A PALEOENVIRONMENTAL ANALYSIS USING FOSSIL INSECTS IN LATE QUATERNARY DEPOSITS IN INDIANA AND OHIO
Melanie L. Bergolc
A Thesis
Submitted to the Graduate College of Bowling Green State University in partial fulfillment of the requirements for the degree of
MASTER OF SCIENCE
December 2004
Committee:
Peg Yacobucci, Advisor
Daniel Pavuk
Jeffrey Snyder
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ABSTRACT
Peg Yacobucci, Advisor
Numerous biological records have been used to infer past climate and
environment. Insects are useful indicators of environmental changes because they are
highly responsive to environmental change. My study examined four sites in Ohio and
Indiana that are dated from the Middle Wisconsinan to the Late Wisconsinan and early
Holocene: Bergendorfer IN (44,000 – 21,640 yr BP), Sidney OH (40,000 yr BP), Snyder
IN (19,700 yr BP) and Sheriden Pit OH (11,557 - <9,844 yr BP). The main focus is on the insect Order Coleoptera (beetles) and how coleopterans may be used as paleoenvironmental indicators. Collection and processing of specimens followed standard techniques. Habitat types for insect species and genera were collected from recent publications. Jaccard and Dice coefficients were used to differentiate insect assemblages. The Mutual Climate Range (MCR) method was used to determine temperature ranges for the sites.
Diversity ranged from very low (Sheriden Pit) to high (Bergendorfer Low) in the sites and horizons. All sites are separate assemblages at genus and family level. Insect habitat, modern biogeography, and MCR envelopes were used for interpretation of the sites’ paleoenvironment. A moist to wetland habitat persisted in the Indiana-Ohio border region in the time intervals represented by the samples, although the temperatures indicate repeated shifts from boreal to slightly warmer conditions. Bergendorfer was most likely a moist to wet, boreal habitat. The Sidney site was near water but no general
iii biome could be determined. Bergendorfer Low may have been a boreal to boreal-tundra ecotone and a very wet environment (bogs or streams). Bergendorfer High was most likely a moist to riparian boreal habitat. Snyder was most likely a boreal forest with spruce and bogs. Sheriden Pit was moist at the time Arpedium cribratum was buried and slightly cooler than today.
This study has demonstrated the utility of insects, especially coleopterans, in determining paleoenvironments. The data correlates quite well with previous pollen and invertebrate studies done at the same sites, and general glaciation patterns during the time period. More studies of ancient insect assemblages will further determine regional habitat shifts throughout the Late Quaternary.
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ACKNOWLEDGMENTS
This project was funded by the Richard Hoare Research Fund, provided by the
Bowling Green State University’s Department of Geology. I would like to thank the following people who have helped me with my thesis. I thank Bob Hall of the Indiana
University Purdue University at Indianapolis Indiana Geology Department and Glenn
Storrs of the Cincinnati Museum Center for allowing me to have access to study sites and recovered sediments for my thesis. I thank Joe Baker, for allowing me to sieve my numerous sediments in a wet and messy fashion in the greenhouse, so I wouldn’t have to worry about being very tidy. Scott Elias sent me Mutual Climate Range files of specific beetles in my study. Donald Schwert identified my fossils when no one else was able to.
Finally, to my committee, Dan Pavuk, Jeff Snyder, and Peg Yacobucci, for reading many revisions and giving me good advice, or cheering me up when things were down.
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TABLE OF CONTENTS
Page
INTRODUCTION ...... 1
Example One...... 2
Example Two...... 3
Example Three...... 4
Quaternary Geologic History: An Overview ...... 5
RESEARCH OBJECTIVES AND HYPOTHESIS...... 13
Significance of Study...... 14
METHODS…..……………...... 16
Study Sites…...... ……… 16
Sample Processing and Specimens ...... 21
Quantitative Analysis...... 22
RESULTS…………… ...... 27
Insects Found at Sites...... 27
General Diversity...... 31
Dissimilarity Tests...... 32
MCR Analysis ...... 32
Summary of Paleoenvironment...... 33
DISCUSSION………...... 35
Changes Through Time...... 35
Specimens Found Often...... 36
Correlations between Glaciation Events and Temperature and Habitat of
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the Sites………...... 36
Correlations with Previous Studies...... 38
Sources of Error ...... 42
CONCLUSION.……...... 46
REFERENCES ...... 48
FIGURES………………...... 57
TABLES………………...... 82
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LIST OF FIGURES
Figure Page
1 Stages of Glaciation in the Tri-State Area ...... 57
2 Glacier Advance Limits During the Pleistocene and Site Localities...... 58
3 Summary of Glaciation and Floral and Faunal Changes Throughout the Late
Pleistocene and Holocene ...... 59
4 Hypothetical Temperature Ranges of Two Insect Species ...... 60
5 Sidney, Ohio Insect Fragments...... 61
6 Sidney, Ohio Insect Fragments Pictures That Were Not Identified...... 64
7 Distribution Map of Helophorus sempervarians ...... 67
8 Distribution Map of Tachinus frigidus ...... 68
9 Distribution Map of Helophorus lacustris...... 69
10 Distribution Map of Lordithon longiceps ...... 70
11 Distribution Map of Polygraphus rufipennis...... 71
12 Insect Fragments from Sheriden Pit , Horizon 50-55 cm Below Base ...... 72
13 Insect Fragments from Sheriden Pit, Horizon 0-5 cm Above Base
and Contact……...... 73
14 Insect Fragments from Sheriden Pit, Horizon 35-40 cm Above Base……….. 74
15 Insect Fragments from Sheriden Pit, Horizon 110-120 cm ...... 75
16 MCR Envelope for Bergendorfer...... 76
17 MCR Envelope for Bergendorfer Low ...... 77
18 MCR Envelope for Bergendorfer High...... 78
19 MCR Envelope for Snyder...... 79
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20 MCR Envelope for Sheriden Pit, 110-120 cm...... 80
21 TMAX and TMAX Calibrated Temperatures (Standard Error ±0.7ºC) Through
Time for the Sites Studied…………………...... 81
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LIST OF TABLES
Table Page
1 Sites and Their Associated Organic Horizons for Fossil Insects...... 82
2 Radiocarbon Dates for All Sites and Material Used...... 83
3 Insect Specimens Found in Bergendorfer...... 84
4 Insect Specimens Found in Sidney ...... 84
5 Insect Specimens Found in Bergendorfer Low...... 85
6 Insect Specimens Found in Bergendorfer High...... 86
7 Insect Specimens Found in Snyder...... 87
8 Insect Specimens Found in Sheriden Pit...... 87
9 Modern Day Biogeography and Ecology of Insect Specimens Found in
Bergendorfer…...... 88
10 Modern Day Biogeography and Ecology of Insect Specimens Found in
Bergendorfer Low...... 89
11 Modern Day Biogeography and Ecology of Insect Specimens Found in
Bergendorfer High ...... 90
12 Modern Day Biogeography and Ecology of Insect Specimens Found in
Snyder………… ...... 90
13 Diversity of Sites at Different Taxonomic Levels ...... 91
14 Dice and Jaccard Coefficients for Genera and Families of all the Sites...... 91
15 MCR Analysis for all Sites and Horizons...... 92
1
INTRODUCTION
Numerous biological records have been used to infer past climate and environment. Microfossils and macrofossils are used frequently, but depending on the type of organism, results can vary. Our detection of cooling and warming events may be skewed due to an organism’s behavior, migration capability, tolerance to temperature change, or other factors. Insects are more useful as indicators of environmental changes than larger or longer-lived organisms because they are highly responsive to environmental change (Schowalter, 2000). The populations of many species can change several orders of magnitude on a seasonal or annual time scale due to their small size, short life spans, and high reproductive rates, minimizing time lags between environmental change and population adjustment to new conditions. Insect fossils can show major climate changes occurring in as little as a few decades within the fossil record, compared to other indicators like pollen that show changes lasting hundreds to thousands of years because of migration lag (Elias, 1994). Insects are particularly vulnerable to changes in temperature, water availability, and air and water chemistry because of their relatively large surface area to volume ratios. The distribution of insects is further influenced by topographic relief, substrate structure and chemistry, and wind.
No individual species is capable of tolerating the entire range of tropical to arctic temperatures or desert to mesic moisture conditions (Schowalter, 2000). These characteristics of insects make them good indicators of climatic changes in the past and present.
The majority of fossil insect studies center on interpreting Quaternary paleoenvironments. Quaternary entomology studies began in the late 1800s and early
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1900s. Most fossil specimens were not given names of extant species because workers believed that most organisms worldwide became extinct during the Pleistocene and the species living today evolved after this extinction (Elias, 1994). Grinnell began to classify a few fossil beetles as extant ones, but it was not until the mid-1900s that the work of
Carl Lindroth and Russell Coope showed that most Quaternary beetles are actually still living today (Elias, 1994). Lindroth revised past studies of fossil insects in Sweden during the 1940’s, giving them extant names and pointing out the taxonomic errors of others. Coope collected fossil beetles in 1955 at the Upton Warren near Birmingham,
UK. He made comparisons with museum specimens and inferred that all the specimens could be identified as modern species. Coope has published over one hundred papers on fossil beetles and their paleoenvironmental significance.
In North America, Quaternary entomology lagged behind Europe, but in the late
1960s workers began to focus on using modern species to identify fossil insects.
Numerous studies have concentrated in the Great Lakes region (especially Canada),
Arctic region, and parts of the southwestern United States (Elias, 1994; Morgan and
Morgan, 1980; NDSU-QEL). The following are several examples of studies from the
United States:
Example One
Morgan et al. (1985) studied a site on the Au Sable River, Michigan, north of
Saginaw Bay near Oscoda, for its Holocene paleoenvironmental and paleoecological record. The site represents the fluvial sediments of the past activity of the Au Sable
River during the Holocene. Two logs from the site were radiocarbon dated at 4,020 ± 30 yr and 4,130 ± 90 yr BP. The beetle fragments were collected by using a 300µm mesh
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screen and washing the silty organic sediment through the screen with kerosene. Over
150 extant beetles were described. Morgan and his colleagues used modern geographical
distributions and habitat of the beetles to infer past conditions at the site. The fossil beetles found are mixed forest species (deciduous and coniferous) and species that live in oxygenated running water and still waters. This environment is similar to northern and central Michigan (of today) where a mean July temperature is 20°C, but the site could have been slightly cooler due to the high number of species found that inhabit northern boreal forest. The ecological habitat of the site around 4,000 yr BP is quite similar to the modern northern mixed forest region within the state. Morgan et al. (1985) interpreted the Au Sable fossil assemblage to represent the initiation of relative ecological stability of this region, during mid-Holocene times.
Example Two
Schwert (1992) combined new sites and previous literature to describe beetle fauna transitions through space and time (28,000-11,000 yr BP) in the mid-continent
United States (Illinois, Iowa, Minnesota, and Wisconsin) as glaciation occurred and climates changed. The beetle fauna described in this study is almost a complete chronological sequence showing how beetles respond to climate change and ice movement prior to, during, and subsequent to the advance of the late Wisconsinan ice sheet. Schwert used a classification scheme, called principle range, based on the northern and southern limits of the species, whether the species were latitudinally restricted or not, and similarity tests of assemblages using the Otsuka similarity coefficient. According to insect records, the environment changed from a closed coniferous forest, to a tundra- forest tundra, to a non-arctic open ground (few trees with insects representing southern
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latitudes below the arctic), and back to a closed coniferous forest. The beetle transitions
were not synchronous throughout the region because of the time-transgressive natures of
glaciation, deglaciation, and community response. Also, some remnants of the insect
assemblages described in the fossil record still exist in parts of the North American
tundra and arctic, to which they possibly migrated after glaciation. Schwert mentions
that future work is needed to understand how refugia and dispersal abilities affect the
response of different species of insects to glacial climates, and including the use
molecular genetics to study genetic changes within a population of a species through
time. Study sites located in the same geographic region are also needed.
Example Three
Cong et al. (1996) studied a site 43,500 (no error given)-39,000±960 yr BP (mid-
Wisconsinan substage) in Titusville, Pennsylvania. The mid-Wisconsinan is generally
known as a relatively warm interval, but climate conditions are poorly known since the
organic sediments are usually stratigraphically discontinuous and poorly dated. Also,
considerable disagreement is expressed with the interpretation of vegetational profiles
consisting mainly of Pinus pollen. Cong et al. (1996) analyzed beetle assemblages from different sediments (peat and silt) and stratigraphic zones within the narrow outcrop belt of the Titusville Till. The sediments are floodplain in origin. They also incorporated the results from a study done in 1967 on pollen and macrofossils. Cong et al.’s (1996) study was done to gain a better understanding of the mid-Wisconsinan paleoclimate in the northeastern U.S. The fossil beetles were collected using a kerosene flotation technique with the peat first soaked in 5% NaOH to disaggregate it. The entire Titusville section spans 3,000-4,000 yrs based on radiocarbon dating and may have been considerably
5 shorter based on interpretation of the sedimentary environment. The fossil sequence is subdivided into three assemblages based on cluster analysis of Otsuka similarity coefficients and shows a cooling, warming, and a cooling again. The beetle assemblages went from a forest-tundra to a boreal-hardwood forest and back to forest-tundra. This study is consistent with studies of beetles in the British Isles and oxygen isotope records from the Greenland Ice Sheet (Cong et al., 1996). Cong et al. (1996) concluded that a significant climate change occurred in which temperatures rose 5-7°C.
This study will incorporate methods and techniques that these researchers and others have used. The current study will use modern day habitats of insects to infer the past. It will also look at several sites but in a more confined geographical area and a wider time range. The beetle assemblages will be evaluated for changes in insect communities and environments through time (Schwert, 1992; Cong et al, 1996), and the
Mutual Climate Range (MCR) method will be used to calculate an actual temperature range.
Quaternary Geologic History: An Overview
Glaciation and Climate
The Quaternary Period, including the Pleistocene Epoch (1.65-0.01Ma) and
Holocene Epoch (0.01Ma to Present), is characterized by repeating cycles of glaciation and interglacial warm periods. During the Pleistocene, the main areas of glaciation were
North America, Greenland, Eurasia, and Antarctica. In the Holocene, climate changes were smaller in amplitude than those of the Pleistocene (Easterbrook, 1999). Modest expansion of glaciers occurred during cooling events. A short cooling event, known as the Younger Dryas, occurred 12,900 to 11,600 cal yr BP (or Late Pleistocene to Early
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Holocene transition, usually radiocarbon dated at 9,900 yr BP; Renssen et al., 2000).
Temperatures were higher on average then they are now throughout most of the globe
during the early and late Holocene (Holocene Optimum and Medieval Warm period;
Lauritzen, 1996; Linacre, 1997; Darby et al., 2001; Siegert, 2001) with another short
cooling between these two warming events. A general cooling event occurred in the Late
Holocene, starting in the thirteenth century A.D. and ending between 1850 A.D. to 1890
A.D., in which glaciers extended their positions globally (Grove, 2001). In North
America, glacier advances occurred in the Canadian Rocky Mountains, east of Calgary,
Alberta (Grove, 2001). The nature and distribution of these Holocene events is still extensively studied.
The Quaternary history of the tri-state area (Indiana-Michigan-Ohio) includes numerous advances and retreats of ice over the area (Figure 1 and 2; Eschman, 1985).
The Late Pleistocene, especially the interval from 24,000 to 10,000 yr BP, experienced multiple cooling and warming events (Figure 2b). Ice would sometimes cover only parts of upper Michigan but reached as far as southern Ohio and central Indiana. During the early Holocene the ice re-advanced several kilometers past the present geographic position of the lakeshore of Superior in the northern portion of the upper Michigan peninsula near Marquette (Eschman, 1985).
Vegetation Changes
There were multiple changes within North America’s vegetation biomes during the Late Quaternary in response to glacial cycles (Adams and Faure, 1997; Adams, 1997
(unpublished)). The following summary will focus on the Great Lakes region, specifically Michigan, Indiana, and Ohio 40,000 yr BP to present. The basic biomes
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occupying the Great Lakes region and climate changes will be discussed here which are described by Adams (1997) and Adams and Faure (1997). The Great Lakes region was most likely spruce and jackpine forest with a cold temperate mixed forest in Tennessee and North Carolina at 40,000 yr BP (Adams 1997). The same boreal forest type was present around 28,000-25,000 yr BP in a large portion of the region, along with tundra and ice in the northern part (Adams, 1997). Around 18,000 yr BP, the region became colder and drier than 28,000-25,000 yr BP, and was mostly covered by ice with open boreal and ‘mid taiga-like’ vegetation (conifer to broadleaved forest with open canopy;
Adams and Faure, 1997). Around 13,000 yr BP a retreat of glaciers allowed more land to
be colonized. During this time, the Great Lakes region seems to be most diverse, having
ice, tundra, open boreal woodland, and mid taiga (Adams, 1997). 11,000 yr BP brought
differential climate patterns throughout North America with certain regions cooling and
others warming, and a general spread of deciduous forest species. In our region (tri-state
area), an open boreal forest, mid taiga, and a cool temperate mixed forest (conifers and
deciduous trees) occurred (Adams, 1997). A warm to slightly warmer climate occurred
in the region around 8,000 yr BP. A cool mixed temperate forest was the dominant
biome with prairie approaching the region on the western side (Adams and Faure, 1997).
This prairie intruded the Great Lakes region 5,000 yr BP (northwest Ohio was prairie and
currently has remnant patches today) when the Midwest experienced a relatively dry
phase (Adams, 1997). Around 4,000 yr BP the region became moist and cooler (Adams,
1997). Presently, we have a moist climate with a cool temperate mixed forest that would
be widespread in the region if no agriculture was present (Adams and Faure, 1997).
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Herpetofaunal Changes
Holman (1992) studied the herpetofauna of the Great Lakes region (Indiana,
Illinois, Michigan, and Ohio) during the late Pleistocene and Holocene. Based on the animals found and the site location, warming and cooling events can be deduced. A warming period occurred 15-14,000 yr BP in southwestern Indiana based on a rich herpetofauna found at the Prairie Creek site south of Indianapolis. The majority of these species are very common in the southeastern and south-central United States. A re- invasion of turtle species occurred 13,000-10,000 yr BP in central Indiana and central
Ohio. In Michigan there was still a pro-glacial climate during this time period. By
11,000 yr BP, modern northwestern Ohio reptiles and amphibians were mainly established (Holman, 1995). Also around 11,000 to 10,000 yr BP, a cold adapted turtle fauna (Chelydra serpentine and Chrysemys picta) was found in Clark Kettle Bog, Darke
County, Ohio (Holman, 1995). In the Holocene, the modern herpetofauna was established throughout the Great Lakes region 6-4,000 yr BP based on a southwestern
Indiana site and three sites in Michigan (Holman, 1992).
Megafauna and Mammals
The main mammalian fauna of North America during the late Pleistocene was part of the Rancholabrean Land Mammal Age (300,000 and 150,000 to 10,000 yr BP), comprising bison, woolly mammoth, caribou, mountain goats, sheep, moose, musk oxen, dire wolves, and humans (Lange, 2002). In the tri-state area and adjacent areas, a small number of studies have been conducted. In Big Bone Lick, Kentucky, 32 km southwest of Cincinnati, Ohio, a mammalian fauna was described in a sequence of three layers aged
18,000 yr BP to near present (Lange, 2002). The late Pleistocene layer (layer three) is
9 comprised of typical megafauna bones: musk ox, stag moose, ground sloth, tapir, mastodon, mammoths, deer, caribou, and more. Layer two has deer, elk, bison, and the older animals. Layer three has both a modern, recent fauna and human artifacts like pottery.
Sheriden Pit, Ohio, east of Findlay, has a rich vertebrate fauna with almost thirty mammal species found dating to between 11,700 to 14,000 yr BP. Giant beaver, short faced bear, flat headed peccary, and elk moose are among the extinct species found at the
Sheriden Pit site (Hansen, 1992). Extant animals recovered that do not live in Ohio now are caribou, porcupine, yellow-cheeked vole, and heather vole (Tankersley and Redmond,
2000). These species were well south of their current range, indicating cooler temperatures. Numerous species currently living in Ohio are also found. Figure 3 summarizes all the organism and glacial activity during the end of the Pleistocene and throughout the Holocene.
Community Ecology Dynamics during the Quaternary
Plant and animal communities may show multiple responses due to climate changes during the Late Quaternary: remaining stable, shifting geographic range, changing internal composition, or creating a refugium. The majority of community dynamics studies have focused on plants, but a few mammal studies have been done.
The two mammal studies done show similar reactions of mammal community responses due to climate changes. In Spain, a study by Rodriguez (2004) showed that the ecological structure of a mammal community was stable throughout climate change episodes during the late Early Pleistocene to the Late Middle Pleistocene, but that the species composition changed and each species behaved individually (Gleasonian
10 community). The FAUNMAP Working Group (1996) looked at species composition of mammal communities and came to the same conclusion that mammals migrate individually in the United States but are that the community is still organized into similar biogeographic patterns (faunal provinces). They also concluded that mammal communities are unpredictable, late Pleistocene communities have no modern analogs, and that modern community assemblages were established only in the last few thousand years.
Plant communities also show a reorganization of species in some studies
(Edwards et al., 2000; Elenga et al., 2000; Jackson and Overpeck, 2000; Prentice et al.,
2000; Williams et al., 2000; Williams et al., 2001; Williams et al., 2004), but also have shown the opposite in a few studies; plant communities can come back again with the same species composition. Few papers have reported biomes and plant communities that retain the same composition or same ecological structure over time. Williams et al.
(2000) only found one biome, cool conifer forests, that stayed structurally the same from the last glacial maximum to present. Tarasov et al. (2000) also indicated only a few biomes and plant communities staying ecologically unchanged (e.g., steppe-like associations in Siberia); structurally similar biomes during the last glacial maximum are still present today in Georgia, Kyrghyzstan, and Mongolia. Tarasov et al. (2000) found that the cryoxerophilic vegetation community has no modern analogs today.
Some studies have shown that even during glacial and interglacial activities, a plant community will stay stable compositionally throughout the whole time but will only change in size, expanding and contracting (Tzedakis et al., 2002; Falcon-Lang, 2004).
However, only one study shows multiple modern analogs for Quaternary plant
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communities. Delcourt and Delcourt (1987) show that full glacial, warm temperate,
mixed conifer-northern hardwood, and boreal forests that occurred in the late Pleistocene
throughout the Holocene in eastern North America have modern analogs. They did show
that during a few times throughout this time period, modern analogs do not exist and that
cool temperate deciduous forests were first created in the Holocene.
Finally, a refugium is sometimes created. A refugium occurs when environmental
changes are occurring throughout an area, and a community or species persists in a small
location whereas the community changes in the rest of the area (e.g., through migration and local extinctions; Ricklefs, 1993). After the environment becomes stable again, migration can occur from refugia and a species can recolonize an area to a large extent
(Gathorne-Hardy et al., 2002). However, if the ecosystem does not return to its original climate, relict populations may persist even today. Tundra species may be found in grasslands, or within temperate forests. Refugia may be able to be distinguished from
biomes migrating by looking at multiple taxa (plants and animals) and sites within a
geographical area. Synchronous and near synchronous cooling or warming at multiple
sites within a close proximity, or large geographic area can permit one to determine
whether an area is a refugium or intact biome.
It is still unclear how terrestrial communities responded to climate fluctuations
during the Late Quaternary glacial intervals. The majority of the studies focus only on
the Last Glacial Maximum (LGM) to present, and the use of plant species. Further
studies still need to be done in this area to better understand community changes due to
climate fluctuations: expand the time interval to include other glacial maximum events,
use other taxa (vertebrates, invertebrates, and microorganisms), and including other
12 factors besides temperature changes. Temperature alone will not determine whether the same community will come back again. Other physical factors should be studied such as precipitation, soil, and topography.
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RESEARCH OBJECTIVES AND HYPOTHESIS
Many parts of North America (especially central Canada and the southeastern
United States) still lack studies describing insect species within Quaternary deposits or do
not apply the Mutual Climatic Range (MCR) method (described further below). In Ohio,
several studies have been done, but they are scattered and are mainly in far northern and
southern Ohio (Coope, 1968; Morgan, 1987; Shane, 1987; Lowell et al., 1990).
Geographical gaps exist within many parts of the tri-state area (Ohio-Michigan-Indiana).
My study will examine four sites in Ohio and Indiana that are dated from the
Middle Wisconsinan to the Late Wisconsinan and early Holocene. Two of these sites have previous data on insects found in them while the others lack insect fossil studies.
The primary analysis focuses on the insect order Coleoptera (beetles), specifically to demonstrate how coleopterans may be used as paleoclimate indicators (other insects will also be considered). Another objective is to determine whether the same assemblage reoccurs through time within a small geographic location. This second analysis will determine if the same assemblage of species returns after climate change or if a community remains compositionally stable through time during relatively stable climate.
My hypothesis is that the species composition of insects stays the same within the same biome and climate at a particular site (if it persists or reoccurs). The null hypothesis is that the species composition of insects changes whether or not the same biome and climate persist through time.
This study will involve identification of the insect species, description of community diversity, and interpretation of habitats and environments. The mutual climate range (MCR) method, which has been used mostly in Europe, will be used to
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reconstruct the past climate. To avoid circularity, the paleoclimate (temperature and seasonality) will be determined using only scavenger and predatory beetles while the fossil assemblages will be defined using all beetle species and other orders of insects.
Significance of Study
This study may help us understand changes within a community of organisms due
to climate and environmental changes. Questions such as whether the same assemblages
of species reoccur throughout time if the appropriate environment reoccurs can be
answered. We can also determine if organisms can migrate back and forth to a specific
area, or if they completely disappear from an area even if other evidence shows that an
area may become suitable for an organism. This information is important to know since
presently global temperatures are rising due to an increase in atmospheric CO2. Humans have caused (and are causing) loss of habitat, changes in water and soil, and air quality.
Experiments have shown that with induced warming of an area, microclimate changes occur within these sites (Shaw and Harte, 2000; Loik et al., 2001, and Scheil et al., 2004).
Long term effects are more recently being realized.
Several possible climate changes may occur within northwest Ohio, eastern
Indiana and central-southern Michigan. Presently this region is a deciduous forest biome
(more precisely, a beech-maple vegetation type) with an annual precipitation in this biome of 80-150 cm and a seasonal temperature span of 38°C to -30°C (Barbour et al.,
1999). If temperatures rise within the local region, a deciduous forest would still be in this region but the more precise vegetation type would shift. If the temperature remains in a similar range to that of present times (45°C to -40°C) but the climate becomes drier
(30-80 cm per year), a grassland will develop (Barbour et al., 1999). If the local climate
15 should become cool and drier (certain places might do this even though the globe as a whole would rise in temperature), a boreal biome may form. A boreal biome is characterized by a temperature range of a summer maximum in the low twenties to -50°C
(for minimum winters) with precipitation of 30-85 cm per year (Barbour et al., 1999). If cooler temperatures do not reach this far south, then a boreal-deciduous mixed forest or ecotone may develop.
A better understanding of past climate and how organisms interact with the environment will help us deal with the environmental changes that occur today. We may learn if certain species can easily travel out of an area and then later re-colonize it due to habitat changes that occur on shorter time scales (decades-centuries). However, now we have to take into consideration human-made obstacles affecting current dispersal and possible future dispersal of organisms.
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METHODS
Study Sites
The sites chosen for this study are located in east central Indiana, west central
Ohio, and northwestern Ohio, and range from the Middle Wisconsinan to Early Holocene
(Tables 1, 2, and Figure 2). These sites represent similar depositional settings and sediments (paleosols near rivers with high silt content) except for Sheriden Pit (a terrestrial pit into which organisms fell). Site locations and sediment samples were provided by Professor Robert Hall from Indiana University-Purdue University
(Indianapolis, IN) and the Cincinnati Museum Center. Professor Hall showed me the
Bergendorfer, Sidney, and Snyder sites. I used previously collected data from one of his student’s theses (Knollenberg, 1996) and sampled Sidney for my own analysis. The
Cincinnati Museum Center loaned me sediments from Sheriden Pit to process and analyze the insect fragments. The following is a general geological history and description of organic layers at the fossil sites:
Bergendorfer
The Bergendorfer section is exposed along the streambanks of Williams Creek.
The age of this section spans from 46,700 ±2100 yr BP to 21,190 ±360 yr BP (Hall and
Zbieszkowski, 2000). Bergendorfer’s base is possibly Illinoian diamicton overlain by alluvium and colluvium that may be Illinoian to Wisconsinan (Hall and Zbieszkowski,
2000). In the first sequence (BD-2) sampled by Knollenberg (1996), the soil (with thick organic layer; 44,000 yr BP) starts off at the base of the river and is 1.5 m thick. This paleosol is overlain by about 8 m of diamicton with a thin light gray clay layer separating the two. The second sequence (BD-4) sampled by Knollenberg (1996) includes the
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younger organic layer (21,190 yr BP). The layers are more diverse, starting off with 5m
of diamicton at the base of the river. The diamicton is overlain by a thin gravelly sand
layer that fines upwards, followed by the organic layer (1.5 m thick). The organic zone is then overlain by 5 m of till, followed by 1.5 m of loess.
The two organic zones are dated to 44,000 ±2000 yr BP and 21,190 ±360 yr BP
(Knollenberg, 1996; Hall, pers. comm. October 17-18 2002) and named in this paper as
Bergendorfer and Bergendorfer Low/Bergendorfer High, respectively. The Bergendorfer organic zone is a 7 cm thick silty loam (70% silt) and has a black color, common organic matter, and a massive (structureless, hard, and appearing or occurring in very large clods) soil structure (Knollenberg, 1996). Bergendorfer Low and Bergendorfer High are so named because the organic layer is very thick (151 cm) and was split up into two sampling sections by Knollenberg (1996) for insect analysis. The organic zone is dark grey, with sparse to common organic matter, a massive soil structure, and a silyt texture
(85% silt; Knollenberg, 1996). General pollen analysis shows the presence of
Pediastrum (green algae), Cyperaceae (grasslike, herbaceous plants), and grasses and ferns.
Sidney
A truncated soil is exposed at Sidney, Ohio along Brushy Creek, 7.5 km southeast from Bergendorfer. The general profile of this site is a base layer of bluish-grey clay overlain by the organic layer (Middle Wisconsinan), which is itself overlain by a Late
Wisconsinan diamicton. The soil is truncated (upper horizon(s) removed by erosion) and possibly tilted and/or folded by glaciation (Hall, pers. comm. October 17-18, 2002). The general age of the organic layer is estimated to be 40,000 yr BP since the dating of
18
material within and on top of the organic layer is scattered: 22,430 yr BP to more than
50,000 yr BP (Forsyth, 1965; Knollenberg, 1996; Hall, pers. comm. October 17-18,
2002). The organic layer is dark grey and highly organic-rich with woody debris
protruding from it. It has a massive soil structure and loam texture (25.5% silt), with
54% Pinus banksiana/P. resinosa (pines), 25% Cyperaceae with Picea (spruce), and
<5% of Gramineae (grass) (Knollenberg, 1996).
Snyder
This section is exposed near Connersville, Indiana within Fall Creek near Snyder
Road and Snyder’s farmhouse. Lowell and Brockman (1994) describe the general sequence of deposits in which the well preserved paleosol is located. A Wisconsinan diamicton is below the paleosol. On top of the paleosol lies a general sequence of Late
Wisconsinan organic silt followed by Late Wisconsinan outwash, till, and lacustrine sediments. The organic silt has some evidence of a weakly developed soil. It is 0-60 cm thick and is characterized by a dark grey color, massive soil structure, common to abundant organic material, and a silty texture (92% silt; Knollenberg, 1996). The organic material also has a banded appearance. It probably originated as loess, but was reworked in shallow ponds (Lowell and Brockman, 1994). The weak soil developed before the late
Wisconsinan glacial advance (Lowell and Brockman, 1994). Abundant plant, wood, and charcoal fragments are found in the sediment; 44% Picea, 25% Cyperaceae, and 14%
Pinus banksiana/P. resinosa (Knollenberg, 1996). Mollusks are also found in this section (freshwater gastropods, bivalves) (Hall and Zbieszkowski, 2000). The Snyder section ranges from 44,440 to 18,160 yr BP with the organic silt dating to 19,700±600 yr
BP (Hall and Zbieszkowski, 2000; Knollenberg, 1996).
19
Sheriden Pit
Sheriden Pit is located in Wyandot County, about 10 miles east of Findlay, Ohio,
adjacent to Indian Trail Caverns, a tourist attraction. In 1989, a depression was
excavated by Dick Hendricks (previous owner) in hopes of finding another below ground
link to the cave system. Instead, he found vertebrate bones from the Pleistocene (Hansen,
1992; Tankersley and Redmond, 2000). In the next several years, numerous bones of
extinct and extant animals were found. Giant beaver, short faced bear, flat headed
peccary, and elk moose are among the extinct species found at the Sheriden Pit site
(Hansen, 1992). Extant animals recovered that do not live in Ohio now are caribou,
porcupine, yellow-cheeked vole, and heather vole (Tankersley and Redmond, 2000). The
sinkhole even contained standing water long enough to support fish, frogs, water snakes,
and aquatic turtles.
In the mid 1990s, an excavation was started to find Paleoindian artifacts. Tools
like bone spear points, a bone rod, flute points, and a uniface side scraper have been found (Sheriden Cave Site, 2001). The site has been interpreted as a hunting and butchering ground since animals fell into the pit and became trapped (Sheriden Cave Site,
2001).
The sinkhole is about 50 ft deep (15.6 m) and 65 ft across (20.3 m). Twenty-nine radiocarbon dates have been taken, ranging from 12,840±100 yr BP at the bottom of the pit to 9,170±60 yr BP at the 0-5 cm a.b (above base) and base layer (Sheriden Cave Site,
2001). In general, the sediments are mostly silt and clay size with low organics, with the exception of the base layer (0-5 cm a.b.) which has a higher amount of organics. The
20
following is a more in depth discussion of each horizon and its contents. The “base”
represents the top of the artifact-bearing layer and the end of the radiocarbon dating
sequence. The following are physical sediment descriptions for each horizon in this
study:
110-120 cm horizon: This sample was incompletely labeled by collectors and it is
unknown if this layer is above or below the base. This layer has silt and clay with a high
amount of gravel. The sediment has a clumpy characteristic to it, and is brown in color.
The organics present include charcoal, microbone fragments and a few insect fragments.
50-55 cm b.b. (below base) horizon: This horizon is also gravelly with silt and clay sized
particles, clumpy, and brown. The organics include plant matter, bones, and a very small
number of insect fragments.
0-5 cm a.b. horizon: This horizon has laminated and unlaminated sections that are
orange-brownish, very silty, and with some gravel. A large amount of organics were
found in this horizon compared to the others. The organics include plant material, charcoal, bone fragments, and several insect fragments (the best preserved fragments compared to the other horizons).
15-20 cm a.b. horizon: This horizon is brown, laminated silt and clay with some sand. A large amount of plant material and charcoal are present, but no identifiable insect fragments. A few fragments may possibly be insect remains but they are too fragmented to be certain.
25-30 cm a.b. horizon: This horizon’s sediment was actually still moist after 8+ yrs in storage so it is assumed little drying took place. The sediment is earthy brown, quite
21
silty, and is laminated with fine layers intermixed with coarser layers. Organics are
mostly charcoal and plant fragments. No identifiable insect fragments were observed.
35-40 cm a.b. horizon: This horizon is silty with clay and sands in lower amounts, and is
also laminated, with an orange-brown color. The sediment was also found to be still
moist. The organics found were plant fragments, charcoal, and a few insect fragments.
Sample Processing and Specimens
Collection of specimens followed standard techniques (Elias, 1994, pp. 25-30).
Samples were taken at 5 cm intervals if the organic layer was thick (> 10cm) or in bulk if the organic layer was thin (<10 cm). The sediment for Sheriden Pit was collected in 1994 by the staff of the Cincinnati Museum and lent to the author for this study. The
Cincinnati Museum sampled Sheriden Pit in 5 cm increments, filling 5 gallon buckets two-thirds to three-fourths full for each horizon. The Sidney, Ohio site was sampled by the author in bulk (two one gallon bags or 3.71 liters) since the organic layer is approximately 10 cm thick at the site. The Bergendorfer and Snyder, Indiana sites samples were taken by Knollenberg (1996) in either bulk or smaller increments depending on layer thickness, and insects were identified by Scott Elias.
Insect specimens for Sheriden Pit and Sidney were processed by the author.
Preparation of specimens followed the guidelines found in Elias (1994, pp. 30-35). Wet sieving was done with 200 and 500 µm sieves. About 15 eight ounce (236 ml) cups of sediment (a total of ≈ 4.6 kg) were taken from each layer to be sieved so a similar sample size was processed for each layer. The method of organic flotation was different than the more common approach. Kerosene is commonly used to float organics by making the organics lighter when the kerosene attaches itself to the organics. Here, 95% ethanol was
22
used to float the organics instead of kerosene, because kerosene is extremely flammable
and potentially hazardous. This new method worked well for collecting small to large
insect fragments along with plant material, and most likely would not contribute to a lack
of insect fragments recovered. Small glass containers filled with 70% ethanol were used to store the insect specimens.
Specimens from Sheriden Pit and Sidney were identified by Donald Schwert of the North Dakota State University Quaternary Entomology Lab. He used morphologic knowledge of various insects, especially the Coleoptera, plus comparison with pinned specimens in a museum reference collection. The specimens were identified down to the species level if possible.
After specimen identification, habitat type and climate tolerances for insect species and genera were collected from recent publications (White, 1983; Downie and
Arnett, 1996; Arnett et al., 2001; Arnett and Thomas, 2001), and online databases
(National Park Service: Northern Cascades; Haarstad, 2002)
Quantitative Analysis
Species Diversity and Faunal Assemblages
Generic and family richness and biodiversity tests were applied to each layer and
each site. Two dissimilarity measures, Jaccard and Dice, were used to see if specimens
from different layers represent the same or different faunal assemblages. Both were used
to look at possible differences between coefficients to get a well rounded statistical view
of the fossil assemblage.
Jaccard Coefficient: J = a/(a+b+c)
Dice Coefficient: D = 2a/(2a+b+c)
23
a = number of species present in both units
b = number of species found only in unit one
c = number of species found only in unit two
J,D = 1 most similar, J,D= 0 least similar
If two sites have a zero value, then they are completely different assemblages. If they have a value of one, then they are the same assemblage. Since a site will show some variation of species seasonally and year by year, a value of one would be nearly impossible. A value of 0.8 was used to compensate for this variation (80% of the species are the same). If a coefficient is under 0.8, then the two sites are separate assemblages.
If the coefficient is 0.8 or higher, then the two sites are the same assemblage.
Paleoclimate Reconstruction
Mutual Climate Range is a technique for reconstructing ancient temperatures based on the geographic ranges of modern species. MCR uses scavenger and predatory beetles to establish a climate range (or envelope) to estimate the past climate by overlapping the envelopes of specific beetles found within the fossil record at a specific site and time period (Figure 4; Elias 1994 and 1997). MCR reflects the summer warmth and degree of seasonality (temperature regime) of the beetles. MCR was performed on each site.
Elias (1994) describes the procedure for MCR analysis. First, modern maps of the scavenger and predatory beetle species found at the site are compiled. Scavenger and predatory beetles are used because they are usually the first to move when climate changes occur. Some feed on carrion or organisms found on carrion. Beetles will also live in mammal and bird nests, feeding on fleas, mites, and other vertebrate parasites.
24
Others feed on arthropods or other invertebrates. Herbivorous beetles respond more
slowly because the plant species (or host plants) take longer to migrate.
Second, the modern climate range for each species is determined. Since over 250
climate envelopes have been developed for eastern and central North American insects, these were used in this study instead of creating new envelopes from scratch (Elias, pers. comm., July 22, 2002). To construct a geographic distribution, a map of the species range and a map of meteorological stations are overlain. The stations that overlap within the species range are used to determine the mean summer temperature in July (TMAX), the mean winter temperature in January (TMIN), and the annual temperature range
(TRange). TMAX describes the warmest month in summer (taking the mean historical temperature) and TMIN incorporates the coldest month in winter for an insect species.
After TMAX and TMIN are calculated for all geographical locations, a species’ TRange can be determined. TRange gives the overall temperature range a species can tolerate,
which is also known as the index of degree of seasonality.
Third, each species’ climate range is numerically stored in a database for easy
retrieval. A 36x60 grid is developed in units of 1°C with the x-axis as TRange and the y-
axis as TMAX. Each x-y coordinate in the grid is coded as a 1 or 0 for presence/absence
of the species tolerance in degrees Celsius. A digitizing tablet (Elias, pers. comm., July
22, 2002) is then used to draw the climate envelopes of each species as image files
(Figure 4).
Fourth, a mutual climate range for all of the known fossil species at a site is
reconstructed by combining all the species grids from step three. A drawing program
such as Corel Draw 7 can be used for this purpose (Elias, pers. comm., July 22, 2002). A
25 potential climate range for the fossil assemblage as a whole is found (Figure 4). The maximum temperature and temperature range of the site can be deduced from this information. Also, if assemblages are studied throughout a stratigraphic sequence, a change of temperature may be seen throughout the fossil record by using this method.
This fourth step will be used in this study by taking previously constructed climate envelopes of insect species (if they are found within the sites) in the form of a Corel
Draw graph from Scott Elias (2000). These envelopes are also available on the web through NOAA: Paleoclimatology, Insecta (2003).
This mutual climate range method tends to overestimate TMAX by less than
1.0°C and underestimate TMIN by <1.0°C or up to ±10°C, depending on the geographic location. The possible reason for TMIN having a larger error from the actual temperature is greater seasonality in North America (Elias, 1997). In Europe, TMIN has a smaller error (1-2°C off from actual temperature) because of the mild winters and maritime climate. Regression equations are used to correct this problem. Modern temperatures at various locations are run against the median reconstructed temperature that is based on the fossil fauna. Coope (Elias, 1997) developed an equation to correct for this error for
European studies:
Median TMAX (calibrated) = [med. predicted TMAX*1.2635] - 4.9059
Median TMIN (calibrated) = [med. predicted TMIN*1.3897] + 2.7012
In these equations r²=0.94, TMAX standard error is ±0.83°C, and TMIN standard error is
±2.42°C. Elias (1997) has recently applied the MCR method to North American sites and has developed these equations,
Median TMAX (calibrated) = [med. predicted TMAX*0.787] + 3.4338
26
Median TMIN (calibrated) = [med. predicted TMIN*0.716] - 4.9381 with r²=0.94 and standard error of ±0.7°C for TMAX and r²=0.87 and a standard error of
±10°C for TMIN. The standard error for TMIN may be quite high due to greater seasonality (more severe winters) compared to Europe.
27
RESULTS
Insects Found at Sites
All specimens were identified to family level (Tables 3-8). Most were identified to genus and a small number to species level. The following is a discussion of the species found, and habitat and ecological characteristics of the species and genera for each site and layer, presented in chronological order from oldest to youngest layers. Information on insect habitats is from Arnett and Thomas (2001) and Arnett et al. (2001) unless otherwise noted (Tables 9-12).
Bergendorfer
Insects were identified by Scott Elias for Knollenberg (1996) at all Bergendorfer sites (Table 3). Coleoptera: Carabidae (ground beetles) and Staphylinidae (rove beetles), and Trichoptera (caddisflies): Limnephilidae were found. The species in this layer tend to be widespread geographically with two contained in North America only (Table 9).
Only one specimen was identified to species level, Acidota crenata. A. crenata may be a predator of small invertebrates (National Parks Service) and is neoarctic in distribution.
Little information can be found on this species. Dyschirius sp. burrows in sandy and clay soils. Olophrum sp. has a boreal to alpine habitat and lives in wet areas. Its habitats include litter, moss, and rodent nests and middens. Overall, this site was cooler than today with a boreal biome and was near water or moist conditions based on the insect genera and species found.
Sidney
Schwert of North Dakota State University identified insects for the author at
Sidney (Table 4). Coleopterans were found, including the families Carabidae,
28
Chrysomelidae (leaf beetles), Curculionidae (weevils), Dystiscidae (predaceous diving beetles), Scarabaeidae (Lamellicorn beetles) and Staphylinidae. Only one specimen,
Bembidion sp., was identified to genus level at this site. This genus contains over 250 extant species. The majority of the species are hygrophilous, but some can be found in crop fields and alkaline areas. They have a very wide distribution, ranging from the
Arctic coast tundra to the subtropical lands of Florida. Specimens were also identified to several families and include leaf beetles (specifically Donaciinae which live on exposed aquatic plants (Arnett, 1985)), predaceous diving beetles, weevils, and rove beetles. This site was near standing water or running water deep enough to allow aquatic plants to grow for the subfamily Donaciinae to be found. No general biome can be deduced from these data. Photographs of the insect specimens found at Sidney can be found in Figures
5 and 6.
Bergendorfer Low
Three orders, Coleoptera, Trichoptera, and Hymenoptera (wasps, ants, bees), and twelve families were identified in this layer, with half of the specimens being
Hydrophilidae (water scavenger beetles), Staphylinidae, and Scarabaeidae (Table 5).
Five species were identified: Bembidion nigripes, Helophorus sempervarians, Acidota subcarinata, Tachinus frigidus, and Aegialia lacustris. Smetana (1988) describes
Helophorus sempervarians. Little is known about the habitat of this species, but in the few places from which data are available, the species appears to be semi-aquatic. It has been found along muddy edges of ponds, in thick vegetation in shallow water, moist mountain habitats, and in muddy flats in a marsh area near a slow flowing stream. The distribution of this species is widespread in Canada and Alaska with isolated populations
29
in mountains of the continental U.S.A. (Pennsylvania, Michigan, South Dakota). A
modern distribution map of this species is shown in Figure 7.
Campbell (1973) describes Tachinus frigidus. Most specimens have been found under animal dung but a few were found in the mouth of mammal burrows and on rotting mushrooms. The species’ distribution in mainly in Canada and Alaska, with relict populations found in the high mountains of New Hampshire and on Mt. Katahdin, Maine.
A modern distribution map is shown in Figure 8.
Most of the other specimens found in Bergendorfer Low are geographically widespread with several restricted to North America (Halilus sp., Eucnecosum sp.,
Olophrum sp., Serica sp., and Graphops sp.) (Table 10). Several species live in boreal and wetland areas (bogs, ponds, riparian areas) and one in tundra (Eucnecosum sp.).
They are also quite diverse in habitat (litter, moss, rodent nests and middens) and feeding preferences (Table 10). Eucnecosum sp. and Geodromicus sp. are predatory. Aegialia
lacustris is a detrivore, eating decaying organic matter (National Park Service). In
summary, the habitats of these species are diverse, including forest and tundra litter,
moss, and living in rodent nests, and the general biome was colder than today, most likely
a tundra-boreal ecotone in a very wet environment with bogs and possibly streams.
Bergendorfer High
Five families of Coleoptera are found in this horizon: Carabidae, Hydrophilidae,
Staphylinidae, Scarabaeidae, and Elateridae (click beetles) (Table 6). Three specimens
were identified to species level: Bembidion nigripes, Helophorus lacustris, and
Helophorus sempervarians (see above for a more detailed discussion on H.
sempervarians). Most species are distributed widely except for Hydrobius sp., which is
30
found only in North America (Table 11). Geodromicus sp. is found in moss and wet litter
in riparian zones and is predaceous. Agonum sp. is hygrophilous and found in alpine forests. Little is known about the habitat of Helophorus lacustris. Some specimens have
been collected in shallow muddy pools, vegetation along slow moving creeks and rivers,
wet swampy meadows, and debris from flooded meadows (Smetana, 1988). In Ohio, it
has been collected at two localities in small temporary ponds (Chapman, 1998). This
species has a northern transcontinental range from Alaska to Newfoundland and in the
central to northern United States (Figure 9) (Smetana, 1988). The overall habitat of this
site is a forest similar to an alpine type (analogous to boreal) and is moist to very wet.
Snyder
Three orders were found in Snyder: Coleoptera, Homoptera (aphids, whiteflies,
cicadas, and hoppers), and Trichoptera (Table 7). A large number of families were also
found in this layer (ten), Carabidae and Staphylinidae being the more frequent families
and predaceous, scavenger, sap feeding, and bark beetles occurring less frequently. Five
species were identified from the specimens found at this site by Scott Elias for
Knollenberg (1996): Helophorus sempervarians (see above for more detailed discussion),
Acidota subcarinata, Tachyporus rulomus, Lordithon c.f. L. longiceps, and Polygraphus
rufipennis. Campbell (1982) discusses Lordithon c.f. L. longiceps. This is a rare species
and very little is known about it except that adults tend to be found on rotten gilled
mushrooms. The distribution is very scattered: north-eastern US, northern British
Columbia, Quebec, and central Alaska (Figure 10).
Bright (1976) describes Polygraphus rufipennis. This species is a bark beetle
(Scolytidae). Adult beetles live on conifers, specifically dead or dying spruce. The
31
distribution in North America is trans-continental in Canada and stretches to the north-
eastern U.S. (Figure 11). In the northeast region, the species can be found south to North
Carolina and Tennessee. In the north-western region, the range stretches from Oregon to
New Mexico (note that Figure 11 only shows Canada and Alaska).
All species are widespread except for Acidota subcarinata, which is found in
eastern North America (Table 12). This species also lives in forests and bogs under litter
and moss. Tachyporus rulomus is found under bark (Haarstad, 2002). Forest habitat
characterizes this site with spruce as one of the tree species and with one or more bogs
present.
Sheriden Pit
One order and one family was found at Sheriden Pit: Coleoptera, Staphylinidae
(Table 8). Only one species was identified by Donald Schwert at this site, Arpedium
cribratum. Its modern distribution is from Iowa to Ohio. The genus as a whole (only three species) is usually found in flood debris, litter, or moss near water, and occasionally in vole nests or beaver houses. No biome can be deduced from this data but the habitat was moist at the time of Arpedium cribratum living in or near the pit. Pictures of the insect specimens from Sheriden Pit can be found in Figures 12 to 15.
General Diversity
Diversity ranged from very low to quite high in the sites and horizons (Table 13).
To produce a more conservative count, unidentified groups within a family or genus were not counted if the same family or genus appeared. If the family was different, then the genus and/or species was counted. The Bergendorfer Low layer is the most diverse in all
32 categories, having three orders and a possible total of twenty-four species. Sheriden Pit is the least diverse in all categories, having one order and possibly two species.
These numbers should be taken as estimates since many factors can affect the diversity within these sites. Some factors will be mentioned in greater detail under
“Sources of Error” section. Factors that may affect diversity measures include type of habit of preservation, amount of sediment sieved (and amount of organics collected), who identified the insects, and the method of identification used.
Dissimilarity Tests
The Dice and Jaccard coefficients show all sites are separate assemblages when compared to each other (Table 14). At the genus level, the sites and layers show low similarity. Coefficients range from zero (any other site compared to Sheriden Pit) to
0.370 (Bergendorfer High compared to Bergendorfer Low). At the family level, half of the site comparisons show low similarity, ranging from 0.083 (Sheriden Pit and
Bergendorfer Low) to 0.667 (close to 0.8; Sidney and Bergendorfer Low). Family level comparisons tend to have higher coefficients than those between genera, which is expected at higher taxonomic levels.
MCR Analysis
MCR envelopes were created for sites containing species with known current envelopes (Figures 16-20 and Table 15). Individual envelopes were provided by Scott
Elias (University of London). Bergendorfer and Sheriden Pit have only one species in the MCR analysis. Snyder and Bergendorfer High have three species and Bergendorfer
Low has five species in the MCR analysis. Snyder and Bergendorfer Low have a very constrained estimate of summer warmth, ranging from 15.3-19.8ºC and from 15.2-
33
19.3ºC, respectively. The other sites show a wide TMAX range. All sites with MCR data have cooler summer temperatures than the modern summer temperature, with calibrated temperatures of 16.4ºC ± 0.7 to 18.4ºC ± 0.7. All calibrated summer temperatures fall within boreal temperature ranges (low to upper teens degrees Celsius;
Barbour et al., 1999 and World Climate, 1996). Even though calibrated TMAXes for
Bergendorfer and Sheriden Pit give a typical boreal temperature, they each only have one species in the envelope, so the biome may range from tundra to boreal to a cool deciduous forest.
Summary of Paleoenvironment
A general paleoenvironment is summarized for each site based on both habitat of insects and MCR analysis. Bergendorfer is most likely a moist to wet boreal habitat based on the presence of Olophrum sp. and the MCR range overlapping a boreal summer temperature. The Sidney site was near water, based on several insect families, but no general biome is known since none of the specimens could be identified to species.
Bergendorfer Low may have been boreal to boreal-tundra ecotone and a very wet environment, based on the MCR and insect analysis. Tundra mean summer temperatures tend to range from 3-12 ºC (Barbour et al., 1999), but may be slightly warmer if at a boundary with a boreal forest. Since Eucnecosum sp. was found in Bergendorfer Low, the site may be part tundra; this genus is found under litter in tundra and boreal habitats.
Bergendorfer High was most likely boreal, since one insect is found today in alpine environments and the MCR summer temperature range and calibrated TMAX fall within a boreal temperature. Bergendorfer High was also riparian (river(s)) and moist in the area based on the insects that were found. Based on MCR analysis, Snyder was most
34
likely a boreal forest with spruce (since Polygraphus rufipennis lives in the bark of spruce) and wet with bogs. Sheriden Pit was moist at the time Arpedium cribratum was buried and slightly cooler than today.
35
DISCUSSION
All sites when compared to each other are separate assemblages at the genus level but a few sites are more similar at the family level. The majority of the sites were wet areas (bogs, running water, standing water, and moist ground) as they are today (rivers at three of the four sites). Even the terrestrial (Sheriden) pit had standing water in it since the genus Arpedium may be found in wet areas. A few sites were boreal based on insect specimens found but are currently highly fragmented deciduous forests within agricultural lands. Also, all of the sites with MCR analysis had cooler summer temperatures than present (≥10º difference).
Changes Through Time
Diversity, temperature and habitat changes will be discussed in this section, from oldest to youngest site and horizon. Diversity starts off low 44,000 to 40,000 yr BP
(Bergendorfer and Sidney), is highest 21,640 yrs BP (Bergendorfer Low) and drops down to a low diversity again 11,557 to <9,844 yrs BP(Sheriden Pit) (Tables 3-8 and Table
13). These differences could be caused by actual habitat or geographical differences, preservation biases, amount of material sieved, or the method of floating organics. Some of these factors will be discussed in greater detail below.
Based on MCR analyses, temperatures at all sites with MCR analysis were cooler than today. The oldest site (Middle Wisconsinan) was 16.4 ºC (61.5ºF), nearly seven degrees Celsius below modern mean summer temperatures (23.2ºC). By 21,000 to
19,000 yr BP, the region warms slightly, reaching a summer maximum of 17.2 ºC
(63.0ºF), six degrees Celsius cooler than today (low 70’s °F). Finally, temperatures
36
continued to warm up to 18.7 ºC (65.7ºF) at the end of the Pleistocene and early
Holocene.
The central region of eastern Indiana and western Ohio contained a boreal and
wetland habitat 44,000 yr BP. A moist to wetland habitat persisted in the area during the
ages of the sites, although the temperatures indicate two slight warming shifts from cooler boreal temperatures to slightly warmer boreal temperatures (Figure 21; calibrated
temperatures and the TMAX range for each site through time).
Specimens Found Often
Specimens found often during different time periods may be an indicator of a
species’ ability to be mobile through time due to changes in the environment or that they recolonize the same site again and again. Recolonization can only be concluded if an actual warm period occurred between sites, but for this study, no dramatic changes
occurred between the sites. There are three species that are found in more than one site
or horizon, Acidota subcarinata, Bembidion nigripes, and Helophorus sempervarians.
These species are found in Bergendorfer Low, Bergendorfer High, and Snyder. These
two sites are very close geographically but also in time period, which may explain the
occurrence of the same species in all sites.
Correlations Between Glaciation Events and Temperature and Habitat of the Sites
The majority of the tri-state area between 55,000 and 24,000 yr BP was ice free,
except for a possible advance around 33,000 yr BP in the Grand Rapids, Michigan region
(Eschman, 1985). The Bergendorfer and Sidney sites fall within this time frame.
Bergendorfer’s boreal habitat and climate would possibly correlate with the geographic position of the glacier during this time period. Boreal forests border the arctic tundra,
37 which in turn border polar glaciers (Barbour et al., 1999). At the time of glacial advances, sometimes only 800 km separated the tundra from the deciduous (oak-hickory- pine) forest region (Barbour et al., 1999). The Bergendorfer and Snyder sites are located near Connersville, Indiana, about 479 km (297 miles) away from Grand Rapids,
Michigan.
The farthest glacial advance of the Nissouri substage (21,000 yr BP to 18,000 yr
BP) nearly borders the Bergendorfer Low and High and Snyder locations (Figure 2b).
Bergendorfer Low and High are just slightly older than the start of maximum extent of glacial activity. These two horizons coincide with cool to cold temperatures associated geographically near a glacier advance (Figure 3). The maximum summer temperatures are mostly boreal but also border on the high end of a low arctic tundra summer average temperature (8-12°C; Barbour et al., 1999). The Snyder site falls within the timespan of the glacier expansion period. It also coincides with cold summer temperatures and a boreal zone expected near glacial activity (Figure 3).
Most of the dated sediments of the Sheriden Pit site date between and after two glacial advances, the Greatlaken (11,000 yr BP) and the Younger Dryas event (9900 yr
BP) in which the Marquette ice sheet advanced into northern Michigan (Figure 2b and
Figure 3). The temperature range for one layer analyzed (110-120cm) could possibly coincide with this activity, since it is cooler than today, being boreal-temperate borderline. However, this estimated temperature is also warmer than the older sites that may have been much closer to glacial activity.
38
Correlations with Previous Studies
The Bergendorfer, Snyder, and Sidney sites have been extensively studied in the
past several decades by numerous researchers, mostly in soil geomorphology and glacial
studies. A few studies have been done at the sites on pollen, mollusks, and insects. The
majority of these studies find similar conclusions to my analysis. Knollenberg (1996)
studied pollen (see METHODS section for list of plant taxa) and came to the conclusion that Bergendorfer, in general, was a cold and damp environment with one of the section profiles showing a fluvial depositional environment. Knollenberg also came to the conclusion based on an insect analysis that the site was Boreal-Montane. Hall (Hall and
Zbieszkowski, 2000) studied mollusks in the upper organic zone (Bergendorfer Low and
High) and found common snails and an ostracod, and came to the conclusion based only on these that cold ponded water must have been present. The Sefton site in Indiana, 11 km southeast of Bergendorfer and aged >44,000 yr BP and >46,100 yr BP, shows a moist terrestrial (fen) boreal habitat based on small mollusks and one insect (Lowell and
Brockman, 1994; Hall and Zbieszkowski 2000). The Wildman site in Indiana, northeast of Bergendorfer (26km) and 42,800±1400 yr BP and 42,900±4000 yr BP, also was boreal, based on fern pollen (Hall and Zbieszkowski, 2000). These studies correlate with my analysis of Bergendorfer, but Bergendorfer may have also been partly tundra to tundra-boreal ecotone, based on Eucnecosum sp. and the low range of the TMAX .
The Sidney site has had extensive pollen analysis done. Forsyth (1965) studied the sequence of pollen within the organic layer and interpreted the habitat as a poorly drained upland with upland depressions and a cool but not cold climate that became slightly cooler and moister. Forsyth found a greater percentage of pine (70%), about 20%
39 shrubs and herbs (mostly alder), and 10% deciduous trees (mostly poplar) in the lower part of the organic layer. The pine decreased to 50% while the shrubs and herbs (mostly alder and salix) and deciduous trees increased (30% and 20% respectively). Knollenberg
(1996) found a different percentage of these main groups (see METHODS section), but the percentages are similar to the top of the organic profile Forsyth studied at Sidney.
Knollenberg came to the conclusion that Sidney was an open pine forest. Porter, 100 km southwest of Sidney and 39,900±3000 yr BP, shows a warm temperate climate and riverine environment, based on snails and mollusks (Gooding, 1963; Hall and
Zbieszkowski, 2000). At multiple other sites within the geographic region 80-90 km southwest of Sidney in eastern Indiana, a cool to cold temperature, possibly a north temperate to boreal environment, was present during this time span, based on moss, arctic herbaceous pollen, coniferous pollen, and mollusks (Gooding ,1963; Hall and
Zbieszkowski, 2000). A few of my insect families indicate moist conditions, which support Forsyth’s (1965) claim that Sidney became more moist.
Several studies of the Snyder site have been done, mainly with mollusks (Lowell and Brockman, 1994; Hall and Zbieszkowski, 2000) and pollen (Knollenberg, 1996).
Lowell and Brockman’s field guide (1994) includes a summary of work performed by
Barry Miller and Mudge Morris (date and journal unknown) in which they studied mollusks in east central Indiana. They found terrestrial and aquatic mollusks that represent a cool to cold climate, possibly boreal. This is contrary to Hall (Hall and D.J.
Zbieszkowski, 2000), who found mollusks that represent a temperate (Mississippi valley) climate (Elliptio cf. dilatata). These mollusks from Hall were analyzed in the diamicton, not the organic layer that Miller and Morris analyzed.
40
Knollenberg (1996) studied the pollen of the Snyder site and soil characteristics
(see METHODS section). Knollenberg came to the conclusion based on the pollen that
Snyder was an open boreal (conifer) forest with spruce predominating (44%). The presence of algae suggests ponded water and pools. The soil characteristics also suggest a fluvial environment due to water lain banded layers.
My study agrees with the findings of Barry Miller and Mudge Morris. They concluded that mollusks indicated a boreal habitat and wet conditions, which corroborates Knollenberg’s pollen analysis. Other sites also show a colder temperature than present around this time in Indiana. Russellville, 140 km west of Snyder and aged
19,510±140 to 21,320±300 yr BP, shows a moist to wet mid to southern boreal ecotone
(16-17ºC) based on multiple paleoenvironmental indicators (moss, insects, mollusks, tree pollen) (Hall and D.J. Zbieszkowski, 2000). The Connersville Interstadial Section, east central Indiana and 18,750±300 to 20,000±800 yr BP, also shows a colder and wet
(boggy) climate based on pollen (Gooding, 1963). Finally, leading up to the Snyder site age, 27,000 to 23,000 yr BP sites in Garfield Heights, Ohio (near Cleveland) and New
Paris, Ohio (west central Ohio on the Indiana border), show a general trend of colder climates starting with the oldest (Garfield Heights) having a severe boreal temperature, to boreal environment, to tundra-forest conditions, to open boreal forest (New Paris), based on pollen, gastropods, and insects (Coope, 1968; Szabo, 1997; Hall and Zbieszkowski ,
2000).
An extensive vertebrate analysis has been done on the Sheriden Pit. This analysis shows many terrestrial and aquatic animals living around and in the pit. Other studies within the pit include pollen, stable carbon isotope values, and fluctuations within
41
magnetic fields created when surface soils accumulate in caves. The pollen analysis was
used to reconstruct the habitat. The carbon isotopes in plant remains and animal bones of
herbivores were used to distinguish dry environments and wet environments through
time. Magnetism studies within the sediment layers were done to verify time changes
and climate fluctuations. A cooler and drier environment produces a lower magnetic
signal, and a higher signal results from warmer, wetter times (Tankersley and Redmond,
2000).
Around 12,000 to 13,000 yr BP, the vegetation consisted of mostly spruce and
pine with possible streams and wetlands (Tankersley and Redmond, 2000). Vertebrate
species that now inhabit northern North America such as caribou, pine marten, and
lemming, were found in Sheriden Pit (Tankersley and Redmond, 2000). Tankersley and
Redmond (2000) found that between 13,000 and 10,000 yr BP, the magnetic variations
show a rapid and profound climate change oscillating back and forth from cold to warm
and warm to cold climates. Near the end of the fluctuation (11,000 to 10,500 yr BP), the
climate became cold and dry. Stable carbon isotope fluctuations within plant and animal
bones also show a colder and drier environment around 11,000 to 10,500 yr BP
(Tankersley and Redmond, 2000). Holman (1995) studied the herpetofauna in the top
bone bearing layer, 11,700 yr BP, and found that by this time, the modern herpetofauna had established itself. My analysis of Sheriden Pit may correlate with one of the time periods that shows a cool and moist habitat, either 12,000 to 13,000 yr BP if the layer is below base or during one of the cooling periods (e.g., Late Holocene Cooling Event) within the Holocene if the layer is above base (Figure 3).
42
Sources of Error
The two major problems this study faces are the lack of identifiable insect
specimens in several horizons and sites, and the lack of information on habitat and
behavior in the literature of insect species found in this study. There are multiple reasons
why there may have been a lack of identifiable insects. One of the main reasons may be that too little sediment was sieved. Enough sediment must be obtained and processed to get adequate organic material for a reasonable number of identifiable insect fragments.
This sample size can range from about 20 kg or less to more than 1,000 kg (to get 1 kg of organic detritus) of sediments (Elias, 1994). For this study, around 15 eight ounce cups
(4.6 kg) of sediment was sieved for each horizon. This smaller amount of sediment may have, therefore, not been sufficient.
Another possible reason for the lack of identifiable insects would be the length of time the collected sediment was stored before processing. If the sediment will not be immediately examined, it must be stored in a cool place and kept moist. The material from Sheriden Pit was collected in 1994, and not processed for this study until late 2002.
The sediment was stored in large plastic buckets, some of which were cracked or
deteriorated, allowing the sediment samples to dry. Drying of sediment will tend to break
up insect remains, thus fragmenting them even more (Elias, 1994). Samples from the
Sidney site were kept moist and processed quickly.
Identifying the insects was only done with light microscopy and macrostructural
features. If SEM was incorporated, some of the insects may have been identified down to
a lower taxonomic level, based on microstructures of the elytra and pronotum. Molecular
studies of Late Quaternary insects have also recently been incorporated into identification
43
and population studies. These two methods are more expensive than the methods used
for this study (little to no expense). Another drawback with the molecular technique is
the lack of available sequence data, specifically for species identification.
Finally, there is a lack of information on habitat and behavior within the literature
for the species identified. This is possibly because of the enormous number of species of
insects, especially coleopterans (the largest order of insects). There are just not enough entomologists and ecologists to describe the behavior, eating habits, and ecology of all beetles and other insects. Certain families (e.g., Staphylinidae) are problem groups with many revisions constantly being made (Pavuk, pers. comm. 2004). Also, when species are collected by entomologists, the habitat and ecology of the specimens are rarely recorded while in the field. Within biology in general, more and more experimentation and manipulation of species is being performed at the expense of basic natural history.
Lovejoy (1988) mentions that “we have such a poor inventory of life on earth, biologists
can say relatively little about specie and that a revitalization of the science of biological
systematics is needed with intense exploration.” Little money and few jobs are out there
for people to focus on collecting organisms and describing their natural history.
Taphonomy of the Fossil Insect Specimens
The taphonomy or preservation of the specimens found may have been a factor
when comparing wet to terrestrial (dry) habitats. Recall that the terrestrial Sheriden Pit had the lowest diversity of insects; however, Bergendorfer also had a very low diversity.
Wet environments, specifically standing or moving water, tend to preserve fossils better due to more rapid sedimentation and deposition. The moisture also keeps the organic tissue from rotting quickly. Lower oxygen levels within wet environments (e.g., bog vs.
44
clear moving stream) promote preservation. Terrestrial (land) environments tend to be
drier, usually have less deposition and are exposed to surface erosion more often than wet
environments, allowing dead organisms to be exposed to air and scavengers for longer
periods.
Insect fragments found in Sidney and Sheriden Pit are of a wide range of size and
body part types. The sizes of known insect fragments range from 0.2 mm to about 2 mm
in length. Many fragments were found, but some intact specimens were also found.
Elytra, pronota, a head section, spines, mandibles, and leg segments with surface
ornament intact were found in Sidney and Sheriden Pit. The best range of well preserved
body parts comes from the 0-5 cm a.b. layer of Sheriden Pit (Figure 13), with no
evidence of chemical degradation. Even though they were not identifiable to species, it
shows what can be preserved in a terrestrial habitat with moist conditions. One of the
smallest fragments also comes from Sheriden Pit, layer 35-40 cm a.b. (Figure 14). This
leg is very tiny, with a total length of 0.63 mm. The Sidney site had many more
fragments and better preserved elytra. The largest fragments were also found in Sidney,
including the 2 mm leg piece. Hence, even though a terrestrial habitat may generally
preserve less organic material, the material that is preserved can be in excellent condition.
Some insect species are more likely to be transported away from their living
habitat than others. Transported specimens can then affect the interpretation of a habitat
or biome at a location. This study found both aquatic and terrestrial insects within the sediment sieved. Aquatic insects may mislead the researcher, especially if they are riverine species, as they could be transported quite far before burial. Closed water sources (ponds, bogs, and lakes) are more likely to have burial in place or transportation
45
merely to another location within the closed water source after death. Several bog species were found and probably would not lead to misleading results. Terrestrial species, especially carabids, may also be more likely to show burial in place. Carabids are closely associated with sediments, living on or within soils. These insects most likely were buried in place, especially since most organic zones sampled for this study were paleosols.
46
CONCLUSION
The first objective of the study, using insects as paleoclimate indicators, was successful. This study has found that insects, especially Coleoptera, are useful in determining the Quaternary paleoenvironments, especially when incorporating not only habitat information but MCR analysis. The results of the present study correlate quite well with previous studies done at the same sites and general glaciation patterns during
the time period studied.
The second objective of the study, seeing whether the same species assemblage
re-occurs through time if the right temperature and habitat came back, found that
assemblages do not recur. When the same temperatures occurred at two sites of different
ages and between two horizons of different ages, the beetle assemblages were somewhat
similar (i.e., had some genera and families in common) but were statistically different
overall. Very few insects were identified to species, but if they were, the similarity
indexes would most likely show a larger difference between sites. Caution should be
taken in interpreting these data however, since a small number of specimens were
identified and a small amount of sediment was processed. Also, Knollenberg (1996)
mentions no estimates of the amount of sediment processed for the insects in her study.
This analysis may show that insects have a more individualistic approach to
migrating when climate changes, similar to the pattern seen in many studies done on
plant communities in the Quaternary. Alternatively, the changes seen may reflect
community turnover and spatial differences within a persisting boreal biome. We must
understand community processes to better anticipate changes associated with current
global warming. Communities may shift geographically and retain the same composition
47
or they may re-mix and have a different composition and/or ecological structure in the future. We should not be concerned so much with keeping whole communities intact, but we should keep an individualistic approach to species conservation. A species may find a niche that is suitable even in a different structured community.
This study found several insect families within two previously undescribed Late
Quaternary sites (Sheriden Pit and Sidney), and one insect was identified to the species level (Arpedium cribratum from Sheriden Pit). If a larger amount of sediment is sieved and processed, and SEM (and possibly molecular techniques) are used, more insects might be identified to genus and species level. Tracking families is a good indicator of the changes of diversity within a site (Ponel et al., 2003), but to get more specific information about habitat and a temperature range, genera and species are needed.
Since the samples processes for analysis were at the low end of the amount of sediment necessary, more studies need to be done. A larger amount of sediment may yield more identifiable fragments within the sites analyzed for this study. There are also many other sites surrounding Connersville, Indiana and Sidney, Ohio that still have not been sampled for insect analysis. These sites are close to the ages of this current study and also, some of the sites are in between the ages. This would make an ideal location for future insect studies to examine a small geographic area and timespan for habitat, climate, and community changes through the Late Quaternary.
48
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Williams, J.W., T. Webb III, P.H. Richard, and P. Newby. 2000. Late Quaternary
biomes of Canada and the eastern United States. Journal of Biogeography 27:
585-607.
Williams, J.W., B.N. Shuman, and T. Webb III. 2001. Dissimilarity analyses of Late
Quaternary vegetation and climate in eastern North America. Ecology 82(12):
3346-3362.
Williams, J.W., B.N. Shuman, T. Webb III, P.J. Bartlein, and P.L. Leduc. 2004. Late
Quaternary vegetation dynamics in North America: Scaling from taxa to biomes.
Ecological Monographs 74(2): 309-334.
World Climate. 1996. Internet 13 August 2004. Available WWW:
http://www.worldclimate.com/
57
Figure 1: Stages of glaciation in the tri-state area (Eschman, 1985). Substages represent times of glacial advance, while interstadials represent times of glacial retreat.
58
a.) b.)
Figure 2: Glacier advance limits during the Pleistocene and site localities. a.) Glacial advance limits of the three stages of the Pleistocene (pre-Illinoian, Illinoian, and Wisconsinan). b.) Glacial advance limits of the four Wisconsinan substages: N-Nissouri: 21,000 yr BP; PB-Port Bruce: 15,500 yr BP; PH-Port Huron: 13,000 yr BP; G- Greatlakean: 11,000 yr BP, and the Marquette (M) re-advance that occurred in the Early Holocene (Eshman, 1985). The stars are this study’s site locations, starting with the lowest star and working up in a northeast direction: Connersville, Indiana (Bergendorfer and Snyder sites); Sidney, Ohio; and 10 miles east of Findlay, Ohio (Sheriden Pit site).
59
Figure 3: Summary of glaciation and floral and faunal changes throughout the Late Pleistocene and Holocene. Cold represents organisms that are found currently in northern latitudes (e.g., boreal) and warm represents organisms that are presently found in the Great Lakes region or farther south (e.g., deciduous forest). The Vegetation line fluctuates to represent changes from cold to colder biomes and warm to warmer biomes through time. The Glacier diagram shows magnitude differences between the Pleistocene and Holocene, since glacier fluctuations were more pronounced during the Pleistocene than the Holocene. References for Vegetation: Adams (1997) and Adams and Fuare (1997); Herpetofauna: Holman (1992 and 1995); Mammals: Hansen (1992) and Lange (2002); and Glaciers: Eschman (1985), Grove (2001), and Spencer (2003).
60
Figure 4: Hypothetical temperature ranges of two insect species a) having a mutual climate range b) not having a mutual climate range.
A B
CD
Figure 5: Sidney, Ohio insect fragments: (a) Carabidae: Bemidion sp. right elytron, (b) Curculionidae: thorax, (c) Carabidae: right 61 elytron, (d) Chrysomelidae: Donociinae Gen. indet. elytral fragment. Black bar length = 1 mm E F
GH
Figure 5 continued… (e, f) Carabidae: Bemidion sp. left elytron, (g) Curculionidae: tribe Bagoini right elytron, (h) Scarabaeidae: Gen. indet. head. Black bar length = 1 mm 62 I
Figure 5 continued… (i) Staphylinidae: Subfamily Aleocharinae pronotum. Black bar length = 1 mm 63 A B C
D E F
Figure 6: Sidney, Ohio insect fragments that were not identified: (a) leg, (b) beetle mandible, (c) mandible, (d) sclerite, (e) spiny sclerite, (f) sclerite. (a) black bar length = 1 mm, (b-f) black bar length = 0.5 mm 64 G H I
JKL
Figure 6 continued… (g) sclerite, (h) sclerite, (i) mandible, (j) elytron, (k) elytron?, (l) sclerite. (g, k, and l) Black bar length = 1 mm, (h, i, and j) Black bar length = 0.5 mm 65 M NO
P
Figure 6 continued… (m) sclerite, (n) elytron, (o) sclerite and elytron, (p) sclerite. (n and o) Black bar length = 1 mm, (m and p) Black bar length = 0.5 mm 66
67
Figure 7: Distribution map of Helophorus sempervarians. (Smetana, 1988, pg. 51).
68
Figure 8: Distribution map of Tachinus frigidus. (Campbell, 1973, pg. 73).
69
Figure 9: Distribution map of Helophorus lacustris. (Smetana, 1988, pg. 88).
70
Figure 10: Distribution map of Lordithon longiceps. (Campbell, 1982, pg. 89).
71
Figure 11: Distribution map of Polygraphus rufipennis. (Bright, 1976, pg. 102).
Figure 12: Insect fragments from Sheriden Pit, horizon 50-55 cm below base: Sclerites of unknown specimen. Black bar length = 1 mm 72 A B
C D E
Figure 13: Insect fragments from Sheriden Pit, horizon 0-5 cm above base and contact: (a) Staphylinidae: Omaliinae Gen. indet. right
elytron, (b-e) unknown fragments: pronotum?, leg, leg and possibly mandible, sclerite. (a) Black bar length = 1 mm, (b-e) Black bar 73 length = 0.5 mm A B
Figure 14: Insect fragments from Sheriden Pit, 35-40 cm above base: (a, b) leg segments that were originally attached to each other. Black bar length = 0.25 mm 74 A B
Figure 15: Insect fragments from Sheriden Pit, 110-120 cm: (a) Staphylinidae: Arpedium cribratum Ful. pronotum, (b) sclerite. Black bar length = 0.5 mm 75 76 Figure 16: MCR envelope for Bergendorfer. 77 Aegialia lacustris Tachinus frigidus Acidota subcarinata Helophorus sempervarians Bembidion nigripes Figure 17: MCR envelope for Bergendorfer Low. 78 Figure 18: MCR envelope for Bergendorfer High. Acidota subcarinata Tachyporus rulomus Helophorus sempervarians
Figure 19: MCR envelope for Snyder. 79 80 Figure 20: MCR envelope for Sheriden Pit, 110-120cm. Sheriden Pit: 110-120cm MCR Temperature Ranges Through Time 30
25
20
15 TMAX low TMAX high TMAX cal.
10 Temperature in Celsius in Temperature
5
50,000 45,000 40,000 35,000 30,000 25,000 20,000 15,000 10,000 5,000 0 Date Before Present 0
Figure 21: TMAX and TMAX calibrated temperatures (standard error ±0.7ºC) through time for the sites studied: 44,000 BP to 10,500 BP (mid-range for Sheriden Pit temperature data). 81
82
Table 1: Sites and their associated organic horizons for fossil insects included in this study.
RADIOCARBON SITE COUNTY STATE HORIZON DATES (YR BP) Sheriden Pit Wyandot Ohio ITC:35-40 cm a.b. < 9,844 Sheriden Pit Wyandot Ohio ITC:25-30 cm a.b. < 9,844 Sheriden Pit Wyandot Ohio ITC:15-20 cm a.b. < 9,844 Sheriden Pit Wyandot Ohio ITC:0-5 cm a.b. 9,844 Sheriden Pit Wyandot Ohio ITC:50-55 cm b.b. 11,557 Sheriden Pit Wyandot Ohio ITC:110-120 cm ?? Snyder Fayette Indiana Snyder 19,700 Bergendorfer Fayette Indiana Berg High 21,640 Bergendorfer Fayette Indiana Berg Low 21,640 Sidney Shelby Ohio Sidney 40,000 Bergendorfer Fayette Indiana Bergendorfer 44,000
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Table 2: Radiocarbon dates of all sites and material used.
Site Section Age Error Material dated Reference (+/-) Bergendorfer BD-2 44000 2000 bulk organics from Ab Knollenberg horizon (1996)* Bergendorfer BD-4 21190 360 bulk organics from silt Knollenberg above paleosol (1996)* Sidney BC-1 42800 4000 wood from Ab horizon Unpublished* Sidney BC-1 41200 3000 bulk organics from Ab Unpublished* horizon Sidney BC-1 >39300 ---- peat form Ab horizon Forsyth (1965)* Sidney BC-1 >37000 ---- peat form Ab horizon Forsyth (1965)* Snyder SN-1 19700 180 bulk organics from Ab Hall (1992)* horizon Sheriden Pit 5a 11610 90 wood charcoal and Tankersley and collagen Sheriden Pit 5a 11570 50 " " Redmond (2000); Sheriden Pit 5a 11570 70 " " Sheriden Cave Site (2001) Sheriden Pit 5a 11480 60 " " " " Sheriden Pit 6 10020 115 " " " " Sheriden Pit 6 9775 70 " " " " Sheriden Pit 6 9190 60 " " " " Sheriden Pit 6 9170 60 " " " "
* Taken from Hall and Zbieszkowski (2000) field guide.
Table 3: Insect specimens found in Bergendorfer.
Bergendorfer ORDER FAMILY GENUS SPECIES NUMBER 44,000BP+-2,000yrs Coleoptera Carabidae Dyschirius sp. 1 Carabidae Amara sp. 1 Staphylinidae Olophrum sp. 1 Staphylinidae Acidota crenata 1 Trichoptera Limnephilidae indetermined species 1 Acari Oribatei indetermined species 1
Table 4: Insect specimens found in Sidney.
Sidney ORDER FAMILY SUBFAMILY/TRIBE GENUS SPECIES NUMBER 40,000 BP Coleoptera Carabidae Bembidion sp. 4 Carabidae indetermined species 2 Chrysomelidae Donaciinae indetermined species 2 Curculionidae Bagoini indetermined species 1 Curculionidae indetermined species 1 Dytiscidae Agabini indetermined species 1 Scarabaeidae indetermined species 1 Staphylinidae Aleocharinae indetermined species 1 84 Table 5: Insect specimens found in Bergendorfer Low.
Bergendorfer Low ORDER FAMILY GENUS SPECIES NUMBER 21,640BP+-150yrs Coleoptera Carabidae Bembidion sp 1 Carabidae Bembidion nigripes 1 Haliplidae Haliplus sp 1 Dytiscidae Hydroporus sp 1 Hydrophilidae Helophorus sempervarians 3 Hydrophilidae Helophorus sp 4 Hydrophilidae Berosus sp 1 Staphylinidae Eucnecosum sp 1 Staphylinidae Olophrum sp 1 Staphylinidae Acidota subcarinata 1 Staphylinidae Geodromicus sp 4 Staphylinidae Tachinus frigidus 2 Staphylinidae Aleocharine sp. 4 Scarabaeidae Aphodius sp 1 Scarabaeidae Aegialia lacustris 2 Scarabaeidae Serica sp 1 Heteroceridae Heteocerus sp 1 Elateridae Ctenicera sp 1 Chrysomelidae Graphops sp 2 Chrysomelidae Pyrrhalta sp 1 Curculionidae Notaris sp 1 Curculionidae indertermined species 3 Trichoptera Limnephilidae indertermined species 5 Hymenoptera Chalcidoidea indertermined species 2 85 Table 6: Insect specimens found in Bergendorfer High. Bergendorfer High ORDER FAMILY GENUS SPECIES NUMBER 21,640BP+-150yrs Coleoptera Carabidae Bembidon nigripes 1 Carabidae Agonum sp 1 Hydrophilidae Hydrobius sp 1 Hydrophilidae Helophorus lacustris 1 Hydrophilidae Helophorus sempervarians 3 Hydrophilidae Helophorus sp 3 Staphylinidae Geodromicus sp 2 Staphylinidae Tachinus sp 1 Staphylinidae Aleocharine sp 2 Scarabaeidae Aphodius sp 2 Elateridae indetermined species 1 Acari Oribatei indetermined species 1 Crustacea Ostracoda indetermined species 1 Daphniidae Daphnia sp 1 86 Table 7: Insect specimens found in Snyder. Snyder ORDER FAMILY GENUS SPECIES NUMBER 19,700BP+-180yrs Coleoptera Carabidae Dyschirius sp. 3 Carabidae indetermined species 1 Dytiscidae Hydroporus sp 1 Hydrophilidae Helophorus sempervarians 1 Staphylinidae Acidota subcarinata 4 Staphylinidae Tachyporus rulomus 1 Staphylinidae Lordithon cf. longiceps 1 Elateridae indetermined species 1 Nitidulidae indetermined species 1 Curculionidae indetermined species 1 Scolytidae Polygraphus rufipennis 1 Homoptera Cicadellidae indetermined species 1 Trichoptera Limnephilidae indetermined species 1
Table 8: Insect specimens found in Sheriden Pit.
Sheriden Pit ORDER FAMILY SUBFAMILY/TRIBE GENUS SPECIES NUMBER 10,500-9,170 B.P. 0-5 cm a.b. Coleoptera Staphylinidae Omalinae indetermined species 5 to 6 110-120 cm Staphylinidae Arpedium cribratum 1 87 Table 9: Modern day biogeography and ecology of insect specimens found in Bergendorfer. Bergendorfer GENUS SPECIES BIOGEOGRAPHY ECOSYSTEM HABITAT Dyschirius sp. geographically widespread hygrophilous; burrows in sandy and clayey soils Amara sp. geographically widespread Olophrum sp. widespread in North America boreal and alpine litter, moss, rodent nests and middens; near water Acidota crenata northern transcontinental: NM to VA; neoarctic 88 Table 10: Modern day biogeography and ecology of insect specimens found in Bergendorfer Low. Bergendorfer Low GENUS SPECIES BIOGEOGRAPHY ECOSYSTEM HABITAT Bembidion sp worldwide distribution Haliplus sp neoarctic; found throughout NA widerange Helophorus sempervarians palearctic and neoarctic Helophorus sp palearctic and neoarctic Berosus sp generally distributed Eucnecosum sp Alaska, Canada, Rocky Mts, boreal and tundra litter, near water northern MI, NY Olophrum sp widespread in North America boreal and alpine litter, moss, rodent nests and middens; near water Acidota subcarinata eastern north america forests and bogs litter and moss Geodromicus sp mostly montane and west, and AK riparian often found in moss and wet litter Tachinus frigidus generally distributed widerange widerange Aphodius sp worldwide distribution widerange multiple Aegialia lacustris throughout NA; Worldwide genus Serica sp widely distributed in Canada and NA Graphops sp all species found north of Mexico Pyrrhalta sp Palearctic Notaris sp generally distributed, in NA and wetlands associated with cattails, Typha 89 Canada (Typhaceae) Table 11: Modern day biogeography and ecology of insect specimens found in Bergendorfer High. Bergendorfer High GENUS SPECIES BIOGEOGRAPHY ECOSYSTEM HABITAT Bembidon nigripes Agonum sp widespread in Americas; mostly neoarctic alpine forests most hygrophilous Hydrobius sp widespread in North America Helophorus lacustris palearctic and neoarctic Helophorus sempervarians palearctic and neoarctic Helophorus sp palearctic and neoarctic Geodromicus sp generally distributed, mostly montane and west, and AK riparian moss and wet litter Tachinus sp generally distributed widerange widerange Aphodius sp worldwide distribution widerange multiple
Table 12: Modern day biogeography and ecology of insect specimens found in Snyder. Snyder GENUS SPECIES BIOGEOGRAPHY ECOSYSTEM HABITAT Dyschirius sp. globally widespread Helophorus sempervarians palearctic and neoarctic Acidota subcarinata eastern north america forests and bogs litter and moss Tachyporus rulomus generally distributed Lordithon cf. L. longiceps generally distributed 90 Polygraphus rufipennis worldwide live in phloem of broken or fallen Picea
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Table 13: Diversity of sites at different taxonomic levels.
Bergendorfer Sidney Berg Berg Snyder Sheriden Pit Low High Total orders 2 1 3 1 3 1 Total families 3 5 12 5 10 1 Total genera 5 5 21 9 12 1 Total possible 5 6 24 11 13 2 species Species 1 0 5 3 5 1 identified
Table 14: Dice and Jaccard coefficients for genera and families of all the sites.
Jaccard Dice Jaccard Dice Genera Genera Families Families Bergendorfer+Snyder 0.222 0.364 0.300 0.461 Bergendorfer+BergLow 0.095 0.174 0.250 0.400 Bergendorfer+BergHigh 0 0 0.333 0.500 BergLow+BergHigh 0.227 0.370 0.417 0.588 Sheriden Pit+Snyder 0 0 0.100 0.182 Sheriden Pit+Bergendorfer 0 0 0.333 0.500 Sheriden Pit+BergLow 0 0 0.083 0.154 Sheriden Pit+BergHigh 0 0 0.200 0.333 Sidney+Snyder 0 0 0.250 0.400 Sidney+Bergendorfer 0 0 0.333 0.500 Sidney+BergLow 0.053 0.100 0.417 0.588 Sidney+BergHigh 0.125 0.222 0.428 0.600 Sidney+Sheriden Pit 0 0 0.200 0.333 Snyder+BergLow 0.130 0.231 0.467 0.636 Snyder+BergHigh 0.071 0.133 0.364 0.533
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Table 15: MCR analysis for all sites and horizons.
Paleo • Paleo Paleo *Modern *Modern TMAX Calibrated TRANGE TMAX TRANGE Site ºC ºC ºC ºC ºC Sheriden Pit ITC:35-40cma.b. - - - 22.5 26.7 Sheriden Pit ITC:25-30cma.b. - - - 22.5 26.7 Sheriden Pit ITC:15-20cma.b. - - - 22.5 26.7 Sheriden Pit ITC:0-5cma.b. - - - 22.5 26.7 Sheriden Pit ITC:50-55cmb.b. - - - 22.5 26.7 Sheriden Pit ITC:110-120cm 13.8 -25.1 18.7 ± 0.7 27.2 - 37.0 22.5 26.7 Snyder Snyder 15.3-19.8 17.2 ± 0.7 23.7-37.9 23.2 26.8 Bergendorfer Berg High 12-20.9 16.4 ± 0.7 21.8-43.9 23.2 26.8 Bergendorfer Berg Low 15.2-19.3 17.0 ± 0.7 22.2-34.7 23.2 26.8 Sidney Sidney - - - 22.7 26.8 Bergendorfer Bergendorfer 9.0-24.0 16.4 ± 0.7 10.5-40.6 23.2 26.8
9ºC=48ºF, 15ºC=59ºF, 17 ºC=63ºF, 20ºC=68ºF, and 23ºC=73ºF
• Equation for Calibrated TMAXes is from Methods section: Paleoclimate Reconstruction * Data from Internet site: Midwest Regional Climate Data. Historical mean temperatures for the month of July.
Modern TMIN is -4.1ºC for Sidney, -4.2 ºC for Sheriden Pit, and -3.6 ºC for Snyder and Bergendorfer. Historical mean temperatures for January.
Boreal mean temperature range for warmest month 10-19 ºC in summers (Barbour et al., 1999 and World Climate, 1996).
Tundra mean temperature range for warmest month 3-12 ºC warmest (Barbour et al., 1999).