peeeiiae aes Site SoS Aa CaS OAS yet pile cat er Grae ghey feasattetebshy per ty cee batt Aaah a Pheer ara eae ie aii miareiatanise fs oe if ‘ae seen citi beta ir mear i iS AS rh tat mes ks bi eu oe sh ore Renae i ME Hoe es rae oats eet, ee opens aos VEN Seve Was fra. patna ey alent ST) fare Era eae pe btn AoL tes Wil Gel Aes, SRS oa iaaaty si tieitn d Pian " et ine ears ret rit Lae oe case Salt a pene oe ate rah ate " ity cl ely tS te ae ats Depart ae Hea ee eat Os eu tH fy a SCOn anna aa Sea 7 eee Pye a i eee ena ee ra a fit rm earn retire a foe ie Fae rc 0 ie Beets ie a ey a ie es ie ee ih a ay ae Ee cian pea af ee) ans shell ser peas Maines tty tt eS yess gaia et arate pine ¥ ie Aces Ate ete ae eh ‘ set aioe ey Deaton . c 0 eee Peraave yrs i an ee x wert ihe eee is me as pa ey Lae bi ‘rare planation itn Ae Behe i Betas 8 a uae tae tat ea ects Abe dee " ig aeoae ae cae a he ed oem Ce he Ret yy ie & if ou & ny i ey 7 * By ild a en Fic: ipl el a ae i Uy iiusreteg st os mee ee i ra: i) re ie = Ly i =) New York State College of Agriculture At Cornell University Ithaca, N.Y. » Library Cornell University Library QL 751.A41 TA 3 1924 002 894 487 wi mann ORIGINAL FOREST AND PRAIRIE AREA IN ILLINOIS (AFTER BRENDEL AND BARROWS ) BULLETIN OF THE ILLINOIS STATE LABORATORY oF NATURAL HISTORY URBANA, ILLINOIS, U. S. A. STEPHEN A. FORBES, Pu.D., L.L.D., DIRECTOR Vou. XI. SEPTEMBER, 1915 ARTICLE IT. AN ECOLOGICAL STUDY OF PRAIRIE AND FOREST INVERTEBRATES BY CHARLES C. ADAMs, Pu.D. -—r & CONTENTS Pace 33 35 35 33 40-56 40 41 43 44 48 49 52 52 35 Deseription of 56-66 L 36 2, 37 3. jation, i 59 4. Lowland or ““second bottom, ”” red oak-elm-sugar maple wood- land association, Station IV, ¢..........-.-.....--..eeeeees 2 5. Supplementary collections from the Bates woods, Station IV.... 6 6. Small temporary stream in the south ravine, Station IV, d...... ci General characteristics of the gross environment. ................. 66-102 L Topography and soils of the State... - 66 2. Climatic conditions ................. 67 3. Climatic centers of imflmemee .......-. 0... ee ee eens 9 4, Relative humidity and evaporating power of the air............ val 5. Temperature relations in the opem and im forests.............. 83 6. Soil moisture and its relation to vegetation. .............--...- 86 7 Ventilation of land habitats......... 2.2.0.2... 0. eee ee 8B 8. The tree trumk as a habitat..... 2.2.2... cee eens 91 9. Prairie and forest vegetation and animal life................. 91 10. Sourees and réle of water used by prairie and forest animals. . . . 98 Animal associations of the prairie and the forest. ..-............... 0.000055 102-158 TL. Introduction ....... 2.2.2... 22.20 eee eee ee ee een eee ene eenees 102 IL The prairie association... ........ 2... 2... 2-0 cee ee ees 103 L Swamp Irie association. ...... 2.2... eee eee eee eee eens 103 2. The cottonwood commumity....... 2... 2... eee eee eee 105 3. Swamp-grass association... .... 2.2... 2.2... eee ee eee eee 107 4. Low prairie association. ... ~~... 2.2.0.2... ee eee eee 108 5. Upland prairie association. .................. 0.2... e eee eee 109 6. The Sokdago commumity............-..-- +... eee eee eens 109 7. Dry prairie grass association. ...............--......66- 111 8 A muilkweed commumity...................- 12 Tit. Relation of prairie animals to their environment, 113 1. The black soil prairie community... ...... 1144 2. The prairie vegetation commumity........................5-- ni7 4. Interrelations within the prairie association ng ITV. The forest associations.........ccsccsse cece vere er seen sree seeeeee 122 Ve Pntrodwetiony, sé cascdtoves anya oeeststaverdisis «chiding s six ate areutiaials ia Weta are 122 2. Dry upland (Quercus and Carya) forest association............ 124 3. Artificial glade community in lowland forest.........-.....0+5 125 4, Humid lowland (hard maple and red oak) forest association... . 126 5. Animal association of a temporary stream.......-..++eee-eeee 127 V. Relation of the deciduous forest invertebrates to their environment.... 128 1. Forest soil community. ........ ccc cece cece esac nace rcersces 129 2. The forest fungus community...........cceeeee ee ee cee eeees 135 3. The forest undergrowth community............ceeeeeeeeeeaee 138 4, The forest crown community..........ccceeeececeeeee ieinenes 139 5. The tree-trunk community............ OE LISS RINNE TG 142 6. The decaying wood community......... 0c cece ee eee eeeeeeee 148 7. Interrelations within the forest association.................5+ - 157 Eeologically annotated list :— : I, Prairie invertebratesic: ccdeciieecdse code etecwa needs eee ee eerie ee 158-201 Il. Forest invertebrates. ........ ccc esc e esc e eee eee eeee suse eestteees 201-238 BUDO BPA PY «ces: :cosicack. sx evenanie des “shi danny wien des gnas re lca ogee ae “td oc vec Nora ara ova tahesens 239-264 Articie Il—An Ecological Study of Prairie and Forest Inverte- brates. By Cuaries C. Apams, Pu.D. INTRODUCTORY In four generations a true wilderness has been transformed into the present prosperous State of Illinois. This transformation has been so complete that in many parts of the state nearly all of the plant and animal life of the original prairie and forest has been completely ex- terminated. Between the degree of change which has taken place in any given area and the suitability of that area for agriculture there has been an almost direct relation. Fortunately, however, for the preser- vation of prairie and forest animals, the state is not homogeneous, some areas being too hilly, rocky, or sandy for prosperous agriculture. The character and mode of transformation which has taken place in the past is instructive in several particulars because it serves to guide our anticipations as to the future of our fauna. The forested southern part of the state (see frontispiece) was first invaded by trap- pers and hunters, who began the extermination of the larger animals. These invaders were in turn followed by others who, with the round of the season, were hunters or farmers, and continued this exterminat- ing process, particularly in the clearings, which began to replace the forest. These pioneers, men of little wealth, possessed a combination of mental and economic habits which was the result of life in a for- ested country, and naturally they’settled in those places most like their former homes—within the forest or near the forest margin. From these settlements they looked out upon the prairies as vast wastes to be dreaded and avoided. As a result of this attitude toward the prai- ries, it required some time, even a new generation, some economic pressure, and a change of habits before the prairies were settled. Mean- while the northern part of the state was yet a wilderness; but, through the influence of the Great Lakes, as a route of communication with the populous East, a rapid invasion of settlers set in from that direc- tion. Though these settlers also came from a wooded country, they were more wealthy, settled upon a very fertile soil which was favorably located with regard to eastward communication, and they therefore progressed more rapidly than the less favored, more isolated southern invaders on the poorer soil; consequently they spread from the forest 34 to the prairie more rapidly than did the settlers in the South. ‘There thus developed two active centers of influence, each of which trans- formed the primeval conditions in the same manner and in the same direction toward an environment suitable for man. The forests and the upland prairie were first changed. ‘Then the fertile wet prairie was drained, so that today it has largely become either the hilly and rocky areas that survive as forests or the low periodically flooded tracts, and the undesirable sand areas which simi- larly preserve patches of sand prairie. All the changes are more rapid and complete upon fertile soil than upon the poorer soils in the southern part of the state. Such considerations as these will aid one in estimating the probable rate of future changes in different parts of the state, and will serve to show in what parts there is urgent need of local studies if ecological records are to be made before extinction of some forms is complete. A study has been made with the idea of reporting upon represen- tative patches of prairie and forest in a manner which would aid others in making similar local studies, and would at the same time preserve some records of the present condition of the prairie and forest. When this work was planned, we had no general or comprehensive discussion of the conditions of life upon the prairie and in the forest. For this reason a general summary of these conditions and a sketch of the gen- eral principles involved are given, so that the reader may gain some conception of the relation of the local.problems to those of a broader and more general character. A section for this report was prepared giving general directions for making such local studies, but later it was decided to publish this ‘ separately, in somewhat extended form, as a “Guide to the Study of Animal Ecology.”* ‘This volume should be regarded as intimately re- lated to this paper, and this report should at the same time be consid- ered as a concrete example of the procedure suggested in that “Guide” for ecological surveys. It will be observed that the study of the Charleston area here referred to has been conducted in much the same way as was my cooperative study of Isle Royale, Lake Superior, en- titled “An Ecological Survey of Isle Royale, Lake Superior” (’09), although certain aspects have been elaborated here which, for lack of time, were not treated there. The time devoted to the study of the Charleston area was also limited, but in the preparation of the report upon it use has been made of many years’ experience and a general knowledge of the prairie and forest. Without such a background *The Macmillan Co. 1913. 35 much greater caution would have been necessary in discussing many phases of the problem. ACKNOWLEDGMENTS The study of the Charleston area was carried, out with the coop- eration of the Illinois State Laboratory of Natural History, through its director, Prof. Stephen A. Forbes, and with the further coopera- tion of Professors E. N. Transeau and T. L. Hankinson, of the East- ern Illinois State Normal School, located at Charleston. Personally I am indebted to Professor Forbes for the opportunity of taking part in this study as the State Laboratory representative, and for the aid he has given in the illustration of the report. To Professor Transeau I am particularly indebted for the plant determinations, for lists of the plants, and for evaporation data. To Professor Hankinson I am under especial obligation for many specimens, which materially added to my lists, and for a large number of photographs. I am indebted likewise to my associates in this study for their hearty cooperation throughout the progress of the work. For the determination of entomological specimens I am indebted primarily to Mr. C. A. Hart, Systematic Entomologist of the State Laboratory of Natural History, who named most of the insects col- lected. For the names of certain flies I am indebted to Mr. J. R. Mal- loch, of this Laboratory. Others who have determined specimens are as follows: N. Banks (Phalangiida), J. H. Emerton (spiders), R. V. Chamberlain (myriapods), F. C. Baker (Mollusca), Dr. W. T. M. Forbes (lepidopterous larve), Dr. M. C. Tanquary (ants), Dr. M. T. Cook (plant galls), J. J. Davis (Aphidide), and Dr. A. E. Ortmann (crawfishes collected by T. L. Hankinson). I am indebted to the U. S. Geological Survey for photographs. Acknowledgments for illus- trations are made under text figures and in explanations of plates. GENERAL DESCRIPTION OF THE REGION AND LOCATION OF THE ECOLOGICAL STATIONS I, GeneraL DESCRIPTION OF THE REGION The town of Charleston, Coles county, Illinois, in the vicinity of which these ecologic studies were made, is situated on the Shelbyville moraine which bounds the southern extension of the older Wisconsin ice-sheet. ‘T'o the south of this moraine lie the poorer soils which char- acterize so much of southern IIlinois; to the north, upon the older Wis- consin drift, are some of the most productive soils found in the upper 36 Mississippi Valley. The economic, sociologic, political, and historical significance of the difference in the soils of these regions is funda- mental to any adequate understandng of man’s response to his ecolog- ical environment within this area. Some of the results of this differ- ence have long been known, but it is only in recent years that their general bearing has been adequately interpreted in terms of the en- vironment. Hubbard (’04) was the first, I believe, to show the sig- nificance of this difference in soils and its influence upon local eco- nomic problems. That such an important influence should affect one animal (man) and not others seems very doubtful, and yet in only one other case do we know that the lower animals respond to this ecologic influence. Forbes (’07b) has shown that certain kinds of fish found in streams on the fertile soils are wanting in streams on the poorer soil. To what degree the land fauna and the native vegetation respond to this distinction is not known, as this subject has not been investi- gated except agriculturally. Here, then, is a factor in the physical surroundings which should be reckoned with in any comprehensive study of the biotiq environment. In this portion of the state, on ac- count of the differences in the soil, the physical environment is prob- ably more favorable to certain organisms and less. favorable to others, ‘and consequently, to a certain degree, the environment selects, or fa- vors, some organisms. Through their activities and through other agencies of dispersal, the animals along the borders between the two soil types transgress these boundaries, and are therefore forced to respond to the new conditions and to adjust themselves, if possible. But the soil is not the only environmental influence which has pro- duced an unstable zone or tension line in this area. A second factor is the difference in the vegetation—the difference between the forest and the prairie. In all probability, Coles county was at one time all prairie, but the Kaskaskia and Embarras rivers, as they cut their valleys through the moraine and developed their bottoms, have led forests within the morainic border from farther south. The forests about Charleston have extended from the Wabash River bottoms. On account of the southerly flow of the Embarras through this county, the forest and prairie tension line is about at right angles to that produced by the differences in the soil. The forests have tended to spread east and west from the streams and to encroach upon the prairie, and thus to restrict its area more and more. The fundamental significance of the tension between the forest and the prairie has long been known within the state. It influenced its economic, social, political, and historic develop- ment as much as any other single factor during its early settlement. And just as Hubbard (’04) has shown the influence of soil upon man 37 within the state, so also has Barrows (’10) shown the influence of the forests and prairie upon the state’s development. While the influencé of the soil upon the animal life of the state is not so well known or es- tablished, the influence of prairie and forest upon the animals is univer- sally recognized, even though the subject has been given relatively little study by naturalists. A third leading agency is the influence of man, who has trans- formed the prairie and forest to make his own habitat. There are thus recognized in the Charleston region three primary environmental in- fluences: first, the relative fertility of the soil (this depending on the geological history) ; second, the kind of vegetable covering, whether prairie or forest (this probably depending largely on climatic condi- tions) ; and third, the agency of man. The general background of the Charleston region, then, ecologically considered, depends on the com- bined influence of five primary and secondary agencies, four of which we may call natural and one artificial. All these are different in kind and so independent that they tend toward different equilibria or dif- ferent systems of unity. Two of these are due to differences in the soil, two others to the character of the vegetation (whether prairie or forest), and the fifth, or artificial one, is due to man. Though the present report does not undertake to include all the problems centered here, as any complete study would, it is desirable to see the relation of our special study to the general problems of the region as a whole. The undulating plain about Charleston, formed as a terminal mo- raine, is broken along the small streams by ravines, which have cut a few hundred feet below the general level of the region as they ap- proached the larger drainage lines. The main drainage feature is the Embarras River, which flows southwest about two to three miles east of Charleston, in a narrow valley partly cut in rock. The wooded areas are mainly near the streams; the remainder of the area is under intensive cultivation. During the preliminary examination of the region, which was made to aid in selecting representative areag for study, it soon became evi- dent that the only samples of prairie which could give any adequate idea of the original conditions were those found along the different railway rights-of-way. Other situations, vastly inferior to these and yet a valuable aid in the determination of the original boundaries of the prairies, were the small patches or strips along the country roads. Most of the patches of prairie along the railway tracks represent the “black soil” type of prairie, which is extensively developed in this part of the state upon the “brown silt loam” soil” (see Hopkins and Pettit, ’08 : 224-231). Much of the region studied was originally wet prairie 38 (which, tae: Since Teen: drained), but some of the higher ground, formed by the undulation of the surface and surrounded by the black soil, is lighter im color and is well draimed. ‘Thus im the black soil areas there are both wet and well-drained tracts, and corresponding differ- ences im the habitats. The originally wooded and the present wooded areas east of Charleston, in the vicinity of the Embarras River, are in a region quite different from the prairie both im topography amd im soil, Here the re- Sie Zé tach mote pronounced, fin arciend of both the piticimity of the river and the greater development of the draimage limes, which have cut a few hundred feet below the general level of the country. ‘The tribu- tary valleys and ravines are mumerous and steep-sided, and im general are wooded, the density varying with the amount of clearimg done. Most of the soil of the wooded areas and along the bluffs is distinctly im color tham that of the black soil prairie, amd is “gray silt loam ” (Hopkins and Pettit, 08 : 238-242), though along the and the river bottom the soils are mixed im character. IL. Tar Ecotoacan Sraroxs In the study of am area or am amimal association of any comsidera- ble size two methods are available. Onme is to examine as much of the area as is possible and secure data from a very wide range of condi- tions. ‘This method is useful in obtaiming the general or broad featmres of a region or am association, though to a corresponding degree it must ignore local imfimences amd details, and by it most of the previous stud- ies mpon prairie animals hawe been made. It seemed, therefore, that m the present study a somewhat more imtensiwe method was desirable, particularly im view of the fact that the extinction of prairie and for- est is rapidly progressing. The method followed was to examime 2 large area im order to select a represemtative sample, and wpom the basis of this sample to make as intensive a study as time and circum- stamoes would permit. ‘Tis method has the advantage of making it possible to preserve at least some record of the local details; amd at the same time, to the depree that the selected area is a tre sample, it also gives the results a much wider applicatiom. The prairie samples examined were all along the rights-of-way, amd the forest was a secomd-growth woods om the bottoms and bluff of the Eimbarras River, om a farm belonging, at that time, to Mir. J. 1 Bates. Practically all of the observations here reported mpom were made during Ampust, 1910. The forest is a modified ome, but tt ap- pears to hawe been cut over so gradually that its comtimmity as a forest habitat was mot completely imternrmpted, although the cuttimg has prob- 39 ably seriously influenced many animals, particularly those which fre- quent mature forests, abounding im dead and dying trees and with an abundance of logs upon the ground im all stages of decay. Such con- ditions are the cumulative product of a fully mature climax forest. Of course the conditions have also been influenced by the extinction, or eee of the original vertebrate population of the corest. The different localities or regions examined are, for brevity and Precision, indicated by Roman mumerals ; the particular minor condi- tions, situations, or habitats, by italic letters. An effort has been made to indicate the location of the place studied with enough precision to enable students to re-examine the habitats at amy future time (PL 1). Se a ee eee eee meas oe ae ae ing the places studied. Had similar photographic records been made fifty years ago, they would have been of much value and imter- est to us im this study, im much the same way as fifty years hence this report will form a part of the very limited record of the conditions found at the present time. List of Ecological Stations, Charleston, Minois, August, 1910 Station L Prairie along the right-of-way of the Toledo, St. Louis and Western, or ““Clover Leaf’ B. B., between one and two miles north of Charleston: Section 2, Township 12 N., Range 9 B., amd 8. 35, T.13N, B.9 EB. (PLT) a Cord or Slough Grass (Sporting) and Wild Bye (Elymus) Asso- ciation. At mile-post marked ““Toledo 318 miles and St. Louis 133 miles’: S. 2, T. 12 N., B. 9 BE. b. Couch Grass (Agropyrom smithii) Association. The distance of two telegraph poles north of Station I, «, and west of the railway truck: $2, T. 12 BE. B.9 EB. ce. Wild Bye (Elymus) Association. East and north of the “Yard Limits” sigm: $2, T. 12 NL, B.9 B. (PL UO, Fig. 1.) dé Swamp Milkweed (Ascdlepias imcoraata) Association. North of first exst-and-west erosroad north of Charleston; east of railway track: &. 35, T. 13 N., B.9 B. A wet area. (PL Ii, Fig. 2; Pl. Til, Fig. 1.) €. Coneflower (Lepachys pinnata) and Bosin-weed (Silphium tere- binthinacewm) Association. Just north of the preeedimg Station ; east of raihway tradk: 8.35, T. 13 N, B.9 BE. (PL V.) f- Couch Grass (Agropyrom smithii) Association. West of railway track: $35, T.13 N., B.9 EB. Moist area. GQ Prairie Grass (Andropogon furcatus and A. virgimicus and Spo- robolus eryptendrus) Association, bordered by Swamp Milkweed (Asdepias imewrnata) and Mountain Mint (Pycmanthemum flex- 40 uosum). This formed the north boundary of the area studied: S. 35, T.13 N., R.9 E. (PI. III, Fig. 2; Pl. IV, Fig. 1 and 2.) Station II. Prairie area west of Loxa, Illinois. Right-of-way along the Cleveland, Cincinnati, Chicago and St. Louis, or ‘‘Big Four,” R. R.: Sections 10 and 11, Township 12 N., Range 8 E. a. From one half mile west of Loxa west to near Anderson Road, to telegraph pole No. 12330: S.11, T.12 N., R.8 E. (Pl. VI. and VIL.) b. Prairie at Shea’s: 8.17, T.12 N., R.8 E. c. Cord Grass (Spartina) Association. East of Shea’s: 8.17, T.12 N., R.8 E. Station III. Prairie east of Charleston. Right of way along the C. C. C. & St. L. R. R.: 8.12, T.12 N., R.9 E.; 8.5, 6, and 7, T.12 N., R.10 E. a. Rosin-weed (Silphium terebinthinaceum) Association. Just west of the place where the Ashmore Road crosses the Big Four track; about one mile east of Charleston: 8.12, T.12 N., R.9 E. b. Blue Stem (Andropogon) and Rosin-weed (Silphium terebinthina- ceum) Association. Three fourths of a mile east of the crossing of the Ashmore Road and the Big Four track: 8.6 and 5, T.12 'N., R.10 E. An area which grades from prairie into transitional for- est conditions. (Pl. VIII and IX.) Station IV. Bates Woods. On the east bluffs and bottom of the Embar- ras River, north of where the Cleveland, Cincinnati, Chicago and St. Louis, or Big Four, R. R. crosses the river. On the farm of J. I. Bates: 8.5, T.12 N., R.10 E. (Pl. X, Fig. 1; Pl. XI, XII, and XIII.) a. Upland Oak-Hickory Association (Quercus alba and Q. velutina, and Carya alba, C. glabra, and C. ovata.) Second-growth forest. (Pl. XII and XIII.) b. Embarras Valley and Ravine Slopes, with Oak-Hickory Associa- tion. c. Red Oak (Quercus rubra), Elm (Ulmus americana), and Sugar Maple (Acer saccharum) Association. Lowland or ‘‘second bot- tom,’’ Embarras Valley. (Pl. XIV; XV; and XVI, Fig. 1 and 2.) d. Small streamlet in South Ravine. This formed the southern bor- der of the area examined. A temporary stream. (Pl. XVII, Fig. 1 and 2.) DESCRIPTION OF THE PRAIRIE HABITATS AND ANIMALS J. Prarie Area NortH of CHARLESTON, StTaTION I This area includes patches or islands of prairie vegetation oc- curring along the right-of-way of the Toledo, St. Louis and West- 41 ern, or “Clover Leaf,” Railway, north of Charleston. The south- ern border began just beyond the area of numerous side tracks and ex- tended north of the first east and west cross-road for a distance of about one mile, to the place where the right-of-way is much narrowed and fenced off for cultivation. This is a strip of land through the level black soil area, which was originally composed of dry and wet prairie. The higher portions have a lighter colored soil, and the lower parts have the black and often wet soil which characterized the original swamp or wet prairie. The railway embankment and the side drain- age ditches have favored the perpetuation of patches or strips of these wet habitats ; the excavations for the road-bed, on the other hand, have accelerated drainage of the higher grounds. The soil taken from these cuts and heaped up on the sides of the tracks reinforces the surface relief noticeably in a region which is so nearly level. Through the depressions fillings have been made in building the railway embank- ment, and as a result the drainage has been interfered with in some places. The disturbances brought about by railway construction and main- tenance have greatly modified the original conditions, so that the prairie vegetation persists usually only in.very irregular areas, some- times reaching a maximum length equal to the combined. distance be- tween three or four consecutive telegraph poles—these poles are gen- erally about 200 feet apart. In breadth the area is usually less than the space between the ditch bordering and parallel to the road-bed or embankment and the adjacent fence which bounds the right-of-way, or about 40 feet. This entire right-of-way is about roo feet wide. In addition to these changes in the physical conditions, a large number of weeds not native to the prairie have been introduced, opportunities for this introduction being favorable, as railways traverse the entire area. In general, attention was devoted solely to the areas or colonies of prairie vegetation and their associated invertebrate animals, the areas of non-prairie vegetation being ignored, not as unworthy of study, but because the vanishing prairie colonies required all the time available. rt. Colony of Swamp Grasses (Spartina and Elymus), Station I, a This colony of slough grass (Spartina michauxiana) and wild rye (Elymus) is located a short distance north of the “Clover Leaf” switch tracks and just south of the telegraph pole marked “Toledo 318 miles and St. Louis 133 miles.” The length of this colony was about 40 aces. : During August, 1910, it was dry, but probably in the spring and early summer, rains make this area a habitat for swamp grasses. 42 Though it was an almost pure stand of slough grass, with this were mixed a few plants of wild rye (Elymus virginicus submuticus and E. canadensis). These grasses reach a height of about four feet. The ground was very hard and dry, and there were large cracks in it. A single collection of animals was made here, No. 179. Common Names Scientific Names Common Garden Spider Argiope aurantia Ambush Spider Misumena aleatoria Differential Grasshopper, adult and nymphs Melanoplus differentialis Red-legged Grasshopper, adult and nymphs Melanoplus femur-rubrum Texan Katydid Scudderia texensis Meadow Grasshopper Orchelimum vulgare, adult, and nymphs of vulgare or glaberri- mum. Dorsal-striped Grasshopper Xiphidium strictum Black-horned Meadow Cricket Cicanthus nigricornis Four-spotted White Cricket Cicanthus quadripunctatus Ground-beetle Leptotrachelus dorsalis Sciomyzid fly Tetanocera plumosa The basic food-supply in such a habitat is of course the grasses, and this fact fully accounts for the presence of large numbers of individ- uals which feed upon grasses, as do the Orthoptera in general. But the Orthoptera listed are not exclusively vegetable feeders, for Forbes (’0§: 147) has shown that Xiphidium strictum feeds mainly upon in- sects, chiefly plant-lice,as well as upon vegetable tissues, including fun- gi and pollen; Orchelimum vulgare (p. 144), largely upon plant-lice and other insects; and Cécanthus quadripunctatus (p. 220), upon plant tissues, pollen, fungi, and plant-lice. These observations were based upon a study of the contents of the digestive tract. The food of the sciomyzid fly is unknown. The garden spider lives exclusively upon ani- mal food; and being abundant, it must exert considerable influence upon other small animals. It not only destroys animals for its food, but many others are ensnared in its web and thus killed. In one of the webs I found a large differential grasshopper. The rank growth of vegetation furnishes the necessary support for the webs of this spider. Some of the insects, as Melanoplus differentialis and M. femur- rubrum, oviposit in the soil, but others—Scudderia texensis, Xiphid- ium strictum, Orchelimum vulgare, and Cicanthus—deposit their 43 eggs in stems of plants or under the leaf-sheaths of grasses (Forbes, 05: 143, 145, 148, 216). The mode of oviposition in these Orthop- tera raises the question whether or not they are able to pass their com- plete life cycle within this habitat. Are the species which oviposit in the soil able to endure submergence during the wet season of the year, or must they each year re-invade this habitat from the more favorable adjacent regions? The sciomyzid fly is a regular inhabitant of such situations, for an allied species, Tetanocera pictipes Loew, has been found by Needham (’o1: 580) to be aquatic, breeding on colonies of bur reed (Sparganium), and Shelford (’13a: 188, 284) also finds plumosa in wet places. The flower spider, Misumena, captures its prey direct, frequenting flowers where its prey comes to sip nectar, With more perfect drainage the character of this habitat wouid change; a more varied growth of vegetation would probably devel- op; and the relative abundance of the various kinds of animals would also change. The present imperfect drainage is more favorable to the accumulation of vegetable debris than if the habitat was connected with a stream which could float it away. The periodical drying hastens decay, and the deep cracks in the soil become burial places for various kinds of organic debris. 2. Colony of Wild Rye, Elymus virginicus submuticus, Station I, c* Wild rye is a swamp grass. This colony was located about half a mile north of the colony of slough grass (Station I, a) and about 222 feet south of the first east and west cross-road north of Charleston. For a general view of this grassy habitat see Figure 1, Plate II. In length this habitat extends about one third the distance between two consecutive telegraph poles, or about 65 feet. The conditions of the habitat are in general similar to those in the colony of Spartina. The black soil was very dry and much cracked when examined, late in Au- gust. Though a few plants of Asclepias sullivantii grew here among the grass, it was a dense, almost pure stand of wild rye, which reached a height of about three and a half feet. Only a very few collections were made here, and these were for the sole purpose of determining the general composition of the asso- ciation. These collections, Nos. 153, 180, and 181, were as follows: *Animals were not emia’ at Station I, 6, and therefore the location will not be discussed here. 44 Common Garden Spider Argiope aurantia No. 153 Differential Grasshopper Melanoplus differentialis Red-legged Grasshopper Melanoplus femur-rubrum No. 180 Dorsal-striped Grasshopper Xiphidium strictum No. 180 Meadow Grasshopper Orchelimum vulgare, adult, and nymphs of vulgare or glaberrimum No. 180 Texan Katydid Scudderia texensis No. 181 These are all abundant species. O. vulgare, by its persistent fid- dling, is noticeable in all such grass spots during hot sunny weather. A live differential grasshopper was found in the web of the garden spider. A comparison of the two colonies of swamp grasses, Spartina and Elymus, will probably help to give one a general idea of the kind of invertebrates which were abundant in the original swamip-grass area of this vicinity. It will be noticed that grass and grass eaters are the dominant species, and that upon these a smaller number of preda- ceous animals depend. The characteristic species are the Orithoptera and the garden spider. This spider, on account of its predaceous hab- its, is able to live in a great variety of open situations, but does not normally live in dense woodlands. 3. Wet Area of Swamp Milkweed (Asclepias incarnata), Station I, d This colony of swamp milkweed was about one eighth of a mile north of the east and west cross-road. This flat, poorly drained black- soil area, about 80 feet long, was wet throughout August, crawfish holes being abundant (Pl. IITA, fig. 2; Pl. IIIB, figs. 1, 2). To the east, beyond the boundary fence, in the adjoining corn field, stood a pool of water surrounded by a zone of yellowish weakened corn, visited occasionally by a few shore birds. Along the east side of the newly formed railway embankment (PI. II], fig. 1) is a shallow trench containing water and a growth of young willows (Salix) and cottonwoods (Populus deltoides), also blue flags (Iris versicolor), bulrush (Scirpus), and sedge (Carex). The characteristic plants over this area were the abundant swamp milkweed (Asclepias incar- nata, Pl. ILIA, fig. 1) and Bidens. A few plants of water horehound (Lycopus) and dogbane (Apocynum medium) were present, and many individuals of a low plant with a winged stem (Lythrum alatum). The collections (Nos. 1, 12, 13, 14, 15, 18, 32, 37, 45, 156, and 157) of animals taken here were as follows: 45 Pond snail Galba umbilicata 18 Prairie Crawfish Cambarus gracilis _ Garden Spider Argiope aurantia _ Ambush Spider Misumena aleatoria 157 Chigger Trombidium sp. —_— Nine-spot Dragon-fly Labellula pulchella _— Stink-bug Euschistus variolarius 12 Small Milkweed-bug Lygeus kalmii 12 Large Milkweed-bug Oncopeltus fasciatus I Ambush Bug Phymata fasctata 12 Tarnished Plant-bug Lygus pratensis 12 Soldier-beetle Chauliognathus pennsylvanicus 156 Black Flower-beetle_ Euphoria sepulchralis 156 Four-eyed Milkweed-beetle Tetraopes tetraophthalmus 12 Milkweed-beetle Tetraopes femoratus (?) I Leaf-beetle Diabrotica atripennis I Dogbane Beetle Chrysochus auratus 14 Celery Butterfly Papilio polyxenes 15, 45 Philodice Butterfly Eurymus philodice 12 Idalia Butterfly Argynnis idalia 33 Milkweed Butterfly Anosia plexippus — Honeysuckle Sphinx Hemaris diffinis 32 Giant Mosquito Psorophora ciliata 13 Giant Fly Mydas clavatus 12 Honey-bee Apis mellifera —_ Pennsylvania Bumblebee Bombus pennsylvanicus 155 Bumblebee Bombus fraternus 12 Bumblebee Bombus separatus 12, 157 Carpenter-bee Xylocopa virginica 1,156 Rusty Digger-wasp Chlorion ichneumoneum I2 The soft, wet, black soil contained large numbers of crawfish holes, and from several of them T. L. Hankinson dug specimens of Cambarus gracilis. Frogs (Rana) were seen but none were secured. A Caro- lina rail was flushed from the ditch along the track, and on the mar- gins of the water in the adjacent corn field Mr. Hankinson recognized some shore birds. The dragon-fly Libellula pulchella was abundant on the wing and resting on the vegetation, and two examples were found in the webs of Argiope aurantia. No nymphs were found, but doubt- less eggs were laid by some of the numerous adults. It was interest- ing to observe the fresh burrows of the crawfish which had traversed the fresh firm yellow clay of the recently reinforced railway embank- 46 ment (shown im Pl. Ul, fig. 2) and appeared upon its surface. ‘The occurrence hereof 4 swall anal. Calbuccaddivata, is of imterest. A very large species of mosquito with conspicuously banded legs, Psoro- phora ciliata, was found here. Though these aquatics and the ground fonms did not receive much attemtion, they are representative of wet places. The presence of certain plants im this habitat has determined. the occurrence of several species of animals. ‘Thus the dogbame Apocy- mm medium accounts for the brilliamtly colored leaf-beetle Chry- sochus auratus, which feeds wpon its leaves and roots. But the most conspicuous feature of this habitat im August is the wariety of imsects which are attracted by the flowers of the swamp milkweed. These flowers may be regarded as so much imsect pasture. A few butterflies were observed, Papiho polyxenes beimg found im am Argiope web; and om the flowers of the swamp milkweed were Papilio cresphomtes, Ewry- mus plilodice, Al gyums daha, Anosia plesippus, and the honeysuckle sphinx (Hemaris defenis). Among the most abundant Hymenoptera were the homey-bee (Apis mellifera) and the common rusty digger- wasp (Chlorion ichmewmoncum). Others were the carpenter-bee (Xylocopa virgimca) and the bumblebees Bombus fratermus and sep- aratus. On the flowers of the thistle (Cirsium) near this station, Bowm- bus pemmsyloomicus was also talsem lr amet ney ours was taken om the flowers of the swamp milkweed. Beetles from these flowers were the spotted muilkweed-beetles (Tetraopes tetraophthalmus amd femoratus?) the flower-bectle Euphoria sepulchralis, amd, late im Amgust, great mumbers of the soldier-beetle Chauliogmathus pemm- SyWoumicus.. The Hemiptera found are equally chararteristic, and in. weed thugs (Oncopelius fasciatus andl Lygeus kalo) and Lygus pratensis. a om the milkweeds, preying mot wpom the plamt, bact tmpom These were tie amiuch img (Pleymata fascioia) and the cinbush sects of this Ihalbntat: att tihis seasom. Amother abundant: animal was tie chigger, a larval mute of the genus Trombidionm, which is brushed from. the vegetation by one’s anms and legs. “Mhese irritating pests were so abomdamt that to worl with comfort im this regiom it was mecessary to ponsice one's chutes and bal wat Sowezs ot seteien: ‘These young sixlegwed mutes are supposed to prey mpom imsents, as do the adults. Acoording to Chittenden (06:4) diggers are most abum- damt im damp places and forest margims, amd among shrubs, grass, 47 and herbage. “The adults are knowm to eat plamt-lice, small caterpil- lars, amd grasshoppers’ eggs. ‘This mite is thus am importamt preda- cross aneuaher of the association. The dragnnfies are swell kinrars 10 feed wpom small insects, which they capture om the wing, and om ac- coumt of thenr abundance they are imfimential imsects here. Am exammation of the list of ammmals secured at this station shows that there is considerable diversity im the conditions under which their breeding takes place. Indeed the breeding habits amd places are almost as diwerse as are the feedimg relations. ‘Thus the smail Galbo breeds im the water ; amd the crawfish, Cambarus gracilis, lives as a tur- rower except for a brief period im sprimg, whem it is foumd im streams. It 1s distimetly a subterrameam species. “The gardem spider, im the fall, leaves its epps im its web. "The life iistory of the ambush spider is mot kmowm. Itt seems probable that the sexes meet mpom flowers, amd as the flowers fade they migrate to fresh omes—a respomse which Ham- cock has observed ("11 : 182-186) im the allied species Miswmmena voto. "The ambush bug, whem found om flowers, is im a large mmmiber of cases copullatimg, but where the eggs are laid amd the young dewel- oped is mnkmowm to me. “Mhougi this bug also must migrate with the fading of the flowers, after the halbit of Miswmenm, it is wimged amd does mot have to go “om foot” as the spider probably does. Wem dis- thurbed these fbmgs do mot as a mule seek to escape by flight, amd it is mot unlikely that they often crawl from ome flower to amotiher winem tie diistameae is short. "Mhe soldiertbeetle is similar to the ambush bog im its propemsitty to copulate om flowers. ‘The milkweed beetles amd the doghane hectle are commonly seem copullatimg mpon the leawes amd stems of tthe plats om winch they lnwe. The tarva of the milkweed beetles bore imo the roots and stems of plamts; the doghame beetle has similiar halts. Of the butiterfiies, Amoma was observed copulatimg om tthe willows, ome sex with the wimgs spread, tthe fore omnes overlappimg im part the immder pane, tthe otter sex with the wings folded ttogetiner werticailly, the heads of the imsects tbheimg ttmnmed im opposite directions. ‘The egws of the mosquito are laid mear the surface of the watter. “Mhe honeybee andl immiblledbees are social, amd the breedimg amd care of the young are quite differemt fhrom those of the other ammmals fomnd an tits soln Xywlocopa cuts the mest for tts brood im solid wood, - 5 rather foreign mpom the pramie, alithomgh posts amd ties wien is ding: itm tthe grrommdl, with warioms grasshoppers; mpom tihese tire egy iis land amd the young larva feeds. “This wasp probally did mot freed! im this moist thalbitat. “Mhe wet suibsthratumm here is probally mm- faworablle for tthe ireedimg of those Orthopitera winch deposut thhenr epys im tie soll. 48 4. Cone-flower and Rosin-weed Colony, Station I, e This station was continuous with and just north of the swamp milkweed area (Station I,d) just described. The surface of the ground sloped gently upward toward the north, but none of it was free from crawfish holes, and the ground-water level was not far below. The soil is very dark in color. The general appearance of this habitat is shown in Plate V. The large-leaved plants are Silphium terebinthinaceum, and the heads of the numerous cone-flowers (Lepachys pinnata) show as black points in the picture. The cone-flower was the dominant plant at this time. There were a few scattered plants of Silphium integrifolium and of wild lettuce (Lactuca canadensis). At the time the collecting was done in this area Silphium was not in blossom, and all the flower-collecting was from Lepachys. The collections of animals taken here (Nos. 8, 40, and 158) are as follows: Crawfish Cambarus sp. (Burrows observed) Garden Spider Argiope aurantia 40 Sordid Grasshopper Encoptolophus sordidus 158 Differential Grasshopper Melanoplus differentialis 40 Red-legged Grasshopper Melanoplus femur-rubrum 40 Texan Katydid Scudderia texensis 40 Dorsal-striped Grasshopper Xiphidium strictum 40 Black-horned Meadow Cricket Gicanthus nigricornis 40 Membracid bug Campylenchia curvata 40 Jassid Platymetopius frontalis 40 Lygeid Ligyrocoris sylvestris 40 Ambush Bug Phymata fasciata 40 Chrysomelid beetle Nodonota convexa 40 Southern Corn Root-worm Diabrotica 12-punctata 40 Beetle Robber-fly Asilide _— Trypetid fly Euaresta equalis 40 Eucerid bee Melissodes bimaculata 8 Eucerid bee Melissodes obliqua 8 Nomadid bee Epeolus concolor 8 Social wasp Polistes sp. _— Collection No. 40 was made by sweeping the vegetation with an in- sect net. No. 8 is a collection made from the flowers of Lepachys pin- nata. The nest of Polistes was across the railway track. from this station. The abundance of Melissodes obliqua and of the pretty 49 Epeolus concolor on the flowers of Lepachys indicates the attractive power of this plant. The coarser plants furnish support for the webs of Argiope, the flowers serve as drinking cups in which Phymata lies in ambush; and the varied vegetation affords food for the numerous Orthoptera. The proximity of ground-water accounts for the pres- ence of Cambarus, and an adjacent corn field explains the presence of Diabrotica. A robber-fly (Asilide) was seen but not captured. It is interesting to see Melissodes obliqua as it hurries round and round the heads of cone-flowers and sweeps up the great masses of yellow poilen. The hind pair of legs, when loaded with pollen, have nearly the bulk of the abdomen. Robertson (’94; 468) says that this is the most abundant visitor to the cone-flower, and more abundant on this flower than on any other. It is probable that the conditions within this habitat were suitable. for the breeding of most of the species listed. Huaresta equalis has been bred from the seed pods of the cocklebur (Xanthium) and prob- ably came from the adjacent corn field. It is most likely on flowers that the strepsipterid parasitic insects find many of their hosts (Pierce ‘og b: 116). These insects are found on the following prairie insects : Polistes, Odynerus, Chlorion ichneumoneum, C. pennsylvanicum, and C. atratum. Robertson (’10) records many important observations on the hosts of Illinois Strepsiptera. 5. Colony of Blue Stem (Andropogon) and Drop-seed (Sporobolus), bordered by Swamp Milkweed, Station I, g* This colony formed the extreme northern part of the prairie area examined along the “Clover Leaf” track. It extended along the track for a distance of about 200 feet. The area is level black soil prairie. Its general appearance and location are indicated in Figure 2, Plate II, and in Figure 2, Plate III, photographs taken at the time of our study, and in Figure 2, Plate IV, a photograph taken by T. L. Hankin- son April 23, 1911. This latter view clearly shows the character of the drainage during the spring wet season. During the late summer, the dry season, the ditch along the railway track concentrates the drainage so that a colony of swamp milkweed (Asclepias incarnata) and small willows flourish in it. Upon the well-drained part of this area there is a rather rich growth of Andropogon furcatus, A. virginicus, and Sporobolus cryptandrus, and many plants of the dogbane Apocynum medium and a few plants of Asclepias sullivantii. This was the larg- est and best colony of the upland prairie grasses seen along the Clover Leaf tracks; and yet when it is compared with the patches of such *No collections were made at Station I, f. 50 grass east of Charleston (Station III) it is a meager colony. Just south of this grassy colony was a large one of the mountain mint. Pycnanthemum flexuosum. This is shown in Figure 1, Plate IV. The collections of animals (Nos. 1, 2, 3, 4, 6, 7, 19, 28a, 36, 39, 44, 157, and 159) are as follows: Pond snail Crawfish Harvest-man Garden Spider Ambush Spider Red-tailed Dragon-fly Nine-spot Dragon-fly Prairie Ant-lion Lace-wing Fly Grasshopper Sordid Grasshopper Differential Grasshopper Red-legged Grasshopper Texan Katydid Meadow Grasshopper Cone-nosed Katydid Four-spotted White Cricket Stink-bug Small Milkweed-bug Large Milkweed-bug Rapacious Soldier-bug Ambush Bug Four-eyed Milkweed Beetle Rhipiphorid beetle Bill-bug Milkweed Butterfly Giant Mosquito Mycetophilid fly Giant Bee-fly Vertebrated Robber-fly Honey-bee Bumblebee Bumblebee Eucerid bee Nomadid bee Leaf-cutting bee Rusty Digger-wasp Myzinid wasp Physa gyrina 19 Cambarus sp. _ Liobunum politum? 7 Argiope aurantia 6, 39 Misumena aleatoria 6, 157, 159 Sympetrum rubicundulum 7 Libellula pulchella — Brachynemurus abdominalis 36 Chrysopa oculata 44 Syrbula admirabilis 3 Encoptolophus sordidus 44 Melanoplus differentialis 39 Melanoplus femur-rubrum 3, 39 Scudderia texensis 2, 44 Orchelimum vulgare —, 3 Conocephalus sp. 159 CEcanthus 4-punctatus 3 Euschistus variolarius 39 Lygeus kalmii 1,6 Oncopeltus fasciatus I Stnea diadema 5 Phymata fasciata I Tetraopes tetraophthaimus I Rhipiphorus dimidiatus 6 Sphenophorus venatus 39 Danais archippus i Psorophora ciliata 44 Sciara sp. 6 Exoprosopa fasciata 6 Promachus vertebratus 39, 44 Apis mellifica I Bombus fraternus I Bombus separatus I Melissodes bimaculata 6 Epeolus concolor 6 Megachile mendica I Chlorion ichneumoneum I Myzine sexcincta 1, 6 51 Physa and Cambarus were found among the milkweeds on account of the wet ground, and the presence of the giant mosquito was prob- ably due to the same condition. The majority of the other animals were attracted to this habitat by the milkweed, particularly by its flow- ers. Among these were the milkweed bugs and beetles, the milkweed butterfly, the honey-bee, and the rusty digger-wasp. The dense growth of the milkweeds does not appear to be so favorable to the garden: spider as is the more open and irregular growth of vegetation else- where. The ambush spider frequented the milkweed flowers for prey and also the flower masses of the mountain mint, on which it was in active competition with the ambush bug and the rapacious soldier-bug, which have similar food habits. The mountain mint, whose flowers are frequented by the predaceous animals just! mentioned, is also vis- ited by rhipiphorid beetles, the bee-fly (Exoprosopa fasciata), the bees Melissodes bimaculata and Epeolus concolor, and the myzinid wasp Myzine sexcincta. ‘The prairie grasses were frequented by a large variety of Orthoptera, which showed a decided preference for them, their abundance being evident in the list. The wide-ranging predators and parasites, such as Liobunum, Libellula, Sympetrum, Chrysopa, Brachynemurus, Promachus, Chlorion, and Myzine, probably forage over extensive areas compared with the relatively sedentary kinds, such as Misumena, Argiope, Phymata, and Sinea. Phymata was cap- tured on a milkweed flower with a honey-bee; Promachus vertebratus was taken on a grass stem with a stink-bug (Euschistus variolarius) ; and Misumena aleatoria was taken with a large, nearly mature female nymph of Conocephalus. The conditions which permit an animal to breed in a habitat have an important influence upon the character of its population. It is evi- dent that many of the animals taken do not breed here. Some of the relatively sedentary kinds, such as Physa, Cambarus, and Argiope, and probably Misumena, do not cover long distances. Good examples of the wider ranging forms are Sympetrum, Libellula, Danais, Proma- chus, Apis, Bombus, and Chlorion. Several of the animals, as the snails, crawfish, and the dragon-flies, require an aquatic habitat. Chrysopa places its eggs among colonies of plant-lice, and Brachyne- murus probably spends its larval life in dry or sandy places, feeding upon ants and other small insects, as do other ant-lions. Several of the Orthoptera deposit their eggs in the soil; and some of the locustids, among grasses and herbaceous stems. Others are found copulating upon the plants on which the young feed, as Tetraopes, Chrysochus, Lygeus, and Oncopeltus; and still others copulate in the flowers mainly, as Phymata. It is probable that on the flowers some of the para- 52 sitic species find their hosts, as Pierce (’04) has shown to be the case in the rhipiphorid genus Myodites. Rhiptphorus is probably parasitic. 6. Supplementary Collections from Station I In addition to the specimens given in the preceding lists for Statio I there are others, general collections from this area, which should b listed for this prairie. For details concerning each species of the fol lowing consult the annotated list. Garden Spider Ambush Spider Chigger Dorsal-striped Grasshopper Coreid bug Ambush Bug Ladybird Leaf-beetle Four-eyed Milkweed Beetle Old-fashioned Potato Beetle Margined Blister-beetle Black Blister-beetle Snout-beetle Snout-beetle Giant Bee-fly American Syrphid Tachinid fly Bumblebee False Bumblebee Eucerid bee Short Leaf-cutting Bee Halictid bee Halictid bee Stizid wasp Rusty Digger-wasp Harris Digger-wasp Digger-wasp Solitary wasp Argiope aurantia Misumena aleatoria Trombidium sp. Xiphidium strictum Harmostes reflexulus Phymata fasciata Hippodamia parenthesis Trirhabda tomentosa Tetraopes 4-ophthalmus Epicauta vittata Epicauta marginata Epicauta pennsylvanica Centrinus penicellus Centrinus scutellum-album Exoprosopa fasciata Syrphus americanus Trichopoda ruficauda Bombus separatus Psithyrus variabilis Melissodes obliqua Megachile brevis Halictus fasciatus Halictus virescens Stigus brevipennis Chlorion ichneumonenm Chlorion harrisi Ammophila nigricans Odynerus vagus 26 31 35 27 24, 26, 43 Hankinson Hankinson Hankinson Hankinson 26, 152 41 Hankinson 24, 31 II 38 22 22 24, 48 Hankinson 26 23 35, Hankinson II. Pratrtz Area NEAR Loxa, ILLINOIS, Station IL This station includes patches of prairie along the Cleveland, Cin- cinnati, Chicago and St. Louis (Big Four) railroad right-of-way be- tween Charleston and Mattoon, Ill, and about one mile west of 53 the small station of Loxa. Along this track the telegraph-pole num- bers were used in locating our substations. This is a rather level black soil area, originally poorly drained and wet, but now considerably modified by the ditching and grading occasioned by railway construc- tion and maintenance. The changes have been similar to those on the prairie north of Charleston, but the ditching has been a few feet deeper and the embankment is higher. The most abundant and characteristic kinds of vegetation are the tall prairie grasses—blue stem (Andropo- gon furcatus), drop- seed (S‘porobolus cryptandrus), and beard grass (Andropogon virginicus)—a rosin-weed (Silphium laciniatum), the flowering spurge (Euphorbia corollata), wild lettuce (Lactuca can- adensis), rattlesnake-master (Eryngium yuccifolium), and beggar-- - ticks (Desmodium). Many other kinds of plants were also present. The general appearance of this habitat is shown in plates VI and VII. Our collections from this prairie (Nos. 47-57 and 176-178) are as follows: Garden Spider Argiope aurantia 49, 179 Ambush Spider Misumena aleatoria 47, 178 Sordid Grasshopper Encoptolophus sordidus 48 Two-lined Grasshopper Melanoplus bivittatus 55 Differential Grasshopper Melanoplus differentialis 48 Meadow Grasshopper Orchelimum vulgare 178 Lance-tailed Grasshopper Xiphidium attenuatum 48 Dorsal-striped Grasshopper Xiphidium strictum 48, 50, 57 Stink-bug Euschistus variolarius 50, 52, 178 Ambush Bug Phymata fasciata 48, 52, 54, 55, 57, 178 Dusky Leaf-bug Adelphocoris rapidus 55 Soldier-beetle Chauliognathus pennsylvanicus 178 Southern Corn Root-worm = Diabrotica 12-punctata 55 Margined Blister-beetle Epicauta marginata 48 Black Blister-beetle Epicauta pennsylvanica 48, 178 Rhipiphorid beetle Rhipiphorus dimidiatus 52 Rhipiphorid beetle Rhipiphorus limbatus 178 Snout-beetle Rhynchites eneus 48 Thoe Butterfly Chrysophanes thoe 55 Dogbane Caterpillar Ammalo eglenensis or tenera 53 Giant Bee-fly Exoprosopa fasciata 47, 57, 176 Robber-fly Deromyia sp. 51 Vertebrated Robber-fly Promachus vertebratus 56 Corn Syrphid Mesogramma politum 177 Syrphid fly Allograpta obliqua 177 54 Tachinid fly Cistogaster immaculata Pennsylvania Bumblebee Bombus a 50, 52, 55, 176 False Bumblebee Psithyrus variabilis Eucerid bee Melissodes bimaculata “43 Nomadid bee Epeolus concolor 48, 52 Halictid bee Holictus obscurus Halictid bee Halictus fasciatus 48, 52 Black Digger-wasp Chlorion atratum 55 Pennsylvania Digger-wasp Chlorion pennsylvamcum 55 Myzinid wasp Myszine sexcincia 52, 55 Ant Formica pallide-fulva schaufussi incerta 52 The general conditions of this prairie appear to have been less dis-~ turbed than at Station I; at least the prairie vegetation is more exten- sive and uniform. The change in the vegetation is apparently greater than the change in the kinds of animals. Their feeding and breeding relations appear to be much like those at the prairie stations previously discussed. In the flowers of the cup-leaved rosin-weed (Silphium integri- folium) was found a giant bee-fly (Exoprosopa fasciata) which had. been captured by the ambush spider (Misumene aleatoria), and on webs in colonies of this same plant the garden spider (Argiope auran- tia) was observed, with a grasshopper (Melanoplus differentialis) en- tangled in the web. From the flowers of this Silphium the following insects were taken: Epicauta marginata and E. pennsylvanica, Rhyn- chites eneus, Phymata fasciata, Encoptolophus sordidus, Melanoplus differentialis (nymph), Xiphidium strictum (adult and nymph), X. attenuatum, Melissodes bimaculata and obliqua, Epeolus concolor, and Halictus fasciatus. ‘The margined blister-beetle (Epicauta marginata) was found both upon the flowers and the leaves of the plant. On the flowers of the purple prairie clover (Petalostemum purpureum), Bom bus pennsylvanicus, Xiphidium strictum, and Euschistus variolarius were taken. Collection 176 was taken from the flowers of Liatris scariosa, and Nos. 55 and 178 from the flowers of Eryngium yucci- folium. Swarms of the small corn syrphid, Mesogramma politum, were present, on one day settling by dozens on my hands and clothes, where they were easily grasped by the wing. It had been a warm day, and this swarming was in the sunshine at about 4:30 p.m. The flies came from a large corn field a few feet away. 35 THE. Prams Agra East or Cnartesron, Sration IT This prairie area is about two miles east of Charleston along the “Big Four” railway track. ‘There were two colonies here. One, sub- station a, was on low black-soil prairie just west of the first morth and south road crossing the railway track east of Charleston. This was largely a colony of the large-leaved rosin-weed, Silthium terebinthi- naceum. ‘The second colony, substation b, was a mile and a half di- rectly east of substation a, and half a mile east of the second north and. south road east of Charleston. Substation or “station” a was originally far out upon the black soil prairie; b, on the other hand, is of special interest because it was origi- nally wooded, has been cleared and maintained as a railroad right-of- way, and contains today, therefore, a practically unique mixture of for- ca and geditis peaitscad gaiicle. wat the pattie Viads denseciteag: The soil, lighter in color than the black soil prairie, is representative of the wooded regions. This colony has every appearance of a cleared forest area invaded by prairie organisms. The animals at station a were not studied, and ihe only record is that of the black blister-beetle, Epicauta pennsylvanica (No. 119), which was abundant on the flowers of Silphinm terzbinthinaceum. At station 6 excavation was necessary to lower the road-bed, and upon the disturbed soil thus thrown up along the track the prairie veg- etation had become established. The general appearance of this region is shown in plates VIII and IX. Here grew large quantities of rosin- weed (Silphium terebinthinaceum) and blue stem (Andropogon) ; in places upon high ground, indeed, this prairie grass was Associated with it was the flowering spurge, Euphorbia corollata, as seen' in Plate VIII The forest near by is shown in the background. This same forest and grass area is shown in the background and mid- dle of Plate IX, and in the foreground of the same picture is shown the mixture of prairie and forest plants. Here are hickory sprouts, crab-apple, grape, sumac, and smilax, intermingled with Silphium, blue stem, and Lactuca canadensis. Not all of these appear in the photograph, but they were present in some parts of the colony. The collections here (Nos. 58-62 and 175) are as follows: Leather-colored Grasshopper § Schistocera alutacea 59 Black-horned Meadow Cricket Gcanthus nigricornis 62 Meadow Grasshopper Orchelimum vulgare 175 Soldier-beetle Chauliognathus pennsylvamcus 175 Spotted Grape-beetle Pelidnota punctata 58 Black Blister-beetle Epicauta pennsylvamica (Sta. TI,a) 119 56 Cabbage Butterfly Pontia rape 61 Vertebrated Robber-fly Promachus vertebratus 62 Pennsylvania Bumblebee Bombus pennsylvanicus 175 Impatient Bumblebee Bombus impaitiens 175 Bumblebee Bombus auricomus 175 (Rose-gall) Rhodites nebulosus 60 No animals were taken here which were dependent upon the sumac, hickory, crab-apple, or smilax. Pelidnota lives upon the grape, and grapes are primarily woodland or forest-margin rather than prairie plants. Schistocerca is also probably a marginal species. On the flow- ers of Silphium terebinthinaceum were taken Orchelimum vulgare, Chauliognathus pennsylvanicus, and Bombus pennsylvanicus, auri- comus, and impatiens. The persistence of woodland vegetation in this locality, in spite of the repeated mowings and burnings, shows that it has much vigor, and would, if undisturbed, in a few years shade out the prairie vege- tation and restore the dominance of the forest. With such a change in the vegetation there would of course be a corresponding change in the animals. DESCRIPTION OF THE FOREST HABITATS AND ANIMALS 1. The Bates Woods, Station IV The Bates woodland area is located about three and a half miles northeast of Charleston on the farm that was owned by Mr. J. I. Bates, and consists of about 160 acres. It includes a bottom-land area near the Embarras River, and extends up the valley slope on to the upland. It is isolated from the trees bordering the river (Pl. X, fig. 1) by a narrow clearing, and from those on the northeast, north, and north- west by another clearing (Pl. XI); on the south and southwest it is continuous with partially cleared areas, which extend south to the Big Four railway track. The river bottom-land is undulating and rises rather gradually toward the base of the bluffs. The bluff line is irregular on account of the ravines which have been etched in it, the largest of which forms the southern boundary of the region examined. The upland is rela- tively level. The soils on the bottom are darker colored, except in places near the base of the bluff, and at the mouths of the ravines where the upland soil has been washed down. The upland soil is pre- sumably the “light gray silt loam” of the State Soil Survey (Moultrie County Soils, Ill. Exper. Sta. Soil Rep., 1911, No. 2, p. 23). All of 57 the area examined was well drained, and all was forested. The region is not homogeneous physically or in its vegetation, and for this reason the area is divided into substations in order that the influences of the local conditions within the forest might be preserved, and their indi- viduality recognized. 2. The Upland Oak-Hickory Forest, Station IV,a The general appearance of this forest is shown in plates XII and XIII. This is an open second-growth forest composed of oaks and hickories—such as white oak (Quercus alba), black oak (Q. velutina), shag-bark hickory (Carya ovata), bitternut (C. cordiformis), pignut (C. glabra), and scattered individual trees of red oak (Q. rubra), wal- nut (Juglans nigra), and mulberry (Morus rubra). The shrubs are sassafras (Sassafras variifolium), sumac (Rhus glabra), Virginia creeper (Psedera quinquefolia), poison ivy (Rhus toxicodendron), rose (Rosa), raspberry (Rubus), moonseed (Menispermum cana- dense), and tree seedlings. The average diameter of the largest trees is 8-10 inches. Most of the small growth consists of the sprouts from stumps, and many of these are 2-3 inches in diameter. The forest crown is not complete, and as a consequence there are more or less open patches in which most of the herbaceous growth is found, such as horse mint (Monarda bradburiana), pennyroyal (Hedeoma pule- gioides), everlasting (Antennaria plantaginifolia), tick-trefoil (Des- modium nudiflorum), and other, less abundant kinds. Even a plant quite characteristic of the prairie, the dogbane Apocynum, was found here in one of the open glades. The forest floor has an unequal covering of dead leaves, largely oak, most of which lie in the low vegetation and in slight depressions. Occasionally there is but little cover and the light-colored soil is ex- posed. There are few stumps and logs in this part of the forest, and no thick layer of vegetable mold, so that one would not expect to find any animals which normally frequent moist soil and vegetable debris. As this is a second-growth forest it lacks the conditions which abound in an original growth, where are old, dead and decaying trees, and numerous decaying logs and stumps. In this respect the woods is not fully representative of an original upland forest on well-drained bluff land. The relative evaporating power of the air of this substation was 54 per cent. of that of the standard instrument in the open garden at the Normal School, a fact which indicates a relative evaporation com- parable to that of the ordinary black-soil prairie ; in producing this con- dition, the glade-like, open character of this forest is undoubtedly an important factor. 58 The characteristics of this habitat may be summed up as follows: upland, open, relatively dry second-growth oak-hickory forest, with little undergrowth of shrubs and herbs, and with a small amount of litter and humus; soil dry and firm; and few decaying stumps and tree trunks. The collections of animals made here (Nos. 64-67, 69, 71, 74-83, 88, 91-93, 102, 103, 107, 109, 118, 120-123, 127, 135, 136, 142, 145, 147, 150, 151, 162, 163, 166, 169, 170, 171, and 183) are as follows: Land snail Predaceous snail Land snail Carolina slug Land snail Harvest-spider Harvest-spider Stout Harvest-spider Island Spider White-triangle Spider Rugose Spider Ground Spider White Ant Ant-lion Dog-day Harvest-fly Periodical Cicada Forest Walking-stick Grouse Locust Short-winged Grouse Locust Green Short-winged Grasshopper Sprinkled Grasshopper Boll’s Grasshopper Lesser Grasshopper Acridiid grasshopper Acridiid grasshopper Forked Katydid Angle-winged Katydid Common Katydid Meadow Grasshopper Meadow Grasshopper Striped Cricket Spotted Cricket Woodland Cricket Polygyra albolabris QI Circinaria concava 71 Zonitoides arborea 71 Philomycus carolinensis 71 Pyramidula perspectiva 71, 88 Liobunum vittatum 82, 123 Liobunum ventricosum 123b Liobunum grande 82 Epeira insularis 70 Epeira verrucosa 70 Acrosoma rugosa 70, 147 Lycosa sp. 142, 150 Termes flavipes 72, 76, 79 Myrmeleonide (Forest border) 183 Cicada linnei 162 Tibicen septendecim Diapheromera femorata 64, 93 Tettigidea lateralis 109 Tettigidea parvipennis 122 Dichromorpha viridis 67, 92, 93, 121, 123 Chloealtis conspersa 67, 93, 122 Spharagemon bolli 67, 150 Melanoplus atlanis Melanoplus amplectens 67 Melanoplus obovatipennis 93 Scudderia furcata 109 Microcentrum laurifolium 135 Cyrtophyllus perspicillatus 145 Orchelimum cuticulare 67, 93 Xiphidium nemorale 93, 103 Nemobius fasciatus 67, 93, 122 Nemobius maculatus 122 Apithus agitator 93 Woodland Tiger-beetle Caterpillar-hunter Carabid beetle Ladybird Splendid Dung-beetle Dogbane Beetle Tenebrionid larva Philenor Butterfly Turnus Butterfly Troilus Butterfly Sphingid larva Arctiid moth Notodontid moth Notodontid moth Notodontid moth Geometrid moth Gelechiid moth (Cecidomyiid gall) (Cecidomyiid gall) (Cecidomyiid gall) Syrphid fly Corn Syrphid Vespa-like Syrphid Pigeon Tremex (Oak Bullet-gally (White Oak Club-gall) (Oak Wool-gall) Formicid ant Formicid ant Formicid ant Mutillid ant Short Caterpillar-wasp 59 Cicindela unipunctata 136 Calosoma scrutator 64 Galerita janus 171 Coccineliide 81 Geotrupes splendidus 120 Chrysochus auratus 103 Meracantha contractea 83 Papilio philenor 69, 166 Papilio turnus — Papilio troilus 163 Cressonia juglandis 102 Halisidota tessellaris 168 Datana angusi 65, 162 Nadata gibbosa 169 Heterocampa guttivitta? 127 Eustroma diversilineata 163 Ypsolophus ligulellus? 76, 78, Hankinson Cecidomyia holotricha 107, 170 Cecidomyia tubicola’ 107 Cecidomyia caryecola 107, 170 Chrysotoxum ventricosum 163 Mesogramma politum 76, 78, Hankinson Milesia ornata 103 Tremex columba 66 Holcaspis globulus 170 Andricus clavula 170 Andricus lana 170 Cremastogaster lineolata 118 Aphenogaster fulva 74-80 Formica fusca subsericea 163 Spherophthalma I5I Ammophila abbreviata 127 3. Embarras Valley and Ravine Slopes, forested by ihe Oak-Hickory Association, Station IV,b This station included the slope of the valley from the river bottom (Station IV, c) to the upland forest (Station IV, a) and the side of the south ravine, the bottom of which forms Station IV, d. This sub- station is not as homogeneous physically as the upland or lowland for- est, because the part along the south ravine is relatively open, is well drained, and has a south exposure, and the southeast slope to the low- 60 land forest on the other hand, is well wooded and shaded, and much more humid. The substation also has a considerable amount of litter, leaves, and humus. This region may be considered as transitional be- tween the upland and lowland forest, but it represents, not one but two transitional stages, the south slope approaching the upland forest type, and the southeast slope approaching that of the lowland forest. Thus, if one walked from the upland forest down the slope of the south ravine, and eastward to the southeast valley slope to the bottom- land forest, he would traverse all the main degrees of conditions found at Station IV. The forest cover consists primarily of the following trees: white oak (Quercus alba), black oak (Q. velutina), walnut (Juglans nigra), pignut (Carya glabra), and, in smaller numbers, mulberry (Morus rubra), red oak (Quercus rubra), shag-bark hickory (Carya ovata), bitternut (C. cordiformis) ; and of the following shrubs: redbud (Cer- dis canadensis), sassafras (Sassafras variifolium), moonseed (Menis- permum canadense), five-leaved ivy (Psedera quinquefolia), grape (Vitis cinerea), prickly ash (Zanthoxylum americanum), and sumac (Rhus glabra), the latter growing in large colonies on the open south ravine-slope. On the more moist and shaded southeast slope lived the clearweed (Pilea pumila), a plant quite characteristic of moist deep- shaded woods. Thus sumac and clearweed may be Considered as in- dex plants to the physical conditions in different parts of these two slopes, one shaded and the other rather open. The atmometer, located on the upper part of the south ravine slope, gave a relative humidity of 31 per cent. of the standard in the garden of the Normal School. It will be recalled that in the upland forest (Station IV,a) the atmometer gave 54 per cent., the comparison showing how much less the evaporating power of the air is on the south ravine slope than it is in the upland forest. The relative evap- oration was not determined for the southeast slopes, but the presence of Pilea clearly indicates that it is less than on the south ravine slope, where the instrument was located. On the lower parts of the valley slope, where this substation grades into the lowland, the layers of dead matted leaves and humus reached to a considerable depth, and looked as if they had been pressed down by drifting snows. Such places were found to contain very few animals. This habitat is characterized by a sloping surface, by relative open- ness on the ravine side and dense shade on the valley slope, by rela- tively humid air, by second-growth forest somewhat transitional be- tween that of the uplands (Station IV, a) and the river bottoms (Sta- tion IV, c), by a relatively large amount of shrubbery, by considerable 61 humus and litter, by moist soil, and by more logs and stumps than are in the upland forest. The collections of animals made at this substation (Nos. 68, 84, 85, 87, 89, 90, 94, 100, 104, 105, 106, 108, 110, III, 124, 125, 131, 132, 133, 140, 149, 161, 164, 165, 166, and 168) are as follows: Land snail Land snail Land snail Land snail Carolina Slug Land snail Milliped Milliped Stout Harvest-spider White Ant Woodland Cockroach Green Short-winged Grasshopper Boll’s Grasshopper Scudder’s Grasshopper Woodland Cricket Caterpillar-hunter Wireworm Horned Passalus Tenebrionid larva Troilus Butterfly Philenor Butterfly Lyczenid butterfly American Silkworm Hickory Horned-devil Arctiid caterpillar Rotten-log Caterpillar Notodontid Notodontid larva Geometrid Slug Caterpillar Pigeon Tremex (Acorn Plum-gall) Old-fashioned Ant Tennessee Ant Formicid ant Polygyra clausa Vitrea indentata Vitrea rhoadsi Zonitoides arborea Philomycus carolinensis Pyranidula perspectiva Cleidogona cesioannulata Polydesmus sp. Liobunum grande Termes flavipes Ischnoptera sp. Dichromorpha viridis Spharagemon boll Melanoplus scudderi Apithes agitator Calosoma scrutator Melanotus sp. Passalus cornutus Meracantha contracta Papilio troilus Papilio philenor Everes comyntas Telea polyphemus Citheroma regalis Halisidota tessellaris Scolecocampa liburna Datana angusit Nadata gibbosa Caberodes confusaria Cochlidion or Lithacodes Tremex columba Amphibolips prunus Stigmatomma pallipes Aphenogaster tennesseensis Myrmica rubra scabrinodis schnecki 140, 89, 84, 100, 68, 163, 164 164 164 84 125 164 140 125 III 125 140 110 133 124 124 149 125 85 140 161 166 I61 163 108 168 125 104, 94 161 165 132 131 140 87 140 62 Carpenter-ant Camponotus herculeanus penn- sylvanicus 84, 85 Rusty Carpenter-ant Camponotus herculeanus penn- sylvanicus ferrugineus 90 Short Caterpillar-wasp Ammophila abbreviata 124 4. Lowland or “Second Bottom,’ Red Oak-Elm-Sugar Maple Wood- land Association, Station IV, ¢ This station includes the part of the forest located upon the upper or higher part of the river bottom. This area is sometimes called the “second bottom” because it is above the present flood-plain. The gen- eral position of the forest is shown in Figure 1, Plate X. The fringe of willows along the river bank is shown at a, the flood-plain area is cleared at b; the substation forest is at c; and part of the forest of the valley slope is seen at d. Other views of this station are shown in plates XIV, XV, and XVI (figures 1 and 2). The general slope is toward the river; minor inequalities are due to the action of the tem- porary streams which are etching into the uplands and depositing their burdens of debris at the mouths of the ravines. Soil, leaves, and other organic debris are washed from the upland, the ravines, and the val- ley slopes, and are deposited upon the bottoms, forming low alluvial fans, which have been built up in successive layers or sorted again and again as the temporary streams have wandered over the surface of the fan on account of the overloading and. deposition which filled up their channels. In this manner the soil in general is not only supplied with moisture, drained from the upland, but the various soils are both mixed as successive layers of organic debris are buried by storms and also mulched by the large amount of this debris which is washed and blown to the lowland. No springs were found upon the southeast valley slope, but in the south ravine pools of water were present dur- ing August, 1910, when my observations were made. The forest, characterized by hard maple (Acer saccharum), red oak (Quercus rubra), and elm (Ulmus americana), forms a dense canopy which shuts out the light and winds, thus conserving the mois- ture which falls and drains into it, and making conditions very favor- able to a rich mesophytic hardwood forest. That the relative humid- ity is high is shown by the moisture found in the humus of the forest floor, and, further, not only by the presence of clearweed (Pilea pu- mila) and the nettle Laportea canadensis, which characterize such moist shady woods, but also by the presence of the scorpion-flies (Bit- tacus). ‘These organisms are permanent residents where such condi- 63 ditions prevail, and their presence is as clearly indicative of certain physical conditions as that of aquatic animals. would be indicative of other physical conditions. In addition to these evidences we have the readings of our atmometer, which showed the evaporating power of the air to be 26 per cent. of the standard in the garden at the Normal School. This shows that the relative evaporation is very low, .and that conditions for the preservation of the moisture which falls and drains into this area are very favorable. The general character of this forest is shown in plates XIV, XV, and XVI, Figure 1. The vegetational cover on the lowland is quite different in its com- position from that on the upland. This is shown mainly by the pres- ence of the elm (Ulmus americana), hard maple (Acer saccharum), and red oak (Quercus rubra), and secondarily, by the presence, in smaller numbers, of the black cherry (Prunus serotina), slippery elm (Ulmus fulva), shingle oak{ (Quercus imbricaria), and the Kentucky coffee-tree (Gymnocladus dioica). Other trees present are walnut (Juglans nigra), mulberry (Morus rubra), and bitternut (Carya cor- diformis). The shrubs and.vines are gooseberry (Ribes cynosbati), prickly ash (Zanthoxylum americanum), redbud (Cercis canadensis), buck-brush (Symphoricarpos orbiculatus), green brier (Smilax), five-leaved ivy (Psedera quinquefolia), moonseed (Mentspermum canadense), bittersweet( Celastrus scandens), and grape (Vitis cine- rea). "The characteristic herbaceous vegetation is nettle (Laportea canadensis), clearweed (Pilea pumila), bellflower (Campanula ameri- cana), Indian tobacco (Lobelia inflata), tick trefoil (Desmodium grandiflorum), Actinomeris alternifolia, maiden hair fern (Adiantum pedatum), beech fern (Phegopteris hexagonoptera), the rattlesnake fern (Botrychium virginianum), and Galium circesans and G. tri- folium. Although the forest is generally dense and therefore deeply shaded, there are some places which are comparatively open. Attention, how- ever was devoted mainly to the denser parts. At one place, near the base of the eastern slope of the valley, a few trees had been cut within a few years, and in this glade the conditions and plants and animals were different from those in the dense forest. (See Pl. XVI, figs. 1 and 2. ids habitat may be characterized as follows: lowland densely cov- ered by sugar maple-red oak forest (climax mesophytic) ; very humid air; a moist soil; relatively few shrubs; herbaceous plants—nettles and clearweed—characteristic of damp, shady, rich woods; and considera- ble litter and humus in places. 64 The collections of animals made here (Nos. 113, 114, 116, II7, 137-139, 141, 143, 144, 173, 182, and 184) are as follows, the itali- cised numbers designating collections from the glade: Predaceous Snail Land snail Slug eggs Alternate Snail Milliped Ambush Spider Tent Epeirid Three-lined Epeirid Spined Spider Rugose Spider Ground Spider Cherry-leaf Gall-mite Clear-winged Scorpion-fly Leaf-hopper Pentatomid Coreid Spined Stilt-bug Short-winged Grasshopper Acridiid grasshopper Acridiid grasshopper Scudder’s Grasshopper Round-winged Katydid Nebraska Cone-nose Meadow Grasshopper Meadow Grasshopper Meadow Grasshopper Striped Cricket Elaterid larva Elaterid Black-tipped Calopteron Reticulate Calopteron Horned Fungus-beetle Common Skipper Imperial Moth (larva) Noctuid moth Asilid fly Vespa-like syrphid Long-sting Black Longtail Cocoanut Ant Circinaria concava Vitrea indentata Philomycus (?) eggs Pyramidula alternata Callipus lactarius Misumena aleatoria Epeira domiciliorum Epeira trivittata Acrosoma spinea Acrosoma rugosa Lycosa scutulata Acarus serotine Bittacus stigmaterus Aulacizes irrorata Hymenarcys nervosa Acanthocerus galeator Jalysus spinosus Dichromorpha viridis Melanoplus amplectens Melanoplus gracilis Melanoplus scudderi Amblycorypha rotundifolia Conocephalus nebrascensis Orchelimum cuticulare Orchelimum glaberrimum Xiphidium nemorale Nemobius fasciatus Corymbites sp. Asaphes memnonius Calopteron terminale Calopteron reticulatum Boletotherus bifurcus Epargyreus tityrus Basilona imperialis Autographa precationis Deromyia discolor Milesia ornata Thalessa lunator Pelecinus polyturator Tapinoma sessile II7, rr7; 117, II7, Lig, iy, 143, II7, 113 113 114 173 113 184 173 138 172 172 I44 116 141 143 113 182 II7 143 143 143 I17 143 117 143 143 I43 143 113 113 173 143 173 173 106 143 117 184 I43 43 139 65 5. Supplementary Collections from the Bates Woods, Station IV Tent Epeirid White-triangle Spider Spined Spider Rugose Spider Mealy Flata Leaf-hopper Pentatomid bug Pentatomid bug — Tarnished Plant-bug Coreid bug Coreid bug Rapacious Soldier-bug Acridiid grasshopper Pennsylvania Firefly Margined Soldier-beetle Soldier-beetle Chrysomelid beetle Clubbed Tortoise-beetle Portlandia Butterfly Eurytus Butterfly Gelechiid moth. (Hairy Midge-gall) Corn Syrphid Fly ( Horned-knot Oak-gall) (Oak Wool-gall) Ichneumon Wasp Formicid ant Rusty Carpenter-ant Spider Wasp Epeira domiciliorum Epeira verrucosa Acrosoma spinea Acrosoma rugosa’ Ormenis pruinosa Gypona pectoralis Euschistus fissilis Mormidea lugens Lygus pratensis Alydus quinquespinosus Acanthoceros galeator Sinea diadema Melanoplus obovatipennis Photuris pennsylvanica Chauliognathus marginatus Telephorus sp. Cryptocephalus mutabilis Coptocycla clavata Enodia portlandia Cissia eurytus Ypsolophus ligulellus Cecidomyia holotricha 167 126 148 126 Hankinson Hankinson 124 Hankinson Hankinson Hankinson Hankinson Hankinson 124 Hankinson Hankinson Hankinson Hankinson Hankinson 63 Hankinson Hankinson (Near collection No. 96) Mesogramma politum Andricus cornigerus Andricus lana Trogus obsidianator Aphenogaster fulva Hankinson (Near 96) (Near 96) Hankinson 125 Camponotus herculeanus penn- sylvanicus ferrugineus Psammochares ethiops 97 Hankinson 6. Small Temporary Stream in the South Ravine, Station IV, d This small temporary stream in a ravine formed the southern boundary of the area examined (PI. XVII, figs. 1 and 2). At the sea- son of our examination it was a series of small disconnected pools. Very little attention was devoted to the collection and study of its life. Most of the collections were secured by T. L. Hankinson. A few aquat- ic animals were collected here. In a small pool were taken numerous specimens of the creek chub (Semotilus atromaculaius), and one stone- 66 roller (Campostoma anomalum). Frogs, toads, and salamanders were also taken in the vicinity by Mr. Hankinson, who dug from their bur- rows specimens of Cambarus diogenes, and also secured immunis and propinquus. On the surface of the pools were numerous specimens of a water-strider, Gerris remigis. The forest cover is undoubtedly an important factor in the preservation of such pools, as it controls the evaporating power of the air. Mr. Hankinson tells me that during the summer of 1912 this tem- porary stream was completely dry, and that no fish have been taken from it since the earlier collection mentioned above. From the mouth of the ravine across the bottom to the river it is only a few hundred feet, and in time of heavy or prolonged rains these pools are in direct communication with the river. Such a stream is an excellent example of an early stage in the development of the stream habitat, and shows its precarious character, and the liability to frequent extermination of these pioneer aquatic animals which invade it in its early stages. This -applies particularly to those animals which have no method of tiding over dry periods. On the other hand, those animals which live in the pools, those parts of temporary streams which persist longest between showers, have better chances of survival, particularly bur- rowing animals, like the crawfish and its associates. It seems prob- able that crawfish burrows harbor a varied population; not only the crawfish leeches (Branchiobdellide) but also the eggs of certain Cor- ixide (Forbes, ’76: 4-5; 778, p. 820; Abbott, 12) may almost cover the body of some crawfishes. By means of this burrow ground-water is reached, and a subterranean pool is formed. For ihe elaboration of the stream series see Adams (’01) and Shelford (’11 and 13a). This temporary stream shows how, by the process of erosion, the upland forest area is changed into ravine slopes, and, later, even into the bed of a temporary stream. ‘Thus progresses the endless transfor- mation of the habitat. GENERAL CHARACTERISTICS OF THE GROSS “ENVIRONMENT 1. Topography and Soils of the State Illinois lies at the bottom of a large basin. This is indicated in part by the fact that so many large rivers flow toward it. The mean elevation of the state is about 600 feet, and about a third of it lies be- tween 600 and 700 feet above sea-level. Except Kentucky, the bor- dering states are from 200 to 500 feet higher. Iowa and Wisconsin are considerably higher, so that winds from the north and northwest 67 reach the state coming down grade. Taken as a whole the land sur- face is a tilted plain sloping from the extreme northern part—where a few elevations exceed a thousand feet—toward the south, bowed in the central part by a broad crescentic undulation caused by a glacial moraine, and then declining gradually to the lowland north of the Ozark Ridge, near the extreme southern part of the state. This east and west ridge occasionally exceeds 1,000 feet, but its average height is between 700 and 800 feet. It is very narrow, only about 10 miles in average width, and rises about 300 feet above the surrounding low- land (Leverett, ’96, ’99). South of this ridge lie the bottoms of the Ohio River. The largest river within the state is the Illinois. The soils of the state are largely of glacial origin. Even the un- glaciated extreme northwestern part and the Ozark Ridge region have a surface layer of wind-blown loess. In some places considerable sand was assorted by glacial water, forming extensive tracts of sandy soil, and locally dune areas are active. Along the larger streams there are extensive strips of swamp and bottom-land soils. The remaining soils, which characterize most of the state, were either produced mainly by the Iowan or IJlinoian ice-sheets, as in the case of the relatively poorer soils, or by the Wisconsin sheet, which formed the foundation for the better soil. The dark-colored prairie soils are due to organic debris. Coffey (’12: 42) has said: “Whether this accumulation of humus is due to lime alone or to the lack of leaching, of which its presence is an indication, has not been definitely determined. Neither do we know whether it is due to chemical or bacteriological action; most probably the latter, an alkaline medium being necessary for the growth of those bacteria or other microorganism which cause this form of decomposi- tion.””* 2. Climatic Conditions The climatic features of a region are generally conceded to have a fundamental influence upon its life. The controlling influences upon climate are elevation above sea-level, latitude, relation to large bodies of water—generally the sea—and the prevailing winds. The eleva- tion and relief of Illinois have but a slight influence. In latitude Illinois is practically bisected by the parallel 3934° in the north tem- perate zone. This position influences the seasons and the amount of heat received from the sun. The sea is far distant, but the Great ‘Lakes are near by, and proximity to the interior of a large continent *Consult Hopkins and Pettit (’08) and the County Soil Reports of the State Soil Survey for a'detailed account of the chemical conditions of Illinois soils. The bacterial, algal, and animal population have hardly been noticed by stu- dents of Illinois soils. : 68 brings the state within that influence. And, finally, it lies in the zone of the prevailing westerly winds, and directly across the path of one of the main storm tracks, along which travel in rapid alternation the highs and lows which cause rapid changes of temperature, wind, and precipitation, and thus produce the extremely variable weather condi- tions. The state is 385 miles long, and as latitude has much influence upon climate, the climate of Illinois differs considerably in the extreme north and south. This is clearly shown in the average annual tempera- ture, which in the northern part is 48.9° F., in the central part is 52.70°, and in the southern part is 55.9° (Mosier, ’03). These aver- ages probably. closely approximate the soil temperatures for these re- ‘gions. The average date of the last killing frost in the northern part is April 29; in the central part, April 22; and in the southern part, April 12. The average date of the first killing frost for the northern part is October 9, central part, October 11, and the southern part is October 18 (Henry). The growing season for vegetation in the northern half of the state averages from 150 to 175 days and for the southern half from 175 to 200 days (Whitson and Baker, 12: 28). The precipitation shows similar differences, increasing from north to south. The annual average for the northern part is 33.48 inches, in- creasing to 38.01 in the central and to 42.10 inches in the southern part (Mosier, 03:62). Mosier has shown that the Ozark Ridge, with an average elevation of about 800 feet, condenses the moisture on its south slope so that it has a precipitation of 7.15 inches more than do the counties just north of the ridge. This same humid area appears to extend up the Wabash Valley to Crawford county, and gives the valley counties a rainfall 3 inches in excess of the adjacent counties to the west. The average annual rainfall for the state is 37.39 inches—nearly one third of it during April, May, and June, and if July is included, more than half. The heaviest precipitation, 8.23 inches, is in May and June. As previously mentioned, the state lies in the zone of prevailing westerly winds and across the path of storms. These have a dominant influence upon the direction of the winds. In the northern part of the state, they are, by a slight advantage, southerly—a tendency which progressively increases toward the south, for in the central part the southerly winds reach 55 per cent., and in the southern part 62 per cent. During the winter the northwest winds predominate throughout the state, to a marked degree in the central part, where they reach 60 per cent., and where also the velocity is greatest, reaching an av- erage of 10.3 miles an hour. The velocity of the wind for the entire 69 state is highest during spring. During the summer, the southwest winds predominate in the northern and central parts, and in the south- ern part 82 per cent. of the winds are southerly. The velocity of the wind is least during the summer, and the greatest stagnation occurs in August. During autumn there is a falling off of the southerly winds and an increased velocity as winter conditions develop. The transition in the fall is in marked contrast with the vigor of the spring transition. The cooler seasons are more strongly influenced by northerly winds, and the warmer seasons by southerly winds. 3. Climatic Centers of Influence In the preceding section the average conditions of temperature, precipitation, and the direction and velocity of the winds have been summarized, but little effort was made to indicate the mode of opera- tion of the determining factors which produce and maintain these aver- age conditions. It is often true that the main factors which explain the conditions seen in some restricted locality can not be found within it because the local sample is only a very small part of a much larger problem. Thus no one attempts to find an explanation of the through- flowing upper Mississippi system within the state of Illinois; a larger unit of study is necessary. The region examined must extend to the headwaters. So, also, with most of the climatic features of Illinois; their approximate sources must be sought elsewhere. Let us there- fore consider some of the broader features which influence the climate of North America, particularly that of the eastern part. The climates of the world have been divided into two main kinds, depending primarily upon the controlling influence of temperature. This is due to the relative specific heat of land and water, that of water being about four times that of land. The sea, which covers three fourths of the earth’s surface, is thus an immense reservoir of heat, which is taken up and given off slowly, at a rate one fourth that of the land. It is therefore relatively equable. The northern hemisphere contains the largest amount of land, and is therefore less under the control of the sea than the southern hemisphere; yet the sea’s influence is very powerful, particularly near the shore. The large land masses, on the other hand, on account of their lower specific heat, receive and give off heat more rapidly to the air above. For this reason the tem- perature changes, as between day and night or summer and winter, are much more rapid and much more extreme over land than over the sea. A climate dominated by the equable sea is oceanic; that dominated by the changeable lands is continental. [Illinois lies far 70 from the sea and is therefore strongly influenced by continental con- ditions. To what degree is the marine influence shown? Meteorologists (cf. Fassig, ’99) have come to look upon the large areas of permanent high and low barometric pressure as among the most important factors in climatic control. There are five of these powerful “centers'of action” which influence our North American climate (Fig. 1), and four of these are at sea. A pair of Jows are in the far north, one in the north Pacific near Alaska, the other in the 80 60° 40 20° aw, aes pes oon NG Owe | ¢ seen Bs - Lea 160° 40° 120° 100° 80° 60° 40 2 Fig. 1. Diagram showing the positions of the relatively stable areas of high and low barometic pressure, and indicating their influences upon the evaporating power of the air and upon the climate in general. north Atlantic south of Greenland. A pair of highs are farther south, one in the Pacific between California and the Hawaiian Islands, and the other centering in the Atlantic near the Azores. The highs and lows in each ocean seem to be paired and to have some reciprocal rela- tion. The fifth center of action is upon the land. It is a high baromet- ric area in the Mackenzie basin of Canada, where it becomes a pow- erful center‘of influence through winter and spring, but with the prog- ress of summer conditions weakens, and through the accumulation of continental heat becomes converted into a low, thus there is a complete seasonal inversion on the continent. These large highs and lows, although relatively permanent, are con- tinually changing in intensity and position. The highs are regions of descending, diverging, warming, and drying air, producing clearing and clear air on their western side, but the reverse on their eastern side. 71 The lows are regions of ascending, converging, cooling air, with in- creasing moisture and clouds on their western side, but are the re- verse on their eastern side (Moore, ’10: 153). ‘These same character- istics apply to the small highs and lows which we are accustomed to see on the daily weather maps. If, now, we consider these large centers of action, such considera- tion will do much toward giving us a graphic idea of our climate. Dur- ing the winter, because of the small amount of heat received in the Mackenzie basin, the temperature becomes very low, and a powerful high barometric area is formed; then the descending air blowing from the eastern part of this high, or from small highs originating from the larger one, produce the cold winters and cold waves in winter which characterize the northeastern United States. If, however, the Atlantic high wanders on the eastern coast of the United States in winter, the western part of this high, with its descending, diverging, warming, and drying air, produces a mild winter. The climate of the eastern United States is thus, in the cold season, under the alternate invasion of these two powerful centers of action. During the warm season the conti- nental winter high is replaced by a low, due to the accumulating warm continental temperatures which thus have produced an inversion or seasonal overturning. But the Atlantic high is permanent and exerts its influence continuously. If the western part of this high encroaches upon the eastern United States during the summer, with its descend- ing, drying, and clear air, it may produce drouth, this depending, of course, on its degree of development. The continental low of sum- mer, with the drying influence of its eastern side, has a similar ten- dency. Thus the character of the summer is determined, to an im- portant degree, by the interplay and relative balance between these two warming and drying centers. ‘The activity of these centers has a pow- erful influence upon the moisture-bearing winds, which influence hu- midity and evaporation in Illinois, and in the eastern United States. 4. Relative Humidity and Evaporating Power of the Air We are now in a position to examine the facts of relative humidity and the relative evaporating power of the air in the eastern United States. The relative aridity on the plains east of the Rocky Moun- tains is due primarily to the removal of moisture from the prevailing westerlies in their passage from the Pacific over the various western mountain ranges which extend across their path, combined with the excessive summer heating of the continental mass. Here, then, is the influence of the continental summer low. Farther east the Atlantic high tends to supplement the continental low and to cause the Gulf 72 winds to brings moisture inland,* and the Great Lakes region adds its uota. : In the storm-track zone, where stagnation of the air is due largely to the balance existing between the continental low and the oceanic high, the aridity of the plains extends the farthest east, and as an arid peninsula it crosses Illinois, giving during August a relative humidity to the prairie area of 60-70 per cent. of saturation (Johnson, ’07). The reality of the arid peninsula across Illinois is further shown by the rainfall-evaporation ratios computed and mapped by Transeau (’o5). These ratios were determined by dividing the mean annual rainfall at each place by the total mean annual evaporation. These mapped percentages show that the prairie region is closely bounded by the region with an evaporation ratio of between 60 and 7o per cent. of the rainfall received. These conditions furnish a general background or perspective for a profitable consideration of the local and more detailed studies which have been made of the relative evap- orating power of the air in different plant and animal habitats. For our purpose it is not necessary to consider the history of meth- ods of measuring relative evaporation. ‘This measurement may be made by evaporating water in open pans or by the porous porcelain-cup method. Such cups have been devised by several students, but a modi- fied form of the Livingston atmometer has been mainly used by plant ecologists, and this was the kind we used at Charleston. Transeau (’08) was the first to use such an instrument and to show its value in studying the relation of intensity-of evaporation to plant societies. His work on Long Island, N. Y., showed very clearly that evaporation in open places was much greater than in dense forests. These obser- vations were enough to show that evaporation is a factor related to the physical conditions of life upon the prairie and in the forest, and there- fore in our cooperative study of the Charleston area in 1910 relative evaporation was made a special feature in the study of representative environments, in order to determine its relation to both the plants and the animals. So far as is known this is the only study yet made in which these determinations have been recorded from the same places where the animals have been studied. Since our data were secured, several papers have been published on relative evaporation in different sorts of habitats in this state and in northern Indiana by plant ecolo- gists Fuller (’11, ’12a, ’12b), McNutt and Fuller (’12), Fuller, Locke, *Zon (’13) has recently asserted that the moisture from the sea does not -make a single overland flight inland, but rather is largely precipitated near the sea, is evaporated and carried farther inland, is precipitated again, and this process repeated again and again, so that its inland flight is a vertical revolv- ing eycle of precipitation and evaporation. If this contention is valid, evapo- ration from the land is a much more important climatic factor than it is usually thought to be. 73 and McNutt (’14), Sherff (’12, ’13a, ’13b), and Gleason and Gates (12). Shelford (’12, ’13a, ’13b, “14a), utilizing the evaporation data of the plant ecologists, has applied the same to animal associa- tions also, and he has further tested some of these ideas experiment- ally in the laboratory. In Ohio, Dachnowski (’11) and Dickey (’09) have made records of data obtained by the use of the porous cup, and in Iowa Shimek (’10, ’11) has used the open-pan method. Mention should also be made of Yapp’s observations (’09) on a marsh in Eng- land. A very important summary of evaporation records, in the open and in forests, is given by Harrington (’93). The effect of wind- breaks upon evaporation has been studied by Bates (’11) and Card (97). Finally, mention should be made of Hesselman’s studies of relative humidity in forest glades in Sweden (’04). Our records from the Charleston region will be given first, and then their significance will be discussed. The unglazed porcelain cups, with a water reservoir, were placed so that the tops of the cups were about six inches above the soil in the habitats examined, and at weekly in- tervals the water loss was measured. The instruments were in opera- tion simultaneously, so that the results are comparable. The standard instrument was located in the open exposed garden of the Eastern Illinois Normal School at Charleston, which was considered as unity, or 100 per cent. For further details as to the conditions where the atmometers were located consult the description of the stations and the photographs. An examination of the diagram (Fig. 2) will show that although ‘based upon a limited amount of data (for less than a month, from 10 20 30 40 50. 60 70, 80 90 i00 Intensity of evaporation..........+. Standard, open garden, Normal School Sta. III, b. Mixed prairie and young forest Sta. II, a. Grassy area, Panicum Sta. II, uw. Grassy area, Euphorbia Sta. IV, w. Upland, open woods Sta. III, a. Silphium on Diack soil Sta. II, a. Colony of 8. laciniatum Sta. IV, b. Ravine slope, open woods Sta. IV,c. Dense climax forest cover Fie. 2, Diagram of the relative evaporation in different prairie and forest habitats, showing the great reduction in evaporation with the development of a closed forest canopy of a climax forest; Charleston, Illinois. 74 August 19 to September 22) the facts are in harmony with similar studies elsewhere covering a much longer period, so that there is valid reason for confidence in them. The standard instrument was located, as already mentioned, in an open, exposed cultivated garden, where the intensity of evaporation was very high. The black soil prairie areas, Stations II and III, a, have an average of 56.1 per.cent.—a condition much like that in the grassy-Euphorbia prairie at Loxa (Station II, a) —or a little more than half that of the standard instrument. The dry upland area of mixed prairie and young forest, on gray silt loam (Sta- tion III, b), has an intensity of 80 per cent. This is in the region of the most extensive grassy prairie about Charleston; the general ap- pearance of the region is shown in Plate XIII. A surprising feature of the table is the evaporation in the open-crowned upland oak-hickory woods (Station IV, a). In this forest perhaps two thirds to three fourths of the ground was shaded, and it was very well drained. The evaporation here reached 54.2 per cent., being very near that of the average of the black soil prairie (56.1 per cent.). I had anticipated much less evaporation than on the prairie, a position more intermedi- ate between the prairie and the lowland forest, or about 42 per cent. (cf. Harvey, 14:95). The ravine slope (Station IV, Db), although somewhat open, has 31.5 per cent.—a very low rate of evaporation— and is remarkably close to that of the densely crowned lowland for- est (Station IV, c), at 26.9 per cent. The decline, however, in the intensity of evaporation with the degree of completeness of the for- Per cent. of standard............+-- 0 40 60 80 (00 120 Sta. 11. Salt marsh outer margin Sta. 3. Gravel slide, open Sta. 1. Carnegie garden, standard Sta. 9 and 10. Upper beach Sta. 12. Salt marsh, inner margin Sta. 2. Garden, high level Sta. 4. Gravel slide, partly invaded Sta. 5. Forest, open Sta. 13. Fresh-water marsh Sta. 6. Forest, typical mesophytic Sta. 7. Forest, ravine type Sta. 8. Forest swamp type Fic. 3. Diagram of the relative intensity of evaporation in the lowest stratum of different kinds of habitats, Long Island, N. Y. (After Transeau.) 75 est crown, is strikingly shown in passing from the open upland woods, at 54.2 per cent., to the ravine slope at 31.5 per cent., and on to the lowland forest at 26.9 per cent. A comparison of these results with those secured by Transeau (08) on Long Island, is instructive. His standard instrument was also in an open garden (Fig. 3), comparable with the Charleston standard. A gravel slide, partly invaded by plants, had an evaporation of 60 per cent., comparable with the open prairie at Charleston; the open forest, 50 per cent., comparable with the upland open Bates woods at 54.2 per cent.; and the mesophytic forest, 33 per cent., com- parable with the ravine and lowland places in the Bates woods at 31.5 and 26.9 per cent. respectively. Association Blowout (basin) 1.56 Blowout (slide) 1.27 Bunchgrass (Leptoloma consoc.) 1.18 Bunchgrass (Eragrostis trichodes con.) 1.04 Standard 1.00 Beach 0.93 Quercus velutina woods 0.66 Quercus velutina 0.55 Willows (Acer part) 0.56 Willows (Salix part) 0.44 Mixed forest (margin) 0.36 Mixed forest (center) 0.29 Fig. 4. Relative intensity of evaporation in different kinds of habitats on sandy soil, Havana, Illinois. (After Gleason and Gates.) Another series of relative evaporation observations was made by Gleason and Gates (’12) on sandy soils at Havana, Illinois. As their methods were similar to those used at Charleston, useful comparisons may again be made. The standard instrument was in an open area comparable to the garden at Charleston. An examination of Figure 4, summarizing the results of their study, shows that upon the grass- covered sand prairie (bunch-grass) the evaporation was about 110 per cent., that in open black oak (Q. velutina) woods (on sand) it was about 60 per cent., and that in a denser hickory-black-oak-hackberry mixed forest (somewhat open) it was about 31 per cent. There is thus a close general correspondence between the conditions at Havana and. Charleston, although the evaporation upon sand prairie appears to be relatively much greater than upon the black-soil prairie. Fuller (’11) and McNutt and Fuller (’12) have made comparative studies in different kinds of forest in northern Illinois and in northern 76 Indiana. Their results are combined and summarized in Figure 5. This diagram shows the relative evaporation near the surface of the soil, the standard of comparison being the evaporation im a maple- beech climax forest, where evaporation is relatively low. The aver- age daily amount, in c.c., shows that there is a progressive increase in evaporation as follows: 8.1 c.c. in a maple-beech forest, 9.35 c.c. in the oak-hickory upland forest, 10.3 c.c. in an oak dune forest, 11.3 c.c. in a pine dune forest, and an increase to 21.1 c.c., on the cottonwood dunes. This expressed on a percentage basis is, in inverse order, re- spectively 260 per cent. in the cottonwoods, 140 per cent. in the pines, 127 per cent. in the oak dunes, 115 per cent. in the oak-hickory for- est, and 100 per cent. in the maple-beech forest. 20.40 60 @0 100 (120 (140 (160180 200 22Q_ 240 260 280 Intensity of evaporation Sta. A. Cottonwood dunes Sta. B. Pine dune Sta. C. Oak dune Sta. D. Oak-hickory Sta. E. Maple-beach forest % Standard Fic. 5. Diagram showing the relative rate of evaporation in different kinds of forest in northern Illinois and Indiana. [Data from Fuller (’11) and McNutt and Fuller (’12).] Shimek (10, ’11) has made valuable observations on the relative rate of evaporation on the prairie of western Iowa. He used the open- pan method in four representative habitats. His results show very clearly that the rate of evaporation is much greater in exposed places than where there is shelter from the sun and wind. I have put his data in a form comparable with those which have just been discussed (Fig. 6), and have made the cleared field area, Station 4, the standard of comparison, as it more nearly approaches the standard used at Charleston and by others. Station 3 is on a high bluff, exposed to the Intensity of evaporation............ . 2040. 6080100120, 40160 180-200 Sta. 3. Open, much exposed prairie vegetation Sta. 1. Open, exposed slope of bluff, prairie Sta. 4. Open, cleared area, partly pro- tected Sta. 2. Bur-oak grove, protected Fic. 6. Diagram of relative evaporation in prairie and forest habitats, in western Iowa. (Data from Shimek.) 77 west and south winds, and, as might be expected, it has an excessive evaporation—184 per cent. Station 1, also covered by prairie vegeta- tion, and exposed to west and southwest winds but sheltered from winds from the south and southeast, also shows a very high evapora- tion—132 per cent. Station 4, which was made the standard, had been cleared of forest, and was an open place protected by a ridge. Station 2 was apparently a dense grove composed of bur oak, basswood, elm, and ash, with considerable undergrowth. Here the rate of evapora- tion dropped considerably—to 36 per cent. The general character of this forest calls to mind the denser oak forests on sand at Havana, Illinois. An important feature of these observations is that they were made far out upon the “prairie”, bordering the plains, most other studies on relative evaporation having been made much farther east. In Ohio, Dachnowski (’11) and Dickey (’09) have recorded the relative evaporation of the air, using a campus lawn as unity. In the central grass-like area of a cranberry bog the evaporation was 69.2 per cent., and in the marginal maple-alder forest it was 51.2 per cent. Harrington (’93: 96-102), in summarizing European studies on the relative evaporation (with a water-surface as standard) in the open and in German forests shows that the “annual evaporation in the woods is 44 per cent. of that in the fields.” Compared with evapora- tion in the open, that under deciduous trees is 41 per cent., and that under conifers is 45 per cent.—a difference most marked in the sum- mer. Ebermeyer’s Austrian observations (I. c. :99) show that the “evaporation from a bare soil wet is about the same as that from a water surface,” both in the open and in the forest. A saturated soil under forest litter gives an evaporation of only 13 per cent. of that of a free-water surface in the open. Harrington (l.c.: 100) con- cludes that “About seven-eighths of the evaporation from the forest is cut off by the woods and litter together.” Sherff (’13a, ’13b) has shown that in the Skokie Marsh, north of Chicago, the absolute amount of evaporation near the soil was less at the center of a Phrag- mites swamp than at its margin (Fig. 7), that a swamp meadow Intensity of evaporation.........-+++ 20.40 60-80 _—100-— 120-40 S0_—*180 200 Sta. D. White oak-ash forest Sta. B. Phragmites swamp, margin Sta. OC. Swamp meadow Standard Sta. A. Phragmites swamp, center Fig. 7. Diagram of relative evaporation in Skokie Marsh area, near Chicago, at 10 inches (25 cm.) above the soil. Recaleulated. (Adapted from Sherff.) 78 was in an intermediate position, and that in an adjacent white oak-ash forest evaporation was about twice as much as in the swamp meadow. Sherff used as standard the forest (D). This gave him for the center of the swamp (A) 38 per cent., for the swamp meadow (C) 54 per cent., and for the outer swamp margin (B) 105 per cent. In Figure 7, I have used his swamp meadow as 100 per cent., and by recalcula- tion this gives the forest (D) 185 per cent., for the swamp margin (B) 105 per cent., and for the center of the swamp (A) 70 per cent. These figures indicate a concentric arrangement of the conditions of evap- oration about the swamp. Intensity of evaporation........6.004. 10.20 30 40 s0 60 70 80 90 1000 1907: 7 Sta. A. Above vegetation. 4 feet, 6 inches above soil Sta. B. Middle of vegetation. 2 feet, 2 inches above soil Sta. C. Lower vegetation. 5 inches above soil 100%,| 1908: Sta. A. Above vegetation. 5 feet, 6 inches above soil Sta. B. Middle of vegetation. 2 feet, 2 inches above soil Sta. C. Lower vegetation. 5 inches above soil 100% Fie. 8. Diagram showing the relative evaporation at different vertical levels in a marsh in England, the evaporation in the lower layers of the vegetation being much greater than in the upper strata or in the air above it. (Data from Yapp.) Thus far, attention has been devoted solely to the horizontal differ- ences in evaporation. There are also important vertical ones, vary- ing above the surface of the substratum. Important observations on this subject have been made, by a porous-cup method, in an open grassy marsh in England, by Yapp (’09). ‘The vegetation grew to a height of two to five feet. From his data the accompanying diagrams (Figs. 8, 8a) have been prepared. This shows that when the stand- ard was made the rate of evaporation above the general level of the vegetation, within the grass layer evaporation was reduced from about one half (Sta. B, 1908, 56.2 per cent.) to one third (Sta. B, 1907, 32.8 per cent.) at 2 feet 2 inches above the soil; and that at 5 inches above the soil it was reduced to between one fourteenth (Sta. C, 1907, 6.6) and one seventh (Sta. C, 1908, 14.7) of that above the vegeta- tion. Yapp (1. c.: 298) concludes from his studies that “In general, the results of the evaporation experiments show that the lower strata of the vegetation possess an atmosphere which is continually very much 79 more humid than that of the upper strata, and farther, that the higher and denser the vegetation the greater these differences are.” This is shown in Fig. 8a. Intensity of evaporation............. jo 20,3040 50 S60 70 S80 S900 ~—s100 Sta. A. 60 inches above ground, above vegetation 100% Sta. B. 12 inches above ground among vegetation mK Sta. C. 3 inches above ground, among vegetation ope Fic, 8a. Diagram showing the zelative evaporation at different vertical levels in a marsh in England, the evaporation in the lower layers of the vegetation being much greater than in the upper strata or in the air above it. (Data from Yapp.) In America only a few records have been made on vertical gra- dients in evaporation, two of these in marsh areas, one in Ohio by Dachnowski (’11), and the other near Chicago by Sherff (’13a, ’13b). The Ohio observations, made upon a small island in a lake, in a cran- berry-sphagnum bog, show that the rate of evaporation above the vege- tation is much greater than among it, and that this diminishes as the soil is approached, these results agreeing with those obtained by Yapp. Sherff’s observations were made in Skokie Marsh, north of Chicago, and show that the relative evaporation also varies with different kinds of swamp vegetation. From his data a diagram has been made (Fig. 9) in which the rate of evaporation in the upper part of the reeds Intensity of evaporation.......-..+6+ to 20 30 40 SO 60 70 680 90 100 Phragmites Sta. A. Within vegetation, 198 em. (77 100% inches) above soil. Standard. Sta. B. Within vegetation, 107 cm. (42 709 inches) above soil Sta. CO. Within vegetation, 25 cm. (10 inches) above soil 53) Sta. D. At soil surface 33% Typha Sta. A. Within vegetation, 175 cm. (69 inches) above soil Sta. B. Within vegetation, 107 cm. (42 inches) above soil Sta. C. Within vegetation, 25 cm. (10 7 inches) above soil Sta. D. At soil surface Fig. 9. Diagram of relative evaporation at different vertical levels above the soil within the vegetation of Skokie Marsh. (Adapted from Sherff.) 80 (Phragmites) at 77 inches is taken as 100 per cent. or the standard. Lower down, at 42 inches, the rate is 70 per cent., at 10 inches, 53 per cent., and at the surface, 33 per cent. Among the cattails (Typha), in the upper part of the vegetation, at 69 inches evaporation was 85 per cent.; at 42 inches it was 36 per cent.; at 10 inches, 20 per cent.; and at the surface, 8.5 per cent. These results show that at successively lower levels in the vegetation the rate of evaporation is greatly re- duced. They tend also to confirm the results of Yapp and Dachnow- ° ski. It seems, then, fair to conclude that the rate of evaporation above the swamp vegetation increases rapidly with downward progression, and probably with upward progression also. A vegetable layer, com- parable to the mulching of straw used by gardeners, thus acts as a pow- erful conserver of moisture. There are great differences within a few vertical feet in the open; what is the condition within the forest ? Intensity of evaporation............. 20 40__ 60 8010012040 160180 Sta. A. Maple-beech forest. 6 feet (2 m.) above soil Sta. B. Maple-beech forest. 10 inches (25 cm.) above soil Sta. O. Maple-beech forest. On slope of ravine 30 feet deep (10 m.) 13.3 feet (4 m.) below general surface. Fig. 10. Diagram showing the relative evaporation in a beech-maple woods, six feet above the soil (A), near the surface of the soil (B), and in a ravine (C). [Adapted from Fuller (’12).] The character of vertical differences in evaporation within the for- est has not been given as much attention as the similar changes in the open; but attention has already been called to the moisture-conserving effect of a forest litter, the evaporating rate in one instance being only 13 per cent. when compared with that from a water surface in the open. McNutt and Fuller (’12) have shown that grazing in an oak-hickory forest changed the average daily rate of evaporation for 189 days from 9.89 c.c., in the ungrazed forest, to 12.74 c.c., in the grazed for- est, at Palos Park, Ill. There are thus, within the forest, changes in evaporation with differences both in the ground cover and in the litter on the forest floor which correspond to the change in the vegetation in open places. Vertical differences in evaporation have been tested in a maple- beech-forest in northern Indiana by Fuller (’12b), who used the po- rous-cup method. His results have been summarized in Figure 10. This diagram shows that the evaporation at six feet above the surface is nearly twice as much as that at 10 inches above the surface, and 81 that in a ravine, 13.3 feet (4 m.) below, it was 80 per cent. of that 10 inches above the surface. The relative seasonal activity from May to November is shown in Figure 11. This diagram shows that after the leaves appear the highest evaporation takes place in July. This is probably the critical season for some animals. MAY JUNE JULY AUGUST | SEPTEMBER | OCTOBER F1¢. 11. Diagram showing the average daily rate of evaporation in beech-maple forest, six feet above soil (a), near the surface of soil (b), and in a ravine (ce). (From Fuller.) In the forest, Libernau (Harrington, ’93: 34) found that the “rela- tive humidity increases and decreases with the absolute humidity, whereas it is known in general, and also at the Station in the open country, that these two climatic elements are inverse. This is ac- counted for by the fact that the forest is a source of atmospheric aqueous vapor as well as of cooling.” (L.c. : 104: “The absolute humidity decreases in the forest from the soil upwards. The rate of decrease is usually the greatest under the trees and the least at the level of the foliage. The rate above the trees is intermediate between the other two. ‘This rate is least in the late hours of the night, when it may be zero. It increases with the increase of the temperature of the air, becoming greatest in the midday hours, when, under exception- ally favorable circumstances, it may make a difference of 10 per cent. 82 or even more. Occasionally, in high winds, the absolute humidity is greater over the trees. Over the field station the daily progress of ab- solute humidity was about the same as in the forest, but the maximum difference was only about half as great. The absolute humidity in and above the forest is greater than that over the open fields, and there is some trace of an increase of this difference to the time of maximum.” A greater relative humidity has been found over evergreen trees than over deciduous trees, which is slight (1.c.: 104), but the psy- chrometer was close to the evergreens and farther above the decidu- ous ones. Intensity of evaporation............. Sta. A. 20 rods (330 ft.) from wind- break, 25 to 40 feet high. Standard Sta. B. 12 rods (198 ft.) from wind- break Sta.C. 3 vods (49.5 ft.) from wind- break (July 15-Sept 15} 62 days, Lincoln, Neb Fig. 12. Diagram showing relative retardation of evaporation by a windbreak, Lincoln, Nebraska. [Adapted from Card (’97).] The border of the Illinois forest and prairie was characterized by tongues and isolated groves of forest and by glades. The forest had the same kind of influence as windbreaks upon the leeward areas and glades, and therefore the influence of windbreaks upon the evaporating power of the air is of interest. Card (’97) made a valuable study of this series of problems at Lincoln, Nebraska. The influence of wind- breaks upon evaporation is summarized in Figure 12. This diagram shows that leeward of a close windbreak ranging from 25 to 40 feet in height, the rate of evaporation in terms of the standard (A), which was 330 feet leeward, was 91 per cent. at a distance of 198 feet (B), and 71 per cent. at 49.5 feet (C), thus showing a marked reduction with proximity to the windbreak. These observations covered 62 days. Nearer to Illinois, similar though very limited observations were made in central Wisconsin by King (’95) which agree with Card’s on the retardation of evaporation by windbreaks. His results are shown graphically in Figure 13. Recently Bates (’11) has made an elaborate study of the effects of windbreaks upon light, soil, moisture, velocity of wind, evaporation, humidity, and temperature. His results confirm those just given and give additional facts which, however, with one exception, will not be mentioned. The paper itself should be consulted. This investigation by Bates shows that in proportion to the perfection of the windbreak 83 a quiet, stagnant air strip is formed to the leeward, and that this fa- vors excessive heating during clear days and low temperatures on clear nights. Years ago Harrington (’93: 119) suggested this idea and called attention to the close relation existing between the leeward con- ditions of windbreaks and forest glades. The glade climate is more rigorous, or extreme, than that upon plains (I.c.: 19, 84-88, 119). Such a climate is thus a bit more “continental” during the spring, sum- Intensity of evaporation............. i029 305060 080900080 Distance from windbreak 12 inches high: Sta. F. 500 feet leeward. Standard Sta. E. 400 feet leeward Sta. D. 300 feet leeward Sta. C. 200 feet leeward Sta. B. 100 feet leeward Sta. A. 20 feet leeward Fie. 13. Diagram showing the relative evaporation, May 31, at different dis- tances leeward of a windbreak, Almond, Wis. [Adapted from King (’95).] mer, and autumn. These glades are very hot in the early afternoon - and cool on clear nights, and the air is relatively stagnant; as Harring- ton says, it is “lee for winds from all directions.” The center of a dense forest may thus possess physical conditions quite different from those of the glade forest margin or in the open. Beginning with the relatively stable conditions within a forest toward its margin, the diur- nal temperature variations are much more extreme (Harrington, 1. c.: 89) “to a distance of a score or so of rods where it reaches a max- imum. The amplitude is greater in glades. Hence the extremes of temperature are exaggerated just outside the forest.” The annual soil temperatures of a glade are intermediate between that of the forest and the plain. The forest margin is thus seen to possess many of the char- acteristics of the glade, for its climate is somewhat more extreme than that in the open, far from the forest. 5. Temperature Relations in the Open and in Forests The temperature relations in open and forested regions are often very different. The density of the vegetable covering in the open and in the forests varies much and may have considerable influence upon animals. Yapp (’09) observed that the marsh vegetation in England 84 caused marked vertical differences in temperature in the vegetational stratum. He summarizes these results as follows (p. 309): “The temperature results show that the highest layers of the vegetation pos- sess a greater diurnal range of temperature than either the free air above or the lower layers of the vegetation. Regularly, especially in clear weather, both the higher day and the lowest night temperatures were recorded in this position.” Dachnowski (’12: 292-297) studied the temperature conditions in a cranberry bog substratum in central Ohio. He found that at a time when ice formed from 8 to 15 inches thick on the adjacent lake, in the bog it was only 3 to 5 inches thick, and there were small patches where it did not form at all. Ata depth of 3 inches in the peat the tempera- ture ranged from 33° to 77° F. (.5 -25.0° C.). In the bordering maple-alder zone, at 3 inches depth it ranged from 33° to 72° F. (.5°- 22.0 C.). His observations indicate that the temperature relations within the maple-alder zone are more stable than those in the open central area. Cox (’10) has also shown that the character of the vegetation in Wisconsin cranberry bogs has much influence upon temperature rela- tions in this habitat. It seems very probable that similar conditions hold over prairie vegetation, but I do not know of any observations on this point. We are all familiar with the common practice of gardeners of using a mulch of straw to retard temperature changes under it; prairie vegetation must have a similar influence. (Cf. Bouyoucos, ’13: 160.) The relative air temperatures within and without the forest show a distinct tendency to reduce the maxima and minima, and to lower the mean annual temperature. Harrington (’93:53) concludes, therefore, that “the forest moderates (by reducing the extremes) and cools (by reducing the maxima more than the minima) the tempera- ture of the air within it. The moderating influence is decidedly greater than the cooling effect.” ‘These effects are not uniform, but are much more marked in the summer, and Harrington further says: ‘The cool- ing effect tends to disappear in winter. The moderating effect is the most important one and it is the most characteristic” (p. 56). The temperature relations within the forest crown show that in general the effects are similar to those found at an elevation of about 5 feet. The maxima are lowered, the minima are elevated, and there is a cooling effect. The differences are most pronounced during the summer, and the temperatures are intermediate in position between ‘those at the five-foot level and those in the open (l.c.:66). Ata height of 24 feet, deciduous trees showed a marked summer cooling 85 effect, while evergreens showed much less, though they are much more uniform for 9 months of the year. Again, he says: “In summer the average gradient under trees is about +2°; that is, it grows warmer as we ascend at the rate of two degrees per 100 feet (31 m.). Out- side in the general average it grows colder by about a quarter of a de- gree.” This warmer air above the cooler in the forest favors its sta- bility or relative stagnation, although as a whole the forest air is cool- er and heavier than the surrounding air and tends to flow outward. ‘The forest thus tends to produce a miniature or incipient barometric high. In conclusion Harrington (p. 72) states that “The surface of , the surface of the forest is, meteorologically, much like the surface of the meadow or cornfield. The isothermal surface above it in sun- shine is a surface of maximum temperature, as is the surface of a meadow or cornfield. From this surface the temperature decreases in both directions.” In the case of a beech forest the warm diurnal layer above the forest crown was only 6.5 feet thick (p. 34). The conditions above the forest are thus representative of the at- mospheric conditions above dense vegetation in general, and are in per- fect harmony with Yapp’s observations upon the temperature above a marsh (’09: 309), quoted on a previous page, to the effect that tem- perature changes are extreme here, and greater than in the free air above or in the lower layers among the vegetation. The forest is thus to be considered as a thick layer of vegetation in its influence upon meteorological conditions. 'The conditions above the forest, there- fore, exemplify a general law. In general terms, the temperature of the soil below the zone of seasonal influence is that of the mean annual temperature for a given locality. The surface zone, however, varies with the season. Har- rington (’93) has summarized the German observations on the rela- tive soil temperatures in the open and in the forest. In the following quotation the minus sign indicates a forest temperature less than a cor- responding observation in the open. These temperatures were taken about 5 feet above the soil. He says (p. 43): “The average of the seventeen stations (representing about two hundred years of observa- tions) should give us good and significant results. It shows for the surface—2°.s9, for a depth of 6 inches (152 mm.)—1°.87, and for a depth of 4 feet (1.22 m.)—2°.02. The influence of the forest on the soil, then, is a cooling one, on the average, and for central Europe the cooling amounts to about two and a half degrees for the surface. The cooling is due to several causes: The first is the shade; the foliage, trunks, branches, and twigs cut off much of the sun’s heat, absorb and utilize it in vegetative processes, or in evaporation, or reflect it away into space. Thus the surface soil in the forest receives 86 less heat than the surface of the fields. The same screen acts, how- ever, in the reverse direction by preventing radiation to the sky, thus retaining more of the heat than do the open fields. The balance of these two processes, it seems from observation, is in favor of the first and the average result is a cooling one... .. . The differences of temperature at the depth of 6 inches (152 mm.) are more than half a degree less than at the surface. In this is to be seen the specific effect of the forest litter; it adds a covering to that possessed by the sur- face, so that while the deeper layer is cooled as much by the protec- tion from the sun’s rays as is the surface, it is not cooled so much by radiation of heat to the sky. Its temperature is, consequently, rela- tively higher, and approximates somewhat more the field tempera- tures.” “The forest soil is warmer than that of the open fields in winter, but cooler in the other seasons, and the total cooling is much greater than the warming one... ... The forest, therefore, not only cools the soil, but also moderates the extremes of temperature” (p. 46). The character of the forest, whether evergreen or deciduous, in- fluences the temperature conditions of the soil, as is seen by a com- parison of these conditions in the forest and in the open. The two kinds of forest are much alike in winter; during the spring the soil warms up more rapidly under conifers. Temperature variations are slightly greater under deciduous trees. 6. Soil Moisture and its Relation to Vegetation The moisture in the soil is derived largely from precipitation, but part of it, in some localities, comes directly from the adjacent deeper. soils or rocks, and thus only indirectly from precipitation. As Illinois lies at the bottom of a large basin, there must be some subsurface flow from the adjacent higher regions, but to what extent is not known. McGee (’13a:177) estimates that the general ground-water level— the level at which the soil becomes saturated—has, since settlement, de- clined 10.6 feet in Illinois. This decline is not limited to drained re- gions but is a general condition. In addition to these changes of level there are seasonal fluctuations. Sherff (’13a: 583) observed in Skokie Marsh that the water-table was at or above the surface in May, then declined until early September, and then rose rapidly to the surface by the middle of October. The wet prairie at Charleston has undergone just such changes as these; the ground-water level has been lowered and there are marked seasonal changes. Harvey (’14) has recently shown that the soil of Eryngium-Sil- phium prairie at Chicago contains a large amount of water during 87 April and until late in May; that the moisture falls and is low during July and August, with a mean of 24 per cent. of saturation for these months; but that in October the soil is again at or near the point of saturation. The blanket of humid air which accumulates under a cover of vege- tation, retards evaporation and conserves soil moisture. The denser the vegetation the more marked is its influence. The litter—the or- ganic debris in an early stage of decomposition—on the forest floor has the same tendency, and has even a greater water capacity than the soil itself. On the other hand, a forest is a powerful desiccator; as Zon (’13:71) has recently put it: “A soil with a living vegetative cover loses moisture, both through direct evaporation and absorption by its vegetation, much faster than bare, moist soil and still more than a free water surface. The more developed the vegetative cover the faster is the moisture extracted from the soil and given off into the air. The forest in this respect is the greatest desiccator of water in the ground.” This drying effect is shown particularly near the surface of the soil, where roots are abundant and where drouth is so marked that it may prevent the growth of young plants here (cf. Zon and Graves, “11: 17-18). Warming (’09:45) says: “It may be noted that, according to Ototozky, the level of ground-water invariably sinks in the vicinity of forest, and always lies higher in an adjoining steppe than in a forest; forest consumes water.” McNutt and Fuller (12) have made a study of the amount of soil moisture at, 3 inches (7.5 cm.) and at 10 inches (25 cm.) below the surface in an oak-hickory forest, at Palos Park, Illinois. They found that the percentage of water to the dry weight of the soil at the 3-inch level averaged 18.9 per cent. and at 10 inches was 12.5 per cent. of the dry weight of the soil. The greater moisture near the surface is due to the humus present in this layer. The grazed part of the forest possessed less soil moisture, and shows the conserving effect of vege- tation. (Cf. also Fuller ’14.) The artificial control of soil moisture is well shown by the effect of windbreaks. Card (’97) studied the moisture content of the soil to leeward of a windbreak and found that in general there is a “de- crease in the per cent. of water as the distance from the windbreak increases.” As the physical conditions leeward of windbreaks are similar in many respects to those in forest glades and forest margins, it is very probable that the conditions of soil moisture also will be very similar in these places. 88 7. Ventilation of Land Habitats The preceding account of the temperature, humidity, and evapo- rating conditions in various habitats forms a necessary basis for an un- derstanding of the processes of ventilation or atmospheric change in land habitats. The differences in pressure due to the different densi- ties of cool and warm air and to the friction and retardation of mov- ing air currents, determine to an important degree the composition of the air in many habitats. In such an unstable medium as air, changes take place very rapidly through diffusion, and through this constant process of adjustment there is a tendency to level off all local differences. These are naturally best preserved where diffusion cur- rents are least developed—in the most stagnant or stable atmospheric conditions; therefore any factor which retards an air current and pro- duces eddies, or slow diffusion, will favor local differentiation of the air. We have seen that any vegetable cover retards air currents, so that the air within the vegetation becomes different from the faster moving air above it. The accumulation of humidity at different levels above the soil within the vegetation, clearly shows this. The denser the vege- tation-the more completely are the lower strata shut off and, to a cor- responding degree, stagnant and subject'to the local conditions. Two factors have an important influence upon these conditions: the charac- ter of the cover itself, and the character of the substratum. If both of these are mineral rather than organic, in general comparatively little local influence is to be expected, although in some localities CO, escapes from the earth and on account of its density may linger in de- pressions and thus kill animals (Mearns ’03). Generally, however, the organic materials are of most importance both as a cover and asa substratum, and are often the source of carbon dioxide. Living vege- tation may also add oxygen to such stagnant air, but the main source of it is the free air itself. The forest litter, on account of its imperfect stage of decay, consumes oxygen and gives off carbon dioxide; in the humus below it, shut off even more from free access to air, the carbon dioxide is relatively more abundant and the oxygen relatively less so or absent; and in the deeper mineral soil the amount of carbon dioxide is relatively less on account of the absence of organic debris, and a small amount of oxygen is present. The aeration of the soil is influenced to a large degree by its poros- ity; the looser it is, the freer the circulation. Buckingham (’04) has shown that “the speed of diffusion of air and carbonic acid through these soils was not greatly dependent upon texture and structure, but was determined in the main by the porosity of the soil. . . . the 89 rate of diffusion was approximately proportional to the square of the porosity . . . . the escape of carbonic acid from the soil and its replacement by oxygen take place by diffusion, and are determined by the conditions which affect diffusion, and are sensibly independent of the variations of the outside barometric pressure.” In the upper, better ventilated, moist, neutral or alkaline layers of vegetable debris decomposition is brought about mainly by the agency of fungi; but in the deeper, poorly ventilated acid layers, lacking oxy- gen, bacteria are the active agents (cf. Transeau, ’05, 06). The higher the temperature the more rapid the circulation, and on this ac- count ventilation in the open is relatively more rapid than in the cooler woodlands. The black soil prairies are thus favorable to a higher tem- perature and better ventilation. Dry soil, according to Hilgard (706: 279) contains from 35 to 50 per cent. its volume of air, and in moist or wet soils this space is replaced by water. Thus the condi- tions which influence the amount of water present have a very im- portant influence upon aeration. As water is drained from the soil, air takes its place; so drainage and the flow of water through the soil facil- itate ventilation. The part of the soil containing air is thus above the water-table; and as this level fluctuates with the season and from year to year the lower boundary of this stratum is migratory. Hilgard states that cultivated garden soil contains much more air than uncul- tivated forest soil, Warming (’09: 43) says that the “production of acid humus in the forest leads to an exclusion of the air.” If lime is present, such an acid condition can not arise. While the source of oxygen in the soil is the air, the reverse is the case with carbon dioxide. The surface layers of the soil, among dense vegetation, constitute an area of concentration of carbon dioxide. Because this is more soluble than other gases, it is found in rain water, according to Geikie, in a proportion 30 to 40 times greater than in the air. Rains thus assist in the concentration of carbon dioxide in the soil. This concentration is well shown by the following table by Baussungault and Lewy (Van Hise, ’04: 474). co, in Character of soil air 10,000 parts by weight 1. Sandy subsoil of forest 38 2. Loamy subsoil of forest 124 3. Surface soil of forest 130 4. Surface soil of vineyard 146 5. Pasture soil 270 6. Rich in humus 543 90 The amount of carbonic acid in the atmosphere is by weight about 4.5 parts in 10,000, The amount in the air is, as Van Hise says, “‘in- significant in comparison with the amount in soils in regions of luxu- riant vegetation. In such regions the carbon dioxide is from thirty to more than one hundred times more abundant than in the atmosphere.” This carbonic acid in the presence of bases, sodium, potassium, cal- cium, and magnesium compounds, forms carbonates and bicarbonates. This is the process of carbonation—one of the most important proc- esses of change in surface soils. In view of the dominance of CO, in soils we may anticipate that many of the animals living in them possess some of the characteristics of the plants, bacteria, fungi, etc., which are active in such soils. The anaerobic forms live without free oxygen; others live only where oxy- gen is present. The animals which thrive in the soil are likely to be those which tolerate a large amount of CO, and are able to use a rela- tively small amount of oxygen, at least for considerable intervals, as when the soil is wet during prolonged rains. This is a subject to which reference will be made later. : The air is the main source of oxygen, and from the air it diffuses into the soil; thus the process of equilibration is constantly in progress. Carbonic acid, also present in the air, is washed down by rain and concentrated in the soil, where it is increased by the decay of organic debris and by respiring animals to such an extent that it exists under pressure and diffuses into the air, thus contributing to the air. In the soil, then, the process of decarbomzation is of great importance to animal life, and must not be neglected. The optimum soil habitat is therefore determined, to a very important degree, by the proper ratio or balance between the amount of available oxygen and the amount of carbon dioxide which can be endured without injury. The excessive accumulation of carbon dioxide, an animal waste product, is compar- able to the accumulation of plant toxins which may increase in the soil to such a degree as to inhibit plant growth. Such substances must be removed from the soil, or changed in it to harmless com- pounds, or plants and animals can not continue to live in certain places. I have used the term ventilation to cover both the oxygena- tion and decarbonization of land habitats, and the same principles are applicable to life in fresh-water habitats. We have just seen how atmospheric ventilation favors the removal of certain injurious waste products from the air and soil. In addition to gaseous waste products there are also liquids and solid kinds which may be equally harmful in a habitat. These are known to exist in con- fined liquids, as in aquaria (Colton, ’08; Woodruff, ’12), where they 91 interfere with the welfare of the animals present, and it is probable that they also exist in soils. The older naturalists elaborated the idea that if organisms were not such active agents in the destruction or transformation of plant and animal bodies such remains would soon encumber the earth. Thus organisms themselves are among the most active agents in influencing directly and indirectly the ventilation of animal habitats. 8. The Tree Trunk as a Habitat A living tree trunk is composed of wood, sap (moisture), and bark, all of which are relatively poor conductors of heat. When the trunks are cooled, as in winter, they are slow in warming, not only because of poor conduction but also because of the slow circulation of sap, which is derived from the cool ground-water. As the season progresses, the trunks warm up, this process being retarded in part by the shade and the cool forest conditions; and in the fall, radiation of the heat accumulated also takes place slowly. The tree trunk therefore changes its temperature slowly, as does the soil. The animals which live within wood thus live in a relatively cool and stable environment. In living trees the humidity is relatively high, as it may also be in fallen, decaying logs. Relatively dry logs, before progress of decay, on the other hand, form a relatively dry and uniform habitat. (Cf. on the temperature of trees: Harrington, ’93, pp. 72-75; Packard, ’90, p. 23; and Jones, Edson, and Morse, ’03, pp. 97-100.) 9. Prairie and Forest Vegetation and Animal Life The dependence of animals upon plants for food is one of the most fundamental animal relations. It is a world-wide relation, but its mode of operations varies greatly in different environments. For ex- ample, many years ago, Brooks gave us a graphic picture of the réle of marine vegetation in the economy of marine animals. In the sea there are no forests or grasslands, and no corresponding animals as- sociated with these conditions, as on land; but in the sea great numbers of minute plants float, and upon these feed an immense number of small crustaceans and other small animals. These small creatures occur in such large numbers that at times the sea is a sort of gruel which sedentary and stationary kinds may appropriate by simply al- lowing the sea to flow into their mouths. The food here circulates in their environmental medium, as plant foods do in the soil and air. This condition has made it possible for vast numbers of plant-like animals to grow over the sea floor as plants do over rocks and plains. The living meadows of animals thus furnish pasture for a host of preda- 92 \ ceous kinds; and upon these still others prey, so that flesh-eating ani- mals make up the most conspicuous classes of marine animals. Quite otherwise are the conditions on land, where no air current carries food to the hungry mouths of animals. Plants with roots in the soil and stems in the air are able, however, to secure their food from the cir- culating medium, but being themselves fixed, they are easy prey to animals—both the sedentary kinds, which live in or upon the plant tis- sues, and the active wandering kinds, which forage over large areas. The predaceous animals, either by active mind or body, must secure their food from the plant-feeding kinds. The great expanses of grass- land and forest tend to be devastated by a vast army of animals which far outnumber the predaceous kinds. The conditions of life, there- fore, found upon grassland areas, like the prairie, and in the forest, are to the farthest possible extent removed from those found in the sea. ‘This, then, is one of the most fundamental contrasts in the con- ditions of existence encountered by animals. These considerations naturally raise the question to what extent and in what particular manner does land vegetation influence animal life? Does a change in the vegetation as great as that between the for- est and the prairie have a marked influence upon animals? In the Charleston region we have just such a difference in the vegetation. Many years ago Bates pointed out repeatedly in his “The Natural- ist on the River Amazons” that the animals of that densely forested region were to a marked degree distinctly arboreal and “adapted” to a forest life. In most densely forested regions like conditions probably prevail, and to a corresponding degree open lands harbor animals equally characteristic and as truly terrestrial in habits. The contrast between the conditions of life in the open and in the forest is one of the most fundamental environmental conditions upon land. The sig- nificance of this contrast seems to have been realized only in part. The prairies or grasslands are representative of only one kind of open; they are caused by many kinds of factors limiting the extension of forests. Open places are formed by lakes, ponds, and swamps; by the avenues through forests formed by different kinds of streams, as brooks, creeks, and rivers; by the small amount of soil on rock sur- faces; and by still other kinds of limiting influences, such as the sea, severe climate, and altitude. Among almost all of the major taxo- nomic groups of land animals is seen the independent origin and pres- ervation of animals suited for life in the forest; this clearly points to the extensive influence and antiquity of this environment. The same is true of animals living in the open. But to assume that it is solely the kinds of forest trees serving as food for animals, or the cor- responding kinds of vegetation in the open, which determines whether 93 an animal lives in the open or in the forest, would be unwarranted in the light of the preceding discussion of the effect of vegetation upon air temperatures, winds, humidity, relative evaporating power of the air, and corresponding changes in the soil. Animal life is’ most abundant in a narrow vertical layer above the earth’s surface, by far the most of it is within a few inches or feet of the surface; and above the level of the forest-crown it diminishes with great rapidity. Be- low the surface of the soil the same general law holds; most of the ground animals are within the first few inches of soil, only a small number extending a few feet below the surface, and those found at greater depths being indeed very few. The rate of decline is many times more rapid below the surface than it is above it. There is, then, above and below the surface a rapid and progressive attentuation of the favorable conditions for animals and plants, and the animals do not establish thriving communities far from those physical conditions which are also favorable to vegetation. Animals are dependent upon plants for food, but both are dependent upon a certain complex of physical conditions near the surface of the earth. It is well to recall at this point how the influence of the climate and the vegetation exemplify certain general laws which operate in all hab- itats. The differentiation of habitats upon the earth is primarily due to temperature and the specific heat relations of the earth, which re- sult in the several media—gases, liquids, and solids. With a higher temperature all would be gas, and with a lower one all would be solidi- fied. The present intermediate conditions, therefore, permit the pres- ent differentiation. These media are further differentiated by tem- perature about as follows: Since the source of solar energy, heat, and light, and the oxygen supply, are above the surface of the earth, the vertical attenuation of these influences is one of the most striking peculiarities of animal habitats, both in water (where the causes have long been recognized) and upon land. Any covering of the earth, even the surface layer of vegetation, soil and water, tends to shut off heat, light, and oxygen. At the same time such a layer tends to shut in those influences which originate primarily in or below it. Thus car- bonic acid originating under the cover, by organic decay, breathing animals, or bacteria, or washed in by the rain, tends to be shut in. Furthermore, heat once reaching here, either in water or on land, tends toward slow radiation. ‘Thus we may look upon the surface layer as a partition which is under pressure from both sides, and through which constant interchange is in progress, as the process of dynamic equili- bration operates. ‘: 94 This attenuation of intensities, above and below the surface, pro- duces vertical layers of relatively equal strength or pressure. Thus the attenuation of temperature in gases (air) and in liquids (water) causes different densities in air and in water which modify to an im- portant degree the physical and chemical conditions in these media. This results in their stratification: when the heavier layers are below, stability is the tendency; and when the reverse order obtains, a change takes place toward the stable condition. With stratification, flowage tends to occur within the strata, and to be horizontal rather than vertical; additional pressure is therefore necessary to cause the vertical currents or circulation under such conditions. This is why carbonic acid accumulates in the soil and in small deep lakes abound- ing in organic debris, this accumulation being largely due, in both cases, to the slow rate of exchange caused by the stratification pro- duced by differences in density. This same relative stagnation is a primary factor in the vertical differences in the relative evaporating power of the air within a vegetable layer of the prairie or the forest. Though on the prairie the vegetational layer is generally but a few inches or a few feet thick, in the forest it is about eighty feet, or more, thick; and the forest thus influences atmospheric conditions solely as a thick layer of vegetation. Differences, then, in the character, structure, or composition of the surface of the substratum are of fundamental importance in under- standing its relative influence upon animals. Primarily these differ- ences are due to temperature, secondarily to temperature in combina- tion with moisture; and they result in the relative humidity and the relative evaporating power of the air. The most important difference in the surface layer in the Charleston region is that of prairie and for- est, and therefore the main features of these habitats will now be sum- marized. It should not be overlooked that conditions on the prairie are likely to be quite representative of open places in general, though they will probably be somewhat unrepresentative in the case of open places having wet or extremely dry substrata. It is also true that the condi- tions produced by the forest are comparable, in some degree, with those due to the influence of an elevation. 95 SUMMARY oF ENVIRONMENTAL FEATURES OF THE PRAIRIE AND THE DECIDUOUS FoREST —TEMPERATURE, HUMIDITY, AND EVAPORATION—DURING THE GROWING SEASON Above the Vegetation Prairie Forest In sun, maximum heated stratum. Above crown, in sun, maximum heated Cooler above and below this stratum. stratum, A thin layer. Cooler above Absolute humidity less than in or over and below this stratum. forest. Absolute humidity greater than.in the open. Among the Vegetation Prairie Forest Temperature lower and higher than in Temperature moderated—not as low or the forest—more extreme. as high as on the prairie. Temperature lower toward the soil, and Temperature lower toward the soil, and warmer than in the forest. cooler than in the open. Absolute humidity progressively increases Absolute humidity progressively increases toward the soil. toward the soil. Relative evaporation decreases toward the Relative evaporation decreases toward the soil; greater than in the forest. soil; less than in the open. In the Soil Prairie Forest Temperature averaging warmer than Temperature cooler on the average and forest, warmer near surface in sum- in summer, and warmer in winter, near mer, and cooler in winter. Warmer in the surface, than in the open. Cooler sun and cooler at night than in forest. in sun and warmer at night than in tha open. Temperature progressively more stable Temperature progressively more stable downward. Soil moisture increases downward. Soil moisture, below the downward. surface layer, increases downward. The conditions on the prairie and in the forest may be graphically shown as in the following diagrams, Figure 14 showing the tempera- ture relations, and Figure 15 showing the relative evaporating power of the. air. 96 Jovsuns qos “norejedoa Aq pootonpur se yso10y pus oturerd Fo suOTyE[et eanyerodue, SuLMoys meiseiq “Pl “PLL SNOILV 194 FANLVAIIW IL YAINWNS NI I'SNONGIAT Wwved JuNnlLvaesdWwIL WH#OsINA 40 YFIAVT ae A JUNLvUIINILY TIGVIYTA JO YFAVT AGNUMS FeiVad Fdvad NIQHVW 189404 Sy nV oe ‘ , - - i” INOW 4F1009 ean - a 187404 XVWITD 97 “aoryejeSoa Jsor0y pue orirerd Aq peouongur se 17e jo somod Zuryerodese oatye[or Surmoys weiseiq “CT ‘SL NIDYVW FlulV dd L$3Y04 XVWITI yav79 1837401 HVO j;coj3404 dWVO FOvVIUNS JIVIUNS JOvIUNS 20VIUNs * Fowsyns ZOE o/GS. 7S ZGE)HiZ SSAA \ “W6 / i ¢ { 4) ‘409 ; LAVIN (\Ce? ter wel (| i : 709 ie NNN lies ee ia Y Ve | 187404 FAOGV %+001 98 10. Sources and Réle of Water used by Prairie and Forest Animals The bodies of animals contain a very large proportion of water— from 60 to 95 per cent. Growing animals in particular require water in relatively large amounts. Practically all foods gain entrance into the body in aqueous solutions, and are transported by water to all parts; and by the same means, the waste products, with the exception of the excretion of carbonic acid, are removed. The methods by which aquatic animals secure water are relatively simple, because they live in a liquid medium; but the conditions upon land are quite different. Here osmotic pressure does not operate as in water, and the air varies from saturation to a very dry condition. This dryness tends to cause strong evaporation from animals living in such a medium, and a proper bal- ance between intake and water-loss is one of the most potent influences in the life of land animals. In this relation lies the importance of the sources of water available to them. These sources are as follows: with the food, by drinking, from the atmosphere, and by metabolism. The loss is by excretion and evaporation, the relative humidity and the evaporating power of the air being, therefore, important considera- tions. The loss of water is retarded in many ways. Some animals possess a relatively impermeable skin, or a covering, as hair or feath- ers, which retards air currents and evaporation through the skin, just as a cover of vegetation retards soil evaporation. Other animals con- serve their moisture by modes of behavior, being active mainly during the cooler night, thus escaping the excessive evaporation of the heated day; and still others live in burrows in the soil, where the humidity is higher than in the air. Many animals can live only where the air is humid. There is thus an almost endless series of conditions relating animals to the supply and loss of water. On account of the herbivorous food habits of so many animals a large number secure much water with the juicy vegetation eaten, and others from nectar or from the sap drawn or escaping from plants. The predaceous animals secure a large amount of water from the fluids of the animals they devour or the juices sucked from their bodies, as in the case of certain Hemiptera and some parasites. In addition to the fluids derived from plants and animals, many animals also drink’ water, some in small amounts and others in large quantities. Innu- merable observations have been made by naturalists on the drinking habits of animals, but I know of no general discussion of this subject, and particularly of none from the standpoint of the variation of their behavior in this respect in different environments. But the sources of water mentioned are not the only ones available to animals, although 99 they are the most obvious, and familiar to us. An important addi- tional source is that formed within the body of the animal by the proc- esses of respiration and dehydration; this is metabolic water. The relation of this source to others and to water-loss has recently been summarized in an important paper by Babcock (’12:87, 88, 89-90, QI, 160, 161, 171-172, 174-175, 175-176, 181). The following quo- tations from this paper will serve to give a concise statement of the general principles involved in this important process. He says (pp. 87-88): “There are, however, particular stages in the life history of both plants and animals in which metabolic water is sufficient for all purposes for considerable periods of time... .. . This is also true in the case of hibernating animals that receive no water from external sources for several months, although water is constantly lost through respiration and the various excretions. In addition many varieties of insects such as the clothes moths, the grain weevils, the dry wood bor- ers, etc., are capable of subsisting, during all stages of development, upon air-dried food materials containing less than ten per cent water ; in these cases, nearly all of the water required is metabolic. . . . Many organisms also, when deprived of free oxygen, are capable of main- taining for a short time, certain of the respiratory functions, and de- riving energy from food material and from tissues by breaking up the molecular structure into new forms of a lower order. This is known as intramolecular respiration, and like direct respiration, results in the production of both water and carbon dioxide.” (Pp. 89-90): “The substances oxidized by both plants and animals, to supply vital energy, consist of carbohydrates, fats, and proteins. All of these substances contain hydrogen, and their complete oxidation produces a quantity of water equal to nine times the weight of hydrogen present in the orig- inal substances. . . . Most of the the fats yield more than their weight of water, while proteins, when completely oxidized, give from 60 to 65 per cent of water. . . . . Animals, however, are unable to utilize the final products of protein metabolism which are in most cases poisonous and must be removed from the tissues by excretion in vari- ous forms, the principal of which are urea, uric acid, and am- monia. . . . . The amount of metabolic water formed by oxidation during any period is proportional to the rate of respiration... ..... (Page 91): “With parasitic plants, and with animals, which derive all of their organic nutrients from chlorophyl producing plants, im- bibed water is not so essential to life; with these the chief function of imbibed water is to aid in the removal of waste products, the metabolic water being in most cases sufficient for transferring nutri- ents and for replacing the ordinary losses incurred by respiration and evaporation.” . . . . . (Page 160): “Another and more im- 100 portant difference is the inability of animals to resynthesize the or- ganic waste products of respiration into substances that may be again utilized as nutrients. . . . . This is especially the case with the soluble products arising from protein metabolism. With most animals these nitrogenous products are excreted in solution through the kid- neys, chiefly as urea, but birds, reptiles, and all insects excrete most of the nitrogenous waste matter as uric acid, or its ammonia salt, which being practically insoluble in the body fluids, is voided in a solid con- dition.” (Page 61): “The need for water is much less for ani- mals that excrete uric acid than for those that excrete urea, since uric acid, being practically insoluble in the the body fluids, is not so poisonous as urea and is voided solid with a minmum loss of water. Many animals that excrete uric acid instead of urea never have access to water and subsist in every stage of their development upon air dried food which usually contains less than 10 per cent water. The most striking illustrations of this kind are found among insects such. as the clothes moths, the grain weevils, the dry wood borers, the bee moths, etc. The larve of these insects contain a high per cent of water, and the mature forms, in spite of the development of wings which are rela- tively dry, rarely contain less than 50 per cent of water.” (Pp. 171- 172): “Serpents and other reptiles that live in arid regions and rarely if ever have access to water, except that contained in their food, are said by Vauquelin to excrete all of the waste nitrogen as salts of uric acid. The same is true of birds that live on desert islands where only salt water is available. It is essential that animals of these types should produce as much metabolic water as possible from the assimilated food, and the waste of water through the excretions should be reduced to a minimum. Since the food is largely protein both of these ends are at- tained by the excretion of uric acid which, as already stated, contains the least hydrogen of any nitrogenous substance excreted by animals so that the maximum amount of metabolic water has been derived from the food consumed.” (Pp. 174-175): “There are many animals that are able to go long periods without having access to water except that contained in their food, iri which water usually amounts to less than 20 per cent of total weight, and the metabolic water derived from oxi- dation of organic nutrients. A notable example of this is the prairie dog which thrives in semi-arid regions. ‘These small animals feed upon the native herbage which for months at a time is as dry as hay. It has been surmised that the burrows in which they live extend to underground water courses, but this does not seem likely since in many of these regions wells must be sunk hundreds of feet before water is reached. It is more probable that they depend chiefly upon metabolic water. They feed mostly at night when the temperature is low and a 101 during the hottest hours of day remain in their burrows where the air is more nearly saturated with moisture and evaporation is relatively small.” (Pp. 175-176): “An application of these principles would undoubtedly serve to prolong life, when suitable water for drinking is not available. In such cases the food should consist of carbohy-: drates and fats. Proteins should not be used... . . The water re- quired for preventing uremic poisoning under these conditions is small and if the relative humidity of the surrounding air is high enough to prevent rapid evaporation of water from the body, the metabolic water arising from the oxidation of nutrients may be ample for the purpose.” (Page 181): “Metabolic water derived from the oxidation of organic nutrients would probably be sufficient for all animal needs were it not for the elimination of poisonous substances resulting from protein de- generation.” | The preceding quotation brings out very clearly the harmful effects of an accumulation of uric acid upon the animal. This is only a special case illustrating a general law, for except water the main end products of metabolism are acid. There is thus a constant tendency for acid to accumulate, as Henderson (’13a: 158-159; see also ’13b) has said: “This tendency toward acidity of reaction and the accumulation of acid in the body is one of the inevitable characteristics of metabolism; the constant resistance of the organism one of the fundamental regulatory processes. Now it comes about through the carbonate equilbrium that the stronger acids, as soon as they are formed, and wherever they are formed, normally find an ample supply of bicarbonates at their dis- posal, and accordingly react as follows . . . . The free carbonic acid then passes out through the lungs, and the salt is excreted in the urine.” Recently Shelford (’13b, see also ’14a) has summarized the phys- iological effects of water-loss by evaporation and other methods. It is probable that the carbonic acid excretion is retarded by drying, and that by this means irritability may be increased. It is not simply loss of water, but loss beyond certain limits that interferes with the life of animals. Thus loss is not an unmixed evil, because, in addition to removing excretions, evaporation is an impor- tant factor in the control of temperature within the bodies of animals. Loss of water also tends to concentrate the body fluids, and when this loss brings about a relatively dry condition, such tissues are in a con- dition which is favorable for the endurance of relatively extreme low or high temperature (Davenport, 97: 256-2 58), and even dryness (see references, Adams, ’13: 98-99). This is a reason why it is dif- ficult to distinguish, in nature, between the effects of aridity and tem- perature extremes, and hence arise the puzzling interpretations of con- 102 tinental climates. These extreme conditions are characteristic of many habitats. It is readily seen how the general principles just summarized apply to the land animals of the prairie. Many of these are active during the day, live in the bare exposed places, or near the level of the vegetation, where evaporation is greatest and water-loss is correspondingly large, and feed upon the dry haylike vegetation. Others remain among the humid layers of the vegetation or in the moist soil, and feed upon juicy plants and other moist food. Predaceous and parasitic animals, deriving their moisture from their prey, occupy both the dry and humid situations. These are representative cases, between which there are a large number of intergradations. In the forest, where evaporation is more retarded than in the open, a large number of animals live in the forest crown, at the forest mar- gin, in glades, and in wood, of all degrees of dryness, and eat food varying similarly from juicy leaves to dry wood. On the other hand, some live in moist logs, among damp humus, or in the soil, and feed upon dripping fungi or soggy wood. Many of these animals possess little resistance to drying. The optimum for prairie and forest animals thus involves a dynamic balance between the intake of water and its loss by evapora- tion and excretion. ANIMAL ASSOCIATIONS OF THE PRAIRIE, AND THE FOREST © I. INTRODUCTION In an earlier chapter of this paper the habitats and animals found at the different stations were discussed, and in the preceding section the general characteristics of the physical and vegetational environ- ment of the prairie and forest have been described and summarized. We are now in a better position to consider the relations of the inverte- brates, not only to their physical environment, but also to the vege- tation, and, furthermore the relations which these animals bear to one another. We wish also to consider both the prairie and the forest as separate units, and to see how the animals are related to their physical and biological environment. As previously stated, the special locali- ties studied were described by stations both to give a precise and con- crete idea of the prairie and its animals, as now existing in a limited area, and also to preserve as much of the local color as the data would permit. I wish now to reexamine these animals from another stand- point, that of the animal association as a unit. The prairie as a whole 103 is not homogeneous from this point of view; it is a mosaic composed of anumber of minor social communities. Each of these smaller units, however, is fairly homogeneous throughout. Our present knowledge of these minor associations is imperfect, and for this reason they are arranged in an order approximating that which we might reasonably expect to be produced if the initial stage were made to begin with a poorly or imperfectly drained area and to advance progressively with corresponding vegetational changes, toward a more perfect condition of drainage. Upon the prairie a perfect series would include every stage from lakes, ponds, and swamps to well- drained dry prairie. But cultivation and drainage have obliterated so much, that now only very imperfect remnants exist in the vicinity of Charleston. Although the sequence followed therefore does not in- clude all stages of the process it is approximately genetic. There are three essential features in every animal association, or community; certain physical conditions; certain kinds of vegetation, which also modify the physical conditions; and representative kinds of animals. Occasionally an effort is made to divorce these, to separate organisms from their normal habitat, but such an effort is deceptive, for no organism can live for any considerable period without a normal environment. I have not attempted to treat these associations with equal fullness. In the sections devoted to the description of the stations it was possi- ble in some cases, on account of the uniform character of a station, to describe the animal association rather fully. In such instances the detailed account is not repeated. In other cases I have elaborated the community relations more fully here than elsewhere. The descriptions of the stations and the associations, and the annotated lists, are in- tended to be mutually supplementary. II. THe Prairie ASSOCIATIONS 1. Swamp Prairie Association The swamp prairie community lives in a habitat characterized by shallow water, which stands approximately throughout the growing season of the vegetation. The soil is black, and rich in vegetable de- bris. The characteristic plants are bulrush (Scirpus), flags (Iris), swamp milkweed (Asclepias incarnata), beggar-ticks (Bidens), and young growths both of willow (Salix) and cottonwood (Populus del- toides). The abundant growth of vegetation and the wet soil are con- ditions favorable for the production and accumulation of organic de- bris, which tends to fill the depressions and to supplement the inwash 104 from the surrounding slopes. At the same time, burrowing animals, particularly the crawfish, also bury debris and work over the soil. In the Charleston area this community was developed at Station I, d, and in part at I, g. The representative animals of this community are those living in the water, such as the prairie crawfish, Cambarus gracilis (PI. XXXVI), the snail Galba umbilicata, and such insects as the nine- spot dragon-fly, Libellula pulchella (Pl. XXXVIII, fig. 2), and the giant mosquito, Psorophora ciliata, whose immature stages are spent in the water. In addition to these are other representative species whose presence is, to an important degree, conditioned by the pres- ence of certain kinds of vegetation—such species, for example, as those which feed upon the dogbane (Apocynum), the brilliantly col- ored beetle Chrysochus auratus; upon milkweed, the milkweed bugs Lygeus kalmit and Oncopeltus fasciatus (Pl. XL, figs. 1 and 3), and the milkweed beetle Tetraopes,; and, finally, the rather varied series of flower visitors feeding upon pollen or nectar, such as the soldier-beetle (Chauliognathus pennsylvanicus), Euphoria sepulchralis, and several species of butterflies, moths, bees and wasps, including the honey-bee, bumblebees, and carpenter-bee (Xylocopa virginica), and the common rusty digger-wasp (Chlorionichneumoneum). Visiting the same flow- ers, but of predaceous habit, were found the ambush spider (Misumena aleatoria) and the ambush bug (Phymata fasciata). Small insects were preyed upon by the dragon-flies (Libellula pulchella), and the ‘dragon-flies in turn were entangled in the webs of the garden spider (Argiope aurantia). No animals were taken on the flags, but Needham (’00) has made an important study of the population inhabitating flags at Lake Forest, Illinois, and shows that it is an extensive one. He gives an excellent example showing how the injury by one insect paves the way for a train or succession of others. For example: the ortalid fly Chetopsis enea Wied. (Pl. XVIII, fig. 1), bores into the stem of the buds and causes them to decay (Cf. Forbes, ’05, p. 164; Walton, Ent. News, Vol. 19, p. 298. 1908). This condition affords a favorable habitat for a pomace-fly (Drosophila phalerata Meig.*), an oscinid (Oscinis coxendix Fitch, Plate XVIII, figures 3 and 4), a beetle, parasitic Hymenoptera, and, after the decaying buds were overgrown by fungus threads, the bibionid fly Scatopse pulicaria Loew. This paper by Need- ham is one of the very few in which the population of a plant has been studied as a biotic community. Forbes (’90, pp. 68-69; 02, p. 444) has shown that snout-beetles (Sphenophorus ochreus Lec., Plate *Mr. J. BR. Malloch infornts me that D. phalerata is not an American species. 105 XVIII, figures 5, 6, and 7) breed in root-bulbs of Scirpus, and that these beetles eat the leaves of Phragmites. Webster (’90, pp 52-55) observed these beetles feeding on the leaves of Scirpus and the larve feeding on its roots. I have found great numbers of these beetles cast up.on the beach of Lake Michigan. Evidently they breed in the swamps about the lake, fall into it when on the wing, and are washed ashore. 2. The Cottonwood Community Ordinarily we are accustomed to think of the prairie as treeless, and yet one large tree was relatively abundant upon the original prairie of Illinois, particularly upon wet prairie, or, when pools were present, even upon the uplands. This was the cottonwood, Populus deltoides. These trees were often important landmarks when isolated; and today the large trees or their stumps are important guides in determining the former extent of the prairie. In the region studied there were no large mature cottonwoods, although saplings were present, but north of Charleston in the adjacent fields mature trees were found. They grow normally at the margins of wet places, as about prairie ponds and swamps, or along the small ill-defined moist sags and small prairie brooks. This tree is usually solitary or in irregular scattered rows when along streams, and does not, as a rule, form clumps. or’ groves. This relatively isolated habit may be a factor in the comparatively small number of invertebrates which are associated with it, or at least in the amount of serious injury which they do to these trees upon the prairie. Many of the larger trees are mutilated, or even destroyed by lightning (Cf. Plummer, ’12), and such injury favors entrance of in- sects on account of the rupturing of the thick bark. The galls on the leaves and twigs of the trees often attract atten- tion. A large irregular gall on the ends of the twigs becomes conspic- uous in winter. This is formed by the vagabond gall-louse, Pemphigus oestlundi Ckll. (Pl. XIX, fig. 1) (vagabundus Walsh, Ent. News, Vol. 17, p. 34. 1906). I have found these galls abundant upon the prairie at Bloomington, Ill. At this same locality I found a large bullet-like gall at the junction of the petiole and the leaf—that of Pem- phigus populicaulis Fitch (Pl. XIX, fig. 2), and at Urbana, Ill, on other large prairie cottonwoods, a somewhat similar gall, on the side of the petioles, caused by P. populi-transversus Riley (Pl. XIX, fig. 3). I have also taken large caterpillars of the genus Apatela on leaves of cottonwood, and September 3, at Urbana, upon its cultivated form, the Carolina poplar, 4. populi Riley (Pl. XX, fig. 6). These caterpillars have bodies covered by yellow hair penciled with black. At dusk swarms of May-beetles (Lachnosterna) can be seen and heard feeding 106 among the leaves of the cottonwood and the Carolina poplar. It is noteworthy that I have made these observations at Urbana, Illinois, upon cottonwoods growing upon what was originally prairie. Forbes (’07a) has shown, as the result of extensive collections of May-beetles from trees, that they have a decided preference for Caro- lina poplar (p. 456) and willow. This same paper also contains im- portant observations on the nocturnal flights to and from the forest, from the normal habitat of the grubs, and from the daytime abode of the beetles in the open fields. Wolcott (’14) has recently emphasized the point that the grubs live only in open places in proximity to wood- land where the beetles can secure food. These observations show very clearly that May-beetles are animals primarily of the prairie or forest margin, and probably lived upon the original prairie, scattered, where cottonwoods or willows grew. A glance at the map of the prairie and forest (frontispiece) shows that the marginal area was very extensive, and must have furnished an optimum habitat for these beetles. This is a good illustration of the fact that the cottonwood exerted an influence upon the prairie far beyond its shadow. In some localities another beetle (Melasoma scripta Fabr.) feeds upon the leaves of the cottonwood, and may become a serious pest to poplars and willows, but I have not seen this species abundant on iso- lated mature trees upon the prairie. I have taken these beetles (July 2) under cottonwoods at Bloomington, Il. Packard (’90, pp. 426- 474) has published a list of the insects known to feed upon Populus. Willows (Salix) are frequently associated with the cottonwoods upon the prairie, but, in marked contrast with these, they generally grow in colonies and are eaten by a great variety of insects. Packard (790, pp. 557-600) lists 186 species of insects on them, and Chitten- den (’04, p. 63) extends the number to 380 species. Of course in any given locality the number of species found will be relatively small, and the number is further limited by the environmental conditions— whether the land is upland or low and flooded. The degree of ‘prox- imity of willows and cottonwood is likely to influence the relative abundance of the insects feeding upon these trees, since a large number of insects which feed upon willow also feed upon the cottonwood. Col- onies of willow are thus likely to become sources of infestation for the cottonwood; this relation, however, is a mutual one. Walsh (’64) and Heindel (’05) have published very interesting studies of the com- munity life of the insect galls on Illinois willows. Cockerell (’97, pp. 770-771) has listed the scale insects found upon willows and poplars. 107 3. Swamp-grass Association The prairie swamp-grasses, slough grass (Spartina), and wild rye (Elymus) were growing in relatively pure stands or colonies in de- pressions which were dry in the late summer. The prolonged wetness of the habitat and the dominance of the few kinds of grasses are char- acteristic features of the environment of this association. These con- ditions were found at Station I, a and c, north of Charleston. As these stations were rather homogeneous and have already been discussed somewhat fully, only a summary will be given here. On account of the grassy vegetation the abundance of Orthoptera is not surprising. Representative species are Melanoplus differen- tialis, M. femur-rubrum, Scudderia texensis, Orchelimum vulgare, Xiphidium strictum, CEcanthus nigricornis, and GE. quadripunctatus. Other representative animals are Argiope aurantia and the swamp fly Tetanocera plumosa. The list of species is probably very incomplete ; during the wet season there are undoubtedly a number of aquatics; furthermore, there are still other species which feed upon Spartina and Elymus, particularly some Hemiptera, and stem-inhabiting Hymenop- tera, and certain Diptera. Thus Webster (’03a, pp. 10-13, 26, 32, 38) has recorded a number of chalcids of the genus Jsosoma which live in the stems of Elymus virginicus and canadensis. In this same paper he discusses their parasitic and predaceous enemies (pp. 22, 27, 33). A fly also breeds in Elymus, the greater wheat stem-maggot, Mer- omyza americana Fitch (Pl. XX, figs. 1-5), as recorded by Fletcher (1. c., p. 48). This species is of economic importance, having spread from grasses to the cultivated grains. It has been studied in Illinois by Forbes (’84). He found a fly parasite of this species, and Webster reports a mite preying on it. Webster (1. c., p. 53) reports another fly, Oscinis carbonaria Loew, bred from Elymus by Fletcher. In another paper Webster (’03b) has published a list of insects in- habiting the stems of E. canadensis and virginicus. Osborn and Ball (‘97b, pp. 619, 622; 97a) have discussed the life histories of certain grass-feeding Jassid@ which feed upon Elymus. Osborn (’92, p. 129) records a plant-louse, Myzocallis, from Elymus canadensis in Iowa, and a species of leaf-hopper has been recorded by Osborn and Ball (’97b, p. 615') from Spartina. On the same plant, Osborn and Sirrine (’94, p. 897) record a plant-louse on the roots. In a list of the plant- lice of the world and their food plants Patch (12) lists a few from Spartina. This same list includes (pp. 191-206) many grasses an the associated aphids, those on Elymus on page 196. : 108 4. Low Prairie Association The moist black soil prairie, a degree removed from the wet or swamp condition, with ground water in the spring relatively near the surface, is fairly well characterized by the rosin-weed (Silphium), par- ticularly S. terebinthinaceum. Other plants likely to be associated with S. terebinthinaceum are Silphium laciniatum and S. integrifolium, Eryngium yuccifolium, Lepachys pinnata, and, to a less degree, Lac- tuca canadensis. In the Charleston area this condition is represented by Station I, a, north of the town, and Station III, a, and in part b, east of the town. The proximity of ground water is shown at Station I, e, by the pres-, ence of crawfish burrows, probably those of Cambarus gracilis. At Station III the proximity of water was also evident where S. terebin- thinaceum was most abundant in the railway ditches. Such perennial plants are indicative of the physical conditions for a period of years, and are thus a fairly reliable index of average conditions—much more so than the annuals. It is difficult to decide which kinds of animals are characteristic of this kind of prairie. Provisionally I am inclined to consider the fol- lowing as being so: Cambarus gracilis; Argiope aurantia; the grass- hoppers Encoptolophus sordidus, Melanoplus differentialis, M. femur- rubrum, Scudderia texensis, and Xiphidium strictum; Gicanthus nigri- cornis; Phymata fasciata; and asilids. The presence of Lepachys was clearly an important factor in determining the presence of Melissodes obliqua and Epeolus concolor. At Station III, b, east of Charleston, Epicauta pennsylvanica and Bombus pennsylvanicus, auricomus, and impatiens were taken on the flowers of Silphinm terebinthinaceum. Robertson (’94, pp. 463-464; ’96b, pp 176-177) has published lists of insect visitors to the flowers of Silphium and Lepachys (’94, pp. 468-469), at Carlinville, Ill. Recently Shelford (13a, p. 298) has published a long list of animals inhabiting Silphium prairie near Chi- cago. Forbes (’g0, p. 75) has reported the snout-beetle Rhynchites hirtus Fabr. as feeding upon Silphium integrifolium. In a colony of prairie vegetation at Seymour, IIl., which included much Silphium and Eryngium, the following insects were taken Octo- ber 7 from the ball-like flower clusters of Eryngium yuccifolium: the bugs Lygeus kalmii, Thyanta custator Fabr., Euschistus variolarius, and Trichopepla semivittata Say (No. 539, C. C. A.), the last named in large numbers, the nymphs in several sizes as well as the adults, a fact which suggests that both may hibernate upon the prairie. Rob- ertson (’89, pp. 455-456) has summarized his collections of insects from Eryngium and on Euphorbia corollata (’96a, pp. 74-75). 109 Upon remnants of prairie vegetation growing at Urbana, Illinois, I have found several kinds of insects centered about a wild lettuce, Lactuca canadensis. Upon the upper, tender parts of this plant, the plant-louse Macrosiphum rudbecki@ Fitch, thrives late in the fall, in very large numbers. Some seasons nearly every plant is infested. The lice become so abundant upon these tender parts that the entire stem for a distance of a few inches is completely covered. They migrate upward with the growth of the stem and keep on the fresh, tender parts. Among the plant-lice, and running about on the stem of the plant, attending ants abound; eggs, larve, and adults of lace-wing flies (Chrysopa) also abound; and several species of coccinellids, syrphid larve, and a variety of small parasitic Hymenoptera are present. 5. Upland Prairie Association The well-drained prairie, a degree removed from the permanently moist prairie, is fairly well represented by the physical and biological conditions in which Euphorbia corollata, Apocynum medium, and Lactuca canadensis, are the representative plants. The plant ecologist would consider the conditions favorable to mesophytic plants. In the Charleston region these conditions are approximated at Station II, where drainage has doubtless changed the area from a somewhat moist, to its present well-drained, condition. : Representative animals of this community are as follows: Argiope aurantia, Misumena aleatoria, Encoptolophus sordidus, Melanoplus bivittatus, M. differentialis, Orchelimum vulgare, Xibhidium strictum, Euschistus variolarius, Phymata fasciata, Chauliognathus pennsylvan- icus, Epicauta marginata and E. pennsylvanica, Rhipiphorus dimidia- tus and R. limbatus, Ammalo, Exoprosopa fasciata, Promachus verte- bratus, Bombus pennsylvanicus, and Myzine sexcincta. On dry prairie at Mayview, Ill., September 26, I found the plant- louse Aphis asclepiadis Fitch on the leaves and stems of the dogbane (Apocynum) and the lice attended by the ant Formica fusca L. A beetle, Languria mozardi Latr., whose larva is a stem-borer, inhabits Lactuca canadensis. Its life history and habits have been discussed by Folsom (’o9, pp. 178-184). 6. The Solidago Community A common community in the late summer and early fall is centered about the goldenrod (Solidago). This plant was‘not abundant or in blossom at any of the stations studied in detail, but it grew in small widely scattered colonies or clumps. Observations were made in two 110 colonies, north of Charleston, both west of Station I, a, andI,g. The collections made (Nos. 20, 26, 42, 43) are as follows: Ambush Bug Phymata fasciata 20, 26 Stink-bug Euschistus variolarius 26 Black Blister-beetle Epicauta pennsylvanica 26 Noctuid moth Spragueia leo 20, 26 Conopid fly Physocephala sagittaria 26 Empidid fly Empis clausa 43 Halictid bee Halictus fasciatus 26 Myzinid wasp Myzine sexcincta 20, 26 Ant Formica fusca subsericea 20 It is important to know that these collections from Solidago were made just as the flowers were beginning to blossom. Collections a few weeks later would probably have given many more kinds. It should be noted, too, that all these plants were far out upon the prairie and far from woodlands—a factor which may influence to some extent the kinds of visitors. As a rule the lists which have been published state little or nothing at all as to the conditions in which the plants were growing. If this factor is neglected, the presence of some vis- itors remains puzzling. Thus on some goldenrods the locust beetle, Cyllene robinie, is abundant; but this is conditioned in part by the proximity of the yellow locust, which is absent on the Charleston prairie. Phymata was found copulating upon the flower, and with an em- pidid fly, Empis clausa (No. 43), in its grasp. Two kinds of galls formed by insects were found on this plant: one formed by the fly Cecidomyia solidaginis (No. 43), which forms a rosette of leaves; and the other the spindle-like stem-gall, formed by a small caterpillar, Gnorimoschema gallesolidaginis (No. 7462 Hankinson). September 20 the moth Scepsis fulvicollis Hiibn. was found in goldenrod flowers near Station I,a. Its larva feeds on grass. ) from the upland forest to the lowland Aug. 22 (No. 133); and a female was taken among leaves on the ground in the upland forest (Sta. IV, a) Aug. 23 (No. 150). T. L. Hankin- son found an adult and a nymph in the Bates woods June 28, 1911 (No. 7678). (Cf. Hancock, ’11, pp. 362-364.) The positive heliotropism or negative geotropic response shown in diseased grasshoppers is of interest. It may be caused either by a fungous or bacterial disease. (Cf. Gillette, Bull. No. 6, n. s., Div. Ent. U. S. Dept. Agr., pp. 89-93. 1896. ) Morse (’04, p. 15) considers this an exceptional ground-inhabiting or geophilous species since it is “an inhabitant of xerophytic forests as well as of open fields, and in the Southern States is found quite as often in the forest as on the open plain.” Melanoplus differentialis Thomas. Differential Grasshopper. Consult the list of prairie invertebrates, p. 167. Melanoplus atlanis Riley. Lesser Grasshopper. (Pl. LVII, fig. 8.) A single specimen was taken on the ground in the upland forest (Sta. IV, a) Aug. 16 (No. 67). The open character of parts of this dry forest affords favorable conditions for this species. Morse (’04, pp. 19, 42) considers this a characteristic species of open country, but “likely to be found anywhere.” Hancock (’11, pp. 415-416) has described the habitat of this species. 214 Melanoplus amplectens Scudd. This locust and nymphs doubtfully regarded as of the same species were taken from the ground, mainly among leaves, in the up- land forest (Sta. IV, a) Aug. 16 (No. 67); other collections are as follows: in the glade in the lowland forest (Sta. IV, c) Aug. 20 (No. 117); on the open ravine slope (Sta. IV, b) Aug. 22, a male (No. 124a) ; and on the same date, in the glade of the lowland forest (Sta. IV, c), a nymph and an adult female (No. 143). This is the largest of the short-winged locusts in the forest, and an abundant species. Morse (’04, pp. 19, 50, Pl. 7) described its haunts as in thickets, forest margins, open forests, and occasionally in grassy clearings and fields. Melanoplus gracilis Bruner. Two males were found Aug. 20 in a glade in the lowland forest (Sta. IV,c) where there was a luxuriant cover of vegetation, and nettles and Eupatorium calestinum abounded; and Aug. 22, in the same location, one female was found (No. 143). The wings are very rudimentary in this species. Hart (’06, p. 82) describes its habitat as follows: “On tall grasses and weeds in ravines and about marshes, masses of wild vines along railroads, weedy growths in the beds of small streams, and in like situations.” These conditions are found in open areas with an abundance of vege- tation. Melanoplus obovatipennis Blatch. This small species, similar to scudderi, was found in the upland forest (Sta. IV,a) Aug. 17 (No. 93). A nymph taken Aug. 22 from the forest (Sta. IV) is doubtfully regarded as of this species (No. 124). Hart (06, p. 81) gives the habitat of this species as “High wooded hillsides throughout Ilinois.”” Blatchley (’03, p. 308) states that it frequents ““for the most part, high, dry, open woods, espe- cially those in which beech and oak trees predominate.” He further states that in a dry season it may be found associated with Dichro- morpha viridis and Truxalis brevicornis “among the reeds and tall rank grasses near the borders of marshes.” Melanoplus scudderi Uhl. Scudder’s Grasshopper. A single female was found in the open glade in the lowland for- est (Sta. IV,c) Aug. 20 (No. 117); and a nymph taken Aug. 22 from the open ravine slope (Sta. IV, b) is doubtfully referred to this species (No. 124). 215 Hart (’06, p. 81) describes the habitat of this grasshopper as “open woods and thickets, and along rail fences and roadsides.” Species which now characterize our open, partly cleared woodlands, in the primeval forest probably frequented forest margins, bluffs, and the borders of streams, or open patches in woods where a tree had fallen, and similar situations. With a thinning out of the for- est (up to a certain degree) their habitat is increased in area, but when by clearing the woods disappear, their habitat vanishes. LocustTIp Scudderia furcata Bruner. Forked Katydid. (Pl. LVI, fig. 5.) One female was taken in an open area in the upland forest on low shrubs (Sta. IV, a) Aug. 20 (No. 109). Another specimen was taken near Vera, Fayette county, III., on a finely developed colony of prairie vegetation among Andropogon, Sept. 1 (No. 185). Blatchley (’03, p. 349) states that it is “most frequently seen on the low bushes a trees about the margin of thickets and along fence rows, but in the prairie country north [in Indiana] it frequents coarse grasses and weeds.” Amblycorypha rotundifolia Scudd. Round-winged Katydid. (PI. LVII, fig. 2.) A single female of this species was taken in the glade in the low- land forest (Sta. IV,c) Aug. 20 (No. 117); and also a freshly emerged female (No. 143). Blatchley (’03, p. 352) states that this is “more of a terrestrial species than oblongifolia, being often seen on the ground, or on clumps of tall grass and weeds which grow in damp ravines.” Hart (’06, p. 84) says that this species is found “On grasses and weeds in damp ground.” Microcentrum laurifolium Linn. Angle-winged Katydid. (Pl. LVII, figs. 1 and 2.) Males were found on hickory sprouts at the cleared margin of the upland forest (near Sta. IV.a) Aug. 22 (No. 135). They were chirping loudly, in the early afternoon, on sprouts less than two feet high. Cyrtophyllus perspicillatus Linn. Common Katydid. (Pl. LVIII, fig. 1.) One male was taken in the partly cleared area bordering the for- est (near Sta. IV,a) Aug. 23 (No. 145). Here, among stump sprouts of hickory, oak, and young sassafras, about two to three feet high, stood this mate stridulating in the sun at 2:30 p. m., but the note did not seem exactly normal, that is, as when heard at night. 216 This species is so distinctly arboreal and nocturnal that I was sur- prised to find it stridulating during the day, and so near the ground. I have camped for days in a grove where these insects made a great din at night, but found none on the low vegetation or on the ground (as at Kappa, Ill). Years ago a large colony flourished in Franklin Park at Bloomington, Ill. Conocephalus nebrascensis Bruner. Nebraska Cone-nose. A female was taken in the glade in the damp lowland forest (Sta. IV,c) Aug. 20 (No. 117). The female of this species has been observed to oviposit “between the stem and root-leaves of Andropogon”, a typical prairie plant, but little appears to be recorded of its habitat. A large nymph of this genus, and probably of this species (No. 159), was taken on the prairie grass Andropogon (Sta. I,g) Aug. 24. It had been captured by the crab-spider Miswmena aleatoria Hentz (No. 159). Orchelimum cuticulare Redt. A specimen was taken in the upland forest (Sta. IV,a) Aug. 16 (No. 67); another, from the open areas of the upland forest (Sta. IV,a) Aug. 17 (No. 93); and a third, from the glade in the damp lowland forest (Sta. IV,c) Aug. 22 (No. 143). All of these were males. Orchelimum glaberrimum Burm. This insect was found in abundance in the glade in the lowland forest (Sta. IV,c) Aug. 20 (No. 117), and a nymph was taken in the same place Aug. 22 (No. 143). The abundance of this species in this damp area, with its pro- fusion of low vegetation, indicates that the conditions were fav- orable. Xiphidium nemorale Scudd. Nymphs and adults were found in the glade in the lowland for- est (Sta. IV, c) Aug. 20 (No. 117) and Aug. 22 (No. 143); in the openings in the upland forest (Sta. IV,a) Aug. 17 (No. 93), and Aug. 20 (No. 103). Blatchley (’03, p. 374) states that it abounds along the “borders of dry, upland woods, fence rows, and roadsides, where it delights to rest on the low shrubs, blackberry bushes, or coarse weeds usually growing in such localities.” GrYLLDZz Nemobius fasciatus DeG. Striped Cricket. (PI. LVIII, fig. 6.) Nymphs of this species were found in the upland forest on the 217 ground (Sta. IV,a) Aug. 16 (No. 67); in the upland forest area also, in an open place, was found a short-winged male Aug. 17 (No. 93); along a path in the upland forest, among dry leaves, a short- winged female was taken Aug. 22 (No, 122); and an abundance of short-winged males and females, and nymphs (No. 143) were found Aug. 22 in the glade in the lowland forest (Sta. IV, c). This small cricket is generally abundant among the litter on the forest floor. Nemobius maculatus Blatch. Spotted Cricket. A nymph was taken in the upland forest (Sta. IV,a) among leaves Aug. 22 (No. 122). Blatchley (’03, p. 425) states “It is found in low open woods, usually in the vicinity of or beneath logs”; Hart, (’06, p. 89) states that it is found “About logs and dead wood in sparse woods and near streams.” Apithus agitator Uhl. Woodland Cricket. A nymph was taken from the open area in the upland forest (Sta. IV, a) Aug. 17 (No. 93); another from an open ravine slope (Sta. IV, b) Aug. 22 (No. 124). No adults were secured. Blatchley (’03, pp. 458-459) records this species as from forests, noting its preference for prickly ash. It is also recorded as from grape-vines and dense shrubbery. The females deposit eggs in the twigs of the white elm, Ulmus americana Linn. HEMIPTERA CicapIDz Cicada linnet Grossb. (Cicada tibicen L.). Dog-day Harvest-fly. PL. LV, fig; 5.) This insect was found at the cleared margin of the upland forest (near Sta. IV, a) on low hickory sprouts Aug. 26 (No. 162). It is said to require two years to mature. T. L. Hankinson re- ports that Tibicen septendecim L, (Pl. LV, figs. 3 and and 4) was found about Charleston in 1907, and branches scarred by the oviposit- ing females were observed in the Bates forest (Sta. IV, a). Felt (’05, pp. 237-238) describes the emergence of the adult Tibicen from the nymph skin. For the recent synonymy see Smith and Grossbeck (Ent. News, Vol. 18, pp. 116-129. 1907). FuLGoRIDA Ormenis pruinosa Say (?). Mealy Flata. (Pl. LVI, figs. 1 and 2.) This insect was taken by T. L. Hankinson June 28, 1911, in the 218 Bates woods (No. 7678). It.appears to feed upon a large variety of trees, shrubs, and herbaceous plants. Its normal habitat is probably in open woods or the forest margin. Swezey (’04, pp. 8-9) gives full references to the life history of this insect and a list of the food plants. TETTIGONIELLIDA Aulacizes irrorata Fabr. (Pl. LVI, fig. 3.) A few specimens were taken, the collection data being as follows: from an open glade in the lowland forest (Sta. IV, c) Aug. 20 (No. 117); and from the smaller branches of sassafras bushes (Sta. IV, c) Aug. 22 (No. 143). This insect is often taken on grapes, and in the South on cotton. Sanderson (Bull. 57, Bur. Ent., U. S. Dept. Agr., p. 58. 1906) describes briefly the egg-laying habits and figures the adult insect. Gypona pectoralis Spangb. This species was taken June 28, 1911, in the Bates woods (Sta. IV) by T. L. Hankinson (No. 7678). PENTATOMIDA Euschistus fissilis Uhl. This bug was taken in Bates forest (Sta. IV) Aug. 22 (No. 124). It has been known to feed upon wheat (Webster, Rep. U. S. Dept. Agr., 1885, p. 317). It also feeds upon corn, and on the moth Aletia. It is parasitized by the proctotrypid Trissolcus euschisti Ashm. (Olsen, in Journ. N. Y. Ent. Soc., Vol. 20, p. 52. 1912). Mormidea lugens Fabr. A nymph of this bug was taken by T. L. Hankinson in the Bates woods (Sta. IV) June 28, 1911 (No. 7678). Hymenarcys nervosa Say. This insect was taken on the ground from among dead leaves and decayed wood which had drifted to the mouth of a ravine in the low- land forest (Sta. IV, c) Aug. 20 (No. 113). In the South this insect feeds upon cotton. Mirpz Lygus pratensis Linn. Tarnished Plant-bug. This bug was taken in the Bates woods (Sta. IV) June 28, IQII, by T. L. Hankinson (No. 7678). See prairie list, page 175. 219 CorEIDz Alydus quinquespinosus Say. This bug was taken by T. L. Hankinson June 28, 1911, in the Bates woods (No. 7678), and July ro (No. 7693) on the under- growth in the woods (Sta. IV). Acanthocerus galeator (Euthoctha) Fabr. (Pl. LVI, fig. 8.) Six large nymphs of this plant-bug were taken on the apical part of a tall herb, Actinomeris alternifolia Linn., growing in the open glade of the lowland forest (Sta. IV, c; Pl. XIV) Aug. 29 (No. 182). This bug has been reported to suck the juice from the plum, and it injures the tender parts of orange plants. Hubbard (Insects Af- fecting the Orange, U. S. Dept. Agr., Div. Ent., p. 163. 1885) gives figures of the adult insect and describes briefly the eggs and young. Forbes and Hart (’00, p. 445) have summarized the little that is known of this insect. Jalysus spinosus Say. Spined Stilt-bug. (Pl. LVI, fig. 7.) This bug was found Aug. 20 in the open glade of the lowland for- est (Sta. IV, c), where there was a luxuriant growth of herbaceous vegetation (No. 117). It was also taken (Sta. IV) by T. L. Hank- inson June 28, 1911 (No. 7678). Lugger reports it from oak woods. It feeds upon plants. GERRDz Gerris remigis Say. Water-strider. (PI. L, fig. 2.) This water-strider was abundant in the pools of the small tem- porary stream in the ravine bordering the southern part of the Bates woods (Sta. IV, d) Aug. 22 (No. 129). It is an important enemy of mosquito larvee. REDUVIDA Sinea diadema Fabr. Rapacious Soldier-bug. A nymph of this predaceous bug was captured by T. L. Hankin- son in the Bates woods (Sta. IV) June 28, 1911 (No. 7678). See list of prairie animals, page 173. CoLEOPTERA CICINDELIDA: Cicindela unipunctata Fabr. Woodland Tiger-beetle. One specimen of this tiger-beetle was taken along the path through the cleared area as it entered the forest (Sta. IV,a) Aug. 22 (No. 136). 220 Tiger-beetles are generally most abundant in open places, but this beetle seems to be a woodland species like the brilliantly colored C. sexguttata Fabr. Wickham (’99, pp. 210-211) records unipunctata from wooded areas. It is rare and difficult to catch, and is said to be nocturnal in habit. CARABIDAR Calosoma scrutator Fabr. Caterpillar-hunter. This common arboreal beetle was taken Aug. 16 (No. 64) in the upland Bates wood (Sta. IV, a), where it attracted attention by the rustling sound it made in crawling among the dry leaves on the ground. Specimens of these beetles I could easily secure by remain- ing quiet and listening for this rustling of the leaves. One specimen was seen to crawl up the trunk of a small oak-tree, three or four inches in diameter, for about seven feet. Another individual I took from my neck. It may have fallen upon me from a tree, but more prob- ably it climbed upon me as it does a tree. In woods adjacent to the Bates forest, a caterpillar-hunter (No. 97) was found Aug. 20 with what appeared to be the damp cast skin of some large bombycid larva, which was also claimed by an ant, Camponotus herculeanus Linn., subsp. pennsylvanicus DeG., var. ferrugineus Fabr. On the ravine slope (Sta. IV, b) Aug. 20 T. L. Hankinson captured one of these beetles (No. 100) with a caterpillar about an inch long, which it had partly mangled in the thoracic region with its formidable jaws. On the upper slopes of the ravine (Sta. IV, b) Aug. 23 another beetle (No. 149) was found on the ground under a hickory tree, eating a Datana larva. Along this same rather open forested slope another individual was observed to run from the ground up the trunk of a small white oak (six or seven inches in diameter) for three or four feet, and then to return to the ground. The climbing individuals ob- served took a relatively straight course up the trunk, making no ef- fort to climb in a spiral direction, and made the descent head fore- most. ‘At Bloomington, Ill, while picking cherries I have taken the beetle in trees. Although the arboreal habit is evidently very well developed in this species, it is also very much at home on the ground. The rapidity and apparent ease with which it ran over dry oak leaves in the upland Bates woods surprised me. The active foraging habits of this beetle are well shown by Her- man’s observations (Journ. Cincinnati Soc. Nat. Hist., Vol. 21, p. 80. 1910) on its killing nestlings of the cardinal grosbeak (Cardin- alis cardinalis) in bushes three feet from the ground. Harris (In- 221 sects Injurious to Vegetation, p. 470. 1869) states that it preys upon canker-worms, both on the ground and by ascending trees. Galerita janus Fabr. A specimen was found under the bark of a decaying log in the upland Bates forest (Sta. IV,a) Aug. 23 (No. 171). This common beetle is frequently found in such situations, and seems to’ have a preference for relatively damp places. I have taken the adult as early as March 23 under bark of logs in the sap-wood stage of decay at Urbana, Ill., where it was found associated with single dealated females of Camponotus herculeanus pennsylvanicus, Passalus cornu- tus, pyrochroid larve, the caterpillar Scolecocampa liburna, and the slug Philomycus carolinensis. This species is a fairly common one. I found it abundant at Bloomington, Ill., where it was taken April 15, May 1, and June 22. The larva has been described by Hubbard (Psyche, Vol. 1, pp. 49-52. 1875). CoccINELLIDA: A species of lady-beetle was found upon a fungus growing on a stump in the upland forest (Sta. IV,a) Aug. 17 (No. 81). Asso- ciated with the beetle on the fungus were large numbers of the snail Pyramidula perspectiva. ELATERIDA Melanotus sp. A larva belonging to this genus (No. 125) was found Aug. 22 under the bark of a decaying stump (Sta. IV. 6) in which the sap- wood was destroyed, the remainder being sound though discolored. It was associated with the slug Philomycus carolinensis and the caterpillar Scolecocampa liburna. Corymbites sp. A larva belonging to this genus (No. 113) was found in drifted leaves and dead wood at the mouth of a ravine in the lowland for- est (Sta. IV, c). Asaphes memnonius Hbst. This click-beetle was taken at the mouth of a ravine in the low- land forest (Sta. IV,c) Aug. 20 (No. 113) in drift composed of dead leaves and rotten wood. LAMPYRIDE Calopteron terminale Say. Black-tipped Calopteron. This interesting beetle was taken in the damp lowland forest (Sta. IV, c) Aug. 26 (No. 173). 222 This species has been mentioned as an instance of mimicry because of its resemblance in shape and color-pattern to the syntomid moth Lycomorpha pholus Drury. Both are found in damp shady woods. Calopteron reticulatum Fabr. Reticulate Calopteron. (PI. LVIII, fig. 4.) A single specimen was taken—in the glade in the lowland forest (Sta. IV,c) Aug. 22 (No. 143). The larva and pupa of this species are described by Coquillett (Can. Ent., Vol. 15, pp. 97-98. 1883). July 10 he found a pupa “suspended by the hind end of its body beneath a log.” Photuris pennsylvanica DeG. Pennsylvania Firefly. (Pl. LVIII, fig. 3.) This large firefly was taken June 28, 1911, in the Bates woods (Sta. IV) by T. L. Hankinson (No. 7678). McDermott (’10, ’11) Knab (’05), and Mast (’12) should be consulted for discussions on the natural history and ecology of our fireflies. McDermott gives many observations on P. pennsylvanica. Chauliognathus marginatus Fabr. Margined Soldier-beetle. This predaceous beetle was taken June 28, 1911, in the Bates woods (Sta. IV) by T. L. Hankinson (No. 7678). (Cf. Lintner, Fourth Rep. Injurious and other Ins. N. Y., 1888, pp. 74-88.) This is a predaceous species in the larval stage, feeding on immature in- sects. The adults feed on pollen (Riley, in Fifth Rep. Ins. Mo., p. 154. 1873). Telephorus sp. This was taken June 28, 1911, in the Bates woods (Sta. IV) by T. L. Hankinson (No. 7678). See T. bilineatus, Pl. XLIV, fig. 1. LucanDa Passalus cornutus Fabr. Horned Passalus. (Pl. LVIII, fig. 5.) This common woodland beetle was found under the bark of a decaying stump on the slope of a ravine (Sta. IV,b) Aug. 17 (No. 85). One specimen, with a chestnut thorax and yellowish wings, had just shed the pupal skin. Another, a fully matured specimen, carried a large colony of mites. Ewing (Univ. Studies, Univ. IIL, Vol. 3, p. 24. 1909) states that nymphs of uropod mites are often attached to insects for transportation. It has generally been as- sumed that they are parasitic. This Passalus seems to be one of the most common insects found in decaying logs and stumps. I have found it very abundant at 223 Bloomington, Ill. The beetles evidently hibernate, for I have taken them at Urbana, IIl., as late as October 18, and as early in the spring as March 23. This beetle invades logs and stumps as soon as the sap-wood be- gins to be well decayed, and evidently advances into the log with the progress of decay. As it invades logs in the sap-wood stage of decay, it is often associated with newly founded colonies of the ant Cam- ponotus herculeanus pennsylvanicus, pyrochroid larve, the slug Phil- omycus carolinensis, and the caterpillar Scolecocampa liburna. For physiological studies of cornutus see Schafer (Mich. Agr. Coll. Exper. Sta., Tech. Bull. No. 11. 1911). ScaRABAIDS Geotrupes splendidus Fabr. Splendid Dung-beetle. This dung-beetle was dug from a hole, an inch or so below the surface, in the hard clay of the pathway near the margin of the for- est bordering the cleared area (Sta. IV, a) Aug. 22 (No. 120). As cattle and horses were pastured in this forest, its presence is readily accounted for. Pelidnota punctata Linn. Spotted Grape Beetle. Only one specimen of this beetle was taken. It was found on a grape leaf (Sta. III, b) Aug. 15 (No. 58). This insect is primarily a forest or forest-margin insect. The larva feeds upon the decaying roots and stumps of oak and hickory. The adult devours leaves of the grape and of the Virginia creeper. Many undetermined scarabeid larve were found in a much-de- cayed stump in the ravine near the small temporary stream (near Sta: IV, d) Aug. 22 (No. 130). CHRYSOMELIDZ Chrysochus auratus Fabr. Dogbane Beetle. This characteristic species of the prairie (No. 103) was taken Aug. 20 in an open place in the upland oak-hickory forest (Sta. IV, a) on the dogbane Apocynum medium. See list of prairie inver- tebrates, p. 178. Cryptocephalus mutabilis Mels. This leaf-beetle was taken June 28, 1911, in the Bates woods (Sta. IV) by T. L. Hankinson (No. 7678). It has been reported on Ceanothus, Viburnum, hazel, and oak by J. B. Smith. Evidently this is a woodland beetle. 224 Coptocycla clavata Fabr. Clubbed Tortoise-beetle. This leaf-beetle was taken in the south ravine of the Bates woods (Sta. IV. b) by T. L. Hankinson June 28, 1911 (No. 7678). It is known to injure the potato, tomato, eggplant, and bittersweet. The larve and adults feed upon the same kinds of plants (Lintner, Sixth Rep. Injurious and other Ins. N. Y., pp. 126-127. 1890). TENEBRIONIDAS Boletotherus bifurcus Fabr. Horned Fungus-beetle. (Pl. LIX, figs. I, 2, and 3.) This curious-looking beetle was found on the shelf-fungus Polyp- orus in the lowland forest (Sta. IV, c) Aug. 26 (No. 173). I have found this species very abundant near Bloomington, TIL, where at times it was difficult to find an example of Polyporus which was not thoroughly honeycombed by the larvz of these beetles. A single shelf has been found to contain several beetles. They were generally discovered within galleries excavated within the fungus. On July 11 in such a shelf I found larve and pupe in abundance. Other dates of capture are June 3 and July 6. Riley and Howard (In- sect Life, Vol. 3, p. 335. 1891) also report it from Polyporus. Fig- ures of the larva and pupa are given by Packard (’83, p. 474) and descriptions by Gissler (On coleopterous larve of the family Tene- brionide, Bull. Brooklyn Ent. Soc., Vol. 1, pp. 85-88. 1878). Meracantha contracta Beauv. Larve of this beetle were taken under dry leaves in the upland forest (Sta. IV,a) Aug. 17 (No. 83); and others from under damp leaves at the base of the wooded slopes of a ravine leading to the low- land forest (Sta. IV, b) Aug. 22 (No. 140). The latter larvee were associated with the ant. Siigmatomma pallipes. These larve are often confused with wireworms (Elateridae). ‘ I found the beetles occasionally in the forest at Bloomington, IIL, June 13; and Aug. 1 on the papaw. I have a specimen of this larva, in very rotten wood, showing the sinuous larval boring (Pl. XXX), from the Brownfield woods, Urbana, Ill. (March g; collector, D. M. Brumfiel). Wickham has described and figured the larva (Journ. N. Y. Ent. Soc., Vol. 4, pp. TIQ—121. 1896). PyYrocHROoDa Pyrochroa sp. A single specimen of a larva belonging to the above family was taken August 22 (No. 130) in the ravine (Sta. IV, b) from under 225 the bark of a decaying stump, in company with numerous scarabzid larve. These larve are very characteristic animals—under bark when decay has loosened it from the sap-wood. The accompanying figure (Pl. LIX, fig. 4) shows the general appearance of this larva and of an adult beetle. I found Dendroides canadensis Latr. fairly abundant at Bloomington, Ill, July 25. Larve belonging to this family have been taken in the Brownfield woods, Urbana, Ill., under the bark of decaying trees. It is a representative animal species in this habitat. See Moody (Psyche, Vol. 3, p. 76. 1880) for descriptions of pyrochroid larve. LEPIDOPTERA PaPILIONIDZ Papilio philenor Linn. Philenor Butterfly. (Pl. LIX, fig. 5.) The caterpillar was found crawling upon the ground in the up- land forest (Sta. IV,a) Aug. 16 (No. 69). Aug. 26 a larva (No. 166) which had attached itself to the stem of a prickly ash (Sta. IV, b), was just entering upon the pupal stage, but had not yet cast the larval skin. The larva feeds upon Dutchman’s pipe, Aristolochia—a plant which was not observed in the forest. Fapilio turnus Linn. Turnus Butterfly. The butterfly was observed on wing Aug. 16 in the open glades of the upland forest (Sta. IV, a). The larva feeds upon Prunus and Liriodendron. Papilio cresphontes Cram. Cresphontes Butterfly. The butterfly was observed in the open spaces of the upland forest on wing Aug. 16. The larva feeds upon Zanthoxylum, Ptelea, Dictamnus, Citrus, etc. Papilio troilus Linn. Troilus Butterfly. The butterfly was taken, on wing, from the open slope of the south ravine (Sta. IV,b) Aug. 22 (No. 161); and in the upland forest (Sta. IV,a) Aug. 26 (No. 163). The larva feeds upon sassafras and Laurus. NyYMPHALIDZ Polygonia interrogationis Fabr. The butterfly was taken in the open glade in the lowland forest (Sta. IV, c) Aug. 20 (No. 117). The larva feeds upon Humulus, Ulmus, and Urtica. 226 AGAPETIDA Enodia portlandia Fabr. Portlandia Butterfly. This woodland butterfly was taken in the Bates woods (Sta. IV) Aug. 15 (No. 63) and on June 28, 1911 (No. 7678), by T. L. Han- kinson. The larva feeds upon grasses. Fiske (’01, pp. 33-34) gives a good description of the haunts of this species. Years ago I found it abundant near Bloomington (Orendorf Springs), Ill., in dense, damp, shady woods. It is as characteristic of shade as most species are of sunshine. Cissia eurytus Fabr. Eurytus Butterfly. This is also a woodland butterfly. It was taken in the Bates woods by T. L. Hankinson June 28, 1911 (No. 7678). The larva feeds upon grass. LyYca&NIDz Everes comyntas Gdt. This small blue butterfly was taken on the open upper slopes of the wooded south ravine in the Bates forest (Sta. IV, 5) Aug. 22 (No. 161). The larva feeds upon red clover and Desmodium. HESPERIIDA * Epargyreus tityrus Fabr. Common Skipper. This caterpillar was found in the open. glade in the lowland for- est (Sta. IV,c), folded within a leaf of sassafras, Aug. 26 (No. 173). I have taken this butterfly many times at Bloomington, IIl.; and have found the larve folded in leaves of the yellow locust, Robinia, upon which they had evidently been feeding. SPHINGIDZ Cressonia juglandis Sm. and Abb. This caterpillar was taken on low branches of the shell-bark hick- Ban Carya ovata, in the upland forest (Sta. IV,a) Aug. 20 (No. 102). The larva feeds upon walnut, ironwood, and hickory. Our speci- men bore a large number of cocoons of a hymenopterous parasite. When handled, this larva makes a peculiar squeaking sound (Bull. 54, Bur. Ent., U. S. Dept. Agr., p. 80. 1905). 227 SaTURNIIDA Telea polyphemus Cramer. American Silkworm. (PI. LIX, fig. 6.) This caterpillar was taken on the ground, under hickories and white oaks on the forested slopes to the valley (Sta. IV, b) Aug. 26 (No. 163). The larva feeds upon walnut, basswood, elm, maple, cherry, etc. CERATOCAMPIDA Citheronia regalis Fabr. Royal Walnut Moth; Hickory Horned-devil (larval name). (PI. LX, figs. 1 and 2.) This larva was found on the valley slope (Sta. IV, b) on sumac Aug. 16 (No. 68); and on walnut Aug. 20 (No. 108). This last specimen was apparently fully mature. The food plants of the larva are butternut, hickory, sycamore, ash, and lilac. See Packard (’05, p. 130) for many figures and a full description of this species. Bastlona imperialis Drury. Imperial Moth. (Pl. LXI, Fig. 1). The larva of this species was found on the leaves of sassafras on the forested slope to the lowland forest (Sta. IV, b) Aug. 20 (No. 106). It feeds upon a large number of forest trees including oak, maple, wild cherry, walnut, hickory, and several conifers. See Packard (’05, p. 125) for figures and full descriptions of this species. ARCTIDA Halisidota tessellaris Sm. and Abb. (Pl. LXI, fig. 4.) These caterpillars were taken on hickory on the wooded slope to the lowland (Sta. IV, b) Aug. 26 (No. 163); and, again, abundantly (No. 168), in the upland forest (Sta. IV, a) on climbing buckwheat, Polygonum convolvulus, which was entwined about a young walnut or butternut. The yellow hairs and the tufts give this caterpillar a striking appearance. I have found moths of this species abundant at Bloomington, IIl. The food plants are recorded as maple, oak, hazel, and button- wood. Though larve were abundant upon leaves of the climbing buckwheat, I did not observe them there eating it. Noctua Autographa precationis Guen. The moth was taken in the open glade in the lowland forest (Sta. IV, c) Aug. 22 (No. 143). The larva feeds upon plantain, burdock, and dandelion. 228 Scolecocampa liburna Geyer. Rotten-log Caterpillar. A single caterpillar (No. 125) was taken Aug. 22 upon the slope of a wooded ravine (Sta. IV, b) under the bark of a stump in an early stage of decay—the sap-wood honeycombed, but the remainder solid though discolored. The larva, with its characteristic excrement, was found in a cell excavated in the rotten sap-wood. This is another species of animal which invades wood in the sap- wood stage of decay and is so often associated with Philomycus carolinensis, Passalus cornutus, and newly established colonies of Camponotus herculeanus pennsylvanicus. The larva winters in logs, as is evidenced by the fact that I found it in such situations late in fall and early in spring (March 23) at Urbana, Ill. The large quantity of excrement often indicates the approximate location of the larva. This larva has been described by Edwards and Elliot (Papilio, Vol. 3, p. 134. 1883). It has been found in chestnut, oak, and other kinds of decaying logs. The moth is recorded in July. The pileated wood- pecker, Phleotomus pileatus, has been known to eat this caterpillar (Beal, in Bull. 37, Biol. Surv., U. S. Dept. Agr., p. 34. 1911). Smith (Ann. Rep. N. Jersey State Mus., 1909, p. 471. 1910) states that the larva is found in “decaying cherry, hickory, oak and chestnut stumps.” i NoToponTIDz Datana angusti G. and R. The caterpillar of this species was found on the valley slope (Sta. IV. b) on bitternut hickory, Carya microcarpa, Aug. 20 (No. 104); in the upland forest (Sta. IV, a) on hickory Aug. 16 (No. 65); and at the margin of this forest Aug. 26 (No. 162). The food plants of the larva are walnut, hickory, linden, and birch. Packard (’95, pp. 110-111) describes and gives figures of the larva and adult. Nadata gibbosa Sm. and Abb. (Pl. LXI, fig. 2.) This larva was taken on white oak, Quercus alba, in a forested ravine (Sta. IV, b) Aug. 19 (No. 94); on leaves of the white oak, upon which it had been feeding, in the upland forest (Sta. IV, @) Aug. 26 (No. 169). Packard (’95, pp. 142-146) gives figures of this species and lists as food plants, oak, birch, and sugar plum. It is also reported on maple. Heterocampa guttivitta Walk (?). (Pl. LXI, figs. 3 and 5.) This larva (No. 127) was captured Aug. 22 by a digger- wasp, Ammophila abbreviata Fabr. which was found dragging it along the ground in the upland forest (Sta. IV,a). See Packard 229 (95, pp. 230-235) for an account of this forest-inhabiting larva. The larva of guttivitta is known to feed upon red maple, oak, and viburnum. GEOMETRDZ Eustroma diversilineata Hiibn. (P1. LXII, fig. 1.) This span-worm moth was taken in the upland forest (Sta. IV, a) Aug 26 (No. 163). Packard (Monogr. Geometrid Moths, p. 128. 1876) states that the larva feeds upon grape and Psedera. ‘These are mainly forest plants, and this is probably a woodland species. Caberodes confusaria Hiibn. This moth was taken near the upper slope of the south ravine in open woods (Sta. IV, b) Aug. 22 (No. 161). The larva feeds upon Trifolium. CocHLDnDZzZ Cochlidion or Lithacodes sp. Slug Caterpillar. This curious larva was found on a stump on the wooded ravine slope (Sta. IV, b) Aug. 26 (No. 165). GELECHIDA Vpsolophus ligulellus Hiibn. (?) These small moths were taken in the upland woods (Sta. IV, a) by T. L. Hankinson June 28, 1911 (No. 7678). The larva is reported on apple, pear, and plum. DIPTERA CECIDOMYIDA Cecidomyia holotricha O. S. (Hairy Midge-gall.) Abundant on the under side of hickory leaves (near Sta. IV) Aug. 20 (No. 96); and on leaves of Carya ovata in the upland for- est (Sta. IV,a) Aug. 26 (Nos. 107 and 170). These brownish, hairy galls may cover large areas on the under side of some leaves. See Cook ’05, p. 840, or Beutenmiiller ’04, p. 172.’ Cecidomyia tubicola O. S. (Hickory Tube-gall.) Immature galls (No. 107) were found Aug. 20 in the upland Bates woods (Sta. IV, a) on the lower side of leaves of Carya ovata. 230 Cecidomyta caryecola O. S. (Hickory Seed-gall.) This gall was taken on Carya ovata leaves in the upland forest (Sta. IV,a) Aug. 20 (No. 107); and Aug. 26 (No. 170). Many galls are formed on hickory and other trees by plant-lice (Cf. Per- gande, 02). ASILIDE Deromyia discolor Loew. This robber-fly was taken in an open area in the lowland forest (Sta. IV,c) Aug. 20 (No. 117). Williston (Kingsley’s Standard Natural History, Vol. 2, pp. 418-419. 1884) states that most robber- flies “rest upon the ground, and fly up when disturbed, with a quick buzzing sound only to alight again a short distance ahead. All their food, which consists wholly of other insects, is caught upon the wing . . . . Other flies and Hymenoptera are usually their food, but flying beetles, especially Cicindelide, are often caught, and they have even been known to seize and carry off large dragonflies. Not only will they feed upon other Asilide, but the female frequently resents the caresses of her mate by eating him up, especially if he is foolish enough to put himself in her power. In an instance the writer observed, a female seized a pair of her own species, and thrust- ing her proboscis into the thorax of the male, carried them both off together. . . . . The larve live chiefly under ground or in rotten wood, especially in places infested with grubs of beetles upon which they will feed. The young larve will bore their way completely within beetle larvze and remain enclosed until they have consumed them. Many, however, are found where they evidently feed upon rootlets or other vegetable substances. They undergo their trans- formations in the ground. The pupz have the head provided with tubercles, and on the abdominal segments there are also spiny pro- tuberances and transverse rows of bristles, which aid the insects to reach the surface when they are ready to escape as flies.” Mar- latt (Proc. Ent. Soc. Wash., Vol. 2, p. 82. 1893) observed D. dis- color preying upon wasps of the genus Vespa. By seizing the head of the wasp it avoids being stung. Deromyta umbrinus Loew. A specimen of this large robber-fly was taken in the south ravine (Sta. IV, d) by T. L. Hankinson, with the eucerid bee Melissodes perplexa Cresson in its grip, Aug. 22, 1910 (No. 7530). 231 SyrpHpz Chrysotoxum ventricosum Loew. This wasp-like fly was found resting on a leaf in the upland for- est (Sta. IV, a) Aug. 26 (No. 163). Mesogramma politum Say. Corn Syrphid. This fly was taken by T. L. Hankinson in the Bates woods (Sta. IV) June 28, 1911 (No. 7678). See the prairie list, p. 188. Milesia ornata Fabr. Vespa-like Syrphid. This beautiful large syrphid was taken on dogbane in an open space in the upland forest (Sta. IV,a) Aug. 20 (No. 103); in the open glade in the lowland forest (Sta. IV,c) Aug. 22 (No. 143); and on Aug. 26 (No. 184) on the flowers of Eupatorium celestinum in the clutches of the flower spider Misumena aleatoria Hentz. It was also taken in the Bates woods by T. L. Hankinson June 28, 1911 (No. 7678). Metcalf (’13, p. 73) quotes Verrall as follows con- cerning the subfamily Milestine: “What little is known about the metamorphism shows that many species live in rotten wood or about the sap flowing from injured tree trunks.” HYMENOPTERA Sirica Tremex columba Linn. Horntail; Pigeon Tremex. This species was taken on wing in the upland forest (Sta. IV, a) Aug. 16 (No. 66); and on the open slope of a ravine (Sta. IV. b) Aug. 22 (No. 132). The larva bores in the trunks of trees, as oak, elm, sycamore, and maple. Consult Packard (’90, pp. 379-381) for a description and figure of the larva. The long-sting, Thalessa lunator, is an external parasite upon this larva (see Riley, 88). I have taken normally colored females at Bloomington, Ill, July 25, Sept. 29, and Oct. 8. Two abnormally colored individuals were taken in September, one of them almost, and the other (taken Sept. 29) completely lacking the usual black markings. A female was taken at Milmine, III, in October. Consult Bradley (’13) for a key to the varieties of this species of Tremex. An interesting feature in the ecological relations of this species is the fact that it appears to frequent only weakened, diseased, and dying trees, and these, not as a primary invader, but as a trailer, following insects which have done previous injury to the trees. Felt (’05, p. 61) shows that in New York successive attacks of the 232 elm leaf-beetle, or injury by the sugar maple borer Plagionotus spectosus Say, prepare the way for the horntail larva. Ecologically considered, the leaf-beetle and the borer initiate a succession of in- sect invasions into the tree trunk; Tremex follows, with its parasite Thalessa; and these in turn lead the way for still others; thus a suc- cession of insects is produced. CynrrPDz Holcaspis globulus Fitch. (Oak Bullet Gall.) This gall was taken on white oak, Quercus alba, in the upland forest (Sta. IV, a) Aug. 26 (No. 170). Consult Cook (’05) and Beutenmiller (’04) for figures and de- scriptions of various kinds of galls mentioned in this list. Amphibolips confluens Harr. (Oak-apple or May-apple Gall. ) These galls were abundant upon the forest floor in the upland Bates woods (Sta. IV, a) during August (No. 101). The galls grow upon the leaves of several species of oaks (Quercus). Amphibolips prunus Walsh. (Acorn Plum Gall.) (Pl. LXII, fig. 2.) A single specimen of this gall was found on the slope of the south ravine in Bates woods (Sta. IV,b) Aug. 22 (No. 131). Another specimen came from the woods northeast of the Bates woods Aug. 20 (No. 96). It grows upon acorn cups. Andricus clavula Bass. (White Oak Club Gall.) (PI. LXTI, fig. 5.) This gall, formed in the terminal bud, was common on white oak, Quercus alba, in the upland Bates woods (Sta. IV, a) Aug. 26 (No. 170). Andricus cornigerus O. S. (Horned Knot Oak Gall.) (PI. LXII, fig. 3. This gall occurred in very large numbers on the branches of shingle oak, Quercus imbricaria, in a forest just northeast of the Bates woods, Aug. 20 (No. 96). The galls are old and apparently decaying. Andricus lana Fitch. (Oak Wool Gall.) (Pl. LXII, fig. 4.) Two examples of this gall were found on leaves of white oak, Quercus alba: one was taken near the Bates woods (Sta. IV) Aug. 20 (No. 96), and the other was found in the Bates woods (Sta. IV, a) on the petiole of a leaf, Aug. 26 (No. 170). Andricus seminator Harr. (Oak Seed-gall.) (Pl. LXIII, fig. 1.) A single specimen of this gall was found upon Quercus. alba (Sta. IV,a) Aug. 20 (No. 101). The cotton-like masses of this 233 gall are conspicuous. They may be tinged with red; when dry they become brownish. IcHNEUMONIDA Thalessa lunator Fabr. Lunate Long-sting. A female ichneumon of this species was found on a tree trunk in the open glade in the lowland forest (Sta. IV, c) Aug. 22 (No. 143). The larva feeds, as an external parasite, upon, the larva of the horntail, Tremex columba, which was also found in the Bates woods (Sta. IV). I found T. lunator, both males and females, abundant on shade trees at Bloomington, Ill., October 1, 1892, and also took it ' July 26, 1895. Riley (’88) gives an excellent account of this species accompanied by figures of the immature stages, and that of its host as well. Trogus obsidianator Brullé. This black ichneumon with fulvous antenne was taken in the Bates woods (Sta. IV) June 28, 1911, by T. L. Hankinson (No. 7678). This wasp is known to be parasitic upon the larva of Papilio polyxenes Fabr. (P. asterias—Insect Life, Vol. 1, p. 161) and upon the caterpillar of Pyrrharctia isabella (?). This species has been taken in central Illinois during June and July (Weed, Psyche, Vol. 5, p. 52). (See also Riley, in Amer. Ent., Vol. 3, p. 134. 1880.) PELECINIDA Pelecinus polyturator Drury. Black Longtail. (Pl. LXIII, fig. 2.) This remarkable looking insect was found in the glade of the lowland forest (Sta. IV,c) Aug. 20 (No. 117) and Aug. 22 (No. 143). Other females were seen in this forest. I have also’ taken this species at Bloomington, Ill. At Evanston, Ill., during July, 1910, this species was very abundant upon some damp lawns. I have counted four or five females in sight at once. They were often found upon blue-grass sod. The male of this species is considered very rare. The only one which I ever captured was taken July 29, 1910, at Evanston, Ill. The larva is parasitic upon the grub of the May-beetle, Lachnosterna (Forbes, Eighteenth Rep. State Ent. Ill, p. 124. 1894). It may also prey upon other scarabzeid larve inhabiting woodlands. Formicipz Stigmatomma pallipes Hald. Old-fashioned Ant. A single wingless queen and four pupe (No. 140) were taken Aug. 22 near the base of a ravine slope (Sta. IV, >) in dense shaded 234 woods, almost devoid of herbaceous vegetation, but with a thick layer of leaves, and other vegetable debris. Wheeler (Biol. Bull., Vol. 2, pp. 56-69. 1901) considers this a rather rare ant, although widely distributed over eastern North America. It is subterranean in habit, and ‘‘does not come to the surface even at night.” Contrary to the habits of must ants this primitive species has retained the carnivorous habits of the ancestral forms, and the young are fed on fragments of insects. They do not feed one another, or the larve by regurgitation, as do the specialized species of ants. They thus furnish us a glimpse at the ancient his- tory of ants. Wheeler (’05, pp 374-375) states that this species oc- curs only in “rich, rather damp woods, under stones, leaf mould, or more rarely under or in rotten logs.” A worker of Myrmica rubra Linn., subsp. scabrinodis Nyl., var. schencki Emery (No. 140) was taken from the same patch of leaves. Cremastogaster lineolata Say. (Pl. LXII, fig. 6.) This ant was taken only once—in the upland part of the Bates woods (Sta. IV, a) Aug. 20 (No. 118). Large numbers of the ants were found in an oak-apple gall (Amphibolips confluens Harr.) lying on the forest floor. When I picked up the gall, many ants came out and ran over my hand, biting vigorously. This is essentially a ground and forest-inhabiting ant, which forms nests in the soil, under stones, and in logs, stumps, etc. It has the peculiar instinct to make a sort of temporary nest out of debris to cover the aphids and coccids which it attends (Wheeler, Bull. Am. Mus, Nat. Hist., Vol. 22, pp. 1-18. 1906). Several carnivorous staphylinid beetles of the genus M. yrmedonia have been taken in the nests of these ants (Wheeler, ’10a, p. 382; Schwarz, ’gob, p. 247). Aphenogaster fulva Roger. A well-rotted stump in the upland Bates woods (Sta. IV, a) was found Aug. 17 to contain a moist, felt-like layer of some fungous growth, and on this was a large colony of snails (No. 71). In an adjacent part of this stump was a small colony of white ants, Termes flavipes Koll. (No. 72). A colony of ants which was in close prox- imity to the white ants, proved to be A. fulva Roger. As the gal- leries were exposed by cutting up the stump, these ants were seen to pick up the termites and carry them away, just as they do their own young on similar occasions. Five pairs—the ant and the termite which it carried—were preserved (Nos. 74-76, and 78-79). One of the termites lacks a head. All of them were workers. Larve and naked pupe (No. 79) were abundant in this nest, and workers (No. 80) were abundant about the stump. On Aug. 22 another 235. colony of this ant (No. 125) was found under the bark of a decaying oak stump (Sta. IV) in which the sap-wood was honeycombed, but the remainder solid, though discolored. Forel (Psyche, Vol. 9, p. 237. 1901) remarks that Aphenogas- ter is “very fond of termites, and when one uncovers and scatters about a nest of termites in a wood, they hasten to feast on the suc- culent morsels.” These observations suggest the possible fate of the captured termites; none of the ants were seen to eat them, how- ever. In the absence of observations, the missing head mentioned above may be variously accounted for. This habit of carrying off termites has been observed in other species of ants. Forbes (19th Rep. State Ent. Ill., p. 198. 1896) re- ports that near Carterville, Mason county, Ill, Mr. John Marten observed Formica schaufussi (=Formica pallide-fulva Linn., subsp. schaufussi Mayr) to pick up and carry away the living termites when its nest under a log in which termites abounded, was disturbed, and McCook (Proc. Acad. Nat. Sci. Phila., 1879, p. 155) has ob- served similar behavior in the case of the mound-building ant, For- mica exsectoides Forel. The histerid beetle Heterius blanchardi Schwarz has been found in nests of this ant (Wheeler, ’10a, pp. 388, 389) ; and European ob- servers have seen ants carrying and rolling them about. Consult also Schwarz (’9ob, 247) for a list of beetles found with this ant. Wheeler (’10a, p. 206) lists A. fulva as a glade species which in the forests utilizes logs and branches as‘substitutes for stones. (See Wheeler, ’05, pp. 372-373.) Aphenogaster tennesseensis Mayr. Tennessee Ant. A colony of this ant (No. 87) was taken Aug. 17 from a decaying stump, situated on the slope (Sta. IV, >) from the upland forest to the lowland on the river bottom. According to Wheeler (Bull. Am. Mus. Nat. Hist., Vol. 20, 1904, p. 362, and Vol. 21, 1905, p. 373) this species normally nests in dead wood in rather open forests. He holds the opinion that the queen of this species can not rear her own brood, and thus establish a new colony, but must utilize a small or weak colony of the allied species A. fulva Roger, which lives under stones. Thus the new colonies are started under stones; later, when they become numerous, they are found in rotten wood. This, Wheeler concludes, indicates that they “migrate away from the fulva workers.” Tanquary (11) has per- formed some interesting experiments which show that queens of tennesseensis are adopted by colonies of other ants, a result which seems to confirm Wheeler’s anticipation. Schwarz (’9ob, p. 247) records two beetles found with this ant. 236 Formica fusca Linn., var. subsericea Say. This ant was taken in the upland Bates woods (Sta. IV, a) Aug. 26 (No. 163). See the list of prairie invertebrates, p. 190. Myrmica rubra Linn., subsp. scabrinodis Nyl., var. schencki Emery. This ant (No. 140) was found Aug. 22 under leaves in a small ravine on a shady slope (Sta. IV. b) from the upland forest to the valley bottoms. The soil under these leaves had been thoroughly tun- neled by small mammals during the preceding winter, but recently the leaves had not been disturbed. The soil was a mixture of sand, clay, and vegetable debris, was moist, and contained few kinds of animals. A single ant of this variety (No. 140) was taken while collecting spec- imens of Stigmatomma pallipes. This species is listed by Wheeler (Bull. Am. Mus. Nat. Hist., Vol. 21, p. 373. 1905) as a field ant which prefers to nest in grassy pas- tures and lawns, in situations exposed to the sun. Our specimen was, therefore, found in an unusual habitat. Tapinoma sessile Say. Cocoanut Ant. This cocoanut ant, so called because of the odor of the workers, which has been compared to that of decayed cocoanuts, was found in the lowland part of the Bates woods, at the base of the slope to the bottoms (Sta. IV,c) Aug. 22 (No. 139). A large colony was found among the surface layers of dry dead leaves; from it were se- cured two queens, vast numbers of eggs, and also larve, pupa, and workers.. Wheeler (’05, pp. 373, 389) states that this ant usually nests in open sunny woods, the borders of woods, and under stones, logs, etc. Schwarz (’9ob, p. 247) records beetles as living with this ant. Camponotus herculeanus Linn., subsp. pennsylvanicus DeG. Carpen- ter Ant. This species was taken from under the bark of a rotting: stump among a dense second-growth, on the valley slope (Sta. IV, b) be- tween the upland and the lowland forest Aug. 17 (No. 84). This stump was in that stage of decay so often utilized by the large Caro- lina slug, Philomycus carolinensis, and the horned Passalus beetle, Passalus cornutus. The colony was recently founded, for the dea- lated female occupied a small cell excavated in the rotten sap-wood. This colony consisted of four pupz and six larve of different sizes. Another colony was taken in the same stump, from the rotted sap- wood zone, in company with the snail Philomycus carolinensis and some kind of pulmonate snail eggs. This colony was in a more ad- vanced stage than the preceding, about a dozen larve, seven pup, 237 and two adult workers being present, and about half a dozen eggs (No. 85). Pricer (’08) has given an interesting account of the life history and habits of this ant in Illinois. He states (p. 197) that the food is largely the honeydew of plant-lice, but is supplemented by plant juices and dead insects. He found a small staphylinid beetle, Xeno- dusa cava, abundant in the nests. I have found pennsylvanicus abundant at Bloomington, Ill., and represented as follows: by a male June 29; by a winged female in June; and by dealated females June 29 and July 2 and 25. McCook (’83) has given an interesting account of the found- ing of colonies of this ant. See also Wheeler, ’o6b, pp. 38-39, Plate VII, and ’1ob, pp. 335-338, for further information concerning it. Camponotus herculeanus Linn., subsp. pennsylvanicus DeG., var. ferrugineus Fabr. This variety was taken a short distance to the northeast of the Bates woods (Sta. FV) Aug. 20 (No. 97). Here the large ground- beetle Calosoma scrutator was found running on the ground with what appeared to be a bunch of greenish moss; a large reddish ant also struggled for possession of the prize. Upon closer examination it was found that the skin of some large lepidopterous larva was the object desired. This skin, recently shed or moistened by a recent rain, was a prize for both ferrugineus and Calosoma. A dead wingless ferrugineus, covered with a fungus growth, was found in a small cell excavated in the rotten wood of a decaying log on the ravine slope (Sta. IV, b) Aug. 17 (No. 90). Apparently this female had died before her colony developed. (See Pricer, ’08; Wheeler ’10b, pp. 338-339.) I have found this form abundant at Bloomington, Ill. Winged females were taken July 26, dealated ones on July 25 and 26, and males June 29, and July 9 and 25. On July 21, 1892, several males were taken at night, being attracted to a lamp located near a small brook. A very large colony, numbering thousands of individuals, was found May 26, under a well-decayed log, in a forest at White Heath, Ill. It contained winged males, females, and workers. The winged forms were present in vast numbers. The far-advanced condition of decay of the log was in marked contrast with that in which the initial colonies are usually found. During the years of development of such a large colony the progress of decay will naturally make some changes in the habitat; reciprocally the ants will doubtless tend to monopolize the logs to the exclusion of some other animals, and 238 also facilitate the decay of the log by their activities. There is an “orderly sequence” of changes in the developing colony, and a simi- lar orderly sequence of changes in the log habitat. An ant colony in its development clearly illustrates the transfor- mation from the individual to the associational phase of ecological relations. Beginning with the fertilized female and her progeny, the colony develops in size and in the division of labor among its members; until, finally, by the possible addition of slaves, commen- sals, parasites, and even predaceous enemies, the colony or associa- tion is built up in an orderly sequence, and the organisms adjust themselves to one another and to the environment in general. MuTILLDA Spherophthalma sp. 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