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Title: The Forest Habitat of the University of Kansas Natural History Reservation
Author: Fitch, Henry S., 1909-2009, McGregor, Ronald L.
Language: English
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Copyright Status: Not copyrighted in the United States. If you live elsewhere check the laws of your country before downloading this ebook. See comments about copyright issues at end of book.

*** Start of this Doctrine Publishing Corporation Digital Book "The Forest Habitat of the University of Kansas Natural History Reservation" ***

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                   University of Kansas Publications
                       Museum of Natural History

    Volume 10, No. 3, pp. 77-127, 2 pls., 7 figs. in text, 4 tables
                           December 31, 1956

                The Forest Habitat of the University of
                   Kansas Natural History Reservation



                          UNIVERSITY OF KANSAS


         Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
                            Robert W. Wilson

    Volume 10, No. 3, pp. 77-127, 2 pls., 7 figs. in text, 4 tables
                      Published December 31, 1956

                          UNIVERSITY OF KANSAS
                            Lawrence, Kansas

                               PRINTED BY
                    FERD VOILAND, JR., STATE PRINTER
                             TOPEKA, KANSAS


                The Forest Habitat of the University of
                   Kansas Natural History Reservation


                 HENRY S. FITCH and RONALD L. MCGREGOR


In northeastern Kansas, before it was disturbed by the arrival of
white settlers in the eighteen fifties, tall grass prairies and
deciduous forests were both represented. These two contrasting types
of vegetation overlapped widely in an interdigitating pattern which
was determined by distribution of moisture, soil types, slope exposure
and various biotic factors.

The early explorers who saw this region, and the settlers who came
later, left only incomplete descriptions, which were usually vague as
to the locality and the species of plants represented. As a result,
there is but little concrete information as to the precise boundaries
between the forests and grasslands, and opinions differ among
ecologists. No representative sample of either type remains.

It may be assumed that the plant communities existing one hundred
years ago and earlier were far more stable than those of the present
that have resulted from man's disruptive activities. This stability
was only relative, however. Within the last few thousand years since
the final withdrawal of the Wisconsinan ice sheet, fairly rapid and
continual change must have occurred, as a result of changing climate,
the sudden extinction of various large, dominant mammals, and finally
the impact of successive aboriginal cultures.

The land north of the Kansas River had been a reserve for the Delaware
Indians. This land was thrown open to settlement as a result of two
separate purchases from the tribe, in 1860 and 1866. The alluvial
bottomlands were fertile and soon were under cultivation.


Because the prairies and forests were soon destroyed or altered by
cow, ax, plow and fire, knowledge of the region's ecology under the
conditions that prevailed in the early nineteenth century and the
centuries before must be gained largely from circumstantial evidence.
Although there were no ecologists among the first settlers in Kansas,
occasional glimpses of the region's ecology are afforded by the writings
of early residents who mentioned native plant and animal life from time
to time. However, such mention was usually casual and fragmentary.

A brief early description of forest in northeastern Kansas, which is
casual and incomplete, and perhaps misleading, since it differs from
later accounts, was included in Major W. S. Long's report of the
exploring expedition that passed through country now included in
Johnson, Douglas, Shawnee, Wabaunsee, Riley, Pottawatomie, Jackson,
Jefferson and Leavenworth counties in 1819. "The catalogue of the
forest trees in this region is not very copious. The cottonwood and
the plane tree [sycamore] everywhere form conspicuous features of the
forests. With these are intermixed the tall and graceful acacia, the
honey locust, and the bonduc, or coffee-tree, and carya [hickory] and
fraxinus [ash] ..." (Taft, 1950:442).

A description of the country in northern Douglas County and adjacent
Leavenworth County, while it was still in virtually undisturbed
condition, was written by Mr. George S. Parks (1854). Travelling up
the Kansas River from the Missouri state line he described the
vegetation and physiography with respect to specific landmarks that
can be easily located at the present time. His descriptions of the
areas he saw that were nearest the Reservation, are quoted below, in

[Travelling west from near the mouth of Stranger Creek 10 miles ESE
Reservation.] "... bluff with open woods and high rolling prairie in
background. On the south side of the river ... grass and scattering
timber forming a green lawn back with high prairie. In this
neighborhood the shore is rocky. We passed a bald bluff on the north,
with a rich bottom on the south side, and a high open lawn in the
rear. A little farther on the elevated prairies strike the river,
giving a charming variety of scenery--while on the north are extended
bottoms of rich timbered lands.

"In this vicinity we saw many Indians along the banks; we also passed
a grape thicket, in the bottom, spread over several thousand
acres--while just above, on our right, rose a rocky bluff, covered
with open woods. A little above this Sugar Creek empties into the
Kansas, from the right; and a little farther up, there is a low
bluff--a short distance beyond, there being another fine grape
thicket, and rich walnut bottom. On the right side of the river ...
rises a beautiful undulating eminence ... open woods and a fine
prairie about a mile back.

"On the left, a short distance above, the Wakarusa flows in--a
considerable stream--with good timber for some way back.

"On both sides of the river, above the Wakarusa, there are excellent
bottom lands; ... farther up on the south bank, the high prairie comes
down to the water's edge.... away as far as the eye could reach in a
southwest direction, the prairies were high and rolling, like the
waves of old ocean--southward, beautiful groves dot the prairie and
the dark line of timber that stretches along the Wakarusa Valley--with
the great Prairie-mound ... fixed there as a landmark of perpetual
beauty--the meandering river with its dark skirting forests of timber
on the north ... Proceeding north, high rich bottoms extend for many
miles and we saw vast thickets of grape-vines, pea-vines etc. and
paw-paws. The timber was principally oak, walnut, ash, hickory,
mulberry, hackberry, linden, cottonwood and coffee-bean.

[Between the Reservation and the mouth of the Delaware River, 10 miles
west.] "A few miles below the mouth of the Grasshopper [Delaware] on
the north the prairie undulates gradually back from the river as far
as the eye can reach ... between the Grasshopper and Mud Creek there
is a prairie bottom where pioneers are making claims."

In 1855 Mrs. Sara T. D. Robinson, wife of Dr. Charles Robinson who was
the first governor of Kansas, described in her diary the environs of
Lawrence (1899). In part, the areas described by her overlap those
described by Parks, and both writers impart similar impressions. Mrs.
Robinson's writing was concerned chiefly with the social and political
affairs of the territory and the occasional comments on the "scenery"
in her voluble accounts must be regarded as impressions rather than
purposeful and accurate descriptions, as certain inconsistencies are
apparent. Excerpts from several of her more significant descriptive
passages are quoted below. [Between Lawrence and Kansas City, April
17, 1855.] "... prairie stretching in all directions, noble forests
marking the line of the rivers and creeks, ... tall oaks and walnuts
grouped in admirable arrangement ... there were deep ravines ...
skirted with graceful trees, while the water in their pebbly beds is
limpid and clear." [North of Wakarusa Crossing.] "... stumps in every
direction in the woods ..." [At Lawrence, April 18, 1855.] "The town
reaches to the river, whose further shore is skirted with a line of
beautiful timber, while beyond all rise the Delaware lands, which in
the distance have all the appearance of cultivated fields and
orchards.... A line of timber between us and Blue Mound marks the
course of the Wakarusa, while beyond the eye rests upon a country
diversified in surface, sloping hills, finely rolling prairies, and
timbered creeks ... to the northwest there is the most delightful
mingling together of hill, valley, prairie, woodland, and river ...
fine grove about a mile west of town, one of Nature's grand old

[On trip to visit a neighbor four miles away from Lawrence.] "There
were high, conical hills, bearing on their tops forest trees, with
dense, thick foliage; at the next moment a little shady nook, with a
silvery rivulet running over its pebbly bed...."

[On trip west toward Topeka.] "Timber was more abundant, not only
marking the line of the creeks, but crowning the summit of many an

[At Lawrence.] "Lawrence and its surroundings, of river flowing
beneath the dim forests two miles deep on the north bank...."

Parks' and Robinson's accounts seem to show that in general
bottomlands and stream courses were wooded, and uplands were mainly
prairie, but that local deviations from this pattern were numerous,
with trees and groves isolated or partly isolated in a variety of
situations. This condition suggests that prairies were then
encroaching into formerly wooded areas. A climatic shift toward hotter
and drier conditions, or a change in native practices, with more
frequent burning, might have brought about the trend.

Further information concerning the distribution and composition of the
forest is afforded by a series of letters from the settlers at
Lawrence, Kansas, that were printed in various Boston newspapers and
in the Milwaukee Daily Sentinel, in 1854, 1855, and 1856. In nine such
letters which discuss, among other things, the availability of timber,
several kinds of trees are listed. Oak (species not mentioned), black
walnut, and cottonwood are each listed in seven of the nine letters,
while elm, hickory and "white walnut" are each listed in two, and ash,
hackberry, sycamore, basswood, willow and locust are each mentioned
only once. Copies of these letters are in the files of Dr. James C.
Malin, to whom we are much indebted for the privilege of examining
them, and for his critical reading of parts of the manuscript.

Early U. S. Government maps of northeastern Kansas show the
distribution of forest in the late eighteen fifties, and in general
the pattern agrees well with that indicated by the accounts of Parks
and Robinson. Through the kindness of Dr. Malin, we have been
permitted to examine his photostatic copies of a series of these early
maps, covering the area discussed in our study, and made in the
period extending from 1855 through 1860. A tracing taken from parts of
two of these maps, showing the Kansas River north and east of
Lawrence, and the area between the river and the north boundary of
Douglas County, is reproduced in Fig. 1. For comparison, a map of the
same area showing the stream courses and the distribution of timber,
as traced from recent U. S. Geological Survey maps, is reproduced in
Fig. 2.

The early maps agree with Parks' and Robinson's descriptions in
showing an extensive belt of timber in the flood plain north of the
river, and narrower belts of timber along its tributary streams. In
Fig. 1 the courses of the Kansas River and of Mud Creek agree fairly
well with those shown on modern maps, but there are gross errors in
the minor drainage systems of the sections of land in the northeastern
part. Other evidence indicates that the distribution of forest was
much different than that shown in this part of the map. Field work by
the map-makers in this marginal area must have been extremely sketchy.
Dr. Malin explains that such inaccuracies are to be expected because
the contracts for mapping were made on a political basis, with little
or no regard for other qualifications of the applicant.

The University of Kansas Natural History Reservation is in the
northeasternmost section (Section 4, Township 12S, Range 20E) of
Douglas County, Kansas. Topographically, it is almost evenly divided
into three parts: (1) peninsular extensions of the Kansas River
Valley, sloping gradually up to a level approximately 100 feet above
that of the flood plain; (2) hilltops 200 feet or more above the level
of the flood plain; (3) steep slopes from the hilltops to the valley

The land that is now the Reservation was part of a tract acquired in
the eighteen sixties by former governor Charles Robinson, after the
Delaware Reserve lands in the northeastern part of Kansas Territory
were sold by the tribe. The section of land now comprising the
Reservation was used primarily for grazing after Robinson acquired it.
However, several squatters settled on the area and cultivated small
acreages for periods of years in the eighteen seventies and eighteen
eighties. In the eighteen nineties parts of the area including some of
the hillsides were still covered with a mixed forest of virgin timber
(_fide_ Frank H. Leonhard in conversation, October 19, 1951). Mr.
Leonhard, who was long in the employ of the Charles Robinson family,
remembered the area as far back as the early eighteen nineties when he
worked on it cutting timber. He remembered, especially, cutting large
walnut trees as much as two feet in diameter, which were valuable
timber, but he thought that elm also was abundant at that time. By
then the area, separated into east and west halves by a rock wall, had
already been heavily grazed, and the original prairie vegetation,
presumably dominated by big bluestem, had been much altered. The open
upland portions were dominated by blue grass.

  [Illustration: Fig. 1. Tracing from early (1855-60) U. S.
      Government maps of northeastern Douglas County, Kansas, and
      adjacent western edge of Leavenworth County, showing stream
      courses and approximate distribution of woodland before
      deforestation had occurred. Section 4 to right of center at
      upper edge of figure, is now mostly included in the University
      of Kansas Natural History Reservation. Note inaccuracies in
      drainage systems on this part of map as compared with Fig. 2.]

  [Illustration: Fig. 2. Tracing from 1950 U.S. Geological Survey
      maps of same area shown in Fig. 1, indicating present
      distribution of woodland, and the pattern of drainage systems.]

By about 1900 control of the area had passed to the J. F. Morgan
family. The homesteads had long since been deserted and the entire
area was used for grazing (_fide_ J. F. Morgan, in conversation,
January 13, 1952). Parts of the bottomland were fenced and broken for
cultivation in 1907, 1912, and 1915, and hilltop fields were first
cultivated in 1909. Tree cutting was more or less continual. Many of
the old stumps still present on the area are remnants of the trees cut
in the "twenties" or even earlier. Several acres of hilltop and south
slope in the northwest corner of the area were protected from
livestock and maintained for harvesting of prairie hay. The hay was
mowed annually, and the vegetation was burned at less frequent
intervals, usually in early spring. This treatment served to kill
encroaching woody vegetation and to maintain a prairie type.

In the mid-thirties control of the area passed to the University of
Kansas. At that time a program of development was launched by the
University and the U. S. Soil Conservation Service with relief labor
(_fide_ C. G. Bayles in conversation, November 10, 1953). The work
included: filling gullies, digging diversion ditches and building
check dams and terraces to prevent erosion; clearing extensive
thickets; bindweed eradication from the cultivated areas; and fencing
off the wooded hillsides from the valley and hilltop pastures for
protection from livestock. This work extended over several years, and
one main objective was to utilize the area for growing timber.
However, plans to make extensive plantings of walnut and other
valuable timber never materialized. In the forties the check dams fell
into disrepair. The area was leased to a farmer and was again heavily
overgrazed. In this period there was some tree-cutting by the
University's Department of Buildings and Grounds and by farmers, but
this cutting was not on a commercial scale and was mainly for firewood
and fence posts. One of the chief results of fencing off the wooded
hillsides was that shrubs and young trees, formerly held in check by
livestock, were allowed to flourish. Understory thickets sprang up
throughout most of the woodland, and especially in edge situations.

Late in 1948, after the area had been made a Reservation, livestock
were excluded. In the years following, the parts of the closely grazed
pastures adjacent to woodland passed through stages similar to those
that had occurred 10 to 12 years earlier in the parts protected by
fences. Young trees and shrubs sprang up in thickets, the numbers and
kinds depending on amount of shade, seed sources, soil, moisture, and
various other factors.

Although most of the tree-cutting was done prior to 1934, annual
growth rings are discernible on many of the old stumps, indicating the
age of the tree at the time it was cut. Occasionally the stumps
produced sprouts which had grown into sizable trees by 1954. In such
instances the year that the tree was cut and the year that it
originally began growing could be determined from a study of the
annual growth rings. In 54 instances ring counts were obtained from
stumps or logs, or from trees that had been split and fallen in wind

Stumps that were otherwise intact often had small central cavities an
inch or more in diameter. For these it was necessary to estimate the
numbers of missing rings in order to obtain a figure for the
approximate total age of the tree at the time it was cut. Many of the
logs and stumps were so much decayed that growth rings were no longer
distinct, and on most there were a few rings that were not clearly
defined. In the majority of instances the time of cutting could not be
determined accurately, but it is known that there was little
tree-cutting after 1934 on most parts of the area. Probably most of
the stumps on the Reservation that were well enough preserved to
provide counts were from 20 to 30 years old. Most of the counts of
growth rings on chestnut oaks were obtained on a hillside adjoining
the Reservation where the trees were cut in the early nineteen

Width of the annual growth rings reflects rapidity of growth in the
tree and is determined, in part, by the amount of annual rainfall,
especially in this region on the western edge of the deciduous forests
where moisture is the chief limiting factor. Periods of drought or of
unusually heavy rainfall may result in growth rings smaller or larger
than average. Because the trees draw moisture from the deeper soil
layer there is a lag in their response to precipitation, and a single
year that is much wetter or much drier than those preceding or
following it may not stand out clearly in the annual rings. In
individual trees the effect of precipitation is often obscured by the
effects of crowding and shading by competitors, injury or disease.
None of the trees examined for growth rings reflected the annual
precipitation accurately for long periods though some indication of
known drought periods or of series of wet years were usually

For 35 black oaks, chestnut oaks, and American elms, growth rings
averaged 3.81 per inch of trunk diameter (according to size of the
tree; 5.1 rings per inch in those trees 9 to 12 inches in diameter,
4.0 in those 13 to 15 inches, 3.6 in those 16 to 24 inches, and 2.8 in
those of more than 24 inches). Data from a few complete counts and
many incomplete counts indicate that in _Gleditsia triacanthos_ growth
is much more rapid, with only 2 to 3 rings per inch of trunk diameter,
whereas in _Juglans nigra_, _Celtis occidentalis_, _Carya ovata_, and
_Fraxinus americana_ growth is much slower, with usually five or more
growth rings per inch of trunk diameter. Individual trees deviate
widely from the average for their species, and those in rich
bottomland soil grow more rapidly than those in shallow soil of
hilltops or those on rocky slopes. If such factors are taken into
account the ages of trees may be estimated from the diameters of their
trunks. In mature trees growth slows; age is likely to be
underestimated rather than overestimated in those of exceptionally
large size.

The belief that this and similar areas in northeastern Kansas were
virtually treeless at the time of occupation by white settlers is
shown to be wholly unfounded by the information obtained from growth
rings. The ring counts show that many trees now growing on the area
and others cut within the last 30 years, but still represented by
stumps, were already present in the eighteen sixties when the area was
first occupied. A few trees on the area probably are much older,
dating back to the early eighteen hundreds. As there are no virgin
stands of timber, and the more valuable trees have been removed by
selective cutting at various times, it is to be expected that there
are few or no trees on the area approaching the potential longevity
for their species.

The many oaks and elms on the area that are more than two feet in
trunk diameter mostly date back to the eighteen sixties or earlier.
The distribution of the larger trees and stumps provides a clue as to
the original distribution of forest and grassland on the area. There
is no description available of the area that is now the Reservation in
its original condition. However, Mrs. Anna Morgan Ward (1945) has
recorded comments on the appearance of the country in the section of
land adjoining the Reservation on the south, as it appeared when her
family settled there in 1864. This land differed from that of the
Reservation, as it consists of low rolling hills, well drained with
predominately south exposure, and with sandy soil. It adjoins the
present flood plain of the Kansas River, and consists partly of the
old Menoken Terrace deposited in the Pleistocene. The following
excerpts from Mrs. Ward's manuscript are selected as most descriptive
of the original vegetation on this section of land.

[In southwest part of section near the Morgan house.] "... some hills
that were covered with Jack Oak trees ... Here we found wild
strawberries on the hillsides. And along the creeks we located
gooseberry bushes, wild grapes, both summer and winter grapes, plums,
and paw paws in the fall. We found a crabapple tree ... Plenty of
walnuts and hazel nuts."

[Hilly south-central part of section, the J. P. Whitney farm.] "... on
a hill among many small trees ... especially on the east were many

[Less hilly southeastern part of section.] "... Was open prairie and
free grazing ground for many years...."

Much of the land in this section is now under cultivation but there
are still hilltop groves of blackjack oak, probably in about the same
places where Mrs Ward noticed them 90 years ago--south of the house
that was formerly Robinson's residence, and west across the county
road, beside the Oakridge School building, and on other knolls to the
east and southeast.

The bottomland areas of the Reservation are mainly grassland and no
old stumps remain to indicate that trees were formerly present.
Nevertheless, it might be expected that under original conditions
these bottomland areas supported forests, as the soil is deep and rich
with abundant moisture. Also most of the early accounts agree that
forests occurred mainly along stream courses in this region.
Presumably these areas were cut over early, because they were most
accessible, and because they supported the best stands of timber.

One of the best indications of the former vegetation on these
bottomland areas is provided by old bleached shells of snails and
certain other mollusks, brought to the surface by plowing in
cultivated fields adjoining the Reservation on the south and west
(Fitch and Lokke, 1956). A high proportion of the shells are of
species limited to humus soil, decaying logs, or leaf litter in moist
woodlands (_Stenotrema leai_, _Retinella electrina_, _Zonitoides
arboreus_, _Vertigo ovata_, _Helicodiscus parallelus_), to wet places
(_Lymnaea parva_, _Succinea avara_) or even to standing pools (_Physa
hawni_, _Helisoma trivolvis_, _Pisidium compressum_). No living
mollusks could be found in these fields and none could be expected to
survive on land that is cultivated annually. As a whole the assemblage
seems to be indicative of a humid, poorly drained forest habitat.
Presumably most of the shells or all of them are more than 100 years
old, antedating the time when the area was first disturbed by human
activities, and also antedating the time when the creeks (now 15 feet
or more below the fields) had begun to erode their channels. That the
shell deposits are of no great antiquity, and represent conditions
prevailing within the last few hundred years, is suggested by the fact
that all are species still living in Douglas County, and with one
exception, all still live on the Reservation.

  [Illustration: Fig. 3. Tracing from a contour map made in 1914,
      of the two small valleys on the Reservation, showing changed
      position of contour lines at gullies by 1952. As a result of
      overgrazing, and cultivation of part of the upland drainage
      area, there was relatively rapid erosion in the 38-year

  [Illustration: Fig. 4. Map of University of Kansas Natural
      History Reservation, with 20-foot contours, showing probable
      approximate distribution of forest in early eighteen hundreds
      (vertical lines show slopes and hilltops that are still wooded;
      grid pattern shows bottomlands that were formerly wooded but
      later cleared for pasture or cultivated crops). Stippled areas
      show those slopes and hilltops now wooded seemingly as a result
      of recent reinvasion, that probably were bluestem prairie
      earlier. Unshaded areas are relatively flat hilltops that are
      still grassland and are thought to have been bluestem prairie.]

Mrs. Ward (_op. cit._) in her manuscript concerning the early history
of Grant Township, mentioned the small creek that drains the east part
of the Reservation. Evidently in the sixties it had a more constant
flow, usually with clear water. Later it eroded its channel, cutting a
deep gully. Presumably the water table has been much lowered. In his
verbal reminiscences of the area, Mr. J. F. Morgan told us that in the
nineties this stream had eroded its channel but little within the
present limits of the Reservation. In a period of years, 1902 to 1905
inclusive, when there was abnormally heavy rainfall, severe erosion
occurred, and the saturated soil of several hillside areas slipped
downhill to the extent of several feet vertical displacement. The
ravine draining into the present pond from the north was known as
"Sunken Canyon" because of such soil slips. However, a map of the
Reservation and surrounding areas made by the University of Kansas
Department of Civil Engineering in 1914, shows that by that time
relatively little gullying had occurred. Comparison of this contour
map with a more detailed one prepared in 1952 shows that the gullies
had eroded their channels to depths more than 15 feet greater in some
places, in the 38-year interval (Fig. 3). In June and July, 1951, when
there was unusually heavy rainfall, gullies deepened perceptibly.
Dozens of trees including many large mature elms, honey locusts, and
osage orange, growing along the banks were undermined and fell into
the gullies.

Composition of the Forest

Under present conditions, every one of the larger tree species
dominates at least some small part of the area. For reasons that are
usually obscure, locations that seem otherwise similar differ in the
kinds, numbers, and sizes of trees they support. Probably most of
these differences have arisen in the varying treatments under human
occupation in the last 100 years.

In the two valley areas, presumably heavily wooded under primitive
conditions, the trees growing at present seem to be secondary
invaders. They include groves and isolated trees of elm, honey locust,
walnut, and osage orange, and an occasional red haw, hackberry, or

The hilltops likewise are chiefly open, but forest of the hillsides
encroaches onto them for as much as 100 yards in some places. The
slopes between the hilltops and the valleys are almost everywhere
wooded, but the aspect of the woods changes from place to place.
Subdivisions on a vertical scale, might be recognized as follows: the
upper limestone outcrop (Plattsmouth member) at the hilltop; the
usually steep slope strewn with rocks, between the upper and lower
(Toronto) limestone outcrop; the lower limestone outcrop; an almost
level terracelike formation often approximately 50 feet wide a few
feet below the level of the Toronto limestone; the slope below the
terrace, variable in steepness, exposure, and soil type, and usually
several times more extensive than the first four subdivisions
combined. Along both the upper and lower outcrops, elm and hackberry
are especially prominent. Chestnut oak is abundant along the outcrops
and on the rocky slope between them in some situations. Ash grows
abundantly on some upper slopes but there are few growing on the upper
outcrop. On the terrace, elm, ash, hackberry, honey locust,
coffee-tree and black oak are abundant. On the lower slopes grow most
of the blackjack oaks, post oaks, red oaks and mulberries.

Even greater differences in the local aspect of woodland on the
hillsides are caused by slope exposure. On south facing slopes,
especially, the woodland is noticeably different from that in other
situations, and of more xeric aspect. The climax species, _Quercus
Muehlenbergii_, _Q. rubra_, _Q. velutina_ and _Carya ovata_ are almost
totally absent. Such trees as are present are of small to medium size.
They are mostly red elm, American elm, walnut, honey locust,
hackberry, and osage orange, with dogwood (_Cornus Drummondii_) and
plum (_Prunus americanus_) forming dense thickets. Occasional patches
of prairie grasses remain in more exposed situations where they have
not been shaded out. These, together with the small size of most of
the trees, indicate that the south slopes have become wooded rather
recently, and originally were prairie. Nevertheless, the small
remaining groves of blackjack oak and post oak are on slopes that face
south, southeast, or southwest, and probably under original conditions
they occupied these situations, separate from the forests of other
hardwoods. Slopes facing east, west, and north, are more similar in
relative abundance of various kinds of trees, and they do not differ
much from hilltop edges that are wooded. Chestnut oak and hickory are
most abundant on north slopes, and ash occurs mainly on north slopes.

    Table 1.--Percentages of Larger Trees (a Foot or More in Trunk
    Diameter) on Different Slope Exposures.

                         |  North   |            |   West  |   South
                         |  slopes  |   Hilltops |  slopes |   slopes
  Elm                    |   35.7   |   38.6     |  25.8   |   51.4
  Chestnut oak           |   22.0   |   18.3     |  17.8   |    2.9
  Hickory                |    8.8   |    4.0     |   3.6   |    5.0
  Walnut                 |    8.8   |    5.8     |  19.6   |   12.1
  Ash                    |    7.1   |     .8     |         |     .4
  Hackberry              |    8.2   |    1.6     |   2.4   |    6.9
  Black oak              |    3.3   |   16.4     |         |    1.0
  Red oak                |    2.2   |            |   23.8  |
  Locust                 |    1.8   |    7.5     |    1.9  |   11.6
  Osage orange           |     .5   |    1.5     |     .2  |    5.3
  Sycamore               |     .5   |            |    2.1  |     .1
  Coffee-tree            |          |    1.2     |    2.4  |    1.0
  Cherry                 |          |    2.4     |         |     .1
  Red haw                |          |     .4     |         |    1.3
  Ailanthus              |          |            |         |     .3
  Mulberry               |          |     .5     |         |     .1
  Cottonwood             |          |            |         |     .1
  Redbud                 |          |     .8     |     .2  |     .1
  Boxelder               |          |     .1     |         |     .3
  Blackjack oak          |          |            |     .2  |
                         |          |            |         |
  Total trees in sample  |  182     |  890       |  467    |  898

Table 1 shows the percentages of different kinds of trees a foot or
more in trunk diameter on different slope exposures sampled. Elm is
almost always the dominant tree, making up from one-fourth to one-half
of the total stand. The other species dominate relatively small areas.
Chestnut oak usually makes up a substantial part of the stand on
hilltops and slopes of north, east, or west exposure. Black oak, red
oak, and walnut may be prominent on the east and west slopes. Walnut
and locust are prominent on south slopes.

Hickory usually has a trunk diameter of less than one foot, and,
therefore, it is not prominent anywhere among the larger trees. Table
2, showing ratios of medium-small trees (more than 6 inches and less
than one foot in trunk diameter) demonstrates that hickory is one of
the more prominent trees on hilltops and on slopes other than those of
south exposure.

Invasion of Fields

In 1948 when the extensive open parts of the Reservation were grazed
and cultivated, small trees were inconspicuous and few. Mature trees,
with trunk diameters of 9 inches to more than two feet, were
distributed over the pastured areas, however, with groves of American
elm, honey locust, and walnut near the edges of the woods, and
occasional scattered trees of these species and of osage orange,
coffee-tree, red haw, hackberry, and ash.

    Table 2.--Percentages of Different Kinds of Small Trees (Six
    Inches to a Foot in Trunk Diameter) on Different Slope Exposures.

                             | North  | Hilltops | West   | South
                             | slopes |          | slopes | slopes
                             |        |          |        |
  Elm                        |  29.6  |   29.9   |  34.6  |  57.9
  Chestnut oak               |  29.6  |   17.5   |  15.5  |    .4
  Hickory                    |  11.1  |   25.4   |  28.4  |    .8
  Walnut                     |   5.6  |     .7   |   7.4  |   5.3
  Hackberry                  |  13.0  |    1.0   |   3.7  |  26.4
  Black oak                  |   1.9  |   16.3   |        |
  Red oak                    |   1.9  |          |   6.8  |
  Locust                     |        |    3.3   |        |   3.0
  Osage orange               |        |    2.0   |        |   1.5
  Coffee-tree                |   1.9  |     .7   |        |   1.1
  Cherry                     |        |          |        |    .4
  Red haw                    |        |    2.4   |        |
  Mulberry                   |        |     .7   |        |
  Redbud                     |   9.3  |          |   3.7  |    .8
  Boxelder                   |        |          |        |   2.6
                             |        |          |        |
  Total trees in sample      |    54  |    295   |   162  |   266

In 1949 soon after the discontinuance of grazing and cultivation, a
large crop of tree seedlings became established. Each year thereafter
the numbers were augmented by new crops of seedlings, but conditions
rapidly became less favorable for their establishment, as the ground
cover of herbaceous vegetation became thicker. The numbers and kinds
of young trees that became established differed markedly in different
situations. The seedlings present in large numbers were those of elm,
honey locust, boxelder, dogwood, walnut, osage orange and crab-apple.
There was none of the climax species--oaks or hickories--in the

    Table 3.--Numbers of Young Trees Per Acre in Fields of the
    Reservation, June, 1952.

                       |          |       |Bottomland|Hilltop |
                       |Bottomland|Hilltop|  fallow  |fallow  |Prairie
                       | pasture  |pasture|  field   | field  |
  No. of 1/100 acre    |    250   |    80 |     70   |    80  |    50
    plots sampled      |          |       |          |        |
                       |          |       |          |        |
  Honey locust         |   83.0   |  58.8 |          |   5.6  |
  Elm                  |   80.0   |  72.5 |  138.8   | 230.0  | 150.0
  Boxelder             |    1.6   |   1.2 |   22.9   |        | 200.0
  Dogwood              |   18.8   |  18.8 |   11.4   |  51.2  |  44.0
  Walnut               |    2.0   |  50.0 |    7.15  |        |
  Osage orange         |   16.0   |  48.7 |          |        |
  Crab-apple           |    7.2   |  93.8 |          |   1.2  |
  Red haw              |    5.2   |  17.5 |    2.8   |   2.5  |   4.0
  Coffee-tree          |    4.8   |   1.2 |          |        |
  Hackberry            |    2.8   |       |          |        |   2.0
  Cottonwood           |     .2   |       |          |        |
  Ash                  |          |   8.8 |          |   3.7  |
  Plum                 |     .8   |       |          |        |
  Peach                |     .2   |       |          |        |
  Cockspur thorn       |     .8   |  21.3 |          |        |
  Sycamore             |     .4   |       |          |   1.2  |
  Cherry               |          |   1.2 |          |        |   2.0
                       |          |       |          |        |
  Total number counted |    236   |   393 |    279   |   296  |   402

Table 3 shows the numbers of young trees counted in a total of 530
plots of 1/100 acre each, in June, 1952. The trees counted included
all those approximately one foot high or larger. A few were up to 12
feet tall, but most were between one foot and five feet in height. Not
included were the many smaller seedlings, which were mostly concealed
beneath the dense layer of low herbaceous vegetation.

Of young trees there were most on the bluestem prairie area, less on
the former pastures and least on the fallow fields. In both the
pasture areas and the fallow fields, the bottomlands had fewer trees
than the hilltops--60 per cent and 94.3 per cent, respectively. In
every instance the abundance of young trees seemed to be inversely
proportional to the amount of competing herbaceous vegetation. The
bottomland fallow fields, which had the fewest tree seedlings, were
dominated by a rank growth of giant ragweed and sunflower, often as
much as ten feet tall, effectively shutting most of the light from the
tree seedlings. By 1954, however, the sunflower was nearly eliminated,
and the giant ragweed, though still abundant, was much stunted.

The bluestem prairie on an area of hilltop and upper slope had not
been burned over or otherwise disturbed for some years prior to 1948,
and probably trees began to invade this area years before they invaded
the fallow fields and pastures accounting, in part, for their greater
abundance in 1952. Approximately half of the young trees on this
prairie area were boxelders, which were relatively scarce on the other
four areas. Elm was either first or second in abundance on each area.
On both types of pasture areas honey locusts were appearing in
abundance and osage orange seedlings were present in somewhat smaller
numbers. However, these two kinds of trees were almost entirely absent
from the other areas sampled, except that a few locusts were recorded
on a hilltop fallow field. In 1948 honey locust seeds were noticed in
great abundance in the droppings of cattle; their dispersal in this
manner probably is in large part responsible for the abundance of
young honey locusts throughout the former pastures. Osage orange may
have been distributed in the same manner. Seedlings of dogwood were
moderately numerous on each one of the areas sampled, and those of red
haw were somewhat less abundant on each area. Crab-apple was the most
abundant species invading the hilltop pastures but was scarce or
absent in the other situations. The remaining species of trees,
including coffee-tree, hackberry, cottonwood, ash, plum, peach,
cherry, cockspur thorn, sycamore, and redbud, each made up only a
small percentage of the tree crop in the situations where they

In late July and early August, 1954, counts of young trees were made
again on the upland pasture area, with a total of 200 1/100-acre plot
samples. This sample was taken at the end of one of the longest and
most severe droughts in the history of the area. Both 1952 and 1953
had drought summers, and up to the end of July the summer of 1954 was
exceptionally dry also. The conditions of the young trees at this
time, in the relatively dry and shallow hilltop soil, was especially
significant. As might have been anticipated, in this 1954 count, young
trees were more numerous than they had been on any of the areas
sampled in 1952. However, the data for 1952 and 1954 are not entirely
comparable, because in 1952 none of the plots sampled was nearer than
50 feet to the edge of the woods, whereas in 1954, the sample was
arranged to be representative of the entire field, including the parts
adjacent to the woods. The numbers per acre of each kind of tree, and
the percentages that were dead or dying, were as follows: crab-apple
167 (33.5 per cent dead); locust 98 (3 per cent dead); elm 69.5 (2.9
per cent dead); osage orange 63.5 (none dead); walnut 36.5 (4.1 per
cent dead); red haw 25.5 (none dead); ash 19.5 (none dead); cockspur
thorn 17 (17.6 per cent dead); wild plum 14 (3.6 per cent dead);
dogwood 9.5 (none dead); prickly ash 2 (25 per cent dead); black oak
1.5 (none dead); boxelder .5 (none dead). Thus, of the species that
were prominent invaders of the field, only crab-apple showed heavy
mortality. In many instances the mortality in crab-apple was due
wholly or in part to attack by cottontails (_Sylvilagus floridanus_),
which had completely girdled many of the stems. In general, mortality
in the young trees was light in this grassland area compared with the
mortality in any part of the woodland.

Competition and Mortality

The ratios of trees of different species and different size groups
reflect, to some extent, the changes to which the area has been
subjected. Under original conditions mature trees of oak and hickory
dominated the forest. With the opening up of the forest that resulted
from cutting most of these mature trees, other kinds of trees
increased and spread. Species relatively intolerant of shading became
established. Chinquapin oak, honey locust, osage orange, cherry,
dogwood, red haw, and crab-apple, being especially intolerant of
shading, cannot grow in close competition with climax species, and
they become established only in fairly open situations. Their presence
in thick woodland, along with climax competitors, usually is an
indication that the woodland is either of recent origin or has been
much disturbed in the past, permitting invasion by them.

About 1934 when approximately half of the Reservation, including
nearly all the woodland areas, was fenced against livestock, shrubs
and young trees sprang up in great abundance, especially in more open
woodland situations, and at the edge of the forest. Sumac (_Rhus
glabra_) often dominated at first in such situations. Crab-apple, wild
plum, red haw, chinquapin oak, prickly ash, dogwood, honey locust, and
redbud also soon came into prominence. By 1954 thickets had grown up
and the intense competition had killed much of the woody vegetation.
Sumac, especially, had been almost entirely killed out by the shading.
By then, however, the adjacent fields had been protected for eight
years from grazing, and sparse sumac thickets were present on the
field sides of the fences, the average sizes of the plants
progressively declining farther from the edge of the woods. Much
mortality had occurred also in all the other species mentioned, with
only a few of the larger surviving in competition with elm, hackberry,
ash and osage orange, and with reproduction practically stopped except
near the edges of the thickets.

In 1954, after approximately 20 years of protection from livestock,
the woodland had become much denser, with a thick understory of
saplings and tall shrubs in most places. From a time soon after
protection was initiated, there was little or no reproduction (except
where the woodland originally was open) in blackjack oak, dwarf or
chinquapin oak, red haw, honey locust, and osage orange. On one south
slope, an open woods with well scattered trees of black oak, American
elm, hackberry, honey locust and osage orange, had by 1954 become so
dense that it was almost impassable except with the aid of a brush
knife to cut or break through the thickets. Saplings of honey locust
made up an important part of the understory vegetation on this slope.
Those of the smallest size group, up to 1½ inches stem diameter,
were mostly dead; in a strip 900 feet long and 50 feet wide there were
29 dead saplings and ten live ones of this size group. In the next
largest size group, up to 2½ inches in stem diameter, there were 17
dead and 53 live saplings, while in the size group 2½ to 3½
inches stem diameter, there was one dead sapling and 51 were alive.

On another south slope, which had more large and medium-sized trees
and less dense underbrush, 233 saplings six inches or less in stem
diameter, counted on a sample strip 530 feet long and 40 feet wide,
included elm 37.3%, dogwood 19.7%, hackberry 16.4%, coffee-tree 15.6%,
honey locust 11.0%, plum 10.3%, chestnut oak 5.5%, crab-apple 3.4%,
osage orange 2.1%, red haw 1.4%, hickory, redbud, mulberry and
cockspur thorn each .7%. There was substantial mortality in the
saplings of several of these species; plum 86.5%, dogwood 69.5%, elm
49.5%, locust 31.2%, chestnut oak 25.0%, coffee-tree 4.4%.

By 1954 several areas of hilltop-edge and north slope, which
presumably had been wooded originally, but which had been subjected to
heavy cutting, supported thriving stands of young hickories mostly two
to six inches in trunk diameter. Most of these saplings seemed to have
originated as stump-or root-sprouts. These numerous and closely spaced
saplings produced a dense and almost continuous leaf canopy, shading
and killing out many of the smaller trees of their own species as well
as competing elms, redbuds, dogwoods, hackberries and others.

On a north slope in the southeastern part of the Reservation, many
large stumps were found in late stages of decay, cut from 20 to 30 or
more years before. Insofar as could be determined, these old stumps
were mostly of oaks, but in 1954 the trees growing on this slope were
chiefly elms and coffee-trees less than one foot in diameter.

Effects of Livestock

Livestock importantly affected the trend of succession. The tendency
of grazing animals to hold back the forest by stripping the foliage
from young trees and killing them is selective, however; the several
kinds of trees differ in their tolerance to browsing and in their
palatability to animals. The kind of animal and the season and
intensity of use also have important bearing on the ultimate effect.
Several kinds of shrubs and small trees seem to be especially
susceptible to damage by browsing; chinquapin oak, crab-apple, plum,
hazel, dogwood, prickly ash, and paw paw were found to be either
absent entirely from the parts of the woodland that were heavily used
by stock, or much scarcer than they were on adjacent unbrowsed areas.
Some woody plants that are even more susceptible may have been
completely eliminated by browsing.

In the thirties when most of the woodland area was fenced off and
protected from grazing, three wooded hillside areas of a few acres
each, were maintained as connecting strips between the pastures of the
hilltops and those of the bottomlands. These areas were utilized only
at certain seasons, but by 1948 the effect of trampling and heavy
browsing by livestock was conspicuous. Herbaceous ground vegetation
was almost lacking and low woody vegetation was also scarce, in
contrast to the parts of the woodland that were adjacent but separated
by fences that excluded livestock. The contrast was perhaps heightened
along the fences because the animals tended to follow along the fence
lines and their effects were concentrated there.

    Table 4.--Numbers of Young Trees of Various Kinds and Sizes in
    1954 on a .919-acre Area Consisting of Six Hillside Strips Each
    20 Feet Wide. Each Strip Was Equally Divided by a Fence Line,
    Excluding Livestock from One Side During the Period 1934
    (Approximately) to 1948.

                  |Less than ½-inch | ½-inch to 4-inch|5-inch to 12-inch
                  |  stem diameter  | stem diameter   | stem diameter
                  |       | Percent-|       | Percent-|       | Percent-
                  | Total |  age in |Total  |  age in | Total |  age in
                  |number | browsed |number | browsed |number | browsed
                  |       |   half  |       |   half  |       |   half
                  |       |         |       |         |       |
  Dogwood         | 556   |   52.1  | 1058  |   16.4  |       |
  Redbud          |  40   |   42.5  |  102  |    5.9  |       |
  Elm             |  30   |   76.7  |  189  |   27.6  |  99   |  47.5
  Hackberry       | 131   |   39.7  |  206  |   13.1  |   5   |  20.0
  Plum            |  26   |   77.0  |   35  |   22.8  |   1   | 100.0
  Crab-apple      |  11   |  100.0  |   46  |   37.0  |       |
  Red haw         |   1   |  100.0  |   33  |   48.5  |   9   |  75.8
  Walnut          |   7   |   28.6  |   32  |   43.7  |  26   |  61.5
  Honey locust    |   2   |  100.0  |   20  |   15.0  |  11   |  27.3
  Osage orange    |   1   |  100.0  |    7  |   57.1  |   2   |  50.0
  Shagbark hickory|   3   |  100.0  |   42  |   73.8  |  44   |  40.9
  Chestnut oak    |       |         |   26  |   30.8  |  24   |  58.2
  Chinquapin oak  |       |         |   12  |  100.0  |   1   | 100.0
  Coffee-tree     |       |         |   11  |   18.1  |   8   |  12.5
  Ailanthus       |   6   |   33.3  |   65  |   26.1  |   3   | 100.0
  Black oak       |       |         |    5  |   40.0  |   7   |  16.6
  American ash    |  21   |  100.0  |    3  |   33.3  |       |
  Paw paw         |  12   |         |   61  |   27.8  |       |

In 1954 ten-foot wide strips were sampled on both sides of the fences.
For both browsed and unbrowsed samples, the strips had a total length
of 4000 feet, each representing an area of .919 acres. Table 4
contrasts the number of young trees per acre on the browsed and
unbrowsed areas, grouped in several size classes. In general the
saplings up to one-fourth inch in diameter were those that had become
established in the five growing seasons since browsing was
discontinued and both areas were protected. For this size group the
numbers were approximately equal, being slightly higher on the browsed
strips. However, in the size group of ½ inch to 4 inches in stem
diameter, the trees were nearly three times as abundant on the
unbrowsed areas, and most trees within this size range must have
become established within the time of differing treatments. The
disparity in numbers was great for hackberry, redbud, elm and dogwood
which made up the bulk of the saplings. In the size range 5 to 12
inches most trees antedated the fence, and the unbrowsed portion had
only a few more than the portion that had been browsed.

On the formerly browsed areas clumps of gooseberry bushes were
conspicuous and were computed to cover 3.81 per cent of the area
sampled, versus 2.87 per cent on the unbrowsed area. These thorny
bushes seem to be resistant to browsing, and elsewhere have been noted
in abundance in woodlands heavily used by livestock. The elimination
of competing undergrowth by browsers may be a factor favoring
development of gooseberry clumps. The trend was just the opposite for
fragrant sumac, which was computed to cover 1.94 per cent of the
browsed sample versus 3.23 per cent of the unbrowsed sample.
Greenbrier (_Smilax tamnoides hispida_) was most abundant on the
unbrowsed strips, with seven large clumps, and 56 smaller clumps (10
stems or fewer) as contrasted with five large clumps and 32 smaller
clumps on the browsed strips. There were 32 grapevines (_Vitis
vulpina_) on the unbrowsed strips and only seven on those that were

Animal Associates

The invertebrates of the University of Kansas Natural History
Reservation have not been intensively studied. Most of the species of
vertebrates are characteristic of the deciduous forest of the eastern
United States, or of the edge of woodland; relatively few kinds are
characteristic of prairies.

Of birds, for example, some 23 species characteristic of the eastern
deciduous forests have been found nesting on the Reservation, as have
14 additional species that are mainly eastern in their distribution
but are most characteristic of forest-edge thickets, clearings, or
marshy places. The ruffed grouse (_Bonasa umbellus_) and wild turkey
(_Meleagris gallopavo_) are not present on the area, although they may
have occurred there earlier. Other forest birds which occur in the
general area, and which have been recorded from time to time on the
Reservation, although they seem not to nest there, are:
chuck-will's-widow (_Caprimulgus carolinensis_), scarlet tanager
(_Piranga olivacea_), Acadian flycatcher (_Empidonax virescens_),
veery (_Hylocichla fuscescens_), parula warbler (_Parula americana_),
oven-bird (_Seiurus aurocapillus_), and orchard oriole (_Icterus
spurius_). For each of these, habitat conditions on the Reservation
seem to be deficient in some respect. On the other hand, the only
typical prairie bird that breeds on the Reservation is the dickcissel
(_Spiza americana_). Others, including the Swainson hawk (_Buteo
swainsoni_), greater prairie chicken (_Tympanuchus cupido_), upland
plover (_Bartramia longicauda_), western kingbird (_Tyrannus
verticalis_) and loggerhead shrike (_Lanius ludovicianus_), occur in
the general area, and may even cross the Reservation at times, but
they do not become established.

In the mammalian fauna, species typical of the deciduous forests
include the opossum (_Didelphis marsupialis_), short-tailed shrew
(_Blarina brevicauda_), eastern mole (_Scalopus aquaticus_), eastern
gray squirrel (_Sciurus carolinensis_), and pine vole (_Microtus
pinetorum_), but the eastern chipmunk (_Tamias striatus_) and southern
flying squirrel (_Glaucomys volans_) are lacking. Also, the present
fauna lacks large mammals that may have been present under original
conditions: the white-tailed deer (_Odocoileus virginianus_), recorded
on the area from time to time but not permanently established there,
the wapiti (_Cervus americanus_), black bear (Ursus americanus), and
bobcat (_Lynx rufus_). Other species on the area, that are
characteristic of the deciduous woodlands, but that occur also far
west into prairie regions, include the little short-tailed shrew
(_Cryptotis parva_), raccoon (_Procyon lotor_), fox squirrel (_Sciurus
niger_), white-footed mouse (_Peromyscus leucopus_), eastern woodrat
(_Neotoma floridana_) and eastern cottontail. On the area, the only
mammals that are sharply confined to grasslands, elsewhere as well as
on the Reservation, are the plains pocket gopher (_Geomys bursarius_)
and plains harvest mouse (_Reithrodontomys montanus_), both of which
are rare on the area, and the hispid cotton rat (_Sigmodon hispidus_).
The following species are typical of the plains, but they range
eastward into the region of deciduous forests: western harvest mouse
(_Reithrodontomys megalotis_), deer mouse (_Peromyscus maniculatus_),
coyote (_Canis latrans_), and spotted skunk (_Spilogale putorius_).
The following mammals, typical of grassland, are absent: black-tailed
jack rabbit (_Lepus californicus_), black-tailed prairie dog (_Cynomys
ludovicianus_), 13-lined ground squirrel (_Spermophilus
tridecemlineatus_), Franklin's ground squirrel (_Spermophilus
franklinii_), southern lemming-mouse (_Synaptomys cooperi_), and of
course, the buffalo (_Bison bison_), and the prong-horned antelope
(_Antilocapra americana_) long extinct in this part of their range.

Of amphibians and reptiles also, the majority are typical forest
species, including: the American toad (_Bufo terrestris_), common tree
frog (_Hyla versicolor_), brown skink (_Lygosoma laterale_), common
five-lined skink (_Eumeces fasciatus_), worm snake (_Carphophis
amoenus_), pilot black snake (_Elaphe obsoleta_), DeKay snake
(_Storeria dekayi_), western ground snake (_Haldea valeriae_),
copperhead (_Agkistrodon contortrix_), and timber rattlesnake
(_Crotalus horridus_). Other typical forest species missing from the
area include the spring peeper (_Hyla crucifer_), Carolina box turtle
(_Terrapene carolina_), coal skink (_Eumeces anthracinus_), and
red-bellied snake (_Storeria occipitomaculata_). Of typical prairie
species only the Kansas ant-eating frog (_Gastrophryne olivacea_) and
the ornate box turtle (_Terrapene ornata_) are common, and, curiously,
each seems to prefer a forest habitat on this area, in the absence of
their closely related eastern representatives, the eastern ant-eating
frog (_G. carolinensis_) and the Carolina box turtle, respectively,
which usually live in forests. The plains spadefoot (_Spea
bombifrons_), garden toad (_Bufo woodhousii_), Great Plains skink
(_Eumeces obsoletus_), prairie skink (_Eumeces septentrionalis_),
slender tantilla (_Tantilla gracilis_), prairie rat snake (_Elaphe
guttata_), bull snake (_Pituophis catenifer_), and blotched king snake
(_Lampropeltis calligaster_) are all scarce on the area. The plains
toad (_Bufo cognatus_), collared lizard (_Crotaphytus collaris_),
except for an introduced colony, plains garter snake (_Thamnophis
radix_), lined snake (_Tropidoclonion lineatum_), and massassauga
(_Sistrurus catenatus_) seem not to occur on the area at all.

Annotated List of Species

#Juniperus virginiana.#--Red cedar, the only native gymnosperm of
northeastern Kansas, occurs in nearly all woodlands of the region,
although individual trees are widely scattered. It has increased
remarkably in the past few years. No mature cedar trees grow anywhere
on the Reservation, but young trees, probably several dozen in all,
are widely scattered in a variety of situations on the area. Probably
in every instance the seeds have reached the area in droppings of
birds. Approximately 15 miles south and a little east of the
Reservation is a stand of cedars some of which are 100 to 300 years
old. Near the southwest corner of the section, at the site of a former
farm house there is a small grove of these trees, probably planted.
These may have been the source for some of the young trees on the

On several occasions cardinals (_Richmondena cardinalis_) were
observed to have nested in the young cedars, whose thick foliage
provided well sheltered nesting sites. This shelter was utilized
especially in early nestings when foliage had only begun to appear on
other trees and shrubs. However, two such nests in cedars, that were
checked repeatedly, were eventually destroyed by predators.

#Salix nigra.#--Black willow is localized in the vicinity of the one
small pond on the Reservation. The pond was made in 1936; at the
upper end of a small valley a dirt bank 100 yards long was built
across a ravine through which an intermittent creek drained. Hilltop
fields draining into this ravine were then under cultivation. In the
next few years heavy erosion occurred in the upland fields, and the
soil carried downstream was deposited in the pond. Most of the pond
was filled up with a silt flat about an acre in area. On the higher
part of this silt flat a dense thicket of saplings of elm, honey
locust and osage orange sprang up. On the lower, wetter part of the
silt bar a willow grove grew up, dominated by _S. nigra_, with _S.
eriocephala_, _S. interior_ and _S. amygdaloides_ in smaller numbers.
By 1955 some of these trees had attained a trunk diameter of eight
inches and a height of thirty feet. Elsewhere on the Reservation,
willow is represented only by a few scattered trees and bushes along
the two intermittent creeks. The silty soil preferred by the willow is
scarce as both streams are actively eroding their channels.

The moist, silty soil beneath the willow grove is covered with a dense
mat of low vegetation including giant ragweed, carpenter's square,
dayflower, and rice cutgrass. Short-tailed shrews, house mice (_Mus
musculus_), harvest mice and cotton rats thrive in this habitat.
Red-winged blackbirds (_Agelaius phoeniceus_), yellow-billed cuckoos
(_Coccyzus americanus_), red-eyed vireos (_Vireo olivaceus_), catbirds
(_Dumetella carolinensis_) and Kentucky warblers (_Oporornis
formosus_) use it for nesting. The high humidity and dense vegetation
in this grove render it favorable habitat for recently metamorphosed
frogs and toads, especially the tree frog, which is sometimes
extremely abundant there in summer.

#Populus deltoides.#--Cottonwood is one of the less common trees on
the area, but it attains a larger size than any of the other kinds.
The larger of the two creeks on the Reservation is lined with mature
cottonwoods along the lower part of its course. Along the smaller
creek large cottonwoods are also present but they are more widely
spaced. A few cottonwoods are present at well scattered points on
slopes and hilltops, usually in forest edge situations or in woodland
where other trees are sparse. By far the largest tree on the
Reservation is a cottonwood of 15-foot circumference (Plate 1),
growing on a hilltop near the south boundary of the Reservation, at
the edge of woodland adjacent to a cultivated field.

The heavy rainfall of 1951 resulted in the establishment of hundreds
of cottonwood seedlings, mostly in places remote from the mature
trees. So far as observed, all these were in recent silt deposits.
Many of them have survived the drought of 1952-1954.

Because of their great height, towering above the level of the
surrounding tree-tops, cottonwoods are preferred look-out perches
of certain of the larger birds, notably red-tailed hawks (_Buteo
jamaicensis_), barred owls (_Strix varia_), and crows (_Corvus
brachyrhynchos_). Flocks of robins (_Turdus migratorius_) and of rusty
blackbirds (_Euphagus carolinus_) preparing to roost have been noted
habitually to gather in the tops of tall cottonwoods. In spring,
large wandering flocks of goldfinches (_Spinus tristis_) have been
seen feeding on the leaf buds of cottonwoods. Baltimore orioles
(_Icterus galbula_) and yellow-billed cuckoos often forage in
cottonwoods. Red-bellied woodpeckers (_Centurus carolinus_) spend a
disproportionately large amount of their time in cottonwoods. These
woodpeckers have been observed nesting in the hollow branches on several
occasions. Downy woodpeckers (_Dendrocopos pubescens_) also have been
noticed foraging in cottonwoods on many occasions. Certain large
isolated cottonwoods along creeks were favorite stopping places of blue
jays (_Cyanocitta cristata_) which, on trips from one wooded hillside to
another, usually perched briefly in the tops of these tall trees.
Calling and looking about, the jays seemed to maintain contact with
distant mates or members of the flocks by using these high perches.
Often after a brief pause in the top of the cottonwood they flew off in
a new direction.

Both woodrats and opossums have been known to utilize hollow
cottonwoods as dens. Fox squirrels have been seen climbing in
cottonwoods occasionally.

#Juglans nigra.#--Black walnut is one of the more prominent hardwoods.
Under original conditions, evidently many of the larger trees were of
this species. Being the most valuable timber species of the area,
walnut has been subjected to heavy cutting over the past 85 years.
Most of the walnut trees still present are small or medium-sized, but
the species is still abundant over much of the area. Along certain
hilltop edges there are groves of walnuts, growing in nearly pure
stands, with an occasional elm, ash, coffee-tree or honey locust.
Elsewhere walnut trees are more scattered, but are distributed
throughout the woodland. Although the walnut trees growing in woods
are of various sizes from those of mature size down to saplings,
seedlings are to be found mainly in fields near the woodland edge. In
these situations it is one of the more prominent of the woody species
invading open lands. The seeds evidently are transported mainly by
rodents, especially fox squirrels.

In autumn every walnut tree that is bearing nuts becomes a focal
point of activity for squirrels. Over a period of weeks the squirrels
concentrate their attention on the walnut crop, continuing until
virtually every nut has been harvested. Walnut seems to be the one
most important food source, for both the fox squirrel and the gray
squirrel. Most of the nuts are stored for future use. Many buried
separately and never retrieved by the squirrels, grow into new trees.

White-footed mice often store the nuts in their nests, in burrows,
beneath rocks or in crevices. In summer, groves and isolated trees of
walnuts are favorite haunts of the yellow-billed cuckoo, which finds
concealment in the thick foliage, and probably feeds upon the tent
caterpillars that commonly infest these trees.

#Carya ovata.#--Shagbark hickory is one of the more important
hardwoods of the area. The trees are relatively small compared with
the larger oaks, elms, ashes and hackberry. However, on several parts
of the area this hickory is dominant. It grows mainly on north slopes
and hilltops. The trees most frequently associated with it are black
oak, American elm and chestnut oak. Scattered through the woodlands
are occasional mature hickories of DBH 18 inches or more. However,
many of the trees are six inches or less DBH and a large proportion of
these have originated as stump sprouts from trees cut in the early
thirties or before.

Shagbark is especially tolerant of shading. Numerous young trees and
seedlings noted all were growing in dense woods of larger hickories,
oaks, or mature elms. None has been found in open fields or even in
edge situations. This hickory is resistant to drought; relatively few
died during the drought of 1952-1954, and these were mostly small
trees in crowded stands.

In parts of the woodland dominated by shagbark hickory the trees are
mostly 5 to 6 inches or even smaller in trunk diameter and 20 to 30
feet high, sometimes growing in nearly pure stands, and with a leaf
canopy so dense that shrubs and herbaceous vegetation are sparse.

The mast crop produced by shagbark is an important food source for
both fox squirrels and gray squirrels. Both kinds of squirrels often
use these hickories as sites for their stick nests. White-footed mice
also store the nuts as a winter food source.

Birds which are most often seen in groves of shagbark include the
yellow-billed cuckoo, tufted titmouse (_Parus bicolor_), black-capped
chickadee (_P. atricapillus_), blue jay, summer tanager (_Piranga
rubra_), and red-eyed vireo. The Cooper hawk (_Accipiter cooperii_)
has been recorded nesting in this hickory. In dead trees of this
species that are still standing, the interiors may decay more rapidly
than the armorlike bark plates. On several occasions tufted titmice
and chickadees have been recorded as nesting in such cavities.

#Quercus stellata.#--Post oak is relatively scarce on the Reservation.
One area of approximately an acre on a south slope is dominated by it.
There are several other small groves and scattered trees. All are on
moderately steep south slopes in poor soil. Trees often found
associated with it include red elm, chestnut oak, chinquapin oak,
blackjack oak, hickory, and dogwood. It seems likely that under
original conditions this species occupied about the same area as it
does at present. It is not spreading, and there are few young trees
anywhere on the area. In every instance the groves are limited to a
rocky clay soil, and edaphic factors obviously are of major
importance. Under original conditions fire was probably a limiting
factor, and at the present time competition with other hardwoods may
be even more important.

#Quercus macrocarpa.#--Less than a dozen individuals of mossycup oak
have been noticed on the area, at well scattered points. Under
original conditions, it probably grew chiefly in the bottomlands that
have been completely cleared of timber for cultivation. The few now
present are all on hillsides, and are medium to large trees.

#Quercus Muehlenbergii.#--Chestnut oak was perhaps the one most
important tree species of the original climax forest on the area.
Because of its slow growth, scanty seed production, and large heavy
fruits with seeds lacking effective dispersal mechanisms, it has lost
ground to other kinds of trees as a result of the unnatural
disturbances which have occurred.

It still dominates on rocky upper slopes that have north, east or west
exposures and forms nearly pure stands in limited areas. Nearly all
the larger trees of this species now present have been cut one or more
times and have regenerated from stump sprouts. Seedlings and young
saplings of this oak are scarce even in parts of the woodland where
the species is most common. It is evident that reproduction is slow,
at least under present conditions. On the lower hill slopes these oaks
are scarce and scattered, but some of the largest are in such
situations. Chestnut oak seems to be relatively resistant to drought.
In the summer of 1954 when elms, and especially black oaks of all
sizes were dying in large numbers, the chestnut oaks growing among
them showed little evidence of injury in mature trees and only a small
percentage of mortality in saplings.

  [Illustration: Fig. 5. Map of Reservation showing present
      distribution of chestnut oak (shaded). The species is not
      spreading  and is thought to be largely confined to the area
      that was wooded  before 1860. Except in minor details,
      shagbark hickory conforms to  the same distribution pattern
      on this area.]

Chestnut oak has a relatively slow growth rate. In 17 that were
recorded, there were, on the average, 4.59 annual rings per inch of
trunk diameter. Near Pigeon Lake, Miami County, Kansas, counts were
obtained from five cut in 1952 from a virgin stand in a habitat
similar to that on the Reservation. The five trees had trunk diameters
of 16½ to 25 inches and ranged in age from 65 to 183 years. Several
still growing on the Reservation are larger and presumably are well
over 100 years old.

As this oak seems to be in process of being replaced by other trees,
is slow-growing, and slow in dispersal, it seems probable that the
areas now occupied by its stands supported stands of it under
original conditions. Whether it can regain dominance under present
conditions of protection from cutting, fire and grazing remains to be

The chestnut oak produces a mast crop which is utilized by many kinds
of animals. Fox squirrels, gray squirrels, and white-footed mice feed
upon the acorns and store them. Blue jays, red-headed woodpeckers
(_Melanerpes erythrocephalus_), and red-bellied woodpeckers also eat
them. The red-eyed vireo, summer tanager and tufted titmouse are among
the birds that most frequently forage for insect food in chestnut
oaks. Relatively few kinds of birds seem to use this tree as a nest

#Quercus prinoides.#--The chinquapin oak on this area is a small
shrubby tree, usually not more than 15 feet high and more typically
only six to eight feet. It occurs chiefly in dry rocky situations
along hilltop edges and upper slopes, usually where the slope exposure
is at least partly to the south. In such situations it may grow in
nearly pure stands. Often it is associated with dogwood. The trunks
are usually two to four inches in diameter, gnarled and twisted. The
crowns are dense and spreading.

This oak is the dominant plant in certain small areas of its preferred
habitat. In other areas of hilltop edge and upper slope it is being
eliminated by stands of hickory, chestnut oak, black oak and elm,
which shade it out. The species is tolerant of moderate to heavy
browsing, but seemingly can be eliminated by more intensive
utilization; even the higher foliage is often within reach of
livestock. In "Horse Woods" one of the hillside areas that was open to
livestock until 1949, this oak was almost absent, but it was abundant
in adjoining parts of the woods that were fenced in the thirties to
exclude livestock.

The thickets formed by this shrubby oak are frequented by cottontails,
which feed upon the bark and foliage. The small acorns are used as
food by rodents, especially the white-footed mouse. On several
occasions, in winter, groups of long-eared owls (_Asio otus_) have
been found roosting in thickets of chinquapin oak. Crows also utilize
these thickets for roosting occasionally. The white-eyed vireo (_Vireo
griseus_), gnatcatcher (_Polioptila caerulea_), and tufted titmouse,
frequent the oak thickets.

#Quercus rubra.#--The red oak is one of the important climax species
of the area. At present it is largely confined to a ravine in the
northeastern part of the section. The woodland here is less disturbed
than on most other parts of the Reservation, and red oak is the
dominant species. There are large trees, rather evenly distributed,
growing on east-facing and west-facing slopes. Just east of the
Reservation, in the "Wall Creek" area, the small valley on either side
of the creek and the adjacent lower slopes are dominated by giant red
oaks larger than any now growing on the Reservation. Farther up the
slope in the area of limestone outcrops, dominance shifts to chestnut
oak. That red oaks of similar size, and even larger, formerly occurred
on the Reservation, at least in the area still dominated by the
species, is shown by the presence of a stump 49 inches in diameter,
now in an advanced state of decay.

  [Illustration: Fig. 6. Map of Reservation showing present
      distribution of black oak (smaller dots) and red oak (larger
      dots). Neither species is spreading and both are thought to
      be largely  confined to the area that was wooded before 1860.]

The large acorns of the red oak are a favorite food of the gray
squirrel, which is most numerous on the parts of the Reservation where
these trees are present. The red-headed woodpecker on the area tends
to concentrate its activities where there are red oaks. The fox
squirrel, white-footed mouse, and blue jay are important consumers of
the acorns of red oak. A pair of barred owls resided in the deep woods
formed by these oaks and the associated trees.

#Quercus velutina.#--Black oak is one of the dominant species of the
original forest climax, and is still one of the more important trees
of the woodland. Like chestnut oak it shows little tendency to spread
beyond its present limits. Wherever there are small trees there are
old mature trees or remains of them nearby. For this reason the
present distribution of black oak on the area is thought to fall
entirely within the area occupied by the original forest. At present
it occurs throughout most of the woodland except in the warmer and
drier situations, such as on south slopes. In some hilltop situations
it is common, with occasional large mature trees. In some parts of the
bottomland and lower slopes it is abundant also, but there are
scarcely any on the upper dry rocky slopes that are the preferred
habitat of chestnut oak.

Growth in the black oak is somewhat more rapid than in the chestnut
oak, as the black oak usually grows on better soil. For 15 the average
growth amounted to 3.21 annual rings per inch of trunk diameter.

In 1954 a study of annual rings in a large, long dead, black oak at
the bottom of a north slope near the Reservation headquarters showed
that the tree was 96 years old, and hence was growing before the area
was settled. Within the period of this study black oak underwent
reduction in numbers more severe than that noted in any other species
of tree on the Reservation. The effect of drought may have been the
primary factor, although undoubtedly disease was involved also. In
1953, the second successive drought year, mortality was noticeable.
Precipitation continued below normal until August 1954. By then the
oaks had been decimated. On a sample strip of hilltop where 29 were
recorded, 21 had recently succumbed, and their leaves were dry and
withered; two were dying, though still having some green foliage, and
only six were surviving, all evidently in critical condition. The
mortality included trees of all sizes, even the largest and oldest. No
further mortality was noted in 1955 when precipitation was only
slightly below normal. On the Reservation there are many old logs, and
snags still standing, of mature black oaks long dead. Earlier drought
periods such as those of 1936-37 and 1925-26 possibly were also times
of unusually heavy mortality. In any case it seems clear that this oak
was originally more prominent in the woodlands than it is at present,
and has been steadily losing ground. Even where the mature trees
remain in greatest numbers the saplings are relatively scarce as
compared with those of elm, ash, hackberry, and hickory. The
westernmost limits of the range are nearly 100 miles west of the

Black oak provides a mast crop which is utilized by various small
mammals, notably squirrels and white-footed mice. Gray squirrels have
often been noticed in or about these trees. Hairy woodpeckers
(_Dendrocopos villosus_), black and white warblers (_Mniotilta
varia_), and brown creepers (_Certhia familiaris_) have often been
noticed foraging on the trunks. Blue jays, myrtle warblers (_Dendroica
coronata_), tufted titmice, and summer tanagers frequently forage
through the crowns. Often black oak trunks are hollow and the cavities
are utilized by various birds and mammals including the screech owl
(_Otus asio_), barred owl, raccoon, opossum, fox squirrel, gray
squirrel, woodrat, and white-footed mouse.

#Quercus marilandica.#--Black Jack oak is localized in four small
compact groves on the Reservation. These sites, though well separated,
are similar. All are on steep lower slopes, where there is dry rocky
clay soil and the exposure is mainly south. Probably all four groves
date back to the time when the area was still in an undisturbed state.
Originally they were perhaps largely separated from the remainder of
the woodland. Black Jack oak is more tolerant of heat and drought than
most of the other hardwoods are. The species is intolerant of fire,
but perhaps was partly protected under original conditions by the
sparseness of herbaceous vegetation on the poor soil where the groves
were situated.

These oaks are relatively slow-growing. One stump of 9-inch diameter,
typical of the larger Black Jack trees, had approximately 60 annual
rings. Under present conditions there is little or no reproduction and
these trees are dying out as a result of competition by other
hardwoods. Under protection from fire and browsing, elms, other oaks,
locust and dogwood have closed in about the groves and seem to be
shading them out.

There are several mature oaks of anomalous appearance, in different
places within a few hundred feet at most of the groves of Black Jack.
Most of these appear to be hybrids between the present species and _Q.
velutina_, as they are somewhat intermediate in size, bark texture,
and leaves.

This oak produces a mast crop used by various birds and mammals, and
groves are frequented by blue jays, fox squirrels, white-footed mice
and woodrats. In the mid-forties when the woodrat population was high,
there were many of the rats' stick houses in the groves, built either
at the bases of the trunks or among the dense branchlets in tops of
fallen trees. By 1952 the population of woodrats was much reduced and
had disappeared entirely from these groves. The houses were collapsed
and decaying.

Horned owls (_Bubo virginianus_) and barred owls often make their day
roosts among the dense interlacing twigs of these trees, and
red-tailed hawks have been known to roost for the night in the same
kinds of situations.

#Ulmus americana.#--On most parts of the area American elm is the
dominant tree. It occurs throughout the woodland, and most of the
larger trees are of this species. In each of the fields that were
formerly cultivated, and in the pasture areas, there are many
saplings. More than one hundred elms of DBH two feet or more have been
recorded. Presumably these mostly date back 90 years or more and were
already growing on the area when it was relatively undisturbed. On the
area the distribution of these large elms corresponds in a general way
with the present distribution of the oak-hickory type. The coinciding
distribution of the climax species and of the largest trees is
believed to reflect the distribution pattern of the original forest,
except that clearing was thorough in the bottomlands so that hardly
any trees of the climax species, or large trees of any kind remain.
Several elms of three feet or more DBH were recorded, and the largest
one measured was 46 inches. The largest elms are in alluvial soil near
small creeks in the two valleys. Also many large elms grow along the
upper slopes, especially along the outcrops of the two main strata of
the Oread Limestone. Such sites along the outcrops on open slopes are
the first to be invaded. The rock strata are relatively impervious to
water, which is held at a depth where it is readily available to the
trees. Along rocky upper slopes between the two outcrops, where
chestnut oak is abundant, elms are relatively scarce and seem unable
to compete successfully. It is noteworthy that elm is not mentioned in
several of the descriptions (Taft, 1950; Parks, 1854; Robinson, 1899)
of the original forest, even in listings of the species present. It
must have been much less prominent until favored by disturbed

  [Illustration: Fig. 7. Map of Reservation showing present
      distribution of the largest American elms, those more than two
      feet in trunk diameter. American elm is increasing and spreading
      on the area, and smaller trees are abundant even in former
      cultivated fields and pastures. Growth rate varies according to
      site, but these larger trees are, in many instances, 90 years or
      more in age and most of them are thought to be in the area
      wooded in the eighteen sixties and before.]

In July and August, 1954, a large proportion of the elms on the area
died. The die-off included trees of all sizes, and evidently the
cumulative effect of drought in 1952 and 1953, continuing into the
spring and summer of 1954, was the primary cause, although
diseases such as phloem necrosis, and insect infestations, may have
intensified its effect. In August of 1954 the bare dead elms stood out
conspicuously in the mass of green foliage surrounding them. Most of
them had survived the two dry summers of 1952 and 1953 with little
evident loss in vitality. However, the continued lack of moisture as
the 1954 growing season progressed, and the extremely hot weather of
June and July caused heavy mortality. In the course of a few days the
foliage of the upper branches would wither, die and turn brown. In
some instances numerous sucker shoots grew from the trunk of the tree
as the top was dying. Mortality was especially heavy on south-facing
slopes. Certain ecologists believe that over the years, as trees
deplete subsoil moisture and periodic droughts make their effects
felt, other species also will die off and eventually prairie will
replace them where the present forests are growing in dry and exposed

Infestations of the introduced bark beetle, _Scolytus multistriatus_,
were common and probably contributed to death of many elms. In the
winter of 1953-54 before much mortality had occurred, the bark beetle
infestations had become conspicuous. Especially on south slopes elms
of about six inches DBH were heavily infested. Woodpeckers, including
the downy, hairy, and red-bellied, habitually resorted to the elm
trunks to forage. As a result of their activities chips of bark
accumulated sometimes to a depth of several inches around the bases of
the trunks, and the exposed inner layers of brown bark caused the
infested trees to contrast with the predominantly gray color of those
that were still healthy and retained the outer layer of bark.

In April and early May seeds of the American elm constitute a major
food source for birds, including the black-capped chickadee, tufted
titmouse, junco (_Junco hyemalis_), red-eyed towhee (_Pipilo
erythrophthalmus_), Harris sparrow (_Zonotrichia querula_), cardinal,
goldfinch, tree sparrow (_Spizella arborea_) and field sparrow (_S.
pusilla_). Birds recorded as nesting in the American elm include the
mourning dove (_Zenaidura macroura_), Cooper hawk, red-tailed hawk,
broad-winged hawk (_Buteo platypterus_), turkey vulture (_Cathartes
aura_), screech owl, horned owl, barred owl, red-bellied woodpecker,
downy woodpecker, tufted titmouse, black-capped chickadee,
gnatcatcher, red-eyed vireo, summer tanager, indigo bunting
(_Passerina cyanea_), field sparrow and cardinal.

Opossums, raccoons, fox squirrels and white-footed mice often live in
cavities in elms.

Insectivorous birds that find their food on foliage and prefer elm or
use it to a large extent are: yellow-billed cuckoo, tufted titmouse,
black-capped chickadee, blue-gray gnatcatcher, red-eyed vireo,
white-eyed vireo and warblers, including the myrtle, Audubon
(_Dendroica auduboni_), yellow (_D. petechia_), black-throated green
(_D. virens_), black-poll (_D. striata_), Tennessee (_Vermivora
peregrina_), orange-crowned (_V. celata_), Nashville (_V.
ruficapilla_) and American redstart (_Setophaga ruticilla_).

#Ulmus rubra.#--The red elm (or slippery elm) is widely distributed
over the area, but only a few trees with a trunk diameter of twelve
inches or more are present. Throughout the woodlands of the
Reservation the saplings of this species constitute a prominent part
of the understory. However, few survive beyond the sapling stage. The
red elm is never abundant in Kansas woodlands. It is intolerant of
drought conditions, and is one of the first trees to die. This fact
probably explains the scarcity of mature trees of this species on the

#Celtis occidentalis.#--Hackberry is widely distributed on the area,
but is not dominant anywhere. Its favorite site is along hilltop
limestone outcrops, especially where there is south exposure. There
are few on hilltops away from the outcrops. Hackberries are scattered
in small numbers over the wooded slopes. There are a few of unusually
large size, along edges of the bottomlands. Hackberries are
slow-growing. Counts of annual rings for four indicated an average of
7.1 rings per inch of trunk diameter. Young hackberries of all sizes
are numerous throughout the woodland. Therefore it seems likely that
this species is in process of spreading and probably has already
extended beyond the situations which it originally occupied.

The fruits of hackberry provide a fall and winter food supply for
various animals. Opossums are especially fond of them. Red-bellied
woodpeckers have been seen storing them. Migrating flocks of robins
may utilize them as a major food source temporarily. White-footed mice
and woodrats store them and eat them.

#Morus rubra.#--Red mulberry is moderately common in certain heavily
wooded areas, especially the lower parts of north slopes. A few are
present on wooded hilltops. Most of the trees are between ten and
twenty feet tall, and generally die before growing larger. Red
mulberry is present in most woodlands of eastern Kansas and is
seemingly distributed by birds. It is never an important component of
woodlands in the area. Catbirds (_Dumetella carolinensis_) and wood
thrushes (_Hylocichla mustelina_) especially have been noted
frequenting the vicinity of mulberry trees in fruit. Probably many
other kinds of birds utilize the fruits to some extent.

#Maclura pomifera.#--Osage orange was not a member of the original
flora, but early settlers in Kansas valued it for windbreaks and fence
posts, and they made extensive plantings. Presumably it was introduced
onto the area of the present study in the eighteen sixties. At the
present time it occurs throughout the woodland, with scattered mature
trees and many young trees on the former pastures. This aggressive
invader spread despite frequent cutting, and now plays an important
part in the ecology of the area. Most of the larger trees have been
cut one or more times, but have regenerated from stump sprouts with
multiple stems and spreading habit. The tough and durable wood is
useful for fence posts. The growth rate is slow, similar to that of
oaks and elms.

Osage orange is intolerant of fire and is easily killed by scorching.
It is damaged by browsing, and cannot grow in deep shade. It is
drought resistant. Mortality was light during the drought period of
1952-1954, although many of the trees were growing on poor soil in the
hotter and drier sites.

Where there are stands of mixed hardwoods, osage orange is relatively
scarce and tends to be on or near the edges of the stands. The osage
orange trees growing in competition with oaks, elms and hickories may
have tall, slender trunks and narrow crowns, in contrast with the
spreading habit of those growing in more open sites. In the woodlands
small and medium-sized trees are scarce and there is hardly any
reproduction. Obviously the osage orange, like honey locust became
established in the forests when the stands were more open, probably
after cutting of the large trees. In contrast to the meager
reproduction in shaded sites is the abundant crop of young saplings
along edges of fields adjacent to woods or about isolated osage orange
trees. Evidently the tree does not become established readily on
bluestem prairie. On a hillside adjoining the northwest corner of the
Reservation, long subjected to heavy grazing, osage orange dominates,
but just across the fence on the Reservation side, it is almost
absent. This area had been maintained as bluestem prairie until about
1934 by occasional burning and since then had partly grown up into
thickets in which dogwood, and saplings of elm and hackberry were

The dense thorny branches provide shelter and nesting sites for many
kinds of animals. On this area the cardinal utilizes it for nesting
sites more frequently than any other kind of tree. Some nests were so
well protected by the thorns that they could scarcely be reached.
Indigo buntings, field sparrows, and yellow-billed cuckoos also use
these trees or young saplings for nesting sites.

In the forties, when the woodrat was common on the area, its local
distribution seemed to be determined mainly by the osage orange. Many
houses of the woodrat were built around old stumps at the bases of
large, spreading osage orange trees. Frequently the houses were in the
main crotch of a tree two to eight feet from the ground.
Characteristically the rats used horizontal or gently inclined, low
branches of the tree as runways to and from the house. In summer and
early autumn these rats stored foliage of the osage orange in large
quantities in chambers adjacent to the nest. The seeds also provided
an important food source. During the period 1948 to 1951 the woodrat
population steadily decreased, and one by one the houses in osage
orange trees were deserted, until the small surviving population of
woodrats was limited to hilltop rock outcrops not associated with
osage orange trees.

The seeds are well liked by other rodents also. In late fall and
winter after the "hedge balls" have fallen, fox squirrels visit the
trees and shred the fruits to gain access to the seeds. Over periods
of weeks heaps of the shredded refuse accumulate at the base of the
tree trunk. The seeds probably constitute the one most important
winter food of the fox squirrel. The tufted titmouse also relies to a
large extent on the seeds for its winter food. Being unable to shred
the bulky hedge balls itself, it depends almost entirely on the seeds
in fruits torn open by the squirrel but not fully utilized by it. At
times when the ground and trees are snow-covered, making unavailable
most other food sources, the osage orange seeds gleaned from refuse
heaps in the sheltered feeding places of the squirrels are probably of
critical importance to the titmouse.

The cottontail and white-footed mouse also eat the seeds.

#Platanus occidentalis.#--Sycamores are few and scattered on the area,
but those present seem to be holding their own if not gaining in
numbers. They include some of the largest trees on the Reservation.
The most typical habitat is along rocky ravines on wooded slopes.
Occasional trees are scattered through the woods away from ravines on
slopes of north, east, or west exposures, or on hilltop edges,
providing strong evidence that these areas were more open at the time
the sycamore seedlings became established. Cutting of the mature trees
in the original forest and subsequent grazing might have created the
conditions favorable for their establishment. Many saplings have
sprung up in the fallow hilltop fields that were formerly cultivated.

Many of the larger sycamores have cavities and these are inhabited by
various animals. A large sycamore in a ravine below a pond had a
cavity in its base within which a raccoon reared its litter of young
one summer. At other times this same cavity was inhabited by woodrats
and by fox squirrels. Seemingly this cavity was the habitat of a
certain chigger which was found on both the squirrels and the woodrat.
Red-bellied woodpeckers excavated a cavity high on this same tree
trunk, in which they reared their brood.

Several large sycamores died as a result of the cumulative effect of
drought in the summers of 1952, 1953 and 1954, but many others

#Prunus americana.#--Wild plum is a small tree, usually not more than
three inches in trunk diameter, nor more than twelve feet high. It
tends to grow in dense thickets which are spotty in distribution.
Several of these thickets are in edges of former pastures at the
woodland edge. Other extensive thickets are in the following
situations: along hilltop rock ledges and encroaching into adjacent
prairie on upper south-facing slope maintained as bluestem prairie by
mowing and burning, until 1934; along a ravine in formerly cultivated
hilltop fields; along tops of steep creek banks at edge of old corn
field. In a few situations within the woodland there are dead and
dying thickets of wild plum, shaded out by the closing in of the tree
canopy, as fast-growing trees such as elm, honey locust, and cherry
sprang up in former clearings.

The woodrat lived in several plum thickets that provided the type of
shelter from predators that it requires. The bark, fruit and foliage
are used as food. In autumn the plums sometimes are the chief food of
the opossum. Plum thickets provide the preferred habitat for the Bell
vireo (_Vireo bellii_). The white-eyed vireo, field sparrow, tree
sparrow, Harris sparrow, and white-throated sparrow (_Zonotrichia
albicollis_) also frequently use these thickets.

#Prunus serotina.#--Isolated trees of black cherry six to fifteen
inches in trunk diameter, have been noted on various parts of the
Reservation at widely scattered points. On a flat hilltop at the
southeastern corner of the Reservation there are many large trees of
black cherry, which make up a major portion of the stand, and trunks
of some are as much as 21 inches in diameter. Other trees in the
vicinity are mostly elms and honey locusts, and seemingly the area was
more open or perhaps entirely treeless in the recent past. The
presence of black cherry in forest often can be interpreted as
indicating more open conditions at the time the seedling became
established. Black cherry prefers a rich soil and an open habitat;
hence it is generally not common in woodlands of northeastern Kansas.

The fruits of black cherry are a favorite food of the opossum, and the
seeds have often been noticed in the scats of this animal.
White-footed mice store and eat the seeds. Two trees of black cherry
well isolated from other trees except for saplings in low thickets,
constituted the headquarters of a Bell vireo's territory each summer
from 1951 through 1955.

#Pyrus ioensis.#--Crab-apple is a small tree, usually less than five
inches in trunk diameter and less than 12 feet high. It grows both in
woodlands and in former pastures, but chiefly along the line of
contact. After removal of livestock in early 1949, crab-apple spread
into the edges of hilltop pastures, from the adjacent protected
woodland. Each year thickets of encroaching crab-apple have extended
farther into the fields, until, in 1955, there were graded series from
the trees along the fence, six feet high or more, to the seedlings 30
to 50 feet out in the fields. Dogwood, red haw, and smooth sumac are
among the most common associates of crab-apple as they share its
tendency to invade open land adjacent to the forest.

Evidently the tree is intolerant of browsing by livestock, as few were
growing in the pastured areas in 1948, but as soon as livestock were
removed these areas were rapidly invaded.

The thickets formed by crab-apple provide shelter for many kinds of
animals. Cottontails, especially, tend to stay in or near these
thickets. In autumn the fruits are eaten by them, and in winter, when
the ground is covered with snow, the bark is a major food source. Most
mature or partly grown trees show old scars near their bases, where
the rabbits have attacked them. Often the trees are completely
girdled. In years when snow lies on the ground for long periods
girdling is extensive and a substantial portion of the trees in the
thickets may be killed, but this mortality has been insufficient to
check the rapid spread of crab-apple.

The crab-apple is one of the trees preferred as a nesting site by the
cardinal. Other birds that frequently use the crab-apple tree as a
nest site include the field sparrow, towhee and indigo bunting.
White-footed mice, prairie voles and pine voles eat the fruit and

#Crataegus mollis.#--Red haw occurs over much of the Reservation, both
in woodland and former pastures. The trees are scattered, and are not
dominant, even on small areas. In the woodland, haw usually grows in
the more open situations. Where there are haws in denser woods, they
are usually large and old; seemingly they are survivors from a time
when the woods were more open. Haw is intolerant of shading, and being
of lesser height than any of the climax species, it cannot compete
with them. The present wide distribution of haw on the area is
secondary, resulting from the extensive cutting of the larger trees
and opening up of the woodland. Haw trees are most numerous on south
facing slopes that have grown up into thickets in the last 30 years.
Here its associates are chiefly honey locust, osage orange, dogwood
and elm.

Red haws have been recorded as nest trees of horned owls,
yellow-billed cuckoos, cardinals, and fox squirrels. Cavities in the
trunks are used by downy woodpeckers, titmice, chickadees and
white-footed mice.

#Cercis canadensis.#--Redbud is abundant in some parts of the
woodland. Trees are up to nine inches in diameter and 25 feet high.
They grow chiefly in rich soil on hillsides in moist situations.
Redbud and dogwood are in part complementary in distribution, each
forming an understory in parts of the woodland where the leaf canopy
of larger trees is not too dense. However, redbud is more tolerant of
shade. In general dogwood grows in the drier, more rocky situations
and redbud in better soil and damper sites. In the southeastern part
of the Reservation, on a west facing slope, redbud dominates, with
smaller numbers of elm, blackjack oak, and dogwood.

Several times nests of yellow-billed cuckoos were found in redbuds.
Titmice, chickadees, and red-eyed vireos forage in redbuds on many
occasions. Brown creepers forage on the trunks. Titmice, chickadees,
and downy woodpeckers used cavities in dead or dying redbuds. However,
there is no evidence that this tree is especially attractive to any
kind of vertebrate, or plays an important part in the ecology of the

#Gymnocladus dioica.#--Kentucky coffee-tree is one of the less
important trees on the area but it is widely distributed. In general
it is absent from the denser woods. On limited areas of certain slopes
it is the dominant species. The groves sometimes are in nearly pure
stands. Slope exposure evidently is not the determining factor in the
local distribution as groves have been found on hillsides of varying
exposure. The tree seems to flourish where the forest has been opened
by cutting of the larger trees. Groves are mainly on the more gently
sloping parts of the hillsides, or on the nearly level terrace. There
are few coffee-trees more than 12 inches in trunk diameter. The
largest tree examined was 27 inches.

In May, groups of orchard orioles (_Icterus spurius_) have been
observed in coffee-trees, seemingly attracted by the blossoms. These
concentrations never lasted more than a few days and seemed to involve
individuals that were still migrating or newly arrived and not yet
established on their territories.

In winter the large pods of this tree are used as food to a limited
extent by cottontails. The large hard shelled seeds resist attack by
most animals. Seemingly they are used by white-footed mice, as they
have often been found stored in the nest cavities of these mice,
beneath rocks or in logs.

#Gleditsia triacanthos.#--Honey locust is at present one of the more
important species of trees on the area. There are scattered locusts
throughout most parts of the woodland. In the bottomland fields there
are groves and scattered trees of medium to large size. On south
slopes honey locust, osage orange and red elm form thickets. On
hilltops, along woodland edges where fences were installed in the
mid-thirties, young honey locusts have become established and are now
abundant. Some have grown to a diameter of 8 inches or more. Honey
locust is the fastest growing of the trees on the area and therefore
has an early advantage in competing with other kinds. A locust of
25-inch diameter cut in 1950 was found to have 32 annual rings, an
average of only 1.3 rings per inch as contrasted with an average of
3.8 for all the trees studied, and more than 9 for some of the slowest
growing. In open fields, both those used for pasture and those
formerly cultivated, young honey locusts have sprung up in abundance
since the discontinuance of grazing in 1948. The species is resistant
to drought. It seems to have been limited on the area mainly by
grazing and shading. The locusts growing in the woods tend to be
concentrated near its edges. Those that are deeper in woodland
evidently became established after heavy tree-cutting had opened
clearings. Locusts in such situations, competing with other hardwoods
are of much different form than those growing in the open; the trunks
are long and slender and the crowns are narrow.

The south slopes that were originally prairie, were evidently only
sparsely clothed with trees up until the thirties when livestock were
fenced out. Then the abundant growth of shrubs and young trees formed
thickets. Honey locust, growing rapidly tended to dominate. The
younger locust saplings that were shaded beneath the leaf canopy died
in large numbers.

Honey locust plays an important part in the over-all ecology of the
area, providing both food and shelter for many kinds of animals. The
foliage is well liked by livestock; consequently young trees have
little chance of surviving in heavily grazed pastures. Rabbits like
both the foliage, and the bark. Often they girdle or injure young
trees, and eat the beans. Both the prairie vole and the pine vole
often feed upon the inner bark and root crowns of small saplings,
sometimes completely undermining them. These voles also store and eat
the seeds. Beneath large mature locusts, runway systems and burrows
of the pine vole are sometimes much in evidence. As ground vegetation
is scanty in these places it seems that the voles are attracted by the
abundant supply of locust seeds.

The spiny branches of locusts provide well protected nesting sites
that are utilized by various kinds of birds; mourning dove, horned
owl, yellow-billed cuckoo, gnatcatcher, cardinal and goldfinch have
been recorded nesting in locusts. The wood is relatively soft. The
hairy woodpecker has been recorded nesting in a cavity which it had
dug in a living honey locust, while the black-capped chickadee and
red-bellied woodpecker have been recorded nesting in cavities in dead
limbs. The summer tanager prefers large locusts near the edge of
woodland as singing stations.

Fox squirrels also often exploit the spiny protection provided by
locust trunks, and build their stick nests in these trees, usually in
a fork of the main trunk eight to twelve feet above the ground. Such
nest trees often are either isolated or are in groves of other
locusts. Presumably the squirrels are attracted to them by the supply
of locust seeds.

#Acer Negundo.#--Boxelder probably was not a part of the original
flora of the Reservation. The trees present now are few and scattered,
and most are not more than eight inches in trunk diameter. The species
seems intolerant of shade and does not grow in the denser woodlands. A
few are present along the banks of the intermittent streams, and there
are others in open woodlands of south slopes. The small patch of
bluestem prairie remaining at the northwest corner of the Reservation
is being invaded by a variety of shrubs and saplings, and boxelder is
by far the most prominent of these invaders, with two hundred
seedlings and saplings per acre.

#Ailanthus altissima.#--Tree-of-heaven is an Asiatic species that was
introduced early into northeastern Kansas, and has become established
locally in the woodland. Most of those on the Reservation are near the
central part of the southwestern one-fourth. Concentrated about the
site of an old homestead, occupied in the eighteen-seventies, within a
few acres, there are dozens of mature trees, up to 22 inches in trunk
diameter, and hundreds of saplings. Elsewhere on the Reservation the
species is scarce and is represented by isolated trees and scattered
clumps at a few places.

#Cornus Drummondi.#--This dogwood is the most abundant tree on the
area. However, it scarcely reaches the size of a tree. Most mature
examples are 1½ to 3½ inches in trunk diameter, and rarely more
than twelve feet high. Dogwood grows in greatest abundance on dry
rocky slopes where other trees are scarce. In small areas it may be
the dominant tree, often closely associated with chinquapin oak and
red elm. In parts of the woodland where there are larger trees,
dogwood may form an understory, its development depending largely on
the amount of light passing through the upper leaf canopy. Where the
canopy is dense and nearly continuous, dogwood tends to be eliminated
by shading. In some situations where forest has recently closed in,
most of the dogwoods are dead or dying. Especially on formerly
cut-over north slopes, where oak and hickory have sprung up in a dense
stand 20 feet high, with a thick canopy, most of the dogwoods have
been eliminated.

On the remaining hillside prairie near the northwest corner of the
Reservation, dogwood is the most prominent of the trees and shrubs
encroaching onto the area since it has been protected from fire--a
period of approximately 20 years. There are dense thickets of dogwood
along the borders of the prairie and the woodland edge.

The white-eyed vireo and Bell vireo both forage and nest in thickets
of dogwood and other shrubs.

#Fraxinus americana.#--White ash is localized on the Reservation and
most of the mature trees are within an area of perhaps three acres on
a steep slope of northwest exposure. Several of the largest trees,
well over a foot in trunk diameter, grow at the lower limestone
outcrop. Ash is most abundant at this level and at the terrace just
below it. On the one slope where it is concentrated, ash is one of the
most common trees, growing in association with American elm, chestnut
oak, black oak, and shagbark hickory. This area is one of the most
mesic on the Reservation. The soil is usually damp, with thick leaf
litter and rich humus. In hilltop fields, formerly cultivated or
pastured, saplings of white ash are among the most prominent invaders.

The leaves of this tree and especially its saplings, are favorite
foraging places for the tree frog. The groves of this tree provide
favorable habitat for the opossum, short-tailed shrew, gray squirrel,
and white-footed mouse. Birds that frequent the same habitat include
the black-capped chickadee, tufted titmouse, blue jay, rose-breasted
grosbeak (_Pheucticus ludovicianus_), yellow-billed cuckoo, red-eyed
vireo, gnatcatcher, hairy woodpecker, Kentucky warbler, and crested
flycatcher (_Myiarchus crinitus_).

  [Illustration: PLATE 7


  Upper figure shows gully in southeastern part of Reservation, which
  has enlarged and deepened greatly in the past 40 years. Heavy
  precipitation in the summer of 1951 resulted in the undermining and
  collapse of many large and medium sized trees, as shown in this
  photograph taken in March, 1956, by H. S. Fitch.

  Lower figure shows Cottonwood fifteen feet in circumference,
  growing on hilltop near south edge of the Reservation. This is the
  largest tree on the area. Several exceptionally large black oaks,
  chestnut oaks, and elms are present on the same hilltop. Photograph
  taken in December, 1954, by H. S. Fitch.]

  [Illustration:  PLATE 8

  Large American elm at edge of bottomland field in west part of the
  Reservation. Photograph taken on April 2, 1955, by H. S. Fitch.]

Summary and Conclusions

The University of Kansas Natural History Reservation, in the
northeastern corner of Douglas County, Kansas, is situated in an area
that originally supported two types of climax vegetation, tall grass
prairie, and hardwood forest. These associations were distinct and
sharply defined. The present distribution of the different species of
trees on the area, supplemented by the data from snails, indicates the
approximate distribution of the two original climaxes. The principal
climax trees of the original forest were mossy-cup oak (mainly in
bottomlands), black walnut, shagbark hickory, hackberry, red oak,
black oak (mainly on hillsides and hilltop edges), chestnut oak
(mainly on rocky upper slopes). Subclimax trees characteristic of
marginal situations include: American elm, red elm, white ash, honey
locust, osage orange, coffee-tree, red haw, dogwood, redbud, cherry,
wild plum and crab-apple. Others characteristic of hydroseral
situations include sycamore, willow (of four species), and cottonwood.

In the Kansas River flood plain and small tributary valleys, rich
mesophytic forest of predominantly oak-hickory type was present. In
somewhat stunted form, and with partial replacement of its species by
those of more xeric habit, it extended up onto hillsides sloping
north, east or west, and onto the adjacent hilltop edges. Slopes
having poor shallow soil and exposures mainly to the south supported
chiefly tall grass prairie, but also had compact clumps of blackjack
oak and post oak, usually more or less isolated from other parts of
the woodland. Hilltops were mostly treeless (except near their edges)
and supported a tall-grass prairie vegetation. Shrubs and various
kinds of small trees must have been a much less conspicuous part of
the woodland flora than they are at present, and occurred in small
ravines where shelter was inadequate for the larger forest trees, and
also along the extensive line of contact between forest and open land.

One of the earliest changes was the destruction of the bottomland
forest. With the rapid settlement of the region in the sixties and
seventies, lumber was in demand and the supply was limited. The
cleared land was productive as pasture. Heavy grazing combined with
drought, gradually altered the original tall grass prairie; the
bluestems and other perennial grasses were replaced by the introduced
blue grass and by various weedy forbs. Prolonged protection from fire
permitted encroachment of trees and shrubs into situations where they
had not grown previously. Heavy grazing however, tended to hold in
check the spread of the woody vegetation.

When the bottomlands had been cut over, lumbering operations were
extended onto those hillsides where the better stands of trees were
located. The cutting of large, mature oaks, walnuts, and hickories
opened up the woodland and permitted large scale encroachment by
subclimax species. American elm, especially, sprang up in thickets.
Ash, honey locust, cherry, red haw, crab-apple, dogwood, and the
introduced osage orange, thrived and spread in the situations to which
they were especially adapted. These species largely replaced the
original climax. Some of the trees cut, the oaks, sycamores, and
hickories, usually produced fast-growing stump sprouts and competed
vigorously with the invaders. At each successive cutting, however, the
climax species lost ground. American elm, being tremendously prolific
of seed, and only a little less tolerant of shading than its climax
competitors, soon became the dominant tree of the woodlands.

Literature Cited

  Braun, E. L.

    1950. Deciduous forests of eastern North America. The Blakiston Co.,
          Philadelphia, xiv + 595 pp.

  Fernald, M. L.

    1950. Gray's manual of botany, 8th edition. The American Book Co.,
          N. Y., lxiv + 1632 pp.

  Fitch, H. S.

    1952. The University of Kansas Natural History Reservation. Univ.
          Kansas Mus. Nat. Hist. Misc. Publ. no. 4: 1-38.

  Fitch, H. S. and D. H. Lokke

    1956. The molluscan record of succession on the University of
          Kansas Natural History Reservation. Trans. Kans. Acad. Sci.

  Flora, S. D.

    1948. Climate of Kansas. Rept. Kansas State Board Agric. 67, no.
          285, pp. xii + 320.

  Leonard, A. B. and C. R. Goble

    1952. Mollusca of the University of Kansas Natural History
          Reservation. Univ. Kansas Sci. Bull. 34: 1013-1053, 2 pls.

  Parks, G. S.

    1854. "The Tourist" [Column]. The Kansas Herald of Freedom, 1
         (no. 1) Wakarusa, Kansas Terr., October 21, 1854.

  Robinson, [Mrs.] S. T. D.

    1899. Kansas; its interior and exterior life including a full view
          of its settlement, political history, social life, climate,
          soil, productions, scenery, etc. Journal Publishing Co.,
          Lawrence, Kansas (10th ed.) xi + 438 pp.

  Taft, R.

    1950. The great sandy desert. Trans. Kansas Acad. Sci., 53: 441-442.

  Ward, [Mrs.] A. M.

    MS.  As I knew them--early settlers of Grant Township--Douglas
         Co., Kansas. Univ. Kansas Library, 26 pp.

_Transmitted April 20,1956._

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Transcriber's Notes

    With the exception of the typographical correction noted below,
    the text in this file is that presented in the original printed
    version. Minor corrections of missing periods or commas may have
    been made; but are not reported here. Some of the text was
    rearranged so that figures and tables do not split paragraphs.

Emphasis Notation

    _Text_  :  Italics
    #Text#  :  Bold-Italics

Typographical Corrections

    Page 92 Para. 4: plaes => places

       *       *       *       *       *       *       *

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