#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
Reservation.
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 16-1/2 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
seen.
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
site.
#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
Reservation.
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
conditions.
[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
situations.
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
Reservation.
#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
abundant.
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
survived.
#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
seed.
#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
area.
#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 1-1/2 to 3-1/2 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