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New York
State College of Agriculture
At Cornell University
Ithaca, N.Y.
» Library
Cornell University Library
QL 751.A41
TA
3 1924 002 894 487
wi
mann
ORIGINAL FOREST AND PRAIRIE AREA IN ILLINOIS
(AFTER BRENDEL AND BARROWS )
BULLETIN
OF THE
ILLINOIS STATE LABORATORY
oF
NATURAL HISTORY
URBANA, ILLINOIS, U. S. A.
STEPHEN A. FORBES, Pu.D., L.L.D.,
DIRECTOR
Vou. XI. SEPTEMBER, 1915 ARTICLE IT.
AN ECOLOGICAL STUDY OF PRAIRIE AND FOREST
INVERTEBRATES
BY
CHARLES C. ADAMs, Pu.D.
-—r &
CONTENTS
Pace
33
35
35
33
40-56
40
41
43
44
48
49
52
52
35
Deseription of 56-66
L 36
2, 37
3.
jation, i 59
4. Lowland or ““second bottom, ”” red oak-elm-sugar maple wood-
land association, Station IV, ¢..........-.-.....--..eeeeees 2
5. Supplementary collections from the Bates woods, Station IV.... 6
6. Small temporary stream in the south ravine, Station IV, d...... ci
General characteristics of the gross environment. ................. 66-102
L Topography and soils of the State... - 66
2. Climatic conditions ................. 67
3. Climatic centers of imflmemee .......-. 0... ee ee eens 9
4, Relative humidity and evaporating power of the air............ val
5. Temperature relations in the opem and im forests.............. 83
6. Soil moisture and its relation to vegetation. .............--...- 86
7 Ventilation of land habitats......... 2.2.0.2... 0. eee ee 8B
8. The tree trumk as a habitat..... 2.2.2... cee eens 91
9. Prairie and forest vegetation and animal life................. 91
10. Sourees and réle of water used by prairie and forest animals. . . . 98
Animal associations of the prairie and the forest. ..-............... 0.000055 102-158
TL. Introduction ....... 2.2.2... 22.20 eee eee ee ee een eee ene eenees 102
IL The prairie association... ........ 2... 2... 2-0 cee ee ees 103
L Swamp Irie association. ...... 2.2... eee eee eee eee eens 103
2. The cottonwood commumity....... 2... 2... eee eee eee 105
3. Swamp-grass association... .... 2.2... 2.2... eee ee eee eee 107
4. Low prairie association. ... ~~... 2.2.0.2... ee eee eee 108
5. Upland prairie association. .................. 0.2... e eee eee 109
6. The Sokdago commumity............-..-- +... eee eee eens 109
7. Dry prairie grass association. ...............--......66- 111
8 A muilkweed commumity...................- 12
Tit. Relation of prairie animals to their environment, 113
1. The black soil prairie community... ...... 1144
2. The prairie vegetation commumity........................5-- ni7
4. Interrelations within the prairie association ng
ITV. The forest associations.........ccsccsse cece vere er seen sree seeeeee 122
Ve Pntrodwetiony, sé cascdtoves anya oeeststaverdisis «chiding s six ate areutiaials ia Weta are 122
2. Dry upland (Quercus and Carya) forest association............ 124
3. Artificial glade community in lowland forest.........-.....0+5 125
4, Humid lowland (hard maple and red oak) forest association... . 126
5. Animal association of a temporary stream.......-..++eee-eeee 127
V. Relation of the deciduous forest invertebrates to their environment.... 128
1. Forest soil community. ........ ccc cece cece esac nace rcersces 129
2. The forest fungus community...........cceeeee ee ee cee eeees 135
3. The forest undergrowth community............ceeeeeeeeeeaee 138
4, The forest crown community..........ccceeeececeeeee ieinenes 139
5. The tree-trunk community............ OE LISS RINNE TG 142
6. The decaying wood community......... 0c cece ee eee eeeeeeee 148
7. Interrelations within the forest association.................5+ - 157
Eeologically annotated list :— :
I, Prairie invertebratesic: ccdeciieecdse code etecwa needs eee ee eerie ee 158-201
Il. Forest invertebrates. ........ ccc esc e esc e eee eee eeee suse eestteees 201-238
BUDO BPA PY «ces: :cosicack. sx evenanie des “shi danny wien des gnas re lca ogee ae “td oc vec Nora ara ova tahesens 239-264
Articie Il—An Ecological Study of Prairie and Forest Inverte-
brates. By Cuaries C. Apams, Pu.D.
INTRODUCTORY
In four generations a true wilderness has been transformed into
the present prosperous State of Illinois. This transformation has been
so complete that in many parts of the state nearly all of the plant and
animal life of the original prairie and forest has been completely ex-
terminated. Between the degree of change which has taken place in
any given area and the suitability of that area for agriculture there has
been an almost direct relation. Fortunately, however, for the preser-
vation of prairie and forest animals, the state is not homogeneous,
some areas being too hilly, rocky, or sandy for prosperous agriculture.
The character and mode of transformation which has taken place
in the past is instructive in several particulars because it serves to
guide our anticipations as to the future of our fauna. The forested
southern part of the state (see frontispiece) was first invaded by trap-
pers and hunters, who began the extermination of the larger animals.
These invaders were in turn followed by others who, with the round
of the season, were hunters or farmers, and continued this exterminat-
ing process, particularly in the clearings, which began to replace the
forest. These pioneers, men of little wealth, possessed a combination
of mental and economic habits which was the result of life in a for-
ested country, and naturally they’settled in those places most like their
former homes—within the forest or near the forest margin. From
these settlements they looked out upon the prairies as vast wastes to
be dreaded and avoided. As a result of this attitude toward the prai-
ries, it required some time, even a new generation, some economic
pressure, and a change of habits before the prairies were settled. Mean-
while the northern part of the state was yet a wilderness; but, through
the influence of the Great Lakes, as a route of communication with
the populous East, a rapid invasion of settlers set in from that direc-
tion. Though these settlers also came from a wooded country, they
were more wealthy, settled upon a very fertile soil which was favorably
located with regard to eastward communication, and they therefore
progressed more rapidly than the less favored, more isolated southern
invaders on the poorer soil; consequently they spread from the forest
34
to the prairie more rapidly than did the settlers in the South. ‘There
thus developed two active centers of influence, each of which trans-
formed the primeval conditions in the same manner and in the same
direction toward an environment suitable for man.
The forests and the upland prairie were first changed. ‘Then the
fertile wet prairie was drained, so that today it has largely become
either the hilly and rocky areas that survive as forests or the low
periodically flooded tracts, and the undesirable sand areas which simi-
larly preserve patches of sand prairie. All the changes are more
rapid and complete upon fertile soil than upon the poorer soils in the
southern part of the state.
Such considerations as these will aid one in estimating the probable
rate of future changes in different parts of the state, and will serve to
show in what parts there is urgent need of local studies if ecological
records are to be made before extinction of some forms is complete.
A study has been made with the idea of reporting upon represen-
tative patches of prairie and forest in a manner which would aid others
in making similar local studies, and would at the same time preserve
some records of the present condition of the prairie and forest. When
this work was planned, we had no general or comprehensive discussion
of the conditions of life upon the prairie and in the forest. For this
reason a general summary of these conditions and a sketch of the gen-
eral principles involved are given, so that the reader may gain some
conception of the relation of the local.problems to those of a broader
and more general character.
A section for this report was prepared giving general directions
for making such local studies, but later it was decided to publish this
‘ separately, in somewhat extended form, as a “Guide to the Study of
Animal Ecology.”* ‘This volume should be regarded as intimately re-
lated to this paper, and this report should at the same time be consid-
ered as a concrete example of the procedure suggested in that “Guide”
for ecological surveys. It will be observed that the study of the
Charleston area here referred to has been conducted in much the same
way as was my cooperative study of Isle Royale, Lake Superior, en-
titled “An Ecological Survey of Isle Royale, Lake Superior” (’09),
although certain aspects have been elaborated here which, for lack of
time, were not treated there. The time devoted to the study of the
Charleston area was also limited, but in the preparation of the report
upon it use has been made of many years’ experience and a general
knowledge of the prairie and forest. Without such a background
*The Macmillan Co. 1913.
35
much greater caution would have been necessary in discussing many
phases of the problem.
ACKNOWLEDGMENTS
The study of the Charleston area was carried, out with the coop-
eration of the Illinois State Laboratory of Natural History, through
its director, Prof. Stephen A. Forbes, and with the further coopera-
tion of Professors E. N. Transeau and T. L. Hankinson, of the East-
ern Illinois State Normal School, located at Charleston. Personally I
am indebted to Professor Forbes for the opportunity of taking part in
this study as the State Laboratory representative, and for the aid he
has given in the illustration of the report. To Professor Transeau I
am particularly indebted for the plant determinations, for lists of the
plants, and for evaporation data. To Professor Hankinson I am
under especial obligation for many specimens, which materially added
to my lists, and for a large number of photographs. I am indebted
likewise to my associates in this study for their hearty cooperation
throughout the progress of the work.
For the determination of entomological specimens I am indebted
primarily to Mr. C. A. Hart, Systematic Entomologist of the State
Laboratory of Natural History, who named most of the insects col-
lected. For the names of certain flies I am indebted to Mr. J. R. Mal-
loch, of this Laboratory. Others who have determined specimens are
as follows: N. Banks (Phalangiida), J. H. Emerton (spiders), R. V.
Chamberlain (myriapods), F. C. Baker (Mollusca), Dr. W. T. M.
Forbes (lepidopterous larve), Dr. M. C. Tanquary (ants), Dr. M. T.
Cook (plant galls), J. J. Davis (Aphidide), and Dr. A. E. Ortmann
(crawfishes collected by T. L. Hankinson). I am indebted to the U.
S. Geological Survey for photographs. Acknowledgments for illus-
trations are made under text figures and in explanations of plates.
GENERAL DESCRIPTION OF THE REGION AND LOCATION
OF THE ECOLOGICAL STATIONS
I, GeneraL DESCRIPTION OF THE REGION
The town of Charleston, Coles county, Illinois, in the vicinity of
which these ecologic studies were made, is situated on the Shelbyville
moraine which bounds the southern extension of the older Wisconsin
ice-sheet. ‘T'o the south of this moraine lie the poorer soils which char-
acterize so much of southern IIlinois; to the north, upon the older Wis-
consin drift, are some of the most productive soils found in the upper
36
Mississippi Valley. The economic, sociologic, political, and historical
significance of the difference in the soils of these regions is funda-
mental to any adequate understandng of man’s response to his ecolog-
ical environment within this area. Some of the results of this differ-
ence have long been known, but it is only in recent years that their
general bearing has been adequately interpreted in terms of the en-
vironment. Hubbard (’04) was the first, I believe, to show the sig-
nificance of this difference in soils and its influence upon local eco-
nomic problems. That such an important influence should affect one
animal (man) and not others seems very doubtful, and yet in only one
other case do we know that the lower animals respond to this ecologic
influence. Forbes (’07b) has shown that certain kinds of fish found
in streams on the fertile soils are wanting in streams on the poorer
soil. To what degree the land fauna and the native vegetation respond
to this distinction is not known, as this subject has not been investi-
gated except agriculturally. Here, then, is a factor in the physical
surroundings which should be reckoned with in any comprehensive
study of the biotiq environment. In this portion of the state, on ac-
count of the differences in the soil, the physical environment is prob-
ably more favorable to certain organisms and less. favorable to others,
‘and consequently, to a certain degree, the environment selects, or fa-
vors, some organisms. Through their activities and through other
agencies of dispersal, the animals along the borders between the two
soil types transgress these boundaries, and are therefore forced to
respond to the new conditions and to adjust themselves, if possible.
But the soil is not the only environmental influence which has pro-
duced an unstable zone or tension line in this area. A second factor is
the difference in the vegetation—the difference between the forest and
the prairie. In all probability, Coles county was at one time all prairie,
but the Kaskaskia and Embarras rivers, as they cut their valleys
through the moraine and developed their bottoms, have led forests
within the morainic border from farther south. The forests about
Charleston have extended from the Wabash River bottoms. On account
of the southerly flow of the Embarras through this county, the forest
and prairie tension line is about at right angles to that produced by the
differences in the soil. The forests have tended to spread east and west
from the streams and to encroach upon the prairie, and thus to restrict
its area more and more. The fundamental significance of the tension
between the forest and the prairie has long been known within the
state. It influenced its economic, social, political, and historic develop-
ment as much as any other single factor during its early settlement.
And just as Hubbard (’04) has shown the influence of soil upon man
37
within the state, so also has Barrows (’10) shown the influence of the
forests and prairie upon the state’s development. While the influencé
of the soil upon the animal life of the state is not so well known or es-
tablished, the influence of prairie and forest upon the animals is univer-
sally recognized, even though the subject has been given relatively
little study by naturalists.
A third leading agency is the influence of man, who has trans-
formed the prairie and forest to make his own habitat. There are thus
recognized in the Charleston region three primary environmental in-
fluences: first, the relative fertility of the soil (this depending on the
geological history) ; second, the kind of vegetable covering, whether
prairie or forest (this probably depending largely on climatic condi-
tions) ; and third, the agency of man. The general background of the
Charleston region, then, ecologically considered, depends on the com-
bined influence of five primary and secondary agencies, four of which
we may call natural and one artificial. All these are different in kind
and so independent that they tend toward different equilibria or dif-
ferent systems of unity. Two of these are due to differences in the
soil, two others to the character of the vegetation (whether prairie or
forest), and the fifth, or artificial one, is due to man. Though the
present report does not undertake to include all the problems centered
here, as any complete study would, it is desirable to see the relation of
our special study to the general problems of the region as a whole.
The undulating plain about Charleston, formed as a terminal mo-
raine, is broken along the small streams by ravines, which have cut a
few hundred feet below the general level of the region as they ap-
proached the larger drainage lines. The main drainage feature is the
Embarras River, which flows southwest about two to three miles east
of Charleston, in a narrow valley partly cut in rock. The wooded
areas are mainly near the streams; the remainder of the area is under
intensive cultivation.
During the preliminary examination of the region, which was made
to aid in selecting representative areag for study, it soon became evi-
dent that the only samples of prairie which could give any adequate
idea of the original conditions were those found along the different
railway rights-of-way. Other situations, vastly inferior to these and
yet a valuable aid in the determination of the original boundaries of
the prairies, were the small patches or strips along the country roads.
Most of the patches of prairie along the railway tracks represent the
“black soil” type of prairie, which is extensively developed in this part
of the state upon the “brown silt loam” soil” (see Hopkins and Pettit,
’08 : 224-231). Much of the region studied was originally wet prairie
38
(which, tae: Since Teen: drained), but some of the higher ground,
formed by the undulation of the surface and surrounded by the black
soil, is lighter im color and is well draimed. ‘Thus im the black soil areas
there are both wet and well-drained tracts, and corresponding differ-
ences im the habitats.
The originally wooded and the present wooded areas east of
Charleston, in the vicinity of the Embarras River, are in a region quite
different from the prairie both im topography amd im soil, Here the re-
Sie Zé tach mote pronounced, fin arciend of both the piticimity of the
river and the greater development of the draimage limes, which have cut
a few hundred feet below the general level of the country. ‘The tribu-
tary valleys and ravines are mumerous and steep-sided, and im general
are wooded, the density varying with the amount of clearimg done.
Most of the soil of the wooded areas and along the bluffs is distinctly
im color tham that of the black soil prairie, amd is
“gray silt loam ” (Hopkins and Pettit, 08 : 238-242), though along the
and the river bottom the soils are mixed im character.
IL. Tar Ecotoacan Sraroxs
In the study of am area or am amimal association of any comsidera-
ble size two methods are available. Onme is to examine as much of the
area as is possible and secure data from a very wide range of condi-
tions. ‘This method is useful in obtaiming the general or broad featmres
of a region or am association, though to a corresponding degree it must
ignore local imfimences amd details, and by it most of the previous stud-
ies mpon prairie animals hawe been made. It seemed, therefore, that m
the present study a somewhat more imtensiwe method was desirable,
particularly im view of the fact that the extinction of prairie and for-
est is rapidly progressing. The method followed was to examime 2
large area im order to select a represemtative sample, and wpom the
basis of this sample to make as intensive a study as time and circum-
stamoes would permit. ‘Tis method has the advantage of making it
possible to preserve at least some record of the local details; amd at the
same time, to the depree that the selected area is a tre sample, it also
gives the results a much wider applicatiom.
The prairie samples examined were all along the rights-of-way,
amd the forest was a secomd-growth woods om the bottoms and bluff
of the Eimbarras River, om a farm belonging, at that time, to Mir. J. 1
Bates. Practically all of the observations here reported mpom were
made during Ampust, 1910. The forest is a modified ome, but tt ap-
pears to hawe been cut over so gradually that its comtimmity as a forest
habitat was mot completely imternrmpted, although the cuttimg has prob-
39
ably seriously influenced many animals, particularly those which fre-
quent mature forests, abounding im dead and dying trees and with an
abundance of logs upon the ground im all stages of decay. Such con-
ditions are the cumulative product of a fully mature climax forest. Of
course the conditions have also been influenced by the extinction, or
eee of the original vertebrate population of the
corest.
The different localities or regions examined are, for brevity and
Precision, indicated by Roman mumerals ; the particular minor condi-
tions, situations, or habitats, by italic letters. An effort has been made
to indicate the location of the place studied with enough precision to
enable students to re-examine the habitats at amy future time (PL 1).
Se a ee eee eee meas oe ae ae
ing the places studied. Had similar photographic records been
made fifty years ago, they would have been of much value and imter-
est to us im this study, im much the same way as fifty years hence this
report will form a part of the very limited record of the conditions
found at the present time.
List of Ecological Stations, Charleston, Minois, August, 1910
Station L Prairie along the right-of-way of the Toledo, St. Louis and
Western, or ““Clover Leaf’ B. B., between one and two miles north
of Charleston: Section 2, Township 12 N., Range 9 B., amd 8. 35,
T.13N, B.9 EB. (PLT)
a Cord or Slough Grass (Sporting) and Wild Bye (Elymus) Asso-
ciation. At mile-post marked ““Toledo 318 miles and St. Louis 133
miles’: S. 2, T. 12 N., B. 9 BE.
b. Couch Grass (Agropyrom smithii) Association. The distance of
two telegraph poles north of Station I, «, and west of the railway
truck: $2, T. 12 BE. B.9 EB.
ce. Wild Bye (Elymus) Association. East and north of the “Yard
Limits” sigm: $2, T. 12 NL, B.9 B. (PL UO, Fig. 1.)
dé Swamp Milkweed (Ascdlepias imcoraata) Association. North of
first exst-and-west erosroad north of Charleston; east of railway
track: &. 35, T. 13 N., B.9 B. A wet area. (PL Ii, Fig. 2; Pl. Til,
Fig. 1.)
€. Coneflower (Lepachys pinnata) and Bosin-weed (Silphium tere-
binthinacewm) Association. Just north of the preeedimg Station ;
east of raihway tradk: 8.35, T. 13 N, B.9 BE. (PL V.)
f- Couch Grass (Agropyrom smithii) Association. West of railway
track: $35, T.13 N., B.9 EB. Moist area.
GQ Prairie Grass (Andropogon furcatus and A. virgimicus and Spo-
robolus eryptendrus) Association, bordered by Swamp Milkweed
(Asdepias imewrnata) and Mountain Mint (Pycmanthemum flex-
40
uosum). This formed the north boundary of the area studied:
S. 35, T.13 N., R.9 E. (PI. III, Fig. 2; Pl. IV, Fig. 1 and 2.)
Station II. Prairie area west of Loxa, Illinois. Right-of-way along the
Cleveland, Cincinnati, Chicago and St. Louis, or ‘‘Big Four,”
R. R.: Sections 10 and 11, Township 12 N., Range 8 E.
a. From one half mile west of Loxa west to near Anderson Road, to
telegraph pole No. 12330: S.11, T.12 N., R.8 E. (Pl. VI. and
VIL.)
b. Prairie at Shea’s: 8.17, T.12 N., R.8 E.
c. Cord Grass (Spartina) Association. East of Shea’s: 8.17, T.12
N., R.8 E.
Station III. Prairie east of Charleston. Right of way along the C. C. C.
& St. L. R. R.: 8.12, T.12 N., R.9 E.; 8.5, 6, and 7, T.12 N.,
R.10 E.
a. Rosin-weed (Silphium terebinthinaceum) Association. Just west
of the place where the Ashmore Road crosses the Big Four track;
about one mile east of Charleston: 8.12, T.12 N., R.9 E.
b. Blue Stem (Andropogon) and Rosin-weed (Silphium terebinthina-
ceum) Association. Three fourths of a mile east of the crossing of
the Ashmore Road and the Big Four track: 8.6 and 5, T.12 'N.,
R.10 E. An area which grades from prairie into transitional for-
est conditions. (Pl. VIII and IX.)
Station IV. Bates Woods. On the east bluffs and bottom of the Embar-
ras River, north of where the Cleveland, Cincinnati, Chicago and
St. Louis, or Big Four, R. R. crosses the river. On the farm of
J. I. Bates: 8.5, T.12 N., R.10 E. (Pl. X, Fig. 1; Pl. XI, XII,
and XIII.)
a. Upland Oak-Hickory Association (Quercus alba and Q. velutina,
and Carya alba, C. glabra, and C. ovata.) Second-growth forest.
(Pl. XII and XIII.)
b. Embarras Valley and Ravine Slopes, with Oak-Hickory Associa-
tion.
c. Red Oak (Quercus rubra), Elm (Ulmus americana), and Sugar
Maple (Acer saccharum) Association. Lowland or ‘‘second bot-
tom,’’ Embarras Valley. (Pl. XIV; XV; and XVI, Fig. 1 and 2.)
d. Small streamlet in South Ravine. This formed the southern bor-
der of the area examined. A temporary stream. (Pl. XVII, Fig.
1 and 2.)
DESCRIPTION OF THE PRAIRIE HABITATS AND ANIMALS
J. Prarie Area NortH of CHARLESTON, StTaTION I
This area includes patches or islands of prairie vegetation oc-
curring along the right-of-way of the Toledo, St. Louis and West-
41
ern, or “Clover Leaf,” Railway, north of Charleston. The south-
ern border began just beyond the area of numerous side tracks and ex-
tended north of the first east and west cross-road for a distance of
about one mile, to the place where the right-of-way is much narrowed
and fenced off for cultivation. This is a strip of land through the level
black soil area, which was originally composed of dry and wet prairie.
The higher portions have a lighter colored soil, and the lower parts
have the black and often wet soil which characterized the original
swamp or wet prairie. The railway embankment and the side drain-
age ditches have favored the perpetuation of patches or strips of these
wet habitats ; the excavations for the road-bed, on the other hand, have
accelerated drainage of the higher grounds. The soil taken from these
cuts and heaped up on the sides of the tracks reinforces the surface
relief noticeably in a region which is so nearly level. Through the
depressions fillings have been made in building the railway embank-
ment, and as a result the drainage has been interfered with in some
places.
The disturbances brought about by railway construction and main-
tenance have greatly modified the original conditions, so that the
prairie vegetation persists usually only in.very irregular areas, some-
times reaching a maximum length equal to the combined. distance be-
tween three or four consecutive telegraph poles—these poles are gen-
erally about 200 feet apart. In breadth the area is usually less than
the space between the ditch bordering and parallel to the road-bed or
embankment and the adjacent fence which bounds the right-of-way, or
about 40 feet. This entire right-of-way is about roo feet wide. In
addition to these changes in the physical conditions, a large number of
weeds not native to the prairie have been introduced, opportunities for
this introduction being favorable, as railways traverse the entire area.
In general, attention was devoted solely to the areas or colonies of
prairie vegetation and their associated invertebrate animals, the areas
of non-prairie vegetation being ignored, not as unworthy of study, but
because the vanishing prairie colonies required all the time available.
rt. Colony of Swamp Grasses (Spartina and Elymus), Station I, a
This colony of slough grass (Spartina michauxiana) and wild rye
(Elymus) is located a short distance north of the “Clover Leaf” switch
tracks and just south of the telegraph pole marked “Toledo 318 miles
and St. Louis 133 miles.” The length of this colony was about 40
aces.
: During August, 1910, it was dry, but probably in the spring and
early summer, rains make this area a habitat for swamp grasses.
42
Though it was an almost pure stand of slough grass, with this were
mixed a few plants of wild rye (Elymus virginicus submuticus and E.
canadensis). These grasses reach a height of about four feet. The
ground was very hard and dry, and there were large cracks in it.
A single collection of animals was made here, No. 179.
Common Names Scientific Names
Common Garden Spider Argiope aurantia
Ambush Spider Misumena aleatoria
Differential Grasshopper, adult
and nymphs Melanoplus differentialis
Red-legged Grasshopper, adult
and nymphs Melanoplus femur-rubrum
Texan Katydid Scudderia texensis
Meadow Grasshopper Orchelimum vulgare, adult, and
nymphs of vulgare or glaberri-
mum.
Dorsal-striped Grasshopper Xiphidium strictum
Black-horned Meadow Cricket Cicanthus nigricornis
Four-spotted White Cricket Cicanthus quadripunctatus
Ground-beetle Leptotrachelus dorsalis
Sciomyzid fly Tetanocera plumosa
The basic food-supply in such a habitat is of course the grasses, and
this fact fully accounts for the presence of large numbers of individ-
uals which feed upon grasses, as do the Orthoptera in general. But
the Orthoptera listed are not exclusively vegetable feeders, for Forbes
(’0§: 147) has shown that Xiphidium strictum feeds mainly upon in-
sects, chiefly plant-lice,as well as upon vegetable tissues, including fun-
gi and pollen; Orchelimum vulgare (p. 144), largely upon plant-lice
and other insects; and Cécanthus quadripunctatus (p. 220), upon plant
tissues, pollen, fungi, and plant-lice. These observations were based
upon a study of the contents of the digestive tract. The food of the
sciomyzid fly is unknown. The garden spider lives exclusively upon ani-
mal food; and being abundant, it must exert considerable influence
upon other small animals. It not only destroys animals for its food, but
many others are ensnared in its web and thus killed. In one of the
webs I found a large differential grasshopper. The rank growth of
vegetation furnishes the necessary support for the webs of this spider.
Some of the insects, as Melanoplus differentialis and M. femur-
rubrum, oviposit in the soil, but others—Scudderia texensis, Xiphid-
ium strictum, Orchelimum vulgare, and Cicanthus—deposit their
43
eggs in stems of plants or under the leaf-sheaths of grasses (Forbes,
05: 143, 145, 148, 216). The mode of oviposition in these Orthop-
tera raises the question whether or not they are able to pass their com-
plete life cycle within this habitat. Are the species which oviposit in
the soil able to endure submergence during the wet season of the year,
or must they each year re-invade this habitat from the more favorable
adjacent regions? The sciomyzid fly is a regular inhabitant of such
situations, for an allied species, Tetanocera pictipes Loew, has been
found by Needham (’o1: 580) to be aquatic, breeding on colonies of
bur reed (Sparganium), and Shelford (’13a: 188, 284) also finds
plumosa in wet places.
The flower spider, Misumena, captures its prey direct, frequenting
flowers where its prey comes to sip nectar,
With more perfect drainage the character of this habitat wouid
change; a more varied growth of vegetation would probably devel-
op; and the relative abundance of the various kinds of animals would
also change. The present imperfect drainage is more favorable to the
accumulation of vegetable debris than if the habitat was connected
with a stream which could float it away. The periodical drying hastens
decay, and the deep cracks in the soil become burial places for various
kinds of organic debris.
2. Colony of Wild Rye, Elymus virginicus submuticus, Station I, c*
Wild rye is a swamp grass. This colony was located about half a
mile north of the colony of slough grass (Station I, a) and about 222
feet south of the first east and west cross-road north of Charleston.
For a general view of this grassy habitat see Figure 1, Plate II. In
length this habitat extends about one third the distance between two
consecutive telegraph poles, or about 65 feet. The conditions of the
habitat are in general similar to those in the colony of Spartina. The
black soil was very dry and much cracked when examined, late in Au-
gust. Though a few plants of Asclepias sullivantii grew here among
the grass, it was a dense, almost pure stand of wild rye, which reached
a height of about three and a half feet.
Only a very few collections were made here, and these were for
the sole purpose of determining the general composition of the asso-
ciation.
These collections, Nos. 153, 180, and 181, were as follows:
*Animals were not emia’ at Station I, 6, and therefore the location will not be
discussed here.
44
Common Garden Spider Argiope aurantia No. 153
Differential Grasshopper Melanoplus differentialis
Red-legged Grasshopper Melanoplus femur-rubrum No. 180
Dorsal-striped Grasshopper Xiphidium strictum No. 180
Meadow Grasshopper Orchelimum vulgare, adult,
and nymphs of vulgare
or glaberrimum No. 180
Texan Katydid Scudderia texensis No. 181
These are all abundant species. O. vulgare, by its persistent fid-
dling, is noticeable in all such grass spots during hot sunny weather.
A live differential grasshopper was found in the web of the garden
spider. A comparison of the two colonies of swamp grasses, Spartina
and Elymus, will probably help to give one a general idea of the kind
of invertebrates which were abundant in the original swamip-grass
area of this vicinity. It will be noticed that grass and grass eaters are
the dominant species, and that upon these a smaller number of preda-
ceous animals depend. The characteristic species are the Orithoptera
and the garden spider. This spider, on account of its predaceous hab-
its, is able to live in a great variety of open situations, but does not
normally live in dense woodlands.
3. Wet Area of Swamp Milkweed (Asclepias incarnata), Station I, d
This colony of swamp milkweed was about one eighth of a mile
north of the east and west cross-road. This flat, poorly drained black-
soil area, about 80 feet long, was wet throughout August, crawfish
holes being abundant (Pl. IITA, fig. 2; Pl. IIIB, figs. 1, 2). To
the east, beyond the boundary fence, in the adjoining corn field, stood
a pool of water surrounded by a zone of yellowish weakened corn,
visited occasionally by a few shore birds. Along the east side of the
newly formed railway embankment (PI. II], fig. 1) is a shallow
trench containing water and a growth of young willows (Salix) and
cottonwoods (Populus deltoides), also blue flags (Iris versicolor),
bulrush (Scirpus), and sedge (Carex). The characteristic plants
over this area were the abundant swamp milkweed (Asclepias incar-
nata, Pl. ILIA, fig. 1) and Bidens. A few plants of water horehound
(Lycopus) and dogbane (Apocynum medium) were present, and many
individuals of a low plant with a winged stem (Lythrum alatum).
The collections (Nos. 1, 12, 13, 14, 15, 18, 32, 37, 45, 156, and
157) of animals taken here were as follows:
45
Pond snail Galba umbilicata 18
Prairie Crawfish Cambarus gracilis _
Garden Spider Argiope aurantia _
Ambush Spider Misumena aleatoria 157
Chigger Trombidium sp. —_—
Nine-spot Dragon-fly Labellula pulchella _—
Stink-bug Euschistus variolarius 12
Small Milkweed-bug Lygeus kalmii 12
Large Milkweed-bug Oncopeltus fasciatus I
Ambush Bug Phymata fasctata 12
Tarnished Plant-bug Lygus pratensis 12
Soldier-beetle Chauliognathus pennsylvanicus 156
Black Flower-beetle_ Euphoria sepulchralis 156
Four-eyed Milkweed-beetle Tetraopes tetraophthalmus 12
Milkweed-beetle Tetraopes femoratus (?) I
Leaf-beetle Diabrotica atripennis I
Dogbane Beetle Chrysochus auratus 14
Celery Butterfly Papilio polyxenes 15, 45
Philodice Butterfly Eurymus philodice 12
Idalia Butterfly Argynnis idalia 33
Milkweed Butterfly Anosia plexippus —
Honeysuckle Sphinx Hemaris diffinis 32
Giant Mosquito Psorophora ciliata 13
Giant Fly Mydas clavatus 12
Honey-bee Apis mellifera —_
Pennsylvania Bumblebee Bombus pennsylvanicus 155
Bumblebee Bombus fraternus 12
Bumblebee Bombus separatus 12, 157
Carpenter-bee Xylocopa virginica 1,156
Rusty Digger-wasp Chlorion ichneumoneum I2
The soft, wet, black soil contained large numbers of crawfish holes,
and from several of them T. L. Hankinson dug specimens of Cambarus
gracilis. Frogs (Rana) were seen but none were secured. A Caro-
lina rail was flushed from the ditch along the track, and on the mar-
gins of the water in the adjacent corn field Mr. Hankinson recognized
some shore birds. The dragon-fly Libellula pulchella was abundant on
the wing and resting on the vegetation, and two examples were found
in the webs of Argiope aurantia. No nymphs were found, but doubt-
less eggs were laid by some of the numerous adults. It was interest-
ing to observe the fresh burrows of the crawfish which had traversed
the fresh firm yellow clay of the recently reinforced railway embank-
46
ment (shown im Pl. Ul, fig. 2) and appeared upon its surface. ‘The
occurrence hereof 4 swall anal. Calbuccaddivata, is of imterest. A
very large species of mosquito with conspicuously banded legs, Psoro-
phora ciliata, was found here. Though these aquatics and the ground
fonms did not receive much attemtion, they are representative of wet
places.
The presence of certain plants im this habitat has determined. the
occurrence of several species of animals. ‘Thus the dogbame Apocy-
mm medium accounts for the brilliamtly colored leaf-beetle Chry-
sochus auratus, which feeds wpon its leaves and roots. But the most
conspicuous feature of this habitat im August is the wariety of imsects
which are attracted by the flowers of the swamp milkweed. These
flowers may be regarded as so much imsect pasture. A few butterflies
were observed, Papiho polyxenes beimg found im am Argiope web; and
om the flowers of the swamp milkweed were Papilio cresphomtes, Ewry-
mus plilodice, Al gyums daha, Anosia plesippus, and the honeysuckle
sphinx (Hemaris defenis). Among the most abundant Hymenoptera
were the homey-bee (Apis mellifera) and the common rusty digger-
wasp (Chlorion ichmewmoncum). Others were the carpenter-bee
(Xylocopa virgimca) and the bumblebees Bombus fratermus and sep-
aratus. On the flowers of the thistle (Cirsium) near this station, Bowm-
bus pemmsyloomicus was also talsem lr amet ney ours
was taken om the flowers of the swamp milkweed. Beetles from these
flowers were the spotted muilkweed-beetles (Tetraopes tetraophthalmus
amd femoratus?) the flower-bectle Euphoria sepulchralis, amd, late
im Amgust, great mumbers of the soldier-beetle Chauliogmathus pemm-
SyWoumicus.. The Hemiptera found are equally chararteristic, and in.
weed thugs (Oncopelius fasciatus andl
Lygeus kalo) and Lygus pratensis. a
om the milkweeds, preying mot wpom the plamt, bact tmpom
These were tie amiuch img (Pleymata fascioia) and the cinbush
sects of this Ihalbntat: att tihis seasom. Amother abundant: animal was tie
chigger, a larval mute of the genus Trombidionm, which is brushed from.
the vegetation by one’s anms and legs. “Mhese irritating pests were so
abomdamt that to worl with comfort im this regiom it was mecessary
to ponsice one's chutes and bal wat Sowezs ot seteien: ‘These
young sixlegwed mutes are supposed to prey mpom imsents, as do the
adults. Acoording to Chittenden (06:4) diggers are most abum-
damt im damp places and forest margims, amd among shrubs, grass,
47
and herbage. “The adults are knowm to eat plamt-lice, small caterpil-
lars, amd grasshoppers’ eggs. ‘This mite is thus am importamt preda-
cross aneuaher of the association. The dragnnfies are swell kinrars 10
feed wpom small insects, which they capture om the wing, and om ac-
coumt of thenr abundance they are imfimential imsects here.
Am exammation of the list of ammmals secured at this station
shows that there is considerable diversity im the conditions under which
their breeding takes place. Indeed the breeding habits amd places are
almost as diwerse as are the feedimg relations. ‘Thus the smail Galbo
breeds im the water ; amd the crawfish, Cambarus gracilis, lives as a tur-
rower except for a brief period im sprimg, whem it is foumd im streams.
It 1s distimetly a subterrameam species. “The gardem spider, im the fall,
leaves its epps im its web. "The life iistory of the ambush spider is mot
kmowm. Itt seems probable that the sexes meet mpom flowers, amd as
the flowers fade they migrate to fresh omes—a respomse which Ham-
cock has observed ("11 : 182-186) im the allied species Miswmmena
voto. "The ambush bug, whem found om flowers, is im a large mmmiber
of cases copullatimg, but where the eggs are laid amd the young dewel-
oped is mnkmowm to me. “Mhougi this bug also must migrate with the
fading of the flowers, after the halbit of Miswmenm, it is wimged amd
does mot have to go “om foot” as the spider probably does. Wem dis-
thurbed these fbmgs do mot as a mule seek to escape by flight, amd it is mot
unlikely that they often crawl from ome flower to amotiher winem tie
diistameae is short. "Mhe soldiertbeetle is similar to the ambush bog im
its propemsitty to copulate om flowers. ‘The milkweed beetles amd the
doghane hectle are commonly seem copullatimg mpon the leawes amd
stems of tthe plats om winch they lnwe. The tarva of the milkweed
beetles bore imo the roots and stems of plamts; the doghame beetle has
similiar halts. Of the butiterfiies, Amoma was observed copulatimg om
tthe willows, ome sex with the wimgs spread, tthe fore omnes overlappimg
im part the immder pane, tthe otter sex with the wings folded ttogetiner
werticailly, the heads of the imsects tbheimg ttmnmed im opposite directions.
‘The egws of the mosquito are laid mear the surface of the watter. “Mhe
honeybee andl immiblledbees are social, amd the breedimg amd care of
the young are quite differemt fhrom those of the other ammmals fomnd
an tits soln Xywlocopa cuts the mest for tts brood im solid wood,
-
5 rather foreign mpom the pramie, alithomgh posts amd ties
wien is ding: itm tthe grrommdl, with warioms grasshoppers; mpom tihese tire
egy iis land amd the young larva feeds. “This wasp probally did mot
freed! im this moist thalbitat. “Mhe wet suibsthratumm here is probally mm-
faworablle for tthe ireedimg of those Orthopitera winch deposut thhenr
epys im tie soll.
48
4. Cone-flower and Rosin-weed Colony, Station I, e
This station was continuous with and just north of the swamp
milkweed area (Station I,d) just described. The surface of the
ground sloped gently upward toward the north, but none of it was free
from crawfish holes, and the ground-water level was not far below.
The soil is very dark in color.
The general appearance of this habitat is shown in Plate V. The
large-leaved plants are Silphium terebinthinaceum, and the heads of
the numerous cone-flowers (Lepachys pinnata) show as black points in
the picture. The cone-flower was the dominant plant at this time.
There were a few scattered plants of Silphium integrifolium and of
wild lettuce (Lactuca canadensis). At the time the collecting was done
in this area Silphium was not in blossom, and all the flower-collecting
was from Lepachys.
The collections of animals taken here (Nos. 8, 40, and 158) are
as follows:
Crawfish Cambarus sp. (Burrows observed)
Garden Spider Argiope aurantia 40
Sordid Grasshopper Encoptolophus sordidus 158
Differential Grasshopper Melanoplus differentialis 40
Red-legged Grasshopper Melanoplus femur-rubrum 40
Texan Katydid Scudderia texensis 40
Dorsal-striped Grasshopper Xiphidium strictum 40
Black-horned Meadow Cricket Gicanthus nigricornis 40
Membracid bug Campylenchia curvata 40
Jassid Platymetopius frontalis 40
Lygeid Ligyrocoris sylvestris 40
Ambush Bug Phymata fasciata 40
Chrysomelid beetle Nodonota convexa 40
Southern Corn Root-worm Diabrotica 12-punctata 40
Beetle
Robber-fly Asilide _—
Trypetid fly Euaresta equalis 40
Eucerid bee Melissodes bimaculata 8
Eucerid bee Melissodes obliqua 8
Nomadid bee Epeolus concolor 8
Social wasp Polistes sp. _—
Collection No. 40 was made by sweeping the vegetation with an in-
sect net. No. 8 is a collection made from the flowers of Lepachys pin-
nata. The nest of Polistes was across the railway track. from this
station. The abundance of Melissodes obliqua and of the pretty
49
Epeolus concolor on the flowers of Lepachys indicates the attractive
power of this plant. The coarser plants furnish support for the webs
of Argiope, the flowers serve as drinking cups in which Phymata lies
in ambush; and the varied vegetation affords food for the numerous
Orthoptera. The proximity of ground-water accounts for the pres-
ence of Cambarus, and an adjacent corn field explains the presence
of Diabrotica. A robber-fly (Asilide) was seen but not captured. It is
interesting to see Melissodes obliqua as it hurries round and round the
heads of cone-flowers and sweeps up the great masses of yellow poilen.
The hind pair of legs, when loaded with pollen, have nearly the bulk
of the abdomen. Robertson (’94; 468) says that this is the most
abundant visitor to the cone-flower, and more abundant on this flower
than on any other.
It is probable that the conditions within this habitat were suitable.
for the breeding of most of the species listed. Huaresta equalis has
been bred from the seed pods of the cocklebur (Xanthium) and prob-
ably came from the adjacent corn field. It is most likely on flowers
that the strepsipterid parasitic insects find many of their hosts (Pierce
‘og b: 116). These insects are found on the following prairie insects :
Polistes, Odynerus, Chlorion ichneumoneum, C. pennsylvanicum, and
C. atratum. Robertson (’10) records many important observations on
the hosts of Illinois Strepsiptera.
5. Colony of Blue Stem (Andropogon) and Drop-seed (Sporobolus),
bordered by Swamp Milkweed, Station I, g*
This colony formed the extreme northern part of the prairie area
examined along the “Clover Leaf” track. It extended along the track
for a distance of about 200 feet. The area is level black soil prairie.
Its general appearance and location are indicated in Figure 2, Plate
II, and in Figure 2, Plate III, photographs taken at the time of our
study, and in Figure 2, Plate IV, a photograph taken by T. L. Hankin-
son April 23, 1911. This latter view clearly shows the character of the
drainage during the spring wet season. During the late summer, the
dry season, the ditch along the railway track concentrates the drainage
so that a colony of swamp milkweed (Asclepias incarnata) and small
willows flourish in it. Upon the well-drained part of this area there is
a rather rich growth of Andropogon furcatus, A. virginicus, and
Sporobolus cryptandrus, and many plants of the dogbane Apocynum
medium and a few plants of Asclepias sullivantii. This was the larg-
est and best colony of the upland prairie grasses seen along the Clover
Leaf tracks; and yet when it is compared with the patches of such
*No collections were made at Station I, f.
50
grass east of Charleston (Station III) it is a meager colony. Just
south of this grassy colony was a large one of the mountain mint.
Pycnanthemum flexuosum. This is shown in Figure 1, Plate IV.
The collections of animals (Nos. 1, 2, 3, 4, 6, 7, 19, 28a, 36, 39, 44,
157, and 159) are as follows:
Pond snail
Crawfish
Harvest-man
Garden Spider
Ambush Spider
Red-tailed Dragon-fly
Nine-spot Dragon-fly
Prairie Ant-lion
Lace-wing Fly
Grasshopper
Sordid Grasshopper
Differential Grasshopper
Red-legged Grasshopper
Texan Katydid
Meadow Grasshopper
Cone-nosed Katydid
Four-spotted White Cricket
Stink-bug
Small Milkweed-bug
Large Milkweed-bug
Rapacious Soldier-bug
Ambush Bug
Four-eyed Milkweed Beetle
Rhipiphorid beetle
Bill-bug
Milkweed Butterfly
Giant Mosquito
Mycetophilid fly
Giant Bee-fly
Vertebrated Robber-fly
Honey-bee
Bumblebee
Bumblebee
Eucerid bee
Nomadid bee
Leaf-cutting bee
Rusty Digger-wasp
Myzinid wasp
Physa gyrina 19
Cambarus sp. _
Liobunum politum? 7
Argiope aurantia 6, 39
Misumena aleatoria 6, 157, 159
Sympetrum rubicundulum 7
Libellula pulchella —
Brachynemurus abdominalis 36
Chrysopa oculata 44
Syrbula admirabilis 3
Encoptolophus sordidus 44
Melanoplus differentialis 39
Melanoplus femur-rubrum 3, 39
Scudderia texensis 2, 44
Orchelimum vulgare —, 3
Conocephalus sp. 159
CEcanthus 4-punctatus 3
Euschistus variolarius 39
Lygeus kalmii 1,6
Oncopeltus fasciatus I
Stnea diadema 5
Phymata fasciata I
Tetraopes tetraophthaimus I
Rhipiphorus dimidiatus 6
Sphenophorus venatus 39
Danais archippus i
Psorophora ciliata 44
Sciara sp. 6
Exoprosopa fasciata 6
Promachus vertebratus 39, 44
Apis mellifica I
Bombus fraternus I
Bombus separatus I
Melissodes bimaculata 6
Epeolus concolor 6
Megachile mendica I
Chlorion ichneumoneum I
Myzine sexcincta 1, 6
51
Physa and Cambarus were found among the milkweeds on account
of the wet ground, and the presence of the giant mosquito was prob-
ably due to the same condition. The majority of the other animals
were attracted to this habitat by the milkweed, particularly by its flow-
ers. Among these were the milkweed bugs and beetles, the milkweed
butterfly, the honey-bee, and the rusty digger-wasp. The dense growth
of the milkweeds does not appear to be so favorable to the garden:
spider as is the more open and irregular growth of vegetation else-
where. The ambush spider frequented the milkweed flowers for prey
and also the flower masses of the mountain mint, on which it was in
active competition with the ambush bug and the rapacious soldier-bug,
which have similar food habits. The mountain mint, whose flowers
are frequented by the predaceous animals just! mentioned, is also vis-
ited by rhipiphorid beetles, the bee-fly (Exoprosopa fasciata), the bees
Melissodes bimaculata and Epeolus concolor, and the myzinid wasp
Myzine sexcincta. ‘The prairie grasses were frequented by a large
variety of Orthoptera, which showed a decided preference for them,
their abundance being evident in the list. The wide-ranging predators
and parasites, such as Liobunum, Libellula, Sympetrum, Chrysopa,
Brachynemurus, Promachus, Chlorion, and Myzine, probably forage
over extensive areas compared with the relatively sedentary kinds,
such as Misumena, Argiope, Phymata, and Sinea. Phymata was cap-
tured on a milkweed flower with a honey-bee; Promachus vertebratus
was taken on a grass stem with a stink-bug (Euschistus variolarius) ;
and Misumena aleatoria was taken with a large, nearly mature female
nymph of Conocephalus.
The conditions which permit an animal to breed in a habitat have
an important influence upon the character of its population. It is evi-
dent that many of the animals taken do not breed here. Some of the
relatively sedentary kinds, such as Physa, Cambarus, and Argiope, and
probably Misumena, do not cover long distances. Good examples of
the wider ranging forms are Sympetrum, Libellula, Danais, Proma-
chus, Apis, Bombus, and Chlorion. Several of the animals, as the
snails, crawfish, and the dragon-flies, require an aquatic habitat.
Chrysopa places its eggs among colonies of plant-lice, and Brachyne-
murus probably spends its larval life in dry or sandy places, feeding
upon ants and other small insects, as do other ant-lions. Several of
the Orthoptera deposit their eggs in the soil; and some of the locustids,
among grasses and herbaceous stems. Others are found copulating
upon the plants on which the young feed, as Tetraopes, Chrysochus,
Lygeus, and Oncopeltus; and still others copulate in the flowers
mainly, as Phymata. It is probable that on the flowers some of the para-
52
sitic species find their hosts, as Pierce (’04) has shown to be the case
in the rhipiphorid genus Myodites. Rhiptphorus is probably parasitic.
6. Supplementary Collections from Station I
In addition to the specimens given in the preceding lists for Statio
I there are others, general collections from this area, which should b
listed for this prairie. For details concerning each species of the fol
lowing consult the annotated list.
Garden Spider
Ambush Spider
Chigger
Dorsal-striped Grasshopper
Coreid bug
Ambush Bug
Ladybird
Leaf-beetle
Four-eyed Milkweed Beetle
Old-fashioned Potato Beetle
Margined Blister-beetle
Black Blister-beetle
Snout-beetle
Snout-beetle
Giant Bee-fly
American Syrphid
Tachinid fly
Bumblebee
False Bumblebee
Eucerid bee
Short Leaf-cutting Bee
Halictid bee
Halictid bee
Stizid wasp
Rusty Digger-wasp
Harris Digger-wasp
Digger-wasp
Solitary wasp
Argiope aurantia
Misumena aleatoria
Trombidium sp.
Xiphidium strictum
Harmostes reflexulus
Phymata fasciata
Hippodamia parenthesis
Trirhabda tomentosa
Tetraopes 4-ophthalmus
Epicauta vittata
Epicauta marginata
Epicauta pennsylvanica
Centrinus penicellus
Centrinus scutellum-album
Exoprosopa fasciata
Syrphus americanus
Trichopoda ruficauda
Bombus separatus
Psithyrus variabilis
Melissodes obliqua
Megachile brevis
Halictus fasciatus
Halictus virescens
Stigus brevipennis
Chlorion ichneumonenm
Chlorion harrisi
Ammophila nigricans
Odynerus vagus
26
31
35
27
24, 26, 43
Hankinson
Hankinson
Hankinson
Hankinson
26, 152
41
Hankinson
24, 31
II
38
22
22
24, 48
Hankinson
26
23
35, Hankinson
II. Pratrtz Area NEAR Loxa, ILLINOIS, Station IL
This station includes patches of prairie along the Cleveland, Cin-
cinnati, Chicago and St. Louis (Big Four) railroad right-of-way be-
tween Charleston and Mattoon, Ill, and about one mile west of
53
the small station of Loxa. Along this track the telegraph-pole num-
bers were used in locating our substations. This is a rather level black
soil area, originally poorly drained and wet, but now considerably
modified by the ditching and grading occasioned by railway construc-
tion and maintenance. The changes have been similar to those on the
prairie north of Charleston, but the ditching has been a few feet deeper
and the embankment is higher. The most abundant and characteristic
kinds of vegetation are the tall prairie grasses—blue stem (Andropo-
gon furcatus), drop- seed (S‘porobolus cryptandrus), and beard grass
(Andropogon virginicus)—a rosin-weed (Silphium laciniatum), the
flowering spurge (Euphorbia corollata), wild lettuce (Lactuca can-
adensis), rattlesnake-master (Eryngium yuccifolium), and beggar-- -
ticks (Desmodium). Many other kinds of plants were also present.
The general appearance of this habitat is shown in plates VI and VII.
Our collections from this prairie (Nos. 47-57 and 176-178) are as
follows:
Garden Spider Argiope aurantia 49, 179
Ambush Spider Misumena aleatoria 47, 178
Sordid Grasshopper Encoptolophus sordidus 48
Two-lined Grasshopper Melanoplus bivittatus 55
Differential Grasshopper Melanoplus differentialis 48
Meadow Grasshopper Orchelimum vulgare 178
Lance-tailed Grasshopper Xiphidium attenuatum 48
Dorsal-striped Grasshopper Xiphidium strictum 48, 50, 57
Stink-bug Euschistus variolarius 50, 52, 178
Ambush Bug Phymata fasciata
48, 52, 54, 55, 57, 178
Dusky Leaf-bug Adelphocoris rapidus 55
Soldier-beetle Chauliognathus pennsylvanicus 178
Southern Corn Root-worm = Diabrotica 12-punctata 55
Margined Blister-beetle Epicauta marginata 48
Black Blister-beetle Epicauta pennsylvanica 48, 178
Rhipiphorid beetle Rhipiphorus dimidiatus 52
Rhipiphorid beetle Rhipiphorus limbatus 178
Snout-beetle Rhynchites eneus 48
Thoe Butterfly Chrysophanes thoe 55
Dogbane Caterpillar Ammalo eglenensis or tenera 53
Giant Bee-fly Exoprosopa fasciata 47, 57, 176
Robber-fly Deromyia sp. 51
Vertebrated Robber-fly Promachus vertebratus 56
Corn Syrphid Mesogramma politum 177
Syrphid fly Allograpta obliqua 177
54
Tachinid fly Cistogaster immaculata
Pennsylvania Bumblebee Bombus a 50, 52, 55, 176
False Bumblebee Psithyrus variabilis
Eucerid bee Melissodes bimaculata “43
Nomadid bee Epeolus concolor 48, 52
Halictid bee Holictus obscurus
Halictid bee Halictus fasciatus 48, 52
Black Digger-wasp Chlorion atratum 55
Pennsylvania Digger-wasp Chlorion pennsylvamcum 55
Myzinid wasp Myszine sexcincia 52, 55
Ant Formica pallide-fulva schaufussi
incerta 52
The general conditions of this prairie appear to have been less dis-~
turbed than at Station I; at least the prairie vegetation is more exten-
sive and uniform. The change in the vegetation is apparently greater
than the change in the kinds of animals. Their feeding and breeding
relations appear to be much like those at the prairie stations previously
discussed.
In the flowers of the cup-leaved rosin-weed (Silphium integri-
folium) was found a giant bee-fly (Exoprosopa fasciata) which had.
been captured by the ambush spider (Misumene aleatoria), and on
webs in colonies of this same plant the garden spider (Argiope auran-
tia) was observed, with a grasshopper (Melanoplus differentialis) en-
tangled in the web. From the flowers of this Silphium the following
insects were taken: Epicauta marginata and E. pennsylvanica, Rhyn-
chites eneus, Phymata fasciata, Encoptolophus sordidus, Melanoplus
differentialis (nymph), Xiphidium strictum (adult and nymph), X.
attenuatum, Melissodes bimaculata and obliqua, Epeolus concolor, and
Halictus fasciatus. ‘The margined blister-beetle (Epicauta marginata)
was found both upon the flowers and the leaves of the plant. On the
flowers of the purple prairie clover (Petalostemum purpureum), Bom
bus pennsylvanicus, Xiphidium strictum, and Euschistus variolarius
were taken. Collection 176 was taken from the flowers of Liatris
scariosa, and Nos. 55 and 178 from the flowers of Eryngium yucci-
folium.
Swarms of the small corn syrphid, Mesogramma politum, were
present, on one day settling by dozens on my hands and clothes, where
they were easily grasped by the wing. It had been a warm day, and
this swarming was in the sunshine at about 4:30 p.m. The flies came
from a large corn field a few feet away.
35
THE. Prams Agra East or Cnartesron, Sration IT
This prairie area is about two miles east of Charleston along the
“Big Four” railway track. ‘There were two colonies here. One, sub-
station a, was on low black-soil prairie just west of the first morth and
south road crossing the railway track east of Charleston. This was
largely a colony of the large-leaved rosin-weed, Silthium terebinthi-
naceum. ‘The second colony, substation b, was a mile and a half di-
rectly east of substation a, and half a mile east of the second north and.
south road east of Charleston.
Substation or “station” a was originally far out upon the black soil
prairie; b, on the other hand, is of special interest because it was origi-
nally wooded, has been cleared and maintained as a railroad right-of-
way, and contains today, therefore, a practically unique mixture of for-
ca and geditis peaitscad gaiicle. wat the pattie Viads denseciteag:
The soil, lighter in color than the black soil prairie, is representative of
the wooded regions. This colony has every appearance of a cleared
forest area invaded by prairie organisms.
The animals at station a were not studied, and ihe only record is
that of the black blister-beetle, Epicauta pennsylvanica (No. 119),
which was abundant on the flowers of Silphinm terzbinthinaceum.
At station 6 excavation was necessary to lower the road-bed, and
upon the disturbed soil thus thrown up along the track the prairie veg-
etation had become established. The general appearance of this region
is shown in plates VIII and IX. Here grew large quantities of rosin-
weed (Silphium terebinthinaceum) and blue stem (Andropogon) ; in
places upon high ground, indeed, this prairie grass was
Associated with it was the flowering spurge, Euphorbia corollata, as
seen' in Plate VIII The forest near by is shown in the background.
This same forest and grass area is shown in the background and mid-
dle of Plate IX, and in the foreground of the same picture is shown
the mixture of prairie and forest plants. Here are hickory sprouts,
crab-apple, grape, sumac, and smilax, intermingled with Silphium,
blue stem, and Lactuca canadensis. Not all of these appear in the
photograph, but they were present in some parts of the colony.
The collections here (Nos. 58-62 and 175) are as follows:
Leather-colored Grasshopper § Schistocera alutacea 59
Black-horned Meadow Cricket Gcanthus nigricornis 62
Meadow Grasshopper Orchelimum vulgare 175
Soldier-beetle Chauliognathus pennsylvamcus 175
Spotted Grape-beetle Pelidnota punctata 58
Black Blister-beetle Epicauta pennsylvamica
(Sta. TI,a) 119
56
Cabbage Butterfly Pontia rape 61
Vertebrated Robber-fly Promachus vertebratus 62
Pennsylvania Bumblebee Bombus pennsylvanicus 175
Impatient Bumblebee Bombus impaitiens 175
Bumblebee Bombus auricomus 175
(Rose-gall) Rhodites nebulosus 60
No animals were taken here which were dependent upon the sumac,
hickory, crab-apple, or smilax. Pelidnota lives upon the grape, and
grapes are primarily woodland or forest-margin rather than prairie
plants. Schistocerca is also probably a marginal species. On the flow-
ers of Silphium terebinthinaceum were taken Orchelimum vulgare,
Chauliognathus pennsylvanicus, and Bombus pennsylvanicus, auri-
comus, and impatiens.
The persistence of woodland vegetation in this locality, in spite
of the repeated mowings and burnings, shows that it has much vigor,
and would, if undisturbed, in a few years shade out the prairie vege-
tation and restore the dominance of the forest. With such a change in
the vegetation there would of course be a corresponding change in the
animals.
DESCRIPTION OF THE FOREST HABITATS AND ANIMALS
1. The Bates Woods, Station IV
The Bates woodland area is located about three and a half miles
northeast of Charleston on the farm that was owned by Mr. J. I. Bates,
and consists of about 160 acres. It includes a bottom-land area near
the Embarras River, and extends up the valley slope on to the upland.
It is isolated from the trees bordering the river (Pl. X, fig. 1) by a
narrow clearing, and from those on the northeast, north, and north-
west by another clearing (Pl. XI); on the south and southwest it is
continuous with partially cleared areas, which extend south to the Big
Four railway track.
The river bottom-land is undulating and rises rather gradually
toward the base of the bluffs. The bluff line is irregular on account of
the ravines which have been etched in it, the largest of which forms
the southern boundary of the region examined. The upland is rela-
tively level. The soils on the bottom are darker colored, except in
places near the base of the bluff, and at the mouths of the ravines
where the upland soil has been washed down. The upland soil is pre-
sumably the “light gray silt loam” of the State Soil Survey (Moultrie
County Soils, Ill. Exper. Sta. Soil Rep., 1911, No. 2, p. 23). All of
57
the area examined was well drained, and all was forested. The region
is not homogeneous physically or in its vegetation, and for this reason
the area is divided into substations in order that the influences of the
local conditions within the forest might be preserved, and their indi-
viduality recognized.
2. The Upland Oak-Hickory Forest, Station IV,a
The general appearance of this forest is shown in plates XII and
XIII. This is an open second-growth forest composed of oaks and
hickories—such as white oak (Quercus alba), black oak (Q. velutina),
shag-bark hickory (Carya ovata), bitternut (C. cordiformis), pignut
(C. glabra), and scattered individual trees of red oak (Q. rubra), wal-
nut (Juglans nigra), and mulberry (Morus rubra). The shrubs are
sassafras (Sassafras variifolium), sumac (Rhus glabra), Virginia
creeper (Psedera quinquefolia), poison ivy (Rhus toxicodendron),
rose (Rosa), raspberry (Rubus), moonseed (Menispermum cana-
dense), and tree seedlings. The average diameter of the largest trees
is 8-10 inches. Most of the small growth consists of the sprouts from
stumps, and many of these are 2-3 inches in diameter. The forest
crown is not complete, and as a consequence there are more or less open
patches in which most of the herbaceous growth is found, such as
horse mint (Monarda bradburiana), pennyroyal (Hedeoma pule-
gioides), everlasting (Antennaria plantaginifolia), tick-trefoil (Des-
modium nudiflorum), and other, less abundant kinds. Even a plant
quite characteristic of the prairie, the dogbane Apocynum, was found
here in one of the open glades.
The forest floor has an unequal covering of dead leaves, largely
oak, most of which lie in the low vegetation and in slight depressions.
Occasionally there is but little cover and the light-colored soil is ex-
posed. There are few stumps and logs in this part of the forest, and
no thick layer of vegetable mold, so that one would not expect to find
any animals which normally frequent moist soil and vegetable debris.
As this is a second-growth forest it lacks the conditions which abound
in an original growth, where are old, dead and decaying trees, and
numerous decaying logs and stumps. In this respect the woods is not
fully representative of an original upland forest on well-drained bluff
land.
The relative evaporating power of the air of this substation was 54
per cent. of that of the standard instrument in the open garden at the
Normal School, a fact which indicates a relative evaporation com-
parable to that of the ordinary black-soil prairie ; in producing this con-
dition, the glade-like, open character of this forest is undoubtedly
an important factor.
58
The characteristics of this habitat may be summed up as follows:
upland, open, relatively dry second-growth oak-hickory forest, with
little undergrowth of shrubs and herbs, and with a small amount of
litter and humus; soil dry and firm; and few decaying stumps and tree
trunks.
The collections of animals made here (Nos. 64-67, 69, 71, 74-83,
88, 91-93, 102, 103, 107, 109, 118, 120-123, 127, 135, 136, 142, 145,
147, 150, 151, 162, 163, 166, 169, 170, 171, and 183) are as follows:
Land snail
Predaceous snail
Land snail
Carolina slug
Land snail
Harvest-spider
Harvest-spider
Stout Harvest-spider
Island Spider
White-triangle Spider
Rugose Spider
Ground Spider
White Ant
Ant-lion
Dog-day Harvest-fly
Periodical Cicada
Forest Walking-stick
Grouse Locust
Short-winged Grouse Locust
Green Short-winged
Grasshopper
Sprinkled Grasshopper
Boll’s Grasshopper
Lesser Grasshopper
Acridiid grasshopper
Acridiid grasshopper
Forked Katydid
Angle-winged Katydid
Common Katydid
Meadow Grasshopper
Meadow Grasshopper
Striped Cricket
Spotted Cricket
Woodland Cricket
Polygyra albolabris QI
Circinaria concava 71
Zonitoides arborea 71
Philomycus carolinensis 71
Pyramidula perspectiva 71, 88
Liobunum vittatum 82, 123
Liobunum ventricosum 123b
Liobunum grande 82
Epeira insularis 70
Epeira verrucosa 70
Acrosoma rugosa 70, 147
Lycosa sp. 142, 150
Termes flavipes 72, 76, 79
Myrmeleonide (Forest border) 183
Cicada linnei 162
Tibicen septendecim
Diapheromera femorata 64, 93
Tettigidea lateralis 109
Tettigidea parvipennis 122
Dichromorpha viridis
67, 92, 93, 121, 123
Chloealtis conspersa 67, 93, 122
Spharagemon bolli 67, 150
Melanoplus atlanis
Melanoplus amplectens 67
Melanoplus obovatipennis 93
Scudderia furcata 109
Microcentrum laurifolium 135
Cyrtophyllus perspicillatus 145
Orchelimum cuticulare 67, 93
Xiphidium nemorale 93, 103
Nemobius fasciatus 67, 93, 122
Nemobius maculatus 122
Apithus agitator 93
Woodland Tiger-beetle
Caterpillar-hunter
Carabid beetle
Ladybird
Splendid Dung-beetle
Dogbane Beetle
Tenebrionid larva
Philenor Butterfly
Turnus Butterfly
Troilus Butterfly
Sphingid larva
Arctiid moth
Notodontid moth
Notodontid moth
Notodontid moth
Geometrid moth
Gelechiid moth
(Cecidomyiid gall)
(Cecidomyiid gall)
(Cecidomyiid gall)
Syrphid fly
Corn Syrphid
Vespa-like Syrphid
Pigeon Tremex
(Oak Bullet-gally
(White Oak Club-gall)
(Oak Wool-gall)
Formicid ant
Formicid ant
Formicid ant
Mutillid ant
Short Caterpillar-wasp
59
Cicindela unipunctata 136
Calosoma scrutator 64
Galerita janus 171
Coccineliide 81
Geotrupes splendidus 120
Chrysochus auratus 103
Meracantha contractea 83
Papilio philenor 69, 166
Papilio turnus —
Papilio troilus 163
Cressonia juglandis 102
Halisidota tessellaris 168
Datana angusi 65, 162
Nadata gibbosa 169
Heterocampa guttivitta? 127
Eustroma diversilineata 163
Ypsolophus ligulellus?
76, 78, Hankinson
Cecidomyia holotricha 107, 170
Cecidomyia tubicola’ 107
Cecidomyia caryecola 107, 170
Chrysotoxum ventricosum 163
Mesogramma politum
76, 78, Hankinson
Milesia ornata 103
Tremex columba 66
Holcaspis globulus 170
Andricus clavula 170
Andricus lana 170
Cremastogaster lineolata 118
Aphenogaster fulva 74-80
Formica fusca subsericea 163
Spherophthalma I5I
Ammophila abbreviata 127
3. Embarras Valley and Ravine Slopes, forested by ihe Oak-Hickory
Association, Station IV,b
This station included the slope of the valley from the river bottom
(Station IV, c) to the upland forest (Station IV, a) and the side of
the south ravine, the bottom of which forms Station IV, d. This sub-
station is not as homogeneous physically as the upland or lowland for-
est, because the part along the south ravine is relatively open, is well
drained, and has a south exposure, and the southeast slope to the low-
60
land forest on the other hand, is well wooded and shaded, and much
more humid. The substation also has a considerable amount of litter,
leaves, and humus. This region may be considered as transitional be-
tween the upland and lowland forest, but it represents, not one but two
transitional stages, the south slope approaching the upland forest type,
and the southeast slope approaching that of the lowland forest.
Thus, if one walked from the upland forest down the slope of the
south ravine, and eastward to the southeast valley slope to the bottom-
land forest, he would traverse all the main degrees of conditions found
at Station IV.
The forest cover consists primarily of the following trees: white
oak (Quercus alba), black oak (Q. velutina), walnut (Juglans nigra),
pignut (Carya glabra), and, in smaller numbers, mulberry (Morus
rubra), red oak (Quercus rubra), shag-bark hickory (Carya ovata),
bitternut (C. cordiformis) ; and of the following shrubs: redbud (Cer-
dis canadensis), sassafras (Sassafras variifolium), moonseed (Menis-
permum canadense), five-leaved ivy (Psedera quinquefolia), grape
(Vitis cinerea), prickly ash (Zanthoxylum americanum), and sumac
(Rhus glabra), the latter growing in large colonies on the open south
ravine-slope. On the more moist and shaded southeast slope lived the
clearweed (Pilea pumila), a plant quite characteristic of moist deep-
shaded woods. Thus sumac and clearweed may be Considered as in-
dex plants to the physical conditions in different parts of these two
slopes, one shaded and the other rather open.
The atmometer, located on the upper part of the south ravine slope,
gave a relative humidity of 31 per cent. of the standard in the garden
of the Normal School. It will be recalled that in the upland forest
(Station IV,a) the atmometer gave 54 per cent., the comparison
showing how much less the evaporating power of the air is on the
south ravine slope than it is in the upland forest. The relative evap-
oration was not determined for the southeast slopes, but the presence
of Pilea clearly indicates that it is less than on the south ravine slope,
where the instrument was located. On the lower parts of the valley
slope, where this substation grades into the lowland, the layers of dead
matted leaves and humus reached to a considerable depth, and looked
as if they had been pressed down by drifting snows. Such places were
found to contain very few animals.
This habitat is characterized by a sloping surface, by relative open-
ness on the ravine side and dense shade on the valley slope, by rela-
tively humid air, by second-growth forest somewhat transitional be-
tween that of the uplands (Station IV, a) and the river bottoms (Sta-
tion IV, c), by a relatively large amount of shrubbery, by considerable
61
humus and litter, by moist soil, and by more logs and stumps than are
in the upland forest.
The collections of animals made at this substation (Nos. 68, 84, 85,
87, 89, 90, 94, 100, 104, 105, 106, 108, 110, III, 124, 125, 131, 132,
133, 140, 149, 161, 164, 165, 166, and 168) are as follows:
Land snail
Land snail
Land snail
Land snail
Carolina Slug
Land snail
Milliped
Milliped
Stout Harvest-spider
White Ant
Woodland Cockroach
Green Short-winged
Grasshopper
Boll’s Grasshopper
Scudder’s Grasshopper
Woodland Cricket
Caterpillar-hunter
Wireworm
Horned Passalus
Tenebrionid larva
Troilus Butterfly
Philenor Butterfly
Lyczenid butterfly
American Silkworm
Hickory Horned-devil
Arctiid caterpillar
Rotten-log Caterpillar
Notodontid
Notodontid larva
Geometrid
Slug Caterpillar
Pigeon Tremex
(Acorn Plum-gall)
Old-fashioned Ant
Tennessee Ant
Formicid ant
Polygyra clausa
Vitrea indentata
Vitrea rhoadsi
Zonitoides arborea
Philomycus carolinensis
Pyranidula perspectiva
Cleidogona cesioannulata
Polydesmus sp.
Liobunum grande
Termes flavipes
Ischnoptera sp.
Dichromorpha viridis
Spharagemon boll
Melanoplus scudderi
Apithes agitator
Calosoma scrutator
Melanotus sp.
Passalus cornutus
Meracantha contracta
Papilio troilus
Papilio philenor
Everes comyntas
Telea polyphemus
Citheroma regalis
Halisidota tessellaris
Scolecocampa liburna
Datana angusit
Nadata gibbosa
Caberodes confusaria
Cochlidion or Lithacodes
Tremex columba
Amphibolips prunus
Stigmatomma pallipes
Aphenogaster tennesseensis
Myrmica rubra scabrinodis
schnecki
140,
89,
84,
100,
68,
163,
164
164
164
84
125
164
140
125
III
125
140
110
133
124
124
149
125
85
140
161
166
I61
163
108
168
125
104,
94
161
165
132
131
140
87
140
62
Carpenter-ant Camponotus herculeanus penn-
sylvanicus 84, 85
Rusty Carpenter-ant Camponotus herculeanus penn-
sylvanicus ferrugineus 90
Short Caterpillar-wasp Ammophila abbreviata 124
4. Lowland or “Second Bottom,’ Red Oak-Elm-Sugar Maple Wood-
land Association, Station IV, ¢
This station includes the part of the forest located upon the upper
or higher part of the river bottom. This area is sometimes called the
“second bottom” because it is above the present flood-plain. The gen-
eral position of the forest is shown in Figure 1, Plate X. The fringe
of willows along the river bank is shown at a, the flood-plain area is
cleared at b; the substation forest is at c; and part of the forest of the
valley slope is seen at d. Other views of this station are shown in
plates XIV, XV, and XVI (figures 1 and 2). The general slope is
toward the river; minor inequalities are due to the action of the tem-
porary streams which are etching into the uplands and depositing their
burdens of debris at the mouths of the ravines. Soil, leaves, and other
organic debris are washed from the upland, the ravines, and the val-
ley slopes, and are deposited upon the bottoms, forming low alluvial
fans, which have been built up in successive layers or sorted again and
again as the temporary streams have wandered over the surface of
the fan on account of the overloading and. deposition which filled up
their channels. In this manner the soil in general is not only supplied
with moisture, drained from the upland, but the various soils are both
mixed as successive layers of organic debris are buried by storms and
also mulched by the large amount of this debris which is washed and
blown to the lowland. No springs were found upon the southeast
valley slope, but in the south ravine pools of water were present dur-
ing August, 1910, when my observations were made.
The forest, characterized by hard maple (Acer saccharum), red
oak (Quercus rubra), and elm (Ulmus americana), forms a dense
canopy which shuts out the light and winds, thus conserving the mois-
ture which falls and drains into it, and making conditions very favor-
able to a rich mesophytic hardwood forest. That the relative humid-
ity is high is shown by the moisture found in the humus of the forest
floor, and, further, not only by the presence of clearweed (Pilea pu-
mila) and the nettle Laportea canadensis, which characterize such
moist shady woods, but also by the presence of the scorpion-flies (Bit-
tacus). ‘These organisms are permanent residents where such condi-
63
ditions prevail, and their presence is as clearly indicative of certain
physical conditions as that of aquatic animals. would be indicative of
other physical conditions. In addition to these evidences we have
the readings of our atmometer, which showed the evaporating power
of the air to be 26 per cent. of the standard in the garden at the Normal
School. This shows that the relative evaporation is very low, .and
that conditions for the preservation of the moisture which falls and
drains into this area are very favorable. The general character of this
forest is shown in plates XIV, XV, and XVI, Figure 1.
The vegetational cover on the lowland is quite different in its com-
position from that on the upland. This is shown mainly by the pres-
ence of the elm (Ulmus americana), hard maple (Acer saccharum),
and red oak (Quercus rubra), and secondarily, by the presence, in
smaller numbers, of the black cherry (Prunus serotina), slippery elm
(Ulmus fulva), shingle oak{ (Quercus imbricaria), and the Kentucky
coffee-tree (Gymnocladus dioica). Other trees present are walnut
(Juglans nigra), mulberry (Morus rubra), and bitternut (Carya cor-
diformis). The shrubs and.vines are gooseberry (Ribes cynosbati),
prickly ash (Zanthoxylum americanum), redbud (Cercis canadensis),
buck-brush (Symphoricarpos orbiculatus), green brier (Smilax),
five-leaved ivy (Psedera quinquefolia), moonseed (Mentspermum
canadense), bittersweet( Celastrus scandens), and grape (Vitis cine-
rea). "The characteristic herbaceous vegetation is nettle (Laportea
canadensis), clearweed (Pilea pumila), bellflower (Campanula ameri-
cana), Indian tobacco (Lobelia inflata), tick trefoil (Desmodium
grandiflorum), Actinomeris alternifolia, maiden hair fern (Adiantum
pedatum), beech fern (Phegopteris hexagonoptera), the rattlesnake
fern (Botrychium virginianum), and Galium circesans and G. tri-
folium.
Although the forest is generally dense and therefore deeply shaded,
there are some places which are comparatively open. Attention, how-
ever was devoted mainly to the denser parts. At one place, near the
base of the eastern slope of the valley, a few trees had been cut within
a few years, and in this glade the conditions and plants and animals
were different from those in the dense forest. (See Pl. XVI, figs. 1
and 2.
ids habitat may be characterized as follows: lowland densely cov-
ered by sugar maple-red oak forest (climax mesophytic) ; very humid
air; a moist soil; relatively few shrubs; herbaceous plants—nettles and
clearweed—characteristic of damp, shady, rich woods; and considera-
ble litter and humus in places.
64
The collections of animals made here (Nos. 113, 114, 116, II7,
137-139, 141, 143, 144, 173, 182, and 184) are as follows, the itali-
cised numbers designating collections from the glade:
Predaceous Snail
Land snail
Slug eggs
Alternate Snail
Milliped
Ambush Spider
Tent Epeirid
Three-lined Epeirid
Spined Spider
Rugose Spider
Ground Spider
Cherry-leaf Gall-mite
Clear-winged Scorpion-fly
Leaf-hopper
Pentatomid
Coreid
Spined Stilt-bug
Short-winged Grasshopper
Acridiid grasshopper
Acridiid grasshopper
Scudder’s Grasshopper
Round-winged Katydid
Nebraska Cone-nose
Meadow Grasshopper
Meadow Grasshopper
Meadow Grasshopper
Striped Cricket
Elaterid larva
Elaterid
Black-tipped Calopteron
Reticulate Calopteron
Horned Fungus-beetle
Common Skipper
Imperial Moth (larva)
Noctuid moth
Asilid fly
Vespa-like syrphid
Long-sting
Black Longtail
Cocoanut Ant
Circinaria concava
Vitrea indentata
Philomycus (?) eggs
Pyramidula alternata
Callipus lactarius
Misumena aleatoria
Epeira domiciliorum
Epeira trivittata
Acrosoma spinea
Acrosoma rugosa
Lycosa scutulata
Acarus serotine
Bittacus stigmaterus
Aulacizes irrorata
Hymenarcys nervosa
Acanthocerus galeator
Jalysus spinosus
Dichromorpha viridis
Melanoplus amplectens
Melanoplus gracilis
Melanoplus scudderi
Amblycorypha rotundifolia
Conocephalus nebrascensis
Orchelimum cuticulare
Orchelimum glaberrimum
Xiphidium nemorale
Nemobius fasciatus
Corymbites sp.
Asaphes memnonius
Calopteron terminale
Calopteron reticulatum
Boletotherus bifurcus
Epargyreus tityrus
Basilona imperialis
Autographa precationis
Deromyia discolor
Milesia ornata
Thalessa lunator
Pelecinus polyturator
Tapinoma sessile
II7,
rr7;
117,
II7,
Lig,
iy,
143,
II7,
113
113
114
173
113
184
173
138
172
172
I44
116
141
143
113
182
II7
143
143
143
I17
143
117
143
143
I43
143
113
113
173
143
173
173
106
143
117
184
I43
43
139
65
5. Supplementary Collections from the Bates Woods, Station IV
Tent Epeirid
White-triangle Spider
Spined Spider
Rugose Spider
Mealy Flata
Leaf-hopper
Pentatomid bug
Pentatomid bug —
Tarnished Plant-bug
Coreid bug
Coreid bug
Rapacious Soldier-bug
Acridiid grasshopper
Pennsylvania Firefly
Margined Soldier-beetle
Soldier-beetle
Chrysomelid beetle
Clubbed Tortoise-beetle
Portlandia Butterfly
Eurytus Butterfly
Gelechiid moth.
(Hairy Midge-gall)
Corn Syrphid Fly
( Horned-knot Oak-gall)
(Oak Wool-gall)
Ichneumon Wasp
Formicid ant
Rusty Carpenter-ant
Spider Wasp
Epeira domiciliorum
Epeira verrucosa
Acrosoma spinea
Acrosoma rugosa’
Ormenis pruinosa
Gypona pectoralis
Euschistus fissilis
Mormidea lugens
Lygus pratensis
Alydus quinquespinosus
Acanthoceros galeator
Sinea diadema
Melanoplus obovatipennis
Photuris pennsylvanica
Chauliognathus marginatus
Telephorus sp.
Cryptocephalus mutabilis
Coptocycla clavata
Enodia portlandia
Cissia eurytus
Ypsolophus ligulellus
Cecidomyia holotricha
167
126
148
126
Hankinson
Hankinson
124
Hankinson
Hankinson
Hankinson
Hankinson
Hankinson
124
Hankinson
Hankinson
Hankinson
Hankinson
Hankinson
63
Hankinson
Hankinson
(Near collection No. 96)
Mesogramma politum
Andricus cornigerus
Andricus lana
Trogus obsidianator
Aphenogaster fulva
Hankinson
(Near 96)
(Near 96)
Hankinson
125
Camponotus herculeanus penn-
sylvanicus ferrugineus
Psammochares ethiops
97
Hankinson
6. Small Temporary Stream in the South Ravine, Station IV, d
This small temporary stream in a ravine formed the southern
boundary of the area examined (PI. XVII, figs. 1 and 2). At the sea-
son of our examination it was a series of small disconnected pools.
Very little attention was devoted to the collection and study of its life.
Most of the collections were secured by T. L. Hankinson. A few aquat-
ic animals were collected here.
In a small pool were taken numerous
specimens of the creek chub (Semotilus atromaculaius), and one stone-
66
roller (Campostoma anomalum). Frogs, toads, and salamanders were
also taken in the vicinity by Mr. Hankinson, who dug from their bur-
rows specimens of Cambarus diogenes, and also secured immunis and
propinquus. On the surface of the pools were numerous specimens
of a water-strider, Gerris remigis. The forest cover is undoubtedly an
important factor in the preservation of such pools, as it controls the
evaporating power of the air.
Mr. Hankinson tells me that during the summer of 1912 this tem-
porary stream was completely dry, and that no fish have been taken
from it since the earlier collection mentioned above. From the mouth
of the ravine across the bottom to the river it is only a few hundred
feet, and in time of heavy or prolonged rains these pools are in direct
communication with the river. Such a stream is an excellent example
of an early stage in the development of the stream habitat, and shows
its precarious character, and the liability to frequent extermination
of these pioneer aquatic animals which invade it in its early stages.
This -applies particularly to those animals which have no method of
tiding over dry periods. On the other hand, those animals which live
in the pools, those parts of temporary streams which persist longest
between showers, have better chances of survival, particularly bur-
rowing animals, like the crawfish and its associates. It seems prob-
able that crawfish burrows harbor a varied population; not only the
crawfish leeches (Branchiobdellide) but also the eggs of certain Cor-
ixide (Forbes, ’76: 4-5; 778, p. 820; Abbott, 12) may almost cover
the body of some crawfishes. By means of this burrow ground-water
is reached, and a subterranean pool is formed. For ihe elaboration of
the stream series see Adams (’01) and Shelford (’11 and 13a).
This temporary stream shows how, by the process of erosion, the
upland forest area is changed into ravine slopes, and, later, even into
the bed of a temporary stream. ‘Thus progresses the endless transfor-
mation of the habitat.
GENERAL CHARACTERISTICS OF THE GROSS
“ENVIRONMENT
1. Topography and Soils of the State
Illinois lies at the bottom of a large basin. This is indicated in
part by the fact that so many large rivers flow toward it. The mean
elevation of the state is about 600 feet, and about a third of it lies be-
tween 600 and 700 feet above sea-level. Except Kentucky, the bor-
dering states are from 200 to 500 feet higher. Iowa and Wisconsin
are considerably higher, so that winds from the north and northwest
67
reach the state coming down grade. Taken as a whole the land sur-
face is a tilted plain sloping from the extreme northern part—where a
few elevations exceed a thousand feet—toward the south, bowed in
the central part by a broad crescentic undulation caused by a glacial
moraine, and then declining gradually to the lowland north of the
Ozark Ridge, near the extreme southern part of the state. This east
and west ridge occasionally exceeds 1,000 feet, but its average height
is between 700 and 800 feet. It is very narrow, only about 10 miles in
average width, and rises about 300 feet above the surrounding low-
land (Leverett, ’96, ’99). South of this ridge lie the bottoms of the
Ohio River. The largest river within the state is the Illinois.
The soils of the state are largely of glacial origin. Even the un-
glaciated extreme northwestern part and the Ozark Ridge region have
a surface layer of wind-blown loess. In some places considerable sand
was assorted by glacial water, forming extensive tracts of sandy soil,
and locally dune areas are active. Along the larger streams there are
extensive strips of swamp and bottom-land soils. The remaining soils,
which characterize most of the state, were either produced mainly by
the Iowan or IJlinoian ice-sheets, as in the case of the relatively poorer
soils, or by the Wisconsin sheet, which formed the foundation for the
better soil. The dark-colored prairie soils are due to organic debris.
Coffey (’12: 42) has said: “Whether this accumulation of humus is
due to lime alone or to the lack of leaching, of which its presence is an
indication, has not been definitely determined. Neither do we know
whether it is due to chemical or bacteriological action; most probably
the latter, an alkaline medium being necessary for the growth of those
bacteria or other microorganism which cause this form of decomposi-
tion.””*
2. Climatic Conditions
The climatic features of a region are generally conceded to have a
fundamental influence upon its life. The controlling influences upon
climate are elevation above sea-level, latitude, relation to large bodies
of water—generally the sea—and the prevailing winds. The eleva-
tion and relief of Illinois have but a slight influence. In latitude
Illinois is practically bisected by the parallel 3934° in the north tem-
perate zone. This position influences the seasons and the amount of
heat received from the sun. The sea is far distant, but the Great
‘Lakes are near by, and proximity to the interior of a large continent
*Consult Hopkins and Pettit (’08) and the County Soil Reports of the State
Soil Survey for a'detailed account of the chemical conditions of Illinois soils.
The bacterial, algal, and animal population have hardly been noticed by stu-
dents of Illinois soils. :
68
brings the state within that influence. And, finally, it lies in the zone
of the prevailing westerly winds, and directly across the path of one
of the main storm tracks, along which travel in rapid alternation the
highs and lows which cause rapid changes of temperature, wind, and
precipitation, and thus produce the extremely variable weather condi-
tions.
The state is 385 miles long, and as latitude has much influence
upon climate, the climate of Illinois differs considerably in the extreme
north and south. This is clearly shown in the average annual tempera-
ture, which in the northern part is 48.9° F., in the central part is
52.70°, and in the southern part is 55.9° (Mosier, ’03). These aver-
ages probably. closely approximate the soil temperatures for these re-
‘gions. The average date of the last killing frost in the northern part
is April 29; in the central part, April 22; and in the southern part,
April 12. The average date of the first killing frost for the northern
part is October 9, central part, October 11, and the southern part is
October 18 (Henry). The growing season for vegetation in the
northern half of the state averages from 150 to 175 days and for the
southern half from 175 to 200 days (Whitson and Baker, 12: 28).
The precipitation shows similar differences, increasing from north to
south. The annual average for the northern part is 33.48 inches, in-
creasing to 38.01 in the central and to 42.10 inches in the southern
part (Mosier, 03:62). Mosier has shown that the Ozark Ridge,
with an average elevation of about 800 feet, condenses the moisture
on its south slope so that it has a precipitation of 7.15 inches more
than do the counties just north of the ridge. This same humid area
appears to extend up the Wabash Valley to Crawford county, and
gives the valley counties a rainfall 3 inches in excess of the adjacent
counties to the west. The average annual rainfall for the state is
37.39 inches—nearly one third of it during April, May, and June,
and if July is included, more than half. The heaviest precipitation,
8.23 inches, is in May and June.
As previously mentioned, the state lies in the zone of prevailing
westerly winds and across the path of storms. These have a dominant
influence upon the direction of the winds. In the northern part of the
state, they are, by a slight advantage, southerly—a tendency which
progressively increases toward the south, for in the central part the
southerly winds reach 55 per cent., and in the southern part 62 per
cent. During the winter the northwest winds predominate throughout
the state, to a marked degree in the central part, where they reach
60 per cent., and where also the velocity is greatest, reaching an av-
erage of 10.3 miles an hour. The velocity of the wind for the entire
69
state is highest during spring. During the summer, the southwest
winds predominate in the northern and central parts, and in the south-
ern part 82 per cent. of the winds are southerly. The velocity of the
wind is least during the summer, and the greatest stagnation occurs
in August. During autumn there is a falling off of the southerly
winds and an increased velocity as winter conditions develop. The
transition in the fall is in marked contrast with the vigor of the
spring transition. The cooler seasons are more strongly influenced
by northerly winds, and the warmer seasons by southerly winds.
3. Climatic Centers of Influence
In the preceding section the average conditions of temperature,
precipitation, and the direction and velocity of the winds have been
summarized, but little effort was made to indicate the mode of opera-
tion of the determining factors which produce and maintain these aver-
age conditions. It is often true that the main factors which explain
the conditions seen in some restricted locality can not be found within
it because the local sample is only a very small part of a much larger
problem. Thus no one attempts to find an explanation of the through-
flowing upper Mississippi system within the state of Illinois; a larger
unit of study is necessary. The region examined must extend to the
headwaters. So, also, with most of the climatic features of Illinois;
their approximate sources must be sought elsewhere. Let us there-
fore consider some of the broader features which influence the climate
of North America, particularly that of the eastern part.
The climates of the world have been divided into two main kinds,
depending primarily upon the controlling influence of temperature.
This is due to the relative specific heat of land and water, that of water
being about four times that of land. The sea, which covers three
fourths of the earth’s surface, is thus an immense reservoir of heat,
which is taken up and given off slowly, at a rate one fourth that of the
land. It is therefore relatively equable. The northern hemisphere
contains the largest amount of land, and is therefore less under the
control of the sea than the southern hemisphere; yet the sea’s influence
is very powerful, particularly near the shore. The large land masses,
on the other hand, on account of their lower specific heat, receive and
give off heat more rapidly to the air above. For this reason the tem-
perature changes, as between day and night or summer and winter,
are much more rapid and much more extreme over land than over
the sea. A climate dominated by the equable sea is oceanic; that
dominated by the changeable lands is continental. [Illinois lies far
70
from the sea and is therefore strongly influenced by continental con-
ditions. To what degree is the marine influence shown?
Meteorologists (cf. Fassig, ’99) have come to look upon the large
areas of permanent high and low barometric pressure as among the
most important factors in climatic control. There are five of these
powerful “centers'of action” which influence our North American
climate (Fig. 1), and four of these are at sea. A pair of Jows are in
the far north, one in the north Pacific near Alaska, the other in the
80 60° 40 20°
aw, aes
pes
oon NG
Owe
| ¢
seen Bs
-
Lea
160° 40° 120° 100° 80° 60° 40 2
Fig. 1. Diagram showing the positions of the relatively stable areas of high and
low barometic pressure, and indicating their influences upon the evaporating power of
the air and upon the climate in general.
north Atlantic south of Greenland. A pair of highs are farther south,
one in the Pacific between California and the Hawaiian Islands, and
the other centering in the Atlantic near the Azores. The highs and
lows in each ocean seem to be paired and to have some reciprocal rela-
tion. The fifth center of action is upon the land. It is a high baromet-
ric area in the Mackenzie basin of Canada, where it becomes a pow-
erful center‘of influence through winter and spring, but with the prog-
ress of summer conditions weakens, and through the accumulation of
continental heat becomes converted into a low, thus there is a complete
seasonal inversion on the continent.
These large highs and lows, although relatively permanent, are con-
tinually changing in intensity and position. The highs are regions of
descending, diverging, warming, and drying air, producing clearing
and clear air on their western side, but the reverse on their eastern side.
71
The lows are regions of ascending, converging, cooling air, with in-
creasing moisture and clouds on their western side, but are the re-
verse on their eastern side (Moore, ’10: 153). ‘These same character-
istics apply to the small highs and lows which we are accustomed to
see on the daily weather maps.
If, now, we consider these large centers of action, such considera-
tion will do much toward giving us a graphic idea of our climate. Dur-
ing the winter, because of the small amount of heat received in the
Mackenzie basin, the temperature becomes very low, and a powerful
high barometric area is formed; then the descending air blowing from
the eastern part of this high, or from small highs originating from the
larger one, produce the cold winters and cold waves in winter which
characterize the northeastern United States. If, however, the Atlantic
high wanders on the eastern coast of the United States in winter, the
western part of this high, with its descending, diverging, warming, and
drying air, produces a mild winter. The climate of the eastern United
States is thus, in the cold season, under the alternate invasion of these
two powerful centers of action. During the warm season the conti-
nental winter high is replaced by a low, due to the accumulating warm
continental temperatures which thus have produced an inversion or
seasonal overturning. But the Atlantic high is permanent and exerts
its influence continuously. If the western part of this high encroaches
upon the eastern United States during the summer, with its descend-
ing, drying, and clear air, it may produce drouth, this depending, of
course, on its degree of development. The continental low of sum-
mer, with the drying influence of its eastern side, has a similar ten-
dency. Thus the character of the summer is determined, to an im-
portant degree, by the interplay and relative balance between these two
warming and drying centers. ‘The activity of these centers has a pow-
erful influence upon the moisture-bearing winds, which influence hu-
midity and evaporation in Illinois, and in the eastern United States.
4. Relative Humidity and Evaporating Power of the Air
We are now in a position to examine the facts of relative humidity
and the relative evaporating power of the air in the eastern United
States. The relative aridity on the plains east of the Rocky Moun-
tains is due primarily to the removal of moisture from the prevailing
westerlies in their passage from the Pacific over the various western
mountain ranges which extend across their path, combined with the
excessive summer heating of the continental mass. Here, then, is the
influence of the continental summer low. Farther east the Atlantic
high tends to supplement the continental low and to cause the Gulf
72
winds to brings moisture inland,* and the Great Lakes region adds its
uota.
: In the storm-track zone, where stagnation of the air is due largely
to the balance existing between the continental low and the oceanic
high, the aridity of the plains extends the farthest east, and as an arid
peninsula it crosses Illinois, giving during August a relative humidity
to the prairie area of 60-70 per cent. of saturation (Johnson, ’07).
The reality of the arid peninsula across Illinois is further shown by
the rainfall-evaporation ratios computed and mapped by Transeau
(’o5). These ratios were determined by dividing the mean annual
rainfall at each place by the total mean annual evaporation. These
mapped percentages show that the prairie region is closely bounded
by the region with an evaporation ratio of between 60 and 7o per
cent. of the rainfall received. These conditions furnish a general
background or perspective for a profitable consideration of the local
and more detailed studies which have been made of the relative evap-
orating power of the air in different plant and animal habitats.
For our purpose it is not necessary to consider the history of meth-
ods of measuring relative evaporation. ‘This measurement may be
made by evaporating water in open pans or by the porous porcelain-cup
method. Such cups have been devised by several students, but a modi-
fied form of the Livingston atmometer has been mainly used by plant
ecologists, and this was the kind we used at Charleston. Transeau
(’08) was the first to use such an instrument and to show its value in
studying the relation of intensity-of evaporation to plant societies.
His work on Long Island, N. Y., showed very clearly that evaporation
in open places was much greater than in dense forests. These obser-
vations were enough to show that evaporation is a factor related to the
physical conditions of life upon the prairie and in the forest, and there-
fore in our cooperative study of the Charleston area in 1910 relative
evaporation was made a special feature in the study of representative
environments, in order to determine its relation to both the plants and
the animals. So far as is known this is the only study yet made in
which these determinations have been recorded from the same places
where the animals have been studied. Since our data were secured,
several papers have been published on relative evaporation in different
sorts of habitats in this state and in northern Indiana by plant ecolo-
gists Fuller (’11, ’12a, ’12b), McNutt and Fuller (’12), Fuller, Locke,
*Zon (’13) has recently asserted that the moisture from the sea does not
-make a single overland flight inland, but rather is largely precipitated near the
sea, is evaporated and carried farther inland, is precipitated again, and this
process repeated again and again, so that its inland flight is a vertical revolv-
ing eycle of precipitation and evaporation. If this contention is valid, evapo-
ration from the land is a much more important climatic factor than it is usually
thought to be.
73
and McNutt (’14), Sherff (’12, ’13a, ’13b), and Gleason and Gates
(12). Shelford (’12, ’13a, ’13b, “14a), utilizing the evaporation
data of the plant ecologists, has applied the same to animal associa-
tions also, and he has further tested some of these ideas experiment-
ally in the laboratory. In Ohio, Dachnowski (’11) and Dickey (’09)
have made records of data obtained by the use of the porous cup, and
in Iowa Shimek (’10, ’11) has used the open-pan method. Mention
should also be made of Yapp’s observations (’09) on a marsh in Eng-
land. A very important summary of evaporation records, in the open
and in forests, is given by Harrington (’93). The effect of wind-
breaks upon evaporation has been studied by Bates (’11) and Card
(97). Finally, mention should be made of Hesselman’s studies of
relative humidity in forest glades in Sweden (’04).
Our records from the Charleston region will be given first, and then
their significance will be discussed. The unglazed porcelain cups, with
a water reservoir, were placed so that the tops of the cups were about
six inches above the soil in the habitats examined, and at weekly in-
tervals the water loss was measured. The instruments were in opera-
tion simultaneously, so that the results are comparable. The standard
instrument was located in the open exposed garden of the Eastern
Illinois Normal School at Charleston, which was considered as unity,
or 100 per cent. For further details as to the conditions where the
atmometers were located consult the description of the stations and
the photographs.
An examination of the diagram (Fig. 2) will show that although
‘based upon a limited amount of data (for less than a month, from
10 20 30 40 50. 60 70, 80 90 i00
Intensity of evaporation..........+.
Standard, open garden, Normal School
Sta. III, b. Mixed prairie and young forest
Sta. II, a. Grassy area, Panicum
Sta. II, uw. Grassy area, Euphorbia
Sta. IV, w. Upland, open woods
Sta. III, a. Silphium on Diack soil
Sta. II, a. Colony of 8. laciniatum
Sta. IV, b. Ravine slope, open woods
Sta. IV,c. Dense climax forest cover
Fie. 2, Diagram of the relative evaporation in different prairie and forest
habitats, showing the great reduction in evaporation with the development of a closed
forest canopy of a climax forest; Charleston, Illinois.
74
August 19 to September 22) the facts are in harmony with similar
studies elsewhere covering a much longer period, so that there is valid
reason for confidence in them. The standard instrument was located,
as already mentioned, in an open, exposed cultivated garden, where the
intensity of evaporation was very high. The black soil prairie areas,
Stations II and III, a, have an average of 56.1 per.cent.—a condition
much like that in the grassy-Euphorbia prairie at Loxa (Station II, a)
—or a little more than half that of the standard instrument. The dry
upland area of mixed prairie and young forest, on gray silt loam (Sta-
tion III, b), has an intensity of 80 per cent. This is in the region of
the most extensive grassy prairie about Charleston; the general ap-
pearance of the region is shown in Plate XIII. A surprising feature
of the table is the evaporation in the open-crowned upland oak-hickory
woods (Station IV, a). In this forest perhaps two thirds to three
fourths of the ground was shaded, and it was very well drained. The
evaporation here reached 54.2 per cent., being very near that of the
average of the black soil prairie (56.1 per cent.). I had anticipated
much less evaporation than on the prairie, a position more intermedi-
ate between the prairie and the lowland forest, or about 42 per cent.
(cf. Harvey, 14:95). The ravine slope (Station IV, Db), although
somewhat open, has 31.5 per cent.—a very low rate of evaporation—
and is remarkably close to that of the densely crowned lowland for-
est (Station IV, c), at 26.9 per cent. The decline, however, in the
intensity of evaporation with the degree of completeness of the for-
Per cent. of standard............+-- 0 40 60 80 (00 120
Sta. 11. Salt marsh outer margin
Sta. 3. Gravel slide, open
Sta. 1. Carnegie garden, standard
Sta. 9 and 10. Upper beach
Sta. 12. Salt marsh, inner margin
Sta. 2. Garden, high level
Sta. 4. Gravel slide, partly invaded
Sta. 5. Forest, open
Sta. 13. Fresh-water marsh
Sta. 6. Forest, typical mesophytic
Sta. 7. Forest, ravine type
Sta. 8. Forest swamp type
Fic. 3. Diagram of the relative intensity of evaporation in the lowest stratum
of different kinds of habitats, Long Island, N. Y. (After Transeau.)
75
est crown, is strikingly shown in passing from the open upland
woods, at 54.2 per cent., to the ravine slope at 31.5 per cent., and on
to the lowland forest at 26.9 per cent.
A comparison of these results with those secured by Transeau
(08) on Long Island, is instructive. His standard instrument was
also in an open garden (Fig. 3), comparable with the Charleston
standard. A gravel slide, partly invaded by plants, had an evaporation
of 60 per cent., comparable with the open prairie at Charleston; the
open forest, 50 per cent., comparable with the upland open Bates
woods at 54.2 per cent.; and the mesophytic forest, 33 per cent., com-
parable with the ravine and lowland places in the Bates woods at 31.5
and 26.9 per cent. respectively.
Association
Blowout (basin) 1.56
Blowout (slide) 1.27
Bunchgrass (Leptoloma consoc.) 1.18
Bunchgrass (Eragrostis trichodes con.) 1.04
Standard 1.00
Beach 0.93
Quercus velutina woods 0.66
Quercus velutina 0.55
Willows (Acer part) 0.56
Willows (Salix part) 0.44
Mixed forest (margin) 0.36
Mixed forest (center) 0.29
Fig. 4. Relative intensity of evaporation in different kinds of habitats on sandy
soil, Havana, Illinois. (After Gleason and Gates.)
Another series of relative evaporation observations was made by
Gleason and Gates (’12) on sandy soils at Havana, Illinois. As their
methods were similar to those used at Charleston, useful comparisons
may again be made. The standard instrument was in an open area
comparable to the garden at Charleston. An examination of Figure 4,
summarizing the results of their study, shows that upon the grass-
covered sand prairie (bunch-grass) the evaporation was about 110 per
cent., that in open black oak (Q. velutina) woods (on sand) it was
about 60 per cent., and that in a denser hickory-black-oak-hackberry
mixed forest (somewhat open) it was about 31 per cent. There is thus
a close general correspondence between the conditions at Havana and.
Charleston, although the evaporation upon sand prairie appears to be
relatively much greater than upon the black-soil prairie.
Fuller (’11) and McNutt and Fuller (’12) have made comparative
studies in different kinds of forest in northern Illinois and in northern
76
Indiana. Their results are combined and summarized in Figure 5.
This diagram shows the relative evaporation near the surface of the
soil, the standard of comparison being the evaporation im a maple-
beech climax forest, where evaporation is relatively low. The aver-
age daily amount, in c.c., shows that there is a progressive increase in
evaporation as follows: 8.1 c.c. in a maple-beech forest, 9.35 c.c. in
the oak-hickory upland forest, 10.3 c.c. in an oak dune forest, 11.3 c.c.
in a pine dune forest, and an increase to 21.1 c.c., on the cottonwood
dunes. This expressed on a percentage basis is, in inverse order, re-
spectively 260 per cent. in the cottonwoods, 140 per cent. in the pines,
127 per cent. in the oak dunes, 115 per cent. in the oak-hickory for-
est, and 100 per cent. in the maple-beech forest.
20.40 60 @0 100 (120 (140 (160180 200 22Q_ 240 260 280
Intensity of evaporation
Sta. A. Cottonwood dunes
Sta. B. Pine dune
Sta. C. Oak dune
Sta. D. Oak-hickory
Sta. E. Maple-beach forest % Standard
Fic. 5. Diagram showing the relative rate of evaporation in different kinds of
forest in northern Illinois and Indiana. [Data from Fuller (’11) and McNutt and
Fuller (’12).]
Shimek (10, ’11) has made valuable observations on the relative
rate of evaporation on the prairie of western Iowa. He used the open-
pan method in four representative habitats. His results show very
clearly that the rate of evaporation is much greater in exposed places
than where there is shelter from the sun and wind. I have put his
data in a form comparable with those which have just been discussed
(Fig. 6), and have made the cleared field area, Station 4, the standard
of comparison, as it more nearly approaches the standard used at
Charleston and by others. Station 3 is on a high bluff, exposed to the
Intensity of evaporation............ . 2040. 6080100120, 40160 180-200
Sta. 3. Open, much exposed prairie
vegetation
Sta. 1. Open, exposed slope of bluff,
prairie
Sta. 4. Open, cleared area, partly pro-
tected
Sta. 2. Bur-oak grove, protected
Fic. 6. Diagram of relative evaporation in prairie and forest habitats, in western
Iowa. (Data from Shimek.)
77
west and south winds, and, as might be expected, it has an excessive
evaporation—184 per cent. Station 1, also covered by prairie vegeta-
tion, and exposed to west and southwest winds but sheltered from
winds from the south and southeast, also shows a very high evapora-
tion—132 per cent. Station 4, which was made the standard, had been
cleared of forest, and was an open place protected by a ridge. Station
2 was apparently a dense grove composed of bur oak, basswood, elm,
and ash, with considerable undergrowth. Here the rate of evapora-
tion dropped considerably—to 36 per cent. The general character of
this forest calls to mind the denser oak forests on sand at Havana,
Illinois. An important feature of these observations is that they were
made far out upon the “prairie”, bordering the plains, most other
studies on relative evaporation having been made much farther east.
In Ohio, Dachnowski (’11) and Dickey (’09) have recorded the
relative evaporation of the air, using a campus lawn as unity. In the
central grass-like area of a cranberry bog the evaporation was 69.2
per cent., and in the marginal maple-alder forest it was 51.2 per cent.
Harrington (’93: 96-102), in summarizing European studies on
the relative evaporation (with a water-surface as standard) in the
open and in German forests shows that the “annual evaporation in the
woods is 44 per cent. of that in the fields.” Compared with evapora-
tion in the open, that under deciduous trees is 41 per cent., and that
under conifers is 45 per cent.—a difference most marked in the sum-
mer. Ebermeyer’s Austrian observations (I. c. :99) show that the
“evaporation from a bare soil wet is about the same as that from a
water surface,” both in the open and in the forest. A saturated soil
under forest litter gives an evaporation of only 13 per cent. of that
of a free-water surface in the open. Harrington (l.c.: 100) con-
cludes that “About seven-eighths of the evaporation from the forest
is cut off by the woods and litter together.” Sherff (’13a, ’13b) has
shown that in the Skokie Marsh, north of Chicago, the absolute
amount of evaporation near the soil was less at the center of a Phrag-
mites swamp than at its margin (Fig. 7), that a swamp meadow
Intensity of evaporation.........-+++ 20.40 60-80 _—100-— 120-40 S0_—*180 200
Sta. D. White oak-ash forest
Sta. B. Phragmites swamp, margin
Sta. OC. Swamp meadow Standard
Sta. A. Phragmites swamp, center
Fig. 7. Diagram of relative evaporation in Skokie Marsh area, near Chicago,
at 10 inches (25 cm.) above the soil. Recaleulated. (Adapted from Sherff.)
78
was in an intermediate position, and that in an adjacent white oak-ash
forest evaporation was about twice as much as in the swamp meadow.
Sherff used as standard the forest (D). This gave him for the center
of the swamp (A) 38 per cent., for the swamp meadow (C) 54 per
cent., and for the outer swamp margin (B) 105 per cent. In Figure
7, I have used his swamp meadow as 100 per cent., and by recalcula-
tion this gives the forest (D) 185 per cent., for the swamp margin (B)
105 per cent., and for the center of the swamp (A) 70 per cent. These
figures indicate a concentric arrangement of the conditions of evap-
oration about the swamp.
Intensity of evaporation........6.004. 10.20 30 40 s0 60 70 80 90 1000
1907: 7
Sta. A. Above vegetation. 4 feet, 6
inches above soil
Sta. B. Middle of vegetation. 2 feet,
2 inches above soil
Sta. C. Lower vegetation. 5 inches
above soil
100%,|
1908:
Sta. A. Above vegetation. 5 feet, 6
inches above soil
Sta. B. Middle of vegetation. 2 feet,
2 inches above soil
Sta. C. Lower vegetation. 5 inches
above soil
100%
Fie. 8. Diagram showing the relative evaporation at different vertical levels in
a marsh in England, the evaporation in the lower layers of the vegetation being much
greater than in the upper strata or in the air above it. (Data from Yapp.)
Thus far, attention has been devoted solely to the horizontal differ-
ences in evaporation. There are also important vertical ones, vary-
ing above the surface of the substratum. Important observations on
this subject have been made, by a porous-cup method, in an open
grassy marsh in England, by Yapp (’09). ‘The vegetation grew to a
height of two to five feet. From his data the accompanying diagrams
(Figs. 8, 8a) have been prepared. This shows that when the stand-
ard was made the rate of evaporation above the general level of the
vegetation, within the grass layer evaporation was reduced from about
one half (Sta. B, 1908, 56.2 per cent.) to one third (Sta. B, 1907,
32.8 per cent.) at 2 feet 2 inches above the soil; and that at 5 inches
above the soil it was reduced to between one fourteenth (Sta. C, 1907,
6.6) and one seventh (Sta. C, 1908, 14.7) of that above the vegeta-
tion. Yapp (1. c.: 298) concludes from his studies that “In general,
the results of the evaporation experiments show that the lower strata
of the vegetation possess an atmosphere which is continually very much
79
more humid than that of the upper strata, and farther, that the higher
and denser the vegetation the greater these differences are.” This is
shown in Fig. 8a.
Intensity of evaporation............. jo 20,3040 50 S60 70 S80 S900 ~—s100
Sta. A. 60 inches above ground, above
vegetation 100%
Sta. B. 12 inches above ground among
vegetation mK
Sta. C. 3 inches above ground, among
vegetation ope
Fic, 8a. Diagram showing the zelative evaporation at different vertical levels in
a marsh in England, the evaporation in the lower layers of the vegetation being much
greater than in the upper strata or in the air above it. (Data from Yapp.)
In America only a few records have been made on vertical gra-
dients in evaporation, two of these in marsh areas, one in Ohio by
Dachnowski (’11), and the other near Chicago by Sherff (’13a, ’13b).
The Ohio observations, made upon a small island in a lake, in a cran-
berry-sphagnum bog, show that the rate of evaporation above the vege-
tation is much greater than among it, and that this diminishes as the
soil is approached, these results agreeing with those obtained by Yapp.
Sherff’s observations were made in Skokie Marsh, north of Chicago,
and show that the relative evaporation also varies with different kinds
of swamp vegetation. From his data a diagram has been made (Fig.
9) in which the rate of evaporation in the upper part of the reeds
Intensity of evaporation.......-..+6+ to 20 30 40 SO 60 70 680 90 100
Phragmites
Sta. A. Within vegetation, 198 em. (77 100%
inches) above soil. Standard.
Sta. B. Within vegetation, 107 cm. (42 709
inches) above soil
Sta. CO. Within vegetation, 25 cm. (10
inches) above soil
53)
Sta. D. At soil surface 33%
Typha
Sta. A. Within vegetation, 175 cm. (69
inches) above soil
Sta. B. Within vegetation, 107 cm. (42
inches) above soil
Sta. C. Within vegetation, 25 cm. (10
7 inches) above soil
Sta. D. At soil surface
Fig. 9. Diagram of relative evaporation at different vertical levels above the soil
within the vegetation of Skokie Marsh. (Adapted from Sherff.)
80
(Phragmites) at 77 inches is taken as 100 per cent. or the standard.
Lower down, at 42 inches, the rate is 70 per cent., at 10 inches, 53 per
cent., and at the surface, 33 per cent. Among the cattails (Typha), in
the upper part of the vegetation, at 69 inches evaporation was 85 per
cent.; at 42 inches it was 36 per cent.; at 10 inches, 20 per cent.; and
at the surface, 8.5 per cent. These results show that at successively
lower levels in the vegetation the rate of evaporation is greatly re-
duced. They tend also to confirm the results of Yapp and Dachnow- °
ski. It seems, then, fair to conclude that the rate of evaporation above
the swamp vegetation increases rapidly with downward progression,
and probably with upward progression also. A vegetable layer, com-
parable to the mulching of straw used by gardeners, thus acts as a pow-
erful conserver of moisture. There are great differences within a few
vertical feet in the open; what is the condition within the forest ?
Intensity of evaporation............. 20 40__ 60 8010012040 160180
Sta. A. Maple-beech forest. 6 feet (2 m.)
above soil
Sta. B. Maple-beech forest. 10 inches
(25 cm.) above soil
Sta. O. Maple-beech forest. On slope of
ravine 30 feet deep (10 m.)
13.3 feet (4 m.) below general
surface.
Fig. 10. Diagram showing the relative evaporation in a beech-maple woods, six
feet above the soil (A), near the surface of the soil (B), and in a ravine (C).
[Adapted from Fuller (’12).]
The character of vertical differences in evaporation within the for-
est has not been given as much attention as the similar changes in the
open; but attention has already been called to the moisture-conserving
effect of a forest litter, the evaporating rate in one instance being only
13 per cent. when compared with that from a water surface in the open.
McNutt and Fuller (’12) have shown that grazing in an oak-hickory
forest changed the average daily rate of evaporation for 189 days
from 9.89 c.c., in the ungrazed forest, to 12.74 c.c., in the grazed for-
est, at Palos Park, Ill. There are thus, within the forest, changes in
evaporation with differences both in the ground cover and in the litter
on the forest floor which correspond to the change in the vegetation in
open places.
Vertical differences in evaporation have been tested in a maple-
beech-forest in northern Indiana by Fuller (’12b), who used the po-
rous-cup method. His results have been summarized in Figure 10.
This diagram shows that the evaporation at six feet above the surface
is nearly twice as much as that at 10 inches above the surface, and
81
that in a ravine, 13.3 feet (4 m.) below, it was 80 per cent. of that 10
inches above the surface. The relative seasonal activity from May to
November is shown in Figure 11. This diagram shows that after the
leaves appear the highest evaporation takes place in July. This is
probably the critical season for some animals.
MAY JUNE JULY AUGUST | SEPTEMBER | OCTOBER
F1¢. 11. Diagram showing the average daily rate of evaporation in beech-maple
forest, six feet above soil (a), near the surface of soil (b), and in a ravine (ce).
(From Fuller.)
In the forest, Libernau (Harrington, ’93: 34) found that the “rela-
tive humidity increases and decreases with the absolute humidity,
whereas it is known in general, and also at the Station in the open
country, that these two climatic elements are inverse. This is ac-
counted for by the fact that the forest is a source of atmospheric
aqueous vapor as well as of cooling.” (L.c. : 104: “The absolute
humidity decreases in the forest from the soil upwards. The rate of
decrease is usually the greatest under the trees and the least at the level
of the foliage. The rate above the trees is intermediate between the
other two. ‘This rate is least in the late hours of the night, when it
may be zero. It increases with the increase of the temperature of the
air, becoming greatest in the midday hours, when, under exception-
ally favorable circumstances, it may make a difference of 10 per cent.
82
or even more. Occasionally, in high winds, the absolute humidity is
greater over the trees. Over the field station the daily progress of ab-
solute humidity was about the same as in the forest, but the maximum
difference was only about half as great. The absolute humidity in and
above the forest is greater than that over the open fields, and there is
some trace of an increase of this difference to the time of maximum.”
A greater relative humidity has been found over evergreen trees
than over deciduous trees, which is slight (1.c.: 104), but the psy-
chrometer was close to the evergreens and farther above the decidu-
ous ones.
Intensity of evaporation.............
Sta. A. 20 rods (330 ft.) from wind-
break, 25 to 40 feet high.
Standard
Sta. B. 12 rods (198 ft.) from wind-
break
Sta.C. 3 vods (49.5 ft.) from wind-
break
(July 15-Sept 15} 62 days, Lincoln, Neb
Fig. 12. Diagram showing relative retardation of evaporation by a windbreak,
Lincoln, Nebraska. [Adapted from Card (’97).]
The border of the Illinois forest and prairie was characterized by
tongues and isolated groves of forest and by glades. The forest had
the same kind of influence as windbreaks upon the leeward areas and
glades, and therefore the influence of windbreaks upon the evaporating
power of the air is of interest. Card (’97) made a valuable study of
this series of problems at Lincoln, Nebraska. The influence of wind-
breaks upon evaporation is summarized in Figure 12. This diagram
shows that leeward of a close windbreak ranging from 25 to 40 feet
in height, the rate of evaporation in terms of the standard (A), which
was 330 feet leeward, was 91 per cent. at a distance of 198 feet (B),
and 71 per cent. at 49.5 feet (C), thus showing a marked reduction
with proximity to the windbreak. These observations covered 62 days.
Nearer to Illinois, similar though very limited observations were
made in central Wisconsin by King (’95) which agree with Card’s
on the retardation of evaporation by windbreaks. His results are
shown graphically in Figure 13.
Recently Bates (’11) has made an elaborate study of the effects of
windbreaks upon light, soil, moisture, velocity of wind, evaporation,
humidity, and temperature. His results confirm those just given and
give additional facts which, however, with one exception, will not be
mentioned. The paper itself should be consulted. This investigation
by Bates shows that in proportion to the perfection of the windbreak
83
a quiet, stagnant air strip is formed to the leeward, and that this fa-
vors excessive heating during clear days and low temperatures on clear
nights. Years ago Harrington (’93: 119) suggested this idea and
called attention to the close relation existing between the leeward con-
ditions of windbreaks and forest glades. The glade climate is more
rigorous, or extreme, than that upon plains (I.c.: 19, 84-88, 119).
Such a climate is thus a bit more “continental” during the spring, sum-
Intensity of evaporation............. i029 305060 080900080
Distance from windbreak 12 inches high:
Sta. F. 500 feet leeward. Standard
Sta. E. 400 feet leeward
Sta. D. 300 feet leeward
Sta. C. 200 feet leeward
Sta. B. 100 feet leeward
Sta. A. 20 feet leeward
Fie. 13. Diagram showing the relative evaporation, May 31, at different dis-
tances leeward of a windbreak, Almond, Wis. [Adapted from King (’95).]
mer, and autumn. These glades are very hot in the early afternoon
- and cool on clear nights, and the air is relatively stagnant; as Harring-
ton says, it is “lee for winds from all directions.” The center of a
dense forest may thus possess physical conditions quite different from
those of the glade forest margin or in the open. Beginning with the
relatively stable conditions within a forest toward its margin, the diur-
nal temperature variations are much more extreme (Harrington,
1. c.: 89) “to a distance of a score or so of rods where it reaches a max-
imum. The amplitude is greater in glades. Hence the extremes of
temperature are exaggerated just outside the forest.” The annual soil
temperatures of a glade are intermediate between that of the forest and
the plain. The forest margin is thus seen to possess many of the char-
acteristics of the glade, for its climate is somewhat more extreme than
that in the open, far from the forest.
5. Temperature Relations in the Open and in Forests
The temperature relations in open and forested regions are often
very different. The density of the vegetable covering in the open and
in the forests varies much and may have considerable influence upon
animals. Yapp (’09) observed that the marsh vegetation in England
84
caused marked vertical differences in temperature in the vegetational
stratum. He summarizes these results as follows (p. 309): “The
temperature results show that the highest layers of the vegetation pos-
sess a greater diurnal range of temperature than either the free air
above or the lower layers of the vegetation. Regularly, especially in
clear weather, both the higher day and the lowest night temperatures
were recorded in this position.”
Dachnowski (’12: 292-297) studied the temperature conditions in
a cranberry bog substratum in central Ohio. He found that at a time
when ice formed from 8 to 15 inches thick on the adjacent lake, in the
bog it was only 3 to 5 inches thick, and there were small patches where
it did not form at all. Ata depth of 3 inches in the peat the tempera-
ture ranged from 33° to 77° F. (.5 -25.0° C.). In the bordering
maple-alder zone, at 3 inches depth it ranged from 33° to 72° F. (.5°-
22.0 C.). His observations indicate that the temperature relations
within the maple-alder zone are more stable than those in the open
central area.
Cox (’10) has also shown that the character of the vegetation in
Wisconsin cranberry bogs has much influence upon temperature rela-
tions in this habitat.
It seems very probable that similar conditions hold over prairie
vegetation, but I do not know of any observations on this point. We
are all familiar with the common practice of gardeners of using a mulch
of straw to retard temperature changes under it; prairie vegetation
must have a similar influence. (Cf. Bouyoucos, ’13: 160.)
The relative air temperatures within and without the forest show
a distinct tendency to reduce the maxima and minima, and to lower
the mean annual temperature. Harrington (’93:53) concludes,
therefore, that “the forest moderates (by reducing the extremes) and
cools (by reducing the maxima more than the minima) the tempera-
ture of the air within it. The moderating influence is decidedly greater
than the cooling effect.” ‘These effects are not uniform, but are much
more marked in the summer, and Harrington further says: ‘The cool-
ing effect tends to disappear in winter. The moderating effect is the
most important one and it is the most characteristic” (p. 56).
The temperature relations within the forest crown show that in
general the effects are similar to those found at an elevation of about
5 feet. The maxima are lowered, the minima are elevated, and there
is a cooling effect. The differences are most pronounced during the
summer, and the temperatures are intermediate in position between
‘those at the five-foot level and those in the open (l.c.:66). Ata
height of 24 feet, deciduous trees showed a marked summer cooling
85
effect, while evergreens showed much less, though they are much more
uniform for 9 months of the year. Again, he says: “In summer the
average gradient under trees is about +2°; that is, it grows warmer
as we ascend at the rate of two degrees per 100 feet (31 m.). Out-
side in the general average it grows colder by about a quarter of a de-
gree.” This warmer air above the cooler in the forest favors its sta-
bility or relative stagnation, although as a whole the forest air is cool-
er and heavier than the surrounding air and tends to flow outward.
‘The forest thus tends to produce a miniature or incipient barometric
high. In conclusion Harrington (p. 72) states that “The surface of ,
the surface of the forest is, meteorologically, much like the surface of
the meadow or cornfield. The isothermal surface above it in sun-
shine is a surface of maximum temperature, as is the surface of a
meadow or cornfield. From this surface the temperature decreases in
both directions.” In the case of a beech forest the warm diurnal layer
above the forest crown was only 6.5 feet thick (p. 34).
The conditions above the forest are thus representative of the at-
mospheric conditions above dense vegetation in general, and are in per-
fect harmony with Yapp’s observations upon the temperature above a
marsh (’09: 309), quoted on a previous page, to the effect that tem-
perature changes are extreme here, and greater than in the free
air above or in the lower layers among the vegetation. The forest is
thus to be considered as a thick layer of vegetation in its influence upon
meteorological conditions. 'The conditions above the forest, there-
fore, exemplify a general law.
In general terms, the temperature of the soil below the zone of
seasonal influence is that of the mean annual temperature for a given
locality. The surface zone, however, varies with the season. Har-
rington (’93) has summarized the German observations on the rela-
tive soil temperatures in the open and in the forest. In the following
quotation the minus sign indicates a forest temperature less than a cor-
responding observation in the open. These temperatures were taken
about 5 feet above the soil. He says (p. 43): “The average of the
seventeen stations (representing about two hundred years of observa-
tions) should give us good and significant results. It shows for the
surface—2°.s9, for a depth of 6 inches (152 mm.)—1°.87, and for
a depth of 4 feet (1.22 m.)—2°.02. The influence of the forest
on the soil, then, is a cooling one, on the average, and for central
Europe the cooling amounts to about two and a half degrees for the
surface. The cooling is due to several causes: The first is the shade;
the foliage, trunks, branches, and twigs cut off much of the sun’s
heat, absorb and utilize it in vegetative processes, or in evaporation, or
reflect it away into space. Thus the surface soil in the forest receives
86
less heat than the surface of the fields. The same screen acts, how-
ever, in the reverse direction by preventing radiation to the sky, thus
retaining more of the heat than do the open fields. The balance of
these two processes, it seems from observation, is in favor of the first
and the average result is a cooling one... .. . The differences of
temperature at the depth of 6 inches (152 mm.) are more than half a
degree less than at the surface. In this is to be seen the specific effect
of the forest litter; it adds a covering to that possessed by the sur-
face, so that while the deeper layer is cooled as much by the protec-
tion from the sun’s rays as is the surface, it is not cooled so much by
radiation of heat to the sky. Its temperature is, consequently, rela-
tively higher, and approximates somewhat more the field tempera-
tures.”
“The forest soil is warmer than that of the open fields in winter,
but cooler in the other seasons, and the total cooling is much greater
than the warming one... ... The forest, therefore, not only cools
the soil, but also moderates the extremes of temperature” (p. 46).
The character of the forest, whether evergreen or deciduous, in-
fluences the temperature conditions of the soil, as is seen by a com-
parison of these conditions in the forest and in the open. The two kinds
of forest are much alike in winter; during the spring the soil warms
up more rapidly under conifers. Temperature variations are slightly
greater under deciduous trees.
6. Soil Moisture and its Relation to Vegetation
The moisture in the soil is derived largely from precipitation, but
part of it, in some localities, comes directly from the adjacent deeper.
soils or rocks, and thus only indirectly from precipitation. As Illinois
lies at the bottom of a large basin, there must be some subsurface flow
from the adjacent higher regions, but to what extent is not known.
McGee (’13a:177) estimates that the general ground-water level—
the level at which the soil becomes saturated—has, since settlement, de-
clined 10.6 feet in Illinois. This decline is not limited to drained re-
gions but is a general condition. In addition to these changes of level
there are seasonal fluctuations. Sherff (’13a: 583) observed in Skokie
Marsh that the water-table was at or above the surface in May, then
declined until early September, and then rose rapidly to the surface by
the middle of October. The wet prairie at Charleston has undergone
just such changes as these; the ground-water level has been lowered
and there are marked seasonal changes.
Harvey (’14) has recently shown that the soil of Eryngium-Sil-
phium prairie at Chicago contains a large amount of water during
87
April and until late in May; that the moisture falls and is low during
July and August, with a mean of 24 per cent. of saturation for these
months; but that in October the soil is again at or near the point of
saturation.
The blanket of humid air which accumulates under a cover of vege-
tation, retards evaporation and conserves soil moisture. The denser
the vegetation the more marked is its influence. The litter—the or-
ganic debris in an early stage of decomposition—on the forest floor
has the same tendency, and has even a greater water capacity than the
soil itself. On the other hand, a forest is a powerful desiccator; as
Zon (’13:71) has recently put it: “A soil with a living vegetative
cover loses moisture, both through direct evaporation and absorption
by its vegetation, much faster than bare, moist soil and still more than
a free water surface. The more developed the vegetative cover the
faster is the moisture extracted from the soil and given off into the air.
The forest in this respect is the greatest desiccator of water in the
ground.” This drying effect is shown particularly near the surface
of the soil, where roots are abundant and where drouth is so marked
that it may prevent the growth of young plants here (cf. Zon and
Graves, “11: 17-18).
Warming (’09:45) says: “It may be noted that, according to
Ototozky, the level of ground-water invariably sinks in the vicinity of
forest, and always lies higher in an adjoining steppe than in a forest;
forest consumes water.”
McNutt and Fuller (12) have made a study of the amount of soil
moisture at, 3 inches (7.5 cm.) and at 10 inches (25 cm.) below the
surface in an oak-hickory forest, at Palos Park, Illinois. They found
that the percentage of water to the dry weight of the soil at the 3-inch
level averaged 18.9 per cent. and at 10 inches was 12.5 per cent. of the
dry weight of the soil. The greater moisture near the surface is due
to the humus present in this layer. The grazed part of the forest
possessed less soil moisture, and shows the conserving effect of vege-
tation. (Cf. also Fuller ’14.)
The artificial control of soil moisture is well shown by the effect of
windbreaks. Card (’97) studied the moisture content of the soil to
leeward of a windbreak and found that in general there is a “de-
crease in the per cent. of water as the distance from the windbreak
increases.” As the physical conditions leeward of windbreaks are
similar in many respects to those in forest glades and forest margins,
it is very probable that the conditions of soil moisture also will be very
similar in these places.
88
7. Ventilation of Land Habitats
The preceding account of the temperature, humidity, and evapo-
rating conditions in various habitats forms a necessary basis for an un-
derstanding of the processes of ventilation or atmospheric change in
land habitats. The differences in pressure due to the different densi-
ties of cool and warm air and to the friction and retardation of mov-
ing air currents, determine to an important degree the composition
of the air in many habitats. In such an unstable medium as air,
changes take place very rapidly through diffusion, and through this
constant process of adjustment there is a tendency to level off all local
differences. These are naturally best preserved where diffusion cur-
rents are least developed—in the most stagnant or stable atmospheric
conditions; therefore any factor which retards an air current and pro-
duces eddies, or slow diffusion, will favor local differentiation of
the air.
We have seen that any vegetable cover retards air currents, so that
the air within the vegetation becomes different from the faster moving
air above it. The accumulation of humidity at different levels above
the soil within the vegetation, clearly shows this. The denser the vege-
tation-the more completely are the lower strata shut off and, to a cor-
responding degree, stagnant and subject'to the local conditions. Two
factors have an important influence upon these conditions: the charac-
ter of the cover itself, and the character of the substratum. If both
of these are mineral rather than organic, in general comparatively
little local influence is to be expected, although in some localities CO,
escapes from the earth and on account of its density may linger in de-
pressions and thus kill animals (Mearns ’03). Generally, however,
the organic materials are of most importance both as a cover and asa
substratum, and are often the source of carbon dioxide. Living vege-
tation may also add oxygen to such stagnant air, but the main source of
it is the free air itself. The forest litter, on account of its imperfect
stage of decay, consumes oxygen and gives off carbon dioxide; in the
humus below it, shut off even more from free access to air, the carbon
dioxide is relatively more abundant and the oxygen relatively less so
or absent; and in the deeper mineral soil the amount of carbon
dioxide is relatively less on account of the absence of organic debris,
and a small amount of oxygen is present.
The aeration of the soil is influenced to a large degree by its poros-
ity; the looser it is, the freer the circulation. Buckingham (’04) has
shown that “the speed of diffusion of air and carbonic acid through
these soils was not greatly dependent upon texture and structure, but
was determined in the main by the porosity of the soil. . . . the
89
rate of diffusion was approximately proportional to the square of the
porosity . . . . the escape of carbonic acid from the soil and
its replacement by oxygen take place by diffusion, and are determined
by the conditions which affect diffusion, and are sensibly independent
of the variations of the outside barometric pressure.”
In the upper, better ventilated, moist, neutral or alkaline layers of
vegetable debris decomposition is brought about mainly by the agency
of fungi; but in the deeper, poorly ventilated acid layers, lacking oxy-
gen, bacteria are the active agents (cf. Transeau, ’05, 06). The
higher the temperature the more rapid the circulation, and on this ac-
count ventilation in the open is relatively more rapid than in the cooler
woodlands. The black soil prairies are thus favorable to a higher tem-
perature and better ventilation. Dry soil, according to Hilgard
(706: 279) contains from 35 to 50 per cent. its volume of air, and in
moist or wet soils this space is replaced by water. Thus the condi-
tions which influence the amount of water present have a very im-
portant influence upon aeration. As water is drained from the soil, air
takes its place; so drainage and the flow of water through the soil facil-
itate ventilation. The part of the soil containing air is thus above the
water-table; and as this level fluctuates with the season and from year
to year the lower boundary of this stratum is migratory. Hilgard
states that cultivated garden soil contains much more air than uncul-
tivated forest soil, Warming (’09: 43) says that the “production of
acid humus in the forest leads to an exclusion of the air.” If lime is
present, such an acid condition can not arise.
While the source of oxygen in the soil is the air, the reverse is the
case with carbon dioxide. The surface layers of the soil, among
dense vegetation, constitute an area of concentration of carbon
dioxide. Because this is more soluble than other gases, it is found
in rain water, according to Geikie, in a proportion 30 to 40 times
greater than in the air. Rains thus assist in the concentration of
carbon dioxide in the soil. This concentration is well shown by the
following table by Baussungault and Lewy (Van Hise, ’04: 474).
co, in
Character of soil air 10,000 parts
by weight
1. Sandy subsoil of forest 38
2. Loamy subsoil of forest 124
3. Surface soil of forest 130
4. Surface soil of vineyard 146
5. Pasture soil 270
6. Rich in humus 543
90
The amount of carbonic acid in the atmosphere is by weight about
4.5 parts in 10,000, The amount in the air is, as Van Hise says, “‘in-
significant in comparison with the amount in soils in regions of luxu-
riant vegetation. In such regions the carbon dioxide is from thirty to
more than one hundred times more abundant than in the atmosphere.”
This carbonic acid in the presence of bases, sodium, potassium, cal-
cium, and magnesium compounds, forms carbonates and bicarbonates.
This is the process of carbonation—one of the most important proc-
esses of change in surface soils.
In view of the dominance of CO, in soils we may anticipate that
many of the animals living in them possess some of the characteristics
of the plants, bacteria, fungi, etc., which are active in such soils. The
anaerobic forms live without free oxygen; others live only where oxy-
gen is present. The animals which thrive in the soil are likely to be
those which tolerate a large amount of CO, and are able to use a rela-
tively small amount of oxygen, at least for considerable intervals, as
when the soil is wet during prolonged rains. This is a subject to
which reference will be made later. :
The air is the main source of oxygen, and from the air it diffuses
into the soil; thus the process of equilibration is constantly in progress.
Carbonic acid, also present in the air, is washed down by rain and
concentrated in the soil, where it is increased by the decay of organic
debris and by respiring animals to such an extent that it exists under
pressure and diffuses into the air, thus contributing to the air. In the
soil, then, the process of decarbomzation is of great importance to
animal life, and must not be neglected. The optimum soil habitat is
therefore determined, to a very important degree, by the proper ratio
or balance between the amount of available oxygen and the amount of
carbon dioxide which can be endured without injury. The excessive
accumulation of carbon dioxide, an animal waste product, is compar-
able to the accumulation of plant toxins which may increase in the
soil to such a degree as to inhibit plant growth. Such substances
must be removed from the soil, or changed in it to harmless com-
pounds, or plants and animals can not continue to live in certain
places. I have used the term ventilation to cover both the oxygena-
tion and decarbonization of land habitats, and the same principles
are applicable to life in fresh-water habitats.
We have just seen how atmospheric ventilation favors the removal
of certain injurious waste products from the air and soil. In addition
to gaseous waste products there are also liquids and solid kinds which
may be equally harmful in a habitat. These are known to exist in con-
fined liquids, as in aquaria (Colton, ’08; Woodruff, ’12), where they
91
interfere with the welfare of the animals present, and it is probable
that they also exist in soils. The older naturalists elaborated the idea
that if organisms were not such active agents in the destruction or
transformation of plant and animal bodies such remains would soon
encumber the earth. Thus organisms themselves are among the most
active agents in influencing directly and indirectly the ventilation of
animal habitats.
8. The Tree Trunk as a Habitat
A living tree trunk is composed of wood, sap (moisture), and
bark, all of which are relatively poor conductors of heat. When the
trunks are cooled, as in winter, they are slow in warming, not only
because of poor conduction but also because of the slow circulation of
sap, which is derived from the cool ground-water. As the season
progresses, the trunks warm up, this process being retarded in part by
the shade and the cool forest conditions; and in the fall, radiation of
the heat accumulated also takes place slowly. The tree trunk therefore
changes its temperature slowly, as does the soil. The animals which
live within wood thus live in a relatively cool and stable environment.
In living trees the humidity is relatively high, as it may also be in
fallen, decaying logs. Relatively dry logs, before progress of decay,
on the other hand, form a relatively dry and uniform habitat. (Cf.
on the temperature of trees: Harrington, ’93, pp. 72-75; Packard,
’90, p. 23; and Jones, Edson, and Morse, ’03, pp. 97-100.)
9. Prairie and Forest Vegetation and Animal Life
The dependence of animals upon plants for food is one of the most
fundamental animal relations. It is a world-wide relation, but its
mode of operations varies greatly in different environments. For ex-
ample, many years ago, Brooks gave us a graphic picture of the réle
of marine vegetation in the economy of marine animals. In the sea
there are no forests or grasslands, and no corresponding animals as-
sociated with these conditions, as on land; but in the sea great numbers
of minute plants float, and upon these feed an immense number of
small crustaceans and other small animals. These small creatures
occur in such large numbers that at times the sea is a sort of gruel
which sedentary and stationary kinds may appropriate by simply al-
lowing the sea to flow into their mouths. The food here circulates in
their environmental medium, as plant foods do in the soil and air. This
condition has made it possible for vast numbers of plant-like animals
to grow over the sea floor as plants do over rocks and plains. The
living meadows of animals thus furnish pasture for a host of preda-
92 \
ceous kinds; and upon these still others prey, so that flesh-eating ani-
mals make up the most conspicuous classes of marine animals. Quite
otherwise are the conditions on land, where no air current carries food
to the hungry mouths of animals. Plants with roots in the soil and
stems in the air are able, however, to secure their food from the cir-
culating medium, but being themselves fixed, they are easy prey to
animals—both the sedentary kinds, which live in or upon the plant tis-
sues, and the active wandering kinds, which forage over large areas.
The predaceous animals, either by active mind or body, must secure
their food from the plant-feeding kinds. The great expanses of grass-
land and forest tend to be devastated by a vast army of animals which
far outnumber the predaceous kinds. The conditions of life, there-
fore, found upon grassland areas, like the prairie, and in the forest,
are to the farthest possible extent removed from those found in the
sea. ‘This, then, is one of the most fundamental contrasts in the con-
ditions of existence encountered by animals.
These considerations naturally raise the question to what extent
and in what particular manner does land vegetation influence animal
life? Does a change in the vegetation as great as that between the for-
est and the prairie have a marked influence upon animals? In the
Charleston region we have just such a difference in the vegetation.
Many years ago Bates pointed out repeatedly in his “The Natural-
ist on the River Amazons” that the animals of that densely forested
region were to a marked degree distinctly arboreal and “adapted” to a
forest life. In most densely forested regions like conditions probably
prevail, and to a corresponding degree open lands harbor animals
equally characteristic and as truly terrestrial in habits. The contrast
between the conditions of life in the open and in the forest is one of
the most fundamental environmental conditions upon land. The sig-
nificance of this contrast seems to have been realized only in part. The
prairies or grasslands are representative of only one kind of open;
they are caused by many kinds of factors limiting the extension of
forests. Open places are formed by lakes, ponds, and swamps; by the
avenues through forests formed by different kinds of streams, as
brooks, creeks, and rivers; by the small amount of soil on rock sur-
faces; and by still other kinds of limiting influences, such as the sea,
severe climate, and altitude. Among almost all of the major taxo-
nomic groups of land animals is seen the independent origin and pres-
ervation of animals suited for life in the forest; this clearly points to
the extensive influence and antiquity of this environment. The same
is true of animals living in the open. But to assume that it is solely
the kinds of forest trees serving as food for animals, or the cor-
responding kinds of vegetation in the open, which determines whether
93
an animal lives in the open or in the forest, would be unwarranted in
the light of the preceding discussion of the effect of vegetation upon
air temperatures, winds, humidity, relative evaporating power of the
air, and corresponding changes in the soil. Animal life is’ most
abundant in a narrow vertical layer above the earth’s surface, by far
the most of it is within a few inches or feet of the surface; and above
the level of the forest-crown it diminishes with great rapidity. Be-
low the surface of the soil the same general law holds; most of the
ground animals are within the first few inches of soil, only a small
number extending a few feet below the surface, and those found at
greater depths being indeed very few. The rate of decline is many
times more rapid below the surface than it is above it. There is, then,
above and below the surface a rapid and progressive attentuation of
the favorable conditions for animals and plants, and the animals do
not establish thriving communities far from those physical conditions
which are also favorable to vegetation. Animals are dependent upon
plants for food, but both are dependent upon a certain complex of
physical conditions near the surface of the earth.
It is well to recall at this point how the influence of the climate and
the vegetation exemplify certain general laws which operate in all hab-
itats. The differentiation of habitats upon the earth is primarily due
to temperature and the specific heat relations of the earth, which re-
sult in the several media—gases, liquids, and solids. With a higher
temperature all would be gas, and with a lower one all would be solidi-
fied. The present intermediate conditions, therefore, permit the pres-
ent differentiation. These media are further differentiated by tem-
perature about as follows: Since the source of solar energy, heat, and
light, and the oxygen supply, are above the surface of the earth, the
vertical attenuation of these influences is one of the most striking
peculiarities of animal habitats, both in water (where the causes have
long been recognized) and upon land. Any covering of the earth,
even the surface layer of vegetation, soil and water, tends to shut off
heat, light, and oxygen. At the same time such a layer tends to shut
in those influences which originate primarily in or below it. Thus car-
bonic acid originating under the cover, by organic decay, breathing
animals, or bacteria, or washed in by the rain, tends to be shut in.
Furthermore, heat once reaching here, either in water or on land, tends
toward slow radiation. ‘Thus we may look upon the surface layer as a
partition which is under pressure from both sides, and through which
constant interchange is in progress, as the process of dynamic equili-
bration operates. ‘:
94
This attenuation of intensities, above and below the surface, pro-
duces vertical layers of relatively equal strength or pressure. Thus
the attenuation of temperature in gases (air) and in liquids (water)
causes different densities in air and in water which modify to an im-
portant degree the physical and chemical conditions in these media.
This results in their stratification: when the heavier layers are below,
stability is the tendency; and when the reverse order obtains, a
change takes place toward the stable condition. With stratification,
flowage tends to occur within the strata, and to be horizontal rather
than vertical; additional pressure is therefore necessary to cause the
vertical currents or circulation under such conditions. This is why
carbonic acid accumulates in the soil and in small deep lakes abound-
ing in organic debris, this accumulation being largely due, in both
cases, to the slow rate of exchange caused by the stratification pro-
duced by differences in density. This same relative stagnation is a
primary factor in the vertical differences in the relative evaporating
power of the air within a vegetable layer of the prairie or the forest.
Though on the prairie the vegetational layer is generally but a few
inches or a few feet thick, in the forest it is about eighty feet, or
more, thick; and the forest thus influences atmospheric conditions
solely as a thick layer of vegetation.
Differences, then, in the character, structure, or composition of the
surface of the substratum are of fundamental importance in under-
standing its relative influence upon animals. Primarily these differ-
ences are due to temperature, secondarily to temperature in combina-
tion with moisture; and they result in the relative humidity and the
relative evaporating power of the air. The most important difference
in the surface layer in the Charleston region is that of prairie and for-
est, and therefore the main features of these habitats will now be sum-
marized. It should not be overlooked that conditions on the prairie are
likely to be quite representative of open places in general, though they
will probably be somewhat unrepresentative in the case of open places
having wet or extremely dry substrata. It is also true that the condi-
tions produced by the forest are comparable, in some degree, with
those due to the influence of an elevation.
95
SUMMARY oF ENVIRONMENTAL FEATURES OF THE PRAIRIE AND THE DECIDUOUS FoREST
—TEMPERATURE, HUMIDITY, AND EVAPORATION—DURING THE GROWING SEASON
Above the Vegetation
Prairie Forest
In sun, maximum heated stratum. Above crown, in sun, maximum heated
Cooler above and below this stratum. stratum, A thin layer. Cooler above
Absolute humidity less than in or over and below this stratum.
forest. Absolute humidity greater than.in the
open.
Among the Vegetation
Prairie Forest
Temperature lower and higher than in Temperature moderated—not as low or
the forest—more extreme. as high as on the prairie.
Temperature lower toward the soil, and Temperature lower toward the soil, and
warmer than in the forest. cooler than in the open.
Absolute humidity progressively increases Absolute humidity progressively increases
toward the soil. toward the soil.
Relative evaporation decreases toward the Relative evaporation decreases toward the
soil; greater than in the forest. soil; less than in the open.
In the Soil
Prairie Forest
Temperature averaging warmer than Temperature cooler on the average and
forest, warmer near surface in sum- in summer, and warmer in winter, near
mer, and cooler in winter. Warmer in the surface, than in the open. Cooler
sun and cooler at night than in forest. in sun and warmer at night than in
tha open.
Temperature progressively more stable Temperature progressively more stable
downward. Soil moisture increases downward. Soil moisture, below the
downward. surface layer, increases downward.
The conditions on the prairie and in the forest may be graphically
shown as in the following diagrams, Figure 14 showing the tempera-
ture relations, and Figure 15 showing the relative evaporating power
of the. air.
96
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98
10. Sources and Réle of Water used by
Prairie and Forest Animals
The bodies of animals contain a very large proportion of water—
from 60 to 95 per cent. Growing animals in particular require water
in relatively large amounts. Practically all foods gain entrance into the
body in aqueous solutions, and are transported by water to all parts;
and by the same means, the waste products, with the exception of the
excretion of carbonic acid, are removed. The methods by which aquatic
animals secure water are relatively simple, because they live in a
liquid medium; but the conditions upon land are quite different. Here
osmotic pressure does not operate as in water, and the air varies from
saturation to a very dry condition. This dryness tends to cause strong
evaporation from animals living in such a medium, and a proper bal-
ance between intake and water-loss is one of the most potent influences
in the life of land animals. In this relation lies the importance of the
sources of water available to them. These sources are as follows:
with the food, by drinking, from the atmosphere, and by metabolism.
The loss is by excretion and evaporation, the relative humidity and the
evaporating power of the air being, therefore, important considera-
tions. The loss of water is retarded in many ways. Some animals
possess a relatively impermeable skin, or a covering, as hair or feath-
ers, which retards air currents and evaporation through the skin, just
as a cover of vegetation retards soil evaporation. Other animals con-
serve their moisture by modes of behavior, being active mainly during
the cooler night, thus escaping the excessive evaporation of the heated
day; and still others live in burrows in the soil, where the humidity is
higher than in the air. Many animals can live only where the air is
humid. There is thus an almost endless series of conditions relating
animals to the supply and loss of water.
On account of the herbivorous food habits of so many animals a
large number secure much water with the juicy vegetation eaten, and
others from nectar or from the sap drawn or escaping from plants.
The predaceous animals secure a large amount of water from the fluids
of the animals they devour or the juices sucked from their bodies, as
in the case of certain Hemiptera and some parasites. In addition to
the fluids derived from plants and animals, many animals also drink’
water, some in small amounts and others in large quantities. Innu-
merable observations have been made by naturalists on the drinking
habits of animals, but I know of no general discussion of this subject,
and particularly of none from the standpoint of the variation of their
behavior in this respect in different environments. But the sources
of water mentioned are not the only ones available to animals, although
99
they are the most obvious, and familiar to us. An important addi-
tional source is that formed within the body of the animal by the proc-
esses of respiration and dehydration; this is metabolic water. The
relation of this source to others and to water-loss has recently been
summarized in an important paper by Babcock (’12:87, 88, 89-90,
QI, 160, 161, 171-172, 174-175, 175-176, 181). The following quo-
tations from this paper will serve to give a concise statement of the
general principles involved in this important process. He says (pp.
87-88): “There are, however, particular stages in the life history of
both plants and animals in which metabolic water is sufficient for all
purposes for considerable periods of time... .. . This is also true
in the case of hibernating animals that receive no water from external
sources for several months, although water is constantly lost through
respiration and the various excretions. In addition many varieties of
insects such as the clothes moths, the grain weevils, the dry wood bor-
ers, etc., are capable of subsisting, during all stages of development,
upon air-dried food materials containing less than ten per cent water ;
in these cases, nearly all of the water required is metabolic. . . . Many
organisms also, when deprived of free oxygen, are capable of main-
taining for a short time, certain of the respiratory functions, and de-
riving energy from food material and from tissues by breaking up the
molecular structure into new forms of a lower order. This is known
as intramolecular respiration, and like direct respiration, results in the
production of both water and carbon dioxide.” (Pp. 89-90): “The
substances oxidized by both plants and animals, to supply vital energy,
consist of carbohydrates, fats, and proteins. All of these substances
contain hydrogen, and their complete oxidation produces a quantity
of water equal to nine times the weight of hydrogen present in the orig-
inal substances. . . . Most of the the fats yield more than their weight
of water, while proteins, when completely oxidized, give from 60 to
65 per cent of water. . . . . Animals, however, are unable to utilize
the final products of protein metabolism which are in most cases
poisonous and must be removed from the tissues by excretion in vari-
ous forms, the principal of which are urea, uric acid, and am-
monia. . . . . The amount of metabolic water formed by oxidation
during any period is proportional to the rate of respiration... .....
(Page 91): “With parasitic plants, and with animals, which derive all
of their organic nutrients from chlorophyl producing plants, im-
bibed water is not so essential to life; with these the chief function
of imbibed water is to aid in the removal of waste products, the
metabolic water being in most cases sufficient for transferring nutri-
ents and for replacing the ordinary losses incurred by respiration
and evaporation.” . . . . . (Page 160): “Another and more im-
100
portant difference is the inability of animals to resynthesize the or-
ganic waste products of respiration into substances that may be
again utilized as nutrients. . . . . This is especially the case with the
soluble products arising from protein metabolism. With most animals
these nitrogenous products are excreted in solution through the kid-
neys, chiefly as urea, but birds, reptiles, and all insects excrete most
of the nitrogenous waste matter as uric acid, or its ammonia salt, which
being practically insoluble in the body fluids, is voided in a solid con-
dition.” (Page 61): “The need for water is much less for ani-
mals that excrete uric acid than for those that excrete urea, since
uric acid, being practically insoluble in the the body fluids, is not so
poisonous as urea and is voided solid with a minmum loss of water.
Many animals that excrete uric acid instead of urea never have access
to water and subsist in every stage of their development upon air dried
food which usually contains less than 10 per cent water. The most
striking illustrations of this kind are found among insects such. as the
clothes moths, the grain weevils, the dry wood borers, the bee moths,
etc. The larve of these insects contain a high per cent of water, and
the mature forms, in spite of the development of wings which are rela-
tively dry, rarely contain less than 50 per cent of water.” (Pp. 171-
172): “Serpents and other reptiles that live in arid regions and rarely
if ever have access to water, except that contained in their food, are
said by Vauquelin to excrete all of the waste nitrogen as salts of uric
acid. The same is true of birds that live on desert islands where only
salt water is available. It is essential that animals of these types should
produce as much metabolic water as possible from the assimilated food,
and the waste of water through the excretions should be reduced to a
minimum. Since the food is largely protein both of these ends are at-
tained by the excretion of uric acid which, as already stated, contains
the least hydrogen of any nitrogenous substance excreted by animals so
that the maximum amount of metabolic water has been derived from
the food consumed.” (Pp. 174-175): “There are many animals that
are able to go long periods without having access to water except that
contained in their food, iri which water usually amounts to less than
20 per cent of total weight, and the metabolic water derived from oxi-
dation of organic nutrients. A notable example of this is the prairie
dog which thrives in semi-arid regions. ‘These small animals feed
upon the native herbage which for months at a time is as dry as hay.
It has been surmised that the burrows in which they live extend to
underground water courses, but this does not seem likely since in many
of these regions wells must be sunk hundreds of feet before water is
reached. It is more probable that they depend chiefly upon metabolic
water. They feed mostly at night when the temperature is low and
a
101
during the hottest hours of day remain in their burrows where the air
is more nearly saturated with moisture and evaporation is relatively
small.” (Pp. 175-176): “An application of these principles would
undoubtedly serve to prolong life, when suitable water for drinking
is not available. In such cases the food should consist of carbohy-:
drates and fats. Proteins should not be used... . . The water re-
quired for preventing uremic poisoning under these conditions is small
and if the relative humidity of the surrounding air is high enough to
prevent rapid evaporation of water from the body, the metabolic water
arising from the oxidation of nutrients may be ample for the purpose.”
(Page 181): “Metabolic water derived from the oxidation of organic
nutrients would probably be sufficient for all animal needs were it not
for the elimination of poisonous substances resulting from protein de-
generation.” |
The preceding quotation brings out very clearly the harmful effects
of an accumulation of uric acid upon the animal. This is only a special
case illustrating a general law, for except water the main end products
of metabolism are acid. There is thus a constant tendency for acid to
accumulate, as Henderson (’13a: 158-159; see also ’13b) has said:
“This tendency toward acidity of reaction and the accumulation of acid
in the body is one of the inevitable characteristics of metabolism; the
constant resistance of the organism one of the fundamental regulatory
processes. Now it comes about through the carbonate equilbrium that
the stronger acids, as soon as they are formed, and wherever they are
formed, normally find an ample supply of bicarbonates at their dis-
posal, and accordingly react as follows . . . . The free carbonic acid
then passes out through the lungs, and the salt is excreted in the urine.”
Recently Shelford (’13b, see also ’14a) has summarized the phys-
iological effects of water-loss by evaporation and other methods. It
is probable that the carbonic acid excretion is retarded by drying, and
that by this means irritability may be increased.
It is not simply loss of water, but loss beyond certain limits that
interferes with the life of animals. Thus loss is not an unmixed evil,
because, in addition to removing excretions, evaporation is an impor-
tant factor in the control of temperature within the bodies of animals.
Loss of water also tends to concentrate the body fluids, and when this
loss brings about a relatively dry condition, such tissues are in a con-
dition which is favorable for the endurance of relatively extreme low
or high temperature (Davenport, 97: 256-2 58), and even dryness
(see references, Adams, ’13: 98-99). This is a reason why it is dif-
ficult to distinguish, in nature, between the effects of aridity and tem-
perature extremes, and hence arise the puzzling interpretations of con-
102
tinental climates. These extreme conditions are characteristic of many
habitats.
It is readily seen how the general principles just summarized apply
to the land animals of the prairie. Many of these are active during the
day, live in the bare exposed places, or near the level of the vegetation,
where evaporation is greatest and water-loss is correspondingly large,
and feed upon the dry haylike vegetation. Others remain among the
humid layers of the vegetation or in the moist soil, and feed upon
juicy plants and other moist food. Predaceous and parasitic animals,
deriving their moisture from their prey, occupy both the dry and humid
situations. These are representative cases, between which there are a
large number of intergradations.
In the forest, where evaporation is more retarded than in the open,
a large number of animals live in the forest crown, at the forest mar-
gin, in glades, and in wood, of all degrees of dryness, and eat food
varying similarly from juicy leaves to dry wood. On the other hand,
some live in moist logs, among damp humus, or in the soil, and feed
upon dripping fungi or soggy wood. Many of these animals possess
little resistance to drying.
The optimum for prairie and forest animals thus involves a
dynamic balance between the intake of water and its loss by evapora-
tion and excretion.
ANIMAL ASSOCIATIONS OF THE PRAIRIE,
AND THE FOREST ©
I. INTRODUCTION
In an earlier chapter of this paper the habitats and animals found
at the different stations were discussed, and in the preceding section
the general characteristics of the physical and vegetational environ-
ment of the prairie and forest have been described and summarized.
We are now in a better position to consider the relations of the inverte-
brates, not only to their physical environment, but also to the vege-
tation, and, furthermore the relations which these animals bear to one
another. We wish also to consider both the prairie and the forest as
separate units, and to see how the animals are related to their physical
and biological environment. As previously stated, the special locali-
ties studied were described by stations both to give a precise and con-
crete idea of the prairie and its animals, as now existing in a limited
area, and also to preserve as much of the local color as the data would
permit. I wish now to reexamine these animals from another stand-
point, that of the animal association as a unit. The prairie as a whole
103
is not homogeneous from this point of view; it is a mosaic composed
of anumber of minor social communities. Each of these smaller units,
however, is fairly homogeneous throughout.
Our present knowledge of these minor associations is imperfect,
and for this reason they are arranged in an order approximating that
which we might reasonably expect to be produced if the initial stage
were made to begin with a poorly or imperfectly drained area and to
advance progressively with corresponding vegetational changes, toward
a more perfect condition of drainage. Upon the prairie a perfect series
would include every stage from lakes, ponds, and swamps to well-
drained dry prairie. But cultivation and drainage have obliterated so
much, that now only very imperfect remnants exist in the vicinity of
Charleston. Although the sequence followed therefore does not in-
clude all stages of the process it is approximately genetic.
There are three essential features in every animal association, or
community; certain physical conditions; certain kinds of vegetation,
which also modify the physical conditions; and representative kinds of
animals. Occasionally an effort is made to divorce these, to separate
organisms from their normal habitat, but such an effort is deceptive,
for no organism can live for any considerable period without a normal
environment.
I have not attempted to treat these associations with equal fullness.
In the sections devoted to the description of the stations it was possi-
ble in some cases, on account of the uniform character of a station, to
describe the animal association rather fully. In such instances the
detailed account is not repeated. In other cases I have elaborated the
community relations more fully here than elsewhere. The descriptions
of the stations and the associations, and the annotated lists, are in-
tended to be mutually supplementary.
II. THe Prairie ASSOCIATIONS
1. Swamp Prairie Association
The swamp prairie community lives in a habitat characterized by
shallow water, which stands approximately throughout the growing
season of the vegetation. The soil is black, and rich in vegetable de-
bris. The characteristic plants are bulrush (Scirpus), flags (Iris),
swamp milkweed (Asclepias incarnata), beggar-ticks (Bidens), and
young growths both of willow (Salix) and cottonwood (Populus del-
toides). The abundant growth of vegetation and the wet soil are con-
ditions favorable for the production and accumulation of organic de-
bris, which tends to fill the depressions and to supplement the inwash
104
from the surrounding slopes. At the same time, burrowing animals,
particularly the crawfish, also bury debris and work over the soil. In
the Charleston area this community was developed at Station I, d, and
in part at I, g.
The representative animals of this community are those living in
the water, such as the prairie crawfish, Cambarus gracilis (PI.
XXXVI), the snail Galba umbilicata, and such insects as the nine-
spot dragon-fly, Libellula pulchella (Pl. XXXVIII, fig. 2), and the
giant mosquito, Psorophora ciliata, whose immature stages are spent
in the water. In addition to these are other representative species
whose presence is, to an important degree, conditioned by the pres-
ence of certain kinds of vegetation—such species, for example, as
those which feed upon the dogbane (Apocynum), the brilliantly col-
ored beetle Chrysochus auratus; upon milkweed, the milkweed bugs
Lygeus kalmit and Oncopeltus fasciatus (Pl. XL, figs. 1 and 3), and
the milkweed beetle Tetraopes,; and, finally, the rather varied series of
flower visitors feeding upon pollen or nectar, such as the soldier-beetle
(Chauliognathus pennsylvanicus), Euphoria sepulchralis, and several
species of butterflies, moths, bees and wasps, including the honey-bee,
bumblebees, and carpenter-bee (Xylocopa virginica), and the common
rusty digger-wasp (Chlorionichneumoneum). Visiting the same flow-
ers, but of predaceous habit, were found the ambush spider (Misumena
aleatoria) and the ambush bug (Phymata fasciata). Small insects
were preyed upon by the dragon-flies (Libellula pulchella), and the
‘dragon-flies in turn were entangled in the webs of the garden spider
(Argiope aurantia).
No animals were taken on the flags, but Needham (’00) has made
an important study of the population inhabitating flags at Lake Forest,
Illinois, and shows that it is an extensive one. He gives an excellent
example showing how the injury by one insect paves the way for a
train or succession of others. For example: the ortalid fly Chetopsis
enea Wied. (Pl. XVIII, fig. 1), bores into the stem of the buds and
causes them to decay (Cf. Forbes, ’05, p. 164; Walton, Ent. News,
Vol. 19, p. 298. 1908). This condition affords a favorable habitat for
a pomace-fly (Drosophila phalerata Meig.*), an oscinid (Oscinis
coxendix Fitch, Plate XVIII, figures 3 and 4), a beetle, parasitic
Hymenoptera, and, after the decaying buds were overgrown by fungus
threads, the bibionid fly Scatopse pulicaria Loew. This paper by Need-
ham is one of the very few in which the population of a plant has been
studied as a biotic community. Forbes (’90, pp. 68-69; 02, p. 444)
has shown that snout-beetles (Sphenophorus ochreus Lec., Plate
*Mr. J. BR. Malloch infornts me that D. phalerata is not an American species.
105
XVIII, figures 5, 6, and 7) breed in root-bulbs of Scirpus, and that
these beetles eat the leaves of Phragmites. Webster (’90, pp 52-55)
observed these beetles feeding on the leaves of Scirpus and the larve
feeding on its roots. I have found great numbers of these beetles cast
up.on the beach of Lake Michigan. Evidently they breed in the
swamps about the lake, fall into it when on the wing, and are washed
ashore.
2. The Cottonwood Community
Ordinarily we are accustomed to think of the prairie as treeless,
and yet one large tree was relatively abundant upon the original prairie
of Illinois, particularly upon wet prairie, or, when pools were present,
even upon the uplands. This was the cottonwood, Populus deltoides.
These trees were often important landmarks when isolated; and today
the large trees or their stumps are important guides in determining the
former extent of the prairie. In the region studied there were no large
mature cottonwoods, although saplings were present, but north of
Charleston in the adjacent fields mature trees were found. They grow
normally at the margins of wet places, as about prairie ponds and
swamps, or along the small ill-defined moist sags and small prairie
brooks. This tree is usually solitary or in irregular scattered rows
when along streams, and does not, as a rule, form clumps. or’ groves.
This relatively isolated habit may be a factor in the comparatively
small number of invertebrates which are associated with it, or at least
in the amount of serious injury which they do to these trees upon the
prairie. Many of the larger trees are mutilated, or even destroyed by
lightning (Cf. Plummer, ’12), and such injury favors entrance of in-
sects on account of the rupturing of the thick bark.
The galls on the leaves and twigs of the trees often attract atten-
tion. A large irregular gall on the ends of the twigs becomes conspic-
uous in winter. This is formed by the vagabond gall-louse, Pemphigus
oestlundi Ckll. (Pl. XIX, fig. 1) (vagabundus Walsh, Ent. News,
Vol. 17, p. 34. 1906). I have found these galls abundant upon the
prairie at Bloomington, Ill. At this same locality I found a large
bullet-like gall at the junction of the petiole and the leaf—that of Pem-
phigus populicaulis Fitch (Pl. XIX, fig. 2), and at Urbana, Ill, on
other large prairie cottonwoods, a somewhat similar gall, on the side
of the petioles, caused by P. populi-transversus Riley (Pl. XIX, fig. 3).
I have also taken large caterpillars of the genus Apatela on leaves of
cottonwood, and September 3, at Urbana, upon its cultivated form, the
Carolina poplar, 4. populi Riley (Pl. XX, fig. 6). These caterpillars
have bodies covered by yellow hair penciled with black. At dusk
swarms of May-beetles (Lachnosterna) can be seen and heard feeding
106
among the leaves of the cottonwood and the Carolina poplar. It is
noteworthy that I have made these observations at Urbana, Illinois,
upon cottonwoods growing upon what was originally prairie.
Forbes (’07a) has shown, as the result of extensive collections of
May-beetles from trees, that they have a decided preference for Caro-
lina poplar (p. 456) and willow. This same paper also contains im-
portant observations on the nocturnal flights to and from the forest,
from the normal habitat of the grubs, and from the daytime abode of
the beetles in the open fields. Wolcott (’14) has recently emphasized
the point that the grubs live only in open places in proximity to wood-
land where the beetles can secure food. These observations show very
clearly that May-beetles are animals primarily of the prairie or forest
margin, and probably lived upon the original prairie, scattered, where
cottonwoods or willows grew. A glance at the map of the prairie and
forest (frontispiece) shows that the marginal area was very extensive,
and must have furnished an optimum habitat for these beetles. This is
a good illustration of the fact that the cottonwood exerted an influence
upon the prairie far beyond its shadow.
In some localities another beetle (Melasoma scripta Fabr.) feeds
upon the leaves of the cottonwood, and may become a serious pest to
poplars and willows, but I have not seen this species abundant on iso-
lated mature trees upon the prairie. I have taken these beetles (July
2) under cottonwoods at Bloomington, Il. Packard (’90, pp. 426-
474) has published a list of the insects known to feed upon Populus.
Willows (Salix) are frequently associated with the cottonwoods
upon the prairie, but, in marked contrast with these, they generally
grow in colonies and are eaten by a great variety of insects. Packard
(790, pp. 557-600) lists 186 species of insects on them, and Chitten-
den (’04, p. 63) extends the number to 380 species. Of course in any
given locality the number of species found will be relatively small, and
the number is further limited by the environmental conditions—
whether the land is upland or low and flooded. The degree of ‘prox-
imity of willows and cottonwood is likely to influence the relative
abundance of the insects feeding upon these trees, since a large number
of insects which feed upon willow also feed upon the cottonwood. Col-
onies of willow are thus likely to become sources of infestation for
the cottonwood; this relation, however, is a mutual one. Walsh (’64)
and Heindel (’05) have published very interesting studies of the com-
munity life of the insect galls on Illinois willows. Cockerell (’97, pp.
770-771) has listed the scale insects found upon willows and poplars.
107
3. Swamp-grass Association
The prairie swamp-grasses, slough grass (Spartina), and wild rye
(Elymus) were growing in relatively pure stands or colonies in de-
pressions which were dry in the late summer. The prolonged wetness
of the habitat and the dominance of the few kinds of grasses are char-
acteristic features of the environment of this association. These con-
ditions were found at Station I, a and c, north of Charleston. As these
stations were rather homogeneous and have already been discussed
somewhat fully, only a summary will be given here.
On account of the grassy vegetation the abundance of Orthoptera
is not surprising. Representative species are Melanoplus differen-
tialis, M. femur-rubrum, Scudderia texensis, Orchelimum vulgare,
Xiphidium strictum, CEcanthus nigricornis, and GE. quadripunctatus.
Other representative animals are Argiope aurantia and the swamp fly
Tetanocera plumosa. The list of species is probably very incomplete ;
during the wet season there are undoubtedly a number of aquatics;
furthermore, there are still other species which feed upon Spartina and
Elymus, particularly some Hemiptera, and stem-inhabiting Hymenop-
tera, and certain Diptera. Thus Webster (’03a, pp. 10-13, 26, 32, 38)
has recorded a number of chalcids of the genus Jsosoma which live
in the stems of Elymus virginicus and canadensis. In this same paper
he discusses their parasitic and predaceous enemies (pp. 22, 27, 33).
A fly also breeds in Elymus, the greater wheat stem-maggot, Mer-
omyza americana Fitch (Pl. XX, figs. 1-5), as recorded by Fletcher
(1. c., p. 48). This species is of economic importance, having spread
from grasses to the cultivated grains. It has been studied in Illinois
by Forbes (’84). He found a fly parasite of this species, and Webster
reports a mite preying on it. Webster (1. c., p. 53) reports another
fly, Oscinis carbonaria Loew, bred from Elymus by Fletcher.
In another paper Webster (’03b) has published a list of insects in-
habiting the stems of E. canadensis and virginicus. Osborn and Ball
(‘97b, pp. 619, 622; 97a) have discussed the life histories of certain
grass-feeding Jassid@ which feed upon Elymus. Osborn (’92, p. 129)
records a plant-louse, Myzocallis, from Elymus canadensis in Iowa,
and a species of leaf-hopper has been recorded by Osborn and Ball
(’97b, p. 615') from Spartina. On the same plant, Osborn and Sirrine
(’94, p. 897) record a plant-louse on the roots. In a list of the plant-
lice of the world and their food plants Patch (12) lists a few from
Spartina. This same list includes (pp. 191-206) many grasses an
the associated aphids, those on Elymus on page 196. :
108
4. Low Prairie Association
The moist black soil prairie, a degree removed from the wet or
swamp condition, with ground water in the spring relatively near the
surface, is fairly well characterized by the rosin-weed (Silphium), par-
ticularly S. terebinthinaceum. Other plants likely to be associated with
S. terebinthinaceum are Silphium laciniatum and S. integrifolium,
Eryngium yuccifolium, Lepachys pinnata, and, to a less degree, Lac-
tuca canadensis.
In the Charleston area this condition is represented by Station I, a,
north of the town, and Station III, a, and in part b, east of the town.
The proximity of ground water is shown at Station I, e, by the pres-,
ence of crawfish burrows, probably those of Cambarus gracilis. At
Station III the proximity of water was also evident where S. terebin-
thinaceum was most abundant in the railway ditches. Such perennial
plants are indicative of the physical conditions for a period of years,
and are thus a fairly reliable index of average conditions—much more
so than the annuals.
It is difficult to decide which kinds of animals are characteristic of
this kind of prairie. Provisionally I am inclined to consider the fol-
lowing as being so: Cambarus gracilis; Argiope aurantia; the grass-
hoppers Encoptolophus sordidus, Melanoplus differentialis, M. femur-
rubrum, Scudderia texensis, and Xiphidium strictum; Gicanthus nigri-
cornis; Phymata fasciata; and asilids. The presence of Lepachys was
clearly an important factor in determining the presence of Melissodes
obliqua and Epeolus concolor. At Station III, b, east of Charleston,
Epicauta pennsylvanica and Bombus pennsylvanicus, auricomus, and
impatiens were taken on the flowers of Silphinm terebinthinaceum.
Robertson (’94, pp. 463-464; ’96b, pp 176-177) has published lists
of insect visitors to the flowers of Silphium and Lepachys (’94, pp.
468-469), at Carlinville, Ill. Recently Shelford (13a, p. 298) has
published a long list of animals inhabiting Silphium prairie near Chi-
cago. Forbes (’g0, p. 75) has reported the snout-beetle Rhynchites
hirtus Fabr. as feeding upon Silphium integrifolium.
In a colony of prairie vegetation at Seymour, IIl., which included
much Silphium and Eryngium, the following insects were taken Octo-
ber 7 from the ball-like flower clusters of Eryngium yuccifolium: the
bugs Lygeus kalmii, Thyanta custator Fabr., Euschistus variolarius,
and Trichopepla semivittata Say (No. 539, C. C. A.), the last named
in large numbers, the nymphs in several sizes as well as the adults, a
fact which suggests that both may hibernate upon the prairie. Rob-
ertson (’89, pp. 455-456) has summarized his collections of insects
from Eryngium and on Euphorbia corollata (’96a, pp. 74-75).
109
Upon remnants of prairie vegetation growing at Urbana, Illinois,
I have found several kinds of insects centered about a wild lettuce,
Lactuca canadensis. Upon the upper, tender parts of this plant, the
plant-louse Macrosiphum rudbecki@ Fitch, thrives late in the fall, in
very large numbers. Some seasons nearly every plant is infested. The
lice become so abundant upon these tender parts that the entire stem
for a distance of a few inches is completely covered. They migrate
upward with the growth of the stem and keep on the fresh, tender
parts. Among the plant-lice, and running about on the stem of the
plant, attending ants abound; eggs, larve, and adults of lace-wing flies
(Chrysopa) also abound; and several species of coccinellids, syrphid
larve, and a variety of small parasitic Hymenoptera are present.
5. Upland Prairie Association
The well-drained prairie, a degree removed from the permanently
moist prairie, is fairly well represented by the physical and biological
conditions in which Euphorbia corollata, Apocynum medium, and
Lactuca canadensis, are the representative plants. The plant ecologist
would consider the conditions favorable to mesophytic plants. In the
Charleston region these conditions are approximated at Station II,
where drainage has doubtless changed the area from a somewhat
moist, to its present well-drained, condition. :
Representative animals of this community are as follows: Argiope
aurantia, Misumena aleatoria, Encoptolophus sordidus, Melanoplus
bivittatus, M. differentialis, Orchelimum vulgare, Xibhidium strictum,
Euschistus variolarius, Phymata fasciata, Chauliognathus pennsylvan-
icus, Epicauta marginata and E. pennsylvanica, Rhipiphorus dimidia-
tus and R. limbatus, Ammalo, Exoprosopa fasciata, Promachus verte-
bratus, Bombus pennsylvanicus, and Myzine sexcincta.
On dry prairie at Mayview, Ill., September 26, I found the plant-
louse Aphis asclepiadis Fitch on the leaves and stems of the dogbane
(Apocynum) and the lice attended by the ant Formica fusca L. A
beetle, Languria mozardi Latr., whose larva is a stem-borer, inhabits
Lactuca canadensis. Its life history and habits have been discussed
by Folsom (’o9, pp. 178-184).
6. The Solidago Community
A common community in the late summer and early fall is centered
about the goldenrod (Solidago). This plant was‘not abundant or in
blossom at any of the stations studied in detail, but it grew in small
widely scattered colonies or clumps. Observations were made in two
110
colonies, north of Charleston, both west of Station I, a, andI,g. The
collections made (Nos. 20, 26, 42, 43) are as follows:
Ambush Bug Phymata fasciata 20, 26
Stink-bug Euschistus variolarius 26
Black Blister-beetle Epicauta pennsylvanica 26
Noctuid moth Spragueia leo 20, 26
Conopid fly Physocephala sagittaria 26
Empidid fly Empis clausa 43
Halictid bee Halictus fasciatus 26
Myzinid wasp Myzine sexcincta 20, 26
Ant Formica fusca subsericea 20
It is important to know that these collections from Solidago were
made just as the flowers were beginning to blossom. Collections a few
weeks later would probably have given many more kinds. It should
be noted, too, that all these plants were far out upon the prairie and
far from woodlands—a factor which may influence to some extent
the kinds of visitors. As a rule the lists which have been published
state little or nothing at all as to the conditions in which the plants
were growing. If this factor is neglected, the presence of some vis-
itors remains puzzling. Thus on some goldenrods the locust beetle,
Cyllene robinie, is abundant; but this is conditioned in part by the
proximity of the yellow locust, which is absent on the Charleston
prairie.
Phymata was found copulating upon the flower, and with an em-
pidid fly, Empis clausa (No. 43), in its grasp. Two kinds of galls
formed by insects were found on this plant: one formed by the fly
Cecidomyia solidaginis (No. 43), which forms a rosette of leaves;
and the other the spindle-like stem-gall, formed by a small caterpillar,
Gnorimoschema gallesolidaginis (No. 7462 Hankinson). September
20 the moth Scepsis fulvicollis Hiibn. was found in goldenrod flowers
near Station I,a. Its larva feeds on grass. snyoaiophy
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155
4. Destruction of trees by animals: the processes of defolia-
tion, borings in branches, bark, trunk, or roots, and the girdling of
trees. Fires started by man, depending on the degree of destruction,
cause new cycles of succession. Both beavers and man build dams,
flood areas, and thus kill trees.
5. Combinations of physical and organic processes; the flood-
ing of river bottoms by driftwood rafts which become converted into
dams and thus submerge large areas.
Since it is most usual for these causes to act, not singly but in
various combinations, and since they also vary greatly in their degrees
of influence, their operation is extremely complex. ‘The drowning of
the forests along the Mississippi River through the sinking of the
land by the New Madrid earthquake, is a good example, showing how
a large tract of forest may be killed and much dead and decayed wood
formed, as has been shown by Fuller (’12)—(Plate XXXII). Tarr
and Martin (’12) have shown how destructive to forests the earth-
quakes are in Alaska. The influence of the New Madrid earthquake
upon animal life has not been investigated, but it is not too late even
today, after more than one hundred years, to make important studies
on this subject. On the other hand, the processes of erosion operate
more continuously than the periodic earthquakes, and tend to degrade
the land, lower the water-level, and to change the habitats in swamp
and other forests.
The results of climatic influences are seen in the amount of injury
done by sleet, which, weighing down the branches, breaks many of
them and leaves the fractured stubs as favorable points for attack
by fungi and insects. Webb (’09) has shown that when a tornado
passed through Mississippi and Louisiana the felled pine forests were
from one to three miles wide. Practically all of this timber became
infested with the larve of Monohammus titillator Fabr. After a
severe frost in Florida the dead wood of the orange-trees became in-
fested by wood-boring larve, which spread from this wood to the
enfeebled living wood, as Hubbard (Howard, ’95) observed. Light-
ning (Plummer, ’12) kills and maims many trees, producing dead
wood, and through fires started in the same manner much more dam-
age is done. Hopkins (’o09) considers that much of the injury at-
tributed to fire is primarily due to insects which made the dead and ~
dry fuel for the destructive fire work.
That competition among trees weakens some of them is well
known. This weakening makes them more susceptible to attack by
fungi and insects. In a forest where the shade-enduring trees can
shade out all competitors, the shrubs and trees which are intolerant
show just such a lack of resistance. As an example of this process
156
the following case may be cited: Mr. W. P. Flint informs me that
he has observed that shaded, suppressed white oaks in southern IIli-
nois are much more heavily infested by the bark-louse Aspidiotus
obscurus Comstock, and by the beetle Phymatodes varius Say than
are the vigorous trees.
Trees may be injured and killed by animals in many ways, as by
defoliating them, boring in the twigs, trunk, or roots, and by the de-
struction of the bark and sap-wood of the trunk. Of injuries caused
by insects the work of defoliators of hardwoods is one of the most
conspicuous kinds. Repeated defoliation of elms by the elm leaf-
beetle Galerucella luteola Mill. will, according to Felt (’05, p. 61),
so weaken a tree that Tremex columba finds suitable food in its dis-
eased and dying substance. With T'reme-x present its parasite Thalessa
also arrives. The maple borer, Plagionotus speciosus Say, may also
weaken a tree and pave the way for Tremex and Thalessa. A study
of the after effects of the prominent defoliators of shade and forest
trees, such as the fall web-worm (Hyphaniria cunea), the white-
marked tussock-moth (Hemerocampa leucostigma, Plate XXXII, figs.
3, 4 and 5), the bag-worm (Thyridopteryx ephemereformis), the
larch saw-fly (Nematus erichsonii), the gypsy moth (Porthetria
dispar), and the brown-tailed moth (Euproctis chrysorrhea), would
doubtless throw much light upon the details of successions caused by
insects. I have not been able to learn that this subject has been studied
carefully in this country. Such injuries are clearly not limited to
hardwoods, for many similar observations have been made in conif-
erous forests. Hewitt (’12, p. 20) has listed some of the beetles
which follow the defoliation of larches by the larch saw-fly. Hop-
kins (’o1, pp. 26-27) found that the spruces of New England were
being killed by the bark-beetle Dendroctonus piceaperda Hopkins;
that following the damage done came other beetles, such as Polyg-
raphus rufipenms Kby., which attacks the weakened tops of the
trees, following the attack of its predecessors on the trunk or base;
and that also, following Dendroctonus, came Tetropium cinnamop-
terum Kby., which mines in the dead trees. The yellow pines of the
West are killed by the bark-beetle Dendroctonus ponderosa, and this
is followed by many kinds of insects which live on the decaying bark
and wood, as Hopkins (’02, pp. 10-16) has shown. He also states
(‘o9, p. 68) that in the Appalachian Mountains Dendroctonus fronta-
lis Zimm. killed a large part of the trees in an area “aggregating over
75,000 square miles.” Such examples of multiple attack show the
complexity of the causes influencing forest life. When the great
amount of influence which insects are able to exert and do exert upon
forests is considered, the question is raised as to what may be their
157
influence in determining the kind of trees that compose what the
plant ecologists (Cowles and others) consider the climax forest of
eastern North America—the maple-beech forest. It has long been
known (Packard, ’90, p. 515) that the beech has remarkably few in-
sect enemies, perhaps about fifty species being recorded. Its associate,
the hard maple (Acer saccharum), has many more, and the oaks and
hickories, which are largely absent from the climax forest and char-
acterize the changing stages preceding the climax, are preyed upon
by more insects than any other of our trees, their number possibly
equaling the sum total of all the other forest-tree insects.
A good example of the combined influence of physical and organic
factors is seen in the huge rafts of driftwood which have accumulated
in the Red River of Louisiana and Arkansas (Veatch, ’06)—(Pls.
XXX and XXXIV)—on such an extensive scale that hundreds of
acres of the bottoms were flooded and the forests killed, producing
vast quantities of dead and decaying wood. With the opening of
the drainage canal, connecting Lake Michigan with the Illinois River,
the bottoms were so flooded that willows, maples, cottonwoods, etc.,
on the lowest ground were killed along the river for many miles, and
presented a view similar to that shown on Plate XXXV._ In this
manner vast quantities of dead and decaying wood have been made
available as food and habitat for wood-inhabiting invertebrates.
7. Interrelations within the Forest Association
The dependence of the animal upon the physical and organic en-
vironment is primarily a phase of the problem of maintenance. In
the forest these relations are so intricate, and involve the lives of so
many kinds of animals, that a forest, like the prairie, must be looked
upon as a mosaic composed of a vast number of smaller animal, or
biotic communities, each one not only interrelated at many angles
within itself, but similarly connected with the other communities of
the forest. Walsh (’64, pp. 549-550) has given us a graphic ac-
count, not of the forest as a_whole but of one of its smallest units—
those which he found clustered about the galls of willow trees, the
willow leaf-gall community. He says:
“Nothing gives us a better idea of the prodigious exuberance of
Insect Life, and of the manner in which one insect is often dependent
upon another for its very existence, than to count up the species which
haunt, either habitually or occasionally, one of these Willow-galls,
and live either upon the substance of the gall itself or upon the bodies
of other insects that live upon the substance of the gall. In the single
gall S [alicis]. brassicoides n. sp. there dwell the Cecidomyia which
158
is the maker of the gall—four inquilinous Cecidomyia—an inquilinous
saw-fly (Hymenoptera)—five distinct species of Microlepidoptera,
some feeding on the external leaves of the gall, and some burrowing
into the heart of the cabbage, but scarcely ever penetrating into the
central cell, so as to destroy the larva that provides them with food
and lodging—two or three Coleoptera—a Psocus (Pseudoneuroptera)
—a Heteropterous insect found in several other willow-galls—an
Aphis which is also found on the leaves of the willow, but pecu-
liarly affects this gall—and preying on the Aphides the larva of
a Chrysopa (Neuroptera) and the larva of a Syrphide (Diptera)—
besides four or five species of Chalcidide, one Braconide Ichneumon
(Hymenoptera) and one Tachinide (Diptera), which prey on the
Cecidomyia and the Microlepidoptera—making altogether about two
dozen distinct species and representing every one of the eight Or-
ders. . . . If this one little gall and the insect that produces it
were swept out of existence, how the whole world of insects would
be convulsed as by an earthquake! How many species would be com-
pelled to resort for food to other sources, thereby grievously disar-
ranging the due balance of Insect Life! How many others would
probably perish from off the face of the earth, or be greatly reduced
in numbers! Yet to the eye of the common observer this gall is noth-
ing but an unmeaning mass of leaves, of the origin and history of
which he knows nothing and cares nothing!’
With this conception of a community in mind it is only necessary
to refer to the following diagram (Fig. 18) to see how immaterial
it is as to where one begins to take up this thread of interrelations,
for sooner or later every animal and plant in the association will have
to be passed in review and its influence recognized as a response to
its conditions of life.
ECOLOGICALLY ANNOTATED LIST
I. Pratrre INVERTEBRATES
An exhaustive study of the animal ecology of a region or an as-
sociation must be based upon a thorough investigation of the ecolog-
ical relations of the individual animals composing it. An ideal an-
notated list in an ecological paper should, therefore, include for each
species a complete account of its life history, its behavior, its physi-
ology, and the structural features which would in any way contribute
to an understanding of the response of the animal to its organic and
inorganic environment. At present we have no such knowledge of
the animals of ‘any locality or of any complex association of animals.
159
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In a preliminary study, like the present one, it is desirable to record
rather fully the observations made in the region studied, because
we have so few descriptions of the conditions of life on our prairies.
An effort has been made to give for each species the date of obser-
vation or collection, the locality or “station’’ where found, observa-
tions on habits and life history, and the field numbers of the speci-
mens secured. These numbers illustrate how observations may be
accumulated, upon a large number of individuals, without the ob-
server's being familiar with them, or even knowing their scientific
names. '
It is really surprising how little is recorded about some of the
commonest animals of the prairie and forest in zoological literature.
Other animals, particularly’ those of economic importance, are
treated rather fully, but generally with little relation to their natural
environment. In this list it has been considered desirable not to
give an extended account of each kind of animal, but to refer to
some of the most important literature concerning it, so that one may
gain some general idea of the ecological potentialities of each kind of
animal.
MOLLUSCA
PHyspz
Physa gyrina Say.
Three half-grown young and an adult shell were taken among
swamp milkweed, Asclepias incarnata (Sta. I, g), Aug. 11 (No. 19).
All show distinct varices; the last one formed on the adult shell is
very distinct. ‘These scars mark a period of rest or slow growth
which was probably due to hibernation or the drying-up of the
swamp. Physa, asa rule, can not endure such extreme desiccation as
can Lymnea, and to that degree is indicative of a more permanent
water supply. Our specimens were all dead, but some of them so
recently that fly maggots came from them.
LYMN2IDA
Galba umbilicata (C. B. Adams).
A single specimen of this small snail was taken among swamp
milkweeds (Sta. I, d) Aug. 11 (No. 18). Mr. F. C. Baker, who de-
termined the specimen, writes me that this is the first record of this
species for Illinois. Baker remarks (’11, p. 240) that this species is
“abundant in still water in sheltered borders of rivers, in small
brooks, ditches, and streams, and in shallow overflows. Clings to
dead leaves or other submerged debris, or crawls over the muddy
161
bottom of its habitat, in shallow water. Associated with Galba
obrussa, Aplexa hypnorum, and the small planorbes (Baker). In
ditches and brooks in pastures (True). Common in damp places
and in ditches along roads where water collects only in rainy weather
(Nylander ).”
Our specimen was taken where the water was very shallow (only
a few inches deep) and overgrown with vegetation. This species ap-
pears to be a strictly shallow-water marginal form, and has consider-
able power of enduring desiccation.
CRUSTACEA
ASTACIDA
Cambarus gracilis Bundy. Burrowing Prairie Crawfish. (Pl. XXXVI.)
The prairie crawfish was abundant at Sta. I, d, on the wet parts
of the prairie. T. L. Hankinson dug some specimens from their
holes, which proved to be of this species. Specimens were captured
Apr. 23, 1911, and Aug. 9, 1910 (No. 7442).
Crawfish burrows were observed to traverse the dense yellow
clay with which the railway embankment had been built over a
swampy place at Sta. I,d. Burrows were also observed at Sta. I, e,
among the colony of Stlphium terebinthinaceum and Lepachys pin-
nata, and also at Station I, g.
I have found the characteristic claw of this species on wet prairie
along the railway track at Mayview, Ill. At this time, September 26,
1912, burrows with fresh earth were numerous, far from any stream.
(No. 482, C. C. A.)
Cambarus diogenes Girard. Diogenes Crawfish.
Crawfish of this species were taken by T. L. Hankinson at Sta.
I, d (No. 8047A). The presence of this chimney builder at this sta-
tion suggests that the numerous chimneys shown in Figure 2, Plate
IIIB are in part the work of this species though they are in part also
the work of gracilis.
ARACHNIDA
PHALANGIIDA
PHALANGIDA
Liobunum politum Weed. Polished Harvest-spider. (Pl. XXXVII,
fig. 3.
Two small phalangiids, both probably of this species, were found
under moist wood upon the prairie (Sta. I, g) Aug. 8. Concerning
162
these specimens, Mr. Nathan Banks writes me that they are “young,
not fully colored, but probably Liobunum politum Weed.”
Weed (’91) reports that this rather rare species occurs in fields
and forests, and is seldom found about buildings. He has found it
among river driftwood, and says (’92a, p. 267): “It sometimes oc-
curs under boards in fields, and is often swept from grass and low
herbage.’’ When disturbed it emits, as do others of its family, a
liquid with a pungent odor. Weed (’91) has made some observa-
tions on its breeding habits. He notes that in confinement it ate
plant-lice.
L. formosum Wood was taken by me upon the lodged drift-
wood of a small brook on the border of a forest at White Heath,
Ill, May 4, 1911. (No. 505, C.C.A.) This species, according to
Weed (’89, p. 92), hibernates as an adult.
ARANEIDA
EPerDz
Argiope aurantia Lucas (=riparia Hentz). Common Garden Spider.
(Pl. XXXVII, figs. 1 and 2.)
This is very abundant, and the most conspicuous spider on the
prairie. Found among the prairie grasses (Sta. I, g) Aug. 8 and 12
(Nos. 6 and 39); in its web among goldenrod, Solidago (Sta. 1),
Aug. 12 (No. 26); among the swamp grasses (Sta, I,a) Aug. 28
(No. 179); and among Elymus (Sta. I,c) Aug. 24 (No. 153);
from sweepings made in the colony of Lepachys pinnata (Sta. I, e)
Aug. 12 (No. 40); and on the Loxa prairie (Sta. II) Aug. 13 (No.
49), Aug. 27 (No. 178), and Aug. 28 (No. 179); in an open
area in the upland Bates woods (Sta. IV,a) Aug. 17 (No. 93);
and in an open glade in the lowland forest (Sta. IV,c) Aug. 22
(No. 143). In its webs in the swamp-milkweed colony (Sta. I, d)
Aug. 9 the large dragon-fly Libellula pulehella Drury was found en-
trapped; a grasshopper, Melanoplus differentialis Thomas, was also
found entrapped (Sta. I,a) Aug. 28 (No. 179); and a large butter-
fly, Papilio polyxenes Fabr., was discovered (Sta. I, d) Aug. 12 (No.
45).
The openness of an area rather than its prairie character appears
to determine the habitat of this spider. This is evidenced by its
presence in open spaces within the forest. It flourishes in gardens
for similar reasons. Years ago I found this species very abundant
in the late summer and fall at Bloomington, IIl., in an asparagus bed,
after the plants had been allowed to grow up and form a rank mass
‘163
of. vegetation. This species has received considerable study.
McCook (’90) and Porter (’06) record many observations on this
species. Howard (’92b) has discussed its hymenopterous parasites
and those of some other spiders.
No specimens of Argiope transversa Emerton, the transversely
black-and-yellow-banded relative of aurantia, were observed at
Charleston, although they are fairly abundant in colonies of prairie
vegetation near Urbana, e. g. at Mayview, IIl:, Sept. 26, and on Nov.
26, 1911. I have seen this species only among colonies of prairie
vegetation along railway rights-of-way.
THOMISIDA a
Misumena aleatoria Hentz. Ambush Spider.
' This crab-like flower spider was abundant upon flowers: on the
mountain mint, Pycnanthemum flexuosum (Sta. I,g), Aug. 8 (No.
6); on the mint, (Sta. 1) with a giant bee-fly, Exoprosopa fasciata
Macq., Aug. 12 (No. 31); on the Loxa prairie (Sta. II) with the
same kind of fly, Aug. 13 (No. 47); on the prairie (Sta. I, g) on the
flower of the swamp milkweed, Asclepias incarnata, Aug. 24 (No.
157) with a male bumblebee, Bombus separatus Cress.; on Andropo-
gon (Sta. I, g) with a large immature female of Conocephalus, Aug.
24 (No. i159); on the Loxa prairie (Sta. II) on flowers of Eryn-
gium yuccifolium, Aug. 27 (No. 178); in the colony of Elymus
(Sta. I, a) Aug. 28 (No. 179); and in the open glade of the low-
land Bates woods (Sta. IV,c) on the flowers of Eupatorium celes-
tinum, with a very large syrphid fly, Milesia ornata Fabr. (=virgin-
iensis Drury), Aug. 26 (No. 184). These insects captured by the
spiders vary from about five to ten times the size of their captor. There
is considerable variation of color in this series of spiders,
It would be well worth while for some one to make a special
study of this spider, and give us an account of its methods of cap-
turing food and finding fresh flowers, with a full account of its life
history. McCook (’90, Vol. 2, pp. 367-369) gives some informa-
tion about the habits of an allied species of spider, but the account is
meager. Some observations on the breeding habits of this species
have been made by Montgomery (’o9g, p. 562); and Pearse (’I1)
has recently published the results of an interesting study of the rela-
tion between the color of these spiders and the color of the flowers
they frequent. He concludes that althouch this spider may change
its color slowly (from yellow to white), it does not do so with
rapidity or in such a way as to match its surroundings, and, further,
that it does not seek an environment or a flower colored like itself.
164
He finds, however, that on white flowers, white spiders occur gen-
erally, that on yellow flowers, yellow spiders occur, and also that
upon flowers of colors other than white and yellow, such as purple,
pink, and blue (p. 93), white spiders predominate.
ATTIDA,
Phidippus sp.
This jumping spider was taken Aug. 12 (No. 34) on the common
milkweed, Asclepias syriaca, along the railway tracks (near Sta.
T, a), and when captured had in its jaws fragments of what seemed
to be Diabrotica 12-punctata Oliv.; but as the fragments were lost
during the process of capture, this determination was not made
certain.
ACARINA
TROMBIDIIDAL
Trombidium sp. Harvest-mites. Chiggers. (Pl. XXI, figs. 1 and 2.)
These are the immature six-legged stage of a mite or mites which
when mature have eight legs. The young are parasitic on insects
(Banks, Proc. U. S. Nat. Mus., Vol. 28, pp. 31-32, 1904); -the
adults prey upon plant-lice and caterpillars; one species also eats
locusts’ eggs.
These mites were very abundant on the prairie north of Charles-
ton (Sta. I), and became such a pest that relief had to be sought,
in a liberal application of flowers of sulphur to our legs and arms,
as is recommended by Chittenden (’06).
INSECTA
ODONATA
LIBELLULIDA
Sympetrum rubicundulum Say. Red-tailed Dragon-fly.
This dragon-fly was taken in the prairie grass zone (Sta. I, g)
Aug. 8 (No. 4.) It is one of our commonest kinds. The nymphs
live in small bodies of standing water. The adults forage for small
insects in open places, along hedge rows, and in open forest glades.
For the habitats of dragon-fly nymphs, reference should be made
te Needham (Bull. 68, N. Y. State Mus., p. 275. 1903). William-
son (00, pp. 235-236) has observed robber-flies carrying this species,
and has found this and other species of dragon-flies in the webs of
the spider Argiope.
165
ep aa Drury. Nine-spot Dragon-fly. (Pl. XXXVIII,
©, 2,
Individuals were abundant in both colonies of swamp milkweeds
(Sta. I, d and g) and several were seen entrapped in webs of Argiope
aurantia (Sta. I,d) Aug. 9. This is one of the most abundant of
our large dragon-flies. It frequents small bodies of water and slug-
gish pond-like streams. Williamson has taken it also in the webs of
Argiope. ‘This large powerful insect is able to do considerable dam-
age to a spider-web and then make its escape. Among the milk-
weeds (Sta. I,d) an individual was seen by T. L. Hankinson to
escape from a web. This dragon-fly, like most of its kind, captures
small insects on wing; one kind, however, is reported to have dug a
cricket out of the ground (Psyche, Vol. V, p. 364. 1890).
NEUROPTERA
MyYRMELEONIDZ
Brachynemurus abdominalis Say. Adult Ant-lion.
A single specimen was taken along the railway track north of
Charleston (near Sta. I,g) Aug. 12 (No. 36). This is a species
which frequents dry habitats. The larva is unknown, but is prob-
ably predaceous—as other ant-lion larve are and as the adult is sup-
posed to be.
Two adult females were taken July 19 and 20, 1907, at Cincin-
nati, Ohio, in my room, to which they were attracted by the electric
light. Another female was taken Aug. 8, 1901, at Gate City, Vir-
ginia (near Big Moccasin Gap). Determined by R. P. Currie.
CHRYSOPIDA
Chrysopa oculata Say. Lacewing. (Pl. XX XVIII, fig. 1.)
A single specimen of this insect was taken among prairie grasses
(Sta. I,g) Aug. 12 (No. 44). The larve feed upon plant-lice, and
the adults are also considered predaceous. Howard (Proc. Ent.
Soc., Wash., Vol. 2, pp. 123-125. 1893) has given a list of their
numerous hymenopterous parasites. Mr. T. L. Hankinson captured
one also (Sta. I) July 3, 1911 (No. 7665). Fitch (’56) published
many observations on the members of this genus; and Marlatt (’94a)
has written on the life history of this species.
166
ORTHOPTERA
AcrIpupz
Syrbula admirabilis Uhler.
One specimen of this grasshopper was found in the tall prairie
grasses blue-stem Andropogon and Pamcum (Sta. I,g) Aug. 8 (No.
3). Morse (’04, p. 29) says this species frequents “open country”
and is “common in upland fields amid Andropogon and other coarse
grasses.”
Encoptolophus sordidus Burm. Sordid Grasshopper. (Pl. XXXIX,
fig. 1.) ;
One nymph of this species was taken in the prairie-grass colony
north of Charleston (Station I, g) Aug. 12 (No. 44); another (No.
158) on Aug. 24 in the colony of Lepachys pinnata (Sta. I, e) ; and
an adult (No. 48) Aug. 13 at Loxa (Sta. II, a) from the flowers of
Silphium integrifolium.
This is a species characteristic of dry open places, where the
vegetation is low. The peculiar snapping sound made by the male
when on wing is quite characteristic. (Cf. Hancock, ’11, pp. 372-
373-)
Dissosteira carolina Linn. Carolina Grasshopper. (Pl. XXXIX,
fig. 4.)
A very reddish specimen of this species was taken in a cleared
bottom forest at River View Park, about three miles southeast of
Charleston, Aug. 19 (No. 95). Many specimens were observed in
the pasture above the “Rocks,” on the Embarras River about three
miles east of Charleston. These individuals exhibited to a marked
degree the hovering, undulating flight which is so characteristic of
this species during the hot days of summer and early autumn. Town-
send (Proc. Ent. Soc. Wash., Vol. 1, pp. 266-267. 1890) has made
interesting observations on this habit, and finds that it is mostly the
males which participate in this courting ceremony, as he considers it.
There appears to be more or less of a gathering of individuals when
one of the locusts performs. There were perhaps half a dozen per-
forming in the colony observed at the “Rocks.” Townsend (Can.
Ent., Vol. 16, pp. 167-168. 1884) has considered this flight as re-
lated to breeding. Some one might study this subject with profit,
and determine its meaning. Poulton’s paper “On the Courtship of
certain Acridiide” (Trans. Ent. Soc. London, 1896, Pt. II, pp. 233-
252) might prove helpful in this connection.
This species seems to have been influenced by man to a marked
degree. Its original habitat appears to have been natural bare spots,
167
such as sandy beaches, banks of streams, sand-bars, and burned
areas. In a humid forested area such places are usually in isolated
patches, or in more or less continuous strips as along shores; but
- since the activities of man produce large cleared areas and bare
spots, such as roads, railways, and gardens, the favorable area of
habitat for this species has been vastly increased. Consult Han-
cock (’I1, pp. 340-347) for observations on the habits of this
species.
Schistocerca alutacea Harr. Leather-colored Grasshopper. (PI.
XXXIX, fig. 3.) :
One specimen of this large grasshopper was taken east of
Charleston, on the prairie which grades into the forest (Sta. ITI, a)
Aug. 15 (No. 59). Morse (’04, p. 39) and Hart (’06, p. 79) rec-
ognize that this species lives among a rank growth of vegetation
and brush. In general the local conditions are open or transitional,
and may be compared to those of a shrubby forest margin, and not
to those of the distant open prairie or to conditions within the for-
est. (Cf. Hancock, ’11, pp. 366-370.)
Melanoplus bivittatus Say. ‘Two-striped Grasshopper. (Pl. XL.
fig. 3.)
This grasshopper was taken from flowers of the rattlesnake-
master, Eryngium yuccifolium, on the prairie at Loxa (Sta. II),
Aug. 13 (No. 55). It is a little surprising that it was so rare this
season on the prairie areas examined, as it is usually a common
species. Hancock (’11, pp. 356-359) has discussed this grasshopper.
Melanoplus differentialis 'Thomas. Differential Grasshopper. (PI.
XXXIX, fig. 5, and Pl. XL, fig. 1.)
This species was generally common in open areas, especially on
the prairie, but was also found in open places in the forest. It was
very abundant in the colonies of swamp prairie grasses, Spartina
and Elymus (Sta. I,a), Aug. 28 (No. 179); in the upland prairie
grasses, as Andropogon and Panicum (Sta. I, g), Aug. 12 (No. 39);
and in colonies of Lepachys (Sta. I,e) Aug. 12 (No. 40); also at
Loxa on Silphium integrifolium (Sta. II, a) Aug. 13 (No. 48).
This must be considered as one of the most common and char-
acteristic of prairie animals. Notwithstanding the destruction of
the original prairie, its habitat has been perpetuated, particularly
upon waste and neglected areas, such as fence rows, roadsides, rail-
way rights-of-way, and vacant city lots.
168
Melanoplus femur-rubrum DeG. Red-legged Grasshopper. (PI.
XXXIX, fig. 2.)
This species also is one of the most common and generally dis-
tributed insects upon open areas. It was found among the prairie
grasses Andropogon and Sporobolus (Sta. I. g) Aug. 8 and 12 (Nos.
3 and 39); in the Lepachys colony (Sta. I,e¢) Aug. 12 (No. 40);
and in Elymus and Spartina (Sta. I, a and c) Aug. 24 and 28
(Nos. 153, 179, and 180). As Hart (’06, p. 8r) has remarked, it
is common in cultivated areas. Cultivation appears to be distinctly
favorable to it; differentialis, on the other hand, seems to thrive best
in waste places. .
Locustipa
Scudderia texensis Sauss.-Pict. Texan Katydid.
This is the common and characteristic katydid of the prairie
areas. It was found (Sta. I, g) among the tall swamp milkweeds
Aug. 8 (No. 2); in the tall blue-stem Andropogon and in Panicum
Aug. 12 (No. 44); in the Lepachys colony (Sta. I, e) Aug. 12 (No.
40); and among the swamp prairie grasses Spartina and Elymus
(Sta. I,@ and c) Aug. 28 (Nos. 179 and 180). Consult Hancock,
*II, pp. 330-331, for the life history of this species.
Conocephalus sp., nymph.
A large female nymph was secured on blue-stem Andropogon
(Sta. I,g) Aug. 24 (No. 159), having been captured by a crab-
spider, Misumena aleatoria Hentz.
Orchelimum vulgare Harr. Common Meadow Grasshopper. (PI.
XL, figs. 2 and 4.)
This grasshopper was taken east of Charleston on the flowers of
broad-leaved rosin-weed, Silphium terebinthinaceum (Sta. III), Aug.
26 (No. 175); on the Loxa prairie (Sta. II) Aug. 27; on the flow-
ers of rattlesnake-master, Eryngium yuccifolium (No. 178); and
on the prairie north of Charleston from the colony of wild rye,
Elymus (Sta. I, a), Aug. 28 (No. 179). A squeaking individual
(No. 180) captured here confirmed observations made in other
places—particularly in the tall prairie grasses Andropogon and
Sporobolus (Sta. I, g), where the first specimen (No. 3) was taken
Aug. 8. Nymphs, very probably of this species, were also in the
prairie grasses Andropogon and Sporobolus (Sta. I, g) Aug. 8 (No.
3); and Aug. 28 (Nos. 179 and 180) in the swamp grasses Elymus
and Spartina (Sta. I,a,c). This species is preeminently a tall-grass
frequenter, whose penetrating zeeing during the sunny hours serves
to locate grass plots and low, rank weedy growths.
169
Blatchley (’03, p. 384) has observed the species feeding on small
moths, and once saw an individual on goldenrod eating a soldier-
beetle, Chauliognathus pennsylvanicus DeG. Forbes (’05, p. 144)
reports that its food consists mainly of plant-lice, and leaves of grass,
fungus spores, and pollen. It is thus evident that it eats both animal
and vegetable food.
me attenuatum Scudd. Lance-tailed Grasshopper. (Pl. XL,
g. 7.)
On the prairie at Loxa (Sta. II), on flowers of the arrow-leaved
rosin-weed, Silphium integrifolium, a single individual of this species
was found Aug. 13 (No. 48).
According to Blatchley (’03, pp. 380-381) it frequents the coarse
vegetation bordering wet places. He also states that the eggs are
placed between the stems and leaves of “tall rank grasses.”
Xiphidium strictum Scudd. Dorsal-striped Grasshopper. (PI. XL,
fig. 6.)
This prairie species was taken on prairie clover, Petalostemum
(Sta. I, b>), Aug. 11 (No. 21); in sweepings among the cone-flower,
Lepachys pinnata (Sta. 1,¢), Aug. 20 (No. 40); on the mountain
mint Pycnanthemum flexuosum (Sta. I) Aug. 12 (No. 35); on P.
flexuosum or P. pilosum (Sta. II) Aug. 13 (No. 57); among the
swamp grasses Elymus and Spartina (Sta. I,a and c) Aug. 28 (Nos.
179, 180) ; on the Loxa prairie on Silphium integrifolium (Sta. II)
Aug. 13 (No. 48); and on purple prairie clover, Petalostemum pur-
pureum (Sta. II), Aug. 13 (No. 50).
Forbes (’05, p. 147) gives its food as plant-lice, fungi, pollen
and, larsely, other vegetable tissues. He also states that it frequents
the “drier slopes in woods and weedy grounds” (p. 148).
’ G@RYLLIDA
Gicanthus nigricornis Walk. Black-horned Meadow Cricket. (PI.
XL, fig. 5, Pl. XLI, figs. 1 and 2.)
This prairie cricket was taken in sweepings from the cone-flower
(Lepachys pinnata) colony (Sta. I,e) Aug. 12 (No. 40); on the
transitional prairie east of Charleston (Sta. HI, b) Aug. 15 (No.
62); and from the swamp cord-grass, Spartina (Sta. I,a), Aug. 28
(No. 179).
Blatchley (’03, p. 451) says: “In August and September, nearly
every stalk of goldenrod and wild sunflower along roadsides, in open
fields or in fence corners, will have from one to a half dozen of these
insects upon its flowers or branches. It is also especially abundant
170
upon the tall weeds and bushes along the borders of lakes and ponds,
and in ‘sloughs and damp ravines.”
Blatchley (l.c., p. 452) made some incomplete observations on
the peculiar courting habits of this species, a subject which has been
elaborated by Hancock (’05). Hancock also describes the method
of oviposition. The female first gnaws the plant stem; then bores a
hole and deposits an egg; and next, again gnaws the stem. The eggs
are laid in stems of blackberry, goldenrod, and horseweed (Leptilon).
Houghton (Ent. News, Vol. 15, pp. 57-61. 1904) has published
interesting observations on the carnivorous habits of nymphs of
G. niveus DeG. Cf, Parrott and Fulton, ’14.
Ashmead (Insect Life, Vol. 7, 241. 1894) reports that G. nigri-
cornis (fasciatus) is preyed upon by the wasp Chlorion harrisi Fernald
(Isodontia philadelphica St. Farg.).
CGicanthus quadripunctatus Beut. Four-spotted White Cricket.
This prairie species was found among the tall prairie grasses
blue-stem Andropogon and Panicum (Sta. I,g) Aug. 8 (No. 3);
and among the colony of cord grass, Spartina (Sta. I,a), Aug. 28
(No. 179). ; ;
Blatchley (’03, p. 453) reports it on “shrubbery and weeds in
fence-rows and gardens; and along roadsides.” ‘This indicates how
a prairie species adjusts itself to the conditions produced by man.
Parrott (Journ. Econom. Ent., Vol. 4, pp. 216-218. 1911) gives
figures of the eggs of this species and describes its method of ovipo-
sition in raspberry stems.
HEMIPTERA
Cicappz
Cicada dorsata Say. Prairie Cicada.
Although this species was not taken at Charleston, a single speci-
men (No. 185) was captured at Vera, Fayette county, Ill., Septem-
ber 1, on a giant stool of blue-stem Andropogon. Osborn (Proc.
Iowa Acad. Sci., Vol. 3, p. 194. 1896) reported one specimen from
Iowa; Woodworth, (Psyche, Vol. 5, p. 68. 1888) says: “On the
prairies, Illinois to Texas”; and MacGillivray (Can. Ent., Vol. 33, p.
81. 1901) adds Missouri, Colorado, and New Mexico.
_ MEMBRACDE
Campylenchia curvata Fabr.
This bug was taken in sweepings made in the colony of cone-
flower, Lepachys pinnata (Sta. I, e), Aug. 12 (No. 40).
171
JassDz
Platymetopius frontalis Van D.
This leaf-hopper was taken in sweepings in the cone-flower col-
ony (Sta. I,¢) Aug. 12 (No. 40). '
APHIDIDA
Microparsus variabilis Patch.
This plant-louse infests the leaves of the Canadian tick-trefoil,
Desmodium canadense, and causes the leaves to curl. Quite a colony
of these plants found infested (near Sta. I, f) Aug. 24, were stunted
and deformed by these plant-lice (No. 160). Consult Patch (Ent.
News, Vol. 20, pp. 337-341. 1909) for a description of the insect
and a plate showing the injury which it causes; also Williams (Univ.
Studies, Univ. Neb., Vol. 10, p. 76, 1910) and Davis (ibid., Vol. 11,
p. 28. 1912).
Aphis asclepiadis Fitch. Milkweed Plant-louse.
Plant-lice of this species were abundant upon the younger ter-
minal leaves of the common milkweed, Asclepias syriaca, along the
railway track north of Charleston (Sta. I) Aug. 12 (Nos. 28, 29,
and 154). Associated with them were workers of the ants Formica
fusca Linn. var. subsericea Say (Nos. 28, 29, and 154) and For-
mica fusca Linn. (No. 28). Ona milkweed plant which lacked the
plant-lice were found associated another ant, Formica pallide-fulva
Latr., subsp. schaufussi Mayr, var. incerta Emery, and the metallic-
colored fly Psilopus sipho Say.
At Urbana, Ill, a very abundant plant-louse on wild lettuce,
Lactuca canadensis, is Macrosiphum rudbeckie Fitch (det. by J. J.
Davis). The upper, tender branches of these plants are in the fall
covered with vast numbers of these lice, both wingless and winged.
That this species feeds upon a number of other prairie plants is a
point of much interest because of their distinctly prairie character.
It is reported from Vernonia, Solidago, Bidens, Ambrosia, Cirsium,
Silphium, and Cacalia (Thomas, Eighth Rep. State Ent. Ill, p. 190.
1879).
PENTATOMIDA:
Euschistus variolarius Beauv. (Pl. XLI, fig. 3.)
This common plant-sucking bug was taken on flowers of the
swamp milkweed, Asclepias incarnata (Sta. I,d), Aug. 9 (No. 12);
from the blue-stem Andropogon colony (Sta. I,g), where a large
robber-fly, Promachus vertebratus, was taken astride a grass stem
with one of these bugs in its grasp Aug. 12 (No. 39); at Station
172
I by T. L. Hankinson, July 3, 1911 (No. 7665) ; on the Loxa prairie
(Sta. II), with insects from flowers of the purple prairie clover,
Petalostemum purpureum, Aug. 13 (No. 50); and on flowers of the
mountain mint Pycnanthemum pilosum or P. flexruosum (Sta. II),
Aug. 13 (No. 52). Consult Forbes (’05, pp. 195, 261) for a sum-
mary of its life history, and references to literature. It feeds upon
a great variety of plants (Olsen, in Journ. N. Y. Ent. Soc., Vol. 20,
Pp. 53. 1912) and on soft-bodied insects.
Stiretrus anchorago Fabr. (Pl. XLI, fig. 5.)
This highly colored bug was taken, Aug. 23 (No. 146), not
upon the prairie proper but at the margin of the Bates woods (near
Sta. IV,a), where the clearing had been so complete that only
sprouts and young trees occurred, associated with many plants which
frequent open, sunny places, such as ironweed (Vernonia) and
Pycnanthemum pilosum.
This bug sometimes feeds upon the larve of the imported as-
paragus beetle, Crioceris asparagi (Chittenden, Circ. No. 102, Bur.
Ent., U. S. Dept. Agr., p. 6. 1908). This circular contains figures
of the nymph and adult. Olsen reports it as feeding upon cater-
pillars and beetle larve and on the plants Asclepias and Rhus (Jour.
N. Y. Ent. Soc., Vol. 20, pp. 55, 56. 1912).
THYREOCORIDA
Thyreocoris pulicarius Germ. Flea Negro-bug. (Pl. XLII, fig. 2.)
This negro-bug was taken on the flowers of goldenrod, Solidago
(near Sta. I,a), Aug. 12 (No. 26). Forbes and Hart (’oo, p. 100)
state that this insect abounds on Bidens, a plant which grew in great
abundance near the goldenrod referred to. Taken (Sta. I) by T. L.
Hankinson July 3, 1911 (No. 7665).
Lyreana
Ligyrocoris sylvestris Linn.
This insect was taken while sweeping vegetation in the cone-
flower (Lepachys) colony (Sta. I, e) Aug. 12 (No. 40).
Lygeus kalmii Stal. Small Milkweed Bug. (Pl. XLII, fig. 1.)
This is one of the commonest insects found upon milkweeds of
the prairie. Specimens were taken on the flowers of the swamp
milkweed, Asclepias incarnata (Sta. I, g), Aug. 8 (No. 1); on flow-
ers of the mountain mint, Pycnanthemum flexuosum (Sta. I, g),
Aug. 8 (No. 6); and on swamp milkweeds (Sta. I,d) Aug. 9
(No. 12).
173
This is another common insect about which very little is known.
Its food plants and life history are worthy of study. I have taken
this species from Mar. 20 (adult, 1894) to Nov. 4.-(adult, 1893) at
Bloomington, Ill.; at Havana, IIl., during August; and at Chicago
June 8 (1902). That it probably hibernates in the adult stage is
shown by the fact that I captured an adult as early as Mar. 22 at
Urbana, Ill. This bug, like the squash-bug (Anasa), may have
an active migratory period in the fall, and only those individuals
survive the winter which happen to be in favorable places when the
cold weather sets in. I have captured this bug in the dense Brown-
field woods (Urbana), where it was crawling on a log Oct. 12 (No.
312, C.C.A.). Hart (’07, p. 237) records it from Asclepias cornuti
ne syriaca) at Havana in the sand area, and:also from Teheran,
Illinois.
Oncopelius fasciatus Dall. Large Milkweed Bug. (Pl. XLII, fig. 3.)
This large red plant-bug I took but once—on flowers of the
swamp milkweed, Asclepias incarnata (Sta. I, g), Aug. 8 (No. 1);
T. L. Hankinson, however, captured another specimen (Sta. I) July
3, 1911 (No. 7665).
I have found it in years past abundant on prairie colonies of
niilkweed at Bloomington, Ill., from June into September, and at
Havana and Chicago during August. On Sept. 26, at Mayview, IIl.,
along the railway among prairie plants this plant-bug was found on
dogbane (Apocynum). ) from the upland forest to the lowland Aug.
22 (No. 133); and a female was taken among leaves on the ground
in the upland forest (Sta. IV, a) Aug. 23 (No. 150). T. L. Hankin-
son found an adult and a nymph in the Bates woods June 28, 1911
(No. 7678). (Cf. Hancock, ’11, pp. 362-364.)
The positive heliotropism or negative geotropic response shown
in diseased grasshoppers is of interest. It may be caused either by a
fungous or bacterial disease. (Cf. Gillette, Bull. No. 6, n. s., Div. Ent.
U. S. Dept. Agr., pp. 89-93. 1896. )
Morse (’04, p. 15) considers this an exceptional ground-inhabiting
or geophilous species since it is “an inhabitant of xerophytic forests as
well as of open fields, and in the Southern States is found quite as
often in the forest as on the open plain.”
Melanoplus differentialis Thomas. Differential Grasshopper.
Consult the list of prairie invertebrates, p. 167.
Melanoplus atlanis Riley. Lesser Grasshopper. (Pl. LVII, fig. 8.)
A single specimen was taken on the ground in the upland forest
(Sta. IV, a) Aug. 16 (No. 67). The open character of parts of this
dry forest affords favorable conditions for this species.
Morse (’04, pp. 19, 42) considers this a characteristic species
of open country, but “likely to be found anywhere.” Hancock (’11,
pp. 415-416) has described the habitat of this species.
214
Melanoplus amplectens Scudd.
This locust and nymphs doubtfully regarded as of the same
species were taken from the ground, mainly among leaves, in the up-
land forest (Sta. IV, a) Aug. 16 (No. 67); other collections are as
follows: in the glade in the lowland forest (Sta. IV, c) Aug. 20 (No.
117); on the open ravine slope (Sta. IV, b) Aug. 22, a male (No.
124a) ; and on the same date, in the glade of the lowland forest (Sta.
IV, c), a nymph and an adult female (No. 143).
This is the largest of the short-winged locusts in the forest, and
an abundant species. Morse (’04, pp. 19, 50, Pl. 7) described its
haunts as in thickets, forest margins, open forests, and occasionally
in grassy clearings and fields.
Melanoplus gracilis Bruner.
Two males were found Aug. 20 in a glade in the lowland forest
(Sta. IV,c) where there was a luxuriant cover of vegetation, and
nettles and Eupatorium calestinum abounded; and Aug. 22, in the
same location, one female was found (No. 143).
The wings are very rudimentary in this species. Hart (’06, p.
82) describes its habitat as follows: “On tall grasses and weeds in
ravines and about marshes, masses of wild vines along railroads,
weedy growths in the beds of small streams, and in like situations.”
These conditions are found in open areas with an abundance of vege-
tation.
Melanoplus obovatipennis Blatch.
This small species, similar to scudderi, was found in the upland
forest (Sta. IV,a) Aug. 17 (No. 93). A nymph taken Aug. 22
from the forest (Sta. IV) is doubtfully regarded as of this species
(No. 124).
Hart (06, p. 81) gives the habitat of this species as “High
wooded hillsides throughout Ilinois.”” Blatchley (’03, p. 308) states
that it frequents ““for the most part, high, dry, open woods, espe-
cially those in which beech and oak trees predominate.” He further
states that in a dry season it may be found associated with Dichro-
morpha viridis and Truxalis brevicornis “among the reeds and tall
rank grasses near the borders of marshes.”
Melanoplus scudderi Uhl. Scudder’s Grasshopper.
A single female was found in the open glade in the lowland for-
est (Sta. IV,c) Aug. 20 (No. 117); and a nymph taken Aug. 22
from the open ravine slope (Sta. IV, b) is doubtfully referred to this
species (No. 124).
215
Hart (’06, p. 81) describes the habitat of this grasshopper as
“open woods and thickets, and along rail fences and roadsides.”
Species which now characterize our open, partly cleared woodlands,
in the primeval forest probably frequented forest margins, bluffs,
and the borders of streams, or open patches in woods where a tree
had fallen, and similar situations. With a thinning out of the for-
est (up to a certain degree) their habitat is increased in area, but
when by clearing the woods disappear, their habitat vanishes.
LocustTIp
Scudderia furcata Bruner. Forked Katydid. (Pl. LVI, fig. 5.)
One female was taken in an open area in the upland forest on
low shrubs (Sta. IV, a) Aug. 20 (No. 109). Another specimen was
taken near Vera, Fayette county, III., on a finely developed colony of
prairie vegetation among Andropogon, Sept. 1 (No. 185).
Blatchley (’03, p. 349) states that it is “most frequently seen on
the low bushes a trees about the margin of thickets and along
fence rows, but in the prairie country north [in Indiana] it frequents
coarse grasses and weeds.”
Amblycorypha rotundifolia Scudd. Round-winged Katydid. (PI.
LVII, fig. 2.)
A single female of this species was taken in the glade in the low-
land forest (Sta. IV,c) Aug. 20 (No. 117); and also a freshly
emerged female (No. 143). Blatchley (’03, p. 352) states that this
is “more of a terrestrial species than oblongifolia, being often seen
on the ground, or on clumps of tall grass and weeds which grow in
damp ravines.” Hart (’06, p. 84) says that this species is found
“On grasses and weeds in damp ground.”
Microcentrum laurifolium Linn. Angle-winged Katydid. (Pl. LVII,
figs. 1 and 2.)
Males were found on hickory sprouts at the cleared margin of
the upland forest (near Sta. IV.a) Aug. 22 (No. 135). They
were chirping loudly, in the early afternoon, on sprouts less than
two feet high.
Cyrtophyllus perspicillatus Linn. Common Katydid. (Pl. LVIII,
fig. 1.)
One male was taken in the partly cleared area bordering the for-
est (near Sta. IV,a) Aug. 23 (No. 145). Here, among stump
sprouts of hickory, oak, and young sassafras, about two to three
feet high, stood this mate stridulating in the sun at 2:30 p. m., but
the note did not seem exactly normal, that is, as when heard at night.
216
This species is so distinctly arboreal and nocturnal that I was sur-
prised to find it stridulating during the day, and so near the ground.
I have camped for days in a grove where these insects made a great
din at night, but found none on the low vegetation or on the ground
(as at Kappa, Ill). Years ago a large colony flourished in Franklin
Park at Bloomington, Ill.
Conocephalus nebrascensis Bruner. Nebraska Cone-nose.
A female was taken in the glade in the damp lowland forest (Sta.
IV,c) Aug. 20 (No. 117).
The female of this species has been observed to oviposit “between
the stem and root-leaves of Andropogon”, a typical prairie plant, but
little appears to be recorded of its habitat. A large nymph of this genus,
and probably of this species (No. 159), was taken on the prairie
grass Andropogon (Sta. I,g) Aug. 24. It had been captured by the
crab-spider Miswmena aleatoria Hentz (No. 159).
Orchelimum cuticulare Redt.
A specimen was taken in the upland forest (Sta. IV,a) Aug. 16
(No. 67); another, from the open areas of the upland forest (Sta.
IV,a) Aug. 17 (No. 93); and a third, from the glade in the damp
lowland forest (Sta. IV,c) Aug. 22 (No. 143). All of these were
males.
Orchelimum glaberrimum Burm.
This insect was found in abundance in the glade in the lowland
forest (Sta. IV,c) Aug. 20 (No. 117), and a nymph was taken in
the same place Aug. 22 (No. 143).
The abundance of this species in this damp area, with its pro-
fusion of low vegetation, indicates that the conditions were fav-
orable.
Xiphidium nemorale Scudd.
Nymphs and adults were found in the glade in the lowland for-
est (Sta. IV, c) Aug. 20 (No. 117) and Aug. 22 (No. 143); in the
openings in the upland forest (Sta. IV,a) Aug. 17 (No. 93), and
Aug. 20 (No. 103).
Blatchley (’03, p. 374) states that it abounds along the “borders
of dry, upland woods, fence rows, and roadsides, where it delights to
rest on the low shrubs, blackberry bushes, or coarse weeds usually
growing in such localities.”
GrYLLDZz
Nemobius fasciatus DeG. Striped Cricket. (PI. LVIII, fig. 6.)
Nymphs of this species were found in the upland forest on the
217
ground (Sta. IV,a) Aug. 16 (No. 67); in the upland forest area
also, in an open place, was found a short-winged male Aug. 17 (No.
93); along a path in the upland forest, among dry leaves, a short-
winged female was taken Aug. 22 (No, 122); and an abundance of
short-winged males and females, and nymphs (No. 143) were found
Aug. 22 in the glade in the lowland forest (Sta. IV, c).
This small cricket is generally abundant among the litter on the
forest floor.
Nemobius maculatus Blatch. Spotted Cricket.
A nymph was taken in the upland forest (Sta. IV,a) among
leaves Aug. 22 (No. 122).
Blatchley (’03, p. 425) states “It is found in low open woods,
usually in the vicinity of or beneath logs”; Hart, (’06, p. 89) states
that it is found “About logs and dead wood in sparse woods and near
streams.”
Apithus agitator Uhl. Woodland Cricket.
A nymph was taken from the open area in the upland forest (Sta.
IV, a) Aug. 17 (No. 93); another from an open ravine slope (Sta.
IV, b) Aug. 22 (No. 124). No adults were secured.
Blatchley (’03, pp. 458-459) records this species as from forests,
noting its preference for prickly ash. It is also recorded as from
grape-vines and dense shrubbery. The females deposit eggs in the
twigs of the white elm, Ulmus americana Linn.
HEMIPTERA
CicapIDz
Cicada linnet Grossb. (Cicada tibicen L.). Dog-day Harvest-fly.
PL. LV, fig; 5.)
This insect was found at the cleared margin of the upland forest
(near Sta. IV, a) on low hickory sprouts Aug. 26 (No. 162).
It is said to require two years to mature. T. L. Hankinson re-
ports that Tibicen septendecim L, (Pl. LV, figs. 3 and and 4) was
found about Charleston in 1907, and branches scarred by the oviposit-
ing females were observed in the Bates forest (Sta. IV, a).
Felt (’05, pp. 237-238) describes the emergence of the adult
Tibicen from the nymph skin. For the recent synonymy see Smith
and Grossbeck (Ent. News, Vol. 18, pp. 116-129. 1907).
FuLGoRIDA
Ormenis pruinosa Say (?). Mealy Flata. (Pl. LVI, figs. 1 and 2.)
This insect was taken by T. L. Hankinson June 28, 1911, in the
218
Bates woods (No. 7678). It.appears to feed upon a large variety of
trees, shrubs, and herbaceous plants. Its normal habitat is probably
in open woods or the forest margin. Swezey (’04, pp. 8-9) gives
full references to the life history of this insect and a list of the food
plants.
TETTIGONIELLIDA
Aulacizes irrorata Fabr. (Pl. LVI, fig. 3.)
A few specimens were taken, the collection data being as follows:
from an open glade in the lowland forest (Sta. IV, c) Aug. 20 (No.
117); and from the smaller branches of sassafras bushes (Sta. IV, c)
Aug. 22 (No. 143).
This insect is often taken on grapes, and in the South on cotton.
Sanderson (Bull. 57, Bur. Ent., U. S. Dept. Agr., p. 58. 1906)
describes briefly the egg-laying habits and figures the adult insect.
Gypona pectoralis Spangb.
This species was taken June 28, 1911, in the Bates woods (Sta.
IV) by T. L. Hankinson (No. 7678).
PENTATOMIDA
Euschistus fissilis Uhl.
This bug was taken in Bates forest (Sta. IV) Aug. 22 (No.
124). It has been known to feed upon wheat (Webster, Rep. U. S.
Dept. Agr., 1885, p. 317). It also feeds upon corn, and on the moth
Aletia. It is parasitized by the proctotrypid Trissolcus euschisti
Ashm. (Olsen, in Journ. N. Y. Ent. Soc., Vol. 20, p. 52. 1912).
Mormidea lugens Fabr.
A nymph of this bug was taken by T. L. Hankinson in the Bates
woods (Sta. IV) June 28, 1911 (No. 7678).
Hymenarcys nervosa Say.
This insect was taken on the ground from among dead leaves and
decayed wood which had drifted to the mouth of a ravine in the low-
land forest (Sta. IV, c) Aug. 20 (No. 113). In the South this insect
feeds upon cotton.
Mirpz
Lygus pratensis Linn. Tarnished Plant-bug.
This bug was taken in the Bates woods (Sta. IV) June 28, IQII,
by T. L. Hankinson (No. 7678). See prairie list, page 175.
219
CorEIDz
Alydus quinquespinosus Say.
This bug was taken by T. L. Hankinson June 28, 1911, in the
Bates woods (No. 7678), and July ro (No. 7693) on the under-
growth in the woods (Sta. IV).
Acanthocerus galeator (Euthoctha) Fabr. (Pl. LVI, fig. 8.)
Six large nymphs of this plant-bug were taken on the apical part
of a tall herb, Actinomeris alternifolia Linn., growing in the open
glade of the lowland forest (Sta. IV, c; Pl. XIV) Aug. 29 (No. 182).
This bug has been reported to suck the juice from the plum, and
it injures the tender parts of orange plants. Hubbard (Insects Af-
fecting the Orange, U. S. Dept. Agr., Div. Ent., p. 163. 1885) gives
figures of the adult insect and describes briefly the eggs and young.
Forbes and Hart (’00, p. 445) have summarized the little that is
known of this insect.
Jalysus spinosus Say. Spined Stilt-bug. (Pl. LVI, fig. 7.)
This bug was found Aug. 20 in the open glade of the lowland for-
est (Sta. IV, c), where there was a luxuriant growth of herbaceous
vegetation (No. 117). It was also taken (Sta. IV) by T. L. Hank-
inson June 28, 1911 (No. 7678). Lugger reports it from oak woods.
It feeds upon plants.
GERRDz
Gerris remigis Say. Water-strider. (PI. L, fig. 2.)
This water-strider was abundant in the pools of the small tem-
porary stream in the ravine bordering the southern part of the Bates
woods (Sta. IV, d) Aug. 22 (No. 129).
It is an important enemy of mosquito larvee.
REDUVIDA
Sinea diadema Fabr. Rapacious Soldier-bug.
A nymph of this predaceous bug was captured by T. L. Hankin-
son in the Bates woods (Sta. IV) June 28, 1911 (No. 7678). See
list of prairie animals, page 173.
CoLEOPTERA
CICINDELIDA:
Cicindela unipunctata Fabr. Woodland Tiger-beetle.
One specimen of this tiger-beetle was taken along the path through
the cleared area as it entered the forest (Sta. IV,a) Aug. 22 (No.
136).
220
Tiger-beetles are generally most abundant in open places, but this
beetle seems to be a woodland species like the brilliantly colored C.
sexguttata Fabr. Wickham (’99, pp. 210-211) records unipunctata
from wooded areas. It is rare and difficult to catch, and is said to
be nocturnal in habit.
CARABIDAR
Calosoma scrutator Fabr. Caterpillar-hunter.
This common arboreal beetle was taken Aug. 16 (No. 64) in the
upland Bates wood (Sta. IV, a), where it attracted attention by the
rustling sound it made in crawling among the dry leaves on the
ground. Specimens of these beetles I could easily secure by remain-
ing quiet and listening for this rustling of the leaves. One specimen
was seen to crawl up the trunk of a small oak-tree, three or four inches
in diameter, for about seven feet. Another individual I took from
my neck. It may have fallen upon me from a tree, but more prob-
ably it climbed upon me as it does a tree. In woods adjacent to the
Bates forest, a caterpillar-hunter (No. 97) was found Aug. 20 with
what appeared to be the damp cast skin of some large bombycid larva,
which was also claimed by an ant, Camponotus herculeanus Linn.,
subsp. pennsylvanicus DeG., var. ferrugineus Fabr. On the ravine
slope (Sta. IV, b) Aug. 20 T. L. Hankinson captured one of these
beetles (No. 100) with a caterpillar about an inch long, which it had
partly mangled in the thoracic region with its formidable jaws. On
the upper slopes of the ravine (Sta. IV, b) Aug. 23 another beetle
(No. 149) was found on the ground under a hickory tree, eating a
Datana larva. Along this same rather open forested slope another
individual was observed to run from the ground up the trunk of a
small white oak (six or seven inches in diameter) for three or four
feet, and then to return to the ground. The climbing individuals ob-
served took a relatively straight course up the trunk, making no ef-
fort to climb in a spiral direction, and made the descent head fore-
most.
‘At Bloomington, Ill, while picking cherries I have taken the
beetle in trees. Although the arboreal habit is evidently very well
developed in this species, it is also very much at home on the ground.
The rapidity and apparent ease with which it ran over dry oak leaves
in the upland Bates woods surprised me.
The active foraging habits of this beetle are well shown by Her-
man’s observations (Journ. Cincinnati Soc. Nat. Hist., Vol. 21, p.
80. 1910) on its killing nestlings of the cardinal grosbeak (Cardin-
alis cardinalis) in bushes three feet from the ground. Harris (In-
221
sects Injurious to Vegetation, p. 470. 1869) states that it preys upon
canker-worms, both on the ground and by ascending trees.
Galerita janus Fabr.
A specimen was found under the bark of a decaying log in the
upland Bates forest (Sta. IV,a) Aug. 23 (No. 171). This common
beetle is frequently found in such situations, and seems to’ have a
preference for relatively damp places. I have taken the adult as
early as March 23 under bark of logs in the sap-wood stage of decay
at Urbana, Ill., where it was found associated with single dealated
females of Camponotus herculeanus pennsylvanicus, Passalus cornu-
tus, pyrochroid larve, the caterpillar Scolecocampa liburna, and the
slug Philomycus carolinensis.
This species is a fairly common one. I found it abundant at
Bloomington, Ill., where it was taken April 15, May 1, and June 22.
The larva has been described by Hubbard (Psyche, Vol. 1, pp.
49-52. 1875).
CoccINELLIDA:
A species of lady-beetle was found upon a fungus growing on a
stump in the upland forest (Sta. IV,a) Aug. 17 (No. 81). Asso-
ciated with the beetle on the fungus were large numbers of the snail
Pyramidula perspectiva.
ELATERIDA
Melanotus sp.
A larva belonging to this genus (No. 125) was found Aug. 22
under the bark of a decaying stump (Sta. IV. 6) in which the sap-
wood was destroyed, the remainder being sound though discolored.
It was associated with the slug Philomycus carolinensis and the
caterpillar Scolecocampa liburna.
Corymbites sp.
A larva belonging to this genus (No. 113) was found in drifted
leaves and dead wood at the mouth of a ravine in the lowland for-
est (Sta. IV, c).
Asaphes memnonius Hbst.
This click-beetle was taken at the mouth of a ravine in the low-
land forest (Sta. IV,c) Aug. 20 (No. 113) in drift composed of
dead leaves and rotten wood.
LAMPYRIDE
Calopteron terminale Say. Black-tipped Calopteron.
This interesting beetle was taken in the damp lowland forest (Sta.
IV, c) Aug. 26 (No. 173).
222
This species has been mentioned as an instance of mimicry because
of its resemblance in shape and color-pattern to the syntomid moth
Lycomorpha pholus Drury. Both are found in damp shady woods.
Calopteron reticulatum Fabr. Reticulate Calopteron. (PI. LVIII,
fig. 4.)
A single specimen was taken—in the glade in the lowland forest
(Sta. IV,c) Aug. 22 (No. 143).
The larva and pupa of this species are described by Coquillett
(Can. Ent., Vol. 15, pp. 97-98. 1883). July 10 he found a pupa
“suspended by the hind end of its body beneath a log.”
Photuris pennsylvanica DeG. Pennsylvania Firefly. (Pl. LVIII,
fig. 3.)
This large firefly was taken June 28, 1911, in the Bates woods
(Sta. IV) by T. L. Hankinson (No. 7678).
McDermott (’10, ’11) Knab (’05), and Mast (’12) should be
consulted for discussions on the natural history and ecology of our
fireflies. McDermott gives many observations on P. pennsylvanica.
Chauliognathus marginatus Fabr. Margined Soldier-beetle.
This predaceous beetle was taken June 28, 1911, in the Bates
woods (Sta. IV) by T. L. Hankinson (No. 7678). (Cf. Lintner,
Fourth Rep. Injurious and other Ins. N. Y., 1888, pp. 74-88.) This
is a predaceous species in the larval stage, feeding on immature in-
sects. The adults feed on pollen (Riley, in Fifth Rep. Ins. Mo., p.
154. 1873).
Telephorus sp.
This was taken June 28, 1911, in the Bates woods (Sta. IV) by
T. L. Hankinson (No. 7678). See T. bilineatus, Pl. XLIV, fig. 1.
LucanDa
Passalus cornutus Fabr. Horned Passalus. (Pl. LVIII, fig. 5.)
This common woodland beetle was found under the bark of a
decaying stump on the slope of a ravine (Sta. IV,b) Aug. 17 (No.
85). One specimen, with a chestnut thorax and yellowish wings,
had just shed the pupal skin. Another, a fully matured specimen,
carried a large colony of mites. Ewing (Univ. Studies, Univ. IIL,
Vol. 3, p. 24. 1909) states that nymphs of uropod mites are often
attached to insects for transportation. It has generally been as-
sumed that they are parasitic.
This Passalus seems to be one of the most common insects found
in decaying logs and stumps. I have found it very abundant at
223
Bloomington, Ill. The beetles evidently hibernate, for I have taken
them at Urbana, IIl., as late as October 18, and as early in the spring
as March 23.
This beetle invades logs and stumps as soon as the sap-wood be-
gins to be well decayed, and evidently advances into the log with the
progress of decay. As it invades logs in the sap-wood stage of decay,
it is often associated with newly founded colonies of the ant Cam-
ponotus herculeanus pennsylvanicus, pyrochroid larve, the slug Phil-
omycus carolinensis, and the caterpillar Scolecocampa liburna. For
physiological studies of cornutus see Schafer (Mich. Agr. Coll. Exper.
Sta., Tech. Bull. No. 11. 1911).
ScaRABAIDS
Geotrupes splendidus Fabr. Splendid Dung-beetle.
This dung-beetle was dug from a hole, an inch or so below the
surface, in the hard clay of the pathway near the margin of the for-
est bordering the cleared area (Sta. IV, a) Aug. 22 (No. 120). As
cattle and horses were pastured in this forest, its presence is readily
accounted for.
Pelidnota punctata Linn. Spotted Grape Beetle.
Only one specimen of this beetle was taken. It was found on a
grape leaf (Sta. III, b) Aug. 15 (No. 58). This insect is primarily
a forest or forest-margin insect. The larva feeds upon the decaying
roots and stumps of oak and hickory. The adult devours leaves of
the grape and of the Virginia creeper.
Many undetermined scarabeid larve were found in a much-de-
cayed stump in the ravine near the small temporary stream (near
Sta: IV, d) Aug. 22 (No. 130).
CHRYSOMELIDZ
Chrysochus auratus Fabr. Dogbane Beetle.
This characteristic species of the prairie (No. 103) was taken
Aug. 20 in an open place in the upland oak-hickory forest (Sta.
IV, a) on the dogbane Apocynum medium. See list of prairie inver-
tebrates, p. 178.
Cryptocephalus mutabilis Mels.
This leaf-beetle was taken June 28, 1911, in the Bates woods
(Sta. IV) by T. L. Hankinson (No. 7678). It has been reported on
Ceanothus, Viburnum, hazel, and oak by J. B. Smith. Evidently this
is a woodland beetle.
224
Coptocycla clavata Fabr. Clubbed Tortoise-beetle.
This leaf-beetle was taken in the south ravine of the Bates woods
(Sta. IV. b) by T. L. Hankinson June 28, 1911 (No. 7678). It
is known to injure the potato, tomato, eggplant, and bittersweet.
The larve and adults feed upon the same kinds of plants (Lintner,
Sixth Rep. Injurious and other Ins. N. Y., pp. 126-127. 1890).
TENEBRIONIDAS
Boletotherus bifurcus Fabr. Horned Fungus-beetle. (Pl. LIX, figs.
I, 2, and 3.)
This curious-looking beetle was found on the shelf-fungus Polyp-
orus in the lowland forest (Sta. IV, c) Aug. 26 (No. 173).
I have found this species very abundant near Bloomington, TIL,
where at times it was difficult to find an example of Polyporus which
was not thoroughly honeycombed by the larvz of these beetles. A
single shelf has been found to contain several beetles. They were
generally discovered within galleries excavated within the fungus.
On July 11 in such a shelf I found larve and pupe in abundance.
Other dates of capture are June 3 and July 6. Riley and Howard (In-
sect Life, Vol. 3, p. 335. 1891) also report it from Polyporus. Fig-
ures of the larva and pupa are given by Packard (’83, p. 474) and
descriptions by Gissler (On coleopterous larve of the family Tene-
brionide, Bull. Brooklyn Ent. Soc., Vol. 1, pp. 85-88. 1878).
Meracantha contracta Beauv.
Larve of this beetle were taken under dry leaves in the upland
forest (Sta. IV,a) Aug. 17 (No. 83); and others from under damp
leaves at the base of the wooded slopes of a ravine leading to the low-
land forest (Sta. IV, b) Aug. 22 (No. 140). The latter larvee were
associated with the ant. Siigmatomma pallipes. These larve are
often confused with wireworms (Elateridae). ‘
I found the beetles occasionally in the forest at Bloomington, IIL,
June 13; and Aug. 1 on the papaw.
I have a specimen of this larva, in very rotten wood, showing
the sinuous larval boring (Pl. XXX), from the Brownfield woods,
Urbana, Ill. (March g; collector, D. M. Brumfiel). Wickham has
described and figured the larva (Journ. N. Y. Ent. Soc., Vol. 4, pp.
TIQ—121. 1896).
PyYrocHROoDa
Pyrochroa sp.
A single specimen of a larva belonging to the above family was
taken August 22 (No. 130) in the ravine (Sta. IV, b) from under
225
the bark of a decaying stump, in company with numerous scarabzid
larve. These larve are very characteristic animals—under bark
when decay has loosened it from the sap-wood. The accompanying
figure (Pl. LIX, fig. 4) shows the general appearance of this larva
and of an adult beetle. I found Dendroides canadensis Latr. fairly
abundant at Bloomington, Ill, July 25. Larve belonging to this
family have been taken in the Brownfield woods, Urbana, Ill., under
the bark of decaying trees. It is a representative animal species in
this habitat.
See Moody (Psyche, Vol. 3, p. 76. 1880) for descriptions of
pyrochroid larve.
LEPIDOPTERA
PaPILIONIDZ
Papilio philenor Linn. Philenor Butterfly. (Pl. LIX, fig. 5.)
The caterpillar was found crawling upon the ground in the up-
land forest (Sta. IV,a) Aug. 16 (No. 69). Aug. 26 a larva (No.
166) which had attached itself to the stem of a prickly ash (Sta.
IV, b), was just entering upon the pupal stage, but had not yet cast
the larval skin.
The larva feeds upon Dutchman’s pipe, Aristolochia—a plant
which was not observed in the forest.
Fapilio turnus Linn. Turnus Butterfly.
The butterfly was observed on wing Aug. 16 in the open glades
of the upland forest (Sta. IV, a).
The larva feeds upon Prunus and Liriodendron.
Papilio cresphontes Cram. Cresphontes Butterfly.
The butterfly was observed in the open spaces of the upland
forest on wing Aug. 16.
The larva feeds upon Zanthoxylum, Ptelea, Dictamnus, Citrus,
etc.
Papilio troilus Linn. Troilus Butterfly.
The butterfly was taken, on wing, from the open slope of the
south ravine (Sta. IV,b) Aug. 22 (No. 161); and in the upland
forest (Sta. IV,a) Aug. 26 (No. 163).
The larva feeds upon sassafras and Laurus.
NyYMPHALIDZ
Polygonia interrogationis Fabr.
The butterfly was taken in the open glade in the lowland forest
(Sta. IV, c) Aug. 20 (No. 117).
The larva feeds upon Humulus, Ulmus, and Urtica.
226
AGAPETIDA
Enodia portlandia Fabr. Portlandia Butterfly.
This woodland butterfly was taken in the Bates woods (Sta. IV)
Aug. 15 (No. 63) and on June 28, 1911 (No. 7678), by T. L. Han-
kinson.
The larva feeds upon grasses. Fiske (’01, pp. 33-34) gives a
good description of the haunts of this species. Years ago I found it
abundant near Bloomington (Orendorf Springs), Ill., in dense, damp,
shady woods. It is as characteristic of shade as most species are of
sunshine.
Cissia eurytus Fabr. Eurytus Butterfly.
This is also a woodland butterfly. It was taken in the Bates
woods by T. L. Hankinson June 28, 1911 (No. 7678). The larva
feeds upon grass.
LyYca&NIDz
Everes comyntas Gdt.
This small blue butterfly was taken on the open upper slopes of
the wooded south ravine in the Bates forest (Sta. IV, 5) Aug. 22
(No. 161).
The larva feeds upon red clover and Desmodium.
HESPERIIDA *
Epargyreus tityrus Fabr. Common Skipper.
This caterpillar was found in the open. glade in the lowland for-
est (Sta. IV,c), folded within a leaf of sassafras, Aug. 26 (No.
173).
I have taken this butterfly many times at Bloomington, IIl.; and
have found the larve folded in leaves of the yellow locust, Robinia,
upon which they had evidently been feeding.
SPHINGIDZ
Cressonia juglandis Sm. and Abb.
This caterpillar was taken on low branches of the shell-bark hick-
Ban Carya ovata, in the upland forest (Sta. IV,a) Aug. 20 (No.
102).
The larva feeds upon walnut, ironwood, and hickory. Our speci-
men bore a large number of cocoons of a hymenopterous parasite.
When handled, this larva makes a peculiar squeaking sound (Bull.
54, Bur. Ent., U. S. Dept. Agr., p. 80. 1905).
227
SaTURNIIDA
Telea polyphemus Cramer. American Silkworm. (PI. LIX, fig. 6.)
This caterpillar was taken on the ground, under hickories and
white oaks on the forested slopes to the valley (Sta. IV, b) Aug. 26
(No. 163).
The larva feeds upon walnut, basswood, elm, maple, cherry, etc.
CERATOCAMPIDA
Citheronia regalis Fabr. Royal Walnut Moth; Hickory Horned-devil
(larval name). (PI. LX, figs. 1 and 2.)
This larva was found on the valley slope (Sta. IV, b) on sumac
Aug. 16 (No. 68); and on walnut Aug. 20 (No. 108). This last
specimen was apparently fully mature.
The food plants of the larva are butternut, hickory, sycamore,
ash, and lilac. See Packard (’05, p. 130) for many figures and a
full description of this species.
Bastlona imperialis Drury. Imperial Moth. (Pl. LXI, Fig. 1).
The larva of this species was found on the leaves of sassafras on
the forested slope to the lowland forest (Sta. IV, b) Aug. 20 (No.
106). It feeds upon a large number of forest trees including oak,
maple, wild cherry, walnut, hickory, and several conifers.
See Packard (’05, p. 125) for figures and full descriptions of
this species.
ARCTIDA
Halisidota tessellaris Sm. and Abb. (Pl. LXI, fig. 4.)
These caterpillars were taken on hickory on the wooded slope to
the lowland (Sta. IV, b) Aug. 26 (No. 163); and, again, abundantly
(No. 168), in the upland forest (Sta. IV, a) on climbing buckwheat,
Polygonum convolvulus, which was entwined about a young walnut
or butternut. The yellow hairs and the tufts give this caterpillar a
striking appearance.
I have found moths of this species abundant at Bloomington, IIl.
The food plants are recorded as maple, oak, hazel, and button-
wood. Though larve were abundant upon leaves of the climbing
buckwheat, I did not observe them there eating it.
Noctua
Autographa precationis Guen.
The moth was taken in the open glade in the lowland forest (Sta.
IV, c) Aug. 22 (No. 143).
The larva feeds upon plantain, burdock, and dandelion.
228
Scolecocampa liburna Geyer. Rotten-log Caterpillar.
A single caterpillar (No. 125) was taken Aug. 22 upon the slope
of a wooded ravine (Sta. IV, b) under the bark of a stump in an
early stage of decay—the sap-wood honeycombed, but the remainder
solid though discolored. The larva, with its characteristic excrement,
was found in a cell excavated in the rotten sap-wood.
This is another species of animal which invades wood in the sap-
wood stage of decay and is so often associated with Philomycus
carolinensis, Passalus cornutus, and newly established colonies of
Camponotus herculeanus pennsylvanicus. The larva winters in logs,
as is evidenced by the fact that I found it in such situations late in fall
and early in spring (March 23) at Urbana, Ill. The large quantity
of excrement often indicates the approximate location of the larva.
This larva has been described by Edwards and Elliot (Papilio, Vol.
3, p. 134. 1883). It has been found in chestnut, oak, and other kinds
of decaying logs. The moth is recorded in July. The pileated wood-
pecker, Phleotomus pileatus, has been known to eat this caterpillar
(Beal, in Bull. 37, Biol. Surv., U. S. Dept. Agr., p. 34. 1911). Smith
(Ann. Rep. N. Jersey State Mus., 1909, p. 471. 1910) states that the
larva is found in “decaying cherry, hickory, oak and chestnut
stumps.”
i
NoToponTIDz
Datana angusti G. and R.
The caterpillar of this species was found on the valley slope (Sta.
IV. b) on bitternut hickory, Carya microcarpa, Aug. 20 (No. 104);
in the upland forest (Sta. IV, a) on hickory Aug. 16 (No. 65); and
at the margin of this forest Aug. 26 (No. 162).
The food plants of the larva are walnut, hickory, linden, and birch.
Packard (’95, pp. 110-111) describes and gives figures of the larva
and adult.
Nadata gibbosa Sm. and Abb. (Pl. LXI, fig. 2.)
This larva was taken on white oak, Quercus alba, in a forested
ravine (Sta. IV, b) Aug. 19 (No. 94); on leaves of the white oak,
upon which it had been feeding, in the upland forest (Sta. IV, @)
Aug. 26 (No. 169).
Packard (’95, pp. 142-146) gives figures of this species and
lists as food plants, oak, birch, and sugar plum. It is also reported
on maple.
Heterocampa guttivitta Walk (?). (Pl. LXI, figs. 3 and 5.)
This larva (No. 127) was captured Aug. 22 by a digger-
wasp, Ammophila abbreviata Fabr. which was found dragging it
along the ground in the upland forest (Sta. IV,a). See Packard
229
(95, pp. 230-235) for an account of this forest-inhabiting larva.
The larva of guttivitta is known to feed upon red maple, oak, and
viburnum.
GEOMETRDZ
Eustroma diversilineata Hiibn. (P1. LXII, fig. 1.)
This span-worm moth was taken in the upland forest (Sta. IV, a)
Aug 26 (No. 163).
Packard (Monogr. Geometrid Moths, p. 128. 1876) states that
the larva feeds upon grape and Psedera. ‘These are mainly forest
plants, and this is probably a woodland species.
Caberodes confusaria Hiibn.
This moth was taken near the upper slope of the south ravine in
open woods (Sta. IV, b) Aug. 22 (No. 161).
The larva feeds upon Trifolium.
CocHLDnDZzZ
Cochlidion or Lithacodes sp. Slug Caterpillar.
This curious larva was found on a stump on the wooded ravine
slope (Sta. IV, b) Aug. 26 (No. 165).
GELECHIDA
Vpsolophus ligulellus Hiibn. (?)
These small moths were taken in the upland woods (Sta. IV, a)
by T. L. Hankinson June 28, 1911 (No. 7678). The larva is reported
on apple, pear, and plum.
DIPTERA
CECIDOMYIDA
Cecidomyia holotricha O. S. (Hairy Midge-gall.)
Abundant on the under side of hickory leaves (near Sta. IV)
Aug. 20 (No. 96); and on leaves of Carya ovata in the upland for-
est (Sta. IV,a) Aug. 26 (Nos. 107 and 170). These brownish,
hairy galls may cover large areas on the under side of some leaves.
See Cook ’05, p. 840, or Beutenmiiller ’04, p. 172.’
Cecidomyia tubicola O. S. (Hickory Tube-gall.)
Immature galls (No. 107) were found Aug. 20 in the upland
Bates woods (Sta. IV, a) on the lower side of leaves of Carya ovata.
230
Cecidomyta caryecola O. S. (Hickory Seed-gall.)
This gall was taken on Carya ovata leaves in the upland forest
(Sta. IV,a) Aug. 20 (No. 107); and Aug. 26 (No. 170). Many
galls are formed on hickory and other trees by plant-lice (Cf. Per-
gande, 02).
ASILIDE
Deromyia discolor Loew.
This robber-fly was taken in an open area in the lowland forest
(Sta. IV,c) Aug. 20 (No. 117). Williston (Kingsley’s Standard
Natural History, Vol. 2, pp. 418-419. 1884) states that most robber-
flies “rest upon the ground, and fly up when disturbed, with a quick
buzzing sound only to alight again a short distance ahead. All their
food, which consists wholly of other insects, is caught upon the
wing . . . . Other flies and Hymenoptera are usually their food,
but flying beetles, especially Cicindelide, are often caught, and they
have even been known to seize and carry off large dragonflies. Not
only will they feed upon other Asilide, but the female frequently
resents the caresses of her mate by eating him up, especially if he is
foolish enough to put himself in her power. In an instance the
writer observed, a female seized a pair of her own species, and thrust-
ing her proboscis into the thorax of the male, carried them both off
together. . . . . The larve live chiefly under ground or in rotten
wood, especially in places infested with grubs of beetles upon which
they will feed. The young larve will bore their way completely
within beetle larvze and remain enclosed until they have consumed
them. Many, however, are found where they evidently feed upon
rootlets or other vegetable substances. They undergo their trans-
formations in the ground. The pupz have the head provided with
tubercles, and on the abdominal segments there are also spiny pro-
tuberances and transverse rows of bristles, which aid the insects to
reach the surface when they are ready to escape as flies.” Mar-
latt (Proc. Ent. Soc. Wash., Vol. 2, p. 82. 1893) observed D. dis-
color preying upon wasps of the genus Vespa. By seizing the head
of the wasp it avoids being stung.
Deromyta umbrinus Loew.
A specimen of this large robber-fly was taken in the south ravine
(Sta. IV, d) by T. L. Hankinson, with the eucerid bee Melissodes
perplexa Cresson in its grip, Aug. 22, 1910 (No. 7530).
231
SyrpHpz
Chrysotoxum ventricosum Loew.
This wasp-like fly was found resting on a leaf in the upland for-
est (Sta. IV, a) Aug. 26 (No. 163).
Mesogramma politum Say. Corn Syrphid.
This fly was taken by T. L. Hankinson in the Bates woods (Sta.
IV) June 28, 1911 (No. 7678). See the prairie list, p. 188.
Milesia ornata Fabr. Vespa-like Syrphid.
This beautiful large syrphid was taken on dogbane in an open
space in the upland forest (Sta. IV,a) Aug. 20 (No. 103); in the
open glade in the lowland forest (Sta. IV,c) Aug. 22 (No. 143);
and on Aug. 26 (No. 184) on the flowers of Eupatorium celestinum
in the clutches of the flower spider Misumena aleatoria Hentz. It
was also taken in the Bates woods by T. L. Hankinson June 28, 1911
(No. 7678). Metcalf (’13, p. 73) quotes Verrall as follows con-
cerning the subfamily Milestine: “What little is known about the
metamorphism shows that many species live in rotten wood or about
the sap flowing from injured tree trunks.”
HYMENOPTERA
Sirica
Tremex columba Linn. Horntail; Pigeon Tremex.
This species was taken on wing in the upland forest (Sta. IV, a)
Aug. 16 (No. 66); and on the open slope of a ravine (Sta. IV. b)
Aug. 22 (No. 132).
The larva bores in the trunks of trees, as oak, elm, sycamore, and
maple. Consult Packard (’90, pp. 379-381) for a description and
figure of the larva. The long-sting, Thalessa lunator, is an external
parasite upon this larva (see Riley, 88). I have taken normally
colored females at Bloomington, Ill, July 25, Sept. 29, and Oct. 8.
Two abnormally colored individuals were taken in September, one
of them almost, and the other (taken Sept. 29) completely lacking
the usual black markings. A female was taken at Milmine, III, in
October. Consult Bradley (’13) for a key to the varieties of this
species of Tremex.
An interesting feature in the ecological relations of this species
is the fact that it appears to frequent only weakened, diseased, and
dying trees, and these, not as a primary invader, but as a trailer,
following insects which have done previous injury to the trees.
Felt (’05, p. 61) shows that in New York successive attacks of the
232
elm leaf-beetle, or injury by the sugar maple borer Plagionotus
spectosus Say, prepare the way for the horntail larva. Ecologically
considered, the leaf-beetle and the borer initiate a succession of in-
sect invasions into the tree trunk; Tremex follows, with its parasite
Thalessa; and these in turn lead the way for still others; thus a suc-
cession of insects is produced.
CynrrPDz
Holcaspis globulus Fitch. (Oak Bullet Gall.)
This gall was taken on white oak, Quercus alba, in the upland
forest (Sta. IV, a) Aug. 26 (No. 170).
Consult Cook (’05) and Beutenmiller (’04) for figures and de-
scriptions of various kinds of galls mentioned in this list.
Amphibolips confluens Harr. (Oak-apple or May-apple Gall. )
These galls were abundant upon the forest floor in the upland
Bates woods (Sta. IV, a) during August (No. 101). The galls grow
upon the leaves of several species of oaks (Quercus).
Amphibolips prunus Walsh. (Acorn Plum Gall.) (Pl. LXII, fig. 2.)
A single specimen of this gall was found on the slope of the south
ravine in Bates woods (Sta. IV,b) Aug. 22 (No. 131). Another
specimen came from the woods northeast of the Bates woods Aug.
20 (No. 96). It grows upon acorn cups.
Andricus clavula Bass. (White Oak Club Gall.) (PI. LXTI, fig. 5.)
This gall, formed in the terminal bud, was common on white oak,
Quercus alba, in the upland Bates woods (Sta. IV, a) Aug. 26 (No.
170).
Andricus cornigerus O. S. (Horned Knot Oak Gall.) (PI. LXII,
fig. 3.
This gall occurred in very large numbers on the branches of
shingle oak, Quercus imbricaria, in a forest just northeast of the
Bates woods, Aug. 20 (No. 96). The galls are old and apparently
decaying.
Andricus lana Fitch. (Oak Wool Gall.) (Pl. LXII, fig. 4.)
Two examples of this gall were found on leaves of white oak,
Quercus alba: one was taken near the Bates woods (Sta. IV) Aug.
20 (No. 96), and the other was found in the Bates woods (Sta. IV, a)
on the petiole of a leaf, Aug. 26 (No. 170).
Andricus seminator Harr. (Oak Seed-gall.) (Pl. LXIII, fig. 1.)
A single specimen of this gall was found upon Quercus. alba
(Sta. IV,a) Aug. 20 (No. 101). The cotton-like masses of this
233
gall are conspicuous. They may be tinged with red; when dry they
become brownish.
IcHNEUMONIDA
Thalessa lunator Fabr. Lunate Long-sting.
A female ichneumon of this species was found on a tree trunk in
the open glade in the lowland forest (Sta. IV, c) Aug. 22 (No. 143).
The larva feeds, as an external parasite, upon, the larva of the
horntail, Tremex columba, which was also found in the Bates woods
(Sta. IV). I found T. lunator, both males and females, abundant
on shade trees at Bloomington, Ill., October 1, 1892, and also took it
' July 26, 1895. Riley (’88) gives an excellent account of this species
accompanied by figures of the immature stages, and that of its host
as well.
Trogus obsidianator Brullé.
This black ichneumon with fulvous antenne was taken in the
Bates woods (Sta. IV) June 28, 1911, by T. L. Hankinson (No.
7678). This wasp is known to be parasitic upon the larva of Papilio
polyxenes Fabr. (P. asterias—Insect Life, Vol. 1, p. 161) and
upon the caterpillar of Pyrrharctia isabella (?). This species has been
taken in central Illinois during June and July (Weed, Psyche, Vol. 5, p.
52). (See also Riley, in Amer. Ent., Vol. 3, p. 134. 1880.)
PELECINIDA
Pelecinus polyturator Drury. Black Longtail. (Pl. LXIII, fig. 2.)
This remarkable looking insect was found in the glade of the
lowland forest (Sta. IV,c) Aug. 20 (No. 117) and Aug. 22 (No.
143). Other females were seen in this forest.
I have also’ taken this species at Bloomington, Ill. At Evanston,
Ill., during July, 1910, this species was very abundant upon some
damp lawns. I have counted four or five females in sight at once.
They were often found upon blue-grass sod. The male of this
species is considered very rare. The only one which I ever captured
was taken July 29, 1910, at Evanston, Ill. The larva is parasitic
upon the grub of the May-beetle, Lachnosterna (Forbes, Eighteenth
Rep. State Ent. Ill, p. 124. 1894). It may also prey upon other
scarabzeid larve inhabiting woodlands.
Formicipz
Stigmatomma pallipes Hald. Old-fashioned Ant.
A single wingless queen and four pupe (No. 140) were taken
Aug. 22 near the base of a ravine slope (Sta. IV, >) in dense shaded
234
woods, almost devoid of herbaceous vegetation, but with a thick layer
of leaves, and other vegetable debris.
Wheeler (Biol. Bull., Vol. 2, pp. 56-69. 1901) considers this a
rather rare ant, although widely distributed over eastern North
America. It is subterranean in habit, and ‘‘does not come to the
surface even at night.” Contrary to the habits of must ants this
primitive species has retained the carnivorous habits of the ancestral
forms, and the young are fed on fragments of insects. They do not
feed one another, or the larve by regurgitation, as do the specialized
species of ants. They thus furnish us a glimpse at the ancient his-
tory of ants. Wheeler (’05, pp 374-375) states that this species oc-
curs only in “rich, rather damp woods, under stones, leaf mould,
or more rarely under or in rotten logs.”
A worker of Myrmica rubra Linn., subsp. scabrinodis Nyl., var.
schencki Emery (No. 140) was taken from the same patch of leaves.
Cremastogaster lineolata Say. (Pl. LXII, fig. 6.)
This ant was taken only once—in the upland part of the Bates
woods (Sta. IV, a) Aug. 20 (No. 118). Large numbers of the ants
were found in an oak-apple gall (Amphibolips confluens Harr.)
lying on the forest floor. When I picked up the gall, many ants
came out and ran over my hand, biting vigorously.
This is essentially a ground and forest-inhabiting ant, which
forms nests in the soil, under stones, and in logs, stumps, etc. It
has the peculiar instinct to make a sort of temporary nest out of
debris to cover the aphids and coccids which it attends (Wheeler,
Bull. Am. Mus, Nat. Hist., Vol. 22, pp. 1-18. 1906).
Several carnivorous staphylinid beetles of the genus M. yrmedonia
have been taken in the nests of these ants (Wheeler, ’10a, p. 382;
Schwarz, ’gob, p. 247).
Aphenogaster fulva Roger.
A well-rotted stump in the upland Bates woods (Sta. IV, a) was
found Aug. 17 to contain a moist, felt-like layer of some fungous
growth, and on this was a large colony of snails (No. 71). In an
adjacent part of this stump was a small colony of white ants, Termes
flavipes Koll. (No. 72). A colony of ants which was in close prox-
imity to the white ants, proved to be A. fulva Roger. As the gal-
leries were exposed by cutting up the stump, these ants were seen to
pick up the termites and carry them away, just as they do their own
young on similar occasions. Five pairs—the ant and the termite
which it carried—were preserved (Nos. 74-76, and 78-79). One
of the termites lacks a head. All of them were workers. Larve
and naked pupe (No. 79) were abundant in this nest, and workers
(No. 80) were abundant about the stump. On Aug. 22 another
235.
colony of this ant (No. 125) was found under the bark of a decaying
oak stump (Sta. IV) in which the sap-wood was honeycombed, but
the remainder solid, though discolored.
Forel (Psyche, Vol. 9, p. 237. 1901) remarks that Aphenogas-
ter is “very fond of termites, and when one uncovers and scatters
about a nest of termites in a wood, they hasten to feast on the suc-
culent morsels.” These observations suggest the possible fate of
the captured termites; none of the ants were seen to eat them, how-
ever. In the absence of observations, the missing head mentioned
above may be variously accounted for.
This habit of carrying off termites has been observed in other
species of ants. Forbes (19th Rep. State Ent. Ill., p. 198. 1896) re-
ports that near Carterville, Mason county, Ill, Mr. John Marten
observed Formica schaufussi (=Formica pallide-fulva Linn., subsp.
schaufussi Mayr) to pick up and carry away the living termites
when its nest under a log in which termites abounded, was disturbed,
and McCook (Proc. Acad. Nat. Sci. Phila., 1879, p. 155) has ob-
served similar behavior in the case of the mound-building ant, For-
mica exsectoides Forel.
The histerid beetle Heterius blanchardi Schwarz has been found
in nests of this ant (Wheeler, ’10a, pp. 388, 389) ; and European ob-
servers have seen ants carrying and rolling them about. Consult
also Schwarz (’9ob, 247) for a list of beetles found with this ant.
Wheeler (’10a, p. 206) lists A. fulva as a glade species which in
the forests utilizes logs and branches as‘substitutes for stones. (See
Wheeler, ’05, pp. 372-373.)
Aphenogaster tennesseensis Mayr. Tennessee Ant.
A colony of this ant (No. 87) was taken Aug. 17 from a decaying
stump, situated on the slope (Sta. IV, >) from the upland forest to
the lowland on the river bottom.
According to Wheeler (Bull. Am. Mus. Nat. Hist., Vol. 20, 1904,
p. 362, and Vol. 21, 1905, p. 373) this species normally nests in dead
wood in rather open forests. He holds the opinion that the queen of
this species can not rear her own brood, and thus establish a new
colony, but must utilize a small or weak colony of the allied species
A. fulva Roger, which lives under stones. Thus the new colonies are
started under stones; later, when they become numerous, they are
found in rotten wood. This, Wheeler concludes, indicates that they
“migrate away from the fulva workers.” Tanquary (11) has per-
formed some interesting experiments which show that queens of
tennesseensis are adopted by colonies of other ants, a result which
seems to confirm Wheeler’s anticipation.
Schwarz (’9ob, p. 247) records two beetles found with this ant.
236
Formica fusca Linn., var. subsericea Say.
This ant was taken in the upland Bates woods (Sta. IV, a) Aug.
26 (No. 163). See the list of prairie invertebrates, p. 190.
Myrmica rubra Linn., subsp. scabrinodis Nyl., var. schencki Emery.
This ant (No. 140) was found Aug. 22 under leaves in a small
ravine on a shady slope (Sta. IV. b) from the upland forest to the
valley bottoms. The soil under these leaves had been thoroughly tun-
neled by small mammals during the preceding winter, but recently the
leaves had not been disturbed. The soil was a mixture of sand, clay,
and vegetable debris, was moist, and contained few kinds of animals.
A single ant of this variety (No. 140) was taken while collecting spec-
imens of Stigmatomma pallipes.
This species is listed by Wheeler (Bull. Am. Mus. Nat. Hist., Vol.
21, p. 373. 1905) as a field ant which prefers to nest in grassy pas-
tures and lawns, in situations exposed to the sun. Our specimen
was, therefore, found in an unusual habitat.
Tapinoma sessile Say. Cocoanut Ant.
This cocoanut ant, so called because of the odor of the workers,
which has been compared to that of decayed cocoanuts, was found
in the lowland part of the Bates woods, at the base of the slope to
the bottoms (Sta. IV,c) Aug. 22 (No. 139). A large colony was
found among the surface layers of dry dead leaves; from it were se-
cured two queens, vast numbers of eggs, and also larve, pupa, and
workers.. Wheeler (’05, pp. 373, 389) states that this ant usually
nests in open sunny woods, the borders of woods, and under stones,
logs, etc.
Schwarz (’9ob, p. 247) records beetles as living with this ant.
Camponotus herculeanus Linn., subsp. pennsylvanicus DeG. Carpen-
ter Ant.
This species was taken from under the bark of a rotting: stump
among a dense second-growth, on the valley slope (Sta. IV, b) be-
tween the upland and the lowland forest Aug. 17 (No. 84). This
stump was in that stage of decay so often utilized by the large Caro-
lina slug, Philomycus carolinensis, and the horned Passalus beetle,
Passalus cornutus. The colony was recently founded, for the dea-
lated female occupied a small cell excavated in the rotten sap-wood.
This colony consisted of four pupz and six larve of different sizes.
Another colony was taken in the same stump, from the rotted sap-
wood zone, in company with the snail Philomycus carolinensis and
some kind of pulmonate snail eggs. This colony was in a more ad-
vanced stage than the preceding, about a dozen larve, seven pup,
237
and two adult workers being present, and about half a dozen eggs
(No. 85).
Pricer (’08) has given an interesting account of the life history
and habits of this ant in Illinois. He states (p. 197) that the food
is largely the honeydew of plant-lice, but is supplemented by plant
juices and dead insects. He found a small staphylinid beetle, Xeno-
dusa cava, abundant in the nests.
I have found pennsylvanicus abundant at Bloomington, Ill., and
represented as follows: by a male June 29; by a winged female in
June; and by dealated females June 29 and July 2 and 25.
McCook (’83) has given an interesting account of the found-
ing of colonies of this ant. See also Wheeler, ’o6b, pp. 38-39, Plate
VII, and ’1ob, pp. 335-338, for further information concerning it.
Camponotus herculeanus Linn., subsp. pennsylvanicus DeG., var.
ferrugineus Fabr.
This variety was taken a short distance to the northeast of the
Bates woods (Sta. FV) Aug. 20 (No. 97). Here the large ground-
beetle Calosoma scrutator was found running on the ground with
what appeared to be a bunch of greenish moss; a large reddish ant
also struggled for possession of the prize. Upon closer examination
it was found that the skin of some large lepidopterous larva was
the object desired. This skin, recently shed or moistened by a recent
rain, was a prize for both ferrugineus and Calosoma.
A dead wingless ferrugineus, covered with a fungus growth, was
found in a small cell excavated in the rotten wood of a decaying log
on the ravine slope (Sta. IV, b) Aug. 17 (No. 90). Apparently this
female had died before her colony developed. (See Pricer, ’08;
Wheeler ’10b, pp. 338-339.)
I have found this form abundant at Bloomington, Ill. Winged
females were taken July 26, dealated ones on July 25 and 26, and
males June 29, and July 9 and 25. On July 21, 1892, several males
were taken at night, being attracted to a lamp located near a small
brook.
A very large colony, numbering thousands of individuals, was
found May 26, under a well-decayed log, in a forest at White Heath,
Ill. It contained winged males, females, and workers. The winged
forms were present in vast numbers. The far-advanced condition of
decay of the log was in marked contrast with that in which the initial
colonies are usually found. During the years of development of
such a large colony the progress of decay will naturally make some
changes in the habitat; reciprocally the ants will doubtless tend to
monopolize the logs to the exclusion of some other animals, and
238
also facilitate the decay of the log by their activities. There is an
“orderly sequence” of changes in the developing colony, and a simi-
lar orderly sequence of changes in the log habitat.
An ant colony in its development clearly illustrates the transfor-
mation from the individual to the associational phase of ecological
relations. Beginning with the fertilized female and her progeny,
the colony develops in size and in the division of labor among its
members; until, finally, by the possible addition of slaves, commen-
sals, parasites, and even predaceous enemies, the colony or associa-
tion is built up in an orderly sequence, and the organisms adjust
themselves to one another and to the environment in general.
MuTILLDA
Spherophthalma sp. Velvet Ant.
This stinging, wingless velvet ant was taken at the margin of the
forest near the cleared area (Sta. IV, a) Aug. 23 (No. 151).
PSAMMOCHARIDAG
Psammochares ethiops Cress. (Pompilus Fabr.)
This large black wasp was taken by T. L. Hankinson July 10,
Ig1I, in the Bates woods (No. 7693). It probably stores its nest
with spiders.
SPHECID
Ammophila abbreviata Fabr. Short Caterpillar Wasp.
This wasp was taken on the open ravine slope (Sta. IV, b) Aug.
22 (No. 124). One example (No. 127) was running on the ground
in the upland forest (Sta. IV,@) with a quiescent bombycine cater-
pillar—probably Heterocampa guitivitta Walk.—in its grip.
I took this species of wasp at Bloomington, III, July 26. Its
copulating habits have been recorded, with figures, by Turner (’02).
BIBLIOGRAPHY
Abbott, J. F.
"12, An unusual symbiotic relation between a water bug and a
crayfish. Am. Nat., 46: 553-556.
Adams, C. C.
’o1. Baseleveling and its faunal significance, with illustrations
from southeastern United States. Am., Nat., 35: 839-852.
‘og. The Coleoptera of Isle Royale, Lake Superior, and their
relation to the North American centers of dispersal. Ann.
Rep. Mich. Geol. Surv., 1908, pp. 157-215.
13. Guide to the study of animal ecology. VII+183 pp. New
York.
715. An outline of the relations of animals to their inland envi-
ronments. Bull. Ill. State Lab. Nat. Hist. 11: 1-32.
Aldrich, J. M.
’7o5. A catalogue of North American Diptera. Smithsonian
Miscel. Coll., 46: No. 1444. 680 pp.
Ashmead, W. H.
’92. Notes on the genus Melittobia. Proc. Ent. Soc. Wash.,
2: 228-232.
Babcock, S. M.
"12. Metabolic water ; its production and réle in vital phenomena.
Research Bull. Wis. Agr. Exper. Sta., No. 22: 87-181.
Baker, F. C.
’o2. The Mollusca of the Chicago area. Part II. The Gastrop-
oda. Bull. Chicago Acad. Sci., No. 3, Part 11: 137-410.
11. The Lymnzidz of North and Middle America, recent and
fossil. Special Pub. Chicago Acad. Sci., No. 3. 539 pp.
Baker, H. B.
711. Mollusca. Ann. Rep. Bd. Mich. Geol. and Biol. Surv..
IQIO, pp. 121-176.
’
240
Banks, N.
04. A treatise on the Acarina, or mites. Proc. U. S. Nat. Mus.,
28:1-114. (No. 1382) ;
’o7. A “census of four square feet.” Science, n. s., 26: 637.
Banta, A. M.
‘07. The fauna of Mayfield’s Cave. Carnegie Inst. Pub. No.
67. I14 pp.
Barrows, H. H.
‘10. Geography of the middle Illinois valley. Bull. Ill. Geol.
Surv., No. 15. 128 pp.
Bates, C. G.
"11. Windbreaks: their influence and value. U. S. Dept. Agr.,
Forest Serv., Bull. 86. 100 pp.
Beutenmiiller, W.
796. Food-habits of North American Cerambycide. Journ.
N. Y. Ent. Soc., 4: 73-81.
04. The insect-galls of the vicinity of New York City. Am.
Mus. Journ., 4: 89-124.
‘oga. ‘The species of Holcaspis and their galls. Bull. Am. Mus.
Nat. Hist., 26: 29-45.
*ogb. The species of Amphibolips and their galls. Bull. Am.
Mus. Nat. Hist., 26: 47-66,
Binney, W. G.
85. A manual of American land shells. Bull. U. S. Nat. Mus.,
No. 28. 528 pp.
Blatchley, W. S.
03. The Orthoptera of Indiana. 26th Ann. Rep. [Ind.] Dept.
Geol. and Nat. Resources, 1901, pp. 123-471.
10. An illustrated descriptive catalogue of the Coleoptera or
beetles (exclusive of the Rhynchophora) known to occur in
Indiana. Bull. No. 1, [Ind.] Dept. Geol. and Nat. Resources.
Also published by Nature Pub. Co., Indianapolis. 1386 pp.
?
Bouyoucos, G. J. ae
713. An investigation of soil temperature and some of the most
important factors influencing it. Mich. Agr. Exper. Sta.,
Tech. Bull. No. 17, pp. 1-196.
241
Bradley, J. C.
13. The Siricide of North America. Journ. Ent. and Zool.,
5: I-30.
Brauer, F.
83. [Habits of i imago of Bittacus tipularius F.] Verhandl. zool. -
bot. Vereins in Wien, 3:.151.
55. Beitrage zur Kenntniss des inneren es und der Ver-
wandlung der Neuropteren. Verhandl. der zool.-bot. Vereins
in Wien, 5: 701-720.
63. Beitrage zur Kenntniss der Panorpiden-Larven. Verhandl.
k. k. zool.-bot. Gesellsch., 13 : 307-324.
"71. Beitrage zur Kenntniss der Lebensweise und Verwandlung
der Neuropteren. (Micromus variegatus Fabr., Panorpa
communis L., Bittacus italicus Klg. und Bittacus Hagenii
Brau.) Verhandl. k. k. zool.-bot. Gesellsch. in Wien, 21:
107-166.
Briggs, L. J., and Belz, J. O.
*11._ Dry farming in relation to rainfall and evaporation. U. S.
Dept. Agr., Bur. Plant Industry, Bull. No. 188. 71 pp.
Buckingham, E.
04. Contributions to our knowledge of the aeration of soils.
U.S. Dept. Agr., Bur. Soils, Bull. No. 25. 52 pp.
Bunge, G.
88. ' Ueber das Sauerstoffbediirfniss der Schlammbewohner.
Zeit. fiir Physiol. Chemie, 12: 565-567.
Busck, A.
’02. Notes on enemies of mushrooms and on experiments with
remedies. U.S. Dept. Agr., Div. Ent., Bull. No. 38, n. s., pp.
32-35-
Cameron, A. E.
13. General survey of the insect fauna of the soil within a lim-.
ited area near Manchester [England]:a consideration of the
relationships between soil insects and the physical conditions
of their habitat. Journ. Econ. Biol., 8: 159-204.
Card, F. W.
‘97. Windbreaks. Bull. Neb. Agr. Exper. Sta., 9, No. 48:
69-96.
242
Caudell, A. N.
‘or. The genus Sinea of Amyot and Serville. Journ. N. Y.
Ent. Soc., 9: I-11.
Chittenden, F. H.
’92. Notes on the food habits of some species of Chrysomelide.
Proc. Ent. Soc. Wash., 2: 261-267.
’98. Twig pruners and allied species. U. S. Dept. Agr., Div.
Ent., Bull. No. 18, n. s., pp. 35-43.
02. The leaf-mining locust beetle, with notes on related species.
U. S. Dept. Agr., Div. Ent., Bull. No. 38, n. s., pp. 70-89.
704. Insects injurious to basket willow. U.S. Dept. Agr., Bur.
Forestry, Bull. No. 46: 63-80.
‘06. Harvest mites, or “chiggers.” Circ. No. 77, Bur. Ent.,
U.S. Dept. Agr. 6 pp.
08. The nut weevils. Circ. No. 99, Bur. Ent., U. S. Dept. Agr.
15 pp.
710, The oak pruner. Circ. No. 130, Bur. Ent., U. S. Dept.
Agr. 7 pp. ;
Cockerell, T. D. A.
’97. The food plants of scale insects (Coccide). Proc. U. S.
Nat. Mus., 19: 725-785. (No. 1122)
Coffey, G. N.
712. A study of the soils of the United States. U. S. Dept.
Agr., Bur. Soils, Bull. No. 85. 114 pp.
Colton, H. S.
’08. Some effects of environment’ on the growth of Lymnza
columella Say. Proc. Acad. Nat. Sci. Phil., 1908, pp. 410-
448.
Cook, M. T.
’os. The insect galls of Indiana. Twenty-ninth Ann. Rep. °
[Ind.] Dept. Geol. and Nat. Resources, 1904, pp. 801-867.
Cook, O. F.
’t1a. Notes on the distribution of millipeds in southern Texas,
with descriptions of new genera and species from Texas,
Arizona, Mexico and Costa Rica. Proc. U. S. Nat. Mus.,
40:147-167. (No. 1810)
3
243
*11b. New tropical millipeds of the order Merocheta, with an
example of kinetic evolution. Proc. U. S. Nat. Mus., 40:
451-473. (No. 1831)
*11c. The hothouse milliped as a new genus. Proc. U. S. Nat.
Mus., 40: 625-631. (No. 1842)
Coquillett, D. W.
’97. Revision of the Tachinide of America north of Mexico.
U.S. Dept. Agr., Div. Ent., Tech. Ser. No. 7. 154 pp.
Coville, F. V.
’14. The formation of leafmold. Smithsonian Report for 1913,
PP. 333-343.
Cowles, H. C.
"11. The causes of vegetative cycles. Bot. Gaz., 51: 161-183.
Cox, H. J.
A
10. Forest and temperature conditions in the cranberry marshes
of Wisconsin. U. S. Dept. Agr., Weather Bur., Bull. T.
I2I pp.
Crosby, C. R., and Leonard, M. D.
"14. The tarnished plant-bug. Cornell Agr. Exper. Sta., Bull.
No. 346: 463-526. ,
Crumley, J. J.
"10. The relative durability of post timbers. Bull. Ohio Agr.
Exper. Sta., No. 219: 605-640.
Currie, R. P.
’o5. Catalogue of the exhibit of economic entomology at the
Lewis and Clark Centennial Exposition, Portland, Oregon,
1905. U.S. Dept. Agr., Bur. Ent., Bull. No. 53. 127 pp.
Dachnowski, A.
"11. The vegetation of Cranberry Island (Ohio) and its rela-
tions to the substratum, temperature, and evaporation. Parts
I and II. Bot. Gaz., 52: 1-33; 126-150.
712. Peat deposits of Ohio: their origin, formation and uses.
Bull. Geol. Surv. Ohio, ser. 4, No. 16. 424 pp. Columbus,
O.
Davenport, C. B.
‘97. Experimental morphology. Part I. Effect of chemical
and physical agents upon protoplasm. 280 pp. New York.
244
‘03. The Collembola of Cold Spring beach with special reference’
to the movements of the Poduride. Cold Spring Harbor
Monogr., IT. 32 pp.
Davis, J. J.
"12. Williams’ ‘““The Aphidide of Nebraska”; a critical review.
Univ. Studies, Univ. Nebr., 11: 253-290.
"13. The Cyrus Thomas collection of Aphidide, and a tabulation
of species mentioned and described in his publications. Bull.
Ill. State Lab. Nat. Hist., 10: 97—121.
Dendy, A.
95. The cryptozoic fauna of Australasia. [Pub.] Austral. Assn.
Adv. Sci., 1895. 21 pp.
Dickey, M. G.
’og. Evaporation in a bog habitat. Ohio Nat., 10: 17-35.
Diem, K.
’03. Untersuchungen tiber die Bodenfauna in den Alpen. Jahrb.
der St. Gallischen Naturwiss. Gesellsch. f. das Vereinsjahr
IQOI—1902, pp. 234-414.
Ellis, M. M.
713. A descriptive list of the cephaline Gregarines of the New
World. Trans. Am. Micr. Soc., 32: 259-296.
Emerton, J. H.
’o2. ‘The common spiders of the United States. 225 pp. Boston.
Enteman, M. M.
‘02. Some observations on the behavior of the social wasps.
Pop. Sci. Monthly, 61 : 339-351.
Ewing, H. E.
og. A systematic and biological study of the Acarina of Illinois.
Univ. Studies, Univ. Tll., 3: 359-472.
Fassig, O. L.
‘99. ‘Types of March weather in the United States. Am. Journ.
Sci. (4) 8: 320-340.
Felt, E. P.
94. On certain grass-eating insects. A synopsis of the species
of Crambus of the Ithaca fauna. Bull. Cornell Univ. Agr.
Exper. Sta., No. 64: 47-102.
245
7
95. a scorpion-flies. Tenth Rep. N. Y. State Ent., pp. 463-
480.
‘05, ’06. Insects affecting park and woodland trees. Mem. 8,
N. Y. State Mus.: Vol. I (’05), 459 pp.; Vol. II (’06),
877 pp.
11. Summary of food habits of American gall midges. Ann.
Ent. Soc. Amer., 4: 55-62.
Felt, E. P., and Joutel, L. H.
’04. Monograph of the genus Saperda. Bull. N. Y. State Mus.,
No. 74. 86 pp.
Fernald, C. H.
96. The Crambide of North America. 81 pp. Mass. Agr.
Coll.
Fernald, H. T.
06. The digger-wasps of North America and the West Indies
belonging to the subfamily Chlorionine. Proc. U. S. Nat.
Mus., 31: 291-423.
Fiske, W. F.
7or1. An.annotated catalogue of the butterflies of New Hampshire.
Tech. Bull. N. H. Agr. Exper. Sta., No. 1. 80 pp.
Fitch, A.
’56. First and second report on the noxious, beneficial and other
insects of the State of New York. 327 pp. Albany.
Fletcher, J.
‘91. Report of the entomologist and botanist. Reports on the
experimental farms of the Dominion of Canada for 1890, pp.
154-188.
Flint, W. P., and Sanders, G. E.
’12. Note on a parasite of white-grubs. Journ. Econ. Ent.
5: 490.
Folsom, J. W.
‘og. ‘The insect pests of clover and alfalfa. Bull. Ill. Agr. Exper.
Sta. No. 134:113-197; Twenty-fifth Rep. State Ent. Ill,
Pp. 41-124.
Forbes, S. A.
76, List of Illinois Crustacea. Bull. Ill. State Lab. Nat. Hist.,
1, No. 1: 1-25.
,
246
78. Breeding habits of Corixa. Am, Nat., 12: 820.
’80. Notes on insectivorous Coleoptera. Bull. Ill. State Lab.
Nat. Hist., Vol. I, No. 3, pp. 1534160; reprint (1903), pp.
167-176.
83a. The food relations of the Carabide and Coccinellide.
Bull. Ill. State Lab. Nat. Hist., Vol. I, No. 6, pp. 33-64; re-
print (1912), pp. 33-60.
’83b. The food relations of predaceous beetles. Twelfth Rep.
State Ent. Ill, 1882, pp. 105-122. Also in Trans. Ill. Dept.
Agr. for 1882, Vol. 20.
84. The wheat bulb-worm. Thirteenth Rep. State Ent. Il,
1883, pp. 13-29. Also in Trans. Ill. Dept. Agr. for 1883,
Vol. 21.
go. Sixteenth report of the state entomologist on the noxious
and beneficial insects of the State of Illinois. Also in Trans.
Ill. Dept. Agr. for 1888, Vol. 26, pp. 1-104.
’94. A monograph of insect injuries to Indian corn. Part I.
Eighteenth Rep. State Ent. Ill. XI+171 pp. Also in Trans.
Ill. Dept. Agr. for 1893, Vol. 31, pp.
02. The corn bill-bugs in Illinois. Bull. Ill. Agr. Exper. Sta.,
No. 79: 435-461. Also (1903) in 22d Rep. State Ent. IIL,
pp. 1-26.
04. The more important insect injuries to Indian corn. Bull.
Ill. Agr. Exper. Sta., No. 95 : 329-399.
7o5. A monograph of insect injuries to Indian corn. Part II.
Twenty-third Rep. State Ent. Ill. 273 pp.
ova. On the life history, habits, and economic relations of the
white-grubs and May-beetles (Lachnosterna). Bull. Ill. Agr.
Exper. Sta., No. 116: 447-480. Also (1908) in 24th Rep.
State Ent. Ill., pp. 135-168.
’o7b. On the local distribution of certain Il!inois fishes: an essay
in statistical ecology. Bull. Ill. State Lab. Nat. Hist., 7: 273-
303.
"11. Some important insects of Illinois shade trees and shrubs.
- Bull. Il. Agr. Exper. Sta., No. 151: 463-529; 26th Rep.
State Ent. Ill, pp. 1-61. .
712. What is the matter with the elms in Illinois? Bull. Ill.
Agr. Exper. Sta., No. 154: 1-22; 27th Rep. State Ent. Ill,
pp. 1-20.
?
7
,
247
Forbes, S. A., and Hart, C. A.
‘oo. The economic entomology of the sugar beet. Bull. Ill. Agr.
Exper. Sta., No. 60: 397-532; 21st Rep. State Ent. Ill, pp.
49-175.
Fox, H.
’14. Data on the Orthopteran faunistics of eastern Pennsylvania
and Southern New Jersey. Proc. Acad. Nat. Sci. Phila.,
1914. pp. 441-534.
Franklin, H. J.
"13-14. The Bombide of the New World. Parts I and II.
Trans. Am. Ent. Soc., 38: 177-486; 39 : 73-200.
Fuller, G. D.
"11. Evaporation and plant succession. Bot. Gaz., 52: 193-208.
*12a. Evaporation and plant succession on the sand dunes of
Lake Michigan. Trans. Ill. Acad. Sci., 4: 119-125.
*12b. Evaporation and the stratification of vegetation. Bot.
Gaz., 54: 424-426.
"14. Evaporation and soil moisture in relation to the succession
of plant associations. Bot. Gaz., 58: 193-234.
Fuller, G. D., Locke, J. R., and McNutt, W.
14. The stratification of atmospheric humidity in the forest.
Trans. Ill. Acad. Sci., 1913, 6: 100-102.
Fuller, M. L.
712. The New Madrid Earthquake. Bull. No. 494, U. S. Geol.
Surv. I19 pp.
Gibson, W. H.
7oo. My studio neighbours. 245 pp. New York.
Gleason, H. A., and Gates, F. C.
‘12. A comparison of the rates of evaporation in certain asso-
ciations in central Illinois. Bot. Gaz., 53: 478-491.
Graenicher, S.
’o5. Some observations on the life history and habits of parasitic
bees. Bull. Wis. Nat. Hist. Soc., 3: 153-167.
706. A contribution to our knowledge of the visual memory of
bees. Bull. Wis. Nat. Hist., Soc., 4: 135-142.
10. The bee-flies (Bombyliide) in their relations to flowers.
Bull. Wis. Nat. Hist. Soc., 8: gI-101.
,
248
Hamilton, J.
’85. Short notes on Coleoptera. Can. Ent., 17: 45-48.
’87. Elaphidion villosum Fab. Can. Ent. 19: 141-145.
‘90. Balaninus—its food habits. Can. Ent., 22: 1-8.
Hamm, A. H.
08. Observations on Empis livida, L. Ent. Mo. Mag. (2),
19: 181-184.
’oga. Observations on Empis ‘opaca, F. Ent. Mo. Mag. (2),
20: 132-134.
’ogb. Setulia grisea, Mg., and Cerceris arenaria, Linn., in the
New Forest. Ent. Mo. Mag. (2), 20: 273-274.
Hancock, J. L.
’o2. The Tettigide of North America. 176 pp. Chicago.
‘o5. The habits of the striped meadow cricket (CEcanthus fas-
ciatus Fitch). Am. Nat., 39: I-II.
711. Nature sketches in temperate America. 451 pp. Chicago.
Harrington, M. W.
93. Review of forest meteorological observations: a study pre-
liminary to the discussion of the relation of forests to climate.
U. S. Dept. Agr., Forestry Div., Bull. No. 7: 23-122.
Harris, J. A.
’03. An ecological catalogue of the crayfishes belonging to the
genus Cambarus. Kan. Univ. Sci. Bull., 2: 51-187.
Hart, C. A.
’06. Descriptive synopsis of insect collections for distribution to
Illinois high schools. IJ. Orthoptera. Pages 67-92. [Illinois
State Laboratory of Natural History, Urbana.
’07. Zoological studies in the sand regions of the Illinois and
Mississippi river valleys. Bull. Ill. State Lab. Nat. Hist.,
7: 195~267.
’11. Miscellaneous economic insects. ‘T'wenty-sixth Rep. State
Ent. Ill., pp. 668-98.
Harvey, E. M.
*14. Evaporation and soil moisture on the prairies of Illinois.
Trans. Ill. Acad. Sci., 1913, 6: 92-99.
Heindel, R. L,
’05. Ecology of the willow cone gall. Am. Nat., 39: 859-873.
249
Henderson, L,. J.
’13a. The fitness of the environment. An inquiry into the bio-
peut significance of the properties of matter. 317 pp. New
ork. ;
’13b. The regulation of neutrality in the animal body. Science,
N. S., 37: 389-395.
Henneguy, L. F.
704. Les Insectes: Morphologie—Reproduction, Embryogénie.
804 pp. Paris.
Herman, W. C.
"10. A new bird enemy. Journ. Cincinnati Soc. Nat. Hist.,
21: 80.
Hesselman, H.
’o4. Zur Kenntniss des Pflanzenlebens schwedischer Laubwiesen.
Beihefte zum Bot. Centralbl., Bd. 17, Jahrg. 1904, pp. 311-
460.
Hewitt, C. G.
*12, The large larch sawfly [Nematus erichsonit], with an ac-
count of its parasites, other natural enemies and means of con-
trol. Can. Dept. Agr., Div. Ent., Bull. No. 10, sec. revise. 42
PP.
Hine, J. S.
798. The North American species of the genus Bittacus. Journ.
Columbus Hort. Soc., XIII. 12 pp.
’or. A review of the Panorpide north of Mexico. Bull. Sci.
Lab. Denison Univ., 11: 241-264.
Hilgard, E. W.
’06. Soils, their formation, properties, composition, and relations
to climate and plant growth in the humid and arid regions.
593 pp. New York.
Holdaus, K.
710. Die Siebetechnik zum Aufsammeln der Terricolfauna (nebst
Bemerkungen iiber die Oekologie der im Erdboden lebenden
Tierwelt). Zeit. f. wiss. Insektenbiol., 6: 1-4; 44-57.
"tra. Ueber die Abhangigkeit der Fauna vom Gestein. Ver-
handl.' des. VIII. Internat. Zool.-Kong. zu Graz, Ig10, pp.
726-745.
250
"trb. Ueber die Abhingigkeit der Fauna vom Gestein. Mém. —
1* Cong. Internat. d’Ent., Bruxelles, 1910, 2: 321-344.
Holmes, S. J.
713. Note on the orientation of Bombilius to light. Science,
n. S., 38: 230.
Hopkins, A. D.
’93a. Catalogue of West Virginia Scolytide and their enemies.
Bull. W. Va. Agr. Exper. Sta., No. 31: 121-168.
’93b. Catalogue of West Virginia forest and shade tree insects.
Bull. W. Va. Agr. Exper. Sta., No. 32: 171-251.
‘99. Report on investigations to determine the cause of unhealthy
conditions of the spruce and pine from 1880-1893. Bull. W.
Va. Agr. Exper. Sta., No. 56: 197-461.
o1. Insect enemies of the spruce in the Northeast. U. S. Dept.
Agr., Div. Ent., Bull. No. 28, n.s. 48 pp.
’o2. Insect enemies of the pine in the Black Hills Forest Reserve.
An account of results of special investigations, with recom-
mendations for preventing losses. U. S. Dept. Agr., Div.
Ent., Bull. No. 32, n.s. 24 pp.
*o4. Catalogue of exhibits of insect enemies of forests and forest
products at the Louisiana Purchase Exposition, St. Louis,
Mo., 1904. U.S. Dept. Agr., Div. Ent., Bull. No. 48. 56 pp.
706. The locust borer. U.S. Dept. Agr., Bur. Ent., Bull. No. 58,
Pt. I. 16 pp.
‘og. Barkbeetles of the genus Dendroctonus. U. S. Dept. Agr.,
Bur. Ent., Bull. No. 83, Pt. I. 169 pp.
Hopkins, C. G., Mosier, J. G., Pettit, J. H., and Readhimer, J. E.
*11. Moultrie county soils. Soil Rep. No. 2, Ill. Agr. Exper.
Sta., pp. 3-38.
Hopkins, C. G., and Pettit, J. H.
08. The fertility of Illinois soils. Bull. Ill. Agr. Exper. Sta,
No. 123: 187-294.
,
Howard, L. O.
’g1. The parasites of the Hemerobiine. Proc. Ent. Soc. Wash.,
2: 123-124,
’92a. The Habits of Melittobia. Proc. Ent. Soc. Wash., 2: 244—
248.
’92b. ‘The hymenopterous parasites of spiders. Proc. Ent. Soc.
Wash., 2: 290-302.
251
’95. A secondary effect of the Florida freeze. U.S. Dept. Agr.,
Div. Ent., Insect Life, 7: 413.
’o5. A jumping gall. U.S. Dept. Agr., Bur. Ent., Bull. No.
54:81.
Howlett, M.
’o7. Note on the coupling of Empis borealis. Ent. Mo. Mag.
(2), 18: 229-232.
Hubbard, G. D.
04. A case of geographic influence upon human affairs. Bull.
Ain. Geogr. Soc., 36: I-13.
Hubbard, H. G.
’85. Insects affecting the orange. U.S. Dept. Agr., Div. Ent.
227 pp.
’92. The inhabitants of a fungus. Can. Ent., 24: 250-255.
‘97. The ambrosia beetles of the United States. U. S. Dept.
Agr., Div. Ent., Bull. No. 7, n. s., pp. 9-30.
Hunter, W. D.
‘or. The Aphidide of North America. Bull. Iowa Agr. Exper.
Sta., No. 60: 63-138.
Jackson, C. F.
708. A synopsis of the genus Pemphigus, with notes on their
enemies, importance, life history and geographical distribu-
tion. Proc. Columbus Hort. Soc., 22: 160-218.
Jager, G.
’74. Deutschlands Thierwelt nach ihren Standorten eingetheilt,
Bd. I. 400 pp. Stuttgart.
Johannsen, O. A.
‘op—12. The fungus gnats of North America. Parts I—IV.
Bull. Me. Agr. Exper. Sta.: Pt. I (1909), No. 172, pp. 209—
276; Pt. II (1910), No. 180, pp. 125-192; Pt. III (1912),
No. 196, pp. 249-328; Pt. IV (1912), No. 200, pp. 57-146.
Johnson, K. §.
‘o7. Mean monthly and annual relative humidity charts of the
United States. Rep. S. African Assn. Adv. Sci., 1906, pp.
161-168.
252
Jones, C. H., Edson, A. W., and Morse, W. J.
‘03. ‘The Maple sap flow. Bull. Vt. Agr. Exp. Sta., No. 103:
43-184. °
King, F. H.
’95. Destructive effect of winds on sandy soils and light sandy
loams, with methods of prevention. Eleventh Ann. Rep. Wis.
Agr. Exper. Sta., pp. 292-326.
Knab, F.
‘o5. Observations on Lampyride. Can. Ent., 37: 238-239.
Knapp, J. B.
"12. Fire-killed Douglas fir: a study of its rate of deterioration,
usability, and strength. U. S. Dept. Agr., Forest Service,
Bull. 112. 18 pp. ,
Kraus, E. J., and Hopkins, A. D.
"11. A revision of the powder-post beetles of the family Lycti-
dz of the United States and Europe. Appendix: Notes on
habits and distribution with lists of described species. U. S.
Dept. Agr., Bur. Ent., Tech. Ser. No. 20, Pt. III, pp. 111-138.
Leverett, F.
96. The water resources of Illinois. Ann. Rep. U. S. Geol.
Surv., 1895-96, Pt. II, pp. 1-155.
’99. The Illinois glacial lobe. Monogr. U. S. Geol. Surv., Vol.
38. 817 pp.
Livingston, B. E.
’06. The relation of desert plants to soil moisture and to evap-
oration. Carnegie Inst. Pub., No. 50. 78 pp.
Lugger, O.
’99._ Beetles (Coleoptera) injurious to fruit-producing plants.
Bull. Minn. Agr. Exper. Sta., No. 66: 85-332.
oo. Bugs injurious to our cultivated plants. Bull. Minn. Agr.
Exper. Sta., No. 69: 1-259; Sixth Ann. Rep. Entomol. Minn.
Exper. Sta., 1900. 259 pp.
McAtee, W. L.
’o7. Census of four square feet. Science, n. s., 26: 447-449.
McCook, H. C.
83. How a carpenter ant queen founds a formicary. Proc.
Acad. Nat. Sci. Phil., 1883, pp. 303-307.
’
7
253
’89, ’90, ’93. American spiders and their spinningwork. Vol. I
(1889), 369 pp.; Vol. II (1890), 479 pp.; Vol. III (1893),
284 pp. and atlas. Philadelphia.
McDermott, F. A.
‘ro. A note on the light-emission of some American Lampyride.
Can. Ent., 42: 357-363.
II. Some further observations on the light-emission of Ameri-
can Lampyride: The photogenic function as a mating adapta-
tion in the Photinini. Can. Ent., 43: 399-406.
McGee, W J
"13a. Wells and subsoil water. U. S. Dept. Agr., Bur. Soils,
Bull. No. 92. 185 pp.
’13b. Field records relating to subsoil water. U.S. Dept. Agr.,
Bur. Soils, Bull. No. 93. 40 pp.
McNeil, J.
86. A remarkable case of longevity in a longicorn beetle (Ebu-
ria quadrigeminata). Am. Nat., 20: 1055-1057.
M’Nutt, W., and Fuller, G. D.
"12. The range of evaporation and soil moisture in the oak-
hickory forest association of Illinois. Trans. Ill. Acad. Sci.,
51 129-137
Malloch, J. R.
709. Miltogramma Germari, Mg., and Amaurosoma flavipes,
Fin., from Oxfordshire. Ent. Mo. Mag. (2), 20: 105-106.
12, The insects of the dipterous family Phoridz in the United
States National Museum. Proc. U. S. Nat. Mus., 43: 411-
529. (No. 1938).
Marlatt, C. L.
‘94a. The hibernation of the chinch-bug. U. S. Dept. Agr.,
Div. Ent., Insect Life, 7: 232-234.
’94b. The pear-tree Psylla in Maryland. U.S. Dept. Agr., Div.
Ent., Insect Life, 7: 175-185.
’o7. The periodical cicada. U.S. Dept. Agr., Bur. Ent., Bull.
No. 71. 181 pp.
Mast, S. O.
’12. Behavior of fire-flies (Photinus pyralis) with special ref-
erence to the problem of orientation. Journ. Animal Behav.,
2: 256-272.
?
7
254
Mearns, E. A.
‘03. Feathers beside the Styx. The Condor, 5: 36-38.
Melander, A. L.
‘04. Destructive beetles: a note on landscape gardening. Ent.
News, 15: 19-20. j
Melander, A. L., and Brues, C. T.
’03. Guests and parasites of the burrowing bee Halictus. Biol.
Bull., 5: 1-27.
Metcalf, C. L.
13. The Syrphidae of Ohio. A biologic, economic, and syste-
matic study of the family in the state. Bull. Ohio State
Univ., 17: No. 31. 122 pp.
Moller, L.
67. Die Abhangigkeit der Insekten von ihrer Umgebung.
107 pp. Leipzig.
Moffatt, W. S.
’o9. The higher fungi of the Chicago region. Pt. I—The Hy-
menomycetes. Bull. Chicago Acad. Sci., No. 7, Pt. I. 156 pp.
Montgomery, T. H., Jr.
703. Studies'on the habits of spiders, particularly those of the
mating period. Proc. Acad. Nat. Sci. Phil., 1903, pp. 50-149.
’og. Further studies on the activities of araneads, II. Proc.
Acad. Nat. Sci. Phil., 55: 548-569.
Moore, W. L.
’10. Descriptive meteorology. 344 pp. New York.
Morgan, H. A., and Dupree, J. W.
703. Development and hibernation of mosquitoes. U. 8. Dept.
Agr., Div. Ent., Bull. No. 40, n. s., pp. 88-92.
Morse, A. P.
’04. Researches on North American Acridiidae. Carnegie Insti-
tution, Pub. No. 18. 55 pp.
Mosier, J. G.
’03. Climate of Illinois. Bull. Ill. Agr. Exper. Sta., No. 6:
45-76.
255
Motter, M. G.
’98. A contribution to the study of the fauna of the grave. A
study of one hundred and fifty disinterments, with some ad-
ditional experimental observations. Journ. N. Y. Ent. Soc.,
6: 201-231.
Murtfeldt, M. E.
’94.__ Acorn insects, primary and secondary. U. S. Dept. Agr.,
Div. Ent., Insect Life, 6: 318-324.
Needham, J. G.
’98. Outdoor studies: a reading book of nature study. 90 pp.
New York.
oo. The fruiting of the blue flag (Iris versicolor L.). Am.
Nat., 34: 361-386.
Needham, J. G., and Betten, C.
?or. Aquatic insects in the Adirondacks. Bull. N. Y. State
Mus., No. 47: 383-612.
Ortmann, A. E.
706. The crawfishes of the State of Pennsylvania. Mem. Car-
negie Mus., 2: 343-523.
Osborn, H.
‘oo. Report on insects of the season in Iowa. U. S. Dept. Agr.,
Div. Ent., Bull. No. 22: 18-41.
g2. Catalogue of the Hemiptera of Iowa. Proc. Iowa Acad.
- Sci., Vol. I, Pt. II, pp. 120-131.
Osborn, H., and Ball, E. D.
97a. Contributions to the hemipterous fauna of Iowa. Proc.
Iowa Acad. Sci., 4: 172-234.
’97b. Studies of the life histories of grass-feeding Jassidae.
Bull. Iowa Agr. Exper. Sta., No. 34: 612-640.
Osborn, H., and Sirrine, F. A.
94. Notes on injurious insects. Bull. lowa Agr. Exper. Sta.,
No. 23: 881-905.
Osten-Sacken, C. R.
’82. Ueber das Betragen des californischen fliigellosen Bittacus
(B. apterus MacLachl.). Wiener Ent. Zeit., I Jahrg., pp.
123-124.
,
a
256
Parrott, P. J., and Fulton, B. B.
"14. Tree crickets injurious to orchard and garden fruits. N. Y.
Agr. Exp. Sta. Bull., No. 388, pp. 417-461. Geneva.
Packard, A. S. Jr.
773. Our common insects. 225 pp. Salem, Mass.
’83. Guide for the study of insects. Eighth ed. 715 pp. New
York,
’90. Insects injurious to forest and shade trees. Fifth Rep.
U. S. Ent. Comm. U. S. Dept. Agr. 957 pp.
“95, 05, "14. Monograph of the Bombycine moths of North
America, including their transformations and origin of the
larval markings and armature :—
Part I. (’95) Family 1—Notodontide. Mem. Nat. Acad.
Sci., Vol. VII. 291 pp.
Part II. (‘o5) Family Ceratocampide, subfamily Cerato-
campinz. Mem. Nat. Acad. Sci., Vol. IX. 149 pp.
Part III. (’14) Families Ceratocampide (exclusive of Cera-
tocampinz), Saturniide, Hemileucide and Brahmeide. Edited
by T. D. A. Cockerell. Mem. Nat. Acad. Sci., Vol. XII.
516 pp.
798.