3C / HARVARD UNIVERSITY LIBRARY OF THE Museum of Comparative Zoology MUa COMP. 200U LIBRARY flPP 1 4 1972 THE UNIVERSITY OF KANSAS I NATURAL HISTORY I RESERVATION -* *l?^"-" "l%^." The University of Kansas Natural History Reservation in 1965 BY HENRY S. FITCH University of Kansas Museum of Natural History editor: E. RAYMOND HALL MisccUancous Publication Nu. 42, pp. 1-60, 2 maps, S plates Published December 10, 1965 Lawrence • Kansas PRINTED BY ROBERT R. (BOB) SANDERS. STATE PRINTER TOPEKA. KANSAS 1 965 30-8140 The University of Kansas Natural History Reservation in 1965 BY HENRY S. FITCH CONTENTS PAGE ixtroduction 5- Objectives 6' The Concept of a Natural Area 8 History 10 The Rockefeller Experimental Tract 22 Research 24 Description of the Area 34 Plant and Animal Life 35- Trees, shrubs and woody vines 48 Flowering plants 48 Grasses, sedges, rushes, lilies 49 Lichens, mosses, liverworts, ferns 50 \Limmals 50 Birds 51 Reptiles 52 Amphibians 52 Fishes 52 Beetles 52 Crickets, grasshoppers, katydids, roaches 53 Ants, bees, wasps 53 Moths 54 Butterflies 55 Spiders 55 Chiggers 5& Crustaceans 55 Earthworms, mollusks 55 ElBLIOGRAPHY 57 (3) The University of Kansas Natural History Reservation in 1965 Introduction In 1947, at the suggestion of Professor E. Raymond Hall of the Department of Zoology, Chancellor Deane W. Malott proposed to the State Board of Regents that a tract of 590 acres of the University's "Robinson Farm" land be set aside as a reservation where native plants and animals might be protected and studied under natural conditions. This action was taken and on July 1, 1948, the writer came to The University of Kansas, Department of Zoology, and began field work as first superintendent and resi- dent naturalist of the newly created natural history reservation. In the following years a residence and small laboratory were com- pleted and the writer and his family moved onto the area on March 1, 1950. As anticipated, the Reservation has provided a valuable adjunct to the several departments of biological science of the University, providing a place for field work to supplement the types of insti-uction carried on in classrooms, laboratories and museums. The Reservation is administered by the Department of Zoology. Early in 1952 a pamphlet was published (Fitch, 1952) briefly describing the Reservation and its plant and animal life, its ob- jectives, the research program during its first three years, and the work planned for later years. The publication is now much out- dated and has long been unavailable. The present report incor- porates the more essential material included in the earlier report, presents some of the more general findings, and ti'aces the trend of the numerous investigations already completed or in progress on the area in the 17 years since it was made a reservation. In the present report attempt has been made to avoid scientific names except in the formal lists of species. For most groups, ex- cept those of some invertebrate animals, widely accepted vernacular names are available. The reader is referred to the following au- thoritative works for both the scientific names and the vernacular names used here: Fernald, 1950, for flowering plants; Hall, Ander- (5) 6 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. son, Jones and Packard (1957) for mammals; Johnston (1960) for birds; Smith (1956) for reptiles and amphibians; Harlan, Speaker and Reece (1956) for fish. Much of the information in this report is not new but is com- piled from previous publications — many of them based on field studies carried on partly or wholly on the Reservation. These publications are all included in the Bibliography section. A large ninuber of persons have contributed to the information used herein. The writer is especialh' indebted to those persons who have donated large amounts of time to the study and identification of specimens. Dr. Ronald L. McGregor and Dr. ^^'ortllie H. Horr identified most of the plants. Mr. A. Noel McFarland identified all the moths or submitted them to specialists for identification. My former research assistants, Mr. William N. Berg, the late Robert M. Hedrick, and Dr. Allen Brady, identified many insects and spiders. Other persons who have identified many specimens in- clude: Dr. George W. Byers, Mr. J. B. Karren, Mr. Bruce Harri- son and Mr. H. L. WilHs (insects), Dr. Paul R. Ehrlich (butter- flies), Mr. Charles D. Miles (snails). Dr. William Murchie (earthworms), Dr. Frank Kurczewski (wasps). Dr. M. R. Smith (ants), and Dr. Robert L. Gering, Dr. Willis J. Gertsch, Dr. T. B. Kurata, Dr. Herbert W. Levi, Mr. Wilton Ivie, and Dr. M. H. Muma ( spiders ) , and Dr. Jerry Tash ( crustaceans ) . The photographs included in this report are intended to illus- trate the vegetation and the general habitats on tlie area as the\- existed in 1964 and 1965, but also are intended to illustrate ecological succession. Most of these photographs had their prede- cessors, taken from the same spot at the same season but years ear- lier, and published in one of the earlier reports concerning the plants and animals of the Reservation. These earlier photographs are not reproduced in the present report, but the publication of each is cited, so tliat the interested reader may compare the scenes and judge the extent of successional change in the interval of year's elapsed between the photographs of each pair. Objectives The objectives of the University of Kansas Natural History Reservation are: to accomplish research in ecology; to teach biol- ogy, with emphasis on natural history; and to conserve a sample of the Kansas flora and fauna. It is intended that the Reservation be maintained permanently as a natural area and a research sta- tion, with a long-term program of study, concerning the inter- Natural History Reservation 7 relations of native animals and plants with their physical and biotic environments, and their popnlation dynamics under natural conditions. Continued collaborativ^e effort by various specialists working on this limited area will ultimately provide unusual in- sight into these interrelations. Another function is to promote a better appreciation of the land and of living plants and animals by the public in general. The area provides practical demonstration of the benefits to be derived by protection of the land from abuse and exploitation, showing, for example, the control of erosion by natural vegetation after cultivation and grazing are stopped, and gradual restoration of fertility and disappearance of noxious weeds after discontinuance of overgrazing. This is one of the few areas within the state of Kansas where native plants and animals are rigidly protected from disturbance by humans and where primitive conditions are to some degree approximated, so that the natural interrelations of species are revealed. Constantly expanding human populations, agricul- ture and industry encroach everywhere except in the few such places that are maintained by constant effort. The animals pro- tected on the Reservation include species of native game such as deer, rabbits, squirrels, and quail, which propagate naturally, attain high population densities on the area, and tend to disperse into depleted surrounding areas and restock them, to the benefit of local sportsmen. Another major function of the Reservation is that of providing a place conveniently near the University campus where classes in the biological sciences may carry on field work. Each year hundreds of students in many different classes visit the area on organized field trips. Traditionally these have included the large class in Biology I, of students experiencing their first contact with a life science. Early in their training these students thereby gain some grasp of ecological principles, and on the field trips each student is required to use field equipment, and to find and study animal life on his own. Many other classes and groups not con- nected with the University visit the Reservation each year. Among these are classes in biology and ecology from various colleges and universities in Kansas, high school biology classes and science clubs, elementary schools, groups interested in bird study, and Boy Scouts. Many interested individuals of the general public, and research workers from other states or from foreign countries visit the area from time to time. 8 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. The nearness (6/2 miles by road) of the Natural History Reser- vation to the campus of the University of Kansas in LawTence strengthens the teaching function of the Reservation over what it would be if the Reservation and Campus were farther apart. The 590-acre Reservation comprises all but the southwestern 50 acres of the northeasternmost section (see Fig. 1) in Douglas County and is only four and a half miles north and one and a half miles east of the City Hall at 11th and Massachusetts streets, Law- rence, Kansas. The Concept of a Natural Area In recent decades unprecedented growth of industry and popula- tion in the United States has been accompanied on the one hand by the rapid and progressive destruction of remaining natural com- munities of plants and animals as they formerly existed by a public callous and largely unaware of them and, on the other hand, by an increasing sense of outrage and urgency concerning this trend by more enlightened segments of the citizenry. Through the efforts of the latter, channeled and directed into effective action by the Nature Conservancy and kindred organizations, hundreds of "nat- ural areas," large and small, in all parts of the country, have been acquired and turned over to the custodianship of institutions or agencies interested in their preservation. Some of the best examples of natural areas are in the larger national parks. Through the far-sightedness of the policy-makers in the National Park Service, such areas were to be preserved un- impaired for posterity. Nevertheless, in national parks and in all other natural areas there has been some compromise between the ideal of preservation of natural conditions and the practical con- siderations of obtaining some short-term benefits to mankind. In late years the provision of recreation on a large scale has been emphasized by the National Park Service; more than 170 areas principally for that purpose now (1965) are administered by the Service, in addition to the 32 National Parks per se in which preservation of natural conditions unimpaired (and therefore nat- urally occurring changes) was the primary and prescribed aim. A degree of dilution of National Park policy, and more especially practice, has permitted massed humanity in such popular places as Yosemite Valley, the basin containing Old Faithful Geyser in Yel- lowstone, and Clingman's Dome in the Great Smoky Mountains, thereby so greatly altering the plant and animal communities locally that these places do not qualify as natural areas. Although these Natural History Reservation 9 are extreme examples, ecosystems everywhere including all so- called natural areas, have been more or less altered from their natural condition by man. The University of Kansas Natural History Reservation is no ex- ception in this regard. In approximately 95 years prior to 1949 the area was grazed by domestic livestock and substantial parts of it were cultivated at one time or another. In administering the area since 1949 the policy has been "hands-off" allowing nature to take its course with no interference, but several notable exceptions have been made. In the first place, fire has not swept over the area in more than 20 years. In the second place large primary consumers, bison and elk (wapiti), have been absent. In the third place the decision to build a dwelling and laboratory near the center of the area, with associated well and pumphouse, garage, small lawn, sewage disposal field, access road, parking area, power lines, and clotheslines has changed a small part of the area eliminating some species but rendering conditions favorable for those which require an edificarian habitat. In 1949 soon after the Reservation was set aside as such, an eroded barren upland field in its northeast quarter was sown to native tall-grasses, and in 1953 a silted pond was deepened and an outlet pipe installed to provide a permanent water supply, and assure the continued presence of certain aquatic plants and animals. While picnicking and camping per se are not encouraged be- cause of potential destruction or disturbance of plant and animal life, visitors who are interested in the flora or fauna, or the area itself, are welcome. Attempt is made to impress visitors with the fact that all plants and animals are protected, even those which are considered undesirable elsewhere. Harvesting of wild crops such as grapes, mushrooms, berries, or nuts, is not allowed. As a general pohcy plants and animals are neither introduced to the Reservation nor removed from it and effort is made to exclude activities that unnaturally influence popula- tions. Nevertheless, limited collecting, for study purposes, of plants and small animals has been permitted when the kinds involved were sufficiently abundant that it seemed no appreciable effect would result on their populations or on the ecosystem as a whole. In this regard a somewhat arbitrary distinction has been made in per- mitting the collecting of insects by classes and individuals but not permitting such collecting of small vertebrates, even those of the more abundant and prolific species. Visitors and workers on the area are admonished to avoid any 10 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. activities that might alter or destroy habitats — notably tearing apart decaying stumps or logs, stripping of loose bark from trees, or careless turning of rocks or logs. Live-trapping constitutes an im- portant means of studying small vertebrates, and has been done on a large scale, sometimes with the use of bait, even though it is recognized that such operations may alter the numbers and be- havior of the animals being studied, and perhaps of other kinds. Domestic animals are rigorously excluded from the area. A major policy decision involving the philosophy of a natural area was that concerning exclusion of fire; the Reservation lies in an area of fire subclimax and the tall-grass prairie found in the region by explorers and early settlers was fire-dependent. Fires started by lightning or set by aboriginal human inhabitants in ac- cordance with long-establislied custom often swept unchecked o\'er hundreds of square miles, destroying the plants and animals except those specially adapted to escape or withstand fire. Preserva- tion of the remaining remnants of this prairie type and the return of other areas to a like condition could have best been accom- plished by periodical controlled burning. But the effects of fires under primitive conditions probably could not be precisely dupli- cated with any program of controlled burning. The policy of pro- tection from fire allows areas that were formerly fire subclimax to develop into the "climatic climax" of deciduous forest with its rich associated fauna. The 160-acre Rockefeller Experimental Tract adjacent to the Reservation but not committed to the same policy of protection, even from experiments, provides an excellent situa- tion for manipulation of fire as a tool in maintaining a topical tall- grass prairie subclimax. History The University of Kansas Natural History Reservation is situated in the broad belt of the ecotone or transition from the eastern hardwood forests to the grasslands of the Great Plains. Its original condition is not well known, but before the coming of the White Man, this area probably consisted, as it still does to-day, of mixed woodland and grassland, with each t\'pe dominating in the local situations more favorable to it. Some ecologists, aware of the rapid invasion of woody vegetation in grassland areas protected, from fire, have adopted the extreme view that the entire area was virtually treeless, but there is much evidence to the contrary. In Major W. S. Long's report of an 1819 exploring expedition Natural History Reservation 11 through the area that is now in Doughis County, Kansas, and nearby counties, various forest trees of kinds still common in the area were mentioned (Taft, 1950:442). Early residents of Lawrence, Kansas, have described the countryside about the recendy settled tow n and between the town and the present Reservation as it ap- peared in the early eighteen-fifties (Robinson, 1899; Parks.^ 1854). The late Mrs. Anna Morgan Ward, daughter of a pioneer family which settled in 1864 on the section of land adjoining the Reserva- tion on the south, has set in writing her girlhood memories of the area (Ward, 1945). A U.S. Government map prepared sometime in the eighteen-fifties showed distribution of forest in Douglas County. Surveying records of the section lines bounding the Reservation are also available and have provided information about the vegetation at regular intervals along these transects, in different soils and slope exposures. Finally, numerous core borings of large trees on the Reservation made in the early nineteen-fifties have indicated approximate ages of the trees and possible minimum ages of the groves where they occur. This e\'idence has been set forth in some detail in earlier pub- lications (Fitch, 1952:4-6; Fitch and McGregor, 1956:79-86). In brief, it may be stated that parts of the flood plain of the Kansas River supported a forest with a climax vegetation of many species of large trees. The forest had oaks, hickories, elms, w^alnut, linden, ash, hackberry, Kentucky coffee tree, honey locust and cottonwood. The broad bottomland forest had attenuate peninsular extensions along tributary streams, but elsewhere, even on some of the bottom land, tall-grass prairie dominated the landscape. Where soil, mois- ture, and slope exposure were favorable, groves extended up onto hillsides or hilltop edges in a sparser, less luxuriant type of forest than that which grew in the bottomlands. Some groves were iso- lated and surrounded by prairie. Usually in autumn, prairie fires burned the dry grass from extensive areas and any woody vegeta- tion beginning to encroach was killed. On the Reservation two small valleys draining to the west and south, respectively, were both heavily timbered, but they were cleared early and used for cultiva- tion of crops and for grazing. North-facing slopes also were sparsely wooded and hilltop edges had at least a thin line of trees along the rock outcrops. Elsewhere, notably on hilltops and slopes of predominantly southward exposure, there was tall-grass prairie, except for occasional open groves of blackjack oak and post oak. For centuries prior to the settlement in the eighteen-fifties and 12 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. sixties, this region was under the influence of a succession of aboriginal cultures. Relatively rapid succession of aboriginal groups resulted from the progressive colonizing of European settlers, fan- ning out from the Atlantic seaboard with resultant westw'ard dis- placement of native tribes. Frequent changes in population density, and varying practices in agriculture, keeping of domestic animals, hunting and burning, must have had pronounced effect on the plant and animal community. From time to time stone-age artifacts have been found on the Reservation attesting to the former presence of primitive man, but as yet none of the artifacts found has been definitely identified with any specific culture or period. In 1829 the Delaware Indians were moved from Ohio to the land granted to their tribe by the Federal Government between the Missouri and Kansas rivers, including the tract that is now the Natural History Reservation. They occupied this land until 1868, when, at their own request, they were moved again to, Oklahoma. Having absorbed much of the White Man's culture in nearly 200 years of contact prior to their move to Kansas, these Indians lived much as did other frontiersmen who were their contemporaries. Earlier the Kansa Indians were in possession of the lower Kansas River Valley, but they too were of more eastern origin. They were a small tribe (estimated at 800, and by another traveler at 1750, in the early eighteen hundreds) of Siouan affinities, most closely related to the more southern Osage. In the seventeen hundreds known \illage sites of the Kansa were near the present towns of Atchison and Leavenworth, but five villages that were known in the interval 1804 to 1836 (mentioned in the accounts of Lewis and Clark, and later travelers) were all situated beside or near the Kansas River between the present cities of Topeka and Manhattan. Economy of the tribe was based to a large extent on primiti\e agri- culture. However these Indians also depended on the buffalo for part of their subsistence, and almost up to the time of their removal to Oklahoma in 1873 they continued to make an annual buffalo hunt, following a regular route to hunting grounds in central Kan- sas far to the west of their villages. Ry 1811 the tiibe had acquired many horses and mules, rendering them more mobile in their hunt- ing, and this may have hastened the extirpation of the buffalo and elk from northeastern Kansas. In 1825, by treaty with the United. States, the Kansa relinquished claim to the part of northeastern Kansas where the Natural History Reservation is located, and from Natural History Reservatiox 13 1847 to 1873 the tribe occupied three villages along the Neosho River near Council Grove (Wedel, 1959:50-54; Chapman and Chapman, 1964:81). With the disappearance of the buffalo, elk and deer no grazers above woodchuck size remained and the annual crop of prairie grass was largely unused. As a result "primary consumers" of the vegetation became relegated to a secondary role and "reducers" and decay organisms became relatively more important. In the absence of effective grazers the accumulation of dead and dry vege- tation over intervals of years might have intensified the effects of fire. At any rate the accounts written in the eighteen-fifties, de- scribing a savannah type with isolated groves of trees surrounded by prairie, seem to suggest a recent encroachment of prairie at the expense of a formerly more extensive woodland. In a more remote period, perhaps before the influence of the White Man had affected the region, even indirectly, Indians of the so-called "Nebraska" Culture occupied northeastern Kansas and lived by primitive agriculture, probably with much less effect on the ecology than the later cultures that utilized the horse and firearms. Attempts to ti'ace ecological trends in the area still farther back in time must be largely speculative. Major ecological changes must have been wrought by: 1. The Hypsithermal Period, from about 3000 to 8000 years before the present, with a climate much warmer than that now prevailing (and drier in its late stages ) resulting in widespread displacement of biota; 2. Rela- tively sudden extinction in the Great Plains region in late glacial ( Wisconsinan ) or postglacial time of various species of large herbiv- orous mammals including buffaloes different from those now liv- ing, and native horses and elephants. It has been speculated on the basis of some circumstantial evidence that early human hunters played some part in this extinction (Wormington, 1957; Martin, 1958; MacGowan and Hester, 1962:196). Dr. Charles Robinson, first governor of Kansas, acquired his farm of more than 1200 acres (in Douglas County) in 1864. The portion that is now the Reservation was long known as the Robin- son Pasture and up to the time when it was acquired by the University was used mainly for grazing. However, four separate homesteads were made on the area, each occupied for a few years, mainly in the eighteen-seventies and eighteen-eighties. A peach orchard was planted at one site. Small acreages were cultivated 14 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. by the homesteaders, who also grazed livestock. Rocks and boulders were piled into walls marking the boundaries of the separate farms. Parts of these walls have been torn downi and removed; the remaining parts, now in a state of collapse, are the only remaining evidence of the fonner farmsites. Usually the homesteader failed to profit from his small scale marginal farming operations, and eventually lost possession of the land, which re- verted to pasture under control of the original owiier. In the eighteen-nineties pai"ts of the area including some of the hillsides still supported a mixed forest of \irgin timber. In 1951 the late Frank H. Leonhard remembered harvesting this timber some 6Q years before, especially large walnut trees. Dr. Robinson bequeathed his entire farm to The University of Kansas, but the transfer of title did not take effect until tlie death of his widow in 1910, and for some years diereafter the University administration did not have any active interest in the area. Several farmers on neighboring land leased it at different times for grazing their livestock. Heavy overgrazing caused de- terioration of the forage crop, originally dominated by a luxuriant cover of native grasses, to a condition in which coarse and un- palatable weeds were prominent. Severe erosion with gullying occurred. About 1900 use of the area now constituting the Reser- vation had passed to the J. F. Morgan family and the entire area was used for grazing. Sheep were pastured on parts of the area, and a corral for them was situated a short distance northeast of the present Reservation headquarters. By 1964 this old corral area had grown up to an almost impenetrable thicket of brush and young trees, especially dogwood, elm, honey locust and osage orange, and with vines of poison ivy, Virginia creeper, grape and bittersweet. Parts of the bottomland were fenced and broken for cultivation in 1907, 1912 and 1915 and hilltop fields were first cultivated in 1909. Throughout this period there was intermittent tree-cutting. Several acres of hilltop and south slope in the northwest corner of the section were protected from livestock. There prairie hay was mowed annually and the remaining dry vegetation was burned off at less frequent intervals, thereby maintaining a prairie type. In the mid-thirties control of the area finally passed to the University. At that time a program of development was launched by the University and the U. S. Soil Conservation Service with relief labor. The work included: filling gullies, digging diversion ditches and building check dams and terraces to prevent erosion; Natural History Reservation 15 clearing extensive thickets; bindweed eradication from the culti- vated areas; and fencing off the wooded hillsides from the valley and hilltop pastures for protection from livestock. This work ex- tended over several years and one main objective was to utilize the area for growing timber. However, plans to make extensi\e 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 Depart- ment 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 slirubs 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 situa- tions. Late in 1948, after the area had been made a Reservation, live- stock 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 pro- tected 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. Much of the northwestern part of the Reservation is drained by a small intermittent creek which leaves the section at the mid-point of its west side. A somewhat larger intermittent creek drains ap- proximately the southeastern half of the section. Both creeks have deeply incised channels; that of the larger is more than 15 feet below bank level in some places. The verbal and written accounts of old-time residents, comparison of early and later maps, and known changes in the fauna, all provide evidence that the creeks formerly flowed in shallow meandering channels only a few feet below bank level, and that their flow was more constant than at the present time. Erosion occurred as a result of cultivation and heavy grazing. In a period of years from 1902 to 1905 when rainfall was unusually heavy, saturated soil of several hillside areas slipped down\\'ard with a vertical displacement of several feet. According to the evidence, gully-cutting was most severe between 1915 and 1948. Even after cessation of cultivation and the removal of live- stock, permitting regeneration of vegetation in formerly bare areas, erosion and gully-cutting was severe in the summer of 1951 as a result of unusually heavy rainfall. Incision of deep gullies in the 16 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. two small valleys has lowered the water table, hastened run-off after storms, and thereby rendered conditions more xeric. Min- nows ( fatheads and stone rollers ) and mollusks ( Vertigo ovafa, Lymnaea porvo, Heliosoma trivoJvis, Physa anatina, Pisidium com- prcssum) which formerly abounded in or near the then more sluggish and more nearly permanent creeks have now completely disappeared or have become scarce and localized. In 1948 when the policy of protection was finally put into prac- tice on the area, some longtime trends resulting from cultivation, overgrazing, erosion, and the harvesting of trees, wild crops, and game animals were in part reversed. Responses to the changed treatment were immediate and in some instances they were striking. The closely grazed pastures became covered with high grass and weeds and by the end of the first growing season accumulated a ground litter of dead vegetation. Small mammals including the prairie vole, cotton rat, harvest mouse, and jumping mouse had become abundant after two growing seasons. Reptiles including the glass "snake," blue-racer, prairie king snake and garter snake, made corresponding gains but with a longer lag. By the fourth growing season the luxuriant weedy vegetation of the pastures, in which ironweed, vervain, germander, nightshades, milkweeds and spurges were prominent, had been largely replaced by grasses (es- pecially awnless brome and Kentucky blue-grass). Concurrently with this shift to a more homogeneous vegetation type, populations of the dominant small rodents, the prairie vole, cotton rat and harvest mouse, trended downward and have never again approached the maximum levels attained in 1951. Numerous small vertebrates that prefer open situations, notably the lark sparrow, kingbird, pocket gopher. Plains harvest mouse, ornate box turtle, six-lined racerunner, Great Plains skink and collared lizard became scarcer, more localized or entirely absent. The ruby-throated hummingbird once common in summer, became rare as weedy flowering species, especially germander, declined. Concurrently with the increase in grass, small trees and shrubs were springing up in great numbers. By 1954 brushy thickets of mixed elm, crab apple, dogwood, sumac, honey locust, Osage orange and red haw, had sprung up in former pastures, cultivated fields, and prairie, and especially in the edges of these areas where they adjoined woodland. Animals preferring this type of habitat including the white-footed mouse, cottontail, indigo bunting, cardinal, cuckoo and brown thrasher, increased. Scattered trees of medium to large size, mostly elm, honey locust, Osage orange, ailanthus and Kentucky coffee tree, occurred through- Natural History Reservation 17 out most of the pasture areas and provided seed sources. In 1949 when the effect of grazing livestock in holding in check the growth of tree seedlings was eliminated, the former pastures in both bot- tomland and hilltop entered a stage similar to that which had oc- curred 10 to 12 years earlier on most hillsides of the Reservation when the hillsides were fenced to exclude livestock. In the summer of 1952 a systematic survey of the vegetation on the Reservation was made. Series of plots of 1/100 acre each were examined in most of the open parts of the Reservation, and tlie percentage of the total vegetative cover made up by each species was estimated. The trend of the records showed striking changes when compared to a more general survey made in 1948 and 1949. The detailed survey of vegetation was repeated in the summer of 1964. Table 1 shows the trends in the more important kinds of plants — those that comprised at least two per cent in one or more of the series. A much larger number of species were represented each of which separately made up less than two per cent, but combined they made up appreciable percentages in some of the series, and have been included under the heading of "miscellaneous." For the sake of brevity groups of species that are similar ecologically, and closely related, have been com- bined under the same heading in the table in some instances: for example, "thistle" includes Cirsium altissimum and C. canadensis; "goldenrod" includes Solidago alflssima, S. lindheimeriana, S. nemoralis, S. uhnifolia, and S. rigida; and perennial tall-grasses include big bluestem, little bluestem, Indian grass, side-oats gramma and switchgrass. Over the 12-year period from 1952 to 1964, some of the main trends were: (1) the reduction of weedy vegetation on the for- merly pastured areas and its replacement by grass, (2) reduction of ragweeds and other annual pioneer weeds on the old-field ai^eas and their replacement by perennials, especially goldenrods, (3) in- crease in woody vegetation, especially young trees of elm, dogwood, Osage orange and honey locust on all the areas. The woodland areas of the Reservation, especially those parts having climax species, have changed much less rapidly than the fields. However in 1953 and 1954 a high proportion of black oaks died from the combined effects of drought and disease. Also in 1954 heavy mortality of American elms was noted and the die-off (probably resulting from attacks of introduced bark beetles and phloem necrosis) has continued with increasing severity up to 1965. The prospect is that this most important kind of tree in the 2—8140 18 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. Table 1. — Vegetation on Unforested Areas of the University of Kansas Natural History Reservation, Sampled on 1 100 Acre Plots, Showing Changes in Percentages From 1952 to 1964. Number of plots . SO 2 Kind of vegetation grasses awnless brome . . 44 3 68 . 8 Japanese chess. . o 7 bluegrass 14 9 12 4 panic three-awn perennial tall- grasses serai weeds aster saw-tooth sunflower bindweed. ... care.x 3 9 fleabane germander 4 5 horseweed legumes {Desnianthus Lespedeza, Melilolus) hemp iron weed 10 6 nightshades . . . polygonum 2 2 ragweeds .1 thistle goldenrod trees and shrubs elm dogwood Osage orange . locust walnut grape sumac coralberry 10 3 miscellaneous 3 5 11 5 13 C C ■ -■sz 1952 1964 9 9 68 s o O -^ y >■ t "-- ■^ w •£! 1952 1964 60 6 7 4 10 9 19 1^9 72 6 5 3 1.2 193 123 CO o O 0- 1952 1964 9.1 6'7 2 0 2.2 46 5 10 0 2 5 15 10 7 2.7 .... 3.1 30 .5 2 4 12 40 3 5.2 .6 2 16 8 100 100 fc "5: a; b£ 10 13 9 4 5 50 -3 O Pi 1952 1964 47 9 61 0 14 9 10 4 12 1 61 Natural History Reservation 19 local woodland will be largely eliminated. Already the general aspect of the woodland on most parts of the area is greatly changed; the dying of elms has been accompanied by an opening up of the canopy which has permitted large scale encroachment of brushy and weedy species. In 1954, 200 elms that were approximately two feet or more in trunk diameter were recorded on the Reservation. A recheck of these same trees in November, 1964, showed that 54 percent were then dead or dying, and most of those remaining appeared to be in poor condition, obviously diseased. In smaller elms the incidence of mortality was even higher. In 1948 there was little decaying woody material on the ground, as the forest consisted mostly of young trees, but in 1964, there was an abundance of decaying logs and of dead snags still standing, creating conditions favorable for the flora and fauna of this type of microsere. Approximately 8.3 acres of bluestem prairie in the northwestern corner of the Reservation was rapidly invaded by woody vegetation after cessation of burning and annual mowing; by 1964 parts of this area had grown up to an almost impenetrable tangle of brush and little of the prairie vegetation remained. Certain major changes noted in the animal populations of the area are due primarily to factors other than ecological succession. The white-tailed deer, for example, has gradually increased. In the first year of field work on the Reservation no deer or signs of them were noticed but in the following years they were seen with increasing frequency, and by 1964 deer sign had become conspicu- ous and deer were seen often, sometimes in groups of ten or more. The trend reflects increase in the population of deer over the entire eastern part of Kansas. In 1963 and 1964 the southern bog lemming was caught frequently in traps set for small mammals, and had be- come moderatelv common on the Reservation, Rockefeller Tract and other areas of northeastern Kansas, whereas none at all had been recorded in 14 years of field work previously. Despite fairly intensive but variable collecting efforts in northeastern Kansas by mammalogists, the bog lemming has seldom been recorded. How- ever, the striking increase in numbers in 1963 and 1964 was not unprecedented; in 1925 and 1926, 87 were trapped in Douglas County by students from the University interested in mammals, 20 others were caught in the years 1924, 1927, 1928, and 1929, and one each in 1894 and 1937 (Cockrum, 1952:194). The reason for the increase is still unexplained. 20 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. The eastern woodrat was phenomenally abundant on the Reserva- tion in 1947 and 1948 but underwent drastic reduction in a period of years thereafter. Although from time to time its population has made gains over periods of years, it has never again approached the high lexel of the late nineteen forties. The muskrat has been present in small but variable numbers each year at the pond, but the colony has not been continuous. In autumn as a result of low water level and reduced food supply the rats have usually been eliminated by predators but in early spring new immigrants arrive to recolonize the area if habitat conditions are favorable. The hispid cotton rat, formerly unknown at the latitude of the Reserva- tion, extended its range northward and reached the area in the early nineteen-hundreds but in this northern part of its range it is subject to decimation by severe winter storms. At times its num- bers have reached remarkably high levels, of several dozen per acre, but at irregular intervals of several years, at times of severe and prolonged low temperatures, it has died off except for a small percentage of survivors in unusually favorable habitat situations. The Carolina wren also is unable to withstand unusually low tem- perature, and twice in a sixteen year period it has been eliminated from the Reservation in severe winters. After its disappearance several years are required for it to return to the area from unknown refugia and to regain its former numbers. The chuck- wills- widow is a common nesting species some years, but in other years it is much scarcer or absent, and the Reservation is near the northern edge of its breeding range. In late May and June, 1964, there was a mass emergence of a periodical cicada (Mogicicada cassini) on the Reservation and for a time these large insects with populations of tens of thousands per acre, dominated the aspect of the habitat and created an abundant food supply for insectivorous animals in general. A more lasting effect was created by the cylindrical soil tunnels extending vertically from feeding places on tree roots to the surface and excavated by the mature nymphs preparing to emerge. Besides aerating the soil and rendering it more porous these tunnels offered ready-made shelters for various small subterranean animals. In 1949 the late Dr. Paul B. Lawson showed the writer extensive damage to honey locust twigs still remaining from the mass ovipositing of periodical cicadas of the 1947 brood. The next major emergence is to be expected in 1981. The "spring cankerworm," the larva of a geometrid moth, varies greatly in abundance from year to year but it is a primary con- Natural History Reservation 21 sumer of major importance ecologically. In late April and May the larvae are growing rapidly and feed voraciously on new leaves, especially those of American elm, crab apple and a few other kinds of trees. The larvae are at times sufficiently numerous to cause extensive defoliation. Arboreal birds of many species, as well as insectivorous small mammals, lizards, and arthropods are provided with an abundant food supply until the larvae mature. Species that prefer tall grass — the eastern cottontail, Hudsonian jumping mouse, western harvest mouse, cotton rat, marsh hawk, slender glass lizard, blue racer, common garter snake, and prairie king snake — have all increased, as have some brush-loving species, notably the long-tailed chat and least vireo. Otlier species that prefer open situations — the Plains harvest mouse, Plains pocket gopher, killdeer, lark sparrow, collared lizard. Great Plains skink, prairie skink, six-lined racerunner and ornate box turtle — have dis- appeared from the Reservation or have become relatively scarce and localized. Species of birds not regularly present on the Reservation, but which sometimes come in large numbers, and have important effect on the ecology, are the starling, the grackle, the red-wing black- bird, the robin and the flicker. The crop of wild grapes constitutes a major food source upon which large visitant flocks of the robin subsist for periods of weeks in late autumn. Although thousands of species of plants and animals occur on the Reservation, a relatively few species are so much more abundant than others that they are the kinds usually noticed by the casual observer; they make up most of the biomass and control the flow of energy through the ecosystem. Some of the most important are the following: American elm black walnut honey locust osage orange shagbark hickory chestnut oak black oak dogwood white-tailed deer eastern cottontail cottonrat prairie vole Producers crab apple wild currant coralberry redbud avens wliite snakeroot awnless brome Missouri goldenrod flowering aster Kentucky blue-grass fox grape Japanese chess switchgrass little bluestem Indian grass Primary Consumers katydids moths ( Oecanthus niveus ) ( Apantesis phalaerata ) (Orcheluuum vulgare) {Orfhnsia ohirina) (Orchelimum nigripes) (Orthosia hibisci) 90 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. deer mouse wood mouse harvest mouse juuco tree sparrow mourning dove grasshoppers (Melanophis bivittatiis) ( M. differentialis ) ( M. fcnnir-rtihrum ) crickets (Grylhts assimilis) ( Nemohius fasciatus ) eastern mole short-tailed shrew downy woodpecker black-capped chickadee tufted titmouse red-eyed vireo yellow-billed cuckoo ornate box turtle five-lined skink ring-necked snake bullfrog cricket frog chorus frog ( Neoconoce))luihts rohitstus) ( Conocephahis ) camel cricket ( CeutJiopIiilus paU-dus ) cicadas ( Magicicada ca-ssini) ( Tibicen pruinosa ) roach ( Farcohlatta holUana) beetles ( Scohjtus multistriatus) (PhyUophaga sp.) Secondary Consumers ground beetles (Colosoma scrutator) (PasiinacJius sp.) ( Harpalus caliginosus ) ants {Caniponotus herculancus) {Lasius niger) (Crematogaster sp.) wheel -bug ( Arilus cristatus ) spiders (Argiopc auranfia) ( OrtJiosUi ruhescens ) ( Palaeacrita vernata ) (Melanolophia signataria ) termites ( Reticulitermes claripennis) leaflioppers {ErytJironeura sp. ) ( Exitianus obscurinervis) ( AgaUia constricta ) earthwonn ( AUolobophora caliginosa ) ( Argiopc trifasciata ) (Neoscona benjamina) ( Micrathena sp. ) (Singa pratensis) ( Tetragnatha laboriosa ) ( Agelenopsis naecia ) {Dapanus mirus) ( Lycosa aspersa ) (Lycosa rabida) (Scliizocosa crassipes) ( FrontineUa pyrainilcla) coyote red fo.x raccoon Dom'uuint Predators opossum horned owl barred owl red-tailed hawk marsh hawk black rat snake copperhead The Rockefeller Experimental Tract In 1956, through the generosity of the late Mr. John D. Rocke- feller, Jr., an upland tract of 160 acres in Jefferson County, con- tiguous on its south edge with the western half of the Reservation, was acquired by the University as the Rockefeller Experimental Tract, which is administered through the Museum of Natural His- tory. Need for such an area on which experimentation was per- missible, had been acutely felt. Along its west edge the Rockefeller Tract contains an 11.4 acre strip of virgin tall-grass prairie, and an additional 107 acres now is regenerated prairie, but with impover- Natural History Reservation 23 ished and eroded soil because of former abuse by cultivation and overgrazing. The northern part of the quarter-section, on hilltops and north slopes, has about 27 acres of forest, somewhat different from any on the Reservation. Two species of climax hardwoods found there, mossy-cup oak and basswood, are almost absent from the Reser\'ation. The southeastern part of the quarter-section, a hilltop and south-facing slope, has a brushy and scrubby woodland. In the spring of 1957 the formerly cultivated and pastured parts of the Rockefeller Tract were sown to seeds of four native prairie grasses, namely big bluestem, little bluestem, Indian grass and switchgrass. The seed was obtained from several sources but was thoroughly mixed, and applied with a mechanical seeder. After several years the tall grass clumps had become well established over the area, but not in uniform stands as anticipated. After each filling of the seeder, the relatively heavy rounded seeds of switch- grass had fed through rapidly, while the light fluffy seeds of little bluestem passed through much more slowly with Indian grass and big bluestem somewhat imtermediate. As a result, these kinds of grasses tended to be arranged in strips with almost pure stands of switchgrass or little bluestem in some places. Big bluestem did not thrive on the impoverished and eroded old fields and its stands were sparse from the start. Indian grass also was much less abun- dant than switchgrass and little bluestem. Differences in limiting factors between the latter two grasses became increasingly ap- parent, in the nineteen-sixties; in low areas where soil was deeper, with silt deposits, switchgrass grew in tall dense stands, to the exclusion of most other vegetation, whereas on sloping ground es- pecially where the soil was poor and eroded, little bluestem domi- nated, but in relatively sparse stands. In 1961 and 1964 the tract of original prairie was burned off. In 1962 the eastern part of the regenerated prairie was divided into four parallel north-to-south experimental strips of approximately 15 acres each, and the eastern- most strip was burned off in early April. A second strip was main- tained as a control and a third was fenced and lightly grazed in summer. These same treatments were repeated in 1963 and 1964, and in these years a fourth strip was mowed in July. By 1964 the experimentally burned strip was almost a pure stand of the dominant prairie grasses, with increasing percentages of big bluestem. The mowed strip was somewhat similar. The grazed strip and especially the control had brush encroaching, mostly dogwood and sumac. In 1957 several species of animals characteristic of open places. 24 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. including the Plains pocket gopher, Plains harvest mouse, horned lark, lark sparrow, loggerhead shrike, and sparrow hawk were prominent on the Rockefeller Tract. In the following years as tall prairie grasses became reestablished, each of these animals became scarcer and eventually disappeared from the area. Meanwhile prairie loving animals formerly scarce or absent came into promi- nence on the area — notably the dickcissel, yellowthroat, marsh hawk, prairie vole, blue-racer and prairie king snake. Although the Rockefeller Experimental Tract is administered with somewhat different policy than the Natural History Reservation, the two areas supplement each other in some respects, and many field studies encompass parts of both areas. Research Over the past 16 years dozens of research projects covering a wide range of subjects have been carried on at the Reservation. Most research has been done by staff members and graduate stu- dents in the departments of Zoology, Botany, Entomology, Geog- raphy and Geology. Many graduate students have carried on re- search on the area for doctoral or master's dissertations. In some instances persons not directly connected with the University of Kansas, have carried on research — either as individuals or repre- senting organizations such as the Soil Conservation Service. Several undergraduate students from the University and even high school students have, with guidance, carried on research projects on the area. Although nearly all studies on the Reservation have been con- cerned with "pure research," many may eventually have practical applications, sometimes unforeseen. Kinds of animals that are most abundant and conspicuous have been given most attention and often these are species which have important effects on human affairs. Some are important as game (the cottontail, gray squirrel, fox squirrel), some are agricultural pests (voles, cotton rat), some are involved in public health problems (mosquitoes, chiggers, and venomous snakes), and some are destroyers of pests (red-tailed hawk, blue-racer). All of these kinds have been intensively studied on the Reservation. With the accumulation and synthesis of data over periods of years, findings that have practical applications emerge. For instance, the roles of certain kinds of animals as car- riers and transmitters of human diseases are clarified, and means of "biological control" of certain undesirable species, such as weeds or rodent pests, are revealed. Natural History Reservation 25 Most of the studies by the writer and his co-workers have been autecological, that is to say the studies have been concentrated on particular species in attempts to determine interrelationships of those species with their physical environment and with their plant and animal associates. Vertebrate animals have been given the most attention. An important aspect of a study of this type is the capture and distinctive marking of individual animals, wliich thereby may be recognized if they are recaptured, or even seen again on a later occasion. From one to 4000 individuals of each of 97 species (10 amphibians, 23 mammals, 26 reptiles, and 38 birds) have been marked. The method of marking varies, of course, according to the kind of animal being studied. Information regarding the growth, movements, and longevity of individuals representative of their species are thereby obtained. One of the simplest and most effective methods of marking is by clipping toes to half length, sometimes in combination with ear-clipping. Ordinarily no more than two toes (these on different feet) are clipped on each individual. After amputation heaHng occurs rapidly and without noticeable handicapping of the animal in most instances, but the metliod needs to be used with discretion on some kinds of animals that have toes specialized for climbing or digging. Thousands of small mammals, Hzards and frogs have been marked by toe-clipping. Birds have been marked with serially numbered metal leg bands issued by the U. S. Fish and Wildlife Service, sometimes supplemented by color bands for sight recognition without recapture. Certain kinds of mammals, notably rabbits and raccoons, have been marked with numbered metal ear tags. Snakes have been marked by clipping the serially arranged subcaudal and ventral scales, leaving permanent scars to yield dis- tinctive formulas. One of the major research efforts by the writer and his graduate students has been directed to a study of small mammals. Grids of hve-traps spaced at intervals of 10 to 75 feet have been estab- hshed on various parts of the Reservation and Rockefeller Tract and some of these have been maintained as much as ten years. Mammals captured have been individually marked by toe- and ear-clipping and many have been recaptured from one to 50 times revealing much concerning their natural history. For a few such as the eastern wood rat and pine vole it was found far more ef- fective to set traps in carefully selected sites, rather than in grids. Table 2 summarizes the information accumulated from many thou- sand catches for several of the most common species. 23 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. In the autumn of 1949 and for the following 14 years, wire fun- nel traps were set along hilltop limestone outcrops to catch snakes where they concentrated in search of suitable hibernacula, and all those captured were marked individually by clipping the sub- caudals and certain other scales. In 1957 and each summer there- after, lines of funnel traps were maintained also in grassland and brush on various parts of the Reservation and Rockefeller Tract. The following table summarizes the information gained from thousands of records including hundreds of recaptures for six of the common species. For all but the ring-necked snake "Number of individuals recorded" refers almost entirely to indi- viduals marked and released, but the majority of the 4846 ring- necked snakes were measured, sexed and released without marking, and some of these ring-necked snakes were caught on neighboring areas rather than on the Reservation or Rockefeller Tract. Seven species of lizards are known from the Reservation; five of these have been intensively studied on the area, by use of funnel traps similar or identical to those employed for snakes along with other special types of traps and capture techniques, and with indi- vidual marks based chiefly on toe-clipping. Table 4 shows some of the findings concerning these common lizards. It is noteworthy that three of these five species have been almost or entirely elimi- nated by successional changes on the area; the fix e-lined skink and glass snake are abundant. Studies by Greenberg (1952), Kardos (1954), Loomis (1956) and Wolfenbarger ( 1953 ) have revealed the extensive chigger fauna of the Reservation (22 species in all) and clarified their taxonomy, life histories, and possible bearing on public health problems. Chiggers are the larvae of trombiculid mites, all of them ectoparasites of vertebrate hosts although in the nymphal and adult stages they are free-living predators. The many genera and species differ in habitats, seasonal timing of life cycles and type of host preferred. They differ in host specificity; for instance, the common pest chigger that annoys man, Tromhicula alfrcddugesi, has been recorded from 83 species of vertebrates in Kansas whereas some other kinds are more or less specific to bats (T. fiventei), squirrels (T. fitchi), lizards (Acomatocarus arizonensis) , and frogs (Hanneman'.a eltoni). Studies on several genera and many species of the native bee fauna by Professor Charles D. Michener and his students, H. V. Daly, Alvaro Willie, Alvin Shinn, and Ellen G. Ordway, and by Professor Karl Stockhammer, have revealed a wide arrav of life Natural History Reservation 27 -^^^— ^.- — V ,— ^ aj -t^ OOlO — S) c3 ^ IC C3 lO lO t-1 -a c o -?* la > ^ cocoes 1 1 L-r c; c c^i c c t^ t^ » X X ^ rt ?- bc - c j: b£ en ~ rt itPi — : ^ X 3 1 CO CO — — -t o; X c ^"■Ti X oj _:: S" ^ >o ■ — ' V - O fa£— ^ CO CO c^) c^i ^ T" l^ CG S< - ~~-a — - t^ r^ a}"3 a. ^ OJ >- X o X rt oj (N 0 o3 o ^■>J'o CO C^) 'bo bo O CO CO 1— t t^ c 1 ^ a= _r '^- X 1 , '-0 CO X c o x" ^ t- X ■ ►— H 03 0 > T LO Ci t^ "" 3 s 00 CO 0 0) C OJ CO CO C I> t^ >.. - X (N " -tJ X -H r— 1 "m _aj .^ lO oc "3 > .2 5 ~ 5 c I~- lO 1 ca O CO -t-= 0) 'C -, 4^ ^ lO ^ c aa '3 :S5 2 j] >3 03 0) -H 00 Z^ <^' '£ X _ X -kJ 0 (N :3 2 rt OJ > T— ■ O -f o " rt C 0 lO tn c3 oj 2. aj O Tf o CO t- 0 0 10 o ^, >.w^ X CO ^^ (M bc J: Tf i-H TS ^_^ -a 1|_ rf ^ ^ ^^ O V i o .-A 3 flj O ,^\ 1 "rt 5 1 CO CC CO o CO (N L.0 ~~^ X X "0 X = '= = ^51 S3 > ^ OQ t^ CC^J " 0) cc ■s^^ a; _w OJ t- iM -f -f '^ CO ^.^-^ X Tf r— X 00 0) — M 3 13 rJ 00 ^ X X O -o r X s 3 0; 1 T CO 1 1 CO ^ S--S s tT Q 1^ gj > r—i CO 00 o^ - a-^ o -f rt ■ • CC l> >iW X ^ '^ »— " 1-H X 1>- f— 1 (U tc ^ — ^ „ 3 lO ^ £ 2 /-. uO -f 1 s 02 HI 1 T o ^^ C C 4 % X X il s 05 c3 ^^ > iC -t ^ (N ^ 00 J5 c3 ci " > wJ 1 ^ X ti £, -I . i=l C x CD •- Z3 -X ■ • c o o ?§ X Z 0 7i bC X -^ rf. 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CO tu to 3 0. " hO bl 0 03 C >i 3 3 .£ _C a -^ 3 o3 O CO 2 !Z b£ ]> C o3 0. CO faj Q, ^ •n -(J 3 3 ^ o 03 o . 1 ^ 8 Is o3 o 03 o -13 c3 O 'I IB "3 IE c c K oi a: c £ Ah H -a o c 3 -♦-3 .a c =^ faC 3 h-5 £ a Natural History Reservation 29 or- O "* 1 1 [ 1 lOt^ 00 >c 00 iM ^ lO od lO 1 CO 1-H t^ -^ ■^ ■ -H .— 1 (N(N »-H CO T-H t— 1 oo I I 9(14 6(21 CO -* lOCO T— 1 iCCO co<^^ bC bC o CO (N r— ( 1-H 8.1) 10.3) 1 1 coco >. 3 coco a _D ^^ o; oi CO 00 00 T^ 'tl 00 S. o 00 00 00 ^ CO -^ C lO IN ^ '^ o3 bC bC s a o bC o T3 o 3 T3 O O 30 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. < e < < ■z < o h a > z U H z o en Q N z o o U z o h < > r t" tc Q >• Z K W H S M > u Z Z OS u u z o u en O z c z o 1/J u CQ 0) C IB a c c 03 c a c o ■-V.S > 02 t- aJ oi b o S m +3 . S-t^ — > 3^ 03 rasshopp nd cricke eetles CD 11 ^ OJ S ST 1-H s .:!! o w bC o3^ CO o CD C^ cc tn Sac O. K o3 O 0) tC' — w _:; — ; bC o o ., to S CJ O I •g^ 03 ^ bb >. 3 aj-T" -<^ is o CO -a o - ^ -fi CO -^ CO *-( b£ 03 o3 O 3 03 0) -c ■5. CO iC co" 05 CO (M 03 C O m 02 C o » o3 i 0 a) V « £ c 3 o3 t-, ^ 03 a -0 0 a c C3 ^^ 03 0 a a> 0 a .S a; > I o c CO 10 hC bC to bC C .3 .3 c 03 O I- 03 b£ I-:; ^ b£ c 11 bC lO on (r4 0 1 0 >o *— 1 y—i 1 (N 0 ■^ C^ CO f—t 00 CO ^-^ 0 0 1—1 lO 00 CD 00 fN 10 1-^ 00 t^ (N 05 Oi CO 00 IM C CO 00 I IM CD CO c3 3 bJC 5 o CM CO d a > I 3 o c 03 i - s s "^ — o3 b£ H^; < Natural History Reservation 31 iC Ol CO iC o o 00 — o CO CC CC OS 'h o • N ^ o= (M lO IC (M CC ^ ^_,_^ ■b Of I?: o c: O r- _c ■^ rt -C 0. •♦^ c lO cc QC CO ^ C' -t iC >* C CC ^- c^ OS iH e3 a; >> ^^ — •' 2 2^ "« 3 2; 1 K a a h c 0 ^ CC o > Hi o c: M >, .? 0. hi £ c £ a. t £ 3 0. £ £ C3 1 a bj C C O h-" S O ;? h 1 32 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. histories, with various stages from those that are truly solitary to those that have developed a social organization of some com- plexity. Dr. A. Ralph Bmr and his student, Harlan L. MacMillan, made a survey of the mosquito fauna on one part of the Reservation — the headquarters' area. Their studies revealed a surprisingly large number of species (several dozen), and also revealed a suc- cession of species during the season, with each common species prominent for a relatively brief span of days or weeks. Professor Robert E. Beer and his students have studied the mite fauna of the Reservation, by taking extensive samples with Berlese funnels from various parts of the area. As in most other areas mites have here been shown to play an important role in the over- all ecology, by feeding as primary consumers, scavengers, predators and parasites. In a survey of the Reservation's spider fauna undertaken by the present writer in 1948 and completed in 1963, with the generous assistance of several leading araneologists, including Dr. Willis J. Gertsch, Dr. Herbert Levi, Mr. Wilton Ivie and Dr. Allen Brady, who identified specimens, a list of 192 species of 114 genera and 21 families was compiled from the area. Spiders as a group are important predators on small animals, especially on the insects. The common large orbweavers, wolf spiders, and funnel-web weaver destroy the larger kinds of insects in great numbers. How- ever, other smaller kinds of spiders, which are less conspicuous but far more abundant, and especially some of the minute erigonids, may have even more important effects on the area's ecology through their destruction of insects. Mr. John George has studied the taxonomy, seasonal cycles, habitats and hosts of the kinds of ticks occurring on the Reservation. The cottontail rabbit is an important carrier of ticks on the area. Mr. Alwan J. El-Wailly studied the fleas of the Reservation and found 15 species, with some kinds of mammalian hosts carrying several species. Dr. James Chilicott, as a graduate student, studied the arthropod fauna in nests of the prairie vole. These nests, situated several inches underground in damp soil, harbor many species of in- vertebrates, some of which are parasites on the voles, whereas others are scavengers feeding on the decaying vegetation of the nest, and still others are predators that feed upon species of the two groups mentioned first. Dipterous flies are especially prominent in the nest fauna. Natural History Reservation 33 Dr. John M. Legler from 1953 to 1957 studied the ecology and population dynamics of the box turtle, Terrapene ornafa, at the Reser\ation and at Damm Farm 12 miles west of Lawrence. The turtles were found to prefer shortgrass areas that were heavily grazed. Females lay clutches averaging 4.7 eggs. Late May and June is the principal nesting season, but some females lay second clutches in July. Males become sexually mature in eight to ten years, on the average, and females in ten to eleven years, but the total growing period extends over 15 to 20 years and the longevity is estimated as approximately 50 years. Box turtles are active for approximately six and one-half months, in the warmer part of the year. They prefer a body temperature of approximately 30° C. Insects that are associated with cow dung, especially scarabaeid and carabid beetles, are the chief food source, but larvae of noctuid and arctiid moths, and grasshoppers (Melanophis) were also frequent items, and some plant food, especially mulberries, were eaten. Box turtles have home ranges averaging 5.6 acres, and in their season of activity usually travel 200 to 300 feet daily. In a period of years following 1948 box turtles became much less conspicuous on the Reservation, and by 1964 they were rarely found although they remained common on adjoining areas of pastureland. The removal from the area of cattle and other livestock, and the increase in rank herbaceous vegetation were factors which rendered the habitat less favorable for them. Dr. R. W. Frederickson surveyed the myriapod fauna of the Reservation and recorded 54 kinds of millipedes and centipedes. The former play an important role as scavengers reducing decay- ing vegetation, while the latter are important as predators on insects and other small animals. The molluscan fauna of the Reservation was surveyed by Leonard and Goble (1952) in 1948 and 1949 soon after the policy of pro- tection was inaugurated, and was reexamined several years later by Fitch and Lokke (1956). A total of 25 species of snails and two clams were found, but some of these were represented only by old shells, their populations having been eliminated by the successional changes that have taken place. The terrestrial and freshwater mollusks of the area are sensitive indicators of environ- mental conditions, and also play significant roles as primary con- sumers in their ecosystems. It is intended that the Reservation be maintained permanently as a natural area and a research station for ecological studies. Each study that is completed provides background for further 3—8140 34 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. studies by which the complex web of ecological interrelationships between species in the local biotic community may be worked out. A large number of species are involved, and no end is in prospect for the studies. For insects especially there are hosts of species to be dealt with, most of them still not studied eco- logically either on the Reservation or elsewhere. For vertebrates the situation is somewhat different, as the number of kinds present is relatively small, and of these some are present irregularly, or are so uncommon that they are not suitable subjects for aute- cological studies. Many of those that are most common and most important in the local ecology have already been the subject of intensive investigation. This is not to imply that opporunities for research on vertebrates have decreased. Every species studied merits further more in- tensive investigation, but new approaches and more refined tech- niques are needed. Undoubtedly future studies will rely to a greater extent on instrumentation. A beginning in this direction was made in the winter of 1964-65, in a collaborative study by Dr. Hampton W. Shirer and the writer, by attaching thermisters on or within various reptiles hibernating under natural conditions, in order to ascertain their tolerances for low temperatures and the nonnal temperature ranges within their hibernacula. Description of the Area The Universitv of Kansas Natural Historv Reservation is situated on the northern edge of the Kaw River Valley, where level or gently sloping bottomland interdigitates with plateaulike upland. The "cuesta" (Schoewe, 1949:282) is a prominent physiographic feature created by a nearly horizontal stratum of limestone, ap- proximating the 1,040-foot contour. It underlies the shallow glacial till soil of the hilltops, but with an exposed edge occurring as a more or less continuous escarpment, be^ow which the teiTain slopes, steeply at first, and then more gradually, to the level of the valley floor. The upper stratiun of the limestone is the Plattsmouth mem- ber of the Oread limestone, of late Pennsylvanian age, in the Shawnee group of the Virgilian Series (Moore, Frye and Jewett, 1944:181-182). Its exposed edge is in places almost covered and concealed by deposited soil, but for most of its length it may be seen as an outcrop from 18 inches to three feet, or in places more than six feet, high. The face of this outcrop is deeply fissured and cracked, with numerous jagged boulders and small rocks, in process of breaking away or strewn over the steep slope below. Natural History Reservation 35 Oxerhanging rocks of the ledge shelter the soil beneath from rain and snow, and partly shade it with the result that it is nearly devoid of vegetation. Many kinds of animals depend on this ledge more or less for shelter. Another member of the Oread limestone, the Toronto limestone, is a conspicuously hard lower stratum about 20 feet below the level of the hilltop and its ex- posed edge also occurs more or less continuously along the upper slope. Upper and lower outcrops are separated by usually 50 to 300 feet of steep wooded slope, with a soil mantle, but with quanti- ties of loose rock. The lower outcrop often appears as a dark brown^ smooth, vertical rock face, from se\eral inches to three feet or more in height. Below this lower outcrop the hillside continues to slope steeply for a few yards, but then in most places, levels off to a natural terrace. The terrace follows the contour of the hillside and is often 30 to 50 feet wide. In places there are other terracelike formations farther down the hillsides, and these may be nearly horizontal, or may slope gently to join the upper terrace. Both upper and lower outcrops total approximately 4.5 miles in length on the Reservation, and 1.1 mile on the Rockefeller Experimental Tract. Below the lower outcrop the hillsides slope at more gradual gradients to the valley floor. Virtually all the hillsides, from the cuesta top to the valley floor, and small areas of upland adjacent to the upper ledge are wooded. On the Reservation this woodland comprises approximately 338 acres. The remaining 252 acres of the Reservation are open land of which about 116 acres was formerly under cultivation. Plant and Animal Life 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 primi- tive conditions, the trees growing at present seem to be secondary inxaders. They include groves and isolated trees of elm, honey locust, walnut, and osage orange, and an occasional red haw, hack- berry, 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. 36 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. SW 1/4 SECTION 33 TOWNSHIP MS RANGE 20E • RESERVATION HEADQUARTERS SECTION 4 TOWNSHIP I2S RANGE 20E Figure 1. Contour map of The University of Kansas Natural History Reserva- tion and the Rockefeller Experimental Tract. Natural History Reservation 37 ^ .INDEN^^^-""' ^:FaD^"^'' ^"; "STRIP" i ^ :;fire.-.:-":::..; FIELD... ■"":. MOWED grazed!"-3j;;^ STflP'^^^v^URNED"' mm^m4^r:': ■•■'''<;trip c ' X ->"■• ^STRIP" ^i^LD-^p;!^'^':?!-.;:": '- ;;;::;„, xfield"y„; C,,r"r.:FIELD.,"^ ■%^ , vii'^N^/OODS / - tr - '.»*-.V''t.V i!mt\ FOREST ROAD _cJ til//. BRUSH k SCRUB TALL- GRASS PRAIRIE ^ PASTURE GRASS WEEDY OLD-FIELD STREAM FENCE MRbulgeIrove te^r:,fe^nE^ ^Ismwrn^^ ^M^- ; ;:.-^^ANGLE^-.7^"^'^^Y<^ •'^' ^itsppR WOODS? IFIELD Jr ./WOOD^^i^- •^:-.-.-,/-- .^'--.lFIELD:f'^-tM^?.^ - ^^~"~?4--WEED FIFin?- ::srwMSftJ!.C ^■'S^^cS^i: SQUARE :l(>:' ,.. WOODS ^y^< -.^A-FiFin F.: <-?^<« - „^OAT FIELD.- }-V"^^^ ^ »'■ ^ ■ Figure 2. Map of The University of Kansas Natural History Reservation and the Rockefeller Experimental Tract showing named subdivisions of these areas, and showing the distribution of each of several major types of vegetation. 38 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. PLATE I Fig. 1. Stiuiip of large cottonwood on liilltop at southeast comer of Bottom Woods on April 1, 196.5. At the time of a previous photograph, taken from the opposite side of the tree in December, 1954 (see Fitch and McGregor, 1956, pi. 7, lower figure) the tree was living and was the largest on the Res- ervation. Hollows in the trunk sheltered manv kinds of animals. V j^^^ \ >'' .■->»■ "x. s,-'*.-^. -^ Fig. 2. South part of pond looking east from dike on January 16, 1965, when surface was frozen to a depth of several inches. Cattails and other succulent riparian or shallow water plants, abundant at the time of a previous photograph November 1, 1951 (Fitch, 1958b, pi. 5, fig. 1), had been almost entirely removed by muskrats, but small willow trees had appeared in the water, changing the aspect of tlie pond. Natural History Reservation 39 PLATE II Fig. 1. Large American elm at nortlieast corner of Ditch Field, still thriving on January 15, 1965, although many others had died presumably as a result of introduced diseases. Since April 2, 1955 ( see Fitch and McGregor, 1956, pi. 8) the tree had become noticeably more massive, and the thicket around it (an open field in 1948) became denser. Fig. 2. Scene on west-facing slope of Skink Woods on January 15, 1965, from outcrop of Toronto Limestone, looking east up rocky hillside dominated by chestnut oaks. The scene remained similar to that in a photograph taken July 22, 1950 (Fitch, 1954a, pi. 1, fig. 2; 1958b, pi. 3, fig. 1) except that a large oak log in the foreground on the earlier date had completely disinte- grated by the latter date and become incorporated in the soil. 40 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. PLATE III ^z \ 1 ... ■*. Fig. 1. Diversion ditch at edge of woodland 100 yards norda of Reservation headquarters, looking west. The formerly barren eroded banks of the ditch provided favorable habitat for species that prefer open spaces ( see Fitch, 1958b, pi. 2, fig. 2) but by January 15, 1965, as shown here, the bottom and banks of the ditch had grown up to thickets of woody vegetation. Fig. 2. Gully in regenerated tall-grass prairie of formerly culti\ated and eroded area of Road Field and High Field, looking north, on April 1, 1965. An earlier photograpli from almost the same spot, taken on November 9, 1954 (see Fitch, 1958b, pi. 4, fig. 1) shows the young trees much smaller and less prominent, especiallv the three cottonwoods in center. Woody vegetation has spread rapidly onto this grassland area and has dominated some parts of it, and erosion has almost stopped in the gully, now choked with dense grass. Natural History Reservation 41 PLATE IV Fig. 1. Northwestern corner of the Reservation, ironi county road, looking nortli on January 15, 1965. The pale-colored central triangular area is thick brush, which has encroached onto a slope that was still dominated by big bluestem and other prairie grasses at the time of a previous photograph in the summer of 1951 (Fitch, 1963a, pi. 22, fig. 2), after cessation of burning and cessation of mowing in 1947. ** ' '••|f»«^^**.A* • 1 .-4; ,. ■^*T' '»•'«>• '^^ . Fig. 2. View of Dry Field looking east down gully toward edge of Point ^^'oods on April 1, 1965. Vegetation in the field is sparse and consists of such weedy plants as three-awn grass, common lespedeza, and goldenrod. In the interval since November 9, 1954 (Fitch, 1958b, pi. 4, fig. 2) erosion had largely ceased in the gully but little successional change had occurred. 42 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. PLATE V Fig. 1. UL'i,Li\atiou licadqumtris \i(.\\cd troiii north on Januai)' 15, 1965. Since the time of a previous photograph on October 13, 1951 (see Fitch, 1952, pi. 1, fig. 1) several large elms around the buildings died and fell, and brusln- vegetation appeared in the field in foreground. l-'iG. 2. House Field looking W'SW toward Reserx ation licadciuartcrs on Janu- ary 15, 1965. Since the time of a previous photograpli on March 3, 1956 (see Janes, 1959, pi. 45, fig. 1) much brush had sprung up in the field. Natural History Reservation 43 PLATE VI Fig. 1. K. U. zoology class studying the home Hfe of small animals, October 13, 1951. Photograph by Robert Rose. Fig. 2. Professor and stiidtnl mark a Hudsonian jumping mouse for release October 13, 1951. Photograph by Robert Rose. 44 Univ. Kansas Mus. Nat. Hist. Miscl. Publ. PLATE VII Fig. 1. Gully in soutlieastern part ot Reservation on April 10, 1963. Since the time of an earlier photograph in March, 1956 (see Fitch and McGregor, 1956, pi. 7, upper figure) the gully eroded more deeply, and a large elm that had been undercut and had fallen, disintegrated and disappeared. ■^ •,> ^ ■i- ». ' , A -' l'"iG. 2. Same gulK' as shown ah;,\i', ticm a point scxeral liundred \ards farther south, looking north, upstream, on April 11, 1965. Since the time of a pre- vious photograph, from approximately the same point, on November 7, 1949 (see Fitch, 19581), pi. 6), configuration of the banks and gully bottom changed somewhat; erosion slowed and banks gained more brushy vegetation, and had less extensive bare areas. Natural History Reservation 45 PLATE VTTT Fig. 1. Same gully as shown in Plate 7, viewed from south boundary of Res- ervation on January 15, 1965. Since the time of a previous photograph from approximately the same point in November, 1950 (see Leonard and Goble, 1952, pi. 99, f ), the channel deepened, banks became more brushy, and a part of the bank enclosed in a loop of the creek (left center) was partly eroded away, so that trees in background were better exposed to view. Fig. 2. Small cedar tree on hilltop (Grove Field), which by January 15, 1965, when this photograph was taken, had the upper part of its trunk girdled and killed by deer rubbing their antlers against it. By 1965 such sign of the activities of deer had become conspicuous on the area. 46 Univ. Kansas Mus. Nat. Hist. Miscl. Pub. Subdixisions on a vertical scale, might be recognized as follows: the upper limestone outcrop at the hilltop; the usually steep slope strewn with rocks, between the upper and lower limestone out- crop; the lower limestone outcrop; an almost level terracelike forma- tion 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 \\'oodland is noticeably different from that in other situations, and of more xeric aspect. The climax species of oaks and hickories 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 dog- wood and plum 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 nearly alike 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. In the following alphabetical lists scientific names are grouped in several major categories that in general correspond with taxo- nomic groupings. Nevertheless, for the plants it seemed desirable to separate the trees and other woody species from the herbaceous types. All the species listed have been definitely identified. For some, notably the birds and a few other vertebrates, sight records are the sole basis for inclusion, but all such sight records were Natural History Reservation 47 made by well-trained and eompetent observers and hence there can be little doubt that the species actually occur on the area. For most of the plants and invertebrates listed, the records are sub- stantiated by preserved specimens in the small museum collection at the Reservation headquarters. In some respects the records for different species are not strictly comparable. Of the birds, for in- stance, relatively few are permanent residents, some are present only in winter, others only in summer, others on migration, and some only as accidental wanderers, perhaps recorded only on one occasion. For each of several species of birds the sole record is based on an individual flying over the Reservation but not known to have stopped there. Status of each of the vertebrates is dis- cussed in a previous publication (Fitch, 1958b). The lists represent the combined data from the Reservation and the Rockefeller Tract. A few species, notably the large conspicuous woodland snails, Triodopsis olbolobris and Anguispira aJtcrna, and the spider, Loxosceles rechisa, occur on the Rockefeller Tract but stop short of the Reservation. Many other species have been recorded on the Reservation but not on the Rockefeller Tract. It may be expected that most of the lists will be enlarged as a result of further fieldwork adding previously undetected species. But in some instances species formerly present have been eliminated (notably certain aquatic mollusks whose habitat has deteriorated as a result of erosion), and other instances may be expected. Some groups of invertebrates, notably the chiggers, mollusks, moths and spiders, have been made the subjects of intensive in- vestigations on the area and specimens have been submitted to leading specialists for authoritative determinations. For other groups such as orthopterans and beetles, study has been more casual and determinations have been by research assistants, comparing material with the reference collection in the Snow Entomological Museum, with only occasional help from specialists. Also, some determinations were made several years before the present report was prepared. Hence, some of the names used may not be in line with the most recent taxonomic revisions. Some major groups of invertebrates on the area, such as the hemipteran bugs, wasps, leafhoppers, mites (other than chiggers), crustaceans and nematode worms have been given little attention or none at all, and hence no lists are available for these groups. To this extent, the treatment herein is unbalanced, and major syne- cological investigations on the area must await detailed studies of such neglected groups. 48 Univ. Kansas Mus. Nat. Hist. Miscl. Pub. Acer negundo Acer saccharinum Ailanthus altissima Ampelopsis cordata Asimina triloba Ceanothus ovatus Carya ovata Celastrus scandens Celtis occidentalis Cercis canadensis Cornus drummondii Cornus amomum Corylus americana Crataegus crus-galli Crataegus mollis Diospyros virginiana Euonymus atropurpureus Fraxinus americanus Gymnocladus dioica Juniperus virginiana Madura pomifera Trees, shrubs and woody vines Menispermum canadense Morus rubra Parthenocissus quinquefolia Platanus occidentalis Populus deltoides Prunus americana Prunus angustifolia Prunus hortulana Prunus nana Prunus persica Pyrus ioensis Quercus macrocarpa Quercus marilandica Quercus muehlenbergii Quercus prinoides Quercus borealis Quercus stellata Quercus velutina Rhamnus lanceolata Rhus aromatica Rhus copallina Rhus glabra Rhus radicans Ribes missouriense Rosa arkansana Rosa setigera Rubus argutus Rubus flagellaris Salix amygdaloides Salix eriocephala Salix interior Salix nigra Sambucus canadensis Smilax hispida Staphylea trifolia Symphoricarpos orbiculatus Tilia americana Ulmus americana Ulmus rubra Vitis cinerea Mtis vulpina Xanthoxylum americanum Floioering plants (herbaceous dicotyledons) Abutilon theophrasti Acalypha virginica Achillea lanulosa Agastache nepetoides Agrimonia parviflora Agrimonia pubescens Amaranthus hybridus Amaranthus tamariscina Amaranthus torreyi Ambrosia artemisiifolia Ambrosia trifida Ammannia coccinea Ampelamus albidus Amorpha canescens Amorpha fruticosa Amphiachyris dracunculoides Androsace occidentalis Anemone canadense Anemone virginicus Antennaria campestris Apocynum cannabinum Arctium minus Arisaema dracontium Arabis canadensis Arabis perstellata Asclepias meadii Asclepias purpurascens Ascelpias stenophylla Asclepias syriaca Asclepias tuberosa Asclepias verticillata Aster ericoides Aster simplex Aster drummondii Baptisia leucantha Baptisia leucophaea Bidens bipinnata Bidens frondosa Bidens polylepis Cacalia atriplicifolia Campanula americana Capsella bursa-pastoris Cassia fasciculata Cassia marilandica Chenopodium album Chenopodium boscianum Chenopodium hybridum Chrysanthemum leucanthemum Circaea canadensis Cirsium altissimum Cirsium discolor Comandra richardsiana Commelina communis Convolvulus arvensis Conyza canadensis Corydalis flavula Croton capitatus Croton monanthogynus Cycloloma atriplicifolium Cynoglossum officinale Dasj^stoma macrophylla Datura stramonium Delphinium virescens Descurainia sophia Desmanthus illinoiensis Desmodium glutinosum Desmodium illinoense Echinacea pallida Ellisia nyctelea Erigeron philadelphicus Erigeron strigosus Eupatorium altissimum Eupatorium purpureum Eupatorium rugosum Euphorbia coroUata Euphorbia dentata Euphorbia heterophylla Euphorbia maculata Euphorbia marginata Euphorbia missurica Euphorbia serpens Fragaria virginiana Galium aparine Galium circaezans Galium tinctorium Gaura biennis Gaura parviflora Geum canadense Geum vernum Geum virginianum Geranium carolinianum Gnaphalium obtusifolium Hedeoma hispida Helenium autumnale Helianthus aunuus Helianthus laetiflorus Helianthus grosseserratus Helianthus tuberosus Hypericum perforatum Natural History Reservation 49 Ipomoea hederacea Iva ciliata Kuhnia eupatorioides Lactuca canadensis Lactuca floridana Lactuca scariola Lamium amplexicaule Laportea canadensis Lappula echinata Lepidium densiflorum Lespedeza capitata Lespedeza striata Liatris aspera Liatris squarrosa I.inum sulcatum Lithospermum arvense Lithospermum canescens Lithospermum incisum Lobelia siphilitica Lobelia spicata Lomatium foeniculaceum Ludwigia palustris Lycopus americanus Lythrum alatum Aledicago lupulina Melilotus alba Melilotus officinalis Mentzelia oligosperma Mimulus alatus Mirabilis nyctaginea Mollugo verticillata IMonarda fistulosa Najas guadalupensis Nepeta cataria Oenothera biennis Oenothera speciosa Onosmodium occidentale Opuntia humifusa Osmorhiza longistylis Oxalis dillenii Oxalis stricta Flowering plants (concluded) Parietaria pennsylvanicus Penthorum sedoides Penstemon cobaea Petalostemon candidus Phlox divaricata Physalis heterophylla Physalis longifolia Physalis macrocarpa Physalis missouriensis Ph3^salis pumila Phytolacca americana Phryma leptostachya Plantago major Plantago virginica Podophyllum peltatum Polanisia trachysperma Polygonum aviculare Polygonum coccineum Polygonum hydropiper Polygonum hydropiperoides Polygonum lapathifolium Polygonum pennsylvanicum Polygonum punctatum Polygonum scandens Potentilla simplex Potentilla norvegica Potamogeton foliosa Potamogeton nodosus Prenantlies aspera Prunella vulgaris Psoralea esculenta Ranunculus abortivus Ratibida pinnata Rhamnus lanceolata Ruellia humilis Ruellia strepens Rumex altissimus Sagittaria latifolia Salvia azurea Salvia reflexa Sanicula canadensis Scrophularia marilandica Sida spinosa Silene stellata Silphium laciniatum Silphium perfoliatum Solanum carolinense Solanum nigrum Solanum rostratum Solidago altissima Solidago angusta Solidago nemoralis Solidago rigida Solidago ulmifolia Specularia perfoliata Stenosiphon linifolius Taraxacum erythrospermum Teucrium canadense Thlaspi arvense Tradescantia bracteata Tragia urticifolia Tragopogon pratensis Trifolium pratense Trifolium repens Triosteum aurantiacum Triosteum perfoliatum Urtica procera Verbascum blattaria Verbascum thapsus Verbena baldwini Verbena canadensis Verbena hastata ^'erbena stricta Verbena urticifolia Viola rafinesquii Viola papilionacea Viola pedatifida Viola sororia Xanthium strumarium Zizia aurea Grasses, rushes, sedges, lilies (monocotyledons) Agrostis alba Andropogon gerardi Andropogon scoparius Andropogon virginicus Andropogon ternarius Aristida oligantha Bouteloua curtipendula Bromus inermis Bromus japonicus Bromus purgans Carex annectens Carex bicknellii Carex blanda Carex brevior Carex emoryi Carex gravida Carex hyalinolepis Carex vulpinoidea Cyperus ferruginescens Dactylis glomerata Diarrhena americana Digitaria ischaemum Digitaria sanguinahs Echinochloa crusgalli Eleocharis macrostachya Elymus virginicus Eragrostis capillaris Eragrostis cilianensis Eragrostis spectabilis Erythronium albidum Glyceria striata Hypoxis hirsuta Juncus interior Koeleria cristata Leersia oryzoides Muhlenbergia schreberi Muhlenbergia sylvatica Panicum capillare Panicum clandestinum Panicum lanuginosum Panicum latifolium Panicum virgatum Paspalum ciliatifolium Phleum pratense Poa pratensis 4—8140 50 Univ. Kansas Mus. Nat. Hist. Miscl. Pub. Grasses, rushes, sedges, lilies (concluded) Polygonatum canaliculatum Scirpus atrovirens Scirpus lineatus Scirpus validus Scleria triglomerata Allarthonia caesia Amphilonia lanuginosum Bilimbia trachona Candelaria concolor Caloplaca aurantiaca Cladonia capitata Graphis scripta Lecanora dispersa Lecanora iowensis Amblystegiella confervoides Amblystegiella serpens Amblystegiella varium Anomodon attenuatus Anomodoii rostratus Astomum muhlenbergianum Atrichum angustatum Barbula cruegeri Brachythecium oxycladon Frullania eboracensis Frullania inflata Setaria faberii Setaria glauca Setaria viridis Sisyrinchium campestre Sorghastrum nutans Lichens Lecanora subfusca Lecanora versicolor Lecidia botryosa Parmelia borreri Pertusaria lecanina Physica adglutinata Physica caesia Physica obscura Physica orbicularis Mosses Brachythecium salebrosum Bryum argenteum Chamberlainia acuminata Entodon cladorrhizans Entodon compressus Entodon seductrix p]urhynchium serrulatum Fabronia wrightii Fissidens taxifolius Grimmia apocarpa Homomallium adnatum Liverworts Frullania riparia Riccia hirta Spartina pectinata Tridens flava Tripsacum dactyloides Typha angustifolia Typha latifolia Physica stellaris Psora demissa Psora russellii Rinodia dakotensis Sarcogyne pruinosa Verrucaria calciseda Verrucaria calkinsiana Xanthoria candelaria Hygroamblystegium irriguum Hygroamblystegium orthocladum Leptodictyum trichopodium Leskea gracilescens ]\Iinum cuspidatum Orthotrichum pumilum Orthotrichum pusillum Physcomitrium turbinatum Thelia asprella Riccia lamellosa Botrychium obliquum Botrychium virginianum Cystopteris protrusa Blarina brevicauda Canis latrans Cryptotis parva Didelphis marsupialis Geomys bursarius Lasiurus borealis Marmota monax Mephitis mephitis Microtus ochrogaster Microtus pinetorum Mus musculus Mustela frenata Ferns Ophioglossum engelmannii Mammals Myotis sj). Xeotoma floridana Odocoileus virginiana Ondatra zibethicus Peromyscus leucopus Peromyscus maniculatus Procyon lotor Rattus norvegicus Reithrodontomys megalotis Reithrotlontomys montanus Woodsia obtusa Scalojnis aquaticus Sciurus carolinensis Sciurus niger Sigmodon hispidus Spermophilus franklini Spilogale putorius Sylvilagus floridanus Synaptomys cooperi Urocyon cinereoargenteus Vulpes vulpes Zapus hudsonius Natural History Reservation 51 Accipiter cooperii Accipiter gentilis Accipiter striatus Actitisi macularia Agelaius phoeiiiceus Aegolius acadicus Aix sponsa Ammodrammiis savannanim Anas acuta Anas carolinensis Anas discors Anas platyrhynchos Anser albifrons Aquila chrysaetos Archilochus colubris Arciea herodias Asio otus Aythya affinis Aythya collaris Bartramia longicauda Bombycilla cedrorum Botaurus lentiginosus Branta canadensis Bubo viiginianus Buteo jamaicensis Buteo lagopus Buteo lineatus Buteo platypterus Butorides virescens Capella gallinago Caprimulgus carolinensis Caprimulgus vociferus C'arpodacus purpureus Cathartes aura Centurus carolinus Certhia familiaris Chaetura pelagica Charadrius vociferus Chen caerulescens Chen hyperborea Chondestes grammacus Chordeiles minor Circus cyaneus Cistothorus platensis Coccyzus americanus Coccj'zus erythrophthalmus Colaptes auratus Colinus virginianus Columba hvia Contopus virens Corvus brachyrhynchos Cj'anocitta cristata Dendrocopos pubescens Dendrocopos villosus Dendroica coronata Dendroica fusca Dendroica petechia Dendroica striata Dendroica virens Birds Dumetella carolinensis tlmpidonax flaviventris Empidonax minimus Empidonax traillii Eremophila alspestris Elreunetes mauri Kluphagus carolinus Falco columbarius Falco mexicanus Falco sparverius Fulica americana (leothlypis trichas Grus canadensis CJuiraca caerulea Haliaetus leucocephalus Ilirundo rustica Hylocichla fuscescens Hylocichla guttata Hylocichla minima Hylocichla mustelina Hj'locichla ustulata Icteria virens Icterus galbula Icterus spurius Junco hyemalis Lanius ludovicianus Larus pipixcan Megaceryle alcyon Melanerpes erythrocephalus Melospiza georgiana Melospiza lincolnii Melospiza melodia Micropalama himantopus Mimus polyglottos Mniotilta varia Muscivora forficata Myiarchus crinitus Xuttallornis borealis Xycticorax nycticorax (^porornis formosus Otus asio Pandion haliaetus Parula americana Parus atricapillus Parus bicolor Passer domesticus Passerculus sandwichensis Passerella iliaca Passerherbulus caudacutus Passerina ciris Passerina cyanea Pelecanus erythrorhynchos Petrochelidon pyrrhonota Pheucticus ludovicianus Philohela minor Pipilo erythrophthalmus Piranga olivacea Piranga rubra Podilymbus podiceps Polioptila caerulea Pooecetes gramineus Porzana Carolina Progne subis Quiscalus quiscula Rallus limicola Regulus calendula Regulus satrapa Richmondena cardinalis Riparia riparia Sayornis phoebe Seiurus aurocapillus Seiurus motacilla Seiurus noveboracensis Setophaga ruticilla Sialia sialis Sitta canadensis Sitta carolinensis Spatula clypeata Sphyrapicus varius Spinus pinus Spinus tristis Spiza americana Spizella arborea Spizella pallida Spizella passerina Spizella pusilla Stelgidopteryx ruficollis Strix varia Sturnella magna Sturnella neglecta Sturnus vulgaris Telmatodytes palustris Thryomanes bewicki Thryothorus ludovicianus Totanus flavipes Toxostoma rufum Tringa solitaria Troglodytes aedon Troglodytes troglodytes Turdus migratorius Tj-rrannus tyrrannus Tyrrannus verticalis Tyto alba Vermivora celata Vermivora peregrina ^'ermivora ruficapilla Vireo bellii Vireo flavifrons Vireo gilvus Vireo griseus Vireo olivaceus Vireo solitarius Wilsonia canadensis Wilsonia pusilla 52 Univ. Kansas Mus. Nat. Hist. Miscl. Pub. Xanthocephalus xanthocephalus Agkistrodon contortrix Carphophis amoenus Chelydra serpentina Chrysemys picta Cnemidophorus sexlineatus Coluber constrictor Crotalus horridus Crotaphytus coUaris Diadophis punctatus Acris crepitans Ambystoma tigrinum Bufo americanus Camptostoma anomalum Acalymma vittata Acanthocinus sp. Achmaeodera pulchella Adalia frigida Agabus disintegratus Agabus stagninus Agonoderus pallipes Agroiconota bivittata Alaus oculatus Alobates pennsylvanica Anisoclavia quatuordecimguttata Anomoea laticlavia Anomola marginata Anoplum cinerascens Anthonomus signatus Aphodius concavus Aphodius distinctus Aphodius stercorosus Aphodius terminalis Araecerus sp. Astenus cognatus Bassareus clathratus Batyleoma suturale Bembidion sp. Berosus sp. Brachynus alterninus Buprestis rufipes CalHgrapha scalaris Calligrapha similis Calosoma cahda Birds (concluded) Zenaidura macroura Zonotrichia albicolUs Reptiles Elaphe guttata Elaphe obsoleta Eumeces fasciatus Eumeces obsoletus Eumeces septentrionalis Lampropeltis caUigaster Lampropeltis doliata Lampropeltis getulus Natrix sipedon Amphibians Bufo woodhousei Gastrophryne olivacea Hyla versicolor Pseudacris nigrita Fishes Ictalurus melas Lepomis cyanellus Beetles Calosoma scrutator Cantharis bilineatus Cantharis carolinus Carabus sylvosus Casnonia pennsylvanica Cassida sp. Ceratoma punctata Ceratoma trifurcata Ceratomegilla fuscilabris Ceratomegilla maculata Chauliognathus marginatus Chelymorpha cassidea Chion cinctus Chirida guttata Chlaenius bicolor Chrysochus auratus Cicindela belfragei Cicindela punctulata Cicindela repanda Cicindela scutellaris Cicindela sexguttata Clivinia bipustulosa Clivinia dentipes Colaspis sp. CoUops sp. Coniontellus sp. Conotrachelus seniculus Copelatus glyphicus Cotinus nitida Cryptocephalus venustus Zonotrichia leucophrys Zonotrichia querula Ophisaurus attenuatus Pituophis catenifer Scincella laterale Storeria dekayi Tantilla gracilis Terrapene Carolina Terrapene ornata Thamnophis sirtalis Virginia Valerias Rana catesbeiana Rana pipiens Spea bombifrons Xotropis lutrensis Pimephales promelas Cryptocephalus quadruplex Cryptorhynchus tristis Cycloneda sp. Deloyala guttata Dermestes sp. Desmocerus palliatus Diabrotica duodecimpunctata Diabrotica longicornis Diabrotica vittata Dineutes sp. Disonycha glabrata Disonycha sp. Disonycha xanthomelaena Dorchaschema alternatum Eburia quadrigeminata Elaphrus sp. Enochrus perple.xus Enoclerus sp. Epicaerus imbricatus Epicaerus sericata Epicaerus unicolor Epicauta occidentalis ■ Epicauta pennsylvanica p]picauta sericans Epicauta unicolor Eucanthus lazarus Eumolops colossus Euphoria fugida Natural History Reservation 53 ' Reservation. Univ. Kansas Publ., Mus. Nat. Hist. 11:63-323. 1960. Autecologv of the copperhead. Univ. Kansas Publ., Mus. Nat. Hist. 13:85-288. 1963a. Natural history of the racer. Coluber constrictor. Univ. Kansas Publ., Mus. Nat. Hist. 15:351-468. 1963b. Natural history of the black rat snake (Eluphe o. obsoleta) in Kansas. Copeia, 1963(4) :649-658. 1963c. Spiders of the University of Kansas Natural History Reservat'on and Rockefeller Experimental Tract. Univ. Kansas Mus. Nat. Hist., Misc. Publ. no. 33:1-202. 1965. An ecological study of the garter snake, Thamnophis sirtalis. Univ. Kansas Publ. Mus. Nat. Hist., 15:493-564. ** Fitch, H. S., and Fitch, V. R. 1955. Observations on the summer tanager in northeastern Kanas. \\'il- son Bull., 67:45-54. ' Fitch, H. S., and Lokke, D. H. 1956. The molluscan record of succession on the Uni\ersity of Kansas Natural History Reservation. Trans. Kansas Acad. Sci. 59(4): 442-454. " Fitch, H. S., and McGregor, R. L. 1956. The forest habitat of the University of Kansas Natural Historv Reservation. Univ. Kansas Publ., Mus. Nat. Hist. 10:77-127. " Fitch, H. S., and Packard, R. L. 1955. The coyote on a natural area in northeastern Kansas. Trans. Kan- sas Acad. Sci. 58:211-221. " Fitch, H. S., and Rainey, D. G. 1956. Ecological observations on the woodrat, Neotoma floridana. Univ. Kansas Publ., Mus. Nat. Hist. 8:499-533. * Fitch, H. S., and Sandidge, L. L. 1953. Ecology of the opossum on a natural area in northeastern Kansas. Univ. Kansas Publ., Mus. Nat. Hist. 7:305-338. * Fredrickson, R. W. 1954. Myriapoda of the University of Kansas Natural History Reservation. Unpublished thesis. University of Kansas Librar\'. * Freiburg, R. D. 1951. An ecological study of the narrow-mouthed toad (MicroJiyla olivacea) in northeastern Kansas. Trans. Kansas Acad. Sci. 56:376- 386. " Greenberg, B. 1952. A review of the New World Acomatacarits (Acarina, Trombiculi- dae). Ann. Ent. Soc. Amer., vol. 45, no. 3, Oct. 25, 1952:473-491. Natural History Reservation 59 Hall, E. K., Anderson, S., Jones, J. K., Jh., and Packahd, H. L. 1957. Vernacular names for North American mammals north of Mexico. Univ. Kansas Mus. Nat. Hist., Misc. Puhl. no. 14:1-16. Harlan, J. R., Speake, E. B. and Reece, M. F. 19.56. Iowa fish and fishing. Iowa State Cons. Com., 377 pp., 22 pis. ' Hartmann, E. L. 1956. The br>oplnte flora of the University of Kansas Natural History Reservation. Trans. Kansas Acad. Sci. 59:57-70. * Janes, D. W. 1959. Home ranges and movements of the eastern cottontail in Kansas. Univ. Kansas Publ., Mus. Nat. Hist. 10:553-572. Johnston, R. F. 1960. Directory to the bird-life of Kansas. Univ. Kansas Mus. Nat. Hist., Misc. Publ. no. 23:1-69. * Kardos, E. H. 1954. Biological and systematic studies of the subgenus Neotrombiciila (Genus Tromhicula) in the Central United States (Acarina, Trombiculidae). Univ. Kansas Sci. Bull. 36 pt. 1 (4):69-123. * Kramer, C. L. 1956. Lichens of the University of Kansas Natural History Reservation. Trans. Kansas Acad. Sci. 58-519-524. * Legler, J. H. 1960. Natural history of the ornate bo.x turtle, Terrapene onuita ornata Agassiz. Univ. Kansas Publ., Mus. Nat. Hist. 11:527-669 * Leonard, A. B., and Coble, C. R. 1952. Mollusca of the University of Kansas Natural History Reservation. Univ. Kansas Sci. Bull. 34:1013-1053, 2 pis. * LooMis, R. B. 1956. The chigger mites of Kansas (Acarina, Trombiculidae). Univ. Kansas Sci. Bull. 38:1195-1444. Macgovvan, K., and Hester, J. A., Jr. 1962. Early man in the New World. Anchor Books, Doubleday and Co., Inc. N. Y. xxiii + 333 pp. * Martin, E. P. 1956. A population study of the prairie vole (Microttis ochrogaster) . Univ. Kansas Publ., Mus. Nat. Hist. 8:361-416. Martin, P. S. 1958. Pleistocene ecology and biogeography of North America. Amer. Assoc. Adv. Sci., Publ. 51:405. * McFarland, a. N. Spring moths ( Macrolepidoptera ) of the University of Kansas Natural History Reservation. In press. * Michener, C. D. and Willie, A. 1961. The bionomics of the primitively social bee, Lasioglossuni incon- spicuiim. Moore, R. C, Frye, J. C, and Jewett, J. J. 1944. Tabular description of outcropping rocks in Kansas. State Geol. Surv. of Kansas, Bull. 52, 1944 Reports of studies. Part 4, pp. 137-212, figs. 1-9. * Ordway, E. 1961. The biology of Augochlorella, a green sweat bee in Kansas. Proc. North Central Branch Entom. Soc. Amer., 16:17. 1964. Sphecodes pimpenellae and other enemies of Augochlorella. Jour. Kansas Ent. Soc, 37(2) : 139-152. 60 Univ. Kansas Mus. Nat. Hist. Miscl. Pub. * Packard, R. L. 1956. The tree squirrels of Kansas: ecology and economic importance. Univ. Kansas Mus. Nat. Hist., Misc. Publ. no. 11:1-57. Parks, G. S. 1854. ' The Tourist" Column. The Kansas Herald of Freedom, 1 (no. 1) Wakarusa, Kansas Terr., October 21, 1854. " Rainey, D. G. 1956. Eastern wood rat, Neotoma floridana: life history and ecology. Univ. Kansas Publ. Mus. Nat. Hist., 8:535-646. " Rettenmeyer, C. W. 1963. Behavioral studies of army ants. Univ. Kansas Sci. Bull., 44: (9): 281-465. 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 Publ. Co., Lawrence, Kansas (10th ed. ) xi + 438 pp. "Sandidge, L. L. 1953. Food and dens of the opossum (Didelphis virginiana) in north- eastern Kansas. Trans. Kansas Acad. Sci. 56( 1) :97-106. Schoewe, W. W. 1949. The geography of Kansas. Part II. Trans. Kansas Acad. Sci. 52:261-333. Smith, H. M. 1956. Handbook of amphibians and reptiles of Kansas. 2nd [revised] edition. Univ. Kansas Mus. Nat. Hist., Misc. Publ. no. 9:1-356. ** Stewart, P. L. 1960. Lung-flukes of snakes, genera Thamnophis and Coluber, in Kansas. Univ. Kansas Sci. Bull. 41, no. 8:877-890. * Stockhammer, K. a. 1961. Aspects of the life history of the sweat bee, Augochlora p. parva (Say). Proc. North Central Branch Entom. Soc. Amer., 16:17-18. Taft, R. 1950. The great sandy desert. Trans. Kansas Acad. Sci., 53:441-442. * TOMASELLI, R. 1958. Plant communities of the western half of the University of Kansas Natural History Reservation (1952-63). Casa Editrice Rengo Cortina, Pavia (Italy), 27 pp., many maps. Wedel, W. R. 1959. An introduction to Kansas archeology. Smiths. Inst. Bur. Ethnol- ogy, Bull. 174:xvii -f- 723 pp. ' Wolfenbarger, K. a. 1953. Systematic and biological studies on North American chiggers of the genus Tromhicula, subgenus Eutrombicula (Acarina, Trombi- culidae). Ann. Ent. Soc. Amer., vol. 45, no. 4:645-677. Wormington, H. M. 1957. Ancient man in North America. Denver Mus. Nat. Hist., Popular Ser. no. 4:xviii -f 322 pp. TransmiUed May 7, 1965. 3 2044 093 361 590 Date Due