2 ml 0.3N KOH, incubated
for 1 hr at room temperature, centrifuged, and the supernatant solution was
saved. The pellet was extracted again and the 2 supernatant fractions were
combined. The residue was reserved for determination of insoluble protein.
The combined supernatants were divided into 2 aliquots, one for determina¬
tion of RNA and the other for determination of soluble protein.

The RNA aliquot (2 ml) was prepared by adding 1.3 ml of IN perchloric
acid and allowing the solution to stand in ice for 10 min. The pellet recovered
by centrifugation as before was washed once with 2.7 ml of 0.2N perchloric
acid, and the washing was added to the first supernatant. The total RNA was
determined by absorption measurement at 260 nm in a Beckman DU spectro¬
photometer. The amount of RNA present was calculated on the basis of a
standard curve prepared from yeast RNA (Sigma Chemical Company).

Soluble protein was determined by the method of Lowry et al., (1951).
The residue was treated with IN NaOH at 100 C for 4.5 min to render the
insoluble protein soluble, and determinations were made as before. The amounts
of protein present in these samples were calculated on the basis of a standard
curve made by using bovine serum album.

The activities of the soluble protein, insoluble protein, and RNA fractions
were determined by liquid scintiallation counting. The counts were corrected
for quenching.

RESULTS

Extraction of RNA

The method of RNA extraction employed in this study effectively re¬
moved all RNA from the tissue as determined by liquid scintillation count¬
ing and absorbance at 260 nm. Extraction for a longer time (Osborne,
1962) or with IN NaOH according to Barker and Hollinshead (1964) gave
greater absorption at 260 nm, but the higher readings were due to gross contam¬
ination of the samples by protein soubilized by the treatments. Corrections for
protein absorbance according to Barker and Hollinshead (1964) gave similar
readings by the 3 methods.

196

THE TEXAS JOURNAL OF SCIENCE

Effect of Prometryne on RNA Content and Synthesis

The data in Table 1 indicate that prometryne had no effect on either RNA
content of the leaf discs or RNA synthesis as measured by the incorporation
of 14 C-uracil. All values are within the standard deviation of the means, thus
no significant differences can be attached to the slight variations.

TABLE 1

Effect of Prometryne on RNA Content3 and 14C-uracil Incorporation^ in Bean Leaf Discs.

Day of
Treatment

RNA Content

1 4

C-uracil Incorporation

h2o

Prometryne

H20

Prometryne

0

133±17

133±17

-

_

1

180±28

166±26

40±8

41±9

2

189±25

175±34

42±7

47±6

3

188+18

216±39

41±7

40±3

5

207±39

194 ±15

39±5

34 ±4

7

149±25

147±11

30±6

32±2

aData are expressed as/Jg RNA/ 10 discs and are the average of 5 replications.
°Data are expressed as cpm/jU RNA, and are the average of 5 replications.

Effect of Prometryne on Protein Content and Synthesis

Protein content or synthesis in bean leaf discs does not appear to be af¬
fected by prometryne until after 3 days of treatment. If the means and standard
deviations as shown in Table 2 are considered, the prometryne treated discs

TABLE 2

Effect of Prometryne on Total Protein Content3 and 14C- alanine Incorporation^

in Bean Leaf Discs.

Day of
Treatment

Protein content

1 4

C- alanine Incorporation

h2o

Prometryne

H20

Prometryne

0

69 3 ±45

69 3 ±45

—

—

1

756±72

862±43

191 ±45

126±32

2

762±75

692 ±8 1

195 ±37

133±20

3

817±74

715 ±72

228 ±50

159 ±18

5

837±70

602±38

170±29

121 ± 2

7

809±50

596±34

173 ±19

106± 9

3 Data are expressed as (Jg protein/10 discs, and are the average of 5 replications.
^Data are expressed as cpm/jltg protein, and are the average of 5 replications.

contained a minimum of 17% less protein at both the 5 and 7 day sample
periods. Although the prometryne treated discs tend consistently to incorporate
less 1 4 C- alanine into protein, again the differences are not significant until

EFFECTS OF PROMETRYNE ON RNA

197

after 3 days of treatment. The data in Table 2 indicate a minimum inhibition
of protein synthesis in the treated discs of 13% at 5 days, and 25% at 7 days.

DISCUSSION

The data indicate that prometryne had no effect on the total RNA content
or synthesis of RNA as measured by incorporation of 14C-uracil in bean leaf
discs under the experimental conditions employed. Such results are not in
agreement with those of Be ruter and Temperli (1970). These investigators
found that total RNA synthesis, as measured by 32 P incorporation in intact
spinach plants grown in solution culture, was reduced within a few hours after
treatment with prometryne. Singh and West (1967) have shown that a reduction
of total RNA in simazine treated oat plants cannot be detected by 6- days
after treatment, but that the synthesis of chloroplastic RNA was reduced by
that time. It is possible, therefore, that prometryne may be more specific in
its action in beans than in spinach, and that minute changes in RNA content
or synthesis in organelles such as chloroplasts could not be detected by the
techniques employed. However, the data of Singh and West (1967) do show
a slight reduction in total RNA content after 7 days of treatment, and a con¬
tinued decline thereafter. Thus the results presented in the present paper are
interpreted to indicate no effect of prometryne on RNA synthesis in bean leaf
discs.

The data concerning protein metabolism are more difficult to interpret.
Several possible explanations exist, but the data do not definitively indicate
whether a primary mechanism is involved, or whether the decline in protein
content and synthesis is the result of a combination of mechanisms. The slow
onset of significant symptoms (5th day of treatment) suggests that the pro¬
metryne does not directly affect protein synthesis, but rather the effect is upon
some cofactor(s) or intermediate(s) essential for protein synthesis. The data
presented in this paper suggest that RNA is not a limiting factor as a decrease
in protein synthesis would be expected to be preceded, or at least accompanied,
by a decrease in RNA content and synthesis. Protein synthesis requires high-
energy phosphate compounds, and ATP is essential for some steps. Thus block¬
age of oxidative phosphorylation as shown by McDaniel and Frans (1969)
is one possible site of action for prometryne, but does not appear to be the
main one since ATP is an essential nucleotide for RNA synthesis. A reduction
in ATP concentration that would inhibit protein synthesis would probably
inhibit RNA synthesis also.

Senescence in plant tissues is characterized in part by a decline in protein
synthesis and content (Goldthwaite and Laetsch, 1967; Osborne, 1962).
Goldthwaite and Laetsch (1967) found that the ability of white light to retard
senescence in bean leaf discs was blocked by concentrations of 3-(3,4-dichloro-
phenyl)- 1 , 1- dime thy lure a that inhibited photosynthesis. Because prometryne

198

THE TEXAS JOURNAL OF SCIENCE

is a known inhibitor of C02 fixation (Sikka and Davis, 1969), the most probable
explanation for its effect on protein metabolism in bean leaf discs is an ac¬
celerated rate of senescence due to blockage of the photosynthetic mechanism.
Senescence might also be expected to result in decreased RNA synthesis
(Goldthwaite and Laetsch, 1967); thus the data presented here further sug¬
gest that pathways for protein metabolism are affected more rapidly than
pathways for RNA metabolism in bean leaf discs.

ACKNOWLEDGMENT

This study was supported by funds for organized research from the State
of Texas.

LITERATURE CITED

Ashton, F.M., G. Zweig, and G.W. Mason, 1960-The effect of certain triazines on C1402
fixation in red kidney beans. Weeds, 8:450.

- , E.M. Gifford, Jr., and T. Bisalputra, 1963-Structural changes in Phaseolus

vulgaris induced by atrazine. II. Effects on the fine structure of chloroplasts. Botan.
Gaz., 125:336.

Barker, G.R., and J.A. Hollinshead, 1964-Nucleotide metabolism in germinating seed:
the ribonucleic acid of Pisum anvense. Biochem. J., 93:78.

Beruter, Jr., and A.T. Temperli, 1970-Influence of prometryne and ioxynil on photo¬
synthesis and nucleic acid metabolism in plants . Experientia, 26:600.

Goldthwaite, J.J., and W.M. Laetsch, 1967— Regulation of senescence in bean leaf discs
by light and chemical growth regulators. Plant Physiol., 42:1757.

Good, N.E., 1961 -Inhibitors of the Hill reaction .Plant Physiol., 36:788.

Gysin, H., and E. Knusli, 19 60 -Chemistry and herbicidal properties of triazine derivatives.
In R.L. Metcalf (Ed.), Advances in Pest Control Research, Vol. III. Interscience Pub¬
lishers, Inc., New York, pp. 289-358.

Hilton, J.L., L.L. Jansen, and H.M. Hull, 196 3 -Mechanisms of herbicide action. Annu.
Rev. Plant Physiol., 14:35 3.

Lowry, O.H., N.J. Rosenbrough, A.L. Farr, and R.J. Randall, 1951 -Protein measurement
with the folin-phenol reagent./. Biol. Chem., 193:265.

Mann, J.D., L.S. Jordan, and B.E. Day, 1965 -A survey of herbicides for their effect upon
protein synthesis. Plant Physiol., 40:840.

McDaniel, J.L., and R.E. Frans, 1969-Soybean mitochondrial response to prometryne
and fluometuron. Weed Sci., 17:192.

Moreland, D.E., 1967 -Mechanisms of action of herbicides. Annu. Rev. Plant Physiol.,
18:365.

- , W.A. Gentner, J.L. Hilton, and K.L. Hill, 1959-Studies on the mechanism

of herbicidal action of 2-chloro-4,6-bis(ethylamino)-s,-triazine./>/ffttf Physiol., 34:432.

EFFECTS OF PROMETRYNE ON RNA

199

- , and K.L. Hill, 1962 -Interference of herbicides with the Hill reaction of isolated

chloroplasts. Weeds, 10:229.

- , S.S. Malhotra, R.D. Gruenhagen, and E.H. Shokraii, 1969-Effects of herbicides

on RNA and protein synthesis. Weed Sci., 17:556.

Osborne, D..J., 1962-Effect of kinetin on protein and nucleic acid metabolism in Xan-
thiurn leaves during senescence. Plant Physiol., 37:595.

Rhodes, M.J. C., and E.W. Yemm, 1963 -Development of chloroplasts and the synthesis
of proteins in leaves. Nature, 200:1077.

Sikka, H.C., and D.E. Davis, 1969-Effect of prometryne on 14C02 fixation in cotton
and soybean. Weed Sci., 17:122.

Singh, R.P., and S.W. West, 1967 -Influence of simazine on chloroplast ribonucleic acid
and protein metabolism. Weeds, 15:31.

Smith, D., and K.P. Bucholtz, 1962-Transpiration rate reduction in plants with atrazine.

Science, 136:263.

Truelove, B., L.R. Jones, and D.E. Davis, 1973-Light and prometryne effects on leucine
uptake and incorporation. Weed Sci., 21:24.

Zweig, G,, and F.M. Ashton, 1962-The effect of 2-chloro-4-ethylamino-6-isopropyl-
amino-s-triazine (atrazine) on distribution of 14C-compounds following 14CC>2 fixa¬
tion in excised red kidney bean leaves./. Exp. Bot., 13:5.

RESULTS OF A MAGNETOMETER SURVEY AT HUECO TANKS,

A PREHISTORIC MOGOLLON VILLAGE IN WESTERN TEXAS

by J. BARTO ARNOLD III and GEORGE B. KEGLEY III

Texas Antiquities Committee and Interpretation and Exhibits Section ,
Texas Parks and Wildlife Departmen t, A us tin 78 744

ABSTRACT

Results of a magnetometer survey at Hueco Tanks State Park (41 EP 2) suggest a re¬
lationship between magnetic anomalies and pit house structures and other areas of cultural
activity at this site. Hueco Tanks is a late prehistoric site of the Jornada Branch of the
Mogollon near El Paso, Texas. Two techniques of magnetometer surveying were employed.
One consisted of a tightly controlled “insite” mode. The data from this technique can be
reduced to a map, which shows the variations in magnetic intensity by contour lines. The
second method employed a “search mode” technique of a more wide-ranging nature.
Anomalies discovered by the proton magnetometer were tested by excavation.

INTRODUCTION

This survey used the magnetometer to locate cultural features for excavation
and to help define the extent of the village area. The survey and subsequent
excavation results are discussed following background information on the
instrument and the site.

The magnetometer can be an important tool for archeology, both on land and
underwater. It is a remote sensing device which enables the archeologist to
make a preliminary assessment of the sub-surface cultural features of a site with¬
out excavation. Data from a careful magnetometer survey can be invaluable in
planning research strategy and in choosing excavation units. Underwater ar¬
cheologists of the Texas Antiquities Committee have successfully used the
magnetometer to produce magnetic contour maps of mid-16th century Spanish
shipwrecks off the Texas coast. These maps are used in association with shore-
based control system to direct the excavation in the murky water (Clausen and
Arnold, in preparation). Interest in the application of magnetometers to ar¬
cheology began with Aitken (1958) and a series of articles published in the
journal, Archaeometry. Black and Johnston (1962) report on a magnetometer
survey at the Angel site, a large Middle Mississippian village. Breiner and Mac-
Naughton (1965) explain how magnetometers work and discuss the application
of these instruments to land and marine archeology. Linington (1966) presents
a discussion of magnetic theory and archeological interpretation of magneto-

Accepted for publication: June 16, 1975.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

202

THE TEXAS JOURNAL OF SCIENCE

meter data. Pioneering work in the positioning control for magnetometer surveys
in underwater archeology was done by Clausen (1966) in Florida. The survey of
the Olmec site of San Lorenzo is a classic example of the use of the magneto¬
meter to guide excavations on land (Breiner and Coe, 1972). Perhaps the most
comprehensive articles on the archeological application of magnetometers are
by Tite (1972) and Breiner (1973).

The site chosen for a magnetometer survey was a small Jornada Mogollon
(Lehmer, 1948) village located thirty miles northeast of El Paso, Texas. Test
excavations in the fall of 1972 revealed 3 shallow pithouses (Kegley, in prepara¬
tion). These houses and additional ones located since the first field season were
identified in order of discovery, House 1, House 2, etc. Three house locations
are shown in Figure 1 , which shows only a portion of the total excavated area.

Like other similar sites in the area, the village is blanketed with midden soil,
seldom deeper than 15 cm. One radiocarbon date of A. D. 1 150 + 50 (TX 1735)
is in general agreement with cross dating of ceramics. Although pit houses could
be located by conventional methods such as trenching, there were no suface
indications to aid in locating the structures. This, then, led to the exploration of
remote sensing techniques.

The instrument utilized for this survey was a Varian Proton Magnetometer
(M-50) with the sensor mounted on a 1.38m aluminum pole. A magnetometer
senses distortions in the earth’s magnetic field caused by the presence of ferrous
compounds. Initially, it was not certain that the instrument would assist in de¬
lineating cultural features at this site. Local geology could be expected to pro¬
duce a masking effect since the site is located about 100m from a massive out¬
crop of syenite porphyry called the Hueco Tanks. This, however, did not prove
to be a problem. Test readings taken directly beside the outcrop recorded a
1500 gamma distortion, but successive stations moving away from the outcrop
indicate that at a distance of 50m there was no noticeable interference.

No metallic iron artifacts could be expected in the prehistoric village, but
the magnetometer is not limited to detecting anomalies caused by metallic
iron. Iron in the form of the compounds hematite, maghemite and magnetite
appears as a trace element in many clays. When the clay is fired as pottery, at
temperatures above the Curie point of 580 C for magnetite and 650 C for hematite
(Tite, 1972), the magnetic domains align themselves with the earth’s magnetic
field during cooling (each potsherd is like a small magnet). The remnant mag¬
netization thus produced will cause a greater anomaly than an equal amount
of randomly aligned magnetite in unfired clay. Therefore, concentrations of
pottery could cause anomalies although the strength of the distortion would
probably be small. This may in part be due to the randomized alignment of the
remnant magnetization of the induvidual sherds. During the first season’s ex¬
cavation, pottery was found in significantly higher quantities in the fill above
the floors of pit houses than about the structures. We believe that this is a

PREHISTORIC MOGOLLON VILLAGE

203

HOUSE 2

S 75

S 75

E 10

W 10

TEST

PIT

HOUSE 4

S 95

S 95

10-k

40 20

METERS

10 y CONTOUR INTERVAL

10 30

MAGNETIC CONTOUR MAP
HUECO TANKS
41 EP 2

HOUSE 6

Figure 1. Computer drawn magnetic contour map showing a portion of 41EP2 with cul¬
tural features superimposed. Area delineated by line is the area of “in-site mode.”

result of the abandoned pit houses having filled in with midden material. Burn¬
ing of the houses could cause similar alignment of any traces of the ferri-magnetic
compounds in the adobe used to line the pits. This could possibly cause a large
anomaly (Tite, 1972).

204

THE TEXAS JOURNAL OF SCIENCE

Another cultural feature which could possible be detected by the magneto¬
meter is the fired adobe hearth which is characteristic of these houses. One of
the houses found in the first season was still exposed and since it had a hearth,
the opportunity was provided for a test. The hearth was about 20 cm in diameter
and caused only a 10 gamma anomaly and that only with the sensor directly
above the hearth. At a distance of 1 m, the hearth did not register at all. This
type of small hearth is not likely to be any help in this particular site in view
of the amount of background noise caused by the overlying midden and local
geological conditions. Experience at this site indicated that random + 5 gamma
deviations are not unusual in an area of thick midden deposits (this noise factor
is well illustrated in the computer drawn perspective rendition of the magnetic
data in Figure 2).

METHOD AND RESULTS

The survey was divided into 2 phases consisting of an in-site mode and a
search mode. The term “in -site” is used to refer to a tightly controlled systematic
survey, with complete coverage of an area already established as part of a site
(Clausen and Arnold, in preparation). Through this method is produced a
magnetic contour map which can be used as a guide for the excavations. The
search mode is of a more speculative nature, with the emphasis on prospecting
for anomalies. The search mode is for initially locating an anomaly or a site
or determining site boundaries.

At Hueco Tanks, the area between and adjacent to the known pit houses
was selected for an in-site survey. Ten m. squares with corners corresponding
to the original grid for the site were gridded with string at 1 m intervals. The
small interval between the points of the data sample was necessary in view of
the expected small magnitude of the anomalies. Two things soon became ap¬
parent: 1st, there would be no trouble picking up and defining anomalies at the
site; and 2nd, setting up the control grid took much longer than acutually taking
the data. The in-site mode was used to cover approximately 450 m2 (Figure 1).
Subsequent preparation of a contour map of the local variations in the earth’s
magnetic field showed 5 well defined anomalies.

Archeological testing in the fall of 1973 was begun after the magnetometer
survey, by excavating test pits near the centers of magnetic anomalies or “peaks”
plotted as contours (Figure 1). Results obtained from this excavation are con¬
sidered significant. The following discussion is keyed to Figure 1. A test pit
about 6 m. north of House 6 exposed a concentrated area of mixed burned
rock and an ashy midden soil at a depth of 15 cm below the surface. House
6 has a contiguous room or rooms to the east which appear to have burned,
raising the possibility that fire has been instrumental in producing the magnetic
anomaly, which we tested, locating the north wall of the house. Burned structures
and fire -cracked rock are rare in other portions of the site. Proceeding next to
the area adjacent to House 2 (excavated in 1972), our excavations uncovered a

PREHISTORIC MOGOLLON VILLAGE

205

/ / i f I

METERS

Figure 2. Computer drawn perspective views of magnetic data.

206

THE TEXAS JOURNAL OF SCIENCE

complex series of prepared superimposed pit house floors. A promising 50 gamma
anomaly led us to test here. However, this pit house turned out to be quite
similar to other pit houses at the site. No evidence of burning was noted, but
a portion of the anomaly was no doubt caused by a 30 lb syenite porphyry
metate found in a shallow depression in the house floor.

House 4 was located during test excavation of a 10 gamma anomaly. The
situation is analogous to other pit houses at the site, the thin blanket-like cover
of midden material thickens to a depth of about 1 m and extends from the ground
surface to the floor of the house. In the pit fill there is a higher concentration
of midden soil, melted adobe, potsherds, and lithic artifacts than in the surround¬
ing midden.

A 20 gamma anomaly lies about 10 m south of House 4. Testing here produced
no evidence of structures or features. This anomaly, however, may prove instruct¬
ive since excavation again produced an unusually rich midden having a high
yield of ceramic and lithic material.

The final anomaly appears in the center of Figure 1 . It is spacially small and
of 20 gammas intensity. The cause of this anomaly was found upon test ex¬
cavation to be an iron reinforcement rod used the previous season to mark a
grid corner.

As time was short, a limited search mode survey of a less precise and com¬
plete but more extensive nature was carried out. It was intended to sample the
remainder of the site and perhaps define its limits. Results of this search mode
survey are more difficult to evaluate, but it did result in locating several more
anomalies and gave some feeling for the extent of the site. The noise gave way
to relative quiet as the midden thinned out.

For the search mode survey, the magnetometer operator stood in the desired
area. His position became the center of 2 circles of data samples, 8m and 4m in
diameter. The samples on each circle were again spaced at lm intervals. The
centers of these circles were plotted. If an anomaly was detected, the center of
the next sample was moved slightly to facilitate the delineation of its extent.
The magnitude was recorded and the center of the anomaly was marked with
a stake later to be shot in with alidade and plane table.

Prospecting in the search mode produced 6 anomalies varying in strength
from 10 gammas to 80 gammas. Since time was not available for testing all
6 anomalies, 3 showing the greatest gamma deviation were chosen for trial
excavations. Results are unclear since none of the anomalies checked produced
cultural features nor offer easy explanation for the magnetic anomalies. The
test pits lacked dense concentrations of artifacts or evidence of fire. Localized
geological conditions can not be ruled out as causes of these anomalies.

CONCLUSIONS

The in-site of survey has proven itself a most valuable guide for planning ex¬
cavation. The search mode survey aided in delineating the extent of the site but

PREHISTORIC MOGOLLON VILLAGE

207

limited test excavations revealed no cultural features. Additional magnetometer
work at Hueco Tanks would no doubt be fruitful. Further excavation would also
provide useful information concerning the magnetic data already in hand. For
example, one large area of magnetic disturbance in the 20 gammas range remains
untested (Figure 1).

Another conclusion drawn from the survey data deals with the strength of
various anomalies. In the area of the tightly controlled in-site survey, it seems
that the magnitude of magnetic anomalies is less significant than the form or
shape of these anomalies.

Breiner and Coe (1972) assert that no site is suited for magnetometer survey
until such suitability is demonstrated. This survey, nevertheless, was successful
in detecting anomalies representing cultural disturbances in a geological situation
which at first glance would contra-indicate a successful magnetometer survey.
However, we agree with them that someone with experience in magnetic survey¬
ing, preferably an archeologist, should be present to carry out the surveying
and interpretation of the resulting data ( ibid .).

LITERATURE CITED

Aitken, M. J., 1958 -Magnetic Prospecting I. Archaeometry , 1 :24.

Black, G. A. and R. B. Johnston, 1962-A test of magnetometry as an aid to archaeology.
American Antiquity , 28:199.

Breiner, S., 191 3 -Applications Manual for Portable Magnetometers. Geometries, Palo Alto,
California.

- , and M. D. Coe, 1972 -Magnetic exploration of the Olmec Civilization. American

Scientist, 60:566.

- , and K. G. MacNaughton, 1965 -The application of magnetometers to underwater

archeology. Sec. Con. on Underwater Archeology. Toronto, Canada.

Clausen, C. J., 1966- The proton magnetometer: Its use in plotting the distribution of the
ferrons components of a shipwreck site as an aid to archeological interpretation. Fla.
Anthropologist, 19:77.

- , and J. B. Arnold III, The magnetometer and underwater archeology: A new con¬
trol technique. In prep.

Kegley, G. B ., Hueco Tanks Village. Tex. Parks and Wildlife Dep. Rep., In. prep.

Lehmer,D. J., 1948-The Jornada Branch of the Mogollon. Univer. of Arizona Bull. XIX.
(Social Science No. 17).

Linington, R. E., 1966-An extension to the use of simplified anomalies in magnetic survey¬
ing. Archaeome try, 9:51.

Tite, M. S., 1912-Methods of Physical Examination in Archaeology . Sem. Press, New York.

■

A SODIUM IODIDE CENTRIFUGATION TECHNIQUE FOR ISO¬
LATION OF NEWLY-REPLICATED DNA FROM RAT LIVER

by ROGER F. BROWN

Department of Biology, Southwest Texas
State University, San Marcos 78666

ABSTRACT

A simple procedure has been described for isolation of newly-replicated DNA from rat
liver. Liver homogenates were lysed in 2% sodium dodecyl sulfate-0.1 M NaOH and centri¬
fuged for 2.5 hr on alkaline sodium iodide step gradients. The procedure involved minimal
handling of DNA and provided high resolution of newly-replicated DNA from bulk DNA.

INTRODUCTION

Replication of mammalian DNA is a discontinuous process. Very short
chains of approximately 100 nucleotides (thought to be analogous with the
Okazaki fragments of bacteria) are formed and subsequently linked with longer,
intermediate strands of DNA (Huberman and Horowitz, 1973). The intermediate
strands join in turn to form bulk DNA (Hyodo, et al., 1970).

Various analytical techniques, including autoradiography (Huberman and
Riggs, 1968), sedimentation in sucrose gradients (Berger and Irvin, 1970; Huber¬
man and Horowitz, 1973; Tsukada, et al., 1968), and chromatography (Sato,
et al, 1972), have been utilized to characterize growing strands of DNA label¬
led with radioactive precursors. Techniques chosen for isolation and analysis
of newly replicated DNA fragments should satisfy a number of criteria includ¬
ing minimal handling to avoid excessive shearing, adequate separation from bulk
DNA, and sufficient yield for analysis. A rapid, simple procedure conforming
to these criteria is described in the present report for isolation of newly repli¬
cated DNA from liver tissue. The procedure involves alkaline treatment of liver
homogenates followed by centrifugation in alkaline sodium iodide step gradients.

MATERIALS AND METHODS
Treatment of Animals

Male albino rats from a colony maintained in this laboratory were used when
they weighed 80-100 g. Synthesis of liver DNA was induced by removal of the

Accepted for publication: November 10, 1975.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

210

THE TEXAS JOURNAL OF SCIENCE

median and left lateral lobes of the liver (Higgins and Anderson, 1931). [Methyl-
3H] Thymidine (17 Ci/mmole, purchased from Amersham-Searle) was admin¬
istered via the tail vein at 19-22 hr after the partial hepatectomy.

Preparation of Tissue Lysates and Nal Step Gradients

Animals were killed by cervical dislocation. Approximately 1 g of liver was
removed, homogenized in 4 ml of ice-cold 0.15 M NaCl with 4 strokes in a loose
Dounce homogenizer, and filtered through 4 layers of cheesecloth. Filtered
homogenate (0.5 ml) was pipetted into a tube containing an equal volume of
2% sodium dodecyl sulfate-0.1 M NaOH. Less than 1 min was required from
the time the animal was killed to addition of homogenate to the alkaline sol¬
ution. The tube was gently rolled to mix the contents and then left undisturbed
for 60 min at room temperature.

Step gradients were formed in cellulose nitrate tubes fitting the Spinco
SW 41 rotor by the successive addition of 2.8 ml layers of NaI-0.1 M NaOH
solutions with densities of 1 .5 , 1 .4, 1 .3, and 1 .2 g/cc, respectively. Lysed homo¬
genates were then poured on top and the gradients centrifuged for 2.5 or 24
hr at 35,500 RPM and 18 C. Twelve 1 ml fractions were collected from the
bottom of each gradient. The refractive index of each fraction was measured
at room temperature with an Abbe refractometer and compared with alkaline
Nal solutions of known densities.

Determinations of Size of DNA Recovered from Nal Gradients

Nal gradients were fractionated after centrifugation for 2.5 hr and bottom
fractions containing bulk DNA were pooled. Fractions containing newly-repli¬
cated DNA collected from the middle of the gradients were pooled separately.
The pooled fractions were dialyzed against 0.1 M NaOH-1 mM EDTA and then
concentrated by dialysis against 2 M sucrose containing 0.1 M NaOH-1 mM
EDTA. Aliquots (0.2 ml) of the concentrated solutions of newly- replicated
and bulk DNA were layered on 5 ml gradients of 10-25% sucrose containing
0.9 M NaCl, 0.1 M NaOH, and 1 mM EDTA. Centrifugation was for T .5 hr in
a Spinco SW 50.1 rotor at 40,000 RPM and 18 C. 1 4 C- Labelled T7 DNA was
used as a marker for estimation of the sedimentation coefficient of the newly-
replicated DNA.

A ssay Pro ced ures

Half of each Nal gradient fraction was removed to 5 ml of ice-cold 10%
trichloracetic acid (TCA). Acid-insoluble radioactivity was precipitated onto
a 2.5 cm disc of Whatman No. 1 filter paper covered with 40 mg of ‘Celite’
(Johns-Manville), dried with ethanol and ether, and counted in a liquid scintil¬
lation solution containing 4 g of PPO and 0.05 g of POPOP/1 of toluene. DNA
was precipitated from the remaining half with cold TCA and measured color-
imetrically by reaction with diphenylamine (Burton, 1955).

ISOLATION OF NEWLY-REPLICATED DNA

211

Ten drop fractions were collected from the bottom of the alkaline sucrose
gradients and adjusted to 1 ml with water. Absorbance was measured at 260
mjU and radioactivity was counted in a solution of toluene and Triton X-100.

RESULTS

Centrifugation of Purified Nuclear DNA and Crude Homogenate

Isolation of newly replicated DNA by Nal centrifugation of purified DNA
was compared with results obtained when lysed whole homogenate was ap¬
plied to the Nal gradients. Liver was removed from one rat 2 min after injection
of 100 juCi of 3 H- thymidine and nuclei were isolated as described previously
(Brown, et al, 1970). DNA, purified from the nuclei by the procedure of
Zamenhof (1957), was redissolved in 0.15 M NaCl. Aliquots (0.5 ml) of the
purified DNA were mixed with equal volumes of 2% sodium dodecyl sulfate-
0.1 M NaOH and layered on two Nal gradients. Liver was removed from a
second rat 2 min after injection of 100 jxC i of 3 H- thymidine, homogenized,
lysed, and layered on two additional Nal gradients. One of the gradients with
lysed homogenate and one with purified DNA were centrifuged for 2.5 hr. The
other two gradients were centrifuged for 24 hr.

Results presented in Figure 1 suggest that physical manipulation of samples
during preparation affected resolution of newly- replicated DNA from bulk
DNA. The bulk of the DNA from lysed homogenate (samples subjected to
minimal handling) sedimented close to the bottom of the gradients during
the 2.5 hr centrifugation and most of the acid-insoluble radioactivity remained
near the top (Figure 1 A). Preparation of purified DNA required more extensive
handling and the bulk of the purified nuclear DNA sedimented less than half¬
way through the gradient during the 2.5 hr centrifugation allowing only partial
resolution of radioactive DNA (Figure IB). Centrifugation of samples for 24 hr
sedimented radioactivity and bulk DNA to the same equilibrium density (Fig¬
ures 1C and ID). This suggest that the radioactive peak at the top of the gradient
shown in Figure 1 A is newly -replicated DNA.

Distribution of Protein in Gradient Fractions

Results presented in Table 1 indicate that Nal centrifugation effectively
separated cellular proteins from DNA. A liver sample was lysed and centri¬
fuged for 2.5 hr in a Nal gradient. Protein was precipitated from each gradient

TABLE 1

Distribution (%) of Protein in Gradient Fractions

Fraction Number

1

2

3

4

5

6

7 8

9

10

11

12

0.1

0.4

0.3

0.2

0.2

0.2

0.2 0.4

0.4

1.6

30.4

65.6

FRACTION NUMBER

212

THE TEXAS JOURNAL OF SCIENCE

ABSORBANCE

o o o

o -* W w

AT 600 my

o o o

O '«■* N3 CO

DENSITY,

b

g/«

Figure 1. Comparison of Nal gradient sedimentation of lysed liver homogenate and
purified liver DNA from rats pulse labelled with 3H-thymidine for 2 min.
A. Lysed homogenate centrifuged for 2.5 hr, B. Purified DNA centrifuged
for 2.5 hr. C. Lysed homogenate centrifuged for 24 hr. D. Purified DNA centri¬
fuged for 24 hr. Radioactivity (o); absorbance (•); density (x).

ISOLATION OF NEWLY-REPLICATED DNA

213

fraction with cold 10% TCA and measured colorimetrically by the procedure
of Lowry, et al., (1951). As shown in the table, 96% of the protein from lysed
homogenate was recovered at the top of the gradient (fractions 1 1 and 12).
Very little protein was recovered with bulk DNA near the bottom (fractions 1-3)
or in fractions containing most of the newly -replicated DNA (fractions 4-10).

Effect of Pulse Duration on Sedimentation Profiles

Sedimentation profiles shown in Figure 2 indicate that as the pulse duration
increased, acid insoluble radioactivity shifted to bulk DNA. Liver samples were
removed from 4 rats labelled with 10-100 juGi of 3 H- thymidine for durations
of 2 to 60 min, lysed, and centrifuged for 2.5 hr in Nal gradients. Newly-repli¬
cated DNA from the animal pulse -labelled for 2 min with 100 /rCi was recovered
in the top fractions of the gradient (Figure 2A). Pulse labelling for 5 min with
50 /iCi resulted in a broad peak of radioactivity in the middle of the gradient
(Figure 2B). Removal of liver 10 min after injection of label (30 juCi) resulted
in a higher peak of radioactivity in bulk DNA than in the middle fractions
(Figure 2C). And, finally, most of the acid-insoluble radioactivity was in bulk
DNA in the sample from the rat pulse -labelled for 60 min with 10 /rCi. Similar
profiles were obtained with preparations centrifuged for 2.5 hr at 41,500 RPM
in a Spinco SW 50.1 rotor.

Alkaline Sucrose Gradient Analysis of Products Recovered from a Nal Gradient
Bulk and newly-replicated DNA prepared by centrifugation of lysed liver
homogenate in a Nal gradient were analyzed by sedimentation in alkaline
sucrose gradients. As shown in Figure 3, radioactive, newly -replicated DNA re¬
covered from the middle of the Nal gradient was smaller than bulk DNA re¬
covered from the bottom of the same gradient. To obtain these results liver
was removed from one rat 5 min after injection of 50 juCi of 3 H- thymidine,
lysed, and centrifuged for 2.5 hr in Nal. The Nal gradient was fractionated
and fractions 1 and 2, containing bulk DNA, were combined. Fractions 6-8,
containing newly -replicated, radioactive DNA, were pooled separately. Samples
of bulk and newly -replicated DNA and T7 DNA were then sedimented in separ¬
ate tubes containing alkaline sucrose gradients. Bulk DNA sedimented close
to the bottom of the first sucrose gradient. Newly-replicated DNA, however,
sedimented less than halfway through the second sucrose gradient indicating
that it was much smaller than the bulk DNA. Comparison of the position of
the peak of radioactivity (fraction 14) with the position of the T7 DNA marker
(identified by the arrow) sedimented in an identical, third gradient yielded
an estimated sedimentation coefficient (S) of 49 for the newly -replicated DNA.
This estimate was based upon a published S value of 37.2 for T7 DNA in alkaline
solution (Studier, 1965).

FRACTION NUMBER

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THE TEXAS JOURNAL OF SCIENCE

ABSORBANCE

AT 600 m fj

Figure 2. Nal gradient sedimentation of lysed liver homogenates from animals pulse-
labelled with 3H-thymidine for: (A) 2 min; (B) 5 min; (C) 10 min; or
(D) 60 min. Sedimentation was from right to left. Radioactivity (o); absorb¬
ance (•).

ISOLATION OF NEWLY- REPLICATED DNA

215

RADIOACTIVITY, CRM PER FRACTION

Figure 3. Alkaline sucrose gradient analysis of DNA’s recovered from a Nal gradient.

Sedimentation profiles of newly-replicated DNA and bulk DNA centrifuged
in separate alkaline sucrose gradients (10-25%) are superimposed. The arrow
represents the position of 14C- labelled T7 DNA centrifuged in an identical
alkaline sucrose gradient. Sedimentation was from right to left. Radioactivity
(o); absorbance (•).

216

THE TEXAS JOURNAL OF SCIENCE

DISCUSSION

The alkaline treatment of liver tissue and centrifugation of crude lysates
on sodium iodide step gradients described in this paper provided good resolution
of newly- replicated DNA from bulk DNA. Two factors appear to have contri¬
buted to separation of the DNA’s. First, it was important to minimize handling
of samples to reduce breakage of DNA. Shearing stresses were minimized by
gentle lysis of liver tissue in alkaline solution. A similar alkaline lysis treatment
was used by Lett et al, (1967) for isolation of high molecular weight DNA from
cultured cells. Bulk DNA from liver lysates sedimented to the bottom of the
Nal gradients. When the bulk DNA was then recovered it was still highly viscous
and sedimented more rapidly through alkaline sucrose gradients than did the
newly-replicated DNA. Bulk DNA from the purified nuclear DNA sample,
however, sedimented less than halfway through the Nal gradient. Separation
from newly -replicated DNA was thus incomplete. This suggests that bulk DNA
may have been sheared during purification to fragments not much greater in
length than newly -replicated DNA.

Secondly, Nal step gradients were quite effective in separating newly-repli¬
cated DNA from bulk DNA if centrifugation was for 2.5 hr. Long durations
of centrifugation sedimented DNA (both new DNA and bulk DNA) to its
equilibrium density. Separation of the DNA’s seemed to be a function of poly¬
nucleotide chain length as evidenced by the results of the sucrose gradient
analysis of DNA recovered from a Nal gradient.

The Nal step gradients were developed as an alternative to conventional
alkaline sucrose gradients. This investigator was not successful with centri¬
fugation of crude lysates in sucrose gradients. The relatively dense lysates
tended to partially sink when poured onto sucrose gradients, but layered very
well on Nal as demonstrated by the fact that protein did not sediment appreci¬
ably into the Nal gradients.

The shift of acid-insoluble radioactivity from the top of the Nal gradient
to the bottom with increased durations of 3 H- thymidine incorporation appears
to reflect growth of new strands of DNA and is consistent with the results
obtained by Huberman and Horowitz (1973). These investigators observed
progressive lengthening of radioactive DNA chains from short to intermediate
and finally to bulk size as the length of the labelling period increased. The
average size estimated (49S) for fragments of newly -replicated liver DNA re¬
covered from the Nal gradients is within the size range observed from growing,
intermediate strands of DNA in cultured cells (Huberman and Horowitz, 1973;
Hyodo ,etal, 1970).

ACKNOWLEDGMENTS

14C-Labelled T7 DNA was a generous gift of Dr. Roger Hewitt, M.D. Ander¬
son Hospital and Tumor Institute, Houston, Texas.

ISOLATION OF NEWLY- REPLICATED DNA

217

This investigation was supported in part by state funds appropriated to
Southwest Texas State University for organized research.

LITERATURE CITED

Berger, H. and J.L. Irvin, 1910-Proc. Nat. Acad. Sci., USA, 65:152.

Brown, R.F., T. Umeda, S. Takai and I. Lieberman, 1910-Biochem. Biophys. Acta, 209:49.
Burton, K., 1955 -Biochem. J 61:473.

Higgins, G.M. and R.M. Anderson, \93\-Arch. Pathol., 12:186.

Huberman, J.A. and H. Horowitz, 1913-Cold Spring Harbor Sym. Quant. Biol., 38:233.
- and A.D. Riggs, 1968-/. Mol. Biol, 32:327.

Hyodo, M., H. Koyama and T. Ono, 1910-Biochem. Biophys. Res. Commun., 38:513.

Lett, J.T., I. Caldwell, C.J. Dean and P. Alexander, 1961 -Nature, 214:790.

Lowry, O.H., NJ. Rosebrough, A.L. Farr and R.J. Randall, 1951-/. Biol. Chem., 193:265.

Sato, S., S. Ariake, M. Saito and T. Sugimura, \91l-Biochem. Biophys. Res. Commun.,
49:270.

Studier, F.W., 1965-/. Mol. Biol., 11:373.

Tsukada, K., T. Moriyama, W.E. Lynch and I. Lieberman, \9 6% -Nature, 220: 162.

Zamenhof, S., 1951 -In S.P. Colowick and N.O. Kaplan (Eds.) Methods in Enzymology
Vol. 3, Academic Press, p. 696.

BARRIERS TO INTERNAL ROTATION IN 2-BROMOPROPENE
AND 2-IODOPROPENE

by G. A. CROWDER and ROYCE W. WALTRIP

Department of Chemistry, West Texas State University,

Canyon 79016

ABSTRACT

The 3-fold and 6-fold barriers to internal rotation of the methyl group were calculated
from torsional frequencies to be 2640 and -106 cal/ mole for 2-bromopropene and 2580 and
-75 cal/ mole for 2-iodopropene.

INTRODUCTION

Vibrational spectra of 2-bromopropene have been published by Meyer and
Gunthard (1967), and Meyer, et al., (1969) have published the infrared spectrum
of 2-iodopropene. The potential energy barrier hindering internal rotation of the
methyl group in 2-bromopropene has been determined by microwave spectroscopy
by Benz, et al, (1966). The far infrared vapor and liquid-state spectra of 2-bromo-
propene had previously been determined by one of the present authors. Obser¬
vation of the 0 -* 1 and 1 ->2 torsional frequencies for both 2-bromopropene and
2-iodopropene has allowed calculation of not only the 3-fold barrier to internal
rotation, but also the 6-fold barrier.

EXPERIMENTAL

Far infrared spectra were obtained for 2-bromopropene with a Perkin-Elmer
model 301 spectrophotometer, located at the Bartlesville Energy Research Center,
through the courtesy of Dr. D. W. Scott. The sample of 2-bromopropene was ob¬
tained from Pfaltz and Bauer, and was used without further purification. No im¬
purities were detected by gas chromatography.

Barriers to Internal Rotation

The observation of both 0 -> 1 and 1^2 torsional bands allows the determin¬
ation of both V3 and V6 in the potential energy expression

V(cl)) = 1/2V3(1 - cos 3<|>) + /4V6(1 - cos 6(f)) + -- -
by the procedure outlined by Fateley and Miller (1963).

Accepted for publication: December 1, 1976.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

220

THE TEXAS JOURNAL OF SCIENCE

The F value for 2-bromopropene was taken from Benz ,et al, (1966). The 3-
fold barrier height is given by

V3=|Fs

and s is obtained from tables of solutions for the Mathieu equation. The value of
V3 for 2-bromopropene is seen from Table 1 to be 923 cm-1 or 2640 cal/mole,
and V6/V3 = -0.04.

TABLE 1

Methyl Torsional Barrier Determination Summary.

Compound

it i

v V

u)a

(cm-1)

F

(cm-1)

S

0.00

V3(cm

-0.01

[) when V6/V3 is:

-0.02 -0.03

-0.04

CH3CBr=CH2

o->i

196.0

5.60

69.12

871

883

896

910

923

1 ~>2

183.6

5.60

70.89

893

901

908

915

923

CH3CI=CH2

o->i

195.4

5.60

68.73

866

878

891

904

918

1 ->2

182.2

5.60

69.99

882

889

896

903

910

aObserved frequencies are from this work for 2-bromopropene and from Meyer, et al, (1969)
for 2-iodopropene.

In order to justify the negative sign of V6 , it must be shown that the accuracy
of the torsional frequencies is high enough that a negative V6 is obtained for the
complete range of uncertainty in the frequency measurement. Meyer and Gunthard
(1967) determined the wavenumber of the 0 ^ 1 transition to be 195.7 cm"1 , with
an uncertainty of 0.5 cm"1 , and they report the wavenumber of the 1 -> 2 transi¬
tion as 183 cm"1, with an uncertainty of lcm"1. Our values of 196.0 and 183.6 cm"1
should be accurate to within these same uncertainties. Therefore, the limits of
these 2 transitions should be 182.6 to 184.0 and 195.5 to 196.2 cm"1 .

The combination 196.2 and 182.6 cm"1 (the high and low extremes, respectively,
of the 2 frequencies) gives V3 = 902 cm"1 , with V6/V3 = -0.022. while the other
combination of extremes, 195.5 and 184.0 cm"1 , gives V3 = 940 cm'1 and V6/V3
= -0.056. Therefore, the maximum uncertainties are approximately 20 cm"1 for
V3 and 0.017 for the V6/V3 ratio, based only on the accuracy of the torsional fre¬
quencies. The uncertainty in the value of F adds to the uncertainties in V3 and
V6, but Hunziker and Gunthard (1965) point out that this is less than the un¬
certainty due to the frequency error. The value of V3 should be accurate to within
100 cal, and our value, therefore, overlaps the microwave -determined value.

Use of the torsional wavenumbers reported by Meyer and Gunthard (1967),
namely 195.7 and 183 cm"1 , yields V3 = 2610 cal/mole and V6/V3 = -0.045.

The potential barrier hindering internal rotation of the methyl group in 2-iodo¬
propene has not been determined by microwave spectroscopy, and structural data
have not been obtained. Therefore, the F value for the internal rotation was taken

BARRIERS TO INTERNAL ROTATION

221

to be 5.60 cm-1, which is the same as for 2-bromopropene. The values for 2-chloro-
propene,5.62 cm-1 (Unland, et al, 1965), and 2-fluoropropene, 5.62 cm-1 (Fateley
and Miller, 1963), are almost the same.

Meyer, et al, (1969) have determined the vapor-state infrared spectrum and
they observed the 0 ->1 transition of the methyl torsion at 195.4 ± 0.5 cm-1 and
the 1 2 transition at 183.2 ± 0.5 cm-1 . The Fateley-Miller procedure was again

used to obtain V3 = 902 cm”1, or 2580 cal/mole (see Table l),andV6/V3 = -0.029.
Use of the 0.5 cm”1 uncertainties yields uncertainties of approximately 40 cal/mole
in V3 and 0.012 in the V6/V3 ratio. The uncertainty in the F value will increase
the overall uncertainty in V3 to approximately 70 cal/mole. The data used for
both compounds is summarized in Table 1 .

The methyl torsional barriers in the four 2-halopropenes are listed in Table 2.
The value of V6 for 2-bromopropene is 4% of V3 , and this is larger than any of
the V6 values for the compounds studied by Fateley and Miller (1963).

TABLE 2

Methyl Torsional Barriers in 2-Halopropenes.

Compound

V3

(cal/mole)

v6

(cal/mole)

Methoda

Reference

ch3cf=ch2

2440

MW

Peirce and O’Reilly (1959)

2340

+4

IR

Fateley and Miller (1963)

ch3cci=ch2

2671

MW

Unland, et al., (1965)

2645

-49

IR

Hunziker and Gunthard (1965)

CH3CBr=CH2

2695

MW

Benz, et al., (1966)

2640

-106

IR

This work

CH3CI=CH2

2580

-75

IR

This work

aMW=microwave, IR=infrared

ACKNOWLEDGEMENT

The authors are grateful to The Robert A. Welch Foundation for financial
support of this work.

LITERATURE CITED

Benz, H. P., A. Bauder, and Hs. H. Gunthard, 1966-/. Mol. Spectrosc., 21:165.

Fateley, W. G., and F. A. Miller, 1963 -Spectrochim. Acta., 19:611.

Hunziker, H., and Hs. H. Gunthard, 1965 -Spectrochim. Acta., 21:51.

Meyer, R., and Hs. H. Gunthard, 1967 -Spectrochim. Acta., 23A:2341.

- , Hunziker, H., and Hs. H. Gunthard, 1969 -Spectrochim. Acta., 25A:29 5.

Pierce, L., and J. M. O’Reilly, 1959-/. Mol. Spectrosc., 3:5 36.

Unland, M. L., V. Weiss, and W. H. Flygare, 1965-/. Chem. Phys., 42:2138.

THE POLAROGRAPHIC REDUCTION OF METAL IONS IN ACE¬
TONE AND ACETOPHENONE IN THE PRESENCE OF LITHIUM
PERCHLORATE AND TETRAETHYL AMMONIUM PERCHLORATE

by IVORY V. NELSON, Ph.D.

Department of Chemistry,

Prairie View A&M University,

Prairie View 77445

ABSTRACT

The polarographic reduction behavior of certain metal ions in acetone and acetophenone
in the presence of selected supporting electrolytes, lithium perchlorate and tetraethylammonium
perchlorate, is observed.

INTRODUCTION

The effect of supporting electrolytes on the polarographic reduction of metal
ions has been examined by several investigators. Larson and Iwamoto (1960) re¬
ported that half-wave potential for the reduction of Mn+2 in acetonitrile is more
positive with 0.1 M tetraethylammonium perchlorate than with 0.1 M Lf1, Na+1
or Mg+2 perchlorate as supporting electrolyte. They also noted similar behavior
with tetraethylammonium perchlorate supporting electrolyte as compared with
Lf 1 and Mg+2 perchlorates as supporting electrolytes for the polarographic re¬
duction of Mn+2 in benzonitrile. In addition, Larson and Iwamoto (1960) also
observed that with the exception of aluminum perchlorate, which is shifted in
the positive direction, the perchlorates of Ba+2 , Cr+3 , Fe+2 , Co+2 , Nf2 , Zn+2 and
Ccf2 are reduced at more negative half-wave potentials in benzonitrile with 0AM
LiC104 than with 0. 1 M tetraethylammonium perchlorate as supporting electrolyte.
Schaap, et al, (1961) studied the polarographic reduction of Tf1, Pb+2, and Ccf2
with 0.1 M NaN03 in ethylenediamine and concluded that the shifts in the half¬
wave potentials were attributed to ion-pair formation between the reducible metal
ion and anion of the inert electrolyte. Similarly, Schober andGutman (1958, 1959,
1960, 1961) examined the polarographic behavior of a number of metal salts in
ethylenediamine, and stated that the half-wave potentials of metal ions depended
upon the supporting electrolyte and the anion of the reducible metal ion.

In an effort to obtain additional information on the effect of inert electrolytes
on the polarographic behavior of metal ions, we have chosen to carry out 2 studies,
the polarographic reduction of metal ions in acetone and acetophenone in the
Accepted for publication: October 25, 1976.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

224

THE TEXAS JOURNAL OF SCIENCE

presence of lithium and tetraethylammonium perchlorates, and the effect of
inert electrolytes on the formation constants of aquo-Cu+2 complexes in acetone.

Acetone, dielectric constant 21, and acetophenone, dielectric constant 17, have
been neglected as solvents for electro -chemical studies.

Only 4 polarographic studies have been reported for acetone as a solvent. Arthur
and Lyons (1952) observed well-defined reduction waves for acid bromides and acid
chlorides in acetone, but an incompletely developed reduction wave for Ccf 2 in acetone
with LC1 as supporting electrolyte. Ozerov and Yakovleva (195 6) obtained polarograms
for the reduction of Sr+2 and Bf3 nitrates, Cu+2 and Zn+2 chlorides, Cdl2 and
CuBr in acetone solutions without supporting electrolyte and in acetone solutions
containing 0.5 M LiN03 as supporting electrolyte. Migal, et al.f (1956) studied
the polarography of HC1 in acetone with Li Cl as supporting electrolyte. They ob¬
served that the diffusion current constant of HC1 increased with increasing con¬
centration of HC1. Coetzee and Siao (1963) investigated the polarographic be¬
havior of the rare earth elements in acetone with 0.1 M tetraethylammonium
perchlorate supporting electrolyte. These investigators concluded that the re¬
duction of the +3 oxidation state of the rare earths led to H2 evolution rather
than amalgam formation, and that acetone is a differentiating solvent for the
polarographic reduction of HC104 and H2 S04 .

EXPERIMENTAL

The organic solvents were purified by treatment with appropriate reagents
and fractional distillation (Weissburger and Proskauer, 1955; Vogel, 1956). Reagent-
grade acetone was treated with KMn04, dried over anhydrous K2C03 and dis¬
tilled from P2 05. Highest available purity acetophenone was dried over anhydrous
CaCl2 and fractionally distilled.

Tetraethylammonium perchlorate was prepared from Eastman tetraethylammonium
bromide as previously described (Kolthoff and Coetzee, 1957). All the metal per¬
chlorates were obtained from G. F. Smith Chemical Company and were dried in
a vacuum over 60 C for 48 hrs. The residual water content of the perchlorate
salts was not determined; however, the colors of the dried meterials in comparison
to the corresponding hydrated salts indicated that the drying operation reduced
the amount of water in the salts. The total water content of the acetone solutions
containing supporting electrolyte and Cu(C104)2 was determined with a modified
Karl Fischer reagent (Mitchell and Smith, 1958). The water content of the aceto¬
phenone solutions was approximately 0.02 M.

Current-voltage curves were obtained with a Kelly, et al., (1959) and Larson
and Iwamoto (1962) controlled potential polarograph. All solutions were examined
at 25 C in a thermostatically controlled bath. An H-type cell, with the 2 compart¬
ments separated by a single glass disc, was used. The dropping mercury electrode
was placed in one side arm, and a saturated colomel electrode along with a platinum-
foil electrode to function as the auxilliary electrode in a 3-electrode system was
placed in the other.

POLAROGRAPHIC REDUCTION OF METAL IONS

225

RESULTS AND DISCUSSION

Effect of Supporting Electrolyte on Half-Wave Potentials

Table 1 lists the half-wave potentials for the reduction of Na+1 , C<f 2 , Mn+2 ,
Zn+2 , Cr+3 , Cr+2 , Nf 2 , Pb+2 , Cu+2 , Cu+1 and Mg+2 ions in acetone and acetophenone
in the presence of 0.1 M IiC104. The behavior observed in acetone for Na+1 , Mg+2 ,
Ccf2 , Cu+2 and Cu+1 in the presence of both supporting electrolytes is similar to
the behavior observed by Coetzee and Siao (1963).

TABLE 1

Half-Wave Potentials of Metal Ions in Acetone and Acetophenone.

Metalb

Acetone

El/2a

Sloped

F a
^1/2

Sloped

(Et4NC104)

Na+1

—

—

-1.90

0.062

Cd+2

-0.20

0.033

-0.21

0.036

Mn+2

-1.11

0.035

-1.12

0.039

Zn+2

-0.57

0.037

-0.61

0.031

Ni"2

-0.54

0.100

-0.57

0.105

Pb+2

-0.12

0.034

-0.12

0.030

Cu+2/Cu+1

+0.58

0.068

+0.55

0.058

Cu+1/Cu°

+0.33

0.065

+0.38

0.650

Mg+2

—

—

-1.73

0.080

Sr+2

—

—

-1.93

0.060

Cr+3/Cr+2

—

—

—

± —

Cr+2/Cr°

—

—

—

—

Metalb

Acetophenone

El/2a

Sloped

EV2a

Sloped

(LiC104)

(Et4NC104)

Na+1

_

_

_

_

Cd+2

-0.17

0.028

-0.16

0.028

Mn+2

-1.05

0.049

-1.05

0.047

Zn+2

-0.55

0.037

-0.5 3

0.040

Nr2

-0.51

0.081

-0.47

0.105

Pb+2

-0.08

0.028

-0.03

0.030

Cu+2/Cu+1

—

—

—

—

Cu ‘/Cu°

—

_

_

_

Mg+2

_

_

_

_

S/2

_

_

_

_

Cr+3/Cr+2

-0.50

0.059

-0.46

0.058

Cr+2/Cr°

-0.93

0.081

-0.97

c

a Versus S.C.E.

b0.1 mmol concentration

c Drawn out wave.
d0.05 9/n

226

THE TEXAS JOURNAL OF SCIENCE

It is also seen from the results presented in Table 1 that the waves of Mn+2, Zn+2,
Sr+2, and Pb+2 are reversible. Nf2 gives one well-defined diffusion plateau but is
somewhat drawn out.

The results obtained with solutions of the perchlorates of Ccf2, Mn+2, Zn+2,
Nf2, Pb+2, Cr+3, and Cr+2 in acetophenone in the presence of both supporting elec¬
trolytes is presented in Table 1. The observed half-wave potentials are in the same
order as those in acetone. C(f 2, Zn+2, and Pb+2 give well-defined waves corresponding
to a 2-electron reduction. Mn+2 and Nf2 give well-defined diffusion plateaus, but
are somewhat drawn out.

Cr+3 gives a well-defined first wave corresponding to a 1-electron reduction,
and a drawn out wave corresponding to the reduction of Cr+2 to the metal. Aceto¬
phenone is reduced at a potential of -1.4V vs S.C.E. The investigation of the
polarographic behavior of any metal ion which is reduced at potentials more
negative than 1.4V vs S.C.E. , therefore, is not possible.

In the presence ofO.l Af LiC104 the average E1/2 (E1/2 acetone-E1/2 acetophenone),
for the reduction of the metal ions is approximately -0.03V, whereas, for 0.1 M
tetraethylammonium perchlorate, the average Ey2 is approximately -0.8V. This
suggests that the solvated metal ion undergoing reduction is acetone and aceto¬
phenone is more nearly the same species when 0.1 M LiC104 is used as the supporting
electrolyte.

Effect of Supporting Electrolyte on Formation Constants

Figure 1 contains the experimental data, in the form of the half-wave potential
for the reduction of Cu+2 to Cu+1 vs the negative logarithm of the total water
concentration, for the polarographic study of the interaction of small amounts
of water in acetone with Cu+2 in 0.1 M LiC104 and 0.1 M tetraethylammonium
perchlorate supporting electrolyte solutions. Curve A shows the experimentally
observed dependency of the half wave potential for the Cu+2/Cu+1 couple on the
total water concentration with LiC104 as supporting electrolyte, and Curve B
shows the experimentally observed dependency of the half-wave potential for
the Cu+2/Cu+1 couple on the total water concentration with tetraethylammonium
perchlorate as supporting electrolyte.

Over the range of water concentrations studied, there was no change in the
liquid-junction potential with either supporting electrolyte. [E^2 Et4NC104 vs S.C.E.
in 0.01 M to 0.5 M water +0.942V, and E^2 Licl04 w S.C.E. in 0.01 M to 0.5 M
water = +0.940V for the oxidation of tris-4,7-dimethyl-l ,1 0-phenanthroline
Fe(C104)2 ] . There appears to be only a slight change in the solvation energy of
Cu+1 ion over the range of water concentration studied, as evidenced by a change
of only 5mV in the half-wave potential of the Cu+1/Cu(Hg) couple. Consequently,
the change in half-wave potential for the Cu+2/Cu+1 wave with water concentra¬
tion is due essentially entirely to the complexation of Cu+2 ions by water mole¬
cules, and due not to the interaction of water with Cu+1, nor to changes in liquid-
junction potential.

POLAROGRAPHIC REDUCTION OF METAL IONS

227

-Log [H2C0

Figure 1. Half-wave potential, Ej/2 for the Cu+2, Cu+1 wave in acetone as a function of
-log [H2O] : Curve A 0.1 M LiC104 supporting electrolyte; Curve B 0.1 M
(C2Hs)4NC104, supporting electrolyte.

The graphically evaluated formation constants from the 2 sets of data are
listed in Table 2.

Because the graphically evaluated constants are at best good approximations
of the dissociation constants, better values were calculated through the method
of successive approximations. The average calculated values, using Equation 1,
for log k are presented in Table 3.

228

THE TEXAS JOURNAL OF SCIENCE

TABLE 2

Graphically Evaluated log Formation Constants
for Aquo-Cu+2 Complexes in Acetone.

0.1 A/LiC104

0.1 MEt4NC104

log k 1 = 2.00

logkj = 1.93

log k2 = 1.52

log k2 = 1.37

log k3 = 1.05

log k3 = 0.84

log k4 = 0.64

log k4 =0.33

v co n run i n x E H2°] [H20]n

•E1/2 = E° - 0.059 log [1 + -jjj- + • • • K[ . . . v

where

[Cuy] [H2Q] _j_

K-l — n ~ ir

[Cu(H20)*] Nl

and

K,

[Cu(H2)N-1 ] [H2Q]

[Cu(H20#n] Kr

and

Kr+1 +2

E°‘ =E°+ 0.059 log C“ ^uf "2°

KCu+2 Cu+1

(1)

(2)

(3)

(4)

TABLE 3

Average Values of pk log K Obtained with Equation 1 by the
Method of Successive Approximations Aquo-Cu+2 Complexes in Acetone.

0.1 M LiC104

0.1 M Et4NC104

log k ! = 1.75

logkj = 1.73

log k2 = 1.52

log k2 = 1.27

log k3 = 0.99

log k3 = 0.80

log k4 = 0.73

log k4 = 0.66

SUMMARY

From the results of the data on the Cu+2/Cu+1, Cu+1/Cu° reactions in the presence
of 0.1 M UC104 or 0.1 M tetraethylammonium perchlorate, the following infer¬
ences can be drawn.

1. Although Lf 1 ion has a higher affinity for water than tetraethylammonium
ion, the formation constants for aquo-Cu+2 complexes in acetone are very
nearly alike in the presence of LiC104 and tetraethylammonium perchlorate.

POLAROGRAPHIC REDUCTION OF METAL IONS

229

2. Similarly, ion-ion interactions between Lf 1 and perchlorate and tetraethyl-
ammonium and perchlorate would be expected to affect the stability constants
of aquo-Cu+2 complexes in acetone containing these perchlorates. No such
effect is observed.

3. Finally, it is of interest to note that Larson and Iwamoto (1962) have obtained
similar results in their study of the formation constants of aquo-Cu+2 complexes
in nitrome thane with and without added electrolyte. These investigators found
that the average individual formation constants of the aquo-Cu+2 complexes
obtained by electrochemical means with 0.1 M tetraethylammonium perchlorate
as supporting electrolyte agreed very well with the average individual formation
constants obtained by spectrophotometric means without any added electrolyte.

LITERATURE CITED

Arthur, P., and H. Lyons, 1952 -Anal. Chem., 24:1422.

Coetzee, J. F., and Wei-San Siao, 1963 -Inorg. Chem., 2:14.

Kelly, M.T., H.C. Jones, andH.C. Fisher, 1959 -Anal. Chem., 31:1475.

Kolthoff, I. M., and J. F. Coetzee, 1957-/. Amer. Chem. Soc., 79:870.

Laitinen, H., 1960 -Chemical Analysis. McGraw-Hill, New York, N.Y., p. 107.

Larson, R. C., and R. T. Iwamoto, 1960-/. Amer. Chem. Soc., 82:3242.

- , and - , 1962 -Inorg. Chem., 1:316.

Migal, P. K., Ya. I. TurVan, and N. I. Bondarenko, 1956 -Zh. Fiz. Khim., 30.

Mitchell, J., and D. M. Smith, 1958 -Aquametry. Interscience Publishers, New York,N.Y.,
pp. 146-51.

Nelson, I. V., and R. T. Iwamoto, 1962-Inorg. Chem., 1:151.

Ozerov, A. M., and A. V. Yakovleva, 1956 -Chem. Abstr., 50:10565.

- — - , and - , 1956 -Zh. Priklad. Chem., 29:124.

Schaap, W. B., R. E. Bayer, J. R. Siefker, J. Y. Kim, P. N. Brewster, and F. C. Schmidt, 1961 —
Rec. Chem. Progr., 22:197 .

Schober, G., and V. Gutmann, 1958 -Monatsh, 89:401.

— , and -

- , 1959-Z. Elektrochem., 63:274

— , and -

- , 1960-Z. Anal. Chem., 173:2.

, and

- , 1961 -Monatsh., 92:292.

Vogel, A. I., 19 5 6 -Practical Organic Chemistry. 3rd Ed., Longmans, London.

Weissburger,. A.W., and E: S. Proskauer, 1955 -Organic Solvents. 2nd Ed., Interscience, New York , N.Y.

EFFECTS OF SIMPLE SUGARS ON CERTAIN SERUM CONSTITU¬
ENTS AND ERYTHROCYTE OSMOTIC FRAGILITY IN RATS1

by M. E.PURDOM,Ph.D.\ M. BAYER, B.S.2, S.J. NORTON, Ph.D.1,
and P. T. SULLIVAN, Ph.D.1

School of Home Economics and Departments of Chemistry and
Basic Health Sciences, North Texas State University, Denton 76203

ABSTRACT

Four groups of rats were maintained on isocaloric, isonitrogenous, chemically defined
diets differing only in carbohydrate source, i.e., glucose, fructose, sucrose, and a combination
of glucose and sucrose. The effects of such dietary alteration were examined with respect to
the following: serum proteins, serum glycoproteins, serum haptoglobin, serum bilirubin, as
well as erythrocyte osmotic fragility. The results of these studies indicate abnormal test values
for those animals on the fructose diet.

INTRODUCTION

In previous studies in this laboratory (unpublished data), we have observed
that rats fed diets containing different amounts of sucrose and fats showed unusual
serum glycoprotein profiles when compared with profiles reported by other workers
(Cockerell and Beisel, 1973). The greatest percentage of protein-bound carbohy¬
drates was found to migrate as a2 glycoproteins, regardless of the amount and
type of sucrose or lipid offered in the diet. Considerable interest has been shown
by other investigators in serum proteins, glycoproteins, and particularly hapto¬
globin, an ce2 glycoprotein, in man and animals (Brus and Lewis, 1959; Deutsch
and Goodloe, 1945; Fink, et al, 1967; Krauss and Sarcione, 1963; Nyman, 1959;
Pearce, et al, 1964; Pigman, 1969; Purdom, et al, 1972, 1973; Spiro, 1963).
However, the influence of a chemically defined high carbohydrate diet on these
parameters has not been reported.

In addition to the high a2 glycoprotein levels found in our previous studies, a
considerable degree of erythrocyte hemolysis in rats maintained on diets contain¬
ing a high sugar content has been observed (unpublished data ; Purdom, e tal., 1 972,
1973). Thus, under the above dietary conditions, there may exist a relationship
between serum haptoglobin, the hemoglobin binding protein with the highest
carbohydrate content of the a2 glycoproteins (Berk ,etal, 1969;Bieri and Evarts,
1975; Bieri and Poukka, 1970; Brus and Lewis, 195 9; Fink ,etal., 1967; Grodsky,

Supported by Faculty Research Grants, No. 35854 and 35 135, North Texas State University.
Research assistant, present address: Dietetic Internship, School of Allied Health, University
of Texas, Southwestern Medical School, Dallas 75219.

Accepted for publication: October 16, 1975.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

232

THE TEXAS JOURNAL OF SCIENCE

etal, 1962; Kraus and Sarcione, 1963; Nyman, 1959; Ostrow, et al., 1962,1963;
With, 1954), serum bilirubin, a metabolite of hemoglobin , and erythrocyte hemo¬
lysis.

The investigation reported herein was designed to determine the effects of
feeding Sprague -Dawley male rats isocaloric, isonitrogenous chemically -defined
diets containing a high percentage of glucose, fructose, and sucrose or a combination
of equal amounts of glucose and sucrose as a sole source of carbohydrate. A
moderate level of corn oil was chosen as the lipid component of diets. The influ¬
ence of these diets was monitored for their effects on the following: serum pro¬
teins; serum glycoproteins, serum haptoglobin; serum bilirubin; and erythrocyte
osmotic fragility.

METHODS

Twenty male Sprague -Dawley rats were divided into 4 groups of approximately
equal average weight. The average starting weight of the entire 20 animals was
152.8 gm with individual weights ranging from 136 to 164 gm. Each animal was
kept in a stainless steel metabolic cage and fed daily (16-20 gm) of isocaloric, iso¬
nitrogenous chemically defined test diets (Table 1) for 28 days. Water was given
ad libitum. All animals were weighed on the same day each week. The daily food
offered to each animal was recorded and the uneaten food from each cage was
collected and weighed to determine the amount actually consumed.

TABLE 1

Composition of Experimental Diets.3

Constituents

Diet

1

2

3

4

Amino acid mixture13

139.95

139.95

139.95

139.95

Alphacel (hydrolyzed)

50.00

50.00

50.00

50.00

Choline chloride

2.00

2.00

2.00

2.00

Salt mixture RHb

40.00

40.00

40.00

40.00

Vitamin mixture0

5.00

5.00

5.00

5.00

Corn oil (Mazola)

101.05

101.05

101.05

101.05

Cerelose (D-glucose)d

662.00

----

331.00

Fructose

662.00

----

----

Sucrose

----

—

662.00

331.00

aNitrogen content of all diets was 2.24%. All diets contained approximately 4.0 kcal/gm.
bRogers and Harper (1965).

cPer kilogram vitamin mixture: Vitamin A, 900,000 I.U.; Vitamin D, 100,000 I.U.; alpha
tocopherol, 5.0 gm; ascorbic acid, 45.0 gm; inositol, 5.0 gm; choline chloride, 75.0 gm;
riboflavin, 1.0 gm; menadione, 2.25 gm; p-aminobenzoic acid, 5.0 gm; niacin, 4.5 gm;
pyridoxine hydrochloride, 1.0 gm; thiamin hydrochloride, 1.0 gm; calcium pantothenate,
3.0 gm; biotin, 20 mg; folic acid, 90 mg; Vitamin Bi2, 1.35 mg.
dCerelose Corn Products Refining Co., Argo, Illinois.

EFFECTS OF SIMPLE SUGARS

233

At the termination of the experiment, all animals were fasted for 14 hr, anesthe¬
tized with absolute ether, and blood was drawn into sterile syringes following
standard cardiac puncture. Approximately 2-3 cc of whole blood from each rat
was transferred to separate vacutainers containing 0.04 ml of a 15% solution of
EDTA (tripotassium salt) and 0.008 mg potassium sorbate and immediately re¬
frigerated. These blood samples were employed in the studies of osmotic fragility.
All other blood drawn from each rat was transferred into individual sterile test
tubes and immersed in a 37 C water bath until firm and fully retracted clots formed.
The clotted blood samples were centrifuged at full speed in a clinical centrifuge
for 30 min. The serum was withdrawn and transferred to separate containers
containing 0.04 ml of the above EDTA solution.

Total serum proteins were determined using a calibrated T/C refractometer.1
Serum proteins were electrophoresed on Titan III cellulose acetate plates2 and
stained with Ponceau S according to a published procedure (Chin, 1970). Serum
glycoproteins were electrophoresed on Titan III cellulose acetate plates in like
manner and stained with Schiffs reagent (Kelsey, et al., 1964). The resulting
electrophoretograms were scanned in a densitometer3 with a digital integrator.

Serum haptoglobin levels were determined by single radial immunodiffusion
employing the Behring Diagnostics M-Partigen Haptoglobin Accupak™ Kit4. A
standard curve was constructed from Behrings Protein Standard Serum B which
is a stabilized pooled human serum consisting of 3 pre-diluted standards contain¬
ing 68 mg%, 136 mg%, and 260mg% haptoglobin respectively. The Mancini simplified
end point methodology was then utilized for quantitation (Mancini, etal., 1965).

The total serum bilirubin and erythrocyte osmotic fragility determinations
were conducted according to the method of Davidson and Henry (1969).

Means and standard deviations are reported in each table to show the variability
within the diets. Means were compared by TuKey’s multiple range test at the 5%
level of significance (Winer, 1971).

RESULTS AND DISCUSSION
Growth Response

The average weight gain per day of 5 animals maintained on the test diets
(Table 1) was: Diet 1 , 5 .2 g; Diet 2, 4.6 g; Diet 3, 5.2 g; and Diet 4, 5.2 g. The
overall total consumption of the food offered animals on Diets 1 , 3, and 4 averaged
96, 91, and 94% respectively, while those animals on Diet 2 averaged 88% food
consumption. Animal weight gain corresponds well with food consumption.

1 Model 10402, American Optical Corporation, Buffalo, New York.

2

Helena Laboratories, Beaumont, Texas.

3Photovolt Model 542A.

4M-Partigen Haptoglobin Kit (No. 12-749-001), Behring Diagnostics, Sommerville, New Jersey.

234

THE TEXAS JOURNAL OF SCIENCE

Serum Proteins

The distribution of serum proteins obtained electrophoretically is presented
in Table 2. Plasma volumes were approximately the same for all animals. The
albumin, alpha! , alpha2, and gamma globulin fractions displayed no statistically
significant changes with respect to the various diets. However, differences were
noted among the following: total protein, beta globulins, and the albumin/globulin
(A/G) ratio. The smallest total protein values were observed for animals on the
glucose diet; the beta globulin fraction varied according to diet, with the least
value observed for Diet 3 animals and the greatest for Diet 2 animals. Perhaps of
significance is the fact that fructose -fed animals (Diet 2) had the lowest A/G ratio,
1 .4, while those animals on the sucrose diet had the highest A/G ratio, 2.2.

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EFFECTS OF SIMPLE SUGARS

235

Serum Glycoproteins

While the alphaj and beta glycoprotein levels did not significantly vary with
dietary regimen, the alpha2 glycoprotein levels exhibited a significant variation
with Diets 2 and 3 (Table 3). Thus, those animals on the fructose diet (Diet 2)
had a higher alpha2 value than those on the sucrose diet (Diet 3). Furthermore,
the alpha2 glycoprotein levels were very high when compared with the levels pre¬
viously reported for rats by Cockerell, et al, (1973) who stated that serum glyco¬
protein distributions from rats maintained on a regular rat laboratory chow, ranged
from 2.7% albumin; 53.4% alpha! glycoprotein; 15.7% alpha2 glycoprotein; 17.4%
beta glycoprotein; and 10.8% gamma glycoproteins. These findings suggest that
alterations in serum alpha2 glycoprotein levels may be affected by the amount
and type of mono- or disaccharide in the diet.

TABLE 3

Effect of Experimental Diets on Serum Levels of
Glycoproteins, Haptoglobin, and Bilirubin.

Serum Concentration of Serum Constituents3

Determination

Diet 1 (G)

Diet 2 (F)

Diet 3 (S)

Diet 4 (G-S)

Alphai Glycoprotein

15.20 ±

1.48

8.8 ±

8.60

17.4 ± 5.59

15.2

±

8.04

Alpha2 Glycoprotein

74.8 ±

2.86

79. 0b ±

4.53

66. 8C ± 3.27

70.6

±

6.69

Beta Glycoprotein

10.0 ±

2.82

12.2 ±

4.83

15.8 ± 5.63

14.2

±

1.92

Haptoglobin

238.2 ±

115.05

463.2 ±

121.46

370.8 ±42.72

286.2

±

76.65

Bilirubin

0.17 ±

0. 14

0.30 ±

0.26

0. 14 ± 0.10

0.00

+

0.00

aGlycoprotein concentrations are expressed as%of total; Haptoglobin and bilirubin concen¬
trations are expressed as mg %.

b,c Means are significantly different from one another (P < 0.05).

Additional electrophoretic studies were conducted to elucidate the nature of
the high alpl^ glycoproteins. Rather high levels of haptoglobin, a hemoglobin¬
binding protein which migrates with the alpha2 fraction were observed (Table 3).
There appears to be no significant correlation of the haptoglobin level with the
type of carbohydrate in the diet, even though there is an apparent trend toward
higher haptoglobin in animals fed fructose (Diet 2).

Serum Bilirubin and Erythrocyte Osmotic Fragility

The levels of serum bilirubin (Table 3) varied from 0.0 to 0.3 mg%. The latter
value, obtained for fructose -fed animals, parallels the results for the haptoglobin
studies. However, this determination may not accurately reflect the true bilirubin
content due to the large standard deviation. Several investigators (Berk, et al,
1969; Grodsky, et al, 1962; Ostrow, et al, 1962, 1963; With, 1954) have shown
that hepatic uptake and biliary excretion of bilirubin are much more rapid in
rats than in man due to the relatively greater size of the liver, a higher metabolic
rate, and differences in hepatic blood flow. Thus, serum bilirubin levels may not
respond to a high dietary intake of mono- or disaccharides.

236

THE TEXAS JOURNAL OF SCIENCE

The response of erythrocytes from rats on the different diets to in vitro hypo¬
tonic saline solutions is presented in Table 4. Although other investigators have
studied rat erythrocyte hemolysis in relation to the ratio of tocopherols to poly¬
unsaturated fatty acids in the diet (Bieri and Evarts, 1975; Bieri and Poukka,
1970), published studies are not available revealing rat erythrocyte osmotic fra¬
gility in hypotonic saline solution. Thus, the only comparison that could be made
in this investigation was to that of hemolysis in man reported by Davidson, etal.,
(1969) shown in Table 4. No erythrocyte response was noted for NaCl concen¬
trations ranging from 0.85 to 0.60%. However, upon varying the NaCl concen¬
tration from 0.55 to 0.30%, erythrocyte hemolysis occurred in all animals. Of all
the groups of animals, those on Diet 2 (fructose) displayed an apparently greater
tendency toward erythrocyte hemolysis (Table 4).

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SUMMARY

The effects of dietary carbohydrates
were studied on certain serum con¬
stituents (e.g., total protein, albumin,
the globulins, glycoprotein distribu¬
tion, haptoglobin, bilirubin) as well
as on erythrocyte osmotic fragility.
Rats were maintained for 28 days on
isocaloric, isonitrogenous, chemically
defined diets which differed only in
the type of carbohydrate (Diet 1,
glucose; 2, fructose; 3, sucrose; 4,
equal amounts of glucose and
sucrose).

Analysis of the serum proteins
revealed: (a) total protein levels
were least for animals on Diet 1 (glu¬
cose), (b) the A/G was lowest for
animals on Diet 2 (fructose); (c) the
A/G was highest for animals on
Diet 3 (sucrose); and (d) beta globu¬
lin was highest for animals on Diet 2
(fructose).

Serum glycoprotein electrophoresis
revealed high alpl^ glycoprotein
levels (% of total) with the highest
being observed for animals on the
fructose diet (Diet 2). The serum
haptoglobin and bilirubin levels were

EFFECTS OF SIMPLE SUGARS

237

also highest for those animals on the same diet. Lastly, it was found that the eryth¬
rocytes of these animals (fructose diet) were the most susceptible to hemolysis
in hypotonic saline solution.

LITERATURE CITED

Berk, P. D., R. B. Howe, J. R. Bloomer, and N. I. Berlin, 1969-Studies of bilirubin kinetics
in normal adults./. Gin. Invest., 48:2176.

Bieri, J. G., and R. P. Evarts, 1975 -Vitamin E adequacy of vegetable oils./. Am. Diet. Assoc.,
66:134.

and R. K. H. Poukka, 191 0-In vitro hemolysis as related to rat erythrocyte con¬
tent of QKocopherol and polyunsaturated fatty acids./. Nutr., 100:557.

Brus, I., and S. M. Lewis, 195 9 -The haptoglobin content of serum in haemolytic anemia.
Brit. J. Haemat., 5:348.

Chin, H. R, 197 0 -Cellulose Acetate Electrophoresis - Techniques and Applications Humphrey
Science Pub., Inc., Ann Arbor, Mich., pp. 46, 67-68.

Cockerell, G. L., and W. R. Beisel, 1973-Comparison of serum protein-bound carbohydrate
and glycoprotein patterns of man, monkey and rat. Brit. J. Exp. Path., 54:49.

Davidson, I., and J. B. Henry, 1969 -(Todd-Sanford), Ginical Diagnosis by Laboratory
Methods, 14th ed. W. B. Saunders Co., Philadelphia, Pa., pp. 149-15 1, 701-702.

Deutsch, H. F., and M. B. Goodloe, 1945 -An electrophoretic survey of various animal plasmas.
/. Biol. Giem., 161:1.

Fink, D. J., M.D., L. D. Petz, M.D., and M. B. Black, M.D., 1967-Serum haptoglobin. J.A.M.A.,
199:615.

Grodsky, G. M., J. V. Carbone, R. Fanska, and C. T. Peng, 1962-Tritiated bilirubin: Prep¬
aration and physiological studies. Am. J. of Physiol., 203:5 32.

Kelsey, P. L., T. P. de Graffenreid, and R.C. Donaldson, 1964-Electrophoretic fractionation
for serum glycoproteins on cellulose acetate. Gin. Giem., 11:1058.

Krauss, S., and E. J. Sarcione, 1963-Synthesis of serum haptoglobin by the isolated perfused
rat liver. Biochemica et Biophysica Acta., 90:301.

Mancini, G., A. D. Carbonara, and J. F. Hermans, 1965 -Immunochemical quantitation of
antigens by single radial immunodiffusion . Immunochem., 2:236.

Nyman, M., 1959-Variation of haptoglobin level and clinical value of haptoglobin determin¬
ation. Scand. J. Gin. and Lab. Invest., ll(supp. 39): 153.

Ostrow, J. D., J. H. Jandl, and R. Schmid, 1962-The formation of bilirubin from hemoglobin
in vivo. J. Gin. Invest., 41:1628.

- — — - , R. Schmid, and D. Samuelson, 1963-The protein binding of C14-bilirubin in

human and murine serum./. Gin. Invest., 42:1286.

Pearce, R. H.,E. M. Watson, R. Stodolski, J. M. Mathieson, and J. J. Theoret, 1964-Effects of age
and sex on the electrophoretic fractions of the serum glycoprotein. Gin. Giem., 10: 1066.

Pigman, W. W., 1969-Glycoproteins: Synthesis in two steps. C&E News, 47:11.

238

THE TEXAS JOURNAL OF SCIENCE

Purdom, M. E., K. Hyder, and M.D.Pybas, 1973-Effects of saccharin on rats fed chemically
defined diets./. Am. Diet. Assoc., 63:635.

- , N. H. Mondragon, W. W. Pryor, and R. W. Gracy, 1972— Effect of carbohydrate

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87:267.

Spiro, R. G., 196 3 -Glycoproteins: Structure, metabolism, and biology. N. Eng. J. Med.,
269:566.

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With, T. K., 1954 -Biology of the Bile Pigments. Arne-Frost-Hansen, Copenhagen, p. 210.

RAYLEIGH- SCHROED IN GER PERTURBATION CALCULATIONS
FOR BERYLLIUM1

by G. P. SAXON2 and D. L. HARDCASTLE

Baylor University, Waco 76703

ABSTRACT

The ground-state energy for beryllium through 2nd order has been calculated using de¬
generate Rayleigh-Schroe dinger perturbation theory. The perturbation terms were the inter-
electronic potentials. The wave functions used were antisymmetric products of 1 -electron
atomic eigenfunctions, and were also eigenfunctions of spin. The eigenvalues and eigenvectors
of the 1st- order perturbation matrix were obtained. The true zero-order ground- state eigen¬
function was then approximated and the 2nd-order energy was obtained. The approximation
was the representation of the ground-state by the state Is2 2s2 The total energy through

2nd-order was E0 = -14.65885 which is 0.06% above the experimental energy.

INTRODUCTION

In the past few years, many new and powerful methods have been applied to
the calculation of energy levels of atoms with small values of z. These investiga¬
tions can be classified according to the type of approximate wave function used
as follows:

1 . Single -particle orbital

a. Hartree-Fock

b. Hartree-Fock -Slater

2. Correlated wave function

a. Configuration-interaction

b. Interelectronic coordinates

3. Perturbation

a. Rayleigh-Schroedinger (R-S)

b. Brueckner-Goldstone (many -body).

The single -particle orbital method can be further classified as to whether the
wave functions used contain variational parameters which are adjusted so as to
obtain the minimum total energy for the system, or the variational principle is
used to obtain a set of integrodifferential equations which are solved self- consistently,

Supported in part by a Faculty Research Grant from Baylor University.

2 Present Address: Amoco, Houston 77001.

Accepted for publication: J une 17, 1975.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

240

THE TEXAS JOURNAL OF SCIENCE

yielding the wave function for the system. Recent many-body calculations have
used the Hartree-Fock orbitals as a basis set for their perturbation expansion. As
would be expected, the perturbation method yields significantly improved results.

The correlated wave function approach is, like the single -particle orbital method ,
a variational method and involves the expansion of the wave function in terms of
either the interelectronic coordinate, ry, or the spherical harmonics, Y™ (0, $),
in addition to single particle terms and variational parameters.

Normally, Hartree-Fock calculations will give a total energy for the system
which differs from the experimental value by less than a few %. Perturbation cal¬
culations and correlated wave function calculations will give results 1 or 2 orders
of magnitude better than this, depending on the number of terms used, basis set,
etc.

Froese (1967) has used the Hartree-Fock method for calculation of energy levels
for 2- and 4-electron systems. Barnett and Shull (1967) and Barnett and Platas (1968)
have used the Reduced Density -Matrix theory to obtain the ground-state energy
of beryllium. Kelly (1963) used the many-body perturbation theory to obtain
the ground state of beryllium to within 0.03% of the experimental energy. Weiss
(1961) used the configuration-interaction approach to obtain results for the ground
state of beryllium which were within 0.05% of the experimental value. Tuan and
Sinanoglu (1964) used a many -electron theory to obtain a value for the ground-
state energy of beryllium which is in excellent agreement with the experimental
value. The above successes together with the supposed difficulty of the Rayleigh-
Schroedinger perturbation method have probably caused the neglect of perhaps
the oldest of the approximate methods. It is the purpose of this paper to show
that the R-S method is indeed viable and can give surprisingly good results, even
when applied only through 2nd order.

DEVELOPMENT OF THE PERTURBATION SERIES

The atomic system chosen is the Be atom, and the level chosen is the ground
state. The Hamiltonian in atomic units is

H = H0 +V,

(i)

where

i 4 4

H0 = -- 2 V!-Z1

O . . i ■ + - ’

2 i= 1 • i=l rj

(2)

and

V

(3)

RAYLEIGH -SCHROEDINGER PERTURBATION CALCULATIONS

241

It is required that the eigenfunctions of H0 be antisymmetric and also be eigen¬
functions of spin. The ground -state of beryllium (spectroscopic designation) is
Is2 2s2 %, indicating a spin of zero and a spin projection of zero. The 4-particle
spin eigenfunctions which satisfy zero spin and zero spin projection are

=2 ~ 0i <*2 /34 -Pi0i2P30i4] , (4)

and

\2 [2 ~ O' ^2^3 @4 1^2^3a4 "01a2a3$4

-@i0i2@3tt4 + 2j31j32a30'4] • (5)

From these spin functions, it is readily deduced that possible eigenfunctions of
H0 which satisfy both the Pauli Exclusion principle and the spin requirements are

where the a, b, c, and d represent 1 electron atomic eigenfunctions and

ajtti a2 ol2 a3a3 a4G!4

f abcdj __ 1 bx jSj b2 /32 b 3 /33 b4/34

^ CiOi! c2a2 c3a3 c4a4

(8)

4i/3i d2/32 d3/33 d4/34

The and N2i are the normalization coefficients. By expanding the determinants,
one can show that

<£>

li

= Nli
v5?

(2A-B-Q + /3 ‘x^(C-B)

(9)

and

$2i = ^i [‘x2 (2/3G-/3H-/3 I)+‘x‘(3H-3I)] , (10)

242

THE TEXAS JOURNAL OF SCIENCE

where

A = abcd + bacd + abdc+badc+cdab + dcab + dcba+cdba,

B = acbd + bead + adbc+bdac+cadb + cbda + dacb + dbca,

C = aedb + bed a + adeb + bdea + cabd + ebad + dabc + dcab,

G = abed + edab - abde-edba - bacd - dcab + bade + deb a,
H=adbc + cbda-acbd - cadb + bead + dacb - bdac - dbca,
and

I = aedb + cabd - adeb - ebad - bcda-dabc + bdca + dbac.

The subscripts indicating the particle coordinates have now been dropped and
the coordinates will be designated by position. Thus

abed = aj b2 c3d4 .

Eigenfunctions have now been found which are both antisymmetric and eigen¬
functions of spin. In particular the eigenfunctions have a spin of zero and a spin
projection of zero.

It is easily shown that and <f>2i are orthogonal only when none of the a, b,
c, d, is equal to another of the a, b, c, d. Whenever any 2 of the a,b,c,d are equal
$li = $2i, but if 3 or 4 of the a, b, c, d are equal then ^ . = $2i = 0. The only other
possibility is for the a, b, c, d to form 2 equal pairs in which case $2 . = 0 but ^ . =£ 0 .

The ground state eigenfunction is obtained by the choice a = Uj 0o> b = Vl 00,
c = U2 o o and d = U2 o o . Then Equations (6) and (7) become

$io” 4Ni o

Ui oo
a

Ui oo

P

U

2 00
a

U

7

O’

and

(ii)

$2 0 ” 0.

The normalization coefficients are easily calculated and are

N. . = 1/(4 - 2A - 2 A, - 2A. . + 4A . + 4A . + 4A KA .

li /v ac be bd ab cd ab cd

(12)

+ 4A A,, + 4A ,A, V

acbd orl ^

ad bc;

and

(13a)

N2j = 1 / [6(2 - 2Aab - 2Acd + Aac + Afed + Abc + Aad + 2AabAcd

+ 2A .A. + 2A A, .

ad be ac bd

)]

1/2

(13b)

RAYLEIGH - SCH ROEDINGER PERTURBATION CALCULATIONS

243

where A u = 5 > 8n n> 8 > whenever a = U n and b = U v >.

ab nn ££ mm n£m n£m

The zero order ground state energy will now be found. Thus

H $> = E
ol o o

(0)

V

(14)

or

E(0) = <WV

0 - - — —

<<t> 1$ >
*01*0

(15)

_ 1

24[<2D-E-F|Ho|2D-E-F> + 3< F-E|H |F-E>], (16)

where

and

D = U(1) U(2) U(3) U<4) + IJ(1) U(2) II(3) II(4)

LJ i o 0 ^10 0 ^2 0 0 ^2 00 T U 2 0 0 ^200 ^100 U j qq ?

(1) (2) (3) (4) (1) (2) (3) tjW)

JC “ U i o 0 '-^200 ^100 ^ 2 0 0 ^200 U j q q U 2 o 0 u 1 00 >

F-II(1) TI° n(3) TT(4) + IT(1) TT(2) i t(3) tt(4)

r “ u j g g ^200 ^200 ^ 1 0 0 ” U 2 g g ^100 U | g g ^200°

Equation (16) is easily evaluated and gives

n

E(o0)=(-

|<2D-E-F|2D-E-F> + 3<F-E|F

2 2 8 8

-E>]

/ 24

_ 5z

and for z = 4,

E(°> = -20.

It is found that the 10 eigenfunctions of Hoisted in Table 1 also correspond
to this energy. The ground state is thus degenerate, and degenerate perturbation
theory must be used. The first order correction (Kelly, 1968) to the energy and
the expansion coefficients for the proper wave function are obtained by deter¬
mining the eigenvalues and corresponding eigenvectors of the perturbation matrix

Vi

-E™ 5 = 0,

ninj

n

(17)

where

244

THE TEXAS JOURNAL OF SCIENCE

TABLE 1

Ground State Eigenfunctions.

a

b

c

d

$

Uioo

Uioo

U200

U200

$01

Uioo

Uioo

U200

U21-1

$02

Uioo

Uioo

U200

U2 10

$03

Uioo

Uioo

0

0

Oi

D

U2 11

$04

Uioo

Uioo

U21-1

U2 i"i

$os

Uioo

Uioo

U21-1

U2 10

$06

Uioo

Uioo

U2 1-1

U21 1

$07

Uioo

Uioo

U2 10

U210

$08

Uioo

Uioo

0

cj

D

U211

$09

Uioo

Uioo

U2U

U211

$10

Using the notation of Table 1 and for i “j = 1 Equation 18 becomes

V,1 = <$01|V|$01>

=2- ^<2D-E-F|V|2D-E-F> + 3<F-E|V|F-E>)

=i [<d|v|d>+<e|v|e> + <f|v|f>-<d|v|e>-<d|v|f>

- <E|V|F>] . (19)

After expansion and integration, Equation (19) becomes

V = 5 z + 77z + 68z 32z

11 8 5 12 81 ~ 129 5

and for z = 4

(20)

Vn =6.28400420 (21)

Equation (18) is given explicitly in Table 2.

The diagonalization is easily performed and the results are given in Table 3.
Using the results of Table 3 the proper ground state eigenfunction of HQcan
now be written. It has the form

i 10

$0 = 2Ci$oi,

where the C2s make up the eignevector corresponding to the lowest eigenvalue

6.284004 0 0 0 0 0 -0.165728 0.117187

0 6.730950 0 0 0 0 0 0

0 0 6.730950 0 0 0 0 0

0 0 0 6.730950 0 0 0 0

RAYLEIGH-SCHROEDINGER PERTURBATION CALCULATIONS

245

co

oo

OOOOOOOOOOO

00

os

Os

m

''t

OOOOOOOOrfO

O

Os

VO

SO (N

SO o

O O os -H o o
in m
p p
a r~

Os <N

O so
<N so

O O CO Os O O
t-~ m

o o

Os

in

'3-

o-^-oooo

o

os

o o o o o

o o o o o o

o o o o o o

o o o o o o

oo r-

<N 00

o o in c-~ o o

SO —I

??

CO oc
fcj W

u

L>

U

U

U

U

oooooooo®— <

OOOOOOOO’-HO

o o o o o

SO

00

CO o o o o o

00

o o o o o

© O O O — < ©

OO—IOOO

© ^ © © © ©

o o o o o

t-~ O O O tJ- Os
Os in wo »n co io
oosososoo^t
r- 0 0 0 00'^-
CO CO CO CO oo O
(N t — ^ t — ; t : Os OS
SO so SO SO *0 so

7.162304 -0.225165 0 0 0 0 0 0.795529 -0.562524

6.988834 0 0 0 0 0 0 0.577350 0.816497

6.904459 0 00000 0 0

6.988834 0 00000 0 0

246

THE TEXAS JOURNAL OF SCIENCE

of Equation (17). Explicitly

=0.9743201 001 + 0.1838465$O7

+ 0.1299991

Uo

The 2nd order ground-state energy may now be obtained from the well-known
equation

e(2) = s'.

0 i,m

E(0) -E(0)
o m

(22)

where i = 1 or i = 2 and the prime on the £ indicates that the sum over the

r ii ii ii

does not include any of the $ .’s. Since V $. >, «J>._ V $. > and <$AQ V <t>. >

J oi on i im 07 i i im os i i im

are all close to the same value, and since approximately 95% of the energy contri¬
bution will come from <$01 |V |$. >, the approximation is made that C } - 1 ,
C7 ■- C8 = 0. Then Equation (22) becomes

Eo2) = 2'

ILL

>

1 m

e(°)_e(°)

o m

+ £'

«i>

QJL

>

2m

E(0) _p(0)

0 m

(23)

where the £ also includes integration over continuum states.
It is possible to calculate E^. The result is

E(°> = -£
cm 2

(24)

where n1? n2, n3, and n4 are the principal quantum numbers of the 1-electron
atomic eigenfunctions which make up the $im .

The matrix elements of Equation (23) may now be expanded and rewritten
using symmetry arguments. The results are

<%.lVl^lm>=^TIL [<DI^I4A-2B-2C>

+ <E|^— |4B-2A-2C>+ <F I— |4C-2B-2A>], (25a)

rl 2 1 rl 2 1 I

and

<*o. |V |$2m> = ^ [<D |^|H-I> + <E \±- |l-G>

+ <F l-L |G-H>
lr12 1

(25b)

RAYLEIGH-SCHROEDINGER PERTURBATION CALCULATIONS

247

The notation U100 = 1 , U200 = 2, U210 = 3, U21 ^ = 4, and U211 = 5 is helpful.
Then

<U1(o1o)U1(oo)U2(oo)U2(04o)|abcd> = <1122 |abcd> = AlaAlbA2cA2d

Since the terms of the perturbation are 2-particle interactions but the eigen¬
functions of HQare 4-particle eigenfunctions, the matrix elements of Equation (23)
will vanish unless 2 of the a,b, c, and d are either U100 orU200. The Equations (25 a)
and (25b) then become

<$o: |vKm> = ^ (Aia [l6<22|v|cd>Alb-8<22|v|bc>Ald
-8<22|v|bd>Alc+A2d(16< 1 2 1 v|bc> -8< 1 2 1 v|cb >)

+ A2c(16< 12 |v|bd>-8< 12 1 v | db >) +A2b(-8< 12 1 v| cd>
-8<12|v|dc>)J +A2a^16<ll|v|cd>A2b-8<ll|v|bc>A2d
-8<ll|v|bd>A2c+Ald(16<12|v|cb> -8 <12 |v|bc>)

+ Alc(16<12 1 v| db > - 8<12 1 v|bd>) + Alb(-8<12 |v|dc>
-8<12 1 v | cd >)j + A^c |l6<22 1 v | ab> A^ - 8<22 |v |ad > Alb
+A2b(16<12|v|da> -8<12|v|ad>) + A2d(-8<12|v|ab>
-8< 12 |v |ba>)J + A2c[l6< 11 |v| ab > A^ - 8< 11 |v|ad> A2b
+Alb(16<12|v|ad> -8<12|v|da>) + Ald(-8<12|v|ab>
-8<12|v|ba>)J +AlbJj-8<22|v|ac>Ald+A2d(16<12|v|ac>
-8< 12 |v |ca>)J +A2b|j8< 11 |v |ac>A2d+Ald(16< 12|v |ac >
-8<12|v|ca>)jJ , (26a)

and

<$oi|v|$

2m

c> -4<12|v|cd >)+4<12|v|db>A2(
-4<12|v|cb>A2b +4<22 |v|cb>Ald -4<22|v|db>AlcJ
+A2a^lb(4<12|v|cd> -4<12 |v|dc>)+4<12|v|bd>Alc

248

THE TEXAS JOURNAL OF SCIENCE

-4<12|v|bc>Ald+4<ll |v|bc>A2d- 4<11 |v|bd>A2cJ
+Alb[4<12|v|ca>A2d-4<12|v|da>A2c+4<22|v|ad>Alc

-4<22|v|ac>AldJ +A2bj4<12|v|ac>Ald-4<12|v|ad>Alc

+ <11 |v |ad >A2c-4< 11 |v|ac>A2dj +A[cA2d (4< 12|v|ba>

-4<12|v|ab>)+A2cAld(4<12|v|ab>-4<12|v|ba>|j(26b)

For the case of a = 1 , c = 2, and d =£ 1 , 2, Equations (23), (24) and (26a) may
be combined to give

E(2) = 2| <22 | v |2d> + 2<12 | v | Id > - <12 1 v|dl>

(27)

2112

where v = — . Equation (27) may be written more explicitly. In particular
r12

E® = 2 2[<U^U<020>£ |u(0Vu(2)m>+2<l410)ui020>|^|U^U<2)m:

n£m

n> 2

Ij ^ l*

"8 2?

]

(28)

There are 16 other cases similar to Equation (28). These are listed in Table 4, and

(29)

E<,2) = 2 E(2)
0 i=l 01

These series have been evaluated for ££4 and the results tabulated in Table 5
The ground-state energy has now been obtained and the result is

e0 = e<°» + e(1) + e® +

= -20 + 6.237097 -0.83595

= -14.65885 (30)

or in terms of z

E0 = | z2 + 1 ,559274z - 0.83595 .

RAYLEIGH-SCHROEDINGER PERTURBATION CALCULATIONS

249

TABLE 4
The Energy Series.

Case

Series

112n

1 Ipn

?(2) -

'01

02

= 2'

<22

2d> + 2 <12

ld> - <12

dl >

8 2i rr

?(2) 1

, 2 j <22 | v|pn> I'

8 2n2

p = 3, 4, 5

linn

22pn

,(2)

"03

r(2)

"04

n>n i + A

2 <22 v cd>

g;l<‘.l|,|Tr P-J.4.S

- 7z_ + z2
8 Trf

22nn

223p

224p

2255

?(2)

"05

?(2)

"06

"07

?(2)

"08

S' _JL

2 <11 v cd>

n>n/ 1 + A . - z2 + z2 1 1

cd — (~r + t)

2 ni

! 2 | < 1 1 | V | 3p > |2

1 + A

3p

- 3z

?(2) _ 1

2 | < 1 1 1 v 1 4p > |2

1+A4P "if
4

| <11 | v| 55 > |2

- 3z2

p = 3, 4, 5

p = 4, 5

122n

12pn

122p

?(2)

"09

>(2)

010o

= 2

, 2 <11 v ld> + 2<12 v d2>- <12 v 2d>

l_ + z2
2 2r?

<12 | v| pn>+ <12 | V | np>

z2 , z2

p = 3, 4, 5

■(2) _
oiob ~

?(2) =
"Oil

T 2^~

3 | <12 | v | np> - <12 | v|pn> |2

— - + — —

2 2n2

2 |<12 | v| 2p> - 2<12 |v|P2>- <11 |v| lp>

li?

P = 3,4,5

250

THE TEXAS JOURNAL OF SCIENCE

TABLE 4 (Continued)

Case

Series

1233 E'

(2)

012

2 | <12 I v | 33>
8

<12 v 3p>+ <12 v p3> r

123p Eqj2 = — - U - - U - L_ P “ 4, 5

a -

8

m 3 I <12 I v I p3> - <12 I v I 3p> I2

- I II _ ''I n - 4 s

°l3b _ 3z2 P

“T

1244 E'

(2) _

014

2 | < 1 2 | v 1 44 > |2

r3?"~

1245 E

(2)

015o

<12 I v 1 45 > + <12 I v 1 54>

^3?

3 <12 v 45>- <12 v 54>

1255 E

(2)

°1Sk Tsz

8

(2) _ 2 | <12 | v 1 55 > |2

016

~3z ?
8"

12nn' e!?2 = S' — ^

<12 v cd>+ <12 v dc>

a 1+Acd -4 + 4(4^>

8 2 n^ nq

(2)

01?b

| < 1 2 I v I dc> - <12 I v | cd >
2 ~2

c^di-Acd -5|i +^(jr. +4

8 2

This series comes from the 4-determinant excited state wave function,
5This series comes from the 6-determinant excited state wave function.

RESULTS AND CONCLUSIONS

The result obtained in Equation (30) is within 0.06% of the experimental value,
and is compared with other methods of calculation in Table 6. While the above
results are not as close to the experimental value as some of those listed in Table 6,
the result was quite close to the experimental value, and the method has several

RAYLEIGH-SCHROEDINGER PERTURBATION CALCULATIONS

251

TABLE 5

(2)

Energy Contributions to Eq .

Case

BB

BC

cc

Extrapolation

Total

Eqi

-0.50535

-0.211732

0

-0.01231

-0.72940

E02

-0.00034

-0.006749

0

-0.000120

-0.007209

E03

-0.0071829

-0.003136

-0.031177

-0.000803

-0.042299

E04

-0.001047

-0.004977

0

0

-0.006024

E05

-0.0004343

-0.004221

-0.004064

-0.000293

-0.009012

E06

-0.003497

0

0

0

-0.003497

E07

-0.006994

0

0

0

-0.006994

E08

0

0

0

0

0

E09

-0.018124

-0.056885

0

-0.00927

-0.075936

E010a

-0.000541

-0.000363

0

-0.000017

-0.000921

d

E010u

-0.000744

-0.002363

0

-0.000052

-0.003159

Eon

0

0

0

0

0

E012

-0.000884

0

0

0

-0.000884

E°13a

0

0

0

0

0

E013b

0

0

0

0

0

E014

0

0

0

0

0

Eoisa

-0.001767

0

0

0

-0.001767

Eoisb

0

0

0

0

0

E016

0

0

0

0

0

E°i7a

-0.000132

-0.000927

-0.006075

-0.000055

-0.007189

E017b

-0.000004

-0.000645

-0.000989

-0.000026

-0.001664

Total

-0.89595

^his series comes from the 4-determinant excited state wave function.
^This series comes from the 6-determinant excited state wave function.

TABLE 6

Comparison of Results for Beryllium.

Author (Reference)

Result

Method

Froese (1967)

-14.57307

Hartree-Fock

Weiss (1961)

-14.66090

Configuration-Interaction

Kelly (1963)

-14.6640

Many-body Perturbation Theory

Sinanoglu (1964)

-14.66737

Many-electron Theory

Barnett (1967)

-14.6861

-14.6609

Reduced Density Matrix

Barr (1969)

-14.659

Consolidated Configuration-
Interaction Perturbation

Saxon (Present Work)

-14.65885

Rayleigh-Schroedinger Perturbation
Theory (Through Second-Order)

Experimental

-14.667306

252

THE TEXAS JOURNAL OF SCIENCE

inherent advantages. There is nothing sophisticated about the method itself in
that the theory is straightforward and the results are not dependent upon a fortui¬
tous choice of beginning eigenfunctions. Large computing machines are not neces¬
sary for the calculations. Preliminary- calculations for the above were performed
on a Honeywell 1200 with a 4000 word memory and the final calculations were
performed on a Honeywell 1250 wihich has a 10,000 word memory. For the 2nd-
order energy, wavefunctions of any z may be used, thus eliminating much duplication
of effort for atoms with different z.

The R-S method is seen to provide a relatively simple way of obtaining the
ground-state energies of 4-electron systems. Excited states should follow similarly,
and some of the matrix elements obtained in the calculation of the ground-state
will be needed in some of the excited states. The above method can easily be applied
to lithium and other few-electron systems.

LITERATURE CITED

Barnett, G. P., and O. R. Platas, 1968-/. Chem. Phys., 48:4265.

- , and H. Shull, 19 67 -Phys. Rev., 15 3:61.

Barr, T. L., andW.T. Simpson, 1969-/. Chem. Phys., 51:1526.

Froese, C., 1967-/. Chem. Phys., 47:1410.

Kelly, H. P., 1963 -Phys. Rev., 131:684.

- , 1968 -Z? Keith A. Bruckner (Ed.), Advances in Theoretical Physics, Vol.II. Academic

Press, Inc., New York, pp. 97-98.

Tuan, D. F., and O. Sinanoglu, 1964-/. Chem. Phys., 41:2677.

Weiss, A. W ., 1961 -Phys. Rev., 122:1826.

SYNTHESIS OF A RADIOLABELED CARRIER TO PRECIPITATE
ANTI-HAPTEN ANTIBODIES

by A. C. SCHRAM and C. P. CHRISTENSON

West Texas State University
Department of Chemistry
Canyon 79016

ABSTRACT

The synthesis of a simple hapten carrier (l,3-dihydroxy-5-iodobenzene) was developed
for routine use in the precipitation of anti-hapten antibodies. Coupling to a model hapten
(lactose) and quantitative precipitation studies were performed, to apply the compound to
the lactoside chicken anti-lactoside antiserum system.

INTRODUCTION

Pauling, et al. , (1942) have shown that some antibodies can be precipitated
by small carrier molecules covalently bonded to at least 3 haptenic groups. These
synthetic precipitating antigens are chemically defined by their synthesis steps,
while natural antigens, such as polypeptides or polysaccharides, have haptenic
groups whose 3-dimensional structure is more difficult to identify; often, several
different haptenic groups may be present on the same molecule. Our studies of
inhibition of precipitation of antigen-antibody complexes required a chemically
defined precipitating antigen (to relate antibody avidity to hapten structural
formula) which could be precisely assayed in very small quantities (to minimize
the amount of serum used). The first problem we wanted to investigate was the
influence of the aglycone portion of lactosides on the binding by anti-lactoside
antibodies. The chicken had been chosen over the rabbit or the guinea pig to pro¬
duce the anti-lactoside antibodies because chicken anti-lactoside antisera with
consistently high titers were obtained at lower cost.

The most convenient way to covalently bind 3 lactoside haptens to a carrier
was through p-diazophenyl bridges, since p-aminophenyl lactoside could be pre¬
pared (Karush, 1957). An activated aromatic carrier such as a phenol analog was
desirable to increase the yield of tri-substituted product.

Phloroglucinol has been used as a carrier for glucoside, galactoside, and lacto¬
side haptens (Yariv, et al, 1962), but radiolabeled phloroglucinol is not available.
Tritiated phloroglucinol could not be used, since the 3 ring hydrogens are substi¬
tuted by the haptens, and the 3 phenolic hydrogens would be lost through ioni¬
zation at the pH most favorable for coupling. Methyl-14 C methoxy analogs of
Accepted for publication: November 2, 1976.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

254

THE TEXAS JOURNAL OF SCIENCE

phloroglucinol are less soluble and less reactive than phloroglucinol,so we selected
1, 3 -dihydroxy -5 -iodobenzene as the carrier. Although less soluble and less reactive
than phloroglucinol, it could be synthesized with relative ease, and the short
half-life of 131I and 125I minimized decontamination procedures. Of several
possibilities which we tried, the simplest overall pathway of synthesis for the
model compound, 2,4,6-tri-(p-diazophenyl lactoside)-l , 3 -dihydroxy-5 -iodo¬
benzene-1 2 5 1, is outlined below.

3,5-Dimethoxyaniline (2, R = NH2 ; 1.53 g, 10 mmoles) was diazotized and
treated with 1.66 g (10 mmoles) of KI (with or without labeled iodide) to give
0.9 g of 1 ,3 -dimethoxy-5 -iodobenzene (2, R = I), mp 70-3 C, after steam distil¬
lation and recrystallization from ether. A 0.1 g sample of 1 ,3-dime thoxy-5-
iodobenzene was stirred for 1 hr at 120 C withl ml of glacial acetic acid and 1 ml
of 57% hydriodic acid, freshly distilled in the presence ofhypophosphorous acid.
The excess acetic and hydriodic acids were removed under reduced pressure and
the residue was dissolved in ether; the ether solution was washed with 50% (w/v)
aqueous hypophosphorous acid, diluted NaHC03 , and water, and dried over an¬
hydrous CaCl2. Distillation of the ether at atmospheric pressure, followed by
vacuum distillation (0.5 torr) at 120 C, re crystallization from aqueous ethanol
and purification by thin layer chromatography (Rf = 0.88; silica gel G; benzene:
acetic acid = 5:1, v/v; Rf of compound 2 (R = I) was 0.99) yielded 0.02 g of 1 ,3-
dihydroxy -5 -iodobenzene (1), mp 110C. The reported melting point of 1 is 95 C.
(Hodgson and Wignall, 1926); the product we prepared had the proper elemental
analysis and the expected NMR and IR spectra for 1 ,3 -dihydroxy -5 -iodobenzene
monohydrate.

The model antigen, 2,4,6-tri-(p-azophenyl lactoside)-l ,3-dihydroxy -5 -iodo¬
benzene-1 25 1 (3, X = p-N2C6H4OC! 2 H2 x O i ! = Lac) was prepared by coupling
4 mg of 1 , labeled with 300,000 cpm of 1 2 5 1 , and 1 29 mg of freshly diazotized
p-aminophenyl lactoside (carrienhapten ratio of 1 :4) at pH 9-10 and 4 C for 5 hrs;
the rate of reaction was followed by the increase in absorbance at 456 nm, char¬
acteristic of triazophenylbenzene. As soon as the absorbance at 456 nm stabilized,
the reaction was stopped by addition of TV HC1 to pH 5. The product which pre¬
cipitated was recrystallized from 50% aqueous ethanol, yielding 5 mg of 3(X= Lac).

Chicken anti-lactoside antibody (Schram, 1970) was precipitated by Compound 3
(X = Lac) following an ordinary antigen-antibody precipitation pattern (Figure 1).
The serum precipitating anti-lactoside antibody level was 1 .625 mg/ml, measured

M9 Antigen Precipitated -

P° Pi >J

SYNTHESIS OF A RADIOLABELED CARRIER

255

Figure 1. Precipitation of 3(X = Lac) by its specific antibody. The amount of 3(X = Lac)
was calculated from the radioactivity of the precipitates, corrected for antigen
controls without serum; the amount of antibody, by determination of the pro¬
tein in the precipitates (Lowry, et al, 195 3), corrected for controls using the
same levels of antigen and pre-immunization serum. Each point, thus, represents
the results of the reaction between the amount of 3 (X = Lac) shown on the
abscissa and 100 £ll of chicken anti-lactoside antiserum (Schram, 1970), under
conditions previously described (Schram, et al., 1971).

by quantitative precipitation with an 12SI-labeled bovine serum albumin (BSA)
carrier, coupled to an average of 1 1 Lac groups/molecule of carrier. The total
level of lactoside -binding antibody, measured by the Farr technique (Farr, 1958),
was 1.77 mg/ml. Only 1.35 mg of the available antibody in 1 ml of serum was
precipitated by Compound 3 (X = Lac). The shape of the precipitation curve,

256

THE TEXAS JOURNAL OF SCIENCE

however, indicated that all of the antibody which could be precipitated by Com¬
pound 3(X = Lac) had been precipitated: a plateau was reached (Figure 1). Thus,
only 83% of the available precipitating antibody was actually precipitated. Anti¬
bodies of varying avidities are probably present in the antiserum, but at first glance,
it did not seem that avidity differences could be responsible for the difference
between the amounts precipitated by Lacn-BSA and by Compound 3 (X = Lac),
since the Lac group was the same determinant in either case.

A probable explanation is based on the 3 -dimensional restrictions imposed by
the 1,3 -dihydroxy -5 -iodobenzene central structure. In Lac proteins, the Lac
groups are distributed on the surface of the protein carrier wherever a tyrosyl or
a histidyl residue is available (Schram, 1970). Since the protein carrier is much
larger than the 1 ,3 -dihydroxy -5 -iodobenzene carrier, the distance between the
Lac groups on the protein carrier is larger: approximately 50A, assuming a random
distribution on the surface of a BSA molecule with the shape of a 160A x 35 A
ellipsoid (Oncley, 1949).

The 6 diazo nitrogen atoms bound to the phenyl ring of 1 are isoplan ar with
the phenyl ring, because of the conjugation with the central ring’s it electrons.
Although the 3 phenyl rings linking the diazo nitrogen atoms and the Lac groups
are probably tilted (steric hindrance), free rotation would be restricted by con¬
jugation; the lactosyl groups themselves, being in a para position with respect to
the diazo nitrogen atoms, are restricted in rotation to the axis going through the
acetal oxygen atom and the diazo nitrogen atoms, toward the center of the 1,3-
dihydroxy -5 -iodobenzene ring (Figure 2). The net result will be to limit the dis¬
tance between neighboring lactosyl groups to approximately 15 A.

The antibody is complementary for the whole lactosyl group, but the phenyl
ring and the atoms para to the lactosyl group still influence the binding to the
antibody (Paktinat, 1976). In other words, the antibody polypeptide chains in
the combining site envelop more than the lactosyl groups. The thickness of the
polypeptide chain enveloping a lactosyl group may hinder the positioning of 2
other polypeptide chains around the other 2 lactosyl groups in Compound 3
(X = Lac). Thus, Compound 3 (X = Lac) will precipitate only that portion of the
antibody population with shallower combining sites.

There is some evidence for this reasoning: the binding of a different sample
of chicken anti-lactoside serum by a reference lactoside was inhibited by p-nitro-
phenyl galactoside (Schram, manuscript in preparation). However, the inhibition
was weak, and its kinetics indicated that it involved a small proportion of the
antibody population. This particular antibody population was still complementary
to lactose, but with specificity for the galactoside portion of lactose only. Thus,
the antiserum must have antibody populations with various depths of combining
site.

In the serum used for this study, there were probably antibodies with combining
sites of various depths, also. These populations may well be responsible for part
of the avidity differences. Antibodies with shallower combining sites would have

SYNTHESIS OF A RADIOLABELED CARRIER

257

fewer potential hydrogen bonding groups; the binding energy for the antigen-
antibody complex would be lower, and the complex would be more easily dis¬
sociated.

The 17% of the precipitating antibody which was not precipitated by Com¬
pound 3 (X = Lac) may represent a portion of the antibody population with the
deepest combining site. To accommodate all of the antibodies present, it would

258

THE TEXAS JOURNAL OF SCIENCE

then be necessary to locate the Lac Groups further from the geometric center of
Compound 3, perhaps by intercalation of several methylene groups between the
Lac groups and the central phenyl ring.

Since any other haptenic group (X), easily bonded to Compound 1 could re¬
place the Lac group of the example outlined, the carrier described is widely adapt¬
able to those cases in which the determinant is chemically defined. We did not
use purified antibody in this report, for the reason that purification by absorption
and elution (which are equilibrium processes) tend to concentrate antibodies of
low avidities in the eluates (Schram, et al. , 1971). The purified preparations,
thus, do not represent the composition of the original serum.

ACKNOWLEDGEMENTS

This investigation, carried out in the Killgore Research Center, West Texas
State University, was supported in part by grants from the Robert A. Welch Founda¬
tion and from the Committee on Organized Research at West Texas State University.

LITERATURE CITED

Farr, R. S., 1958- A quantitative immunochemical measure of the primary interaction be¬
tween 131I-BSA and its antibody./. Infect. Dis., 103:239.

Hodgson, H.H., and J. S.Wignall, 1926-Preparation of 3, 5-dihalogenophenols. / Chem. Soc.,
(London), 2:2077.

Karush, F., 1957 -The interaction of purified anti-beta-lactoside antibody with hapten. / Am.
Chem. Soc., 79:3380.

Lowry, O. H., N. J. Rosebrough, A. L. Farr, and R. J. Randall, 1953-Protein measurement
with the Folin reagent./. Biol. Chem., 193:265.

Oncley, J. L., 1949-Conference on the preservation of the cellular and protein components
of blood. American National Red Cross, Washington, D.C.

Paktinat, J., 1976-Avidity of chicken anti-lactoside antibody for synthetic lactosides. Masters
Thesis, West Texas State Univ., Canyon, Texas.

Pauling, L., D. Pressman, D. H. Campbell, C. Y. Ikeda, and M. Ikawa, 1942-The serological
properties of simple substances. I. Precipitation reactions between antibodies and sub¬
stances containing two or more haptenic groups. /. Am. Chem. Soc., 64:2994.

Schram, A. C., 1970-Heterogeneity of the antigenic centres of albumin toward chicken
antibodies. Immunology, 18:7.

- , S. J. Hwong, and C. P. Christenson, 1971 -Purification of chicken anti-bovine

serum albumin antibody. Immunology, 20:637.

Yariv, J., M. M. Rapport, and L. Graf, 1962 -The interaction of glycosides and saccharides
with antibody to the corresponding phenylazo glycosides. Biochem. J., 85:383.

CORROSION STUDIES IN A WATER FLOOD OIL FIELD SYSTEM

by LARRY G. SPEARS

Division of Science, University of
Houston, Downtown College,

Houston 77002

ABSTRACT

Corrosion studies of a water flood oil field system were undertaken. Experimental evidence
indicated that a CaCC>3 and Fe containing film was forming on the interior surfaces of the
mild steel pipe that handled production fluids. The injection water used for water flooding
contained an insufficient amount of Ca to allow formation of a protective CaCC>3 film and
was thus more corrosive than the production water.

INTRODUCTION

The use of water flooding for the recovery of oil, after primary production
techniques are exhausted, is now very common. Water flooding as a deliberately
planned operation for secondary oil recovery had its origin in the Bradford and
Allegany fields of Pennsylvania and New York, in 1920 (Muskat, 1949). This type
of secondary recovery operation involves the forcing of water, under pressure,
into the ground via a number of injection wells to force out any remaining oil
not removed by a primary recovery operation. Permeation of this injection water
through the surrounding formation results in the formation of an oil bank ahead
of the advancing water which is subsequently removed by a producing well.

In water flood operations, attention must be given to the character and treat¬
ment of the injection water in order to prevent corrosion and/or plugging of the
formation. Because of the large volumes of water used in this type of recovery,
the unit cost of treatment/barrel of water injected must be small. In many instances
the source of the injection water used is the local producing oil wells. This water
is usually very corrosive because of its high NaCl content and the presence of either
02 , H2S, or C02 , or a combination of these gases.

This study concerns a water flood oil recovery system of more than 100 pro¬
ducing shallow oil wells. The water flood operation is necessary to obtain the oil
from the sand formations. The injection water used is a mixture of water from
large water wells outside the oil field and water that has been separated from pro¬
duction fluids. For this discussion, injection water obtained from the external
water wells will be referred to as Well Injection (W.I.) water. Water from these
wells is transported via a common handling system to the oil field where it is
Accepted for publication: November 20, 1975.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

260

THE TEXAS JOURNAL OF SCIENCE

then pumped into the ground at various sites. Water that has been separated from
combined production fluids of all the producing wells will be referred to as Com¬
posite Production (C.P.) water. Four other water samples, taken from 4 different
sites in the oil field, were also studied. They will be referred to as Production Water
P-1, P-2, P-3, and P-4.

When this water flood operation was started approximately 10 years ago, the
C.P. water contained almost all natural occuring formation water. This was also
true for Production Water samples- taken at the same location samples P-1, P-2,
P-3, and P-4 were taken. Very little corrosion was observed in the production
fluid handling system. However, significant corrosion problems were observed in
the W.I. handling system. These problems were solved by the use of cement lined
tubing. During the past 10 years, the composition of the production fluids in
some portions of the field have become more like that of the W.I. water.

This study was initiated to determine what future corrosion problems might
occur in this field, as the composition of the formation water approaches that of
the W.I. water.

METHODS AND RESULTS

Corrosion rates for mild steel (No. 1018) in samples of W.I. , C.P., 50-50 mix¬
ture of W.I. and C.P., and P-1 , P-2, P-3, and P-4 water were made using direct
weight loss measurements and corrosion rate meter measurements. The steel
specimens used were cylindrical with a surface area of 8.74 cm2. All specimens
were sanded lightly with steel wool, polished with an emory cloth, washed in hot
benzene, rinsedi in acetone, and dried in a vacuum dessicator prior to use.

For direct weight loss measurements, each specimen was weighed using an
analytical balance, and then placed in the corrosion test solution for 24, 48, or
72 hrs. The test solution was closed to the atmosphere and stirred using a Sargent
magnetic stirrer set on a scale reading of 4. When a specimen was removed from
the test solution, any corrosion products remaining on the surface of the specimen
were removed using a hard gum eraser. The specimen was then washed in hot
benzene, rinsed in acetone, and dried in a dessicator. The steel specimen was then
weighed and the weight loss determined by comparison with the initial weight.
Using this weight loss data, the corrosion rates in mils/yr (mpy) were calculated
using the equation

3440W

mpy^“D^r

where W = weight loss in mg

D = density of the test specimen in g/cm3
A = area of the test specimen in cm2
T = time of exposure in hrs
1 mil ~ 0.001 in

CORROSION STUDIES

261

Corrosion rate meter measurements were made using a Petrolite Corrosion
Rate Meter, Model M-103. This instrument utilized 3 electrodes; a reference,
auxiliary, and test electrode. For this instrument, the reference and auxiliary elec¬
trodes have the same composition as the test electrode. The instrument electronics
are designed to utilize the equation developed by Stern and Geary (1957)

AE app = B a B c

Al 2 .3 ( I co rr) (Ba + Bc)

where Ba and Bc are the Tafel slopes of the anodic and cathodic reactions, respectively,
and

AEapp =10 mV for this instrument

AI = change in current as a result of AEapp

Icon = corrosion current.

The instrument automatically converts Icorr to mils/yr.

The corrosion rates obtained using these methods are listed in Table 1 .

TABLE 1

Corrosion Rates of Different Waters.

Water Type

Time

Weight Loss

Average

Corrosion
Rate Meter

Average

(hrs)

(mpy)

(mpy)

(mpy)

(mpy)

W.I.

24

36.3

34.0

48

37.8

36.3

35.0

33.3

72

34.7

31.0

C.P.

24

22.7

22.0

48

18.5

21.4

20.5

22.0

72

23.0

21.5

50-50 Mixture

24

14.9

of W.I.

48

14.7

14.3

and C.P.

72

13.3

Production

24

19.7

Water, P-1

48

16.0

18.0

72

18.4

Production

24

14.3

Water, P-2

48

13.4

13.9

72

14.1

Production

24

22.3

Water, P-3

48

24.3

24.8

72

27.7

Production

24

19.9

Water, P-4

48

16.8

17.5

72

15.7

262

THE TEXAS JOURNAL OF SCIENCE

All water samples were analyzed by a private testing laboratory. These results
are given in Table 2. It is well know that in water systems, Cl- and S04~4 enhance
corrosion while Ca+2, Mg+2, and HC03~ ions act as mild corrosion inhibitors. In
Table 2, it is easy to see that natural formation water is high in Cl", S04~2, Ca+2,
and Mg+2, and the HC03~ level in the W.I. water is slightly higher than that of the
natural formation water.

TABLE 2
Water Analysis
Well Injection Water

Total Dissolved Solids, 14,976 mg/1; Resistivity, 0.4233 £2-m at 78 F;
Specific Gravity, 1.0128 at 60 F; pH, 8.8 at 79 F

Constituents

meq/1

mg/1

Constituents

meq/1

mg/1

Sodium

242.11

5,566.0

Chloride

211.55

7,499.0

Calcium

1.92

38.5

Bicarbonate

11.23

685.0

Magnesium

3.06

37.2

Sulfate

22.82

1,096.0

Iron

0.16

4.5

Carbonate

1.65

49.5

Barium

0.00

0.0

Hydroxide

0.00

0.0

Composite Production Water

Total Dissolved Solids, 81,55 1 mg/1; Resistivity, 0.0885 12-m at 79 F;

Specific Gravity, 1.0689 at 60 F; pH, 7.55 at 79 F

Constituents

meq/1

mg/1

Constituents

meq/1

mg/1

Sodium

1,259.12

28,947.00

Chloride

1,300.99

46,118.0

Calcium

86.98

1,743.00

Bicarbonate

4.51

275.0

Magnesium

41.79

508.00

Sulfate

82.43

3.959.0

Iron

0.04

1.25

Carbonate

0.00

0.0

Barium

0.00

0.00

Hydroxide

0.00

0.0

Production Water, P-1

Total Dissolved Solids, 39,233 mg/1; Resistivity, 0.20000 H-m at 75 F;

Specific Gravity, 1.0248 at 60 F; pH, 7.35

Constituents

meq/1

mg/1

Constituents

meq/1

mg/1

Sodium and

Chloride

456.7

16,194.0

Potassium

492.9

11,337.0

Bicarbonate

10.2

620.0

Calcium

58.6

1,174.0

Sulfate

98.4

4,730.0,

Magnesium

13.8

168.0

Carbonate

--

—

Iron

Trace

Trace

Hydroxide

--

—

Barium

0.0

0.0

CORROSION STUDIES

263

Table 2 (Continued)

Production Water, P-2

Total Dissolved Solids, 125,962 mg/1; Resistivity, 0.062 £2-m at 75 F;
Specific Gravity, 1.0911 at 60 F; pH, 7.3

Constituents

meq/1

mg/1

Constituents

meq/1

mg/1

Sodium and

Chloride

2,052.6

72,786

Potassium

1,934.6

44,496.0

Bicarbonate

8.0

488

Calcium

117.1

2,347.0

Sulfate

86.9

4,180

Magnesium

95.8

1,165.0

Carbonate

—

—

Iron

Trace

Trace

Hydroxide

—

—

Barium

0.0

0.0

Production Water, P-3

Total Dissolved Solids, 57,215 mg/1; Resistivity, 0.125 £2-m at 75 F;
Specific Gravity, 1.0412 at 60 F; pH, 7.80

Constituents

meq/1

mg/1

Constituents

meq/1

mg/1

Sodium and

Chloride

859.3

30,473

Potassium

865.8

19,913.0

Bicarbonate

3.2

195

Calcium

80.9

1,622.0

Sulfate

100.2

4,818

Magnesium

16.0

194.0

Carbonate

—

—

Iron

Trace

Trace

Hydroxide

—

—

Barium

0.0

0.0

Production Water, P-4

Total Dissolved Solids, 131,548 mg/1; Resistivity, 0.061 12-m at 75 F;
Specific Gravity, 1.0934 at 60 F; pH, 6.70

Constituents meq/1 mg/1 Constituents meq/1

mg/1

Sodium and

Chloride

2,160.6

76,617

Potassium

2,068.4

47,573.0

Bicarbonate

7.0

429

Calcium

149.1

2,988.0

Sulfate

75.5

3,630

Magnesium

25.6

311.0

Carbonate

__

—

Iron

Trace

Trace

Hydroxide

—

—

Barium

0.0

0.0

Since the ion concentrations varied in the different waters, it was thought that
these various concentrations might have a noticeable effect on the corrosion rate.
Two hundred ml of C.P. water was evaporated and the remaining salts added to
200 ml of W.I. water. The corrosion rate of mild steel in the W.I. water decreased
from 36 mpy to 18 mpy after addition of these salts. It was then decided to vary
the different ion concentrations selectively and to measure the resulting corrosion
rates.

264

THE TEXAS JOURNAL OF SCIENCE

As shown in Table 1 , P-2 was the least corrosive water. Ca+2, Mg+2, and Cl” ions
were added to 3 samples of W.I. water to make the respective ion concentration
equal to that of P-2 , and the corrosion rates of mild steel in these samples measured.
Table 3 gives the results of these measurements. The Ca+2 ions and Cl” ions caused
a major decrease in the corrosion rate. No HC03” ions were added to the W.I.
water since it has a high HC03” concentration. So by varying the concentrations
of the Ca+2 and Cl” ions, the corrosive properties of the W.I. water could be made
similar to that of the production waters.

TABLE 3

Corrosion Rates of Well Injection Water with Ions Added.

W.I. water with CaCl2 added:

Time

Corrosion Rate

Average

(hrs)

(mpy)

(mpy)

24

7.99

48

7.58

7.31

72

6.35

W.I. water with NaCl added:

Time

Corrosion Rate

Average

(hrs)

(mpy)

(mpy)

24

15.8

48

21.0

18.7

72

19.3

W.I. water with Mg Cl 2 added:

Time

Corrosion Rate

Average

(hrs)

(mpy)

(mpy)

24

36.0

48

38.2

35.9

72

33.7

Calcium carbonate is well known for its ability to act as a cathodic corrosion
inhibitor (Bregman, 1963). CaC03 deposits on a metal result in an ennoblement
of the metal for the cathodic reaction (thus it is called a cathodic inhibitor). Anodic
and cathodic potentiostatic polarization curves for mild steel in samples of W.I. ,
C.P. and 50-50 mixtures of W.I.-C.P. water were made. An Elron potentiostat
(Model CHP-1) was used as a constant potential source and aKeithley electrometer
(Model 602) was used to monitor the potential change versus a saturated calomel
electrode. An ammeter, built into the potentiostat, was used to record the current.
The electrolytic solution was at room temperature and stirred using a Sargent
magnetic stirrer set at 5. Current readings were recorded at each potential after
they had maintained a constant value for 5 min. The polarization curves obtained
are shown in Figure 1 . The cathodic portions of the C.P. and 50-50 mixture in¬
dicate an ennoblement of the steel, with the 50-50 mixture producing the greatest
effect.

CORROSION STUDIES

265

Film analyses were made on mild steel specimens exposed to W.I. water with
CaCl2 added to it. The film was dissolved from the specimen by immersing it in
10% HC1. The solution was then analyzed for Ca+2 using a Beckman, Model 979,
atomic absorption system. Assuming that all the Ca+2 was due to CaC03, the film
contained approximately 85% by weight CaC03. When the specimen was placed
in the HC1 solution, vigorous bubbling accompanied the decomposition of the
film (probably C02).

As mentioned above, CaC03 is an effective corrosion inhibitor in some systems.
A thin, adherent, protective film of CaC03 is effective in some types of water
treatment. The equilibrium between HC03" and C03~2 ions is represented by the
following:

266

THE TEXAS JOURNAL OF SCIENCE

2HC03" t C03”2 +C02 + H20.

A possible mechanism for the corrosion observed in this particular study is as follows:

at the anodic sites: Fe -> Fe+2 + 2e“

Fe+2 + 20HT (from cathodic sites) J Fe(OH)2

at the cathodic sites:

in the absence of 02 : 2H2 0 + 2e“ -* 20H” + H2

in the presence of 02 : Vi02 + H20 + 2e” -> 20FT
Ca+2 + 20H" % Ca(OH)2
Ca(OH)j + HC03' t CaC03 + H20 + OPT
or

HCCV + OH” t H20 + C03"2
Ca+2 + C03"2 t CaC03 .

Evans (1960) points out that a protective film of CaC03 is only formed from
bicarbonate waters if the C02 content is limited to the amount needed to stabilize
the Ca(HC03)2 that is present. In this manner, the smallest increase in pH suffices
to render the liquid next to the metal supersaturated with CaC03 .

A saturation point of HC03” and Ca+2 must be maintained for cathodic in¬
hibition to occur. Evans (1960) and Bregman (1963) have stated that formation
of a CaC03 film will occur even if the bulk solution is not at its saturation point.
This is because the Ca+2 ions will migrate to the negative cathodic sites, where
the pH is higher than that of the bulk solution due to formation of OH” ions.
This results in formation of a protective CaC03 film.

The control of pH is an essential part of the water treatment. A pH of 6.0 to
6.5 is the minimum value that must be maintained, since below this value corrosion
proceeds rapidly. Also a maximum value of 7.5 to 8.0 must not be exceeded,
since at higher pHs scaling will occur. In practice, for better corrosion inhibition
the pH is kept as high as possible without having scaling occur. Langeleir (NACE,
1971) has devised an index which is the difference between the actual pH value
of water and the pH value reached when water is brought into equilibrium with
CaC03 . It indicates when the possibility of a CaC03 protective film will be formed .
Finally, other suspended solids or deposits present in the system can act as a seeding
material for the precipitation of the CaC03 .

A graph was constructed of the corrosion rate of mild steel versus the product
of the Ca+2 ion and the square of the HC03” ion concentrations in meq/1. This
graph is shown in Figure 2. An important trend can be seen in this graph. As the
product of the ion concentration increases, the corrosion rate decreases. W.I. water
has the greatest corrosion rate while having the smallest concentration product
value. The production waters containing the most original formation water con¬
tains the highest Ca+2 ion concentration. W.I. water has only small amounts of

CORROSION STUDIES

267

[Ca++] [HC03 ]2 mEQ3/l3
Figure 2. Effect of Ca+2 and HCO3 ions on corrosion rate.

Ca+2 but is rich in HC03” ions. Thus, by adding Ca+2 to W.I. water, its corrosion
rate should decrease significantly. The HC03” concentration of W.I. is very high
and thus the equilibrium between HC03~ and C03“2, described above, is shifted
to the right when Ca+2 ions are added to the system. This favors the formation
of CaC03 .

As shown in Table 3, the NaCl that was added to the W.I. water caused the
corrosion rate to decrease. Foley (1970) describes the role 02 and Cl” ions play
in corrosion processes. He states that 02 acts as a cathode depolarizer, causing
Fe+2 to go into solution, oxidizes the dissolved Fe(OH)2 to Fe(OH)3, and restores
an oxide film. In the absence of 02 , saturated NaCl solutions are essentially non-
corrosive to Type 1020 steel. In many studies it has been amply shown that in

268

THE TEXAS JOURNAL OF SCIENCE

neutral solutions and at temperatures up to about 80 C, the 02 effect is important.
A good discussion on the effects of 02 on metal dissolution rates is given by Bockris
and Reddy (1970). The fact that concentration effect and 02 availability are inter¬
related has been observed by many investigators. As Cl" ion concentration increases,
the solubility of 02 decreases, thus decreasing the corrosion rate. Therefore, the
reduction in corrosion rate when NaCl was added to W.I. water was probably
due to the decreased solubility of 02 (even though it may be present in low con¬
centrations) in the water.

Alexander and Foley (1975) present a good discussion on the anion (S04~2, Cl",
Br", and I" ) dependence of the activation energy for Fe corrosion. The mild steel
used in this study was approximately 99% Fe.

Also, from Table 3 it can be seen that the addition of Mg+2 ions had little effect
on the corrosion rate. This is because MgC03 is much more soluble in water than
CaC03. Therefore, very little protective MgC03 would be available to form a pro¬
tective film on Fe .

When discussing ion concentrations present in the waters, it is not completely
advantageous to look at one ion concentration by itself. The corrosive properties
of waters are due to a composite of all ions and dissolved gases present in them.
There exists no defined points where one ion concentration may override the effect
of another ion present. This sometimes leads to a complicated system which is
often difficult to explain. The presence or absence of one ion may or may not
enhance the inhibition properties of the other. This is also true for the gases that
may or may not be present. Also, the degree of effectiveness of the ions as in¬
hibitors must be taken into account. Therefore, we must always keep in mind
the total picture when discussing these corrosive waters.

CONCLUSIONS

Corrosion studies of a water flood oil recovery system were undertaken. Past
field experiences had indicated that portions of the oil field handling mainly W.I.
water were experiencing corrosion problems. Mild steel in contact with production
waters did not undergo serious corrosion.

Our data indicates that a thin, protective CaC03 film is being formed on the
steel that handles production fluids. In the W.I. system, a low Ca+2 concentration
prevents formation of this film. As the ratio between W.I. water and the original
formation water increases in the future, the ratio of the concentration of Ca+2 and
HC03“ ions will decrease. This will cause a decrease in the amount of shift of the
HC03-C03"2 equilibrium to the right, thus decreasing the amount of CaC03
formed at the metal surface. Thus while the current production waters cause rel¬
atively low corrosion rates on mild steel, this will change as more W.I. water enters
the formation.

CORROSION STUDIES

269

LITERATURE CITED

Alexander, B. J., and R.T. Foley, 1975 -Anion dependence of the activation energy for iron
corrosion. Corrosion, 31:148.

Bockris, J. O’M., and A.K.N.Reddy, 1970 -Modem Electrochemistry, Vol. 2. Plenum Press,
New York, N.Y., pp. 1296-1308,

Bregman, J. I., 1963 -Corrosion Inhibitors. Macmillan Co., New York, N.Y., pp. 94.

Evans, U. R., 1960-77ze Corrosion and Oxidation of Metals. Edward Arnold Publishers, Ltd.,
London, pp. 161-162.

Foley, R. T., 1970-Role of the chloride ion in iron corrosion. Corrosion, 26:58.

Muskat, M., 1949 -Physical Principles of Oil Production. McGraw Hill, New York, N.Y., pp. 732.

NACE, 191 1— Cooling Water Treatment Manual TPC Publication No. 1, Nat. Assoc, of Cor¬
rosion Engineers, Houston, pp. 7.

Stearn, M., and A. L. Geary, 195 7 -Electrochemical polarization, I. A theoretical analysis
of the shape of polarization curves./. Electrochem . Soc., 104:56.

THE SALINITY AND TEMPERATURE TOLERANCE AND THE
GROWTH OF MACROBRACHIUM OHIONE (SMITH) 1874 REARED
IN LABORATORY TANKS1

by KYUNG-SUK CHUNG

Texas A&M University ; Department of Wildlife and

Fisheries Sciences , College Station 77843

ABSTRACT

The salinity and temperature tolerances and the growth of Macrobrachium ohione was
observed in laboratory tanks at the National Marine Fisheries Service, Galveston, Texas.

Salinities over 13 ppt and temperatures over 31.5 C increased mortality, especially during
molting periods.

In an effort to determine maximum size of M. ohione the relationship between initial
body length (lt) and terminal body length (Lt+T) was plotted and the Walford line was found
Lt+T “ 96.79431 (1-e-2,40192) + e”2‘40192lt. The calculated maximum body length, carapace
length, and body weight were 96.79 mm, 28.23 mm, and 11.47 g, respectively.

INTRODUCTION

The continuing population growth of the world has initiated a strong demand
to increase agricultural as well as aquacultural products and to examine the eco¬
nomic feasibility of culturing many invertebrate fishery resources. River shrimp
{Macrobrachium) is one of the prospective groups of protein foods which is re¬
ceiving much attention and many researchers are studying intensive culture prob¬
lems. Salinity , temperature, and growth on the Malaysian giant shrimp, M. rosen-
bergii, as it relates to the aquaculture and larval development have been reported
by Fujimura (1966, 1970), Lewis (1962, 1965,1966), Ling (1962,1967a, 1967b,
1971), Ling and Merican (1961), Uno and Kwon (1969), and Wickins (1969, 1972).
Larval development of M. acanthurus and M. carcinus has been studied by Choudhury
(1970, 1971a, 1971b, 1971c). Ingle and Edward (1960) have noted the possibility
of artificially culturing them. Ecology, larval development, and aquaculture prob¬
lems relating to salinity, temperature, and breeding cycle of M. nipponense were
reported by Chung (1972), Kwon and Uno (1969), and Uno (1971), respectively.

Four species of Macrobrachium (M. ohione, M. olfersi, M, acanthurus, and M.
carcinus) in Texas are reported by Hedgpeth (1947, 1949) and Reimer and Trudeau
(1975). Macrobrachium ohione is the most common and smallest of the 4 species.

1 A contribution of the Department of Wildlife and Fisheries Sciences, Texas Agricultural
Experiment Station, Texas A&M University.

Accepted for publication: January 20, 1975.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

272

THE TEXAS JOURNAL OF SCIENCE

It is found along the coastal line and inland waters of the United States from
Virginia (Hobbs and Massmann, 1952) to Central Texas (Holthuis, 1952).

Dugan (1971 , 1972) reported on salinity tolerances in the larval stages of this
species and Reimer ,etaL, (1974) have discussed the general ecology of M. ohione
collected from Galveston Bay, in natural environments. Reimer, et al, (1974)
suggest that juvenile and adult M. ohione live in the freshwater and avoid salinities
above 15 ppt. Temperatures near 30 C represent their upper tolerance level.

This work was done in 2 parts. The 1st experiment was for evaluating salinity
and temperature effects on survival and ran for 30 days. The 2nd experiment was
concerned with growth in freshwater and ran for 30 days.

MATERIALS AND METHODS

M. ohione were collected from the San Bernard River, Brazoria County, Texas.

The shrimp were held in aged tap water until the beginning of each experiment.
Forty-eight shrimp of only 1 species were transported in an ice cooler (60x35 x35 cm)
with aeration on 22 June 1973 for temperature and salinity experiments. An addi¬
tional 30 shrimp were transported, as in the above method, on 15 July 1973 for
growth experiments. All groups of shrimp were fed an experimental hard pellet-
type food of Louisiana State University, about 5% of the total body weight of the
group every day. A distinction between sex was not made.

Salinity - Temperature

Shrimp changed from fresh to salt water were first allowed to acclimate for
6 hrs in 2 ppt salinity. Each group of shrimp to be held at higher salinities was
allowed to acclimate for 6 hrs at each of the test salinities (4, 6, 10, 13 ppt) until
they reached the specific salinity they were to be held at. Salinity levels are con¬
tinuously fluctuating in estuaries with diurnal tides. The median of 6 hrs-salinity-
acclimation was arbituary chosen. Due to the variable chemical characteristics of
natural seawater, synthetic sea salt, Instant Ocean, was used in the experiments.
This type of solution was found adequate for penaeid shrimp culture by Cook and
Murphy (1969) and Mock and Murphy (1971). Salinities were regulated by use
of an American Optical Goldberg T/C refrectometer. The salt solution was aged
for at least 24 hrs in a 500 gal (1 .89 m3) tank before the experiment began. Eight
shrimp were kept in each of 6 experimental tanks, 80 x 50 x 45 cm in dimension
with each tank containing approximately 150 1 of water. A biological filter using
oyster shells was connected with each tank so that the water was continually cir¬
culated. In addition, water was circulated in each tank by air dispensed by a Mil¬
lion Aire air stone (Figure 1). Dissolved oxygen levels were not measured but were
assumed to be adequate.

The experimental tanks were located in a green house that was roofed with
gray corrugated fiber glass sheeting. Each tank was covered with fiber glass sheeting
to prevent the shrimp from jumping out of the tank during the experiment.

MA CROBRA CHIUM OHIONE

273

Figure L Experimental tank with biological filteration and air circulation (A, air tube; B,
shell tube; C, MillionAire air stone, c, air stone; D, PVC pipe; E, lead combiner).

Water was not changed during experiments, but distilled water was added every
day to correct salinity changes as a result of evaporation.

Temperature was measured with a Fisher’s 15 -043 A thermometer twice daily
at 8 am and 5 pm. The temperature in each tank was allowed to fluctuate with
atmospheric temperature.

274

THE TEXAS JOURNAL OF SCIENCE

Growth

Since good survival was found to occur in freshwater, this medium was used
for the growth study. The tank and filter system used in the first experiment were
also used here . Temperature was allowed to fluctuate with atmospheric conditions.

Shrimp were measured according to total body length (tip of rostrum to distal
end of telson), carapace length (posterior margin of orbit to the dorsal posterior
margin of carapace) and body weight. Measurements were to the nearest 0.1 mm
for body length , 0 .0 1 mm for carapace length , and were made with caliper . Length
measurements were taken before and after experiments. For weighing the shrimp,
individuals were removed from the tank, carefully wiped with paper towel to
remove excess water, and placed on the balance.

The body length obtained after 30 days was plotted against that of the initial
length to find maximum length to be expected in ponds. A Walford (1946) type
line was used for this determination. The instantaneous growth rate using body
length was determined based on the procedures by Gulland (1969).

Estimate of daily survival rate was calculated by S = Nt/N0 (Ricker, 1975)
assuming constant survival rate during experimental days where S is survival rate,
N0 is initial population size, and Nt is population size time t. Instantaneous mor¬
tality rate (Z) was calculated by e_zt=Nt/N0 assuming mortality rate is constant
during experimental days.

RESULTS

Salinity - Temperature

The % survival in each experimental tank after 30 days was as follows: 87.5%,
in aged tap water; 84.1% in 2 ppt; 82.5% in4ppt; 80.8% in 6 ppt; 72.8% in lOppt
and 52.0% in 13 ppt. An analysis of variance was run to determine if these sur¬
vival rates were significantly different (Table 1).

TABLE 1

Analysis of variance of survival rate of test animals in 6
salinities and Tukey’s honestly significant difference test
based on residual (mean not spanned by the line is sig¬
nificantly different).

ANOVA

Source

d.f.

SS

MS

F

Salinities

5

25,465

.63

5

,093.13

12.75**

Survival rate

174

69,497.

.77

399.41

in salinities

Total

179

94,693

.40

Tukey’s Test

Salinity (ppt)

0.0

2.0

4.0

6.0

9.0

13.0

Mean survival (%)

87.5

84.1

82.5

80.8

72.8

52.0

** Significance at 1%

MACROBRACHIUM OHIONE

275

The total survival rate of the 6 experimental groups was significantly different
at 1% level. To determine which group or groups were significantly different, a
Tukey’s honestly significant difference test was run. The survival rate of those
shrimp in 13 ppt salinity was significantly different from that of other salinities
at the 1% level (Table 1). The relationship between salinity (s) and survival rate (S)
was calculated with S = 94.14559s" 1515 (Figure 2).

Figure 2. Relationship between salinity (s) and survival rate (S) of M. ohione.

Most deaths occurred during ecdysis at temperatures of 31.5 C and greater.
The relationship between temperature (T) and % of mortality (Z) was calculated
with Z= (0.53604 x 10~34)T23*7949. Afternoon temperatures ranged from 28.5 C
to 32.9 C and morning temperatures from 25.7 C to 28.7 C (Figure 3). All shrimp
deaths were reported in the afternoon and when temperatures were above 31.5 C.
In every case where death was recorded some shrimp were observed to be molting.

temperature

276

THE TEXAS JOURNAL OF SCIENCE

MACROBRACHIUM OHIONE

277

Growth

The relationship between initial (lt) and terminal (Lt+T) body length was plotted
using the Walford type formula (Walford, 1946), Lt+T = 96.79431 (l-e~2*40192) +
e -2.40192 ^ (FigUre 4). Calculated maximum carapace length was 28.23 mm.

Figure 4. Relationship between M. ohione body length before (lt) and after experiments

(Lt+T)-

Relationship between carapace length (C) and body length (L) was presented
with L= 10.49223 + 3.05742 C (Figure 5).

Relationship between body length (L) and body weight (W) was calculated to
be W = (0.18625 x 10-6)L3*9225 (Figure 6).

DISCUSSION
Salinity - Temperature

Laboratory experiments showed that differences in survival rates of M. ohione
in freshwater and at salinities of 2, 4, 6 or lOppt were not significant at 1% level.
However, the mortality of shrimp at 13 ppt was significantly different from fresh¬
water as well as the other salinities (Table 1). Hobbs and Massmann (1952) have

278 THE TEXAS JOURNAL OF SCIENCE

CARAPACE LENGTH in mm

Figure 5 . Relationship between carapace length (C) and body length (L) of M. ohione.

reported seasonal movement of M. ohione with maximum abundances in 5 ppt
salinity. Gunter (1937) found M. ohione in a salinity range of 1.38 - 14.24 ppt
in bay waters. Viosca( 1957) noted that M ohione also inhabited freshwater lakes

MA CROBRA CHIUM OH ION E

279

X

o

LU

5

>-

D

o

co

BODY LENGTH in mm

Figure 6. Relationship between body length (L) and body weight (W) of M. ohione.

in the floodplains down to the edge of salt water in Louisiana. Dugan (197 1 , 1972)
discussed its salinity tolerances in the laboratory and indicated that gravid females
would spawn and the larvae hatch in freshwater. The larvae needed brackish water.
Reimer, et al, (1974) have suggested that salinity near 15 ppt acted as a barrier
to movement of shrimp into Galveston Bay.

Hughes, et al, (1973) reported that current chamber experiments indicated
gravid M. acanthurus tend to swim consistently downstream whereas nongravid
females tend to swim upstream. This would seem to substantiate the general feeling
that the gravid female is stimulated to return to brackish water for the release of

280

THE TEXAS JOURNAL OF SCIENCE

her spawn. This probably occurs in M. ohione and accounts for the large number
taken in bay systems (Gunter, 1937; Reimer, et al, 1974). It would seem most
likely that M. ohione spawn and the eggs hatch in the mouth of the rivers or in
the upper estuary where the salinity is lower than 15 ppt. The larvae move to the
bay area where salinities are higher (between 10 - 20 ppt), for the larvae need more
than 15 ppt salinity during larval development periods (Dugan, 1972). After meta¬
morphosis into the post-larval stage, they presumably migrate contranatant to the
river where salinities are low (less than 15 ppt). Growth continues in freshwater
until the next spawning when they again migrate to the estuaries.

High mortality was associated with high water temperatures occurring during
molting of the shrimp . The mortality during molting occurred when late afternoon
temperatures reached 31 .5 C and higher. Experimental temperatures reached 32 C
for several days and non-molting animals were able to survive. None survived at a
peak temperature of 32.9 C after molting. In general, temperature and dissolved
oxygen would be highly interacted in molting and mortality of crustacean, however,
a biological filter system and a Million Aire air stone (Figure 1) presumably pro¬
vided adequate dissolved oxygen in this experiment. Temperature was considered
to be the primary factor affecting molting and mortality.

Reimer, et al, (1974) have suggested that temperature near 30 C represented
the upper limit of tolerance of M. ohione in the Galveston Bay area. My data showed
that 31.5 C was the upper tolerance limit of M. ohione in laboratory tanks. It
would appear that temperature regulates the distribution of this species by inter¬
fering with the actual molting process. Since Macrobrachium must go through a
prespawning molt, temperature control is critical.

M. ohione lives in the intake canal of the P. H. Robinson Generating Station,
Bacliff, Texas, during periods of low salinity. Temperatures of intake water during
June and July of 1974 ranged from a daily minimum of 24.4 to 30.0 C and a
maximum of 26.6 to 32.2 C. These temperatures are similar to those recorded in
my tanks (25.7 to 28.7 C in the morning and 28.5 to 32.9 C in the afternoon)
and they indicate that M. ohione needs to seek cool area to survive during the
hot part of the summer.

The relationship between temperature (T) and the % mortality (based on number
of shrimp in all 6 tanks after molting) is presented graphically in Figure 7. It shows
that temperature is directly correlated with mortality. It would seem that tem¬
peratures for juvenile and adult M. ohione, in ponds, will need to be under 30 to
31 C.

Growth

The relationship between initial (lt) and terminal body length (Lt+T) was plotted
and a Walford line was found Lt+T = 96.79431(l-e~2,40192) + e"2,40192lt. Calcu¬
lated maximum body length was 96.79 mm.

The calculated maximum body length, 96.79 mm, compared favorably with
most reports (Table 2) from the literature. It would seem, therefore, under arti¬
ficial conditions, one might expect the size of adult populations to be similar to
natural populations.

MACROBRACHIUM OHIONE

281

Figure 7. Relationship between temperature (T) and percent mortality (Z) of M. ohione.

Relationship between carapace length (C) and body length (L) was plotted
and showed a linear equation, L = 10.49223 +2.05742 C. Calculated maximum
carapace length substituting from this formula, was 28.23 mm. The body length
(L) and body weight (W) relationship was plotted and formulated with W =
(0.18625 x 10“6)L3*9225 . Calculated maximum body weight was 1 1.47 g.

282

THE TEXAS JOURNAL OF SCIENCE

TABLE 2

Maximum Lengths of M. ohione Reported From the Literature.

Author

Length

Sex

Schmitt (1933)

103 mm

F

McCormick (1934)

96 mm

?

Gunter (1937)

93 mm

?

Hedgpeth (1959)

100 mm

F

Dugan (1971)

100 mm

F

Reimer, et al. , ( 1 974)

85 mm

F

ACKNOWLEDGEMENTS

Dr. Rollin D. Reimer, Department of Wildlife and Fisheries Sciences, Texas
A&M University, assisted in this work and Mr. Cornelius R. Mock and Mr. Billy R.
Salser, Fishery Biologist, National Marine Fisheries Service, Galveston, provided
research facilities and cooperated in performing laboratory experiments. Mr. Gary
Jones, Texas A&M University, Agricultural Experiment Station, Angleton, Texas,
provided shrimp and Dr. Kirk Strawn, Department of Wildlife and Fisheries Sciences,
Texas A&M University, aided with the manuscript.

LITERATURE CITED

Choudhury, R C., 1970 -Complete larval development of the palaemonid shrimp, Macrobrach¬
ium acanthurus (Wiegmann, 1836), reared in the laborabory. Cruataceana, 18(2): 113.

- , 1971a— Complete larval development of the palaemonid shrimp, Ma cro brachium

carcinus (L.) reared in the laboratory (Decapoda, Palaemonidae). Crustaceana, 20(1):51.

- , 1971b-Responses of larvae of Macrobrachium carcinus (L.) to variations in sal¬
inity and diet (Decapoda, Palaemonidae). Crustaceana, 20(2): 113.

— - -, 1971c-Laboratory rearing larvae of the palaemonid shrimp , Macrobrachium acan¬

thurus (Wiegmann, 1836). Crustaceana, 21(2): 113.

Chung, K. S., 1972— Biological studies on the freshwater shrimps in Korea 4. The ecology of
Macrobrachium nipponensis (De Haan). Bull. Korea Fish. Soc., 5(3):83.

Cook, H. L., and M. A. Murphy, 1969-The culture of larval penaeid shrimp. Trans. Amer.
Fish. Soc., 98(4):75 1.

Dugan, C. C., 1971 -Floridan research: Freshwater shrimp. Amer. Fish. Farm., 2( 1 0) : 8 .

- , 1972— Culture of brackish-freshwater shrimp, Macrobrachium acanthurus, M. car¬
cinus, and AT ohione. Proc. World Mariculture Soc., 3:185.

Fujimura, T., 1966-Notes on the development of a practical mass culturing technique of
the giant prawn Macrobrachium rosenbergii. FAO Rep. Conseil Indo-Pacific. 12 Sess.
IPFC/WP, 47:1.

MA CR OBRA CHIUM OH ION E

283

- , and H. Okamoto, 1970-Notes on progress made in developing a mass culturing

technique for Macrobrachium rosenbergii in Hawaii. IPFC/C70/Sym., 53:1.

Gulland, J. A., 1969-Part 1. Fish population analysis. Manual of Methods for Fish Stock
Assessment. FAO Manuals Fish. Sci Vol. 4, 154 p.

Gunter, G., 19 37 -Observations on the river shrimp , Macrobrachium ohione (Smith). Amer.
Midi. Nat., 18:1038.

Hedgpeth, J. W., 1947-River shrimps. Prog-. Fish Cult., 10:181.

- , 1949-The North American species of Macrobrachium. Tex. J. Sci., 1 (3) :28 .

Hobbs, H. H., and W. H. Massmann, 195 2 -The river shrimp , Macrobrachium ohione (Smith)
in Virginia. Virginia J. Sci., 3:206.

Holthuis, L. B., 195 2-A general revision of the Palaemonidea (Crustacea, Decapoda, Natantia)
of the Americas. 2. The sub-family Palaemoninae. Occ. Pap. Allan Hancock Found., 12: 396.

Hughes, D. A., and J. D. Richard, 1973-Some current-directed movements of Macrobrachium
acanthurus (Wiegmann, 1836) (Decapoda, Palaemonidae) under laboratory conditions.
Ecol, 54(4):927.

Ingle, R. M., and B. Edward, 1960-Notes on the artificial cultivation of freshwater shrimp.
West Indies Fish. Bull., 4:1.

Kwon, C. S., and Y. Uno, 1969 -The larval development of Macrobrachium nipponense (De
Hann) reared in the laboratory. La mer., 7(4):30.

Lewis, J. B., 1962 -Preliminary experiments on the rearing of the freshwater prawn Macro¬
brachium carcinus (L ).Proc. Gulf Caribb. Fish. Inst., 14:1.

- , and J. Ward, 1965 -Developmental stages of the Palaemonid shrimp Macrobrach¬
ium carcinus (Linnaeus, 1758). Crustaceana, 9(2): 137.

- , - , and A. Mclver, 1966-The breeding cycle, growth and food of the

freshwater shrimp Macrobrachium carcinus (Linnaeus). Crustaceana, 10(1):48.

Ling, S. W., 1962— Studies on the rearing of larvae and juveniles and culturing adults of Macro¬
brachium rosenbergii (de Man). FAO Curr. Affair Bull. Mp., 35:1.

- , 1967a-Methods of rearing and culturing Macrobrachium rosenbergii (de Man).

FAO Rep., 57(3):607.

- , 1967b-The general biology and development of Macrobrachium rosenbergii (de

Man ). FAO Rep., 57(3):589.

- , 1971 -Some brief notes on the status and problems of shrimp and prawn farming

development in Asia. 48th Meeting, IPFC/EXCO, 48(6): 1 .

- , and A. B. O. Merican, 1961 -Notes on the life and habits of the adults and larval

stages of Macrobrachium rosenbergii (de Man ). IPFC Proc., 9(1 1) :5 5 .

McCormick, R. N., 1934 -Macrobrachium ohionis, the large freshwater shrimp. Proc. Ind.
Aca. Sci., 43:218.

Mock, C. R., and M. A. Murphy, 1971 -Techniques for raising penaeid shrimp. Proc. 1st Ann.
Workshop World Mariculture Soc., 143.

284

THE TEXAS JOURNAL OF SCIENCE

Reimer, R. D., K. Strawn,and A. Dixon, 1974-Notes on the river shrimp, Macrobrachium
chime (Smith, 1874) in the Galveston Bay system of Texas. Trans. Amer. Fish. Soc., 103(1):120.

- , and R. Trudeau, 1975 -Range Extension of Macrobrachium olfersi (Wiegmann

1836) into Texas. Tex. J. Sci., 26(3&4):620.

Ricker, W. E., 1975 -Computation and interpretation of biological statistics of fish popula¬
tions./. Fish. Res. Bd. Can. Bull., 191:382.

Schmitt, W. L., 1933-Notes on shrimps of the genus Macrobrachium found in the United
States./. Wash. Acad. Sci., 23:312.

Uno, Y., 1971 -Studies on the aquaculture of Macrobrachium nipponense (De Haan) with
special reference to breeding cycle, larval development and feeding ecology. Lamer., 9(2):123.

— - - — , and C. S. Kwon, 1969-Larval development of Macrobrachium rosenbergii (de

Man) reared in the laboratory./. Tokyo Univ. Fish., 55:179.

Viosca, P., 1957-Shrimp potpourri. Toms. Cons., 9(7) :20.

Walford, L. A., 1946-A new graphic method of describing the growth of animals. Biol. Bull.,
90(2):141.

Wickins, J. F., 1969— Preliminary experiments in the culture of the prawn Pandalus platy ceros
(Brandt) and the giant prawn ( Macrobrachium rosenbergii ) (de Man). ICES, C.M.E:11,
8 p. (mimeo).

- , 197 2 -Experiments on the culture of spot prawn, Pandalus platy ceros (Brandt)

and giant freshwater prawn, Macrobrachium rosenbergii (de Man). Fish. Invest. London.
Ser. 2, 27(5): 1023.

THE ECOLOGY OF A BRANCHIOBDELLID (ANNELIDA : OLIGO-
CHAETA) FROM A TEXAS POND

by STEPHEN J. KOEPP1 and EDGAR A. SCHLUETER

Department of Biological Sciences, North Texas State University,
Denton 76201

ABSTRACT

Seasonal changes in population structure of the branchiobdellid Cambarincola vitrea in¬
habiting crayfishes of a Texas farm pond were investigated. Worm population density and
reproduction were correlated with fluctuations in the physical habitat, the individual host
size class, and the apparent host molt condition. Mean worm load and cocoon deposition
reflected several meteorological disturbances of the pond during the period of study (J une
1972 to May 1973). Regardless of month of collection, molting hosts showed significantly
reduced branchiobdellid populations compared to intermolt crayfishes. A curvi-linear rela¬
tionship exists between increasing host size and worm load in favor of larger animals, indicating
the high molt frequency among younger crayfishes. Cocoon deposition fluctuated in a pattern
similar to that of the adult worms. The branchial cavities and pleopods appeared to be pre¬
ferred sites for cocoon deposition.

INTRODUCTION

Branchiobdellids are annelid ecto-inhabitants of the gills and exoskeleton of
freshwater crustaceans, chiefly crayfishes. Although quite common in freshwater
environs of North America, knowledge of these oligochaetes is limited. In particular
the extent of interaction between host and annelid remains obscure. While several
branchiobdellid species have been reported to be truly parasitic (Holt, 1963), the
majority are considered commensals. Young (1966) has demonstrated that a viable
host is a necessary condition for branchiobdellid reproduction.

Previous investigations have been primarily limited to taxonomic considerations.
Goodnight (1940) revised the family, while reviewing the scattered early literature.
Holt (1954, 1955, 1960, 1963) and Hoffman (1963) have added new species de¬
scriptions. Although several ecological reports have been published (Berry and
Holt, 1959; Penn, 1959; McMannus, 1960; Young, 1966; Bishop, 1968), specific
information is lacking concerning branchiobdellid activities within a habitat prone
to seasonal meteorological extremes. Such habitats are common to Texas in the
form of small farm ponds. The branchiobdellid Cambarincola vitrea Ellis is fre¬
quently found inhabiting crayfishes of this region, including the pond-dwelling
Procambarus simulans Faxon.

1 Present address: Biology Department, Montclair State College, Upper Montclair, New J ersey
07043.

Accepted for publication: October 8, 1975.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

286

THE TEXAS JOURNAL OF SCIENCE

It was the purpose of this investigation to study seasonal changes in branchiob-
dellid population structure within a Texas pond. Branchiobdellid response was
assessed on the basis of worm load and reproduction correlated with host factors
and fluctuations of the physical environment.

METHODS AND MATERIALS

The study site was located 3.5 mi north of Denton, Texas, on public land ad¬
jacent to Interstate Highway 35. The pond is a man-made depression with a silty
black mud substrate. Water depth varied from 2.2 m in the spring to complete
dryness in the late summer and early fall.

A prominent member of the aquatic fauna is the crayfish Procambarus simulans
(family Astacidae, subfamily Cambarinae), which is among the most common of
the Texas crayfishes. The only branchiobdellid foynd within the study site was
Cambarincola vitrea. Keys prepared by Hoffman (1963) and Goodnight (1940)
aided in the identification.

Crayfishes were collected with a hand net and immediately placed into individ¬
ual bottles containing a saturated solution of chlorobutanol. Monthly collections
were carried out (except September when the pond was dry). At each sampling,
50 crayfishes of various size classes were treated for removal of resident branchiob-
dellids, and the worm load determined. All specimens were subsequently fixed
and stored in 70% isopropyl alcohol.

Crayfishes were also individually analyzed for carapace length (in cm) and
apparent molt condition. Determination of molt condition was restricted to molt
and intermolt categories. The former included postmolt animals. Criteria included:
(1) evidence of pronounced imbibition; (2) presence of encrusting algae and/or
deposited insect ova on the exoskeleton; and (3) extent of gastrolith development.

Branchiobdellid cocoon deposition and distribution on the host body were
also investigated. Accordingly, all animals were first screened for the presence of
cocoons. Crayfishes were then grouped into 6 size classes (0-1 .9, 2. 0-2 .9, 3.0-3 .9,
4.0-4.9, 5.0-5.9, and 6.0-6 .9 cm carapace length), and cocoon deposition measured
as a function of this parameter.

Finally, 10 cocoon-bearing hosts from each monthly sample were carefully
biopsied under a dissecting microscope to determine the relative distribution of
cocoons. Areas examined included branchial chamber, cephalothorax, pereiopods,
abdominal segments, and pleopods. Relative percentages were calculated for each
area.

RESULTS

Mean worm load per host (Figure 1) decreased significantly when the pond
dried up in September and during a freeze in early January. Infestation varied
from a high of 98.8 worms/host in August to 16.6 worms/host in March. Gradual
recovery was noted when sampling was terminated in June.

BRANCHIOBDELLID (ANNELIDA : OLIGOCH AETA)

287

J2

&

X

H

X

O

r*

isoh uad samaaaoiHDNVdfl m wvaw

Figure 1. Monthly fluctuation in branchiobdellid number/given host. Each mean (± 1
Standard Deviation) represents a sample size of 50 crayfishes.

Observed differences in worm number between successive samples suggested
that host factors were also involved. When mean worm load was tested against

Intermolt

288

THE TEXAS JOURNAL OF SCIENCE

host molt status (Figure 2), intermolt crayfishes were observed to maintain larger
branchiobdellid populations than the corresponding molted animals. Greater vari¬
ation in worm number was also noted for intermolt hosts. When mean worm load
was plotted against host size (Figure 3), a curvi-linear relationship was obtained
in favor of larger crayfishes.

0 • •

6 #

o

isom md somiaaoiHDNViie ** nviw

Figure 2. The relationship of branchiobdellid number to host molt condition. Each mean
(± 1 Standard Deviation) represents a variable number of crayfishes.

BRANCHIOBDELLID (ANNELIDA : OLIGOCH AET A)

289

isoh nad samiaaoiHONVda **= nviw

Figure 3. The relationship of branchiobdellid number to host size. Each mean (± 1 Standard
Deviation) represents a variable number of crayfishes.

As with the adult worms, cocoon deposition (Figure 4) reflected seasonal
changes in the abiotic habitat. In August, nearly 90% of all crayfishes sampled
were found to bear cocoons. The subsequent October sample showed a decline
to 63%. A second and much sharper decrease (to 22%) in cocoon-bearing animals
was observed in March, with subsequent recovery indicated in June.

Cocoon deposition was further assessed as a function of host size (Table 1).
Invariably, lst-year crayfishes (0-1.9 cm) were devoid of cocoons. Only isolated

001

290

THE TEXAS JOURNAL OF SCIENCE

T

O

if)

SN00303 HUM S3HSIJAVH3 %

Figure 4. Percent occurrence of branchiobdellid cocoons on crayfishes from monthly
collections.

JUL AUG OCT MOV DEC JAN iPEfi MAR APR MAT JUN
1972 MONTH 1973

BRANCHIOBDELLID (ANNELIDA :OLIGOCHAET A)

291

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hosts in the 2.0-2 .9 cm range possessed cocoons. Medium-sized crayfishes (3.0-
4.9 cm) appeared to vary as suitable choices for deposition, while larger hosts
(5 .0-6 .9 cm) maintained a high % throughout the year.

Relative distribution of branchiobdellid cocoons on the host body is presented
in Table 2 (since each monthly sample was limited to 10 crayfishes, results are
grouped to coincide with the 4 seasons). The branchial chambers and pleopods
were employed as primary deposition sites throughout the year. The first abdominal

292

THE TEXAS JOURNAL OF SCIENCE

TABLE 2

Relative Distribution of Cocoons on Crayfishes by Seasons.

Body Region

Percent of total deposited cocoons

SUMMER

FALL

WINTER

SPRING

OVERALL

Cephalothorax

0.0

0.0

5.9

0.0

1.5

1st Abdominal seg.

2.3

1.3

6.8

11.0

5,3

2nd Abdominal seg.

1.6

16.4

13.8

13.3

11.3

34d Abdominal seg.

0.3

12.1

5.3

1.8

4.8

4th Abdominal seg.

0.4

1.9

1.9

1.8

1.5

5th Abdominal seg.

0.3

0.0

0.1

0.1

0.2

6th Abdominal seg.

0.0

0.0

0.0

0.0

0.0

7th Abdominal seg.

0.0

0.0

0.0

0.0

0.0

Pleopods

13.5

16.6

31.6

25.2

21.7

Branchial cavity

81.5

50.9

28.6

46.9

52.0

Pereiopods

0.3

0.9

6.0

0.0

1.8

segments ranked as secondary sites. Pereiopods, terminal abdominal segments, and
cephalothorax generally were less favored for deposition.

DISCUSSION

Major decreases in worm load and reproduction of Cambarincola vitrea reflect
meteorological disturbances of the pond in September and early January. Seasonal
reductions in branchiobdellid population density have been reported previously
(Young, 1966; Bishop, 1968), with selective mortality and diminished reproduc¬
tion cited as possible mechanisms. It is likely that such severe disturbances of the
habitat medium accelerate normal selective processes (e.g., juvenile mortality, de¬
creased reproductive rates) that govern the observed fall and spring reductions in
branchiobdellid population density.

In addition to abiotic factors, the results indicate a preferential host selection
by Cambarincola vitrea in favor of larger intermolt crayfishes. Replacement of
the host exoskeleton has been shown to cause irreversible cessation of cocoon
development (Young, 1966) and the mass evacuation of the affected host by adult
worms (Koepp, 1975). Accordingly, the lengthy intermolt period of larger cray¬
fishes makes them ideal for long-term branchiobdellid colonization. The increased
body size also provides a greater surface area for worm inhabitation, cocoon depo¬
sition, and availability of food.

Contrary to previous reports relative to the genus Cambarincola (McManus,
1960; Young, 1966; Bishop, 1968), the branchial cavities and pleopods appear
to be the principal sites for cocoon deposition. While not specifically investigated,
the use of these ventilative loci may reflect an adaptation to satisfy the respiratory
needs of the developing embryos.

BRANCHIOBDELLID (ANNELIDA : OLIGOCH AETA)

293

LITERATURE CITED

Berry, J. W., and R C. Holt, 1959-Reaction of two species of Branchiobdellidae (Oligochaeta)
to high temperature and low oxygen tensions. Va. Agr. Exper. Sta. Tech. Bull., 141:1.

Bishop, J. E., 1968-An ecological study of the branchiobdellis commensals (Annelida-
Branchiobdellidae) of some mid -western Ontario crayfishes. Can. J. Zool., 46:835.

Goodnight, C. J., 1940-The Branchiobdellidae of North American crayfishes.///. Biol. Monogr.,
17:1.

Hoffman, R. L., 1963- A revision of the North American annelid worms of the genus Cam-
barincola (Oligochaeta : Branchiobdellidae). Proc. U. S. Natl. Mus., 114:271.

Holt, R C., 1954-A new branchiobdellid of the genus Cambarincola (Oligochaeta, Branchiob¬
dellidae) from Virginia. Virginia J. Sci., 5:168.

- , 1955 -A new branchiobdellid of the genus Cambarincola from Kentucky./. Tenn.

Acad., 30:27.

- , 1960-The genus Ceratodrilus Hall (Branchiobdellidae, Oligochaeta) with the de¬
scription of a new species. Virginia J. Sci., (N.S.), 11:53.

- , 1963-A new branchiobdellid (Branchiobdellidae: Cambarincola). J. Tenn. Acad.

Sci., 38:97.

Koepp, S. K., 1 975 -Effects of host ecdysis on population structure of the epizootic branchiob¬
dellid Cambarincola vitrea (Annelida: Oligochaeta). Sci. Biol. J., 1(2) :39.

McManus, L. R., 1960-Some ecological studies of the Branchiobdellidae (Oligochaeta).
Trans. Amer. Microsc. Soc., 79:420.

Penn, G. H., 195 9 -Survival of branchiobdellid annelids without a crayfish host. Ecology, 40:514.

Young, W., 1966 -Ecological studies of the Branchiobdellidae (Oligochaeta). Ecology, 47:571.

LIMNOLOGICAL THESES AND DISSERTATIONS
CONCERNING TEXAS WATERS, 1897-1976

by JOHN M. PETTITT and RONALD H. DUTTON

Route 2, Box 328 P, Midlothian 76065

ABSTRACT

Seven hundred seventy-seven titles of limnologically- oriented theses and dissertations
were collected over a five-year period by the author. Universities and colleges in Texas
and bordering states were surveyed by various methods. The titles are indexed according to
key words in the respective titles. Although the major period of time covered is from 1897
through 1975, there is a 1976 ADDENDUM which includes those listings currently in prog¬
ress.

INTRODUCTION

In order to provide easier access to information on subjects related to limn¬
ology in Texas, this compilation has been made of theses and dissertations in
that area. This report is the result of five years of continuous literature surveys
conducted among the various educational institutions in Texas and contiguous
states. The survey methods included a letter addressed to the research section
of each institutional library, letters to the departmental chairmen of all fields
related to limnology, contacts by telephone, and personal visits. Additional
listings were obtained from commencement programs of various graduation
proceedings. A base for the work was provided by two publications:

Clark, W.J., 1966— Publications, personnel, and government organ¬
izations related to the limnology, aquatic biology, ichthyology of
the inland waters of Texas. Technical Report No. 5, Water Re¬
sources Institute. Texas A&M University. 102 pp.

Ferguson, S., 1970— A bibliography of the water-related theses and
dissertations written at the University of Texas at Austin 1897-
1970. Report No. 68, Center for Research in Water Resources,
University of Texas at Austin.

The listing of theses and dissertations is grouped by college or university.
To facilitate use of the list, however, the entries have been numbered and are
indexed below (SUBJECT INDEX). In most cases this index is based upon key

Accepted for publication: January 21, 1977.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

296

THE TEXAS JOURNAL OF SCIENCE

words contained in the titles, as it has not been possible for the author to review
each publication.

It should be noted that copies of the theses and dissertations compiled
herein are available from the library of the institution where the work was
completed. Correspondence suggesting additions, corrections, and deletions
to this report is welcome, and should be addressed to the author: John Pettitt,
Route 2, Box 328 P, Midlothian, TX 76065.

ANGELO STATE UNIVERSITY

1. McClung, G. Dan, 1975 -The phylogenetic relationships of some species of Gambusia

occurring in Texas. M.S.

BAYLOR UNIVERSITY

2. Alexander, T.C., 1940-A biological survey of the Bosque River system, Waco,

Texas. M.S.

3. Bane, Carol, 1975 -The limnology of aquatic systems of Big Bend National Park

with emphasis on water quality. M.S. (In progress)

4. Bothner, G.L., 1957-An age and growth study of Ictiobus bubalus in Lake Waco,

Texas. M.S.

5. Chen, T., 1969-Age and growth of the white crappie, Poxomis annularis (Rafin-

esque)in Lake Waco, Texas. M.S.

6. Davidson, F.F.,1933-The algae of McLennan County, Waco, Texas. M.S.

7. Harrell, H.L., 1975 -Fishes of Devil’s River. M.S.

8. Kennedy, S., 1975 -The ecology of the lean springs pupfish, Cyprinodon bovinus.

M.S. (In progress)

9. Lade, O.R., 1934-Food of fish in McLennan County. M.A.

10. Lewand, R.L., 1967-The geomorphic evolution of the Leon River system. M.S.

11. Lukins, David L., 1975 -Production and diversity of aufwuchs populations in

thermal effluents. M.S. (In progress)

12. Moore, T.H., 1968 -Geochemistry of waters of Hog Creek Basin; Hamilton, Coryell

and Bosque Counties, Texas. M.S.

13. Moore, T.D. , 1965 -Growth and development of the Brazos River Authority. M.S.

14. Proctor, C.V., 1967-The North Bosque watershed: Inventory of a drainage basin.

M.S.

15. Stewart, M.B.W., 1967-An ecological study of the bacterial flora of Comanche

Spring and Harris Creek. M.S.

16. Scott, C., 1975 -Factors influencing transverse stability in fishes. M.S.

17. Venables, B.J., 1972 -Some effects of thermal history on the large mouth black bass,

Micropterus salmoides (Lacepede) of two central Texas reservoirs receiving
power plant effluents. M.S.

TEXAS WATERS

297

18. Waldorf, G.W., 1942-Fall and winter food habits of fish of the Bosque River system,

Waco, Texas. M.A.

19. West, G.H., 1932-Fishes of McLennan County. M.A.

20. Wilson, K.R., 1975 -Phosphate uptake kinetics of natural phytoplankton populations

as affected by temperature. M.S. (In progress)

21. Wysong, M., 197 3 -The ostracods Cypridopsis vidua: Physiological response due

to thermal stress. M.S.

EAST TEXAS STATE UNIVERSITY

22. Bradley, M.J., 1975-Species diversity of fish populations in three Northeast Texas

rivers. M.S.

23. Campbell, D., 1975-A study of feeding habits and condition of Micropterus salmoides

in Lake Davy Crockett, Texas. M.S.

24. Clay, David, 1975 -An age-growth study of black bass in Lake Davy Crockett, Texas.

M.S.

25. Duncan, C.W., 1971 -Species diversity of benthic macroinvertebrates in the South

Sulphur River. M.S.

26. Glueck, D., 1975-A vegetative analysis of beaver habitats along the South Sulphur

River. M.S.

27. Henry, D., 1975 -Food competition between channel catfish and black bullheads

in newly constructed farm ponds. M.S.

28. Long, H., 1963-A study of the algae population of the Hefley-Taylor Lake of

Commerce, Texas. M.S.

29. Patterson, B., 1975-Population analysis of red-eared turtles in an East Texas farm

pond. M.S.

30. Roberts, D., 1975 -An ecological study of red-eared turtles in an East Texas farm

pond. M.S.

31. Schultz, G.G., 1968 -Chlorinated hydrocarbon residues in aquatic plants. M.S.

32. Seglem, M., 1975 -Effects of Toxic metals from industrial plating solutions on large

mouth bass, Micropterus salmoides. M.S.

LAMAR UNIVERSITY

33. Ashcraft, J.A., 197 3 -Physio-chemical conditions and macrobenthic community

structure in the lower Neches River. M.S.

34. Chaiyarach, S., 1974-Acute toxicity of toxaphene and ordram to four species of

aquatic organisms. M.S.

35. Duplechin, J., 1975-The effects of municipal sewage effluent, municipal surface

runoff, and rice fields irrigation water on community structure of macrobenthos
in Hillebrandt Bayou. M.S.

298

THE TEXAS JOURNAL OF SCIENCE

36. Gibson, G.E., 1974-Seasonal variations in physico-chemical conditions of sewage

effluent through the Beaumont, Texas, treatment plant and receiving stream. M.S.

37. Howard, R., 1973-Physico-chemical conditions and communities of macrobenthos

in the lower Neches River.

38. Lewis, S.P., 1974-Physico-chemical conditions and macrobenthos of Village Creek.

M.S.

39. Marsh, C.E., Jr., 1973-Seasonal abundance and community structure of algae and

rotifers in Massey Lake, Southeast Texas. M.S.

40. Olson, K.R., 1971 -Salinity effects on the TLm for mercury, copper, and chromium

for Rangia cuneata. M.S.

41. Patterson, L.D., 1971 -Physico-chemical conditions and community structure of

benthic macroinvertebrates in the Neches River in the vicinity of the salt water
barrier. M.S.

42. Ratananun, V., 1974-Toxicity of the insecticide Sevin and the herbicide Propanil

to four aquatic species. M.S.

43. Sala, G.M., 1975— Benthic macroinvertebrates in the Beaumont tertiary sewage

treatment ponds and receiving stream. M.S.

44. Siegal, B.A., 1975— Physico-chemical conditions and community metabolism in

a Southeast Texas oil refinery holding pond. M.S.

45. Welch, M.O., 1973-Seasonal variation of physico-chemical characteristics and

benthic macroinvertebrates in a southeast Texas meander scar lake. M.S.

MIDWESTERN UNIVERSITY

46. Bustetter, L.A., 1972-Membrane filter characteristics of fecal streptococcal pol¬

lution in Holliday Creek, Wichita County, Texas. M.S.

47. Whitt, J.W., 1971— An ecological study of Cladoceran populations and selected lit¬

toral regions of Lake Wichita. M.S.

NORTH TEXAS STATE UNIVERSITY

48. Abshire, Robert Louis, 1970-Immunoflorescence as a method for the rapid iden¬

tification of Streptococcus foecalis in water. Ph.D.

49. Alam, I.M., 1976-Phytohormone effects in algae. Ph.D.(In progress)

50. Allison, Richard C., 1966 -Microbio tic cycles in Lake Hefner. M.S.

51. Bagwell, Russel L., 1938-A study of the phytoplankton population of Lake Dallas.

M.S.

52. Beddow, D.G., 1966-Some physiological effects of chlorine upon two chlorine

resistant algae. M.S.

53. Benson, D., 1976— Production and energy metabolism in Midge and Mayfly pop¬

ulations in North Central Texas pond ecosystem. M.S. (In progress)

TEXAS WATERS

299

54. Boswell, James T., 197 6 -The use of adenosine triphosphate (ATP) assays in de¬

scribing the Limnology of Moss Reservoir. Ph.D. (In progress)

55. Brooks, Benjy F., 1940-A bio-chemical comparative study of the plankton in Lake

Dallas and Pecan Creek. M.S.

56. Brown, A.V., 1974 -Ecological energetics of the Dobsonfly, Corydalus cornutus.

Ph.D.

57. Brown, J.H., 195 7 -An investigation of methods for the concentration of chemical

compounds produced by actinomycetes and their relation to tastes and odors
in municipal water supplies. M.S.

58. Cahoon, R., 1976-Decomposition rates in a small North Central Texas pond eco¬

system. M.S. (In progress)

59. Chapman, R., 1973-Effects of ethephon on Scenedesmus quadricauda. M.S.

60. Cloud, Thomas J., 1973-Drift of aquatic insects in the Brazos River, Texas. M.S.

61. Cooper, W.A., 1950-Age, growth, and feeding habits of the large-mouthed black

bass ( Micropterus salmoides ) and the spotted bass {Micropterus punctulatus sp.)
in North-Central and East Texas lakes. M.S.

62. Cook, V., 1946-A quantitative and qualitative bacterial analysis of Pecan Creek.

M.S.

63. Dougherty, J.H., 1957-Survival and growth of bacteria in chlorine treated water.

M.S.

64. Durrett, Charles Wade, 1973-Density, distribution, production, and drift of benthic

fauna in a reservoir receiving thermal discharges from a steam-electric generating
plant. M.S.

65. Dill, W.S., 1951 -The chemical compound produced by actinomycetes and their

relation to tastes and odors in a water supply. M.S.

67. Elliott, J.M., 1948-The age and rate of growth of the black crappie, Pomoxis nigro-

maculatus (Le Seur), and the white crappie, Pomoxis annularis (Rafinesque),
in the Koon Kreek Klub Lakes, Texas. M.S.

68. Estes, C.M.,1949-The fecundity of the bluegill ( Lepomis macrochirus ) in certain

small East Texas reservoirs. M.S.

69. Evans, A. A., 1936-Study of plankton dilution in source streams compared with

that of Lake Dallas proper. M.S.

70. Faggard, J.M., 1940-An analysis of the seasonal food habits of two species of Texas

Centrarchids. M.S.

71. Fair, Helena Juengerom Ann., 1952-Toxicity studies of aquatic actinomycetes. M.S.

72. Farrell, R., 1975-Sporulation control in heterocystous blue-green algae. M.S. (In

progress)

73. Foster, R.M., 1937 -A yearly study of the bacterial flora of Lake Dallas water with

special reference to sanitation. M.S.

74. Foerster, J.W., 1964-The synecology of phy co-periphyton in oligotrophic lakes. M.S.

300

THE TEXAS JOURNAL OF SCIENCE

75. Friday, Gary Phillip, 1971-Spring food habits of Corydalus cornutus in the Brazos

River, Texas. M.S.

76. Goodman, J.W., 1959-A water quality study of Lake Texoma. M.S.

77. Gough, L.C., 1949-Chlorophyceae of Erath County. M.S.

78. Graham, E., 1937-The ecology of Chlorophydra viridissima in a small perennial

pond. M.S.

79. Grizzle, Walter Ray, 1951 -Pathogenic bacterial survey in the Trinity River, from

Ft. Worth, Texas, to South Dallas, Texas. M.S.

80. Hansard, John D,, 1969-Distribution of phosphates in a sewage plant and its re¬

ceiving water. M.S.

81. Harmon, J.C., 1956-An investigation of algae and common tastes and odors in

in freshwater. M.S.

82. Hemphill, Louis, 1952-The application of chlorine dioxide to tastes and odors

in water supplies. M.S.

83. Hendricks, Albert Cates, 1970-Eutrophic levels of different areas of a reservoir:

a comparative study. Ph.D.

84. Henley, Don E., 1968— Isolation and identification of an odor compound produced

by a selected aquatic actinomycete. M.S.

85. - , 1970-Odorous metabolite and other selected studies of Cyanophyta.

86. Higgins, M.L., 1964-The life cycle of an aquatic actinomycete. M.S.

87. Hill, J.F., 1939-The food and growth of the large-mouthed black bass (. Huro

salmoides) in north Texas. M.S.

88. Horkel, J.D., 1974-Effects of turbidity on gilling rates and oxygen comsumption of

green sunfish. M.S.

89. Jenkins, J.D., 1940-Bottom fauna of Lake Worth. M.S.

90. Johns, W.B., 1939-A limnological study of Lake Worth. M.S.

91. Johnston, P.M., 1942-The seasonal cycle in the testis of the large-mouthed black

bass, Huro salmoides (Lacepede). M.S.

92. Jones, R.J., 1969-Quantitative aspects of the microflora of an overland flow spray

irrigation sewage disposal system. M.S.

93. Jones, B.J., 1976 -Hematological parameters of the Bluegill, Lepomis macrochirus

(Rafinesque) including effects of turbidity, chloroamines and Flexibacter col-
umnatis. M.S.

94. Jones, F.V., 1975 -Population dynamics and production of seven fishes in a small

pond. M.S.

95. Kelly, M.H., 1975 -Primary productivity and community metabolism in a small

North Central Texas pond ecosystem. M.S.

TEXAS WATERS

301

96. Koen, L.D., 197 2 -The food habits of bluegills in a lake receiving heated effluent.

M.S.

97. Lamb, L.D., 1938-A comparative study of the bottom fauna of four Texas lakes.

M.S.

98. Lawley, Gary G., 1973-Biological nitrogen fixation in two Southwestern reservoirs.

Ph.D.

99. Legett, J.H., 1940-A net plankton survey of a small perennial pond. M.S.

100. Maki, Alan Walter, 1971 -Influence of sublethal pesticide levels on respiratory

activity of selected aquatic animals. M.S.

101. Maran, J.E., 1964-Benthic algae of selected thermal springs of Yellowstone Na¬

tional Park. M.S.

102. Mardcastle, Ronald Van, 1970-A screening of fungi for metabolites inhibitory to

the growth of bloom-forming blue-green algae. M.S.

103. McClellean, W.G., 195 3- A study of the southern spotted channel catfish, Ictalurus

punctatus (Rafinesque). M.S.

104. McCormick, W.C., 195 4 -The cultural, physiological, morphological, and chemical

characteristics of an actinomycete from Lake Waco. M.S.

105. McDaniel, M.D., 1973-Trophogenic ecology of selected Southwestern reservoirs.

Ph.D.

106. McIntosh, J., 1976-Energy flow in seven fish populations in a small North Central

Texas pond ecosystem. M.S. (In progress)

107. McNeely, D.L., 1973-Distribution, size, condition, and food habits of selected

fishes in a reservoir receiving heated effluent from a power plant. M.S.

108. Micks, D.W., 1942 -The age, growth, and food habits of the Lake Dallas white bass,

Lepibema chrysops (Rafinesque). M.S.

109. Milliger, L.E., 1969-Studies of the passive dispersal of viable algae and protozoa

by aquatic and terrestial beetles. Ph.D.

110. Mitchell, G.C., 1941-An analysis of the seasonal food habits of black and white

crappie taken from Texas lakes. M.S.

111. Mitterer, G., 1976-Distribution, abundance and food habits of larval fishes in a

reservoir receiving heated effluents from a power plant. M.S. (In progress)

112. Neal, G.C., 1941 -The phototrophic properties of Lactuca ludoviciana (Nutt.)

DC and Silphium laciniatum L. M.S.

113. Newton, C.E., 1973-Carbon flux in reservoir sediments. Ph.D.

114. Northam, F.E., 1975 -Pecan Creek: some effects of urbanization on a small stream.

M.S. (In progress)

115. Parsons, W.M., 1966-Dispersal of algae and protozoa by selected Odonata. M.S.

116. Peterson, J.A., 1 97 2 -Indole-3-acetic acid effects on the nucleic acids of Chlorella

pyrenoidosa. M.S.

302

THE TEXAS JOURNAL OF SCIENCE

117. Pipes, W.O., 1955 -An investigation of naturally occurring tastes and odors from

freshwater. M.S.

118. Redden, D.R., 1949-The physical, chemical and biological factors affecting algae

blooms in freshwater reservoirs. M.S.

119. Revill, D.L., 1966— Transport of viable dissemules of algae and protozoa by selected

diptera. M.S.

120. Rhame, R.E., 1973-Life cycle and ecology of the Hydropsy chidae (Trichoptera)

of the Brazos River, Texas. Ph.D.

121. Richey, Harvey, 1973-Studies of selected Cyanophyta response to varying geo sm in

concentrations. M.S.

122. Russell, J.C., 195 2 -Biological indices of stream pollution. M.S.

123. Sams, Barry L., 1976 -Comparative chemistry of thermally stressed North Lake and

its water source, Elm Fork of the Trinity River. M.S.

124. Sherman, R.C., 1946-Seasonal food habits of five species of Texas Centrarchids.

M.S.

125. Sivels, H.C., 1945 -A preliminary investigation of the effects of various fertilizers

on plankton and fish production in small Texas ponds. M.S.

126. Smith, J.A., 1973-Primary productivity and nutrient relationships in Garza-Little

Elm Reservoir. Ph.D.

127. Smith, C.G., 1948-The reproductive cycles of five species of Texas Centrarchids.

M.S.

128. Smith, B.A., 1938-A physical and chemical investigation of Eagle Mountain Lake

with reference to biological productivity. M.S.

129. Smith, G.A., 1976-Population dynamics and energy metabolism of microcrusta¬

cean zooplankters in a North Central Texas pond ecosystem. M.S.

130. Solon, B.M., 1970-Passive dispersal of algae and protozoa internally and externally

by selected aquatic insects. Ph.D.

131. Stark, W.P., 1972-The stoneflies (Plecoptera) of Oklahoma. M.S.

132. Stiles, J.C., 1968- A bacteriological survey of a freshwater reservoir. M.S.

133. Stuart, T.J., 1975-Effects of entrainment and passage through an electric power

generator condenser and thermal effluent on phytoplankton populations and
their photosynthetic activity. M.S. (In progress)

134. Szczytko, Stanley W., 1978 -The biology and systematics of the genus hoperla

(Plecoptera) of western North America. Ph.D. (In progress)

135. Trotter, D.M., 1969- A comparison of the carbon dioxide to oxygen rate of change

methods for measuring primary productivity. M.S.

136. Tsai,C.E.J., 1973-Methylglyocal metabolism in Scenedesmus quadricauda. M.S.

137. Vaught, G.L., 1972 The life history and ecology of the stone fly Neoperla clyment

(Newman). M.S.

TEXAS WATERS

30.3

138. Venables, B., 1976-Thermal ecology of large-mouth bass in North Central Texas.

Ph.D.

139. Welch, H., 1939-A chemical investigation of Lake Dallas to determine the factors

influencing plankton growth. M.S.

140. Wilcox, D.P., 1976-The relationship between biomass and primary production

in the phytoplankton community of a mesotrophic Texas reservoir. M.S. (In
progress)

141. Woods, A.M., 1937-A comparative investigation of the water of Lake Bridgeport

with reference to plant and animal life. M.S.

142. Wyatt, J.T., 1969-Selected physiological and bio-chemical studies on blue-green

algae. Ph.D.

OKLAHOMA STATE UNIVERSITY

143. Whiteside, Bobby Gene, 1964-The biology of the white crappi e Poxomis annularis,

in Lake Texoma, Oklahoma. M.S.

PAN AMERICAN UNIVERSITY

144. Penn, G.J., 1974-Seasonal periodicity of dominant, benthic, marine macroalgae of

the north jetty, Brazos-Santiago Pass, Texas, as related to environmental factors.
M.S.

RICE UNIVERSITY

145. Crowley, P.H., 197 2 -Filtering rate inhibition of Daphnia pulex in Wintergreen Lake

water. M.S.

146. Keegan, G.J., 1970-Water quality management. Optima allocation of pollutant

discharges constrained by quality and equity considerations. M.S.

147. McRitchie, R.G., 1968-Effects of thermal and osmotic acclimation on an estuarine

gastropod. Ph.D.

SAM HOUSTON STATE UNIVERSITY

148. Altaras, R.M., 1972-The effects of domestic sewage on the distribution of the fish

fauna of Harmon Creek, Walker County, Texas. M.A.

149. Barrett, S.R., 1974-A morphological and meristic study of Menidia (Bonaparte)

populations in the Trinity River system, East Texas (Pisces, Atherinidae). M.A.

150. Boyd, J.F., 1968-A taxonomic and ecological survey of the fish fauna of Bedias
Creek, Texas. M.A.

Cline, E.M., 1949-A study of the pH variations in the Sam Houston Park pond
water. M.A.

151.

304

THE TEXAS JOURNAL OF SCIENCE

152. Harper, A.L., 1969-Salinity tolerances of five species of East Texas teleosts. M.A.

15 3. King, J.M., 1968 -The investigation of freshwater algae populations in Walker and
Montgomery counties. M.A.

154. Lasswell, J., 1968 -A taxonomic and ecological survey of the fish fauna of the West

Fork of the San Jacinto River, Texas. M.S.

155. Thurston, N.I., 1969-An ecological survey of a pondside region in Walker County,

Texas. M.A.

156. Tipton, J.L., 1974-The removal of selected elements from a germination medium

by Anabaena akinetes. M.A.

157. Twidwell, S.R., 1971 -Effects of condenser cooling water on community structure

of net phytoplankton and related physicochemical features of Lewis Creek Res¬
ervoir, Texas. M.A.

158. Wiley, E.O., III, 197 2 -Geographic variation and systematics of the Fundulus nottii

(Agassiz) species complex (Pisces, Cyprinodontidae). M.A.

SOUTHERN METHODIST UNIVERSITY

159. Allen, M.R., 1964-An ecological survey of the recent land and freshwater gastro¬

pods of Wichita County, Texas. M.S.

160. Anderson, D.S., 1968 -The aquatic Coleoptera in the vicinity of Oak Ridge, Ten¬

nessee. M.S.

161. Callahan, W.P., 195 3 -The distribution of coliform bacteria in a reservoir lake as

related to dissolved oxygen. M.S.

162. Dean, P.N., 1967 -An ecological survey of the living terrestrial and freshwater

gastropods of El Paso County, Texas. M.S.

163. Dust, J.V., 1963-A quantitative comparison of chlorophyll “a” with algae count

and classification in sewage treatment oxidation ponds. M.S.

164. Erickson, S.J., 1952-Effects of weather conditions on the Turtle Creek plankters

near Dallas, Texas. M.S.

165. Ferguson, A.K., 1937-Systematic and ecological notes on the Odonata of the vi¬

cinity of Dallas, Texas. M.S.

166. Fullwood, P.D., 1950-A survey of parasitic copepods infesting fish in Dallas County,

Texas. M.S.

167. Fullington, R.W., 1969-A study of the distribution, ecology, and taxonomic status

in the gastropod families Physidae and Lymnaeidae in Texas. M.S.

168. Flook, J.M., 1970-A survey of Metazoan parasites in Unionid bivalves of Garza-

Little Elm Reservoir, Denton County, Texas, with descriptions of a near species
of Rhopalocercous cercaria.

169. Glass, V., 1949 -Report on the aquatic Coleoptera of the Dallas area. M.S.

170. Goff, J.D., 1958-A study of the water quality of the Trinity River in Dallas County.

M.S.

TEXAS WATERS

305

171. Hall, C.C., 1950-The Trichoptera of Dallas County, Texas. M.S.

172. Haynes, F., 1942-A study of the food habits of large-mouth black bass fingerlings

( Huro salmoides ) in rearing ponds. M.S.

173. Hasty, M.A., 1950-Plankton studies on Denton Creek, Texas. M.S,

174. Logsdon, W.T., 1967 -An ecological survey of the recent land and freshwater gas¬

tropods of Kaufman County, Texas. M.S.

175. Majors, E.C., 1964-A partial survey of the freshwater and land gastropods in the

area of Harlingen, Cameron County, Texas. M.S.

176. McCleskey, O., 1949-The bionomics of the Culicinae of the Dallas, Texas, area.

M.S.

177. Millspaugh, D.D., 19 36 -Bionomics of the aquatic and semi-aquatic Hemiptera of

Dallas County, Texas. M.S.

178. Moore, L.E., 1950-Distribution of mayfly nymphs (Ephemeroptera) in a stream

of Dallas County, Texas. M.S.

179. Oliver, K.H., 1952-The effect of herbicides on selected zooplankton. M.S.

180. Patterson, M., 1942-A study on fluctuations in plankton populations in White

Rock Lake in winter and spring. M.S.

181. Read,L.B., 195 3 -The Pelecypoda of Dallas County. M.S.

182. Robbins, D.M., 1968-A seasonal study of certain zooplankton in Turtle Creek,

near Dallas, Texas. M.S.

183. Smith, K.K.L., 1972-A limnological survey of Lake Granbury (DeCordova Bend),

Texas. M.S.

184. Zischkale, M.A., 1951 -The effect of rotenone and some common herbicides on fish

food organisms. M.S.

SOUTHWEST TEXAS STATE UNIVERSITY

185. Arnold, C.R., 1962-Oxygen consumption as a factor in the distribution of certain

Trichoptera larvae. M.A.

186. Anderson, B.T., 1968-A comparative primary productivity study of three ponds

bearing different primary producers. M.A.

187. Barra, T.C., 1976-The effects of a heated effluent on the macroinvertebrate com¬

munity in Calaveras Lake, Texas. M.S.

188. Barker, E., 1969-A comparative study of the Hydracarina (Acarina) of three per¬

manent ponds. M.A.

189. Bayer, C., 1975 -The distribution of Odonates of the Guadalupe River Basin, Texas.

M.A.

190. Becker, P.R., 1969-The systematics and seasonal distribution of the Cladocera

of Hays County, M.A.

306

THE TEXAS JOURNAL OF SCIENCE

191. Broz, L., 1974-The influence of a deep-storage reservoir and an underground res¬

ervoir on the physico-chemical limnology of a Central Texas River. M.A.

192. Bruchmiller, J.P., 197 3 -Description and keys to the macrophytes of Spring Lake

(excluding algae). M.S.

193. Carraway, Joe, 1975-The ecological distribution of water mites- of the genus Ar-

renurus in Hays County, Texas. M.A.

194. Chandler, T., 197 6 -Influence of rain runoff and sewer effluents on the Blanco and

San Marcos Rivers. M.S.

195. Colbert, B.K., 1973-Plankton periodicity in two man-made ponds in Central Texas.

M.S.

196. Conrads, C., 1972-The effects of selected environmental parameters on the hatching

of Eulimnodia texana. M.A.

197. Conrads, L.M., 1969 -Demography and ecology of the Fern Bank salamander,

Eurycea pterophila. M.A.

198. Cover, L., 1975-The drift ecology of the Trichoptera of the San Marcos River.

M.A.

199. Davis, J., 1975-Ecology of the helminths of Gambusia affinis in the area of San

Marcos, Texas. M.S.

200. Devall, L.L., 1940 -A comparative study of plant dominance in a spring-fed lake.

M.S.

201. Dowden, D.L., 1968 -Population dynamics of the San Marcos salamander, Eurycea

nana. M.A.

202. Hamilton, A., 1973-Some taxonomic aspects of certain paedogenetic Eurycea of

the Blanco River drainage system in Hays and Blanco Counties, Texas. M.A.

203. Henderson, E., 197 2 -Population dynamics of the Lepomis cyanellus (male) X

Lepomis microlophus (female) hybrid as compared with its reciprocal cross and
parental species. M.A.

204. Kent, David, 1971 -The effects of a deep-storage reservoir on the benthic macro¬

invertebrate community of the Guadalupe River, Texas. M.A.

205. Kneupper, A.C., 1971 -An investigation of specific taxonomic feature used in the

description of Loptestheria compleximanus . M.A.

206. Kolb, J., 1975 -Age and growth of large-mouth bass in Canyon Lake, Texas. M.S.

207. Litchfield, K., 1975-The macrophytes of the upper San Marcos River. M.A.

208. Lockett, C., 1975— On the classification of 18 Central Texas reservoirs. M.A. (In

progress)

209. Martin, M., 1975 -A survey of the helminths of the shiners Notropis venustus and

Notropis lutrensis in the area of San Marcos, Texas. M.A.

210. Mayhew, JJ., 1970-Nitrogen and phosphorus dynamics in a 153-kilometer stretch

of the Guadalupe River, Texas. M.A.

211. McNatt, R., 1968-Fish species diversity in relation to stream order and physico¬

chemical conditions in the Plum Creek drainage basin. M.A.

TEXAS WATERS

307

212. Munson, J.W., 1946 -The distribution of plankton in the San Marcos River, San

Marcos, Texas. M.S.

213. Parsons, D., 1970-A limnological study of the New Braunfels-Gonzales stretch of

the Guadalupe River. M.A.

214. Peters, M.S., 1975 -The Ephemeroptera nymphs of the Guadalupe River basin,

Texas. M.S. (In progress).

214a Phillips, T.R., 1975 -Seasonal succession of plankton in a cooling reservoir. M.S.
(In progress)

214b Prentice, J.A., 197 3 -Validation of aging techniques for large-mouth bass, Micro-
pterus salmoides and channel catfish, Ictalurus punctatus, in Central Texas farm
ponds. M.S.

214c Ralph, JJ., 1975 -Adsorption and desorption of cations in aquatic sediments. M.S.
(In progress)

2l4d Rhodes, C., 1973 -The influence of oxygen concentrations on the activity and sur¬
vival of Hydracarina. M.A.

214e Schenck, John, 1975-Ecology of the fountain darter, Etheostoma f on ticola (Oste-
ichthyes: Percidae). M.S.

214f Strenth, N.E., 1970-A morphological study of the post-embryonic stages of Eu-
limnadia texana (Conchostraca, Crustacea). M.A.

214g Tatum, J.W., 1970 -Physico-chemical limnology and chlorophyll “a” of the New
Braunfels-Gonzales, Texas, stretch of the Guadalupe River. M.A.

214h Warren, J., 1975 -Subterranean aquatic organisms as potential indicators of ground-
water pollution. M.S. (In progress)

214i Woerner, H., 1971 -Multiple regression analysis of 19 variables influencing eutrophi¬
cation in a 153-km stretch of the Guadalupe River, Texas. M.A.

214k Young, W.J., 1971 -The influence of impoundment and thermal stratification in
Canyon Reservoir on the physicochemical limnology and chlorophyll a of the
Guadalupe River, Texas. M.A.

STEPHEN F. AUSTIN STATE UNIVERSITY

215. Adler, P.M., 1972-A study of eutrophication in upper Sam Rayburn Reservoir. M.S.

216. Alford, H.W., 1950-Bacteriological analysis of rural water supplies in Nacogdoches

County, Texas. M.S.

217. Allard, D.W., 1974-Zooplankton population dynamics at Sam Rayburn Reservoir.

M.S.

218. Bachtel, H.J., 1940 -Freshwater mussels of East Texas. M.S.

219. Bailey, T.F., 1970-Food habits of adult Pomoxis annularis and Poxomis nigro-

maculatus in Sam Rayburn Reservoir, Texas. M.S.

220. Bowers, J.H., 1962-Food habits of water snakes, genus Matrix, in Bowie and Red

River Counties, Texas. M.S.

308

THE TEXAS JOURNAL OF SCIENCE

221. Castleberry, W.B., 1942- A biological investigation of the common fish of Central

East Texas. M.S.

222. Cox, D., 1975 -Benthic macroinvertebrates as indicators of water quality in the

Angelina River, Texas. M.S. (In progress)

223. Dawson, J.E., 197 3 -The influence of Sam Rayburn Reservoir on the water quality

of the Angelina River and Attoyac Bayou. M.S.

224. Dickens, F.A., 1950-A distributional study of fishes in the Nacogdoches area. M.S.

225. Foster, R.E., 1972-A survey of aquatic beetles in the city of Nacogdoches, Texas,

and environs. M.S.

226. Hale, J.N., 1973-Existing water quality for the proposed site of Lake Nacogdoches.

M.S.

227. Hanicak, E., 1946-Black bass of East Texas. M.S.

228. Harris, M.A., 1973 -Analysis of the Trinity River coliform and streptococcus bacteria

as indicators of fecal pollution. M.S.

229. Harry, D.N., 1974-A limnological comparison of selected Texas reservoirs. M.S.

230. Johnson, P.D., 1975 -Analysis of water chemistry in Striker Reservoir. M.S. (In

progress)

231. Lock, J.T., 1967 -The minimal dissolved oxygen requirements of Ictalurus punctatus

and Ictalurus melas. M.S.

232. Mulvihill, S.R., 1976-Physico-chemical limnology of the Angelina River, Texas. M.S.

233. Riley, M., 1975 -Some of the effects of thermal discharges on benthic macroinverte¬

brates in a cooling reservoir. M.S.

234. Rodgers, G.D., 1976-Comparison of zooplankton communities in Murval Reservoir

and Striker Reservoir. M.S.

235. Rodgers, J.S., 1967 -Interspecific hybridization in sunfishes ( Lepomis ) of Eastern

Texas. M.S.

236. Rudy, K.C., 1975-The effects of hot water effluents from an electric generating

plant on phytoplankton primary productivity. M.S.

237. Snelgrove, R.E., 1975 -A survey of organochlorine pesticide residues in the Angelina

River and Attoyac Bayou. M.S. (In progress)

238. Sniffen, R.P., 1973-A linear water quality study of Bayou Lanana from Nacag-

doches, Texas, to the Angelina River. M.S.

239. Underwood, H.T., 1975 -A comparative study of the parasitism of Micropterus

salmoides, large-mouth bass, in two arms of Sam Rayburn Reservoir. M.S.

240. Wahrer, R., 1976-The periphyton community of Striker Reservoir. M.S.

241. Walker, W.T., 1973-Physical and chemical limnology of the Trinity River, Texas,

and the effects of impoundment upon water quality. M.S.

242. Whitman, R.L., 1973-Survey of pesticide residues in sediments from the Upper

Trinity River Basin, Texas. M.S.

TEXAS WATERS

309

243. Wood,J.M., 1972-Feeding habits of large-mouth bass, Micropterus salmoides (Lace-

pede), from the headwaters of Toledo Bend Reservoir. M.S.

TEXAS A & I UNIVERSITY

244. Perez, R.G., 1971 -Relationships between phytoplankton and chlorophyll standing

crop in two permanent freshwater ponds in a marine supratidal environment. M.S.

245. Pineda, P.H.A.K., 1975 -A study of fishes of the lower Nueces River. M.S.

246. Serota, T., 1971 -Relationships between primary productivity and chlorophyll

standing crop in two permanent freshwater ponds in a marine supratidal en¬
vironment. M.S.

TEXAS A & M UNIVERSITY

247. Adams, C.S., 1935 -Annual rainfall and runoff relations on the Trinity River water¬

shed above, Dallas, Texas. M.S.

248. Albaugh, D.W., 1973-Life histories of the crayfishes Procambarus acutus and

Procambarus hinei in Texas. Ph.D.

249. Allard, D., 197 6 -Productivity of littoral microcrustaceans in a South Texas pond.

Ph.D. (In progress)

250. Allison, R.C., 1970 -Algal uptake of selected organic compounds. Ph.D.

251. Anderson, A.J., 1974- A method for determining the optimization of alternate

uses of river oxbows. Ph.D.

252. Bedford, W.B., 1972-The physiological ecology of the estuarine clam Rangia cuneata

(Gray). Ph.D.

253. Bedinger, Jr., C.A., 1974-Seasonal changes in condition and biochemical consti¬

tuents of the brackish water clam Rangia cuneata (Gray). Ph.D.

254. Bellamy, J.D., 1941 -Organography of the Notropis potteri (Hubbs). M.S.

255. Bennett, J., 1975 -Polarized light reception in turtles. M.S. (In progress)

256. Berti, A.L., 19 39 -A survey of anopheline larvae and breeding places at College

Station, Texas. M.S.

257. Brooks, J.M., 1970-The distribution of organic carbon in the Brazos River basin.

M.S.

258. Brown, P.J., 1974-An investigation of the toxin of Microcystis aeruginosa: Effects

of varying the concentration of selected salts upon toxicity and the effects of
the toxin on some physiological parameters. Ph.D.

259. Burchell, L.C., 1974-Rural reservoir recreation impact: perceptions of knowledge¬

able leaders. Ph.D.

260. Calnan, T., 1975-Systematics and distribution of the metazoan parasites in the

Unionidae in the Navasota River, Texas. M.S.

310

THE TEXAS JOURNAL OF SCIENCE

261. Calvin, T.L., 1974-An ecological study of the amphibians and reptiles of the Nava-

sota River, Texas. M.S.

262. Carillo, S.J., 1941-A survey of anopheline larvae in 39 stock tanks in Brazos County,

Texas. M.S.

263. Champ, M.A., 1973— Organic and inorganic carbon cycles in a pond ecosystem. Ph.D.

264. Clements, G., 1975 -Relation of phytoplankton population to pond fertilization.

M.S. (In progress)

265. Colle, D.E., 1976-The food habits of three centrachids in a Central Texas farm pond.

M.S.

266. Cowan, W.L., 1933-A study of the rainfall and the stream flow of the Navasota

River system. M.S.

267. Crew, Henry, 1970- A numerical model of the dispersion of a dense effluent in a

stream. Ph.D.

268. Cole, D.R., 1973-The effects of DDT and other insecticides on accumulation,

growth, and photosynthesis in Chlorella pyrenoidosa (CHICK). Ph.D.

269. Daen, Philip, 197 6 -Polarized light reception in fish and sharks. M.S. (In progress)

270. Davy, F.B., 1973-The role of chemoreception in teleost locomotion behavior as

affected by a pesticide. Ph.D.

271. Drier, Thomas, 1976-Use of the scanning electron microscope in quantitative es¬

timates of aquatic bacterial populations. M.S. (In progress)

272. Dry, Eddie Maurice, 1975 -An economic analysis of the Brazos River waterway.

Ph.D.

273. Dziuk, Larry J., 1971 -A study of pesticide residue levels and insecticide resistance

in selected aquatic organisms occurring around the Bryan-College Station ag¬
ricultural production areas. M.S.

274. Evans, J.W., 1975 -Longitudinal distribution of fish in an East Texas stream. M.S.

(In progress)

275. Farrell, WJ., 1965 -Distribution of the Cladocera of Brazos County. M.S.

276. Gallaher, W.B., 1974-A limnological investigation of the relationship between the

Navasota River, Texas, and a selected floodplain. Ph.D.

277. Gauntt, W.C., 1939-Studies of winter hibernation of Anopheles larvae at College

Station, Texas. M.S.

278. Glaze, F.M., 1973-An analysis of rural subdivision development on selected res¬

ervoirs in the Trinity River Basin Texas. Ph.D.

279. Gleastine, Billy W., 1974-A study of the cichlid Tilapia aurea (Steindachner) in

a thermally modified Texas reservoir. M.S.

280. Gillard, R.M., 1974-Distribution, abundance, and species diversity of macrobenthic

and meiobenthic invertebrates in relation to Houston Ship Channel pollution
in upper Galveston Bay and Tabbs Bay, Texas. Ph.D.

TEXAS WATERS

311

281. Hammerschmidt, P.C., 1974-The use of fish in cages as biological monitors of the

quality of water passing through a power plant. M.S.

282. Harry, David, 1976-Factors contributing to variation in productivity among Texas

reservoirs. Ph.D. (In progress)

283. Hodson, R.G., 1973-A comparison of occurrence and abundance of fishes within

three Texas reservoirs which receive heated water discharges. Ph.D.

284. Hutton, W.S., 1970-A quantitative and qualitative survey of benthal deposits con¬

tained in the Houston Ship Channel. M.S.

285. Helm, J.C., 1972-Methodology for systematic planning of regional water manage¬

ment. Ph.D.

286. Islam, M.A., 1972-The effect of temperature on reproduction of the red shiner

Notropis lutrensis (Baird and Girard). Ph.D.

287. Johnson, K.W., 197 3 -Occurrence and abundance of fishes in the intake and dis¬

charge areas of the Cedar Bayou power station before and during the first year
of plant operation. Ph.D.

288. Joyce, J.A., 1973-The effects of stocking density and feeding rate on the growth

of channel catfish Ictalurus punctatus (Rafinesque), in floating areas. M.S.

288a Jones, T.L., 1975 -Species composition, distribution and abundance of macro¬
zooplankton in the intake and discharge areas after construction and operation
of the Cedar Bayou electric power station. M.S.

288b Jackson, W.B., 1974-Distribution and abundance of shrimps, crabs, and fishes
in the cooling lake of the Cedar Bayou electric power station. M.S.

289. Kirk, William, 1976-The effect of varied Ca, Fe, Mg, and pH on growth and tax¬

onomic considerations for four desmid species. Ph.D. (In progress)

290. Klussmann, W.G., 1973-An evaluation of the utilization of anhydrous ammonia as

fisheries management technique in ponds. Ph.D.

291. Landry, Andre M., 1971 -Number of individuals and injury rates of economically

important fish passing through the P.H. Robinson generating station. M.S.

292. Lehmberg, Richard Verne, 1974-Nutrient limitation determinations for a Texas

watershed. Ph.D.

292a Linder, D.R., 1974-The culture of marine fish and their use as biological monitors
of water quality in ponds receiving heated discharge from a power plant. M.S.

293. Littleton, G., 1976-Growth characteristics of the Unionidae found in a stretch of

the Navasota River. M.S. (In progress)

294. Lock, J.T., 1971 -The effects of fertilization on the fishes of a Central Texas farm

pond. Ph.D.

295. Lukins, D., 1976-Ephontic algal productivity in an East Texas stream. Ph.D. (In

progress)

296. Malar, T., 1972-The locomotion and orientation of goldfish in a sound field. Ph.D.

297. Malouf, J.B., 1974-The development of a mathematical model to predict runoff

water quality from watersheds. Ph.D.

312

THE TEXAS JOURNAL OE SCIENCE

298. Maiquis, S., 1976-Survey of the Mallomonas (Chrysophyta) found in South Central

Texas. M.S. (In progress)

299. McBee, Jr., J.T., 1975 -Species composition, distribution, and abundance of macro-

benthic organisms in the intake and discharge areas before and after the con¬
struction and operation of the Cedar Bayou electric power station. Ph.D.

300. McCullough, Jr., J.D., 1970-Carbon fixation and community dynamics of a phyco-

periphyton community. Ph.D.

301. McCullough, M.M., 1971-A study of the juvenile fish fauna associated with the

cooling water of a steam electric generating station. M.S.

302. Medlen, Ammon Brown, 1952-Studies on the development of Gambusia affinis.

Ph.D.

303. Meyers, D.G., 197 3 -Comparative physical limnology of farm ponds in South Central

Texas. M.S.

304. Mitchell, T.M., 1970-The dispersion of dense effluent from an inclined jet discharg¬

ing into still fluid. M.S.

305. - , 1974- A parametric model for a dense plume near a stream bed. Ph.D.

306. Moore, J., 1973-The effect of water quality on chemical and physical properties

of a Hoban silty clay loam soil in situ. Ph.D.

307. Moskovito, G., 1961-Studies on diatom-bacteria interactions. Ph.D.

308. Moulton, B.A., 1976-Limnological study of a small watershed draining into the

Navasota River. Ph.D. (In progress)

309. Murrell, J.L., 1973-The intensive cage culture of channel catfish Ictalurus punctatus

(Rafinesque), in the intake and discharge canals of a steam electric generating
station, Trinidad, Texas. M.S.

310. Parmer, David Lee, 1970-Some mechanisms of organism limitations in the inland

Houston ship channel. M.S.

311. Petrocelli, S.M., 1973-The interactions of a chlorinated hydrocarbon insecticide

among the water, sediments and biota in an estuarine system. Ph.D.

312. Phantumvanit, D., 1974-Simulation and statistical quality control for water pol¬

lution abatement. Ph.D.

313. Phelps, R., 1976-Repopulation dynamics of the macrobenthos in a ripple area

of the Little Brazos River, Texas. Ph.D. (In progress)

314. Prather, J.E., 1976-The influence of thermal inflow on the distribution and con¬

dition of freshwater drum in Lake Trinidad, Texas. M.S. (In progress)

314a Quarberg, D.M., 1974-Report on the culture of brown shrimp, Penaeus aztecus, in
ponds receiving thermal effluent from a power plant. M.S.

315. Ramsey, D.B., 1973-The effects of a single application of fertilizer on the fish

population of Post Oak Lake. M.S.

316. Rand, G., 1976-Effect of pesticides on fish locomotion. M.S. (In progress)

TEXAS WATERS

313

317. Rennie, T.H., 1975-Zooplaknton studies in the Cox Bay, Texas, area before and

during early operation of an electric power plant. Ph.D.

318. Respess, Richard O., 1970-Seasonal variations in selected physical and chemical

conditions of a small impoundment in Brazos County, Texas. M.S.

319. Rozenburg, E.R., 1970-The composition and distribution of the fish fauna of the

Navasota River. M.S.

320. Rubee, Peter, 1976 -Behavioral studies in catfish. M.S. (In progress)

321. Runnels, W.C., 1969-Relationships between planktonic epiphytic and epipelic

diatom populations in selected ponds of Brazos County, Texas. Ph.D.

322. Reed, Joel R., 1972-Social and environmental characteristics of an urban river

recreation system. M.S.

323. Sarker, M.A.L., 1970-Patterns of feeding of the bluegill sunfish Lepomis macro-

chirus (Rafinesque), in two heated reservoirs of Texas. Ph.D.

324. Schwebel, M.D., 1973-Remote measures of turbidity and chlorophyll through

aerial photography. M.S.

325. Serns, S.L., 1972-Age, growth, and condition of bluegill sunfish Lepomis macro-

chirus (Rafinesque), in four heated reservoirs in Texas. M.S.

326. Strenth, N.E., 1974-A review of the systematics and zoogeography of the fresh¬

water species of Palamonetes heller (Crustacea, Decapoda) of North America.
Ph.D.

327. Smith, Francis William, 1973-A study of waterfowl habitats, populations and

fluctuations in the lower Trinity River and Upper Trinity Bay, Texas. M.S.

328. Schwebel, J.A., 1973-Hormonal control of growth and development in Lemna

minor L., with special emphasis on the role of abscisic acid (ABA). M.S.

329. Timms, A.M., 1972-Some aspects of intraspecific communication and of the role

of vision in the control and modulation of locomotor behavior of fish. Ph.D.

330. Wardle, W.J., 1974-A survey of the occurrence, distribution and incidence of

infection of helminth parasites of marine and estuarine Mollusca from Gal¬
veston, Texas. Ph.D.

331. Waxman, J.B., 1974-The locomotor behavior of fish in response to a subacute

concentration of copper alone and in combination with other environmental
factors. Ph.D.

332. Westlake, Garson F., 1973-Some effects of thyroxine on locomotor behavior in

fish. Ph.D.

333. Whitaker, R.E., 1974-Drag coefficient at hurricane wind speeds as deduced from

the numerical simulation of dynamic water level changes in Lake Okeechobee.
Ph.D.

333a Wiesepape, L.M., 1974-Thermal resistance and acclimation rate in young white
and brown shrimp, Penaeus setiferus Linn, and Penaeus aztecus Ives. Ph.D.

334. Wood, C.E., 1969-Relationship between primary productivity (C14 method) and

phytoplankton in a mesotrophic lake. Ph.D.

314

THE TEXAS JOURNAL OF SCIENCE

335. Yeh, C.F., 1971-Mark and recovery estimates of fish populations in three heated

reservoirs of Texas. M.S.

336. - , 1972-Population studies of selected fishes in three heated reservoirs in

Texas. Ph.D.

337. Zengerle, M.W., 1972-Age, growth, and condition of white crappie, Pomoxis an¬

nularis (Rafinesque) in Lake Nasworthy, Texas, a reservoir receiving a heated
effluent. M.S.

TEXAS CHRISTIAN UNIVERSITY

338. Brooks, A.W., 1928 -Freshwater Cladocera of North Texas. M.S.

339. Bryan, J., 1974-The distribution of benthic organisms in Nueces Bay, Texas, with

regard to the effects of thermal loading and hurricane Fern. M.S.

340. Cappel, S.M., 1970-A comparative study of the net plankton populations in the

tributaries and the main reservoir of Benbrook Lake, Tarrant County, Texas.
M.S.

341. Clavin, S., 1975 -The ichthyofauna of Benbrook Lake. M.S.

342. Coldiron, Donna Lee Roe, 1976-Biology: Some aspects of the biology of the

exotic mollusk Corbicula (Bivalvia: Corbiculidae). M.S.

343. Coolridge, J., 1975 -Acute toxicity of the herbicide Diquet to minnows.

344. Forsyth, J.W., 1937-Ecology of sewage filter bed with special reference to Psychoda

flies. M.S.

345. Froehlich, K., 1975 -Acute toxicity of organophosphorus insecticide, Malathion

to the fathead minnows ( Pimephales promelas) (Rafinesque) and golden shiners
( Notemigonus crysoleucas ) (Mitchell) M.S.

346. Green, D., 1968-Food and feeding habits of the white crappie Pomoxis annularis

(Rafinesque) in Benbrook Lake, Texas. M.S.

347. Gruninger, T., 1975 -Macroparasites of certain fishes of Eagle Mountain Lake. M.S.

348. Gunn, F.A., 195 3- A limnological investigation of a freshwater impoundment in

Wise County, Texas. M.S.

349. Hawley, J.B., 1926-A preliminary study of the microscopic life in Texas waters. M.S.

350. Holland, J.L., 1954-Geological investigations of Honey Creek watershed. M.S.

351. Jones, D.E., 1939— A limnological survey of Lake Como. M.S.

352. Kerby, H., 1966-Life history of largemouth bass , Micropterus salmoides, (Lacepede)

in Benbrook Lake, Tarrant County, Texas. M.S.

353. Lawrence, J.L., 1965 -Parasites of five fishes from Benbrook Lake, Tarrant County,

Texas. M.S.

354. Mahon, M.F., 1928-The Phylum Rothelminths in North Texas waters. M.S.

355. Mauldin, V.L., 1972-The bivalve Mollusca of selected Tarrant County, Texas,

reservoirs. M.S.

TEXAS WATERS

315

356. Moore, W.S.L., 1971 -Trace metal concentrations in water and sediments of the

Trinity River system, Tarrant County, Texas. M.S.

357. Paxton, J.E., 1973-A seasonal study of diurnal physico-chemical changes in a small

pond in Tarrant County, Texas. M.S.

358. Pettitt, J.M., 1973-Net plankton population of Eagle Mountain Lake, Tarrant

County, and Possum Kingdom Lake, Palo Pinto County, Texas. M.S.

359. Pruett, J., 1975 -The ichthyofauna of Eagle Mountain Lake. M.S.

360. Rose, D.L., 1941 -A limnological investigation of the Trinity River at Ft. Worth.

M.S.

361. Sale, C.M., 1957-Geology along the Clear Fork of the Trinity River southwest of

Ft. Worth, Texas, including Benbrook Lake. M.S.

362. Simmons, Barbara Hudson, 1976-Biology: Toxicity of selected industrial waste

effluents to channel catfish eggs and fry and largemouth bass fry as determined
through bioassay. M.S.

363. Taylor, R., 1975 -Water quality of Sycamore Creek, Ft. Worth, Texas: A non¬

point pollution source. M.S.

364. Ten Eyck, J., 1975 -Organic fractionization and selected trace metal content of

sludges. M.S.

TEXAS TECH UNIVERSITY

365. Ali, M.M., 1970-Unlined treated sewage storage ponds as sources of nitrate pol¬

lution of the Ogallala. M.S.

366. Anderson, J.F., 197 3 -Variation of urban runoff quality with duration and intensity

of storms. M.S.

367. Austin, T.A., 1971-A simulation model for the urban runoff process. Ph.D.

368. Brownlee, R.C., 1970 -Pollution of storm runoff from a small residential water¬

shed. M.S.

369. Carter, C.K., 1973-The effect of irrigation demand forecasting on the optimal

operation of the Elephant Butte-Caballo Reservoir system. M.S.

370. Chapman, S., 1974-Nitrogen mass balance determination for simulated wastewater

land-spreading operations. M.S.

371. Chitwood, J.R., 1965 -Hydraulics of groundwater flow. M.S.

372. Compton, J.L., 1975 -Diatoms of the Lubbock Lake site, Lubbock County, Texas.

M.S.

373. Cook, B.C., 1973-The effects of recreation pool size on irrigation and power gen¬

eration at Elephant Butte Reservoir. Ph.D.

374. Foerster, E.P., 1965 -A study of the effects of detergents on seepage rates. M.S.

375. Forehand, C.E., 1966-Fate of pesticides applied to lake beds and surrounding water

sheds. M.S.

316

THE TEXAS JOURNAL OF SCIENCE

376. Griffin, D.G., 1962-A population study of Chara zeylanica in Texas, Oklahoma

and New Mexico. M.S.

377. Hejl, Henry R., 1964-A study of the time of concentration for small natural water¬

sheds. M.S.

378. Henderson, G.T., 1961-Some factors affecting oospore germination in Chara zeylan¬

ica (Willdenow). M.S.

379. Keenum, BJ.B., 1975 -Comparisons of approximate F tests and a procedure for

the development of an alternative. M.S.

380. Knowles, T.R., 1971 -The development of a computer model for confined and

unconfined aquifers. M.S.

381. Knowles, T.R., 1972-The development of a computerized procedure to determine

aquifer parameters. Ph.D.

382. Lee, M.M., 1965 -Percolation of saltwater in an unconfined aquifer. M.S.

383. Lovell, T.L., 1969-A comparison of system designs and operations for the trans¬

mission of large volumes of water to West Texas. M.S.

384. Malone, C.R., 1965 -Dispersal of freshwater gastropods by water birds. M.S.

385. Mocek, M.J., 1971-A study of the compatibility of conjunctive use and Texas

groundwater law. M.S.

386. Pettit, G.M., 1974 -Evaluation of parameters for the production of water avail¬

ability from ungaged watersheds in Texas. M.S.

387. Rauschuber, D.G., 1972-A mathematical model to correlate rainfall and runoff.

M.S.

388. Roberts, D.W., 1970 -Mathematical model: A tool for water management. M.S.

389. Stephens, D.W., 1966-The economics of a constructed pit for groundwater re¬

charge on the high plains of Texas. M.S.

390. Thompson, G.B., 19 74 -Variation of urban runoff quality and quantity with dura¬

tion and intensity of storms. M.S.

391. Ward, C.R., 1964 -Ecological changes in modified playa lakes with special emphasis

on mosquito production. M.S.

392. Webster, R.D., 197 3 -Development and application of coaxial correlation tech¬

niques to predict runoff from small watersheds. M.S.

393. Williams, N.R., 1969-Population ecology of Natrix harteri. M.S.

394. Winn, W.T., 1973-A pilot study of the Canyon Lakes project. M.S.

395. Yeager, C.C., 1965 -A study of a salt-water wedge trapped in a freshwater aquifer.

M.S.

TRINITY UNIVERSITY

406. Albright, P.N., 195 2 -Contributions to the knowledge of the Odonata of Texas with
particular attention to the area surrounding San Antonio. M.S.

TEXAS WATERS

317

407. Giesler, R.S., 1972-A comparison of the benthic macroinvertebrates of Calaveras

and Braunig Lakes. M.S.

408. Smalley, H.E., 1956-The mosquito fauna (Diptera-Culicidae) of San Antonio,

Texas. M.S.

UNIVERSITY OF OKLAHOMA

409. Ausburn, B.E., 1961 -Subsurface disposal of natural brines in western Oklahoma and

northern Texas. M.S.

410. Baglin, Raymond E., Jr., 1976— Age and growth characteristics of white bass Morone

chrysops (Rafinesque) in Lake Texoma, Oklahoma. Ph.D.

411. Campbell, J.W., 1955 -A description of Lissorchis gullaris n. sp. (Trematoda: Lis-

sorchilidae) from the buffalo fish of Lake Texoma, Oklahoma. M.S.

412. Dowell, V.E., 1956-Activity patterns and distribution of the fishes in the Buncombe

Creek Arm of Lake Texoma, Oklahoma. Ph.D.

413. Echelle, A. A., 1966-The food of young-of-year gars, Lepisosteus, in Lake Texoma

with notes on spawning and development. M.S.

414. Grinstead, B.G., 1965 -The vertical distribution of the white crappie, Poxomis an¬

nularis, in the Buncombe Creek Arm of Lake Texoma. M.S.

415. Houser, A., 1958-A study of the commercial fishery of Lake Texoma. M.S.

416. Kilpatrick, E.B., 1959-Seasonal cycle in the gonads of the white bass ( Roccus

chrysops ), in Lake Texoma, Oklahoma. Ph.D.

417. McDaniel, J.S., 1961-A survey of the parasites of the genus Lepomis (Centrarchidae)

from the Buncombe Creek Arm of Lake Texoma. M.S.

418. Martin, J.M., 1952-Age and growth of the goldeye, Hiodon alosoides (Rafinesque)

of Lake Texoma, Oklahoma. M.S.

419. Mense, J.B., 1967-Ecology of the Mississippi silversides, Menidia audens Hay, in

Lake Texoma. M.S.

420. Moser, B.B., 1968-Food habits of the white bass in Lake Texoma with special

reference to the threadfin shad. M.S.

421. Riggins, M.A., 195 3- A study of the helminth parasites of certain shore birds from

Lake Texoma, Oklahoma. M.S.

422. Saunders, R.P., 1959-A study of the food of the Mississippi silversides Menidia

audens Hay, in Lake Texoma. M.S.

423. Shelton, W.L., 1972— Comparative reproductive biology of the gizzard shad,

Dorosoma cepedianum (Lesueur), and the threadfin shad, Dorosoma petenense
(Gunther), in Lake Texoma, Oklahoma. Ph.D.

424. Sublette, J.E., 1953-The ecology of the macroscopic bottom fauna in Lake Texoma.

Ph.D.

425. Szalwinski, A. A., 1967 -The highland lakes of Texas: a recreational study. M. A.

318

THE TEXAS JOURNAL OF SCIENCE

426. Taber, C., 1969-The distribution and identification of larval fishes in the Buncombe

Creek Arm of Lake Texoma with observations on spawning habits and relative
abundance. Ph.D.

427. Whiteside, B.G., 1964-Biology of the white crappie, Poxomis annularis, in Lake

Texoma. M.S.

428. Wilson, C., 1950— Age and growth of the white crappie Poxomis annularis (Rafin-

esque) in Lake Texoma, Oklahoma. M.S.

UNIVERSITY OF TEXAS AT ARLINGTON

429. Aldridge, D., 1975 -Biometrics and population dynamics of Corbicula manilenais

in Lake Arlington. M.S. (In progress)

430. Barnett, J., 1974-Littoral macrobenthos of Lake Arlington. M.S.

431. Bird, J.L., 1975 -Relations of zooplankton to water stratification due to density

differences resulting from thermal effluents. M.S. (In progress)

432. Carr, L., 1972-The effect of a thermal outfall on the phytoplankton of Lake Arling¬

ton. M.S.

433. Hall, F., 1972-Species diversity of the benthic macroinvertebrates inhabiting a

reservoir receiving a heated effluent. M.S.

434. Karbach, A., 1975 -Ecological survey of insular plants and vertebrates of Lake

Arlington. M.S. (In progress)

435. Kelly, M., 1975-Fishes of the headwaters of the Trinity River. M.S. (In progress)

436. Parker, F.R., 1971-Reduced metabolic rates in fishes as a result of induced school¬

ing. M.S.

437. Rutledge, C.J., 1975 -Fishes of Lake Arlington. M.S. (In progress)

438. Smith, M.D., 197 3 -Water quality survey ofBenbrook Lake. M.S.

439. Speegle, M., 197 3 -Limnology and ichthyology of Lake Fairfield, Texas. M.S.

440. Thompson, W., 197 2 -Potential storm water pollution from downtown Dallas. M.S.

441. Tommey, W.H., 1975 -Fishes of Monticello Reservoir. M.S. (In progress)

UNIVERSITY OF TEXAS AT AUSTIN

442. Aguirre, Jorge, 1967 -Nitrification and denitrification in a model waste stabilization

pond. M.S.

443. - , 1971 -Influence of the anaerobic sludge layer location on facultative

waste stabilization pond performance. Ph.D.

444. Archibald, Patricia A., 1969-A study of the algal flora of bogs with special reference

to the genus Chlorococcum. Ph.D.

445. Baker, Ailsie F., 1969-Taxonomic studies in the Oscillatoriaceae. Ph.D.

TEXAS WATERS

3.19

446. Barnhill, Betty Louise, 1970 -Aggressive interactions in reproductively active Cyprin-

odon variegatus. M.S.

447. Batterton, John Clyde, Jr., 1967-Studies of the phosphorus-deficient state in the

blue-green alga Anacystis nidulans. M.A.

448. - , 1970-A study of halotolerance in blue-green algae. Ph.D.

449. Bechtel, Timothy Joseph, 1970-Fish species diversity indices as pollution indicators

in Galveston Bay, Texas. M.S.

450. Beyers, Robert John, 1962-The metabolism of twelve aquatic laboratory micro¬

ecosystems. Ph.D.

451. Bhagat, Suriner Kumar, 1966-Transport of nitrosylruthenium in an aquatic environ¬

ment. Ph.D.

452. - , 1962-Estimating algal concentrations. M.S.

453. Birke, Lawrence Edward, Jr., 1968-Development of a blue-green algal assay for

vitamin B- 12 application to an ecological study of the San Antonio estuary. M.S.

454. Bischoff, Harry W., 1963-1. The soil algal flora of Enchanted Rock, II. Thorea

riekei sp. nov. and related species. Ph.D.

455. Bishop, Neil Erasmus, 1975— Studies on mixing and heat exchange in aerated lagoons.

Ph.D.

456. Blystone, Robert Vernon, 1968-The hemoglobin of Daphnia magna (Straus). M.S.

457. Bohmfalk, Clyde Edward, 1971 -Differentiation in temperate and tropical zone

populations of Typha under transplant garden and controlled photoperiod and
thermoperiod conditions. Ph.D.

458. Bolch, William Emmett, Jr., 1964-Behavior of ruthenium in algal environments.

M.S.

459. Brack enridge, Genie, 1968-The ultraviolet spectral analysis of coumarins. M.S.

460. Brock, David A., 1975 -Niche response structure of competition between Scene-

desmus obliquus and Chlorella vulgaris . M.S.

461. Bronaugh, Richmond Lee, 1950-Geology of Brazos River terraces in McLennan

County, Texas. M.A.

462. Brown, Richard Malcolm, Jr., 1964 -A comparative study of the algal genera Tetra-

cystis and Chlorococcum . Ph.D.

463. Brown, Billy Austin, 1975— Exchange of nitrogen and phosphorus with sediments

of the brackish water marshes of Lavaca Bay, Texas. M.S.

464. Buchan, Glenn Carl, 1972-A cybernetic approach to thermal pollution decision¬

making. Ph.D.

465. Bullock, Charles William, 1972— Measurement and prediction of abnormal reservoir

operations on Lake LBJ’s water quality. M.S.

466. Burch, Marvin Chandler, 1950-A history of the lower Trinity River region to 1836.

M.S.

320

THE TEXAS JOURNAL OE SCIENCE

467. Cain, Brother Joseph R., 1963-The morphology, taxonomy and physiology of

certain Chlamydomonas-like algae. Ph.D.

468. Calhoun, Sam Harlan, 1970-An investigation of the oxygen balance in the Colorado

River, Lake Austin, Texas. M.S.

469. Carlisle, Clara, 1924-The rate of disappearance of chlorine in drinking water. M.A.

470. Carr, Roseanne, 1972-Eield and culture studies of the diatom genus Biddulphia

in the Gulf of Mexico. M.S.

471. Carroll, William Erwin, Jr., 1972-Hypolimnetic dissolved oxygen simulation model.

M.S.

472. Chambers, John Edward, 1962-Development of a polarographic system for the

measurement of oxygen exhange in algal suspensions. M.S.

473. Chavez, Pedro Islas, 1973-Variation in sesquiterpene lactones within a Trinity

River, Texas, population of Xanthium strumarium. M.S.

474. Cech, Joseph Jerome, Jr., 1970 -Respiratory responses of the striped mullet, Mugil

cephalus, to three environmental stresses. M.S.

475. Chen, Tony Gwokwong, 1972-Selective withdrawal at Lake Livingston. M.S.

476. Clark, Thomas Phillips, 1972-Hydrology, geochemistry, and public health aspects

of environmental impairment at an abandoned landfill near Austin, Texas. M.S.

477. Clark, George Gregory, 1973-Evaluation of loading on water quality attributable

to non-point sources. M.S.

478. Cobb, Howell Dee, 1963-Studies on nitrogen fixation in a blue-green alga. Ph.D.

479. Collier, Robert Gressett, 1939-A survey of the water fauna of Houston and its

vicinity and an analytical study of its use in teaching biology. M.Ed.

480. Cooper, B.D., 1954-The feeding habits of the largemouth bass (Micropterus sal-

moides). M.S.

481. Cooper, David Clayton, 1970-Responses of continuous-series estuarine micro¬

ecosystems to point source input variations. Ph.D.

482. Cox, E.R., 1966-Taxonoinic, morphological and physiological studies of the algal

genus Stigeoclonium. Ph.D.

483. Cramer, Marian Lenore, 1952-Studies on growth and photosynthesis of Euglena

with emphasis on the replacement of CO2 in metabolism. Ph.D.

484. Crawley, Richard Alvin, 1969-Flood effects on Sanderson Creek, Trans-Pecos

Texas, June 11, 1965. M.S.

485. Crow, Larry Isaac, 1974-A numerical study of the velocity and temperature fields

in a stratified reservoir. M.S.

486. Crowley, Dennis John, 1971 -Evaluating the effects of return flows on stream

quality in terms of total dissolved solids. M.S.

487. Cumming, Robert Bruce, 1964 -Cytogenetic studies in the order Odonata. Ph.D.

488. Darnell, Paul Edward, 1972-Upflow filtration of waste stabilization pond effluent.

M.S.

TEXAS WATERS

321

489. Davenport, James B., 1975 -Phytoplankton succession and the productivity of

individual algal species in a Central Texas reservoir. M.S.

490. Davidson, F.F., 1941 -Taxonomy of Texas Zygnemataceae. Ph.D.

491. Dawson, Anita Joan, 1976-The role of plants in nutrient exchange in the Lavaca

Bay brackish marsh system. M.S.

492. Deason, Temd Robert, 1960-Exploratory studies of Texas soil algae. Ph.D.

493. Delco, A., 1962-Reproductive behavior and sexual isolation mechanisms between

two sympatric cyprinid fishes. M.S.

494. Dharmikarak, Sawasdi, 195 7 -Coagulation of Lake Austin water. M.S.

495. Diehl, W.H.S., 1959-Peak discharge predictions for Waller Creek at 23rd Street

at Austin, Texas. M.S.

496. Drewry, George Earl, 1962-Some observations of courtship behavior and sound

production in five species of Fundulus. M.S.

497. - , 1967 -Studies of relationships within the family Cyprinodontidae. Ph.D.

498. Drynan, Walter Ronald, 1956-Methods of concentrating algae. M.S.

499 - , 1961 -Radioactivity in surface waters: a statistical approach to establish¬

ing base levels. Ph.D.

500. Duke, M.E.L., 1967-A production study of a thermal spring. Ph.D.

501. Dykstra, Richard F., 1966-An investigation of some algae of the Texas Gulf Coast.

M.A.

502. de la O, Ernesto Espino, 1968-Sulfide production in waste stabilization ponds. Ph.D.

503. Eddleman, Charles Douglas, 1965 -A morphological study of the early instars of

mosquito larvae. M.S.

504. Edwards, Linda Kay, 1969-Some non-aquatic epiphytic and lithophilous algae. M.S.

505. Edwards, Peter, 1969-Field and cultural studies of the seasonal periodicity of

growth and reproduction of selected Texas marine algae. Ph.D.

506. Eley, James Henry, Jr., 1967-Enhancement of photosynthesis in Chlorella and

a kinetic model for photosynthesis. Ph.D.

507. Elias, Robert William, 197 3 -The role of seagrasses and benthic algae in the bio¬

geochemistry of trace metals in Texas estuaries. Ph.D.

508. Espey, William Howard, Jr., 1965 -A study of some effects of urbanization on storm

runoff from a small watershed. Ph.D.

509. Felsing, William August, Jr., 1962-Sr90 and Ra226 in a domestic water supply. M.S.

510. Fisher, Charles Parker, 1963 -Stabilization of biosorption activated sludge in waste

stabilization ponds. Ph.D.

511. Floyd, Bert Allen, 1969 -The use of algal cultures to assess the effects of nutrient

enrichment on the Highland Lakes of the Colorado River, Texas. M.S.

512. Fogg, John Brian James, 1973-A dissolved oxygen model for the hypolimnion of

Lake LBJ. M.S.

322

THE TEXAS JOURNAL OF SCIENCE

513. Ford, Davis Lee, 1965 -Kinetics of aerobic oxidation in the thermophilic range. M.S.

514. Garner, Lee Edwin, 197 3 -Environmental geology of the Austin area, Texas. M.S.

515. Gazda, Lawrence Paul, 1969-Land disposal of municipal solid wastes in selected

standard metropolitan statistical areas in Texas. M.S.

516. Gearing, Patrick James, 1970-A study of the effect of TC-99 Pertechnetate on

blue-green algae. M.A.

517. Gebhard, Thomas Granville, Jr., 1964-Streamflow frequency distributions on se¬

lected streams in Texas. M.S.

518. Gerald, Jerry Wayne, 1970-Species isolating mechanisms in the genus Lepomis. Ph.D.

519. Govin, Charles Thomas, Jr., 197 3 -Sedimentation survey, Lake Buchanan, Texas,

1973. M.S.

520. Gravel, Alan Charles, 1969-The distribution of coliform bacteria in stratified im¬

poundments. M.S.

521. Green, George Edward, 1975 -An assessment of phosphorus data collected from

Lake Lyndon B. Johnson. M.S.

522. Groover, Robert Don, 1968 -Experimental studies on Chlorosphaeracean and other

edaphic algae. Ph.D.

523. Guaqueta, Camilo Hernando, 197 3 -Environmental and economic impact of pol¬

lution abatement by SIC 28 and 29. M.S.

524. Hagen, Donald Warren, 1963-Some aspects of thermal tolerance in three species of

Gambusia. M.S.

525. Harris, James Carl, 1971 -Pollution characteristics of channel catfish culture. M.S.

526. Haughton, Edward Ronald, 1970-Oxygen depletion and prediction in the hypo-

limnion of a southwestern reservoir. M.S.

527. Henderson, G.G., 1960-Stomach contents of Central Texas specimens of Percina

caprodes. M.S.

528. Herman, Edward Robert, 195 7 -Development of design criteria for waste stabiliza¬

tion ponds. Ph.D.

529. Herman, William Hoffman, Jr., 1976-A continuous equation to describe bioassay

results for acute exposure of fish to toxins. M.S.

530. Higgins, Robert Brown, 1969 -Enzymatic method for the detection of luxury phos¬

phorus uptake. M.S.

531. Hoffman, Larry R., 1961-Studies on the morphology, cytology and reproduction

of Oedogonium and Oedocladium. Ph.D.

532. Hu, Vicky Ray-Shen, 1971-Nutrient studies in Texas impoundments. M.S.

533. Huang, George S., 1968— Scheduling of releases to meet downstream irrigation

requirements. M.S.

534. Huang, Ju-Chang, 1967 -Effects of toxic organics on photosynthesis reoxygen¬

ation. Ph.D.

TEXAS WATERS

323

535. Hung, Yung-Tse, 1970-Biological reaction rate constant in natural water. Ph.D.

536. Ingram, Lonnie O’Neal, 1971-Mutations and cell division in blue-green algae. Ph.D.

537. James, Stephen Nelson, 1974- An analysis of factors affecting the distribution of

nekton in and around a power generating station in an estuarine area. M.S.

538. Jetton, Elden Verner, 1973-Climatology of the upper Rio Grande Basin and the

development of spring runoff forecast equations. Ph.D.

539. Jones, Larry Warner, 1964-Photosynthetic enhancement phenomena in a blue-

green alga, Anacystis nidulans. Ph.D.

540. Jurgens, K.C., 195 1 -The distribution and ecology of the fishes of the San Marcos

River (Austin, Texas). M.S.

541. Kantz, Paul Thomas, Jr., 1966-Some South-Texas soil algae with special reference to

the genus Nos toe. M.A.

542. - -, 1968-Taxonomic, morphological, and physiological studies of the algal

genera Nostoc and Anabaena. Ph.D.

543. Kapraun, Donald Frederick, 1969-Field and cultural studies of the genera Ulva

and Enteromorpha . Ph.D.

544. Kemp, Robert James, Jr., 1950-A comparative study of brackish water fishes. M.S.

545. Kikuchi, Noboru, 1975 -A study of variational inequalities with a class of seepage

flow problems. M.S.

546. Killam, Allen Page, 1955 -Special effects of light on Gilorella. M.S.

547. Koch, Charles Thomas, 1966-Organic constituents in surface waters. M.S.

548. Kratz, William A., 1954-Studies on the physiology of blue-green algae. Ph.D.

549. Kreitler, Charles Wrightman, 19 74 -Determining the source of nitrate in groundwater

by nitrogen isotope studies. Ph.D.

550. Krise, George Martin, Jr., 1952-A study of nitrogen excretion of ciliate protozoa.

Ph.D.

552. Ledbetter, Joe Overton, 1963-Predictive techniques for water quality-inorganics.

Ph.D.

553. Lee, Chin-Yuan, 1968 -Macro- turbulence in wind waves. Ph.D.

554. Lennington, Richard Kent, 1975 -The analysis of clustering methods as applied

to biological population data. Ph.D.

555. Lindstrom, Bruce M., 1975-An analysis of niche response of Lake LBJ phyto¬

plankton to environmental levels of light and temperature. M.S.

556. La Montagne, John Ring, 1967 -A procedure for the isolation of heterocysts from

blue-green alga Anabaena cylindrica (Fogg). M.S.

557. Laplante, Joseph Michael, 1975 -Investigation of uptake characteristics of various

clays for selected heavy metals. M.S.

558. Lytle, Thomas Foster, 1966-The biogeochemistry of trace metals in the near shore

environment. Ph.D.

324

THE TEXAS JOURNAL OF SCIENCE

559. MacRae, John R., 1975 -Some effects of temperature on the population dynamics

of planktonic rotifers. M.S.

560. Martin, Floyd Douglas, 1967 -Osmotic shock survival in certain cyprinodont fishes.

M.S.

561. - , 1968 -Some factors influencing penetration into rivers by fishes of the

genus Cyprinodon. Ph.D.

562. Mattox, Karl R., 1962-The taxonomy of certain ulotrichalean algae. Ph.D.

563. Matusi, Saburo, 1972 -Biological accumulation of mercuric and methyl mercury

by Chlorella pyrenoidosa. Ph.D.

564. McCarver, C.T., 1950-The effects of pollution by water purification sludge on

freshwater fishes (Austin, Texas). M.S.

565. McClure, Jerry Weldon, 1964-Taxonomic significance of the flavonoid chemistry

and the morphology of Lemnaceae in axenic culture. Ph.D.

566. McClure, John Robert, 1962-Analysis of the stationary and death phases of the

growth curve of Tetrahymena pyriformis. M.S.

567. McGowen, Joseph Hobbs, 1969-Gum Hollow Fan Delta. Nueces Bay Texas: mode

of development and sedimentation characteristics. Ph.D.

568. McLaurin, Banks, 1928-The silt problem in Texas. M.S.

569. McHale, John Thomas, 1965— The reduction of heavy metal salts by chloroplasts,

with reference to ascorbic acid. Ph.D.

570. McNaughton, Samuel Joseph, 1 964 -Ecotypic patterns in Typha and their significance

in ecosystem integration. Ph.D.

571. Melbard, Albert Roy, 19 74 -Inactivation of virus in bench-scale oxygenerated waste

stabilization ponds. M.S.

572. Meier, W.L., 1964-Analysis of unit hydrographs for small watersheds in Texas. M.S.

573. Milliger, Larry Edward, 1965-Some soil algae from Bastrop State Park, Texas. M.S.

574. Mills, Jimmy Tom, 1972-The biology of two species of haptophycean algae (Chryso-

phyta). Ph.D.

575. Mims, Charles Wayne, 1969-An ultrastructural study of the life cycle of the Myxo-

mycete Arcyria cinerea. Ph.D.

576. Moore, Richard Byron, 1966-Survival studies on the blue-green alga, Anacystis

nidulans. Ph.D.

577. Moore, Harry Grady, Jr., 1969-Studies on the surplus phosphorus uptake phenome¬

non in algae. Ph.D.

578. Moran, Robert Edward, 1974-Trace element content of a stream affected by metal-

mine drainage, Bonanza, Colorado. Ph.D.

579. Nations, M.A., 1959-Multiple correlation of rainfall and other factors to runoff. M.S.

580. Neill, William Edward, 1972-Effects of size -selective predation on community

structure in laboratory aquatic microcosms. Ph.D.

TEXAS WATERS

325

581. Nelson, Robert Young, 1964-Atmospheric emissions from waste stabilization

ponds. Ph.D.

582. Nokes, Mark Alan, 1972-Effects of x-irradiation on photosynthesis in Chlorella

pyrenoidosa . M.S.

583. Palmer, Neil Meredith, 1949 -An approach to the ecology of the lower Rio Grande

Valley. M.S.

584. Parker, Bruce C., 1960 -Environmental investigations of the relationships of certain

soil algae with their associated microorganisms. Ph.D.

585. Patyrak, Richard Clark, 197 2 -The distribution of DDT in a laboratory soil- water

system. M.S.

586. Pearson, G.P., 1955 -Water resources of Texas (Austin, Texas), M.S.

587. Pearson, R.G., 1938-The aquatic oligochaeta of the Austin region. M.S.

588. Penumalli, Bhaskara Reddy, 1975 -Large systems approach to water quality model¬

ing and management. Ph.D.

589. Pessoney, George Francis, 1968-Field and laboratory investigations of Zygnemat-

aceous algae. Ph.D.

590. Phillips, Jesse Neal, Jr., 1953-The growth rate of Chlorella pyrenoidosa as a function

of intensity and intermittency of illumination. Ph.D.

591. Pittman, Dwight Lee, 1969-Determination of the effects of shoreline development

on impoundment water quality. M.S.

592. Pocock, Mary Avis, 1966 -Chemical discrimination in Procambarus clarkii: behav¬

ioral responses to electrolytes.

593. Poole, John Alex, 19 70 -A study of the death rate of Escherichia coli in natural

waters. M.S.

594. Proffitt, Thomas Oliver, 1971 -Variations in daily solar radiation at National Weather

Service observing sites in Texas. M.S.

595. Pursley, Lewis Ferguson, 1970-Impoundment model for predicting water quality

changes with depth in a stratified reservoir. M.S.

596. Raul, Cuellar- Chavez, 1976-Regional methodology to estimate point and non¬

point pollutional loads discharged into a stream.

597. Redford, Edward Logan, 1950-Some effects of ultraviolet radiation on Chlorella.

M.S.

598. Reimers, Robert Stolt, 1968 -A stable carbon isotope study of a marine bay and

domestic sewage plant. M.S.

599. Renfro, W.C., 1958-The effect of salinity on the distribution of fishes in the Ark¬

ansas River. M.S.

600. Ruane, Richard James, 1970-Effects of environmental development on water

quality and variation of water quality with depth and location-lower Highland
Lakes. M.S.

601. Ruppersberger, John Sidney, 1971-Studies on nutrients released from a Trinity

River sediment under oxic conditions. M.S.

326

THE TEXAS JOURNAL OF SCIENCE

602. Samejima, Hirotoshi, 1957-Heterotrophic metabolism of unicellular green algae.

M.S.

603. Sauer, S.P., 1963-Multiple correlation estimates of runoff as affected by areal

distribution of rainfall. M.S.

604. Sealey, Jesse Q., 1951 -The biology of Clostridium nigrificans. Ph.D.

605. Sell, John Leroy, 1960-Study of effects of low temperature on Anacystis nidulans.

606. Sever, Julia Rebecca, 1968 -The geochemistry of fatty alcohols in recent and ancient

sediments. M.S.

607. Shen, Eugene Y.F., 1966-Morphogenetic and cytological investigations of Chara

contraria and C. zeylanica. Ph.D.

608. Shull, Roger Don, 19 68 -Radioactivity transport in water; simulation of sustained

releases to selected river environments. Ph.D.

609. Smith, Michael Alexis, 1975 -Geology and trace element geochemistry of the Fort

Davis area, Trans- Pecos, Texas. Ph.D.

610. Smith, Cecil Hiawatha, 1965 -The synthesis of storm runoff hydrographs from

unsteady, non-uniform rainfall. Ph.D.

611. Smith, Beulan Sands, 1938-A study of the algae of the Gulf Coast of Texas. M.A.

612. Smith, R.L., 1965 -Morphological, taxonomic and physiological investigations of

the algal genera Eremosphaera and Oocystis. M.S.

613. Sorensen, Elsie Mae Boecker, 1974-Thermal effects on the biological magnification

of arsenic in green sunfish, Lepomis cyanellus. Ph.D.

614. Sorokin, Constantine Alexis, 1955 -Studies on high-temperature algae. Ph.D.

615. Steed, David Lewis, 1967 -Metabolic responses of some estuarine organisms to an

industrial effluent. M.S.

616. Stern, Milton Harold, 1956-A study of generation time in Tetrahymena pyriformis.

M.S.

617. Stewart, Jeanette Louise Kershaw, 1971 -The biology of the green alga Characio-

siphon rivularis (Iyengar). Ph.D.

618. Strawn, R.K., 1957-The influence of environment on the meristic counts of the

fishes, Etheostoma grahami and E. lepidum. Ph.D.

619. Stuessy, Tod Falor, 1968- A systematic study of the genus Melampodium (Com-

positae-Heliantheae). Ph.D.

620. Sun, Elaina Si-chen Liu, 1966-Studies of East Texas soil algae with special reference

to Botrydium. M.S.

621. Suwannakarn, Verachai, 1963-Temperature effects on waste stabilization pond

treatment.

622. Thirumurthi, Dhandapani, 1966-Relative toxicity of organics to Chlorella pyrenoi-

dosa. Ph.D.

623. Thomas, Dempsey L., 1968-Morphological and biochemical taxonomy of the

algal genus Protosiphon. Ph.D.

TEXAS WATERS

327

624. Tilton, John E., 1961 -Ichthyological survey of the Colorado River of Texas. M.S.

625. Tischler, Lial Frederick, 1966-A mathematical study of the kinetics of biological

oxidation. M.S.

626. Tomme, Michael Henry, 1974-Algal bioassays applied to Central Texas waters.

M.S.

627. Tribbey, Bert Allen, 1965-A field and laboratory study of ecological succession

in temporary ponds. Ph.D.

628. Trevino, D.B., 1955 -The ichthyofauna of the lower Rio Grande River, from the

mouth of the Pecos to the Gulf of Mexico. M.S.

629. Tu, Catharina Tse-Ying Yang, 1971 -The study of a phosphate- containing fluore¬

scent compound from spinach chloroplasts. M.S.

630. Uhlik, David James, 1969 -Studies in Chlamydomonas . M.A.

631. Urban, L.V., 1966-Estimates of physical exchange in Galveston Bay, Texas. M.S.

632. - , 1971 -A laboratory investigation of temperature and velocity fields

in a stratified cooling reservoir. Ph.D.

633. Valencia-Montoya, Guillermo, 1970-Oxygen consumption by benthic deposits.

M.S.

634. Van Dover, Byron, 1972-Studies of the ultrastructure and reproduction of several

species of Chlamydomonas and Catena. Ph.D.

635. Volkmann, Carol Mary, 1960-The microbial decomposition of organic carbon in

surface sediments. M.S.

636. Voswinkel, Linda Louise, 1969-Taxonomy and ultrastructure of the algal genus

Pyrobotrys. M.S.

637. Wadsworth, Albert Hodges, 1941 -The lower Colorado River, Texas. M.A.

638. Wallace, James William, Jr., 1967 -Investigations of flavone biosynthesis in the

Lemnaceae. Ph.D.

639. Walz, David Henry, 1974-Sewage renovation and surface-water quality, Lakeway

Resort Community , Travis County, Texas. M.S.

640. Watkins, G.M., 19 30 -Vegetation of San Marcos Springs. M.S.

641. Watson, Melvin W., 1970-Ultrastructural aspects of development in Microthamnion .

Ph.D.

642. Weber, Gerald Eric, 1968-Geology of the fluvial deposits of the Colorado River

Valley, Central Texas. M.S.

643. Wells, Dan Moody, 1966-Management of return flows in Texas. Ph.D.

644. Wenzel, Roger Allen, 1975 -Effects of floods on selected sand bars, San Bernard

and Brazos Rivers, Austin County, Texas. M.S.

645. White, Dale Wesley, 1974-Survey of pollution characteristics of intensely cultured

fish. M.S.

646. Wiedeman, Varley Earl, 1964-Some aspects of algal ecology in a waste-stabilization

pond system. Ph.D.

328

THE TEXAS JOURNAL OF SCIENCE

647. Wiland, Bruce Lane, 1975 -Transient state biological kinetics. M.S.

648. Wilkins, Lowell H., 1973-Wave spectra and associated statistical characteristics

of free-surface waves on a freshwater lake. Ph.D.

649. Williams, George Jackson, III, 1969-Comparative phenology, physiology, and

biochemistry of populations of Liquidambar styraciflaa L. Ph.D.

650. Williams, John Gregor, 1974-A comparison of development rates and temperature

tolerances of two species of Cyprinodont fishes (Family Cyprinodontidae):
Cyprinodon variegatus and Cyprinodon bovinus. M.S.

651. Wilson, Joe Robert, 1967-Field investigation of mixing and dispersion in a deep

reservoir. Ph.D.

652. Woodruff, Charles Marsh, Jr., 197 3- Land-use limitations related to geology in the

Lake Travis vicinity, Travis and Burnet Counties, Texas. Ph.D.

65 3. Wu, Madeline Chang Sun Wang, 1966-Pentose inhibition of the growth of Chloro-
coccum echinozygotum. Ph.D.

654. Yang, Julie Tsu-yu, 1968- A study of selected isoenzymes in Lemnaceae. M.S.

655. Yantis, Hugh Cleveland, Jr., 1965 -Water quality criteria for Brushy Creek inWilliamson

County, Texas. M.S.

656. Yardley, Darrell Gene, 1972-An ecological and electrophoretic analysis of the inter¬

action between Gambusia a f finis and G. heterochir. M.S.

657. Yarish, Charles, 197 2 -Experimental taxonomic studies of certain marine chaeto-

phoracean algae. M.A.

658. Young, W.C., 1960-Ecological studies of commensal Branchiobdellidae (Oligochaeta)

and Entocytheridae (Ostracods) on Procambarus simulans simulans (FAXON)
(Decapoda). Ph.D.

659. Young, Donald Ray, 1972-The relationship of land use to water use in San Antonio,

Texas. M.S.

660. Young, Frederick LeRoy, Jr., 1965 -Vacuum filtration of algae. M.S.

661. Young, Joncie Harvey, 1974-Facultative waste stabilization pond processes, per¬

formance, design, and predictive models. M.S.

662. Yousef, Ariz Yousef, 1962-Coliform in waste stabilization ponds. M.S.

UNIVERSITY OF TEXAS AT DALLAS

663. Gary, M., 1976-Factors controlling the eutrophication of Lake Ray Hubbard.

Ph.D. (In progress)

664. Meyers, M., 1976-An evaluation of the factors influencing water quality of White

Rock Lake. M.S. (In progress)

665. Rast, Walter, 1976-An evaluation of the relationship between aquatic plant nutrient

loads and the degree of fertilization of lakes and impoundments. Ph.D. (In prog¬
ress)

TEXAS WATERS

329

UNIVERSITY OF TEXAS AT EL PASO

666. Ferguson, J.A., 1961-Water for international rivers; the Rio Grande as a model. M.S.

667. Haney, P.L., 1948 -The international controversy over the waters of the Rio Grande.

M.A.

668. Kirby, J.W., 1968-Water resources, El Paso County, Texas: past, present, future.

M.A.

669. Nicoll, M.C., 1951-Brief history of the El Paso water system from 1881-1921. M.A.

670. Sedillo, F.J., 1969 -Investigations on the recharge to groundwater by the sewage

effluent of WSMR, New Mexico. M.S.

671. Stachowiak, A.S., 1969-Groundwater resources of the river alluvium. M.S.

672. White, Alice M., 1950-History of the development of irrigation in El Paso Valley.

M.A.

UNIVERSITY OF HOUSTON

673. Powell, R.W., 195 3-Diatoms of Bray’s Bayou, Houston, Texas. M.S.

WEST TEXAS STATE UNIVERSITY

674. Anderson, G., 1975 -Bacterial quality of Lake Meredith. M.S. (In progress)

675. Canada, W.B., 1975 -Water quality of Lake MacKenzie, M.S. (In progress)

676. King, E.D., 1969-Coliform bacteria in Lake Meredith, Texas. M.S.

677. Hestand, R.S., III, 1966-Quality of water in Lake Meredith, Texas. M.S.

678. Haralson, J.E., 1972-The problem of providing a dependable quality municipal

water supply for the City of Winfield, Kansas. M.S.

679. Jones, O.R., 1970-The effect of net plankton on the recharge rate of wells and

spreading basins. M.S.

680. Lutz, J.D., 1970-Age, growth, and condition of Northern Pike ( Esox lucius ) in

Greenbelt Lake in the Texas Panhandle. M.S.

681. Meriage, M., 1975 -Checklist on amphibians and reptiles of Randall County. M.S.

(In progress)

682. Newton, C.E., 1968-Phytoplankton succession and seasonal periodicity in a ma¬

turing lake. M.S.

683. Whitney , W.E., 1967-Canadian River dam project in Texas. M.S.

1976 ADDENDUM
ANGELO STATE UNIVERSITY

Fischer, Lyndal, 1976-The Caddisflies of the Concho River System. M.S. (In prog¬
ress)

684.

330 THE TEXAS JOURNAL OF SCIENCE

BAYLOR UNIVERSITY

685. Jacobs, Robert, 197 6 -The combined relationship of temperature and molybdenum

concentration to nitrogen fixation by Anabaena cylindrica. M.S.

686. Neely, Robert, 197 6 -The contribution of aquatic macrophytes to primary product¬

ivity in two Texas reservoirs. M.S. (In progress)

687. Short, James, 1976-Effects of dissolved organics on nutrient availability. M.S.

(In progress)

688. Muschler, Lee, 1976-The phosphorus budget of two Texas reservoirs. M.S. (In

progress)

LAMAR UNIVERSITY

689. Griffith, Wliiam H., 1975 -Acute toxicity of copper and lead to the blue crab,

Callinectes sapidus Rathbun. M.S.

690. Linney, George K., 1975 -Minimum temperature tolerance of Rhizophora mangle

saplings in culture. M.S.

SAM HOUSTON STATE UNIVERSITY

691. Hand, S.S., 1976-A comparison of three methods of establishing primary product¬

ivity of surface water of Lake Conroe. M.S.

PAN AMERICAN UNIVERSITY

692. Atkinson, Eduardo, 1976-Comparative hematology of the Rio Grande cichlid,

Cichlasoma cyanoguttatum, and the redear sunfish, Lepomis microlophus. M.S.

STEPHEN F. AUSTIN UNIVERSITY

693. Allen, Charles E., 1976-Bioeconomics and feeding habits of ducks in flooded bot¬

tomlands of Eastern Texas. M.S.

694. Lindsey, Larry R., 1976-An analysis of coliform and streptococcus bacteria as

indicators of fecal pollution in the Attoyac River. M.S.

695. Richardson, Larry J., 197 6 -Analysis of the coliform and streptococcus bacteria

as indicators of fecal pollution in the Angelina River. M.S.

TEXAS A&M UNIVERSITY

696. Grosmaire, Eric G., 1976-Food analysis and population dynamics of fresh water

turtles in the Rio Grande Valley of south Texas. M.S.

697. Hays, Bill, 1976-The comparative ethology of various genera of Texas crayfish. Ph.D.

TEXAS WATERS

331

UNIVERSITY OF HOUSTON

698. Carr, Barbara Ann, 1972™ Assumption of L- valine and D-glucose by the mid-gut

gland of the shrimp .Penaeus aztecus. M.S.

699. Castille, Frank, 197 6 -The uptake of hexose from seawater by post-larval Penaeid

shrimp. M.S.

UNIVERSITY OF TEXAS AT DALLAS

700. Foute, Steve, 1976-Land usage patterns within the Lake Ray Hubbard, Texas,

watershed. Ph.D. (In progress)

701. Meckel, Elaine, 197 6 -Biological responses of Lake Ray Hubbard, Texas, to phos¬

phorus and nitrogen loadings. M.S. (In progress)

702. Moore, Mike, 197 6 -Sedimentation patterns as they relate to the phosphorus and

nitrogen budgets of Lake Ray Hubbard, Texas. M.S. (In progress)

703. Rahman, Magda, 197 6 -Development of a eutrophication model for Lake Ray

Hubbard, Texas. Ph.D. (In progress)

THE UNIVERSITY OF TEXAS

SCHOOL OF PUBLIC HEALTH AT HOUSTON

THESES AND PROJECTS COMPLETED

704. Abd-Eldayen, Mohamed Sabry, 1973-A feasibility study of the non- chlorination

methods for potable water disinfection - - (with reference to proposed disinfec¬
tion of the Holy Water of Zamzam well in Mecca). M.P.H.

705. Baird, Justus N., 197 3 -Salmonella, turtles, and public health. M.P.H.

706. Bean, Judy A., 1973-Behavior of replication and linearization variance estimators

for complex multistage probability samples. Ph.D.

707. Casserly, Dennis M., 1976-Indicator and pathogenic bacteria dynamics of urban

and rural stormwater runoff. M.S.

708. DiPietro, Walter P., 1975 -Bacteriological investigation of private well water sup¬

plies in Acres Homes (Houston). M.S.

709. Dunn, William F., 1971 -Nutrient Flux in the San Antonio Bay complex as measured

by water column analysis. M.S.

710. Fosbury, Walter Jay, 1972-The use of aerial remote sensing for the detection of

specific water pollution parameters. M.S.

711. Gomez, Sabino, 1971 -The significance of viruses in water. M.P.H.

712. Gothard, Clair S., 1974— Bacteriological analysis of surface water by the technique

of a continuous monitoring system. M.P.H.

713. Gray, William G., 1973- Ammonia levels in Texas streams receiving municipal waste-

water effluents during periods of low flow. M.P.H.

332

THE TEXAS JOURNAL OF SCIENCE

714. Hall, Robert C., 1970-Study of sanitary landfills as a potential health hazard. M.P.H.

715. Henderson, Marilyn, 1976— Some factors affecting the enumeration of viruses iso¬

lated from water and wastewater. M.S.

716. Hulka, Steven C., 1972-The occurrence of indicator organisms in two Texas est¬

uaries. M.S.

717. Jackson, Gaines B., 1972-A treatise on the community health aspects of water

quality in Chocolate Bayou. M.S.

718. Keen, Spruce R., 1972-Postchlorination bacterial survival and aftergrowth in sec¬

ondary treated wastewater. M.S.

719. Langdon, William C., 1972-Primary productivity and nutrient budget of San An¬

tonio Bay - a Texas estuary. M.S.

720. Moore, James D., 1975 -Multiple regression analysis of bacterial populations in a

natural stream. M.S.

721. Reeve, Gordon R., 1975 -Cholinesterase levels of aerial applicators exposed to

organophosphate pesticides. M.P.H.

722. Simpson, Diane M., 1974-Bacteria in treated domestic sewage used for irrigation:

identification of the enterobacteriaceae emitted from a sprinkler and observa¬
tions of the coliform concentrations present within the system. M.P.H.

723. Smith, George Franklin, 1973-A preliminary investigation into the nutrient re¬

quirements of four Texas estuarine systems. M.S.

724. Smith, Billy R., 1972-A comparison of two bacteriological analysis techniques

in brackish waters. M.P.H.

725. Tapp, Mark, 1975— An evaluation of the eutrophication rate and some health re¬

lated factors in the Clear Lake Basin (Texas). M.S.

726. Valentine, Jane L., 197 3 -Distribution of trace elements in water in the Houston

environment: relationship to mortality from arteriosclerosis, heart disease. Ph.D.

727. Weese, Thomas H., 1973-The study of the possible effects of salinity, water tem¬

perature and hydrological conditions on Penaeas setiferus (Linnaeus). M.S.

728. Yates, Matthew M., 1974-Field and laboratory observations of possible interactions

between some acquatic algae and Culex pipiens quniquefasciatus (The Southern
House mosquito). M.S.

NORTH TEXAS STATE UNIVERSITY

729. Atmar, Gerald L., 1970-Food, feeding and ecological efficiencies of Fundulus

notatus (Rafinesque) (Osteichthyes; Cyprinodontidae) Ph.D.

730. Bokich, J.M., 1976-Demography and life history energetics of White Bass, Morone

chrysops. Ph.D. (In progress)

731. Burleson, J., 1976-Effect of chlorine on amino acids and polypeptides in municipal

wastewaters. M.S. (In progress)

TEXAS WATERS

333

732. Childress, M., 1976-Systems model of energy flow and trophic dynamics of a

North Central Texas pond ecosystem. M.S. (In progress)

733. Dermanci, AM, 1976-Size distribution and chlorine contents of treated waste-

water effluents. M.S. (In progress)

734. Fagan, I., 1976-Life history energy allocation to growth and reproduction in Giz¬

zard Shad. M.S.

735. Fullington, Richard W., 1976-The aquatic gastropods of Texas. Ph.D. (In progress)

736. Fullington, Kate E., 1976-Nymphs of the stonefly genera, Taeniopteryx and Zea-

leuctra of North America. M.S. (In progress)

737. Gash, Stephen L., 1971 -Effects of Carbaryl (1-napthyl-N-Methylcarbamate)

on Trichocorixa reticulata (Hemiptera:Corixidae) and Glyptotendipes barbipes
(Diptera : Chironomidae). Ph.D.

738. Grant, Peter, 1976-Life cycle of the mayfly Isonychia sicca manca (Eaton) in

Clear Creek and the Brazos River, Texas. M.S. (In progress)

739. Glidewell, J., 197 6 -Life history, population and niche analyses of Red Slider turtles

in North Central Texas. Ph.D. (In progress)

740. Huang, Francis, 1976-Photolytical oxidation of polynuclear aromatic hydrocarbons

in aqueous solutions. M.S. (In progress)

741. Henderson, James E., IV, 1976— GC/MS studies of organochlorine compounds

formed by the chlorination of municipal wastewaters. Ph.D. (In progress)

742. James, Gary L., 1976-Genetic variation in red shiners, Notropis lutrensis in the

Brazos and Trinity Rivers. M.S. (In progress)

743. Jones, J., 1976-Biochemical variation in the top minnow Fundulus. M.S. (In prog¬

ress)

744. Lin, Simon, 1976-Photolytic oxidation of trihalomethanes in aqueous solutions.

M.S. (In progress)

745. McCullough, W., 197 6 -Phytoplank ton- salinity relationships in Lake Texoma.

M.S. (In progress)

746. Merritt, R., 1976-Genetic variation and the effects of isolation on the Stoner oiler,

Campostoma anomalum. M.S. (In progress)

747. McClure, Richard G., 1974-The life history and ecology of the mayfly Neochoro-

terpes mexicanus Allen (Ephemeroptera: Leptophlebiidae) M.S.

748. Perry, W.B., 1976-Biomass and carbon relationships in water and sediment of

a meso trophic Texas reservoir. M.S. (In progress)

749. Oberndorfer, Reed Y., 1976-The life cycle of Hydroperla crosbyi (Needham Claas-

sen) (Plecoptera : Perlodidae) M.S.

750. Poole, Walton C., 1973-The vertical stratification of the macrobenthos in the

Brazos River, Texas. M.S.

751. Richmond, M.C., 1 97 6 -Evolutionary genetics of red shiners, Notropsis lutrensis ,

inhabiting a cooled effluent from a reservoir. M.S. (In progress)

334

THE TEXAS JOURNAL OF SCIENCE

752. Shelton, Bonnie K., 1976-The use of scanning electron microscopy in Limnology.

M.S. (In progress)

753. Snellen, Rosalyn K., 1976-Life cycle of Zealeuctra claasseni, Zealeuctra lutei,

Perlesta placida (Plecoptera) and Chimarrha obscura (Trichoptera) in North
Texas. Ph.D. (In progress)

754. Widner, D., 1976-Stream ecology of White Bass Morone chrysops in North Central

Texas. M.S. (In progress)

755. Ziegler, David D., 1976-Drumming behavior of selected Nearctic Stoneflies (Ple¬

coptera). M.S.

UNIVERSITY OF TEXAS AT AUSTIN

756. French, Dorian, 1976-Heavy metal analysis of raw water supply and wastewater

discharge for the city of Austin, Texas. M.S.

756a Hoffman, H.W., Jr., 1976-A continuous equation to describe bioassay results for
acute exposure of fish to toxins. M.S.

757. Neuse, David W., 1976-The removal of algae from an oxidation pond effluent

through the use of a tertiary water hyacinth pond system. M.S.

758. Roques, Pat F., 1975 -Effects of fluctuating nutrient and regimes on autotrophic

succession in microecosystem. M.A.

TEXAS CHRISTIAN UNIVERSITY

759. DeLuca, Robert C., 1976-A study of the interrelationship of naturally collected

bacteria and several species of algae. M.S.

760. O’Kane, Kevin D., 1976-A population study of the exotic bivalve Corbicula man-

ilensis (Philippi, 1841) in selected Texas reservoirs. M.S.

TEXAS A&M UNIVERSITY

761. Camejo, Daniel, 1941-Typical anopheles breeding places formed by irrigation.

M.S.

762. Crocker, B.R., 1975 -An inventory of the original vegetation of the Palmetto Bend

reservoir site in Jackson County, Texas with respect to potential eutrophica¬
tion. M.S.

763. Coffey, J.E., 1956-Flood in the Nueces, Guadalupe, Lavaca and Mission River

Basins: magnitude and frequency. M.S.

764. Hayes, W.A., 1976-Comparative studies of agonistic behavior in cambarid cray¬

fish. Ph.D.

765. Holt, G.J., 1976-Community structure of macrozooplankton in Trinity and upper

Galveston Bays, with special reference to the cooling water system of Cedar
Bayou Electric Generating Station. Ph.D.

TEXAS WATERS

335

766. Hall, E.R., 1976-Water quality of some coastal canal communities in the Galveston

County area. M.S.

767. Janssesn, H.E., 1976 -Distribution and fate of technical Chlordane and Mirex resi¬

dues in a central Texas aquatic ecosystem. M.S.

768. Langford, T.R., 1948-A study of transition of flow in open channels with flow

at a sub- critical depth. M.S.

769. Maldonado, R.J., 1976-Steady state estuarine modeling of the Brownsville Ship

Channel. M.S.

770. Priebe, W.F., 1972— Oxygen uptake of bentic systems. M.S.

771. Rahman, K., 1963-Study of the effect of rate of sediment transport on the stability

of open channels. M.S.

772. Reavis, M.W., 1976-The environmental management of a ship channel complex.

M.S.

773. Smith, R.H., 1968-Survey of organic content in bottom of the Houston Ship

Channel. M.S.

774. Vigil, S.A., 1974-Nutrient sources and loading for the proposed Millican Lake.

M.S.

775. Withers, R.E., 1969-Salinity patterns in the Houston Ship Channel. M.S.

776. Walsh, Don, 1968 -The Mississippi River outflow, its seasonal variations and its

surface characteristics. Ph.D.

SUBJECT INDEX

Aquatic Arthropoda (General-Ostracoda, Hydracarina, etc.)

21, 109, 188, 193, 214d, 288b, 314a, 326, 333a, 658,689, 698, 699, 727, 728, 764

Aquatic Vertebrates

26, 29, 30, 197, 201, 202, 203, 212, 220, 255, 261, 327, 421, 681, 693, 696, 697, 705,
739

Benthos (Aquatic insects, etc.)

25, 33, 35, 37, 38, 41, 43, 45, 53, 56, 60, 64, 75, 89, 97, 100, 115, 119, 120, 130, 131,

134, 137, 144, 160, 165, 169, 171, 176, 177, 178, 185, 187, 189, 196, 198, 204, 210,

214, 214f, 214j, 222, 224, 233, 248, 256, 262, 277, 280, 284, 299, 313, 339, 344, 406,
407, 408, 424, 430, 433, 481, 503, 507, 587, 592, 658, 684, 735, 736, 737, 738, 746,
747,749,750,753,761,770

Biological and Ecological Surveys (General)

2, 3, 15, 30, 35, 47, 58, 78, 90, 95, 99, 105, 118, 128, 141, 144, 150, 154, 155, 162,

165, 174, 183, 193, 210, 213, 214e, 222, 224, 261, 276, 308, 349, 351, 358, 434, 453,

48 L, 540, 580, 583, 584, 627, 658, 694, 695, 729, 732, 739, 747, 748, 754, 755, 758,
759,762,767,772,773

Computer Science

199, 200, 206, 207, 211, 214i, 267, 297, 305, 312, 333, 367, 379, 499, 579, 603, 706,
720

336

THE TEXAS JOURNAL OF SCIENCE

Energetics, Biochemistry

55,56, 106, 129, 142,459,623,649,698,699,729,730,732,743

Geology

10,350,361,461,514,609,642,652,770

Groundwater

199, 200, 201, 204, 208, 214, 214h, 374, 409,545,670, 671,679

Hazardous Materials (Metals, Pesticides, Bioassays, Chlorinated hydrocarbons, Chlorine,
Toxicity studies)

31, 32, 34, 40, 42, 44, 52, 59, 63, 71, 82, 88, 93, 100, 116, 121, 179, 184, 196, 237,

242, 258, 268, 270, 273, 281, 311, 316, 331, 332, 343, 345, 356, 362, 364, 375, 450,

474, 483, 507, 511, 529, 534, 557, 558, 563, 569, 578, 585, 592, 613, 615, 622, 626,

653, 685,689, 721, 726, 737, 740, 744, 756, 756a, 767, 773

Hurricane, Weather
164,333,339

Ichthyology

I, 4, 5, 7, 8, 9, 16, 17, 18, 19,22,23,24,27,32,61,67,68,70,87,88,91,93,94,96,

103, 106, 107, 108, 110, 111, 124, 125, 127, 138, 143, 148, 149, 150, 151, 154, 158,
166, 172, 184, 199 , 2 03, 206 , 209 , 211, 2 1 4b, 214e, 219, 221, 224, 227, 231, 235, 239,2 243,
254, 265, 269, 270, 274, 279, 281, 283, 286, 287, 288, 288b, 290, 291, 292a, 294, 296,
301, 302 , 309 , 314 , 315 , 3 1 6, 319 , 32 0 , 32 3 , 325 , 329 , 331, 335 , 33 6 , 337 , 341, 343 , 345,

346 , 347 , 35 3 , 359 , 362 , 4 1 0 , 411, 4 1 2 , 4 1 3, 4 1 4 , 415 , 4 1 6, 4 1 7 , 4 1 8 , 4 19 , 4 20 , 4 22,

423, 426, 427, 428, 435, 436, 437, 439, 441, 446, 449, 474, 480, 493, 496, 497, 518,

5 24 , 525 , 5 2 7 , 5 37 , 5 40 , 544 , 5 60 , 561, 5 64 , 5 99 , 613, 618 , 624 , 628 , 645 , 650 , 656,

680, 729, 730, 734, 742, 743, 751, 754, 756a

Macrophytes and Periphyton

II, 31, 74, 100, 192, 200, 207, 240, 300, 321, 328, 376, 378, 434, 482, 457, 459, 491,
570, 607,638, 640, 649, 654, 665,686, 690, 757

Management, General Hydrology, History, Law, Economics, Education

13, 146, 202, 204, 205, 207, 251, 259, 272, 278, 285, 290, 304, 322, 369, 370, 371,

373, 425, 475, 476, 479, 484, 464, 465, 466, 495, 517, 523,532,5 38,586,588,637,

644, 652, 655, 659, 666, 667, 668, 669, 672, 678, 683, 700, 702, 763, 768, 771, 772,

773,775

Microbiology, Bacteriology, Mycology

15, 46, 48,57, 62, 63, 65, 71, 73, 74, 79, 84, 86, 92, 102, 104, 132, 161,216,228,271,
307, 520, 584, 593, 635, 662, 674, 676, 694, 695, 704, 705, 707, 708, 711, 712, 714,
715,716,718,720,722,724,752,759,

Modeling, Biological Indexes

122, 222, 292a, 465, 471, 495, 512,526,535,538,552,554,567,572,588,595,596,
625,647,700, 703, 706, 720, 732, 734, 756a, 769, 775 , 776

TEXAS WATERS

337

Mollusca

40, 147, 159, 162, 167, 168, 174, 175, 181, 203, 218, 252, 253, 260, 293, 330, 342,
355,429,735,760

Nutrients , Eutrophication , Water Quality

3, 20, 54, 76, 80, 83, 98, 113, 123, 125, 126, 139, 141, 170, 183, 210, 214i, 214k, 215,
229, 238, 241, 245, 257, 263, 264, 276, 289, 290, 292, 294, 303, 306, 308, 310, 315,

348, 351, 360, 363, 365, 370, 374, 438, 439, 442, 453, 478, 463, 465, 468, 491, 502,

511, 521, 530, 532, 547, 549, 550, 577, 581, 591, 598, 600, 601, 609, 629, 635, 639,

655, 663, 664, 665, 675, 677, 685, 687, 688, 694, 695, 700, 701, 702, 703, 709, 710,

713, 714, 717, 719, 723, 725,748, 758, 762, 766, 774

Physico-chemical, Geochemistry , Water Chemistry, Tastes and Odors
12, 14, 33, 36, 37, 38, 40, 41, 44, 45, 57, 65, 81, 82, 84, 85,88,93, 117, 118, 123, 128,
151, 157, 183, 191, 204, 208, 211, 213, 214c, 214g, 214k, 223, 226, 229, 230, 232,
241, 276, 284, 289, 303, 304, 305, 306, 310, 318, 348, 351, 357, 360, 438, 439, 451,

476, 485, 486, 494, 519, 553, 567, 568, 591, 594, 599, 606, 631, 632, 633, 639, 648,

651, 655, 664, 745,770, 775,776

Phytoplankton (Algae)

6, 20, 28, 39, 49, 50, 51, 52, 55, 59, 69, 72, 77, 78, 81,-85, 98, 99, 100, 102, 109, 115,
116, 118, 119, 121, 125, 130, 133, 136, 139, 142, 144, 153, 155, 157, 163, 164, 173,
180, 195, 212, 214a, 236, 244, 246, 250, 258, 264, 268, 289, 295, 298, 300, 307, 321,
334, 340, 358, 372, 432, 434, 444, 445, 447, 448, 452, 453, 454, 478, 483, 458, 460,

462, 467, 470, 472, 489, 490, 492, 498, 501, 504, 505, 506, 507, 511, 516, 522, 531,

536, 539, 541, 542, 543, 546, 548, 555, 556, 562, 563, 565, 569, 573, 574, 575, 576,

577, 582, 589, 590, 597, 602, 604, 605, 611, 612, 614, 617, 619, 620, 622, 623, 629,

630, 634, 636, 641, 646, 653, 657, 660, 673, 679, 682, 685, 701, 728, 745, 757, 758,

759

Primary Production (Chlorophyll a, Biomass, Photosynthesis)

54, 95, 126, 133, 135, 140, 163, 186, 214g, 214k, 236, 244, 246, 282, 295, 300, 324,
334,483,500,506,534,539,686,691,719

Radioactive

499,509,516,582,598,608

Sewage, Effluents, Treatment (Oxidation ponds)

35, 36, 43, 80, 92, 146, 148, 163, 194, 280, 312, 344, 365, 442, 443, 481, 488,502,
510, 515, 528, 564, 571, 581, 598, 615, 621, 639, 645, 661, 670, 713, 715, 718, 722,
731,733,741,756,757

Thermal , Electric Power

11, 17, 20, 21, 64, 96, 100, 107, 111, 123, 133, 138, 147, 157, 187, 214a, 214k, 233,
236, 279, 281, 283, 286, 287, 288a, 288b, 29 1, 292a, 299, 301, 309, 314, 314a, 317, 323,
325, 333a, 335, 336, 337, 339, 358, 373,431,433,464,500,5 13,524,537,559,605,
613,614,621,631,632,650,685,689,727,751,765

338

THE TEXAS JOURNAL OF SCIENCE

Urban Runoff, Storm Runoff, Non- point Source

35, 114, 194, 206, 209, 211, 247, 266, 297, 363, 366, 367, 368, 377, 440, 477, 508,
579,596,603,610,707

Zooplankton

39, 47, 99, 109, 115, 119, 129, 130, 145, 166, 179, 182, 190, 212, 214f, 214j, 217,234,
249, 275, 288a, 317, 326, 338, 340, 354, 358,431,456,550,559,566, 616, 765

m

IS CANOPY INTERCEPTION AN ACCURATE MEASURE OF LOSS
FROM THE HYDROLOGIC BUDGET?

by MINGTEH CHANG

School of Forestry,

Stephen F. Austin State University,

Nacogdoches 75961

INTRODUCTION

Interception is the process by which the aerial portions of vegetation intercept
and retain precipitation until it is evaporated. The amount of precipitation that
is intercepted by vegetation, estimated at 10-20% of the annual total (Linsley,
et al, 1975), varies with storm intensity and duration, precipitation type, wind
speed, type of plant cover, and season of the year. Since intercepted water does
not reach the ground and contribute to soil moisture or stream flow, it is widely
considered as a loss to the hydrologic budget.

The evaporation of intercepted water is similar to water loss from an open water
surface. Due to the small diffusive resistance at the evaporating surface, it is ex¬
pected that evaporation of intercepted water occurs at a greater rate than trans¬
piration under similar atmospheric conditions. During evaporation of intercepted
water, the energy available for transpiration is reduced, and a reduction in trans¬
piration is expected. Interception therefore should not be a complete loss from
hydrologic budgets. This idea has been discussed by Christiansen (1942), Goodell
(1963), Penman (1963), Lull (1966), and others, and studied by Rakhmanov (1958),
Burgy and Pomeroy (1958), McMillan and Burgy (1960), and more recently by
Harr (1966), Thorud (1967), and Nicolson,ef al, (1968).

Investigators agree that evaporation and transpiration are energy dependent
processes, but appear to differ on the availability of energy for them. One group
feels that evaporation of intercepted water preempts energy which would have
been used for transpiration. Others appear to believe that more energy is available
for evaporation over a wet canopy as compared to a dry canopy. Thus, Murphy
and Krioerr (1975) state that ‘Interception of precipitation does represent a loss
of water to the soil and stream flow under field conditions.” It is the purpose of
this paper to evaluate the question: “Should canopy interception be considered
as a loss in the hydrologic budget, or not?”

Accepted for publication: December 3, 1976.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

340

THE TEXAS JOURNAL OF SCIENCE

INTERCEPTION MEASUREMENTS

The interception process is composed of 2 phases: 1) interception storage, or
retaining precipitation in the vegetative canopy, and 2) evaporation of the inter¬
cepted water. Interception loss, properly defined, is the amount of evaporation
minus the amount of transpiration reduction. Since direct simultaneous measure¬
ment of transpiration and interception loss is not possible, estimates must be based
on indirect methods.

Usually, interception is estimated by comparing precipitation measured in the
open, P0, and that under the vegetative canopy, Pc, with a correction for stemflow,
i.e., for the water which runs down the vegetative stems, Ps. Thus the canopy in¬
terception, Ic, can be calculated by the following equation

Ic = (P0-pc)-ps 0)

P0 and Pc in Equation 1 are also refered to as gross precipitation and net precipi¬
tation respectively. Water intercepted by forest litter, a related phenomenon re¬
quiring other measurement approaches, is not included in this discussion.

The concept of Equation 1 has been widely employed in scientific research
and for practical purposes by Beall (1934), Rowe and Hendrix (1951), Semago
and Nash (1962), Leonard (1961), Miller (1964), Rogerson and Byrnes (1968),
and many others. Stemflow (Ps) can be measured by a rolled gutter attached to
the tree trunk, the catch from which is drained into a container. The volume of
water collected in containers is measured and converted into depth over the sam¬
pled areas. Precipitation in openings (gross precipitation, P0) and under vegetative
canopies (throughfall, Pc) is measured with multiple conventional rain gages, under
careful sampling techniques (Helvey and Patric, 1965; Lawson, 1967; Zinke, 1967).

INTERCEPTION STUDIES
New Findings

Rakhamanov (1958) observed the transpiration rates of sprinkled and unsprinkled
excised plant branches, whose bases were sealed into a water filled bottle. Water
loss due to transpiration was determined by weighing, and expressed as the quantity
of water transpired per unit fresh weight of the branch. He found that transpira¬
tion of aspen, downy birch, hazels, yellow acacia, oak, and sedge basket willow
was reduced from 15 to 40% by spraying. The author concluded: “The precipita¬
tion intercepted by the tree crowns . . . cannot be considered as a loss to the water
balance of the forest soils.”

Burgy and Pomeroy (1958) studied interception by grassy species grown in
nutrient solutions in a greenhouse. Rates of evapotranspiration from sprayed and
unsprayed pans were compared. “The evaporation of a given amount of inter¬
cepted moisture,” the authors found, “was accompanied by a like reduction in
the amount of evapotranspiration from the plants.” The total moisture loss in

CANOPY INTERCEPTION

341

sprayed and unsprayed pans was about the same. Thus, the authors suggested
that the terminology of interception loss should be re-examined on the basis of
the net effect.

Later McMillan and Burgy (1960) measured evapotranspiration from sprayed
and unsprayed vigorous grass covers in floating lysimeters in a grass field. They
concluded that, on the average, the difference between sprayed and unsprayed
plots was not significant, a result consistent with that of the earlier greenhouse
study. Lee (1968), however, called attention to the fact that, during the first
hours after spraying, evapotranspiration increased by 16%.

Thorud (1967) evaluated interception- transpiration relationships of small
trees. Treatment and control groups were randomly selected from ten 6- to 7 -yr-
old ponderosa pines well established in 10-qt plastic pails. The pail, and the base
of each tree, were sealed with plastic to prevent evaporation from the soil, and
transpiration was determined by weight differences. Before applying water to the
trees, the relationship of transpiration rates between control and treatment groups
were calibrated. Water of known quantities was then sprayed on the treatment
trees and the concurrent transpiration losses of control and treatment trees were
measured simultaneously. The amount of transpiration reduction was estimated
with an equation based on the calibration data. The results showed that about 9%
of the applied water was conserved in the soil. In other words, about 91% of the
applied water was a net interception loss to soil moisture. The author concluded
that transpiration reduction was small compared with the losses implied by earlier
studies of interception by grass.

In similar studies with Colorado blue spruce and Austrian pine in small pots,
Harr (1966) found average reductions in transpiration of about 6%. Nicolson,
et al., (1968), using white spruce and white pine in Minnesota, measured average
savings as about 12% of the intercepted water applied.

The studies cited above provided evidence that intercepted water reduced
transpiration and consequently conserved soil moisture. The amount of water
conserved depended in part on weather conditions (Nicolson, et al. , 1968), and
varied among species.

Energy Source

Since evaporation is an energy dependent process, many have attempted to
interpret interception loss in terms of available energy. For example, in discussing
the special difficulties of estimating evaporation, Penman (1963) states:

. . . while the leaves are wet the evaporation is effectively
as from an open water surface rougher (sic) than a nor¬
mal plane surface so that the evaporation rate will be
greater during the day than that of transpiration of soil
water, and can go on at night. But the same energy can¬
not be used twice, and while the intercepted water is
being evaporated the drain on soil water is checked. Inter¬
cepted water is not wasted; though its use may be slightly
luxurious.

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THE TEXAS JOURNAL OF SCIENCE

Penman’s (1963) statement on the evaporation rate from a wet canopy is supported
by several studies. In a stand of Scots pine in southeast England, Rutter (1967)
found that intercepted water evaporated, on average, about 4 times as fast as the
transpiration rate under the same environmental conditions. Murphy and Knoerr’s
(1975) simulation study indicates that evaporation of wet leaves may be from
1.5 to more than 2 times greater than that of dry leaves. They further explain
that the energy source for the greater evaporation of a wet canopy comes from
an increase in net radiation through “a decreased long-wave radiation and a de¬
creased or even negative sensible heat flux.” Thus, they do not agree on the hypothe¬
sis that the amount of energy available for evaporation is fixed in a particular
environment, and claim that precipitation interception does represent a hydrologic
loss under field conditions. Apparently, they ignore the 10% compensating reduction
in evapotranspiration caused by the evaporation of intercepted water.

EVALUATION

These studies agree that evaporation of intercepted water is greater than the
transpiration from dry leaves under the same environmental conditions. The inter¬
pretation of intercepted water in the hydrologic budget, however, remains con¬
troversial. This report will consider the interception process from three viewpoints:
available energy, water vapor diffusive resistance, and plant physiology.

Available Energy

The energy required to change water from liquid to vapor at ordinary air tem¬
perature is about 580 cal/g. The energy for sustained vaporization of water from
a vegetative surface, such as a forest is primarily from solar radiation. The net
radiation, S, of a stand is given by

S = (I + H)(1 -r) + G- aT4 (2)

where I is direct solar radiation, H is sky radiation, r is the reflectivity of incoming
short-wave radiation or so-called albedo, G is long-wave radiation from the atmos¬
phere, and a T4 is the long-wave radiation emitted by the forest, in which T is
surface absolute temperature in °K, and a is Stephen-Boltzmann constant =
8.26 (lCT11 ) cal/cm2min°K4. The albedo, r, is affected by the nature of a surface,
and to some extent by its moisture content. For example, Leyton, et al., (1967)
found that wetting sharply reduced the albedo of a spruce canopy in England
and the albedo gradually increased as the foliage dried out.

From Equation 2 it is evident that albedo, r, and surface temperature, T, are
the only 2 surface parameters that can change the net radiation. Albedo, on average,
is about 0.14 for coniferous forests, and up to 0.25 for deciduous trees; it is about
0.05-0.07 for a free water surface (Bruce and Clark, 1966) at high sun angles. A
canopy wetted by intercepted water probably has a lower albedo than a dry can¬
opy. On the other hand, the canopy temperature is lowered when intercepted

CANOPY INTERCEPTION

343

water is evaporated. Rutter (1967) found that the wet leaves of Scots pines were
1.0 C cooler than the air. Consequently, with a lowered canopy temperature,
the canopy outgoing radiation, qT4, is reduced. Equation 2 shows that negative
changes in r and aT4 caused by the presence of intercepted water will cause a
positive change in S.

The net energy delivered to a forest stand is dissipated primarily by convection,

H, vaporization, L, and conduction, B, as indicated by the equation:

S = H + L + B (3)

From Equation 3, the heat flow of vaporization is given by:

L = S - (H + B) (4)

Equation 4 indicates that vaporization increases with S and decreases with H + B.
But B is usually a small portion of the total available energy in a forest. Therefore,
L is mainly affected by the available energy, S, and the heat exchange between
the canopy and the atmosphere, H. According to the above analysis, a wet canopy
would cause a positive change in S, and, because of a reduction of surface tem¬
perature, the sensible heat flux, H, is reduced or even negative. This will result
in a higher rate of vaporization, L.

Diffusive Resistance

That the evaporation of intercepted water is faster than the transpiration from
the leaf can be explained in terms of relative resistances in the water vapor diffusion
pathways. Consider the following equation,

T (g/cm2 sec) = (Cs - Ca)/(Rf + Ra) (5)

where T is the transpiration rate, Cs and Ca are the water vapor concentrations
(g/cm3) at the evaporating surface within a leaf and in the bulk air respectively,
and Rf and Ra are the diffusive resistances (sec/cm) in the leaf stomates and in
the surface boundary layer of air outside the leaf. Equation 5 is mainly a function
of temperature if diffusive resistance remains constant, because Cs is uniquely
determined by temperature. However, if weather conditions remain constant,
the diffusive resistance will play an important role in determining the transpira¬
tion rate. When water is intercepted at the leaf surface, Rf = 0, and the total re¬
sistance is reduced, and Equation 5 becomes

E (g/cm2 sec) = (CS-Ca)/Ra (6)

where E is the evaporation rate of intercepted water. Rutter (1967) in his Scots
pine study found that the resistance to water vapor diffusion from a wet leaf was

I. 24 sec/cm, while that for transpiring leaves with dry surfaces was 9.1 sec/cm. The
large difference in diffusive resistances explains the higher evaporation rate from
wet leaves.

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THE TEXAS JOURNAL OF SCIENCE

Plan t Physiology

Physiologically, water movement through the soil -plant-atmosphere system is
mainly controlled by water potential gradients in the soil and plants, and by the
vapor pressure gradients between evaporating surfaces and the air (Kramer, 1969).
As water is transpired from leaves, the water potential in the leaf cells is reduced.
This causes water to move from the area of higher water potential in the xylem
of the leaf veins into leaf cells. Removal of water from the xylem reduces its water
potential, and this reduction is transmitted through the water conducting systems
of the plant and into the soil. The energy required for* these water movements,
together with the fact that transpiration from leaves is largely limited to interior
surfaces of stomatal openings, account for the greater diffusive resistance to trans¬
piration.

Rutter’s (1967) study demonstrated that “the rate of evaporation of intercepted
water is, on average, about 4 times as great as the transpiration rate in the same
environmental conditions.” Intercepted water, presenting an open surface to the
air is obviously more available to evaporation than the water in leaf cells, and must
be evaporated before transpiration can occur. Since one source of energy cannot
be used in two different places, energy dissipated by evaporating the intercepted
water at the leaf surface reduces the energy available for leaf transpiration. It should
be noted, however, that the energy available for evaporating the intercepted water
is not necessarily a fixed fraction of solar radiation. The reduction in transpira¬
tion caused by intercepted water, ranging from 100% (Burgy and Pomeroy, 1958;
McMillan and Burgy, 1960) to about 10% (Harr, 1966;Thorud, 1967; Nicolson,
et al, 1968), depends on the duration of intercepted water on the leaf’s surfaces,
and varies with species and weather conditions.

For an adequate understanding of the significance of transpiration reduction
in interception of rainfall, a thorough analysis of heat transfer and water dissipa¬
tion over natural wet and dry canopies is necessary.

CONCLUSION

Evaporation and transpiration are energy dependent processes. A vegetative
canopy wet with intercepted water has more energy available for evaporation by
increased absorption of short-wave radiation, decreased emission in long-wave
radiation, and a decreased or even negative sensible heat flux between canopy
and the adjacent air. With more available energy, and a much lower diffusive re¬
sistance at the evaporation surface, evaporation of intercepted water occurs at a
greater rate than transpiration under similar atmospheric conditions. So long as
intercepted water is available for evaporation on a leafs surface, transpiration is
reduced or inhibited. Thus intercepted water is not a complete loss to the hydro-
logic cycle. Measurement of these two types of water loss has proved difficult,
and the relative importance of reduced transpiration, especially under field con¬
ditions, is uncertain. Obviously, however, it is too large to be ignored in precise
studies.

CANOPY INTERCEPTION

345

The variability of results obtained by different investigators illustrates the diffi¬
culty of measuring the relative rates of water movement in vapor state, and the
multiplicity of factors affecting them. These problems also suggest the need for
highly instrumented research designed to measure such rates under natural con¬
ditions and within specific vegetative types, together with data on temperature,
humidity, radiation patterns, energy flows, wind movement, and rainfall charac¬
teristics. Intensive study of the interaction of transpiration , interception , and evap¬
oration with these and other factors will be essential to an accurate understanding
and evaluation of interception losses in forests and similar vegetative types.

LITERATURE CITED

Beall, H. W., 1934-The penetration of rainfall through hardwood and softwood forest canopy.
Ecology , 14:412.

Bruce, J. P., and R. H. Clark, 1966 -Introduction to Hydrometeorology . Pergamon Press,
New York, N.Y., 319 pp.

Burgy, R. H., and C. R. Pomeroy, 1958-Interception losses in grassy vegetation. Trans.
Amer. Geoph. Union, 39(6): 1095.

Christiansen, J. E., 1942-Irrigation by sprinkling. Univ. of Calif. Agr. Exp. Sta. Bull., 670:115.

Goodell, B. C., 1963-A reappraisal of precipitation interception by plants and attendant
water loss./. Soil and Water Corner., 11-12:231.

Harr, R. D., 1966-Influence of intercepted water on evapotranspiration from small potted
trees. Ph.D. dissertation, Colo. St. Univ., Fort Collins.

Helvey, J. D., and J. H. Patric, 1965 -Canopy and litter interception of rainfall by hardwoods
of eastern United States. Water Resources Res., 1:193.

Kramer, P. J., 1969 -Plant and Soil Water Relationships: A Modern Synthesis. McGraw-Hill
Book Company, 482 pp.

Lawson, E. R., 1967-Throughfall and stemflow in a pine-hardwood stand in the Ouachita
Mountains of Arkansas. Water Resources Res., 3(3) :7 3 1 .

Lee, R., 1968-Reply to Dr. Idso’s letter. Water Resources Res., 4(3):667.

Leonard, R. E., 1961 -Interception of precipitation by northern hardwoods. U.S. Forest Ser.
N.E. Forest Exp. Sta. Paper 159, 16 pp.

Leyton, etal, 1967-Rainfall interception in forest and moorland. In Sopper and Lull (Eds.),
International Symp. on Forest Hydro. Pp. 163-178.

Linsley, etal. , 197 5 -Hydrology for Engineers. McGraw-Hill Book Co ., New York , N.Y., 482 pp.

Lull, H. W., 1966-Ecological and silvicultural aspects. In V.T. Chow (Ed.), Handbook of
Applied Hydrology. Pp. 6-9.

McMillan, W. D., and R. H. Burgy, 1960-Interception loss from grass./ Geoph. Res., 65(8):2389.

Miller, D. H., 1964-Interception processes during snow storms. U.S. Forest Ser. Pacific S.W.
Forest and Range Exp. Sta. Res. Paper 18.

346

THE TEXAS JOURNAL OF SCIENCE

Murphy, C.E., Jr., and K. R. Knoerr, 1975-The evaporation of intercepted rainfall from a
forest stand: An analysis by simulation. Water Resources Res., 1 1(2) :27 3-280.

Nicolson,ef al., 1968 -The interception-transpiration relationship of white spruce and white
pine./. Soil and Water Conserv. 9-10:181.

Penman, H. L., 1963 -Vegetation and Hydrology. Tech. Comm. No. 5 3, Commonwealth
Bureau of Soils, Harpenden.

Rakhmanov, V. V., 1958-Are the precipitations intercepted by the tree crowns a loss to the
forest? USDC. Washington, D.C. Translated from Russian, Botanicheskii Zhurnol, 43:1630.

Rogerson, T. L., and W. R. Byrnes, 1968-Net rainfall under hardwoods and red pine in cen¬
tral Pennsylvania. Water Resources Res., 4(1) :5 5 .

Rowe, P. B., and T. M. Hendrix, 195 1 -Interception of rain and snow by second-growth pon-
derosa pine. Amer. Geophys. Union Trans., 32:903.

Rutter, A. J., 1967 -An analysis of evaporation from a stand of scotch pine. In Sopper and
Lull (Eds.), International Symp. on Forest Hydro. Pergamon Press, pp. 403-417.

Semago, W. T., and A. J. Nash, 1962-Interception of precipitation by a hardwood forest
floor in the Missouri Ozark s. Univ. Missouri Agr. Exp. Sta. Res. Bull, 796:31.

Thorud, D. B., 1967-The effect of applied interception rates on potted ponderosa pine.
Water Resources Res., 3(2):443.

Zinke, P. J., 1967-Forest interception studies in the United States. In Sopper and Lull (Eds.),
International Symp. on Forest Hydro. Pp. 137-161.

CRITICAL LEVELS OF PHOSPHORUS AND NITROGEN IN
TEXAS IMPOUNDMENTS1

by G. FRED LEE

Center for Environmental Studies,

University of Texas, Dallas 75080

INTRODUCTION

The 1972 amendments to the Federal Water Pollution Control Act require
that each state classify its lakes and impoundments with respcet to the degree of
aquatic plant fertility. Further, this Act requires the state to develop a plan for
remedial action for those bodies of water that are determined to be excessively
fertile. Texas, as well as other states, is in the process of examining the available
information on nitrogen and phosphorus loadings, the sources of these nutrients,
and the resultant concentrations in the State’s impoundments. Further, they are
attempting to assess the effect of these loads on water quality in the impoundments
and the element(s) or other factors that exert primary control on aquatic plant
growth in the water body. In the State of Texas the responsibility for making
this assessment falls with the Water Quality Board. This organization must devel¬
op and implement an aquatic plant nutrient management plan which will restore
water quality in those impoundments that are determined to be receiving exces¬
sive loads of the critical nutrients) that control aquatic plant growth. The
results of these evaluations could be of great significance to the State of Texas in
that aquatic plant nutrient control could cost State residents very large amounts
of money for advanced waste treatment and altered agricultural and urban prac¬
tices. Because of the large amounts of funds needed for aquatic plant nutrient
control, it is important that the evaluation of the process governing algal growth
be carefully done in order to develop technically sound, economically feasible
managament plans for the problems of excessive fertilization of Texas’ impounded
waters.

One of the most critical aspects of the evaluation process is the determina¬
tion of the critical levels and the loads that cause these levels of aquatic plant nu¬
trients in Texas’ impoundments. Because of the significance of this topic to the
residents of Texas, it is important to discuss the approach that should be used to
establish critical loading criteria for Texas’ impounded waters. Presented below
is a discussion of the information available on critical levels of nitrogen and
phosphorus in the State’s impoundments.

^ES Occasional Publication No. 2.

Accepted for publication: Date unknown.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

348

THE TEXAS JOURNAL OF SCIENCE

Previous Studies on Critical Levels of Aquatic Plant Nutrients

The classical work on the critical concentrations of aquatic plant nutrients
was conducted by C. N. Sawyer in the 1940’s. Sawyer studied a group of South¬
ern Wisconsin lakes and concluded that whenever the phosphorus exceeded 0.01
mg/ 1 P and the inorganic nitrogen exceeded 0.3 mg/ 1 N the lake was likely to
have deteriorated water quality in the summer due to excessive algal and other
aquatic plant growth.

Dr. R. A. Vollenweider has reviewed the data obtained on several European
lakes and concluded that the 0.01 mg/1 P and 0.3 mg/1 N is appropriate for these
lakes. The author has spent 13 years in Wisconsin studying problems of excessive
fertilization of Wisconsin and other mid-western lakes and impoundments. As a
result of these studies, he generally agrees with Sawyer on the critical levels of
nitrogen and phosphorus in Wisconsin waters. During the past year, however,
the author has had the opportunity to review the information on nutrient loads-
algae response in Texas impoundments and concludes that at the present time
there is insufficient information to establish critical nitrogen and phosphorus
loadings or in-lake concentrations for these waters. While the critical concentra¬
tions cannot be established, it is clear that it is not technically sound to apply
the 0.01 mg/1 P and the 0.3 mg/1 N criteria to Texas impoundments without a
thorough evaluation of these criteria. The basis for this judgment is presented
below.

Water Quality and Water Use

The Sawyer criteria are based on subjective judgment of impaired water qua¬
lity due to excessive algal and other aquatic plant growth in the lakes that he
studied. The large number of small lakes in Wisconsin affords the opportunity
for someone to readily compare water quality in lakes within a relatively short
distance. Within one day’s drive, an individual in Wisconsin can observe lakes with
almost distilled water quality which are essentially free of aquatic plant growth
to extremely fertile lakes where ducks and turtles can walk on the algal scum
that accumulates along the shore. Texas impoundments also have a wide variety
of water quality although rarely are waters encountered that would be classified
as oligotrophic by Wisconsin criteria.

From an aesthetic point of view, deep, crystal-clear water where it is possible
to see the bottom in 30 to 40 ft. of water is highly desirable; however, a lake with
this type water is generally so nutrient-poor that fish growth is seriously impaired
and, as a result, a poor fishery is present. It is readily apparent in both Wisconsin
and especially in Texas that a lack of high degree of water clarity is not a signifi¬
cant detriment to extensive recreational use of waters.

An important difference between Wisconsin lakes examined by Sawyer
and many Texas impoundments is the turbidity of the water. In Wisconsin
high water turbidity (reduced water clarity) is generally due to algal growth.
In Texas, however, some impoundments have high level of turbidity due to ero¬
sion al material. This background turbidity permits a higher level of algal growth

CRITICAL LEVELS OF PHOSPHORUS AND NITROGEN

349

without serious impairment of water quality which would alter water-use patterns
by the residents of the area. As discussed in the subsequent section, the erosional
turbidity also affects nutrient load-lake response relationships in several additional
ways.

It is clear from the information available that one of the first steps that must
be taken to develop meaningful aquatic plant nutrient criteria for Texas surface
waters is a study of the relationship between algal and other aquatic plant prob¬
lems in Texas impoundments and the use of this water for municipal and indus¬
trial water supplies and recreational activities. This study should focus on the
question of what is the aquatic plant nutrient load that causes a significant water
quality deterioration to the water users of the area. This type of information is
not available today, and it is certainly inappropriate to apply Wisconsin criteria
to Texas impoundments.

Cycling and Transport of Nutrients

The Sawyer aquatic plant nutrient criteria are applicable to Wisconsin lakes
nutrient concentrations at ‘ice out’ in the spring. It is inappropriate to apply
these criteria to nutrient concentrations in Wisconsin lakes at other times of the
year without a careful review of the characteristics. At the ‘ice out’ period in
Wisconsin lakes and in other North American lakes that experience extensive ice
cover, the nutrient concentrations are at a maximum in their available form. On
the other hand, warm-water lakes such as Texas impoundments do not experience
the ice cover and, therefore, do not have the long winter period of slow mineral¬
ization under the reduced light conditions which bring about the accumulation
of nutrients in the spring of the year. In general, this could mean that the overall
cycling of nitrogen and phosphorus in Texas impoundments would tend to be
faster, and therefore, with those waters where the morphology, thermal regime,
and hydrology are favorable, the lake would tend to utilize these nutrients to a
greater extent and produce more algae per unit mass of nutrient input. However,
to counter this is the fact that each time a unit mass of nutrient passes through
the aquatic plant system, a certain fraction becomes refractory and, therefore,
unavailable for future growths of algae.

In September 1973, an international workshop on warm-water lakes was
held in Israel where it was concluded by the participants that warm-water
lakes, such as those in Israel, Africa, and the southern U.S., utilize their aquatic
plant nutrients differently than the cold temperate lakes, which have generally
been studied in previous limnological investigations. There was general agree¬
ment that there was an urgent need for studies on nutrient flux-algal response
relationships in warm-water lakes. The author fully supports this conclusion,
and, based on his review of the information available on Texas impoundments,
an extensive series of studies on nutrient transport and cycling must be made to
determine how much algae and other aquatic plants develop per unit mass of
aquatic nutrient input. If Texas is to properly manage these waters, considerable

350

THE TEXAS JOURNAL OF SCIENCE

amounts of federal and state funds will have to be appropriated to conduct
the studies necessary to develop technically sound management policies.

Turbidity, Water Quality and Algal Growth

In addition to affecting how the public perceives algal growth in Texas
impoundments, the turbidity due to erosional materials in these waters could
have a pronounced effect on how a certain nutrient load affects algal growth
in the impoundment. Turbidity affects light penetration with the result that
less algal growth will occur per unit mass of nutrients in turbid lakes and im¬
poundments than in clear lakes. Also, the turbidity typically present in Texas
impoundments would tend to make nutrients less available by sorption of
certain forms of the nutrients onto the surface of the solid particles. In
addition, a substantial part of the nutrient load that enters Texas impound¬
ments is in a particulate form, an unknown part of which is available for algal
growth. There is need for a study on Texas impoundments in order to determine
what fraction of the total nitrogen and phosphorus entering the waterbody can
become available for algal growth and how much of this available nutrient is
actually utilized within the water body.

CONCLUSION

Based on the experience of the author, after review of the information
available on aquatic plant nutrient water quality problem response in lakes
and impoundments throughout the U. S., it is concluded that there is insuf¬
ficient information available today to establish critical nutrient concentrations
or loads for Texas impoundments. Because of the importance of these criteria
in developing water quality management plans for these waters, it is recom¬
mended that no attempt be made to develop criteria at this time without add¬
itional study. These criteria must be based on the situations that develop in
the Texas impoundments. Consideration must be given to the public’s res¬
ponse to water quality in this region, the recreational and other uses of the
waters as a function of algal and other aquatic plant growths, and the limno¬
logical and environmental chemical aspects of nutrient transport and cycling
within these waters.

WOLVES, COYOTES, DUCKS, AND HYBRIDISM

by BOB H. SLAUGHTER

Shuler Museum of Paleontology ,

Southern Methodist University,

Dallas 75221

INTRODUCTION

We of the Texas Academy of Science are indeed fortunate to have the Dialectic
section in our journal where we may share very speculative thoughts. In keeping
with this “food for thought” vein , I wish to present a possible triggering mechanism
for hybrid swarming. One of the most fascinating examples of range adjustment
and/or usurpation, in my opinion, is that of the American Red Wolf, Canis niger.
During the early part of this century, the coyote {Canis latrans ) and the Red Wolf
had a minimum of range overlap. The coyote prefers the more-or-less open prairies
and the Red Wolf the more humid woodlands of the southeast.

In 1962, McCarley called attention to the replacement of true Red Wolves in
much of East Texas by forms intermediate between and presumably hybrids be¬
tween them and the coyotes. Paradise and Nowak (1971) developed a technique
to distinguish Red Wolves from coyotes on the basis of skulls and then proceeded
to evaluate the taxonomic status and adjusted ranges of Red Wolves. This was based
on the extensive collections at the U.S. National Museum, Washington, D.C.

The changing zoogeography of this endangered species was plotted (see Figure 1)
as follows: In the northern and southern (coastal) areas of range abuttment be¬
tween the 2 species, little or no hybridization occurred prior to 1930, and a
pocket of nothing but hybrids separated the pure coyotes’ range from that of
the Red Wolf in southcentral Texas. Sometime after 1932 and at least by 1942
the population of hybrids had expanded northward and eastward until it had re¬
placed pure Red Wolf populations in Oklahoma, Arkansas, and eastcentral Texas,
leaving the only areas containing pure Red Wolves in southeast Texas, Louisiana,
and possibly eastern Missouri. Range retraction of Red Wolves continued accom¬
panied by replacement by hybrids until 1963-1969 when pure Red Wolves could
be found only along the Gulf Coast in extreme southeast Texas and Louisiana.
Before the work of Paradiso and Nowak (1971), it was recognized that hybrids
had been replacing Red Wolves in East Texas but it was generally considered inter¬
breeding of coyotes, that were expanding their range eastward, with the resident
Red Wolves. It is now apparent, however, that interbreeding was not taking place
in the northern and extreme southern portions of the species’ range abuttment.
Instead a viable population of hybrids (hybrid swarm) that probably had its origin
Accepted for publication: October 15, 1975.

The Texas Journal of Science, Vol. XXVIII, Nos. 1-4, March, 1977.

352

THE TEXAS JOURNAL OF SCIENCE

^C.n.ger
C.latrans
hybrid

Figure 1. For precise numerical data and records by date and counties, see Paradiso and
Nowak (1971), from which these maps were extrapolated.

WOLVES, COYOTES, DUCKS, AND HYBRIDISM

353

in southcentral Texas was either usurping the Red Wolves’ range or was rapidly
replacing the Red Wolf as its numbers decreased. This probably resulted from man-
induced changes in the range. The hypothesis seems well founded but as such
events are very rare, the intriguing question concerns the triggering mechanism
of such massive interbreeding in certain areas. The creation of these hybrid swarms
is rare, but not because hybrids are inferior and unable to compete with parent
types. For reasons of brevity, I shall not go into such things as hybrid vigor but
it is well known that hybrids are often very competitive. Such happenings are rare
because mating behavior is broken down and the hybrids do not continue to have
genetic input to either of the parent populations. If, on the other hand, for one
reason or another, an area produced an abnormal number of hybrids at a given
time, those very competitive individuals could establish a very adaptive and therefore
competitive population. But what would cause the sudden increase in hybridization?

Several years ago I began to observe ducks on a small (3 acres) artificial lake
in Dallas, Texas, that is completely surrounded by homes. The fauna of the lake
included a flock of 9 white Peking ducks, added 1 or 2 at a time after they grew
out of the cute Easter duckling stage and a flock of 8 wild mallard ducks that
had stopped by during fall migration and, no doubt enjoying the safety, stayed
on to become permanent residents. The mallard ducks and the Pekings did not
integrate, either socially or sexually. One Peking, that turned out to be a hen, had
been ostrasized by the flock, apparently because of its runtiness and/or lack of
feathers on the back of its head and neck.

With the introduction of the plastic loop 6-pac holders, that annually kill
thousand of water fowl (mostly ducks and gulls), the ducks of this lake began
to have the same problems of all other lake fowl. The dastardly “loops” that
find their way into lakes and streams float and hold a deadly fascination to surface
swimming birds. They peck at them until 1 or more loops are around their necks.
Attempt at removal often winds additional loops either around the necks or feet
until the duck either strangles, or is rendered easy prey, or starves. I captured all
of these unfortunates that I could and removed the plastics. However, 1 handsome
drake elluded me for over a week until he^was starved down and could no longer
offer resistance. During this time, his antics had caused him to be ostrasized by
the remaining mallards and he had taken to swimming with the ostrasized Peking
hen. After removal of the plastic, he remained with the white hen.

The following spring, one clutch each was produced by the Peking flock and
the mallard flock as usual. Also, as usual, all ducklings were gone within a week
due to predation by local domestic cats. The ostrasized couple also produced a
clutch, 2 drakes of which survived. For some unknown reason, the following
fall, 1 of the 2 mallard hens took up with the ostrasized Peking hen and mallard
and the two hybrid male offspring of the previous season. So, within 2 years the
“ragtag” flock surpassed both the mallard and Peking flocks which had remained
stable due to unsuccessful survival of offspring. I hesitate to speculate on the
reason for the apparent more successful survival of cat preditation although I am
tempted to think along the lines of “hybrid vigor” and other rather vague adaptive
phenomena.

354

THE TEXAS JOURNAL OF SCIENCE

Could it be that some sudden event causing an abnormal injury rate , not affecting
viability, caused enough individuals of coyotes and Red Wolves to be ostrasized
and cling together and start a hybrid population, or “swarm”? Increased use of
steel traps by government trappers might be one candidate.

There are other possible reasons for such an event. Habitat changes in the area
due to farming techniques, dam building, etc., but such changes were taking place
all along the coyote -Red Wolf range abuttment and mixing was minimal. Apparently
swarming did not take place during this period, while the area was inhabited by
Indian populations, or before man moved into this area. The triggering mechanism
(unidentified as of now) for the production of the hybrid swarm appeared some¬
time during the 3rd decade of the 20th century. In any case, these observations
are offered as “food for thought” and with an invitation to comment or share
the reader’s own observations and ideas on this fascinating subject.

LITERATURE CITED

McCarley, Howard, 1962-The taxonomic status of wild Canis (Canidae) in southcentral
United States. Southw. Nat., 7(3):227.

Paradiso, J. L., and Ronald M. Nowak, 1971 -A report on the taxonomic status and distri¬
bution of the Red Wolf. U.S. Fish and Wildlife Service, Special Scientific Report, No. 145,
36 pp.

Notes Section

355

PROCOELUS VERSUS OPISTHOCOELUS VERTEBRAE - Evolutionary
selection for procoelus or opisthocoelus vertebrae (ball-and-socket) over amphicoelus
(concave-concave) or platycoelus (flat ended) vertebrae is obviously to increase flex¬
ibility. Selection for procoelus over poisthocoelus, on the other hand, is not so easily
explained. The vertebrae of modern crocodillians are procoelus (i.e.. ball-and-socket
with ball to the rear). They are aquatic and use their tails for propulsion through the
water. Troxell (1925) analyzed the mechanics of crocodile vertebrae and concludes the
procoelus has 2 distinct advantages over opisthocoelus vertebrae for this function. One,
although not the first presented by Troxell, is that the column of procoelus vertebrae
allows greater action for the ammount of angular movement. He states that bending
a procoelus vertebral column describes an arc of smaller diameter than bending a column
of opisthocoelus vertebrae. Our analysis disagrees with Troxell’s and suggests that they
produce arcs of equal radii.

The other advantage offered by Troxell, protection of the cup, appears valid. When
a crocodile is swimming forward, the tail is the means of propulsion and the means of
force transmission. Analysis of the procoelus caudel vertebrae demonstrates that the
propulsive force is transmitted along its axis to the caudel next most anterior, which in
turn duplicates the action by transmitting the received force along its axis to the caudel
in front of it. Figure 1 shows forward force transmission in procoelus vertebrae versus
that of opisthocoelus vertebrae. Force transmission from the cup onto the ball appears
mechanically more sound than transmission from the ball into the cup. The danger of
shearing the rim of the cup is increased if the cup is the receiver. Our investigation of
this possibility involved several alligator vertebrae mounted on epoxy bases at various
angles. They were tested for fracture tendencies under loads from a hydrolic press. The
results demonstrate that a configuration where force is directed axially from the cup
onto the ball can withstand 3 times greater pressure than one where force is transmitted
from the ball into the cup. When the ball was advanced axially into the cup, the cup’s
rim sheared off. On the other hand, when the cup was advanced onto the inclined ball
50% more strokes of the hydrolic pump were required to break the vertebra. Furthermore,
when it did break, it broke in the center, (compressibility of the green bone made
accurate recording of the absolute force required impossible)

Animals with relatively long necks (sauropod dinosaurs, camels, giraffes, etc.) which
hold their heads erect or nearly so, commonly have opisthocoelus cervical vertebrae. Even
bird’s cervical vertebrae might be considered opisthocoelus although the “ball” is more
of a cylinder. These animals offer an interesting analogy with the mechanical analysis
of crocodile vertebrae. In this case, the force acting along the cervical vertebrae is the
acceleration of gravity which is directed down the column (to the rear) toward the body.
As in the crocodillian tail the force is transmitted axially through the cup onto an
inclined ball.

Our conclusion is that selection for opisthocoelus, or procoelus vertebrae is dependent
upon the direction of forces applied to the vertebral column and that protection of the
cup rim is the primary concern.

LITERATURE CITED

Troxell, E. L., 1925 -Mechanics of crocodile vertebrae. Bull. Geol. Soc. Amer., 36:605.
-William D. Gosnold and Bob H. Slaughter, Shuler Museum of Paleontology , Southern
Methodist University, Dallas 7522L

356

DIRECTION

OF

FORCE

THE TEXAS JOURNAL OF SCIENCE

A

OPISTHOCOELUS VERTEBRA

Figure 1. Schematic drawing showing opisthocoelus vertebrae (A) and procoelus ver¬
tebrae with force transmission from the same direction.

A NOTEWORTHY RECORD OF THE SILVER-HAIRED BAT IN SOUTH¬
EAST TEXAS — On the evening of 24 September 1975, a silver-haired bat, Lasiony-
cteris noctivagans, was captured in a residential area near the intersection of 69th Street
and Stewart Road in the city of Galveston, Galveston County, Texas. The specimen
(Texas Cooperative Wildlife Collection No. 29002) was an adult female weighing 13.6 g.
External measurements in mm were: total length, 101; tail, 40; hindfoot, 10; ear from
notch, 16; forearm, 42.

NOTES

357

This is the southeasternmost record of L. noctivagans in Texas and the first occurrence
of the species in the Gulf Coast prairie and marsh region of the state. McCarley (1959,
Texas J. Sci., 15:385) did not report the silver-haired bat as part of the East Texas bat
fauna. The nearest records to the Galveston locality are from Medina County, 250 mi
to the west on the Edwards Plateau (Blair, 1952, Texas J. Sci., 4:95), and from Winn
Parish, approximately 300 mi to the northeast in the pine-hardwood forest of north-
central Louisiana (Lowery, 1974, The Mammals of Louisiana and its Adjacent Waters,
Louisiana State Univ. Press). Other documented localities from Texas include those
from Lubbock, Hockley, and Culberson counties (Davis, 1975, The Mammals of Texas,
Bull. 41-rev., Texas Parks and Wildlife Dept.).

According to Davis ( op . cit.), L. noctivagans is migratory, spending the summer in
northern latitudes and wintering toward the south, but wintering locations are largely
unknown. All specimens taken from Texas, except for 17 June males captured in the
Guadalupe Mountains by LaVal (1973, Southwestern Nat., 17:357), probably represent
migrating individuals (Davis, op. cit.). The Galveston specimen was probably also migrating;
this was suggested by its September occurrence in an atypical environment south of its
normal range and by heavy layers of fat found on the specimen. The discovery of l.
noctivagans on the Texas coast during September and a recent January record of the
species in the San Carlos Mountains of Tamaulipas, Mexico (Schmidly,/« Press, J. Mamm.)
may indicate that certain populations migrate through Texas to winter in the forested
regions of Mexico.

I would like to thank David J. Schmidly for reviewing the manuscript and making
helpful suggestions. Special thanks goes to Bill Proctor of Galveston, Texas, who cap¬
tured the specimen and donated it for my examination.- Chester O. Martin, Environ¬
mental Resources Branch, U.S. Army Corps of Engineers, Galveston District, P.O. Box 1229,
Galveston 77550.

OCCURRENCE OF STRIPED BASS (MO RONE SAXATILIS) IN COASTAL
WATERS OF TEXAS— A commercial net fisherman caught a striped bass in the
Gulf of Mexico surf 1-1/4 mi east of the North Galveston Jetty, Texas, on 21 January
1975. He sold the fish to James Hornbeck, the proprietor of Milt’s Seafood (Port Bolivar,
Texas), who notified officials of the Texas Parks and Wildlife Department. The authors
examined the specimen.

The dressed weight was 5.35 kilograms (14 pounds). The total length was 80.5 cm
(31.7 in); standard length was 68.0 cm (26.8 in). Spine and ray counts were; first dorsal
dorsal IX; second dorsal 1,12; anal III , 11; pectoral 15 ; pelvic 1,5. The testes are preserved
at the Parks and Wildlife Department Laboratory, Seabrook, Texas. We examined scales
from its left side near the tip of the pectoral and above the lateral line. There appeared
to be 6 to 7 “annuli” on each scale.

To our knowledge this is the second authentic occurrence of a striped bass in the
coastal waters of Texas. E. G. Simmons (Parks and Wildlife Department, Rockport,
Texas, personal communication) examined a 3 lb specimen caught off the south jetty
at Port Aransas, Texas during the late 1960s. Talbot (1966, Araer. Fish. Soc., Spec. Publ.
No. 3:37) reports the occurrence of striped bass in major streams of the Gulf of Mexico
from the Florida coast to the Tchefuncta River, Louisiana. Baughman (1950, Texas J. Sci.,
2:242) reported that its presence in Texas is doubtful; however, he cited Taylor (1878,
Forest and Stream, 2:7) who reported Roccus saxatilis in Texas. It is not listed by Hoese
(1958, Pubis. Inst. Mar. Sci., 5:312) and Parker (1965, Pubis. Inst. Mar. Sci., 10:201).

358

THE TEXAS JOURNAL OF SCIENCE

Parks and Wildlife Department biologists have stocked fry in fresh waters of Texas
since 1967 and the Louisiana Wildlife and Fisheries Commission stocked fry in Lake D’
Arbonne and Toledo Bend (Hughes and Walker, 1974, La. Conservationist, 26:18 ).-R.L.
Benefield, A. W. Moffett, and L. W. McEachron, Parks and Wildlife Department, Seabrook
77586.

A SURVEY OF ECTOPARASITES OF HARES AND RABBITS IN GRANT
COUNTY, NEW MEXICO- In 1934, Ward (1934, Proc. Okla. Acad. Set, 14:31)
published a study of rabbit parasites in central Oklahoma. Randolph and Eads (1946,
Ann. Entomol. Soc. Amer., 39:597) and Stannard and Pietsch (1958 , Nat Hist. Sur. Div.
Biol., Notes, 38) listed some ectoparasites from Sylvilagus Floridanus in Texas and Lee
County, Illinois, respectively. In New Mexico, Morlan (1955, Texas Rep. Biol. Med., 13:93)
mentioned fleas from Lepus and Sylvilagus taken during a survey of mammals of Santa
Fe County. Likewise, Kartman (1960, Zoonoses Res., 1:1) included information on the
ectoparasites of both these lagomorpha in eastern New Mexico.

The purpose of this study was to examine ectoparasites of Lepus and Sylvilagus in a
different region and environment in southwestern New Mexico. Twospecies of lagomorph,
Lepus californicus Gray, the blacktailed jack rabbit, and Sylvilagus auduboni (Baird), the
desert cottontail, were collected from the Upper Sonoran life zone of Grant County, New
Mexico. The jack rabbit is associated with grasslands and is also found in areas of pinon-
juniper and of oak woodland. The cottontail is more abundant in woodlands of the foot¬
hills and elevated mesas in the central area of Grant County. Twenty -eight cottontails and
14 jack rabbits were collected and examined for ectoparasites. over a span of 8 months at
localities selected on the basis of relative host population densities and distributions. The
parasites were preserved in 95% alcohol, treated with 10% potassium hydroxide, 0.02 N
HCL, and beechwood creosote then mounted in Canada balsam.

Ninety-eight percent of the lagomorpha were infested with ectoparasites. Five species
of fleas, 3 species of ticks, and a single bot-fly larval species were recovered. The fleas
Cediopsylla inaequalis inaequalis (Baker) and Hoplopsyllus glacialis a f finis (Baker) were
recovered from 2 and 26 desert cottontails, respectively. E chidnophaga gallinacea (West-
wood) which is usually found on birds, and Meringis dipodomys Kehls, common on wild
rodents, proved to be incidental fleas of this species of rabbit. Thirteen H. glacialis
affinis and 5 E. gallinacea were taken from 6 jack rabbits. Adult Haemaphysalis leporis-
palustris (Parckard), was found on only 2, or 7%, of thejack rabbits but was more abundant
on cottontails (21%). However, 13, or 93%, of thejack rabbits examined were infested
with the ticks Dermacen tor parumapertus Neumann. Two adults and 1 nymph of Otobius
megnini (Duges) were collected from 2 different cottontail hosts but were not found on
jack rabbits. Only 5 larvae of Cuterebra were taken from 3 different cottontail host.
Based on distribution records, these bot-fly larvae are probably C. cuniculi (Clark) or
C. leporivora Coquillett. These observations compare favorably with results obatined by
Ward (1934) in Oklahoma, Randolph and Eads (1946) in Texas, Morlan (1955) in Santa
Fe County, New Mexico, and Stannard and Pietsch (1958) in Lee County, Illinois.

The author wishes to thank Professor C. Clayton Hoff for his advice and suggestions
during this study. - Paul H. Rodriguez, Division of Allied Health and Life Sciences,
University of Texas at San Antonio, San Antonio 78285

A KI AM INCISED POT FROM LOUISIANA - The Central Louisiana Electric
Company cleared for a new power line right-of-way last year on the Louisiana side of

NOTES

359

Toledo Bend lake and bulldozed out a crushed Indian pot. Mrs. Melva L. Marsh re¬
covered the pieces. The John Guy fmaily at Anacoco put me in contact with Mrs.
Marsh and I undertook reconstruction of the pot with fair results and a lot of experi¬
ence. About 2/3 of the vessel was present.

The pot was found on a hilltop in Sabine Parish. Coordinates are 45.1 NS-346.2
EW, U. S. Geological Survey, Alexandria Series, V502; Add. IAMS, NH 15-2. There
was no indication of a burial present but occupational debris is scattered over the surface.
It is unusual to find this large a vessel in this much completeness.

The restored is 37.46 cms high (14.75 in). The mouth is 33 cms (13 in) on the out¬
side. The greatest width is 38.1 cms (15 in), 17.78 cms (7 in) above the base. The bottom
is slightly concave and is 12.7 cms (5 in) wide. The neck is slightly flared. The capacity
is 25.12 1 (6.64 gal.). The color is reddish brown with darker areas near the base.

The temper is clay and grit with some fiber. Thickness of the wall averages 7.94 mm
(5/16 in). The rim is rounded and flush with the sides.

The decoration consists of irregularly incised lines around the neck. Punctations,
made with a small wedge-shpaed object, begin 12.7 cms (5 in) above the base and cover
the whole pot up to the neck. The finish is fine on the lower part and coarser above.

According to Suhm and Jelks (1962, Editors Handbook of Texas Archaeology:
Type Descriptions., The Texas Archaeological Society and The Texas Memorial Museum,
Austin, Texas), this vessel is a Kiam Incised pot belonging to the Alto Focus and dated
between 500 and 1000 A. D.. McClurkan, Field, and Woodall (1966, Excavations in
Toledo Bend Reservoir, 1964-65., Papers of the Texas Archaeological Salvage Project,
No. 8, Austin, Texas.), found sherds of Kiam Incised at the Salt Lick Site, 7 mi west of
Many, also in Sabine Parish.

The vessel is in the personal collection of Mrs. Melva L. Marsh, RR3, Box 62, Florien,
Louisiana, 71429. Photographs by Steven Lewis. -Russell J. Long, Department of
Biology, Lamar University, Beaumont 77710.

Figure 1. Kiam incised pot. A, detail of neck; B, detail of body.

360

THE TEXAS JOURNAL OF SCIENCE

RESOLUTION OF 2-(0-CHLOROPHENYL)-2-(P-CHLOROPHENYL) ACETIC

ACID— Mitotane is best known by its trivial name, o,p’DDD, and is chemically 1,1— di-
chloro-2-(o-chlorophenyl)-2-(/?-chlorophenyl)ethane. This drug is used in the treat¬
ment of inoperable adrenal cortical carcinoma of both functional and non-functional
types (Hutter, 1966, Amer. J. Med., 41:575).

Mitotane can best be described as an adrenal cytotoxic agent, although it can cause
adrenal inhibition, without cellular destruction. Its biochemical mechanism of action is
unknown. Mitotane has been shown to inhibit adrenal glucose- 6-phosphate dehydrogen¬
ase in the dog and corticotropin stimulation of steroidogenesis in vivo but not in vitro
(Hart, 1971, Biochem. Pharmacol., 20:1679; Grady, 1965, Proc. Soc. Exp. Biol. Med.,
119:238). These findings suggest that a metabolite of miotane rather than the compound
itself is responsible for its activity. The major metabolite of mitotane appears to be 2-
(o-chlorophenyl)-2-(/?-chlorophenyl) acetic acid,o,pT)DA (Sinsheimer and Guilford, 1972,
J. Pharm. Sci., 61:314). We wish to describe an improved preparative method for the
synthesis of 2-(o-chlorophenyl)-2-(p-chlorophenyl) acetic acid, o,/?’DDA, and its chemical
resolution.

A mixture of 80 ml diethylene glycol and 10.0 g of 1 ,l,l-trichloro-2-(o-chlorophenyl)-
2 -(p-chlorophenyl) ethane was added to a solution of 12.6 g of KOH in 7 ml of water.
The mixture was stirred for 4 days at 110 C. The mixture was allowed to cool and was
poured, with vigorous stirring, into 250 ml of cold water. The insoluble material was filtered
and washed twice with 10 ml portions of warm water. The filtrate was boiled gently for
5 min with 2 g of decolorizing carbon; the carbon was removed by filtration. The filtrate
was acidified to litmus with 20% H2SO4 ( ca 50 ml), and then an additional 6 ml of the
acid was added. The mixture was cooled in an ice bath for 15 min. The precipitate was
collected by suction filtration, washed free of sulfate ions with water, dried at 85 C, re¬
crystallized from methanol, and gave 3.5 g of product (44% yield, mp 104-105 C). The
old method involved a Ba(OH)2 hydrolysis (Cristol and Haller, 1945, J. Am. Chem.
Soc., 67:2222) and gave a yield of 13% (lit. mp 107-108 C corrected).

o,p' DDA (2.0 g) was dissolved in 40 ml of ethanol with warming to 50 C. Quinine
(2.3 g) was added over a period of 10 min. The solution was heated to boiling and 25 ml
of water was added (to turbidity). The mixture was allowed to cool to room temperature
and stand for 2 days. The product was re crystallized from methanol-water to constant
rotation, air dried at 60 C to yield 1.3 g, mp 156-157 C; (0) j}5 (ethanol)-22.5 (c 2.48).

A 2-ml portion of concentrated HC1 was added to a mixture of 1.3 g of the diaster-
eoisomer and 40 ml of water. The solid precipitate was filtered and washed with 5 ml
portions of 50% HC1, re crystallized from ethanol to constant rotation, air dried and then
dried at 80 C to give 0.470 g of o,/?’DDA, mp 105-106 C, (a) j^5 (chloroform) +11.5
(c 1.00); Umax (KBr) 3000-2500 (OH 1710 (C-O) cm-1. The essential data for racemic
o,p’DDA (III), appear in Table 1.

TABLE 1

Intermediates in the Resolution of 2-(o-Chlorophenyl)-2-(p-Chlorophenyl) Acetic
Acid and (-)-2-(o-Chlorophenyl)-2-(p-Chlorophenyl) Acetic Acid.

No.

M.P. C

Formula

c

H

N

c

H

N

I

104 - 1051 2

c14h10o2ci2

59.80

3.58

59.90

3.60

-

II

156-157

C34H34N204C12

67.43

5.66

4.62

67.52

5.56

4.51

III

105 -106

C14H10O2C12

59.80

3.58

l ■ p n

59.89

3.52

-

1. Melting points are uncorrected.

2. A mixed melting point with an authentic sample showed no depression.

NOTES

361

This work was supported by the National Institutes of Health (Grant RR-08061)
through the National Cancer Institutes-/. Guilford, E. Hickman^ and D. Ghosh, Texas
Southern University, School of Pharmacy, Houston 77004.

OBSERVATIONS ON THE ECOLOGY OF MICROPTERUS TRECULI IN
THE GUADALUPE RIVER— Hook and line collections and skindiving observations
were made of Micropterus treculi in approximately 28 km of the Guadalupe River from
State Highway 473 (Sultanfuss crossing) to U.S. Highway 281 bridge, Kendall County,
Texas, March 12 through 15, 1975. A representative sample of the catch was catalogued
in the Oklahoma State University zoological museum following confirmation of identifi¬
cation by R.J. Miller.

During the daytime hours of this period, surface water temperatures ranged from 14
to 17 C and the maximum air temperature was 26 C under partly-cloudy skies. Under¬
water visibility estimates ranged from approximately 1 m in mainstream areas to 6 m
in a spring-fed tributary stream. In upstream areas M. treculi were taken on spinners in
riffle areas and swift water. Farther downstream where riffles were widely separated by
deep pools, Guadalupe bass were caught in deep eddies in riffle tail races. Largemouth
bass, Micropterus salmoides were taken only in deep pools, while no spotted bass, Microp¬
terus punctulatus, were caught. Hurst, Bass and Hubbs, 1975, Natl. Symp. on the Biol,
and Mgmt. of the Centrarchid Basses, 47-53, reported that “trophy” sized Guadalupe
bass would be about 250 mm SL. This is approximately 310 mm or greater. Two Guadalupe
bass were taken having a TL of 330 mm. The authors are aware of no information avail¬
able on the movements of M. treculi. However, Gerking, (1953, Ecology, 34:347) reported
a home range of 200 to 400 linear ft (approximately 61-122 m) for the closely related
spotted bass M. punctulatus in a small Indiana stream. The stretch of river from which
these specimens were taken was approximately 12.9 km to 40 km upstream from Canyon
Reservoir. It therefore appears likely that the large specimens taken in our study were
lentic residents and not upstream migrants which might have exhibited accelerated growth
rates in a newly formed reservoir.

With 5 of 7 and 2 of 13 male M. treculi caught, it was possible to cause apparently viable
sex products to be emitted from the genital openings by application of gentle pressure
to the abdomen. Only 1 of 4 female and none of 4 male largemouth bass taken in the same
period were similarly determined to be in “ripe” spawning condition. Hurst, Bass and
Hubbs (1975) suggested a May through June spawning period for M. treculi based upon
size-class evaluations and the presence of large ova in a 70 mm SL female collected in
May. Our results, however, indicate a spawning period for large-sized Guadalupe bass be¬
ginning much in advance of this time.

In skindiving observations of several mainstream areas and the lowest pool areas of 3 tribu¬
tary streams, no large-mouth or spotted bass nests were found. However, 1 actively guarded
M. treculi nest was observed. The individual guarding this nest was approximately 280 mm
TL and moved a short distance away when approached. A second individual, thought to
be a spawning female was seen nearby beneath an overhanging creek bank. The nest site
was located approximately 75 m upstream of the mouth of a springfed tributary at a depth
of 69 cm. It was situated 1 m from the shore on a 5 to 10 degree slope with the current
speed at this point being approximately 0.3 m per second. The nest was an oval shaped,
bowl-like depression 51 cm by 43 cm and 10 cm in depth. It was swept into the hard black
soil of the creek bank and was free of silt. The interior of the nest was lined with 5 cm
diameter limestone rubble which was partially covered by a few scattered sticks and leaves.
The eggs within the nest were adhered to the sticks and leaves with only a few on the tops
and sides of the rubble. All eggs were collected in a cloth bag and preserved in 10% formalin.
Later enumeration showed the nest to contain 1,406 eggs. This was fewer than the 5,016

362

THE TEXAS JOURNAL OF SCIENCE

average number of eggs reported by Vogele (1975 , U.S . Fish and Midi. Serv. Tech. Pap. 84: 1)
for nests of spotted bass in Bull Shoals Reservoir, Arkansas. The average diameter of 200
eggs from this M. treculi nest was 2.058 mm with a standard deviation of .078 mm. This
was larger than the 1.93 mm average diameter reported by Vogele (1975) for eggs collected
from nests of M. punctulatus. Stomachs of 5 M. treculi were preserved and later examined.
Three were empty, 1 contained an immature aquatic insect, and the remaining 1 contained
1 crayfish and 2 partially digested Notropis sp.-R.L. Boyer, G. W. Luker, and R.J. Tafanelli,
School of Biological Sciences, Oklahoma State University, Stillwater 74074.

AN AMBICOLORATE WINDOW? ANE ( SCOPHTHALMUS AQUOSUS ), WITH
NOTES ON OTHER ANOMALOUS FLATFISH— An almost totally ambicolored
windowpane ( Scophthalmus aquosus-, Texas Cooperative Wildlife Collection 0419.1;
130 mmTL, 101 mm SL, 22 g) was trawled off Waccamaw Sound, South Carolina 16 May 1973
(Figure 1). Pigmentation similar to that of the ocular side covers approximately 75% of the
right- or blind-side surface, with a majority of the head region, proximal portions of the
first 8 dorsal fin rays and a wedge-shaped area extending dorso-posteriad from above the
preopercle to the vicinity of the 27th dorsal fin ray exhibiting typical blind-side coloration.
Premaxillary and dentary bones, exhibiting faint coloration on anteriormost portions, are
the only regions of the head bearing pigmentation. Comparison of eyed- and blind-side
coloration indicates no symmetry of pigmentation patterns.

Our specimen represents the first recorded instance of abnormal coloration in Scoph¬
thalmus aquosus. Pigment abnormalities in other species of the genus Scophthalmus are
common (Hussakof, 1914, Bull. Am. Mus. Nat. Hist. 33:95; Norman, 1934, British Mus.
Nat. Hist., 1:22), especially among European species and particularly turbot (<S. maximus).
The piebald condition of the lower surface is so common among turbot that it was consid¬
ered by early French zoologists as characteristic of the species (Giard, 1892, Compt. Rend
et Mem. Soc. Biol., 9(IV):31). Norman (op. cit.) found that although ambicolorate examples
have been recorded for a number of scophthalmids, color anomalies seem to be much more
common in certain species than in others. Norman demonstrated that total ambicoloration
is commonplace in turbot and generally rare in brill (S. rhombus ), a sympatric species.
Our specimen increases the number of bothids exhibiting abnormal pigmentation and
strengthens Norman’s conclusion that ambicoloration is rarer in some species than in others.

The remainder of this paper describes several examples of pigment and associated morph¬
ological abnormalities in southern flounder ( Paralichthys lethostigma ) taken from Texas
waters. A partially albinistic southern flounder (TCWC 4032.1; 255 mm TL, 205 mm SL,
168 g) was collected 3 December 1968 from the cooling-water intake canal of Houston
Lighting & Power Company’s P. H. Robinson Generating Station on Galveston Bay, Texas.
Right-side coloration is typical of the species, while the left-side of the caudal peduncle
and caudal fin is largely albinistic. Lack of pigmentation extends from the caudal pe-
uncle area 25 mm anterior to the caudal fin base and through dorsal rays 85 to 92 and anal
rays 70 to 73. Small, scattered splotches of color exist on the distal portion of the caudal
fin rays. A transitional zone of partial pigmentation exists between normal and albinistic
areas. Although no visible morphological abnormalities are evident, a radiograph (Figure 2)
does reveal osteological malformation in the hypural plate region. The proximal portion
of the first autogenous epural is lodged tightly between an abnormally thickened last neural
spine (penultimate vertebra) and a ventrally-bowed, second autogenous epural. The sec¬
ond autogenous epural and first 2 epural elements of the urostyle are broken into approx¬
imately equal halves, with the distal halves dislodged ventrally. The base of the last hae¬
mal spine appears to have been notched posteriorly by ventral flexure of the urostyle.
Location and extent of both color and osteological abnormalities in our specimen closely
resemble those described by Dawson (1967, Trans. Am. Fish. Soc., 96:400) for a 64 mm
SL southern flounder. However, Dawson’s specimen exhibits more extensive damage to the

NOTES

363

Figure 1. Right side of an almost totally ambicolored Scophthalmus aquosus (TCWC
0419.1). Left side is normally pigmented.

364

THE TEXAS JOURNAL OF SCIENCE

Figure 2. Radiograph of a partially albinistic Paralichthys lethostigma (TCWC 4032.1)
illustrating osteological damage to the hypural plate region.

posterior caudal vertebrae and caudal fin and a proportionally larger albinistic area. Our
findings and other records of albinistic flatfishes exhibiting morphological aberrations
(summarized in Dawson, 1962, Copeia, 1962:138) provide supporting evidence for Norman’s
(op. cit .) theory that partial albinism in heterosomates is generally associated with osteo¬
logical injury.

A totally ambicolored southern flounder (TCWC 0420.1; 318 mm TL and 266 mm SL)
was trawled from East Matagorda Bay, Texas 23 October 1973. This specimen fits Dawson’s
(1962, op. cit.) Type D anomaly classification in exhibiting almost total ambicoloration,
incomplete eye rotation and hooked dorsal fin. Typical pigmentation covers the entire
blind side except for outermost portions of the pectoral fin and the lower, anteriormost
point of the dorsal fin hook. The migrating eye is positioned atop the dorsal crest in an
extensive depression of the frontal region and is in the same plane as the right edge of the
dorsal fin and right nasal openings. Other atypical features exhibited by this specimen in¬
clude variation in pelvic fin length and position toward symmetry and an anomalous con¬
cave posterior half of the left lateral-line arch. A radiograph of the head region reveals
several osteological abnormalities (Figure 3). Median portions of the left frontal bone
encompassing the right orbit exhibit an incomplete ventro -lateral migration and, as a result,
are positioned along the dorsal midline. Premature arrestment of counter-clockwise twist¬
ing of this medial sector has caused the posterior oblique ridge of the right orbit to be

NOTES

365

Figure 3. Radiograph of the head region of a totally ambicolored Paralichthys letho-
stigma (TCWC 0420.1) illustrating incomplete migration of left frontal bone.

directed dorso-posteriad instead of dorso-anteriad. This incomplete deflection of the central
skull region also has inhibited forward growth of the dorsal fin and has resulted in crowd¬
ing of anteriormost dorsal fin rays and associated pterygiophores between the posterior
oblique ridge of the right orbit and the first neural spine. Although dorsal fin rays and
pterygiophores are on a 1-to-l basis, the first pterygiophore is in contact with the oblique
ridge of the right orbit and does not function in direct support of the first dorsal fin ray.
The radiograph of our ambicolorate flounder closely resembles that taken by Powell and
Schwartz (1972,7. Elisha Mitchell Sci. Soc., 83(3): 155) of a totally ambicolorate southern
flounder exhibiting hooked dorsal fin and rearward projection of the oblique crest of the left
frontal bone. These radiographs support Norman’s ( op . cit .) finding that frontal bones
arrive in final position only after eye migration is complete. Furthermore, it appears that,

366

THE TEXAS JOURNAL OF SCIENCE

although forward extent of dorsal fin growth in flatfish is dependent upon degree of eye
migration, number of dorsal fin rays and pterygiophores is determined before complete
eye rotation is attained.

An almost totally ambicolored southern flounder (TCWC 4033.1; 293 mm TL, 240
mm SL, 337 g) was taken 7 August 1972 in the Houston Ship Channel sector of Galveston
Bay, Texas. This flounder exhibits pigmentation on approximately 85% of its blind side.
The head, pectoral fin and portions of the first 13 dorsal rays exhibit normal blind-side
albinism. Although blind-side pigmentation is slightly lighter than that on the eyed side,
there is a sharp distinction between pigmented and non-pigmented areas on the lower
surface. Except for a strongly convex blind-side contour, this specimen exhibits no other
morphological or osteological abnormalities.

We wish to thank Houston Lighting & Power Company for financing the study during
which the partially albinistic specimen was collected. We also wish to thank Messers. E.A.
Bruguiere, W.L. Oliver, J. Poncik and H.A. Smith for making the anomalous specimens
available to us. -Andre M. Landry, Jr., Department of Marine Sciences, Texas A&M Univer¬
sity, Galveston 77550, and Kenneth W. Johnson, Department of Biology, Mary Hardin-
Bay lor College, Bel ton 76513.

ANALYSIS OF STUMP VEGETATION IN AN EAST TEXAS POND-Plants
growing in and around water are becoming more significant as reservoirs are proposed
and built and as more environmental impact studies are initiated. It is necessary, when
discussing the effects a particular reservoir project might have on the vegetation present,

TABLE 1

Density, Frequency and Importance Value Data for Species Found Growing on
Stumps in an East Texas Pond.

Species

Density
no /stump

Relative

density

%

Frequency

%

Relative

frequency

%

Importance

value*

Juncus repens

7.5

33.9

47.5

12.6

46.5

Ly copus rubellus

4.0

18.1

45.2

12.0

30.1

Rhynchospora glomerata

2.3

10.4

36.8

9.7

20.1

Xyris difformis

2.2

9.8

35.2

9.3

19.1

Proserpinaca palustris

.9

4.0

30.3

8.0

12.0

Ascyrum stans

.8

3.4

20.1

5.5

8.9

Bidens polylepis

.7

3.1

21.5

5.7

8.8

Aster lateriflorus

.8

3.4

19.1

5.1

8.5

Myrica cerifera

.4

2.0

23.0

6.1

8.1

Hypericum walteri

.5

2.2

17.6

4.7

6.9

Others* *

9.7

2 1.3

31.0

Total

100.0

100.0

200.0

* Sum of relative density and relative frequency.

** Other species recorded on stumps listed in order of decreasing importance value:

Andropogon virginicus, Panicum verrucosum, Pluchea foetida, Sium suave, Pinus taeda,
Hydrolea ovata, Hydrocotyle verticillata, Cephalanthus occidentalis, Scirpus cyperinus,
Cyperus haspan, Ludwigia glandulosa, Acer rubrum, Ludwigia palustris, Nymphaea
odorata, Persicaria coccinea, Amorpha paniculata, Eupatorium perfoliatum, Liquidam-
bar styraciflua, Paspalum longipilum, Rubus sp., Smilax sp., Bulbostylis ciliatifolia,
Desmodium sessilifolium, Rhus copallina, Solidago rugosa, Baccharis halimifolia, Heli-
anthus angustifolius, Lobelia cardinalis, Panicum agrostoides, Spiranthes laciniata,
Mikania scandens.

NOTES

367

to submit possible immediate or long-term changes in the vegetation of that impound¬
ment area. The object of this study was to provide information on plants likely to be
found on logs and stumps in reservoirs or other aquatic habitats.

The pond studied was located near Nacogdoches, Texas. Three-fourths of the pond
was surrounded by trees and shrubs whereas one side of the pond fronted a highway and
is characterized by herbaceous species. Imperical observation indicated that loblolly pine
(Pinus taeda L.)s sweetgum (Liquidambar styraciflua L.), black gum (Nyssa sylvatica Marsh.)
and red maple (Acer rubrum L.) were prevalent species bordering the pond while short-
leaf pine (Pinus echinata Mill.) and white oak (Quercus alba L.) were occasionally present.
Local populations of common buttonbush (Cephalanthus occidentalis L.) and wax myrtle
(Myrica cerifera L.) were also observed.

There were 161 stumps in the pond. Plants present on each stump were identified
and counted and density and frequency data were obtained. Counts for rhizomatous species
included individual shoots or plants along each rhizome. Nomenclature followed Correll
and Johnston (1970; Texas Research Foundation, Renner, Texas). Stumps were period¬
ically sampled from March to October.

Forty -one species of plants were found growing on the stumps. These species are listed
in order of importance value in Table 1. Penfound (1949, Proc. La. Acad. Sci. 12;47)
recorded 26 species of plants on logs in Lake Chicot located in Louisiana. Ten of these
species were found on stumps in our study.-E’.S'. Nixon, W.F. Trotty, B.L. Batesyand
D.L. Wilkinson, Department of Biology, Stephen F. Austin State University, Nacogdoches
75961.

The Texas Journal of Science

Index to Volume XXVIII
1977

Printed in San Angelo, Texas U.S.A.

By

The Talley Press

A

Aboul-Ela, M. M., see Aune, J.

Acer

rubrum; 367

Alexander, E. 3., see Cannon, M. S.
Amblycera; 147
Anoplura; 145
Aplysia

brasiliana ; 102

Arnold, J. B., G. B. Kegley, “Results
of a Magnetometer Survey at Hueco
Tanks, a Prehistoric Mogollon Vil¬
lage in Western Texas”; 201
Aune, J., B. J. Smith, M. M. Aboul-Ela,
“The Effect of Lysosomal Enzymes
on Chloroplasts in vitro ”; 85
Austroriparian Province; 127
auxotrophic organism; 120, 125

B

Bacillus

subtilis; 94

Bates, B. L., see Nixon, E. S.

Bayer, M., see Purdom, M. E.
Benefield, R. L., A. W. Moffett, L. W.
McEachron, “Occurrence of Striped
Bass ( Morone Saxatilis) in Coastal
Waters of Texas”; 357
Boopidae; 147
Borcher’s ash; 54, 55,59
Boyer, R. L., G.W. Luker, R. J.Tafanelli,
“Observations on the Ecology of
Micropterus treculi in the Guadalupe
River”; 361

2-Bromopropene; 219, 220, 221
Brown, R.F., “A Sodium Iodide Cen¬
trifugation Technique for Isolation
of Newly-Replicated DNA from
Rat Liver”; 209
Brueckner-Goldstone; 239

c

Caelonopsis

latus; 110, 111

Cagle, G. D., G. R. Vela, “A Simple,
Inexpensive Freeze-Etch Device
for Use in Electron Microscope
Laboratories”; 91
Cambarincola

vitrea; 285, 286, 292
Canis

latrans; 35 1
niger; 35 1

Cannon, M. S., E. S. Alexander, “The
Flower Gardens Coral Reefs”; 99
Cediopsylla
inaequalis

inaequalis; 358
Cephalanthus

occidentals; 367
Chaetodon
ay a; 101

Chang, M., “Is Canopy Interception
an Accurate Measure of Loss from
the Hydrologic Budget?”; 339
Christenson, C. P., see Schram, A. C.
Chromis

enchrysurus; 101

Chung, Kyung-Suk, “The Salinity and
Temperature Tolerance and the
Growth of Macrobrachium ohione
(Smith) 1874 Reared in Laboratory
Tanks”; 271

clinopyroxene; 54,56,59
Cosmolaelaps

passali; 110, 111

Crowder, G. A., R.W.Waltrip, “Barriers
to Internal Rotation in 2-Bromo-
propene and 2-Iodopropene”; 219
Cry oultramicro tome; 91
Cuter ebra; 358
cuniculi; 358
leporivora ; 358

D

Dermacentor

parumapertus; 358
Diploria; 100

DNA; 7, 8, 9, 10, 11, 13, 14, 15, 16, 17,
209, 210, 211, 212, 213, 215, 216
polymerase I; 10, 13, 14, 16, 17

INDEX TO VOLUME XXVIII 1977

Dorsey, S. L., “A Reevaluation of Two
New Species of Fossil Bats from
Inner Space Caverns”; 103
Dorson, J. W., R. E. Moses, “Excision
Repair Following Ultraviolet Ir¬
radiation in Toluene-Treated Esch¬
erichia coli 7

E

Echidnophaga
gallinacea; 358
Eptesicus
fuscus; 131
fuscus; 131
Escherichia
coli; 8, 193
Eunice; 101

aphroditois; 101

F

ferroaugite; 54

Flippen, J. H., see Whitenberg, D. C.
FORTRAN IV program; 72

G

Ghosh, D., see Guilford, J.

Gibson, G. E., “Seasonal Distribution
of Mites Associated with Popilius
disjunctus (IlligerXColeoptera ;Pas-
salidae) in Hardin County, Texas”;
109

Gonioplectrus
hispanus; 101

Gosnold, W. D., B. H. Slaughter, “Pro-
coelus Versus Opisthocoelus Verte¬
brae”; 355

Guilford, J., E. Hickman, D. Ghosh,
“Resolution of 2.-{o -Chlor op henyl) -2-
(p-Chlorophenyl)Acetic Acid”; 360

H

Haemaphysalis

leporispalustris; 358
Haematopinidae; 146
Hardcastle, D. L., see Saxon, G. P.
Hartree-Fock; 239, 240, 251
Hartree-Fock -Slater; 239
Haywood, C. T., “Some Comments
Concerning the Impact Ball Appar¬
atus”; 45
Heterochelytus

fusiformis; 109, 110, 111
Hickman, E., see Guilford, J.

Hicks, D. R., see Lenhart, J. D.

369

Ho, Chi-Yun, see Lenhart,!. D.

Honeycutt, R. L., see Schmidly, D. J.

Hoplopsyllus

glacialis

a f finis; 358

hornblende; 54, 55, 56, 58, 59

Huffman, D. G., see Tuff, D. W.

Huygens’ Principle; 61, 62

Hypopi; 110, 111

I

Illiger; 109

Ischnocera; 147

J

Jeschofnig, P.,“Petrography of Pleisto¬
cene Volcanic Ash from Palo Duro
Canyon, Texas”; 5 1

Johnson, K. W., see Landry, A. M.

K

Kegley, G. B., see Arnold, J . B.

Kirchoff’s surface integral; 61,62,63,
64,65,66

Koepp, S. J., E. A. Schlueter, “The
Ecology of a Branchiobdellid (An¬
nelida : Oligochaeta) from a Texas
Pond”; 285

L

Landry, A. M., K. W. Johnson, “An
Ambicolorate Windowpane ( Scoph -
thalmus aquosus), with Notes on
Other Anomalous Flatfish”; 362

Lasionycteris

noctivagans; 131, 142, 356, 357

Lasiurus

borealis; 111, 132, 142
borealis; 132
cinereus; 133, 142
cinereus; 133
intermedius; 133
florid anus; 133
intermedius; 133
seminolus; 132

Lee, G. F., “Critical Levels of Phos¬
phorus and Nitrogen in Texas Im¬
poundments”; 347

Lenhart, J. D., Chi-Yun Ho, D. R. Hicks,
“Crown Positions Within Unthinned
Loblolly Pine Plantation Canopies”;
113

370

Lepus; 358

calif ornicus; 358
Leuconostoc

mesenteroides ; 119
Linognathidae; 146
Liquidambar

styraciflua; 367

Long, R. J., “A Kiam Incised Pot from
Louisiana”; 358
Luker, G. W., see Boyer, R. L.
Lutjanus; 100

campechanus; 101

M

Macrobrachium

acanthurus; 271, 279
carcinus; 271
nipponense; 271

ohione; 271, 272, 276, 277, 278,
280,282
rosenbergii; 271
Macrocheles

tridentatus; 111
Mallophaga; 146, 147
Martin, C. O., “A Noteworthy Record
of the Silver-Haired Bat in South¬
east Texas”; 356

McEachron, L. W., see Benefield, R. L.
McLemore, E. W., I. L. Wright, “Ap¬
plication of Huygen’s Principle to
the Reflection of Seismic Waves
at a Free Surface”; 61
Meringis

dipodomys; 35 8
Megathanus

floridanus; 111
Menoponidae; 147
Micro -vertebrate
fauna; 5 1
Micropterus

punctulatus; 361, 362
salmoides; 361
treculi; 361, 362

Moffett, A. W., see Benefield, R. L.
Mollusk
fauna; 51
Montastrea; 100
Moses, R. E., see Dorson, J. W.

My otis; 103

austroriparius; 130, 133
keenii

apache; 104
lucifugus; 142
lucifugus; 130
occultus; 104, 130
magnamolaris; 103, 104, 105 , 106,
107

THE TEXAS JOURNAL OF SCIENCE

rectidentis; 103, 104, 105 , 106, 107
thysanodes; 107

velifer; 103, 104, 105, 106, 107,
108

incautus; 103
magnamolaris; 106
volans; 107
yumanensis; 104
Myrica

cerifera; 367

N

Nelson, I. V., “The Polarographic Re¬
duction of Metal Ions in Acetone
and Acetophenone in the Presence
of Lithium Perchlorate and Tetra-
ethylammonium Perchlorate”; 223
Nixon, E. S., W. F. Trotty, B. L. Bates,
D. L. Wilkinson, “Analysis of Stump
Vegetation in an East Texas Pond”;
366

Norton, S. J., see Purdom, M. E.
Nycticeius

humeralis; 133, 134, 142
humeralis; 134

Nyssa

sylvatica; 367

o

Oppelia; 110, 111
Otobius

megnini; 358

P

Paralichthys

lethostigma; 362, 364, 365
Passalacarus
sylvestris; 111
Pearlette

ash; 51,57
Pediculidae; 146
Pediococcus

cerevisiae; 119

Pettitt, J. M., “Limnological Theses
and Dissertations Concerning Texas
Waters, 1897- 1976”; 295
Phaseolus

vulgaris; 194
Philopteridae; 148
Pinus

echinata; 367
taeda; 113, 367
Pipistrellus

subflavus; 127, 131, 142
Platygonus; 103

INDEX TO VOLUME XXVIII 1977

Plecotus

rafinesquii; 136
polA ; 10, 12, 13, 17
polA?; 14, 17
Polychaeta; 101
Popilius

disjunct us; 109, 111
Porifera; 101
Pontes; 100

Prasad, J. S., A.G.Walvekar, “Markov-
Renewal Programming Under In¬
complete Information-I”; 19
See Walvekar, A. G.

Procambarus

simulans; 285, 286
Prometryne; 193, 196
Pseudomonas; 94, 96
Purdom, M. E., M. Bayer, S. J. Norton,
P. T. Sullivan, “Effects of Simple
Sugars on Certain Serum Constitu¬
ents and Erythrocyte Osmotic
Fragility in Rats”; 231

Q

Quercus
alba; 367

R

Ramos-Cormenzana, A., see Ruiz-
Berraquero, F.

Rayleigh-Schroedinger; 239, 240, 251,
252

Reduced Density -Matrix; 240, 251
Rhomboplites
aurorubens; 101

RNA; 193, 194, 195, 196, 197, 198
Rodriguez, P. H., “A Survey of Ecto¬
parasites of Hares and Rabbits in
Grant County, New Mexico”; 358
Ruiz-Berraquero, F., A. Ramos-Cor¬
menzana, “Production of Amino
Acids by Soil Microorganisms”; 119

s

Saxon, G. P., D. L. Hardcastle, “Ray¬
leigh-Schroedinger Perturbation
Calculations for Beryllium”; 239
Schlueter, E. A., see Koepp, S. J.
Schmidly, D. J., K. T. Wilkins, R. L.
Honeycutt, B. C. Weynand, ‘The
Bats of East Texas”; 127
Schram, A. C. C. P. Christenson, “Syn¬
thesis of a Radiolabeled Carrier to
Precipitate Anti-Hapten Antibod¬
ies”; 253

371

Scophthalmus

aquosus; 362, 363
maxirnus; 362
rhombus ; 362
Seiodes

trifidus; 111
Sesbania; 194

Slaughter, B. H., “Wolves, Coyotes,
Ducks, and Hybridism”; 351
See Gosnold, W. D.

Smith, B. J., see Aune, J.

Spears, L. G., “Corrosion Studies in a
Water Flood Oil Field System”; 259
Sullivan, P. T., see Purdom, M. E.
syenite

porphyry; 202, 206
Sylvilagus

auduboni; 358
Floridanus; 358

T

Tadarida

brasiliensis; 136, 137, 139, 142
cynocephala; 136, 137, 139,
140, 141

mexicana; 136, 137, 138, 139,
140, 141

Tafanelli, R. J., see Boyer, R. L.

Trichodectidae; 148

Trotty, W. F., see Nixon, E. S.

Tuff, D. W., “A Key to the Lice of Man
and Domestic Animals”; 145
D. G. Huffman, “An Index to the
Genera of Hosts in Yamaguti’s
Sy sterna Helminthum" \ 161
Tyroglyphidae; 110, 111

u

Uroobovella ; 110, 111
levis; 111
setosa; 111
spinosa; 111
Uroseius

quercus; 111
uvr; 1

uvrA ; 10, 12, 14

V

Vela, G. R., see Cagle, G. D.

w

Waltrip, R. W., see Crowder, G. A.
Walvekar, A. G., J. S. Prasad, “Markov-
Renewal Programming Under In¬
complete Inforniation-H”; 33
See Prasad, J. S.

Wevnand

372

THE TEXAS JOURNAL OF SCIENCE

Weynand, B. C., see Schmidly, D. J.
Whitenberg, D. C., J. H. Flippen, “Ef¬
fects of Prometryne on RNA and
Protein Metabolism in Bean Leaf
Discs”; 193

Wilkins, K. T., see Schmidly, D. J.
Wilkinson, D. L., see Nixon, E. S.
Wright, I. L., see McLemore, E. W.

z

Zercon

passalorum; 109, 111

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EXECUTIVE COUNCIL

President: HERBERT H. HANNAN, Southwest Texas State University

President-Elect: JAMES R. UNDERWOOD, JR., West Texas State University

Secretary-Treasurer: EVERETT D. WILSON, Sam Houston State University

Sectional Vice-Presidents:

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State University

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BOARD OF DIRECTORS

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ARTHUR E. HUGHES, Sam Houston State University
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COVER PHOTO

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TVo/e: A check must accompany this order. This amount includes postage and mailing costs. Texas residents
add 5% sales tax.

Volume XXIX, Nos. 1 and 2

September, 1977

CONTENTS

Instructions to Authors . . . . 3

Effect of Balcones Faults on Groundwater Movement, South Central Texas. By Patrick L.

Abbott . . . . . . . . 5

Baking of Shale by a Basaltic Dike: A Chemical and Strontium Isotopic Study. By

E. Julius Dasch, Hannes K. Brueckner, and J. M. Rhodes ................ 15

Influence of Anisotropies on the Shear Strength and Field Behavior of Heavily Over¬
consolidated, Plastic and Expansive Clay-Shales. By Robert G. Font . 21

Sex Ratios and Spawning of White Bass ,Morone chrysops, from the Red and Washita

River Segments of Lake Texoma. By Raymond E. Baglin, Jr. . 29

Density and Distribution of the Black-Tailed Prairie Dog in Texas. By Lloyd K.

Cheatheam . . . . . . . 33

Yield Estimates Derived from Active and Passive Creel Surveys of a Small Pond Fishery.

By Fredrick V. Jones, William D. Pearson, and Lloyd C. Fitzpatrick ......... 41

Age and Growth of Largemouth Bass in Canyon Reservoir, Texas. By Joe W. Kolb

and B. G. Whiteside . . . . . 49

The Effects of Amniotic Fluid on the Growth of Certain Gram-Negative Bacteria. By

John R. McGill and Caroline P. Benjamin . 59

Dentitional Phenomena and Tooth Replacement in the Scabbard Fish Trichiurus

lepturus Linnaeus (Pices : Trichiuridae). By Edward C. Morgan . . . 71

Evidence for the Species Status of Baird’s Ratsnake. By R. Earl Olson . 79

Antigenic Analysis of the Genus Aeromonas. By VijayaB. Rao and B. G. Foster .... 85

Growth Responses of Tilapia aurea to Feed Supplemented with Dried Poultry Waste.

By Robert R. Stickney, Herbert B. Simmons, and Lenton O. Rowland . . 93

Dog Flesh as a Potential Food Resource for Carnivores: An Exploratory Study. By

Dr. James U. McNeal and Mr. William L. Griffin ..................... 101

Estimations of the Ratio of Induced Fission to Spontaneous Fission in Uranium Ores.

By Moses Attrep, Jr., K. S. Tasa, and J. D. Sherwood . . . . 109

Mass Spectrometry Studies of Norcamphor and Norbornyl Acetate. By James L.

Marshall and Steven R. Walter . . . 121

Chromatographic Studies of Interaction Coefficients of Benzophenone and Benzhydrol

with Various Compounds. By Reeves B. Perry and Terry K. Henson . 131

The Temperature of Maximum Density of Deuterium Oxide in Capillaries. By J. A.

Schufle and Hsieh-Sui Hao . . . . . . 137

NOTES SECTION

Equus tau Owen from the Pleistocene of Mitchell County, Texas. By Walter W.

Dalquest . . . . . 141

The Structure of the Retina in Four Species of Whistling Ducks, Dendrocygna. By

Stephen M. Womack, Michael K. Ry lander, and Eric G. Bolen . . 141

If

'

• - ■ •

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.

INSTRUCTIONS TO AUTHORS

Papers intended for publication in The Texas Journal of Science are to be sub¬
mitted to Dr. Roland Vela, Editor, P. 0. Box 13066, North Texas State University,
Denton, Texas 76203.

The manuscript submitted is not to have been published elsewhere. Triplicate
typewritten copies (the original and 2 reproduced copies) MUST be submitted.
Typing of both text and references should be DOUBLE-SPACED with 2-3 cm
margins on STANDARD 8V2 X 11 typing paper. The title of the article should be
followed by the name and business or institutional address of the author(s). BE
SURE TO INCLUDE ZIP CODE with the address. If the paper has been
presented at a meeting, a footnote giving the name of the society, date , and occasion
should be included but should not be numbered. Include a brief abstract at the
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introduction (understandable by any scientist) and then whatever paragraph
headings are desired. The usual editorial customs, as exemplified in the most
recent issues of the Journal, are to be followed as closely as possible.

In the text, cite all references by author and date in a chronological order, i.e.j
Jones (1971); Jones (1971 , 1972); (Jones, 1971); (Jones, 1971 , 1972); Jones and
Smith (1971); (Jones and Smith, 1971); (Jones, 1971; Smith, 1972; and Beacon,
1973). If there are more than 2 authors, use: Jones, etal. (1971); (Jones, et al,
1971). References are then to be assembled, arranged ALPHABETICALLY, and
placed at the end of the article under the heading LITERATURE CITED. For a
PERIODICAL ARTICLE use: Jones, A. P., and R. J. Wilson, 1971-Effects of
chlorinated hydrocarbons./. Comp. Phys. , 37:116. (Only the 1st page number
of the article is to be used.) For a PAPER PRESENTED at a symposium, etc., use
the form: Jones, A. P ., 1971— Effects of chlorinated hydrocarbons. WMO Sym¬
posium on Organic Chemistry, New York,N.Y. For a PRINTED PAPER use: Jones,
A.P., 1971— Effects of chlorinated hydrocarbons. Univ. of Tex., Dallas, or Jones,
A. P., 1971— Effects of chlorinated hydrocarbons. Univ. of Tex. Paper No. 14,46 pp.
A MASTERS OR Ph.D THESIS should appear as: Jones, A. P., 1971 -Effects of
chlorinated hydrocarbons. M.S. Thesis, Tex. A&M Univ., College Station. For a
BOOK, NO EDITORS, use: Jones, A. R, 1971— Effects of Chlorinated Hydrocarbons.
Academic Press, New York, N.Y., pp. 13-29. Fora CHAPTER IN A BOOK WITH
EDITORS: Jones, A. R, 1971— Structure of chlorinated hydrocarbons.//? A. R Jones
and T. S. Gibbs (Eds.), Effects of Chlorinated Hydrocarbons. Academic Press,
New York, N.Y.,pp. 3-12. For an IN PRESS PERIODICAL reference, use: Jones, A.P.,
1971— Effects of chlorinated hydrocarbons./, of Org. Chem ,, In Press. For an
IN PRESS BOOK reference, use: Jones, A.P., 1911— Effects of Chlorinated Hydro¬
carbons. Academic Press, New York, N.Y. In Press.

All tables are to be typed with a carbon ribbon, free of error, without hand¬
written notations, and be prepared for photographic reproduction. Tables should
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(see Table 2).

4

THE TEXAS JOURNAL OE SCIENCE

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All photographs, line drawings, and tables are to be provided with self-
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Galley proof of each article will be submitted to the author. This proof must
be carefully corrected and returned within 3 days to the Managing Editor’s Office
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IMMEDIATELY SO WE CAN SEND YOUR GALLEY PROOF TO YOU
WITHOUT LOSS OR DELAY.

EFFECT OF BALCONES FAULTS ON GROUNDWATER MOVE¬
MENT, SOUTH CENTRAL TEXAS

by PATRICK L. ABBOTT

Department of Geological Sciences,
San Diego State University,

San Diego, California 92182

ABSTRACT

The subparallel normal faults of the Balcones system have created an extensive fracture sys¬
tem in Lower Cretaceous limestones in south central Texas. The permeability avenues pro¬
vided by the fractures have allowed surface water to enter already porous limestone and create
extensive cavern systems that are especially pronounced in the Edwards Group. Some of the
elongate, fault-bounded blocks of carbonate rock contain porosity systems that are partly
independent of those in adjoining fault blocks. The channelization of groundwater flow within
some fault blocks is indicated by preferential concentration of sinkholes, cave plans, and
separate but parallel spring systems.

INTRODUCTION

Heterogeneous dissolution of Lower Cretaceous limestones (Aptian to Albian)
has created an extensive network of caverns in south central Texas. The Edwards
Group has been most affected as has, to a lesser extent, the subjacent Walnut and
Glen Rose Formations. These limestone bodies have minimal thicknesses that
total several hundred feet and they extend over several thousand square miles, yet
the significant development of caverns occurs mainly within the Balcones Fault
zone (Fig. 1).

The carbonate rocks were formed with high initial porosities. During an inter¬
val of subaerial exposure at the close of Edwards Group deposition, some caverns
were created. Nonetheless, the modern cavern system is primarily due to dissolution
associated with faults. The Early Miocene subparallel, down-to-the -coast, normal
faults of the Balcones system improved the access of surface water into the void
systems of the subsurface rocks, but more importantly, the fault -rejuvenated
erosion exposed discharge sites that allowed initiation of a continuously circulating
groundwater system (Abbott, 1975). Groundwater undersaturated with respect
to calcite and dolomite dissolves the carbonate rocks it flows through. Thus a
self-ramifying process is set in motion whereby groundwater continuously enlarges
its initial flow paths, thereby receiving ever-increasing amounts of groundwater.
The original fault-created porosity and permeability have been greatly increased
through time.

Received for publication: October 25, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

6

THE TEXAS JOURNAL OF SCIENCE

Figure 1 . Location map. Reading counterclockwise from the lower left, the 7.5 quadrangles
outlined in Comal County are: Bulverde, Bat Cave, New Braunfels West, Sattler
and Smithson Valley.

A broad-scale understanding of the genetic relationship between the Balcones
Faults and the extensive artesian aquifer has been well explained by Sayre and
Bennett (1942) and discussed effectively by Hill and Vaughan (1898), Livingston,
et al., (1936), George (1948, 1952) and other workers. The purpose of this study
was to examine the effects of individual faults on groundwater flow in order to
better understand the limestone aquifers in Comal and Bexar Counties.

LOCAL RELATIONSHIPS BETWEEN FAULTS, GROUNDWATER
FLOW AND CAVERN DEVELOPMENT

There are several questions that might be raised about the role of the faults in
affecting the flow of groundwater and thus the development of caverns. Have the

GROUNDWATER MOVEMENT

7

faults acted as conduits for lateral movement of groundwater along strike and, as
a consequence, been enlarged into caverns? Have the faults acted as barriers that
impeded the flow of groundwater by having juxtaposed impermeable rock masses
against updip aquifer intervals? Have the faults placed different aquifer horizons
together so that groundwater moves downdip across faults without noticeable
hindrance? Or have all of the above occurred in different local areas? Several
lines of evidence are developed that bear on these questions.

Evidence From Plots of Groundwater Levels

One way to evaluate the effect of faults on groundwater is to examine water
levels in wells to see if marked differences exist on respective sides of a fault that
would indicate a damming or compartmentalization . Conversely, the water-level
elevations may present an evenly -sloping surface that apparently is unaffected
by the presence of faults.

Several books of water-level elevations in wells have been published by the
U.S. Geological Survey and the Texas Water Development Board. The data con¬
sist primarily of multiple readings on wells in the San Antonio and confined aquifer
areas or of one-time water-level readings in the unconfined aquifer northwest of
the main fault-line scarp of the Balcones system. Although many wells are available
for mapping the groundwater flow pattern, the right kind of data is difficult to
obtain. There are insufficient continuous hydrograph records, and it is impossible
to utilize the typical water-level readings which were taken in widely varying
months and years and thus reflect monthly and yearly fluctuations in rainfall.
However, it is apparent from the water-level data in George (1952), Follett (1956),
and Pettit and George (1956) that a field program was undertaken in May of 1945
to record the levels of as many wells as possible in as short a time as possible.
Although the water levels in' general declined during May, 1945, these records
afford us as close an approximation to a map of the potentiometric surface as is
available (Fig. 2).

The water level elevations plotted in Figure 2 are too scattered to allow definitive
pronouncements about local groundwater movements. There is an overall slope
of the potentiometric surface toward the southeast, but this is also the direction
of the regional dip. An imperfect secondary trend of declining, water-level eleva¬
tions to the northeast within fault blocks is also discernible.

The wells north of the Waco Springs Fault draw water primarily from the
Glen Rose Formation and those to the south from the Edwards Group. The
potentiometric surface has a steeper slope north of the Bat Cave Fault which
suggests that the cavernous porosity networks are not as well developed in the
Glen Rose Formation. The few water levels available near the Bat Cave Fault do
not indicate a barrier effect. Yet the Natural Bridge Caverns are developed on
the upthrown block and abut directly against this fault (Jan Knox and Charles M.
Woodruff, Jr., personal comm.).

Analysis of the effect of the Waco Springs fault suffers from a similar paucity
of data points. The slope of the potentiometric surface in the block between the

8

THE TEXAS JOURNAL OF SCIENCE

Figure 2. Structural and hydrologic map for a portion of Comal County. Well locations
plotted as small circles; water levels recorded in May, 1945 .

Waco Springs and Bat Cave Faults is steeper than in the blocks south of the
Waco Springs Fault. An anomalously high value occurs at Krueger ranch (spot
elevation 759 on Figure 2 just south of the Waco Springs Fault about 1 1/2 mi
east of Dry Comal Creek) where large-scale cavern collapses have disrupted the
strata. Note that water emerges from West Waco Spring at 649 ft elevation which
is about 12 ft higher than the downdip water level. This suggests that the Waco
Springs Fault has acted as a dam that concentrated groundwater along the fault.

GROUNDWATER MOVEMENT

9

Comal Springs arise along a 500-yd length of the Comal Springs Fault. At the
springs, the throw along the fault has entirely displaced the Edwards Group and
caused it to abut a relatively impermeable seal of Washita Division and Gulf Series
strata. Southwest along the Comal Springs fault the throw decreases to about
200 ft at the southern boundary of the Bat Cave quadrangle (Newcomb, 1971).
Groundwater apparently crosses the Comal Springs Fault easily in the southwestern
part of the county where the Edwards is only partially offset.

Notice that the bad-water line pays no heed to the nearby presence of the
largest fault in the Balcones Fault zone (Fig. 2). The bad-water line is a distinct
boundary between the potable water of the Edwards artesian aquifer and the
nonpotable water immediately downdip. The irregularity of this demarcation
line and its disregard for faults suggest that it is the dissolution-ingrained flow
boundary of the early groundwater that flowed to the initial discharge sites (Abbott,
1975).

Evidence From Cave Maps

The relationship of cavern formation to faults can probably best be visualized
by inspection of caves. Two large caves (Bracken Bat Cave and Natural Bridge
Caverns) developed near the Bat Cave Fault occur in the report area (Fig. 2). Maps
of these caverns have been published by Red dell (1964) and adapted plans are
shown in Fig. 3.

Both caverns have developed in the Glen Rose Formation on the up thrown
block next to the Bat Cave Fault. The long segments of the caves are preferentially
developed in near-horizontal fashion along bedding. In one stretch of Natural
Bridge Caverns there are 2 subhorizontal passages developed at different levels
within the 120-ft thick, upper dolomite subdivision of the Glen Rose Formation
(Abbott, 1973). The different levels of cave formation in the Glen Rose Formation
occur within a thick unit of relatively homogeneous lithology above the “alter¬
nating beds” interval. These interconnected, subhorizontal cavern passages probably
make a good analogue to conditions within the generally similar rock types that
comprise the Edwards Group to the southeast.

The trend of both cavern systems is largely perpendicular to and terminated
near the Bat Cave Fault. The trend of the fault is roughly N60E and Bracken Bat
Cave has formed along a N30W course. The right-angle orientation of the cave
relative to the trend of the fault may have been inherited from a joint system
formed at the time of the faulting.

The longest passages of Natural Bridge Caverns run approximately N30W,
parallel to the trend of Bat Cave, and at right angles to the fault. Other large cave
segments have developed along subparallel trends of about N30E. A lesser cavern
branch has developed near to and parallel with the Bat Cave fault on aN60E course.
The repeated subparallel cave passage trends suggest dissolution along fault -associated
joint systems. Both caverns are developed on the updip side of the fault and seem
to indicate that groundwater did not easily flow across that segment of the fault.

10

THE TEXAS JOURNAL OF SCIENCE

Figure 3. Maps of major passages of Bracken Bat Cave and Natural Bridge Caverns adapted
from Reddell (1964). Cavern locations are shown in Fig. 2.

Evidence From Preferential Development of Sinkholes

The fault block between the Bat Cave and Waco Springs faults contains a pro¬
fusion of sinkholes and some mega-collapse areas where beds have been chaotically
deformed by the collapse of large caverns in regions of about 2 mi2 each. The
recognition of abundant karstic features in this fault block is partly a function of
the surface stratigraphy but is probably also due to some groundwater channeli¬
zation within the fault block.

GROUNDWATER MOVEMENT

11

The outcrops on this fault block commonly include a 22 -ft thickness of clayey
biomicrites of the Regional Dense Member which occurs about midway through
the Edwards Limestone. The Edwards Group has been divided by Rose (1972)
into the Kainer (lower) and Person Formations by separation at the base of the
Regional Dense Member which is the basal unit of the Person Formation. Although
the Regional Dense Member does not create an impassable seal, it seems to create
locally perched groundwater bodies which flow laterally through the overlying
strata. This process explains the profuse sinkholes, but the mega-collapse areas
seem too large to be explained solely by this mechanism.

A mega-collapse area has been mapped in the northern part of the fault block
on the former John Classen ranch that lies astride the Bulverde and Bat Cave
quadrangles (Abbott, 1973). Here erratically dipping, petrified wood-bearing
beds of the Person Formation (upper Edwards) have collapsed down through the
Regional Dense Member into the Kainer Formation (lower Edwards). Prominent
caverns still exist in this area.

An even more striking mega-collapse area occurs in a relatively narrow part of
the Bat Cave -Waco Springs fault block on the Dietz Ranch in the northwestern
part of the New Braunfels West quadrangle (Abbott, 1973). The ground surface
is largely a mesquite -covered, cherty, terra rosa regolith which tends to obscure
the detailed relationships of the collapses. However, it is clear that beds of the
Georgetown and Del Rio Formations that overlie the Edwards Group are found
at the general stratigraphic level of the Regional Dense Member and are completely
surrounded by Edwards strata. This stratigraphic association, accompanied by
some steep dips, suggests a downward collapse of about 200 ft. The abundant
collapses in this 2 1/2 by 1 1/4 mi are a may have followed the increased dissolution
accomplished by groundwater forced to move through a narrower part of the
fault block.

No evidence of similar mega-collapse features on adjacent and nearby fault
blocks has been demonstrated to date. Thus, their presence in the Bat Cave-Waco
Springs Fault block may be attributed partly to the long-term flow of groundwater
confined largely to this fault block.

Evidence From Spring Flow

Waco and Comal Springs are major outlets that drain a large part of Comal
and Comal/Bexar Counties, respectively (Fig. 2). They appear to be independent
and unconnected systems although they are separated by only 3 mi in a north-
south direction. These separate but parallel delivery systems strongly suggest
that faults have aided the channelization of groundwater flow.

Waco Springs. The Waco Springs rise along the Waco Springs Fault at 2 sites
in the floodplain just west of the Guadalupe River in the southeastern part of
the Sattler quadrangle. Water emerges from outlets about 5 and 10 ft above the
level of the Guadalupe River, at flows ranging from 0 to 96 ft3 /sec, with temper¬
atures fluctuating between 68 and 71 F, and with turbid flow after heavy rains.

12

THE TEXAS JOURNAL OF SCIENCE

The Waco Springs Fault, and the dipping beds near it, appear to have had an
impeding or channelizing effect on groundwater flow that has been responsible,
in part, for the location of Waco Springs. Near the Guadalupe River the Waco
Springs Fault splits into a sliver (Bills, 1957) from which the Waco Springs emerge
out of arched beds that dip back to the northwest. This roll-over, plus the possible
juxtaposition of less permeable beds on the downthrown side against more per¬
meable beds on the upthrown side, has caused groundwater to be diverted along
the fault. This diversion has been reinforced by development of a cavern system
parallel to the fault which has grown in dominance through time.

The Waco Springs recharge and delivery system was partly explained by Bills
(1957) who followed up the disposition of a localized August rainstorm during
the drought of 1956 when the springs were not flowing. An isolated thunder
storm dumped up to 5 in of rain in the vicinity of Smithson Valley (Fig. 2). The
surface runoff went bank full down the west fork of Dry Comal Creek until
reaching the Bear Creek Fault where it went underground. Approximately 24 hrs
after this rain, the West Waco Spring began discharging about 2 ft3/sec of slightly
muddy water which subsided and ceased 2 days later.

The floodwater that went underground on the west fork of Dry Comal Creek
at the crossing with the Bear Creek Fault apparently descended through the alluvial
cover and the Walnut Formation into the upper dolomite subdivision of the Glen
Rose Formation. The groundwater possibly moved toward the Bat Cave fault
through a cavern system similar to the nearby Bat Cave and Natural Bridge Caverns.
The groundwater apparently crossed the Bat Cave fault which juxtaposes the
cavernous upper dolomite of the Glen Rose against the Kainer Formation (lower
Edwards). Then the groundwater may have moved more or less diagonally across
the narrow fault block between the Bat Cave and Waco Springs Faults, passed
beneath the Dietz Ranch mega-collapse area, and emerged through contorted
beds within a fault sliver along the Waco Springs Fault to flow out of West Waco
Spring. Of course the floodwater that flowed down Dry Comal Creek may not
be the same water which came out of West Waco Spring. Local flood recharge
increased the head within the aquifer and the transmitted effect of the increased
head may have caused the spring to flow.

The Waco Springs water-supply system apparently runs in large part to the
east and northeast, or approximately at right angles to the surface drainage. The
direction of flow seems to be subparallel to the faults and illustrates the effect
of the faults on the groundwater flow.

Comal Springs . The flow of Comal Springs has varied between 0 to 420 ft3/sec.
It is one of the largest springs in the U.S. but the current and future discharge is
lessened by the voluminous extractions of groundwater from wells in the San Antonio
area. The average discharge is greater than the runoff from the 1,432 mi2 drainage
area of the Guadalupe River above the springs. The groundwater emerges in New
Braunfels along a 500-yd length of the main fault-line scarp of the Balcones system
(Comal Springs Fault) about a mile from the Guadalupe River, which is 40 ft

GROUNDWATER MOVEMENT

13

lower. The water is never turbid and its temperature remains steady at 74 F which
is 6° higher than the average annual temperature at New Braunfels. This increased
temperature has been accounted for by circulation paths between 300 to 500 ft
below the surface (George, 1948, 1952).

The immense discharge of clear, constant temperature water indicates that
Comal Springs is the outlet for groundwater collected over a vast area. To the
north lies the effluent Guadalupe River and the isolated Waco Springs system
which draws from the northwest. To the east and south is the highly mineralized
water across the bad -water line. Thus the groundwater must be derived from the
southwest. This means the subsurface water flow is subparallel to the Balcones
Faults, about at right angles to the surface drainage, and at oblique angles to the
regional dip and to the apparent regional piezometric surface contours. A master
conduit system, or a greater volume of honeycombed and cavernous beds, has
formed subparallel to the fault beneath a cover of younger rocks in analogous
and parallel fashion to the Waco Springs system.

CONCLUSIONS

Mapped caves trend at right angles to faults, mega-collapse regions are developed
in certain fault blocks rather than along faults, and spring-supply systems com¬
monly cross faults. These are evidence that fault surfaces are not the major sites
of cavern development and that the various fault-defined blocks have had differ¬
ent degrees of cavern formation. The larger faults in Comal County (he., Comal
Springs and Waco Springs Faults, and to a lesser degree the Bat Cave Fault) have
acted in part as barriers that have diverted groundwater flow to the northeast.
The location of most major cavern systems was determined largely by the porosity
and permeability, plus structural controls incident to Balcones faulting, created
by the early groundwater flow patterns established millions of years ago (Abbott,
1975). Thus the modern cavern locations are largely inherited and may display
no easily discernible relationship to modern topography and stream courses.

ACKNOWLEDGEMENTS

I would like to thank reviewer Charles M. Woodruff, Jr., for his helpful comments
and Enos J. Strawn for drafting the figures.

LITERATURE CITED

Abbott, P. L., 1973-The Edwards Limestone in the Balcones fault zone, south central Texas.
Unpub. Ph.D. diss., Univ. Tex. at Austin, 122 p.

i— t-— , 1975 -On the hydrology of the Edwards Limestone, south central Texas. J. Hydrol¬
ogy , 24:251.

Bills, T. V., Jr., 1957-Geology of Waco Springs quadrangle, Comal County, Texas. Unpub.
MA thesis, Univ. Tex. at Austin, 106 p.

14

THE TEXAS JOURNAL OF SCIENCE

Follett, C. R., 1956-Records of water-level measurements in Comal and Guadalupe Counties,
Texas. Tex. Board Water Engrs. Bull. 5610, 32 p.

George, W. O., 1948-Development of limestone reservoirs in Comal County, Texas. Amer.
Geophys. Union Trans., 29:503.

- — , 1952-Geology and groundwater resources of Comal County, Texas. U.S. Geol.

Survey Water Supply Paper 1138, 126 p.

Hill, R. T., and T. W. Vaughan, 1898-Geology of the Edwards Plateau and Rio Grande plain
adjacent to Austin and San Antonio, Texas, with reference to the occurrence of under¬
ground waters. U.S. Geol. Survey 18th Ann. Rept., 2: 193.

Livingston, P., et. al., 1936-Water resources of the Edwards Limestone in the San Antonio
area, Texas. U.S. Geol. Survey Water Supply Paper, 773-B:5 9.

Newcomb, J. H., 1971-Geology of the Bat Cave quadrangle, Comal and Bexar Counties,
Texas. Unpub. MA thesis, Univ. Tex. at Austin, 104 p.

Pettit, B. M., Jr., and W. O. George, 1956-Groundwater resources of the San Antonio area,
Texas. Tex. Board Water Engrs. Bull. 5608 (2 vols.), 842 p.

Reddell, J. R., 1964- A guide to the caves of Texas. In J. R. Reddell (Ed.), Guidebook , 1964.
Natl. Speleol. Soc. Convention, New Braunfels, 61 p.

Rose, P. R., 1972-Edwards Group, surface and subsurface, central Texas. Univ. Tex. at Austin,
Bur. Econ. Geol. Rept. Inv. 74, 198 p.

Sayre, A. N., and R. R. Bennett, 1942-Recharge, movement, and discharge in the Edwards
Limestone reservoir, Texas. Amer. Geophys. Union Trans., 23:19.

BAKING OF SHALE BY A BASALTIC DIKE: A CHEMICAL AND
STRONTIUM ISOTOPIC STUDY

by E. JULIUS DASCH

Department of Geology , Oregon State University
Corvallis 97331

and HANNES K. BRUECKNER1

Queens College of the City University of New York,
Flushing 11367

and J. M. RHODES

Geoscience Section, Lockheed Electronics Company,
Houston 77058

ABSTRACT

Baking of friable, calcareous shale by approximately 1 m-thick alkalic basalt dikes in
Trans-Pecos Texas resulted in prominent but relatively thin selvages of densely lithified shale
adjacent to these dikes. Analyses of a sequence of baked and unaltered shale samples from
one of the outcrops shows an infiltration of Fe, Mn, Mg, and possibly Sr into the host shale.
The effects of baking of the shale, reflected in the clay mineralogy, also are shown by varia¬
tions in the Sr isotopic compositions of whole-rocks, acid-leached residues, and acid supernates
from samples of shale successively closer to the dike contact. Thermal effects by the intrusion
apparently caused an approach to equilibrium of Sr isotopes between the marine carbonate
and aluminosilicate fractions of the shale, as a function of increasing heat from the dike.
However, complete homogenization was not achieved in a shale sample less than 1 cm from
the dike contact. Thus, internal homogenization of Sr isotopes in detrital sediments is not
achieved even at the relatively high temperatures (A 1000C) found near hypabyssal intrusions.

INTRODUCTION

Although baking (mild, contact metamorphic effects resulting from hypabyssal
intrusions and lava flows) is a common geologic feature, there are few chemical
studies of the interaction between the magmatic and baked phases. In this study
we report and discuss certain chemical and mineralogic features of calcareous
shale baked by a basaltic dike.

1 Present address: Lamont-Doherty Geological Observatory of Columbia University, Palisades,
New York 10964.

Accepted for publication: October 2, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

16

THE TEXAS JOURNAL OF SCIENCE

Baked tuff, conglomerate, and especially shale are prominent features in the
Rim Rock dike swarm of Trans-Pecos Texas (Dasch, 1959; Dasch, et al, 1969).
The alkalic basaltic dikes, which range in age from about 18 to 23 m.y. (Dasch,
et al, 1969), transect Upper Cretaceous (Gulfian) calcareous shale and sandstone,
and Tertiary volcanic strata (Vieja Group), mainly tuff, tuffaceous sedimentary
rock, ignimbrite, and silicic flow rock. The approximately 100 dikes that have
been mapped range in thickness from several cm to several m, but the majority
are close to about 1 m in thickness. Baked host rock selvages range in thickness
according to dike thickness; in shale exposures, the average, 1 m-thick dike has
baked zones about 35 cm thick. Contacts between baked and unbaked shale are
surprisingly sharp in most outcrops. In many exposures, baked rocks are more
resistant to weathering than the basalt and stand as prominent parallel walls.

The yellow-weathering, calcareous shale adjacent to the dikes is densely lithified
and somewhat darkened as a result of baking. Some thin sections show euhedral
crystals of feldspar, pyroxene, and amphibole which apparently formed by infil¬
tration of magmatic fluids into the shale. The dense, baked shale exhibits well
developed cooling joints in most outcrops.

The dike selected for study (Dasch, et al, 1969) has been analyzed in some
detail (Dasch, 1969; Dasch, et al, 1969). It is an alkalic basalt porphyry with
large, partly resorbed phenocrysts of anorthoclase, kaersutite, biotite, plagioclase,
and apatite. Gas vesicles in this amygdaloidal rock are filled with analcime and
calcite. The fine-grained matrix consists of plagioclase, pyroxene, and scattered
olivine crystals, with patches of poorly crystallized alkalic feldspar and analcime.
The secondary minerals apparently result more from late -stage magmatic, or deu-
teric, alteration rather than from weathering (Dasch, 1969).

METHODS

Samples were obtained from the dike (about 12 cm into the dike from the dike-
shale contact), from baked shale 2.5 cm (S-5) and 23 cm (S-8) from the contact,
and from apparently unmetamorphosed shale 183 cm (S-10) and 610 cm (S-l 1)
from the contact.

Carbon was determined at Oregon State University on water-rinsed samples
utilizing a Leco carbon analyzer. Rock powders for the other major elements
were ignited at 1000 C for several hours to liberate H2 O and C02 , and to oxidize
the Fe. Excepting Sample S-10, element concentrations then were determined
by X-ray fluorescence spectrometry at the Manned Spacecraft Center (NASA),
on a glass disc prepared by fusing a 280 mg aliquant of the sample with a lanthanum¬
bearing, lithium borate fusion mixture (Norrish and Hutton, 1969). Calibrations
were based on primary synthetic standards supplemented by previously analyzed
U.S.G.S. and N.B.S. rock and mineral standards. Na was analyzed by atomic
absorption spectrometry on a separate 20 mg aliquant of the sample. All samples
were analyzed in duplicate.

BAKING OF SHALE

17

Sr was separated from the samples by standard techniques of dissolution and
ion exchange. Sr isotope ratios were measured on rhenium triple filaments on
the 12 -in radius of curvature, 60° sector mass spectrometer at Lamont-Doherty
Geological Observatory. Precision on 'Sr87 /Sr86 ratios = ± 0.0002 except as noted
in Table 2. Sr87 /Sr86 values were normalized to a Sr88 /Sr86 value of 8.3752.

RESULTS AND DISCUSSION

As with other dikes in the Rim Rock dike swarm (Dasch, 1959) the analysis
of Dike 188 (Table 1) shows it to be a soda-rich, alkalic basalt; the large amount
of apparently primary analcime indicates a composition approaching ph on olite.
Calcite from one of the large, calcite -analcime arnygdales was analyzed by G. D. Garlick
(personal communication) for O and C isotopic composition. The results, 60 18 =
+17.6%o(SMOW), SC13 = -3.8°/oo(PDB), indicate that the calcite is clearly sec¬
ondary. Textural relations in matrix analcime, however, suggest that much of this
feldspathoid is deuteric in origin. A calcite amygdale from this dike has a Sr87 /Sr86
ratio of 0.7056, whereas the dike itself is much less radiogenic, 0.7041 (Dasch,
1969); Sr from the secondary calcite thus is at least partly derived from sources
other than the original magma- -such as sedimentary calcite from the adjacent
shale.

TABLE 1

Major element concentrations of Dike Rock 188, and unbaked (S-ll)
and baked (S-5 , S-8) samples of shale.

188

(interior
of dike)

S-5

(baked shale,

2.5 cm from
contact)

S-8

(baked shale,

23 cm from
contact)

S-ll

(unaltered shale,
610 cm from
contact)

Si02

47.11

48.26

55.93

48.10

Ti02

2.28

0.65

0.56

0.64

A1203

16.76

15.21

12.48

14.51

Fe203

10.28

6.54

4.28

4.82

MnO

0.13

0.09

0.06

0.02

MgO

4.88

2.91

1.84

1.93

CaO

8.35

17.34

17.03

18.43

Na20

4.79

4.97

4.29

3.17

k2o

2.76

0.86

0.64

2.09

P205

1.43

0.19

0.18

0.16

s

0.10

0.03

0.02

1.57

c*

0.91

3.37

3.63

3.20

Totals

99.78

100.42

100.94

98.64

*Total C measured prior to ignition.

18

THE TEXAS JOURNAL OF SCIENCE

Apparently unmetamorphosed shale (S-10, S-ll) and progressively more intensely
baked shale (S-8, S-5) have compositions as shown in Table 1. Initial chemical
variations within this detrital unit undoubtedly mask many minor changes brought
about by baking. However, the data suggest rather marked increases in Fe,Mn,
and Mg as a result of the contact effects. As euhedral, apparently igneous minerals
are apparent in thin sections of the more highly baked rocks, it is obvious that
some infiltration of magmatic fluid has occurred.

Mineralogic analysis of whole-rock samples of S-ll, S-8, and S-5 were made
by E. A. Perry (personal communication). Sample S-l 1 is mainly composed of a
mixed-layer illite/smectite with about 20% smectite and ordered (allevardite-type)
interlayering. It also contains about 15 to 20% chlorite and kaolinite and appears
to be completely unmetamorphosed. Samples S-8, from the baked zone, contains
no mixed-layer clay, illite, or potassium mica; it is comprised largely of chlorite,
quartz, low albite, and potassium feldspar. Sample S-5 is much like S-8, but the
chlorite is better crystallized, and, along with feldspar, is somewhat more abundant
than in S-8. The mineralogy of Samples S-8 and S-5 thus exhibit the effects of
progressive contact metamorphism.

Strontium isotope analysis (Table 2; Figure 1) of the samples indicates that
comparatively nonradiogenic Sr of the dike (Sr87/Sr86 = 0.7040) infiltrated the
more radiogenic shale, lowering the Sr87/Sr86 ratios in whole -rock samples of the
shale from about 0.709 to about 0.707 adjacent to the dike. Our interpretation
of these data is consistent with other observations: the dike is a ready source of
Sr (1425 ppm), and euhedral minerals occur in the baked rocks, which apparently
formed as a result of fluid infiltration.

TABLE 2

Strontium isotopic composition of dike, and of whole-rock, acid
supernate, and acid-leached residue from shale samples. Sr87/Sr86
ratios normalized to Sr88 /Sr86 = 8.3752. Analytical uncertain ties
(2 a) on Sr87/Sr86 = ±0.0002 except for acid supernate for S-l 1
(±0.0003) and leached residue for S-8 (±0.0004).

Distance From Dike-

Sample Number Shale Contact, cm Whole Rock

Sr87/Sr86 Acid-leached

Acid Supernate Residue

Dike Rock (188)
S-5
S-8
S-10
S-ll

12.0

2.5

23.0

183.0

610.0

0.7040

0.7075

0.7076

0.7084

0.7090

0.7072

0.7072

0.7074

0.7080

0.7083

0.7086

0.7104

0.7122

Of more significance, Sr between carbonate and aluminosilicate fractions in
the shale has become partially equilibrated as a function of increasing degree of
baking. In addition to whole -rock samples, Sr isotope analyses of the shale were

-188 S-5 S-8 S- 10

BAKING OF SHALE

19

Figure 1. Sr87/Sr86 ratios of Dike Rock (188), and unbaked (S-ll) and baked (S-5, S-8)
samples of shale vs. distance from dike-shale contact. Except for the acid-leached
residues of Samples S-10 and S-ll, the measured ratios are the same as the ratios
at the time of formation (initial) of the rocks. Initial ratios of the acid-leached
residues of S-10 and S-ll are about 0.001 less than the measured values.

made on supernate and acid-leached sample aliquants. In all samples (Figure 1),
Sr in the aluminosilicate residue is more radiogenic than Sr associated with the

Distance from Dike- Shale Contact (cm)

20

THE TEXAS JOURNAL OF SCIENCE

leached carbonate. Carbonate (acid supernate) Sr of the Upper Cretaceous, un¬
metamorphosed shale (S-l 1) has an Sr87 /Sr86 ratio of 0.7080, close to the value
reported for Sr dissolved in Upper Cretaceous seawater (Peterman, et al, 1970).
However, in samples closer to the dike-shale contact, the spread between Sr87 /Sr86
ratios of Sr in carbonate and Sr in aluminosilicate phases is reduced from about
4 parts in 700 to 1 part in 700, ostensibly by equilibration resulting from increasing
metamorphism.

These data have some relevancy to the problem of dating shales by Rb-Sr tech¬
niques. Many geochronologists (eg. W. Compston/^ Bofmger and Compston, 1968)
believe that isochronism in detrital sediments may be brought about by early Sr
homogenization, owing to relatively mild burial metamorphism or diagenesis (see
also Fbrry and Turekian , 1974). Our data show that homogenization was approached
but not achieved, in the specific case of this baking study, but only at temperatures
near 1000 C at the dike-shale contact. Of course, the baking process probably is
short-lived; much longer periods of, say, burial metamorphism, even at much
lower temperatures, probably is more effective as a homogenizing process.

ACKNOWLEDGEMENTS

We thank G. D. Garlick (Humbolt State College) and E. A. Perry (University
of Massachusetts) for the isotopic and clay mineral analyses discussed herein.

LITERATURE CITED

Bofinger, V. M., and W. Compston, 1968-A reassessment of the age of the Hamilton Group,
New York and Pennsylvania, and the role of inherited radiogenic Sr87 Geochim. et Cosmo -
chim. Acta. 31 :2353.

Dasch, E. J., 1959-Dike swarm of northern Rim Rock country, Trans-Pecos Texas. Univ. of
Texas at Austin, Masters Thesis, 62 p.

- , 1969-Strontium isotope disequilibrium in a porphyritic alkali basalt and its bearing

on magmatic processes./. Geophys. Res., 74:560.

- , R. L. Armstrong, and S. E. Clabaugh, 1969-Age of Rim Rock dike swarm, Trans-

Pecos Texas. Bull. Geol. Soc. Amer., 80:1819.

Norrish, K., and J.T. Hutton, 1969-Preparation of samples for analyses by X-ray fluorescence
spectrography. C.S.I.R.O. Div. Soils Rept., 3/64.

Perry, E. A., and Turekian, K. K., 1974-The effects of diagenesis on the redistribution of
strontium isotopes in shales. Geochim. et Cosmochim. Acta, 38:929.

Peterman, Z. E., C. E. Hedge, and H. A. Tourtelot, 1970-Isotopic composition of strontium
in sea water throughout Phanerozoic time. Geochim. et Cosmochim. Acta, 34:105.

INFLUENCE OF ANISOTROPIES ON THE SHEAR STRENGTH
AND FIELD BEHAVIOR OF HEAVILY OVERCONSOLIDATED,
PLASTIC AND EXPANSIVE CLAY -SHALES

by ROBERT G. FONT

Department of Geology,

Baylor University, Waco 76703

ABSTRACT

Heavily over consolidated, highly plastic and expansive clay- shales crop out in the north cen¬
tral Texas region. These shales have given rise to problems related to engineering construction.
Slope instability, high potential volume change, and low shear strength characterize these
materials.

The highly anisotropic character of the clay-shales has been found to have a pronounced
effect on their field behavior. Their fractured and fissured character provides them with pre¬
existing planes of weakness which enhance slope instability and related problems. Field in¬
vestigations and laboratory testing demonstrate that if these anisotropies are not taken into
account, it is possible to greatly overestimate the shear strength of the shales, a fact that can
lead to disastrous results.

INTRODUCTION

The shales of the Taylor, Eagle Ford, Woodbine, and Washita groups of the
Cretaceous System crop out in the vicinity of the Waco urban area in north cen¬
tral Texas. These shales have become well-known for giving rise to problems re¬
lated to engineering construction. Detailed field studies and a rigorous program of
laboratory testing have demonstrated that all of the above mentioned shales are
highly anisotropic. These anisotropies are found to have a pronounced effect on
their field behavior.

The purpose of this study is to document the effects of these anisotropies on
the shear strength and field behavior of the Waco shales. It is expected that this
knowledge may help prevent problems in the future urbanization of the region.

LOCATION AND PREVIOUS WORK

The study area lies in and around the Waco urban area within McLennan
County , Texas (Fig. 1).

In 1972, Wright and Duncan recognized the highly anisotropic character of
the Pepper Shale in Waco (Table 1). The authors stated in their paper concerning
the analysis of the Waco Dam slide, that the Pepper Formation of the Woodbine

Received for publication: December 28, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

22

THE TEXAS JOURNAL OF SCIENCE

i

i

Figure 1. Location Map.

INFLUENCE OF ANISOTROPIES

23

TABLE 1

Stratigraphic Relationship, Depositional History and Maximum Thickness
of the Cretaceous Strata in the Local Area

Formation Depositional
Series Group and Symbol History

Maximum
Local
Thickness
in feet

Maximum
Probable
Thickness
at one time
in feet

Gulf Taylor

Austin

Taylor

Marl

Austin

Chalk

Eagle South

Ford Bosque

Shale

Woodbine

Lake

Waco

Fm.

Klw

Pepper

Shale

Se

Comanche Washita

Buda

Limestone

K

bu

Del Rio
Clay

George¬
town Lime¬
stone

K

ge

Deposited dur¬
ing marine trans¬
gressions and
regressions.

Deposited dur¬
ing marine trans¬
gression with pos¬
sible tluctuations
of the strandline
Unconformably un¬
derlies the K, .

ta

Deposited in a
neritic marine
environment with
poor circulation.
Unconformably un¬
derlies the K •
au

Deposited in a la-
goonal environment.
Conformably under¬
lies the Ks^.

Deposited in brackish
environment. Uncon¬
formably underlies

theKiw

Nearshore marine
deposit. Almost
completely eroded
locally prior to depo¬
sition of K .

pe

Marine regression and
transgression. Re¬
stricted environment.
Conformably underlies

the Ku .
bu

Shallow marine
deposit.

250

250

160

80

70

2

85

210

1170

295

160

145

100

35

85

210

24

THE TEXAS JOURNAL OF SCIENCE

Group is horizontally fissured. They also demonstrated that on the average, the
strength along these fissured planes is 40% lower than the strength obtained by
testing samples in a conventional style, so that the shear plane does not coincide
with a preexisting plane of weakness. Wright and Duncan (1972), in fact, related
the failure of the Waco Dam to the highly anisotropic character of the Pepper
Shale.

Studies by Font from 1969 to 1976 have led to the conclusion that all of the
Waco shales are highly anisotropic. Data supporting this conclusion are presented
in this paper.

Previous works regarding the engineering geology of the Waco urban region
include those of Burket (1965), Font (1969), Font and Williamson (1970), and
Font (1973a, 1973b, 1974, 1975, 1976a, 1976b).

GEOLOGIC SETTING

Most of the bedrock exposed within Waco include marine limestones and shales
of Cretaceous age. The Waco region is transgressed by the Balcones Fault Zone, a
zone of normal faults that cuts across Texas (Fig. 1). The Balcones Faults in Waco
trend NNE and are commonly downthrown to the east. Maximum displacement
along these faults is approximately 300 ft.

The stratigraphic relationship, depositional history and maximum thickness
of the Cretaceous strata are summarized in Table 1 .

Stress History >

The Waco shales are all heavily preloaded. Studies indicate that 500 to 2000 ft
of overburden overlain the top of these shales at one time (Font, 1976a). Since
overburden on top of these shales is now either totally absent or at the most only
a few feet thick, the Waco shales are heavily overconsolidated. The field instability
and stress history of the Waco shales is reflected by their engineering properties
as indicated in Table 2.

EFFECT OF ANISOTROPIES ON THE SHEAR STRENGTH AND

FIELD BEHAVIOR OF THE WACO SHALES

In order to study the effect of anisotropies on the shear strength and field
behavior of the Waco shales, a series of consolidated-undrained direct shear
tests simulating field conditions have been conducted on remolded and undis¬
turbed samples of these materials. Most of the tests have been performed in a
model D-120b direct shear apparatus manufactured by Soil Test Inc. Sensitivity
and unconfined compressive strength values have been obtained by running un-
confined compression tests on undisturbed and remolded samples at the same
water content. The apparatus used in these tests is a model U-560 manufactured
also by Soil Test Inc. The results obtained for all of the Waco shales are similar.
In view of this, results obtained for the Del Rio Formation of the Washita Group

INFLUENCE OF ANISOTROPIES

25

TABLE 2

Index Properties of the Waco Shales.

Liquid

Plasticity

Potential

Unified Soil

Limit

Index

Volume Change

System Category

60

30

Marginal

to

to

to

CH

80

52

Very Critical

Percent

Percent

Dry Unit

Liquidity

Clay

Calcium

Weight

Index

Carbonate

PCF

75

2

Zero

to

to

97

to

95

30

Negative

Sensitivity

Residual Strength Parameters

1.0

0r = i

5° - 10°

to

1.2

cr = 0

are the only ones presented here. The results obtained for the shales of the Taylor,
Eagle Ford, and Woodbine groups have been found to be nearly identical, but
even more pronounced than those obtained for the Del Rio Formation.

Figure 2 depicts 3 Mohr rupture envelopes obtained in testing the Del Rio
Formation. The upper rupture envelope has been obtained by testing undisturbed
samples, where the shear plane does not coincide with a preexisting plane of
weakness or anisotropic plane. The middle envelope depicts the results obtained
by testing normally consolidated remolded samples. The lower envelope has been
obtained by testing undisturbed samples of the Del Rio Formation, where the
samples have been oriented, so as to insure that the shear plane developed in the
test coincides with preexisting planes of weakness or planes of anisotropy. It can
be seen that the strength along preexisting planes of weakness is drastically lower
than that which could be expected by using the shear strength parameters of the
upper envelope of Fig. 2. On the average, it has been found that the strength
along preexisting planes of weakness is from 2.5 to 4 times lower than that expected
from the upper Mohr envelope. The important point to establish now is which
strength actually operates in the field. In order to accomplish this goal, several
slope failures within Waco have been analyzed. Slope stability analyses have been
performed on these slopes using the shear strength parameters shown in the upper
and lower Mohr envelopes of Fig. 2. It should be recalled that if a slope is stable,

26

THE TEXAS JOURNAL OF SCIENCE

Figure 2. Mohr rupture envelopes for the Del Rio Formation.

the slope stability analysis must yield a factor of safety greater than 1.0. It should
also be indicated that all of the slopes that have been analyzed by the author in
this study had already failed. In using the shear strength parameters which define

INFLUENCE OF ANISOTROPIES

27

the upper rupture envelope in Fig. 2, factors of safety considerably greater than
1.0 are obtained. This would indicate that the slopes are stable. However, since
the slopes have already failed, this could not be true. The shear strength parameters
that actually predict the instability of these slopes have been found to be those
of the lower envelope of Fig. 2.

CONCLUSIONS

It is found in analyzing several slope failures in the Waco shales, that the shear
strength at failure for these materials is very close to that which coincides with
the strength along preexisting planes of weakness. Thus, it is highly possible to
greatly overestimate the field strength of these shales if these anisotropies are ignored.
It is imperative to take into account the influence of these anisotropies in future
engineering endeavors if we wish to prevent catastrophic failures, such as that of
the Waco Dam.

ACKNOWLEDGEMENTS

I would like to express my gratitude to Wayne Mudd and Patrick Millegan for their
help in some of the laboratory testing. Special thanks are offered to Dr. G. A. Morales
of Baylor University for reviewing this manuscript. I am thankful to Mrs. Viola Shivers
for typing the manuscript.

LITERATURE CITED

Burket, 1965 -Geology of Waco in urban geology of greater Waco-Part I. Baylor Univ.

Geol. Stud. Bull. 8.

Font, R. G., 1969-Engineering geology of the greater Waco area. Unpub. thesis, Baylor Univ.

- , 197 3a-Engineering geology study of the instability of the South Bosque Shale

and the Del Rio Clay in the Waco area. Unpub. dissertation, Texas A&M Univ.

- , 1973b-Engineering geology study of the instability of two north central Texas

shales and the development of a reconnaissance tool in engineering geology. G.S.A. abstract,
Program with abstracts, National G.S.A., Dallas.

- , 1974-Influence of fundamental geologic factors on landslide genesis and mor¬
phology in overconsolidated Cretaceous shales— a study in urban engineering geology.
A.E.G. abstract, Program with abstracts, National A.E.G., Denver.

- — , 1976a-Relationship between the geologic history and engineering properties of

two Cretaceious shales. Tex. J. of ScL, 27(2) : 267 .

- , 1976b-Influence of anisotropies on the shear strength and field behavior of

certain heavily overconsolidated, highly plastic and expansive clay-shales. G.S.A. abstract,
Program with abstracts, National G.S.A., Denver.

- , and P. S. Millegan, 1975 -Effect of overconsolidation on the undrained strength

of an unstable Cretaceous shale -a geomechanical application to urban geology. G.S.A.
abstract, Program with abstracts, Regional, South Central G.S.A., Austin.

28

THE TEXAS JOURNAL OF SCIENCE

- , and E. F. Williamson, 1970 -Geologic factors affecting construction in Waco-Part IV.

Baylor Univ. Geol. Stud. Bull. 12.

Wright, S. G., and J. M. Duncan, 1972-Analysis of the Waco Dam slide./, of Soil Me ch. and
Found. Div., SM9:869.

SEX RATIOS AND SPAWNING OF WHITE BASS, MORON E CHRYSOPS,
FROM THE RED AND WASHITA RIVER SEGMENTS OF LAKE
TEXOMA1

by RAYMOND E. BAGLIN, JR.

National Oceanic and Atmospheric Administration,

National Marine Fisheries Service,

Southeast Fisheries Center-Miami Laboratory,

75 Virginia Beach Drive, Miami, Florida 33149

ABSTRACT

Sex ratios of white bass, Morone chrysops, from the Red and Washita River segments of
Lake Texoma were studied with reference to seasonal variations. The gonadosomatic index
was used to determine the duration of the spawning season.

INTRODUCTION

Fish production in many reservoirs declines markedly after the First few years
(Ellis, 1937). The white bass is a valuable species because it often maintains a
sport fishery in older reservoirs as the initial high production of game fishes declines.

In Lake Texoma, a 36,423 ha impoundment of the Red and Washita Rivers
in southern Oklahoma and northern Texas, the white bass is probably the most
sought-after species. To formulate a rational management program, knowledge
of its life history is necessary. This paper provides additional information on the
sex ratios and spawning of this species in Lake Texoma.

MATERIALS AND METHODS

From September 1969 to July 1972, 1,540 white bass were collected from
the Red and Washita Rivers which form Lake Texoma, primarily using gill nets
with 1.5 -in to 2-in square mesh. An electroshocker mounted on al4-ft flat-bottom
boat with a 220V generator also was used. Seining and angling were other methods
of capture.

Immature fish were sexed by removing a thin cross-section from one gonad,
placing it on a glass slide with a drop or two of aceto-orcein, and examining with

1 This is part of a dissertation presented in partial fulfillment of the requirements for the de¬
gree of Doctor of Philosophy at the University of Oklahoma.

Received for publication: April 28, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

30

THE TEXAS JOURNAL OF SCIENCE

a microscope. This technique allowed the detection of immature ova in the fe¬
males. Mature fish could readily be identified by external examination. The dur¬
ation of the spawning period was determined by studying the gonadosomatic index,
defined as the ovary weight divided by the total body weight times 100.

RESULTS AND DISCUSSION

Most Red River samples were taken approximately 24 km (15 mi) upstream
from the lake, and most white bass were taken from the Washita River about 16 km
or 10 mi from Texoma. On April 14, 1970, however, a collection of 8 males and
2 females was taken by electroshocking in Wild Horse Creek, a tributary of the
Washita River approximately 161 km (100 mi) upstream from Texoma. Ranchers
say they catch white bass there only in the spring.

Schooling by sex occurred during the white bass’s migration. For example, on
April 4, 1970, a mass migration passed through Cumberland Cut where the Washita
River enters Lake Texoma. Of 86 fish collected, all were males. Unisexual schooling
has also been reported for the white bass by Riggs (1955) in Lake Shafer and
Lake Texoma.

The Red River samples show that for 6 mo out of 1 1, males outnumbered
females; in the Washita River, males were more numerous than females 2 mo out
of 7 (Table 1). However, significant deviation from 1:1 sex ratios occurred in
Red River samples only for the month of March and significant deviation occurred
in the Washita River only in February.

TABLE 1

Monthly Sex Ratios for White Bass from the Red and Washita River
Segments of Lake Texoma for 1969-72.

Location

Month

Number of
White Bass

Sex Ratio
(Females/Males)

df1

2 2

Red River

J anuary

8

0.33

1

2.00

February

182

0.80

1

2.20

March

137

0.36

1

30.843

April

51

1.43

1

1.58

May

25

1.27

1

0.36

June

18

0.80

1

0.22

July

99

1.20

1

0.82

August

6

0.50

1

0.66

September

62

1.21

1

0.58

October

24

1.40

1

0.66

November

30

0.76

1

0.54

Washita River

J anuary

65

1.03

1

0.02

February

184

2.07

1

22.263

March

87

0.89

1

0.28

April

470

0.84

1

3.76

May

23

1.30

1

0.40

June

13

5.50

1

6.24

November

56

2.73

1

2.70

1 Degrees of freedom
1 Chi-square values
Significant at 0.01 level

SEX RATIOS AND SPAWNING OF WHITE BASS

31

Casselman (1975) found that sex ratios of northern pike , Esox lucius, from 3
Ontario populations showed seasonal trends in his samples. He believed that these
peaks of availability were related to activity and were independent of locality and
method of capture. The seasonal differences observed in white bass in this study
may be related to sexual differences in prespawning activity.

The gonadosomatic index (Fig. 1) showed that the ovaries of white bass from
Lake Texoma were most developed in March and April, and that by May they
were mostly spent. The spawning season, therefore, probably started in late March
and was completed by early May. Taber (1969), studying larval fish in the Buncombe
Creek arm of Lake Texoma,gave the first of April to early May as the white bass’s
Spawning season.

is — *

14-

13-

Figure 1 . Seasonal variation in mean gonadosomatic index (ovary weight/body weight x 100)
of Lake Texoma white bass for 1969-72. Numbers of fish are indicated.

ACKNOWLEDGEMENTS

This research was supported in part by a rese arch fellowship from the Oklahoma
Game and Fish Council and was subsidized by the U.S. Army Corps of Engineers.

I thank Grant Beardsley, Luis Rivas, Thomas Costello, and Lynn Pulos of the
National Marine Fisheries Service, Miami, Florida, for their helpful comments.

32

THE TEXAS JOURNAL OF SCIENCE

LITERATURE CITED

Casselman, J. M., 1975 -Sex ratios of northern pike,/iS0x lucius Linnaeus. Trans. Amer. Fish.
Soc 104(1) :60.

Ellis, M. M., 1937 -Some fishery problems in impounded waters. Trans. Amer. Fish. Soc.,
66:63.

Riggs, C. D., 195 5 -Reproduction of white bass, Morone chrysops. Invest. Ind. Lakes and
Streams, 4(3): 87.

Taber, C. A., 1969-The distribution and identification of larval fishes in the Buncombe Creek
arm of Lake Texoma with observation on spawning habits and relative abundance. Doctoral
Dissertation, Univ. of Oklahoma, 120 pp.

DENSITY AND DISTRIBUTION OF THE BLACK-TAILED PRAIRIE
DOG IN TEXAS1

by LLOYD K. CHE ATHE AM

Texas A&M University Agricultural Extension Service,
Rodent and Predatory Animal Control Service,

35 05 Haw thorne Drive, A marillo 79109

ABSTRACT

More than 30,000 Agricultural Stabilization and Conservation Service aerial photographs
providing coverage in 108 counties in central and western Texas were examined to provide
management data on the current population size and distribution of the black -tailed prairie
dog ( Cynomys ludovicianus) . In areas of incomplete photographical coverage, individuals
knowledgeable of black -tailed prairie dog colony locations were interviewed. A total of 1,336
prairie dog colonies inhabiting 36,432 ha were located in 89 counties in the study area. On¬
site inspections were made at 319 colonies (23.8% of total) to verify the colony’s presence,
size, and location. The average colony size was 27.27 ha. Croplands presently utilize 29.4%
of the study area’s prairie dog habitat. A total of 1,268 burrowing owls ( Speotyto cunicularia )
were observed in 91 of the 319 colonies inspected.

INTRODUCTION

In literature describing the black-tailed prairie dog, the species’ historic pop¬
ulation, circa 1900, is frequently compared with the present population. The
historic population in Texas is impressive and few authors dispute a considerable
decline; but the degree of the decline is questionable and the present population
has been difficult to ascertain. Control efforts are emphasized by many authors
as the major factor involved in the population’s decline while land use changes,
if acknowledged, are not well described.

The historic prairie dog population in Texas was estimated by Bailey (1905)
at 800,000,000 animals inhabiting 233,100 km2. “Their range extends from
Henrietta, Fort Belknap, Baird, and Mason west almost to the Rio Grande, north
over the Staked Plains and the Panhandle region, and south to the draws of Devil’s
River, to 10 mi south of Marathon and 25 mi south of Marfa” (Fig. 2).

Cottam and Caroline (1965) provided some conservative estimates on the present
population. They polled reliable individuals residing in counties containing prairie

XThis management survey was suported and administered by the U.S. Fish and Wildlife Ser¬
vice, Division of Wildlife Services, cooperating with Texas A&M University Agricultural
Extension Service.

Received for publication: April 1, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

34

THE TEXAS JOURNAL OF SCIENCE

dogs and concluded that Texas contained a minimum of 300 colonies. Other
authors have described a more tenuous abundance and some factors responsible
for the population’s decline.

Rue (1967) said, “Today, poison, traps, and guns have all but wiped out what
was once one of our most common western animals.” Drimmer (1954) reported,
“A few towns still persist in out-of-the-way places, but their numbers are limited.”
Davis (1960) reported on the prairie dog in Texas as, “Now rare throughout most
of its former range.” These undocumented descriptions do not adequately reflect
the prairie dog population size in Texas.

The need for documented management data on this species and the capability
to census colonies over vast areas provided the impetus for this study.

SURVEY AREA

The survey area encompasses 108 counties in central and western Texas total¬
ing 34,388,179 ha (343,615 km2) (Soil Conservation Service, 1970) and covers
the majority of the Kansan, Balconian, and Chihuahuan biotic provinces (Blair,
1950) (Fig. 1). Most of these counties are within the historical range of the prairie
dog in Texas as determined by Bailey (1905) (Fig. 2).

Mesquite (Prosopis juli flora), juniper (J unipenis spp.), sand shinnery oak ( Quercus
havardii), sand sage {Artemisia filifolia), and scattered riparian vegetation occur
throughout the area. Yucca ( Yucca galuca), prickly pear {Opuntia tuna ), and

BLACK-TAILED PRAIRIE DOG

35

Figure 2. Prairie dog range in Texas and one continuous prairie dog colony described by
Bailey (1905).

cholla ( Opuntia imbricata) are common. The major grasses are 5 species of bluestem
( Andropogon spp.), buffalo grass ( Buchloe dactyloides), and 6 species of grama
{Bouteloua spp.).

METHODS

Agricultural Stabilization and Conservation Service (ASCS) aerial photography
at a scale of 1:7,920 was used to locate colonies. Each photograph covers 10.36 km2
(4 mi2) and its accuracy is corrected to ±1%. Finding that black-tailed prairie
dog colonies could be recognized on these photographs led to a survey technique
for this species (Cheatheam, 1972).

Prairie dog colonies on the photographs appear as a grouping of small white
dots; each dot represents a large mound or a large area denuded of vegetation
around a prairie dog mound. Occasionally, short vegetation within the colony
perimeter is reflected on the photographs as a white haze through the colony.
Each colony’s size was measured with a polar planimeter and the colony was then
plotted on Texas Highway Department county maps with a notation of the colony’s
size.

An estimated 30,000 plus ASCS aerial photographs were examined in the
study area for prairie dog colonies. Photographs were available for most of the

36

THE TEXAS JOURNAL OF SCIENCE

study area; however, complete county -wide coverage was not available in a few
counties, because they had only small amounts of improved rangeland, and in
the 9 counties west of the Pecos River. No problems were encountered in photo¬
graphically censusing colonies in this xeric habitat as several colonies were recog¬
nized on the available photography and inspected on-site.

Because of incomplete photo coverage, this management survey was discussed
with many reliable individuals including the: County Agricultural Extension
Service, Texas Parks and Wildlife Department, Soil Conservation Service, ASCS,
State and Federal Animal Damage Control personnel, and landowners. These in¬
dividuals provided valuable information concerning many colonies that were not
recognizable on photographs or that were in areas not photographed.

Of the total colonies located, 319 (23.9% of the total) were inspected on-site
to verify the colony’s presence, size, and location. The involved are a was measured
by automobile odometer or by pacing.

RESULTS

The black -tailed prairie dog in Texas continues to occupy the same general
area described by Bailey (1905); the present boundary was determined to be west
and north of aline from Henrietta south through Palo Pinto, Coleman, and Chero¬
kee and west through Sonora, Marathon, and Sierra Blanca (Fig. 3).

Figure 3. The present range parallels the species’ 1905 range by Bailey (1905).

BLACK-TAILED PRAIRIE DOG

37

In the 108 county survey area, 1 ,336 prairie dog colonies were recorded totaling
36,432 ha in 89 counties (Fig. 3). Of this number, 1,144 colonies comprising
31,215 ha were located from aerial photographs. The remaining 192 colonies,
consisting of 5,217 ha were located through contacts with individuals familiar
with the area or while traveling to or from on-site inspections of other colonies.
In counties with prairie dogs, the mean number of colonies was 15.16 ± 3.70,
and the average area occupied by prairie dogs/county was 409.3 ha. The average
colony occupied 27.27 ha. The smallest colony contained a few burrows. The
largest was a Schleicher County colony that was estimated by the landowner to
cover 7.8 km2 ; accurate ground measurement of this colony was impossible due
to its meandering boundary.

Counties in the Texas Panhandle bordering New Mexico have the greatest
number of colonies. Dallam County (106 colonies, 3,5 14 ha) and Bailey County
(80 colonies, 2,458 ha) have the largest populations in the State. Each county’s
population decreased progressively in an easterly direction to the fringe of the
specie’s range.

On-site inspections of 229 colonies located on the photographs indicated that:
45 colonies totaling 2,650 ha (8.5% of total) were no longer active; 13 colonies
had decreased from 1 ,174 to 492 ha; and 30 colonies had increased from 1,068
to 2,899 ha. During on-site inspections, all the burrowing owls {Speotyto cunicu-
laria) observed were tabulated because of their interest to scientists and their
status as a colony -associated species.

Burrowing owls were not found on the colony surface in all seasons or weather
extremes. This temporary absence may be further extended by their reproductive
processes (Martin, 1973a), diurnal and crepuscular habits (Thomsen, 1971), mi¬
gratory activities (Brenckle, 1936), or nocturnal activities; nevertheless, a total of
1,268 owls were counted in 91 colonies totaling 3,064 ha— an average of 1 owl/2.39 ha.
That colonies containing owls were widely scattered throughout the study area
indicates this average may be representative of the study area.

Time available for the study prevented lengthy investigations for the black¬
footed ferret ( Mustela nigripes). During on-site colony inspections, no ferret or
ferret signs, as described by Fortenberry (Biologist, U.S. Fish and Wildlife Service,
pers. comm., 1971) and Hillman (1968), were observed.

DISCUSSION

Many authors have not adequately described all components responsible for
the reduction of the prairie dog from its historical population. It is a fact that
land-use changes were, and continue to be, a factor in the decline.

The 89 counties in which prairie dog colonies were located comprise a total
of 25,497,831 ha (254,978 km2 ). Water areas (over 16.2 ha) and urban build-up
totaled 657,177 ha or 2.6% of the land area. The remaining “Inventoried Area”
(IA) (Fig. 4), 24,838,042 ha, is further divided as follows: cropland, 7,235,324 ha

38

THE TEXAS JOURNAL OF SCIENCE

(29.1% IA); rangeland, 16,539,1 15 ha (66.6% IA); and improved pasture, forest,
and other, 1,063,603 ha (4.2% IA) (Soil Conservation Service, 1970). The range-
land figures for the 89 counties would not, in their entirety, be optimum prairie
dog habitat due to land slope and soil composition. More importantly, the present
land-use patterns outlined indicate a great loss of prairie dog habitat ; neverthe¬
less, on the basis of the rangeland figures, 1 ha of prairie dog colony exists per
182 ha of rangeland throughout the prairie dog range in Texas.

Figure 4. Percent of Inventoried Area (excludes water, urban build-up) of county in
agricultural crops in study area (Soil Conservation Service, 1970).

Bailey (1905) described a single colony stating that, “They (prairie dogs) cover
the whole country . . . from San Angelo (Tom Green County) north to Clarendon
(Donley County) in a strip approximately 100 mi wide by 250 mi long (64,750 km2 ).”
This area was estimated and plotted on Fig. 3. The estimated area covers 65,783 km2 .
Present urban build-up and water cover 1.9% of the area. The IA is: improved
pasture, forest, and other uses, 2.6%; cropland, 27.5%; and rangeland, 69.9%
(Soil Conservation Service, 1970). It is evident that this colony would be seriously
disrupted under the present land-use patterns.

During the on-site inspections, it became apparent that colonies were continuing
to be destroyed by changes in land use. The largest was a colony in Lynn County.

BLACK-TAILED PRAIRIE DOG

39

The 1969 aerial photos showed a 607 ha colony in a mesquite thicket. The 1972
site inspection showed an excellent stand of cotton in the former 7.8 km2 pasture.
Without nearby pastureland, this colony probably died from a lack of suitable
habitat for dispersal or emigration.

However, the Kansan biotic province (Fig. 1) in Texas contains thousands of
playa lakes in intensely farmed agricultural counties (Fig, 4). These unfarmed
playas are inundated many times each year. Most of them contain some grass
slopes between the lake bottom and plowed fields. In intensely farmed areas,
such as Castro County, these playa edges constitute the majority of the available
prairie dog habitat.

Castro County contains a total of 223 ,483 ha with only 28 ,382 ha of rangeland
(Soil Conservation Service, 1970). Luther Geiger (Soil Scientist, Soil Conservation
Service, pers. comm., 1973) reports that the county has over 500 playa lakes and
that 6,459 ha are inundated many times each year. This leaves about 21 ,923 ha
of rangeland for prairie dogs. The county has 41 colonies totaling 699 ha; most
are small and occur in the fringe vegetation around playa lakes. Since prairie dogs
in playa edge colonies do little harm, they are not controlled. The colony cannot
expand due to the barriers of plowed fields and the lake bottom, and thus remain
relatively static over the years. Extremely drastic land-use changes in playa lakes
must occur to remove these reservoirs of prairie dog populations in this biota.

Several cities in the study area, such as Amarillo and Lubbock, and several
cities outside the study area, such as Ft. Worth and Tyler, have small transplanted
colonies in local parks that were not included in the survey. However, 6 cities in
the study area, such as Plainview and Sweetwater, have colonies that appeared to
naturally occur on vacant lots or grasslands within the cities’ limits. A total of
10 colonies occupying 31 ha were located and included in this survey. Colonies
in such locations have high esthetic value and their natural existence contributes
to the State’s native population. That such colonies occur may be viewed as an
inherent tenacity of the species to continue perpetuation.

The State’s native prairie dog population can only be estimated as the number
of animals collectively inhabiting any colony are constantly fluctuating. Some
relative influences affecting density/ha are: the abundance of lack of vegetation,
weather, and counts made before or after parturition or winter mortality or the
emigration of some adults. Merriam (1902) reported, “The number of holes
on each acre varies from a few to upward of a hundred, and probably averages at
least 25. It is certainly a conservative estimate to assume the average number of
animals/acre to be 25.” Assuming this estimate is presently valid, it may be stated
that 2,249,000 prairie dogs inhabit the State and that they continue to occupy
the same general area in Texas described at the turn of the century.

Such a population certainly cannot be classified as rare within its range in
Texas or endangered as a species. That the prairie dog has continued is evident—
that it will continue, is reasonable.

40

THE TEXAS JOURNAL OF SCIENCE

ACKNOWLEDGEMENTS

I am indebted to many personnel of the Agricultural Stabilization and Conser¬
vation Service, the Soil Conservation Service, and the Texas Parks and Wildlife
Department for their cooperation. I am also indebted to U.S. Fish and Wildlife
Services colleagues within the Division of Wildlife Services and particularly Gilbert
Marrujo, Howell Ellard, Stephen Atzert, Steve Smith and Milton Caroline who
provided inestimable support.

LITERATURE CITED

Bailey, V., 1905 -Biological survey of Texas North American Fauna. U.S. Dept, of Agri.,
Washington, No. 25.

Blair, W. F., 1950-The biotic provinces of Texas. Tex. J. ofSci., 2:93.

Brenckle, J. F., 1936-The migration of the western Burrowing Owl. Bird Banding, 7:166.

Cheatheam, L. K., 1972-Censusing prairie dog colonies using aerial photographs. Proc. Black¬
footed Ferret /Prairie Dog Workshop, S. Dakota S. Univ., Brookings, p. 78.

Cottam, C., and M. Caroline, 1965 -The Black-tailed Prairie Dog in Texas. Tex. J. Sci., 17(3):294.

Davis, W. B., 1960-77m Mammals of Texas. Tex. Game and Fish Comm., Austin, 252 pp.

Drimmer, G., 1954-77?e Animal Kingdom. Vol. 1. Graystone Press, New York, N.Y., 680 pp.

Hillman, C. N., 1968-Field observations of Black -footed Ferrets in S. Dakota. Trans, of 33rd
N.Am. Wildl. and Nat. Res. Con/. Pub. by : Wildl. Mgmt. Inst. p. 433.

Martin, D. J., 1973a-Selected aspects of Burrowing Owl ecology and behavior. Condor,
73:446.

Merriam,C., 1902-The Prairie Dog of the Great Plains. Yearbook on Agriculture, U.S. Dept.

of Agri., Washington, pp. 257-270.

, 1 ^

Rue, L., Ill, 19 67 -Pictorial Guide to the Mammals of North America. Thos. Crowell Co.,
New York, N.Y., 299 p.

Soil Conservation Service, 1970-Conservation needs inventory. Soil Conservation Service,
Temple, 297 p.

Thomsen, L., 1971 -Behavior and ecology of Burrowing Owls on the Oakland Municipal
Airport. Condor, 73:177.

YIELD ESTIMATES DERIVED FROM ACTIVE AND PASSIVE
CREEL SURVEYS OF A SMALL POND FISHERY1

by FREDRICK V. JONES2 , WILLIAM D. PEARSON3 , and
LLOYD C. FITZPATRICK

Department of Biological Sciences,

North Texas State University , Denton 76203

ABSTRACT

Estimates of angler harvest from a small pond were made by active and passive (voluntary¬
reporting) survey techniques. According to the active survey, anglers removed 789 fish weigh¬
ing 61.3 kg (64.5 kg/ha) from the pond, made 577 fisherman visits to the pond, and caught
1.06 fish/hr during 1973-74. The passive survey method underestimated fishing pressure
(no. anglers) and yield (no. fish) but gave an estimate of catch rate similar to that of the
active surveys.

INTRODUCTION

Fishery managers often rely upon either voluntary reporting of catch informa¬
tion or direct, in-the-field, creel surveys to estimate fish harvest rates. Such studies
are usually confined to the largest and most important fisheries in the manager’s
region. Comparisons of harvest data collected by voluntary reporting, and an
active creel survey in which the angler is actually observed by the biologist as a
census technique, usually confirms the well-known tendency for anglers to exag¬
gerate (Carlander, et al., 1958; Carline, 1972; and Radford, 1973). During an in¬
tensive study of energy flow in a small pond ecosystem (Jones, 1975) we made
estimates of fish harvest rates by both passive (voluntary reporting) and active
creel survey methods and were subsequently able to compare the two sets of
data in light of concurrent measurements of fish production and biomass in the
pond.

THE STUDY AREA

The study was conducted on a 0.95 ha pond located on the North Texas
State University golf course. The pond is located just inside the city limits of

financial aid was supplied to William D. Pearson and Lloyd C. Fitzpatrick by faculty re¬
search grants at North Texas State University.

2Present address: Department of Wildlife and Fisheries Sciences, Texas A&M University,
College Station 77843.

3 Present address: Water Resources Laboratory, University of Louisville, Louisville, Kentucky
40208.

Received for publication: October 8, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

42

THE TEXAS JOURNAL OF SCIENCE

Denton, Texas, off Interstate Highway 35 and on Avenue D. Avenue D, a gravel
road, runs along the east side of the pond, 1 .5-6.1 m from the shore. The other
sides of the pond are bordered by the course fairways.

In most years the pond was capable of supplying all irrigation water needed
by the golf course, but drawdowns in summer months sometimes removed 70%
of the total storage volume. Prior to 1973, natural runoff to the pond was supple¬
mented by water purchased from the City of Denton. In 1972, a well was drilled
30.6 m southeast of the pond to provide pumped water for the increased demands
of the golf course. During this study , well water was pumped into the pond, but
problems with the pumping mechanism caused drastic fluctuations in water level
in summer and fall months. By late fall the pump was working properly and draw¬
downs were less severe.

The pond was constructed in 1947 and Bill White, fisheries manager at the
Lewisville, Texas, fish hatchery, stocked bluegill sunfish (. Lepomis macro chirus),
largemouth bass (Microp terns salmoides), and channel catfish (Ictalurus punctatus )
in unknown numbers in 1948. Channel catfish were last taken from the pond in
1970, when two 4.5-kg specimens were captured, marked, and returned to the
pond. No catfish were collected during this study. No additional fish have been
stocked by the hatchery, but the superintendent of the golf course added 7 large
(0.7 -0.9 kg) white crappie ( Pomoxis annularis) from Lake Dallas in 1970, and
other additions are assumed to be accidental. During this study there were 7 species
of fish in the pond: black bullhead (Ictalurus melas), largemouth bass, white
crappie, bluegill, green sunfish ( Lepomis cyanellus), longear sunfish ( Lepomis
megalotis), and golden shiner ( Notemigonus chrysoluecas). Fishing in the pond
was prohibited for many years until the summer of 1973, when fishermen were
allowed to fish on the east side of the pond next to the road.

METHODS AND MATERIALS

The passive creel survey began in July 1973 with the installation of two data
boxes or registers on the east side of the pond in full view of anyone approaching
from the road and concluded in May, 1974. The registers were equipped with cards
for recording number of fishermen, gear used, number of hours fished, and num¬
ber, species, and length of all fish caught. Pencils and rulers for measuring the
fish were also provided. Lengths were converted to weights using length -weight
relationships derived from seine samples. Signs posted near the boxes asked
fishermen to fill in the cards and deposit them in the boxes with any fish tags
collected. We tried to contact as many fishermen as possible and asked them to
cooperate in the voluntary reporting.

The active creel survey employed a stratified monthly sampling regime similar
to one described by Best and Boles (1956). At least 2 weekdays and 1 weekend
day of each week were sampled in each month from June 1973 to May 1974.
The selection of days within these restrictions was made randomly. Both mornings

ACTIVE AND PASSIVE CREEL SURVEYS

43

and evenings were sampled on each day selected. Sampling was from 0600 hrs to
2100 hrs. At each 3 -hr period we would travel to the pond and observe the fisher¬
men for 1 hr. Data were collected on type of gear used, bait, number of fisher¬
men, and amount of gear. The fishermen were checked for catch and those still
fishing from the last period were questioned as to how long they had fished, and
if any other fishermen had come and gone during the interval. Night fishing was
prohibited by the golf course attendants and was not monitored beyond 2100 hrs.
We believe there was very little illegal night fishing. Each category (weekday and
weekend) value was then expanded to estimate total harvest figures for each
month.

Fish collected for research purposes (stomach analysis, respiration studies,
etc.) were tabulated and included in the total harvest figures.

RESULTS

Active Creel Survey

Although fishing was prohibited until June 1973, enforcement of the prohibi¬
tion had always been rather lax and some fishermen seemed to make a practice
of fishing for 30-60 min until asked to leave by the course attendants. When fish¬
ing was no longer prohibited after June 1973, these fishermen began to fish the
pond steadily, and were joined by newcomers. Catches were high in June and
July 1973, when 538 fish (68 .2% of the annual catch of 789), amounting to 38.1 kg
(62.2% of the total harvest weight of 61.3 kg) were caught (Table 1). The catch
decreased steadily during the rest of the summer and fall. In winter months the
catch rate increased slightly and then fluctuated somewhat during the late winter
and spring.

Black bullhead made up 75.3% of the total catch by numbers (594 fish) and
83.2% (51.0 kg) of the catch by weight according to the active census. Adult
bass(l 1) and crappie (19) were the largest fish captured, with a 2.36 kg crappie
being the largest fish recorded. Although sunfish were the second most abundant
fish captured, they accounted for only 3.3 kg of weight removed.

The number of fishermen using the pond decreased through the year. Of some
738 fishermen visits to the pond, 415 were in June and July 1973. The number
of fishermen visits decreased during fall and win ter, then increased again in spring,
but never reached the high of the previous summer. Fishing occurred primarily
in the afternoon from 1400 hr to sunset. Only 2 fishermen were observed fishing
at night, and very little fishing occurred during the morning (6.5%). Weekend
fishing was more popular than weekday fishing throughout the study. Summer
fishermen used both cane poles and rod-and-reel combinations with natural or
preserved food baits. During the fall and winter most fishermen used only 1 pole,
while in summer and spring the average fisherman used 1.1 -1.5 poles. One man
tended 10 cane poles at one time. The highest catch rates (2.31 fish/hr) occurred
in May. The average catch rate for the entire year was 1 .06 fish/hr.

44

THE TEXAS JOURNAL OF SCIENCE

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Passive Creel Survey

Table 2 presents the data derived from the passive creel survey in which the
fishermen voluntarily supplied information on catches. Only 1 12.8 hr of fishing
and 138 fish, weighing 42.6 kg, were reported. Black bullhead accounted for

ACTIVE AND PASSIVE CREEL SURVEYS

45

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most of the catch (72 fish; 26.4 kg), followed by sunfish (39 fish; 5.4 kg), large-
mouth bass (8 fish; 7.9 kg) and crappie (19 fish; 2.9 kg). No fish were reported
in October and January, while only 1 fish/month were reported in July, November,
and December. The low value in July is attributed to the fact that many fisher¬
men were still unaware of the survey.

46

THE TEXAS JOURNAL OF SCIENCE

The late summer catch rates reported in the passive survey (1 .26-1 .64 fish/hr)
were comparable to those determined in the active survey. During the fall and
winter catch rates declined to 0-0.33 fish/hr and then increased steadily to 2.03
fish/hr in April. The mean catch rate over the entire year was 1 .22 fish/hr.

Fish Removed for Research

Many fish were removed from the pond for research purposes (i.e., stomach
analysis and respiration studies conducted by other students). Table 3 presents
a summary by species of the 5,662 fish removed for research purposes (28.9 kg),
which was almost half as much as that cropped by fishermen (61 .3 kg).

TABLE 3

Number and Weight of Fish Removed from the Pond for Experimental Purposes.

Species

Number

Weight (kg)

Black Bullhead

2,992

5.9

Bluegill

554

4.0

Green Sunfish

824

7.7

Longear Sunfish

39

0.5

Largemouth Bass

246

2.3

White Crappie

842

5.9

Golden Shiner

165

2.6

Total

5,662

28.9

Of the fish removed for research purposes, green sunfish accounted for the
largest weight removed (7.7 kg), followed by bullhead and crappie (5.9 kg each).
The total removals of bluegill, golden shiner, largemouth bass, and longear sun-
fish were much lower (0.5 -4.0 kg).

DISCUSSION

The pond illustrated a classic example of a new fishery, with very high initial
fishing and catch rates, followed by sharp reductions as fishing increased above
optimum for. the system. From catch data it was obvious that many of the large
bullheads were fished out by late August, as were the bass and crappie.

The active creel survey has usually been considered to be more reliable than
the passive or voluntary-reporting survey (Best and Boles, 1956; Carlander,ef al.,
1958; Stewart and Trout, 1973). In this study the active survey resulted in an
estimated harvest of 789 fish weighing 61.3 kg, while the passive survey reported
only 138 fish weighing 42.6 kg. Thus the passive survey reported only 17% of
the number of fish estimated by the active survey, but the weight reported in
the passive survey was nearly 70% of that obtained in the active survey. These?
discrepancies probably result from the tendency of fishermen to report only
catches of large fish, to overestimate the size of fish taken, or both. The passive

ACTIVE AND PASSIVE CREEL SURVEYS

47

survey also underestimated the total amount of fishing pressure on the pond (5 1
fisherman visits compared to 577 for the active survey). Catch rate ranges reported
by the 2 methods were comparable (0-2.31 fish/hr for the active and 0-2.03 for
the passive).

The active survey estimate of total annual harvest rate was 830 fish/ha or
64.5 kg/ha. These figures are several times greater than the 62.8 fish/ha and 16.4
kg/ha mean annual sport fishing harvest rates reported for 130 U.S. reservoirs by
Jenkins and Morais (1971). The reservoirs surveyed by Jenkins and Morais were
all much larger (<202 ha) than the golf course pond and might be expected to
yield fewer fish/ha. Bennett (1971) and Childers and Bennett (1967) have reported
annual angler harvest rates of 3-76.7 kg/ha for fishes in ponds of 1-10 ha in the
midwestern and southwestern U.S., although yields exceeding 50 kg/ha were rare.
The annual harvest from the golf course pond in 1973-74 ranks among the highest
for an uncultured system in the United States.

Jones (1975) reported a total annual production for all species of fish in the
pond of 1,986 kg or 2,090 kg/ha, and a mean standing crop of 1 ,050 kg or 1 ,105
kg/ha. Therefore, fishermen removed only 3.1% of the total annual production.
If we include the removals for experimental purposes in the annual harvest, it
still amounts to only 4.5% of the annual production. However, Jones (1975) has
also pointed out that most (83.8%) of the total annual production in the pond
was laid on by young-of-the-year fishes which were not susceptible to the fisher¬
men’s gear. The annual yield to fishermen was 28.5% of the annual production
of adult fish alone (Age Class I+).

We suggest that the passive creel survey method will greatly underestimate
fishing effort and total harvest in a sport fishery, although it may give an accurate
estimate of the catch rate. Yield to anglers in the pond was very high in the first
year of legalized fishing, but probably cannot be expected to be maintained in
subsequent years.

LITERATURE CITED

Bennett, G. W., 1971 -Management of Lakes and Ponds. Van Nostrand Reinhold Co., New York,
N.Y., 375 pp.

Best, E. A., and H. D. Boles, 1956-An evaluation of creel census methods. Calif. Fish and
Game, 42(2):109.

Carlander, K. D., C. J. DiCostanzo, and R. lessen, 1958— Sampling problems in creel census.
Prog. Fish-Cult., 20(2):73.

Carline, R. F., 1972-Biased harvest estimates from a postal survey of a sport fishery. Trans.
Amer. Fish. Soc., 101(2): 262.

Childers, W. F., and G.W. Bennett, 1967 -Hook -and-line yield of largemouth bass and redear
x green sunfish hybrids in a one-acre pond .Prog. Fish-Cult. , 29(1): 27.-35.

Jones, F. V., 1975 -The population dynamics and trophic relationships of seven species of
fish in a small Southwestern pond: with special attention toward young-of-the year fish.
Unpubl. M.S. Thesis, North Texas State Univ., Denton, 192 pp.

48

THE TEXAS JOURNAL OF SCIENCE

Jenkins, R. M., and D. I. Morais, 1971 -Reservoir sport fishing effort and harvest in relation
to environmental variables. In G. E. Hall (Ed.), Reservoir Fisheries and Limnology. Spec.
Publ. No. 8, Amer. Fish. Soc., Washington, D.C., pp. 371-384.

Radford, D. S., 1973- A comparison between actual and reported angling times. Trans. Amer.
Fish. Soc., 102(3): 641.

Stewart, R. W., and J. R. Trout, 197 3 -Evaluation of a voluntary angler creel census pro¬
cedure. New Jersey Dept. Envir. Port. Div. Fish Game Shell Fish., Freshwater Fish.
Lab., N.J. Mis. Rept. No. 38. 52 pp.

AGE AND GROWTH OF LARGEMOUTH BASS IN CANYON
RESERVOIR, TEXAS

by JOE W. KOLB1 and B. G. WHITESIDE
Aquatic Station,

Southwest Texas State University ,

San Marcos 78666

ABSTRACT

The method of aging largemouth bass by their scales was validated for a central Texas
reservoir. From March 1974 to April 1975, 838 fish were collected from Canyon Reservoir
with bass tournaments being the major source of fish 305 mm or longer in total length. No
substantial difference was found in the growth rates of males and females, however, the oldest
females found were 6 years old and the oldest male was 5 years old, with age groups 4 and 5
containing mostly females. Mean total lengths (mm) of combined sexes for age groups 1-6
were 161, 269, 326, 392, 459, and 510, respectively. Mean weight increments (g) of com¬
bined sexes for age groups 1-6 were 45, 175, 179, 308, 446 and 446, respectively. Growth
rates of largemouth bass from Canyon Reservoir are compared to those from other states.
The length-weight relationship was log W = -5.2005 + 3.1039 log L, where W = weight in
g and L = total length in mm, and the bass exhibited allometric growth. A condition factor
of 4.0 (SE = 0.02) indicated the fish were in poor flesh.

INTRODUCTION

Age and growth studies have been used as fisheries management tools for many
years and in many parts of the U.S. They have been used not only to determine
the existence of a problem, but also to evaluate the results of applied manage¬
ment practices.

The scale method of determining the age and growth of fishes is generally
accepted as valid for fishes in northern states. However, its validity for fishes in
southern states has been in question for many years due to mild winters (Prather,
1967), since for an annulus to be formed, growth during the winter must cease
for at least 14 days (Bennett, 1970).

The objectives of this study of largemouth bass, Micropterus salmoides, in
Canyon Reservoir were to validate the use of the scale method for aging large¬
mouth bass in Canyon Reservoir; to determine the age composition, length -weight
relationship, and condition factor.

Present address: Texas Water Quality Board, 2318 Center Street, Deer Park 77536.
Received for publication: November 22, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

50

THE TEXAS JOURNAL OF SCIENCE

METHODS AND MATERIALS

Canyon Reservoir is a deep-storage impoundment in Comal County about 26 km
west of San Marcos, Texas. It was impounded in June 1964 and has a 3,300 ha
surface area and 40 m maximum depth.

Specimens were collected from March 1974 to April 1975 by seining, hook
and line, rotenoning of coves and gill netting. Bass tournaments were the major
source of fish 305 mm (12 in) or longer; most smaller fish were taken by seining.
Letters sent to bass clubs within a 210-km radius of the study area requested
permission to obtain data from fish caught in their Canyon Reservoir tourna¬
ments. Many clubs responded to this request and were very cooperative at the
tournament weigh-ins.

When possible, total length (mm), weight (g) and sex were determined for each
specimen, recorded on a numbered scale envelope and a scale sample from that
individual placed in the envelope. The scale sample contained 5 to 8 scales removed
from a key area located at the tip of the depressed left pectoral fin and 2 to 4
scale rows below the lateral line. Scale impressions were made on a cellulose ace¬
tate slide with a roller press (Smith, 1954) and the slide identified by number.
The impressions were examined using a scale reading machine (Van Oosten, £/•#/.,
1934), without knowledge of the size of the fish from which the scales came. All
slides were reread without knowledge of the first or subsequent readings until
there was agreement between two readings for each slide. The slides were examined
in groups of at least 75 to reduce the possibility of bias due to remembering pre¬
vious examinations of the same slide. According to Carlander (1961), repetition
of readings may increase the accuracy of age determinations.

After the age of fish was determined, the positions of the focus, each annulus
and the margin of a representative scale measured along the midanterior field of
the scale at 60x were marked on a strip of paper along with the fish identification
number, total length and sex. Each strip of paper was used in a nomograph method
for the back -calculation of the total length of the fish at the end of each year of
life (Carlander and Smith, 1944). To determine the y -intercept on the nomograph,
a body -scale relationship was calculated on measurements from 686 largemouth
bass from Canyon Reservoir using the following equation:

L = a + bS

where L is the total length (mm) of the fish , S is the distance from the focus to
the anterior margin of the scale (mm x 60), a is they -intercept on the nomograph
and b is the slope. Since scale measurements were used to derive total lengths of
fish, the scale measurements were designated the independent variable (Whitney
and Carlander, 1956). Characteristics used for the recognition of annuli and
false annuli were according to Prentice and Whiteside (1975).

A length-weight relationship was calculated on data from 838 largemouth
bass of both sexes with total lengths which ranged from 34 to 563 mm with a

LARGEMOUTH BASS

51

mean of 179 mm and weights which ranged from 0.6 to 2523 g with a mean of
64 g by using the formula:

log W = a + blogL

where W is the weight (g), L is the total length (mm), a is they -intercept and b is
the slope. A two-tailed student’s Mest was used to test the hypothesis that the
slope of the length -weight relationship line was equal to 3.0 (Steel and Torrie,
1960).

A condition factor for all fish was calculated according to the following equation:

G F = 1 0,000 If1
L3

where W is the weight (g) and L is the total length (mm) (Bennett, 1970).

RESULTS AND DISCUSSION

Length-Weight Relationship and Condition Factor

If the slope of a length -weight relationship line is close to 3.0, fish growth is
isometric (no change in body shape with a change in length), and the condition
factor should not vary with changes in length of the fish. If the slope is not equal
to 3.0 then growth is allometric (a change inhody shape with a change in length),
and the condition factor should vary with changes in length of the fish. However,
in both types of growth there is individual variability (Tesch, 1968; Rounsefell
and Everhardt, 1962). The length -weight relationship equation for largemouth
bass from Canyon Reservoir was:

log W= -5.2005 + 3.1039 logL.

The difference between the slope of the length-weight relationship line for large¬
mouth bass in Canyon Reservoir (b = 3.1039) and 3.0 was highly significant (p<
0.001 ; 836 df), indicating that there should have been an increase in the condition
factor with an increase in length. An increasing condition factor trend with an
increase in length was also found, further indicating that growth was allometric.

Bennett (1970) gave the following ranges of condition factors for largemouth
bass between 127 and 381 mm in total length: 3.5-4 .5 = a fish in poor flesh,
4.6-5 .5 = a fish in average flesh, 5. 6-6.5 = a very fat fish. The mean condition
factor for 462 largemouth bass between 127 and 381 mm in total length in Canyon
Reservoir was 4.2 (SE = 0.03). According to Bennett’s ranges, the bass population
in Canyon Reservoir was below average condition, indicating a potential growth
problem. The overall mean condition factor was found to be 4.0 (SE = 0.02) for
804 largemouth bass from Canyon Reservoir between 34 and 563 mm in total
length.

A total length (TL) to standard length (SL) conversion equation, SL = 1 .7379
+ 0.8195 TL, was calculated from 128 Canyon Reservoir largemouth bass ranging
in total length from 60 to 522 mm.

52

THE TEXAS JOURNAL OF SCIENCE

Aging Trials

Although criteria from recognizing annuli and supernumerary marks are well
established, some scale marks were difficult to interpret- Age analysis results
(Table 1) show that, although agreement of two readings was obtained by the
second reading for a majority of fish (70%), many fish required a third reading
(26%) and a few fish required a fourth reading (4%). The necessity for more read¬
ings with an increase in age was due to increased difficulty in distinguishing annuli
that formed closer together in later years of life. This same phenomenon was found
by Prentice and Whiteside (1975) in a study of largemouth bass of known age in
central Texas farm ponds, yet they were able to attain a 94% aging accuracy.

TABLE 1

Number and % Agreement of Aging Trials for Age Groups of Largemouth Bass
Collected from Canyon Reservoir, Texas, in 1974-75.

Age

No. of fish in

age group

No.

of individuals with agreement of 2 readings by

2nd Reading

3rd Reading

4 th Reading

1

111

79

(71)a

29

(97)a

3

(100)a

2

312

244

(78)

61

(98)

7

(100)

3

115

69

(60)

39

(94)

7

(100)

4

42

19

(45)

19

(90)

4

(100)

5

9

2

(22)

4

(67)

3

(100)

6

5

0

( 0)

3

(60)

2

(100)

Total

594

413

(70)

155

(96)

26

(100)

aN umbers in parentheses are % agreement by that reading.

Validation of Scale Method

Many fisheries workers assume that Texas fishes do not form annuli each year,
therefore, the scale method has seldom been used in the area. Silvey and Harris
(1948) found that the scale method of age determination for largemouth bass
was reliable in east Texas provided that a sufficient number of fish were examined.
Prentice and Whiteside (1975) studied the validity of aging largemouth bass in
central Texas farm ponds and showed that the criteria for validation of the scale
method were satisfied.

To validly use the scale method for age determination it is necessary to deter¬
mine the approximate time and regularity of annulus formation each year (Hile,
1941 ; Berg and Grinnaldi, 1967). A small percentage of Canyon Reservoir large¬
mouth bass appeared to have formed annuli by the latter part of January and
February, a large percentage formed annuli during March and most had formed
annuli by the end of April or May (Table 2). A trend in the time of annulus forma¬
tion for different age and length groups could not be determined due to small
numbers of fish collected each month.

To check for regularity of annulus formation, back-calculated total lengths of
bass at different ages were examined for similarity to the empirical total lengths

LARGEMOUTH BASS

53

TABLE 2

Time of Annulus Formation for Largemouth Bass Collected
from Canyon Reservoir, Texas, in 1974-75.

Collection

Date

Total No.
Collected

% Fish With
Annulus

March 29 & 30, 1974

19

47

May5, 1974

150

100

May 17, 1974

14

100

January 22 & 24, 1975

18

11

February 21, 1975

27

11

March 19, 1975

18

33

March 29,1975

8

38

April 6 & 9, 1975

56

59

April 12, 1975

53

70

April 27, 1975

34

94

of bass at that age. Back -calculated total lengths of fish were obtained by using
a nomograph method with ay-intercept correction of 36.7 mm ( L = 36.715 +
1.087 S). The mean empirical total lengths (mm) of the bass in age groups 1 +
through 6+ were found to be 245, 303, 351, 409, 469 and 517 respectively,
and were similar to the back -calculated grand mean total lengths (weighted for
individuals) of the bass at each corresponding annulus (Table 3).

Comparison of Age and Growth Between Sexes

No substantial difference in growth rates between male and female bass was
found when using a common y -intercept correction factor for back-calculating
total lengths (Table 4). Separate y-intercept correction factors for back -calculation
of total lengths for each sex were not calculated since few small bass could be
sexed. Stroud (1948), in a study of Norris Reservoir, Tennessee, found that older
and larger largemouth bass tended to be females. Padfield (195 1), in a study in
Alabama, found that male largemouth bass were few or absent in the older age
groups. In Canyon Reservoir the relative abundance of males showed a sharp de¬
crease in age group 4 and males were absent in age group 6 (Table 4).

Comparison of Growth Rates in Canyon Reservoir with Those of Other Areas in
the United States

In many age and growth studies of largemouth bass, a zero y-intercept was
assumed when back-calculating lengths of fish. For the purpose of comparison
with these studies and studies that used a corrected y-intercept, total lengths
were back -calculated using both a zero y-intercept and a corrected y-intercept
(Table 3). Largemouth bass in age groups 1 through 4 from Canyon Reservoir
had lower growth rates than largemouth bass from Louisiana, Oklahoma, Arkansas
and Tennessee, while older age groups had growth rates similar to those from Okla¬
homa, Arkansas and Tennessee (Table 5). However, the growth rates of largemouth
bass in Canyon Reservoir were greater than those in more northern states.

54

THE TEXAS JOURNAL OF SCIENCE

TABLE 3

Back -calculated total lengths (mm) at the end of each year of life for large-
mouth bass collected from Canyon Reservoir, Texas, in 1974-75. Total lengths
were obtained using a corrected y -intercept of 36.7 mm. Numbers of indi¬
viduals used in calculations are given in parentheses.

Year

Age Groups

Class

1

2

3

4

5

6

1974

181

(16)

1973

177

283

(248)'

(153)

1972

155

281

352

(191)

(191)

(32)

1971

137

237

325

448

(94)

(94)

(94)

(ID

1970

135

229

310

373

478

(35)

(35)

(35)

(35)

(4)

1969

162

247

304

390

440

(5)

(5)

(5)

(5)

(5)

1968

140

251

340

404

462

510

(5)

(5)

(5)

(5)

(5)

(5)

Grand mean total length (weighted for individuals)

161

269

326

392

459

510

Grand mean total length increment

161

108

57

66

67

51

aGrand mean weight increment

45

175

179

308

446

446

Mean annual total length increment (weighted for year classes)

161

104

77

84

71

48

Sum of mean annual total length increment

161

265

342

426

497

545

^Grand mean total length (weighted for individuals)

142

260

324

389

457

509

Total number of fish

594

483

171

56

14

5

a

b

These weight increments were calculated using the length-weight relationship equation.
These back -calculated total lengths were obtained using an assumed zero y -intercept.

The general increase in the rate of growth of fish from north to south is usually f,
attributed to an increase in the length of growing season (Bennett, 1937; Viosca, !
1942; Stroud, 1948; Jenkins and Hall, 1953). However, Jenkins (1974), in an
analysis of large reservoirs in the southern U.S., made some general statements ;1!
concerning the factors governing the productivity of water. Generally the highest

LARGEMOUTH BASS

55

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production of sport fishes occurs where all of the following criteria are found:

(1) mean annual concentration of total dissolved solids is from 120-350 ppm,

(2) dominant ions are calcium-magnesium and carbonate-bicarbonate and (3) mean
annual suspended sediment load of inflowing streams is less than 2000 ppm.
Hannan and Young (1974), Jenkins (1974) and Rawson (1968) indicated that
the Canyon Reservoir environment met the three above criteria. Jenkins (1974),
further stated that if the above criteria are met then the length of the growing
season is the major factor controlling sport fish production and Canyon Reservoir
has a growing season of approximately 260 days.

56

THE TEXAS JOURNAL OF SCIENCE

TABLE 5

Comparison of grand mean total lengths weighted for individuals (mm)
of largemouth bass in U.S. waters. An assumed zero y -intercept was used
in calculating lengths.

Age Groups

Location

Reference

1

2

3

4

5

6

7

8

Central Texas

This study

142

260

324

389

457

509

Louisiana

Carlander, 1950

193

287

368

478

531

597

630

658

Oklahoma
(Reservoir mean)

Jenkins and

Hall, 1953

170

290

361

417

465

498

528

554

Bull Shoals
Reservoir, Ark .

Bryant and
Houser, 1971

176

297

377

427

457

492

519

524

Norris Reservoir,
Tenn.

Stroud, 1948

175

310

371

406

445

490

528

Nebraska

Carlander, 1950

91

193

277

343

401

447

480

503

Ohio

Carlander, 1950

89

178

257

318

368

409

450

480

Wisconsin

Bennett, 1937

94

188

267

318

356

384

414

442

Minnesota

Carlander, 1950

88

170

236

292

333

384

414

447

It should be pointed out that just because the younger age groups of large-
mouth bass in Canyon Reservoir have slower growth than other southern reservoirs,
that this does not mean that the bass production in Canyon Reservoir is low. In
fact, based upon Jenkins’ (1974) criteria a high production of bass would be ex¬
pected. Thus, it should be kept in mind that production is dependent upon many
factors (i.e ., growth rates, mortality rate, population size, etc.).

During the years immediately after the impoundment of Canyon Reservoir,
catches of “large” largemouth bass were commonly reported by fishermen. How¬
ever, during the present study many fishermen reported catching large numbers
of bass but few large enough to weigh-in at bass tournaments. Gill-net data from
Canyon Reservoir (J. Dean, Texas Parks and Wildlife Department, personal com¬
munication) showed a decline in the mean weight of largemouth bass from 748 g
in 1969 to 227 gin 1975. Bennett (1970) stated that “in the production of fishes
above average size, a large source of available food per individual fish seems to
exceed in importance the length of the growing season.” Many fisheries managers
assume that in southern waters available food is usually not limiting, which may
not be true, since a problem in many southern reservoirs is the overpopulation of
game and rough fish (Thompson, 1954) causing high intraspecific and Interspecific
competition.

It is our understanding that in referring to the forage fishes available to the
sport fishes, that many biologists include adult forage fishes of which many are
too large to be utilized by most sport fishes. Thus, a large portion of the rough
fish population is not available as a food source but is actually competing with
sport fishes for certain types of food. The small average size of bass in Canyon

LARGEMOUTH BASS

57

Reservoir may be due to a large population of bass and a limited supply of food
for which the bass must compete.

ACKNOWLEDGEMENTS

The authors wish to thank W. C. Young and D. G. Huffman of the Aquatic
Station, Southwest Texas State University for reviewing the manuscript, and
J. Dean and W. Butler of the Texas Parks and Wildlife Department for their assist¬
ance in collecting data.

LITERATURE CITED

Bennett, G. W., 1937 -The growth of the largemouthed black bass ,Huro salmoides (Lacepede),
in the waters of Wisconsin. Copeia, 1937:104.

- , 1970 -Management of Artificial Lakes and Ponds. Reinhold Pub. Corp., New York

and London, 375 pp.

Berg, A., and E. Grinnaldi, 1967 -A critical interpretation of the scale structures used for
the determination of annuli in fish growth studies. Memorie 1st. Ital. Idrobiol., 21:225.

Carlander, K. D., 1961 -Variations on rereading walleye scales. Trans. Amer. Fish. Soc., 90:230.

- and L. L. Smith, Jr., 1944-Some uses of nomographs in fish growth studies.

Copeia, 1944:157.

Hannan, H. H., and W. C. Young, 1974-The influence of a deep-storage reservoir on the
physicochemical limnology of a central Texas river. Hydrobiologia, 43:419.

Hile, R., 1941 -Age and growth of rock baiSS,Ambloplites rupestris (Rafinesque), in Nebish
Lake, Wisconsin. Trans. Wise. Acad. Sci. Arts Lett., 33:189.

Jenkins, R. M., 1974 -Reservoir management prognosis: migraines or miracles. Proc. 27th
Ann. Conf. SE Assoc. Game Fish Comm., (1973): 374 .

- , and G. Hall, 195 3-The influence of size, age and condition of waters upon the

growth of largemouth bass in Oklahoma. Okla. Fish. Res. Lab. Rep. No. 30, 43 pp.

Padfield, J. H., Jr., 1951 -Age and growth differentiation between the sexes of the large¬
mouth black bass Micropterus salmoides (Lacepede)./. Tenn. Acad. Sci., 26:42.

Prather, E. E., 1967-A note on the accuracy of the scale method in determining the ages of
largemouth bass and bluegill from Alabama waters. Proc. 20th Ann. Conf. SE Assoc.
Game Fish Comm., (1966):483.

Prentice, J. A., and B. G. Whiteside, 1975 -Validation of aging techniques for largemouth
bass and channel catfish in central Texas farm ponds. Proc. 28th Ann. Conf. SE Assoc.
Game Fish Comm., (1974):414.

Rawson, J., 1968-Reconnaissance of the chemical quality of surface waters of the Guada¬
lupe River Basin, Texas. Rep. No. 88, U. S. Geol. Surv., Austin, 36 pp.

Rounsefell, G. A., and W. H. Everhart, 1962 -Fishery Science, Its Methods and Applications.
John Wiley and Sons, New York, N.Y., 444 pp.

58

THE TEXAS JOURNAL OF SCIENCE

Silvey, J. K. G., and B. B. Harris, 1948-A ten-year management program on an east Texas
lake. Trans. 12th Amer. Wildl. Conf, (1948) : 25 8 .

Smith, S. H., 1954-Method of producing plastic impressions offish scales without using heat.
Prog. Fish -Cult., 16:75.

Steel, R. G. D., and J. H. Torrie, 1960 -Principles and Procedures of Statistics. McGraw-Hill
Book Co., New York, N.Y., 481 pp.

Stroud, R. H., 1948-Growth of the bass and black crappie in Norris Reservoir, Tennessee.
J. Tenn. Acad. Sci., 23:31.

Tesch, F. W., 1968-Age and growth. In W. E. Ricker (Ed.), Methods for Assessment of Fish
Production in Fresh Waters. Int. Biol. Prog., London, pp. 93-123.

Thompson, W. H., 1954-Problems of reservoir management. Trans. Amer. Fish. Soc., 84:39.

Van Oosten, J., H. H. Deason, and F. W. Jobes, 1934-A microprojection machine designed
for the study of fish scales./, de Conseil, 9:241.

Viosca, P., Jr., 1942 -Phenomenal growth rates of largemouth black bass in Louisiana waters.
Trans. Amer. Fish. Soc., 72:68.

Whitney, R. B., and K. D. Carlander, 195 6 -Interpretation of the body-scale regression for
computing body length of fish./. Wildl. Mgmt., 20:21.

THE EFFECTS OF AMNIOTIC FLUID ON THE GROWTH OF
CERTAIN GRAM-NEGATIVE BACTERIA

by JOHN R. McGILL and CAROLINE P. BENJAMIN

Biology Department,

Southwest Texas State University,

San Marcos 78666

ABSTRACT

The ability of bovine amniotic fluid to support the growth of 4 species of gram-negative
bacteria was examined and compared with the ability of control laboratory medium to do
so. The results indicate that growth is minimal in amniotic fluid and that this restriction is
due to the presence of unidentified inhibitory substance(s) in the amniotic fluid rather than
to the absence of essential nutrients in it. Furthermore, the antibacterial substance(s) remain
stable when stored over long periods of time and when exposed to extreme temperatures.

INTRODUCTION

Amniotic fluid, the watery medium which surrounds the embryo and fetus,
has been shown to be of significant value to the developing unborn (Fairweather
and Eskes, 1973; Moore, 1973; Natelson, et al, 1974). Its protective role for the
embryo or fetus is well documented (Balinsky, 1975). It is a dynamic fluid with
large volumes moving in both directions between the fetal and maternal circula¬
tions, mainly via the placental membrane (Barnes and Seeds, 1972).

Amniotic fluid samples may be collected relatively easily from the amniotic
cavity by transabdominal amniocentesis. This common clinical procedure has
not been accompanied by an increased incidence of amnionitis (Galask and Snyder,
1968). Thus, the antibacterial properties of amniotic fluid have become the sub¬
ject of many scientific investigations.

Walsh, et al, (1965) and Sarkany and Gaylarde (1968) have suggested that
amniotic fluid will support bacte rial growth as well as will control laboratory
medium. Kitzmiller, et al, (1973) have claimed that amniotic fluid, in itself, is
nutritionally deficient and will not support bacterial growth unless it is enriched
with added nutrients. Galask, et al, (1974) and Bergman, et al, (1972) maintain
that amniotic fluid is nutritionally sufficient for bacterial growth, but will not
support bacterial growth due to the presence of antibacterial factors found in
the fluid. These include such potential antibacterial substances as lysozyme, trans¬
ferrin, immunoglobulins, and the beta-lysin, and the hormone progesterone.
Received for publication: January 6, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

60

THE TEXAS JOURNAL OF SCIENCE

In view of the many conflicting reports in the literature concerning the anti¬
bacterial activity of amniotic fluid, this investigation was undertaken to deter¬
mine the nature of the inhibitory effects of bovine amniotic fluid on the growth
of four species of gram-negative bacteria.

MATERIALS AND METHODS
Amniotic Fluid

Amniotic fluid was collected aseptically from pregnant cattle. The fluid was
collected into sterile plastic containers, and transferred into sterile Erlenmeyer
flasks and immediately refrigerated at 5 C.

A 1-ml aliquot from each amniotic fluid sample collected was transferred into
each of 4 culture tubes containing nutrient media to confirm the sterility of the
amniotic fluid samples.

Organisms

The test organisms, Pseudomonas fluorescens, Proteus vulgaris, Escherichia
coli, and Serratia marcescens, were obtained from the microbiology stock culture
collection of Southwest Texas State University. These organisms were subcultured
and maintained on tryptic soy agar slants. From these slants, the organisms were
cultured in tryptic soy broth.

Media

Dehydrated Difco-Bacto tryptic soy broth and Difco-Bacto thioglycollate
medium, without indicator, were rehydrated according to the manufacturer’s
instructions.

Two % (w/v) Difco-Bacto agar was added to the tryptic soy broth for making
pour plates. Following sterilization, the media was placed in a 43-45 C water
bath until used (within 2-3 hrs). Pour plates were incubated 24 hrs at 37 C.

Physiological saline (0.85% NaCl) and phosphate buffer (0.1 M; pH 8.0), used
to dilute amniotic fluid and other media, were sterilized by filtration through
0.45 micron millipore filters. These sterile solutions were refrigerated at 5 C and
were used within 1 week.

Experimental Procedures

To determine the inhibitory capacity of amniotic fluid, growth studies were
conducted on 4 species of bacteria by both direct (standard plate count) and in¬
direct (photometric) methods of measurement.

Dilutions of amniotic fluid. Dilutions of amniotic fluid were made to deter¬
mine whether the inhibitory properties of amniotic fluid could be diluted out
before the loss of nutritional adequacy. Serial dilutions were made in 1-ml incre¬
ments from Oto 10 ml of amniotic fluid/culture tube. Phosphate buffer or physio¬
logical saline was pipetted aseptically into each of the culture tubes in the series
to a final volume of 10 ml/tube.

GROWTH OF GRAM-NEGATIVE BACTERIA

61

Enrichment of amnio tic fluid. Dilutions of amniotic fluid were made and en¬
riched with tryptic soy broth to determine whether the amniotic fluid was nutri¬
tionally deficient. Serial dilutions were made in 1 -ml increments form 0 to 10 ml
of amniotic fluid/tube. Tryptic soy broth was pipetted aseptically into each of
the series of culture tubes to a final volume of 10 ml.

Effect of the inocula. To minimize the lag phase, some species of bacteria
used for inoculation were cultured in amniotic fluid instead of tryptic soy broth.
These organisms were inoculated into tubes containing amniotic fluid or dilutions
thereof. The E. coli, Ps. fluorescens, and P. vulgaris used in these experiments
were cultured in 100%, 50% and 10% solutions of amniotic fluid. Sterile physio¬
logical saline was used to dilute all tubes to a final volume of 10 ml.

Effect of prolonged incubation time. Dilutions of amniotic fluid were made
using sterile physiological saline and these dilutions were inoculated and cultured
in the same manner as described previously, with the exception that the incubation
period was extended over a time period of 72 hrs. Growth of the bacteria was
measured photometrically and by the standard plate count method.

Heat resistance of amniotic fluid. The inhibitory activity of amniotic fluid
was measured after the fluid was heated either to 100 C for 45 min or to 121 C
for 15 min. These results were compared with the amniotic fluid samples that
had been refrigerated at either 5 C or -20 C. E. coli was grown in 100%, 50%,
and 10% solutions of each sample of amniotic fluid. Sterile physiological saline
was used as the diluent in order to obtain a final volume of 10 ml/tube. Each tube
was inoculated with 0.3 ml of E. coli. Bacterial growth was determined photo¬
metrically.

Effect of fetal gestation age on amniotic fluid. An attempt was made to corre¬
late the inhibitory activity of amniotic fluid wfth the gestation age. Comparisons
were made of the antibacterial activities of 3 different aged samples of amniotic
fluid. All tubes were inoculated with 0.3 ml of E. coli from a young broth culture.
Bacterial growth was determined photometrically.

Long-Term stability of the amniotic fluid inhibitor! s). Several samples of
amniotic fluid were stored at 5 C for a period of 15 weeks to determine the sta¬
bility of the bacterial inhibitory factor(s). Prior to storage, growth curves of
P. vulgaris were made by inoculating the amniotic fluid with 0.3 ml of a young
broth culture .

RESULTS

Dilutions of Amniotic Fluid

Amniotic fluid dilutions were made and were inoculated with the 4 species of
bacteria used in this study. The results of these experiments are shown in Table 1 .
E. coli showed maximum log growth in the 50-60% dilutions of amniotic fluid
while P. vulgaris and S. marcesens demonstrated maximum log growth in the 80-
90% dilutions. Ps. fluorescens grew maximally in the 40-50% dilutions of amniotic

62

THE TEXAS JOURNAL OF SCIENCE

fluid. E. coli diluted with physiological saline demonstrated maximum growth in
the 80% dilution of amniotic fluid.

TABLE 1

The effect of the dilution of amniotic fluid with phosphate buffer (PB) and
physiological saline (S) on bacterial growth. _ _

_ NET KLETT UNIT CHANGE PER 24 HRS

% of Amniotic Fluid

0

10

20

30

40

50

60

70

80

90

100

E. coli (PB)

5

10

21

32

48

67

71

65

62

67

60

Ps. fluorescens (PB)

9

27

33

42

54

47

38

38

30

32

37

P. vulgaris (PB)

14

11

11

14

12

20

29

33

39

35

10

S. marcescens (PB)

5

16

25

35

38

51

57

58

61

63

56

E. coli (S)

4

17

30

37

53

54

50

50

75

45

50

Growth rates of the organisms cultured in 100% amniotic fluid were compared
to the tryptic soy broth controls (Fig. 1).

Figure 1. Growth curves of (A) P. fluorescens, (B) P. vulgaris, (C) S. marcescens, and
(D) E. coli in amniotic fluid and in control tryptic soy broth. -A- amniotic
fluid; -a- tryptic soy broth.

&LETT UNITS KLETT UNITS

GROWTH OF GRAM-NEGATIVE BACTERIA

63

Viable counts of the bacteria also were made on several of the dilutions of
amniotic fluid which were inoculated with E. coli to confirm the results of the
indirect method. These results are seen in Table 2.

TABLE 2

The viable count of E. coli grown in amniotic fluid diluted with saline.

% of Amniotic Fluid

12 hrs

No. of Organisms

24 hrs

10

8

10

50

23

19

80

92

78

90

49

54

100

23

18

Enrichment of Amniotic Fluid

The results of the amniotic fluid samples enriched with tryptic soy broth are
found in Fig. 2. These results demonstrated that amniotic fluid retained its inhibitory

HOURS HOURS

HOURS H 0 U RS

Figure 2. Growth curves of (A) P. fluoresccns, (B) P. vulgaris, (C) S. marcencens , and
(D) E. coli in amniotic fluid, control tryptic soy broth, and in amniotic fluid
enriched with tryptic soy broth. -A- amniotic fluid; -a- tryptic soy broth;
— amniotic fluid plus 10% tryptic soy broth; amniotic fluid plus 20%
tryptic soy broth.

64

THE TEXAS JOURNAL OF SCIENCE

capacity when it was supplemented with varying amounts of tryptic soy broth.
If amniotic fluid were nutritionally deficient and contained no antibacterial
factors, the addition of tryptic soy broth should have resulted in better growth.
The inhibitors present in amniotic fluid could be diluted out when the samples
contained over 50% tryptic soy broth (Table 3). In some cultures containing
mainly tryptic soy broth, enhancement of bacterial growth was observed. In these
cultures the amount of inhibitor present was negligible.

TABLE 3

The effect of the enrichment of amniotic fluid with tryptic soy broth on bacterial growth.

NET KLETT UNIT CHANGE PER 24 HRS
% of Amniotic Fluid

0

10

20

30

40

50

E. coli

111

101

103

95

99

98

Ps. fluorescens

244

194

187

174

160

143

P. vulgaris

157

164

162

154

139

124

S. marcescens

149

166

181

164

160

148

Effect of the Inocula

Culturing bacteria to be used for inoculating purposes in amniotic fluid did
not prove successful. The inhibitors present in the amniotic fluid prevented suffi¬
cient growth of the organisms. With every species of bacteria studied, the amniotic
fluid-grown inocula exhibited less growth than the inocula grown in tryptic soy
broth (Table 4).

TABLE 4

The effect of inoculating amniotic fluid samples with amniotic fluid-grown cells.

NET KLETT UNIT CHANGE PER 24 HRS

Organism

%of

Amniotic Fluid

Amniotic Fluid-
Grown Cells

Control

Cells

E. coli

100

60

83

E. coli

50

38

50

E. coli

10

8

9

Ps. fluorescens

100

20

64

Ps. fluorescens

50

2

28

P. vulgaris

100

27

32

P. vulgaris

50

17

25

P. vulgaris

10

9

16

Effect of Prolonged Incubation Time

Extending the incubation period of the organisms studied demonstrated that
amniotic fluid and dilutions thereof supported minimal bacterial growth, and
within 24 hrs peak numbers of organisms were obtained. The results in Fig. 3 are
those of the E. coli strain studied.

GROWTH OF GRAM-NEGATIVE BACTERIA

65

Figure 3. The effect of prolonged incubation time on the growth of E. coli in amniotic
fluid and dilutions of amniotic fluid. -A- 100% amniotic fluid; 50% am¬
niotic fluid; 10% amniotic fluid; —A— tryptic soy broth control.

Heat Resistance of Amniotic Fluid

Fig. 4 shows the growth curves of E. coli grown in 100% amniotic fluid samples
that were exposed to different temperatures. No differences are seen in these
growth curves, indicating that the antibacterial factor(s) associated with amniotic
fluid are quite stable to temperature extremes. Other experiments in which di¬
lutions of amniotic fluid were used produced similar results. In each case examined
the inhibitory factor(s) remained stable.

Effect of Fetal Gestation Age on Amniotic Fluid

The gestation age becomes an important factor when the antibacterial activity
of amniotic fluid is considered. Fig. 5 shows that as the fetal age at which the
amniotic fluid was collected increases, the amount of bacterial growth decreases.

Long-Term Stability of the Amniotic Fluid Inhibitors )

The effect of the storage for extended periods of time on the amniotic fluid
inhibitor(s) is seen in Fig. 6. These results demonstrate that the inhibitor(s) are
quite stable when refrigerated at 5 C for 15 weeks.

THE TEXAS JOURNAL OF SCIENCE

4 8 12 16 20 24

HOURS

Figure 4. Growth of E. coli in amnio tic fluid exposed to temperature variations. 5 C;

-a- 100C; -o- 121 C; -A- -20 C.

CO

H

Z

Z3

H

I-

(U

_i

HOURS

Figure 5 . The effect of fetal gestation age on the growth of E. coli in amniotic fluid. — A-
early embryo; mid-gestation fetus; — ■— term fetus.

GROWTH OF GRAM-NEGATIVE BACTERIA

67

H

LU

HOURS

Figure 6. The long-term stability of the amniotic fluid factors in terms of their effects
on the growth of P. vulgaris. - m amniotic fluid stored for 24 hrs; -A— am¬
niotic fluid stored for 15 weeks.

DISCUSSION AND SUMMARY

This study has shown that bovine amniotic fluid will not support the growth
of bacterial species as well as control laboratory medium will and that it will only
support poorly any growth at all. Dilutions made of amniotic fluid indicated that
the inhibitor substance(s) could be diluted out before the loss of nutritional ade¬
quacy occurred. Moreover, the addition of nutrients to the amniotic fluid was
not beneficial to supporting growth of the bacteria, presumably because of potent
inhibitor(s) present in high concentrations of amniotic fluid. Two aspects of the
stability of the antibacterial factor(s) were investigated and revealed that the in¬
hibitory substance(s) remained stable when exposed to extremes of temperature
and when stored over long periods of time.

Much of the controversy in the literature regarding the antibacterial activity
of amniotic fluid cannot be resolved easily (Gusjdon, 1962; Walsh, etal., 1965;
Sarkany and Gaylarde, 1968; Galask and Snyder, 1968, 1970; Bergman, et al,
1972; Cherry, et al., 1973; Kitzmiller, et al., 1973; Galask, et al., 1974; Larsen,
et al, 1974a, 1974b); however, the data obtained in the course of this investigation
should lend support to certain of these findings. Furthermore, the use of bovine
amniotic fluid in this study circumvented some of the problems encountered by
other workers who used human amniotic fluid; namely, small aliquots of amniotic
fluid, contamination of the fluid with blood, and the necessity of filter sterilization.

68

THE TEXAS JOURNAL OF SCIENCE

Throughout the course of this study it was demonstrated that amniotic fluid
did not support significant growth in any of the bacterial species used because of
the antibacterial activity of the fluid. These results do not agree with those of
other investigators who claim either that amniotic fluid will support bacterial
growth or that it will not support bacterial growth due to nutritional deficiences.

ACKNOWLEDGEMENTS

The authors wish to acknowledge gratefully Mr. Edward Villegas, USDA and
the Hughson Meat Company of San Marcos, Texas, for enabling us to procure
the bovine amniotic fluid.

LITERATURE CITED

Adamsons, Karlis, 1968 -Diagnosis and Treatment of Fetal Disorders. Springer-VerlagNew York,
Inc., New York, N.Y.

Andrews, B. F., 1970-Amniotic fluid studies to determine fetal maturity. Pediat. Gin. N.
Amer., 17:49.

Balinsky, B. I., 1975-Arc Introduction to Embryology. 4th Ed., W. B. Saunders Co., Phila¬
delphia, Pa.

- — - , and A. Elmore Seeds, 1912-In Charles C. Thomas (Pub.), The Water Metabolism

of the Fetus. Springfield, Ill.

Behrman, R. E., 1967 -Placental growth and formation of amniotic fluid. Nature, 214:678.

Bergman, Nira, Bruno Bercovici, and Theodore Sacks, 1972- Antibacterial activity of human
amniotic fluid. Am. J. Obstet. Gynecol., 114:520.

Bonsnes, R. W., 1966-Composition of amniotic fluid. Gin. Obstet. Gynecol, 9:440.

Cattaneq, P., 1949 -Ricerche Sperimentol. Gin. Obstet. e Ginecol, 51:60.

Cherry, S. H., M. Filler, and H. Harvie, 1973-Lysozyme content of amniotic fluid. Am. J.
Obstet. Gynecol., 116:639.

Crosignani, P. G., and G. Pardi, 197 1- Fetal Evaluation During Pregnancy and Labor. Academic
Press, Inc., New York, N.Y.

Dancis, J., J. Lind, and P. Vara, 1960-Transfer of proteins across the human placenta. In
C. A. Villee (Ed.), The Placenta and Fetal Membranes. Williams and Wilkins, Baltimore, Md.

DIFCO Manual of Dehydrated Culture Media and Reagents for Microbiological and Clinical
Laboratory Procedures, 1967 -9th Ed., Difco Laboratories, Inc., Detroit, Mich.

Dorfman, A., 1912-Antenatal Diagnosis. The Univ. of Chicago Press, Chicago, Ill.

Emery, Alan E. H., 1912-Antenatal Diagnosis of Genetic Disease. Williams and Wilkins Co.,
Baltimore, Md.

Fairweather, D. V. I., andT. K. A.B. Eskes, 197 3- Amniotic Fluid. Excerpta Medica, Amsterdam.

Ferreira, Antonio J., 1969 -In Charles C. Thomas (Pub.), Prenatal Environment. Springfield, Ill.

GROWTH OF GRAM -NEGATIVE BACTERIA

69

Florman, A, L., and D. Tuebner, 1969-Enhancement of bacterial growth in amniotic fluid
by meconium./. Ped., 74:111.

Fort, Arthur T., and A. Roberts, 1970— Amniocentesis: its diagnostic and research applica¬
tions./. Term. Med, Assoc., 63:913.

Fuchs, Fritz, and Arnold Klopper, 191 1 -Endocrinology of Pregnancy. Harper and Row
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Galask , Rudolph P., Bryan Larsen, and Irvin S. Snyder, 1974-Amniotic fluid-induced sur¬
face ultramicrocytopathology of Escherichia coll Am. J. Obstet. Gynecol, 119:492.

— — — — , and Irvin S. Snyder, 1968 -Bacterial inhibition by amniotic fluid. Am.J. Obstet.
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70

THE TEXAS JOURNAL OF SCIENCE

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DENTITIONAL PHENOMENA AND TOOTH REPLACEMENT IN THE
SCABBARD FISH TRICHIURUS LEPTURUS LINNAEUS (PICES :
TRICHIURIDAE)

by EDWARD C. MORGAN

Department of Biology,

Temple Jr. College,

Temple 76501

ABSTRACT

Dentitional phenomena and tooth replacement were analyzed in a series of Scabbard fish,
Trichiurus lepturus, from the Gulf Coast of Texas. The dentition of this species is adapted
for impaling and holding prey. Teeth in the premaxilla and dentary may be simple, compressed
blades or elongate, recurved, barbed fangs. Replacement on the premaxilla involves 2 units,
a medial and a lateral unit. Replacement in the medial unit involves primary and accessory
alveoli at 3 tooth positions, which are occupied by fangs. Replacement teeth form in accessory
alveoli and migrate to the primary alveoli where they become ankylosed. Replacement in
the lateral unit of the premaxilla is typical of most predaceous polyphyodont vertebrates.
The dentary has 2 replacement units, an anterior unit of 2 alveoli which are occupied alter¬
nately by fangs, and a posterior unit in which replacement operates as it does in the lateral
premaxillary unit.

INTRODUCTION

Our knowledge of tooth replacement in lower vertebrates is incomplete. In
the works of Edmund (1960, 1969), the general pattern of tooth replacement
was described (primarily in reptiles) as typically polyphyodont with rear to front
waves of replacement throughout life.

In this paper, I analyze the dentitional phenomena in Trichiurus lepturus, a
member of the predaceous fish family Trichiuridae.

Trichiurus lepturus possess a very specialized dentition characterized by fangs
in the anterior portions of the premaxillary and dentary bones.

The trichiurids were reviewed taxonomically by Tucker (1956), who used
only a few characters of the dentition in his classification. Tucker proposed a
scheme which would include the Trichiuridae and Gempylidae as evolutionarily
related scombroid fishes, with the Trichiuridae having been derived from the
Gempylidae. Starks (1911) discussed the osteology of the family and included
comments on the shape of the premaxillary and dentary fangs, on the occurrence
of alveoli, and on the development of replacement teeth within the jaw bones.

Received for publication: March 10, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

72

THE TEXAS JOURNAL OF SCIENCE

MATERIALS AND METHODS

Fourteen specimens, 56 dentigerous bones, of T. lep turns were examined.
Specimens were obtained from a fish kill in Galveston County, Texas, and are
retained in my private collection.

The dentigerous bones were dissected and dried. Because of the decomposition,
no sex, weight, length, or age data were taken.

Data taken were: length of the dentigerous bones, extent of regional differ¬
entiation of the teeth, number of primary and accessory alveoli, number of tooth
positions, extent of embryogenesis, reabsorption and ankylosis of teeth, sequence
of tooth replacement, and number of replacement waves.

Measurements were made to the nearest millimeter. Replacement teeth, and
embedded portions of functional teeth were observed with a dissecting micro¬
scope with a glass insert in the base. A strong light was directed through the base
upon which the bones were positioned. Due to the extremely thin bone of the
jaws, even the smallest replacement teeth could be observed by this method.

Terminology for different surfaces of the teeth is that of Peyer (1968). Use
of the term fang is a retention of that proposed by Tucker (1956). The term
accessory alveolus was suggested by Edmund (1960). My interpretation of cranial
osteology follows Starks’ (1911).

TOOTH BEARING BONES AND REGIONAL DIFFERENTIATION
OF TEETH

Premaxilla

The premaxilla contains a lateral dentitional unit of 29-36 tooth positions,
and a medial dentitional unit of 3 positions which are occupied by fangs. Teeth
of the medial unit are ankylosed into alveoli and those of the lateral unit are
embedded in a longitudinal dental groove. Fangs are elongate, barbed teeth
which curve first caudad, then recurve anteriad toward the tip (Fig. 1). Cutting
surfaces are present on the entire labial border and On both the labial and lingual
border of the barbed tip. The lingual border, excluding the barbed tip, is rounded.

Teeth of the lateral unit posterior to Position 1 0 or 1 1 are laterally compressed
with cutting surfaces on the mesial and distal borders. Anterior to this position,
the size of the teeth is abruptly reduced and the teeth are more attenuated with¬
out cutting edges. The long axis of each tooth is perpendicular to the long axis
of the premaxilla up to Position 19 or 20, then it is directed somewhat caudad.
Vertical height of these teeth increases to about halfway on the bone, then de¬
creases to the end of the series.

Dentary

The dentary contains a posterior unit of 18-24 positions in a longitudinal
dental groove, and an anterior unit of 2 alveoli, one of which is occupied by a
fang. One of the alveoli is anterior medial with respect to the other. There is a

THE SCABBARD FISH

73

Figure 1. Anterior medial view of the right premaxilla of ECM-D-9, Trichiurus lepturus.

short diasterna of about 5 tooth positions separating the units. Fangs of the den¬
tary are shorter and less curved than those of the premaxilla.

Teeth of the posterior unit, in the anterior and posterior portions, are similar,
although less expanded mesio-distally, to those of the lateral premaxillary unit.
In the middle portions of this unit, the teeth develop barbed tips and cutting sur¬
faces similar to those of the anterior unit. Numbers of barbed dentary teeth (0-3)
seem to increase during ontogeny. No specimen with a dentary length of less
than 35 mm possessed a functional barbed tooth in the posterior dentary series.
Two specimens possessed 3 barbed teeth (ECM-D-8,44mm andECM-D-6,49 mm).
A specimen with a dentary length of 50 mm (ECM-D-4) has one barbed dentary
tooth.

TOOTH DEVELOPMENT, ANKYLOSIS AND REABSORPTION

Tooth development begins at the tip and proceeds to the base. Ankylosis
appears to be by a cementum-like substance rather than by fibrous connective
tissue. Tooth development occurs within the dentigerous bones.

Reabsorption begins at the base and proceeds toward the tip, eventually attack¬
ing the ankylosing substance.

74

THE TEXAS JOURNAL OF SCIENCE

REPLACEMENT
Medial Premaxillary Unit

Replacement in this unit involves a primary and an accessory alveolus at 3
tooth positions. Functional teeth are ankylosed into primary alveoli which are
nearly perpendicular to the long axis of the premaxilla (Fig. 1). Replacement
teeth form in an accessory alveolus which opens dorsal and posterior to the primary
alveolus on the medial aspect of the bone. Replacement teeth completely form
in the accessory alveolus. The fangs make an anterior-ventral migration into the
primary alveolus after reabsorption of the bone separating the 2 alveoli. Reforma¬
tion of the 2 alveoli occurs after the migration. Re absorption of the functional
tooth occurs simultaneously with the development of a replacement at each
position. The usual sequence of replacement (20 of 28 premaxilla) appears to be
in an alternating pattern. In the 8 premaxilla which do not appear to have con¬
tinuous replacement, the condition is probably one of a tooth in the middle
alveolus being followed by a replacement in alveolus 1 or 3. This is a typical re¬
placement sequence which would be expected in 1/3 of the specimens. Replace¬
ment at Position 1 may proceed faster or tooth retention may be longer at that
position. When right and left premaxilla of 14 specimens were considered, only
3 primary alveoli were vacant at Position 1. In 12 dentigerous bones, the primary
alveoli at Position 2 were vacant, and in 9 the primary alveoli at Position 3 were
vacant (Table 1). No synchrony of replacement, between right and left premaxilla,
was evident in 13 of 14 specimens.

TABLE 1

A summary of the replacement phenomena in the medial premaxillary unit of
Trichiurus lep turns.

Position 1

Position 2

Position 3

Tooth

Right

Left

Right

Left

Right

Left

Condition

Alveolus

Alveolus

Alveolus

Alveolus

Alveolus

Alveolus

Fully ankylosed

7

10

8

3

4

9

Unankylosed

1

0

0

1

2

0

Partially ankylosed

2

2

2

2

0

2

Reabsorption evident

2

1

0

0

0

2

Socket vacant

2

1

4

8

8

1

Broken ankylosed

0

2

1

2

1

1

Dentary and Lateral Premaxillary Units

Replacement teeth of the anterior dentary unit form deep within the alveoli.
Formation of a replacement tooth begins before the loss of the old functional
tooth at that position. In the right dentary of ECM-D-5 both alveoli have at least
partially functional teeth. Usually, loss of an old functional tooth occurs prior to
ankylosis of a replacement in the other alveolus. Each alveolus is occupied alter¬
nately without any synchrony between jaws.

THE SCABBARD FISH

75

The posterior dentary and lateral premaxillary units exhibit posterior to an¬
terior replacement waves. About half the tooth positions in any dentigerous
bone possess at least partially functional teeth (Table 2). Replacement waves in
the dentary operate for the full length of the odd series as well as for the full
length of the even series of teeth. Replacement in the premaxilla operates in 2
waves in the odd and 2 waves in the even series of teeth. The usual line of demar¬
cation between the 2 waves is at a level lateral to the posterior edge of the third
premaxillary fang. One specimen (ECM-D-10) exhibits a reversal in the direction
of replacement at about tooth Position 16. Usually, about 4 generations of teeth
are evident in any particular dentary or premaxilla, and no more than 2 teeth
occur at any one tooth position.

TABLE 2

A summary of tooth positions and functional teeth in the lateral premaxillary unit
and the posterior dentary unit of Trichiurus lep turns.

Functional Teeth

Tooth Positions

Bone

Range

Mean

Range

Mean

Premaxilla

13-18

15.2

29-36

31.1

Dentary

8-11

9.9

18-24

22.2

I have chosen the right dentary of ECM-D-9.to use as an example here (Fig. 2).
Four generations of teeth, in rear to front waves are evident, when the exposed
height of each tooth is considered.

12 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22

Figure 2. Semischematic representation of the replacement sequence in the posterior unit
of the right dentary of ECM-D-9, Trichiurus lep turns. Tooth generation 1 is
represented by diagonal lines, 2 by solid symbols, 3 by stippling, and 4 by open
symbols. The horizontal line between teeth indicates the jaw line, and numbers
indicate tooth positions.

DISCUSSION

Development of the cutting surfaces on the labial border of the fangs and the
presence of barbed tips are probably adaptations toward penetration and holding

76

THE TEXAS JOURNAL OF SCIENCE

rather than shredding of tissue, since it would seem likely that a tooth adapted
for shredding would possess a greater cutting surface on the lingual aspect. The
greater mesio-distal expansion of the lateral premaxillary dentition may retard
penetration. These teeth may function in holding the prey in a fixed position
while the upper jaw- forces the prey down onto the barbed teeth of the posterior
dentary series. Evidence that the number of barbed dentary teeth increases during
ontogeny may reflect a change in feeding habits with age. The reduced size of
the teeth in the anterior end of the lateral premaxillary unit and their position,
which is well removed from the occlusal surfaces of the premaxillary fangs which
they flank, may indicate that these teeth do not function in the feeding activities
of this fish.

The development of each replacement tooth occurs in a manner well docu¬
mented (Bogert, 1943; Edmund, 1960, 1969;Peyer, 1968). Development of teeth
within the jaw bones is an adaptation for the protection of growing teeth from
being dislodged by a struggling prey. Similar development of teeth occurs in
predaceous fish such as Lepidotes (Peyer, 1968) and most of the characoid genera
(Roberts, 1967). Re absorption of teeth occurs in a manner similar to other poly-
phyodont vertebrates (Edmund, 1960, 1969; Peyer, 1968).

The occurrence of accessory alveoli well away from the occlusal surfaces of
the functional premaxillary teeth, seems to be an adaptation for predation. Ac¬
cessory alveoli have been reported by Morgan (1973) in the sibynophiine genus
Scaphiodontophis; Edmund (1960) in ornithiscian dinosaurs; and by Roberts
(1967) in the characoid genera Hydro cynus and Alestes.

Longer tooth retention or more rapid tooth replacement at Position 1 in the
premaxilla would be a selective advantage in a predaceous animal since it is this
tooth position which contacts the prey first. Asynchrony between right and left
premaxillary units probably maintains at least 1 functional tooth at 3 levels in
the series when both premaxilla are considered as a functional unit. If longer
tooth retention is a fact, then there will often be 2 teeth at alveolus 1 , due to an
overlap in the replacement sequence between right and left premaxilla.

The replacement sequence on the dentary and lateral premaxillary units is
typical of polyphyodont vertebrates (Bogert, 1943; Edmund, 1960, 1969; Peyer,
1968). The low number of replacement teeth suggests that replacement may be
slow. The holostean Lepisosteus, with intense replacement, exhibits a high num¬
ber of replacement teeth (Peyer, 1968). The condition in T. lepturus may be feasible
because of the well-protected position of the replacement teeth, the deep implan¬
tation of the functional teeth, and the cementum type of ankylosing substance.

CONCLUSIONS

The dentition of T. lepturus is adapted for impaling and holding prey. Teeth
in the premaxilla and dentary are either laterally compressed blades or elongate,
recurved, barbed fangs. Replacement on the premaxilla involves primary and

THE SCABBARD FISH

77

accessory alveoli at 3 tooth positions, which are occupied by fangs. Replacement
teeth form in accessory alveoli and migrate to the primary alveoli where they be¬
come ankylosed. Replacement in the lateral unit of the premaxilla is typical of
most predaceous polyphyodont vertebrates. The dentary has 2 replacement units,
an anterior unit of 2 alveoli which are occupied alternately by fangs, and a posterior
unit in which replacement operates as it does in the lateral premaxillary unit.

ACKNOWLEDGEMENTS

I am grateful to Pat H. Simpson and my wife , Marie , for their helpful comments
and criticism of the manuscript.

LITERATURE CITED

Bogert, C. M., 1943-Dentitional phenomena in cobras and other elapids, with notes on the
adaptive modifications of fangs. Bull. Amer. Mus. Nat . Hist., 81(3) :285 .

Edmund, A. G., 1960-Tooth replacement phenomena in lower vertebrates. Royal Ontario
Mus., Life Set Div., Contr 52:1.

— - 1969-Dentition. In C. G. Cans and A. d’A. Be Hairs (Eds.), Biology of the Reptilia.

Academic Press, London.

Morgan, E.C., 1973-Snakes of the subfamily Sibynophiinae. Ph.D. Thesis, Univ. Southwestern
Louisiana.

Peyer, B., 1968 -Comparative Odontology. Univ. Chicago Press, Chicago, Ill.

Roberts, T. R., 1967-Tooth formation and replacement in characoid fishes .Stanford Ichthyol
Bull, 8:231.

Starks, E. C, 1911 -Osteology of certain scombroid fishers. The Osteological Characters of
the Scombroid Fishes of the Families Gempylidae, Lepidopidae and Trichiuridae. Stanford
Univ. Pubh No. 5 , pp. 17-26.

Tucker, D. W., 1956-Studies on the trichiuroid fishes - 3: A preliminary revision of the
family Trichiuridae. Bull. Brit. Mus. Nat. Hist. Zool, 4:73.

' : i V v •; ‘

.

•

.

.

.

!

.

EVIDENCE FOR THE SPECIES STATUS OF BAIRD’S RATSNAKE

by R. EARL OLSON1

Department of Biology,

University of Colorado,

Boulder 80302

INTRODUCTION

When allopatric populations of similar taxa have ranges which narrowly over¬
lap geographically, the systematist must decide whether he is dealing with sub¬
species of the same species or with 2 species. Mayr, etal, (1953) in review of the
characteristics of allopatric populations with narrow zones of overlap, showed
that in those cases where there is complete interbreeding over relatively broad
or very narrow zone of contact, one is dealing with subspecies of the same species,
but where only occasional hybrids or no interbreeding occur at the zone of con¬
tact, one is dealing with different species.

In 1880, Yarrow (in Cope) described Coluber bairdi from Ft. Davis, Jeff Davis
County, Texas. Dowling (1951), citing only 2 specimens of bairdi , but alluding
to others, referred it to a subspecies of Elaphe obsoleta. He argued that although
bairdi has more subcaudals and dorsal blotches, those same features in obsoleta
increase in number towards the west; he envisioned bairdi as representing the
western extremes of clinal variation, stating that a few Kerr County specimens
represented an intergrade population on the basis of dorsal blotch number.

Wright and Wright (1957) treated Elaphe bairdi as a species, mentioning that
it could be separated from Elaphe obsoleta by its usually greater number of sub¬
caudals and dorsal blotches.

The present study is based on specimens of Elaphe bairdi from throughout its
known range, and of Elaphe obsoleta from nearby localities in central Texas.
Specimens have been examined from the following collections: Chicago Academy
of Sciences; Strecker Museum, Baylor University, Waco, Texas; Sul Ross State
University, Alpine, Texas; Texas Cooperative Wildlife Collection, Texas A&M
University, College Station; Texas Natural History Collection, University of Texas,
Austin; and the University of Michigan Museum of Zoology, Ann Arbor.

DIAGNOSTIC CHARACTERS

Several characters have been used to separate Elaphe bairdi from Elaphe obsoleta.
Not all of those are reliable, while others that have not been previously mentioned

Present address: Department of Biology, Cornell College, Mount Vernon, Iowa 52314.
Received for publication: September 8, 1975.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

80

THE TEXAS JOURNAL OF SCIENCE

are found to be distinctive, the most useful characters being ventral counts, sub-
caudal counts, and pattern.

Ventrals

In the specimens examined, ventral counts for both sexes of Elaphe hairdi
average distinctly higher than in adjacent populations of Elaphe ohsoleta. Eleven
male bairdi have 233-260 (X = 247.8) ventrals, whereas 11 male obsoleta have
220-232 ( X = 225 .6). The one example with 233 ventrals (other male bairdi have
no fewer than 242) is from southern Brewster County, Texas. Nine bairdi females
have 235-247 (X = 241.9) ventrals, whereas those counts in 9 obsoleta females
are 223-238 (X = 229.0).

Males of Elaphe bairdi have a greater mean number of ventrals than do females,
but in central Texas, ventral counts for female Elaphe obsoleta slightly exceed
those for males. Ventral counts of female obsoleta from Kansas (Smith, 1956)
and Illinois (Smith, 1961) also exceed those of males.

Subcaudals

Although some overlap is exhibited in subcaudal number, the counts average
distinctly higher in Elaphe bairdi than in Elaphe obsoleta. In bairdi , subcaudal
counts are 88-100 (X = 94.0) in 1 1 males, and 82-93 (X = 88.6) in 8 females.
The counts for obsoleta are 75-89 ( X = 8 1 .8) in 12 males, and 72-79 ( X = 74.0)
in 7 females.

Pattern

A comparison of color pattern presents difficulties because of the extreme
ontogenetic variation exhibited by Elaphe bairdi, and also because subadult and
adult specimens tend to fade in preservative, revealing blotching that is masked
in life. By contrast, central Texas Elaphe obsoleta change relatively little in pattern,
except that the ground color occasionally becomes suffused- with black, obscuring
or almost obliterating the blotches.

Dorsal and lateral blotches are apparent only in juvenile Elaphe bairdi, but
they remain visible throughout life in central Texas Elaphe obsoleta (Fig. 1). Dial
(1965) illustrated the juvenile pattern of bairdi. Dorsal body blotches are 44-61
(X = 50.5) in 8 bairdi, and 27-37 (X = 32.0) in 20 obsoleta. Dorsal and lateral
blotches in obsoleta are conspicuously longer than are those in bairdi. In obsoleta
the dorsal blotches are 4-6 scales long, the lateral blotches 4-5 scales long. In
bairdi, when blotches are present, they extend 2-2.5 scales, rarely 3, and lateral
blotches 1.5-2 scales. Dorsal blotches in obsoleta are typically broadly H-shaped,
usually transversely oblong in bairdi.

In Elaphe bairdi, the blotched dorsal pattern of the juvenile gives way to the
dorsolateral and lateral dark stripes of the young adult. The striped snake has a
gray ground color, each scale being orange anteriorly in slightly older individuals.
Later the stripes disappear, and the ground color becomes orange, deepening

BAIRD’S RATSNAKE

81

Figure 1 . The midbody patterns of the juvenile Elaphe bairdi from Real County, Texas (left),
and of an adult E. obsoleta from west-central Bexar County, Texas (right).

posteriorly to red-orange (Olson, 1967). The posterior 2/3 of each dorsal scale is
suffused with black peppering, so that larger snakes are grayish orange in appear¬
ance.

Freshly preserved specimens (1370-1480 mm) were orange cast with faint
striping; a 1571 mm male was completely without pattern in life (Olson, 1967),
though, after 7 years in preservative, very faint blotch outlines are discernible.
Assuming that the sexes are similar in pattern ontogeny, the juvenile pattern per¬
sists in animals about 700-800 mm long, the young adult pattern in the size range
between 880-1200 mm, and adult pattern appears in snakes more than 1250 mm.
In central Texas Elaphe obsoleta the pattern of dorsal and lateral blotches remains
conspicuous throughout life.

Adults of Elaphe obsoleta have a uniformly black head dorsally and laterally
with the exception of the lower parts of the supralabials (Fig. 2). In contrast,
the sides of the head are light in Elaphe bairdi, only the dorsal head plates being
dark gray or gray -brown (Fig. 3).

Figure 2. Lateral view of the head of an adult male Elaphe obsoleta (WMM 650) from
south of Kerrville, Kerr County, Texas.

82

THE TEXAS JOURNAL OF SCIENCE

Figure 3. Lateral view of the head of an adult male Elaphe bairdi { REO 687) from near
Alpine, Brewster County, Texas.

Other characters which have been utilized to separate the snakes taxonomically
are felt to be unreliable: genials, posterior scale row reduction, relative tail length,
and number of supralabials.

Genials

Wright and Wright (1957) mentioned the divided posterior genials in Elaphe
bairdi, a condition not seen in Elaphe obsoleta. The character is variable in the
series examined. A few specimens of bairdi have one or both posterior genials
divided; many have undivided scales. Briefly, the genial condition seems constant
in obsoleta, variable in bairdi.

Posterior Scale-Row Reduction

Reductions to 19 scale rows occurs in 90% of the Elaphe bairdi examined, but
only 40% of central Texas obsoleta, the others having 21.

Relative Tail Length

Elaphe bairdi has a proportionately longer tail than does Elaphe obsoleta. Male
bairdi have tail/total length values of 18.6-23.0% (X = 20.6%), female bairdi
18.3-21.8% (X = 19.8%), male obsoleta 16.0-19.5% (X = 17.9%), and female
obsoleta 16.0-19.2% ( X = 17.5%).

Supralabials

Supralabial counts have been used as a basis for distinguishing these snakes,
but are unreliable. Milstead, et al., (1950) suggested that bairdi might some day
be found to be a subspecies of Elaphe obsoleta on the basis of specimens of bairdi
with 8 supralabials.

BAIRD’S RATSNAKE

83

In central Texas, 92% of the Elaphe obsoleta examined have 8 supralabial
scales on each side. However, Elaphe bairdi may possess 8 or 9 supralabials, or a
combination thereof (Table 1).

TABLE 1

Frequency distribution of supralabial number in Elaphe bairdi.

Number of Supralabials1

16

6

4

17

2

1

18

2

3

1 Combined count for both sides of head.

DISCUSSION

Differences between Elaphe bairdi and Elaphe obsoleta are not diminished by
geographic proximity. Such would not be expected in an intergrade situation. I
am aware of 2 areas where the ranges of these forms overlap; south of Kerrville,
and at Ingram, in Kerr County, Texas. A third possibility exists along the eastern
edge of Medina County, but only a striped female Elaphe bairdi is extant from
there. (Specimens of Elaphe obsoleta are available from 5 mi east of Medina Lake
in Bexar County.)

The series from south of Kerrville includes 5 Elaphe bairdi (2 males, 3 females),
and 2 male Elaphe obsoleta. The female bairdihzve 235,242, and 243 ventrals,
86+ , 89, 90 subcaudals, and 21, 19, 19 scale rows posteriorly, while the males
have 248, 250 ventrals, 89+, 100 subcaudals, and 19 scale rows posteriorly. The
Elaphe obsoleta have 225, 229 ventrals, 89, 88 subcaudals, and 21 scale rows
posteriorly. These obsoleta are adults with prominent blotches (30, 33) and with
head uniformly dark above the lower portions of the supralabials. All bairdi, on
the other hand, show the typical body and head patterns previously described
for that form. Thus, no hybridization in apparent in that area.

From Ingram, 5 adult and subadult males are available. Of those, two are
typical of obsoleta, having 230, 231 ventrals, 83,83 subcaudals, 21,19 posterior
scale rows, and 34, 32 dark dorsal body blotches. Three show what is regarded
to be intermediacy. One (UMMZ 102843) has 225 ventrals and 84 subcaudals as
in obsoleta , but 44 blotches arranged in' a pattern resembling the condition in
juvenile bairdi The other 2 hybrids (UMMZ 102844, 102845) have patterns
similar to that of obsoleta, with 239, 235 ventrals, 91, 81 subcaudals, and
40, 42 dorsal blotches. All of these specimens have 19 posterior scale rows.

The question of whether one is dealing with a species or with subspecies in
this case can be viewed with regard to 3 features of the population: the presence
of an area of overlap of ranges, the nature of the population in the area(s) of
overlap, and the nature of populations adjacent to the areas of overlap.

84

THE TEXAS JOURNAL OF SCIENCE

Two known areas of overlap of ranges have been mentioned above. Were inter¬
gradation occuring between subspecies of the same species, it would be expected
that the entire population within the area of range overlap would reflect inter¬
mediacy in many or all of the characters. Such is not the case here. Specimens
from the same localities have all characters of either Elaphe obsoleta or Elaphe
bairdi or only some show intermediacy (at Ingram). This reflects a hybrid situation,
not intergradation.

The populations adjacent to the areas of range overlap differ from each other
100% in characters regarded to be reliable. They do not reflect intermediacy.
Again this reflects species, not subspecies status.

Thus, although some hybridization probably occurs at Ingram, it is unknown
elsewhere where the ranges meet, nor is there evidence of introgression. I feel
that Elaphe bairdi and Elaphe obsoleta are acting as good species, and should be
so regarded.

ACKNOWLEDGEMENTS

I am grateful to Drs. Hobart Smith and Philip W. Smith for reading the manu¬
script and offering very useful suggestions.

LITERATURE CITED

Cope, E. D., 1880-On the zoological position of Texas. Bull. U. S. Natl. Mus., 17:1.

Dial, B. E., 1 965 -Pattern and coloration in juveniles of two West TexasElaphe. Herpetologica,
21:75.

Dowling, H.G., 1951-A taxonomic study of the American representatives of the genusElaphe
Fitzinger. Dissertation, Univ. Michigan, Ann Arbor. 194 p.

Mayr, E.,E.G. Linsley and R. L.Usinger, 1953-Methods and Principles of Systematic Zoology.
McGraw-Hill, New York, N.Y., 328 p.

Milstead, W. W., J. Mecham, and H. McClintlock, 1950 -The amphibians and reptiles of the
Stockton Plateau, northern Terrell County, Texas. Tex. J. Sci., 2:543.

Olson, R. E., 1967— Peripheral range extensions and some new records of Texas amphibians
and reptiles. Tex. J. Sci, 19:99.

Smith, H. M., 1956-Handbook of Amphibians and Reptiles of Kansas. Univ. Kansas Publ.,
Mus. Nat. Hist., 356 p.

Smith, P. W., 1961— The amphibians and reptiles of Illinois .III. Nat. Hist . Surv. Bull. No. 28,
298 p.

Wright, A. H., and A. A. Wright, 1951 -Handbook of Snakes. Vol. 1. Comstock Publ. Assoc.,
Ithaca, N.Y., 1105 p.

ANTIGENIC ANALYSIS OF THE GENUS AEROMONAS

by VIJAYA B. RAO and B. G. FOSTER

Department of Biology,

College of Science,

Texas A&M University,

College Station 77843

ABSTRACT

Particulate and soluble heat stable antigens were prepared from autoclaved cultures of
the recognized species and subspecies of the genus Aeromonas grown on solid media. Anti¬
sera were prepared in New Zealand white rabbits. Cross reactions and cross adsorptions were
performed in attempts to show relationships between the members of the genus.

There was a relationship between A. liquefaciens, A. punctata, A. punctata caviae and
A. hydrophila when agglutinins were demonstrated in antisera prepared with whole cells and
a cell-free supernatant. In addition to species-specific antigens^the analysis showed an antigen
that was shared by A. liquefaciens, A. punctata , and A. punctata caviae . In addition, A. li-
quifaciens also shared another antigen with A. hydrophila and yet another with ,4. punctata.

INTRODUCTION

The genus Aeromonas has been known and studied for the past 80 years. Still,
its classification is not clear. Several investigators have used serological methods
in an attempt to classify these organisms. However, their studies did not include
all of the species and subspecies in this genus. Guthrie and Hichner (1943), when
working with strains of A. hydrophila, found them to constitute an antigenic
heterogeneous group. Miles and Miles (1951) also showed antigenic heterogeity
when testing 12 strains from different sources and antisera to 7 of them. Kjem
(1955) carried out cross agglutination experiments with 5 Aeromonas strains
using O (Somatic) antigens and H (flagella) antigen preparations. He found 2 of
these strains had 0 related antigens but were otherwise unrelated. Alodova and
Geizer (1968) worked with 14 strains of A. shigelloides. Cross O-agglutination
showed that most of the strains had a distinct O-antigen or O-antigen complex.
Three strains proved to be more related to each other than to the remaining strains
examined. In H-antigen cross agglutinations, they found 5 groups having specific
antigens for their group, one of which could be divided into subgroups. Ewing,
et al., (1961), using several strains of Aeromonas from the cultural collection of
CDC-RH, established 12 previsional O-antigen groups and 9 H-antigen groups
and differentiated a number of serotypes within each group. Among the inter¬
relationships between species shown by them include antigens shared by the fol¬
lowing 5 pairs: A. salmonicida and A. hydrophila; A. formicans and A. liquefaciens;

Received for publication:

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

86

THE TEXAS JOURNAL OF SCIENCE

A. formicam and A. shigelloides; A. hydro phila and A. formicans; and A. shigel-
loides and ,4. punctata.

This paper reports the antigenic relatedness between all available species and
subspecies of Aeromonas with the use of 2 heat stable somatic antigen preparations.

METHODS AND MATERIALS

Antigens

Representatives of available species and subspecies of the genus Aeromonas
were used in this study. These included: A. hydrophila ATCC 0971, A hydro-
phila formicans ATCC 13137, A. liquefaciens ATCC 14715, A. proteolytica
ATCC 15338, A. punctata ATCC 11163, A. punctata caviae ATCC 14486, A sal-
monicida # 61 Italy MS-320 West Fish Disease Lab, and A. shigelloides ATCC
14029.

Excepting ^4. salmonicida and A. proteolytica, all cultures for antigen prepara¬
tion were grown in Brain Heart Infusion (BHI) Broth (Difco) starter cultures
which were used to inoculate BHI agar in Roux bottles. Growth was for 24 hrs
at 37 C. A. proteolytica requires sea water for growth (Merkel, et al, 1964) and
consequently was grown under the above conditions except on 2% casitone made
with artificial sea water. A. salmonicida requires a growth temperature of 22 C.
After incubation, the cells were harvested in saline in a flask containing glass
beads and agitated on a reciprocal shaker for 1 hr. After separation, the cells were
washed in saline and killed by autoclaving at 15 lb psi and 121 C. The heat-killed
suspension was then centrifuged and the supernatant filtered. Both the superna¬
tant (soluble antigen) and suspended cells (6 x 106) (particulate antigen) were
refrigerated at 4 C.

Antiserum

Two types of antisera were prepared against the described strains. One type
was directed against the particulate antigen and the other against the soluble
supernatant antigens. New Zealand white rabbits of mixed sex and weighing 2-3
kg were pre-bled and tested for natural antibody. The antigens were mixed with
Freund’s (1957) incomplete adjuvant in 1:1 proportions and 2 ml of this material
given in each hip once a week for 2 weeks. On the third week, the animals were
given 2 ml of 1 :1 mixture of antigen and Freund’s complete adjuvant.

Serological Cross Reactions

A tube agglutination test, utilizing both homologous and heterologous sero¬
logical systems were used to assess the antigenic relatedness of the various species.
Two-fold dilutions of antiserum were prepared. The saline used for dilution con¬
tained 2% gelatin to minimize auto-agglutination. Incubation was at 56 C for 5-6
hrs (Kabit and Meyer, 1948) followed by overnight refrigeration.

GENUS AEROMONAS

87

Agglutinin Adsorption and Antigenic Analysis

Adsorption studies were conducted as dictated by results observed in the sero¬
logical cross reactions. The serum was diluted 1:10 andmixed with heavy suspen¬
sions of appropriate cells. This mixture was incubated in the 56 C water bath for
4 to 5 hrs without shaking. After overnight refrigeration, the mixture was centri¬
fuged and the clear supernatant removed. The adsorbed serum was tested for
homologous and heterologous agglutination using the above procedure.

RESULTS AND DISCUSSION
Relationship of Particulate Antigens

Agglutination tests were used to analyze the homologous as well as hetero¬
logous antigen-antibody systems. Cross agglutinations of the heat stable particu¬
late systems showed that most of the strains have distinct O-antigens or 0 -antigen
complexes. The O-antigenic relationship was only observed within 4 groups as
seen in Table 1. Low titer reciprocal agglutination reactions were observed in the
heterologous system of A. salmonicida and A. liquefaciens, but these were not
considered significant. Some organisms reacted only in one system but not in the
reciprocal (Table 1). The nature of the O-antigenic relationship between^, lique¬
faciens, A. hydrophila, A. punctata and A. punctata caviae is seen in Table 2.
According to these adsorption experiments, A. liquefaciens shares an antigenic
component with both A. punctata andA punctata caviae and another in common
with A. hydrophila, but lacked a distinct O-antigen. A. punctata was shown to
have in addition to the antigen shared with A. liquefaciens, a species specific
antigen. A. punctata caviae also had a distinct antigen in addition to one shared
with A. punctata and A. liquefaciens. A. hydrophila was also found to have a
distinct antigen in addition to the one shared with A. liquefaciens.

Relationship with Soluble Antigens

Homologous and heterologous relationships of antisera to soluble antigens
were analyzed by reacting antisera with the various particulate (whole heat-
killed cells) antigens. The results are presented in Table 3. It seems that soluble
antigenic complexes of A. liquefaciens, A. punctata and A. punctata caviae are
capable of stimulating antibodies to related antigens, thus share common antigens,
whereas A. hydrophila, A. hydrophila formicans, A. proteolytica, A. salmonicida
and A. shigelloides possess unique antigens since the latter organisms did not
stimulate cross reacting antibodies. The relatedness of A. liquefaciens, A. punc¬
tata and A. punctata caviae as determined by cross adsorption experiments is
seen in Table 4. Again, A. liquefaciens was found to share an antigen with A. punc¬
tata and A. punctata caviae. However, the adsorptions also indicate that A. lique¬
faciens does have a distinct antigen and an antigen shared with A. punctata in
addition to the antigen shared with A. punctata and A. punctata caviae. Ad¬
sorptions with A. punctata verified the presence of these antigens and also its

88

THE TEXAS JOURNAL OF SCIENCE

6

3

03

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type specific antigen. As before, studies with A. punctata caviae showed the
presence of antigen shared with A. liquefaciens and the species-specific antigen.

GENUS AEROMONAS

89

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Unlike in the particulate antigen adsorptions, there was no relationship demonstrated
between A. liquefaciens and A. hydrophila. In the 8 strains studied, each had a
distinct species-specific 0 -anti gen or O-antigenic complex and in addition, 4 had
interrelated O-antigens. They were: A. liquefaciens, A. punctata, A. punctata
caviae, and A. hydrophila . However, some difference existed between the 2 analyses.
In the particulate antigenic analysis, the relationship between^. hydrophila and
A. liquefaciens was observed, a relationship that was not observed in the soluble
antigen analysis.

In the particulate antigenic analysis, a specific antigen to A. liquefaciens and a
common antigen between A. liquefaciens and A. punctata were not demonstrated,
whereas in the soluble antigen analysis, they were observed.

Serological Cross Reactions: Antisera to Soluble Antigens

90

THE TEXAS JOURNAL OF SCIENCE

•5 R

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GENUS AEROMONAS

91

TABLE 4

Cross Adsorptions: Antisera to Soluble Antigen

Serum

Adsorbed With

Reacted With

A. liquefaciens

A. punctata

A. punctata
caviae

A. liquefaciens

1:160

1:80

1:40

A. punctata

1:20

0

0

A. punctata
caviae

1:40

1:20

0

A. punctata

1:40

1:160

1:20

A. liquefaciens

0

1:40

0

A. punctata
caviae

1:20

1:40

0

A. punctata
caviae

1:20

1:20

1:160

A. liquefaciens

0

0

1:40

A. punctata

0

0

1:40

The analysis recorded here should be regarded only as an indication of the re¬
lationship between the recognized species and subspecies. Inclusion of more strains
from each Aeromonas group would reveal a more complete antigenic relationship.

LITERATURE CITED

Aldova, E., and D. Geizer, 1968-A contribution to the serology of Aeromonas shigelloides.
Zentrobl. Bacteriol. Parasitenk. Infect. Krankh. Hyg., 207:41.

Ewing, W. H., R. Hugh, and J. G. Johnson, 1961— Studies on the Aeromonas group. U.S.
Dept, of Health, Education and Welfare, Communicable Disease Center, Atlanta, Ga.

Freund, J., 1957-Adjuvant action./. Allergy, 28:18.

Guthrie, R., and E. R. Hitchner, 1943— Studies on microorganisms classified as Proteus
hydrophilus. J. Bacteriol, 45:52.

Kabit,E., and M. Mayer, 1948 -Procedure for quantitative microdetermination of agglutinins.
Experimental Immuno chemistry . Charles C. Thomas, Springfield, Ill., p. 115.

Kjems, E., 1955-Studies on five bacterial stains of the genus Pseudomonas. ACTA Path.
Microbiol Scand., 36:531.

Merkel, J. R., E. D. Traganza, B. B. Mukkeijee, T. B. Griffin, and J. M. Prescott, 1964 -Proteo¬
lytic activity and general characteristics of a marine bacterium. Aeromonas proteolytica sp.
M.J. Bacteriol, 87:1227.

Miles, E. M., and Miles, A. A., 1951 -The identity of Proteus hydrophilla. Bergey, et al, and
Proteus melanovogenes. Miles and Hainan, and their relation to genus Aeromonas. Kluyver
and Van Niel. /. Gen. Microbiol, 5:298.

mm.

■

■

GROWTH RESPONSES OF TILAPIA A UREA TO FEED SUPPLE¬
MENTED WITH DRIED POULTRY WASTE

by ROBERT R. STICKNEY, HERBERT B. SIMMONS,

Department of Wildlife and Fisheries Sciences,

Texas ASM University, College Station 77843

and LENTON 0. ROWLAND

Department of Poultry Science,

Texas ASM University, College Station 77843

ABSTRACT

Blue tilapia, Tilapia aurea, fed for a period of 10 wks at 3% of body weight daily on a
commercial trout diet control and the trout diet supplemented at 10, 20 and 30% with dried
poultry waste (DPW)., demonstrated decreasing rates of growth as a function of increasing
DPW levels although the differences were not significant (P >0.10). Food conversions for fish
fed the control and 10% DPW -supplemented diets were similar (1.02 and 1.06, respectively).
In a second, 8-wk experiment, T. aurea fed trout feed supplemented with. 20% DPW failed
to grow when offered feed at 1% of body weight daily. Fish fed the 20% DPW -supplemented
diet at 5% daily grew at approximately the same rate as those fed 3% of that diet in the
initial experiment.

INTRODUCTION

The suitability for aquaculture of various species of Tilapia (family Cichlidae)
has been demonstrated in many parts of the world (Chimits, 1955, 1957; Kirk,
1972; Fryer and lies, 1972), but only recently have Tilapia spp. been considered
as aquaculture candidates in the United States. Tilapia spp. tolerate relatively
high temperatures (Allanson and Noble, 1964), low dissolved oxygen (Denzer,
1968; Fryer and lies, 1972), a wide range of salinity (Lotan, 1960; Chervinski,
1961, 1966; Chervinski and Zorn, 1974) and high levels of ammonia (Stickney,
et al., In Press).

Dried poultry waste (DPW) has been used to feed a variety of terrestrial ani¬
mals including poultry, ruminants and swine (Couch, 1972). Six states have
approved the use of DPW in livestock feed as of March, 1976, although the fed¬
eral government has not cleared DPW in the feed of animals used in interstate
commerce. Channel catfish have been successfully reared on diets heavily supple¬
mented with DPW (Joe Lock , personal communication).

Received for publication: April 13, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

94

THE TEXAS JOURNAL OF SCIENCE

This research represents preliminary studies to determine the feasibility of sup¬
plementing tilapia feed with DPW, and was supported, in part, by Project HM-283 1
of the Texas Agricultural Experiment Station, Texas A&M University. Tilapia
aurea, the blue tilapia, was chosen for study owing to its presence in Texas in
several power plant cooling reservoirs.

MATERIALS AND METHODS

Two experiments were conducted indoors at the Aquaculture Research Center
of the Texas Agricultural Experiment Station, near College Station, Texas. Alumi¬
num troughs (3 .05 m X 5 1 cm) containing water 20 cm in depth were each stocked
with 100 T. aurea averaging 5.5 g. Individual fish weights were not obtained;
however, the fish were of approximately the same age and were selected to be of
uniform size. Well water at ambient temperature (approximately 26 C) was sup¬
plied through flow regulators at a constant rate of 1 .9 1/min in each trough. Ven¬
turi drains were located at the ends of the troughs opposite the Inflow. Supple¬
mental aeration was provided through air stones. The water exchange rate in
each trough was approximately 2.7 hrs. Dissolved oxygen was maintained in
excess of 5.0 mg/1 throughout the study with the exception of one 3-day period
during the first experiment when pump failure occurred and the fish were sub¬
jected to static water until repairs were made. The fish were not fed during that
period.

Three experimental rations were prepared by mixing DPW at 10, 20 and 30%
of the total with No. 3 pellets of Master Mix trout feed (Central Soya and Sub¬
sidiaries, Ft. Wayne, Indiana). The DPW, composed of chicken manure and
spilled chicken feed, was collected from under laying hens and was air dried
prior to use. Unaltered trout feed was used as a control diet. Although the DPW
was not incorporated into the trout feed, the fish did not discriminate against
the DPW portion of any of the diets. Virtually all of the feed offered was con¬
sumed within minutes following presentation.

During the 1st experiment, 4 troughs received the control^ 10, 20 and 30%
DPW -supplemented diets, respectively, at 3% of the biomass of the trough daily
for 10 wks. In the 2nd experiment, 2 additional troughs received the 20% DPW-
supplemented diets at 1% and 5% of biomass daily for 8 wks. All fish were weighed
at 2-wk intervals and the feeding rates were adjusted accordingly.

At the initiation of each experiment and at intervals of 2 wks thereafter, total
biomass determinations were made on the fish in each trough. The average
weight of the fish was determined by dividing the total weight by the number of
fish present (survival was 100% in all troughs during both experiments). In order
to keep handling to a minimum, individual weights were not obtained.

A proximate analysis was run on a sample of DPW utilizing standard analytical
procedures (A.O.A.C., 1975). Amino acid analyses were run on a Beckman Model
120C amino acid analyzer.

TIL API A A UREA

95

RESULTS

The mean weights of T. aurea fed at 3% of body weight daily for 10 wks on
control, 10, 20 and 30% DPW -supplemented diets are presented in Fig. 1. Fish
receiving the control and 10% DPW -supplemented diets demonstrated similar
growth, whereas after the first 4 wks of the experiment, slower growth was
demonstrated by the other groups. Statistical comparison of the slopes of the re¬
gression lines presented in Fig. 1 (Snedecor and Cochran, 1967) indicated no sig¬
nificant differences among the 4 groups of fish (P > 0.10), although the trends
followed a pattern of decreased growth with increased DPW-supplementation.

Figure 1. Mean body weight of T. aurea fed 3% of biomass daily for 10 wks on the con¬
trol diet and diets supplemented with 10, 20 and 30% DPW.

96

THE TEXAS JOURNAL OF SCIENCE

Food conversions (dry weight of feed/wet weight of fish gain) were 1.02 and
1.06 for the control and 10% DPW -supplemented fish, respectively. The food
conversion for fish supplemented with 20% DPW was 1 .25, while that of the 30%
DPW -supplemented fish was 1 .40.

Growth curves of fish fed for 8 wks at 1% and 5% of body weight daily on
the 20% DPW -supplemented ration are presented in Fig. 2. Included in that figure
is the growth curve for the 20% DPW-supplemented fish fed at 3% of body weight
daily during the 10-wk experiment. The slight suppression in growth between 4
and 6 wks for fish on the 3% feeding level corresponded with the 3 -day pump
failure, during which time the fish were not fed. A similar depression did not
occur in the other 2 groups of fish (Fig. 2) since they were stocked subsequent
to the pump failure.

TIME (WEEKS)

Figure 2. Mean body weight of T. aurea fed for 8 wks at 1, 3 and 5% of biomass daily
on a diet supplemented with 20% DPW.

TIL APIA A UREA

97

Proximate analysis of the DPW indicated that the material used in this study
contained 26.2% protein, 12.5% fiber, 2.3% fat and 24.0% ash. Calcium and
phosphorous levels were 7.7 and 1.5% of the total material, respectively. A par¬
tial amino acid analysis and the ammonia level found in the DPW are presented
in Table 1 .

TABLE 1

Amino Acid and Ammonia Content as a Dry Weight % of DPW

Amino Acid

% of DPW

Alanine

0.95

Arginine

0.33

Aspartic Acid

0.72

Cystine

0.32

Glutamic Acid

1.16

Glycine

1.90

Histidine

0.28

Isoleucine

0.49

Leucine

0.90

Lysine

0.72

Methionine

0.07

Phenylalanine

1.82

Proline

0.64

Serine

0.25

Threonine

0.32

Tyrosine

0.53

Valine

0.62

Ammonia

1.08

DISCUSSION

The growth curves obtained from the diets fed at 3% of body weight daily
demonstrated that increases in the level of dietary DPW resulted in slower growth
compared with the control diet. Since these differences were not significant, higher
levels of DPW -supplementation may have been possible without seriously retarding
growth rate. Food conversions were outstanding for fish reared on the control
and 10% DPW-supplemented diets. Even the poorest food conversion value obtained
(1.40 from fish reared on the 30% DPW-supplemented diet) was excellent compared
to other species of fish and to livestock.

With 20% DPW-supplementation, T. aurea can be maintained without growth
if fed 1% of body weight daily (Fig. 2). A 5% daily feeding level appears to be
excessive, since no increase in growth over feeding at 3% daily was apparent, al¬
though the higher feeding level might have resulted in faster growth at higher

98

THE TEXAS JOURNAL OF SCIENCE

temperatures than the average of approximately 26 C which occurred during this
study. Further studies comparing controls at 1% and 5% daily feeding levels are
required, but the lack of significant differences in growth with various levels of
DPW -supplementation implies that the data in Fig. 2 would be representative for
all diets used in the study.

DPW contains an average of about 10% true protein (Couch, 1974) in addition
to other required nutrients. A potential problem associated with DPW is its high
ammonia content, although elevated levels of ammonia were not observed during
this study, nor in a study in which chicken droppings fell directly into a pond
stocked with T. aurea (Stickney,er al., InPress). Similarly, when catfish were fed
high levels of DPW in their feed, no ammonia toxicity was reported (Joe Lock,
personal communication). T. aurea appear to have a high tolerance for ammonia.
Stickney, et al., (In Press) found that some survival occurred in a pond where
ammonia levels in excess of 20 mg/1 occurred consistently over a period of several
weeks.

Among the essential amino acids present in DPW, methionine appeared to be
the most limiting (Table 1). If DPW were utilized as a major source of protein in
commercial T. aurea diets, additional methionine would probably be required. At
present the amino acid requirements of T. aurea have not been determined, so
further research is required before an assessment of the amino acid restrictions
of DPW can be made.

This study gives a preliminary indication of the value of DPW in fish feed.
Further studies to determine the highest levels of DPW supplementation possible
without significant growth retardation are indicated. In addition, the DPW was
not actually incorporated into the feed, even though the consumption rate and
particle size of the control and DPW -supplemented feeds appeared to be similar.
After the protein requirements of T. aurea are determined, the possibility of
replacing significant portions of dietary protein with DPW should be investigated.
Savings in feed costs without significantly reducing growth rate may be possible
as long as the nutritional value of the feed is maintained.

LITERATURE CITED

Allanson, B. R., and R. G. Noble, 1964-The tolerance of Tilapia mossambica (Peters) to
high temperature. Trans, of the Am. Fish. Soc., 93:323.

A.O.A.C., 1915 -Official Methods of Analysis of the Association of Official Analytical
Chemists. 12th Ed. A.O.A.C., Washington, D.C., 1094 p.

Chervinski, J., 1961 -Study of the growth of Tilapia galilaea (Artedi) in various saline con¬
ditions. Bamidgeh , 13:71.

- — , 1966-Growth of Tilapia aurea in brackish water .Bamidgeh, 18:81.

- , and M. Zorn, 1974-Note on the growth of Talapia aurea (Steindachner) and

Tilapia zillii (Gervais) in sea-water ponds. Aquaculture, 4:249.

Chimits, P., 1955 -Tilapia and its culture. A preliminary bibliography. FAO Fishery Bull., 8:1.

TIL API A A UREA

99

- , 195 7 -The tilapias and their culture. A second review and bibliography. FAO

Fishery Bull, 10:1.

Couch, J. R., 1972-Feeding poultry manure to animals. Feedstuffs, 44:24.

- , 1974-Evaluation of poultry manure as a feed ingredient. World’s Poul. Set J.,

30:279.

Denzer, H. W., 1968 -Studies on the physiology of young Tilapia. FAO Fishery Reports,
44:357.

Fryer, G., and T. D. lies, 1912-The Cichlid Fishesof the Great Lakes of Africa. T.F.H. Publi¬
cations, Inc., Neptune City, N.J., 641 p.

Kirk, R. G., 1972- A review of recent developments in Tilapia culture with special reference
to fish farming in the heated effluents of power stations. Aquaculture , 1:45.

Lotan, R., 1960- Adaptability of Tilapia nilotica to various saline condition s.Bamidgeh, 12:96.

Snedecor,G. W., and W.G. Cochran, 19 67 -Statistical Methods, 6th Ed. The Iowa State Univer¬
sity Press, Ames, 593 p.

Stickney, R. R., L. O. Rowland, and J. H. Hesby, 3977-Water quality Tilapia aurea inter¬
actions in ponds receiving swine and poultry wastes. Proc. of the World Mariculture Soc.,
In Press.

DOG FLESH AS A POTENTIAL FOOD RESOURCE FOR CARNIVORES:
AN EXPLORATORY STUDY

by JAMES U. McNEAL and WILLIAM L. GRIFFIN

Department of Marketing, College of Business Administration,

Texas A &M University, College Station 77843

ABSTRACT

The unowned dog population in the U.S. has become so large that it presents serious social
and economic problems. Utilization of these dogs as a food resource for carnivores could
eliminate the problems they cause while contributing to the total food supply. This paper is
the first known effort to discuss this possible solution to the unowned dog problem and offer
some exploratory research findings to support its thesis.

INTRODUCTION

There are now between 10 and 20 million unowned dogs in the U.S. and their
population growth is much more rapid than that of humans. At a meeting of the
Texas Municipal League, it was noted that problems with unowned dogs repre¬
sent the largest number of complaints to city mayors and are of major concern
to public health authorities (Blackwell, 1976).

Problems from unowned dogs are numerous and serious. Among the problems
are (1) livestock damage, (2) wildlife damage, (3) communication of several human
diseases, some resulting in long-term illness and even blindness, (4) dog bites that
may require painful treatment and cause physical disfigurement, (5) contamination
that breeds flies, cockroaches and brown rats, (6) and of course, disposal of the
dogs (Beck, 1974, 1975). The costs to the public of these problems exceed
$500,000,000 annually (Blackwell, 1976).

Solutions to the unowned dog problem have been mainly 2, both of which
add to the costs of the problem while producing minimal results. One is the dis¬
posal of the dogs by animal humane groups that “put them to sleep.” Eighteen
million dogs are put to sleep each year at a cost of $125,000,000 (Djerassi, et al.,
1973). The other solution is birth control among owned dogs including spaying,
neutering and many birth control devices (Seager, 1976).

Converting this public liability to an asset should be given serious considera¬
tion. One possible solution may be to view the unowned dog as a food resource
for other carnivores such as household pets and many zoo animals. If unowned
dogs were slaughtered (instead of converted to ashes) and prepared as a food for

Received for publication: March 28, 1977.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

102

THE TEXAS JOURNAL OF SCIENCE

household pets and zoo animals, 2 positive results would be (1) a reduction of
surplus dogs, and (2) an increased availability of human foods by a reduction in
the edibles now consumed by pets and the addition of the meats that are now
eaten by feral dogs. This net increase in food available for human consumption
should be of interest to those concerned with our hungry world.

The existence of aversion to feeding dog flesh to household pets is quite ap¬
parent. In spite of the fact that dog flesh is consumed by humans around the
world, feeding that same flesh to treasured pets is likely to be unacceptable to
many people. However, this aversion should not deter us from considering the
feasibility of the idea. Desensitization therapy (Schaefer and Martin, 1975) is
now in the realm of scientific status and probably could reduce the aversion over
time through its application in the mass communications media. Also, aversion
might be reduced by changing the name of the product just as we have done
with fish roe (caviar) or calf thymus (sweetbread).

Recognizing the seriousness of the unowned dog problem, the investigators
began an exploration into the feasibility of utilizing these dogs as a food resource.
The slight possibility of solving the unowned dog problem while making a con¬
tribution to the world hunger problem were adequate incentives for the research
effort.

METHOD

The investigators assumed that food scientists had studied dogs as a food
resource since dogs have been consumed for centuries by some humans around
the world. Therefore, the first step taken in the exploration was an exhaustive
search of the literature. The search was essentially unproductive. It produced
some minor notes indicating some consumption of dog flesh by humans (Simoons,
1961). Two sources offered a 1-line presentation of estimated nutritional values
of dog flesh with no supporting discussion (Spector, 1956; HEW, 1972). No em¬
pirical reports were found, however, about dog flesh as a food resource for carni¬
vores.

The estimated nutritional values for dog flesh found in the literary research
are presented in Table 1 along with values for lean beef and pork. A visual com¬
parison among the 3 meats suggests that dog flesh generally possesses nutritional
characteristics comparable to beef and pork. For example, protein and iron are
slightly higher for dog, while calcium and niacin are lower. If one examines the
nutritional contents of wet and moist commercial dog foods, he can see that the
use of dog flesh in lieu of other meats such as beef and pork would not change
the nutritional value of the feeds.

The consequence of the unproductive literary search combined with the en¬
couraging interpretation of Table 1 was a mail survey conducted during the
month of November, 1976, with 62 authorities in the field of nutrition. These
experts were suggested by the Food and Nutrition Board of the National Academy
of Sciences. Forty-four of the experts were affiliated with 36 universities in the

DOG FLESH

103

TABLE 1

Nutritional Composition of Lean Cattle, Pork and Dog*
(100 grams, edible raw portions)

Component

Measure

Cattle

Pork

Dog

Water

%

70-71. 8a

50.1-65a

60.8-76a

Ash

g

1.0

0.8

1.2

Calcium

mg

9-lla

8

3.3-26a

Carbohydrate, Total

g

0

0

0

Chlorine

mg

55b

NA

50-75

Cholesterol

%

0.24b

0.06b

0.3

Choline Phospholipid

%

2.3b

NA

2.9

Fat

g

7.2

35

23.5

Fiber

g

0

0

0

Food Energy

cal

150

376

274

Iron

mg

3.0

2.1

3.6

Lipid

%

10-14b

8b

14

Magnesium

mg

32b

o

o

cr

44

Niacin

mg

6.7

3.7

1.8

Phospholipid

%

3.7

3.1b

5.8

Phosphorus

mg

1 7 1-2 1 0a

151

1 85-21 9a

Potassium

mg

489

NA

292-320a

Protein

%

19. 3b

19b

21

Riboflavin

mg

0.34

0.16

0.08

Sodium

mg

93

NA

74—1 3 1 a

Thiamine

mg

0.07

0.69

0.04

* Developed from Spector (1956) and HEW (1972)

Combination of Both Sources
bNot Necessarily Lean

U.S., 2 in Canada, 1 in Taiwan, and 1 in Nigeria, 12 worked as nutritional specialists
for U.S. food companies, and 6 were associates of hospitals in the U.S. They were
asked to furnish any information or opinions regarding (1) the nutritional value
of dog flesh, and (2) the feeding of dog flesh to household pets. Responses were
received from 70% of the nutritionists.

The responses of the nutritionists were tabulated accordingly: (1) nutritionists
who thought that dog flesh was nutritious, (2) nutritionists who felt dog flesh
could be used as a pet food source, and (3) nutritionists who were concerned
with the aversion factor. Also, an attempt was made to ascertain the degree of
general knowledge about the nutrition of dog flesh held by the nutritionists.

Concurrently, a personal survey was conducted with a random sample of 121
pet owners during a 3-week period in November, 1976, in Bryan and College
Station, Texas. The interviews were conducted by the authors and one student

104

THE TEXAS JOURNAL OF SCIENCE

in 3 supermarkets. The respondents were identified as a result of their selection
of a cat or dog food from the supermarket shelf. The 2 objectives of this investi¬
gation were (1) to obtain an estimate of the amount of aversion held by pet
owners to feeding canine flesh to their pets, and (2) to obtain estimates of aver¬
sion to other meats as pet foods for comparison. The respondents’ aversive attitudes
were categorized according to the respondents’ age, gender, and social class.1 The
degree of aversion to dog flesh was expected to be high, but confirmation was
desired. With a good estimate of the amount of aversion to dog flesh, particularly
in comparison with other meats, some decision could be made about the possibility
of reducing it through applications of desensitization therapy.

FINDINGS
Survey of Nutritionists

Approximately 2/3 of the responding nutritionists believed dog flesh to be
nutritious. Most of the respondents compared the nutrition of dog flesh to that
of pork, and 17.5% supported their belief with the HEW dog nutrition table men¬
tioned above. Over 40% of the nutritionists felt that dog flesh could be used as
an additional food source for household pets. A few of the respondents were, in
fact, enthusiastic about the idea.

Half of the respondents noted the aversion factor. Some felt it might be cir¬
cumvented; some felt it could not. A few voiced strong objections to the idea on
the basis of being pet owners.

An interesting aspect of the survey was that 1/3 of the sample members said
that they had no knowledge about the subject. Since mail surveys often are un¬
answered, particularly when the respondents have nothing to offer or are uninter¬
ested, this response was unexpected. It apparently demonstrates the profession¬
alism of the sample members, and the interest in the topic by the group.

The mail survey appeared to produce 2 conflicting findings. It confirmed the
unproductive literary search since no respondent referred to any research on the
topic. However, the survey produced the impression that many nutritionists have
knowledge and interest about dog flesh as a food resource. The interest was con¬
firmed by the fact that 10 of the nutritionists offered to act as consultants on a
funded project to study the topic in depth.

Survey of Pet Owners

The main goal of surveying the pet owners was to obtain an estimate of their
aversion to certain foods, including dog meat, that might be fed to their pets.
Questions were asked about 3 groups of foods based on expected levels of aver¬
sion. The group of meats expected to elicit low aversion (<. 25% of n) consisted
of pork, lamb and poultry. Fish and horsemeat, the second group, were expected
to receive mild to strong aversion scores (>25%, < 75% of n) from dog owners.

Occupation provided a serviceable index for determining social class.

DOG FLESH

105

Similar scores for cat owners were expected for horsemeat, but fish was antici¬
pated to produce a lower measure. The high aversion group of meats (>75% of
n) consisted of dog and armadillo. Table 2 presents a summary of the aversion to
these meats according to respondents’ age, sex and social class, and Table 3 sum¬
marizes the aversion by cat and dog ownership.

TABLE 2

Degree of Aversion to 7 Meats as Possible Ingredients in Dog and/or Cat Food
by Age, Sex, and Social Class of Respondents

Meat

Age

Sex

Social Class

20-29

30-39

40 & Over

Male

Female

Lower

Lower

Middle

Upper

Middle

(%)

(%)

(%)

Poultry

7

6

7

4

9

8

8

5

Lamb

5

20

23

17

16

24

24

11

Pork

32

40

44

17

30

48

44

35

Fish

22

17

16

19

20

20

18

20

Horse

27

23

21

15

29

24

26

21

Armadillo

22

37

40

29

36

45

44

26

Dog

42

57

67

46

64

68

70

47

TABLE 3

Degree of Aversion to 7 Meats as Possible Ingredients in
Dog and/or Cat Food by Kind of Pet Owned

Meat

Dog Owner

Cat Owner

(%)

(%)

Poultry

8

5

Lamb

18

14

Pork

39

40

Fish

27

7

Horse

21

28

Armadillo

35

30

Dog

54

61

The aversion to lamb and poultry was low regardless of age, sex or social class.
However, there was much more aversion to pork (about 40% of sample) than to
lamb (16%) and poultry (7%) for both cat and dog owners. The amount of aver¬
sion to fish and horse meat as pet foods was lower than expected. Aversion to
fish for dogs was expected to be mild to strong. Actually, 73% of the sample
said they would feed fish to dogs. The aversion to horsemeat was about the same
for dog and cat owners (around 25%).

106

THE TEXAS JOURNAL OF SCIENCE

The expected very strong aversion to armadillo and dog meat was less than
anticipated. Aversion to armadillo was only 26% among upper middle social class,
and around 45% among lower and lower-middle pet owners. Aversion to armadillo
increased with age, but only to a high of 40%. There was not much difference
between male and female respondents’ aversion to armadillo. Acceptance of arma¬
dillo for pet dogs was about the same as its acceptance for pet cats.

The major concern of the pet owner study, that is, the degree of aversion to
dog meat as a pet food, proved to be a surprise. About 60% of people age 20-30
found dog meat acceptable, but only 39% of people over 30 accepted it. Over 1/2
of upper middle class respondents found dog meat acceptable for pet food, but
only around 30% of lower and lower middle class pet owners accepted it. Aver¬
sion did not differ significantly between dog and cat owners. In general, aversion
to dog meat as pet food was strong but not extreme.

SUMMARY AND IMPLICATIONS

The research reported here was an initial exploratory study about the nutritional
aspects of dog flesh and the amount of aversion among scientists and pet owners
toward using it as a food for other carnivores. It is a small beginning, but offers
more than presently exists in the literature.

A significant % of the nutritionists surveyed felt that dog flesh was a feasible
food resource for house pets. In spite of the paucity of data about the nutritional
value of dog flesh, a majority of those nutritionists in the study indicated it to
be nutritious. Only 1/2 of the nutritionists were concerned with the aversion
factor of consuming dog flesh. Since it is clearly stated in much literature how
socially important the dog is in our society, the investigators intuitively antici¬
pated the % to be much higher.

The survey results made it apparent that some nutritionists have knowledge
and much interest in the food potential of dog flesh. However, the literature does
not support this finding. Probably the aversion factor has caused the food scien¬
tists to not study dogs as a food source, or at least not to publish any results. As
one respondent said, “I do not want to be associated with such an idea even
though it may have merit.”

Aversion to dog flesh as a pet food is not extreme among pet owners. In fact,
the aversion to dog flesh among young people holding professional and managerial
positions is only slightly higher than the aversion to pork as a pet food. Middle-
aged women in lower middle class families had higher aversion to dog meat as
pet food. In general, the research results reported here do not indicate that attempt¬
ing to utilize dogs as a food resource for household pets is a futile task.

Besides the aversion factor, there is other opposition to utilizing dogs as an
animal feed. Dog owners may feel that such a program is a threat to dog owner¬
ship. To the contrary, elimination of unowned dogs will remove “legal eyes”
that are reviewing the dog population (Blackwell, 1976). Another probable element

DOG FLESH

107

of opposition to the proposed idea is pet food manufacturers. They may errone¬
ously view it as a threat to their industry. However, at most what is being suggested
is a change in some raw materials used by the pet food industry.

Finally, animal humane groups may be opposed to this suggested means for
alleviating the unowned dog problem. Hopefully, these groups will see that the
proposed program will terminate unwanted dogs as humanely as they would while
having more effect on the unowned dog population. Hence, this proposed pro¬
gram is congruent with the goals of these 3 groups that might oppose it.

Using dog flesh as a food resource may be a viable alternative to solving the
unowned dog problem. Its other merit is its contribution to the world hunger
problem. It may be that experiments using unowned dogs as a food resource
could be the beginning of a new industry comparable in nature to the beef or
pork industry. An indication of the economic possibility is the fact that consumers
spend $2 billion annually on commercial cat and dog foods.

Many serious questions must be answered, however, in addition to the nutritional
and aversion aspects.

Some of these are concerned with

(1) Collecting and containing the dogs

(2) Deleting diseases in the dog carcasses

(3) Developing feeding and breeding procedures

(4) Developing slaughtering procedures

(5) Determining the final forms of the meat to be offered

(6) Determining the economic feasibility of the undertaking

(7) Dealing with local and national laws

(8) Convincing the pet food industry to use dog flesh as a food substitute or
additive in lieu of other edibles.

In sum, the dearth of research reported on dog flesh as a food source is dis¬
appointing. The interest shown by the nutritionists surveyed on this subject, and
the less-than-expected amount of aversion by pet owners, are encouraging. They
suggest that basic interdisciplinary research should be undertaken to judge the
value of using dog flesh as a food resource for carnivores. The social and economic
pay-offs for such an effort could be substantial.

LITERATURE CITED

Beck, A. M., 1974-Ecology of unwanted and uncontrolled pets. Proceedings of the National
Conference on the Ecology of the Surplus Cat and Dog Problem. American Human Society,
Denver, pp. 31-39.

- , 1975 -The public health implications of urban dogs. A mer. J. Pub. Health, 29:1315.

Blackwell, W. R., 1976-The politics of dog control. Tex. Town & City, 43:4.

Djerrassi, C., A. Israel, and W. Jochle, 197 3- Planned parenthood for pets? Bull. At. Sci.,
29:10.

Health, Education and Welfare, 197 2-Food Composition Table for Use in East Asia. Washington,
D.C.

108

THE TEXAS JOURNAL OF SCIENCE

Schaefer, H. H., and P. L. Martin, 1975 -Behavioral Therapy, 2nd Ed. McGraw-Hill, New York,
N.Y.

Seager, S. W. J., 1976- Review of reproduction control in cats and dogs. Proceedings of the
National Conference on Dog and Cat Control American Humane Society, Denver,
pp. 228-249.

Simoons, F. J., 1961 -Eat Not This Flesh. The University of Wisconsin Press, Madison, Wise.

Spector, W. S., 19 56 -Handbook of Biological Data. W. B. Saunders Company, Philadelphia,
Penn.

ESTIMATIONS OF THE RATIO OF INDUCED FISSION TO SPON¬
TANEOUS FISSION IN URANIUM ORES1

by MOSES ATT REP, JR., K. S. TASA and J. D. SHERWOOD

Department of Chemistry, East Texas State University,

Commerce 75428

ABSTRACT

The mass spectrometric analysis of noble gases in uranium ores have been previously re¬
ported for geologically old uranium minerals. These data have been treated in a new method
to determine the ratio of thermal neutron induced fission of 23SU to the spontaneous fission
of U. The results indicate that xenon isotopes are more suitable for this method of treat¬
ment than the krypton isotopes. There is evidence to believe that in uranium minerals the
basic modes of nuclear fission are the U neutron-induced fission and the U spontane¬
ous fission. The results indicate that the ratio, R, for individual uranium mineral samples
range from ^0.5 to ^uO. Results are also consistent with previously reported radiochemical
methods and the general geochemical composition of certain uranium minerals. Also, the
self-sustaining nuclear reactor character of the Oklo Mine deposit is evident. The method is
simple and provides a rapid means of determining this nuclear geochemical property of uranium
minerals.

INTRODUCTION

There have been no reports on the generalization of the nuclear fission phenom¬
enon in uranium ores and minerals with respect to the spontaneous and induced
fission processes. Several studies concerning the ratio of neutron induced fission
of 235 U to the spontaneous fission of 2 38 U in uranium systems have been re¬
ported for natural uranium laboratory systems or a specific uranium ore (Kenna
and Kuroda, 1960; Kenna and Attrep, 1966; Attrep and Sherwood, 1972; Raut
and Attrep, 1975).

In discussing the nuclear geochemical parameter of the induced fission con¬
tribution in natural uranium systems, consideration must be given to the sources
contributing to the total neutron population in the ore. These are neutrons arising
primarily from the spontaneous fission of 2 3 8 U, neutron-induced fission of 2 3 5 U,
and from (a, n) reactions on low Z elements. Cosmic-ray neutron contributions
have been shown to be negligible (Montgomery and Montgomery, 1939).

The isolation and identification of element 94, plutonium, from pitchblende,
brought attention on the requirement that an additional neutron contribution
must be present in addition to that supplied from the spontaneous fission of 238 U.

Presented in part at the National A.C.S. Meeting, Los Angeles, 1974.

Received for publication: July 1, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

110

THE TEXAS JOURNAL OF SCIENCE

It was then speculated that (a, n) reactions on low Z elements might give the ex¬
planation (Peppard, et al., 1951; Levine and Seaborg, 1951). Recently, in this
laboratory (Attrep and Sherwood, 1972; Raut and Attrep, 1975) it has been
shown that the ratio of induced fission to spontaneous fission in natural uranium
can be increased by (a, n) reactions on beryllium as had been previously expected.

Some earlier mass spectrometric works (Macnamara and Thode, 1950; Fleming
and Thode, 1953; Wetherill, 1953; Young and Thode, 1960) have investigated
the fissiogenic isotopic composition of krypton and xenon in a number of uranium
and thorium samples. The work of Young and Thode (1960) presented spontan¬
eous fission spectra for both fissiogenic krypton and xenon isotopes. These mass
spectrometric measurements of the noble gas fission products have enabled us
to project the nuclear geochemical history of these minerals with respect to the
induced fission of 235 U.

METHODS AND EQUATIONS

It will be assumed at this point that the total number of any fissiogenic iso¬
tope of krypton or xenon arises primarily from the spontaneous fission of 238 U
and the thermal neutron-induced fission of 235 U. With this basic assumption, the
average ratio of the neutron-induced fission of 235 U to the spontaneous fission
of 238 U can be estimated using the concentrations of these fissiogenic noble gas
nuclides produced in uranium ores. The model presented here will allow a nuclear
geochemical description of uranium ores considering the fission phenomena. In
effect, this will reflect on the neutron population for the particular uranium
mineral since the amount of neutron induced fission of 235 U is a direct consequence
of the neutron fluence.

The number of atoms of a particular fissiogenic nuclide produced from the
spontaneous fission process of 238 U can be expressed according to the following
relationship:

a 238

Ns = N238 [exp(X2t38)T-l]-|iFYs (1)

At

where Ns is the number of atoms of the particular fissiogenic nuclide produced
by the spontaneous fission of 238 U; N238 is the number of 238 U atoms present
today in the sample; T is the fissiogenic retention time of the ore sample; Y§ is
the spontaneous fission yield for this fissiogenic nuclide; X2|8 is the spontaneous
fission decay constant for 238 U; and A238 is the total decay constant for 238 U.

In a similar manner, the number of atoms of the same fissiogenic nuclide
produced by the thermal neutron induced fission of 235 U in the mineral can be
expressed as follows:

Nj = N235 [exp(Af5)T- 1]T£Yj

(2)

RATIO OF INDUCED FISSION TO SPONTANEOUS FISSION

111

where Ni is the number of atoms of the fissiogenic nuclide produced from the
thermal neutron induced fission of 235 U atoms initially present in the ore sample;
$ is the neutron fluence; a is the thermal neutron fission cross-section; Yj is the
induced fission yield for this nuclide; and A235 is the sum of all the decay constant
process for 235 U.

The overall ratio, R, for the thermal neutron-induced fission of 235 U to the
spontaneous fission of 238 U experienced in the uranium mineral is expressed as
the total number of induced fission events in relation to the total number of
spontaneous fission events for the same time period, T, the retention time of
the ore. This is expressed as follows:

N23s[exp(Xf5)T-l]-^-

R- _ (3)

N238 [exp(A2t38)T - 1]— llg-

At

For one fissiogenic nuclide, the total concentration is expressed as N[. This
will represent the concentration of that nuclide in the uranium ore. This concen¬
tration is assumed to be fissiogenic and is assumed to be comprised of 2 compon¬
ents: spontaneous fission produced, Ng, and induced fission produced, Nj. Thus,
N[ = Nj + N| and the total concentration of a specific fissiogenic nuclide in a uranium
ore is

X238

N't = N238 [exp(A238)T - + N235 [exp(A23S)T - 1] -fg-Y| (4)

At At

By combining equations (3) and (4) and making appropriate rearrangements, Nj
is expressed as a function of the value R as shown below:

X238

n; = N238 [exp(A238)T - + RYJ). (5)

At

If the absolute concentration value of the particular fissiogenic nuclide is known,
along with the other parameters including the retention time for the ore, the
ratio of induced fission to spontaneous fission can be obtained. However, for the
most part, these values are unattainable, and some of those which probably could
be estimated may be unreliable.

Upon inspection, it is noted that a similar expression can be written for another
fissiogenic nuclide in the same ore :

X238

N't' =N238 [exp(A2t38)T- 1]-|C-(Y'S'+ RY”). (6)

At

If equation (5) is divided by equation (6), the ratio of the total number of atoms
of one fissiogenic isotope in relation to another can be expressed as follows in

112

THE TEXAS JOURNAL OF SCIENCE

which this relationship is a function of R the fission yields of the isotopes in
question, and the ratio of 2 fissiogenic nuclides. This is expressed as follows:

k _ (y;+ryd (7)

N? (Y”+ RY")

As shown in equation (7), the observed ratio of fissiogenic products is not a func¬
tion of T, but is a function of single variable, R, the average ratio of thermal neu¬
tron-induced fission of 235 U to the spontaneous fission of 238 U. The fissiogenic
nuclides used in this study are the krypton and xenon isotopes. The fission yields
for the spontaneous fission process of 238 U and the thermal neutron-induced
fission process of 235 U are shown in Table 1. The yields for 129 Xe have been
omitted because the spontaneous fission yield is set at a lower limit at <0.012%
(Wetherill, 1953). The spontaneous fission yields used in this study are those of
Young and Thode (1960).

TABLE 1

Percent fission yields of xenon and krypton isotopes form 238U spontaneous
fission and 23 5 U thermal neutron induced fission.

Mass Number

238U Spontaneous

Fissiona
% Yield

235U Neutron Induced
Fission^

% Yield

Krypton Isotope

83

0.0327 ±0.0028

0.561 ±0.012

84

0.1220 ± 0.0121

1.032 ±0.022

86

0.9510 ±0.0570

2.036 ±0.019

Xenon Isotope

131

0.524 ±0.031

2.92 ±0.02

132

3.630 ±0.220

4.35 ±0.04

134

5.140 ±0.310

7.93 ±0.24

136

6.300 ±0.380

6.39 ±0.11

aFrom Young and Thode, (1960).

bThe % yields presented are an average of 5 reported values, as presented in Hyde (1964);
the errors associated are “standard deviations.”

Using equation (7) and the fission yields given in Table 1, the relationship be¬
tween R, the ratio of induced to spontaneous fission, and the fissiogenic noble
gas pairs are produced. Variation of R with the ratios of the total number of
fissiogenic isotopes of krypton pairs are shown in Fig. 1 for the 3 combinations:
86Kr/83Kr, 86Kr/84Kr and 84Kr/83Kr. In a similar manner, Fig. 2 shows the
ratio of the total number of atoms of one xenon isotope to that of another versus R.
From inspection of Fig. 1 it appears that a small variation of R, the ratio of in¬
duced to spontaneous fission, is more readily reflected in the relatively large

Krypton Isotopic Ratio (Atom/Atom)

RATIO OF INDUCED FISSION TO SPONTANEOUS FISSION

113

Figure 1. The average ratio of induced fission of 235U to spontaneous fission of 238U
as a function of ratio of krypton isotopic ratios.

change of the 86/84 and 86/83 krypton ratios as compared to that for the krypton
pair 84/83. Also, from Fig. 2 the same conclusion can be drawn from the xenon
isotopes. The xenon isotope pairs of 134/131, 136/131 and 132/131 exhibit the
most useable relationships.

RESULTS AND DISCUSSION

Treatment of the reported mass spectrometric analyzed noble gas nuclides in
a number of uranium minerals using the relationship devised (equation 7) yields

Xenon Isotopic Ratio (Atom/Atom)

114

THE TEXAS JOURNAL OF SCIENCE

Figure 2. The average ratio of induced fission of 235u to spontaneous fission of 238U
as a function of ratio of xenon isotopic ratios.

values of R. The evaluated R values from the 3 krypton isotope pairs and from
the 3 xenon isotope pairs are presented in Table 2. The % U308 and the age of
the ores are those given by the authors from which the noble gas values were
taken. The type of uranium sample analyzed is predominately pitchblende; how¬
ever, there are a few uraninite samples. The name and location of the uranium
minerals are also those reported in the original papers.

The R values derived from the noble gas isotopic ratios exhibit some features
which are of importance for further discussion of the nuclear fission phenomenon
in uranium minerals. There is consistency of the value of R for the 3 sets of xenon

RATIO OF INDUCED FISSION TO SPONTANEOUS FISSION

115

TABLE 2

Derived values for the ratio of thermal neutron induced fission of 235U to the spontaneous

2 38

fission of U in uranium ore samples.

Sample

'AI3Ob

Age'

(million

years)

R for Xenon Isotopes
132 134 136

1 31 131 131

Average

R for Krypton Isotopes
86 86 84

84 83 83

Average

Ref.

Pitchblende.

Katanga,

Belgian Congo

65.21

640

0.42

0.42

0.40

0.41

0.42

0.77

>1

0.73

Fleming and
Thode (1953)

Pitchblende,

Belgian Congo

-

650

0.42

0.42

0.44

0.43

0.44

0.32

0.17

0.31

Wetherill

(1953)

Pitchblende,

Belgian Congo

66.5

635

0.46

0.48

0.48

0.47

0.39

0.37

0.30

0.35

Young and
Thode (1960)

Pitchblende,

Fagle Mine.
Beaverlodge

45.5

1630

0.34

0.33

0.34

0.34

Fleming and
Thode (1953)

Pitchblende,
l ag le Mine

53.4

1700

0.26

0.25

0.26

0.26

0.26

0.22

0.16

0.21

Young and
Thode (1960)

Pitchblende.

Ace Mine,
Beaverlodge

16.96

1600

0.27

0.28

0.28

0.28

--

--

Fleming and
Thode (1953)

Pitchblende,

Beaverlodge

Lake

48.6

785

0.08

0.09

0.09

0.09

0.02

0.08

0.19

0.10

Young and
Thode (1960)

Pitchblende.

Nesbitt Labinc.
Beaverlodge ■

27.55

0.13

0.15

0.13

0.14

— -

Fleming and
Thode (1953)

Pitchblende,

Lake Athabaska

13.99

1690

0.12

0.12

0.12

0.12

....

Fleming and
Thode (1953)

Pitchblende,

Great Bear

Lake

36.46

1370

0.1 1

0.13

0.12

0.12

0.33

0.08

" 0

0.14

Fleming and
Thode (1953)

Pitchblende,

Great Bear

Lake

0.12

0.13

0.12

0.12

0.30

0.09

0.0

0.13

Macnamara and
Thode (1950)

Pitchblende.

Great Bear

Lake

51.1

1400

0.23

0.24

0.24

0.24

0.19

0.19

0.19

0.19

Young and
Thode (1960)

Uraninite.

Jahala Lake

35.8

1740

0.04

0.04

0.04

0.04

0.09

0.05

0.04

0.06

Young and
Thode (1960)

Uraninite,

Cardiff Township,
Ontario

71.0

0.03

0.03

0.03

0.03

Fleming and
Thode (1953)

Uraninite,

Cinch Lake

34.2

1120

0.04

0.04

0.04

0.04

0.05

0.05

0.08

0.06

Youne and
Thode (1960)

Pitchblende,

Rabbit Lake

5.3

1200

0.16

0.16

0.16

0.16

0.0

0.08

>1

0.36

Drozd, ct al..
(1974)

Pitchblende,

Rabbit Lake

3.2

1200

0.04

0.04

0.04

0.04

0.0

0.03

0.13

0.05

Drozd, et al.,
(1974)

Pitchblende,

Rabbit Lake

10.3

1200

0.05

0.05

0.05

0.05

0.0

"0

'0

0.0

Drozd, ct al.,
(1974)

Pitchblende,

Oklo Mine

7.6

1700

>1

>1

: >r

>1

>1

>1

>1

>1

Drozd, ct al..
(1974)

Pitchblende,

Oklo Mine

17.5

1700

>1

>1

; >1

>1

>1

>1

>!

>1

Baudin. ct al. ,
(1972)

116

THE TEXAS JOURNAL OF SCIENCE

i

isotopes for all samples. This is not the case for those values of R arising from
the krypton isotope ratios. In general, however, there is poor agreement between
those R values from the krypton isotopes and those from the xenon isotopes. It
should be noted that the experimental data used for the krypton isotopes have
been reported with errors in some cases up to 100%. In view of the experimental
error involved our discussion concerning the ratio, R, will be drawn primarily from
the xenon isotopic values.

The 3 Belgian Congo pitchblende samples show similar R values that are very
high in relation to the other samples. For these 3 samples the values of R are 0.47,
0.43, and 0.41. It is not known if this is one single sample analyzed by the 3
separate laboratories. Aside from the Oklo Mine sample which had undergone a
self-sustaining nuclear fission reaction, these samples have the highest contribution
of thermal neutron induced fission. From the determination of 36 Cl(t1/2 = 3 x 10s
years) from African (Belgian Congo) pitchblende and Canadian (Great Bear Lake)
pitchblende, Kenna and Kuroda (1960) reported R values of 0.33 and 0.19,
respectively. The African pitchblende samples have somewhat higher values of R
than that reported for the 36C1 work. However, the radiochlorine value of R= 0.19
is in good agreement with the Great Bear Lake samples reported in Table 2, i.e.,

R = 0.12, 0.12, and 0.24. Two considerations must be made, however. First, be¬
cause the samples for mass spectrometric analysis and radiochemical analysis are
probably not identical there maybe major chemical composition factors bringing
about these differences if they are significant. Secondly, the radiochlorine work
represents the ratio of induced fission to spontaneous fission over the past 3 million
years whereas the mass spectrometric xenon isotope value for R includes essentially
the lifetime of the mineral, up to over 600 million years. The ratio of induced
to spontaneous fission measured by stable isotopes is an average value over the
retention age of the sample. Due to the shorter half-life of 235 U with respect to
238U, the 235U concentration would be enriched in the geological past and the
ratio of induced fission to spontaneous fission measured in this manner would be
different from that determined by a radionuclide.

Non-homogeniety of either chemical or physical properties of the mineral can
yield variations in the value of R. For example, the variation of R within a partic¬
ular pitchblende deposit is well pronounced in the Beaverlodge pitchblende sam¬
ples. R ranges from 0.09 to 0.34. Significant increases in the contribution of ther¬
mal neutron induced fission of 235U is probably due to the presence of low Z
elements like beryllium which are capable of increasing the total neutron flux by
(a, n) reactions (Attrep and Sherwood, 1972; Raut and Attrep, 1975). Variations
in chemical composition of elements capable of undergoing (a, n) reactions would
account for such a variation of induced fission contribution in the samples.

The presence of elements such as the lanthanides may decrease this neutron
flux due to their characteristically high neutron absorption cross-sections. This
effect is observed in the 3 uraninite samples. Uraninites usually contain lanthanides
and would exhibit low values of induced fission. For these samples, R = 0.03,

RATIO OF INDUCED FISSION TO SPONTANEOUS FISSION

117

0.03, and 0.04, respectively. These results are consistent with those obtained by
Young and Thode (1960) using complete spectral analysis. Slight variations of
the concentrations of either group of elements from location to location will
definitely alter the thermal neutron-induced fission factor.

Figure 3 is a plot for the xenon isotopes 134/131 versus 136/131. Similarily
in Fig. 4 a xenon isotopic composition graph for 134/132 versus 136/132 is
shown. In each figure the points which represent the pure spontaneous fission of

^Xe/*3'xe (Atom/Atom)

Figure 3. 1 34Xe/1 31Xe versus 1 36Xe/1 31Xe plot for uranium minerals.

238U and thermal neutron-induced fission of 235 U are designated. We have also
included the point representing 239Pu neutron-induced fission (Hyde, 1964). The
individual points are not identified except for the Oklo mine samples.

In Fig. 3, it is quite evident that all points lie on a line between the values of
pure spontaneous fission of 238 U and the thermal neutron- induced fission of 235 U.

Figure 4. 134Xe/132Xe versus 136Xe/132Xe plot for uranium minerals.

This trend strengthens our original assumption that in uranium ores there are
basically only the 2 types of fission processes previously stated. Also, if the graph¬
ical method of analysis of Podosek (1970) is employed for Fig. 3 in order to establish
the amounts of the individual xenon isotopes originating from the thermal neutron-
induced fission of 235 U and the spontaneous fission of 238U, the same R values
are obtained for each sample which are derived by the first proposed method.

The Oklo Mine sample shows the induced fission component to be extremely
dominant. From Table 2 the R value is >1. This agrees with the evidence for a
naturally self-sustaining chain reaction taking place in this unique uranium deposit
(Baudin, et al, 1972). (An excellent review of this natural nuclear reactor has
been recently published by Cowan (1976).) The chain reactor aspect of the Oklo
Mine is clearly indicated from our proposed treatment of the mass spectrometric
data of the xenon isotopic composition and the isochronic treatment of the data
of Fig. 3 and 4.

RATIO OF INDUCED FISSION TO SPONTANEOUS FISSION

119

In Fig. 4 using the xenon isotopic pairs 134/132 versus 136/132, the same
trend is expressed as in Fig. 3. However, this trend is not as neat. The purpose of
plotting the 239Pu value, especially in Fig. 3, is to indicate the possibility of this
mode of fission in the self-sustaining chain reactor ore, the Oklo Mine. Drozd,
et al, (1974) and Baudin, et al, (1972) have both indicated that this type of
fission occurred in this uranium deposit. Although it is not evident in Fig. 3 be¬
cause the 239Pu(n, f) point lies close to the 235U(n, f) point, it does become some¬
what convincing in Fig. 4 that the 239Pu neutron induced fission process occurred
in this deposit.

CONCLUSION

This treatment of the noble gas mass spectrometric analysis of uranium ores
using our initial approach indicates that our basic approach to the nuclear geo¬
chemical phenomena of fission is correct. The use of xenon isotopes is more re¬
liable than those data of the krypton isotopes. The following are concluded from
this study: (a) there are predominately 2 modes of fission in normal uranium
minerals: the spontaneous fission of 238U and the therman neutron-induced
fission of 235 U; (b) the thermal neutron-induced fission of 2 35 U contributes up
to about 30% of the total fission events in the ore; (c) the Oklo Mine samples are
the only uranium mineral samples which clearly demonstrate a predominance of
235U thermal neutron induced fission giving support to the idea that it was a
natural nuclear chain reactor; (d) there appears to be no correlation between age,
uranium content and the ratio of induced to spontaneous fission in the uranium
samples.

ACKNOWLEDGEMENT

This work was supported by the Robert A. Welch Foundation, Grant T-291.
LITERATURE CITED

Attrep, M., and J. D. Sherwood, 1972-The effect of (a, n) reactions on the ratio of induced
to spontaneous fission in natural uranium. J. Inorg. Nucl. Chem., 34:435.

Baudin, C., C. Bain, R. Hagemann, M. Kremer, M. Lucas, L. Merlivat, R. Molina, G. Nief,
F. Prost-Marcehal, F. Regnaud, and E. Roth, 1972-Quelques donnees nouvelles sur les
reactions nucleaires en chain que se sont produites dan le gisement d’Oklo. C. R. Acad.
Sci. Paris, 275:2291.

Cowan, G. A., 1976-A natural fission reactor. Sci. Am., 235(1): 36.

Drozd, R. J., C. M. Hohenberg, and C. J. Morgan, 1974-Heavy rare gases from Rabbit Lake
(Canada) and Oklo mine (Gabon): natural spontaneous chain reactions in old uranium
deposits. Earth Planet. Sci. Lett., 23:28.

Fleming, W. H., and H. G. Thode, 1953-Neutron and spontaneous fission in uranium ores.
Phys. Rev., 92:378.

120

THE TEXAS JOURNAL OF SCIENCE

Hyde, E. K. (Ed.), 1964— 77ze Nuclear Properties of the Heavy Elements, Vol. 3. Prentice-
Hall, Inc., Englewood, N.J.

Kenna, B. T., and M. Attrep, Jr., 1966 -The ratio of induced fission vs. spontaneous fission
and the trace element analysis in pitchblende. J. Inorg. Nucl. Chem., 28:1491.

- , and P. K. Kuroda, 1960-The ratio of induced fission vs. spontaneous fission in

pitchblende and the natural occurrence of radiochlorine. J. Inorg. Nucl. Chem., 16:1.

Levine, C. A., and G. T. Seaborg, 1951 -The occurrence of plutonium in nature./ Am. Chem.
Soc., 73:3278.

Macnamara, J., and H. G. Thode, 1950-The isotopes of xenon and krypton in pitchblende
and the spontaneous fission of U238 Phys. Rev., 80:471.

Montgomery, C. G., and D. D. Montgomery, 1939— The intensity of neutrons of thermal
energy in the atmosphere at sea level. Phys. Rev., 56:10.

Peppard, D. F., M. H. Studier, M. V. Gergel, C. W. Mason, J. C. Sullivan, and J. F. Mech, 1951-
Isolation of microgram quantities of naturally occurring plutonium and examination of
its isotopic composition. J. Am. Chem. Soc., 73:2529.

Podosek, F. A., 1970-The abundance of 244Pu in the early solar system. Earth Planet. Sci.
Lett., 8:183.

Raut, M. K., and M. Attrep, Jr., 1975-(a, n) Reactions in uranium solutions. J. Inorg. Nucl.
Chem., 37:274.

Wetherill, G. W., 195 3 -Spontaneous fission yields from uranium and thorium. Phys. Rev.,
92:907.

Young, B. G., and H. G. Thode, 1960- Absolute yields of xenon and krypton isotopes in
U238 spontaneous fission. Can. J. Phys., 38:1.

MASS SPECTROMETRY STUDIES OF NORCAMPHOR AND NOR-
BORNYL ACETATE

by JAMES L. MARSHALL and STEVEN R. WALTER

Department of Chemistry,

North Texas State University,

Denton 76203

ABSTRACT

Six nor camphor ( 1 ) derivatives variously specifically deuterated were synthesized and
studied by mass spectrometry to ascertain fragmentation patterns. Extensive deuterium
scrambling occurs, attesting to facile hydride migrations, but the fragmentation pattern of
norcamphor closely parallels that of the trimethyl analog camphor. A brief look at 2-norbornyl
acetate ( 3 ) and a deuterio derivative indicates less scrambling occurs in this system than in
1 and that the ketone function actually promotes hydride migrations.

INTRODUCTION

The mass spectra of acyclic and monocyclic ketones have been studied"exten-
sively and commonly involve an initial a-cleavage with subsequent hydride shifts
and rearrangements (Budzikiewicz, et al., 1964). Studies for compact polycyclic
ketones are fewer, but an initial a-cleavage is indicated (Weinberg and Djerassi,
1966; Sydow, 1964; Dimmel and Wolinsky, 1967). In general, polycyclic com¬
pounds are capable of more complicated and less predictable reactions, and pre¬
vious studies of bicyclo [2.2.1 ] heptanones demonstrate this possible complexity
in such systems (Weinberg and Djerassi, 1966; Sydow, 1964; Dimmel and Wolinsky,
1967). However, these studies have dealt with methyl-substituted systems, and
it has not been clear to what extent these methyl substituents directed the re¬
arrangement process. Accordingly, in the present study, unsubstituted norcamphor
( 1 ) was studied. In this study, various deuterated norcamphor derivatives ( 1 b-f )
were synthesized and subjected to mass spectral analysis. Undeuterated norcamphor
itself ( la ) has been briefly analyzed by mass spectrometry (Goto, et al, 1966),
but no conclusions were reached other than it appeared that a CH3CO radical
was lost.

Received for publication: April 19, 1977.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

122

THE TEXAS JOURNAL OF SCIENCE

RESULTS AND INTERPRETATION

The mass spectral peaks for Ja-f are given graphically in Fig. 1. Mechanisms
are given below to account for these patterns and are then compared with those
proposed for camphor 2 (Weinberg and Djerassi, 1966; Dimmel and Wolinsky,
1967).

MASS SPECTROMETRY STUDIES

123

Figure 1. Mass spectra (relative abundance vs. m/e) for norcamphor ( fa ) and deuterio
derivatives lb-f\ The origins of peaks specifically identified are discussed in
the text.

In the first spectral region (m/e 81), norcamphor loses CO (but not ethylene
because the deuteriums are retained in lb) and a hydrogen to form the me thy 1-
cyclopentenyl cation 4; this mechanism is mirrored closely in the fragmentation
patterns for camphor (2), which gives as abase peak m/e 95, proposed (Weinberg
and Djerassi, 1966; Dimmel and Wolinsky, 1967) as 5. The mass spectral patterns
for Jj-f indicate that C-7 hydrogen is lost, consistent with the proposed mech¬
anism for camphor (Weinberg and Djerassi, 1966; Dimmel and Wolinsky, 1967)
that a C-7 methyl group is lost.

©

124

THE TEXAS JOURNAL OF SCIENCE

m/je 109

In the next spectral region ( m/e 66, 67) norcamphor is proposed to lose a
ketene to give 6, and then a hydride to give 7. Similarly, camphor loses ketene
and a methyl group to give 8, m/e 95. For camphor, it was shown (Dimmel and
Wolinsky, 1967) that the ketene arises from the C2-C3 linkage, and likewise the
patterns for Ja-f indicate the same type of loss for norcamphor (for example,
the 2 deuteriums are retained in lb). A remarkable similarity between camphor
(Weinberg and Djerassi, 1966, Dimmel and Wolinsky, 1967) and norcamphor is
the abstraction of a hydride by the C1-C2 a-cleavage intermediate in a 6,2-shift
process to give 9 and 10 respectively (Bartlett, 1965). The necessity for proposing
such a 6,2-shift for norcamphor is prompted because it appears that sometimes
C7 hydrogens are retained (see le,f ).

The next spectral region {m/e 54) corresponds to a C4H6 fragment. For cam¬
phor a precisely corresponding fragment does not exist, but there does exist a peak
at m/e 81 that corresponds to the retention of two methyl groups but the loss
of an extra hydrogen. Thus, it is not clear whether the correlation between the
fragmentation patterns of camphor and norcamphor extends into this region. The
spectral patterns for Ja-f indicate that the origin of this m/e 54 peak is the €3,
C4, C5, C7 assemblage.

Finally, the origin of the m/e 41 peak for norcamphor is proposed as below.
For camphor (2) this same process was proposed to give the fragment 13, which
through a series of hydride transfers and loss ofC3H7 gave 15, m/e 81 (Weinberg
and Djerassi, 1966; Dimmel and Wolinsky, 1967). In norcamphor, however, no
such C3H7 fragment could be lost; instead, a more direct sequence would be to
symmetrically cleave 11 as shown. If this were correct, then an analogous sequence
should be observed for camphor— 13 would cleave to give 14, with the positive
charge preferentially on the lower half, m/e 83. This peak is indeed observed for
camphor (Weinberg and Djerassi, 1966; Dimmel and Wolinsky, 1967;Sydow, 1964)
and precisely this mechanism has been proposed to account for its existence.

COMPARISON OF NORCAMPHOR WITH NO RBORNYL ACETATE

The norcamphor system 1 appears to exemplify that relatively simple terpenes
can undergo quite complicated fragmentations in their compact framework in
the absence of powerful directing groups. In fact, the carbonyl group seems to
complicate matters. To demonstrate this, the mass spectral data of mio2-norbornyl
acetate 3a and its exo,exo- 5,6-dideuterio derivative 3b (Figure 2) can be inter¬
preted rather extensively, as shown in the scheme below. The only indication of
scrambling is apparent at m/e 80 for 16; if 5x or 6x migrates and is lost, then
overall only one deuterium will be lost.

126

THE TEXAS JOURNAL OF SCIENCE

The fragmentation pattern given in the scheme on the following page is consistent
with mechanisms proposed for 1- and 2-methylnorbornenes (Reed, 1963) in that the
retro Diels-Alder reaction is not the principal method of fragmentation in norbornenes
(a retro Diels-Alder reaction for 3 would give m/e 66 directly with no fragments
m/e 67-69).- Apparently norbornene derivatives need abetter dienophile leaving
group; hence, norbornenyl acetate gives virtually quantitative retro Diels-Alder
reaction (Cristol, et al, 1966). A retro Diels-Alder mechanism has been proposed
to be general for norbornenes and benzonorbornenes, (Goto, et al, 1966) but
we suggest that the loss of two carbon fragments from these compounds is a step¬
wise process, as indicated in the scheme.

CONCLUSIONS

Although fragmentation patterns for small, compact molecules may be complex,
and although the ketonic functionality (at least in the present study) does not
promote clean fragmentation patterns, mass spectral analysis for norcamphor

MASS SPECTROMETRY STUDIES

127

m/e. 79 a m/e 79 a

SO b 81 |d

*Alternatively, H<jx migration and subsequent hydride shift will give the same overall effect.

may be done by means of various deuterated derivatives. The fragmentation pat¬
terns that appear to emerge from this analysis are quite analogous to those for
the trimethyl substituted analog camphor. Therefore, this study indicates that
fragmentation patterns for the bicyclo [2.2.1] heptan-2-one system remains rela¬
tively the same with or without methyl substitution and attests to the mechanistic
integrity of the carbon skeleton itself in mass spectral fragmentation patterns.

EXPERIMENTAL
Norcamphor Derivatives la

This synthesis of these derivatives has been previously described (Marshall
and Walter, 1974) and a very close proton nmr study (Marshall and Walter, 1974)
of lb-f proves the deuteriums are cleanly substituted with negligible admixture
of isomers or compounds with lesser amounts of deuterium. Mass spectral analysis
gave the following compositions: lb, 86.6% d2 , S.9%dl , 4.4% d0 ; lc, 88.2% d3,
7.8% d2 , 4.0% d ! , Id, 85.5% J4~~10.0% d3, 4.7% d2 ; le, 95% d~<~ 4% d5 ; If,
91% d5 , 9% <74.

128

THE TEXAS JOURNAL OF SCIENCE

Figure 2. Mass spectra (relative abundance vs. m/e ) for endo- 2-norbornyl acetate ( 31a,)
and deuterio derivative 31b. The origins of peaks specifically identified are
discussed in the text.

Norbornyl Acetates 3a, b

The synthesis of these derivatives is described in the same source as for the
norcamphor derivatives (Marshall and Walter, 1974). Owing to the very low in¬
tensity of the parent peaks for 3a, b, an isotopic composition was not determined;
however, 3b preceeds lb in the synthetic sequence (Marshall and Walter, 1974)
and hence must have an isotopic purity equal to or greater than that for lb
( > 86.6 %d2). Analysis of the norbornene peak(m/e 94) in 3a, b indicated Sl%d2 .

Mass Spectra

Mass spectra were run on a Hitachi-Perkin-Elmer RMU-6E mass spectrometer
at 70 ev at 2-3 x 10~6 torr.

ACKNOWLEDGEMENTS

The authors wish to acknowledge the Robert A. Welch Foundation, Grant
No. B-325, and North Texas State University Faculty Research for financial
support of this research. The assistance of Mr. Robert Owens in obtaining the
mass spectra of Ja-f is also gratefully acknowledged.

MASS SPECTROMETRY STUDIES

129

LITERATURE CITED

Bartlett, P. D., 1965 -Nonclassical Ions. W. A. Benjamin, Inc., New York.

Budzikiewicz, H., C. Djerassi, and D. H. Williams, 1961 -Mass Spectrometry of Organic
Compounds. Holden-Day, Inc., San Francisco, pp. 129-173.

- , - , and - — , 1964-Interpretations of Mass Spectra of Organic

Compounds. Holden-Day, Inc., San Francisco, pp. 17-22.

Cristol, S. S., T. C. Morrill, and R. A. Sanchez, 1966-/. Org. Chem., 31:2738.

Dimmel, D. R., and J. Wolinsky, 1967-/ Org. Chem., 32:410.

Goto, T., Y. Hata, R. Muneyuki, H.Tanida, A.Tatematsu,K.Tori, 1966-Tetrahedron, 22:2213.

Marshall,!. L., and S. R. Walter, 1974-/. Amer. Chem. Soc., 96:6358.

Reed, R. I., 1963-Mass spectra of terpenes. In F. W. McLafferty (ED.), Mass Spectrometry
of Organic Ions. Academic Press, New York, N.Y., pp. 655-658.

von Sydow, R., 1964 -Acta Chem. Scand., 18:1099.

Weinberg, D. S., and C. Djerassi, 1966-/ Org. Chem., 31:115.

'■

■

CHROMATOGRAPHIC STUDIES OF INTERACTION COEFFICIENTS
OF BENZOPHENONE AND BENZHYDROL WITH VARIOUS COM¬
POUNDS1

by REEVES B. PERRY and TERRY K. HENSON

Department of Chemistry ,

Southwest Texas State University,

San Marcos 78666

ABSTRACT

The interaction coefficients of benzophenone and benzhydrol with various test compounds
were determined. The stationary chromatographic columns were Fluoropak 80 supported
benzophenone, benzhydrol, and mixtures of the two compounds. The test compounds in¬
cluded acetic acid, allyl ether, 1-butanol, 1-decene, ethanol, heptanal, methyl acetate, and
pyrrole. The interaction coefficients were determined as compared to a hypothetical n-alkane
of the same molecular weight as the test compound. The interaction coefficients of column
mixtures tend to be weighted averages of the coefficients obtained with columns of the pure
compounds.

The results indicate that the interaction coefficients might be useful in the characteriza¬
tion of mixtures as to properties such as polarity. However, they do not promise to give
qualitative or quantitative data related to functional groups of a mixture.

INTRODUCTION

Retention behavior related to solute-solvent interactions in gas-liquid chroma¬
tography (GLC) depends on chemical properties of functional groups (Wehrli
and Kovats, 1959; James and Martin, 1954; and Heubner, 1962). Kovats (1958)
proposed a method of using retention indices for expressing qualitative GLC
values.

Equivalent chain-length numbers which are parameters similar to the retention
index have been used to characterize fatty acids (Miwa, et at, 1960). A combi¬
nation of 2 values, one from a polar and the other from a non-polar stationary
column, was sufficient for characterization.

Brown (1963) used retention volumes of 3 selected compounds to classify
stationary phases according to their polarity and behavior as electron donors or
acceptors. Brown suggested the elimination of hydrocarbon chain length to study
the behavior of functional groups.

1 This publication is a part of the M.A. thesis of Terry K. Henson, whose present address is
E. I. du Pont de Nemours and Company, Textile Fibers Division, Chattanooga, TN 38403.

Received for publication: March 31, 1977.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

132

THE TEXAS JOURNAL OF SCIENCE

Rohrschneider (1966) characterized stationary liquids by retention index
differences of 5 selected compounds. The method was based on additivity of inter-
molecular forces. Kaiser (1963) suggests that the greater retention times will result
from the greater sum of interacting forces.

Davis, et al, (1966) proposed the technique of inverse gas-liquid chromatography
(IGLC) to characterize asphalts. The primary interest is placed on the stationary
phase rather than the mobile phase as in conventional GLC. The asphalt was placed
on the support column and characterized by determining corrected retention
volumes of a series of selected test compounds of various functional groups. The
retention data were quantified by relating the retention volume of the test com¬
pound to the behavior of /7-alkanes of the same molecular weight. The retention
data were expressed as interaciton coefficients (7^) and were calculated using the
equation

Ip ~ [log (test compound) - log V°^ (hypothetical //-alkane)] x 100 (1)

where V°^ is the corrected retention volume.

Air is injected with the test compound, and the retention time (f^) is measured
from the appearance of the air peak to the appearance of the peak of the test
compound. V°^ is determined from and the flow rate (F):

V°r = tRF. (2)

The values of Ip depend on the interactions of test compounds with the func¬
tionality of the asphalt and were useful in showing differences among asphalts
but did not provide definite conclusions on the composition of the asphalts.

The correlation of the interaction coefficients of various functional groups of
pure compounds with the interaction coefficients of mixtures was studied for
benzhydrol, benzophenone and mixtures of these 2 compounds. These compounds
are similar except for the functional groups, and any differences in interaction
coefficients with a specific test compound would be due to the interaction with
the hydroxyl group of the benzhydrol or the carbonyl group of the benzophenone.
The study was to determine if qualitative and/or quantitative data could be derived
for functional groups in mixtures of organic compounds.

The vapor pressure of the stationary phase should not exceed 0.5 Torr at the
column temperature or the stationary phase would bleed from the column (Kaiser,
1963). Benzhydrol and benzophenone and mixtures of these 2 compounds
yielded a stable stationary phase for the column.

EXPERIMENTAL

The procedure was similar to that of Davis, et al, (1966). All retention data
were obtained on a F&M Model 700 gas chromatograph equipped with a thermal
conductivity detector and a Leeds and Northrup Speedomax G recorder.

INTERACTION COEFFICIENTS

133

Fluoropak 80 was used as the solid support, and the test compounds listed
in Table 1 were of ACS reagent grade or better. Benzhydrol, benzophepone, and
mixtures of these 2 compounds were used as the stationary phase. The total

TABLE 1

Interaction Coefficients of Various Systems

Test

Compounds

A

Interaction Coefficients (Ip)
BCD

E

Acetic Acid

163.1

164.5

176.5

187.1

191.7

Allyl Ether

45.5

49.1

52.0

57.7

60.3

1-Butanol

117.5

119.2

131.7

137.1

143.3

1-Decene

5.8

20.0

13.3

10.2

13.5

Ethanol

110.5

114.4

125.2

132.2

139.2

Heptanal

92.0

90.5

98.9

105.4

108.5

Methyl Acetate

56.9

67.7

71.0

77.2

82.5

Pyrrole

196.9

196.1

188.7

190.5

188.1

Stationary Phase (weight % supported on Fluoropak 80):

System A- 10.0% Benzophenone
System B - 7.5% Benzophenone - 2.5% Benzhydrol
System C - 5.0% Benzophenone -5.0% Benzhydrol
System D - 2.5% Benzophenone - 7.5% Benzhydrol
System E - 10.0% Benzhydrol

amount of the 2-component stationary phase was 10% (by weight) of the column
material (1 part stationary phase and 9 parts Fluoropak 80). The interaction
coefficients of 1 -component stationary phases did not show a large deviation
when the weight of stationary phase ranged from 5 to 15% of the column mass.
With the exception of 1-decene, the deviation was within 10% of the Ip at a sta¬
tionary phase of 10% of the column mass. Therefore, a stationary phase of 10%
of column mass was selected. The packing was prepared by making a homogeneous
slurry of Fluoropak 80 and the desired amount of stationary phase dissolved in a
volatile solvent. The slurry was spread out in a shallow pan and allowed to dry
overnight (@ 50 C). Four-ft columns were prepared from 1/4-in diameter aluminum
tubing. The column temperature was 100 C, and helium was the carrier gas. The
column was maintained at 100 C with continual helium flow for a minimum of
2 hrs before the injection of the test compounds. The volume of the test compound
was 0.5 jul.

V°g is obtained for a series of n-alkanes and the test compounds for each sta¬
tionary column. Ip is calculated by Equation (1).

RESULTS AND DISCUSSION

The test compounds which gave different interaction coefficients for benzo¬
phenone and benzhydrol are given in Table 1. The interaction coefficients are

134

THE TEXAS JOURNAL OF SCIENCE

listed for the stationary columns of pure compounds and those which are mixtures
of benzophenone and benzhydrol. Each column contains 90% inert support
(Fluoropak80) and 10% stationary phase. If the column contains 2% benzhydrol,
then it is 8% benzophenone. Test compounds used which gave the same inter¬
action coefficients are not included in Table 1. These compounds are benzene,
cyclohexane, methylcyclohexane , and toluene. The ^-alkanes which were used
as references are also omitted as these would automatically give interaction coef¬
ficients of zero.

The interaction coefficients of the mixtures tend to be weighted averages of
those of the pure stationary phases. This is shown for representative examples
in Figs. 1 and 2. The molecular weights of the 2 components of the stationary

Figure 1. Dependence of interaction coefficient {Ip) with the composition of the station¬
ary column. Legend of test compounds: ■ Ethanol, O 1-Butanol, •Heptanal.

INTERACTION COEFFICIENTS

135

Figure 2. Dependence of interaction coefficient (Ip) with the composition of the station¬
ary column. Legend of test compounds: • Allyl ether, O Methyl acetate.

phase are approximately equal (182.2 for benzophenone and 184.2 for benz-
hydrol). If the weight % of the stationary phase is 2.5% benzhydrol then the
mole fraction of benzhydrol in the stationary phase is 0.25, if 5.0% the mole
fraction is 0.5, etc. Therefore, the % by weight of one component of the column
is linearly proportional to its mole fraction of the stationary phase. The lines
drawn in Figs. 1 and 2 are the theoretical lines if the interaction coefficients were
linearly dependent on the composition of the stationary phase. The experimental
values of Ip are reproducible within 5%. The interaction coefficients of the 2-
component stationary phases seem to be more of a weighted average related to
the composition of the stationary phase rather than the sum of interacting forces
as might be expected from previous works of Rohrschneider (1966) and Kaiser
(1953).

These results indicate that interaction coefficients might be useful in the char¬
acterization of mixtures as to properties such as polarity. However, they do not
indicate that this is a method for obtaining qualitative or quantitative information
related to functional groups present in such complex mixtures as petroleum asphalts.

136

THE TEXAS JOURNAL OF SCIENCE

ACKNOWLEDGEMENT

This research was supported by Organized Research Funds of Southwest Texas
State University.

LITERATURE CITED

Brown, I., 1963-The role of the stationary phase in gas chromatography. J. Chromatog.,
10:284.

Davis, T. C., J. C. Petersen, and W. E. Haines, 1966-Inverse gas-liquid chromatography-A
new approach for studying petroleum asphalts. Anal. Chem., 38:241.

Henson, T. K., 1968-Inverse gas-liquid chromatography: An approach to studying functional
groups in organic compounds. M.A. thesis, Southwest Texas State University, San Marcos,
78666.

Heubner, V. R., 1962-Determination of the relative polarity of surface active agents by gas-
liquid chromatography. Anal. Chem., 34:488.

James, A. T., and A. J. P. Martin, 1954-Gas-Liquid chromatography-A technique for the
analysis and identification of volatile materials. Brit. Med. Bull, 10:170.

Kaiser, R., 1963 -Gas Phase Chromatography, Vol. 1. Butterworth, Washington, D.C., pp. 50, 53.

Kovats, E., 1958-Gas-Chromatographische charakterisierung organischer verbindungen, teil 1 :
Retentionsindices aliphatischer halogenide, akohole, aldehyde, and ketone. Helv. Chim.
Acta., 41:1915.

Miwa, T. K., K. L. Mikolajczak, F. R. Earle, and I. A. Wolff, 1960-Gas chromatographic
characterization of fatty acids. Anal. Chem., 32:1739.

Rohrschneider, L., 1966-Eine methode zur charakterisierung von gaschromatographischen
trennflussigkeiten. J. Chromatog., 22:6.

Wehrli, A., and E. Kovats, 1959-Gas-Chromatographische charakterisierung organischer
verbindunge. Helv. Chim. Acta., 42:2709.

THE TEMPERATURE OF MAXIMUM DENSITY OF DEUTERIUM
OXIDE IN CAPILLARIES1

by J. A. SCHUFLE and HSIEH-SUI HAO

Chemistry Department,

New Mexico Highlands University,

Las Vegas 8 7701

ABSTRACT

In earlier work we have described the discovery of a shift in the temperature of maximum
density (TMD) of water in fine capillaries.

We have applied the same technique to heavy water (deuterium oxide) and we find a
similar shift in the TMD from a value of 11.7 C for bulk D2O to 5.8 C for D2O in a quartz
capillary of a diameter of 2 micrometers (pm).

INTRODUCTION

The temperature of maximum density (TMD) of water has been shown to
change from the normal value of 4.0 C in bulk water to a temperature of -0.3 C
in a capillary 4 pm (Schufle and Venugopalan, 1967; Muller and Schufle, 1968;
Schufle and Huang, 1972). It was of interest to determine if a similar effect was
observed with deuterium oxide which exhibits a TMD at 1 1 .4 C ± 0.3 C.

Normal TMD for D20

The TMD for D20 is given in the Landolt-Bornstein (1950) Tables as 1 1.6 C.
However, when the same data was treated by our computerized, least-squares
method (Muller and Schufle, 1968) for determining TMD it produced a value of

1 1.1 C. We then consulted the original data of G. N. Lewis and R. T. MacDonald
(1933), who give a value for the TMD of “about 1 1.6 C.” The computer treat¬
ment of this data yielded a value for the TMD of 1 1 .4 C . Kell ( 1 967) gives a value
of 11.185 C. Finally, the value determined by the present authors in their own
apparatus described in Schufle and Venugopalan (1967) on D2 0 in a 0.5 mm diam¬
eter capillary was 11.7 C. We believe the best value may be somewhere around

11.2 to 11.4 C. Karasev, et al, (1971) have noted that D20 has a less pronounced
maximum than H20.

1 Paper delivered at meeting of Southwestern and Rocky Mountain Division of American
Association for the Advancement of Science, Laramie, Wyoming, 24 April 1974. Part of
the work was done by Mr. Hao as part of the requirements for the Master of Science degree
in Chemistry at New Mexico Highlands University.

Received for publication: January 2, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 1 and 2, September, 1977.

138

THE TEXAS JOURNAL OF SCIENCE

Determination of TMD in Capillaries

The variation of volume with temperature was measured for 99.90% D20. The
D20 was distilled in vacuo until a resistivity of 0.7 megohm-cm was attained.
The capillaries were filled with D20 and sealed immediately. Quartz capillaries
of diameters 500 pm, 20 pm, 10pm, 1 pm, and 2 pm were filled at the same time.
The capillaries were then introduced into the apparatus described in Schufle and
Venugopalan (1967) and the length of the column of liquid measured for temper¬
atures from approximately 40 C down to temperatures as low as -36 C for the
7 pm capillary. Liquid H2 O was carried to -40 C in a 4 pm capillary (Schufle and
Venugopalan, 1967), but with liquid D2 Owe were able to reach only -36 C before
freezing occurred spontaneously.

Xfhe following results (Table 1) were obtained, given in terms of the coefficients
of the polynomial of degree 3 of the form:

1/L = A(0) + A(l)t + A(2)t2 + A(3)t3 (1)

where L is the length of the water column, t is the temperature, and the TMD is the
temperature of maximum density obtained by the computer from the experimental
data.

TABLE 1

Coefficients for Equation (1).

Cap. Diam (pm)

500

20

10

7

2

10 A(0)

1.893385

2.145448

2.021029

2.094744

1.978843

10s A(l)

4.055660

3.832834

3.110212

4.917084

2.590978

106 A(2)

-1 .633188

-2.460285

-2.679592

-4.333409

-2.487299

108 A(3)

-0.568127

2.375391

3.454090

4.274537

3.082243

TMD

11.7 C

9.0 C

6.7 C

6.3 C

5.8 C

DISCUSSION OF RESULTS

The data show a continued shift of the TMD for D20 from the normal value
of about 1 1.4 C for '‘bulk” D20 down to 5.8 C for the smallest capillary with a
diameter of 2 pm. This corresponds to the results previously obtained for ordinary
H20 in Schufle and Venugopalan (1967), Muller and Schufle (1968) and Schufle
and Huang (1972).

The model we have proposed (Schufle, et al , 1976) is that of “vicinal water,”
a layer of water in the vicinity of the surface. We have found that the thickness
of the vicinal layer may be of the order of 1 pm thick. As the water droplet, or
the capillary containing water, becomes of smaller diameter the surface layer be¬
comes of relatively greater importance. When the diameter reaches the order of
a few micrometers the properties exhibited by the water are principally those of
the vicinal layer rather than those of bulk water. This is an alternative model for
explaining such anomalous behavior as that described by Derjaguin (1970).

THE TEXAS JOURNAL OF SCIENCE

139

In our model it is proposed that the TMD of vicinal water is at a lower temper¬
ature than that of ordinary water. The model seems to be true for D20 as well
asH20.

It is of interest to note that Karasev, et al , (1971) have shifted their emphasis
from their earlier model of anomalous bulk behavior to one of anomalous behavior
in small pores. For example they have found a shift in the TMD for H20 to a
temperature below OC, and for D20 to a temperature of 6 C. Derjaguiri’s abstract
of this article reads as follows:

“Water in bodies with small pores is found not to have a density maximum
at +4 C. Heavy water under the same conditions gives a less pronounced
maximum, which is shifted towards lower temperatures. Water released from
the pores is found to have the properties of ordinary bulk water.”

The results of this present work confirm these conclusions of Derjaguin with
respect to the shift in TMD for D20 from 11.4 C to about 6.0 C, for water in
small capillaries.

Derjaguin’s data give a TMD of 6.2 C for D20 in small pores when his data
are treated by our computer analysis (Muller and Schufle, 1968).

LITERATURE CITED

Derjaguin, Boris, 1970-Superdense water. Set Amer., 223:52.

Karasev, V. V., B. V. Derjaguin, and E. N. Khromova, 1911-In B. V. Derjaguin (Ed.), Re¬
search in Surface Forces, Vol. 3. Consultants Bureau, New York, N.Y., pp. 25-28. (English
Translation of a report of a conference held in Moscow, 1966, originally published in Russian
by Nauka Press, Moscow, 1967).

Kell, G. S., 1967 -Precise representation of volume properties of water at one atmosphere.
J. Chem. and Engineering Data, 12:66.

Landolt-Bornstein, 1950-ih A. Euken (Ed.), Zahlenwerke und Runctionen aus Physik,
Chemie, Astronomic, Geophysik und Technik, 6th Ed. Springer Verlag, Berlin, Band II,
Teill, p.457.

Lewis, G. N., and R. T. MacDonald, 1933-Some properties of pure H2 H2Q. J. Amer. Chem.
Soc 55:3057.

Muller, Ralph H., and J. A. Schufle, 1968-Shift in temperature of maximum density of
water in capillaries. / Geophys. Res., 73:3345.

Schufle, J. A., Chin-tsung Huang, and Walter Drost-Hansen, 1976-Temperature dependence
of surface conductance and a model of vicinal (interfacial) water. J. Coll, and Interface
Set, 54:184.

- , and Sun-yi Huang, 1972-Relative volume of water in quartz capillaries. Tex. J.

Set, 24:197.

- , and M. Venugopalan, 1967 -Specific volume of liquid water to ~40 C.J. Geophys.

Res., 72:3271.

Notes Section

141

EQUUS TAU OWEN FROM THE PLEISTOCENE OF MITCHELL COUNTY,
TEXAS — In the course of compiling records of the fossil remains of small horses found in
Pleistocene deposits in North America, I was impressed by the resemblance of the figure of
the lower jaw referred to Equus littoralis Hay by Hay and Cook (1930, Proc. Colorado Mus.
Nat. Hist., 9:4, Plate 2) to the lower jaw (FC 672), from Aguascalientes, Mexico, referred
to Equus tau (Mooser and Dalquest, 1975, J. Mamm., 56:781, Fig. 6). Dr. K. Don Lindsey
loaned me the fossil (Denver Museum of Natural History Number 616), found in the “bison
quarry” on Lone Wolf Creek, near Colorado, Texas (for details of this site and fauna see
Cook, 1925, Sci., 62:459; and 1926, Sci. Amer., 11:334). Direct comparison of the fossil
with the referred jaw of Equus tau, from Mexico, shows that both belong to the same species.

The rami of the 2 jaws are of the same size and shape. The pattern of the enamel in the
teeth of the specimens is similar except that the teeth of the specimen from Texas are more
worn, and the metastylids and metaconids appear more rounded. The slight differences in
measurements of the 2 specimens are less than the individual variation to be expected between
2 specimens of a single species of extinct horse.

The measurements that follow are in mm, and those of the Mexican horse are in parentheses.
Depth of ramus under P3, 58.3 (58.6). Length P2-M2 (M3 missing in the Texas fossil),
101.2 (100.5). Individual teeth, anteroposterior diameter followed by transverse diameter
(measurements of the Texas fossil are from Hay and Cook, loc. cit., p. 14): P2, 22 x 12.5
(24.7 x 11.4); P3, 20 x 13 (20.2 x 12.6); P4, 20 x 14(19.8 x 12.6); Ml, 18 x 12.5 (19.4 x
1 1.2); M2, 19 x 11.5(17.2x11.0)

The jaw from Mitchell County, Texas, is referable to Equus tau, and apparently constitutes
the first definite record of this species for the State of Texas.- Walter W. Dalquest, Depart¬
ment of Biology, Midwestern State University, Wichita Falls 7630 7.

THE STRUCTURE OF THE RETINA IN FOUR SPECIES OF WHISTLING
DUCKS , DEN DR OCYGNA - Four of the 8 species of whistling (or tree) ducks ,Dendro-
cygna, may be placed in 2 groups on the basis of their ecology. The black-bellied whistling
duck, D. autumnalis, and the plumed whistling duck,/). eytoni, are highly terrestrial; in con¬
trast the fulvous whistling duck, D bicolor, and the wandering whistling whistling duck,/).
arcuata, are highly aquatic. Previous studies have shown anatomical differences between the
2 groups correlated with the 2 niches (Rylander and Bolen, 1970, 1974). The feet and legs,
for example, are better adapted for walking in the terrestrial type, and the feeding apparatus
is more suited for straining in the aquatic type. What makes the study of these 4 species par¬
ticularly interesting is that one member of each ecological type, D. autumnalis and D. bicolor,
respectively, is found in North America, including Texas, and the second member of each
type is found in Australia. Convergent evolution of characters associated with the 2 niches
appears, therefore, to provide the best explanation for the anatomical similarities between
ecological counterparts that are so widely separated geographically; a correlation phenogram
of these relationships supports this contention (Bolen and Rylander, 1974).

Since the 2 cursorial species as well as the 2 diving species feed commonly at night, it
was hypothesized that the proportion of rods to cones in their retinas might be similar, thus
supporting the contention that the feeding niche is largely maintained by locomotor and
feeding features rather than by differences in visual abilities. A contrary situation was suggested
by a previous study that showed that the percentage of rods was greater in the black-bellied
whistling duck than in the mallard, Anas platyrhynchos, a predominately diurnal species
(Wells, et al, 1975); both are species not anatomically equipped for feeding in deep water.
This paper reports a histological study of the retinas of the 4 species of whistling ducks that
fall into the 2 ecological types described above.

One retina trom at least one individual of the 4 species was removed, fixed in 10% formalin,
embedded in parattin, sectioned at 15 microns, and stained with hematoxylin and eosin.
Rods and cones were identified on the basis of the location of their nuclei in relation to the

142

THE TEXAS JOURNAL OF SCIENCE

outer nuclear layer (Polyak, 1957). Twenty-five counts of rods and cones in each retina
were taken of sample areas (12 x 100 microns) measured with an ocular micrometer. The
sample areas were picked at random, but the fovea was avoided. The results indicate no sig¬
nificant differences in the proportion of the rods and cones between the 4 species (Table 1).

TABLE 1

Mean rod and cone counts from the retinas of 4 species of whistling ducks.

Species

No. Ducks

Examined3

Mb

%

Rods

Rods

Sd

Cones

Sd

D. autumnalis

3

33.9

3.36

17.0

1.14

65.4

D. bicolor

1

41.2

3.96

21.2

3.80

66.0

D. eytoni

1

39.4

5.47

20.6

2.38

65.6

D. arcuata

2

44.6

8.08

19.0

0.71

70.0

aCounts from 25 randomly selected areas from each duck’s retina.
bRods and cones per square millimeter = M x 104.

We therefore conclude that there are no retinal differences reflecting differences in visual
abilities between the cursorial and diving types of whistling ducks, and that sight is not an
important feature contributing to the maintenance of the respective niches of these 2 eco¬
logical types.

ACKNOWLEDGEMENT

This research was supported in part by the Welder Wildlife Foundation.

LITERATURE CITED

Bolen, E. G., and M. K. Rylander, 1974-Foot adaptations in four species of whistling
duck Dendrocygna. Wildfowl, 25:81.

Polyak, S., 1951 -The Vertebrate Visual System, University of Chicago Press, Chicago.

Rylander, M. K., and E. G. Bolen, 19 70- Ecological and anatomical adaptations of North
American tree ducks. Auk, 87:72.

— - — , and - , 19 74- Feeding adaptations in whistling ducks (. Dendrocygna ).

Auk, 91 :86.

Wells, M. C., P. N. Lehner, E. G. Bolen, and M. K. Rylander, 1975-Comparison of scotopic
sensitivity in diurnal ( Anas platyrhynchos) and crepuscular ( Dendrocygna autumnalis )
ducks./ Comp, and Physiol Psych., 88:940.

- Stephen M. Womack, Michael K. Rylander, Dept, of Biological Sciences, Texas Tech Uni¬
versity, Lubbock 79409, and Eric G. Bolen, Rob and Bessie Welder Wildlife Foundation,
Sinton 78387.

EXECUTIVE COUNCIL

President:
President-Elect :

Vice President:

Im mediate Past President:
Secretary- Treasurer:

JAMES R. UNDERWOOD, JR., West Texas State University
J. D. McCULLOUGH, Stephen F. Austin State University
J. L. POIROT, North Texas State University
HERBERT H. HANNAN, Southwest Texas State University
EVERETT D. WILSON, Sam Houston State University

Sectional Chairpersons:

I -Mathematical Sciences: WILLIAM D. CLARK, Stephen F. Austin State University

II -Physical and Space Sciences: ROBERT W. GRUEBEL, Stephen F. Austin State Univ.

III -Earth Sciences: JAMES B. STEVENS, Lamar University

IV -Biological Sciences: RICHARD H. RICHARDSON, University of Texas at Austin

V -Social Sciences: RAYMOND TESKE, JR., Sam Houston State University

VI -Environmental Sciences: ELRAY S. NIXON, Stephen F. Austin State University

VII -Chemistry Section: BILLY J. YAGER, Southwest Texas State University

VIII -Science Education: JOEL E. BASS, Sam Houston State University

IX -Computer Sciences: GRADY G. EARLY, Southwest Texas State University

X -Aquatic Sciences: WILLIAM J. CLARK, Texas A&M University

XI -Forensic Sciences: RAYMOND WM. MIRES, Texas Tech University

Manuscript Editor: G. ROLAND VELA, North Texas State University
Managing Editor: MICHAEL J. CARLO, Angelo State University

Chairperson, Board of Science Education: PAUL J. COWAN, North Texas State University
Collegiate Academy: ROBERT V. BLYSTONE, Trinity University
Junior Academy: FANNIE M. HURST, Baylor University

BOARD OF DIRECTORS

JAMES R. UNDERWOOD, JR., West Texas State University
J. D. McCULLOUGH, Stephen F. Austin State University
HERBERT H. HANNAN, Southwest Texas State University
EVERETT D. WILSON, Sam Houston State University
G. ROLAND VELA, North Texas State University
MICHAEL J. CARLO, Angelo State University
ARTHUR E. HUGHES, Sam Houston State University
WILLIAM K. DAVIS, Southwest Texas State University
ROBERT BOYER, University Station
JAMES D. LONG, Sam Houston State University
JOHN W. FITCH, Southwest Texas State University
LAMAR JOHANSON, Tarleton State University
FREDERICK GEHLBACH, Baylor University

COVER PHOTO

(Top) Lateral view of the head of an adult male Elaphe ob sole ta (WMM 650)
from south of Kerrville, Kerr County, Texas. (Bottom) Lateral view of the
head of an adult male Elaphe bairdi (REO 687) from near Alpine, Brewster
County, Texas.

by R. Earl Olson, pp. 79-84.

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Manuscript Editor, P.O.Box 13066, North Texas State University, Denton, Texas 76203.

Published quarterly by The Talley Press, San Angelo, Texas, U.S.A. (Second Class
Postage paid at Post Office, San Angelo, Texas 76901.) Please send 3579 and returned
copies to the Editor (P.O.Box 10979, Angelo State University, San Angelo, Texas 76901.)

Volume XXIX, Nos. 3-4

December, 1977

CONTENTS

Instructions to Authors ..................................... 144

Loss of Efficiency Due to Estimated Weights in the Recovery of Inter-Block

Information. By Luisa L. Sia and Anant M. Kshirsagar. . . 147

On the State of Saturation of Groundwater with Respect to Dissolved Carbonates,

Edwards Artesian Aquifer, South-Central Texas. By Patrick L. Abbott . ....... 159

Fitting Soil Temperature by a Periodic Regression Model. By Mingteh Chang

and Douglas G. Boyer . . . 169

Trace Fossils from the Pecan Gap Formation (Upper Cretaceous), Northeast

Texas. By William C. Dawson, Donald F. Reaser, and Jimmy D. Richardson . 175

Preliminary Field Study of the Fracture Patterns Associated with the Balcones
Fault Zone in North Central Texas. By R.G. Font, J.C. Y elder man, C.T.

Hayward, and E.E. Baldwin . . 187

Late Pleistocene Pollen and Sediments: An Analysis of a Central California

Locality. By Eric W. Ritter and Brian W. Hatoff . . . . . . 195

Vegetation Types of Chambers County, Texas. By P.A. Harcombe and J.E. Neaville. . . . 209

Geomyid Interaction in Burrow Systems. By Graham C. Hickman . . . 235

Mitotic Chromosomes of Turtles. IV. The Emydidae. By Flavius C. Killebrew . 245

Effects of Hypophysectomy in the Lizard Holbrookia Propinqua. By Jon T. Watson . . . 255

Cretaceous (Albian) Ammonites from Puerto Rico and St. Thomas. By Keith Young . . . 263

Ideal Gas Thermodynamic Functions of the 2-Halopropenes. By G.A. Crowder

and Royce W. Waltrip . . . . 279

A Protostegid (Sea Turtle) from the Taylor Formation of Texas. By L.W. Osten ..... 289

Choice of Minicomputers’ Floating Point Base. By James L. Poirot ............ 293

NOTES SECTION

First South Texas Records of Pappogeomys Castanops. By Arthur G. Cleveland . .... 299

The Level of Understanding of Algebra and Trigonometry by Students ofFresh-

man Physics. By H.T. Hudson and RM. Rottmann . ................... 299

The Occurrence of Argulus Bicolor Bere, 1936 on Trachinotus Carolinus (Linnaeus)

from the Texas Coast. By Timothy L. Jones and Ronald C. Circe . . 301

INDEX Volume XXIX - 1977 . . . . 304

INSTRUCTIONS TO AUTHORS

Papers intended for publication in The Texas Journal of Science are to be sub¬
mitted to Dr. Roland Vela, Editor, P. 0. Box 13066, North Texas State University,
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Academic Press, New York, N.Y., pp. 13-29. Fora CHAPTER IN A BOOK WITH
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and T. S. Gibbs (Eds.), Effects of Chlorinated Hydrocarbons. Academic Press,
New York, N.Y.,pp. 3-12. For an IN PRESS PERIODICAL reference, use: Jones, A.P.,
1971— Effects of chlorinated hydrocarbons./, of Org. Chem., In Press. For an
IN PRESS BOOK reference, use: Jones, A. P., 1 97 1 — Effects of Chlorinated Hydro¬
carbons. Academic Press, New York, N.Y. In Press.

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145

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WITHOUT LOSS OR DELAY.

LOSS OF EFFICIENCY DUE TO ESTIMATED WEIGHTS IN THE
RECOVERY OF INTER-BLOCK INFORMATION

By LUISA L.SIA

Waller Hall

Rutgers University

New Brunswick, New Jersey 08 9 03

and AN ANT M. KSHIRSAGAR

Department Bio-Statistics
School of Public Health
University of Michigan
Ann Arbor 48109

ABSTRACT

In the recovery of inter-block information the ratio p of the inter-block variance to the
intra-block variance is used as weight, p being unknown, the ratio of the corresponding
variance estimates obtained from the analysis of variance is usually used. It is, however, not
unbiased for p, and often results in biased estimators of treatment effects. Roy and Shah
(1962) suggested several alternative estimators of p which yielded unbiased estimators of
treatment effects. In this paper, the loss of efficiency in the estimation of treatment ef¬
fects due to sampling error of the weight estimators are calculated using Rao’s (1965) delta
technique. The same technique is also used for the estimated weights proposed by Kshir-
sagar (1971) in the recovery of inter-row and inter-column information of a two-way
design.

INTRODUCTION

Let N be the vxb incidence matrix of an experimental design involving v treat¬
ments, b blocks of k plots each and r replications. The elements ny(i=l, 2,...v,
j=l,2,...,b) of N are 1 or 0 according as the i-th treatment does or does not occur
in the j-th block. Assume the design to be connected, then rank C=v-1 , where
C=rlv - -I- NN;. This implies that there is exactly one eigenvalue of NN; which is
equal to rk, and all other eigenvalues are strictly smaller than rk. Suppose rank
NN' =q + 1, and let s=l,2,...,q, be a set of orthonormal eigenvectors of NN*
corresponding to the q positive eigenvalues 0S all smaller than rk, and let j^,
s=q + l,q + 2,...,v-l, be a set of (v-l)-q orthonormal vectors, each orthogonal
to and also to EVl . It is well known that the intra-block, inter¬

block, and combined intra- and inter-block estimates of any treatment contract

Accepted for publication: January 28, 1977.

The Texas Journal of Science, Vol. XXIX, Nos. 3-4, December, 1977.

148

THE TEXAS JOURNAL OF SCIENCE

ji are respectively x £_' £_*, and x where xx* and Tare any solutions
of the following equations:

Cx - Q. (1)

NN' r_* = NB - t-GEvl (2)

(C+^NN’)T = a + f(NB - tGEvl) (3)

where

P =

a2 + ka£

(4)

QL = T -

(5)

and T, B, G denote the vector of treatment totals, the vector of block totals,
and the grand total of a 11 the yields respectively, o 2 is the intra-block vari¬
ance/plot, and is the variance of the random block effects.

When p is known, the minimum variance unbiased estimator of an eigenvector
contrast is given by

ts(p) =i-'s£- for s = 1 , 2,...,q, (6)

hence the estimator is a function of p. For s = q + 1, q + 2,..., v - 1, T_
coincides with JlsX and ts=l_,sr_doesnot depend on p. (Kshirsagar and Sia,1974).
p being usually unknown, an estimate P is often substituted for p in (3), and the
resulting estimator is denoted by ts(P). Roy and Shah (1962) proposed esti¬
mators of the form

q

aSi + 2 bsz2

P= - X1 - +c (7)

where S0 is the residual sums of squares from the analysis of variance,

%-v^2i.-*XF-s’NX (8)

q 2

Sj = adjusted block S.S. - 2 _$s^s_

s=l rk

(9)

RECOVERY OF INTER-BLOCK INFORMATION

149

and a, c, bs are constants to be determined for each P of specified optimum
property. The increase in the sampling error of the treatment effects due to
sampling fluctuations of P was shown to be

Var[ts(P)] = Var[ts(p)] + dsVar(u>s) for s = l,2,...,q, (10)

where

ds -

k

0S(P + Cs5)

(11)

cos

(P-P)z*

1 + CSP

(12)

rk-0c
Cc = VS

b 0S

(13)

S0 and S! are respectively distributed as y2 o2 and x2 o\ with corresponding
degrees of freedom

c0 = b(k-l) - (v— 1 )

(14)

O

il

'a*

1

'Y

hQ

(15)

zs ~ NID(0,Us), o l = o2 + cs(7i , for s = 1 , 2V

-»q- (16)

S0, Si and zs, s = 1,2,. ..,q, are all independent.

LOSS OF EFFICIENCY

Since the variance increment Var(cos) has not been given in an explicit form,
the loss of efficiency due to P cannot be computed. An approximation is ob¬
tained in this paper using the delta technique (Rao, 1965)-.

For algebraic convenience P can be written as

p cSn+ aSi + u + b<z?

P" S0

(17)

where

u = 2 bjZ? — 2'bjZj
rFs

(18)

(18)

150

THE TEXAS JOURNAL OF SCIENCE

and cos = 'I' (Si , S0, u, zs), a function of Si , S0, u and zs. Using the delta tech¬
niques, first take the total differential of cos,

dws = ~:-dS1 +^-dS0 + |^-du +^-dzs (19)
oS i oS0 ou ozs

where dSt stands for Si - E(Si), and d— is evaluated at the expected values of

9Si

Si, SQ, u and zs, etc. It is noted that for the proposed P, E(cos) = 0. Squaring
and taking the expected value yields

+ 0^)2 Var(u) +0f^) 2 Var^’ (2°)

since all the covariances vanished. We have

Var(<os)= 0^ (o2 +csa?) (21)

where

Vi = (c-p) e0 + apej + S'bi(l + qp) (22)

V2 = (1 + ccs)e0 + apcse! + csS'bi. (23)

Among the estimators of p proposed by Roy and Shah (1962), we consider
3 which will be denoted by P! , P2 and P3 below.

1. To make the classical estimator of p unbiased, take

k(e0-2)

(24)

v(r-l)

= -(v-k)

(25)

v(r-l)

(eo-2)0s s=i 2,...,q.

(26)

bk(r-l)

P now takes the form

(27)

RECOVERY OF INTER-BLOCK INFORMATION

151

where R is the classical estimator of p. ?i is unbiased for p, and the resulting
treatment contrast estimator ts (Pi) is also unbiased for ts(p) = iL'sll. Applying
(21), the loss of efficiency due to substitution of P: for p is dsVpl(cos), where

vp, fa2 +csai) (28)

v„ = [r(k-v)eo + (eo-2)2V>i]0s+ {vr(r-l)e0 + (e0-2) [rk(b-2)-Z'0i]0sP (29)

V21 = rke0[r(k-v) + (b-l)0s] + (e0-2) (rk-<^s)Z'0j
+ (ec-2) (rk-0s) [rk(b-2)-2'0j] p. (30)

2. The variance of the unbiased estimator P of p is a quadratic function in p.
When p is large, the term involving p2 in Var(P)is dominant (Roy and Shah, 1962).
The constants can thus be chosen to make the coefficient of p2 in Var(P) a min¬
imum, giving

, _ 3(eQ-2)

(31)

3ei +(e, + 2)q

O

II

)

M.O

c r

(32)

(e0-2) s=l

u _ (ei +2)a
bs " 3cs

(33)

P now takes the form

1

<N

1

O

<L>

1 _

3S[ + (e[ + 2)E ( ^ ) /z?+ S° )

3et + (e ! + 2)q _

L kk-*J v e^vJ

W

P2 is unbiased for p. Applying (21), the loss of efficiency due to substitution of
P2 for p is ds • Vp2 (cos), where

VP2 (ws) =? (a2 + csa^ ) (34)

V21 = 2(e0 + 2)2 cl1 + (e0e! +4ej +4q + 26^)136! +(ei +2)q]
a=l

+ [4(e1-l) + (e1 +2)(ec + 2q)]p (35)

152

THE TEXAS JOURNAL OF SCIENCE

V22 =60(36! + (e, +2)q]-2(e, +2)05.^ cj1 - (e0-2) (e, +2)
+ 3(e0-2)cs[(e, +2)q-2(e,-l)]p. (36)

3. Another unbiased estimator of p proposed by Roy and Shah is of the form

P3 =(1 (1-1) (37)

eo v0 e0(b-l) E

where

E = (v-1) / + 2 1

rk ' L rk s=l rk-&

(38)

is Kempthorne’s efficiency factor.

= So

and

(39)

C Q

vx = _ 2

e0(b-l)s= 1

1+_L_

cs (b— 1)

q

Si +2

S= 1

(40)

are unbiased estimates of and o\ respectively; and they are optimum in the
sense that the term involving p2 in their respective variances are minimized.
Shah (1962) suggested these estimates from intuitive considerations. Applying
(21), the loss of efficiency due to substitution of P3 for p is ds • Vp3 (cos), where

VP3(w5)=02i_y (o2+csa?) (41)

V3 1 = 2(e0-3)2 cl1 - (e0-2>i1 - (b- 1) (e0 + 2(b-2))p (42)

1=1

V32 = (b-2)e0 + 2(e0-3)cs c"1 + 2-(b-2)(e0-2)csp (43)

Since Px , P2 , and P3 all give unbiased combined inter- and intra- block esti¬
mations of an eigenvector contrast; i.e., ts(Px), ts(P2), and ts(P3) are all unbiased
for L'sr, the choice among the 3 can therefore be guided by the computed loss
of efficiency for a particular design.

RECOVERY OF INTER-BLOCK INFORMATION

153

APPLICATION OF THESE RESULTS TO SOME DESIGNS

In calculating the loss of efficiency using equations (24), (27) and (29), the
parameters needed are q, e0, ex , 0S, Z'0i> cs, and These are given for 3 de¬

signs in this section.

1) BIBD (b, v, r, k, X)

q = v-1

(44)

e0 = bk - b - v + 1

Qi = b-v

(45)

0s = r.X = Kv-k)

V-1

s= 1, 2,...,v-l

(46)

S'0i=?0i = (v-2)(r-X)=r(v-2)fv-k)
i=Fs v-1

(47)

r. = rk -0S _ v(k-l)
0s v-k

s = 1, 2,...,v-l

(48)

£ 1 = (v-1) (v-k)
i=l cj v(k-l)

(49)

2) GDD (b, v, r, k, Xi , X2 , m, n)

We consider here a group divisible PBIBD with two associate classes, where
the v treatments are divisible into n groups of m treatments each.

v = mn

(50)

q = mn- 1

(51)

e0 = bk-b-mn+ 1

(52)

e1 = b-mn

(53)

^ = { rk-mnXa

for s = 1 , 2,...,m(n-l)
for s = m(n-l)+l,...,mn-l.

(54)

If s < m(n-l)

2'0i = (mn-2)r + [m(m-2)(n-l)+ l]X1-(m-l)(mn-m+l)X2 (55)

154

THE TEXAS JOURNAL OF SCIENCE

r = r(k-l)-Xj
r-Xi

2 1 =(mn-l)(r-X1)
i""1 q r(k-l)-Xi

(56)

(57)

If m(n-l) < s < mn

2*01 = (mn-2)r+m(m-3)(n-l)X1- [m(m-2)(n- l)+2(m-l)] X2
P = mnX2

Cg . — - - — -

rk-mnX2

(58)

(59)

2 1 mn(mn-l)X2

i= 1 q rk-mnX2

(60)

3) Confounded sm Factorial Experiment

Consider a factorial experiment in which there are m factors each at s levels
(s being the power of a prime). Assume that rank C = v-1, and that out of the
(sm-l)/(s-l) pencils, nu pencils are confounded in r-u replications and uncon¬
founded in u replications, u = 1, 2,...,r. Then the eigenvalues of C are Ty = i,
[j = l, 2,...,nj(s-l); i = 1, 2,...,r] , (Kshirsagar and Sia, 1974). Therefore the
eigenvalues of NNf are

0ij = k(r-i) j= l,2,...,ni(s-l) (61)

i= 1, 2,...,r.

i-1

Note that the single subscript notation 0S = 0y may be used with s = 2 nj(s-l)+j;

u=l

however, a double subscript is more convenient in this case.)

In computing the approximate value of Var(copq), the following results are
needed:

q = sm~l

(62)

e0 = bk-b~sm + 1

(63)

Qy = b“Sm

(64)

2 V 2 2 0*1=^ (k2-k+l)-rk-p

i=Fp j^q k

(65)

c - P
pq "TF

(66)

RECOVERY OF INTER-BLOCK INFORMATION

155

2 ni^_1) J_= l-sm+r(s-l) 2 (67)

i=l j=l qj i

EXTENSION TO TWO-WAY DESIGNS

As in one-way designs the precision of the estimates of treatment effects in
two-way designs can be increased by the use of information available from

inter- row and inter- column comparisons. The ratio pr

a2 + bo2

of the

inter-row variance to the intra-row and column variance, and the ratio pc = °Z..+^°C .

o 2

of the inter- column variance to the intra-row and column variance play im¬
portant roles in the combined inter- and intra- estimations of treatment effects.
However pr and pc are usually unknown, and the usual estimates of them ob¬
tained from an analysis of variance of the data are biased; as a result the final
estimates of the treatment effects are, in general, no longer unbiased. Kshir-
sagar (1971) extended the results of Roy and Shah (1962) for a particular case
of the general two-way design; namely, the case in which the C -matrices of the
row and column designs have the same eigenvectors. In the following discussions
we shall use the same notations used by Kshirsagar in order to make the present¬
ation and cross-referencing easy.

Let F be the C-matrix of the design, L and M be the row and column inci¬
dence matrices respectively. It is assumed that rank LL'= % + 1 and rank MM' = qc + 1 .
Rank F = v-1 since the design is connected. Denote the common eigenvectors of

LL/ and MM' to be -L=Evi
VT

iLi ,... jLv-i • Then the combined inter- and intra¬

estimators of an eigenvector contrast is a function of pr and pc; namely,

itl*(Pr>Pc) =

^Q-VJL'sQr + 7ciL'sQ(

(68)

0S

i-k?

uPc

gs

where u = number of rows, u = number of columns, 0§, es, and gs are the respec¬
tive eigenvalues of F, LL' and MM' corresponding to ls; Q, Qr, and Qc are the
respective vectors of adjusted treatment totals in the intra-row and column,
inter-row, and inter-column estimations.

Kshirsagar (1971) proposed estimators of the weights pr and pc with the
forms

qr

aEr + 2 bsco2

(69)

156

THE TEXAS JOURNAL OF SCIENCE

qc 0
h Ec + 2 dsXs

Pc= _ £ _ + f (70)

Ej

where

ws = l_asQ)-^-(i'sQr), (7i)

0s es

Xs= L_ Oi^-iLOiOc), (72)

0s gs

and the inter-row, inter-column and combined intra-row and column error S.S.
are respectively:

Ej ~ X2 o2

with v = n-v-u-u' + 2 d.f.,

(73)

Er ~ X2(u2 + u'a2)

with vx - u-qr-l d.f.

(74)

Ec ~ X2 (a2 +ua2)

with uc - u'-qc-l d.f.;

(75)

a, c, h, f, bs and ds are constants to be determined.

Kshirsagar showed that when Pr and Pc are substituted for the unknown
weights pr and pc in l/si^Pr* Pc)> the resulting estimator i.^(Pr, Pc) is unbiased
for the treatment contrast isL_; moreover,

Var[l'stJ;(Pr, Pc)] = Var[&*(pr, pc)] + Var(zs) (76)

where

zs =hi - if)w, j- - r ) (^S-Xs)

U rr uu pc rr

+ M_(i- i )Xs-^s_( i- ix«5-x.) (77)

u Pc Pc uu pr Pc Ec

Thus, the effect of substituting Pr and Pc for pr and pc is to increase the variance
oflsl?(pr, pc) by Var(zs).

When the delta technique is applied to zs, an approximation to its variance is
found to be

Var(zs) = [A2(JL + 3L£l) + B2 ( -J- + + 2AB( -L)] a2

0s es 0s 8s 0s

(78)

RECOVERY OF INTER-BLOCK INFORMATION

157

where

a=tM| 7-m^s+iw) + ni'(|57)1

(79)

(80)

i / §r

m = [a i^Pr + ov + (—-+ u Pl ) .2 fy] _1

0s es

(81)

n = [h vcpc + fv + ( J— + uPc ) .? dj] _I

0s gs i=Fs

(82)

LITERATURE CITED

Kshirsagar, A.M., 1971 -Recovery of inter-row and inter-column information in two-way
designs. Ann. Inst. Statist. Math. 23:263.

- , and L.L. Sia, 1974-C-matrix of confounded designs in symmetrical factorial

experiments. Communications in Statistics, 3(11): 1015.

Rao, C.R., 1965 -Linear Statistical Inference and Its Applications. John Wiley and Sons,
New York.

Roy, J., and K.R. Shah, 1962-Recovery of interblock information. Sankhya, A, 24:269.

Shah, K.R., 1962-An alternative estimate of interblock variance in incomplete block
designs. Sankhya, A, 24:281.

ON THE STATE OF SATURATION OF GROUNDWATER WITH
RESPECT TO DISSOLVED CARBONATES , EDWARDS ARTESIAN
AQUIFER, SOUTH-CENTRAL TEXAS

by PATRICK L. ABBOTT

Department of Geological Sciences, San Diego State University,
San Diego, California 92182

ABSTRACT

Groundwater flowing through the Edwards Limestone in the Balcones fault zone has cre¬
ated extensive cavern systems. Much of the groundwater in the Edwards artesian aquifer has
traveled distances exceeding 100 mi. Long distances of flow seemingly should result in water
saturated or supersaturated with respect to calcite and dolomite and hence unable to further
dissolve host rock. Thus cavern development should be most pronounced near the recharge
sites where water is likely to be most undersaturated. Yet caverns in the Edwards aquifer are
most extensive near the distal, or discharge-dominated, end of the flow system. The well
developed caverns are the result of the general undersaturation of Edwards artesian aquifer
water. At least seasonal undersaturation is maintained by introduction of relatively small
volumes of flood recharge in Medina, Bexar and Comal Counties to the large groundwater
body.

INTRODUCTION

Groundwater in carbonate aquifers generally flows through void systems that
have been created largely by the solvent action of the water on the host rock. In
the Balcones fault zone of south-central Texas an extensive system of caverns
has been developed in the Lower Cretaceous Edwards Limestone and to a lesser
extent in the subjacent Walnut and Glen Rose formations. To evaluate whether
cavern development is occurring at present, the state of saturation of the ground-
water must be known with respect to the common minerals of older carbonate
rocks- -calcite and dolomite. If the water emerging at springs is undersaturated
then it is aggressive enough to dissolve the rocks it has passed through. If the
water is supersaturated at the springs, and for considerable distance into the
aquifer from the springs, then dissolution is probably concentrated nearer the
recharge areas where the water is still undersaturated.

The purpose of this study was to calculate the saturation of aquifer water
samples with respect to calcite and dolomite and then examine their areal trends

Accepted for publication: September 17, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 3-4, December, 1977.

160

THE TEXAS JOURNAL OF SCIENCE

to see if any tentative hypotheses about aquifer development could be made.
The water samples examined as to their state of saturation were drawn from the
wells and springs whose locations are plotted in Fig. 1 with respect to the bound¬
aries of the Edwards artesian aquifer. Groundwater in the aquifer beneath Bexar
and Comal Counties moves dominantly to the northeast into Hays County. For
example, groundwater near well 804 in southwestern Bexar County travels toward
San Marcos Springs in Hays County. The straight-line distance between these
points, which is not to be taken as the actual path of the water, is about 68 mi.
Along the way some of the excess water spills out at San Pedro, San Antonio
and Comal Springs. The majority of the groundwater in the Edwards aquifer seg¬
ment shown in Fig. 1 does not come from the local rivers and creeks depicted in
Fig. 1 . Instead it is derived as underflow within the aquifer from as far as the
groundwater divide near Brackettville in Kinney County which is a straight-line
distance of about 95 mi to the west of well 804 in Bexar County (Garza, 1962;
Arnow, 1963).

GEOCHEMICAL EVIDENCE ON WATER SATURATION

The key evidence necessary to evaluate the interaction between groundwater
and carbonate host rock is the state of saturation of groundwater with respect
to calcite and dolomite.

Method of Calculating States of Saturation

Unpublished chemical analyses of spring waters in Comal, Bexar and Hays
Counties and of well waters in Bexar County were used in this study; all had pH,
bicarbonate concentration, temperature, and sometimes specific conductance de¬
termined in the field. The water samples selected for analysis all had a close bal¬
ance of charges between anions and cations. Ionic strengths (I) were calculated
both by taking half the sum of the molalities multiplied by the ion charge squared
and by the empirical relationship of Jacobson and Langmuir (1970) where I
equals 1.88 x 10”5 times the specific conductance. Both methods of calculating
ionic strength yielded the same activity coefficients from Figure 4.5 of Garrels
and Christ (1965).

The method of calculating the state of saturation of water with respect to cal¬
cite and dolomite followed the procedure in case 3, chap 3, (Garrels and Christ, 1965).
Solubility constants for calcite and dolomite were adjusted for water temperature
according to the values listed in Table 3.2 (Garrels and Christ, 1965) and Table 1
(Thrailkill, 1972), respectively.

The observed and calculated molalities of calcium and magnesium were com¬
pared using the calcite saturation index (Sip;) and the dolomite saturation index
(SIp>). Sip; and Sip) are defined as:

_ [Ca++] [C03— ] observed
C ~ ^Calcite calculated

SATURATION OF GROUNDWATER

161

Figure 1. Map of major hydrologic features in Bexar, Comal, and parts of Medina and
Hays Counties. Well location numbers are the last 3 digits of the identification
numbers in Table 1. Base map for this figure was adapted from Alexander,
etal., (1964).

162

THE TEXAS JOURNAL OF SCIENCE

ciD = LCa++l fMg++l [C03~~] 2 observed
^Dolomite calculated

Negative values of SI^ and SIj) indicate the water samples are undersaturated
and positive values show the water is supersaturated.

The saturation indices listed in Table 1 slightly overestimate the water satura¬
tion because the molalities of ion pairs were not subtracted. Although the pre¬
ponderance of water samples in this study were undersaturated, the application
of ion-pairing corrections would lower the level of saturation slightly further.

Results of Water Saturation Calculations

Examination of the data in Table 1 shows than no entirely consistent general¬
izations are possible. One area where care must be exercised is in comparing sat¬
uration indices of waters drawn from the aquifer on different days, months, and
years. Seasonal variations in the state of saturation of groundwater in carbonate
terranes have been demonstrated in Pennsylvania by Shuster and White (1971)
and in Kentucky by Thrailkill (1972). Thus the only direct comparisons that can
be made are between water samples collected at the same time. Notice, for in¬
stance, that the least saturated samples were all collected in mid-December 1974
and that most of the supersaturated samples were collected in late March, 1972.
It would be most instructive to be able to examine the chemical analyses of a
large number of samples extracted from throughout the aquifer in as short a
time as possible.

Despite the inadequacies of the presently available data, which are from an
aquifer that has non-homogeneous porosity development and anisotropic ground-
water flow, some significant hypotheses may be suggested with caution. In¬
spection of the chemical analyses listed in Table 1 shows that the groundwater
and spring water from the Edwards aquifer are predominately undersaturated.
Several additional imperfect generalizations may be drawn from these data:

1) water nearer the southeastern boundary, i.e., the bad-waterline,of the aquifer
is more saturated, 2) concentration of Ca++ and HC03” may increase toward
the northeast, 3) higher temperature water is more saturated, 4) water from lower
yield wells is more saturated, and 5) spring water is the least saturated.

One difficult-to-assess variable that bears directly on the foregoing generaliz¬
ations is the size of the caverns intersected by a given well and their significance
in the overall pattern of groundwater movement. Groundwater flowing rather
rapidly through interconnected large openings is likely to be less saturated than
slower-moving water wending its way through less direct, smaller openings. This
concept may be of value in explaining the location of some of the more saturated
groundwater samples. Water from wells near the bad-water line seems to com¬
prise more than a proportional share of the saturated samples. The bad-water
boundary, on the down-dip side of the Edwards aquifer, is a dissolution-engrained
bypass boundary that marks the original extent of groundwater drawn to the

Well1 or Spring SIq Sljy Calcium Magnesium Bicarbonate Date Water Depth Well

(mg/I) (mg/1) (mg/1) collected temperature (feet) yield

(gpm)

SATURATION OF GROUNDWATER

163

o

o

O

o

o

o

o

o

o

o

o

o

o

o

o

o

O 1

1 MO

o

o

o

o

CO

co

o

o

00

O 1

I NO

o

CO

CNl

NO

»— <

pH

CO

CO

pH

CO

i—H

r-l

00

m

oo

c-~

5— <

CO

CO

*3-

o

o

o

NO

o

o

pH 1

CO 1

i r-

O

oo

MO

MO

CO

CO

CO

ON

o

NO

MO |

1 oo

c-

s>

r--

oo

U

MO

U

MO

r-

p-

MO

t"-

r-

O"

MO

cd

MO

MO

MO

wo

d

CO

wo

MO

cd

MO

cd

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

CO

Tf

•cf

CO

Nt

CO

p-

P-

r~-

t-

t"

r~

r-

I

r-

C-'

t*'-

1

I."*"

1

o-

1

r-

r~-

IN-

r~

r-

co

1

CO

1

NT

1

r-

1

r-~

1

r-

1

cn

1

CO

CO

1

CO

1

CO

1

co

ND

NO

MO

NO

M0

CO

CO

o

o

o

o

CN

■

(N

CO

CO

CO

CN

1

CO

T

pH

T

T

CO

CO

1

as

l

CO

1

CO

i

CO

1

1

rH

CO

CO

CO

1

CO

CO

CO

CO

MO

CO

MO

1— 1

pH

< — 1

r-H

pH

pH

r— <

CO

00

_H

oo

NO

NO

C"

NO

oo

o

NT-

NO

CO

CO

CO

CO

NO

N"

"ct*

M0

MO

Nj-

MO

NO

OO

NT-

NO

CO

oo

oo

pH

o

CO

CO

CO

co

CO

CO

CO

CO

CO

CO

CO

CO

CO

co

CO

CO

CO

CO

NO

NT-

NO

CO

o

MO

NO

NT

MO

MO

r-

C--

CO

NO

NO

r-

P-

p- 1

pH

CO

CO

pH

pH

*H 1

H

pH

^=*H

f— (

1— 1

P“<

1—1

CO

00

NO

W")

NO

o

CO

NO

ON

NO

o

oo

N5f

c->

p-

o

NO

NO

NO

NO

NO

NO

NO

P~

p-

r~<

oo

NO

c-

NO

ON

ON

l'-

00

oo

oo

00

NO

NO

NO

CO

CO

CO

CO

o

NO

•'T"

©

CO

■sj*

NO

•*fr

p- <

o 9

o

pH

NO

CO

CO

CO

1— 1

CO

■St

MO

©

CO

©

o

o

o

©

o

o

o

©

o

©

d

d

pH

H

d

r-H

d

+

1

1

+

1

1

1

1

+

1

+

1

1

1

1

1

1

NO

,-H

oo

p-

r-i

ON

o

CO

cn

CO

cn

ON

00

NO

NO

©

pH

CO

CO

CN

©

o

pH

©

pH

wn

t"

©

NO

o

©

©

o

d

©

d

o

d

©

©

d

d

©

d

©

©

©

©

+

1

1

1

+

1

+

1

1

c«

1

1

MS

so

&0

.s

c«

e

"»H

ao

so

CO

m

*rT

CO

a

C

c

-Nt

pH

oo

-Nt

NO

c

CO

CO

CO

•sf

on

”h

‘S

o

pH

o

o

o

o

©

o

pH

©

©

e «

c «

ex.

P,;

oo

1

CO

cn

1

CO

NO

1

CO

ON

1

NO

r->

p>“

r-

fH,

a,

m

.2

"S

t-

1

ON

CO

p-

as

I

ON

00

©

1

o

'o'

o

a>

W)

.s

bJD

.3

c n

wa

O

O0

«a

O

'Nf

O')

CO

CO

o

o

CO

CO

CO

m

cn

3

a

a

O

o

I

1

1

1

1

1

1

DO

m

?H

fH

oo

00

00

00

oo

c©

g

s

00

00

oo

oo

oo

KJ

ed

NO

NO

NO

NO

NO

NO

eu

<

NO

NO

NO

NO

NO

O

13

s

s

>

>

>

>■<

>

IX

§

c

ed

>

><

>

>

O

cs

S

o

s

o

G

cd

G

cd

<

<

<

<

<

<

m

00

<

<

<

<

<

U

U

m

m

Wells located by system of Texas Water Development Board.

164

THE TEXAS JOURNAL OF SCIENCE

earliest discharge sites (Abbott, 1975). As the down-dip portion of the aquifer
seems to have received less overall flow it follows that wells nearer the bad- water
line are more likely to penetrate smaller, less significant caverns that contain
more saturated water.

Wells with the lowest water yields furnished most of the supersaturated water
samples. This trend is sharply reversed by the supersaturated state of the water
from the highest yield well (202). Remember that even the most saturated waters
are only slightly supersaturated. Well 202 is about 2 mi from a salient in the bad-
water line and was sampled during late March 1972 when seasonal saturation ap¬
pears to have been high. At any rate, lower yield wells are probably drawing
from lesser developed conduits with slower water flow, increased contact with
the wall rock and, hence, higher levels of saturation.

The least saturated waters are those discharged from springs. This observation
may be explained by invoking the presence of large conduits leading more or less
directly from recharge areas to discharge sites. The large volumes of undersat¬
urated water directed to springs dissolve even larger conduits en route which sets
up a self-perpetuating system. As the size of caverns increases, an ever-decreasing
percentage of the groundwater is in contact with the walls at any instant. Thus
aggressive waters necessary for dissolution tend to be available and their volume,
compared to the surface area of the conduit walls, is great.

Effect of Mixing of Waters

The occurrence of rock dissolution in the Edwards aquifer may be appreci¬
ated by comparing the concentrations of Ca++, Mg++, and HCOT listed in Table 1
with the positions of the wells and springs on Fig. 1. Mg++ shows no systematic
changes in concentration but Ca++ and HCO3 do. The concentration of Ca++ in
mg/1 is generally in the 60’s in western Bexar County, in the 70’s in central and
eastern Bexar County, and in the 80’s in Comal and Hays Counties. The increase
in Ca++ concentration is probably due to dissolution of host rock by undersat¬
urated water moving northeast in the aquifer. This trend is most pronounced in
the central portion of the aquifer where water flow is presumably more volum¬
inous. In the vicinity of the bad-water line the Ca++ concentrations of water
samples are similar which further suggests that water does not circulate as freely
there.

The northeastward increase in HCO3 concentration parallels the increase in
Ca++ concentration. HCOJ concentrations in mg/1 are largely in the 240’s in
western Bexar County, in the 250’s and 260’s in central Bexar County, and in
the 300’s in Hays County.

Considering that the majority of the groundwater in the Edwards aquifer in
Bexar and Comal Counties has come in as underflow from the counties to the
west (Garza, 1962; Arnow, 1963), it is surprising to find groundwater undersat¬
urated with respect to calcite and dolomite at the distal end of the aquifer. Yet

SATURATION OF GROUNDWATER

165

the least saturated waters found in these analyses were collected from Comal and
San Marcos Springs which are over 150 mi from the proximal, or recharge-
dominated, end of the aquifer and despite the apparent increase in Ca++ and
HCOJ concentrations through Bexar, Comal and Hays Counties.

The processes that occur to create this unexpected undersaturation have been
explained best by Thrailkill (1968). Meteoric water that descends through the
soil and unsaturated zones (vadose seepage) may enter the aquifer in a saturated
or supersaturated state, but if it is cooled upon entering the main groundwater
body or is mixed with groundwater which is in equilibrium with a lower partial
pressure of C02 , the net effect is to cause undersaturation in the main water
body. The significant process here is the introduction of C02 into the ground-
water body which allows further dissolution to occur.

Probably the most important factor in keeping the main body of groundwater
undersaturated is the introduction of large volumes of flood water in Medina,
Bexar and Comal Counties that are undersaturated with respect to calcite and
dolomite. Arnow (1963) has estimated that about 1/6 of the groundwater in the
Edwards aquifer in Bexar County is derived as local recharge from the Medina
River, Cibolo Creek and other smaller streams such as Salado and Leon Creeks.
George (1952) calculated that about 1/4 to 1/3 of the water flow from Comal
Springs is obtained as recharge from the Cibolo and Dry Comal Creek basins; the
Guadalupe River flows across the aquifer without significant changes in its dis¬
charge. Saturation indices for water in creeks that recharge the aquifer were not
calculated because no chemical analyses with field-measured pH were available.
However, the concentrations of Ca++, Mg++, and HCOJ are available for surface
water samples collected under variable discharge conditions from these creeks
(e.g. Land, 1972, Steger, 1973). The bulk of the waters that flow in these creeks
have Ca++, Mg++, and HCOJ concentrations that are markedly less than those in
the aquifer. During times of low discharge, when the creeks are fed by small
springs, the ion concentrations in the water may be higher than in the aquifer.
But the stream waters that flow in the high discharge conditions that occur after
heavy rains commonly have ion concentrations that are only 1/4 to 1/2 of those
in the aquifer (Tables in Land, 1972, and Steger, 1973). More recharge occurs
from floods than from base flow. The addition of these high volume, low ionic
concentration waters into the Edwards aquifer seems to be the major factor in
maintaining a general state of undersaturation.

For purposes of comparison it is interesting to note the ion concentrations in
Waco Springs water (Table 1). Although this spring system discharges just 3 mi
north of Comal Springs it is not connected to the major aquifer and its recharge
area appears to be entirely within Comal County to the west and northwest of
the springs (Fig. 1). The mid-December, 1974 analysis showed the spring water
had a 1/8 to 1/4 higher concentration of Ca++ and HCOJ than the waters dis¬
charged from the major aquifer at Comal and San Marcos Springs. The Waco
Springs are a much simpler system where undersaturated flood waters recharge

166

THE TEXAS JOURNAL OF SCIENCE

a separate but parallel aquifer, flow several mi in a large cavern system without
dilution by significant amounts of other recharge, and emerge with higher ion
concentrations than the long distance waters discharged at Comal and San Marcos
Springs. The higher ion concentrations developed in short flow distances in this
local, although large, system are probably attributable in part to the absence of a
significant mixing- of- waters effect.

CONCLUSIONS

Despite straight-line travel distances exceeding 160 mi for some of the ground-
water in the Edwards aquifer the groundwater appears to be undersaturated at
times. This undersaturation is due partly to large volumes of groundwater flow¬
ing in pipe- like voids where little of the water is actually in contact with the
host rock, and partly to the mixing effect that occurs with addition of signif¬
icant volumes of undersaturated recharge. The undersaturated state of some of
the groundwater in the aquifer implies that caverns are probably being enlarged
during seasonal conditions. Those areas transmitting the greatest amounts of
water probably are receiving the greatest amounts of dissolution.

ACKNOWLEDGEMENTS

This paper was made possible by the chemical analyses of water samples
furnished by Robert W. Maclay of the U.S. Geological Survey in San Antonio.
Special thanks go to Paul Rettman of the U.S.G.S. for collecting the water samples
and making field measurements of pH, specific conductance and bicarbonate
concentration. The maiiUscript was improved by the comments of Charles M.
Woodruff, Jr. of the Bureau of Economic Geology at the University of Texas.
The figure was drafted by Enos. J. Strawn.

LITERATURE CITED

Abbott, P.L., 1975 -On the hydrology of the Edwards Limestone, south-central Texas.
Jour. Hydrology , 24 : 25 1 .

Alexander, W.H., Jr., B.N. Myers, and O.C. Dale, 1964— Reconnaissance investigation of the
ground- water resources of the Guadalupe, San Antonio, and Nueces river basins, Texas.
Texas Water Comm. Bull , 6409, 106 pp.

Arnow, T., 1963— Ground- water geology of Bexar County, Texas. U.S. Geol. Survey Water-
Supply Paper, 1588, 36 pp.

Garrels, R.M., and C.L. Christ, 1965 -Solutions, Minerals , and Equilibria. Harper and Row
Pub., New York, 450 pp.

Garza, Sergio, 1962— Recharge, discharge, and changes in ground-water storage in the Ed¬
wards and associated limestones, San Antonio area, Texas, a progress report on studies,
1955-59. Texas Board Water Engin. Bull, 6201, 42 pp.

SATURATION OF GROUNDWATER

167

George, W.O., 1952-Geology and ground- water resources of Comal County, Texas. U.S.
Geol. Survey Water-Supply Paper 1138, 126 pp.

Jacobson, R.L., and D. Langmuir, 1970 -The chemical history of some spring waters in
carbonate rocks. Ground Water, 8:5.

Land, L.F., 1972— Annual compilation and analysis of hydrologic datq for urban studies in
the San Antonio, Texas metropolitan area, 1970. U.S. Geol. Survey, Water Resources
Div. Pub., 178 pp.

Shuster, E.T., and W.B. White, 1971 -Seasonal fluctuations in the chemistry of limestone
springs: a possible means for characterizing carbonate aquifers. Jour Hydrology , 14:93.

Steger, R.D., 197 3 -Annual compilation and analysis of hydrologic data for urban studies
in the San Antonio, Texas metropolitan area, 1971. U.S. Geol. Survey, Water Resources
Div . Pub., 109 pp.

Thrailkill, J., 1968-Chemical and hydrologic factors in the excavation of limestone caves.
Geol. Soc. Am. Bull., 79:19.

- , 1972— Carbonate chemistry of aquifer and stream water in Kentucky. Jour.

Hydrology, 16:93.

FITTING SOIL TEMPERATURE BY A PERIODIC REGRESSION
MODEL

by MINGTEH CHANG

School of Forestry, Stephen F. Austin State University,

Nacogdoches 75962

and DOUGLAS G. BOYER

Division of Forestry, West Virginia University,

Morgantown 26505

ABSTRACT

A sine curve regression model is presented to which monthly mean soil temperature may
be fitted from mean air temperature. The model was separately tested at Nacogdoches,
Texas and Kearneysville, West Virginia with predictabilities greater than 98% and standard
error of estimates less than 1 C.

THE PROBLEM

Soil temperature is one of the important factors governing biological, chemical,
and physical processes in soil. For example, the rates of organic-matter decom¬
position and evaporation loss increase with soil temperature, while seed-germin¬
ation and root-growth are impossible for most species when temperature is less
than 40 F. Below freezing there is little biological activity, no chemical weather¬
ing, no liquid water movement, and, as a consequence, no soil formation — time
stands still for the soil. As Smith, et al., (1964) expressed it, soil comes to life
when its temperature exceeds 42 F. For these reasons, mean annual soil temper¬
ature (at 50 cm depth) is used as one of the criteria to differentiate soil families
in the new U.S. soil classification system (USDA, 1975).

Soil temperature varies in time and space due to the difference in 1) heat
flux into the soil, 2) the heat transfer in the soil, and 3) the heat exchange
between soil and air. Since soil temperature data are usually available at only
a few stations in each state, estimates based on air temperature or other climat¬
ological elements become necessary for practical purposes. The Soil Conserva¬
tion Service (Smith, et al., 1964) estimates annual soil temperature by adding

Accepted for publication: January 28, 1977.

The Texas Journal of Science, Vol. XXIX, Nos. 3-4, December, 1977.

170

THE TEXAS JOURNAL OF SCIENCE

2 F (about 1 C) to the annual air temperature. For a farmer or forester, however,
monthly temperature is more important than annual as it reflects seasonal pat¬
terns of evaporation, transpiration, snowmelt, and plant growth. The objective
of this study was to develop a periodic regression model to estimate monthly
soil temperature at Nacogdoches, Texas with acceptable accuracy.

THE PREDICTION MODEL

Heat transfers in soil are governed by the nature of the soil, the type of sur¬
face cover, and incoming radiation. For estimating long-term average monthly
soil-temperature at a particular station, one can assume that basic soil prop¬
erties are constant and surface cover varies in a relatively fixed pattern during
the year. Thus the seasonal fluctuation of soil temperature is controlled pri¬
marily by incoming radiation, which can be indexed as a function of air temper¬
ature (TA) and time angle:

STt = a0 + a! TAt + a2SIN(27rt/12) + a3Cos(27rt/12) + et (1)

where

ST = monthly soil temperature, C,

2*1/12 = the monthly units of time in a single cycle to
angular measure in radians,
t = month, from 0 (January) to 1 1 (December),
et -- error term,

and a0, a! , a2 , and a3 are constant. Equation 1 is a sine curve with modification
of air temperature. For estimating annual soil temperature (STy), the Equation
becomes

t = 1 1

STy - "t &i( 2 ^ TAt)/ 12 + €y (2)

where ey is the error of the predicted annual soil temperature. Once Equation 1
is developed, the only climatological data required for solving the equations are
monthly mean air temperatures, TA. Values of the time angle, 27rt/12, for each
month are shown in Table 1.

DATA, ANALYSIS, AND RESULTS

The soil temperature data used in the analysis were observed at the Stephen F.
Austin Experimental Forest, 15 miles southwest of Nacogdoches, Texas from
May 1957 to September 1959. The soils are loams and sandy loams of the
Shubuta series (formerly mapped as Boswell series), a member of the clayey,
mixed, thermic family of Aquic Hapludults, supporting a mixed stand of sec¬
ond-growth hardwoods and loblolly-shortleaf pine. The annual rainfall during

SOIL TEMPERATURE

171

TABLE 1

Values of the Monthly Time Angles

Month

t

Sin(27Tt/12)

Cos(2tt/12)

January

0

0.0000

1.000

February

1

0.5000

0.8660

March

2

0.8660

0.5000

April

3

1.0000

-0.0000

May

4

0.8660

-0.5000

June

5

0.5000

-0.8660

July

6

-0.0000

-1.0000

August

7

-0.5000

-0.8660

September

8

-0.8660

-0.5000

October

9

-1.0000

0.0000

November

10

-0.8660

0.5000

December

11

-0.5000

0.8660

the period was 47.58 in (1209 mm), or 0.46 in (12 mm) below normal (193 1-60).
Soil temperature was recorded tri-weekly using Colman fiberglass units with
thermisters at two different depths (6 and 12 in) in nine 1/10 acre circular plots.
Air temperature at 4 ft above the ground was observed at the weather station in
the SFA Experimental Forest. The data have been published by Schneider and
Stransky (1966).

Equation 1 was employed to fit the 29 monthly soil temperature values
for each depth observed at the Experimental Forest. The analysis showed that
this equation explained more than 98% of soil- temperature variation, with a
standard error of estimates less than 3.7% or 0.69 C (Table 2). Since the thermal
capacity, conductivity, and diffusivity of a soil are affected by its moisture con¬
tent, precipitation (use as an index of soil moisture) was added to Equation 1 in
the regression analysis. However, the inclusion of a precipitation variable did not
improve the accuracy of the prediction.

TABLE 2

The Constants and Simple Statistics of Equation 1 for Two Different Locations

Location

Depth

(In)

a0

Constants
al a2

a3

Statistics1
R2 SE,%

Nacogdoches, Texas

6

2.82

0.91

-0.84

1.49

0.987

3.5

20

5.05

0.82

-1.73

1.72

0.980

3.7

Kearneysville, W.V.

8

7.85

0.48

-2.22

-6.25

0.987

8.2

20

9.57

0.47

-3.66

-4.39

0.976

8.7

1 2

R : Coefficient of multiple determination; SE: standard error of estimate.

172

THE TEXAS JOURNAL OF SCIENCE

The average observed and estimated values at the 6-in depth were plotted in
Fig 1 for further illustration. Greatest errors of estimates at the 6- and 12-in
levels were -1.26 C and -1.40 C, respectively, both in September 1959. Of the

Month

Figure 1. Estimated and observed monthly mean soil temperatures at the 6-in level and the
observed air temperatures at the SFA Experimental Forest, Nacogdoches, Texas.

SOIL TEMPERATURES

173

58 estimates, only 4 at the 6-in level and 6 at the 12-in level had errors of esti¬
mate greater than ± 1 C. Except for the January soil temperature at the 12-in
level in 1958, all the greatest errors were in the summer months (i.e., July,
August, and September). Normally the 3 summer months are the driest of the
year in Nacogdoches. As soils become drier their thermal capacity tends to be
reduced, thus increasing the fluctuations of the soil temperatures. This would
account, at least in part, for the greatest errors in estimates of soil tempera¬
tures occurring during the summer.

The soil temperatures regime, as shown in Figure 1, at the SFA Experimental
Forest followed the general patterns in humid regions described by Chang (1958).
Heat Conduction to and from the air-soil interface causes monthly mean soil
temperature to increase with depth during winter and to decrease with depth
during summer. Soil temperatures at 6 and 12 in were generally warmer than
mean air temperature during the colder months and cooler during the warmer
months. Temperature differences in colder months, however, were less than in
observations made in West Virginia (Chang and Lee, 1977).

APPLICATION

Applicability of Equation 1 was treated by fitting 36 monthly mean soil
temperature values (1963-65) at two depths (8 and 20 in) observed at Kearneys-
ville, West Virginia (Weedfall, et al., 1967). The soil is a Hagerstown series of
fine, mixed, mesic family, Typic Hapludalfs. The analysis (Table 2) showed
that the R2 values at these two soil levels were greater than 0.97 with an average
absolute error less than 7% (0.97 C). The great accuracy suggests that Equation 1
is an applicable model for fitting monthly mean soil temperature data at a part¬
icular station. If data are available at sufficient number of stations for spatial
analysis, it could be also employed to estimate monthly soil temperatures at un¬
aged sites in a region.

ACKNOWLEDGEMENTS

The authors wish to thank Mr. George K. Stephenson and Dr. K.G. Watterston,
Stephen F. Austin State University, for their review of this manuscript, and
to Dr. J.J. Stransky, U.S. Forest Service, for his temperature data of Nacog¬
doches, Texas.

LITERATURE CITED

Chang, J-H., 1958 —Ground Temperature, Vol. I. Blue Hill Meteo. Obs., Harvard Univ. 300 pp.

Chang, M., and R. Lee, 197 7 -On the adequacy of hydrologic data in West Virginia. Water

Res. Inst., West Virginia Univ. (In Press).

Schneider, G., and J.J. Stransky, 1966-Soil moisture and soil temperature under a post

oak-shortleaf pine stand. School of Forestry, Stephen F. Austin State Univ., Bull. 8.

24 pp.

174

THE TEXAS JOURNAL OF SCIENCE

Smith, G.D., F. Newhall, L.H. Robinson, and D. Swanson, 1964-Soil temperature regimes-
their characteristics and predictability. Soil Conserv. Service, U.S.D.A. SCS-TP-144,
13 pp.

USDA, 191 5 -Soil taxonomy, USD A Agr. Handbook 436. Washington, D.C. 754 pp.

Weedfall, R.O., W.H. Dickerson, and W.H. Stirm, 1967 -The agroclimate of university ex¬
periment farm, Kearney sville, West Virginia. Current Report 52, West Virginia Univ¬
ersity, Agri, Exp. Sta., Morgantown, W.V. 19 pp.

TRACE FOSSILS FROM THE PECAN GAP FORMATION (UPPER
CRETACEOUS), NORTHEAST TEXAS1

by WILLIAM C. DAWSON

Eason Oil Company, 5225 North Shartel,

Oklahoma City 73118

DONALD F. REASER

Department of Geology,

The University of Texas at Arlington 76019

and JIMMY D. RICHARDSON

Phillips Petroleum Company- UK Branch,

Portland House, Stag Place, London, England SW1E 5DA

ABSTRACT

The Pecan Gap Formation in northeast Texas exhibits varied lithologies and several
distinctive sedimentary features that provide significant information concerning the de-
positional history of the Taylor Group. Trace fossils are the characteristic megafossil and
consequently furnish the most promising source of paleoenvironmental data.

Two distinct assemblages of ichnofossils were identified: 1) small Thalassinoides , hori¬
zontal Rhizo cor allium, and Stipsellus occur within a thinly bedded glauconitic facies;
2) large Thalassinoides, vertical Rhizo cor allium, Gyrolithesfl), Pseudobilobites, and nu¬
merous vertical burrows are found in a “massive” quartzose calcarenite facies. Both as¬
semblages are indicative of inner shelf deposition. Morphologic differences displayed by
these 2 groups of trace fossils probably resulted from local variations in energy as suggested
by physical sedimentary features.

INTRODUCTION

Our knowledge of the Taylor Group in northeast Texas has evolved gradually
to a picture of complex stratigraphic relations which consist of a myriad of inter¬
calated facies. It is generally accepted that the Taylor Group is a neritic marine

^his work was taken in part from a M.S. thesis in geology completed by the senior author
at The University of Texas at Arlington, August, 1976.

Presented at the 79th annual meeting of the Texas Academy of Science, February, 1976,
College Station, Texas.

Accepted for publication: January 23, 1977.

The Texas Journal of Science, Vol. XXIX, Nos. 3-4, December, 1977.

176

THE TEXAS JOURNAL OF SCIENCE

sequence deposited during the Late Cretaceous (Beall, 1964; Powell, 1970).
However, a comprehensive depositional model including all local facies has not
yet been established.

Presented herein is paleoenvironmental information derived primarily from
studies of trace fossils collected at exposures of the Pecan Gap Formation in
Collin and Rockwall counties, Texas (Fig. 1). The present writers hope that our
results will be integrated with future surface and subsurface studies to develop
a complete history for Late Cretaceous sedimentation in northeast Texas.

Figure 1. Location of study area and outcrop of the Wolfe City and Pecan Gap form¬
ations (Geology modified from Barnes, 1967, 1972).

TRACE FOSSILS

177

GENERAL STRATIGRAPHY

The general stratigraphy of the Taylor Group is essentially as described origi¬
nally by Stephenson (1918). The Pecan Gap Formation disconformably over-
lies the Wolfe City Sandstone in its type area and- is a relatively thin medial
carbonate unit within a thicker, dominantly argillaceous sequence.

Although the Taylor Group can be subdivided into lithologically distinct
rock units on a regional scale, localized facies complicate the stratigraphy

(Fig-1)-

Near Farmersville in Collin County the Pecan Gap changes laterally from a
biocalcarenite to a densely burrowed glauconitic and phosphatic calcarenite
within a distance of about a mile (Rouse, 1944; Brown, 1949). Brezina (1974)
proposed the name “Farmersville Member” for this bioturbated glauconitic
facies.

The member is bounded laterally on the north by the Wolfe City Formation
(Brezina, 1974) and disconformably overlies the Ozan Formation (Brown, 1949;
Dawson, 1976). The basal contact is marked by a discontinuous phosphate con¬
glomerate.

According to Brezina (1974, Fig, 5), the Farmersville Member is laterally
juxtaposed with a silty marl that is present in Rockwall and southern Collin
counties. Maluf (1975) undertook a study of this area and subsequently divided
the Pecan Gap Formation exposed there into the following lithologic units:
Rockwall Member (silty marl) and Bear Creek Member (calcarenite). Based
on microfauna, Maluf (1975, Fig. 3) correlated the Farmersville Member with
the Bear Creek Member. Much of the area between these rock units is covered
with alluvium and the details of lithostratigraphy remain in question.

DESCRIPTION

Previously, trace fossils from the Taylor Group of northeast Texas were
described systematically by Richardson (1972, 1975) and Dawson (1976). In
the descriptions that follow ichnofossils are treated as biogenic sedimentary
structures, and their stratigraphic position and orientation are compared with
those of the physical sedimentary features. Biogenic structures are described
using the stratinomic (preservational) terminology of Chamberlain (1971, Fig. 3)
and descriptions of Hantzschel (1975) are followed for established ichnogenera.

Farmersville Member

The Farmersville Member (Brezina, 1974, Fig. 1) is exposed in several quarries
northwest of Farmersville in Collin County. It has an areal extent of about
1.5 km and an average thickness of 9 m. This lenticular rock body consists
of sharply-defined, laterally continuous, thin beds (5-10 cm) composed of alter¬
nating quartzose-glauconitic calcarenite and montmorillonitic claystone. Even

178

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TRACE FOSSILS
(cont.)

179

g

Figure 2. a) Quarried slab from Farmersville Member composed of a “mat” of interwoven
thalassinoidean burrows. MS C-3X. Scale in cm. b) Rhizo cor allium sp., frag¬
ment of a vertically protrusive horizontal structure. Unit 5, MS C-2. c) Stip-
sellus sp., MS C-2X. d) Stipsellus sp., MS C-3X. e) Thalassinoides sp., Unit 3,
MS C-3. Scale in cm. 0 Thalassinoides sp., MS C-2X. g) Smooth-walled verti¬
cal tube. Unit 5, MS C-2.

parallel laminations (1-2 mm) are the dominant internal sedimentary structure.
These well-developed sedimentary features are unique to the Farmersville Mem¬
ber; generally the rock units of the Taylor Group are internally structureless.

Bioturbation is characteristic of the Farmersville Member (Rouse, 1944;
Brown, 1949; Brezina, 1974). Trace fossils occur in most of the well-cemented
calcarenites but remain undetected, if present, in the interbedded claystones.
Biogenic reworking of the calcarenites is commonly so dense that it results
in a rock composed of a “mat” of interwoven burrows (Fig.2a).

Although the Farmersville Member has generally been described as “densely-
burrowed,” detailed examination reveals that the degree of bioturbation is
highly variable. Some beds are completely homogenized with all evidence of
primary sedimentary features obliterated; in others, burrows and delicate la¬
minations occur together. These variations occur vertically and also within single
beds along an exposure.

Most trace fossils within the Farmersville Member are rather nondescript
smooth-walled forms: intergenic and endogenic full-relief burrows. These
burrow systems consist . of interwoven horizontal cylindrical tubes (5-8 mm in
diameter) that can generally be traced for several m across bedding surfaces.
Burrow fillings consist of inoceramid prisms, foraminifers, and Fine quartz sand
cemented with sparry calcite. These traces commonly have an external rim of
glauconite.

Recognizable ichnogenera in the Farmersville Member include horizontal,
vertically-protrusive Rhizo cor allium (Fig. 2b), Stipsellus (Fig. 2c, d), and Thalas-

180

THE TEXAS JOURNAL OF SCIENCE

sinoides (Figs,2e,f). Thalassinoidean structures in the Farmersville Member con¬
sist of simple horizontal tubes with Y-bifurcations. These structures lack many
features associated with typical Thalassinoides sp. (Hantzschel, 1975): lon¬
gitudinal ridges, vertical shafts, swellings at branching points, and bulbous “dead¬
end” burrow terminations.

Rhizocorallium, Stipsellus, and Thalassinoides occur within laminated cal-
carenites throughout most of the Farmersville Member. However, the upper part
of the member contains only vertical burrows (Fig42g), which are associated
with bimodal cross-laminations.

Bear Creek Member

The Bear Creek Member consists of about 9 m of “massive” quartzose cal-
carenite composed primarily of inoceramid prisms and foraminifers with only
minor amounts of glauconite and collophane. In contrast to the Farmersville
Member, the Bear Creek Member is intensely bioturbated throughout, and com¬
pletely homogenized, leaving no evidence of primary depositional features.

The trace fossil assemblage includes relatively large and robust Thalassinoides
systems with both horizontal and vertical components (Fig.3, a, b, c, d, e); ver-
tically -protrusive Rhizocorallium oriented perpendicular to bedding (Fig. 3f);
Gyrolithes (?); and numerous vertical tubes with “scratch marks” (Fig, 3g).
These forms occur together through the total thickness of the member. The cal-
carenite burrow fillings are better sorted and contain more sparry calcite cement
than the surrounding sediment. Hence, the networks weather into resistant frag¬
ments. Also, phosphate nodules (Fig. 3d) are concentrated in thalassinoidean
structures.

Pseudobilobites sp. (Fig.3h) is abundant in the subjacent Rockwall Member
(Maluf, 1975) and in the upper part of the Wolfe City Formation throughout
northeast Texas (Richardson, 1972). However, this ichnogenus is scarce in the
Bear Creek Member.

Rhizocorallium (Fig. 3f) is similar to U-shaped, spreiten-bearing, vertical
burrows described as Diplocraterion by Fursich (1974). However, the forms in
the Bear Creek Member display external striae, a significant feature of many U-
shaped burrows that was not discussed by Fursich (1974). According to Hantz¬
schel (1975), external striae (scratch marks) are characteristic of Rhizocorallium ;
the terminology of Hantzschel is retained herein.

DISCUSSION AND INTERPRETATION

Trace fossils are difficult to classify phylogenetically because: 1) organisms
responsible for the structures are rarely preserved; 2) a single animal is capable
of producing a variety of traces; and 3) animals which differ taxonomically may
create similar traces. Therefore, trace fossils should be considered independently
of their producers (Hantzschel, 1975). Even though the biologic affinities of

TRACE FOSSILS

181

most ichnofossils remain in question, they are potentially valuable in environ¬
mental reconstructions.

Creatures dwelling in the sediment interact with the sedimentation processes
of the environment. Seilacher (1964, 1967) suggested that these processes
largely control the behavior of an animal which in turn controls the morphology
of its traces. The underlying assumption of Seilacher’s classification is that ani¬
mals with similar life habits produce similar traces, regardless of their anatomical
features. This concept has been used to recognize trace fossil assemblages and ex¬
plain their distribution by variations in bathymetry. The associations illustrated
by Seilacher (1967) show a gradation from vertical (protective) burrows in the
shallowest marine environments to highly patterned, horizontal (feeding) burrow
systems in deep areas: these traces are said to reflect a change from suspension-
to sediment-eating habit.

Seilacher’s model predicts that the lateral and vertical distribution of trace
fossils in ancient sediments should reflect a bathymetric sequence on a sub¬
marine slope. However, later workers have demonstrated that factors other than
depth also influence the distribution of trace fossils. In shallow marine environ¬
ments the complexities of sedimentation can result in local energy conditions
or a substrate that is generally associated with another depth zone (Crimes, 1970;
Frey, 1971; Hill, 1976). Thus, trace fossils can occur locally in sediment de¬
posited outside the normal depth limits of that assemblage. Using these con¬
cepts an interpretation of the Pecan Gap trace fossils will now be undertaken.

The ichnocoenoses of the Pecan Gap Formation in the study area belong to
the shallow marine ( Cruziana ) trace fossil assemblage of Seilacher (1964, 1967).
The physical and biogenic sedimentary structures of the Farmersville Member
are similar to many features that occur in modern and ancient inner shelf de¬
posits (Howard, 1972; Goldring and Bridges, 1973; Cotter, 1975; and Kumar
and Sanders, 1976). The dominant primary structure in each case is even parallel
laminations. According to Reineck and Singh (1972) laminated. shallow marine
sands are commonly produced by a process of “settling-out” from suspension
following storm-generated turbulence. Cumulative-frequency grain size curves
presented by Dawson (1976) are similar to curves illustrated by Visher (1969)
and Kumar and Sanders (1976) for Holocene sands deposited from suspension.

Stratigraphic relations indicate that the Farmersville sediments accumulated
in a trough shoreward of a submarine high developed in the Ozan Formation.
This positive feature acted as a velocity barrier to storm currents (Dawson, 1976).
During fair-weather periods, normal suspension clays and silts were deposited,
and the storm-generated sands were reworked by burrowing organisms. Longer
interstorm periods permitted more thorough churning of the sediment. When the
biogenic (disruptive) processes were interrupted by storm activity, previously-
deposited storm layers remained only partly reworked as the surviving organisms
“escaped” upward to colonize the new sediment horizon. The irregular alterna¬
tions in sedimentation rate explain the variable degree of bioturbation observed
within the Farmersville Member.

182

THE TEXAS JOURNAL OF SCIENCE

TRACE FOSSILS

183

Figure 3. a) Thalassinoides sp., note swollen node at branehing point, b) Thalassinoides
paradoxica, MS R-5X. c)Thalassinoides sp., “dead-end” burrow termination,
d) Thalassinoides sp., showing horizontal and vertical components. Note phos¬
phate pebbles (p). e) Thalassinoides sp., MS R-5X. f) Rhizo cor allium sp..
Unit 1, MS R-5. g) Vertical burrow displaying “scratch marks.” h) Pseudo-
bilobites sp., Unit 3, MS R-5.

Agerand Wallace (1970) described an association of horizontal Rhizocorallium
and small “atypical” Thalassinoides from a protected area shoreward of a shal¬
low marine barrier: a situation similar to the trace fossil assemblage of the Farm-
ersville Member. The density of Stipsellus and Thalassinoides suggests an in¬
tensive feeding behavior in an area of low turbulence and generally slow sedi¬
mentation rates (Frey. 1971). The textural immaturity of the Farmersville sedi¬
ments and the abundance of glauconite also support a low energy depositional
environment (Dawson, 1976).

The upper part of' the Farmersville Member displays bimodal cross-lamina¬
tions and vertical burrows suggestive of a tidally-dominated zone. Apparently,

184

THE TEXAS JOURNAL OF SCIENCE

as deposition progressed, the Farmersville sediments built upward into a level
of higher energy and more rapid sedimentation.

The Bear Creek Member is internally structureless making environmental in¬
terpretations relatively difficult. Extensive reworking of these carbonate sands
suggests that generally slow and continuous deposition prevailed. The trace fossil
assemblage of the Bear Creek Member is similar to crustacean communities
found in lower shoreface environments (Howard, 1972). These interpretations
are consistent with those proposed by Richardson (1975) for the Wolfe City
Formation.

CONCLUSIONS

Trace fossils are abundant in the Pecan Gap Formation of northeast Texas
and because of their autochthonous nature, they provide the most reliable
source of paleoenvironmental information. The combined sedimentologic and
paleontologic data suggest that Pecan Gap sediments in the study area were de¬
posited on the inner shelf. Farmersville sediments accumulated shoreward of a
submarine barrier in an area influenced by storm and tidal currents. Bear Creek
sedimentation occurred in the subtidal environment.

Strata of the Taylor Group in northeast Texas are perhaps most similar to
Holocene nearshore deposits off the Gulf and Atlantic coasts described by Shepard
and Moore (1955) and Howard and Reineck (1972). These sediments occur on
a tide-dominated shelf where normal physical and biologic processes are period¬
ically interrupted by storm activity.

ACKNOWLEDGEMENTS

We are indebted to Dr. Bob F. Perkins, Dean of the Graduate School, Uni¬
versity of Texas at Arlington and Dr. Robert W. Scott, Amoco Production Re¬
search, Tulsa for their helpful suggestions and valuable criticism of the manu¬
script.

The senior author extends his sincerest gratitude to the following persons:
especially, Dr. Donald F. Reaser for his encouragement and guidance, his assist¬
ance in the field, and his critical review of the manuscript; Mr. Jimmy D. Richard¬
son provided fossil specimens and other unpublished data gathered during his
thesis research at UT- Arlington; Ms. Kathy Nelms edited the original manu¬
script and provided encouragement during every phase of the study; Ms. Wanda
Slagle typed the final manuscript; and Mr. Robert Mathis aided in the collection
of specimens.

LITERATURE CITED

Ager, D.V., and P. Wallace, 1970-The distribution and significance of trace fossils in the
uppermost Jurassic rocks of the Boulonnais, northern France. In T.P. Crimes and J.C.
Harper (Eds.), Trace Fossils, Liverpool, Steel House Press, pp. 1-18.

TRACE FOSSILS

185

Beall, A.O., 19 64 -Stratigraphy of the Taylor Formation, Upper Cretaceous, east-central
Texas. Baylor Geol. Studies Bull No. 6, 34 pp.

Brezina, J.L., 19 74 -Stratigraphy and petrology of the Pecan Gap Formation (Taylor Group,
Upper Cretaceous) in its type area. Unpubl. M.S. Thesis, Univ. Texas, Arlington, 63 pp.

Brown, O.C., 1949-Geology of the Verona area, Collin County, Texas. Unpubl. M.S.
thesis, Southern Methodist Univ., 21 pp.

Chamberlain, C.R., 1971— Morphology and ethology of trace fossils from the Ouachita
Mountains, southeastern Oklahoma. J. Paleontology, 45:212.

Cotter, E., 1975 -Late Cretaceous sedimentation in a low-energy coastal zone; the Ferron
Sandstone of Utah. J. Sed. Petrology, 45:669.

Crimes, T.P., 1970-The significance of trace fossils in sedimentology, stratigraphy, and
paleoecology with examples from Lower Paleozoic strata. In T.P. Crimes and J.C.
Harper (Eds.), Trace Fossils, Liverpool, Steel House Press, pp. 101-126.

Dane, C.H., and L.W. Stephenson, 1928— Notes on the Taylor and Navarro formations in
east-central Texas. Am. Assoc. Petroleum Geologists Bull., 12:41.

Dawson, Wm. C., 1976-Petrography and sedimentation of the Upper Cretaceous Farmers-
ville Member, Pecan Gap Formation, Taylor Group of northeast Texas. Unpubl. M.S.
thesis, Univ. Texas, Arlington, 158 pp.

Frey, R.W., 1971— Ichnology, the study of fossil and recent lebensspuren. In B.F. Perkins
(Ed.), Trace Fossils, Louisiana State Univ., Misc. Publ. 71-1, pp. 91-125.

Fursich, F.T., 1974-On Diplocraterion Torell 1870 and the significance of morphological
features in vertical, spreiten-bearing, U-shaped trace fossils. J. Paleontology, 48:952.

Goldring, R., and P. Bridges, 1973 -Sublittoral sheet sandstones. /. Sed. Petrology , 43:736.

Hill, G.W., 19 76 -Biogenic sedimentary structures of south Texas outer continental shelf.
Am. Assoc. Petroleum Geologists Abs. with Programs, 1976 Annual Meeting, pp. 72.

Hantzschel, W., 1975-Trace fossils and problematics. In C. Teichert, (Ed.), Treatise on In¬
vertebrate Paleontology, Part W, Supplement I: Geol. Soc. America, Univ. Kansas
Press, 269 pp.

Howard, J.D., 1972-Trace fossils as criteria for recognizing shorelines in the stratigraphic
record. Soc. Econ. Paleontologists and Mineralogists, Spec. Pub. 16, pp. 215.

- and H.E. Reineck, 1972-Georgia coastal region, Sapelo Island, U.S.A.: Sedi¬
mentology and biology IV, physical and biogenic structures of the nearshore shelf.
Senckenbergiana maritima, 4:81.

Kumar, N., and J.E. Sanders, 1976 -Characteristics of shoreface storm deposits; modern
and ancient examples./. Sed. Petrology, 46:145.

Maluf, F.W., 1975 -Stratigraphy of the Rockwall Member of the Upper Cretaceous Pecan
Gap Formation, Collin and Rockwall counties, Texas. Unpubl. M.S. thesis, Univ. Texas
Arlington, 65 pp.

Powell, J.D., 1970-Summary of Upper Cretaceous stratigraphy . Interamerican Micropaleon-
tological Colloquium Field Trip Guidebook, July 19-30, 91 pp.

186

THE TEXAS JOURNAL OF SCIENCE

Reineck, H.E., and EB. Singh, 1972-Genesis of laminated sands and graded rhythmites in
storm- sand layers of shelf mud. Sedimentology , 18:123.

Richardson, J.D., 1972-Stratigraphy and depositional environment of the Wolfe City Form¬
ation (Upper Cretaceous) northeast Texas. Unpubl. M.S. thesis, Univ. Texas, Arling¬
ton, 151 pp.

- , 1975-Trace fossils from the Wolfe City Formation, Upper Cretaceous, north¬
east Texas. Texas J. Sci., 26:339.

Rouse, J.T., 1944 -Correlation of the Pecan Gap, Wolfe City, and Annona formations in
east Texas. Am. Assoc. Petroleum Geologists Bull, 28:522.

Seilacher, A., 1964-Biogenic sedimentary structures, In J. Imbrie and N.D. Newell (Eds.),
Approaches to Paleoecology , New York, John Wiley, pp. 296-316.

- , 1967-Bathymetry of trace fossils. Marine Geology, 5:413.

Shepard, F.P., and D.G. Moore, 1955 -Central Texas coast sedimentation; characteristics of
sedimentary environments, Recent history, and diagenesis. Am. Assoc. Petroleum Geolo¬
gists Bull., 39:1463.

Stephenson, L.W., 19 18 -A contribution to the geology of northeastern Texas and southern
Oklahoma. U.S. Geol. Survey Prof. Paper 120, pp. 129-163.

PRELIMINARY FIELD STUDY OF THE FRACTURE PATTERNS
ASSOCIATED WITH THE BALCONES FAULT ZONE IN NORTH
CENTRAL TEXAS

by R. G. FONT, J. C. YELDERMAN,

C. T. HAYWARD, and E. E. BALDWIN

Department of Geology, Baylor University,
Waco 76703

ABSTRACT

A study of the fracture patterns associated with the Balcones Fault Zone in north central
Texas has been conducted. The study has revealed the following information: (1) Two general
types of fractures are seen in the field; those that cut through strata disregarding lithologic
boundaries, and those that are confined to individual beds; (2) well-developed fracture pat¬
terns are not confined to limestone sections only; they are also well-developed in the shales;

(3) fracture density increases from a few fractures per tens of yards to many fractures per
yard in the proximity of major faults, thus providing a reconnaissance tool in field studies;

(4) the NE fracture trend of the Balcones Fault Zone is dominant in the carbonate sections.
Most shales reflect this trend, but also show a well-developed and sometimes dominant NW
fracture trend. Many of these NW trending fractures appear to be genetically unrelated to
the Balcones faulting episode.

INTRODUCTION

The purpose of this study is to investigate the attitude of the fracture patterns
in the vicinity of the Balcones Fault Zone in north central Texas. This knowledge
will prove advantageous in future engineering geology, structural and tectonic
studies, and in our understanding of the tectonic history and structural evolution
of the area.

GEOLOGIC SETTING

Location and Previous Work

The study area is centered in the Waco urban region and encompasses McLennan
County, Texas and its immediate vicinity (Fig. 1).

Previous works discussing Balcones faulting are numerous. Those that proved
to be most useful in this study include that of Hill (1901), Foley (1926), Link
(1929), Melton (1934), Weeks (1945), 0. T. Hayward (1957), Burket (1965),

Received for publication: February 28, 1977.

The Texas Journal of Science, Vol. XXIX, Nos. 3-4, December, 1977.

188

THE TEXAS JOURNAL OF SCIENCE

k

"A'\

\

Figure 1. Location Map.

BALCONES FAULT ZONE

189

Goodson(1965), Font (1969), Hudson (1972), Font (1973, 1976), and C. Hayward
(1976).

Structure and Stratigraphy

McLennan County is underlain by eastward dipping beds of the Gulf Coastal
Plain. The typical homoclinal structure of the plain is disrupted in the Waco area
by the Balcones Fault Zone, a zone of normal faults which extends from west of
Uvalde to the Dallas area (Fig. 1). Major faults in the area trend north-northeast
and exhibit vertical displacements up to 300 ft. The faults are commonly down-
thrown to the east, although some are downthrown to the west. Precise dating
of the faulting in McLennan County is impossible but there is no evidence of dis¬
placement along any of the faults since the Middle Pleistocene (Burket, 1965).
The complexity of fault and fracture patterns in the study area as seen from high
altitude aerial photographs is shown in Fig. 2. The stratigraphic relationships,
depositional history and maximum thickness of the Cretaceous strata in the region
of investigation are summarized in Table 1.

SELECTED REVIEW OF PROPOSED MECHANISMS FOR

BALCONES FAULTING

The Balcones Fault System was named by R. T. Hill. Hill (1901) suggested
that the faulting probably resulted from adjustments of the Cretaceous strata
to the sope of the pre-Cretaceous surface. Foley (1926) concluded that the fault¬
ing resulted due to tensional stresses created by subsidence in the East Texas
Basin. Link (1929) proposed that faulting resulted from tensional stresses caused
by differential settlement of sediments over and around pre-Cretaceous topo¬
graphic features. Weeks (1945) separated the Balcones, Mexia, and Luling fault
zones into 3 separate fracture systems. 0. T. Hayward (1957) proposed that Bal¬
cones faulting resulted from the inclination of a deep salt “glide plane” causing
Gulf Coast Mesozoic and Cenozoic rocks to be fractured over the Ouachita fold-
belt by normal gravitational forces. He postulated that the Balcones faults might
not penetrate the pre-Cretaceous surface. Goodson (1965) believed that the
Ouachita foldbelt was responsible for the Balcones faulting, but never stated if
the foldbelt was active or passive in this respect. Hudson (1972) described the
complexity of aerial photo alinements in north central Texas and concluded that
fracturing resulted from the action of “multiple geologic processes” throughout
Cretaceous and early Tertiary time. C. Hayward (1976) has shown that tectonic
activity in the area decreased continuously from earliest Cretaceous to pre-George¬
town deposition. Beginning with Georgetown deposition, tectonic activity in¬
creased steadily through Austin time. Hayward (1976) has also suggested that
the concept of a Cretaceous hinge line may be in error, since deposition apparently
prograded into the East Texas Basin.

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THE TEXAS JOURNAL OF SCIENCE

Figure 2. Alinements as seen on high altitude aerial photographs, (after Hudson, 1972).

FIELD ANALYSIS OF FRACTURE PATTERNS AND CONCLUSIONS

In 1973, Font initiated a systematic study of fracture patterns in some of the
shale sections in the Waco area. He recognized 6 prominent fracture trends in the
Del Rio Clay (Font, 1973, 1976). Field investigations by the authors of this study
have led to the following conclusions regarding fracture patterns in McLennan
County :

1. Fractures that cut through lithologic boundaries are related to the Balcones
faulting episode. Some of the fractures confined to individual beds are un¬
doubtedly younger and unrelated to the faulting episode. Some may be the
result of residual stresses or of swelling and heaving of the clay-shales.

BALCONES FAULT ZONE

191

Series

TABLE 1

Stratigraphic Relationship, Depositional History and Maximum Thickness
of the Cretaceous Strata in the Local Area.

Maximum

Maximum Probable
Local Thickness

Formation Depositional Thickness at one time

Group and Symbol History in feet in feet

Gulf

Taylor

Austin

Eagle

Ford

Taylor

Marl

Kta

Austin

Chalk

K

South

Bosque

Shale

Ksb

Deposited dur- 250

ing marine trans¬
gressions and
regressions.

Deposited dur- 250

ing marine trans¬
gression with pos¬
sible fluctuations
of the strandline
Unconformably un¬
derlies the Kt .

Deposited in a 160

neritic marine
environment with
poor circulation.
Unconformably un¬
derlies the K .

1170

295

160

Woodbine

Comanche Washita

Lake

Waco

Fm.

Klw

Pepper

Shale

Kpe

Buda

Limestone

Kbu

Del Rio
Clay

Kdr

George¬
town Lime¬
stone

Kge

Deposited in a la- 80 145

goonal environment.

Conformably under¬
lies the Ksk*

Deposited in brackish 70 100

environment. Uncon¬
formably underlies
theKiw

Nearshore marine 2 35

deposit. Almost
completely eroded
locally prior to depo¬
sition of K .

pe

Marine regression and 85 85

transgression. Re¬
stricted environment.

Conformably underlies

theKbu'

Shallow marine 210 210

deposit.

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THE TEXAS JOURNAL OF SCIENCE

2. Locally well-developed fracture patterns in shales suggest brittle behavior
during deformation. This may be related to high strain rates or to a high degree
of consolidation before the fracturing took place.

3. Locally well-developed fractures (antithetic, synthetic, feather, etc.) in the
vicinity of faults indicate sense of shear and active and passive members. Some
locally dense fractures are undoubtedly a result of local stress concentrations
related to the curvature of fault planes.

4. Fracture density increases drastically in the proximity of major faults. Fracture
spacing can increase from a few fractures per tens of yards to many fractures
per yard. This increase in fracture density can be used successfully as a recon¬
naissance tool in field studies.

5. The NE trend of the Balcones Fault Zone is dominant in limestone sections.
The NE trend is reflected in all shales, but a well-developed and sometimes
dominant NW trend is recorded in these (Figs. 3-8). The prominent NW frac¬
ture trend exhibited by the shale sections appears to be unrelated to the Bal¬
cones faulting episode. A great number of the NW trending fractures are short
and abut against longer, more prominent NE trending fractures. This indicates
that the NW trending fractures are younger than the NE set. In cross-section,

Figure 3. Fracture patterns in the Taylor
Marl (based on 179 fractures).

Figures. Fracture patterns in the South

Bosque Shale (based on 135
fractures).

Figure 4. Fracture patterns in the Austin
Chalk (based on 794 fractures).

Figure 6. Fracture patterns in the Lake
Waco Formation (based on 145
fractures).

BALCONES FAULT ZONE 1 93

Figure 7. Fracture patterns in the Del Rio Figure 8. Composite diagram showing frac-
Clay (based on 267 fractures). ture patterns in north central

Texas (based on 1525 fractures).

NE trending fractures extend through large portions of the shale sections and
appear to be closely related to local faults. In contrast, the NW trending
fractures are largely confined to individual beds and do not penetrate deeply
into the shale sections. These observations lead to the conclusion that most of
the NW trending fractures are probably a result of local stresses generated by
the swelling and heaving and subsequent contraction of the bentonitic, smectite-
rich clay shales.

6. The complexity of the fracture patterns and their modification across major
lithologic boundaries indicate the necessity of further, more detailed studies.

7. Lack of outcrops of Pepper Shale and Buda Limestone have not allowed the
mapping of sufficient fractures to make any concrete statements. As for the
Georgetown Limestone, practically no fractures are recognized in this unit in
the study area.

ACKNOWLEDGEMENTS

The authors express their gratitude to all of those students who helped in the
field measurements of faults and fractures. Thanks are expressed to Art Bishop and
Don Little for assistance in the field. Special thanks are given to Dr. G. A. Morales
for reviewing this manuscript. Our deep appreciation goes to Mrs. Viola Shivers
who typed the manuscript.

LITERATURE CITED

Burket, J. M., 1965-Geology of Waco in urban geology of Greater Waco-Part I. Baylor
Univ. Geol. Stud. Bull., 8:1.

Foley, L. L., 1926-Mechanics of the Balcones and Mexia faulting. Amer. Assoc, of Petrol.
Geol. Bull., 10:1261.

194

THE TEXAS JOURNAL OF SCIENCE

Font, R. G., 1969-Engineering geology of the Greater Waco area. Unpub. M.S. thesis, Baylor
Univ.

- , 1973-Engineering geology study of the instability of the South Bosque Shale

and the Del Rio Clay in the Waco area. Unpub. Ph.D. dissertation, Tex. A&M Univ.

- , 1976-Relationship between the geologic history and engineering properties of

two cretaceous shales. Tex. J. of Sci., 27(2) : 267 .

Goodson, J. L., 1965 -The Balcones Fault Zone, central Texas. Unpub. senior thesis, Baylor
Univ.

Hayward, C., 1976-Structural evolution of the Waco region. Unpub. senior thesis, Baylor
Univ.

Hayward, O. T., 1957-Structural significance of the Bosque Escarpment, McLennan County,
Texas. Unpub. Ph.D. dissertation, Univ. of Wisconsin at Madison.

Hill, R. T., 1901 -The geography and geology of the Black and Grand Prairies, Texas. U.S.
Geol. Surv., 21st Ann. Rept., Pt. 7, 666 pp.

Hudson, P. C., 1972— Interpretation of alinements visible on aerial photographs of central
Texas. Unpub. M.S. thesis, Baylor Univ.

Link, T. A., 1929 —En echelon tension fissures and faults. Amer. Assoc. of Petrol. Geol . Bull.,
13:627.

Melton, F. A., 1934-Fracture systems in central Texas. Univ. of Tex. at Austin Bull., 3401:118.

Weeks, A. W., 1945-Balcones and Luling and Mexia Fault Zone in Texas. Amer. Assoc, of
Petrol. Geol. Bull., 38:306.

LATE PLEISTOCENE POLLEN AND SEDIMENTS: AN ANALYSIS
OF A CENTRAL CALIFORNIA LOCALITY

by ERIC W. RITTER and BRIAN W. HATOFF1

United States Department of the Interior,

Bureau of Land Management, 1695 Spruce St.,

Riverside, CA 92507

ABSTRACT

Palynological and sedimentary analyses were conducted on samples obtained from late
Pleistocene bog sediments and contiguous-superimposing glacio-fluvial silt loams. These
sediments were exposed during gravel quarrying operations just east of Sacramento, Cal¬
ifornia, at the Teichert site. The bog sediments may represent the remnant of an oxbow
lake of an ancient (abandoned) channel of the American River.

Uranium and actinium series dates from faunal remains in the overlying and surrounding
sediments indicate these deposits are about 100,000 years in age (upper Riverbank Form¬
ation).

Reconstructions from the pollen record, faunal remains, and macroscopic plant frag¬
ments suggest local bog and oak-riparian vegetation and regional pine parkland vegetation.
Climate was apparently cooler than present. This trend is in accordance with a proposed
cold climatic period occurring about this time, and these deposits probably indicate cor¬
respondence with the Mono Basin glaciation of the Sierra Nevada. Stratigraphic data tend
to support the hypothesis. The surrounding and overlying silt-loam layers of the upper
Riverbank Formation were apparently deposited during late glacial times by glacial outwash.

Comparisons with modern surface and near- surface pollen from related locations in the
Valley and adjoining Sierra indicate a lowering of present vegetation belts along the western
Sierra flank 100,000 ya by about 600 to 750 meters.

INTRODUCTION

As an adjunct to recent radiometric dating of Quaternary soils and sediments
(Hansen and Begg, 1970), the opportunity was presented to interpret paleo-
environments by palynological analysis of bog sediments found near Sacramento,
in central California. This investigation focused on a late Pleistocene bog ap¬
parently formed in an oxbow lake of an ancestral channel of the American
River of Riverbank Formation age (e.g. Shlemon, 1967).

The bog and adjacent sediments, exposed during gravel quarrying operations,
have been called the Teichert site by Hansen and Begg (1970). The bog is about
10 kilometers (6.2 mi) east of central Sacramento, California (Fig 1).

Present Address: United States Dept, of the Interior, Bureau of Land Management, 1050 East
William St., Carson City, NV 89 701.

Accepted for publication: September 11, 1975.

The Texas Journal of Science, Vol. XXIX, Nos. 3-4, December, 1977.

196

THE TEXAS JOURNAL OF SCIENCE

EXPLANATION

Rw Riverwash (Post Modesto chan¬
nel deposits)

Ha Hanford very fine sandy loam and
loamy fine sand (Upper Modesto
Formation) i

Ho Honcut loam (Lower Modesto
Formation)

Pe Perkins gravelly loam (River-
bank Formation)

Sa San Joaquin loam (Riverbank
Formation)

Figure 1. Map of study area east of Sacramento, California. Inset map shows more
precise site location and local geology (after Hansen and Begg, 1970).

Sampling Mechanics

A series of samples was secured for analysis from the exposed cut. Collecting
procedures followed those suggested by Mehringer (1967). In short, the wall
was cleaned back about 30 cm, then sample locations were marked with a
nail and flagging. Samples of about 200-300 grams each were removed by trowel
in peds from bottom to top. Samples were placed in Nasco Whirl-Pak ster¬
ilized plastic bags. Subsequently each sample was thoroughly mixed before ex¬
traction.

Pollen was sampled from 2 loci in the bog sediments: “A” near the central
portion, and “B” near the apparent northern edge (Fig 2). Three samples also
were collected from the overlying silt loams, but apparently oxidation in the
alkaline sediments destroyed the pollen. In contrast, the bog samples were rich
in pollen, wood and other organic material.

The sampling was designed to provide a general overview of the local and
regional vegetation (and possibly associated climate) of this central California
locality during a specified time range of the late Pleistocene. Samples were
taken from the relatively thin and narrow black stratum to (1) acquire pollen
spectra of the early, middle and late “bog” center deposits and (2) to deter-

LATE PLEISTOCENE POLLEN AND SEDIMENTS

197

mine intra-bog differences between peripheral and center bog pollen. We hoped
that these samples would provide a handle on the relative ranges of variance in
the deposit pollen. Our aim was not to determine major climatic or vegetational
changes. Our sampling is viewed as only a preliminary step toward better defin¬
ing our working hypotheses on late Pleistocene vegetation in this region; to en¬
able us to ask more explicit and informative questions.

DISTANCE 01 2 3 48 49 50 51 5? (meters)

• Nun s e I I color notations 0 - d i y M - mo ■ s t

Figure 2. Stratigraphic section of the Teichert Site showing location of pollen samples
and radiometrically dated vertebrate fossils and wood (after Hansen and
Begg, 1970).

Laboratory Mechanics

In following Mehringer (1967), 6 basic steps are required to isolate the fossil
pollen from the sediment matrix: (1) HCL swirl for removal of carbonates;
(2) soaking and boiling in HF to remove silicates; (3) HN03 treatment to oxidize
organic colloids; (4) acetolysis to remove cellulose; (5)NaOH boiling to destroy
humates; and (6) staining the residue with Sulfannin 0 and mounting on a slide
with glycerine. Two hundred pollen grains were counted for each sample. We
concur with Martin (1963) who indicates this is a significant number statistically
since he found only a difference of 4% when compared with slides counted to
2000 grains.

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Stratigraphy of the Teichert Site

The upper 2 meters consist of modern overburden of variable color and
texture and the San Joaquin soil, a brown silty loam with an iron-silica cemented
hardpan (Al, B2t, Clm and C2 of Fig 2). Pale brown and light brownish gray
silty clays intermingled with 2 discontinuous lime-silica cemented hardpans
extend some 2.5 m below this soil (IIAllb, II Al 2b, and IlClrnb of Fig 2).
The next lower 3 to 4 meters include light gray silty clay loam underlain by a
light gray silt loam (IIIC2; IIIC3 of Fig 2) both of which contain a faunal as¬
semblage of Rancholabrean age (for a detailed list see Hansen and Begg, 1970).

The bog deposit occurs approximately 7.5 to 9.0 m below the surface (IVC5
of Fig 2). Underlying the bog sediments are unsorted pebbles, cobbles and
'sands, portion of an ancient gravel bar (VC5 of Fig 2), derived from granitic
and metamorphic terrain of the higher Sierra. The bog is about 75 m wide, while
the length is presently unknown. Macroscopic plant remains from the bog sedi¬
ments include wood from Douglas fir (. Pseudotsuga menziesii), cottonwood
( Populus sp.), sycamore ( Platanus racemosa) and willow (Salix sp.) as well as
unidentified leaf impressions and other plant remains.

Leopold (1964) suggests that detailed study of macroscopic plant remains
in peat bogs is useful for indicating local flora and floral succession. These
macroscopic plant remains suggest a wet, cool riparian location, while the re¬
mains of many mammals, a goose, pond turtles, frogs, snakes and fish in the im¬
mediately overlying sediments (5 .6-7.5 m depth) indicate the location remained
wet with much aquatic life and may also have provided a watering place for non-
aquatic mammals. Recovery of Bison , Camelops, Equus, and Mammathus,
and possibly several rodent species indicate nearby or later grasslands. Sylvilagus
sp. and Odocoileus sp. appear discrepant but could be accomodated by a lake-
shore or riparian woodland zone in the vicinity. The overlying fauna seemingly
reflects a continuance of wet conditions, but it also suggests that the proposed
pine oak-grass community was replaced by a prairie similar to the historic one,
perhaps during the last interglacial (cf. Suggate, 1974).

While we cannot in toto rule out the possibility that some macroscopic plant
and animal remains and pollen floated in from the high Sierra, we believe such
contamination was minimal as inferred from the evidence presented below.

Dating

Age determinations have been discussed in some detail by Hansen and Begg
(1970). The Rancholabrean fauna such as found in the sediments overlying
bog deposits for the most part probably began to die out from 10,000 to 30,000
years ago.

Data of uranium and actinium series nuclides within the fossils from the
Teichert site have provided more accurate age calculations. Five uranium series
and 3 uranium-actinium series dates were obtained on the fossils (Hansen and
Begg, 1970). The average age of these 8 dates is 98,000 ± 12,000 years BP.

LATE PLEISTOCENE POLLEN AND SEDIMENTS

199

This age is supported by a radiocarbon date of greater than 38,300 years B.P.
for wood taken from the bog (Shlemon and Hansen, 1969). This age is also
consistent with the 30,000 to 600,000 yr old Rancholabrean faunal assemblage
contained within the sediments.

The local geomorphology has been analyzed by Shlemon (1972). The basal
gravels at this locality (VC5, Fig 2) are attributed to the upper Riverbank Form¬
ation (Shlemon, 1972). These gravels fill a well-defined channel. Shlemon (1972)
notes that “There is not disconformity or buried soil separating the channel
gravels from the overlying radiometrically dated sediments. Because of this
close stratigraphic proximity, inferentially, the gravels were laid down by an an¬
cestral American River, perhaps 1 10,000 yrs ago, a time of world-wide cooling
as deduced from deep sea cores” (Emiliani, 1961; also see Broecker, 1965). The
fine-grained sediments are apparently glacial flour derived from late glacial
outwash in the Sierra Nevada (Arkley, 1962; Janda, 1963; Wahrhaftig and Bir¬
man, 1965; and Shlemon, 1972).

Dalrymple (1964) has dated interglacial basalt underlying Tahoe (Altonian
substage of the Wisconsin Stage) glaciation material in the nearby Sierra by
K-Ar as about 60,000 to 90,000 yrs BP. On this evidence it appears the bog
deposit antedates the Tahoe glaciation. It may correlate with Sharp and Birman’s
(1963) Mono Basin (early Wisconsin) glaciation or Birkeland’s (1964) equivalent
Donner Lake Glaciation (compare Birkeland, et al, 1971). It also may compare
with one of the last “interglacial” cooling periods about 100,000 yrs B.P. as
discussed by Suggate (1974). Various other workers have found evidence of
such “interglacial” cooling trends (cf. Dansgaard, et al, 1972; Emiliani, 1972;
Hayes and Perruza, 1972; Kennett and Huddlestun, 1972; McIntyre and Ruddi-
man, 1972; Richmond, 1972; and Sancetta, et al, 1972).

Pollen Analysis

At Locus A (Fig 3 — depths are from upper bog contact with stratum IIIC3)
there is a shift through time in pine pollen from 9% near the base to 28% near
the upper limits, whereas the oak pollen decreases from 47% to 32%. Nymphaea,
the water lily, increases from 1% in the bottom sample to 16% in the middle
sample and back to 1% near the top. Compositae show a decrease from 19%
in the lowest sample to 11% in the middle, increasing to 14% near the top of
the bog sediments. Graminae (grass); Cyperaceae (sedge), TCT (Taxodiaceae,
Cupresaceae, and Tacaceae, probably mostly Libocedrus--ce dar and perhaps
some Juniper us — juniper, Sequoiodendron — redwood, Taxus — yew, and Cupres-
sus-- cypress), cheno-ams and the other conifers ( Abies — fir, Pseudotsuga —
Douglas fir, etc.) and deciduous trees (Populus — cottonwood, Salix — willow,
Platanus — sycamore, etc.) remain fairly stable or have insignificant representation.

Douglas fir ( Pseudotsuga menziesii), although present in fair amounts macro-
scopically in the bog sediments, is poorly represented in the pollen spectra and
has been lumped in the other conifer category (Fig 3). Leopold (1964) indicates
such pollen is usually under-represented, which is a possible explanation.

Figure 3. Comparative pollen diagram of the Teichert Site and select east-central Calif¬

ornia locations.

All the categories in Locus B (Fig 4) show insignificant variation. Quercus
pollen dominates with 45 and 43% from the lower and upper sample respectively,
while Pinus is present in 20 and 18% frequency. All others, except Nymphaea ,
are similar in percentile to Locus A.

Nymphaea show no significant variation with only 4 and 5% represented at
Locus B. The percentile does not approach the 16% of Nymphaea found in the

EAST-CENTRAL CALIFORNIA

LATE PLEISTOCENE POLLEN AND SEDIMENTS

201

middle sample of Locus A. It is probably that pollen variation between the 2
loci is explainable by intra-bog locational differences and minor variation in
pollen dispersal and filtration as discussed by Janssen (1966), Tauber (1967) and
others. A possible explanation for some of these differences, based on a study
by Maher (1970), is that along the edges of certain lakes vesiculate pollen such
as pine accumulates at a greater density than in the center because even light
winds can push the floating pollen to the shore. This would account for the
higher overall percentages of pine pollen in Locus B (probably bog-lake margin)
compared to the lower 2 samples at Locus A. Maher (1970) suggests oxidation-
destruction of vesiculate pollen at lake margins is possible under certain condi¬
tions (cf. Havinga, 1971). Whether this occurred at Locus B is uncertain, but its
potential significance appears minimal owing to the wet anaerobic acidic soil
conditions favorable for preservation at the bog locality. The high pine pollen
percentage from the upper sample of Locus A may be representative of the pen¬
ultimate bog environment when free standing water presumably was low, an in¬
crease in local or regional pine, or a change in the filtration- dispersion rates.

The higher percentage of Nymphaea in the middle of the deposit seemingly
is less difficult to explain. Janssen (1966) indicates that most of the lowland
herbs are best represented in the local pollen rain. It is noteworthy with respect
to later discussion that an over-representation of local pollen will distort the
regional pollen rain representation making inferences on climatic (vegetation)
changes difficult (Janssen, 1966). Apparently a high number of water lilies
were present during the bog climax, about midway in its formation. According
to Dapples (1959), during the genesis of bog lakes the false bottom extends
for some distance from shore into open water before terminating with a zone
of water lilies and similar floating hydrophytes. This false bottom gradually ex¬
tends lakeward until it covers the pond. Debris then builds up and the pond mat
increases until the bog is eventually terminated. Adam (1966), interpreting pol¬
len spectra from Osgood Swamp in the central Sierra Nevada, attributes an in¬
crease in Nymphaea to cool (Neoglacial) conditions and increased productivity
of the lake-swamp. A similar cool period with increased bog productivity at
the Teichert site is also a possibility.

Several problems are evident in the palynological analysis of the Teichert
site. Is the fossil pollen an adequate measure of regional vegetation and climate?
What percent of local, regional and extra-local pollen is present? How much
contamination has occurred from fluvial transported pollen?

Potter (1964), Janssen (1966), Tauber (1967), Havinga (1971), and many
others mention such basic problems in palynological-environmental recon¬
struction as (1) amount of pollen production/species, (2) distance of pollen
transport, (3) mechanisms of initial and secondary deposition, (4) differential
dispersal and filtration of pollen rain, (5) relative preservation of different species
in various sediments, and (6) the relation of pollen composition to features of
climate, landform, and vegetation. These problems have for the most part been

202

THE TEXAS JOURNAL OF SCIENCE

considered and it is our conclusion that only a general reconstruction of the lo¬
cal and regional vegetation about 100,000 yrs ago is possible.

Havinga (1971) has noted that where wet anaerobic conditions prevail good
pollen preservation is guaranteed. Haynes (1964) has indicated that probably the
best opportunity for obtaining a pure pollen assemblage is furnished by sedi¬
ments representing semi-permanent or intermittent ponding along the stream
channel as inferred at the Teichert site. Bogs themselves have long been used
as prime locations for paleoecological reconstruction because of good preserva¬
tion and relative lack of contamination. If this bog constitutes the remnant of
an ancestral oxbow lake, this feature was fairly effectively sealed off from mech¬
anical intrusion of new water, except perhaps during rare extreme flooding. Sev¬
eral small thin lenses of light gray silty loam possibly derived from such flooding
were, in fact, noted in the bog sediments, but pollen samples were not collected
from these thin sections nor near contacts. Otherwise the bog sediments appear
“undisturbed”. It must be noted, however, that a single marine foraminifera was
identified from the bog sediments by personnel of the Department of Geology,
University of California, Davis. It is possible this foraminifera was redeposited
from older sediments. Scanning the pollen slides revealed no other foraminifera
and this may represent an isolated specimen.

Tauber (1967) has criticized the idea a pollen rain represents the regional
pollen production. For instance, in closed deciduous forests pollen carried
through the trunk space, above the canopy and in the rain will affect dispersal
as well as distance gradients, and pollen carried through the vegetation will be
subject to differential filtering. In fact, Tauber (1967) has said“...it is also ob¬
vious that various dispersion processes will have a great effect on the deposition
of pollen on lake and bog surfaces and that information on these matters is es¬
sential for the interpretation of pollen diagrams.” Tauber (1967) concludes
that in minor lakes and bogs (perhaps like that at the Teichert site), where the
trunk-space component is likely to dominate, the area effectively represented
will be much smaller than is usually thought. Tsukada (1958) reports that in
a forested region most of the pollen precipitation comes from an area within
a few kilometers.

It appears the pollen from the Teichert bog is a fair representative of the
local late Pleistocene vegetation, that is the bog and surrounding vegetation with¬
in several kilometers. The spectra have important implications to late Quater¬
nary events in the Sierra Nevada and Central Valley.

To better interpret the pollen results it is worthwhile to compare the Teichert
site pollen with modern pollen spectra obtained from surface- sample transects
and recent archaeological sites. Such samples are assumed by the researchers
to be representative of the local and regional pollen rain/vegetation.

Adam (1967) conducted a pollen transect (Fig 3) from the Central Valley
over the Sierra crest at Carson Pass which provides significant comparisons.

Adam’s (1967) Central Valley sample (Fig 3) compares closely with that
of surface samples from the Blodgett Site (CA-Sac-267) (Fig 3; Germeshau-

LATE PLEISTOCENE POLLEN AND SEDIMENTS

203

sen, 1969). This archaeological midden is located 7 kilometers (4.3 mi) east of
the Teichert site on the first terrace above the Consumnes River near Slough-
house, California. Each spectrum is representative of the modern open grass¬
lands with scattered oak. This is also the present vegetation pattern at the
Teichert site locality and extends to the present riparian belt along the American
River to the north (Fig 1). As the elevation increases in Adam’s transect ascend¬
ing the Sierra, the Pinus pollen significantly rises from 12% in the Valley to 30%
at 760 m (2500 ft), to 57% at 1520 m (5000 ft), increases slightly to 60% at
2280 m (7500 ft), and finally to 90% at 3.040 m (10,000 ft). Quercus, Graminae,
and Compositae, on the other hand, correspondingly decrease, with the latter
2 not represented at 1500 m and higher. TCT and other conifers, notably Abies
(fir), oscillate, with TCT increasing to its highest frequency (28%) at 1500 m
before dropping off, and the other conifers peaking at 2280 m with 30%.

Modern pollen rain samples from the Spring Garden Ravine archaeological
site (CA-Pla-101) in the Sierra upper foothills (elevation 760 meters or 2500
ft) 60 kilometers (42 mi) east-northeast of the Teichert site (Fig 3) correlate
well with the present pine-oak woodland (Matson, 1970). This site adjoins a
bog. Subsurface samples radiometrically dated at 3350 ± 110 BP. (GaK-2246)-
still within the present day climatic regime (Medithermal)-suggest vegetation
was a lightly wooded oak grassland with or without low pollen-producing chap¬
arral species. Changes in pollen spectra/vegetation from the older pattern to the
modern pattern can be attributed to cessation of aboriginal practices of burn¬
ing (Matson, 1970). The results of the modern pollen rain are thought by Matson
to better reflect natural vegetation conditions during the Medithermal.

What is believed diagnostic in comparing results of these other studies with
those of the Teichert bog is the similarity at about 750 m of the pine and grass
pollen. High Quercus values at the Teichert site (compared to the samples ob¬
tained from the higher elevation locations) we attribute to local riparian-like
growth not present at either Adam’s or Matson’s test locations. Adam (1967),
in fact, notes that neither riparian nor montane-chaparral communities are well
represented in his transects.

INTERPRETATIONS AND EXPLANATIONS

It appears that the local and regional vegetation around the Teichert locality
at the time of bog formation, about 100,000 yrs ago, differed from present or
historic Valley vegetation, a reflection of glacio-climatic conditions in the ad¬
jacent Sierra Nevada. The 98,000 yr old average age of these upper Riverbank
Formation sediments corresponds closely to a cold period which occurred about
this time (cf. Broecker, 1965). We assume that vegetation zones of the Sierra,
as noted by Adam (1967), have retained their integrity and relative positions
during vertical movements imposed by Pleistocene climatic changes, that lag was
minimal, and that glacial climates depress vegetational belts relative to their

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interglacial positions, as do minor climatic fluctuations such as have occurred
during the Holocene. Localized conditions such as riparian belts and bogs add
complexity to this concept.

The changes within the bog are a result of vegetational responses to evolving
bog conditions and perhaps a minor shift in climate (e.g .Pinus increases at Locus
A). At the end of bog life cooler conditions may have prevailed than during the
beginning, but it may also be that during the early stages of bog formation winds
moved the vesiculate pine pollen from the center of the oxbow lake (Locus A)
to the margin (Locus B). In either case, a higher pine percentage is found in the
bog sediments than prevails in surface samples in the area today. The silt loam
layers superposing the bog sediments are characteristic of glacial flour, an indica¬
tion of rapid aggradation of glacio- fluvial outwash and hence a glacial recession
with overloading of the ancestral American River. As early as 1885 and 1889,
Russell demonstrated that pluvial events in western North America could be
linked stratigraphically to local glaciation in the Sierra and elsewhere. The
Teichert bog was thus probably formed just prior to deposition of the overlying
sediments during a cold episode corresponding to high Sierra glaciation (i.e., Mono
Basin) or part thereof.

If the present vegetation in the Sierra Nevada can be extrapolated to the bog,
then a lowering of the present vegetation belts by about 600 to 750 meters
(2000 to 2500 ft) is postulated for the period of bog formation. Local channel
conditions during this period apparently favored a heavy oak stand and local
aquatic plants such as sedge and water lily. The aquatic plants decreased as the
bog lost its effective hydrophilous attributes and became filled. The riparian
vegetation was apparently symbiotic with a surrounding pine parkland. The tree
remains of Douglas fir, which presently is found on moist slopes of the west
central Sierra below 1500 m (5000 ft) and above 300 m (1000 ft), seemingly
add further confirmation.

Martin (1964) utilizing data from 9 pollen spectra thought to be of full
glacial age in the American Southwest, notes that to find a modern pollen count
similar to the full glacial pollen spectra it is necessary to sample lake sediments
or soils 800 to 1200 m (2625 to 3937 ft) higher in elevation than the fossil local¬
ities. Furthermore, Martin (1964) indicates the pollen type most likely to reveal
the full glacial change is pine. It is believed that these results from the California
Central Valley are analogous to those in the Southwest. While this model is still
conjectural, empirical evidence is accumulating, and further effort should be
made to find correlative or non-correlative data. Along this line it is worth men¬
tioning that a few undated relict soil structures possibly characteristic of cooler
areas have been found along the east side of the Central Valley. These as yet un¬
reported features include probable stone nets and stripes. However, Shlemon,
Begg and Huntington (1973) have noted that some such structures along the
Sacramento Valley edge are structurally controlled.

This study reaffirms the importance of interdisciplinary cooperation needed
to reconstruct paleoenvironments.

LATE PLEISTOCENE POLLEN AND SEDIMENTS

205

ACKNOWLEDGEMENTS

We would like to acknowledge the comments, criticisms, and help put forth
by Dr. Delbert True, Dr. Robert Kautz and Mr. Peter Schulz of the Department
of Anthropology at the University of California, Davis; by Dr. Roy Shlemon of
the Department of Geography, Mr. Eugene Begg of the Department of Soils and
Plant Nutrition and Dr. Jack Major of the Botany Department of the same
institution. The authors, however, assume full responsibility for the article’s
contents.

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VEGETATION TYPES OF CHAMBERS COUNTY, TEXAS

by P. A. HARCOMBE

Biology Department, Rice University , Houston 77001
and J. E. NEAVILLE

U. S. Fish and Wildlife Service, Angle ton 77515
ABSTRACT

The vegetation of Chambers County, Texas, is described and mapped on the basis of field
reconnaissance. Eight vegetation types are recognized. Moisture availability, salinity, and soil
texture are identified as probable determinants of the pattern of vegetation distribution.

INTRODUCTION

Chambers County, Texas, lies just inland from the Gulf of Mexico on the north
and east shores of Galveston Bay, and in many respects, it is typical' of the coastal
environment of eastern Texas and western Louisiana. Because Chambers County
contains a wide range of vegetation— from prairie to forest— it is of interest to de¬
scribe the vegetation pattern and to attempt to identify the contributing environ¬
mental factors.

Documentation of the present distribution of vegetation types seems particu¬
larly worthwhile, given that Chambers County is located between two rapidly
growing metropolitan centers (Beaumont— Port Arthur and Houston— Galveston),
and will undergo change with increasing industrial and residential development.
Thus, the data reported here are intended to provide both a baseline for evalua¬
tion of the impact of development and a regional overview for more intensive
studies of coastal ecosystems.

This work is part of a larger attempt to describe the natural environment of
Chambers County and to develop mechanisms facilitating land-use decision-making
(Rowe, et al., 1974). Primary emphasis here is description of existing native (i.e.,
not cultivated) plant assemblages.

Three sources have been used: published descriptions of the vegetation of the
Gulf Coast, published information on the effects of environmental factors on vege¬
tation, and personal observation and field reconnaissance. Quantification of vege¬
tation composition was outside the scope of the present study.

Site

Chambers County is a broad, flat, poorly drained plain, of about 221 ,000 ha
with slopes less than 3% and elevation mostly less than 11 m. It is composed of

Accepted for publication: December 3, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 3-4, December, 1977.

210

THE TEXAS JOURNAL OF SCIENCE

Pleistocene and Holocene deltaic and fluviatile deposits (Rowe, et al., 1974;
USDA, 1976). The soils are predominantly clays and silt-loams of the Beaumont-
Morey-Lake Charles and Morey-Anahuac-Frost Associations. The climate is warm
and wet with 1309 mm average annual rainfall and a growing season of approxi¬
mately 261 days (USDA, 1976). f

Chambers County lies entirely within the Gulf Prairie and Marsh Vegetation
Area of Texas (Gould, 1969), which is characterized by “level grassland, low flat
woodlands, especially near the streams, swamps and . . . marshes” (Correll and
Johnston, 1970). The types of vegetation mapped by Kiichler (1964) included
Bluestem-Sacahuiste Prairie, Oak-Hickory-Pine Forest, Southern Floodplain Forest,
and Southern Co rdgrass Prairie. Fisher, etal., (1972, 1973) mapped 12 assemblages
of rooted vegetation, including 4 types of marsh, a single prairie grassland type,
2 types of swamp, 3 types of woodland, and a successional forest type. The Soil
Conservation Service recognized 7 types of prairie and marsh in Chambers County
(USDA, 1976); the U.S. Fish and Wildlife Service recognized 8 distinct marsh zones
(U. S. Department of Interior, 1952).

These publications, plus the earlier surveys of Tharp (1926, 1939), constitute
most of the available information on the vegetation of Chambers County, and
they were used as the basis for reinterpretation of the vegetation. A reinterpretation
was deemed necessary because none of the above works comprehensively describe
the vegetation. Kiichler’s work (1964) is very general, Fisher, et al., (1972, 1973)
erred in assemblage descriptions (see below) and in setting assemblage boundaries
(Solomon and Smith, 1973); and the U.S. Government information does not treat
all assemblages uniformly.

To compile the new. description, we followed Kiichler (1964) in dividing the
landscape first according to physiognomy (i.e., whether forest, woodland, brush-
land), and second according to species composition (identity of prominent or
predominant species).

Once the landscape had been divided into physiognomic types, each type was
subdivided into assemblages, or groups of species commonly found together. Eight
assemblages were identified (Table 1): floodplain cypress swamp, floodplain hard¬
wood forest, upland oak -pine forest, streamside woodland, bluestem prairie, cord-
grass prairie, fresh marsh, and brackish marsh. They are described in Section II.
For convenience, these assemblages are discussed as discrete entities; actually in
most cases they appear to intergrade with one another.

Vascular plant species lists for the assemblages (Tables 2-5) were compiled from
personal records and from observations in nearby areas (Lohse, et al., 1973). They
are not necessarily complete; they should be regarded as preliminary checklists.
Plant nomenclature follows Correll and Johnston (1970).

A map (Figure 1) was produced by copying forest, prairie, and marsh boun¬
daries from 1:8,000 SCS aerial photographs flown in 1970, and subdividing forest,
prairie, and marsh according to species composition determined by field recon¬
naissance. Environmental factors which appear to strongly influence assemblage
distribution were identified on the basis of field observations.

VEGETATION TYPES

211

TABLE 1

Plant Assemblages of Chambers County.

£

I. Forest

Major Species

Cypress Swamp

bald-cypress, water elm, buttonbush,
water hickory

Floodplain Hardwood Forest

cedar elm, sugarberry, hawthorn, water
oak, sweetgum, green ash

Upland Oak -Pine Forest

loblolly pine, sweetgum, willow oak,
water oak, southern red oak, magnolia

Steamside Woodland

sugarberry, hawthorn, water oak , willow
oak

II. Prairie

Major Species

Bluestem Prairie

little bluestem, brownseed paspalum,
switchgrass, indiangr ass, eastern gamagrass

Cordgrass Prairie

gulf cordgrass, knotroot bristle grass

I II. Marsh

Major Species

Fresh Marsh

common rked, saw grass, cutgrass, big
cordgrass, California bulrush

Brackish Marsh

marsh-hay cordgrass, seashore salt grass,
salt-marsh bulrush, olney bulrush

TABLE 2

Scientific names of plant species referred to in text. (After

Correll and Johnston, 1970, except where authority is cited.)

Alligatorweed - Altemanthera philoxeroides
American beautyberry - Callicarpa americana
Ash, green - Fraxinus pensylvanica
Bald-cypress - Taxodium distichum
Bluestem - Andropogon spp.

Bluestem, Big — A. gerardi
Bluestem, Little - Schizachyrium scoparium
Bluestem, Silver - Bothriochloa saccharoides
Bulrush, California - Scirpus califomicus
Bulrush, Salt Marsh - S. maritimus
Bulrush, Olney - S. olneyi
Bushybeard - Andropogon glomeratus
Button bush - Cephalanthus occidentalis
Carpetgrass - Axonopus affinis
Cattail - Typha latifolia, T. dominguensis
Chinese Tallow - Sapium sebiferum
Common Reed - Phragmites communis
Cordgrass, Big - Spartina cynosuroides
Cordgrass, Gulf - S. spartinae

212

THE TEXAS JOURNAL OF SCIENCE

Table 2 (Continued)

Cordgrass, Marsh -hay - S. patens
Cordgrass, Smooth - S. alterniflora
Crossvine - Bignonia capreolata
Cutgrass - Zizaniopsis miliacea
Eastern Gamagrass - Tripsacum dactyloides
Elm, Cedar - Ulmus crassifolia
Elm, Water — Planera aquatica
Greenbriar - Smilax spp.

Hawthorn - Crataegus spp.

Hickory — Carya spp.

Hickory, Water - C. aquatica

Huisache - Acacia farnesiana

Indiangrass — Sorghastrum avenaceum

Knot-root Bristlegrass - Setaria genicultata

Longtom - Paspalum lividum

Magnolia - Magnolia grandiflora

Needlegrass - J uncus roemerianus

Oak, Basket - Quercus prinus

Oak, Laurel - Q. lauri folia

Oak, Live — Q. virginiana

Oak, Post - Q. stellata or Q. similis

Oak, Southern Red - Q. falcata

Oak, Water - Q. nigra

Oak, Willow - Q. phellos

Palmetto — Sabal minor

Paspalum, Brownseed - Paspalum plicatulum

Paspalum, Seashore - P. vaginatum

Poison Ivy - Rhus toxicodendron

Pine, Loblolly - Pinus taeda

Pine, Longleaf - P. palustris

Pine, Shortleaf — P. echinata

Rattan - Berchemia scandens

Rattle Bush - Sesbania drummondii

Sacahuiste - Spartina spartinae

Savanna panicum — Panicum gymnocarpon

Saw Grass - Cladium jamaicense

Sea Myrtle - Baccharis halimifolia

Seashore Saltgrass - Distichlis spicata

Smutgrass - Sporobulus indicus

Sugarberry - Celtis laevigata

Sweetgum - Liquidambar styraciflua

Switch Grass - Panicum virgatum

Sycamore - Platanus occidentalis

Trumpet Creeper - Campsis radicans

Tupelo - Nyssa aquatica

Vasey Grass — Paspalum urvillei

Willow - Salix nigra

Willow, Water - Justicia lanceolata

Woodgrass - Chasmanthium sessiliflorum, C. laxum

Yaupon - Ilex vomitoria

VEGETATION TYPES

213

TABLE 3

Checklist of Forest Species in Chambers County

Floodplain

Upland

Hardwood

Cypress

Oak-Pine

Streamside

Trees

Forest

Swamp

Forest

Woodlands

Acer negundo - box elder

X

X

A. rubrum - red maple

X

X

Carpinus caroliniana — ironwood

X

X

Cary a aquatica — water hickory

X

X

C. cordiformis - bitternut hickory

C. illinoiensis — pecan

X

X

C. laciniosa - shellbark hickory

X

Celtis laevigata - sugar hackberry

X

X

X

Crataegus brachyacantha - blueberry hawthorn

C. spathulata - littlehip hawthorn

X

X

X

C. uniflora - oneflower hawthorn

X

C. viridis - green hawthorn

Diospyros virginiana — common persimmon

X

X

X

Fraxinus americana - white ash

F. pensylvanica - green ash

X

X

X

Gleditsia aquatica - water honeylocust

X

X

G. triacanthos - common honeylocust

X

X

Ilex opaca — American holly

X

Juglans nigra - black walnut

Juniperus silicicola - southern red cedar

X

X

J. virginiana - eastern red cedar

Liquidambar styraciflua - sweetgum

X

X

X

Magnolia grandiflora - Southern magnolia

X

Melia azedarach - chinaberry tree

Morus alba — white mulberry

X

X

M. rubra - red mulberry

Parkinsonia aculeata - retama

X

X

Pinus taeda - loblolly pine

Platanus occidentalis — American sycamore

X

X

Populus deltoides - eastern cottonwood

Prunus angustifolia - chickasaw plum

X

X

P. mexicana — Mexican plum

X

X

P. serotina - black cherry

X

P. umbellata - flatwood plum

X

Quercus falcata - southern red oak

X

X

Q. lauri folia - laurel oak

X

X

Q. lyrata - overcup oak

X

Q. nigra - water oak

X

X

X

Q. phellos - willow oak

X

X

X

Q. prinus - chestnut oak

X

X

Q. shumardii - shumard oak

X

Q. similis - bottomland post oak

X

X

Q. Stella ta - post oak

Q. virginiana - live oak

X

X

X

Salix nigra - black willow

X

X

Sapindus saponaria - western soapberry

Taxodium distichum - bald-cypress

Tilia caroliniana - Carolina basswood

X

X

X

Ulmus alata - winged elm

X

X

X

U. americana - American elm

X

U. crassifolia - cedar elm

X

X

U. rubra - slippery elm

Zanthoxylum clava - herculis - toothache tree

X

X

214

THE TEXAS JOURNAL OF SCIENCE

Table 3 (Continued)

Floodplain

Upland

Hardwood

Cypress

Oak-Pine

Streamside

Shrubs

Forest

Swamp

Forest

Woodlands

Amorpha fruticosa — bastard indigo

X

Aralia spinosa - Devil’s walking stick

X

Ascyrum hypericoides - St. Andrew’s cross

X

X

A. stans - Atlantic St. Peter’s wort

X

X

Asimina triloba - pawpaw

X

Baccharis halimifolia - sea myrtle

X

Bumelia lanuginosa - gum bumelia

X

X

Callicarpa americana - American beautyberry

X

Cephalanthus occidentalis - common buttonbush

X

Chionanthus virginica - old-man’s beard

X

Citrus trifoliata - bitter orange

X

X

X

Cornus drummondii - roughleaf dogwood

X

C. florida - flowering dogwood

X

Erythrina herbacea - coral bean

X

X

Forestiera acuminata - swamp privet

X

X

F. ligustrina - privet

X

X

Halesia dip ter a - silverbell

X

Ilex decidua - p 0‘s sum-haw

X

I. vomitoria - yaupon

X

X

X

Ligustrum quihoui - wax-leaf ligustrum

X

Malvaviscus arboreus - turks cap

X

X

Myrica cerifera - wax -myrtle

X

Ptelea trifoliata - woolly hop tree

X

X

Rhamnus caroliniana - Carolina buckthorn

X

X

Rhus copallina - flameleaf sumac

X

Rivina humilis - rougeplant

X

X

Sabal minor - dwarf palmetto

X

X

Sambucus canadensis - common elder-berry

X

Sassafras albidum - sassafras

X

Symphoricarpus orbiculatus - coral-berry

\

X

Symplocos tictoria - horse-sugar

X

Vaccinium arboreum - farkleberry

X

Viburnum dentatum - southern arrow-wood

X

V. nudum - possum-haw

X

V. prunifolium - black -haw

X

V. rufidulum - rusty black -haw

X

Vines

Ampelopsis arborea - peppervine

X

X

X

Aristolochia tomentosa - Wooly Dutchman’s pipe

X

Berchemia scandens - rattanvine

X

X

Bignonia capreolata - crossvine

X

Campsis radicans - trumpet creeper

X

X

Cissus incisa - ivy tree vine

X

Cocculus carolinus - snailseed

X

Gelsemium sempervirens - Carolina jessamine

X

X

VEGETATION TYPES

215

Table 3 (Continued)

Vines (Continued)

Lonicera japonica - Japanese honeysuckle
L. sempervirens - trumpet honeysuckle
Lygodium japonicum - Japanese climbing fern
Matelea gonocarpa - angle-pod milkvine
Melothria pendula - melon cito
Mikania scandens - climbing hempweed
Parthenocissus quinquefolia - Virginia creeper
Passi flora incarnata - may pop
P. lutea - dwarf passion flower
Rhus toxicodendron - poison ivy
Rubus louisianus - blackberry

R. trivialis - southern dewberry
Smilax bona-nox - saw greenbriar

S. glauca - catbriar

S. hispida - bristly greenbriar
S. lauri folia - blaspheme vine
S. pumila - sarsaparilla vine
S. rotundifolia - common greenbriar
Trachelospermum difforme - star jasmine
Vitis aestivalis - summer grape
V. mustangensis - mustang grape
V. rotundifolia - muscadine grape

Floodplain Upland

Hardwood Cypress Oak-Pine Streamside

Forest Swamp Forest Woodlands

x

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

Herbs

Argemone albiflora - white prickly poppy x

Arisaema triphyllum - Jack-in-the-pulpit x

Aster lateriflorus - starved aster x

Athryium filix-femina - Southern ladyfern x

Bidens bipinnata - Spanish needles x

Boehmeria cylindrica - smallspike false nettle x

Cassia occidentals - coffee senna x

Clematis crispa - blue jasmine x

Clitoria mariana - Atlantic pigeonwings x

Coreopsis cardiminae folia - Manzanilla silvestre x

Crepis pulchra - showy hawksbeard x

Cuscuta compacta - dodder x x

Diclipera brachiata - dicliptera x x

Desmodium paniculatum - panicle tick clover x

Elephantopus carolinianus - leafy elephantfoot x

E. nudatus - naked elephantfoot x

E. tomentosus - hairy elephantfoot x

Eupatorium coelestinum - mist flower x x

E. serotinum - late flowering eupatorium x

Euphorbia bicolor - Snow-on-the -prairie

E. cyathophora - wild poinsettia x

Galactia volubilis downy milkpea x

Geum canadense - white avens x

Habenaria clave llata - white fringe orchid x

Hydrolea ovata - hairy hydrolea x

Pan tana horrida - Texas lan tan a x

Lithospermum caroliniense - Puccon x

x

x

x

x

x

216

THE TEXAS JOURNAL OF SCIENCE

Table 3 (Continued)

Floodplain

Upland

Hardwood

Cypress

Oak-Pine

St ream side

Herbs (Continued)

Forest

Swamp

Forest

Woodlands

Lobelia cardinalis - Cardinal flower

L. puberula - downy lobelia

X

X

X

Mitchella repens - Partridgeberry

X

X

Mollugo verticillata — green carpetweed

X

X

Onoclea sensibilis — sensitive fern

Opuntia spp. - prickly pear

X

X

Osmunda regalis - royal fern

Oxalis dillenii - yellow wood sorrel

X

X

X

O. violacea - violet wood sorrel

Persicaria hydro pip eroides - swamp smart weed

X

X

X

P. punctata - water smartweed

Podophyllum peltatum - common mayapple

X

X

Polianthes virginica - false aloe

X

X

Polygala polygama - bitter milkwort

Polypodium polypodioides - Resurrection fern

X

X

Rhexia mariana - Maryland meadowbeauty

Salvia lyrata - lyreleaf sage

X

X

X

Sanicula canadensis - Canada snakeroot

X

Scutellaria ovata - egg skullcap

X

Senecio obovatus - ragwort

X

Spermacoce glabra - smooth buttonweed
Sprianthes gracilis - Texas ladies’ tresses

X

X

X

X

Stachys crenata — shade betony

X

S. drummondii - Drummond betony

Urtica urens - dog nettle

X

X

X

Verbesina virginica - frostweed

X

X

Vernonia texana - Texas ironwood

X

Vicia levenworthii — leavenworth vetch

X

Viola missouriensis - Missouri violet

X

V. primuli folia - violet

X

V. sagittata - arrow-leaved violet

Xanthium strumarium - cocklebur

X

X

X

Grasses and Sedges

Andropogon glomeratus - bushy bluestem

X

X

X

A. virginicus - broom sedge

X

X

Aristida purpurascens - arrowfeather three-awn

X

X

Arundinaria gigantea - giant cane

X

X

Axonopus affinis - carpet grass

X

X

Bromus unioloides - rescue grass

Carex amphibola - amphibious sedge

X

X

X

C. cephalophora - woodbank sedge

X

C. cherokeensis - Cherokee sedge

X

X

C. debilis - spindlefruit sedge

X

C. hyalinolepis - thinscale sedge

X

C. muhlenbergii - Muhlenberg sedge
Chasmanthium latifolium - inland sea oats

X

X

C. laxum - wood grass

X

X

C. sessiliflorum - wood grass

X

X

Digitaria filiformis - slender crabgrass

Eleocharis obtusa - blunt spikesedge

X

X

VEGETATION TYPES

217

Table 3 (Continued)

Floodplain

Upland

Hardwood

Cypress

Oak-Pine

Streamside

Grasses and Sedges (Continued)

Forest

Swamp

Forest

Woodlands

Elymus canadensis - Canada wildrye

X

X

E. virginicus - Virginia wildrye

Erianthus contortus - Bentawn plumegrass
Fimbristylis puberula - Fimbry

X

X

X

X

Leersia virginica - white grass

Melica mutica - two-flower melic

X

X

X

Muhlenbergia schreberi - niblewill muhly

X

X

Oplismenus hirtellus - basketgrass

X

X

0. setarius - basketgrass

Panicum anceps - beaked panicum

X

X

X

P. gymnocarpon - Savannah panicum

P. lanuginosum - wooly panicum

X

X

X

Paspalum bifidum - pitchfork p asp alum

X

P. dilatatum - Dallis grass

X

X

P. langei - rustyseed paspalum

X

P. setaceum - thin paspalum

Poa autumnalis - autumn bluegrass

X

X

Schizachyrium scoparium - little bluestem

X

Sorghastrum elliottii - slender indiangrass

X

X

St ip a avenacea - black seed needlegrass

X

Tridens flavus - purpletop

Tripsacum dactyloides - Eastern gamagrass

X

X

X

TABLE 4

Checklist of Prairie Species in Chambless County

Bluestem

Cordgrass

Grasses and Sedges

Prairie

Prairie

Agrostis hyemalis - spring bentgrass

X

A. scabra - rough bentgrass

X

X

Alopecurus carolinianus — Carolina foxtail

X

Andropogon gerardi - big blue stem

X

A. glomeratus - bushy beard bluestem

X

X

A. temarius - splitbeard bluestem

X

X

A. virginicus - broomsedge bluestem

X

Anthaenantia rufa - purple silky scale

X

Aristida longespica - slim spike three-awn

X

X

A. oligantha - prairie three-awn

X

A. purpurascens — arrowfeather three-awn

X

Axonopus affinis - carpet grass

X

X

Bothriochloa ischaemum - king ranch bluestem

X

B. saccaroides - silver bluestem

X

X

Briza minor - little quaking grass

X

X

Bromus unioloides - rescue grass

X

X

Care: c spp. - sedges

X

Cenchrus echinatus - southern sandbur

X

C. incertus - coast sandbur

X

Chloris petraea - still-leaf chloris

X

X

C. verticillata - tumble windmill grass

X

X

Cynodon dactylon - Bermuda grass

X

X

218

THE TEXAS JOURNAL OF SCIENCE

Table 4 (Continued)

Grasses and Sedges (Continued)

Bluestem Cordgrass

Prairie Prairie

Cyperus articulatus - jointed Hat sedge
C. erythrorhizos - red root flatsedge ,

C. esculentus - chufa
C. globulosus - Baldwin Hatsedge
C. iria - rice field sedge

C. strigosus - false nut grass
Dichanthium spp. — gordo bluestem
Dichromena colorata - white-top star rush
Digitaria adscendens - southern crabgrass

D. sanguinalis - northern crabgrass
Echinochloa crusgalli - barnyard grass
Eleocharis acicularis - needle spikesedge
Eleusine indica - goosegrass
Eragrostis lugens - mourning lovegrass

E. oxylepis - red lovegrass

Erianthus giganteus - sugarcane plumegrass
Eriochloa contracta - prairie supgrass
Fimbristylis spp. - fimbry
Hordeum pusillum - little barley
Juncus effusus - common rush
J. polycephalus - Hat leaf rush
Leptoloma cognatum - fall witchgrass
Muhlenbergia capillaris - gulf muhly
Panicum anceps - beaked panicum
P. capillare - common witchgrass
P. filipes — filey panicium
P. hians - gaping panicium
P. lanuginosum - woolly panicium
P. oligosanthes - heller panicium
Paspalum dilatatum - dallis grass
P. floridanum - Florida paspalum
P. praecox - cypress paspalum
P. notatum - Bahia grass
P. plicatulum - brownseed paspalum
P. setaceum - thin paspalum
Phalaris caroliniana - Carolina canarygrass
Poa annua - annual bluegrass
Rhynchospora fascicularis - stout beakrush
Schedonnardus paniculatus - tumblegrass
Schizachyrium scoparium - little bluestem
S. tenerum - slender bluestem
Scleria spp. - nutrush
Setaria geniculata - knotroot bristlegrass
S. glauca - yellow foxtail
Sorghastrum avenaceum - Indian grass
Sorgum halepense - Johnson grass
Spartina patens - marshhay cordgrass
S. pectinata - prairie cordgrass
S. spartinae - gulf cordgrass
Sphenopholis obtusata - prairie wedgescale
Sporobolus asper - tall dropseed
S. indicus - rattail smutgrass
S. pyramidatus - whorled dropseed
S. virginicus - coastal dropseed

x

x

x

x

x

x

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

VEGETATION TYPES

219

Table 4 (Continued)

Grasses and Sedges (Continued)

Bluestem

Prairie

Cordgrass

Prairie

Stenotaphrum secundatum - St. Augustine grass

X

X

Stipa leucotricha - Texas winter-grass

X

X

Tridens albescens - white tridens

X

T. st rictus - longspike tridens

X

X

Tripsacum dactyloides - eastern gamagrass

X

Trisetum interruptum - prairie trisetum

X

X

Vulpia octo flora - six-weeks fescue

X

X

Willkommia texana - Texas willkommia

X

Forbs

Agalinis maritima - seaside gerardia

X

X

A. purpurea -

X

Allium spp. - onion

X

X

Ambrosia artemisiifolia - eastern ragweed

X

A. psilostachya - western ragweed

X

X

A. trifida - giant ragweed

X

Amsonia illustris - blue star

X

Asclepias linearis - slip milkweed

X

A. viridiflora - green antelope-horn

X

X

A. viridis - antelope-horn

X

Aster ericoides - Heath aster

X

A. subulatus - salt marsh aster

X

X

Baptisia leucantha - plains wild indigo

X

B. sphaerocarpa - green wild indigo

X

X

Cacalia lanceolata - Lance-leaf Indian plantain

X

Callirhoe involucrata - poppy-mallow

X

Capsella bursa-pastoris - shepperd’s purse

X

Cassia fasciculata - partridge pea

X

X

C. obtusifolia - sickle-pod senna

X

Castilleja indivisa - Indian paintbrush

X

Centaurea americana - basket-flower

X

Chenopodium album - Lamb Vquarters

X

X

Cirsium horridulum - yellow thistle

X

C. texanum - southern thistle

X

Clematis drummondii - Texas virgin’s bower

X

Commelina erecta - erect day flower

X

Convolvulus arvensis - field bindweed

X

Conyza canadensis - mare’s tail

X

X

Coreopsis cardaminaefolia - Cardamine coreopsis

X

Croton capitatus - woolly croton

X

X

C glandulosus - northern croton

X

C. monanthogynus - one-seeded croton

X

C. punctatus - gulf croton

X

X

Cuscuta cuspidata - cusp dodder

X

X

Desmanthus acuminatus — sharp-pod bundleflower

X

D. illinoense - Illinois bundleflower

X

X

Dichondra carolinensis - pony-foot

X

Erigeron geiseri - basin fleabane

X

Erodium texanum - Texas filaree

X

X

Erythrina herbacea - coral bean

X

X

Eryngium hookeri - hooker eryngo

X

X

E. yuccifolium - button snakeroot

X

220

THE TEXAS JOURNAL OF SCIENCE

Table 4 (Continued)

Forbs (Continued)

Bluestem Cordgrass

Prairie Prairie

Eupatorium coelestinum - mist flower
E. compositifolium - yankee weed
E. serotinum - late flowerine eupatorium
Euphorbia bicolor - snow-on-the-prairie
E. maculata - spotted euphorbia
E. serpens - mat euphorbia
E. spathulata - warty euphorbia
Eustoma exaltatum - tall prairie gentian
E. grandiflorum - bluebell
Eustylis purpurea - purple pleat leaf
Evolvulus sericeus - silky evolvulus
Gaillardia pulchella - firewheel
Gaura brachycarpa - plain gaura

G. parvi flora - small-flower gaura
Geranium carolinianum - Carolina geranium
Gnaphalium obtusifolium - fragrant cudweed
Hedeoma hispidum rough hedeoma
Helenium amarum - sneezeweed
Helianthus angustifolius - swamp sunflower

H. annuus - common sunflower

H. argophyllus - silverleaf sunflower

H. hirsutus - stiff-haired sunflower
Hydrocotyle verticillata - whorled penny wort
Hy dr ole a ovata - hairy hydro lea
Indigofera miniata - coast indigo

I. suffruticosa - indigo

Iva angustifolia — narrowleaf sumpweed
7. annua — seacoast sumpweed
Krameria lanceolata - trailing rattany
Krigia oppositifolia - weedy dwarf dandelion
Lactuca canadensis — wild lettuce
L. serriola — prickly lettuce
Lantana horrida - Texas lantana
Lepidium virginicum - Virginia pepperweed
Lespedeza capitata - round-head bush clover
L. virginica - slender bush clover
Liatris acidota - sharp gayfeather
L. elegans - pink scale gayfeather

L. pycnostachya - Kansas gayfeather
Linum rigidum - stiffstem flax
Lobelia appendiculata - earflower lobelia
Machaeranthera phyllocephala - camphor daisy
Medicago arabica - spotted bur-clover

M. lupulina - black medick

M. polymorpha - California bur-clover
Melilotus albus - white sweet clover
M. officinalis - yellow sweet clover
Mimosa strigillosa - powderpuff
Mollugo verticillata - green carpetweed
Monarda fistulosa - wild bergamont
Neptunia lutea - low neptunia yellow-puff
TV. pubescens - tropical nuptunia
Nothoscordum bivalve - crow-poison

x

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

VEGETATION TYPES

221

Table 4 (Continued)

Forbs (Continued)

Bluestem Cordgrass

Prairie Prairie

Oenothera laciniata - cut-leaved evening primrose x

O. speciosa - pink evening primrose x

Oxalis dillenii — yellow wood-sorrel x

O. violacea - violet wood-sorrel x

Palafoxia rosea - rose palafoxia x

Parthenium hysterophorus - false ragweed x

Passiflora incarnata - Maypop x

Persicaria hydropiperoides - swamp smartweed x

P. punctatum - dotted smartweed x

Phyla nodiflora - common frog-fruit x

Physalis angulata - ground cherry x

Physostegia intermedia - intermediate lion’s heart x

Phytolacca americana - poke weed x

Plantago elongata - slender plantain x

P. rhodosperma - red-seeded plantain x

Pluchea foetida - stinking fleabane x

P. rosea — marsh fleabane x

Polanisia dodecandra - clammy-weed x

Polygonum aviculare - knotweed x

Portulaca mundula - shaggy portulaca x

Pyrrhopappus multicaulis - many stem false dandelion x

Ranunculus muricatus - rough seed buttercup x

Ratibida peduncularis - Mexican hat x

Rhexia mariana - Maryland meadowbeauty x

Rhynchosia texana - Texas snoutbean x

Rudbeckia grandi flora - large flower coneflower x

R. hirta - black-eyed Susan x

Ruellia humilis - low ruellia x

Rumex crispus - curly dock x

R. pulcher - fiddle dock x

Sabatia campestris - prairie rose-gentian x

Salvia coccinea - tropical sage x

S. lyrata - lyreleaf sage x

Samolus ebracteatus - coast brookweed x

Schrankia uncinata - catclaw sensitive brier x

Scutellaria drummondii - Drummond skullcap x

Senecio imparipinnatus - groundsel x

Sesbania marcrocarpa - bequilla x

S. vesicaria - bag-pod sesbania

Sida lindheimeri — showy sida x

S. rhombifolia - arrowleaf sida x

S. spinosa - prickly mallow x

Silene antirrhina - sleepy catchfly x

Silphium asperrimum - rough stem rosin-weed x

Sisyrinchium angustifolium - blue-eyed grass x

S. exile - blue-eyed grass x

Solanum elaeagnifolium - silver-leaf nightshade x

S. rostratum - buffalo bur x

Sonchus asper - common sow thistle x

S. oleraceus - sow thistle x

Spigelia texana - Texas pinkroot x

Spirant hes cernua - nodding ladies’ tresses x

Stellaria media - chick weed x

x

x

X

X

X

X

222

THE TEXAS JOURNAL OF SCIENCE

Table 4 (Continued)

Forbs (Continued)

Bluestem

Prairie

Cordgrass

Prairie

Stillingia sylvatica - queen’s delight

X

Strophostyles helvola - trailing wildbean

X

Tephrosia lindheimeri - tephrosia

X

T. onobrychoides - multi-bloom tephrosia

X

Thelesperma spp. - greenthread

X

Tidestromia lanuginosa - woolly tidestromia

X

Tillaea aquatica - water pygmy-weed

X

Tradescantia ohioensis - Ohio spiderwort

X

Tragia glanduligera - roseburn

X

Trifolium amphianthum - peanut clover

X

T. repens - white clover

X

Triodanis perfoliata - Venus’ looking-glass

X

Verbena halei - Texas vervain

X

V. hast at a - blue vervain

X

V. xutha - Gulf vervain,

X

Vicia leavenworthii - leavenworth vetch

X

V. ludoviciana - Louisiana vetch

X

Vigna luteola - cowpea

X

X

Xanthocephalum texanum - Texas broomweed

X

X

Woody Plants

Acacia farnesiana - sache

X

X

Amorpha fruticosa - bastard indigo

X

Asimina triloba - pawpaw

X

Ascyrum hypericoides - St. Andrew’s cross

X

Baccharis halimifolia - sea myrtle

X

X

Bumelia lanuginosa - gum bumelia

X

X

Cornus drummondii - roughleaf dogwood

X

Diospyros virginiana - common persimmon

X

Ilex vomitoria - yaupon

X

Iva frutescens - bigleaf sumpweed

X

X

Melia azedarach - Chinaberry tree

X

Myrica cerifera - southern wax-myrtle

X

Opuntia lindheimeri - Texas prickly pear

X

X

0. macro rhiza - plains prickly pear

X

Parkinsonia aculeata - retama

X

Quercus marilandica - blackjack oak

X

Rosa bracteata - MacCartney rose

X

Rubus louisianus - blackberry

X

R. trivialis - Southern dewberry

X

Sapindus saponaria - Western soapberry

Sapium sebiferum - Chinese tallow

X

Sesbania drummondii - rattlebox

X

Tamarix gallica - salt cedar

X

Zanthoxylum clava - herculis - toothache tree

X

Z. fagara - lime prickly ash

X

TABLES

Checklist of Marsh Species in Chambers County

Fresh

Brackish

Grasses - Sedges - Rushes

Marsh

Marsh

Brachiaria platyphylla - broadleaf signalgrass

X

Carex amphibola - Amphibious sedge

X

C. vulpinoidea - fox sedge

X

VEGETATION TYPES

223

Tab le 5 (Continued) _

Fresh

Grasses - Sedges - Rushes (Continued) _ Marsh

Cladium jamai cense - J arnica sawgrass x

Cyperus acuminatus - tapeleaf sedge x

C. articulatus - jointed sedge x

C ochraceus - flatsedge x

C. virens - green flatsedge x

Distichlis spicata - seashore saltgrass

Echinochloa colonum - jungle rice x

E. crusgalli - barnyard grass x

E. cruspavonis - Gulf cockspur x

Eichhomia crassipes - water hyacinth x

Eleocharis acicularis - needle spikesedge x

E. cellulosa - Gulfcoast spikesedge , x

E. montevidensis - sand spikesedge x

E. parvula - dwarf spikesedge

E. quadrangulata - squarestem spikesedge x

Eragrostis reptans - lovegrass x

Fimbristylis castanea - fimbry x

F. vahlii - Vahl fimbry x

Juncus acuminatus - knotleaf rush x

J. bufonius - toad-rush x

J. effusus - common rush x

J. interior - interior rush x

J. roemerianus - needlegrass rush

J. validus - roundhead rush x

Leersia lenticularis - catchfly grass x

L. monandra - bunch cutgrass x

L. virginica - white grass x

Leptochloa filiformis - red sprangletop x

L. nealleyi - Nealley sprangletop , x

L. uninervia - Mexican sprangletop x

Monanthochloe littoralis - shoregrass

Panicum anceps - beaked panicum x

P. geminatum - x

P hemitomon - maidencane x

P. paludivagum - water panicum x

P. repens - torpedo grass x

P. virgatum - switchgrass x

Paspalum acuminatum - brook paspalum x

P. bifidum - pitchfork paspalum x

P. distichum - knotgrass x

P. lividum - longtom x

P. monostachyum - Gulf'd une paspalum
P. vaginatum - seashore paspalum

Phalaris caroliniana - Carolina canarygrass x

Phragmites communis - common reed x

Polypogon monspeliensis - rabbitfoot grass x

Rhynchospora corniculata - horned beakrush , x

Sacciolepis striata - American cupscale x

Scirpus americanus - American bulrush

S. californicus - California bulrush x

S. maritimus - salt-marsh bulrush
S. olneyi - Olney bulrush

Scleria sp. - nut rush x

Brackish

Marsh

x

x

x

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

X

224

THE TEXAS JOURNAL OF SCIENCE

Table 5 (Continued)

Grasses - Sedges - Rushes (Continued)

Fresh

Marsh

Brackish

Marsh

Setaria geniculata - knotroot bristlegrass

X

X

S. Magna - giant bristlegrass

Spartina alterniflora - smooth cordgrass

X

S. cynosuroides - big cordgrass

X

S. patens - marshhay cordgrass

X

X

S. spartinae - Gulf cordgrass

X

X

Sporobolus virginicus - seashore dropseed

X

Zizaniopsis miliacea — giant cutgrass

X

Forbs

Acnida tamariscina - Nuttall’s water-hemp

Alternanthera philoxeroides - Alligator-weed

X

Amaranthus greggii - Gregg amaranthus

X

A. tamariscinus - water hempweed

X

X

Ammannia coccinea - purple ammania

X

Asclepias perennis - shore milkweed

X

Aster spinosus - devilweed aster

X

X

A. subulatus - slim aster

X

X

A. tenuifolius - saline aster

X

Azolla caroliniana - water fern

X

Bacopa monnieri - water-hyssop

X

X

B. rotundifolia - disc water-hyssop

X

Borrichia frutescens - bushy sea oxeye

X

Cakile fusiformis - sea rocket

X

Callitriche heterophylla - large waterwort

X

C. terrestris - annual waterstarwort

X

Calystegia sepium - hedge-bindweed

X

Canna glauca - Indian-shot

X

Cardiospermum halicacabum - common balloon-vine

X

X

Centaurium calycosum - Buckley centaury

X

Centunculus minumus - chaffweed

X

Corchorus hirtus - Orinoco jute

X

' Desmanthus illinoensis - Illinois bundleflower

X

X

Diodia virginiana - rough buttonweed

X

Echinodorus cordifolius - burhead

X

X

E. parvulus - burhead

X

X

E. ro stratus - burhead

X

X

Eclipta alba - Yerba de tajo

X

Glinus radiatus -

X

Heliotropium curassavicum - seaside heliotrope

X

H. procumbens - spike heliotrope

X

Heteranthera limosa - blue mud-plantain

X

H. reniformis - mud-plantain

X

Hydrocotyle bonariensis - large leaf pennywort

X

X

H. umbellata - Umbrella pennywort

X

Hydrolea ovata - hairy hydrolea

X

Hymenocallis liriosme - fragrant white spiderlily

X

Ipomoea sagittata - salt marsh morning glory

X

I. trichocarpa - sharppod morning glory

X

Iris virginica - Southern blueflag

X

Justicia lanceolata - lance-leaved water willow

Kosteletzkya virginica - saltmarsh mallow

X

VEGETATION TYPES

225

Table 5 (Continued)

Forbs (Continued)

Fresh Brackish

Marsh Marsh

Lemna gibba - swollen duckweed x

Limnobium spongia - common frog’s-bit x

Limonium nashii - Carolina sea-lavender

Lindernia anagallidea - false pimpernel x

Lobelia cardinal is - Cardinal flower x

Ludwigia glandulosa - creeping seedbox x

L. octovalvis - water-primrose x

L. palustris - marsh purslane x

L. peploides - verdolaga de agua x

Machaer anther a phyllocephala — camphor daisy x

Malachra capitata - Malva de caballo x

Malvastrum coromandelianum - threelobe falsemallow x

Melochia corchorifolia - redweed x

Modiola caroliniana — Carolina modiola x

Myriophyllum pinnatum - green parrot’ s-feather x

Najas guadalupensis - Southern naiad x

N. mariana - spiny naiad x

Persicaria coccinea - big root smartweed x

P. hydropiperoides - swamp smartweed x

P. punctata — dotted smartweed x

Physostegia angustifolia - lion’s heart x

P. intermedia — intermediate lion’s heart x

Pistia stratiotes - water-lettuce
Pluchea purpurascens - purple pluchea
P. rosea - marsh fleabag
Polygonum aviculare - Prostrate knotweed

Potamogeton diversifolius - waterthread pondweed x

P. nodosus - longleaf pondweed x

Proserpinaca palustris - mermaid-weed x

Rivina humilis - rougeplant x

Rorippa islandica - bog marshcress x

Rumex chrysocarpus - dock x

R. pulcher - fiddle dock x

Ruppia maritama - widgeon-grass x

Sabatia campestris - prairie rose-gentian x

Sagittaria graminea - grassy arrowhead x

S. latifolia - common arrowhead x

S. longiloba - longlobe arrowhead x

S. papillosa - nipplebract arrowhead x

Samolus ebracteatus - coast brookweed x

S. parviflorus - brookweed x

Sesuvium maritimum - sea purslane

S. portulacastrum - sea purslane x

Solanum nodiflorum - nightshade x

Solidago sempervirens var. Mexicana - seaside goldenrod x

Spergularia echinosperma — bristleseed sand-spurry
S. marina - marsh sand-spurry

Sphenoclea zeylanica - pie fruit x

Suaeda linearis - annual seepweed

Teucrium canadense - wood sage x

Thelesperma filifolium - green-thread x

Thurovia triflora - three flowered thurovia x

x

x

x

x

x

x

x

X

X

X

X

X

X

X

226

THE TEXAS JOURNAL OF SCIENCE

Table 5 (Continued)

Forbs (Continued)

Fresh

Marsh

Brackish

Marsh

Typha angustifolia - narrow-leaved cattail

X

X

T. latifolia - common cattail

X

Utricularia gibba - cone-spur bladderwort

X

U. purpurea - purple bladderwort

X

Wolffia columbiana - Columbia water-meal

X

Wolffiella gladiata - mud-midget

X

Xanthium strumarium - cocklebur

X

Woody Plants

A triplex spp. - saltbush

X

Baccharis halimifolia - sea myrtle

X

X

Bat is maritima - saltwort

X

Bumelia lanuginosa - gum bumelia

X

Cephalanthus occidentalis - common buttonbush

X

Diospyros texana - Mexican persimmon

X

Hibiscus lasiocarpos - woolly rose-mallow

X

X

H. militaris - scarlet rose-mallow

X

Iva frutescens - bigleaf sumpweed

X

Lycium carolinianum - Carolina wolfberry

X

Salicomia bigelovii - biglow glasswort

X

S. virginica - Virginia glasswort

X

Sapium sebiferum — Chinese tallow

X

Sesbania drummondii - rattlebox

X

X

S. macrocarpa - coffeebean

X

X

Tamarix gallica - salt cedar

X

X

Zanthoxylum clava-herculis - toothache tree

X

X

PLANT ASSEMBLAGES OF CHAMBERS COUNTY
Oak-Pine Forest

Oak-pine forests are identified by presence of loblolly pine in relative abun¬
dances of 25-100%. Associated species usually include sweetgum, (see Table 2
for scientific names), willow oak, and water oak. Less frequently, laurel oak,
basket oak, southern red oak, or hickories may be encountered. Forest height
ranges from 20 to 30 m; tree density is in the vicinity of 500 trees per ha (trees
> 10 cm diameter at breast ht.); basal area is 20 to 30 m2/ha. The understory
shrubs may be dense or sparse, depending on height and density of the tree
canopy. Yaupon and American beautyberry are predominant shrubs. Greenbriar,
trumpet creeper and Virginia creeper are the common vines.

In Chambers County, oak-pine forests occur on upper terraces of the Trinity
River floodplain, along the major streams emptying into upper Trinity Bay
(Double Bayou, Turtle Bayou), and along Cedar Bayou down to Baytown. This
assemblage is replaced by floodplain hardwood forest or cypress swamp on the
lower terraces of the Trinity River. Under natural conditions, the boundary be¬
tween oak -pine forest and grassland may have been determined by soil differences,

VEGETATION TYPES

227

since forests are often limited to alluvial soils with better drainage and a more
constant moisture supply (Kilburn, 1959; Severson and Arneman,T973). Periodic
drought and subsequent fires may also have played a role in determining this
vegetation boundary, but our observation of successional patterns in the prairie
and our unpublished data from Harris County suggest that the soil difference
is of primary importance in Chambers County.

Edmiston (1963), Monk (1965), and others indicate that forests such as these
are often successional — the predominance of loblolly pine and sweetgum is indi¬
cative of fire or cutting — and that 75-150 yrs after disturbance they will revert

228

THE TEXAS JOURNAL OF SCIENCE

to mixed forests containing much less pine. On relatively dry sites in Chambers
County, southern red oak, post oak and hickories will probably become more
prominent; on moist sites magnolia and basket oak probably will predominate.
This assemblage probably represents the southernmost extension of the Southern
Mixed Hardwood Forest (Monk, 1965). McLeod (1971) calls it the “Lower Big
Thicket Forest” and Kuchler (1964) maps it as Oak-Hickory-Pine Forest.

The oak-pine forest along both forks of Double Bayou was erroneously mapped
as oak motte by Fisher, et al, (1972). Fisher, et al, (1972) also mistakenly in¬
cluded shortleaf and longleaf pine as components of the mixed Pine-Hardwood
Forest, which roughly corresponds ta our oak-pine forest. Neither species of pine
is present in Chambers County.

Cypress Swamp

Cypress swamps are relatively open stands of bald-cypress about 30 m high,
with high tree density (about 500 per ha) and high basal area (50 m2 /ha). Other
important woody species include willow, water elm, and buttonbush. Along the
edges of the cypress swamps, water hickory is common.

Cypress swamps are common in the Trinity River floodplain where standing
fresh water exists for most of the year. Cypress trees appear to be very sensitive
to saline encroachment (O’Neil, 1949), so they do not extend onto the present
Trinity delta. Since regeneration appears to occur only when the swamp is dry
(Hall, et al, 1946) permanent flooding could have a substantial impact on these
forests.

Cypress swamps are well-known in the southeastern United States (Penfound,
1952), and those in Chambers County appear to be typical.

Floodplain Hardwood Forest

Water oak, cedar elm, sugarberry, green ash, and hawthorn characterize the
forests of the lower Trinity River floodplain and the mouth of Cedar Bayou.
Willow and sycamore are common on recent sand bars. Cypress and tupelo occur
in the sloughs. The forests are relatively open, with widely spaced overstory oaks
above a subcanopy of cedar elm, sugarberry, and hawthorn (15-20 m tall). Tree
density is about 700 stems per ha; basal area is about 15 m2 per ha. In northern
Chambers County, the understory varies from dense stands of palmetto to con¬
tinuous grass and herb cover. Savannah panicum, smartweed and water willow
are dominant in the wet grassy areas; woodgrass and sedges dominate in drier
grassy areas. Along the Trinity Bay margin and at the mouth of Cedar Bayou,
palmetto is replaced by dense stands of yaupon, and the herb layer contains
more upland grasses and herbs. Vines are a conspicuous component everywhere.
Rattan, poison ivy and trumpet creeper are common in the Trinity floodplain;
greenbriar and blackberry are important along the Trinity Bay margin and at the
mouth of Cedar Bayou.

VEGETATION TYPES

229

Increased seasonal flooding in floodplain forests usually enhances tree growth,
but permanent flooding is detrimental to many floodplain hardwood species
(Broadfoot and Williston, 1973). Saline encroachment may also influence these
forests: we observed tree death at the mouth of Cedar Bayou, apparently due to
subsidence-related salt encroachment.

Kiichler (1964) maps the Trinity River floodplain as Southern Floodplain
Forest. However, the species composition is quite different and does not fit any
of Kiichler’s mapping units. The Trinity River floodplain forest bears a stronger
resemblance to the vegetation along rivers further south along the central Texas
coast described by Tharp (1926).

Streamside Woodlands

Sugarberry, hawthorn, and low-growing water oak are the characteristic spe¬
cies in the streamside woodlands. The trees are usually small (less than 15 m tall),
and are frequently tied together in a dense tangle by greenbriar and blackberry.

Streamside woodlands occur along Oyster Bayou, Elm Bayou and Spindletop
Bayou. In contrast, the larger bayous emptying into Trinity Bay support oak-pine
forest along the streambanks. Streamside woodlands are a conspicuous feature of
prairie regions (Weaver, 1960).

Bluest em Prairie

Characteristic species of this assemblage are big bluestem, Indiangrass, switch-
grass, little bluestem, and eastern gamagrass. Under natural conditions, these
species form a uniform stand of grasses 1-2 m tall, interrupted by individuals and
clumps of legumes and other forbs. In Chambers County, species composition
appears to vary with soil texture. Little bluestem predominates on sandy soils
that are drier; the others are more common on moist sites (clay and loam soil).
Brownseed paspalum is also common on moist sites. The Soil Conservation Service
(1976) divides this assemblage into 4 types on the basis of soil texture (sandy
prairie, loamy prairie, blackland prairie, lowland prairie). However, species com¬
position differences are not great, so we prefer to lump them into a single assemblage.

Today, this assemblage has been almost completely replaced in eastern Cham¬
bers County by rice fields and pastures. The fields are normally planted 2 years
in rice and then the fallow for 2-3 years. The first year following cultivation,
sedges, slim aster, and barnyard grass are dominant. Subsequently, bushybeard,
annual sumpweed, rattlebush, and sea myrtle invade. On heavily grazed sites,
carpetgrass and smutgrass predominate. Vasey grass becomes dominant on prop¬
erly grazed sites and may persist for many years. In western Chambers County
most of the prairie is no longer in agricultural use, but due to protection from
fire, extensive brushland has emerged. Sea myrtle is the dominant brush species.
Chinese tallow and huisache are common in some areas.

This assemblage is the same as the Eastern Tallgrass Prairie Formation of Weaver
(1954) or the Bluestem Prairie of Kiichler (1964). Kiichler (1964) mapped it

230

THE TEXAS JOURNAL OF SCIENCE

as Bluestem-Sacahuiste Prairie, but since sacahuiste (Gulf cordgrass) is restricted
to the coastal rim in Chambers County, we prefer to designate this assemblage as
Bluestem Prairie.

Cordgrass Prairie

Gulf Cordgrass (sacahuiste) is the characteristic species of the cordgrass prairie,
though little bluestem may also occur. This assemblage was probably once more
extensive as an intermediate type between upland prairie and brackish marsh,
but it now exists primarily on th;e natural levee along the north edge of East Bay
as a nearly pure stand of Gulf cordgrass. It is equivalent to the Salty Prairie Range
Site of the Soil Conservation Service.

Cordgrass prairie is more extensive further south along the Texas Coast; other
cordgrass species are common on saline soils of northern prairies (Weaver, 1954).

Fresh Marsh

Marshland can be divided into shallow and deep phases of fresh, brackish, and
salt assemblages (Shaw and Fradine, 1956; Shiftlet, 1963). However, here the
only divisions we recognize are fresh and brackish marsh, primarily because the
others are either absent or are of very limited extent in Chambers County. The
principal distinguishing feature is presence (in brackish marsh) or absence (in
fresh marsh) of marsh-hay cordgrass ( Spartina patens).

The most common species of fresh marshes are common reed, sawgrass, cut-
grass, torpedograss, longtom, cattail, and big cordgrass, in various combinations.
Alligator weed is a common successional species. It occurs on the active areas of
the Trinity River delta between Interstate Highway 10 and the northern boundary
of the county.

As mentioned earlier, landward sides of the coastal marshes, especially near
the mouth of Oyster Bayou, once supported extensive fresh marsh assemblages
(USDI, 1952). Large stands of common reed may, in fact, have persisted until
the mid 1950’s (F. M. Fisher, personal communication). However, at present
there is little evidence of fresh marsh vegetation. At least 3, and possibly 4, factors
are involved. First, freshwater marsh on Beaumont Clay soils has been claimed
for rice production. Second, impoundment of water for rice farming probably
has reduced freshwater input into the marshes. Third, land subsidence probably
has resulted in increased salt water intrusion. Fourth, disturbances (e.g., burning
to improve muskrat habitat) seems to favor fresh marsh species (O’Neil, 1949),
so changed management objectives may have resulted in succession from fresh-
marsh species to brackish-marsh species without any actual change in soil or
water salinity. Marsh drainage and conversion to rice fields are undoubtedly the
most significant factors.

Brackish Marsh

This assemblage is dominated by marsh-hay cordgrass. and/or seashore saltgrass.
Usually one or more rush or sedge species occur in. isolated clumps (especially

VEGETATION TYPES

231

salt-marsh bulrush and olney bulrush). Near tidal drains needlegrass rush may
occur. On levees, fresh-marsh species, especially common reed and big cordgrass,
are common. Seashore paspalum and longtom are also common in the fresher
areas.

Smooth cordgrass is common in Chambers County, but it occupies such a
narrow fringe along East Galveston Bay that it is not mappable as a distinct
assemblage.

Brackish marsh is the predominant marsh assemblage in Chambers County. It
occurs in the mouth of Cedar Bayou, in the Trinity River Bottom south of Inter¬
state Highway 10, along Smith Point, and all along the southeastern part of the
county.

Any alteration of overland water flow will undoubtedly affect species compo¬
sition and productivity in the coastal marshes. The area of the marshes has prob¬
ably now been reduced by drainage and reclamation as much as is possible, mostly
at the expense of the fresh-marsh types, so further reduction of fresh water input
will result in slightly increased salinity, and possibly increased frequency of dry
conditions in the marsh. This probably will reduce productivity and may result
in a shift in species composition toward gulf cordgrass.

Brackish marsh appears equivalent to Kiichler’s (1964) Northern Cordgrass
Prairie (marsh-hay cordgrass and seashore saltgrass dominant). It bears little re¬
lationship to his Southern Cordgrass Prairie (smooth cordgrass dominant). Hence
we believe the term brackish marsh is a more appropriate name until additional
data are published on the relationship of Kiichler’s Northern Cordgrass Prairie
type to the southern marshes. This marsh type is probably predominant from
the Trinity River eastward at least to the Mississippi (Chabreck, 1972).

The extreme southeastern tip of the county fronts on Galveston Bay, and
supports dune vegetation. Because of the small area involved, and the presence
of brackish marsh species in the assemblage, the area is included as brackish
marsh. It is actually a distinct assemblage, dominated by bitter panicum and
marsh-hay cordgrass. It has much in common with the dune vegetation of Mustang
Island, Texas (Gillespie, 1976).

VEGETATION - ENVIRONMENT RELATIONSHIPS

Water is the factor with most obvious influence on the vegetation of Chambers
County. The boundaries between marsh and prairie are set by limits of periodic
flooding, and the boundary between prairie and forest is indirectly related to
water through the effect of soil moisture in preventing upland prairie fires from
burning into lowland forest. Water is important in determining variation within
the physiognomic types, also, though other factors may also be involved. In
marshes, variation in water level and salinity can account for most of the varia¬
tion in species composition (Penfound and Hathaway, 1938; Shiflet, 1963).

In forests and prairies, water availability depends on topographic position and
on soil texture, so these landscape factors are the primary determinants of species

232

THE TEXAS JOURNAL OF SCIENCE

composition (Caplenor, 1968; Rice, 1965; Weaver, 1954; Wells, 1942). The mois¬
ture dependence is readily apparent in the forests of Chambers County, where 4
of the 5 forest assemblages (cypress swamp, floodplain hardwood, flatwoods,
and streamside woodlands) can be identified from topographic information alone.

In the prairies, variation in species composition is less apparent— certainly
vegetation physiognomy changes much less— with changes in moisture availability.

Species composition may also be influenced by catastrophic events like fires
or floods. For example, long-term fluctuations in climate may cause marked
changes in prairie species composition (Albertson and Tomanek, 1965; Lynch,
1971), or storm tide flooding which brings in salt water can affect species com¬
position of marshes (Shiflet, 1963). Thus, species composition of plant assem¬
blages is probably constantly changing in response to long- and short-term climatic
fluctuations.

In addition to responding to environmental factors, vegetation also responds
to human disturbance. All stages of prairie, forest, and marsh succession can be
fjound in Chambers County, and they are mentioned briefly in the climax assem¬
blage descriptions above.

None of the variation in physiognomy or species composition of Chambers
County vegetation is obviously tied to soil chemical differences (apart from salin¬
ity), though variations in soil nutrients (i.e., variations in fertility) undoubtedly
exist and influence growth rates and species composition within assemblages.

ACKNOWLEDGEMENTS

The work reported here was completed in consultation with the Rice Center for
Community Design and Research and was financed in part through a grant to the
Southwest Center for Urban Research from the National Science Foundation,
Research Applied to National Need. The work was completed while Mr. Neaville
was employed as Range Conservationist, Soil Conservation Service.

We wish to thank P. L. Marks for critically reading the manuscript, and pro¬
viding his characteristic insightful, penetrating comments.

LITERATURE CITED

Adams, D. A., 1963-Factors influencing vascular plant zonation in North Carolina salt
marshes. Ecol. , 44:445.

Albertson, F. W., and G. W. Tomanek, 1965-Vegetation changes during a 30-year period in
grassland communities near Hays, Kansas. Ecol., 46:714.

Broadfoot,W. M., andH. L. Williston, 1973-Flooding effects in southern forests. J. Forestry,
71:584.

Caplenor, D., 1968-Forest composition on loessal and non-loessal soils in west-central
Mississippi. Ecol., 49:322.

Charbreck, R. H., 1972-Vegetation, water, and soil characteristics of the Louisiana coastal
region. Bull. No. 664, L.S.U. Agr. Expt. Sta., Baton Rouge.

VEGETATION TYPES

233

Correll, D. S., and M. C. Johnston, 1910-Manual of Vascular Plants of Texas. Texas Research
Foundation, Renner, Texas.

Edmiston, J. A., 1963-The ecology of the Florida pine flatwoods. Ph.D. thesis. Univ. Fla.,
Gainesville.

Fisher, W. L., J. H. McGowen, L. F. Brown, Jr., and C. G. Groat, 197 2-Environment al Geo¬
logic Atlas of the Texas Coastal Zone-Galveston-Houston Area. Bureau Econ. Geol.,
Univ. Texas, Austin.

- , 1 973 -Environmental Geologic Atlas of the Texas Coastal Zone- Beaumont -Port

Arthur Area. Bureau Econ. Geol., Univ. Texas, Austin.

Gillespie, T. S., 1976-The flowering plants of Mustang Island, Texas-An annotated check¬
list. Tex.J. Sci., 27(1): 131.

Gould, F. W., 1969-Texas plants: A checklist and ecological summary. Texas Agr. Expt.
Sta. Bull. MP- 58 5 1 Rev.

Hall, T. F., W. T. Penfound, and A. D. Hess, 1946-Water-level relationships of plants in the
Tennessee Valley. Tenn. Acad. Sci., 10:18.

Jefferies, R. L., 1972-Aspects of salt-marsh ecology with particular reference to inorganic
plant nutrieiton. In Barnes, R.S.K., and J. Green (Eds.), The Estuarine Environment.
Applied Publishers, London, pp. 61-85.

Kilburn, P. D., 1959-The forest -prairie ecotone in Northeastern Illinois. Amer. Midi. Nat.,
62:206.

Kiichler, A. W., 1964 -Potential Natural Vegetation of the Conterminous United States
(Manual to accompany the map). Amer. Geogr. Soc. Spec. Publ. 36.

Lohse, A., Joe Tyson, and Consultants, 191 3 -Environmental Resources Inventory and
Evaluation Clear Creek, Tx. U.S. Corps of Engin., Galveston.

Lynch, Br. Daniel, 1971-Phenology, community composition, and soil moisture in a prairie
at Austin, Texas. Ecol., 52:890.

McHarg, I. L., 1969 -Design with Nature. Natural History Press, Garden City, N.Y.

McLeod, C. A., 1971-The Big Thicket Forest of East Texas. Tex. J. Sci., 23(2) : 221 .

Monk,C. D., 1965 -Southern mixed hardwood forest of north central Florida. Ecol. Monogr.,
32:335.

O’Neil, Ted, 1949-The muskrat in the Louisiana coastal marshes, a study of the ecological,
biological, tidal, and climatic factors governing the production and management of the
muskrat industry in Louisiana. New Orleans, La., Dept, of Wildlife and Fisheries.

Penfound, W. T., 1952-Southern swamps and marshes. Bot. Rev., 18(6):413.

- , and E. S. Hathaway, 1938-Plant communities in the marshlands of southeastern

Louisiana. Ecol. Monogr., 8:1.

Phleger, C. F., 1971 -Effect of salinity on a salt marsh grass. Ecol., 52:908.

Rice, E. L., 1965 -Bottomland forests of north-central Oklahoma. Ecol., 46:708.

Rowe, P. G. (Ed.), 1974- An approach to natural environmental analysis for development
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Severson, R. C., and H. F. Arneman, 1973-Soil characteristics and the forest-prairie ecotone
in northwest Minnesota. Soil Sci. Soc. Amer. Proc., 37:593.

Shaw, S. P., and C. G. Fredine, 1956-Wetlands of the United States. U.S. Dept. Interior
Fish and Wildlife Service, Circular 39, pp. 67.

Shiflet, T. N., 1963-Major ecological factors controlling plant communities in Louisiana
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Solomon, D. E., and G. D. Smith, 1973-Seasonal assessment of the relationship between
the discharge of the Trinity River and the Trinity Bay Ecosystem. Coastal Ecosyst. Mgt.,
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Tharp, B. C., 1926 -Structure of Texas vegetation east of the 98th meridian. Univ. Texas
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Ungar, I. A., 1966-Salt tolerance of plantsgrowing in the saline areas of Kansas and Oklahoma.
Ecol., 47:154.

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Bot. Rev., 8:533.

GEOMYID INTERACTION IN BURROW SYSTEMS

by GRAHAM C. HICKMAN

Department of Zoology, University of Natal,
Pietermaritzburg, South Africa

ABSTRACT

A 5.5 m long and 7.5 mm wide glass-walled observation chamber restricted construction
of burrow systems to 2 dimensions. Introductions of representatives of 3 genera of North
American geomyids were made in various combinations: size was an important factor in
combat. Geomyids were dominant over other vertebrates and invertebrates tested. Ecological
factors are probably more important than aggressiveness in determining range boundaries
for pocket gophers.

INTRODUCTION

A variety of parasitic and nonparasitic invertebrates, herptiles, and small
mammals invade burrow systems. The responses between pocket gophers and
this diverse fauna is speculative. Documentation of several adult pocket gophers
in a single burrow system is sparse. Nonetheless, burrow systems do intersect
and cases of plural occupancy reported (Hansen and Miller, 1959). Encounters
within burrow systems probably occur regularly in areas of high population
density. The degree and nature of intra- and interspecific interaction is little
known, even though the aggressive nature of geomyids is demonstrated in that
pocket gophers are solitary. Interspecific aggressiveness between pocket gopher
species may be important in mutual competitive exclusion and parapatry char¬
acteristic of geomyids.

Lack of knowledge in these areas revolves around the difficulty of visually
monitoring the subterranean activities of this most fossorial of rodent families
in North America. This study attempted to solve this problem by construction
of a large dirt-filled, glass-walled observation chamber which permitted con¬
struction of a burrow system in only 2 dimensions.

MATERIALS AND METHODS

The burrow simulator measured 5.5 m long, 1.2 m high, and 7.6 cms wide
(Fig 1). Construction of a similar observation chamber has been described by

Accepted for publication: January 17, 1977.

The Texas Journal of Science, Vol. XXIX, Nos. 3-4, December, 1977.

236

THE TEXAS JOURNAL OF SCIENCE

Figure 1. Position of components used in the construction of the burrow simulator.

B = bolt, BB = back brace, F = frame, FB = front brace, P = plywood, PG = plate
glass, TB = triangular brace, W = wire. Soil within the chamber was never
changed, remaining “fresh” after a year and a half of use.

Hendricksen and McGaugh (1975) after a design by Hickman (1974). Pocket
gopher movements and interactions were observed in the chamber with 2 in-
flourescent and 2 incandescent infra-red lamps.

Pocket gophers were live-trapped (Baker and Williams, 1972): Thomomys
bottae from localities near Sacramento, New Mexico; Geomys bursarius from
Kermit, and Slaton, Texas; and Pappogeomys castanops from Lubbock, Texas.
Animals were fed crimped oats and laboratory rabbit chow, occasionally lettuce,
and individually caged in covered 20 gal garbage cans filled with moist sandy
soil.

A minimum of 3 observations of individuals of each species weight class
was made for inter- and intraspecific interaction of pocket gophers, and for
each geomyid interaction with various non-geomyid vertebrates and inverte¬
brates. Pocket gophers were permitted 3 to 5 days in the burrow simulator
before being considered resident.

RESULTS

Pocket gophers yielded no sign of ticks, fleas, warbles, or any evidence of in¬
fection. Mites were found on pocket gophers, but the animals did not appear
irritated by their presence.

Various beetles, size 20 to 40 mm, were ignored in the burrow system. Camel
crickets (Ceuthophilus) hung from ceilings and walls, avoiding contact with
pocket gophers. Earthworms were occasionally severed during excavation, but
the wriggling remains of earthworm were treated with indifference. Ants (abund¬
ant in the surrounding trapping areas) were never found in burrow systems.

Pocket gophers showed little reaction to various amphibians. Likewise, Am-
bystoma tigrinum did not react to being walked on by pocket gophers while
passing in the narrow tunnelway. In one instance, Pappogeomys castanops arrived

GEOMYID INTERACTION

237

at the end of a tunnel, apparently to begin excavating, only to find a tiger sala¬
mander at rest. The pocket gopher picked the salamander up by the mid-dorsal
skin with the incisors, turned, and deposited the animate obstruction 0.6 m
away from the plug. The pocket gopher then returned to the plug to begin ex¬
cavating. The salamander did not appear agitated or physically harmed. In an¬
other incident, a Geomys bursarius pushed a tiger salamander into a plug 5 con¬
secutive times before the caudate was able to squeeze to one side allowing the
persistent pocket gopher to work without further interference. Large Bufo
cognatus were pushed aside or packed into plugs. A jumping toad would cause
a pocket gopher to retreat backwards for a short distance before it returned
to investigate further. Scaphiopus couchi, a smaller toad, were passively toler¬
ated.

Lizards moved faster than amphibians in the burrow. An occasional pause by
pocket gophers was the only reaction to the passing of the fast moving lizard
(Cnemidophorus sexlineatus) in the tunnelway. Pocket gophers isolated the
slower hog-nose snake, Heterodon nasicus , behind a dirt plug. After several
min to several hrs, pocket gophers unplugged the restricted area for possible
reclamation of the barricaded portion of the system. The same response was
elicited by introducing a bull snake, Pituophis melanoleucus, into burrow sys¬
tems. Dirt was excavated from the floor, walls, and ceiling of the burrow to
block the advancing bull snake. Strikes by the snake appeared to bounce off the
wall of incisors and foreclaws of the pocket gopher, and wall of dirt pushed at
the snake.

Perognathus flavus, the silky pocket mouse, and Onychomys leucogaster, the
grasshopper mouse, avoided pocket gophers with apparent ease. Nonetheless,
both species never refused an opportunity to leave the burrow system. Larger
rodents such as Dipodomys spectablis, the banner- tailed kangaroo rat, entered
burrows of Pappogeomys castanops and Geomys bursarius , but not Thomomys
bottae, because of the smaller tunnel diameter. Larger and slower mammals
were herded to the surface or into a blind tunnel where retreat was impossible.
Intruders were then sealed off with dirt. Kangaroo rats either dug a tunnel to
the surface, or through the plug. However, one Dipodomys spectablis and a
13-lined ground squirrel died as a result of being buried alive. The 1 3— lined
ground squirrels retreated as the pocket gopher “splashed” dirt in their direction.

Pocket gophers investigated any disturbance. If the pocket gopher acted
dominant, the intruder was chased and any openings to the burrow plugged.
In extreme cases, submissive pocket gophers would live on the surface. If equal,
the resident or intruding pocket gopher quickly engaged in excavating dirt as
a buffer between the pocket gopher and the disturbance. It was not unusual for
2 pocket gophers to add dirt to an intervening plug. Pocket gophers of large
differential size could not interacf within burrows because burrow walls re¬
stricted the movement of the large pocket gopher.

238

THE TEXAS JOURNAL OF SCIENCE

Interspecific interaction between adult pocket gophers favored the large
pocket gopher (Table 1). Smaller pocket gophers intruding into larger burrows
were invariably intimidated by the larger pocket gophers. Territorial behavior
varied with the location of the encounter in the burrow system; for example,
submissive pocket gophers became aggressive when trapped in a blind tunnel.
Small residents were usually more submissive to larger pocket gophers in areas
where the burrow diameter was dilated. Burrow diameters larger than normal
may be caused by cave-ins, or by occupation of an old system. Thomomys
bottae , being smaller than Pappogeomys castanops and Geomys bursarius , was
at size disadvantage. In one instance, a 353 g Pappogeomys castanops trapped
a small Thomomys bottae in a blind runway and killed the smaller pocket
gopher by biting through the skull. Even a young Pappogeomys castanops was
dominant over a mature Thomomys bottae because of size differences. Most
encounters between pocket gophers were not mortal battles, usually ending
instead with one pocket gopher retreating to the surface, or walling itself off in
a small portion of the system.

DISCUSSION

Pocket gophers may be afflicted by ectoparasites. English (1932) reported
mites and lice, 'while Wilks (1963) did not find any external parasites on G bur¬
sarius (= breviceps ) from Texas. No fleas or ticks were found on Geomys, Pap¬
pogeomys, of Thomomys.

Many non-parasitic forms of invertebrates have been first described from
pocket gopher systems. Pseudoscorpions, dipterans, beetles, Collembolla, cave
crickets, scorpions, spiders, centipedes, thysanurans, and moths (Ingles, 1952;
Hermann, 1950; Hubbell and Goff, 1940) offer no serious obstruction to the
activities of a pocket gopher (except as possible reservoirs for disease), or com¬
pete indirectly for habitat or food. The only inconvenience for a pocket gopher
may be a bite from a large arthropod, whereby the pocket gopher may recipro¬
cate by offering the temporary inconvenience of burial. Contact with ants are
apparently avoided by rerouting of the tunnel system.

Vertebrates taken from geomyid burrow systems are surprisingly diverse.
A frequently mentioned amphibian is the tiger salamander, Ambystoma tigrinum
(Seton, 1929; Howard and Childs, 1959; Vaughan, 1961). Other amphibians
found in the burrow systems of Geomys bursarius in Texas include Bufo speci-
osus , B. valliceps, Scaphiopus couchi , and S. holbrooki (Wilks, 1963). Amphibia
pose no threat to the Geomyidae and vice-versa. Perhaps the poisonous epi¬
dermal glands of toads inhibits geomyids from biting or otherwise injuring
amphibian intruders. A problem could arise for a pocket gopher if large num¬
bers of anurans and caudates congested the tunnelways following a rain, but this
has never been reported.

TABLE 1

Interaction of Pocket Gophers Released into the Observation Chamber (Intruders) with Pocket Gophers which Occupied an Established
Burrow System (Residents). Three Weight Classes of Pocket Gopher were Tested 3 Times Each for Each of the 3 Species. An X Indicates
no Interaction. In the Confrontations, the Resident was either Dominant (D), Equal (E), or Submissive (S) to the Intruder

GEOMYID INTERACTION

239

240

THE TEXAS JOURNAL OF SCIENCE

Reptiles have been found in burrow systems. Lizards are likely only occasional
visitors and are a little more than a nuisance when present. A whiptail lizard,
Cnemidophorus tigris , has been found in a live- trap set for Thomomys bottae
(Howard and Childs, 1959). The hognose snake, Heterodon platyrhinus , which
has been taken from a burrow system of Geomys bursarius in Texas (Wilks, 1963),
poses no threat to the adult pocket gopher, since Heterodon subsist exclusively
on other reptiles and amphibians. Nonetheless, the pocket gopher is inconven¬
ienced at least by a temporary loss of a portion of the tunnel system. The ability
of the bull or gopher snake ( Pituophis melanoleucus) to dig through the dirt¬
pushing defenses of a pocket gopher is one reason for this snake being a major
predator of geomyids (Howard and Childs, 1959; Morse, 1927). The glossy snake
( Arizona elegans), listed by Stebbins (1954) as a predator on pocket gophers,
was found by Wilks (1963) in the burrow system of Geomys bursarius. Howard
and Childs (1959) observed that Thomomys bottae did not retreat from a rattle¬
snake, but instead pushed dirt towards it. Snakes found in this study were re¬
stricted in striking ability because of limitations to coiling imposed by a narrow
tunnelway.

An efficient mammalian predator entering geomyid burrow systems is the
long- tailed weasel, Mustela frenata (Scheffer, 1945; Moore, 1945 ; Vaughan, 1961;
Cummings, 1946). Most of the other carnivores such as bobcats, striped skunks,
foxes, coyotes, badgers, and housecats known to prey on pocket gophers are too
large to enter a geomyid burrow system. However, Scheffer (1910) recorded a
spotted skunk trapped from a pocket gopher system.

Insectivores may infrequently intrude on geomyids. Scheffer (1945) found
the mole, Scalopus townsendii , and the pocket gopher, Thomomys talpoides,
caught in a runway constructed by the mole. Shrews may utilize the burrows
of both moles and pocket gophers (Scheffer, 1954).

Rodents are frequently found in the burrows of pocket gophers. Howard
and Childs (1959) live-trapped the deer mouse ( Peromyscus maniculatus),
pocket mouse ( Peromyscus inornatus ), and the kangaroo rat ( Dipodomys
heermanni ) from the burrows of Thomomys bottae. Wilks (1963) thought that
Peromyscus hispidus in burrows of Geomys bursarius in Texas robbed the food
caches of the pocket gopher. Perognathus and the grasshopper mouse ( Ony -
chomys leucogaster) were similarly evasive in the present study, but left the
burrow system given the opportunity. While Vaughan (1961) occasionally found
ground squirrels forcing pocket gophers from pocket gopher burrow systems,
Scheffer (1945) notes Spermophilus townsendii expelled from the burrow of
Thomomys talpoides. Spermophilus tridecemlineatus might evict a smaller
pocket gopher if the diameter of the burrow was dilated, but not under usual
circumstances. The fact that geomyids lack the speed to apprehend smaller
antagonists does not appear to be a serious handicap, since the ultimate goal
of solitude is attained.

GEOMYID INTERACTION

241

By plugging all openings to the surface, the pocket gopher not only excludes
predators, but inhibits a host of interlopers seeking the safety and compatible
environment of a burrow system (Vaughan, 1961). The small eyes and ears,
short hair, large incisors which are permanently exposed, and large foreclaws
all contribute to making the pocket gopher the dominant form of the subter¬
ranean environment in North America.Yet, the herbivorous dietary regimen of
the Geomyidae precludes anything but a commensal or parasitic relationship
with various arthropods and other moderate-sized fauna which benefit from the
diversification of a subterranean habitat (Howard and Childs, 1959; Ken¬
nedy, 1964).

Questions as to which of the 3 North American genera of geomyids is most
aggressive has been touched upon by Miller (1964) and Best (1973). Pocket
gophers have the same physical appearance; differences are mainly quanti¬
tative, notably the proportionately shorter foreclaws of Thomomys bottae and
the comparatively larger body sizes of Geomys bursarius and Pappogeomys cast-
anops. A large pocket gopher has 2 advantages: the bite of the incisors, which
is limited by the extreme length of the teeth, is proportionately larger and much
more effective for a larger pocket gopher; and, a large pocket gopher can with¬
stand wounds which would ordinarily prove fatal to a smaller pocket gopher.
One Pappogeomys castanops used its large incisors to bite through the skull
of one Thomomys bottae. Behavioural differences manifested in combat in the
3 species are few. Upon initial contact, pocket gophers assumed a threat posture
with the anterior portion of the body slightly elevated, the mouth open, and one
or both claws extended forward. Forward lunges with flailing claws and clashing
incisors accompanied panting sounds. This commotion lasted less than a minute,
when one animal would suddenly retreat. Deaths resulted only rarely. Soil
acted as a buffer not only from surface environmental conditions, but also be¬
tween the pocket gopher and intruders in the burrow system. If the intruder
was not removed by force, a barrage of dirtloads isolated the disturbance. Nor¬
mally, geomyid body size correlates with burrow diameter (Best, 1973) so that
smaller pocket gophers are protected from geomyids too large to enter their
burrows.

It appears from these studies that dominance is not measured only in terms
of aggressiveness and success in combat. The competitiveness of interspecific
and intraspecific interaction in geomyids may more importantly be determined
by most efficient utilization of the habitat (Vaughan and Hansen, 1964; Reich-
man and Baker, 1972; Best, 1973).

ACKNOWLEDGEMENTS

Robert L. Packard gave help and advice throughout the study; Robert Baker
made available pocket gopher live-traps; John Capeheart helped trap animals;
and Suzie Hickman typed the manuscript. I am also grateful to Texas Tech

242

THE TEXAS JOURNAL OF SCIENCE

University for use of facilities throughout the study. This study was aided with
financial support from the Society of the Sigma Xi and the Theodore Roosevelt
Memorial Fund of the American Museum of Natural History.

LITERATURE CITED

Baker, R.J., and S.L. Williams, 1972-A live trap for pocket gophers. J. Wildlife Manage¬
ment., 36:1320.

Best, T.L., 1973 -Ecological separation of three genera of pocket gophers (Geomyidae).
Ecology, 54:1311.

Cummings, M., 1946 -Grand Mesa pocket gopher study. Progress Report, 1946, Filed at
Denver Wildlife Research Center, U.S. Dept. Interior, 27 pp.

English, F., 1932-Some habits of the pocket gopher, Geomysbreviceps. J. Mammal., 13:126.

Hansen, R.M., and R.S. Miller, 1959-Observations on the plural occupancy of pocket go¬
pher burrow systems. J. Mammal., 40:557.

Hendricksen, R.L., and M.H. McGaugh, 1975 -A display unit for a live pocket gopher. Muse¬
ology, 1:1.

Hermann, J.A., 1950-The mammals of the Stockton Plateau of Northeastern Terrell
County, Texas. Texas J. Sci., 2:368.

Hickman, G.C., 1974-A behavioral study of representatives of the three genera of North
American geomyids. Ph.D. Dissertation, Texas Tech University.

Howard, W.E., and H.E. Childs, Jr., 1959— Ecology of pocket gophers with emphasis on
Thomomys bottae mewa. Hilgardia, 29:277 .

Hubbell, T.N., and C. Goff, 1940-Florida pocket gopher burrows and their arthropod in¬
habitants. Proc. Fla. Acad. Sci., 4:127.

Ingles, L.G., 1952— The ecology of the pocket gopher, Thomomys monticola. Ecology, 33:87.

Kennedy, T.E., Jr., 1964— Microenvironmental conditions of the pocket gopher burrow.
Texas J. Sci., 16:395.

Miller, R.S., 1964— Ecology and distribution of pocket gophers (Geomyidae) in Colorado.
Ecology, 45:256.

Moore, J.C., 1945-Life history notes on the Florida weasel. Proc. Florida Acad. Sci., 7:247.

Morse, A.P., 1927-A way of snake with a pocket gopher. Copeia, 164:71.

Reichman, O.J., and R.J. Baker, 19 72 -Distribution and movements of the species of
pocket gophers (Geomyidae) in an area of sympatry in the Davis Mountains, Texas,
/. Mammal., 53:21.

Scheffer, T.H., 1910-The pocket gopher. Kansas State Agr. Coll. Exp. State Bull, 19 72, 39 pp.

- , 1945-Burrow associations of small mammals. Murrelet, 26:24.

- , 1954 -Concerning the pocket gopher in mole range. Washington Agr. Exp. Sta.

Circ., 242, 4 pp.

GEOMYID INTERACTION

243

Seton, E.T., 1929 -Lives of Game Animals. Doubleday, Doran and Company, New York,
4:395.

Stebbins, R.C., 19 5 A -Amphibians and Reptiles of Western North America. McGraw-Hill,
Inc. New York. 563 pp.

Vaughan, T.A., 1961 -Vertebrates inhabiting pocket gopher burrows in Colorado. J. Mam¬
mal., 42:171.

- , and R.M. Hansen, 1964 -Experiments on interspecific competition between two

species of pocket gophers. Amer. Midland Nat., 72:444.

Wilks, B.J., 1963— Some aspects of the ecology and population dynamics of the pocket
gopher ( Geomys bursarius ) in southern Texas. Texas J. Sci., 15:241.

MITOTIC CHROMOSOMES OF TURTLES. IV. THE EMYDIDAE

by FLAVIUS C. KILLEBREW1

Department of Biology , West Texas State University
Canyon, 79016

ABSTRACT

Karyotypic data are presented on 30 species and subspecies of emydid turtles. Macro¬
chromosome morphology and diploid numbers were compared to determine possible phylo¬
genetic relationships. All New World (emydine) species had a diploid number of 50, while
Old World (batagurine) species had 52 chromosomes. The largest 13 pairs of chromosomes
were similar in morphology for all species examined. The major karyotypic difference noted
was in the number of microchromosomes. The New World species had 12 pairs of micro¬
chromosomes, while, the Old World species had 13 pairs.

The karyotypes of those emydids examined were also compared to those of several chely-
drid and testudinid species. The macrochromosomes comprising the karyotypes of these 3
families were found to be morphologically similar. Comparison of diploid numbers indicates
that testudinids, chelydrids, and Old World (batagurine) emydids are similar, while New World
(emydine) emydids differ by possessing one fewer pair of microchromosomes. Thus, the karyo¬
typic data reaffirms the karyotypic stability previously noted for cryptodiran turtles.

INTRODUCTION

Chelonians, until recently, have been relatively neglected with respect to karyo-
logical research. Previously reported diploid numbers range from 26 to 66 (Ayres,
et al., 1969; Ohno, 1967) with increases in diploid number primarily attributable
to increases in the number of microchromosomes.

The most common chromosome number for emydids is 50 (Huang and Clark,
1967; Matthey and van Brink, 1956; McKown, 1972; Stock, 1972). Emydids
with a diploid number of 52, however, include the following: Chinemys reevesii,
Clemmys japonica (Sasaki and Itoh, 19 67), Mauremys nigricans, Cyclemys dentata,
Kachuga tecta (Stock, 1972), Graptemys barbouri and G. pulchra (McKown, 1972).
There are a number of emydids, however, which have not been karyotyped, or
which have been incorrectly karyotyped. Thus, the purpose of this paper is to
present additional karyotypic data for use in analyzing relationships among mem¬
bers of this and other closely related families of turtles.

Present address: Department of Zoology, University of Arkansas, Fayetville 72701.
Accepted for publication: July 21, 1975.

The Texas Journal of Science, Vol. XXIX, Nos. 3-4, December, 1977.

246

THE TEXAS JOURNAL OF SCIENCE

MATERIALS AND METHODS

The method used for the study of emydid chromosomes is basically that of
Gay and Kayfman (1950), with modifications for use on turtles (McKown, 1972).
Karyotypes were prepared and examined from approximately 50 spreads per speci¬
men. The chromosomes were classified according to Levan, et al, (1964) and
paired on the basis of size and the position of the centromere. Differences in size
and morphology of the elements beyond the 13th pair were often difficult to
ascertain ; thus, only the largest 13 pair of chromosomes were considered as macro¬
chromosomes (Killebrew, 1975).

Specimens were obtained primarily from pet dealers and by field collection of
basking turtles using a long-handled dip net and a boat. All specimens used in this
study are preserved and catalogued in the museum collection, Department of
Biology, West Texas State University.

RESULTS

Specimens of 30 species and subspecies, representing 15 emydid genera, were
karyotyped (Table 1). All New World (emydine) genera had the same diploid
number of 50, while all Old World (batagurine) genera had a diploid number of
52 (Table 1). The largest 13 pairs of chromosomes included 8 metacentric,3 sub-
metacentric, and 2 telocentric chromosome pairs in all specimens examined. The
remaining 12 to 13 pairs were usually recognizable as metacentric or telocentric
elements (Fig. 1).

The 50 chromosome karyotype reported for Ocadia sinensis mdSiebenrockiella
crassicollis (Stock, 1972), and the diploid number of 52 for Graptemys barbouri
and G. pulchra (McKown, 1972) are not supported by this study. The 52 chromo¬
some karyotype presented for Ocadia sinensis is similar to that reported by Nakamura
(1949).

DISCUSSION

Data on emydid karyotypes presented in this study may be taxonomically im¬
portant. Comparison on diploid numbers separates emydids into New World (emy¬
dine) genera (2n = 50), including Emys orbicularis (van Brink, 1959), and Old
World (batagurine) genera (2n = 52), including Rhinoclemys. Analysis of the
macrochromosome morphology, however, reveals that all emydids are karyotypically
similar. Thus, the karyotypic datum supports the division of the Emydidae into
2 subfamilies, the Emydinae and the Batagurinae (McDowell, 1964).

Comparison of emydid (present study) and testudinid (Ohno, 1967; Sampaio,
et al., 1971; Stock, 1972) karyotypes reveals a similarity between the diploid
number (2n = 52) and morphology of batagurine and testudinid karyotypes. Emy¬
dine genera (present study) are also similar to testudinids in the morphology of

MITOTIC CHROMOSOMES OF TURTLES

247

TABLE 1

Diploid number and collection source for those species
of emydids examined. Abbreviations in the table rep¬
resent the following: M = macro chromosomes, m = micro-
chromosomes, 2n = diploid number.

Species Diploid Number Source

Mm 2n

Chinemys reevesii
Chrysemys picta
Clemmys caspica leprosa
Clemmys marmorata pallida
Cuora amboinensis
Cyclemys dentata
Deirochelys reticularia reticularia
Graptemys caglei
Graptemys barbouri
Graptemys flavimaculata
Graptemys geographica
Graptemys kohni
Graptemys nigrinoda
Graptemys oculifera

Graptemys pseudogeographica sabinensis

Kachuga smithi

Mauremys mutica

Ocadia sinensis

Pseudemys concinna texana

Pseudemys floridana hoyi

Pseudemys scripta callirostris

Pseudemys scripta elegans

Pseudemys nelsoni

Malayemys subtrijuga

Rhinoclemys funeria

Rhinoclemys punctularia melanosterna

Siebenrockiella crassicollis

Terrapene Carolina triunguis

Terrapene coahuila

Terrapene ornata

26

26

52

Midwest Reptile

26

24

50

R. McKown

26

24

50

Midwest Reptile

26

24

50

R. McKown

26

26

52

R. McKown

26

26

52

Midwest Reptile

26

24

50

Midwest Reptile

26

24

50

Guadalupe River

26

24

50

Chipola River

26

24

50

Pascagoula River

26

24

50

Buffalo River

26

24

50

Deri Brooks

26

24

50

Alabama River

26

24

50

Pearl River

26

24

50

Sabine River

26

26

52

Midwest Reptile

26

26

52

Midwest Reptile

26

26

52

Midwest Reptile

26

24

50

Guadalupe River

26

24

50

Sabine River

26

24

50

Moses Department Store

26

24

50

Sabine River

26

24

50

Midwest Reptile

26

26

52

Midwest Reptile

26

26

52

Midwest Reptile

26

26

52

F. Medem

26

26

52

Midwest Reptile

26

24

50

Pike Co., Arkansas

26

24

50

Cuatro Cienegas, Mexico

26

24

50

Randall Co., Texas

the macrochromosomes but differ by possessing one fewer pair of microchromo¬
somes. Thus, the close relationship between emydid and testudinid genera is sup¬
ported by the karyotypic data. This is interesting since ancestral emydids are thought
to be of Asiatic origin (Loveridge and Williams, 1957). If one assumes that these

248

THE TEXAS JOURNAL OE SCIENCE

Bum

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61 eo lltllliu ax«*

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At' (It) -

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t-i

V,** ::k .i:. *3

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fid 00 IS OQIm XX XX IiAaau

UlloflfliW

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(in Ktl 00 X» icxtexiin^ *%

Figure 1. Karyotypes are as follows: 1. Chinemys reeve sii, 2n = 52; 2. Chrysemys picta,
2n = 50; 3. Clemmys caspica leprosa, 2n = 50; 4. Clemmys marmorata pallida ,
2n = 50; 5. Cuora amhoinensis, 2n = 52; 6. Cyclemys dentata, 2n = 52; 7. Deiro-
chelys reticularia reticularia, 2n = 50.

MITOTIC CHROMOSOMES OF TURTLES

249

8

IIKnh

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p ♦

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A* AO f& M** afar m mA

&«&

10

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II

fttok

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ft (Mill Kiumu m

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12

tf> 28 Off

sss

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1311

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XX At

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Figure 1. (Continued) 8. Graptemys caglei, 2n = 50; 9. Graptemys barbouri, 2n =50;

10. Graptemys flavimaculata, 2n = 50; 11. Graptemys geographica, 2n=50;
12. Graptemys kohni, 2n = 50; 13. Graptemys nigrinoda, 2n = 50; 14. Graptemys
oculifera, 2n = 50; 15. Graptemys pseudogeographica sabinensis, 2n = 50.

250

THE TEXAS JOURNAL OF SCIENCE

A* V.

16

iri) il 0® t§ is M u n »

«ft Q§

17

lift! ttft ftft ml UMUt XX VM

A4M Xk

18

It

'<ri»

***^

HS «><> iM » •', :{» 4« >1* S*

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%%*•

V * Ri| ?r*7

19 il« md() IttllfUf* <>;! kK y

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«#

20

•• 4

i’^41

«W IQ S£ #U IX Xs 3« ** a* it

21

tlafiiu

22

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i\ li ®S xxntsx Ha sat *• «• «« ’*■$

23

1*19 ft! XX iff a* X* AA

st%»*

Figure 1. (Continued) 16. Kachuga smithi, 2n = 52; 17. Mauremys mutica, 2n = 52;

18. Ocadia sinensis, 2n = 52 ; 19. Pseudemys concirma texana, 2n = 50 ; 20. Pseudemys
floridana hoyi, 2n = 50; 21. Pseudemys scripta callirostris, 2n = 50; 22. Pseudemys
scrip ta elegans, 2n = 5 0; 23. Pseudemys nelsoni, 2n = 50.

MITOTIC CHROMOSOMES OF TURTLES

251

24

era*

UN IS HU

25

26

27

Uilfefc

8k

AV,

• • *■ ««

Mm(M) ••»«■«■••

** r%

II NXilft Si stit »•«*

•;V

28

flbu

DlSNHU NHKrmuu

29

QUm

30

Sttii

*»:?$•

nmumumu

mm as at

HNUMHRHmuu

Figure 1. (Continued) 24. Malayemys subtrijuga, 2n = 52; 25. Rhinoclemys funeria,
2n = 52; 26. Rhinoclemys punctularia melanosterna, 2n = 52; 27. Siebenrockiella
crassicollis, 2n = 52; 28. Terrapene Carolina, 2n = 50; 29. Terrapene coahuila,
2n = 50; 30. Terrapene ornata, 2n = 50.

252

THE TEXAS JOURNAL OF SCIENCE

ancestral emydidshad a 52 chromosome karyotype, as do those present-day Asiatic
genera that were examined, then derivation of the 50 chromosome emydid karyo¬
type would require either the deletion of 1 pair of microchromosomes or the
fusion of 2 pair of these elements. If one assumes, however, that the ancestral
emydids had 50 rather than 52 chromosomes, then some other mechanism, such
as chromosomal breakage, would be required to explain the extra pair ofbatagurine
microchromosomes. Present analyses of microchromosome morphology, however,
are not sufficient to resolve this question. Another alternative is that the ances¬
tral emydids had a diploid number other than 50 or 52. However, this is not sup¬
ported by available evidence.

The emydid karyotypes presented are also very similar to those of chelydrids
(Stock, 1972). Chelydrids, until recently, were thought to be more closely related
to the kinosternids (Williams, 1950) than to the emydids. Recent studies of serum
proteins (Friar, 1962, 1964, 1972), however, indicate that the chelydrids are
closer to the New World emydids than to kinosternids. Analysis of the morphology
of the macrochromosomes of these groups indicates that the chelydrids, emydids,
and staurotypines are karyotypically similar, while kinosternines are more diver¬
gent (Killebrew, In Press ; Stock, 1972), and the second pair of macrochromosomes
is submetacentric in the kinosternines and metacentric in the chelydrids, emydids,
and staurotypines.

A different grouping is obtained, however, when diploid numbers are compared.
According to this grouping, New World emydids (2n = 50) form 1 group, chelydrids
and Old World emydids (2n = 52) form a 2nd group, staurotypines (2n = 54) form
a 3rd group, and kinosternines (2n = 56) form a 4th group. Thus, a grouping based
on diploid number supports the theory that the chelydrids are more closely related
to emydids than to the kinosternids. The karyotypic evidence also indicates that
the chelydrids share karyotypic affinities with the kinosternids, since staurotypines
have a morphologically similar karyotype and a diploid number that differs from
the chelydrid karyotype by one pair of microchromosomes. Thus, the karyotypic
data supports the contention that the kinosternids and chelydrids arose from a
common stock which is close to the emydids.

ACKNOWLEDGEMENTS

I thank Dr. Ronald McKown for his assistance and support throughout this
study. I also thank Drs. Deri Brooks, Robert Wright, and David LaBrie for their
editorial comments on this manuscript in its several forms. Most of the turtles
examined were provided by Larry N. Lantz of Midwest Reptile and Animals
Sales, Fort Wayne, Indiana. Aid in the field colleciton of other specimens was
given by Drs. Federico Medem, Deri Brooks, and Ronald McKown.

LITERATURE CITED

Ayres, M., M. M. Sampaio, R. M. S. Barros, L. B. Dias, and O. R. Cunha, 1969-A karyological

study of turtles from the Brazilian Amazon Region. Cytogenetics, 8:401.

MITOTIC CHROMOSOMES OF TURTLES

253

Friar, W., 1962 -Comparative serology of turtles with systematic implications. Ph.D. Disserta¬
tion, Rutgers University, New Brunswick, New Jersey.

- , 1964-Turtle family relationships as determined by serological tests. In C. A. Leone

(Ed.), Taxonomic Biochemistry and Serology. Ronald Press, New York, N.Y.

- , 1972-Taxonomic relations among chelydrid and kinosternid turtles elucidated

by serological tests. Copeia, 1972:97.

Gay, H., and B. P. Kaufman, 1950-The corneal epithelium as a source of mammalian somatic
mitosis. Stain Tech., 25:209.

Huang, C. C., and H. F. Clark, 19*67 -Chromosome changes in cell lines of the box turtle
( Terrapene Carolina ) grown at two different temperatures. Canadian J. Genet. Cytol.,
9:449.

Killebrew, F. C., 1975-Mitotic chromosomes of turtles. I. The Pelomedusidae. J. ofHerp.,
9:281.

- , 197 6 -Mitotic chromosomes of turtles. III. The Kinosternidae. Herpetologica,

In Press.

Levan, A., K. Fredga, and A. Sandberg, 1964-Nomenclature for the centromeric position
on chromosomes. Hered. (Lund.), 52:201.

Loveridge, A., and E. E. Williams, 195 7 -Revision of the African tortoises and turtles of the
suborder Cryptodira. Bull. Mus. Comp. Zool., Harvard, 115:162.

Matthey, R., and J. M. van Brink, 1956— Sex chromosomes in amniota .Evol., 11:163.

McDowell, S. B., Jr., 1964-Partition of the genus Clemmys and related problems on the
taxonomy of the aquatic Testudinidae. Proc. Zool. Soc. Lond., 143:239.

McKown, R. R., 197 2 -Phylogenetic relationships within the turtle genera Graptemys and
Malaclemys. Ph.D. Dissertation, University of Texas, Austin, Texas.

Nakamura, K., 1949-A study in some chelonians with notes on chromosomal formula in
the Chelonia. Kromosomo, 5:205.

Ohno, S., 1967-Sex chromosomes and sex-linked genes. In A. Labhart and T. Mann (Eds.),
Monographs on Endocrinology, Vol. 1. Springer-Verlag, New York, N.Y., pp. 34-37.

Parsons, T: S., 1968-Variation in the choanal structure of recent turtles. Canadian J. Zool.,
46:1235.

Sampaio, M. M., R. M. S. Barros, M. Ayres, and O. R. Cunha, 1971-A karyo typical study of
two species of tortoises from the Amazon Region of Brazil. Cytol, 36:199.

Sasaki, M., and M. Itoh, 1967 -Preliminary notes on the karyotype of two species of turtles,
Clemmys japonica and Geoclemmys reevesii. Chrom. Info., 8.

Stock, A. D., 1972-Karyological relationships in turtles (Reptilia: Chelonia). Canadian J.
Genet. Cytol., 14:859.

van Brink, J. M., 1959-L’expression morphologique de la digame'tie chez les Sauropsides et
les Monotremes. Chromosoma (Berl.), 10:1.

Williams, E. E., 1950-Variation and selection in the cervical central articulations of living
turtles. Bull. Amer. Mus. Nat. Hist., 94:504.

' ■

■

'

.

EFFECTS OF HYPOPHYSECTOMY IN THE LIZARD
HOLBROOKIA PROPINQUA

by JON T. WATSON

College of Optometry, University of Houston, 77004
ABSTRACT

With more interest in comparative studies of various interests, such as reproduction, the
importance of the phylogenetic position of the reptiles is once again pointed out. Although
relatively ignored, the reptiles are being subjected to an increasing scrutiny in the labor¬
atory, but even more work is needed to get the reptilian “state of the art” to levels com¬
parable with the other vertebrate classes. This investigation provides additional information
on yet another species.

INTRODUCTION

In this investigation the effects of hypophysectomy on the testis and sperm¬
atogenesis were studied in the lizard Holbrookia propinqua. Secondary effects,
such as those concerning the androgen-sensitive epithelial tissues of the epididy-
• mus and renal sex segment, as well as fat body weight, which shows cyclicity co¬
incident with reproductive cycles, were also studied. This study provides quanti¬
fication of some effects of hypophysectomy in this species by considering the re¬
lationship of time and the appearance of these effects.

MATERIALS AND METHODS

The animals used were keeled, earless lizards, Holbrookia propinqua , collected
on Padre Island1 off the Texas coast during July and August, 1973. The experi¬
ments were begun after the animals had been held 2 wks. The lizards were housed
in plywood cages containing several cm of sand. The cage floors measured approx¬
imately 60 to 70 cm and 20 to 30 lizards were kept in a qage. Photothermal re¬
quirements were supplied by a bank of lights, controlled by a timer, providing
14 hrs of light and 10 hrs of darkness. The lights were 75 watt flood lights di¬
rected toward the floor of the cage about 15 cm from one side. This provided
a temperature gradient across the cage floor. Maximum temperature was main-

collecting was done on the Padre Island National Seashore with permission of the U.S.

National Park Service.

Accepted for publication: January 14, 1977.

The Texas Journal of Science, Vol. XXIX, Nos. 3-4, December, 1977.

256

THE TEXAS JOURNAL OF SCIENCE

tained at 35-37 C, with the lights on, falling to 20-24 C when the lights were
off. Some animals did not feed well in the laboratory, so the entire group was
force-fed meal worms to maintain a constant nutritional state.

In H. propinqua the pituitary is visible in the roof of the mouth and con¬
sequently can be removed with relative ease intraorally. A small hole was made
in the cartilage just below the gland with a dental burr, size #l/2-round. The
pituitary was then aspirated through this hole with a blunt 20 gauge needle at¬
tached to a vacuum source and no further treatment was necessary.

Testes, epididymi, kidneys, and heads were fixed in Bourns’ fluid, embedded
in paraffin and sectioned at 7 pm. Staining was done using Harris’ hematoxylin
and an eosin-phloxine counterstain. The heads were sectioned serially at 10 pm
to confirm the completeness of hypophysectomy and where pituitary tissue was
identified the data were not used.

At autopsy, total body weight was recorded as well as weights of the testes
and fat bodies. Measurements of epithelial cell heights were made randomly
using a graticule scale with 5 measurements recorded of each tissue for each ani¬
mal. Means and standard errors were calculated and t-tests performed on the
mensural data.

This study was designed, in part, to determine the time needed for responses
to pituitary ablation to occur. Two groups of animals were hypophysectomized
and examined at various intervals to determine the effects. The first group was
autopsied over a longer time base which gave an overview of the temporal re¬
lationships. This suggested the second group study, which produced additional
data, expanding the first half of the original study. The first group was autopsied
in subgroups at 0, 5, 10, 15 and 20 days and the second group at 0, 3, 5, 7 and
10 days.

RESULTS

Results of these studies are presented in Tables 1 and 2. The data show a pat¬
tern of testicular regression, and following closely, a similar decline in tissues
that are dependent upon androgen stimulation. Time periods required for the
effects of endocrine ablation to appear on reproductive structures are shown.

In the 20-day study, total body weight dropped, inexplicably, in both the
control and hypophysectomized groups at about day 10. This weight loss ap¬
peared in the organ weights and measurements for the control group but how
much it is reflected by the hypophysectomy data is not as clear. With the ex¬
ception of epididymal epithelial height, at day 10, the other measurements grad¬
ually decreased (except for the increase in fat body weight).

Testes (Fig. 1) began to lose weight between days 3 and 5 after hypophysec¬
tomy. Testis weight showed similar values in the 2 hypophysectomized groups
and was significantly reduced by day 10 in both studies. Weight loss continued
until approximately day 15, whep the testes reached their minimum weight. The

Hypophysectomy Effects - Males - 20 Days

EFFECTS OF HYPOPHYSECTOMY

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EFFECTS OF HYPOPHYSECTOMY

259

Figure 1. Effects of hypophysectomy on the testis. A is from 20 day post-hypophys-
ectomy, B is control. Note reduction in seminiferous tubule size and loss of
germinal epithelium. 500X

spermatic stage, a system of ranking spermatic activity (Table 3) showed a sim¬
ilar rate of involution, seen as a loss of organization of the germinal epithelium
and lack of sperm in the epididymus, occurring between days 3 and 5 with com¬
plete disorganization occurring by day 10 (Fig. 4).

TABLE 3

Spermatic Stage Criteria1

Stage

Histological Condition

Seminiferous Tubules

Epididymus

0

Involuted with epithelium in disarray

Atrophic, empty

1

Involuted with only spermatogonia

Atrophic, empty

2

Primary spermatogonia appearing

Atrophic, empty

3

Secondary spermatocytes and early
spermatids

Atrophic, empty

4

Transforming spermatids with a few
spermatozoa

Atrophic, empty

5

Spermatids and spermatozoa abundant

Hypertrophied, empty

6

Spermatozoa abundant and
spermatocytes reduced

Hypertrophied with
many sperm

Modified from Licht, 1967.

The effects of hypophysectomy on the androgen-maintained epithelium of
the epididymis and the renal sex segment appeared at day 5 or soon after. Cell
height reached minimum values in the epididymi by day 10 and appeared to do

260

THE TEXAS JOURNAL OF SCIENCE

HYP

0246 0246 024-6 0246 0246

0 3 5 7 10

DAYS AFTER HYPOPH YSECTOM Y

Figure 4. Effects of hypophysectomy on spermatic stage. Each dot represents the place¬
ment of 1 animal with regard to the criteria numbered 0 through 6 in Table 3.
Notice the control group (c) values tend to occur most often at 5 and 6 with a
few at the low end. The treated group (Hyp) shows the same pattern at day 3
as the controls but from day 5 on shows the reverse, suggesting the effects on
the germinal epithelium are manifested before day 5.

so in the sex segment at or soon after that time (Figs. 2 and 3). Fat body weight
did not change as rapidly as the other parameters but increased by day 20 after
hypophysectomy.

DISCUSSION

Effects of hypophysectomy on reproduction in reptiles has been described
primarily as gonadal involution. This is accompanied by regression of tissues that
are dependent upon steroid secretion by the gonad (Cieslak, 1945; Wagner, 1955;
Eyeson, 1971; Tsui and Licht, 1974). From the data presented in this study, it
can be seen that hypophysectomy in Holbrookia propinqua produces the same
general effects on the reproductive system that occur in other reptiles. Further,
it is possible to make some- statement on the time element needed to manifest
these effects.

Minimum time required to see the effects of gonadotropin depletion was ap¬
proximately 3 days. Panda and Thapliyal (1964) reported that 13 days were nec¬
essary for complete regression of the germinal tissues of testes in the lizard Calotes
versicolor and pointed out the contrast to the 9 days necessary in snakes reported
by Cieslak (1945). Data presented here show considerable reduction of the germ-

EFFECTS OF HYPOPHYSECTOMY

261

Figure 2. Effects of hypophyscctomy on the epididymus. The epithelium is much re¬
duced and the tubules are empty at 20 days after operation. A is from 20 day
post-hypophysectomy, B is control. 500X

Figure 3. Effects of hypophysectomy on the renal sex segment. There is loss of secret¬
ory material in the cells and reduction in cell height. A is from 20 day post-
hypophysectomy, B is control. 500X

inal epithelium (described as spermatic stages) beginning between 3 and 5 days
after hypophysectomy with complete involution at or soon after day 10.

Rapid regression in testicular size and spermatogenic activity after hypophys¬
ectomy was reported in the lizard Anolis carolinensis when the animals were
kept at 31 C. Only slight changes occurred in the testes of animals kept at 20 C.

262

THE TEXAS JOURNAL OF SCIENCE

Short daylengths at 31 C produced the same testicular effects as hypophys-
ectomy (Licht and Pearson, 1969). In this study, temperature was maintained
above 31 C at temperatures nearer those of the natural habitat and photoperiod
was 14 hrs light, 10 hrs dark which also approximates natural conditions.

Effects of hypophysectomy on the interstitial tissue are shown indirectly
as reductions of androgen- sensitive epithelial tissues. These tissues, the epididymis
and renal sex segment, regressed to minimal values at day 10 following hypo
physectomy. Ignoring any time needed to deplete circulating hormones (i.e. temp¬
oral differences between cessation of androgen secretion and involution of and¬
rogen dependent tissues), one could speculate that the interstitial tissue had in¬
voluted within 10 days after hypophysectomy.

In this study, as reproductive tissues regressed to conditions compatible with
those of non-reproductive lizards, the fat bodies began to increase in weight.
These changes are consistent with reports that fat body cycles correlate with
yearly reproductive cycles where fat body weights are at minimal values during
periods of gonadal activity (Dessauer and Fox, 1959; Hahn and Tinkle, 1966).

LITERATURE CITED

Cieslak, E.S., 1945 -Relations between the reproductive cycles and the pituitary gland in
the snake Thamnophis radix. Physiol. Zool., 18:299.

Dessauer, H.C., and W. Fox, 1959-Changes in ovarian follicle composition with plasma
levels of snakes during estrus. Amer. J. Physiol., 197:360.

Eyeson, K.N., 1971 -The role of the pituitary gland in testicular function in the lizard
Agama agama. Gen Comp. Endocrinol., 16:342.

Hahn, W.E., and E.W. Tinkle, 1966-Fat body cycling and experimental evidence for its
adaptive significance to ovarian follicle development in the lizard Uta stansburiana.
J.Exp. Zool, 158:79.

Licht, P., 1967 -Environmental control on annual testicular cycles in the lizard Anolis
carolinensis . I. Interaction of light and temperature in the initiation of testicular re-
crudescense. J. Exp. Zool, 165:505.

- , and A.K. Pearson, 1969-Effects of adenohypophysectomy on testicular func¬
tion in the lizard Anolis carolinensis. Biol. Reprod., 1:107.

Pandha, S.K., and J.P. Thapliyal, 1964 -Hypophysectomy in Indian garden lizard, Calotes
versicolor. Naturwissenschaften, 51:201.

Tsui, H.W., and P. Licht, 1974— Pituitary independence of sperm storage in male snakes.
Gen. Comp. Endocrinol, 22:277 .

Wagner, E.M., 1955— Effect of hypophysectomy in the turtle Chrysemys d’orbignyi. Acta.
Physiol. Latinoamer., 5:219.

CRETACEOUS (ALBIAN) AMMONITES FROM PUERTO RICO
AND ST. THOMAS

by KEITH YOUNG

The University of Texas at Austin

ABSTRACT

Except for Cuba, ammonites from the Caribbean Islands have been less than
abundant, although a few have been described from Jamaica, Curacao, Haiti,
and Trinidad. Herein are described Albian ammonites from St. Thomas of the
Virgin Islands and from Puerto Rico. Manuaniceras sp. cf. M. supani (Lasswitz)
is from the Middle Albian Robles Formation of the Comerio Quadrangle, Puerto
Rico. Eogaudryceras (?) sp., Puzosia sp. cf. P. welwitschi Choffat and de Loriol,
and Engonoceras spp. are from the pre-Robles beds, but also are most probably
Middle Albian.

Mortoniceras sp. juv. cf. M. scobinum (Van Hoepen), M. sp. juv. cf. M. nanum
Spath, and Hypophylloceras sp. juv. cf. H. ellipticum (Kossmat) are from the
Upper Albian part of the Tutu Formation, west end of St. Thomas, Virgin Islands.

INTRODUCTION

Very few ammonites have been described from the Caribbean Islands, except
for Cuba. Spath (1925a) described Campanian ammonites from Jamaica, and
Matsumoto (1966) described a Cretaceous ammonite from Curacao. Reeside (1947)
described a few Lower Senonian ammonites from Haiti, and Spath (1939) de¬
scribed latest Jurassic ammonites from Trinidad. Cuban ammonites are largely
late Jurassic (Imlay, 1942; O’Connell, 1920). Meyorhoff (1933) mentions Middle
Cretaceous ammonites from Puerto Rico, identified by J.B. Reeside, Jr. Still
another ammonite from Puerto Rico was identified as belonging to the genus
Manuaniceras, perhaps M. carbonarium, by W.A. Cobban (Sohl, 1975); it was
collected from the Fajardo Shales at the northeast end of the island. Although
many Cretaceous formations can be assigned to stages by rudists and foramini-
ferans, still, corroborative evidence is always desirable. In areas of good ammo¬
nite collecting the forms described herein would not merit a paper. Since they
constitute all but 2 of the ammonites known from the islands of St. Thomas
and Puerto Rico, their importance is emphasized by their uniqueness.

Accepted for publication: June 15, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 3-4, December, 1977.

264

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Ammonites From Puerto Rico

Early in the 1960’s I received from Norman Sohl 2 ammonites from the Cre¬
taceous of Puerto Rico. One was an oxytropidoceroid of which the Middle
Albian age was never in doubt. The second was a puzosiid, which I compared to
Austiniceras dibleyi Spath (1922) and considered either Turonian or Coniacian.
Further stratigraphic work in Puerto Rico indicated that the horizon of the
puzosiid was older than that of the oxytropidoceroid, which was Middle Albian;
the stratigraphers correctly followed the dictates of superposition from field
work and ignored my identification.

In 1968 Erie Kauffman, Norman Sohl, Bob Perkins, and I revisited this
locality (U.S. Geological Survey Mesozoic Locality 26087 = U.S. Geological
Survey Mesozoic Locality 26804) and collected more specimens of the puzosiid,
plus specimens of engonocerids and Eogaudryceras (?) sp. Upon seeing the
eogaudrycerine I changed my identification of the puzosiid at the site. I now
consider that meager and poorly preserved fauna to consist of the following forms:

Puzosia sp. cf.P. welwitschi Choffat and de Loriol, 1888.

Eogaudryceras (?) sp.

Engonoceras spp. indet.

The rock containing the fossils is a dirty sandstone, somewhat clayey, that ap¬
pears to be composed of fragments reworked largely from mafic bentonites.
It is an environment in which ammonites are seldom found. The coarse, granular,
clayey rocks preserve no shell material, and the internal molds are devoid of su¬
tures, growth lines, and most other ornamentation. A few faint ribs appear on
some specimens. Under such circumstances of preservation all identifications
are precarious and suspect.

In 1972 I suggested that this fauna was Lower Albian. This was on the basis
of work by stratigraphers indicating that the containing beds underlay undoubted
Middle Albian, and also on the identification of Eogaudryceras sp. Further study
has shown that the Puerto Rican Eogaudryceras (?) cannot be demonstrated to
be Lower Albian. Furthermore, the genus Puzosia s.s. is not supposed to extend
below the Middle Albian, which is the most likely age of Puzosia welwitschi
Choffat and de Loriol (1888) and P. cuvervillei (Meunier, 1888), the 2 species
to which the Puerto Rican puzosiid is most easily compared. This places a Mid¬
dle Albian date on this locality of the pre-Robles beds, some 3 mi north of
Aiboneto, Barranquitas quadrangle, Puerto Rico.

The previously mentioned oxytropidoceroid is from the Robles Formation,
about the middle of a very thick sequence. It is a Middle Albian Manuaniceras ,
probably M. supani (Lasswitz, 1904), well known in the Texas Comanche Peak
and Goodland Formations. The ribbing is that of older M. supani, rather than
younger supani , and indicates an age high in the Middle Albian, according to
zonations of Breistroffer (1947), and Young (1966). Thus we have an undoubted
Middle Albian fauna from the Robles Formation (Berryhill, et al., 1960; Don¬
nelly, 1970a), overlying by as much as 300 or more meters of rock an indicated
Middle Albian fauna (pre-Robles beds), but in different areas.

CRETACEOUS (ALBIAN) AMMONITES

265

St. Thomas Ammonites

The 3 micromorphs (Plate 2, Figs 19-24), all silicified, from the Tutu Form¬
ation (Donnelly, 1970b), western end of the island of St. Thomas, Virgin Islands,
are all juveniles. The specimen of Hypophylloceras, herein compared with//, sp.
group of H. ellipticum (Kossmat), is too generalized and too juvenile to identify
with precision. The 2 juvenile specimens of the genus Mortoniceras are likewise
too small to identify specifically, but small specimens of this genus, particularly
the binodose forms with ribs bifurcating at the umbilical node, are so unique
they can be dated as Upper Albian.

SYSTEMATIC DESCRIPTIONS
Phylum Mollusca
Class Cephalopoda
Order Ammonoidea
Suborder Phylloceratina Arkell, 1950
Superfamily Phylloceratacea Zittel, 1884
Family Phylloceratida Zittel, 1884
Subfamily Phylloceratinae Zittel, 1884
Genus Hypophylloceras Salfeld, 1924

Hypophylloceras sp. cf. group of//, ellipticum
(Kossmat, 1895)

PI. 2, Figs. 23,24

=. Hypophylloceras sp. juv. cf. H. velledae (Michelin) in Young, 1972, p. 469.
cf. Phylloceras ellipticum Kossmat, 1895, PI. 6, Figs, lab

Dimensions.

D H W U H/W

16.0 70.5 53.0 6.7 1.33

Remarks. The specimen illustrated on PI. 2, Figs. 23 and 24 is about the same
size as the pyritic micromorphs illustrated by Pervinquiere [1910, PI. 1 (10)] ,
but is much less well preserved. This makes it impossible to identify, although,
because it has a more rapidly expanding whorl than those illustrated by Pervin¬
quiere, it would appear to be more closely related to H. ellipticum (Kossmat, 189 5,
PI. 6, Figs, lab) and H. subalpinum (d’Orbigny, 1850 = Ammonites alpinus
d’Orbigny, 1840-42, p 283, PI. 83). The latter species was discussed by Fallot (1910)
and Gignoux (1920), and was illustrated by Spath (1923, PI. 1 , Figs, lab, 2) and
Wiedmann (1962a, PI. 16, Figs. 2ab). The St. Thomas specimen has the greater
width of the whorl more ventrad than does H. subalpinus (d’Orbigny), and in
this way also resembles//, ellipticum (Kossmat).//. cypris (Fallot and Termier, 1923,
PI. 3, Figs. 1 and 2) is less involute than the St. Thomas specimen.

The St. Thomas specimen is, then, unlike those specimens Wiedmann (1962a,
p 254) groups together as related to H. onoense (Stanton, 1895), but is very

266

THE TEXAS JOURNAL OF SCIENCE

CRETACEOUS (ALBIAN) AMMONITES

267

DESCRIPTION OF PLATE 2

1-8

Eogaudryceras ? sp. 1-6, 8, ventral and lateral views of WSA 16307;
7, WSA 16301; both from pre-Robles beds about 3 mi north of Aiboneto,
Puerto Rico. U.S.G.S. Mesozoic locality 26087.

9-12,14,15

Engonoceras ? spp. 9, 10, 12, WSA 16298; 11. WSA 16300 14, 15, WSA
16299; all from pre-Robles formation beds about 3 mi north of Aiboneto,
Puerto Rico, U.S.G.S. Mesozoic locality 26087.

13, 16-18

Puzosia sp. cf. P. welwitschi Choffat and de Loriol, 1888, 13, 16, lateral
and ventral views of WSA 16300, 17, ventral view of WSA 16302 (see also
PL 1, Figs. 7-9); 18, ventral view of WSA 16306; all from pre-Robles beds,

3 mi north of Aiboneto, Puerto Rico, U.S.G.S. Mesozoic locality 26087.

19-20

Mortoniceras sp. juv. cf. M. nanum Spath, ventral and lateral views of speci¬
men at Syracuse University from the Tutu Formation, west end of St.
Thomas, Virgin Islands, collected by T.W. Donnelly.

21-22

Mortoniceras sp. juv. cf. M. scobinum (Van Hoepen), lateral and ventral
views of a specimen at Syracuse University from the Tutu Formation,
west end of St. Thomas, Virgin Islands, collected by T.W. Donnelly.

23-24

Hypophylloceras sp. cf. ellipticum (Kossmat), ventral and lateral views of
a specimen at Syracuse University from the Tutu Formation west end of
St. Thomas, Virgin Islands, collected by T.W. Donnelly.

1-16, 18, X 1; 17, X 0.5; 19-24, X 2.

268

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similar to H. escragnollense (Breistroffer, 1947, p 71 = Phylloceras ellipticum
Parona and Bonarelli, 1897, PI. 1, Figs. 7abc, non Kossmat, 1895). All that can
be said about the St. Thomas specimen, then, is that it belongs to that Albian
group of Hypo phylloceras spp., which is characterized by a more rapidly ex¬
panding whorl that does not taper ventrad as does the whorl of H. subalpinus
(d’Orbigny), but is fully rounded as in H. ellipticum (Kossmat) and//, escragnolles
(Breistroffer). In 1972 I incorrectly compared this specimen to Hypo phylloceras
sp. cf. H. velledae (Michelin, 1838).

Horizon and Locality. The specimen of Hypophylloceras from St. Thomas,
Virgin Islands, is from the Tutu Formation, west end of the Island. The age is
Albian, as already pointed out by Donnelly (1970b), but can be determined as
Upper Albian on the basis of 2 associated specimens of Mortoniceras.

Suborder Lytoceratina Hyatt, 1889
Superfamily Lytocerataceae Neumayr, 1875
Family Tetragonitidae Hyatt, 1900
Subfamily Gaudryceratinae Spath, 1927
Genus Eogaudryceras Spath, 1927
Eogaudryceras (?) sp.

PI. 1, Figs. 4, 5; PI. 2, Figs. 1-8

Dimensions.

Specimen

D

H

W

U

H/W

WSA- 16301

48.0

54.5

58.5

24.0

0.93

33.5

49.0

51.0

18.7

0.96

23.5

49.0

62.0

27.5

0.79

Remarks. According to Wiedmann (1962b) the primary difference between
Eogaudryceras and Tetragonites is the suture, not recoverable on the Puerto
Rican specimens, and according to Murphy (1967) the extremely prosiradiate
restrictions in Tetragonites. There are no restrictions on the Puerto Rican spec¬
imens, which would indicate their assignment to Eogaudryceras , except that
many specimens of Tetragonites , much better preserved than those from Puerto
Rico, do not show the constrictions (Wiedmann, 1962b, PI. 14). Perhaps their
absence is due to poor preservation of the fossils.

That Tetragonites has a more symmetrical first lateral lobe and a broader and
better developed suspensive lobe than Eogaudryceras seems undoubted (Wied¬
mann, 1962b). Furthermore, Eogaudryceras lacks a good internal lateral lobe,
whereas Tetragonites has the well developed internal lateral lobe. But these
features cannot be recovered on the material from Puerto Rico.

Some lineages of Tetragonites become more depressed through the Albian
[e.g., lineage of T. timotheanus (Pictet, 1848)] , whereas Eogaudryceras, leading
to Gaudryceras , becomes less depressed. The Puerto Rican specimens are less

CRETACEOUS (ALBIAN) AMMONITES

269

depressed than T. timotheanus, but more depressed than most specimens of
Eogaudryceras, and some of them have slightly trapezoidal whorls sections,
leading one to believe that they are not far from the divercation of the 2 genera,
and thus early in the generic lineages (e.g., probably Lower Albian or very
earliest Middle Albian).

Horizon and Locality. All specimens are from the same locality and horizon,
pre-Robles beds, about 3 mi north of Aiboneto, Puerto Rico, U.S.G.S. Meso¬
zoic locality 26087 (= U.S.G.S. locality 26804), from a road cut on secondary
road 0.85 mi ESE of Escuela Segunda Unidad de la Lama and about 2.9 mi
airline due north of Aiboneto, Borrio Hunduras, Municipio de Barranquitas,
Puerto Rico meter grid 170, 175 X 38, 700, Barranquitas Quadrangle (Sohl, 1975).

Suborder Ammonitina
Superfamily Desmocerataceae Zittel, 1895
Family Desmoceratidae Zittel, 1895
Subfamily Puzosiinae Spath, 1922
Genus Puzosia Bayle, 1878

Puzosia sp. cf. P. welwitschi Choffat and de Loriol, 1888.

PI. 1, Figs. 3, 6-9; PI. 2, Figs. 13, 16-18
=. Puzosia sp. cfr. planulata Bayle, 1878 in Young, 1972, p. 469
cf. Puzosia welwitschi Choffat and de Loriol, 1888, PI. 2, Fig. 3.
cf. Desmoceras cuvervillei Meunier, 1888, PI. 1 , Fig. 3.

Dimensions.

Specimen

D*

H

W

U

H/W

WSA- 16302

160.0

42.0

33.7

24.5

1.25

100.0

45.0

35.5

24.5

1.27

60.0

48.5

36.5

24.5

1.33

WSA- 16304

49.5*

39.0*

1.27

WSA- 16306

76.5

49.0

36.0

28.0

1.36

45.5

48.5

37.5

25.0

1.29

*Figures marked with an asterisk are in mm. Other figures are percent¬
ages or a ratio (H/W).

Remarks. Matsumoto’s (1954) remarks on the differences between Puzosia
(Anapuzosia) and Puzosia (Puzosia) give one the impression that the two sub¬
genera can only be differentiated on large specimens. If the statement concern¬
ing the age of the Puerto Rican specimens of Eogaudryceras (?) sp. is correct,
then the Puerto Rican fauna from Aiboneto is too old to include Puzosia plan¬
ulata Bayle, or most other specimens of Puzosia. Matsumoto (1954) does point
out that P. planulata has, at least in some younger stages, a higher whorl section
than do specimens that can definitely be referred to Anapuzosia (e.g .,P. mayor-

270

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7

CRETACEOUS (ALBIAN) AMMONITES

271

DESCRIPTION OF PLATE 1

1, 2 Manuaniceras sp. cf. M. supani (Lasswitz), lateral views of latex peels of 2
uncollectable specimens (WSA 16297) from the Robles Formation, near
the middle of the bedded sequence on the Convento Road north of Com-
erio, Comerio Quadrangle, Puerto Rico.

3, 6-9 Puzosia sp. cf. P. welwitschi Choffat and de Loriol, 1888, 3, 6, lateral
and ventral views of WSA 16304; 7-9, lateral and ventral views of WSA
16302 (see also PI. 2, Fig. 17); both from pre-Robles formation beds
about 3 mi north of Aiboneto, Puerto Rico, U.S.G.S. Mesozoic locality
26087.

4, 5 Eogaudryceras ? sp. lateral and ventral views of WSA 16308, from the pre-
Robles beds about 3 mi north of Aiboneto, Puerto Rico, U.S.G.S. Mesozoic
locality 26087.

1-5, X 1.0; 6, X 0.5; 7-9, X 0.4.

272

THE TEXAS JOURNAL OF SCIENCE

iana Jacob, 1908, PL 16, Fig. 2, not d’Orbigny). The Puerto Rican specimens
are all higher than wide, H/W ranging from 1.25 to 1.35, although the spec¬
imens with higher ratios may be flattened by sedimentary load. Constrictions
are more frequent than in Puzo sia (Puzosia) mayoriana (d’Orbigny) (Spath, 1923,
Text-fig. 10, PL 1 , Figs. 9ab, lOab). The Puerto Rican specimens have an evenly
arched venter, compressed whorl section, and vertical umbilical wall. Further¬
more, the constriction is a definite sulcus following a major rib that truncates
normal ribbing dorsad. The constrictions do not have the linguiform ventrad
projection of P. mayoriana (Spath, 1923, pp 43, 44, Text-Fig. 10), P. planulata
Bayle, or P. crebrisulcata Kossmat (1898, PL 17, Figs. 4ab), but are more like
those of P. welwitschi Choffat and de Loriol (1888, PL 2, Fig. 3), but less sig¬
moid. They are nearly straight up to a diameter of 80 mm., more like those of
P. cuvervillei (Meunier, 1888, PI. 1, Fig. 3), but the latter specimen is so small
as to prevent proper identification.

Horizon and Locality. All specimens are from the same locality and horizon,
pre-Robles beds, about 3 mi north of Aiboneto, Puerto Rico, U.S.G.S. Meso¬
zoic locality 26087 [see more complete description under Eogaudryceras (?) sp.] .

The species with which the Aiboneto specimens are compared are mostly
Upper Albian, and Puzosia does not range below the Middle Albian. Choffat and
de Loriol (1888, p. 23) list P. welwitschi as occurring in the same beds with
Pervinquieria inflata and Douvilleiceras [Acanthoceras mamillare (Schloth.)? of
Choffat and de Loriol, 1888, p 24, PL 3, Figs, labc] , without indication of rela¬
tive age. It could be as low as Middle Albian, but they list no Lower Albian fos¬
sils. Meunier (1888, p 62) likewise lists P. cuvervillei with uSchloenbachia in¬
flate i. Sow”, but his “S. inflata ” (PL 1, Fig. 1, not Fig. 2) appears to be a Mos-
jisovicsia, and is probably Middle Albian. Since there is no evidence of large ribs
or horns in the adult stages of the Puerto Rican form, it does not seem advisable
to assign it to Anapuzosia. The 2 species to which it can best be compared both
seem to be Middle Albian, although an Upper Albian age for P. welwitschi Chof¬
fat and de Loriol is not eliminated.

Superfamily Hoplitaceae H. Douville, 1890
Family Engonoceratidae Hyatt, 1900
Genus Engonoceras Neumayr & Uhlig, 1881
Engonoceras (?) spp.

PL 2, Figs, 9-12, 14, 15

Remarks. There are at least 2 species of Engonoceras in the collection from
near Aiboneto, P.R., a species with an acute keel (PL 2, Figs. 9-12) and a tabu¬
late species (PL 2, Figs. 14, 15). The smaller specimen of the species with an
acute keel shows very faint, ventrad costae, as in the juveniles of Engonoceras
stolleyi Bohm (Hyatt, 1903, PL 24, Fig. 4), but unlike E. stolleyi this specimen
would appear to have an acute keel. Many engonocerids have tabulate venters,
and the fragment illustrated herein (PL 2, Figs. 14, 15) is not further identifiable.

CRETACEOUS (ALBIAN) AMMONITES

273

Horizon and Locality. All specimens are from the same locality and horizon,
pre-Robles beds, about 3 mi north of Aiboneto, Puerto Rico, U.S.G.S. Mesozoic
locality 26087 [see more complete description under Eogaudryceras (?) sp.] .

Superfamily Acanthocerataceae Hyatt, 1900
Family Brancoceratidae Spath, 1933
Genus Manuaniceras Spath, 1925a

Manuaniceras sp. cf .M. supani (Lasswitz)

PL 1, Figs. 1,2

=. Manuaniceras sp. cf. supani (Lasswitz) in Young, 1972, p 469.

For synonomy of M. supani (Lasswitz) see Young, (1966), p 103.

Remarks. Manuaniceras supani (Lasswitz) is differentiated from other species
of Manuaniceras by its ribs overhanging the adjacent sulcus during some stage
of growth. Otherwise, the shells are moderately evolute with high whorls (H/W
ranging from 1.5 in juveniles to 2.4 or more in larger specimens), and they have
ribs that range from sigmoid to straight, with a forward swing near the venter;
ribs do not join the keel.

The pictures of the Puerto Rican specimens are lateral views only, but show
at some growth stages the walls of the ribs overhanging the sulcus, a feature di¬
agnostic of the species.

In older members of the species the overhang is restricted to a rather narrow
growth stage, but in younger members rib overhang may occur throughout that
ontogeny beyond a diameter of 60 mm. The Puerto Rican form belongs to the
older part of the lineage and is from high in the Middle Albian, according to zon-
ation schemes of Breistroffer (1947) or Young (1966), or older, but not Lower
Albian.

Horizon and Locality. From the Robles Formation near the middle of the
bedded sequence on the Convento Road north of Comerio, Comerio Quadrangle,
Puerto Rico.

The indicated age is Middle Albian and most likely the upper part of the Mid¬
dle Albian.

Subfamily Mortoniceratinae Spath, 1925
Genus Mortoniceras Meek, 1876

Mortoniceras sp. juv. sp. cf. M. nanum Spath, 1933
PI. 2, Figs. 21,22

= Mortoniceras sp. juv. cf. M. minor Spath in Young, 1972, p 469.
cf. Mortoniceras (Pervinquieria?) nanum Spath, 1933, PI. 43, Figs. 6ab.

Dimensions.

H W H/W

4.2* 5.4* 0.78

*These figures are in mm.

274

THE TEXAS JOURNAL OF SCIENCE

Remarks. The specimens of Mortoniceras from St. Thomas are too small to
identify specifically. Although not readily visible in the figure, this specimen
has binodose ribs bifurcating rectiradially from the umbilical node — a feature
very typical of Mortoniceras.

Horizon and Locality. From the Tutu Formation, west end of the island of
St. Thomas, Virgin Islands. The specimen is in the possession of T.W. Donnelly.

Mortoniceras sp. juv. sp. cf .M. scobinum
(Van Hoepen, 1942)

PI. 2, Figs. 19,20

-Mortoniceras sp. juv. cf. scotia (typographical error) (Van Hoepen) in
Young, 1972, p 469.

cf. Pervinquieria scobina Van Hoepen, 1942, pill, Figs. 92 and 93.
Dimensions.

H W H/W

5.1* 6.1* 0.83

*These figures are in mm.

Remarks. This specimen is more densely costate than the specimen compared
to M. nanum Spath; its ribs are slightly sigmoid, and swing more ventrad near the
venter. It likewise is too juvenile to identify accurately, but is typical of minute
specimens of Mortoniceras that can be found in western Europe, Angola, Mad¬
agascar, and Texas. They are always Upper Albian. One reason that identifica¬
tion is difficult is because such small binodose juveniles may become trinodose
prior to a diameter of 50 mm — Pervinquieria and some Deiradoceras — or they
may remain binodose to diameters of 200 mm or more — Mortoniceras (=Leonites
Spath, 1932).

Horizon and Locality. From the Tutu Formation, west end of the island of
St. Thomas, 4 Virgin Islands. The specimen is in the possession of T.W. Donnelly.

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/ / / ,

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O’Connell, M., 1920-The Jurassic ammonite fauna of Cuba. Am. Mus. Nat. Hist. Bull.,
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Liquria occidentale. Palaeontographia Italia, 2:53, plates 10-14.

Pervinquiere, L., 1910-Sur quelques ammonites du Cretace' Algerien. Mem. Soc. Geol.
France, (Paleont.) 1 7 (Mem. 42), Paris, 86 pp., 7 tables.

Pictet, F.J., 1848 -Description des Mollusques fossiles qui se trouvent dans les Gres Verts
des environs de Geneve. I. Cephalopodes. Mem. Soc. Phys. and Hist. Nat., Geneve, 1 1:257,
plates 1-15.

Reeside, J.B., Jr., 1947-Upper Cretaceous ammonites from Haiti. U.S. Geol. Survey, Prof.
Paper 214-A, 5 pp., 3 plates.

Salfeld, H., 1924 -Die Bedeutung der Konservativstamme fur die Stammesentwicklung
der Ammonoideen , Leipzig, Verlag von Max Weg, 16 pp., 16 plates.

Sohl, N., 1975-Letter to Keith Young dated May 22, 1975.

Spath, L^F., 1922-On the Senonian ammonite fauna of Pondoland. Roy. Soc. So. Africa,
Trans., 10(2): 1 13, plates 5-9.

- , 1923-A monograph of the Ammonoidea of the Gault, Part 1; Palaeontological

Society, 75:1, figures 1-14, plates 1-4.

- , 1925a— On Senonian Ammonidea from Jamaica. Geol. Mag., 62:28, 1 plate.

- , 1925b -On the Upper Albian Ammonoidea from Portuguese East Africa, with

an appendix on Upper Cretaceous ammonites from Maputoland. Ann. Transvall Mus.,
11:179, 10 plates.

— - — , 1927-Revision of the Jurassic cephalopod fauna of Kachh (Cutch). Part I:

Mem. Geol. Surv. India, Paleontologia Indica, Memoir 2, 9(1): 1 , plates 1-7.

- 1932-A monograph of the Ammonoidea of the Gault, Pt. 9 . Palaeonto graphical

Soc., 84:381, figures 125-140, plates 37-42.

- , 1933-A monograph of the Ammonoidea of the Gault, Pt. X. Palaeontographical

Soc., 85:41 1, figures 141-152, plates 43-48.

- , 1939-On some Tithonian ammonites from the northern range of Trinidad,

B.W.I. Geol. Mag., 76:187.

Stanton, T.W., 1895-The fauna of the Knoxville beds. U.S. Geol. Survey Bull. 133, 85 pp.,
20 tables.

Wiedmann, J., 1962a-Die systematische stellung von Hypophylloceras Salfeld. N. Jb.
Geol. Paldont. Abh., 115:243, 5 figures, plate 16.

- , 1962b-Ammoniten aus der Vascogotischen Kreide (Nordspanien) I. Phyllo-

ceratina, Lytoceratina. Palaeonto graphica, Vol. 118, Lieferung4-6, pp. 119-237, 58 figures,
plates 8-14.

CRETACEOUS (ALBIAN) AMMONITES

277

Young, K., 1966-Texas Mojsisovicziinae (Ammonoidea) and the zonation of the Freder¬
icksburg. Geol. Soc. Amer., Mem. 100, 225 pp., 21 figures, 38 plates, 5 tables.

- , 1972-Ammonites from Puerto Rico and St. Thomas, Isla Margarita, Venezuela,

VI Conferencia Geologica del Caribe, p. 469, figure 1.

'

ii M

IDEAL GAS THERMODYNAMIC FUNCTIONS OF THE 2-HALO-
PROPENES

by G.A. CROWDER and ROYCE W. WALTRIP

Department of Chemistry ,

West Texas State University , Canyon 79016

ABSTRACT

The thermodynamic functions Gibbs free energy function, enthalpy function, entropy,
and heat capacity were calculated for 2-fluoropropene, 2-chloropropene, 2-bromopropene,
and 2-iodopropene in the ideal gas state at one atm pressure and selected temperatures.

INTRODUCTION

Thermodynamic properties of the 2-halopropenes have not been reported in
the literature. Vapor-state infrared spectra and barriers to internal rotation of
the methyl groups have recently become available for the 2-halopropenes, and
these data have made possible the calculation of ideal gas thermodynamic func¬
tions for these compounds. The functions (G°-Hg)/T, (H°-H§)/T,S°, and C°
were calculated for each of the four 2-halopropenes in the ideal gas state at 1
atm pressure and selected temperatures. The contributions of translation, over-all
rotation, and vibration were calculated with standard formulas of statistical
thermodynamics (Pitzer, 1953). These calculations were based on the rigid ro¬
tator-harmonic oscillator model. The equations that were used are summarized
in the Appendix. The contributions of restricted internal rotation of the methyl
groups were taken from the tables of Pitzer and Gwinn (1942). 1 No corrections
were made for anharmonicity, etc., because no experimental heat capacity or en¬
tropy data were available for use in making those corrections.

RESULTS

2-Fluoropropene

Pierce and O’Reilly (1959) have obtained moments of inertia of this molecule
from the microwave spectrum, and have calculated a methyl torsional barrier of
2432 cal/mole. However, they used the value 3.11 amu A2 for the moment of

1 A computer program that obtains the contributions of restricted internal rotation from
the Pitzer and Gwinn tables has been described by Crowder and Riley (1972).

Accepted for publication: May 13, 1977.

The Texas Journal of Science, Vol. XXIX, Nos. 3-4, December, 1977.

280

THE TEXAS JOURNAL OF SCIENCE

inertia, Ia, of the methyl group about its axis, a value they assumed to be the
same as in propene. However, Fateley and Miller (1963) pointed out that an ac¬
curate value of the structural factor F can be obtained from the torsional fre¬
quency and the value of the reduced barrier height, s, obtained from microwave
work. Fateley and Miller’s (1963) value of F = 5.620 cm-1 yielded Ia = 3.205
amu A2 and a methyl barrier (V3) of 2340 cal/mole. The reduced moment of
inertia2 of the methyl group was calculated from their Ia to be 3.00 amu A2 . (An
uncertainty of 0.05 amu A2 in the reduced moment results in an uncertainty of
0.04 cal or less in the thermodynamic functions.) Thfe vibrational assignment was
taken from Crowder and Smyrl (1971). The molecular data used in the calcula¬
tions are listed in Table 1 and the thermodynamic functions for 2-fluoropropene
at selected temperatures are given in Table 2.

TABLE 1

Molecular Constants of 2-Fluoropropene

Property

Value

Molecular weight

Principal moments of inertia

60.071

IA (amu A2)

49.71

Ig (amu A2)

55.91

1^ (amu A2)

102.60

Reduced moment of inertia (amu A2)

3.00

Potential barrier, V3 (cal/mole)

2340

Vibrational frequencies (cm-1)

3141,3059,3012

2975,2942, 1687

1448, 1448, 1429

1401, 1270, 1048

1008,944, 862

846,629,472

404,352

2- Chloropropene

Unland, et al., (1965) have determined the principal moments of inertia of
CH2 = C35C1CH3 and CH2 = C37C1CH3. These authors gave a value of 3.110
amu A2 for la, which is the result of their assumed value of 111.20° for the
C-C-H angle involving the methyl hydrogens. From their F value (5.62 cm-1),
we calculated a reduced moment of inertia of the methyl group of 3.00 amu A2.
The barrier height obtained by Unland, et al., (1965) was used in the calculations,
2

The relation between the reduced moment of inertia, If, and I^is

lr = la[l-la|(VIg)l

where Ag is the cosine of the angle between the axis of the internal top and the gth principal
axis and Ig is the g*h principal moment of inertia.

THERMODYNAMIC FUNCTIONS, OF THE 2-HALOPROPENES

281

TABLE 2

Thermodynamic Functions for 2-Fluoropropene in the Ideal Gas State at 1 atm

T

O—

K

-(G°-Hg)/T

(H°-H§)/T

S°

C°

cal/deg/mole

cal/deg/mole

cal/deg/mole

cal/deg/mole

273.15

55.53

11.64

• 67.17

17.19

298.15

56.59

12.14

68.73

18.25

300

56.66

12.18

68.84

18.33

400

60.44

14.22

74.66

22.25

500

63.83

16.17

80.00

25.64

600

66.94

18.00

84.94

28.50

700

69.85

19.67

89.52

30.91

800

72.56

21.22

93.78

32.99

900

75.15

22.62

97.77

34.78

1000

77.61

23.91

101.52

36.32

and the vibrational assignment was taken from Hunziker and Gunthard (1965).
These latter authors used a value of F = 5.502 cm"1 to calculate the barrier
height, but they calculated this value from structural data transferred from pro¬
pylene and vinyl chloride. The F value obtained by Unland, et al., (1965) is con¬
sidered more accurate since it is the same as for 2-fluoropropene and very nearly
the same as for 2-bromopropene (next section).

The molecular constants used for 2-chloropropene are listed in Table 3 and
the thermodynamic functions are given in Table 4.

TABLE 3

Molecular Constants of 2-chloropropene

Property

Value

Molecular weight

Principal moments of inertia

76.526

l\ (amu X2)

54.52^

54.52b

Ig (amu X2)

101. 43a

104.22b

\q (amu X2)

152. 99a

155.78b

Reduced moment of inertia (amu X2)

3.00

Potential barrier, V3 (cal/mole)

2671

Vibrational frequencies (cm"1)

3121, 3025, 2992, 2973

2940, 1645, 1450, 1450

1424, 1382, 1184, 1046

999, 926, 879, 692

641,434, 396, 343

aMoments of inertia for CH2 = CC135CH3
^Moments of inertia for CH2 - CC137CH3

282

THE TEXAS JOURNAL OF SCIENCE

TABLE 4

Thermodynamic Functions for 2-Chloropropene in the Ideal Gas State at 1 atm

H

o

-(G°-Hg)/T

(H°-Hg)/T

S°

cp

cal/deg/mole

cal/deg/mole

cal/deg/mole

cal/deg/mole

273.15

57.36

11.76

69.12

17.62

298.15

58.42

12.29

70.71

18.72

300

58.48

12.34

70.82

18.78

400

62.33

14.46

76.79

22.72

500

65.79

16.47

82.26

26.17

600

68.94

18.33

87.27

28.98

700

71.91

20.02

91.93

31.33

800

74.68

21.57

96.25

33.36

900

77.31

22.98

100.29

35.10

1000

79.79

24.26

104.05

36.62

2-Bromopropene

Benz, et al, (1966) determined, by microwave spectroscopy, the principal
moments of inertia of CH2 = C7 9 BrCH3 and CH2 = C8 1 BrCH3 . The reduced
moment of inertia was calculated from their F value of 5.60 cm-1, and their
barrier height was used for the methyl torsion.The barrier has also been calculated
from the infrared torsional frequency (Crowder and Waltrip, 1977), and the 2
values agree within the uncertainties involved.

A vibrational assignment has been made for 2-bromopropene by Meyer and
Gunthard (1967). The far infrared spectrum has also been obtained by one of
the present authors in the region below 650 cm-1 with a Perkin-Elmer 301 far
infrared spectrophotometer, through the courtesy of Dr. Donald W. Scott. Our
frequencies agree well with those of Meyer and Gunthard, except for the C=C-C
bending vibration. This is a type B band with branches at 343 and 355 cm"1 and
a maximum between the 2 branches, at 350 cm"1 in the vapor-state spectrum.
Our liquid-state value is 351 cm"1 , and the liquid-state Raman value is 354 cm"1
(Kirrman, 1939). The value reported by Meyer and Gunthard, (1 967) is 335 cm"1 ,
which differs significantly from our value. The spectrum shown by these authors
shows this band at ca. 350 cm"1 , so the 335 cm"1 value they reported is probably
a typographical error. The other frequencies below 650 cm"1 differ by 2 cm"1
at the most.

The molecular constants used in the calculations are listed in Table 5. The vi¬
brational frequencies above 600 cm"1 were taken from Meyer and Gunthard (1967)
and those below 600 cm"1 were taken from this work. The resulting thermo¬
dynamic functions are given in Table 6.

2-Iodopropene

Structural data have not been obtained for 2-iodopropene, so the molecular
parameters were transferred from 2-chloropropene (Unland, et al., 1965), and

THERMODYNAMIC FUNCTIONS OF THE 2-HALOPROPENES

283

TABLE 5

Molecular Constants of 2-Bromopropene

Property

Value

Molecular weight

Principal moments of inertia

120.98

IA (ainu A2)

54.62a

54.62b

Ig (amu A2)

161.1 0a

162. 40b

I(- (amu A2)

212.7 8a

214. 08b

Reduced moment of inertia (amu A2)

3.01

Potential barrier, V3 (cal/mole)

2695

Vibrational frequencies (cm-1)

3115, 3010, 2987

2972, 2930, 1640

1443, 1439, 1405

1379, 1170, 1045
996,925,883,680

550,412, 350, 299

aMoments of inertia for CH2 = CBr79CH3
^Moments of inertia for CH2 = CBr81CH3

TABLE 6

Thermodynamic Functions for 2-Bromopropene in the Ideal Gas State at 1 atm

T

°K

-(G°-Hq)/T

(H°-Hq)/T

S°

C°

c.p

cal/deg/mole

cal/deg/mole

cal/deg/mole

cal/deg/mole

273.15

59.75

12.13

71.88

18.12

298.15

60.84

12.67

73.51

19.19

300

60.91

12.71

73.62

19.26

400

64.86

14.85

79.71

23.17

500

68.40

16.85

85.25

26.47

600

71.64

18.69

90.33

29.22

700

74.64

20.37

95.01

31.54

800

77.47

21.88

99.35

33.52

900

80.12

23.28

103.40

35.25

1000

82.64

24.55

107.19

36.74

the C-I bond length was assumed to be the same as in vinyl iodide, 2.089 A
(Chemical Society, 1958). The principal moments of inertia were calculated from
those structural parameters. For internal rotation of the methyl group, the F
value was taken to be 5.60 cm-1 , which is the value for 2-bromopropene. The
F value for 2-fluoropropene and 2-chloropropene, 5.62 cm"1, is only slightly
higher. The barrier height has been calculated by Crowder and Waltrip (1977)

284

THE TEXAS JOURNAL OF SCIENCE

from the observed torsional frequency. The vibrational frequencies were taken
from Meyer, et al., (1969). The molecular constants are listed in Table 7 and the
thermodynamic functions are given in Table 8.

TABLE 7

Molecular constants of 2-Iodopropene

Property

Value

Molecular weight

Principal moments of inertia

167.98

I^(amu A2)

53.23

Ig(amu A2)

218.83

I^(amu A2)

268.87

Reduced moment of inertia (amu A2)

3.01

Potential barrier, V3 (cal/mole)

2580

Vibrational frequencies (cm-1)

3103,2994,2974

2967, 2928, 1627

1457, 1436, 1406

1377, 1159, 1056

991,926,893,701

514,389,318,271

TABLE 8

Thermodynamic Functions for 2-Iodopropene in the Ideal Gas State at 1 atm

T

°k

-(G°-Hq)/T

(H°-Hq)/T

S°

C°

P

cal/deg/mole

cal/deg/mole

cal/deg/mole

cal/deg/mole

273.15

61.47

12.38

73.85

18.35

298.15

62.58

12.92

75.50

19.37

300

62.67

12.96

75.63

19.44

400

66.68

15.07

81.75

23.25

500

70.26

17.04

87.30

26.50

600

73.54

18.84

92.38

29.24

700

76.56

20.50

97.06

31.55

800

79.39

22.01

101.40

33.54

900

82.07

23.39

105.46

35.26

1000

84.61

24.65

109.26

36.75

CONCLUSION

Ideal gas thermodynamic functions have been calculated for the 2-halopro-
penesat 1 atm pressure. There are no experimental data available for comparisons,
but calculations have yielded accurate values for other compounds. Recently, for

fHERMODYNAMIC FUNCTIONS OF THE 2-HALOPROPENES

285

example, Chen, et al, (1975) used this method to calculate the thermodynamic
functions for CF3CH3, which has only 1 atom fewer than the 2--halopropenes.
Their calculated entropy of 63.82 cal K"1 mol-1 at 224.4K agrees well with the
experimental value of 63.89 cal K'1 mol-1 determined by Russell, et al, (1944).
The calculated entropy of CF3CF3 at 2 temperatures (Chen, et al, 1975) also
agrees well with the experimental values of Pace and Aston (1948).

Thermodynamic functions were calculated for both isotopic species of 2-chloro
and 2-bromopropene and were averaged according the isotopic abundance. The
difference between the functions of the 2 species amounted to only 0.04 cal for
2-chloropropene and 0.01 cal for 2-bromopropene.

ACKNOWLEDGEMENT

The authors are grateful to The Robert A. Welch Foundation, Houston, Texas,
for financial support of this work.

APPENDIX

The equations used to calculate the thermodynamic functions are summarized
here. All equations result in calorie units.

Heat Capacity

translation, rotation, and vibration:

20

CJ = 7.9487 + 2 1 .4388 (£/ T)e

i=l

1 .43 8 8

^i/T/(e 1 *43 8 8ZVT_i)2

Enthalpy function

translation, rotation, and vibration:

20

H0-H: =7.9487 + 2 1.9872T/(l-e
- i=l

T

i=l

Entropy

translation, rotation;free internal rotation, and vibration:

S° = (18.3025 + 2.2878n)log T-2.3493 + 3.0346n + 6.8634 log M
+ 2.2878 log {I ! I2 13 x 101 1 7 [ H (Irxl038)l 1-4.5756 log (a II ar)

+ 2 — — — —

i=lel .4388Fj/T

20 2.859lFi/T

-4.5756 log (l-e-':4388t'i/T)

286

THE TEXAS JOURNAL OF SCIENCE

Gibbs Free Energy Function

The Gibbs free energy function was obtained from the entropy and enthalpy
function:

_(0-Ho‘) = s--(H-Ho-)

T T

Restricted Internal Rotation

The restricted internal rotational contributions were taken from the tables
of Pitzer and Gwinn (1942), which have as variables V/RT and 1/Qf, where V
is the barrier to internal rotation and Qf is the partition function for free internal
rotation. The tables for heat capacity and enthalpy function lists the direct con¬
tribution of the internal rotation, but the table for entropy lists the decrease
from free internal rotation.

Definitions of Variables in the Equations
n = number of internal rotations
T = absolute temperature
vx = the i^1 vibrational wavenumber (cm"1 )

M = molecular weight (g)

I ! I2 13 = product of the three principal moments of inertia (g cm2)

Ir = reduced moment of inertia of the r^ rotating group (g cm2 )

o = symmetry number for overall rotation

ar = symmetry number for the r^1 internal rotation

LITERATURE CITED

Benz, H.P., A. Bauder, and Hs.H. Gunthard, 1966-/. Mol. Spectrosc 21:165.

Chemical Society Special Publication No. 11, 1958 -Tables of Interatomic Distances and
Configuration in Molecules and Ions, London.

Chen, S.S., A.S. Rodgers, J. Chao, R.C. Wilhoit, and B.J. Zwolinski, 1975-/. Phys. Chem.
Ref Data, 4:441.

Crowder, G.A., and P. Riley, 1972-/. Chem. Educ., 49:30.

- -, and N. Smyrl, 1971-/. Mol. Spectrosc., 40:117.

— - , and R.W. Waltrip, 1911 -Texas J. ScL, 28:219.

Fateley, W.G., and F.A. Miller, 1961 -Spectrochim. Acta., 17:857.

- , and - — — — , 1963 -Spectrochim. Acta., 19:611.

Hunziker, H., and Hs.H. Gunthard, 1965 -Spectrochim. Acta., 21:51.

Kirrman, A., 1939 -Bull. Soc. chim. France, 6:841.

Meyer, R., H. Hunziker, and Hs.H. Gunthard, 1969 -Spectrochim. Acta., 25A:295.

THERMODYNAMIC FUNCTIONS OF THE 2-HALOPROPENES

287

- , and Hs.H. Gunthard, 1967 -Spectrochim. Acta., 23A:2341.

Pace, E.L., and J.G. Aston, 1948 -7. Amer. Chem. Soc., 70:1326.

Pierce, L., and J.M. O’Reilly, 1959-/. Mol. Spectrosc., 3:536.

Pitzer, K.S., 1953-Quantum Chemistry, Prentice-Hall, Englewood Cliffs, N.J.

- , and W.D. Gwinn, 1942-7. Chem Phys., 10:428.

Russell, H., D.R.V. Golding, and D.M. Yost, 1944-7. Amer. Chem. Soc., 66:16.
Unland, M.L., V. Weiss, and W.H. Flygare, 1965-7. Chem. Phys., 42:2138.

A PROTOSTEGID (SEA TURTLE) FROM THE TAYLOR FORM¬
ATION OF TEXAS

by L.W.OSTEN

Shuler Museum of Paleontology ,

Southern Methodist University , Dallas 75275

ABSTRACT

A right hypoplastron recovered from the Pecan Gap Chalk of the Taylor Formation in
Collin County, Texas is the fourth reported chelonioid in the Cretaceous of Texas. This
specimen is referred to the family Protostegidae but generic assignment is not made. The
other Texas specimens are from the Eagle Ford Formation (Zangerl, 1953), the Austin
Chalk (McNulty and Slaughter, 1964) and the Taylor Formation (Zangerl, 1953).

INTRODUCTION

In 1974, the author received a well preserved portion of a right hypoplastron
which was recovered from the Taylor Formation (Gulfian) of Collin County,
Texas. The locality is an outcrop 500 m east of Lake Lavon Dam where the
Pecan Gap Chalk is exposed. It is on deposit in the collections of the Shuler
Museum of Paleontology at Southern Methodist University (SMP-SMU 63472).

DESCRIPTION

The specimen consists of a major portion of the right hypoplastron and in¬
cludes a wedge of the right xiphiplastron (Fig. 1). The dorsal concavity is pre¬
served with a relief of 4.3 mm and is nearly symmetrical about a central point
on the hypoplastron. From this point, narrow elongate ridges radiate out to
the extremeties of the bony plate. These ridges can be traced on the ventral
surface as well. The insertion of the xiphiplastron into the hypoplastron pro¬
duces a ridge on the dorsal and ventral surfaces. This ridge originates along
the suture where a portion of the xiphiplastral element is exposed on the dorsal
surface. This insertion, as traced by the ridge, cannot be traced to the central
point but extends about half way (Table 1). This ridge is delineated in Figure 1
by narrow, elongate depressions.

Accepted for publication: November 3, 1975.

The Texas Journal of Science, Vol. XXIX, Nos. 3-4, December, 1977.

290

THE TEXAS JOURNAL OF SCIENCE

5 mm

Figure 1. Dorsal view of Protostegidae right hypoplastron with dorsal concavity ex¬
posed; arrow points anterior.

TABLE 1
Measurements

Anterior-Posterior Diameter

37.6

mm

Lateral Diameter

30.7

mm

Maximum Thickness

1.86

mm

Minimum Thickness

0.5 2

mm

Length of Xiphiplastoral Insertion Ridge

9.0

mm

The hypoplastron is not complete as preserved. The internal edge which
extends into the central cavity of the plastron is relatively thin, indicating a
termination and demonstrating that this is a natural edge (Fig. 2b). The external
edge is damaged and the greatest thickness is measured along this fracture.
The natural and fractured edges are indicated in Figure 2a.

COMPARISON

The properties of the Superfamily Chelonioidea are self evident. A com¬
parison of the right hypoplastron with a representative of each of the 3 families

A PROTOSTEGID

291

Figure 2. Dorsal view of upper Cretaceous Chelonioidea plastral plates. (Modified from
Zangerl, 1953) a. Protostegidae showing the alignment of our fossil specimen
(shaded); dotted line indicates natural edge. Chelosphargis advena. b. Outline
of Protostegidae plastron; right hypoplastron darkened, e. Cheloniidae right
hypoplastron. Corsoehelys haliniches. d. Toxochelyidae right hypoplastron.
Toxochelys latiremis.

suggests a protostegid affinity. This relationship was established using the over¬
all geometry and the character of the insertion of the xiphiplastron. The right
hypoplastron of the Protostegidae, Toxoehelyidae, and Cheloniidae are re¬
produced from Zangerl (1953) (Fig. 2). A photograph of our specimen was
aligned over these drawings using the xiphiplastral insertion as a guide.

The toxochelyid xiphiplastral insertion is developed in a proximal position
to the internal edge (Fig. 2d). The cheloniid xiphiplastron exhibits a relatively
wider insertion and the anterior-posterior length is not as well developed as in
our fossil specimen (Fig. 2c).

The fossil right hypoplastron is assigned to the Protostegidae of which three
genera are described (Zangerl, 1953). Generic assignment of this specimen must
wait for additional Gulfian protostegids.

292

THE TEXAS JOURNAL OF SCIENCE

ACKNOWLEDGEMENTS

Appreciation is due to Ronald Ritchie, collector of the fossil studied, and
to the United States National Park Service under whose program the fossil
was discovered and collected (PX 7000-3-0343).

LITERATURE CITED

McNulty, C.L., and B.H. Slaughter, 1964-A protostegid ramus from the upper Cretaceous
of Texas. Copeia, 2:454.

Zangerl, R., 1953-The vertebrates of the Selma Formation: Part III, The turtles of the
Family Protostegidae., Part IV, The turtles of the Family Toxochelyidae., Part V, An
advanced cheloniid sea turtle., Fieldiana: Geology Memoirs, 3(3,-5):59.

CHOICE OF MINICOMPUTERS’ FLOATING POINT BASE

by JAMES L. POIROT

Department of Computer Sciences,

North Texas State University, Denton 76203

ABSTRACT

The large increase in usage of the minicomputer for scientific applications has generated
a new flurry of discussion of the relative numerical merits of various choices of the base
for floating point arithmetic systems. Most of the papers on this subject compare usage of
binary, quarternary, and hexadecimal formats with the range of the exponent remaining
a common factor. This then leads to comparison of 32 bit (Fraction Bits X Exponent Bits)
floating point formats of (23 X 9), 24 X 8), and (25 X 7) for binary, quarternary, and hexa¬
decimal bases respectively. This paper compares the numerical precision attainable using
various bases with a common word structure for all bases. In addition, comparison of ac¬
curacies for common existing floating point systems are given.

INTRODUCTION

Over the past few years, the minicomputer has progressed from being a hard¬
ware device used primarily for monitor and control, to a widely accepted com¬
puter for general applications. This change has been primarily generated either
because of its low cost or because of small physical size requirements as is found
in many military applications. In any case, scientific programmers are often find¬
ing the minicomputer with its small word size their primary tool for problem
solving. Thus, discussion of the numerical accuracy of these computers has been
active. In particular, several recent papers, e.g. Brent (1973) and Cody (1973),
discuss the relative merits of various floating point formats and choices of base
for a 32 bit floating point word.

Several floating point bases are currently being used, including binary, octal,
and hexadecimal. In addition, a base 4, or quarternary number representation
scheme has been proposed. Comparisons of numerical accuracy as a function of
the base used have, for the most part, been conducted with the basic assumption
that the range of the exponent for all bases be equivalent. This leads to com¬
parisons then of formats where the number of bits set aside for fraction and ex¬
ponent vary depending upon the base being used. We believe that this type of
comparison, although theoretically pleasing and possibly applicable for future
design, does not aid the computer buyer in determining which existing com-

Received for publication: December 9, 1976.

The Texas Journal of Science, Vol. XXIX, Nos. 3 and 4, December, 1977.

294

THE TEXAS JOURNAL OF SCIENCE

puter would be numerically best for his application. The majority of existing
computer systems using 32 bit floating point formats, either allow 24 bits for
the fraction, including sign bit, and 8 bits for the exponent, (24 X 8), or 25
bits for fraction and 7 for exponent (25 X 7), independent of the base being
used. Moreover, hardware configurations, byte addressability and compatibility
with existing systems will most probably dictate the continued usage of such
formats. This paper is intended to give more insight on the attainable numerical
accuracy of floating point systems with the word format remaining constant.
Static properties of the (24 X 8) word format are compared with various choices
of base. Finally, a table comparing various word formats and bases, along with
examples of systems using such formats, are given.

Preliminaries

We will assume that we have a normalized sign-magnitude floating-point
representation scheme with 32 bits/word. Unless specified otherwise each word
contains a 24 bit fraction, f and an 8 bit exponent, e, with a base of 0. Moreover,
when using a base of 2 we will consider the case where the first bit is implicit,
Brent (1973). Thus, if the implicit first bit technique is used, we will let the
variable p = 2, otherwise p = 1 . We will then consider systems of the following
type (j3,p) = (2,1), (2,2), (4,1) and (16,1), where (3 is the base being used.

Static Characteristics

Cody (1973) has studied floating point systems with the range of exponents
remaining the same, comparing (23 X 9), (24 X 8) and (25 X 7) systems with
(/3,p) = (2,1), (4,1) and (16,1) respectively. In this section, we wish to make the
same type comparisons with a common (24 X 8) format for all bases. In our
study of numerical errors, we will let r represent an equivalence class of real
numbers and assume our error occurs in representing an element of r. This error
will be referred to as the representation error. Assuming the logarithmic prob¬
ability distribution for floating point numbers (Hamming, 1970), the average
relative representation error ARRE, is given by

(1) ARRE (ff) = if1--

4 - 2lln/3

where t = number of bits used for fraction, not including sign bit.

For the (24 X 8) word structure t = 23 if p = 1 and t = 24 if p = 2, that is if
the implicit bit system is used. The maximum relative representation error
(MRRE) is given by

(2) MRRE (ft) = 2_t _10.

Table 1 gives the values of ARRE and MRRE for the (24 X 8) word format
with (j3,p) equal to (2, 2), (2, 1), (4, 1) and (16, 1). As can be seen, and we might

MINICOMPUTERS

295

add as expected, ((3, p) = (2, 2) .gives the best results followed by the (2, 1),
(4, 1), and (16, 1) system.

TABLE 1

ARRE and MRRE for Fixed Format

Format

t

P

P

ARRE

MRRE

(24 X 8)

24

2

2

.180 X2"23

1 X 2"24

(24 X 8)

23

2

1

.361 X 2"23

2 X 2~24

(24 X 8)

23

4

1

.541 X 2~23

4 X 2“ 2 4

(24 X 8)

23

16

1

1.353 X 2"23

16 X 2“24

As a point of comparison, Table 2 shows values for ARRE and MRRE where
the range of the exponent remains constant.

Here t = 23 if /3 = 2 , p = 2
t = 22 if j3 = 2, p = 1
t = 23 if /3 = 4, p = 1
t = 24 if j3= 16, p = 1.

Notice that comparisons using Table 2 might lead to entirely different con¬
clusions than if using Table 1. From Table 2 the MRRE of the (2, 1) and (4, 1)
systems are the same and the ARRE of the (2, 1) system is greater than that for
the (4, 1) and (16,1) systems.

TABLE 2

ARRE and MRRE for Formats with Same Exponent Range

Format

t

0

p

ARRE

MRRE

(23 X 9)

23

2

2

.361 X 2~23

2 X 2“24

(23 X 9)

22

2

1

.722 X 2~23

4 X 2"24

(24 X 8)

23

4

1

.541 X 2“23

4 X 2~24

(25 X 7)

24

16

1

.676 X 2-23

8 X 2"24

Conclusions using Table 2 might be that there is little difference between the
(2, 1) and (16, 1) systems and a slight advantage in the ARRE of (4, 1) over
(2, 1). This conclusion obviously could not be reached using Table 1.

Another quantity used in comparison of floating point systems is the variance
of the relative error

e = E/f

where for our case E is the error in representation due to rounding the fraction

296

THE TEXAS JOURNAL OF SCIENCE

to t bits. E is assumed to be uniformly distributed in [-2-t -1, 2~t~1]. The
equation for the variance is given by [2]

0) a2(e)

10 -22t ln/3

it should be noted that the mean of e is 0. Also, if the particular system truncates
rather than rounds, the mean is not zero and equation (3) changes. However,
the comparisons of these variables give similar results.

Table 3 allows comparisons of a2(e) for the various (24 X 8) systems, while
Table 4 gives similar results maintaining equivalent sized exponents. As in the
cases of comparing the ARRE and MRRE, two different conclusions could be
obtained using the separate tables.

TABLE 3

Variance of Relative Error for f ixed Format

Format

t

&

P

o2(e)

(24 X 8)

24

2

2

.045 X 2~46

(24 X 8)

23

2

1

.180 X 2"46

(24 X 8)

23

4

1

.451 X 2"46

(24 X 8)

23

16

1

3.832 X 2~46

*

TABLE 4

Variance of Relative Error for f ormats with Same Exponent Range

Format

t

p

P

02(€)

(23 X 9)

23

2

2

.180 X 2“46

(23X9)

22

2

1

.721 X 2~46

(24 X 8)

23

4

1

.451 X 2~46

(25 X 7)

24

16

1

.958 X 2-46

Comparison of Common Formats

Because of the various choices of word formats and bases, comparison of
numerical accuracy of all systems cannot be simply performed. Choosing as
standard a common range for the exponent or common word format, leads to
results which for the computer buyer, may be somewhat misleading. For exist¬
ing systems, however, comparison of attainable accuracy can be simply accom¬
plished since the format and base have been set.

MINICOMPUTERS

297

Table 5 gives values for the ARRE, MRRE and a2(e) for 3 of the most com¬
mon floating point formats and bases. In addition, examples of computer sys-

TABLE 5

Comparisons of Characteristics of Various Existing Systems

Format

Base

ARRE

MRRE

a2(e)

Examples

(25 X 7)

16

.676 X 2"23

8 X 2-24

.958 X 2"46

NOVA, UNIVAC 1616

(24 X 8)

2

.361 X 2“23

2 X 2-24

.180 X2~46

Honeywell 316

(24 X 8)

2

(Implicit

Bit)

.180 X 2"23

1 X 2"24

0.45 X 2"4 6

PDP-11

terns using such floating point structures are listed. These examples are certainly
not exhaustive.

LITERATURE CITED

Brent, Richard P., 1973 -On the precision attainable with various floating-point number
systems. IEEE Transactions on Computers, C-22:601.

Cody, William J., 1973 -Static and dynamic numerical characteristics of floating-point
arithmetic. IEEE Transactions on Computers, C-22:598.

Hamming, R.W., 1970-On the distribution of Numbers. Bell System Tech. J., 49:1609.

How to Use the Nova Computers, 1970-Data General Corp., Southboro, Mass.

PDP 11/45 Processor Handbook, 1971 -Digital Equipment Corporation, Maynard, Mass.

Programmer’s Introduction to the IBM System 360, Assembler Language, 1969 -IBM.

Series 16 Software Documentation, 1970-Honeywell, Inc., Framington, Mass.

’

Notes Section

299

FIRST SOUTH TEXAS RECORDS OF PAPPOGEOMYS CASTANOPS- Although
the chestnut-faced pocket gopher has been reported in west Texas (Davis, 1974, Tex. Parks
and Wildlife Dept. Bull. 41:170; and Thornton and Creel, 1975, Southwestern Naturalist ,
20:272), no previous records have been noted from south Texas. Blair (1952, Tex. J. Sci.
4:230) failed to record this species. Three specimens of Pappogeomys castanops were col¬
lected during a mammal survey of the National Audubon Society Sabal Palm Grove Sanc¬
tuary at Southmost, Texas. These animals are in the collection of the Museum of Zoology
at Texas Wesleyan College.

Two adult females (TWCMZ-1449 and TWCMZ-1674) and 1 juvenile female (TWCMZ-
1833) were collected 6 mi (9.7 km) southeast of Brownsville, Cameron County, Texas. The
first adult catalogued was taken on 11 October 1974. The second adult and the juvenile
were collected on 8 January 1976.

The gophers were all taken in mixed short grass with deep sandy soil north of the Sabal
Palm forest. Numerous burrows were observed although none were noted during a similar
visit to the site in 1972. The nearest previously reported site was south of the Rio Grande
River in Tamaulipas, Mexico (Russell, 1968, Univ. Kans. Publ. Mus. Natur. Hist., 16:580).
Associated mammals taken from the site include Rattus rattus, Mus musculus, Reithro-
dontomys fulvescens, Cryptotis parva, Liomys irroratus, Peromyscus leucopus, Perognathus
hispidus, and Sigmodon hispidus. —Arthur G. Cleveland, Department of Biology, Texas
Wesleyan College, Fort Worth 761 05.

THE LEVEL OF UNDERSTANDING OF ALGEBRA AND TRIGONOMETRY
BY STUDENTS OF FRESHMAN PHYSICS— The high school graduate is required
to use his education in a variety of ways, and in many cases can minipulate circumstances
to avoid having to expose academic deficiencies. However, those students who attend col¬
lege in one of the preprofessional courses of study must encounter a survey course in physics in
a head on confrontation. This is generally considered to be a backbreaking experience, and
dropout and failure rates in these courses higher than 50% are accepted by college faculty
without concern or response.

A few years ago we became concerned that factors other than the topics covered in the
course were contributing substantially to the difficulty, and began to seek those parameters
which might fall into this category. As a result, a 26 question multiple choice test of 13
identifiable algebraic and trigonometric skills was developed. This paper reports the result
of that test when given to 843 students of freshman physics in the fall of 1976.

DISCUSSION

The topics tested are given in Table 1. Each topic had 2 questions, and those students
who correctly answer both questions on a given topic are said to “Understand” that topic.
Those who miss both questions are classified as “Do not understand”, and those who have
only one correct are called “Not sure”.

Table 1 also lists the % response of the students who finished in the top 20 percentile of
one section of 233 students. The other sections did not use computer grading so the correl¬
ation of final standing with the mathematics test is not readily available. We feel that the
sample of 233 students is large enough to have meaning.

The responses should be evaluated in terms of the test being multiple choice, 5 answer
choices/question. Guesses would fall on top of correct answers and skew figures up. To
counter that, the test was given on the first class day without prior warning and many stu¬
dents may have done better if some review had preceded the examination.

300

THE TEXAS JOURNAL OF SCIENCE

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Students Who Finished

Group As A Whole In Top 20 Percentile

NOTES

301

We also surveyed the mathematics background on the students in the section of 233. Of
that number, every student had had at least 1 semester of high school algebra and only 9
had less than 1 year. Fifty did not take high school trigonometry, but only 5 had not com¬
pleted trigonometry in college before taking this test. We feel that the ability to use the
mathematics expected to be a part of an algebra and trigonometry course was inadequate,
especially for topics 6, 8, 9, 10 and 11. Topic 11 stands out because even better students
did not fare well.

CONCLUSION

The University of Houston is a large, urban, commuter campus. The mean SAT of enter¬
ing freshman was 922 in 1975 (the last year for which figures are available). Many of these
students come from some of the most reputable school districts in the state of Texas. There¬
fore the sampling is considered to be typical. It is obvious that, for this group, the lack of
ready skills in algebra and trigonometry is contributing to their difficulties in the study of
physics.

The authors will be happy to send a copy of the test on request.-//. T. Hudson, and R.M.
Rottmann, Physics Department, University of Houston, 77004.

THE OCCURRENCE OF ARGULUS BICOLOR BERE, 1936 ON TRACHINOTUS
CAROLINUS (LINNAEUS) FROM THE TEXAS COAST-During July 1975 1 male
Argulus bicolor (Bere, 1936) was found on the first gill arch of a Florida pompano, Trach-
inotus carolinus (Linnaeus). The pompano came from a sample taken in the surf off South
Padre Island, Texas.

This is the first occurrence of A. bicolor from the Texas coast, the previous range being
from North Carolina to Louisiana (Cressey, 1972, Water Pollution Control Research Series,
18050 ELD-2171:14 pp.) T. carolinus is a new host for this species. Cressey (1972) re¬
ported the following genera as hosts: Strongylura, Morone, Gobionellus, Micropogon,
Scomboromous, Dorosoma, and Rhinoptera.

Figure 1. Argulus bicolor, a) Supporting rods sucking discs; b) Respiratory areas - vential
view; c) Basal plate of maxilla; d) Base of mouth tube with spines.

The principal character used to identify the animal was the respiratory areas (Fig. lb).
The supporting rods of the sucker wall (Fig. la), the teeth on the basal plate of the maxilla
(Fig. lc) and the spines on the base of the mouth tube (Fig. Id) were instrumental in the
final identification. The keys used for identification were Cressey (1972) and Wilson (1940,
Proc. U.S. Natl. Mus., 8&(30&1):459 .-Timothy L. Jones, Southwest Research Institute,
3600 Yoakum Blvd., Houston 77006, and Ronald C. Circe, Box 6010, No. 339, Texas A&I,
Corpus Christi 784 1 1 .

The Texas Journal of Science

Index to Volume XXIX

1977

Printed in San Angelo, Texas U.S.A.

By

The Talley Press

A

Acanthoceras
ma mil la re ; 212
Aeromonas; 85, 86, 91
formicans; 85, 86
hydrophila; 85, 86, 87, 88, 89, 90
formicans; 86, 87, 88, 90
liquefaciens; 85, 86, 87, 88, 89, 90, 91
proteolytica; 86, 87, 88, 90
punctata; 85, 86, 87, 88, 89, 90, 91
caviae; 85, 86, 87, 88, 89, 90, 91
salmonicida; 85, 86, 87, 88, 90
shigelloides; 85, 86, 87, 88, 90
Albian; 5
Alestes; 76
Ambystoma

tigrinum; 236, 238
Ammonites
alpinus; 265
Anapuzosia ; 269, 272
Anas

platyrhynchos; 141
Andropogan; 35
Anolis

carolinensis; 261
Aptian; 5
Argulus

bicolor; 301
Arizona

elegans; 240
Artemisia
fill folia; 34

Attrep, M., K.S. Tasa, J.D. Sherwood,
“Estimations of the Ratio of In¬
duced Fission to Spontaneous Fis¬
sion in Uranium Ores”; 109
Austiniceras
dibleyi; 264

B

Baglin, R.E., “Sex Ratios and Spawn¬
ings of White Bass, Morone Chry-
sops, from the Red and Washita
River Segments of Lake Tex-
oma”; 29

Bairdi; 79, 80, 83, 84
Baldwin, E.E., see Font, R.G.

Bat Cave Fault; 7, 8, 9, 10, 1 1, 12, 13

batagurine; 246

Bear Creek Fault; 12

Benjamin, C.P., see McGill, J.R.

beta-lysin; 59

Bison; 198

Bolen, E.G., see Womack, S.M.
Bouteloua; 35

Boyer, D.G., see Chang, M.
2-bromopropene; 279, 281, 282, 283
Brueckner, H.K., see Dasch, E.J.
Buchloe

dactyloides; 35
Bufo

cognatus; 237
speciosus; 238
valliceps; 238

c

Calotes

versicolor; 260
Camelops; 198
Ceuthophilus; 236

Chang, M., D.G. Boyer, “Fitting Soil
Temperature by aPeriodic Regres¬
sion Model”; 169

Cheatheam, L.K., “Density and Distri¬
bution of the Black-Tailed Prairie
Dog in Texas”; 33

INDEX TO VOLUME XXIX 1977

Cheloniidae; 291
Chinemys

reevesii; 245, 248

2-chloropropene; 279, 280, 281, 282,
283

Chrysemys
picta; 248

Circe, R.C., see Jones, T.L.
Clemmys
caspica

leprosa; 248
japonica; 245
marmorata
pallida; 248

Cleveland, A.G., “First South Texas
Records of Pappogeomys Cast-
anops”\ 299
Cnemidophorus
sexlineatus; 237
tigris; 240
Coluber
bairdi; 79

Comal Springs Fault; 9, 11, 12, 13
Cretaceous

chelonioidea ; 291

Crowder, G.A., R.W. Waltrip, “Ideal
Gas Thermodynamic Functions of
the 2-Halopropenes”; 279
Cruziana; 181
Cuara

amboinensis; 248
Cupressus; 199
Cyclemys

dentata; 245, 248
Cynomys

ludovicianus; 33

D

Dalquest, W.W., Equus tau Owen
from the Pleistocene of Mitchell
County, Texas”; 141
Dasch, E.J., H.K. Brueckner, J.M.
Rhodes, “Baking of Shale by a
Basaltic Dike: A Chemical and
Strontium Isotopic Study”; 15
Dawson, W.C., D.F. Reaser, J.D. Rich¬
ardson “Trace Fossils from the
Pecan Gap Formation (Upper Cre¬
taceous), Northeast Texas”; 175
Deiradoceras; 274

305

Deirochelys

reticularia

reticularia; 248
Dendrocygna; 141
arcuata; 141, 142
autumnalis; 141, 142
bicolor; 141, 142
eytoni; 141, 142
Desmoceras

cuvervillei; 269
Diplocraterion; 180
Dipodomys

heermanni; 240
spectablis; 237
Dorosoma; 301
Douvilleiceras; 272

E

Eagle Ford groups; 21, 25
Edwards Group; 5, 7, 9, 1 1
Elaphe

bairdi; 79, 80, 81, 82, 83, 84
obsoleta; 79, 80, 81, 82, 83, 84
Emys

orbicularis; 246

Engonoceras; 263, 264, 267, 272
stolleyi; 272

Eogaudryceras; 263, 264, 267, 268
269,271,272,273
Equus; 198
littoralis; 141
tau; 141
Escherichia

coli; 60, 61, 62, 63, 64, 65, 66
Esox

lucius; 31

F

Fitzpatrick, L.C., see Jones, F.V.
2-fluoropropene; 279, 280, 281, 283
Font, R.G., “Influence of Anisotro¬
pies on the Shear Strength and
Field Behavior of Heavily Over-
consolidated, Plastic and Expansive
Clay-Shales”; 21

Font, R.G., J.C. Yelderman, C.T.
Hayward, E.E. Baldwin, “Prelim¬
inary Field Study of the Fracture
Patterns Associated with the Bal-
cones Fault Zone in North Cen¬
tral Texas”; 187

306

THE TEXAS JOURNAL OF SCIENCE

G

Gaudryceras; 268
Gempylidae; 7 1
Geomys; 238

bursarius; 236, 237, 238, 239,
240, 241.

Glen Rose Formation; 5, 7, 9, 12

Gobionellus; 301

Graptemys

barbouri; 245, 246, 249
caglei; 249
flavimaculata; 249
geographica; 249
kohni; 249
nigrinoda; 249
oculifera; 249
pseudogeograph ica
sabinensis; 249
pulchra; 245

Griffin, W.L., see McNeal, J.U.
Gyrolithes ; 175, 180

H

Hao, H., see Schufle, J.A.

Harcombe, P.A., J.E. Neaville, “Veg¬
etation Types of Chambers County,
Texas”; 209

Hatoff, B.W., see Ritter, E.W.
Hayward, C.T., see Font, R.G.
hematoxylin; 142
Henson, T.K., see Perry, R.B.
Heterodon
nasicus; 237
platyrhinus; 240

Hickman, G.C., “Geomyid Inter¬
action in Burrow Systems”; 235
Holbrookia

propinqua; 255, 260
Hudson, H.T., R.M. Rottmann, “The
Level of Understanding of Algebra
and Trigonometry by Students of
Freshman Physics”; 299
Hydrocynus; 76

Hypophylloceras; 263, 265, 267, 268
ellipticum; 263, 265, 267, 268
escragnollense; 268
onoense; 265
subalpinum; 265

subalpinus ; 268
velledae; 265, 267
hypoplastron; 289, 290, 291

I

Ictalurtus
melas; 42
punctatus; 42
immunoglobulins; 59
2-iodopropene; 279, 282, 283

J

Jones, F.V., W.D. Pearson, L.C.
Fitzpatrick, “Yield Estimates De¬
rived from Active and Passive
Creel Surveys of a Small Pond
Fishery”; 41

Jones, T.L., R.C. Circe, “The Occur¬
rence of Argulus bicolor Bere,
1936 on Trachinotus carolinus (Lin¬
naeus) from the Texas Coast”; 301
Juniperus; 34, 199

K

Kachuga
smithi; 250
tecta; 245

Kainer Formations; 11, 12
Killebrew, F.C., “Mitotic Chromo¬
somes of Turtles. IV. The Emy-
didae”; 245

Kolb, J.W., B.G. Whiteside, “Age and
Growth of Largemouth Bass in
Canyon Reservoir, Texas”; 49
Kshirsagar, A.M., see Sia, L.L.

L

Lepidotes; 76
Lepisosteus; 76
Lepomis

cyanellus; 42
macrochirus; 42
megalot is; 42
Libocedrus; 199
lysozyme; 59

INDEX TO VOLUME XXIX 1977

M

Malay emys

subtrijuga; 25 1
Mamma thus; 198

Manuaniceras ; 263, 264, 271, 273
carbonarium; 263
supani; 263, 264, 271, 273

Marshall, J.L., S.R. Walter, “Mass
Spectrometry Studies of Norcam-
phor and Norbornyl Acetate”; 121
Mauremys
mutica; 250
nigricans; 245

McGill, J.R., C.P. Benjamin, “The
Effects of Amniotic Fluid on the
Growth of Certain Gram-Nega¬
tive Bacteria”; 59

McNeal, J.U., W.L. Griffin, “Dog
Flesh as a Potential Food Resource
for Carnivores: An Exploratory
Study”; 101
Micropogon; 301
Micropterus

salmoides; 42, 49

Morgan, E.C., “Dentitional Phenomena
and Tooth Replacement in the
Scabbard Fish Trichiurus lep turns
Linnaeus (Pices: Trichiuridae)” 71
Morone; 301
chrysops; 29

Mortoniceras; 263, 265, 267, 268,
273, 274

nanum; 263, 267, 273
scobinum; 263, 267, 274
Mustela

frenata ; 240
nigripes; 37

N

Neaville, J.E., see Harcombe, P.A.
Notemigonus

chrysoluecas; 42
Nymphaea; 199, 200, 201

o

obsoleta; 79, 80, 83, 84
Ocadia

sinensis; 246, 250

307

Odocoileus; 198

Olson, R.E., “Evidence for the Species
Status of Baird’s Ratsnake”; 79
Onychomys

leucogaster; 237, 240
Opuntia

imbricata; 35
tuna; 34

Osten, L.W., “A Protostegid (Sea
Turtle) from the Taylor Forma¬
tion of Texas”; 289

P

Pappogeomys; 238

castanops; 236, 237, 238, 239,
241, 299

Pearson, W.D., see Jones, F.V.
Perognathus; 240
flavus; 237
Peromyscus
hispidus; 240
inornatus; 240
maniculatus; 240

Perry, R.B., T.K. Henson, “Chroma¬
tographic Studies of Interaction
Coefficients of Benzophenone and
Benzhydrol with Various Com¬
pounds; 131
Person Formations; 11
Pervinquieria

inflata; 272, 274
Phylloceras

ellipticum; 268
Pinus; 200, 203, 204
Pituophis

melanoleucus; 237, 240
Platanus

racemosa; 198

Poirot, J.L. “Choice of Minicomputers’
Floating Point Base”; 293
Pomoxis

annularis; 42
Populus; 198, 199
pre-Cretaceous; 189
progesterone; 59
Prosopis

juliflora; 34
Proteus

vulgaris; 60, 61, 62, 63, 64, 67
Protostegidae; 289, 290 291

308

Pseudemys

concinna

texana; 250
floridana
hogi; 250
nelsoni; 250
script a

callirostris; 250
elegans; 250

Pseudobilobites; 175, 180, 183

Pseudomonas

fluorescens; 60, 61, 62, 63, 64

Pseudotsuga

menziesii; 198, 199

Puzosia; 26 3, 264, 267 , 269, 271, 272
cuvervillei; 264, 272
planulata; 269, 272
(Puzosia)

mayoriana; 272

w el wits chi; 263, 264, 267, 269,
271, 272

Q

Quercus ; 200, 203
havardii; 34

R

Rao, V.B., B.G. Foster, “Antigenic
Analysis of the Genus Aero-
monas 85

Reaser, D.F., see Dawson, W.C.

Rhinoclemys; 246
funeria; 251
punctularia

melanosterna; 251

Rhinoptera; 301

Rhizocorallium; 175, 179, 180, 183

Rhodes, J.M., see Dasch, EJ„

Richardson, J.D., see Dawson, W.C.

Ritter, E.W., B.W. Hatoff, “Late Ple¬
istocene Pollen- and Sediments:
An Analysis of aCentral California
Locality”; 195

Rottman, R.M., see Hudson, H.T.

Rowland, L.O., see Stickney. R.R.

Rylander, M.K., see Womack, S.M.

s

THE TEXAS JOURNAL OF SCIENCE

Scalopus

townsendii; 240
Scaphiodontophis; 76
Scaphiopus

couchi; 237, 238
holbrooki ; 238
Schloenbachia
in flat a

Sow; 272

Schufle, J.A., H„ Hao, “The Tempera¬
ture of Maximum Density of Deu¬
terium Oxide in Capillaries”; 137
Scomboromous; 301
Sequoiodendron; 199'

Serratia

marcescens; 60, 61, 62, 63, 64
Sherwood, J.D., see Attrep, M.
Sia, L.L., A.M. Kshirsagar, “Loss of
Efficiency due to Estimated
Weights in the Recovery of Inter-
Block Information”; 147
Siebenrockiella

crassicollis; 246, 251
Simmons,- H.B., see Stickney R.R.
Speotyto

cunicutaria; 33, 37
Spermophilus
townsendii; 240
tridecemlineatus; 240
Stickney, R.R., H.B. Simmons, L.O.
Rowland, “Growth Responses of
Tilapia mrea to Feed Supplemented
with Dried Poultry Waste”; 93
Stipsellus; 175, 179, 180, 1-83-
Strongylura; 301
Superfamily Chelonioidea; 290
Sylvilagus; 198

T

Tasa, K.S., see Attrep, M.

Taxus; 199
Taylor groups; 21, 25
Terra pene

Carolina; 25 1
coahuUa; IS 1
ornata ; 251
Tetragonites; 268

timotheamis; 268, 269
Thalassinoides; 175, 179, 180, 183
Thomomys ; 238

home; 236, 237, 238, 239, 240, 241

Salix; 198, 199

INDEX TO VOLUME XXIX 1977

309

talpoides; 240
Tilapia; 93

aurea; 93, 94,95,96, 97,98
Toxoehelyidae; 291
Trachinotus
carolinus ; 301
transferrin; 59
Trichiuridae; 71
Trichiurus

lepturus; 71, 72, 73, 74, 75, 76

V

Vieja Group; 16

w

Waco Springs Eault; 7, 8, 10, 11, 12, 13

Walnut Formation; 5, 12

Walter, S.R., see Marshall, J.L.

Waltrip, R.W., see Crowder, G.A.

Washita groups; 2 1 , 24

Watson, J.T., “Effects of Hypophys-
ectomy in the Lizard Holbrookia
propinqua"', 255

Whiteside, B.G., see Kolb, J.W.

Womack, S.M., M.K. Rylander, E.G.
Bolen, “The Structure of the
Retina in Four Species of Whistling
Ducks , Dendrocygiw, 141

Woodbine groups; 21, 25

X

Xiphiplastron; 289, 291

Y

Yelderman, J.C., see Font, R.G.

Young, K., “Cretaceous (Aibian) Am¬
monites from Puerto Rico and
St. Thomas”; 263

Yucca

galuca; 34

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EXECUTIVE COUNCIL

President:

President-Elect :

Vice President:

Immediate Past President:

Secretary- Treasurer:

Sectional Chairpersons:

I -Mathematical Sciences: WILLIAM D. CLARK, Stephen F. Austin State University

II -Physical and Space Sciences: ROBERT W. GRUEBEL, Stephen F. Austin State Univ.

III -Earth Sciences: JAMES B. STEVENS, Lamar University

IV -Biological Sciences: RICHARD H. RICHARDSON, University of Texas at Austin

V -Social Sciences: RAYMOND TESKE, JR., Sam Houston State University

VI -Environmental Sciences: ELRAY S. NIXON, Stephen F. Austin State University

VII -Chemistry Section: BILLY J. YAGER, Southwest Texas State University

VIII -Science Education: JOEL E. BASS, Sam Houston State University

IX -Computer Sciences: GRADY G. EARLY, Southwest Texas State University

X -Aquatic Sciences: WILLIAM J. CLARK, Texas A&M University

XI -Forensic Sciences: RAYMOND WM. MIRES, Texas Tech University

Manuscript Editor: G. ROLAND VELA, North Texas State University
Managing Editor: MICHAEL J. CARLO, Angelo State University

Chairperson, Board of Science Education: PAUL J. COWAN, North Texas State University
Collegiate Academy: ROBERT V. BLYSTONE, Trinity University
Junior Academy: FANNIE M. HURST, Baylor University

BOARD OF DIRECTORS

JAMES R. UNDERWOOD, JR., West Texas State University
J. D. McCULLOUGH, Stephen F. Austin State University
HERBERT H. HANNAN, Southwest Texas State University
EVERETT D. WILSON, Sam Houston State University
G. ROLAND VELA, North Texas State University
MICHAEL J. CARLO, Angelo State University
ARTHUR E. HUGHES, Sam Houston State University
WILLIAM K. DAVIS, Southwest Texas State University
ROBERT BOYER, University Station
JAMES D. LONG, Sam Houston State University
JOHN W. FITCH, Southwest Texas State University
LAMAR JOHANSON, Tarleton State University
FREDERICK GEHLBACH, Baylor University

JAMES R. UNDERWOOD, JR., West Texas State University
J. D. McCULLOUGH, Stephen F. Austin State University
J. L. POIROT, North Texas State University
HERBERT H. HANNAN, Southwest Texas State University
EVERETT D. WILSON, Sam Houston State University

COVER PHOTO

Location map. The study area is centered in the Waco urban region and
encompasses McLennan County, Texas and its immediate vicinity.

by R.G. Font, pp. 187-194

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