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diff --git a/34353.txt b/34353.txt new file mode 100644 index 0000000..c8b7f9f --- /dev/null +++ b/34353.txt @@ -0,0 +1,2569 @@ +The Project Gutenberg EBook of The Forest Habitat of the University of +Kansas Natural History Reservation, by Henry S. Fitch and Ronald L. McGregor + +This eBook is for the use of anyone anywhere at no cost and with +almost no restrictions whatsoever. You may copy it, give it away or +re-use it under the terms of the Project Gutenberg License included +with this eBook or online at www.gutenberg.org + + +Title: The Forest Habitat of the University of Kansas Natural History Reservation + +Author: Henry S. Fitch + Ronald L. McGregor + +Release Date: November 17, 2010 [EBook #34353] + +Language: English + +Character set encoding: ASCII + +*** START OF THIS PROJECT GUTENBERG EBOOK FOREST HABITAT OF U. OF KANSAS *** + + + + +Produced by Chris Curnow, Tom Cosmas, Joseph Cooper and +the Online Distributed Proofreading Team at +https://www.pgdp.net + + + + + + + + + + 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 + + BY + + HENRY S. FITCH AND RONALD L. MCGREGOR + + + UNIVERSITY OF KANSAS + LAWRENCE + 1956 + + + + + UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY + + 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 + 1956 + + 26-3855 + + + + + The Forest Habitat of the University of + Kansas Natural History Reservation + + By + + HENRY S. FITCH and RONALD L. MCGREGOR + + + + +Introduction + + +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. + + + + +History + + +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 +part. + +[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 +forests." + +[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 +elevation." + +[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 +floor. + +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 +storms. + +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 +forties. + +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 +discernible. + +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 +trees." + +[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 + interval.] + + [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 +coffee-tree. + +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 +sample. + + 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 +occurred. + +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 1-1/2 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 2-1/2 inches in stem diameter, there were 17 +dead and 53 live saplings, while in the size group 2-1/2 to 3-1/2 +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 1/2-inch | 1/2-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 1/2 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 +browsed. + + + + +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 +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 + + EXPLANATION OF 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. + 59(4). + + + 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._ + + + * * * * * * * + + +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 + + * * * * * * * + + + + + +End of the Project Gutenberg EBook of The Forest Habitat of the University +of Kansas Natural History Reservation, by Henry S. Fitch and Ronald L. McGregor + +*** END OF THIS PROJECT GUTENBERG EBOOK FOREST HABITAT OF U. 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