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An Upland Forest Continuum in the Prairie-Forest Border Region of Wisconsin Author(S): J. T. Curtis and R. P. Mcintosh Source: Ecology, Vol

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An Upland Forest Continuum in the Prairie-Forest Border Region of Wisconsin Author(S): J. T. Curtis and R. P. Mcintosh Source: Ecology, Vol An Upland Forest Continuum in the Prairie-Forest Border Region of Wisconsin Author(s): J. T. Curtis and R. P. McIntosh Source: Ecology, Vol. 32, No. 3, (Jul., 1951), pp. 476-496 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/1931725 Accessed: 01/07/2008 17:00 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=esa. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit organization founded in 1995 to build trusted digital archives for scholarship. We work with the scholarly community to preserve their work and the materials they rely upon, and to build a common research platform that promotes the discovery and use of these resources. For more information about JSTOR, please contact [email protected]. http://www.jstor.org AN UPLAND FOREST CONTINUUM IN THE PRAIRIE- FOREST BORDER REGION OF WISCONSIN1 J. T. CURTIS University of Wisconsin, Madison and R. P. MCINTOSH Middleburv College, Middlebury, Vermont INTRODUCTION generously placed his files on forest corn- It has been said that the desirable position at our disposal; to Professors order of ecological research in a given Grant Cottam, H. C. Greene, N. C. Fas- region is first, the study of communities, sett, R. S. Muckenhirn, M. L. Partch, second, the study of the individual spe- and S. A. Wilde for valued advice and cies, and last, the study of the habitat consultation; to Messrs. Orlin Anderson, (Yapp 1922). Sociological investiga- and R. S. Brown for indispensable aid tions of constituent species and their in the collection of field data; to Miss numerical relations should precede aute- Margaret Gilbert for aid in the computa- cological studies if the latter are to have tion; to Messrs. A. M. Fuller, Neil maximum meaning. This sequence Harrington and Fred Wilson for help in many should lead to a greater efficiency in re- the location of stands; and to the Serv- search, in that an initial knowledge of any members of the Soil Conservation natural groupings of species may enable ice, the Regional Foresters of the Wis- the autecology of the groups to be studied consin Conservation Department and simultaneously, with a resultant saving in the advisers of the Junior Wisconsin time and travel expenses. The grossly Academy of Sciences, who so willingly condi- inadequate state of autecological knowl- imparted their knowledge of local edge of even our most common species tions in their regions. The authors are indicates that the current haphazard also indebted to Professor K. J. Arnold, method of attack is faulty; if aggrega- of the University Computing Service, for tions of species occur together under aid in the statistical phases of the work. in similar environmental conditions in na- The taxonomic nomenclature used Manual ture, they could be studied in a systematic this paper follows that of Gray's program. A long-term investigation of of Botany, 8th edition, 1950. the communities of Wisconsin has been LITERATURE REVIEW underway since 1946 with a view to de- to the vegeta- termining the existence and the floristic The earliest references were nature of such aggregations (Curtis and tion of southwestern Wisconsin the explorers and pioneer trav- Greene 1949). The present paper deals made by 1698. For with the upland hardwood forests of the elers in the years following border of the state the most part, they contained only gen- p)rairie-forest region remarks about un- and their interrelations with each other eral descriptions, with first account a and with certain physical factors of the usual species. The by was that of Increase environment. Deep appreciation is ex- competent botanist in l)ut he dealt pri- presse(l to Professor P. B. Whitford, hlio A. Lapham 1852, marily with the distribution of individual 1 This work wvas supported in part by the species. The first description of plant Research Committee of the radiatete School of communities as such was given by T. C. the University of Wisconsin from funds sup- plied by the Wisconsin Alumrni Research Chamiberlainin 1877. This eminent geol- Foundation. ogist, whlo was later to exert such great 476 July, 1951 UPLAND FOREST CONTINUUM IN WISCONSIN 477 influence on H. C. Cowles at the Univer- and the changes which have taken place sity of Chicago, was at that time the State in them since settlement. He attributed Geologist of Wisconsin. He distin- major importance to fire in the mainte- guished three forest communities in nance of oak openings and to cessation southern Wisconsin, but made clear that of fire as the cause of the subsequent de- no abrupt line of demarcation existed be- velopment of the present closed canopied tween them, with repeated statements stands. illustrating "a gradual transition from The most comprehensive paper on the one to the other" and "an almost imper- forests of southern Wisconsin was by ceptible merging of one into the other." Whitford (1951). Although this study His three communities were the "oak was primarily concerned with the relation group" including Quercus alba (white between the clone size of herbs and the oak), Q. mctcrocarpa (bur oak), Q. rubra age of the forest stands, considerable at- (red oak), and Q. velutina (black oak), tention was devoted to the establishment with Populus tremuloides (aspen), Carya of a method for classifying the stands. ovata (shagbark hickory), Prunus sero- In the course of the investigation, Whit- tina (black cherry) and Pyrus ioensis ford studied a considerable number of (wild crab apple) as lesser members; stands and presented detailed data on the "oak and maple group" in which Acer tree, shrub, and herb composition. He saccharum (sugar maple) was most con- arranged his series of 28 stands into spicuous but Quercus alba and Q. rubra three groups on the basis of shade toler- were still common, with Acer rubrum ance of the tree species. He assigned (red maple), Ulmus rubra (slippery each species to one of three tolerance elm), and Tilia americana (basswood) in classes, and then determined the relative small numbers; and the "maple group" fraction of all trees in a stand which be- with A. saccharum dominant, but with longed to each of these classes, on a sum- Tilia amnericanca,Ulmus rubra, and Ostrya mation basis. This method resulted in virginiana (ironwood) as characteristic a sequence of stands from those com- and abundant associates. He comments posed entirely of intolerant species to about the "oak and maple group" as fol- those made up mostly of tolerant species. lows: "This group is not characterized by Whitford divided his stands into three the exclusive presence of any prominent groups-"oak-hickory, intermediate, and plant but by a distinctive association of maple-basswood," but the division was plants common to several groups." made for pragmatic reasons and was not Bruncken (1900) also described three based on any internally apparent separa- forest associations, but only two were tion of the data into three groups. found on uplands the "hemi-xerophytic Studies in Illinois, Iowa and Minme- oak-hickory" and the mesophyticc bass- sota indicate that a series of forests simi- wood-maple." Additional observational lar to those in Wisconsin are found in at studies on the forests of the region were least portions of these states. No quan- reported by Pammel (1904), Marks titative investigations in northern Illinois (1942) and Stout (1944), but the first have come to our attention, but the gen- account in which quantitative data on eral descriptions of Fuller and Stras- actual composition were presented was baugh (1919), and DeForest (1921) that of Wagner (1947), who studied a indicate a series of forest types from stand in Jefferson County which fits into those dominated by Quercus and Carya Chamberlain's oak and maple group. to those dominated by Acer saccharum. Cottam (1949) investigated an oak In Iowa, the studies of Clark (1926) and woods in Dane County by quantitative of Aikman and Smelser (1938) show a methods and reported in detail on the succession from Quercus macrocarpa and historical development of oak openings Q. velutina in the initial stages through 478 CURTIS AND MCINTOSH Ecology, Vol. 32, No. 3 Q ucrcus rlibra-Tilia am1tericanastands to Carya ova/a, and Jitglans nigra reach the terminal forests of Acer saccharun their northern boundaries in a zone which and Tilia am erican-a. Daubenmire (1936) also includes the southern limits of such reported on the structure of the Acer- trees as Betfila lutea, A bies balsarnea, Tilia. community and on its relation to and Populls balsamnifera. This same the surrounding Qutercus forests in the zone marks the northward or southward "Big Woods" of Minnesota. extent of a great many herbaceous spe- While the results of the various inves- cies, both native and exotic; it also sepa- tigations in these four states are by no rates closely related varieties of single means in complete agreement, there does species, as in the case of Acer saccharunm appear to be a general uniformity of saccharum and its subspecies A. s. nigrum opinion as to the topographic distribu- (Desmarais 1948) and Braclyelytrum tion of the forests, with some type of erectau erectur and B.
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