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Xerox University Microfilms 300 North Zoob Road Ann Arbor, Michigan 48103 76-3469 KEFERL, Eugene Paul, 1940- AN ECOLOGICAL STUDY OF THE MOLLUSCA OF THE CEDAR BOG NATURE SANCTUARY, CHAMPAIGN COUNTY, OHIO. The Ohio State University, Ph.D., 1975 Zoology

■ Xerox University Microfilmst Ann Arbor, Michigan 48106

THIS DISSERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED. PLEASE NOTE: Page 66 is not available for photography. UNIVERSITY MICROFILMS AN ECOLOGICAL STUDY OF THE HOLLUSCA OF THE CEDAR BOG

NATURE SANCTUARY, CHAMPAIGN COUNTY, OHIO

DISSERTATION

Presented In Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University

By

Eugene Paul Keferl, B.Sc. ******

The Ohio State University

1975

Reading Committee: Approved By

Dr. N. Wilson Britt

Dr. David H. Stansbery £ Co-Advisor Dr. John L. Crltes Department of Entomology

Co-Advisor Department of Zoology ACKNOWLEDGMENTS

I would like to express my sincere gratitude and thanks to my advisers, Dr. N. Wilson Britt and Dr. David H. Stansbery for help and guidance throughout this study. I wish to thank Dr. Clara May Frederick for the use of her original Cedar Bog maps; Mr. Kenneth L. Powell, for the use of his personal notes on the soils of the region; Mr. Ted

Smith, for his invaluable assistance with the statistics; Dr. William

J. Clench, for identifying the'Physidae; Dr. Dorothea S. Franzen, for much help identifying many of the Succineidae; Dr. William Heard, for his identification of the Sphaeriidae; and Mr. Leslie Hubricht, for his verification of Stenotrema barbatum. I also wish to thank the Ohio

Historical Society and Lottie McDaniel for allowing me to make this study at Cedar Bog. Finally, I wish to extend special thanks to Dr.

Charlsie A. Keferl, my wife; Mr. John M. Condit; and Dr. Carol B. Stein for their unrelenting pressure to complete this study.

ii VITA

September 17, 1940 ..... Born - Dayton, Ohio

1963...... B.Sc., University of Dayton, Dayton, Ohio

1964-1969 ...... Teaching Assistant, Department of Zoology, The Ohio State University

1970 ...... Teaching Associate, CORE Biology Pro­ gram, Department of Zoology, The Ohio State University

1975 ...... Technical Assistant, Museum of Zoology, Department of Zoology, The Ohio State University

PUBLICATIONS

"Some Mollusca of Cedar Bog, Champaign County, Ohio." American Malacological Union, Annual Report of 1969, pp. 14-16, 1969.

"AdansonTs Slite Shells." Of Sea and Shore, Vol. IV, No. 1, p. 27, 1973.

"Terrestrial Mollusca of Cedar Bog." Cedar Bog Symposium, Ohio Academy of Science, Ohio Biological Survey Informative Circular No. 4, pp. 46- 48, 1974.

FIELDS OF STUDY

Major Field: Invertebrate Zoology

Studies in Invertebrate Zoology. Professors Joseph N. Miller, A. Carter Broad, Paul A. Colinvaux, John L. Crites, N. Wilson Britt, and Barry D. Valentine

Studies in Ecology. Professors David H. Stansbery, Gareth E. Gilbert, Lionel G. Klikoff, Jerry H. Hubschman, Tony J. Peterle, and N. Wilson Britt

ill TABLE OF CONTENTS

Page

ACKNOWLEDGMENTS...... 11

VITA ...... ill

LIST OF T A B L E S ...... v

LIST OF PLATES AND F I G U R E S ...... vli

LIST OF M A P S ...... x

LIST OF G R A P H ...... xv

INTRODUCTION...... 1

THE CEDAR BOG NATURE SANCTUARY STUDY AR E A ...... 4

GEOLOGY OF CEDAR B O G ...... 6

SOILS ...... 8

CLIMATOLOGY OF CEDAR B O G ...... ■ ...... 20

CEDAR R U N ...... 22

VEGETATION OF CEDAR B O G ...... 26

METHODS AND MATERIALS...... 48

RESULTS...... 52

DISCUSSION OF DEFINITIONS AND DATA ANALYSIS OF PROCEDURES .... 78

DISCUSSION OF THE SPECIES COLLECTED AT CEDAR B O G ...... 84

DISCUSSION OF VARIOUS MOLLUSCAN F A U N A S ...... 237

DISCUSSION OF THE DIVERSITY I N D E X ...... 271

THE MOLLUSCA* OF THE ARBOR VITAE ASSOCIATION...... 275

REFERENCES ...... 282

lv LIST OF TABLES

Table Page

1. Plant Diversity as Recorded by Recent Investigators at Cedar Bog, Champaign County, Ohio ...... 2

2. The Basic Characteristics of the Soils Found in the Cedar Bog Nature Preserve, Champaign County, Ohio ...... 14

3. A Summary of the Arbor Vitae Association at Cedar Bog Nature Preserve, Champaign County, Ohio ...... 30

4. Species Composition of the Plant Associations at the Cedar Bog Nature Preserve, Champaign County, Ohio ...... 32

5. Species Composition and Relic Plant Diversity in the Plant Associations at the Cedar Bog Nature Preserve, Champaign County, Ohio as Interpreted by the Environment Control Corporation ...... 40

6. Collections made at Cedar Bog Nature Preserve, Champaign County, Ohio ...... 55

7. Phylogenetic Listing of the Mollusca Collected at the Cedar Bog Nature Preserve, Champaign County, Ohio.... 74

8. Mollusca Collected at Cedar Bog Nature Preserve, Champaign County, Ohio by Other Investigators ...... 76

9. Aquatic Molluscs Living in and Around the Cedar Bog Nature Preserve, Champaign County, Ohio ...... 93

10. Habitat Diversity Indices for the Mollusca Found at Cedar B o g ...... 238

11. Distribution of Molluscs in the Various Vegetation Zones at Cedar Bog Nature Preserve, Champaign County, Ohio . . . 240

12. A Comparison of the Major Statistics Concerning the Molluscan Fauna of each of the Frederick Vegetation Z o n e s ...... 244

v Table ■Page

13. Distribution of Molluscs in the Various Vegetation Zones at • Cedar Bog Nature Preserve, Champaign County, Ohio .... 253

14. A Comparison of the Major Statistics Concerning the Molluscan Fauna of Each of the Environment Control Corporation Vegetation Zones ...... 257

15. Distribution of Molluscs in the Soil Zones at the Cedar Bog Nature Preserve, Champaign County, Ohio ...... 259

16. A Comparison of the Major Statistics Concerning the Molluscan Fauna of Each of the Various Soil Zones .... 262

17. Distribution of Molluscs in the Soil Zones at the Cedar Bog Nature Preserve, Champaign County, Ohio ...... 263

18. Mollusca Recorded from the Arbor Vitae Associations in Ohio, Michigan, Wisconsin and Ontario ...... 276

vi LIST OF PLATES AND FIGURES

Plate Figure Page

I. 1 Collection Frequencies for Pomatlopsis l a p i d a r i a ...... 95

2. Collection Frequencies for Lymnaea humilis .... 95

3. Collection Frequencies for Gyraulus parvus .... 95

4. Collection Frequencies for Physa integra ...... 95

II. 1. Collection Frequencies for Carychium exiguum . . . 112

2. Collection Frequencies for Carycium exile .... 112

3. Collection Frequencies for Carychium nannodes . . 112

4. Collection Frequencies for Gastrocopta c o n t r a c t a ...... 112

III. 1. Collection Frequencies for Gastrocopta pentodon...... 134

2. Collection Frequencies for Gastrocopta tappaniana...... 134

3. Collection Frequencies for Vertigo morsel .... 134

4. Collection Frequencies for Vertigo ovata ...... 134

IV. 1. Collection Frequencies for Vertigo pygmaea .... 147

2. Collection Frequencies for Vertigo tridentata . . 147

3. Collection Frequencies for Vertigo gouldi .... 147

4. Collection Frequencies for Vertigo milium .... 147

V. 1. Collection Frequencies for Columella edentula . . 158

2. Collection Frequencies for Strobilops aenea . . . 158 •

3. Collection Frequencies for Strobilops labyrinthica ...... 158

4. Collection Frequencies for Succinea ovalis .... 158 vli Plate Figure Page

VI. 1. Collection Frequencies for Catinella vermeta . . . 170

2. Collection Frequencies for Oxyloma spp...... 170

3. Collection Frequencies for Philomycus carolinianus ...... 170

4. Collection Frequencies for Anguispira a l t e r n a t a ...... 170

VII. 1. Collection Frequencies for Helicodiscus parallelus...... 182

2. Collection Frequencies for Punctum minutissimum . 182

3. Collection Frequencies for Deroceras Laeve .... 182

4. Collection Frequencies for Nesovitrea binneyana . 182

VIII. 1. Collection Frequencies for Nesovitrea e l e c t r i n a ...... 193

2. Collection Frequencies for Glyphyalinia indentata ...... 193

3. Collection Frequencies for Hawaiia minuscula . . . 193

4. Collection Frequencies for Euconulus chersinus . . 193

IX. 1. Collection Frequencies for Euconulus fulvus . . . 204

2. Collection Frequencies for Zonitoides arboreuB...... 204

3. Collection Frequencies for Guppya sterkii .... 204

4. Collection Frequencies for Striatura milium . . . 204

1. Collection Frequencies for Striatura exigua . . . 214

2. Collection Frequencies for Haplotrema concavum . . 214

3. Collection Frequencies for Stenotrema barbatum . . 214

4. Collection Frequencies for Stenotrema leai .... 214

viii Plate Figure Page

XI. 1 Collection Frequencies for Mesodon clausus ...... 223

2. Collection Frequencies for Mesodon thyroidus . . . 223

3. Collection Frequencies for Mesodon inflectus . . . 223

4. Collection Frequencies for Triodopsis multilineata ...... 223

XII. 1. Collection Frequencies for Allogona profunda . . . 233

2. Collection Frequencies for Pisidium casertanum...... 233

ix LIST OF MAPS

Map Page

I. Topo Map, Cedar Bog Nature Preserve, Champaign County, Ohio ...... 5

II. Soil Map of Cedar Bog (Modified from Ritchie et al., 1971: Map 3F-206)...... 9

III. Soil Map of Cedar Bog (Modified from Environment Control Corporation, 1973: C-9) ...... 17

IV. Vegetation Map of Cedar Bog (Modified from Frederick, 1967: 50-53) ...... 28

V. Vegetation Map of Cedar Bog (Modified from Environment Control Corporation, 1973: B-2 and B - 3 ) ...... 38

VI. Vegetative Cover of Cedar Bog in 1938 (Adapted from a 1938 Aerial Photograph Published by the Clark County Audubon Society, 1972: 17 ) ...... 46

VII. Collection Locations at Cedar Bog, Champaign County, O h i o ...... 54

VIII. Collection Location for Valvata tricarlnata (Say) at Cedar Bog ...... 85

IX. Collection Locations for Valvata sineera Say at Cedar B o g ...... 87

X. Collection Locations for Amnicola limosa (Say) at Cedar B o g ...... 89

XI. Collection Locations for Pomatiopsis lapidaria (Say) at Cedar Bo g ...... 92

XII. Collection Locations for Oxytrema semlcarinata (Say) at Cedar Bo g ...... 97

XIII. Collection Locations for Lymnaea humilis (Say) at Cedar B o g ...... 100

x Collection Locations for Hellsoma anceps (Menke) at Cedar Bog ...... 102

Collection Locations for Gyraulus Parvus (Say) at Cedar Bog ...... 104

Collection Locations for Planorbula armlgera (Say) at Cedar Bog ...... 106

Collection Locations for Physa Integra Haldeman at Cedar Bog ...... 108

Collection Locations for Carychium exiguum (Say) at Cedar Bog ...... 111

Collection Locations for Carychium exile H. C. Lea at Cedar Bog ...... 115

Collection Locations for Carychium nannodes Clapp at Cedar Bog ...... 117

Collection Locations for Cochlicopa lubrica (0. F. Mllller) at Cedar B o g ...... 120

Collection Locations for Vallonia excentrica Sterki at Cedar Bog ...... 122

Collection Locations for Pupoldes albilabris (C. B. Adams) at Cedar Bog ...... 124

Collection Locations for Gastrocopta procera (Gould) at Cedar B o g ...... - . 126

Collection Locations for Gastrocopta armifera (Say) at Cedar Bog ...... 128

Collection Locations for Gastrocopta contracta (Say) at Cedar Bog ...... 131

Collection Locations for Gastrocopta pentodon (Say) at Cedar Bog ...... 133

Collection Locations for Gastrocopta tappaniana (C. B. Adams) at Cedar Bog ...... 137

xi Collection Location for Gastrocopta corticaria (Say) at Cedar Bog ...... 139

Collection Locations for Vertigo morsel SterkL at Cedar Bog ...... 141

Collection Locations for Vertigo ovata Say at Cedar B o g ...... 143

Collection Locations for Vertigo pygmaea (Draparnaud) at Cedar Bog ...... 146

Collection Locations for Vertigo tridentata Wolf at Cedar Bog ...... 149

Collection Locations for Vertigo elatlor Sterki at Cedar Bog ...... 151

Collection Locations for Vertigo gouldi (Binney) at Cedar Bog ...... 152

Collection Locations for Vertigo milium (Gould) at Cedar B o g ...... - ...... 155

Collection Locations for Columella edentula (Draparnaud) at Cedar Bog ...... 157

Collection Locations for Strobilops aenea Pilsbry at Cedar Bog ...... 160

Collection Locations for Strobilops labyrinthica (Say) at Cedar Bog ...... 163

Collection Locations for Succinea ovalis Say at Cedar Bog ...... 166

Collection Locations for Catinella vermeta (Say) at Cedar Bog ...... 169

Collection Locations for Oxyloma spp. at Cedar Bog ...... 172

Collection Locations for Philomycus carolinianus (Bose.) at Cedar Bog ...... 176 xii Map Page

XLIV. Collection Locations for Anguispira alternata (Say) at Cedar B o g ...... 178

XLV. Collection Locations for Helicodiscus parallelus (Say) at Cedar B o g ...... 181

XLVI. Collection Locations for Puneturn minutissimum (Lea) at Cedar B o g ...... 184

XLVII. Collection Locations for Deroceras Laeve (Mtlller) at Cedar Bog ...... 187

XLVIII. Collection Locations for Nesovitrea binneyana (Morse) at Cedar B o g ...... 190

XLIX. Collection Locations for Nesovitrea electrina (Gould) at Cedar B o g ...... 192

L. Collection Locations for Glyphyalinia indentata (Say) at Cedar B o g ...... 196

LI. Collection Locations for Hawaiia minuscula (Binney) at Cedar B o g ...... 198

LII. Collection Locations for Euconulus chersi'nus (Say) at Cedar B o g ...... 200

LIXI. Collection Locations for Euconulus fulvus (Mtlller) at Cedar B o g ...... 203

LIV. Collection Locations for Zonitoides arboreus (Say) at Cedar B o g ...... 206

LV. Collection Locations for Guppya sterkii (Dali) at Cedar B o g ...... 208

LVI. Collection Locations for Striatura milium (Morse) at Cedar B o g ...... 211

LVII. Collection Locations for Striatura exigua (Stimpson) at Cedar B o g ...... 213

LVIII. Collection Locations for Haplotrema concavum (Say) at Cedar Bog ...... 216

xiii Map Page

LIX. Collection Locations for Stenotrema barbatum (Clapp) at Cedar B o g ...... 218

LX. Collection Locations for Stenotrema leai (Binney) at Cedar B o g ...... 220

LXI. Collection Locations for Mesodon clausus (Say) at Cedar B o g ...... 222

LXXI. Collection Locations for Mesodon thyroidus (Say) at Cedar B o g ...... 226

LXIII. Collection Locations for Mesodon inflectus (Say) at Cedar B o g ...... 228

LXIV. Collection Locations for Triodopsis multilineata (Say) at Cedar Bog...... 230

LXV. Collection Locations for Allogona profunda (Say) at Cedar B o g ...... 232

LXVI. Collection Locations for Pisidium compressum Prime at Cedar B o g ...... 235

LXVII. Collection Locations for Pisidium casertanum (Poli) at Cedar B o g ...... 236

LXVIII. Distribution of Carychium nannodes in the United States ...... 280

LXIX. Distribution of Vertigo morsei in the United States . . 281

xiv LIST OF GRAPH

Graph Page

I. The Relation between Diversity Index (H), Based on the Shannon-Wiener Equation, and the Number of Species (Diversity) for Various Zonation Schemes in Cedar Bog Nature Preserve, Champaign County, Ohio ...... 272

xv INTRODUCTION

The Cedar Bog Nature Sanctuary in Champaign County,' Ohio is an area of special significance. For several thousands of years since the Wisconsin Glacier receded from this area, a constant source of alkaline water has surfaced where the Bog stands today. These alkaline springs have helped maintain an environment found nowhere else in Ohio. Within this Bog several unique communities continue to persist far from their normal range. Many of these plants are normal constituents of the northern coniferous forests, the western wet prairies and the southeastern coastal plain.

Cedar Bog has attracted many botanists since the early 1830fs (Stuckey, 1973). The first published account of this area was pro­ vided by Dachnowski (1910) in which he describes the vegetation and soils. Since Dachnowski numerous studies have been made concerning the vegetation at Cedar Bog (Table 1). As a result of these studies much is understood about the geology, soils and vegetation of Cedar Bog. The diversity of plant species found at Cedar Bog is high and perhaps unmatched in any other area of comparable size in Ohio.

The first faunal study at Cedar Bog was made by Franks (1931) on the vertebrates. Since then Cavender and Yoder (1973) made some studies on the fishes found in Cedar Run; Edward Thomas (1973) on some Orthoptera; Carl Albrecht (1973 and 1974) on some Lepidoptera; Jon Williams (1966) on the aquatic invertebrates, mainly insects; Keferl (1969 and 1973) on mollusks; and the Environment Control Corporation (1973) on the aquatic invertebrates. Each investigator who has studied a particular animal group at the Bog has found unusual, rare and even endangered species living there.

Except for unpublished records of snails collected by Mrs. R. P. Klein in 1951 and Jon Williams (1966), the molluscs of" the Cedar Bog Nature Preserve are unknown. In fact, very few molluscs were recorded previously from any where in Champaign County. This study attempts to determine which species of mollusks Inhabit Cedar Bog and, Insofar as possible, what factors determine their distribution.

This is an ecological study of the mollusca of the Cedar Bog Nature Preserve. Some of the objectives of this study hinge greatly on the unique nature of the area and the extensive Information available concerning its flora and soils. Every reasonable effort has been made to determine what molluscs are found in the Bog and where. Correlations between the density and diversity of the mollusc species, the vegetation and the soils are investigated.

1 2

TABLE 1

Plant diversity as recorded by recent Investigators at Cedar Bog, Champaign County, Ohio.

Number of Plant Group Studied Investigator Taxa Recorded

Myxomycetes Braun (1973: 40-43) * 26 (Slime Molds) -

Fungi Cooke (1973: 31-35) 116

Algae (Exclusive of Diatoms) Briggs (1972: 113-128) 84

Algae (Diatoms Only) Hufford (1972: 130-137) 302

Lichens Rudolph (1973: 38-40) 49

Mosses and Environment 39 Liverworts Control Corporation (1973: Appendix B) Vascular Plants Frederick (1974: 114) 546

Living Taxa 516

Extinct Taxa 30

Disjunct Species Frederick (1973: 16-20) 53

Northern 33

Southern 8

Prairie 5

Other 7 Total Number of Taxa Recorded 1162 3

Several ecological surv.eys concerning terrestrial molluscs have been made in the United States. Some of these studies have attempted to correlate terrestrial mollusc distribution and density with the vegetation and soils present. F. C. Baker (1939) in Illinois; H. B. Baker (1911 and 1922) in Michigan; Burch (1955 and 1956) in Virginia; Karlin (1961) in Colorado, Montana and New Mexico; Jacot (1935) in North Carolina; Ingram (1941 and 1946) in New York; Shimek (1930) in Iowa; and Bash, Bainer and Wilhm (1961) in Kansas are some notable examples of this kind of study. However, all these studies have dealt with plant associations which were normal for the area. No one as yet has intensively studied the molluscs of a plant association far from its normal range.

My investigation of the molluscs of one particular relic plant association, the arbor vitae or northern white cedar association, was extended beyond Cedar Bog, Ohio. Visits were made to Michigan, Wisconsin and Canada where the arbor vitae association is a natural part of the vegetation. It was anticipated that these extended Inves­ tigations would help to interpret the mollusc populations of the arbor vitae association in Ohio. It was hoped that some insight as to the true nature of a relic population could be made.

Ecological data on most terrestrial molluscs is limited. Much of the Information available comes from surveys made in Virginia, Michigan, Illinois and Kansas. It is far from being complete. For some species the habitat Information is based upon one or two collections. The goal of this study is to make available needed ecological information con­ cerning some of the terrestrial molluscs found in Ohio. A special effort was made to secure information concerning habitats and limiting factors. THE CEDAR BOG NATURE SANCTUARY STUDY AREA

The Cedar Bog Nature Sanctuary is located In sections 31 and 32, Urbana Township, Champaign County, Ohio in the Urbana West 7.5 minute' topographic quadrangle (Map I). The Bog lies at approximately 40° 3* 11" north latitude and 83° 47* 30" west longitude. The study area is accessible via Woodburn Road and United States 68, 4.3 miles south southeast of Urbana.

The Bog has been under the supervision of the Ohio Historical Society since 1942. This study took place in the original 98.45 acres and approximately 15 acres of the McDaniel property immediately east of East Branch of Cedar Run. The original acreage for the Bog was pur­ chased in 1941 by the state of Ohio. In 1968 the Nature Conservancy purchased an additional 105.41 acres and added it to the preserve. This extension Is west of the original tract and was not Included in this study.

4 M A P I

Topographic map of Cedar Bog Nature Sanctuary, Champaign County, Ohio

•951 970 930

HICKOR GROY.E If 000

992

965

*'"970

900

WOODBURN ROAD

978

Current Cedar Bog Nature Sanctuary boundary.

Study area boundary. GEOLOGY OF CEDAR BOG

Historical Geology

Cedar Bog lies in the Mad River Valley Train which is the largest Wisconsin outwash deposit in Ohio (Jansens, 1964: 33). Cummins (1959: Plate I) shows that Cedar Bog lies directly over the junction of the old Pre-Kansan Teays River valley and the Deep Stage stream, the Middletown River. The Teays River flowed north at an elevation of about 530 feet above sea level, on Ordovican bedrock. This bedrock is composed of alternating layers of shale and limestone (Harker and Bernhagen, 1943: 15). A south-moving Pre-Kansan or the Kansan glacier blocked the north-flowing Teays River and impounded the water over much of the southern part of Ohio (Stout et al., 1943: 80). As the Pre- Kansan or Kansan glacier receded the Middletown River was formed. This river flowed south at an elevation of about 830 feet on Pre-Kansan glacial deposits (Stout$ Ver Steeg and Lamb, 1943: 78). The Middletown River followed approximately the present course of the present Mad River Valley (Harker & Bernhagen, 1943: 10). Much of this Deep Stage Middletown River is filled with various types of glacial deposits from the Illlnoian glacier (Stout et al., 1964: 86).

The last glacier, the Wisconsin, was relatively thin in this region of Ohio and its affect on the area was quite different than the pre­ ceding glaciers (Frederick, 1964: 12 and 1967: 19). The thin Wisconsin ice was slowed by a high elevation of bedrock at Beliefontaine, Logan County, Ohio. This high bedrock divided the glacier in two lobes, the western Miami lobe and the eastern Scioto lobe. Together these two lobes formed the parallel moraines which border the Mad River valey today. This situation resulted in the area between the Miami and Scioto moraine systems being a natural drainage basin for melt waters. Therefore, the resultant glacial deposits were primarily outwash rather than glacial till (Jansens, 1964: 4). Glacial outwash is composed of sorted sands and gravels and with very little silt and clay, hence is a potential aquifer with a great water storage capacity.

Hydrology

The existence of Cedar Bog and its boreal relicts is dependent upon a continuous flow of cool, highly calcareous ground water. Accord-' ing to Cummins (1959: 1-2) the Teays River and the Deep Stage Middletown River valleys are good sources of ground water. Cedar Bog lies over the junction of these two buried river valleys. Since outwash of the Mad River Valley Train is the immediate sustrate, this underground water can 6 7 readily move through these sorted sands and gravels to the surface. This view does not completely explain why spring water is coming to the surface only in this particular area. Forsyth (1974: 116) states,

Regionally the water table in the Mad River Valley Train lies only about 10 feet below the surface, but is too deep to create such springs. However, ground water moving through a higher outwash level, the Urbana Outwash, to the northeast, provides so much additional ground water, at the place where this outwash has been truncated by the lateral erosion of the Mad River, that the water table in the Mad River Valley Train is locally raised enough to reach the land surface and produce these springs. Contours drawn on the water table near the Bog demon­ strate that the main source of ground water feeding the Bog is the Urbana Outwash (this water becomes super­ imposed on the moving down through the Mad River Valley Train) and not the very deeply burled fill of the pre­ glacial Teays valley, which lies almost directly below the bog.

Environment Control Corporation (1973: 111-12) found that 37% of the water is supplied by regional flow and 63% of the water is sup­ plied by local flow. The regional flow comes from the deep outwash drainage and is little affected by seasonal climatic conditions. The local water flow is above the regional water and is affected by seasonal oscillations of precipitation. A change in either water source would cause alterations at the Bog (Environment Control Corpor­ ation, 1973: 111-13). SOILS

United States Department of Agriculture Interpretation

According to the U.S.D.A. soil survey (Ritchie, Powell and Siegenthaler, 1971) there are three basic soils in the Cedar Bog study area, the Lippincott silty-clay loam, the Carlisle muck and the Lin- wood muck. This survey does not recognize peat and calcareous tufa or marl as soils. According to Mr. K. L. Powell (1971), these substrates are regarded as inclusions in the Carlisle muck. Therefore, I modi­ fied the original U.S.D.A. soil map map to include the peat and marl deposits (Map II). Table 2 gives a brief summary of some of the gen­ eral characteristics of each of these substrates as taken from Ritchie et al. (1971), Dachnowski (1912) and Stout (1940).

Lippincott silty-clay loam

The Lippincott silty-clay loam (formerly called the Abington silty-clay loam) is a dark colored, very poorly drained, neutral to moderately alkaline and highly productive soil. It is high in fertility, moisture holding capacity and organic matter. It is formed in silty deposits underlain by stratified calcareous sand and gravel at 50 inches or more on depressed areas of terraces (Ritchie et al., 1971: 60). This soil is found in the grassy meadows west of the study area, in parts of the beech-maple association and in the shrub zones in the southwestern section of Cedar Bog. The best developed beech-maple stand corresponds well with this soil type.

The following Is a typical profile of Lippincott silty-clay loam taken In Champaign County as recorded in Ritchie et al. (1971: 60).

Ap— 0 to 7 inches, black (10YR 2/1) silty-clay loam; moderate, medium and coarse, granular structure; friable when moist; neutral; abrupt, smooth boundary. A3— 7 to 12 inches, black (10YR 2/1) silty-clay loam; moderate, coarse, angular blocky structure; firm when moist;, neutral; clear, wavy boundary. B21tg— 12 to 16 Inches, very dark gray (10YR 3/1) silty-clay loam; many, fine, distinct, yellowish-brown (10YR 5/6) mottles; moderate, fine and medium, prismatic structure breaking to strong, medium, angular blocky structure; very firm when moist; some crayfish channels; thin,

8 9

m a p n

Soil map of Cedar Bog (modified from Ritchie et al., 1971: Map 3F-206).

Lippincott Silty-clay Loam

Carlisle Muck

Linwood Muck

Peat

Marl map n Soil map of Cedar Bog (modified from Ritchie et a l., 1971: Map 3F-206).

w

P « ir.r <

320 11

continuous clay coatings and coatings of organic matter on the peds; neutral; clear wavy boundary. B22tg— 16 to 27 Inches, dark grayish-brown (2.5Y 4/2) silty clay; common, medium, distinct, strong-brown (7.SYR 5/6) mottles; strong, medium, prismatic structure breaking to strong, medium, angular blocky structure; very firm when moist; moderate, patchy clay films on ped faces; some crayfish channels; neutral; clear, wavy boundary. IIB3t— 27 to 36 inches, grayish-brown (2.5Y 5/2) gravelly silt loam; massive; friable when moist; many,' light- gray (2.5Y 7/2), partly weathered dolomite pebbles; a few crayfish channels; few dark-gray (10YR 4/1) clay films on vertical surfaces; moderately alkaline and calcareous; clear, wavy boundary. IlCi— 36 to 50 Inches, grayish-brown (2.5Y 5/2) gravel and sand; common, medium, distinct, yellowish-brown (10YR 5/6) mottles; single grain; loose; calcareous; clear, smooth boundary. IIC2— 50 inches +, gray (5Y 5/1) gravel and sand; loose; calcareous.

Linwood muck

Linwood muck (formerly called Tawas muck) is a poorly drained, slightly acid to mildly alkaline and highly productive soil. It develops on woody and fibrous material underlain at 42 inches or more by sand and gravel. It appears as If Linwood muck is somewhat Inter­ mediate between Carlisle muck and Lippincott silty-clay loam. Linwood muck Is shallower than Carlisle muck and is generally underlain with a loamy material rather than peat or marl. The Linwood muck is an organic soil whereas the Lippincott silty-clay loam Is a soil derived from a mineral parent material with later addition of much organic matter. These three soils are frequently found together.

At Cedar Bog, Linwood muck is found in parts of the beech-maple association, the shrub communities, and the arbor vitae association. The arbor vitae growing in this soil occurs in small isolated groves. There are numerous small springs In these areas which provide a suit­ able habitat for arbor vitae. I believe the natural vegetation for this soil Is some type of swamp forest.

The following is a typical profile of Linwood muck taken in Champaign County as recorded in Ritchie et al. (1971: 59-60).

1— 0 to 6 Inches, black (N 2/0), well-decomposed muck; very fine, weak, granular structure; very friable when moist; slightly acid; clear, smooth boundary. 1— 6 to 14 Inches, black (N 2/0), well-decomposed muck; strong, fine, angular blocky structure; firm when moist; slightly acid; clear, wavy boundary. 12

3— 14 to 22 inches, black (10YR 2/1) and yellowish-brown (10YR 5/8) muck; massive; firm when moist; calcareous; clear, smooth boundary. 4— 22 to 29 inches, dark grayish-brown (2.5Y 4/2) sedimentary' peat; common, medium, distinct, dark-brown (7.SYR 4/4) mottles; weak, thick, platy structure; friable when moist; some shells; calcareous; clear wavy boundary. IIC1— 29 to 38 inches, light olive-brown (2.5Y 5/4) silt loam; massive; friable when moist; a few shells; calcareous; gradual, irregular boundary.

Carlisle muck

Carlisle muck is a very poorly drained, slightly acid to neutral, and highly productive soil. It develops on 36 inches or more of mixed woody and grassy materials on flat to depressed bogs (Ritchie, 1961: 21). This is the most predominant soil at Cedar Bog. This soil is presently found in all but two of the vegetation zones at Cedar Bog. This, however, is not an accurate picture, because marl and peat are recorded as Carlisle muck. As mentioned before, all peat and marl deposits in this area are considered inclusions of the surrounding soil which is Carlisle muck.. It is my opinion that Carlisle muck is not the dominant substrate forming in the bog meadow, marl meadow or arbor vitae associations. The natural vegetation for Carlisle muck is probably some type of swamp forest.

The following is a typical profile of Carlisle muck taken in Champaign County as recorded by Ritchie et al. (1971: 49).

1— 0 to 7 inches, black (10YR 2/1), well-decomposed muck; moderate, fine, granular structure; friable when moist; slightly acidic; gradual, smooth boundary. 2— 7 to 13 Inches, black (N 2/0), well-decomposed muck that breaks out in chunks; firm when moist; slightly acidic; gradual, smooth boundary. 3— 13 to 38 inches, black (10YR 2/1), partly decomposed muck; friable when moist; some plant remains visible"; slightly acidic; gradual, smooth boundary. 4— 38 to 42 inches, black (10YR 2/1), and dark-brown (7.SYR 3/2), disintegrated peat; friable when moist; neutral; gradual, smooth boundary. 5— 42 to 72 inches, dark-brown (7.5YR 4/4), raw peat; .mas­ sive; a few fragments of wood; neutral. IIC— 72 to 100 inches, light olive-brown (2.5Y 5/4), compact silty-clay loam glacial till; calcareous.

Peat

Peat is not recorded in any surface soil survey for Champaign County by Ritchie et al. (1971). However, Dachnowski (1912: map) 13

records numerous deposits of peat.in Champaign County Including Cedar Bog. Dachnowski took numerous samples of peat in various unspecified parts of what is now called Cedar Bog. At that time Dachnowski recorded that there was about 600 acres of peat deposits in Champaign County.

Peat is an organic material consisting of an accumulation of plant remains in varying degrees of preservation or decomposition (U.S.D.A., 1938: 1129). If Individual plant species can be identified In the substrate, it is called peat. If the individual plant species cannot be identified, it is called muck (U.S.D.A., 1938: 1130). Peat is really the parent material for most muck soils.

The peat deposits predominate in the bog meadow and arbor vitae associations. In these two areas peat is now being formed. In both of these areas the water level is much too high most of the year to permit the organic material which is being deposited to decompose much. The soil forming processes have hardly begun here.

Considering the definition of peat, there Is no well defined line between a peat and a muck soil. This being the case, there will be an infinite variation from pure raw undecomposed organic matter to peat to well decomposed muck soil.

Many peat deposits are very acidic and low In minerals. However, at Cedar Bog the peat's natural acidity is partially neutralized by the water which is supersaturated with calcium carbonate.

The following peat profile was taken by Dachnowski (1912: 40) at Cedar Bog in Champaign County.

0 to 24 Inches, compact, well-decomposed, non-fibrous peat. 24 to 36 inches, dark-brown, somewhat gibrous, with a con­ siderable admixture of marl below; a number of well pre­ served logs and branches were found. 36 to 48 inches, brown and compact peat, fibrous with frag­ ments of rhizomes and roots. 48 and below, coarse gravel with stones showing glacial striatlons.

Marl

The marl deposits at Cedar Bog are frequently referred to as calcareous tufa (Dachnowski, 1912: 41). A calcareous tufa is a par­ ticular kind of calcium carbonate deposit originating by chemical deposition through springs, streams, caverns and waterfalls (Stout, 1940: 9). The water which flows from the springs at Cedar Bog Is supersaturated with calcium bicarbonate and calcium carbonate. When the water becomes exposed to the surface heat and evaporation occurs, a calcium carbonate deposit is left called calcareous tufa. Exposed TABLE 2

The basic characteristics of the soils found in the Cedar Bog Nature Preserve, Champaign County, Ohio

Lippincott Silty-clay Loam Linwood Muck Carlisle Muck Peat Calcareous Tufa

Naisral Drainage Very Poor Very Poor Very Poor Very Poor Very Poor

I Surface Silty-clay Loam Muck Muck Non-Fibrous, Fine Fine Grained (A Horizon) Grained, Compact Peat CaC03 DrlGT rrOTlX© Description Subsoil Silty-clay or Sedimentary Peat Muck and Peat Fibrous Peat

(B Horizon) Silt Loam *

Substratum Calcareous Stratified Silt Loam and Raw Peat and Calcareous Tufa or Glacial Outwash (C Horizon) Sand and Gravel Glacial Outwash Glacial Outwash Glacial Outwash

Normal Slope Range ($) 0 to 1 0 to 2 0 to 2 0 to 2 Level

Depth to Substratum (inches) 50 to 60 30 to 38 42 to 60 12 to 72 36

Depth to Bedrock (inches) Deep Deep Deep Deep Deep

Depth to Seasonal High Water 0 to 12 0 to 6 0 to 6 0 to 6 At or Above Table (inches) the Surface

Surface O.63 to 2.0 0.2 to O.63 O.63 to 6.3 - -

Permeability Subsoil O.63 to 2.0 0.2 to O.63 O.63 to 6.3 - - (inches/hour)

Substratum 6.3 to 12* O.63 to 2.0 -- - TABLE %• -Continued

Lippincott Silty-clay Loam Linwood Muck Carlisle Muck Peat Calcareous Tufa

4 Surface 0.l6 to 0.28 0.20 to 0,30 0.20 to O.3O a* -

Available Moisture Capacity Subsoil 0.06 to 0.11 0.20 to 0.30 0.20 to O.30 - (inches/inch of soil)

Substratum 0.05 0.l6 to 0.20 - - -

Surface Usually Neutral Usually Slightly Acidic Usually Slightly Acidic Acidic Alkaline 6.1 to 7*4 6.1 to 7.4 6.1 to 7.3

Reaction (pH) Subsoil 7.4 to 8.4 6.1 to 7*4 6.1 to 7.3 Acidic Alkaline

Substratum Calcareous Calcareous Calcareous Calcareous Calcareous

Surface Color Black Black Black Dark Brown Cream

Surface Organic Matter Very High Very High Very High Very High Very Low (Organic Soil) (Organic Soil)

Productivity Very High High High Variable Low Usually Low

Building Recaonendations Subsoil - Poor Surface and Subsoil Surface and Subsoil Unfit Unfit Substratum - Excellent Fair to Unfit Unfit

Extent In Champaign County ■% 6.4 0.6 1.0 0.2 (in 1912) ?

Acres 17.*? 1,797 2,719 600 (in 1912) Very Small

Sourest Alexander Ritchie Jr. (1961) and Alexander Ritchie Jr., Kenneth Powell and V. L. Siegenthaler (1971) 16 marl or calcareous tufa Is found -predominately in the marl meadow association. Here the water is almost always at or above the surface.

Environment Control Corporation Interpretation

The modified soil map based on Ritchie et al (1971) (Map II) and the soil map produced by the Environment Control Corporation (1973: B-7) (Map III) have very little in common. The Environment Control Corporation distinguished three soil groups at Cedar Bog, marly soils, loam and organic soils. I found it impossible to correlate all their soil core data and their well core data with their own soil map. The E.C.C. soil map (Map III) represented in this paper has been modified. I changed the scale to make It consistent with the other maps In this paper and I realigned the stream to make it consistent with the aerial photographs of Cedar Bog.

Loam

The E.C.C. loam best correlates with the U.S.D.A. Lippincott silty-clay loam. Environment Control Corporation (1973: IV-29) con­ siders this soil to be well drained, well-aerated with moderate to good stability. Normally, Lippincott silty-clay loam is a poorly drained and poorly aereated soil as evidenced by its gleying and mottling. At Cedar Bog this loam is found west of the West Branch at Cedar Run which has been dredged. This dredging has lowered effectively the water table In this soil. The typical soil profile presented by the U.S.D.A. probably does not accurately represent the Lippincott silty-clay loam at Cedar Bog today. The E.C.C. map (Map III) has different boundaries for this soil and consequently includes different vegetation zones. According to the E.C.C. (1973: IV-29) this soil is dominated by the entire mixed hardwood-beech forest, scattered short-grass sedge, scattered sycamore-basswood-cherry and scattered sycamore-maple- cottonwood segregants.

Organic Soils

The organic soils were subdivided Into muck, fibrous peat and woody peat. These soils are moderately to poorly drained, usually poorly aerated and relatively unstable. Map correlations between these organic soils and the Carlisle and Linwood mucks are not very good. I was not able to differentiate In the field the difference between woody and fibrous peat, therefore they are not differentiated on the map. Woody peat is formed under trees and shrubs and fibrous peat is formed under bog meadow or short-grass sedge associations. The muck is formed from the breakdown of these peats. Muck is usually occupied by some type of swamp forest. 1 rarely found any substrate which I could call peat in a swamp forest community. I did find that the peat deposits 17

m a p m

Soil map of Cedar Bog (modified from Environment Control Corporation, 1973: C -9).

Lippincott Silty-clay Loam

Muck

Fibrous Peat • > iii i

Woody Peat

M arl map in Soil map of Cedar Bog (modified from Environment Control Corporation* 1973: C-9).

!!!,

Wanilt.|j»l C n n r l

220 4 4 0 19 seem to correlate well with the bog meadow and the arbor vitae associ­ ations. There does not seem to be any particular organic soil and vegetation correlations with the E.C.C. interpretation. The distri­ bution patterns of the organic soils, whatever they are called, needs to be greatly efined at Cedar Bog.

The E.C.C. (1973: IV 41-42) also made some detailed analyses on several Thuja hummocks where peat deposits predominate. They listed eighteen correlations. Listed below are those correlations which apply to the substrates or soil conditions found in these hummocks.

1. The older the trees, the greater the accumulated leaf mulch was, particularly near the bases of the trees.

2. Peat deposits correlated with canopy area.

3. Soil stability increased with hummock size.

4. Hummock soils generally had a lower pH than the surround­ ing area.

5. Subsurface water temperatures were cooler in the hummocks than the surrounding area.

6. The upper layers of the hummock soils maintained aerobic conditions.

Marl

The E.C.C. divided the marly soils at Cedar Bog into two cate­ gories, organic marl and silty marl. They did not, however, differ­ entiate these two types on their map. They rated the drainage and stability as being moderate to poor. The two marl categories are apparently separable by the amounts of organic matter found in them. Their interpretation of marl seems very broad. It comprises a large area on the map (Map III). I found marl deposits to be sporadic everywhere except in the marl meadow association. Sometimes I would find interfaces which could be termed marly-muck or mucky-marl. It appears the E.C.C. called these intermediate areas marl. CLIMATOLOGY OF CEDAR BOG

All the climate information in this summary was taken directly from Frederick (1964, 1967 and 1974) or the Environment Control Corpor­ ation (1973). Additional detailed information concerning local climatic conditions can be obtained from these studies.

Air Temperature

In a comparison of the air temperatures at the 5-foot height at Cedar Bog and the Urbana weather station, Frederick (1967: 34) found the bog meadow averaged 3.1°F. higher for the maximum temperatures and 3.3°F. lower for the minimum temperatures.

The arbor vitae areas had less extremes and more constant temper­ atures than any other area of the Bog. The minimum temperatures were higher in the winter and maximum temperatures were lower In the arbor vitae area than in the bog meadow. The temperature variation was greater in the swamp forest than the arbor vitae forestand less than the bog meadow (Frederick, 1967: 39).

Frederick (1967: 41-42) also compared the number of frost-free days for Cedar Bog with those of the Urbana weather station. She found that the frost-free period in Cedar Bog was 2 1/2 months shorter than the surrounding areas. July was the only consistently frost-free month during her study,

Frederick (1967: 43) states, "There can be no question that the microclimate in Cedar Bog Is much more severe than in adjoining areas. Surely this microclimate has been an importantfactor in the survival of a boreal bog in this part of Ohio."

Soil Temperature

Frederick (1974: 69) states, "The soil temperatures never went below 32°F. in the bog meadow and swamp forest areas. In contrast, during 17 weeks in 1966 and 1967, the soil in the arbor vitae tract was at or below freezing. During the six warmest months, however, soil temperatures in the bog meadow averaged 3.1°F. higher, and in the swamp forest 2.2°F. higher, than those in the arbor bitae stand. One factor in maintaining a boreal habitat may be the effect on soil temperature resulting from the presence of arbor vitae in the area." The average minimum and average maximum soil temperature range for the bog meadow

20 21 was 36.3°F. to 64.1°F.; for the arbor vitae It was 32.2°F. to 61.0°F.; and for the swamp forest it was 38.2°F. to 63.0°F.

Humidity

Environmental Control Corporation (1973: II-7) states, "Humidities were contently high over Cedar Bog's discharge area. The high relative humidities and dense night fogs indicate that evaporation is in equilib­ rium with condensation in the form of dew, that the net water removal from Cedar Bog during the night is negligible, and that effective evaporation occurs only during the day."

Temperature Inversions

Gilbert and Hennen (1973: 14-15) reported that temperature inver­ sions occurred each day within the swamp and adjacent old-field. These temperature Inversions usually occurred from mid-morning to late afternoon and 23% of the time over the swamp habitats. "The signifi­ cance of inversions lies in the lack of vertical mixing of air across the interface between upper and lower air layers." (E.C.C., 1973: II-9) This would cause any pollutants, water vapor and heat to become trapped in the air layer near the ground. CEDAR RUN

General Description

There are two branches of Cedar Run which pass through the Cedar Bog Nature Preserve. The West Branch is approximately 5900 feet long and passes completely through the hardwoods and swamp forest on the west side of the study area. The East Branch is approximately 3200 feet long and passes through most of the arbor vitae, bog meadow and marl meadow associations. From the confluence of the two branches to the end of the Cedar Bog Nature Preserve, Cedar Run is approximately 1000 feet long. From the confluence of the two branches to its termination at the Mad River, Cedar Run is approximately 4.7 miles long. The West Branch drains about .66 sq. miles (422.4 acres) which is about 22% of the Cedar Run drainage basin. The East Branch drains .44 sq. miles (281 acres) which is about 15% of the Cedar Run drainage basin (E.C.C., 1971: Table C-l).

The West Branch was dredged about 5 feet deep north toward Hickory Grove Road around 1912 (E.C.C., 1971: 111-13). The West Branch is now a very straight stream and heavily shaded within the Bog by deciduous hardwoods. Williams (1966: 18-19) found it be 61 inches wide and 2.5 inches deep at the Woodburn Road bridge, with a flow rate of .68 cubic feet per second (cfs). This stream has a large amount of aquatic vegetation dominated by Nasturtium, Potamogeton, Sagittaria and Alisraa. The bottom substrate is a thick layer of silt and decay­ ing vegetation over gravel.

The East Branch has never been dredged and still meanders natur­ ally. It is shaded only where it passes through arbor vitae groves. Williams (1966: 16-18) found it to be about 62 inches wide and 6.5 inches deep in a run and 3 inches deep in a riffle at the Woodburn Road bridge, with a flow rate of .73 cfs. The stream has a variable sub­ strate which ranges from silt and decaying organic matter, to marl, to rounded gravel at the bridge. The aquatic vegetation is spotty and Is dominated by Spirogyra in the headwaters and Chara, Cladophora and Batrachospermum at the bridge.

The E.C.C. (1973: Table C-6) measured the flow rate of Cedar Run at various places and found the following averages.

East Branch above the McDaniel pond o u t l e t ™ . 21 cfs East Branch below the McDaniel pond outlet .67 cfs East Branch at the Woodburn Road bridge— — 1.33 cfs 22 23

West Branch at the north end of the Bog------.09 cfs West Branch at the Woodburn Road bridge 1.14 cfs Cedar Run below the confluence — — — ---2.81 cfs

Cedar Run was dredged in 1912 from the Mad River to about 50 yards south of the confluence of the east and west branches to a depth of 10 feet. This dredging lowered the water table of the entire drainage basin including the Bog. The rate of flow probably also in­ creased (E.C.C. , 1973: 111-13). The E.C.C. (1973: 111-25) believes that this dredging has decreased the stability of the soil and vegeta­ tion in the Bog. This instability is characterized by banked hummocks, small meander cuts, fallen mature trees and bare marly soil. Probably many of these phenomena are natural, but the rate at which they occurred increased after the dredging.

Water Temperature

The seepage areas or springs at Cedar Bog continue to flow throughout the year even though ice may form at the surface. Cedar Run itself never freezes. A thin sheet of ice may form along the banks. Frederick (1974: 70-71) found that "The average monthly tem­ perature of the water in Cedar Run varied from a low of 38.6°F in January to a high of 63.3°F in August. Monthly deviation was greatest from August to September, with a drop in the monthly average of 8.7°F. However, deviation from month to month through the remainder of the year averaged only 3.6°F. The highest water temperature recorded during 1966 was 73,0°F in July and the lowest was 26.0°F in January." She found that in 1965 the temperature extremes were 22°F and 78°F but, even at 22°F the water did not freeze. Large quantities of calcium carbonate and other salts along with a constant flow of water kept the water from freezing (Frederick, 1974: 73). All the tempera­ tures given for Cedar Run by Williams (1966) and the E.C.C. (1973) fall within the ranges given by Frederick.

Chemistry

Detailed information concerning the chemistry, pH, water flow and other physical parameters on both branches of Cedar Run can be found in Frederick (1964), Williams (1966) and E.C.C. (1973).

The pH of Cedar Run varies with location and temperature. E.C.C. (1973: 111-23) found the average pH of the West Branch of Cedar Run to ■ be 7.5, and the East Branch of Cedar Run to be 7.35. Williams (1966: 103) found the West Branch to have an average pH of 7.8 and the East Branch an average pH of 7.9. All the pH readings I have found for any part of Cedar Run fall between the pH range of 7.35 to 8.4.

One of the most important characteristics of the water flowing through Cedar Bog is the high concentrations of dissolved calcium and 24 magnesium carbonate. Environment Control Corporation (1973: 111-24) measured the calcium content as about 100 ppm and the magnesium about 40 ppm. The concentration of calcium carbonate in water is dependent upon the temperature and dissolved carbon dioxide. Temperature and carbon dioxide solubility are inversely proportional. As the temper­ ature rises the carbon dioxide concentration decreases and the dis­ solved calcium carbonate decreases. Environment Control Corporation (1973: 111-20) states that at Cedar Bog the

. . . supersaturation conditions for calcium carbonate are maintained in the cold groundwater. Apparently, the groundwater is almost immediately loaded with calcium carbonate which remains in solution as long as the temper­ ature of the water is low (12°C or 54°F) and pH is low (7.2). With discharge into the stream, precipitation occurs due to water temperature and pH increase. Stream water is saturated with calcium carbonate and is actively precipitating calcite as tufa.

The deposition of tufa is very important at Cedar Bog. Tufa deposited by the action of plant photosynthesis or by direct chemical action becomes mixed with organic matter. This mixture begins to form some characteristics of a primitive soil is called marl.

The E.C.C. also measured the hardness (carbonate-bicarbonate) of the Cedar Bog waters and found the ground water to have a range of 415 to 1680 ppm and surface water 378 to 476 ppm. All these readings indicate a supersaturated solution.

The dissolved sulfates in the Cedar Bog water are probably derived from gypsum and anhydrite in underlying deposits. The E.C.C. (1973: III-23) found quite a range from 8 to 1080 ppm. In the marl meadow the smell of hydrogen sulfide is sometimes very strong. This is due to the action of anaerobic bacteria on the sulfates and organic matter (E.C.C., 1973: 111-23). The high amount of sulfates present in the waters around the marl meadow could be concentrated there because of the Sphagnum growing there. Reid (1961: 194) mentions that Sphagnum accumulates sulfur from the atmosphere and releases it as sulfuric acid. The E.C.C. (1973: IV-38) found the marl areas to be more alkaline during wet periods and more acidic during dry periods. In dry periods, water stopped flowing in some areas and the soil water stagnated. Anaerobic bacteria can work on the sulfates and organic matter present to produce hydrogen sulfide which can directly or in­ directly lower the pH, In wet periods the downward movement of oxygen­ ated rain water increased soil aeration and stopped anaerobic activity. Rein (1961: 195) states that some other bacteria can work on the hydrogen sulfide to produce sulfur, water and energy. This same bacteria can release sulfuric acid by utilizing the sulfur originally produced, If the supply of hydrogen sulfide runs out. Reid also men­ tions a third group of bacteria which regularly produces sulfuric acid from elemental sulfur. Whenever sulfuric acid is released the pH of 25 the surrounding area is lowered. I do not know what bacteria are active in the sulfur cycle at Cedar Bog. It appears to me that the sulfur cycle is somewhat complicated in the marl areas of Cedar Bog. What effect this has on the plants and animals living there is unknown.

Environment Control Corporation (1973: 111-23) found the chloride concentrations to be 7 to 28 ppm and the nitrates to be 1 to 21 ppm. The nitrate concentration is high and they mention that It is probably due to fertilizer and sewage In the groundwater.

Williams (1966: 99-102) measured the dissolved oxygen In both branches and the main stream of Cedar Run. He found the headwaters of the East Branch had the greatest fluctuations from 57% to 95% of satur­ ation. The East Branch at Woodburn Road bridge varied from 70% to 90% saturation and the West Branch at Woodburn Road bridge varied from 77% to 90% saturation. The main stream near the Clark-Champaign County line south of Cedar Bog varied from 79% to 100% saturation, with most of the readings showing 100% saturated with oxygen. Cedar Run at the Clark-Champaign County line is open and has a great deal of submerged aquatic vegetation. This could account for the consistently higher dissolved oxygen readings in the waters there during daylight hours. VEGETATION OF CEDAR BOG

General Considerations

The floral composition of the Cedar Bog Nature Preserve Is very rich and varied. In an area of about 220 acres which makes up this Bog and some adjacent private property, 1162 plant taxa have been found by various workers (Table 1). The Bog has a unique assemblage of boreal, prairie and southern coastal plain plant species, many of which are absent or very rare elsewhere In Ohio. Frederick (1973: 16-20) summarizes and lists all the known disjunct plant species in Cedar Bog.

The boreal species present today in the Bog are relics of a wet and cool period which predominated after the recession of the Wiscon­ sin glacier. This conclusion is drawn from studies made by Braun (1928: 298-299), Sears (1930: 212-217 and 1932-: 4-5) and Ogden (1966: 394-395). About three to four thousand years after the glacier re­ treated northward the climate became much warmer for a while. This Is called the Xerothermic period (Sears, 1942: 708). During this period many western prairie species became established in parts of Ohio (Frederick, 1964: 13 and 1967: 21; Braun, 1928: 289). As the climate changed some of the wet prairie species continued to survive in the undisturbed wet areas like Cedar Bog (Braun, 1928: 299). The coastal plain species probably became established since the Xerothermic period, but no explanation has been found.

One of the primary reasons for initiating this study on the terrestrial molluscs at the Cedar Bog Nature Preserve was the exten­ sive vegetation information available from earlier studies. Frederick (1964 and 1967) made surveys of the vascular plants at Cedar Bog. Host of the vegetation descriptions presented are summarized from her studies. The Environment Control Corporation (1973) also made a study of the vegetation at Cedar Bog. Some of their interpretations differ significantly from Dr. Frederick*s.

The vegetation studies made by Dr. Frederick have taken place over a long period of time, her formal studies have taken place at least since 1962. Her published work indicates the use of qualitative methods. The E.C.C. studies took place during a five month period within one year. Their results were collected by both qualitative and quantitative techniques. Within this limited period of time they probably did not see all the annuals. They have also not witnessed the drastic changes, observed by Dr. Frederick, which have occurred at Cedar Bog in the last 20 years.

26 27

Frederick Vegetation Study

Frederick (1967) divided the study area into five plant associ­ ations, the arbor vitae, the bog meadow, the marl meadow, the swamp forest and the hardwood forest. The swamp forest and the hardwood associations were subdivided into seven segregants. Map IV shows the various vegetations zones (associations and segregants) as presented by Frederick (1974: 75-86). Tables 3 and 4 list some of the dominants found in these zones. I will summarize her descriptions concerning each of these vegetation zones.

Arbor Vitae Association

This association is dominated by northern white cedar (Thuja occidentalis L.). It Is frequently referred to as the white cedar association. This association occupied about 13 acres of 12% of the study area in 1967, north and south of Woodburn Road. There Is good evidence that the arbor vitae area is expanding and may be larger today.

Table 3 summarizes some of the descriptive text provided by Frederick (1964 and 1967) concerning various parts of the arbor vitae association In the Bog. Frederick (1967: 70-71) found some significant differences between the cedar stands north and south of Woodburn Road. The two cedars are generally smaller on the south side and have been subjected to more wind damage. Apparently, elm trees west of the cedar groves sheltered these trees from high winds until the elms died of the blight. In general, there are fewer herbaceous plant species in the arbor vitae south of the road. The development of an herbaceous layer in the various arbor vitae stands varies according to the age and density of the trees and also the water level of the stand (Frederick, 1964: 63).

Bog Meadow Association

The bog meadow is one of the relic plant communities within the Cedar Bog Nature Preserve. This community comprises about 15 acres or 13.3% of the study area. The bog meadow is characterized by many species of grasses, sedges and composites along with shrubby cinquefoil (Fotentllla fruticosa L.) and swamp Birch (Betula pumila L.) The water is always near the surface through the association and there are many small pools and streams. The substrate is primarily peat (fibrous or sedge-formed peat). Sometimes this peat Is mixed with muck or marl. This vegetation zone has the greatest diversity of plants in the Bog (Frederick, 1964: 53). There are good indications that white cedar is taking over various parts of this community by becoming established in the hummocks of grasses and sedges. 28

MAP IV Vegetation map of Cedar Bog (modified from Frederick, 1967: 50-53).

Arbor Vitae Association

-A' Bog Meadow Association

M arl Meadow Association

Swamp Forest Maple-Ash-Basswood Segregant

Swamp Forest Elm-Maple Segregant

Swamp Forest Tuliptree Segregant

Swamp Forest Silver-Red Maple Segregant

Shrub Communities iWifjii’siiUH

ij' ' 1 i r Hardwood Forest 11 tit i Oak-Maple Segregant ill I I 1*1

Hardwood Forest Beecii-Maple Segregant MAP IV Vegetation map of Cedar Bog (modified from Frederick, 1907: 50-53).

W»niHinfH Biml

220 440 TABLE 3

A summary of the arbor vitae association at Cedar Bog Nature Preserve, Champaign County, Ohio*

Hap Relative Age and Lichens, Liverworts, Mosses, Horsetails Some Dominant or Characteristic Herbaceous Remarks Coordinates Density and Ferns Plants, Shrubs and Small Trees

12 W Very Young; Even None Small Patches of Bog Meadow Plants Successional; Aged; Dense Betula pumila L. Patches of Bog Meadow Still Ainus ruqosa (DuRoi) Exist. Potentilla fructicosa L. Cornu3 sp.

10 W Young; Even Aged None None 10 X Dense 6 Y

9W Up to 14 inches Lycopodium lucidulun Michx. Li paris loeselii (L.) Richards Parts of These Stands Badly low DBH; Variable Dryopteris cristata Gray. Mitchclla rcperis L. Damaged By High Winds; 11 V, w Age; Dense Thelypteris palustris Schott. Andropooon qerardi Vitman Some Parts Are a Dense 12 V, W Alnus ruqosa (Du Roi) Tangle Now; Rubus pubescens Raf. Some Parts Are Drier Now Physocarpus opulifolius (L.) Maxim. Because Of Dredging; Taxus canadensis Marsh. Cornus sp.

6 X Variable Age; Equisetum arvense L, Habenaria clavellata (Michx.) 7 x Less Dense; Cystopteris bulbifera (L.) Bemh. Anemonelln thalictroides (L.) Spach, a x Osmunda rcqalis L. Hepatica americana (DC) Ker, Polypodiura virqinianun L. Viola sp. West of Camptosorus rhizophyllus (L.) Link. Mitella diphylla L. East Branch Trillium qrandiflorun (Michx.) Maianthemum canadense Desf. Hvdranqea arborescens L. Lindcra benzoin (L.) Blumo TABLE 3- -Continued

Hap Relative Age and Lichens, Liverworts, Mosses, Horsetails' Some Dominant or Characteristic Herbaceous Remarks Coordinates Density and Ferns Plants, Shrubs and Small Trees

Age Quite Equisetura arvense L, Cirsium muticurn Michx, Has Been Cut, Several Variable; Up to Cystopteris bulbifera (L.) Bemh. Parnassia glauca Raf, Large Stumps Remain; 20 inches DBH) Habenaria clavollata (Michx.) Some Plants Have Been Less Dense Mitchclla rcpens L, Introduced In This Area. Drosera rotundifolia L. Physocarpus opulifolius (L.) Maxim,

7 x, Y Mostly Old Trees; Conoccphalum sp. Maianthenum canadense Desf. Herbaceous Plants Are 8 X Up To 17 inches fin Low and Wet Areas) Hitchella repens L. Found Only In The Drier East of DBH; Less Dense. Arioacna atrorubens (Ait.) Blume Areas; The Shrubs Are East Branch Lichens and Mosses Aralia raceiaosa L. Along The Edge. (in Drier Areas) Viola sp. Cypripcdium reginae Walt. Cystopteris bulbifera (L.) Bernh, Rhus vcrnix L. (Only In Drier Areas) Rosa setigera Michx. Cornus sp. Xanthoxylum amcricanum Mill. Ptelea trifoliate L.

Sumarized from Frederick (1964 and 1967).

to H 3 2 TABLE 4

Species composition of the plant associations at the Cedar Bog Mature Preserve, Champaign County, Ohio

Plant Associations Some Characteristic Plant Species Some Common Associate Plants

Arbor vitae Thuja occidentalis L. See Table 3

Bog Meadow Andropoqon gerardi Vitman. Allium cernuum Roth. Rhynchospora alba (L.) Vahl. Zioadanus glaucus Nutt. Potentilla fruticosa L. Tofieldia glutinosa (Michx.) Pers. Sorqhastrum nutans (L.) Campanula aparinoides Pursh. Betula punila L. Gentiana crinita Froel. Thuia occidentalis L. Thelypteris palustria Schott. Many grasses, sedges and composites Lobelia kalmii L. Parnassia qlauca Raf. Cornus spp.

Marl Meadow Triqlochin maritima L. Calopoqon pulchellus (Saliab.) Eleocharis pauciflora (Lightf.) Cacalia tuberosa Nutt. Eleocharis rostellata Torr. Krioia biflora (Walt.) Blake. Utricularia cornuta Michx. Drcsera rotundifolia L. Helenium autumnale L. Camoanula aparinodes Pursh. Rhynchospora capillacea Torr. Potentilla fruticosa L. Carex spp. Juncus spp.

Swamp Forest Acer rubruin L. Carpinus caroliniana Walt. Fraxinus nigra Marsh. Quercus bicolor Willd. Maple-Ash-8asswood Tilia americana L. Liriodendron tulipifora L. Segregant Jualans cinceroa L. Platanus occidentalis L. Celtis occidentalis L. Lindera benzoin (L.) Blums Senecio aureus L. Symplocarpus foetidus (L.) Nutt. Caliha palustris L.

Swamp Forest Acer rubrum L. Liriodendron tulipifera L. lllmus rubra Muhl. Quercus imbricaria Michx. Elm-Maple Urtica dioica L. Quercus muehlenberqii Engelm. Segregant Cornua spp. Fraxinus niqra Marsh. Rubus spp. Platanus occidentalis L. Celtis occidentalis L. Prunus serotina Ehrh. Allaria officinalis Andrz. Ambrosia trifida L, 33 TABLE 4- -Continued

Plant Associations Some Characteristic Plant Species Some Common Associate Plants

Swamp Forest Ulmus rubra Muhl. Lindera benzoin (L.) Blume Fraxinus nigra Marsh. Taxus canadensis Marsh. Elro-Ash-Maple Acer rubrum L. Platanus occidentalis L. Segregant Acer saccharinum L. Rubus occidentalis L. Fraxinus pennsylvanica Marsh. Urtica dioica L. Fraxinus americana L. Laportea canadensis (L.) Wedd. Athyrium thelypteroides Desv. Allaria officinalis Andrz. Urtica procera Muhl.

Swamp Forest Acer rubrum L. Allaria officinalis Andrz. Acer saccharinum L. Polemonium reptans L. Silver— Red Maple Anemone quinquefolia L. Segregant Impatiens capensis Meerb. Thaspium barbinode (Michx.) Nutt. Osmorhiza lonqistylis (Torr.) DC. Viola spp.

Swamp Forest Liriodendron tulipifera L. Symplocarpus foetidus (L.) Nutt. Hepatica americana (DC.) Ker. Anemonella thalictroides (L.) Spach. Oxypolis riqidior (L.) C. & R. Eupatorium ruqosum Houtt. Carpinus caroliniana Walt. Lindera benzoin (L.^ Blume

Hardwood Forest Faqus qrandifolia Ehrh. Carya ovata (mill.) K. Koch Acer saccharum Marsh. Quercus bicolor Willd. Beech-Maple Quercus muehlenqerqii Enqelm. Segregant Quercus schumardii Buckl. Acer rubrum L. Ulmus rubra Muhl. Celtis occidentalis L. Aesculus qlabra Willd. Galium triflorum Michx.

Hardwood Forest Quercus bicolor Willd. Quercus borealis Michx. Quercus muehlenqerqii Enqelm. Quercus imbricaria Michx. Oak-Maple Acer rubrum L. Juglans cinerea L. Segregant Acer saccharinum L. Prunus serotina Ehrh. Many Hybrid Quercus Fraxinus pennsylvanica Marsh. Fraxinus niqra Marsh. Carpinus caroliniana Walt.

Shrub Community Rhus vernix L. Rosa setiqera Michx. Swamp Forest and Xanthoxylum americanum Mill. Bog Meadow Ecotone Ptelea trifoliata L. Cornus spp. 34 TABLE 4— —Continued

Plant Associations Some Characteristic Plant Species Some Common Associate Plants

Shrub Community Salix discolor Muhl. Lindera benzoin (L.) Blume Betula pumila L. Sambucus canadensis L« Arbor vitae and Rhannus alnifolia L'Hex. Cornus spp. Marl Meadow Ecotone Rubus allcqhenienais Port. Gaylussacia baccata (Want.) K. Koch. Rhus vernix L.

Shrub Community Rubus occidentalis L.

Swamp Forest North of Woodburn Road

Shrub Community Lindera benzoin (L.) Blume Thalictrum polyoamum Muhl. Rubus pubescens Raf. Blephilia hirsute (Pursh.) Benth. Swamp Forest South Cornus alternifolia L. of Woodburn Road

Source; Summarized from Frederick {19^4 , 19^7 Qnc* 1 3 7 4). 35

Marl Meadow Association

The marl meadow association makes up about 5 acres or 4.6% of the Bog. It is characterized by a substrate of marl where the water is always at or above the surface. The dominant plants are grasses and sedges. Frederick (1964: 69) found the appearance of this community changes with the seasons and what appeared dominant at one time did not appear so at another time. The apparent seasonal changes are caused by the different flowering times of the many common plant species found in the marl meadow.

Swamp Forest Association

The entire swamp forest association occupies about 43 acres or 38% of the study area. This is the largest association in the Bog. Assuming there are no major changes of water level or species composi­ tion, this association should become larger and more mature.

Originally, the swamp forests at Cedar Bog were dominated by American elm (Ulmus americana L.). Since 1962 these large canopy dominants have all died due to the Dutch elm disease (Frederick, 1967: 72). There has been a great deal of change in the swamp forest associ­ ation since the elms died. These changes have not been uniform through­ out the Bog. Now most of the canopy dominants are red elm (Ulmus rubra Muhl.), red maple (Acer rubrum L.) and black ash (Fraxinus nigra Marsh.).

In some areas where American elm had flew canopy associates, drastic changes occurred after the elm died. In the swamp forest segregant called elm-maple by Frederick and savana by E.C.C., the area left the few canopy associates such as red maple, sycamore, cottonwood and red elm standing isolated. Dense stands of grasses, ragweed, nettles and shrubs filled in the open spaces.

Prior to 1962 the swamp forest south of Woodburn Road (Map IV, T,U,V,-10, 11, 12) was probably dominated entirely by American elm. When the elm died it became a dense stand of shrubs dominated by spicebush, blackberry and dogwood. Now this same area is beginning to differentiate and various young swamp forest dominants are becoming established over the shrubs in some areas (V-10,11, 12). This part of the shrub community is now called elm-ash-maple segregant by both Frederick (1974) and E.C.C. (1973).

When Frederick visited Cedar Bog in the 1940's, she found a few trees and a lot of grasses in what is now a silver-red maple segregant (V-9, 10). This stand has formed in the last 30 years. This segregant is a young dense stand which shows little diversity now. Eventually this segregant is expected to become a mature swamp forest and join with the elm-ash-maple swamp forest developing further south. 36

The swamp forest segregant Immediately north of Woodburn Road has changed since the American elm died, but not drastically. Apparently there was more diversity in the canopy at the outset. When the elms died it left fewer openings and grasses and shrubs never "took over." This swamp forest segregant is also very wet, being flooded in many areas in the early spring. This segregant is now called maple-ash- basswood by Frederick (1974) and tuliptree-basswood-ash by E.C.C. (1973). The basic difference lies in the recognition of tuliptree as a separate segregant by Frederick.

The tuliptree segregant as recognized by Frederick (1967 and 1974) is most distinct in several areas of the Bog. Tuliptree is found throughout the swamp forest association, but in several areas it exists in almost pure stands. I believe that its separation as a segregant is justified. This area is very wet in the spring, and I have seen standing water over one foot deep in April. The soil is frequently a dense packed muck when dry and has almost no accumulated leaf litter. The buttressed trunks and the long exposed roots of the tuliptrees makes this area difficult for both walking and sampling.

Hardwood Forest Association

Frederick (1967: 85) divided the hardwood forest association in two segregants at Cedar Bog, the beech-maple and the oak-maple.

The beech-maple segregant comprises about 14 acres or 12% of the study area. This forest is dominated by American beech (Fagus grandifolia Ehrh.) and young sugar maple (Acer saccharum Marsh.). In 1967 Frederick called this an oak-hickory-beech-maple forest. This beech-maple does have many oaks and hickories in the canopy. The shrub level is indistinct within the forest, but it is well developed around the edges bordering the open fields. The sub-canopy is dom­ inated by young sugar maple, buckeyes and oaks. The herbaceous level is also well developed with good diversity.

The oak-maple segregant Is found only along the west bank of the West Branch. It comprises about 4.5 acres of 4.0% of the study area. This segregant is characterized by widely separated mature hardwoods, a well developed shrub level and a poorly developed herbaceous level. In spite of the fact that a swamp forest is across the stream, this segregant does not have many of the typical swamp forest species. This narrow band is somewhat drier than the swamp forest, but still has numerous springs and seepage areas. In a few places white cedar occupies the seepage areas. This entire segregant has been greatly disturbed by dredging and the clearing of the adjacent field.

Shrub Communities

Shrubs usually form an Intermediate community between arbor vitae, deciduous forests, bog meadow and marl meadow associations (Frederick, 37

1967: 58). There are several different areas at Cedar Bog where shrubs dominate. They are not always composed of the same species (Table A). Some shrub communities are successional and others are more ecotonal. In some cases shrubs (Lindera benzoin, Rubus pubescenB, Rubus occidentalis and Cornus alternifolia) became the dominant vegeta­ tion forms after the canopy was destroyed by wind or disease (Table A). Once the canopy is restored many of these shrubs will die out. In other cases shrubs form ecotones between one distinct association and another. Ecotonal shrub communities are never static because of con­ stant changes of the associations which they border. The various shrub communities dominated about 16 acres or 15% of the study area at Cedar Bog in 1967. This has undoubtedly changed in several areas now.

Environment Control Corporation Vegetation Survey

The Environment Control Corporation (1973: IV-11) reported and mapped (Map V) five distinct plant communities, the short grass-sedge, the tall grass-herb, the savanna-, the evergreen and evergreen-mixed deciduous, and the deciduous. Some of these communities are subdivided into segregants.

Evergreen and Evergreen-Mixed Community

The evergreen and evergreen-mixed community was divided into three segregants, the Thu ja occldentalis-mlxed hardwoods, the Thuja occidentalis-marginal, and the Thuja occidentalis-hummock (E.C.C., 1973: IV-13). These three segregants are not separated on the map in this paper (Map V). The Thuja-mixed hardwood is the most prevalent segregant of this community. It is composed of the Thuj a stands along the eastern edge of the deciduous forest and along the East Branch and main stream of Cedar Run. The Thuja-marginal segregant is the narrow white cedar area south of Woodburn Road and east of Cedar Run (Map V, W-10, W-12, X-9 & 10). The Thu.ja-hummock segregant is composed of the isolated white cedar groves within the marl short grass-sedge community. Some of these are indicated on the vegetation map (Map V). All the segregants of the evergreen and evergreen-mixed community are the same as the arbor vitae association of Frederick (196A, 1967 and 197A).

The Thuj a-mixed hardwoods contain a substantial number of deciduous tree species (Table 5). The Thu.j a-marginal is 95% Thuja with a few deciduous trees (E.C.C., 1973: IV-13). "Frequency of occurrence of hardwoods tended to decrease as the age of the stand increased; however, older stands were commonly over-topped by deciduous members, such as Acer rubrum, Liriodendron tulipifera, Ulmus rubra, and Platanus occidentalis" (E.C.C., 1973: IV-13-1A). The Thuja-hummock communities are almost pure stands of Thuj a occidentalis. All the evergreen and evergreen-mixed communities have a large number of relic species (Table 5). 38

MAP V Vegetation map of Cedar Bog (modified from Environment Control Corporation, 1973: B-2 and B-3).

Marl Short Grass-Sedge

Non-Marl Short Grass-Sedge

Tall Gras s-Herb

Savanna Sycamore-Bass woo d - Cherry

Savanna Sycamore -Maple -Cottonwood

Evergreen and Evergreen-Mixed

Deciduous Forest Tuliptree -Basswood-Ash

Deciduous Forest Maple-Elm #1

Deciduous Forest Maple-Elm #2 \

Deciduous Forest Maple -Ash -Elm

Deciduous Forest Mixed-Beech - MAP V Vegetation map of Cedar Bog (modified from Environment Control Corporation, 1973: B-2 and B-3).

» I 5 I t ~l U 1 V 1 W \ X \ Y | Z

t

3

3

4

5

«

7

I Wocrfhmn. tla a iL

9

10

n

13

» ... I » I T t U I V 1 W | K | V | 1 40 TABLE 5

Species composition and relic plant diversity in the plant associations at the Cedar Bog Nature Preserve, Champaign County, Ohio as interpreted by the Environment Control Corporation.*

Plant Associations Dominant Plants Number of Number of Number of Northern Prairie Southern Components Components Components

Marl Triqlochin maritime 21 6 3 Short Grass-Sedge Utricularia cornuta Eleocharis spp. Carex spp. Juncus spp. Potentilia fruticosa Betula pumila Rhynchospora capillacea

Non-Marl Cirsium arvense 4 3 l Short Grass—Sedge Solonum nigrum Ambrosia artemisiifolift Festuca elatior

Tall Grass-Herb Andropogon qerardi 5 2 l Ambrosia trifida Urtica dioica

Savanna Platanus occidentalis 7 2 l Tilia americana Sycamore-Basswood— Prunus serotina Cherry Segregant Andropoqon qerardi Ambrosia trifida Urtica dioica

Savanna Platanus occidentalis 7 1 l Acer rubrum Sycamore— Mapie- Populus deltoides Cottonwood Segregant Andropoqon qerardi . Ambrosia trifida Urtica dioica

Evergreen and Thuja occidentalis 13 1 1 Evergreen-Mixed Substantial Numbers of Hardwoods Hardwood Acer rubrum Liriodendron tulipifera Ulmus rubra Platanus occidentalis Fraxinus nigra Lindera benzoin 41 TABLE 5 “ -Continuod

Plant Associations Dominant Plants Number of Number of Number of Northern Prairio Southern Components Components Components

Evergreen and Thuja occidentalis l6 5 2 Evergreen-Mixed Infrequent Occurence of Hardwoods Hardwood Acer rubrum Liriodendron tulipifera Marginal Ulmus rubra Fraxinus niqra Platanus occidentalis

Evergreen and Thuia occidentalis 12 5 2 Evergreen-Mixed Hardwood

Hummock

Deciduous Hardwood Liriodendron tulipifera B l 0 Tilia americana Tuliptree-Basswood- Fraxinus niqra Ash Segregant Acer rubrum Celtis occidentalis Prunus serotina Quercus bicolor Ulmus rubra Platanus occidentalis

Deciduous Hardwood Acer rubrum 3 0 1 Ulmus rubra Maple-Elm Site #1 Lindera benzoin Segregant Prunu3 serotina Celtis occidentalis

Deciduous Hardwood Acer rubrum 1 0 0 Ulmus rubra • Maplo-Elm Site #2 Prunu3 serotina Lindera benzoin Sanbucus canadensis Morus spp.

Deciduous Hardwood Acer rubrum 11 0 1 Fraxinus niqra Maple-Asb-Elm Ulmus rubra Segregant Quercus bicolor Celtis occidentalis Liriodendron tulipifera Acer saccharum Lindera benzoin 42 TABLE -Continued

Plant Associations Dominant Plants Number of Number of Number of Northern Prairie Southern . Components Components Components

Deciduous Hardwood Faqus qrandifolia 5 0 0 Tilia americana Hixed-Beech Acer rubrum Segregant Aesculus glabra Acer saccharum Prunus serotina Quercus borealis Quercus bicolor Carya ovata

* Summarized from Environment Control Corporation (1973s IV—11 to l 6 and Tables B-3 to S—7 J• 43

Short Grass-Sedge Community

Within the study area the short grass-sedge community was found on all marly soils and grazed and cultivated areas on humus. The E.C.C. distinguished between the marl and the non-marl short grass- sedge areas on their map (Map V). A part of this marl short grass- sedge community correlates reasonably well with what Dr. Frederick calls the marl meadow. The non-marl short grass-sedge community is part of the bog meadow association of Frederick.

In addition to the large number of relic species found in the marl short grass-sedge community (Table 5), there is a rich assemblage of other plants, many of which are rare in Ohio. The non-marl short grass-sedge community is dominated by grasses and herbs characteristic of agricultural disturbances (E.C.C., 1973: IV-19). With such differ­ ences I find it difficult to place them in the same short grass-sedge community. The non-marl segregant appears to be the result of man's activities in the area and not a natural association.

Tall Grass-Herb Community

The tall grass-herb community is characterized by big bluestem (Andropogon gerardi), great ragweed (Ambrosia trifida) and stinging nettle (Urtica dioica). This community as indicated by the E.C.C. is found in a very small part of the study area (X-8). It was Included as a part of the bog meadow association by Frederick (1964). Table 5 compares the number of relic components in this community with other communities. "The tall grass-herb community in the Bog is not uniform in composition, and it is difficult to assess the relative Importance of species present" (E.C.C., 1973: IV-20).

Savanna Community

The savanna is separated In two segregants, the sycamore-basswood- cherry (Platanus occidentalis-Tilia americana-Prunus serotina) segregant and the sycamore-maple cottonwood (Platanus occidentalis-Acer rubrum- Populus deltoides) segregant. The sycamore-basswood-cherry segregant is the largest segregant in the study. The sycamore-maple-cottonwood is represented by a small section west of the West Branch of Cedar Run (V-4,5).

These segregants are termed savannas by the E.C.C. because of the open areas between the trees which are dominated by big bluestem, stinging nettle and great ragweed. These plants are the characteris­ tic components of the tall grass-herb community. Frederick (1967) designated most of the savanna segregants as part of the elm-maple swamp forest. I am not sure what this area should be called. There does not appear to be enough living trees now to call it a forest and with the dense growth of grasses and herbaceous plants between the trees the term 44 savanna may be Che most appropriate. Eventually this area is expected to become a part of the swamp forest association.

Deciduous Communities

"Five segregant forest types are identified— Liriodendron tulipifera-Tilia americana-Fraxinus nigra, Acer rubrum-Ulmus rubra Site 1, Acer rubrum-Ulmus rubra Site 2, Acer rubrum-Fraxinus nigra- Ulmus rubra and Mixed-Fagus grandifolia" (E.C.C., 1973-: IV-14).

The tuliptree-basswood-ash (Liriodendron-Tilia-Fraxinus) stands are fairly dense and have a rich diversity of canopy species (Table 5). The small tree and shrub levels are well developed. This deciduous segregant is generally what Frederick (1974) called maple-ash-basswood. The E.C.C. includes the tuliptree zone to the east and the oak-maple zone on the west bank of the West Branch of Cedar Run as a part of this segregant, Frederick does not.

The Acer Ulmus Site 1 is a young dense stand of trees with very low canopy diversity. This community is what Frederick (1974) re­ ferred to as the silver maple-red maple swamp forest segregant. The E.C.C. (1973: IV-12) refers to all the maple in this segregant as Acer rubrum. They concluded that these trees may be hybrids of Acer rubrum and Acer saccharinum, but none were Acer saccharinum. Frederick (1967 and 1974) recognizes both species and no hybrids.

The Acer-Ulmus Site 2 is a young community, similar to Acer-Ulmus Site 1, but less dense and with a greater canopy diversity. Frederick (1974) basically agrees with the E.C.C. on this segregant.

The maple-ash-elm (Acer-Fraxinus-Ulmus) segregant was observed by the E.C.C. (1973: IV-15) to be a diverse and relatively mature area. They listed several other associate species in the canopy (Table 5). This community has a dense shrub zone dominated by Lindera benzoin (spicebush) which is common throughout. Frederick (1967) subdivided this same area four ways. Where tuliptree dominates the canopy she called that a tuliptree segregant, where the oaks and maples dominated the canopy she made that part of the beech-maple segregant; where the shrubs dominated in the center, the shrub zone; and where the white cedar dominates along the eastern edge, the arbor vitae community. Frederick (1974: 83) mentions that the eastern part of the shrub com­ munity is a swamp forest dominated by elm, ash and maple. Some of the differences between these two studies can be attributed to time. The E.C.C. study occurred several years after Frederick's study and changes could have occurred. Some of the fast growing maples could change the aspect of the area significantly.

The mlxed-Fagus segregant is considered a mature stand. Several species of trees are canopy associates. Frederick (1967) originally 45

called this the oak-hickory~beech-maple community which indicated this diversity. She now calls this a beech-maple segregant.

Vegetational Changes

Through the previous text I have mentioned various vegetational changes which have taken place or possibly will take place at Cedar Bog. Some of these changes have been closely examined by the Environment • Control Corporation and Dr. Frederick. The arbor vitae association is expanding into the bog meadow and marl meadow associations. The swamp forest has changed drastically since the American elm died. Also, in several areas open fields are well on their way to becoming forested again. Clues to these changes are evident throughout the Bog. How­ ever, the most startling evidence of change is revealed by comparing aerial photographs of Cedar Bog published by the Clark County Audubon Society (1972: 17). They reproduced two aerial photographs of Cedar Bog, a 1938 aerial map and a 1968 aerial map. I could not differenti­ ate many specific vegetation types, but it is not difficult to tell which areas were heavily forested, lightly forested or open in 1938. Map VI shows the forest coverage as I interpreted it from the 1938 aerial photograph. The significance of these changes, as far as terrestrial molluscs are concerned, may not be fully understood. Con­ sidering the known habitats of many species these changes probably have had a significant effect on their distribution and density at the Bog. MAP VI Vegetative cover of Cedar Dog in 1938 (Adapted from a 1938 aerial photograph published by the Clark County Audubon Society, 1972: 17).

Forested

Scattered Trees

Open 47

MAP VI Vegetative cover of Cedar Dog in 1938 (Adapted from a 1938 aerial photograph published by the Clark Couuty Audubon Society, 1972: 17).

■ I s | t ) u | v j w | > | r | I

W a adhitia_8ocd.

1cm. o330ll.

V T 'I "x j Q I V "w~ | X | T | I METHODS AND MATERIALS

The first collections made at Cedar Bog were random and not according to any particular plan. The objective was to determine the feasibility of studying molluscs at the Bog. My initial plans were to investigate only the molluscs found In the relic plant communities studied by Frederick (1964). Therefore, most of the early collections were taken in the arbor vitae, bog meadow and marl meadow associations. Many of these early collections are not mapped. A map for the Bog did not exist at the time and poor field notes did not help matters later when I did have a map. The only accurate locality data known about each collection site was which vegetation zone it had been made in.

The molluscs were collected In three principal ways. One col­ lecting method which Involved lifting logs and loose bark, and search­ ing any available habitat in the field Is called a random collection. A random collection could only be mapped if I concentrated ray efforts In a relatively small area having a defined vegetation. However, some random collections were made while simply walking through a particular vegetation zone and all specimens were lumped into one lot. These collections could not be pinpointed on any map. Some of the early collections are called qualitative litter samples. These were random collections of soil and leaf litter which were taken back to the labor­ atory for processing. Again, some of these samples could be mapped and some could not. The third type of collection was a quantitative litter sample. These varied in size from one square foot to one square meter. All these samples were mapped accurately.

In order to make a reasonably accurate comparison of the mol- luscan diversity among the various vegetation zones, some type of standard collecting procedure had to be used. In my search of the literature I found that a standard procedure for collecting molluscs has not been established. Jacot (1935) made one of the first ecologi­ cal studies devoted to molluscs which used a quantitative technique. Jacot used a cylinder sharpened on one side to take shallow cores of leaf litter and soil. He took three cores per location which totaled about one square foot. Several other Investigators used quantitative sampling methods to obtain various terrestrial invertebrates. Blake (1931) and Smith (1928) studied all the invertebrates in the soil and litter. They made two square feet samples ten centimeters deep. Foster (1937) and Strandine (1941) used one square meter quadrats to study terrestrial snails, but they studied only the large species and hand- picked them in the field. I tried all these methods. Samples larger than one square foot were not practical. They were too large to process efficiently and did not allow enough time in the field to sample

48 49 a highly variable substrate. All. the quadrats which were hand-picked in the field were only reasonably accurate with regard to the large species. I could not accurately count any species of mollusc in the field if it were smaller than 8 millimeters. This completely eliminated most of the diversity. As a result of several field trials 1 decided that one square foot quadrats were as large as I could handle efficiently.

I used a piece of galvanized metal formed into a tube one foot high having a cross-sectional area of one square foot. The top edge of this tube was covered with a section of garden hose-. This pro­ tected my hands from the sharp metal edges. In the field I would rotate this coring device and I could usually cut through small sticks and roots. I tried to take each sample four Inches deep. Due to the variable nature of the substrates involved I had to alter this at times. Sometimes large roots would block the way and the soil and litter had to be dug out from around the roots. This reduced the depth accuracy. In the winter collections the soil was frequently frozen to a depth of six inches. I had to sample all frozen substrates down to an inch below the frozen layer. The amount of soil and organic debris collected in each sample varied considerably. The idea behind this procedure was to collect all the living molluscs in one square foot. I believe I accomplished that. Table 6 lists all the collec­ tions made at Cedar Bog during this study. It also records where and when each collection was made.

The question as to how many quadrats should be collected in each vegetation zone was not easily answered. Likhachev and Rammel'meier (1952: 88) state that large mollusks should be collected or counted in quadrats of 4 to 16 m^ per sample area and small mollusks in leaf and soil samples should be collected in 3 to 4 quadrats of 1/8 to 1/4 each. This, they said, would allow the investigator to subject the data to a statistical analysis. In non-metric terms the soil quadrats would cover an area which varied from 4 ft^ to 11 ft^ per sample area. Krutzsch (1952) collected five square feet of soil and litter per sample area. Jacot (1935) collected one square foot per sample area. I sub­ mitted my preliminary data to several graduate students participating in a Statistics Department seminar. They arrived at a figure of 5 one square foot quadrats per sample area by methods unknown to me. I felt that since this figure was within the range of the recommendations made by Likhachev and Rammel’meier I would start with 5 one square foot samples per vegetation zone.

Blake (1931) found the number of soil and litter inhabiting snails fluctuated with the season. There apparently was no one time which would be best to sample all possible species present in the soil and litter. It was clear that I had to collect samples from all vegetation zones several times a year. Initially, I decided to collect 5 one square foot samples of litter and soil from each vegetation zone at Cedar Bog four times in one year. Since Frederick (1967) defined ten distinct vegetation zones at Cedar Bog, this meant that 1 would collect 50 quadrats four times or 200 ft^. 50

I was strongly advised by several people that each sample be taken with as little bias as possible. I felt that by using Dr. Frederick’s original vegetation map I could reliably find in the field, within-a radius of 25. feet, any specific point marked on the map. On this map 1/8 inch equaled 50 feet. I took 1/8 translucent graph paper and * traced all the vegetation boundaries. 1 numbered both coordinates and cut equal size pieces of paper to represent each 1/8 mark along the two coordinates. I put all the numbers representing each coordinate in . separate jars. I then selected a number from each jar. These two num­ bers would be the coordinants for a point on the map. I repeated this until I had five points in each of the ten vegetation zones in the Bog. I repeated this whole process four times to select quadrat sites for the four different collection times.

In the field I attempted to find each preselected collection site on the map by using vegetation zone boundaries, known trails and the streams as landmarks. Most of the time 1 had to locate a point from more than one direction. I started my quantitative collecting in December of 1968 because it would be the easiest time to get around the Bog and spot the various landmarks.

. Once I had located each preselected point on the map, I then had to determine exactly where the quadrat sample would be taken. This was accomplished by rolling a die. On the first roll, one meant 5 feet north, two meant 10 feet north, three meant 15 feet north, four meant 5 feet south, five meant 10 feet south and six meant 15 feet south. The second roll would determine the east or west distances. After each direction was selected I marked off the distance and took the sample. Naturally, several factors interfered with this procedure. If the end point was in the middle of a tree or a log I would select the nearest side as close as possible to the three or log. Also, if by marking off the distance for each point, I moved out of the vegeta­ tion zone in which the sample was intended to be taken, I rolled the die a second time.

At the same time each quadrat was being taken I recorded the temperature of the soil at a depth of six inches, the .substrate sur­ face and the air at 6 feet above the substrate. I also recorded the relative humidity at ground level and at six feet. The general nature of the substrate was also recorded.

All the soil and litter samples were placed in large plastic bags, labeled on the outside and stored until they could be dried out. The samples were brought to the laboratory and dried out by placing them in large open plastic trays under incandescent lights. The soil and litter was periodically stirred up and all large pieces of bark, rotted wood and leaves were broken apart to facilitate drying. Once the sample was dry, it was stored in plastic boxes or jars with the appro­ priate label until it could be processed further. 51

The dried samples were, sorted by means of a series of four screens, 1/4 inch, 1/8 inch, 1/16 inch and 1/32 inch. The first two separations were searched carefully without any magnifying aids, the third separa­ tion was searched with the aid of a lighted magnifying lens and the fourth separation was searched under a 10X binocular dissecting micro­ scope. The material which passed through the last screen was not examined. This fine material was examined when I first started this study. I found that it takes as long to search this fine material as it does to process the rest of the sample. The results were not very rewarding. The few snails recovered were all protoconchs of the smallest species which were, in most cases, impossible to identify. I felt that the knowledge gained was not worth the time spent obtaining it.

The material I examined under the binocular microscope was put in a standard petri dish a tablespoon at a time and spread evenly over the bottom of the dish. I would scan the material in zig zag pattern across the dish, using a fine brush to separate clumps. The snails found in this separation were usually less than two millimeters in height. These very small molluscs are most practically handled by means of a wetted 000 camel hair brush. The snails were scraped off the brush by means of a V-shaped piece of stiff paper mounted on a small plastic funnel. A gelatin capsule was fitted over the end of the funnel and the snails fell into the capsule.

The length of time needed to completely separate all the molluscs from a given quadrat varied from one hour to two weeks. By the time I had collected the third fifty quadrats in August of 1969 I had not yet completely processed the first set of quadrats collected seven months earlier. The entire process of drying, sifting, sorting and identify­ ing molluscs from soil and leaf litter samples took almost three years. The fourth set of fifty quadrats were never collected.

After several quantitative samples were completely analyzed it became apparent that several species were being selected against. Several species which appeared to be common by random collections were quite rare in the litter samples. On order to make sure that all the mollusc populations were being sampled in each vegetation zone, I re­ sumed the random collections periodically. These random stratified collections were made in some specific habitats which were never col­ lected by the quadrat method. This type of data could not be analyzed in the same way, but it would insure a better measure of diversity.

When the molluscs were identified I also noted whether the species was dead, alive, mature or immature. It is very important to know which' species were living in each vegetation zone. It is my experience that all the snail shells smaller than 5 millimeters are translucent if collected alive and one could easily see the dried up body inside. I realized that this number would probably be wrong, but it was done con­ sistently. All samples were processed this way and the data obtained represents the minimum number of molluscs that were living when the sample was collected. RESULTS

The results of this study will be presented in several different forms. Most of the results are presented in form of tables, graphs and maps. The data concerning a specific species, vegetation zone or soil zone will be presented in connection with a discussion of the same. Table 6 lists all the collections made within the study area, when they were made, where they were made and which of the six zona- tion schemes they are in.

All the maps except the topographic map (Map I) and the United States distribution maps for Carychturn nannodes and Vertigo morsel have the same marginal coordinates and the same scale. There is a distribution map for each species collected in the study area with two exceptions. Oxyloma retusa and Oxyloma decampi gouldi are com­ bined into one distribution map because I cannot separate the shells of these two species. The outlines represented on these distribution maps correspond to the various vegetation zones delineated by Frederick (1967). It is important when considering the distribution of each species to remember all the different collections sites that the species was not found in. The presence of one or two points on a map can be very important when placed in perspective with the total number of sites in which the species was not found.

A systematic listing of all the species of mollusca collected in the Bog is presented in Table 7. This listing will give the reader a general idea of the relationships of these mollusca to each other. The order in which the various families appear is not based upon any evolutionary scheme, but on how they appeared in various Identification references. In the case of the terrestrial mollusca the family order is taken from Burch (1962: 188-202).

I found very little information concerning previous collections of mollusca made at Cedar Bog. It is very possible that numerous people have picked up molluscs from this area over the last century. Considering that botanists have been visiting Cedar Bog for quite some time, it seems certain that the numerous molluscs had to have been noticed by someone before Mrs. R. P. Klein collected some in 1951. Table 8 lists the mollusca collected by Mrs. R. P. Klein which were deposited in the Ohio State University Museum of Zoology, those aquatic species recorded by Jon Williams in his Master's Thesis (1966) and the terrestrial molluscs given to me by Jon Williams. The aquatic molluscs collected by Jon Williams were identified by Dr. Henry van der Schalie of the Museum of Zoology at the University of Michigan. At this time,

52 53

there are no mollusca from Cedar Bog catalogued in the collections of the Ohio State University Museum of Zoology. Most of the material collected during this study will be deposited in the Ohio State Uni­ versity Museum of Zoology collections.

i MAP VII Collection locations at Cedar Bog, Champaign County, Ohio.

Dec. 23 & 25, 1968 Jan. 5, 1969 April 23 & 25, 1969 August 20 & 30, 1969 Miscellaneous Collections Oct. 1965 to Jul. 1973

©

WuiihUfn

O 220 440 TABLE 6

Collections node at Cedar Bog Nature Preserve, Champaign County, Ohio

Collection Map Type of Date Vegetation Zone Soil Type Observed Vegetative Number Coordinates Collection Collected Substrate Cover In Frederick (1974) E.C.C. {1973) u.s.o.a, (1971) E.C.C. (1973) Type 1938

1 ft Random 6 Jul. 66 Arbor vitae Evergreen and Carlisle Muck Marl Forested Evergreen-Mixed *

2 'i* Random 6 Jul. 66 Maple-Ash Tuliptree- Carlisle Muck Marl - Forested Basswood Basswood-Ash

3 8Y Qual. Litter 6 Jul. 66 Tuliptree Non-Marl Peat Woody Peat - Open Short Grass

■ 4 7X Random 19 Jul. 66 Tuliptree Tuliptrea- Carlisle Muck Marl Forested Basswood-Ash

5 8X 1 a 19 Jul. 66 Bog Meadow Non-Marl Peat Marl - Forested Short Grass

6 7* Random 25 Aug. 66 Arbor vitae Evergreen and Peat Marl - Forested Evergreen-Mixed

7 7* 1 « 2 29 Aug. 66 Maple-Astv* Tuliptrea- Carlisle Muck Fibrous Peat - Forested Basswood Basswood-Ash

8 8X 1 B2 29 Aug. 66 Tuliptree Non-Marl Peat Woody Peat - Open Short Grass

9 ft I B 2 l6 Sept. 66 Bog Meadow Non-Marl Peat Fibrous Peat m Open Short Grass

10 ft 1 B2 l6 Sept. 66 Marl Meadow Evergreen and Marl Marl - Open Evergreen-Mixed

n ft 1 f t 2 19 Oct. 68 Arbor vitae Evergreen and Peat Marl - Forested Evergreen-Mixed

12 7X 1 ft2 19 Oct. 68 Arbor vitae Evergreen and Peat Marl Peat Forested Evergreen-Mixed TABLE 6- -Continued

Collection Hap Type of Date Vegetation Zone Soil Type Observed Vegetative Number Coordinates Collection Collected Substrate Cover In Frederick (1974) E.C.C. (1973) U.S.D.A. (197I) E.C.C. (1973)

13 7* 1 f t 2 15 Oct. 68 Arbor vitae Evergreen and Peat Marl Muck Forested Evergreen-Mixed

14 8tf 2 ft2 15 Oct. 68 Maplo-Asl>- Tuliptree- Carlisle Muck Fibrous Peat Muck Forested Basswocd Basswood-Ash

15 . 8* 2 ft2 15 Oct. 6b Maple-Ash- Tuliptree- Carlisle Muck Fibrous Peat Muck Forested Basswood Basswood-Ash

16 7Y 2 ft2 15 Oct. 68 Tuliptree Tall Grass Peat Fibrous Peat Muck Forested « 17 1GW 1 ft2 5 Nov. 68 Arbor vitae Maple-Elra # 1 Peat Fibrous Peat - Forested

18 1 ft2 Jl No v . 68 Arbor vitae Maple-Elm #1 Peat Fibrous Peat - Open

1 9- 5 V 2 ft2 5 Nov. 68 Silver & Red Maple-Elm #1 Linwood Muck Muck - Open Kaple

20 9V 2 ft2 5 Nov. 68 Silver & Red Maple-Elra #1 Carlisle Muck Muck - Open Maple

21 7V Random 14 Hay 69 Maple-AsHn Tuliptree Carlisle Muck Fibrous Peat - Forested Basswood Basswood-Ash

22 H R Random 14 May 69 Beech-Maple Mixed-Beech Lippincott Lippincott - Forested Silty-clay Loan Silty-clay Loam

23 10s Random 14 toy 69 Beoch-Maple Mixed-Beech Lippincott Lippincott - Forested Silty-clay Loam Silty-clay Loam

24 7V Randan 14 Hay 69 Qak-Mapla Tuliptree Carlisle Muck Muck - Open Basswood-Ash

25 7* Randan 26 Jun. 69 Arbor vitae Evergreen and Peat Marl • - Forested Everareen-Mixed TABLE 6- -Continued

Collection Map Typo of Date Vegetation Zone Soil Type Observed Vegetative Nuaber Coordinates Collection Collected Substrate Cover In Frederick (1974) E.C.C. (1973) U.S.D.A. (I57I) E.C.C. (1973) Type 1938

26 8W Random 26 Jun. 63 Maple-Asfw- Tuliptrco- Carlisle Muck Fibrous Peat Forested Basswood Basswood-Ash

27 7V Random 26 Jun. 69 Oak-Maple Tuliptree- Carlisle Muck Fibrous Peat Scattered Basswood-Ash Trees

28 6W Random 26 Jun. 63 Vest Branch Cedar Run

29 8V Random 4 Jul. 69 Maple-Astv- Tuliptreo- Carlisle Muck. Fibrous Peat Forested Basswood Basswood-Ash

30 7X Random 4 Jul. 69 Marl Mead cw Evergreen and Peat Marl Open Evorgrocrv-Mixed

31 ior Random 4 Jul. 69 Beech-Maple Mixed-Beech Lippincott Lippincott Open Silt^wlay Loam Silty-clay Loam

32 10T Random 4 Jul. 69 Beech-Maple Mixed-Beech Lippincott Lippincott Open Silty-clay Loam Silty-clay Loam

33 91 Random 4 Jul. 69 Old Field Non-Marl Lippincott Lippincott Open Short Grass Silty-clay Loam Silty-clay Loam

34 ex Random 4 Jul. 69 East Branch Cedar Run

35 4 V Qual. Litter 20 Aug. 69 Arbor vitae Marl Carlisle Muck Marl Open Short Grass

36 7* Qual. Litter 20 Aug. 69 Maple-Asb- Sycamores Carlisle Muck Fibrous Peat Forested Basswood Basswood-Cherry

37 8X Qual. Litter 20 Aug. 69 Tuliptree Evergreen and Peat Marl Forested Evergreen-Mixed TABLE 6- -Continued

Collection Map Typo of Date Vegetation Zone Soil Type Observed Vegetative timber Coordinates Collection Collected Substrate Cover In Frederick (1974) E.C.C. (1973) U.S.O.A. (1971} E.C.C. (1973) Type 1338

38 9* Qual. Litter 30 Aug. 63 Tuliptree Tuliptree- Carlisle Muck Fibrous Peat - Forested Basswood-Ash

39 7Y Random 30 Aug. 69 Tuliptree Tall Grass Carlisle Muck Woody Peat - Open

40 8X Random 30 Aug. 69 Gog Meadow Tall Grass Peat Fibrous Peat - Open

41 10V Random 30 Aug. 69 Beech-Maple Maple^Ash-Elm ■ Linwood Muck Muck - Forested

42 9V Random 30 Aug. 69 Silvor-Red Maple-Elm £1 Carlisle Muck Muck - Open Maple

43 5“ Qual. Litter 20 Oct. 69 Arbor vitae Sycaraore- Carlisle Muck Woody Peat - Forested Basswood-Cherry

44 ex Random 20 Oct. 69 Bog Meadow Tall Grass Peat Fibrous Peat - Open

45 5 * Random 20 Oct. 69 Marl Meadow Marl Peat tori - Open Short Grass

46 6v Random 16 Apr. 70 Oak-Maple Tuliptree- Carlisle Muck Muck Scattered - Basswood-Ash Trees

47 3V Random 16 Apr. 70 Oak-Maple Sycamore Carlisle Muck Muck - Scattered Basswood-Cherry 1 Trees

48 5* Random 18 Jul. 73 Marl Meadow Marl Peat Marl - Open Short Grass

49 SY Random 18 Jul. 73 Bog Meadow Maple-Elm £2 Peat Muck - Open

50 10U Random 18 Jul. 73 Beech-Maple Maple-Ash-Elm Lippincott Muck - Forested Silty-clay Loan TABLE &- -Continued

Collection Map Type of Date Vegetation Zone Soil Type Observed Vegetative Nunber Coordinates Collection Collected Substrate Cover In Frederick (1974) E.C.C. (I973) U.S.D.A. (1971) E.C.C. (1973) Type 1938

51 (M Random 15 Oct. 68 Maplfr^sb- Sycamore- Carlisle Muck Muck - Open Basswood Basswood-Cherry

52 7V Random 26 Jun. 69 Oak-Maple Tulip tre&- Carlisle Muck Muck - Scattered Basswood-Ash Trees

53 ?x Random IB Jun. 73 Maple-Asfv- Tuliptree- Carlisle Muck Marl - Forested Basswood 8asswood-Ash

54 9V Random 14 M^y 69 Silver-Red Maple-Elm #1 Linwood Muck Muck - Open Maple

55 4 V 1 ft2 5 Jan. 69 Arbor vitae Evergreen and Carlisle Muck Marl Marl Scattered Evergreen-Mixed Trees

56 5 * 1 ft2 5 Jan. 69 Arbor vitae Evergreen and Feat Marl Peat Scattered Evergreen-Mixed Trees

57 6x 1 f t 2 ■ 5 Jan, 69 Arbor vitae Marl Peat Marl Peat Forestod Short Grass

58 1 W 1 ft2 23 Dec. 68 Arbor vitae Maple-Elm #1 Peat Marl Marl Forested

59 10X 1 ft2 23 Dec. 68 Arbor vitae Maplo-Elra #1 Peat Muck Peat Open 1 60 6w 1 ft2 30 Deo. 68 Maplo-AtJv* Tuliptree* Carlisle Muck Fibrous Peat Muck Forested Basswood Basswood-Ash ■

61 7v 1 f t 2 30 Dec. 68 Maple-Asb- Tuliptrea- Carlisle Muck Fibrous Peat Muck Forested Basswood Basswood-Ash

62 8W 1 ft2 3® 60 Maple-Asf^ Tuliptree- Carlisle Muck Fibrous Peat Muck Forested Basswood Basswood-Ash

63 BH 1 ft2 30 Dec. 68 Maple-Astv* Tuliptreo* Carlisle Muck Fibrous Peat Muck Forested Basswood Basswood-Ash TABLE 6- -Continued

le c tio n Map Type o f Date Vegetation Zone S o il Type Observed V egetative ber Coordinates C o lle ctio n C ollected Substrate Cover In Frederick (1974) E.C.C. 1973 ( ) U.S.D.A. (1971} E.C.C. (1973)

& 5V 1 f t 2 30 Dec. 68 Maple-Asf^ Sycamore Carlisle Muck Fibrous Peat Muck Forested Basswood Basswood-Ash

65 6x 1 f t 2 30 Dec, 68 T u lip tre e Evergreen and Carlisle Muck Marl Muck Forested Evergreen-Mixed

66 5V 1 f t 2 30 Dec. 68 T u lip tre e Sycanore- Carlisle Muck Fibrous Peat Muck Forested Basswood-Cherry

• 67 ■ 3 * 1 f t 2 23 Dec. 6B T u lip tre e T u lip tre e - Carlisle Muck Fibrous Peat Muck Forested Basswood-Ash

6s 9* 1 f t 2 23 Dec. 68 Tuliptree Tuliptree- Carlisle Muck Fibrous Peat Muck Forested Basswood-Ash

69 11V 1 f t 2 23 Dec. 68 T u lip tre e tople-Ash-Elm Carlisle Muck Fibrous Peat Muck Forested

70 2X 1 f t 2 23 Dec. 68 Bog Meadow This sample was lo s t.

n 2X 1 f t 2 5 Jan. 69 Bog Meadow * Evergreen and Peat Woody Peat Peat Open Evergreen-Mixed

72 3X 1 f t 2 3 Jan. 69 Bog Meadow Evergreen and Peat Woody Peat Peat Scattered Evergreen-Mixed Trees

73 low 1 ft2 23 Dec. 68 Bog Meadow Evergreen and Peat t o r i Peat Open Evergreen-Mixed

74 SW 1 ft2 23 Dec. 68 Bog Meadow M arl Peat Fibrous Peat Peat Open Short Grass

75 3X 1 ft2 5 Jan. 69 M arl Meadow M arl M arl Woody Peat t o r i Open Short Gra$3 o\ 76 4X 1 ft2 5 Jan. 69 M arl Meadow M arl M arl . M arl M arl Open o Short Grass TABLE 6- -Continued

Collection Map Type of Date Vegetation Zone Soil Type Observed Vegetative Number Coordinates Collection Collected Substrate Cover In Frederick (1974) E.C.C. (1373) U.S.D.A. (1971) E.C.C. (1973) Type 1938

77 5 * 1 ft2 30 Dec: 68 Marl Meadow Marl Peat Marl Marl Open Short Grass

78 & 1 ft2 30 Dec. 68 Marl Meadow Evergreen and Marl Marl Marl Open Evergreen-Mixed

79 1ft2 30 Dec. 68 Marl Meadow Marl Marl Marl Marl Open Short Grass

80 10S 1ft2 23 Dec. 68 Beech-Maple Mixed-Beech Lippincott Lippincott Loan Open Silty-clay Loam Silty-clay Loan

81 10T 1 ft2 23 Dec. 68 Beech-Maple Mixed-8eech Lippincott Lippincott Loan Open Silty-clay Loan Silty-clay Loan

82 10U 1ft2 23 Dec. 68 Beech-Maple Haple-AsWLIm Lippincott Muck Loan Forested Silty-clay Loan

83 10V 1 ft2 23 Dec. 68 Beech-Maple Maple-Ash-Elm Linwood Muck Muck Muck Forested

84 10V 1 ft2 23 Dec. 68 Beech-Maple Maple-Elm #1 Linwood Muck Muck Muck Forested

85 5 V 1 ft2 30 Dec. 68 Oak-Maple Sycamore-Maplfr. Linwood Muck Muck Muck Open Cottonwood

86 5 V 1ft2 30 Dec. 68 Oak-Maple Sycamore- Carlisle Muck Muck Muck Open Basswood-Cherry

87 6v 1ft2 30 Dec. 68 Oak-Maple Tuliptre&- Carlisle Muck Muck Muck Open Basswood-Ash

88 8V 1ft2 30 Dec. 68 Oak-Maple Tuliptrefr- Lippincott Fibrous Peat Muck Scattered Basswood-Ash Silty-clay Loan Trees TABLE 6- -Continued

Collection Map Type of Date Vegetation Zone Soil Type Observed Vegetative Number Coordinates Collection Collected Substrate Cover In Frederick (1974) E.C.C. (1973) U.S.D.A. (1971) E.C.C* (1973) Type 1938

89 8V 1 ft 30 Dec. 68 Oak-Maple Tuliptree Carlisle Muck Fibrous Peat Muck Scattered Basswood-Ash Trees

50 9V 1 ft 23 Dec. 68 Silver-Red Maple-Elm #1 Linwood Muck Muck Muck Open Maple

91 *9V 1 ft 23 Dec. 68 Silver-Red Maple-Elm #1 Linwood Muck • Muck Muck Open Maple

92 9V 1 ft 23 Dec. 68 Silver-Red Maple-Elm # 1 Carlisle Muck Muck Muck Open , Maple

93 9V 1 f t 23 Dec. 68 Silver-Red Maple-Elm #1 Carlisle Muck Muck Muck Open Maple

94 10V 1 ft 23 Dec. 68 Silver-Red Maple-Elm #1 Carlisle Muck Muck Muck Forested Maple

95 IV 1 ft 30 Dec. 68 Shrub Sycamore- Carlisle Muck Muck Muck Scattered Basswood-Cherry Trees

96 11T 1 f t 23 Dec. 68 Shrub Mixed-Beech Linwood Muck Lippincott [tick Forested Silty-clay Loam

97 11T 1 ft‘ 23 Dec. 68 Shrub Mixed-Seech Lippincott Lippincott Muck Forested Silty-clay Loam Silty-clay Loam

98 n u 1 ft 23 Dec. 68 Shrub Maple-Ash-Elm Lippincott Lippincott Muck Forested Silty-clay Loam Silty-clay Loam

99 12V 1 ft 23 Dec. 68 Shrub Maplo-Ash-Ela Carlisle Muck Fibrous Peat Peat Forested

100 zx 1 f t 3 Jan. 69 Elm-Maple Sycanor&- Carlisle Muck Woody Peat Muck Scattered Basswood-Cherry Trees TABLE 6- -Continued

Collection Hap Typo of Date Vegetation Zono Soil Type Observed Vegetative Number Coordinates Collection Collected Substrate Cover In Frederick (1974J E.C.C. (1973) U.S.O.A. (1971) E.C.C* (1973) Type 1938

101 ZX 1 ft2 5 Jan. 69 Elm-Maple Marl Linwood Muck Marl Muck Scattered Short Grass Troes

102 2V 1ft2 30 Dec. fe Elm-Maple Sycamore- Carlisle Muck Muck Muck Open Basswood-Cheriy

103 3V 1 ft2 30 Dec. 68 Elm-Maple Sycamore- Carlisle Muck Muck Muck Open Basswood-Cherry

104 5Y 1 ft2 23 Dec. 68 Elm-Maple Maple-Elm #2 Peat Muck Muck Open

105 7 X 1 ft2 23 Apr. 69 Arbor vitae Evergreen and Carlisle Muck Marl Muck Forested Evorgreen-Mixed

106 4 X 1 ft2 25 Apr. 69 Arbor vitae Evergreen and Peat Woody Peat Peat Scattered Evergreen-Mixed Trees

107 9X 1 ft2 25 Apr. 69 Arbor vitae Evergreen and Peat Fibrous Peat Peat Open Evergreen-Mixed

108 10X 1ft2 25 Apr. 69 Arbor vitae Evergreen and Peat Muck Peat Open Evergreen-Mixed

109 11V 1 ft2 25 Apr. 69 Arbor vitae Maple-Asb-Elo Peat Fibrous Peat Muck Forested

110 8W 1ft2 23 Apr. 69 Maple-Ask* Tuliptreo- Carlisle Muck Muck Muck Forested Basswood Basswood-Ash

111 7* 1 ft2 23 Apr. 69 Maple-Asb- Tuliptree- Carlisle Muck Marl Muck Forested Basswood Basswood-Ash

112 to 1 ft2 23 Apr. 69 Haple-AsJv- Tuliptree- Carlisle Muck Woody Peat Muck Open Basswood Basswood-A&h TABLE 6- -Continued

Collection Hap Type of Date Vegetation Zone Soil Type Observed Vegetative Number Coordinates Collection Collected Substrate Cover In Frederick (I974J E.C.C. (1973 U.S.D.A. (1971) E.C.C. (1973) Type 1938

113 Q i i n 2 23 Apr. 69 Maple*J\sf^ Sycaraor&» Carlisle Muck Muck Muck Forested Basswood Basswood-Cherry

114 8W 1 ft2 23 Apr. 69 Maple-Asb- Tuliptree Carlisle Muck Muck Muck Forested Basswood Basswood-Ash

115 8X 1ft2 23 Apr. 69 Tuliptree Tuliptree- Carlisle Muck Marl Muck Forested Basswood-Ash

116 5V 1ft2 23 Apr. 69 Tuliptree Sycaraoro- Carlisle Muck Muck Muck Scattered Basswood-Cherry Trees

117 5 M 1ft2 25 Apr. 69 Tuliptree Tuliptree- Carlisle Muck Muck Muck Forested Basswood-Ash

118 9* 1ft2 25 Apr. 69 Tuliptree Tuliptreo- Carlisle Muck Marl Muck Forested Basswood-Ash

113 1ft2 ‘ 25 Apr. 69 Tuliptree Tuliptree- Peat Marl Muck Forested Basswood-Ash

120 8Y 1 ft2 23 Apr. 69 Bog Meadow Non-Marl Peat Woody Peat Muck Forested Short Grass

121 4 W 1 ft2 23 Apr. 69 Bog Meadow M arl Marl Marl Marl Open Short Grass

122 4 X 1ft2 23 Apr. 69 Bog Meadow Non-Marl Peat Fibrous Peat Marl Open Short Gras3

123 2X 1ft2 23 Apr. 69 Bog Meadow Marl Peat Woody Peat Peat Open Short Grass TABLE -Continued

Map Typo of Date Vegetation Zone Soil Type Observed Vegetative Coordinates Collection Collected Substrate Cover In Frederick (1274) E.C.C. (1273) U.S.D.A. (1271) E.C.C. (1573) Type 1338 ■ 124 9* 1 ft2 25 Apr. 63 Bog Meadow Maple-Elm #2 Peat Muck Marl Open

125 6X 1 ft2 23 Apr. 65 Marl Meadow Marl Marl Marl Marl Qpen Short Grass

126 5 * 1 ft2 23 Apr. 63 Marl Meadow Marl Peat Marl Marl Open Short Grass

127 5* 1 ft2 23 Apr. 63 Marl Meadow Evergreen and Peat Marl Marl Open Evergreen-Mixed

128 4 X 1 ft2 23 Apr. 63 Marl Meadow Marl Marl Marl Marl Open Short Grass

12? 3X 1 ft2 23 Apr. 63 Marl Meadow Evergreen and Marl Marl Marl Open Evergreen-Mixed

130 10s 1 ft2 • 25 Apr. 63 Beech-Maple Mixed-Beech Lippincott Lippincott Muck Open Silty-clay Loam Silty-clay Loom

131 IOR 1 ft2 25 Apr. 63 Beech-Maple Mixed-Beech Lippincott Lippincott Muck Open Silty-clay Loam Silty-clay Loam

132 ior 1ft2 25 Apr. 63 Beoch-Maple Mixed*43eech Lippincott Lippincott Loom Forested Silty-clay Loam Silty-clay Loam

133 11T 1 ft2 25 Apr. 63 Beech-Maple Mixed-Beech Lippincott Lippincott Muck Forested Silty-clay Loam Silty-clay Loam

134 12S 1 ft2 25 Apr. 63 Beech-Maple Mixed-Beech Lippincott Lippincott Muck Forested Silty-clay Loam Silty-clay 1 fWBI

8V l.ft2 23 Apr. 63 Oak-Maple Tuliptree Linwood Muck Muck Muck Open Basswood-Ash TABLE 6- -Continued

Collection Hap Type of Date Vegetation Zone Soil Type Observed Vegetative Number Coordinates Collection Collected Substrate Cover In Frederick (1974) E.C.C. (1973) U.S.D.A. (1971) E.C.C. (1973) Type 1938

136 1 ft Tuliptree- Carlisle Muck Muck Muck Open 77 23 Apr. 69 Oak-Maple Basswood-Ash

137 6v 1 ft 23 Apr. 69 Oak-Haple Tuliptre&- Carlisle Muck Muck Kx:k Scattered Basswood-Ash Trees

138 4 V 1 ft 23 Apr. 69 Oak-Maple Sycamore- Carlisle Muck Muck Muck Scattered Basswood-Cherry Trees

• 133 4 V 1 ft 23 A pr. 59 Oak-Maple Sycamore- Carlisle Muck Muck Muck Scattered Basswood-Cherry Trees

140 9V 1 ft 25 Apr. 69 Silver-Red Maple-Elm #1 Linwood Muck Fibrous Peat Muck Open Maple

141 9* 1 ft‘ 25 Apr, 69 SilveM?ed Maple-Elm #1 Linwood Muck Fibrous Peat Muck Open Maple

142 9V 1 ft 23 Apr. 69 Silver-Rod Maple-Elm #1 Linwood Muck Muck Muck Open Maple

143 10U 1 ft 25 Apr. 69 Silver-Red Maple-Elm #1 Linwood Muck Muck Muck Open Maple

144 10V 1 ft 25 Apr. 69 Silver-Red Maple-Elm #1 Linwood Muck Muck Muck Forested Maple

145 2V 1 ft 23 Apr. 59 Shrubs Sycamore- Carlisle Muck Muck Peat Scattered Basswood-Cherry Trees

145 10T 1 ft 29 Apr. 59 Shrubs Maple4 teh-Elra Lippincott Lippincott Muck Open Silty-clay Loam Silty-clay Loam-

147 10U 1 ft 29 Apr. 69 Shrubs Kaple-Ash-Elm Lippincott Muck Muck Forested Silty-clay Loam TABLE to -Continued

Collection Hap Type of Date Vegetation Zone Soil Type Observed Vegetative Number Coordinates Collection Collected Substrate Cover In Frederick {1974) E.C.C, (1373) U.S.D.A, (1971) E.C.C. {1973} Type 1938

161 (M 1ft2 20 Aug. 69 Maple-Asf>- Sycamore- Carlisle Muck Fibrous Peat Muck Forested Basswood Basswood-Cherry

162 to 1ft2 20 Aug. 69 Maple-Asb- Tuliptree- Carlisle Muck Fibrous Peat Muck Forested Basswood Basswood-Ash

163 8W 1ft2 20 Aug. 69 Haplo-Asb- Tuliptroo- Carlisle Muck Fibrous Peat Muck Forested Basswood Basswood-Ash

164 to 1 ft2 20 Aug. 69 Maple-Asfr- Sycamore- Carlisle Muck Fibrous Peat Muck Open Basswood Basswood-Cherry

165 u v 1ft2 30 Aug. 69 Tuliptree Haple-Ash-Elo Carlisle Muck Fibrous Peat Muck Forested

166 SW 1ft2 30 Aug. 69 Tuliptree Tuliptreo- Carlisle Muck Fibrous Peat Muck Forested Basswood-Ash

167 9 > 1 ft2 30 Aug. 69 Tuliptree Tuliptree- Carlisle Muck Fibrous Peat Muck Forested Basswood-Ash

160 7Y 1ft2 30 Aug. 69 Tuliptree Non-Marl Carlisle Muck Fibrous Peat Muck Open Short Grass

169 ‘ 9" 1 ft2 30 Aug. 69 Tuliptree Evergreen and Carlisle Muck Marl Muck Forested Evergreen-Mixed

170 8X l ‘ft2 30 Aug. 69 Bog Meadow Non-Marl Peat Fibrous Peat Marl Open Short Grass

171 8Y 1 ft2 30 Aug, 69 Bog Meadow Non-Marl Peat Woody Peat Muck Forested Short Grass

172 2X 1 ft2 20 Aug. to Bog Meadow Evergreen and Peat Woody Peat ' Muck Open Evergreen-Mixed TABLE 6- -Continued

Hap Type of Date Vegetation Zone Soil Type Observed Vegetative Coordinates Collection Collected Substrate Cover In Frederick (1374) E.C.C. (1373) U.S.O.A. (1371) E.C.C. (1573) TyP0 1938

173 8X 1 ft 30 Aug. 63 Bog Meadow T a U Grass Peat Fibrous Peat Muck Open

274 1GW 1 ft 30 Aug, 6? Bog Meadow Marl Peat Fibrous Feat Marl Open Short Grass

175 & 1 ftc 20 Aug, 63 Marl Meadow Marl Marl Marl Marl Open Short Grass

176 5* 1 ft 20 Aug. 63 Marl Meadow Marl Marl Marl Marl Open Short Grass

177 3* 1 ft 20 Aug. 63 Marl Meadow Evergreen and Marl Woody Peat Marl Open Evergreen-Mixed

178 4 X 1 ft" 20 Aug, 63 Marl Meadow Marl Marl Woody Peat Marl Open Short Grass

179 4 X 1 ft 30 Aug. 63 Marl Meadow Marl Marl Woody Peat Marl Open Short Grass

180 10s 1 ft‘ 30 Aug. 63 Beech-Maple Mixed-Beech Lippincott Lippincott Loan Forested Silty-clay Loan Silty-clay Loam

181 US 1 ft 3P Aug. 63 Beech-Maple Mixed-Beech Lippincott Lippincott Loam Forested Silty-clay Loan Silty-clay Loam

182 11R 1 ft 30 Aug. 63 Beech-Maple Mixed-Beech Lippincott Lippincott Muck Open SUty-day Loao Silty-clay Loam

183 12T 1 ft 30 Aug. 63 Beech-Maple Mixed-Beech Linwood Muck Lippincott Muck Forested Silty-clay Loam

UR l f t ‘ 30 Aug, 63 Beech-Maple Mixed-Beech Lippincott Lippincott Loao Open Silty-clay Loao Silty-clay Loom TABLE 6- -Continued

Collection Map Type of Date Vegetation Zone Soil Type Observed Vegetative Number Coordinates Collection Collected Substrate Cover In Frederick (1974) E.C.C. (1573) U.S.D.A. (1971) E.C.C. (1973) Type 1938

185 5V 1 ft 20 Aug, 69 Oak-Maple Sycomore- Carlisle Muck Muck Muck Scattered Basswood-Cherry Trees

186 8V 1 ft 20 Aug. 69 Oak-Maple Tuliptree- Carlisle Muck Fibrous Peat Muck Scattered Eosswood-Ash Trees

187 4 V 1 ft 20 Aug. 69 Oak-Maple Sycanore- Linwood Muck Muck Muck Open 8asswood-Cheny

188 3V 1 ftc 20 Aug. 69 Oak-Maple Sycamore Carlisle Muck Muck Muck Scattered Basswood-Cherry Trees

18? 8V 1 ft 20 Aug. 69 Oak-Maple Tuliptree- Carlisle Muck Fibrous Peat Muck Scattered Basswood-Ash Trees

190 SW 1 ftc 30 Aug. 69 Silver-Red Maple-Elm #1 Carlisle Muck Fibrous Peat Muck Open Maple

191 10V 1 fte 30 Aug. 69 Silver-Red Maple-Elm f f l Carlisle Muck Fibrous Peat Muck Forested Maple

192 9V 1 ft 30 Aug. 69 Silver-Red Maple-Elm # 1 Carlisle Muck Muck Muck Open Maple

193 9V 1 ft 30 Aug. 69 Silver-Red Maple-Elm f f l Linwood Muck Muck ' Muck Open Maple

194 9V 1 fte 30 Aug, 69 Silver-Red Maple-Elm # 1 Linwood Muck Muck Muck Open Maple

195 2v 1 ft 20 Aug. 69 Shrubs Sycamore- Carlisle Muck Muck Muck Scattered Basswood-Cherry Trees TABLE 6- -Continued

Collection Mop Type of Date Vegetation Zone Soil Type Observed Vegetative Number Coordinates Collection Collected Substrate Cover In Frederick (1974) E.C.C. (1973) U.S.D.A. (1971) E.C.C. (1973} Type 1938 • 156 IV 1 ft2 20 Aug. 69 Shrubs Sycamore- . Carlisle Muck Muck Muck Scattered Basswood-Cherry Trees

157 11T 1 f t 2 30 Aug. 69 Shrubs Maple-Ash-Elm Linwood Muck Lippincott Muck Forested Silty-clay Loao

15S 2V 1 f t 2 20 Aug. 69 Shrubs Sycamore- Carlisle Muck Muck Muck Cpen Basswood-Cherry

199 11T 1 ft2 30 Aug. 69 Shrubs Maple-Ash-£Ira Linwood Muck Lippincott Muck Forested Silty-clay Loam

200 6v 1 ft2 20 Aug. £9 Elm-Maple Sycamore- Carlisle Muck Fibrous Peat Muck Scattered Basswood-Cherry Trees

201 ur 1 ft2 20 Aug. 69 Elm-Maple Sycamore- Linwood Muck Marl Muck Scattered Basswood-Cherry Trees

202 4V 1 ft2 20 Aug. 69 Elm-Maple Sycamore Carlisle Muck Marl Muck Scattered Basswood-Cherry Trees

2C0 IX 1 ft2 20 Aug. 69 Elm-Maple Sycamore- Carlisle Muck Woody Peat Marl Scattered Basswood-Cherry Trees 1 204 a t 1 ft2 20 Aug, 69 Elm-Maple Sycamore- Linwood Muck Marl Muck Scattered Basswood-Cherry Trees

205 Not Happed Qual. Litter 25 Oct. 65 Arbor vitae - M - - -

206 Not Mapped Qual. Litter 25 Oct. 65 Maple-Aah- - - - - Basswood

207 Not Happed Qual. Utter 25 Oct. 65 Tuliptree _ m - «• TABLE 6- -Continued

Collection Map Type of Date Vegetation Zone Soil Type Observed Vegetative Number Coordinates Collection Collected Substrate Cover In Frederick 1974 ( ) E.C.C. (1973) U.S.D.A. ( 1971) E.C.C. (1973) Type 1938

206 Not Mapped Qual, Litter 25 Oct. 65 Elm-Maple - m - -

209 Not Mapped Random 5 Mar. 66 Arbor vitae - M - -

210 Not Mapped Random 3 Mar. 66 Bog Meadow - - - - -

211 Not Mapped Random 6 Jul. 66 Tuliptree - - - --

212 Not Mapped Qual. Litter 19 Jul. 66 Arbor vitae - - - - -

213 Not Mapped Random 19 Jul. 66 Marl Meadow - - --

M 214 Not Mapped Random 3 Aug, 66 Maple-Ash-Basswood -- -

215 Not Mapped Random 3 Aug. 66 Tuliptree - - - - -

216 Not Mapped Random 3 Aug. 66 Marl Meadow -- - -

217 Not Mapped Random 25 Aug. 66 Maplo-Ash-Basswood - - - - -

M 218 Not Mapped Random 3 Feb. 68 Maple-Ash-Basswood - -- -

mm 219 Not Mapped Random 3 Feb. 68 Bog Meadow - - -

220 Not Mapped Random 1 Jun. 6b Maple-Ash-Basswood - - - - -

221 Not Mapped Random 1 Jun. 68 Tuliptree - - - - -

222 Not Mapped Random 1 Jun. 68 Marl Meadow - - - - -

223 Not Mapped Random 1 Jun. 68 Bog Meadow «* • _ _ - TABLE 6- -Continued

C o lle ctio n Map Type o f Oats Vegetation Zone S o il Type Observed V egetative Humber Coordinates C o lle ctio n C ollected Substrate Cover In Frederick 1974 ( ) E.C.C. ( 1973) U.S.D.A. ( 1973) e .c .c . 1973 ( ) Type 1938

225 Not Mapped Random 8 J u l. 6 b Bog Meadow - -- - -

226 Not Mapped Random 5 Nov. 68 Oak-Maple - - mm - -

227 Not Mappod Random 14 May 69 Arbor vitae *■ - mm mm mm

228 Not Mapped Random 14 May 69 Tuliptree - - - - -

22? Not Mapped Random 26 Jun. 69 Bog Meadow - mm m -

230 Not Mapped Random 4 J u l. 69 Bog Meadow - - mm m -

231 Not Mapped Random 20 O ct. 69 Arbor vitae - - mm - mm

232 Not Mapped Random 20 Oct. 69 Tuliptree - •• - - «■

233 Not Mapped Random 7 Apr. 70 Oak-Maple - - - -

234 Not Mapped Random l6 Apr, JO Maplo>Jtsh-8asswood - - m - -

235 Not Mapped Random 16 Apr. 70 Elo-Maple - mm - - -

■ki w TABLE 7

Phylogenetic listing of the Mollusca collected at the Cedar Bog Nature Preserve, Champaign County, Ohio.

Class Gastropoda Subclass Prosobranchia Order Mesogastropoda Family Valvatidae Valvata tricarinata (Say, 1817) Valvata sincera (Say, 1824) Family Hydrobiidae Amnicola limosa (Say, 1817) Family Pomatiopsidae Pomatiopsis lapidaria (Say, 1817) Family Pleuroceridae Oxytrema semicarinata (Say, 1829) Subclass Pulmonata Order Basommatophora Family Lymnaeidae Lymnaea caperata Say, 1829 Lymnaea humllis Say, 1822 Family Planorbldae Helisoma anceps (Henke, 1830) Gyraulus parvus (Say, 1817) Planorbula armigera (Say, 1818) Family Physidae Physa Integra Haldeman, 1841 Family Carychildae Carychium exiguum (Say, 1822) Carychium exile H, C. Lea, 1842 Carychium nannodes Clapp, 1905 Order Stylommatophora Suborder Orthurethra Family Cochlicopidae Cochlicopa lubrica (Mtlller, 1774) Family Valloniidae Vallonia excentrica Sterki, 1893 *■ Family Pupillidae Pupoides albllabris (C. B. Adams, 1841) Gastrocopta procera (Gould, 1840) Gastrocopta armifera (Say, 1821) Gastrocopta contracta (Say, 1822) Gastrocopta pentodon (Say, 1821) Gastrocopta tappaniana (C. B. Adams, 1842) . Gastrocopta corticaria (Say, 1817) Vertigo morsel Sterki, 1894 Vertigo ovata Say, 1822 Vertigo pygmaea (Draparnaud, 1801) Vertigo elatior Sterki, 1894) Vertigo gouldl (Binney, 1843) TABLE 7— Continued

Vertigo milium (Gould, 1840) Vertigo tridentata Wolf, 1870 Columella edentula (Draparnaud, 1805) Family Strobilopsidae Strobilops aenea Pilsbry, 1926 Strobilops labyrlnthica (Say, 1817) Suborder Heterurethra Family Succlneidae Succinea ovalis Say, 1817 Catinella vermeta (Say, 1829) Oxyloma retusa (H. C. Lea, 1834) Oxyloma decampi gouldi Pilsbry, 1948 Suborder Sigmurethra Family Philomycidae Philomycus carolinianus (Bose, 1802) Family Endodontidae Anguispira alternata (Say, 1817) Helicodiscus parellelus (Say, 1821) Punctum minutissimum (H. C, Lea, 1841) Family Limacidae Deroceras laeve (MUller, 1774) Family Zonitidae Nesovitrea binneyana (Morse, 1864) Nesovitrea electrina (Gould, 1841) Glyphyalinia indentata (Say, 1823) Hawaiia minuscula (Binney, 1840) Euconulus chersinus (Say, 1821) Euconulus fulvus (Mllller, 1774) Zonitoides arboreus (Say, 1817) Guppya sterkii (Dali, 1888) Striatura milium (Morse, 1859) Striatura exigua (Stimpson, 1850) Family Haplotrematidae Haplotretna concavum (Say, 1821) Family Polygyridae Stenotrema barbatmn (Clapp, 1904) Stenotrema leai (Binney, 1821) Mesodon clausus (Say, 1821) Mesodon thyroidus (Say, 1817) Mesodon inflectus (Say, 1821) Triodopsls multilineata (Say, 1821) Allogona profunda (Say, 1821) Class Bivalvia Subclass Lamellibranchia Order Heterodonta Family Sphaerildae Pisidium compressum Prime, 1851 Pisidium casertanum (Poli, 1791) TABLE 8

Mollusca collected at Cedar Bog fiature Preserve, Champaign County, Ohio by other investigators.

q b Collected by Collected by R. P. Klein J, L. Williams Sept. 12, 1951 toy to June, 1965

Carychium exile H.C. Lea, 1842 X

Carychium exiquum (Say, 1822)X

Gastrocopta contracts (Say, 1822)X

Gastrocopta pentodon (Say, 1821) X

Gastrocopta tappaniana (C.B. Adams, 1842)X

Gastrocopta tappaniana form curta (Sterki, 1894) X

Vertiqo milium (Gould, 1840 ) X

Strobilops labyrinthica (Say, 1817) X

Succinea ovalis Say, 1817 X

Succinea sp. X

Helicodiscus parallelus (Say, 1821) X

Retinella indentata (Say, 1823) X

Retinella binneyana (Morse, 1 8 & 0 X

Retinella olectrina (Gould, 1841 ) X

Zonitoides arboreus (Say, 18l6) X

Striatura milium (Morse, 1859) X

Haplotrema concavum (Say, 1821) X

Stenotrema hirsutum (Say, 1817)X

Stenotrema monodon (Rackett, 1821) X

Mesodon thyroidus (Say, 18l6)X

Mesodon inflectus (Say, 1821) X

Triodopsis multilineata (Say, 1821) X

Alloqona profunda (Say, 1821) X

Lymnaea humilis Say, 1822 77 TABLE 8— Continued

Collected by a Collected by R. P. Klein J, L. Williams Sept. 12, 1951 May to June, ‘19&5

Planorbula armiqera (Say, 1818) X

Goniobasis livescens (Msnko, 1830} X

Pisidium casertanum (Poll, 1791)X

Q Specimens deposited in the Ohio State University Musoum of Zoology, k Records taken from 1966 Master's thesis and personal communication. DISCUSSION OF DEFINITIONS AND DATA ANALYSIS PROCEDURES

While comparing the molluscan faunas of the various plant communi­ ties I became aware of several dimensions of my data. These data gave me information on what species were present, how many species were present, how often each species was encountered in the collections and how many of each species were present. When dealing with molluscs, all. the above mentioned aspects of the data can be expressed in terms of the total number collected or the living number collected. It will become evident that the conclusions made are greatly affected by whether total numbers or living numbers are used. In an attempt not to distort the facts I will, where possible, present an analysis of both aspects of these data.

When comparing the faunas of various vegetation zones and the habitats of the various species I will be using a number of terms such as density, frequency, diversity, species diversity Index and habitat diversity index. Each term is defined below in the manner In which it is used in this study.

Density

Density can be defined In more than one way. Smith (1966: 342) considers two types of density measurements, a crude density and an ecological density. Crude density refers to the number collected per unit area and ecological density is the number collected per available living space. I am not able to adequately define the available living space for each species. The living space available for each species Is not necessarily the same. Until the habitat requirements for each species are better known this type of density measurement cannot be made for terrestrial mollusca. Therefore, when I refer to density I mean only crude density. Odum (1959: 150) states that the population density of many Invertebrate groups can change rapidly or are very difficult to accurately measure. Because of this, density measurements other than crude, can have little meaning In this study. This certainly could be true, but most terrestrial mollusca native to this area have one or two generations each year and some take two or three years to mature. I believe terrestrial mollusca respond to environmental changes, but not so rapidly that density information cannot be of value. In this study the sampling procedure used Is only accurate for some species. If the density data are qualified properly, I believe it to be useful for comparing habitat preferences. Most of the density data used are averages of collections taken at different times of the

78 79 year and only from quantitative samples. This reduces the variations- present in individual samples and variable collection techniques.

Frequency

Frequency is defined as the number of collections in which each species appears from a given area. Frequency is usually expressed as a percentage of total number of collections made from that same area. Basically, the measurement refers to the incidence of occurrence for each species or how often a given species will be encountered in a given habitat or vegetation zone. Burch (1955: 156-157) used a fre­ quency measurement in his descriptions of the distribution of terres­ trial molluscs in several counties in Virginia. Burch used frequency terms such as abundant, common, frequent, occasional, and rare in a very particular way. I am not assigning categories to the various frequency levels.

Plates I to XII graphically illustrate how the frequency of most species varies in the different vegetation zones as they are defined by Frederick (1974). These frequency comparisons are based on all the 234 collections made within the study area. Hopefully, this will reduce or perhaps average the bias innate to a single collecting pro­ cedure. Density is more dependent on a standardized collecting pro­ cedure or quantitative sampling, whereas frequency is not.

Diversity

Species diversity generally refers to the number of species found in a given area. This is the simplest means of measuring diversity. Many times this is as far as one can go. However, if the organisms involved can be sampled in some way to indicate density, then diversity can take on other meanings. Lloyd, Zar and Karr (1968: 257) incorpor­ ate the number of individuals for each species into their definition for diversity. Actually, Lloyd et al (1968) were not the first investiga­ tors to use species numbers in the definition of diversity, Margalef (1957), MacArthur and MacArthur (1961), Lloyd and Ghelardi (1964), Pielou (1966), Goulden (1969), McNaughton (1974) and Getz (1974) have also used a revised definition of diversity. In my opinion they are not really altering the definition as much as they are adding a new dimension. What they are using is properly called a diversity index. Diversity and a diversity index are not necessarily the same. Within the context of this paper diversity will refer to the number of species of a given area. I will refer to a total diversity, which includes all the dead and living specimens collected, and a living diversity which includes only the living specimens collected. 80

Diversity Index

Pennak (1964: 160) defined diversity index as the "ratio between number of species and the number of individuals in a community." This is a very generalized definition for what can be a highly complex measurement. First of all, there are many diversity indices. Odum (1963: 32) defined two diversity indices as the ratio of the cumula­ tive number of species and the logarithm of the individuals counted or the ratio of the cumulative number of species and the square root of the individuals counted. In some studies diversity increases on a straight line basis as the logarithm of the individuals counted or the square root of the individuals counted, Odum (1959: 150) considered these ratios as being characteristic of the community. Pielou (1966a) defines diversity (diversity index) as "the amount of uncer­ tainty that exists regarding the species of an individual selected at random from a population. The more species there are and the more nearly even their representation, the greater the uncertainty and hence the greater the diversity." Some of the formulas used for measuring this degree of uncertainty were developed by communication engineers for predicting correctly the name of the next letter in a message (Krebs, 1972: 506). Pielou (1966b) reviews the various divers­ ity indices developed and indicates when each index should be used. Pielou states that the type of diversity index used is dependent upon what kind of sampling can be performed and whether the number of species is known or unknown. According to Ted Smith (1975) my data best fits the Shannon-Wiener formula for a diversity index. I can sample the soil and litter species randomly and I can know the total number of species present. I found two molluscan studies which used the Shannon-Wiener index. Randolph (1973) studied the influence of environment on two species of land snails in Texas and Getz (1974) studied the mollusca in various vegetation zones in the Smoky Mountains.

The only data I used to calculate all the diversity indices were the quantitative samples (55 through 204) which were made in a com­ pletely random and uniform way. This leaves out certain valuable data and causes certain problems. These problems will be discussed as they apply to a specific species or vegetation zone.

The Shannon-Wiener diversity index is defined as:

S H *» - X (pi)(log2 p4 > 1 = 1

H ** information content of sample (bits/individual) ** index of species.diversity.

S = number of species.

P£ “ proportion of total sample belonging to 1 th species. 81

The question as to why this diversity index is a better measure­ ment than diversity for expressing the number of species for a commun­ ity is illustrated by MacArthur and MacArthur (1961: 594-598). They used this example. A one species community has a diversity index of zero. A community with two species naturally has a higher diversity, but in proportion to the number of each species present. If n^ ™ 99 and n2 “ 1 then the diversity index (H) would be 0.056 which is close to zero. However, If n^ *= 50 and n2 ** 50 then the diversity index (H) would be 0.694. The second example is also the maximum diversity index possible for a two species community since the species are evenly distributed. The first example is the minimum diversity index pos­ sible for a two species community since it is closest to a one species community. If only the number of species found were used, the differ­ ence between the two communities would not be detected. When dealing with many species in a community, this diversity Index can make It possible for an investigator to compare the fauna of various communi­ ties by a single reading. Pielou (1966a) cautions that the Shannon- Wiener diversity index is to be used only when the population is too large for all the members to be counted and that the number obtained by this method is only an estimate.

The most useful paper for actually calculating the diversity Index is by Lloyd, Zar and Karr (1968). This paper presents a working formula and a table of useful functions used in the formula. The working formula I used is:

H ° - I Pi Log2 Pi 55s- ^ (N LoglO N -■ Log1 0 ni)

c *» a conversion factor of 3.321928 which Is used to change logarithms from a base of 1 0 to a base of 2 .

N = the total number of specimens found in the sample.

n^ “ the number of specimens found for each species.

Eni ** N

In order to help clarify to someone not familiar with these func­ tions, I will present an example.

Species Collected Number Collected Log^n^ n^Logionl

Species A (n^) 10 1.0 10.0 Species B (n2 ) 20 1.30103 26.0206 Species C 0 *3 ) 30 1.47712 44.3136 Species D (n4 ) 40 1.60206 64.0824 Total Collected (N)«* 100 Zn^Logn^ • 144.4166

Log10N « 2.0 N togi0N - 200 82

H « It-3.2.1.?2 .8 (200 - 144.4166) =■ .0321928 (55.5834) = 1.7894 100

The maximum diversity index (Hmax.) for the four species A, B, C, and D would be 2.00. This can be calculated by assuming that all species were found in equal numbers. This maximum diversity index is only useful as a reference point. A sample with an index reading of 1,7894 with four species has a more equal distribution of the species than a sample with 10 species and an index reading of 1.7894. The maximum diversity index for 10 species is 3.3219. Graph I has a maxi­ mum diversity index line (line A) plotted to show its relationship to the actual indices calculated for the various zones in the study area.

What does all this mean when comparing one community with another? Unfortunately I cannot answer that question completely. Krebs (1972: 536-537) mentions that six hypotheses have been proposed to explain variations in species diversity. Some of these hypotheses do not apply to an area such as Cedar Bog; One hypothesis proposes that species diversity increases as the number of habitats available per unit of area increases. Another hypothesis links environmental stability with diversity, as the stability increases, the diversity increases. A third hypothesis suggests that high productivity results in a high diversity. There may be a great deal of truth to all of these hypotheses although no one has used terrestrial mollusca to confirm them. Odum (1963: 34) states that the highest diversity usually occurs in areas where the physical factors are moderate. Odum also supports the belief that a high diversity is of survival value to the community by increasing its stability. Goulden (1969: 96) studied cladocerans in various lakes throughout the world. He found that the diversity index remained high as long as the lake was undisturbed. When any changes in the lake occurred the index decreased, but the number of species or diversity remained the same. This supports the belief that a stable environment is Important to a high diversity index or vice- versa. McNaughton (1974: 351-353) studied the energetics of plants in an old field. He found that the diversity index increased as the succession progressed, but the net productivity of the community re­ mained the same. Successional communities are less stable than climax communities. MacArthur (1965: 531) believes that the diversity index derived from the information theory can be used to measure habitat diversity and the differences between communities or habitats. When changes occur in the pattern of species diversity these changes can be compared with changes in the environment. Lloyd et al (1968: 261) states that in the general view, the average species diversity index is an intrinsic parameter of a whole community.

Habitat Diversity Index

In addition to comparing various communities through the use of the diversity index, one can also compare the habitat breadth of each species by using the same data in a different way. Lloyd et al (1968) 83 and Getz (1974) compared the breadth of habitats for birds and ter­ restrial molluscs in various plant communities studied. The higher the index the more habitats a particular species is found in. Again, the evenness of the distribution of each species within each community is as important as the number of different communities in which it is found.

The habitat diversity index can be calculated the same way the community diversity index is calculated. Instead of measuring how many species of molluscs are present in the community and how evenly they are distributed in that same community, the data are arranged to show the number and evenness of each species in each community.

I grouped the habitat diversity indices into categories, 0 to .99 is low; 1.00 to 1.99 is medium; 2.00 to 2.99 is high; and 3.00 to 3.99 is very high. A low habitat diversity index can mean several things. Usually it means the species has a specialized habitat niche and occurs in a narrow range of habitats being studied. However, it could mean the species is rare and there were not enough specimens collected to present an accurate comparison. A high or very high habitat divers­ ity index means the species has a rather wide habitat niche and is found In many different habitats in relatively even densities. Natur­ ally there is a continuum from zero to the maximum habitat diversity index possible for a given set of habitats.

Only the quantitative quadrat data were used to figure these habitat Indices. Species not usually found in the soil and litter will not have an accurate Index. I calculated indices for all the species and they are listed In descending order as they were figures by using the total number collected in the vegetation zones made by Frederick (1974). These data are found in Table 10. DISCUSSION OF THE SPECIES COLLECTED AT CEDAR BOG

General Remarks

Each molluscan taxon collected at Cedar Bog will be discussed. I will give its current known distribution and review its general ecology as it is known from the literature. I will make remarks as to its taxonomic status only if there are changes from what is generally accepted for the Ohio fauna. Finally, I will discuss its occurrence in Cedar Bog. One very important point concerning my reference to vegeta­ tion zones. The original collection scheme was designed around Dr. Clara Frederick's dissertation of 1967. All the distribution maps have outlines of her vegetation zones. In my discussion of where each species is found, I will refer to vegetation zones that she outlined in 1967 and refined in 1974. Only occasionally will I make reference to the Environment Control Corporation (1973) zonatlon. Usually only when the habitat diversity Indices are greatly affected by the differ­ ences caused by the two interpretations.

Valvata tricarinata (Say, 1817)

Distribution: "New Brunswick to Virginia, west in the St. Lawrence and Upper Mississippi River drainage areas to Iowa and Nebraska, north in the Canadian Basin to James Bay and Hudson Bay, northwest within treeline to the mouth of the Mackenzie River, and west to Alberta" (Clarke, 1973: 237). LaRocque (1968: 367) records V. tricarinata widespread in Ohio both living and fossil.

General Habitat

In the Canadian Interior Basin V. tricarinata Is characteristically associated with large perennial bodies of water (Clarke, 1973: 237). It is also abundant in subarctic muskeg pools. Clarke found them in waters with all types of bottoms, usually with vegetation, and where the current, If any, was moderate to slow.

H. B. Baker (1922: 30) in upper Michigan found this snail most abundant along lake shores with a sandy marl bottom. LaRocque (1968: 367) states that V. tricarinata is "found in shallow water to depths exceeding 9m.; in lakes and streams with or without vegetation; on sand and gravel, rock, clay and mud bottom." Leonard (1950: 11) reiter­ ates the fact that this species lives in permanent bodies of water where the vegetation is abundant.

84 85

M A P V III Collection locations for Valvata tricarinata (Say) at Cedar Bog.

1— 'vT"Tr I

D e c . 23 & 2 5 , 1968 Jan. 5, 1969 • Collected Alive ® C ollected D ead

April 23 & 25, 1969 A C ollected Alive d C ollected D ead

August 20 & 30, i969 ■ C ollected Alive 0 C ollected D ead

Miscellaneous C ollections © C o lle c te d A liv e © C ollected D ead

Woarfhufn. BoccL. 86

Occurrence at Cedar Bog

Only three dead specimens of Valvata tricarinata were collected at Cedar Bog. All the specimens found were along the West Branch of Cedar Run. I am quite certain that these specimens were dredged from the bottom of Cedar Run many years ago. I found no evidence that this species is currently living in the West Branch or any other part of Cedar Run today.

Valvata sincera Say, 1824

Distribution

Most of Canada and the United States north of 40° north latitude. It is recorded by Sterki (1907: 387) as Valvata lewisi Currier, 1868 from Summit County, Ohio. Valvata sincera is recorded from Ohio, but it is not recorded from any particular place.

General Remarks

F. C. Baker (1928), Leonard (1950) and LaRocque (1968) refer to this species as Valvata lewisi. However, Clarke (1973: 225) states that V, lewisi is a smooth variant of V. sincera. Clarke has examined specimens from the type locality and definitely considers V. lewisi as a synonym of V. sincera.

General Habitat

In Wisconsin, V. sincera (as V. lewisi) Is found in shallow water vegetation where there Is a sandy bottom (F. C. Baker, 1928: 28). Leonard (1950: 11) states that "Valvata lewisi inhabits ponds and lakes, especially the latter, living in water little more than 1 meter in depth, crawling about on the mud or on aquatic vegetation."

Clarke (1973: 223-224) referring to Valvata sincera, usually associated the species with permanent bodies of water, either lakes or rivers, where the substrate was sand and vegetation was present. He did collect it a few times in small swamp streams and muskeg pool.

Occurrence at Cedar Bog

Valvata sincera was never collected alive at Cedar Bog. Three out of the four times it was found were in soil samples taken along the West Branch of Cedar Run. These specimens were dredged up from the West Branch some time ago. However, its presence in a soil sample taken in the middle of the Bog is a little puzzling. There is some evidence that Cedar Bog was once a lake or at least ponded for a time after the 87

MAP IX Collection locations lo r Valvata sincera Say at Cedar Bog.

I s " I -T u | ~ w r *

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

A p r i l 23 & 2 5 , 1969 * Collected Alive a C ollected D ead

August 20 & 30, 1969 ■ C ollected Alive o C ollected D ead

Miscellaneous Collections Q C ollected Alive @ C ollected D ead

ter. >230". 88

Wisconsin glacier left. These specimens could be fossils from that time. All the specimens collected were old and I found no evidence that the species is still living within the Bog today.

Amnicola limosa (Say, 1817)

Distribution

"New England and New Jersey west to Utah, Manitoba south to Texas" (F. C. Baker, 1928: 97).

General Habitat

"Amnicola limosa has a wide range in distribution and is found in creeks, rivers, and fresh and brackish water lakes. It Is most abund­ ant where there are thick beds of Chara, Potamogeton. Valljsneria and Elodea. These plants, however, are not used as food by the amnicolid, but they do harbor rich colonies of diatoms that the snails eat" (Berry, 1943: 26). H. B. Baker (1922: 31), F. C. Baker (1928: 97) and Leonard (1950: 12) found that Amnicola limosa lives in permanent bodies of water and In shallow water, where vegetation Is generally abundant. This species can be found in waters with a wide variety of substrates, but many investigators (R. B, Baker, 1911: 170, F. C. Baker, 1928: 97, Goodrich and van der Schalie, 1944: 299 and Clarke, 1973: 259) associ­ ated it with mud, Clarke (1973: 259) found It in waters where the current was imperceptible to moderate.

Clarke (1973: 259) summarizes this species habitat by stating that it can be found In "all unpolluted, permanent aquatic habitats where macroscopic aquatic vegetation grows."

Occurrence at Cedar Bog

Amnicola limosa Is another aquatic species I did not find living at Cedar Bog. I did find it living in the aquatic vegetation in Cedar Run at the Dallas Road bridge south of the study area. X did find dead specimens of this species in the soil along the West Branch, in the West Branch at the Woodburn Road bridge, and in several pools and springs in the arbor vitae, bog meadow and tuliptree vegetation zones. There Is a good possibility that this species still lives within the Bog, either in the West Branch were there Is plenty of aquatic vegeta­ tion or possibly, in some of springs and pools bordering either branch of Cedar Run.* M A P X

Collection locations for Amnicola limosa (Say) at Cedar Bog.

w

D e c . 23 & 2 5 , 1968 Jan. 5, 1969 o • Collected Alive ° C ollected D ead

A p r i l 23 & 2 5 , 1969 A C ollected Alive G C ollected D ead

August 20 & 30, 1969 ■ C ollected Alive o C ollected D ead

Miscellaneous C ollections ©Collected Alive © C ollected D ead

Weaibutn_6ocd,

220

r 5 u X ZV 90

Pomatlopsls lapidaria (Say, 1817)

Distribution

New York west to southern Minnesota; south to eastern Oklahoma in the west; southwest from New York to North Carolina; northern Georgia and Alabama in the east. It is absent from Mississippi, Louisiana and South Carolina (Dundee, 1957: 7).

General Habitat

"Pomatiopsis lapidaria is distinctly an amphibious snail, though seemingly in many localities to prefer wet ground to actual immersion in water'* (F. C. Baker, 1928: 166). Baker continues to mention that he had not personally collected lapidaria under water, but he had found it under leaves and damp or wet mud in areas subject to overflow. Dundee (1957) has made the most extensive study of this species, particularly in Michigan. Many aspects of this species' ecology, life history and morphology are discussed. Dundee (1957: 11-12) states that _P. lapidaria is found in a number of different habitats such as marshes, wooded flood plains along small streams, grassy hummocks in low, wet pastures and in upland artesian-fed marshes. She summarizes: "The features common to all P^. lapidaria habitats seem to be: (1) a very moist substratum with enough sand to prevent it from becoming muck, (2) shade, (3) fresh-water."

Shimek (1930: 681) made the remark that ]?. lapidaria should not be referred to as a fresh-water snail, but is "strictly terrestrial." F. C. Baker (1930: 495) took exception to Shimek*s comments and stated, "Pomatiopsis lapidaria is not wholly and exclusively a land mollusk as maintained by some conchologists, but is truly an amphibious species, living immersed in water at different times and in different places. In the same habitat it may be found in water in the spring and under leaves or wet places in summer and fall."

Leonard (1959: 35-37), in Kansas, found this species living in a marsh on the ground and on the stems and leaves of sedges, reeds and cattails. Berry (1943: 60), in Kentucky, collected it 500 feet from water on a mountain side. In Indiana, Goodrich and van der Schalie (1944: 301) found it under wet logs, in grass, in flood-plain debris and around pools on mud flats.

Van der Schalie and Dundee (1955: 120) also summarized some aspects of the ecology of P^. lapidaria.

1. It is amphibious and confined to wet marginal regions

2. It can withstand submergence In water during high water periods., 91

3. It can occur in large numbers in optimal conditions.

4. It can withstand drought and over winter well.

5. It occurs In Typha swamps, and grassy hummocks In low pastures.

Occurrence at Cedar Bog

Pomatiopsis lapidaria is an occasional species in many areas of Cedar Bog. This snail was not collected in either branch of Cedar Run, but it was taken In the mud along the banks of the West Branch. Host of the bank area bordering the East Branch is not suitable for this species. The most common habitat for P_. lapidaria Is in the pools, springs and seasonal ponds of the wet vegetation zones (see Table 9). Pomatiopsis lapidaria was not found at all In the beech-maple and shrub zones (Map XI). It is most often found in the Carlisle muck, marl and peat substrates. The distribution of this species at Cedar Bog is associated with the presence of standing or flowing water at some time during the year. Map XI shows that most collections are concen­ trated In three general areas, along the West Branch, in the seasonal ponds of the swamp forest and in the marl meadow where water is always present. Living specimens are found in over 20% of all collections made in the maple-ash-basswood swamp forest, the bog meadow and the marl meadow (Plate I, Figure 1). The habitat breadth measured by the Shannon-Wiener formula Indicates a fairly restricted habitat prefer­ ence for the living specimens collected, but a rather wide habitat preference when considering all the areas where empty shells were found (Table 10). Moisture conditions at Cedar Bog have changed. Most aquatic and terrestrial species which live In very wet habitats are less extensive now than they were before the dredging occurred.

Oxytrema semicarlnata (Say, 1829)

Distribution

"Tributaries of Ohio River, Scioto River, Ohio, to Big Blue River, Indiana; Licking River to Salt River in Kentucky; two creeks of Green River of Kentucky" (Goodrich, 1940: 18).

General Remarks

Originally, I listed this species as Goniobasis llvescens (Menke, 1830) for Cedar Bog (Clark County Audubon Society, 1972: A4). However, based on distribution records given by Goodrich (1920: 73-74) this species should be called Goniobasis semicarlnata. 92

MAP XI l4 Collection locations for Pomatiopsis lapidaria (Say) at Cedar Bog.

D ec. 23 St 25, Jan, 5, 1969 C ollected A live f t ° C ollected D ead

April 23 St 25, 1969 C ollected Alive d C ollected D ead

August 20 St 30, 1969 C ollected Alive ° C ollected D ead

Miscellaneous Collections G Collected Alive 0 C ollected D ead

W o adhuifl-JLa c d TABLE 9

Aquatic molluscs liv in g in and around the Cedar Bog Nature Preserve, Champaign County, Ohio.

Species Collected East Branch West BranchCedar Run Temporary pond Pools and Springs or OT 60UXD Or WGst of Cedar Run Cedar Run Cedar Bog Cedar Bog Arbor Vitae Bog Meadow Marl Meadow Maplo-Asfv- T ulip tree OaM-Maple Basswood

Amnicola limosa - D A - 0 D - - D 0

Pomatiopsis la p id a ria - --A A A A AA • Oxytrema semicarinata AAA -A - - - -

Lymnaea caperata - - A A --- - -

Lyranaoa humilis - A -A A AADA

Gyraulus parvus - 0 A - D D A 0 D 0

Physa inteqra AA A- 0 A A A 0 A

Pisidium compression -A- - - - - D - -

Pisidium cascrtanum A A A - A 0 AA D A

A Found living at one time or other. D Only found dead. «• Not found at all.

vo u> PLATES I - XII

— Total Frequency £55 fMi - Living Frequency

Vegetation Zones

A Marl Meadcw Association

B Bog Meadow Association

C Tuliptree Swamp Forest Segregant

D Maple-Ash-8asswood Swamp Forest Segregant

E Arbor vitae Association

F Oak-Maple Forest

G Elm-Maple Swamp Forest Segregant

H Silven-Red Maple Swamp Forest Segregant

J Shrub Communities

K Beoch-Maple Association 95

PLATE I

ICO 100 - ?0 JO « 80 00 - ai 70 7 0 - o* 5* Go- cr> 6o Cl «■ 50 ” £ 50 40 ^ 40 - 30 30 - 20 20 -

10 i&i 10 - '•-f. ■T+‘. ‘jtzi**.»* 0 *?•S3::' S-vnl 0 'M .r.y; A 8 C 0 E F G H i) E r 6 H K Vegetation Zones Vegetation Zones

Figure 1. Collection frequencies for Figure 2. Collection frequencies for Pomatiopsis lapidaria Lymnaea humilis

100 100

80 CO

VI y cX> 3 Qtr t- 40 u.

K C 0 CFGHJK Vegetation Zones Vocptxtion Zones

Figure 3. Collection frequencies for Figure 4. Collection frequencies for Gyraulus parvus Physa integra 96

Morrison (1954: 359) states, "Oxytrema Rafinesque is the earliest and correct name for one of the most widespread 'Melanian* genera in the world. This genus includes numerous North American species whose ranges extend from the Atlantic to the Pacific coasts and from southern Canada to Florida and Texas." "All the species called 'Pleurocera' by Bryant Walker, and other authors who followed him blindly, and all the species called 'Goniobasis1 (with very few exceptions) belong to this genus." Hargreaves (1971: 39) personally communicated with Morrison as to the status of Goniobasis semicarinata and Morrison called it Oxytrema semicarinata based on its egg-laying characteristics.

General Habitat

There is very little specific ecological data available for this species. La Rocque (1968: 425) found it to be common in many small streams and creeks in southern Indiana. In my experience, many mem­ bers of the family Pleuroceridae in easthern North America live in moving water, usually highly oxygenated and unpolluted.

Occurrence at Cedar Bog

Oxytrema semicarinata is found in both branches of Cedar Run. It is found on the soft organic sediments in the quiet waters as well as the gravel bottom and submerged cedar logs. Its presence in Cedar Run is to be expected since these waters are cool, unpolluted and highly oxygenated. I found it living in a small spring in an arbor vitae stand (Map XII, X-7). The spring was very small and contained no firm substrate. Several dead specimens were found in the soil samples taken along the dredged West Branch. This species survived the dredging.

An interesting difference occurs between the East and West Branch specimens. The East Branch shells are heavily encrusted with calcium carbonate deposits and the West Branch shells are clean. This deposit is probably caused by the algae growing on the shells and the super­ saturated state of calcium bicarbonate in the East Branch. This algae changes the pH balance in the immediate area by taking in carbon dioxide. This in turn causes calcium carbonate to precipitate out of the supersaturated waters of the East Branch.

Lymnaea caperata (Say, 1829)

Distribution

"Lymnaea caperata occurs In a broad belt across North America, from Yukon Territory and James Bay, on the north, to Maryland, Indiana, Colorado and California, on the south" (Leonard, 1950: 14). MAP XU CoUection locations for Oxytrema semicarlnata (Say) at Cedar Bog.

Dec. 23 fit 25, 1968 Jan. 5, 1969 o • C ollected A live ° C ollected D ead

April 23 fit 25, 1969 A C ollected Alive A C o lle c te d D ead

August 20 fit 30, 1969 ■ C ollected Alive D C ollected D ead

Miscellaneous C ollections ©Collected Alive © C ollected D ead

W otulbuU-Jacd.

230 440

-- r - . --- j.. s u X VV z 98

General Habitat

F. C. Baker (1911: 233) states that Lymnaea caperata Is one of the most distinct and uniform American lymnaeids and also one of the com­ monest lymnaeids in America.

"In the Mississippi Valley this species seems to almost invariably occupy intermittent streams or small pools, ponds and ditches which dry up in the summer. In Illinois it is usually found in association' with Aplexa hypnorum and Sphaerium oceidentale, either in small streams, pools or sloughs, or in spring pools In the woods which become completely dry in late spring and summer" (F. C. Baker, 1911: 233).

Goodrich (1932: 51-52), Goodrich and van der Schalie (1944: 284), Robertson and Blakeslee (1948: 56), Leonard (1950: 14) and Clarke (1973: 315-316) concur with F. C. Baker concerning the habitat prefer­ ences of Lymnaea caperata. Clarke (1973: 315-316) also collected this species in Canada in some permanent lakes and rivers. He usually found L^ caperata associated with muddy bottom and aquatic vegetation or flooded grass.

Occurrence at Cedar Bog

Actually, Lymnaea caperata was not collected in the study area, either dead or alive. However, this species is found in the Cedar Bog Nature Preserve. Within the preserve I found it in a small tempor­ ary pond in an old field west of the study area. I also found it in Cedar Run at the Dallas Road bridge. It may be present in the study area in a seasonal swamp forest pond, but it would have to be quite rare. If it existed in any numbers dead shells would most probably have turned up in the soil samples at one time or another. I do not believe it exists in either branch of Cedar Run within the study area. The nature of Cedar Run at the Dallas Road bridge Is very different from Cedar Run in the study area. Cedar Run at the Dallas Road bridge had a muddy bottom and a great deal of aquatic vegetation. This fits the Clarke (1973) description very well.

Lymnaea humills Say, 1822

Distribution

Canada, most of the United States except the southern coastal plain and Florida (Hubendlck, 1951: 126-128).

General Remarks

I am In doubt about the exact identity of this species. By fol­ lowing F. C. Baker (1928: 289), La Rocque (1968: 469) and Clarke (1973: 276) this species should probably be called Lymnaea modlcella 99

(Say, 1825). If I follow F. C. Baker (1911: 259), Goodrich and van der Schalie (1944: 285) or Leonard (1959: 53) it should be referred to as Lymnaea humllls modicella and/or Lymnaea humilis rustica. Rubendiqk (1951: 126-128) calls it Lymnaea humilis and believes that his species has a lot of variation. I am following Rubendick at this time because the Cedar Bog lymnaeids appear to show many intergraded shell charac­ teristics. I believe there is only one species in this complex. Jon Williams (1966) collected a few aquatic mollusks at Cedar Bog while studying the aquatic insect fauna. He sent this specimens to Dr. Henry van der Schalie at the University Michigan. Van der Schalie identified them as Lymnaea humilis.

The Lymnaea catascopium I reported in the list published in Cedar Bog Report II by the Clark County Audubon Society (1972: A4) is in error. L. catascopium has not been recorded from Cedar Bog.

General Habitat

The ecological notes for this species have been taken from publi­ cations which refer to species under the names Lymnaea(Fossaria) humilis, Lymnaea(Fossaria) modicella and Lymnaea(Fossaria) rustica.

F. C. Baker (1911: 266; 1928: 290) and Goodrich and van der Schalie (1939: 15; 1944: 285) found that Lymnaea(Fossaria) modicella was usually found along the moist mud flats or mud beaches along streams, rivers and small lakes. F. C. Baker (1911: 270; 1928: 292) states that L. rustica had a habitat preference similar to L. modicella. Clarke (1973: 278-279) collected _L. modicella from several diverse types of waters from small pools and streams to large lakes and rivers, but chiefly where there was a mud bottom and aquatic vegetation.

Leonard (1959: 53), in Kansas, associated L. humilis rustica with moist ground on the edge of or on the mud bottom of shallow streams. F. C. Baker (1911: 258) found L. humilis In a tide meadow ditch In New Jersey and vacant lots and ditches in South Carolina.

Occurrence at Cedar Bog

As can be seen on Map XIII, Lymnaea humilis Is found predominantly in the vegetation zones around the two branches of Cedar Run. I never found L. humilis common in the streams proper, but in the many springs and pools within the vegetation zones surrounding Cedar Run (Table 9). This species is common in the marl meadow. It is also frequent as dead shells in several other vegetation zones. The percentage of liv­ ing specimens in most collections is fairly low, yet it appears to be one of the characteristic species of the marl meadow and springs in several other zones. The habitat breadth of this species is fairly low (Table 10) owing to its dependence on a fairly wet environment. The 100

MAP XIII Collection locations for Lymnaea humilis (Say) at Cedar Bog.. I

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

A p r i l 23 & 2 5 , 1969 A C ollected Alive A C o lle c te d D ead

A u g u s t 20 & 3 0 , 1969 ■ C ollected Alive tJ C ollected D ead

Miscellaneous Collections G C ollected Alive © C ollected D ead

W n m l k i r n D

to

H

n lim.altOll 101 marl within the Bog has a nature much like fine mud and this species seems to be occupying the same type of environment as recorded by other investigators.

Helisoma anceps (Menke, 1830)

Distribution

’’Canada and United States, east of the Rockies, f.rom about the latitude of Hudson Bay south to Western Mexico" (La Rocque, 1968: 500),

General Habitat

Helisoma antrosa (Conrad, 1834) and Helisoma anceps (Menke, 1830) are synonymous (Clarke, 1973: 428). F. C. Baker (1928: 319) stated that in Wisconsin "antrosa is primarily a river and creek species, not living in the large lakes.” Goodrich (1932: 62) found that in Michi­ gan, H. antrosa occurs in lakes and streams and can withstand heavy currents. Goodrich and van der Schalie (1939: 15-16), in the Upper Peninsula of Michigan, found H. anceps in brooks, creeks, rivers, lakes and ponds. Archer (1939: 15), also in Michigan associated this species with roots and decaying vegetation, open pools and woodland pools. Leonard (1950: 16), in Kansas, even collected it in temporary ponds. Clarke (1973: 431-432), in Canada, found li. anceps in all types of permanent aquatic habitats, in all types of bottoms, usually with vegetation.

La Rocque (1968: 500) states, "The typical form lives in rivers and creeks as do a number of more or less recognizable varieties; num­ erous varieties have been named which live in lakes of various sizes.” The early investigators associated the typical subspecies with a specific habitat and later their concept on this species widened and so did its habitat preferences.

Occurrence at Cedar Bog

Helisoma anceps has not been found living in the Bog. Dead shells have been found in soil samples in several areas, mainly along the West Branch of Cedar Run in the stream dredgings (Map XIV). It is very possible that H. anceps is still living in Cedar Run south of the Bog. The presence of dead specimens in areas far removed from the West Branch Indicates that the aquatic habitats have changed somewhat at Cedar Bog. More than likely these changes took place since the dredging which occurred around 1912. MAP XIV Collection locations for Helisoma anceps (Menke) at Cedar Bog.

Dec. 23 & 25, 1968 Jan. 5, 1969 o • C ollected A live o C ollected D ead o

April 23 & 25, 1969 * C ollected Alive 6 C ollected D ead

August 20 & 30, 1969 ■ C ollected Alive ° C ollected D ead

Miscellaneous Collections G C ollected Alive © C ollected D cad

WoaafctLun-JLoed.

r — ;— r — r~"T— r— a— r— v— l— w— t ~ '* — T — 1— f 103

Gyraulus parvus (Say, 1817)

Distribution

Most of North America north to the treellne In Canada; Cuba (Taylor, 1960: 58 and Clarke, 1973: 403). It Is widespread throughout Ohio (La Rocque, 1968: 491-492).

General Habitat

F. C. Baker (1928: 376) usually associated Gyraulus parvus in Wisconsin, with small quiet bodies of water. In Michigan, it has been collected by several different investigators in many habitats. Goodrich (1932: 65) found it common in sluggish waters; Goodrich and van der Schalie (1939: 18) in lakes and rivers; H. B. Baker (1911: 166) in a perennial swamp; Kenk (1949: 54) in temporary ponds; and Archer (1939: 16) in bog-lake pools and tamarack-bog pools. All men­ tion its close association with dense aquatic vegetation. In Indiana, Goodrich and van der Schalie (1944: 288) found it common in ponds, lakes and slow-moving streams and in Kansas, Leonard (1959: 61) found it common in all kinds of lakes, quiet pools and stream beds. Robert­ son and Blakeslee (1948: 63), in New York, collected it in small pools. In Ohio, Dexter (1953: 29) collected Gyraulus parvus in all types of ponds. La Rocque (1968: 491) generally associated (3. parvus with small, quiet, shallow bodies of water which have diverse sub­ strates. Clarke (1973: 403) frequently collected parvus in the Interior Basin of Canada. He found it in many diverse aquatic habi­ tats, but almost always where there was a substantial amount of aquatic vegetation and the current, if any, was slow to moderate. He found it 60% of the time in habitats with a mud bottom.

Occurrence at Cedar Bog

I found dead shells of Gyraulus parvus in every vegetation zone at Cedar Bog except the beech-maple forest (Plate I, Figure 2). Its dis­ tribution is much the same as that of Lymnaea humilis, but I only found it living in four places in the study area, all of them in or near the marl meadow. This species was apparently fairly common throughout the Bog at one time. The habitat preferences for this species appear to be broad enough to enable it to live in the springs or pools and particu­ larly in the West Branch of Cedar Run. However, I did find it living in Cedar Run at the Dallas Road bridge, south of the Bog. 104

MAP XV Collection locations for Gyraulus parvus (Say) at Cedar Bog.

} i w t x i r

D ec . 23 8c 25, 1968 Jan, 5, 1969 • Collected Alive ° Collected Dead

A p r i l 23 8' 2 5 , 1969 A C ollected Alive & CollectedDead

August 20 & 30, 1969 ■ Collected Alive ° C ollected D ead

Miscellaneous C ollections 9 C ollected Alive © C ollected D ead

Wflrtdhurtl -I

TO 10

ta Itrrv -a a o u I V 105

Planorbula armigera (Say, 1818)

Distribution

New England west to Nebraska, south to Georgia and Louisiana,, north to the Great Slave Lake (F. C. Baker, 1928: 354)*

General Habitat

La Rocque (1968: 507) summarizes the findings of many investi­ gators concerning the habitat of Planorbula armigera. "This is a species of small stagnant bodies of water, found even in marshy areas, in disconnected pools of water. It seems to prefer mud bottoms but can accommodate Itself to a bottom of meadow grass and can also live on logs. It seldom lives in water more than three feet deep."

Clarke (1973: 422), Canada, found P. armigera in muskeg; Archer (1939: 15), in Michigan, in many kinds of pools including bog lake pools; Kenk (1949: 54), also in Michigan, found it in temporary ponds; and Dexter (1953: 29), in Ohio, collected it in pasture and woodland pools.

Occurrence at Cedar Bog

I found one dead specimen of Planorbula armigera in the bog meadow near the East Branch of Cedar Run. Williams (Table 9) also found two adult specimens in the same area of the East Branch. He did not mention whether his specimens were dead or alive. It is possible that armigera is still living in Cedar Run in small numbers.

Physa Integra Haldeman, 1841

Distribution

New York and Ontario west to North Dakota, Wyoming and Colorado; southeast to Texas and Louisiana in the west and south from New York to Virginia in the east. Except for the Virginia records, it appears as if the distribution of this species follows the Mississippi River and the St. Lawrence River drainages (La Rocque, 1968: 547). La Rocque also records it from twelve counties in the southern half of Ohio.

General Remarks

Doctor William Clench identified most of the specimens collected at Cedar Bog as Physa Integra. He also mentioned that they appeared most like Physa michiganensis Clench, 1926 which he now feels Is a form of Physa Integra. However, several investigators have collected MAP XVI Collection locations for Planorbula armigera (Say) at Cedar Bog.

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live ° Collected Dead

A p r il 23 & 2 5 , 1969 A Collected Alive A C ollected D ead

August 20 & 30, 1969 ■ C ollected Alive 0 C ollected D ead

Miscellaneous Collections Q C ollected Alive © C ollected D ead

Wood hum toed 107

Physa michiganensis and have recorded its specific hahitat preferences. Clench (1926: 4-6) originally collected P^ michiganensis In small creeks at the edge of the quiet water under leaves. Goodrich (1932: 71-72), Goodrich and van der Schalie (1944: 291) and Robertson and Blakeslee (1948: 78) found ]?. michiganensis in small creeks and brooks.

General Habitat

Clampitt (1970: 132) studied Physa Integra in a lake in Iowa. He found that this species was characteristic of rocky, algae covered, cobble shores where the wave action was slight to heavy, and in vegetated offshore areas where the depth varied from 50 cm to 3 m and the substratum was sand or mud. Goodrich and van der Schalie (1944: 291) found it in Indiana in streams. Goodrich (1932: 71-72), in Michigan, usually associated it with brooks, creeks, and rivers, but occasionally in lakes and the quiet waters of the Great Lakes. Robertson and Blakeslee (1948: 78) found it in New York in streams and ponds.

La Rocque (1968: 548) summarizes the habitat preferences of Physa Integra. It is "found on a variety of bottoms (mud, sand, gravel, boulders, clay) in lakes, bays of the Great Lakes, and streams, in water less than 3 inches to 6 feet or more deep. A species widely adaptable to different environments and depths."

Occurrence at Cedar Bog

The distribution of Physa integra parallels that of Lymnaea humilis and Gyraulus parvus (Map XVII). I have found it living In several areas including both branches of Cedar Run, many temporary pools, springs and in the marl meadow (Table 9). Ten to fifteen per­ cent of the samples taken in the bog meadow, marl meadow and oak-maple zones contained living Physa integra (Plate I, Figure 4).

Summary of the Aquatic Gastropods of Cedar Bog

I have found evidence of ten aquatic gastropods which have lived or are now living In Cedar Bog. Only five species have been found living. I personally feel that this number is low. There are several quite diverse aquatic habitats within the Bog. These are seasonal woodland ponds, permanent springs, pools in the marl meadow, and several diverse areas In both branches of Cedar Run. I did search the stream habitats, but not as extensively as I did terrestrial habitats. It is possible that several species which I could only find dead, are living in Cedar Run today.

In general, the distribution patterns of the five living species are fairly easy to explain. Lymnaea humilis, Gyraulus parvus and 108

MAP X V II

Collection locations for Physa integra Haideman at Cedar Bog.

Dec. 23 &c 25, 1968 Jan. 5, 1969 o • C ollected A live ° C ollected D ead

A p r il 23 & 2 5 , 1969 * Collected Alive A C o lle c te d D ead

A u g u s t 20 & 3 0 , 1969 * C ollected Alive ° C ollected D ead

Miscellaneous Collections © Collected Alive © C ollected D ead

V/o ttdLbiXLO— R d c d.

TU

*40

1cm. *J J0h 109

Physa Integra are common near the constant sources of water such as the springs and streams. These numerous springs are found in the oak- maple, bog meadow, marl meadow and arbor vitae vegetation zones which also border the streams. Pomatiopsis lapidaria has a little broader distribution pattern because it can also be found in the seasonal ponds in the maple-ash-basswood and tuliptree swamp forest segregants in addition to the spring areas. Oxytrema semicarinata is confined to permanent running water which is found mainly in the streams.

Carychium exiguum (Say, 1822)

Distribution

This species is recorded from most states and providences east of the Rocky Mountains, from southern Canada to southern Texas. It is also found on Vancouver Island and in Alaska (La Rocque, 1970: 560). It is widespread in Ohio, but most current records are from the west- central counties.

General Habitat

Harry (1951) did his dissertation on Carychium exiguum. In Michigan, he found this species to prefer arbor vitae forests, grass­ lands which were formerly arbor vitae forests, arbor vitae-hardwood transitions, grassland-hardwood transitions, and those areas where Populus preominates. H. B. Baker (1922: 26) found exiguum common in ash-arbor vitae swamps in the northern penninsula of Michigan. Smith (1928: 490) found C t exiguum to be quite common in a sugar maple-red oak-elm climax forest in his study area in Illinois. Burch (1956: 61) associated this species with several different plant communi­ ties on the Piedmont Plateau in Virginia, all had oak as one of the dominants. H. B. Baker (1911: 160-161) collected this species along the borders of lakes and swamps behind the dunes with sugar maple, oaks, ashes, alder and dogwoods dominating. He also found them under Isolated masses of decaying vegetation and driftwood along a sandy beach of Lake Huron.

One of the most Important factors governing the distribution of C_. exiguum on the microclimatic level is moisture. Most collectors make note of the wet conditions in which this species is found. Oughton (1948: 95) states that this snail is "found in margins of ponds, streams and marshes; seeping hillsides; also sandy flats which receive water by percolation." F. C. Baker (1902: 255) found them plentiful in the vicinity of ponds and creeks. In fact, he states that they are "almost as frequently found In water as out of It, that is, under logs and stones in such moist localities that the animal is Immersed In water." Goodrich and van der Schalle (1944: 282) state, "This minute snail is to be found under logs in damp places which, In spring, are partially under water." Pilsbry (1948: 1054) and Leonard 110

(1959: 194) also mention this snail's preference for very wet places, around ponds and streams, and frequently submerged. Harry (1951: 17-18) does offer some contradictory information regarding its affinity for water. He states that £. exiguum is not found in areas frequently flooded or on the margins of temporary ponds. He does state that this species requires high and constant moisture throughout the year.

Harry (1951: 141) found that C, exiguum requires an abundance of decaying plant material and darkness. The decaying plant material provides food, shelter and a consistently high substrate moisture. Pilsbry (1948: 1054) states that this species lives in the crevices of rotten logs or in dead leaves. La Rocque (1970: 558) states that it is "found in shady, protected situations, such as in moist dead leaves, in crevices of rotten logs and under sprung bark of dead trees and stumps." F. C. Baker (1902: 255) found it plentiful under and about old logs, sticks and frequently in wet moss. However, Harry (1951: 20) points out that he never found it associates with Sphagnum moss.

Occurrence at Cedar Bog

Living specimens of Carychium exiguum were found in eight of the ten vegetation zones at Cedar Bog. I found no trace of this species in the beech-maple and only dead shells in the shrub zones. £. exiguum appeared in 40% of all the collections made at the Bog. This species must be considered a major constituent of the molluscan fauna of the Bog. The habitat diversity index for £. exiguum is 2.18 for total specimens and 2.38 for living specimens (Table 10). These high Indices show that this species was found in a rather wide variety of habitats within the Bog.

Map XVIII shows that this species is generally distributed over the wetter parts of the swamp forests, marl meadow, bog meadow and along the streams. It is almost totally absent from the drier southern part of the study area. This distribution pattern is similar to that of Pomatlopsis lapidaria.

Even though it is a frequent constituent^ of many vegetation zones, this species definitely shows preferences for the wettest vegetation zones. Carychium exiguum shells appear in 60% of the marl meadow samples, 54% of the tuliptree samples and 52% of the bog meadow samples (Plate II, Figure 1). This graph also shows that this species is found living in a high percentage of the collections made in these three vegetation zones.

In addition to the distribution and frequency data, the average densities for this species indicate a definite preference for wet habitats. The average density of the total specimens collected in the bog meadow was 227.9/ ft^ and for the marl meadow It was 203.7/ ft . M A P X V U I

D ec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live a ° C ollected D ead

April 23 & 25, 1969 A Collected Alive A C ollected D ead

August 20 & 30, 1969 » C o lle c te d A liv e ° C ollected D ead

Miscellaneous C ollections Q C ollected Alive @ C ollected D ead

W eodbuia_Jtiicd_ ti. Frequency (ffi) 100 100 • 100 °-| - 2° - * 80 50 ao 0 3 40 70. 20- 0." - 20 . 20 50. 60 • 10 vJL o 0 - - . - -

Figure 3. Collection frequencies for frequencies Collection Figure 3. Figure Figure i'-y ABCDtfGHJ

. 1 .frequencies for Collection 1

Carychium nannodesCarychium Carychium exiguumCarychium m k . . . Vegetation Zones Vegetation Vegetation Zones Vegetation § i PLATE II PLATE 3 55 . u I i* 5- 5* 0 . 70 70 . 100 100 • 100 40 40 o- go - 0 2 Go 50 20 10 30 40 30 30 80 50 - 60 10 0 7 0 0

-

- - - - - • - - . Figure

Figure

& } ABCDEFGHJ . m 4 2 . Collection frequencies for .frequencies Collection . Collection frequencies . Collection m s.*:-: Gastrocopta contracts Gastrocopta Carychium exile Carychium D F h G F E D C Vegetation Zones Vegetation ,Vr- i-v-'- Vegetation Zones Vegetation $ 4 Wl m m . w W.fj- TV-.':’:' •V a V* i £ vTT V*..‘ ;L'v S“v. £i i£ for 112 •‘Mi i W 'X- i'tX 113

The average density for living specimens in the bog meadow was 7.6/ ft^ and the marl meadow it was 9.5/ ft^. No other area had densities near this high for this species.

A wet habitat does not appear to be essential for this species, but it is frequently collected in the wettest habitats, I collected £. exiguum in many areas where it had to be living under water part of the time. I have also observed specimens crawling over wet logs in a film of water. I found this species in the same habitat that Pomatiopsis, Lymnaea and Physa are found. I find it very difficult to draw a line between the habitats of many aquatic, amphibious and terrestrial mollusca.

Carychium exile H. C. Lea, 1842

Distribution

"Maine and Ontario to Manitoba, south to Mobile, Alabama and Texas" Pilsbry (1948: 1058).

General Remarks

I consider Carychium exiguum and Carychium exile distinct species because when the two species occur together they usually can be read­ ily separated morphologically. Harry (1951) and Burch (1962) consider these two species synonyms because, in Michigan, he found no consist­ ent and significant morphological differences in the' internal anatomy or the shell of what has been called two species. I was usually able to separate these two species without any trouble. In one collection (# 156) there were in my opinion some individuals having intergrading of shell characters.

General Habitat

Carychium exiguum and Carychium exile are frequently recorded to­ gether. F. C. Baker (1902: 256), H. B. Baker (1911: 161), Smith (1928: 490) and Leonard (1959: 195) record both of these species from the same microhabitats. Carychium exile is strongly associated with moist areas whether it is with C, exiguum or not. It has been recorded from a wide variety of plant communities. In Michigan, H. B. Baker (1911: 161) reports it from swamp edges, beaches, mesic ash- maple woods, very wet woods dominated by black oak, swamp maple, elm and arbor vitae, drier woods dominated by populars, birches, beech, elm and maples and also grassy meadows. In Minnesota, P. C. Baker (1935: 273) collected

Michigan. All these Investigators collected exile from the moist areas within the various plant communities mentioned.

Carychium exile appears to have a wider habitat tolerance than does £. exiguum. Sterki (1907: 381) states that Carychium exile "seems to be more in elevated and dry places than exiguum, although the two are found associated." Oughton (1948: 94) places £. exile in damp woodlands composed mainly of deciduous trees, whereas he placed C. exiguum In much wetter habitats along margins of small bodies of water and seeping hillsides.

Hubricht (1964: 34) found C\. exile in seven different caves in Kentucky, Tennessee and Alabama with sufficient numbers to indicate that they lived and bred there.

Regardless of what specific communities C. exile is found in and how wet those communities are, most investigators found this species associated with abundant leaf mould, forest debris, under.old logs, started bark, stumps, saturated moss and roots of ferns.

Occurrence at Cedar Bog

Carychium exile and Carychium exiguum are frequently found to­ gether. From the data I have obtained at Cedar Bog I would definitely say their habitat niches overlap. However, when comparing the dis­ tribution patterns (Map XVIII and Map XIX), these two species are quite different. C, exile is not found in the marl meadow and rarely found in the bog meadow. Within the various swamp forest segregants it is found in the drier sections. This is the only member of the genus which is found living in the beech-maple forest.

When comparing the frequency graphs (Plate II, Figures 2 and 3) of C. exiguum and C. exile, C\ exile is quite different. £. exile is most frequently found in arbor vitae followed by maple-ash-basswood and silver-red maple swamp forest segregants. Shells of this species were found in 58% of the collections made in arbor vitae and living specimens were found in 45% of the arbor vitae samples" exile had a very low frequency in four of the five vegetation zones where C. exiguum was most common. The greatest zone of overlap is in the maple-ash-basswood swamp forest. This understandable because It was composed of many diverse microhabitats.

The habitat diversity index (Table 10) is high In every way In which It was figured. C . exile has rather wide niche requirements and an even disttibution throughout much of the vegetation zones in which it is present.

The average density for C. exile is low except in the arbor vitae association. In the 18 arbor vitae samples In which £. exile w s b collected, it had an average total density of 34.2/ ft^ and an average living density of 14.6/ ft*. 1 1 5

MAP XIX

Collection locations for Carychium exile H. C. Lea at Cedar Bog.

13 ec. 23 & 25, Jan. 5, 1969 • Collected Alive ° C ollected D ead

A p r i l 23 & 2 5 , 1969 C ollected Alive a C ollected 13 ead

A u g u s t 20 & 3 0 , 1969 C ollected Alive ° C ollected D ead

Miscellaneous Collections 9 C ollected Alive © C ollected D ead 9 116

Carychium exile is a common species throughout the Bog, but It Is characteristic of the arbor vitae association. It can withstand a drier range of habitats and appears to be more closely associated to leaf liter substrate than exiguum. Judging from where some of the samples were collected, it can also withstand prolonged immersion.

Carychium nannodes Clapp, 1905

Distribution

Southwestern corner of Michigan, southern Ontario and west- central Ohio, Kentucky, Tennessee, West Virginia, and western half of Virginia and North Carolina, northern Alabama and Georgia (Map LXVIII).

General Remarks

Carychium nannodes has not been previously recorded from Ohio. It was first described by George H. Clapp in 1905 from Madison County, Alabama. Since that time, many records from other states have been made, but the distribution of this species is still somewhat sporadic in the northern part of Its range. It is a very small species (1.4 mm long and ,5 mm wide) and could be easily overlooked by most collectors.

General Habitat

The habitat notes in the literature for C. nannodes are almost non-existent. This species was recorded at the type locality by H. H. Smith as "abundant among leaves in a shady ditch in damp forest near the top elevation 1600 feet" Pilsbry (1948: 1056).

In 1922, Winslow reviewed the internal lamellae of several species of Carychium, including C. nannodes and a C. exiguum form collected in Michigan which she suspected as being £. nannodes. Pilsbry (1948: 1055) and Oughton (1948: 80) supported her belief that the specimens in question were £. nannodes. These specimens were collected "in moist ground and debris of a wooded ravine between sand dunes, near the Lake Michigan beach" (Winslow, 1922: 5).

Occurrence at Cedar Bog

Carychium nannodes was collected in nine litter samples. All the collections north of Woodburn Road contained living specimens and all the collections south of the road contained dead specimens (Map XX). I am not sure whether this is coincidental though. The main differ­ ence between the two sides of the road Is in the degree of past 117

MAP X X

Collection locations for Carychium nannodes Ciapp at Cedar Bog.

z = z o n

D e c . 23 & 2 5 , 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

A p r il 23 &t 2 5 , 1969 * Collected Alive A C o lle c te d D ead

A u g u s t 2 0 L 3 0 , 1969 ■ C ollected Alive d Collected Dead

Miscellaneous C ollections Q C ollected Alive @ C ollected D ead

10

0 320 M O

1cm. .230*1 13 r z : r z i E 118

disturbances. The south side has been disturbed more by nan, farming, and wind. % I found _C. nannodes In only three vegetation zones, the arbor vitae association, the maple-ash-basswood and the tupllptree swamp forest segregants. Six of the nine collections were taken from the arbor vitae association.

Carychium nannodes Is relatively rare at the Bog. This makes it difficult to make many correlations. It Is most often found In the arbor vitae association In the leaf litter, in peat or muck substrates and in areas forested since 1938 (Map VI). I did not find it living in the more disturbed sections of the arbor vitae. The low habitat diversity index indicates a fairly limited habitat and a fairly limited distribution.

Cochllcopa lubrica (0. F. Mtlller, 1774)

Distribution

Holarctic; Throughout Europe, U.S.S.R., North China, Korea, Japan, North Africa, Azores, Madiera and North America (Bequeart and Miller, 1973: 72). In North America It occurs from Alaska to Mexico in the Rocky Mountains; throughout Canada and south to Maryland, Tennessee and Missouri. Cochllcopa lubrica is absent from the Pacific slope, Atlantic Coastal Plain and most of the dry southwestern plains area (Leonard, 1950: 27). It is common and widespread In Ohio (La Rocque, 1970: 770).

General Remarks

Bequaert and Miller (1973: 193) explain their current change to the genus Cochllcopa and Cionella. I am following their reasoning and that of most European malacologists In using the older Cochllcopa and Cochllcopldae.

General Habitat

Cochllcopa lubrica does not seem to be found in the same habitat throughout its wide range. Some investigators find it more in open areas and others in dense shaded areas.

In Russia, Llkachev and Rammel'meler (1952: 126) recorded it liv­ ing in humid places in grass, moss and leaf litter. In Great Britain, Boycott (1934: 14, 22 and 26) frequently found £. lubrica in damp meadows and marshy places, gardens and beech-ash woods growing in oolitic limestone. Atkins and Lebour (1923: 234) made correlations 119 between snail habitats and pH and found £. lubrica occurred in a pH range of 6.0 - 7.5, but most were found in the 6.5 - 7.0 range.

In Ontario, Canada, Oughton (1948: 94-95) occasionally found C. lubrica in deciduous woods, but its chief habitat was the moist to wet areas along streams, ponds, lakes and rivers. Dimelow (1962: 52), in New Brunswick, found it under fallen logs in a marshy meadow which was periodically flooded.

In Michigan, H. B. Baker (1911: 160 and 1922: 29.and 37) collected Cochllcopa lubrica in several locations along a floodplain. Harry (1951: 31) collected It In an arbor vitae swamp and deciduous woods dominated by Populus. Archer (1936: 12-13), also in Michigan, found it preferred leaf mold and rotting logs in shady deciduous woods.

Goodrich and van der Schalie (1944: 280-281) collected it under logs, leaves, stones and in grass from various places in Indiana. In Illinois, F. C. Baker (1939: 120) found it in ravines and woodlands under dead leaves and loose bark. In Kansas, Leonard (1950: 25) found it under leaves and decaying wood in damp, densely shaded areas.

In the southeastern part of its range in North America* several investigators found Its habitat preferences a little more variable. Grimm (1959: 126) collected It In Maryland along railroad tracks, In fields, and In the debris around the ruins of a house. In New York, Ingram (1946: 91-92) found £. lubrica typically under fallen fence rails and old lumber piles around a lake. He also collected it at the bases of blackberry bushes, in bark and humus in a young maple stand, in cleared forest areas and at the bases of grass roots in a field. Muchmore (1959: 86), also in New York, states that this species seeks shelter under rocks.

In Ohio, Archer (1937: 121) found Cochliocopa lubrica more common in open country rather than wooded areas. La Rocque (1970: 769) states that "in the Columbus area (Ohio), I have often found it abundant in disused quarries, bedding planes widened by erosion. It is also an abundant snail In stream drift since It inhabits river and creek floodplains."

Actually, the most interesting thing about Cochllcopa lubrica is its aggregation behavior in open areas such as sidewalks and concrete pavements. Roscoe (1962: 111-115) has summarized the data available on this aggregation behavior and investigates several explanations.

Cochllcopa lubrica is frequently found In unnatural habitats made by man. Pilsbry (1948: 1049) states that it is found under board­ walks and stone walls. Boycott (1934: 22) actually called this an "anthropophile" because of its frequent association with gardens and human habitation. Goodrich and van der Schalie (1944: 280-281) stated that this snail is occasionally found in damp cellars. MAP X X I

Collection locations for Cochllcopa lubrica (O. F. Mllller) at Cedar Bog.

w

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live o C ollected D ead

April 23 & 25, 1969 * Collected Alive A C o lle c te d D ead

August 20 & 30, 1969 ■ Collected Alive ° C ollected D ead

Miscellaneous Collections 0 C ollected Alive 0 C ollected D ead

Waaitbii rr>_ Eccd.

i d 10

Q 230 440 12 u m .-a a o it 11

■— r ~ ; h k i x 121

Occurrence at Cedar Bog

Cochllcopa lubrica was never found living at Cedar Bog. It was found in four different samples all in different vegetation zones (Map XXI). A total of eleven specimens were collected, all in a muck substrate.

Vallonia excentrlca (Sterki, 1893)

Distribution

Europe; Madeira; South Africa; Newfoundland southwest to Michigan and Illinois in the west and south to Virginia in the east. It Is also recorded from Oregon, Washington, California and Mexico (La Rocque, 1970: 759-760).

General Remarks

Vallonia excentrlca and Vallonia pulchella are sometimes con­ fused. Oughton (1948: 70-72) found that In the specimens he examined from Ontario the difference between the two species was very small. Oughton considered them the same species. Pilsbry (1948: 1026) and Hubricht (1969: 107-108) considered _V. excentrlca and V. pulchella distinct species. Both authors could readily separate the two species even though they frequently associated with each other.

General Habitat

Boycott (1934: 14 and 17) called V. excentrlca a facultative xerophile. He found them in some localities in England where there was neither shelter nor any accumulated water. Boycott also found them in damp meadows and marshy places, but not in woods.

Sterkl (1907: 378), in Ohio, found them frequently In lawns. He states, "This seems to be one of the few species which” have rather increased in numbers in consequence of deforestation and cultivation of the land."

Occurrence at Cedar Bog

A single fresh dead specimen of Vallonia excentrica was collected in the elm-maple swamp forest. This area of the Bog is open today and has been at least since 1938. This species is probably more common outside of the Bog in the grasses along the edges of cultivated fields and the roadsides. A survey of the mollusca which are present around the Bog would yield valuable ecological data concerning the require­ ments of both the Bog fauna and the non-Bog fauna. I believe 122

M AP X X II

Collection locations for Vallonia excentrica Sterki at Cedar Bog.

u | v w T T

D e c . 23 & 2 5 , 1968 Jan. 5, 1969 • Collected Alive ° C ollected D ead

A p r i l 23 & 2 5 , 1969 * C ollected Alive & C o lle c te d D ead

August 20 & 30, 1969 ■ C ollected Alive 0 C ollected D ead

Miscellaneous C ollections Q Collected Alive 0 C ollected D ead

l«rf. -320 M 123

Cochllcopa lubrica belongs in the same category as Vallonia excentrlca and is more common outside the Bog.

Pupoides albilabrls (C. B. Adams, 1841)

Distribution

Main and southern Ontario south to Florida, west to the Dakotas and south deep into Mexico. It has also been recorded from Bermuda, the Bahamas, the Greater Antilles, Curacao, Venzuela and Columbia (Bequeart and Miller, 1973: 78). Most Ohio records are confined to the southern and western counties (La Rocque, 1970: 732).

General Habitat

"Pupoides albilabrls owes its wide range to its eurytopic toler­ ance to a wide variety of environments. In the Nearctic Region it lives from arid near desert in west Arizon and Sonora to moist mesic forest in east Texas, and from the severe winters of the northeast United States to the subtropical heat of south Texas" (Bequaert and Miller, 1973: 79). Leonard (1950: 29) also mentions this species1 tolerance to a wide range of environmental conditions. In Kansas, he found It thrived In woodlands, prairies, and even short-grass pastures. Basch et al. (1961: 193-194) found It present In 15 of 16 terrestrial stations studied in Kansas with the greatest densities being found in very dry prairie conditions. Cheatum and Fullington (1973: 28), in Texas, records It from under rocks, In grass roots -and leaf litter of sparse woodlands.

In the eastern part of Its range, Pupoides albilabrls is associ­ ated with more mesic environments. Goodrich and van der Schalie (1944: 277) found its habitat in Indiana to be "under sticks, logs and fallen leaves near bodies of water, and on damp limestone cliffs." In Illinois, F. C. Baker (1939: 108) also found it common in wooded bluffs and hillsides along rivers and streams, and occasionally in grassy areas. In lower Michigan, Archer (1939: 23) associated ]?. albilabrls with grassy areas on slopes, roadsides and in wild grape thickets. His investigations in Ohio (1937: 121) found It abundant in fields, rocky open country and railroad embankments. Archer specifi­ cally states that It was not found in woods, either In Michigan or Ohio.

Grimm (1959: 29), F. C. Baker (1939: 108) and Archer (1937: 121) found albilabrls along railroad embankments. Pilsbry (1948: 923), Goodrich and van der Schalie (1944: 277), Leonard (1950: 29), Basch et al. (1961: 193-194) and Cheatum and Fullington (1973: 28) associ­ ated Pupoides albilabrls with limestone bedrock. 1 2 4

MAP X X III

Collection locations for Pupoides albilabris (C. B. Adams) at Cedar Bog.

TTT H | W I I

D ec. 23 & 25, 1968 Jan. 5, 1969 • Collected Alive ° C ollected D ead

April 23 St 25, 1969 * C ollected Alive 6 C ollected D ead

August 20 fit 30, 1969 ■ C ollected Alive ° C ollected D ead

Miscellaneous C ollections © Collected Alive © C ollected D ead

We Qi£iiitLaL_locd \

■ 230*1 125

Basically, P_. albilabrls is characteristic of open, dry, hot, limestone regions. It is more typical of grasslands than wooded areas and not associated with :coniferous forests.

Occurrence at Cedar Bog

I found two dead specimens of Pupoides albilabrls at Cedar Bog. One was found in the oak-maple zone and the other in an isolated arbor vitae grove surrounded by bog meadow (Map XXIII). I was very sur­ prised to find either of these specimens in the study area. However, the specimen found in the oak-maple forest was very near the old field west of the study area. This field is likely the source and is a more typical habitat for albilabrls. The presence of this species in the Isolated arbor vitae grove is a mystery to me.

Gastrocopta procera (Gould, 1840)

Distribution Southern Ontario and Maryland south to Alabama, west to South Dakota, southeastern Wyoming, Colorado and Arizona (Leonard, 1950: 32). Goodrich and van der Schalie (1944: 276) stated that the northern range of (J. procera was limited by the north-south drainage divide In Ohio and Indiana. Most Ohio records are from the southwestern counties. There are a few records from northern Ohio (Reigle, 1963: 16-17).

General Habitat

Very little habitat information is available for Gastrocopta procera is typically an inhabitant of timbered slopes near streams, where it lives in leaf mold, beneath fallen logs or loosened bark, or beneath stones, but it sometimes is found In meadows in dead grass. Its wide distribution indicates an ability to tolerate wide extremes of temperature and humidity." Basch et al. (1961: 193), also in Kansas, found it in 13 of 16 terrestrial stations Investigated and frequently with Pupoides albilabrls in dry open prairie conditions. They found that "G. procera seemed to favor fairly dry environments since it was not abundant in the more mesic areas of hedgerows and woods. Shells were generally collected under limestone rocks and cow dung although many were burled among the roots of prairie grasses."

In Texas, Cheatum and Fullington (1973: 20) associated this species with "open as well as woodland areas and it is universally more abundant in-the latter. It Is frequently found under limestone rock on sloping hillsides with sparse trees and shrubs." 126

MAP X X IV

Collection locations for Gastrocopta procera (Gould) at Cedar Bog.

\ t \ ~ w ) * | r I

Dec. 23 & 25, 1968 Jan. 5, 1969 \ • Collected Alive ° Collected Dead

April 23 & 25, 1969 A C ollected Alive d C ollected D ead

August 20 Sc 30, 1969 * Collected Alive 13 Collected Dead

Miscellaneous C ollections ©Collected Alive © C ollected D ead

Wnmlhifn t.rrf

10 10

o aao 440

13 1

Occurrence at Cedar Jog,

I found one dead and one living specimen of Gastrocopta procera at Cedar Bog. The living specimen was taken In an arbor vitae stand near Cedar Run. The dead specimen was found in the oak-maple forest (Map XXIV). This specimen was found in the same sample in which Pupoides albilabrls was found. Considering procera is such a rare species at Cedar Bog, little can be added to its ecology from this study. Finding this species living within the arbor vitae association was most unexpected.

Gastrocopta armifera (Say, 1821)

Distribution

Southern Canada and the United States east of the Continental Divide. Not found in southern New Mexico, Texas or Florida (Pilsbry, 1948: 875). In Ohio, it is found in all counties except the north­ east and east ones (La Rocque, 1970: 718-719).

General Habitat

Gastrocopta armifera appears to have fairly distinct habitat preferences. Few authors (Leonard, 1959: 170 and F. C. Baker, 1939: 95-96) associated this species with a wide variety of habitats. F. C. Baker (1939: 95-96) did say that G. armifera was characteristic of the prairie region of Illinois. Smith (1928: 487), in Illinois, and Archer (1937: 121 and 1939: 22-23), in Michigan, found this species in exposed areas such as fields. Archer goes even further and specifi­ cally says that (J. armifera was rarely found in any wooded areas. In the western part of its range, such as in Kansas, Basch, Balner and Wilhn (1961: 193), Leonard (1950: 29-30) and Krutzsch (1952: 130-132) collected it in various wooded areas. I did not find any references which links this species to a coniferous forest of any kind.

Smith (1928: 487), Archer (1937: 121 and 1939: 22-23) and Grimm (1959: 125) found G. armifera in rocky areas. F. C. Baker (1939: 95-96), Pilsbry (1948: 875), Oughton (1948:.50), Leonard (1950: 29-30), Burch (1955: 66), La Rocque (1970: 718) and Cheatum and Fullington (1973: 11) associated this species with not only rocky areas but limestone bedrock. Burch collected it in Virginia only in- areas where the pH was 7.3-7.7 with a very high calcium, magnesium, potas­ sium and phosphorous concentration.

Gastrocopta armifera is a species characteristic of dry, rocky, exposed, high calcium habitats. Because of these preferences It is frequently collected in quarries, railroad embankments, old house foundations and some other urbanized environments (F. C. Baker, 1939: 128

MAP X X V

Collection locations for Gastrocopta armifera (Say) at Cedar Bog.

I ~ ~ " T w

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

A p r i l 23 & 2 5 , 1969 * C ollected Alive A C o lle c te d D ead

A u g u s t 20 & 3 0 , 1969 ■ C ollected Alive Q Collected Dead

Miscellaneous Collections Q Collected Alive © C ollected D ead

WouhuuUbicd

10

O 220 **0

lcm. *.010*1 11 129

95-96, Grimm, 1959: 125, Gugler, 1963: 196-197, and La Rocque, 1970: 718).

Occurrence at Cedar Bog

I found one dead specimen of Gastrocopta armifera in the oak- maple forest (Map XXV). This specimen was quite old. After reviewing the ecology of this species, Vallonia excentrlca, Pupoides albilabrls and Gastrocopta procera, I find all these species are more character­ istic of the prairies and the dry grasslands of the middle states. All these species are much too common in Ohio to be called prairie relics, but their rarity in the Bog today seems to lead one to that conclusion. Actually, all these species are probably far more common in the surrounding fields, hedgerows and road berms. If they are not, then maybe some fossil digging at Cedar Bog would yield some interest­ ing history concerning past molluscan changes. As far as I am aware, no one has examined the fossil record at Cedar Bog.

These snails are very small, much smaller than many seeds. They are probably transported in the same manner as some seeds. These snails could easily be carried Into the Bog on a boot, tools or live­ stock in a clod of dirt. Wind could also carry small species like these Into the Bog.

Gastrocopta contracta (Say, 1822)

Distribution

Quebec to Manitoba, south to northern Mexico in the west and south to Florida In the east. In Ohio, it is more commonly collected In the western counties (La Rocque, 1970: 721).

General Habitat

Gastrocopta contracta has a very wide ecological amplitude. It Is not confined to any specific vegetation zone or habitat. In the west, it is frequently associated with Gastrocopta armifera (Leonard, 1959: 174, and Cheatum and Fullington, 1973: 13). (3. contracta can be found in dry exposed, rocky areas, but not in the density that C5« armifera is found.

£. contracta is found in numerous habitats that G^. armifera is usually not associated with. Oughton (1948: 95) found it along the "margins of ponds, streams and marshes; seeping hillsides; also sandy flats which receive water by percolations." F. C. Baker (1939: 97) collected It on river and creek floodplains where the moisture was high. H. B. Baker (1911: 159) found (G. contracta abundant In Michigan 130

In both wet and dry habitats, even In a damp meadow along a northern white cedar thicket. Archer (1939: 22), also In Michigan, found G . contacts in many habitats, some which included deep marsh grasses and blrch-maple swamp woods.

Gastrocopta contracta is rarely associated with coniferous forest, but it is common in numerous types of deciduous forests, open fields and in both wet and dry habitats. It is not limited to rocky areas or limestone areas.

Burch (1955: 66) found it to be scarce, but in areas where the pH was 6.3 to 7.2 and the soil was high to very high in calcium, magnesium and potassium, low to high in phosphorous and very high in organic matter. Burch also found it more or less restricted to com­ munities dominated by oak.

Occurrence at Cedar Bog

The distribution map of Gastrocopta contracta (Map XXVI) shows a wide, almost even distribution, over the study area with very little clumping. Dead shells of this species have been found in very vege­ tation zone in the Bog with a fairly high frequency (Plate II, Figure 4). Living specimens have not been found In the bog meadow or the shrub zones, in spite of a reasonably high frequency of occurrence. It is interesting to note that two highest frequencies for dead shells are two of the lower frequencies for living shells. These two vegeta­ tion zones, the shrub and the elm-maple zones show a lot of disturb­ ances. Most of the living specimens are found.In the central parts of the study area where the disturbances have been low compared to the northern and southern areas. The major exception to this is the high frequency of living G. contracta in the silver-red maple zone which was an open field in 1938. It is now relatively undisturbed.

A look at the various habitat diversity indices for (3. contracta support its distribution patterns (Table 10). This species has a high index reading of 2.83 for the total collected and 2.24 for the living collected in the Frederick vegetation zonation scheme. It has a habitat diversity index of 3.02 for total collections in the En­ vironment Control Corporation vegetation zonation scheme. This last index reading Is very high, the highest index for any species in this particular zonation scheme. Gastrocopta contracta does not appear to show any habitat preferences at Cedar Bog. Living specimens are more frequently collected in the less disturbed areas of the Bog. MAP X X V I

Collection locations for Gastrocopta contracta (Say) at Cedar. Bog.

April 23 & 25, 1969 * C ollected Alive a C ollected D ead

A u g u s t 2 0 & 3 0 , 1969 ■ Collected Alive D C ollected D ead

Miscellaneous C ollections Q C ollected Alive © C ollected D ead

Wonrfhiifn tried 132

Gastrocopta pentodon (Say, 1821)

Distribution

Southern Canada from Prince Edward Island west to British Columbia, south to Mexico in the west, and south to Florida in the east. It is not found on the Pacific Slope. In Ohio, it has been recorded from eight widely distributed counties (La Rocque, 1970: 723-724).

General Habitat

Gastrocopta pentodon is an animal of mesic deciduous forests and fields. Oughton (1948: 52 and 94) found it in southern Ontario on damp deciduous hillsides or in well drained groves among leaves in the underbrush. Burch (1955: 160), in Virginia, and Blake (1931), in Illinois, found it in leaf litter of deciduous woods. F. C. Baker (1939: 100), Leonard (1959: 176) and Cheatum and Fullington (1973: 18) associated G^. pentodon with good cover on deciduous hillsides and grassland areas which were not too dry. Call (1900: 396-397) states, "This species in particular may be found all over Indiana, under bark, logs, fallen trees, and often under flat stones, on the edges of forests and in open lands in river bottoms. It is also to be found in gardens, about dwellings, under old boards, stones, andunder refuse piles of plant stems."

H. B. Baker (1911: 159) and Archer (1939: 22), in lower Michigan, found this snail in some wet areas, such as a swamp forest, In addi­ tion to the mesic fields and wooded hillsides.

I have found no records which places this snail In dry, rocky areas, in coniferous forests or on floodplains.

Occurrence at Cedar Bog

The distribution pattern of Gastrocopta pentodon Is to that of Carychium exile (Map XIX and Map XVII). (J. pentodon has a wide dis­ tribution over the Bog, but not extensively in the wettest areas, such as the ponded areas of the swamp forests and In the marl meadow. This snail has been collected alive in every vegetation zone in the study area. In the elm-maple swamp forest, specimens were found in 89% of the samples, in the shrub zones, 80% and in the arbor vitae associa­ tion 58% (Plate III, Figure 3). The habitat diversity indices range from 2.02 to 2.62 which are in the high range. The difference between this range is actually the difference between the total number col­ lected (2,02) and the living number collected (2.62). Usually readings are reversed with the greatest habitat diversity being from the total collected. In this case, the difference indicates a very uneven dis­ tribution of dead specimens. For example, in the elm-maple swamp forest where the total number collected was 2048, the frequency was 89% 1 3 3

MAP XXVU t. Collection locations for Gastrocopta pentodon (Say) at Cedar. Dog.

i—5 I ; r

Dec. 23 & 25, 1968 Jan. 5, 1969 • Collected Alive 0 C ollected D ead

April 23 & 25, 1969 * C ollected Alive 6 C ollected D ead

August 20 & 30, 1969 C ollected Alive D C ollected D ead

Miscellaneous Collections 0 Collected Alive 0 Collected Dead

1cm. xlSOft

-f Frequency (SS) Frequency (# ) 100 100 30 80 20 70 90 10 40 80 20 0

- - - • - - - Figure Figure Figure Figure B A 1 3 . Collection frequencies for for frequencies Collection . * Collection frequencies for for frequencies * Collection G Gastrocopta pentodon. Gastrocopta Vertigo morsei. Vertigo VrrS Vegetation Zones Vegetation 0 Vegetation Zones Vegetation r-4;: ovs . , r.V E -.’■'■'■j vi/>: Kill » c * v#-V VS V ,fV T'»*

* H G ». J • ;* ( s**.* V'V*" i<' * \ V* J *f \ PLATE III PLATE K li- c. o 5* cr C - ICO o- so 20 30 40 50 6o 0 7 - 90 10 80 „ ------

Figure Figure

Figure Figure

[wL fax*. ABCDEFGHJK 4 2 . Collection frequencies for for frequencies Collection . . Collection frequencies for for frequencies Collection . Vertigo ovate. Vertigo Gastrocopta tappaniana. Gastrocopta C C ■ 1 i Vegetation Zones Vegetation 0 F H J H G F E rrrr V I • —I I— .— I— 4 3 1 K i . i 135 and in the shrub zones where the total number collected was only 389, the frequency was still very high at 80%.

The total average density for Gastrocopta pentodon in the elm- maple swamp forest was 138.1/ ft2, in the shrub zones it was 32.4/.ft2 , in the bog meadow 31.5/ ft2, in the silver-red maple swamp forest it was 18.4/ ft2 and in the arbor vitae 12.3/ ft2 . In these same vegeta­ tion zones the density of the living snails varied only from .8/ ft2 in the arbor vitae to 1.6/ ft2 in the bog meadow and silver-red maple zones.

In spite of the wide distribution pattern for this species, it is usually found in the greatest numbers dead and alive in the mesic areas. It is not common in the wettest or the driest areas. This supports the previous investigators. Gastrocopta pentodon does live in the arbor vitae association which is a coniferous forest.

Gastrocopta tappaniana (C. B. Adams, 1842)

Distribution

Maine and southern Ontario south to Florida, west to South Dakota in the north and Texas in the south (La Rocque, 1970: 726). He also lists records for seven counties in Ohio, Most of these are in the western half of the state.

General Remarks

It is unfortunate that Bequaert and Miller (1973: 88-89) have lumped Gastrocopta pentodon and Gastrocopta tappaniana together under G. pentodon. Although the range and habitats of these two species overlaps, they can be consistently separated, at least at Cedar Bog. It is true, as in the case of Carychium exile and Carychium exiguum, that some shell characters intergrade, but I believe all four of these species should remain separated. The preferred habitat for each of these species appears to be distinct and keeping them separate . would be far more useful to the ecologist than lumping them, since they are good habitat indicators. These species could be used as indicators of varying moisture conditions and are definitely useful when comparing various plant communities.

General Habitat

H. B. Baker (1911: 159 and 1922: 37) in Michigan, F. C. Baker (1939: 101) in Illinois, Goodrich and van der Schalie (1944: 276) in Indiana, Oughton (1948: 95) in Ontario, and Leonard (1950: 32) in Kansas associate Gastrocopta tappaniana with moist, wet or swampy habitats. It is usually found in or on the margins of streams, rivers, 136

ponds, lakes, marshes and swamps; In open areas, coniferous or decidu­ ous forests. Basch et al..^l961: 193), in Kansas, has also found it around springs. H. B. Baker (1911: 159) and Harry (1951: 32) collected G. tappaniana in and around arbor vitae bogs. La Rocque (1970: 725) states that (J. tappaniana is associated with Vertigo ovata.

Occurrence at Cedar Bog

Gastrocopta tappaniana is found most often in the marl meadow, the bog meadow and in ponded areas and springs in the swamp forest segre- gants (Map XXVIII). Except for the four living specimens collected along the stream in the silver-red maple swamp forest, all the living and most of the fresh dead specimens were found in the bog and marl meadows.

The habitat diversity indices for G. tappaniana vary from high to low (Table 10). This is due mainly to the great difference in the wet vegetation zone interpretation between Dr. Frederick and the E.C.C. investigators. In this case, I believe that the Frederick zonation scheme is more accurate and the boundaries between the bog meadow, the marl meadow and the arbor vitae are distinct enough. I do know where I collected each sample. The habitat diversity index for living specimens in the Frederick interpretation is 0.73. This is low as it should be for a species with such a specialized niche.

The graph (Plate III, Figure 2) showing differences in the fre­ quency of occurrence in the various vegetation zones shows the very high frequency of Gastrocopta tappaniana in the marl meadow both dead and alive. Density figures show this species' preference for a wet habitat. The average density for total number collected is 158.2/ ft^ in the bog meadow and 146.1/ ft^ in the marl meadow. No other zone comes near those figures. The density for living specimens is very low, 0.1/ ft2 in the bog meadow and 1.8/ ft^ in the marl meadow.

Gastrocopta corticaria (Say, 1816)

Distribution

Southeastern Canada south to Florida, west to Minnesota in the north and Texas in the south (La Rocque, 1970: 728). LaRocque also records £. corticaria from nine widely scattered Ohio counties.

General Habitat

Gastrocopta corticaria is a climber. F. C. Baker (1939: 101), Pilsbry (1948: 894), Oughton (1948: 51) and Leonard (1959: 179) state that this snail is frequently found up off the ground on large logs and the sides of trees, as much as two feet. This habit is unusual 137

MAP xxvrn

Collection locations for Gastrocopta tappaniana (C.B. Adams) at Cedar Bog.

D e c . 23 & 25, 1968 Jan. 5, 1969 Collected Alive ® C ollected D ead

April 23 25, 1969 C ollected Alive A C o lle c te d D ead

August 20 & 30, 1969 C ollected Alive o C ollected D ead

Miscellaneous Collections Q C ollected Alive © C ollected D ead

t«*n. m i lOh 138 for such a small species. G. corticaria is also most often observed associated with trunks of trees, loose bark or decayed wood (Call, 1900: 399; H. B. Baker, 1911: 159; F. C. Baker, 1939: 101 and Leonard, 1959: 128).

In Michigan, H. B. Baker (1911: 159) found this species seemed to prefer "a wet, rich habitat,M and Archer (1939: 22) found it "com­ mon in the fecal mush of arthropods under started bark on rotten logs in the oak-hickory woods." F. C. Baker (1939: 101) found this species on hillsides, river bluffs and deciduous woods in Illinois.

Most researchers who did habitat studies that Included Gastrocopta corticaria found this species rare or localized in small specific niches.

Occurrence at Cedar Bog

Gastrocopta corticaria has- remained consistent with the notes of previous workers on terrestrial mollusca. I found one colony of this species under loose bark of a large fallen elm tree in the maple-ash- basswood swamp forest. I collected seven living and two dead speci­ mens. This species never turned up in any of the soil and litter samples.

The specialized habitat and rarity of this species presents some interesting questions. How does this species get from one dead tree to another? Why did not at least one dead shell appear'in the soil and litter samples? Do the adults ever leave their' normal habitat? If not, how do the immatures or the eggs get carried around? The unfortunate thing about this species is that its apparent rarity makes it difficult to study.

Vertigo morsel Sterki, 1894

Distribution

The range of Vertigo morsel is described by La Rocque (1970: 736-737) as "New York and New Jersey west to Grand Rapids, Michigan, Indiana and Illinois. In Ontario it Is not known north of Hastings County." In actuality, this snail is known from about 15 collections made since Its description in 1894 by Victor Sterki from Michigan. V. morsel is now known in the Pennsylvania fauna, it is only recorded from one recent collection in New Jersey (Alexander, 1952: 58), Illinois (Baker, 1939: 109), Ohio (Sterki, 1907: 379) and Ontario (Oughton, 1947: 61). There are two records each from New York (Robertson and Blakeslee, 1948: 36) and West Virginia (Brooks, 1937: 98-98 and Briscoe, 1963: 41). The remainder of the records are from Michigan (Walker, 1906: 516) and Indiana (Blatchley and Daniels, 1903: 139

MAP X X IX

Collection location for Gastrocopta corticaria (Say) at Cedar Bog.

t I u 1 v

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live C $ \ o C ollected D ead

A p r i l 23 & 2 5 , 1969 * C ollected Alive 0 C ollected D ead

August 20 & 30, 1969 ■ Collected Alive o C ollected D ead

Miscellaneous Collections 9 C ollected Alive © C ollected D ead 140

587). Since I believe this species to be somewhat of particular interest, I have included a United States distribution map (Map LXIX).

The type locality for Vertigo morsei Sterki, 1894, is Kent County, Michigan (Winslow, 1926: 7 and Sterki, 1894: 89-90). Pilsbry (1948: 952), and La Rocque (1953: 335 and 1970: 735-738) recorded the type locality for V. morsei as Joliet, Will County, Illinois. This is incorrect.

Vertigo morsel is recorded from many Pleistocene deposits. I examined the records of 139 Pleistocene collections from eight states and Canada (Walker, 1906; La Rocque, 1966; Nave, 1969 and Bickel, 1970) and only found this species recorded in 15 collections. Eleven of these collections were made from five Ohio counties (Green, Cham­ paign, Stark, Erie and Ottawa),

General Habitat

The habitat of V. morsei is poorly known. Blatchley and Daniels (1903: 587) found V. morsei around three different lakes of glacial origin. They found it living under plant rubbish, such as fallen weeds, grass stems and pieces of wood. Eight of the 15 recent collec­ tions were definitely from around ponds or lakes. Three of these recent records and twelve of the Pleistocene records were associated with marl deposits. Some of the other records could have been associ­ ated with marl deposits, but the collectors were not always specific about where the collections were made.

Until a lot more collecting is done, the range and habitat for V. morsei will not be well known. Preliminary indications lead me to believe that \7. morsei could possibly be a Pleistocene relic cur­ rently living in wet habitats around lakes and ponds of glacial ori-4 . gin. It may even be restricted to areas near marl deposits.

Occurrence at Cedar Bog

One look at the distribution map (Map XXX) will show its associ­ ation with the marl meadow area. All living specimens were found on marl in the marl meadow. Vertigo morsei is of common occurrence only in the marl meadow (Plate III, Figure 3). Actually, dead specimens have been collected in 7 different vegetation zones, but 152 of the 174 collected were taken from the marl meadow. The habitat of this species at Cedar Bog supports what little information is available. V. morsei is a species of wet areas and commonly associated with marl deposits. The habitat diversity indices are medium to low (Table 10) which reinforces its narrow habitat niche requirements. 141

MAP X X X

Collection locations for Vertigo morsei Sterki at Cedar Bog. i— r r f ii j " ' v~ w | % | r [ t

D ec. 23 8c 25, 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

A p r i l 23 8c 25, 1969 A Collected Alive & C ollected D ead

A u g u s t 20 8c 3 0 , 1969 ■ C ollected Alive D C ollected D ead

Miscellaneous Collections Q C ollected Alive 6 Collected D ead

Woadhurrt Incd

1cm. •030ft 142

The presence of Vertigo morsel as a living member of the fauna may be the first real proof that this snail is living in Ohio today. It is now known with certainty whether Sterki1s specimens from Erie and Ottawa counties were living. He collected them in an area were there are numerous marl deposits. He may have had Pleistocene fossils. His specimens should be reexamined and the collection site revisited.

If Vertigo morsei is a Pleistocene relic living in a few scat­ tered areas which still have wet marl, it would take much more collecting and careful searching to establish this theory more con­ clusively.

Vertigo ovata Say, 1822

Distribution

Vertigo ovata is recorded from most of North America. It has not been found in Nevada and is sparsely collected in many of the western states (Bequaert and Hiller, 1973: 92-93). It has been found In a few localities in Mexico and the greater Antilles. Pilsbry (1948: 953) states that V. ovata "is common in the eastern and middle western states, rarer and local in the southern and western peripheral states of its range." Oughton (1948: 62) indicates that Vertigo ovata has the greatest range of latitude and climate of any Vertigo or other pupillid in the world.

General Habitat

F. C. Baker (1902: 244 and 1939: 105), In Illinois, and Cull (1900: 401) in Indiana, found V. ovata plentiful in moist places under sticks, stones and miscellaneous debris usually along rivers, creeks and ponds. H. B. Baker (1911: 159-160), in upper Michigan, collected this species in six different localities; among sedges and driftwood on a sandy beach, among leaves and logs along the shores of lakes and swamps; under logs and driftwood in swampy meadows along a river; among leaves and under logs in a swamp forest and under driftwood in a sedge marsh along a lake. Archer (1939: 22), in lower Michigan, found the habitat of V. ovata to be in sedges, grasses, leaves and plant debris along the edges of pools and rotting leaves in a tamarack bog. Harry (1951: 31), also in lower Michigan, found this species to be quite common in northern white cedar swamps.

In Ontario, Oughton (1948: 95) recorded V. ovata from the "margins of ponds, streams and marshes; seeping hillsides; also sandy flats which receive water by percolation." In Georgia, Teskey (1955: 70-71) collected it in a "cattail swale at the edge of a man-made pond." In Kansas, Leonard (1959: 186) collected this snail from moist or marshy areas and found enormous populations near artesian springs. MAP XXXI Collection. locations for Vertigo ovata Say at Cedar Bog.

s I t | u 1 v j w t > 1 r | z

Dec. 23 & 25. 1968 Jan. 5, 1969 • Collected Alive ° C ollected D ead

A p r i l 23 & 2 5 , 1969 * Collected Alive A C o lle c te d D ead

August 20 & 30, 1969 « C o lle c te d A liv e ° C ollected D ead

Miscellaneous Collections O C ollected Alive 0 C ollected D ead

Waqrfhurn Sncd. 144 springs. In Texas, Cheatura and Fullington (1973: 32) associated V. ovata with considerable moisture near springs, creeks and marshy areas.

There seems to be little doubt that this species is partial to wet or moist habitats. However, it appears that most species with large distributions also have fairly wide ecological requirements.

Occurrence at Cedar Bog

The distribution pattern for Vertigo ovata is somewhat unclear at Cedar Bog (Map XXXI). I found it living in only two areas, the maple-ash-basswood swamp forest and the bog meadow. The frequency of occurrence for V_. ovata was around 20% in the bog meadow and marl meadow. Its density was generally quite low, only 85 speciems were found among the 18 samples in which it appeared. Forty-six specimens were taken in the bog meadow.

The habitat diversity indices were medium to high for total number collected and low for the living number collected. Vertigo ovata is assocaited with the wetter areas of the Bog along the streams and springs, and is totally absent from the driest areas.

Vertigo pygmaea (Drapernaud, 1801)

Distribution

This species is Holarctic, being found in Europe, Siberia, northern Caucasia, Transcaucasia, central Asia, and eastern North America (Likhachev and Rammel'meier(1952: 137). In Canada it has been found in Nova Scotia, Quebec and Ontario (La Rocque, 1970: 743). In the United States, La Rocque records it from Maine to Virginia west to Ohio. In Ohio it has been recorded from Franklin and Summit counties.

General Habitat

In Asia, Likhachev and Rammel’meier (1952: 137) stated that Vertigo pygmaea lived "both on plains and mountains; it is encountered in humid meadows in moss and grass, in forests under rotting sylvan vegetation and under leaf litter." In England, Boycott (1934: 14) found V. pygmaea usually associated with damp meadows and marshy places, though they live equally well or better in other habitats. He found it to be indifferent to lime.

"This species, at least in North America, seems to be partial to lime-rich Paleozoic soils; it is a hardy species, capable of surviving 145

In the rigorous climate of the Hudson Bay area, but it has spread as far south as Virginia so that it is probable that lime content of sub­ strate is a more important factor for this species than cool climate" (La Rocque, 1970: 743-744).

In Ohio, Archer (1937: 121) found it with Vallonia pulchella and Vallonia costata in open country. Grimm (1959: 126) found it in Maryland near a railroad track.

Occurrence at Cedar Bog

At Cedar Bog Vertigo pygmaea is definitely associated with the wettest areas (Map XXXII) and closely associated with marl (high lime). These findings are right in character for this species. Only one dead specimen was collected some distance from the bog or marl meadows (Map XXXII, W-8). The frequency graph (Plate IV, Figure 1) shows a very strong association with the marl meadow. The habitat diversity indices vary from medium to low and are fairly consistent.

Vertigo tridentata Wolf, 1870

Distribution

Southeastern Canada west to Minnesota, south into northern Texas in the west and south to Virginia in the east (La Rocque, 1970: 745). It is absent from the Coastal Plain area. In Ohio, Vertigo tridentata has been recorded from Summit, Tuscarawas, Franklin, Miami and Hamilton counties (Sterki, 1907: 380).

■■General Habitat W When Wolf (1870: 198) discovered this species In Illinois he found it "abundant In shady copses on green weeds, climbing as high as three feet from the ground." At the time, he collected 12,000 from standing weeds and not one from the ground. Since that time very little has been added to our knowledge of its habitat. Leonard (1959: 188) stated that this species was found "in and among ground debris in forested areas as well as grass and weeds in more open situations," In Maryland, Grimm (1959: 126) found it in a quarry and around the ruins of a house. In Ontario, Oughton (1948: 95) said it was probably on creek and river flood plains.

Occurrence at Cedar Bog

Vertigo tridentata has been found dead In every vegetation zone in the Bog, but that hardly reflects its true distribution or habitat MAP X X X II

Collection locations for Vertigo pygmaea (Draparnaud) at Cedar Bog.

w

D e c . 23 & 2 5 , 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

April 23 & 25, 1969 A C ollected Alive a C ollected D ead

August 20 & 30, 1969 ■ C ollected Alive o C ollected D ead

Miscellaneous Collections Q C ollected Alive @ C ollected D ead

W e aztibuLQ—Bn c

I cm.-320ft Frequency (jfi) . Frequency (JS) 100 100 40 -40 - 0 7 •80 50- 30. 0 6 20 - 0 5 40 ■ 40 70- ■ 80 50- 0 6 90 30 - 30 20 10 0

- ■ -I * - . .

Figure 3* Collection frequencies for for frequencies Collection 3* Figure Figure 1. Collection frequencies for for frequencies Collection 1. Figure *!.’*» w U fi-Tf. r.V* ABCOtFGHJ • Y.&21 •tai‘ Vertigo pygmaea. Vertigo Vertigo gouldi. Vertigo Vegetation Zones Vegetation Vegetation Zones Vegetation LT IV PLATE 5 w « 1 d> c S*

- - - . - Figure 2, Collection frequencies for for frequencies Collection 2, Figure Figure 4. Figure h f'Ti- $ 8 ; * r** .s4§» B Vertigo milium. Vertigo Vertigo tridentata. Vertigo Collection frequoncios frequoncios Collection C C Vegetation Zones Vegetation Vegetation Zones Vegetation 0 E F G H H G F E m m 147 148 preferences. When comparing this species' distribution (Map XXXIII) with the soil patterns determined by Ritchie et al. (Map II) only two samples occur in the marl zone, one in the peat and none in the loam. This species is most frequently collected in the muck soils of the silver-red maple, shrub and elm-maple vegetation zones (Plate IV, Figure 2). There are no springs or ponded areas in these three zones. The muck soil in these areas is loose, granular and moist most of the time. These vegetation zones are also in a high state of change and not what I would call stable communities.

The habitat diversity indices are medium to low (Table 10). This shows the uneven distribution easily seen on the distribution map and the habitat specificity of this species.

All the specimens I collected were part of the soil and litter. It is possible that I missed this species if it climbed the sur­ rounding vegetation. It is a small brown species and easy to overlook.

Vertigo elatior Sterki, 1894

Distribution

Southern Canada from the east to the west coasts, southeast through Montana to New Mexico in the west, and south to West Virginia in- the east. Vertigo elatior is generally absent from the Rocky Mountains (La Rocque, 1970: 740-741). In Ohio, Taft (1961: 44) recorded from Summit, Stark, Tuscarawas and Erie counties.

General Remarks

Oughton (1948: 64-65) and Bequaert and Miller (1973: 96) believe this species to be synonymous with Vertigo ventricosa or refer to it as a subspecies of Vertigo ventricosa. I have not thoroughly investi­ gated the validity of either position. I am referring to it as a distinct species as did Pilsbry (1948: 956).

General Habitat

Little is known about the ecology of Vertigo elatior. La Rocque (1970: 741) states that i. elatior is "a hardy species typical of rigorous climate zones but able to survive In mountainous areas in the south." In upper Michigan, H. B. Baker (1922: 29 and 36) found it on the floodplain of a creek which was dominated by grasses and sedges. In Ontario, Oughton (1948: 95) lumps this species with V. ventricosa and indicates its affinity for margins of ponds, streams and marshes and other wet areas. F. C. Baker (1939: 106), in Illinois, found It rare, but associated with Vertigo ovata which is a snail of wet habitats. MAP XX XIII

Collection locations for Vertigo tridentata Wolf r.t Cedar Bog.

D ec. 23 Sc 25, Jan. 5, 1969 • C ollected A live ° C ollected D ead

April 23 Sc 25, 1969 A C o lle c te d A liv e A C o lle c te d D ead

August 20 & 30, 1969 Collected Alive D C ollected D ead

Miscellaneous C ollections © C ollected Alive © C ollected D ead

Woqrfhmp .gncd ISO

Occurrence at Cedar Bog .

I found one dead specimen of Vertigo elatior In the elm-maple swamp forest segregant just north of the bog meadow. I cannot draw any conclusions about this species. I would not expect to find it. common in the areas surrounding habitats outside the Bog.

Vertigo gouldi (Blnney, 1843)

Distribution

Newfoundland west to British Columbia southeast to Mexico in the west and south to Maryland in the east. Vertigo gouldi is found in Tennessee, northern Alabama and Georgia (La Rocque, 1970: 749). It is sporadic in the western part of its range. It has been recorded from Ashtabula, Summit, Portage, Tuscarawas and Washington counties in eastern Ohio (Taft, 1961: 45).

General Habitat

"Ecological data on this species are not available, beyond the fact that it requires a moderate amount of moisture, and does not easily survive periods of high temperature and low humidity" (Leonard, 1950: 27). Oughton (1948: 95), in Ontario, associated V. gouldi with wet areas around ponds, streams, marshes, springs and occasion­ ally Sphagnum bogs.

Most of the habitat data available for this species comes from studies made in Michigan. H. B. Baker (1911: 160) found V. gouldi in two mixed deciduous and coniferous swamps and a dry deciduous woods. All three habitats had a substantial amount of northern white cedar or arbor vitae. Later, H. B. Baker (1922: 27-28) found V. gouldi In a virgin mixed deciduous and coniferous forest with a good accumulation of leaf litter and dead wood. In the same survey he also found this species quite common In a cedar-tamarack bog which was entirely carpeted with Sphagnum. Walker and Ruthven (1906; 97J collected V. gouldi among the dead leaves in a deciduous woods and in moss in a tamarack swamp.

Vertigo gouldi has a wide distribution, but It appears to be most common in the northern states and southern Canada in moist.to wet deciduous and mixed deciduous and coniferous forests.

Occurrence at Cedar Bog

Vertigo gouldi shows a strong association with the bog meadow and the marl meadow zones (Map XXXV). All the living specimens collected 151

MAP XX XIV

Collection locations for Vertigo elatior Sterki at Cedar Bog.

I I w T * i T \

D ec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live ° Collected Dead

A p r il 23 & 2 5 , 1969 * C ollected Alive A C o lle c te d D ead

A u g u s t 20 81 3 0 , 1969 ■ C ollected Alive n C ollected D ead

Miscellaneous C ollections © C ollected Alive © C ollected D ead

10 10

u o aao IJ 1cm. clJOft. 13

U ' V 1 w 1 5 2

MAP XXX V

Collection locations for Vertigo gouldi (Binney) at Cedar Bog.

D e c . 23 & 2 5 , 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

April 23 25, 1969 C ollected Alive C ollected D ead

A u g u s t 20 & 3 0 , 1969 Collected Alive ° C ollected D ead

Miscellaneous C ollections 0 C ollected Alive 0 Collected Dead

Woarfham Kncd

120’i 153 were from areas along the East Branch of Cedar Run. These collections show a strong association with w6t places, though not necessarily the exposed marl. V. gouldi was frequently collected in the shrub zones and the elm-maple swatnp forest, but only dead specimens (Plate IV, Figure 3).

The habitat diversity index for the total number collected in the various E.C.C. vegetation zones was 2.59 which is pretty high. This would seem to indicate that V. gouldi has a rather wide habitat range. The habitat diversity indices for the total collected and the living collected in the various Frederick vegetation zones are much lower. This would Indicate a much more restricted habitat preference.

A look at the density data for V. gouldi in the bog and marl meadows show some Interesting contrasts. In the bog meadow, where it was found in 32% of the collections, I found an average of 7.9/ ft,^ for total number collected and an average of only 1.1/ ft.2 for living number collected. In the marl meadow, where It was found In 68% of the collections, the average density for total number col­ lected was 29.1/ ft.2 and for the living number collected it was 2.3/ ft.2. This shows a high degree of clumping in the bog meadow.

The general association of V. gouldi with wet habitats agrees with the findings of previous workers, but its association with deciduous or mixed deciduous and coniferous forests does not. ,1 found only 53 specimens in 5 collections from areas which were heavily forested in 1938. I found 1232 specimens In AO collections In open or lightly forested areas. One living specimen was found in a forested area In Y-7 and two living specimens in an arbor vitae stand next to the bog meadow in W-A.

Vertigo milium (Gould, 18A0)

Distribution

Maine and southern Ontario south to the Florida Keys, west to Minnesota, South Dakota, Colorado and eastern Arizona (Bequaert and Miller, 1973: 9A-95). Also recorded from Jamaica and Hispaniola In the West Indies, these records are probably accidental introductions. La Rocque (1970: 735) records it from seven counties in Ohio.

General Habitat

"Its habitat is under sticks and debris in moist floodplain areas and In woods on bluffs bordering the large rivers of the state" (F. C. Baker, 1939: 107). In Kansas, Leonard (1950: 26) states that "Vertigo milium is an inhabitant of humid situations, such as those afforded by marshes and wooded slopes near streams. It is absent 154 from regions characterized by low humidity or by high extremes of summer temperatures." In Texas, Cheatum and Fullington (1973: 30) found milium living "In moist situations usually along heavily- wooded streams and associated with rock and litter." Oughton (1948: 95), in Ontario, recorded that it may live on floodplains of creeks and rivers confined to limestone regions. La Rocque (1970: 735) states that Vertigo milium is "apparently confined almost entirely to lime-rich areas.

From the small amount of Information available, it is evident that Vertigo milium is a snail of moist habitats usually under the cover of a forest. This species may be restricted to limestone regions in Canada, but I believe more information must be gathered before this species can be classified as a calciphile.

Occurrence at Cedar Bog

The distribution of Vertigo milium shows an association with moist to wet areas, but not the wettest areas such as the marl meadow or the tuliptree swamp forest (Map XXXVI). It is found most often in peat and muck substrates. It is also most frequently found in the arbor vitae association, the bog meadow association and the elm- maple swamp forest areas which border either the arbor vitae or bog meadow (Plate IV, Figure 4). Most living specimens were made in the bog meadow and arbor vitae associations. This species is definitely absent from the driest portions of the study area.

The habitat diversity indices (Table 10) vary from medium to high. This index variability Is the result of very different Inter­ pretation of the arbor vitae association zonation by Frederick and E. C. C. It is evident from these results that vegetation boundaries are very important In the determination of habitat preferences for this species.

Even though the frequency data are fairly close for total and living numbers collected in the arbor vitae and bog meadow zones, the densities are very different. The total average density is 8.1/ ft^ for the arbor vitae and 41.8/ ft^ for the bog meadow. The living aver­ age density was 1.6/ ft2 for the arbor vitae and 1.4/ ft^ for the bog meadow.

Columella edentula (Drapernaud, 1805)

Distribution

This species is Kolarctic, being widespread in Russia, Europe and North America (Likhachev and Rammel'meier, 1952: 144). In North America it is found from Laborador south to North Carolina in the east, and Alaska south to California in the west and from northern X 5 5

MAP X X X V I

Collection locations for Vertigo milium (Gould) at Cedar Bog.

J I w T

D e c . 23 & 2 5 , 1968 Jan. 5, 1969 • C ollected A live ( t ° C ollected D ead

April 23 & 25, 1969 A Collected Alive A C o lle c te d D ead

August 20 & 30, 1969 ■ C o lle c te d A liv e D C ollected D ead

Miscellaneous Collections © Collected Alive © C ollected D ead

Wnfiifkntft tn cd

10 to

2 2 0 - ' 440

12 12 --- r :r w 156 parts of Georgia, Alabama and Texas (La Rocque, 1970: 754). It is generally absent from the southwestern and southeastern parts of the United States. This species is most common in the northern Mississippi drainage. In Ohio it has been recorded from ten counties (La Rocque, 1970: 754).'

General Habitat

In Russia, "the species lives mainly in plains and mountain valleys, though it is capable of ascending to 2000 meters above sea level. It favors humid places with a rich grass cover but also lives in forests and shrubs'* (Likhachev and Rammel’meier, 1952: 144). In England, this species is found in any area which has adequate shelter and moisture (Boycott, 1934: 33). Boycott also found that this species could inhabit Sphagnum bogs as well as chalk beech woods.

E. C. Baker (1939: 109), in Illinois, and Goodrich and van der Schalie (1944: 277), in Indiana, found this snail in small numbers under organic debris on floodplalns. Archer (1936: 13) and Harry (1951: 32), in Michigan, Oughton (1948: 94) in Ontario, and Burch (1954: 31) in Virginia, associated jC. edentula most often with decidu­ ous forests. In some areas Harry found this snail associated with arbor vitae.

On Mackinac Island, Michigan, Archer (1934: 138-140) and in Maryland (Grimm, 1959: 126) found this species in rocky areas such as a limestone cliff and a marble quarry.

Occurrence at Cedar Bog

Columella edentula is a snail of Intermediate moisture conditions at Cedar Bog. I found it most often in the arbor vitae and elm-maple swamp forest, two very different plant communities. The distribution of C^. edentula is fairly even throughout most of the Bog (Map XXXVII). However, it is not found in the wettest area, the marl, meadow.

Only a total of 317 specimens were collected in the 65 samples it occurred. The highest density occurred in the elm-maple swamp forest with an average total density of 9.5/ ft^. The arbor vitae association with about the same collection frequency only had an average total density of 4.3/ ft^. The density and collection fre­ quency for living specimens was low throughout the Bog.

The hab'itat diversity index is high for this species. The dif­ ferent indices varied from 2.39 for living specimens collected to 2.80 for the total number collected in the various vegetation zones. These high indices reflect its wide habitat diversity coupled with a relatively even distribution. Boycott's comment about this species 157

m a p xxxvn

Collection locations for Columella edentula (Draparnaud) at Cedar Bog.

T I T w T

Dec. 23 & 25. 1968 Jan. 5. 1969 • Collected Alive o C ollected D ead

April 23 & 25, 1969 A Collected Alive a C ollected D ead

August 20 & 30, 1969 * C ollected Alive ° C ollected D ead

Miscellaneous C ollections Q C ollected Alive 0 C ollected D ead

nott^hum Eacri

Icnv Frequency (Si) Frequency (?Q 100 100 40 30 -30 80 6o 20 0 5 10 H 70 i ° 5 20 80 0

- -

- - . Figure Figure Figure Figure

ABODE 3 1 »frequenciesfor Collection .frequencies for Collection Strobilops labyrinthica. Strobilops Colunellaedentula. Vegetation Zones Vegetation Vegetation Zones Vegetation i *.i n iiir >.Kb: rGrtJ i' ■‘ ii • wm K FLATE V FLATE • S U. SR lu 5* 6o- S t. a cr 3 5- 100 100 A 20 80 10 - 0 3 0 7 - 0 5 0 5 6o 0 7 80 10 90 -90 0 0 ------• - - - - .

Figure

Figure ABCUEFGHJK 2 4 .frequencies Collectionfor ,frequencies for Collection .v%O/V'T Strobilop3 oenea. c Succinea avails.Succinea y- 1 ? Vogotation Zones Vogotation Vegetation Zones Vegetation E U r 0 J H o 158 159

living- in anything from a Sphagnum bog -to a chalk beech woods appears to be true at Cedar Bog. Goodrich and van der Schalie (1944: 277) mentioned it being found in low numbers and that is also true.

Strobilops aenea Pilsbry, 1926

Distribution

Massachusetts, southern Ontario west to Wisconsin, south to Louisiana in the west, and south to Florida in the east (La Rocque, 1970: 716). Pilsbry (1948: 863) states that this species is the common Strobilops on the Gulf Coastal Plain. La Rocque (1970: 716) records it from eleven counties in Ohio, most of these are in the northwest quarter of the state.

General Habitat

In Illinois, "the usual habitat of Strobilops aenea is in higher uplands, in forests of oak, elm, hickory, dogwood, walnut, sassafras and ironwood. Only rarely is it found in a floodplain valley and then only when the situation Is dry. It is often abundant under loose or started bark, on the underside of old wood and on chips from forest cuttings. It occurs in old wood and.In recent cut-overs wherever logs and decaying wood have accumulated" (F. C. Baker, 1939: 114-115).

Oughton (1948: 94), In Ontario, and Michelson (1953: 51), In Florida, both associated j3, aenea with deciduous forests. In Florida It was also found in scrub, sinks, and rocky outcrops. In Ohio, Dexter (1950: 26) and In Indiana Goodrich and van der Schalie (1944: 279-280) found J3. aenea in deciduous forests in upland areas.

Archer (1941: 10) states that the usual habitat for this species Is under loose bark on logs. Burch (1955: 161) in Virginia, supports Archer and also associated it with soils with a pH range of 6.3 to 6.7 and very high calcium.

Occurrence at Cedar Bog

Strobilops aenea occurred in only eight samples. Six of these eight samples were made by random collecting which means that areas other than soil and litter were searched. This is not a soil and litter species at the Bog, and Its usual habitat is under loose bark and logs. Ohly 17 specimens were found in these eight samples.

I found _S. aenea only in forested regions of the arbor vitae, swamp forest and beech-maple (Map XXXVIII). This follows, because of its usual habitat is under bark of dead or fallen trees. Plate V, M A P X X X V IU

Collection locations for Strobilops aenea Pilsbry at Cedar Bog.

I I w I

Dec. 23 St 25, 1968 Jan. 5, 1969 • Collected Alive ° C ollected D ead

April 23 St 25, 1969 * C ollected Alive £ C ollected D ead

August 20 & 30, 1969 ■ C ollected Alive D C ollected D ead

Miscellaneous Collections © Collected Alive 0 C ollected D ead

We oitbm a-lne 4

5lo 440

i* m . «*asoit w r 161

Figure 1 shows the collection frequencies for the four vegetation zones in which it was foundi It was most frequently collected in the tullptree segregant of the swamp forest. The habitat diversity index is low for this species (Table 10) because it was based only on the soil and litter samples.

Strobilops labyrlnthica (Say, 1817)

Distribution

Southeastern Canada west to Manitoba, south to Texas in the west and south to Georgia in the east. It is found in many widely scattered localities in Ohio (La Rocque, 1970: 713).

General Habitat

Oughton (1948: 94) considered Strobilops labyrlnthica a species of damp deciduous woodlands in southern Ontario. In the upper peninsula of Michigan, Walker and Ruthven (1906: 94) found S_. labyrlnthica In a hardwood forest, an aspen forest, a tamarack swamp and other coniferous forest communities associated with damp leaves, logs, roots of ferns and humus. H. B. Baker (1922: 26) also working in the upper peninsula of Michigan, found this species to be the most common small land snail in the study area. He found it in conif­ erous and deciduous forests in 13 of 14 vegetation zones examined. He stated that It was abundant in hardwoods and the drier habitats. F. C. Baker (1935: 257-274) working in Minnesota, also found S>. labyrlnthica everywhere, deciduous forest, coniferous forests and mixed deciduous and coniferous forests. It was common in the northern white cedar bogs.

Further south In this species' range Archer (1936: 13) working in the lower peninsula of Michigan, found that "this is one of the com­ monest of the small species, and occupies hardwoods,aspens, and shrubby areas. It lives in leaf mold, under fallen bark, rotten logs, and at the base of stumps. In open, shrubby areas it inhabits plant debris at the bases of stumps, and also occurs under boards and palings." Harry (1951: 32) found it quite common in the northern white cedar bogs and deciduous forest In lower Michigan.

In Ohio, Sterki (1907: 378) stated that £5. labyrlnthica preferred damp places. In Indiana, Goodrich and van der Schalie (1944: 279) associated it with wet ground under rotting wood, very open places within a few feet of standing water. In Illinois, F. C. Baker (1939: 111-112) found that "its habitat is in river valleys, on large river bluffs, in woodland areas and less often in second growth woods and on cut over lands. Oak, elm, hickory, basswood, walnut and in places pine are its natural cover." "It is not common In floodplain areas of Illinois. 162

Leonard (1959: 167) associated jS. labyrlnthica with organic debris in the forested areas of Kansas. In Georgia, Teskey (1955: 69-71) found j^. labyrlnthica in leafmold on loose shale near a water falls, in detritus in the holes of a stone wall and in the rotted wood of an old mill*

In the northern part of this species1 range it is associated with a wide variety of habitats and Is very common. In the southern part its range is most often associated with deciduous forests and fre­ quently in and around decaying wood and bark.

Occurrence at Cedar Bog

I found specimens of Strobilops labyrlnthica in every vegetation zone within the study area. The distribution map (Map XXXIX) for £>_. labyrlnthica indicates a fairly wide distribution over the northern part of the study area and along Cedar Run in the southern part of the study area with few collections sites in the southwestern section. The habitat diversity Index is medium to low (Table 10). This shows a habitat specificity which is not readily apparent on the distribution map. It becomes more noticeable when one looks at the collection frequency data (Plate V, Figure 3). J3. labyrinthicais present In 72% of all the collections made in the marl meadow. The density data indicates why this species has such a low habitat diversity index. Of the 1283 specimens collected in all the quantitative samples, 805 were found in the marl meadow and 285 were found in the bog meadow. There was a total of 111 living snails collected and 92 were from the marl meadow, Strobilops labyrlnthica appears to be strongly associ­ ated with the wettest habitats and especially the marl meadow. At Cedar Bog, this snail is not common in organic debris in the deciduous forests.

Succlnea ovalis Say, 1817 % Distribution

Newfoundland west to Alaska, southeast to Montana, Nebraska, Kansas, Oklahoma, Arkansas then east to Georgia. It is found in numerous counties throughout Ohio (La Rocque, 1970: 709).

General Habitat

Succinea ovalis is one of the largest and most commonly collected members of the family Succlneldae in eastern North America. It has a surprising adaptability to many diverse habitats.

In Michigan, F. C. Baker (1910: 459) and H. B. Baker (1911: 142- 143, 148-149) found It in swamp forests. In New York, Ingram (1946: MAP X X X IX

Collection locations for Strobilops labyrlnthica (Say) at Cedar Bog.

* | s i T ! U I V I W '} X I 1 I I

• Collected Alive ° C ollected D ead

A p r i l 23 & 2 5 , 1969 * C ollected Alive 6 Collected Dead

A u g u s t 20 & 3 0 , 1969 ■ C ollected Alive D C ollected D ead

Miscellaneous C ollections © C ollected Alive © C ollected D ead

WmujWirr. tn.H 164

92) found it characteristic.on floodplains and common in beech-hemlock forests and bogs. In Illinois, F. C. Baker (1939: 122) associated this snail with river valleys and floodplains. In Indiana, Goodrich and van der Schalie (1944: 281) found it inhabited damp woods. In Ontario, Oughton (1948: 75) associated this species with damp and dry deciduous woods and fields. Gugler (1963: 197-198) collected it in Iowa under heavy leaf litter on north facing slopes of ravines.

"Succinea ovalis is an Inhabitant of moist situations near ponds, swamps, and streams, often among trees or shrubs. It is abundant along the floodplains of the Missouri River in western Iowa, eastern Nebraska, and northeastern Kansas, where it lives among grasses and hedges in mud flats, but often ascends the wooded bluffs where mois­ ture is abundant. Its preference for moist environments is so charac­ teristic that its distribution on a wooded slope may be suddenly truncated above a horizon where contact springs emerge" (Leonard (1950: 24-25).

Strandine (1941: 86-91) studied a population of Succinea ovalis in Illinois in some detail. He found this species on old floodplains not subject to flooding anymore. He also found that "the fluctuations in the density of the Succinea ovalis populations coincides with the fluctuations in the soil moisture, the organic matter and the pH; the snail population increasing when the pH is increasing and the soil moisture and organic matter are decreasing." He also found that "temperature and rainfall are Important influences In determining the size of the Succinea populations."

It appears from Strandine’s work that too much moisture is detrimental to the population and from Leonard’s statements that too little moisture also limits their distribution. Since most investi­ gators did not monitor moisture conditions in the areas S^. ovalis was collected, it is difficult to say what exactly the moisture require­ ments are throughout its range. The mental picture I have concerning this species' habitat preferences at this point are not contradictory. However, this is clouded by the following habitat Information regard­ ing this species.

In Iowa, Shimek (1935: 6-9) found Succinea ovalis in many scattered areas in all wooded parts of the state. In the eastern part of Iowa it was in low alluvial woods subject to overflow. In the western part of the state this species was distinctly xerophilous and usually associ­ ated with upland bur oak groves or in thickets bordering them.

Archer’s works in the lower peninsula of Michigan add further inconsistencies. In Cheboygan county, Michigan, Archer (1939: 24) found ji. ovalis "in grassy marshes; In the oak-hickory woods; in grass, in the axils of burdocks, and on fence posts in the open fields and along roadsides. This snail avoids summit areas of all kinds." In Livingston county, Michigan, Archer (1936: 14) states that "this 165

Succinea seems to avoid damp habitats. It lives in leaf mold In hardwoods. In open fields it inhabits grasses and weeds, and along roadsides congregates under boards and rubbish."

Climatic conditions and genetic variation in the various popula­ tions probably account for its habitat inconsistencies throughout its range.

Occurrence at Cedar Bog

Succinea ovalis may not be the densest or most abundant species in the Bog, but in many areas it is one of the most commonly encoun­ tered species. This snail Is fairly large and it climbs up several feet on all types of vegetation. Its size, color and position on the vegetation gives the appearance of the most dominant species in the Bog. The distribution of Succinea ovalis at Cedar Bog is most inter­ esting and quite different from that of any other succineid (Map XL). This snail is so easy to collect that after I made a collection of them In a particular vegetation zone I would move on to other things. Consequently, its distribution Is far more extensive within the maple-ash-basswood, tuliptree and beech-maple forest than it appears on the map. Even so, Succinea ovalis has definite habitat preferences within the study area.

I collected it most often in the drier forests of the silver-red maple and beech-maple forests. Second In importance were the wetter swamp forests. This snail is largely absent from the wettest areas of the study area. I did not even find dead shells in the marl meadow and rarely found specimens in the bog meadow and arbor vitae zones. Succinea ovalis was not commonly encountered in the northern half of the study area. At Cedar Bog this species definitely associated with forested regions and not open areas such as the meadows and elm-maple swamp forest.

The habitat diversity index for the total number collected is 2.14 and the habitat diversity index for the living number collected Is 0.81. This difference could be Interpreted in at feast two differ­ ent ways. Succinea ovalis Is a climbing species, its presence in soil and litter samples is fairly consistent, but usually in low numbers and almost always dead. The diversity indices are based only on the data collected by quantitative soil and litter samples, therefore this method selected for dead specimens and against living specimens. If this species were normally a soil and litter inhabitant, then this great difference might be interpreted as being due to a habitat change. This last interpretation might still be true, but the methods I used to sample this species could not possibly detect this. 166

MAP X L

Collection locations for Succinea ovalis Say at Cedar Bog.

I ! » r 1— W—T

D e c . 23 & 2 5 , 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

April 23 & 25, 1969 * C ollected Alive o C ollected D ead

A u g u s t 20 & 30, 1969 ■ C ollected Alive ° C ollected D ead

Miscellaneous Collections © C ollected Alive © C ollected D ead

W f t a d k o r n Vn~

10

O 1 2 0 440

I cm. la n z E Z i 167

Catinella vermeta (Say, 1829)

Distribution

Tentatively, the distribution of this species is New Jersey west to North Dakota, south to Arizona in the west and south to Georgia in the east (extracted from Bequaert and Miller, 1973: 157; La Roque, 1970: 700 and 704). Most states within these boundaries have records for Succinea avara vermeta, Catinella vermeta or Catinella vagans (Pilsbry, 1948: 838 and 844). Catinella vermeta was referred to as Succinea avara vermeta in Taft (1961: 56) and was found in Hamilton and Tuscarawas counties. La Rocque (1970: 703-704) listed S^. avara as being widespread in Ohio with the exception of northeastern Ohio. The range of this species is at best uncertain at this time.

General Remarks

According to Hubricht (1958: 60-61), Thomas Say's Succinea vermeta of 1829 from New Harmony, Posey county, Indiana belonged to the genus Ouickella (C. R. Boettger, 1939) based on internal anatomy. Since that time it has been discovered that Quickella is the same as Catinella (W. H. Pease, 1871) according to Bequaert and Miller (1973: 154). Therefore, Succinea vermeta should now be referred to as Catinella vermeta (Say, 1829).

This sounds simple and really uncomplicated except that Pilsbry (1948: 837) and F. C. Baker (1939: 124) synonymized Succinea vermeta with Succinea avara (Say, 1824). Hubricht (1958: 61) states that the type of S^. avara is an immature shell and appear to be different from the types of JS. vermeta. The type locality for ji. avara is presumed to be Minnesota. Hubricht believes that the exact identity of avara is uncertain and the avara-like snails from the midwest are Quickella vermeta (Succinea vermeta). However, since vermeta and £». avara were synonymized many people have referred to all avara-like shells as S^. avara and it is not clear in the literature if all the records for distribution and habitat really belong to S^. avara or C. vermeta. Bequaert and Miller (1973: 157) consider S_. avara and C. vermeta distinct species. They also believe Catinella vagans (Pilsbry, 1900) is conspecific with vermeta.'

I have dissected a few of the specimens I found at Cedar Bog and have confirmed that they are indeed members of the genus Catinella.

General Habitat

In the literature, Succinea avara is found in wet or dry deciduous and coniferous habitats. It has been found on dry limestone mountains of Alabama and in grass along lake shores of Michigan (Archer, 1936: 168

14). F. C. Baker (1939: 124) who called _S. avara and S^. vermeta absolute synonyms states that the "varied habitats of these snails include oak, elm, walnut, and ironwood in hillside regions; oak, elm, birch, beech and maple in floodplain localities; grass, weeds and low bushes in railroad embankments."

In Michigan, H. B. Baker (1911: 158) found S^. avara vermeta in sand pools and sandy beach drift along a lake. Pilsbry (1948: 839) states, "S^. avara is usually found in vegetable debris thrown up on muddy shores, or crawling on muddy banks of ditches, often exposed to the sun; also in swampy places in pastures, as I have found it in Essex County, N.Y. But it is an upland species as well, to be seen under stones with Pupillidae, or occasionally after rains crawling up the trunks of trees. In such relatively dry places it does not reach the size of individuals living in humid situations."

"Though conspicuous differences in texture, size and shape, at least partly correlated with station, are seen In specimens from most parts of the wide range of avara, I have not been able to satisfy myself that there are any subspecific forms. I am disposed to view the variations as direct reactions to environment, the small upland form (typical avara) being to some extent a 'hunger form,1 while that called avara vermeta is the full development of the species in humid places. However, the variations observed in the genitalia may indicate the existence of several species or races."

Occurrence at Cedar Bog

I have collected very few living specimens of this species any­ where In the study area. Dead shells of jC. vermeta do occasionally turn up in soil and litter samples. The distribution map (Map XLI) for jC. vermeta shows a strong association with the wet areas along the East Branch of Cedar Run. The frequency graphs (Plate VI, Figure 1) show the same correlations. Why I could not find more living speci­ mens within the marl meadow is a mystery.

The best collection of this species was not made "in the study area. I collected a series around a temporary pond in the old field west of the study area. This snail is not a forest species, but more common in open areas such as the bog meadow, marl meadow and the old field. The habitat diversity Index is low.

Oxyloroa retusa (Lea, 1834)

Distribution

"Ohio west to Montana, south to Kansas" (Hibbard and Taylor, 1960: 141). 169

MAP X L I

Collection locations for Catinella vermeta (Say) at Cedar Bog.

w T I

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

A p r i l 23 & 2 5 , 1969 A C ollected Alive A C ollected D ead

August 20 & 30, 1969 ■ Collected Alive D C ollected D ead

Miscellaneous Collections © C ollected Alive © C ollected D ead

Wonrfhuto_gonA

to

0 220 » 1cm. alJOIt 12 , ---- ,---- p .— - r ---- r ~ " Frequenqy (ji) Frequency (%) 100 100 80 80 30. 30. 40 ■ 60 70 90 90 50 20 10 0 - -

. ■ • -

Figure Figure Figure Figure ASCDEFGHJ 3 1 * Collection frequencies for *frequencies Collection .for frequencies Collection C D S F G H J J H G F S D C Catinella vermeta.-Catinella Philooycus carolinianus. Philooycus 1 Vegetation Zones Vegetation Vegetation Zones Vegetation PLATE VI PLATE K ti. U O 13 u o’ n iT c_ a* cr* 100 100 - 20 - 20 0 5 80 100 - 0 3 (So- -- 0 1 - 0 9 20 0 3 40 80 80 50 0 5 6o 70 10 0 — j

0 0 -

- - - .

. - - Figure 1 1 Figure Figure .*-sr i.v? ViVt , A t *

•* i k I .-ji: -•V*. • i *« d 2 4 • . frequencies for Collection .for frequencies Collection Oxylona cpp. Oxylona C C iwA Anguispira alternate. Anguispira Vegetation Zones Vegetation Vegetation Zones Vegetation 0 0

E F G G F E £ C J H C F m v». ;.' ,ii*I J K J H a ‘ 170 5 W A>* 171

General Remarks

The taxonomy of the Succineidae is not uniform. A number of • species have been described only on the basis of shell characters. Shell characters are not reliable in this family. Franzen (1963: -82) states that because of this dependence on the shell, "a great deal of the systematics within this family are in a state of error and con­ fusion." Therefore, data on the distribution and ecology may also be in error. The specimens in this study were tentatively identified by Dr. Dorothea S. Franzen as Oxyloma retusa and Oxyloma decampi gouldi. Originally, I thought I had only one species of Oxyloma, therefore the distribution map and all the data reflects this. It may be impossible to separate all the dead shells collected in any event.

General Habitat

Oxyloma retusa is a species of vet habitats. H. B. Baker (1922: 15), Goodrich (1932: 38), Archer (1939: 23), Oughton (1948: 95) and Hibbard and Taylor (1960: 142) found this species common around bodies of water, either as ponds, marshes, bogs, swamps, pasture pools, streams or springs. The source or type of water does not seem to be important. The vegetation present can also be varied, H. B. Baker (1922: 15) found 0 . retusa abundant in an ash-cedar swamp, Archer (1939: 23) found them near an oak-hickory woods on Sphagnum and sedges. Goodrich (1932: 38) and Hibbard and Taylor (1960: 142) state that 0. retusa is not strictly an aquatic species, but its habitat overlaps with that of various Lymnaea.

Shrader (1972: 12-13) did an interesting study on the feeding behavior of three species of succineid snails in lower Michigan. She studied Oxyloma retusa, Succinea ovalis and Catinella vermeta, the same three species found at Cedar Bog. In her study area <). retusa hardly ventured from the water. She found them all tolerant of moisture and sensitive to desiccation. Her study also concluded that each species ate a different amount and/or kind of diatoms, green algae, blue-green algae and pollen. .

Goodrich (1932: 38) and Archer (1939: 23) indicate that 0. retusa is a climbing species, especially on sedges.and cattails.

Occurrence at Cedar Bog

At Cedar Bog, Oxyloma retusa and/or Oxyloma decampi gouldi are found in the' wet habitats. Most of the living specimens are obtained around springs, temporary ponds, along the stream banks in several vegetation zones, in addition to the bog meadow and marl meadow. I did not find either of the Oxyloma in the drier parts of the Bog (Map XLII). Most of the specimens collected, dead and alive, were from the 172

MAP X L II

Collection Locations for Oxyloma spp. at Cedar Bog

w T I '* ' '"1

D e c . 23 & 2 5 , 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

April 23 & 25, 1969 * C ollected Alive & C ollected D ead

A u g u s t 20 & 3 0 , 1969 ■ C ollected Alive D C ollected D ead

Miscellaneous Collections © C ollected Alive © C ollected D ead

I cm. aillQll 173 bog meadow, marl meadow and bordering arbor vitae. These snails were most often found climbing on the sedges and cattails.

The habitat diversity indices are medium to low, but cannot be too meaningful because of the climbing nature of these species and the fact that there are two species mixed together. The habitat diversity index would probably be even lower for each of these species.

At Gedar Bog, all four succlneids may be found in the same habitat, but probably not very often. Succinea ovalis and the two Oxyloma species are climbing species and in some areas of the bog meadow they occur together, Catinella vermeta is not a frequent climber and is usually found on the substrate or some low wet log. At times all the succlneids may be on the substrate at the same time.

When looking at the distribution maps of all the succlneids, it is evident that Succinea ovalis is much more common in the drier por­ tions of the Bog where the other species are rare. Succinea ovalis has not been collected in the marl meadow where the other three species are common. Additional work needs to be done on the distribution patterns and ecology of all the Succineidae of Cedar Bog. A better understanding of the taxonomy will lead to a better understanding of their ecology.

Oxyloma decampi gouldi Pilsbry, 1948

Distribution

Maritime Provinces of Canada, Ontario and New England, the middle states and south to Maryland, west to Colorado and Montana (Pilsbry, 1948: 782).

General Remarks

This species is a recent addition to the Cedar Bog list. Doctor Franzen (1975) identified these snails after a lengthy investigation. I could not separate the two species of Oxyloma, so the data reprer- sents both species. Pilsbry (1948: 782) included Ohio in his gener­ alized distribution for JO. decampi gouldi. La Rocque (1970: 695-697) also included this species in his work on the Pleistocene mollusca. He gave five uncertain fossil localities for Ohio. Apparently this is the first authenticated living record for this species in Ohio. Pilsbry (1948: 782) considered many of the Oxyloma retusa records to really be those of Oxyloma decampi gouldi. 174

General Habitat

"This is the Oxyloma of marshy places, being common on and around the aquatic vegetation of muddy pond and river margins and ditches" (Pilsbry, 1948: 782).

Occurrence at Cedar Bog

See the discussion on Oxyloma retusa.

Philomycus carolinianus (Bose, 1802)

Distribution

Maine to Florida, west to Iowa, Kansas and Oklahoma (La Rocque, 1970: 691).

General Habitat

Ingram (1949: 86-90) worked In New York and probably took the most intensive look at Philomycus carolinianus. He found this species commonly associated with beech-hemlock forest and never In abandoned fields or human habitation. It specifically avoided the hemlock trees for beech trees. This slug was active all day, it had definite arboreal tendencies and was gregareous. In Kansas, Leonard (1959: 139) found carolinianus in forests beneath trunks of fallen trees, started bark and in deep leaf litter. He never found it associated with urban areas, basements, houses or greenhouses.

Goodrich and van der Schalie (1944: 279), in Indiana, Oughton (1948: 94) in Ontario, Michelson (1953: 5) in Florida, F. C. Baker (1939: 132) in Illinois and H. B. Baker (1911: 157) in Michigan, all associated jP. carolinianus with established wooded areas.

In Michigan, Archer (1936: 9) and Archer (1942: 96) in Alabama, found some P^. carolinianus associated with pine communities. Rapp and Rapp (1945: 125) found this slug to be the most common mollusc in the Great Swamp of New Jersey where the pH of the substrate was as low as 4.

"P. carolinianus Is a relatively inactive slug. It Is widespread but rarely locally abundant. Although native to our region, its biology is not well known. It Inhabits both deciduous and coniferous woods where'it is usually found In decaying logs. It Is more common in deciduous forests, especially beech and basswood" (Chichester and Getz, 1973: 32-33). 175

In Ohio I have never collected £. carolinianus outside of a deciduous woods and almost always on fallen logs under loose bark.

Occurrence at Cedar Bog

Philomycus carolinianus is unique at Cedar Bog, it is the only mollusc which does not have a shell. Consequently, it is never found dead. This species never turned up in any of the soil and litter samples. I found it only while making random collections under loose bark on fallen trees. This is definitely not a soil and litter species at Cedar Bog. I do not have any data on density and I could not cal­ culate a habitat diversity index.

The distribution of this species (Map XLIII) and the frequency graph Plate VI, Figure 3) show that I never found it dead and only in four contiguous vegetation zones north of Uoodburn Road. I believe there are two factors which regulates the distribution of this species at Cedar Bog, humidity and cover. I found this species only in forested areas which have fallen trees and only where there are enough springs and temporary ponds to keep the humidity high.

Anguispira alternata (Say, 1817)

Distribution

In the north it ranges from southeastern Canada and Maine west to the eastern edge of Minnesota, Nebraska and Kansas and in the south to ranges from North Carolina west to Alabama and Texas. It is absent from South Carolina, Georgia and Florida (La Rocque, 1970: 673). It is one of the most common snails found throughout'Ohio.

General Habitat

Compared to most species of terrestrial gastropods, Anguispira alternata Is biologically well known. The most Intensive study was done by Elwell and Ulmer (1971). They studied its life history in some detail. Gugler (1963) observed its egg laying habits, Jones (1935) its burrowing habits and Douglas (1963) various behavior patterns and growth rates.

Anguispira alternata is one of the most common large snails found in eastern United States. It has been recorded from a wide variety of habitats. Although, several investigators were quite specific as to where this species was found. Sometimes this information was contra­ dictory. Upon close examination I find that Anguispira alternata does have some specific habitat preferences In different parts of Its range. In New York, Ingram (1946: 90) stated that A. alternata 176

M A P X L I1 I

Collection locations for Philomycus carolinianus (Bose.) at Cedar Bog.

r i w T » i r:

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

April 23 & 25, 1969 * C ollected Alive o C ollected D ead

August 20 & 30, 1969 ■ C ollected Alive ° C ollected D ead

Miscellaneous C ollections 9 C ollected Alive 0 C ollected D ead

Woqdihmn toed.___ 177

preferred floodplain areas and specifically avoided bogs and fields. In Iowa, Elwell and Ulmer (1971: 204) found that this species was predominantly found In well established deciduous forests. Douglas . (1963: 186) and Leonard (1959: 130) state that in Kansas it is most common in wooded areas whether they were floodplains or uplands.

Host of the habitat information available is from Michigan, Ohio and Illinois. In these states Anguispira alternata is not only common in floodplains, and upland woods, but from old fields, pastures, swamps and marshes.

In Ohio, Dexter (1950: 24) found this species common In the alder shrub, willow and swamp forest zones surrounding a bog lake and Archer (1937: 120) considered A. alternata an abundant snail along walls, roadsides and railroad embankments.

In lower Michigan, H. B. Baker (1911: 156) found it In 12 of 21 habitats investigated, these included swamps, driftwood along beaches, under stones In dry meadows and dry woods. In later studies in upper Michigan, H. B. Baker (1922: 25) found It in schist rock outcroppings along a river valley, stream valleys dominated by white and red pines, and several mixed deciduous forests in upland areas. Archer (1936: 9) found A. alternata in a wide variety of habitats, in deciduous forest hillsides, pine groves, bog woods dominated by arbor vitae and spruce and in old fields.

F, C. Baker (1939: 84) summarized A. alternata habitats In Illinois. "It occurs under many conditions, in wet or dry habitats, in forests, on limestone ledges, on floodplains and railway embank­ ments. It is characteristic of the started or loose bark habitat." From my experience, I believe that Baker's summary would fit Anguispira alternata in Ohio very well.

Burch (1955: 66 and 1956: 61) found this snail in Virginia mainly in lowland deciduous forests where the pH varied from 5.3 to 7.7, the calcium and organic matter content of the soil was very high and magnesium and phosphrous content was medium.

The habitat of Anguispira altenate is not the same In all parts of its range, but some of its behavioral patterns are distinctive and consistent. Goodrich (1932: 34-35), Call (1900: 380), Jones (1935: 141-142), Archer (1936: 9), MacMillan (1940: 380-381), Leonard (1959: 130) and Douglas (1963: 186) found this species to be quite gregareous at times, frequently during hibernation. Jones (1935: 141-142), Douglas (1963: 192-193) and Gugler (1963: 198-199) also noted Its intensive burrowing activities before hibernation. Archer (1939: 19) and F. C. Baker (1939: 84) noted Its frequency in climbing trees. In collecting this species in Ohio and keeping them for extended periods in the laboratory I have found it to be a burrower especially when conditions are dry or when they are laying eggs and a climber when the 178

MAP X L IV

Collection locations for Anguispira alternata (Say) at Cedar Bog.

1 " C T - T ! V~I w i— r

D e c . 23 & 2 5 , 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

A p r il 23 & 2 5 , 1969 A C ollected Alive & C o lle c te d D ead

August 20 & 30, 1969 ■ Collected Alive ° C ollected D ead

Miscellaneous C ollections Q C ollected Alive © C ollected D ead

Woa£buxn_2csd. 179 conditions are wet and humid. X have also found large numbers under loose flat rocks and loose bark.‘

Occurrence at Cedar Bog

At Cedar Bog there are two factors which seem to govern the distribution of Anguispira alternate, moisture and cover. Anguispira alternata is not found in areas that are wet all the time or flooded part of the year. It can occur in some fairly dry habitats. The other factor is cover or shelter. Anguispira alternata is like many of the large species of terrestrial mollusca, it must have sufficient cover. At Cedar Bog, cover for this species is usually loose bark on or off the tree. Since this snail is relatively large, the bark must be fairly loose and easy to crawl under. This combination was not common throughout the Bog when the study was started. As the dead elms age and rot, this species may become more common in some areas.

Anguispira alternata does pot lend itself to the soil and litter sampling procedure. It is a log species and a climber. Living specimens do not turn up in the soil and litter very often at Cedar Bog.

Helicodiscus parallelus (Say, 1821)

Distribution

Northeastern Canada west to Manitoba, south to northern Florida in the east and south to Texas in the west (La Rocque, 1970: 685). Generally absent from the Gulf Coastal Plain. La Rocque also records this species from Idaho, California, Colorado and New Mexico, This species is distributed throughout Ohio.

General Habitat

Helicodiscus parallelus Is a fairly common woodland snail. Oughton (1948: 94) and Harry (1951: 32) associated this species with unspecified deciduous forests. Burch (1956: 61) collected it in 21 of the 27 plant communities examined in his Virginia studies. Most of these communities were deciduous forests. F. C. Baker (1910: 478 and 1939: 89), H. B. Baker (1911: 157 and 1922: 29-30), Ingram (1941: 15) and Goodrich and van der Schalie (1944: 275) found H. parallelus in deciduous floodplaln communities in Michigan, Illinois, Indiana and New York. In certain parts of this species’ range It is found associ­ ated with swamps and bogs. H. B. Baker (1922: 29-30) found It in a drained tamarack-cedar swamp and Walker and Ruthven (1906: 94) found it in a tamarack swamp. H. B. Baker (1911: 157) in Michigan, collected H. parallelus around the borders of swamps, under driftwood around a lake, under logs in swampy and dry woods, and in a moist meadow. 180

Ingram (1946: 91) stated that H. parallelus was the dominant mollusk in bogs from around the fern hummocks in New York. The general opinion is that this species is most often associated with meslc to hydric environments.

There are, however, several studies which stretch this last generalization some. Grimm (1959; 125) found it in quarries, railroad tracks, ruins of old buildings besides woodlands in Maryland. In northern Florida, Michelson (1953: 51) found it in sinks, rock out­ crops, hammocks and cultivated fields. These all could be wet or mesic habitats, but it is not specified.

Archer (1939: 17) investigated several large areas in lower Michigan and his habitat information differs somewhat from most workers. Helicodiscus parallelus is found "In leaf mold and under rotten logs of oak-hickory woods; in grass, moss, In grassy gullies, rock piles, bramble thickets, wild-grape patches, In the open fields; under logs and fallen bark In the fields and on the roadsides. This snail is commonest in the open country." This last statement makes me wonder whether we are talking about the same animal.

Occurrence at Cedar Bog

Helicodiscus parallelus is a species of the moist to dry sub­ strates at Cedar Bog. Its distribution pattern is almost an opposite of many other species whose distribution Is concentrated along the East Branch. H. parallelus is most frequently collected along the West Branch of Cedar Run in the oak-maple, silver-red maple, elm-maple, shrub and beech-maple vegetation zones (Map XLV). These are the five driest vegetation zones studied. I did not find this snail in the marl meadow at all.

The frequency of living specimens is fairly low in most areas (Plate VII, Figure 1). It is moderately common in the oak-maple and beech-maple communities, but I never found it alive there. At Cedar Bog, H. parallelus is not a swamp and bog species. My findings differ significantly from those of several other investigators.

The habitat diversity Index is medium to high for living specimens collected high for the total number collected (Table 10). These read­ ings indicate a rather wide habitat preference. This snail is found in most of the vegetation zones, but almost always in very low numbers. I found only 175 specimens In 51 samples. I always found this species in the soil and litter samples. In other places In Ohio, I usually find this species in rotting logs. 181

MAP X L V

Collection locations for Helicodiscus parallelus (Say) at Cedar Bog.

! j l ZJ w T

D e c . 23 & 2 5 , 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

April 23 & 25, 1969 A Collected Alive A C o lle c te d D ead

August 20 & 30, 1969 ■ C ollected Alive D C ollected D ead

Miscellaneous Collections © C ollected Alive © C ollected D ead

W e odtfautil—Id o d

I cm. fviaolv Frequency ($S) . Frequency (JS) 100 80 80 90 AO 20 6o - 30 30 70 50 10 ;#iv 0

. -

- Figure 3. Collection frequenciesfor 3.Collection Figure Figure Figure 1 .frequencies for Collection W-S Deroceras laeve. Deroceras Helicodiscus parallelus. COE' Vegetation Zones Vegetation Vegetation Zones Vegetation F E VIA-. F^G^ ^ H G ^ F 0 H

PLATE VII PLATE 5* u cr ■»• 100 80 100 30 40 50 60 80 90 70 - - Figure Figure Figure A A : vv-V h C E F E 0 C B '.V.* 2 0 A. .forfrequencies Collection tv*\ Collection frequencies for frequencies Collection Punctum ninutissimura. V . W Nesovitrea binneyana. Nesovitrea c .V .If:! Vegotation. Zones __ Vegetation Zones Vegetation 0 3# ?:■& w i M ir'r" fvr E y F 6 K G I S *?••Tu'i '• »*! Kt,,- H ?.y

*&■ 182 ‘,*V> K 183

Puneturn, mlnutissimum (Lea, 1841)

Distribution

This snail is widespread in North American from 55° Latitude in Canada south to Mexico (L. Rocque, 1970: 689). It is also widely . distributed in Ohio. Bequaert and Miller (1973: 85) believe that many of the records on which the distribution of Puneturn minutissimum is based may be in error. "At present it can only be stated that true P. minutissimum is basically an eastern Nearctic snail, that extends, perhaps sporadically, west of the 100th Meridian."

General Habitat

Punctum minutissimum is one of our smallest North American snails. I am sure it is frequently overlooked. Few investigators have taken care to records it habitat. Leonard (1959: 137) states, "Punctum minutissimum'is a forest snail to be looked for on damp leaves around decaying logs, sticks, and bits of bark." In North Carolina, Jacot (1935: 604-605) collected it In nine different plant associations, but only In the cove hardwoods did he find It common. In Illinois, F. C. Baker (1939: 90) associated I?, minutissimum with forests and forest debris. In Ontario, Oughton (1948: 94) considered It an inhabitant of damp and dry deciduous woods and fields.

Walker and Ruthven (1906) surveyed the northern part of the upper peninsula of Michigan and Isle Royale and collected ]P. minutissimum in an arbor vitae swamp, aspen forest, in tamarack trees in a peat bog and abundantly in the leaves of a deciduous forest. H. B. Baker (1911: 157), also In Michigan, found it in several swamps. Harry (1951: 32) discovered it quite abundant in an arbor vitae bog. Archer (1939: 18) states that P. minutissimum is "abundant In the moist zone of the leaf-mold carpet in oak-hickory woods, especially concentrated in leaf pockets; rare in grass and moss on slopes of old fields along the highway, and in wild-grape thickets on rotten boards of the roadsides."

Most people have associated this snail with moist to wet habitats and find It In various types of organic debris.

Occurrence at Cedar Bog

Punctum minutissimum Is a very common snail of the soil and litter ■ at Cedar Bog. I found it living in every vegetation zone and soil zone in the study area. P. minutissimum has a distribution pattern very much like Gastrocopta pentodon (Map XLVI and Map XXyil). This snail is very common in the arbor vitae association and the elm-maple swamp forest. The total collection frequency for IP. minutissimum Is 89% in 184

MAP X L VI

Collection locations for Punctum minutissimum (Lea) at Cedar Bog.

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

April 23 & 25, 1969 C ollected Alive £ C ollected D ead

August 20 & 30, 1969 ■ C ollected Alive o C ollected D ead

Miscellaneous C ollections © C ollected Alive © C ollected D ead

W e a i i h u r n Bnrrf

Wnv _330ll 185

the elm-maple swamp forest and 74% In the arbor vitae association (Plate VII, Figure 2). This species is not common In the beech-maple or marl meadow zones. Frequency and distribution data indicate a rather wide mesic or Intermediate moisture preference.

The habitat diversity indices are high which Is to be expected with such a wide distribution within the Bog. A look at the density data illustrates how common this species is In some areas. In the arbor vitae association I found a total average of 72.1/ ft^ and a living average of 12.5/ ft^. In the elm-maple swamp forest segregant I found a total average of 56.4/ ft^ and a living average of 11.2/ ft^. In the shrub zones the total average density was 63.8/ ft^, but the living average was very low. In the oak-maple forest, the frequency was high, but the density was quite low. This density data indicates that the arbor vitae association is currently the most ideal habitat for ]?. mlnutissimum at Cedar Bog followed by the elm-maple swamp forest and the tuliptree swamp forest which has an average living density of 12.9/ ft . The high living density found in the tulip­ tree swamp forest must be qualified. In most areas of the swamp forest segregant it Is very wet and even seasonally flooded. However, In one section in coordinates X, Y - 7, 8 (Map XLVI) this tuliptree zone Is higher and not particularly wet. It also has a consistently good leaf litter. JP. minutissimum is found In very high densities there.

Deroceras laeve (Mtlller, 1774)

Distribution

Its natural range is North America, northern Europe and northern Asia (extracted from Likhackev and Raramel'meier, 1952: 333 and Pilsbry, 1948: 540). It is also recorded from southern Mexico, Guatemala and Nicaragua, but these localities are probably introduced parts of its range (Bequaert and Miller 1973: 68). They also believe it has definitely been Introduced Into Hawaii, Australia, and South Africa.

General Habitat

Deroceras laeve has a wide ecological range and lives In many seemingly diverse habitats. It is found hiding In any number of natural or unnatural articles on the ground. It is frequently re­ corded in areas occupied by man. Deroceras laeve appears to prefer a high humidity for activity whether In a naturally dry or wet environ­ ment. Most of the records for this species in a natural environment are from marshes, swamps or are near bodies of water.

In northern Michigan, H. B. Baker (1922: 26) found D^. laeve in a marsh around a lake; in a pine flat on an outwash plain; in an ash-cedar 186

swamp; a dry alluvial hardwood forest and on floodplains of two rivers. Goodrich and van der Schalle (1944: 278) found it in woods and edges of marshes in addition to numerous man-made environments. In lower Michigan, H.. B. Baker (1911: 156) recorded D. laeve abundant in driftwood, among sedges, floating driftwood in a sedge marsh, under logs along lakes, swamps, wet and dry woods, meadow borders and in a tamarack swamp. In Illinois, Dexter (1953: 29-30) found it along the edges of many temporary ponds.

Getz (1959: 485-498) studied three species of slugs in some detail. He found D. laeve in many diverse habitats, old fields, pastures, hardwood swamp forest, spruce swamp, a sphagnum bog, a sedge marsh and an upland deciduous woods. Getz found that Deroceras laeve was more tolerant to desiccation, subfreezing temperatures, high tem­ peratures and had a greater diversity of food preferences than 15. reticulatum or Arion circumscriptus.

Leonard (1959: 127) states, "Deroceras laeve lives beneath sticks, stones, bark and leaves in woodlands, and beneath boards, logs and even sidewalks In urban communities." "It requires a rather moist habitat, and has been observed crawling about submerged for periods of an hour or more."

La Rocque (1970: 667) summarizes, "This slug prefers humid areas and is particularly partial to floodplains and low terraces of streams; It is also found in or near marshes, in wet weather crawling quite far up the vegetation, in dry weather, it hides under logs, twigs, leaves grass roots, and moss. It Is much hardier than might be Inferred from the above data and will live also in apparently dry situations with good cover and protection from desiccation. In rainy weather, It emerges from cracks between sidewalks or buildings and lawns and its slime trails attest to Its abundance in apparently unlikely situations that are usually dry but afford sufficient moist cover to suit this species."

"D.* laeve is undoubtedly the most ecologically ubiquitous slugs in in northeastern North America. It can be found In greenhouses, gardens, fields, marshes, and deciduous and coniferous woods, "in cultivated areas population sizes approach those of some colonial imported slugs. In wild areas population sizes are generally small" (Chichester and Getz, 1973: 37).

Occurrence at Cedar Bog

When I 'first found this slug at Cedar Bog, I then expected to find it everywhere. After looking at the literature I would also have expected a much greater distribution then it has. Deroceras laeve has a distribution pattern similar to the aquatic gastropods and some Pupillidae. This animal is definitely found more often in the bog meadow, marl meadow and wet parts of the swamp forest. 187

MAP X L VII

Collection locations for Deroceras laeve (Mllller) at Cedar Bog.

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

April 23 & 25, 1969 A Collected Alive ^ C ollected D ead

August 20 & 30, 1969 ■ C ollected Alive n C ollected D ead

Miscellaneous Collections © C ollected Alive 0 C ollected D ead

1«m. - a i o l l 188

This slug has an internal shell which frequently shows up in soil and litter samples. The only living specimens were found by random collecting and in very low numbers. My previous experience with this slug outside of Cedar Bog is quite different. This species is usually quite common when found under loose bark, logs and stones. In the marl meadow, where its shells are most often found, there are very few logs and no bark or stones which might serve as cover. In fact, the only hiding places are In grass or sedge hummocks and Sphagnum. Apparently the perpetually high humidity and an absence of high concentration of predators reduces the necessity of finding this type of cover.

Nesovitrea binneyana (Morse, 1864)

Distribution

Maine and Quebec west to Montana and Colorado, south to West Virginia in the east (La Rocque, 1970: 627). There are two recent records from Ohio (Taft', 1961: 76).

General Habitat

The only ecological information available for Nesovitrea binneyana is from southern Ontario, Minnesota and Michigan. Five of the seven papers I examined that contained habitat data on N. binneyana were from collections made in Michigan.

In southern Ontario, Oughton (1948: 94) records this species from damp deciduous woods and sometimes from Sphagnum bogs. In the lower peninsula of Michigan, Archer (1936: 10) found it only "within the limits of the hardwoods," H. B. Baker (1911: 154) found it among sedges and beach drift along a lake, in wooded flats and wet woods on a river, and Harry (1951: 32) found it in mixed deciduous and conifer­ ous woods, especially in arbor vitae bogs. In the upper peninsula of Michigan, H. B. Baker (1922: 29) found N. binneyana in a cleared and drained tamarack-arbor vitae swamp and Walker and Ruthven (1906: 96), on Isle Royale, found it in alder thickets along a river, In moss in a coniferous forest bordering a river, in arbor vitae swamps, humus under spruce logs and in the moss of a tamarack swamp. F. C. Baker (1935: 257-274) records N. binneyana in Minnesota from many second growth mixed deciduous-coniferous forest and birch-arbor vitae- tamarack forests.

Nesovitrea binneyana appears to be able to survive quite well in coniferous communities. Most of this species' known range is within the mixing area between the coniferous dominated forest of the north and the deciduous dominated forest of the south. It also seems to be partial to wet habitats. 189

Occurrence at Cedar B o r

X am not sure what the distribution pattern (Map XLVIII) or the frequency data (Plate VII, Figure 4) show concerning this species. Its occurrence at Cedar Bog is sporadic and in low numbers* It occurs in the arbor vitae and bog meadow associations which are typical of the habitats it occupies further north. It is also just as common in the tuliptree swamp forest and oak-maple forest. All these zones are very wet in places. Most of the living specimens were collected in random samples taken from arbor vitae litter or from litter very near the arbor vitae stands.

Ohio is on the southern edge of the distribution of Nesovitrea binneyana. I found this species fairly common in the arbor vitae forests in Michigan. It is possible that Cedar Bog is the southern limit of this species. Maybe only relic populations still occur in Ohio. It will take extensive distributional work before this could be proved one way or another.

Nesovitrea electrina (Gould, 1841)

Distribution

Eastern Canada to Alaska, south to South Carolina in the east and south to Arizona and Utah in the west (La Rocque, 1970: 625)". La Rocque also recorded it from several counties throughout Ohio.

General Remarks

"Retinella (Nesovitrea) electrina has usually appeared in our literature under one of the three names commonly applied to a closely allied European snail (Helix hammonis Str8 m, 1767, Helix viridula Menke, 1830 or Helix radiatula Alder, 1830), and in fact the separa­ tion of the American race specifically is more a matter of conven­ ience than definable constant diversity; the subspecific rank seems more fitting" Pilsbry (1946: 258).

Bequaert and Miller (1973: 67) used Nesovitrea hammonis elecrrina (Gould, 1841). At this point in time I am not sure which name to use. I will currently follow the traditional American name Nesovitrea electrina.

Nesovitrea hammonis hammonis (Stroem, 1765) is found throughout Europe and northern Asia. If Nesovitrea electrina is really just a subspecies of Nesovitrea hammons, then this species is really Holarctlc. 190

MAP X D VIH

Collection locations for Nesovitrea binneyana (Morse) at Cedar Bog.

I Z w T

Dec. 23 & 25, 1968 Jan. 5, 1969 • Collected Alive ° C ollected D ead

A p r i l 23 & 2 5 , 1969 A C o lle c te d A liv e C> C ollected D ead

August 20 8i 30, 1969 ■ C ollected Alive ° C ollected D ead

Miscellaneous Collections O Collected Alive 0 C ollected D ead 191

General Habitat

Nesovitrea electrina has been recorded from many different habitats, but it is most frequently associated with fairly moist habitats such as floodplains, edges of marshes, stream and lake margins. Oughton (1948: 27) states, "In southern Ontario, this species is quite restricted, being confined to margins of rivers and lakes and to marshy and oozy ground." H. B. Baker (1911: 154) found N. electrina in several wet habitats in lower Michigan, under driftwood along sandy beaches, around swamps and lakes, wooded flats, swamp forests, damp meadows and in Sphagnum in a tamarack swamp. He also recorded it from a few drier wooded areas. Archer (1939: 19) also in Michigan, found this species in "marsh grass, in leaf mold and under logs of oak— hickory woods; in herbaceous tall weeks, in grass, and under burdocks in open fields; and under boards and shingles in the open slopes and summits."

Leonard (1959: 112-113) summarized the habitat data for N. electrina in Kansas. "This snail has been found in both upland wooded areas and along the margins of streams. It seems to require a fairly moist environment, not occurring out in open grasslands as does Zonltoides arboreus with which electrina is frequently elsewhere associ­ ated. It is to be looked for under sticks, logs, rocks and in crevices of started bark as well as leaf litter."

Occurrence at Cedar Bog

Dead shells of Nesovitrea electrina were found in all vegetation zones at Cedar Bog (Map XLIX). Most of the samples which contained living specimens were collected in some of the wettest parts of Cedar Bog. The general distribution of even the dead shells is similar to that of some of the aquatic species. N. electrina appears to have a wide tolerance for moisture conditions with a definite preference for the wet habitats. Very few were collected in the driest southwestern parts of the study area.

The habitat diversity indices varied from medium to high (Table 10). These readings are probably close to reality, because I did not find many specimens in places other than soil and liter samples. N. electrina is most frequently collected in the marl meadow (Plate VIII, Figure 1). Density data also substantiate this. Of the 971 specimens collected in the quantitative samples, 498 were collected in the marl meadow and 234 were collected in the bog meadow. The elm-maple swamp forest which showed a high frequency had a total of 45 specimes. Seventy-five percent of the specimens collected were from either the marl meadow or the bog meadow. 1 9 2

MAP X L IX

Collection locations for Nesovitrea electrina (Gould) at Cedar Bog.

Dec. 23 & 25, Jan. 5, 1969 C ollected A live u \ o C ollected D ead

A p r il 23 & 2 5 , 1969 A C ollected Alive A C ollected D ead

A u g u s t 20 & 3 0 , 1969 C ollected Alive Q C ollected D ead

Miscellaneous Collections Q C ollected Alive © C ollected D ead

W . n J h i m

l’ Nesovitrea electrina. Hawaiia minuscula. w Vegetation Zones Vegetation E G H G F E 0 Vegetation Vegetation Zones i-.V PLATE VIII fA>, 3 LL. - 3 C 4) O* 5* 100 60 60 - 10 20 30 70 40 50 0 5 Go 0

- - - iue2 CollectionfrequenciesFigure 2. for im Figure Collectionfor frequencies 4. A B C D E F G H J K ■■■■ •if'.’.*- ftrWf! Glyphyalinia indentsta. Glyphyalinia f’F. Euconulus chersinus. m m ■«ai m ft!??,- :Uifc Vegetation Zones Vegetation Vegetation Zones Vegetation SlK'.B i'o!'* E F

Glyphyallnia Indentata (Say, 1823)

Distribution

Southern Canada and most states east of the 100th median and also from Utah, Arizona, Nevada, Baja California, Mexico and Central Guatemala (La Rocque, 1970: 620). This species is recorded throughout Ohio.

General Habitat

Glyphyallnia indentata may have some particular habitat prefer­ ences in various parts of its wide range. However, these preferences cannot be summarized from the literature. After examining 18 publica­ tions which mention ecological notes about indentata, in 8 states and Ontario Canada and by 14 different authors, I can find no habitat patterns.

Burch (1956: 61) found this species in 18 of 27 different plant associations in Virginia. After examining the literature this number could easily be extended to over 30 plant associations in eastern United States alone. It has been recorded from alder-dogwood thickets in Michigan (H. B. Baker, 1922: 29), jack pine in Michigan (van der Schalie, 1939: 368), former prairie and oak-hlckory-elm-maple woods in Illinois (F. C. Baker, 1939: 71) and floodplains and upland woods in Kansas (Leonard, 1959: 113-114).

It has been recorded from floodplains, ravines, open slopes, wooded slopes, meadows, marshes, old fields, and deep woods dominated by either deciduous or coniferous trees.

H. B. Baker (1922: 24) found G. indentata on quartzite cliffs, Teskey (1955: 70) found It in loose shale, van der Schalie (1939: 368) found It on limestone and Grimm (1959: 124) found It under sandstone.

Glyphyallnia indentata can apparently withstand wide variances in moisture and cover. Most authors mentioned that this species is found in damp or moist areas, but H. B. Baker (1911: 155) also collected it in dry woods dominated by mixed deciduous and coniferous trees. The types of cover In which G. Indentata can be found seems to be endless. It is frequently under rocks, and in living grasses and ferns besides, the usual kinds of plant debris.

Burch (1955: 6 6 ) analyzed the soil he found G. indentata on and found it had a pH range of 5.3 to 7.7 in Virginia, but was most common in the 6.3 to 6.7 range. It was also most common where the calcium, magnesium and potassium was high to very high, the phosphorous varied from low to high and the organic matter content was very high. 195

Gugler (1963: 199) studied some aspects of the life history of G. indentata and suggests that this animal has an annual cycle in which the adults die after laying the eggs. Blake (1931: 516-517) detected a biannual vertical migration of this species between the soil and the leaf litteri

Occurrence at Cedar Bog

Glyphyallnia indentata is the most common mollusc within the study area. One look at the distribution map (Map L) shows that there is hardly an area in the Bog where shells were not found. Dead shells are common in all vegetation zones, but there is a greater collection frequency in the mesic zones and a slightly lower frequency in the wettest and the driest portions of the study area (Plate VIII, Figure 2). All habitat diversity Indices are high to very high. G. indentata had the highest habitat diversity index for both the total number and the living number collected in the study area..

Glyphyallnia indentata can be found in soil, litter, under bark, rotten wood and under logs. A total of 4165 specimens were collected in 162 samples. The average density for the total number varied from 6.4/ ft^ in the beech-maple forest to 76.9/ ft^ in the shrub zones. The densities for the living number collected for all zones varied from 1/ ft^ in the arbor vitae and oak-maple forests to 5.3/ ft^ in the shrub zones.

My data substantiates previous workers, this snail has few habitat restrictions in eastern North America.

Hawaiia minuscula (Binney, 1840)

Distribution

Its natural range is probably most of the North America (Bequaert and Miller, 1973: 76). This species has been introduced into Bermuda, Bahamas, Cuba, Lesser Antilles, Greater Antilles, Mexico, Costa Pica, Ecuador, Peru, Hawaii, the southern maritime Province of Siberia, Great Britain, Ireland, the Netherlands and, Switzerland (Bequaert and Miller, 1973: 76-66). Hawaiia minuscula is distributed over the state of Ohio (La Rocque, 1970: 640).

General Habitat

The habitat information summarized below is taken from workers who were concerned with Hawaiia minuscula within its natural range. This species is frequently associated with wet or moist habitats. H. B. Baker (1911: 156) in Michigan, Goodrich (1932: 33) in Michigan, Oughton (1948: 95) in Ontario, and Leonard (1959: 120) in Kansas, 196

MAP L

Collection locations for Glyphyallnia indentata (Say) at Cedar Bog.

D e c . 23 L 2 5 , 1968 Jan. 5, 1969 • C ollected A live © C ollected D ead

April 23 & 25, 1969 A C o lle c te d A liv e a C ollected D ead

August 20 St 30, 1969 ■ C ollected Alive ° C ollected D ead

Miscellaneous Collections G C ollected Alive © C ollected D ead

WctufhiiTn tncd

lcnv anOfl 197 mention river and and stream floodplains as one of the common H. minuscula habitats. H. B. Baker (1911: 156) also found this species in debris along sandy beaches of a lake. In Ohio, Dexter (1950: 23) found H. minuscula in a tamarack community, an alder community and Dogwood-buttonwood community around a bog lake. Dexter mentioned that some of the communities were occasionally flooded.

Hawaii minuscula has also been recorded from numerous mesic forest associations in various parts of North America. In Illinois, F. C. Baker (1939: 72) states that "its most common habitat is in woodlands of oak, hickory and sycamore." Burch (1956: 61) found H. minuscula In ten of the twenty-seven vegetation associations examined In Virginia and it was most often collected in the oak-maple and the oak-sycamore associations. Smith (1928: 490) named this species as an important "subinfluent" In a red oak-maple climax forest in Illinois.

Although H. minuscula is found throughout much of the west, very little habitat data seems to be available. Karlin (1961: 61) having examined numerous forest associations in Montana, Colorado and New Mexico, found it once In an engelmann spruce-lodgepole pine-aspen association.

Grimm (1959: 124) collected H. minuscula frequently in Maryland along railroad tracks under various sorts of debris, around an old burned house foundation and under wet sandstone In a field. Goodrich and van der Schalie (1944: 271), in Indiana, Michelson (1953: 51) in Florida, and Archer (1939: 19) in Michigan, also collected this species in fields.

Archer (1939: 19) summarized the H. minuscula habitat in lower Michigan, "In leaf mold of the aspen woods and willow-red osier- dogwood thickets; In leaf mold pockets of oak-hickory woods; In grass, on till, in rock piles of open fields; and in wild grape thickets along the roadsides."

Burch (1955: 6 6 ) found II. minuscula in soils with a pH range of 5.8 to 7.7, however, 50% were collected In the pH range of 6.3 to 6.7. The species was also most frequently found where the soil was very high in calcium, magnesium, potassium and organic matter.

Hawaiia minuscula is conspicuously absent from most coniferous communities and from xeric deciduous communities.

Occurrence at Cedar Bog

Hawaiia minuscula was collected alive In all the vegetation zones In the study area. This species is not very common in the wet meadows and springs around the East Branch of Cedar Run. The distribution map (Map LI) definitely shows a high concentration of collection sites in the western half of the study area. The collection frequency data 198

MAP L I

(Collection locations for Hawaiia minuscula (Binney) at Cedar Bog.

D e c . 23 & 2 5 , 1968 Jan. 5, 1969 C ollected A live ° C ollected D ead

A p r il 23 & 2 5 , 1969 C ollected Alive a Collected Dead

August 20 & 30, 1969 C ollected Alive ° C ollected D ead

Miscellaneous C ollections ©Collected Alive 0 C ollected D cad

Woarlhiil.DL Bocrf

L_* 199

(Plate VIII, Figure 3) show a high incidence of occurrence in the shrub zones, silver-red maple and elm-maple swamp forests. These three zones have a loose drier muck than other zones and generally fewer fallen logs than the denser swamp forests and hardwood forests. Its high frequency in the very wet tuliptree swamp forest cannot be ignored.

One very striking thing about the frequency data is the high frequency of living specimens in many of the zones. The habitat diversity indices taken for this species are all high (Table 10) with an average of 2.7. This is second only to Glyphyallnia indentata. H. minuscula has a wide habitat range leaning toward dry substrate conditions.

Density data show the highest average concentrations of living H. minuscula in the shrub zone with 18.5/ ft^ and 15.8/ ft^ in the elm-maple swamp forest. The total density for the shrub zone averaged 45.6/ ft^, the elm-maple swamp forest averaged 80.1/ ft^ and the bog meadow average 59.8/ ft^. This last statistic is very surprising for a species usually associated with much drier conditions. A close look at this bog meadow data shows that H. minuscula was found in a few collections in very high numbers, mostly dead. The distribution map shows that most of these collections with very high number of dead shells were very close to the elm-maple swamp forest. It is possible that conditions have changed along this interface between these two vegetation zones.

Euconulus chersinus (Say, 1821)

Pistribution

Southeastern Canada and throughout the United States east of the Mississippi River and west of the Mississippi River In Missouri, Kansas, Oklahoma, Louisiana and eastern Texas (La Rocque, 1970: 611). It is generally distributed throughout Ohio.

General Habitat

Euconulus chersinus is apparently a very adaptable species. It has been associated with both coniferous and deciduous forests. Much, of the information available refers to a subspecies IS. chersinus polygratus. So far, I have not found enough Information to be able to separate the habitat of E. chersinus chersinus and E. chersinus polygratus. * The following habitat information applies to both sub­ species.

Archer (1939: 19) states that in Michigan, this species is found "in tall grass in marshes; in ferns in meadow; in leaf mold and under MAP LH

Collection locations for Euconulus chersinus (Say) at Cedar Bog.

w T t

Dec. 23 25, 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

April 23 & 25, 1969 A C ollected Alive o C ollected D ead

August 20 & 30, 1969 • C ollected Alive ° C ollected D ead

Miscellaneous C ollections Q C ollected Alive 0 C ollected D ead

Wnmthurn- 201 rotten logs in oak-hickory woods, aspen woods, and willow-red osier- dogwood thickets; in grass and plant trash on open slopes; fields and orchards; under rock slabs, cement, and boards on site of an old house. This snail is rather characteristic on the grassy slopes of old gullies;" In Kansas, Leonard (1959: 111) found that "this species is to be found in and about decaying wood, under stocks, logs, the loosened bark of fallen trees, beneath stones and in leaf litter in the forest floor."

Archer (1936: 10) in Michigan, F. C. Baker (1939: 76) in Illinois, Oughton (1948: 94) in Ontario, and Burch (1956: 61) in Virginia, found it mainly in hardwood communities. Whereas h. B. Baker (1922: 28) in northern Michigan and Archer (1942: 97) in Alabama, found IS. chersinus in arbor vitae and pine communities.

Occurrence at Cedar Bog

Euconulus chersinus occurs in too few collections to draw any possible distribution conclusions. It has always turned up in the litter and soil samples. All the collection sites for this species occur in the Carlisle muck and they are all found in or near areas which were unforested in 1938. With such few collections this Informa­ tion is possibly coincidence. Except to show a general relationship with other species, the habitat diversity Indices are not too informa­ tive. The indices range from zero to 1.2 which indicates a low habitat diversity. This is expected for a relatively rare species.

Euconulus fulvus (Mllller, 1774)

Distribution

"Euconulus fulvus (Mllller) is holarctic in distribution, but is wanting in the Gulf and southern Atlantic states from North Carolina to Texas" (Leonard, 1950: 37-38).

General Habitat

Euconulus fulvus is recorded by numerous researchers from many different plant communities. In Michigan, van der Schalie (1939: 368) found it in a jack pine forest on limestone, Harry (1951: 32) found it in several arbor vitae bogs, and H. B. Baker (1922) found it in numerous hardwood and mixed hardwood-conifer communities. F. C. Baker (1935) recorded E. fulvus from several coniferous communities dominated by pine, spruce, arbor vitae, fir and hemlock. It was particularly abundant in a birch-arbor vitae-tamarack forest. Oughton (1948: 94) even found it occasionally in Sphagnum bogs. Karlin (1961: 161) in his work in Colorado, Montana and New Mexico, associated E. fulvus with various coniferous and deciduous communities. 202

There does not appear -to be any specific moisture requirements. Goodrich (1932: 31), Oughton 1948: 94), Leonard (1950: 37-38) and La Rocque (1970: 608) all recorded this species from damp or wet habitats. However, in Russia, Likhachev and Rammel'meier (1952: 287- 288) found it in both damp as well as dry places.

Euconulus fulvus does require good cover, most frequently in the form of "downed" organic matter, such as started bark, rotting logs, leaves, decomposing humus and boards. Muchmore (1959: 8 6 -8 8 ) fre­ quently found it under stones in various deciduous woods in New York.

Occurrence at Cedar Bog

Shells of E. fulvus were found most often in the elm-maple swamp forest, bog meadow and shrub zones (Map LIII). Except for several sites in the bog meadow near Woodburn Road, this species is concen­ trated at the north end of the study area. Moisture preferences are on the wet side. The range of habitat diversity indices is from 1.72 to 2.12 (Table 10). This shows a moderately wide habitat range. Densities are all low.

Zonitoides arboreus (Say, 1817)

Distribution

North America; it is found in every state in the United States and every providence in Canada. It has been recorded from Mexico, Costa Rica, Guatamela and the West Indies (La Rocque, 1970: 653-654). It is found throughout Ohio.

General Habitat

Pilsbry (1946: 482) states, "In the eastern states and Mississippi valley this snail is everywhere abundant, to‘ be found wherever there are trees or shelter of any kind; on or under the bark of logs, under boards, bricks, stones in the grass, or in any like situation offering protection from the sun and a reasonable degree of moisture." Un- doubtably Pilsbry was summarizing the numerous records of the various investigators of his time. More recently, La Rocque (1970: 652) states that "this species is able to occupy almost any kind of environ­ ment; it is equally at hone in dense woods and open plains, in culti­ vated fields and gardens and city yards, even where there is a minimum of cover and moisture." After reviewing 34 papers which specifically refer to habitat preference for Zonitoides arboreus, I would agree with some qualifications.

Since Zonitoides arboreus is found in many diverse habitats, I am not going tp summarize where it Is found, but rather state where, in 203

MAP L IU

Collection locations for Euconulus fulvuB (Mllller) at Cedar. Bog.

v j w I

Dec. 23 & 25, 1968 Jan, 5, 1969 • Collected Alive ° C ollected D ead

A p r i l 23 & 2 5 , 1969 * C ollected Alive O C ollected D ead

August 20 Sc 30, 1969 ■ C ollected Alive g C ollected D ead

Miscellaneous Collections © C ollected Alive © C ollected D ead

Womfimm_Bocri

to

■ ■ 0 230

1. *330t) 13 Frequency {%) Frequency ($) ICO 100 GO 70 CO 30 40 20 50 CO 90 10 0 Figure 1. Collection frequencies for Figure 2. Collection frequencies for frequencies Collection 2. Figure for frequencies Collection 1. Figure Figure 3* Collection frequencies for for frequencies Collection 3* Figure M m L Z 'I 1 *1 Euconulus fulvus. Zonitoides arboreus. Zonitoides fulvus. Euconulus Guppya sterkii. Guppya

Vi-’ Vegetation Vegetation Vegetation Zones Vegetation E 0 M m ivyx 1 $ -I® l t V'W '•vf-y F Zenos enpi. e r Vegetation Zones Vegetation .74 It . L Z M K J PLATE IX PLATE S yi u. u <0 S* o u rr. ;j Ul o’ i) * 100 100 40 50 80 80 70 XO 60 GO 20 40 - 40 XO - XO 30 50 Co 70 .90 0 •j

- -

. • Figure 4. Collection frequencies for for frequencies Collection 4. Figure A 1 it * ■’-VI 3 ,s'4fi c Striatura milium. Striatura Vegetation Zenos Vegetation 0 H G F E •.■rjS siLv1;- s.-Ji 204 205 my experience and that of o.thers it is found in some environmental extremes and where it is conspicuous by its absence.

Zonitoides arboreus is not characteristic of wet or swampy places (Lee, 1952: 59). Ingram (1946: 90) specifically mentions that this species was not found on floodplains where there was a dearth.of cover. H. B. Baker (1922: 26) did not find it common in the swamps he investigated in northern Michigan. This species is found in swamps In some parts of its range and drier habitats in other parts of its range.

Karlin (1961: 61) Investigated 15 different vegetation associ­ ations in Montana, Colorado and New Mexico at 72 different stations. He found Z . arboreus common in six of these vegetation zones which included deciduous or mixed deciduous and coniferous trees. He found but one individual in a purely coniferous vegetation zone. In the eastern part of its range, Z. arboreus has been recorded numerous times from purely coniferous plant associations. Walker and Ruthven (1906: 93 and 96) found it under the bark of decaying tamarack, arbor vitae and pine: F. C. Baker (1935: 257-279) found it associated with many coniferous forests including arbor vitae; van der Schalie (1939: 368) found it in a jack pine forest; and Archer (1942: 96-97) found it In old field pine community composed of short-leaf pine.

Rapp and Rapp (1946: 124-125) surveyed the mollusca of the Great Swamp of New Jersey. They found only three species of molluscs. One was Zonitoides arboreus. They recorded some of the substrates as having a pH as low as 4.

Hubricht (1964: 34) collected Z_. arboreus living in ten different caves in Kentucky, Tennessee and Alabama. Lindeborg (1949: 130) and Oughton (1948: 94) found this snail in Sphagnum bogs in Ontario, Canada.

Zonitoides arboreus is not a desert species nor a prairie species. It is principally a deciduous forest Species. It can adjust to most any kind of cover. In Ohio, it is usually found under loose bark, rotting logs or thick leaf litter.

Occurrence at Cedar Bog

The occurrence of Zonitoides arboreus at Cedar Bog brought to light a very important point about collecting. In my early random collections I could always get Z . arboreus by lifting the loose bark on ■ dead elm trees wherever they occurred. Later on, I concentrated on quantitative soil and litter sampling. After analyzing these samples I found arboreus only infrequently, in small numbers and usually dead. I realized then that I was consistently missing a part of the molluscan fauna and resumed some random collecting around logs and other debris. At Cedar Bog, Zonitoides arboreus is usually not a soil 206

MAP L IV

Collection locations for Zonitoides arboreus (Say) at Cedar.Bog.

* I T u | v 1 w 'j x | if

D e c . 23 & 2 5 , 1968 Jan. 5, 1969 • C ollected A live 0 C ollected D ead

A p r i l 23 & 2 5 , 1969 * C ollected Alive a C ollected D ead

A u g u s t 20 & 3 0 , 1969 ■ Collected Alive ° C ollected D ead

Miscellaneous C ollections Q C ollected Alive © C ollected D cad

W e od ^m u i— a d

iu 10 ti

Q 370 440

17 1«m. -310H 11

I V 207 and litter Inhabitant. It is a climbing species and its usual habitat is under loose bark on fallen elm trees.

The distribution map (Map LIV) and the collection frequencies (Plate IX, Figure 2) show that it is not found in the bog or marl • meadows at all. Habitat diversity indices are meaningless because the quantitative data used to figure these indices are based solely on soil and litter samples.

Guppya sterkil (Dali, 1888)

Distribution

Southern Ontario and New York south to Florida in the east, and west to Ohio, Kentucky, Arkansas and Louisiana (La Rocque, 1970: 614). There are records from five counties in eastern Ohio (Taft, 1961: 6 8 ).

General Habitat

Very little information is available concerning Guppya sterkli. Oughton (1948: 94) found it in deciduous woods on limestone bedrock. La Rocque (1970: 612) found it "on grassy slopes with moss and small beeches."

Occurrence at Cedar Bog

Guppya sterkii inhabits those vegetation zones which occupy the intermediate moisture conditions. The distribution map (Map LV) indicates few collection sites In the drier southwest part of the study area and in the bog and marl meadows. Collection frequency data (Plate IX, Figure 3) shows the same distribution pattern. Guppyasterkii is most frequently collected In the arbor vitae associ­ ation with a high incidence of living specimens.

Guppya sterkii is strictly a soil and litter species at Cedar Bog. I have noticed that the highest densities occur in those forested areas which have a fairly thick moist leaf litter. The habitat divers­ ity index is high and with very little variation between the total number and the living number using either vegetation Interpretation (Table 10).

An analyses of where the consistently high densities occurred is of some value for this species. I found 1242 specimens of (3. sterkii in the study area. Five hundred and fifty-five of these were from miscellaneous litter and soil samples. Most of these miscellaneous litter and soil samples came from the arbor vitae association and a small segment of the tuliptree swamp forest (X, &-7,8). These two 208

MAP L V

Collection locations for Guppya sterkii (Dali) at Cedar Bog.

I I / | ' w ‘ I x | r~

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

April 23 & 25, 1969 A C ollected Alive a C ollected D ead

A u g u s t 20 & 3 0 , 1969 • C ollected Alive o C ollected D ead

Miscellaneous Collections © C ollected Alive © C ollected D ead

Wcac£faaiO—Eosd.

10

0 330 440

Icm. .330*1. 13 209 areas accounted for 539 of the 555 specimens. I checked all the tullptree collections which’ had substantial numbers of G. sterkli and all these were from the drier tullptree stands which had a good accumulation of leaf litter. Most of the arbor vitae stands have a thick carpet of litter because it does not decompose readily. Many of the arbor vitae samples had large numbers of (3. sterkii. In fact, Guppyasterkii is characteristic of the arbor vitae association at Cedar Bog.

It is my opinion that G. sterkii has to have a constant, loose, and moist substrate. The arbor vitae is one of the few plant com­ munities which can consistently provide this type of habitat at Cedar Bog. Most of the tullptree swamp forest is very wet at one time or another and occasionally flooded, but in those few narrow bands which are on higher ground, the leaf litter does not get compacted and saturated with water. Guppya sterkii also lives quite well In the loose moist tullptree litter in these special areas.

Most of the wet swamp forest areas do not have an accumulated leaf litter. It gets compacted by standing water or thoroughly wetted and decomposes quickly. The open elm-maple swamp forest and the shrub zones do not have a good leaf litter because of the low density of the trees and the beech-maple forest Is probably too dry.

The total average density of Guppya sterkii in the arbor vitae is 31.1/ ft^ and for the drier tullptree areas It is 34.8/ ft^. The average living density if 5.6/ ft^ for the arbor vitae association and 21.4/ ft^ for the drier tullptree areas.

Striatura milium (Morse, 1859)

Distribution

Northeastern Canada west to the eastern edge of Manitoba, south to North Dakota in the west and south-to North Carolina and Alabama in the east (La Rocque, 1970: 661). It is recorded from six widely scattered counties in Ohio (Taft, 1961: 82; La Rocque, 1970: 661).

General Habitat

Striatura milium has been recorded from wet and dry habitats in both deciduous and coniferous vegetation associations. In North Carolina, Jacot (1935: 604-605) found S^. milium In all the plant com­ munities Investigated, old field pine, white pine, pine-oak, yellow pine, yellow plne-dogwood, yellow pine-oak, scarlet oak- black oak and cove hardwoods. Three of these communities were burned and living specimens were still found in the samples collected. Most of the Striatura milium were found in the cove hardwood and the yellow pine- oak communities. 210

In the upper peninsula-and Isle Royale of Michigan, Walker and Ruthven (1906: 93 and 96) found S_. milium in communities which con­ tained arbor vitae, hemlock, spruce and birch trees in bark, moss and humus. Birch (1956: 61) recorded this snail, in Virginia, from seven different deciduous associations, most were dominated by oak. Many other investigators found it in deciduous and coniferous forests, especially in Michigan.

There does not appear to be any definite pattern in any part of this species1 range. Most records Indicate that the species was collected In moss, humus, under bark or leaf litter. Most of the communities mentioned were mesic to dry in moisture conditions.

Occurrence at Cedar Bog

Striatura milium is a common soil and litter species in the moist to dry parts of the study area (Map LVI). It has not been found living in the marl meadow. Its collection frequency was lowest in the four wettest vegetation zones in the Bog (Plate IX, Figure 4). In the shrub zones, 94% of the samples contained dead shells of jS. milium. High collection frequencies also occurred in the elm-maple swamp forest and the arbor vitae association. The habitat diversity indices are high (Table 10).

The average total density for j3. milium in the elm-maple zone was 8 6 / ft^, in the shrub zones 33.9/ ftzt in the bog meadow 45.8/ f2 and the arbor vitae 18/ ft^. The average living density for S. milium In the elm-maple was 5.2/ ft2, in the bog meadow 32.3/ ft^", in the shrub 19.5/ ft^ and in the arbor vitae association 2.3/ ft^. As you can see there are several inconsistencies between density, dis­ tribution and frequency data. The major inconsistency is In the bog meadow. S_. milium was not found there often, but when It did occur, it occurred in large numbers. In other words, there is a high degree of clumping of this species in the bog meadow. Possibly isolated areas that were not too wet all the time. ‘S. milium had a high collection frequency and a low living density In the beech-maple forest and the shrub zone of the southwestempart of the study area. It appears as if the conditions changed there and jS. milium is no longer common there.

Striatura exigua (Stimpson, 1850)

Distribution

Northeastern Canada west to Manitoba, south to Virginia in the east and northwest from Virginia on an angle to Minnesota (La Rocque, 1970: 658). In Ohio, this species has been recorded from five counties in the south and east (Taft, 1961:80). MAP L V I

Collection locations for Striatura milium (Morse) at Cedar Bog.

s I ! _“_TJ w

D e c . 23 & 2 5 , 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

A p r i l 23 & 2 5 , 1969 A C ollected Alive A C o lle c te d D ead

August 20 & 30, 1969 » C o lle c te d A liv e d C ollected D ead

Miscellaneous Collections Q C ollected Alive © C ollected D ead

W fln i lh n H I

r :

0 220 440 twUOli 2X2

General Habitat

The only habitat notes available apply to Ontario and Michigan. In Ontario, Canada, Oughton (1948: 28 and 94) considered this species abundant and generally from damp deciduous woods and occasionally from Sphagnum bogs. La Rocque (1970: 657) collected this species In moss around stumps and logs in shaded woods.

In the lower peninsula of Michigan, Harry (1951: 32) collected It from several northern white cedar (arbor vitae) swamps and deciduous woods, and Archer (1939: 19) took it "in leaf mold In log rafts in the birch-maple swamp woods." In the northern part of the upper peninsula of Michigan, Walker and Ruthven (1906: 93) found S^. exigua in the leaf litter from hemlock and maple woods, and in decaying logs of arbor vitae and tamarack In a tamarack swamp. On Isle Royale, Walker and Ruthven (1906: 96) found it In moss in several coniferous communi­ ties including arbor vitae and tamarack swamps and in the leaves of an alder thicket.

This species is found in both deciduous and coniferous forests under logs, leaf litter and moss. Most of the forest communities mentioned were either along waterways, swamps or bogs. This snail seems to be most common in wet habitats.

Occurrence at Cedar Bog

Based on the general ecology notes, I would have expected to find this species common in the arbor vitae association, but I did not. Except for a few scattered collections north of Woodburn Road, Striatura exigua is confined to a small area south of the road (Map LVII). It was not found in any wet vegetation zones, dead or alive. Only 37 specimens were collected and 24 of those specimens were taken In two samples from the arbor vitae association. I cannot offer any reason why S. exigua is confined to such a small area at the Bog.

Haplotrema concavum (Say, 1821)

Distribution

Southeastern Canada to Florida and west to Minnesota in the north and Texas in the south (La Rocque, 1970: 607). La Rocque also recorded in many localities throughout Ohio.

General Habitat

"This species has long been known to be a carnivore, feeding on other snails. This fact restricts its distribution to areas where snails are abundant, specifically lime-rich areas; it may therefore be 2 1 3

MAP LV H

Collection locations for Striatura exigua (Stimpson) at Cedar Bog.

s | t j u | v j w 1 x

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

A p r i l 23 & 2 5 , 1969 A C ollected Alive O C ollected D ead

August 20 & 30, 1969 ■ Collected Alive D C ollected D ead

Miscellaneous C ollections © C ollected Alive © C ollected D ead

Wotirihiirn tncit

10

Itm, H 2 3 0II ^ ~i t ji ; v r w 214 PLATE X

100 100

80 80

c 3o CT II CP t- U-

10 - 10 -

Vegetation Zones Vegetation Zones

Figure 1. Collection frequencies for Figure 2. Collection frequencies for Striatura exigua. Haplotrcma concavum.

100 • 100

80

cS* 3o * . tr cr, CD t. Lu 40

20

10 - mt&'j. m

Vegetation Zones Vegetation Zones

Figure 3* Collection frequencies for Figure 4. Collection frequencies for Stenotrema barbatum. Stonotrema lcai. 215 described as an indirect calciphile" (La Rocque, 1970: 605). Burch (1954: 32) found H. concavum "extremely common, being found wherever the habitat is favorable for other snails." So far H. concavum has not been associated with any particular plant community or vegetation association. Most researchers have found it associated with various hardwood communities.

In Indiana, Goodrich and van der Schalle (1944: 269) found it "at the edges of woods and shaded margins of marshes, on limestone bluffs and occasionally in grass heaps in town gardens." In Michigan, Archer (1936: 8 ) found it associated with ferns in bog woods.

It is found in many kinds of cover. F. C. Baker (1939: 92) found it under forest debris, old logs, leaves or most any object which could hide and shelter it. He also thought this species to be rather solitary. That is generally the case with predators.

Occurrence at Cedar Bog

Haplotrema concavum is a rather uncommon snail at Cedar Bog. It does not show any particular distribution pattern (Map LVIII). H. concavum appeared in 18 samples with a total of 29 specimens collected. Only one living specimen was found. The habitat diversity index is between 2.28 and 2.38 which is moderately high and indicates a rather even and wide habitat preference. Most of the specimens collected were from soil and litter samples. I did not find this snail in the logs and under bark as I have many times in other localities. Also, H. concavum did not appear in the wettest or the driest areas.

Stenotrema barbatum (Clapp, 1904)

Distribution

"This species ranges from Massachusetts to southern Minnesota (including the southern tip of Ontario), south in the west through Iowa to Kansas, in the east to Mississippi and Alabama, north on the Coastal Plain and Piedmont through the Carollnas to southern New England" (Grimm, 1971: 17).

General Remarks

Initially I called this species Stenotrema hirsutum (Say, 1817). After reviewing much of the ecological data concerning hirsutum I realized that other investigators such as Grimm (1971) and Hubricht (1973) and Archer (1948) were right. There are definitely two species lumped together. Pilsbry (1940: 664) states, "It has long been recognized by collectors that S^. hirsutum is represented In many localities by a small form of upland or dryer woodland, and a larger MAP L V ffl

Collection locations for Haplotrcma concavum (Say) at Cedar Bog.

w T I

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

A p r i l 23 & 2 5 , 1969 A C ollected Alive A C o lle c te d D ead

A u g u s t 20 & 3 0 , 1969 ■ C ollected Alive ° C ollected D ead

Miscellaneous Collections © Collected Alive © C ollected D ead

W o odhtufk-JEa e d 217 lowland form." Pllsbry states that he found intermediate forms, but that further anatomical study was in order. Archer (1948: 30-32) studied the genus Stenotrema and found the shell and the genitalia distinct. Grimm (1971: 16-17) again reviewed the status of jS. barbatum and j>_. hirsutum. He also concluded that ji. barbatum was a distinct species.

X sent my specimens to Leslie Hubricht and he identified them as Stenotrema barbatum. He also sent to me some specimens of S^. hirsutum for comparison. I personally believe that S^. barbatum and _S. hirsutum are distinct species.

General Habitat

I could review all the ecological data concerning S_. hirsutum which would include most of what is known about S^. barbatum. At this time, I would have no way of separating what habitat data belonged to S^. hirsutum and what belonged to S^. barbatum. I do know that the data I reviewed fell into two categories, the wet habitats and the dry habitats.

Pllsbry (1940: 666) reports that the types of S^. hirsutum barbatum from Wetumpki, Alabama were found on the floodplain of the Tallapoosa River. Archer (1948: 31-32) states that this species is usually restricted to low mesic or hydric woods in stream valleys in the southern part of its range. However, in the north, J3. barbatum is found on bluffs and hills bordering rivers as well as in marshes. He states further that it is found in hardwood cover, under logs, under rocks and in leaf litter, and in the roots of grasses and ground plants of open fields, roadsides, and suburban gardens. Grimm (1971: 17) states that "throughout most of its range Stenotrema barbatum is a hygrophile, preferring shaded floodplains and marshes. It has been called the 'lowland form' of J3. hirsutum. It is most abundant where the calcium content of the soil is high, but it is found in acid soils as well. In scattered upland areas, usually near floodplains, it occupies ravines, clearings and fields."

Occurrence at Cedar Bog

The distribution of S^. barbatum correlates loosely with the drier vegetation zones (Map LIX). At Cedar Bog, £>. barbatum is not particu­ larly a hygrophile. However, I did find dead specimens in all the vegetation zones in the study-area. I found it in the soil and litter as well as under logs and bark. I usually found living j>. barbatum under logs or loose bark, and not in the soil and litter. This appar­ ent when the proportion of living specimens collected by random methods is compared with the proportion of living specimens collected by soil and litter samples. Most living specimens are collected by random collecting. MAP L IX

Collection locations for Stenotrema barbatum (Clapp) at Cedar Bog.

w

Dec. 23 Sc 25, 1968 Jan. 5, 1969 • Collected Alive ° C ollected D ead

April 23 8c 25, 1969 A C ollected Alive & C o lle c te d D ead

August 20 & 30, 1969 ■ Collected Alive ° C ollected D ead

Miscellaneous Collection 0 C ollected Alive © C ollected D ead

W ouiliuutjbicil

0 320 440 1cm .110h ---- (---- _ ---- r ---7---- r ------■ 219

Frequency data (Plate X, Figure 3) shows a higher incidence of dead shells in the drier vegetation zones. The frequency data for living specimens collected does not show any patterns. The habitat diversity indices are high, but since they are are based solely on the soil and litter collections, they are most likely incorrect. Densi­ ties are very low in most collection for both dead and living speci­ mens.

My data show that S^. barbatum can live in many types of habitats over a fairly wide range of moisture conditions. I found it living in some wet areas of the raaple-ash-basswood and tuliptree swamp forest segregants and in some fairly dry beech-maple and silver-red maple forests. It was rare or absent in most of the wettest parts of the study area.

Stenotrema leal (Binney, 1878)

Distribution

New Brunswick and southern Ontario west to Minnesota and South Dakota, south to Texas in the west, and south to North Carolina in the east (La Rocque, 1970: 569). It has not yet been recorded from South Carolina, Georgia and Florida (La Rocque, 1970: 569). In Ohio, this species has many records, but most of them occur in the south­ western quarter of the state.

General Habitat

Stenotrema leai is found predominately in moist to wet habitats In and around river floodplains, swamps, marshes, lakes and ponds (H. B. Baker, 1911; 154, F. C. Baker, 1939: 60, Pilsbry, 1940: 678, Marsh, 1941: 114, Oughton, 1948: 95, and La Rocque, 1970: 568). H. B. Baker (1911: 154) also found It in some higher woods and Archer (1936: 11, 1937: 120 and 1939: 20-21) found £. leai frequently associated with grassy areas in meadows, orchards, railroad embankments and ditches.

S. leai Is not associated with any particular plant community. It has been recorded from woodlands of oak, hickory, elm and willow by F. C. Baker (1939: 60) and near arbor vitae by H. B. Baker (1911: 154).

This snail does require a certain amount of cover. In forest con­ ditions It is found under logs, boards, and stone piles (H, B. Baker, 1911: 154, Archer, 1936: 11, 1937: 120, and F. C. Baker, 1939: 60). Archer (1939: 20-21) found that this species used grass hollows for cover. 220

MAP L X

Collection locations for Stenotrema leai (Binney) at Cedar Bog.

T I w ' j x ' f

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

April 23 & 25, 1969 A C ollected Alive ^ C ollected D ead

August 20 & 30, 1969 ■ C ollected Alive ° C ollected D ead

Miscellaneous Collections © C ollected Alive © C ollected D ead

WoodkujUtoed 221

Occurrence at Cedar Bog

A comparison of distribution maps (Map LIX and Map LX) between Stenotrema leai and Stenotrema barbatum shows how much of a hygrophile S. leai really is. At Cedar Bog, S^. leai is found mainly in the bog meadow and marl meadow associations. Only one living specimen was found in a drier wooded area. Collection frequency data (Plate X, Figure 4) shows that 91% of the marl meadow collections contained shells of J3. leai and 56% of the bog meadow collections contained shells of S^. leai.

This is one of the polygyrid species in which the habitat divers­ ity indices would be accurate. This species is usually found on the substrate and in small sedge and grass hummocks. In the bog and marl meadows all areas have an equal change of being samples. There are no logs and trees to affect the sampling procedure. The habitat diversity indices are 1.16 and 1.70 for the total number collected and .61 and ,93 for the living number collected (Table 10). These low indices show a narrow habitat preference.

Mesodon clausus (Say, 1821)

Distribution

New York and Virginia west to eastern Kansas, north to southern Michigan, Wisconsin and Minnesota, south to Texas, Mississippi, Alabama and northern Florida (La Rocque, 1970: 576). Taft (1961: 16) records M. clausus widespread in Ohio.

General Habitat

Mesodon clausus, in the eastern part of its range, appears to be an inhabitant of ecotonal areas and successional communities. Good­ rich (1932: 18) and Archer (1937: 119) found this snail abundantly along roadsides and railroad embankments. Call (1900: 392), in Indiana, found it frequently associated with low areas with "abundant growths of weeds." They also associated this species with open areas. F. C. Baker (1902: 159) and (1939: 57) found M. clausus in forests, especially in river valleys, as well as railroad embankments and road­ side areas. Leonard (1959: 94) in Kansas, however, refers to this species as essentially a forest species. Perhaps in the western part of its range, it is.

Occurrence at Cedar Bog

The occurrence of M. clausus at Cedar Bog was puzzling for awhile. I infrequently collected dead specimens in the soil and litter samples. This in Itself was not unusual for a polygyrid snail except 222

MAP L X I

Goilectloti locations for Mesodon clausus (Say) at Cedar Bog.

I w i— r

Dec. 23 & 25, 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

April 23 & 25, 1969 * C ollected Alive A C ollected D ead

August 20 & 30, 1969 ■ Collected Alive ° C ollected D ead

Miscellaneous C ollections © C ollected Alive © C ollected D ead

Wo

- - -

- . - Figure 3* Collection frequencies Collectionfor 3* Figure Figure 1. Collection frequencies1. for Collection Figure

A3CUEFGH 'Mi Mesodon inflectus. Mesodon Mesodon clausus.Mesodon Vegetation Zones Vegetation Vegetation Zones Vegetation ■i:# F m H PLATE XI PLATE tn 3 t- CJ ’ C O e a* 100 80 100 40 40 - 0 5 - 0 7 50 - 10 20 0 3 6q 0 3 40 - 0 5 • 80 10 20 6 70. 0 9 0 0 o ------• ------

Figure 2. Collection frequenciesfor 2. Collection Figure Figure 4. Collection frequenciesfor Figure 4.Collection '2 V-Wii- & s A Si M /> If B ~i ..rrm. Mesodon thyroidus. Mesodon Triodopsis nultilineata.Triodopsis r??,, § ! * Vegetation Zones Vegetation Vegetation Zones Vegetation 0 VilXi 0 E F G H J J H G F E 0 £

F G H H G F i$F' I 223 m 224

I rarely found it under either logs or bark. On several occasions after most of the collections were made* I returned to the study area to collect succineids for identification purposes. Adult M. clausus were found abundantly during July and August in border areas of the beech-maple -and silver-red maple forests. These snals were three to six feet high on nettles, jewelweed, and a few other tall herbaceous plants. I also found it on cattails along with some Succinea ovalis at the edge of the bog meadow. The adults of this species are definitely climbers. The lmmatures are probably lumped with all. the other polygyrid immatures which cannot always be accurately identified. It is possible that the immatures, which were not identified, live primarily in the soil and litter.

Initially the distribution of M. clausus also had me wondering (Map LXI). I now believe that M, clausus, as an adult at Cedar Bog, is an ecotonal species as I previously mentioned. This snail was usually collected at the edge of forested areas bordering open fields and meadows. As adults this snail may have some specific food plant which usually occurs in abundance only in ecotonal areas.- This species requires more study. A life history study of this species would be interesting.

Mesodon thyroidus (Say, 1817)

Distribution

Maine, southern Ontario west to Minnesota and Nebraska, south to Texas In the west, and northern Florida in the east (La Roque, 1970: 574). It is distributed over the entire state of Ohio (Taft, 1961: 21).

General Habitat

Mesodon thyroidus is a large snail commonly collected throughout most of its range. It is recorded from numerous collections (Pilsbry, 1940: 706-709). Several aspects of this species* life history and ecology have been investigated by Foster (1936: 478-482) and (1937: 545-546), Van Cleave and Foster (1937: 50-54), Blinn (1961) and (1963: 498-505).

Not only is M. thyroidus widespread In eastern United States, it is also found in a wide variety of habitats. Like most terrestrial molluscs it has not been found associated with any one particular plant community. F. C. Baker (1910: 496-498), H. B. Baker (1911: 154), Dexter (1950: 25-26), Burch (1954: 32) and Blinn (1963: 498) all record this species from communities where various species of oak (Quercus) are among the dominants. It has been listed from a willow community by Dexter (1950: 25-26), an old field pine and sassafras community by Archer (1942: 97), and weedy and grassy fields by Archer (1937: 119). 225

Most investigators do not elaborate as to the composition of the plant communities in which M. thyroidus is found, but simply call them hardwoods or deciduous woods.

Mesodon thyroidus is frequently found on river and stream flood­ plains (H. B. Baker, 1911: 154; Foster, 1936: 987-982; F. C. Baker, 1939; 56; Ingram, 1941: 14; Hubricht, 1950: 7; and Leonard, 1959: 60). Clench (1954: 23) states, "Its distribution will probably always remain spotted owing to its own ecological demands for rather moist hardwood areas, particularly along stream margins."

H. B. Baker (1911: 154) surveyed a sand dune region along the south shore of Saginaw Bay, Michigan and found M. thyroidus in nine of the 40 different habitats looked at. Summarizing these habitats, he found this snail in swampy woods, higher woods, on the sand beach, in a dry meadow, under driftwood, in wooded flats, at the edge of meadow and a swamp. All the areas mentioned had good forms of cover or shelter.

Cover appears to be a necessary ingredient for many large ter­ restrial molluscs in eastern North America. Foster (1936: 978-982) found M. thyroidus associated with nettles (Laportea conadense), but only when there was a good ground cover nearby. Many investigators noted that Mesodon thyroidus is associated with rotten logs, loose bark, leaf mold, old stumps and old railroad ties (F. C. Baker, 1902: 158, 1910: 496, and 1939: 56; H. B. Baker, 1911: 154; Goodrich, 1932: 19; Archer, 1936: 12 and 1937: 119; Dexter, 1950: 25-26; Grimm, 1959: 123; and Leonard, 1959: 90). Grimm (1959: 123) and Ingram (1941: 14) also found it used stones as shelter.

Burch (1955: 66), in Virginia, found M. thyroidus in soils with a pH range of 5.3 to 7.2, high to very high in calcium, magnesium, and potassium, low to medium in phosphorous and very high in organic matter.

Occurrence at Cedar Bog

Mesodon thyroidus does not occur in the open bog and marl meadows, the shrub zones or the elm-maple savanna like swamp forest segregant. It occurs only in the forested areas and most often in the deciduous forests.

This snail is not usually collected in soil and litter samples. I usually found it under some form of rigid shelter or crawling up on vegetation. M. thyroidus is a climber at times and is occasionally active during the day. At Cedar Bog, the type of cover or shelter available is probably more important than the moisture conditions, although it seems to be found more often in the intermediate or mesic habitats. 226

MAP L X II

Collection locations for Mesodon thyroidus (Say) at Cedar Bog.

I II w T I

Dec. 23 & 25, 1968 Jan. 5, 1969 • Collected Alive ° C ollected D ead

A p r il 23 & 2 5 , 1969 A C ollected Alive A C o lle c te d D ead

A u g u s t 20 St 3 0 , 1969 ■ Collected Alive ° C ollected D ead

Miscellaneous Collections Q C ollected Alive @ C ollected D ead

W oedtuiUL-ftoc d 227

Considering the habitat of this snail and the sampling procedure for the data used in the diversity indices, these indices are mean­ ingless for this species.

Mesodon inflectus (Say, 1821)

Distribution

New York and southern Ontario west to southern Michigan, south­ west to southern Illinois, eastern Kansas and Oklahoma, south to Louisiana (La Rocque, 1970: 587). In the eastern part of its range It is found south to northernFlorida (Pllsbry, 1940: 772). It appears to be sporadic in the eastern coastal states and largely absent from the northern New England states. It is distributed over most of Ohio, but it has not been collected from the northeastern counties (Taft, 1961: 18; La Rocque, 1970: 586-587).

General Habitat

Mesodon inflectus was studied in some detail by Archer (1933: 5) and states the following about its habitat preferences. "In the Middle West its favorite habitat is a depression in the ground filled with fallen leaves and plant debris. As a rule such spots are not in thick woods, but in open groves, very open woods chiefly of oak, on dry hillsides, or In upland country." Lee (1952: 59) found Mesodon inflectus, in southern Michigan, inhabited dry upland areas far from the wetter situations frequented by other snails. In Ohio, Archer (1937: 120) found it abundant in orchards and fields under boards and rocks and common In roadside thickets. In southern United States, Mesodon inflectus is found "in open and cut-over country, river bluffs, and dry open, rocky places supporting a growth of oak or cedar" (Archer, 1933: 5).

This snail's preference for rocky areas has been recorded by Archer (1941: 9), F. C. Baker (1939; 47), Call (1900: 387), Teskey (1955: 70) and Leonard (1959: 94). Hubricht (1964: 33) even found this snail in caves in Kentucky and Alabama.

Occurrence at Cedar Bog

Mesodon inflectus is a rare species at Cedar Bog. Considering its habitat preferences recorded by other investigators, this is no surprise. I found 16 specimens In 9 collections from five vegetation zones. Most specimens were found In the drier parts of several swamp forest segregants. This species does not occupy the soil and litter habitat at Cedar Bog. Only one living specimen was collected in the beech-maple zone (not mapped) where I would have expected it, If at all. In fact, this was the only specimen found south of Woodburn Road. MAP LXni

Collection locations for Mesodon inflectus (Say) at Cedar Bog.

w ! r

Dec. 23 fit 25, 1968 Jan. 5, 1969 • Collected Alive o C ollected D ead

April 23 &t 25, 1969 A C ollected Alive o C ollected D ead

August 20 8t 30, 1969 • C ollected Alive D C ollected D ead

Miscellaneous Collections © C ollected Alive © C ollected D ead

Weaithnm-Hiicd

1«ik. hIIOii 229

Trlodopsis roultilineata (Say, 1821)

Distribution

Maryland, New York and southern Ontario, west to Minnesota and Kansas, south down the Mississippi Valley to Mississippi in the west, but only down to Tennessee In the east (La Rocque, 1970: 600). It is found in many widely scattered localities in Ohio (Taft, 1961: 30.

General Habitat

Trlodopsis multillneata is a gregareous species (Call, 1900: 390; Goodrich and van der Schalie, 1944: 269; Leonard, 1959: 100; and La Rocque, 1970: 598-599). It also migrates to particular areas to hibernate (Goodrich and van der Schalie, 1944: 269; Vagvolgyl, 1968: 222; and La Rocque, 1970: 598-599). This species is found most abundantly In wet habitats. Call (1900: 390), H. B. Baker (1911: 153-154), F. C. Baker (1939: 51), Pllsbry (1940: 848), Goodrich and van der Schalie (1944: 269), Leonard (1959: 100) and La Rocque (1970: 598-600) all recorded this species from the edges of marshes or swamps, floodplains, and all sorts of wet communities. This species shows a great deal of consistency with regard to habitat preferences, more so than any other polygyrid snail in Ohio.

Occurrence at Cedar Bog

Trlodopsis multillneata Is not common at Cedar Bog. It occurred almost exclusively in the bog and marl meadows. Of the 37 specimens collected, only three dead specimens were collected in other vegeta­ tion zones (Map LXIV). The preferred habitat is the bog meadow, probably because of the better cover there. The specific habitat of this species at Cedar Bog is the dense grass and sedge hummocks in the bog meadow. The habitat diversity indices are low and probably accurate (Table 10).

Allogona profunda (Say, 1821)

Distribution

Southern Ontario, New York, Maryland, and Virginia, west to southern Minnesota, eastern Nebraska and Kansas, south to northern Alabama on the east (La Rocque, 1970: 602). There is also an exten­ sion of Its southern range into Louisiana and Mississippi. It is found throughout Ohio (Taft, 1961: 14-15). MAP L X IV

Collection locations for Triodopsis multilineata (Say) at Cedar Bog.

> |s | t j u | v j w 'f x | r| i

Dec. 23 & 25, 1968 Jan. 5, 1969 • Collected Alive ° C ollected D ead

April 23 & 25, 1969 * C ollected Alive A C ollected D ead

August 20 & 30, 1969 ■ C ollected Alive D C ollected D ead

Miscellaneous C ollection Q C ollected Alive @ C ollected D ead

Wflmftiiifn Bn-rf 231

General Habitat

Allogona profunda Is predominately a forest snail. In most parts of its range it occurs in well shaded habitats. F. C. Baker (1902: 150 and 1939: 48), Goodrich (1932: 13), Shimek (1936: 71), Goodrich and van der Schalie (1944: 269) and Solem (1952: 127-129) associated A. profunda with deep woods. Shimek (1936: 71), Archer (1937: 119) and Goodrich and van der Schalie (1944: 269) also found this species on rocky bluffs. Goodrich (1932: 13) and Archer (1937: 119) report find­ ing A. profunda in areas that have dense weedy cover.

Most researchers found this species in fairly moist environments and usually associated with rocky areas or where there is an abundance of fallen timber.

Goodrich (1932: 13) considered A. profunda a disappearing species associated with the original hardwood forests. This species is probably not as common as it once was, but it is still found in many areas that provide good dense cover.

Blinn (1963: 505) recorded this species as a migrater which moves seasonally to winter hibernating quarters.

Occurrence at Cedar Bog

Allogona profunda is one of the three gastropods which never turned up in the soil and litter samples. This is a rare species at Cedar Bog. I only found five specimens. The only living specimen was collected under a fallen tree just outside the beech-maple forest in a grassy field. This snail is probably left from the time when the beech-maple "forest was much more extensive. This snail requires more cover than is currently available in most of the study area.

Pisidium compressum Prime, 1851

Distribution *"

"Found throughout most of Canada and the United States, and into Mexico" (Burch, 1972: 7).

General Habitat

"PisiddLum compressum is an inhabitant of streams, rather than ponds, but Is occasionally found in the latter. It has been taken at depths up to 20 feet or more, on mud or clay bottoms. This species flourishes in a brook fed by artesian springs In Meade County, Kansas, here it lives in fine to coarse sand in swiftly flowing water, though it may also be found creeping about on aquatic vegetation" (Leonard, MAP L X V

Collection locations for Allogona profunda (Say) at Cedar Bog.

w I x | r "

Dec. 23 & 25. 1968 Jan. 5, 1969 • C ollected A live ° C ollected D ead

April 23 & 25, 1969 * C ollected Alive 6 C ollected D ead

A u g u s t 20 & 3 0 , 1969 ■ C ollected Alive ° Collected D ead

Miscellaneous Collections 9 C ollected Alive © C ollected D ead

Waarfkufn

1

1950: 40-42). Herrington (1962: 35) states that this species prefers sandy bottoms with vegetation In shallow water. Clarke (1973: 176) found it most frequently on mud bottoms in the Canadian Interior Basin. Apparently the substrate is not important. Zimmerman (1960: 23) reports P. compressum from sandy, sandy silt or mud bottoms.

Taylor (1960: 48) states, "This species inhabits only perennial water bodies with some current action, such as lakes, rivers and creeks; it is never found in ponds, swamps, lagoons or bogs."

Occurrence at Cedar Bog

Pisidium compressum is, so far, limited to one collection from the West Branch of Cedar Run. All the dead specimens collected on land are probably the result of the dredging which occurred there some time ago.

Pisidium casertanum (Poll, 1795)

Distribution

"Nearly cosmopolitan in distribution. It has been reliably recorded from all the United States except Hawaii, Kentucky and North Dakota" (Burch, 1972: 7).

General Habitat

"The wide distribution of Pisidium casertanum reflects a cor­ responding adaptability, greater than any other American sphaerlid. It lives where any other species of Pisidium can live, except for deep water" (Taylor, 1960: 47). ' Harrington (1962: 34) found it in bog ponds, ponds, seasonal swamps, swamp creeks, swift flowing creeks, rivers, and lakes, even the Great Lakes. Clarke (1973: 174) found it predominately in mud bottoms. Zimmerman (1960: 22) reports it from water 0.5 m to 3.0 m deep in mud or sand bottoms.

Occurrence at Cedar Bog

Pisidium casertanum is a common species throughout most of the wet areas of the Bog. Its distribution follows Cedar Run and its feeding springs (Map LXVII). It Is occasionally found in the seasonal wooded pools, but it is consistently collected in the springs and streams at Cedar Bog. I am not sure why this bivalve is so infre­ quently collected alive, especially in the marl meadow where its total frequency is 63%. It was found dead in numerous areas where standing water or springs no longer occur. This all may be supportive evidence that the water sources for Cedar Bog have changed significantly in the last 100 years. Possibly due to the dredging of the West Branch of Cedar Run. 2 3 5

MAP LXVI Collection locations for Pisidium compressum Prime at Cedar Bog.

_L "I 5 T 5" I

D ec. 23 & 25, 1968 Jan. 5, 1969 • Collected Alive ° C ollected D ead

A p r il 23 & 2 5 , 1969 * C ollected Alive 6 C ollected D ead

August 20 & 30, 1969 ■ C ollected Alive a C ollected D ead

Miscellaneous Collections © C ollected Alive © C ollected D ead

WoaJhnr n tn.J

10 10

0 220 *40 ia l«m. m2201* ■3 T “t 717 o' 1" v x _ w MAP LXVII Collection locations for Pisidium casertanum (poll) at Cedar Bog.

Dec. 23 & 25, 1968 Jan. 5, 1969 o • C ollected A live ° C ollected D ead

A p r i l 23 & 2 5 , 1969 A C ollected Alive A C ollected D ead

A u g u s t 20 L 3 0 , 1969 * C ollected Alive ° C ollected D ead

Miscellaneous C ollections ©Collected Alive 0 C ollected D ead

Wa&tibu£n_KaccL.

□ O

330

R S uT X V Z DISCUSSION OF VARIOUS MOLLUSCAN FAUNAS

Frederick Vegetation Zones

Arbor Vitae Association

The xnolluscan fauna of the arbor vitae association is unexpectedly diverse. A total of 44 species was collected, of which 31 species were found living in this association (Table 11). The characteristic feature of this fauna is that the most abundant species are small. Very few specimens of the physically large species were collected in the arbor vitae association. This correlates well with the small, granular leaf litter available for cover. There are fallen trees in the arbor vitae stands, but most of these trees continue to live. There would be some space for large species around the interface between the trunk and the litter, but rotting wood and loose bark is almost non-existent. Very few species of Philomycus carolinianus, Anguispira alternata, Haplotrema concavum and the large polygyrids were collected in the arbor vitae association at Cedar Bog. The general lack of a well developed herbaceous level may also be a factor, especially as a food source for some of the large polygyrids.

The arbor vitae association is not a particularly uniform habi­ tat. It is broken up by many springs and wet depressions which provide microhabitats for several aquatic species and some hygrophilic terrestrial species. The arbor vitae stands throughout the Bog show extensive individual variation in age. A wind damage stand (V, W - 11, 12) also adds to the habitat diversity of the entire association.

The most frequently collected living species in the arbor vitate association are those which are fairly widespread throughout the Bog in the moist areas. Carychium nannodes is the only species which is almost totally restricted to this zone. Nesovitrea binneyana and Guppya sterkii are more common in this association than any other single vegetation zone or association in the study area.

Relative to the other vegetation zones at Cedar Bog the diversity index for the arbor vitae association Is high for both the total num­ ber of species collected and the living species collected (Table 12). There are no standard criteria for diversity Indices. A high divers­ ity index is relative to all the other diversity indices calculated for the various other vegetation zones in the same area.

Since most of the mollusca living In the arbor vitae are small soil and litter Inhabitants, the quantitative sampling procedure works

237 238 TABLE 10

Habitat diversity indices -for the Molluscs found at Cedar Bog.

Species Habitat Diversity Index Calculated Habitat Diversity Index Calculated by Using the Frederick (19&7) by Using the Environment Control Vegetation Zones Corporation (1973) Vegetation Zones

Immature Polygyridae 3-13 , % 2.80 Glvphvalinia indentata 3*05 (2.99) 2.62 (2.31) Hawaiia ninuscula 2.90 (2.70) 2.87 (2.31) Gastrocopta contracta 2.83 (2.24) 3.02 (2.34) Zonitoides arboreus 2.B2 (1.55) 2.64 (1.55) helicodiscus parallelus 2.81 (1.99) 2.60 ( I. 99) Colunella edentula 2.80 (2.39) 2.60 (2.47) Stenotrena barbatum 2.79 (2.06 ) 2-55 (1-83) Carychiun exile 2 . 0 (2.54) 2.54 (2.59) Fonatiopsia lapidaria 2.60 (1.94) 2.67 (1.58 ) Punctun ninutissimum 2.96 (2.24) 2.45 ( 1.85 ) Mesodon clausus 2.44 (0.00) 2.45 (0.00) Striatura milium 2.43 (2.43) 2.32 (2.08) Guppya sterkii 2.38 (2.51) 2.44 ( 2.56 ) Gyraulus parvus 2.36 (0.22) 2.43 (0.22) Fisidium casertanum 2.28 (0 .77) 2.08 (O.77) Haolotrena concavum 2.28 2.38 Carychium cxiguum 2.18 (2.38) 2.26 (2.16 ) (.’esovitrea clectrina 2.14 (2.14) 1*95 (1.54) Succinea ovalis 2.14 (0.81) 2.05 (0.81) Pnyr-a integra 2.03 (0.00) 1.81 (0 .00) Gastroccpta pentodon 2.02 (2.62 ) 2.19 (2.49) Immature Pupillidao 2.01 (1.99) 2.34 (2.02) Deroceras laeve 1.94 2.33 Lynnaea humilis 1-93 (0.65) 1.90 (0.92) Vertigo ovata 1.86 (0.00) 2.14 (0.00) Nesovitrea binneyana 1.83 (0.00) 1.46 (0.00) Vertiqo qouldi 1.82 (I.56 ) 2.59 (1-78) Vertiqo milium 1.82 (1.46) 2.69 *(1.92) Helisona anceps 1.79 1.24 Anauispira alternata 1.79 (0.00) 1.79 (0 .00) Gastroccpta iappaniana 1.77 (0-73) 2.02 ( I. 37) Immature Zonitidae 1.75 2.10 Vertiqo tridentata 1.73 (1.35) 1.63 (0.59) Euconulus fulvus 1.72 (2.03) 1.78 (2. 12) Strobilops labyrinthica 1.67 (0.58 ) 1.99 (0.90) Striatura exigua 1.66 (0.92) 1.66 (0.92) Mosodon thyroidus 1.52 (1.00) 1.52 (1.00) Cochliocopa lubrica 1.49 1.06 Amnicola limoca 1.44 1.49 Triodccsis nultilineata 1.38 (0.00) 2.16 (0.00) Oxytrena semicarinata 1-37 1.41 Catinella vermeta 1.35 (0.00) I .89 (0.00) Euconulus chersinus 1.20 (0 ,00) 1.20 (0.00) Stenotrena leai 1.16 (0. 61 ) 1.70 (0.93) Oxyloma spp. 1.16 (0.41) I .69 (1. 19) Kesodon inflectus 1.00 0.00 Pupoides albilabria 1.00 1.00 Pisidium compressum 0.98 0.98 Vertiqo jiycjmaea 0.87 (0.81) 1.16 (0.81) Vertiqo morsei 0.83 (0.00) 1.24 (0.00) Strobilops aonea 0.72 0.72 Valvata sincera 0.64 1.14 239 TABLE 10— Continued

Specios Habitat Diversity Index Calculated Habitat Diversity Index Calculated by Using the Frederick (1967 ) by Using the Environment Control Vegetation Zones Corporation (1973) Vegetation Zones

Carychium nannodes 0.62 (0.00) 0.53 (o.oo) Valvata tricarinata 0.00 0.00 Planorbula armiqcra 0.00 0.00 Vallonia excentrica 0.00 0.00 Gastrocopta procera 0.00 (0.00) 0.00 (0.00) Gastrocopta arnifera 0.00 0.00 Vertiqo elatior 0.00 0.00 Gastrocopta corticaria 0.00 (0.00) 0.00 (0.00)

Numbers in parentheses are habitat diversity indices for living specimens only. TABLE 11

Distribution of molluscs in the various vegetation zones at Cedar Bog Nature Preserve, Champaign County,. Ohio. Vegetation zones according to Frederick (1974).

Species Arbor Vitae Bog Meadow Marl Meadow Maple-Ash-8asswood Elm-Maple Silver-Red Maple Tuliptree Beecb-Maple Oak-Maple Shrub Collected

Valvata tricarinata

Valvata sincera - D D

Acnicola limosa D D D D D

Pomatiopsis lapidaria A A D A A

Cxytreroa semicarinata A D D D D

Lynnaea hunilis A A A D D A

Heliscma ancops - D D D D

Gyraulus parvus D D D D D

Planorbula armiqera - D fe*

Physa integra D A A A D D D A D

Carychium cxiquura A A A A A A A A D

Carychium exile A A A A A A A A

Carychium nannodes A A D

Cochliocopa lubrica - D

Vallonia excentrica -

Pupoidea albilabris < m TABLE 11— Continued

Species Arbor Vitae Bog Meadow Marl Meadow Maple-Asb-Basswood Elm-Maple Silver-Red Maple T u lip tre e Beecb-Maple Qak-Maple Shru Collected

Gastrocopta procera A --- -- D -

Gastrocopta arraifera ------0 -

Gastrocopta contracta ADA A AA A A A 0

Gastrocopta pentodon AAAA AA A A A A

Gastrocopta tappaniana D A A 0 0 A 0 DD D

Gastrocopta corticaria -- - A «• - - - - -

Vertiqo morsei 0 0 A D - - D - D 0

Vertiqo ovata D A . 0 A D - - - 0 0

Vertiqo pygmaca - AA D 0 - D - - -

Vertiqo tridcntata DD D D A A A 0 D A

Vertiqo elatior - - - - D - - - - -

Vertiqo qouldi A A A . D A 0 A - D 0

Vertiqo nilium A A D AA A A - D A

Columella edentula A A - AA AAAA 0

Strobilops aenca 0 - - A «■ -A D - r

Strobilops labyrinthica AAAA 0 AA D DD

Succinea ovalis D A - AQ A A A A D

Catinella veraeta D A A A D 0 m A 0 TABLE 11— Continued

Species Arbor Vitae Bog Meadow Marl Meadow Maple-Ash-8assvfood Elra-Maple Silver-Red Maple Tuliptree Beecb-Maple Oak-Maple Shrub Collected

Qxyloffia ' A A D A

Philonycus carolinianus A A A A

Anquispira alternata A A A A A A

Helicodiscus parallelus A 0 A A A 0 D D A

Punetun minutissimimi A A A A A A A A A A

Deroceras laeve A A A A 0 A A 0

Nesovitrea binneyana A A D D A A

Nesovitrca electrina A A A A 0 A 0 D D

Gl^gh^alinia indentata A A A A A A A A A

Hawaiia niinuscula A A A A A A A A A

Euconulus chersinus D 0 A M A

Euconulus fulvus A A A 0 A D

Zonitoides arboreus A A A A A A A .A

Guppya sterkii A 0 D A A A A 0 A A

Striatura niliura A A 0 A A A A A A A

Striatura exigua A D A 0 A tiaplotrema concavum D D A D 0 242 Stenotrema barbatum A A A A A TABLE 11— Continued

Species Arbor Vitae Bog Meadow Marl Meadow Maple-Ash-Basswood Elm-Maple Silver-Red Maple Tuliptree Beecb-Maple Oak-Maple Shrub Collected

Stenotrema leai 0 AA- D A -- - 0

Mesodon clausus . A - _ A A D A AD

Mesodon thyroidus A- - A - AA A A -

Mesodon inflectus - - -A D - D - A -

Triodopsis nultilinoata - A A - 0 - 0 - D a -

Alloqona profunda - - D - 0 DA- -

Pisidium compressum _ - - D - D -- D-

Pisidium casertanum A 0 . AAD 0 0 - A D

A Found alive at one time or another. 0 Oily fouid dead. - Not found at all. 243 TABLE 12

A comparison of the major statistics concerning the molluscan fauna of each of the Frederick vegetation zones

All Samples Collected Predetermined Quantitative Samples Only

Frederick Total Living Total Total Total Diversity Total Total Living Diversity Number of Total Vegetation Zones Diversity Diversity Number of Number of Diversity Index Number of Number of Diversity Index Living Number of Specimens Samples Specimens Samples Specimens Samples

Marl Meadow 30 23 2 0 ,407 22 29 3-41 20,196 15 21 3 . 4 5 521 15

Bog Meadow 40 28 25 3 9 3 . 8 0 17,692 14 20 3.31 4 2 9 14 1 7 , 9 H ■

Tuliptreo 46 26 4 , 4 2 3 28 4 0 4 . 2 2 2 ,684 15 1 4 2.50 171 15

Maple-Ash-Basswood 46 34 2 ,664 31 36 4 . 2 4 1,836 15 12 2.24 2 48 15

Arbor vitae 44 31 6 , 5 4 9 3 1 38 4 . 0 9 3,991 15 15 2.86 2 3 5 15

Oak-Maple 48 27 2,246 22 43 4 . 2 5 2,107 1 5 14 3 .03 109 15

Elm-Maple 41 20 10,302 17 41 3.88 10,122 15 15 2.78 363 15

Silver-Red Maple 37 23 1,203 19 35 3-85 1,076 15 20 3 . 6 5 138 1 5

Shrub 36 14 5,806 15 36 3 . 7 8 5,806 1 5 1 4 2 .53 360 1 5

Beech-Maple 23 15 807 22 21 -3.49 5 59 15 12 2.25 139 15

------244 245 well. The biggest cause for differences between the total diversity (44) by all collection methods and total diversity by just the quantitative litter samples (38) was mainly the addition of aquatic species and hygrophilic terrestrials from the spring areas. The diversity of living species collected by just quantitative methods (15) was much lower than the living diversity obtained by all methods of collecting (31). This difference is probably indicative of too few samples collected to compensate for the great habitat diversity in the various arbor vitae stands. In this case 16 additional random samples added 16 living species.

Maple-Ash-Basswood Swamp Forest Segregant

The molluscan fauna of the maple-ash-basswood swamp forest has a high diversity. A total of 46 species have been recorded from this zone and 34 species were found living there at one time or another (Table 11). No other vegetation zone exceeds the living diversity. The diversity index, based on the total number collected, is also very high. The remarkable contrast is that the number of living species collected by quantitative methods (1 2 ) and the diversity index (2.24) representing those same specimens is very low (Table 12).

The total diversity is high because there are many diverse microhabitats in this vegetation zone. The moisture conditions are highly variable. There are seasonal ponds, a few perpetually wet spring areas, and drier areas which are never inundated by water. The ground is uneven with numerous mounds of well rotted wood, fallen logs and piles of bark. The water condition and the available shelter is highly variable. In addition to this, the maple-ash-basswood swamp forest was not greatly affected by the elm blight as were some of the other swamp forests. It shows a certain amount of stability. This total habitat variability is expressed by a high diversity and a high total diversity index.

Why the very low diversity index.for living snails? This is a product of not having taken enough quantitative samples to adequately sample all the habitats. There is a fairly large number of species in this fauna that are climbers or log species which do not lend themselves to the kind of quantitative methods used. All three species which were never collected by quantitative methods were present in this zone.

The characteristic snails of this zone are Glyphyalinia lndentata, Philomycus carolinianus, Mesodon inflectus, Mesodon thyroidus, Zonitoides arboreus and Carychium exile. Four of these species are usually associated with fallen trees and loose bark. Gastrocopta corticaria is the only snail restricted to this vegetation zone at the present time. Seven of the forty-six species are aquatic forms. In some samples aquatic mollusca were the most abundant. The densities for most species in this zone are quite low. In spite of the 31 samples made, only 2664 specimens were collected. 246

Tuliptree Swamp Forest Segregant

Frederick (1967 and 1974) distinguished a tuliptree segregant of the swampt forest. The Environment Control Corporation (1973) com­ bined this zone with several other swamp forest segregants. This swamp forest segregant is broken up into five isolated stands. Three of these stands are very wet and two are much drier. The drier stands are small, narrow and probably greatly influenced by surrounding vegetation.

The difference between the total diversity (46) and the living diversity (26) is striking. Why is there such a large number of species found only as dead shells? I am not sure of the answer, but several other vegetation zones show the same thing. The oak-maple, shrub and elm-maple zones all show a large number of dead species. What do all these areas have in common? They are areas which appear to be in the state of change or have changed sometime in the past. The tuliptree zone appears to be relatively stable, but it may not be now or may not have been in the past.

I found nine aquatic species in the tuliptree swamp forest, but only Pomatiopsis lapldaria was found alive. Most of the aquatic species probably live only in the seasonally ponded areas for a short time. It is possible that they are short lived and I missed them when they were alive. Four of the five tuliptree stands are adjacent to arbor vitae stands, this could explain the high fre­ quency of Guppya sterkii and Nesovitrea binneyana which are more characteristic of the arbor vitae association. Carychium exiguum, Strobilops labyrinthica and Pomatiopsis lapidaria appear to be the characteristic species of the wet sections of this swamp forest.

Bog Meadow Association

The molluscan fauna of the bog meadow shows a total diversity of 40 species and a living diversity of 28 species (Table 11). I con­ sider these number fairly high. The overall density of the mollusks Is very high. I found 17911 specimens in 25 samples, a mean of 716.4 speci­ mens per sample.

The bog meadow is a fairly uniform and stable community. The substrate, vegetation, cover and moisture conditions are not nearly as variable as those found in many of the forested areas. This uni­ formity shows up in the sampling results. A comparison between the total diversity (39) found in the quantitative substrate samples and the total diversity (40) found by all collection methods are almost the same. The quantitative substrate sampling procedure works well for this community. There are no fallen trees, miscellaneous logs, piles of bark and deep water pools to bias the collections made. The number of different molluscan habitats is small in comparison to other vege­ tation zones in the study area. 247

My reasons for believing that the present bog meadow has been a fairly stable community, is based on the lack of composition changes in the molluscan fauna and the low number of species represented only by dead shells. The water table dropped many years ago after a regional dredging operation, but most of these changes have been since stabilized. This dredging probably reduced the overall area of the bog meadow, but there is no evidence that it drastically changed the bog meadow that persists today. Arbor vitae is becoming established in several areas, but this is normal and a very slow process. This process will eventually reduce the bog meadow even more.

There are 12 species which are found only as dead shells. Six of these are aquatic species and six are terrestrial species which are more characteristic of adjacent vegetation zones.

What really characterizes the bog meadow molluscan fauna is the large number of species which have a high frequency of occurrence in this association. There are at least 15 species which are characteris­ tic of the bog meadow. All of these species show distribution patterns which link them to wet or very wet habitats. Vertigo milium and Triodopsis multilineata are the most characteristic species of the bog habitat. The molluscan fauna of the bog meadow association is unique because of its 1) high diversity, 2) high density, 3) uniformity and 4) distinctive composition.

Marl Meadow Association

The marl meadow association has the most distinctive molluscan fauna at Cedar Bog. I only found 30 species present, but 23 species were found living (Table 11). The only vegetation zone with a lower diversity is the beech-maple forest. The total diversity index for the marl meadow is the lowest of all zones (Table 12). The diversity may be low, but the density is very high, 20,196 specimens were ob­ tained in 22 collections, a means of 918 specimens per collection.

There are 18 species commonly collected in, or characteristic of, the marl meadow. Of those 18 species, 16 are collected more often in the marl meadow than any other vegetation zone in the Bog. For example, there are five aquatic species found in the marl meadow, all five are found living there and all five species are more frequently collected in the marl meadow than any other vegetation zone in the Bog.

The difference between the total diversity (30) and the living diversity (23) is only 7 species, the smallest difference of any zone in the Bog. In my opinion the marl meadow is the most stable, slowest changing vegetation zone in the Bog. This low number of dead shells Indicates that there is very little evidence of past changes. There are very few species, if any, that were living in this zone in the recent past that are not now living there. 248

Another important aspect of the marl meadow is its habitat uni­ formity. I found very little difference between the diversity recorded by quantitative methods and the diversity recorded by all collection methods. This evidence demonstrates the thoroughness of quantitative sampling procedure in the marl meadow.

Almost all species recorded from the marl meadow are those which can thrive in constantly very wet conditions. The only species which is unique to the marl meadow is Vertigo morsel. The fauna is dominated by aquatics, succineids, pupillids and very few polygyrids. The marl meadow molluscan fauna is characterized by a 1 ) low diversity, 2 ) high density, 3) uniformity in composition and 4) many distinctive, specialized species.

Beech-Maple Association

If the marl meadow is one extreme, the beech-maple forest is the other. This is the driest vegetation zone in the study area. I did not find one shell of an aquatic species. The total diversity (23) and the total diversity index (3.49) are very low (Table 11 and Table 12). The living diversity (15) and the living diversity index (2.25) are the second lowest in the study area. The overall density is also the lowest in the study area with 807 specimens In 22 collections.

There are at least two possible reasons for this low density and diversity, the dryness and the lack of variable types of shelter. There are no springs, seasonal pools or streams in this area and this entire area is the greatest distance from a permanent source of water. The general shelter is not good. A leaf litter does accumulate In some areas, but apparently dries out. There are a few fallen logs and dead trees, but not nearly the number that are found in some swamp forest segregants. The habitat diversity in the beech-maple forest Is fairly low. Compared with other zones at Cedar Bog, the beech-maple forest provides a relatively uniform molluscan habitat. The differ­ ence between the diversities determined by all collecting methods and quantitative collecting methods is low. Most of the species found in the beech-maple forest were collected by quantitative methods.

This beech-maple forest has little evidence of recent changes. There are few species (8 ) recorded only as dead specimens. The charac­ teristic species of this vegetation zone are Stenotrema barbatum and Succinea ovalis. There are only six species living In the beech-maple forest that are also living in the marl meadow. Most of the species found living in the beech-maple forest have rather wide habitat tolerances or are usually associated with drier habitats. The molluscan fauna of this area is characterized by a 1 ) low diversity, 2) low density, 3) no aquatics, 4) few pupillids and 5) species which can survive drying conditions. 249

Oak-Maple Hardwood Forest .

The total diversity (48) of molluscs in the oak-maple forest is the highest for any vegetation zone in the study area (Table 11). The living species diversity is much lower at 27. There are several reasons for this high diversity and large number of dead species.

The oak-maple forest is bordered on the east side by the West Branch of Cedar Run. This branch was dredged many years ago. The dredging operation drastically altered this vegetation zone (Fred­ erick, 1967: 90-91). The water table was lowered making the immedi­ ate area drier, but the spoil bank, composed of some glacial outwash, was also piled along the banks of this area. This made some parts of this narrow zone much drier. The numerous springs along this bank were probably reduced and today are confined to small seepage areas. These are good evidences that this area was once much wetter. Eight of the dead species were those which are typical of the bog and marl meadows today. Many of the dead species are also aquatics brought up with the dredgings.

In the oak-maple forest today the molluscan habitats are fairly diverse for such a small area. There are numerous seepage areas, moist to wet muck and dry spoil banks scattered with some leaf litter and logs.

Some of the characteristic molluscs of the oak-maple forest are Philomycus carolinianus, Zonitoides arboreus, Pomatiopsis lapidaria, Guppya sterkii, and several aquatic species. The living fauna is similar to that found in the maple-ash-basswood swamp forest across the stream.

One specimen each of Gastrocopta armifera, Gastrocopta procera and Pupoides albilabris were also found in the oak-maple forest. These species are more typical of dry open areas. The west side of this forest is bordered by an open meadow which could be the main habitat and source for these species.'

The total molluscan fauna is an odd mixture of many dead species typical of much drier areas and much wetter areas. This molluscan assemblage was a 1 ) moderately high diversity, 2 ) low density, and 3) a swamp forest composition.

Silver-Red Maple Swamp Forest Segregant

The silver-red maple swamp forest is very different from the other swamp forest segregants. Here the trees are very young, even aged and the canopy diversity is very low. There are few large dead trees and fallen logs. There are no springs or seasonal ponds. The leaf litter is moderate and does provide good shelter. The entire area is very uniform. 250

The total diversity (37) and the living diversity (23) are inter­ mediate compared with other zones in the study area (Table 11). The total diversity index is also intermediate, but the living diversity index is the highest in the study area (Table 12). This high index is the result of both an intermediate diversity and a very even distri­ bution of this diversity. There is very little clumping of the species in this zone. The general density is low, with only 1203 specimens collected in 19 samples, a mean of 63 specimens per col­ lection.

The data substantiate the visual uniformity of this zone. It has already been mentioned that the diversity index for living speci­ mens is very high with a diversity of only 23. This could be attributed to the evenness of the species throughout the zone. The difference between the diversity of quantitative collections and all the collections is not very large. This also shows that most of the species found there can be collected by quantitative methods. The quantitative collection method used works best In the most uniform habitats.

There are eight aquatic species recorded for the silver-red maple swamp forest. All of these were found dead and most were found along the spoil bank left from dredging of the West Branch of Cedar Run. This zone was not as adversely affected as the oak-maple forest because it was a larger area and not as much was damaged.

The species most often collected In this zone are small soil and litter inhabitants. For a zone with few dead trees and fallen logs there are several large species which can definitely be considered characteristic. Stenotrema barbatum appears to be able to survive in the loose muck soil. Succinea ovalis is a common inhabitant, usually climbing on trees and low vegetation. Mesodon thyroidus, Mesodon clausus and Anguispira alternata are common in the few logs and fallen bark habitats that do exist here.

This vegetation zone will, of course, change a great deal as the trees mature and thin out. This maturing process will change the uniformity by adding fallen logs and increasing the overall habitat diversity. The diversity of the mollusca should increase with successional changes. The molluscs found here are now typical of moist to dry habitats. There are very few wetf-habitat species there now. The basic composition of the fauna should not change unless the current vegetation and/or water table changes.

Shrub Zones

There are six designated shrub stands in the study area. Only the two largest stands were sampled. The shrub zones sampled provided little cover in the form of logs, and leaf litter. The only substan­ tial cover was the loose granular muck soil. 251

The total species diversity is somewhat intermediate with 36 species recorded, but the living species diversity of 14 is the lowest in the Bog (Table 11). The largest species found living in these zones was Stenotrema barbaturn and Glyphyalinia indentata, both of these snails find loose, granular muck adequate cover. All the other living molluscs were very small soil and litter inhabiting species. Striatura milium, Hawaiia minuscula, Helicodiscus parallelus, Gastrocopta pentodon, Punctum minutissimum and Glyphyalinia indentata are the most commonly collected species in the shrub zones. Dead shells of four aquatic species were collected, but I found no evidence that any of these snails are currently living in the shrub zones examined.

The diversity indices are medium to low and are probably accur­ ate because of the uniformity of the shrub zones examined (Table 12). The overall density is intermediate with a total of 5806 specimens collected in 15 samples, having a mean of 387 specimens per collec­ tion.

The difference between the total diversity and the living divers­ ity is most significant. There were 22 species found only as dead shells. Host of the dead shells (15) were those now found in the wetter areas, such as the bog and marl meadows. This is good evidence that these shrub zones are drier today.

The two shrub zones examined are successional communities. More than likely they will become some type of swamp forest in the future. In general these shrub communities possess a low diversity special­ ized molluscan fauna capable of living in a moist, loose, granular muck soil.

Elm-Maple Swamp Forest Segregant

The elm-maple swamp forest currently has an uneven, high divers­ ity, high density fauna which gives evidence of some recent changes. The total diversity is 41 species and the living diversity is 20 species (Table 11). The composition of the living species is very different from the composition of the dead species. Host of the common living species are small soil and litter inhabitants. Many are the same as those currently found in the shrub zones. There are also several species which are common as dead shells, but are rare or absent as living members of the fauna. These are Vertigo milium, Vertigo gouldi, Gastrocopta tappaniana, Pomatiopsis lapidaria, Nesovitrea, Carychium exiguum and Oxyioma spp. These are wet habitat species particularly common today in the bog and marl meadows. I found only one of the seven aquatic species living. There is very good evidence that this area was much wetter in the recent past (50 to 100 years?). The large number of species only found dead, the composition changes and the high density indicates an area which was once similar to the bog meadow. 252

The overall density is high with 10,302 specimens collected in 17 samples, having a mean of 606 specimens per collection. This density is not even. The high density samples were found only in areas imme­ diately north of the present bog meadow (W, X, Y- 1, 2). If Dr. Frederick's conation lines are correct, this would indicate that the bog meadow is or has receded substantially in the recent past.

The elm-maple swamp forest is not a very uniform habitat for terrestrial mollusca. It has been affected in the western part by past dredging, lowering of the water table, an American elm die off and several changes in the dominant vegetation. The dominant cover today is a loose granular muck soil with patches of fallen leaves and some fallen logs.

Environment Control Corporation Vegetation Zones

I do not feel that a complete analyses of the molluscan fauna of each of the vegetation zones designated by the E. C. C. (1973) would substantially increase the understanding of the relationships between the molluscs and the vegetation zonation. I will, however, make some general comparisons. Table 13 shows the general distribution of each species in the various E. C. C. vegetation zones. Table 14 summarizes the various statistics concerning the molluscan faunas as applied to each of the E.C.C. vegetation zones. These tables are set up in the same manner and order as those summarizing the molluscs in the Frederick vegetation zones.

Where one draws the line between one vegetation zone and another can be very important. A comparison of the general statistics between the mixed-beech forest of the E. C. C. and the beech-maple forest of Frederick shows only small differences. The basic definition of this forest area is the same, but the boundaries are not. A comparison of data between Tables 12 and 14 show the boundaries are not too important here because a change from a beech-maple forest to a shrub or swamp forest is not abrupt; at least as far-as molluscan habitats are con­ cerned.

On the other hand, a comparison of the arbor vitae association of Frederick with that of the evergreen and evergreen-mixed hardwood forests of the E.C.C. show some significant' differences. In the field, I did not find it difficult to differentiate the boundaries of the various arbor vitae stands. A liberal interpretation of these forest areas by the E.C.C. includes some very diverse molluscan habitats under one zone. The E. C. C. Included a lot of bog meadow and tulip­ tree swamp forest in their interpretation. This changed the total number of specimens collected from 6549 to 18914 with two less samples collected. The result is an average density change from 211 specimens per sample to 652 specimens per sample. The diversity and the divers­ ity index for living species collected also changed, from 31 to 36. TABLE 13

Distribution of molluscs in the various vegetation zones at Cedar Bog Nature Preserve, Champaign County, Ohio. Vegetation zones according to Environment Control Corporation (1973)*

Species * Marl Non-Marl Tall Sycamore- Sycanore- Evergreen Tuliptree- Maple-Elm Maple-Elm Maple- Mixed-Beech Collected Short Short Grass Basswood- Maple- Evergreen- Basswood-Ash # 1 # 2 Ash-Elm Grass Grass Cherry Cottonwood Mixed

Valvata tricarinata ------D mm - - -

Valvata sincera - - - - - D D D - - -

Amnicola linosa D -- D D -DD mm --

Pomatiopsis lapidaria A A AA D AA 0 D D -

Qxytrema semicarinata D mm - DD A 0 D - - -

Lymnaea hunilis AD 0 0 -A A DD D -

Helisoma anceps - - - D -D 0 tm - - -

Gyraulus parvus A ■ D DA - AD DD DD

Planorbula armigera - - - - D - -- -

Physa integra A D - D D A A D D D -

Carychium exiquurn A A • D A -AAA D D D

Carychium exile A A A AA AAA A A

Carychium nannodes A - - A - 0 D - * m D -

Cochliocopa lubrica m m - D - - 0 D mm D -

Vallonia excentrica - - D ------253

Pupoides albilabris - -- D - 0 --- TABLE 13— Continued

Species Marl Non-Marl Tall Sycamore- Sycanore- Evergreen Tuliptree- Maple-Elra Maple-Elra Kaple- Mixed-Beech Collected Short Short Grass Basswood- Maple- Evergreerv- Basswood-Ash # 1 # 2 Ash-Elm Grass Grass ' Cherry Cottonwood Mixed

Gastrocopta procera - _ - D - - - A ---

Gastrocopta artnifera - - - - - m 0 - - --

Gastrocopta contracts A A 0 AA A A A D 0 A

Gastrocopta pentodon AA AA A AAA A 0 A

Gastrocopta tappaniana A 0 D 0 - AD A D D D*

Gastrocopta corticaria - - A ------

Vertiqo morsei A 0 - D - D D 0 0 0 -

Vertiqo ovata 0 D b D a* A A D - - m

Vertiqo pyqmaea A D - D - A D - - - -

Vertiqo tridentata D D - A •- A D A 0 D -

Vertiqo elatior 0 - - - a* - --- -

Vertiqo qouldi A A A A m A 0 0 D D -

Vertiqo milium AAA AAAA 0 0 -

Columella edentula A AA A 0 A A A D A A

Strobilops aenea - - . _ - - D A - -- 0

Strobilops labyrinthica A A AA 0 A AA 0 D 0

Succinea ovalis - D - A - D A AAAA 254 TABLE 13— Continued

Species Marl Non-Marl T a ll Sycanore- Sycanore- Evergreen Tulip tre& - Maple-Elm Maple-Elm Maple- Mixed Beech Collected Short Short Grass Basswood- Maple- Evergreer>- Basswood-Ash #1 #2 Ash-Elm Grass Grass Cherry Cottonwood Mixed

C a tin e lla vermeta D 0 - 0 - A AD 0 D - 1 Oxylcma A AA DA AAD A D -

Philoraycus carolinianus - - - A - A A - ■ - -

Anquispira altornata D - 0 A - - A A - - A I Helicodiscus parallclus - 0 - A 0 A 0 A 0 0 A

Pune tun ninutissiraum A AAAAA AAD A A

Deroceras laeve D A «• D - AA DD 0 -

Nesovitrea binncyana A AA A - A D A - D -

Nesovitrea electrina A D - A- A A A 0 D 0

Glyphyalinia indentata A A AA D AAA D A A

Hawaiia ninuscula A A D AA A A AD AA

Euconulus chersinus - D AA *» - - D - - -

Euconulus fulvus A D D A -A A 0 0 A -

Zonitoides arboreus AA 0 A - A A A -A A

Guppya s te r k ii A AA A A A A A - A D

S tria tu ra milium AA AA AA AA AA A 255 S tria tu ra exiqtta - - w D «■ D 0 A — A • TABLE 13— Continued

Species Marl Non-Marl T a ll Sycamore- Sycaraore- Evergreen T u lip tre e - Maple-Elm Maple-Elm Maple- Mixed Beech Collected Short Short Grass Basswood- Maple- Evergreen- Basswood-Ash # 1 #2 Ash-Elm Grass Grass Cherry Cottonwood Mixed

Haplotrema coacavun D -- 0 - 0 A - D D

Stenotrema barbatun AA 0 A - A A A 0 0 A

Stenotrema le a i AAA D - A - A 0 - -

Mesodon clausus D D - 0 - «■ A AA A A • Mesodon thyroidus mm A - - - A AA- 0 A

Mesodon in fle c tu s - 0 - A - - A - - mm -

Triodopsis multilineata DAA D - A - - - - -

Allcgona profunda ------0 D - DA

Pisidium compressum - - mm -- D 0 - mm -

Pisidium casertanum AD - D 0 A A 0 D D -

A Found alive at one time or another. 0 Only found dead. Not found a t a l l . 256 TABLE 14

A comparison of the major statistics concerning the molluscan fauna of each of the Environment Control Corporation vegetation zones.

A ll Samples Mapped Predetermined Q uantitative Samples Only

E. C. C. Total Living T otal T otal T otal D iversity Total Total Living D iversity Number of Total D iv e rs ity D iversity Number of Number of D iversity Indox Number of Number of D iversity Index Living Number of Vegetation Zones Specimens Samples Specimens Samples Specimens Samples

Marl Short Grass 40 29 19,754 20 37 4.03 19,301 17 25 3.63 507 17

Non-Marl Short Grass 39 22 3,486 11 32 4.02 2,586 6 11 2.90 132 6

T a ll Grass 27 16 936 5 12 2 .6 l 393 l 2 .92 3 1

T u lip tre e - BasswoocLAsh 51 32 3,782 41 45 4.26 3,129 27 12 1.94 336 27

Evergreen and Evergreen-Mixed 46 36 18,914 29 42 3 .82 17,397 20 26 3.65 558 20

Maple-Elm-Ash 38 12 2,616 16 - 36 4.08 2,491 14 10 2.00 112 14

Maple-Elm § 1 45 26 2,485 23 41 4 .25 1,574 17 21 3.68 139 17

Maple-Elm § 2 29 5 5,522 6 29 3.50 5,501 5 3 1.31 25 5

Sycamore- Basswood-Cherry 51 29 13,184 31 49 4.03 12,871 27 22 3.11 730 27

Sycamore— Maple-Cottonwood 17 8 266 1 17 3.3S 266 : i 8 2.41 34 l

Mixed-Seech 24 16 683 18 22 3.52 560 14 13 2.33 137 14 257 258

These changes are due to the Inclusion of small scattered arbor vitae trees which are currently invading the bog meadow as part of the ever­ green and evergreen-mixed hardwood forest. This makes the habitat diversity much broader for both terrestrial and aquatic molluscs.

Similar results are obtained by making comparisons in several, other vegetation zones. Only seven of the eleven E. C. C. vegetation zones can be compared, because of the uneven distribution of samples collected. This uneven sample distribution for the E. C. C. vegeta­ tion zones lowered the average diversity, the diversity Indices, the average number of samples collected per zone and widened the diversity index range.

Several points can be made from a comparison of these two inter­ pretations. I would need to collect samples in the E. C. C. vegeta­ tion zones in the same manner as I did in the Frederick vegetation zones in order to fairly compare them species by species. The position of a boundary between two very diverse vegetation zones is very important, more important than between two less diverse zones. I am currently under the opinion that a general vegetation interpretation, which includes many very diverse habitats, is not as helpful in determining molluscan ecology as a more rigid or restricted interpre­ tation would be. Indistinct ecotonal areas, such as some of the shrub zones, should be separated from more pure vegetation zones. It is possible that we may find some species are restricted to or more dense in ecotonal areas. From a malacological point of view, the separation of an area into many different vegetation zones is far more informative than lumping them all under a few plant associations.

Substrate Or Soil Zonation

After comparing the soil maps produced by Ritchie et al. (1971) (Map II) and E. C. C. (1973) (Map III) It is difficult to believe they are describing the same area. Yet a look at some general statistics show some similarities and molluscan diversity patterns do exist. Tables 15 and 16 make it possible to compare each species collected with either soil interpretation. Table 17 summarizes the general statistics obtained for all substrate interpretations. The various soil types are arranged from top to bottom in each subdivision from the wettest to the driest.

Marl is easy to identify in the field. It does grade Into some muck and peat, but in general I found It readily separable. A com­ parison between the total and living diversity and the total and living diversity indices of the E. C. C. Interpretation and my field observation are very.close. The same comparisons made by using my Interpretation of the marl zone on the Ritchie et al. (1971) soil map are not close at all. This leads me to the conclusion that my marl interpretation of the Ritchie et al. map (Map II) Is too restrictive. 259 TABLE 15

Distribution of molluscs in tho soil zones at the Cedar Bog Nature Preserve, Champaign County, Ohio, Soil zones modified from Ritchie et al.

Species Lippincott Silty-clay Loam Linwood Muck Carlisle Muck Peat Marl Collected

Valvata tricarinata - - D - ••

Valvata sincere - 0 0 - -

Amnicola limosa - D D 0 -

Pomatiopsis lapidaria D 0 A AA

Oxytrema semicarinata - D D A -

Lymnaea humilis DD A AA

Helisoma anceps - - D D mm

Gyraulus parvus D D A A A

Planorbula armiqera - -- D -

Physa inteqra D D A AA

Carychium exiquum D A A A A

Carychium exile A A AA-

Carychium nannodos -A D -

Cochliccopa lubrica - D D - m m

ValIonia excentrica - - D - -

Pupoides albilabris -D mm -

Gastrocopta procera - • D mm A -

Gastrocopta armifera - - D - -

Gastrocopta contracts A A A A A

Gastrocopta pentodon AA A A A

Gastrocopta tappaniana D A 0 AA

Gastrocopta corticaria - - A - -

Vertigo morsei D - D AA

Vertigo ovata 0 A A 0

Vertiqo pyqmaoa - D D A A

Vertiqo tridentata - A A D D

Vertiqo elation tm D - - - 260 TABLE 15— Continued

Species Lippincott Silty-clay Loam Linwood Muck Carlisle Muck Peat Marl Collected

Vertiqo qouldi D D A AA

Vertiqo milium 0 A AA D

Columella edentula AA AA -

Strobilops aenea 0 - A 0 -

Strobilops labyrinthica DD AA A

Succinea ovalis A AA A

Catinella vermeta m m 0 A 0 A

Qxyloma 0 AA AA

Philomycus carolinianus - - A - -

Anquispira alternate A AA ** 0

Helicodiscus parallelus 0 A A A -

Punctum minutissimum D A A AA

Derocerae laevo D - A A A

Nesovitrea binneyana - D A A ««

Nesovitrea electrina D D A AA

Glyphyalinia indentata A A A A A

Hawaiia minuscula AA A AA

Euconulus chersinus - - A A p*

Euconulus fulvus - AA A A

Zonitoides arboreus A AA A -

Guppya sterkii D A A A 0

Striatura milium A A A A 0

Striatura oxiqua - A A A -

Haplotrema concavum - 0 AD -

Stenotrena barbatum A A A A 0

Stenotreraa leai - 0 A AA

Mesodon clauses A AA A -

Mesodon thyroidus A A A A - 261 TABLE 15— Continued

Species Lippincott Silty-clay Loam Linwood Muck Carlisle Muck Peat Marl Collected

Mesodon inflectus - - A D -

Triodopsis multilineata - - 0 A A

Allopona profunda •■A 0 0 - -

Pisidium compressum D D - -

Pisidium casertanum m m 0 A A A

A Found alive at one time or another. D Only found dead. - Not found at all. TABLE l6

A comparison of the major statistics concerning the molluscan fauna of each of the various soil zones.

Soil Zone All Samples Mapped Predetermined Quantitative Samples Only •

Modified Total Living Total Total Total Diversity Total Total Living Diversity Number of Total U. S. 0. A. Diversity Diversity Number of Number of Diversity Index Number of Number of Diversity Index Living Number of Specimens Samples Specimens Samples Specimens Samples

Marl 30 23 16,410 14 3 2 3 . 5 8 16,374 13 21 3*39 513 13

Peat 48 39 27,677 50 4 2 3 . 9 0 2 4,060 3 1 2 4 3 . 4 8 58 o 3 1

Carlisle Muck 57 42 20,913 88 54 4 . 2 6 1 9 , 3 2 2 65 2 7 3.51 1,279 65

Linwood Muck 46 23 5 , 3 4 3 26 4 3 4 . 1 4 5 , 2 7 3 2 3 18 3.16 171 23

Lippincott S ili y - C l a y Loam 31 14 1,292 23 30 3 . 7 4 1 , 0 4 7 17 12 2.09 1 64 17

E. C. C.

Marl 46 -38 2 8 , 5 7 7 4 6 4 5 3 .83 2 6,384 3 0 28 4 . 0 5 612 30

Fibrous Peat 54 39 10,999 53 4 9 4 . 4 4 9 ,240 38 22 3-38 448 38

Muck 53 33 15,046 59 51 4 . 1 6 1 4,676 4 7 27 3 . 0 8 GO 4 7

Woody Peat 45 31 , 16,005 19 40 4.01 14,891 15 2 7 3.60 636 15-

Lippincott Silty-Clay Loam 26 16 1 ,008 2 4 24 3 . 4 8 885 1 9 1 4 2.27 1 54 19 262 263 TABLE 17

Distribution of molluscs in the soil zones at the Cedar Bog Nature Preserve, Champaign County, Ohio. Soil zones modified from Environment Control Corporation (1973)•

Species Lippincott Silty-clay Loam Muck Woody Peat Fibrous Peat Marl Collected

Valvata tricarinata - D - -

Valvata sincera - 0 - D D

Amnicola limosa - D - D -

Pomatiopsis lapidaria - A AA A

Oxytrema semicarinata - D - 0 A

Lynnaea humilis - A A AA

Helisoma anceps - D DD D

Gyraulus parvus D D AD A

Planorbula armigera - - D - -

Physa Integra - A D AA

Carychium exiguum DA A A A

Carychium exile AAAAA

Carychium nannodes -- AAA

Cochliocopa lubrica - 0 - 0 -

Vallonia excentrica - - D --

Pupoides albilabria - D - - -

Gastrocopta procera -• D - A -

Gastrocopta armifera -- D -

Gastrocopta contracts A A AAA

Gastrocopta pentodon AA A AA

Gastrocopta tappaniana DD A .A A '

Gastrocopta corticaria - A - - -

Vertiqo morsei - D D D A

Vertigo ovata - D AA 0

Vertiqo pygmaea - - A D A

Vertiqo tridentata D A A D A 264 TABLE 17— Continued

Species Lippincott Silty-clay Loam Nuck Woody Peat Fibrous Peat Marl Collected

Vertiqo elatior - - - - 0

Vertiqo qouldi 0 DAA A '

Vertigo milium - A A A A

Columella edentula A A A A A

Strobilops aenea 0 - 0 AA

Strobilops labyrinthica D A A A A

Succinea ovalis A A 0 A 0

Catinella vermeta - 0 A A 0

Oxyloma - AD A A

Philomycus carolinianus - A - AA

Anguispira alternate A A 0 AA

Helicodiscus parallelus AAAA A

Punctum minutissimum 0 AAA A

Deroceras laeve - - 0 0 AA s Nesovitrea binneyarta - A AA A

Nesovitrea electrina 0 A A A A

Glyphyalinia indentata A A A AA * Hawaiia minuscula A AAAA * Euconulus chersinus - A A -

Euconulus fulvus - 0 A A A

Zonitoides arboreus A A A A A

Guppya storkii A A A •A A

Striatura milium A A A A A

Striatura exigua - A -A 0

Haplotrcma concavum 0 DDA -

Stenotrema barbatum A A A A A

Stenotrema leai AAAA 2 6 5 - TABLE 17— Continued

Species Lippincott Silty-clay Loam Muck Woody Peat Fibrous Peat Marl Collected

Mesodon clausus A A D 0 -

Mesodon thyroidus A A A AA

Mesodon inflectua - A DD D

Triodopsis multilinoata - 0 0 A A

Allogona profunda A D - D-

Pisidium compressum - 0 - 0 -

Pisidium casertanum - A A 0 A

A Found alive at one tine or another. D Only found dead. - Not found at all. 266

There is definitely a distinct molluscan fauna associated with marl at Cedar Bog which is discussed under the marl meadow association.

Muck covers much of the study area. The molluscan diversity and the diversity indices for muck soils are very high no matter whose interpretation is used. This soil type is variable and grades into loam, peat and marl, but in general the molluscs which dominate this soil type are swamp forest species typical of a consistently moist or a seasonally very wet substrate. In general, this is the middle area between the extremes of marl and loam.

The molluscs of the peat substrate are not easily compared. All the peat substrate are not easily compared-. All the peat interpreta­ tions show significant differences in the diversity and diversity index statistics. There is very little similarity between the two interpre­ tations presented on Maps II and III. My field observations do not seem to help distinguish which interpretation is most correct.

The molluscan fauna of the loam soils is essentially the same as that of the beech-maple or mixed beech vegetation zones. In the field there is a gradual change from loose granular muck to a definite compacted silty loam. My interpretation of this loam in the field is restricted and this shows up by a low total diversity and a low diversity index.

There are 18 species of molluscs not found living in the loam soil zones and only one species, Allogona profunda, which was only found living in the loam. Mesodon inflectus and Gastrocopta corticaria, two rare species, were limited to muck soils and Gastrocopta procera was limited to a peat substrate. Vertigo pygmaea and Triodopsis multilineata were found living only In marl and peat substrates, whereas Euconulus chersinus, Striatura exigua and Haplotrema concavum were found only in muck and peat soils. It is difficult to consider these correlation as firm, because most of these species are uncommon or rare In the study area.

In general the species diversity is highest in the muck, followed by the peat, marl and then loam soils. The composition of the living fauna varies with the substrate but these composition changes are probably due to moisture changes rather than substrate changes.

At Cedar Bog, vegetation, soil, cover and water conditions are interrelated. All vegetation affects the soil forming beneath It, and eventually the soil moisture, and cover, which is available for the molluscs. The water and parent material initially determines which course of silccessional changes will occur. It is very difficult, if not Impossible, to isolate any of these factors as being an absolute in determining the distribution of the molluscs at Cedar Bog. 267

Factors Affecting the Distribution of Mollusca

There are many physical as well as biological factors which effect the distribution of mollusca within a given area. At this time, it is not possible to know the importance and effects of most of them for even a small fraction of the species. The factors I believe are the most important or the most obvious, are the avail­ ability of calcium, pH, food, predators, moisture or water conditions, the amount and type of shelter available and climate. The signifi­ cance of some of these factors, such as food and predators are beyond this type of study. Some of these factors probably do' not have any serious effects on the distributions of molluscs in Cedar Bog today.

Cedar Bog is an area of some dramatic extremes, particularly with regard to moisture and temperature. The climatic conditions recorded within the Bog show greater temperature variations, fewer frost free days and higher humidities than the surrounding country­ side. This must have some effect on the molluscan populations, but it is almost, if not, impossible to single out any one of these factors as being most important." It is likely that the shrub, swamp forests and beech-maple forest in the southwestern portion of the study area has microclimatic conditions significantly different from the bog meadow, arbor vitae, and marl meadow zones in the northeastern portion of the study area. These difference could have some effects on diversity, density and composition of the molluscan fauna. Unfortunately, not enough is known about these differences to measure their effects.

Calcium and pH as Limiting Factors

In general a source of calcium can be very Important in limiting the distribution of some species. Boycott (1934: 29-34) and Burch (1955: 62-68) studied the effects of calcium as a limiting factor on terrestrial mollusca. One usually finds higher densities and greater diversity in areas with abundant calcium and the opposite in those areas with little calcium. At Cedar Bog, the entire areas is under­ lain with a calcareous outwash. The water moving up through this outwash is supersaturated with dissolved calcium compounds. Much of the highly organic soils of Cedar Bog, which would naturally be acidic, are close to neutral due to this water. Even in the areas where there is not a source of alkaline spring water, the soils show only a slightly acidic to neutral reaction. The peat which is frequently elevated above the marl Is probably the most acidic subs tract in the Bog, except where springs are located. The exact effects of this highly alkaline calcium laden water are not known, but in general I am convinced that it is one of the prime reasons for the high overall diversity of Cedar Bog Nature Preserve. It is again not clear whether the availability of calcium effects the specific distribution of any species within the study area. The difference between the amount of 268

available calcium in the marl meadow and the beech-maple forest is very great. Only Vertigo mOrsei and Vertigo pygmaglare strongly associated with marl at Cedar Bog. They may be calclphiles.

A major factor which clouds the calcium issue is water. All the marl or high calcium areas are saturated continually with water and the lower calcium areas have the lowest amount of water. Another complicating factor is the type of shelter available. In the wet high lime areas the cover is mainly clumps of sedges and grasses. In the drier forested areas it is mainly leaf litter, logs and bark. This In itself is a significant limiting factor for some species. However, I am not convinced that the availability of calcium is a major limiting factor affecting the distribution of mollusca in the Bog. I am fairly sure that pH is not a major limiting factor at Cedar Bog, At this time I believe that the amount of water, the amount and type of cover are the most Important factors affecting the distribution of molluscs at Cedar Bog.

Water As a Limiting Factor

The variations of water and cover within the study area are almost infinite. I made an attempt to rank the various Frederick vegetation zones from the wettest to the driest for a comparison with the diversity and diversity index data (Table 12).

Evaluating the water conditions of Cedar Bog is a very complex problem. It is not a simple question as to whether a particular area is wet or dry, but how wet, for how long and how dry, for how long. This is a study in itself. For many species of molluscs a constant source of water is very important. Previously reviewed habitat data indicate that many of the aquatic species and some of the so-called amphibious species found at Cedar Bog require water much of the time. Distribution patterns for these species show a close correlation with ponded areas, springs and streams. The springs are common in four vegetation zones, the marl meadow, bog meadow, arbor vitae and the oak-maple zones. The seasonal ponds are present In the tuliptree and raaple-ash-basswood swamp forests, usually from late fall through part of spring. These ponded areas are quite dependent on how much and when local precipitation occurs. Some of the swamp forest segregant3 bordering the arbor vitae or bog meadow zones have wet areas due to influences by nearby springs.

The most difficult vegetation zones to classify are those which have uneven ground and some springs and ponded areas. The tuliptree and raaple-ash-basswood swamp forest are ponded in some areas and never ponded in other areas. The differences in its molluscan fauna are dramatic. The greater these differences, the greater the molluscan diversity. The oak-maple forest borders the West Branch. It has high areas next to low wet areas with springs. The total molluscan fauna reflects these differences. ' 269

It appears that if it were possible to map out accurately the water conditions of Cedar Bog, the distribution patterns for many living terrestrial and fresh-water molluscs could easily be related to it. The distribution maps constructed for the species found in this study show varying degrees of association with the general moisturp conditions within the study area.

Shelter As a Limiting Factor

Another significant limiting factor, which appears to be very important at Cedar Bog, is shelter or cover. Areas which have many types of cover generally have a greater diversity or a different faunal composition. There are no rock outcroppings at Cedar Bog. Fallen trees and loose bark are limited to forested areas. Most of the dead and fallen trees were American elm. Most of the elm were part of the swamp forest complex.

There are many species of molluscs which occur only In and around fallen logs and bark. This is most evident when the fauna of soil and litter samples is compared to the fauna of the random col­ lections. Three species were never recorded in the soil and litter samples. These were Fhilomycus carolinianus, Gastrocopta corticaria and Allogona profunda. Several other species were usually far more common or only found living in random collections. These were Strobilops aenea, Anguispira alternata, Mesodon thyrodus, Mesodon inflectus, Stenotrema barbatum and Zonitoides arboreus. The distri­ bution and density of these species is probably governed more by where and how many trees die and fall. The American elm kill will be very beneficial to several species of molluscs for years to come.

Another form of shelter significant at Cedar Bog Is leaf litter. Leaf litter does not accumulate in all the forested areas in the Bog. In the wet forested areas the leaf litter gets compacted and de­ composes rapidly. Only in the drier forested areas does this litter stay around long enough to provide adequate shelter. Some parts of the tuliptree and maple-ash-basswood swamp forest rarely have any significant litter carryover from year to year. The arbor vitae association presents a much different picture as far as litter is concerned. Here the litter decomposes very slowly and It is con­ stantly building up. The top one or two centimeters are loose and granular. This provides good shelter, but only for the smallest species. Very few large snails occur in the arbor vitae, probably because of the type of shelter found there.

The third type of shelter Is the soil itself. It is my experi­ ence that the surface consistency of the Linwood and Carlisle mucks is not uniform. This is principally due to springs and ponded areas where the muck is more compact. It Is very soft when saturated, but when it dries out it becomes quite firm. In the elm-maple, shrub and 270 silver-red maple zones the muck is usually moist, but also loose and granular. This muck would be very easy for molluscs to burrow into.

In the bog meadow and the marl meadow cover takes the form of clumps or hummocks of grasses, sedges, small shrubs and even small arbor vitae. These would be the only places the snails could get out of the constant water in the marl or peat.

Ground cover or shelter is almost impossible to plot on a map. As can be seen, water conditions, pH and soil conditions are also difficult to accurately plot on a map for an area as large as Cedar Bog. However, plants react to some of the same limiting factors as do snails. The correlations between soil and water conditions at Cedar Bog are not perfect, but they do exist. Each vegetation zone is the result of many factors. By measuring the diversity of the molluscan fauna of each vegetation zone we also reflect the uniformity and stability of that same vegetation zone. DISCUSSION OF THE DIVERSITY INDEX

In this study I did not depend on the diversity index to make comparisons between various species and various habitats. Lloyd et al. (1968: 261) states that the average species diversity index is an intrinsic parameter of a whole community. This has' never been proved by using molluscan populations. I believe Lloyd et al. are probably correct, diversity indices are intrinsic to a given community, but certainly with some qualifications when molluscs are used. The sampling procedure is very Important. In a fairly uniform area a random quadrat system probably will work well, but in an area where there are many unsampled types of habitats which contained unsampled molluscan populations, a dual quadrat system should be devised. One tailor-made to the specific habitats and the species encountered should be used. This system must be unbiased and produce results which can be reduced to the same terms as that of the soil and litter quadrat system.

Some vegetation zones had many diverse habitats which were not adequately sampled by the quadrat methods and many species lived In habitats never sampled by the soil and litter sampling procedure. Some of the species diversity indices and habitat diversity indices are not accurate. Therefore, all the species diversity Indices cannot be considered as standard intrinsic parameters. Wi.th the large soil zones they are probably very accurate.

Graph I shows the relationship of the species diversity and the corresponding species diversity index. Line A is the maximum divers­ ity index possible for the species diversity. It can be seen that the higher the species diversity becomes the further away the index and the maximum become. This shows a leveling off which will not change. Any increase in diversity will probably not increase the diversity index. Each community or habitat studied will level off after enough samples have been collected and the species diversity index intrinsic for that particular community or habitat will have been reached. At this point the species diversity Indices for all communities or habitats can be compared.

At Cedar Bog some communities can be compared because there is good evidence that they were adequately sampled. Species diversity indices for the beech-maple, bog meadow and marl meadow zones probably are close to their intrinsic values. The silver-red maple, shrub and elm-maple zones were probably adequately sampled, but they are changing. The diversity and diversity indices will also change as these communi­ ties change and become more stable. The oak-maple, maple-ash-basswood 271 272

GRAPH I

The relation between diversity index (H), based on the Shannon-Wiener equation, and the number of species (diversity) for various zonation schemes in Cedar Bog Nature Preserve, Champaign County, Ohio.

• Frederick (1974) Vegetation Zones (Total Number of Species)

° Frederick (1974) Vegetation Zones (Number of Living Species)

A Environment Control Corporation (1973) Vegetation Zones (Total Number of Species)

A Environment Control Corporation (1973) Vegetation Zones (Number of Living Species)

■ Soil Zones Modified from Ritchie et al (1971) (Total Number of Species)

O Soil Zones Modified from Ritchie ot al (1971) (Number of Living Species)

6 Environment Control Corporation (1973) Soil Zones (Total Number of Species)

O Environment Control Corporation (1973) Soil Zones (Number of Living Species)

® 193S Vegetative Cover (Total Number of Species)

0 1 9 3 9 Vegetative Cover (Number of Living Species)

* Observed Substrate at each Collection Site (Total Number of Species)

X Observed Substrate at each Collection Site (Number of Living Species)

© Cedar Bog Nature Preserve (Total Number of Species)

© Cedar Bog Nature Preserve (Number of Living Species) 273 GRAPH I

relation between diversity index (H), based on the Shannon-Wiener equation, and tho number of species (diversity) for various zonation schemes in Cedar Bog Nature Preserve, Champaign County, Ohio.

6o

60

55

50' A * 9 9 m a 45' *■» ■ a 40 ■ 9 •* • AID • A * 35-

30' •A a ) x & 25- a OA 0 3 o A D * o & 20 - Boo X . a • A 15 o o O 0 0 0 a A O 0 0 « A 10. A

5 .

0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 4.0 4 .5 5.0 5-5 Xo Shannon-Wiener Diversity Index (H) 274 and tuliptree zones need an alternative sampling procedure because of their internal habitat diversity. The arbor vitae association does not need an alternative sampling procedure, but because of the high habitat diversity present in the various cedar areas sampled I needed to collect more soil and litter samples. My data indicate that molluscs are highly aggregated in the various cedar habitats and more samples may increase the known diversity.

The average species diversity index for all species at Cedar Bog is about 4.3 and the average species diversity index for living species is about 3.95. Better sampling over the entire Bog may alter these Indices a little by the discovery of some rare populations, but I believe they can be used to compare Cedar Bog with pther areas surveyed in the same way. THE MOLLUSCA OF THE ARBOR VITAE ASSOCIATION

One of the unique vegetation zones at Cedar Bog is the arbor vitae association. The natural range of arbor vitae (Thuja occidentalis L.) is nearly all further north. It is found in bog or swamp conditions from about 43° north latitude to the southern part of James Bay in Canada. This places its southern boundary range all along Lake Michigan in Wisconsin, about the middle of the lower penin­ sula of Michigan, southern Ontario and the middle of New York state. One of the objectives of this study was to determine whether any of the molluscs found at Cedar Bog were relics or disjuncts. I had to first find out what molluscs live in the arbor vitae, bog meadow and marl meadow in their natural range. A thorough Investigation would have involved studying the molluscs in all three of these disjunct plant associations. However, the easiest association to find in its natural range Is the arbor vitae association. It is also the only one of the three in which there are records available from the literature.

I made collections In ten different counties in the lower peninsula of Michigan, five different counties in the upper peninsula of Michigan, one county in Wisconsin and six different localities in Ontario, Canada. Table 18 compares the different species of molluscs collected by myself and various other investigators in the arbor vitae association in Ohio, Michigan, Wisconsin and Canada. The data obtained from the literature does not specify whether each species was collected dead or alive, so I just indicated Its presence. In those collections which I have indicated whether living specimens were collected and how many localities it was collected in. A total of 77 species were collected In all of the arbor vitae forests surveyed. Fourteen species were found only at Cedar Bog, Ohio and 33 species were not found at Cedar Bog, Ohio.

The following list of molluscs represent what are the most common inhabitants of the arbor vitae association in its natural range; Carychium exile, Carychium exiguum (Harry, 1951 did not recog­ nize £. exile as a separate species), Planogyra asteriscus, Columella edentula, Strobilops labyrinthica, Anguispira alternata, Helicodiscus parallelus. Puneturn minutissimum, Discus cronkhltel catskillensis, Nesovitrea binneyana, Euconulus fulvus. Striatura milium, Striatura exigua, Striatura ferrea and Zonltoides arboreus. Only Planogyra asteriscus, Discus cronkhitei catskillensis ajid Striatura ferrea were not in the Cedar- Bog, Ohio arbor vitae association. Anguispira

275 TABLE 13

llallusca recorded fron the arbor vitae association in Ohio, Michigan, Wisconsin and Ontario*

Oiiio Lcwfcr Peninsula of richigan Upper Peninsula of Michigan Wisconsin Ontario

Xoferl Keferl Harry (1351) Jewel and Arclier (133^) 3ak«r (1322) Walker and Buthvon ( 1906) Keferi Kefcrl Koferl Brom (1329}

Collections Collections Collections Chacpaigrt Co. Tan Cheboygan Co. Essot Co. Cheboygan Co. Cheboygan Co. Dickinson Co. Chtoragcn Co. Keweenaw Co, Nine Marinette Co* Six Cedar Sog L o ca lities (Bog Pools) I s le Rpyale L o ca lities L o ca litie s

- - --* ------V5iva*a sine^ra X 0 4------.■ - - - — Pc*vtict.-,i* iasidaria A -_ ------C^vn --a sr :isa r ir a ta A - - - «------L>r-'’.ca uwr::Iis A -- • ------L v - —- J s U i X _* —_ •-• • Lv ~ jzi "pp* ..------0(2) D D (l) Gyrli.iiis GSr.US 0 0(1) - - X- - -- -" G-'T-jiuiJS •:Lf*7.i.“stria tu s -- X X • ------GvraulilS ^ fl-’CTJS - - - - X ------“ Gvrr.;lu‘ '.ir-utos —_-- X - - _ - -- P ij-.-r r jli j-rrlfj^ra - - - X X - X -- - - 0 ( 1 ) Pn,-?^ ir .tc v a D ------Fnyra rr;.'i:'.i - -- - X - X - --- hvorort-i ** XXX • • p. -- 0 - Car-.criiw c m I l: A A (0 ) - • -- X -- A (2 ) A A l l ) Carvchi'n e-»icj^Ln A A (5 ) X X* XX - m - A A ( l) Carvcni.j* nar.nodo*: A » _ •• --- » ** CoitiiioccM iuL^ica • 0 ( 4 ) _ X ••• 0(2) 0 _ VoIIrria oe'rtrici • _ - p. - p. - 4 (1 ) • - Plart'Twra ir.i*-/ iscJS - A (6 ) X -- X - X - A {4 ) - A (3 ) rjroa - -4 -- •--X A ( l ) - - Pu*:3i i

Ca^irDrcri’u co rtica ri* ------0 ( 1 ) Vertv*sc r*cr«:r:i. 0 --- ** - •» --- V ertico ovi-.ti D _ X X p. • •* . X? - ~ - Vcr-icro ov'T^sa « 0 ( 3 ) - w - w - - m 0(1) A m Vertioo tridental* D M• .- • - - 0(2) - 0(2) Vertico oouldi A A(2) X A ( l ) A A (3 ) 276 TABLE 18— Continued

O hio Lower Peninsula of Michigan Upper Peninsula of Michigan Wisconsin Ontario

Koferl Kafcrl Harry (1951) Jewel and Archer (1 9 3 6 ) Baker (1322) Walkor and Ruthven (1906) Keferl Kcfcrl Koferl Brown ( 1J2 3 )

Collections Collections Collections Champaign Co. Ten Cheboygan Co. Ecaet Co. Cheboygan Co. Cheboygan Co. Dickinson Co. Ontonagon Co. Keweenaw Co. Nino Marinette Co. S i x Codar Bog L o c a l i t i e s (Bog Pools) Islo Royals Localities Localities

Vcrtino riliui A 0 ( 2 ) •--- * to • 0 ( 1 ) A * r v la r d r r i - - -— * * - *• to - A ( l ) Cell.".. I 1 a c/2 -_r.tula A A (8 ) X X - - - X- A (3 ) A 0 ( 2 ) Strorilevs aarca to - ■» -*- * to - to Str*cyilcr;> lafcyrinthica A A (1 0 ) XX - - XX* A (6 ) A A ( l ) Sucrir.ca ovalia 0 ( 1 ) - X -- to to to — • Succireidao 0 ( 2 ) ------A (2 ) - 0 ( 1 ) C*vlera sro* A ----- X to - - -- Fiiiicrvc;j5 .larelinicnus A - to • * to to * - - - Paiiif-ra dcrcilis A ( l ) X to - to to *- - A ( l ) Pallitcr.i K .-cnilli -- to > •« to to X A ( l ) -- to Ar.n-:i:cirj il\jrr.ata. A A {3 ) XX - XX to M 2 ) 0 0 ( 1 ) HcliL'Cdiscjs t^rallelus A A {6) X to > XX A (4 ) 0 A (3 ) Py^ctu': rirruti r titji A A (1 0 ) XX to to X * A (7 ) A A (2 ) > 0; "cu- crcrxhit«i» A (0 ) X X to to X A {6) A A (5 ) Ccrcccras liiive A A (3 ) to -- > X to X A (2 ) to to O^^cc^ra^ r^^ictilatua . to • - to to to - - A ( l ) Aricfl ta*,2»a*u5 A ( l ) --- - to - to a ( 3 ) - ?^rnvitr«a birrr.rvcuia A A(flJ XX • to to to X A (7 ) A A (4 ) - to Letcvitrca clcatriaa A A (5 ) --- to to - D A (2 ) Gtvtr'valir’ia indontata A D (4) - . to X to A ( l ) •• - - to to to - > paravilrc*! r’jltiacntata a ( 3 ) to • Hawaiia -linyieuia A A ( l ) -- - - to to to -- * Eyacrjltn chcrsinua • to - * - X to to - to to Euctrvjlus. fu!vus A A (9 ) XX - to to to A (7 ) A A (2 ) ZcrEtoiJcs arborcus A A (6 ) - X -- XXX A (5 ) A A (6 ) Gucova ctcrkii A • — - to to to - * 0 ( 1 ) Striatura riliua A A la ) X - . to X to A (7 ) A A (3 ) Striaty-a vxi

Ohio Lower Peninsula o f Michigan Upper Peninsula c f Michigan Wisconsin Ontario

Kofcrl Kcferl Harry (1551) Jewel and Archer {193&) Baker (1322) Walker and Ruthven ( 1306) Knforl Keferl Keferl Brown (1329)

Collections Collections Collections Cbaspaign Co. Ter. Cheboygan Co. E&aat Co. Cheboygan Co. Cheboygan Co. Dickinson Co. Ontonagon Co. Keweenaw Co. Nina Marinette Co. Six Cedar Bog L o ca lities (Beg Pools) Isle Rqyale Localities L o ca litie s

Mesodon th/roldus A — _ m Ikscdcti zal*'tu". - -• - - - -- A{1) - - Mssaiin ^var.io • A (l) - - - X ------Triodcrs:' altslocris • A (l) XX - X --- D{Z) -- Sc-'-aorit.-? c c c id ^ ta io - - XX ------oartu^eija ------X -“- -- Pisidiua cascrUn:s A ------* Pisidit:a cororcssi^j - •-- -- D - Pisidiuft spp. —"- — X — ■ X D(2) “

— Absent, X Present, but it is not known whether the species was found dead or alive, A Collected living. 0 Collected dead only. {) Nwber of localities collected in. 279 alternata and Zonitoides arboreus are more common in the northern arbor vitae forests, probably because there are more fallen logs pres­ ent, particularly birches (B e t u l a ) . These birches rot quickly and provide very good habitat for the large species in an arbor vitae forest.

Planogyra asteriscus, Nesovitrea binneyana, Discus cronkhitei catskillensis and Straitura exigua have northern distributions and Ohio would be considered the southern edge of their natural range. Only Nesovitrea binneyana and Striatura exigua are found at Cedar Bog, Ohio. These snails are also found in other localities in Ohio and I do not believe they should be considered relics or disjunct species until extensive knowledge concerning their distribution in Ohio is known. Vertigo nylanderi, Vltrina limpida, Planogyra asteriscus, Zoogenetes harpa, Vertigo morsel and Discus cronkhitei catskillensis might be considered northern disjuncts if they were found at Cedar Bog, Ohio, but only Vertigo morsel is. It is actually a typical marl meadow species, not an arbor vitae species. It has never been recorded from any arbor vitae forest other than at Cedar Bog, Ohio.

Carychium nannodes is most commonly associated with the arbor vitae association at Cedar Bog. This species has a southern distri­ bution (Map LXVIII) and it might be considered a southern disjunct. The only other species found at Cedar Bog which appear to be on the edge of its natural range is Vertigo pygmaea. It has an eastern distribution. Before any final conclusion can be made concerning any disjunct or relic species, better distributional data for these species must be obtained. All these species may be far more common in Ohio than our present knowledge indicates. 280 HAP LXVIII

Distribution of Carycliium nannodos in the United States. 281 MAP LXIX

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