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Master's Theses Graduate College

4-1979

Postglacial Environmental History of a Marl Lake Site in Kalamazoo County, Southwestern Michigan

Valerie A. Naeve

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Recommended Citation Naeve, Valerie A., "Postglacial Environmental History of a Marl Lake Site in Kalamazoo County, Southwestern Michigan" (1979). Master's Theses. 2043. https://scholarworks.wmich.edu/masters_theses/2043

This Masters Thesis-Open Access is brought to you for free and open access by the Graduate College at ScholarWorks at WMU. It has been accepted for inclusion in Master's Theses by an authorized administrator of ScholarWorks at WMU. For more information, please contact [email protected]. POSTGLACIAL ENVIRONMENTAL HISTORY OF A MARL LAKE SITE IN KALAMAZOO COUNTY, SOUTHWESTERN MICHIGAN

by

Valerie A. Naeve

A Thesis Submitted to the Faculty of The Graduate College in partial fulfillment of the Degree of Master of Science

Western Michigan University Kalamazoo, Michigan April, 1979

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. POSTGLACIAL ENVIRONMENTAL HISTORY OF A MARL LAKE SITE IN KALAMAZOO COUNTY, SOUTHWESTERN MICHIGAN

Valerie A. Naeve

Western Michigan University, 1979

The depositional and ecological history of Oakland Bog, a relic

Wisconsinan (16,500+300 years B.P.) lake site is interpreted from

pollen stratigraphy, molluscan faunal assemblages and sediment facies.

The lake basin, which probably formed after the melting of a late

Wisconsinan buried ice block in the Kalamazoo Moraine Complex, is

irregular, steep-sided and floored by glacial till. The till is

overlain by as much as 7.6 meters of lacustrine marls which extend 2 over an area of nearly 0.5 km . Basal marls are characteristically

medium gray, clayey, and contain abundant calcified stem fragments

of charophytes; however, the overlying marls display alternating

pale carbonate-rich and darker organic-rich laminae separated by

massive or graded unlaminated beds.

The molluscan faunal assemblages occur in vertically restricted

associations that probably reflect changing environmental conditions

influencing the local habitat. The oldest molluscan assemblage is

interpreted to record a pioneering stage because it is characterized

by low species diversity and low intra-specific frequency. The

pioneering stage was associated with a creek or intermittent ponded

setting that was rapidly transformed to a relatively silt-free, per­

manent waterbody with a soft substrate and considerable vegetation

in the shallow areas.

ii

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. There is no evidence in either the pollen record or the mollus­

can assemblages to indicate any reversals in the general trend of

climatic warming. Throughout late-glacial and early post-glacial time,

the climate of southwestern Michigan appears to have been cool and

relatively continental with no clear record of temperature oscil­

lations correlative with ice margin movements. The tripartite nature

of the pollen record strongly resembles that of other Great Lake

region pollen stratigraphies. Differential migration rates may help

to explain the basic pattern of postglacial vegetation succession.

Ecotones of mollusks and vegetation appear to have a south to north

zonation which is time transgressive.

i i i

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NAEVE, VALERIE ADRIENNE POSTGLACIAL ENVIRONMENTAL HISTORY OF A MARL LAKE SITE IN KALAMAZOO COUNTY, SOUTHWESTERN MICHIGAN.

WESTERN MICHIGAN UNIVERSITY, M.S., 1979

University . M iapnlm s International 300N. ZEES ROAO. ANN ARBOR. Ml 48106

@ 1979

VALERIE ADRIENNE NAEVE

ALL RIGHTS RESERVED

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Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ACKNOWLEDGEMENTS

I would like to thank the many people whose efforts made

possible this study. I am indebted to Dr. J. Terasmae (Brock

University) who not only sponsored the radiocarbon dates at Brock

University Geochronology Laboratory but also provided instruction

in techniques of quantitative pollen analysis and permitted me to

use the facilities at Brock University. I would like to extend

my sincere appreciation to Dr. L. Kalas (Canada Centre for Inland

Waters, CCIW) for his time and efforts with the molluscan research

and making available to me unpublished data collected by the CCIW

on molluscan ecology. Much of my understanding of palynology stems

from the assistance of Mr. Chris McAtee (Brock University graduate),

to whom I am most grateful. I would like to extend my thanks to

Dr. W. Benninghoff (University of Michigan) and Dr. R. Kapp (Alma

College) who dedicated many hours of assistance helping me to recog­

nize palynomorphs and to Dr. W. Bailey (Central Michigan University)

for the unpublished pollen date from Wintergreen Lake. I am most

grateful to Dr. B. Wilkinson (University of Michigan) for his tech­

nical assistance and insightful discussions on marl depositional

environments.

A special thanks to Bill French who not only built most of the

coring equipment but also spent many hours in the field helping to

collect samples. I would also like to extend my thanks to the many

hands and strong backs, without whose help land and water sampling

iv

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. might not have been possible: Bill Dennison, Steve Tripp, Nat

Fuller, Ron Parker, Jeff Reinhart, Norman Lovan, Richard Oxhandler,

Tom Laux, Greg Paulx, Mai H., Ron Penekamp, Kevin Kincare, Robert

Ramsey, Chuck Denien, Jim Hahnenburg, Craig Payne and Maggie

Prechincki. I would also like to acknowledge the support of the

geology department staff (Western Michigan University) - Dr. W.

Harrison (my advisor), for his time, patience and guidance through

the periods of research and writing; Dr. W.D. Kuenzi, for his in­

valuable assistance both in the field and technical counseling;

Dr. W.T. Straw, for his advice and field assistance; and Dr. R.

Passero, for helping to edit the final copy. This work was supported

in part by a Graduate Research Grant from Western Michigan University.

Many thanks to Lynda Vallier for typing and Douglas Dillon for

assisting in proofreading the manuscript.

This manuscript is dedicated in the memory of Mr. Frank Allen,

head librarian at the Physical Sciences Library, Western Michigan

University whose efforts and interest were gratefully appreciated.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. TABLE OF CONTENTS

PAGE

I INTRODUCTION...... 1

The Study Site...... 3

Location and general description...... 3

Pleistocene geology of southern Michigan. . . . 5

Recent land survey...... 9

Origin of m a r l ...... 10

Sedimentary deposits in Oakland B o g ...... 15

Historical Reviews of Palynological and Molluscan Research...... 18

History of palynology ...... 18

Previous palynological investigations in the m i d w e s t ...... 19

Previous work with post-glacial mollusks. . . . 20

II METHODS AND MATERIALS ...... 24

F i e l d w o r k ...... 24

The Laboratory...... 28

Subsampling for pollen analysis...... 28

Palynormorph identification and counting. . . . 30

Molluscan analysis...... 31

Radiocarbon Dating...... 32

III NATURE OF THE SEDIMENT...... 34

Discussion of Marl Deposition ...... 40

Sedimentation Rates ...... 45

vi

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IV MOLLUSCA RESULTS AND DISCUSSION...... 49

Paleoecological Reconstructions with Freshwater M o l l u s c a ...... 49

Oakland Bog Fossil Molluscan Description ...... 52

Discussion of Molluscan Stages ...... 57

V POLLEN STRATIGRAPHY...... 66

Principles of Palynology ...... 66

Palynomorphic data presentation and interpretation ...... 66

Statistical methods for pollen analysis...... 71

Construction of the pollen diagrams...... 75

The pollen zone...... 76

Pollen results ...... 77

Zone 1_ (interval 381cm to 2 7 5 c m ) ...... 78

Zone II (interval 275cm to 180cm)...... 79

Zone III (interval 180cm to 41cm)...... 80

Distribution of additional palynomorphs...... 81

Discussion of Pollen Zones ...... 82

Zone 1^ sprue e-herb z o n e ...... 82

Zone II the pine pollen z o n e ...... 100

Zone III the deciduous hardwood z o n e ...... 109

VI SUMMARY AND CONCLUSIONS...... 118

VII RECOMMENDATIONS...... 123

VIII REFERENCES CITED ...... 124

vii

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LIST OF FIGURES Page

Figure 1. Topographic map showing the study areas and the surrounding terrain...... 4

Figure 2A. Map showing the surfical geology of Kalamazoo County and location of Oakland Bog ...... 6

Figure 2B. Cross-section A-A* (from Figure 2A)...... 7

Figure 3. Map of moraines in the study area...... 8

Figure 4. Site map showing the location of pollen cores in the Great Lakes region ...... 21

Figure 5. Map of sampling sites at Oakland B o g ...... 25

Figure 6A. Calcified cortical tube casts from charophytes . . . 35

Figure 6B. Laminated marl displaying alternating light and dark laminae separated by massive shelly layers. . .36

Figure 6C. Thread or root-like plant debris and shells in m a r l ...... 37

Figure 6D. Nitella reproducing bodies in marl ...... 38

Figure 6E. Peat fragments in marl ...... 39

Figure 7. Cross-section (from x to x', Fig. 5) of Oakland Bog based on core s a m p l e s ...... 41

Figure 8. Sequential reconstruction of site II based on sediment facies and molluscan assemblage ...... ^4

Figure 9. Mean sedimentation rates of marl accumulation at Oakland Bog plotted against time and pollen zones. . 48

Figure 10. Molluscan habitats of a typical marl lake...... 51

Figure 11. Freshwater molluscan diversity curves and total populations from site II, Oakland Bog...... 53

Figure 12. The transition of stages of molluscan assemblages and relative water depth recorded at site II .... 56

Figure 13. Molluscan stages and lake history of Oakland Bog as inferred from Molluscan assemblages...... 59

with permission of the copyright owner. Further reproduction prohibited without permission. Page

Figure 14. Two suggested environmental changes occurring during the Second Enrichment Stage...... 61

Figure 15. Reconstruction of full-glacial vegetation .... 93

Figure 16. Holocone migrations of ecotones ...... 103

Figure 17. Index map of sites in Minnesota where postglacial pollen studies have been conducted...... 104

Figure 18. Summary chart: pollen vegetational reconstruc­ tion, lake history, climate, molluscan community in relative time perspective...... 117

ix

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LIST OF TABLES Page

Table 1. Principal theories of the origin of freshwater marls...... 14-

Table 2. Study site sampling thickness of auger recon­ naissance...... 26

Table 3. Pollen extraction procedure...... 29

Table 4. Radiocarbon dates from Oakland B o g ...... 33

Table 5. Sedimentation rates for site I based on radio­ carbon dates and pollen stratigraphic marker h o r i z o n s ...... 46

Table 6. Environmental evolution of lake at Oakland Bog • . 54

Table 7. Oak pollen grain size...... 80

Table 8. Modern pollen rain data from Canada...... 86

Table 9. Correlation table of pollen stratigraphic sub­ zones of the deciduous hardwood zone III across the northeast...... 110

LIST OF PLATES

Plate 1. Molluscan absolute diagram of Oakland Bog, site I I ...... 139

Plate 2. Relative pollen diagram of Oakland Bog, site I . .

Plate 3. Pollen concentration diagram ...... 3^3

Plate 4. Pollen influx diagram...... l^3

APPENDIX

Appendix 1. Sediment log and description ......

Appendix 2. Molluscan environmental table...... ^33

Appendix 3. Molluscan habitat survey ...... 333

x

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. INTRODUCTION

The climate and geologic history of Oakland Bog, a relic

Wisconsinan marl lake site in southwestern Michigan, can be re­

constructed from pollen stratigraphy, molluscan faunal assemblages

and sedimentary facies.

Late-glacial and early post-glacial pollen sequences are use­

ful for interpreting the ecology and dynamics of forest succession

under the influence of a changing climate. Analysis of the pollen

stratigraphy in the marl sequence from Oakland Bog provides insight

into the following questions:

1. Was the post-glacial climatic cycle of sufficient intensity

and duration to cause changes in the vegetation at or near

the study site?

2. Did tundra conditions exist along the glacial ice margin?

3. Can vegetational changes be correlated with climatic fluc­

tuations such as those involved with ice-margin correla­

tions?

4. What influenced the major transition of vegetational

assemblages at the study site and how does the local site

fit into the regional herb-spruce to pine to oak-deciduous

forest succession?

Local environmental conditions recorded by marl sequences can

also be interpreted from certain species of freshwater mollusks which

are sensitive to physical and chemical factors in their environment.

As a paleoecological tool, mollusks are useful indices of substrate,

1

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 2

plant growth, sedimentation rate, water depth, chemistry and temper­

ature. Vegetation-depth zones in postglacial lakes can also be

delineated by fossil mollusk assemblages (Camp, 1974). The geographic

distribution of mollusks today, as compared to their fossil forms,

records migration patterns which were sensitive to climatic change

(Clarke, 1974). These correlations are possible because many of

the taxa preserved as fossils in the postglacial sediments are still

living and under study today (LaRocque, 1968).

The sedimentary history of a lake is dependent on local envir­

onmental conditions (Brunskill, 1969 and Camp, 1974). The color,

texture, laminations, and enclosed macrofossils of the marl sequence

at Oakland Bog can be used to reconstruct depositional environments

and to interpret the developmental history of the lake.

Comparison of the pollen stratigraphy, molluscan faunal assem­

blages and sedimentary facies of Oakland Bog with other studies in

the Great Lakes provides some insight into the regional vegetation

and climatic history. This results in a new approach to paleoecol-

ogical investigations of Quaternary deposits. Ultimately this

information will be added to the storehouse of data by which a

better understanding of environmental dynamics can be achieved. This

furthers the possibilities of forecasting environmental change.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 3

The Study Site

Location and general description

Oakland Bog, located in Kalamazoo County, Portage Township;

NW k, section 20, T3S, R11W; 42011' N, 85°35' W, was located from

marl exposures along the drainage ditches of Oakland Bog (Fig. 1).

Infrared photographs and topographic maps do not clearly reveal the

"boggy" nature of the site; however, they do show marl excavation pits

with the water table near or at ground level. Although no evidence

of a stream is apparent at present, a study of older topographic maps

of the area shows that Oakland Bog was once part of the Portage Creek

drainage system which has its headwaters in Hampton Lake, west of

the study site. The original drainage of Oakland Bog has been sub­

sequently modified by artificial drainage ditches.

The lake basin, which probably formed after the melting of a

late Wisconsinan ice block in the Kalamazoo Moraine Complex, is

irregular, steep-sided and floored by glacial till (Fig. 1). The

basal molluscan fauna of Oakland Bog indicate that running water was

first associated with the basin before the transition to a permanent

lake. As much as 7.6 meters of lacustrine marl overlie the till, 2 extending over an area of nearly 0.5km . The bog is surrounded by

sandy knobs of an outwash plain formed from the meltwaters of the

Lake Michigan and Saginaw Ice Lobes. A thin veneer of till was found

overlying the outwash just to the east of the bog in excavations made

for a housing development. This veneer of till is quite different

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Kalamazoo County

' x m Portage 1 Township <3 ■ Section 20 Location of Oakland Bog

Cehtce stre et <; K

&

v'clh.

jlL

Figure 1. Topographic map showing the study area (outlined) and the surrounding terrain. Area underlain by marl is cross-hatched and waterfilled marl pits are indicated in black.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 5

from the till underlying the marl. The upper till is more red in

color, friable and contains large angular fragments; whereas, the

lower till is a densely compacted sandy till with a gray clay matrix.

Pleistocene geology of southern Michigan

The earliest comprehensive study of the Pleistocene geology of

Michigan was done by Leverette and Taylor (1915). More recent

accounts of the glacial features and Pleistocene geologic history

of Michigan have been written by Wayne and Zumberge (1965) and Shah

(1969). Regional surficial geologic maps have been prepared by

Leverette and Taylor (1915), Martin (1959) and Flint et al (1959).

The glacial geology in Kalamazoo County, southwestern Michigan was

remapped in 1968 by Shah (Fig. 2).

A distinctive feature of these surficial maps are the many small

lakes of glacial origin in southern Michigan. Many of these lakes

have resulted from the hummocky nature of the end moraine while others

began as ice block depressions on the outwash plains.

The study area is located in the interlobate region of Lake

Michigan Lobe and Saginaw Lobe ice (Loven, 1977) (Fig. 3). The re­

treat of the two ice lobes in Kalamazoo County, left the study area

free of ice between 16,000 and 15,000 years B.P. Opinions differ

however, as to when all of southern Michigan was first relieved of

the ice burden. Kelly and Farrand (1967) show the region as com­

pletely exposed 16,000 years ago. Wilson (1967) reported that

southern Michigan was free of the ice sheet about 15,000 years ago,

while Cleland (1966) stated, "at about 12,000 years B.P., the retreat

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 6

O a l l u v iu m © 0UTWA8H CLAY ON OUTWASH THIN TILL ON OUTWASH TILL, LOW RELIEF T IL L , HUMMOCKY

Figure 2. Map of surficial geology in Kalamazoo County (modified after Passero, 1978), Portage township cornered, X shows location of Oakland Bog. Cross-section A-A' depicted in Figure 2B.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. •vj Outwash Sand Sand lens

CO 3 < H Jk z < ui * ____ O*

* a- »- < H Z 5 o z UI a UI CO o _L— ^■?*j _ _ . ^j ^ ■..y • Till Clay ^ • 6 • 6

Z Z < 1 ° 5 * O £ N < 0 = COLDWATER SHALE -I < 2 o < a. z 5

h >- z < o z No O3 < o Alluvium I _l _J ui Z o < 3 Z >- H- 8

4 s & ui t- < O > CO CO -J 3E 0

Bedrock Till Till —

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:

/ ice margin SAGINAW LOBE ERIE LOBE Figuie B 3 \") V/,:) V/,:) MICHIGAN LOBE Figure Figure 3 A c u e

a b a t t l e COLDWATER \ < wl wl -iwnk -iwnk I Bog Oakland

£ \ ’* •n i\ x ’/ V Figure 3. A. Great Lakes region during the Valparaiso and Fort Wayne phases of the Cary : v . M - Glacial Substage, showing distribution of ice (cross-hatching). B. Moraines of southwestern Michigan in the study region (black box in 3A). * *

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 9

of the Cary Substage of the Wisconsinan Glaciation opened up the

southern part of the Great Lakes region to plant and animal coloni­

zation." Based on radiocarbon dates, pollen stratigraphy and mollus­

can faunal assemblages, this study demonstrates that the region of

investigation must have been ice-free between 15,000 and 16,000 years

B.P. By 12,000 years ago, the ice retreat had exposed most of the

Great Lakes with the exception of the northeastern-most part of Lake

Superior and the northern, or Georgian Bay, part of Lake Huron

(Terasmae, 1967).

Approximately 11,500 years ago, a glacial readvance, termed the

Two Rivers Ice Advance (Evenson et al., 1976), put ice well south in

the Michigan basin, over Lake Superior and into the northern part of

Lake Huron. With the retreat of the Two Rivers Ice, all of lower

Michigan became ice-free (Evenson et al., 1976). And some time be­

tween 9,500 and 10,000 years ago the Great Lakes region became com­

pletely ice-free (Door and Eschmann, 1971). The changes in ice

position and water levels in the Great Lakes were major factors in­

fluencing the opportunities for and colonization by plants and

animals alike.

Recent land survey

The earliest land surveys show that pre-settlement plant com­

munities in Kalamazoo County and southwestern Michigan were forests

of oak, maple, ash and walnut surrounding isolated islands of prairie-

grassland and oak savanna (Durant, 1880; Kenoyer, 1930, and 1934;

and Peter, 1970).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 10

Today, well-drained soils in southern Michigan support oak-

history forests while beech-maple forests tend to dominate the more

clay-rich soils. Most of this deciduous forest region is presently

used as farmland, fields, meadows and occasional woodlot. Farther

to the north, in the Lower Peninsula, forests on sandy soils are rich

in pine, but where there has been extensive logging or burning,

poplar and birch are common. Fine-grained till soils in the upper

Lower Peninsula and Upper Peninsula support an assemblage of beech,

birch, maple and hemlock better known as the "Northern Hardwoods."

Surrounding Oakland Bog today are stands of oak, hickory and

maple. The bog itself has previously been used to grow beans and

dredged for marl to supply lime for local farm fields. A golf course

occupies the southern part of the site, Gourdneck State Preserve is

to the east, and housing developments are located on the hills north

and west of the bog.

Origin of marl

The origin of freshwater marls, a fine-grained loosely consoli­

dated material composed mainly of calcium carbonate, has been a

perplexing problem since their recognition before the turn of the

century. The impetus to study the marls in Michigan and Indiana came

from the Portland Cement (Manufacturing) Company. A few studies were

published during the early 1900's when the cement industry searched

for suitable marl deposits to mine. Some of the comprehensive papers

resulting from marl exploitation include: Davis, 1900, 1901; Blatchely

and Ashley, 1900; Hale, 1903; Lane, 1903; and Russel, 1901.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 11

Carbonate deposition in freshwater lakes is known to occur:

1) in relatively shallow lakes in arid and semi-arid settings, where

carbonate production is the result of evaporative processes; and

2) in lakes occurring in glaciated regions under temperate climatic

conditions. Temperate region marl lakes will be examined in more

detail below because Oakland Bog was once the site of such a lake.

Modern temperate marl lakes are restricted in their distribu­

tion by geologic and climatic controls. Carbonate deposition in

this system requires: 1) a source of carbonate from the surrounding

drainage basin and 2) a seasonal variation in lake temperature.

Hardwater marl lakes are common to portions of the glaciated

midwest particularly southern Michigan and northern Indiana (Wetzel,

1973). Carbonate deposits can be extensive, forming thick marl

banks that extend some distance outward from the shoreline before

abruptly dropping off to the central flat bottom of the basin

(Wetzel, 1973 and Wilkinson, personal communication). Most of the

marl lakes in southern Michigan lie within areas of highly calcerous

till. The carbonate-rich glacial drift around Oakland Bog apparently

served as a suitable and plentiful source of carbonate material for

the Oakland Bog marl.

Evidence suggests that surface and ground water flowing through

glacial drift dissolve carbonate ions from the till reservoir and

supply the freshwater lake with suitable ions for marl production

(Hale, 1903; Brunskill, 1969; and Wetzel, 1973). Hale (1903) re­

ported that cold lime-rich waters entering lakes by way of springs

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did not release calcium carbonate until the water was warmed by

overturning. Thermal stratification is also considered to be an

important process for the development of marl lakes in temperate

glaciated regions (Kindle, 1927; and Brunskill, 1968).

Little modern research has been directed towards the relative

importance of the processes leading to marl accumulation. White

and Wetzel (1975) and Wetzel (1970, 1975) have argued that the

photosynthetic activities of cyanophytes, charophytes and phyto­

plankton are primarily responsible for spring and summer marl

deposition. On the other hand, Terlecky (1974) and Brunskill and

Lundam (1969) have shown that physio-chemical processes such as

direct degassing of carbon dioxide as the surface waters warm are

dominant in several New York lakes.

Largely on the basis of the work of Minder (1926) and Nipkow

(1920, 1927), some paleolimnologists have called upon the seasonal

cycle of phytoplankton production and temperature to account for

autocthonous laminations of calcium carbonate and organic matter

in both recent and ancient lake sediments (Deevey, 1953 and Brunskill,

1969), but the mechanism and processes of precipitation and sedimen­

tation have not been documented with sufficient detail to permit

paleolimnologic interpretation.

The study of the physical chemistry of calcium and carbon

dioxide in sea water provides a fruitful approach to the problems

of calcium carbonate precipitation in the marine environment, but

these techniques have not been widely applied to the question of

"seasonal deposition" of calcium sediments in freshwater.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 13

Brunskill (1969) investigated the activities of calcium and

carbonate in Fayetteville Green Lake, a freshwater marl lake in

New York and estimated seasonal values of such variables as: 1)

dissolved carbon dioxide chemistry, 2) activity products of calcium

and bicarbonate ions, 3) suspended crystalline calcium carbonate in

the water-column, and 4) total sedimentation rain. Examination

of marl deposits in Fayetteville Green Lake by Takahashi and others

(1968) and Brunskill (1969) led to the conclusion that the bulk of

the marl deposits were precipitated physio-chemically in the

epilimnion. On the other hand, Wetzel and his workers (1969, 1973),

investigating the factors related to productivity in marl lakes in

Michigan and Indiana considered biogenic contributions or regula­

tions as primary factors in the production of marl.

Whether the production of marl in the hardwater lakes of the

temperate, glaciated northeast is a result of physio-chemical

processes, biogenic processes or a combination of both, the forma­

tion of varved laminations is restricted to either the hypolimnion

in dimictic lakes or the monolimnion inmeromictic lakes (Brunskill,

1969; Wetzel, 1970; and Williams, 1974).

Fine-grained marl deposits with little organic matter are

thought to characterize an open, deep-water environment of

deposition; whereas, the presence of increasing amounts of organic

matter and coarsening texture point to a marginal marsh environment

(Terlecky, 1974). Massive, unlaminated beds have been observed

forming from rapid deposition associated with turbidity currents

(Terlecky, 1974). These beds commonly display scour marks and

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graded bedding. Other massive beds reported from shallow terraces

where waters are often turbid are characterized by root or root­

like plant structures and a prevalence of shallow-water mollusks.

Brunskill and Lundlam (1969) have found that marl does not accumu­

late to any great extent in water depths greater than 9 meters,

and any marls collected in waters deeper than the chemocline do not

show horizontal stratification.

No one theory for the origin of marl accounts fully for all

marls. Most marls appear to be a mixture of components from several

origins. Restrictions proposed by the various theories for the

origin of marl are useful in interpreting ancient systems and may

eventually lead to a better understanding of paleoenvironments and

tolerances of flora and fauna in past and present temperate hard­

water lake systems.

Table 1. Principal theories of the origin of freshwater marls.

Theories Studies

Physio-chemical precipitation Blatchley and Ashely, 1901 Theil, 1933 Brunskill, 1968, 1969 Takahashi et al., 1968 Terlecky, 1974

Biochemical precipitation Davis, 1900, 1901

Goldich et al.,•» 1959 Wetzel, 1972

Diagenesis Twenhofel, 1933

Mechanical sedimentation Theil, 1933

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Sedimentary deposits in Oakland Bog

The surface of the bog is covered by approximately one meter of

peat and underlain by as much as 7.6 meters of marl. Since paly-

nomorphs and mollusk shells were poorly preserved in the past, this

study concentrated on the marl sequence.

Based on the preceding research, it seems reasonable that the

mechanism for the production of sltsmsting light and dsxk tsarl

couplets at Oakland Bog required: 1) that hydrologic and chemical

budgets be maintained; 2) a climate that provided seasonal thermal

stratification; and 3) windwork not to disrupt meromictic stability.

The laminations and variations in color and texture displayed by

the marl sequences at Oakland Bog reflect variations in depositional

conditions; however, some controversy exists over the interpretation

of these features.

The marl deposits at Oakland Bog, as well as other marl deposits

in the midwest, appear to have been localized on shallow littoral

banks and lake mounts (Wetzel, 1970). Around the perimeter and

shallow areas of the lake, rich growths of lime-tolerant plants

(Chara and Potamogeton, in particular) seemed to be major contributors

to littoral marl through photosynthetic activities that produced

calcareous coatings on the plant.

The accumulation of marl in the deeper parts of the lake basin

(below the photic zone) appeared to have occurred at a slower rate.

This was probably a function of the physio-chemical regime rather than

biogenic factors. Evidence of littoral molluscan fauna and organic

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 17

debris also indicate that some of the marl was produced in shallow

areas and washed into deeper waters.

In general, the marl units at Oakland Bog record the progres­

sive progradation of carbonate banks filling the lake. Gray to

dark gray, fine grained marls, containing little organic matter but

often rich in clay are commonly associated with cold or deep-water

molluscan fauna. The marls progressively coarsen upward, increasing

in organics and shallow water molluscan fauna. The light brown or

buff color marls occurring in shallow waters today are comparable

to the fossil marls at Oakland Bog. The shallow water marls contain

abundant shells and often have root-like plant material preserved in

the sediment.

Fossiliferous marls at Oakland Bog are similar to the marls

Wetzel (1970) has described forming on shallow terraces. Mollusk

shells form the purest marl units at Oakland Bog but in general,

mollusks compose only a minor constituent of most marls (Wetzel,

1973). The occurrence of mollusks and their plentiful growth depend

upon many of the same factors that are responsible for the formation

of marl (Wetzel, 1973); however, more research is necessary before

the complexities of the CaC03 cycle are fully understood.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 18

Historical Reviews of Palynological and Molluscan Research

Analysis of the palynomorphs preserved in the sedimentary mater­

ial from lakes have long been used to reconstruct ancient environ­

mental conditions. In addition to palynomorphs, mollusks preserved

in sediments provide valuable evidence for environmental interpreta­

tion.

History of palynology

The first recorded observations of fossil pollen grains and

spores were made by Goppert in 1836 in Miocene brown coal from

Germany. Postglacial palynomorphs were first recognized by Geintz

in 1887. Although pollen analysis has been practiced as a micro­

scopic technique since before the turn of the century, its full po­

tential as a research tool for studying vegetational history was not

realized until the early 1900's, when the Swedish geologist and

founder of modern pollen analysis, Lennart von Post introduced the

technique of representing the analytical results by pollen percentage

diagrams. Von Post found that he could demonstrate the similarity

of the pollen sequences within a small region and the differences and

trends among widely spaced localities. In 1921, Erdtman, a student

of von Post, made an historic contribution to palynology by quanti­

fying the science.

Since the middle twenties, pollen analysis has been the dominant

method for investigating late Quaternary vegetation and climatic

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 19

development. It is being perfected into a very refined method of

research, highly versatile and giving surprisingly intimate glimpses

into the contributions of life during earlier periods. Moreover,

palynology has become one of the most important auxiliary sciences

for archeology, adding to information derived from human relics and

macrofossils.

Previous palynological investigations in the midwest

The earliest pollen studies conducted in the midwest by Voss

(1934), Houdek (1935), Sears (1942) and Potzger (1946, 1948) re­

constructed the general succession of tree species from early post­

glacial times until the present. Pioneering studies by Sears (1948)

and his students on the vegetational history in Ohio, lead to the

development of a five-fold vegetational division of postglacial

time for the mid-west region. The divisions, patterned after the

Blytt-Serander post-glacial pollen sequence for northwestern Europe,

consist of: I, basal spruce-fir; II, overlying pine; III, beech-

maple deciduous forest; IV, oak-hickory-prairie; and V, mixed

deciduous forest.

Before 1950, few pollen studies included non-aboreal pollen

(NAP) and many only counted 100 tree pollen grains. Anderson (1954)

investigated the pollen at George's Reserve in southeastern Michigan;

however, his analysis was statistically limited, and redeposition of

older pollen grains and the lack of radiocarbon dates prevented accu­

rate correlation with other sites.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 20

Recently, more extensive study of herbaceous plant pollen, par­

ticularly the pollen of various aquatics and grasses has aided in

the interpretation of open areas such as open (spruce) woodland,

parkland, prairie, tundra and marshland. New research on pollen

rain and productivity from surface and atmospheric sampling in the

modern environment provides better insight into the interpretation

of the pollen record (Ritchie and Lichti-Federovich, 1967; Terasmae,

1976; Davis, 1969; and Webb, 1974). The introduction of new tech­

niques by David (1965) and Benninghoff (1962) have not only increased

laboratory precision but also statistical reliability.

Comparison of pollen diagrams from recent palynological investi­

gations in Michigan, Indiana, Ohio, New York, and Minnesota (Fig. 4)

provides further insight into the migration and variation of ecotones

in the northeast since the last glaciation. Within Kalamazoo County,

three other locations have had some pollen studies; including two

sites where pollen was extracted from sediment associated with

mastodon remains found in marl pits near Climax (Benninghoff and

Hibbard, 1961, Benninghoff, 1961) and Scotts (Semken, Miller and

Stevens, 1964). More recently a core taken from Wintergreen Lake

was analyzed for pollen by Bailey (personal communication).

Previous work with post-glacial mollusks

Walker (1879, 1895, 1906) was a pioneer in the study of

Pleistocene and living mollusks of Michigan, having published a

catalogue of Michigan Mollusca as early as 1879. Baker (1916, 1918,

1920) collected data on fossil and modern mollusca from numerous

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•BD • LM

0 7

f i M

•HL •PL _ •S 3S« •SB

km 3 0 0

Figure 4. Site map showing the location of pollen cores in the Great Lakes Region. Key: A, Alfies Lake (Saarnisto, 1975); VB, Victoria (Terasmae, 1973); H, Harrowsmith Bog (Terasmae, 1973); VN, van Nostrand Lake (McAndrews, 1970); C. Crieff Kettle Bog (Terasmae, 1963); PT, Protection Bog (Miller, 1973); CR, Crystal Lake (Walker and Hartman, 1960); MY, Myrtle Lake (Janssen, 1968); LC, Lake of the Clouds (Craig, 1972); W, Weber Lake (Fries, 1962); BD, Bog Pond (McAndrews, 1966); T, Terhell (McAndrews, 1966); HS, Horseshoe Bog (Wright, 1968); CB, Cedar Bog (Wright, 1968); KM, Kirchner Marsh (Wright, 1968); LCL, Lake Carlson (Wright, 1968); M, Madelia (Jelersgma, 1962); RZ, Rutz Lake (Waddington, 1969); P, Pickerel Lake (Watts and Bright, 1968); LM, Lake Mary (Webb, 1974); SD, Seidel Lake (West, 1961); D, Disterhaft Bog (West, 1961); BM, Blue Mounds Creek (Davis, 1977); GR, Green Lake (Lawrence, 1975); V, Vestaburg Bog (Gilliam et al., 1967); F, Frains (Kerfoot, 1974); W, Wintergreen Lake (Bailey et al., unpublished); CL, Climax Bog (Benninghoff et al., 1961); SC, Scotts Bog (Smeken et al., 1964); SH, South Haven (Zumberge and Potzger, 1956); S, Silver Lake (Ogden, 1966); PL, Pretty Lake (William, 1974; SB, Sunbeam Prairie Bog (Kapp and Gooding, 1964); KB, Kohoma Bog (Englehardt, 1960); 0B, Oakland Bog (this study); BS, Bacon Swamp (Englehardt, 1960).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 22

sites in the midwest including several sites in Michigan. The

continuing research of Baker resulted in his Freshwater Mollusca of

Wisconsin in 1928, a standard reference in the field.

Working with marl deposits at Castalia and along Tinker's Creek

in Ohio, Sterki (1920) was among the first to make interpretations

of depositional environment from molluscan faunal assemblages. More

extensive ecological studies began in the 1930's when Baker (1930,

1937) examined the effects of glaciation on a molluscan fauna. He

not only used Pleistocene mollusks as ecological indices, but also

as time indicators.

Archer (1939), Berry (1943), Goodrich (1931, 1943) and van der

Schalie (1953) carried out experiments to document ecology and dis­

tribution of living mollusks. Cain and others (1950) conducted a

study at Sodon Lake, Oakland County, Michigan, and used pollen and

molluscan data to reconstruct the depositional and ecological his­

tory of the lake and lacustrine community.

Since the early 1950's, LaRocque and his students have supplied

much useful information on the distribution and ecology of Pleistocene

mollusks through the study of numerous lacustrine deposits in the

midwest, particularly Ohio and Canada (LaRocque, 1966). LaRocque

(1966, 1967, 1968, 1969, 1970) has summarized and revised the classi­

fication, ecology and distribution of Pleistocene mollusca in the

midwest, other northern states and parts of Canada, in his Pleistocene

Mollusca of Ohio monographs.

Other workers whose contributions have helped advance molluscan

study include Dexter (1950), Herrington and Taylor (1958), Mowery

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(1961) and Clovers (1966). Quantitative distribution of species

was used by Zimmerman (1960) to demonstrate changes in the environ­

ment, particularly climate. Nave (1969) researched the ecology,

environment and distribution of fossil mollusks common to Ohio. The

ecology of modern analogues of midwest late and post-glacial fossil

molluscan fauna has been summarized by Clarke (1973). The distri­

bution, ecological requirements and habitats of living mollusk species

in Canada are currently under study by Kalas at the Canada Centre for

Inland Waters.

Studies by Camp (1974) and Wootten (1975) tie directly to the

molluscan analysis at Oakland Bog. Camp's (1974) observations in

several marl lakes in Michigan and Ohio have provided useful informa­

tion for determining molluscan zones. Data on terrestrial gastropods

from Pleistocene marl deposits in St. Joseph County, just south of

the study area have helped to complete the climate-mollusk relation­

ship (Wootten, 1975).

Paleoecological interpretations of molluscan faunas are based

on analysis of the vertical distribution and succession of significant

molluscan species within the sedimentary sequence. Changes in

climatic conditions, since the retreat of Wisconsinan ice, are also

distinguishable from the molluscan time-distribution pattern. Many

species indicate a more southerly distribution during early post­

glacial time (La Rocque, 1968). Living representatives of the late

glacial and early post-glacial mollusks of Ohio, Indiana and Michigan

are today common in northern Wisconsin and Manitoba (LaRocque, 1968).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. METHODS AND MATERIALS

Fieldwork

Fieldwork was designed to collect sufficient, uncontaminated

materials to permit pollen analysis, molluscan faunal study, recon­

struction of sedimentary facies and radiocarbon dating. The field­

work at Oakland Bog was done in three stages and involved: 1) pre­

liminary reconnaissance and mapping the extent of the marl; 2)

equipment testing; and 3) sample and core recovery.

Reconnaissance fieldwork was done in May, 1977 when the vege­

tational growth in the bog was minimal. Sequential air photos were

used to estimate the approximate extent of the bog. Peat thickness

was measured and the nature and distribution of underlying sediments

were determined with the aid of a soil auger and jet down rig (Fig. 5).

However, limited access due to quick muck, standing water, thicket

tangles and landowner refusal of access onto land prevented the mapping

of Oakland Bog in its entirety.

After preliminary reconnaissance and equipment testing, three

sites were chosen for core recovery with a modified fixed-piston

corer. Before coring, jet-down borings enabled the thickness and

gross lithology of the marl to be determined (Table 2). Because the

coring apparatus could not handle core barrels greater than 10 feet

in length, several cores were taken in sequence, labeled and brought

back to the laboratory for extrusion.

24

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. N3 Ln

12 • • • DIRT ROAD PAVED PAVED ROAD DRAINAGE DRAINAGE DITCH •CONTOUR LINE •CONTOUR 20 14 14 • 3 7 # km

671 ci h o l e

o l e H down

j e t auger 10 • • • X JJCORE 38Jj # 22 32 33 523ci * . 5 0 4 ® 2 8 34 Figure 5. Map of Oakland Bog showing location of cores, jet-down holes and auger holes. Materials core and jet-down holes are indicated in centimeters. Cross-section X Y used in Figures 7 and 13. are encountered in auger holes (numbered sites) are listed in Table 3. Depths to base of marl in 26

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Table 2 Study site sampling thickness of auger reconnaissance

number of peat (cm) contact below

1 61 till 2 61 till 3 30 till 4 30 till 5 30 till 6 61 marl 7 91 marl 8 122 marl 9 122 marl 10 91 marl 11 61 marl 12 30 till 13 30 till 14 61 marl 15 61 marl 16 91 marl 17 61 marl 18 61 marl 19 30 marl 20 61 marl 21 122 marl 22 122 marl 23 91 marl 24 91 marl 25 61 marl 26 30 till 27 61 till 28 61 marl 29 61 marl 30 61 marl 31 91 marl 32 122 marl 33 61 marl 34 30 till 35 30 till 36 30 till 37 30 marl

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 27

Additional jet-down borings were made in September, 1977 in

an effort to better establish the basin morphology. Unfortunately,

the borings had to be limited to accessible areas where standing

surface water was readily available for pumping.

Equipment

Two-inch diameter schedule 40 PVC pipe was used as core barrel.

Five and ten foot sections were fitted with male and female couples

to facilitate taking sequential cores. After taking the first section

of core, the upper part of the hole was supported by a casing of

three-inch diameter schedual 40 PVC pipe, five feet in length. Be­

fore reentry for successive cores, the hole was washed out to obtain

suitable reentry passage. The core was recovered with a modified

fixed-piston coring device (Wright, Livingston and Cushing, 1965).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 28

The Laboratory

Subsampling for pollen analysis

The core sections were extruded, split in half lengthwise with

a steel guitar string and carefully logged. Cores from sites I

and II were processed to extract the pollen present in the sediment

using a method modified from Faergi and Iverson (1975) (Table 3) *

One cubic centimeter of sediment was removed every 10 cm through­

out the length of the cores from sites I and II. After preliminary

counting of site I pollen slides, selected intervals were further

subsampled for additional control and detail.

Each sample was weighed wet, dried at 200°C for 12 hours and

reweighed to determine the water content. The one cubic centimeter

volume, which had been recommended in the literature for absolute

pollen frequency (APF) calculations, was found to be too time con­

suming and introduced a mixing error in the final residue because of

the long digestion time for marls.

Additional samples were further subsampled to a weight of less

than one gram but more than 0.5 grams of marl, this however limited

the precision of the volumetric determination. No APF calculations

were possible on these samples because the marl had become to dry

for determination of the water content and bulk density.

An exotic spike (Lycopodium) was added to each sample prepared

for pollen extraction. Samples with less than three spike tablets

had limited exotic recovery. The unreliable recovery of the spike

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 29 further limited APF calculations because the confidence limits were

too broad for statistical analysis.

Table 3 Pollen extraction procedure

1. One cc subsampled, sample weighted wet, dried 12 hours at 200°C, reweighed dry sample

2. Addition of exotic tablet with Lycopodium spike (each tablet contains 12,500+500)

3. 10% HC1 digestion to dissolve CaCOg

4. Centrifuge and decant

5. KOH water bath for peats (optional for marl), deflocculating procedure

6. Wash with water

6a. Remove samples for diatom analysis 7. HF treatment, one week digestion or heat to steaming in nickel crucibles, omit for peats and non-mineral materials

8. Wash with water 3 times

9. Centrifuge and decant

10. 10% HC1 water bath to prevent precipitation of silica fluoride

11. Wash with water 3 times

12. Centrifuge and decant

13. Drain well, acetylation with glacial acetic acid, drain

14. Acetolysis: solution of 9 parts acetic anhydride, 1 part sulfuric acid, water bath for 10 minutes or start at 40°C and heat to 99°C, centrifuge and decant

15. Wash with glacial acetic acid. Prevents reprecipitation of cellulose, centrifuge and decant

16. Wash with water 3 times, centrifuge and decant, drain

17. Mix organic residue with glycerol, mount in mounting media (cornsyrup).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 30

Palynomorph identification and counting

The mounted slides were traversed using a Leitz binocular micro­

scope and the palynomorphs were identified, counted and tabulated.

Exotic Lycopodium spores (Stockmarr, 1971) introduced to each sample

were counted and recorded along with the other palynomorphs for

determination of absolute fossil pollen concentration. The major

taxonomic entities identified and tabulated include: aboreal pollen

(AP), non-aboreal pollen (NAP), spores, aquatics, and indeterminables.

Indeterminables group are those pollen and spores which "cannot be

differentiated into distinct types amenable to precise description

and recognition" (Cushing, 1967). Grains whose essential features

were obscured as to make identification impossible were also placed

in the indeterminable group.

Palynomorphs were identified by comparison with the reference

collections from Alma College (Kapp, Dept, of Botany), University of

Michigan (Benninghoff, Dept, of Botany), Brock University (Terasmae,

Geological Sciences Dept.) and Western Michigan University (Naeve,

Geology Dept.). Technical assistance was attained from Drs.

Benninghoff, Kapp, Terasmae and Ms. Winn and Mr. McAtee. Keys of

Wodehouse (1935), Kapp (1969), Richards (1970) and McAndrews et al.,

(1973) were also utilized in the identification of the palynomorphs.

Pollen slides are housed at the Western Michigan Museum (numbers

3022-3110).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 31

Molluscan analysis

One cup of marl collected from the homogenous wash boring over­

flow was sieved, washed and allowed to air dry. A representative

sampling of shells were then picked with a fine paintbrush for gen­

eral identification, labeled, and photographed and placed in capsules

for future reference. The total number of individuals in a particular

species was tabulated for each sample interval (Plate 1).

A malacological diagram is organized by the mollusk groups:

pelecypods, operculates and aquatic pulmonates. The absolute number

of individuals in a species were plotted against the sampling depth

(Plate 1). Diversity curves of the three mollusk groups were also

constructed (Fig. 11) to help interpret changing environmental con­

ditions through time.

Ostracod, diatom and floral macrofossils were also preserved

in the sediment samples; however, no quantitative analysis was made

of these elements.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 32

Radiocarbon Dating

The term "radiocarbon" refers to the naturally occurring isotopes

of carbon. The isotope used in radiocarbon dating is carbon-fourteen

which has an atomic weight of 14 atomic mass units and a half-life

of 573CHF40 years (Godwin, 1962).

Six marl samples and one peat sample were submitted for radio­

carbon dating at the Radiocarbon Dating Laboratory of Brook University

(Table 4). The samples were processed for analysis by Howard Melville

with the Picker Nuclear Benzene Synthesizer and the counts were made

using the Picker Nuclear Liquimat 220.

The marl sediment and enclosed shells were digested with dilute

HC1 (10%). The recovered organics (excluding roots or plant fibers)

were used for dating purposes. At least 50 grams of marl were neces­

sary for the dating procedure; however, only 10 centimeters of sedi­

ment could be processed without spreading the dated interval too far.

Dating of the peat required 15 to 20 grams of material. The

general procedure for the pretreatment of peats involves (Melville,

1972):

1. removal of rootlets, stones and other inorganic materials

2. treatment for humic acid with 0.2 N NaOH for 1/2 hour, followed by 15 minutes with 0.2 N HC1.

3. wash until neutral and dry thoroughly.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 33

Table 4

Radiocarbon dates from Oakland Bog

sample number age depth cm location material

BGS 446 16,420+570 370.8-381 site I, marl A

BGS 447 6,060+100 45.7- 44 c-i ho T noof D

BGS 448 16,500+330 670.5-642.6 site II marl JF

BGS 449 13,300+230 520.7-505.5 site III„ marl with D peat fragments BGS 495 12,100+400 254 -264 site I marl

BGS 496 10,530+400 184 -194 site I marl

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. NATURE OF THE SEDIMENT

The sediment in each core taken from Oakland Bog was measured,

described in terms of sedimentary structures, color indexed with

the Geological Society of America rock color chart and observed

for macrofossils and textural variations (Appendix 1). Sections of

the cores were photographed before samples were removed (Fig. 6).

Three sediment types were observed:

1. basal till 2. marl 3. surface peat 4. coarse sand (dike).

The till, a gray fine sandy loam, floored the depositional basin.

At site II, a coarse sand dike, approximately 30 cm long was found

just above the mar1-till contact. The till was washed of the clay

fraction and compared to the sand dike; the dike sand was better

sorted and coarser than the grains derived from the till. Marl clasts

incorporated into the dike, ranged from % cm to 1 cm in length, and

the dike-marl contact was sharper and darker than either material. It

is possible that the sand dike could have resulted from frost activity

during the early stages of the lake.

The basal marls were medium gray, clayey, and contained abundant

calcified cortical tube casts from charophytes (Fig. 6a), mosses and

fine plant debris. Ostacods, diatoms and mollusk shells were common

through the marl section. Some sections of the marl were distinctly

laminated (Fig. 6b), displaying alternating light (carbonate-rich)

and dark (organics greater than carbonate fraction) laminae separated

34

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 35

H

Figure 6a. Calcified cortical tube casts from charophytes found in washings from basal marls (sites I, II, III, IV, V, VI, and VII). Sample photographed from washings of site VI (762cm).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 36

Figure 6b. Laminated marl displaying alternating light and dark laminae separated by massive and shelly layers. (Site I, core A: 330cm)

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. HONI

Figure 6c. Thread or root-like plant debris and shells in marl. (Site I, core C: 80cm)

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 38

Figure 6d. Nitella reproducing bodies in marl. (Site II, core B: 430cm)

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 39

Figure 6e. Peat fragments in marl. The lower wavy peat fills a scour. (Site III, core A: 278-277cm.)

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 40

by massive or graded unlaminated beds. Each light lamina grades

gradually into that of the overlying darker lamina; the contact

between a dark lamina and the overlying light lamina is sharp.

Darker layers often appear streaked or mottled. Laminae range in

thickness from a few millimeters to one centimeter with the lighter

laminae commonly being the thickest in a sequence. Thread-like or

root-like plant fragments (Fig. 6c), Nitella reproducing bodies

(Fig. 6d), oogonia of Chara and peat fragments (Fig. 6d) were observed

at certain intervals in the cores (Appendix 1).

Preliminary scanning electron microscopic observations have

shown the marl to be composed largely of calcium carbonate aggregates.

Although no detailed geochemical tests have been performed on the

marls, reactions observed during the acidilizing (10% HC1) stage for

pollen extraction suggest the lighter marl contains more carbonate

because it reacted more vigorously and took longer for the digestion

of calcium carbonate to go to completion. This general observation

agrees with the extensive geochemical research of Brunskill (1969)

who has shown the light laminae at Fayetteville Green Lake, New York

to contain more carbonate than darker laminae.

Discussion of Marl Deposition

The depositional history of the Oakland Bog lake basin has been

reconstructed by comparing cores from three sites (Fig. 5, Appendix 1)

and six jet down samples (Fig. 5, Appendix 1) with the sediment and

associated fauna and flora of other studies involving temperate gla­

ciated marl environments. Jet down sites other than core locations

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. SITE d o i I I / BANK CARBONATE

' DEEP Figure 7. Cross-section (Z to Y, figure 5) of Oakland Bog based on core (I, II, III). present ground surface

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 42

provided additional control on the morphology of the lake basin.

Site I is located on a prominent rise (Fig. 7) which was prob­

ably initiated by some irregular deposit in the ice block, resulting

in a configuration that served as a nucleus for the calcareous depos­

its forming what is interpreted to be a lakemount. Early accumula­

tions of organic matter and pest fragments (Fig. 6c) in this littoral

area were probably associated with a relatively high input of

allochtonous debris during the transitional period between tiaga-bog

conditions and the forests developing under warmer climatic conditions.

Relatively shallow water conditions prevailed at site I as evidenced

by in situ root-like structures enclosed in the marl throughout much

of the core.

The thick layers of coarse, light colored and shelly marl be­

tween darker fossil-poor layers have been suggested by Brunskill (1969)

to represent marl formed in the epilimnion. The biogenic-rich layers

appear to represent the summer depositional conditions when the

epilimnion becomes supersaturated with respect to CaC0 3 thus promot­

ing the production of marl and shell formation (Brunskill, 1969).

The marl sequence at site II is interpreted to represent the

progradation of a carbonate bank. Similar variations in the marl

laminations recorded at Oakland Bog have been reported by Lane (1903),

Brunskill (1969) and Wetzel (1979) as reflecting the amount of par­

ticulate and organic matter deposited with the carbonate which in

turn indicates the quality and quantity of overlying water.

Deep-water laminations are abruptly terminated at a scoured sur­

face (interval 574cm-579cm) becoming a sequence of alternating massive

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 43

and laminated beds indicating a period of rapid deposition followed

by quiet settling similar to the turbidite association observed by

Terleckly (1974). Abundant carpaces of Nitella in mottled to faintly

laminated sediments above this sequence record open-water slope

conditions. The bank margin is recognized as the purest part of the

marl bed at site II, being composed largely of molluscan shells,

charophyte fragments and indeterminable aggregates of calcium car­

bonate .

Water depth at site II is interpreted to have fluctuated from

deep to shallow and back to deep again reflecting the drier Pine

Period (see Section V, on Pollen Stratigraphy) interval (275cm-180cm).

The shallow interval is recorded by grayer marls with more organics,

plant debris, roots and peat fragments. As progradation proceeded,

increasing amounts Of organic matter and shallow water species of

mollusks associated with the inner littoral and shoreline became

dominant components in the marl (Fig. 8).

The persistence of deep-water conditions at site III are indi­

cated from thick accumulations of dark, faintly laminated to laminated,

fine grained marl sediment. A comparison of these sediments to those

described by Brunskill (1969) suggest that the laminations represent

annual layers that formed in either the hypolimnion of a dimictic

lake or the monomolimnion of a meromictic lake.

Above the laminated sequence, the sediment grades to a darker

marl containing peat fragments with no distinct laminations suggesting

site III was also influenced by the drier climate of the Pine Period.

This shallow water interval is immediately overlain by sediment

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 44

cm sediment description sitelE environment interval and interpretation community

Littoral. Abundant shallow water molluscan fauna, 183 plentiful plant debris, peat fragments and many root struc tures in laminated marl. rrTTTCT^KOURISHING Carbonate bank. Light lamina 305 gradually grades into over- lying darker lamina, sharp contact between dark to light laminae. Lamina few mm to transition 366 one cm. 1 deeper Deep water 12'. The assoc­ 458 iation of Nitella and mass­ ive carbonate reflects deep water conditions such as along the slope. 488 'a BLJSHMEN'T

Prograding bank sequence be­ shallowing 518 gins. Time of shallower con­ ditions, suggested by roots, peat fragments and shallow­ ing molluscan assemblage. 548 CHMENT IT Slope. Graded bedding, bro­ ken shells of deep and shal­ ^JgkeJev«l low water molluscan fauna, scoured surfaces intermittent 6 4 0 with gray massive carbonates- turbidite deposition on pro­ grading slope surface. ENRICHMENT I Sand, moss and riverine form Pisidium suggest moving water occurred during this interval.

670 till carbonate PIONEER

Figure 8. Interpretation of site II based on vertical sequence of sediment facies and molluscan assemblages in core (11) II.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 45

containing Nitella carpaces which suggests that deeper water conditions

returned. The final shallowing stage of the lake is indicated by

abundant root-like and plant fragments in the coarser upper layers of

site III, and the development of a peaty marl suggesting a marginal

marsh depositional environment.

The presence of increasing amounts of organic matter and abundant

shallow water mollusks at the top of each core indicates shallow-water

conditions existed across the entire basin just prior to the sudden

drainage of the lake. The sharp marl-peat contact indicates the

abrupt transition from lake to bog, perhaps recording the influence of

the drier climate associated with the hypsithermal.

Sediment deposited in the shallow areas were stabilized by

marcophytic colonization. The increased sedimentation rates within

the littoral zone reflected greater productivity associated with

warmer climatic conditions.

Sedimentation Rates

Several estimates of sedimentation rates were calculated for the

marl sediments at Oakland Bog (Table 5, Fig. 9). Assuming a constant

sedimentation rate for the entire marl sequence, the mean sediment

rate for site I approximated ancient mean sediment rates for similar

lakes (Wetzel, 1973; and Terleckly, 1974). During the first 10,000

years (16,000 to 6,000 years B.P.), the mean sediment rate for site I

at Oakland Bog was 0.038 cm/year. Terleckly (1974) calculated the

mean sediment rate of a relic marl lake near Rochester, New York to

be 0.032 cm/year. Pretty Lake, Indiana has been examined by Wetzel

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10,500 - 8,500 Table 5 and pollen stratigraphic marker horizons 0.047 9,800- 8,500 0.077 400 0.045 Sedimentation rates for site I based on radiocarbon dates * ±400 6,060 ± 106 8,500 - 6,000 0.048 10.530 ± 400 10.530 * 400 0.032 9,800 - 9,000 - 7,400 6,000 0.063 0.054 12.100 16,420 i 570 12.100 0.027 15,000 - 10,500 0.028 from Oakland Bog rate (cm/year) horizon date (years B.P.) rate (cm/year) Radiocarbon dates Calculated sediment Pollen stratigraphic Inferred sediment

45 275 180 Sediment zone Zone I 381 and depth cm Zone II 275 Zone III 180

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 47

(1970) and Williams (1974) to determine ancient and present carbonate

sedimentation rates respectively. Williams (1974) determined the

ancient mean rate to be 0.039 cm/year while the current rate as cal­

culated by Wetzel (1970) is 0.113 cm/year.

Radiocarbon dates from Oakland Bog and established pollen marker

horizons are used to estimate sedimentation fluctuations at site I

(Table 5, Fig. 9). Variations in sedimentation rates are apparent

at Oakland Bog; however, detailed geochemical analyses, chlorophyll

degradation product analyses and additional sediment rates are nec­

essary for their proper interpretation.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 48

.08

.0 6

Ul % Radiocarbon data OT O Pollan markar * Today

TIME |4,0 < !)0 10,000 6,000y«araa-K l i HERS I SPRUCE i HARDWOOOS I PRAIRIE I l

Figure 9. Mean sedimentation rates of marl accumulation at Oakland Bog plotted against time and pollen zones. The bar ( |------1 ) indicates the time period for which the sedimentation rates were esti­ mated. (Table 5). Modern rates after Wetzel (1970) and Williams (1974).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. MOLLUSCAN RESULTS AND DISCUSSION

Paleoecological Reconstructions with Freshwater Mollusca

Fossil freshwater molluscan taxa and their relative abundances

can be used to interpret environmental conditions and reflect the

changes in a lake environment. Paleoecological reconstructions of

the Pleistocene have an advantage over those of earlier geologic

time since many of the species are still part of the present environ­

ment.

If one assumes that the species occupied the same type of en­

vironment in the Wisconsinan as they do today, one needs only to

study the ecology of living species to obtain an understanding of

their probable ecologic niches during the Wisconsinan. Regrettably,

only limited data is available for present species. Because the

lacustrine environment involves a complex interaction of physical,

chemical and biological parameters, reliable interpretations of

post glacial conditions may be tenuous.

The fossil fauna of lacustrine deposits are predominantly

thanatocenosic assemblages, being mixtures of both indigenous species

and intruders that have been deposited together after death. The

abundance of a species at any time is partly determined by the

availability of its specific ecological requirements and the effec­

tive community of its competitors and predators. However, this

relative abundance of a particular species in the natural environ­

ment may be difficult to estimate on the basis of its frequency in

49

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 50

a fossil assemblage. Currents and waves may lead to transport and

hydrodynamic sorting of empty shells in the shore zone.

The lacustrine sediments and their enclosed fossils are the key

to the geologic history of a lake basin. Molluscan fauna reflect

the changing ecological and depositional conditions of their habitat

(Camp, 1974; Kalas, personal communication). Water level fluctuations

respectively increase and decrease the range of freshwater mollusks

living in the nearshore areas. Distribution data from post glacial

marl deposits (LaRocque, 1968) and surveys of living Mollusca (Camp,

1974) support the presence of a series of concentric molluscan habitat

zones around lakes (Fig. 10) (Baker, 1928; Dexter, 1950; and Camp,

1974). The range of the terrestrial mollusks living in the fringing

marsh would also be affected by the lake level fluctuations; their

habitat increases during periods of low water and decreases in times

of high water. These changes in distribution are recorded in the

sediment column as variations in the abundance of species with depth.

The regional distribution of mollusks have also changed since the

last glaciation. The assemblage of mollusks which lived just south

of the full-glacial ice boundary, fossils which have been found in

late glacial and early post-glacial deposits of Ohio, Indiana and

Michigan, are found today in similar assemblages in Minnesota, Wiscon­

sin, northern Michigan and Canada (Baker, 1928; La Rocque, 1967;

Clarke, 1973; Camp, 1974; and Kalas, personal communication). Experi­

mental studies with living species of mollusks confirm the hypothesis

that they are sensitive to temperature conditions which further sup­

ports the assumption that fossil mollusk assemblages reflect climatic

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. i UPLAND SLOPE SHORELINE INNER LITTORAL OUTER LITTORAL DEEP WATER CO LAND SNAILS LAND SNAILS LAND SNAILS AQUATIC MOLLUSKS AQUATIC m ollusks AQUATIC MOLLUSKS i S /VOODLAND (MOISTURE (SUC CINEIDS) MANY PHYSA AM NICOLA LIMOSA VALVATA TOLERANT) COMMON ONCHARA io forms ) MANY GYRAULUS MARSTONIA U KJ AQUATIC MOLLUSKS i/» 'N HELISOMA GYRAULUS DECEPTA 3 PH YSA PISIDIUM VALVATA GYRAULUS FOS S ALIA O CINCINNATI A z GYRAULUS A M N I C 0 S & a l k Z

SWAMP SHRUB MARSH FLOATING-LEAF CO} BANK-OPEN DEEP SUBMERGENT £ g EMERGENT SUBMERGENT ARROWHEAD POND LI LY MILFOIL NITELLA a.* sN CAT T AI LS PONDWEED C MARA

ut * < ui PEAT _i -f z i ° 5< *0. Z GLACIAL SUBSTRATE

Figure 10. Molluscan Habitats of a Typical Marl Lake (after Camp, 1974). Ul 52

conditions (Berry, 1943).

As the Wisconsinan ice retreated across the area of the Great

Lakes, a series of spectacular changes took place; the effects of

which are reflected in the molluscan community in the Great Lakes

region today. The drainage of preglacial streams and the early

Great Lakes towards the Mississippi River permitted the invasion of

Mollusca from that great southern drainage into the Great Lakes.

The Mohawk-Hudson drainage outlet permitted the invasion of Mollusca

from the Atlantic drainage into the Great Lakes area (La Rocque, 1968).

These events help to explain the anomalous zoogeographical distri­

bution of some Mollusca in the States bordering the Great Lakes and

they are also useful in explaining the presence of other forms of

life in the Great Lakes drainage as it exists at present.

Oakland Bog Fossil Molluscan Description

The general trend of the fossil molluscan assemblages recorded

at Oakland Bog (Plate 1 and Fig. 11) is one of increasing diversity

and frequency through time. Good preservation permitted the identi­

fication of the mollusks to the species level. Most shells of the

mollusks are whole, bleached white or mineral stained and filled with

the same marl as the matrix. In some cases shells were pitted, many

the result of boring algae or fungi, while large holes may be the

result of predator activity, post-mortem solution or mechanical destruc­

tion. Upon closer examination of the deposit, the patchy distribution

of mottles in the marl were found to represent the accumulation of

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 73 CD "O o CQ. o CD Q.

■D CD

C/) 00

tc S iu o H o Z o T 3 Ul _J X Q. t- s Q. < Ul lu (/> co Q Q l < CD .183 a. R 244 3- CD * 3 0 5 CD ■D O ■ 3 6 6 Q. -4 2 7 O ZD 1 4 8 8 O ZD" ■ 5 4 9

CD -6 1 0 Q. ®670 O O 0 C CD 1500 ■O CD 1 1

00 C/5 o' = 3 Figure 11. Freshwater Molluscan diversity curves and total populations from site II, Oakland Bog.

Ul CO 54

shell fragments. It was also noted that the preserved shells are

now very fragile suggesting some chemical disintegration.

Site II (Fig. 12) was selected for molluscan faunal study be­

cause its interpreted proximity to land would favor faunal elements

most sensitive to water level fluctuations and temperature changes.

Using the principles of Walther's Laws for facies reconstruction, the

uninterrupted vertical changes are used to reconstruct lateral shifts

in the environment through time.

The molluscan fauna consists of 29 species representing 14 genera

(Plate 1). The faunal assemblages were composed entirely of aquatic

mollusks and consist of species that are characteristically confined

to a relatively silt-free permanent water body associated with cir­

culating water, indicating some current and/or wave action in portions

of the lake. Most faunae have a preference for soft substrate, con­

siderable vegetational growth and shallow water depths.

Comparison of Oakland Bog molluscan data with available modern

molluscan observations suggests that the faunal occurrences can be

subdivided into five stages based on diversity and frequency (Table 6

and Fig. 12). These stages represent the environmental evolution of

the lake at the study site.

Table 6 Environmental Evolution of Lake at Oakland Bog

STAGE I Pioneer Stage (670 cm) is characterized by low molluscan abundance and diversity in the basal marl. The marl is the grayest at this interval, containing abundant calcified Charophyte frag­ ments, fine sand, mosses and plant debris.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 55

Table 6 continued

STAGE II The First Enrichment Stage is designated at the (670cm-640cm) sample interval. Pulmonate gastropods are now included in the growing number of species and correspondingly increase in frequency. No distinct lamination are recorded in the brown to gray marl.

STAGE III A four-fold increase in abundance (518cm-549cm interval) while the number of different species occurring increased by only one. This interval is referred to as the Second Enrichment Stage. Unonid shell fragments, Nitella carps, Chara oogonia, some decayed leaf material and grass­ like plant debris were recovered from the brown- gray marl. The sedimentary structures include a mixture of alternating dark and light lami­ nations with scoured-irregular boundries cutting across several laminations grading into massive light colored marl units with peat fragments and root-like plant debris.

STAGE IV The 458cm-488cm and 305cm-366cm intervals record a significant decline in pulmonates and burrow­ ing species of mollusks. No Chara oogonia were observed but carps of Nitella were present in great abundance. Stage IV is the Community Establishment Stage which maintains approxi­ mately the same assemblages indicating the estab­ lishment of a permanent lake. The marl in this stage is dark and massive.

STAGE V The remainder of the sampled interval 304cm-183cm records a progressive increase in abundance while diversity remains nearly constant. This is the Flourishing Community Stage featuring the greatest abundance with maximum diversity. Well preserved laminations, averaging 1 to 2 cm, with peat inclusions at the irregular boundries and abundant grass-like threads characterize the generally lighter colored marl.

UNSAMPLED MOLLUSCA INTERVAL OBSERVATIONS 183 cm TO PEAT

No wash boring collections were made at this inter­ val due to mechanical difficulties. The correspond­ ing interval of core showed progressive darkening of the marl sediment, peat inclusions and shell-rich layers (packstone).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Ul as

ponded J)

* ------

Grydulus Grydulus parvus Valvata sincere hellcolldea Valvata trlcarlnata Gyraulus parvus - - - - Marstonla decepta Psidlum Psidlum compressum (riverine-form) Physia soyll soyll Physia Hellsoma anoeps _ _ < st Amnicola walkeri Amnicola walkeri •« Gyraulus Gyraulus parvus Valvata hellcolldea sincere (low diversity, low frequency) cinnatiensis Cincinnotia | trlcarlnata Valvata Pisidium lllljeburgi 11 11 Marstonia decepta Amnicola llmosa — Amnicola limnosa i d i o l i i k i a STAGE ELEMENTS PIOMPFR tC TA O i icuiiCTKiT Marstonla decepta to ADUIO1 nM tN 1 Valvata tricarinata ENRICHMENT i?i s \i FLOURISHING MOLLUSCAN DOMINANT MOLLUSCAN SITE II POSITION Figure 12. The transition of stages of molluscan assemblages and relative water depth recorded at Site II.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 57

Discussion of Molluscan Stages

The occurrence of Pisidium compressum, commonly associated within

a modern creek habitat (Kalas, personal communication) and the low

diversity of the pioneering stage may represent a ponded or inter­

mittent creek during the initial stage of Oakland Bog's development.

Sand and mosses at the base of the core from site II further support

a creek environment (Kalas, personal communication). The nature of

the early drainage is highly speculative since the pro-glacial drainage

of the Kalamazoo area is not well understood and these conditions

could reflect meltwater drainage or debris associated with the ice.

Pisidium lilljeburgi, Gyraulus circumstraitus, and the riverine form

of Pisidium compressum together suggest oligotrophic and cold

climatic conditions persisted into the First Enrichment Stage at

Oakland Bog.

The basin at Oakland Bog was most likely interconnected with other

depressions in the vicinity during the early stages of deglaciation.

The 1926 topographic maps show the site to be at the junction of

several small creeks within the wetland. The basin must have filled

rather rapidly to allow the sudden transition from ponded or inter­

mittent creek to permanent lake molluscan species which were apparently

already established in the region.

The invasion of pulmonate gastropods marks the First Enrichment

Stage (interval 640cm-670cm) but more importantly reflects the growth

of aquatic vegetation (Kalas, personal communication). The water must

have been relatively cold and well oxygenated during this stage because

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the presence of Sphaerium striatum and Marstonia decepta required such

conditions. The occurrence of Pisidium nitidium throughout the vertical

profile indicates that Oakland Bog was a hardwater lake from the initial

stages to the final in-filling with peat.

Once the lake became established, marl rapidly accumulated in

shallow areas, building carbonate banks upon which vegetation developed,

further enhancing the marl sedimentation. The mollusk evidence (Plate 1)

suggests there were several coexisting habitats and these varied com­

ponents together support the development of a shallow, hardwater lake.

Many new niches were also created when the basin began to fill.

The lake took on new dimensions and the additional microhabitats sup­

ported a variety of floral and faunal assemblages (Fig. 13). Marshes,

beaches, mudflats and protected bays developed along the shoreline

providing new habitats for operculates and especially pulmonates as

vegetation further developed in the shallows. Inlets, mounds-and islands

further modified the water circulation. The shallow bays offered pro­

tection from waves and supported extensive aquatic plant growth. These

quiet areas not only were favorable sites for mollusks inhabitation

but were also accumulation sites for empty shells swept in during times

of high water.

The enrichment stages I and II recorded the immigration of shallow

water species tending to require cool climates or warmer, such as

Gyraulus parvus, Amnicola limosa, Marstonia decepta, Valvata sineera

helicoides and Valvate tricarinata. The deeper parts of the lake

simultaneously acted as a refugia for the previously existing cold-loving

species Amnicola walkeri, Pisidium ferrugineum and Gyraulus deflectus.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 59 SITE LOCATIONS O (ne

HYPOTHETICAL

sffallowing FLOURISHING

ESTABLISHMENT

SAND ' • E n r ic h m e n t ii permanent lake

MARL ENPflCHMENT intermittent creek-pond t

. , ® TILU 7 % PIONEER

Figure 13. Molluscan stages and lake history of Oakland Bog as inferred molluscan assemblages.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 60

The dominance of the Gyraulus parvus, Marstonia decepta and Valvata

tricarinata assemblage during the Second Enrichment Stage (interval

548.6cm-518cm) suggests that extensive vegetated shallows had developed

in the marl lake. These shallows may be reflecting two possible changes

in the local environment. The gradual build-up of carbonate banks in

a generally enlarging lake would create suitable shallows for inhabi­

tation (Fig. 4 and 12). Alternatively, a drier climate would also be

reflected as a lowering of the water table (Fig. 14).

In the case of Oakland Bog, both conditions appear to have in­

fluenced the lake. Radiocarbon dates, pollen stratigraphy and assoc­

iated sediment facies correlate the molluscan faunal shift with a

warmer, drier climate and carbonate bank progradation. Although the

date associated with this phenomenon is too old for direct comparison

at other sites, it does however approximate the beginning of the Pine

Period relative to Oakland Bog's dated pollen stratigraphy (Plate 2).

The Pine Period, or zone II has been interpreted to be the result of

a cool but much drier climate than previously encountered since de­

glaciation (see section on pollen stratigraphy, zone II).

The marl sediment is distinctly laminated prior to the Gyraulus

parvus population explosion (interval 154cm-158cm), becoming darker

with more plant debris but never indicating that the lake completely

dried up.

Warmer conditions were also needed to support the immigration

of species such as Amnicola limosa, Marstonia decepta, Helisoma anceps,

and Pomatiopsis lapidaria (Plate 1). The occurrence of Musculium

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.

TIME TIME TIME I Figure 14. Environmental reconstructions of conditions present during the Second relationship and ii. vertical relationship. Enrichment Stage. A. Assumes position on the bank is maintained and water level results in shallower water conditions at the same reference point — i. lateral decreases due to drier climatic conditions. B. Progradation of carbonate bank

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 62

lacustra and increase of Promenetus exacuous are associated in the

modern with the dropping of lake levels further supporting a dry

period hypothesis for Oakland Bog (Clarke, 1973; Camp, 1974; Kalas,

personal communication) (Fig. 13).

The significant decline in pulmonate species during the Commun­

ity Establishment Stage (intervals 488cm-457cm and 366cm-305cm) along

with the general decline of mollusks may reflect deeper water con­

ditions (Fig. 13). The site previously located near the bank margin

was flooded with rising water levels as suggested by the darker marls

and the abundance of carpaces from the deep water charophyte, Nitella

(Fig. 6d). The molluscan assemblage is diverse but with low frequen­

cies suggesting several habitats are in transition and under the

stresses of a changing environment.

The Oakland Bog molluscan community was rebuilt with species

that were able to adjust to large permanent lake conditions including

Marstonia decepta, Valvata tricarinata, Pisidium casertarum and

Gyraulus parvus (Plate 1; Fig. 12, 13). Species associated with the

arctic or subarctic become insignificant, most likely surviving in

deeper sections of the lake during the Community Establishment Stage.

Warmer climatic conditions can be postulated since these species

demonstrate a preference for mixed forest climatic regime instead

of the tundra or boreal forests of the earlier bottom assemblages.

The continued build-up of marl produced a wide terrace upon which

vegetation proliferated as the climate warmed. The transition of the

Established Lake Community to a Flourishing Lake Community is marked

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 63

by the increase in frequencies of groups while the diversity remains

nearly constant (Plate 1, Fig. 12).

The dominance of species such as Gyraulus parvus and Marstonia

decepta suggest lake conditions became increasingly eutrophic from

the Establishment to the Flourishing Community (Fig. 12, 13).

Cincinnatia cincinnatiensis, Marstonia decpta, Pomatiopsis, Physa

sayii, and Helisoma anceps occur today in mixed forest or hardwood

deciduous forest climatic belts (Kalas, personal communication).

These warmer associated species replaced the cold-cool molluscan

assemblages as the dominant community. The cold-loving species such

as Amnicola walkeri, Gyraulus circumstraitus, Valvata sineera helicoides

and even Amnicola limosa, a cool boreal indicator, display a time-

transgressive migration northward as the lakes filled with sediment

decreasing the lake depth and cold water refugia sites. The migration

pattern and modern locations of these mollusks also support the theory

of a gradual climatic warming.

The presence of Fossaria parva during the Flourishing Community

Stage records the development of mudflats resulting from the shallow­

ing of the lake, as do the replacement of Physa sayii by Physa skinneri

and corresponding increases in Promenetus exacuous and Musculium

lacustra populations (Clarke, 1973; Camp, 1974; and Kalas, personal

communication) (Plate 1). The increase of Gyraulus parvus in this

stage at the expense of other mollusks also indicates that the environ­

ment was steadily changing to shallow water, near swampy conditions

(Plate 1). The large populations of Marstonia decpta, Valvata tri-

carinata, and Gyraulus parvus are commonly associated together today

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along marl benches where there is luxuriant aquatic plant growth which

suggests that the fossil assemblage at Oakland Bog reflects the devel­

opment of a broad, shallow and highly vegetated shelf in the lake

basin before the sudden transition to peat (Fig. 13).

The upper portions of cores and wash borings indicate shallow

water conditions at all sites. The light tan sediment is similar

to the littoral bank marl observed at Littlefield Lake (Central

Michigan) and reported from Pretty Lake in Indiana (Williams, 1974)

and Fayetteville Green Lake in New York (Brunskill, 1969). The

darker marl contains much organic material including plant debris

and many peat fragments which are also commonly observed in shallow

water zones in modern marl lakes (Williams, 1974; and Wilkinson,

personal communication).

No terrestrial gastropods were observed in either the marl or

overlying peat at Oakland Bog. Wootten (1975), working in St. Joseph

County, just south of Kalamazoo County recorded the transition of

aquatic mollusks to terrestrial mollusks in a mar1-peat bog. The

terrestrial gastropods there suggest the transition to a warmer cli­

mate as also postulated by the pollen stratigraphy at Oakland Bog.

The final stages of the lake records a rapid sediment accumula­

tion. Sedimentation rates were between 0.05 cm and 1.0 cm per year.

Warmer conditions probably promoted macrophyte and algal photosynthetic

activities which in turn stimulated the production of marl. The

abrupt transition from marl to peat suggests that the lake basin

drained suddenly, interrupting the mollusk record. This may be a

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function of the introduction of the hypsithermal interval (6,000

years B.P.).

As more information is made available regarding the distribution,

habitats and ecology of living mollusk species, depositional and

climatic reconstruction of fossil communities becomes more accurate.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. POLLEN STRATIGRAPHY

Principles of Palynology

Palynomorphic data presentation and interpretation

Vegetation has left a fossil record unrivalled in stratigraphic

completeness and detail in the form of pollen grains and spores. The

versatility of palynomorphs is dependent upon four factors:

1. The greater resistance of pollen and spores to degradation

than most other plant parts, thus facilitating their survival as

fossils.

2. The small size of pollen grains and spores (most less than

20yOf- permits them to be transported and deposited as sedimentary

particles.

The morphological complexity permits various characteristics

of the pollen grains and spores to be distinguishable.

4. The enormous production of pollen grains and spores

facilitates recovery in statistically significant quantities.

Disagreements are frequent concerning the proper format with

which to present the pollen data and the corresponding vegetational

assemblage interpretation. Further consideration of what the deposited

pollen represents and how best to evaluate the pollen rain are neces­

sary to determine the method(s) of pollen analysis. The possibilities

as well as the difficulties of pollen analysis are described by

Faegri and Iverson (1975) and Wright (1970).

66

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 67

The first step is to present the pollen data in such a manner

that each pollen spectrum can be translated into terms of a vegata-

tional assemblage. Pollen stratigraphy is based on the assumption

that pollen assemblages preserved in lake sediments should reflect the

surrounding vegetation and changes in the pollen record through time

reflect regional changes in vegetation with which climatic history

can be inferred (Davis. 1967b: Wright. 1968; and Faegri and Iversen.

1975). Absolute chronology and pollen influx data provide information

on vegetational migration, establishment and succession (Davis and

Deevey, 1963; and Wright, 1976).

Most pollen diagrams illustrate the pollen sequence taken from

deposits in the bottom of low areas often formed by melting ice blocks.

These areas were often filled with ice long after the glacial front

had receded and plant succession had begun. Therefore, it is difficult

to find pollen samples that represent the earliest vegetation which

grew in an area. Also, if ground cover such as the mosses now present

in the subarctic-arctic regions of Canada exemplified the postulated

ice front vegetation, lake deposits may not have recorded those pollen

and spores since they are usually not trapped within the moss polysters

(Fredskill , 1975; Nichols, 1975; and Terasmae, 1976).

Ideal deposits for pollen analysis are those in which the surround­

ing vegetation producing the majority of the pollen has not influenced

the state of preservation of pollen grains. Bog plants record the

local hydrologic conditions which may not reflect the regional climate.

Pollen and spores are also subject to corrosive oxidation in peat thus

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 68

destroying some of the pollen recorded. Regional vegetation approxi­

mations are best reconstructed from pollen and spores preserved in

the sediments recovered from closed, moderate-sized lakes (Faegri

and Iversen, 1975).

The interpretation of post glacial pollen diagrams in terms of

vegetational change has long been recognized as a complex problem

requiring not only a thorough knowledge of the present-day ecology

of the taxa involved, but also information on the relationship

between modern pollen rain and the vegetation from which it is derived.

Because of factors such as the physiological and ecological condi­

tions affecting the flowering and pollen production of the individual

plant; the abundance of the taxon within the vegetation, the structure

of the community in which it occurs, the mode of pollen dispersal,

the meterological factors influencing pollen transportation, and the

physical, chemical and biological conditions that control pollen

sedimentation and preservation at the site of deposition, the pro­

portion of the pollen of different species preserved in the sediment

is usually not the same as the proportion of the species in the

vegetation (Havri, 1967; Tauber, 1967; David, 1969; and Birks, 1973).

A quantitative approach to rationalize the discrepancy through

the application of correction factors to the components of the pollen

rain has been advocated by Davis (1963). For most purposes it is

generally assumed that the pollen frequency of a given taxon is

roughly proportional to the abundance of the taxon in the surrounding

vegetation. Gross changes in the numbers of pollen grains and spores,

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commonly expressed as either the relative pollen frequency (RPF) or

absolute pollen frequency (APF), can be correlated with real changes

in thecomposition of the surrounding vegetation. Pollen spectre

give no direct information about the spatial composition and distri­

bution of the plant communities that comprise the vegetation of an

area, thus the reconstruction of ancient vegetation can only be in­

ferred by comparison with modern pollen studies.

More promising is the semiqualitative approach whereby geo­

graphically distinguishable vegetational units and associated physio­

graphic features (landform-vegetational zones) are characterized by

their contemporary pollen assemblages (Lichti-Federovich and Ritchie,

1965). Working with atmospheric pollen samples and surface samples,

these workers have shown that the several landform-vegetational zones

within the transition from boreal forest to prairie in southern Manitoba

can be distinguished by their pollen spectra. They applied knowledge

of the relationship between vegetation and pollen assemblages to

identify periods of grassland and deciduous forest in pollen diagrams

from what is now the southern boreal forest.

As more information becomes available on modern pollen deposition

rates and their relationship to the vegetation, it may become possible

to read the pollen deposition rate diagrams directly as a record of

the vegetational density on the landscape. Many refinements remain

to be made, and the lack of adequate testing of the basic ecological

assumptions continues to invite criticism from the skeptics of pollen

analysis.

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Recent investigations of modern pollen influx to lakes emphasizes

the complex relations between the pollen in-put and vegetational

formation (Ritchie and Lichti-Federovich, 1967; Tauber, 1967; Davis,

Brubaker and Webb, 1971; Richards, 1971; and Terasmae, 1973, 1976).

Although some pollen types in surface samples are apparently repre­

sented accurately in the relation to the abundance of the parent

formation, others are either overrepresented or underrepresented. The

causes of disproportionate representation are complex and many, those

of greater significance include:

1. variability in pollen production by different species,

2. the type of pollination,

3. ease with which pollen dispersal takes place,

4. preservation-differential susceptibility of pollen degrada­

tion by chemical and biological activity after deposition,

5. long distance transport of pollen and spores,

6. reworking and vertical displacement of palynomorphs, and

7. concentrating of palynomorphs by limnological processes.

High variance in modern surface sample influx measurements-

indicate that not all vegetational communities are distinguishable

from one another in the pollen record (Davis, 1967; and Ritchie

and Hare, 1971). Influx is not necessarily the same at different

sites surrounded by similar vegetation (Davis, 1967). However, influx

differences from region to region were shown to be significant by

Davis, Brubaker and Webb (1971) inferring that the vegetational differ­

ences at the community level are discernable in the fossil record.

The total pollen influx reflects the total pollen productivity

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of the vegetation, the influx assumes characteristic values in each

of the major plant formations, permitting identification of tundra,

boreal forest, mixed deciduous forest and deciduous forest (Ritchie,

personal communication). Transition zones and open areas are more

difficult to separate since pollen dispersal occurs well beyond the

range of growth limits causing measurable influx in regions where

trees are absent or scarce. The gradient of influx might provide

an estimate of the distance of a fossil site from the edge of the

major formation boundary. The pollen percentage interpreted with

the aid of modern analogues together with pollen influx data provides

a means for the reconstruction of the contributing vegetational

community.

The major limitation of influx data is that it is difficult to

distinguish changes in influx due to sedimentation from changes that

reflect pollen production of the vegetation. The sedimentary pro­

cesses affecting pollen influx are of great importance and should also

be taken into account (Cushing, 1964; Davis, 1968; and Craig, 1972).

Statistical methods for pollen analysis

Changing relative percentages of pollen grains reveal information

regarding the changing proportions of types of pollen in the pollen

rain. Relative pollen frequency (RPF) in this study is expressed as

the proportion of different taxa relative to the aboreal pollen (AP)

sum: (Equation 1) RPF = individual number of taxa/AP.

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The difficulty with compositional percentages is that they only

record the pollen types which are best represented in the sample;

however, it does not reflect the actual increase in abundance in the

pollen rain over time.

Because a pollen assemblage based on relative frequencies can

be derived from more than one vegetational type, vegetational inter­

pretation may be distorted by the presence of a prolific pollen

producer(s). Relative numbers also tend to exaggerate compositional

differences between samples. These difficulties may be aleviated by

determining actual numbers of pollen grains as done with absolute

pollen frequency (APF) determinations.

The main advantage of APF is that it can, at least in principle,

reflect a given floristic community with sufficient accuracy so that

the masking effects resulting from differential pollen production can

be overcome. APF reflects the change in pollen influx rates and has

proven to be a useful statistic for biostratigraphic correlations

(Faegri and Iversen, 1975). APF permits the consideration of pollen

curves as independent variables rather than as interdependent per­

centages or proportions. Absolute pollen analysis involves the rela­

tion of the number of fossil pollen grains to a unit volume or weight

of sediment, in order to compare throughout the profile the number of

grains deposited per unit area in unit time: (aquation 2) APF = influx 2 ^ concentration grains/ml grains/cm /yr grains/cm^ or or gram grams dry weight of sediment

In order that the vegetational community be reflected with any

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degree of accuracy, it is necessary to standardize the pollen count

on a meaningful factor, independent of the rate of pollen production.

The most meaningful standard is the sedimentation rate (Davis and

Deevey, 1963). Permitting APF to be expressed as "grains per unit

area per unit time." This however contains the assumption that the

sedimentation rate can be interpolated accurately between radio­

carbon dates.

Two commonly used APF methods are the Exotic Pollen Method

(Benninghoff, 1962; Waddington, 1968; and Stockmarr, 1971) and the

Weighing Method (Traverse and Ginsberg, 1966). In the weighing

method, a known weight of sediment is treated, the resulting con­

centration is weighed, a slide is prepared and a fraction of the

concentration is observed. The proportion of the weight of concen­

trate scanned under the microscope to the weight of treated sample

gives the number of grains per unit weight of sediment.

The more common technqiue to determine the actual number of

pollen grains in quantitative samples is based on the addition of a

pre-determined quantity of pollen added to a unit volume or weight of

sediment before preparation. Erdtman (1943) first made the pollen

counts with a haemocytometer. The Aliquot Slide Technique was intro­

duced by Muller (1959) with prepared standard drops taken from a

glycerine-alcohol suspension of the pollen samples. In 1962,

Benninghoff proposed the addition of a known quantity of exotic grains

(spike) be used to estimate the concentration of indigenous grains.

Although any of these methods can be used to calculate absolute

pollen data, each depends upon the removal of an exact quantity of

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material from the sediment core. It is well known that weight can

be determined more precisely than volume, but because sediments

differ greatly in density, sediment weight is not always useful as a

substitution sediment volume. Only APF with respect to dry weight

can be expected to be a meaningful and repeatable measure of absolute

pollen frequency unless time-stratigraphic control is both rigorous

and extensive (Faegri and Iversen, 1975).

Standardized pellets produced by Stockmarr (1971) permit the

easy addition of pre-counted exotics to the sediment being processed,

simplifying the once tedious and exacting procedure involved in

absolute pollen analysis. The exotics are easily recognized, behave

as do the native pollen and spores during the processing and can be

supplied in large quantities, increasing statistical precision.

There are some statistical errors involved with APF determina­

tions; including the error inherent in the datings and the fact that

the rate of sedimentation is never constant throughout a core between

dated levels. In addition, the repeatability of sampling may be a

major factor in the validity of a measure (Fletcher and Clapham, 1974).

Confidence intervals for absolute pollen values of pollen concentra­

tion and pollen influx have been constructed to evaluate the precision

of the recovery of native grains to exotics added (Bonny, 1972; and

Maher, 1972). It should also be mentioned that factors such as poor

pollen preservation, faulty laboratory technique or misidentification

of of grains are bound to result in misleading diagrams and interpre­

tations which are unreliable.

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Construction of the pollen diagrams

Relative pollen frequency (RPF) percentage values and pollen

concentrations were calculated with the aid of a computer program

adapted by Chris McAtee at Brock University. RPF method of calcu­

lation permits direct comparison with published data. RPF percentage

values were based on the portion of individual taxa to the arboreal

pollen (AP) sum: (equation 3)

RPF = number of population/sum of AP.

The pollen concentration was calculated from the formula;

(equation 4)

pollen concentration = exotics added X fossil grains counted (grains/gram) exotics counted gram dry wet sediment

using the ratio of exotics added to exotics counted times the ratio

of the fossil grains counted per gram of dry weight sediment

(Benninghoff, 1962; Stockmarr, 1971; and Davis, 1969).

Graphical representation of the total pollen influx through time

needs accurate concentration values and reliable sediment rates (Davis,

1969). (equation 5)

pollen influx = pollen concentration X sedimentation rate 2 3 grains/cm /year = (grains/cm or grains/gram dry weight)x(cm/year) or grains/g dry weight/year

Areal extent of influx can be calculated from the weight of

sediment if the bulk density is known (Davis, 1969) by the equation

below:

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(equation 6)

influx = concentration X bulk density ^ grains/cm^ grains/gram wet weight/cm dry weight or measure specific volume (cm )

Sedimentation rates were approximated for Oakland Bog with the

use of inferred pollen time-stratigraphic levels and radiocarbon

dates (Table 5). Where sedimentation rates have been determined,

Davis (1969) has demonstrated that relatively stable pollen concen­

tration in the sediment can be converted to an approximation of the

number of pollen grains contributing to the fossil pollen rain 2 deposited in time over a specific area (grains/cm /year). Only the

total pollen influx was approximated for Oakland Bog since poor

exotic recovery limited the precision of calculated pollen concen­

trations and too few radiocarbon dates were available to calculate

reliable sedimentation rates.

The pollen zone

The pollen zone is a sequence depicted on a pollen diagram that

is characterized by its flora (Faegri and Iversen, 1975). The com­

position of the flora is dependent upon four interacting factors:

1. geography,

2. climatic developments,

3. successional stages of the vegetation, and

4. ecological conditions at the site.

In principle pollen zones can be correlated from region to region

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and through time. Similar climates may be inferred from pollen zones

that are not alike in pollen composition. Depending on the localities

relationship to glacial vegetation refugia, distinguishing pollen

spectra may vary with migration rates and local soil and topographic

conditions.

Pollen zones are presently subjective delineations of pollen

diagrams chosen by changes in AP and NAP frequencies. Comparing the

fossil pollen spectra with modern pollen rain from known vegetational

assemblages whose climate is known, ancient climates can be inferred.

Current research by Ritchie (personal communication) may stan­

dardize the zoning of pollen objectively with computerized multi-

component analysis. The problem of interpretation still, however,

retains a personal bias.

Pollen Results

The pollen stratigraphy from the sediments of Oakland Bog (site I)

is illustrated as the relative pollen frequency (RPF) (Plate 2). The

RPF pollen sum is the total aboreal pollen (AP), calculated in this

study as:

AP = conifer pollen + deciduous pollen + Alnus + Corylus/Cornus + Salix.

This method of calculating the pollen percentages permits the direct

comparison of the pollen spectra with previously published palynological

investigations from the northeast

The pollen diagram from Oakland Bog (Plate 2) has been subdivided

into three pollen zones; each of which represents a pollen sequence in

the sediment characterized by a distinct vegetational assemblage. This

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tripartite zonation of the pollen assemblages found recorded in the

sediments of Oakland Bog resembles that of previously published Great

Lakes pollen stratigraphy (Sears, 1948; Benninghoff, 1961; Kapp and

Gooding, 1964; Ogden, 1966, 1969; Waddington, 1969; Wright, 1971;

Webb, 1974; Williams, 1974; and Bailey personal communication).

The composition and delineation of each pollen zone are further

described under the following separate section headings.

Zone I_ (interval 381cm-275cm)

The pollen assemblage in zone I is characterized by the dominance

of Pices (spruce) and herbs (non-aboreal pollen (NAP)). Spruce com­

prises up to 80% of the pollen grains counted in the basal portion

of the zone, declining irregularly to less than 30% at the upper limit

of the zone. Picea mariana (Black spruce) and Picea glauca (White

spruce) are both recognized as contributing to the spruce pollen

(Benninghoff, personal communication and Terasmae, personal communica­

tion) . NAP fluctuates irregularly between less than 5% and 60% with

Cyperaceae (sedge) pollen and the composites being the major contribu­

tors.

Pinus (pine) pollen gradually increases (10% to 30%) through zone

I with a sharp peak from 310 to 311 cm of 70% to 60% respectively.

Only Pinus banksiana (Jack pine) and Pinus resinosa (Red pine) have

been observed in zone I, but since they are difficult to distinguish

from one another and have similar ecologies they are considered as

one entity in the pollen spectra.

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Salix (willow) is best represented in the middle of zone I pollen

profile, where it reaches the greatest frequency from 20% to 40%.

There is also a scattered representation of other tree pollen, herbs,

and aquatics within zone I. Many spores were noted in zone I, par­

ticularly from the basal sediments.

The lowest pollen concentrations (Plate 3) and derived pollen

influx values occur in zone I. The pollen influx (Plate 4) averages

2300 grains per dry weight gram, ranging from 285 to 6554 grains per

dry weight gram. The basal pollen grains are poorly preserved, often

broken or corroded. Preservation improves remarkably at the top of

zone I, remaining excellent through the remainder of the marl.

Zone II (interval 275cm to 180cm)

Zone II is delineated by the abrupt decrease in Picea (spruce)

pollen and a corresponding sharp rise in Pinus (pine) pollen from

20% to 80%. NAP is generally less than 5%. A 20% Abies (fir) peak

is recorded midway in zone II (240cm) along with a minor decrease in

the pine curve.

Quercus (oak) is the only hardwood tree pollen to attain any

significant count in zone II. The oak grains are larger in zone II

and III as compared with the oak identified in zone I (Table 7),

suggesting a change in oak species. A few Castanea (chestnut) pollen

grains were counted and Populus (poplar) pollen is first noted in

zone II (Plate 2).

The pine during the beginning of zone II is either Jack and/or

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Red pine; however, at the 230cm level, Pinus strobus (White pine) is

found in the assemblage. Increasing to as much as 50% of the total

pine pollen, White pine becomes a significant element of the flora

in the rest of zone II.

The pollen concentration and pollen influx values reach their

highest values in zone II (Plate 3). The average influx in zone II

is approximately 12,000 grains per dry weight gram, ranging from

3470 to 39,501 grains per dry weight gram (Plate 4). Zone II records

the greatest pollen production in the units examined.

Table 7 Oak pollen grain sizes

ZONE______small (24^0______large ( 3 6 . ^ 0

I abundant none

II common common

III none abundant

Zone III (interval 180cm to 41cm)

Zone III represents a time interval dominated by Quercus (oak)

pollen. The lower boundary of zone III is distinguished by the decline

of pine pollen from nearly 80% to 30% and an increase of oak pollen

from 20% to 40% (Plate 2). Fagus (beech) pollen is first observed in

the lower portion of zone III, becomes a significant component (5% to

10%) in the pollen profile above the 80cm interval. In addition, Acer

(maple), Ostrya/Carpinus (ironwood), Corylus/Cornus (hazelnut/dogwood)

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and Carya (hickory) also have a significant representation (greater

than 5%) in the upper part of zone III. The lower boundary of zone

III is also marked by a sudden increase in NAP. NAP values rise, but

do not attain the high percentages recorded earlier in zone I, averag­

ing a nearly constant 20%.

The oak pollen in zone I were relatively small, zone II had a

mixture of large and small oak pollen grains, but in zone III only

the large pollen type is present (Table 7). One Ephedra (morman tea)

pollen grain and one grain of Tsuga (hemlock) pollen were counted on

slide C-2 at the 162cm interval.

The pollen concentrations are somewhat higher than those of zone I

but significantly lower than zone II (Plate 3). The average influx

of pollen grains in zone III was approximately 7,700 grains per dry

weight gram. Zone III has the greatest variance in influx, ranging

from 1,000 to nearly 12,000 grains per dry weight gram.

Distribution of additional palynomorphs

Throughout all three of the pollen zones there is a patchy distri­

bution of herb pollen (generally less than 3%) including: Thalictrum

(meadowrue),, Rosaceae (rose family), Ranunculaceae (crowfoot family)

and Plantago (plantain). Monolete fern spores are present throughout

the core, reaching their greatest representation (40%) in the middle

of zone I.

Aquatics counted in the core, averaged less than 3% throughout

the pollen zones. The total aquatic sum reaches a maximum of 5%

between the top of zone I and the middle of zone II. Aquatics counted

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at site I include: Myriophyllum (milfoil), Typha (cattail), Nuphar

(yellow pond-lily), Nymphaea (white pond-lily), Potamogeton (pond-

weed) , Sagittaris (arrowhead) and Utricularia (bladderwort).

Cyperaceae (sedges and rushes) include upland variety and wetland

species. The Compositae were identified as Artemisia (wormwood)

Ambrosia (ragweed) and other. Together the Cyperaceae and Compositae

dominate the NAP sum at site I. No significant accumulation of

Gramineae (grasses) or Chenopodiaceae/Amaranthaceae (goosefoot/amaranth

families) were recorded in the site I pollen slides but they are

present throughout the profile.

Cupressaceae (Taxus and Juniperus family) has a patchy distribution

of generally less than 1%. The hardwood tree pollen counted included

minor amounts of Celtis (hackberry), Juglans (butternut/walnut) and

Alnus (alder).

Fern spores are found throughout the core, reaching significant

values (10% to 40%) only in the upper portion of the first pollen

zone (I). Fungal spores, not included in the pollen count are found

in large numbers especially in the basal portion of pollen zone I.

The unidentified group includes those grains which are too broken,

corroded or obscured for recognition or unknown to the analyst.

Discussion of Pollen Zones

Zone I spruce-herb pollen zone

Elements of tundra and open spruce parkland are represented in

the basal sediments of Oakland Bog's pollen zone I (Plate 2). Low

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pollen influx values indicate tree flora was sparce on the local

landscape and high RPF values for spruce infer that the pollen rain

was influenced by the long distance transport of pollen grains (Nichols,

1975). Four possible vegetational assemblages have been suggested

for similar data in the literature:

1) Tundra vegetational assemblages characterized by high values

of locally produced Artemi sia (wormwood), Cyperaceae and Ericaeaceous

pollen and records the long distance transport of over produced spruce

pollen. Dwarf shrubs and trees are scattered across the landscape but

the modern pollen rain is dominated by NAP (Kapp and Gooding, 1964;

Davis, 1967; Ritchie and Lichti-Federovich, 1967; Lichti-Federovich

and Ritchie, 1968; and Williams, 1974).

2) Tundra/Boreal Forest transitional vegetation assemblage also

have high NAP values and large spruce pollen values. Modern surface

samples from Canada have shown this transitional assemblage to take

on vastly different profiles depending on local conditions (Wright et.

al., 1963; McAndrews, 1966; Ritchie and Litchi-Federovich, 1967; and

Ritchie and Hare, 1971).

3) Prairie/Boreal Forest transitional vegetational assemblage

has considerable NAP dominated by Artemisia, Cyperaceae, Graminae and

Chenopodiaceae/Amaranthaceae. This assemblage is difficult to dis­

tinguish from the tundra/boreal transition vegetation since both are

open communities strongly influenced by the proximity of the boreal

forest (Cushing, 1965; Litchi-Federovich and Ritchie, 1965, 1968; and

Ritchie and Hadden, 1975).

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4) Parkland-Open Spruce Forest vegetational assemblage records

high NAP values but tends to have a better representation of hardwood

pollen (particularly Quercus, oak). Local variations in the open

community are characteristic since patchy distribution of trees in­

fluences pollen rain (Benninghoff, 1961 and Miller, 1973).

Each analogue has its own limitations, and the merits will be

further examined by comparing equivalent stratigraphic levels from

other fossil pollen records and modern pollen surface samples with the

pollen profiles from zone I at Oakland Bog. The fact that satisfactory

analogues cannot be found for the late-glacial vegetation provides

a challenge for evaluating the complexities of ecological succession

on deglaciated terrain during a phase of pronounced climatic change.

The fossil pollen spectra from Oakland Bog most likely represents

a mosaci of several coexisting vegetational assemblages. Relatively

high NAP values (30% to 60%) are dominated by Artemisia and Cyperaceae

pollen. The prominance of Artemisia, 20% of the NAP value, may indi­

cate either tundra (cold) or prairie (dry) conditions at Oakland Bog

during the late-glacial. Considering the ecology of Artemisia and the

availability of glacial meltwater, the basal sediments at Oakland Bog

suggest the prevailing conditions were cold and perhaps frozen in

places. The particular Cyperaceae pollen at Oakland Bog represents

upland vegetation as opposed to marginal marsh typical of parkland or

tundra conditions, now found in northern Canada (Ritchie and Lichti-

Federovich, 1967 and Terasmae, personal communication). The small

size variety of oak pollen is a conspicious element in zone I suggesting

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the local source was either dwarf and/or living in a stresses environ­

ment (Terasmae, personal communication).

The late-glacial vegetation recorded in the lacustrine sediments

at Oakland Bog is most similar to modern surface samples from northern

Canada (Table 8) (Terasmae, 1964, 1965, 1967, 1968; Bartely, 1967;

Ritchie and Litchi-Federovich, 1967; Birks, 1973; and Nichols, 1975).

However, the Ericaceous pollen common to the subarctic and arctic

tundra is not recorded at Oakland Bog which suggests tundra as it

is known in northern Canada did not exist when pollen was being

deposited in the early lake stages of Oakland Bog (Love, 1959 and

Terasmae, 1976).

The particular kind of pioneer vegetation at Oakland Bog was

determined by the availability and aggressiveness of the species close

at hand. The preglacial and postglacial migrations are two very dif­

ferent matters. In the latter case the plants have a chance of rapidly

advancing over freshly denuded ground, but in the former they must

overcome the resistance of an already established vegetation while at

the same time experiencing a growth retardation due to the deteriorat­

ing conditions produced by the onset of glaciation.

As a matter of fact, Love (1959) found evidence that living trees

(southern hardwoods) were overrun in southern Ohio, indicating that

there could not have been a long time lag between a relatively warm

climatic period and the arrival of the glacier there. Lawrence (1958)

on the other hand has been able to demonstrate that a period of 170

years was necessary after the ice melted for a mature Sitka spruce

forest to develop at Glacier Way, Alaska.

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Table 8 Modern Pollen Rain Data from Canada

Vegetational community Location Reference

tundra Northern Canada Ritchie & Lichti- Federovich, 1967

Northern Canada Terasmae, 1965

Northern Quebec Terasmae, 1967

Arctic Quebec Bartely, 1967

Western interior Lichti-Federovich of Canada (north) & Ritchie, 1968

Northern Canada Birks, 1973

Northern Canada Nichols, 1975

forest/tundra Northern Canada Ritchie & Lichti- Federovich, 1967

Northwest Canada Nichols, 1975

spruce woodland Northern Canada Ritchie & Lichti- Federovich, 1967

boreal forest Ottawa, Canada Terasmae & Mott, 1964

Northern Great Terasmae, 1968 Lakes region

Western interior Lichti-Federovich of Canada & Ritchie, 1968

mixed forest Eastern Ontario King & Kapp, 1963

parkland Manitoba, Canada Lichti-Federovich & Ritchie, 1965

Western interior Lichti-Federovich of Canada & Ritchie, 1968

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The spruce-herb pollen zone is often subdivided in many pollen

diagrams, in part due to attempts to correlate vegetational sequences

with inferred late-Wisconsinan ice-margin fluctuations. The erractic

nature of both the tree and herb pollen curves from Oakland Bog make

such interpretations difficult.

The irregular fluctuations in the pollen curves at Oakland Bog

are most likely associated with the progressive colonization and

establishment of the local vegetation. The fine details of any climatic

change associated with glacial activity are incomplete. A better per­

spective of the vegetational development at Oakland Bog can best be

achieved by examining the pollen stratigraphy from the surrounding

region.

The low pollen influx and relatively high, irregular herb pollen

values supplemented by macrofossil evidence from the late-glacial

of New England are interpreted by Davis (1965, 1967b, 1969) to represent

tundra such as found in northern Canada today. The occurrence of

macrofossils of tundra plants in late-glacial deposits from northeastern

Minnesota suggests the existence of at least a narrow area of tundra

between the proglacial lakes and the late-glacial boreal forest (Watts

and Winter, 1965). Late-glacial pollen stratigraphy from other sites

in Minnesota suggest treeless to open spruce woodland bordered the

retreating ice-sheet (Wright, 1974). The conditions were also appar­

ently favorable for light-demanding grasses, herbs and shrubs (Love,

1959, 1970). Only the Salix pollen curve at Oakland Bog reaches

maximum values in zone I, indicating willow stands on the surrounding

slopes and shores but grasses are poorly represented in the pollen

profile.

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Compared to modern pollen in studies from the tundra of northern

Canada, pollen profiles from the late-glacial in New England tend

to have less spruce while the localities in the midwest have more

spruce. Cyperaceae RPF values are similar in the late-glacial of

New England and the midwest but are lower than in the modern tundra

pollen rain, suggesting the late-glacial is only reflecting tundra­

like conditions (Wright, 1964; Ogden, 1965; Davis, 1967b; and Terasmae,

1976).

The spruce dominated the vegetational assemblage in the northeast

15,000 to 12,000 years ago; however, this assemblage was clearly not

of uniform composition across the region of New England to Minnesota.

The most obvious difference is the presence of high values for Pinus

pollen in western New York and New England and their absence from

sites in Michigan, Wisconsin, and Minnesota. Apparently pines were a

part of the late-glacial vegetation in the East, but did not occur in

the contemporaneous vegetation of the Midwest. The available data

(Wright, 1964, 1968) indicate that the Appalachian region served as

a full and late glacial refugium for the three main pine species.

Pinus banksiana (Jack pine) Pinus resinosa (Red pine), and Pinus

strobus (White pine), which participated in the revegetation of the

glaciated Northeast.

The relative numbers of temperate deciduous tree pollen types

also vary from site to site within the Great Lakes region. An accurate

assessment of the variability in terms of climate, however, depends

in part on a detailed knowledge of the modern pollen rain from boreal

forest and woodland, the forest-tundra transition, and the tundra

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itself. Also, it must be kept in mind that modern analogues for

certain late-glacial pollen assemblages may never be found because

the vegetation which produced them may have been a mixture of species

brought together by differing migration rates and may thus represent

chance combinations of species which coexisted for varying periods of

time following the withdrawal and disappearance of the ice.

The regional stratigraphy of New York contrasts with the situation

west of the Great Lakes, where significant amounts of pine pollen are

not found in the profile until near the end of, or following the spruce

zone (Wright, 1964, 1968, 1969; Miller, 1973). Pollen diagrams from

southwestern New York (Miller, 1973) closely resemble the Oakland Bog

pollen diagrams in terms of diversity and frequency. A comparison

of the pollen profiles from a small region in southwestern New York

records divergences from the basic pollen profile pattern implying

the occurrence of localized park-tundra vegetation such as suggested

for Oakland Bog prior to the development of a regional spruce dominated

woodland. Once spruce trees became established, the total picture

most likely resembled open boreal woodland such as that Which occurs

in the subarctic of northern Quebec today (Terasmae and Mott, 1965;

Ritchie and Litchi-Federovich, 1967; and Nichols, 1975).

There is at present no pollen analytical evidence of a true

tundra element having been recorded in the late-glacial of the midwest

(Wright, 1964, 1971; Watts and Winter, 1965; Brinks, 1973; and Williams,

1974). Instead, the early spruce-herb pollen sequence is commonly

interpreted as tundra-like. Ericaceous pollen grains and pollen from

flowering plants common to the arctic and subarctic tundra have not

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 90

been found in the interior regions of the continental ice-margin (Fries,

1962; Wright, 1964; and Davis, 1967). Some arctic species were how­

ever able to exist in unique ecological sites such as along the bluffs

of Lake Superior and exposed river banks (Terasmae, 1967).

If a narrow belt of tundra did exist around the ice-margin per­

haps it either 1) lasted a short time and was not recorded in the

pollen that was deposited in ice-blocked basins, or 2) tundra-like

conditions existed but only in a patchy distribution along the parkland-

boreal elements. The record may also be incomplete because of the

lag between freeing the depression from ice, allowing deposition of

pollen from the invading pioneer vegetation which had a low produc­

tion of pollen grains and spores. The fresh, open, "humus-free" soils

left by Pleistocene glaciers most likely had a significant influence

on germination. Even from a very early period, pollen grains have

been found of plants which were decidedly not arctic in their require­

ments (Love and Love, 1963).

Scientists have disagreed over the possible types of vegetation

which first grew along the front of the retreating glacier. The

character and width of the transitional belt between glacier ice and

forest is highly debated. One of the interesting problems in Pleis­

tocene pollen studies is whether the Laurentide Ice Sheet during the

Wisconsinan full-glacial completely split the Boreal Forest and in­

vaded the prairie or whether it was bordered by tundra and/or forest

along the front.

Maps prepared by Martin (1958) showing the extension of tundra

or taiga westward from New England to Minnesota are highly speculative

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 91

west of Pennsylvania since they were only based on two pollen diagrams,

both from southern Michigan (Andersen, 1954; and Zumberge and Potzger,

1956). Andersen’s (1954) pollen diagram is controversial because

of a redeposition problem, having no radiocarbon dates nor any APF

values. Although no suggestion of tundra is indicated by Zumberge

and Potzger (1956), their diagram is also unreliable since detection

of NAP was inadequate.

Martin’s map (1958) of Michigan during the valders time showed

tundra or taiga covering all of the lower peninsula. On the other

hand, by comparing present day tree lines with climatological and

meterological data, Manley (1955) suggested that a forest could have

been established within 80 miles of the ice-margin. Fitting (1970)

describes a tundra-like zone in northern Michigan with richer flora

and fauna than found in the modern arctic tundra due to richer soils

and warmer summer temperatures.

The basal portion of zone I at Oakland Bog records pollen de­

posited during the Cary-Port Huron Interstadial. The region to the

south of the study area was characterized by high NAP values, rela­

tive spruce pollen values between 30% and 50% and low depositional

rates (Kapp and Gooding, 1964; Ogden, 1965, 1969; Garrison, 1967;

Williams, 1974; and Shane, 1975). Wayne and Zumberge (1964) saw no

indication of tundra in Illinois but pollen and mollusks from inter­

till siltbeds suggest a narrow tundra-like zone had existed at the

ice-margin. Intensive frost-action recorded as polygonal features

have also been discovered near ice-front margin positions in western

Indiana (Wayne, 1966).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 92

The Wisconsinan ice-sheet was not headed by arctic flora in the

interior of the North American continent, because the ice developed

in such a way as to separate the arctic flora from the area to the

south of it (Love, 1959). The arctic flora was unable to migrate

easily into an already closed community. A reconstruction of the

distribution of full-glacial biomes during the late Wisconsinan

(Fig. 15 and Table 9) indicate an azonal displacement of vegetation

(Whitehead, 1973; and Bernado and Webb, 1977). The vigor and adapt- >

ability which allowed spruce to migrate from its unglaciated refugium

south of the ice-margin is readily demonstrated from studies by

Wright and others (1974), recording the rapid invasion of spruce onto

deglaciated terrain in Minnesota during the late-glacial. The wide­

spread distribution and tolerance range of spruce today, further

supports its adjustment to the conditions during the late-glacial and

post-glacial times. Vegetation of unglaciated regions to the south

of the ice-margin are reported to have contained both boreal and

hardwood forest elements at the time of the full-glacial (Gruger, E.,

1972; Gruger, J., 1972; Bryant, 1975; King, 1977; and Wright, 1977).

Southward draining late-glacial waterways provided the pioneer­

ing vegetation easy access to the deglaciated terrain. A definite

south to north vegetational zonation has been recognized as time-

transgressive during the late-glacial (Wright, 1964, 1971; Terasmae,

1967; Cushing, 1967b; Davis, 1967, 1969, 1977; Whitehead, 1973;

Bryant, 1975; Webb and McAndrews, 1975; King, 1977; and Bernado and

Webb, 1977).

As the glacier melted back across the Great Plains and the Great

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 93

Drift Border I #1# LAURENTIDE ICE SHEET Tundra and / p. Boreal Woodland 4 0 ° S

'13 ■ y0 / Borealour mil Fores* vj' a ' • • / Ay ' 7 J (Spruce) ^

; - V 7 ------1------• |.g '; 0 5 00 Boreal For'est !••• A ; "J km (Pine) f • ' .30° S D-eciduous** Communities 1

Full-Glacial 10 7 5 ° W Shoreline \ \

Figure 15. Reconstruction of full-glacial vegetation. Key: 1, Marsh (Martin, 1958); 2, New Paris (Guilday et al., 1964); 3, Buckles Bog (Maxwell and Davis, 1973); 4, Round Glade (discussion by Martin in Guilday et al., 1964); 5, Shenandoah Valley Ponds (Craig, 1970); 6, Chesapeake Bay borings (Harrison, et al., 1965); 7, Dismal Swamp (Whitehead, 1974); 8, Roakyhock Bay (Whitehead, 1973); 9, Bladen Lakes ( Frey, 1951, 1953, 1955, Whitehead, 1963, 1964, 1967); 10, Lake Annie (Watts, unpublished in Whitehead, 1973); 11, Bartow County ponds (Watts, 1970); 12, Southern Illinois (E. Gruger, 1972); 13, Missouri Ozarks (King, 1973); 14, Northeastern Kansas (J. Gruger, 1977).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 94

Lakes-St. Lawrence flanks, the vegetational zones shifted. The

uppor portion of zone I from Oakland Bog records an increase of pine

and hardwood pollen at the expense of spruce and sedge pollen, indi­

cating the shift from an open spruce dominated community to a pine

woodland as the climate is less moderated by the glacial environment.

Fossil pollen extracted from the marl in the cranial cavity of

a woodland musk oxen (Symbos cavifrons (Leddy)) records an open spruce

woodland some 13,000+600 years B.P. in Climax, Michigan (Kalamazoo

County) (Benninghoff and Hibbard, 1956, 1961). Another mastodon

found in nearby Scotts (Kalamazoo County) lived in a pine dominated

forest at least 11,000 years ago (Semken, Miller and Stevens, 1964).

Other local findings of the forest browsing mastodons, herbivrous

grazing mammoths and giant beavers in association with their respec­

tive pollen spectra suggest a variety of vegetational assemblages

coexisted including upland boreal forest, wetland and parkland and

that the mosaic community was transitional (Benninghoff, 1961; Skeels,

1962; Dorr and Eschman, 1970; and Brewer, 1972). The landscape in

Kalamazoo County 13,000 years ago had the aspect of the boreal forests

that now exist north of Lake Superior (Benninghoff, 1964; Terasmae,

1967, 1968). The spruce forest contained local openings on the

gravelly flood plains, rich in herbs and with stands of willow.

Pine pollen begins to dominate over spruce at the 300cm level

at Oakland Bog. At approximately the same time-stratigraphic level

at Wintergreen Lake, some 20 miles north of the study site, Jack/Red

pine and birch began to replace spruce in the vegetation 11,000

years B.P., interpreted by Bailey (personal communication) as

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coincident with the retreat of Valders Ice. Increasing pine pollen

values at the expense of spruce in zone I at Oakland Bog can be

interpreted as:

1) pollen production variability in a changing forest scene

2) blown-in pollen differences.

Absolute pollen studies suggest that the change in pine pollen per­

centages represents the deterioration of the spruce forest (Davis,

1967).

A wave of forest transformation began about 12,000 years B.P.

affecting the boreal forests or parkland forests that had covered the

entire midwest during the Two Creek Interstade (Whitehead, 1973). By

10,000 years B.P. spruce no longer shows a continuous band of domi­

nance from east to west and attains its highest frequencies northward

in the wake of deglaciation (Bernado and Webb, 1977).

The vast differences in the pollen representation at equivalent

stratigraphic levels of neighboring sites in the midwest may be only

reflecting local variations rather than regional sensitivity. During

the time Oakland Bog was experiencing a gradual but continuous decline

in spruce pollen, Myers Lake in northern Indiana records a spruce

decline followed by a strong peak, interpreted by Frey (1959) to

represent the Two Creeks Interval of warmer climate and subsequent

Valders Ice advance. However, at Fox Prairie Bog in northcentral

Indiana and Bacon's Swamp in central Indiana, spruce was nearly absent

(Engelhardt, 1960) during the same time interval. On the other hand

at Sunbeam Prairie Bog, Ohio (150 miles south of the study site) spruce

accounts for more than 90% of the tree pollen during the Two Creeks

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 96

time (Kapp and Gooding, 1964). Tippecanoe Lake in northeastern

Indiana (Potzger and Wilson, 1941) and nearby Cabin Creek (Friesner

and Potzger, 1946) reported similar findings while just 25 miles to

the west at Reed Bog, Indiana, Griffin (1960) described a decline in

both pine and spruce pollen percentages during Two Creeks time. The

preliminary pollen stratigraphy of Sears (1942) and Potzger (1948)

in the midwest correlated vegetational shifts with climatic changes

associated with glacial events. Many of the interpretations may also

easily reflect the succession of vegetation as the climate gradually

warmed.

Wright (1964) and Whitehead (1973) examined the regional trends

of pollen data and concluded that the northeast developed patches

of closed forest like those of the modern boreal forest but they

were not regionally continuous. The pollen curves of Oakland Bog

and southwestern New York (Miller, 1973) indicate that the patchy

park-tundra in the late-glacial was succeeded by a spruce dominated

woodland while New England recorded a rise in pines and hardwoods.

In southern New England, the spruee-hardwood subzone was followed

by a spruce-fir subzone (Davis, 1965, 1967; and Davis and Deevey

1963). On the basis of absolute pollen counts, Davis (1967) con­

cluded that although the rise of hardwood tree pollen (especially

Quercus were most likely windblown from distant forests) may repre­

sent a warming climate, the fall of hardwoods was a reflection of the

sharp rise in pine pollen frequency. The pollen sequence was inter­

preted by Davis (1969) to be a response to a steadily warming climate

and a progressive change in the forest composition, starting about

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12,000 years B.P. correlating with the end of the Two Creeks Inter-

slade. The hardwoods were not a significant portion of the midwest

forest during that time thus are not sensitive indicators of climate

in the pollen profile.

High hardwood tree pollen anomalies in the northeast have been

used as further evidence for a late-glacial climatic oscillation

(Davis, 1967). The distinction between two limiting factors, mois­

ture and temperature, is important because alone they imply reverse

correlations with glacial events. Consideration of temperature and

moisture conditions is important for stratigraphic correlation of

vegetation-climate-glacial events which will lead to more precise

reconstruction of the paleoecology of the late-glacial and early

post-glacial. The importance of hardwood pollen may be interpreted

in more than one way so further consideration is necessary.

The maximal hardwood tree pollen percentage may not only reflect

an actual increase in the numbers of hardwood trees as postulated by

Davis (1967) but may also record a decline in local pollen produc­

tivity that was due to the replacement of the local forest by parkland

(West, 1961). Redeposition of hardwood tree pollen from older inter­

glacial deposits as a result of solifluction also suggests such an

interval is correlated with the cold (Andersen, 1954).

An alternative to the parkland interpretation comes from the

data published by Leopold (1956) and Wright and others (1963, 1974)

collected in the northeast and Minnesota. Both correlated the rise

of hardwood tree pollen percentage to the Two Creeks Interval. The

increased numbers of hardwood tree pollen was interpreted to reflect

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 98

an actual increase in production commonly associated with warmer

climate. However, increased production is not necessarily due to a

warmer climate. Increased temperatures also increase evaporation,

this may lead to drought conditions such as found in the prairie

environment. Subsequent to the decrease in the number of trees

associated with the dry prairie environment, an increase in tree

pollen frequency is correlated with a return to a cooler and moister

climate such as that presumably associated with glacial advance.

In light of the new appraisal of climatic implications consider

the problem at Gillis Lake (northeastern Quebec) (Livingston and

Livingston, 1958 ). The birch maximum within the herb zone at Gillis

Lake has been previously interpreted to represent a temporary ad­

vance of the forest margin into tundra in response to the Two Creeks

interval of climatic warming (Livingston and Livingston, 1958). Com­

parison of the late- to early post-glacial profile from Gillis Lake

with a modern analogue, surface-sample data from Labrador (Wiltson,

1964) the birch maxima has been reinterpreted to represent simply a

stage in succession from tundra to woodland, occurring during a time

of gradual climatic warming rather than a time of climatic oscilla­

tion correlated with glacial retreat and advance (Litchie-Federovich

and Ritchie, 1965). Local depositional conditions influenced the

pollen record at this site and should be a consideration at all sites

evaluated.

There is no evidence of reversals in the general trend of climatic

warming from Oakland Bog pollen profile in zone I. Throughout the

late-glacial and early post-glacial time the climate appears to have

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been cool and relatively continental without any clear record of

temperature oscillations correlative with the well documented Allerod

of northern Europe or other North American pollen profiles (Cushing,

1967).

The controversy concerning the recognition of glacial events

such as the Two Creeks and Valders Stages in the pollen profile and

their climatic implications may be a function of:

1. The vegetation's sensitivity to record the particular

intensity and duration of the climatic changes associated with such

glacial events.

2. The local conditions at depositional sites to record vege­

tational changes.

3. The modification of climates away from the direct influence

of the ice sheet.

The vegetational record substantiates the mathematical predic­

tions of Webb and Bryson (1972) that a non-glacial mode of climate

for the southern Great Lakes region became established 10,000 to

11,000 years B.P. The transition of vegetational assemblages in the

midwest are more directly related to differential migration rates.

The shifts in population dominance within the vegetational community

are not directly correlated with climatic changes associated with the

ice-margin front as noted in New England but rather the pollen spectra

records the influence of the glacial ice and relative water levels of

the predecessors to the Great Lakes which acted as barriers to immi­

gration in an already warming climate.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 100

Zone II the pine pollen zone

The top of the Spruce Pollen Zone from Oakland Bog and elsewhere

in the midwest and New England is marked by a sudden decline in spruce

pollen (Appendix 2). At Oakland Bog, pine replaces spruce as the

dominant element delineating Zone II, the Pine Pollen Period. The

post-glacial Pine Period Zone defined by Deevey (1939) has a fossil

pollen assemblage similar to the modern pollen rain of central Canada

(Ritchie and Litchie-Federovich, 1967) and northern Minnesota (Webb,

1974 and Webb and Andrews, 1975).

Spruce pollen declines from more than 50% to less than 5% over

a stratigraphic interval of only a few decimeters at Oakland Bog.

This is the sharpest vegetational transition described in the litera­

ture of any pollen stratigraphic level in North America, including the

level of forest clearance and agricultural development (Wright, 1964

and Ogden, 1967). The abrupt decline of spruce pollen has been docu­

mented as occurring approximately 10,000 years ago in the Great Lakes

region (Fries, 1962; Wright et al.; and Ogden, 1967).

Determination of the time involved in this transformation depends

on the rate of sedimentation which can only be measured through radio­

carbon dating. Compilations by Ogden (1967) and Wright (1968) have

indicated a range of 100 to 600 years for the spruce decline. Abso­

lute pollen counts by Davis (1967) in New England eliminates any doubt

that the pine pollen zone, in New England at any rate, represents a

true immigration of pine trees as the spruce forest deteriorated.

The spruce pollen decrease probably reflects a comparable change

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in forest composition in the midwest. Summers had probably become

too warm for the regeneration of spruce (Wright, 1964). Openings made

by fire or by blown down senile spruce trees along the southern mar­

gin of the spruce forest permitted available seeds to germinate.

Pines are similar to each other in most of their life-history

characteristics, but there are just enough differences in their en­

vironmental requirements to bring about significant ecological separa­

tions (Curtis, 1974). Pinus banksiana (Jack pine) is generally

assumed to be the species involved early in the Pine Zone because of

its more northerly distribution today, its ease in invading open

areas (especially after fires), its lighter seeds, and early maturity

and thus relatively migration. The pollen of Pinus banksiana (Jack

pine) and Pinus resinosa (Red pine) are difficult to distinguish and

their similar ecological demands permits grouping them together as

one pollen entity (Terasmae, personal communication). Jack/Red pine

are less shade tolerant than Pinus strobus (White pine) but can ger­

minate on drier sites with a lower nutrient content. White pine is

the most exacting with respect to the moisture and nutrient supply

required for optimum development (Fowells, 1965).

An Abies (fir) maximum is represented midway in zone II at

Oakland Bog. Balsam fir was growing suppressed in a spruce-dominated

woodland, deterioration of the spruce overstory allowed fir seedlings

and saplings to develop in the understory. The period during which

fir thrived must have been relatively short because its pollen disap­

pears from the count shortly after the maximum is reached. At present

Abies balsamea persists under dense forest cover but nearly full

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sunlight is needed for best development (Fowells, 1965). High fir

percentages near the end of the spruce zone occur over a wide area

in the northeast, although the peak is sometimes within the spruce

zone and other times into the pine zone (Deevey, 1943; Cox, 1959;

Davis, 1967b; Wright, 1968; Miller, 1973; and Bailey, personal com­

munication) .

Pine was not a major component at Oakland Bog during the late-

glacial nor anywhere else in the Great Lakes region (Ogden, 1967;

Wright, 1964; and Williams, 1974). The absence of pine appears to

be a result of its slow migration (Wright, 1964). If this is the

case, the unique spruee-hardwood pollen zone (which has no modern

analogue) found between the Spruce and Pine Pollen Zones in the post­

glacial did not reflect a major climatic shift. Instead, the spruce-

hardwood assemblage of the midwest represents an interval during which

hardwoods were most readily available to replace the failing spruce.

When the spruce forest began to deteriorate in southern Minnesota

some 12,000 years ago, pine was still confined to New England and the

Appalachian Highlands (Fig. 16) so it was primarily birch and alder

that succeeded at Madelia (Fig. 17) (Jelgersma, 1962). The wave of

destruction of spruce forests reached the southeast of Minnesota about

11,000 years B.P. as seen from the pollen profiles of Kirchner Marsh

(Fig. 17) (Wright, Wintert and Patten, 1974) and Lake Carlson (Fig. 17)

(Wright, Wintert and Patten, 1974). To the northeast, Weber Lake

(Fig. 17) (Fries, 1962) recorded the replacement of spruce pollen by

pine-oak-elm pollen at a radiocarbon date of 10,230. Farther north, c birch and ash were already present, so they flourished briefly before

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. RED,JACK PINE 0 HEMLOCK 0 BEECH © WHITE PINE © HICKORY @ CHESTNUT

Figure 16. Holocene migrations of ecotones (after Davis, 1974).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 104

C_QNIFER HARDWOODS

DECIDUOUS

PRAIRIE /

Figure 17. Index map of sites in Minnesota where postglacial pollen studies have been conducted. Key: 1, Ithica transect (McAndrews, 1966); 2, Myrtle Lake (Janssen, 1968); 3, Lake of the Clouds (Potzger, 1953); 4, Weber Lake (Fries, 1962); 5, Kotiranata Lake (Watts, 1967); 6, Horseshoe Lake (Cushing, 1967); 7, Cedar Bog Lake (Cushing, 1967); 8, Rutz Lake (Waddington, 1969); 9, Kirchner Marsh (Wright et al., 1963); 10, Lake Carlson (Wright et al., 1963); 11, Madelia (Jelgersma, 1962).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 105

the other hardwoods migrated up from the south (Watts and Winter,

1965; Wright, 1964). There were still favorable conditions for spruce

growth north of Horseshoe Lake (Fig. 17), so the forest composition

changed gradually rather than abruptly (Cushing, 1965).

The transformation of spruce to pine or hardwoods can reasonably

be attributed to the gradual warming of the climate resulting in

conditions apparently too warm for the survival of spruce seedlings

along the southern margin of the spruce forest. The sensitive outer

fringe of forests such as in the Nebraska Sandhills (Wright, 1968)

and southwest Minnesota (Wright, 1968) first felt the effects of

the warming climate 12,500 years B.P. and what replaced the spruce

depended on what seed sources were available nearby. Thermophiluos

deciduous trees immediately succeeded the spruce forest at Madelia

and Kirchner where pine had not interceded, reaching full development

9,700 years B.P. (Jelgersma, 1962; and Wright, Winter and Patten, 1963).

Oak, apparently was a component of the forests south of the boreal

woodland, migrated quickly with the warming climate and was soon joined

by elm, ironwood and other deciduous hardwood trees (Gruger, E. 1972)

at Oakland Bog and in southwestern New York (Miller, 1973). But

neither Oakland Bog and Wintergreen Lake (Bailey, personal communica­

tion) in southwestern Michigan nor southwestern New York (Miller, 1973)

display an intercedent hardwood prevalence between the decline of

spruce pollen and the dominance of pine pollen.

South of Oakland Bog, Myers Lake in Indiana also records the

disappearance of spruce pollen from the profile, followed by a sharp

rise in pine and finally the replacement of coniferous elements oak

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 106

and other hardwoods (Frey, 1959). The same coniferous to deciduous

hardwood transition can be recognized at Bacon Swamp (Engelhardt,

1960) and Kokoma Bog (Potzger, 1946) in central Indiana. This suc­

cession is again similar to that found in Ohio. Sunbeam Prairie

Bog, in western Ohio was reported by Kapp and Gooding (1964) to record

the replacement of conifers by deciduous trees (principally oak and

elm) at a radiocarbon date younger than 10,600 years B.P. Silver

Lake in west central Ohio, diverges from the spruce-pine-hardwood

deciduous succession in that the pine zone follows the spruce zone by

several hundred years with a Quercus (oak) maxima between (Ogden,

1966).

A factor in the immigration of pine across the Great Lakes region

from the Appalachian refugia were the Great Lakes and their prede­

cessors which changed dimensions during the late-glacial and early

post-glacial. During the Two Creeks Stage of the Wisconsinan ice

retreat, about 11,800 years B.P., all the present Great Lakes were

drained to low levels (Wright and Frey, 1965). The adjacent areas,

still covered with spruce, excluded the first invasion of the more

durable Jack/Red pine into a closed, healthy spruce forest but allowed

the immigration of these pines across the lowlands at the southern

end of Lake Huron (Terasmae, 1967).

To explain the subsequent rapid immigration of pine, Wright

(1968) postulated that the heavy seeds of pine did not have to travel

very far but rather the pines occurred in stand so small as not to be

reflected in the pollen rain but large enough to provide local seed

sources when the opportunity for expansion was provided.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 107

Jack/Red pine was prominent in the early Spruce Pollen Zone at

Douglas Lake, Cheboygan County , northcentral Lower Michigan (Wilson

and Potzger, 1943) and Vestaburg, central Lower Michigan (Gilliam

et al., 1967) but does not appear until the end of the late-glacial

at George's Reserve, Livingston County in southeastern Michigan

(Andersen, 1954) and at Oakland Bog and Wintergreen Lake (Bailey,

personal communication) in southwestern Michigan.

In the Valders Stage that followed, ice readvanced far down the

lake basins and lakes rose to a new high level. Both the lakes and

the ice lobes must have provided barriers to pine immigration from

the east. Final withdrawal of the ice from the basins about 11,000

years ago, associated with the climatic change which also caused the

deterioration of the spruce forests, opened the terrain north of the

Great Lakes to Jack/Red and later White pine immigration. Perhaps

this event also accounts for the abrupt increase in pine pollen west­

ward of Michigan.

To the south of Oakland Bog, barriers of a different nature pre­

vented the immigration of pine. The calcareous tills of western and

central Ohio were detrimental factors limiting the migration of pine

(Ogden, 1967). Pollen records from this region shows no "Pine Period",

and only a brief interval when the contribution of pine pollen ex­

ceeded 10% (Ogden, 1967). Some pines may have spread along the south

side of the Great Lakes perhaps as far west as Indiana, but they did

not occur in large quantities because of the greater importance of oak

in a climate that was already too warm for pines to successfully

compete with prairie elements. The differential migration rates of

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species best explains the basic pattern of post glacial succession

(Fig. 16).

An influx of 50% White pine by the middle of zone II at Oakland

Bog and a similar trend recorded at Wintergreen Lake (Bailey, personal

communication) in Kalamazoo County, southwestern Michigan and Pretty

Lake, Indiana (Williams, 1974) dated at 9,800 years B.P., reflect the

immigration of White pine from the east as the climate becomes wetter

(Fig. 16). White pine first appeared in quantity in the northeast

about 9,000 years B.P., some 2,000 years after the dominance of

Jack/Red pine (Davis, 1967b). It was not until 7,000 years B.P.,

that eastern Minnesota records the arrival of White pine (Wright,

1968). Further migration was limited by the eastward expansion of

the prairie and oak savanna which began 8,000 years ago in the upper

midwest (Davis, A., 1977). White pine finally reached Kirchner

Marsh, (Wright, 1968) 1,600 years B.P. but no pine rise has been

recorded at Cedar Bog Lake in northern Minnesota (Cushing, 1963).

Zone II at Oakland Bog represents a time of more or less complete

closure of the boreal forest, since the NAP is reduced to less than

5%. This was also a time of forest transition from boreal spruce to

deciduous hardwood. The climate was not only becoming warmer with

the retreat of the glacier but drier also, creating prairies in the

west and led to the Pinus maxima in Michigan. The top of zone II is

marked by an abrupt rise of NAP components, decrease in pine and an

increase in mosaic hardwood tree elements indicating a change to a

more open deciduous forest.

The pollen record of zone II is one of progressively changing

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 109

forest composition. Evaluation on a broad scale demonstrates that

forest zones did not migrate as entities in response to climatic

change but rather individual species moved independently (Davis, 1974)

(Fig. 16). Soils became progressively leached and depressions in­

filled favoring new lowland communities, complicating the migration

record and climatic interpretation.

Zone III the deciduous hardwood zone

The abrupt rise of pine in zone II is accompanied by slower rises

of Ulmus (elm), Ostrya-Carpinus (ironwood) Carya (hickory) and Quercus

(oak). Oak apparently succeeded pine at Oakland Bog (Appendix 2) as

greater warmth and dryness caused pine to decrease (Wright, 1968; and

Terasmae, personal communication). This transition took place about

the same time in the central midwest as evidenced by pollen diagrams

from Wintergreen Lake in southwestern Michigan (Bailey, personal com­

munication) and in the northeast and northwest Indiana (Williams,

1974).

Representation of Acer (maple), Fagus (beech), Juglans (walnut),

and Tilia (basswood) at Oakland Bog begin at the boundary between

zones II and III, along with an abrupt rise of NAP. Pollen influx of

the boreal species decreases, indicating as Davis (1969) suggested,

that the change in the pollen percentages are not merely a reflection

of dilution but represents a real decline in the number of boreal trees.

Regional diversity in equivalent pollen stratigraphic subzones

of the deciduous-hardwood zone III across the northeast record local

variations of the deciduous forest (Table 11).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 110

elm ostrya/carpinus prairie prairie 5 5 4

(southern) (Wright, 1968) l c l c l c Minnesota

prairie hickory (Sears, 1942) (southern) - beech Michigan

Table 9 hickory oak elm hickory beech oak maple oak - oak beech C (northern) (Ogden, 1966) 3e 3d 3C 3a 3b Indiana

savanna maple basswood basswood maple maple deciduous-hardwood zone III across the northeast. (northern) (Ogden, 1966) Cb Cc ca Ohio Correlation table of pollen stratigraphic subzones of the

elm hickory elm oak oak oak hickory hickory (southern) (Deevey, 1939) New England

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Ill

The beginning of zone III at Oakland Bog is marked by the two­

fold increase of oak pollen at the expense of pine (Appendix 2). At

this level and on, the oak pollen is comprised entirely of the large

grain variety suggesting only mature oak tree pollen was being depos­

ited. The oaks and herbs may have occupied the more xeric sites in

hilly, sandy areas around Oakland Bog which is also the case today

in southwestern Michigan (Wright, 1964, 1971; and Brewer, personal

communication).

Hemlock is notably absent from the pollen profile at Oakland Bog

and throughout the central midwest (Cushing, 1965; and Wright, 1976).

The comparable interval in New England contained much hemlock but no

hickory until 5,000 years ago (Davis, 1974, 1977) (Fig. 16). In

today’s Hemlock-White Eine Northern Hardwood Forest region of Michigan

and Wisconsin, hemlock pollen was weakly represented at the C-l strati­

graphic level (Deevey, 1939). Wilson and Potzger (1942) showed hemlock

reaching the Douglas Lake region of central Michigan early in the

oak-hardwood domination, contemporary with Deevey's C-l interval of

the East (Table 19). Hemlock is consistently part of the pollen

record in central Michigan above a radiocarbon date of 7982+250 years

B.P. (Gilliam et al., 1966) but it accounts for only 5% or less of

the pollen sum. West's diagram (1961) from Seidel Lake in eastern

Wisconsin similarly shows that hemlock appeared fairly early during

the period of oak domination that followed the Pinus maxima but never

exceeded about 3% of the total pollen until much higher in this section.

Western New York, north and central Michigan and northern

Wisconsin (Sears, 1935; Broecker and Farrand, 1963; Cushing, 1965;

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 112

Miller, 1973; and Bailey, personal communication) all record hemlock

in the pollen record at the C-l stratigraphic level, approximately

8,000 years B.P., but the dominance such as found east of western

New York is absent. During the same period, the prairie-oak savanna

expanded eastward in Minnesota (Wright, 1968). It has not yet been

established whether Wisconsin and Michigan were yet affected by the

drier, warmer climate but if they were this might explain the meager

representation of hemlock in the sediments accumulated during zone

C-l. That hemlock grows best in a cool, humid climate and is sen­

sitive to drought is a helpful clue in unraveling clamatic factors

where it is prevalent (Fowells, 1965).

When Minnesota was undergoing a xerothermic phase, western

New York appears not to have (Wright, 1968; and Miller, 1973). From

8,500 to 4,300 years ago, New York was apparently cooler and moister

than the midwest, which allowed hemlock to migrate in (Wright, 1968;

and Miller, 1973). The abrupt decline of hemlock 4,390 years ago in

southwestern New York (C-2 interval) signifies the warmer, drier

climate which effected Michigan and the midwest earlier allowed the

expansion of oak and hickory at the expense of hemlock in New England

(Miller, 1973).

Hemlock appears to have entered Michigan from the east, north

of Lake Erie, and not from the south through the wedge of prairie

vegetation commonly referred to as the Prairie Peninsula. The Prairie

Peninsula apparently acted as an effective barrier to the migration

of not only hemlock but beech as well (Benninghoff, 1963). This

theory is supported by the zone III pollen data from Oakland Bog and

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 113

Wintergreen Lake (Bailey, personal communication) in southwestern

Michigan, Silver Lake in western Ohio (Ogden, 1966), and Pretty Lake

in Indiana (Williams, 1974), all of which were comparable to the

modern surface samples from Clearwater Lake (Janssen, 1968) which

is situated in a predominantly xeric oakwoods in westcentral Minnesota.

The presence of the Prairie Peninsula proved a physical barrier

to the invation of deciduous forests, altered the species composition

of the invading elements and slowed the rate of succession from

pioneer oaks and hickories to more mesophytic hardwood species.

Once the warm and dry climate established the prairie regime, it was

difficult to dislodge (Benninghoff, 1964). The persistence of the

prairie was most likely a function of low winter precipitation and

summer droughts which hindered the development of seedlings by rigor­

ous competition for soil moisture with the many grass roots. The

streams and rivers were probably the main avenues of forest migration

into the prairie (Davis, A., 1977).

Pollen in the upper portion of zone III at Oakland Bog reflects

the establishment of more mesic species, Acer, Tilia, Fagus and

Juglans, dated at nearby Wintergreen Lake as occurring 8,500 years

B.P. (Bailey, personal communication). The change in the vegetational

community suggests that there was a gradual development of a moister

regional climate than that of the previous Pine Period.

The increase in frequency of Fagus (beech) and other mesic hard­

wood tree pollen along with a decline in NAP (Appendix 2) marks the

beginning of subzone III at Oakland Bog. This trend parallels the

collection of beech pollen elsewhere in the central Great Lakes region

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 114

7,400 years B.P. (Ogden, 1966, 1969; Williams, 1974; and Bailey,

personal communication).

The regional patterns of vegetation change in zone III appear

to have been associated with changes in the precipitation regimes

(Wright, 1964; Davis, 1969; and Bailey, personal communication).

Pollen of the more mesic species suggest an increase in moisture

accompanied the general trend of climatic warming although in the

case of beech, its immigration from the east was also a function of

the development of a protective forest cover of oak and hickory

(Benninghoff, 1963).

There is additional evidence from molluscan studies at Oakland

Bog (see Molluscan Section IV) and alteration between littoral and

planktonic diatom assemblages at nearby Wintergreen Lake (Bailey,

personal communication) to support the hypothesis that water levels

fluctuated somewhat, perhaps as a function of the moisture regime.

Corresponding changes in the abundance of angiosperm seed, leaves and

diatoms have been recorded at other sites in the western Great Lakes

during a warm interval referred to as the hypsithermal (8,000-4,000

years B.P.) (Watts and Winter, 1966; and Watts and Bright, 1968).

The earliest prairie was recorded as early as 10,000 years B.P. in

northeastern Kansas, corresponding to the general warming trend asso­

ciated with the replacement of spruce by pine in the central Great

Lakes region (Gruger, J., 1972).

Prior to the transition from lake to bog at Oakland Bog, lakes

in Minnesota were intermittently dried-up, permitting the spread of

herbs over the basin floor as evidenced by:

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 115

1. shallow rooted aquatics (Watts and Winters, 1966)

2. oxidation of pollen grains in dried beds (Watts and Winters,

1966)

3. badly corroded grains at levels corresponding to moss layers

of Drepanocladus sendtneri, which inhabits swamps, bogs and marl wet

places (Watts and Winters, 1966)

4. extremely high chenopod/amaranth values implying much drier

climatic conditions than present today in southeastern Minnesota

(Watts and Winters, 1966)

5. seed studies and supplementary diatom work (Haworth, 1972),

mollusks (Watts and Bright, 1968) and oxygen isotope ratios in mollusk

shells (Stuiver, 1970) all indicating increasing salinity at the time

of the prairie period.

Pollen diagrams from Kirchner and Carlson Lakes in southeastern

Minnesota are the first to show with any detail the progression of

the prairie invasion during post glacial time, reaching full expres­

sion some 7,200 years ago when herbs were favored over oaks (Wright,

1968).

Stratigraphically, the ragweed maximum recorded at Rogers Lake,

Conn. (Davis, 1969) is correlated with the Prairie Period of the Great

Lakes region. Progressive drying in interpreted by increased NAP and

oak pollen values at the expense of other hardwood tree pollen. This

hypsithermal or xerothermic interval appears to be time-transgressive,

west to east (Davis, 1967; Wright, 1976).

Different abundances of species prevailed in the Great Lakes

region as compared with New England once the hardwood transition began.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 116

Factors such as geology, topography, soil development and differen­

tial migration rates, indirectly related to, but nevertheless a

function of climate, appear responsible for the variation of vege-

tational assemblages in the northeast. It seems probable that even

in an identical climate, the vegetational assemblages would not be

the same across the northeast.

Pollen zone III records the development of hardwood forest

under a climatic regime warmer than the Pine Period but cooler than

present. The mesic species also suggest the latter part of zone III

recorded at Oakland Bog was wetter than the climate today. Unfor­

tunately the marl action examined at Oakland Bog included sediments

no younger than 6,000 years B.P. and the overlying peat had extremely

low pollen counts with poor preserved grains and had been disturbed

by agricultural activity thus proving useless for pollen analysis,

limiting the scope in time of the climatic development in south­

western Michigan.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 117

*r 0 ' N u fo rest

« « \ V\ V\ M i«Tip^rvf III // // * TO *■ \ *■ » z' HYPOTHETICAL * ' / ' / / / / 5,0 0 0 MARSH-BOG-FEN DETERIORATION d e c id u o u s PRAIRE-SAVANNA

lake ••

6 ,0 0 0

0 III 12 flourishing permanent

/MOIST • •• •• • COOL/ WARMING 8,0 0 0 MESIC HARDWOODS large establishment

9.00 0 9.00

/DRY s' II < lake E N RICH ME NRICH ME N E T H PINE PINE PERIOD s-N W ARMER /

10.0 0 0

SHALLOW PERMANENT SHALLOW

BOREAL FOREST

> ) 1 11.0 0 0 LAKE 1 /MOIST 5 '-7 ' WOODLAND 13.0 0 0 COOL/ PERMANENT ENRICHMENTI OPEN SPRUCE s t , 14.0 0 0 0 14.0 1 m o / r - Y TUNDRA PIONEER A INTERMITTENT INTERMITTENT 16,0 0 0 c o l CREEK -PONDED CREEK PARKLAND-

Figure 18. Summary of pollen zones, molluscan stages and inferred climatic conditions during

stage

«. b Z uu z *

CLIMATE the development of Oakland Bog, SouthwesternMichigan.

I o MOLLUSCA 8 1 Ul s? - Z ¥ 0 % 5 | « >. ZONE POLLEN POLLEN _ >_.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. SUMMARY AND CONCLUSIONS

Paleoecological reconstructions of life during the late Pleistocene

are fragmentary and generalized. The nature, extent and movements of

the Laurentide ice sheet are also controversial. New information from

Oakland Bog has strenghtened and enlarged upon previous post glacial

paleoecological interpretations. The palynomorphs and molluscan

shells preserved in the marl sediment at Oakland Bog were used to re­

construct the environmental and ecological evolution of the lake and

climatic history of southwestern Michigan (Fig. 18).

The vicinity of Oakland Bog became free of ice nearly 16,000

years ago. The pioneering molluscan community recorded in the initial

lake stage of Oakland Bog was characterized by arctic species such as

Pisidium lill.j eburgi and Valvata sineera helicoidea. The occurrence

of the riverine form of Pisidium compressum further suggests Oakland

Bog resembled a pond or intermittent creek shortly after the depression

became ice-free.

Pollen recovered from the basal marl were dominantly herbs (par­

ticularly Cyperceae and Artemisia) and spruce pollen, interpreted to

represent tundra and parkland elements. The late-glacial pollen

spectra at Oakland Bog suggests a mosaic of vegetational communities

existed and not a continuous tundra.

The climate was relatively cool and continental with no evidence

of climatic reversals associated with ice-margin fluctuations. The

general climatic trend suggested from pollen and mollusk study at

118

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 119

Oakland Bog is one of a gradual climatic warming. What have pre­

viously been interpreted to represent climatic shifts, appear now to

be a function of differential immigration rates.

The pioneering Mollusk Stage at Oakland Bog was followed by a

sequence of stages adapting to changing environmental conditions of

the lake as it evolved and the climate drew warmer. The low diversity

of the Pioneer Mollusk Stage rapidly expanded as the pond was trans­

formed into a permanent hardwater lake, providing new niches for occu­

pation.

The Second Enrichment Molluscan Stage (interval 518cm-549cm) is

correlated with the dry, warm climate of the Pine Pollen Period (pollen

zone II). Shallow water mollusks, Marstonia decepta and Helisoma

anceps, associated with luxuriant vegetation and warmer waters re­

placed the cold-loving species (Fig. 18). The sedimentary record at

site II also records the shoaling of the water on the interpreted

prograding carbonate bank when root-like structures and peat fragments

interrupt the deep water laminations with Nitella.

The development of the closed boreal forest (pollen zone II) is

interpreted to represent warming climate associated with the Valders

Ice retreat which resulted in the deterioration of the spruce domi­

nated woodland nearly 12,000 years ago. The ice and low lake levels

no longer blocked the westward path of immigration of pine from their

glacial refugia in the Appalachians. 10,000 years ago the Great

Lakes region was completely ice-free and White pine was able to immi­

grate into southwestern Michigan following the earlier wave of hardier

Jack/Red pine.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 120

The abrupt rise in pine is accompanied by slower rises in elm,

ironwood and oak. The warmer dry climate was also responsible for

the development of prairie conditions in the west. Southwestern

Michigan was already too dry 8,000 years B.P. for the survival of

hemlock as it began to move from its eastern refugia. The prairie

vegetation and associated soil development in Ohio and Indiana acted

as a barrier to the further immigration of the more mesic southern

hardwood species.

At the time the pollen spectra records the transition from pine

to hardwood vegetational assemblage, permanent large lake conditions

became established as evidenced by the Cincinnatia cincinnatiensis

association (Plate 1). Deeper parts of the lake still acted as a

refugia for the cold-loving species of the late- and early post­

glacial.

The molluscan community flourished (interval 305cm-183cm) as

extensive, highly vegetated shallows developed on the carbonate banks.

Marstonia decepta proliferated in the warm, eutrophic waters. These

more mesic conditions were also recorded in the vegetation 7,400 years

ago, with the -immi g r a t i o n of beech into the oak dominated forests

in southwestern Michigan.

In the final stages of the lake, broad shallows had developed as

the lake filled with marl. Sedimentation rates increased with the

accelerated plant growth in the warm shallow waters. Gyraulus parvus

dominated the shallow community at the expense of other mollusks,

indicating the luxuriant growth of vegetation in the lake.

During the life of the lake, from its earliest oligotrophic,

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 121

cold water ponded stage to the shallow, warm water eutrophic lake

stage, the continued presence of Pisidium nitidium indicates hard-

water conditions existed throughout the history of the lake. The

associated molluscan fauna at Oakland Bog also indicate the lake was

relatively silt-free with a soft bottom.

It appears the lake was drained rapidly, terminating marl pro­

duction while organic deposition continued forming peat. There was

no evidence that terrestrial mollusks had inhabited the site. The

transition from lake to fen-bog may be reflecting more than just the

natural aging process of a lake. The poor preservation of paly-

nomorphs in the peat, restrict vegetational reconstruction at Oakland

Bog; however, regional studies indicated that there was a transition

to prairie conditions (Hypsithermal Zone) coincident with the peat

deposition 6,000 years B.P. at Oakland Bog.

The following conclusions can be reached based on the available

evidence from Oakland Bog and the surrounding region as to some of

the late-glacial and post-glacial environmental changes in south­

western Michigan and the study site:

1. The late-glacial pollen spectra records a mosaic of tundra

and parkland communities.

2. No continuous tundra developed along the ice-margin in the

c entral midwes t.

3. Climate was relatively cool and continental during the

late-glacial with no evidence of climatic reversal associated with

ice-margin fluctuations.

4. Pioneer Molluscan Stage represents cold, oligotrophic,

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 122

ponded to intermittent creek conditions.

5. The Pine Pollen Period represents the deterioration of

spruce woodland as the climate warms and immigration barriers are

breached.

6. Enrichment Molluscan Stage II and the Pine Pollen Period

are coincident with the same dry, warm climate.

7. Oak replaced pine as the climate continued to warm.

8. More mesic conditions occurred 7,400 years B.P. allowing

the immigration of beech into southwestern Michigan.

9. Vegetational and molluscan assemblages suggest the climate

gradually warmed during the post glacial.

10. Physical barriers of ice, water, and closed vegetational

communities were obstacles to immigration.

11. Differential immigration rates of species influenced vege­

tational communities.

12. Site I sediment facies are interpreted to represent a lake-

mount.

13. Site II sediment facies are interpreted to represent a

prograding carbonate bank.

14. Site III sediment facies are interpreted to represent the

deep water basin at Oakland Bog.

15. The molluscan communities during the lake stage of Oakland

Bog record relatively silt-free, soft bottom and hardwater conditions

as prevalent during the development of the lake.

16. Ecotones show a south to north transition which is time-

transgressive.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. RECOMMENDATIONS

Preliminary results indicate that Oakland Bog is a fruitful site

for postglacial paleoecological investigations since the site is

relatively undisturbed with good control.

A major question to be investigated is whether the valley or

linear trend of which Oakland Bog is part of is a continuous car­

bonate system. Cores or wash-boring lakes and between lakes would

permit better geological evolution of the glacial environment in

this region.

Geochemical studies would provide a better insight into the

conditions under which the marl formed. Pigment studies and

chlorophyll degradation production studies would contribute to the

understanding of the productivity of the lake and habitat recon­

struction.

Absolute pollen frequency studies in other similar lakes would

supplement the Oakland Bog study. Within the Oakland Bog site, more

molluscan and pollen investigations would increase the accuracy of

environmental reconstructions. Additional jet-down transects across

Oakland Bog would provide a better handle on the true morphology of

the depositional site.

Further investigation of the general glacial geology of the area

would probably solve the questions about earlier drainage in the

area and geologic evolution of the region. This may ultimately aid

in the better understanding of the natural resources and ecology of

southwestern Michigan.

123

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. REFERENCES CITED

Andersen, S., 1954. A late-glacial pollen diagram from southern Michigan. Danmarks Geologiske Undersogelse, ser. II, 80:140-155.

Archer, A., 1939. The ecology of the Mollusca of the Edwin S. George Reserve, Livingston County, Michigan. Occ. Papers Mus. Zool. Univ. Michigan no. 398:1-24.

Baker, F., 1916. The relation of mollusks to fish in Oneida Lake. New York Coll. Forestry. Tech. Publ. 4 366 pp.

Baker, F., 1918. Post-glacial molluscs from marls of central Illinois. Jour. Geol. 26:659-671.

Baker, F., 1920. Pleistocene Mollusca from Indiana and Ohio. Jour. Geol. 28:439-452.

Benninghoff, W., 1962. Calculation of pollen and spore density in sediment by addition of exotic pollen in known quantities. Pollen et Spres 4:332-333.

Benninghoff, W., 1963. The Prairie Peninsula as a filter barrier to postglacial plant migration. Proc. Ind. Acad. Sci. 73:116-124.

Benninghoff, W., 1968. Biological consequences of Quaternary gla­ ciations in the Illinois region. Quaternary of Illinois. Spec. publ. 14:70-77.

Benninghoff, W. and Hibbard, A., 1961. Fossil pollen associated with a late-glacial woodland musk ox in Michigan. Michigan Acad. Sci., Arts, and Letters i ipers 46:155-159.

Bernado, J. and Webb, T., 1977. Changing patterns in the Holocene pollen record of northeast North America: A mapped summary. Quat. Res. 8:64-96.

Berry, E., 1943. The Amnicolidar of Michigan: distribution, ecology, and taxonomy. Misc. Publ. Mus. Zool., Univ. Mich. 57:1-68.

Birks, H., 1973. Modern pollen rain studies in some arctic and alpine environments, in Quaternary Plant Ecology (H. Brinks and R. West, Eds.) pp 143-168. John Wiley, New York.

124

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 125

Blatchely, W. and Ashely, G., 1900. The lakes of northern Indiana and their associated marl deposits. Ann. Rept. Ind. Dept. Geol. and Nat. Resources, Indianapopi.

Bonney, A., 1972. A method for determining absolute pollen fre­ quencies in lake sediments. New Phytol 71:393-405.

Brewer, R., (Ed.), 1972. The ecology of Kalamazoo County. Western Michigan University.

Brinks, H., 1973. Modern macrofossil assemblages in lake sediments in Minnesota. In Quaternary Plant Ecology (G. Brinks and R. West, Eds.) pp. 173-189. John Wiley, New York.

Broecker, W. and Farrand, W., 1963. Radiocarbon age of the Two Creeks Forest bed, Wisconsin. Geol. Soc. Amer. Bull. 74:795-802.

Brunskill, G., 1968. Fatetteville Green Lake, New York: I. Physical and chemical limnology, II. Precipitation and sedimentation of calcite in a meromictic lake with laminated sediments. Unpubl. PhD. thesis, Cornell Univ. 171p.

Brunskill, G., 1969. Fayetteville Green Lake, New York: II. Pre­ cipitation and sedimentation of calcite in a meromictic lake with laminated sediments. Limnol. and Ocean. 14:830-847.

Brunskill, G. and Ludlam, S., 1969. Fayetteville Green Lake, New York: I. Physical and chemical limnology. Limnol. and Ocean. 14:814-829.

Bryant, V., 1975. A 16,000 year pollen record of vegetational change in central Texas. Palynology 1:143-156.

Cain, S., Segadas-Vianna, F., and Bunt, F., 1950. Mollusks of Sodon Lake, Oakland County, Michigan. II. The winter occurrence of certain species. Ecology 31: 546-553.

Camp, M., 1974. Pleistocene Mollusca of three southeastern Michigan marl deposits. Sterkiana 56:21-64.

Clarke, A., 1973. The freshwater Mollusca of the Canadian Interior Basin. Malacologia 13:1-509.

Cleland, C. 1966. The prehistoric animal ecology and ethnozoology of the upper Great Lakes region. Anthropological Papers 10:1-294.

Clowers, S., 1966. Pleistocene Mollusca of the Box March deposit, Admaston Township, Renfrew County, Ontario, Canada. Sterkiana 22:31-59.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 126

Cox, D., 1959. Some postglacial forests in central and eastern New York State as determined by the method of pollen analysis. Bull. New York State Mus. Sci. Ser. 377. 52pp.

Craig, A., 1970. Vegetational history of the Shenandoah Valley, Virginia. Geol. Soc. of Amer. Spec. Paper 123:283-296.

Craig, A., 1972. Pollen influx to laminated sediments: A pollen diagram from northeastern Minnesota. Ecology 53:46-57.

Curtis, J., 1974. The vegetation of Wisconsin: An ordination of plant communities. Univ. Wisconsin Press, Madison, Wise. 657p.

Cushing, E., 1963. Late-Wisconsin pollen stratigraphy in east- central Minnesota. PhD. Thesis. Univ. Minn., Minneapolis, Minn. 165p.

Cushing, E., 1965. Problems in the Quaternary phytogeography of the Great Lakes region. In The Quaternary of the United States (H. Wright and D. Frey, Ed.) pp. 403-416. Princeton Univ. Press, Princeton, New Jersey.

Cushing, E., 1967. Evidence for differential pollen preservation in Late Quaternary sediments in Minnesota. Rev. Palaeobot. Palynol. 4:87-101.

Cushing, E., 1967b. Late-Wisconsinan pollen stratigraphy and the glacial sequence in Minnesota. In Quaternary Paleoecology (E. Cushing and H. Wright, Eds.) pp. 59-88. Yale Univ. Press, New Haven.

Davis, A., 1977. The Prairie-Deciduous Forest ecotone in the Upper Middle West. Ann. Assoc. Amer. Geogr. 64:204-213.

Davis, C., 1900. A contribution to the natural history of marl. Geol. Survey Mich. 8:65-99.

Davis, C., 1901. A second contribution to the natural history of marl. Jour. Geol. 9:491-506.

Davis, M., 1965. A method for determination of absolute pollen frequency. In Handbook of paleontological techniques (B. Kummel and D. Raup, Eds.) pp. 674-686. W.H. Freeman, San Francisco, Calif.

Davis, M., 1967. Pollen accumulation rates at Rogers Lake, Connecticut, during late- and post-glacial time. Rev. Paleobot Palyn. 2:219-230.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 127

Davis, M., 1967b. Late-glacial climate in northern United States: A comparison of New England and the Great Lakes Region. In Quaternary Paleoecology (E. Cushing and H. Wright, Eds.) pp. 11-44. Yale Univ. Press, New Haven.

Davis, M., 1968. Pollen grains in lake sediments: Redeposition caused by seasonal water circulation. Science 162:796-799.

Davis, M., 1969. Climatic changes in southern Connecticut recorded by pollen deposition at Rodgers Lake. Ecology 50:409-422.

Davis, M., 1974. Holocene migrations of ecotones. AMQUA (Abstract) Amer. Quat. Assoc. Meeting (July 30-Aug. 1, 1974). Univ. Wise, at Madison. 18-22.

Davis, M., Brubaker, L. and Beiswerger, J., 1971. Pollen grains in lake surface sediments from southern Michigan. Quat. Res. 1:450-467.

Davis, M., Brubaker, L. and Webb, T., 1973. Calibration of absolute pollen influx. In Quaternary Plant Ecology (H. Birks and R. West, Eds.) pp. 9-25. John Wiley, New York.

Davis, M. and Deevey, E., 1963. Estimation of absolute pollen rain from pollen frequencies in sediment of known accumulation rate. Ecol. Soc. Amer. Bull. 44:81.

Davis, M. and Deevey, E., 1964. Pollen accumulation rates: Estimates from late-glacial sediments of Rodgers Lake. Science 145:1293-1295.

Deevey, E., 1939. Studies on Connecticut lake sediments. I. A post-glacial climatic chronology for southern New England. Amer. Jour. Sci. 237:691-724.

Deevey, E., 1943. Additional pollen analysis from southern New England. Amer. Jour. Sci. 241:717-752.

Dexter, R., 1950. Distribution of the mollusks in a basic bog nad its margins. Nautilus 64:19-26.

Dorr, J. and Eschmann, D., 1971. Geology of Michigan. The Univ. of Michigan Press, Ann Arbor. 476p.

Durant, S., 1880. History of Kalamazoo County, Michigan. Everts and Abbott, Philadelphia, Penn. 552p.

Engelhardt, D., 1960. A comparative study of two early Wisconsin bogs in Indiana. Ind. Acad. Sci. Proc. 69:110-118.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 128

Erdtman, G., 1943. An introduction to pollen analysis. Chronica Botanica Waltham, Mass. 231p.

Erdtman, G., 1969. Handbook of Palynology: An introduction to the studies of pollen grains and spores. Hafner Pub. Co., New York. 486p.

Evenson, E., Farrand, W., Eschemann, D., Mickelson, D. and Maher, L., 1976. Greatlakean Substage: A replacement for Valderan Substage in the Lake Michigan Basin. Quat. Res. 6:411-424.

Faegri, K. and Iverson, ., 1975. Textbook of pollen analysis. Hafner Press, New York. 295p.

Fitting, J., 1970. The archeology of Michigan. Natural History Press, Garden City, New York. 274p.

Fletcher, M. and Clapham, W., 1974. Sediment density and the limits to the repeatability of absolute pollen frequency determinations. In Geoscience and Man IX (B. Perkins, Ed.)

Flint, R., Colton, R., Goldthwait, R. and Willman, H., 1959. Glacial map of the United States east of the Rocky Mountains (scale 1/750,000). Geol. Soc. Amer.

Flint, R., 1971. Glacial and Quaternary Geology. Wiley, New York.

Fowells, H., 1965 (Ed.) Silvics of forest trees of the United States. U.S. Dept. Agr. Forest Service Agr. Handbook 271. 762p.

Fredskild, B., 1975. A late-glacial and early post-glacial pollen- concentrating diagram from Langeland, Denmark. Geologiska Foreningens i Stockholm. 97:151-161.

Frey, D., 1951. Pollen succession in the sediments of Singletary Lake, North Carolina. Ecology 32:518-533.

Frey, D., 1953. Regional aspects of the late-glacial and post­ glacial pollen succession of southeastern North Carolina. Ecological Monographs 23:289-313.

Frey, D., 1955. A time revision of the Pleistocene pollen chronology of southeastern North Carolina. Ecology 36:762-763.

Frey, D., 1959. The Two Creeks Interval in Indiana pollen diagrams. Indiana Lakes and Streams Inv. 5:131-139.

Fries, M., 1962. Pollen profiles of late Pleistocene and Recent sediments from Weber Lake, Minnesota. Ecology 43:295-308.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 129

Friesner, R. and Potzger, J., 1946. The Cabin Creek raised bog, Randolph County, Inc. Butler Univ. Bot. Studies 8:24-41.

Garrison, G., 1967. Pollen stratigraphy and age of an early post­ glacial beaver site near Columbus, Ohio. Ohio Jour. Sci. 67:96-105.

Gilliam, J., Kapp, R. and Bogue, R., 1967. A post-Wisconsin pollen sequence from Vestaburg Bog, Montcalm County, Mich. Papers Mich. Acad. Sci. Arts and Letters 52:3-17.

Griffin, C., 1950. A pollen profile from Reed Bog, Randolph County, Indiana. Butler Univ. Bot. Studies 9:131-139.

Gruger, E., 1972. Late Quaternary vegetational development in southcentral Illinois. Quat. Res. 2:217-231.

Gruger, J., 1972. Studies on the late Quaternary vegetation history of northeast Kansas. Geol. Soc. Amer. Bull. 83:

Godwin, H., 1962. Half-life of radiocarbon. Nature 165:984.

Goldich, S., et al., 1959. The chemical composition of Minnesota Lake marl - Comparison of rapid and conventional chemical methods. Econ. Geol. 54:285-300.

Goodrich, C., 1932. The Mollusca of Michigan. Univ. Mich. Handbook 5. 120p.

Guilday, J., Martin, P. and McCrady, A., 1964. New Paris No. 4: A late Pleistocene cave deposit in Bedford, County, Pennsylvania. Bull. Nat. Speleological Soc. 26:121-194.

Hale, D., 1903. Theories of origin of bog lime or marl. Geol. Surv. Mich. 8:41-47.

Harrison, W . , Malloy, R., Rusnak, G. and Terasmae, J., 1965. Possible late-Pleistocene uplife, Chespeake Bay entrance. Jour. Geol. 73:201-229.

Havinga, A., 1967. Palynology and pollen preservation. Rev. Palaeobot Palynol. 2:81-98.

Haworth, E., 1972. Diatom succession in a core from Pickerel Lake, northeast South Dakota. Geol. Soc. Amer. Bull. 83:157-172.

Herrington, H. and Taylor, D., 1958. Pliocene and Pleistocene Sphaeriidae (Peleceypoda) from the central United States. Occ. Papers Mus. Zool., Univ. Mich. 596:1-28.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 130

Houdek, P., 1935. Pollen statistics from two bogs in southwestern Michigan. Papers Mich. Acad. Sic. Arts and Letters 20:49-56.

Janssen, C., 1968. Myrtle Lake: A late and postglacial pollen diagram from northern Minnesota. Can. Jour. Bot. 46:1397-1410.

Jelgersma, S., 1962. A late-glacial pollen diagram from Madelia, southcentral Minnesota. Amer. Jour. Sci. 260:522-529.

Kapp, R., 1969. How to know pollen and spores. WMC Brown Co. Publ., Iowa.

Kapp, R. and Gooding, M., 1964. A radiocarbon-dated pollen profile from Sunbeam Prairie Bog, Drake County, Ohio. Amer. Jour. Sci. 262:259-266.

Kelley, R. and Farrand, W., 1967. The glacial lakes around Michigan. Michigan Geological Survey Bull. 4:1-23.

Kenoyer, L., 1930. Ecological notes on Kalamazoo County, Michigan, based on the original land survey. Paper Mich. Acad. Sci., Arts and Letters 11:211-217.

Kenoyer, L., 1934. Forest distribution in southwest Michigan, as interpreted from the original land survey. Paper Mich. Acad. Sci., Arts and Letters 19: 107-111.

Kerfoot, W., 1974. Net accumulation rates and history of cladoceran communities. Ecology 55:51-61.

Kindle, E., 1929. A comparative study of different types of thermal stratification in lakes and their influence on the formation of marl. Jour. Geol. 37:150-157.

King, J., 1973. Late Pleistocene palynology and biogeography of the western Missouri Ozarks. Ecological Monographs (in press). In Late-Wisconsin Vegetational Changes in Unglaciated Eastern North America. (D. Whitehead, 1973) Quat. Res. 3:624-631.

King, J. and Kapp, R., 1963. Modern pollen rain studies in eastern Ontario. Can. Jour. Bot. 41:243-252.

King, J. and Allen, W., 1977. A Holocene vegetation record from the Mississippi River Valley, southeastern Missouri. Quat. Res. 8:307-323.

Lane, A., 1903. Notes on the origin of Michigan bog limes. Geol. Surv. Mich. 8:199-223.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 131

Lawrence, D., 1958. Glaciers and vegetation in southeastern Alaska. Amer. Scientist 46:89-122.

LaRocque, R., 1966. Pleistocene Mollusca of Ohio. Ohio Div. Geol. Surv. Bull. 62 (part 1) (General) 1-112.

LaRocque, A., 1967. Pleistocene Mollusca of Ohio. Ohio Div. Geol. Surv. Bull. 62 (part 2) (Pelecypoda) 113-356.

LaRocque, A., 1968. Pleistocene Mollusca of Ohio. Ohio Div. Geol. Surv. Bull. 62 (part 3) (Aquatic Gastropods) 357-554.

LaRocque, A., 1970. Pleistocene Mollusca of Ohio. Ohio Div. Geol. Surv. Bull. 62 (part 4) (Terrestrial Gastropods) 554-800.

Leverett, F. and Taylor, F., 1915. Pleistocene of Indiana and Michigan and the history of the Great Lakes. U.S. Geol. Surv. Monograph 53:1-529.

Leopold, E., 1956. Two late-glacial deposits in southern Connecticut. Proc. Nat. Acad. Sci. 42:863-867.

Livingston, D. and Livingston, B., 1958. Late-glacial and post­ glacial vegetation from Gillis Lake in Richmond County, Cape Breton Island, Nova Scotia. Amer. Jour. Sci. 256:341-359.

Lichti-Federovich, S. and Ritchie, J., 1965. Contemporary pollen spectra in central Canada. II. The forest-grassland transition in Manitoba. Pollen et Spores 7:63-87.

Lichti-Federovich, S. and Ritchie, J., 1968. Recent pollen assemblages from the western interior of Canada. Rev. Paleobot. Palyn. 7:297-344.

Lindeman, J., 1941. The developmental history of Cedar Creek Bog, Minnesota. Amer. Midi. Nat. 25:101-112.

Love, D., 1959. The postglacial development of the flora of Manitoba: A discussion. Can. Jour. Bot. 37:547-585.

Love, D., 1970. Subarctic and subalpine: where and what? Arctic and Alpine Res. 2:63-73.

Love, A. and Love, D., 1963. North Atlantic biota and their history. Pergamon Press, New York. 430p.

Loven, N., 1977. Analysis of an Interlobate Boundary in the Wisconsinan Drift of Kalamazoo County and Adjacent areas in southwestern Michigan. Western Michigan University Master Thesis (Unpublished) 129p.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 132

Ludlam, S., 1969. Fayetteville Green Lake, New York. III. The laminated sediments. Limnol. and Ocean. 14:848-857.

Maher, L., 1972. Absolute pollen diagram of Redrock Lake, Boulder County, Colorado. Quat. Res. 2:531-553.

Manley, G., 1955. A climatological survey of the retreat of the Laurentide ice-sheet. Amer. Jour. Sci. 253-256-273.

Martin, P., 1958. Taiga-tundra and the full-glacial period in Chester County, Pennsylvania. Amer. Jour. Sci. 256:470-502.

Martin, H., 1959. Pleistocene ecology and biogeography of North America. In Zoogeography (C. Hubbs, Ed.) Amer. Assoc. Adv. Sci. Publ. 51:375-420.

Maxwell, J. and Davis, M., 1972. Pollen evidence of Pleistocene and Holocene vegetation of the Allegheny Plateau, Maryland. Quat. Res. 2:506-530.

McAndrews, J., 1966. Post-glacial history of prairie, savanna, and forest in northewestern Minnesota. Torrey Bot. Club Mem. 122:1-72.

McAndrews, J., Berti, A. and Norris, G., 1973. Key to the Quaternary pollen and spores of the Great Lakes Region. Life Sci. Misc. Publ., Royal Ont. Mus. Univ. Toronto Press, Toronto. 61p.

Melville, H., 1972. Notes on theory and practice of radiocarbon dating and description of the C-14 Laboratory at Brock Univ. Research Report Series No. 9: Studies in geoghronometry no. 1. Brock Univ. 37p.

Miller, N., 1973. Late-glacial and postglacial vegetation change in southwestern New York State. New York State Mus. and Sci. Ser. Bull. 420. The Univ. of the State of New York, New York.

Minder, 1926. Biologisch-chemische Untersuchungen im Zurichsee. Rev. Hydrol. 3:1-70. In Brunskill, G., 1969. Fayetteville Green Lake, New York II: Precipitation and sedimentation in a meromictic lake with laminated sediments. Limnol. and Ocean. 14:830-847.

Mowery, 1961. Pleistocene mollsucan fauna of the Jewell Hill deposit, Logan County, Ohio. Sterkiana 4:1-21.

Muller, G., 1970. Aragonite precipitation in a freshwater lake. Nat. Phys. Sci. 229:18.

Nave, F., 1969. Pleistocene Mollusca of southwestern Ohio. Sterkiana 34:1-48.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 133

Nichols, H., 1975. Palynological and paleoclimatic study of the late Quaternary displacement of the Boreal Forest-Tundra Ecotone in Keewatin and MacKenzie, N.W.T., Canada. Occ. Paper No. 15 Institute of Arctic and Alpine Res., Univ. Col. 87p.

Nipkow, 1920. Vorlaufige Mitteilungen uber Utersuchungen des Schlammabsatzes im Zurichee. Z. Hydrol (Budapest) 1:1-23. In Brunskill, G., 1969. Fayetteville Green Lake, New York, II: Precipitation and sedimentation in a meromictic lake with laminated sediments. Limnol. and Ocean. 14:830-847.

Nipkow, 1927. Uber das Verhalten der Skelette planktischer Kieselal- gen im geschichteten Tiefenschlamm des Zurich und Baldeggersees. Z. Hydrol. (Budapest) 4:71-120. In Brunskill, G., 1969. Fayetteville Green Lake, New York, II: Precipitation and sedimentation in a meromictic lake with laminated sediments. Limnol. and Ocean. 14:830-847.

Ogden, J., 1965. Pleistocene pollen records from eastern North America. Bot. Rev. 31:481-504.

Ogden, J., 1966. Forest history of Ohio. I. Radiocarbon dates and pollen stratigraphy of Silver Lake, Logan County, Ohio. Ohio Jour. Sci. 66:387-400.

Ogden, J., 1967. Radiocarbon and pollen evidence for a sudden change in climate in the Great Lakes region approximately 10,000 years ago. In Quaternary Paleoecology (E. Cushing and H. Wright, Eds.) pp.117-130. Yale Univ. Press, New Haven.

Ogden, J., 1969. Correlation of contemporary and late Pleistocene pollen records in the reconstruction of postglacial environ­ ments in northeastern North America. Mitt. Inter. Ver. Limnol. 17:64-77.

Passero, R. (principal ed.) 1978. Kalamazoo County: Geology and the environment, Western Michigan University 144p.

Peters, B., 1970. Pioneer evaluation of the Kalamazoo County landscape. Mich. Acad. 3:15-25.

Potzger, J., 1948. A pollen study in the tension zone of Lower Michigan. Butler Univ. Bot. Studies 8:161-177.

Potzger, J., 1953. History of forests in the Quetico-Superior country from fossil pollen studies. J. Forestry 51:560-565.

Potzger, J. and Wilson, I., 1941. Post-Pleistocene forest migration as indicated by sediments from three deep inland lakes. Amer. Midland Nat. 25:270-289.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 134

Richard, P., 1970. Atlas pollinique des arbres et quelques arbustes indigenes du Quebec. I. Intro generale, II. Gym- nospermes, III. Angiosperxnes, and IV. Angiospermes. Le Naturaliste Canadien 97:1-306.

Richard, P., 1971. Two pollen diagrams from the Quebec City area, Canada. Pollen et Spores 13:523-559.

Richard, P., 1977. Vegetation tardiglaciaire au Quebec meridonal et implications paleoclimatiques. Geogr. Phy. Quat. 31: 161-176.

Ritchie, J. and Lichti-Federovich, S., 1967. Pollen dispersal phenomena in arctic-subarctic Canada. Rev. Palaeobot. Palyn. 3:255-266.

Ritchie, J. and Hare, F., 1971. Late Quaternary vegetation and climate near the Arctic tree line of northwestern North America. Quat. Res. 1:331-342.

Russel, I., 1901. The Portland Cement industry in Michigan. USGS 22nd Ann. Report, part III, pp.635-684.

Saarnisto, M., 1975. Stratigraphic studies on the shoreline dis­ placement of Lake Superior. Can. Jour. Earth Sci. 12:300- 319.

Sears, P., 1942. Forest sequences in the north central states. Bot. Gaz. 103:751-761.

Sears, P., 1948. Forest sequence and climatic change in the north­ east North America since Wisconsin time. Ecology 29:326-333.

Semken, H., Miller, B. and Stevens, J., 1964. Late Wisconsin woodland musk oxen in association with pollen and invertebrates from Michigan. Jour. Paleo. 38:823-835.

Shah, B., 1971. Evaluation of natural aggregates in Kalamazoo County and vicinity, Michigan. Unpubl. Ph.D. dissertation, Mich. State Univ. 216p.

Shane, L., 1975. Palynology and radiocarbon chronology of Battaglia Bog, Portage County, Ohio. Ohio Jour. Sci. 75:96-102.

Skeels, M., 1962. The mastodons and mammoths of Michigan. Papers Mich. Acad. Sci., Arts and Letters XLVII:101-126.

Sterki, V., A preliminary catalogue of the land and freshwater Mollusca of Ohio. Proc. Ohio Acad. Sci. 4:365-402.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 135

Stockmarr, J., 1971. Tablets with spores used in absolute pollen. Pollen et Spores 14:615-621.

Stuiver, M., 1970. Oxygen and carbon isotope ratios of fresh­ water carbonates as climatic indicators. Jour. Geophys. Res. 75:5247-5257.

Takahashi, T., Broecker, W., Li, H. and Thurber, D., 1968. Chemical and isotopic balances for a meromictic lake. Limnol. and Ocean. 13:272-292.

Tauber, H., 1967. Investigations of the mode of transfer of pollen in forested areas. Rev. Paleobot. Palynol. 3:277-286.

Terasmae, J., 1965. Modern pollen deposition in the Nichicum Lake area, Quebec. Can. Jour. Bot. 43:393-404.

Terasmae, J., 1967. Postglacial chronology and forest history in the northern Lake Huron and Lake Superior" Regions. Quaternary Paleoecology (E. Cushing and H. Wright, Eds.) pp.45-58 Yale Univ. Press, New Haven.

Terasmae, J., 1968. A discussion of deglaciation and the boreal forest history in the northern Great Lakes region. Proc. Entomological Soc. Ontario 99:31-43.

Terasmae, J., 1973. Notes on late Wisconsin and early Holocene history of vegetation in Canada. Arctic and Alpine Res. 5:201-222.

Terasmae, J., 1976. In search of a palynological tundra. Geo- Science and Man. 40:77-82.

Terasmae, J. and Mott, R., 1964. Pollen deposition in lakes and bogs near Ottawa, Canada. Can. Jour. Bot. 42:1355-1363.

Terasmae, J. and Mott, R., 1965. Modern pollen deposition in the Nichicum Lake area, Quebec. Can. Jour. Bot. 43:393-404.

Terlecky, P., 1974. The origin of a late Pleistocene and Holocene marl deposit. Jour. Sed. Pet. 44:456-465.

Thiel, G., 1933. The marls of Minnesota - part II. Office of Iron Range Res. and Rehab., illp.

Traverse, A. and Ginsburg, R., 1966. Palynology of the surface sediments of Great Bahama Bank, as related to water movement and sedimentation. Marine Geology 4:409-416.

Twenhofel, W., 1933. The physical and chemical characteristics of the sediments of Lake Mendota, Wisconsin. Jour. Sed. Pet. 7:68-76.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 136

Van der Schalie, H., 1953. Mollusks from an interglacial deposit (Sangamon ? age) in Meade County, Kansas. Nautilus 66:80-90.

Voss, J., 1934. Postglacial migration of forests in Illinois, Wisconsin and Minnesota. Bot. Gaz. 96:3-43.

Waddington, J., 1969. A stratigraphic record of the pollen influx to a lake in the Big Woods of Minnesota. Geol. Soc. Amer. Spec. Paper 123:263-282.

Walker, 1895. Review of our present knowledge of the molluscan fauna of Michigan. Privately printed 27p. Reprinted Sterkiana 17:10-25.

Walker, 1906. An illustrated catalogue of the Mollusca of Michigan.

Walker, P. and Hartman, R., 1960. The forest sequence of the Hartstown Bog in western Pennsylvania. Ecology 41:461-474.

Watts, W., 1967. Late-glacial plant macrofossils from Minnesota. In Quaternary Paleoecology (E. Cushing and H. Wright, Eds.) pp.89-98. Yale Univ. Press, New Haven.

Watts, W., 1970. The full-glacial vegetation of northwestern Georgia. Ecology 51:17-33.

Watts, W. and Winter, T., 1965. Plant microfossils from Kirchner Marsh, Minnesota - a paleoecological study. Geol. Soc. Amer. Bull. 77:1334-1359.

Watts, W. and Bright, R., 1968. Pollen, seed and mollusk analysis of a sediment core from Pickerel Lake, northeast South Dakota. Geol. Soc. Amer. Bull. 79:855-876.

Wayne, W., 1966. Ice and Land. In Natural features of Indiana (A. Lindsey, Ed.) pp.21-39. Indiana Acad. Sci.

Wayne, W., and Zumberge, J., 1964. Pleistocene geology of Indiana and Michigan. In The Quaternary of North America (H. Wright and D. Frey, Eds.) pp.63-84. Princeton Univ. Press, New Jersey.

Webb, T., Ill, 1974. A vegetational history from northern Wisconsin: evidence from modern and fossil pollen. Amer. Midland Nat. 92:12-34.

Webb, T., Ill and Bryson, R., 1972. Late- and postglacial climatic change in the northern Midwest, U.S.A.: quantitative estimates derived from fossil pollen spectra by multivariate statistical analysis. Quat. Res. 2:70-115.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 137

Webb, T., Ill and McAndrews, J., 1975. Corresponding patterns of contemporary pollen and vegetation in central North America. Quat. Res. 5:395-434.

West, R., 1961. Late-glacial and postglacial vegetation history in Wisconsin, particularly associated with the Valders readvance. Amer. Jour, of Sci. 259:766-783.

Wetzel, R., 1969. Factors influencing photosynthesis and excretion of dissolved organic matter by aquatic macrophytes in hard- water lakes. Vrh. Int. Verein. Theor. Angew. Limnol. 17: 72-85.

Wetzel, R., 1970. Recent and postglacial production rates in a marl lake. Limnol. and Ocean. 15:491-503.

Wetzel, R., 1972. The role of carbon in hard-water marl lakes. Nutrients and eutrophication special symposia. Limnol. and Ocean. 1:84-97.

Wetzel, R., 1973. Productivity investigations of interconnected marl lakes I: The eight lakes of the Oliver and Walters chains, northeast, Indiana. Hydrobiol. Studies 3:91-143.

White, W. and Wetzel, R., 1975. Nitrogen, phosphorus, particulate and colloidal carbon content of sedimenting seston of a hard- water lake. Verh. Int. Ver. Limnol. 19:330-339.

Whitehead, D., 1963. "Northern" elements in the Pleistocene flora of the Southeast. Ecology 44:403-406.

Whitehead, D., 1964. Fossil pine pollen and full-glacial vegetation in southeastern North Carolina. Ecology 45:767-776.

Whitehead, D., 1967. Studies of full-glacial vegetation and climate in southeastern United States. In Quaternary Paleoecology (E. Cushing and H. Wright, Eds.) pp.237-248. Yale Univ. Press, New Haven.

Whitehead, D., 1972. Developmental and environmental history of the Dismal Swamps. Ecol. Monogr. 42:301-315.

Whitehead, D., 1973. Late-Wisconsin vegetational changes in unglaciated eastern North America. Quat. Res. 3:621-631.

Williams, A., 1974. Late-glacial and postglacial vegetational history of the Pretty Lake region, northeast Indiana. Prof. Pap. United States Geol. Surv. 686-B:l-23.

Wilson, I. and Potzger, J., 1943. Pollen study of the sediments from Douglas Lake, Cheboygan County and Middle Fish Lake, Montmorency County, Michigan. Ind. Acad. Sci. 52:87-92.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 138

Wilson, 1967. Pleistocene vertebrates of Michigan. Papers Mich. Acad. Sci., Arts and Letters. 52:197-234.

Wiltson, W., 1964. The forests of Labrador. Department of Forestry Publ. No. 1066. Queen's Printer and Controller of Stationary, Ottawa.

Wodehouse, R., 1965. Pollen grains: their structure, identifi­ cation, and significance in science and medicine. Hafner Publ. Co., New York.

Wootton, C., 1975. Pleistocene Mollusca of the Colon deposit, St. Joseph County, Michigan. Sterkiana 57:1-17.

Wright, H., 1964. Aspects of the early postglacial forest succession in the Great Lakes region. Ecology 45:439-448.

Wright, H., 1968. The roles of pine and spruce in the forest history of Minnesota and adjacent areas. Ecology 49:937-955.

Wright, H . , (Ed.) 1969. Quaternary geology and climate. Nat. Acad. Sci., Washington, D.C. 162p.

Wright, H., 1971. Retreat of the Laurentide ice sheet from 14,000 to 9,000 years ago. Quat. Res. 1:316-330.

Wright, H., 1976. The dynamic nature of Holocene vegetation. A problem in paleoclimatology, biogeography and stratigraphic nomenclature. Quat. Res. 6:581-596.

Wright, H. and Frey, D., 1965. The Quaternary of the United States. Princeton Univ. Press, New Jersey. 922p.

Wright, H., Livingstone, A. and Cushing, E., 1965. Coring devices for lake sediments. In Paleontological Techniques Handbook (B. Kummel and D. Raup, Eds.) pp.495-520. W.H. Freeman, San Francisco.

Wright, H., Winter, T. and Patten, H., 1963. Two pollen diagrams from southeastern Minnesota: Problems in the regional late- glacial and postglacial vegetational history. Geol. Soc. Amer. Bull. 74:1371-1396.

Zimmerman, J., 1960. Pleistocene molluscan faunas of the Newell Lake deposit, Logan County, Ohio. Ohio Jour. Sci. 60:10-39.

Zumberg, J. and Potzger, J., 1956. Late Wisconsin chronology of the Lake Michigan basin correlated with pollen studies. Geol. Soc. Amer. Bull. 67:271-288.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Plate 1. Molluscan absolute diagram of Oakland Bog (site II).

139

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141

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143

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145

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147

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KEY: Sediment log description

p plant debris

R root-like structures

Sh abundant shells visible

Sd abundant Nitella bodies

irr irregular boundary

m mixed sediment

W wavey and scoured boundary

> and (combination of hues)

/ with (separate hues)

faint poorly distinguishable

peat peat material evident

yg (5Y 7/2) yellowish gray ay (5Y 6 A ) dusky yellow

log (5Y 5/2) light olive gray pgy (10Y 8/2) pale greenish yellow og (5Y 3/2) olive gray bb (5YR 2/1) brownish black

gy (5Y 8 A) grayish yellow ab (5YR 2/2) dusky brown

mob (5Y k/k) moderate olive brown mlg (N6) medium light gray lob (5Y 5/6) light olive brown mg (W5) medium gray

po (10Y 6/2) pale olive mbg (5B 5/1) medium bluish gray go (10Y k/2) grayish olive mag ( m medium dark gray lo (10Y 5 A ) light olive lg (H7) light gray

ob (5Y 2/1) olive black vlg (N8) very light gray

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 149 og 3h og log og Sh log ys log.yg log og/log peat:dh =S log acg Big 1=3 iog og og ys OS is log ad log log c ag Bg.Og os =S Bg.og ag log mg OS yg.icg.og los Big ag .7 COBS 3 COBS 180 -118 - I',’ 2

-73 *. 31.5 -78.5 31.5 37.5 -8537.5 35 -31.5 ys 77 77 -75 ’ 8 73 -71.5 -7071.5 =5 113 113 -90.5 90.5 37.5 90.5 -3.5 -77 -3.5 135 135 -133.5 119 119 -113 133 133 -136 136 136 -135 133.5-120 120 11*5.5-139.5 139.5-138 150.5-11*5.5 153.5-152.5 152.5— 51.5 159 159 -157.5 156.5-153.5 loO loO -159 157.5-157 172 172 -171 171 171 -160 178.7-172.7 COBS *.. SITS 191.2-139.2 181 139-2-181*. 2 log 139-2-181*. ISO.7-178.7 221.7-191*. 2 227.7-221.7 221.7-191*. 19**.2-191.2 ys 239-7-229.2 229.2-227.T SITS SITS 21*1.7-21*0.7 21*0.7-239.7 Og log log ys og log los iog Sh ag A log =S logLrr =S 3g =S =S loslog 157 -156.5 og irr og iog irr Sh irr iog log yS irr yS 151.5-150.5 icgirr os og ys yg irr Sh irr yg yg irr yg log og yg irr yg ys ys =8 ag core, 281.5 11 5-298 5-286 . . -295.5 -285.5 - *. 5-291* *. I 298 298 -297 sirs sirs 296 295.5-295 296.5-296 og 5 295 -291*. 291 297 297 -296.5 293 293 -292.5 292.5-292 292 -291 291 -290.5 291* -293.5 291* 293.5-293 290.5-290 290 -239 log 289 -238.5 238.5-238 log 287.5-287 286 282.5-232irr log 288 288 -287.5 237 -286.5 286 285.5-285 285 -281* 231* -283 231* 2c3 2c3 -232.5 282 281.5-231 231 231 -280.5 230.5-280 280 280 -279.5 279.5-279 279 279 -278.5 278.5-278 278 278 -277.5 21*2.7-21*1.7 SITS 1t COKE1t SITS 3 -250.7261* iog 2-21*2.7250.7-21*5.2 leg 251*. S3JE42BT LOG S3JE42BT SS a log/po og log =8 og =S log og log log olg 1=S log log =S log 1=6 log og/log =S og log log log log log leg 1=S ys ys ys log ys =s ys ys ys ys =S I* 31 .3 0 3 - 5 -305.5 - 307.5 - 307 307 -306.5 303 -302.5 308 302.5-302 302 -301 301 -300.5 300.5-299.5 298.7-293.5 306 321 321 319.5 311*. 310.5-310 307.5-307 5 305.5-301*. 301* 299.5-299 299 -298.7 303.5-308 olg 325. 5-325 325. 321*.5-323.5 los 323.5-321 313 -312.5 312.5-312 312 -311.5 311.5-311 325 -321*. 1* 325 -321*. 315.5-315 -313.5 311* 313.5-313 311 -310.5 309 -308.5 328.5-328 326 326 -325.5 319.5-315.5 327 327 -326.5 326.5-326 5 315 -311*. 328 328 -327.5 SITS I, CORE A CORE I, SITS 4

z 1=5 log o, I°S= log Sh 1=5 log loglog 306.5-306log log os 5-30U 301*. log log ys log los leg log log Sh log irr log 310 -309 ys log ys ys ys 05 log log 05 ys ay ys ys ys ys ys 35 ys nog2 327.6-327

337.5-336.5 336.5-335.5 331 -330.5 329 323.5 31*1.5-339-5 339-5-339 339 338.5 335.5-331* 31*3.5-31*2.5 -332.5 331* 332.5-331 330.5-329.5 329.5-329 355 355 -351* -353-5 351* 353 353 -352.5 352 -350 350 -31*8.5 31.2.5-31*1.5 338.5-337.5 31*6 -31*3.5 31*6 356 356 -355 353-5-353 352.5-352 371*. 5-372-5 371*. 358.5-353 358 -357-5 357-5-357 357 -356 31*8.5-31*6 331 331 aarl/till 365 365 -362.5 372.5-369.5 369.5-363 363 -366.5366.5-365 log 362.5-360.5 360.5-359.5 359-5-358.5 SITE SITE A COES 331 331 -373.5 5 378.5-376.5 376.5-371*. Hou.vcxn Reproduced with permission ofthe copyright owner. Further reproduction prohibited without permission. 150

S g 25 3 _ s "a — e £ es tr4 *j O — II 0 a. a 1 ~ £ 5! 2 = £ 5 = £ £ = X 2 as g O p. T4 5 3 | s § 8 § 2 | 3 3 £ 3 a — — a a a a SITS I, CORE c SITS n , CORE D SITE II, CORE 3-1 SITE II, CORE A-l TO -63.5 log 600 -538 aob/loo 180 -170 log 267 -231 abg 63.5 -67.5 7 8 600 -598 so 170 -160 SO 231 -279 po 67-5 -6l 7 8 . 1 ° a 598 -595-5 so 160 -155 log 279 -275 abg/po R 61 58.5 = 8 595-■5-591.5 go 155 -119 Og 275 -271 abg/po 53.5 -1*6 yg.aig.log - 591*.-5-533 So 119 -113 log 271 -271 abg,dag 16 -1*5 =8 538 -531.5 faint la=. 118 -116 SO 271 -269-5 abg,=g i*; -13-5 78 588 -587.5 lob 116 -111 eg peatirr 13.5 -37 is 587-5-537 =ob 111 -112 yg,aob,go,05269 .5-267.5 afcg/po,dy SITE I. CORE D 537 -5S6.5 aob faint,Sd 267-■5-267 abg 37 -35 rig 536. 5-536 sob 112 -110 go 267 -266 a'og/po,dy 35 -30 peat:db 536 -535.5 07 H O -395 log.yg faiataSo -265 abg 30 -1 peat: bb 535-5-535 07 Sh, Sd 265 -261 abg/po,ay 535 -531.5 lob 395 -367 og,log 261 -262 nbg irr SITE II, CORE; J 531*.-5-531 lob 337 -333 og 262 -261 =bg 563 -539 log/nob/go 531 -533-5 go 333 -377 7S.-og.og 261 -251 log a Sd 0 0 . 3 - 0*0 sand diice 533--5-582.5 lob,lb 377 -373 og faint lea SITE II, CORE 0 532.5-575.5 go,lob v 373 -369 og,ys 251 -250 go, abg/ 525 523 nob 576 -575 SO 372 vooa po 523 -622 ncc,dy 575 -563.5 =cb SITS II, CORE 3-2 250 -219.5 abg 622 -620.5 lob 5o3. 5-563 lob 369 -353 og/yg 219.■5 -217 abg,go/pc 620.5 -619 nob 563 -565 aob,go irr. 353 -352 log peat irr 619 -613 nob 565 -561 ay,lob 352 -350.5 og 217 -213 log/po 5lS t617 lob 561* -563 dy,iob 350.5-319 log peat irr 617 -5l5 dy 563 -555 go,aob 319 -317 Og 213 -236 abg/po Sd 6 16 -515-5 lob 555 -551 SO 317 -316 log 236 -228 abg irr 615.5-611.5 nob 551* -553-5 &o,g 316 -311 Og nreasy 5li*. j-6il nob 553- 5-552 aob 311 -337 og,log.yg 228 -221 og,go/dy oil* -6l** nob 552 -551 aob,lob 337 -313,309 yg.iog 221 -222 iob.dy 613 -610.5 lob Sh 551 -51*9 aob,cy irr. irr ? 222 -220 so 610 .5-610 noo/lob 519 -518 aob irr. 309 -296 adg,ag 220 -216 log/yg,po . a 610 .5-ol0 0 51*3 -517.5 go,=ob 296 -291 sag R peatirr 510 -609.5 5^7 - 5—51*6.5 SO 29a -293 log 216 -212 po,log/yg 609.5 -609 0 51*6.3-515 50 , g 293 -292 sag 212 -211.5 log 609 -608.5 s 51*5 -511.5 pog 292 -288 og 211. 5 -201 3 log.yg 503.5-603 a = 1 1 . 5-511 iog 283 -291 og/log 20h -136 75, ?o <>2 503 -607.5 511 -513.5 aob 281 -281 log.yg 1S6 -180 log/dy.yg 607-5-607 C 51*3.5-5H log ,=ob, pc 231 -279 og 130 -176 log/yg 607 -606.5 j s 51*1 -5l0 3 7 SITE II, CORE A—1 oreaay 506.5 -606 5I1C -535 aob,lob,pgy 302 297 io,,po Sh 176 -171.5 log/yg Sh 606 -603 lob,nob 535 -533.5 go.po peat bary 603 -602.5 nob 533- 5-533 lob 297 -291 abg Sh SITE 11, CORE A-2 602 .5-602 lob 533 -532 acb peat 171. 5 -151.5 faint iao 502 -501.5 nob 532 -531 lob irr 291 -292.5 abg/po peat ? R Sh 601.5 -6 01 lob 531-5-531-526 gO/=ob/pgy =292.5-290.5 po Sh Sd R 151.5 - H 9 .5 is 601 -600.5 nob 290.5-289.5 abg 119. 5 -111.5 log 6 OO.5-0OO lob 239.5-289 abg/po ill. 5 -1U.5 ag 239 -287 PO ill. 5 -131.5 yg/iog

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 151

SITE II, CORE A-2 SITE III, COIRE 3-2 SITE III, CORE A-l SITE III, CORE A-2 131.5-110.5 =S 379 -376 go 262 -261.5 log,go 131.5-130 og creaay 110.5-103.5 los 376 -370 og.po 261.5-260.5 iog 130 -123 yg.log irr 103.5-31.5 og irr 370 -368.5 og 260.5-259 los,SO 123 -118 peaty aarl ? sand 363.5-367.5 SO 259 -253-5 log ob P R Si 57-5 -102.5 i'8 »l°8 367.5-367 aob 258.5-256.5 log, go 118 - H i og.log r P. 102.5-51.5 bb,og,nob 367 -361.5 OS 256.5-253.5 log.po . peaty ? a 361.5-36 I aob 253.5-252 SO 111 -109-5 peaty aarl 51-5 -19.5 og, peat 361 -359.5 os,go 282 -219 Og.go ob ? P. So Sh 359-5-358.5 SO 219 -all.; ^og/iy.go 109.5-105.5 peaty sari 19.5 - peat 358.5-335-5 log,go 213.5 «ca peat ob ? ?. Sh §10 lea 211.5-213 SO 05,iog irr SHE II, CORE n laa. ■ereaay 106.5-37.3 log ? 91 -91.5 3lg 335.5-331 log 213 -215-5 faint !aairrS7.5 -30 og ? a Sh 91.5 -37 log 331 -333-5 aob 210 peat lea 30 peat 37 -76 3lg 333-5-332 so 236 peat lea 76 -57 Bg 332 -315 log,go 232 peat lea 57 - U log irr Sh 232 -227 Sd irr S h aid.laa. 220 peat lea 11 -23 og 315 -313 los 210 -236.5 log,lob grainy 313 -311-5 aob,lob 230.5-231.5 go,lob 2S -11 Og,2g 311.5-307 lcg.og laa 231-5-229 go Sh 3h ? 307 -302 log P 229 -225 log.oy u -10.5 bb Sh 302 -301 aob 225 -215 og.lob irr peat 301 -295 aob,log 215 -209-5 go ? Sh 13.5 - peat 295 og Sh 209-5-206 go,log 295.5- disturbed 206 -199 og ? R SITE III, CORE D 199 -197 go,aob ? R 523 -522 till SITS III, CORE A-i 197 -195 og.aob ? R 522 -513 a s 231 -278.5 lob,cob Sh 195 -192.5 og ? ?. 513 -516 dg vood 278.5-273 og 192.5-138 lob,eg,go 516 -510 yg/lg,dy 273 -277.- peat faint laa ? vood 277.1-271-3 og.log ? R Sh 510 -191 PS.5° 3h 271.3-271.2 log.lob 188 -182 lob,Iog,go 271.2-273-2 log ? P. Sh SITE III, CORE 3-1 273.2-272.5 log,lob 162 -18! log.yg 177 -373.5 faint lea. 272.5-271.2 log coarse 177 -173 log 271-2-266 log,lob 173 -153 log.dy faint laa. SITE III, CORE A-2 153 -139 po,aob 266 -265 lob 157-5-112 faint laa. 139 -108 log,aob 265 -263.5 log creasy og 1C3 -378 lob,po,leg 263.5-262.5 lob ? P. Sd a og Sh 262 .5-262 log 112 -131.5 log.yg irr Sh

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. JET-DOKS SEDIMENT RECORD

DEPTH CM DESCRIPTION (from field observations)

i33-2bt yellowy’, shell hash and whole shells

2 h k -2 7 k gray, N ite ll a. few obviously large shell3, clumpy m arl

271-305 gray, shinny Unoid shell fragments

305-366 gray, Nitella. grass-like threads and plant debris, shara fragments

^57-^23 gray, many large Unoid shell fragments, .litalia, grass-like threads

513-5“9 gray, Uniod shell fragments, NTtella. bark, clumpy marls, few shells

520-671 gray, shara ooids abundant, scarce shell, clumpy, few sand grains of very fine subrounded cuartz

o71 fine sand washed up from the till bi-i v; 152 yellowy

305 gray

157 sreen gray, clumpy

610 light brown, large aggregrates, scarce shells, few shara rods and pine needles

732 7charcoal, fine grassy or thread-like plant debris in dark gray marl, chara rods very abundant, few grains of subrounded very fine sand

152 yellowy, molluscan-rich (great diversity), paint debris

305 gray, few shells

157 gray, clumpy, few shells, Nitella

610 gray, few shells, Nitella abundant

7*9 ?charcoal, dominated by chara rods, pieces of pine needles light gray marl contains mica, very fine quartz sand and scattered shells

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Appendix 2. Oakland Bog (site II) Molluscan Environmental Table.

Each species occurring in a sample interval is evaluated in terms of present day ecology. Plot of RSO triangle indicates environment of assemblage. R = running water; S = stagnant; 0 = open water.

153

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 154

MOLLUSCA ENVIRONMENT c o •H ■£ t e bD j &| f s C > ol -P s O 3 8 -P 0 rH & •H D $ $ s 03 Pu CO 03 PS fee. 670 cm 2 Pisidium eompressum x x x 28.57 1 Amnicola limosa x X X 14.29 Valvata sine, hel. 14.29 R V. tricarinata X X 42.86 670-640 cm 2 Pi sidium lill.jeburgi x X 1.18 35 P- nitiaum x X X X X 20.59 2 Sphaerium striatium x X X 1.18 24 Marstonia deeepta x X X 14.12 17 Valvata sine. hel. 10.00 50 V. tricarinata x X X 29-41 3 Fossaria decampi 1.76 1 Gyraulus aeflectus 0.59 22 G_. parvus x X X X X X X 12.94 3 G. circumstriatus X X 1.76 2 Helisoma anceps x X 1.18 « 1 H. campanulatum x X 0.59 3 H.- trivolvi s XXX X 1.76 2 Physa skinneri 1.18 3 Promenetus exacuous X X 1.76 522-218 cm 1 Anoaonta sp. x X 0.14 5 Pisidium eompressum x X X 0.7 24 P. nitiaum X X X X X 3.37 1 Musculium lacustrinex XXX 0.14 2 Sphaerium striatinum x X X 0.28 73 Amnicola limosa x X X 10.25 11 A. walkeri x 1.54 106 Marstonia deeepta x X X 14.89 95 Valvata tricarinata X X X 13.34 5 Foss aria decampi 0.7 22 Gyraulus aeflectus 3.09 314 G. parvus X X X X X X 44.10 14 Helisoma anceps X 1.97 19 Physa sayii 2.67 3 P. skinneri 0.42 17 Promenetus execuous X X 2.39

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 155

MOLLUSCA ENVIRONMENT

G o •H a 0 * -P CD 60 -P > G G > O -p CD S CD 05 O 0 CO CD aj G W G (D -P EH 0 aJ •H -p O G s •rH 3 rH (— 1 -P c -P 0 G M G O >3 a} 'G CO G 0 CD rH (D X! G CD -p G EH r> 60 rH -P CO 60 ■G 3 CD C(H M g G 0}O G a} G 0 •rH > O 3 3 a) F~ g ai •P O rH 3 •H D 3 i-O UJ Ph s CO Oh CO CO 05 ^5.

It88-lt57 cm 1 Anoaonta X X 0,62 I Pisidium eompressum XXXX 0.62 12 P. nitiaum XXXXX X 7 .15 3 Musculium lacustra XXXX X 1.86 5 Amnicola walkeri XX 3.10 s 28 Marstonia deeepta XX 17.39 II Valvata sine, hel. 6.83 / *\ k2 V. tricarinata XXX 26.09 y/ . */X 5 Fossaria decampi 3.10 R £- ^0 5 Gryaulus aeflectus X X 3.10 2k G. parvus XX X XXXXXX lit.91 k Helisoma anceps X X X 2.18 1 H. trivQlvis XXXX 0.62 9 Physa sayii X X 5.59 10 Promenetus exacuous X X 6.21 366-301 cm 1 Anoaonta sp. X X 0.39 1 Pisidium eompressum X XXX X 1.57 17 P. nitiaum XXXXXX 6.67 3 Sphaerium striatinum XX X 1.18 6 Amnicola limosa XXX 2.35 6 A. walkeri XX 2.35 1 Cincinnatia cinn. XXX XXX 0.39 s 65 Marstonia deeepta X XX 25.1+9 3 Valvata sine, hel. 1.18 36 V. tricarinata X XX lit.12 f/ • *\ h Fossaria decampi 1.57 R ^ ^ 0 76 Gyraulus parvus XXXX XXXXX 29.80 6 Helisoma anceps XX X 2.35 1 H. trivolvis XX XX 0.39 1 Physa gyrina X X 0.39 6 P. sayii XX 2.35 2 Promenetus exacuous X X 0.78 17 Pisidium casertarum 6.67

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 156

MOLUJSCA ENVIRONMENT

a o •H S o -p 0 60 si 0 0 > G AS ol -P 0 0) 0 d) m ?H 0 +3 E-t o I I ■H -P O u •rH a rH rH ■P G -p G !h o >j cS to G 0 0 rH D C UJ -P G. 60 rH + 3 W 6 0 O a % « 03 -P O rH > -H J c§ £ s CO PL, CQ CO « Vi.

30U'-27it cm 1 Anoaonta sp. x X 0.33 6 Pisidium casertarum x XXXXXXX 0.99 10 P. nitiaum x XXXXX 3.30 1 P. sp. 0.33 5 Amnicola walkeri x X 1.65 7 Cincinnatia cinn. x XX XX 2.31 126 Marstonia deeepta x X X 1*1.58 13 Pomatiopsis lap. x x XXX It.29 2 8 Valvata tricarinataxx XX 9.2lt 3 Fossaria parva X X 0.99 It Gyraulus aeflectus x X 1.32 73 G. parvus x x XXXXXXX 2U.09 8 Helisoma anceps x x X 2.6U 3 H. trivolvis X XXX 0.99 15 Physa sayii X ^•95 27*r--250 cm - - 1 Anoaonta sp. x X 0.08 12 Pisidium casertariumx X XXXXXX 0.90 83 P. nitiaum x X XXXX 6.25 1 Musculium lacustrax x XXX 0.08 3 Sphaerium striatimunx XX 0.23 It 6 Amnicola limosa x X X 3.U7 h8 A. walkeri x X 3.62 115 Cincinnatia cinn. x X XXXX 8.67 h29 Marstonia deeepta x XX 32.33 U Pomatiopsis lap. x x XXX 0.30 237 Valvata tricarinataxx XX 17.86 lfc3 Gyraulus parvus x x XXXX XXX 10.78 8 0 Helisoma anceps x x X 6.03 26 H. trivolvis XXX X 1.96 101 Physa sayii x X 0.81* U Promenetus exacuous XX 0.30

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 157

MOLLUSCA ENVIR03MENT

a o •H a "cO o P h A! C o| p Vi 0 V -P Eh o co ro *H P O V P -H S H H p C P O C M Vi o >> <0 co 3 CD 0 H O JEf C 0 P tn EH £ 6O 1—IP C 0 60rd: * <*-iO

250-183 cm 2 Pisiaium casertarum XX XX XX X XX 0.17 13 P. ferrugineum XX X X 0.58 7 P. nitiaum X X XXX X 0.17 2 Amnicola limosa XX X 0.17 5 A.. .'walkeri X o.l+l 83 Cincinnatia cinn. X X XXXX 6.87 68 Marstonia decepta XX X 5.68 3 Pomatiopsis lap. X 0.25 1 Valvata sihc. hel. 0.08 313 V. tricarinata XX X 25.91 1 Fossaria decampi 0.08R 7 Gyraulus aeflectus X X 0.58 1+86 G_. parvus XX XX XXXX X 1+0.23 137 Helisoma anceps X X X 1 1 .31* 5 H. campanulatum XXX 0.1+1 6 H. trivolvis XX X X 0.50 2 Physa athearni 0,17 1+ P. gyrina X X 0.33 59 P. sayii X It.88 2 P. skinneri XX 0.17 2 Promenetus exacuous 0.17

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Appendix 3. Molluscan Habitat Survey.

Plot of environmental conditions indicated by the presence of a species.

158

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WATER CONDITIONS

pH

6 7 8 fast-clear l cti a > 0 W 3 rH ■■ 1 H 0 O u cti < D p c to CO 1 •p helicoidea cincinnatiensis OPERCULATE MOLLUSCAN GROUP Cincinnatia Valvatatricarinata Pomatiopsis lapidaria Amnicola limosa Amnicola walkeri Valvata sineera Marstonia deeepta

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. 160 o o o o o o 3^567 water depth (M) water temperature 0 1 2 WATER CONDITIONS pH 6 7 7 6 8 AQUATIC MOLLUSCAN MOLLUSCAN GROUP PUIMONATES Fossaria decampi Fossaria parvus Helisoma anceps Gyraulus deflectus Gyraulusatusircumstri c Gyraulus parvus Helisoma campanulatum Helisoma trivolvis Physa athearni Physa gyrina Physa sayii Physa skinneri Promenetus exacuous

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7 8 WATER CONDITIONS pH water depth (M) fast-clear ai 0 1 o OT H M I a X (V O rH in -p

MOLLUSCAN GROUP PELECYPODA Sphaerium striatinum Pisidium nitiaum Pisidium s p . Pisidium ferrugineum Pisidium lill.jeburgi Musculium lacustra Pisidium eompressum Anodonta sp. Pisidium casertanum

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