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Transactions

JAR 1 6 138!

of the

Illinois

State Academy
of Science

Volume 55
No. I
1962

Springfield, Illinois

TRANSACTIONS of the ILLINOIS STATE ACADEMY of SCIENCE

Editorial Board:

Wesley J. Birge, University of Illinois, Editor and Chairman

Robert S. Bader, University of Illinois

Russell S. Drago, University of Illinois

Francis Kruidenier, University of Illinois

John McGregor, University of Illinois

Wayne J. McIlrath, University of Chicago

Howard C. Roberts, University of Illinois

Theodore Schmudde, Southern Illinois University

Timothy Whitten, Northwestern University

The current Transactions may be obtained by payment of annual dues.

Previous volumes may be obtained by addressing Willard D. Klimstra,
Southern Illinois University, Carbondale.

Exchanges may be arranged by addressing Thorne Deuel,

Illinois State Museum, Springfield.

(49177 — 1-62)

TRANSACTIONS

OF THE

ILLINOIS STATE
ACADEMY OF SCIENCE

VOLUME 55 - 1962

No. 1

Illinois State Academy of Science

AFFILIATED WITH THE

Illinois State Museum Division
Springfield, Illinois

PRINTED BY AUTHORITY OF THE STATE OF ILLINOIS

Otto Keener, Governor

December 17, 1962

CONTENTS

Skrjabingylus chitwoodorum (Nematoda : Metastrongylidae) in Skunks
in Illinois. By N. D. Levine, V. Ivens, T. R. B. Barr, and B. J.
Verts . 3

Additional Faunal Records from the Kingston Lake Site, Illinois. By

Paul W. P arm alee . 6

Response of Bobwliite Quail to Management on some Illinois Strip-

mined Lands. By Paul Vohs, Jr. and Dale E. Birkenholz . 13

On Multiply Mutant Sets. By A. A. Multan . 20

Comparative Methods of Trapping Small Mammals in an Illinois Woods.

By Raymond L. Will . 21

Absence of Rabies in Some Bats and Shrews from Southern Illinois.

By E. W. Pearson and T. R. B. Barr . 35

Root Growth of Transplanted Lobloll}" Pine ( Pinus taeda L.) Seedlings

in Relation to Chemical Root Reserves. By A. R. Gilmore . 38

The Mirabilis — Insect Community in Illinois. By W. V. Balduf . 42

The Effects of Red and Far-red Irradiation on the Vegetable Develop¬
ment of Pea and Cocklebur. By Robert H. Kupelian . 48

Some Comparative Aspects of Organ Weights in Canada Geese

( Brant a canadensis interior). By Harold C. Hanson . 58

Junior Academy Publications. By John C. Frye . 70

The Hatching Muscle in the American Coot. By James R. Fisher . 71

Food Habits of the Leopard Frog ( Rana pipiens sphenocephala) in a

Minnow Hatchery. By Robert Jeffery Lewis . 78

Stomach Contents of Bullfrogs ( Rana catesbeiana) Taken from a

Minnow Hatchery. By William M. Lewis, Jr . 80

SKBJ ABINGYLUS CHITWOODOBUM (NEMATODA:
MET ASTEON GYLIDAE ) IN SKUNKS IN ILLINOIS

NORMAN D. LEVINE, VIRGINIA IVENS, THOMAS R. B. BARR

and B. J. VERTSi

University of Illinois , Urbana and Illinois Natural History Suy'vey, Urbana

Skrjabingylus chitwoodorum lias
been found in the frontal sinuses of
skunks in several localities in the
United States, but it has not hereto¬
fore been reported from Illinois. The
purpose of the present paper is to
record its occurrence in this state
and to summarize earlier reports on
its occurrence, life cycle and patho¬
genesis.

In connection with a study of
sylvan rabies, the brains of 184
striped skunks ( Mephitis mephitis )
from northern Illinois were exam¬
ined. Skrjabingylus chitwoodorum
was found in the sulci and fissures
beneath the dura mater in two of
these animals. Two male and two
female nematodes were found in an
adult male skunk trapped March 19,
1958 in Carroll County, Illinois, and
two males and two females were
found in another adult male skunk
trapped June 24, 1958 in Cook
County.

In addition to the above, a single
male S. chitwoodorum was found
February 23, 1959 on the brain of
a road-killed male skunk near Pom¬
ona, Jackson County, and a single
female nematode was found on the
brain of a female skunk caught in
a trap March 11, 1959 by a fox trap-

1 This study was supported in part by
National Institutes of Health Grant E-
1349.

per near Alto Pass, Union County.
Both these counties are in southern
Illinois.

All the worms were not intact,
so that it was not possible to make
complete measurements. Three fe¬
males were 27 to 42 mm long* and
842 to 1070 microns wide. Five
males were 16 to 24 mm long and
502 to 816 microns wide. The
spicules of four males were 800, 816,
846 and 875 microns long, respec¬
tively, and the gubernacula of two
males were 90 and 97 microns long.
These may be compared with spic¬
ule lengths of 540 to 710 (average
631) microns reported by Hill
(1939) and of 715, 795 and 830 mi¬
crons reported by Goble (1942), and
to gubernaculum lengths of 72 to 88
(average 80) microns reported by
Hill (1939).

Skr jabingylus chitwoodorum Hill,
1939 was originally described from
two striped skunks, Mephitis mephi¬
tis mesomelas (syn., M. mesomelas
mesomelas) , and three eastern
spotted skunks, Spilogale put or i us
interrupta (syn., S. interrupta ), in
Oklahoma by Hill, (1939). (The
names of the hosts in our paper are
those used b}^ Hall and Kelson,
1959).

S . chitwoodorum has also been
found in Mephitis mephitis occi-
dentalis (syn., M. occidentalis) in
California by Hobmaier (1941), in

[3]

Transactions Illinois Academy of Science

18 of 25 M. m. nigra (syn., M. nigra)
in New York by Goble (1942) and
Goble and Cook (1942), in a dead
skunk (presumably M. mephitis) in
Pennsylvania by Bell and Chalgren
(1943), and in M. m. nigra, pre¬
sumably from the vicinity of Belts-
ville, Maryland by Dikmans and
Goldberg (1949). In addition, Stege-
man (1939) found bone lesions sug¬
gestive of S. chitwoodorum infection
in the skulls of 26 out of more than
150 M. m. nigra from New York, and
Tiner (1946) found similar lesions
in the skulls of 3 M. m. varians out
of 400 skulls of this species and
Spilogale g. gracilis (syn., S. leu-
coparia) in Texas. Finally, Miller
(1899) found nematodes which were
identified as Filaroides mustelarum
but which were undoubtedly S. chit¬
woodorum in the frontal region of
the skulls of skunks ( M . m. mephi¬
tis) at North Bay, Ontario, and
stated that all the adult skunks which
he took in this area were infected.

The only record of S. chitwoodo¬
rum in Spilogale putorius is that of
Hill (1939) in Oklahoma.

S. chitwoodorum has been found
in the western spotted skunk, S.
gracilis, in California by Grinnell,
Dixon and Linsdale (1937), in S.
gracilis phenax in California by
Hobmaier (1941), and in S. gracilis
latifrons in British Columbia by
Cowan (1941). In addition, Tiner
(1946) found bone lesions sugges¬
tive of S. chitwoodorum infection in
the skulls of 3 S. g. gracilis (syn.,
S. leucoparia) out of 400 skulls of
this species and M. m. varians in
Texas.

Hobmaier (1941) found that third
stage larvae developed in slugs and
snails. Development was best in

Limax maxima s, L. cinereus, L.
flaws, L. niger, Agriolimax agrestis
and Milax sp., while Epiphragmo-
pliora sp. and Helix pomatia were
seemingly less favorable hosts. Hob¬
maier apparently infected skunks
with larvae from slugs or snails. In
addition, he found a natural infec¬
tion with third stage larvae in a
Pacific garter snake, Thamnopliis
sirtalis infernalis, and he infected
frogs artificially. He was unable to
obtain adult nematodes in artificially
exposed guinea pigs, dogs, cats or
ferrets, but found some larvae in
the stomach wall and mesenteries of
mice and rats. He considered that
the normal route of migration of the
larvae in skunks was directly from
the mouth thru the nasal passages to
the sinuses.

S. chitwoodorum ordinarily occurs
in the frontal sinuses and may de¬
form the skull. Bell and Chalgren
(1943) stated that the skunk which
they examined “had bulbous en-
largments of the frontal sinuses con¬
taining” the nematode. Miller
(1899) reported that the parasites
he found disfigured the frontal re¬
gion of the skulls of a large propor¬
tion of the specimens. Stegeman
(1939) found that 26 of more than
150 skulls of skunks from New York
had a porous, relatively soft, fre¬
quently discolored dilation on the
dorsal surface at the junction of the
frontal and parietal bones ; the cavi¬
ties in these swellings were contin¬
uous with the frontal sinuses ; he
found no worms, since he was study¬
ing skulls alone. The lesions which
Tiner (1946) found in the sknlls of
S. gracilis and M. mephitis and
which he assumed were caused bv
S. chitwoodorum varied from marked

Skrjabingylus Chitwoodorum in Skunks

bulging and osteitis over the frontal
sinuses to actual holes in the walls
of one or both of them.

Lesions suggestive of S. chitwood¬
orum infection were seen in the
frontal region of the skull of a fifth
skunk collected in northern Illinois ;
the frontal sinuses of this animal
were not examined, and no worms
were seen. However, no lesions were
seen in the skulls or brains of the
Illinois skunks in which S. chit¬
woodorum was found. Our report
is apparently the first record of this
nematode’s occurrence in the brain
cavity.

Summary

Skrjabingylus chitwoodorum was
found in the sulci and fissures of
the brain beneath the dura mater
of 2 out of 184 striped skunks {Me¬
phitis mephitis ) in northern Illinois
and in 2 other M. mephitis in south¬
ern Illinois. This is apparently the
first record of this nematode’s oc¬
currence in the brain cavity. The
hosts were apparently healthy, and
their skulls were not noticeably de¬
formed. Earlier reports on S. chit¬
woodorum are summarized.

Literature Cited

Bell, J. F., and W. S. Ciialgren. 1943.

Some wildlife diseases in the eastern

United States. J. Wildlife Manag.,
7: 270-278.

Cowan, I. McT. 1941. Report upon
some diseases and parasites of game
birds and game and fur-bearing mam¬
mals in British Columbia. Rep. Prov.
Game Comm. Brit. Col., 40-45.

Dikmans, G., and A. Goldberg. 1949.
A note on Arthrocephalus lotoris
(Schwartz, 1925) Chandler, 1942 and
other roundworm parasites of the
skunk, Mephitis nigra. Proc. Helm.
Soc. Wash., 16: 9-11.

Goble, F. C. 1942. Skrjabingylus chit¬
woodorum from the frontal sinuses of
Mephitis nigra in New York. J. Mam¬
mal., 23: 96-97.

Goble, F. C., and A. H. Cook. 1942.
Notes on nematodes from the lungs
and fronted sinuses of New York fur
bearers. J. Parasitol., 28: 451-455.

Grinnell, J., J. S. Dixon and J. M.
Hinsdale. 1937. Fur-bearing mam¬
mals of California. 310 pp.

Hall, E. R., and K. R. Kelson. 1959.
The mammals of North America. Vol.
II. New York, Ronald Press Co.. 1083

pp.

Hill, W. C. 1939. The nematode
Skrjabingylus chitwoodorum n. sp.
from the skunk. J. Parasitol., 25: 475-
478.

Hobmaier. M. 1941. Extramammalian
phase of Skrjabingylus chitwoodorum
(Nematode. J. Parasitol., 27: 237-

239.

Miller G. S., Jr. 1899. Notes on the
mammals of Ontario. Proc. Boston
Soc. Nat. Hist., 28: 1-44.

Stegeman, L. C. 1939. Some parasites
and pathological conditions of the
skunk ( Mephitis mephitis nigra) in
central New York State. J. Mammal.,
20: 493-496.

Tiner, J. D. 1946. Some helminth para¬
sites of skunks in Texas. J. Mammal.,
27:82-83.

Manuscript received June SO, 1001 .

ADDITIONAL FAUNAL RECORDS FROM THE
KINGSTON LAKE SITE, ILLINOIS

PAUL W. PARMALEE
Illinois State Museum, Spring-field

Recent acquisitions by the Illinois
State Museum of private archaeol¬
ogical collections from the Kingston
Lake (Village; Kitchen Midclen)
Site has provided quantities of bone
and shell of noteworthy interest. Re¬
covery of this material was made
periodically from 1932 to about 1943,
and through the kindness of Mr.
L. P. Elliott, Dr. V. H. Chase and
the late Dr. Donald E. Wray, all of
Peoria, animal remains found by
them at this site were made avail¬
able for study. A small sample of
bone collected by the late George
and Ethel Schoenbeck during this
time period has also been included
in this report.

This large Middle Mississippi site
(primarily Spoon River Focus:
1,100-1,400 A.D.), once covering ap¬
proximately 15 acres, was situated
on the north bank of Kingston Lake
(parallel and adjacent to the Illi¬
nois River) 15 miles southwest of
Peoria, Peoria County, Illinois. The
village and the one large platform
mound were the property of the
Kingston Lake Gravel Company, and
with the commencing of dredging
operations in 1931, the site was grad¬
ually destroyed until, by 1938, most
of it was gone. A general account
of the artifacts, burials and other
materials recovered during these
years has been presented by Simp¬
son (1939). Included in Simpson’s

report is an article by F. C. Baker
on a sample of the faunal materials
found at the beginning of the ex¬
cavations ; this same article appeared
in the Transactions of the Illinois
Academy of Science (Baker, 1936).
A few of the first specimens re¬
covered from this site (“Simpson
Village”) were reported in an ear¬
lier paper by Baker (1931).

The variety of both molluscan and
vertebrate species represented pro¬
vides an excellent index of the food
habits of these people as well as to
the prehistoric distribution and pos¬
sible abundance of the local fauna.
As suggested by the remains of cer¬
tain species, both the marshy flood-
plain areas to the south and upland
areas to the north were hunted for
game. As evidenced by the variety
of fresli-water mussels, fish, turtles,
and aquatic and semi-aquatic species
of birds and mammals, however, the
Illinois River and its associated back¬
waters and bottomlands constituted
the major areas hunted. The re¬
cently acquired faunal materials con¬
sidered in this report add to the
number of species previously record¬
ed, as well as to the quantity, thus
presenting a more complete study of
the early fauna and its use by the
Indians occupying this site. Table
1 is a list of the species identified
from the Kingston Lake Site, in¬
cluding those recorded by Baker

[6]

Faunal Records from Illinois

(1931, 1936) and those recently ob¬
tained from Elliot, Chase and Wray.

Accounts of Species

Mollusks. — Compared with the
quantity and variety of marine
shells recovered at the Cahokia Site
(Parmalee, 1958a), located along the
Mississippi River (Madison Co., Illi¬
nois) and probably contemporaneous
with this site, there was little evi¬
dence found to suggest a like use of
such marine forms by the Kingston
Lake people. Apparently trade
routes or contacts with the south¬
eastern coastal areas were meager,
and the only reference to the use of
marine shells states that ‘ ‘ . . . several
tools made from the columnella of
conch shells . . . were collected”
(Simpson, 1939).

Fresh-water mussels, however,
were used extensively for food and,
to a lesser degree, for ornaments
and/ or implements. Twenty-four
species were identified and nearly all
were typical large-river forms. Simp¬
son {op. cit.) mentions the finding
of two “clam bakes” in which mus¬
sels had been placed in the ground
for baking but never removed. Mus¬
sels were undoubtedly collected lo-
cally in the Illinois River beds ; with
the possible exceptions of P. lineo-
lata, 0. olivaria and E. crassidens,
all species represented apparently
still inhabit that section of the river.
Alasmidonta marginata is a head¬
water or small-stream form and nor¬
mally does not occur in a large riv¬
er environment, and its presence with
all large-river species is unusual.

Valves of mussels were used in
the manufacture of beads, spoons,
and hoes. Simpson {op. cit.) re¬

cords two spoons (probably L. ven-
tricosal ), 10 shell ornaments, 695
disc beads (possibly from marine
whelks rather than mussels?) and
170 shell hoes, and states that “With
two exceptions, all [hoes] were made
from one species of clam.” This
would probably be the large, thick-
shelled washboard, M. gigantea; 15
shell hoes in the Elliott collection
were referable to this species.

Vertebrates. — Twenty species of
mammals were identified from the
Kingston Lake Site, bones of the
white-tailed deer forming 65 per¬
cent of the total. Although the per¬
centage of deer bone to that of other
species was not so large as it is at
most sites {e.g. Cahokia : Parmalee,
1957), the quantity of remains at¬
test to the use of this animal as a
basic meat staple. Numerous deer
bone artifacts were recovered, in¬
cluding cut antler, antler tines and
projectile points, awls (from ulnae
and splinters), jaw bone “hoes” and
beamers (from metatarsals).

Several species of the smaller
mammals (raccoon, beaver, muskrat,
cottontail, squirrel) were also taken
in considerable numbers, their re¬
mains comprising 18 percent of the
mammalian bone. Remains of the
mole and small rodents {Peromyscus,
Citellus, Oryzomys) are probably
from animals that died naturally at
the site location and were incidental
to human occupation. However, as
Baker (1936) has indicated, the
former occurrence of the marsh rice
rat in Peoria County is of special
significance since it is now restricted
to the southern most part of the
state. This rodent has since been
identified from several other sites in
the Illinois-Mississippi river valley

Transactions Illinois Academy of Science

(Parmalee, 1957) and its presence
at these sites is indicative of dense
vegetation along marsh and swamp
margins.

Elk still inhabited the Illinois re¬
gion in early historic times and bones
have been recovered at numerous
aboriginal village sites, but only rare¬
ly in any quantity. Apparently elk
were far less numerous than the
white-tailed deer in prehistoric times.
One of the more interesting aspects
of the mammalian complex from
this site is the occurrence of bison.
Griffin and Wray (1945) have sug¬
gested that this animal did not
cross the Mississippi River into east-
tern United States until 1600 or
shortly before. This appears correct
as bison remains in prehistoric Illi¬
nois sites are typically rare or non¬
existent ; only in the historic com¬
ponents of the Starved Rock and
Zimmerman sites in La Salle county
have bison bones been encountered
in quantity (over 150 specimens in
the Illinois State Museum archae-
ozoological collections). The Sclioen-
becks reportedly obtained a scapula
hoe “thought” to be bison from this
site and, although the number of
bones recovered are few, they may
suggest a beginning of the eastward
bison migration in the late 15th or
early 16th century.

Birds were apparently an im¬
portant source of food to these In¬
dians, and the Illinois River, King¬
ston Lake and associated backwaters
formed an excellent habitat for a
large variety of species. At least 36
species were represented, and bones
of waterfowl (ducks, geese, swans)
comprised 54 per cent of the total
identified birds. Other aquatic and
semi -aquatic species (sora, cormor¬

ant, coot, sliorebirds, etc.) formed an
additional 12 percent. Only two
species, the bobwhite and the prairie
chicken, suggest an upland prairie
habitat.

The trumpeter swan is now ex¬
tirpated in Illinois. In prehistoric
times, however, it was a common mi¬
grant along the Mississippi River
and apparently, but to a lesser ex¬
tent, elsewhere in the state, so re¬
mains of this species at the King¬
ston Lake Site are not unexpected.
Although far less numerous than at
Cahokia (Parmalee, 1957), bones of
0. buccinator at this site establishes
it as a former migrant through cen¬
tral Illinois. A single, cut distal
humerus end was recovered at King¬
ston Lake ; these cut ends were com¬
mon at the Cahokia Site (Parmalee,
op. cit.), the bone shaft having ap¬
parently been used for tools or sec¬
tioned into beads. The majority of
the other avain species identified
from this sample still occur locally
or as migrants through the area.
However, the wild turkey is now
extirpated in Illinois and the prairie
chicken is no longer found in that
area. Both the whistling swan and
sandhill crane occur rarely during
migration, and the long-billed cur¬
lew (now accidental in Illinois) has
been identified from only two other
sites (Parmalee, 1958b; plus an his¬
toric Crawford Farm Site, Rock Is¬
land Co. record).

Turtles were utilized to a limited
extent by the Indians who occupied
this site, probably mainly for food,
although sections of worked cara¬
pace (scraped interior) were re¬
covered which indicate that shells
were occasionally fashioned into
bowls or dishes. A minimum of six

Faunal Records from Illinois

Table 1. — Enumeration of the Animal Species Identified from the Kingston Lake
Site, Peoria Co., Illinois.

Species

Number of Specimens

Elliott, Chase,
Wray,

Fresh-water mussels Schoenbeck

Amblema peruviana (rariplicata) , Blue-point . 29

Megalonaias gigantea, Washboard . 28

Actinonaias carinata, Mucket . 20

Elliptio dilatatus, Spike . 17

Fusconaia undata, Pig-toe . . . 14

Quadrula jmstulosa, Pimple-back . . 12

Elliptio crassidens, Elephant’s Ear . 10

Fusconaia ebenus, Niggerhead . 7

Lampsilis ventricosa, Pocketbook . 7

Lampsilis siliquoidea, Fat Mucket . 5

Quadrula quadrula, Maple-leaf . 5

Pleurobema ( cor datum ) pyramidatum and

P. c. coccineum. Small Niggerhead . 4

Proptera alata, Pink Heel-splitter . . . 2

Plethobasus cyphyus, Bullhead . 2

Ligumia recta, Black Sand-shell . 2

Quadrula metanevra, Monkey-face . 1

Obovaria olivaria, Hickory-nut . 1

Tritogonia verrucosa, Buckhorn . 1

Quadrula' nodulata, Warty-back . 1

Cyclonaias tuberculata, Purple Warty-back.. .

Alasmidonta marginata, Elk-toe . .

Arcidens confragosus, Rock Pocketbook .

Plagiola lineolata, Butterfly .

Lampsilis fallaciosa, Slough Sand-sliell . ......

Snails

Campeloma integrum . . 1

Campeloma rufum . .

Pleurocera acuta .

Fishes

Bowfin, Amia calva . 29

Fresh-water Drum, Aplodinotus grunniens . 24

Channel and/or Blue Catfish, Ictalurus sp . 20

Bullhead, Ictalurus ( Ameiurus ) sp . . 19

Suckers and Buffalofish, Catostomidae . 18

Buffalofish, Ictiobus sp. . . 10

Gar, Lepisosteus sp . 7

Longnose Gar, Lepisosteus osseus . . . 3

Pike, Esox sp . 3

Northern Pike, Esox lucius . . 2

Redhorse, Moxostoma sp . 2

Bass, Micropterus sp . 1

Sturgeon, Scaphirhynchus sp.? . 1

Flathead Catfish, Pylodictis olivaris . 1

Smallmouth Buffalofish, Ictiobus bubalus .

Turtles

Box Turtle, Terrapene sp. . . . 31

Soft-shelled Turtle, Trionyx (Amy da) sp . 26

Turtle spp . 20

Pond Terrapin, Pseudemys scripta . 12

Turtle, Pseudemys, Graptemys, Chrysemys group . 8

Snapping Turtle, Chelydra serpentina . 1

Map Turtle, Graptemys geographica . 1

Blanding’s Turtle, Emys blandingii . 1

Baker

(1931; 1936)

14 +

Transactions Illinois Academy of Science

Birds

Mallard, Anas platyrhynchos, and/or Black Duck,

A. rubripes . 25 3

Turkey, Meleagris gallopavo . 17 7

Canada Goose, Branta canadensis . 11 2

Prairie Chicken, Tympanuchus cupido . 10 5

Duck spp . 7

American Coot, Fulica americana . 5 2

Trumpeter Swan, Olor buccinator . 5 2

Wood Duck, Aix sponsa . . 5 1

Bobwhite, Colinus virginianus . ... 4 2

Redwinged Blackbird, Agelaius phoeniceus . 4 1

Green-winged Teal, Anas carolinensis . 4 1

Bufflehead, Bucephala albeola . 3 2

Lesser Scaup, Aythya affinis, and/or

Ring-necked Duck, A. collaris . 2 3

Canvasback, Aythya valisineria . . 2 2

Blue-winged Teal, Anas discors . 2 3

Long-billed Curlew, Numenius americanus . 2 1

Sora, Porzana Carolina . 2 2

Snow and/or Blue Goose, Chen sp . . 2

Pintail, Anas acuta . 2 1

Flicker, Colaptes cf. auratus . 1 1

Blue Jay, Cyanocitta cristata . 1

Rusty Blackbird, Euphagus carolinus ? . 1

Sparrow Hawk, Falco sparverius . 1

Double-crested Cormorant, Phalacrocorax auritus . 1

Black-crowned Night Heron, Nycticorax nycticorax . . . . 1 2

Whistling Swan, Olor columbianus . 2

Hooded Merganser, Lophodytes cucullatus . 2

Bald Eagle, Haliaeetus leucocephalus . 2

American Bittern, Botaurus lentiginosus . 1

Red-tailed Hawk, Buteo jamaicensis . 1

Red-shouldered Hawk, Buteo lineatus . . 1

Broad-winged Hawk, Buteo platypterus . 1

Sandhill Crane, Grus canadensis . . . 1

Woodcock, Philohela minor . 1

Short-billed Dowitcher, Limnodromus griseus . 1

Shoveller, Spatula clypeata . 1

Grackle, Quiscalus quiscula . . . . . 1

Mammals

White-tailed Deer, Odocoileus virginianus . 404 64-

Beaver, Castor canadensis . 37 84-

Raccoon, Procyon lotor . 25 2

Elk, Cervus canadensis . . 24 1

Canids: Canis sp., and Dog, C. familiaris . 22 13-4-

Muskrat, Ondatra zibethica . 17 7

Cottontail, Sylvilagus floridanus . 13 2

Fox Squirrel, Sciurus niger . .... 12 4

Mink, Mustela vison . . 4 64-

Bison, Bison bison . 4

Bobcat, Lynx rufus . . . . . 2 1(?)

Gray Squirrel, Sciurus carolinensis . 2 1

Striped Skunk, Mephitis mephitis . 2

Franklin Ground Squirrel, Citellus franklinii . 2

Marsh Rice Rat, Oryzomys palustris . 1 3

River Otter, Lutra canadensis . 1 1

Gray Wolf, Canis lupus . . . 1

Opossum, Didelphis marsupialis . 1

Common Mole, Scalopus aquaticus . 2

White-footed Mouse, Peromyscus cf. leucopus . 1

Faunal Records from Illinois

species were determined, and 69 per¬
cent of the remains were those of
aquatic forms. With the possible
exception of Emys blandingii, all of
the turtles represented are still com¬
mon in that area.

Fish were well represented in the
general midden deposit and in re¬
fuse pits, with at least 12 species
being* identified. Judging from the
size of many specimens, and from
the species involved, most of the fish
were taken in the Illinois River. Re¬
mains of the bowfin were the most
numerous (29), while bones of sev¬
eral species of other “ rough fish,”
the gar and catastomids, comprised
33 percent of the total. Catfish and
bullheads were taken in considerable
numbers, while drum ranked sec¬
ond in the number of bones re¬
covered ; compared with specimens
of known weight, the majority of
drum caught by these Indians
weighed between 6 and 10 pounds.
Considering the large size of many
of the drum, catfish and buffalofish,
fish were an important source of
food to these people.

Apparently few ‘ ‘ game fish ’ ’ were
caught by the Indian as evidenced
by paucity of remains. In addition
to the bass and pike listed in Table
1, Simpson (1939) mentions many
erappie ( Pomoxis sp.) scales found
in a grave fill (identified by Dr.
D. F. Hansen, Dept of Zool., U. of
I., Urbana). Through the courtesy
of Dr. Donald F. Hoffmeister, Di¬
rector of the Museum of Natural
History, University of Illinois, Ur¬
bana, the author was able to examine
the faunal specimens described by
Baker (1936). One apparent error
is the identification of a dentary as
Stizostedion ; this jaw section ap¬

pears to be Esox rather than sauger
or walleye. The presence of north¬
ern pike at this site is noteworthy
since this fish is now restricted (ex¬
cept where re-introduced) in Illi¬
nois to the northern sections of the
Mississippi River. Two large jaw
sections of the pike recovered by Mr.
Elliott were from fish that weighed
6 to 8 pounds. Parts of three small¬
er mandibles are also probably E.
Indus, but may be referable to the
redfin pickerel, E. americanus.

Summary

Faunal remains recovered at the
Kingston Lake Site in Peoria Coun¬
ty, Illinois, between 1932 and 1943
were discussed. A minimum of 12
species of fish, 6 species of turtles,
36 species of birds and 20 species of
mammals were identified, the ma¬
jority of which still occur in the re¬
gion. Several, such as the bison,
elk, otter, gray wolf, turkey, and
trumpeter swan, are now extirpated
in Illinois while the prairie chicken,
long-billed curlew, bobcat, white¬
tailed deer and marsh rice rat are
no longer present locally. Extensive
river and bottomland habitat pro¬
vided a variety of game species which
were utilized by the Indian. Fresh¬
water mussels were also important
food items, and 24 species were iden¬
tified from this site.

Literature Cited

Baker, Frank C. 1931. Additional
notes on animal life associated with
the mound builders of Illinois. Trans.
Ill. Acad. Sci., 23: 231-235.

Baker, Frank C. 1936. Remains of
animal life from the Kingston kitchen
midden site near Peoria, Illinois.
Trans. Ill. Acad. Sci., 29:243-246.

Transactions Illinois Academy of Science

Griffin, John W., and Donald E. Wray.
1945. Bison in Illinois archaeology.
Trans. Ill. Acad. Sci., 38: 21-26.
Parmalee, Paul W. 1957. Vertebrate
remains from the Cahokia site, Ill.
Trans. Ill. Acad. Sci., 50: 235-242.
Parmalee, Paul W. 1958a. Marine
shells of Illinois Indian sites. Nau¬
tilus, 71 (4) : 132-139.

Parmalee, Paul W. 1958b. Remains of
rare and extinct birds from Illinois
Indian sites. Alik, 75(2): 169-176.
Simpson, A. M. 1939. The Kingston
village site. Peoria Acad. Sci., Arch.
Sect., 15 pp.

Manuscript received November 21, 1961.

RESPONSE OF BOBWHITE QUAIL TO MANAGEMENT
ON SOME ILLINOIS STRIP-MINED LANDS

PAUL VOHS, JR. and DALE E. BIRKENHOLZ
Iowa State University and University of Florida

Population changes of bobwhite
quail, Colinus virginianus, in re¬
sponse to habitat development on
strip-mined lands have been studied
in southern Illinois since 1954. This
has been conducted as a part of a
cooperative research project to eval¬
uate the potential of spoil banks as
recreational areas with primary
emphasis on hunting and fishing.
Initially, population levels of bob-
whites precluded productive hunting
on stripmine lands and it was con¬
sidered important to determine the
feasibility of developing such areas
to increase resident game popula¬
tions. Earlier efforts toward recla¬
mation on strip-mined lands have
most often been directed towTard
forestry, horticultural and grazing
practices (Klimstra, 1959).

Acknowledgments

Truax-Traer Coal Company, Mid¬
west Coal Producers Institute, Illi¬
nois Department of Conservation
and the U. S. Fish and Wildlife
Service aided in the development
and continue to provide support for
the study. Dr. W. D. Klimstra, Di¬
rector, Cooperative Wildlife Re¬
search Laboratory and Professor of
Zoology, Southern Illinois Univer¬
sity, supervised a portion of the re¬
search. These data are a contribu¬
tion from project No. 26, Coopera¬
tive Wildlife Research, Southern Il¬

linois University and No. W-64-R,
Illinois Department of Conservation.

Description of Area

A 920-acre tract, located 6 miles
south of Pinckneyville and 3 miles
west of Pyatts, Perry County, Illi¬
nois was deeded to Southern Illinois
University for research purposes by
the Truax-Traer Coal Company.
Small agricultural fields interspersed
with woods border on the west and
partially on the south ; similar strip -
lands lie adjacent to all other bound¬
aries.

The spoilbanks on the research
area, formed from 1932 through 1941
(excluding 1934), vary in direction,
length and height. Erosion has
rounded the crests and deposited al¬
luvial materials in the valleys be¬
tween the ridges. Soils, at or near
the surface, generally have sufficient
quantities of available phosphorus
and potash ; nitrogen is available at
the rate of 30 to 40 pounds per acre
(Birkenholz, 1958). Except for
shale and gob deposits, soils are
neutral or basic. Analysis of soils
and vegetation indicate that soil fer¬
tility is better than adjacent farm
lands not under proper management
(Klimstra, 1959).

Approximately one-half of the to¬
tal area, including the south and
western portions, was planted to
trees in 1942 and 1943. Pine planta-

Transactions Illinois Academy of Science

tions contain short-leaf pine ( Pinus
echinata) and jack pine ( Pinus
banksiana) ; deciduous species in¬
clude black locust, osage-orange
(. Madura pomifera) , catalpa ( Catal -
pa speciosa ), silver maple (Acer
saccharinum ) and oaks ( Quercus
spp.). Excluding the black locust
plantings, these plantations, charac¬
terized by a dense stand of trees and
sparse understory, are of little value
to upland game and require exten¬
sive management to produce suitable
habitat.

The main herbaceous species (sci¬
entific nomenclature after Jones,
1950) on naturally revegetated spoil-
banks include sweetclover ( Melilotus
alba and M. Officinalis ), goldenrod
(Solidago spp.), woody aster ( Aster
pilosus ) and cheat (Bromus secali-
nus). Interspersed widely with
these herbs are clumps of sumac
(Rhus glabra and R. copallina),
poison ivy (Rhus radicans) and
blackberry (Rubus frondosus) . Cot¬
tonwood (Populus deltoides), syca¬
more (Plantanus occidentals), wil¬
low (Salix interior), elm, (Ulmus
rubra), box elder (Acer negundo)
and persimmon (Diospyros virg'ni-
ana) are common trees.

Previous studies (Brewer and
Triner, 1956; Verts, 1957) showed
that the vegetation in the naturally
revegetated areas is similar in each
age group of spoils; species composi¬
tion, number of stems, average height
of vegetation, and percentage of bare
ground varies little from areas mined
in 1932 to spoilbanks formed in 1941.
The only significant difference in the
vegetation on the oldest spoils as
compared with those more recently
formed is the increase in diameter
and height of the trees.

Typical of many Southern Illi¬
nois strip-mined areas, there are two
unmined tracts on this research area.
A 51-acre plot near the center and a
smaller one along the west border
provide about 60 acres of tillable
land though the clay soils are of low
fertility and poorly drained.

On the basis of initial studies
(Brewer and Triner, 1956; Verts,
1956) it was believed that spoilbank
habitat could be improved for up¬
land game. The uniformity of vege¬
tative cover as well as the lack of
it in some areas, the absence of plant
species which provide food, and the
lack of openings or breaks in plant
distribution were considered major
limiting factors.

Techniques

Roads, totaling 4.2 miles in length,
were constructed. A total of 12.1
miles of spoil crests were leveled ;
0.9 mile of spoil valley was graded
and widened. Selected areas of
spoilbanks were leveled to provide
plots ranging from 0.25 acre to 1.5
acres in size. These developments
not only created critically needed
“edge” but also bare ground for es¬
tablishing plant species which would
benefit bob whites.

Korean lespedeza (Lespedeza stip-
ulacea) was broadcasted on road¬
sides, leveled areas, non-mined lands
and spoilbanks lacking in ground
cover each winter after 1955. Es¬
tablishment has been successful on
over 100 acres of the research area ;
natural reseeding occurs each year.

Sericea and bicolor lespedezas
(Lespedeza sericea and bicolor) were
planted on spoils, roadsides and in
prepared food plots. Food patches

Quail Management on Strip-mined Lands

containing: combinations of Korean
lespedeza, German millet ( Setaria
italica), sorghum ( Sorghum vul¬
gar e), soybeans ( Glycine max), corn
(. Zea mays) and buckwheat ( Fago -
pyrum esculentum) were established
on the unmined areas and in spoil -
banks on leveled areas. Row crops
were planted most years on the in¬
terior and perimeter areas that were
not mined.

Since 1954, quail populations have
been censused each year just prior
to the hunting season (Table 1) ; in
addition, daily observations of coveys
have been maintained. Except for
1958, censuses were accomplished in
2 or 3 days by 5 to 12 persons walk¬
ing abreast along the crests of ad¬
jacent spoilbanks. Dogs were utilized
where possible, but dense patches of
briars and the undulating terrain
restricted their effectiveness. The
fall population in 1958 was esti¬
mated on the basis of the number of
quail present the following spring.

Crops were obtained from 49 quail
harvested from November 14 to De¬
cember 23, 1959, on the research
area. Food items from the crops
were identified and compared with
contents from crops of six quail col¬
lected during the fall of 1955.

Presentation
and Analysis of Data

The number of bobwhites on the
research area (Table 1) increased
from 46 birds in 1954 to 279 in the
fall of 1959. The population nearly
doubled from 1954 to 1955 and from
1955 to 1956 ; increase was more
gradual in subsequent years. The
population buildup continued on the
research area through 1958 and 1959

Table 1. — Bobwhite Quail Populations
on the Pyatts Striplands Re¬
search Area, November, 1954-
59.

Year

Number

Coveys

Mean

Covey

Size

Total
Number
of Quail

1954...

11.5

1955...

12.0

1956. . .

16.3

163

1957...

13.9

167

1958...

16*

13.5*

216*

1959...

13.9

279

* Estimated on basis of prenesting
population and routine field observa¬
tions.

despite decreases in regional popu¬
lations (Klimstra, 1958, 1959).

Excluding the pine and hardwood
plantations not used by quail, the
population density was about one
bird per 2.3 acres of habitat after
4 years of management ; nearly
250% more than on unmanaged
farm lands in southern Illinois
(Klimstra, 1959). This increase in
population, when compared with ad¬
jacent unmanaged, but similar, spoil-
bank areas, is striking. The high
population level afforded hunting in
1959 even though the terrain was
more difficult to traverse than non-
mined areas.

Roads and leveled crests of spoils
were important in both management
and hunting of quail. The roads
were valuable to quail as loafing
sites when vegetation was wet and
in providing edge. Food plots were
developed adjacent to roads and
leveled crests of spoils to facilitate
access for management and hunting.
Quail did not include in their ranges
areas where there had been extensive
leveling of spoils or the isolated

Transactions Illinois Academy of Science

leveled plots when no food producing1
plants were provided. Regrowth of
sweetclover, briars, cottonwoods, and
cheat was more rapid on these
cleared and unplanted areas than on
areas planted to favorable plant
species following disturbance of the
natural vegetation.

With the exception of black locust
plantings, tree plantations were of
little value as quail habitat at any
time during the year. The open
canopy resulting from destruction
by the locust borer on the trees pro¬
duced an understory vegetation.

With food available from the locust
and cover from the tangled under¬
story, quail utilized segments of the
locust plantations; however, hunt¬
ing was next to impossible because
of the dense growth. Large planta¬
tion plantings (exceeding 5 acres in
size) are not recommended for spoil -
banks which are managed for wild¬
life. Limited pine plantings may be
used to enhance aesthetic qualities
of striplands and to provide edge,
but large block plantings offer little
food or protection for bobwhites.

Table 2. — Food Items Identified from Crops of Bobwhite Quail Collected on the
Pyatts Stripland Research Area, Fall, 1959.

1959

Food Item

Per cent
occurrence

49 crops

Rank

Per cent
volume

Korean lespedeza

(Lespedeza stipulaceae) .

94.0

73.0

Common lespedeza

( Lespedeza striata) .

26.0

trace

German millet

(Setaria italica) .

24.0

17.4

Sweet clover

( Melilotus spp.) .

20.0

trace

Small wild bean

( Strophostyles leiosperma ) .

18.0

trace

Common ragweed

(. Ambrosia elatior ) .

16.0

trace

Tick-clover

( Desmodium spp.) .

16.0

1.2

Lance-leafed ragweed

( Ambrosia bident at a ) .

14.0

trace

Dwarf sumac

{Rhus copallina ) .

12.0

1.2

Trailing wild bean

( Strophostyles helvnla) .

8.0

trace

Beggar-ticks

(Bide ns spp.) .

8.0

trace

Wild black cherry

(. Prunus serotina) .

8.0

1.2

Grit .

8.0

trace

Short horned grasshopper

( Locustulae ) .

6.0

trace

Cheat

( Bromus secalinus ) .

6.0

trace

Quail Management on Strip-mined Lands

Table 2. — Continued.

1959

Food Item

Per cent
occurrence

49 crops

Rank

Per cent
volume

Pennsylvania smartweed

{Polygonum pennsylvanicum ) .

6.0

trace

Leafy material .

6.0

trace

Hemiptera

( Redviidae ) .

6.0

trace

Sorghum

( Sorghum vulgare ) .

4.0

trace

Partridge-pea

( Cassia fasciculata ) .

4.0

trace

Leaf beetles

{Chrysomelidae) .

4.0

trace

Soybeans

{Glycine max) .

2.0

trace

Smooth sumac

{Rhus glabra) .

2.0

trace

Black locust

{Robina pseudoacacia) .

2.0

2.5

Rough but ton weed

{Dioda teres) .

2.0

trace

Panic grass

{Panicum spp.) .

2.0

trace

Paspalum spp .

2.0

trace

Yellow foxtail

{Setaria lutescens) .

2.0

trace

Spiders

(Arachnidae) .

2.0

trace

Total Number of Food Items .

Thinning* and block cutting (or
using* chemical herbicides) within
plantations might enhance their
wildlife values, but are expensive
and time consuming. Management
efforts on older striplands may be
more profitably directed toward re¬
tarding succession on naturally re-
vegetated areas and establishing food
producing plants.

The importance of providing food
was evident in the analysis of crops
of bobwhites collected on the re¬
search area (Table 2). Crops of
quail collected in 1959 contained Ko¬
rean lespedeza, German millet, black

locust, tickclover, dwarf sumac, and
wild black cherry in amounts in ex¬
cess of 1% of the total volume. Ko¬
rean lespedeza was the most im¬
portant single food, occurring in
94% of the crops and yielding 73%
of the total volume.

Though a small sample for com¬
parative purposes, the crops of six
bobwhites collected on the area in
1955 contained seeds of corn (avail¬
able in a food plot), trailing wild
bean ( Strophostyles helvola), small
wild bean ( S . leiosperma), rushfoil
( Crotonopsis elliptica), lance-leafed
ragweed ( Ambrosia bidentata) , beg-

Transactions Illinois Academy of Science

gar-ticks (Bidens spp.) and grass¬
hoppers in amounts exceeding 1%
of the total volume (Verts 1956).
Davison (1958) classified trailing
and small wild beans and ruslifoil as
inferior quail foods. Even though
available in very limited amounts,
cultivated species comprised 50% of
the volume of the crops (primarily
corn). Korean lespedeza was not
recorded in any of the crops.

The establishment of several covey
ranges could be correlated with the
development of food plots in the
spoilbanks, especially where Korean
lespedeza had been planted. A total
of 227 observations of coveys was
recorded during the winters of 1957-
58 and 1959-60; 70% of these oc¬
curred in or immediately adjacent
to patches of Korean lespedeza.
Quail began using this lespedeza as
the seeds matured in late September
and early October. Continued and
intensive use of lespedeza plantings
was recorded until other foods be¬
came available in late March and
early April even when harassed by
hunters during the legal season.

Broods of quail utilized food
patches on the unmined areas dur¬
ing the summer ; this use continued
until the hunting season in Novem¬
ber, indicating the importance of
managing unmined areas associated
with stripped lands. Following the
initial exposure to hunting, how¬
ever, the birds could only infrequent¬
ly be located near the food patches
or even on the unmined tracts.
Stoddard (1931) reported that dog
handlers spoke highly of small food
patches as being a great help in
locating coveys when they started
to train their dogs in the fall. In
Stoddard’s study, general use of the

food patches ceased soon after work
with the dogs began. Utilization of
the food plots on the unmined areas
followed a similar pattern, but con¬
tinued use of introduced foods within
the spoilbanks was noted even after
constant harassment by hunters.

One of two coveys utilizing a food
patch in 1959-60 was completely an¬
nihilated after being located on six
separate occasions during the hunt¬
ing season in the same food patch ;
the other covey was reduced to one-
half of the original number, but con¬
tinued to utilize the food plot. The
food plots within the spoils served
as starting points for hunters with
dogs and high success was recorded
in locating coveys during 1959-60.
Quail were not always present with¬
in the plots, but were often trailed
from the plots by dogs. Though
Stoddard (1931) warned against lo¬
calization of coveys during the hunt¬
ing season to prevent over harvest
unless shooting is controlled, localiza¬
tion is necessary for management of
quail on strip-mined lands.

A few of the coveys on the entire
area may bear the brunt of the hunt¬
ing pressure, but the terrain and
availability of escape cover gen¬
erally makes the hunting of singles
unprofitable. Careful observation of
flushed birds sometimes makes a
second contact with a segment of a
covey possible. The distribution of
the coveys throughout the research
area in 1959-60 was such that a num¬
ber of coveys were not located by
hunters during the season. The
danger of over harvest is considered
to be very minor.

Summary

Responses of bobwhite quail to

Quail Management on Strip-mined Lands

management practices applied on
920 acres of strip-mined land in
southern Illinois have been studied
since 1954. Practices employed in¬
cluded road construction, grading of
spoils and widening of spoil valleys,
manipulation of cover and intro¬
ducing plant species which produce
food for quail. An increase of bob-
whites from 46 prior to incorpora¬
tion of management to 279 in 1959-
GO was recorded. An analysis of the
crops of 49 quail obtained in 1959-
GO indicated a dependence of the
birds on plant species introduced
through management.

Huntable populations of quail on
naturally revegetated spoilbanks de¬
pends largely on the establishment
of a suitable food supply and to a
lesser extent upon altering the vege¬
tative pattern to provide diversity.
Large tracts with extremely homo¬
geneous vegetative cover are as de¬
trimental to quail populations as are
farmlands where cover is wanting.
Efforts should be directed toward re¬
tarding or disrupting the natural
plant establishment by bulldozing,
burning or application of herbicides.
The selection of specific spoilbanks
to manage would depend upon the
topography and the accessibility for
hunting. Less expense would be in¬
curred if management practices were
initiated as soon after mining as pos¬
sible. This would result in less com¬
petition as pioneer species would not
have become firmly established.

Literature Cited

Birkenholz, D. E. 1958. Reclamation
of a spoil-bank area for wildlife pur¬
poses. Master of Arts Thesis, South¬
ern Illinois University Library, 70 pp.
(Unpublished) .

Brewer, R. and E. D. Triner. 1956.
Vegetational features of some strip-
mined land in Perry County, Illinois.
Trans. Ill. State Acad. Sci., 48: 73-84.

Davison, Verne E. 1958. A summary
and reclassification of bobwhite foods.
Journ. Wildl. Mgmt., 22(4): 437-8.

Jones, G. N. 1950. Flora of Illinois, 2nd
ed. Amer. Midi. Nat. Monog. 5, Uni¬
versity of Notre Dame Press, Notre
Dame, Ind., 368 pp.

Klimstra, W. D. 1958. Progress quar¬
terly report of the Cooperative Wild¬
life Research Laboratory, Oct. -Dec.,
Cooperative Wildlife Research Labo¬
ratory, Southern Illinois University,
Carbondale, Illinois. (Typewritten).

Klimstra, W. D. 1959. Progress quar¬
terly report of the Cooperative Wild¬
life Research Laboratory, Oct.-Dec.,
Cooperative Wildlife Research Labo¬
ratory, Southern Illinois University,
Carbondale, Illinois. (Typewritten).

Klimstra, W. D. 1959. The potential
of wildlife management on strip-
mined areas. Illinois Wildlife, 14(2):
5-9.

Stoddard, Herbert L. 1931. The bob-
white quail — its habits, preservation,
and increase. Charles Scribner’s Sons,
N. Y., 559 pp.

Verts, B. J. 1956. An evaluation of
wildlife and recreational values of a
strip-mined area. Master of Science
Thesis, Southern Illinois University
Library, 61 pp. (Unpublished).

Verts, B. J. 1957. The population and
distribution of two species of Pero-
myscus on some Illinois strip-mined
land. Journ. Mammal., 38(1): 53-59.

Manuscript received April 26, 1661.

ON MULTIPLY MUTANT SETS

A. A. MULLIN

University of Illinois, Urbana

In this paper we will extend and
develop some of the results of two
earlier papers (Mullin, I960, 1961).
First we consider non-empty sets on
which there is defined some non¬
empty index set of closed binary
composition laws. Secondly, we
show some concrete and general
properties of sets that satisfy an anti-
closure condition relative to all of
the elements of some non-empty sub¬
set of the index set of composition
laws.

In the additive and multiplicative
monoids of non-negative integers the
set of all odd primes is a doubly
mutant set, i.e., it is a mutant with
respect to both addition and multi¬
plication.

Results

Prelemma : Consider the algebraic

system determined by the additive and
multiplicative monoids of non-negative
integers. Then there exists, in a con¬
structive sense, a maximal doubly mu¬
tant set of that system.

Proof: Put E equal to the set of all
odd primes, i.e., 1 3,5,7... 1 . Consider

the following infinite sequence Aj of
infinite sets:

demonstrate that result recall that the
set of all odd integers is a maximal
mutant under addition and make use
of the Unique Factorization Theorem
of arithmetic.

Definition: Consider an algebraic sys¬
tem (A, *j) where j £ J and J is non¬
empty. A set is said to be p-tuply mu¬
tant in (A, *j) where 0<p<card J,
provided that the set is a mutant with
respect to p and at most p of the com¬
position laws.

Lemma: Let <p be a homomorphism
from (A, *j) onto (B, ok) for all j e J
and k e K with, say, card J<card K.
Let M be a maximal p-tuply mutant set
of (A, *j) where o<p<card J and j e J.

If <p(M) c<p(M) then there exists a
cardinal number q where p<q<card K
such that <p(M) is a maximal q-tuply
mutant set of (B, ok) where k e K.

Proof: Use the prelemma as an ex¬
istence proof for multiply mutant sets.
Then apply lemma 1.7 of (Mullin, 1961)
q times.

Summary

Two new and fundamental prop¬
ositions, relating mutant sets to ele¬
mentary number theory and general
algebraic systems, are given.

Literature Cited

pi.

1 3

:Pi

P. P

H x2

12n+l

Then, clearly, A4 is a maximal

doubly mutant set of the system, rela¬
tive to all positive odd integers. To

Mullin, A. A. 1960. A Concept Con-
cerning a Set with a Binary Composi¬
tion Law. Trans. Ill. St. Acad. Sci.,
53 (%) : 144-145.

Mullin, A. A. 1961. Some Remarks
on a Relative Anti-closure Property.
Zeit. f. Math. Logik u. Grund. d. Math.,
Bd. 7 (2) : 99-103.

Manuscript received February 9, 1962.

COMPARATIVE METHODS OF TRAPPING SMALL
MAMMALS IN AN ILLINOIS WOODS

RAYMOND L. WILL

Illinois Department of Conservation, Macomb

The two basic methods for cen-
susing small mammals are live-trap¬
ping and snap-trapping. Because
of its relative convenience, the lat¬
ter method is the one most com¬
monly used. However, Bole (1939)
has reported an inverse relationship
between the size of the sample area
snap-trapped and the population
density obtained, suggesting a pos¬
sibility of serious error in the
method.

Stickel (1946) attempted to de¬
termine the magnitude of error in¬
volved, if any, in an experiment in
bottomland forest on the Patuxent
Research Refuge, Maryland, in Sep¬
tember, 1945. She live-trapped a
circular 17-acre area for 7 nights
with 293 Sherman metal box traps
spaced at 50 foot intervals. Fol¬
lowing this, a circular acre in the
center of the 17-acre area was snap-
trapped for 3 nights with 200 traps.
Live-trapping indicated a density of
6 to 7 white-footed mice ( Peromyscus
leucopus novel) or acensis) per acre;
snap-trapping, 23 per acre. The
population density obtained by snap¬
trapping appeared to be significant¬
ly erroneous. Therefore, she con¬
cluded that densities obtained by
snap-trapping should be used only
as relative indices.

Other workers drew conclusions
that were contradictory to Stickel’s.
Goodnight and Ivoestner (1942) com¬
pared live-trapping and snap-trap¬

ping on two plots of Illinois prairie
and concluded that, in general, 6 to
7 days were required to determine
the population density by live-
trapping, with 3 days of snap-trap¬
ping giving the same density. How¬
ever, their plots were only 62%
meters long by 10 meters wide, and
their conclusions were based on
cumulative totals of 7 species. Also,
spacing of live-traps and snap-traps
was alike.

Buckner (1957) compared live-
trapping and snap-trapping in south¬
eastern Manitoba. He used three
trapping methods : live-traps set
with a 66 foot grid spacing, snap-
traps with the same spacing, and a
standard line of snap-traps. He
concluded that the results of all
three methods were reliable, except
that, perhaps due to habitat condi¬
tions, snap-trapping in early sum¬
mer gave a population density only
one-half that of live-trapping. No
white-footed mice were taken in his
study.

Wetzel (1949) made several com¬
parisons of live-trapping and snap-
trapping in undisturbed woods near
Champaign, Illinois. His methods
were essentially similar to those of
Stickel. His results, however, were
directly contradictory. He found
the population densities of white-
footed mice obtained by live-trap-
ping and snap-trapping to be nearly
identical.

Transactions Illinois Academy of Science

In view of these findings, the
present study was undertaken to ob¬
tain additional evidence concerning
the reliability of population densi¬
ties obtained by snap-trapping. Field
work involving live-trapping of 13.9
acres was carried out in an upland
deciduous woods in Illinois in July
and August, 1960. Peromyscus leu-
copus novel) or acensis was used as the
experimental animal.

Acknowledgments

Sincere gratitude is due my wife
for extensive help with the field
work and manuscript, to Dr. Donald
F. Hoffmeister for advice, encourage¬
ment, criticism of the manuscript,
and aid in obtaining field equip¬
ment, to Dr. Robert A. Evers for aid
in plant identification, to my father,
Martin Will and cousins, Leonard
and Andrew Will, for permission
to use the study area, to Mr. Harry
Henriksen and Miss Alice Boatright
for art work. Results presented
herein represent a contribution from
the Department of Zoology and Mu¬
seum of Natural History, University
of Illinois.

Study Area

Geographic location and climate.
The tract of land comprising the
study area is the EV2> SE^, SE^,
Sec. 1, T.8N., R.6E., in the north¬
eastern part of Effingham County,
in south-central Illinois. The lati¬
tude is 37°10' N; the longitude,
88°28' W. The Shelbyville glacial
moraine, the dividing line of the
dark soils to the north and the light
soils to the south, is 20 miles north
of the area. Drainage is to the Lit¬
tle Wabash River and thence into
the Wabash and Ohio rivers.

The climate is characterized by a
wide range of temperatures. The
maximum summer temperature is
100° F or more. During the sum¬
mer, there are often extended peri¬
ods of hot, dry weather. The mini¬
mum winter temperature may be be¬
low -10° F. The temperature may
fluctuate widely during the winter,
with an occasional extended cold
spell. The average monthly tem¬
perature of January is 31.5° F ; that
of July, 77.4° F (Illinois Clima¬
tological Data, 1958). These repre¬
sent the low and the high average
monthly temperatures. The average
annual rainfall is about 40 inches.
The high average monthly rainfall
(4.51 inches) occurs in June; the
low (2.16 inches), in December (Illi¬
nois Climatological Data, 1958).

Geologic history , topography, and
soil types. The present appearances
and characteristics of the study area
resulted primarily from the effects
of Pleistocene glaciation. Of the
four advances of the ice-slieet into
Illinois during the Pleistocene, only
the third, or Illinoian, reached the
study area. It left drift deposits
that constitute the present subsoil.
The topsoil is formed of loess de¬
posits blown in during interglacial
periods. It varies in thickness, but
averages only a few inches.

Drainage of the area occurs in 3
directions. To the south, drainage
occurs through an east-west ditch,
with two fingers extending 300 to
500 feet northward. A large per¬
centage of the remaining area drains
westward to the creek. Some drains
northward through a shallow ravine
extending to the creek. A small
area near the center is relatively
flat. The greatest difference in ele-

Trapping Small Animals

ration is about 30 feet, the highest
elevation being a little more than
600 feet above sea level.

Management history. The study
area has been the property of the
present owners for 35 years. Dur¬
ing that time, it has never been
pastured and none of the boundaries
have been fenced. Occasionally
some of the larger trees were cut
for lumber. Even then, only the
logs were removed from the woods,
the rest of the tree being left to rot.
Dutch elm disease has caused high
mortality to the American elms.
High winds annually bring down
many of these dead elm trees along
with other branches and trees. As
a result of wind action and logging
operations, a great amount of rotting
debris is scattered throughout the
woods. A system of logging roads,
most of it overgrown with herbs and
small saplings, branches out from
an exit located near the center of
the eastern border of the woods.

Fauna. No attempt was made to
define the invertebrate community.
Vertebrate species observed during
the trapping operations were listed.
The Fowler toad ( Bufo woodhousei
fowleri) and the leopard frog ( Rana
pipiens ) were the only amphibians
noted. The eastern box turtle ( Ter -
rapene Carolina ) and the five-lined
skink ( Eumeces fasciatus) were the
commonly observed reptiles. The
avian species noted most often was
the tufted titmouse ( Varus bicolor).
The cardinal ( Richmondena cardi-
nalis), blackcapped chickadee ( Pa¬
nts atricapillus) , blue jay ( Cy -
anocitta cristata), white-breasted
nuthatch ( Sitta carolinensis) , whip-
poor-will ( Caprimulgus vociferus)
and red-bellied woodpecker ( Cen -

turns carolinus) were also common¬
ly observed.

Three species of mammals were
caught in the live-traps : white¬
footed mouse (Per omy sens leucopus
noveboracensis) , short-tailed shrew
(Blarina brevicauda) and eastern
chipmunk ( Tamias striatus). In¬
dividuals of three other species, the
eastern cottontail (Sylvilagus flori-
danus), eastern gray squirrel (Sciu-
rus carolinensis) and eastern fox
squirrel (Sciurus niger) were ob¬
served. Four species were known
by sign only. These were the red
fox (Vulpes fulva), striped skunk
(Mephitis mephitis), opossum (Di-
delphis marsupialis) and raccoon
(Procyon lotor) . The southern fly¬
ing squirrel (Glaucomys volans) has
occasional^ been observed during
logging operations.

Flora. A dense stand of shrubs
and trees was present over most of
the study area. White oak (Quer-
cus alba), black oak (Quercus velu-
tina), shagbark hickory (Carya
ovata), bitternut hickor}^ (Carya
cordiformis) and green ash (Fraxi-
nus pennsylvanicus) were the domi¬
nant tree species. Other species
were present, but insignificant as re¬
gards crown cover. Poison-ivy
(Rhus radicans) was the most preva¬
lent shrub, occurring in dense
patches, as scattered erect plants,
and as climbing vines. Other com¬
mon shrub species were buckbrush
(Symphoricarpos orbiculatus) , Vir¬
ginia creeper (Parthenocissus quin-
quefolia) and riverbank grape (Vitis
riparia). White snakeroot (Eupa-
torium rugosum ) was the most prev¬
alent herb, being present over most
of the area. Other common herbs
were the Virginia knotweed (Poly-

Transactions Illinois Academy of Science

gonum virginianum) , false Solo¬
mon ’s-seal ( Smilacina racemosa),
mayapple (Podophyllum pelt at um )
and common wood-sorrel (Oxalis
cymosa ) .

Materials and Methods

Traps ayid baits. Two types of
live-traps were used in this study.
One was constructed mostly of wood,
with a hardware cloth top and gal¬
vanized metal door. The trigger
mechanism was of the type described
by Fitch (1950). The other was
constructed of hardware cloth. Its
trigger mechanism was a modified
and improved version of the former,
a swinging hardware cloth partition
and brass wire fulcrum replacing
the heavy wire mechanism. No more
than 6 of the latter type were used
at any one time. Museum Special
snap -traps were used.

Preliminary trapping to find a
convenient and effective bait was be¬
gun on July 1. After 4 days of ex¬
perimental trapping, shelled corn
soaked in peanut oil was determined
to be a suitable bait for the live-
traps. To have a somewhat com¬
parable bait for the snap -traps,
cornmeal, mixed with peanut oil to
a crumbly consistency, was used.

Trap layout. All of the live-trap
locations employed during this study
are shown in Figure 2A. For con¬
venience, all of the locations are
shown as spaced at 50 foot inter¬
vals. Such preciseness was not the
case, however, for no attempt was
made to space the traps at exact 50
foot intervals. The trap locations
were marked by means of strips of
white cloth hung from convenient
branches. When setting the traps,

they were not all set just below the
markers, but at favorable locations
within 10 feet. There were 242 trap
locations, but no more than 177 were
utilized at any one time. Total ef¬
fective coverage was 13.9 acres, each
trap location being assigned an area
of 2500 square feet.

The location of the circular acre
in which snap-trapping was carried
out is also shown in Figure 2 A.

Field techniques. Marking was
accomplished by clipping the distal
2 joints of a toe, or combination of
toes, on the front feet. Age was de¬
termined by use of the juvenile molt
pattern described by Gottschang
(1956). Three age classes were rec¬
ognized : juveniles — those not yet
starting the juvenile molt ; sub-
adults — those in the process of the
juvenile molt; adults — those hav¬
ing completed the juvenile molt.
Two breeding conditions were rec¬
ognized for males : testes descended
and testes not descended. Three
breeding conditions were recognized
for females : not visibly pregnant,
pregnant and nursing.

Trapping methods. The essence
of the trapping procedure was to
live-trap a large area until the ap¬
parent home range of supposedly all
the individuals was known, and then
to snap-trap a central acre for a 3-
day period. In live-trapping, the
traps were set in late afternoon or
early evening and checked the fol¬
lowing morning, at which time they
were snapped. The decision to leave
the traps unset during the midday
was influenced by two factors. Ants
quickly carried off unprotected bait.
With the traps closed, it was not
possible for the ants to remove the

Trapping Small Animals

Fig. 1. — Live-trapping and snap-trapping results for Peromyscus leucopus.
A and B, location and date of each live-capture for all individuals. C, composite
of apparent ranges (note amount of overlap). D, snap-trapping results, showing
date and location of last previous live-capture. Point at which individual was
taken in snap trap is shown by an X. No. 36 not previously taken in a live-trap.
Small numbers indicate date of capture in July except where “A” indicates date
of capture in August. Where ranges of Numbers 3 and 38 overlap, lower dates
are for No. 3.

Transactions Illinois Academy of Science

kernels of corn. Also, the midday
heat would very likely have caused
the death of any animal captured
shortly after the traps had been
checked in the morning.

After a suitable bait was found,
live-trapping in rows 6 through 19
(see Fig. 2A) was continued through
July 9. Trapping records for mice
captured more than once indicated
that all but one seemed to have home
ranges near the periphery of the
trapping area. All but 2 were near
the southern edge. A large rec¬
tangular barren area of nearly 5
acres seemed to exist. Eighty-four
traps were set within this area on
the afternoon of July 11. There
were no captures the following morn¬
ing. It was felt that snap-trapping

a central acre, most of which was
apparently unoccupied, would be of
little or no value. Consequently,
the traps in rows 18 and 19 were
taken up and 19 of them placed in
the form of an “E” within the area
of rows 1 through 5. It was felt
that this would probably reveal the
presence of any new individuals
within that area. However, in 3
nights of trapping, July 13, 14, and
15, only 3 captures were made, all
of the mice being individuals pre¬
viously marked in the adjacent area.
After a one-week delay, the traps in
rows 14, 15, 16, 17, 18, and 19 were
used to fill in rows 1 through 5.
which bordered the area where most
of the mice were caught. This ar¬
rangement of traps was thought to

Fig. 2. — Trapping results for Blarina brevicauda and Tamias striatus. A indi¬
cates date in August. Number preceding the hyphen is the number of the indi¬
vidual ; that following, the date of capture. A, date and location of each capture
of a short-tailed shrew. Capture in a snap-trap indicated by an X. Dots represent
live-trap locations used during this experiment. Circle represents 1-acre snap¬
trapping area. B, date and location of each capture of a chipmunk.

Trapping Small Animals

permit the obtainment of more ade¬
quate data on home range. Live-
trapping in rows 1 through 13 was
then carried on for 4 nights, be¬
ginning July 25. An exception oc¬
curred on July 26 when, due to a
thunderstorm, only the traps in rows
2. 3, 4, 5, 6, and 7 had been set the
previous afternoon. After this peri¬
od, the location and apparent home
range of all or a majority of the in¬
dividuals was known.

Snap-trapping was begun on July
29. A circular acre was marked off,
using the eighth trap from the east
end of row 7 as the center (Fig. 2A).
This was one trap position removed
from the center of the area covered
by the live-traps in rows 1 through
13. The reason for this dissymmetry
was practical, rather than scientific.
The author wished to avoid an ex¬
tensive patch of poison-ivy. A total
of 160 traps was set within this acre.
In each quarter of the acre, 40 traps
were set at random. Snap-trapping
was carried on for 5 nights. The
first three nights were used to de¬
termine population density, the last
2 only to see if the rate of ingress
changed. As a further check on
movements, live-trapping was con¬
tinued in rows 1 through 13 on Au¬
gust 3, 4, and 5, at which time
thievery by a squirrel hunter forced
a halt to trapping operations.

Results

Trapping success. The results of
live-trapping are summarized in
Table 1. Individuals of 3 species
were captured. P. leucopus will be
treated extensively. Data for the
other 2 species will be treated later
in summary form. Live-trapping
was conducted 15 nights before and

3 nights after snap-trapping, for a
total of 18 nights. Of the former,
the 8 nights included in the periods
of July 6 through 9 and 25 through
28 are of primary importance. Dur¬
ing these 8 nights, which represented
only 71.6 per cent of the total trap
nights (1679), 91.4 per cent of the
total captures (58) occurred. July
1, 2, and 5, represented 21 per cent
of the trap nights, but accounted for
only 3.4 per cent of the captures.
This period occurred before a suit¬
able bait was found. The period of
July 12 through 15 represented 7.5
per cent of the trap nights, and ac¬
counted for 5.2 per cent of the cap¬
tures. This was a period of ex¬
ploratory trapping.

Eleven white-footed mice were tak¬
en a total of 38 times, or an average
of nearly 3.5 captures per individual.
By July 9, the fourth day of effec¬
tive trapping in rows 6 through 19,
all of the mice taken were recaptures.
In the effective trapping of rows 1
through 13, no new individuals were
found after the second day.

A summary of snap-trapping re¬
sults is given in Table 2. Of the 6
marked mice captured, 4 were taken
the first night, one the third night,
and one the fourth night. An un¬
marked individual was taken the
third night. No individuals were
caught the second and fifth nights.

Population composition. Of a to¬
tal of 12 individuals taken during
the present study, 7 were males and
5 were females. The age classes were
somewhat difficult to categorize, since
some individuals progressed to a suc¬
ceeding age class during the trap¬
ping period. For convenience, the
age class of an animal at its initial
capture has been given in Table 3.

Transactions Illinois Academy of Science

Table 1. — Live-trapping results.

Date

No. of
traps set

P. leucopus

B. brevicauda

T . striatus

Total

New

individuals

Recaptures

Total

New

individuals

i -

Recaptures

Total

New

individuals

Recaptures

Total

July 1....

Pre-snap-trapping

177

2. . .

104

5. . .

6. ..

7...

177

8...

177

9. . .

177

12. . .

13...

14. . .

15. ..

25...

169

26. ..

27...

169

28...

169

Total. .

1679

Aug. 3 -

Post-snap-trapping

169

4. . . .

169

5. . . .

163

Total . .

501

Table 2. — Snap-trapping results.

Date

Total

individuals

P. leucopus

B. brevicauda

Marked

Unmarked

Total

Marked

Unmarked

Total

Julv 29 .

30 .

31 .

Aug. 1 .

2 .

Trapping Small Animals

Of 7 males, 3 were juveniles, 3 were
sub-adults and one an adult. None
of the males had descended testes un¬
til the last week of July, when all
males captured had descended tes¬
tes. One juvenile, 2 subadult, and
2 adult females were taken. None
was pregnant or nursing.

Distribution. The location and
date of capture for each mouse tak¬
en are shown in Figures 1A and IB.
From these data, an approximation
of the home range of each individual
has been drawn. All of the home
ranges are shown in Figure 1C. A
large amount of apparently unoc¬
cupied space is evident. Clumping
is so pronounced that every individ¬
ual captured more than once shows
some degree of overlap of home
range. The home ranges vary con¬
siderably in size and shape. The
numbers involved are too few and
the variations too extreme to make
data on size of home range meaning¬
ful.

The location of the one-acre plot
that was snap-trapped is shown in
Figure ID. Each of the 6 mice taken
in a snap-trap is noted. The date of
capture and the approximate loca¬
tion of the trap in which it was
taken is given, along with the loca¬
tion and date of the last capture in
a live-trap.

Movement. The maximum distance
between recaptures is given for each
individual in Table 4. The greatest
distance was 570 feet (12). The
least was 100 feet (14). The aver¬
age for the 9 individuals captured
more than once was 234 feet. For
6 males it was 229 feet ; for 3 fe¬
males, 243 feet. One mouse (12)
moved 525 feet during a 3 night

period, and the next night moved
390 feet back toward the original
point of capture.

The distance traveled from the
last previous capture to the point
at which the animal was taken in a
snap-trap is given in Table 5. Only
2 animals were taken on consecutive
dates. One (3) traveled the greater
distance, while the other (12) trav¬
eled the shortest distance. The re¬
maining animals traveled intermedi¬
ate distances over periods varying
from 3 to 21 days.

Population densities. From the
13.9 acres live-trapped, only 11
white-footed mice were taken. This
gives a population density of 0.8
mice per acre. By employing a buf¬
fer strip, one-half of the average
maximum distance between capture
locations (117') added to the peri¬
phery of the trapping area, the pop¬
ulation density is lowered to 0.5
mice per acre.

A 3-niglit period of saturation
trapping with snap-traps is gen¬
erally thought to take all of the resi¬
dent animals in a one-acre trapping
area. The first 3 nights of snap-
trapping in the present study yielded
6 animals, giving a population densi¬
ty of 6 mice per acre. Adding the
same width of buffer strip as before
lowers the population denshy to 1.5
mice per acre.

Oth er Mammals. Trapping data
for the short-tailed shrew ( Blarina
hrevicauda) are summarized in Ta¬
bles 1 and 2. Five individuals were
taken a total of 6 times in the live-
traps, the single recapture occur¬
ring in the same trap as the initial
capture and only 2 nights later.
Three of the shrews died in the live-

Transactions Illinois Academy of Science
Table 3. — Population composition ( P . leucopu. s* and T. striatus)

Total

Sex

Age group

P. leucopus

T. striatus

Juvenile

Male

Subadult

Adult

Juvenile

Female

Subadult

Adult

Table 4. — Maximum distance between recaptures of P. leucopus

# of individual

Sex

Age*

# of times
captured

Max. distance
between recaptures

3 .

male

J to SA

320'

5 .

female

6 .

female

260'

7 .

male

SA to A

115'

8 .

male

160'

12 .

male

J to A

570'

14 .

male

SA to A

100'

15 .

female

16 .

female

320'

26 .

female

150'

38 .

male

110'

*J = juvenile; SA — subadult; A — adult; “to” indicates change in age class
over trapping period.

traps. Two unmarked shrews were
taken in the snap -traps, one the first
night and the other the fourth night.

The point of capture is given in
Figure 2A for all of the animals
taken. Live-trapping yielded a pop¬
ulation density of less than 0.4 in¬
dividuals per acre ; snap-trapping,
one per acre. Five of the 7 shrews
captured apparently were females.
Based on size, all of them were
thought to be adults. One had been
nursing young.

Data on live-trapping* of the east¬

ern chipmunk ( Tamias striatus) are
summarized in Table 1. Eleven in¬
dividuals were taken alive, and an¬
other was accidentally captured and
later found dead in the trap. The
former were captured a total of 22
times with 3 animals accounting for
all of the recaptures. None of the
3 was recaptured until the seventh
day of effective trapping, at which
time all but one individual had been
taken at least once.

The sex and age composition of
the chipmunk population is shown

Trapping Small Animals

Table 5. — Distance from last previous
live-capture to point at which
taken in a snap trap. ( P . leu-
copus)

# of

individual

Distance

traveled

# of days

3 .

300'

6 .

160'

7 .

120'

12 .

50'

15 .

150'

38 .

200'

in Table 3. Age was subjectively
based on size. Differential growth
rates of males and females might
have caused unreliable age class
data, since no males were classified
as subadults. One male appeared
to have undescended testes, only a
shriveled scrotal sac being present.

Distribution of the chipmunks is
shown in Figure 2B. Approximate
home ranges have been drawn for
the 3 animals that were recaptured.
The maximum distance between cap¬
ture locations was 250 feet for an
adult male (27), 140 feet for an¬
other adult male (24), and 110 feet
for a subadult female (25). The
average was 167 feet. The popula¬
tion density was less than 0.9 chip¬
munks per acre.

Discussion

Trapping success. It was felt that
all or nearly all of the white-footed
mice within the trapping area were
caught at least once during the live-
trapping operation. This conclusion
is supported by the fact that only
one unmarked mouse was caught in
the snap-traps. The 2 white-footed
mice (3 and 12) caught most fre¬
quently also had the largest home

ranges, indicating the presence of
little or no trap proneness. Neither
was caught more than 2 nights con¬
secutively in the same trap.

There is evidence that the recent
capture of one white-footed mouse
in a live-trap predisposes that trap
in some way for the capture of an¬
other individual. Three different in¬
dividuals were caught in one trap
within 3 consecutive nights, while in
4 other traps different individuals
were caught during 2 consecutive
nights. In still another trap, 2 dif¬
ferent individuals were captured
within 3 nights. These instances ac¬
counted for nearly one-third of the
total captures. Thus, in many cases,
the overlap of home ranges would
seem to be primarily the result of
an artifact. Even in nature, how¬
ever, this same sort of overlap very
likely occurs due to some stimulus,
such as urinating posts. It may be
assumed, therefore, that the data ob¬
tained from the live-trapping were
reasonably reliable.

Distribution and movement . Home
ranges of white-footed mice, as
shown in Figure 1C, were drawn
by using a 50 foot strip between
successive captures. The sharp
angles thus formed were then sub¬
jectively rounded off. For the pur¬
poses of this study, only an approxi¬
mation of the position of the home
range was needed. The variations
in size and shape of home ranges
was perhaps a reflection of the low
population density.

The distribution pattern of the
11 mice taken was unexplainable.
Large barren areas existed. Where
animals were present, much overlap
of home range occurred. The topog-
graphy and the amount of debris

Transactions Illinois Academy of Science

present were thought to be deter¬
mining factors. The mice seemed to
be in close proximity to the drainage
ways and in areas containing rela¬
tively larger amounts of debris. Yet
a similar situation existed in the
northeastern part of the trapping
area, with no mice being found.
Therefore, some other factor, or a
combination of factors, may have
caused the clumped distribution.

The average maximum distance
between recapture locations was 229
feet for 6 males and 243 feet for 3
females. The corresponding dis¬
tances found by Stickel (1946) were
146 feet and 93 feet. Nicholson
(1941) found that most of the mice
he studied moved less than 200 feet.
The animals in the present study
traveled considerabty greater dis¬
tances, especially the females. The
number of animals is so small that
this variation might be due to chance
alone. However, Blair (1940), in a

studv of the meadow vole in south-

ern Michigan, found greater move¬
ments in areas of lower population
densities.

Population densities. A total of
11 animals were taken in the live-
traps. Five of 7 animals captured
3 or more times had a home range
extending to the periphery of the
trapping area. Therefore, some in¬
dividuals may have ranged a con¬
siderable amount outside of the 13.9
acre trapping area. Some workers
(Dice, 1938; Stickel, 1946; Wetzel,
1949) have used a buffer strip in
their calculations in order to lessen
the amount of error involved. Stick¬
el (1946) used a buffer strip based
on the average maximum distance
between capture locations. In this
study, the average maximum dis¬

tance between capture locations was
234 feet. The addition of one-half
of this amount to the periphery of
the trapping area as a buffer strip
gives a corrected trapping area of
24.0 acres. By this method of cal¬
culation, the population density is
lowered from 0.8 to 0.5 mice per
acre.

Saturation trapping of a circular
acre for a period of 3 nights is a
method of determining population
density by the use of snap-traps.
Within the 3 night period of snap¬
trapping in the present study, 6 mice
were taken. Employing the same
type of buffer strip as before, the
population density obtained by snap¬
trapping is lowered from 6.0 to 1.5
animals per acre.

A comparison of the corrected
densities shows the population densi¬
ty obtained by snap-trapping to be
300 per cent larger than that ob¬
tained by live-trapping. Wetzel
(1949) found the density obtained
by snap-trapping to be nearly iden¬
tical to that found by live-trapping.
The results of the present study,
however, concur with and corrobo¬
rate the findings of Stickel (1946).
She found adjusted densities of 6
to 7 per acre by live-trapping and
23 per acre by snap-trapping, a dis¬
crepancy only slightly larger than
that found by the author. The an-
tlior agrees with her proposal that
snap-trapping be used only as a rela¬
tive index of population densities.

The low population density found
in this study is worthy of special
note. Wetzel (1949) found a densi¬
ty of 4 mice per acre in a central
Illinois woods in August. Burt
(1940) has given monthly popula¬
tion densities for the white-footed

Trapping Small Animals

mouse in southern Michigan. He
found the lowest density in May,
with another low point of the cycle
occurring in July or August. Al¬
though the latter period corresponds
to the time of the present study, his
population density of slightly more
than 5 animals per acre is much high¬
er than that reported here. The rea¬
son for this unusually low summer
population is not known.

Other mammals. Efficiency of the
live-traps in taking the short-tailed
shrew ( B . brevicauda) was low.
Five individuals were captured. One
of these was taken in the snap-trap-
ping area. Two unmarked individ¬
uals were taken in the snap -traps,
giving a total of 3 animals within
the circular acre. More than 4
shrews were surely present over the
remaining 12.9 acres. Moreover,
only a single shrew was recaptured.

Distribution of the shrew seemed
to be scattered (Fig. 2A), individ¬
uals being found in the area left
unoccupied by the white-footed
mouse. No records of movement
were obtained. Live-trapping yield¬
ed a population density of less than
0.4 animals per acre ; snap-trapping,
one per acre. The similarity might
make both methods seem reliable.
However, the presence of 3 shrews
within the circular acre and only 4
outside of it is good evidence that
at least one of the trapping methods
was in error.

Efficiency of the live-traps in tak¬
ing chipmunks (T. striatus) was
similar to their efficiency in taking
white-footed mice. No unmarked
chipmunks were taken after 8 days
of effective trapping. Three chip¬
munks (24, 25, and 27) seemed to
be trap prone, being the only 3 re¬

captured of a total of 11 taken alive.
One (24) was captured 4 times, the
other 2 each 5 times. None showed
any great movement, the average
maximum distance between recap¬
tures being 167 feet. The greatest
distance traveled was 250 feet by
an adult male.

A comparison of figures 1C and
2B indicates that the distribution of
the chipmunk closely paralleled that
of the white-footed mouse. This
would suggest that surface debris
might be the factor influencing the
clumped distribution of both, since
the shrew, primarily a sub-surface
dweller, occurred where both mouse
and chipmunk did not.

Twelve chipmunks, including one
found dead in a trap, were taken on
the study area. Only 3 individuals
were recaptured, and they seemed
to be trap prone. Hence, data on
home ranges were meager and prob¬
ably unreliable, making the calcula¬
tion of a buffer strip impractical.
Thus, all that can be said is that the
population density was less than 0.9
chipmunks per acre.

Summary

Live-trapping of small mammals
was conducted over a total of 13.9
acres of upland woods in south-cen¬
tral Illinois in July and August,
1960. Snap-trapping of a circular,
centrally located acre was carried
on for 5 nights with 160 Museum
Special traps. White-footed mice
( Peromyscus leucopus noveboracen-
sis ), short-tailed shrews ( Blarina
brevicauda) , and eastern chipmunks
( Tamias striatus) were captured in
live-traps. Individuals of the former
2 species were also taken in snap-
traps.

Transactions Illinois Academy of Science

Summarization of the results may
be categorized by species.

P. leucopus. — Eleven individuals
were captured a total of 38 times
in the live-traps. Approximate
home ranges were drawn, and great
variations in size and shape were
noted. The average maximum dis¬
tance between locations of capture
was 229 feet for 6 males and 243
feet for 3 females. The population
density determined by live-trapping
with a 50 foot grid spacing was 0.8
animals per acre ; the population
density obtained by saturation snap-
trapping for 3 nights, 6 animals per
acre. The adjusted population den¬
sities were 0.5 for live-trapping and
1.5 for snap-trapping. A serious
error in the results obtained from
the snap-trapping method is indi¬
cated.

B. brevicauda. — Five individuals
were taken in the live-traps and 2
in the snap-traps. No data were
obtained on home ranges or move¬
ments. The population densities ob¬
tained were less than 0.4 animals
per acre by live-trapping and one
animal per acre by snap-trapping.
The density figures are felt to be in
error, especially that of live-trap-
ping.

T. striatus. — Twelve individuals
were taken, giving a population den¬
sity of less than 0.9 individuals per
acre. Only 3 animals were recap¬
tured. Of these, one was taken 4

times ; the other 2, 5 times. The
greatest distance traveled was 250
feet by an adult male.

Bibliography

Blair, W. F. 1940. Home ranges and
populations of the meadow vole in
southern Michigan. Jour. Wildl. Mgt.,
4(2) : 149-161.

Bole, B. P., Jr. 1939. The quadrat
method of studying small mammal
populations. Cleve. Mus. Nat. Hist.,
Pub., 5(4) : 15-77.

Buckner, C. H. 1957. Population
studies on small mammals of south¬
eastern Manitoba. Jour. Mamm.,
38(1): 87-97.

Burt, W. H. 1940. Territorial behavior
and populations of some small mam¬
mals in southern Michigan. Mus.
Zool., Univ. Mich., Misc. Pub. 45.

Dice, L. R. 1938. Some census meth¬
ods for small mammals. Jour. Wildl.
Mgt., 2(3) : 119-130.

Fitch, H. S. 1950. A new style live-
trap for small mammals. Jour.
Mamm., 31(3) : 364-365.

Goodnight, C. J., and E. J. Koestner.
1942. Comparison of trapping meth¬
ods in an Illinois prairie. Jour.
Mamm., 23(4): 435-438.

Gottschang, J. L. 1956. Juvenile molt
in Peromyscus leucopus noveboracen-
sis. Jour. Mamm., 37(4): 516-520.

Illinois Climatological Data, Annual
Summary. 1958. Volume LXIII, No.
13.

Nicholson, A. 1941. The homes and
social habits of the wood-mouse ( Pero -
myscus leucopus noveboracensis ) in
southern Michigan. Am. Midi. Nat.,
25(1) : 196-223.

Stickel, L. F. 1946. Experimental
analysis of methods for measuring
small mammal populations. Jour.
Wildl. Mgt., 10(2): 150-159.

Wetzel, R. M. 1949. Analysis of small
mammal population in the deciduous
forest biome. Unpublished Ph.D.
thesis, Univ. Ill.

Manuscript received December 12, 1961.

ABSENCE OF RABIES IN SOME BATS AND SHREW S

FROM SOUTHERN ILLINOIS

E. W. PEARSON1 and THOMAS R. B. BARR-
Illinois Natural History Survey, Jlrbana

During1 July, 1958, a project was
initiated in southern Illinois to de¬
termine the relationship of distribu¬
tion, abundance, and ecology of
wildlife vectors to the incidence of
rabies (National Institutes of Health
Project E-1349). Two of several
related studies involved the collec¬
tion and testing of bats and shrews
for rabies.

The fact that bats sometimes carry
rabies is becoming well known ; each
year several bats are received for
rabies determination in the labora¬
tories of the Illinois Department of
Health. The history of chiropteran
rabies in the United States and else¬
where has been well summarized by
Martin (1959). In October, 1959,
Illinois became the twenty-third state
to report bat rabies (U. S. Dept, of
Hltli., Ed., and Welfare, 1959), and
Kansas recently became the thirty-
first state to report rabies in bats
(U. S. Dept, of Hltli., Ed., and Wel¬
fare, 1961).

Insectivores as possible natural
hosts for rabies are discussed by
Verts and Barr (1960), but no refer¬
ence concerning* rabid shrews has
been found. Two hundred and
sixty-six shrews (Verts and Barr,
1960) and 559 bats (Verts and Barr,

1 Present address: Wildlife Research
Center, Fish and Wildlife Service, Den¬
ver, Colorado .

'l Present address: Department of

Poultry Science, University of British
Columbia, Vancouver, B. C.

1961) from northwestern Illinois
have been tested for rabies, all with
negative results.

In the present study, 93 bats,
representing seven species, were col¬
lected from abandoned silica mines
and natural caves in three counties
(Union, Alexander and Pulaski) in
southern Illinois between December,
1958, and March, 1959 (Table 1).
All bats were captured within a
radius of less than 16 miles from a
site where rabid skunks were found
in June and September, 1958.
Twenty-five shrews and one mole
were caught from August, 1958 to
April, 1959 (Table 2) ; nine of the
shrews were trapped within 14 mile
and the remainder less than 20 miles
from recent sites of rabies.

Brain and salivary gland tissues
from all insectivores and 35 of the
bats were removed in the laboratory
and stored in a deep-freeze cabinet
at minus 20° C in separate tubes
containing a solution of equal vol¬
umes of glycerine and twice-normal
physiological saline until all tests
had been completed. Laboratory
technique for testing these tissues
was the same as described by Verts
and Barr (1960) .

The remaining 58 bats were tested
by Mr. Nathan Nagle, Director, Illi¬
nois Department of Health Labora¬
tory, Carbondale. Some of the tis¬
sues were preserved in the manner
previously described, but most brains

35 ]

Transactions Illinois Academy of Science

Tarle 1. — Species of bats and numbers of tissues tested for rabies
from southern Illinois.

Species

Number of
brains
examined

Number of
salivary glands
examined

Pipistrellus subflavus .

Myotis keenii .

Myotis lucifugus .

Myotis sodalis .

Eptesicus fuscas .

Plecotus raf inesquii .

Lasionycteris noctivagans .

Totals .

Table 2. — Species of insectivores and numbers of tissues tested for
rabies from southern Illinois.

Species

Number of
brains
examined

Number of
salivary glands
examined

Blarina brevicauda .

Sorex longirostris .

Scalopus aquaticus .

Totals .

and salivary glands were placed in
sterile glycerine when removed from
the animal. They were allowed to
stand for 24 hours at room tempera¬
ture before being used in mouse in¬
oculation tests. Sections of brain
and salivary gland tissues from each
bat were ground with pestle and
mortar into an emulsion, mixed with
nine parts of 0.9 per cent saline and
allowed to stand for 5 minutes at

room temperature. Quantities of
0.03 ml of the supernatant fluid
from each suspension were injected
intra cerebrally into three 4- to 6-
week old mice using a 0.25 ml
tuberculin syringe and a 27 gauge
needle ; this is essentially the mouse
inoculation test described by Ko-
prowski (1954). Mice dying within
4 days after inoculation were con¬
sidered to have succumbed for rea-

Absence of Rabies in Rats and Shrews

sons other than rabies infection. The
inoculated mice were observed for
28 days.

Results of tests of all tissues were
negative for rabies. The possibility
of finding rabid shrews and bats by
random collections is apparently
remote, but those exhibiting unusual
behavior will continue to be ex¬
amined.

Literature Cited

Koprowski, H. 1954. Laboratory tech¬
niques in rabies. Wld. Hlth. Org.
Mono. Ser., No. 23, pp. 56-68.

Martin, R. L. 1959. A history of
chiropteran rabies with special refer¬
ence to occurrence and importance in

the United States. Wildl. Dis., 3, 75 .

pp.

U. S. Department of Health, Educa¬
tion, and Welfare. 1959. Veteri¬
nary Public Health Newsletter; pre¬
pared by Veterinary Public Health
Section of Epidemiology Branch, Com¬
municable Disease Center, Atlanta,
Ga. November. 16 pp. Mimeo.

U. S. Department of Health, Educa¬
tion, and Welfare. 1961. Veteri¬
nary Public Health Section of Epi¬
demiology Branch, Communicable Dis¬
ease Center, Atlanta, Ga. January.
14 pp. Mimeo.

Verts, B. J. and Thomas R. B. Barr.

1960. Apparent absence of rabies in
Illinois shrews. J. Wildl. Mgmt.,
24(4): 438.

Verts, B. J. and Thomas R. B. Barr.

1961. An effort to identify rabies in
bats from northwestern Illinois. Cor¬
nell Vet. (in press) .

Manuscript received May 28, 1961.

ROOT GROWTH OF TRANSPLANTED LOBLOLLY PINE
( PINUS TAEDA L.) SEEDLINGS IN RELATION TO
CHEMICAL ROOT RESERVES

A. R. GILMORE

University of Illinois. Dixon Springs Experiment Station, Robbs, Illinois

The ability of a seedling* to pro¬
duce new roots after being trans¬
planted from the nursery affects the
possibility of its survival. The gen¬
eral practice in nurseries growing
loblolly pine seedlings is to run a
lifting blade eight inches below the
soil surface to remove the seedlings
with the least amount of damage.
Most of the fine rootlets and all roots
more than eight inches below the
soil surface are lost during this op¬
eration. This reduction in the
amount of moisture absorbing sur¬
face as compared to the transpiring
surface is critical and undoubtedly
often determines if a seedling will
survive.

Stone (1955) demonstrated that
new root growth is necessary for
survival of ponderosa pine and
Douglas-fir when transplanted, but
he did not determine which factors
were associated with the production
of new roots. Reed (1939) observed
that root growth of field-planted pine
seedlings in North Carolina is hin¬
dered by low soil moisture before the
wilting point is reached. This
means that a seedling transplanted
into the field must produce new roots
before soil moisture becomes low in
early summer if the plant is to sur¬
vive.

Many investigators have reported
the presence of sugars, starches and
other carbohydrate fractions in

plants and plant parts, but the litera¬
ture is nearly devoid of references
to the role that carbohydrates play
in the survival of transplanted
seedlings. A high carbohydrate
content in the roots of a seedling at
the time of transplanting could in¬
dicate a high capacity to supply
energy necessary for the production
of new roots.

Wakeley (1954) reported that
shading southern pine seedlings in
nursery beds, reduced the survival
of outplanted longleaf pine seedlings
26 per cent and slash pine seedlings
79 per cent. He attributed the high
mortality to the failure of the
shaded seedlings to develop new roots
promptly after transplanting.

Wassink and Richardson (1951)
reported that root growth of first
year seedlings of sycamore maple
was directly related to light intensi¬
ty, while root growth of red oak
seedlings of the same age appeared
to depend on stored food rather than
on products of current photosyn¬
thesis. In another study, Richard¬
son (1953) reported that when pho¬
tosynthesis was curtailed, root
growth of silver maple proceeded at
the expense of food stored in the
roots. He also found that the leaves
supply a stimulus essential to root
formation and growth.

Reines (1957) in his comprehen¬
sive review of rooting southern pine

[38]

Root Growth of Loblolly Pine

states that many studies have been
reported where the effects of car¬
bohydrates on rooting of plant cut¬
tings is mentioned.

Although little work has been
done on root growth of transplanted
pine seedlings, results from several
studies indicate that root growth of
seedlings might be dependent on the
carbohydrates in the plant. In an
early study using root growth cham¬
bers, the author found that a weak
statistical relationship existed be¬
tween the carbohydrates in the roots
and root growth of transplanted
loblolly pine seedlings; but, because
of the wide variation in root growth,
it was difficult to reach a positive
conclusion regarding root growth
and reserve root carbohydrates.

Carbohydrates are known to be
translocated from the leaves through
the stem phloem and into the roots.
If this path of translocation could
be blocked at the time of transplant¬
ing, carbohydrates would be pre¬
vented from entering the roots and
a test could be made to determine
the correlation between the carbohy¬
drates in the roots at this time and
during subsequent root growth. A
scheme was devised that fulfills these
requirements and enabled the in¬
vestigator to test the results of the
early study more thoroughly.

Methods

Twenty 1-year-old loblolly pine
seedlings that were used in the ex¬
ploratory phase of the experiment
were girdled in the late spring of
1959. These seedlings and 20 control
seedlings (not girdled) were planted
in cans. All seedlings were removed
after 30 days and the roots visually

examined to determine whether or
not root growth had begun.

Year-old seedlings used in the
main part of the experiment were
removed from the nursery bed each
month from October, 1959 through
April, 1960. Forty seedlings were
girdled each month and then planted
in root growth chambers. To serve
as a control, each month 20 un¬
girdled seedlings were planted in
gallon containers. All seedlings were
observed over a 30-day period for
root growth.

Results

During the exploratory phase of
the experiment in the spring of 1959
a light pressure with the thumbnail
was all that was required to remove
the bark around the seedling stem.
Both seedlings that were girdled and
ungirdled developed new branch
roots in addition to elongation of old
roots after they were transplanted.
Root growth was approximately four
times greater on ungirdled seedlings
than on girdled ones.

Seedlings removed from the nurs¬
ery bed in October, 1959 and dur¬
ing the next 6 months, appeared to
be in a dormant state. The bark ad¬
hered to the stem very tightly in all
7 groups of seedlings. To girdle a
seedling it was necessary to double
ring the stem with a razor blade and
then scrape away the bark.

None of the seedlings girdled dur¬
ing the period from October, 1959
through April, 1960 showed any
visible root growth after they were
transplanted. In contrast, seedlings
that were not girdled during this
period developed new roots in addi¬
tion to elongation of old roots.

Transactions Illinois Academy of Science

Only a few of the girdled or un-
girdled seedlings in the experiment
died within 30 days after they were
transplanted. A large percentage of
seedlings girdled and transplanted
in March and April of 1960 started
height growth and the seedlings ap¬
peared to be healthy except for the
girdle on the stem.

Discussion

The primary difference between
seedlings transplanted in the late
spring 1959 and those transplanted
during the period from October,
1959 to April, 1960 must have been
in their growing status. It is well
known that easy mechanical removal
of bark from trees occurs only when
the cambium is physiologically ac¬
tive (Kramer and Kozlowski, 1960,
p. 8). The fact that bark could be
removed easily from the stems of
seedlings girdled in the spring of
1959 but was difficult to remove dur¬
ing the other periods indicates that
the former were physiologically ac¬
tive, whereas the other girdled
groups of seedlings were in a dor¬
mant state.

Root growth of seedlings girdled
in the spring of 1959, that were
thought to be physiologically active
at time of girdling, is not a signifi¬
cant event in itself. However, when
compared with dormant girdled
seedlings that produced no new roots
after being transplanted, the results
suggest that before a seedling can
produce new roots, the roots must
receive some stimulus from the leaves
or buds.

The supposition that a root growth
stimulus is translocated from the top
of the seedlings to the roots is fur¬

ther substantiated by the two groups
of seedlings tested in the spring of
1959. As previously stated, all seed¬
lings in the spring of 1959 appeared
to be in an active state of growth ;
seedlings from both groups devel¬
oped new roots. If root growth did
not depend upon a stimulus from
the needles or buds, the total amount
of root growth should be the same
in both groups of seedlings. The
fact that this did not result sug¬
gests that it is necessar}^ to replenish
some substance in the roots before
root growth can proceed at the same
rate as in ungirdled seedlings. This
substance cannot be stored in roots,
otherwise dormant girdled seedlings
would produce new roots. This ma¬
terial probabty is not one of the
major carbohydrates (reducing sug¬
ars, sucrose, or starches) found in
pine seedling roots, as loblolly pine
roots contain relative large quanti¬
ties of such carbohydrates during
periods in which transplantings are
made.

As previously stated, none of the
girdled seedlings during the 7 month
period produced either new roots or
elongation of old roots. For this
reason the carbohydrate content in
the seedlings roots was not deter¬
mined, as no correlation could possi¬
bly be established between root
growth and food reserves of the
roots.

Summary

No correlation was shown between
root growth of transplanted seed¬
lings and food reserves of roots.
However, certain results indicate
that before root growth can begin,
the roots must receive a stimulus
from the leaves or buds.

Boot Growth of Loblolly Pine

Acknowledgments

The work reported here embodies
a portion of a thesis submitted to the
School of Forestry of Duke Univer¬
sity in partial fulfillment of the re¬
quirements for the degree of Doctor
of Forestry.

Literature Cited

Kramer, P. J. and T. T. Kozlowski. 1960.
Physiology of trees. McGraw-Hill,
New York, 642 pp.

Reed, John F. 1939. Root and shoot
growth of shortleaf and loblolly pines
in relation to certain environmental
conditions. Duke Univ. Sch. For Bull.,
4, 52 pp.

Reines, M. 1957. Rooting of southern
pines. Proc. Fourth Sou. Conf. For.
Tree Improv., Athens Ga., pp. 116-121.

Richardson. S. D. 1953. Studies on
root growth in Acer saccharinum L.
II. Factors affecting root growth
when photosynthesis is curtailed.
Proc. ned. Akad. Wet., 56: 346-353.

Stone, Edward C. 1955. Poor survival
and the physiological condition of
planting stock. For. Sci., 1: 90-94.

Wakeley, P. C. 1954. Planting the
southern pine. Agr. Monograph No.
18. For. Ser., U.S.D.A., 233 pp.

Wassink, E C. and S. D. Richardson.
1951. Observations on the connection
between root growth and shoot il¬
lumination in first-year seedlings of
Acer pseudoplatanus L. and Quercus
borealis maxima (Marsh) Ashe. Repr.
from Proc. Koninklyke Nederlandsche
Akademie van wetenschappen (Ser.
C), Amsterdam, 54(5): 503-10. For.
Abst., 13, No. 3682.

Manuscript received December 31, 1961.

THE Ml RAM LIS— INSECT COMMUNITY IN ILLINOIS

W. V. BALDUF

University of Illinois, Urbana

Like the human community, the
natural communities consist of pro¬
ducers, parasites, predators, scaven¬
gers, visitors and vagabonds. In the
Mirabilis- insect community, as now
known in central Illinois, only the
producing, predacious and parasitic
classes, groups or strata have been
observed. However, more extensive
and intensive studies, at other times
and places, would almost certainly
reveal additional classes and species,
for the community is not static. The
thirteen organisms now known for
this community represent four eco¬
logical groups. The umbrella-wort,
Mirabilis nyctaginea (Miclix.), Nye-
taginaceae, constitutes the primary
producing center of the community
in that it directly supplies the food
and shelter for five phytophagous
insects, and indirectly makes possi¬
ble the life of a predator and six
species of parasites that subsist on
the phytophags, the latter thus be¬
ing secondary producers in this or¬
ganic complex.

/Specific Components of the Groups.
(1) Mirabilis nyctaginea, the pri¬
mary producing center, or sine qua
non of the community. (2) Phy¬
tophags: Heliodines nyctaginella

Gibson and II. ionis Clarke, Heli-
odinidae, Lepidoptera; Celerio line-
ata Fabr., Sphingidae, Lepidoptera;
Onychobaris subtonsa LeConte, Cur-
culionidae, Coleoptera ; and Cator-
hintha mendica Stal, Coreidae, He-
miptera. (3) Predator: Orius in-

sidiosus (Say), Anthocoridac, He-
miptera. (4) Parasites: Bracon
caulicola (Galian) and B. gelechiae
Aslim., Braconidae ; Tetrastichus
coerulescens Aslim., Eulophidae ; Ea-
pelmus allynii (French) and E.
cyaniceps Aslim., Eupelmidae; Neo¬
cat olaccus tylodermae (Aslim.),
Pteromalidae, (all Hymenoptera) .

Characteristics of the Species. Al¬
though the species composing each
group have food preferences in com¬
mon, each also has distinctive struc¬
tural characteristics that implement
its specific role in the community.
The concrete habits and structures
that appear to fit each species to
perform its own role are described
briefly here. More details on cer¬
tain life histories may be had in the
articles by Balduf and Wester, as
cited among the references below.

Mirabilis nyctaginea, as the pro¬
ducing center of the community, is
well suited to support its phyto¬
phagous insect dependents. It is a
perennial herb with a fleshy root-
stock, sturdy stems and lush ovate
leaves. Even the clusters of small
pink flowers afford some food to the
insects. At least on railroad em¬
bankments, where section crews mow
it, the plant grows two crops of
leaves and seeds annually, in May -
June and in August - September,
with a period of bare woody stalks
intervening. Through the natural
spread of seeds and extension of its
root system, and especially when

[42]

Insects Associated with Mirahilis in Illinois

aided by human conveyances, M.
nyctaginea is capable of rapid
spread ; this feature accounts for the
mobility of the community. In the
process of spreading*, it is favored by
its predilection for well-drained
gravelly soil, such as prevails on
railroad trackways, where it is now
common in Illinois. Indigenous to
the Great Plains west of the Missis¬
sippi river and the western part of
Illinois, its spread eastward began
about 1855, when the first railway
spanned the river, connecting those
new agricultural regions with the
populous urban centers along the
Atlantic (Balduf, 1957).

The Phytophagous Species of the
Community. Catorhintha mendica
is unique among the phytophags in
being heterometabolous, and that all
the stages live exposed on the sur¬
faces of the leaves or branches. Both
adults and nymphs feed on plant
sap obtained by piercing-sucking
mouth parts. Both stages move rapid¬
ly on foot, and the adult flies readi¬
ly. So far as known, C. mendica
feeds only on M. nyctaginea, but
possibly also utilizes other species
of Mirahilis, three of which occur
infrequently in Illinois (Jones and
Fuller, 1955). It is the only species
of the community that Hart (1909)
included in his list of all insects
found in the sand regions of the
Illinois and Mississippi river valleys.
Stal (1870) described C. mendica
from specimens obtained, possibly
by Belfrage (Geiser, 1933), from
Texas and Mexico. Probably em¬
ploying its own means of locomo¬
tion, it escaped the Great Plains
since 1850, moving down the east-
bound railroads where its food plant
was previously established. It is

common wherever M. nyctaginea oc¬
curs on trackways in Illinois north
from Quincy and Urbana, and since
Evers (1960) discovered the host
plant on railroads south from Madi¬
son county, the bug is probably ex¬
tending its range also in that direc¬
tion.

Celerio lineata (the white-lined
sphinx, Clemens, 1859). In the
years 1938-1941, I swept M. nyc¬
taginea extensively in Champaign
county while making a study of Ca¬
torhintha mendica (Balduf, 1942).
Larvae of C. lineata were taken on
only two occasions. Five almost full-
grown individuals, all about two
inches long, were obtained near To-
lono on October 5 and 11, 1941, and
other large ones near Mahomet, along
railroads in both cases. The identi¬
fying specialist commented that the
larvae are darker than usual. Hav¬
ing a long list of food plants of
divers taxonomic relations, this com¬
mon hornworm is a facultative mem¬
ber of the Mirahilis community.

Heliodines nyctaginella. This leaf-
skeletonizing micromoth was de¬
scribed by Gibson (1914) from
adults reared by Criddle at Aweme,
Manitoba, from larvae found feed¬
ing on M. nyctaginea, “a widely oc¬
curring representative of the Nyc-
taginaceae in Canada.” This brief
account shows that both the food
plant and the insect occur even in
the extreme northern part of the
Great Plains. I found it commonly
on railways in the northern half of
Illinois wherever its host occurred.
While investigating its life history,
Wester (1956) found it in 20 coun¬
ties across central Illinois. During
the seasons of bud clusters, most of
the eggs are laid on the outside of

Transactions Illinois Academy of Science

the involucres ; otherwise usually on
leaves near the top of the plant.
Excepting the first instars, the lar¬
vae live under webs spun over the
clusters of flower-buds and on the

leaves. Usuallv one larva inhabits

«/

a web. When it has consumed the
tissue, excepting the veins, enclosed
by its web, the larva emerges through
the opening left at the base of the
leaf, crawls down to the next leaf
and spins a second web where feed¬
ing is resumed. Such changes in
location and web-spinning continue
until the larva is full-grown, where¬
upon it apparently descends to de¬
bris on the ground for pupation.
Wester concluded that the chry-
palises of the last, i.e. fifth, genera¬
tion pass the winter.

Heliodines ionis (a stem borer).
Previous to the studies by AVester
(1954, 1956), only one species of
Heliodines, i.e. nyctaginella, was rec¬
ognized from ill. nyctaginea. He
discovered larvae similar to those of
H. nyctaginella boring in the stems.
Moths reared from such larvae by
AATester proved to be a new species,
which Clarke (1952) named H. ionis.
It was reported from Champaign,
Macoupin and Mason Counties, Illi¬
nois, and Wyandotte County, Kan¬
sas.

Although the adults are very simi¬
lar, the two species possess striking
differences in habits. H. nyctaginel¬
la attach their eggs to leaves and
clusters of buds, where the larvae
develop mostly under webs, then
pupate on the ground, and winter
as pupae. In contrast, H. ionis
oviposits in such secluded sites as
the paired axial buds, old exits from
which previous adults emerged from
the stem, and in natural splits in

stems. From these positions, the
small larvae enter the stems and de¬
velop by feeding on the succulent
pithy center. When almost full-
grown, the borer enlarges its tunnel
locally and packs its accumulated
excrement in both ends of the en¬
largement, thereby forming its pupal
chamber. But before transforming,
the larva lines the chamber with silk
and chews a hole through the stem
wall to provide for its subsequent
exit as an adult. AVester calculated
that H. nyctaginella, the leaf-skele-
tonizer, develops five generations in
a year as against only three for its
boring congener.

Onychobaris snbtonsa (a stem
borer). Like those of Heliodines
ionis, the larvae of 0. snbtonsa live
as stem borers. However, their ac¬
tivities in relation to the community
center differ sharply in several re¬
spects. The siphon of H. ionis limits
it to ingesting free fluids, if it feeds
at all, and, lacking a penetrating
ovipositor, this micromoth lays the
eggs on surfaces. In constrast, the
sturdy proboscis of the adult 0. snb-
tonsa, bearing mandibles at its apex,
serves both to feed and as an aid to
oviposition. Apparently both sexes
feed in the spring on incipient
leaves, clusters of flower buds, green
seeds and eventually only on the
stems (Wester, 1956a). The female
places the eggs only in the stems.
A pit is prepared with the proboscis
for reception of the egg; after the
egg is placed, the snout is employed
again, probably, in Wester’s opin¬
ion, to force the egg deeper. Plant
sap emanating from the wounded
tissue is said to envelope the egg and
congeal to form a protective cov¬
ering.

Insects Associated with Mirabilis in Illinois

The young larvae first feed in the
epidermis and the cortex of the stem,
then tunnel into the pith where they
feed until full-grown. Throughout
its development, the larva is white,
with head light brown, robust,
curved and legless. So they are
easily distinguished from the above
boring lepidopterous larvae. Their
last functions are (1) to prepare a
pupal cell in the tunnel, employing
excrement, and splinters of the
woody cortex glued together with
excretions from the malphighian
tubes, and (2) to chew an exit hole
through the cortex to, but not in¬
cluding, the epidermis. From this
opening the young adult emerges
the following spring. The full-
grown larva winters in the tunnel ;
pupation is delayed until April of
the next year. Only one generation
is completed annually.

I found Q. subtonsa to be common
wherever M. nyctaginea occurred in
the northern half of Illinois and
Wester collected it from 16 counties
in Central Illinois. LeConte (1876)
describes this species from a single
specimen taken in Texas by G. W.
Belfrage (see Geiser, 1933).

The Entomophagous Species of the
Community. The predatory bug,
Orius insidiosus and the six hy-
menopterous parasites, reported by
Wester (1956b), occurred in small
numbers compared with their phy¬
tophagous hosts alread}^ described
above. Both Orius and the para¬
sites are facultative in this communi¬
ty since they have previously been
shown to utilize also other species of
prey and hosts living outside the
Mirabilis center. (Muesebeck et al,
1951.) This means they have prob¬
ably entered the Mirabilis communi¬

ty from others, where they attacked
alternate hosts.

The parasitic forms are equipped
with exserted or exsertile oviposi¬
tors which enable them to implant
their eggs upon or into the host in¬
visible beneath a web or in a tunnel
in the stem of Mirabilis. Also a high
level of sensitivity must be presumed
to account for this ability. Previous
to oviposition, certain of these para¬
sites inject a venom via the hollow
terebra into the host, thus paralyzing
it temporarily or permanently, par¬
ticularly in case of ectoparasites.
Wester found that five of the six
Hymenoptera are solitary and ecto-
parasitic on host larvae, i.e. a single
larva feeds and grows to maturity
from the outer surface of a given
individual host. Only Tetrastichus
coerulescens deviates from that pat¬
tern of relationship. Here numbers
of the larvae develop, all internally.
Moreover, the host is not a larva,
but the pupa or chrysalis of Heli-
odines ionis. Such are described as
gregarious endoparasites.

The host relations and habits of
the six species, as reported by Wester
(1956b), are briefly summarized
herewith.

I. Braconidae.

1. Bracon caulicola, solitary ecto¬
parasite on the boring larvae of Heli-
odines ionis , micromoth, and Ony-
chobaris subtonsa, snout beetles, in
tunnels of Mirabilis stems. Host lar¬
vae suffer permanent paralysis. Pu¬
pation occurs in a cocoon in the host
tunnel, and the new adult chews a
hole in the tunnel wall for escape.

2. Bracon gelechiae. In all bi-
onomic respects like B. caulicola,
but attacks the advanced larvae of

Transactions Illinois Academy of Science

Heliodines nyctaginella, which live
more accessibly under webs on leaves.

II. Eulophidae.

3. Tetrastichus coerulescens, a
gregarious endoparasite in pupae of
H. ionis in tunnel of Mirabilis stem.
Wester believed the adult female en¬
ters the host tunnel via the hole
chewed in wall of stem tunnel by
the host larva, and there parasitizes
the mature host. He obtained from
10 to 40 adult coerulescens per host
pupa.

III. Eupelmidae.

4. Eupelmus allynii, a solitary
ectoparasite on the boring larvae of
Heliodines ionis in burrow in stem
of Mirabilis. The host larva is per¬
manently paralyzed. The parasite
winters as a mature larva in the
host’s burrow, where it also pupates
in the spring. The new adults
emerge through holes which they
chew in the plant stem.

5. Eupelmus cyaniceps, a solitary
ectoparasite of H. ionis. Its relations
to the host, also its life cycle, are in
all respects as stated above for E.
allynii.

IV. Pteromalidae.

6. Neocatolaccus tylodermae , a
solitary ectoparasite on larvae of the
black snout beetle, Onychobaris sub-
tonsa, in burrow in stem of Mirabilis.
All hosts previously reported for
this species are larvae of snout
beetles (Muesebeek, 1951).

7. Orius insidiosus. When per¬
forming in the Mirabilis community,
this small predatory bug attacked

chieflv the first instars of Heliodines
*/

nyctaginella, which do not spin a
protective web. Both nymphs and

adults engaged in the attack. Wester
observed them circle around a pro¬
spective larva several times, before
they rammed it with the extended
proboscis and started to feed. Many
predatory Hemiptera first employ
the forelegs to capture their victims.

Acknowlegments

I am indebted to Dr. Clifford
Wester for basic bionomic data on
several species concerned in this ar¬
ticle. See References Cited.

References Cited

Balduf, W. V. 1942. Bionomics of Ca-
torhintha mendica St&l. Bui. Brook¬
lyn Ent. Soc., 37: 158-166.

Balduf, W. V. 1957. The spread of
Catorhintha inendica Stal. Proc. Ent.
Soc. Wash., 59: 176-185.

Clarke, J. F. G. 1952. A new Heliodinid
from Illinois. Proc. Ent. Soc. Wash.,
54 (3): 138-139.

Clemens, J. B. 1859. Synopsis of North
American Sphingidae. Jour. Acad.
Nat. Sci. Phila., 4 (ser. 2): 145.

Evers, R. A. and J. S. Winterringer.
1960. New records of Illinois vascular
plants. Illinois State Museum, Spring-
field, Scientific Papers, Vol. 11, pp. 135.
Geiser, S. W. 1933. G. W. Belfrage’s
Texas localities. Ent. News, 44: 127-
138.

Gibson, A. 1914. A new Elachistid
moth from Manitoba. Can. Ent. 46:
423-424.

Hart, C. A. 1907. Zoological studies in
the sand regions of the Illinois and
Mississippi river valleys. Bui. Ill. St.
Lab. Nat. Hist., 7: 195-272.

Jones, G. N. and D. G. Fuller. 1955.
Vascular plants of Illinois. Universi¬
ty of Illinois Press, pp. 593.

LeConte, J. L. 1876. The Rhynchophora
of America north of Mexico. Proc.
Amer. Phil. Soc., 15: 29.

Muesebeck, C. F. W., C. V. Krombein
and H. K. Townes. 1951. Hymenop-
tera of America north of Mexico.
Synoptic Catalog, U.S.D.A., Agr. Mono¬
graph No. 2, 1420 pp.

StAl. C. 1870. Enumeratio Hemip-
terorum. Svenska Vetensk. Akad.
Handl., p. 187.

Insects Associated with Mirabilis in Illinois

Tietz, H. M. 1952 The Lepidoptera of
Pennsylvania. Penn State College,

Agr. Exp. Sta., 194 pp.

Wester, C. 1954. Mirabilis nyctaginea
(Michaux) MacMillan and certain in¬
sects as a microcommunity. Ph.D.
thesis, University of Illinois, Urbana.

Wester, C. 1956. Comparative bionomics
of two species of Heliodines on Mi¬

rabilis. Proc. Ent. Soc. Wash., 58:
43-46.

Wester, C. 1956a. Notes on the bionom¬
ics of Onychobaris subtonsa LeConte.
Proc. Ent. Soc. Wash., 58: 105-108.

Wester, C. 1956b. Notes on the bionom¬
ics of the natural enemies of the in¬
sects on Mirabilis. Proc. Ent. Soc.
Wash., 58: 283-286.

Manuscript received April 30. 1961.

THE EFFECTS OF RED AND FAR-RED IRRADIATION
ON THE VEGETATIVE DEVELOPMENT OF
PEA AND COCKLEBUR

ROBERT H. KUPELIAN

University of Chicago

The vegetative growth of many
plants is regulated significantly by
light quality and intensity. Several
responses are controlled by light
quality; these include seed germina¬
tion, pigment formation, unfolding
of the plumular hook, elongation of
stems, and leaf expansion. Recent
studies have shown that the basic
underlying photoreaction involved
in each case is the same (Borthwick
and Hendricks, 1960 ; Butler and
Downs, 1960 ; Downs, 1959 ; Hend¬
ricks, 1959; Meijer, 1959). Light is
absorbed by a blue-green pigment,
phytochrome, which appears to exist
in two forms. The photochemical
reaction may be written as given
below.

Pigment RH^ + A

6500-6600 A max
- ^

7200-7400 A max

Pigment PR -(- AH2

In this formula. R and FR refer to
the red and far-red absorbing pig¬
ment forms, respectively (Hend¬
ricks, 1959). It has been shown that
the effects of red radiation can be
markedly reduced or even entirely
eliminated in some cases by a sub¬
sequent exposure to wavelengths in
the far-red portion of the spectrum
(Hendricks, 1959 ; Meijer, 1959 ; Van
der Veen and Meijer, 1959).

The present investigation was un¬
dertaken to examine the effects of
red and far-red light on certain as¬
pects of the vegetative development
of garden pea and cocklebur.

Materials and Methods

Cultural procedure. Seeds of gar¬
den pea ( Pisum sativum L.), variety
Thomas Laxton, were soaked for 19
hours in flowing tap water at
8-10° C and then sown approximately
4 cm apart in unglazed clay pots or
plastic flats containing vermiculite,
thoroughly wetted with tap water.
At the end of a germination period
of 72 hours in a dark controlled-
environment room at 21 °C, approxi¬
mately 50% of the seedlings had
emerged. At this time the containers
were divided into groups, each con¬
taining a minimum of twenty uni-
form tagged seedlings, and subjected
to the various light treatments. The
remaining plants in the containers
were permitted to grow with the
tagged seedlings but were not used
in the experiment.

Achenes of cocklebur ( Xanthium
pensylvanicum Wallr.) were germi¬
nated by the procedure described by
Vergara and Me II rath (1960) and
planted approximately 8 cm apart
in quartz sand in plastic flats. The
seedlings were grown under non-
inductive conditions of 20 hours of
light per day in a controlled-environ-

[48]

Effects of Irradiation on Plant Growth

ment room at 70±2°F and received
approximately 2000 ft-c. of light
from General Electric Power Groove
fluorescent lamps supplemented with
60-watt incandescent bulbs (approxi¬
mately 12% of the total wattage).
When two nodes became visible, the
plants were subjected to the various
light treatments. Six plants con¬
stituted a series in each treatment.

Both cocklebur and pea plants
were watered three times per week
during the course of the experiment,
the former with a complete nutrient
solution (Hoagland and Arnon,
1950) and the latter with distilled
water.

Irradiation procedure. All seed¬
lings were exposed to an 11 -hour
photoperiod between 9 a.m. and 8

p.m. daily. In the six treatment
series plants were exposed to various
light qualities for the following num¬
ber of hours : Group A — 11 incan¬
descent (I) ; Group B — 91 and 2
far-red (FR) ; Group C — 91 and 2
red (R) ; Group D — 11 fluorescent
(F) ; Group E — 9F and 2FR ; Group
F — 9F and 2R. A temperature of
21±2°C and a relative humidity of
60-75% were maintained. When
red or far-red light constituted a
portion of the treatment, it was al¬
ways given during the final 2 hours
of the light period.

The lamps and filters utilized to
produce each type of light regime
are indicated in Table 1. The filters

were similar to those described bv

•/

other workers (Liverman, 1959 ; Na-

Table 1. — Light Sources and Energies Utilized in the Various Experiments.

Light Energies, [jlW /cm2

Experiment

Irradiation

Lamp

Filters

II-III

Incandescent .

1000- watt
incandescent,

General Electric,

RB 52 (I)*

None

1565

3700

Fluorescent .

40-watt warm white
fluorescent,

General Electric (F) *

None

1132

3700

Red .

40- watt warm white
fluorescent,

General Electric

Two layers
of red
cellophane

1827

3700

Far-red .

1000- watt
incandescent,

General Electric

RB 52

Two layers
of red and
two layers
of blue
cellophane

1152

3700

* Percentage spectral energy distribution: 4000-5000 A — I 10.1, F 19.2:
5000-6000 A — I 24.1, F 48.8; 6000-7600 A — I 65.7, F 32.0.

Transactions Illinois Academy of Science

kayama, et al., 1960). The radiant
energies for the various types of
light were determined with a Weston
Illumination Meter (type 756) which
had been previously calibrated with
a thermopile for each type of irra¬
diation (Van der Veen and Meijer,
1959). The Weston meter readings
were taken at the level of the plants.
The spectral energy distribution of
the visible light from the incan¬
descent and fluorescent lamps (Ta¬
ble 1) was taken from the tables
presented by Weitz (1956) .

Harvest procedure. The pea seed¬
lings were permitted to grow under
the light conditions described for 14
days, at which time the morphologi¬
cal age of the plants in tenths of
nodes was determined using a modi¬
fication of the method of Higgins
(1952). This technique is based on
scoring the plant ’s morphological age
in terms of number of nodes pro¬
duced and it permits stages beyond
the last discernible node to be desig¬
nated in tenths of nodes (Figs. 1
and 2 ) .

After the morphological ages of
the plants in each group had been
determined, the shoots were cut at
the level of the cotyledons, and the
lengths were recorded to the nearest
millimeter of the third internode.
The widths of the stipules and leaf¬
lets at node five were also deter¬
mined. The total surface area of
the leaflets and stipules at node five
was measured with an Aminco leaf
area meter.

The length of the second inter¬
node of cocklebur was measured to
the nearest millimeter. These plants

were irradiated simultaneouslv with

the pea seedlings.

Statistical analyses were carried

out according to established proce¬
dures (Snedecor, 1946).

Results and Discussion

Influence of Quantity and Quality
of Light on Internodal Lengths of
Pea anid Cocklebur. Internodal
lengths were greatest when irradia¬
tion with either incandescent or
fluorescent lamps was followed by
far-red radiation (Table 2).

With respect to pea, it is of inter¬
est that the multifold increase in
light energy during Experiment II,
compared with that in Experiment
I, had a pronounced effect in the
fluorescent series, groups D-F, and
a slight but statistically insignifi¬
cant effect in the incandescent series,
groups A-C, (Table 2).

Fig. 1. — Garden pea, variety Thomas
Laxton. Nodes are numbered; terminal
growing point located between stipules
at node 9.

Effects of Irradiation on Plant Growth 51

Fig. 2. — Stages in leaf development of garden pea, variety Thomas Laxton.
(1.0) illustrates a completed node at the tip of the plant at which point occurs
a mature leaf composed of two leaflets, two stipules, and a tendril. Between the
pair of stipules is an immature stem which supports a small, tightly closed leaf
bud; (0.1) bud develops, increases in size, and the tendril unfolds from between
the pair of stipules; (0.2) leaflets, closely held together, begin to show between the
stipules; (0.3) leaflets and tendril elongate; (0.4) leaflets separate, both leaflets
and tendril elongate; (0.5) separated leaflets and tendril continue to elongate;
(0.6) leaflets begin to separate from the stipule; (0.7) leaflets become completely
separated from the stipules; (0.8) leaflets begin to unfold, become separated further
from the tightly closed stipules; (0.9) leaflets unfold completely, stipule begins
to unfold; and (1.0) leaflets and stipules have attained maximum expansion and
between the stipules is a tightly closed leaf bud.

The greater energy from the flu¬
orescent light source in Experiment
II, as compared with Experiment
I, resulted in internode lengths which
where 51, 27, and 34 per cent less
for plant groups D, E, and F, re¬
spectively. The difference in inter-
nodal growth of peas at the different
light energies under fluorescent and
incandescent lamps can probably be
explained on the basis of the wave¬
lengths of light emitted by these
sources. Fluorescent lamps, with
high red and almost no far-red emis¬

sion, would be expected to maintain
the phytochrome system predomi¬
nantly in the far-red absorbing form
and hence less elongation should re¬
sult (Downs, 1959). In Experiment
I, however, internodes were as long
or longer in plants grown under flu¬
orescent lamps as in those grown
under an incandescent source (Ta¬
ble 2). Apparently the quantity of
red light for groups D-F in Experi¬
ment I was not sufficient to maintain
enough of the pigment in the far-
red form to limit elongation. In

Transactions Illinois Academy of Science

Experiment II, however, in groups
D-F the increased energy from the
fluorescent lamps increased the rela¬
tive amount of red (and also blue)
light received by the plants without
any appreciable increment in the
far-red, thus maintaining enough
pigment in the far-red absorbing
form to cause less growth of the in¬
ternodes. The increased energy in
the blue wavelengths may have also
been important since Wassink and
Stolwijk (1956) have demonstrated
that at high energies, in the order
of 3700 fxW/cm.2, blue light is very
active in inhibiting elongation. In
Experiment I, plants subjected onl}"
to light from a fluorescent source
(group D) were appreciably shorter
than those (group F) given such
light plus a supplementary treat¬
ment of red (Table 2). Incandescent
lamps produce considerable red as
well as far-red light, and the in¬
creased energy from this source in
Experiment II for groups A-C did
not change the ratio of these two
light qualities. Thus one would not

expect to get appreciably great¬
er internodal elongation with in¬
creased energy from this source, as¬
suming that the initial light energy
was not seriously limiting other proc¬
esses required to sustain growth,
such as photosynthesis.

That the pea plants received an
appreciable quantity of red light
from the incandescent source was
apparent from the fact that no sig¬
nificant difference was found in the
internodal lengths of plants grown
under incandescent lamps with or
without supplementary treatment
with red light (groups A and C, Ex¬
periments I and II) ; plants receiving
supplementary treatment of far-red
(group B) were, however, signifi¬
cantly different from those (group
A) exposed only to light from in¬
candescent lamps (Table 2).

Although no significant differences
were found among the internodal
lengths of the cocklebur plants ex¬
posed to the various incandescent
light treatments (groups A-C, Ex¬
periment III), the values are in the

Table 2. — Effect of Light Treatment on the Length of Internodes of
Pea and Cocklebur Plants.

Plant Group

Treatment

Pea

Internode 3, mm

Cocklebur
Internode 2, mm

Expt. I

Expt. II

Expt. Ill

A .

49.2 ± 1.3

53.3 ± 1.6

111.7 ± 1.7

B .

I + FR

57.5 ± 1.7

58.5 + 1.9

112.5 ± 2.9

C .

I + R

52.1 ± 1.8

53.6 ± 1.7

100.2 ± 5.4

D .

59.1 ± 1.9

29.1 + 1.2

26.0 ± 1.2

E .

F + FR

65.7 ± 2.1

48.2 ± 1.0

87.3 + 1.8

F .

F + It

62.8 ± 1.9

41.5 ± 1.4

24.3 ± 0.4

Effects of Irradiation on Plant Growth

sequence of magnitude to be pre¬
dicted if internodal elongation in
tli is species were a red - far-red con¬
trolled response (Table 2). The
failure of cocklebur plants to show
significant differences among the
various incandescent treatments, as
contrasted with pea, is indicative of
the variability among species in re¬
sponding to a given light treatment.

The difference in internodal
lengths of cocklebur plants (Experi¬
ment III) grown under fluorescent
(group D) and incandescent (group
A) lamps was found to be highly
significant (Table 2) ; internodal
elongation was drastically curtailed
bv fluorescent illumination. This
inhibition by light from fluorescent
lamps, however, was partially over¬
come by exposure to far-red light ; a
highly significant difference was evi¬
dent between group D or F and
group E. A supplementary treat¬
ment of red light caused a slight but
insignificantly greater inhibition of
internodal elongation than light from
fluorescent lamps alone (Table 2).

Downs (1959), working with
loblolly pine, soybean, and tomato,
found that longer internodes were
produced when the plants entered
the dark period with the pigment
system predominantly in the red ab¬
sorbing form which would occur un¬
der incandescent supplemental light.
Additional work with several vari¬
eties of beans further supported the
general statement that a brief ex¬
posure to far-red radiation before
the beginning of the dark period
promoted internodal elongation. Mei-
jer (1959) observed that far-red
(near infrared) stimulated elonga¬
tion in Petunia, Calendula, Perilla,
Helianthus, bean, and tomato plants.

The results with pea and cocklebur
plants were consistent with the ob¬
servations of these workers.

Influence of Quantity and Quality
of Light Upon the Morphological
Age of Pea Plants. An acceleration
of the morphological aging of pea
plants resulted from the greater
light energy used in Experiment II
(Table 3). In the incandescent
series, groups A-C, the morphologi¬
cal age was 3.6, 12.0, and 1.8 per
cent greater, respectively, in Ex¬
periment II than in Experiment I.
Greater maturity in the fluorescent
groups, D-F, was even more pro¬
nounced, the values showing a 10.5,
18.0, and 8.6 per cent increase over
those of Experiment I.

Supplemental far-red irradiation
not only resulted in longer inter¬
nodes in pea plants but also de¬
pressed the rate of node initiation.
This was true whether this light
quality followed illumination from
incandescent or fluorescent lamps
(Table 3). It was slightly more ef¬
fective, however, when given after
light treatment from a fluorescent
source. Although supplemental treat¬
ment with red light appeared to
have a slight influence on the rate
of node initiation, statistically it
proved to be insignificant.

In the discussion of the influence
of fluorescent illumination on inter¬
nodal length it was pointed out that
perhaps the blue light from this
source was of importance, for plants
grown under fluorescent light only
were significantly shorter than plants
receiving fluorescent plus a supple¬
mental red-light treatment. With
respect to morphological age, how¬
ever, no significant difference existed
between these two treatments, indi-

54 Transactions Illinois Academy of Science

Table 3. — Morphological Age of Pea Plants Expressed as Mean Node Number.

Average number of nodes

Plant Group

Treatment

Experiment I

Experiment II

A .

5.5 ± 0.07

5.7 ± 0.03

B .

I + FR

5.0 ± 0.07

5.5 ± 0.03

C .

I + R

5.6 ± 0.09

5.7 ± 0.03

D .

5.7 ± 0.06

6.3 ±0.10

E .

F + FR

5.0 ± 0.08

5.9 ± 0.06

F .

F + R

5.8 ± 0.11

6.3 ± 0.10

Table 4. — Average Widths of Leaflets and Stipules and Average Area of
Leaflets and Stipules at Node Five.

Plant Group

Treatment*

Stipule
Width, mm

Leaflet
Width, mm

Area of leaflets
and stipules, cm2

A .

10.4 ± 0.4

16.4 ± 0.5

10.4

B .

I + FR

9.5 ± 0.2

13.6 ± 0.7

8.4

C .

I + R

11.4 ± 0.3

16.9 ± 0.7

10.8

D .

13.6 ± 0.4

17.2 ± 0.6

12.0

E .

F -f FR

12.3 ± 0.4

16.9 ± 0.8

11.7

F .

F + R

15.1 ± 0.5

19.6 ± 0.5

12.4

* Plants received light energies of 3700 ^W/cm^.

eating that blue light was not a de¬
termining factor in the rate of node
production.

Influence of Light Quality on the
Growth of Leaflets and Stipules of
Pea. Although in general the widths
of the stipules on plants of the flu¬
orescent series tended to be greater
than those on plants in the incan¬
descent group, the stipules of plants
of group D were not significantly
wider than those of group C (Fig.

3; Table 4). Stipule enlargement
as a consequence of supplementary
irradiation with red light was quite
apparent. It was likewise obvious
that supplementary irradiation with
far-red light inhibited stipule ex¬
pansion (Table 4) .

Red light given subsequent to in¬
candescent illumination appeared to
be of little consequence in leaflet ex¬
pansion, since such supplementary
treatment did not result in a signifi-

Effects of Irradiation on Plant Growth

Fig. 3. — Leaf and stipule size at node five. From left to right: top row, groups
A through C; bottom row, groups D through F.

eantly greater leaflet width than
that which occurred under incan¬
descent lamps (Table 4). A signifi¬
cantly increased leaflet width did
result from red light treatment in
the fluorescent series, however. With
far-red supplementary illumination,
on the other hand, significantly nar¬
rower leaflets were produced in the
incandescent series while no signifi¬
cant effect could be found in the
fluorescent group. The reasons are
obscure for this apparent red-light
effect in the fluorescent group but
not in the incandescent series, and
also for the far-recl effect in the in¬
candescent group but not in the flu¬
orescent series. It is interesting that
the stipules responded differently
than did the leaflets to the various
light treatments.

The area of the stipules and leaf¬
lets combined did not show the clear
cut relationships exhibited when
these organs were considered individ¬
ually (Fig. 3). This is to be ex¬

pected in view of the differential
responses of these organs to the vari¬
ous light treatments. The general
tendency was apparent, however, for
the stipules and leaflets to have a
greater area when produced under
fluorescent rather than under in¬
candescent lamps ; under each of
these types of illumination these or¬
gans exhibited a greater area with
red light treatment and a smaller
area with far-red illumination.

These results appear to be con¬
sistent with the observations of other
workers (Liverman, 1959; Parker
et at. , 1949; Went, 1941).

Summary

Common garden pea {Pi sum sa¬
tivum L.), var. Thomas Laxton, and
cocklebur (Xanthium pensylvanicum
Wallr.) plants were grown under
various light treatments in 11-hour
photoperiods. These included 11
hours of incandescent or warm white

Transactions Illinois Academy of Science

fluorescent, and 9 hours of incan¬
descent or fluorescent followed by a
supplementary illumination of 2
hours of red or far-red irradiation.
The plants were harvested after a
2-week exposure to the various light
treatments. The criteria selected as
indices of vegetative growth in pea
were internodal elongation, mor¬
phological age of plants measured
in tenths of nodes, width of stipules
and leaflets, and total area of leaf¬
lets and stipules. Only internodal
elongation was measured in cockle-
bur.

Internodes were generally shorter
on pea and cocklebur plants illumi¬
nated with fluorescent lamps only.
Exposure to red light following in¬
candescent or fluorescent illumina¬
tion did not result in significantly
shorter internodes, but far-red light
following such illumination resulted
in significantly longer internodes.

The morphological age of pea
plants was greatest for plants grown
under fluorescent illumination. Ex¬
posure to supplemental red light did
not result in a significant increase in
morphological age following either
fluorescent or incandescent illumina¬
tion. Far-red light, however, de¬
pressed the rate of node initiation
significantly under both conditions.

The widths of stipules on plants
illuminated with incandescent or
fluorescent light alone were signifi¬
cantly different from those given
subsequent treatment with either
red or far-red light.

The widths of leaflets on plants
under incandescent illumination
were significantly different from
those receiving supplemental treat¬
ment with far-red light but not from
those receiving red light. For plants

receiving fluorescent illumination,
the reverse was true.

Acknowledgments

The author wishes to express his
gratitude to Dr. W. J. Mcllrath for
his valuable counsel and interest
throughout the course of this inves¬
tigation. Appreciation is also ex¬
pressed for fellowship support re¬
ceived through the Charles L. and
Francis K. Hutchinson Fellowship
of the Department of Botany, Uni¬
versity of Chicago and a National
Science Foundation Summer Fellow¬
ship.

This investigation was supported
in part by grants from the National
Science Foundation (G-4018) and
from the Dr. Wallace C. and Clara
A. Abbott Memorial Fund of the
University of Chicago.

Literature Cited

Borthwick, H. A. and S. B. Hendricks.
1960. Photoperiodism in plants. Sci¬
ence, 132: 1223-1228.

Butler. W. L. and R. J. Downs. 1960.
Light and plant development. Sci¬
entific American, 203(6): 56-63.

Downs, R. J. 1959. Photocontrol of
vegetative growth. In: Photoperiod-
ism and Related Phenomena in Plants
and Animals. R. B. Withrow, editor,
Amer. Assoc. Adv. Sci. Pub. No. 55.
Wash. D. C., pp. 129-135.

Hendricks, S. B. 1959. The photoreac¬
tion and associated changes of plant
photomorphogenesis. In: Photoperi-
odism and Related Phenomena in
Plants and Animals. R. B. Withrow,
editor, Amer. Assoc. Adv. Sci. Pub.
No. 55, Wash. D. C., pp. 423-437.
Higgins, J. J. 1952. Instructions for
making plienological observations of
garden peas. Publications in Clima¬
tology. V(2). The Johns Hopkins
University Press.

Hoagland, D. R. and D. I. Arnon. 1950.
The water-culture method for growing
plants without soil. Calif. Agr. Exp.
Sta. Circ., 347.

Effects of Irradiation on Plant Growth

o i

Liyerman, J. L. 1959. Control of leaf
growth by an interaction of chemicals
and light. In: Photoperiodism and
Related Phenomena in Plants and Ani¬
mals. R. B. Withrow, editor, Amer.
Assoc. Adv. Sci. Pub. No. 55, Wash.
D. C., pp. 161-180.

Meijer, G. 1959. The spectral depend¬
ence of flowering and elongation. Acta.
Bot. Neerl., 8: 189-246.

Nakayama, S., H. A. Borthwick and
S. B. Hendricks. 1960. Failure of
photoreversible control of flowering in
Pharbitis nil. Bot. Gaz., 121 : 237-243.

Parker. M. W., S. B. Hendricks, H. A.
Borthwick and F. W. Went. 1949.
Spectral sensitivities for leaf and stem
growth of etiolated pea seedlings and
their similarity to action spectra for
photoperiodism. Am. J. Bot., 36: 194-
204.

Snedecor, G. W. 1946. Statistical Meth¬
ods. 4th ed., Iowa State College Press,
Ames, 485 pp.

Van der Veen, R. and G. Meijer. 1959.
Light and Plant Growth. Macmillan
Co., New York, 161 pp.

Vergara, B. S. and W. J. McIlrath.
1960. Influence of photoperiod on wa¬
ter uptake of Xanthium. Bot. Gaz.,
122: 96-102.

Wassink, E. C. and J. A. J. Stolwijk.
1956. Effects of light quality on plant
growth. Ann Rev. Plant Physiol., 7:
373-400.

Weitz, C. E. 1956. G. E. Lamps LD-1.
General Electric, Cleveland, Ohio, 79

pp.

Went, F. W. 1941. Effects of light on
stem and leaf growth. Am. J. Bot.,
28: 83-95.

Went, F. W. 1957. Experimental Con¬
trol of Plant Growth. Chronica Bo-
tanica, Waltham, Mass., 340 pp.

Manuscript received January 30, 1962.

SOME COMPARATIVE ASPECTS OF ORGAN
WEIGHTS IN CANADA GEESE ( BRANT A
CANADENSIS INTERIOR)1

HAROLD C. HANSON

Illinois State Natural History Survey , Urbana

Physiological studies of the Cana¬
da geese wintering at Horseshoe
Lake, Alexander County, Illinois,
were initiated in 1954 with a study
of organ weights. A series of papers
on condition factors (in press), blood
chemistry, tissue chemistry, the en¬
docrine organs, nasal glands, and
histology of the long bones of the
leg are currently under preparation.
For background information on the
population of geese under discus¬
sion, the reader is referred to Han¬
son and Smith (1950).

Materials and Methods

Organ weights were obtained from
142 geese. The majority of the or¬
gans weighed were from geese shot
by hunters between dawn and 10.00
A.M. during November and early
December.

Prior to weighing the heart, all
major blood vessels were excised as
completely as possible, and the
chambers were cut open and all
blood was removed. Spleens were
merely freed of excess fascia. The
proventriculus was left attached to
the gizzard because its function is
so intimately related to that of the
gizzard and because the juncture of
the proventriculus with the esopha¬
gus provided a clear-cut margin that
could be consistently followed in dis¬
section. All excess fascia and fat

1 Based on part of a thesis submitted
for the Ph.D. degree at the University
of Illinois, 1958.

attached to the exterior of these
organs were removed. They were
cut open, rinsed clean of contents,
and then excess water was removed
by blotting with absorbent paper.
Before weighing the liver, the gall
bladder was removed, as well as all
clotted blood present in the exposed
sections of the major veins. The
lobes of the pancreas were stripped
from the intestine and freed of fat.
All weighings were made to one-
tenth gram on a triple-beam balance.
Age and sex classes of geese were
distinguished according to criteria
previously presented by Hanson
(1949). Immature geese were 5-8
months old, yearlings 17-20 months,
and adults 29 or more months of
age at the time collections were
made. Dr. Horace W. Norton of
the College of Agriculture, Univer¬
sity of Illinois was consulted on the
statistical analyses.

Results

Heart W eights. Weights were ob¬
tained of hearts of 142 Canada geese
at Horseshoe Lake in late autumn
(Table 1). In both sexes a signifi¬
cant absolute increase in heart
weight with age is indicated but, as
body weight is also higher for each
successive age class, there does not
appear to be a significant increase
in heart weight relative to body
weight of age classes except in males

[58]

Organ Weights in Canada Geese

{Table 1). In respect to the correla¬
tion of heart weight with body
weight, the age-sex class samples
proved to be homogeneous ; the
pooled correlation coefficient was .65
(P = <.01). The correlation of
heart weight with body weight in
each sex-age class was found to be
highly significant. Coefficients of
variation for heart weights were also
the lowest recorded for the five vis¬
ceral organs studied.

Quiring and Bade (1943) also
found the highest correlation of
heart-body weight of any organ
studied in the house sparrow ( Pas¬
ser domesticns) . Marsden (1940)
found a similarly high correlation
for domestic turkeys. Latimer
(1927), studying the turkey hen,
also found heart weights to have the
highest correlation with body weight
of the visceral organs. Souba’s
(1923) data showed a slightly high¬
er correlation of liver weight to body
weight than heart weight to bod}"
weight in 100 white leghorn cock¬
erels.

The heart weight of the Canada
geese studied averaged 0.90 to 0.94
per cent of average body weight.
In chickens, heart weight was 0.55
per cent of body weight ; in turkeys,
0.53 per cent (Latimer and Rosen¬
baum, 1926). In white Pekin ducks,
heart weight was 0.64 to 0.66 per
cent of the body weight (Salgues,
1939). Sixteen passerine species
studied by Renscli (1948) had hearts
that averaged 0.82 per cent of body
weight. Presumably the relatively
small heart of domestic chickens,
turkeys, and ducks reflects artificial
selection and/or their sedentary ex¬
istence.

Hartman (1955) has reviewed

heart weight studies in wild birds
and presented much new data for
a wide variety of birds. Heart
weights varied from 0.2 to 2.4 per
cent of body weight. Wide varia¬
tion was found in the same families
of birds. No significant differences
were found to be present between
age-sex classes, but activity was be¬
lieved to have an important influ¬
ence on heart size between species.
Mitchell, et al. (1926) reported that
heart weights in white plymouth
rock chickens were higher in cock¬
erels than in both pullets and capons,
a difference they attributed to the
greater muscular activity of the
cockerels. However, it is more like¬
ly that these differences, as in the
case of pectoral muscles in Canada
geese, can be attributed to the ni¬
trogen-conserving effect of the male
sex hormones (Hanson, 1961). The
relative weight of the heart in Cana¬
da geese does not change significant¬
ly with age in females despite re¬
peated migrations, but consistently
is relatively greater in the older age
classes of males than in immatures
(Table 1).

Heart weights of white leghorn
chickens were also greater relative
to body weight in cockerels than in
pullets (Mitchell, et al ., 1931). Rid¬
dle (1947) reported that male ring¬
doves ( Strejjtopelia decaocto) have
a slightly larger heart than females,
but failed to find a sex difference
in the domestic pigeon. Kirkpatrick
(1944) found that in both sexes of
the ring-necked pheasant ( Phasianus
colchicus ) the weight of the heart
increased with age.

Smith (1928) concluded that in
humans the relative weight of the
heart does not increase with age,

Transactions Illinois Academy of Science

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Table 2. — Spleen weights and their relation to body weights in Canada geese
at Horseshoe Lake, Illinois in late autumn.

Organ Weights in Canada Geese

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irrespective of the weight of the
body. The weight of the human
heart does, however, increase with
increases in weight of the body. In
males, the heart is 0.43 per cent of
the body weight ; in females, 0.40
per cent of the body weight. This
latter difference might also be at¬
tributable to the differential effect
of the sex hormones.

Spleen Weights. The spleens of
137 Canada geese were weighed (Ta¬
ble 2). The spleen in Canada geese
tends to decrease in size with age.
In immatures and yearlings there
is no difference between sexes in the
relative size of the spleen ; in the
adult age class, the males have a
markedly larger spleen, both ab¬
solutely and relatively. The dif¬
ferentially larger spleens in adult
males may reflect an accommodation
to the vascular requirements of the
more highly developed musculature
system of this age-sex class. While
size of spleen is not directly related
to the red cell count of the blood
and hemoglobin level, it is interesting
to note that the sex differential in
size of the spleen in the adults also
corresponds with higher hematocrit
and hemoglobin values found for
adult males (Hanson, 1958). Har¬
mon, et al. (1932) have demonstrated
that the spleen of birds apparently
can function as a blood reservoir.

The weight of the spleen is ex¬
tremely variable in Canada geese
(Table 2). In nearly all studies of
organ weights in chickens and tur¬
keys, the spleen was reported to be
the most variable in weight of the
visceral organs. Because of the blood
reservoir function of the spleen, the
variability of spleen weight in Cana¬
da geese may reflect the circum¬

stance of death rather than an ex¬
treme variability in the mass of the
organ tissue per se. Geese killed in¬
stantaneously may be expected to
have a spleen containing more blood
than one shot in such a way that its
death was in large measure due to
loss of blood — whether bleeding oc¬
curred internally or externally.
Spleen weight in Canada geese
tended to be significantly correlated
with body weight in males (im-
matures, P = .01 ; adults, P = .1 -
.05) ; in females no significant cor¬
relation was found.

Mitchell, et al (1926, p. 126) re¬
ported a sex differential in the rela¬
tive weight of the spleen of white
Plymouth rock chickens reverse to
that found in Canada geese : ‘ ‘ The

weights of spleen were consistently
heavier for pullets than for cock¬
erels. ’ ’ A similar relationship is not
evident in their data for white leg¬
horn chickens (Mitchell, et al, 1931).

A highly significant increase in
the weight of the spleen in male ring¬
necked pheasants from 87 to 172
days of age was observed by Kirk¬
patrick (1944). Spleen weights in
captive ringdoves are three times as
large in spring and summer as in
winter and autumn and the spleens
of males are larger than those of
females (Riddle, 1929). Opposite-
seasonal trends in the size of the
spleen in ring-necked pheasants
have been reported by Kirkpatrick
(1944). Male white-crowned spar¬
rows (Zonotrichia leucophrys gam-
helii) have spleens that average
larger, absolutely and relatively,
than those of the females (Oakeson.
1953). Males of this subspecies at¬
tained their lowest average spleen
weights in May upon their arrival

Organ Weights in Canada Geese

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( )rgan Weights

in Alaska (Oakeson, 1953). A com¬
parable seasonal trend in spleen
weights for a non-migratorv race of
the white-crowned sparrow (Zono-
trichia leucophrys mitt alii) was also
reported by Oakeson (1956).

Weights of Proventricnlus and
Gizzard. The weights of gizzards
with the proventricnlus attached
were obtained from 103 Canada
geese (Table 3). Second to the heart,
the combined weight of these organs
was the least variable of those
studied. The correlation of the
weights of these organs to body
weight tended to increase with age,
but only in adult females did rela¬
tionships approach the significant
level (P = .1) .

As the proventricnlus accounts for
a relatively small per cent of the
combined weight of this organ and
the gizzard in Canada geese, it is
evident from their combined relative
size, 4.1 to 4.7 per cent of the body
weight, that the gizzard in this
species is large. These data suggest
that the Canada goose is adapted to
feeding on relatively coarse, indiges¬
tible foods as well as tender stems
and leaves of grasses, sedges, and
domestic forage crops. For example,
prior to the establishment of the
Horseshoe Lake Refuge, the Canada
geese which then wintered along the
Mississippi River utilized the bark
and cambium of the willow shoots
growing on the river bars. When
food is not readily available on the
Horseshoe Lake Refuge, the geese
will enter woodlands and eat acorns.
In northern Ontario in early spring
while there is still a deep snow cover

and little food available, the furry

«/

catkins of willows are sometimes
taken.

in Canada Geese 65

Latimer and Rosenbaum ( 1926 j
found the gizzard to be the least
variable (V = 6.35) of the visceral
organs of the hen turkey. In tur¬
keys, the gizzard forms 2.4 per cent
of the body weight; in chickens, the
gizzard forms 2.1 per cent (Latimer
and Rosenbaum, 1926). Mitchell
et at. (1926, p. 109) have stated re¬
garding white leghorn chickens that
“the females consistently exceeded
the males in weights of gizzard,” a
finding which the present data on
Canada geese support. The gizzard
of the starling (Stum us rut gun's)
averages 2.79 and 2.97 per cent of
the body weight for males and fe¬
males respectively (Ltegman, 1954).

Liver Weights. Liver weights were
obtained for 133 Canada geese (Ta¬
ble 4). In autumn, the liver aver¬
ages higher in absolute weight in
males than in females, but when
compared on the basis of body
weight there were no significant dif¬
ferences. In both sexes, the relative
weight of the liver in immatures was
markedly higher than in the older
age classes, but in both sexes it was
relatively greater in adults than in
yearlings. However, in adults, the
liver is notablv more variable in size
than in the younger age classes. The
best estimates that could be made of
the correlation of liver weight to
body weight (data pooled by the
method of least squares, using the z
transformation) were .12 for imma¬
tures, .51 for yearlings and .77 for
adults, no difference between the
sexes being found. While the sig¬
nificance of the values for immatures
is nil, the correlation coefficient in¬
creases with age and in the adults
the values are highlv significant

Transactions Illinois Academy of Science

{ P = .01). These comparisons be¬
tween age-sex classes are considered
valid because nearly all of the geese
studied were shot by hunters in the
morning before the geese had much
opportunity to feed. Liver weights
of Canada geese at other times of
the year will be reported by Hanson
(1961, in press).

Marsden (1940) found no signifi¬
cant correlation of liver weight to
body weight in a sample of ten tur¬
keys studied. The livers of 12
bronzed turkey hens studied by Lati¬
mer and Rosenbaum (1926) com¬
posed 2.25 per cent of the body
weight as compared with 2.40 per
cent for chickens. The coefficient
of variability for liver Aveight in
these turkeys was 29.5.

.V highly significant correlation of
liver weight to body weight in white
leghorn cockerels was reported by
Souba (1923). Mitchell, et al. (1931)
found that in white leghorn cock¬
erels between 0.5 and 5.0 pounds,
the liver decreased from 3.36 to 2.15
per cent of body weight with in¬
creasing body weight. In pullets
between 0.5 to 4.0 pounds, the rela¬
tive weight of the liver decreased
from 2.98 to 1.89 per cent of body
weight. Between the weights of 0.5
and 7.0 pounds, the livers of cockerel
Plymouth rock chickens decreased
from 3.7 to 1.3 per cent of body
weight with increasing body weight ;
in pullets weighing from 2 to 5
pounds, the liver decreased from 2.5
to 1 .9 per cent of body weight
(Mitchell, et al., 1926).

Mitchell, et al. (1931, p. 107) have
stated that “Beyond the 1.5 pound
weight, white leghorn pullets pos¬
sessed a larger average weight of
digestive apparatus, both absolute

and relative, than the cockerels/'
They found a decrease in the weight
of the viscera with age. particularly
in the younger age groups, noting
(p. 136) that “The cockerels were
clearly distinguished from the pul¬
lets by a more rapid decrease in the
percentage weight of the digestive
tract.” The data presented in this
paper for the visceral organ weights
of Canada geese reflect these same
general trends.

It is generally recognized that
small animals have proportionately
larger organs than large animals
(Rensch, 1948). The heart weight
of geese appears to be exceptional in
this respect, but liver weights are
consistent with the rule. Most of
the 46 species of small birds (large¬
ly passerines) studied by Rensch
(1948) had livers that ranged from
3 to 5 per cent of body weight. A
single swan ( Cyynus olor) studied
had a liver weight equal to 1.85 per
cent of body weight. Liver weights
constituted 2.7 to 2.9 per cent of
body weight of Pekin ducks (Sal-
gues, 1939) and 5.01 to 5.02 per cent
of body weight of the European
starling (Stegman, 1954).

The liver weights of migratory
white-crowned sparrows ( Zonotri -
chia leucophrys gambelii) increased
from November through April, high¬
est and lowest values coinciding with
the beginning and ending of migra¬
tion (Oakeson, 1953). The pattern
of liver weight change in the n on-
migratory race ( Zonotrichia leu¬
cophrys null alii) was essentially the
same as in the migratory race (Oake¬
son, 1956). Female ringdoves 15 to
18 months of age have a significantly
larger liver than males, and both
sexes show an increase of liver

Organ Weights in Canada Geese

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Transactions Illinois Academy of Science

weights in summer (10.4 per cent
in males, 6.1 per cent in females)
over winter weights (Riddle, 1928).
In a later study, Riddle (1947, p.
122) stated that “In both ring doves
and pigeons the livers of females
are relatively (usually also absolute¬
ly in doves) heavier than those of
males. In data from doves this ex¬
cess averaged 10-14 per cent ; in
pigeons it was somewhat smaller and
more variable. ’ ’

Pancreas Weights. The weights of
the pancreases of 119 Canada geese
were obtained (Table 5). The data
indicate that pancreas weight de¬
clines slightly with age, both ab¬
solutely and relatively, and that rela¬
tively, the females have a slightly
larger pancreas than males. How¬
ever, no significant correlation be¬
tween pancreas weight and body
weight was found within any age-
sex class.

The pancreas is a highly variable
organ in birds, being second only
to the spleen in this respect. Studies
of pancreas weight in chickens by
Oakberg (1949) and in turkeys by
Latimer (1927) and Marsden (1940)
showed no significant correlation be¬
tween pancreas weight and body
weight. However, Souba’s (1923)
data on 100 white leghorn cockerels
indicate a highly significant correla¬
tion between pancreas weight and
body weight. Perhaps this may be
explained by the sample being high¬
ly uniform in age and uniformity of
the experimental conditions. Ac¬
cording to Salgues (1939), the pan¬
creas of Pekin ducks constitutes 0.22
per cent of the total body weight in
males and 0.26 per cent in females,
values that are considerably lower

than for Canada geese. Latimer and
Rosenbaum (1926) give 0.15 and 0.20
as values for the turkey hen and
chicken respectively. Values for the
white plymoutli rock and white leg¬
horn chickens given by Mitchell, et
al. (1926, 1931) are similar to those
for the Pekin duck.

Summary

Heart weight was the least variable
of the visceral organs of Canada
geese and showed the highest cor¬
relation with body weight. The
spleen was the most variable of the
five organs weighed. In respect to
body weight, the weight of the spleen
of adult males was markedly greater
than that of adult females. The
proventriculus and gizzard, particu¬
larly the latter, were found to be
remarkably large in Canada geese,
constituting 4.1 to 4.7 per cent of
body weight. Liver weights aver¬
aged between 1.43 and 1.79 per cent
of body weight, values for imma-
tures being highest and those for
yearlings lowest of the three age
classes. No correlation between liv¬
er weight and body weight was found
for immatures ; in adults the correla¬
tion was highly significant. The
weight of the pancreas was highly
variable and showed no significant
correlation with body weight. In
most respects, organ weight relation¬
ships in Canada geese showed good
agreement with findings for other
species of birds.

Literature Cited

Hanson, Harold C. 1949. Methods of
determining age in Canada geese and
other waterfowl. Jour. Wildl. Mgmt.,
13: 177-183.

Organ Weights in Canada Geese

Hanson, Harold C. 1958. Studies on
the physiology of Canada geese ( Bran -
ta canadensis interior). Ph.D. thesis,
University of Illinois, 125 pp.

Hanson, Harold C. 1961. The dy¬
namics of condition factors in Canada
geese and their relation to seasonal
stresses. Arctic Institute of North
America, In press.

Hanson, Harold C. and Robert H.
Smith. 1950. Canada geese of the
Mississippi Flyway, with special refer¬
ence to an Illinois flock. Ill. Nat.
Hist. Surv. Bull., 25: 67-210.

Harmon, I. W., E. Ogden and S. F. Cook.
1932. The reservoir function of the
spleen in fowls. Am. Jour. Physiol.,
100: 99-101.

Hartman, Frank A. 1955. Heart weight
in birds. Condor, 57: 221-238.

Kirkpatrick, C. M. 1944. Body weight
and organ measurements in relation
to age and season in ring-necked
pheasants. Anat. Rec., 89: 175-194.

Latimer, Homer B. 1927. Correlation
of weights and lengths of the body,
systems and organs of the turkey hen.
Anat. Rec., 35: 365-377.

Latimer, H. B. and J. A. Rosenbaum.
1926. A quantitative study of the
anatomy of the turkey hen. Anat.
Rec., 34: 15-23.

Marsden, S. J. 1940. Weights and
measurements of parts and organs of
turkeys. Poul. Sci., 19: 23-28.

Mitchell, H. H., L. E. Card and T. S.
Hamilton. 1926. The growth of
white Plymouth rock chickens. Univ.
Ill. Agr. Exp. Sta. Bull., 278: 71-132.

Mitchell, H. H., L. E. Card and T. S.
Hamilton. 1931. A technical study
of the growth of white leghorn chick¬
ens. Univ. Ill. Agr. Exp. Sta. Bull.,
367: 83-139.

Oakberg, E. F. 1949. Quantitative
studies of pancreas and islands of
Langerhans in relation to age, sex
and body weight in white leghorn
chickens. Am. Jour. Anat., 84: 279.

Oakeson, Barbara B. 1953. Cyclic
changes in liver and spleen weights
in migrating white-crowned sparrows.
Condor, 55: 3-16.

Oakeson, Barbara B. 1956. Liver and
spleen weight cycles in non-migratory
white-crowned sparrows. Condor, 58:
45-50.

Quiring, Daniel P. and Paul H. Bade.
1943. Organ and gland weights of
the English sparrow. Growth, 7: 299-
307.

Rensch, Bernhard. 1948. Organ pro-
portionen und Korpex grosse bei
Yogeln und Saugetieren. Zoologische
Jahrbucher. Abteilung. fur Allgemeine
Zoologie und Physiologie, 61: 337-412.

Riddle, Oscar. 1928. Sex and seasonal
differences in weight of liver and
spleen. Proc. Soc. Exp. Biol, and Med.,
25: 474-476.

Riddle, Oscar. 1929. Endocrine regula¬
tion of reproduction. Endocrin., 13:
311-319.

Riddle, Oscar. 1947. Endocrines and
constitution in doves and pigeons.
Carnegie Inst, of Wash., Publ. No.
572., 306 pp.

Salgues, R. 1939. Contributions nou-
velles a la physiologie de la plume F.
L’osteomalaise aviare et ses repercus¬
sions sur le plumage. L’Oiseau et la
revue Francais d’ornithologie. New
Series, 9: 33-51.

Smith, H. L. 1928. The relation of the
weight of the heart to the weight of
the body and of the weight of the
heart to age. Am. Heart Jour., 4:
79-93.

Souba, A. J. 1923. Variations and cor¬
relation of the organs of single comb
white leghorn cockerels. Anat. Rec.,
26: 291-297.

Stegman, Leroy. 1954. Variation in a
flock of the European starling. Auk,
71: 179-185.

Manuscript received February 2<8. /.%‘2.

JUNIOR ACADEMY PUBLICATIONS

John C. Frye

President , Illinois State Academy of Science
121 natural Resources Bldg., Urbana

The Illinois Junior Academy of
Science, since its formation, has been
a major project of the State Acade¬
my of Science. In recent years the
Junior Academy has had a substan¬
tial growth. The success of the dis¬
trict and state science expositions,
which testifies to the increasing in¬
terest among junior and senior high
school students, has prompted the
Council of the Academy to explore
additional means of assisting in the
development of young scientists. To
this end a publication program was
approved.

A limited number of Junior Acade¬
my papers will be published in an
appropriately designated section of
the Transactions of the Illinois State
Academv of Science. There are two
methods by which a student’s paper
can be submitted for consideration,
but in either case the manuscript
must conform to acceptable stand¬
ards. Source material must be prop-
erlv documented, references and ci-
tat ions accurately given in accord¬
ance with the style currently in use
in the Transactions, and the text
typed double spaced. Illustrations
should be used only where essential
and must be prepared so that they
can be clearly reproduced within

the limitations of the page size. The
student’s name and school should be
accompanied by his home address
so that he can be contacted for nec¬
essary editorial changes.

The first method of submitting a
paper for possible publication is by
way of the paper sessions of the
State meeting of the Junior Acade¬
my. The highest ranking papers at
the State meeting will automatically
be given consideration.

The second method is by sponsor¬
ship by a member of the State Acade¬
my of Science who has personal
knowledge of the project. Although
any member of the Academy may
serve as such a sponsor, it is pre¬
ferred that the teacher under whom
the student is working be the spon¬
soring member. In such cases the
sponsor should send the manuscript,
accompanied by a supporting letter
of transmittal, directly to the Editor
of the Transactions.

It is unfortunately true that limi¬
tations of space make it likely that
it will not be possible to publish all
of the worthy papers. Final decision
for acceptance for publication rests
with the Editor of the Transactions,
and his Editorial Board.

[70]

THE HATCHING MUSCLE IN THE AMERICAN COOT

JAMES R. FISHER

Carbondale, Illinois

It is believed that a young’ bird
hatches by using a muscle which ex¬
ends from the neck to the middle of
the back of the head. This muscle
increases in size during development
but most rapidly just before hatch¬
ing. After hatching, it decreases in
size. In grown birds the muscle, al¬
though relatively small, is used for
other purposes. Inside the egg the
embryo is so positioned with its bill
against the shell, and with the neck
curled, that it can use this muscle
to extend the head and bill upward
to pip the egg and eventually hatch
out.

There has been some work done
on this muscle in the chicken (Kei-
bel, 1912 ; Pohlman, 1919 ; Fisher,
1958), in North American grebes
(Fisher, 1961a) and in Franklin’s
Gull (Fisher, 1961b). The work of
Fisher supports this idea of hatch¬
ing, as outlined above.

My purpose in this paper is to
point out and explain the phases of
the gross development of this muscle
in the American Coot, Fulica ameri-
cana, from the time the embryo
weighs just a few grams to the age
of two weeks.

Materials and Methods

One hundred fifty-eight eggs were
taken from nests in the Delta Marsh¬
es at Delta, Manitoba, Canada. Im¬
mediately upon return to the Delta
Waterfowl Research Station, the
eggs were incubated, usually within

two or three hours of collection. The
eggs in each clutch were numbered
and the clutches kept separate in
the incubator. Incubation was at
99.5 degrees F and from 60 to 70%
relative humidity. The young coots
were removed from the incubator at
one day of age and kept in indoor
pens. Their food was a higli-protein,
turkey pre-starter mix. An abun¬
dance of water was supplied for
drinking and swimming. Never
were more than seven or eight young
coots kept in any one pen.

Since the female coot lays one egg-
each day and starts incubating upon
the laying of the first egg, the eggs
in any one clutch were in different
stages of development at the time
of collection. It was thus not pos¬
sible to age the embryos by back¬
dating from the time the first egg
hatched. Sometimes two eggs in the
same clutch hatched on the same
day. For these embryos which were
not permitted to complete their de¬
velopment I used body weight as an
index to the stage of development.
After hatching, both age and weight
were used to indicate development.
1 attempted to get a series of em¬
bryos representing all weight stages.

Removal of unhatched embryos
was done within 30 minutes of re¬
moval of the egg from the incubator.
The egg was cracked on a line around
the end containing the air pocket so
as not to damage the specimen in
any way and to permit examination

[71]

Transactions Illinois Academy of Science

of the position of the head. The
membranes were removed from the
embryos, including the yolk sac in
embryos in which it had not yet
been completely drawn into the body.
The embryos were dried by gently
rotating them on paper towels for
not more than one minute. After
drying, the embryos were weighed
on a balance to the nearest tenth of
a gram. The muscles were removed
with iridectomy scissors and weighed
immediately on a Roller-Smith Pre¬
cision Balance to the nearest ten-
thousandth of a gram. The birds
were then preserved in formalin for
later study of the egg-tooth.

The thickness of the egg shell was
measured at two different places on
the edge of the breakage line of
eggs which had produced normally-
hatched young. A dial micrometer,
calibrated to thousandths of an inch,
was used.

Results

Segmentation begins to appear
faintly at approximately the 7-gram
stage of embryonic development.
Segments are definitely visible by
the 9-gram stage and are in all cases
visible throughout incubation. But
just before hatching the segments
may be difficult to see because of the
large lymph content of the muscle.
Segmentation gradually disappears
after hatching. At six or seven days
of age it is very faint, and in many
specimens is not visible thereafter.
In some embryos only parts of the
posterior and anterior boundaries of
the individual segments remain.

The development of segmentation
starts anteriorly (Fig. 1). The an¬
terior segment appears first and is
the largest throughout the history of

the muscle. The posterior segment
is always the smallest. Usually three
pairs of segments are present be¬
fore hatching. In only one instance
was there any variation in the num¬
ber of segments; one chick had a
slight enlargement of a fourth pair
of segments.

The first medial contact between
segments of opposite sides is be¬
tween the anterior pair; this is the
condition in one-fourth of the 6-
gram specimens. By the time the
embryos reach 7 grams in body
weight most show medial contact at
least between the first pair of seg¬
ments. At the 8- or 9-gram stage
the medial contact has proceeded
posteriorly to include the first two
pairs of segments in approximately
50% of the specimens. There is
medial contact between the first two
pairs by the prepip stage in most
specimens. And some individuals
also have limited contact between
the anterior parts of the third pair
of segments. After hatching there
is a progressive decrease, from pos¬
terior to anterior, in the amount of
medial contact, until at 17 days of
age only part of the inner margins
of the first pair are touching (Fig.

i).

The muscle increases in absolute
weight from 0.020 grams at 8 grams
of body weight to 0.24 grams at 15
grams of body weight, at which time
the embryo is ready to pip. Muscle
weight decreases from the pipping
stage, when it weighs 0.16 grams, to
approximately 0.05 or 0.07 grams at
two or three days of age. After this
it slowly increases in weight.

There are two times in develop¬
ment when the muscles are very
large, if the muscle weight is com-

Hatching Muscle

PIPPING JUST HATCHEO

3 DAYS

7 DAYS

17 DAYS

Fig. 1. — Diagrams of development of the egg tooth and hatching muscle in
the American coot. Weights refer to body weights.

puted in relation to body weight
(Fig. 2). The first time is at the
4-gram stage or earlier when the
muscle constitutes about 1.56% of
bod}" weight. The second time is
between the body weights of 13 and
16 grams. The first instance is due
to the more rapid development of
the anterior part of the embryo.
The latter time is during the prepip
stage. As the development pro¬
gresses from the 4-gram stage, the
muscle weight becomes relatively less
until about the 7- or 8-gram stage
when the muscle may make up as
little as 0.2% of body weight (Fig.
2). During the rest of the incuba¬
tion period weight increases rapidly,
in relation to body weight, and
reaches a maximum of 1.64% of
body weight. During the pipping

period the muscle begins to decrease
very rapidly in weight, and this con¬
tinues through the second day of
age. Beginning at about the third
day of age the decrease in relative
weight becomes more gradual and
continues to decline at least until
14 days of age. At five or six days
of age the muscle is about the same
relative size as it was when the em¬
bryo weighed only 7 or 8 grams.

The lymph glands situated on
either side of the pair of muscles on
the neck are believed to have some
influence on the action of the mus¬
cle. It is known that they produce
lymph which moves into the muscle,
and Pohlman (1919) states that this
hinders the action of the muscle.
H. I. Fisher (1961a) believed that
this infiltration of lymph may ac-

Transactions Illinois Academy of Science

1.64-

1.56
1.44
1.40
1.36
1.32
128
1.24
1.20
116
I 12
1.08
1.04
1.00
96
.92
88
84
80
76
72
68
64
60-
56
52
48
44
40

H P

P* PIPPED

H* HATCHED

NOS. - DAYS OF AGE

36-
32 -
28-
24-
20-
16-

9 9

8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 50 58

BODY WEIGHT IN GRAMS

— * — i - » —

88 100

-l - i - > -

120 132

Fig. 2. — Scattergram to illustrate changes in relative size of the hatching
muscle in the American coot.

tually help the action of the muscle,
perhaps through the provision of fats
and sugars for energy. Because of
these varying views the lymph glands
will be discussed in this paper.

The lymph glands are not grossly
apparent before the embryo reaches
7 grams of body weight, and of 21
specimens, 3 days or more of age,
the glands were completely absent
or too small to measure in 12. There
is not much change in the dimensions

of the glands between the 7-gram
and 16- to 18 -gram stages. At the
latter time the width decreases and
the glands become more elongate.
During the pipping process the
glands become thin and string-like.

The lymph glands frequently con¬
sist of scattered lobes or kernels.
This scattered condition may appear
at any stage, but it is less frequent
in the pipping and hatching stages.
Triangularly shaped glands occur in

Hate h in g Muse l e

about 8% of the birds prior to hatch¬
ing, but are not seen after hatching.

In only 4 of 97 birds were the
glands absent or less than 1 mm. in
diameter on one side or the other.
There was no difference in the fre¬
quency of occurrence or in size be¬
tween the glands of the two sides.

I noted earlier that the muscle
has a large lymph content. This re¬
sults in a clear to yellowish “lymph
color ’ ’, as I shall call it ; lymph color
first appears in the muscles at the
9-gram stage, but only to a slight
degree and only in 50% of the speci¬
mens. When the 13-gram stage is
reached nearly all the specimens
show a strong lymph color which
continues through the hatching peri¬
od. In this period the muscle ap¬
pears lymph-filled, a condition which
appears in only about one-fourth of
the specimens at the 12-gram stage.
Lymph-filled muscles were never
found in specimens more than 12
hours of age, but the muscles retain
a pinkish-yellow color until about
the 3-day stage when the “normal”
reddish color begins to appear. By
5 to 7 days after hatching, the mus¬
cles are reddish.

Since the development of the egg-
tooth, as a part of the body aiding
in hatching, is related to the develop¬
ment of the hatching muscle, I shall
indicate here a few of the more ob¬
vious stages. The smallest embryo
in which I could find any indica¬
tion of an egg-tooth weighed 0.78
grams. There is a definite bump on
the end of the bill by the 3-gram
stage, and by 10-grams in body
weight this protuberance is pointed
upward and forward (Fig. 1). The
egg-tooth is largest at the time of
pipping. It diminishes in size very

rapidly in the later stages of pipping
and immediately after hatching.
There is no evidence of an egg-tooth
after three days of age.

Since there may be a relationship
between the size of the muscle and
the thickness of the shell, I shall
give here the measurements of the
thickness of the zone of breakage
during hatching. In 34 eggs this
averaged 0.0105 ±0.0001 inches, with
a range from 0.0095 to 0.0122 inches.

Discussion and Summary

Development of segmentation in
the hatching muscle is the same in
coots as in the chick (Keibel, 1912,
Pohlman, 1919, Fisher, 1958) and
in the Franklin’s Gull (Fisher,
1961b). Segments or blocks of tissue
which will form the muscle first ap¬
pear distinctly to the unaided eye at
the 0.5-gram stage. The first seg¬
ments in the formed muscle appear
at 7 grams, are definite by the 9-
gram stage, and consist of 3 pairs
in most instances. A 4th segment
was observed more often in the
North American grebes (Fisher,
1961) than in the coots. The de¬
velopment starts anteriorly and pro¬
gresses toward the posterior. Thus
the anterior segments are the largest
and the posterior segments are the
smallest.

Segmentation is not visible in some
birds after 6 or 7 days of age and
only traces of the segmental bound¬
aries remain visible in others. The
first medial contact becomes visible
between the anterior pair of seg¬
ments. The area of contact increases
between other segments, reaching the
3rd pair just before hatching. After
hatching, the line of medial contact

Transactions Illinois Academy of Science

decreases progressively. This same
developmental pattern was noted in
the above literature.

Absolute weight of the muscle in
the coot increases from the 8-gram
stage of body weight to the pipping
stage. Once the chick is hatched, the
weight of the muscle decreases for a
few days and then starts to increase
again. There is no apparent de¬
cline of the weight during or after
hatching in the grebes, but the data
were few.

Relative weight is great at first
observation ; it then decreases until
the 8-gram stage. During the rest
of the incubation period the weight
increases relatively, up to the pre¬
pip and pipping periods when it
starts to decline rapidly. This de¬
crease is more gradual from about
3 days to at least 14 days of age.

There is not much change in the
size of the lymph glands from the
time they first become apparent,
when the bird weighs 7 grams, un¬
til it weighs 16 to 18 grams. The
width decreases and the glands be¬
come string-like at 16 to 18 grams
when the chick is pipping. These
glands are absent after approxi¬
mately 3 days of age. Although
there are no visible lymph glands in
the area of the muscle of the North
American grebes, Fisher (1961a)
said there is an obvious infiltration of
lymph. In the Franklin’s Gull
(Fisher, 1961b) the glands start to
elongate at 10 grams. This elonga¬
tion increases more rapidly than the
width, reaching its maximum just
after hatching.

The color of the muscle is changed
by the infiltration of the lymph.
This “lymph color” first appears at

9 grams in 50% of the specimens.
It then increases in amount and num¬
ber of specimens in which it is visi¬
ble, showing strong lymph discolora¬
tion up through hatching. After 12
hours of age the muscle slowly re¬
turns to its normal reddish color.

The egg-tooth first appears at 0.8
grams of body weight and grows
larger and more pointed up to pip¬
ping. During pipping and hatch¬
ing the egg-tooth decreases in size.
There is no sign of it after three
days of age.

Thickness of the egg shell at the
time of hatching was 0.0105 ±0.0001
inches.

Success in maintaining and raising
young coots, which has been very
difficult in the past, was thought to
be mostly the result of three factors :
(1) putting newly hatched young in
with a few older birds (ducks or
coots) that were feeding success¬
fully; (2) allowing at least five
square feet for each young bird in
the pen; and (3) using a very high
protein diet.

Acknowledgments

I am grateful to Mr. Albert Hoch-
baum, Director, Delta Waterfowl
Research Station, Delta, Manitoba,
Canada, who gave me permission to
collect eggs and coots on the research
station area and who also provided
the space and facilities to carry out
the research for this paper. Many
thanks go to my father for his con¬
stant help and advice in the labora¬
tory and field. To my mother I am
indebted for aid in collecting eggs
which were badly needed.

Hatching Muscle

1 1

Literature Cited

Fisher. Harvey I. 1958. The “hatching
muscle” in the chick. Auk, 75: 391-
399, 12 figs.

- 1961a. The hatching mus¬
cle in North American grebes. Con¬
dor, 63(3) : 227-233, 4 figs.

- 1961b. The hatching mus¬
cle in Franklin’s Gull. Wilson Bull.,
in press.

Keibel, Franz. 1912. Wie zerbricht der
ausschliipfende Vogel die Eischale?
Anat. Anz., 41: 381-382.

Pohlman, A. G. 1919. Concerning the
causal factor in the hatching of the
chick, with particular reference to the
musculus complexus. Anat. Rec., 17 :
89-104, 2 figs., 8 tables.

Manuscript received June ), 1961.

FOOD HABITS OF THE LEOPARD FROG
{RAN A PI PIE NS SPHENOCEPHALA )

IN A MINNOW HATCHERY

ROBERT JEFFERY LEWIS
University School, Carbondale, Illinois

This paper is based upon an Illi¬
nois Junior Academy of Science
project.

Due to the abundance of ideal
habitat free from predaceous fishes,
the leopard frog is unusually abun¬
dant in most minnow hatcheries. In
view of its abundance it is desirable
to know whether or not it feeds on
minnows.

Other studies have been made on
the food habits of the leopard frog,
but not on specimens taken where
small fishes were known to be abun¬
dant. In a study done by Kilby
(1935), out of 443 stomachs con¬
taining food, the bulk of the food
consisted of insects and spiders. The
insects were beetles, lepidopterous
larvae, crickets, grasshoppers, and
some aquatic insects. Knowlton
(1944) examined 97 stomachs and
found the following : shorthorned
grasshoppers, field crickets, leaf
hoppers, false chinch bugs, pea
aphid, English grain aphid, click
beetles, lepidopterous larvae, house¬
flies, blowflies, mosquitoes, mosqui¬
to eggs, deer flies, and ants. In a
study done by Drake (1914) out of
209 stomachs, 60 percent contained
insects. Other food items included
mollusks, crustaceans, myriapods,
and spiders.

Study Area

The hatchery at which the study
was done is in the Mississippi River
bottoms of Jackson County, Illinois
near the town of Gorham. The whole
study area covered 6 acres. These
6 acres were divided into 19 ponds
ranging from 1/20 to 1 acre and
ranging in depth from 2 to 4 feet.
On the levees and their shorelines,
there was a heavy growth of grass
and weeds. There were water plants
and algae growing on the surface of
some ponds.

Method of Procedure

The 46 specimens for the study
were collected by the use of an air
gun. The stomach contents were
determined after dissection.

During the course of the study a
record was kept of the availability
of the more noticeable food organ¬
isms with special reference to tin*
minnow population of the ponds.

Results

Under the conditions of the study
area, the leopard frog fed exclusive¬
ly on insects (Table 1). The ma¬
jority of these insects were terres¬
trial. Some aquatic larvae were

Food of Leopard Frog

Table 1. — Stomach contents of 46 leop¬
ard frogs i.

Percent

Food Item Occurrence

Terrestrial Beetles . 43

Unidentified Insect Remains.. 24

Miscellaneous Insects . 17

Crickets . 11

Aquatic Insect Larvae . 9

Empty . 7

1 Collected April 25 to September 12
in a southern Illinois minnow hatchery.

found indicating that the frog’s may
feed underwater. Minnows, smaller
frogs, tadpoles, and crayfish were
available in the study area. None
of these items were found in the
stomachs.

Acknowledgments

The author wishes to thank his
father, Dr. William M. Lewis, Sr.
and mother, Mrs. Sue D. Lewis for
assistance in identifying food items
and in preparation of this paper.
The science project upon which this
paper is based was sponsored by Mr.
Carol Hampton, Lecturer, Univer¬
sity School, Carbondale, Illinois.

Literature Cited

Drake, Carl J. 1914. The Food Habits
of Rana pipiens, Schrebar, Ohio Nat¬
uralist, 14: 259-269.

Kilby, John D. 1935. Feeding Habits
of Rana pipiens sphenocephala , Jour,
of Fla. Acad, of Sci., 8(1): 71-104.
Knowlton, G. F. 1944. Some Insect
Food of Rana pipiens, a Utah Re¬
source. Copeia, 2: 119.

Manuscript received March Ilf, 1961 .

STOMACH CONTENTS OF BULLFROGS (RAN A
CATESBE1ANA ) TAKEN FROM A MINNOW HATCHERY

WILLIAM M. LEWIS, JR.

Carbon dale , Illinois

In the southern Illinois region,
the bullfrog is one of the most abun¬
dant vertebrates in minnow hatch¬
eries. It is therefore desirable to es¬
tablish its status as a predator on
minnows. This was the objective of
the present study.

The study was done as a science
project for the Illinois Junior Acade-

mv of Science.

«/

Several studies have been made of
the natural feeding habits of the bull¬
frog. Among the more exhaustive is
that of Korschgen and Moyle (1955),
who examined the contents of
455 stomachs from specimens coll-
lected from central Missouri farm
ponds. The percent volume for the
principal food items were as fol¬
lows: insects, 32; crayfishes, 26;
frogs, 11 ; tadpoles, 10 ; meadow mice,
3 ; fishes, 2 ; snails, 2 ; toads, 2 ; mis¬
cellaneous invertebrates, 2 ; and
snapping turtles, 1.

Frost (1935) reported that in¬
sects comprised the greatest part of
the diet of 25 smaller frogs. Ants
were eaten in considerable numbers.
Spiders and snails formed the largest
part of the diet by volume. Upon
examining the larger bullfrogs,
Frost found crayfishes in 4, frogs in
2 and mice in 2.

Needham (1905) concluded that
snails and insects were the main
items of the diet. In 16 bullfrogs
collected from New York, he found
18 snails, 3 spiders, 3 crustaceans
and 2 vertebrates.

A study by Perez (1951) in Puer¬
to Rico showed the following princi¬
pal food items in terms of percent
volume of total stomach content :
insects, 19; plant matter, 11; and
bullfrogs, 4.

Pope (1947) gives the combined
results of five workers who tabu¬
lated the contents of about 200
stomachs and more casual informa¬
tion from others. The diet of young
frogs was comprised of insects and
other small invertebrates, at least
half of which were non-aquatie.
Larger frogs preyed upon numerous
invertebrates and vertebrates such
as fishes, frogs, salamanders, young
turtles, snakes, moles, mice and
birds. Frogs and crayfishes seemed
to be the chief food items of larger
frogs.

Other more casual observers add
to the widely varied list of food
items. Baker (1940) gave crayfishes
as the chief food item of the bull¬
frog in eastern Texas. Dickerson
(1906) lists fishes, small turtles,
young water birds, and frogs as food
items forming the greatest part of
the diet. Breckenridge (1944) in¬
cludes insects, fishes, crayfishes,
birds, and other frogs as popular
food items. Morris (1945) states
that the bullfrog is cannibalistic to
a marked degree. Other food items
listed by Morris are mice, crayfishes,
salamanders, snails, small fishes,
worms, and various insects, both lar-

[80]

Feeding Habits of Bullfrog

val and adult, that are found among
the water weeds.

Dyche (1914) examined 30 stom¬
achs from bullfrogs taken from a
Kansas fish hatchery. He found 32
fishes in 14 of the 30 stomachs. Oth¬
er items found included crayfishes,
insects, spiders, and snails.

Study Area

The frogs utilized in this study
were taken from a minnow hatchery
located in the Mississippi River Bot¬
toms of Jackson County, Illinois.
The collecting area covered 6 acres.
Within this area were 19 ponds
ranging in size from 0.05 to 1.0 acre.
Throughout the summer and fall,
the levees of the ponds were covered
with a dense growth of vegetation.

Method

The original plan of sampling
called for collecting ten frogs per
week. This plan was followed from
spring to midsummer. However,
after this time, the frogs became so
scarce and vegetation cover so dense
that it was not possible to obtain the
desired number. It was also noted
that the larger frogs became rela¬
tively more scarce. A total of 123
frogs were utilized in the study.

The specimens were collected pri¬
marily during the daylight hours by
use of a .22 caliber rifle. They were
all collected in the water or at the
water’s edge. As each frog was col¬
lected it was marked as to which
pond it was taken from and stomach
contents were determined by dissec¬
tion. The size of frog was measured
from the snout to the posterior end
of the body. The abundance of vari¬
ous food items in the habitat was

noted throughout the course of the
study.

Results

Plant matter constituted only a
minor portion of stomach contents
and was considered to be accidentally
ingested with food. For this reason
it was not listed with stomach con¬
tents (Table 1) .

Insects were found to be in 68
percent of the stomachs examined.
Terrestrial beetles (32 percent),
aquatic beetles (24 percent), and
dragon flies (13 percent), were the
most numerous food organisms in
this group. Dragonflies constituted
a large part of the diet in July, Au¬
gust, and September, probably be¬
cause of availability. Twenty-five
other kinds of insects were recog¬
nized but were relatively unimpor¬
tant as food items.

Crayfishes (especially Orconectes
immunis ) found in 30 percent of
the stomachs examined constituted
the second most prevalent food item.
A decrease in the number of cray¬
fishes eaten in late summer was prob¬
ably due to a poisoning program
which reduced the number of cray¬
fishes present in the ponds.

Frogs (especially Ran a pipiens)
were the third most prevalent food
item and were found in 24 per cent of
the stomachs examined. Although
cricket frogs ( Acris gryllus were
known to be abundant in the study
area at the time the bullfrogs were
being collected they occurred in only
1.6 percent of the stomachs ex¬
amined. Young Fowler’s toads ( Bufo
woodhousei ) were also abundant in
May and June, but none were found
in the stomachs of the bullfrogs ex¬
amined.

Table 1. — Percent Occurrence of Food Items in Stomachs of 123 Bullfrogs Collected
From Minnow Hatchery Ponds, Gorham. Illinois.

Transactions Illinois Academy of Science

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Feeding Habits of Bullfrog

Since the primary objective of
this study was to determine the
status of the bullfrog as a predator
on fishes, the data collected for this
study were analyzed with this pur¬
pose in mind. Fishes were the fourth
most prevalent food item. Minnows
( Notemigonus crysoleucas and Pime-
phales promelas) were known to be
abundant in most ponds on the hatch¬
ery but were found in only 14 per¬
cent of the stomachs examined. On
comparing size of frog with stomach
contents, it Avas found that larger
frogs fed more heavilv on minnows.
They occurred in 25 percent of the
stomachs from frogs 6 inches or over,
15 percent in 5-incli to 6 -inch frogs,
7 percent in 4-inch to 5-inch frogs,
and 9 percent in frogs 3 inches or
less in body length.

Conclusions

Under conditions existing in a bot¬
tomland minnow hatcherv of the
southern Illinois region, the bullfrog
appears to utilize primarily insects,
crayfish, frogs, and minnows as food.
Of these principal items minnows
are utilized the least. It is ques-
tionable if the bullfrog constitutes
a very serious predator on minnows
under hatchery conditions.

Acknowledgments

The writer is indebted to his fa¬

ther Dr. William M. Lewis and his
mother Mrs. Sue D. Lewis for aid in
identification of food items and prep¬
aration of this report and to Mr.
Carol Hampton, Lecturer, Universi¬
ty School, Carbondale, Illinois, who
sponsored the science project upon
which this paper is based.

Literature Cited

Baker. R. H. 1942. The Bullfrog, a
Texas Resource. Texas Game and
Fish Comm. Bull., 23, pp. 3-7.

Breckenridge, W. J. 1944. Reptiles and
Amphibians of Minnesota. Univ. Minn.
Press, Minneapolis, Minn., 202 pp.

Dickerson. Mary C. 1907. The Frog-
Book. Doubleday, Page and Co., New
York, 253 pp.

Dycite, L. L. 1914. Ponds, Pond Fish,
and Pond Fish Culture. State Dept.
Fish and Game, Kan. Bull., 1: 149-158.

Frost, S. W. 1935. The Food of Rana
catesbeiana Show. Copeia. 1: 15-18.
Korschgen, J., and D. L. Moyle. 1955.
Food Habits of the Bullfrog in Cen¬
tral Missouri Farm Ponds. Amer.
Mid. Nat., 54(2) : 332-341.

Morris, Percy A. 1945. They Hop and
Crawl. Jaques Cattell Press, Lan¬
caster, Pa. 253 pp.

Needham, J. G. 1905. The Summer
Food of the Bullfrog ( Rana cates¬
beiana) Shaw at Saranac Inn. N. Y.
State Mus. Bull., 86: 9-15.

Perez, M. E. 1951. The Food of Rana
catesbeiana Shaw in Puerto Rico.
Jour. Agric. Univ. Puerto Rico, 35(4) :
136-142.

Pope, C. H. 1947. Amphibians and
Reptiles of the Chicago Area. Chi¬
cago Nat. His. Mils., 275 pp.

Manuscript received March l!h 1961.

PREPARATION OF MANUSCRIPTS FOR
THE TRANSACTIONS

For publication in the Transactions, articles must present significant
material that has not been published elsewhere. Review articles are ex¬
cepted from this provision, as are brief quotations necessary to consider
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Titles should be brief and informative. The address or institutional
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literature are to be placed in footnotes. Citations under Literature Cited
are as shown below:

Doe, John H. 1951. The life cycle of a land snail. Conchol., 26(3):
21-32, 2 tables, 3 figs.

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XT'

R33%

Transactions

of fhe

Illinois

State Academy
of Science

Volume 55
No. 2
1962

Springfield, Illinois

TRANSACTIONS of the ILLINOIS STATE ACADEMY of SCIENCE
Editorial Board:

Wesley J. Birge, University of Illinois, Editor and Chairman

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♦

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TRANSACTIONS

OF THE

ILLINOIS STATE
ACADEMY OF SCIENCE

VOLUME 55 - 1962

No. 2

Illinois State Academy of Science

AFFILIATED WITH TIIE

Illinois State Museum Division
Springfield, Illinois

PRINTED BY AUTHORITY" OF THE STATE OF ILLINOIS

Otto Kerner, Governor

May 1, 1963

CONTENTS

Observations on a Colony of Big-Eared Bats, Plecotus rafinesquii.

By Donald F. Hoffmeister and AVoodrow AAt. Goodpaster . 87

A Second Porcupine Record for Illinois. By Paul W. Parmalee . 90

Geographic and Host Distribution of Blood Parasites in Columborid
Birds. By Norman D. Levine . 92

Dental Anomalies of the Raccoon. By Robert L. Martin . 112

Leaf Characteristics of Two Hybrid Junipers. By Margaret Ivaeiser. . Ill

Germination Capacity in American Basswood. By William C. Ashby. . 120

Comparative Effectiveness of DDT Selection Methods in Drosophila
melanogaster meigen. By Thom as R. Kallstedt and
Jack Bennett . 124

Oxygen Consumption in The Small Short-Tailed Shrew (Crypt of is

parva) . By Carl J. Pfeiffer and George H. Gass . 130

Nomenclature of The Late Mississippian White Pine Shale and

Associated Rocks in Nevada. By R. L. Langenheim, Jr . 133

Ground Water Geology of The DeKalb and Sycamore quadrangles.

By Loren T. Caldwell . 146

Anomalous Alar Plate Regulation in the Early Chick Neural Tube.

By Wesley J. Birge . 164

Academy Business . 167

Treasurer’s Annual Report . 176

Officers, Committees, and Section Chairmen, 1962-63 . 178

OBSERVATIONS ON A COLONY OF BIG-EARED BATS,

PLECOTUS RAFINESQUII1

DONALD F. HOFFMEISTER and WOODROW W. GOODPASTER

University of Illinois, Urbana, and Cincinnati, Ohio

Colonies of big-eared bats, Pleco-
tus rafinesquii, are so uncommonly
found that our limited observations
of a small colony are worthy of
recording. A small colony occupies
an unused cistern, now far removed
from any buildings, about one mile
northeast of Reelfoot Lake, Obion
County, western Tennessee. The bats
were first discovered in the cistern
in May 1950, but they were present
“many years” before that, accord-
ing to local residents. During the
months we have visited the cistern —
February, March, April, May, and
September — from 1 to 64 Plecotus
rafinesquii, and only this species,
have been present.

The cistern is located in slightly
rolling country among the bluffs
back from Reelfoot Lake. The im¬
mediate area has a sparse second
growth of trees, most of which are
under 30 feet high. To the north
there is a thicket of blackberries but
no trees. The mouth of the cistern
projects 2 feet above ground level
and is 30 inches in diameter. It is
always open. The cistern is about
25 feet deep and “bells-out” inside
(see Fig. 1). The cistern is built
of bricks and stone but the inner
face is smoothed with cement. Water,
always present in the bottom, fluc¬
tuates in depth but is always less
than 8 inches deep in our exerience.

Plecotus rafinesquii hang in the
top-half of the cistern in the warmer

1 Aid for this study was furnished in
part hv N.S.F. grant G 19392.

parts of the year (May 25, Septem¬
ber 2 and 3). In the colder months
they hang in the bottom half. On the
occasion when 64 bats were present,
most of them hung in three clusters,
4% to 6% feet above the water (see
Fig. 1) with about 20 in each clus¬
ter. Two or three individuals were
hanging separately higher on the
walls. On cold days, these bats are
apparently in hibernation, for they
are cold to the touch, their ears are
rolled down, and the bats are so
sluggish that bands could be attached
to most individuals before they
aroused.

The hibernating chamber must be¬
come cold at certain times and must
receive considerable light since there
is no cover on the cistern. On the
night of March 21, 1960, the tem¬
perature outside the cistern Avas as
low as 8°F ; earlier in the winter
the temperature was lower. On
April 2, 1961, at 8 :30 a.m., the tem¬
perature near the bottom of the cis¬
tern was 49°F; at ground surface,
51 °F. We do not know if the tem¬
perature drops below freezing within
the cistern but suspect that it must
on occasions. Light filters in the top
of the cistern, especially on bright
days, and it must be considerably
lighter here than just inside the en¬
trance of most caves.

Big-eared bats must move in and
out of this colony during the winter
months. For example, on February
14, 1961, about 50 Plecotus were
present of which 13 had bands previ-

[87]

Transactions Illinois Academy of Science

ously affixed. On April 1, 1961, one
and one-half months later, only 3 of
the banded bats were present yet it
was too cold for them to be active
within the cistern. On April 1, the
ground surface temperature was
about 34°F. ; earlier on February 14,
it was about 60° F. On the latter
date, the bats were active within the
cistern and it appeared as if they
might be copulating.

Some Plecotus use the cistern more
than one winter, but new individ¬
uals move in with the ‘ ‘ resident ' ’
bats on succeeding years. On March
21, 1960, 60 bats Avere banded in the
cistern. About 11 months later, on
February 14, 1961, of about 50 bats
present, only 13 had been banded
previously. On April 1, 1961, of 28
bats, 11 had been banded a little over
a year before. However, among these
11, only 3 AArere from the 13 wearing
bands observed on February 14. On
April 4, 1962, of 3 bats present, one
was banded 2 years earlier and one
was banded one year earlier. It ap¬
pears that throughout the cold peri¬
od of late winter and early spring,
there is a movement of bats in and
out of this colony.

A colony of Plecotus rafinesquii
can be greatly reduced or depleted
in size and it will successfully re-
build. Early in our observations,
most of the colony was removed. On
May 25, 1950, only 1 bat was pres¬
ent and it was removed ; on Septem¬
ber 2, 1950, 16 bats were present and
15 were removed; on May 31, 1951,
28 bats Avere present and 25 were
removed. Within a year’s time, 41
of 45 bats AArere removed and pre-
served as skins. Nevertheless, on
March 21, 1960, 9 years later and
the time of our next thorough sur-

Fig. 1.- — Cross section of abandoned cis¬
tern which houses Plecotus rafinesquii.
The position of the 3 large clusters, X,
and single individuals, s, of bats as on
March 21, 1960, is shown. The black
square indicates one square foot. Draw¬
ing by H. C. Henrikson.

ATey, the largest concentration of bats
ever noted, 64 in all, Avas present.

Nine bats AArere taken from the cis¬
tern to Cincinnati, Ohio, on August
4, 1957, banded, and released. This
Avas 345 miles away from the cistern.
None Avas ever recovered back at the
cistern or, so far as Ave knoAv, any-
Avere else.

Six of the eleATn big-eared bats
taken in the cistern on September 2
and preserved as skins and skulls
appear to be young of the year. TIoav-
ever, these young, thought to be
about 3 months old, haAye the epiphy¬
ses of the fingers nearly closed and
are as large as winter-taken speci¬
mens. These 6 differ from adults

Colony of Big-Eared Bats

primarily in having’ shorter fur with
the hairs on the back lacking the
glossy brown tips. An age of 3
months is estimated for these young
on the basis of specimens in the col¬
lection from Mammoth Cave, Ken¬
tucky, one of which appears to be
about 2 weeks old, collected June 17,
and another taken on July 6 which
could be about 5 weeks old. The
birth date for both of these was near
June 1.

Twenty-eight adults (14 males, 14
females) from the one locality near
Reelfoot Lake, only 164 miles south¬
west of the type locality of Plecotus
rafinesquii rafinesquii, provide the
following averages and extremes, all
in millimeters. Measurements of the
skull are taken according to Handley
(1959 :98 ) , with the males listed first.
Total length, 97.8 (94-102), 99.6
(93-105) ; tail length, 49.8 (46-53),

50.3 (47-54) ; hind foot length, 10.7

(9-12), 11.2 (9.5-12) ; ear from

notch, 33.5 (29-35), 34.1 (27-37);
length of forearm, 43.0 (41.8-44.4),

43.3 (41.7-45.0) ; tragus (as taken

by collector), 12.8 (10-16), 13.6 (11-
16) ; greatest length of skull, 16.14
(15.8-16.5), 16.35 (16.1-16.65) ; zygo¬
matic breadth, 8.80 (8. 4-9.1), 8.87
(8.5-9.15) ; interorbital breadth,
3.57 (3.45-3.7), 3.60 (3.45-3.7);

breadth of braincase, 7.85 (7. 6-8. 8),
7.90 (7. 7-8.1) ; depth of braincase,
6.03 (5.9-6.25), 6.03 (5.85-6.25);

maxillary toothrow length, 5.32
(5.25-5.45), 5.35 (5. 2-5. 5) ; postpala¬

tal length, 5.98 (5. 7-6.1), 6.04 (5.8-
6.4) ; palatal breadth, 6.12 (5.85-
6.3), 6.15 (6.0-6.35). All specimens
have a well-developed secondary cusp
present on the first upper incisor
except for one specimen. In this,
the incisor is much as in Plecotus
townsendii.

Summary

Big-eared bats, Plecotus rafine¬
squii, may use as roosting places cer¬
tain cavities that are well lighted
and become cold during the winter.
Such is the case for big-eared bats
inhabiting an abandoned, unused,
uncovered cistern in westernmost
Tennessee. Only Plecotus rafinesquii
occupies this cistern and in numbers
varying from 1 to 64 individuals.
They are present in winter, spring,
and summer, and probably through¬
out the year. During the winter,
bats move in and out of the hiber¬
nating chamber, with individuals
from different localities taking places
of those moving out. The same indi¬
viduals may be present for parts of
two consecutive winters, but not

continuouslv in the winter nor in
«/

the summer. When only about 3
months old, big-eared bats are adult-
size.

Literature Cited

Handley, C, O., Jr. 1959. A revision of
American bats of the genera Enderma
and Plecotus. Proc. U. S. Nat. Mus.,
110:95-246.

Manuscript received May 17, 1962.

A SECOND PORCUPINE RECORD FOR ILLINOIS

PAUL W. PARMALEE
Illinois State Museum, Springfield

Approximately 3,200 bones and
over 33,000 freshwater mussel valves
(37 species) were identified from the
faunal samples recovered at three
Archaic Indian sites situated along
the Wabash River in east-central
Illinois. The field work at these sites
(Riverton, Swan Island, Robeson
Hills) took place during April, Majq
June and October, 1961, and was
supervised by Mr. Howard D. AVin-
ters of the Illinois State Museum.
The sites were spaced nearly equal
distance (10 miles) from one an¬
other : Riverton, two miles northeast
of Palestine, Crawford County, and
Swan Island, located at the present
Crawford-Lawrence county line were
situated on the river flood plain,
while the Robeson Hills village (two
miles north of Vincennes, Indiana,
Lawrence County, Illinois) occurred
on the river bluff (isolated aggraded
upland). Artifact assemblages, the
faunal complex and radio-carbon
dates suggest that the sites were oc¬
cupied by the same group at differ¬
ent seasons or by contemporaneous
groups.

Fish remains were more abundant
at the Riverton and Swan Island
sites, those of catfishes, bowfin and
freshwater drum being the most
numerous. Turtles had been taken
in considerable numbers (six species
represented). Although a minimum
of 17 species of birds were identified,
only the turkey had been utilized
extensively. Of all vertebrate groups,
mammals (22 species) were the most

important source of food to these
peoples, the white-tailed deer con¬
stituting the basic meat staple in
their diet. Numerous bones of rac¬
coon, gray squirrel, beaver, cotton¬
tail, muskrat and other small mam¬
mals from these sites were indicative
of former populations and of their
extensive use by these Archaic
groups.

Recovery of one skull and four
mandible sections of the porcupine
( Erethizon dorsatum) from the Riv-
ton Site constitutes a noteworthy
zoological record for the area. There
are no historic accounts which defi¬
nitely establish the porcupine as a
former resident of Illinois and, previ¬
ous to the recovery of the Riverton
Site material, no remains have been
found in any of the other numerous
archaeological sites investigated thus
far. The first record to indicate a
former population of porcupine in
Illinois consisted of several cranial
bones and jaws recovered from a nat¬
ural cave in Monroe County (Parma-
lee, Bieri and Mohrman 1961). This
isolated find probably represents an
early southern range extension of
this rodent along the Mississippi
River bluffs.

The Riverton Site material con¬
sisted of portions of four lower man¬
dibles (three right jaws, consequent¬
ly at least three individuals
represented) and the right maxilla
containing the premolar and molars
1 and 2. Two jaws contained the pre¬
molar and molars 1 and 2 ; one con-

Porcupine Record

tained all three molars, and the
fourth jaw retained only molars 2
and 3. Fragments of an incisor and
the proximal end of a radius from
this site may also be referable to
E. dorsatum. Two jaws found in
the 36-42 inch level were dated,
based on C 14 tests of charcoal from
the levels, at 3,100 ± 200 years B.P. ;
the date from the 60-66 inch level,
in which a third jaw was recovered,
was 3,200 ± 200 years B.P.

The question arises as to whether
the Indians obtained porcupines on
the Illinois or Indiana side of the
Wabash River. Lyon (1936) lists
the published records with the dates
of observation and the specimens in
collection of the porcupine in Indi¬
ana ; three were from counties (Vigo,
Knox, Posey) bordering the Wabash
River. There was no record of E.
dorsatum from Sullivan County, In¬
diana, the county immediately east
of Crawford County, Illinois, and
the Riverton Site. This rodent was
recorded from \rigo and Knox coun¬
ties which border Sullivan County
on the north and south, respectively,
and it could have been quite possible
for the Indians to have easily cov¬
ered these short distances (minimum
of about 15 miles) while hunting.
However, the Archaic inhabitants of
the Swan Island and Robeson Hills
sites were closer to the porcupine’s

known range in Indiana (Knox
County, directly across the river),
yet no remains of it were recovered
at either site.

Rising over 150 feet above the Wa¬
bash River flood plain, Merom Bluff
(about 1 Yj miles northeast of the
Riverton Site, Sullivan County, In¬
diana) is a 2i/2 mile long section of
wooded, sandstone bluff that could
have served as suitable habitat for
the porcupine. Approximately 2%
miles south of the Riverton Site
(Crawford County, Illinois), and
paralleling the river for about 1%
miles, is another wooded bluff (over
120 feet above the flood plain) that
may also have been inhabited by
this animal. Future archaeological
excavations throughout the lower
Wabash River Valley may establish
conclusively the presence of a pre¬
historic population of porcupine on
the Illinois side of the river. How¬
ever, these remains of E. dorsatum
from the Riverton Site are indica¬
tive of its former occurrence locally
in the Crawford County, Illinois —
Sullivan County (?), Indiana area.

Literature Cited

Lyon, Marcus W., Jr. 1936. Mammals
of Indiana. Amer. Mid. Nat., 17(1):
384 pp.

Parmalee, Paul W., R. A. Bieri and
R. K. Moiirman. 1961. Mammal re¬
mains from an Illinois cave. Jonr.
Mamm., 42(1): 119 pp.

GEOGRAPHIC AND HOST DISTRIBUTION OF BLOOD
PARASITES IN COLUMBORID BIRDS

NORMAN D. LEVINE

University of Illinois, Urbana

Synopsis

AArailable data on geographic dis¬
tribution and incidence of blood
parasites in different host species of
columborid birds in the wild are
assembled from 174 reports in the
literature, analyzed, and informa¬
tion on life cycles and pathogenesis
is given. The most common genus is
Haemoproteus, of which two species
occur in naturally affected birds.
The more common species is H.
columbae, the gametocytes of which
extend along one side of the host
erythrocyte and curve around its
ends. It has been found in 14 host
species in 24 countries. II. sacha-
rovi has gametocytes which com¬
pletely fill the host cell when mature.
It has been found in 3 host species
in the United States and Italy.

Trypanosoma avium has been re¬
ported from 7 host species in 6 coun¬
tries.

Three species of Plasmodium have
been reported. P. relict um is by far
the most common ; it has been found
in 3 host species in 5 countries.
P. elongatum lias been found in 1
host species ( Zenaidnra macroura )
in the United States. P. hexamerium
lias also been found in Z. macroura
in the United States and possibly
also in Columbigallina talpacoti in
Colombia.

Leucocytozoon marchouxi, which
has rounded gametocytes, has been
reported from 13 host species in 15
countries. A species of Leucocyto¬

zoon with elongate gametocytes has
been seen once in Uganda. Leucocy¬
tozoon has not been reported from
South or Central America despite
the fact that a number of surveys
have been made there.

Toxoplasma gondii has been iden¬
tified bv mouse inoculation or dve
«/ «/

test or both in 8% of 176 domestic
pigeons in 4 surveys in different
parts of the United States. What
may have been Toxoplasma, Lanke-
sterella or possibly some other genus
has been found in 3 host species in
4 countries. The possible role of the
domestic pigeon as a reservoir of
Toxoplasma gondii is mentioned.

The distribution of blood protozoa
in columborid birds is poorly known.
Haemoproteus has been reported
from only 20% of the known 61
genera and 8% of the known 320
species of these birds, Plasmodium
has been found in 11% of tlm genera
and 3% of the species, Leucocytozoon
in 10% of the genera and 4% of the
species, Trypanosoma in 8% of the
genera and 2% of the species, and
Toxoplasma or a similar form in 3%
of the genera and 1% of the species.
The parasite species has been iden¬
tified in only 58% of 138 reports of
Haemoproteus, 62% of 26 reports of
Plasmodium, 35% of 26 reports of
Leucocytozoon, 60% of 10 reports of
Trypanosoma, and 77% of 13 re¬
ports of Toxoplasma and similar
forms. In 43% of the total of 213
records. Hie species name of the para¬
site was not given. In only 41 (24% )

Blood Parasites of Columborid Birds

of the 173 papers were 10 or more
birds of a single species examined,
and in only 12 (7%) of them were
100 or more birds examined.

Introduction

Our present information on the
blood parasites of birds of the order
Columborida is, with a few excep¬
tions, both scattered and scanty. It
has been tabulated by Levine and
Kantor (1959), and the analyses in
the present paper are based pri¬
marily on the raw data in their
tables. Additional information from
other papers has been included as
indicated. The only references given
at the end of this paper are to the
papers specifically discussed in the
text ; all others and the sources of
all data can be found in Levine and
Kantor (1959). Records of blood
parasites from birds in zoos have
not been tabulated, since they do
not necessarily mirror the true situ¬
ation in the wild. The surveys dis¬
cussed in the text are for the most
part only those in which 100 or more
birds were examined. The results of
surveys in which fewer birds were
examined are given in the tables.

Trypanosoma. Trypanosomes
were first found in columborid birds
by Novy and MacNeal (1905a, b) in
mourning doves ( Zenaidura macrou-
ra ) in Michigan ; they thought that
they were probably Trypanosoma
avium. De Mello and Bras de Sa
(1916) found a trypanosome which
they named Trypanosoma hannai in
domestic pigeons ( Columba livia ) in
Portuguese India, and Sergent
(1941a, b) found a trypanosome
which he named T. oenae in Oena
capensis in the Nigerian Sahara.

Wood and Herman (1943) and Coat-
n ey and West (1938) used the name
T. avium for the trypanosomes which
they found in Zenaida asiatica
mearnsi in Arizona and in Zenaidura
sp. in Nebraska, respectively.

Trypanosoma avium Danilewsky,
1885 was first described from owls
(scientific name not given) and
roller-birds ( Coracias garrulus ) in
Europe, and has since been reported
from a wide variety of birds. Vari¬
ous other names have been given to
trypanosomes in other birds, largely
on the basis of their presence in a
different host from those previously
reported. The validity of most of
these names, including T. hannai
and T. oenae, is highly questionable.

Bennett (1961) studied the mor¬
phology of trypanosomes from 25
species of birds belonging to 11 fam¬
ilies (but not including any colum¬
borid birds), and concluded that
they were all the same species except
for a trypanosome which he found
once in a chipping sparrow ( Spizella
passerina ) ; he considered the chip¬
ping sparrow form to be Trypano¬
soma paddae Laveran and Mesnil,
1904 and all the others to be T.
avium.

Cross transmission studies on
avian trypanosomes have been car¬
ried out by Baker (1956a, b) and
Bennett (1961). The former trans¬
mitted T. avium from the rook ( Cor -
vus frugilegus) and jackdaw (C.
monedula) to canaries, but failed to
transmit it to a single 3-day-old
chick. Bennett (1961) transmitted
strains of T. avium by means of
simuliids or Aedes aegypti from 9
species of birds belonging to 4 orders
into 15 species of birds belonging
to 5 orders. Many of these trans-

Transactions Illinois Academy of Science

missions were from birds of one
order to those of another. Bennett
made no attempt to infect every
receptor host species with a strain
from every donor host species, but
used enough combinations to show
that it would have been possible.
Among them, he infected the pigeon
with strains from the blue jay and
saw-whet owl.

On the basis of the above morpho¬
logic and cross-transmission studies,
it is safe to conclude that Trypano¬
soma avium occurs widely in many
different orders of birds and that,
unless they are proven to be differ¬
ent, all avian trypanosomes should
be referred to this species.

Blood-sucking arthropods such as

blackflies, mosquitoes and hippo-
boscid flies are the vectors of avian
trypanosomes. Baker (1956a, b)
worked out the life cycle of T. avium
from rooks and jackdaws. He found
that in England the hippoboscid fly,
Ornithomyia avicularia, acts as the
vector and that birds become in¬
fected when they eat insects which
have become infected by sucking-
blood. According to Baker, there
is no multiplication in the avian
host, the trypanosomes simply be¬
coming larger; multiplication takes
place only in the arthropod host.

Bennett and Fallis (1960) com¬
pared the incidence and level of
parasitemia of Trypanosoma with
the occurrence and feeding habits of

Table 1. — Known Geographic Distribution and Prevalence of Trypayiosoma

in Columborid Birds.

Prevalence (%)

Surveys in

Present but

which more

which 100

no reliable

than 10

or more

figures on

birds were

Country

Bird Species

prevalence

examined

EUROPE

Germany .

Columba palumbus .

ASIA

Portuguese India. .

Columba livia .

West Java .

Streptopelia chinensis ....

AFRICA

Nigerian Sahara. . .

Oena capensis .

Gambia .

Streptopelia vinacea .

NORTH AMERICA

Arizona .

Zenaida asiatica mearnsi.

U. S. A .

Zenaidura macroura

carolinensis .

0.5

Michigan .

Zenaidura macroura

carolinensis .

Nebraska .

Zenaidura macroura

carolinensis .

Blood Parasites of Col urn bo rid Birds

Table 2. — Known Geographic Distribution and Prevalence of Plasmodium

in Columborid Birds.

Prevalence (%)

Surveys in

Present but

which more

which 100

no reliable

than 10

or more

Plasmodium

figures on

birds were

Country

Bird Species

Species

prevalence

examined

EUROPE

Germany .

Columba palumbus

relictum .

Czechoslovakia. . . .

Columba palumbus

sp .

ASIA

Japan .

Streptopelia

orientalis .

sp .

AFRICA

Egvpt .

Columba livia .

relictum .

French Congo. . . .

Columba livia ....

relictum .

Algeria .

Streptopelia

turtur .

sp .

Belgian Congo ....

Treron calva .

sp .

NORTH AMERICA

U.S. A .

Columba livia ....

relictum .

California .

Columba livia ....

relictum .

Iowa .

Columba livia. . . .

relictum .

Arizona .

Zenaida asiatica

mearnsi .

sp .

U.S. A .

Zenaidura

macroura .

elongatum ....

U.S. A .

Zenaidura

macroura .

relictum .

U.S. A .

Zenaidura

macroura .

sp .

0.5

0 . 5

Illinois .

Zenaidura

macroura .

hexamerium . .

Nebraska .

Zenaidura

macroura .

relictum .

D. C. & vicinity. . .

Zenaidura

macroura .

sp .

California .

Zenaidura

macroura .

relictum .

Mexico .

Leptotila

verreauxi .

sp .

SOUTH AMERICA

Colombia .

Columba

cayennensus ....

sp .

Uruguay .

Columba livia ....

relictum .

Colombia .

Columbigallina

talpacoti .

hexamerium . .

(?)

Transactions Illinois Academy of Science

various ornithophilic flies in Algon¬
quin Park, Canada, and concluded
that the vectors are probaly simu-
liids. Their study did not include
any columborid birds. Bennett
(1961) found that several species
of simuliid flies and also Aedes
aegypti could act as vectors of T.
avium. Leptomonad, crithidial and
metacyclic trypanosome forms devel¬
oped in the midgut and hindgut of
the simuliids, and infection took
place when trypanosomes which had
been passed in the feces entered the
host thru breaks in the skin pro¬
duced by the feeding flies. Bennett ’s
experiments indicated that birds
could be infected by eating flies only
if the insects were crushed enough
to release flagellates from the hind-
gut into the birds’ mouths.

Nothing is known of the patho¬
genicity of avian trypanosomes. They
are presumably non-pathogenic.

The known geographic distribu¬
tion and prevalence of T . avium in
columborid birds are shown in Table
I. Only three surveys have been
reported in which 100 or more birds
were examined, and only four in
which more than 10 birds were ex¬
amined. Boing (1925) found Try¬
panosoma sp. in 24% of 128 Columba
palumbus in Germany; Ivraneveld
and Mansjoer (1954) reported it in
8% of 2100 Streptopelia chinensis
tigrina in West Java; Huff (1939)
reported it in 0.5% of 188 Zenaidura
macroura carolinensis in the United
States; and Wood and Herman
(1943) reported Trypanosoma avium
in 8% of 12 Zenaida asiatica mearnsi
in Arizona. (As the result of a
printing error, the host species was
listed by Levine and Ivantor (1959)
as Zenaidura macroura ; actually,

Wood and Herman found no trypa¬
nosomes in the 27 Z . macroura which
the}7 examined.)

To these surveys should be added
several in which no trypanosomes
were found, such as that of Hanson
ei al. (1957) in mourning doves in

Illinois : these can be identified bv

«/

comparison with the data on Haemo-
proteus (Table 3).

Plasmodium. Three species of
Plasmodium have been reported
from naturally infected columborid
birds in the wild. P. relict urn is by
far the most common. Boing (1925)
found it in 48% of 128 Coin tuba
palumbus in Germany; Herman et
al. (1954) found it in 5% of 43
Columba livia and 1% of 383 Zenai¬
dura macroura in Kern County,
California ; Mathey (1955) found it
in 3 C. livia in the Sacramento area
of California ; Becker, Hollander and
Pattillo (1956) found it in 20% of
15 C. livia in Iowa; and Coatney
(1938) found it in 9% of 11 Zenai¬
dura macroura in Nebraska. In ad¬
dition, it has been reported from
Columba livia in Egypt, the French
Congo and Uruguay.

Plasmodium elongatum was found
in 1% of 188 Zenaidura macroura
carolinensis from the United States
by Huff (1939), and P. hexamerium
was found in 1% of 134 Z. macroura
carolinensis in Illinois by Huff
(1935). Renjifo-Salcedo, Sanmartin
and Zulueta (1952) found a Plas¬
modium which they thought might
be P. hexamerium in 17% of 6 Co-
lumbigallina talpacoti in Colombia.

In addition to the above 3 species,
mention should be made of the form
which Carini (1912) described under
the name Plasmodium columbae from
a pigeon which had died of toxoplas-

Blood Parasites of Columborid Birds

mosis following’ experimental infec¬
tion with a canine strain of Toxo¬
plasma gondii. Its gametocytes were
halter-shaped and resembled those of
Haemoproteus columbae ; in addi¬
tion to these, Carini found many
round, oval or halter-shaped para¬
sites with slender cytoplasmic ex¬
tensions or pseudopods. Carini was
not sure that he was actually deal¬
ing’ with a Plasmodium, since he saw
no schizonts. In the absence of con¬
firmation in the ensuing 49 years,
Plasmodium columbae cannot be ac¬
cepted as a valid species or even as
a Plasmodium. I consider the name
a nomen nudum.

In addition to the named species,
Plasmodium sp. has been found in
Coi n m b a palumbus in Czecho¬
slovakia, in 13% of 111 Streptopelia
orient alis in Japan by Og’awa
(1912). in Streptopelia turtur in
Algeria, in Treron calva in the Bel¬
gian Congo, in Zenaida asiatica
mearnsi in Arizona, in 0.5% of 188
Zenaidura macroura carolinensis in
the U. S. by Huff (1939), in Lepto-
tila vereauxi in Mexico, and in Co¬
lumba cayennensis in Colombia.

Wolf son (1937, 1940) infected do¬
mestic pigeons ( Columba livia) ex¬
perimentally with P. cathemerium,
and Huff et al. (1950) and Huff and
Marchbank (1955) infected domestic
pigeons experimentally with P. fal-
lax.

The known geographic distribution
and prevalence of Plasmodium in
columborid birds are shown in Table
2. The results of surveys in which
100 or more birds were examined
have been given above. There have
been only 5 ; in Germany, Japan,
California, Illinois and the U. S. in

general. To these should be added
several surveys in which no Plas¬
modium was found, such as that of
Hanson et al. (1957) in mourning
doves in Illinois ; these can be iden¬
tified by comparison with the data
on Haemoproteus (Table 3).

In sum, Plasmodium relict inn has
been found in the wild in Columba
livia, C. palumbus and Zenaidura
macroura. It has also been reported
in zoos in Columba argentia, Ducula
concinna, Leptotila crumeniferus,
Megaloprepia magnifica Scardafella
squammata, Tympanistria bicolor,
and T. tympanistria. Plasmodium
elongatum has been reported only
from the mourning dove, Zenaidura
macroura. Plasmodium hexamerium
has been reported with certainty
from Z. macroura and questionably
from Columbigallina talpacoti. Un¬
identified species of Plasmodium
have been reported in Columba
palumbus, C. cayennensis, Leptotila
verreauxi, Streptopelia orient alis, S.
turtur, Zenaida asiatica mearnsi and
Zenaidura macroura in the wild, and
in Columba squamosa, and Tympa¬
nistria tympanistria in zoos. In all,
Plasmodium lias been found in a
total of 10 identified species of co¬
lumborid birds in the wild, and in 9
others in zoos.

Haemoproteus. This is by far the
most common genus of blood proto-
zoon in columborid birds. Two spe¬
cies occur in naturally infected birds
in the wild. The more common one
is H. columbae, which has so-called
halter-shaped gametocytes which ex¬
tend along one side of the host cell
nucleus and curve around its ends.
This species was originally described
by Celli and Sanfelice (1891a, b)
from the domestic pigeon and has

Transactions Illinois Academy of Science

Table 3. — Known Geographic Distribution and Prevalence of Haemoproteus*

in Columborid Birds.

Prevalence (%)

Surveys in

Present but

which more

which 100

no reliable

than 10

or more

figures on

birds were

Country

Bird Species

prevalence

examined

EUROPE

Greece .

Columba livia .

Italy .

Columba livia .

Spain .

Columba livia .

Italy .

Columba oenas .

Germany .

Columba palumbus .

England .

Columba palumbus .

Greece .

Streptopelia decaocto .

Italy .

Streptopelia turtur .

Spain .

Streptopelia turtur .

Germany .

Streptopelia turtur .

ASIA

Lebanon .

Columba livia .

“Plains,” India. . . .

Columba livia .

100

Delhi, India .

Columba livia .

Portuguese India . .

Columba livia .

Manila, Philippines

Columba livia .

Wellesley Province

India .

Columba sp .

too

Palestine .

Columba sp .

Philippines .

Columba sp .

Mukteswar, India. .

Sphenurus sphenurus . . . .

Formosa .

Streptopelia chinensis. . . .

Tonkin, Vietnam. .

Streptopelia tranquebarica

India .

Streptopelia turtur .

AFRICA

French Sudan .

Columba guinea .

Algeria .

Columba livia .

Egypt .

Columba livia ....

French Morocco. . .

Columba livia .

French Congo .

Columba livia .

Belgian Congo. . . .

Columba livia .

Portuguese Guinea .

Columba livia .

Un. S. Africa .

Columba livia .

Ethiopia .

Gena capensis .

Un. S. Africa .

Gena capensis .

Mozambique .

Plectopterus gambiensis . .

Un. S. Africa .

Streptopelia capicola. . .

Ethiopia .

Streptopelia decipiens ....

Belgian Congo. . . .

Streptopelia semitorquata .

Liberia .

Streptopelia semitorquata .

French Sudan .

Streptopelia senegalensis .

Gambia .

Streptopelia senegalensis .

Algeria .

Streptopelia turtur .

Blood Parasites of Columborid Birds

Table 3. — Continued

Country

Bird Species

Prevalence (%)

Present but
no reliable
figures on
prevalence

Surveys in
which more
than 10
birds were
examined

Surveys in
which 100
or more
birds were
examined

French Morocco. . .

Streptopelia turtur .

French Sudan .

Streptopelia vinacea .

Gambia .

Streptopelia vinacea .

Belgian Congo. . . .

Treron calva .

Gambia .

T reron calva .

NORTH AMERICA

Ariz. & Calif .

Columba fasiata .

Colorado .

Columba fasciata .

California .

Columba livia .

D.C., Md., Va .

Columba livia .

Florida .

Columba livia .

Hawaii .

Columba livia .

Iowa .

Columba livia .

Iowa .

Columba livia .

15**

Nebraska .

Columba livia .

22**

Pennyslvania. .

Columba livia .

South Carolina. . . .

Columba livia .

U.S.A .

Columba livia . .

100

U.S.A .

Columba livia .

_j_ **

Mexico .

Columba livia .

Arizona .

Zenaida, asiatica .

Mexico .

Zenaida asiatica. .......

Ariz. & Calif .

Z enaidura macroura .

Ariz. & Calif .

Zenaidura macroura .

41**

Northern Calif .

Z enaidura macroura .

Northern Calif .

Zenaidura macroura .

Southern Calif .

Zenaidura macroura .

D. C. & vicinity. . .

Zenaidura macroura .

+**

Georgia .

Zenaidura macroura .

Georgia .

Zenaidura macroura .

Illinois .

Zenaidura macroura .

Illinois .

Zenaidura macroura .

25-43

Illinois .

Zenaidura macroura .

-i_ **

Illinois .

Zenaidura macroura .

43-58**

Massachusetts .

Zenaidura macroura .

y**

Massachusetts .

Zenaidura macroura .

Michigan .

Zenaidura macroura .

Michigan .

Zenaidura macroura .

_p**

Nebraska .

Zenaidura macroura .

Nebraska .

Zenaidura macroura .

Nebraska .

Zenaidura macroura .

_i_**

Nebraska .

Zenaidura macroura .

67**

Texas .

Zenaidura macroura .

Texas .

Z enaidura macroura .

27**

Texas .

Zenaidura macroura .

U.S.A. (mostly

Illinois) .

Zenaidura macroura .

U.S.A. (mostly

Illinois) .

Zenaidura macroura .

56**

100

Transactions Illinois Academy of Science

Table 3. — Concluded

Country

Bird Species

]

Present but
no reliable
figures on
prevalence

Prevalence (%

Surveys in
which more
than 10
birds were
examined

)

Surveys in
which 100
or more
birds were
examined

SOUTH AND CEN

TRAL AMERICA

Colombia .

Columba cayennensis ....

Colombia .

Columba cayennensis ....

Brazil .

Columba livia .

Brazil .

Columba livia .

Brazil .

Columba livia .

French Guiana. . . .

Columba livia, .

Uruguay .

Columba livia .

Argentina .

Columba picazuro .

Brazil .

Columba picazuro .

Brazil .

Columba rufina .

Brazil .

Columba rufina .

French Guiana. . . .

Columba rufina .

Brazil .

Columba sp .

Brazil .

C olumbigallina talpacoti. .

Colombia .

Columbigallina talpacoti. .

Venezuela .

C olumbigallina talpacoti..

Argentina .

Columbigallina picui . . . .

Brazil .

Columbigallina picui. . . .

Brazil .

Leptoptila sp .

Brazil .

Scardafella squammata. . .

El Salvador .

Zenaida asiatica .

Argentina .

Zenaidura auriculata ....

Colombia .

Zenaidura auriculata ....

AUSTRALIA .

Ptilinopus superbus .

* H. columbae unless otherwise indicated.
** H. sacliarovi.

since been found in many other co-
lumborid birds.

Separate specific names have been
given to morphologically indistin¬
guishable forms in some hosts : II.
maccallumi by Novy and MacNeal
(1905a) to the form in Zenaidura
macroura, H. melopeliae by Laveran
and Petit (1909) to the form in
Zenaida asiatica, H. turtur by Co-
valeda Ortega and Gallego Beren-
gner (1950) to the form in Strepto-

pelia turtur, and H. vilhenai by
Santos Dias (1953) to the form
in Plectopterus gambiensis. Huff
(1932) transmitted H. maccallumi
from the mourning dove to the do¬
mestic pigeon, but Coatney (1953)
was unable to transmit H. columbae
from the pigeon to the mourning
dove. Both used the hippoboscid
fly, Pseudolynchia canariensis, as
the vector. There may be strain
differences between the different

Blood Parasites of Columborid Birds

101

hosts, but until greater differences
than these are brought out, it is
probably best to use the name H.
columbae for all species with halter¬
shaped gametocytes from columborid
birds.

Another name, H. danilewskyi,
was used by some earlier authors
for H. columbae in columborid birds.
This name was originally given by
Kruse (1890) to the species in the
crow, Corvus cornix, and should not
be used for parasites of birds of other
orders in the absence of proof that
they are the same.

One reservation should be kept in
mind regarding reports of halter¬
shaped Haemoproteus from birds.
This is that in some cases these pro¬
tozoa may not be Haemoproteus at
all but a Plasmodium species with
halter-shaped or elongate gameto¬
cytes, such as P. fallax or P. circum-
flexum, which does not have schi-
zonts in the blood at the time of
examination.

Haemoproteus columbae was
found by Giovannoni (1946) in 58%
of 159 C. livia in southern Curitiba,
Brazil ; by Singh, Nair and David
(1951) in 22% of 214 C. livia in
Delhi, India; by Huff (1939) in
47% of 188 Z. macroura in the
United States (mostly in Illinois) ;
by Couch (1952) in 56% of 213
Z. macroura in Texas; and by Han¬
son et at. (1957) in 30% of 392 im¬
mature Z. macroura and in 43% of
72 adult Z. macroura in Illinois.

In addition to the above studies
in which at least 100 birds of each
species were examined, H. columbae
has been found by various workers
in Columba fasciata in Arizona and
California, in Columba guinea in
the French Sudan, in Columba livia

in various parts of the world, in
Columba oenas in Italy, in Columba
rufina in Brazil, in Columbigallina
talpacoti in Venezuela, in Plectop-
terus gambiensis in Mozambique, in
Streptopelia senegalensis and S.
vinacea in the French Sudan, in
S. turtur in French Morocco, in
Zenaida asiatica in Arizona and El
Salvador, in Zenaidura macroura on
Cape Cod and in Arizona, California
and Nebraska, and in Ptilinopus
iozonus and Turtur brehmeri in
zoos.

The second species, Haemoproteus
sacharovi, was first described by
Novy and MacNeal (1904a, b) in the
mourning dove, Zenaidura macrou¬
ra, in Michigan. Its gametocytes
differ from those of most species of
Haemoproteus in that when mature
they completely fill the host erythro¬
cyte, enlarging and distorting it, and
often pushing the host cell nucleus
to the edge of the cell.

H. sacharovi has been found in
both mourning doves and domestic
pigeons. It was found by Huff
(1939) in 56% of 188 Z. macroura,
mostly from Illinois ; by Couch
(1952) in 27% of 213 Z. macroura
in Texas ; and by Hanson et al.
(1957) in 58% of 392 immature and
43% of 72 mature Z. macroura in
Illinois.

In addition to the above studies
in which at least 100 birds of each
species were examined, H. sacharovi
has been found by various workers
in Columba livia in Iowa and Ne¬
braska and in Zenaidura macroura
on Cape Cod and in Arizona, Cali¬
fornia and Nebraska.

What was almost certainly the
same species was described by Fran-
chini (1924) in 3% of 36 Strepto-

102

Transactions Illinois Academy of Science

pelia tnrtur in Italy. He called his
form Leucocytozoon sp., but his de¬
scription and figures fit H. sacharovi
better than they do Leucocytozoon.

The above findings refer to studies
in which the species of Haemopro-
teus was named. In addition, Hae-
moproteus sp. lias been reported
without further identification in Co¬
lumba cayennensis in Colombia, C.
picazuro and Columbina picui in
Argentina and Brazil, Oena capensis
in South Africa and Ethiopia,
Ptilinopus superbus in Australia,
Scardafella squammata in Brazil,
Sphenurus sphenurus in India,
Streptopelia capicola in South Afri¬
ca, S. chinensis in Formosa, S. de-
caocto in Greece, S. decipiens in
Ethiopia, S. semitorquata in the Bel¬
gian Congo and Liberia, S. tran-
quebarica in Tonkin, Treron calva
in the Belgian Congo and Gambia,
Zenaidura auriculata in Argentina
and Colombia, and in Capoenas
nicobarica, Columba argentina, Co-
lumbigallina passerina, Geophaps
smithii , Leptoptila crumeniferus ,
Megaloprepia magnifica, Ptilinopus
melanospila, P. periatus, P. wallacei,
Treron curvirostra, T. delalandi,
Turacoena manadensis and Tympa-
nistria tympanistria in zoos.

The known geographic distribu¬
tion and prevalence of Haemopro-
teus in columborid birds are shown
in Table 3. The results of surveys
in which 100 or more birds were ex¬
amined have been given above. There
have been only 8 — of Columba livia
in Hawaii, Brazil and Delhi, India,
of C. palumbus in Germany, of Co-
lumbina picui in Argentina, and of
Zenaidura macroura in Illinois, Illi¬
nois and other states, and Texas.

There have been 32 surveys in

•/

which 10 or more birds were ex¬
amined. They involved 11 species of
columborid birds in 11 countries,
including Columba cayennensis in
Colombia, C. fasciata in Colorado,
C. livia in the United States (Flori¬
da, Hawaii, Iowa, Nebraska), Brazil,
Italy, Greece, India, French Morocco
and the Philippines, C. oenas in
Italy, C. palumbus in Germany and
French Morocco, C. rufina in Brazil,
Columbina picui in Argentina,
Streptopelia decaocto in Greece, S.
turtur in Italy, Spain and French
Morocco, Zenaida asiatica in Ari¬
zona, and Zenaidura macroura in
the United States (Arizona, Cali¬
fornia, Georgia, Illinois, and other
states, Massachusetts, Nebraska,
Texas) .

The only study of the relation of
age to prevalence of Iiaemoproteus
was that of Hanson et al. (1957) in
Zenaidura macroura. They found
H. columbae in 30% of 392 immature
and 43% of 72 adult birds in Illinois.
The incidence of this species in the
immature birds increased steadily
with age, from 7 to 8% in very young
birds to 70% in older ones. The lat¬
ter rate wTas higher than that in the
adults.

H. sacharovi was present in 58%
of the immature and 43% of the
adult birds. Its incidence in the
immature birds did not increase
nearly so sharply with age as did
that of H. columbae. It was present
in 31% of the very young doves, and
its incidence fluctuated between 52%
and 69% in older immature birds.

Hanson et al. (1957) also studied
the incidence of Haemoproteus in
different years from 1948 thru 1954
and in different parts of Illinois. It
varied markedly in both categories.

Blood Parasites of Columborid Birds

103

Table 4. — Known Geographic Distribution and Prevalence of Leucocytozoon*

in Columborid Birds.

Country

Bird Species

Prevalence (%)

Present but
no reliable
figures on
prevalence

Surveys in
which more
than 10
birds were
examined

Surveys in
which 100
or more
birds were
examined

EUROPE

Germany .

Columba palumbus .

England .

Columba palumbus .

Corsica .

Streptopelia turtur .

Italy .

Streptopelia turtur .

Spain .

Streptopelia. turtur .

100

ASIA

Mukteswar, India..

Sphenurus sphenurus ....

Japan .

Streptopelia orientalis ....

Tonkin, Vietnam . .

Streptopelia tranquebarica

AFRICA

Pretoria, Un. S.

Africa .

Columba livia .

French Morocco. . .

Columba palumbus .

Transvaal, Un. S.

Africa .

Oena capensis .

Pietermaritzburg,

Un. S. Africa. . . .

Streptopelia capicola .

Uganda .

Streptopelia semitorquata .

_j_ **

Upper Senegal and

N igeria .

Streptopelia senegalensis. .

French Morocco. . .

Streptopelia turtur .

NORTH AMERICA

Ariz. & Calif .

Columba fasciala .

Colorado .

Columba faseiata .

California .

Streptopelia chinensis ....

Ariz. & Calif .

Zenaidura macroura .

D. C. & vicinity. . .

Zenaidura macroura .

Georgia .

Zenaidura macroura .

Illinois .

Zenaidura macroura .

1 2***

j 9***

Illinois .

Zenaidura macroura .

65****

OTHER

Mauritius .

Geopelia striata .

* Leucocytozoon marchouxi unless otherwise indicated.
** Leucocytozoon sp with elongate gametocytes.

*** Adults.

**** Immature birds.

104

Transactions Illinois Academy of Science

That of Id. sacharovi in the immature
birds varied from 45% in 1948 to
78% in 1954; in the adults it ranged
from 20% in 1953 to 75% in 1951,
but these latter figures are based on
insufficiently large samples. The in¬
cidence of H. columbae in the im¬
mature birds ranged from 6% in
1950 and 1954 to 43% in 1952; in
the adults it ranged from 30% in
1949 and 1950 to 75% in 1952, but
this last figure is based on too small
a sample. The incidence of H. sacha¬
rovi in immature birds ranged from
41% in east central in Illinois
(Champaign County) to 78% in
west central Illinois (Hancock Coun¬
ty in 1954) ; in the adults it ranged
from 34% in northeast Illinois
(Cook County) to 56% in west cen¬
tral Illinois (Hancock County in
1952-53 ; no adults were studied from
this county in 1954). The incidence
of H. columbae in immature birds
ranged from 6% in west central Illi¬
nois (Hancock County in 1954) to
42%, also in west central Illinois
(Hancock County in 1952-53) ; in
the adults it ranged from 28% in
northeast Illinois (Cook County) to
78% in west central Illinois (Han¬
cock County in 1952-53).

There is no consistent pattern here
in the relation of incidence either to
year or to location within the state.
One can conclude, however, that the
results of any survey made at any
particular time and place do not
necessarily hold true for the same
place in a different year or even for
a different time in the same year,
nor do they necessarily hold true for
a different place not too far away
during the same time of the same
year.

The life cycle of Ilaemoproteus

columbae has been studied by Ara-
gao (1908), Adie (1915, 1924) and
Huff (1942) among others. The only
proven vector is the hippoboscid
fly, Pseudolynchia canariensis (syns.,
Lynchia maura, L. liviclicolor, L.
capensis) . In addition, Aragao
(1916) stated that Microlynchia
pusilla is a vector in South America,
but gave no experimental evidence.
Baker (1957) found that Id. colum¬
bae from the English wood pigeon
( Columba palumbus ) would undergo
sporogony in the hippoboscid, Orni-
thomyia avicularia, but 6 attempts
to infect domestic pigeons by bite or
injection of infected louse-flies
failed.

Huff (1932) found that Pseudo¬
lynchia canariensis was a vector of
H. sacharovi and used it to transmit
this parasite from the mourning
dove to the pigeon.

It is highly unlikely, however,
that hippoboscids are the only vec¬
tors of either H. columbae or H.
sacharovi. As Hanson et al. (1957)
pointed out, hippoboscids are ex¬
tremely rare on mourning doves,
especially in the northern states,
yet both species of Ilaemoproteus
are common in them. The discovery
by Fallis and Wood (1957) that bit¬
ing midges ( Culicoides ) are vectors
of H. nettionis of ducks suggests that
they may also transmit H. columbae
and H. sacharovi.

Altho the natural vectors of H.
columbae and H . sacharovi in Illi¬
nois are unknown, the findings of
Hanson et at. (1957) of a much high¬
er incidence of II. sacharovi in con¬
siderably younger mourning doves
than H. columbae permits one to
conclude either that the vectors of
the two species are different or that

Blood Parasites of Columborid Birds

105

Tabi.e 5. — Known Geographic Distribution and Prevalence of Toxoplasma gondii

in Columborid Birds.

Prevalence (%)

Surveys in

Present but

which more

which 100

no reliable

than 10

or more

figures on

birds were

Country

Bird Species

prevalence

examined

NORTH AMERICA

Dist. Columb .

Columba livia .

Ohio .

Columba livia .

New York .

Columba livia .

Tennessee .

Columba livia .

(Identifications Confirmed by Mouse Inoculation, Dye Test or Both)

Table 6. — Known Geographic Distribution and Prevalence of Toxoplasma.
Lankesterella or Similar Protozoa in Columborid Birds*.

Country

Bird Species

Prevalence (%)

Present but
no reliable
figures on
prevalence

Surveys in
which more
than 10
birds were
examined

Surveys in
which 100
or more
birds were
examined

ASIA

Portuguese India. .

Colu mba livia .

AFRICA

Belgian Congo. . . .

Columba livia .

SOUTH AND CEN

Brazil .

Panama .

Brazil .

TRAL AMERICA

Columba livia .

Columba rufina .

Columbigallina talpacoti..

* (Identifications by Microscopic Examination Only.)

II. sacharovi lias a shorter prepatent
period than H. columbae. However,
the marked difference in relative in¬
cidence in immature doves in the
same locality (Hancock County) in
different years (60% for H. sacha¬

rovi and 42% for H. columbae in
1952-53 as compared with 78% for
H. sacharovi and 6% for H. colum¬
bae in 1954) makes it possible to
speculate that the vectors may be
different.

106

Transactions Illinois Academy of Science

Haemoproteus columbae is only
slightly pathogenic. Infected birds
usually show no signs of disease.
In relatively heavy infections, the
birds may appear restless and go off
feed, and anemia may result from
destruction of erythrocytes, but this
is unusual. The schizonts occur in
the endothelial cells of the blood ves¬
sels of the lungs, liver and spleen.
The liver and spleen of affected birds
may be enlarged and dark with
pigment.

H. sacharovi appears to be only
slightly if at all pathogenic in the
mourning dove. Becker, Hollander
and Pattillo (1956) considered that
it caused the enlarged, purplish livers
which they encountered in dressing
domestic pigeon squabs from an in¬
fected flock ; there was apparently
no other evidence of disease.

Leucocytozoon. A single valid spe¬
cies of Leucocytozoon, L. marchouxi
Mathis and Leger, 1910 has been de¬
scribed from columborid birds (Le¬
vine, 1954). This species has round¬
ed gametocytes. In addition, Mincliin
(1910) described but did not name a
form with elongate gametocytes from
a collar-dove, Streptopelia semitor-
quata, in Uganda ; it has not been
encountered since.

Leucocytozoon marchouxi was
found in 30% of 128 C. palumbus
in Germany by Boing (1925) ; in
5% of 111 S. orient alis in Japan by
Ogawa (1912) ; and in 1.2% of 392
immature and 6.5% of 72 adult
Zenaidura macroura in Illinois by
Hanson et al. (1957).

The known geographic distribu¬
tion and prevalence of Leucocyto¬
zoon in columborid birds are shown
in Table 4. The results of surveys
in which 100 or more birds were ex¬

amined have been given above. There
have been only 3 such surveys — of
C. palumbus in Germany, of S. ori-
entalis in Japan, and of Z. macroura
in Illinois. There has been a total
of only 10 surveys in which 10 or
more birds were examined. They
involved 7 species of columborid
birds in 6 countries — Columba fasci-
ata in the United States, C. livia in
South Africa, C. palumbus in French
Morocco and Germany, Streptopelia
chinensis in the United States, S. ori-
entalis in Japan, S. turtur in French
Morocco and Spain, and Zenaidura
macroura in Illinois, Arizona and
California.

In addition to these surveys, Leu -
cocytozoon has been found in Colum¬
ba fasciata in Colorado and Arizona
or California, in C. livia in Pretoria,
South Africa (the only record of
this genus from the domestic pi¬
geon), in C. palumbus in England
and French Morocco, in Geopelia
striata on Mauritius, in Oena ca-
pensis in South Africa, in Sphenurus
sphenurus in India, in Streptopelia
capicola in South Africa, in S. chi¬
nensis in California, in S. senegal-
ensis in Upper Senegal and Nigeria,
in S. tranquebarica in Vietnam, in S.
turtur on Corsica and in French
Morocco, Spain and Italy, in Zenai¬
dura macroura in Georgia, the Dis¬
trict of Columbia area, Arizona and
California, and in Columba argen-
tina, C. vitiensis and Megaloprepia
magnifica in zoos. To these surveys
should be added quite a few others
in which Leucocytozoon was not
found ; those can be identified by
comparison with the data on Haemo¬
proteus (Table 3).

Altho Hanson et al. (1957) found
L. marchouxi in a higher proportion

Blood Parasites of Columborid Birds

107

of adults than of immature mourn¬
ing doves, they pointed out that their
figures are misleading. Of the 10
infected birds in their survey, 5
were adults, 1 was a juvenile 3 to
4 months old, and 4 were nestlings.
Since the great majority of immature
doves in their survey were juveniles,
the prevalence of patent infections
with L. marchouxi is probably con¬
siderably less than 1% in juveniles,
while that in nestlings is probably
considerably more. The youngest-
positive dove was only 14 days old
(Levine, 1954).

The vectors of L. marchouxi are
unknown. They are presumably spe¬
cies of Simulium like the vectors of
other species of Leucocytozoon. How¬
ever, the absence of Leucocytozoon
in columborid birds in South and
Central America despite the rela¬
tively large number of surveys which
have been carried out there suggests
that suitable vectors may not exist
in this area.

Nothing is known about the patho¬
genicity of L. marchouxi. There
were no signs of illness in the in¬
fected mourning doves seen by Le¬
vine (1954) and Hanson et al.
(1957), even tho 4 of them were
nestlings and 1 was only 14 days old.

Toxoplasma, Lankesterella, and
Similar Protozoa. There have been
a number of reports of Toxoplasma,
Lankesterella or morphologically
similar protozoa in columborid birds
(Tables 5, 6). Most have been in
the domestic pigeon. In the great
majority of cases, these organisms
have been assigned to the genus
Toxoplasma, altlio de Mello (1915)
called one form, which he found in
a domestic pigeon in Portuguese
India, a hemogregarine and de Mello

et al. (1917) named another form
from the same host Leucocytogrega-
rina francae. (This generic name
is no longer accepted ; it is a syno¬
nym of Hepatozoon.) However, due
to the confusion which is only now
being resolved regarding the identity
of these parasites, one cannot accept
any identification of Toxoplasma in
birds unless it has been confirmed
by animal inoculation or by serologic
means.

There have been four reports of
Toxoplasma gondii in domestic pi¬
geons which fulfill this requirement.
Feldman and Sabin (1949) found
T. gondii in 5% of 20 pigeons in
Cincinnati, Ohio, confirming their
identification by mouse inoculation.
Manwell and Drobeck (1951) found
T. gondii in 1% of 60 pigeons in
Syracuse, New York, confirming
their identification by use of the dye
test. Jacobs, Melton and Jones
(1952) found T. gondii in 12% of
80 pigeons in Washington, D. C.,
confirming their identification by the
dye test and mouse inoculation. Gib¬
son and Eyles (1957) found T.
gondii in 6% of 16 pigeons in Mem¬
phis, Tennessee, confirming their
identification by mouse inoculation.

In other reports, what may have
been either Toxoplasma or Lanke¬
sterella or possibty some other genus
have been found in the domestic pi¬
geon in Portuguese India, the Bel¬
gian Congo, Brazil and Panama, in
Columha rufina and Columhigallina
talpacoti in Brazil, and in Ducula
concinna in a zoo.

Discussion

A total of 174 papers is included
in the present analysis. Of these, 22
are from Europe, 22 from Asia, 32

108

Transactions Illinois Academy of Science

from Africa, 64 from North America,
32 from South and Central America,
and 2 from Australia. This number
of papers might lead one to believe
that the blood parasites of colum-
borid birds are rather well known.
This is far from the case. In his
Check-list of biids of the world, ,
Peters (1937) listed 61 genera and
320 species of birds in the order.
The most common parasite genus in
these birds is Haemoproteus. Levine
and Kantor (1959) pointed out on
the basis of their compilation that
Haemoproteus had been reported
from 19 genera and 45 species of
the order Columborida, but that
these comprise only 31% of the
known host genera and 14% of the
known host species. Plasmodium had
been reported from 20% of the
genera and 7% of the species, Leu-
cocytozoon from 11% of the genera
and 5% of the species, Trypanosoma
from 8% of the genera and 2% of
the species, and Toxoplasma or some¬
thing similar from 5% of the genera
and 1% of the species. No parasites
at all have been reported from two
genera, Oreopelia and Gallicolumba,
which contain 15 and 18 species,
respectively, and only two cases have
been reported from the genus Ducu-
la, which contains 37 species.

Levine and Kantor ’s compilation
included birds in zoos. If these are
omitted, then Haemoproteus has
been found in only 20% of the known
host genera and 8% of the known
host species, Plasmodium in 11% of
the genera and 3% of the species,
Leucocytozoon in 10% of the genera
and 4% of the species, Trypanosoma
in 8% of the genera and 2% of the
species, and T oxoplasma or some¬
thing similar in 3% of the genera

and 1% of the species.

But this is not all. A great many
of these records were more or less
casual. The authors examined a
series of blood smears from a mis¬
cellany of birds and made no at¬
tempt to identify the parasites be¬
yond genus. Levine and Kantor
(1959) found on analysis of their
compilation that the parasite species
had been named in 51% of 180 re¬
ports of Haemoproteus, 64% of 47
reports of Plasmodium, 32% of 31
reports of Leucocytozoon, 60% of 10
reports of Trypanosoma and 75% of
16 reports of T oxoplasma and simi¬
lar forms. If reports on birds in
zoos are omitted, these figures become
58% of 138 reports of Haemopro¬
teus, 62% of 26 reports of Plas¬
modium, 35% of 26 reports of
Leucocytozoon, 60% of 10 reports of
Trypanosoma, and 77% of 13 reports
of Toxoplasma and similar forms. In
43% of the total of 213 records, the
species name of the parasite was not
given. Furthermore, in only 41
(24%) of the 174 papers in the pres¬
ent analysis were 10 or more birds
of a single species examined and the
prevalence of infection given, and
in only 12 (7%) of them were 100
or more birds examined and the
prevalence of infection given.

Further light can be thrown on
the reliability of our present infor¬
mation on geographic distribution
and prevalence of these protozoa by
considering the number of examina¬
tions on which it is based. I have
done this for Haemoproteus, the
genus on which we have most infor¬
mation. In quite a few reports, the
number of birds examined was not
stated and may have been relatively
small. The data assembled for the

Blood Parasites of Columborid Birds

109

remainder are based on the examina¬
tion of blood smears from 2515 birds
—296 from Europe, 356 from Asia,
53 from Africa, 1391 from North
America, and 419 from South and
Central America. These examina¬
tions. may I remind you, were made
between 1891 and 1957. A geogra-
ph ic* distribution map might appear
to be fairly well filled in, but it
would be based on a pitifully small
population sample.

A great deal thus remains to be
done before we can claim to have
really good information on the blood
parasite situation in the great ma¬
jority of columborid birds in most
parts of the world. Casual observa¬
tions are all very well, but extensive,
careful, thoro surveys would be much
more valuable. Furthermore, sur¬
veys made at one time of year or on
one age group of host may not repre¬
sent the situation at another time of
year or on another age group of the
same host. We lack information on
all this.

Not only is our information on the
geographic distribution and inci¬
dence of blood parasites of colum¬
borid birds scattered and superficial,
but our information on their life
cycles, vectors and pathogenesis is
also poor. AVe do not know the role
of these parasites in the interplay
of favorable and unfavorable factors
on which their hosts’ survival in
nature depends. AVe suspect that
Trypanosoma and Haemoproteus
may be relatively non-pathogenic ;
we think that Plasmodium and Toxo¬
plasma may be more or less patho¬
genic — we know that they can be in
the laboratory, at least ; but we do
know what to say about Leucocyto-
zoon and Lankesterella. Here, too,

more information is needed.

Another problem which needs fur¬
ther study is the role of the domestic
pigeon as a possible reservoir of
Toxoplasma. The surprisingly high
mean incidence of 8% in a total of
176 birds examined by Feldman and
Sabin (1949), Manwell and Drobeck
(1951), Jacobs, Melton and Jones
(1952) and Gibson and Eyles (1957)
in different surveys suggest that this
bird may well be an important reser¬
voir.

Literature Cited

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proteus columbae. Inch J. Med. Res..
2:671-680.

Adie, H. 1924. The sporogony of Haemo¬
proteus columbae. Bull. Soc. Pathol.
Exot., 17:605-613.

Aragao, H. de B. 1908. uber den Ent-
wicklungsgang und die ubertragung
von Haemoproteus columbae. Arch.
Protist., 12:154-167.

Aragao, H. de B. 1916. Pesquisas sobre
o “Haemoproteus columbae.” Brazil
Med., 30:361-362.

Baker, J. R. 1956a. Studies on Trypano¬
soma avium Danilewsky 1885. II.
Transmission by Ornithomyia avieu-
laria L. Parasitology, 46:321-334.
Baker, J. R. 1956b. Studies on Trypano¬
soma avium Danilewsky 1885. III.
Life cycle in vertebrate and inverte¬
brate hosts. Parasitology, 46:335-352.
Baker, J. R. 1957. A new vector of
Haemoproteus columbae in England.
J. Protozoo!., 4:204-208.

Becker, E. R., W. F. Hollander and
W. H. Pattlllo. 1956. Naturally oc¬
curring Plasmodium and Ilaemojwo-
teus infection in the common pigeon.
J. Parasitol., 42:474-478.

Bennett, G. F. 1961. On the specificity
and transmission of some avian trypa¬
nosomes. Can. J. Zool., 39:17-33.
Bennett, G. F. and A. M. Fallis. 1960.
Blood parasites of birds in Algonquin
Park, Canada and a discussion of their
transmission. Can. J. Zool., 38:261-273.
Boing, W. 1925. Untersuchungen fiber
Blutschmarotzer bei einlieimischen
Vogelwild. Zentr. Bakteriol. I. Orig.,
95:312-327.

Cartni. A. 1912. Sur un nouvel hemato-
zoaire du pigeon. Compt. Rend. Soc.
Biol., 73:396-398.

110

Transactions Illinois Academy of Science

Celli, A. and F. Sanfelice. 1891a. Sui
parassiti del globulo rosso nell’ uomo
e negli animali. Ann. 1st. Sper., Univ.
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Celli. A. and F. Sanfelice. 1891b. Ueber
die Parasiten des rothen Blutkorper-
chens im Menschen und in Thieren.
Fortschr. Med., 9:499-511, 541-552, 581-
586.

Coatney. G. R. 1933. Relapse and as¬
sociated phenomena in the Haemo-
proteus infection of the pigeon. Am.
J. Hyg., 18:133-160.

Coatney, G. R. 1938. A strain of Plas¬
modium relictum from doves and
pigeons infective to canaries and the
common fowl. Am. J. Hyg., 27:380-389.

Coatney, G. R. and E. West. 1938. Some
blood parasites from Nebraska birds.
II. Am. Midland Naturalist, 19:601-
612.

Couch, A. B., Jr. 1952. Blood parasites
of some common Texas birds. Field
and Lab., 20:146-154.

Covaleda Ortega, J. and J. Gallego
Berenguer. 1950. Hemoproteus aviares.
Rev. Iber. Parasitol., 10:141-185.

Fallis, A. M. and D. M. Wood. 1957.
Biting midges (Diptera: Ceratopogoni-
dae) as intermediate hosts for Haemo-
proteus of ducks. Can. J. Zool., 34:425-
435.

Feldman, H. A. and A. B. Sabin. 1949.
Skin reactions to toxoplasmic antigen
in people of different ages without
known history of infection. Pediat¬
rics, 4:798-804.

Francfiini, G. 1924. Observations sur
les hematozoaires des oiseaux d’ltalie.
(2e note). Ann. Inst. Pasteur, 38:470-
515.

Gibson, C. L. and D. E. Eyles. 1957.
Toxoplasma infections in animals as¬
sociated with a case of human con¬
genital toxoplasmosis. Am. J. Trop.
Med. Hyg., 6:990-1000.

Giovannoni, M. 1946. Fauna parasi-
tologica paranaense. I. Haemoproteus
columbae Celli e Sanfelice. 1891 em
Colurntta livia domestica nos pombais
de Curitiba. Arq. Biol. Tecnol., 1:19-24.

Hanson, H. C., N. D. Levine, C. W.
Kossack, S. Kantor and L. J. Stan-
nard. 1957. Parasites of the mourn¬
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43:186-193.

Herman, C. M., W. C. Reeves, H. E.
McClure, E. M. French and W. McD.
Hammon. 1954. Studies on avian ma¬
laria in vectors and hosts of encepha¬
litis in Kern County, California. I.
Infections in avian hosts. Am. J. Trop.
Med. Hyg., 3:676-695.

Huff, C. G. 1932. Studies on Haemopro¬
teus of mourning doves. Am. J. Hyg.,
16:618-623.

Huff, C. G. 1935. Plasmodium hexa-
vierium n. sp. from the bluebird, inoc-
ulable to canaries. Am. J. Hyg.,
21:274-277.

Huff, C. G. 1939. A survey of the blood
parasites of birds caught for banding
purposes. J. Am. Vet. Med. Assoc.,
94:615-620.

Huff, C. G. 1942. Schizogony and garne-
tocyte development in Leueocytozoon
simondi, and comparisons with Plas-
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Dis., 71:18-32.

Huff, C. G. and D. F. Marchbank. 1955.
Changes in infectiousness of malarial
gametocytes. I. Patterns of oocyst
production in seven liost-parasite com¬
binations. Exp. Parasitol., 4:256-270.

Huff, C. G., D. F. Marchbank, A. H.
Saroff, P. W. Schrimsfiaw and
T. Shiroishi. 1950. Experimental in¬
fections with Plasmodium fallax
Schwetz isolated from the Uganda
tufted guinea fowl Numida meleagris
major Hartlaub. J. Nat. Malaria Soc.,
9:307-319.

Jacobs, L., M. L. Melton and F. E.
Jones. 1952. The prevalence of toxo¬
plasmosis in wild pigeons. J. Para¬
sitol., 38:457-461.

Kraneveld, F. C. and M. Mansjoer.
1954. Onderzoekingen over bloedpara-
sitien. VIII. Over een Trypanosoma
van de tortelduif ( Streptopelia chinen-
sis tigrina) . Hemera Zoa, 61:9-20.

Kruse, W. 1890. Ueber Blutparasiten.
Virchow’s Arch., 121:359.

Laveran, M. A. and A. Petit. 1909. Sur
une hemamibe du Melopelia leucoptera
L. Compt. Rend. Soc. Biol., 66:952-954.

Levine, N. D. 1954. Leueocytozoon in
the avian order Columbiformes, with
a description of L. marchouxi Mathis
and Leger 1910 from the mourning
dove. J. Protozool., 1:140-143.

Levine, N. D. and S. Kantor. 1959.
Check-list of blood parasites of birds
of the order Columbiformes. Wildl.
Dis., 1(1) : 38 pp.

Manwell, R. D. and H. P. Drobeck.
1951. Mammalian toxoplasmosis in
birds. Exp. Parasitol., 2:221-235.

Mathey, W. J., Jr. 1955. Two cases of
Plasmodium relictum infection in do¬
mestic pigeons in the Sacramento
area. Vet. Med., 50:318.

Mello, I. F. de. 1915. Preliminary note
on a new haemogregarine found in the
pigeon’s blood. Ind. J. Med. Res.,
3:93-94.

Blood Parasites of Columborid Birds

111

Mello. I. F. de and L. J. Bras de Sa.
1916. A contribution to the study of
haemoprotozoa in Portuguese India.
Ind. J. Med. Res., 3:731-737.

Mello, I. F. de, L. J. Bras de Sa, L. de
Sousa. A. Dias and R. Noronha. 1917.
Hematozoaires et pseudo-hematozo-
aires de l’lnde Portugaise. Ann. Sci.
(Fac. Med.) Porto, 3:181-200.
Minchin, E. A. 1910. Report on a col¬
lection of blood-parasites made by the
Sleeping Sickness Commission, 1908-
1909, in Uganda. Rep. Sleep. Sick.
Comm. Roy. Soc., 10:73-86.

Novy. F. G. and W. J. MacNeal. 1904a.
Trypanosomes and bird malaria. Am.
Mprl 8- 939-934

Novy, F. G. and W. J. MacNeal. 1904b.
Trypanosomes and bird malaria. Med.
News. N. Y., 85:1144-1146.

Novy, F. G. and W. J. MacNeal. 1905a.
Trypanosomes and bird malaria. Proc.
Soc. Exp. Biol. Med., 2:23-28.

Novy. F. G. and W. J. MacNeal. 1905b.
On the trypanosomes of birds. J. Inf.
Dis., 2:256-308.

Ogawa, M. 1912. Notizen tiber die blut-
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126.

Peters. J. L. 1937. Check-list of birds
of the world. Vol. 3. Harvard Univ.
Press.

Renjifo-Salcedo, S., C. Sanmartin and
J. de Zulueta. 1952. A survey of the
blood parasites of vertebrates in east¬
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Santos Dias, J. A. T. 1953. Resultados
de um reconhecimento zoologico no
Alto Limpopo efectuado pelos Drs.
F. Zumpt e J. A. T. Santos Dias.
V. Hematozoarios das aves: generos
Haemoproteus Kruse e Carpanoplasma
n. gen. Mocambique, 73:61-99.

Sergent, Ed. 1941a. Trypanosome de la
tourterelle “a longue queue” Strepto-
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Arch. Inst. Pasteur Algerie, 19:29.

Sergent, Ed. 1941b. A propos du trypa¬
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Inst. Pasteur Algerie, 19:431.

Singh, J., C. P. Nair and A. David.
1951. Five years’ observation on the
incidence of blood protozoa in house
sparrows (Passer domesticus Lin¬
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5:229-233.

Wolfson, F. 1937. Experimental infec¬
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Hyg., 26:53-59.

Wolfson, F. 1940. Exo-erythrocytic
schizogony associated with the wood-
thrush strain of Plasmodium cathe-
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the host. Am. J. Hyg., 31(0:26-35.

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J. Parasitol., 29:187-196.

Manuscript received March 1, 1962.

DENTAL ANOMALIES OF THE RACCOON

ROBERT L. MARTIN*

University of Illinois, Urbana

Anomalies in raccoon dentition are
not commonly recorded in the litera-
tnre. To obviate the conclusion that
such anomalies are rare, the follow¬
ing cases are reported. Normal den¬
tition for Procyon lotor, the raccoon,
as given by Goldman (1950) and
by Hall and Kelson (1959), is

* Present address : Department of Sci¬
ence, State University College, Platts¬
burgh, New York.

3 14 2

i — , c — , p - — ■ , m — . No mention
3 1 4 2

is made in either of these standard
works of variations from this pat¬
tern.

The two skulls illustrated in Plate
I, Figures 1-3, present an upper den¬
tal formula of 3-1 -3-2, reflecting the
absence of the first premolar on each

Plate 1. — Dental anomalies in the raccoon. See text for explanation of figures.

[ 112 ]

Denial Anomalies

113

side, though the lower jaw denti¬
tions of these were normal. The
skull shown in Figure 1, RLM No.
17, was found 5% feet down in the
gray clay of a stream bed “several
miles west” of Abilene, Dickinson
County, Kansas. The skull shown
in Figures 2 and 3, CCS No. 2663,
is from a male specimen taken in
Allerton Park, near Monticello, Piatt
County, Illinois.

The skull in Figure 4, GCS No.
1773, has an unusual diastema be¬
tween the first and second upper
premolars, the spacing causing un¬
usually severe wear on the second
upper right premolar. This was a
female specimen collected several
miles east of Lodge, Piatt County,
Illinois.

In Figure 5, a large opening in the
right maxilla exposing the base of
the right canine tooth is shown. This
skull, GCS No. 1788, is from a fe¬
male specimen taken in Chautauqua
Wildlife Refuge, Mason County,
Illinois. On the left side of this
skull, though not illustrated here,
the third premolar is absent with a
small rounded piece of tooth pro¬
truding from a tiny socket in the
anterior portion of the space which
would normally be filled by the third
premolar, no other remnants of an
alveolus being visible. This condi¬
tion undoubtedly resulted from in-

«/

jury, an explanation not applicable
for the former cases of missing teeth
and alveoli.

The skull illustrated in Figure 6,
RLM No. 51, was found with other
skeletal remains about 40 feet inside
the entrance of one of the mines in
the Blackball Mine system, 1 % miles
west of Utica, La Salle County, Illi¬
nois. The opening leading from the
alveolus of the left canine to the
external surface of the maxilla is
indicated by a horse hair. Although
the bone surrounding this opening
and that shown in Figure 5 does not
have the spongy appearance char¬
acteristic of diseased bone tissue, the
openings could have resulted from
alveolar abscesses when the animals
were younger, as the edges of the
openings are rounded inwardly by
extension of the compact bone into
the openings.

The GCS skulls are from the col¬
lection of Glen C. Sanderson of the
Illinois Natural History Survey, to
whom the author is indebted for their
loan.

Literature Cited

Goldman, Edward A. 1950. Raccoons
of North and Midd’e America. North
American Fauna 60, 153 pp.

Hall, E. Raymond, and Keith R. Kel¬
son. 1959. The Mammals of North
America. Vol. II. The Ronald Press
Company, New York

Manuscript received April 20, 1002.

LEAF CHARACTERISTICS OF TWO HYBRID JUNIPERS

MARGARET KAEISER
Southern Illinois University, Carbondale

The purpose of this study has been
to compare and contrast the whip
and scale leaves of two junipers in
a natural population located ap¬
proximately seven and one-half miles
south of Carbondale, Illinois on
limestone outcroppings on a hill near
U. S. Highway 51. The stand of
trees from which the two specimens
were selected is a genetically mixed
“swarm hybrid population” (Hall,
1952) of Juniperus virginiana L.
(eastern red cedar) J. Ashei Buchh.
(Ozark white cedar). The tree in¬
dicated as “ virginiana ” was chosen
to represent the maximum combina¬
tions of the more typical eastern
red cedar growth habit ; the other
tree, indicated as “ hybrid ”, ex¬
hibits more apparent Ozark white
cedar characteristics. Macroscopic as
well as microscopic differences in
leaf structure were studied.

According to Florin (1931, 1951),
species of Juniperus exhibit four
kinds of leaves : cotyledons, juvenile,
transitional and mature. The transi¬
tional and mature types are fre¬
quently referred to as the whip and
scale leaves respectively. Combina¬
tions of certain leaf structures of
juvenile, whip and scale leaves for
J. virginiana, and other combina¬
tions for J. Ashei have been assem¬
bled by Hall (1952). Structural
variations of leaves from trees in
natural stands deviating from these
two sets of combinations are inter¬
preted as indicators of hybridity.
The combined degree of deviations

is expressed as a greater or lesser
tendency to resemble one or the other
original parent in the presumed
cross (Anderson, 1949). The selec¬
tion of the two segregates used in
this study rests on the assumption
that it is advantageous to have plants
from a similar genetic background.
The two trees are the same ones
sampled to point out the differences
in types of shoot apices (Kaeiser,
1960). Specimens from collections
are on deposit in the Herbarium of
Southern Illinois University.

Acknowledgments

The writer wishes to thank Dr.
Edgar Anderson and Dr. M. T. Hall
for their interest in the study. As¬
sistance from a research grant of
the Illinois State Academy of Sci¬
ence is gratefully acknowledged.

Materials and Methods

Fresh as well as herbarium col¬
lections were used for macroscopic
studies and for those under low mag¬
nification (X10-X50). For micro¬
scopic study leaves and branches
were killed and fixed in F.A.A., as¬
pirated, dehydrated in the tertiary
butyl alcohol series and embedded
in paraffin. Sections were cut ap¬
proximately 10 microns in thickness
and were stained in the usual man¬
ner with Safranin O and Fast Green
FCF.

Figure 1 shows the general mode
of branching pattern of J. Ashei

Hybrid Junipers

115

^ \iZi. J**4$*~ Vrfj j

U r*n>

«*■ $<*«<*

Fig. 1. — Left, J. Ashei; and right, J. virginiana.

and J . virginiana growing at the Kas-
kaskia Experimental Forest (Shaw¬
nee National Forest) in Hardin
County, Illinois. Known seed source
was from near Lebanon, Tennessee.
Other materials of J. Ashei, as in¬
dicated in Table 1, were provided by
G. J. Goodman from the Arbuckle
Mountains of Oklahoma. Herbarium
materials collected by W. W. Ashe
in Arkansas and verified by Bucli-
liolz (1930) in his proposed naming
of the species have been studied.
Living specimens growing in Okla¬
homa, Arkansas and Missouri have
also been observed. “Near Ashei ”
material from McYey Knob, Ozark
County, Missouri was provided by
Hall. The two young trees listed in

Table 1, together with the mature
trees indicated as “ virginiana ” and
“ hybrid”, were all growing in the
same stand. The older trees were
approximately thirty feet in height.

Ranges in lengths of sheaths and
blades of leaves found in specimens
of unmixed populations of J . Ashei
and J. virginiana are given by Hall.
The measurements were used for
comparison in the present analysis.

Figure 2 represents surface, medi¬
an longitudinal and median trans¬
verse aspects of the mature stomatal
apparatus on the abaxial surfaces
of spur leaves of the “ hybrid ” and
‘ ‘ virginiana ’ ’ trees used in the study.
Terminology follows that of Florin.

116

Transactions Illinois Academy of Science

Fig. 2. — Diagram of surface view of
matu e stomatal apparatus of “virgini-
ana.” 1. From the dorsal surface of
spur leaf: a, auxiliary cell lateral to
guard cell; e, encircling cell; p, polar
cell. X292. 2. “Hybrid.” 3. Longitudinal
median section through mature stomatal
apparatus of “virginianci.” Guard cell,
g. X292. 4. “Hybrid.” 5. Transverse

section of mature stomatal apparatus of
“virginiana.” From the dorsal surface
of spur leaf: a, auxiliary cell; e, en¬
circling cell; g, guard cell. X292.
6. “Hybrid.”

Observations and Discussion

Among the most easily observable
field characteristics in a mixed popu¬
lation of eastern red and Ozark
white cedar are : 1 ) the color of

foliage, 2) the proportionate lengths
of the whips or terminal branches
and 3) the relative amount of crowd¬
ing of the lateral branches (Fig. 1;
Table 1). The yellow-green rather
than blue-green color, the much
shorter whip branches and the very
crowded appearance of lateral
branches are all notable features of
Asliei influence, and are features in
the “ hybrid ” specimen chosen for
detailed study. Consistent with these

characteristics there are also other
structural features of the whip and
spur leaf types, as summarized in
Table 1.

Color of foliage alone is often de¬
ceptive, and in young trees with
only juvenile foliage it is not a re¬
liable character. The relatively
crowded appearance of lateral
branches, as pointed out in an earlier
study, is correlated with relative size
and rate of growth of the shoot
apices. In whip branches the shoot
apex in this “ hybrid ” in vigorously
growing shoots, has a wider diameter,
the pith tissue is closer to the apex,
and the flanking tissue (derivatives
of which contribute to leaf primor-
dia) is closer to the apex than in the
“ virginiana ” specimen (Ivaeiser,
1960).

Hall (1952) has shown that typi¬
cal Ashei whip leaves have sheaths
4.0 mm and blades 3.0 mm in length,
in contrast to typical “virginiana”
whip leaves with sheaths 9.0 mm
and blades ranging from 4. 0-5.0 mm
in length. Spur leaves of Asliei
show 1.5 mm for sheath and 1.5
mm for blade lengths, whereas typi¬
cal “virginiana” spur leaves have
sheaths 1.0 mm and blades 3.0 mm
in length. These are averages
occurring in unmixed populations.
When the proportionate ratios of
sheath to blade lengths of whip
leaves of the two trees are compared
(Table 1), there is no significantly
close correlation. However, there is
a decided tendencv exhibited in the
“virginiana” specimen to have con¬
sistently longer whip leaves, so that
total leng’th is more closelv related
to typical eastern red cedar. Fur¬
thermore, there is a lack of any ser-

Table 1. — External Features of Leaves of Selected Specimens of Juniperus.

Hybrid Junipers

117

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Transactions Illinois Academy of Science

ration on this kind of leaf, and an
absence of keels on the sheaths. The
presence of the latter two features
are Ozark white cedar characteris¬
tics.

When the proportionate ratios of
sheath to blade lengths of the spur
leaves are compared, this type of
leaf is found to be more like Ashei
in both specimens. Other structures
when compared indicated that the
“hybrid77 spur leaves occasionally
showed both round and raised glands
and had blades slightly but detec-
tably humped. All of these charac¬
teristics are indicative of Ashei and
all are absent in the “ virginiana77
specimen.

Both of the young trees tended
to show more eastern red cedar char¬
acteristics in: 1) greater total
lengths of juvenile leaves; 2) pro¬
portionate lengths of sheath to
blade; 3) lack of serration; 4) lack
of keels; 5) lack of round glands;
and 6) glands not raised.

Aside from the differences in
shoot apices referred to above, there
are also consistent differences in the
mature stomatal apparatus on the
dorsal surfaces of the spur leaves.
In following Florin’s terminology
the apparatus is of the haplocheilic
type, and within this category is
classified as amphicyclic. Surface
views always disclose two polar cells,
and generally two lateral auxiliary
cells on either side of each pair of
guard cells (Figure 2). Surround¬
ing all of these are the encircling
cells. All cells mentioned belong to
the apparatus. The tendency for
greater over-arching of both polar
and lateral auxiliary cells in “vir¬
giniana77 can be seen readily from
Figures 3-6. There is also a ten¬

dency for greater size of this appara¬
tus in the “Ashei77 specimen. This
is consistent with other measure¬
ments taken of ordinary epidermal
cells of the dorsal surfaces of spur
leaves of known J. Ashei.

Summary

1. In the two mature trees selected,
i.e. “ virginiana 77 and “hybrid77,
the former showed the tendency
for whip leaves to resemble those
of the genetically unmixed east¬
ern red cedar in being of greater
total length. The spur leaves,
although with proportionately
shorter blades than eastern red
cedar, were unhumped and pos¬
sessed oblong, unraised glands,
all “ virginiana 77 features. The
“hybrid” whip leaves were
shorter in length, some exhibit¬
ing serration and keels on the
sheaths, all Ozark white cedar
features. The spur leaves ex¬
hibited occasionally round and
raised glands, and blades at least
slightly humped, all Ashei fea¬
tures.

2. Both young trees selected tended
to show more “virginiana” char¬
acteristics, the older one especi¬
ally, as evidenced by the assem¬
blage of six detectable morpho¬
logical similarities.

3. The mature stomatal apparatus
from the dorsal surfaces of spur
leaves of the “virginiana7 7 speci¬
men were smaller in size than in
the “hybrid77 ; both the polar and
lateral auxiliary cells overarched
the guard cells more in the for¬
mer than in the latter. The stoma¬
tal apparatuses of “virginiana77
are more similar to those of east-

Hy b rid Jun ipers

119

ern red cedar, while those of the
‘ ‘ h ybrid ’ ’ specimen correspond
closely in both size and shape to
those of Ozark white cedar.

Literature Cited

Anderson, E. 1949. Introgressive hybri¬
dization. John Wiley and Sons, Inc.
New York, 109 pp.

Buciiholz, John T. 1930. The Ozark
white cedar. Bot. Gaz., 90: 326-332.

Florin, R. 1931. Untersuchungen zur
Stammesgeschichete der Coniferales
und Cordaitales. Svenska Vetensk.
Akad. Handl. Ser. 5, 10: 1-588.

Florin, R. 1951. Evolution in Cordaites
and Conifers. Acta Horti Bergiana,
15: 285-388.

Hall, M. T. 1952. Variation and hybri¬
dization in Juniperus. Ann. Mo. Bot.
Gard., 39: 1-64.

Kaeiser, M. 1960. Shoot apices in two
hybrid junipers. Trans. Ill. St. Acad.
Sci., 53: 132-140.

GERMINATION CAPACITY IN AMERICAN BASSWOOD

WILLIAM C. ASHBY

Southern Illinois University

In the course of several years work
on basswood ( Tilia americana L.)
a number of fruit collections were
obtained, from many of which seed
was extracted and germinated. The
yield of sound seed and germination
characteristics of each fruit lot were
assessed as germination capacity.
This capacity was related to year of
collection, insect damage, geo¬
graphic origin, and other factors.

Acknowledgments

This investigation was aided by
grants from the National Science
Foundation and from the Dr. Wal¬
lace C. and Clara A. Abbott Me¬
morial Fund of the University of
Chicago. I wish to thank the follow¬
ing for furnishing basswood fruits
or seeds : W. L. Ashby, W. D. Bell,
H. J. F. Gall, 0. Vaartaja, P. D.
Voth, J. C. Warden and the Central
States, Northeastern and South¬
eastern Forest Experiment Stations
of the U.S. Forest Service. Seed
was purchased from the F. W. Schu¬
macher and Herbst seed companies.

Materials and Methods

One approach was to plant un¬
treated fruits in the garden with
and without straw mulch or in flats
with potting soil. The flats were
retained in the greenhouse or, more
usually, placed out in a coldframe.
A few embryos were also dissected
from seed and grown in the labora¬
tory. The great majority of fruits

was treated with concentrated nitric
acid for approximately three hours,
the seed shelled out, dried, treated
with concentrated sulfuric acid for
15 minutes, stratified in moist
vermiculite or sphagnum for three
or four months at 36° F, and sown
in the greenhouse or garden under
presumably favorable germination
conditions (Spaeth, 1934; U. S. De¬
partment of Agriculture, 1948). The
germination figures reported are for
the appearance of the cotyledons
above ground. Sprouting of the
hypocotyl which took place during
stratification was generally cor¬
related with the subsequent emer¬
gence of the cotyledons and is not
reported. A trial was made of the
Johnson (1946) method in which
fruits were soaked in water for sev¬
eral days, treated with concentrated
sulfuric acid for 40 minutes, rinsed,
and planted.

Forty-five collections obtained
during the years 1955 through 1959
included fruits from various parts
of the natural distribution of Amer¬
ican basswood, North Carolina to
Manitoba and Maine to Minnesota.
The fruits varied in size, extent of
hairiness and persistence of the
style. Samples from the various
sources were retained as herbarium
material by the author. The num¬
bers of fruits within the several
collections varied greatly. This in
part reflected variations in fruit
production from tree to tree, year
to year and locality to localitv.

[ 1-0 ]

Amer ica n Bass wood

121

Results

Embryos removed from the seed
soon started growth on moist filter
paper. In contrast, no germination
was observed from untreated fruits
planted in the greenhouse for pe¬
riods up to 18 months. During this
period the fruit coats disintegrated,
but the seed remained hard. Un¬
treated fruits planted outdoors in
flats or in the garden in spring or
fall of one year germinated in late
April of the first, second or (in lesser
numbers) third spring after plant¬
ing. Germination in the first season
reached 15%, in agreement with
Bailey (1961).

Considerably higher germination
values were found the same spring
as planting for acid-treated and
stratified seeds (Spaeth, 1934;
Johnson 1946). In Table 1 the high¬
est value found is 78%. Appreciable
variation was noted from one seed
lot to another. For those studies
with acid-treated and stratified
seed continued for a second spring,
less than 5% additional germina¬
tion was recorded. The average first
season germination percentages
were 39% for 1822 seed in 1957 and
34% for 1695 seed in 1958. The
seedlings from various seed lots
were usually very similiar in ap¬
pearance. One accession of 177 seed
from southeastern Wisconsin had
over 10% tricot and one tetracot
seedling. No other unusual types of
seedlings were observed. Treat¬
ments given to stratified seed after
spring planting, such as mulching
with straw or shading (Ashby,
1961), resulted in less rapid emer¬
gence but approximately equal final
numbers of seedlings.

Additional factors were of im¬
portance in determining basswood’s
regenerative capacity (Table 1).
The several collections totalling sev¬
eral thousand fruits varied in yield
of sound seed from zero to nearly
100%. Filled fruits were usually
one - seeded. Collections yielding
high percentages of fruit with seed
often had several percent of fruit
with double or even triple seed.
Seed quality varied from nearly
100% sound to 100% which rotted
when stratified after acid treatment.
Some otherwise sound seed lots had
up to 30% insect infestation. An
apparent yearly correlation in Table
1 of high percentages of multiple-
seeded fruit and high values of
damaged or unfilled seed was not
true for individual seed lots. Ground
collections, while sometimes good,
tended to give poorer seed yields
than tree collections. Average seed
weights for 13 collections in 1958
varied from 12 to 38 mg with an
average of 31 mg.

Fruit production, which was
found on some trees in the Chicago
region each year of the study, oc¬
curred with greatest abundance in
1957. Collections made from a
single tree in southern Wisconsin
had many poorly-filled seed and no
multiple-seeded fruits in 1956, while
in 1957 fruits were greater in
amount with no poor seed and some
multiple-seeded fruits. On a trip
in October, 1958 to parts of Iowa,
Minnesota, and Wisconsin, I found
only one tree with fruit out of hun¬
dreds of trees examined.

Comparisons of fruit quality for
individual trees are illustrated by
data from four trees in a small val¬
ley in the St. Lawrence River sec-

Transactions Illinois Academy of Science

1 22

Table 1. — -Characteristics of Basswood Fruits and Seeds.

Year of Collection

Characteristic

1955

1956

1957

1958

Number of collections with

Seed yield 0-33% .

Seed yield 34-67% .

Seed yield 68-100% .

Percent of the above collections with

Multiple-seeded fruit .

33%

21%

11%

Seed poorly filled, insect damage, etc .

25%

83%

57%

44%

Percent germination of

Apparently sound acid-treated and stratified seed

0-70%

0-78%

0-52%

0-20%*

* Germination in garden rather than greenhouse.

tion of New York State. Seed yield
in 1958 ranged from 100 to 5% and
emergence in the garden for ap¬
parently sound stratified seed from
15 to 0%. The tree yielding 100%
seed per fruit had multiple-seeded
fruit and some poorly filled seed.
Most of the seed from the poorest
yielding tree were poorly filled.

Basswood germination was ob¬
served each spring in several field
areas of the Chicago region. The
numbers varied from year to year.
Only one instance of very abundant
basswood seed germination was
found. This occurred in a presum¬
ably protected area in southwestern
Michigan which had been invaded
by campers the previous autumn.
The nature of the altered seed bed
was not determined.

Discussion

Germination capacity in basswood
is related to the several factors
studied: fruit production, numbers
of seed per fruit, percent of sound

seed, and germination percentage
(Spaeth, 1934). The delayed germi¬
nation leads to annual appearance
of seedlings despite the yearly var¬
iations in fruit production and qual¬
ity. Other investigators (Den Uyl,
et al., 1938 ; Hart, 1958, 1959 ; Mc-
Conkey, 1960, 1961; Rudolph, 1950-
61 ; Spaeth, 1934) have reported
marked annual and geographic
variation in fruiting of forest tree
species, including basswood. Factors
outside the scope of the present in¬
vestigation which determine regen¬
eration capacity include animal use
of seed (which I have observed on
occasion to be extensive), distribu¬
tion of seed to favorable sites, and
suitable soil and climatic conditions.
My present concept is that in the
Chicago region the production and
germination capacity of American
basswood seed is rarely a substan-
tial limiting factor where mature
trees are found. 1 am not yet satis-
tied that such is the case in southern
Illinois.

America n Bass wood

A second type of regeneration
in basswood involves sprouting from
the base of the trunk. This serves
to maintain the species, once estab¬
lished. In only one instance was
such a sprout found as far as one
foot from a trunk. If true root
sprouts occur, they are rare in com¬
parison to the common trunk
sprouts.

Length of the stratification period
was not systematically studied for
its effect on germination. This prob¬
ably affected the absolute perform¬
ance of one seed lot versus another.
My criterion for removing the seed
from stratification conditions and
planting them was evidence of
growth by the radicle. As a rule
several seed lots were removed at,
one time. The percentages of seed
in which growth of the radicle was
evident might differ for the several
seed lots. Seed for comparative
tests were sorted for response to
stratification and the treatments
were given to representative group¬
ings. I did find that “hard” seed
may give very low emergence per¬
centages after a stratification period
which leads to good emergence by
seed on which sprouting is evident.
Thus intra-seed-lot differences need
to be considered in evaluating inter¬
seed-lot performance. A require¬
ment for prolonged stratification
(Spaeth, 1934) would influence the
germination capacity of individual
seed.

Summary

Basswood fruit collections varied
in seed yield from 100% to 0%.
Germination of acid-treated and
stratified selected seed ranged from

123

78% to 0%. Fruiting alone is not
a satisfactory measure of basswood
seed production and quality. Delay¬
ed germination can lead to annual
appearance of seedlings despite the
yearly variations in seed production
and quality.

Literature Cited

Ashby, W. C. 1961. Responses of Ameri¬
can basswood seedlings to several
light intensities. Forest Science, 7 :
273-281.

Bailey, C. V. 1961. Early collection and
immediate sowing increase germina¬
tion of basswood seed. Tree Planters’
Notes, 46:27-28.

Den Uyl, D., O. D. Diller, and R. K.
Day. 1938. The development of nat¬
ural reproduction in previously grazed
farmwoods. Purdue Univ. Agric. Exp.
Sta. Bui. 431, 28 pp.

Hart, A. C. 1958. Report on 1957 forest
tree seed crop in New England. U. S.
Forest Service. Northeastern For.
Exp. Sta., For. Res. Note No. 79, 2 pp.
Hart, A. C. 1959. Reports on 1958 for¬
est tree seed crop in New England.
U. S. Forest Service. Northeastern

For. Exp. Sta., For. Res. Note No. 86,

2 pp.

Johnson, L. P. V. 1946. A practical
method of overcoming seed dormancy
in Tilia americana L. Forestry Chroni¬
cle 22:182-190.

McConkey, T. W. 1960. Report on 1959
forest tree seed crop in New England.
U. S. Forest Service, Northeastern

For Exp. Sta,, For. Res. Note No. 96,

3 pp.

McConkey, T. W. 1961. Report on 1960
forest tree seed crop in New England.
U. S. Forest Service, Northeastern

For. Exp. Sta., For. Res. Note No. 115,
3 pp.

Rudolph, P. O. 1950-1961. Reports on
forest tree seed crop in the Lake
States. U. S. Forest Service, Lake
States For. Exp. Sta. Tech. Notes No.
333, 349, 370, 393, 412, 426, 447, 501,
540, 565, 574, and 598.

Spaeth, J. N. 1934. A physiological

study of dormancy in Tilia seed.
N. Y. Agric. Exper. Sta. (Ithaca).
Memoir 169, 78 pp.

U. S. Dept. Agric. 1948. Woody-Plant
Seed Manual. Misc. Publ. 654. G. P. O.,
Washington, D. C., 416 pp.

COMPARATIVE EFFECTIVENESS OF DDT SELECTION
METHODS IN DROSOPHILA MELANOGASTER MEIGEN

THOMAS R. KALLSTEDT and JACK BENNETT

Northern Illinois University

Insect resistance to a variety of
poisons has been known since the
early part of the century. Brown
(1957, 1958) has summarized the
widespread resistance of various in¬
sects to the newer synthetic insec¬
ticides. This study bears on two
questions concerning evolution of
resistance to DDT by Drosophila
melanogaster Meigen. It is of im¬
portance (1) to know whether a
mixed population of flies known to
contain genes (from several sources)
conferring resistance to DDT, would
show different rates of gain and level
of resistance under different meth¬
ods of selection applied under simi¬
lar culture conditions, and (2) to
determine the maximum degree of
resistance obtainable by selection of
such a population.

Methods and Materials

Three strains of DDT-resistant
Drosophila melanogaster were taken
from stock culture in October, 1959
and mixed to form one heterogene¬
ous stock. The three lines, HL2-top
(Bennett, 1960), Brown eye-R
(Crow, 1954), and ORS-1001 (King,
1957), had been cultured and tested
earlier by Bennett (1960, and un¬
published data). The mixed stock
was then divided among three popu¬
lation cages.

The flies used in this study were

raised in 8 drain straight-walled
glass shell vials (25 x 95 mm), each

of which contained approximately
3A °f an inch of food medium, and
polyethylene population cages (Ben¬
nett, 1956). The standard food medi¬
um consisted of 18 g of agar, 60 cc
of sugar, 100 cc of brewer’s yeast
in 1000 cc of water. For the first
five generations, 5 cc of propionic
acid was added as a mold inhibitor,
for the last 5 generations, 15 cc of
10% Moldex in alcohol was substi¬
tuted. Culture vials were seeded
with dry yeast (Schlitz, brewer’s
yeast) before use. This study cov¬
ered a ten month period (October,
1959 to July, 1960).

In the ninth generation tests, a
disproportionately large kill oc¬
curred when the temperature in the
incubator rose from the normal of
25° to 29° C (a known cause of in¬
creased mortality, Barker, 1957).

The “holding food” was similar
to the above described “regular”
food except that the yeast was omit¬
ted. The test vials contained filter
paper impregnated with DDT crys¬
tals deposited from acetone solution.
A test set consisted of three such
vials with concentrations of 1, 25,
625 /x g DDT/cm2. Data from each
test set covered three concentrations
and LD50 value were thus estab¬
lished. General testing procedures
followed those used by Bennett
(1960), and Coomes and Bennett
(I960). The LD50 value of a par¬
ticular test set was based on the
performance of at least nine and nor-

Comparative Selection Methods

125

mally 18 flies. At least three flies per
vial were tested and not more than
six flies per vial were used.

In the sib-selection line, values
were obtained for each sibship based
on one test set. These values pro¬
vided the basis for selection of the
sibships to provide parents for the
next generation. The values pre¬
sented here are based on summation
of survival values at each concen¬
tration in all of the test sets used
for a particular line or cage in each
generation.

All testing was done on female
flies as Crow (1954) has shown that
the results obtained are more re¬
producible than when males are
used. The females tested were of
varying ages. Those from the proge¬
ny of pair matings raised in stand¬
ard food vials were approximately
two days old.

Methods of Selection

Tests were made on each of the
three cages for two consecutive gen¬
erations to determine a base point
of DDT resistance for each cage.

Cages 1 and 2 were designated as
control cages and cage 3 as the per¬
manent DDT cage A DDT vial
(3,050 fig DDT/cm2) was attached
to this cage throughout the period
of study, beginning with test gen¬
eration one. A new DDT paper was
introduced twice, in the fifth and
ninth generations. Eight to twelve
food vials were attached to each
cage. Samples were obtained from
each cage at every generation by
etherizing the whole cage.

After determining the base point
of resistance, a direct selection line
was established by taking the top

30 females which survived the high¬
est concentrations of DDT from the
sets from cage 3. Males were taken
directly from cage 3. Of the 30 fe¬
males, six were placed in each of
five vials along with males. After a
two-day period of egg laying, the
flies were transferred into another
five vials, providing a larger popu¬
lation of flies for testing. Each suc¬
ceeding generation was established
in the same way without going back
to cage 3. Thus, female survivors of
a given test were used as parents of
the next generation and were mated
to untested males of the same gen¬
eration. This provided direct selec¬
tion for DDT-resistance exhibited by
survivors (and male offspring of the
previous generation’s survivors) that
had been raised in standard food
vials.

The sib-selection line was estab¬
lished with 60 pair matings in stand¬
ard food vials from cages 1 and 3.
Thirty females from cage 1 were
mated to 30 males from cage 3. The
reciprocal crosses were also made,,
insuring a comparable sample from
both cages. Thus, each of 60 pairs
of flies was allowed to produce one
progeny (sibship, family) in one
food vial. Of the 60 the 40 largest
progenies were tested for DDT tol¬
erance. Nine to 18 females were
taken from each progeny and tested.
After the results were recorded, the
progenies were ranked according to
their DDT tolerance, based on the
performance of the females tested.
The top 20 progenies were broken
down into the ‘‘top five” and the
“next 15”. From the top five cul¬
tures showing most resistance, 30
pair matings were made, using six
pairs from each progeny. The next

Transactions Illinois Academy of Science

126

15 progenies each contributed two
males and two females (30 pair mat¬
ings) to the next generation. For
the first five generations males and
females were taken from the same
progenies, thus inbreeding by broth¬
er-sister mating. In the last five
generations, the males and females
from each progeny were mated seri¬
al lv. In each case the flies were

mated with one pair to each food
vial. The result was 60 vials, each
with a pair of parents, to produce
60 progenies, 40 of which would be
tested the next generation. This in¬
direct selection was repeated in each
generation. Individuals of the germ
line were never exposed to DDT.

In each generation, 40 test sets
were made from the sib-lines raised
in vials, 20 test sets from the direct
lines raised in vials, 20 test sets
from control cage 1, and 20 test sets
from cage 3, the permanent DDT
cage. Any remaining test sets were

Fig. 1. — Results of testing sib-selection
line for DDT tolerance for 18 hour
period. Mean and 95% confidence limits
indicated.

g LD^q bogg Doae DDT)

Fig. 2.- — Results of testing direct se¬
lection line for DDT tolerance for 18
hour period. Mean and 95% confidence
limits indicated.

used for the testing of control cage

2. Cages 1 and 3 never required
20 test sets, so in each generation a
number were used for control cage 2.

Results

The principal results of this study
are summarized in Figures 1, 2, and

The course of selection through 10
generations of the sib-selection line
is shown in Figure 1. During the
first five generations of selection,
brother-sister pairs were used as par¬
ents. This inbreeding prevented any
net gain in resistance. Starting with
the parents of generation 6, random
mating was instituted. All of the
observed gain of resistance in this
line occurred following this change
of mating system.

The direct selection results are
shown in Figure 2. No change of
mating system occurred in this line

Comparative Selection Methods

127

• U>g10 ( U)50 DDT/om?, diffsrsnoe plus 100)

DDr oage O— — O
oib-Line O — - — ©

Direct-Line Q - ©

Pig. 3.— Results of the difference (plus
100) between the values (In w g DDT/
cm-) of the control cages and the ex¬
perimental lines tested for DDT toler¬
ance for 18 hour period.

and, despite gross fluctuations, the
gain was more evenly distributed
through the selection period.

Figure 3 presents the material in
different form, and with the results
from cage 3 for comparison. The
base line in Figure 3 was estab¬
lished by taking the combined test
results of the two control (unselect¬
ed) population cages (1 and 2), sub¬
tracting that value for each genera¬
tion from the values of the other
indicated lines, adding 100 to elimi¬
nate negatives, and expressing the
results as logarithms to the base 10.

Discussion

The second objective of this study,
to produce a selected line of Droso¬
phila melanogaster more resistant

than any previously tested, was
clearly achieved. The LD50 of the
direct selection line in the 10th gen¬
eration was 5.70 (log5 unit dose
DDT; 385.6 p g DDT/cm2) and of
the sib-selection line, 4.81 (log5 unit
dose DDT; 189.92 p g DDT/cm2).
In a comparative study of resistant
lines from Japan and two labora¬
tories in the U.S. (Bennett, 1960)
the two most resistant stocks showed
values of 4.50 and 5.49 (log5 unit
dose DDT ; 55.9 and 276.8 p g DDT/
cm2). Thus the most resistant line
in this study was nearly 40% more
resistant than any previously re¬
ported line. The three DDT resist¬
ant lines that were the progenitors
of the starting population in this
study had the following LDd0-s when
tested in 1957 (Bennett, 1960) :
ORS 1001, 5.49 (276.8 p g DDT/
cm2) ; Brown-eye-R, 3.68 (14.99 p g
DDT/cm2) ; HL2-Top, 3.35 (log*
unit dose DDT ; 67.8 p g DDT/cm2).
The values achieved by direct selec¬
tion surpassed the highest value
shown by these lines in the past.
This is interpreted as indicating that
some degree of integration of the
resistance factors of the parental
lines had been achieved, combining
separate resistance mechanisms for
a superior total resistance.

Comparative tests using the mos¬
quito test kits provided by the
World Health Organization showed
that the least resistant parental
line used here required several times
the DDT exposure recommended for
resistant mosquitoes to achieve a
significant kill. Thus it is apparent
that Drosophila melanogaster has
achieved much higher tolerance to
this insecticide than have mosquitoes
(Coomes and Bennett, 1960).

128

Transactions Illinois Academy of Science

The primary objective of this pa¬
per has been only conditionally sat¬
isfied. The final degree of resistance
exhibited by the direct selection line
was considerably higher than that
of the sib-selection line. The in¬
crease in DDT resistance over that
of the starting population was 23-
fold for the direct-selection line and
13-fold for the sib-selection line.
However, the inbreeding in the first
five generations of selection in the
sib line prevented any net increase
in resistance, so the 13-fold gain was
attained in the final 5 generations.
This compares with a 5.5-fold in¬
crease for the direct line in the first
five generations and a 4-fold increase
in the final five generations. Thus
one could argue that sib-selection had
demonstrated greater rate of gain
during the final five generations than
direct-selection produced in either
5 generation period.

The evidence thus does not provide
a delineation of the relative effec¬
tiveness of the two selective methods.
It is clear that both methods can be
highly effective under the conditions
used.

Figure 3 shows that the popula¬
tion in the DDT Cage did not gain
in resistance during this study. This
population was highly resistant at
the start and it appears that the rate
of kill (observed to be very low)
produced by the DDT lined vial in
the cage was so low as to provide no
effective degree of selection. This
is of interest because Bennett (1960)
had attempted earlier to compare
effectiveness of sib-selection in pair
matings in vials with direct-selection
in population cages (of a different
design than those used here). The
comparison did not seem a good one

at the time, but was used as the only
one available in the data at hand.
In this study we have been able to
make a partial comparison using
flies raised in vials on the same
batches of food. We know that a
difference in population density ex¬
isted in the culture vials of the
direct and sib-selection lines. In
future work the differences of popu¬
lation density and of mating pattern
will have to be dealt with.

Acknowledgment

The authors wish to acknowledge
the support of the National Science
Foundation, through Grant No. 8708,
and of the Northern Illinois Uni¬
versity Biology Department, in pro¬
viding laboratory facilities.

Summary

A heterogeneous population of
Drosophila melanogaster was pro¬
duced by mixing three DDT-resist-
ant strains. This hybrid population
was divided into snb-populations by
culturing them in two control popu¬
lation cages and a third population
cage containing a permanent DDT
lined vial. A sib-selection line and
a direct-selection line were tested for
DDT resistance each generation. Se¬
lection was carried on for ten genera¬
tions. During the first five genera¬
tions the sib-line was inbred (broth¬
er-sister pair matings) whereas the
last five generations out-breeding
(between familities within the line)
was used.

At the end of the selection period,
the DDT cage population showed a
DDT tolerance 0.46 times that of the
starting population. The sib-selected

Comparative Selection Methods

129

line showed no increase in the first
five generations due to inbreeding,
but a 13-folcl increase in tolerance
was attained by five generations of
out-breeding and selection. The di¬
rect-selection line reached a 23 -fold
increase in resistance at the end of
ten generations of selection. How¬
ever, in the five generations of ef¬
fective selection, the sib-selection line
increased resistance by more than
twice as much as the direct-selection
line in a comparable five generations.

The direct-selection line attained
a higher degree of tolerance to DDT
than any single resistant strain
tested earlier. Thus the different re¬
sistant genotypes were recombined
in a way which yielded a higher
degree of resistance than had been
attainable by any one selected line.

In this study sib-selection appears
to be a more effective method of
selecting DDT-tolerant genotypes
than the direct-selection method.

Literature Cited

Barker, Roy J. 1957. Some Effects of
Temperature on Adult House Flies
Treated with DDT. Journal of Eco¬
nomic Entomology 50 (4): 446-450.

Bennett, Jack. 1956. Inexpensive Popu¬
lation Cages. Drosophila Inf ormation
Service 30: 159-60.

Bennett, Jack. 1960. A comparison of
Selective Methods and a Test of the
Pre-Adaptation Hypothesis. Heredity
15(1): 65-77.

Brown, A. W. A. 1957. Insecticide Re¬
sistance and Darwinism. Botyu-Kaga-
ku , 22: 277-282.

Brown, A. W. A. 1958. Insecticide Re¬
sistance in Arthropods. W. H. O.
Monograph Ser., No. 38, 240 pp.

Coomes, R. K. and Jack Bennett. 1960.
Use of World Health Organization
Mosquito Test Kit with DDT Resist¬
ant Drosophila. Trans. III. St. Acad.
Sci. 52 (3 & 4) : 151-155.

Crow, J. F. 1954. Analysis of a DDT-
resistant Strain of Drosophila. Jour¬
nal of Economic Entomology 47: 393-
398.

King, James C. 1957. Investigation of
the Genetic Nature of Resistance to
Insecticides Developed by Populations
of Drosophila melanogaster. Final re¬
port of research carried out by Long
Island Biological Association for Medi¬
cal Research and Development Board,
Office of the Surgeon General, Depart¬
ment of the Army.

OXYGEN CONSUMPTION IN THE SMALL, SHORT-TAILED

SHREW (CRYPT OTIS PART A)

CARL J. PFEIFFER and GEORGE H. GASS
Southern Illinois University, Carbondale

Interest in the increase of meta¬
bolic rate with decrease in body size
of mammals and other organisms
has stimulated investigation of oxy¬
gen consumption in very small mam¬
mals. Several species of the shrew
are representative of the smallest
living mammals, and consequently
are unique for such experimenta¬
tion.

Comparative studies of oxygen
consumption of the wandering shrew
( Sorex vagrans vagrans), the Mon¬
terey shrew (Sorex irowbideii mon-
teryensis) , and the Sonoma shrew
(Sorex pacificus sonomae) have been
reported by Pearson (1948). Also,
the rate of oxygen consumption has
been determined for the long-tailed
shrew (Sorex c. cinereus) by Mor¬
rison (1948). Reports are lacking,
however, in regard to one of the
smallest species of shrews, the small,
short -tailed shrew (Cryptotis par-
va) . Thus, it is the purpose of the
present communication to report on
the oxygen consumption of three
small, short-tailed shrews, as deter¬
mined in the apparatus of Watts
and Gourley (1953).

Materials and Methods

Three captive wild specimens of
the small, short-tailed shrew (Cryp¬
totis parva) were utilized for all
experiments. One adult female of
undetermined age was tested in the
post-lactational stage, and two, six-

to seven-week old weanlings from the
litter of the adult female were
studied initially. Oxygen consump¬
tion determinations were subsequent¬
ly carried out on one of the shrews
from the litter at the age of about
four months. At this time the shrew
was full-grown and was proven to
be a male. A total of eighteen ex¬
periments were run. All animals
were sustained on live insects,
ground beef, and water. However,
the shrews were not fed two to three
hours prior to the experiments in
order to provide a post-absorptive
state.

The apparatus employed was simi¬
lar to that used by Watts and Gour¬
ley (1953), except for the following
modifications. A 375 ml dark-tinted
jar with a number 10 rubber stopper,
a standard 5 ml pipette, and an
aluminum screen wire grid were
utilized. In addition, the cylindrical
wire compartment was eliminated so
that the shrew was allowed freedom
of movement. This apparatus util¬
izes a layer of soda lime (8 mesh)
beneath the wire grid to absorb car¬
bon dioxide. The rate of oxygen
consumption can be determined since
the volume of oxygen consumed is
measured by the excursion of a move-
able soap film in the pipette. Watts
and Gourley (1953) have demon¬
strated with the rat that this ap¬
paratus is of adequate reliability and
sensitivity for determination of oxy¬
gen consumption in small mammals.

[ 130]

Oxygen Consumpt w n

131

Ten minutes were allowed for tem¬
perature equilibration, and the dura¬
tion of each experiment was equal
to the time for the utilization of 5 ml
of oxvgen.

Results

The results are given in Table 1.
It is evident that oxygen consump¬
tion increases greatly with increased
activity. Indeed, the rate of oxygen
consumption in the adult male shrew
almost doubled from a basal rate of
7.0cc/g/hr to 13.2cc/g/hr under con¬
ditions of vigorous activity. Due to
the normal incessant activity of
shrews in the waking state, it is
probably impossible to simulate con¬
ditions with the shrew that are char¬
acterized as basal with other less
active mammals. It was found that
slight changes in activity of the test

animals caused immediate alterations
in oxygen uptake, as registered with
the soap film excursions. Shrews in
the respirometer generally were very
active and frequently chewed on the
screen grid or rubber stopper. In
a few instances general activity was
relatively low and as indicated in
Table 1, a subjective rating of slight
activity was applied to shrews at
comparative rest.

Discussion

It has been demonstrated by Ham¬
ilton (1944) and others that the rate
of digestion in shrews is exceedingly
rapid. In one instance Hamilton
(1944) observed that the passage of
chitin through the alimentary tract
in a captive, non-fasted shrew re¬
quired only 95 minutes. The shrews
used in the present investigation

Table 1. — Oxygen Consumption of Small, Short-Tailed Shrew
( Cryptotis pay'va) as Related to Activity.

Animal

Weight

(g)

Air

Temp.

(°C)

Number

Experiments

Standard

Deviation

Activity1

Mean O2
Consum.
(cc/g/hr)

Adult

Female .

6.02

.39

Moderate

9.4

Adult

Female .

6.02

.97

Vigorous

11.4

Adult

Male .

6.36

.78

Slight

7.0

Adult

Male .

6.36

.57

Vigorous

13.2

Immature

Shrews2 .

a 4. 59

Vigorous

11.9

b 4. 73

• •

Vigorous

10.4

1 Activity was subjectively rated : slight, shrew at rest ; moderate, shrew walking or
chewing on grid half time ; vigorous, shrew constantly and vigorously chewing on grid.

2 Sex undetermined.

132

Transactions Illinois Academy of Science

probably were in a post-absorptive
state, since they had not been fed
two to three hours prior to each de¬
termination. However, true basal
conditions were approached only
during five experiments with the
adult male shrew at the periods of
least activity. This lack of muscular
repose in the shrew during metabo¬
lism determinations has also been
encountered by other investigators
(Morrison, 1948).

The results of the present report
are in accord with those of other
investigators (Morrison, 1948 ; Pear¬
son, 1938). The oxygen consumption
of 7.0cc/g/lir for the 6.36g Crypto-
tis parva at basal conditions falls on
the curve constructed by Pearson
(1948) where oxygen consumption of
small mammals is plotted as a func¬
tion of body weight. Also, the high
oxygen utilization of 13.2cc/g/hr
obtained during those experimental
runs characterized by vigorous shrew
activity can be compared to the oxy¬
gen utilization of 13.7cc/g/hr for
the long-tailed shrew (Morrison,
1948).

Summary

The normal rate of oxygen con¬
sumption was determined in three
small, short-tailed shrews ( Cryptotis
parva), one of the smallest species
of shrews. Determinations were ob¬
tained for both immature and adult
male and female shrews in the post-
absorptive state, and in varying de¬
grees of activity. A simple closed
chamber basal metabolism apparatus
was utilized for the determinations.
Mean oxygen consumption rates of
7.00cc/g/hr in the resting state and
as high as 13.2cc/g/hr in states of
vigorous activity were calculated.

Literature Cited

Hamilton, W. J. 1944. The biology of
the little short-tailed shrew, Cryptotis
parva. J. Mammal., 25:1.

Morrison, P. R. 1948. Oxygen consump¬
tion in several mammals under basal
conditions. J. Cell, and Comp. Physiol.,
31:281-292.

Pearson, O. P. 1948. Metabolism of
small mammals, with remarks on the
lower limit of mammalian size. Sci¬
ence, 108: 44-46.

Watts, D. T., and D. R. H. Gourley.
1953. A simple apparatus for deter¬
mining basal metabolism of small ani¬
mals in student laboratory. Proc.
Soc. Exp. Biol, and Med., 84:585-586.

NOMENCLATURE OF THE LATE MISSISSIPPIAN WHITE
PINE SHALE AND ASSOCIATED ROCKS IN NEVADA

R. L. LANGENHEIM, JR.

University of Illinois, Urbana

Late Mississippian detrital rocks
in eastern and southern Nevada and
adjacent California have been as¬
signed to at least nine different for¬
mations and there is no concensus
regarding their classification. Al¬
though our current investigations
are incomplete (Langenheim, 1956a,
1956b, 1960) my own prior use of
the classification proposed in this
paper (Langenheim and Tischler,
1960; Langenheim et al., 1960), im¬
minent use by fellow workers and
needs arising from preparation of a
correlation chart for the Great Ba¬
sin by a committee of the Eastern
Nevada Geological Society require
preliminary publication of nomen-
clatorial problems regarding these
rocks.

The Late Mississippian detrital
rocks of the central Great Basin
consist of a basal calcareous silt-
stone unit, a black fissile shale unit,
a sandy shale unit and an upper
sandstone and conglomerate unit
(Figs. 1, 2). The sequence as a whole
generally thickens toward the west
and the upper, coarser members in¬
crease in relative importance. The
basal, calcareous siltstone unit, how¬
ever, thickens eastward and at least
one interbedded limestone unit oc¬
curs within the black fissile shale
unit in eastern Nevada. The entire
detrital sequence rests clisconform-
ably on Early Mississippian or older
rocks and appears gradational with

overlying later Paleozoic carbonate
rocks. A disconformity may be
present either at the base of the
sandy shale unit or within the con¬
glomerate and sandstone unit, but
this has not been fully demon¬
strated.

History of the Nomenclature

The first significant account of the
late Mississippian rocks in the
Great Basin is that of the King
Survey (Hague, 1870; Hague and
Emmons, 1877; King, 1876), in
which Hague (1870) mapped and
described the White Pine Mining
District. Here he recognized, in as¬
cending order, a calcareous shale,
a siliceous limestone, an argillaceous
shale divided into a lower “ bitum¬
inous ” portion and an upper sandy
portion, a “ reddish yellow” sand¬
stone and Carboniferous limestone.
Hague ’s map and descriptions
(1870: 409-421, Atlas Sheet 14) per¬
mit ready identification of his units
on the ground and correlation with
Humphrey’s (1960) recent descrip¬
tion and map. Furthermore, Hague
appears consistent in his own term¬
inology and apparently always re¬
fers to these units as “ calcareous
shale”, “siliceous limestone”, etc.
in later publications and in con¬
junction with formally proposed
stratigraphic names. Thus the
“White Pine Shale” of Hague is

[133]

134

Transactions Illinois Academy of Science

always described as composed of
argillaceous and sandy shale in the
White Pine District.

Hague (1882) first used the name
“White Pine Shales" in a brief ad¬
ministrative report in which he re¬
fers to “black argillaceous and
arenaceous shales which overlie the
limestone” to the “White Pine
Shales.” He also states that these
rocks are named “White Pine shales
from the locality where they were
first recognized in Eberhardt Can¬
yon, and underlying the town of
Hamilton” (Hague, 1882: 28). In
later work Hague refers to sections
in Applegarth Canyon which is a
strike valley in the White Pine
Shale trending south from Hamil¬
ton. Applegarth Canyon is shown
as the upper part of Cathedral Can¬
yon in the Treasure Hill (edition
1950) and Illipah (edition 1951)
15 minute series topographic maps
of the U. S. Geological Survey. One
year later, Hague (1883), in an
“Abstract” of his forthcoming
monograph on the geology of the
Eureka District, described the
White Pine Shale as a “heavy body
of black shale . . . having been first
recognized as a distinct horizon in
the White Pine Mining District.”
This latter reference is cited in Wil-
martli (1938) as the first reference
to the White Pine Shale. Although
these two citations and the 1870
description of the White Pine Dis¬
trict clearly identify the rocks upon
which Hague’s concept of the White
Pine Shale rests, some confusion has
arisen because the 1882 and 1883
papers were chiefly concerned with
the geology of the Eureka District.
This has led some to accept expo¬
sures in the southern Diamond

Range as the type locality or the most
important reference locality for the
White Pine Shale concept.

In 1883 Hague first applied the
name, “Diamond Peak Quartzite”,
to the sandstone and conglomerate
resting on the White Pine Shale at
Eureka and in the AYliite Pine Dis¬
trict. Previously these rocks had
been referred to as “reddish yellow
Sandstone” (Hague, 1870), “Ogden
quartzite” (Hague in King, 1880:
27) and “Weber quartzite”
(Hague, 1882).

In 1892 Hague fully described the
Eureka District, redescribed the
rocks of the White Pine District and
discussed correlation between these
two areas and other localities. The
section at Sugar Loaf in Packer
Basin near Eureka, important in
nomenclatorial problems, is de¬
scribed on page 81 as paraphrased
below :

Top

1. Shaly sandstone with interbedded
shale and conglomerate. .1,000 feet.

2. Black argillaceous shale with gra¬
dational upper contact.. 400 feet.

3. Gray crinoidal, sandy limestone

with Chonetes . 50 feet.

4. Yellow-weathering, black, argilla¬

ceous and calcareous fossiliferous
shale . 300 feet.

5. Blue, fossiliferous limestone..

. 250 feet.

6. Siliceous limestone . 150 feet.

Bottom

Hague (1892: 80-81) somewhat am¬
biguously remarks that the “beds
directly underlying the shale are of
course the uppermost members of
the Nevada limestone.” From this
and from his correlation of the
Packer Basin section with the sec¬
tion in Applegarth Canyon in the
White Pine District, it is apparent
that units 1-4 are considered part of

White Pine Shale

135

the White Pine Shale and 5-6 are
part of the Nevada limestone.

Thus, in regard to the Applegarth
Canyon Section in the White Pine
District, Hague (1892: 193) states,
“A more characteristic White Pine
fauna is preserved in the black
shale than has yet been obtained in
the corresponding beds at Eureka,
and a belt of intercalated limestone
in the shale similar to that found
east of Sugar Loaf at Eureka bears
equal evidence of its Devonian age.
Here the limestone appears as a
lenticular body in the shale, with
beds identical in composition both
above and below.” These state¬
ments taken alone are also somewhat
ambiguous and it is impossible, out
of context, to be absolutely certain
whether the ‘Denticular body” of
limestone is in Applegarth Canyon,
if so it is certainly a body of lime¬
stone wholly within the “argil¬
laceous shale” and, therefore, part
of the White Pine Shale as typified
by Hague in the White Pine Dis¬
trict. If, however, one assumes that
the “lens” is in Packer Basin and
that the limestone in Applegarth
Canyon is a continuous bed, then it
may be argued that Hague had
modified his conception of the White
Pine Shale in Applegarth Canyon
and elsewhere in the White Pine
District to include the “calcareous
shale” and “siliceous limestone”.
This conclusion may be justified by
rigorous analysis of Hague’s gram¬
matical construction on page 193,
but it is rejected for the following
reasons: (1) If Hague included the
“siliceous limestone” and “calcar¬
eous shale” in Applegarth Canyon
within the White Pine Shale as
suggested by his phraseology on

page 193 of Monograph 20, this is
the only place wherein such a cor¬
relation is suggested in his writings.
Otherwise he is consistent in his
use of lithologic terminology and
restricts the White Pine Shale of
the AVliite Pine District to rocks
described as either “argillaceous,”
“arenaceous,” or “bituminous”
shale. (2) On page 193 Hague states
that the shale above and below the
“lenticular body” is of the same
composition, but on page 81 the
shale layers above and below the
limestone with Chonetes in Packer
Basin are described differently.
Thus the “lenticular body” must
be the limestone in Applegarth Can¬
yon.

This means that Hague, in 1892,
correlated units one through four
of his Packer Basin Section in the
Eureka District with the White
Pine Shale of the type area in the
AVliite Pine Mining District. Inas¬
much as Nolan et al. (1956) have
identified the limestone with Cho¬
netes in Packer Basin as the Joana
Limestone and unit 4 as the Pilot
shale, this means that Hague’s 1892
correlation is incorrect, according
to our persent knowledge of these
rocks, because he equates the upper
Mississippian shale of the AVliite
Pine District with the uppermost
Devonian shale, lower Mississippian
limestone and upper Mississippian
shale of the Eureka District. Fur¬
thermore, this miscorrelation has
been the source of much confusion
regarding the limits of the AVliite
Pine Shale, the upper Mississippian
shale unit in question.

Lawson (1906) followed Hague’s
correlations in describing the AVliite
Pine Shale in the Robinson (Ely)

Transactions Illinois Academy of Science

136

District. Lawson (1906:296) states,

‘ ‘ This shale formation, with its in¬
cluded limestone bed agrees well
with the descriptions that have been
given by Hague for the White Pine
shale of the neighboring White Pine
and Diamond Ranges . . . Even the
thick bed of limestone in the midst
of the shale has its analogue in the
White Pine shale of the Eureka and
White Pine sections.” Lawson re¬
fers to Hague’s 1892 discussion on
pages 192-193, thus making it plain
that he is referring to the lens of
limestone in Applegarth Canyon
and the limestone with Chonetes at
Packer Basin. Spencer (1917 : 25-26)
accepted Lawson and Hague’s cor¬
relations and assigned formal strati¬
graphic names to the three units
described by Lawson. The lower
shale, equivalent to Hague’s “cal¬
careous shale” of the White Pine
District, was named, “Pilot Shale.”
The middle limestone, equivalent to
Hague’s “siliceous limestone”, was
named, ‘ ‘ Joana Limestone, ’ ’ and the
upper shale, equivalent to Hague’s
“argillaceous shale”, was named,
“Chainman Shale.” Thus Lawson
(1906) and Spencer (1917) correct¬
ly correlate rock units in the Ely
District with those of Hague’s Pack¬
er Basin section near Eureka, but
are incorrect in company with
Hague, in comparing these sections
with the type White Pine Shale in
the White Pine Mining District
The Chainman Shale in the Rob¬
inson Mining District consists al¬
most entirely of black, fissile shale
(Fig. 2). Thus the poorly exposed,
thin calcareous siltstone unit is not
mentioned by Lawson (1906) or by
Spencer (1917). In addition, the
relatively thin sandy shale unit is

Fig. 1. — Location Map.

only briefly discussed by Spencer
(1917) who also points out the ab¬
sence of the Diamond Peak Quart¬
zite in the mining district.

In 1932 Westgate (Westgate and
Knopf, 1932: 19-21) described the
sequence at Silverhorn in the Pioclie
District and at Dutch Jolm Moun¬
tain about 40 miles north of Pioche.
Here Westgate assigned the detrital
rocks between the Lower Mississip-
pian Bristol Pass (Joana or “sili¬
ceous”) Limestone and the Late
Mississippian or Early Pennsylvan¬
ian Bailey Spring Limestone to the
newly-described Peers Spring For¬
mation and Scotty Wash Quartzite.
Although the Scotty Wash Quart¬
zite is analagous to the Diamond
Peak Formation, it is of slightly
differing composition, is presumably

White Pine Shale

137

derived from a different source
area, and probably is geographically
separated from the Diamond Peak
Quartzite (James, 1954). The Peers
Spring Formation, however, occu¬
pies the same stratigraphic position,
includes rocks of the same sort and
is presumably geographically con¬
tinuous with the Chainman Shale
as defined by Spencer (1917) and
the White Pine Shale in the White
Pine Mining District as defined by
Hague (1882, 1883, 1892) (Langen-
heim and Peck, 1960).

Westgate describes the Peers
Spring Formation as follows : ‘ ‘ The
most noticeable type (of rock ) is a
black, dense, fine-grained limestone,
much of it gray-white on the weath¬
ered surface, very thin bedded,
locally almost shale-like in its lam¬
ination. Probably a more common
facies, though not so often seen in
actual outcrop because it is a softer
rock, is a brown calcareous shale
that effervesces slightly in warm
hydrocholoric acid. The formation
as a whole weathers easily, so that
outcrops are scarce, the surface being
covered with fine gray, lavender, or
rusty debris. Interbedded in the
shale and thin limestones of the
lower part of the formation are
blue-black limestones, some layers
of which are 4 feet thick. These
seem to form a transition by inter¬
calation to the underlying Bristol
Pass limestone.” (Westgate and
Knopf, 1932: 20). Re-examination
of the type and reference areas
(Langenheim and Peck, 1960) has
shown that accidents of exposure
make outcrops of the calcareous silt-
stone unit (Fig. 2) most extensive in
the Peers Spring area and, as sug¬
gested by Westgate (Westgate and

Knopf, 1932: 20), give an erroneous
impression regarding the formation.
Thus, although Westgate ’s descrip¬
tion pertains almost entirely to the
calcareous siltstone unit, all three
units — calcareous siltstone, black
fissile shale, and sandy shale — are
present. These relationships are
difficult to detect at Peers Spring
but are well displayed at Dutch
John Mountain (Langenheim and
Peck, 1960).

Problems of exposure also led
Westgate (Westgate and Knopf,
1932) to misinterpret the basal con¬
tact of the Peers Spring Formation
at Peers Spring and the Peers
Spring-Scotty Wash contact at
Dutch John Mountain (Langenheim
and Peck, 1960). Lower Mississip-
pian fossils reported from lenses in
the basal Peers Spring Formation
at Peers Spring are actually from
infaulted blocks of Bristol Pass
Limestone and the great thickness
of Scotty Wash Quartzite reported
at Dutch John Mountain appears
largely to result from inclusion of
talus-covered shale slopes within
the quartzite formation (Langen¬
heim and Peck, 1960).

In 1953 the stratigraphic correla¬
tion committee of the Eastern Ne¬
vada Geological Association (Easton
et al., 1953) accepted Hague’s
(1883) designation of the White
Pine Mining District as the type
area for the White Pine Shale. The
Pilot Shale, Joana Limestone and
Chainman Shale, however, were in¬
cluded as members within the White
Pine Formation at Ely and Eureka
(Easton et al., 1953, fig. 2) and, by
implication, within the White Pine
Mining District as well. In their
column for the Pioche District

138

Transactions Illinois Academy of Science

(Easton et al ., 1953, fig. 2) the Pilot
Shale and Joana Limestone (Bristol
Pass Limestone) are shown as mem¬
bers of the White Pine Formation,
but the Peers Spring Formation is
treated separately because, “It is
probable that the Peers Spring for¬
mation includes the Chainman shale
and part of the Diamond Peak for¬
mation.7’ (Easton et al ., 1953: 149).
Thus, Hague's (1892), Lawson’s
(1906) and Spencer’s (1917) mis-
correlation of the Eureka and Ely
sections with the White Pine Shale
of the White Pine District was re¬
inforced. Also, by implication, the
type section in the White Pine Dis¬
trict was revised to include the
“siliceous limestone” and “cal¬
careous shale” previously assigned
to the Nevada Formation by Hague
(1870, 1892) and King (1878).

McAllister (1952: 22-26) created
local formations in dealing with
Late Mississippian detrital rocks in
the Quartz Spring Area, northern
Panamint Range, California. Cal¬
careous siltstone and shale with
interbedded limestone is included
in the upper part of the Perdido
Formation. Black argillaceous shale
resting on a Cravenoceras biostrome
at the top of the Perdido Formation
is assigned to the Rest Spring Shale.
The upper part of the Rest Spring
Shale is silty, includes minor inter¬
beds of cpiartzite and is succeeded
by the Pennsylvanian Tilivipah
Limestone. Langenheim and Tiscli-
ier (1960: 110, fig. 5) have rede¬
scribed the Perdido Formation in
greater detail and correlated the
Upper Perdido Formation with the
regional calcareous siltstone. The
Rest Spring Shale is correlated with
a Chainman Shale concept restricted

to the “argillaceous and arenaceous
shale” of the White Pine District.

Nolan et al. (1956) review the
nomenclature of the White Pine
Shale in a restudy of the Paleozoic
section in the Eureka District.
Nolan et al. (1956: 54) cite the
White Pine Mining district as the
type locality of the White Pine
Shale. They appear, however, in¬
decisive regarding Hague’s concep¬
tion of the AVhite Pine Shale in the
Eureka District. On one hand they
state, in reference to the Joana
Limestone in the Eureka District,
that “although the unit was clearly
recognized in a section measured in
Packer Basin, southeast of Eureka
(Hague, 1892: 81). It is not clear
from this reference, however, if
Hague intended to include the Jo¬
ana with the underlying Devonian
sedimentary rocks, or with his
White Pine Shale.” (Nolan, et al.,
1956: 54). Elsewhere, they state in
regard to the Pilot Shale, “The
lowest beds of the White Pine Shale,
as defined by Hague (1892: 68-69),
differ in lithologic character from
the rest of the beds that were as¬
signed to that unit and appear to
be equivalent in stratigraphic posi¬
tion and relationships to the Pilot
shale of the Ely district (Spencer,
1917 : 26).” (Nolan et al., 1956 : 52).
Although Hague’s (1892: 81) dis¬
cussion of the Packer Basin section
regarding assignment of the lime¬
stone with Chonetes (Joana Lime¬
stone), appears ambiguous out of
context, his correlation of this sec¬
tion with the Applegarth Canyon
section (Hague, 1892: 193) clearly
indicates that he placed the lime¬
stone with Chonetes , and the shale
below in the White Pine Shale, as

White Pine Shale

139

Duckwater

Hamilton

Secret

Canyon

Diamond

Peak

Bock

Pass

Permian

Pennsylvanian Is

Devonian Ls.

Lund

3kT sholy interval

Crystal Spring Kane Spring Devonian
Wash Wash

(After Duley,

1957)

Pennsylvanian Is.

quortzite

1060

y?''

270'

sandy shale

650'

black fissile shale

220'

— ”

... .

—

I701

240'

□vP

crinoidal limestone

Cerro Ubehebe

Gordo Mine

I /

(After Ross. 1956)

Quartz

Spring

( Rest Spring Shale

3000

. 2000

_ 1000

Fig. 2. — Columnar sections of Late Mississippian detrital rocks

in the central Great Basin.

140

Transactions Illinois Academy of Science

Pig. 3. — Graphic summarization of the nomenclatorial history of the Late
Mississippian rocks of the central Great Basin.

White Pine Shale

141

he conceived it in the Eureka Dis¬
trict.

Nolan et al. (1956) further point
out that most workers were con¬
fused regarding the nature of the
White Pine Shale and that many
used the name, “White Pine” in¬
correctly. Therefore, they proposed
rejection of the White Pine Shale
concept and adopted the Chainman
Shale concept for use in the Eureka
District, because :

1) Hague’s White Pine Shale and
Diamond Peak Quartzite were not
satisfactory mapping units for their
work in the Eureka District. These
units show extreme differences in
thickness and lithologic character
both between and within individual
thrust plates.

2) The White Pine Shale has been
applied to black shale sequences of
excessively wide range as, for ex¬
ample. in the Eureka District where
it became necessary to remove the
Pilot Shale and Joana Limestone
from the unit mapped as White Pine
Shale by Hague.

3) Inclusion of the Pilot Shale,
Joana Limestone and Chainman
Shale as members within the White
Pine Formation, as employed by the
stratigraphic committee of the East¬
ern Nevada Geological Association
(Easton et al., 1953) “has the ad¬
vantage of retaining the name,
“White Pine” for the dominant
black shale sequence, (hut) does not
provide for satisfactory treatment
of the thick gradational zone be¬
tween the black shales and the
coarser elastics characteristic of
Hague’s Diamond Peak.” (Nolan,
et al., 1956:57). Nolan et al. (1956)
solve this problem by separately
recognizing the Chainman Shale and

Diamond Peak Formation where
they can be satisfactorily distin¬
guished and elsewhere referring to
the undivided sequence as “Chain-
man and Diamond Peak Formations
undifferentiated. ’ ’

Johnson and Hibbard (1957 : 356-
360) introduce another set of local
formations in mapping the A. E. C.
Proving Grounds. The Narrow Can¬
yon Limestone consists of platv,
buff -weathering silty limestone which
appears similar to the Lower Pilot
Shale of Langenheim (1961) and is
tentatively correlated with the Pilot
Shale by Johnson and Hibbard
(1957: 356). The Camp Mercury
Limestone is described as a prob¬
able correlative of the Joana and Tin
Mountain Limestones, but the Eleana
Formation is less readily compared
with rocks of other areas. Although
the Eleana Formation was not ob¬
served in stratigraphic contact with
the Camp Mercury Limestone, John¬
son and Hibbard (1957) assume it
to be stratigraphically above the
limestone. The lower part of the
Eleana Formation consists of black
shale and is compared with the
Chainman Shale of the Eureka Dis¬
trict as recognized by Nolan et al.
(1956) (Johnson and Hibbard.
1957). The middle quartzitic por¬
tion of the Eleana Formation is ten¬
tatively correlated with the Diamond
Peak Formation of Noland et al.
(1956), but no attempt was made
to compare the upper, shaly portion
of the Eleana Formation to rocks
elsewhere (Johnson and Hibbard,
1957). Poole et al. (1961) have re¬
vised this treatment of the Eleana
Formation after discoverv of a lo-
cality in which the Eleana Forma¬
tion rests disconformably on Devon-

142

Transactions Illinois Academy of Science

ian limestone. Thus they correlate
the basal Eleana Formation with the
Narrow Canyon and Camp Mercury
Limestone and approximately equate
the remainder of the Eleana Forma¬
tion with the Chainman Shale, Dia¬
mond Peak Formation and, possibly,
basal Ely Limestone of the Eureka
District (Poole et al., 1961, table
328.2).

Suggested Regional Terminology
for the Late Mississippian Rocks

of the Central Great Basin

Part of the confusion regarding
Late Mississippian stratigraphic no¬
menclature in the central Great Ba¬
sin arises from conflicting needs of
geologic mapping and those of basin¬
wide stratigraphic synthesis. Local
formational concepts are needed in
many localities for efficient mapping
within a mountain range or mining
district. These local concepts, how¬
ever, may not coincide with forma¬
tional units of greatest regional sig¬
nificance. In some instances, growth
of knowledge may “overtake” such
local concepts and it may be useful
either to abandon or revise them to
bring them into conformity with
regional units having the same local
value. The terminology of Johnson
and Hibbard (1957) and Poole et al.
(1961) refers to a set of such pres¬
ently useful local formational con¬
cepts. Spencer’s (1917) units at Ely
and Westgate’s (West gate and
Knopf, 1932) units at Pioche are
other examples in which the advance
of stratigraphic knowledge has made
the need for a local terminology
more or less obsolete.

Units for regional synthesis should
reflect the distribution of major rock
types within the basin because only

units such as these are of environ¬
mental and paleogeographic signifi¬
cance. In the central Great Basin,
the Late Mississippian detrital se¬
quence includes five major rock units
of this sort. They are the basal
calcareous siltstone unit, the black
fissile shale unit, the sandy shale
unit and the two geographically dis¬
tinct quartzitic and conglomeratic
units mentioned in the introduction
to this paper. Three of these units
were recognized by Hague in 1870
as the bituminous and sandv mem-
bers of the black argillaceous shale
and the reddish yellow sandstone.
The basal calcareous siltstone unit
is thickest in the Pioche District and
is the dominant lithologic type de¬
scribed by Westgate (Westgate and
Knopf, 1932) in his discussion of
the Peers Spring Formation. The
black fissile shale unit is widespread
throughout the area and constitutes
almost all of the Chainman Shale
as described by Spencer in the type
area. The sandy shale sequence is
thickest in the western part of the
area but has not been as vet the
basis of a separate formational con¬
cept. The uppermost quartzitic and
conglomeratic sequence includes the
Diamond Peak Quartzite. This for¬
mation is thickest in the Eureka Dis¬
trict and is separated by an area
in which quartzite is absent from
exposures of the Scotty Wash
Quartzite. Distribution of the Scotty
Wash Quartzite centers on the
Pioche District.

A rational classification for re¬
gional synthesis should include all
of these units and should also fol¬
low priority in definition and nam¬
ing of stratigraphic concepts. Thus
the White Pine Shale, raised to

White Pine Shale

143

Group status, is retained for the
entire fine-grained detrital sequence
and the Scotty Wash Quartzite and
Diamond Peak Quartzite concepts
are applied to the appropriate, sepa¬
rate coarse-grained rock bodies.

Retention of the White Pine Shale
as a Group is justified for the fol¬
lowing reasons :

1) Hague’s original statements in
1882 and 1883 clearly designate the
argillaceous shale of the White Pine
Mining District as the basis for this
formation concept. His map of 1870
and the more extensive discussions
of 1892 establish the type section of
the White Pine Shale as including
the entire body of fine-grained detri¬
tal rock stratigraphically above the
“siliceous limestone” and below the
Diamond Peak Quartzite.

2) The recommendation of the
Eastern Nevada Geological Asso¬
ciation Stratigraphic Committee
(Easton et al., 1953) that the White
Pine Shale be treated as comprising
the Pilot Shale, Joana Limestone
and Chainman Shale is rejected as
based on miscorrelations by Hague
(1892), Lawson (1906) and Spencer
(1917) of rocks at Eureka and Ely
with the type section of the White
Pine Shale in the White Pine Dis¬
trict.

3) The recommendation of Nolan
et at. (1956) that the name “White
Pine " be suppressed and the name,
“Chainman” be applied to the up¬
per shale is rejected because the
Chainman Shale of Spencer (1917)
is a junior synonym of the White
Pine Shale of Hague (1882, 1883,
1892) in the type area. The argu¬
ments of Nolan et al. (1956) to the
effect that Hague’s formational con¬
cept does not satisfactorily serve the

needs of geologic mapping in the
Eureka District and elsewhere are
considered in large part equally ap¬
plicable to use of the same forma¬
tional concept under a junior name.
These arguments are valid in respect
to the needs of local mapping and,
it may be pointed out, have been met
by Nolan et al. (1956) through rec¬
ognition of a local, undifferentiated
Chainman and Diamond Peak unit.

4) Finally, it has been widely
argued that the Chainman Shale con¬
cept is untainted by past confusion
regarding its application and that
current usage favors use of the
Chainman concept and name. This
argument is rejected because it is
felt that the White Pine concept of
Hague is in fact clearly recognizable
and that differences of opinion re¬
garding its application are capable
of resolution by comparison of the
rocks in question with those of the
type locality. In fact, the White
Pine Mining District section, select¬
ed as a reference section by Hum¬
phrey (1960), is superior to the
Chainman type section in that it is
better exposed, less metamorphosed
and deformed, and has a better rep¬
resentation of the three recognized
lithologic subunits within the se¬
quence. The appeal to general usage
is rejected as not being capable of
objective resolution.

The three lithologic units within
the White Pine Group now require
definition and designation of type
or reference sections. It is suggest¬
ed that the Peers Spring Formation
concept be restricted to the basal
calcareous siltstone unit with units
2-5 in the section south of Dutch
John Mountain (Langenheim and
Peck, 1960: 541, fig. 3) designated

144

Transactions Illinois Academy of Science

as a reference or type section for
the restricted formation concept.
This seems appropriate inasmuch as
the calcareous siltstone is best de¬
veloped in this area and Westgate’s
(Westgate and Knopf, 1932) origi¬
nal description is actually almost
entirely a description of the cal¬
careous siltstone assemblage. Fur¬
ther, it is recommended that the
Chainman Shale concept be restrict¬
ed to the black fissile shale unit with
section V (Langenheim et al., 1960:
149, 151, fig. 1) on the west side of
Ward Mountain south of Ely desig¬
nated as a reference section for the
restricted formational concept. This
seems appropriate inasmuch as Spen¬
cer ’s concept enjoys priority and the
section in the Ely area consists al¬
most entirely of the black fissile
shale unit.

Inasmuch as the uppermost sandy
shale unit is not presently recognized
as a formal stratigraphic unit it is
proposed that the unit be referred
to as the Hamilton Canyon Forma¬
tion. The type section is designated
as the appropriate portion of the
White Pine Shale section shown for
the White Pine District in Figure 2.
This section was measured along a
traverse starting at the White Pine-
•Toana contact on the north side of the
water gap through the Joana Lime¬
stone in the NE sec. 31, T. 17 N.,
R.58 E., Illipah Quadrangle, White
Pine County, Nevada. The traverse
proceeds eastward through cover to
the end of a spur in the Chainman
Shale Formation and thence con¬
tinues roughly along the crest of
the spur to the base of the Ely Lime¬
stone near the Hamilton-Illipah
Road. The base of the Hamilton
Canyon Formation is well-marked

by a soil and vegetation change at
the top of the black, fissile shale to
open grassland with scattered out¬
crops of silty shale and fine-grained
sandstone. The upper contact is
taken at the base of the lowest sig¬
nificant sandstone layer. The thick¬
ness and general character of the
Hamilton Canyon Formation in its
type locality and elsewhere in east¬
ern Nevada are shown on Figure 2.

The Diamond Peak Quartzite and
Scotty AVasli Quartzite concepts are
retained for the dominantly quartz-
itic and conglomeratic units spread¬
ing eastward from the Diamond
Range and centering on the Pioche
District respectively.

Acknowledgments

Special thanks are due Philip
Playford for his energetic defense
of the Eastern Nevada Geological
Association classification in a pro¬
longed running debate on the ques¬
tions discussed herein. J. H. Lan¬
genheim and L. J. Stensaas served
as field assistants and the Depart¬
ment of Paleontology and the Facul¬
ty Research Fund of the University
of California at Berkeley supplied
field expenses for the field studies
upon which this work ultimately
rests. H. R. AVanless kindly read
and criticized the manuscript. Final¬
ly the author begs forgiveness for
any excessive zeal in pressing his
argument and invites reply in the
same spirit.

Literature Cited

Duley, D. H. 1957. Mississippian Stra¬
tigraphy of the Meadow Va’ley and
Arrow Canyon Ranges, Southeastern
Nevada. Unpub. M. A. Thesis, Dept.
Paleontology, University California,
Berkeley, 103 pp., 3 pis., 7 figs.

White Fine Shale

145

Easton, W. H., et al. 1953. Revision
of stratigraphic units in Great Basin.
Bull. Amer. Assoc. Petrol. Geol.,
37(1) : 143-151, 2 figs.

Hague, Arnold. 1870. Geology of the
White Pine Mining District, U.S. Geol.
Exploration Fortieth Parallel (King),
3: 409-421, atlas sheet 14.

- . 1882. Report of Mr. Arnold

Hague, 2nd Ann. Rept. U.S. Geol. Sur¬
vey, 1880-81: 21-35.

- . 1883. Abstract of report on

geology of the Eureka District, Ne¬
vada. 3rd Ann. Rept. U.S. Geol. Sur¬
vey, 1881-82: 237-290, pis. 24-25.

- . 1892. Geology of the Eureka

District, Nevada. U.S. Geol. Survey
Mon. 20, xvii + 499 pp., 8 pis., 9 figs.,
13 Atlas Sheets.

- , and S. F. Emmons. 1877. De¬
scriptive Geology. U.S. Geol. Explora¬
tion Fortieth Parallel (King), xiii +
890 pp., 26 pis.

Humphrey, F. L. 1960. Geology of the
White Pine Mining District. Nevada
Bur. Mines Bull. 57, xiv + 119 pp.,
2 pis., 25 figs.

James. J. W. 1954. Upper Mississip-
pian-Lower Pennsylvanian rocks,
southern Egan Range, Nevada (Ab¬
stract). Bull. Geol. Soc. Amer., 65(12)
(2) : 1268.

Johnson, M. S. and D. E. Hibbard. 1957.
Geology of the Atomic Energy Prov¬
ing Grounds Area, Nevada. U.S. Geol.
Survey Bull. 1021-K: 333-384.

King, Clarence. 1876. Paleozoic subdi¬
visions on the 40th Parallel. Amer.
Jour. Sci., Ill: 475-482.

- . 1878. Systematic Geology.

U.S. Geol. Exploration Fortieth Paral¬
lel (King), xii + 803 pp., 23 pis.

- . 1880. First Annual Report

of the U.S. Geological Survey, 62 pp.

Langenheim, R. L., Jr. 1956a. Lower
Mississippian stratigraphic units in
southern Nevada (Abstract). Bull.
Geol. Soc. Amer., 67 (12) (2) : 1773.

- . 1956b. Mississippian stratig¬
raphy in eastern Nevada (Abstract).
Bull. Geol. Soc. Amer., 67(12) (2):

1714.

- . 1960. Early and Middle Mis¬
sissippian stratigraphy of the Ely
Area. Guidebook Geol. East-central
Nevada, Intermtn. Assoc. Petrol. Geol.,
11th Ann. Field Conf., pp. 72-80, 6 figs.

- . 1961. The Pilot Shale, the

West Range Limestone, and the Devon-
ian-Mississippian boundary in Eastern
Nevada. Trans. Illinois Acad. Sci.,
53(3-4) : 122-131, 3 figs.

- , et al. 1960. Preliminary re¬
port on the geology of the Ely No. 3
Quadrangle, White Pine County, Ne¬
vada. Guidebook Geol. East-central
Nevada, Intermtn. Assoc. Petrol. Geol.,
11th Ann. Field Conf., pp. 148-156,
3 figs.

- and J. H. Peck. 1960. La

Formation de Peers Spring dans le
Pioche District, Nevada, Caliiers
Geol., 56:537-548, 3 figs.

- and Herbert Tischler. 1960.

Mississippian and Devonian paleontol¬
ogy and stratigraphy, Quartz Spring
Area, Inyo County, California. Univ.
Calif. Pub. Geol. Sci., 38 (2) : 89-150,
pis. 14-15, 18 figs.

Lawson, A. C. 1906. The copper de¬
posits of the Robinson Mining Dis¬
trict, Nevada. Univ. Calif. Publ. Bull.
Geol., 4(14) : 287-357.

McAllister, J. F. 1952. Rocks and
structure of the Quartz Spring Area,
Northern Panamint Range, California.
Calif. Div. Mines Spec. Rept., Vol. 25,
38 pp., 3 pis., 13 figs.

Nolan, T. B., et al. 1956. The strati¬
graphic section in the vicinity of Eu¬
reka, Nevada. U. S. Geol. Survey Prof.
Paper 276, 77 pp., 2 pis., 2 figs.

Poole, F. G., et al. 1961. Eleana Forma¬
tion of Nevada Test Site and vicinity,
Nye County, Nevada. U. S. Geol. Sur¬
vey Prof. Paper 424-D : 104-111, 2 figs.,
2 tables.

Ross, W. A. 1956. Mississippian rocks
in eastern California and adjacent
Nevada (Abstract). Bull. Geol. Soc.
Amer., 67(12) (2) : 1729.

Spencer, A. C. 1917. The geology and
ore deposits of Ely, Nevada. U. S. Geol.
Survey Prof. Paper 96, 189 pp., 4 figs.,
15 pis.

Westgate, L. G. and Adolph Knopf.
1932. Geology and ore deposits of the
Pioche District, Nevada. U. S. Geol.
Survey Prof. Paper 171, 79 pp., 8 pis.,
13 figs.

Wilmarth, M. G. 1938. Lexicon of geo¬
logic names of the United States (in¬
cluding Alaska), Part 2, M-Z. U. S.
Geol. Survey Bull., 896 (2 ): 1245-2396.

GROUND WATER GEOLOGY OF THE DEKALB AND

SYCAMORE QUADRANGLES

LOREN T. CALDWELL

Northern Illinois University , DeKalb

The DeKalb and Sycamore quad¬
rangles lie principally in central
DeKalb county. Their east and west
margins extend into the western tier
of Kane county townships and the
eastern tiers of Ogle and Lee coun¬
ties townships. Both Leighton
(1958) and Frye (1960) described
the surface of the study area as al¬
most entirely covered by Tazewell'
glacial drift materials, with the ex¬
ception of the very narrow flood
plains of the streams. The surface
of Tazewell drift is covered almost
entirely by Peorian loess and silts.
Locally the loess and silt covers
Sangamon soil and Illinoian glacial
drift.

This writer (1936) previously de¬
scribed the glacial surfaces. They
consist of two morainal ridges which
cross the area in a northeast-south-
west direction. These ridges belong
to the general Bloomington morainic
system. The western or outer ridge
covers the west half of the DeKalb
quadrangle. This ridge is the north¬
ern limit of the Bloomington mo¬
raine. The inner ridge or Arlington
moraine covers the southeast corner
of the DeKalb quadrangle and much
of the Sycamore quadrangles.

The study area (Figure 1) varies
in topography from flat to slightly
rolling. The streams all flow in
shallow valleys. The relief is due
largely to irregular deposition of
glacial materials super-imposed upon
bedrock relief. The moraines are

characterized by an irregularly
rounded, undulating topography.
These moraines vary in width from
3 to 6 miles.

The southeast part of the Syca¬
more quadrangle has a gradual sur¬
face which slopes toward the Fox
River valley. This area is drained
into the Illinois River. The remain¬
ing surface is drained to the north
through the south branch of the
Kishwaukee river, which empties
into the Rock river south of Rock¬
ford. A small area in the northwest
part of the DeKalb quadrangle
drains to the northwest in the east
branch of Killbuck Creek.

The streams have done very little
cutting into the glacial blanket since
the retreat of the Wisconsin ice.
Stream-cuts seldom occur with
depth greater than 15' to 20'. This
slight amount of post-glacial ero¬
sion has done very little to change
the glacial topography except in the
immediate valleys of the streams.
Narrow alluvial flood plains occur
irregularly along the stream banks
only a few feet above the average
water level.

Procedure

This is a study of the glacial drift,
bed-rock topography, and ground-
water geology in the DeKalb and
Sycamore quadrangles. Data are
secured largely from drilled-well
records located in and near the study

[ 146 ]

Ground Water Geology

147

Fig. 1. — Topographic Map of DeKalb and Sycamore Quadrangles. One inch equals 5 miles;

contour interval is 50 feet. Datum is mean sea level.

148

Transactions Illinois Academy of Science

area. The United States Geological
Survey topographic maps for the
DeKalb and Sycamore quadrangles
are used as a base upon which to
chart the well log data. A large
portion of the well logs are secured
directly from well-drillers’ records.
25 well records are secured from the
files of the Illinois State Geological
Survey well record file library.

All well records studied include
the location of the well, the owner’s
name, the description and thickness
of the rock formations through
which the well penetrated, and
ground water geology data made
available through test pumping of
the wells. These data include the
piezometric levels and pump down
capacity of the water in the aquifer
strata tapped by the well. All well
locations are charted on topographic
maps, which show the following :
altitude of well ’s surface, bedrock
surface levels, and thickness of rock
strata penetrated by each well. A
bedrock surface contour map, and
an areal geology map are made from
these data. Since some wells did
not penetrate the glacial drift to
the bedrock surface, data from the
deepest of these wells are used for
their negative value in determining
the elevation of the bedrock surface.

Glacial Drift Cover

The field data showing the glacial
drift cover came from a total of
1100 well records. More than 930 of
these well logs are located in the
DeKalb and Sycamore quadrangles,
while the remaining 150 well logs
are located marginally outside the
area. There are 542 wells which
penetrate bedrock and 338 wells

which end in the glacial drift.

The initial study of this area was
started in 1936 by this writer. The
present study includes subsequent
well log data. The well log data
collected have made possible the
charting of a bedrock surface map.
These map data show drift thick¬
ness and buried bedrock surface in¬
formation. The average thickness of
the glacial drift is about 150'. Its
greatest thickness is more than 350'.
The minimum thickness is found in
Cortland township. Here its mini¬
mum thickness is 15'. Loess and
silt covers the drift over the entire
area except in present stream chan¬
nel and flood plain areas with thick¬
nesses of 1.5' to 2.7'.

Buried Bedrock Surface

Bretz (1923), Ekblaw (1938),
Poster (1956), Hackett (1960), and
Horbert (1946), have separately re¬
ported that the buried bedrock sur¬
face in northern Illinois consists of
deep valleys separated by high
ridges. Buried bedrock river-chan¬
nels and their directions of slope
were charted on Figure 2. Locally
it was difficult to decide the direc-
tion-of-flow and the elevation of di¬
vides for some minor preglacial
tributaries due to a lack of well log
data. For example, in the south¬
west part of the DeKalb quadrangle
in Willow Creek township, section
14, a well penetrates glacial drift
to an elevation of 552'. Similarly,
levels of 600', 610', 632', and 640'
have been reached without encoun¬
tering bedrock in section 23, 10, 10
and 2, respectively. In addition, in
Shabbona township, section 6, a well
reached a 580' level in drift and in

R. 2 E. R.3E. R.4E. R.5E. R.6E.

149

Ground Water

Geology

—Well Location and Bedrock Surface of DeKalb and Sycamore Quadrangles. Bedrock surface contour interval is 50 ft.
Circles represent wells ending in drift; solid dots represent wells ending in bedrock. One inch equals 5 miles.

150

/ nutsact ton.

<£>

lT)

]. — Areal Geology of DeKalb and Sycamore Quadrangles. One inch equals 5 miles. Vertical hatching represents
St. Peter Sandstone; horizontal hatching represents Maquoketa Shale; right oblique hatching represents
Niagaran Limestone; left oblique hatching represents Galena-Platteville Dolomite.

Ground Water Geology

151

Alto township, section 36, a well
reached a 630' level. Since wells
have entered rock at levels from 700'
to 750' in west central Shabbona,
south central Alto, and southwest
Milan townships, it is postulated
that a short tributary here led into
the Old Rock River instead of en¬
tering* the Shabbona tributary. This
conclusion was made because the
stream valley should logically face
in that direction at that location in
order to be attached to the Old Rock
River Valley. This valley was re¬
ported by Ivnappen and others
(1926) to be located immediately to
the west of this location.

Two wells in Alto township, sec¬
tion 12, enter rock at 640' and in
section 12 at 690'. These wells are
surrounded by higher bedrock levels,
consequently a tributary probably
entered the Old Rock River to the
west.

Well log records indicate two
southeast facing valleys to the west
and northwest of Creston. These
two valleys probably joined near
the north and west edges of the
quadrangle due to the proximity of
the Old Rock River only two or three
miles west of this quadrangle. The
presence of a steep stratigraphic dip
in this part of the quadrangle is a
reason for expecting steep gradient
valleys which face east. Also, a
bedrock ridge extends from south to
north across the DeKalb quadrangle,
west of the Shabbona tributary.

Most of the buried bedrock topog¬
raphy of the DeKalb and Sycamore
quadrangles is determined by two
dendritic streams valleys which flow
south across the two quadrangles
and south west just beyond the
south edge of the area. One of

these streams is located in the De¬
Kalb and the other in the Sycamore
quadrangle. The west and east
tributaries will be called the Shab¬
bona and Hinkley tributaries respec¬
tively.

The Shabbona tributary valley
extends lengthwise through the cen¬
ter of the DeKalb quadrangle, ex¬
tending south out of the quadrangle
near Shabbona. It has a gradient of
3' to 5' per channel mile; and its
lowest level is less than 500' above
sea level. This channel floor lies near
the bottom contact of the Galena-
Platteville with older rocks. Its
drainage divide to the west has ele¬
vations of 750' in the northwest part
and 700' in the southwest part of
the quadrangle. The drainage di¬
vide to the east of this valley at¬
tains elevations of more than 800'
to the northeast and 700' to the
southeast.

The Hinkley bedrock valley is lo¬
cated in the south and central parts
of the Sycamore quadrangle. This
stream has two main branches, one
flowing south from the center of the
quadrangle and the other flowing
southwest along the south edge of
the quadrangle. These two branches
will be called the Maple Park and
the Big Rock branches respectively.
The pattern of the Maple Park and
the Big Rock branches indicates
that they join to the south of the
Sycamore quadrangle. These two
streams have average gradients near
5' per mile in their lower courses ;
their lowest levels are 450' for the
Maple Park branch and 400' for the
Big Rock branch. Their drainage
divides vary in altitude from more
than 800' above sea level in the
north to 650' in the south.

152

Transactions Illinois Academy of Science

The buried bedrock topography of
these two quadrangles was formed
by pre-glacial streams which eroded
rock strata with differing hardnesses.
The pre-glacial Rock river, located
near the west edge of this region,
may have caused the high gradients
of this region’s pre-glacial drainage
channels. The DeKalb quadrangle’s
western half has its bedrock topog¬
raphy eroded into the Galena for¬
mation, resulting in the formation
of its flat divides and abrupt valley
walls. Two erosional remnants of
the soft lower Maquoketa formation
occur in parts of Malta and Milan
townships. The middle horizon of
the Maquoketa formation dominates
the east side of the sloping valley
in the east half of the DeKalb quad¬
rangle. This Maquoketa-covering of
shaly dolomite is very thin. It is
immediately underlain by the top
of the Galena dolomite formation.
Along the extreme east edge of the
DeKalb quadrangle, the more resist¬
ant middle horizon of the Maquoketa
formation, occupies the steep pregla¬
cial valley wall. The western half
of the Sycamore quadrangle con¬
tains the Niagaran limestone which
caps the softer upper portion of the
Maquoketa formation. This resist¬
ant dolomite cap-rock is located prin¬
cipally in Cortland township, and
it extends into the northwest part
of Pierce township. Relatively steep
slopes border this cap-rock. These
slopes extend downward through the
resistant middle Maquoketa. Four
wells (Figure 2) located in the
northeast portion of the DeKalb
quadrangle may have been drilled
into solution cavities or sink holes
which were formed by glacial waters
dissolving the dolomite of the middle

Maquoketa. This resistant rock ho¬
rizon is located at the 750' level in
the north and near the 700' level in
the south portion of the quadrangle.

The eastern half of the Sycamore
quadrangle has bedrock surfaces
capped largely by 25' to 30' of
Niagaran limestone. The streams
which cut through this Niagaran
capping to the south and west
formed steep valleys. One valley
south of Maple Park has a crest to
crest width of 3 miles and a depth
of more than 150'.

Pre-Glacial Drainage
Relations

This area has a buried bedrock
topography which has been reported
by Horbert (1950) to be in accord
with the presence of the pre-glacial
Rock river, located west of this area.
This buried Rock river channel-
floor lies at an elevation near 400',
while the buried stream channels of
the study area have average levels
of less than 500' for both the Shab-
bona and Hinckley tributaries. As¬
suming that they enter the channel
of the buried Rock river some miles
to the south and slightly west, this
difference in level would be expected.
The tendency of these tributary
streams to parallel the pre- glacial
Rock river channel may have re¬
sulted from the influence of strati¬
graphic uplift associated with the
sandwich fault which is known to
occur immediately to the west of the
study area in Ogle County.

Pre-Glacial Areal Geology

A pre-glacial areal geology map
was constructed for the study area.
Figure 3 was constructed from data

Ground Water Geology

153

• rH

bJO

6
i— i

154

Transactions Illinois Academy of Science

supplied largely by 542 bedrock well
records. The Shabbona tributary
stream, flowing from north to south
through the center of the DeKalb
quadrangle, follows closely along the
bedrock surface boundary between
the Galena and the Maquoketa for¬
mations. The Galena formation lies
west and the Maquoketa formation
lies east of this buried outcrop con¬
tact line. Long tongues of the Ga¬
lena dolomite are exposed in the
lower portions of the stream valleys
in the south and the central portions
of the Sycamore quadrangle. Most
of the east half of the DeKalb quad¬
rangle and the west half of the
Sycamore quadrangle have Maquo¬
keta. rock comprising the bedrock
surface ; one exception is the long
narrow cap rock of Niagaran lime¬
stone in Cortland and Pierce town¬
ships. Most of the northeast one-
third of the Sycamore quadrangle
is capped with a very thin remnant
of the Niagaran formation, while
the southeast corner of the quad¬
rangle has the valley slopes com¬
prised of the Maquoketa and Galena
formations.

Since, the pre-glacial surface dis¬
tribution of rock formations are de¬
termined by well records, the ques¬
tion always exists about the extent
of these strata between wells. To
determine this extent, the bedrock
surface contour map and two struc¬
tural sections were employed. The
main areas of rock were fairly easily
determined. Certain localities of¬
fered problems. For example : the
patches of the Maquoketa formation,
shown west of the Shabbona tribu¬
tary stream as capping the upland
flats of the Galena dolomite ridge,
were suggested by well records num¬

bers 8, 9, 34, and 35 in sections 20,
21, 4, and 5, Milan township. The
extent of the formation was postu¬
lated in part by widely scattered
deep well records and by using
known dip slope values and the bed¬
rock surface contour map. The line
of contact between the Maquoketa
and the Galena formations in the
southeast part of the DeKalb quad¬
rangle and the southwest part of
the Sycamore quadrangle is derived
largely from known stratiographic
dip values, and bedrock surface
contours.

The arm of Niagaran limestone
shown in Cortland township has its
presence established by well records
numbers 16, 17, 19, 20, and 46 in
sections 5, 6, 7, and 18, Pierce town¬
ship. The areal extent of the forma¬
tion is determined by widely sepa¬
rated deep well records and upon
calculated structure and bedrock
contours for the area.

One well record number 28 in
section 23, Creston township, indi¬
cates that the floor of the pre-glacial
valley may be in St. Peter sand¬
stone. Due to the steep east dip of
the formation at this location and
the probably steep gradient of the
pre-glacial stream, it is difficult to
determine the bedrock surface ex¬
tent of this formation. However it
has been postulated to lie in a very
narrow strip in the bed of this valley
as well as in the floor of the pre¬
glacial valley located in Willows
Creek township.

Rock Formations

The nature and extent of rock

formations has been determined by

«/

using data from about 60 deep well

Ground Water Geology

155

logs (Table 1). These wells vary
in depth from more than 3,000' to
500'. A few wells, located outside
the area, are included in this group
where deep well records are not
available within the region. A few
well logs have been selected which
show the lithologic nature of the
rock formations in widely separated
areas. A few of the deepest wells
furnish most of the correlating
stratigraphic data. A deep well lo¬
cated in Sycamore township, section
35, SWy4, NW%, SE14 was drilled
to a depth of 3105'. This well pene¬
trates (Cambrian) sediments with a
total thickness of 2080'. These Cam¬
brian strata include : upward ; the
Mt. Simon sandstone formation with
a 1380' thickness, Eau Claire with
420', Ironton-Galesville with 145',
Franconia with 80', and Trempeal¬
eau with 55'. These Cambrian for¬
mations were practically all sand¬
stone with the exception of the
Trempealeau, which was largely
cherty dolomite.

Other deep well records show sev¬
eral Ordovician rock formations.
The Prairie du Chien formation
varies in thickness from 55' to more
than 80'. An unconformity occurs
at the top of this formation. The
St. Peter sandstone rests upon the
eroded surface of the lower forma¬
tions. This formation varies in
thickness from as much as 330' at
Creston to 80' at Elburn. The Glen-
wood formation at the top of the
St. Peter, varies in thickness from
95' in the west part of the area to
55' in the east. The Platteville and
Galena formations are considered
together in this discussion. They
are variable in thickness throughout
this area due to pre-glacial erosion,

with an original average thickness of
345'. In a few places, the Maquo-
keta formation is shown to possess
considerable thickness. This- thick¬
ness in the east portion of the area
is 127'. The lower 35' of the Maf
quoketa formation consists of a soft
black shale, interbedded with dolo¬
mite, the middle 37' is fairly com¬
pact dolomite, while the upper 35'
of the formation is composed of cal¬
careous slialy dolomite.

A very thin capping of Niagaran
limestone (Silurian) is found in the
Sycamore quadrangle. This com¬
pact dark gray dolomite varies in
thickness from 15' in the western
portions to 25' in the eastern por¬
tion of this quadrangle.

Structural Geology and
Sections A-B and C-D

Section A-B crosses the area along
an east-west line at latitude 45 de¬
grees, 55 minutes (Figure 4). Deep
well records at Creston, Malta, De-
Kalb, Cortland, Maple Park, and
Elburn are used in deriving the sec¬
tion. In general, the section was
constructed without difficulty. Some
questions of well log interpretation
arose. For example : well logs at
Malta, Creston, and Elburn did not
recognize the Glenwood formation ;
however, the description of the stra¬
ta in the log led the writer to give
the formation, the thickness indi¬
cated on the section. The lithologic
nature of the Maquoketa is very
dolomitic.

Section C-D is located along a
north-south line across the area on
west longitude 88 degrees 44 min¬
utes (Figure 4). This longitudinal
line crosses the north portion of the
study area. The structural data of

Transactions Illinois Academy of Science

156

section C-D is based upon deep well
logs which are located largely in the
north portion of the section.

Two sections have been made to
show the salient features of struc¬
ture. Section on line A-B crosses
the area on a line through Creston,
Malta, DeKalb, Cortland, Maple
Park, and Elburn. Section on line
C-D crosses the area on a line
through DeKalb to one mile east of
Waterman.

In general, the strata have a dip
slope to the southeast. These strati¬
graphic dip values have been deter¬
mined by the Maquoketa-Galena
strata contact levels. However, dip-
slope values have been measured in
east-west and north-south directions.
The average stratigraphic dip of the
formations from Malta to Elburn,
an east-west distance of about 20
miles, is from 7' to 10' per mile. In
contrast to this value, the strata dip
from Creston to Malta, an east-west
distance of 5 miles, averages 60' per
mile. The direction of dip in this
section is to the east. From north
to south along the west edge of the
DeKalb quadrangle, these strata dip
to the south at 10' per mile over a
distance of more than 17 miles. On
a line parallel to the above, but along
the west edge of the Sycamore quad¬
rangle, the average southerly strati¬
graphic dip is about 4' per mile.
Along the east edge of the Sycamore
quadrangle the average dip is 2.5'
per mile the south. From the north¬
west corner of the area to the south¬
east corner, a distance of above 40
miles, the stratigraphic contact level
(true dip) drops 560'. This dip
direction indicates an average true
dip of 14' per mile. The local true
dip values are greatest along the

northwest edge of the area and less
steep to the southeast. The true
average stratigraphic dip in the
southeast portion of the area is 9'
per mile.

Ground Water and Aquifer
Strata

594 of the wells take water from
the glacial drift. 338 wells take
water from various bedrock aquifers.
A majority of the wells which pump
water from bedrock aquifers are lo¬
cated in the population centers of
the area. The wells which pump
water from the glacial drift forma¬
tion are widely scattered over the
entire area.

Wells ending in the glacial drift
varied in depth from place to place.
In the area from Creston to south
of Lee, many of these glacial drift
wells are 200' to 350' deep. In the
vicinity of Steward and west of Lee,
many of the glacial drift wells are
driven wells penetrating to depths
of 10' to 30'. Such shallow driven
wells also occur northwest of Cres¬
ton to the northwest corner of the
quadrangle. Glacial drift wells in
the vicinity of Waterman are near
130' to 140' in depth, giving satis¬
factory amounts of water. Wells
near Hinkley vary in depth from
60' to 80'. Wells north and east
of Malta and west and north of De¬
Kalb which do not penetrate bed¬
rock, have depths of 80', 120', and
160'. A good share of these wells are
no deeper than 80' with small to
average quantities of water for farm
uses. Wells taking water from the
bedrock formations usually pene¬
trate the overlying strata until one
of the following aquifer strata is

Ground Water Geology

157

Table 1. — Well Log Data for 60 Drilled Well Records Which Penetrate

Bedrock Formations.

Bedrock

Top of

Bottom

Well

Surface

Rock

Township

Location

Elevation

Formation

Well

Willow Creek T.38N., R.2E., (1-38) 38 drift wells,

0 bedrock wells.

Shabbona T.38N., R.3E., (1-64) 62 drift wells,

2 bedrock wells.

Clinton T.38N., R.4E., (1-72) 46 drift wells,

26 bedrock wells.

Squaw Grove T.38N., R.4E., (1-40) 21 drift wells,

19 bedrock wells.

Big Rock T.38N., R.6E., (1-48) 27 drift wells,

21 bedrock wells.

10.

S12, SW, NE, SW

710

640

510

505

shale

limestone

11.

S12, SW, SW, SC

700

630

580

470

shale

limestone

12.

S13, SE, NW, SE

735

635

555

limestone

37a.

S24, NE, NW, NE

735

627

395

limestone

47.

S35, NE, NE, NE

700

640

600

shale

limestone

48.

S34, SW, SE, SE

700

630

470

shale

limestone

Alto T.39N., R.2E., (1-37) 27 drift wells,

10 bedrock wells.

Milan T.39N., R.3E., (1-57) 33 drift wells,

24 bedrock wells.

SI, NW, NE, SW

880

472

430

410

shale

limsetone

40.

S23, SW, SW, SW

885

484

400

limestone

sandstone

Afton T.39N., R.4E., (1-76) 48 drift wells,

28 bedrock wells.

61.

S31, NE, NW, NW 880

665

shale

655

limestone

285

Pierce

T.39N., R.5E., (1-76) 24 drift wells,
52 bedrock wells.

158

Transactions Illinois Academy of Science

Table 1. — Continued.

Bedrock

Top of

Bottom

Well

Surface

Rock

Township

Location

Elevation

Formation

Well

S2, SW, SE, SW

880

712

645

587

shale

limestone

10.

S4, SE, SW, SW

870

710

685

shale

limestone

12.

S5, SE, SE, SE

890

711

625

590

shale

limestone

27.

S12, NW, NE, NE

870

660

620

568

shale

limestone

30.

S13, NW, NW, NE

850

686

624

shale

limestone

33.

S14, SE, SE, NE

810

646

553

shale

limestone

53.

S24, NW, NW, NW

810

664

600

shale

limestone

58.

S27, NE, NE, NE

800

660

625

575

shale

limestone

74.

S35, SE, NW, NW

770

613

455

240

shale

limestone

280

St. Peter
sandstone

Kaneville

T.39N., R.6E., (1-62) 31 drift wells,

31 bedrock

wells.

S5, SE, SE, SE

860

725

685

580

shale

limestone

Glen wood shale

655

Prairie du Chien dolomite

580

36.

S22, NE, SW, SW

800

675

635

limestone

575

615

shale

575

limestone

38.

S26, NE, SW, SW

780

655

640

520

limestone

shale

520

limestone

48.

S30, NW, NE, NW

790

670

550

526

shale

limestone

57.

S35, NW, NW, NW

770

680

668

485

limestone

shale

485

limestone

Dement T.40N., R.2E., (1-38) 24 drift wells,

14 bedrock wells.

24.

S23, NW, SE, SE

910 730

660

325

shale

limestone

St. Peter sandstone

325

25a.

S23, SW, SE, NE

925 675

510

185

shale

limestone

Ground Water Geology

159

Table 1. — Continued.

Bedrock

Top of

Bottom

Well

Surface

Rock

Township

Location

Elevation

Formation

Well

Malta T.40N., R.3E., (1-62) 47 drift wells,

15 bedrock wells.

37a.

S22,

SE, SE, NW 890 655

655

-363

limestone

Galena

Platteville

St. Peter sandstone

460

Trempeleau dolomite

Franconian dolomite

Ironton & Galesville sandstone

-85

Eau claire sandstone

-255

39.

S23,

NW, SW, NE 920 675

420

-300

limestone

St. Peter

sandstone

Trempeleau & Franconian dolomite

100

Ironton & Galesville sandstone

—300

39a.

S23,

SW, NW, NW 900 685

400

—100

limestone

sandstone

—100

limestone

DeKalb T.40N., R.4E., (1-118) 59 drift wells,

59 bedrock wells.

12.

S4, NW, SW, NW

870

700

687

665

shale

limestone

19.

S6, SE, NW, NE

860

680

435

405

limestone

sandstone

37.

S12, SE, SW, NW

880

730

655

shale

limestone

37a.

S12, NE, NE, NW

860

685

640

625

shale

limestone

56a.

S15, SW, SW, SE

855

714

673

—230

limestone

shale

limestone

Glenwood shale

325

St. Peter sandstone

270

Trempeleau dolomite

—62

Franconian

dolomite

—148

Galesville sandstone

—230

61.

S23, SW, NW, NW

880

710

670

—441

shale

limestone

St. Peter sandstone

260

Prairie du Chien dolomite

67.

S23, NW, SE, NW

890

685

660

—400

shale

limestone

St. Peter sandstone

340

Prairie du Chien dolomite

320

68.

S23, NE, SE, SW

890

755

320

—438

limestone

St. Peter

sandstone

160

Transactions Illinois Academy of >$cie?ice

Table 1. — Continued.

Bedrock

Top of

Bottom

Well

Surface

Rock

Township

Location

Elevation

Formation

Well

78.

79a.

93.

S26, NE, NE, NE

S26, NW, SE, NW

S31, SW, SW, SE

Prairie du Chien dolomite

-55

890 710

680

shale

limestone

Glenwood shale

334

St. Peter sandstone

239

Ironton Galesville sandstone

—256

888 718

698

shale

Galena

limestone

Platteville limestone

458

Glenwood shale

333

St. Peter sandstone

263

Prairie du Chien limestone

Trempeleau dolomite

-37

Franconian dolomite

—168

Ironton sandstone

-272

Galesville sandstone

—352

Eau claire sandstone

—432

870 486

460

limestone

Platteville

shelf

limestone

St. Peter sandstone

378

—485

—432

358

Cortland T.40N., R.5E., (1-73) 34 drift wells,

39 bedrock wells.

S4, SW, SW, SE

880

725

690

590

shale

limestone

22.

SI 7, SE, SW, WC

880

607

660

215

shale

limestone

St. Peter sandstone

215

28.

S19, NW, NW, NE

900

716

660

525

shale

limestone

35.

S20, SE, SW, NW

880

715

540

shale

limestone

37.

S20, NE, SW. SE

890

765

740

725

shale

limestone

40.

S20, NW, NE, NE

895

740

690

635

shale

limestone

41.

S20, NE, NW, NW

895

745

207

shale

limestone

51.

S26, NE, SW, NE

860

700

690

shale

limestone

53.

S29, NW, NE, SE

900

765

755

745

shale

limestone

65.

S3 4, NW, NE, SW

870

790

720

shale

limestone

T.40N., R.6E., (1-54) 33 drift wells,

21 bedrock

wells.

36.

S29, SE, SW, NW

870

710

630

610

limestone

Ground Water Geology

161

Table 1. — Concluded.

Bedrock

Top of

Bottom

Well

Surface

Rock

Township

Location

Elevation

Formation

Well

Lynnville T.41N., R.2E., (1-5) 4 drift wells,

1 bedrock well.

South Grove

T.41N., R.3E., (1-18) 11 drift wells,

7 bedrock wells.

S29, NE, NW, NW

870

770

730

shale

limestone

13.

S33, SW, SE, NW

930

760

750

600

shale

limestone

15.

S33, SE, SE, SW

940

770

750

660

shale

limestone

Mayfield

T.41N., R.4E., (1-22) 20 drift wells,

2 bedrock wells.

S28, NW, NE, NE

890

760

745

505

shale

limestone

13.

S32, SE, NE, NE

890

690

600

500

shale

limestone

14.

S33, SW, NE, NE

900

700

620

limestone

21.

S34, SW, NW, NW

870

720

490

limestone

Sycamore

T.41N., R.5E., (1-22) 15 drift wells,

7 bedrock

wells.

S32, NE, NE, SC

850

705

315

1002

limestone

St. Peter

sandstone

19.

S35, SW, NW, SE

910

740

660

—2195

shale

Galena limestone

Glenwood shale

330

St. Peter sandstone

265

Trempeleau dolomite

105

Franconian

dolomite

155

Ironton Gales ville sandstone

—240

Eau Claire sandstone

-380

Granite

-2195

Burlington

T.41N., R.6E., (1-14) 11 drift wells,

3 bedrock wells.

14.

S35, NE, SE, SE

925

715

707

545

shale

limestone

Total Number of Well Log Records
Drift Deepened to Bedrock Logs 50
Drift Wells 542
Rock Wells 338

930

Total

162

Transactions Illinois Academy of Science

reached, the Ordovician dolomites,
the St. Peter sandstones or the Cam¬
brian sandstone aquifers. Most of
the deep city, factory, and railroad
wells take water from these aquifers
horizons. This group of wells num¬
ber about 60 in the total group of
wells studied. 100 of the remaining
338 bedrock wells are located on
farms. Most cases these farm wells
were drilled through the Niagaran
and Maquoketa formations and into
the Galena dolomite 10' to 50' before
being completed. The jointed and
fractured Galena formation fur¬
nishes easy means for the circula¬
tion of good quality artesian water.

In general, the water supply is
excellent from both bedrock and gla¬
cial drift aquifers throughout the
entire study area. In agreement with
earlier reports by Suter and others
(1959), and Templeton (1950), wells
less than 80' deep have proven to be
unsatisfactory, except those shallow
wells which are fed by spring water.
The temperature of the water from
wells which are drilled to 80' depth,
varies from 48 to 52 degrees Fahren¬
heit, while those deeper wells have
water slightly warmer.

Acknowledgments

The author is indebted to North¬
ern University for the use of equip¬
ment and facilities in making and
reporting on this research project.
I wish to express gratitude to the
Illinois State Geological Survey well
log record library for much data
used in this study. I also extend
thanks to Mr. Kenneth Prentice, a
former student in my classes and a
well driller who furnished more than
100 recent bedrock well records

taken largely from rural areas lo¬
cated in these two quadrangles. Spe¬
cific thanks are offered to Mr. John
Ross as the cartographer who made
the final line work on figures one
through four.

Summary

A bedrock surface map and an
areal geology map are formulated
from 930 drilled well logs. Two
structure cross sections are made
from the data given by the deeper
well records, one section extends
across the study area in an east to
west direction and the other from
north to south across the central por¬
tion of the area. Interpretations
are made for the data of the two
sections and from the two maps.
Glacial drift covers all the study area
with thicknesses from 40' to 350'.
The glacial drift aquifer which is
the most productive, occurs at the
base of the drift layer. The buried
bedrock surface has an average dip-
slope to the south, including one
large and two small parallel stream
channels with 5' per mile channel
gradients southward. The areal ge¬
ology consists of Galena-Platteville
surfaces in the DeKalb quadrangle
and Maquoketa shale slopes with
Niagaran dolomite uplands in the
Sycamore quadrangle. The rock stra¬
ta have an average structural dip
to the south-east of about 9' per mile.
Bedrock aquifers occur in the Or¬
dovician (Galena-Platteville) dolo¬
mite, Ordovician (St. Peter) sand¬
stone, Cambrian (Ironton-Gales-
ville) sandstone, Cambrian (Eau
Claire) sandstone, and Cambrian
(Mt. Simon) sandstone. The Iron-
Galesville aquifer is the best water-

Ground Water Geology

163

resource for water wells in the study
area. The study is intended to fur-
nish local detailed geologic data
which could prove useful in further
studies of ground water reserves in
this area.

References Cited

Bretz, J. Harlen. 1923. Geology and
mineral resources of the Kings Quad¬
rangle. Illinois Geol. Survey Bull.
No. 43, pp. 205-304.

Caldwell, Loren T. 1936. A Study of
the Stratigraphy and the PreGlacial
Topography of the DeKalb and Syca¬
more Quadrangles. Unpublished Mas¬
ters Thesis, University of Chicago,

pp. 1-22.

Ekblaw, George E. 1938. Kankakee
Arch in Illinois. Geol. Soc. America
Bull., 49: 1425-1430. (Reprinted as

Illinois Geol. Survey Circ. 40, 1938).
Foster, John W. 1956. Groundwater
geology of Lee and Whiteside Coun¬
ties, Illinois. Illinois Geol. Survey
Rept. Inv. 194, p. 9.

Frye, J. C., and H. B. Willman. 1960.
Classification of the Wisconsinan
Stage in the Lake Michigan Glacial
Lobe. Illinois Geol. Survey, Circ. 285,

pp. 1-16.

Hackett, J. E. 1960. Ground-Water
Geology of Winnebago County, Illinois.
Illinois Geol. Survey, Rept. Inv. 213,
pp. 40-49.

Horbert, Leland. 1946. Preglacial ero¬
sion surfaces in Illinois. Jour. Geolo¬
gy, 54(3) 179-192. (Reprinted as Illi¬
nois Geol. Survey Rept. Inv. 118, 1946,
pp. 183-186.)

Horberg, Leland. 1950. Bedrock topog¬
raphy of Illinois. Illinois Geol. Sur¬
vey Bull. 73, pp. 65-67.

Knappen, R. S. 1926. Geology and Min¬
eral Resources of the Dixon Quad¬
rangle, Illinois Geol. Surv. Bull. 49,
p. 93.

Leighton, M. M. 1958. Important ele¬
ments in the classification of the Wis¬
consin glacial stage. Jour. Geology,
66 (3) p. 288-309.

Suter, Max, Robert E. Bergstrom, H. F.
Smith, Grover H. Emricii, W. C. Wal¬
ton and T. E. Larson. 1959. Prelimi¬
nary report on groundwater resources
of the Chicago region, Illinois. Illi¬
nois Water Survey and Illinois Geol.
Survey Coop. Ground-water Rept. 1,
pp. 48-65.

Templeton, J. S. 1950. The Mt. Simon
sandstone in northern Illinois. Trans.
Ill. Acad. Sci., 43: 151-159. (Reprinted
as Illinois Geol. Survey Circ. 170,
1951.)

ANOMALOUS ALAR PLATE REGULATION IN THE
EARLY CHICK NEURAL TUBE

WESLEY J. BIRGE

University of Minnesota, Morris

In previous studies attention has
been given to numerous problems
concerning regulatory mechanics and
proliferation control mechanisms in
the chick neural tube. In a number
of instances studies were made of
the regulative development, includ¬
ing proliferation patterns, occurring
in response to induced neural tube
defects (Birge and Hillemann, 1953;
Birge, 1959a; 1959b; 1960; 1962).
However, most attention has been
given to the mesencephalic alar
plate system of the early chick brain.

Subsequent to the removal of one
mesencephalic alar plate, cells soon
migrate into the deficient area from
the opposite intact alar plate.
Twelve to twenty-four hours after
the operation, mitotic rate has in¬
creased throughout the intact alar
plate tissue to 30-35% above normal.
Proliferation continues at an acceler¬
ated rate, giving rise by day 7 to
an optic tectum with a cell popula¬
tion approaching that of two normal
mesencephalic alar plates of compa¬
rable development. By day 8 an
overshoot occurs in the growth of
the regulating system, and this con¬
dition is sustained through day 9.
During this period of over-growth
the cell population averages 10%
greater than that of two normal
alar plates. Coincident with this
overshoot, mitotic activity drops be¬
low normal. Accordingly, the cell
population decreases on day 10,
showing a slight undershoot before

leveling off at the normal value (for
two plates) by day 12.

The mechanism controlling pro¬
liferation, presumably a feedback
system, appears quite sensitive to
the size and/or density of the cell
population, compensating for in¬
creases or decreases therein, by con-
comitant shifts in mitotic rate. Ap¬
parently the alar plate system is
normally self -limiting or self-regu¬
lating in the control of proliferation.
The regulative responses to hemi-
lateral alar plate ablation are sum¬
marized in Table 1.

Though it has been well estab¬
lished that the regulatory pattern
noted above is the usual response to
early hemilateral alar plate abla¬
tion, a few exceptions to this pattern
have been noted. They will be treat¬
ed in this paper, as they bear on
the subject of proliferation control.

Methods and Materials

This study concerns three chick
embryos out of 138 which under¬
went hemilateral alar plate abla¬
tions during 28-38 hours of devel¬
opment. After each operation the
embryo was reincubated and they
were subsequently sacrificed at 2 to
20 days of development.

The three specimens in question
were sacrificed at 4-5 days of devel¬
opment. In each instance the brain
lesion was less complete than usual,
leaving a significant amount of dam¬
aged tissue intact. All operations

[164]

Alar Plate Regulation

165

Table 2. — Extent of alar plate regulation occurring subsequent to hemilateral

ablation.

Age of Embryos

No. of
Embryos
Sacrificed

Overall
Range of
Regulative
Growth

Average
Extent of
Regulative
Growth

% size of
Intact Alar
Plate Area
Compared
With 2
Normal Plate
Regions

Days

2 .

8(8)

0-3

3 .

7(4)

-16

4 .

5(4)

18-38

5 .

5(4)

35-52

6 .

7(4)

47-68

7 .

4(4)

60-72

8 .

8(4)

102-120

116

108

9 .

6(4)

108-123

120

110

10 .

10(4)

75-93

11 .

7(0)

12 .

8(4)

76-123

13 ... .

4 (0)

14 .

8(4)

96-109

102

101

15 . . . _

8 (0)

16 .

8(4)

97-104

100

18 . . .

6(4)

97-112

106

103

20 .

5(2)

91-101

The extent of regulative growth is given as the percentage volume of tissue
produced in excess of that normally formed by one alar plate. The number of
embryos used in each age group for volumetric studies is given in parenthesis.
(From Birge, 1959a.)

were performed as previously de¬
scribed (Birge, 1959a). Also, all
other experimental procedures were
maintained as previously noted.

Results

Histological examinations of the
three embyros in question revealed
excessive infoldings of the dorsal
half of the mesencephalon, including
the a]ar plate system. In each case,
the extensive infolding nearly tilled
or occluded the mesocoele, rendering
the mesencephalon essentially solid
in appearance. Also in each instance,
the cellular population of the alar

plates greatly exceeded that nor¬
mally found to occur during the
post-operative period.

Estimates of the population size,
based on methods previously used
(Birge, 1959a,), indicated a five to
eight-fold overproduction of cells, as
compared to normal embryos of cor¬
responding development. As noted
above, cell production normally bears
a distinct relationship to population

size and/or densitv. When modest

' «/

over-production results, cell division
rate normally declines, presumably
in response to such proliferation con¬
trol mechanisms as are discussed
above.

166

Transactions Illinois Academy of Science

However, in the three instances
noted herein, proliferation rate ap¬
parently was not subjected to such
limitations or restrictions. It would
seem that the mechanisms which
normally limit proliferation in such
systems were inoperative in these
three cases, at least in part. This
suggests that in a low percentage
of cases, proliferation control mecha¬
nisms normally operative in the
mesencephalic alar plate system of
the chick may, under certain cir¬
cumstances, break down, at least to
some extent. As a consequence ex¬
cessive over-proliferation may re¬
sult. The three cases in point bear
at least a superficial resemblance to
carcinogenetic systems in this re¬
spect.

Literature Cited

Bikge, W. J., and H. H. Hillemanx.

1953. Metencephalic development and
differentiation following experimental
lesions in the early chick embryo. J.
Exp. Zool., 124:545-570.

Bikge, W. J. 1959a. An analysis of dif¬
ferentiation and regulation in the
mesencephalon of the chick embryo.
American Journal of Anatomy, 104:
431-463.

Bikge, W. J. 1959b. Spontaneous alar
plate hyperplasia in the chick embryo.
Anatomical Record, 135:135-140.

Birge, W. J. 1960. Regulatory poten¬
tiality in the forebrain of the chick
embryo. Anatomical Record, 137:475-
484.

Birge, W. J. 1962a. Wound healing in
the chick neural tube. Trans. Ill.
Acad. Sci., 54:130-134.

Bikge, W. J. 1962b. Proliferation pat¬
terns and control mechanisms in the
mesencephalic alar plate system of the
chick embryo. American Zoologist,
2:63.

Manuscript received December 11, 1961.

ACADEMY BUSINESS

SECRETARY’S REPORT FOR THE YEAR
APRIL 29, 1961 -APRIL 27, 1962

G. ROBERT YOHE, Secretary

Council Meetings

The Council held four meetings dur¬
ing the year, as stipulated in the Con¬
stitution.

First Council Meeting. This was held
iu the Conference room of the Library
of Eastern Illinois University, Charle¬
ston, Illinois, on Saturday morning,
April 29, 1961, with the President, Dr.
Walter B. Welch, presiding. Eleven
persons, including eight members of the
Council and a designated representative
of another, were present.

The Secretary reported that 263 per¬
sons had registered for the 54tli Annual
Meeting; 177 of these were Academy
members. The report on the Section
meetings showed that all twelve had met
on April 28, 1961; that 106 papers were
presented, two were read “by title,” and
that the total of maximum attendance
figures reported by the Section Chairmen
was 409.

Dr. W. W. Grimm was appointed Chair¬
man of the Sustaining Membership Com¬
mittee.

The Educational Films Evaluation
Committee was appointed; see TRANS¬
ACTIONS 54, 207 (1961).

The general idea of holding seminars
for high school teachers on “Recent Ad¬
vances in Biological Science,” for which
a National Science Foundation grant of
$14,090 has been approved, at the four
sites proposed by the Planning Commit¬
tee (see under November 18 Council
Meeting) was approved, and Dr. Norman
D. Levine, Director of the project, was
empowered to appoint an ad hoc com¬
mittee to organize and operate these
seminars. The Council approved the
organization of a second series of simi¬
lar seminars, these to be in the area of
the Earth Sciences, and named Dr. J. C.
Frye to designate someone to prepare a
proposal.

Dr. Wesley J. Birge was reappointed
Editor of the TRANSACTIONS, and the
President and Dr. Birge were empowered
to appoint the Board of Editors.

Mr. Arthur R. Wildhagen was reap¬
pointed as the Academy’s Publicity Ad¬
visor.

The Planning Committee was reap¬
pointed except that Dr. W. J. Birge re¬
placed Dr. P. C. Silva, and a new Com¬
mittee called the Junior Academy Re-
evaluation Committee was appointed;
see TRANSACTIONS 54, 207 (1961).

The Council approved requests sub¬
mitted by two of the Sections that their
Section names be changed. These were :
“Science Education, Psychology, and So¬
cial Science Section” becomes the “Sci¬
ence Teaching Section,” and the “Me¬
teorology Section” becomes the “Me¬
teorology and Climatology Section.”

Approval was given to Dr. Klimstra’s
suggestion that $3,500 of the reserve
fund derived from Patron and Sustain¬
ing member dues and designated for
use in the Junior Academy work should
be invested in short-term interest-bear¬
ing securities.

Dr. Klimstra was instructed to inves¬
tigate the matter of the bonding of the
Treasurer, and to report his findings to
the Council.

Second Council Meeting. This was
held in Room 433 of the Hotel St. Nicho¬
las in Springfield on Saturday, Novem¬
ber 18, 1961. Ten members of the Coun¬
cil were present, as well as fourteen
Committee and Section Chairmen and
some 25 others, mostly Junior Academy
officers who were present only for the
first part of the meeting.

The first hour of the meeting was de¬
voted to a discussion of Junior Academy
reports and problems, after which the
Junior Academy officers withdrew to a
separate meeting room.

President Welch announced and the
Council approved the names of those
who were appointed to the Editorial
Board; see inside front cover of the
TRANSACTIONS Vol. 54, Nos. 3 and 4.

The Secretary reported the following
items which had been transacted by the

[ 167]

168

Transactions Illinois Academy of Science

Council by mail ballot since the last
meeting:

1. Approval of research grants to:
Evan K. Oyakawa, Illinois State Normal
University ($250.00); Sister Mary Mari¬
na, B.V.M., Mundelein College ($200.00) ;
Howard G. Applegate, Southern Illinois
University ($100.00); Boris Musulin,
Southern Illinois University ($200.00);
and William C. Ashby, Southern Illinois
University ($49.00).

2. Approval of mailing out a letter
asking members to support House Bill
1689.

3. Approval for the Illinois Society
of Medical Research to use the Acade¬
my's mailing list to send information
about Senate Bill 719.

4. Approval of a joint meeting co¬
sponsored by the Illinois Archaeological
Survey and the Anthropology Section of
the Academy to be held during the fall
or winter of 1961-62.

In the absence of Dr. Klimstra, the
treasurer’s report was read by Dr. S. E.
Harris and received by the Council.
There were progress reports from sev¬
eral of the Section Chairmen.

Dr. S. E. Harris, Chairman of the
Budget Committee, presented the budget
for 1962. During discussion, several re¬
visions were made; Dr. Harris then
moved and Mr. Bamber seconded the
adoption of the budget of $11,240, ex¬
clusive of National Science Foundation
funds allocated to specific uses. This
was approved.

Dr. Levine announced the following
dates and sites for the high school teach¬
ers’ seminars on “Recent Advances in
Biological Science:”

February 9-10, 1962 —

Lorado Taft Field Campus.

Northern Illinois University

February 23-24, 1962 —

Allerton House,

University of Illinois

March 16-17, 1962—

Augustana College

April 13-14, 1962 —

Little Grassy Lake Campus,

Southern Illinois University

Acting on a recommendation received
from Dr. Lloyd Bertholf, President of
Illinois Wesleyan University, the Coun¬
cil appointed Mr. Matthew Prastein, As¬
sociate Professor of Physics at Illinois
Wesleyan as Second Vice-president in

charge of local arrangements for the
56tli Annual Meeting at Bloomington,
April 25-27, 1963.

The Council approved a recommenda¬
tion that the Junior Academy should
amend its constitution so as to make
possible the charging of entry fees on
projects entered in the District and State
Expositions.

Dr. Kenneth E. Damann’s resignation
as Chairman of the Teacher Training
Committee was accepted, and Dr. Glen
Q. Lefler of Eastern Illinois University
was appointed as Dr. Damann’s suc¬
cessor.

The Council expressed a favorable at¬
titude toward the Junior Engineering
Technical Society (JETS), but referred
the matter of recommending possible
cooperative action to the Junior Acade¬
my Re-evaluation Committee.

The question of bonding the Treasurer
was tabled until the next meeting when
the Treasurer himself could be present
and explain the details.

The following changes in the current
budget were approved :

1. The expenditure of $2,344.00 of the
Academy’s regular funds for Jun¬
ior Academy activities.

2. An increase of $37.25 in the Coun¬
cil Activities item.

3. An increase of $153.69 in the Mis¬
cellaneous category.

The question of adding an item to
future budgets to care for expenses of
the President’s office and of other of¬
ficers of the Council was referred to the
Planning Committee.

The Council recommended that the
Junior Academy enforce all safety regu¬
lations strictly at all District and State
Expositions.

Mr. Carlock spoke of attempts that
were being made to encroach upon the
lands of the Illinois Beach State Park,
near Waukegan, and the Council in¬
structed the Chairman of the Conserva¬
tion Committee to write to the Governor
and to the Director of the Department
of Conservation, voicing the opinions of
his Committee on this matter.

Third Council Meeting. This meeting
was called to order at 9:40 A.M., Feb¬
ruary 17, 1962, in the Illini Union Build¬
ing in Urbana, Illinois, by President
Walter B. Welch. There were present
ten Council members, five Committee
Chairmen, ten Section Chairmen, and
twenty-five other officers, guests, and
officers of the Junior Academy.

Academy Business

16<)

Mr. Hopkins reported that there were
currently 552 Science Clubs affiliated
with the Junior Academy.

After further discussion of the Jun¬
ior Academy matters, the Council and
Academy officers adjourned to reconvene
separately in Room 155, Altgeld Hall.

The Secretary gave a report showing
the downward trend of Academy mem¬
bership, and pointed out the need for
attracting new members.

The Treasurer’s report was presented
and accepted.

The reports made by Second Vice-
president Green and the Section Chair¬
men indicated satisfactory progress in
the making of arrangements for the
55th Annual Meeting at Wheaton Col¬
lege.

The Audit Committee (S. E. Harris,
Chairman, W. M. Lewis, and R. W.
Kelting) presented their report, dated
February 16, 1962. This was accepted.

Reports submitted by Dr. C. L. Kanat-
zar, Delegate to the A.A.A.S. and Miss
Elnore Stoldt, Academy Conference Dele¬
gate, were received.

President Welch announced the ap¬
pointment of three committees: (1)
Nominations: F. J. Kruidenier, Chair¬
man, H. B. Mills, Robert J. Smith, and
Elnore Stoldt; (2) Resolutions: C. L.
Kanatzar, Chairman, G. H. Boewe, Har¬
old M. Kaplan, and Walter M. Scruggs;
(3) Audit: William Lewis, Chairman,
Ralph Kelting, and Miss Ellen Abbott.
The Council approved these appoint¬
ments.

Dr. Klimstra reported on several avail¬
able plans for the bonding of the Treas¬
urer, and said that he would prefer to
have the office bonded. The Council
approved this bonding, and empowered
Dr. Klimstra to make the necessary ar¬
rangements.

The Council authorized the Treasurer
to reimburse Miss Elnore Stoldt, Acade¬
my Conference delegate, for her trans¬
portation and hotel expenses for the
Academy Conference meeting in Denver
in December 1961.

The Secretary was instructed to pre¬
pare and submit to the membership for
action at the 55th Annual Meeting
amendments to the Constitution as giv¬
en below:

The first of these amendments was
prompted by discussion of an obviously
unscientific and unsuitable paper which
was submitted to one of the Sections,
concerning which the Council approved
a motion that “the Council place on
record a statement to the effect that

such papers . . . are not suitable for
presentation at any of the Academy’s
Section Meetings.’’

1. Article III, Section 2, to be amend¬
ed so as to provide for the rejection of
papers submitted for oral presentation
if the Section Chairman deems them un¬
suitable, provided that the member sub¬
mitting such rejected paper shall have
the right to appeal the decision to the
Council.

2. Article V, Section 1, to be amended
so as to provide for eliminating the of¬
fice of Collegiate Section Coordinator
and further to increase the number of
elected Councilors from 3 to 4, to be
elected in rotation, each for a 4-year
term, and that the terms of the present
elected Councilors be extended as neces¬
sary to fit into this rotation.

3. Article X, Section 6, to be amended
so as to provide that the President, as
well as the Secretary and Treasurer,
may be reimbursed for expenses while
attending Council meetings and Annual
meetings.

The Council granted approval for the
Anthropology Section to hold another
cooperative meeting with the Illinois
Archaeological Survey as was done in
December 1961.

Chairman Hopkins of the Junior Acad¬
emy entered to report that the Junior
Academy had approved the plan sug¬
gested by David Reyes-Guerra, State Di¬
rector of the Junior Engineering Techni¬
cal Society that the J.E.T.S. State meet
and the University of Illinois Engineer¬
ing Open House be held on the same
week-end as the Junior Academy State
Exposition in 1963. The Council ap¬
proved this plan on a trial basis.

Questions concerning the suitability
of the name “Transactions” for the
Academy’s journal, the possible estab¬
lishment of emeritus membership in the
Academy, and the matter of creating the
rank of “Fellow” of the Academy as sug¬
gested in Dr. Kanatzar’s report as
A.A.A.S. delegate, were referred to the
Planning Committee.

Fourth Council Meeting. Following
a dinner in the “Twenty-seven Room”
of the College Dining Hall at Wheaton
College, Wheaton, Illinois, on April 26,
1962, President Welch called the meet¬
ing to order at 7:00 P.M. Eleven Coun¬
cil members, nine Section Chairmen,
and ten other officers and Committee
Chairmen were present.

The Secretary’s report included the
following membership statistics as of
April 23, 1962:

170

Transactions Illinois Academy of Science

Life Members . 36

Student Members . 4

Regular Annual Members .... 1242

Sustaining Members . 22

Patron Members . 25

Total . 1329

Excluding Sustaining and Patron
Members, the membership by Sections
is:

Anthropology . 40

Aquatic Biology . 30

Botany . 151

Chemistry . 276

Conservation . 9

Geography . 69

Geology . 108

Meteorology and Climatology. 20

Microbiology . 13

Physics . 100

Science Teaching . 109

Zoology . 256

No Section Designated

Individuals . 64

Clubs . 18

Libraries . 19

Total . 1282

The Secretary’s report also called at¬
tention to the fact that of the 160 au¬
thors and co-authors listed on the Sec¬
tion programs for the 55th Annual Meet¬
ing, 57.5% were not members of the
Academy.

Mr. Hopkins, Chairman of the Junior
Academy, reported that there were 617
Science Clubs registered with the Junior
Academy; that over 500 papers had been
submitted in the competition for the
papers sessions at the State Exposition;
and that the National Science Founda¬
tion had rejected our application for a
grant for the coming year.

Mr. Milton Thompson, reporting for
Dr. Deuel, Librarian, stated that Volume
54, Nos. 3 and 4 had been mailed to the
members on April 19, and that about
$5900 remains in the printing fund.

Dr. Wesley J. Birge, Editor of the
TRANSACTIONS, submitted a report
which was summarized by the Secre¬
tary. It showed that 13 papers had
been published since November 17, 1961
(in Volume 54, Nos. 3 and 4); that 13
manuscripts are currently in press (Vol¬
ume 55, No. 1); and that during this
period, 8 additional manuscripts had
been accepted, 4 rejected, 6 are being
revised, and 7 are being reviewed. Be¬
ginning with Volume 55, four separate
issues per volume will be printed.

Reports of the Section Chairmen and
the Committee Chairmen were largely
routine preliminaries to the functioning
of and reporting to the meetings of
April 27.

Dr. Elaine Bluhm reported that in
view of the fact that the National Sci¬
ence Foundation had not approved the
Academy’s most recent request for a
grant, the Junior Academy Re-evaluation
Committee had met and approved the
following recommendation: “ . . . that
the Junior Academy of Science set up
a system for collecting a state registra¬
tion fee, prorated on the basis of school
size, which would guarantee an income
of $11,000 to $12,000 per year to cover
the cost of the State Chairman’s office
and the expenses of the State Exposition
at the University of Illinois, Urbana.”
After some discussion, this was amended
to provide that each school so registered
should receive the TRANSACTIONS for
the year involved; the recommendation
was then approved by the Council.

Dr. Levine’s report on the four semi¬
nars for high school biology teachers
was summarized by the Secretary; it
indicated that these seminars were
deemed “highly successful,” and that
such seminars could well be continued
in future years.

A question of the matter of Academy
policy in regard to permitting equip¬
ment manufacturers to exhibit their
products at Annual Meetings was re¬
ferred to the Planning Committee.

Professor Matthew Prastein, Second
Vice-president for the 1963 meeting, re¬
ported that the construction of the new
Science building at Illinois Wesleyan
was behind schedule; that it obviously
would not be available for the 1963
meeting, and that it would, therefore,
be necessary for Illinois Wesleyan to
cancel the invitation for the Academy
to hold its 56th Annual Meeting there.

The 55th Annual Meetings

General Meeting. The general session
of Friday morning, April 27, 1962, held
in the Main Auditorium of Pierce Chapel,
Wheaton College, Wheaton, Illinois, was
called to order by President Walter B.
Welch at 10:00 A.M.

Dr. V. R. Edman, President of Wheat¬
on College, gave the address of wel¬
come. This was followed by Dr. Welch’s
Presidential address, “This I Would Like
to Know,” and the address, “An Ameri¬
can Educator in Afghanistan,” by Dr.

Academy Business

171

Elbert H. Hadley, Professor of Chemis¬
try at Southern Illinois University.

Section Meetings. On Friday after¬
noon, April 27, all Sections held meeting
as set forth in the printed program.
One hundred sixteen papers were pre¬
sented, six were read “by title,” and
the sum of the maximum attendance
figures reported by the Section Chair¬
men was 404. The new Chairmen named
by the Sections are listed elsewhere in
the TRANSACTIONS, under “Officers for
1962-63.”

THE ANNUAL BUSINESS MEETING

The Business Meeting was called to
order at 5:00 P.M., April 27, 1962, in the
Main Auditorium of Pierce Chapel,
Wheaton College, Wheaton, Illinois, by
President Walter B. Welch. About 55
members were present.

President Welch called attention to
the fact that the minutes of the 54th
Annual Business Meeting had been pub¬
lished in the TRANSACTIONS, Volume
54, pp. 196-208. The Secretary said, “Mr.
President, I move the acceptance of the
report of the 54th Annual Meeting as
published, and also of the summary of
the year’s activities as given in the
President’s Annual letter.” Dr. Evers
seconded and the motion carried.

Reports of Officers

President Welch reported that Illinois
Wesleyan University had found it neces¬
sary to withdraw the invitation for the
Academy to meet there in 1963, and that
the Academy would welcome invitations
for both 1963 and 1964.

Secretary Yohe reported items which
are recorded in the foregoing reports of
Council meetings.

Treasurer Klimstra’s report is pub¬
lished herewith. This report was ac¬
cepted by passage of a motion made
by Dr. Klimstra and seconded by Dr.
Evers.

Librarian Deuel’s report, presented by
Milton Thompson, was that recorded
under the April 26 Council meeting.

Reports of Standing Committees

Dr. Kaplan (Animal Experimentation
in Research), Dr. Klimstra (Conserva¬
tion), and Dr. Lefler (Teacher Training)
referred to resolutions to be presented
later in the meeting.

President Welch summarized the budg¬
et for 1962, which had been approved

by the Council at the November IS, 1961,
meeting.

The following report submitted by Dr.
Van Lente, Chairman of the Research
Grants Committee, was read by the
Secretary:

The Research Grants Committee rec¬
ommends that the following grants be

awarded :

Wm. C. Ashby, Southern

Illinois University . $ 75.00

Sister M. Paulita Springer,

Rosary College . 160.00

Benedict J. Jaskoski, Loyola

University . 50.00

J. Alan Holman, Illinois

State Normal University. 279.00

George Seketa, Southern

Illinois University . 150.00

Aristotel J. Pappelis, South¬
ern Illinois University... 150.00

Total . $ 864.00

Available

from the A.A.A.S . $ 864.00

A motion for the approval of these grants
was made by Dr. Yohe, seconded by Mr.
Bamber, and passed.

The report of the Science Talent
Search Committee was summarized by
the Secretary. It contained the names
of 2 National winners, 25 National and
11 State Honorable mentions in the
Westinghouse Science Talent Search,
and named Gerald Ralph Smith of Green¬
ville High School, Greenville, as winner
of the Frank H. Reed Memorial Award.

Dr. Grimm, reporting for the Sustain¬
ing Membership Committee, stated that
the income from sustaining and patron
memberships since November, 1961, was
approximately $4,000.00, which is about
$1,000.00 less than for the corresponding
period a year ago.

Reports of Special Committees

The report of the Audit Committee
was received by the Council at the Feb¬
ruary 17 meeting; the report of the
Educational Films Evaluation Commit¬
tee will be submitted later to the
TRANSACTIONS for publication.

The report of the Junior Academy Re-
evaluation Committee, which had been
amended and then approved by the Coun¬
cil on April 26 (see above), was ap¬
proved in this final form by the Academy.

The following Constitutional amend¬
ments, which had been sent to all mem¬
bers on March 20, 1962, were read and

172

Transactions Illinois Academy of Science

explained by the Secretary, and voted
upon separately. Deletions are in paren¬
theses; additions are italicized.

Article III, Section 2.

Regular individual members in
good standing shall have the privi¬
lege of voting at the annual meet¬
ing, holding office, offering papers
for presentation at meetings sub¬
ject to the approval of the ap¬
propriate Section Chairman and
with right of appeal to the Coun¬
cil , having (such) papers pub¬
lished in the Transactions if ac¬
cepted by the Board of Editors,
and receiving one copy of the cur¬
rent Transactions of the Academy.
No member in arrears shall re¬
ceive the Transactions for any
year for which he is or remains
in arrears.

Dr. Frye moved and Dr. Klimstra sec¬
onded that this amendment be adopted.
Carried.

Article V, Section 1.

The Council shall consist of the
President, First Vice-president,
Second Vice-president, Secretary,
Treasurer, Librarian, General
Chairman of the Junior Academy,
(Coordinator of the Collegiate
Section), the immediate past Pres¬
ident, the immediate past Secre¬
tary, the immediate past Treas¬
urer, each for a term of one year,
and (three) four Councilors-at-
large. These last shall be elected
for (three-) four- year terms, only
one being elected each year, ex¬
cept that the first year this pro¬
vision is put into effect (one Coun¬
cilor-at-large shall be elected for
a one-year term, one for a two-
year term, and one for a three-
year term.) the terms of the
three incumbent Councilors-at-
large shall each be extended one
year and a fourth Councilor-at-
large shall be elected for a four-
year term.

Dr. Yohe moved and Mr. Bamber sec¬
onded that this amendment be adopted.
Carried.

Article X, Section 6

The President , the Secretary and
the Treasurer shall be reimbursed
for their expenses while attending
Council meetings and annual meet¬
ings . . .

Dr. Evers moved and Dr. Grimm sec¬
onded the approval of this amendment.

After discussion and explanations by
Dr. Bennett, Ekblaw, Grimm, Klistra.
and Green, the motion carried.

The Resolutions Committee (C. L.
Kanatzar, Chairman, G. H. Boewe, H. M.
Kaplan, and W. M. Scruggs) submitted
the following resolutions read by Dr.
Kanatzar; all were approved by the
Academy:

1. APPRECIATION TO HOST
Whereas the Administration, Faculty,

and Staff of Wheaton College have pro¬
vided the arrangements for this Fifty-
fifth Annual Meeting of the Illinois State
Academy of Science,

Be it resolved that the Academy ex¬
press its thanks to all who have served
in any capacity in promoting the in¬
terests and activities of the members
during this meeting, and especially to
Dr. V. Raymond Edman, President of
the College,

Dr. Frank 0. Green, Second Vice-Pres¬
ident of the Academy, in charge of
local arrangements,
the Committee on Local Arrangements,
including Doctors Leedy, Brand,
Kraakevik, Wright, Boardman. and
Mack, and Mr. Haddock, and
the Divisions of Food Service, Build¬
ings and Grounds, and Publicity.
Be it further resolved that the Secre¬
tary be directed to send copies of this
resolution to those specifically named.

2. APPRECIATION OF SERVICE

Whereas

the Departments of Biology, Chemis¬
try, and Geology, of Wheaton Col¬
lege,

Mr. Swink, of the Morton Arboretum,
and

the Illinois State Geological Survey
have accepted the responsibility for
sponsoring the Field Trips to the Ar-
gonne National Laboratory, the Morton
Arboretum, and the surrounding geo¬
logical areas, as a significant portion of
the program for the Fifty-fifth Annual
Meeting of the Academy.

Be it resolved that the Academy ex¬
press its thanks to those responsible for
organizing and conducting these Field
Trips.

Be it further resolved that the Secre¬
tary be directed to send copies of this
resolution to those specifically named.

3. APPRECIATION OF

DISTINGUISHED SERVICE
Whereas Dr. G. R. Yohe, of the Illi¬
nois State Geological Survey, has served

Academ y Busin ess

the Academy in a diligent and produc¬
tive manner as Secretary for the past
three years,

Be it resolved that the members of
the Academy express their appreciation
for his service, and that the Secretary-
elect be directed to send him a letter of
commendation.

4 PROFESSIONAL EDUCATION RE¬
QUIREMENT FOR CERTIFICA¬
TION OF TEACHERS IN THE
SECONDARY SCHOOLS

Whereas a communication from the
Office of the Illinois State Teachers Cer¬
tification Board, dated June 19, 1961,
was received by the Teacher Training
Institutions, County Superintendents of
Schools, and Local School Administra¬
tors, in the state of Illinois, and

Whereas in this letter concerning sec¬
ondary school certification in the state
of Illinois appears the recommendation
that the present requirement of sixteen
(16) semester hours for professional ed¬
ucation courses be increased to eighteen
(18) semester hours by legislative ac¬
tion,

Be it resolved that the Illinois State
Academy of Science oppose this recom¬
mendation.

Be it further resolved that the Acade¬
my authorize its Committee on Teacher
Training to support by any appropriate
means the retention of the present re¬
quirement of sixteen (16) semester
hours of professional education courses
fcr the secondary school certificate.

Be it further resolved that the Secre¬
tary be directed to send a copy of this
resolution to the Illinois State Teachers
Certification Board, Springfield, Illinois;
to the State Superintendent of Public
Instruction, Springfield, Illinois; and to
the academic societies, associations, and
organizations in the state of Illinois, re¬
questing them to support this resolu¬
tion, and to direct a letter of support
to the Illinois State Teachers Certifica¬
tion Board and to the State Superintend¬
ent of Public Instruction, and to any
other persons or groups concerned.

5. NATURE CONSERVANCY

Whereas , it is most apparent that
natural areas are rapidly disappearing
from our State, and that efforts to pre¬
serve many of these for posterity are
frequently thwarted because of financial
difficulties, ignorance, and irresponsi¬
bility, and

173

Whereas the Illinois Chapter of Na¬
ture Conservancy has been partially suc¬
cessful in stemming this trend towards
the loss of natural areas, particularly
along Rocky Branch, northwest of Mar¬
shall in Clark County,

Be it resolved that the Academy rec¬
ognize the efforts of this agency by in¬
structing the Secretary to write to Dr.
Lewis J. Stannard, Chairman of the Illi¬
nois Chapter of Nature Conservancy,
indicating the Academy’s recognition of
the outstanding contributions which Na¬
ture Conservancy is making in our State.

Be it further resolved that the mem¬
bers of the Academy support Nature
Conservancy in its campaign for the
preservation of additional natural areas.

6. PRAIRIE CHICKEN FOUNDATION

Whereas during the past few years the
populations of prairie chickens in Illi¬
nois have shown a drastic decline in
numbers, and

Whereas the recently formed Illinois
Prairie Chicken Foundation has been
making extensive efforts through the
contributions of time, money, and tal¬
ents of many persons, agencies, and or¬
ganizations throughout the State to es¬
tablish refuge areas to insure the preser¬
vation of this native bird, and

Whereas these efforts have resulted
in the purchase of a 77-acre tract near
Bogota in Jasper County,

Be it resolved that the Academy ex¬
tend to the Prairie Chicken Foundation
appreciation of the Foundation’s efforts
to save the prairie chicken, and that the
members of the Academy support the
Foundation’s program.

Be it further resolved that the Secre¬
tary be instructed to send a copy of this
resolution to the Chairman of the Board
of Directors of the Prairie Chicken
Foundation.

7. NATURE PRESERVES

Whereas, a bill prepared for the pur¬
pose of establishing the necessary pro¬
cedure and organization for obtaining
and administering nature preserves was
approved unanimously to both houses of
the 1961 General Assembly, and

Whereas this bill was subsequently
vetoed by the Governor because of his
objection to certain aspects of the form
in which the bill was prepared, and

Whereas Governor Kernel* indicated
his sympathy with the objectives of the
bill, and

174

Transactions Illinois Academy of Science

Whereas it is in the best interests of
the citizens of Illinois to present a re¬
vised nature preserves bill to the 1963
Illinois General Assembly,

Be it resolved that the Academy in¬
dicate its support of the preparation of
a revised bill.

Be it further resolved that the Secre¬
tary write to Mr. George Fell, Chair¬
man of the Citizens Committee for Na¬
ture Conservation, commending him in
his many efforts relating to this bill as
well as to other activities concerned with
the conservation of natural resources in
Illinois.

8. ILLINOIS BEACH STATE PARK

Whereas the Illinois Beach State Park
represents not only one of an insufficient
number of parks in Illinois but also an
unusual and unique natural area, and

Whereas the Illinois Department of
Conservation’s Advisory Board has rec¬
ommended to Governor Kerner that the
demands and interests of the city of
Waukegan to purchase or in some way
obtain 160 acres not be entertained,

Be it resolved that the Academy com¬
mend the Governor and the Director,
William T. Lodge, Illinois Department
of Conservation, on the stand which they
have taken to preserve this unique and
important Illinois park.

Be it further resolved that the Secre¬
tary be directed to send a copy of this
resolution to the Governor and the Di¬
rector.

9. ANIMAL STUDIES IN ILLINOIS
PUBLIC SCHOOLS

Whereas the basic aim of biological
studies that involve animals is to achieve
an understanding of life and to advance
our knowledge of the processes of life,
and

Whereas the ethical and educational
benefits to be gained thereby lead to a
respect for life, and

Whereas biological studies involving
animals are an important means of il¬
lustrating biological principles and in¬
spiring elementary and secondary school
students to consider careers in the bi¬
ological, medical, and veterinary sci¬
ences, and

Whereas there is great need for fu¬
ture scientists in all of the veterinary,
medical, and biological sciences; that
this area is important to the national
defense, and

Whereas the Science Curriculum
Studies of the American Institute of

Biological Sciences contain numerous
projects that involve the use of animals
at the pre-college level; and

Whereas guiding standards for pre-
college animal studies have been ap¬
proved by a number of professional sci¬
entific societies, including the National
Academy of Sciences-National Research
Council, the American Institute of Bi¬
ological Sciences, the National Society
for Medical Research, and the Animal
Care Panel, and

Whereas the Illinois School Code
stating that animals shall not be used
for study or demonstration in Illinois
public schools handicaps proper teach¬
ing of biology,

Be it resolved that the Illinois State
Academy of Science support the princi¬
ple of the need of animal studies in the
public schools of Illinois; that the por¬
tion of the school code forbidding the
use of animals for such studies be re¬
pealed; and that a bill comparable to
House Bill 1689, introduced into the
72nd Illinois General Assembly in 1961,
permitting and regulating the use of
animals in biological studies in the pub¬
lic schools of Illinois, should become
law.

Be it further resolved that the Com¬
mittee on Animal Experimentation in
Research be authorized to cooperate with
other groups to effect a change in the
statute.

10. UNCLAIMED POUND ANIMALS
FOR RESEARCH

Whereas the basic aim of scientific
studies that involve animals is to achieve
an understanding of life, and to advance
our knowledge of the processes of life,
and

Whereas the progress of medical sci¬
ence has contributed greatly to the
health, happiness and longevity of man¬
kind and promises to be equally reward¬
ing in the future, and

Whereas further progress cannot be
achieved without the use of living ani¬
mals for experimentation, and

Whereas animal experimentation is
conducted by humanely motivated in¬
vestigators and with every care to avoid
undue discomfort to the animals in¬
volved, and

Whereas the Illinois State Academy of
Science endorses the use of living dogs
and cats and other animals for research
purposes by responsible investigators in
approved laboratories,

Academy Business 175

Be it resolved that the Illinois State
Academy of Science lend its support to
legislation at the state level that would
make available to approved research and
teaching institutions animals that other¬
wise would be killed in public pounds.

Be it further resolved that the Com¬
mittee on Animal Experimentation in
Research be authorized to cooperate with
other groups to effect such legislation.

11. NECROLOGY

Whereas the Academy has lost by
death within the past year the following
members:

C. C. Burford (September 9, 1961)
John E. Coe (Life Member) (April
11, 1961)

John W. Cralley (February 4, 1961)
L. 0. Gill (July 11, 1961)

Harriet F. Holmes (Life Member)
Edith Putnam Parker (October 10,
1961)

A. J. Throop (1961)

Horatio C. Wood

and

Whereas death has come to two Past
Presidents of the Academy,

Fay-Cooper Cole (September 3,
1961), serving in 1931-32, and
George D. Fuller (Life Member)
(November 22, 1961), serving in
1938-39,

Be it resolved that the members of
the Academy express their sorrow by
rising for a moment of silence.

Dr. Kanatzar moved that all members
present should stand for a moment of
silence. The unanimity of this action
constituted approval of this resolution.

The report of the nominating com¬
mittee (Dr. F. J. Kruidenier, Chairman,

Dr. H. B. Mills, Dr. J. W. Neckers, and
Dr. R. J. Smith) was read by the Secre¬
tary in the absence of Dr. Kruidenier;
in Committee nominations, only the
names of the Chairmen were read. Mr.
Bamber moved and Mr. Austin seconded
that the report be received. Carried.
Dr. Welch called for any nominations
that might be made from the floor.
There being none, Dr. Ekblaw moved
and Dr. Schoffman seconded that the
nominations be closed and that the Sec¬
retary cast a unanimous ballot for this
slate of officers. Carried. The names
of the officers and committees thus
elected, together with others elected by
the Council or the Sections, or appointed
by the President, are published else¬
where in the TRANSACTIONS.

Dr. Ekblaw moved that the members
present stand in recognition of the faith¬
ful efforts made by the officers of the
Academy during the past year. There
were numerous seconds, and those pres¬
ent rose to their feet.

The meeting was adjourned at 6:15
P.M.

Secretary’s note: Complete and de¬
tailed minutes and committee report’s,
of which the above is an abstract, are
on file in the Secretary’s office.

EVENING PROGRAM

The Academy Banquet was held in the
College Dining Hall of Wheaton College
at 6:15 P.M., Friday, April 27, 1962.

The Annual Public Lecture, “The Host
as a Growth Medium for the Parasite,”
by Dr. E. D. Garber, Professor of Micro¬
biology and Plant Pathology at the Uni¬
versity of Chicago, was delivered in the
Main Auditorium of Pierce Chapel at
8:00 P.M.

TREASURER’S ANNUAL REPORT

January 1 - December 31, 1961

Balance Carried Forward, January 1, 1961

$ 7,118.04

Receipts:

Dues

Regular . $ 6,917.00

Sustaining and Patron . 4,692.00

Registration (Charleston) . 111.50

AAAS Research Grants . 1,449.00

Interest, U. S. Bonds . 88.25

Interest, Savings and Loan Assn . 223.50

Return of Loan to Hoffman . 285.32

Refunds from Districts . 340.42

Expenditures :

Council . $ 337.25

Secretary’s Office . 850.50

Treasurer’s Office . 791.22

Editor’s Office . 200.06

Librarian . 319.25

AAAS Research Grants . 1,449.00

Honoraria . . 350,00

Membership Committee . 17.65

Sust. and Patron Membership Comm . .

Planning Committee . .

Publications .

Refunds . 6.00

State Savings and Loan Assn . 4,500.00

Miscellaneous . 303.69

Junior Academy

General . . . $ 2,300.44

Science Talent Search . 501.80

- $ 2,802.24

Grants:

$ 14,106.99

$ 11,926.86

NSF G-12402

Balance as of January 1, 1961 . $ 3,470.19

Refund from Districts . . 148.17

Expenditures:

Salary and Wages . $ 398.28

Speaker Expense . 136.00

Travel . 82.63

Publications . 40.20

Photography . ... 380.09

Research Paper Program . . 200.00

State Chairman Elect . 54.81

Judging Chairman . 89.45

Telephone and Postage . 71.63

Substitute Teacher . 25.00

District Expenses . 2,140.27

NSF G-17016

3,618.36

$ 7,650.00

Receipts:

June 1, 1961

[ 176 ]

Academy Business

177

Expenditures:

Salary and Wages . $ 496.00

Supplies . 1,194.05

Travel . 267.10

Publications . 2,154.87

Telephone and Postage . 320.00

Substitute Teacher . 55.00

Other Expense . 100.00

- $ 4,587.02

Balance as of December 31, 1961 . $ 3,062.98

NSF G-17291
Receipts:

May 15, 1961 . $ 7,045.00

December 27, 1961 . 7,045.00

- $ 14,090.00

Expenditures:

Salary and Wages . $ 17.50

Supplies . 156.10

Travel . 77.60

Publications . 60.90

Other Expense . 17.60

- $ 329.70

Balance as of December 31, 1961 . $ 13,760.30

Permanent Fund

Life Membership

Cash . $ 250.00

Bonds . 1,700.00

- $ 1,950.00

Reserve Funds

U. S. Bonds . $ 1,800.00

State Savings and Loan Assn . 6,000.00

- $ 7,800.00

Frank Reed Memorial Fund

Balance Carried . $ 641.98

Council . 50.00

Interest . 44.45

Honorarium, Yohe . 150.00

- $ 886.43

Cash in Carbondale National Bank as of December 31, 1961 . $ 22,096.37

Balance NSF 17016 . $ 3.062.98

Balance NSF 17291 . 13,760.30

Outstanding Checks . 398.38

Permanent Fund Cash . 250.00

- .$ 17,471.66

Total Uncommitted Cash as of December 31, 1961

$ 4,624.71

ILLINOIS STATE ACADEMY OF SCIENCE
OFFICERS, COMMITTEES, AND SECTION CHAIRMEN

FOR 1962-63

president: John C. Frye, State Geologi¬
cal Survey, Urbana.

first vice-president: Elnore Stoldt, 759
S. Church Street, Jacksonville.

♦second vice-president: Elbert H. Had¬
ley, Southern Illinois University, Car-
bondale.

secretary: Andreas A. Paloumpis, Illi¬
nois State Normal University, Normal.

treasurer: Willard D. Klimstra, South¬
ern Illinois University, Carbondale.

librarian: Thorne Deuel, Illinois State
Museum, Springfield.

GENERAL CHAIRMAN, JUNIOR ACADEMY OF

science: Donald G. Hopkins, Carl

Sandburg High School, Orland Park.

The Council

The Council consists of the above

named officers and the following persons:

IMMEDIATE PAST PRESIDENT: Walter B.

Welch, Southern Illinois University,
Carbondale.

immediate past secretary: G. Robert
Yohe, State Geological Survey, Urbana.

councilors: H. W. Gould (to 1963),

Northern Illinois University, De-
Kalb.

G. H. Boewe (to 1964), State Natural
History Survey, Urbana.

Lyle E. Bamber (to 1965), 101 Burrill
Hall, University of Illinois, Urbana.

Norman D. Levine (to 1966), 143 Vet.
Medicine Building, University of Illi¬
nois, Urbana.

Other Officers

CHAIRMAN-ELECT, JUNIOR ACADEMY '. Wil¬
liam A. Hill, Naperville Community
High School, Naperville.

♦editor: Wesley J. Birge, 318 Natural
History Building, University of Illi¬
nois, Urbana.

♦publicity advisor: Arthur R. Wild-
liagen, 222a Illini Hall, University
of Illinois, Urbana.

♦delegate to the aaas: C. Leplie Kanat-
zar, MacMurray College, Jackson¬
ville.

♦delegate to the aaas academy confer¬
ence: Miss Elnore Stoldt, 759 S.

Church Street, Jacksonville.

Standing Committees

affiliations: George E. Ekblaw, Chair¬
man , State Geological Survey, Ur¬
bana.

Thorne Deuel, State Museum, Spring-
field.

Elbert H. Hadley, Southern Illinois
University, Carbondale.

ANIMAL EXPERIMENTATION IN RESEACH :

Harold Kaplan, Chairman . Southern
Illinois University, Carbondale.

Garwood A. Braun, Highland Park
High School, Highland Park.

N. R. Brewer, 951 East 58th Street,
Chicago.

Robert Sclioffman, Spalding Institute,
Peoria.

F. R. Steggerda, University of Illinois,
Urbana.

ARCHAEOLOGICAL & HISTORICAL SITES: Jo¬
seph Caldwell, Chairman, State Mu¬
seum, Springfield.

William H. Farley, Box 433, Harris¬
burg.

Mary Grant, 805 Randolph, Oak Park,
Illinois.

B. G. Johnson, 1512 Quinton Road, S. E.
Rockford.

John C. McGregor, University of Illi¬
nois, Urbana.

Carroll L. Riley, Southern Illinois Uni¬
versity, Carbondale.

Sol Tax, 1126 East 59th Street, Chi¬
cago.

Daniel A. Throop, Call Printing Com¬
pany, (Third and Broadway) East
St. Louis.

budget: Stanley E. Harris, Chairman,
Southern Illinois University, Car¬
bondale.

Walter A. Brown, Illinois State Normal
University, Normal.

James H. Grosklags, Northern Illinois
University, DeKalb.

J. W. Neckers, Southern Illinois Uni¬
versity, Carbondale.

* Appointed by the President or by the
Council.

[178]

Academy Business

179

conservation: Willard Klimstra, Chair¬
man , Southern Illinois University,
Carbondale.

Stanley A. Changnon, Jr., State Water
Survey, Urbana.

D. H. Ferris, University of Illinois,
Urbana,

John C. Frye, State Geological Survey,
Urbana.

Loring M. Jones, 513 Normal Road,
DeKalb.

Harlow B. Mills, State Natural His¬
tory Survey, Urbana.

Ruben L. Parson, Northern Illinois
University, DeKalb.

Henry Satlier, Western Illinois Uni¬
versity, Macomb.

LEGISLATION AND FINANCE! W. W. Grimm,
Chairman, Bradley University, Pe¬
oria.

John C. Frye, State Geological Survey,
Urbana.

Percival Robertson, The Principia,
Elsali.

Glenn H. Stout, State Water Survey,
Urbana.

Loren P. Woods, Chicago Museum of
Natural History, Chicago.

local conventions: Frank O. Green,
Chairman , Wheaton College,
Wheaton.

Robert R. Brinker, St. James Trade
School, Springfield.

L. R. Hedrick, Illinois Institute of
Technology, Chicago.

A. A. Paloumpis, ex officio, Illinois
State Normal University, Normal.

Robert J. Smith, Eastern Illinois Uni¬
versity, Charleston.

membership: Max R. Matteson, Chair¬
man, University of Illinois, Urbana.

Clyde Anderson, Box 303, Enfield.

J. Bennett, Northern Illinois Univer¬
sity, DeKalb.

Wesley Calef, University of Chicago,
Chicago.

Robert L. Carmin, University of Illi¬
nois, Urbana.

John C. Downey, Southern Illinois
University, Carbondale.

D. Franzen, Illinois Wesleyan Univer¬
sity, Bloomington.

Esther Griffith, Illinois State Normal
University, Normal.

John Harrison, State Geological Sur¬
vey, Urbana.

Ernest L. Karlstrom, Augustana Col¬
lege, Rock Island.

Russell L. Mixter, Wheaton College,
Wheaton.

I. Edgar Odom, State Geological Sur¬
vey, Urbana.

Walter E. Parham, State Geological
Survey, Urbana.

Herbert Priestley, Knox College, Gales¬
burg.

Charles D. Proctor, Loyola University,
Chicago.

Yale S. Sedman, Western Illinois Uni¬
versity, Macomb.

Ben T. Shawver, Monmouth College,
Monmouth.

A. F. Silkett, University of Illinois,
Navy Pier, Chicago.

research grants: Kenneth A. Van

Lente, Chairman, Southern Illinois
University, Carbondale.

Eleanor Dilks, Illinois State Normal
University, Normal.

Ralph J. Miller, Greenville College,
Greenville.

Mark Paulson, Bradley University,
Peoria.

Fr. William J. Slionka, St. Procopius
College, Lisle.

Elnore Stoldt, 759 S. Church Street,
Jacksonville.

H. F. Thut, Eastern Illinois University,
Charleston.

science talent: G. J. Froehlich, Chair¬
man, University of Illinois, Urbana
(Sci. Ed.).

Hal. F. Fruth, 5032 W. Morse, Skokie
(Physics) .

Leland Harris, Knox College, Gales¬
burg (Chemistry).

Charles K. Hunt, Hinsdale (Chemis¬
try).

Donald P. Rogers, University of Illi¬
nois, Urbana (Botany).

John D. Roslansky, University of Illi¬
nois, Urbana (Zoology).

sustaining membership: C. Leplie Ka-
natzar, Chairman, MacMurray Col¬
lege, Jacksonville.

Robert A. Evers, Illinois State Nat.
History Survey, Urbana.

Wilbur W. Grimm, Bradley University,
Peoria.

Milton Thompson, Illinois State Mu¬
seum, Springfield.

Walter B. Welch, Southern Illinois
University, Carbondale.

180

Transactions Illinois Academy of Science

teacher training: Glenn Q. Lefler,
Chairman, Eastern Illinois Universi¬
ty, Charleston.

E. R. Erickson, Augustana College,
Rock Island.

A. Frances Johnson, Rockford College,
Rockford.

Herbert F. Lamp, Chicago Teachers
College, Chicago.

David R. Lauck, Chicago Academy of
Sciences, 2001 N. Clark, Chicago.

R. Maurice Myers, Western Illinois
University, Macomb.

Sister M. Jane Freising. College of
St. Francis, Joliet.

Robert C. Waddell, Eastern Illinois
University, Charleston.

Special Committees
(All appointed by the President
or by the Council)

audit: William M. Lewis, Chairman .

Southern Illinois University, Car-
bondale.

Ellen Abbott, Southern Illinois Uni¬
versity, Carbondale.

Boris Musulin, Southern Illinois Uni¬
versity, Carbondale.

EDUCATIONAL FILMS EVALUATION: MiltOll

D. Thompson, Chairman , Illinois
State Museum, Springfield.

Joseph Caldwell, Illinois State Mu¬
seum, Springfield.

Carleton Condit, Illinois State Mu¬
seum, Springfield.

Melvin O. Foreman, Eastern Illinois
University, Charleston.

Matthew Prastein, Illinois Wesleyan
University, Bloomington.

James M. Sanders, Chicago Teachers
College, Chicago.

Walter M. Scruggs, Eastern Illinois
University, Charleston.

Frank Wittwer, Office of Public In¬
struction, Springfield.

junior academy re-evaluation: (Chair¬
man to be elected by the Commit¬
tee).

Elaine A. Bluhm, 137 Davenport Hall,
University of Illinois, Urbana.

Robert A. Evers, Illinois Natural His¬
tory Survey, Urbana.

C. Leplie Kanatzar, MacMurray Col¬
lege, Jacksonville.

Elnore Stoldt, 759 S. Church Street,
Jacksonville.

Norman D. Levine, 143 Vet. Medicine
Bldg., U. of Illinois, Urbana.

George R. Abraham, Lincoln-Way Com¬
munity High School, New Lenox.

William A. Hill, Naperville Communi¬
ty High School, Naperville.

Donald G. Hopkins, Carl Sandburg
High School, Orland Park.

Sister Mary Ivo, BVM, Immaculata
High School, Chicago 13.

Seichi Konzo, 126 M. E. Bldg., Uni¬
versity of Illinois, Urbana.

Paul M. Wright, Wheaton College,
Wheaton.

nominations: G. R. Yohe, Chairman,

State Geological Survey, Urbana.

William C. Ashby, Southern Illinois
University, Carbondale.

Kenneth H. Harmet, Northern Illinois
University, DeKalb.

F. J. Kruidenier, University of Illinois,
Urbana.

R. Maurice Myers, Western Illinois
University, Macomb.

planning: (Chairman to be elected by

the Committee).

Wesley J. Birge, 318 Natural History
Bldg., U. of Illinois, Urbana.

Kenneth E. Damann, Eastern Illinois
University, Charleston.

John A. Harrison, State Geological
Survey, Urbana.

Joan Hunter, West Senior High
School, Aurora.

F. J. Kruidenier, 322 Natural History
Bldg., U. of Illinois, Urbana.

N. D. Levine, 143 Vet. Medicine Bldg.,
U. of Illinois, Urbana.

Ralph J. Miller, Greenville College,
Greenville.

W. W. Wantland, Illinois Wesleyan
University, Bloomington.

Carl Weatherbee, Millikin University,
Decatur.

resolutions: (For 1962; new commit¬

tee to be appointed at fall Council
meeting) .

C. Leplie Kanatzar, Chairman, Mac¬
Murray College, Jacksonville.

G. H. Boewe, Illinois Natural History
Survey, Urbana.

H. M. Kaplan, Southern Illinois Uni¬
versity, Carbondale.

W. M. Scruggs, Eastern Illinois Uni¬
versity, Charleston.

Academy B usiness

181

Section Chairmen
(Elected by the Sections)

anthropology: Morris Freilich, North¬
ern Illinois University, DeKalb.

aquatic biology: Leo F. Rock, Illinois
Department of Conservation, 705 11th
Street, Sterling.

botany: Dr. Frank A. Crane, University
of Illinois, College of Pharmacy, 833
S. Wood Street, Chicago 12.

chemistry: Dr. Robert E. Van Atta,

Southern Illinois University, Carbon-
dale.

conservation: Dr. Donald T. Ries, Illi¬
nois State Normal University, Normal.

geography: Dr. Stanley Shuman, Illi¬
nois State Normal University, Normal.

geology: Dr. R. L. Langenlieim, Jr.,

University of Illinois, Urbana.

meteorology and climatology : Dr.

James E. Carson, Argonne National
Laboratory, Argonne.

microbiology: Dr. Leslie R. Hedrick,

Illinois Institute of Technology, 3300
S. Federal Street, Chicago.

physics: Dr. Howard H. Claassen,

Wheaton College, Wheaton.

science teaching: Otto Ohmart, Anna-
Jonesboro High School, Anna.

zoology: Dr. Jack Bennett, Northern

Illinois University, DeKalb.

Note: The CHEMISTRY Section named
as Chairman-elect, Dr. Bruce M.
Campbell, MacMurray College,
Jacksonville.

1962-63

PREPARATION OF MANUSCRIPTS FOR
THE TRANSACTIONS

Titles should be brief and informative. The address or institutional
connection of the author appears just below the author’s name. Subtitles
or center headings should be used; ordinarily one uses substitles such as
Introduction, Acknowledgments, Materials, Methods, Results, Discussion,
Summary, and Literature Cited. All papers should have a summary.

No footnotes are to be used.

Doe, John H. 1951. The life cycle of a land snail. Conchol., 26(3):
21-32, 2 tables, 3 figs.

Doe, John H. 1951. Mineralogy of Lower Tertiary deposits. New
York, McGraw-Hill Book Co., iv -(- 396 pp.

Quoted passages, titles, and citations must be checked and rechecked
for accuracy. Citations to particular pages in text are Doe (1908, p. 21)
or (Doe, 1908, p. 21); general citation in text is Doe (1908) or (Doe, 1908).

Photographs should be hard, glossy prints of good contrast. Graphs,
maps and other figures reproduce best when prepared for at least one-half
reduction; lettering, numerals, etc. on all figures in a manuscript should
be worked out to proper size for such reduction. Line widths, letter size
etc. should be uniform from figure to figure within a published paper.
Figures should be drawn on good quality white paper or on drawing
board. Use only India ink. Use a lettering device (Leroy or Wrico) for
numerals and words; do not print “free-hand.”

Authors will receive galley proofs; these should be read carefully and
checked against the original manuscript. Reprints may be ordered at

time galley proofs are returned to the Editor.

Wesley J. Birge,

University of Minnesota, Morris
Morris, Minn.

Xa/idofXincofon

LlbKAKV

NEW YORE

jjOTANlCXL*
GARDE**

Transactions

of the

JUL 2 3 19©

Illinois

State Academy

of Science

Volume 55
Nos. 3 and 4
1962

Springfield, Illinois

TRANSACTIONS of the ILLINOIS STATE ACADEMY of SCIENCE

Editorial Board:

Wesley J. Birge, University of Illinois, Editor and Chairman

Robert S. Bader, University of Illinois

Russell S. Drago, University of Illinois

Francis Kruidenier, University of Illinois

John McGregor, University of Illinois

Wayne J. McIlrath, University of Chicago

Howard C. Roberts, University of Illinois

Theodore Schmudde, Southern Illinois University

Timothy Whitten, Northwestern University

The current Transactions may be obtained by payment of annual dues.

Previous volumes may be obtained by addressing Willard D. Klimstra,
Southern Illinois University, Carbondale.

Exchanges may be arranged by addressing Milton Thompson,

Illinois State Museum, Springfield.

(71558 — 4-63)

TRANSACTIONS

OF THE

ILLINOIS STATE
ACADEMY OF SCIENCE

VOLUME 55 - 1962

Nos. 3 and 4

Illinois State Academy of Science

AFFILIATED WITH THE

Illinois State Museum Division
Springfield, Illinois

PRINTED BY AUTLIORITY OF THE STATE OF ILLINOIS

Otto Kerner, Governor

July 10, 1963

CONTENTS

Presidential Address — This I Would Like to Know.

By Walter B. Welch . 185

Some Coactions of Canada Geese and Small Mammals.

By Lawrence J. Blus and W. D. Klimstra . 191

Dietary Pattern of the Virginia Opossum ( Didelphis marsupialis
virginianus Kerr), Late Summer-Winter, Southern Illinios.

By Walter 0. Stieglitz and W. D. Klimstra . 198

A Late Pleistocene Musk-Ox from East-Central Illinois.

By Edwin C. Galbreath . 209

Current Problems Bearing on the Metabolic Stability of

Deoxyribonucleic Acid (DNAL By Wesley J. Birge . 211

Correlation Between Phenology and Caloric Content in Forest Herbs.

By Barbara J. Kieckhefer . 215

A Re-Examination of Retail Trade in the “Dispersed City”
Southern Ill. By Thomas R. Glennon .

224

Shope’s Fibroma in Illinois Cottontails. By D. H. Ferris, R. D. Lord,

and D. L. Huxsoll . 230

Cerebroid Oolites. By Albert V. Carozzi . 239

Constitution and By-Laws of the Illinois State Academy of Science. . . . 250

[ 184 ]

PRESIDENTIAL ADDRESS

THIS I WOULD LIKE TO KNOW-

W ALTER B. WELCH
Southern Illinois University , Carbondale

At the 54th Annual Meeting of
the Illinois State Academy of Sci¬
ence, the introduction for this ad¬
dress was sounded. Dean Hobart
Heller at Eastern Illinois University,
acting in the absence of President
Quincy Doudna, said that there had
been a number of well known sci¬
entists who, at one time or another
during their careers, had been at
Eastern as students or professors.
The Botanists present thought of
such as E. N. Transeau. He gave
an address, as retiring president of
the Botanical Society of America,
under the title of ‘ ‘ The Fifty Golden
Years of Botany.” It is the fail¬
ing of retiring presidents to make
speeches that act as summaries of a
lifetime of activities in that presi¬
dent’s field of endeavor, or a history
of the organization being addressed.
The history of the Illinois State
Academy of Science is being written.
It is not my desire to anticipate that
work.

Those of us who are in the teach¬
ing of science use this method as an
introduction to a science, or as we
say “To give the historical prospec¬
tive to our subject.” This may be
done so well and completely that the
students get the idea that all the
problems are solved, that all the an¬
swers are given, and they will turn
away from science to other fields

WALTER B. WELCH
President, 1961-1962

where there is still some of the un¬
known to be found. This is an over¬
statement and is not necessarily true
of all sciences nor of any particular
science but it may be true of a par¬
ticular scientist in a particular sci¬
ence.

[ 185 ]

186

Transactions Illinois Academy of Science

Part of the function of an in¬
structor before a class should be to
raise doubts and questions in the
minds of his students as well as to
give answers to questions and to
solve problems. This asking of ques¬
tions is the introduction to research.
It was with this in mind that the
title “This I Would Like To Know
— 77 was chosen. It will be evident
at once that this title is wrong and
it should read “These I Would Like
To Know — ”.

Some of you will remember that
the 1955 Annual Meeting of this
Academy was held at Southern Illi¬
nois University in what was then the
new Life Science Building. Around
that building were mounds of mud
that the landscape architect had left.
The remark was made that the archi¬
tect was attempting to hasten organ¬
ic evolution. When asked “How?”
the answer was “By providing an
environment that might be conducive
for the development of mountain
goats with webbed feet.” A mem¬
ber of the staff of a sister science was
heard to growl, “It would be well for
him to stick to his own field.” I
shall try to heed that admonition.

What is hinted at here is that oth¬
ers of the biological sciences have
advanced much farther in the ex¬
planations of the development of tis¬
sues and organs than have the plant
sciences. These explanations can¬
not be applied to plants without ex¬
tensive modifications. I shall try to
stay with the study of plants.

The questions asked here are not
especially new. Some of them
were asked by the first persons
who examined plants and plant
structures. Some of the questions

were “Where?,” “When?,” “How
many?,” and “How come?” or
‘ ‘ Why ? 7 7 The ‘ ‘ where 7 7 and ‘ ‘ when 7 7
have been rather well investigated
and recorded. The “how many77 is
easily counted, analyzed, and re¬
corded but the “how come77 or
“why77 may still elude us.

To many students of the struc¬
ture of plants, the idea that sim¬
plification or reduction of a struc¬
ture, as an advance in the develop¬
ment of an organ or a plant, is
presented. This idea is illustrated
in reproductive organs, vegetative
organs and in the number of chromo¬
somes in the cell. One example
might be the reduction in the num¬
ber of neck canal cells of the arche-
gonium as seen in such plants as
those of the Bryophyta. The greater
number of cells in the neck canal of
the Mosses means that in this char¬
acter the Mosses are more primitive
than the Liverworts. How did the
Mosses get the longer neck and great¬
er number of neck canal cells? Those
plant scientists who have proposed
the theory that Mosses are more
primitive than the Liverworts have
had trouble with this. The Mosses
could have, by reduction of num¬
bers of cells, produced a neck and
neck canal of the Liverworts but
how did the Mosses get the greater
number in the first place?

A similar situation is found in the
theories that explain the structures
of the stems of plants. We teach
that those plants that have a re¬
duction in the amount of secondary
growth are more advanced in this
characteristic than those plants
which have a greater amount of sec¬
ondary growth. The theories of

Presidential Address

187

simplification and reduction of tis¬
sue or organs are rather well known
but the enlargement of an organ or
tissue, or the theory of accretion, is
seldom expressed.

One of the usual ways in which
to explain the larger structures is
to say that over a great period of
time these structures, organs, or tis¬
sues are built up. It does not take
a student of the plants of the past
very long to discover that the plants
with the larger and more compli¬
cated structures, organs, or tissues
had their origin about the same time
as those with reduced structures.
One attempt to review the informa¬
tion we have on the accretion theory
was made by a member of the Illi¬
nois State Academy of Science, Wil¬
son Stewart (1960). He proposed
that more than one stem was involved
in the development of the stems of
some fossil plants. Several stems
came together to make one stem with
more than one stele. These separate
stems might at first be just that,
separate stems growing very close
together. Later they might “fuse”
making a stem with several steles.
Still later the structure of the stele
may be altered until only the vascu¬
lar bundles are left and a stem ap¬
pears with many vascular bundles
in what is apparently only one stele.
Thus the theories of simplification
and reduction come into play. But
first, the theory of accretion is neces¬
sary to get the larger structures that
can be reduced. Other attempts
should be made to account for the
ideas of simple vs. complex or rather
from the complex to the simple or
reduced in plant structures.

The greater number of chromo¬

somes per nucleus of the cells of
some of the Ferns as compared
with the numbers of chromo¬
somes in the Angiosperms has caused
wonderment. It has been proposed
that there has been a doubling or at
least an addition of chromosomes to
build up a greater number, but here
our timetable seems to be in reverse
again. The plants with the higher
number of chromosomes seem to have
been the ancestors of the plants with
the fewer chromosomes. How can a
plant lose chromosomes without
losing the genetic information neces¬
sary to keep it alive?

Techniques are now in use that
will help us identify the chromo¬
somes of different plants that had a
common ancestry. The chromosome
number, 256, of some of the Ferns
seems to be out of line with the re¬
duced number, 16, of some of the
Angiosperms and they bear little
similarity to the chromosomes of the
Flowering Plants. Do the Ferns
require so much more genetic in¬
formation that they have to have so
many more chromosomes than the
so-called higher plants? Or will the
size of the chromosome make up for
the lower number in the Angio¬
sperms ?

“This I would like to know — ”
How does differentiation take place
in two sister cells such that they
produce different tissues or become
a part of different tissues. The sis¬
ter cells come from the same mother
cell ; presumably they have the same
heritable characteristics. Some con¬
temporary botanists theorize that
there is enough genetic information
in the make-up of the chromosomes
to provide all the possible moro-

.188

Transactions Illinois Academy of Science

phological variations. This may be
an oversimplification that will here
lead ns to obscure some of the de¬
tails of differentiation. Would not
both daughter cells of the same
mother cell have the same genetic
information under normal condi¬
tions? One cell may give rise to
very different cells and tissues than
its sister cell. Then is there some
intrinsic factor which will cause a
difference? We have not been able
to show that they receive different
chemicals from the environment, ex¬
ternally. It does not seem that the
position of the cells can account for
the difference.

Examples of the differences in cell
behavior are pointed up in the work
on Junipers by Margaret Kaeiser
(1960), as reported in the Transac¬
tions of the Illinois State Academy
of Science. Here not only were dif¬
ferent tissues produced from the
derivatives of the same mother cell
but in different amounts.

As Kaeiser points out, some buds
of a leafy shoot produce elongated
stems and other produce the dwarf
stems. Others have shown, in some
plants, that the removal of the apical
bud may cause the dwarf laterals to
assume the characteristics of the bud
that produces an elongated branch.
Is there a reversal of genetic infor¬
mation here or will other intrinsic
or extrinsic factors be responsible
for this change? When will this re¬
versibility cease? Is there, again,
chemical or physical factors involved
that we do not know ?

The first person to work out a
mathematical explanation of the ar¬
rangement of the leaves on a stem
was Leonardo Da Vinci (Welch,

1933) over 400 years ago. He saw
“ where,” “when,” and “how
many.” But “why” he did not
answer. Since Da Vinci’s time oth¬
ers have found the phyllotaxy is a
family characteristic and that it is
transmitted from parent to offspring.
This would indicate that genetic in¬
formation is responsible for the ar¬
rangement of leaves on a stem. Much
work has been done on many plants
of many families with some clear re¬
sults. We know what cells are in¬
volved with the first development of
anything that looks like a leaf— the
leaf primordium. But why these
cells and not the cells next to them?
Why these cells and not those that
are 4 or 5 off to the right or left?
Here again we may have to go be¬
yond our ideas of what is inherited
in order to explain these cell dif¬
ferences.

The work with vegetative buds has
had its counterpart in work with
flower buds. And right away we
could ask the question — Why are
flower buds ever produced? In this
Academy Barbara Palser (1958) has
presented several papers with her
students. Here, it has been pointed
out, there are great differences in
the development of the cells that go
through microsporogenesis and those
that go through a megasporogenesis.
Yet these cells had their inheritance
from the same cells and tissues.
These original tissues were developed
from the same cells that produced
the vegetative organs. How do these
certain cells in the flowers of the
Heath family differ from all other
cells that these will produce micro-
spore mother cells and megaspore
mother cells but no other cells will

Presidential Address

189

produce similar initials? The ge¬
netic information may be the same
in all cells. Then is there some oth¬
er factor necessary? Here again we
ask is there an intrinsic factor in¬
volved? Is this factor in the cell
itself or is it from surrounding cells ?

Some research is being carried out
by a member of this Academy to see
if certain chemicals can be made to
perform as a substitute for an in¬
trinsic factor or, in this case, be an
extrinsic factor. Are the growth
regulating substances the excitants
that will initiate morphogenesis in
plant structures? The point is ini¬
tiate. We know that the auxin re¬
lated growth regulator substances
will enhance the development but
will they initiate the development of
cells, tissues, or organs? This is one
of the things this researcher is at¬
tempting to illustrate. He is using
growth regulating, auxin-like chemi¬
cals on woody cuttings to determine
what happens when adventitious
roots are developed.

There has been some discussion
between plant physiologists and
morphologists as to whether the
growth centers are already present
or developed by the auxin related
chemicals. There is no clear evi¬
dence one way or the other. If
there was sufficient evidence, we
would not have the question. The
work cited may not answer the ques¬
tions but this and other research
may bring us nearer the answer.

‘‘This I would like to know — ”
What initiates cell division? Many
of the cells of the meristems of plants
seem to have all that is necessary to
support mitosis or for that matter —
meiosis. There seems to be enough

food. The structure of the nucleus
and cytoplasm seem to be such that
mitosis and cell division could take
place. Then why don’t these cells
divide more often or sooner than
they do? Some cells of a meristem
have established a rhythm that has
been recorded. Others have not or
at least it is not reported. Is the
rhythm or lack of rhythm determined
by some chemical that needs to be
brought in to stimulate the activi¬
ty? There seems to be enough of
all chemicals necessary including
enough of the ribonucleic acid.
There seems to be more than enough
deoxynibonucleic acid present to
provide for all the chromosome ma¬
terial needed. In fact for some
theory-makers there seems to be too
much DNA in the “resting cells.”

Then why does the cell not divide
sooner or more often? One of the
popular explanations for such ac¬
tivities, control or rhythm, is that
there is an inhibitor present. No
activity can take place until this in¬
hibitor is removed or inactivated.

A member of this Academy is at¬
tempting research on the process of
mitosis to see if there is, indeed, an
inhibitor present and if it is, can it
be inactivated. There is some evi¬
dence that some chemicals are pres¬
ent in the “resting cell” that will
disappear as the cell divides. Are
these inhibiting substances? At¬
tempts are now being made to make
the suspected inhibitor disappear.
If it can be made to disappear or
be inactivated, then it will have to
be determined if cell division is
speeded up.

These are some of the things I
would like to know. You may have

190

Transactions Illinois Academy of Science

some of the answers ; others will come
as the result of research. And this
research will be replaced by still
further research.

The first requirement of this re¬
search should be an idea. The idea
should be dealt with with imagina¬
tion. Too often we are apt to let a
piece of equipment do all the inter¬
pretation of the information gath¬
ered. Too often the measure of the
stature of a man in the scientific com¬
munity is the measure of the number
of dollars he can bring in from out¬
side foundations. Those who assign,
assist, direct or otherwise encourage
research should encourage the use
of the idea equally, if not in greater
measure, than the use of equipment.

Research will go on continually.
Research is as much of a compromise
as is the passing of time and it can’t
be stopped. If the philosopher can
say “today is a compromise between

yesterday and tomorrow,” then we
can paraphrase to say “we teach to¬
day what we did not know yester¬
day, only to have it disproved tomor¬
row. ” Such is our imperfect state
of knowledge. We can hope that
with the results of research we will
teach fewer falsehoods tomorrow
than we teach today.

Literature Cited

Kaeiser, Margaret. 1960. Shoot Apices
in Two Hybrid Junipers. Trans. Ill.
State Acad. Sci., 53:132-140.

Palsar, Barbara. 1958. Studies of Floral
Morphology in the Ericales — IV, Ob¬
servations on Three Members of the
Gaultherieae. Trans. Ill. State Acad.
Sci., 51:24-34.

Stewart, Wilson. 1960. More About the
Origin of Vascular Plants. Plant Sci¬
ence Bui., 6: No. 2, p. 1-5.

Welch, Walter B. 1933. Leonardo Da
Vinci’s Study of Plant Form. The Re¬
sources of Leonardo Da Vinci (An Ab¬
stract) S.I.U. Press.

SOME COACTIONS OF CANADA GEESE AND

SMALL MAMMALS

LAWRENCE J. BLUS AND W. D. KLIMSTRA

Nebraska Game, Forestation and Parks Commission

and

Southern Illinois University i

Where Canada geese concentrate
their feeding and resting activities,
virtually all forage and seeds are
consumed in a relatively few days ;
remaining vegetation is trampled to
ground level or even into the ground.
Because grazing geese tend to avoid
or shy from roads, fence rows, field
edges, or certain physical structures
(Bell, 1957 ; Biehn, 1951 ; Helm,
1951 ; Washington State Game De¬
partment, 1953), the last available
foods are found adjacent to these
sites. On occasion, under extreme
food shortages, these sites may be
utilized by late winter ; however,
such is an exception. It appears that
these feeding and resting activities
of the geese might alter the avail¬
ability of food and cover to small
mammals so as to result in a reduc¬
tion of rodent populations. To
evaluate this and other coactions, a
study of small mammals was con¬
ducted during the winter of 1961
by operating of traplines in wheat,
pasture, and corn fields of the Union
County Refuge. A similar agricul¬
tural area with little or no goose ac¬
tivity, approximately 10 miles north
of the Refuge, was included for com¬
parative studies.

The Union County Wildlife Ref¬
uge comprises 6,201 acres of bottom¬
land 4 miles east of the Mississippi
River and about 1 mile south of

1 Represents a contribution from Project No. 45,
Cooperative Wildlife Research Laboratory.

Ware. This sanctuary was originated
in 1947 to provide a feeding and
loafing area for part of the popula¬
tion of Canada geese, Brant a cana¬
densis, which winters in southern
Illinois. The Refuge, as operated
by the Illinois Department of Con¬
servation, provides wheat, Triticum
aestivum, corn, Zea mays, and pas¬
ture plants such as ladino clover,
Trifolium repens, fescue grasses,
Festuca spp., and orchard grass,
Dactylis glomerata. Johnson grass,
Sorghum halopense, common crab
grass, Dig it aria sanguinalis, and the
panic grasses, Panicum spp. are com¬
mon weeds in corn and pasture fields.

Trapping was initiated January 6
and terminated March 31, 1961.
Hence, populations of small mam¬
mals were not studied prior to the
arrival of geese in September nor
after their departure in March. Mu¬
seum special and regular mouse traps
were utilized in determining the dis¬
tribution and number of small mam¬
mals ; a mixture of peanut butter and
rolled oats was used as bait. In all
areas sampled, intervals of 10 feet
were maintained between stations ;
one trap was placed at each station.
The straight-line method of trapping
was used in Fields la and 3 through
9 (Table 1) ; grid trapping was em¬
ployed in Fields lb and 2. Fields 2,
5, and 7 were located near AVolf
Lake, Illinois; the remaining fields
including 8, which was subjected to

[191 ]

Table 1. Results of Small Mammal Trapping in Agricultural Fields, Union County Wildlife Refuge and Private Property, Illi¬
nois, January-Marck, 1961.

192

Transactions Illinois Academy of Science

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194

Transactions Illinois Academy of Science

little use by geese, were within the
confines of the Refuge.

Results and Discussion

Trapping success during the three
months was low as 11,284 trap nights
yielded 194 captures of small mam¬
mals, a success of 1.7 per cent (Table
1). Four species of mice were rep¬
resented in the captures for the three
types of fields (Table 1). Peromys-
cus maniculatus (81) was the most
abundant and widely distributed,
making use of all but one con¬
trol harvested cornfield. Mus mus-
culus (64) ranked second although
it was not represented in the cap¬
tures in pasture or one field of goose-
harvested corn. Microtus ochrogas-
ter (39), third most abundant, was
not recorded in cornfields or un¬
grazed pasture. P. leucopus (10)
yielded less than 5 per cent of the
catch and was taken in an ungrazed
wheat field.

In Field lb, a lightly grazed strip
of wheat 6 inches in height ex¬
tended from a roadside 30 feet
into the field ; in this area 23 of a
total of 32 captures for the entire
field were recorded. In Field 2,
where wheat was 12 inches in height
throughout, 9 of a total of 40 cap¬
tures were recorded in a strip 30
feet wide, parallel to the edge of the
field. Of 12 captures of Microtus
ochrogaster in Field lb, 11 were
recorded in the lightly-grazed strip
of wheat at the field border; 11 of
15 Mus were captured in that sec¬
tion ; and, of five captures of P. man-
iculatus, one was taken from that
area. P. leucopus was collected only
in Field 2.

In Field 3, a “no trespassing”
sign was located 50 feet from a field

corner bordered by the junction of
two roads. The only Microtus taken
in the field and one of two Mus were
captured in this corner where the
wheat was 8 inches high ; the other
Mus was taken near the corner. All
P. maniculatus were collected out¬
side this area in sections where
grazing was more intense ; two were
taken in areas of bare ground.

In Field la, Microtus were caught
only in the row of traps set 10 feet
from the field border in a lightly
grazed strip of wheat (6 inches high)
which extended from a roadside
20 feet into the field; most of the
Mus and a few P. maniculatus were
taken in this area. Some Mus and
the majority of P. maniculatus were
collected 150 feet from the field edge
where the wheat was 1 inch high.

Microtus seemed to show the most
direct relationship to goose use of
wheat as 27 of 28 specimens were
taken in lightly-grazed areas in
which the wheat was 6 inches or
more in height. Although Mus was
taken in heavily-grazed sections, it
seemed to prefer taller, denser cover,
as 20 of 27 specimens were taken in
lightly-grazed areas. P. maniculatus
was least affected by grazing; 29 of
35 captures were recorded where
goose utilization was most intense.

Although direct evidence was lack¬
ing, some competition probably ex¬
isted among small mammals for the
greater supply of food and cover
present in areas of light grazing.
P. maniculatus , although found
mainly in areas which lacked cover,
were apparently able to compete suc¬
cessfully with other small mammals
where heavy cover and an abundance
of food were available (Table 1,
Field 2). Linduska (1946) found

Canada Geese and Small Mammals

195

that numbers of P. maniculatus were
somewhat higher toward the center
of cultivated fields than nearer field
edges which were adjacent to heavy
fencerow cover. Johnson (1926)
noted that this species was char¬
acteristically an animal of cultivated
fields and was not found in relative¬
ly undisturbed areas which were in¬
habited by Microtus. The possibili¬
ty that at the Union County Refuge
P. maniculatus might have been for¬
aging into denuded areas from liv¬
ing quarters in areas of light grazing
was not substantiated by trapping
data.

The majority of captures of P. leu-
copus in Field 2 (Table 1) was
juveniles which ranged farther into
the field than did the adults. Pres¬
ence of a shrubby area at the field
border probably accounted for the
occurrence of this species. It has
been suggested that P. leucopus enter
open fields due to population pres¬
sure in adjacent woods (Blair, 1940).

All cornfields sampled were picked
mechanically, much grain being lost
during the harvesting process. Waste
grain was abundant in the control
fields as entire ears and individual
grains were observed. On the Ref¬
uge, geese were thorough in gleaning
the fields as no grain was found,
although cobs were plentiful. Only
P. manicidatus and Mus were cap¬
tured, the former being most abun¬
dant in cornfields on the Refuge
and the latter in control fields.

Canada geese have been found to
eat the seeds and, in some cases, the
dried stems of weeds which are found
in cultivated fields (Bell, 1957 ;
Helm, 1951). Thus, geese probably
not only compete with small mam¬
mals for cultivated crops, but also

for most other foods which are ac¬
ceptable to the rodents; conversely,
droppings of geese might have con¬
stituted a source of food if the ro¬
dents were coprophagous.

In cornfields on the control areas,
a large quantity and variety of food
were available to small mammals in¬
cluding corn, weed seeds, and a small
amount of green vegetation. As a
result, the rodents were possibly not
readily attracted by artificial food
used as bait in the traps. This may
have been responsible for the low
catch in control fields and the rela¬
tively higher catch in goose-used
acreages.

Pastures contained many of the
weeds found in cornfields ; and, as in
cornfields, geese would be expected
to be in direct competition with ro¬
dents. Field 9 (Table 1) was per¬
haps the most heavily goose-utilized
area at the Refuge ; cover and food
in particular were scarce. However,
several litters of P. manicidatus and
one litter of Microtus were brought
off in this area during the latter part
of March. This indicated that some
rodents not only were able to survive
a period of apparent stress but were
able to reproduce.

Microtus were taken only at the
border of Field 9 near a ditch and
heavily traveled road where cover
was heavier than in the rest of the
field ; also, some green vegetation
was present. This situation seems
similar to that Martin (1956) re¬
ported where Microtus were appar¬
ently absent in pasture which was
over-grazed by cattle but occupied
a nearby ungrazed area. As in
wheat fields on the Refuge, a strip
at the edge of Field 9 may have
served as a sanctuary for Microtus

196

Transactions Illinois Academy of Science

as little goose activity occurred
there. This restriction of Microtus
to a small area not utilized by geese
indicated the importance of heavy
cover. Although adequate cover is
an important habitat requirement of
this species, Dice (1922) concluded
that sources of food — green vegeta¬
tion and roots or tubers — are more
necessary.

Fields la, lb, and 3 each exceeded
100 acres in size, but suitable habitat
available to Microtus was limited to
a few acres of lightly grazed wheat.
If it can be assumed that Microtus
would occupy the entire area of each
field if grazing were lacking, then
it can also be assumed that Microtus
populations were reduced to a great
extent. There is little evidence to
suggest that P. maniculatus and Mus
populations were greatly reduced in
heavily used fields.

When cattle were removed from
an over-grazed pasture in Kansas,
succession from coarse weeds to suc¬
culent grasses progressed ; and, as an
apparent result, Microtus increased
from a very low population to a
level of abundance in approximately
2 years (Martin, 1956). In fields of
intense goose utilization at the Un¬
ion County Wildlife Refuge, the
ecological situation and time factor
are somewhat different from that de¬
scribed in the above study. The ef¬
fects of geese are temporary as they
are absent during summer and most
of spring when vegetative growth is
greatest. In addition, the food sup¬
ply of all small mammals is renewed
each year by planting and growth of
weed species; thus rodent numbers
probably increase to a normal level
as habitat restrictions terminate

shortly after the geese migrate north
and the vegetation is able to resume
normal growth.

Summary

Species type and population level
of rodents both seemed to show a
relationship to the feeding activities
of the Canada geese. However, this
relationship could not readily be di¬
agnosed as a direct result of goose
utilization and destruction of cover.
Land use and management practices
of the previous summer and fall de¬
termined the kinds and amounts of
cover and food available to the small
rodents irrespective of the presence
of the geese. Hence, differential de¬
grees of effectiveness of goose utili¬
zation in enhancing or discouraging
the occurrence of a given species of
mouse could not be established
with finality. Further, lack of
data on stored food supplies as
noted for several rodents by Wood
(1910), Fisher (1945), and Kenni-
cott (1857), prohibited a complete
understanding of the food complex
for each species captured.

In many cases the presence or
absence of a given species of mouse
showed a relationship to the nature
of habitats immediately adjacent to
areas which were grazed heavily.
When grazed areas were bordered
by roadsides or other natural field
boundaries, the number of captures,
distribution, and species of rodents
seemed to reflect this. Frequently,
due to the avoidance response of
geese to field edges or isolated struc¬
tures in a field, a lightly grazed or
non-grazed spot or strip of wheat
or pasture was present. The pres-

Canada Geese and Small Mammals

197

ence or absence of such areas was
apparent in the trapping- success of
most species of mice.

In wheat fields where Microtus
were restricted to lightly grazed
areas, the depression on population
size was most pronounced. On the
other hand, the majority of captures
of P. maniculatus was recorded in
those areas of wheat which were
heavily grazed by the geese ; Mns
usually preferred areas of light
grazing. Of two pastures and four
cornfields sampled, Microtus were
taken only at the lightly-used edge
of a pasture which provided heavy
cover and green vegetation. In both
pastures and cornfields, a greater
number of captures was recorded
from heavily-utilized fields than in
control areas which were subjected
to little or no use by the geese. The
great quantity and variety of food
available to the small mammals prob¬
ably reduced the attractiveness of
bait in control fields.

In a pasture which showed greater
goose use than any other fields
trapped, P. maniculatus and Micro¬
tus were able to bring off litters in
late March. However, populations
of rodents in all utilized fields were
probably reduced.

Literature Cited

Bell, R. Q. 1957. Pood coactions of Can¬
ada geese, Branta canadensis interior ,
Todd, in southern Illinois. Unpublish¬

ed M. A. thesis, Southern Ill. Univ.
Libr., Carbondale, Ill., 89 pp.

Biehn, E. R. 1951. Crop damage by wild¬
life in California with special empha¬
sis on deer and waterfowl. Calif. Fish
and Game Bui., 5:1-71.

Blair, W. F. 1940. A study of prairie deer
mouse populations in southern Michi¬
gan, Amer. Midi. Nat., 24:273-305.

Dice, L. R. 1922. Some factors affecting
the distribution of the prairie vole,
forest deer mouse and prairie deer
mouse. Ecology, 3:29-47.

Fisher, H. J. 1945. Notes on voles in cen¬
tral Missouri. Jour. Mammal., 26:435-
37.

Helm, L. G. 1951. Effects of Canada geese
on crops and soils in central Missouri.
Unpublished M. A. thesis, Univ. Mo.
Libr., Columbia, Mo., 107 pp.

Johnson, M. S. 1926. Activity and distri¬
bution of certain wild mice in relation
to the biotic community. Jour. Mam¬
mal., 7:245-77.

Kennicott, R. 1857. The quadruped of
Illinois injurious and beneficial to the
farmer. Trans. Ill. State Agr. See,
2:615-84.

Linduska, J. P. 1946. Edge effect as it
applies to small mammals on southern
Michigan farmland. Trans. N. Amer.
Wildl. Conf., 11:200-204.

Martin, E. P. 1956. A population study
of the prairie vole ( Microtus ochrogas-
ter) in northeastern Kansas. Univ.
Kansas Mus. Nat. Hist. Publ., 8 Nel¬
lie.

Washington State Game Department.
1953. Do wild geese injure wheat
crops? Wash. State Game Bui., 5 (3):
12.

Wood, F. E. 1910. Mammals of Cham¬
paign County. Ill. State Lab. Nat. Hist.
Bui., 8:501-613.

DIETARY PATTERN OF THE VIRGINIA OPOSSUM,
DIDELPHIS MARSUPIALIS V1RGIN1ANUS KERR,
LATE SUMMER-WINTER, SOUTHERN ILLINOIS

WALTER O. STIEGLITZ AND W. D. KLIMSTRA

Bureau of Sport Fisheries and Wildlife, Delray Beach , Florida and
Cooperative Wildlife Research, Southern Illinois University.

The opossum ( Didelphis marsu-
pialis virginianus Kerr) is one of
the most common furbearers in Illi¬
nois, and is especially abundant in
the southern one-third of the State.
Because of its wide range of toler¬
ance, omnivorous diet, and high re¬
productive rate, this marsupial
shows general population increases
and continuing widening range, util¬
izing previously unoccupied areas
not only in the United States but
in Canada, as well. Success among
the mammalian fauna is further en¬
hanced by its low-valued pelt, ex¬
panding urban developments, and
decrease of emphasis on it as a food
item or for sport hunting.

This study of the opossum was
undertaken to (1) ascertain what
food items are utilized and the fre¬
quency of occurrence and volume
of each, (2) relate food utilization
with seasons, and (3) evaluate this
mammal as a predator. Although
the literature failed to reveal a de¬
tailed study of the diet of the opos¬
sum in Illinois, several investiga¬
tions of its food habits have been
published for other states (Reynolds,
1945; Hamilton, 1958; Taube, 1947 ;
Wiseman and Hendrickson, 1950;
Sandidge, 1953; Lay, 1942; Wheel¬
er, 1939; and Llewellyn and Uhler,
1952).

Standard laboratory techniques
were utilized in analyzing digestive

tracts and in identifying their con¬
tents. The volume of each item of
food was determined by water dis-
p]acement; frequency of occurrence
was computed on a percentage basis.
For standardization of nomenclature
of food items, Fernald (1950) was
used for plants, Hall and Kelson
(1959) for mammals, American Or¬
nithologists’ Union (1957) for birds,
Conant (1958) for reptiles and am¬
phibians, and Comstock (1947) for
insects.

Mr. Erwin Pearson, formerly Ad¬
junct Research Associate with the
Laboratory, supplied many of the
opossums. This paper represents a
contribution from Projects No. 45
and 52, Cooperative Wildlife Re¬
search Laboratory, Southern Illinois
University, and is abridged from
a thesis submitted in partial fulfill¬
ment of the requirements for the
Master of Arts degree in Zoology.
The data contained herein, and the
preparation of this paper are in no
way associated with the U. S. Bureau
of Sport Fisheries and Wildlife.

Results

The digestive tracts from 131
opossums collected in seven southern
Illinois counties from August, 1958,
through February, 1960, were used
in this investigation. Number of
samples according to month was as
follows: January (11), February

[198]

Food Habits of Opossum

199

Table 1. Major Food Items Occurring in Digestive Tracts of 131 Opossums, Southern
Illinois, 1958-1960.

Food Item

Per Cent
Volume

Per Cent
Frequency of
Occurrence

ANIMAL FOODS .

76.2

100.0

Mammals .

48.7

76.3

Opossum .

16.3

52.7

Cottontail .

14.7

15.3

Prairie Vole .

6.4

5.3

Gray Fox .

2.8

3.1

Short-tailed Shrew .

1.8

4.6

Raccoon . . .

1.7

1.5

N orway Rat .

1.5

0.8

Eastern Mole . . .

1.3

0.8

Deer Mice .

1.0

3.1

Striped Skunk .

0.5

3.1

Pine Vole .

0.5

1.5

Other Mammals . . . . .

0.2

Birds .

14.5

19.1

Domestic Chicken .

7.1

4.6

Grackle .

4.7

2.3

Towhee .

1.3

0.8

Meadowlark .

1 . 1

3.1

Other Birds .

0.3

....

Reptiles .

1.6

14.5

Blue Racer .

0.7

4.6

Other Reptiles .

0.9

....

Amphibians .

3.0

5.3

Frogs .

2.5'

3.8

Toads .

os,;

1.5

Fishes . . . . .

0.1

1.5

Unidentified Scales .

Insects .

6.3

93.1

Scarabaeidae Larvae .

2.0

11.5

Short-horned Grasshoppers . . .

1.5

54.2

Unidentified Lepidoptera Larvae .

0.9

9.2

Other Insects .

1.9

Other Invertebrates .

1.9

Earthworms .

1.3

3.8

Snails .

0.5

31.3

Miscellaneous Invertebrates .

0.1

....

Undetermined Animal Materials .

0.1

2.3

PLANT FOODS .

23 . 8

100.0

Persimmon .

8.1

21.4

Pokeberry .

5.1

25.2

Grapes . . .

1.8

11.5

Tree Leaf Fragments .

1.3

87.0

Corn .

J 1

3.1

Gramineae Leaves and Stems .

0.9

71.0

Plums (Prunus spp.) .

0.9

3.1

Nightshade (Solanum sp.) .

0.9

25.2

Unidentified Fleshy Fruits .

0.9

3.8

Bark and Woody Twigs .

0.8

37.4

Unidentified Seeds and Seed Pods .

0.7

9.2

Other Plant Foods .

1.3

....

200

Transactions Illinois Academy of Science

(16), August (12), September (22),
October (29), November (35), and
December (6). One hundred and
eleven individuals were taken by
trapping, 4 by hunting, and 16 as
DOR,

A total of 75 animal and 66 plant
foods were recorded;1 of these, only
24 animal and 11 plant foods indi¬
vidually constituted 0.5% or more
by volume (Table 1). Animal foods
yielded 76.2% of the total volume
arid plant foods 23.8% ; both groups
appeared in 100% of the tracts.
Four occurrences of miscellaneous
items (string, sacking material, cot¬
ton filter) were recorded ; these were
believed to have been ingested acci¬
dentally.

Animal Foods. Mammals repre¬
sented the most important group
of foods, constituting 48.7% of the
total volume (Table 1). This is in
close agreement with other studies
with the exception of Lay (1942)
who reported mammals yielding 7%
by volume ; however, his sample was
small (16 stomachs) and represented
a single month (September).

The highest ranking food item vol-
umetrically (16.3%) was opossum
(Table 1), suggesting considerable
evidence of cannibalism. Although
remains of opossums appeared in
52.7% of the tracts, only a small
percentage of the occurrences was
considered to be actual food items
as hair and nails were the principle
evidences ; these probably were in¬
gested during preening or trap-
fighting. Four stomachs, which were
filled with remains of opossum, some¬
what distorted the volumetric value

1 For detailed tabular data consult thesis
by senior author on file in Library, South¬
ern Illinois Universty, Carbondale.

for this food item. The majority of
previous records of cannibalism were
among captive animals (Pray, 1921 ;
Seton, 1929 ; Raven, 1929 ; Wheeler,
1939; Wood, 1954). For wild-living
forms Reynolds (1945) and San-
didge (1953) reported that volu-
metrically, opossum contributed
4.9% and 10.9%, respectively.

Cottontail rabbit ( Sylvilagus ftori-
danus) was the second most impor¬
tant food consumed, yielding 14.7%
by volume and occurring in 15.3%
of the tracts. In Kansas (Sandidge,
1953), Missouri (Reynolds, 1945)
and Michigan (Taube, 1947), cotton¬
tail was the primary food.

The prairie vole ( Microtus ochro-
g aster) represented the third rank¬
ing mammalian food item and was
fifth among all foods. It is note¬
worthy that this species constituted
6.4% by volume whereas deer mice
( Peromyscus sp.) comprised only
1.0%.

Large mammals did not contribute
a significant portion of the diet as
by volume gray fox ( Urocyon cine-
reoar gent eus) furnished 2.8%, rac¬
coon ( Procyon lotor) 1.7%, and
striped skunk ( Mephitis mephitis)
0.5%.

Short-tailed shrews ( Blarina
brevicauda) ranked fifth and eastern
moles ( Scalopus aquaticus) eighth
among mammalian foods ; collective¬
ly they represented 3.1% of the total
volume (Table 1). Hamilton (1958)
recorded insectivores from 46 of 461
stomachs. Opossums apparently do
not share the distaste of the red fox
(Vulpes fulva) for this group of
mammals (Murie, 1936 ; Scott and
Klimstra, 1955).

Domestic chicken ( Gallus gallus)

Food Habits of Opossum

comprised 7.1% of the volume of all
foods taken and ranked fourth. It
is doubtful if this represented actual
depredation on poultry flocks; more
likely it reflected availability of car¬
rion.

Among* other birds, the grackle
( Quiscalus sp.) was taken on three
occasions (ranked seventh by vol¬
ume), meadowlark ( Stumella, mag-
na) four times, domestic pigeon
( Columbia livia) twice, and towliee
(Pipiio erythrophthalmus) , junco
( Junco hyemalis ), cardinal ( Rich -
mondena cardinalis ), and Carolina
wren ( Thry othorus ludovicianus )
once each. Tremendous numbers of
grackles winter in this region, and
in localized situations dead grackles
could be one of the most available
foods, especially in roosting areas.

Reptiles occurred in 14.5% of the
tracts, but were significant only dur¬
ing late summer ; only sporadic oc¬
currences were recorded throughout
the remainder of the study period.
The blue racer {Coluber constrictor )
was the most important reptile, con¬
stituting 0.7% of the total volume.

Frogs ( Rana sp.) were taken on
five occasions and constituted 2.5%
by volume. Toads ( Bufo sp.) were
less important, appearing in two di¬
gestive tracts and comprising 0.5%
of the total volume.

Insect remains appeared in 93.1%
of the digestive tracts, but consti¬
tuted only 6.3% of the total volume
(Table 1). Larvae were taken in
considerable numbers, possibly being
more desirable and vulnerable than
adults. Reynolds (1945) reported
that insects comprised 34.2% by vol¬
ume December through May; Wheel¬
er (1939) found that insects fur¬

201

nished 60.3% of the volume for 95
stomachs collected throughout the
year; Sandidge (1953) noted that
insects comprised 42.7% by volume
in 60 digestive tracts taken from
September through Man'll. In con¬
trast, Taube (1947), found 6% of
the diet during Nepfombor-I )ocomber
to be insects while Hamilton (1958)
reported 7.9% of the total volume
of 461 stomachs col leefcd throughout
the year.

The larvae of soarnbaoid beetles
(Sea rabaeidae) , the most important,
insect representative in the current
study, constituted 2.0% of the total
volume and showed a frequency of
11.5%. Although short, -horned grass¬
hoppers ( Locust idao) occurred fre¬
quently (54.2%), they constituted
only 1.5% voliimefrieally. Undeter¬
mined lepidoptcrous larvae had a
frequency of 9.2%; no adults were
recorded.

Plant Foo<ls. IVrsimmou (Pio-
spryos viryiniana) , wh ich was the
most utilized plant, food, ranked
third among all foods, comprising
8.1% of the total volume (Table 1).
I >okeberry ( Phytolacca amcrica na )
occurred in 25.2%; of tin* tracts and
furnished 5.1%; of the total volume,
ranking as tin* sixth most, important,
food. Various other fleshy fruits
constituted 4.9%; of tin* total volume.
Wild grapes (F//wspp.) wen* eaten,
but the volume furnished was rela¬
tively low (1.8%;).

Dried fragments of tree leaves
appeared in 87.0%; of the tracts
while bark and twigs appeared in
37.4%; this probably did not consti¬
tute important, food. The majority
of leaves wore believed ingested acci¬
dentally during feeding activities on

202

Transactions Illinois Academy of Science

Fig. 1 Seasonal trends in the utilization of major food groups by opossums, southern
Illinois, 1958-1960.

ground dwelling insects, snails, etc.
Smith (1941) observed opossums to
place leaves in their mouth during
nest building activities which pos¬
sibly accounts for the appearance of
small quantities of leaves. Also, ani¬
mals under the stress of being steel-
trapped have been observed to in¬
gest leaves, sticks, etc.

Kernels of corn ( Zea mays) com¬
prised 1.1% of the total volume
(Table 2), and had a low frequency
of occurrence (3.1%).

Seasonal Trends. In an effort to
evaluate seasonal aspects of the
opossum’s diet, all digestive tracts
were categorized into late summer
(August and September), fall (Oc-

Food Habits of Opossum

208

tober and November), or winter (De¬
cember, January and February) ; 34,
64 and 33 samples were available,
respectively, for each season.

Marked changes were noted in the
seasonal utilization of plant and ani¬
mal foods (Fig. 1). The volume of
animal materials increased from
52.2% during late summer to 93.4%
in winter; the importance of plant
foods diminished in almost exact
proportion.

Noticeable seasonal trends in the
consumption of the various general
food groups were also noted. The
appearance of mammalian foods in
the diet increased by over 300% be¬
tween late summer and winter. As
the variety of available foods de¬
creased with the advent of severe
weather, opossums possibly resorted
more frequently to mammalian car¬
rion. This is suggested by the marked
increase in utilization of large forms
such as gray fox, striped skunk,
raccoon, woodchuck (Mar mot a mo-
nax) , and opossum as the season pro¬
gressed. Cottontail appeared infre¬
quently during warm weather, but a
major increase in utilization oc¬
curred with the onset of colder tem¬
perature and the beginning of the
hunting season. The decline in con¬
sumption of cottontail during the
winter period might be indicative
of reduced cottontail populations.

Small mammals would tend to be¬
come more vulnerable to predation
as the vegetative cover is reduced
and they are exposed when in quest
of food. Such is suggested by the
utilization of prairie voles which
showed an increase from 2.5% in
the summer to 14.0% during the
winter ; however, predilection and

reduced availability of other foods
are probably reflected as well.

Birds furnished 12.3% of the to¬
tal volume during August and Sep¬
tember, but their use dropped to
virtually zero in the fall. With the
advent of cold weather and probable
increased avian winter mortality,
utilization of birds increased to
29.8% by volume; over four-fifths
of this total was domestic chicken
and grackle.

Reptiles appeared most frequent¬
ly during mild weather with seven
species of snakes and one species of
turtle being recorded. The relative
importance of reptiles decreased as
low temperatures reduced the ac-
tivitv of these cold-blooded forms ;
three occurrences were noted in the
fall sample and two in winter. Am¬
phibians showed a similar pattern
as they comprised 4.0%, 5.4%, and
0.2% of the volume for late sum¬
mer, fall, and winter, respectively.

The volumetric importance of in¬
sects varied between the seasons.
During late summer they furnished
5.9% of the total volume but had
a 97.1% frequency of occurrence;
in fall the volume was 10.9% and
frequency 96.9% and in winter these
measurements were 1.8% and 78.8%,
respectively.

In decreasing order of importance
by volume, short -horned grasshop¬
pers, scarabaeid larvae, adult ground
beetles, and larval ground beetles
were the chief insect items identified
in late summer tracts. Scarabaeid
larvae and short-horned grasshop¬
pers made up over two-thirds of the
total insect volume for fall. The rela¬
tive importance of insects sharply
decreased with the onset of lowered

204

Transactions Illinois Academy of Science

temperatures during winter ; lepi-
dopterous larvae constituted over
80% of all insect materials recorded
for this period. By volume and by
frequency of occurrence, various
Hemiptera were next in importance.
Short-horned grasshoppers, an im¬
portant food item through fall, ap¬
peared in almost one-fourth of the
winter tracts, but constituted only
a trace by volume.

Volumetrically, Gr amine ae was
the leading plant family consumed
during late summer ; corn was the
most important species. The amount
of grasses consumed diminished
markedly in October and November,
but increased noticeably during win¬
ter ; corn was the most prevalent rep¬
resentative.

The utilization of fleshy fruits re¬
flected strongly the time of ripen¬
ing. Grapes were the second most
important plant food in late sum¬
mer ; utilization remained fairly con¬
stant in fall, but declined sharply
during winter. Persimmons were
heavily utilized when present; the
volumetric importance varied from
5.1% in late summer, 15.4% in fall,
to 1.4% during winter. Fruits of
this species normally drop when ripe,
and would thus be most available in
late fall. Although opossums are
adept climbers, it is believed that
they avoid doing so if possible, and
probably prefer to eat persimmons
on the ground ; no green fruits were
noted.

By volume, pokeberry was relative¬
ly important in late summer (6.0%)
and fall (9.8%). Field observations
indicate that the berries shrivel and
dry with the onset of low tempera¬
tures, and they probably lose their

appeal as a food item ; this fruit was
entirely absent in winter.

Discussion

Opossums, like many predatory
animals, are opportunists in the pro¬
curement of food as they probably
use the most readily available, ac¬
ceptable foods. Certain food prefer¬
ences seem to be reflected by the pre¬
ponderance of particular items;
however, selectivity is probably
geared to restrictions imposed by
season, physical ability, familiarity,
and predilection (Scott and Klim-
stra, 1955).

The consumption of carrion was
more pronounced with the advent
of winter and the subsequent re¬
duced availability of favored foods.
However, it was difficult to accu¬
rately determine the percentage of
the diet which resulted from carrion.
Sandidge (1953) suggested that cot¬
tontail, muskrat, and opossum re¬
mains reflected carrion feeding be¬
cause of the association of silphid
beetles (Silphidae) in 12 of 19 oc¬
currences of these mammals. Silphid
beetles appeared in two tracts col¬
lected in September and November
in southern Illinois ; one contained
cottontail remains, but the other only
insect fragments. The conclusion of
Wheeler (1939), who felt that sil¬
phid beetles were taken too promis¬
cuously to be an accurate indicator
of carrion feeding activities, seems
quite logical. In the current study
Muscidae larvae appeared in several
tracts, but were not always associ¬
ated with carrion. Seventy-five fly
larvae were recorded from a single
stomach which contained only insect
and plant materials, suggesting that

Food Habits of Opossum

205

the larvae were taken as a food and
not secondarily with carrion. Ham¬
ilton (1958) postulated that opos¬
sums may consume copious quanti¬
ties of fly larvae when available, dis¬
regarding the carrion on which the
larvae might be feeding.

It seems possible that prairie voles
are more vulnerable to predation or
are more preferred by the opossum
than are deer mice; most trapping
studies of the Cooperative Wildlife
Research Laboratory show the latter
forms much the more numerous. The
home ranges of opossums in south¬
ern Illinois normally encompass
habitats frequented by both of these
mice. The relative abundance of
pine voles ( Pitymys pinetorum),
which is quite low in relation to
prairie voles and deer mice, is re¬
flected in that the former appeared
only twice. Because pine voles and
prairie voles inhabit like habitats
and have similar habits, their vul¬
nerability as prey should be approxi¬
mately the same. The use of large
mammals probably resulted from
carrion as it is doubtful if an opos¬
sum could subdue a healthy, adult
animal of these species. A lead shot
pellet noted in one stomach which
contained gray fox suggested that
hunting injuries might have been
the actual cause of death. A low
frequency of occurrence indicates
that large mammals were chance
items in the diet.

It is noteworthy that meadowlarks
or other ground nesting and roost¬
ing species were not taken more fre¬
quently. This suggests that opos¬
sums may lack the agility to be an
efficient predator on wild birds, even
ground nesters. This is borne out

by the fact that the remains of non¬
domestic birds were recorded from
only 17 digestive tracts.

Results indicated that reptiles
were an acceptable item when avail¬
able, and the species taken reflects
the diverse habitat frequented by
the opossum. Some snakes were prob¬
ably picked up as road kills, but the
opossum is capable of capturing
snakes of considerable size (Lewis,
1929).

Earthworms ( Lumbricus sp.) were
unimportant in this study as they
appeared in only five tracts, yield¬
ing 1.3% by volume. The consump¬
tion of this invertebrate seems to
be a regional variable in regard to
importance in the diet. Dexter
(1951) found that earthworms com¬
prised 15.3% volumetrically of 13
opossums collected in Northeastern
Ohio during the winter. Hamilton
(1958) reported that earthworms
constituted 10.3% of the total vol¬
ume in New York. Michigan studies
indicated that earthworms com¬
prised about 8.0% of the total vol¬
ume (Taube, 1947).

Insects present an example of the
fallacy of relying entirely on per
cent volume as a basis for rating
foods in dietary studies. These in¬
vertebrates ranked low volumetrical¬
ly (6.3%) in the current study
(Table 1), but occurred in 93.1%
of the tracts examined ; this possibly
reflected some predilection. Con¬
versely, the high frequency of oc¬
currence and low volume furnished
by insects may reflect in some cases
chance or promiscuous feeding hab¬
its. This proposition would account
for some of the incongruous values
obtained for frequency of occur-

206

Transactions Illinois Academy of Science

rence and per cent volume in this
study.

The majority of plant foods re¬
corded were available only during
restricted seasons, but were heavily
utilized at such times as they were
present. This availability -consump¬
tion relationship is best illustrated
by the example of pokeberry. Poke-
berry fruits mature early in Septem¬
ber in this area and remain avail¬
able to opossums for an extended
period. The tendency of pokeberry
stems to bend and break makes the
terminal fruiting heads available at
a height conducive to feeding by the
opossum. Dried fruits frequently
adhere to the plant as late as March,
but these were seemingly not as
palatable to the animals as the juicy
berries.

It is surprising that there was not
greater utilization of corn in view
of the abundance of this plant in
southern Illinois. Hartman (1952)
stated that opossums may procure
corn in the manner of raccoons, i.e.,
by pulling up the seedling corn.
Hamilton (1943) suggested that
corn in the milk stage is a favorite
food of opossums, and Reynolds
(1945) reported corn as comprising
7.3% of the opossum diet. Wiseman
and Hendrickson (1950) noted its
occurrence in 46 of 87 scats collected
in Iowa.

Grasses (Gramineae), other than
corn, were recorded in 71% of the
tracts, but in most cases probably
reflect accidental ingestion or food
of secondary origin. Leaves and
stems made up the greater part of
the volume, although 18 species of
seeds were recorded. Some grasses
are probably ingested as the opossum

feeds on certain insects, particularly
grasshoppers which commonly rest
on this type of vegetation. Vege¬
tative material or seeds of grasses
were found in all tracts containing
prairie voles, pine voles, or deer mice,
possibly reflecting secondary inges¬
tion.

Results indicate that predation by
opossums during the period covered
by this study was probably not an
important influence on populations
of game species. Of the major game
animals found in southern Illinois,
only the cottontail formed a signifi¬
cant part of the opossum’s diet.
Bobwhites {Col in us virginianus ) ,
tree squirrels ( Sciurus sp.), white-
tailed deer ( Odocoileus virginianus) ,
and ducks and geese (Anseriiformes)
were not recorded from digestive
tracts examined.

Probably a portion of the cotton¬
tail remains reflected actual opossum
kills ; but, there is a strong possibil¬
ity that the larger percentage repre¬
sented rabbits killed on highways
or by hunters, or those deceased
through natural causes. Hamilton
(1958) proposed that opossums are
capable of killing cottontails in their
dens during winter ; he reported five
instances of predation by opossums
on nestling rabbits. Unfortunately,
the current study did not include
sufficient quantities of tracts from
the peak of the cottontail breeding
season to determine the effects of
predation on the young.

The absence of quail in the diet
of opossums is an interesting facet.
Field observations indicate that pre¬
dation by various mammalian forms
on quail nests is extensive, and opos¬
sums probably contribute to these

Food Habits of Opossum

207

losses. There is a considerable differ¬
ence of opinion as to the extent of
opossum predation on nesting birds ;
Wheeler (1939) found that they
were not destructive in this regard,
but Roberts and Early (1952) felt
that high opossum populations as¬
sociated with concentrations of ring¬
necked pheasant ( Phasianus colchi-
cus ) or rabbit nests might prove to
be detrimental to these game species.
Allen (1940) reported that in Michi¬
gan less than 5% of the nesting losses
of ducks and pheasants were caused
by opossums, but pointed out that
opossum populations were low dur¬
ing the investigations.

Summary

Digestive tracts from 131 opos¬
sums, taken from August 1, 1958-
March 1, 1960, in seven southern
Illinois counties, were examined ; all
tracts contained food. Three sea¬
sons were represented ; late summer
(August and September) : fall (Oc¬
tober and November) ; winter (De¬
cember, January, and February).

Animal materials made up 76.2%
of the total food volume, and plant
materials 23.8%. Both major food
groups appeared in all of the tracts.
Seventy -five animal and 66 plant
foods were recorded. The ten most
important foods by per cent volume
were opossum, 16.3 ; cottontail, 14.7 ;
persimmon, 8.1 ; domestic chicken,
7.1 ; prairie vole, 6.4; pokeberry, 5.1 ;
grackle, 4.7 ; gray fox, 2.8 ; frogs,
2.5 ; and scarabaeid larvae, 2.0. Ac¬
cording to per cent frequency of oc¬
currence the ten top-ranking foods
were grasses, 82.4; short -horned
grasshoppers, 54.2 ; opossum, 52.7 ;
ground beetles, 38.9; snails, 31.3;

pokeberry, 25.2 ; nightshade, 25.2 ;
stinkbugs, 22.9; persimmon, 21.4;
and cottontail, 15.3.

Noticeable variations in the utili¬
zation of foods seemed to reflect sea¬
sonal availability. The most marked
variation was a steady decrease in
consumption of plant materials from
fall through winter, and an increase
in utilization of mammalian foods
during the same period.

Predatory activities apparently do
not have a strong adverse effect on
game populations. However, preda¬
tion on cottontails might, on occa¬
sion, constitute a minor curb on
populations of this species.

Literature Cited

Allen, D. L. 1940. Nobody loves the ’pos¬
sum. Michigan Cons., 9 (6): 5, 10.
American Ornithologists’ Union. 1957.
Check-list of North American birds.
5th ed., Lord Baltimore Press, Balti¬
more. 691 pp.

Comstock, J. H. 1947. An introduction to
entomology. Comstock Publ. Co., Itha¬
ca, New York, 1064 pp.

Conant, R. 1958. A field guide to reptiles
and amphibians. Houghton Mifflin Co.,
Boston, 366 pp.

Dexter, R. W. 1951. Earthworms in the
winter diet of the opossum and rac¬
coon. J. Mammal. 32 (4) : 464.
Fernald, M. L. 1950. Gray’s manual of
botany. 8th ed. American Book Co.,
New York, 1632 pp.

Hall, E. R. and K. R. Kelson. 1959. The
mammals of North America. Ronald
Press Co., New York, vol. 1, 546 pp.
Hamilton, W. J., Jr. 1943. The mammals
of eastern United States. Comstock
Publ. Co., Ithaca, New York, 432 pp.
Hamilton, W. J., Jr. 1958. Life history
and economic relations of the opossum,
Didelphis marsupialis virginianus, in
New York State. Cornell Univ. Agri.
Exp. Sta. Memoir 354: 1-48.

Hartman, C. G. 1952. Possums. Univer¬
sity of Texas Press, Austin, 174 pp.
Lay, D. W. 1942. Ecology of the opossum
in eastern Texas. J. Mammal., 23 (2):
147-159.

208

Transactions Illinois Academy of Science

Lewis, J. B. 1929. Opossum in captivity.
J. Mammal., 10 (2): 167-168.

Llewellyn, L. M. and P. M. Uhler. 1952.
The foods of fur animals of the Pa¬
tuxent Research Refuge, Maryland.
Am. Midi. Nat., 48 (1) : 193-203.

Murie, A. 1936. Following fox trails.
Univ. Mich. Misc. Publ. Mus. Zool.,
32:1-45.

Pray, L. L. 1921. Opossum carries leaves
with its tail. J. Mammal., 2 (2): 109-
110.

Raven, H. C. 1929. A case of matricide in
the opossum. J. Mammal., 10 (2): 168.

Reynolds, H. C. 1945. Some aspects of
the life history and ecology of the
opossum in central Missouri. J. Mam¬
mal., 26 (4): 361-379.

Roberts, H. A. and R. C. Early. 1952.
Mammal survey of southeastern Penn¬
sylvania. Pennsylvania Game Comm.,
Harrisburg. Final Report PR Project
43-4, 70 pp.

Sandidge, L. L. 1953. Food and dens of
the opossum ( Didelpliis virginianus)
in northeastern Kansas. Trans. Kans.
Acad. Sci., 56 (1) : 97-106.

Scott, T. G. and W. D. Klimstra. 1955.
Red foxes and a declining prey popu¬
lation. Southern Ill. Univ. Monog. Ser.
No. 1, 123 pp.

Seton, L. 1929. Lives of game animals.
Doubleday, Doran and Co., Garden
City, New York, vol. 4, 949 pp.

Smith, L. 1941. An observation on the
nest-building behavior of the opossum.
J. Mammal., 22 (2) : 201-202.

Taube, C. M. 1947. Food habits of Michi¬
gan opossums. J. Wildl. Mgt., 11 (1):
97-103.

Wheeler, R. J. 1939. Food habits of the
opossum in Sumter County, Alabama.
Unpublished M. S. Thesis, Library, Al¬
abama Polytechnic Institute, Auburn,
56 pp.

Wiseman, G. L. and G. O. Hendrickson.
1950. Notes on the life history and
ecology of the opossum in southeast
Iowa. J. Mammal., 31 (3): 331-337.

Wood, J. E. 1954. Food habits of furbear-
ers of the upland post oak region of
eastern Texas. J. Mammal., 35 (3):
406-415.

A LATE PLEISTOCENE MUSK-OX FROM
EAST-CENTRAL ILLINOIS

EDWIN C. GALBREATH
Southern Illinois University , Carbondale

Recently, Mr. R. E. McClusky of
the Ambraw Gravel Company, Law-
renceville, Illinois, presented the axis
of a musk-ox, Symbos cavifrons, and
fragments of a proboscidean limb
bone to the Zoology Department of
Southern Illinois University for in¬
clusion in the Vertebrate Paleon¬
tology Collection.

These fossils were collected in the
NW % of Sec. 27, T 11 N, R 4 W,
Lawrence County, Illinois, which
would be on the west edge of George
Field, an abandoned Army airfield
located approximately four miles
northeast of Lawrenceville. My in¬
spection of the beds at this locality
and the nature of the preservation
of the bones suggest that the fossils
came from a thin layer of brownish
gravels, sands, or silts that overlay
the thick, widespread beds of gravel
and sand in the area. Dr. George E.
Ekblaw, of the Illinois Geological
Survey, informs me that the gravel
in this area that composes the “ sec¬
ond bottom” terrace at Lawrence¬
ville was derived from glaciers ter¬
minating to the north in Indiana
and were much later in time than the
Shelbyville glacier.

I11 my opinion, this dating is in
keeping with age determinations for
other late Pleistocene fossils, includ¬
ing musk-ox, found in the Embarass

drainage system almost 60 miles to
the north (Galbreath, 1938).

The axis (No. P200, Vert. Paleont.
Coll., Zool. Dept., S.I.U.) and the
proboscidean fragments are heavy,
massive, and completely mineralized
fossils, well-charged with iron oxide
that gives them a reddish orange
color. The axis is damaged. Only
the centrum and neural arch are
preserved, but this is enough to en¬
able one to recognize that the bone
belonged to a musk-ox. Comparison
of the specimen with the axis of
a musk-ox reported by Hibbard and
Hinds (1960) and identified by them
as Symbos cavifrons leaves no doubt
that the two bones belong to the same
species. Fortunately, comparable
bones or other parts from related
musk-oxen are known (Kitts, 1953)
that enables one to eliminate these
species from possible consideration
in determining the identity of this
bone. The measurements of this
Lawrenceville specimen are so close
to those reported by Hibbard for
his specimen that I see no reason
to do more than report that the two
specimens are similar in size.

This discovery of a musk-ox adds
one more record to the list of known
kinds of musk-oxen found in Illinois,
and is the southernmost reliable rec¬
ord for the distribution of Symbos
cavifrons in Illinois.

210

Transactions Illinois Academy of Science

Cited References

Gajlbreath, E. C. 1938. Post-glacial fossil
vertebrates from east-central Illinois.
Geol. Ser. Field Mus. Nat. Hist., 6
(20) : 303-313, 2 figs.

Hibbard, C. W. and F. J. Hinds. 1960. A
radiocarbon date for a woodland musk¬

ox in Michigan. Papers Michigan Acad.
Sci., Arts, and Letters, 45:103-111, 2
pis.

Kitts, D. B. 1953. A Pleistocene musk-ox
from New York and the distribution
of the musk-oxen. Amer. Mus. Nat.
Hist. Novitates, No. 1607: 1-8, 2 figs.

CURRENT PROBLEMS BEARING ON THE METABOLIC
STABILITY OF DEOXYRIBONUCLEIC ACID (DNA)

WESLEY J. BIRGE

University of Minnesota, Morris

In considering the orderliness and
precision involved in the develop¬
ment of an organism, such as in the
structural and functional differen¬
tiation of cells and tissues in multi¬
cellular forms, it becomes immedi¬
ately apparent that there must be
some system on which final order
and form are based.

Furthermore, when we consider
the transmission of heritable char¬
acters from parent to offspring, it
is still further apparent that such
a system must be capable of retain¬
ing a storehouse of information or
a memory of specificities from gen¬
eration to generation. In this ca¬
pacity, deoxyribonucleic acid
(DNA) is generally regarded as be¬
ing the principal encoding mecha¬
nism for genetic information
(Beadle, 1957; Brachet, 1957;
Hotchkiss, 1955). During recent
years, with the advent of reliable
information as to the structure of
the genetic material (Watson and
Crick, 1953), it has been possible
to give much greater meaning to
the term “ genetic information.”
Such information is visualized as
being represented in the specific mo¬
lecular organization of DNA.

Also, in reference to the template
hypothesis (Brachet, 1955; 1957),
it is possible to visualize mechanisms
which can facilitate the translation
of the specific information stored
in the genetic material into the
equally specific structural identity

of macromolecules such as those syn¬
thesized during periods of growth
and differentiation. In this connec¬
tion, an intermediate substance act¬
ing in the effective transfer of ge¬
netic information from the gene to
the specific end products of genic
action is usually acknowledged. At
least for the most part, this inter¬
mediate substance would seem to be
ribonucleic acid (Brachet, 1957 ;
Spiegelman, 1957).

With specific reference to DNA,
this material is generally regarded
as being a very stable substance. Ac¬
tually, many investigators regard
the relative constancy of the deoxy¬
ribonucleic acid in the “resting”
cell nucleus as constituting a gen¬
erally accepted hypothesis in mod¬
ern biology. This assumption of con¬
stancy arises from a number of
observations. First, it is accorded
support by the fact that, except for
periods of duplication, the DNA con¬
tent per chromosome set is supposed¬
ly constant for any one species. This
was first suggested by Boivin, Ven-
drely and Vendrely (1948), Mirsky
and Ris (1949) and was later sup¬
ported by numerous other investiga¬
tions (Alfert and Swift, 1953; Swift,
1950). A second supporting evi¬
dence for this hypothesis lies in the
general acceptance that, except for
periods of gene replication, the low
rate of turnover exhibited by deoxy¬
ribonucleic acid is indicative of high
metabolic stability (Kihara, et al.,

[ 211

212

Transactions Illinois Academy of Science

1956 ; Smellie, 1955 ; Swick, et al.,
1956). Finally, such data, of course,
tend to fit in with the general belief
that DNA, as the genetic encoding
material, must be maintained at a
constant level and carefully con¬
served in interest of the genetic in¬
tegrity of living organisms.

Although much support has been
amassed in favor of the constancy
hypothesis, the question may still be
raised as to the absolute universality
of this concept for all biological sys¬
tems and for all physiological cir¬
cumstances. Indeed, a considerable
amount of data has been accumu¬
lated over recent years which neces-

«/

sitates a re-examination of this con¬
cept, at least in certain instances.
Inconstancy has been reported in
various developing and secretory tis¬
sues (Finamore and Yolkin, 1958;
Leuchtenberger and Schrader, 1952 ;
Moore, 1957 ; Pelc, 1959 ; Rudkin
and Corlette, 1957 ; Stich and Nay¬
lor, 1958, and others) and has al¬
legedly been induced by cold treat¬
ment (LaCour, et al., 1956; Stich
and Naylor, 1958), hormonal
changes (Common, et al., 1951;
Lowe, 1955; McShan, et al., 1950),
etc. It is quite apparent that in
many cases where instability in the
metabolic activity of DNA has been
reported, such behavior has been di¬
rectly related to concomitant varia¬
tions in cellular proliferation and,
therefore, to DNA synthesis involved
in chromosomal replication. Such
data, of course, do not stand in refu¬
tation of the constancy concept.
However, at least some of the studies
referred to here {e.g., Finamore and
Yolkin, 1958 ; Moore, 1957 ; Pelc,
1959; Stich and Naylor, 1958) ap¬

parently are not resolvable on this
basis and, indeed, seemingly stand
in contradiction to the original con¬
text of the constancy hypothesis.

It is not the principal intent in
this study to present a comprehen¬
sive review of the literature which
stands in contradiction to the con¬
stancy hypothesis, as this has been
done by various other investigators
(Brachet, 1957; Govaert, 1957;
Moore, 1957). Instead, chief concern
will rest with an approach to the
causal analysis of factors which may
possibly underlie certain cases of
DNA instability and the possible
functional significance of such re¬
ported phenomena in nucleic acid
biology.

In this connection, reference
should be made to the extra DNA
known to occur in the cytoplasm of
many yolk-laden animal eggs
(Fraenkel-Conrat, et al., 1952; Hoff-
Jorgensen and Zeuthen, 1952; Solo¬
mon, 1957). It is thought that this
material may represent a general
storage reservoir which functions to
support DNA synthesis during early
embryonic development (Hotchkiss,
1955; Solomon, 1957). In reference
to this “cytoplasmic DNA”, Solo¬
mon (1957, p. 589) states, “The
nucleic acids (or similar highly
polymerized compounds) may be a
convenient means of storing nucleic
acid precursors, which could be ob¬
tained by degradation when required
by the embryo.” Commenting on
the same point, Hotchkiss (1955)
suggests that this substance may
very likely exist as a genetically
nonspecific precursory form of DNA.

Of interest here is the recent work
of Foster and Stern (1958, 1959)

Metabolic Stability of DNA

213

which indicates that extra sources
of DNA are exploited to support
DNA replication in developing pol¬
len of lily anthers. They have shown
that the breakdown products from
DNA of certain neighboring tissues
serve as a source of deoxynucleosides
for DNA synthesis in the micro-
sporocytes and microspores. These
findings stress the possible worth
of “extra sources” of DNA in pro¬
viding prescursory substances for
nucleic acid synthesis.

Attention should also be directed
to studies on the “puffs” of the
salivary gland chromosomes of cer¬
tain species (Beermann, 1959 ; Rud¬
kin and Corlette, 1957 ; Stich and
Naylor, 1.958, and others). Puff for¬
mation seems to be quite specific for
particular chromosomal segments,
varying characteristically with dif¬
ferent cell types and developmental
stages. As noted by Beermann
(1959), such behavior perhaps re¬
flects specific genic activity. Of spe¬
cial interest here is the localized
build-up of DNA known to occur
during puff formation in certain
species. As shown by Stich and Nay¬
lor (1958), certain puffs in Glypto-
tendipes (Chironomidae) show as
much as an 8-fold increase in DNA
content at certain developmental
periods. Also, it is apparent that
the DNA content of a particular
puff varies independently from other
segments of the same chromosome.
It seems possible that in some in¬
stances specific fractions of DNA
(or high molecular weight polyde-
oxyribonucleotides) may form at
certain chromosomal sites and, upon
being released, perhaps serve in
transmitting genetic information,

similar to messenger RNA. As DNA
does not commonly occur in cellular
cytoplasm, such polydeoxyribonu-
cleotides would perhaps function in
the intranuclear synthesis of certain
specific macromolecules, presumably
by playing an intermediate role in
information transfer from specific
genic loci to specific end-products
of genic action.

It is acknowledged that the sug¬
gestions noted above are largely ten¬
tative, and that the pertinence of
such ideas to nucleic acid biology
cannot be fully determined at pres¬
ent.

Reference

Alfert, M. and H. Swift. 1953. Nuclear
DNA constancy: A critical evaluation
of some exceptions reported by Lison
and Pasteels. Exptl. Cell. Res., 5:
455-460.

Beadle, G. W. 1957. The role of the
nucleus in heredity. In The Chemical
Basis of Heredity, edited by W. D.
McElroy and B. Glass. Johns Hopkins
Press, Baltimore, pp. 3-22.

Beermann, W. 1959. Chromosomal dif¬
ferentiation in insects. In Develop¬
mental Cytology, edited by D. Rud-
nick. Ronald Press, New York, pp. 83-
103.

Birge, W. J., G. W. Salisbury, L. de la
Torre, and J. R. Lodge. 1960. Diminu¬
tion of the deoxyribonucleic acid con¬
tent of bovine spermatozoa during in
vitro storage. Anat. Rec., 137: 340.
Birge, W. J., G. Anklesaria and F. D.
Tibbitts. 1961. Differential response of
DNA of fresh and stored bovine sper¬
matozoa to Feulgen (HC1) hydrolysis.
Trans. Ill. Acad. Sci., 54: 107-111.
Boivin, A., R. Vendrely, and C. Vend-
rely. 1948. L’acide desoxyribonuclei-
que du noyau cellulaire, depositaire
des characteres hereditaires; argu¬
ments d’ordre analytique. Compt. Rend.
Acad. Sci., 226: 1061-1063.

Brachet, J. 1955. The biological role of
the pentose nucleic acids. In The Nu¬
cleic Acids. Vol. II, edited by E. Char-
gaff and J. N. Davidson. Academic
Press, New York, pp. 475-519.

214

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Brachet, J. 1957. Biochemical Cytology.
Academic Press, Inc., New York, 1st
Ed.

Common, R. H., D. G. Chapman, and
W. A. Maw. 1951. The effect of gona¬
dal hormones on the nucleic acid con¬
tent of liver and serum in the im¬
mature pullet and the difference be¬
tween the nucleic acid content of
livers of sexually mature pullets and
cockerels. Can. J. Zool., 29: 265-275.

Finamore, F. J. and E. Volkin. 1958.
Nucleotide and nucleic acid metabol¬
ism in developing amphibian embryos.
II. Composition and metabolic activ¬
ity of ovarian egg nucleic acids. Exptl.
Cell. Res., 15: 405-411.

Foster, T. S. and H. Stern. 1958. Solu¬
ble deoxyribosidic compounds in rela¬
tion to duplication of deoxyribonucleic
acid. Science, 128: 653-654.

Fraenkel-Conrat, H., N. S. Snell and
E. D. Ducay. 1952. Avidin I. Isola¬
tion and characterization of the pro¬
tein and nucleic acid. Arch. Biochem.
Biophys., 39: 80-96.

Govaert, J. 1957. Etude quantitative
de la teneur en acide desoxyribonu-
cleique des noyaux des cellules somati-
ques et germinatives chez Fasciola
hepatica. Arch. Biol. (Liege), 68: 165-
200.

Hoff- Jorgensen, E. and E. Zeuthen.
1952. Evidence of cytoplasmic deoxy-
ribosides in the frog’s egg. Nature,
169: 245-246.

Hotchkiss, R. D. 1955. The biological
role of the deoxypentose nucleic acids.
In The Nucleic Acids, Vol. II, edited
by E. Chargaff and J. N. Davidson.
Academic Press, New York, pp. 435-
473.

Kihara, H. K., N. Amano, and A. Siba-
tani. 1956. Stability of deoxypentose
acid in growing and non-growing liv¬
ers of young rats. Biochem. et Biophys.
Acta, 21: 489-499.

LaCour, L. F., E. M. Deely, and J.
Cheyen. 1956. Variations in the
amount of Feulgen stain in nuclei of
plants grown at different tempera¬
tures. Nature, 177: 272.

Leuchtenberger, C., and F. Schrader.
1952. Variation in the amounts of
desoxyribose nucleic acid (DNA) in
cells of the same tissue and its cor¬
relation with secretory function. Proc.
Natl. Acad. Sci., 38: 99-105.

Lowe, C. V. 1955. Effects of cortisone
on nucleic acid composition of rat
liver. J. Natl. Cancer Inst., 15: 1619-
1622.

McShan, W. H., J. S. Davis, S. W. Souk-

up, and R. K. Meyer. 1950. The nu¬
cleic acid content and succinic dehy¬
drogenase activity of stimulated pi¬
geon crop gland tissue. Endocrinology,
47: 274-280.

Mirsky, A. W. and V. Allfrey. 1957.
The role of the cell nucleus in develop¬
ment. In The Chemical Basis of De¬
velopment, edited by W. D. McElroy
and B. Glass. Johns Hopkins Press,
Baltimore, pp. 94-102.

Mirsky, A. and H. Ris. 1949. Variable
and constant components of chromo¬
somes. Nature, 163: 666-667.

Moore, B. C. 1957. DNA in diploid and
androgenetic amphibian hybrids. J.
Morph., 101: 227-274.

Pelc, S. R. 1959. Metabolic activity of
DNA as shown by autoradiographs.
Lab. Invest., 8: 225-236.

Rudkins, G. T. and S. L. Corlette. 1957.
Disproportionate synthesis of DNA in
a polytene chromosome region. Proc.
Natl. Acad. Sci., 43: 964-968.

Salisbury, G. W., W. J. Birge, L. de la
Torre, and J. R. Lodge. 1960. Decrease
in nuclear Feulgen-positive material
(DNA) upon aging in in vitro storage
of bovine spermatozoa. J. Biophysical
and Biochemical Cytology, 10: 353-

359.

Smellie, R. M. S. 1955. The metabolism
of the nucleic acids. In The Nucleic
Acids, Vol. II, edited by E. Chargaff
and J. N. Davidson. Academic Press,
New York, pp. 393-434.

Solomon, J. B. 1957. Nucleic acid con¬
tent of early chick embryos and the
hen’s egg. Biocliimica et Biophysica
Acta, 24: 584-591.

Spiegelman, S. 1957. Nucleic acids and
the synthesis of proteins. In The
Chemical Basis of Heredity, edited by
W. D. McElroy and B. Glass. Johns
Hopkins Press, Baltimore, pp. 232-267.

Stern, H. 1959. Mitosis and metabolic
organization. The Botanical Review,
25: 351-384.

Sticti, H. F. and J. M. Naylor. 1958.
Variation of desoxyribonucleic acid
content of specific chromosome reg¬
ions. Exptl. Cell. Res., 14: 442-445.

Swick, R. W., A. L. Koch, and D. R.
Handa. 1956. The measurement of
nucleic acid turnover in rat liver.
Arch. Biochem. and Biophys., 63: 226-
242.

Swift, H. 1950. The desoxyribose nu¬
cleic acid content of animal nuclei.
Physiol. Zool., 23: 169-198.

Watson, J. D. and F. H. C. Crick. 1953.
Molecular structure of nucleic acids.
Nature, 171: 737-738.

CORRELATION BETWEEN PHENOLOGY AND CALORIC

CONTENT IN FOREST HERBS

BARBARA J. KIECKHEFER

University of Illinois, Uj’bana

Several phenological studies have
been made in which factors such as
breaking of dormancy, flowering,
fruiting and seed dispersal of forest
herbs have been recorded (Wolfe,
Wareham and Scofield, 1949; Leo¬
pold and Jones, 1947 ; Deam, 1920-
1952). Smith (1915) and Hopkins
and Murray (1933) found that the
time of occurrence of major events
in plant development remained rela¬
tively constant from year to year.
Lindsey and Newman (1956), on
the other hand, found considerable
variation for at least the first flow¬
ering dates of many herbaceous spe¬
cies in Indiana. They considered
temperature to be the most impor¬
tant factor controlling time of flow¬
ering.

Closing of the canopy and leaf
abscission cause considerable changes
in the environment of the under¬
story plants. Expansion of tree
leaves causes a great reduction in
sunlight, temperature fluctuation,
precipitation, transpiration rate and
wind velocity at the forest floor.
These microenvironmental changes
presumably cause readjustments in
the physiological activity and phe¬
nological sequence of the plants in
the lower synusia. Meyer and An¬
derson (1952) reported that excep¬
tionally high respiration and assimi¬
lation rates occur in floral meristems
and that various foods, inorganic
compounds and water are translo¬
cated to developing flowers. They

stated, however, that little is actually
known about metabolic activities
during flower and fruit production.

Golley (1958) compiled a list of
caloric determinations for various
plants from contributions by several
investigators. However, none of these
determinations were done on a phe¬
nological basis. Golley (1960) made
caloric determinations on some of the
species comprising an old field com¬
munity in southern Michigan but
found little seasonal variation. While
considerable research is being done
on the consumption and expenditure
of energy in certain animal species
during various phases of their life
cycles, little or no research is being
done on the energy dynamics of in¬
dividual species of plants. The au¬
thor is unaware of any studies relat¬
ing changes in caloric content with
plant development. The objective of
this study was to determine whether
any changes in energy content occur
during the growing season, and if
so, whether they are correlated with
changes in plant phenology.

Methods

Native spring and early summer
flowering herbs were collected from
a 10 x 10 m plot in Trelease AVoods
(Section 1 T19N R9E) Champaign
County, Illinois. The species collect¬
ed included : Claytonia virginica
(spring beauty), Dicentra canaden¬
sis (squirrel corn), Trillium recurva-
tum (purple trillium), Osmorhiza

[215]

216

Transactions Illinois Academy of Science

longistylis (sweet cicely) and Hydro-
ph yll urn c a n a de nse ( waterleaf ) .
Nomenclature is that of Jones
(1950). Voucher specimens are on
deposit in the University herbarium.

Only one plot was used in order
to confine the collecting to a rela¬
tively uniform habitat. With re¬
duced variation in habitat, varia¬
tions in caloric content could be
more easily correlated with phe-
nological variation. Collections were
made at approximately one week
intervals from March 31 to June 1,
1960, and at approximately two week
intervals from the latter date until
August 3. Both roots and shoots
were collected. The number of
plants collected depended upon the
species and upon the plienological
condition. Approximately 50 g
(fresh weight) samples were collect¬
ed for each species. After extrane¬
ous material was removed and roots
were washed to remove soil, the sam¬
ples were oven dried for about 48
hours at 80° C and ground twice in
a Wiley mill (20 mesli/in).

Caloric content determinations
were made using a Parr adiabatic
oxygen bomb calorimeter. Two de¬
terminations were made on each sam¬
ple (except where widely divergent
results were obtained, in which case
a third determination was made).
Values were not corrected to an ash¬
free weight basis. Average caloric
values for carbohydrates (4100 cal/
g), proteins (5700 cal/g) and fats
(9400 cal/g) obtained from Fruton
and Simmons (1953) were used as
a standard for the interpretation of
experimental data.

A slightly modified form of
Nessler ’s Procedure (Umbreit, Bur¬

ris and Stauffer, 1957) was used for
analysis of total nitrogen content.
Duplicate 4mg samples were digested
in Nessler tubes by adding 1 ml of
20% H2S04 (no copper selenite

added) and heating on an electric
plate for about an hour. After cool¬
ing for at least a minute, 2 drops
of 30% H202 were added, and heat¬
ing was resumed for 15 minutes.
Cooling, adding of H2G2 and heating
were repeated a second time in order
to completely clear the samples.
When the tubes had cooled to room
temperature, 20 ml of deionized wa¬
ter were added and the samples
stirred vigorously. Stirring was re¬
peated after addition of 4 ml of 4N
KOH and again after addition (by
blowing) of 2 ml Nessler ’s reagent.
This solution was diluted to 35 ml
with deionized water and allowed
to stand for 15 minutes. Optical
density was measured with a spectro¬
photometer at 490 m/A. These read¬
ings were then converted to mg
protein/g dry weight (Nx6.25).

Though no quantitative environ¬
mental data were obtained, general¬
ized observations of climatic condi¬
tions were recorded along with
observations of plienological condi¬
tion of the species collected. The
latter included observations of the
time and amount of vegetative
growth, flower bud initiation, flower¬
ing, fruit development and vegeta¬
tive drying. A record was also kept
of the initiation, the gradual devel¬
opment and the complete closing of
the tree canopy.

Phenological Observations

The first observations of the study

Caloric Content of Forest Herbs

217

area were made on March 29. Al¬
though a heavy snow had just melted
three days previously, all of the spe¬
cies mentioned above had broken
dormancy. In fact, Claytonia , Hy-
drophyllum and Osmorhiza had al¬
ready penetrated the leaf litter. By
March 31, some Dicentra were be¬
ginning to appear above the litter.
Abundant etiolated stems of Clay¬
tonia were found below the humus,
and flower buds were already pres¬
ent. Trillium ranged in height from
4 to 6 cm, Osmorhiza from 2 to 3 cm.

On April 3 about 10% of Clay¬
tonia flowers were open with the
remaining buds showing pink color.
Only a small amount of vegetative
growth occurred in all species be¬
tween March 29 and April 8 due to
abnormally low temperatures (Illi¬
nois State Water Survey data,
March - April, 1960) . By this date,
however, large buds were present on
Trillium. Claytonia was in full
bloom on April 14 and continued to
be so for about 10 days. Although
flower buds were visible on Dicentra
on the 14th, full bloom was not at¬
tained until the 20th. Flowering
had just begun in Trillium, while
considerable leaf expansion had oc¬
curred in Osmorhiza and Hydro-
phyllum. Leaf buds were just be¬
ginning to expand in the canopy, but
leaf development was almost com¬
pleted on buckeye seedlings and sap¬
lings.

By April 27 most Dicentra and
Claytonia were through blooming,
while Trillium was in full flower.
Osmorhiza and Hydrophyllum con¬
tinued vegetative growth. Hydro¬
phyllum, rather than Claytonia and
Dicentra, now formed the predomi¬
nant herbaceous cover.

Although Dicentra produced very
few fruits, those which were formed
were mature on May 4. Leaves of
this species were turning yellow on
this date, and many new corms were
observed. The fruits of Claytonia
were nearly all mature, and the
petals of many Trillium flowers had
dropped. Flower buds were pres¬
ent on about 50% of Osmorhiza. The
leaves of Hydrophyllum had at¬
tained a height of about 45 cm. In
areas where Hydrophyllum was
sparse, Laportea canadensis had be¬
come very predominant in the her¬
baceous layer.

During the week from May 4 to
May 11, little change took place due
to cold, wet weather. However, Clay¬
tonia seeds were dispersed ; Dicentra
leaves were dying ; most Trillium
were through blooming. The tree
canopy began to close during this
week. By May 18 the canopy was
almost completely developed, Dicen¬
tra and Claytonia had disappeared
from the herbaceous layer and a
few Hydrophyllum were in bloom,
although flower buds were not yet
visible on many individuals.

On May 25 the canopy was fully
developed. Laportea was 0.7 to 1 m
in height, and yellowing was ob¬
served in Trillium. By June 1 Tril¬
lium was drying. The majority of
Osmorhiza were in fruit by the latter
date, while approximately 90% of
Hydrophyllum were in bud.

The author was not in Urbana
after June 1. Therefore, no further
field observations of phenological
condition were recorded. However,
while preparing samples for grind¬
ing, it was noted that fruits of Os¬
morhiza were green on June 23 and
had matured by July 7.

218

Transactions Illinois Academy of Science

Dates

486 .
5
■d

389

292 c

194 2
CL

97 O*
0

I486

• 389

■ 292. c

194

Fig. 1. — Caloric and protein determina¬
tions for Claytonia virginica, (fl-flower-
ing, fr-fruiting).

Dates

Fig. 2. — Caloric and protein determina¬
tions for Dicentra canadensis.

Kesults and Discussion

Caloric content was found to vary
with plant development. Claytonia
virginica (Fig. 1) and Dicentra ca¬
nadensis (Fig. 2) showed a peak
in caloric content at maximum flow¬
ering, while the peak for Trillium
recur vatum (Fig. 3) occurred at
maximum flower bud development
(general curves were drawn from
sight inspection). Caloric values for
Claytonia dropped from 4100 cal /g
at flowering to 3500 cal/g two weeks
after flowering. By the time of vege¬
tative die back, caloric content rose

to about 3750 cal/g. Dicentra and
Trillium showed a continuous drop
in caloric content following peak
values. Dicentra dropped from 4250
cal/g at flowering to about 4000
cal/g at vegetative die back, while
Trillium dropped from 4200 cal/g
at maximum floral bud devel¬
opment to about 3950 cal/g at dor¬
mancy. Osmorhiza longistylis (Fig.
4) and Hydrophyllum canadense
(Fig. 5), on the other hand, showed
a continuous increase in calories per
gram over the entire growing sea¬
son. Caloric values for Osmorhiza

Caloric Content of Forest Herhs

219

486

389 ^
cn

292 c
o>

■ 194 O
Q.

Dates

Fig. 3. — Caloric and protein determina¬
tions for Trillium recurvatum.

increased from about 3500 cal/g at
the beginning of the season to ap¬
proximately 4100 cal/g at the end.
Values for Hydrophyllum rose from
3250 cal/g to 3800 cal/g followed
by a slight drop (to 3700 cal/g) at
the end of the summer.

Corrections of caloric values to>
an ash-free weight basis were made
on limited amounts of material at
a later date. A rise in caloric values
occurred in every case, but the gen¬
eral seasonal pattern was main¬
tained. The extent to which the val¬
ues increased varied with the species.

The variation in time of maximum
caloric content between the species
could possibly be explained by in¬

herent differences. Differences in the
manner and time of floral develop¬
ment may be associated with differ¬
ences in time of peak caloric content
in Claytonia, Dicentra and Trillium.
Claytonia produces flowers and
leaves on the same stem. Flower
buds are already present when the
stem appears above the litter. The
same is true for Trillium except that
only one large flower bud is pro¬
duced per plant. Dicentra differs in
that flowers are produced on a scape
when vegetative growth is nearly
complete. Claytonia, Dicentra and
Trillium grow, reproduce and die
back within two months, while vege¬
tative growth occurs for a much
longer period of time in Osmorhiza
and Hydrophyllum. The ratio of
organic to inorganic matter probably
increases due to continuous vegeta¬
tive growth. This could account for
the continuous rise in caloric content
in the latter two species during the
growing season.

Total protein was also found to
vary with phenological development
in Claytonia, Dicentra and Trillium.
Maximum protein content in Clay¬
tonia (Fig. 1) occurred at maximum
flowering. Total protein dropped
from about 412 mg/g dry weight
at flowering to 170 mg/g a week
before complete die back. A slight
rise in protein occurred during the
last week resulting in a curve simi¬
lar to that for caloric content. Ap¬
parently the rise in caloric values is
related to a relative increase in pro¬
tein content. The latter may be due,
in Claytonia, to continued produc¬
tion of vegetative and floral struc¬
tures until flowering occurs.

The peak in caloric content for

100 cal /g d.w. 100 cal/g d.w.

220

Transactions Illinois Academy of Science

Fig. 5

Dates

Caloric and protein determinations for Osmorhiza longistylis.

486

309 d
o>

292 ^

194 O

i—

97 O'
J 0

Dates

Caloric and protein determinations for HydrophyTlum canaclense.

Caloric Content of Forest Herbs

221

Trillium lagged slightly behind the
peak in protein content. The latter
occurred during maximum vegeta¬
tive and floral bud development.
Total protein dropped continuously
from the peak of about 435 mg/g
to about 122 mg/g at vegetative die
back. It appeared that most tissue
production and differentiation was
completed somewhat before flower¬
ing and that further growth was
due to expansion of existing cells.
The relative increase in amount of
total protein during production and
differentiation of vegetative and flo¬
ral structures could account for most
of the increase in caloric content.
However, the higher caloric values
in Trillium probably indicate for¬
mation of some lipids.

In Dicentra the peak in protein
content occurred, as in Trillium,
during maximum vegetative and flo¬
ral bud development. Protein con¬
tent dropped continuously from a
maximum of about 365 mg/g at this
time to about 90 mg/g at vegeta¬
tive die back. A lag, greater than
that in Trillium, occurred between
peak protein content and peak calor¬
ic content. Leaf and flower bud
production and differentiation were
completed well before flowering.
Petiole and scape length increased
to the time of flowering, but leaves
simply unfolded and flower buds
expanded for a period of at least a
week prior to flowering. Increase
in total protein appeared, therefore,
to be related to the period of maxi¬
mum tissue formation and differ¬
entiation. Since the lag between
maximum protein and maximum
caloric content was considerable, rel¬
ative increase in the amount of pro¬

tein probably did not account for
the peak in caloric content in Dicen¬
tra. Lipid formation may account
for the peak, a possibility which is
supported by the relatively high
caloric values.

The initial values for caloric and
protein content for Claytonia, Dicen¬
tra and Trillium were well above
the values at vegetative die back.
Thus, some conversion of carbohy¬
drates to proteins or lipids must take
place in the bulbs, corms or root
systems of these species during the
dormant period.

Protein content per gram in Os-
morhiza and Hydrophyllum did not
vary with plant phenology. Both
species had values close to 97 mg/g
throughout the growing season.
Therefore, rise in caloric content can
not be attributed to a rise in protein
content. If an increased organic to
inorganic matter ratio occurs, the
increase must be in the form of
carbohydrates or fats. Since the to¬
tal rise in caloric content was great,
increased lipid formation is prob¬
ably the causal factor. Caloric val¬
ues for the last collection date were
well above initial values for both
of these genera. However, drying
had not occurred in either species
when collection ceased. A drop in
caloric values would probably occur
with the shedding of fruits and the
drying of leaves.

An analysis of variance was run
on this factorial experiment in a
completely randomized design. The
results show that caloric values for
different species, for different dates
and for the effect of species upon
dates were significant at the 1%
level.

222

Transactions Illinois Academy of Science

Summary

Three species of spring flowering
herbs ( Claytonia virginica, Dicentra
canadensis and Trillium recurva-
tum ) and two species of summer
flowering herbs ( Osmorhiza longi-
stylis and Hydrophyllum canadense )
were studied to determine whether
changes in plant development are
correlated with changes in caloric
and protein values. The plants were
collected from a 10 x 10 m plot in
Trelease Woods (Section 1 T19N
R9E) Champaign County, Illinois,
at 1 to 2 week intervals from March
31 through August 3, 1960. Time of
maximum vegetative growth, flower
bud development, flowering, fruiting
and vegetative die back were record¬
ed. Notes were also taken on canopy
development and on general cli¬
matic conditions. Caloric determina¬
tions were made on oven dried,
ground samples using a Parr adia¬
batic oxygen bomb calorimeter. A
modified form of Nessler’s Proce¬
dure was used to determine total
nitrogen content. The results were
then converted to total protein con¬
tent.

A correlation was found between
time of maximum leaf and flower
development and peak caloric con¬
tent in Claytonia, Dicentra and Tril¬
lium. These species grow, flower,
fruit and die back in a two month
period. Caloric values for Osmor¬
hiza and Hydrophyllum rose almost
continuously throughout the grow¬
ing season. These species are physio¬
logically active during most of the
summer and, thus, there appears to
be no correlation between peak calor¬
ic values and time of maximum
growth.

The curves for caloric and protein
content were quite similar in Clay¬
tonia and Trillium ; both reached a
peak at flowering. The peak in pro¬
tein content occurred well ahead of
the peak in caloric content in Dicen¬
tra, while in Osmorhiza and Hydro¬
phyllum, protein remained constant
(and low) during the entire grow¬
ing season. Since no significant in¬
crease in protein was found in the
latter two species, increased lipid
content may account for the rise in
caloric values. Some conversion of
carbohydrates to proteins or fats
must occur during dormancy in the
corms, bulbs or root systems of Clay¬
tonia, Dicentra and Trillium in or¬
der to bring caloric values back to
their initial value by the following
spring.

Acknowledgment

The author wishes to express her
gratitude to Dr. Lawrence C. Bliss
for his help and advice in all phases
of this study. She is also indebted
to Dr. Robert W. Howell for help
with protein determinations.

Literature Cited

Deam, C. C. 1920-1952. Unpublished
phenological records as cited in Lind¬
sey and Newman.

Fruton, J. S. and Sofia Simmons. 1953.
General biochemistry. New York, John
Wiley and Sons, Inc., 940 pp.

Golley, F. B. 1958. Table of caloric
equivalents. Unpub. data.

- . 1960. Energy dynamics of

a food chain of an old-field commun¬
ity. Ecol. Monog., 30:187-206.
Hopkins, A. D. and M. Murray. 1933.
Natural guides to the beginning,
length, and progress of the seasons.
Acta Phaenol., 2:33-43.

Illinois State Water Survey Division.
1960. Champaign-Urbana weather sum¬
mary. March - April - May.

Caloric Content of Forest Herbs

223

Jones, G. N. 1950. Flora of Illinois.
Notre Dame, Indiana, Univ. of Notre
Dame Press, 368 pp.

Leopold, A. and Sara E. Jones. 1947. A
plienological record for Sauk and Dane
Counties, Wisconsin, 1935-1945. Ecol.
Monog., 17:81-122.

Lindsey, A. A. and J. E. Newman. 1956.
Use of official weather data in spring
time — temperature analysis of an In¬
diana plienological record. Ecology,
37:812-823.

Meyer, B. S. and D. B. Anderson. 1952.
Plant physiology (2nd Ed.) Princeton,
New Jersey, D. Van Nostrand Co.,
784 pp.

Smith, J. W. 1915. Plienological dates
and meteorological data recorded by
Thomas Mikesell between 1873-1912
at Wauseon, Ohio. Month. Weath. Rev.
Sup. No. 2., pp. 23-93. (as cited in
Wolfe et al. 1949).

Umbreit, W. W., R. H. Burris and J. F.
Stauffer. 1957. Manometric tech¬
niques. Minneapolis, Minnesota, Bur¬
gess Pub. Co., 338 pp.

Wolfe, J. N., R. T. Wareham and H. T.
Scofield. 1949. Microclimates and
macroclimates of Neotoma, a small
valley in central Ohio. Ohio Biol.
Surv., Bull. 41, 267 pp.

A RE-EXAMINATION OF RETAIL TRADE IN
THE “DISPERSED CITY” OF SOUTHERN ILLINOIS

THOMAS R. GLENNON

Southern Illinois University

Whereas southern Illinois lacks an
urban center of more than 20,000
persons, several of its larger cities
have been said to function as a single,
albeit dispersed city. Reference to
this dispersed city was first made by
Oliver Beimfohr (1953:100), within
the context of a study focusing on
the industrial potential of southern
Illinois. A more detailed study by
Ian Burton (1959:145), on the basis
of traffic flow and population densi¬
ty, set the limits of the dispersed
city in a four county area composed
of Williamson, Perry, Franklin, and
Jackson counties. This area includes

an urban core of thirteen major
cities ranging in population size
from more than 14,000 to slightly
over 1,000 inhabitants. (Fig. 1)
Earlier works have failed to for¬
mulate a concise definition of the
term “dispersed city,” but we may
gather, especially from the study of
Burton (1959:145), that it refers
to a group of politically discrete
cities, separated by rather large
tracts of rural land, which function¬
ally inter-act to the extent that they
may be referred to as a single unit
lacking a downtown shopping dis¬
trict. The purpose of this paper is

Figure 1. — A Re-Examination of Retail Trade in the “Dispersed City.”

[224]

Trade in the “Dispersed City ”

225

to analyze the retail functions of the
major centers of the “dispersed
city” in an effort to ascertain if
inter-action among them is sufficient
to warrant calling them a single unit.

Methods

Inter-action among the various
centers is the key to the entire con¬
cept of the dispersed city. The best
insights into this phenomenon prob¬
ably could be gained from detailed
field work in each of the cities in¬
volved. Limitations in time and re¬
sources placed such an investigation
beyond the scope of this study.

Burton (1959:148-9) approached
inter-action by analyzing the rela¬
tionship between population and
several types of retail sales. He as¬
sumed that a relatively high degree
of association normally exists be¬
tween the variables examined and
proceeded to search for anamolies
in the existing pattern. For exam¬
ple, unless inter-action is remark¬
ably uniform, certain cities would
have disproportionately large vol¬
umes of retail trade because they are
able to attract business from other
sections of the dispersed city; spe¬
cialization of certain retail commodi¬
ties would be especially likely to
develop under such circumstances.

This study makes use of Burton’s
method, but goes beyond his and
other previous works by comparing
the retail patterns of the “dispersed
city” to similar groups of urban
units. This approach is based on the
assumption that if inter-action is
unique within the “dispersed city,”
it is probable that the associations
of population with the various cate¬
gories of retail trade in the “dis¬

persed city” differ markedly from
associations found in groups of cities
of similar sizes.

To form a basis for comparison
four sample groups of cities in Illi¬
nois were selected. Three of these
(A, B, and C) consisted of thirteen
cities each, and were structured to
approximate as closely as possible
the rank order of population of the
units found in the dispersed city.
Thus, the largest city in each group
approximated the population of Car-
bondale, the biggest unit in the ‘ ‘ dis¬
persed city;” the second largest city
in each group approached the popu¬
lation of Marion, the second largest
city in the “dispersed unit.” Sample
group D consisted of thirty-five cities
and was chosen at random from all
Illinois cities having populations of
more than 1,000. (Table I) Although
the small sizes of the samples neces¬
sitate caution when using statistical
analysis, a quantitative indication of
the variability between population
and several categories of retail trade
was obtained by correlation analysis.
(Table II)

Discussion

Burton (1959:147-8) pointed to
a weak association between popula¬
tion and total retail sales of the
larger units of the dispersed city
and cited this as evidence of inter¬
action. His conclusion was based on
a comparison of 1950 population fig¬
ures and 1957 sales tax data; on the
other hand when this author com¬
pared 1960 population statistics to
1960 sales tax data, a high positive
correlation was apparent. (Fig. 2)
In fact, the statistical relationship
between these variables was higher

226

Transactions Illinois Academy of Science

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Trade in the “ Dispersed City”

Table II. Correlation coefficients of population with retail sales tax receipts.

227

Categories of Retail Sales

II.

III.

IV.

VI.

VII.

VIII.

IX.

Dispersed City.

.99

.80

.98

.94

.88

.95

.79

.98

.93

.97

Sample groups

.97

.91

.92

.82

.93

.92

.54

.95

.93

.96

.92

.77

.93

.91

.77

.81

.51

.86

.85

.71

.98

.94

.98

.89

.80

.94

.82

.89

.97

.92

.86

.97

.92

,73

.93

.87

.71

.95

.98

Identification of categories: I — Total sales tax receipt, II — General merchandise,
III — Food, IV — Drinking and eating places, V — Apparel, VI — Furniture, radio, and
household goods, VII — Lumber, building, and hardware, VIII — Automotive, IX—
Filling stations, X — Other.

POPULATION

FIGURE 2

FIGURE 3

228

Transactions Illinois Academy of Science

FIGURE 4 FIGURE 5

than that found in any of the sample
groups.

Again using incomparable data,
Burton (1959:148-9) found a rela¬
tively low degree of association be¬
tween total population and two types
of retail sales — general merchan¬
dise and radio, furniture, and house¬
hold goods. These relationships were
cited as evidence of retail specializa¬
tion based upon inter-action. Yet the
use of 1960 figures showed a rela¬
tionship between radio, furniture,
and household goods and total popu¬
lation which was relatively high and
failed to differ significantly from the
associations found in the sample
groups. (Fig. 3) General merchan¬

dise also showed a relatively high
degree of association with total pop¬
ulation in the dispersed city, differ¬
ing little from the relationships in
the sample groups. (Fig. 4)

Examination of additional cate¬
gories of retail sales which might
be considered “non-convenience”
goods, and therefore prone to spe¬
cialization, yielded similar results.
Automotive sales showed a very
strong correlation with population
in the study area (Fig. 5) ; apparel
sales and lumber and building ma¬
terials also were associated with pop¬
ulation in the “dispersed city” to a
higher degree than in some of the
sample groups.

Trade in the “Dispersed City”

229

Summary

The above analysis has indicated
that population is very closely re¬
lated to retail sales in the dispersed
city area. Various commodities in
which retail specialization seemed
likely, are also quite closely corre¬
lated with total population. Previ¬
ous conclusions to the contrary must
be attributed largely to the use of
incomparable statistics. Further¬
more, although earlier workers im¬
plied that the retail patterns of the
dispersed city were unique, they
failed to compare those patterns with
others found outside the study area.
Such a comparison has yielded little
to support the concept that the re¬
tail trade patterns of the dispersed
city are unique. It is therefore con¬
cluded that neither the degree of
retail inter-action, nor the unique¬
ness of the retail trade patterns is
sufficient to warrant the amalgama¬
tion of thirteen politically discrete

cities of southern Illinois into a sin¬
gle unit, known as a “ dispersed
city”.

Acknowledgments

The author would like to express
his appreciation to Dr. Howard A.
Stafford, Jr., for his helpful sug¬
gestions and critical comments made
during the preparation of this manu¬
script, and also to the Mississippi
Valley Investigations, under the di¬
rection of Dr. Charles C. Colby, for
drafting of the graphs used in this
paper, and to Joseph Shramovich for
drafting the map.

Literature Cited

Beimfohr, O. W. 1953. Some factors
in the industrial potential of southern
Illinois. Trans. Ill. Acad. Sci., 1946:
97-103.

Burton, Ian. 1959. Retail trade in a
dispersed city. Trans. Ill. Acad. Sci.
52: 145-150.

Illinois Retailers' Occupation Tax Re¬
turns. 1960. State of Illinois Depart¬
ment of Revenue. Springfield, 30 pp.

SHOPE’S FIBROMA IN ILLINOIS COTTONTAILS

D. H. FERRIS, R. D. LORD, and D. L. HUXSOLL

University of Illinois, Urbana

Shope’s fibroma in cottontail rab¬
bits, Sylvilagus florid anus, has been
found frequently in states east of
the Mississippi (Herman, Kilham,
and Warbach, 1956) but to our
knowledge, this is the first time it
has been reported from Illinois. Al¬
though experimental transmission of
the virus by fleas, (Kilham and
Woke, 1953) mosquitoes, (Dalmat,
1959; Dalmat and Stanton, 1958;
and Kilham and Woke, 1953) redu-
viid bugs, (Dalmat, 1959) and bed¬
bugs (Dalmat, 1959) has been ac¬
complished, the life cycle, reservoir
and vector complex of the natural
disease are not yet known.

To gain a better understanding of
potential vectors and other aspects
ot the epizootiology of fibromatosis,
an investigation of the prevalence
of fibromas in cottontails from two
different habitats was undertaken.
The first habitat (in Allerton Park
near Monticello, Illinois) was basi-
caJJy sylvan, with thick woods and
fallow fields, but with no cultivated
land. The second habitat was basi¬
cally campestral, composed of agri¬
cultural fields almost completely
devoid of woody cover. It consisted
of cultivated fields in Piatt and
three nearby counties.

Ackn o wledgment

Thanks are due Dr. A. M. Watrach
for histologic examination of tumor
materials and to Mrs. Marion Wa¬
trach for the microscopic prepara¬
tions. It is also a pleasure to ac¬

knowledge the encouragement and
suggestions of Dr. Richard E. Shope
and Dr. Herbert J. Dalmat.

Materials and Methods

This study was carried out be¬
tween March 1956 and February
1960.

Rabbits from the sylvan habitat
were trapped and shot, largely dur¬
ing the fall and winter months.
Those from the campestral habitat
were collected each month of the
year from Champaign, Piatt, Mc¬
Lean and Ford Counties in central
Illinois. A few rabbits from the
same type of habitat were included
from the southern part of the state.
Those from the campestral habitat
were collected entirely by shooting.

The fibromas were examined his¬
tologically and attempts were made
to isolate viruses from them. Tumors
from both live and dead rabbits were
removed aseptically and triturated
either in sterile broth containing
2500 units of penicillin and 2500
micrograms of streptomycin per 0.1
ml of inoculum, or in sterile broth
without antibiotics; 0.1 ml of this
material was inoculated intradermal-
ly into the base of the external ears
or the scrotums of domestic rabbits
( Oryctolagus cuniculus) and cot¬
tontails ; 0.2 ml were inoculated onto
the chorio-allantoic membrane of 9
to 11 day-old embryonating* chicken
eggs.

Portions of tumors from living
and dead rabbits were fixed in 10%

[ 230 ]

8 'hope’s Fibroma in Cottontails

Table 1. — Fibromas Found in Rabbits Collected from Sylvan Habitats.

231

Period

Rabbits

Collected

Rabbits with
Fibromas

March 1, 1956

to February 28, 1957 . . .

261

March 1, 1957

to February 28, 1958 .

344

March 1, 1958

to February 28, 1959 . . .

381

March 1, 1959

to February 28, 1960 . . .

268

Totals . . . .

1244

Prevalence for Period 1956 to 1960: 0.72 per cent.

* Including one rabbit later in 1960.

formalin. Sections were made from
these and examined histologically.

Serial passages of the virus were
made in cottontails and domestic
rabbits as well as in chicken em¬
bryos. Tumor materials and mem¬
branes from infected chicken em¬
bryos were stored at — 10 C.

Results

A total of 1,506 cottontail rabbits
collected over a 4-year period from
the campestral habitats yielded no
fibromas. In contrast, 9 infected
rabbits were found among 1,244 rab¬
bits collected from the sylvan habi¬
tat in the same part of the state
(Table 1). The disease was much
more prevalent in 1959 than in previ¬
ous years. A single case was found
in 1956, another in 1958, while 7
were found in the winter of 1959-60.
While more rabbits were collected
from the campestral areas than from
the sylvan areas, no fibromas were
found in the campestral group.

A study of the seasonal prevalence
(Table 2) shows that by far the
greatest number of fibromas were
found in the fall months. Eight
were found during this period, while
only one was obtained in the early
winter and none in the late winter.
The affected rabbits were about
equally divided by sex (Table 3).
Only one adult female had a fibroma ;
all others were found on juveniles.
Most of the tumors were found on
the feet or parts of the legs touching
the ground (Fig. 1). One rabbit
had a tumor on the left ear and
another had one on the nose (Fig.
1). One rabbit had four tumors,
two had two each, and the other six
had one each.

Serum neutralization tests were
carried out on domestic rabbits us¬
ing convalescent serum kindly sup¬
plied by Dr. Richard Shope and also
from our own infected rabbits. The
supernatant fluid from triturated
tumor material centrifuged at 1000
RPM for 10 minutes was allowed to

Table 2. — Seasonal Distribution of Fibroma Infections in Rabbits from the

Sylvan Habitat.

232

Transactions Illinois Academy of Science

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Shope’s Fibroma in Cottontails

233

Table 3. — Age and Sex of Rabbits Affected with Fibroma Tumors
and Anatomical Distribution of the Tumors.

Date

Sex

Age

wt. (g)

No. Tumors

Part Affected

10/24/56

Juv.

1219

Left front foot

9/3/58

Juv.

1389

Foot

9/2/59

Adult

1276

Leading edge, left ear

9/24/59

Juv.

794

Right front foot

9/25/59

Juv.

1191

Both hind feet

Right front foot

Right front knee

10/1/59

....

Hind feet

10/7/59

Juv.

1361

Left hind foot

11/23/59

Juv.

1134

Left front leg

Left hind leg

9/15/60*

Juv.

....

Nose

Left front leg

* Collected after the main period of investigation, but included in 1959 and 1960 figures.

Fig. 1. — Natural Case of Shope’s Fibroma in the Cottontail Rabbit. There is a
lesion on the nose and another on the fore foot, which is held against the rabbit’s
body.

234

Transactions Illinois Academy of Science

Fig. 2. — Plaques from Shope’s Fibroma
Virus on the Chorio- Allantoic Membrane
of an Embryonating Chicken Egg.
(From a color transparency.)

incubate at room temperature with
an equal amount of serum for one
and one-half hours after which 0.1
ml of the mixture was injected intra-
dermally into the shaven sides of
young domestic rabbits. An equal
amount of the triturate diluted to
the same extent with sterile saline
was inoculated into the other side
of the rabbit. In addition, rabbits
of various ages were inoculated in¬
dividually with the virus or with
the virus-antigen mixture. Addi¬
tional controls consisted of rabbits
inoculated with normal rabbit tissue
suspension. Both the homologous
serum and serum supplied by Dr.
Shope neutralized the virus as shown
by the appearance of fibromas on

Fig. 3. — Shope’s Fibroma on the Ear of Cottontail. The lesion above resulted from
intradermal inoculation of triturated fibroma material into the ear of a cottontail
and is typical of first, second and third passage lesions.

/S hope’s Fibroma in Cottontails

235

all sites where the virus alone was
inoculated. No firomas appeared
where the antibody-antigen mixture,
as described above, was inoculated
or in those rabbits receiving the nor¬
mal tissue-saline inoculation.

Histologic examinations were
made of 2 field cases and three ex¬
perimentally infected animals. In all
cases tissue changes characteristic of
Shope’s fibroma were found. Nu¬
merous inclusion bodies were seen
in epithelial cells of the tumor. The
virus was transmitted easily to 11
day-old embryonating chicken eggs,
but less easily to 9 day-old chicken
embryos. Serial passages in 10 and
11 day-old embryonating eggs were
made. The virus did not kill the
embryos but both large and small
plaques formed on the chorio-allan-
toic membranes (Fig. 2). The in¬
fection was readily transmitted to
domestic and wild rabbits (Fig. 3).
In young domestic rabbits the tu¬
mors appeared in 3 to 4 days, reached
a peak in 2 weeks and regressed
within another 2 weeks. Tumors in
cottontails were visible at 8 to 10
days and did not regress for 2 to
3 months or longer. Tumors surgi¬
cally removed from each species
were examined histologically and
found to be indistinguishable from
those of naturally occurring cases
although the gross appearance of
the experimentally induced tumors
was different from that of the nat¬
ural lesions (Figs. 1 and 3).

The virus was readily transmitted
from either natural or experimental
lesions from the cottontail, but not
from the domestic rabbit. When
experimentally induced tumors were
removed from wild rabbits they were
quickly replaced by proliferation of

tumor material at the site, but this
did not occur in domestic rabbits.

Discussion

The natural mode of transmission
of Shope ’s fibroma remains un¬
known, although a considerable
amount of experimental work has
been done on the problem. Analyses
of the data, correlated with the find¬
ings reported here may give clues
for further research. The largest
previous investigation was done by
the Rose Lake Wildlife Experiment
Station in Michigan (Annual Re¬
ports, 1940-41, 1941-42, 1942, 1943,
1944, 1945). Lesions were found in
2.7% of 1071 rabbits examined be¬
tween 1940 and 1945. Herman et. al.
(Herman, Kilham, and Warbach,
1956) found 4.1% of 359 Maryland
rabbits at the Patuxent Research
Refuge involved between 1947 and
1953. Reilly (Herman, Kilham and
Warbach, 1956) found 8.5% of 174
rabbits from the Three Rivers Game
Management Area, Baldwinsville,
N. Y. with fibromas in 1953.

From the standpoint of numbers
and regularity of collection this in¬
vestigation was biased in favor of
finding more fibromas in rabbits
from the campestral habitats. That
fibromas were found in rabbits only
from the wooded terrain suggests
the possibility of a sylvan vector or
reservoir.

Dalmat (Dalmat, 1959) found the
following mosquitoes capable of
transmitting the fibroma virus under
laboratory conditions: Acles aegypti,
A. triseriatus, Culex pipiens and
C. quinquefasciatus. Aedes aegypti
is found only in such sites as ware¬
houses in Illinois and then only oc-

236

Transactions Illinois Academy of Science

casionally. The other species men¬
tioned are common throughout the
state. The flying range of the three
species is such that they were prob¬
ably present in both habitats during
the period of study. It is possible
that the sylvan habitat might have
had larger numbers of Aedes triseri-
atus, the treehole mosquito. Field
records show that mosquitoes did
not appear to be more numerous in
the sylvan habitat than in the cam¬
pestral. The bedbug, Cimex lec-
tularius L is found throughout the
state, but has not been recorded as
an ectoparasite of cottontails (Shope,
1959). No triatoma bugs have been
reported in central Illinois. The two
fleas which were used primarily in
the experiments of Ivilham and
Woke, (1953) Cediopsylla simplex
and Odontopsyllus midtispinosus,
are found on Illinois cottontails. The
former (the common eastern rabbit
flea) was far more abundant. It
was found in abundance on rabbits
of both habitats throughout the year.
Cottontails of central Illinois are
occasionally parasitized by the fol¬
lowing chiggers : Euschongastia per-
omysci (Ewing) , T r o ml) i c u l a
( Eutrombicula ) alfreddugesi ( On -
demans ) and Trombicula ( Neotrom -
bicida) whartoni (Ewing). The
species used unsuccessfully in trans¬
mission by Dalmat (Dalmat, 1959),
T rombicida splendens, has not been
reported from Illinois cottontails.

The continental rabbit tick, Hae-
maphysalis leporispalustris (Pack¬
ard) (Herman, 1938) is another
common ectoparasite of Illinois cot¬
tontails ; it is also the chief carrier
of Pasteurella tularensis, which has
in the past caused mortality among

cottontails of this state (Stannard
and Pietsch, 1958). Adults are
found on the cottontails largely
from March to June. Nymphs have
two peak periods, one in May and
another in September and October.
The peak for the larvae comes in
August and September, at which
time the cottontails are more heavily
parasitized with larvae and nymphs
than in the spring. Since Larson
et. al . (Dalmat, 1958) were able to
transmit rabbit papillomatosis virus
by nymphs of the rabbit tick, this
arthropod should also be considered
as a possible vector.

In correlating the experimental
findings with this investigation, it
would appear that, with the excep¬
tion of Aedes triseriatus, mosquitoes
are not the major suspects in spite
of their proven capability in the
laboratory. The experimental evi¬
dence in favor of fleas is much less
extensive ; from the abundance of
fleas on cottontails at all seasons of
the year in both habitats, it would
appear that they are not the vectors
in central Illinois. Population peaks
of the rabbit tick nymph and larva
as determined by a previous investi¬
gation in northern Illinois correlated
well with the prevalence of Shope ’s
fibroma in the study reported here.
However, few ticks were encountered
in these collections and no differ¬
ences between the two habitats in
this respect were recorded. There
is a strong possibility that more
larval ticks would be found in the
sylvan habitat if efforts were made
to find them.

The location of the lesions favors
the assumption that the vector
reaches the rabbit by way of the

Shope’ s Fibroma in Cottontails

237

ground. The sites of all except two
lesions were on the feet or legs as
shown in Table 3. One was on the
nose and one on the ear. Chigger
mites are usually found in Illinois
cottontails on, or in, the ears (Stan-
nard and Pietsch, 1958). Lesions
caused by fleas, mosquitoes and
reduviid bugs appeared on other
parts of the body in the experimental
disease. There is also the possibility
that another animal found in sylvan
habitat may be the reservoir of the
virus. Tumors similar to or even
immunologically related to Shope ’s
fibroma have been found on other
animals (Herman and Bigchoff,
1950; Herman and Reilly, 1955; and
Kilham, Herman and Fisher, 1953),
including deer, grey squirrels, fox
squirrels, woodchucks and porcu¬
pines.

Subdivision of the sylvan habitat
into dense woods and the immedi¬
ately adjoining fallow fields in Table
2, yielded differences which were not
so striking as those between the agri¬
cultural and wooded areas. Ap¬
proximately 1.8% of 279 rabbits
from the woods had fibromas while
only about 0.4% of 956 rabbits from
the uncultivated fields were affected.
This is another indication that the
wooded area might have been the
major habitat of the vector or reser¬
voir. The seasonal prevalence of
the disease in Illinois supports the
hypothesis that an arthropod vector
is responsible, since the large major¬
ity of cases were found in the fall.
There is an abrupt diminution in
the number of tick larvae and
nymphs found on rabbits after frost
(Shope, 1959). The few cases found

in early winter are readily explained
on the basis of the time required
for the regression of natural lesions,
which can be at least 10 months
(Shope, 1959). It is also possible
that under certain circumstances a
few vectors might live into the win¬
ter months.

The age distribution of the affected
rabbits in Table 3 was one adult to
8 juveniles. This is approximately
the proportion of juveniles to adults
found in the fall and winter. Ninety-
two per cent of the rabbits trapped
in these months were juveniles.

The differences between the preva¬
lence of Shope’s fibroma in cotton¬
tail rabbits on cultivated fields and
from heavily wooded habitats indi¬
cate that foci of the disease in cen¬
tral Illinois are to be found in the
wooded regions.

Summary

An investigation of Shope’s fibro¬
ma in cottontail rabbits was carried
out between 1956 and 1960 in cen¬
tral Illinois. Nearly 3,000 cotton¬
tails, largely from four adjoining
counties, were examined at monthly
intervals. A total of 1,506 cotton¬
tails from cultivated fields were
negative; in contrast, 0.72% of 1244
rabbits taken during the same period
from a heavily wooded area were
positive. A much larger proportion
of the positives were taken from
dense woods than from nearby uncul¬
tivated fields. All fibromas except
one were found on body parts usu¬
ally or frequently in contact with
the ground. All but one case were
found in the fall ; the exception was
found in early winter.

238

Transactions Illinois Academy of Science

Literature Cited

Dalmat, H. T. 1959. Arthropod transmis¬
sion of rabbit fibromatosis (Shope).
J. of Hyg., 57:1-32.

Dalmat, H. T. 1958. Arthropod trans¬
mission of rabbit papillomatosis. J. of
Exp. Med., 108:9-20.

Dalmat, H. T. and M. F. Stanton. 1958.
A comparative study of the Shope
fibroma in rabbits in relation to trans-
missibility by mosquitoes. J. of Natl.
Cancer Institute, 22:593-615.

Herman, C. M. 1938. Occurrence of
larval and nymphal stages of the rab¬
bit tick, Haemaphy salis Leporis-Palus-
tris , on wild birds from Cape Cod.
Bull, of the Brooklyn Entomological
Soc., 33(3) : 133-134.

Herman, C. M., and A. I. Bischoff. 1950.
Papilloma, Skin Tumors in Deer. Cali¬
fornia Fish and Game, 36:19-20.

Herman, C. M., L. Kilham and O. War-
bach. 1956. Incidence of Shope’s rab¬
bit fibroma in cottontails at the Patux¬
ent Research Refuge. J. of Wildlife
Management, 20: 85-89.

Herman, C. M. and J. R. Reilly. 1955.

Skin tumors on squirrels. J. of Wild¬
life Management, 19: 402-403.

Kilham, L., C. M. Herman and E. R.
Fisher. 1953. Naturally occurring
fibromas of grey squirrels related to
Shope’s rabbit fibroma. Proc. Soc.
Exp. Biol, and Med., 82:298-301.

Kilham, L. and P. Woke;. 1953. Labora¬
tory transmission of fibromas (Shope)
in cottontail rabbits by means of fleas
and mosquitoes. Proc. Soc. Exp. Biol,
and Med., 83:296-301.

Rose Lake Wildlife Experiment Sta¬
tion, Michigan. 1940-45. Dept, of Con¬
servation, Annual Reports: 2nd (1940-
41); 3rd (1941-42); 4th (1942); 5th
(1943); 6th (1944); and 7th (1945).

Shope, R. E. 1959. The roles of virus
and host in determining the host re¬
action to the fibroma-myxoma virus
complex. Genetics and Cancer, Texas,
pp. 311-323.

Stannard, L. J. and L. R. Pietsch. 1958.
Ectoparasites of the cottontail rabbit
in Lee County, northern Illinois. Nat¬
ural History Survey Publication, Ur-
bana, Illinois, Biological Notes, 38:1-
19.

CEREBROID OOLITES

ALBERT V. CAROZZI

University of Illinois, Urbana

The oolitic deposits along the
shores of the Great Salt Lake, Utah
described by Eardley (1938) do not
seem to have attracted the attention
they deserve as remarkable examples
of application of the principle of
nniformitarianism.

These sediments display several
peculiar types of oolites, the equiva¬
lent of which occur in the geological
column. Among them are the broken
and regenerated oolites described
from limestones and iron ores which
may be seen forming today along
the shoreline deposits of the Great
Salt Lake as the result of reciprocal
mechanical impacts (Carozzi, 1961).

The present paper describes and
discusses the significance of the so-
called “cerebroid oolites” also form¬
ing at the present time and frequent¬
ly reported in the Ste. Genevieve
Formation (Mississippian) of Illi¬
nois and adjacent states.

Oolitic deposits of the Great Salt
Lake, Utah. Our investigation of
the Great Salt Lake oolitic deposits
has revealed six distinct types of
oolites. These are :

1. Spherical body with pearl-smooth
surface showing in section regu¬
lar and fine concentric rings of
impure aragonite consisting of
small radial needles with inter¬
stitial clay minerals (Fig. 1,A).
These layers are developed
around nuclei of detrital miner¬
als or faecal pellets of brine
shrimps. This is the original type
of oolite as formed in the shallow

and gently agitated waters of
the Great Salt Lake mainly from
June to November (Eardley,
1938: 1327-8).

2. Spherical body with pearl-
smooth surface showing in sec¬
tion a few concentric layers of
impure aragonite coating an in¬
ternal spherical part which is
calcitic and displays an interfer¬
ence between fibro-radiated and
concentric structures (Fig. 1,B).
This is the incipient result of the
inversion to calcite and of the
recrystallization which takes
place from February to May in
correspondence with the period
of maximum inflow during which
the resulting dilution of the wa¬
ters generates conditions of little
or no precipitation of calcium
carbonate. In these oolites the
inversion and recrystallization
processes have not yet affected
the newly deposited aragonite
layers. The central portion of
calcite shows small prismatic
crystals in radial position and
usually limited to a given con¬
centric layer. The degree of re-
crystallization is rather uniform,
however some radial areas are
better crystallized than others
but their distribution is random
and different from one layer to
another. The outside shape of
the recrystallized central part is
not appreciably affected and re¬
mains spherical.

3. Spherical body with finely granu¬
lar and crystalline surface ap-

[ 239 ]

240

Transactions Illinois Academy of Science

Figure 1. Schematic segments of the six types of oolites occurring in the present
deposits of the Great Salt Lake, Utah.

A: aragonitic oolite with normal concentric rings
B: aragonite concentric rings over fibro-radiated calcite
C: calcitic oolite entirely fibro-radiated
D: aragonite concentric rings over cerebroid calcite
E: calcitic oolite entirely cerebroid
F: cerebroid calcite over fibro-radiated calcite

Cerbroicl Oolites

241

pearing in section as entirely cal-
citic and displaying an interfer¬
ence between fibro-radiated and
concentric structures (Fig. 1,C).
This is the completed stage of
the preceding type.

4. Spherical body with pearl-smooth
surface showing in section a few
concentric layers of impure arag¬
onite of variable thickness coat¬
ing an internal portion of calcite
displaying an interference be¬
tween a peculiar type of fibro-

radiated structure and the origi¬
nal concentric one. This
interference generates a fes¬
tooned or “cerebroid” outline
(Fig. 1,D) which consists of
juxtaposed convex and concave
segments corresponding to the
alternation of radial bundles of
clear calcite with darker areas
of cryptocrystalline calcite.

5. Mottled and speckled spherical
body resulting from numerous
light-colored and well-crystal-

Figure 2. Cerebroid oolites from the present deposits at the northern tip of Stans-
bury Island, Great Salt Lake, Utah. Nicols not crossed, X 88. See text for
additional explanations.

242

Transactions Illinois Academy of Science

lizecl bulging areas separated by
darker and concave zones with
cryptocrystalline structure. (Fig.
1,E) In section such an oolite
displays a festooned or cerebroid
aspect representing the complet¬
ed stage of the preceding type.
These bodies have been described
by Eardley (1938: 1380-84, pi. 8,
10 a, 12 b) under the name of
“ mottled oolites” and are the
main subject of the present pa¬
per.

6. In this last type (Fig. 1,F), the
bundles of clear calcite crystals,
instead of originating at the nu¬
cleus boundaries, start on top of
one or several concentric layers
displaying uniform fibro-radiat-
ed structure.

Description of cerebroid oolites in
the Great Salt Lake deposits. Typi¬
cal sections of cerebroid oolites show,
starting from the nucleus or from
a given concentric ring, wedge-
shaped bundles of radial crystals of
clear calcite (Fig. 2, A and B) or
occasionally single wedge-like crys¬
tals (Fig. 3,C and E). Both types
reach the external surface where
they distinctly protrude as flat-
topped or convex bulges displaying
often a well-developed fan-shaped
structure (Fig. 2,C and E).

The bundles of clear calcite fibers
are composite structures which in¬
crease irregularly in width toward
the outside of the oolite and consist
of a superposition of constricted and
expanded segments with straight or
fan-shaped boundaries (Fig. 2, C ;
Fig. 3,D). These segments always
correspond to the intersection with
a given concentric ring or a particu¬
lar set of them. In some of the more

constricted parts, the prismatic bun¬
dles may be reduced to a few fibers
of calcite or even to a single one.
The modifications of section of the
bundles express actually a variable
lateral extension of the prismatic
recrystallization which may in some
cases develop laterally far enough
to join adjacent prismatic areas
along one or several concentric rings.

Inside the bundles of clear calcite
crystals are abundant narrow and
irregular radial zones of crypto¬
crystalline calcite also closely related
to the intersection with given con¬
centric layers (Fig. 3,G). Some of
these radial zones appear limited to
the basal portion of the bundles and
wedge out toward the outside where¬
as others indent the terminal parts
of the bundles which appear as ir¬
regularly crenulated. These darker
areas contain a concentration of im¬
purities, particularly clay minerals
which also line the boundaries of
many prisms as discontinuous films.
Most of the bulging areas are irregu¬
larly convex and some of the flat-
topped ones not corresponding to
single crystal terminations may have
undergone a slight abrasion.

It is important to point out for
the understanding of the genesis
of the cerebroid oolites that in the
bundles of clear calcite fibers the
original concentric rings are wider
than in the adjacent areas and their
boundaries appear as “ ghost lines”
displaced toward the outside with
an increased curvature (Fig. 2, A;
Fig. 3, D and G).

The intermediate radial spaces be¬
tween the projecting bundles of
clear calcite consist of dark crypto-
crystalline calcite containing an ex-

Cerbroid Oolites

243

Figure 3. Cerebroid oolites from the present deposits at the northern tip of Stans-
bury Island, Great Salt Lake, Utah. Nicols not crossed, B and D :X 55,
all the others X 88.

A: concentric rings of aragonite over cerebroid calcitic core
B: broken and recoated oolite
C and E: wedge-like calcite crystals
F: tangential section
G: details of cerebroid structure

cess of impurities and clay minerals.
In these darker areas the original
concentric rings are barely visible,
they are in a depressed position,
thinner compared to those preserved
in the prismatic bundles and con¬

cave toward the outside (Fig. 2, A;
Fig. 3, A and G). The shape of the
cryptocrystalline areas is actually a
function of the adjacent bundles
being either conical with the apex
directed toward the outside of the

244

Transactions Illinois Academy of Science

oolite, or straight or irregular.

The darker zones may be occasion¬
ally interrupted by recrystallized
concentric rings when the prismatic
crystals of the bundles spread later¬
ally and eventually join. The periph¬
eral boundary of the darker areas
is not always a regularly curved line
parallel to the concave concentric
rings, but is quite irregular and cuts
across some of the thinnest concen¬
tric laminae indicating that some
solution or abrasion has taken place
(Fig. 3,G).

It is not possible to determine the
reason for the localization of the
bundles of clear calcite crystals par¬
ticularly in oolites with well-rounded
cores where they seem to be quite
regularly spaced. In oolites with
irregular cores, the bundles are defi¬
nitely located in greater proportion
on the protruding areas of the nu¬
clei which display the smallest radii
of curvature. In both cases, the
final result in cross-section is to
change the original circular outline
of the oolite into a cerebroid one
consisting of a juxtaposition of light-
colored convex areas and dark-col¬
ored concave ones.

Formation of cerebroid oolites in
the Great Salt Lake deposits. The
sections across cerebroid oolites in¬
dicate that they result from a type
of fibro-radiated recrystallization
which, instead of being uniformly
distributed in the spheroidal bodies,
has preferentially taken place along
certain radii generating wedge-like
bundles of calcite fibers.

Let us consider one layer of im¬
pure aragonite deposited by the
oolitization processes over a nucleus

Fig. 4. — Diagrams illustrating the forma¬
tion of the cerebroid structure in the
oolites of the present deposits of the
Great Salt Lake, Utah.

The aragonite layers (A, B and C) are
left blank, the bundles of prismatic cal¬
cite alternate with dark areas of cryp¬
tocrystalline calcite (stippled). See text
for additional explanations.

and inverting to calcite (Fig. 4,
stage 1 ) . The growth of the bundles
of radial crystals corresponds to an
expansion of the original layer at
these particular places expressed by

Cerbroid Oolites

245

an increased curvature and periph¬
eral bulging (Fig. 4, stage 2).

Such a process of local growth
requires an addition of calcite which
has taken place at the expense of
the adjacent cryptocrystalline areas
which contain an addition of clay
impurities expulsed by the growth of
the calcite fibers. Since the loss of
calcite predominates in the darker
areas over the addition of impuri¬
ties, a decrease of volume has taken
place in them accompanied by a
partial destruction of the original
concentric structures. This exchange
is demonstrated by the thinning of
the original concentric layers which
become concave outwards (Fig. 4,
stage 2).

The final result of this process of
segregation during which clear cal¬
cite crystals have grown along cer¬
tain radii and impurities have been
concentrated in the poorly crystal¬
lized intermediate areas, is to change
the original circular outline of the
concentric rings into a juxtaposi¬
tion of convex and concave segments
expressed in the cerebroid aspect
(Fig. 4, stage 2).

This differential recrystallization
into fibro-radiated bundles appar¬
ently takes place from February to
May. It corresponds to the time of
maximum inflow in the Great Salt
Lake during which the related dilu¬
tion of the waters allows little or
no precipitation of calcium carbon¬
ate ; solution of the smallest crystals
could even take place (Eardley,
1938: 1373). This solution would
account for some of the peripheral
irregularities of the darker crypto¬
crystalline areas described above.
Agitation and rolling around of the

oolites during the same period could
also be responsible for slight erosion
in the same poorly crystallized de¬
pressions as well as for the flat-
topped ends of some calcite bundles.

The differential fibro-radiated re¬
crystallization appears completed be¬
fore deposition of the next set of
aragonite layers during the follow¬
ing June to November period (Fig.
4, stage 3). These new layers of
impure aragonite have a tendency
to re-establish spherical outlines and
are normally thicker above the con¬
cave areas and thinner over the
bulges.

During the following winter¬
spring season, inversion to calcite
and recrystallization of these new
concentric layers take place. The
underlying prismatic bundles offer¬
ing better conditions for renewed
growth than the poorly crystallized
intermediate areas, the new groups
of calcite prisms tend to grow in
the same particular places as the
underlying ones but with a variable
width related to the properties of
the concentric layers involved. In
such a manner, the cerebroid struc¬
ture is perpetuated from one set
of layers to the overlying one (Fig.
4, stage 4).

The process just described is then
repeated after deposition of every
set of aragonite layers with possible
intermediate episodes of slight ero¬
sion or solution and new segments
are added to the composite bundles
of calcite fibers (Fig. 4, stages 5 and
6). As mentioned above, the latter
are in fact quite complex and con¬
tain numerous radial patches of
poorly crystallized material (Fig.
4, 7).

246

Transactions Illinois Academy of Science

In conclusion, cerebroid oolites in
the Great Salt Lake deposits result
from the seasonal alternation of
summer-fall periods of oolitization
with winter-spring periods of inver¬
sion to calcite and differential fibro-
radiated recrystallization. It is pos¬
sible that the inversion of aragonite
to calcite may have no genetic rela¬
tion with the cerebroid structure
which could then develop also in
oolites originally calcitic.

The rapid internal change of the
oolites from a concentric to a pre¬
dominant radial structure of any
kind is of great interest because it
explains the shape of broken and
regenerated oolites found in the
Great Salt Lake deposits as well as
in the geological column (Berg,
1944; Carozzi, 1961). In both in¬
stances oolites undergoing mechani¬
cal impacts during reworking proc¬
esses penecontemporaneous with dep¬
osition are broken in most of the
cases in clean-cut halves or quarters
of spheres indicating a stronger con¬
trol of the fibro-radiated structure
over the concentric one in the pat¬
tern of rupture.

Description of cerebroid oolites in
the Ste. Genevieve Formation (Mis-
sissippian) . To our knowledge cere¬
broid oolites have been reported for
the first time in the Fredonia Mem¬
ber of the Ste. Genevieve Formation
of Southern Illinois by Graf and
Lamar (1950: 2327, Fig. 7), and
their interpretation will be discussed
later.

The specimens described in this
paper have been collected from the
Levias Member of the Ste. Genevieve
Formation on the east wall of a road
cut on the northbound lane of In¬

terstate Route 37, SW *4 sec. 6, T.13
S, R.l E, three miles north of Don-
gola, Illinois. The particular lime¬
stone containing the cerebroid oolites
is the basal bed of the Levias Member
and corresponds to unit 7 of stop 7,
of the 25th Annual Tri-State Geo¬
logical Field Conference (Harris,
1961).

In general cerebroid oolites are
rather rare in the Ste. Genevieve
oolitic limestones and this applies
to the investigated bed in which they
represent never more than 2 to 5%
of the total number of oolites present
in a given thin section. The Missis-
sippian cerebroid oolites display all
the major characters of those de¬
scribed above from the Great Salt
Lake and their detailed description
appears superfluous. However a few
small differences may be noticed
(Fig. 5). The prismatic bundles do
not appear so well crystallized in
spite of their distinct peripheral
bulging as they are in the specimens
from the Great Salt Lake and also
they do not show accentuated con¬
strictions and expansions. This last
observation indicates that the super¬
posed concentric layers were rather
uniform in original composition and
structure.

It is interesting to mention here
that one calcitic oolite with an outer
shell of concentric rings coating a
central portion with cerebroid out¬
line has been described from the
Rogenstein of Vienenburg, Germany
(Berg, 1944: 48, Fig. 67). In this
specimen similar to our type 4 of
the Great Salt Lake deposits, the
fibro-radiated structure has clearly
affected the deposition of the last
set of concentric rings. More recent-

Cerbroid Oolites

247

Figure 5. Cerebroid oolites from the Levias Member, Ste. Genevieve Formation
(Mississippian), Dongola, Illinois. Nicols not crossed, A to C: X 22, D to
F: X 55.

ly Usclowski (1962: 167, Fig. 20)
described oolites with “Kegelstruk-
tur” from the Lower Buntsandstein
of Northern Germany which some¬
what resemble cerebroid oolites. He
assumed for their formation the ex¬
istence of several nuclei which
would have interfered during*
growth. This mechanism does not
seem to apply to the oolites dis¬
cussed in this paper.

Formation of cerebroid oolites in
the Ste. Genevieve Formation ( Mis-
sippian). In their original descrip¬
tion Graf and Lamar (1950: 2327,
Fig. 7) considered the cerebroid
oolites to result essentially from a
marginal replacement of the brown
calcite of the oolites by the inter¬
stitial clear calcite. However they
also assumed that a few broader
arcs of the oolite margins might be

248

Transactions Illinois Academy of Science

of primary origin or due to com¬
paction effects.

In the light of the observations
of Berg (1944) and of the preceding
description it is difficult to visualize
a replacement mechanism because
the cerebroid outline is not a super¬
ficial feature but only the peripheral
expression of an internal modifica¬
tion of the oolites. Indeed many
oolites display marginal irregulari¬
ties reaching a depth of 0.1 mm or
more attributable to mechanical im¬
pacts during deposition, or replace¬
ment by the interstitial cement or
compaction. However the designa¬
tion of cerebroid should be restricted
to those owing their peculiar fes¬
tooned shape to differential fibro-
radiated recrystallization which
displays a certain number of transi¬
tional terms to uniformly fibro-
radiated types as would be expected
whenever the bundles of radial cal-
cite are poorly developed.

On the basis of the evidence from
the Great Salt Lake deposits, it is
suggested that the cerebroid struc¬
ture is generated during phases of
interruption of the accretion proc¬
ess and consecpiently before any pre¬
cipitation of clear interstitial calcite.

The abundance of cerebroid oolites
(30 to 50% of the oolites in a given
thin section) in the Great Salt Lake
deposits compared to the relatively
rare occurrence (2 to 5%) in the
Mississippian oolitic rocks may be
an expression of the peculiar pres¬
ent-day physico-chemical conditions
(high salinity and temperature) in
the Great Salt Lake.

Summary

A large proportion of the arago¬
nite oolites forming at present in
the Great Salt Lake, Utah display
a peculiar type of fibro-radiated
structure generated during their sea¬
sonal inversion to calcite. The re-
crystallization instead of being uni¬
formly distributed in the spheroidal
bodies has preferentially taken place
along certain radii generating wedge¬
like bundles of clear calcite fibers
separated by intermediate areas of
dark cryptocrystalline calcite en¬
riched with clay impurities. The final
result in cross section is to change
the original circular outline of the
concentric rings into a juxtaposi¬
tion of convex and concave segments
self-perpetuating from one season to
another. The name of cerebroid
oolite is applied to these peculiar
bodies which are the modern equiva¬
lents of similar forms described in
the Ste. Genevieve Formation (Mis¬
sissippian) of Illinois and adjacent
states. It is suggested that the differ¬
ential recrystallization which gener¬
ated cerebroid oolites in the past may
not necessarily have been restricted
to oolites originally aragonitic.

Literature Cited

Berg, G. 1944. Vergleichencle Petrog-
raphie oolitischer Eisenerze. Reich-
samt f. Bodenf., Archiv fiir Lager-
stattenforschung, Heft 76, 128 pp.
Carozzi, A. V. 1961. Oolithes remaniees,
brisees et regenerees dans le Missis-
sipien des chalnes frontales, Alberta
Central, Canada. Archives des Sci¬
ences, Geneve, 14 (2): 281-296.

Graf, D. L. and J. E. Lamar. 1950.
Petrology of Fredonia oolite in South-

Cerbroid Oolites

249

ern Illinois. Am. Assoc. Petroleum
Geologists Bull., 34 (12): 2318-2336.
Eardley, A. J. 1938. Sediments of the
Great Salt Lake, Utah. Am. Assoc.
Petroleum Geologists Bull., 22 (10):
1305-1411.

Harris, S. E., Jr. 1961. 25th Annual
Tri-State Geological Field Conference

Guidebook. Southern Illinois Univer¬
sity, Carbondale, Illinois, mimeo¬
graphed, 28 pp.

Usdowski, H. E. 1962. Die Entstehung
der kalkoolithischen Fazies des nord-
deutsclien Unteren Buntsandsteins.
Beitrage zur Min. und Petr., 8 (3):
141-179.

CONSTITUTION AND BY-LAWS OF THE
ILLINOIS STATE ACADEMY OF SCIENCE

(As of April 27, 1962)

CONSTITUTION

Article I. Name

This Society shall be known as the Illi¬
nois State Academy of Science.

Article II. Objects

1. The objects of the Academy shall be
the promotion of scientific research, the
diffusion of scientific knowledge and sci¬
entific spirit, and the unification of the
science interests of the state.

Article III. Membership and Dues

1. Any person of good character, pro¬
posed by two members of the Academy,
and recommended bv the chairman of
the Membership Committee may be
elected to any class of membership in the
Academy by a majority vote of the Coun¬
cil upon payment of scheduled dues. Any
organization approved by the Chairman
of the Sustaining Membership Commit¬
tee may become a sustaining member or
a patron upon payment of appropriate
dues. Individuals who attended the or¬
ganization meeting of the Academy in
1908, paid dues for that year, and signed
the original constitution of the Academy
are designated charter members.

2. Regular individual members in
good standing shall have the privilege
of voting at the annual meeting, holding
office, offering papers for presentation
at meetings subject to the approval of
the appropriate Section Chairman and
with right of appeal to the Council, hav¬
ing papers published in the TRANSAC¬
TIONS if accepted by the Board of Edi¬
tors, and receiving one copy of the cur¬
rent TRANSACTIONS of the Academy.
No member in arrears shall receive the
TRANSACTIONS for any year for which
he is or remains in arrears.

3. Any college or high school student
who has not received a bachelor’s degree
and who is certified as such by an ad¬
visor or instructor may become a Stu¬
dent Member with all rights and privi¬
leges of membership except that of vot¬
ing and holding office.

4. The dues shall be as follows:

Regular member, annual $ 5.00

Student member, annual. 2.00

Life member,

single payment . 100.00

Sustaining member,
annual . . . . 10.00-$49.00

Patron member,

annual . 50.00-more.

5. Proceeds from life memberships
shall constitute an inviolate permanent
investment fund from which only the
annual income may be used. Proceeds
from sustaining and patron member¬
ships shall be used to finance the activi¬
ties of the Junior Academy and to ad¬
vance its program as shall be determined
by the Council.

6. The fiscal year of the Academy
shall be from January 1 through Decem¬
ber 31. Members who fail to pay dues
for any fiscal year shall be considered
in arrears for that year, and unless they
pay dues by the succeeding December 31
they shall be dropped from membership
as of that date.

Article IV. Officers

1. A President, a First Vice-President,
a Secretary, and a Treasurer shall be
elected by the members of the Academy
at its annual meeting, shall hold office
for one year or until their successors are
elected and take office, and shall perform
the duties usually pertaining to their re¬
spective offices. The First Vice-President
shall also assist the Secretary in formu¬
lating the program for the annual meet¬
ing.

2. If between annual meetings of the
Academy the offices of both President
and First-Vice-President shall become va¬
cant, the last available Past President
shall act as President until the next an¬
nual meeting. Interim vacancies in the
offices of Secretary and Treasurer shall
be filled by pro-tem officers elected by
the Council.

3. A Second Vice-President, who shall
preferably be a resident of the commu¬
nity in which the next annual meeting is
to be held, shall be elected by the Coun¬
cil each year and shall serve as Chair¬
man of Local Arrangements for that
meeting. When possible, a Second Vice-

Constitution and By-Laws

251

President Elect, who shall preferably be
a resident of the community in which
the second next annual meeting is to be
held, shall also be elected by the Council,
subject to confirming election as Sec¬
ond Vice-President by the next succeed¬
ing Council.

4. The Director (or Acting Director)
of the State Museum Division of the De¬
partment of Registration and Education
of the State of Illinois, or a member of
the State Museum Staff designated by
the Director to represent him on occa¬
sion of his temporary absence, shall be
the Librarian of the Academy and as
such shall have charge of all books, col¬
lections, and similar property of the
Academy, shall serve as archivist of all
official records and documents of the
Academy, and shall have charge of the
distribution, sale and exchange of publi¬
cations of the Academy, in accordance
with policies determined by the Council.

Article V. Council

1. The Council shall consist of the
President, First Vice-President, Second
Vice-President, Secretary, Treasurer, Li¬
brarian, General Chairman of the Junior
Academy, the immediate Past President,
the immediate Past Secretary, the imme¬
diate Past Treasurer, each for a term of
one year, and four Councilors-at-large.
These last shall be elected for four-year
terms, only one being elected each year,
except that the first year this provision
is put into effect the terms of the three
incumbent Councilors-at-large shall each
be extended one year and a fourth Coun¬
cilor-at-large shall be elected for a four-
year term.

2. Except as otherwise herein pro¬
vided, the members of the Council shall
be elected annually by the members of
the Academy at its annual meeting and
interim vacancies in these positions shall
be filled by pro-tem elections by the
Council.

3. The Council shall manage the af¬
fairs of the Academy between annual
meetings of the Academy and shall be re¬
sponsible for the general planning, poli¬
cies, program, and arrangements for an¬
nual meetings. It shall meet on call by
the President ordinarily four times each
year — once as soon as convenient after
the annual meeting of the Academy, once
each normally in November and Febru¬
ary, and once just prior to the next an¬
nual meeting of the Academy.

Article VI. Technical Personnel

1. A Publicity Adviser shall be elected
each year by the Council and he shall be
responsible for appropriate publicity
concerning all meetings and affairs of
the Academy.

Article VII. Meetings of the Academy

1. The regular annual meeting of the
Academy shall be held at such time and
place as the Council may designate.
Meeting places shall be determined at
least two years in advance.

2. Special meetings of the Academy
shall be called by the President upon
written request by twenty members.

3. No meeting of the Academy shall
be held without 30 days previous written
notice by the Secretary to all members.

Article VIII. Junior Academy

1. In order to foster science among
youth, the Academy is committed to
sponsorship of an organization of junior
high-school and high-school students in¬
terested in science and directed by quali¬
fied teachers in the schools of Illinois,
which organization is termed the Junior
Academy.

2. The affairs of the Junior Academy
shall be conducted in accordance with
policies and principles determined by the
Council of the Academy.

3. The directing head of the Junior
Academy shall be a General Chairman
who shall be elected by the members of
the Academy at its annual meeting. A
General Chairman Elect shall also be so
elected. If between annual meetings the
position of General Chairman becomes
vacant, the General Chairman Elect shall
assume the position, and in this event or
if for any other reason the position of
General Chairman Elect becomes vacant,
it shall be filled by pro-tem election by
the Council.

4. No expenditures may be made and
no bills or other liabilities may be in¬
curred on behalf of the Junior Academy
without prior approval of the Council.

Article IX. Publications

1. The regular publication of the
Academy shall be the Transactions of
the Illinois State Academy of Science.

2. Other publications may be author¬
ized by the Council.

3. Papers presented at annual meet¬
ings of the Academy by non-members at
the invitation or with the approval of
the Council shall be eligible also for pub¬
lication by the Academy.

252

Transactions Illinois Academy of Science

Article X. Business and Finance

1. Life membership payments shall
constitute an inviolate endowment fund
which shall be invested in guaranteed se¬
curities and of which only the income, in
lieu of annual dues of the life members,
shall be used by the Academy.

2. A second investment fund consist¬
ing of donations, bequests not otherwise
restricted, allotments transferred by the
Council from surplus reserve in the
treasury, and similar sources shall be
maintained as an emergency reserve, and
only the income from this fund shall nor¬
mally be available for the expenses of
the Academy.

3. No expenditures may be made and
no bills or other liabilities may be in¬
curred by any officer, individual, or com¬
mittee on behalf of the Academy, unless
they are included in the budget, without
prior approval of the Council.

4. At its first meeting each Council
shall approve a budget of expenditures
that shall not exceed the total of the cur¬
rent balance and the anticipated receipts,
which budget may be modified by the
Council at its subsequent meetings. The
Treasurer shall not make payments for
approved expenditures in excess of the
covering item in the approved budget
without express approval of the Presi¬
dent and the Secretary. Such excesses
must be covered by modifications of the
budget at the next Council meeting.

5. The Secretary and Editor shall
each receive an honorarium in amounts
to be determined by the Council. These
honoraria shall be paid by the Treasurer
annually in March.

6. The President, the Secretary, and
the Treasurer shall be reimbursed for
their expenses while attending Council
meetings and annual meetings. Other
members of the Council may be reim¬
bursed for their expenses while attend¬
ing Council meetings other than those
held in connection with annual meetings.

Article XI. Affiliations

The Academy may enter into such re¬
lations or affiliation with other organi¬
zations of appropriate character as may
be recommended by the Council and ap¬
proved by the members of the Academy
at its annual meeting.

Article XII. Amendments

Proposed amendments to this consti¬
tution shall become effective upon ap¬
proval by three-fourths of the members

present and voting at the annual busi¬
ness meeting of the Academy, provided
that notice of the proposed changes has
been sent by the Secretary to all mem¬
bers of the Academy at least twenty days
before such meeting.

BY-LAWS OF THE
ILLINOIS STATE ACADEMY
OF SCIENCE

I. Order of Business

1. The business meetings of the Acad¬
emy and the meetings of the Council
shall be conducted in accordance with
Robert’s Rules of Order.

2. Fifteen members shall constitute a
quorum of the Academy and six coun¬
cillors shall constitute a quorum of the
Council.

II. Committees and Delegates

1. The standing committees of the
Academy shall be Affiliations, Animal
Experimentation in Research, Budget,
Conservation, Archaeological and His¬
torical Sites, Legislation and Finance,
Local Conventions, Membership, Re¬
search Grants, Science Talent, Sustain¬
ing Memberships, and Teacher Training.
Any of these committees may be abol¬
ished and additional ones may be estab¬
lished upon recommendation of the
Council at any annual meeting of the
Academy.

2. The chairmen and members of the
standing committees shall be elected by
the members of the Academy at its an¬
nual meeting. Interim vacancies on the
committees shall be filled by the Coun¬
cil. who shall have authority also to elect
additional members.

3. The Local Conventions Committee
shall consist of the last three available
Second Vice-Presidents, the Secretary of
the Academy ex-officio, and a representa¬
tive of the Junior Academy.

4. There shall also be committees on
Audit, Nominations, and Resolutions of
which the members shall be appointed by
the President not later than the Febru¬
ary Council meeting.

5. The duties of the committees shall
be as defined by the Council, except as
hereafter provided.

6. The budget committee shall submit
at the last Council meeting for its con¬
sideration and for review at the annual
meeting of the Academy a budget of an¬
ticipated income and recommended ex¬
penditures for the following year. The

Constitution and By-Laws

253

budget shall be approved by the succeed¬
ing Council at its first meeting.

7. The Local Conventions Committee
shall prepare and maintain up-to-date a
“hand book” for the guidance of Second
Vice-Presidents in discharging their du¬
ties as Chairmen of Local Arrangements
and shall further advise such officers
when requested.

8. The membership of the Audit,
Nominations, and Resolutions commit¬
tees shall be announced to the members
of the Academy by the President or the
Secretary not less than 30 days prior to
the annual meeting.

9. The Nominations Committee shall
present at the annual business meeting
of the Academy a list of candidates for
all offices and committees which shall be
filled by election by members of the
Academy.

10. The Resolutions Committee shall
present at the annual business meeting
of the Academy all resolutions for con¬
sideration by the members of the Acad¬
emy, with recommendations for or
against approval. Members of the Acad¬
emy who wish to sponsor resolutions
shall submit them to the Resolutions
Committee for consideration not less
than 24 hours and preferably longer be¬
fore the annual meeting. No resolution
not submitted to the Committee may be
presented at the meeting of the Academy.
The Committee shall draft any appropri¬
ate resolutions as directed by the officers
of the Academy or Council, may redraft
any resolution submitted by any member
or members to place it in more accept¬
able form, and may draft resolutions to
incorporate suggestions offered by mem¬
bers of the Academy.

11. One or more delegates to the
American Association for the Advance¬
ment of Science or to any other organi¬
zation with which the Academy is af¬
filiated, when such delegates are re¬
quired, shall be elected by the Council
for such a term or terms as the Council
may designate.

III. Technical Sections

1. Technical papers presented at the
annual meeting of the Academy will be
distributed among Sections representing
various fields of science as follows:
Aquatic Biology; Anthropology; Botany;
Chemistry; Geography; Geology; Me¬
teorology and Climatology; Microbiol¬
ogy; Physics; Science Teaching; and Zo¬
ology.

2. Any of these sections may be abol¬
ished, divided, or combined by the Coun¬
cil as seems desirable, and other sec¬
tions may be established by the Council
upon request of ten active members.

3. Members of the Academy shall in¬
dicate in which section or sections they
are particularly interested.

4. The members present at any sec¬
tion meeting during the annual meeting
of the Academy shall constitute a quo¬
rum of the section. They shall elect a
chairman for the ensuing year, who shall
be responsible for assembling a program
for the section for the next meeting. In
the event that an elected chairman can
not serve, a substitute shall be elected by
the Council.

5. No paper shall be entitled to a
place on the Section programs unless the
manuscript or an abstract of the same
shall have been previously delivered to
the Secretary in accordance with instruc¬
tions. No paper shall be presented at any
Section meeting by any person other
than the author, except with approval
of a majority of the members present at
such meeting. No paper shall be accepted
for publication unless the author or a
co-author is a member of the Academy
or an approved applicant for member¬
ship or unless it has been presented at
the invitation of or with the approval of
the Council, nor shall any paper publish¬
ed wholly or in large part elsewhere be
acceptable for publication.

IV. Publications

1. The publications of the Academy
shall be under the supervision of a Com¬
mittee on Publications, which shall con¬
sist of an Editor and a Board of Editors.

2. The Editor shall be appointed an¬
nually by the Council at its first meet¬
ing. He shall be chairman of the Com¬
mittee on Publications and Technical
Editor.

3. The Board of Editors shall consist
of eight Associate Editors, of which one
shall be for the Junior Academy, one
shall be for News and Notes, and six
shall represent various areas of science.
The Associate Editors shall be appointed
annually by the Council at its first meet¬
ing.

4. Sometime between May 1 and Sep¬
tember 1 of even-numbered years the Edi¬
tor and/or the members of the Board of

254

Transactions Illinois Academy of Science

Editors shall confer with the Director of
the Illinois State Museum concerning
the amount of appropriations that should
be requested for publication of the
Transactions during the next State Bien¬
nium.

V. Suspension or Amendment of
By-Laws

1. These by-laws may be suspended or
amended at any annual meeting of the
Academy by a majority vote of the mem¬
bers present.

PREPARATION OF MANUSCRIPTS FOR
THE TRANSACTIONS

For publication in the Transactions , articles must present significant
material that has not been published elsewhere. Review articles are ex¬
cepted from this provision, as are brief quotations necessary to consider
new material or varying concepts. All manuscripts must be typewritten,
double spaced, with at least one-inch margins. The original copy, not the
carbon copy, is to be submitted.

Titles should be brief and informative. The address or institutional
connection of the author appears just below the author’s name. Subtitles
or center headings should be used; ordinarily one uses substitles such as
Introduction, Acknowledgments, Materials, Methods, Results, Discussion,
Summary, and Literature Cited. All papers should have a summary.

No footnotes are to be used.

Doe, John H. 1951. The life cycle of a land snail. Conchol., 26(3):
21-32, 2 tables, 3 figs.

Doe, John H. 1951. Mineralogy of Lower Tertiary deposits. New
York, McGraw-Hill Book Co., iv -(- 396 pp.

Photographs should be hard, glossy prints of good contrast. Graphs,
maps and other figures reproduce best when prepared for at least one-half
reduction; lettering, numerals, etc. on all figures in a manuscript should,
be worked out to proper size for such reduction. Line widths, letter size
etc. should be uniform from figure to figure within a published paper.
Figures should be drawn on good quality white paper or on drawing
board. Use only India ink. Use a lettering device (Leroy or Wrico) for
numerals and words; do not print “free-hand.”

Authors will receive galley proofs; these should be read carefully and
checked against the original manuscript. Reprints may be ordered at
the time galley proofs are returned to the Editor.

Wesley J. Birge,

University of Minnesota, Morris
Morris, Minn.

JUutdof<£tmo&t/

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