MIDDLE AND LATE WISCONSINAN (PLEISTOCENE)

INSECT ASSEMBLAGES FROM ILLINOIS

by

Kristine D. Carter

Bachelor of Science, North Dakota State University, 1981 B~chelor of Arts, Moorhead State University, 1978

A thesis submitted to the graduate faculty

of the University of North Dakota

in partial fulfillment of the requirements for the degree of

Master of Arts

Grand Forks, North Dakota

May

1985

I"

This thesis submitted by Kristine D. Carter in partial fulfillment of the requirements for the degree of Master of Arts from the University of North Dakota is hereby approved by the Faculty Advisory Committee under whom the work was done.

This thesis meets the standards for appearance and conforms to the style and format requirements of the Graduate School of the University of North Dakota, and is hereby approved.

Dean the Graduate School

55297:1 l. l.

Permission

Title Middle and Late Wisconsinan (Pleistocene)

Insect Assemblages from Illinois

Department Geolo -'---'---'--""'--------------------------Degree Master of Arts

In presenting this thesis in partial fulfillment of the requirements for a graduate degree from the University of North Dakota, I agree that the Library of this University shall make it freely available for inspection. I further agree that permission for extensive copying for scholarly purposes may be granted by the professor who supervised my thesis work, or in his absence, by the Chairman of the Department or the Dean of the Graduate School. It is under­stood that any copying or publication or other use of this thesis or part thereof for financial gain shall not be allowed without my written permission. It is also understood that due recognition shall be given to me and the Univer­sity of North Dakota in any scholarly use which may be made of any material in my thesis.

Signature ~ :1;...:.,. LJ. 01,~ Date 3-//-1£;-

iii

TABLE OF CONTENTS

LIST OF ILLUSTRATIONS .v

LIST OF TABLES .. .vii

ACKNOWLEDGMENTS. .viii

ABSTRACT •...• . . . .. . .. . . . . .. . . . . . . . . . . . . . . . . . . . . . . • • • • • • • • • • • .. X

INTRODUCTION. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . ~ . .1

Objectives. Theory ••.•• Previous Work.

REGIONAL SETTING ..••••.•

Modern Environment •.•.••...•. Wisconsinan Geologic History. Vegetational History .•••...••

• 1 . 4 .8

• .10

.10

.10

.19

LOCATION AND DESCRIPTION OF SITES ..••..••...•....•....•.• 21

Quarry Site. Athens North Biggsville Site . ......... .

21 23

METHODS OF AN ALYS IS ••.. ................................................ • 2 7

Sample Collection. Laboratory Methods. Mathematical Treatment.

RESULTS ..... .............. .

Sediment Analysis. Faunal Analysis •••

DISCUSSION ••.••..•••.•.

at at

the the

Athens North Quarry Site. Biggsville Site.

Local Environment Local Environment Regional Climate Interpretation . ........................ .

CONCLUSIONS . .. . . .. . . . . . . .. .. .. .. . . .. . . . . . .. . . .. . . .. . . . . . . .. . .. . . .. .. . APPENDIX .•••

.27

.28

.31

.34

.34

.34

.65

.65

.80 • 91

• 107

• 109

Appendix A. Sample interval depths, and numbers of beetle individuals

sample and taxa

weights, per

interval .... ............................................. . .110

REFERENCES ..• . . .. .. . . . . . . . . .. .. . . . .. .. . . . . . . . .. . .. . . .. . .. . . .. . . . . . .115

iv

LIST OF ILLUSTRATIONS

Figure

1. Location of Wisconsinan and Holocene midcontinental North American fossil beetle sites ..••...•...•••...•• 2

2. Physiographic divisions of Illinois •..•••.•.••••••.. 11

3. Diagrammatic cross-section showing the formations and members of Wisconsinan age in northern and western Illinois ............................................................ 13

4. Quaternary deposits of Illinois ...•....•...••....... 14

5. Time-stratigraphic and rock-stratigraphic units and absolute dating of the Wisconsinan deposits in central Illinois ........................................................... .. 15

6. Measured stratigraphic section, Athens North Quarry site, Illinois •.••.••••.. , •...••...•....•.... 22

7. Measured stratigraphic section, Biggsville site, Illinois .............................................................. 24

8. Sediment size, amount of organic carbon, and percent of fossils that are corraded, Athens North Quarry site, Illinois •.••••.•.••.•••.•••....•. 35

9. Species diversity and numbers of aquatic taxa, Athens North Quarry site, Illinois ..•..••.•.•••..••. 44

10. Trellis diagram of Jaccard similarity coefficients showing faunal zones, Athens North Quarry site, Illinois ...•...•••.••••..•.•..•....••......•.•....•. 46

11. Clustering of sample intervals, Athens North Quarry site, Illinois ............................................ 47

12. Species diversity and numbers of aquatic taxa, Biggsville site, Illinois ••.••••..••••....•.•••..•.• 62

13. Trellis diagram of Jaccard similarity coefficients showing fauna! zones, Biggsville site, Illinois ••••. 63

14. Clustering of sample intervals, Biggsville site, Illinois .................................................................... 64

15. Sample intervals, lithology, and faunal zones of the Athens North Quarry and Biggsville sites, Illinois .. 73

16. Present distribution of~ borealis (Coleoptera: Scolytidae) ••...•••...•....••..••...... 92

17. Present distribution of Chlaenius alternatus (Coleoptera: Carabidae) •..••...••....••.•...•....... 93

18. Present distribution of Bembidion morulum (Coleoptera: Carabidae) •.•••.•.••.••••.•.•......•... 94

19. Present distribution of Orthotomicus caelatus (Coleoptera: Scolytidae) ...••••••.....•..••..••...•. 95

20. Present distribution of Bembidion frontale (Coleoptera: Carabidae) •••••.••....••...•........••. 96

21. Present distribution of Agonum corvus (Coleoptera: Carabidae) .......•••..........••....... 97

22. Present distribution of~ latidens (Coleoptera: Scolytidae) •.....••••••...•.•.......••. 99

23. Present distribution of Micropeplus cribratus {Coleoptera: Micropeplidae) •...•.•••••••••.••..•..• 100

24. Present distribution of Sphaeroderus nitidicollis brevoorti (Coleoptera: Carabidae) •.•••••..••••.•.•. 101

25. Present distribution of Carphoborus andersoni (Coleoptera: Scolytidae) ..•.••..•....••...•..•..•.. 102

26. Numbers of species represented in the Athens North Quarry and Biggsville beetle assemblages which pr~sently occur in the provinces of Canada and the s =es of the United States ..•••••.•.•..•...•.••... 104

Plate

I. Fossil Coleoptera from the Athens North Quarry and Biggsville sites .....•..•...••....•...•........••... 42

II. Fossil Coleoptera from the Biggsville site ..••.••..• 60

LIST OF. TABLES

Table

1. Midcontinental North P ,rican fossil beetle sites .... 3

2. Taxonomic list of the coleopteran fauna from the Athens North Quarry site, Illinois •.•....•.......... 37

3. Total numbers of coleopteran individuals of each taxon per sample interval, Athens North Quarry site, Illinois ....•••.....•....•..••....•••.••••••.. 40

4. Taxonomic list of the coleopteran fauna from the Biggsville site, Illinois ••..••.•......••..••.•.••.• 49

5. Total numbers of coleopteran individuals of each taxon per sample interval, Biggsville site, Illinois ........................................................................... 54

6. Sample interval depths and sample weights, Athens North Quarry site, Illinois ..•..••.•......•..•...•. 111

7. Sample interval depths and sample weights, Biggsville site, Illinois ••.•••...••••...••..••.... 112

8. Numbers of identified beetles and minimum number of taxa per interval, Athens North Quarry site, Illinois ................................................................... 113

9. Numbers of identified beetles and minimum number of taxa per interval, Biggsville site, Illinois .•..... 114

ACKNOWLEDGMENTS

I would like to thank the many people who gave me help

and support during this undertaking, and made this thesis

possible. I am grateful to Dr. John R. Reid, the chairman

of my thesis committee, for his guidance, patience and

understanding during the preparation of this manuscript. I

thank the members of my committee, Dr. F. D. Holland, Jr.

and Dr. Allan c. Ashworth for their help and cooperation.

I am especially grateful to Dr. Ashworth for introducing me

to fossil beetles and directing my research. Dr. Donald P.

Schwert also gave me invaluable advice and assistance

during this project.

I am grateful to the following people for helping with

sample collection: Dr. R. G. Baker, Dr. G. R. Hallberg, J.

W. Hoganson, Dr. F. B. King, and Dr. J.E. King. Dr. R. G.

Baker and Dr. G. R. Hallberg allowed me access to their

description of the Biggsville site. Dr. R. J. Stevenson

did the SEM photography of my fossils, and Dr. R. D.

LeFever helped me with statistical methods. I also thank

Dr. w. D. Gosnold for the use of his microcomputer.

The use of the facilities and the assistance of the

staff of the Canadian National collection is gratefully

acknowledged. I am indebted to Dr. D. E. Bright, Dr. J.M.

Campbell, Dr. H. Goulet, Dr. J. McNamara, and Dr. A.

Smetana for their help in the identification of my beetle

fossils. I am also grateful to Dr. J. v. Matthews, Jr. of

the Canadian Geological Survey for his help in beetle

determinations, and Dr. E. U. Balsbaugh, Jr. allowed me

access to the North Dakota State Insect Collection at North

Dakota State University. Thanks also go to Dr. A. M.

Cvancara for his determination of my gastropod fossils.

I thank the National Science Foundation (grant number

DEB 80 2392 -- A. C. Ashworth and D. P. Schwert, principal

investigators; grant number BSR-840070 -- D. P. Schwert and

G. R. Hallberg, principal investigators) and the Graduate

School of the University of North Dakota (Graduate Student

Research Grant) for providing funding for this study. I am

also grateful for the use of the facilities of the North

Dakota Geological Survey and the Geology Departments of the

University of North Dakota and North Dakota State

University.

My mother, Stella Ziegler, and family, especially

Debra Sannes and Jack and Imogene Carter, have given me

considerable support throughout my education and I am deep­

ly grateful. Finally, a very special thanks for his sup­

port, patience, and confidence in me goes to my husband,

Brian.

ix

ABSTRACT

Two highly organic sediment sequences from central and

western Illinois (the Athens North Quarry and Biggsville

sites, respectively) of Middle and Late Wisconsinan age

were studied to gain knowledge about the environment and

climate of this area during the time immediately prior to

the maximum extent of the last glaciation. Paleoenviron­

mental and paleoclimatic inferences are based principally

upon knowledge of the habitat preferences and geographic

distributions of extant species of beetles that are repre­

sented as fossils. These data are supplemented by informa­

tion from sedimentological and taphonomic studies and from

previously published palynological and plant-macrofossil

studies.

The Athens North Quarry site is a 2.8 m-thick sequence

of organic silt, ranging in age from about 25,000 to 22,000

11 c yr B.P. Two faunal zones based upon beetle fossils

were recognized; a third was defined using gastropods. The

environment, inferred from the fauna, was the margin of a

eutrophic lake surrounded by a moist coniferous forest.

The sequence of events at the Athens North Quarry site

began with the expansion of a nearby lake or pond over the

forest floor, and ended with the infilling of the water

body by Woodfordian loess and organic material.

The Biggsville site is a channel-fill deposit con­

sisting of fluvial deposits, organic sediments, peat, and

loess. Samples for beetle analysis were collected from a

2.2 m-thick section of organic loams and peat ranging in

age from 27,870 ~ 290 to 21,410 ~ 290 ~Cyr B.P. Two

coleopteran faunal zones were recognized. Habitats in­

ferred from the beetle species present in these zones are

interpreted as representing the development and demise of a

bog in an abandoned stream channel, surrounded by a thin,

spruce-dominated coniferous forest. The bog and forest

were eventually choked by peat and loess deposits. This was

followed by a period of pedogenesis.

An interpretation of the climate of central and

western Illinois between about 28,000 and 21,000 '~c yr

B.P. is based upon the present ranges of the species repre­

sented by fossils in the Athens North Quarry and Biggsville

sites. The majority of these species now occur in the

southern boreal forest region of southern Canada and

northern United States. The climate during this period

apparently was relatively stable, in contrast to the inter­

pretation of previous workers in this area; compositional

changes in the coleopteran assemblages are interpreted as

reflections of local changes in the environment rather than

as indications of the shift from interstadial to full­

glacial conditions.

xi . .

;

INTRODUCTION

Objectives

A wealth of information has been gathered about the

environmental and climatic conditions during the Wiscon­

sinan of midcontinental North America from the interpreta­

tion of fossil insects (Figure 1, Table 1). Schwert and

Ashworth (1982) proposed that at the time of maximum Late

Wisconsinan glaciation, a beetle fauna with tundra and

tree-line affinities inhabited the Midwest. Further, they

said that this fauna subsequently became regionally extinct

as the climate ameliorated. Little is known, however,

about the fauna, environment, and climate in this region

during .the period immediately prior to the last glaciation.

To gain knowledge about this period of environmental

history, two highly organic sediment sequences of Middle

and Late Wisconsinan age in central and western Illinois

were examined for beetle fossils. They are referred to as

the Athens North Quarry site and the Biggsville site,

respectively. These sites were chosen because of their

accessibility, overlapping ages, and their proximity to the

boundary of maximum Wisconsinan glaciation. The specific

objectives of this study are:

1. To describe the coleopteran fauna of each site.

2. To interpret the paleoenvironmental and

paleoclimatic conditions based upon knowledge of

' I

,'---­---I

... __ -, I

I I I I

2

' ' \

0 100ml

1. Gervais 2. Biggsville 3. Athens North Quarry 4. Garfield Heights 5. Champaign County 6. Elkader 7. Conklin

a. Norwood 9. Two Creeks

1 O. Johns Lake 11. Mosbeck 12. Seibold 13. Bongards 14. Stanton

Maximum extent of Late Wlsconsinan ice sheets (Ruhe, 1983, p. 133)

---

Figure 1. Location of Wisconsinan and Holocene midcontinental North American fossil beetle sites.

3

Table 1

Midcontinental North American fossil insect sites.

Site name Age ( Mc yr B.P.) Describing authority

Gervais >46,900 Ashworth (1980) ?Early Wisconsinan

Garfield Heights ca. 24,000

Athens North Quarry ca. 25,170 -22,170

Biggsville 27,870 - 21,410

Champaign County Middle

Elkader

Conklin

Norwood

Two Creeks

Johns Lake

Mosbeck

Seibold

Bongards

Stanton

Wisconsinan

20,530

17,170

12,400

11,860

10,800

9,940

9,750

3,500

325

- 11,200

Coope (1968)

this study

this study

Wickham (1917)

Schwert (1982)

Schwert (1982), Schwert et al. --(1981)

Ashworth et al. (1981)

Morgan and Morgan ( 19 79)

Ashworth and Schwert (1981)

Ashworth et al. (1972)

Ashworth and Brophy (19721

Schwert and Ashworth (in press)

Fischer (1980)

4

the requirements of the identified beetle taxa

and of the lithology of the sediments.

3. To compare the inferred environmental and

climatic data with that derived from other

paleoenvironmental and paleoclimatic indicators

in the region.

Theory

Fossil beetle assemblages have been used as indicators

of past environments and climates since pioneering studies

in the British Isles by G. R. Coope in 1959. Coope (1967,

1970, 1975, 1977a, 1977b, 1978, 1979) summarized the

reasons why beetle fossils are valuable paleoenvironmental

indicators:

1. Fossil beetles occur commonly in Quaternary

lacustrine and fluvial deposits.

2. Fossil beetles may be separated easily from

the sediment by kerosene flotation.

3. The robust coleopteran exoskeleton is often well­

preserved with great morphologic detail and may

be identified by comparison with modern species.

4. The taxonomy and geographic distribution of beetles

are known for the palearctic and nearctic biogeo­

graphic provinces.

5. Beetles are very precisely adapted to a wide

variety of habitats.

6. Adult beetles are often highly mobile and can

5

respond rapidly to changes in the physical

environment.

Ecological and climatic interpretations based upon

fossil beetle assemblages are dependent upon three basic

assumptions. The first of these concerns species con­

stancy. Beetle species have undergone no detectable mor­

phological evolution during at least the last half of the

Quaternary period (Coope, 1970, 1975, 1977a, 1977b, 1978,

1979). Coope (1977b) considered morphological stability of

beetle species an essential prerequisite for their use as

environmental indicators. He said gradual morphological

differences from the species' modern counterparts would

almost certainly have been associated with physiological

change, which would in turn have changed their environ­

mental tolerances. The only evidence so far for evolu­

tionary change during the Pleistocene is Matthews' (1974)

analysis of progressive wing reduction in the staphylinid

species Tachinus apterus. Later work by Matthews (1976a,

1976b), on deposits of the Beaufort Formation from Meighen

Island {Canadian Arctic Archipelago) and Lava Camp, Alaska,

demonstrated that speciation did occur during the late

Tertiary.

Extinctions of beetle species during the Quaternary

are rare. Most species of beetles named from Pleistocene

faunas have been put in synonomy with extant species.

6

Spilman (1976) described a species of Ptinidae, ftinus

priminidi, from fossil packrat middens of late Pleistocene

age in California and Arizona. This species, in the opin­

ion of Ashworth (1979), could represent an undiscovered

member of the extant fauna. Miller et al. {1981) discussed

two extinct scarab beetles, Copris pristinus and

Onthophagus everestae, from late Pleistocene asphalt de­

posits in California. They also believed that one or both

of these species may yet be discovered in inadequately

collected regions of Mexico, where their closest relatives

are presently found.

A second assumption is that the habitat requirements

of species are essentially the same as they were in the

past (Coope, 1967, 1970, 1975, 1~77a). Because ecological

needs are determined by an organism's physiological make­

up, Coope (1977a) considered it important to examine the

possibility of physiological evolution without associated

morphological change. He said that gradual physiological

changes would produce ecologically mixed fossil assemblages

which would become increasingly dissimilar to present day

populations with increasing age. Although ecologically

mixed assemblages do occur, Coope (1977a) maintained that

they are relatively rare and usually represent transitional

faunas characteristic of periods of rapid environmental

change.

7

The third assumption is that the geographic ranges of

the species are controlled by factors in the physical

environment (Coope, 1967, 1975). Factors which influence

distribution are summarized from Coope (1967). He pointed

out that, in most cases, the suitability of a habitat

involves a complex combination of these conditions:

1. Thermal environment -- Species must have an

adequate amount of heat to complete their life

cycles or reach a stage of development suitable

for hibernation before seasonal temperatures

reach the threshhold below which the beetles

become inactive. The impact of the thermal

environment may be influenced by wind exposure

and the conductivity of the substrate, or may

be buffered in habitats such as heat-generating,

decomposing vegetable matter, water, or burrows.

2. Soil conditions -- The particle size and organic­

matter content of the soil are particularly

important to ground-dwelling and burrowing species.

3. Hygric fa~tors -- Water energy, organic-matter

content, pH, and temperature of the water directly

influence species which spend all or part of their

life cycles in water. Other moisture-dependent

species live beside water or in damp or marshy

places.

4. Chemical factors -- The pH and salinity of the

soil and water are determining factors in the

8

habitat choices of some beetle species.

5. Botanical factors -- The distribution of species

dependent upon specific plants for food or shelter

is in turn dependent upon the availability of those

plants. Other related requirements may include the

presence of vegetable litter, trees, and shade.

6. Special factors -- This category includes such

factors as the dependence of predatory species on

certain prey, and the presence of carcasses or

dung.

Previous Work

Morgan et al. (1983) have reviewed the recent North -.- --American paleoentomological literature and have summarized

the results of studies in a region-by-region account. The

older literature and its significance has been reviewed by

Ashworth (1979) and Morgan and Morgan (1980).

Few paleoentomological sites of Middle Wisconsinan age

have been examined in the Midwest. Fossil beetles from

peat deposits in Champaign County, Illinois (Figure 1),

were studied by Wickham (1917). He interpreted these de­

posits td be of Sangamonian age, and deduced from the

beetle assemblage a boreal environment with climatic condi­

tions more rigorous than at present. Ashworth (1979),

however, believed that, from their geographic position,

these deposits may be of Farmdalian age.

9

The only recent study of a Middle Wisconsinan beetle

fauna is Coope's (1988) description of fossils from 24,000

14c yr B.P.-old, plant-rich silts at Garfield Heights, Ohio

(Figure 1). He identified 26 beetle individuals from a

sample weighing about two kilograms. His interpretation of

this fauna suggested an environment that was open and

rather barren, with a scattered vegetation of low plants,

carpets of moss, and a few trees. The presence of well­

drained, as well as damp soil, and the proximity of

standing, or slowly flowing water were also implied. The

climatic conditions suggested by this fauna indicated that

conditions in front of the advancing glacier in early

Woodfordian time (24,000 14c yr B.P.) were more similar to

those present1y near the northern fringes of the boreal

forest of North America than to those presently existing in

the region.

REGIONAL SETTING

Modern Environment

The Athens North Quarry and Biggsville sites are in

the Till Plains Section of the Central Lowlands Province

(Figure 2). This section has a subdued glacial topography

developed largely on Illinoian drift. The Athens North

Quarry site is in the Springfield Plain District and the

Biggsville site, in the Galesburg Plain District. The

Springfield Plain is characterized by its flatness and

shallowly entrenched drainage, whereas the Galesburg Plain

includes a level to undulatory till plain with few morainic

ridges, and more sharply incised valleys (Leighton et al.,

1948).

Espenshade and Morrison (1976) described the modern

environment of Illinois. The climate of Illinois is humid

continental. Average annual precipitation is 700 to 1,000

mm per year. The average length of the frost-free period

is 160 to 200 days. Pre-settlement vegetation consisted of

oak-hickory forest mixed with tall-grass prairie. Present

land usage in Illinois is predominantly feed-grain and

liv.estock production.

Wisconsinan Geologic History

The Late Quaternary geologic histories of the Athens

CENTRAL LOWLAND PROVINCE

OZARK

PLATEAUS

PROVINCE

, ,. .. ---==-"'

ILLINOIS STAr~ tiZOJ.OIZICAL SURVCY

11

INTERIOR LOW PlATE:AUS PROVINCE:

COASTAL PLAIN PROVINCE:

Figure 2. Physiographic divisions of Illinois (modified from Leighton et al., 1948, p. 18).

12

North Quarry and Biggsville sites are similar. Neither

site was glaciated during the Wisconsinan. The strati­

graphic units analyzed in this study were deposited during

the Farmdalian and early Woodfordian Subages of the Wis-

consinan Age (Follmer ~., 1979; Hallberg and Baker,

written communication to D. P. Schwert, 1981). The strati­

graphic relationships of the Wisconsinan Stage are sum­

marized in figure 3. A map of the surficial deposits of

Illinois is presented as figure 4.

Mayewski al. (1980) summarized the time-stratigra-

phic units of the Late Quaternary in the Great Lakes

region. The Wisconsinan Stage in Illinois has been sub­

divided (Figure 5) into five substages: Altonian, Farm­

dalian, Woodfordian, Twocreekan, and Valderan (Frye and

Willman, 1960; Willman and Frye, 1970; Follmer ~.,

1979). As a result of their reinterpretation of the type

locality, Evenson et al. (1976) suggested that the Valderan

Substage be replaced with the Greatlakean Substage. This

change has yet to gain universal acceptance. Dreimanis and

Karrow (1972) found the classification of Frye and Willman

to be unsatisfactory and suggested that the Wisconsinan

Stage be subdivided into three substages: the Early,

Middle, and Late Wisconsinan. Dreimanis and Karrow placed

the boundary between the Early and Middle Wisconsinan at

the base of the Port Talbot I beds in the eastern Great

Lakes region (pre-50,000 years B.P.); the boundary between

., "' "' ... "' z < :!': ., z 0 u V,

3'

TWOCREEKAN ANO VALO(RAN SUSSTAGES

w "' " ... "' .. " "' z " i: 0 ... ~

"

. • • ~

Juiu s~1 o TT'l1TfTTfrn; ;

• ~

13

... ··: cQO;¢;; 'nir.Nle~··;;,:::·::: .:;:.;· ~ Ptat10 SI It M«mbe, Q

••.' ... ::•A ,9·., if• r/1i.M e~ ~-~-,~-:::;·-~::·-:·:.: :::=-::~::.~ l tilts ~ ~·

__,':: ~;,' ': .. ::: .t!>.f ~{i.~ l $1lt'.i . '

Figure 3. Diagrammatic cross section showing the formations and members of Wisconsinan age in northern and western Illinois (modified from Willman and Frye, 1970, p. 70).

14

QUATERNARY DEPOSITS OF ILLINOIS

0

Jerry A. Lineback 1981

O 50km 'j F3 Fi'

AGE UNIT

Hofoeerttt t=:=::j .... Cahokia Allt.ivium. Parldand S.nd, and Henry Fo,mation COfllbined; aUuvium, windblown 'IGl'ld, al'ld sand and gravfl outwash.

Wisconsinan

Wlsconiinan

= i;,_;_;_;_;i EQualitv Formation; 1ili:, ctav. and tand in glacial and 1lack"water lakes.

tJ~M Moraine ~Grau~ ~ maralne

Wfldron and Trafalgar Formations combined; gJ~jal till with some sand, grayef, and .1Ht.

-Winn.tttMgo 4nd Gl&tfcrd Formations combined: gladal lili with some sand. gtavel, and silt; .191;: 11u.ignmenu ol some units is uncen:ain.

T~lfflffe Silt, Pearl Formation, and Hagatnawn Membff of 1ha Gla.fal'd formation i::ombined; lak• 1ilt and day, outwnb t.and, grawl, and s,IL

Wolf Cr~k Formotion; glocial till with 9'~. sand~ and Ult.

CJ BMStoek.

Figure 4. Quaternary deposits (modified from Lineback, 1981).

tSGS 1981

of Illinois

15

CHRONOSTRATIGRAPHY 14c YR LITHOSTRATIGRAPHY B.P.

(I)

C Ul (I) Q) o.-o-~ :::c

7,000

Valderan Substage

Twocreekan 11,000

E Substage Richland l

<I> 12,500 Loess -rl) >, (I)

>, ... Peoria k•,·· al C: rl) <I) Loess ation ... (I) Cl Woodfordian <I) ·- al ... Substage - <I) -al (J) (I) .

::, 0 (I)

C al C: C

Morton Loess (I) 0 (/)

0 C - 0 (I) 0 ·- (I) 25,000 (I)

0. s: Farmdalian Robein Slit Substage

28,000

Alton Ian Roxana Slit Substage

Figure 5. Time-stratigraphic and rock-stratigraphic units and absolute dating of the Wisconsinan deposits in central Illinois (modified from Follmer t al., 1979, p. 4). -

16

Middle and Late Wisconsinan was placed at the base of the

Catfish Creek drift, ·at about 24,000 years B.P. Finally,

they placed the Late Wisco.nsinan-Holocene boundary at

10,000 years B.P.

During the first of the Wisconsinan subages, Altonian

glaciers of the Lake Michigan and Green Bay Lobes entered

Illinois (Willman and Frye, 1970). Follmer et al. (1979)

described the lithologic units deposited during this sub­

age. Tills of Altonian age comprising the Winnebago Forma­

tion were deposited only in the northern part of the state.

Meltwater and valley train deposits extended southward down

the ancient Mississippi Valley and provided source areas

for the uplarid loess deposits of the Roxana Silt. Most of

this formation was deposited between 45,000 and 30,000

years B.P. According to Watts (1983), the Altonian Subage

lasted from about 60,000 to sometime before 28,000 years

B.P.

During the warmer Farmdalian Subage, glaciers re­

treated to at least within the Lake Michigan Basin, but

there is no direct evidence for the deglaciation of the

continent (Frye and Willman, 1973). Watts (19831 suggested

that because there is limited correlation of the deposits

of this subage outside of Illinois and little evidence

for a Farmdalian interstadial phase in marine cores, the

use of "Farmdalian" should be restricted to the margins of

17

the Lake Michigan Lobe at present. The age range of the

Farmdalian, as defined at its type section (28,000 to

22,000 I~ Cyr B.P, Frye and Willman, 1960) has been

shortened by McKay (1979) to 28,000 to 25,000 ~c B yr .P.

based upon more recent radiocarbon dates.

The Farmdalian Substage represented a brief pause in

loess accumulation and a period of pedogenesis (Follmer et

~., 1979). Evidence discussed by McKay (1979) from sites

along the Mississippi Valley in southwestern Illinois sug­

gested that loess deposition in near-source areas slowed

substantially during this substage but never ceased entire­

ly. According to Frye and Willman (1973), the peats and

organic-rich silts of the Robein Formation were also depos­

ited during this time. The Robein Formation overlies the

Altonian Roxana Silt and Winnebago Formations. The Farm­

dale Soil, described by McKay (1979), is weakly developed

in the Robein and Roxana Formations (Figure 3). It lacks a

structural or textural B horizon and often consists of a

thin weakly-expressed A horizon over a carbonate-leached C

horizon. Such a soil indicates that the Farmdalian inter­

stadial was brief and that the peak intensity of pedo­

genesis during this substage was at a low level.

During the Woodfordian Subage, Illinois was invaded

again by the glaciers of the Lake Michigan and Green Bay

Lobes from a northerly d~rection, and by the Erie Lobe and

18

perhaps the Saginaw Lobe, from the east and northeast,

respectively (Willman and Frye, 1970). The ice advance was

preceded by the accumulation of proglacial loess deposits,

the Morton Loess within the Wisconsinan glacial limit, and

the Peoria Loess beyond that limit. At several sites in

Illinois, the lower sections of these formations contain

organic zones consisting predominantly of wood, twigs, and

needles (McKay, 1979). Radiocarbon dates for the base of

these units indicate that the glaciers reentered the

ancient Mississippi drainage basin at about 25,000 1~c yr

B.P., 6,000 years before reaching their maximum extent at

about 19,000 14 c yr B.P. in the Peoria area of central

Illinois (Follmer et al., 1979).

Tills of the Wedron Formation overlie the Morton

Loess (Figure 3). Deposition of the Peoria Loess continued

throughout the Woodfordian beyond the limits of the glacier

(Follmer et~., 1979). The Jules Soil, developed in the

Peoria Loess, implies a period of slowed loess accumulation

that coincided with a major retreat of glacial ice of the

Lake Michigan Lobe (McKay, 1979). Loess, deposited over

the till units as the glacier receded, comprises the

Richland Loess. Other Woodfordian deposits include large

areas of outwash (Henry Formation) and lacustrine deposits

formed in backwater lakes (Equality Formation). The Wood­

fordian Subage ended about 12,500 l4c yr B.P. {Follmer,

et al., 1979).

19

According to Willman and Frye {1970), events of the

Twocreekan and Valderan (or Greatlakean) Substages are

recognized in Illinois only by the fluctuation of lake

stages and by terraces along major valleys. Loess deposi­

tion probably continued but at a greatly reduced rate, and

the amount of loess of post-Woodfordian age in the Peoria

and Richland loesses is probably minor.

Vegetational History

Pollen and plant macrofossil studies from a number of

sites in Illinois {Gruger, 1972a, 1972b; F. B. King, 1979;

J. E. King, 1979, 1981; Whittecar and Davis, 1982), Wis­

consin (Dirlam, 1979), and Iowa (Hallberg et al., 1980;

Mundt and Baker, 1979; Van Zant et al., 1980) have provided

evidence for the presence of a closed-canopy Picea-Pinus

boreal forest during the late Altonian, Farmdalian, and

early Woodfordian Substages. Berti (1975) found no evi­

dence for forest-tundra or tundra. The forest was dom­

inated by Pinus during the late Altonian and early Farm­

dalian time. Subsequently, species dominance shifted to

Picea (Dirlam, 1979; Hallberg et al., 1980; Mundt and

Baker, 1979; Van Zant et~-, 1980; Whittecar and Davis,

1982). This change from pine to spruce dominance was time­

transgressive, occurring at about 36,000 14c yr B.P. in

northernmost Illinois (Whittecar and Davis, 1982) and cen­

tral Wisconsin (Dirlam, 1979), and at approximately 25,000

•4c yr B.P. in southeastern Iowa (Hallberg, et al., 1980).

20

Picea did not appear until about 21,000 '4c yr B.P. in

south-central Illinois (Gruger, 1972a). Whittecar and

Davis (1982) suggested that the compositional shift within

the coniferous forest may have been part of a southward and

westward expansion of the spruce-dominated boreal forest,

presumably in response to climatic deterioration prior to

the advance of Woodfordian ice. Macrofossils of Larix and ----Abies studied by F. B. King (1979) from central Illinois

made their first appearance in the fossil record of the

spruce-dominated forest between about 23,000 and 22,000 14 c

yr B.P. They also provide evidence for a shift from mild

interstadial conditions to colder, moister full-glacial

conditions.

LOCATION AND DESCRIPTION OF SITES

Athens North Quarry Site

The Athens North Quarry site is at the north end of

the Indian Point Quarry of Material Services Corporation,

in the SEl/4, NEl/4, sec. 18, T.18 N., R.5 w. of Menard

County, Illinois (Figure 1). The section studied is

located approximately 90-145 m north and east of that des­

cribed by Follmer et £1.l· (1979).

The sampled section is illustrated in figure 6. The

base of the sequence (6.2 m below ground surface) is dark

gray (5Y4/1) clayey silt of unknown thickness. Overlying

this unit is a distinct, 0.3 m-thick bed of black

(10YR2.5/l) silty muck. Above the muck is 2.2 m of dark

grayish brown (2.SY4/2) to black (10YR2.5/1) organic-rich

silt. Organic fragments are present throughout the silt

but abundant spruce wood, needles and cones are concen­

trated in certain layers. The organic-rich silt unit is

overlain by 3.6 m of greenish-gray (5GY5/1), limonite­

stained, inorganic silt.

The time-stratigraphic, rock-stratigraphic, and soil­

stratigraphic nomenclature, and the ages of the principal

boundaries in the Athens North Quarry site (Figure 6)

follow Follmer et al. (1979). The bedded clayey silt

and overlying silty muck unit belong to the Robein Silt

21

CD Cl (/)

"' (/) -.. (I)

.Q 0 :::, ...I

(I)

C CG ·.:

"' 0 'O (I) ..

Q. 0 -"C 0 0 3:

-C en al CD - Cl C: iii "' ,,- (I)

e1: .0 .. :::, 0 "'(I) a: IJ..

Depth Cm)

- 1 -

-2-

c.. 3 .

- 4-

- 5 -

- 6 -

'" 7 -

22

inorganic

silt

organic

silt

muck

clayey

silt

-

14 C YR B.P.

?-22,170 i200

?-25, 170 .t.450

Figure 6. Measured stratigraphic section, Athens North Quarry site, 'Illinois.

.. (I)

C. E

"' ., (I) -(I) (I)

al

23

of Farmdalian age. The muck unit has been correlated with

the Farmdale Soil (Follmer et~., 1979). The overlying

organic and inorganic silt units belong to the Peoria Loess

of Woodfordian age. A wood and muck sample, reported by

Follmer et al. (1979), from the contact zone of the muck

unit and the overlying organic-rich silt was dated at

25,170 ~200 14c yr B.P. (Illinois State Geological Survey,

ISGS-536) and a wood sample from near the top of the

organic-rich silt unit was dated at 22,170 +450 14 c yr B.P.

{ISGS-534). The underlying Roxana Silt of Altonian age and

the Glasford Formation of Illinoian age described by

Follmer et al. (1979) were not exposed in the sampled

section. The Quaternary deposits rest on Pennsylvanian

carbonates ot the Modesto Formation.

Biggsville Site

The Biggsville site is on the south overburden face of

the Biggsville Quarry, in the SWl/4, sec. 17, T.10 N., R.4

w. of Henderson county, Illinois (Figure 1). The lithology

of the section was described by G. R. Hallberg and R. G.

Baker of the Iowa Geological Survey (Hallberg and Baker,

written communication to D. P. Schwert, 1981).

The sampled section is illustrated in figure 7.

Measured depths begin at the upper surface of the modern

soil (beneath the spoil). The base of the sequence (6.2 m

below the surface) is coarse cobble gravel with a dark

Depth (m)

-0

- 1 -

- 2 .

-3·

-4 -

.. 5 -

!

-6-

- 7 -

24

spoil

modern soil

Peoria Loess

loess paleosol

organic silty loam

peat

silty peat

organic sllty loam

organic Sandy loam

gravel

14C YR B.P.

,_ 21,410.t.290

22,980 .t. 200

-24,040 .t.250

27,870.t.420

.. <II ii E "' .. .2! -<II <II Ill

Figure 7. Measured stratigraphic section, Biggsville site, Illinois (from Hallberg and Baker, written communication to D. P. Schwert, 1981).

25

greenish-gray (5GY4/1) sandy loam matrix of unknown thick-

A ness. The gravel is underlain by Mississippian limestone,

not exposed in the measured section. Overlying the gravel

is a 0.16 m-thick unit of dark olive-gray (5Y3/2) bedded,

organic, sandy loam with an admixture of pebbles near the

base, overlain in turn by 0.3 m of very dark grayish-brown

(2.5Y3/2 and 10YR3/2), bedded, organic, s~lty loam. Well­

preserved moss mats occur along the bedding planes of this

loam. The unit is overlain by 0.45 m of dark brown

(7.5YR3/2) to dark reddish-brown (5YR3/4) and very dark

grayish-brown (10YR3/2) silty peat containing some thin

beds of fine sand and some mats of decomposed moss. Above

the silty peat is a 0.76 m-thick unit of peat, dominantly

fibrous, bedded, and dark reddish-brown (5YR3/4) to very

dark brown (10YR2/2), and very dark gray (10YR3/1). It

contains woody fragments which increase in abundance toward

the top, where some beds are almost entirely of wood. A

thin (0.18 m) unit of very dark gray (5Y3/1) to black

(5Y2.5/1) organic-rich, mucky, silty loam to silty clay

overlies the peat. The uppermost sampled unit is a 0.16 m­

thick basal loess paleosol, a silty loam. The dominant

colors of the paleosol are very dark gray (5Y3/ll, dark

olive-gray (5Y3/2) and black (5Y2.5/1). It is organic­

rich, has some fibrous organic matter and contains a few

wood fragments. Some zones are clearly bedded. Above the

basal loess paleosol is a thick (3.35 m) unit of light

grayish-brown (2.5Y6/2) deoxidized and leached Peoria

26

Loess, overlain by 1.07 m of truncated and disturbed modern

soil. Spoil and crushed rock cover the modern soil unit.

Four samples from the Biggsville section were radio­

carbon dated by the Iowa Geological Survey (Figure 7).

Their ages range from 27,870 ..:420 (Beta 4129) to 21,410

_:290 (ISGS-1231) 14c yr B.P. (Hallberg and Baker, written

communication to D. P. Schwert, 1981, 1984).

METHODS OF ANALYSIS

Sample Collection

The Athens North Quarry site was sampled in June 1979

by J. E. King, F. B. King, A. C. Ashworth, D. P. Schwert

and me. An exposure face in the east wall of the quarry

was cleaned to eliminate the possibility of contamination

of the fossil assemblage by living forms. Bulk samples of

sediment weighing between 3 and 12 kg were collected at 20-

cm intervals, beginning at a depth of 3.4 m and continuing

to 6.2 m. All samples were sealed in plastic bags and

stored at room temperature until processing. Two hundred

grams from each sample were reserved for textural and

organic carbon content analyses and sealed in plastic bags.

The Biggsville site was sampled in June 1981 by G. R.

Hallberg, R. G. Baker, A. C. Ashworth, D. P. Schwert, and

J. w. Hoganson. Bulk samples of sediment weighing between

2 and 6 kg were collected at 10-cm intervals from a cleaned

section of the south overburden face in the quarry. Sam­

pling began at a depth of 4.42 m and continued to 6.62 m.

All samples were sealed in plastic bags and stored at room

temperature until they were processed.

27

28

Laboratory Methods

Sediment Analysis

Grain-size analysis was performed to determine the

proportions of sand, silt and clay in the units of the

Athens North Quarry site. The sediment was first treated

with household bleach to remove the organic material

present. The percentages of each textural category were

determined using the method described by Perkins (1977);

the sand content was determined by wet sieving, and the

clay content by hydrometer. The silt content was calcu­

lated by subtracting the weight of the sand and the weight

of the clay from the total weight of the sample.

Organic carbon content was determined by hydrogen

peroxide oxidation (Gross, 1971). Approximately two grams

of sediment were treated with a 10% solution of hydrogen

peroxide over heat until a minimal reaction was reached.

The weight loss was calculated and converted to a percent­

age of the total weight of the sample.

Insect Analysis

Insect fragments were recovered using the flotation

technique described by Ashworth (1979). Sediment was

first washed through a 300 micron sieve. Disaggregation of

peat samples was accomplished by first boiling them in a

solution of Calgon and sodium carbonate for about one hour.

The resulting residue was drained, transferred to a plastic

29

tub, and covered with kerosene. Kerosene adheres to chitin

fragments, and because it is less dense than water, it will

rise to the surface when mixed with water. The kerosene­

mud mixture was gently stirred for about five minutes,

after which the excess kerosene was poured off, and cold

water was slowly added to fill the tub. The mixture was

allowed to settle for 15 minutes. Floating remains were

decanted onto the 300 micron sieve. The tub was again

filled with cold water and allowed to settle for 15

minutes. Floating remains were decanted onto the same

sieve and washed with household detergent to remove the

kerosene from the specimens. Beetle fragments were sorted

under a binocular microscope and stored in 95% ethanol.

Representative samples of non-coleopteran insect fossils

and gastropods were also retained. Identifiable beetle and

gastropod specimens were dried and mounted on micropaleon­

tological slides using a water soluble glue of gum traga­

canth and gum acacia. All slides were labelled with

reference numbers indicating the site name, interval number

and slide number and have been deposited in the Fossil

Beetle Laboratory of the Geology Department, North Dakota

State University, Fargo, North Dakota.

The mounted fragments were identified by comparison

with identified museum specimens in the insect collections

of the Fossil Beetle Laboratory in Fargo and the Canadian

National Collection (CNC), Department of Agriculture, in

30

Ottawa, Ontario, Canada. Recent taxonomic literature con­

taining keys and illustrations of species was also helpful.

Several specimens were referred to taxonomic specialists

for identification or for confirmation of identification.

ln cases where a species or generic name could not be

assigned with absolute certainty due to either the incom­

pleteness of the fossil or to the absence of markedly

diagnostic characteristics in the modern or fossil speci­

men, the taxonomic name was preceded by cf. Generic and

specific identification was attempted only for the Coleop­

tera. The presence of small numbers of oribatid mites, and

specimens of the Hymenoptera, and of the Arachnida was

noted but they were not included in counts. Because a

beetle has fou·r potentially identifiable parts (head,

thorax, left elytron, right elytron), diffulculties arise

when counting the minimum number of individuals for use in

statistical analysis. These problems may be resolved by

equating the maximum number of any one skeletal part of a

taxon with the minimum number of individuals of that taxon.

For example, a collection of 12 heads, 23 thoraces, 19 left

elytra and 13 right elytra would represent a minimum number

of 23 beetle individuals. Unidentified beetles were not

counted or included in mathematical treatments. Certain

intervals in the Athens North Quarry section contained a

large proportion of corraded fragments. Counts were made

of the corraded and non-corraded fragments and percentages

of them were calculated for each interval.

31

Previous collecting localities for each definitely

identified species from the Athens North Quarry and

Biggsville sites were plotted on maps to facilitate the

comparison of the geographic distribution of that species

with others in the faunas studied. Locality information

was collected from the literature (Ashworth S:! al., 1972;

Bright, 1976; Campbell, 1968, 1982; Gordon, 1976; Lindroth,

1961, 1963, 1966, 1968, 1969; Wood, 1982) and from specimen

labels in the insect collections at the Canadian National

Collection (CNC) and the United States National Museum

(USNM) in Washington, D.C. State or provincial records

were used where specific collecting localities were not

available. Species from both sites with similar geographic

ranges were combined into categories called distribution

groups for the purpose of inferring a modern analog for the

distribution of the majority of the species.

Mathematical Treatment

Counts of the number of individuals of each taxon per

interval for the Athens North Quarry and Biggsville sites

were made. The percentages of aquatic Coleoptera per

interval were also calculated. Aquatic beetles, as defined

by Leech and Chandler (1971), are beetles that spend a

significant portion of their life cycles in water.

According to Schaak and Franz (1978), species diversi­

ty in an ecosystem may be' expressed by either the number of

32

species in a sample (simple species diversity) or by more

complex measures that are sensitive to the numbers of

individuals of each species in a sample (equitability).

The latter interpretive values are dependent upon adequacy

of sampling, whereas simple species diversity is a bias­

free measure of diversity. Because of the high number of

taxa represented by a single individual per interval, com­

plex diversity measures, such as the Simpson Index, were

unusable here. In such cases, the value of the index would

be undefined mathematically. Therefore, a simple species

diversity curve using the number of taxa in an interval

versus the depth of that interval was constructed for each

site. As used here, a taxon is the lowest possible level

of taxonomic identification.

Patterns of similarity among the fossil assemblages

from sampling intervals were established by cluster analy­

sis of similarity coefficients. Jaccard's coefficient of

similarity was chosen because of its simplicity and wide

use in ecological studies. Following the example of

Valentine (1966), this coefficient was calculated as the

number of species common to two samples divided by the

total number of different species present in both samples:

CJ= c/(a+b-c)

a and b = number of species in each sample, respectively

c = number of species common to both samples

The sample units were the sampling intervals of each

33

fossil site. Sample interval number 1 of the Athens North

Quarry site ~as excluded from statistical analysis because

it contained no beetle fossils. A trellis diagram was con­

structed to illustrate the degree of similarity between

pairs of sample intervals.

The coefficients were then subjected to cluster analy­

sis by the weighted pair-group method described by Davis

(1973). In this method, intervals were arranged into a

hierarchy such that intervals with the highest degree of

mutual similarity were placed together. Then, groups or

clusters were associated with other groups which they most

closely resembled. The clustering process continued until

all of the intervals were grouped together in the form of a

dendrogram.

RESULTS

Sediment Analysis

Sample interval depths and sample weights for the

Athens North Quarry and Biggsville sites are listed in

tables 6 and 7, respectively, of appendix A. The tex­

tural composition of the sediments of the Athens North

Quarry site is illustrated in figure 8.

less than 2% sand and less than 16% clay.

sediment, 83 to 93%, is silt.

All units contain

Most of the

The organic carbon content of the Athens North Quarry

samples is plotted against depth in figure 8. The organic

carbon content of the sequence reaches its highest value,

18%, within the lower part of the organic silt lithologic

unit.

Faunal Analysis

Athens North Quarry site

At least 279 coleopteran individuals representing 17

families have been identified from the Athens North Quarry

site. A total of 61 taxa was determined below the level of

family, including 14 identified to species. Of the number

of beetles, 25% was classified as aquatic following the

usage of Leech and Chandler (1971). A summary of the

sample data is found in table 8 of appendix A.

,... E .... .c .. C. CD C

4·

5·

6·

:,, Ol

.2 Ill 0 > .. .c CD :!: .. ..J .5

Inorganic 1 sltt

2

3

4

5

6 organic

silt 7

8

9

10

1 1

12

muck 13

clayey slit 14

f.7.!71 t..i..:I

'll, sand

10 20

D % ailt El 'll,

clay

70 80 90

-- % organic carbon

10 20

\"; i

--- % corraded loaalla

10 20 30 40 50

Figure 8. Sediment size, amount of organic carbon, and percent of fossils that are corraded, Athens North Quarry site, Illinois.

w en

36

A taxonomic list of the beetles is included in table 2. A

quantitative summary of the insect taxa and their numbers

per sample interval is given in table 3.

Scanning electron photomicrographs Of some represent­

ative fossil species from the Athens North Quarry site are

shown in plate I. Fossils from both sites are included to

demonstrate the quality of preservation of the fossil mate­

rial, rather than for any systematic purpose.

Six geographic distribution groups were established to

describe the ranges of the species from both sites. The

groups to which the species from the Athens North Quarry

site belong are listed in table 2. Those species tor which

collecting locality information was not available are indi­

cated by U. Descriptions and illustrations of these dis­

tribution groups will be included in the discussion of the

paleoclimate of the region.

Maximum species diversity and density was found within

sample interval number 4 from the upper part of the organ­

ic-rich silt unit. Simple species diversity, as number of

taxa per sample interval, is illustrated in figure 9.

' I

YI

Table 2

Taxonomic list of the co!eopteran fauna from Athens North Quarry site, Illinois. Order of genera tollows Arnett (1968). Order of species in the Carabidae and the Scolytidae follows that of Lindroth (1961, 1963,

1966, 1968, 1969) and Wood (1982), respectively.

Taxa

Carabidae Sphaeroderus nitidicollis ---i3revoorti Lee. Dyschirius integer Lee. Dyschir~ sp:--~­Patrobus sp. Bembidion morulum Lee. Bembidion cf. morulum Lee. Bembidion sp. ~~-Pterostichus patrue!is DeJ. Pterostichus sp. Diplocheila sp. Chlaenius alternatus Horn Gen. indet·.

Dytiscidae Hydropor1n1 gen. 1ndet. Ilybius sp. Agabiini gen. indet. Gen. indet.

Hydrophilidae Gen. indet.

Hydraenidae _!Iydraena sp.

Staphylinidae Bledius sp. Eucnecosum brunnescens J. Sahlb. Eucnecosum tenue (Lee.) or

brachypterum (Grav.). Eucnecosum spp. Olophrum rotundicolle (C.

Sahlb.) Olophrum spp. Acidota crenata (Fab.J Pycnoglypta ~ptera Cpbl. Omaliinae gen. indet. Stenus spp. Euaesthetus sp. cl::-Pnilonthus sp.

Skeletal elements 1

LR PLR PLR

H HPLR H L HPL

p

H L p p

HPLR

HPLR PLR

HPLR HPLR

HP

p p

p

LR

HPLR

HPLR LR

HP HPLR

PLR HPLR HP

LR

Distribution group2

C A

A

A

A

A

A

A*

- -·· -----------.

38

Table 2--Continued

Taxa

Quedius (Raphirus) sp. Tachinus sp-.----Gymnusa sp. Aleocharinae gen. indet. Gen. indet.

Pselaphidae Reichenbachia sp.

Silphidae Thanatophilus sagax Mann.

Leiodidae Agathidium sp.

Scydmaenidae Gen. indet.

Scarabaeidae Aegialia sp. Aphodius sp.

Helodidae Cyphon or Paracyphon sp. Gen. indet.

Byrrhidae Cytilus sp.

Elateridae Hypolithus bicolor (Esch.)

Cucujidae Narthecius sp.

Chrysomelidae Donacia sp. ct. Phaedon sp.

Curculionidae sftona sp. Hylobius sp. Notaris sp. Erirhin1n1 gen. indet. Magdalis sp. Ceutorhynchini gen. indet. Gen. indet.

Skeletal Distribution elements

HP p

HP P R p

H LR

p

LR

LR

R PLR

R p

PL

p

LR

R LR

H PLR

HPLR H

PLR LR

HPLR

group

u

u

39

Table 2--Cont1nued

Taxa Skeletal Distribution

scolytidae Phloeotribus ~E?~ sw. Carphoborus carr1 Sw. c'arpnoborus cf. carri Sw. Polygraphus rufipennis (Kby.) Scolytus cf. p1ceae Sw. Scolytus sp. Ips latidens (Lee.) Pityophtfiorus cf. nitidus Mann. Pityophthorus sp. cf. Pityophthorus sp. Gen. indet.

elements

R R

L LR

R LR L LR

PLR HPLR H R

1 Skeletal elements: H L -- lett elytron; R

head; P -- pronotum; right elytron.

group

A A

A*

B*

2 Distribution groups: A -- generally northern, trans­american, may also occur in the Rocky Mountains as tar south as ·colorado; B -- generally northern, trans­american,but not north of so• N; C -- eastern; D -­northwestern; A* and B* indicate groups whose d1str1-butions are comparable to those of A and B respectively, but the southern limits extend into southern United States; U -- distribution unknown.

40

Table J

Total numbers of coleopter~n indiv1dugls of edCh tdxon per sample interval, At.hens North Quarry, Ill1no:ui.

'?'ax.a sample intervals

--------· t l 4 6 7 8 • 10 1t 12 tJ " ·---·---------------------~-·

CARABIDAE Sphaerode<us nit1dieollis

brevoort1 Lee. oyach1f1us integer Lee. oysch;.rius sp. l Pattobus sp. Bern610ion morulum t.,c. 2 2 l:!emb1Q1on cf. ~~ Lee. 2 Bemb1(hon sp. ) ' Pterostichus ~truelis Dej, ' ' li'Eeios~t1chus sp.--- ' ~locheda sp. Ch aenius alternatus Horn Gen. 1ndet-.-·--- 6

O'iT I SC l DA"£ Hydroporini gen. indet. ) ) ) ' I lXbius •P• ) ' l 2 l Agabuni. gen. indet. 2 l 7 l l 4 ' Gen. i.ndet:.. l 1

HYOROPHILI0AE Gen. indet.

HYDRAENIOAE Hydrae~ sp.

STAPHYllN!DAE Bledius sp. Eu'c'n"ecci sum brunnescens 2 5

J, sahlb. Eucnecosum tenue !Lee.) or · ~rachyote~um (Crav.} Eucnecosum spp. 5 OL:>enrum r-ot:undicolle 4 2 9 l

jC, Sahlb, l Olo£ht:um spp, Ac1dota crenata (Fab.J ~OSilY:~td aptera Cpbl. l Omal11nae qen. ~net. 4 4 l 5 2 Stenus spp. l l 4 ] ' l ' 4 5 itu.i'esthetus sp. 2 5 2 ct~1lonthu-s sp. l Ouedius !Raeh:i.rusJ sp. ~h~sp. Gymnu~ ~P· 1 Aleccnarinae gen. irtdet. l 1 Cen. indet.

FSELAPHIOAE Rekhenbdchia sp. 2 ' S [ :..?H!OAE ~~atop~~ §~ Mann. LE roe tDAE A5ath:.di.'J!:1: sp . ., .3C:YJ14AENtOAC c~n. tndet. 2

SCAR.'l.BAE I DAE Aegi.at!J! S\'.L

~~ Sp.

41

Table l--Continu~d

;u::.ooroA£ .9:.?hon or Pas_~~.D sp. G;;;n. 111det.

f!YR!<H1DA£ c·1ti,uz sp.

ELATERrDA£ Hynotitnus ~Le.£ {E1,,ctt.J

CUCUJtDAE ~arthecius sp.

CHRYS0M£L IC·A.E P~:.! sp. ct. ?taectot\ sp.

O:tJRCt::LIONIDAE ~SP, :rylcbius sp, Notc'lrts sp. ~r1r~in1n1 gen. 1::d~t. 'tc:!-1:da_l~ Sp, .:"""utcrh~·nch1n1 gen. :_ndet. -:;~n. ~ncet ..

SCGLYT!OAE ?~:oe~tr:buu occBae Sw. ,::,,~·onc~orua car-t! Sw. ::.:.r,mob::>r:.is ct". :arti. S;.,·, Poi•tqrapnus r-ufl~s !KDy. l Sc::d•1e.c:s ct. pL::eaoe 3,..;. Seo l'tt.us sp. 1£! ~dens IL.ac:.J ?tZYO!'ht').'.J(US cf. :'.i:..Ldu.s )l!ann. ~:o r: nor:us so. c::. 1' o~o.r::s 5p. C,;,r.. ;; .,;c.

: 2

2

2 !2

6

7 J l

' ;

42

PLATE I

Fossil Coleoptera from the Athens North Quarry and Biggsville sites. Bar scale represents 100 micronsr anterior is up in all cases. Numbers refer to North Dakota State University Geology Department accession numbers.

a. Pycnoglypta aptera, head, dorsal view, xlOO (ANQ 04-11-03).

b. Pterostichus patruelis, pronotum, x33 (BGV 10-02-08).

c. Olophrum consimile, pronotum, x50 (BGV 10-21-08).

d. Polygraphus rutipennis, left elytron, X50 (ANQ 12-07-03).

e. Olophrum rotundicolle, pronotum, x50 (BGV 13-09-04).

f. Carphoborus andersoni, right elytron, x75 (BGV 14-03-03).

g. Carphoborus carri, right elytron, x75 (ANQ 06-07-03) •

h. corraded pronotum of Bembidion sp., characteristics of corrasion include pitting and thinning of the chitin, xlUO (ANQ 12-04-06).

i. Bembidion morulum, pronotum, xlOO {ANQ 07-05-02).

PLATE I

I

I

4

,... e ..... .c: ,5 -Q. a, 0

6

44

Sample Interval

1 2 \ 3 ~ 4 > 5 ,( 6 , 7 f 8 ; 9 \ 10 ; 1 1 I

12 ~ 13 14

10 20 30 40 50

Number of coleopteran taxa

- .. - Number of aquatic tax a

Figure 9. Species diversity and numbers of aquatic tax a, A U1c,1;s North Quarry site, Illinois.

-------·····--45

A trellis diagram (Figure 10) based upon calculations

of Jaccard coefficients of similarity illustrates the

degree ot similarity between the beetle assemblages ot

pairs ot sample intervals. Each square in the matrix repre­

sents the comparison of two sample intervals which are

numbered along the diagonal. Possible values tor coeffi­

cients range trom O.O, for complete dissimilarity, to 1.0,

for identity. Each coefficient in this sequence has a

value of less than 0.7. The patterns which represent the

degree of similarity become progressively denser as the

value of the coefficient increases. Thus, the darker areas

in the trellis diagram mark the faunally related groups of

intervals. Intervals 2 through 10 form a large similarity

group, and intervals 11 through 14 form a group ot mutually

dissimilar intervals.

The dendrogram (Figure 11), generated by the applica­

tion ot the weighted pair-group method to the similarity

coefficients, shows the order of clustering of the sample

intervals. Intervals 2 through 10 form a large cluster to

which intervals 11 through 14 are added one at a time,

being no more similar to each other than they are to the

major cluster. These dissimilar intervals were grouped

into faunal zone A-1. The intervals comprising the large

cluster were designated faunal zone A-2. Sample interval

number 1, which was not included in the statistical analy­

sis, is designated as fauna! zone A-3. The faunal zones

., 4l = 0 ... GI C ::J <II

LI.

46

Jaccard similarity coefficients

{!] 0.6-0.7

A-3 I!] 0.5-0.6

T [!] 0.4-0.5 0 0 & 6 0 • 0 0 @ A-2 0.3-0.4

A-1

J_

0 •O 0 (9

0 0 0 (9 0 & @] 0.2-0.3 0 . 0 (9 (9 (9 • 9

0 0 0 0 0 0 0 10 [J 0.1-0.2

0 . 0 0.0-0.1 0 0 0 0

Figure 10. Trellis diagram of Jaccard similarity coefficients showing faunal zones, Athens North Quarry site, Illinois.

t, ,1:

1.0

.. 0.9 . ... C: .. 0.8 .!:! --.. 0.7 0 Q

>, 0.6 -;: !'! 0.5 § ..

0.4 "O

-... m

0.3 (.) . (.)

m .., 0.2

0.1 .

47

I

I I

I I •

l

--

--

2 10 3 4 5 6 7 8 9 13 12 14 11

Sample intervals

Figure 11. Clustering of sample intervals Athens North Quarry site, Illinois.

- !l,"

ii'

,, J

48

are illustrated in figure 10.

The percentage of corraded fossils for each interval

in the Athens North Quarry site is plotted against interval

depth in figure 8. The highest percentages occur in inter­

vals 13 and 12, corresponding to the mµck unit and the

lowermost part of the organic silt unit.

Sample interval number 1 yielded 6 juvenile gastro­

pods. These were identified by A. M. Cvancara as terres­

trial snails, probably belonging to the family Succineidae

(Cvancara, oral communication, 1984).

Biggsville site

At least l,395 coleopteran individuals, representing

25 families, have been identified from the Biggsville site.

A total of 114 taxa was identified to the family level or

lower, including 26 identified to species. Of the total,

31 percent was classified as aquatic according to the

criteria established by Leech and Chandler (1971). A sum­

mary of the fauna! data is found in table 9 of appendix A.

A taxonomic list of the beetle fauna, including designation

of aquatic taxa is presented in table 4. A quantitative

summary of the insect taxa and their numbers per sample is

given in table 5.

Scanning electron photomicrographs of several fossil

.. .,' ...

49

Table 4

Taxono~ic list at the coleopteran fauna tram the Biggsville site, Illinois. Order of genera tallows Arnett (1968). Or~er at species in the Carabidae, M1cropepl1dae, and scolyti~ae follows that of Lindroth (1961, 1963, 1966,

1968, 1969), \Campbell (1968), and Wood (1982), respectively.

Taxa

Carabidae I Notio hi lu!s sp. l:ffet isa s • Elaphius a.ericanus Dej. or

ripariusbL. Elapnruss • Dyi;-cEirTus \ .. t!m!_dus Lth. Dyschirius,sp. Patrobus Sf· Bembidion sp. (planiusculum grp.) Bemb1:~:ion <i,f. nigripes Kby. Bembidion !'lp. (dorsale grp.) Beinl:>idion qt. mimus Hay. ?embidion 1p. (versicolor grp.) Bembidion ~rulum Lee. Bembidion of:-morulum Lee. Bembidion s\p. (musicola grp. J Bembidion t,ransparens Gebl. Bembidion c~. transpar_ens Gebl. Bembidion cbncretum Csy. Bembidion fprtestriatum Mts. Bembidion cf. fortestri_<1_tum Mts. Bembidion p~eudocautum Lth. Bembidion frontale Lee. Bembidion s~. (anguliferum grp.) Bembidion sop. PferosITchU§ pensylvanicus Lee. or

adstrictu~ Esch. Pterostichu ~..!E.!:!elis DeJ. Pterostichu~ cf. £,atruelis Dej. Pterostichu spp. · Agonum-sp-.-(1)1elanarium grp.) Agonum corvus Lee. Agonum cf. dfcentis Say Agonum spp. IITplocheila PP• Chlaenius spl Gen. indet. \

Dytiscidae · Bidessini ge!. indet. Hydroporini ien. indet.

Skeletal elements'

R H

HP R H HPLR HPLR HPL

p p PLR p p p

H p

HPLR PL p

HPLR HP R HPLR

p HP HPLR

p PLR PLR

H LR p

p p

PLR L

p HPLR

R p

Distribution group1.

A

A

A

A A

B B

A

B

50

Taxa

Table 4--continued

Skeletal elements

_Ilybius sl. Agabiini 9en. indet. cf. Rhantu.s sp. Col mbete sp. Gen. inde.

Hydrophilidii!e Helophoru' sp Hydrochus sp. Hydrobius fuscipes L. Hydrobius:sp. Cercyon sp. Gen. indef.

Hydraenidae1 Ochthebiu$ sp. Hydraena fP· Gen. indef.

I Staphylinid<iie

Car elimu' sp. cf. Carpe imus sp. cf. Plat tethus sp. Bledius s . Eucnecosum brunnescens J. Sahlb. Eucnecosu tenue Lee. or

brach erum Grav. Eucnecosu spp. Olophrum ~onsimile (Gyll.J Olophrum otundicolle (C. Sahlb.J Olophrum pp. Pse hidon s sp. cf. Pycnowlypta sp. Omaliinae gen. indet. Stenus sp'. Euaesthet s sp. cf. Philo,thus sp. Bryoporusisp.

Tachypori ae gen. indet. Tachinus 1P· Gymnusa s. Aleochari~ae gen. indet. Gen. indet.

' Micropeplidle

Micropeplls Micropepllls Micropeplys

I

tesserula Curtis sculptus Lee .. cribratus Lee.

HPLR HPLR

L L

PLR

HPLR P R

LR R

LR HPLR

H LR HPLR

R

p p

H p p

p HPLR HPLR HPLR HPLR

p p

HPLR HPLR HP HPLR

LR L LR

H LR HPLR HPLR

LR LR

PLR

Distribution group

u

A

A A

A* B* C

51

Table 4--continued

Taxa

Pselaphidai Reichenbachia sp.

Silphidae i Thanatop~ilus sp.

I Leptodirid<j.e

Catoos s~.

Leiodidae ' I

Hyclnobiu~ sp. Agathidiujm sp. Gen. inde1t.

Scydmaenida~ Gen. indef.

Histeridae ' Gen. indef.

I

Scarabaeida'f Aegialia $p. Aphodius sp. Aphodiini[gen. Gen. indett.

HeloC!idae \ Gen. inde~.

i Byrrhidae I

Cytilus sp'. Gen. indet\·

Heteroceridaf' Gen. indetj.

Elmidae ·1

Dubiraphia.sp.

Cantharidae \ Gen.· indet

1

Nitidulidae Gen. indet

Cucujidae Narthecius 'sp. --~-------·-- I Gen. indet.\

indet.

Skeletal Distribution elements

HPLR

HP

L

R LR

LR

HP

LR H LR H L H

p

HPLR H L

p

PL

p

L

R R

p

group

52

Table 4--Continued

Taxa Skeletal Distribution elements group

cryptophagidae Anchicera ephippiata Zimm. L B

coccinellidae Nephus ornatu~ naviculatus (Csy. l R A*

Lathridiidae ct. Corticaria sp.

Chrysomelidae Macroplea sp. D0nac1a sp. Plateumaris_ sp. ct. Al1=i_~ sp.

Curculionidae Lepidophorus sp. Sitonini gen. indet. Dorytomus sp. Grypus~sp. . Notaris aethiops F. Notaris sp. &1.·r11Inini gen. indet. Bagous sp. Magdalis sp. ct. Magdalis sp. Hypol_<=~schtis-sp. Ceutorhynchus sp. Ceutorhynchini gen. indet. Cryptorhynchini gen. indet. Gen. indet.

Scolytidae Hylesinus sp. Phloeotribus piceae Sw. Carphoborus andersoni sw. Carphoboru~ carri Sw. Polygraphus rufipennis (Kby.) Polygraphus sp. Scolytus sp. Orthotomicus caelatus (Eich.) ~ borealis Sw. Crypturgus borealis Sw. Pithophthorus sp. ct. Pithophthorus sp.

PLR

H LR LR LR

p

H HP LR

R H LR H HPL

PL R

L L

p R R

HPLR

R R

LR R

H LR L LR L

R PLR

HPLR p

A

A D A A*

A* A A*

1 Skeletal elements: H L -- left elytron1 R

53

head; P -- pronotumi right elytron

~Distribution groups: A -- generally northern, trans­american, may also occur in the Rocky Mountains as far south as Colorado; B -- generally northern, trans­american, but not north of 50° N1 C -- eastern, D -­northwestern, A* and B* indicate groups whose distri­butions are comparable to those of A and B, respectively, but the southern limits extend into southern United States; (Ul distribution unknown.

th.

Tax.a

Table 5

Total nu/l'bers of coleoptecut indt7tdqals of ea.ct\ ta~on per sample interval, Siqgsv1l\e, Ill1ncis.

5&.mp!~ tnter,,al::.

1 2 J 4 5 n 7 a 9 10 11 12 13 14 1s 16 11 1a l9 20 21 22

CAI.U;lHDAE Notiophi lus sp. Bleth~- sp. Elaphrus americ.arrns Oej, o,

nearius L. Elap(1.r:.!.! Sp Dysch1rius timidus Llh. 5~sctunus :s:p. 1 1 1 J l Paaoous sp. 1 1 Bembid1on sp. ~iusculum grp.} Bemb{dion cf. nigr1pes Kby. Bemhtdion sp. (dorsa. le grp.) Bem~ cf, ~!!>. Hay. Bembtdton :,p. (verucolor qrp. I Bemb1d1on morulom Lee. l Bembidion er, morulum Lee. 1 '13emb1d1on sp. lmus1co-la grp.! Ben'lb1aiori transparens GebL Bemb1dton ~ranspatens Gebl. Bembtd100 concretum Csy. Bemb1d1on forte;str1-atum Mb1. 1 ' Semo1dion ct. fortestrtdtum Ml3. Bemb1dion pseudo~al,tum Lth. eemb1dion front.ale Lth, Bemb1.:.hon ~(~f:S_uliferum g:rp.J Bemhtdlon spp. l 1 1 l 1 Pterost1chus pensvlvarncus Lee.

or adstncrus t:"sch. Pterost1ch':!.:! p,.,1uuelis Oo?j. Pterostlc~us cf. oatru~Ji~ De), 2 Pterc1t1chus spp. l 2 Agonum sp.'"('me 1,mar i 11m q:.:p.) Aqonum corvws"'l:eC:----~:7oni1~ ct. decentis Say A~onum spp. UieToCh~lJ~ sp. Chlaz,r,iqs sr:,, Gen 1 nder, l 1 l l

0\11' l S1~ I D/1£ B1de1;5:;n1 :ien, indet. l!ydrc.p'..•r;:u q""n" u;det. '

7 5 2

l 2

I

6 l '

l l 1

2 2 t

I l

1

1 1

1 l l

l

2

1 7

7 10

l

1

1

t

l 3

l 7 l

l 11

' l

l

l

2

2 l

2

l

'

,. '

1 ' 1 3 1 2 l 1

l l 1

2

l ' J 1

J ' ' l 1 2

l 2 2 J 5

t 2

1 l

l l l

V' .,,

l

Table 5--Continued

Sample intervals Taxa

2 3 ' s 6 ] a 9 10 ll 12 13 14 15 16 17 18 19 20 21 22

Ilybiu~ sp. l l l l l l 3 l 3

AgabiLn1 gen. indet~ l 1 1 1 l 1 2 l 2 l l l

cf. Rhantus sp.

l

Collinbetes sp.

l l

Coen. 1ndeL l l ) 2 2 1

HYDROPHILIDA£ Heloehorus sp.

l l 1 ' 1 • 3 4 7 3 • 7 5 5

Hydr(?chus sp. l '

Hydrob1us fuscipes L. 1

Hydro61us sp. 1

1:ercyoo sp, 1 2 1 1 1 1

11 s 2 • 8 2 l 3 I • 4 4

Gen. iodet.

H'fOFA.ENIDi\t Ochthebios sp.

l l l l

J 4 1 8 21 18 23 15 54 11 1S 6 25 14 18 8

Hydraena sp. Gen. 1ndet,

l U1

l 2 l U1

STA.PHYLINlDAE car:eelimus sp, c!. careeiimus sp. cf. Platysteth~s sp. l tHedius sp. Eucnecosum brunnescens

J, Sahlb. Eucnecosum tenue Lee. or

1 l

hrachypterum Grav. Eucnecosum spp.

l l 7 20 6 21 17 5 23 l l 2 2 3

olophrum consirnile (Gyll.i l 3 13 2 5 2 l ' 2 l '

61o~hrum rotund1colle 1 4 l 2 l 2 24 2 10 11 'll 2 4 l l l

- ( • SahlP. • l ; ' l l l l l

l l Olophrum spp. Pseeh 1dom1s sp. cf. Pycnoglypta sp.

l

Oma l i inae gen. l ndet. l 2 l 4 ' J 2 2 6 2 l 2 3 2 3 •

~~ spp. l l l l 4 2 13 4 4191413 7 4 10 • 17 7 S 11 5 •

Euaesthetus sp. 1 l 3 l 3 5 2 6 l 17 2 6 l 6 • 5

Ct.Phi lonthus sp. 2 2 l 3 ' l 2

2 1

Brzoeorus sp. Ti;1,ch1nus sp. l l

TaChyporin~e gen. indet .• l l

G'fmnusa :;p. 2 l 2 2 J 2 5

A. ~0Cti"~r1nae gen. 1.ndet. l l ' " l 3 l I l 2 2 l 2 1

2 ' 2 l 1 l 2 1 ' Gen. ind et..

~ ___ _,,'

Tabte s--~~1:1nued

raxa Sample 1.ntervals

2 3 ' s ' 7 8 9 10 11 12 13 " 151b1118 192021?2

MfCROPEPLlDAE ~icropeplus tesserula Curtis Micropeplus sculptus Lee, 1 2 ~1cropeplus cr1bratus Lee. 2

PSELAPHIDAE Reichenbachia sp. 1 ' 2 4 28 8 • l l 2 2

51 LPHIOAE Thanatophilus sp.

LEPTODIRI.DAE Catops sp. 2

L£10D10AE H~dnobius sp. l As;ieth1dium sp. 1 2 5 14 l 2 l 3 l ' t J Gen.a.ndet, l

SCYDMAEN I DAE en

Gen. indeL l l 3 °' Hl5TER1DA.E ,,en, indet.

SC'.ARMH,EIDAE A~11/l11a sp. ru?hclh us sp. l I l 1 1 T\Phod1 ini gen. indet.. l 2 Gen. 1ndet.

HELODIDA.E Gen. 1 ndet. ' 7 ' ' l ' B'lRRH!DA.E Cyt1.lus sp. l 2 ' Gen. indet. l 2 ' HETf.ROCEFlIDAE Gen. H,det. 2

EV-1!:iAE i)u:::1:aoh1a sp. 1 l

:: kWriu, Rt DAE :;,..:n. i r,cet.

,ck,.tL • ..... ,>

Tabie S--Continueq

T21xa Sdmplc •ntervals

2 5 6 7 a 9 10 11 12 13 14 15 16 17 18 19 ;,'O 21 22

----- - ------------------------------NIT IDUL lDAE. Gen. rndat.

CUCUJ!DAE Nart.hecius sp. Gen. tndet.

CR'lPTOPttAG ! DAE ~D~ ephipp121t21 Z1mm.

COCCINELL!OAE Nephus ornatus

nav1cuiam-1csy.}

{.A'I'HRIDI IOAE cf. Corticat<d sp.

CHR'lSOHE.I,IOAE Macrcplea sp. Donac ia sp. Flateumaris sp. rr.--nt tea $p. DonacITna'e gen. indet.

CU!tCiJLlONIDAE ~pi do_E.Q£!.:.:i_!! sp. Sttonin1 gen. indet. Dorytomus sp. '!rn~.~ sp. ~;..! ~__E.hiops r. Notari$ Sf', Enrn1nin1 gen. indet. Ba9ous sp. ~dal1._~ sp.

cf. ~aqd~:!_ sp. !:!.z:poles;;~ sp. Ceutorhvnchus sp. Ccutorhynchini gen. indet, Cryptorhynchint gen. indet. Gen. indeL

Sw. Sw,

' J 4 3 8 4

2 2 3 3

, 2

l

2 12 6 1

'

U1 .-J

'l'axa

Cdrphoborus carr1 Sw~ Polygr6phus rufi'eemnu i Kby. J Potygr6phus sp. Scr.>lytus sp. Orthotomicus cael~tus tE1ch,l ~ boreall_! Sw~-Crypturqus bore~lis Sw~ P1tyoehthorus sp. cf. P1tyophthorus sp.

·rctble 5--Cont_~[\!lt!d

Sample 1ntetV6ls

1 2 l 4 5 6 7 8 9 10 11 12 B 14 15 16 17 18 19 2CJ 21 22

1 3 l 2 1

1 1 4

2 l 2 l l l l 2 5 l

en 0:,

59

species tram Biggsville are presented in plates I and II.

The named species are assigned to distribution groups in

table 4. The distribution groups of the species from the

Biggsville site will be described and illustrated along

with those from the Athens North Quarry site in the discus­

sion ot the climate in the region.

Maximum species diversity occurs in intervals numbered

19 and 20 in the organic silty loam unit. Maximum species

density occurs in interval number 10 in the peat unit.

Minimum species diversity occurs in interval number 5 of

the basal loess paleosol. Simple species diversity tor

each sample is illustrated in figure 12.

A trellis diagram {Figure 13) of similarity coef­

ficients shows two main areas with denser patterns. Inter­

vals 1 through 4 form a small group, and intervals 5

through 21 form a much larger group of intervals with

similar beetle assemblages. Interval 22 shows little

similarity with any other unit, with similarity coetticient

values of less than 0.1. The dendrogram (Figure 14)

reflects the same division of intervals. Intervals

5 through 22 are designated faunal zone B-1, and interval

numbers 1 through 4 are included in taunal zone B-2 (Figure

13) •

., .

< ...

D r :n::mrrrrr:

60

PLATE II

Fossil Coleoptera from the Biggsville site. Bar scale represents 100 microns: anterior is up in all cases. Numbers refer to North Dakota State University Geology Department accession numbers.

a. Bembidion frontale, pronotum, xlOO (BGV 20-01-09).

b. Micropeplus sculptus, right elytron, xlOO {BGV 18-04-03).

c. _Micropeplus .tesserula, right elytron, xlOO {BGV 11-02-01).

d-f. Dyschirius timidus

d. head, dorsal view, xlOO (BGV 10-03-13).

e. pronotum, xlOO (BGV 10-03-14).

f. right elytron, x55 (BGV 10-03-09).

g-h. Micropeplus cribratus

g. pronotum, xlOO {BGV 22-01-01)

h. right elytron, xlOO (BGV 22-01-02).

i. Bembidion fortestriatum, pronotum, xlOO {BGV 15-12-06).

j. Bembidion transparens, pronotum, xlOO (BGV 15-12-02).

·nwmwr·w·· mm

PLATE II

I I

C

I

h , I

.-)<,

62

Sample interval

1 ' 2 ' I 3 t 4 5

5 .6 T 7 t 8 ) 9 (

,.... ' e 10 > - ' .c 1 1 '( ... Q. 12 ' Q)

0 13 ; 14 t 15 '\

6 16 lo I

17 ; 18 + 19 ~

' 20 )

" 21 (

' 22 ... 10 20 30 40 50

Number of coleopteran taxa

- .. - Number of aquatic taxa

Figure 12. Species diversity and number of aquatic taxa, Biggsville site, Illinois.

.. II> C: 0 N

"' C:

T B-2

" B-1 "' II.

1 >-- 2 ()

0 • 3

(S) 0 © 4 . . 5

0 . 0 . . 6

• . . . 0 © 7

0 . 0 0 0 • • a 0 . . . 0 0 0 () . . . . 0 0 ()

0 . . . ·0 () ()

. 0 . . 0 () ()

. 0 © 0 0 ()

0 . . & () ()

. . . 0 () 0 0 . . . . 0 0 0 . . . 0 0 & . . . • 0 0 0 . . . . . 0 0 & . . . .

. . 0 0 0

63

Jaccard

9

() 10

() () 11

similarity coeffic ients

[!] o.

~ o .

[§] o.

5-0.6

4-0.5

3-0.4

2-0.3 @] o .

• 0 () 12 D o. 1-0.2

0-0.1

() () 0 () 13

() & • () () 14 0 0. 0 0 IS) 0 & • 15

0 () 0 () () 0 0 16

0 0 0 0 () 6) (9 0 17

0 (;) 0 0 0 & 0 () (9 18

0 0 () 0 & () () () i$) () 19

0 . 0 . ·O 0 ·O . 20

0 0 0 0 0 0 0 0 0 0 (9 (9 21

I 22

Figure 13. Trellis diagram of Jaccard similarity coefficients showing faunal zones, Biggsville site, Illinois.

,-;"

<I) -C .! .!:1 --II> 0 1,1

>, -·.: ~ .s

iJ, <I)

" .. I~ OS

1,1 1,1 OS ..,

1.0

0

o.

o.

o.

0.5

0.4

0.3

0.2

0.1

9.

I·

•.

'

.

.

1

r---a-

--~~

~-

2 3 4 5 6 8

I

I I

I _._

I .

I J_ I J_

_J_

9 12 13 10 7 11 14 15 16 19 18 17 21 20 22

Sample Intervals

Figure 14. Clustering of sample intervals, Biggsville site, IllJnois.

"' ..

DISCUSSION

Paleoenvironmental interpretations of the Athens

North Quarry and Biggsville sites are made to describe the

local environment that existed at each site and to infer

the climate of the region. A sequence of events is pro­

posed for each site.

Paleoenvironmental and paleoclimatic inferences are

based principally upon knowledge of the habitat preferences

and geographic distributions of extant species of beetles

that are represented as fossils. These data are supple­

mented by data from sedimentological and taphonomic studies

and from previously published paleontological and palyno­

logical studies. General knowledge of beetle habitat pref­

erences was gained from Arnett (1968), Blatchley and Leng

(1916), Dillon and Dillon (1961), Doyen and Ulrich (1978),

Leech and Chandler (1971), Moore and Legner (1979), and

White (1983), and from conversations with entomologists.

To avoid excessive repetition, general habitat data derived

from these sources is not cited in the text. The sources

of specific habitat information, however, are indicated.

Local Environment at the Athens North Quarry Site

The Athens North Quarry sequence is divided into three

faunal zones (Figure 10). Zone A-1 includes sample inter­

vals 11 through 14. Zone A-2 comprises sample intervals 2

65

66

through 10. Interval number 1 is assigned to zone A-3.

zone A-1

Faunal zone A-1 is defined on the presence of a total

of 58 beetle individuals belonging to at least 19 taxa.

Beetles from this zone are represented by species of

aquatic, hygrophilous, forest and miscellaneous habitats.

In general, though, zone A-1 is characterized by low spe­

cies diversity and density in which hygrophilous species

dominate.

The aquatic element of this zone is represented by one

individual of the dytiscid tribe Agabiini. Members of this

tribe are generally found in temporary pools or marshy

areas.

The modern counterparts of the majority of the taxa of

zone A-1 are hygrophilous, occupying moist terrestrial

habitats. The carabid genus Qyschirius burrows in moist

soil with sparse or no vegetation, usually near water

(Lindroth, 1961, p.132). Most species of ~emb~dion inhabit

water-marginal areas. Pterostichus patruelis is a very

eurytopic swamp species, occurring in Sphagnum bogs, as

well as more or less eutrophic Carex mires (Lindroth, 1966,

p. 505).

The rove beetle species of zone A-1 commonly inhabit

' ' "-'(\

67

moist organic habitats. Species of Eucnecosum dwell in

leaf litter under shrubs. as well as among sedges. Olophrum

rotundicolle lives in wet mosses and sedges in pond­

marginal habitats (Ashworth, 1980, p. 209). Species of

Stenus are usually found in wet situations such as bogs,

moist meadows, and marshes. The weevil genus Notaris and

members of the minute seed weevil tribe Ceutorhynchini are

also common in pond-marginal situations, feeding on sedges

and other types of vascular hydrophytes.

The presence of coniferous trees in the area during

the deposition of zone A-1 is indicated by several tree­

inhabiting species. The weevil Hylobius is known to feed

on and live beneath the bark of Pinus (Blatchley and Leng,

1916, p. 186). Hosts of the scolytids Carphoborus and

Polygraphus rufipennis include Picea, Abies, Pinus, and

other conifers (Wood, 1982, p. 369-374, 389).

Several species in zone A-1 represent other terres­

trial habitats: the ?ill beetle Cytilus is often found

inhabiting mosses; dry sandy areas near water are indicated

by the presence of the scarab Aegialia. Phytophagous chry­

somelids and weevils are also present. ~or example, Sitona

is often found on plants in open areas.

Zone A-2

Fifty taxa, including a total of 226 individuals, were

68

identified from zone A-2. The number of identified indi­

viduals per kilogram and the average number of taxa present

per interval in zone A-2 are approximately double that of

A-1 (Table 7, Appendix A). All but five species of the 27

taxa identified from zone A-1 are also present in A-2.

Similar to zone A-1, the beetle species of zone A-2 are

predominantly representative of aquatic, hygrophilous and

forest habitats. Also present are inhabitants of several

miscellaneous habitats. Compared to zone A-1, this zone is

generally characterized by higher species diversity and

density, an increase in the aquatic component, but con­

tinued dominance by hygrophilous species.

The aquatic fauna, represented by five families, shows

an increase in diversity from that of zone A-1. Members of

the family Hydrophilidae, like the Dytiscidae, are commonly

found swimming in shallow weedy ponds. The hydraenid

Hydraen~ is closely related to the hydrophilids but cannot

swim. Most species of Hydraena are found crawling in moist

matted vegetation, The chrysomelid beetle Donacia and

species within the Helodidae inhabit thick growths of

sedges and emergent vegetation.

The fauna of zone A-2, like that of A-1, is dominated

by hygrophilous forms. Taxa common to both zones include

the ground beetles Dyschirius, Bembidion, Pterostichus

£>_atruelis, the rove beetl.es, two species of Eucnecosum,

69

Ologhrum rotundicolle, Stenus, and curculionids of the I

tri~e Ceutorhynchini. The species Dyschirius integer bur-

rows in moist clayey soil at the margins of fresh water ' I I

whete the vegetation is depressed and the ground is bare in

spots (Lindroth, 1969, p. 149). Bembidion morulum has been I

obs4rved on moist peaty soil in coastal tundra and inhab-I

iti~g pond-marginal sedges (Lindroth, 1963, p. 389). Most I

spe~ies of Diplocheila occur near stagnant freshwater with

dense vegetation (Lindroth, 1969, p. 939). Similarly, I

Chlfenius alternatus has been found among sedges, grasses,

mos$es, and other water-marginal plants (Lindroth, 1969, p.

992) • I

I Species of the staphylinid Bledius commonly occur

togbther with species of the carabid Dyschirius (Lindroth,

196~, p. 132). Bledius also burrows into moist sand or I

mud!. The rove beetles Acidota crenata and Pycnoglypta

oft1en occur with Olophrum and other staphylinids of the

su9family Omaliinae in wet areas with sedges, grasses, moss

tus 1socks, accumulations of plant debris, and willow shrubs.

Sp~cies of the staphylinid Gymnusa are also associated with

swaimps, swampy places beside ponds and lakes, and bogs

(K~imaszewski, 1979, p. 14). Weevils of the tribe I I

Erfrhinini are marsh dwellers: many of the larvae feed on

aqtjatic plants and the adults are often found near open i

water.

' "

70

Forest-dwelling species increase in numbers and di-,

\iersity in zone A-2 • .!£§_ latidens is known (Bright, 1976,

~· 152) to inhabit shaded-out or broken limbs of Pinus,

T•uga canadensis (hemlock), and Pseudotsuga menziesii I

1pouglas fir). The habitats of other scolytids of this

zbne were described by Wood (1982). Hosts of Carphoborus

c~rri include Picea engelmannii (Engelmann spruce), Picea I

giauca (white spruce), and Picea rubens (red spruce). I ,

Pfloeotribus piceae prefers the shaded-out branches of

l~rge living Picea glauca trees. Polygraphus rufipennis I

uJually lives in dead or dying spruce, but is known to

o~cur in Abies and Pinus. Pityophthorus and Scolytus are '

fdund in both coniferous and broad-leafed trees. Bright

(~976, p. 179) noted that Pithyophthorus prefers dead or

d~ing twigs or small branches.

Several other taxa, present in zbne A-2, also indicate

th~ presence of a forest. The cucujid beetles, such as

Narthecius, live under bark of trees. Members of the click '

beltle family, Elateridae, are found on the foliage of

trtes and bushes as well as in rotten tree trunks. The

pi~e-feeding weevil Hylobius is present in this zone, as is

MaQdalis, a weevil which feeds in broken branches of dead

or ldying conifers or deciduous trees. The ground beetle,

~~roderus nitidicollis brevoorti lives on the forest

flgor, preferring moist places with mosses and dead leaves,

corri1monly near water (Lindroth, 1961, p. 29).

' I•,

' ':,1

71

Other beetle species from zone A-2 represent miscel­

la~eous habitats, several of which may also be related to

th~ forest floor. Members of the rove beetle genera I

Qufdius and Tachinus are generally found in moss, leaf

liltter, debris and decaying organic material (Smetana,

1971, p. 6; Moore and Legner, 1979, p. 270). The leiodid

Agathidium is known to inhabit fungi and slime molds on

decaying vegetation or under the bark of trees. Some mem­

bers of the family Scydmaenidae and the pselaphid genus

Reichenbachia are reported (Grigarick and Schuster, 1967,

p. 1) to live together with ants. Others inhabit leaf

mold, leaf litter and moss, or live under bark, logs, or

stones. The byrrhid Cytilus commonly lives in moss. The

scarab Aphodius is usually found in dung but also dwells in

fungi and beneath leaves and logs. Carrion beetles such as

Thanatophilus sagax commonly feed on carrion.

Zone A-3

This fauna! zone is composed solely of interval number

1, which contains no beetle fossils and therefore is not

included in similarity and cluster calculations. It does

however contain several gastropods. Unfortunately, the

individuals are juveniles and therefore very difficult to

identify with certainty. They probably belong to the ter­

restrial gastropod family Succineidae (Cvancara, oral

communication, 1984). Most terrestrial snails in this

region require a soil cover of moist organic debris in

'"

• :1

. .

72

which to reproduce. Both eggs and immature snails are

highly susceptible to drying and to the effects of direct

sunlight. In addition, most common terrestrial gastropods

depend on decaying organic matter and fungi for food

(Leonard and Frye, 1954, p. 400). Therefore, the occur­

rence of these snails in zone A-3 indicates the presence of

a moist organic habitat.

Sedimentology and taphonomy

Additional clue~ concerning the character of the local

environment were gathered from the characteristics of the

sediment. The relationship of the sample intervals to the

lithologic units and faunal zones at the Athens North

Quarry site are shown in figure 15. Intervals 13 and 14 of

zone A-1 are within the muck unit, which represents a

period of pedogenesis (Follmer et~., 1979). The over­

lying two intervals of zone A-1 and all of zone A-2 are

included in the organic-rich silt unit of the Peoria Loess.

Zone A-3 (interval number 1) belongs to the inorganic silt

unit of the Peoria Loess.

The degree of corrasion of the beetle fossils also

provided information about the environment at the Athens

North Quarry site. The highest proportions of corraded

beetle fragments are found in sample intervals 12 (54%) and

13 (49%) of zone A-1 (Figure 8). Characteristics of these

fragments include pitting and thinning of the chitin frag-

. • ,,

,

: ii

ATHENS NORTH QUARRY

-.c Ill

> ........ ... c. E

Lithology G)

G) ...... ... Q .!:

Inorganic slit 1

2

3 ~4-

4

5

organic silt s

7

8 '--5-

9

10

11

12

muck 13 ~a-

clavey silt · 14

Fauna! zones .­A-3

A-2

A-1

l

.c ... ,_ c. E G> .... C

~5

~s-

-1-

BIGGSVILLE

Lithology

---

loess paleosol

organic silly loam

,

peat

silty peat

organic silty loam

organic sandy loam

gravel

Figure 15. Sample intervals, lithology and fauna! zones of the Athens North Quarry and Biggsville sites, Illinois.

~;;,. ~· ~ - ;- "~'.

"' > ... Ill ... C:

1 2 s 4 5 e 7 8 e

10 11 12 13 14 15 18 17 18 19 20 21 22

Fauna I zones

T B-2

B-1 "" w

..

. '

74

ments (Plate re). Although chemical alteration of chitin by

the leaching of lipids may also cause post-mortem pitting,

the pitting and thinning of the fossils may be predomi­

nantly the result of microbial degradation of the chitin in

the soil. According to Atlas and Bartha (1981), chitin in

the soil is subject to attack by various fungi and bac­

teria, including actinomycetes. The role of actinomycetes

in the degradation of complex organic substrates such as

chitin was discussed by Lacy (1973). These bacteria are

widespread in all soils. Their numbers and importance are

dependent upon the soil type, depth, moisture, aeration, pH

and organic matter content. Soil moisture and pH appear to

be the most important factors. Most soil actinomycetes are

aerobic and prefer neutral or slightly alkaline soils.

The preservation of chitin in an aerated soil occurs when

the degradation process is inhibited by protective layers

of wax and protein on the chitin fragments. Lacy also

pointed out that there are generally few actinomycetes in

acidic peaty soils, especially if they are water-logged.

Pollen and plant macrofossils

J.E. King (1979) described the palynology of the

Athens North Quarry site, and F. 8. King (1979), the plant

macrofossils. Pollen samples were studied from a SO-cm

thick section from the lower organic-rich part of the

Peoria Loess, dating between about 25,000 and 23,000 14c yr

8.P. The dominant taxa, which comprised about 70 percent

75

Other of the total upland flora, were Picea and Pinus.

common taxa throughout the section were Quercus (oak),

Graminae (grasses) and Tubuliflora (the sunflower group)•

Components which were present only in the lower part of the

section include lix (willow), and Morus

(mulberry). J.E. King interpreted the pollen as repre­

sentative of a forest composed of pine and spruce with a

grass and herb understory. Occasional oak trees may also

have grown in the area, or their pollen grains may have

been transported in by prevailing winds from source areas

to the southwest. Although the disappearance upward of

birch, willow, and mulberry might imply a shift to a

slightly colder climate, the pollen from this site did not

suggest any type of major climatic change during this

period.

F. B. King (1979) examined collections of plant macro­

fossils from a bulk sample of the organic-rich zone of the

Peoria Loess, dating between about 25,000 and 22,000 1~C yr

B.P. Present within this sample was wood of both Picea and

Larix laricina (tamarack), and needles of Picea, Abies

balsamea (balsam fir) and cones of Picea mariana (black

spruce). Seeds of Cyperus (sedge), Hypericurn (St. John's

wort) and Viola (violet) were also identified. According

to F. B. King, all of these genera include species adapted

to growth in bogs or under moist coniferous forests. A

change in the composition of the forest during the deposi-

' ' .

76

tion of the upper part of the organic silt unit (Figure 6)

was indicated by the addition of macrofossils of Larix

Be-laricina and Abies balsamea to the floral assemblage.

cause these species were not represented by pollen grains

in the lower part of the organic silt unit, dating between

about 25,000 to 23,000 ~Cyr B.P., F. B. King inferred

that they may have been growing on the site at a slightly

later time than the pollen record, that is, between about

23,000 to 22,000 '~c yr B.P. Both of these species, along

with ficea mariana, are known to occur together frequently

under moist or peaty conditions. Therefore, evidence from

the plant macrofossils supported J. E. King's (1979)

interpretation of the site, but suggested a moister coni­

ferous forest than suggested by J.E. King.

Sequence of events

By combining the data derived from the sediment, vege­

tation (J. E. King, 1979; F. B. King, 1979), beetles and

gastropods, an hypothetical model of the environment and

the events at the location of the Athens North Quarry site

for the period of about 25,000 to 22,000 14 c yr B.P. can be

constructed.

The .muck unit of zone A-1 probably represents a soil

zone in a moist coniferous forest adjacent to the marshy

margin of a pond or lake. High proportions of corraded

fragments indicate microbial activity in the soil; this

' ' •

77

argues against a bog environment with acidic or water­

logged conditions. The forest was composed dominantly of

pine and spruce with lesser components of birch, willow,

mulberry, and possibly oak, with an understory of grasses,

herbs, and shrubs. Nearby marshy areas of sedges, rushes,

wet mosses, and leaf litter were interrupted by patches of

moist soil with sparse or no vegetation. The lithologic

change from muck to organic silt perhaps indicates the

onset of loess deposition in response to the expansion of

glacial ice into the Lake Michigan basin. Slow loess

accumulation (approximately 7 cm per 100 years if a con­

stant rate of deposition is assumed) continued through the

deposition of the upper half of zone A-1 and throughout

zone A-2.

Expansion of the nearby body of water occurred some­

time near the end of zone A-1 and the beginning of zone A-

2. Open water habitats were available at or very near the

site as evidenced by the presence of dytiscids and hydro­

philids in zone A-2. Low percentages of corraded beetles

may be an indication of reduced microbial activity due to a

water-logged substrate. The forest at the site was domi­

nated by pine and spruce with a grass and herb understory

during the deposition of zone A-2. Several beetle species

from other terrestrial habitats in this zone provide evi­

dence for a forest floor with rotting logs, decaying plant

debris, fungi, leaf molds, s.lime molds, mosses, dung, and

' . . '

.j

78

rotting carrion. The addition of tamarack to the forest

was perhaps a reflecti-0n of an increase in soil moisture.

The frequency of occurrence of scolytid beetles that live

in dead and dying spruce increases upward in zone A-2.

This increase may also indicate increased soil moisture

beyond the tolerance of some of those trees.

A possible shift to slightly colder climatic condi­

tions during the deposition of zone A-2 was suggested by J.

E. King (1979) to explain the disappearance of birch,

willow, and mulberry near the top of the pollen profile of

the Athens North Quarry site. overall, however, he be­

lieved the pollen evidence indicated a stable vegetation

between 25,0oo- and 23,000 14c yr B.P. The appearance of

tamarack and balsam fir within the forest during the period

23,000 to 22,000 14 c yr B.P. was interpreted by F. B. King

(1979) as perhaps representative of the beginning of the

shift from a mild interstadial climate to colder and moist­

er, full glacial conditions.

Hygrophilous beetle species dominate the fauna

throughout zone A-2. They indicate the presence of a

thickly vegetated pond-marginal zone composed of sedges,

emergent vegetation, willow shrubs, tussocks of moss, and

matted plant debris. Expansion of the pond-marginal habi­

tat toward the center of the probable lake near the end of

zone A-2 is inferred from_the general decrease in species

" ' '

79

density and diversity of the beetles from open-water habi­

tats. Areas of exposed peaty or clayey soil are also

implied by the presence of burrowing carabids and staphy­

linids.

Zone A-3 is set apart from the preceding zones by the

complete lack of beetle fossils, the appearance of terres­

trial snails, and an abrupt change in lithology. The cause

of the sudden termination of the organic sequence is un-

clear. It may be related to increased loess deposition as

Woodfordian glaciers approached the area. The presence of

the gastropods would seem to indicate a continued moist

organic habitat, but no other evidence for the character of

the local environment is present.

Overall, the interpretations of the beetle assemblage,

the pollen, and the plant macrofossils are in agreement.

The shift to colder, moister conditions proposed by F. B.

King for the period between 23,000 and 22,000 1~C yr B.P.

at the Athens North Quarry site, however, is not reflected

by the beetle data.

Ill .• -

'

' ,'

•

80

Local Environment at the Biggsville Site

The compositions of the beetle assemblages from the

Biggsville site and Athens North Quarry site are very

similar. Only seven species from the Athens North Quarry

site are missing at Biggsville. Four other Athens North

Quarry species may have been present at Biggsville but

corresponding specimens from the Biggsville site were iden­

tifiable only at higher taxonomic levels. Species diversi­

ty and density, however, is much greater at the Biggsville

site. The Biggsville beetle fauna includes 114 taxa

whereas only 61 taxa were identified at the Athens North

Quarry site. Variations in numbers of fossils between the

two sites are shown in tables 8 and 9 of appendix A. The

Biggsville site also has a higher overall percentage of

aquatic taxa than does the Athens North Quarry site (31% at

Biggsville vs 24% at Athens North Quarry).

The Biggsville section is divided into two faunal

zones. Zone B-1 includes sample intervals 5 through 22.

Zone B-2 includes intervals 1 through 4 (Figure 13). The

break between the two zones occurs within the basal loess

paleosol unit but is not reflected in a change in lith­

ology. The relationship of the sample intervals to the

lithologic units and the faunal zones of the Biggsville

site is shown in figure 15.

' . .

' ,,

8 J.

Zone B-1

Faunal zone B-1 is based upon a total of 1,347 beetle

individuals belonging to a minimum of 114 taxa. Beetle

fossils from this zone represent aquatic, hygrophilic,

forest and miscellaneous habitats. Zone B-1 is charac­

terized by high species diversity, dominance of the fauna

by hygrophilic forms, a moderate open-water habitat com­

ponent, and a small but diagnostic forest habitat

component.

The aquatic habitats interpreted for zone B-1 include

open-water, pond-marginal, and riparian situations. A

shallow weedy body of open water bordered by thick growths

of sedges and other vascular hydrophytes is indicated by

fossils of dytiscids, hydrophilids, helodids, hydraenids,

chrysomelids of the subfamily Donaciinae, and an aquatic

curculionid. The proximity of a stream is inferred from

the presence of several aquatic and hygrophilic species;

all of the members of the Bembidion planiusculum species

group described by Lindroth (1963) are strictly riparian,

usually confined to the stoney-gravelly banks of running

waters. The hydrophylids Hydrochus and Helophorus are

known to dwell in streams as well .as ponds. The riffle

beetle Dubiraphia is found chiefly on aquatic vegetation at

the margins of slow-moving streams, as well as in lakes and

ponds (Leech and Sanderson, 1959, p.986). The hydraenid

Ochthebius is common in both riparian habitats and lakes.

82

Hygrophilic forms comprise the largest component of

the coleopteran fauna of zone B-1. A vegetation-rich,

pond-marginal habitat is indicated by several families of

these. Weevils, such as Grypus, are commonly found on

Equisetum (horsetail). Dorytomus commonly occurs on willow

(Blatchley and Leng, 1916, p. 193) and species of

Hypoleschus feed on the leaves of willow and other decid­

uous trees. The presence of additional weevils of the

curculionid tribes Erirhinini and Ceutorhynchini also indi­

cate a wet pond-marginal environment. Rove beetle species,

including Olophrum rotundicolle, Olophrum consimile,

Eucnecosum, Psephidonus, Stenus, and Gymnusa, also indicate

a swampy or very wet pond-marginal habitat composed of

sedges, grasses, tussocks of moss, and accumulations of

moist plant debris.

The hygrophilic component of zone B-1 also includes a

rich Bembidion fauna. Bembidion morulum and other members

of the Bembidion musicola species-group usually occur among

mosses and leaves on firm soil (Lindroth, 1963, p. 389,

387). Bembidion frontale prefers clayey soil mixed with

organic material, often shaded by dense vegetation, at the

margin of relatively small bodies of standing water

(Lindroth, 1963, p. 402). Bembidion pseudbcautum occurs in

rich swampy vegetation (Lindroth, 1963, p. 399). Bembidion

transparens is found at the border of standing waters of a

eutrophic or mesotrophic character, with a rich vegetation

,;,.

83

of carex (sedge), Eleocharis palu.stris (spike rush), Typha

latifolia (cattail), Comarum, and Menyanthes (buckbean)

(Lindroth, 1963, p. 394). Bembidion concretum occurs in

wet, eutrophic or mesotrophic swamps, with organic rich

soil, and luxuriant vegetation including Carex and Typha

latifolia (Lindroth, 1963, p. 396). Bembidion

fortestriatum most often occurs in swamps with Carex,

Comarum, Menyanthes, and Similacina (Lindroth, 1963, p.

397). The habitat of both~- ~~parens and~- concretum

often includes a carpet of mosses, particularly

Drepanocladus aduncus, but never Sphagnum (Lindroth, 1963,

p. 393-396).

Other carabids found within this zone, such as Agonum

corvus, Pterostichus patruelis, Ch.laenius, and Blethisa,

occur in swamps or along standing water among sedges,

rushes, and other pond-marginal plants. Patches of bare

mud in this area are indicated by burrowing species of the

Heteroceridae, Staphylinidae, and Carabidae. Heterocerids

and the rove beetles Carpelimus and Bledius are found in

wet sand, mud, or clay on the banks of ponds or streams

(Moore and Legner, 1979, p. 22). The ground beetle

Dyschirius commonly occurs together with Bledius and

heterocerids, digging burrows in soil with sparse or no

vegetation, usually near water (Lindroth, 1969, p. 132).

A diverse group of scolytid beetles is present in zone

'.l./ I -~

84

B-1. Their ecology was described by Wood (1982).

Phloeotribus piceae, Ips borealis, and Carphoborus

andersoni prefer the shaded or broken lower branches of

Picea glauca. The hosts of Hylesinus belong to the genus

Fraxinus (ash). Carphoborus carri, Polygraphus rufipennis,

and Crypturgus borealis live in species of Picea, Abies, or

Pinf:rs. The hosts of Orthotomicus caelatus probably include

any species of Picea, Pinus, or Larix. Species of

Pityophthorus are found in dead or dying twigs or small

branches of both coniferous and deciduous trees. Scolytus

feeds in both broad-leafed and coniferous trees.

Other forest indicators are also present in zone B-1.

The weevil Magdalis feeds in broken branches or in dead or

dying trees. The cucujid Narthecius lives under the barks

of both conifers and deciduous trees. The Nitidulidae feed

on tree sap, and the Cantharidae are found on herbage and

foliage. Several taxa in zone B-1 live in habitats charac­

teristic of a forest floor. Most species of the hydro­

philid genus Cercyon are strictly terrestrial but live in

damp places, especially in decaying vegetal debris, or dung

(Leech and Chandler, 1971, p. 345). Leiodids, such as

Agathidium, live in fungi or slime mold on decaying plants

or under bark, and Hydnobius has been collected in rotting

fungus. Species of the micropeplid, Micropeplus, the

short-winged mold beetle, Reichenbachia, the pill beetle,

Cytilus, the minute brown scavenger beetle, Corticaria, the

, 85

silken fungus beetle, Anchicera ephippiata, and the

Scydmaenidae indicate the presence of decaying vegetable

debris, fungi, moss, mold, and rotting logs. The

Histeridae and the scarab Aphodius occur in dung as well as

fungi and decomposing leaf litter. The members of the

chrysomelid subfamily Alticinae feed on roots of plants,

and the adults and larvae of the lady-bird, Nephus ornatus

naviculatus, are often found on stems or leaves of plants,

preying on mealybugs (Gordon, 1976, p. 278).

Other members of the fauna of zone B-1 are represent­

ative of more open areas. The weevil Lepidophorus and the

scarab Aegialia are characteristic of open sandy areas,

often near water. Species of the ground beetle Notiophilus

are heliophilous and rather xerophilous, occurring in open

country, where the vegetation is sparse and the soil con­

sists of gravel, or in thin forest land (Lindroth, 1969, p.

90). Members of the ground beetle Pterostichus adstrictus

species-group occur independently of open water (Lindroth,

1966, p. 485), whereas Patrobus inhabits open country but

is more or less hygrophilous (Lindroth, 1969, p. 177).

zone B-2

Zehe B-2 exhibits a sharp drop in both species diver­

sity and fossil density. The average number of taxa per

sample interval is less than half of that of zone B-1, and

an average of less than one-sixth as many identified indi-

86

viduals per kilogram are present (Table 9, Appendix A).

Only four additional species made their first appearance

during the time period encompassed by this zone.

A total of 50 coleopteran individuals is present in

this zone, representing at least 27 taxa. The majority of

the beetles from this zone represent aquatic and hygro­

philic habitats, plus several miscellaneous habitats. The

general trends which characterize this zone are lower spe­

cies diversity, continued dominance by hygrophilous forms,

continued presence of open-water aquatic beetles, and a

sharp decrease in forest taxa.

Although the density of water beetles decreases

sharply in zone B-2, aquatic habitats remain important.

The dytiscids and hydrophilids present are common inhabi­

tants of shallow weedy ponds. Representatives of a ripar­

ian habitat were not found in this zone.

The greatest percentage of beetle species represented

by fossils in zone B-2 lives in water-related habitats.

Weevils in the tribes Ceutorhynchini and Cryptorhynchini,

as well as the rove beetles Olophrum and Stenus, are com­

monly found feeding on sedges and other pond-marginal

plants. The ground beetle Diplocheila lives near stagnant

fresh water where the vegetation is dense (Lindroth, 1968,

p. 939). The Elaphrus species represented at the

Biggsville site, either Elaphrus americanus or E. riparius,

( ' '

87

lives on moist open ground where the vegetation is discon­

tinuous or lacking (Lindroth, 1969, p. 115-116) · The

carabids Dyschirius and Bembidion also indicate the proxi­

mity of water and bare patches of soil. Patrobus is more

or less hygrophilous but usually inhabits open country

(Lindroth, 1969, p. 177).

only. one forest-dweller, Pithophthorus, is represented

in zone B-2; as in zone B-1, this scolytid indicates the

presence of wood. Forest floor habitats, indicated by the

staphylinid Tachinus, the pselaphid Reichenbachia, and the

byrrhid Cytilus, include mold, moss, leaf litter, and other

decaying material.

Sedimentology

Hallberg and Baker (written communication to D. P.

Schwert, 1981) concluded that the Biggsville lithologic

sequence (Figure 7} represents an infilled channel cut into

Mississippian bedrock. The fill consists of fluvial depo­

sits, organic sediments, a peat sequence, and finally by

Late Wisconsinan Peoria Loess. Fluvial deposits of coarse

cobble gravel form the base of the sequence; these are

overlain by beetle-bearing organic sediments. The grain

size of the sediments decreases upward as the organic

component of the units increases. A trend of decreasing

mineral material and increasing organic matter continues

upward through a sequence of silty peat and finally into a

88

woody fibrous peat unit, Formation of the peat probably

began sometime near 24,000 14 c yr B.P. At about 22,000 14 c

yr B.P., an influx of mineral material became mixed with

organic fragments and formed the organic silty loam to

silty clay unit. The overlying basal loess paleosol indi­

cates a period of pedogenesis which ended at about 21,000

14c yr B.P., terminated by the deposition of the overlying

massive loess unit.

Sequence of events

Combination of fossil beetle habitat data and con­

clusions drawn from the character of the sediment resulted

in the construction of the following interpretation of the

events at Biggsvitle during the period of approximately

28,000 to 21,000 14 c yr B.P.

The fossil fauna of zone B-1 indicates a setting that

included a shallow weedy body of open water surrounded by

sedges, rushes, reeds, cattails, and other aquatic plants.

Two species of Bembidion indicate that carpets of moss at

the site were possibly composed of Drepanocladus rather

than Sphagnum. Areas of firm soil covered by moss and

leaves, patches of bare mud, and accumulations of plant

debris were also present near the water. The possible

proximity of a stream with sandy or gravelly banks is

inferred from the presence of several riparian species. A

nearby coniferous forest with open areas was probably domi-

89

nated by Picea, and included species of Fraxinus, and

possibly Pi~, /\.bi~, and Lari~. The floor of the forest

was moist, composed of moss, slime molds, fungi, molds,

rotting logs, and decaying vegetal debris.

The environment during the deposition of zone B-2 also

included a body of open water bordered by sedges and other

aquatic plants; no indicators of a riparian habitat are

present. Areas of moist open ground with moss, molds, and

decaying organic material plus bare patches of soil were

present; but there is little evidence for the persistence

of the coniferous forest in this zone.

The most likely modern analog for the Biggsville site

during the period of 28,000 to 21,000 ~Cyr B.P. is the

margin of a bog in a spruce-dominated coniferous forest.

The development of such an environment was discussed by

Smith (1966, p. 182-185). Bog development begins with the

colonization of open water by submerged and then floating

plants. The accumulation of organic material raises the

pond or lake bottom, and plants such as reeds, sedges,

cotton grass, buckbean, and marsh cinquefoil begin to grow

in the shallow margins. Mos·ses creep in and fill the space

between the plants. As the mass of marginal vegetation

thickens and expands, peat is formed and the mat extends

out over the water. Shrubs surrounded by moss grow on

thickened areas at the shoreward edge of the mat, followed

90

by the invasion of forest species, especially Picea and

Larix, which advance as the mat closes toward the center of

the lake. Larix is eventually replaced by Picea mariana,

Thuja (aborvitae), and Abies balsamea.

The Biggsville bog meets many of Smith's (1966) re­

quirements for a bog, such as the accumulation of peat and

the presence of cushion-like vegetation. It was atypical

perhaps in the composition of the mosses, because two

Bembidion species indicate that the presence of Sphagnum

was unlikely. The inference that the site was at the

margin of the bog is based upon the possible proximity of

forest and forest floor habitats. The forest surrounding

the bog may not have been very dense, and it appears to

have diminished near the top of the sequence. This decline

may have been in response to the infilling of the bog by

Woodfordian loess.

The sequence of events at the Biggsville site ended

with a period of pedogenesis. Destruction of chitin during

this process may account for the decline in numbers and

diversity of beetle fossils near the top of the section

(interval numbers 1 through 6).

·'

1 ~

1 ~

'

I. al I ¥

91

Regional Climate Interpretation

Because of the geographic proximity and overlapping

ages of the Athens North Quarry and Biggsville sites, and

similar ranges of their species, the faunas of the sites

were combined in a study of the distribution patterns

within the beetle assemblages. Maps of beetle distribu­

tions representative of the six distribution groups de­

scribed below illustrate that many of the beetle taxa

recorded as fossils do not live in the area at the present

time. The distribution groups are described as follows:

Group A species {Figures 16 - 18) have northern

transamerican distributions ranging from Alaska and the

northern Yukon to British Columbia in the west, and from

Newfoundland to the Great Lakes region in the east. Some

spec may also occur in the Rocky Mountains as far south

as Colorado. Group A* (Figure 19) has similar northern

limits, but the ranges of the species of this group extend

farther south into the United States, in some cases as far

south as Arizona and Florida.

Group B taxa (Figures 20 and 21) also have generally

northern transamerican distributions, from southern British

Columbia and Washington in the west, to the Great Lakes

region in the east. However, the ranges of these species

do not extend north of 50° North Latitude. Species with

similar northern limits but with expanded southern distri-

0 I 0

ml syo l\ffl 800

92

• !I!.!! borealla • Athens North Quarry

'Q Blggsvllle o S'lale or provlnclal record

Figure 16. Present distribution of~ borealis (Coleoptera: Scolytidae). Locality data from Bright (1976, p.163) and wood (1982, p.681-683). Distribution group A; Athens North Quarry site.

0 ml 500

o km eoo

• Chlaenius alternatus

93

A Athens North Quarry

Q Blggsvllle

Figure 17. Present distribution of Chlaenius alternatus {Coleoptera: Carabidae). Locality~ data from Lindroth (1969, p. 993), CNC and USNM. Distribution group A; Athens North Quarry site.

0 mi 500

o km 800

• Bembldlon morulum

o S1a1e or provincial record

94

• Athens North Quarry

'v Biggsville

Figure 18. Present distribution of Bembidion morulum (Coleoptera: Carabidae). Locality data from Lindroth (1963, p. 389). Distribution group A; Athens North Quarry and Biggsville sites.

95

' ' I I o ml 500

o km aoo " ..,' __ ,__J

·- - ' 0 ' '·''""\

' '

• Orthgtomlcus caelatus A Athens North Quarry

o Stale or provincial record ~ Blggsvllle

Pigure 19. Present distribution of Orthotomicus caelatus (Coleoptera: Scolytidae) Locali.ty data from Bright (1976, p. 149) and Wood (1982, p. 663-664) Distribution group A*; Biggsville site.

0 ml 500

O km 800

• sembldlon trontale

I I I

96

.. J.:--"-.J •. , ... -. ...

"\

6 Athens North Quarry

'iJ Biggsville

Figure 20. Present distribution of Bembidion frontale (Coleoptera: Carabidae). Locality data from Lindroth (1963, .p. 402) and CNC. Distribution group B; Biggsville site.

97

• Agonum corvus A Athens North Quarry

o S1a1e or provlnclal record 'v Biggsville

Figure 21. Present distribution of Agonurn corvus (Coleoptera: Carabidae). Locality data from Lindroth (1966, p. 603). Distribution group B; Biggsville site.

.1

98

butions are assigned to distribution Group B* {Figure 22).

Group C taxa (Figures 23 and 24) have eastern distri­

butions, occurring only as far west as Lake Winnipegosis in

Manitoba and Texas. The eastern limit of this group ex­

tends from Newfoundland in the north to Georgia in the

south.

Distribution group D has one member, Carphoborus

andersoni. Its present range is in northwestern North

America, from Alaska in the west to northern Northwest

Territorities, and northern Saskatchewan to the east

(Figure 25). Although this species is rare at the present,

it is a relatively common fossil in Wisconsinan assem­

blages. The modern and fossil distribution of this species

was described by Ashworth (1977, p. 1627; 1980, p. 208),

Morgan and Morgan (1980, p. 1121-1122), and Schwert and

Morgan (1980, p. 107). Although Wood (1954) had indicated

that this species may include another similar species,

Carphoborus both have been retained as separate

species in his recent (1982, p. 380-382) revision of the

family Scolytidae. Schwert and Morgan. (1980, p. 107)

pointed out that the occurrence of C. andersoni is rare and

that its modern distribution may be extended upon more

intensive sampling in eastern Canada.

0 I 0

'"":"tf 0 : 1

,_ ... _~ \ 0

' ' . -""''"' ' 0 , r-..J---.. 4 o

' ' I.. t

99

t , ,_ __ _,__ \. 0 , ' ~-----

\. I ' t o,, tOs \ '\ ,-----t-----1.------' ---,! I r-~ '; .' Q f Q I "'-- !

-~ : I -- i -.. • r ' ..: _,..__ ' - " " ,J....$~"""") •. ,.,."", ·,

'

• lps 11111gens 4 Athena North Quarry

o State or provincial record 'Q Biggsville

Figure 22. Present distribution of Ips latidens (Coleoptera: Scolytidae). Locality data from Bright (1976, p. 155) and Wood (1982, p. 677). Distribution group B*; Athens North Quarry site.

100

• Mlerooeolus crtbratus • Athens North Ouarry

0 State or provincial record <;J Blggsville

Figure 23. Present distribution of Micropeplus cribratus (Coloeptera: Micropeplidae). Locality data from Campbell (1968, p. 246). Distribution group C; Biggsville site.

0

0

ml 500

km 800

101

............ J I ' -- ..J I 1- "- i t , ,---...L_ ' j ' t. __ ... __

, I f I , J f I \

'\ ,---... -f-----l.------' --"'1 1 r---: r-, ' t.. • -~ ' -... __ .. ,

'·.. : J r-,, _ _, ... --,-...! ;"-.!sii.----1 •. ft ,.~-...

"·

• Sphaeroderus nltldlcollls brevoorli

& Athens North Quarry

Q Biggsville

Figure 24. Present distribution of Sphaeroderus nitidicollis brevoorti (Coleoptera: Carabidae). Locality data from Lindroth {1961, p. 30). Distribution group C; Athens North Quarry site.

0

0

ml 500

km eoo

• Carpnoborus andetsonl

102

6 Athans North Quarry

Q Biggsville

Figure 25. Present distribution of Carphoborus andersoni (Coleoptera: Scolytidae). Locality data from Bright (1976, p. 101) and Wood (1982, p.380-382). Distribution group D; Biggsville site.

103

Figure 26 shows the number of modern taxa represented

in the Athens North Quarry and Biggsville beetle assem­

blages which presently occur in the provinces of Canada and

the states of the United States. The greatest density of

occurrences is in the southern boreal forest region of

southern Canada and the northern Great Lakes region. With

the exception of Carphoborus andersoni, the mixing of spe­

cies with disjunct ranges is minimal,

During the initial stages of this study, it was ex­

pected that the transition from warmer interstadial condi­

tions to cold full-glacial conditions would be reflected by

a corresponding transition from a boreal beetle fauna to a

fauna with tundra or tree-line affinities. However, no

marked change in the species composition of the beetle­

bearing faunal zones was apparent. The abrupt demise of

the fauna of both sites by loess burial was perhaps more

related to the depositional processes associated with wind

than with a regional change in temperature. Therefore, the

temperature change which allowed a tundra fauna described

by Schwert et~- (1981) to survive in eastern Iowa at

about 17,000 11c yr B.P. was not det~cted at the Athens

North Quarry or Biggsville sites.

No clear pattern of the disappearance of thermophilous

species and the appearance of cold-adapted species resulted

from the study of the Biggsville fauna. Further, the first

! I I I I I l I I

104

A Athens North Quarry

Q B!ggsvllle

Figure 26. Numbers of taxa represented in the Athens North Quarry and Biggsville beetle assemblages which presently occur in the provinces of Canada and the states of the United States.

105

or last appearances of a species may be the result of

insufficient sampling rather than a significant climatic

change. In the lowermost part of the sequence (sample

intervals 19 and 20) three species of distribution group B

made their last appearances: Bembidion frontale, Bembidion

pseudocautum, and Anchicera ephippiata. Two southerly

species, Micropeplus cribratus and Micropeplus

made their last appearances in intervals 18 and 19. The

northern species Bembidion morulum appeared around 24.,000

l"\c yr B.P. in interval number 12. But because both zones

at Biggsville contain species with both northern and

southern boreal forest distributions, a relatively stable

climate is proposed for this area for the period of 28,000

to 21,000 0 I<\ C yr B. P.

The same is true of the less diverse fauna of the area

of the Athens North Quarry site. Bembidion morulum made

its first appearance in interval number 7 at about 24,000

I"\ C B P yr .. (if a constant rate of deposition of about 7 cm

per year is assumed). For the most part, however, both

northern and southern boreal elements are present in zones

A-1 and A-2, and climatic conditions would appear to have

been stable at Athens North Quarry between 25,000 and

22,000 I'\ C yr B.P.

The climatic stability indicated by the beetle assem­

blages of these two sites supports J. E. King's (1979)

r 1

t I l

106

pollen-based interpretation of a stable climate for central

Illinois. Minor changes in the composition of the local

vegetation (F. B. King, 1979; J.E. King, 1979) and the

beetle fauna may have been the result of local changes

within the plant and insect communities. The influence of

the climatic shift to colder moister glacial conditions was

perhaps not yet great enough at this time to produce a

sharp change in the faunal assemblages at the two sites.

This would imply that the transition from interglacial

conditions to the full-glacial conditions described by

Schwert et al. (1981) for east-central Illinois took place

some time after 21,000 '"c yr B.P. Cold steppe and proba­

bly tundra conditions existed at Iowa City by about 17,000

•'IC yr B. P. (Schwert et ~-, 1981).

' . '

CONCLUSIONS

Based upon the fossil beetle assemblages from two

Middle to Late Wisconsinan sediment sequences, supplemented

by sediment, gastropod, pollen, and plant macrofossil data,

the following conclusions may be made.

1. The Athens North Quarry site represents the margin

of a pond or eutrophic lake bordered by a moist

coniferous forest. The sequence of events began

with the expansion of a nearby lake or pond over

the forest floor, and ended with the infilling of

the water body by Woodfordian loess and organic

material.

2. The Biggsville sequence reveals the development

and demise of a bog in an abandoned stream chan­

nel, surrounded by at least a thin coniferous

forest. The bog and forest were eventually choked

by peat and loess deposits, and these were fol­

lowed by a period of pedogenesis.

3. The climate of central and western Illinois

between about 28,000 and 21,000 t4c yr B.P. was

relatively stable, and was similar to that of the

southern boreal forest region of southern Canada

and northern United States. The possible indica­

tions of cooler, wetter conditions beginning some

" '

r t

I

..

108

time between 23,000 and 22,000 14 c yr B.P., in­

ferred by F. B. King (1979) and J.E. King (1979)

from vegetational studies at Athens North Quarry,

were not reflected by corresponding changes in the

beetle fauna. Compositional changes in the floral

and coleopteran fossil assemblages during this per­

iod are more likely reflections of changes in the

local environment than of a regional climatic

change. The major shift from interstadial condi­

tions to full-glacial conditions occurred sometime

between the deposition of the beetle-bearing

Biggsville section (ca. 21,000 years ago) and the

development of the tundra conditions described by

Schwer·t et al. (1981) in east-central Iowa

(ca. 17,000 years ago) .

••

APPENDIX

I

• [

...

Appendix A

Sample interval depths, sample weights, and numbers of beetle individuals

and taxa per interval.

• •

•

111

Table 6. Sample interval depths and samp weights, Athens North Quarry site, Illinois.

Interval Depth Weight number (m) (kg)

1 3.4 - 3.6 3.925

2 3.6 - 3.8 10.026

3 3.8 - 4.0 9.457

4 4.0 - 4.2 10.420

5 4.2 - 4.4 10.378

6 4.4 - 4.6 8.584

7 4.6 - 4.8 9.835

8 4.8 - 5.0 11. 624

9 5.0 - 5 • 2 11.257

10 5. 2 - 5.4 10.240

11 5.4 - 5.6 7.887

12 5. 6 - 5.8 11.180

13 5.8 - 6.0 11.120

14 6.0 - 6.2 11.685

,. 112

Table 7. Sample interval <:iepths and sample weights, Biggsville site, Illinois.

Interval Depth Weight number (m) (kg)

1 4.42 - 4.52 4.52

2 4.52 - 4.62 4.08

3 4.62 - 4.72 3.60

4 4.72 - 4.82 3.25

5 4.82 - 4.92 3.43

6 4.92 - 5.02 3.40

7 5.02 - 5.12 3.72

8 5.12 - 5.22 2.73

9 5.22 - 5.32 3.24

10 5.32 - 5.42 3.17

11 5.42 - 5.52 3.36

12 5.52 - 5.62 3.34

13 5.62 5.72 3.76

14 5.72 - 5.82 4.02

15 5.82 - 5.92 3.13

16 5.92 - 6.02 4.01

17 6.02 - 6.12 3.94

18 6.12 - 6.22 4.44

19 6.22 - 6.32 5.50

20 6.32 - 6.42 4.95

21 6.42 - 6.52 5.12

22 6.52 - 6.62 5.50

113

Table 8. Numbers of identified beetles and minimum number of taxa per interval, Athens North Quarry site, Illinois.

Interval Minimum number of Minimum number of Minimum number identified beetle identified beetle number

individuals individuals per kg of taxa

1 0 0 0

2 19 { 10) * 1. 9 12

3 14 ( 43) 0.7 10

4 50 (20) 4.8 20

5 35 ( 2 9) 3.4 17

6 21 ( 2 9) 2.4 14

7 16 ( 12) 1. 6 11

8 34 ( 6) 2.9 12

9 20 ( 2 5) 1.8 12

10 17 (24) 1. 7 10

11 4 0) 0.5 4

12 18 6) 1. 6 11

13 28 0) 2.5 10

14 3 0) 0. 3 2

* Figures in parentheses are the percentages of aquatic beetle individuals.

~ 114

Table 9. Numbers of identified beetles and minimum number of taxa per interval, Biggsville site, Illinois.

Interval Minimum number of Minimum number of Minimum number identified beetle identified beetle number

individuals individuals per kg of taxa

1 9 ( 22) * 2.0 9

2 22 ( 3 6) 5.4 10

3 10 (20) 2.8 9

4 8 ( 12) 2.5 8

5 10 ( 10) 2.9 6

6 9 ( 3 3) 2.6 7

7 36 ( 11) 9.7 13

8 28 ( 21) 10.3 17

9 46 ( 4) 14.2 20

10 203 (11) 64.0 29

11 76 ( 32) 22.6 19

12 114 ( 2 7) 34.1 27

13 107 (33) 28.5 28

14 56 ( 4 3) 13.9 22

15 180 (40) 57.5 25

16 S3 ( 45) 13.2 24

17 81 (43) 20.6 21

18 42 (38) 9.5 23

19 95 ( 3 9) 17.3 34

20 96 (38) 19.4 34

21 54 ( 50) 10.5 22

22 60 ( 40) 10.9 28

* Figures in parentheses are the percentages of aquatic I beetle individuals. .

.... . T 15'1 I .....

~ I

REFERENCES

---

C r

REFERENCES

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Ashworth, A. C., 1977, A Late Wisconsinan coleopterous assemblage from southern Ontario and its environmental significance: Canadian Journal of Earth Sciences, v. 14, p. 1625-1634.

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Blatchley, W. S., and Leng, C. W., 1916, Rhyncophora or weevils of northeastern America: Indianapolis, The Nature Publishing Company, 682 p.

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Coope, G. R., 1968, Insect remains from silts below till at Garfield Heights, Ohio: Geological Society of America Bulletin, v. 79, p. 753-756.

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Coope, G. R., 1975, Climatic fluctuations in northwest Europe since the last interglacial, indicated by fossil assemblages of Coleoptera, in Wright, A. E., and Moseley, F., eds., Ice ages-;-ancient and modern: Liverpool, Seal House Press, p. 153-168.

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