THE COLEOPTERAN FAUNA OF SULTAN CREEK-MOLAS LAKE AREA WITH SPECIAL EMPHASIS ON CARABIDAE AND HOW THE GEOLOGICAL BEDROCK INFLUENCES BIODIVERSITY AND COMMUNITY STRUCTURE IN THE SAN JUAN MOUNTAINS, SAN JUAN COUNTY, COLORADO

Melanie L. Bergolc

A Dissertation

Submitted to the Graduate College of Bowling Green State University in partial fulfillment of the requirements for the degree of

DOCTOR OF PHILOSOPHY

August 2009

Committee:

Daniel Pavuk, Advisor

Kurt Panter Graduate Faculty Representative

Jeff Holland

Rex Lowe

Moira van Staaden

© 2009

Melanie L. Bergolc

All Rights Reserved iii

ABSTRACT

Daniel Pavuk, Advisor

Few studies have been performed on coleopteran (beetle) biodiversity in mountain ecosystems and relating them to multiple environmental factors. None of the studies have examined geologic influences on beetle communities. Little coleopteran research has been performed in the Colorado Rocky Mountains. The main objectives of this study were to catalog the coleopteran fauna of a subalpine meadow in the San Juan

Mountains of Colorado and investigate the role geology had in the community structure of the Carabidae (ground beetles). The study site, a 160,000 m2 plot, was located near

Sultan Creek and Molas Lake in San Juan County, Colorado. Five sites were in each bedrock formation: Molas, Elbert, and Ouray-Leadville. Insects were collected via pitfall trapping in 2006 and 2007, and identified by comparison with museum specimens, museum and insect identification websites, and by taxonomic experts. Biological and physical factors were recorded for each site: detritus cover and weight, plant cover and height, plant species richness, aspect, elevation, slope, soil temperature, pH, moisture, and compressive strength, and sediment size distribution. Quantitative analyses were performed on the Carabidae community and its relationship to bedrock formations and environmental factors measured, and on the entire coleopteran abundance and its relationship to the environmental factors. I sampled 7,316 coleopteran individuals in 27 families and nearly 100 species (61 completely identified). Coleopteran abundance was influenced by plant cover, plant height, soil moisture, aspect, slope, collection site, and week collecting took place. A total of 1,236 carabid individuals representing 30 species were collected. Elevation, detritus cover, and soil temperature were significantly distinct iv

between formations. Species richness and abundance of several species (Agonum placidum, Bembidion mutatum, Carabus taedatus, Cymindis cribricollis, Harpalus animosus, and Harpalus laticeps) differed significantly between formations, but overall there were no distinct carabid communities associated with the three formations. Carabid species richness, abundance, and biodiversity were significantly influenced by plant factors, and frequency was significantly influenced by sediment sorting. The carabid species composition was influenced (not significantly) by compressive strength, pH, plant cover, soil moisture, slope, and aspect. In conclusion, geology had less of an effect on carabidae then other environmental factors. v

ACKNOWLEDGMENTS

This project was partially funded by The Explorers Club Exploration Fund (2006;

$1,149), Mountain Studies Institute and their Mini-Grant Program (2006; $500 and 2008;

$1,000), Geological Society of America Student Research Grant (2007; $2,800), Bowling

Green State University Graduate College’s Katzner Award (2008; $960), and Bowling

Green State University Biological Sciences Department’s Oman Graduate Scholarship

(2008; $750). Permits were provided by the San Juan National Forest (2006 and 2007) to allow me to trap insects at the Molas Lake region in San Juan County, Colorado. I would like to thank the following people for field and lab assistance: Candace Brendler, Minna

Swanson-Theisen, Mike Carey, Matt Territ, and Imtiaz Rangwala. Joanna Lemly and

Kerry Cutler identified several of my plant species. The following people helped identify my beetles: Daniel Duran (Cicindelidae; Ph.D. student currently at Vanderbilt

University), Bill Warner (Scarabaeidae; Farnam Companies, INC), Dan Pavuk (Michigan

State University-The Ohio State University Extension), Donald E. Bright (Curculionidae and Scolytidae; Colorado State University), Boris Kondratieff (Colorado State

University), Virginia Scott (The University of Colorado), Shawn M. Clark

(Chrysomelidae and several uncommon families; Brigham Young University), Paul J.

Johnson (Byrrhidae; South Dakota State University), Stewart Peck (Leiodidae; Carleton

University, Canada), Sam Wells (Elateridae; Bayer CropScience LP, Fresno, CA), Bob

Davidson (Carabidae; Carnegie Museum of Natural History), and Foster Purrington

(Carabidae; The Ohio State University). I would like to thank Boris Kondratieff for access to the Colorado State University’s Entomology Collections and Virginia Scott for access to the University of Colorado’s Entomology Collections, and for allowing me to work in the collections to identify my beetle specimens. Peg Yacobucci and Jeff Holland vi

helped with a few of the statistical tests, CCA and ANISOM, respectively. Helen

Michaels lab allowed me to use their oven to dry my detritus samples. I’d also like to thank the Department of Geology at Bowling Green State University (geological maps, access to Sedimentary Lab for sieving my soils, and borrowing a Brunton compass), The

Mountain Studies Institute (low cost lodging and lab space), my good friends in Silverton

(especially Mike Geryak), and my Mayflower Mill (Jim Cole and Bob Boeder) and Avon

Hotel buddies (too many to list here). Finally, I would especially like to thank my advisor Dan Pavuk for all his encouragement, advice, and friendship, and allowing me to create my very own interdisciplinary project to combine my love of geology and ecology. vii

TABLE OF CONTENTS

Page

INTRODUCTION ...... 1

CHAPTER I. The Influence of Geology on the Environmental Factors Related to Carabidae

Distribution…………...... 18

Introduction…………...... 18

Materials and Methods ...... 19

Results………...... 23

Discussion……...... 25

Conclusion…………… ...... 30

References…...... 32

Tables and Figures…...... 39

CHAPTER II. The Impact of Bedrock Formations on Carabidae (Coleoptera) Community

Structure ….…………...... 53

Introduction…………...... 53

Materials and Methods ...... 54

Results………...... 59

Discussion……...... 62

Conclusion…………… ...... 66

References…...... 67

Tables and Figures…...... 75

CHAPTER III. Influences of Environmental Factors on a Carabidae Community in a Subalpine

Meadow ..……………...... 91 viii

Introduction…………...... 91

Materials and Methods ...... 92

Results………...... 95

Discussion……...... 97

Conclusion…………… ...... 100

References…...... 101

Tables and Figures…...... 106

CHAPTER IV. Coleopteran Biodiversity within the Sultan Creek-Molas Lake Study Area in the

San Juan Mountains, San Juan County, Colorado …………...... 120

Introduction…………...... 120

Materials and Methods ...... 120

Results………...... 123

Discussion……...... 125

Conclusion…………… ...... 128

References…...... 129

Tables and Figures…...... 131

GENERAL SUMMARY AND FUTURE STUDIES ...... 144 ix

LIST OF TABLES

Table Page

CHAPTER I

1 Sediment Size Ranges Measured for Soil Analyses ...... 39

2 One-way ANOVAs for all Variables between Formations in 2006 ...... 40

3 One-way ANOVAs for all Variables between Formations in 2007 ...... 41

4 P-values for Linear Regressions Run on Soil Temperature and Time Taken ...... 41

5 Plant Species Found in each Formation ...... 42

6 Jaccard Coefficients for Plant Communities between Formations ...... 43

CHAPTER II

1 Total Abundance Caught at each Site for each Carabid Species ...... 75

2 Total Frequency Calculated for each Carabid Species at each Site ...... 76

3 Carabidae Values Calculated for each Site ...... 77

4 Means and ANOVA Results of Carabid Values within each Formation ...... 77

5 Absolute Total of Carabid Community Measurements for each Formation ...... 77

6 Abundance Statistics between Formations for the Common Species ...... 78

7 Frequency Statistics between Formations for the Common Species ...... 78

8 Relative Abundances of the Species Composition between each Formation ...... 79

9 Relative Frequencies of the Species Composition between each Formation ...... 80

10 Jaccard Coefficients Calculated between Individual Sites ...... 81

11 Overall Jaccard Coefficients Calculated between Formations ...... 81

12 ANOSIM Output for the Jaccard Coefficient ...... 82

x

CHAPTER III

1 Environmental Variables and Their Measurements Taken at each Site ...... 106

2 Carabidae Frequency Data Used for the CCA Matrix ...... 107

3 P-values of the Environmental Factors for each Carabidae Community and

Biodiversity Measurements ...... 108

4 CCA Results of the First Four Axes for the Environmental Variables ...... 109

CHAPTER IV

1 Coleopteran Families Identified at the Sultan Creek-Molas Lake Region ...... 131

2 Species Identified and Unidentified at the Sultan Creek-Molas Lake Region ...... 132

3 ANOVA and Regression Analyses Performed on Abundance for 2006 Variables ... 135

4 ANOVA and Regression Analyses Performed on Abundance for 2007 Variables ... 136

xi

LIST OF FIGURES

Figure Page

CHAPTER I

1 General Map of San Juan Mountains and Silverton Colorado ...... 44

2 Topographic and Aerial Maps of the Study Area and Site Locations ...... 45

3 Cumulative Curve Example for Site 9 ...... 46

4 Mean Soil Moisture in Bedrock Formations, Week 2, 2006 ...... 47

5 Overall Mean Soil Temperature in Bedrock Formations, 2006 ...... 47

6 Overall Mean Soil Temperature in Bedrock Formations, 2007 ...... 48

7 Mean Soil Temperature in Bedrock Formations; Week 4, 2006 ...... 48

8 Mean Soil Temperature in Bedrock Formations; Week 6, 2006 ...... 49

9 Mean Soil Temperature in Bedrock Formations; Week 1, 2007 ...... 49

10 Mean Soil Temperature in Bedrock Formations; Week 5, 2007 ...... 50

11 Linear Regressions for Soil Temperature and Time Taken in 2006 ...... 50

12 Linear Regressions for Soil Temperature and Time Taken in 2007 ...... 51

13 Mean Detritus Cover in Bedrock Formations, 2006 ...... 51

14 Mean Detritus Cover in Bedrock Formations, 2007 ...... 52

15 Mean Elevation of Bedrock Formations ...... 52

CHAPTER II

1 General View of Study Area ...... 83

2 Study Area Showing Bedrock Formations and Where Sites were Located ...... 83

3 Pitfall Trap Set-up Design ...... 84

4 Site Twelve Showing the Typical Pitfall Trap Set-up ...... 84 xii

5 Typical Pitfall Trap used in Study Area ...... 85

6 Mean Species Richness between Bedrock Formations ...... 85

7 Mean Abundance, Frequency, and Biodiversity in Formations ...... 86

8 Individual Carabid Species Mean Abundance in Formations ...... 87

9 Individual Carabid Species Mean Frequency in Formations ...... 88

10 Graphical Output of Analysis of Similarity for Formations using Jaccard Index ..... 89

11 Cluster Analysis of Jaccard Coefficients for the Carabidae Fauna ...... 90

CHAPTER III

1 Topographic Map of the Study Area and Site Locations ...... 110

2 Linear Regression Analysis for Carabid Species Richness versus Plant Species

Richness……… ...... 110

3 Mean Carabid Species Richness in Bedrock Formations ...... 111

4 Linear Regression Analysis for Carabidae Abundance versus Plant Cover ...... 111

5 Linear Regression Analysis for Carabidae Biodiversity versus Plant Cover ...... 112

6 Linear Regression Analysis for Carabidae Evenness versus Plant Cover ...... 112

7 Linear Regression Analysis for Carabidae Frequency versus Sediment Sorting ...... 113

8 Sites and Their Distribution on the First and Second Axes for the CCA ...... 114

9 Cluster Analyses Results for the Sites based on the Environmental Variables ...... 115

10 CCA Results for the Environmental Variables, Axis 1 and 2...... 116

11 CCA Results for the Environmental Variables, Axis 2 and 3...... 117

12 CCA Results for Carabidae Species, Axis 1 and 2 ...... 118

13 CCA Results for Carabidae Species, Axis 2 and 3 ...... 119

xiii

CHAPTER IV

1 Topographic Map of the Study Area and Site Locations ...... 137

2 Total Coleopteran Caught each Week in 2006 Season ...... 137

3 Mean Coleopteran Abundance Caught each Week in 2006 Season ...... 138

4 Mean Coleopteran Abundance Caught at each Site in the 2006 Season ...... 138

5 Linear Regression of Coleopteran Abundance versus Plant Cover in 2006 ...... 139

6 Linear Regression of Coleopteran Abundance versus Plant Height in 2006 ...... 139

7 Total Coleopteran Caught each Week in 2007 Season ...... 140

8 Mean Coleopteran Abundance Caught each Week in 2007 Season ...... 140

9 Mean Coleopteran Abundance Caught at each Site in 2007 Season ...... 141

10 Linear Regression of Coleopteran Abundance versus Plant Cover in 2007 ...... 141

11 Linear Regression of Coleopteran Abundance versus Soil Moisture in 2007 ...... 142

12 Seasonal Correlation between Coleopteran Abundance and Soil Moisture, 2007 .... 142

13 Mean Coleopteran Abundance in Different Aspects in 2007 ...... 143

14 Mean Coleopteran Abundance in Different Slopes in 2007 ...... 143

GENERAL SUMMARY AND CONCLUSIONS

1 Whole Meadow in the Molas Lake Vicinity ...... 147

1

INTRODUCTION

Geoecology is the study of the structure and function of geoecosystems, which include

the interactions of the abiotic components (lithosphere, pedosphere, atmosphere, and

hydrosphere) with that of the ecosystem (biosphere) (Huggett 1995). Studying all the spheres

allows for a more comprehensive view of the world and how everything interacts. In modern

ecological studies, the interactions between the lithosphere and biosphere have been studied less

often than the interactions of the hydrosphere, pedosphere, and atmosphere with the living world.

General patterns in biodiversity, types of communities/biomes, and soil formation processes are

known for climate, rainfall, elevational, and latitudinal differences. Patterns and general

interactions between the lithosphere and biosphere are less known.

The majority of the studies on the interaction of geology and ecology have investigated

how plant species and community structure were related to specific geological bedrocks,

geomorphologic features, and the geological history of an area (e.g., glaciation), which is also

known as the sub-discipline of geobotany (e.g., Read 1952, Whittaker 1954, Tomans 1977,

Romans 1981, Wentworth 1981, Osterkamp and Hupp 1984, Cole 1986, Friedman et al. 1996,

Forsyth 2003, Kruckeberg 2004). Most of these studies have shown distinct plant communities

and/or biodiversity differences when comparing between formations or specific rock types (e.g.,

sandstone versus limestone). Read (1952) observed tree species on chert, limestone with a

cherty surface, limestone, and sandstone, and found a strong quantitative relationship between

geology and dominant tree species in the Ozark Highlands, Arizona. Wentworth (1981) studied

plant communities along elevational and topographic gradients within limestone and granite and

found that the granite had an overall higher number of species compared to limestone at the same

elevation. Also, herbaceous and arborescent species dominated in granite outcrops, while the 2

limestone was dominated by shrubby species (Wentworth 1981). Osterkamp and Hupp (1984) found that woody plant species were distinct on different geomorphic surfaces along riparian streams in northern Virginia.

Fewer studies have included animals and their interactions with the geology of an area

(e.g., Gereben 1995, Moore et al. 2001, Barr and Babbitt 2002, Southworth et al. 2002, Garrison-

Johnston et al. 2003, and Grant et al. 2005). Animals present more of a challenge in studying

geological linkages due to their mobility and dependence on primary productivity. They may

migrate to another area if their habitat becomes unsuitable. Two studies on salamanders and the

relationship to geology showed no difference in biodiversity but one study did show differences

within a population of a particular species. Moore et al. (2001) found that even though the

species did not differ with geomorphic surface, the size and weight of Desmognathus

ochrophaeus differed significantly between floodplain and hillslope compared to Plethodon cinereus, whose weight differed only between cover type (rock cover versus log/leaf litter). In another study, bedrock had no influence on salamander species and abundance in Shenandoah

National Park, Virginia (Grant et al. 2005).

A few studies have shown significant influences of bedrock on animals. Garrison-

Johnston et al. (2003) concluded that Douglas-fir beetle infestations occurred significantly more on outcrops of metasedimentary rocks (Prichard and lower Wallace formations), intrusive dikes, and sills then other rock types (Libby Formation, Striped Peak Formation, and Ravalli group, and glacial deposits) during non-outbreak years within an area in the Panhandle National Forests of

Idaho and Washington. Southworth et al. (2002) studied how geology and geomorphology

influenced the ecology within the Great Smoky Mountains National Park and found that distinct

flora and fauna were found on debris flows on sulfidic slate, transported regolith exposing 3

bedrock, and on barren mafic and carbonate rocks. The third study examined how

geomorphology affected the genus Nebria (Coleoptera: Carabidae) (Gereben 1995). Gereben

(1995) found that the six Nebria species she studied were active in different areas of the glacier

retreat zone defined by geomorphological and ecological conditions. These studies may infer that the presence and activity of animals may or may not be related to the geology of an area.

The reasons for this may possibly be based on the life history, mobility, and specialization of a particular species.

Objectives

The main objectives of this study were to catalog the coleopteran fauna of a subalpine environment within the San Juan Mountains of Colorado and to investigate how the physical environment, specifically geology, influenced the species composition of the ground dwelling beetle (Coleoptera: Carabidae) community. None of the studies mentioned previously (Gereben

1995 and Garrison-Johnston et al. 2003) have analyzed how the geology of an area influenced beetle biodiversity and community structure. Ottensen (1996) mentioned that soil humidity, soil chemistry and nutrient status, substrate porosity, shadiness of habitat and altitude may be the main factors that contribute to the species distribution of Carabidae. Since the type of geologic bedrock can affect the soil chemistry, particle size, porosity, and to a lesser extent influence the amount of weathering, then geology should affect the carabid community.

Significance

1. This study will investigate the role of geology as a determinant of Carabidae biodiversity.

2. This study will improve our understanding of geological factors and animal distributions. 4

3. A new study on relating abiotic factors to Carabidae distribution in a mountainous system.

4. A first coleopteran biodiversity study in the San Juan Mountains.

General Hypotheses

Hypothesis 1: Geology influences the soil properties.

Hypothesis 2: Bedrock formation influences the community structure of Carabidae.

Hypothesis 3: Bedrock has a stronger effect on the Carabidae community and composition

than other environmental factors.

General Background

The Ground Dwelling Beetles. The ground dwelling beetles, Family Carabidae, were

chosen since they are common, live most of their lives in, or on, the ground, and the majority are

predators and scavengers. These traits should make them ideal candidates to see if geological

bedrock, sediment characteristics, and other physical parameters linked to the geology will have

effects on their habitat preferences. Carabids habitat preference for the ground was the main

reason I chose them. The geology of an area can affect multiple soil factors (see next section)

that have been shown to also influence carabid communities. Larvae and adults are found under

objects and the larvae burrow in the ground, creating tunnels (Pavuk pers. comm.). Adults will

burrow into the soil for finding prey, storing food, concealment, and hibernation, and their

activities within the soil can aid in drainage, aeration, and movement of nutrients (Evans 1991).

Majority of studies also point to soil characteristics (e.g., soil moisture content, pH, soil density,

sediment sizes) for their habitat preferences (e.g., Thiele 1977, Paje and Mossakowski 1984, 5

Eyre et al. 1990, Holmes et al. 1993, Desender et al. 1994, Holopainen et al. 1995, Sanderson et

al. 1995, Ottensen 1996, Hosoda 1999).

The other two reasons for using carabids are their abundance and their feeding ecology.

Since they are common, a large sample size can be taken to investigate geological affects. They are the third largest beetle family (40,000 species worldwide), are usually quite abundant in

ecosystems, and are found in nearly all terrestrial habitat types (foothills thru the alpine; tropical

to boreal forests; and in grasslands and tundra) (Thiele 1977 and Larochelle and Larviviere

2003). Ground beetles often are predatory or feed on dead or dying insects, but some can be

omnivorous, seed feeders, parasitic, or even phytophagous or mycophagous (Thiele 1977 and

Lovei and Sunderland 1996). If herbivorous beetles are used, they may be more influenced by

the presence of host plants rather than soil characteristics. Plants can also be very specialized to

the geology of an area, so to limit confounding effects, carabids were chosen.

Geology and Soil Characteristics. Bedrock can weather by mechanical (e.g., frost wedging)

and chemical (e.g., dissolution) forces. The rate of weathering depends on parent material,

mineral stability, climate, vegetation, topography, and time (Easterbrook 1999). The first two mentioned are based on the geologic material, and topography is based on the geological processes (e.g., mountain building via tectonics). The bedrock itself will affect the minerals found since the weathered material breaks up the rock and leaves behind the minerals, along with decayed organic matter, and windblown material deposited in the area. One of the main byproducts of weathering is soils.

The physical properties of a soil are affected by its mineralogy, texture of mineral grains, and organic matter (Montgomery 1997). The texture of soil and sediments are the shape of mineral grains, the size range present, amount of rock fragments in the material, and how the 6

grains fit together (Montgomery 1997). The texture influences drainage: porosity and permeability of the soil, which in turn can affect moisture content and humidity of a soil. Gravel tends to have low porosity and very high permeability, fine sand tends to have medium porosity and permeability, and clays have high porosity and very low permeability (Montgomery 1997).

The thickness of the soils depends on the bedrock, slope, and climate of an area (Boggs

1995). The development of soils is placed into two main categories, residual and transported.

Residual soils develop in place and the soil characteristics depend on the bedrock itself (e.g., granite, limestone, shales, and sandstones) (Rahn 1996). Transported soils are created from surficial deposits that have been moved and deposited (e.g., colluvium, alluvium, glacial drift, eolian, marine, and lacustrine deposits) (Rahn 1996). The main composition of a soil tends to be controlled by the bedrock and the way it is weathered (Montgomery 1997), but from studies of soil chemistry, the mineral composition and bulk chemical composition may differ greatly from bedrock (Boggs 1995). Igneous and metamorphic rocks tend to create unconsolidated materials whose surface and small particles are often quite different chemically and structurally, whereas sedimentary rocks and their soils are chemically and physically similar (Bohn et al. 1985).

Mountain Ecosystem Beetle Studies. Studies on beetle communities and abiotic factors in

mountainous environments are rare. Most of the studies have been conducted in Europe (e.g.,

Focarile 1986, Thingstad 1987, Gereben 1995, Ottensen 1996, , Hosoda, Kaufmann 2001, Faïek

2004, Loffler and Finch 2005, Lobo et al. 2007) with other studies in Central and South America

(Sota 1996), Asia (Hosoda 1999), South Africa (Botes et al. 2007), and Mexico (Lobo and

Halffter 2000). The majority of these studies focused on altitude as a main abiotic factor, with a

few that focused on soil factors. There are only a few studies in the United States relating 7

Coleoptera to abiotic factors in mountain environments (e.g., Rieske and Buss 2001, Jones 2004,

Apigian et al. 2006).

Colorado Rocky Mountain Beetle Studies. Most articles found for Colorado Coleoptera were surveys of beetles within the mountains (LeConte 1878 and 1879, and Schmoller 1971a and

1971b) or of Colorado in general (Wickham 1902, Kippenhan 1990, and Heffern 1998). A few studies have examined how altitude affects the distribution of Carabidae with the majority of the focus in Boulder County (Haubold 1951, Armin 1963, and Elias 1987). Most beetle studies performed in the mountains of Colorado are population-based and relate to Spruce beetle outbreaks (e.g., McCambridge and Knight 1972, Veblen et al. 1991, Roovers and Rebertus 1993,

Veblen et a. 1994, Kulakowski et al. 2003, Ryerson et al., 2003).

Only a few insect studies have been performed or surveyed in the San Juan Mountains.

Strecker (1878) identified insect specimens collected from a survey conducted by the Corps of

Engineers for the San Juan Mountain region in 1878. There are only a few locality collections in

San Juan County made by Rotger, a Reverend who collected 19,000 insect specimens in southern

Colorado, with 970 specimens from the family Carabidae (Elias 1987). The other insect studies performed in the San Juan Mountains relate to the endangered Uncompahgre fritillary butterfly

(Britten et al. 1994, Britten and Riley 1994) and on spruce beetle outbreaks (Ryerson et al.,

2003).

San Juan Mountains: The San Juan Mountains are known for their ecological, geological, hydrological, and climatological diversity, allowing relationships and interactions between these to be studied (Mountain Studies Institute 2006). A brief geological and ecological description follows. The geological units in the western San Juan Mountains span from the Precambrian to

Quaternary deposits, with bedrock layers from each main Era. The types of rocks range from 8

hard resistant igneous and metamorphic rocks, to alternating soft and hard sedimentary rocks

(Blair 1996). The landscape itself was formed from episodic uplift and deformation, late

Cretaceous plutonic activity (65 to 67 Ma), late Tertiary volcanic activity (35 to 22 Ma), and

multiple glaciations (last 2 My) and post glacial processes (Blair 1996).

The ecology of the western San Juan Mountains is just as diverse as the geology. There

are habitats and species found here that occur nowhere else in the world. For example, the San

Juan Mountains contain last known locations of certain arctic mosses, relics of the last ice age,

and rare alpine fens (Mountain Studies Institute 2006). The plant communities are placed into

three main categories: semi-arid basins and foothills, cool mountain communities, and riparian

communities. The semi-arid communities consist of the greasewood-shadscale shrub steppe and

the Great Basin sagebrush shrub steppe for the basin, and the foothills have pinon-juniper

woodland, mountain shrub community, and ponderosa pine-oak-Douglas fir forest (Blair 1996).

The cool mountain communities consist of the mid-slope vegetations: mixed conifer forests,

aspen groves, and mountain parks or meadows. The highest peaks and ridges have alpine

meadows and fellfields (Blair 1996). The riparian communities can be found along rivers, lakes,

and ponds, and also moist canyon bottoms and wetlands (Blair 1996).

Dissertation Overview

This manuscript is organized into four main chapters. Chapter One documents how

geology influenced the abiotic and biotic factors (shown in previous studies to affect carabids)

measured within the study area. Chapter Two examines how the geology influenced carabid

community structure and individual species. Chapter Three analyzes how the carabid

community was related to the environmental factors (studied in Chapter One), overall. Chapter 9

Four is a taxonomic survey of all the beetles identified so far in this study and how the

environmental factors affected beetle abundance. 10

References

Apigian, K.O., D.L. Dahlsten, and S.L. Stephens. 2006. Biodiversity of Coleoptera and the

importance of habitat structural features in a Sierra Nevada mixed-conifer forest.

Environmental Entomology. Vol. 35 (4): 964-975.

Armin, L.C. 1963. A study of the family Carabidae (Coleoptera) in Boulder County, Colorado.

Dissertation, University of Colorado.

Barr, G.E. and K.J. Babbitt. 2002. Effects of biotic and abiotic factors on the distribution and

abundance of larval two-lined salamanders (Eurycea bislineata) across spatial scales.

Oecologia. Vol. 133: 176-185.

Blair, R. 1996. The western San Juan Mountains: Their geology, ecology, and human history.

University Press of Colorado, Niwot, Colorado.

Boggs Jr., S. 1995. Principles of sedimentology and stratigraphy. Prentice Hall, Upper Saddle

River, New Jersey.

Bohn, H., B. McNeal, and G. O’Conner. 1985. Soil chemistry. John Wiley and Sons, New

York.

Botes, A. M.A. McGeoch, and S.L. Chown. 2007. Ground-dwelling beetle assemblages in the

northern Cape Floristic Region: Patterns, correlates and implications. Austral Ecology.

Vol. 32 (2): 210-224.

Britten, H.B., P.F. Brussard, and D.D. Murphy. 1994. The pending extinction of the

Uncompahgre fritillary butterfly. Conservation Biology. Vol. 8 (1): 86-94.

Britten, H.B. and L. Riley. 1994. Nectar source diversity as an indicator of habitat suitability

for the endangered Uncompahgre fritillary, Boloria acrocnema (Nymphalidae). Journal

of the Lepidopterists Society. Vol. 48 (3): 173-179. 11

Cole, M.M. 1986. The savannas: Biogeography and geobotany. London: Academic Press,

England.

Desender, K. 1994. Carabid beetles: Ecology and evolution. Kluwer Academic Publishers,

Boston MA.

Easterbrook, D.J. 1999. Surface processes and landforms. Prentice Hall, Upper Saddle River,

New Jersey.

Elias, S.A. 1987. Colorado ground beetles (Coleoptera: Carabidae) from the Rotger Collection,

University of Colorado Museum. Great Basin Naturalist. Vol. 47 (4): 631-637.

Evans, M.E.G. 1991. Ground beetles and the soil: their adaptations and environmental effects.

In The Environmental Impact of Burrowing Animals and Animal Burrows. Ed. by P.S.

Meadows and A. Meadows. Clarendon Press, Oxford England. 119-132.

Eyre, M.D., M.L. Luff, and S.P. Rushton. 1990. The ground beetle (Coleoptera, Carabidae)

fauna of intensively managed agricultural grasslands in northern England and southern

Scotland. Pedobiologia. Vol. 34 (1): 11-18.

Faïek, E., P. Jay-Robert, J. Lumaret, and O. Piau. 2004. Composition and structure of dung

beetle (Coleoptera: Aphodiidae, Geotrupidae, Scarabaeidae) assemblages in mountain

grasslands of the Southern Alps. Annals of the Entomological Society of America. Vol.

97 (4):701-709.

Focarile, A. 1986. Altitudinal zonation and structural characteristics of phytosaprobic Cenoses

of Coleoptera in the Upper Seriana Valley Northern Italy Lombardy Bergamo District.

Giornale Italiano di Entomologia. Vol. 3 (14): 229-256.

Forsyth, J.L. 2003. Linking Ohio geology and botany: Papers by Jane L. Forsyth. Ed. and pub.

by R.L. Stuckey, Ohio. 12

Friedman, J.M., W.R. Osterkamp, and W.M. Lewis Jr. 1996. The role of vegetation and

bed-level fluctuations in the process of channel narrowing. Geomorphology. Vol. 14:

341-351.

Garrison-Johnston, M.T., J.A. Moore, S.P. Cook, and G.J. Niehoff. 2003. Douglas-fir

beetle infestations are associated with certain rock and stand types in the inland

northwestern United States. Environmental Entomology. Vol. 32 (6): 1354-1363.

Gereben, B.A. 1995. Co-ocurrence and microhabitat distribution of six Nebria species

(Coleoptera: Carabidae) in an alpine glacier retreat zone in the Alps, Austria. Arctic and

Alpine Research. Vol. 27 (4): 371-379.

Grant, E.H.C., R.E. Jung, and K.C. Rice. 2005. Stream salamander species richness and

abundance in relation to environmental factors in Shenandoah National Park, Virginia.

The American Midland Naturalist. Vol. 153 (2): 348-356.

Haubold, V.L. 1951. Distribution of the Carabidae (Coleoptera) of Boulder County, Colorado.

American Midland Naturalist. Vol. 45 (No. 3): 683-710.

Heffern, D.J. 1998. A survey of the Cerambycidae (Coleoptera) or longhorned beetles of

Colorado (Insects of Western North America). Gillette Museum of Arthropod Diversity,

Dept. of Bioagricultural Sciences and Pest Management, Colorado State University, Fort

Collins Colorado.

Holmes, P.R., D.C. Boyce, and D.K. Reed. 1993. The ground beetle (Coleoptera: Carabidae)

fauna of Welsh peatland biotypes: Factors influencing the distribution of ground beetles

and conservation implications. Biological Conservation. Vol. 63 (2): 153-161. 13

Holopainen, J.K., T. Bergman, E.L. Hautala, and J. Oksanen. 1995. The ground beetle

fauna (Coleoptera: Carabidae) in relation to soil properties and foliar fluoride content in

spring cereals. Pedobiologia. Vol. 39 (3): 193-206.

Hosoda, H. 1999. Altitudinal occurrence of ground beetles (Coleoptera, Carabidae) on Mt.

Kurobi, Central Japan, with special reference to forest vegetation and soil characteristics.

Pedobiologia. Vol. 43 (4): 364-371.

Huggett, R. 1995. Geoecology: An evolutionary approach. Routledge, London England.

Jones, M. E., T.D. Paine, M.E. Fenn, and M.A. Poth. 2004. Influence of ozone and nitrogen

deposition on bark beetle activity under drought conditions. Forest Ecology and

Management. Vol. 200 (1-3): 67-76.

Kaufman, R. 2001. Invertebrate succession on an alpine glacier foreland. Ecology. Vol. 82

(8): 2261-2278.

Kippenhan, M.G. 1990. A survey of the tiger beetles (Coleoptera: Cicindelidae) of Colorado.

Entomological News. Vol. 101 (5): 307-315.

Kruckeberg, A.R. 2004. Geology and plant life: The effects of landforms and rock types on

plants. University of Washington Press, Seattle, Washington.

Kulakowski, D., T.T. Veblen, and P. Bebi. 2003. Effects of fire and spruce beetle outbreak

legacies on the disturbance regime of a subalpine forest in Colorado. Journal of

Biogeography. Vol. 30 (9): 1445-1456.

Larochelle, A. and M.C. Larviviere. 2003. A natural history of the ground beetles (Coleoptera:

Carabidae) of America north of Mexico. Pensoft Publishers, Sofia Bulgaria. 14

LeConte, J.L. 1878. The Coleoptera of the alpine regions of the Rocky Mountains. Bulletin of

the United States Geological and Geographical Survey of the Territories. Vol. 4 (2): 447-

480.

LeConte, J.L. 1879. The Coleoptera of the alpine Rocky Mountain regions - Part II. Bulletin

of the United States Geological and Geographical Survey of the Territories. Vol. 5: 499-

520.

Lobo, J.M. and G. Halffter. 2000. Biogeographical and ecological factors affecting the

altitudinal variation of mountainous communities of coprophagous beetles (Coleoptera:

Scarabaeoidea): a comparative study. Annals of the Entomological Society of America.

Vol. 93 (1): 115-126.

Lobo, J.M., E. Chehlarov, and B. Gueorguiev. 2007. Variation in dung beetle (Coleoptera:

Scarabaeoidea) assemblages with altitude in the Bulgarian Rhodopes Mountains: A

comparison. European Journal of Entomology. Vol. 104: 489-495.

Loffler, J., and O-D. Finch. 2005. Spatio-temporal gradients between high mountain

ecosystems of central Norway. Arctic, Antarctic, and Alpine Research. Vol. 37 (4): 499-

513.

Lovei, G.L. and K.D. Sunderland. 1996. Ecology and behavior of ground dwelling beetles

(Coleoptera: Carabidae). Annual Review of Entomology. Vol. 41: 213-256.

McCambridge, W.F. and F.B Knight. 1972. Factors affecting spruce beetles during a small outbreak. Ecology. Vol. 53 (5): 830-839.

Montgomery, C.W. 1997. Fundamentals of geology. Wm. C. Brown Publishers, Dubuque,

Iowa. 15

Moore, A.L., C.E. Williams, T.H. Martin, and W.J. Moriarity. 2001. Influence of season,

geomorphic surface, and cover item on capture, size, and weight of Desmognathus

ochrophaeus and Plethodon cinereus in Allegheny Plateau riparian forests. The

American Midland Naturalist. Vol. 145 (1): 39-45.

Mountain Studies Institute. Accessed January 31, 2006. Internet website:

www.mountainstudies.org/sanjuanmountains/default.htm

Osterkamp, W.R. and C.R. Hupp. 1984. Geomorphic and vegetative characteristics along

three northern Virginia streams. Geological Society of America Bulletin. Vol. 95: 1093-

1101.

Ottesen, P.S. 1996. Niche segregation of terrestrial alpine beetles (Coleoptera) in relation to

environmental gradients and phenology. Journal of Biogeography. Vol. 23: 353-369.

Paje, F. and D. Mossakowski. 1984. pH-preferences and habitat selection in carabid beetles.

Oecologia. Vol. 64: 41-46.

Rahn, P.H. 1996. Engineering geology: An environmental approach. Prentice Hall, Upper

Saddle River, New Jersey.

Read, R.A. 1952. Tree species occurrence as influenced by geology and soil on an Ozark north

slope. Ecology. Vol. 33 (2): 239-246.

Romans, R. 1981. Geobotany II. Plenum Publishing Corporation, New York.

Rieske, L.K. and L.J. Buss. 2001. Influence of site on diversity and abundance of ground- and

litter-dwelling Coleoptera in Appalachian Oak–Hickory forests. Environmental

Entomology. Vol. 30 (3): 484-494. 16

Roovers, L.M. and A.J. Rebertus. 1993. Stand dynamics and conservation of an old-growth

Engelmann spruce-subalpine fir forest in Colorado. Natural Areas Journal. Vol. 13 (4):

256-267.

Ryerson, D.E., T.W. Swetnam, and A.M. Lynch. 2003. A tree-ring reconstruction of western

spruce budworm outbreaks in the San Juan Mountains, Colorado, U.S.A. Canadian

Journal of Forest Research/Revue Canadienne de Recherche Forestiere. Vol. 33 (6):

1010-1028.

Sanderson, R.A., S.P. Rushton, A.J. Cherrill, and J.P. Byrne. 1995. Soil, vegetation, and

space: An analysis of their effects on the invertebrate communities of a Moorland in

northeast England. Journal of Applied Ecology. Vol. 32: 506-518.

Schmoller, R.R. 1971a. Habitats and zoogeography of alpine tundra Arachnida and Carabidae

(Coleoptera) in Colorado. The Southwestern Naturalist. Vol. 15 (3): 319-329.

Schmoller, R.R. 1971b. Nocturnal arthropods in the alpine tundra of Colorado. Arctic and

Alpine Research. Vol. 3 (4): 345-352.

Soto, T. 1996. Altitudinal variation in life cycles of carabid beetles: Life-cycle strategy and

colonization in alpine zones. Arctic and Alpine Research. Vol. 28 (4): 441-447.

Southworth, S., P. Chirico, D. Denenny, A. Schultz, J. Triplett, and J. Young. 2002. The

role of geology in the ecology of the Appalachian Blue Ridge. Abstracts with Programs-

Geological Society of America. Vol. 34 (6): 546.

Strecker, H. 1878. Section II: Lepidoptera. In Annual Report upon Explorations and Surveys

in the Department of the Missouri being Appendix SS of the Annual Report of the Chief

of Engineers for 1878, by E.H. Ruffner. 1847-1867. 17

Thiele, H.U. 1977. Carabid beetles in their environments: A study on habitat selection by

adaptation in physiology and behaviour. Springer-Verlag, New York.

Thingstad, P.G. 1987. Pitfall trapping of the carabid fauna in alpine and subalpine habitats at

Saltfjellet North Norway. Fauna Norvegica Series B. Vol. 32 (2): 105-116.

Tomans, R.C. 1977. Geobotany I. Plenum Publishing Corporation, New York.

Veblen, T.T., K.S. Hadley, S.M. Reid and A.J. Rebertus. 1991. The response of subalpine

forests to spruce beetle outbreak in Colorado USA. Ecology (Washington D C). Vol. 72

(1): 213-231.

Veblen, T.T., K.S. Hadley, E.M. Nel, T. Kitzberger, M. Reid, and R. Villalba. 1994.

Disturbance regime and disturbance interactions in a Rocky Mountain subalpine forest.

Journal of Ecology. Vol. 82 (1): 125-135.

Wentworth, T.R. 1981. Vegetation on limestone and granite in the Mule Mountains, Arizona.

Ecology. Vol. 62 (2): 469-482.

Whittaker, R.H. 1954. The ecology of serpentine soils. Ecology. Vol. 35 (2): 258-288.

Wickman, H.F. 1902. The Coleoptera of Colorado. Bulletin of the State University of Iowa.

No. 5 (3): 217-310.

18

CHAPTER I

The Influence of Geology on the Environmental Factors Related to Carabidae Distribution

The family Carabidae is composed of the ground dwelling beetles. These beetles tend to

make their living on the ground underneath rocks, logs, detritus or in the soil creating burrows for hunting prey, storing food, and hibernation (Evans 1991). Ottensen (1996) mentions that soil humidity, soil chemistry and nutrient status, substrate porosity, shadiness of habitat and altitude

may be the main factors that contribute to carabid species distribution. In Ottensen’s study, soil

water content was most influential in Carabidae community species composition, though species

differences changed on a continuum rather than communities being distinct. In other studies,

several abiotic and biotic factors (e.g., soil pH, plant structure and composition, soil density,

sediment sizes, geomorphology, soil temperature) have been shown to play a role in habitat

preferences (Thiele 1977, Paje and Mossakowski 1984, Eyre et al. 1990, Quinn et al. 1991,

Holmes et al. 1993, Desender et al. 1994, Gereben 1995, Holopainen et al. 1995, Sanderson et al.

1995, Hosoda 1999, Ings and Hartley 1999, Magura et al. 2003, XiaoDong et al. 2003, Magura et

al. 2005, Long and Medina 2006). Several of these factors (e.g., soil moisture, pH, chemistry,

porosity, nutrients, soil density, plant composition, sediment composition) were influenced by

geology and the geologic history of an area in past studies at other localities (Read 1952,

Whittaker 1954, Plaster and Sherwood 1971, Tomans 1977, Romans 1981, Wentworth 1981,

Bohn et al. 1985, Huggett 1995, Rahn 1996, Montgomery 1997, Burke 2002, Forsyth 2003,

Kruckeberg 2004, Neatrour et al. 2006, Neff et al. 2006, Courjault-Radé 2007). The objective of this study was to determine whether the underlying geology influences any of the environmental 19

factors that have been known to affect Carabidae, especially the soil properties above each

formation.

Materials and Methods

Site Location. The field site was located in the San Juan Mountains of Colorado, several

kilometers south of Silverton, a small mining town in southwest Colorado (Fig. 1). The Sultan

Creek-Molas Lake region was chosen for its several bedrock outcroppings that are mostly

parallel to each other and occur at similar elevations at around 3,170 m (10,400 ft). The study

area itself is an open meadow with remnants of the spruce-fir forest, just south of Sultan Creek

and approximately one kilometer north of Molas Lake on the east side of Highway 550 (Fig. 2).

The open meadow borders the heavily forested area that lies on the Precambrian bedrock within

the Animas River watershed. The entire study area was about 400 m X 400 m. Five sites were

chosen within each formation, with three general formations used. Each site was 25 m2. Sites

were at last 50 m apart from each other and ran in a north-south direction (Fig. 2).

Geology. The bedrock formations chosen for this study were Molas, Ouray-Leadville, and

Elbert, all sedimentary rocks. The Elbert formation is the oldest (Devonian Period) and

composed of sandstone (quartz), siltstone, and paper thin shales. The Ouray-Leadville is two

formations lumped together due to similar rock type (carbonates made up of limestone and

dolomite). They are mainly composed of limestone and dolomite, with minor shale and

sandstones and localized cherty areas. Both limestone and dolomite are made up of the minerals

calcite and dolomite (CaCO3 and CaMg(CO3)2, respectively) which are structurally similar to

each other. The Ouray Formation was formed in the Devonian Period and the Leadville

Formation in the Mississippian Period. The Molas Formation is the youngest (Pennsylvanian) 20

and is composed of shale, siltstone, and conglomerate. It is believed to be an ancient soil

(paleosol). These formations were deposited on top of each other during the Paleozoic Era.

Uplifting of the region caused these bedrock units to be tipped at an angle (≈ 10-25° W) and

formations striking at an N-S direction. Erosion and glaciation followed. Presently, the area is

made up of ridges, with a layer-cake topography of the bedrock units due to its geological

history. Rich et. al’s (unpublished) geologic map was used to set up this research project.

Abiotic Variables. Multiple physical factors of the soils were studied over each bedrock

formation. Soil temperature, pH, moisture, compressive strength, and sediment size distribution

were gathered at each site. These variables were measured to see how much these factors vary

within and between soils developed over different bedrock substrates. In theory, soil and

sediment above each bedrock formation should be different from each other in terms of these

factors if they are different in mineralogy and rock type, especially in a low topographic area.

These factors would most likely cause residual soils to develop. The Sultan Creek-Molas lake

region is low in topographic differences (42 m difference between lowest and highest site), and is

made up of sedimentary rocks, making this area a prime candidate for residual soils and there

with having characteristics similar to the bedrock from which it was weathered (Rahn 1996).

Soil temperature was measured with an HBE International Bi-Metal Dial (accuracy of

1%, or ± 1.2° C) each week in the morning on a Wednesday in 2006 and 2007 (eight weeks and

six weeks total, respectively). In 2006, measurements were taken between 8:30am to noon and

generally took 2 - 2.5 hrs to collect. In 2007, measurements were taken between 8am-10:30am.

The author had help from a field assistant and the time it took to collect the data was 1 - 1.5 hrs.

Temperature was taken in a specific route, site five to one, site six to ten and site fifteen to

eleven or vice versa. This was reversed to make sure temperature at most sites did not 21

correspond to the order temperature was taken. Site eight was the only site that had the same

corresponding number in the routes. Three temperature readings were taken at each site.

The moisture content (% relative saturation) at each site was measured with the Kelway

Soil Acidity and Moisture Tester (accuracy of ± 10%) in situ each week simultaneously with

temperature. Three moisture readings were taken at each site.

Compressive strength and pH were taken during weeks one, five, and eight in 2006 and

once midway through the season in 2007. Compressive strength was measured with a

penetrometer (accuracy of 0.25 kg/sq cm) and the pH was measured with the Kelway instrument

(accuracy of ± 0.2). Three readings were taken per site.

Sediment samples were taken at each site in 2006 and sorted with sieves to calculate sediment size distribution, mean grain size, and type of sorting. Standard phi units (Φ) were used in sieving, ranging from -2 Φ to +4 Φ and a few rocks were measured by hand (Table 1).

Weight (g) of sediments was calculated for each size class (sediment found in each individual sieve) and was transformed into a percent (size class weight divided by total sample weight).

The sediment distribution was then plotted on a cumulative weight curve graph (Fig. 3). The equations used for graphic sediment mean and sediment sorting (the standard deviation) are shown below (Boggs 1995). The numbers in the equations represent percents where the phi

(size) was recorded from the cumulative curve. Extreme measurements (e.g., Φ95) were

estimated by extending the line on the cumulative graph to reach 0% and 100%.

Graphic Sediment Mean = Φ16 + Φ50 + Φ84 3

Sediment Sorting (Standard Deviation) = Φ84 – Φ16 + Φ95 – Φ5 4 6.6 22

Biotic Variables. Percent plant cover, percent detritus cover, average plant height, and dominant plant species richness were measured three times throughout the season during weeks one, five, and eight in 2006 and once midway through the trapping season in 2007. Percent plant

cover, percent detritus cover, and average plant height were measured by dividing a site into four

quadrants and calculating the average for the entire site. Cover was estimated visually and

height of plants was measured with a ruler by visually finding a plant representing the average

height in each quadrant. Plant species were identified by the author in the field or specimens

were collected for later identification. Identification was performed with field guides (Carter

1988, Kershaw et al. 1998, Guennel 2004, and Dahms 2005), an internet website (Southwestern

Colorado Wildflowers, Ferns, and Trees), and by Joanna Lemly (Wetland Ecologist, Colorado

Natural Heritage Program, Colorado State University) and Kerry Cutler (botanist fen technician from CSU during 2006 field season). A small sample of litter detritus was collected near each

site midway through the trapping season in 2007 (45 cm X 45 cm area). The detritus was dried

and weighed in the laboratory to calculate the amount deposited at each site.

Geographical Variables. Latitude and Longitude, elevation and aspect were measured with a

GPS unit for each site. The slope was estimated for each site with a Brunton compass and board.

Statistical Analyses. One way ANOVAs were performed on the 2006 and 2007 abiotic and

biotic variables against bedrock formations to find significant differences between formations.

One way ANOVAs were performed on separated weekly collections, combined weekly data for overall trends, and on data collected just once in a season. If an ANOVA produced a significant

P-value a Tukey test was performed to find which formations were different from each other.

Soil temperature data was further analyzed by running linear regressions on the relative time

pattern soil temperature was taken against the soil temperature itself to see if route by which 23

measurements were recorded affected the resulting data. Plant species between bedrock

formations were analyzed by species richness and the Jaccard Coefficient similarity index. The

Jaccard Coefficient was calculated to see how similar or different plant communities were on different formations.

Jaccard Coefficient: J = a/(a+b+c)

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 = 1 most similar, J = 0 least similar

Results

Abiotic Variables. Soil moisture during week two in 2006 was significantly different between formations (F = 5.42, df = 2, 12, P = 0.021). The Elbert Formation had the lowest moisture content whereas the Molas had the highest moisture content (Fig. 4). Overall differences were not significant in 2006. In 2007, no significant differences weekly or overall were found for bedrock and soil moisture.

Overall trends for soil temperature in both years showed significant differences between bedrock formations with Elbert having the lowest temperature (F = 3.32, df = 2, 312, P = 0.037 in 2006 and F = 12.59, df = 2, 267, P = <0.001 in 2007) (Figs. 5 and 6). In 2006, all weeks except week three soil temperatures were significantly different between formations (Table 2).

In 2006, depending on the order temperature was taken, either the Molas Formation or the Elbert

Formation had the lowest soil temperature (Figs. 7 and 8 gives an example of each route 24

direction taken and soil temperature outcome). In 2007, during weeks one and five, soil

temperature was significantly different between formations (Table 3). The Elbert Formation had

the lowest temperature whereas the Molas Formation had the highest (Figs. 9 and 10). Linear

regressions were performed on the relative time readings soil temperatures were taken for each

route during both years (Figs. 11 and 12). Three of the four routes showed a significantly

positive relationship for temperature during the time soil was taken (Table 4). The route not

affected in 2007 (weeks two, four, and six) by time showed no significant differences in

temperature between formations.

Sediment analyses, compressive strength, and pH were not significantly different

between bedrock formations in 2006 and 2007 (Table 2 and 3).

Biotic Variables. Detritus cover was significantly different between bedrock formations in 2006

and 2007 (Tables 2 and 3). The overall trend in 2006 showed all bedrock units were

significantly different from each other (F = 17.94, df = 2, 42, P = < 0.001) whereas in 2007

Elbert and Ouray-Leadville were significantly different from each other (Figs. 13 and 14). The

Elbert Formation was always the lowest in detritus cover in both years.

An overall total of 34 plant species were found with 29 species in 2006 and 32 species in

2007 (Table 5). The Ouray-Leadville Formations had the highest overall plant species richness

with 22 species whereas the Molas Formation had the lowest with 18 species (Table 5). The

Jaccard Index generally showed low plant species similarity between the formations, typically

about one third of the species shared except in 2007, the Elbert and Ouray-Leadville became

more similar, sharing 50% of the plant species composition (Table 6). Overall, formations

compared to Molas have one-third of the species in common, whereas Elbert and Ouray- 25

Leadville share 43% of their species. Only 15% of plant species are shared (seven species)

across all formations when taking the whole study area into account.

Plant cover, plant height, plant species richness, and detritus amount were not

significantly different between formations (Tables 2 and 3).

Geographical Variables. Elevation was significantly different between bedrock formations (F

= 49.97, df = 2, 12, P = < 0.001). Elbert Formation has the lowest elevation, whereas Molas has

the highest elevation (Fig. 15). Aspect and slope were not significantly different between

bedrock formations (Tables 2 and 3).

Discussion

Abiotic Variables. Temperature was significantly different between bedrock formations on a

continuous basis. However, this was reflected in the order in which temperature was taken

instead of bedrock formation. In 2006, the bedrock the author started in tended to be the lowest

temperature compared to the last unit. When factoring in the sequence the sites were measured,

the 2006 routes were linearly significant with soil temperature. In, 2007, with the help of a field

assistant, time it took to gather the soil temperature data was cut in half and was gathered in a

more consistent basis. Even so, this also resulted in most weeks the first bedrock measured was

lowest and one of the two routes was linearly significant with time sequence. The only

exception was in 2007, during weeks two and six, the Ouray-Leadville was the lowest in

temperature, though not significantly different. In conclusion, this variable most likely was not

affected by geology but by sequence of collection. It is interesting to point out when all weeks

are combined regardless of the route taken the Elbert Formation was always significantly lowest 26

in temperature. This may have been by chance or one of the other factors took part in this

formation being overall cooler (e.g., plant cover, topography).

Soil Moisture content was the second abiotic factor that showed a significant difference

between bedrock formations. Overall, significance was very weak since there was only one

week where significance was observed. This occurred during the driest part of the season in

2006. When P-values between dry and wet seasons were compared, there was no distinction in these values. Sediment size and sorting were very similar between units, so this may have played a role in the moisture not being significantly different. Differences in sediment sizes and type of sorting are well known to influence water infiltration and retention due to porosity and

permeability (Bengt-Owe 1967 and Montgomery 1997). Other factors may also have had an

influence. Aspect, slope, or even plant structural factors might have controlled moisture

differences (Breshears and Barnes 1999, Qiu et al. 2001, Svetlitchnyi et al. 2003). Aspect, slope

and plant cover in this study showed a diverse range within formations and overlap in type of

aspect, slope, and plant cover between formations.

Sediment analyses, soil compressive strength, and pH were not significantly different

between bedrock formations. The sediment analysis resulted in a mean of fine sand size and

very poorly sorted material for each formation, with a mode of silt/clay (sediments found in pan).

Similar grain size most likely has to do with each formation sharing similar rock types. The

Molas Formation and Elbert Formation share similar rock types: shales, siltstones, and possibly

sandstones (conflicting references for Molas if it has sandstones). Ouray-Leadville is mainly a

mix of limestone and dolomite, with minor shale and sandstones and localized cherty areas.

Limestones and dolomites typically have small grain sizes when weathered, although they can

also have a range of grain sizes: clay, silt, and sand particles due to impurities from pure 27

carbonate rock. Soil developed over carbonate rocks is the insoluble portion of the original rock

(quartz, iron and manganese oxides and clay minerals) and the weathered products form a

residual soil that is usually clayey but occasionally silty and sandy (Tugrul and Zarif 2000).

These diverse ranges of rock types mentioned within each formation may have resulted in my

sites having a mix of grain sizes (poorly sorted) and a similar mean (fine sand size).

The pH was acidic in all formations with a mean range of 5.7 - 6.7 for sites. The pH was

unexpected for the Ouray-Leadville Formations. Limestones and dolomites tend to be basic due

to the chemical structure (carbonate), yet the average of the formations was acidic, ranging from

6.11 - 6.43. It is possible that the elements that cause alkalinity have been leached out, but the

soils tend to be shallow and near bedrock outcroppings. Hydrothermal alteration, mining, and

coniferous trees in the area may be a more logical explanation affecting the soil pH. The rocks

in this region have been re-crystallized due to hydrothermal activity which led to secondary

mineralization of acidic minerals (Fetchenhier 1996, Sares and Gleason 2000, and Church et al.

2007). Mining activity also exposes and releases heavy metals that have been shown to acidify water/sediments in the Animas River watershed (Besser and Leib 2007, Nimick et al. 1999, and Church et al. 2007). The study area is near mining activity (Fetchenhier 1996) with abandoned small mines scattered in the Molas Lake region. Finally, conifers produce acidic soil conditions and they have

been shown to alter soil pH (Madeira and Ribeiro 1995, Alfredsson 1998, Binkley and Giardina

1998, Ste-Marie and Paré 1999). Since this meadow is surrounded by coniferous trees and was

until recently a covered forest (during the late 1870’s a fire swept through the area creating the

open meadows), the pine needles, lignin, etc. may have influenced the pH of the study area.

Other factors may have also influenced the soil measurements. Soil development processes or stages have been known to affect soil factors instead of parent material. Shure and

Ragsdale (1977) showed trends or significant differences in soil temperature, moisture, cation 28

activity, amount of organic matter, and soil density within different stages of soil development

on granite outcrops. Sites within each of my formations had different levels of soil development,

with very shallow soil depth that were adjacent to bedrock outcroppings, to deeper, more

organically rich developed soils. Soil depth and development of sites were not studied but

should be quantified in future studies to see if this has an effect on soil factors studied.

Finally, trees have been known to influence soil factors ranging from how the soil was

developed to affecting acidity (see above), organic content, weathering, soil temperature and

water content (Binkley and Giardina 1998). Even though my site was an open meadow, it had

trees scattered throughout and some sites were next to small remnant patches of the forest. Also,

due to the fire history of the area, there may be lag effects. Finally, if trees have been known to

affect soil properties, the herbaceous layer might also affect these properties.

Biotic Variables. Detritus cover was significantly different between formations in both years.

This may have to do with the chemical composition of these formations, plant structure, the type

of detritus within the formations, or detritivore activity or a combination of these and other

factors interacting with each other (e.g., Lang and Orndorff 1983, Beloin et al. 1988, Spehn et al.

2000, Bohlen et al. 2001, Pouyat and Carreiro 2003, Neatrour et al. 2006). The type of detritus

was observed but not quantified. The Elbert Formation detritus was mostly grass and had

smaller quantities of cones, moss, and woody debris. The Ouray-Leadville Formations had

grass, moss, pine needles, then cones, woody debris and a large rotting stump. Detritus of the

Molas Formation consisted of needles, cones, grass, moss, and twigs (at one site). Depending on

the type of detritus, this might have to do with how fast breakdown occurs due to the chemistry

of the litter (e.g., lignin, cellulose, nitrogen dynamics; Pouyat and Carreiro 2003). The chemical

and detritivore compositions were not studied, but should be examined in future studies. The 29

bedrock and its properties might be affecting the microbial and/or invertebrate fauna that break

down litter. Finally, pH has been correlated with the breakdown of detritus in other studies

(Neatrour et al. 2006). Even though the pH was not significantly different between formations, it still may have had an influence on detritus cover and its breakdown.

The only plant related factor observed to have a difference between bedrock was community composition. Only seven species were shared across all three formations specifically, Achillea alpicola, Astragalus alpines, Potentilla hippiana, Potentilla pulcherrima,

Pseudocymopteris montanus, Solidago simplex var nana, and Taraxacum sp. The Molas

Formation was the most distinct when compared to the other formations. This difference was not statistically tested, but most formations during the two seasons tended to only share one-third of the species composition. Differences in plant communities between bedrock were not surprising since most studies done previously at other locations have shown correlations with specific species and/or distinct plant communities based on geology, geomorphology, and geologic history (e.g., Read 1952, Wentworth 1981, Osterkamp and Hupp 1984, Forsyth 2003, and

Kruckeberg 2004).

Geographical Variables. The geographic variables were recorded to check for equivalent range of slopes and aspects for a biodiversity study (Chapters 2 through 4) that was not significantly different between bedrock formations to make sure there were few other confounding factors to separate from bedrock effects. Elevation, though, was significantly different between formations. This significance was most likely caused by the depositional sequence of the formations; Molas is youngest and highest in elevation whereas Elbert is oldest and lowest in elevation.

30

Conclusion

In conclusion, formations were significantly different from each other in terms of

elevation, detritus cover, and soil temperature. The Elbert Formation generally had the lowest

elevation, detritus cover, and soil temperature. The Molas Formation generally had the highest

elevation, temperature, and detritus in 2006, but medium in 2007. The Ouray-Leadville

Formations generally had the medium elevation, soil temperature, and detritus in 2006 but higher

in 2007. Most of these differences were likely associated with the geology and geologic history

of the area, except for soil temperature, which was most likely caused by sequence of the soil

measurements. Plant community composition also appeared distinct between formations and

should be studied in more depth.

Other factors that tend to be associated with geology that were not significantly different

may have to do with using only sedimentary rocks or other factors (e.g., vegetation effects, or rate of soil development). A few of these factors, such as soil pH and sediment size and sorting, although not significantly different between formations, were most likely affected by the geology

(e.g., hydrothermal alteration or sharing a range of rocks with different sized grains like shale and siltstones).

The carabid fauna at this locality may not be closely tied to bedrock since most of the soil variables did not show any differences between formations. However, in past studies, carabids have been known to be influenced by detritus cover and elevation. Carabids can also be influenced by other factors that may be associated with geology, but were not studied here (e.g.,

amount of clay content and calcium content of the soil).

Future studies should incorporate the rest of the meadow and a few other localities for

these units. These formations continue southward in the meadow reaching Molas Creek. They 31

re-appear (as forested sites) in the Lime Creek region between Highway 550 and Old Lime

Creek Road and on a much smaller scale at Coal Bank Pass off of Highway 550. The significant

factors can be studied in other locations to see if they are repeated on a larger geographical area.

Other units are also in the vicinity: metamorphic and igneous rocks (gneiss, quartzite, schist, slate, granite, igneous intrusion), other sedimentary rocks, and glacial deposits. Differences in soil properties and possibly other environmental factors may be occurring at a different scale

than formation, from localized bedding planes (e.g., shale, sandstone, conglomerate) to general rock types (igneous, metamorphic, and sedimentary). 32

References

Alfredsson, H., L.M. Condron, M. Clarholm and M.R. Davis. 1998. Changes in soil acidity

and organic matter following the establishment of conifers on former grassland in New

Zealand. Forest Ecology and Management. Vol. 112 (3): 245-252.

Beloin, R.M., J.L. Sinclair, and W.C. Ghiorse. 1988. Distribution and activity of

microorganisms in subsurface sediments of a pristine study site in Oklahoma. Microbial

Ecology. Vol. 16: 85-97.

Bengt-Owe, J. 1967. The significance of grain size and pore water content for the interstitial

fauna of sandy beaches. Oikos. Vol. 18 (2): 311-322.

Besser, J.M., and K.J. Leib. 2007. Toxicity of metals in water and sediment to aquatic biota.

In Integrated investigations of environmental effects of historical mining in the Animas

River watershed, San Juan County, Colorado. Eds. S.E. Church, P. von Guerard, and

S.E. Finger. U.S. Geological Survey Professional Paper 1651: 837–849.

Binkley, D. and C. Giardina. 1998. Why do tree species affect soils? The warp and woof of

tree-soil interactions. Biogeochemistry. Vol. 42: 89-106.

Boggs Jr., S. 1995. Principles of sedimentology and stratigraphy. Prentice Hall, Upper Saddle

River, New Jersey.

Bohlen, P.J., P.M. Groffman, C.T. Driscoll, T.J. Fahey, T.G. Siccama. 2001. Plant–soil–

microbial interactions in a northern hardwood forest. Ecology. Vol. 82 (4): 965-978.

Bohn, H., B. McNeal, and G. O’Conner. 1985. Soil chemistry. John Wiley and Sons, New

York. 33

Breshears, D.D. and F. J. Barnes. 1999. Interrelationships between plant functional types and

soil moisture heterogeneity for semiarid landscapes within the grassland/forest

continuum: a unified conceptual model. Landscape Ecology. Vol. 14: 465–478.

Burke, A. 2002. Properties of soil pockets on arid Nama Karoo inselbergs-the effect of geology

and derived landforms. Journal of Arid Environments. Vol. 50 (2): 219-234.

Carter, J.L. 1988. Trees and shrubs of Colorado. Johnson Books, Boulder CO.

Church, S.E., J.R. Owen, P. von Guerard, P.L. Verplanck, B.A. Kimball, and D.B. Yager.

2007. The effects of acidic mine drainage from historical mines in the Animas River

watershed, San Juan County, Colorado: What is being done and what can be done to

improve water quality? In Understanding and Responding to Hazardous Substances at

Mine Sites in the Western United States. Ed. DeGraff, J.V. Geological Society of

America Reviews in Engineering Geology. Vol. XVII: 47–83.

Courjault-Radé P., M. Munoz, N. Hirissou and E. Maire. 2007. Geology, key factor for high

quality wine production: An example from the Gaillac Appellation Region (Tarn, SW

France). XXXth World Congress of Wine, Budapest 10-16, June.

Dahms, D. 2005. Rocky Mountain wildflowers. Paragon Press, Windsor, CO.

Desender, K. 1994. Carabid beetles: Ecology and evolution. Kluwer Academic Publishers,

Boston MA.

Eyre, M.D., M.L. Luff, and S.P. Rushton. 1990. The ground beetle (Coleoptera, Carabidae)

fauna of intensively managed agricultural grasslands in northern England and southern

Scotland. Pedobiologia. Vol. 34 (1): 11-18. 34

Evans, M.E.G. 1991. Ground beetles and the soil: their adaptations and environmental effects.

In The Environmental Impact of Burrowing Animals and Animal Burrows. Ed. by P.S.

Meadows and A. Meadows. Clarendon Press, Oxford England. 119-132.

Fetchenhier, S. 1996. CH 7: Ore deposits and minerals. In The western San Juan Mountains:

Their geology, ecology, and human history. Ed. R. Blair. University Press of Colorado,

Niwot, Colorado. 80-95.

Forsyth, J.L. 2003. Linking Ohio geology and botany: Papers by Jane L. Forsyth. Ed. and pub.

by R.L. Stuckey, Ohio.

Gereben, B.A. 1995. Co-ocurrence and microhabitat distribution of six Nebria species

(Coleoptera: Carabidae) in an alpine glacier retreat zone in the Alps, Austria. Arctic and

Alpine Research. Vol. 27 (4): 371-379.

Guennel, G.K. 2004. Guide to Colorado wildflowers. Vol. 2: Mountains. Westcliffe

Publishers, Englewood, CO.

Holmes, P.R., D.C. Boyce, and D.K. Reed. 1993. The ground beetle (Coleoptera: Carabidae)

fauna of Welsh peatland biotypes: Factors influencing the distribution of ground beetles

and conservation implications. Biological Conservation. Vol. 63 (2): 153-161.

Holopainen, J.K., T. Bergman, E.L. Hautala, and J. Oksanen. 1995. The ground beetle

fauna (Coleoptera: Carabidae) in relation to soil properties and foliar fluoride content in

spring cereals. Pedobiologia. Vol. 39 (3): 193-206.

Hosoda, H. 1999. Altitudinal occurrence of ground beetles (Coleoptera, Carabidae) on Mt.

Kurobi, Central Japan, with special reference to forest vegetation and soil characteristics.

Pedobiologia. Vol. 43 (4): 364-371.

Huggett, R. 1995. Geoecology: An evolutionary approach. Routledge, London England. 35

Ings, T.C. and S.E. Hartley. 1999. The effect of habitat structure on carabid communities

during the regeneration of a native Scottish forest. Forest Ecology and Management.

Vol. 119: 122-136.

Kershaw, L., A. MacKinnon, and J. Pojar. 1998. Plants of the Rocky Mountains. Lone Pine

Publishing, Auburn, WA.

Kruckeberg, A.R. 2004. Geology and plant life: The effects of landforms and rock types on

plants. University of Washington Press, Seattle, Washington.

Lang, G.E. and K.A. Orndorff. 1983. Surface litter, soil organic matter, and the chemistry of

mineral soil and foliar tissue: Landscape patterns in forests located on mountainous

terrain in West Virginia. Proceedings, 4th Central Hardwood Conference. University of

Kentucky, Lexington, Kentucky. 303–321.

Long, J.W. and A.L. Medina. 2006. Consequences of ignoring geologic variation in

evaluating grazing impacts. Rangeland Ecology and Management. Vol. 59: 372-382.

Madeira, M. and C. Ribeiro 1995. Influence of leaf litter type on the chemical evolution of a

soil parent material (sandstone). Biogeochemistry. Vol. 29 (1): 43-58.

Magura, T., B. Tothmeresz, and Z. Elek. 2003. Diversity and composition of carabids during

a forestry cycle. Biodiversity and Conservation. Vol. 12: 73–85.

Magura, T., B. Tothmeresz, and Z. Elek. 2005. Impacts of leaf litter addition on carabids in a

conifer plantation. Biodiversity and Conservation. Vol. 14: 475-491.

Montgomery, C.W. 1997. Fundamentals of geology. Wm. C. Brown Publishers, Dubuque,

Iowa.

Neatrour, M.A., M. Lynch, A.G. Sammarco, and R.L. Fuller. 2006. The role of local

geology and soil properties in mediating the effects of episodic acidification on leaf 36

breakdown in stream ecosystems of the Adirondack region, New York, USA. NABS

Annual Meeting, Anchorage, Alaska.

Neff, J. C., R. Reynolds, R.L. Sandford Jr., D. Fernandez, and P. Lamothe. 2006. Controls

of bedrock geochemistry on soil and plant nutrients in southeastern Utah. Ecosystems.

Vol. 9: 879–893.

Nimick, D.A., S.E. Church, T.E. Cleasby, D.L. Fey, B.A. Kimball, K.J. Leib, A.M. Mast,

and W.G. Wright. 1999. Characterization of metals in water and bed sediment in two

watersheds affected by historical mining in Montana and Colorado. In U.S. Geological

Survey Toxic Substances Hydrology Program—Proceedings of the Technical Meeting,

Charleston, South Carolina, March 8-12, 1999: Vol. 1-3. Eds. D.W. Morganwalp, and

H.T. Buxton. Contamination from Hardrock Mining: U.S. Geological Survey Water-

Resources Investigations Report 99-4018A: 11-18.

Osterkamp, W.R. and C.R. Hupp. 1984. Geomorphic and vegetative characteristics along

three northern Virginia streams. Geological Society of America Bulletin. Vol. 95: 1093-

1101.

Ottesen, P.S. 1996. Niche segregation of terrestrial alpine beetles (Coleoptera) in relation to

environmental gradients and phenology. Journal of Biogeography. Vol. 23: 353-369.

Paje, F. and D. Mossakowski. 1984. pH-preferences and habitat selection in carabid beetles.

Oecologia. Vol. 64: 41-46.

Plaster, R.W. and W.C. Sherwood. 1971. Bedrock weathering and residual soil formation in

central Virginia. GSA Bulletin. Vol. 82 (10): 2813-2826.

Pouyat, R.V. and M.M. Carreiro 2003. Controls on mass loss and nitrogen dynamics of oak

leaf litter along an urban-rural land-use gradient. Oecologia. Vol. 135: 288–298. 37

Qiu, Y., B. Fu, J. Wang, and L. Chen. 2001. Spatial variability of soil moisture content and

its relation to environmental indices in a semi-arid gully catchment of the Loess Plateau,

China. Journal of Arid Environments. Vol. 49: 723–750.

Quinn, M.A., R.L. Kepner, D.D. Walgenbach, R.N. Foster, R.A. Bohls, P.D. Pooler, K.C.

Reuter, and J.L. Swain. 1991. Effect of habitat characteristics and perturbation from

insecticides on the community dynamics of ground beetles (Coleoptera: Carabidae) on

mixed grass rangeland. Environmental Entomology. Vol. 20 (5): 1285-1294.

Rahn, P.H. 1996. Engineering geology: An environmental approach. Prentice Hall, Upper

Saddle River, New Jersey.

Read, R.A. 1952. Tree species occurrence as influenced by geology and soil on an Ozark north

slope. Ecology. Vol. 33 (2): 239-246.

Rich, C., L. Walters, D. Stewart, R. Anderhalt, C. Onasch, D. May, and J. Evans.

Unpublished. Geologic map of the Molas Lake-Sultan Creek area, San Juan County,

Colorado. Bowling Green State University, Geology Department, Bowling Green, Ohio.

Romans, R. 1981. Geobotany II. Plenum Publishing Corporation, New York.

Sanderson, R.A., S.P. Rushton, A.J. Cherrill, and J.P. Byrne. 1995. Soil, vegetation, and

space: An analysis of their effects on the invertebrate communities of a Moorland in

northeast England. Journal of Applied Ecology. Vol. 32: 506-518.

Sares, M.A., and J.A. Gleason. 2000. Geology, Water Quality, and Avalanche Hazards of

the Ouray – Silverton Area, Southwest Colorado: Earth Science Week Field Trip October

9, 2000. Colorado Geological Society, Denver Colorado.

Shure, D.J. and H.L. Ragsdale. 1977. Patterns of primary succession on granite outcrop

surfaces. Ecology. Vol. 58 (5): 993-1006. 38

Southwestern Colorado Wildflowers, Ferns, and Trees. http://www.swcoloradowildflowers.com/

Spehn, E.M., J. Joshi, B. Schmid, J. Alphei, and C. Körner. 2000. Plant diversity effects on

soil heterotrophic activity in experimental grassland ecosystems. Plant and Soil. Vol.

224: 217–230.

Ste-Marie, C. and D. Paré. 1999. Soil, pH and N availability effects on net nitrification in the

forest floors of a range of boreal forest stands. Soil Biology and Biochemistry. Vol. 31

(11): 1579-1589.

Svetlitchnyi, A.A., S.V. Plotnitskiy, and O.Y. Stepovaya. 2003. Spatial distribution of soil

moisture content within catchments and its modeling on the basis of topographic data.

Journal of Hydrology. Vol. 277: 50–60.

Thiele, H.U. 1977. Carabid beetles in their environments: A study on habitat selection by

adaptation in physiology and behaviour. Springer-Verlag, New York.

Tomans, R.C. 1977. Geobotany I. Plenum Publishing Corporation, New York.

Tugrul, A. and I.H. Zarif. 2000. Engineering aspects of limestone weathering in Istanbul,

Turkey. Bulletin of Engineering Geology and the Environment. Vol. 58: 191–206.

Wentworth, T.R. 1981. Vegetation on limestone and granite in the Mule Mountains, Arizona.

Ecology. Vol. 62 (2): 469-482.

Whittaker, R.H. 1954. The ecology of serpentine soils. Ecology. Vol. 35 (2): 258-288.

XiaoDong, Y., L. TianHong, and Z. HongZhang. 2003. Species diversity of litter-layer

beetles in the Fengtongzhai National Nature Reserve, Sichuan Province. Acta

Entomologica Sinica. Vol. 46 (5): 609-616. 39

Table 1. Sediment size ranges measured for soil analyses.

Phi Actual Size Size Class -4 Φ to -2 Φ ≥ 4mm to 16mm pebble -1 Φ 2mm granule 0 Φ 1mm very coarse sand +1 Φ 0.5mm coarse sand +2 Φ 0.25mm medium sand +3 Φ 0.125mm fine sand +4 Φ 0.0625mm very fine sand pan < 0.0625mm silt & clay

40

Table 2. One-way ANOVAs for all variables against bedrock formation taken in 2006.

Variable P df F Abiotic Sediment Size 0.584 2,12 0.56 Sediment Sorting 0.331 2,12 1.22 Soil Compressive Strength W1 0.952 2,12 0.05 Soil Compressive Strength W5 0.812 2,12 0.21 Soil Compressive Strength W8 0.590 2,12 0.55 Soil Moisture W1 0.312 2,12 1.29 Soil Moisture W2 0.021 2,12 5.42 Soil Moisture W3 0.869 2,12 0.14 Soil Moisture W4 0.403 2,12 0.98 Soil Moisture W5 0.821 2,12 0.20 Soil Moisture W6 0.993 2,12 0.01 Soil Moisture W7 0.873 2,12 0.14 Soil Moisture W8 0.868 2,12 0.14 Soil pH W1 (dry season) 0.654 2,12 0.44 Soil pH W5+W8 (wet season) 0.483 2,12 0.77 Soil Temperature W1 0.025 2,12 5.11 Soil Temperature W2 0.022 2,12 5.29 Soil Temperature W3 0.116 2,12 2.60 Soil Temperature W4 0.006 2,12 7.99 Soil Temperature W5 0.004 2,12 8.88 Soil Temperature W6 0.002 2,12 11.68 Soil Temperature W7 0.000 2,12 36.18 Soil Temperature W8 0.002 2,12 11.00 Biotic Detritus Cover W1 0.003 2,12 9.61 Detritus Cover W5 0.033 2,12 4.59 Detritus Cover W8 0.063 2,12 3.52 Plant Cover W1 0.997 2,12 0.00 Plant Cover W5 0.984 2,12 0.02 Plant Cover W8 0.999 2,12 0.00 Plant Height W1 0.966 2,12 0.03 Plant Height W5 0.080 2,12 3.13 Plant Height W8 0.201 2,12 1.84 Plant Species Richness 0.241 2,12 1.61 Geographical Aspect 0.799 2,12 0.23 Elevation 0.000 2,12 49.97 Slope 0.796 2,12 0.23

41

Table 3. One-way ANOVAS for all variables against bedrock formation taken in 2007.

Variable P F df Abiotic Soil Compressive Strength 0.612 0.51 2,12 Soil Moisture W1 0.223 1.71 2,12 Soil Moisture W2 0.997 0.00 2,12 Soil Moisture W3 0.995 0.05 2,12 Soil Moisture W4 0.480 0.78 2,12 Soil Moisture W5 0.889 0.12 2,12 Soil Moisture W6 0.811 0.21 2,12 Soil pH 0.486 0.77 2,12 Soil Temperature W1 0.030 4.76 2,12 Soil Temperature W2 0.807 0.22 2,12 Soil Temperature W3 0.111 2.65 2,12 Soil Temperature W4 0.711 0.35 2,12 Soil Temperature W5 0.005 8.38 2,12 Soil Temperature W6 0.945 0.06 2,12 Biotic Detritus Cover 0.035 4.51 2,12 Detritus Weight 0.383 1.04 2,12 Plant Cover 0.978 0.02 2,12 Plant Height 0.306 1.31 2,12 Plant Species Richness 0.752 0.29 2,12

Table 4. P-values for linear regressions run on soil temperature and the relative time it

was taken. Route 1: site five to site eleven; Route 2: site eleven to site five.

Year Route P df F 2006 1 0.004 1,13 11.91 2006 2 0.000 1,13 35.82 2007 1 0.003 1,13 13.36 2007 2 0.668 1,13 0.17

42

Table 5. Overall plant species found in each formation.

Plant Species Elbert Ouray-Leadville Molas 1 Achillea alpicola X X X 2 Anemone multifida globosa X 3 Astragalus alpinus X X X 4 Blepharoneuron tricholepis X 5 Bromus anomalus or rhizomitous X 6 Castilleja occidentalis X 7 Danthonia intermedia X 8 Delphinium glaucum or barbeyi X 9 Elymus trachycaulus X X 10 Ergsimum capitatum X 11 Erigeron simplex or leiomerus X X 12 Erigeron ursinus X X 13 Festuca saximontana X X 14 Festuca thurberi X X 15 Fragaria virginiana X X 16 Geranium richardsonii X 17 Juniperus communis X 18 mushrooms (Fungi) X 19 Packera diphormophylla X X 20 Pentaphylloides floribunda X X 21 Phlox condensata X X 22 Picea engelmannii? X 23 Poa sp. X X 24 Potentilla hippiana X X X 25 Potentilla pulcherrima X X X 26 Pseudocymopteris montanus X X X 27 Ranunculus eschscholtzii X 28 Solidago simplex var nana X X X 29 Taraxacum sp. X X X 30 Thalictrum fendleri or sparsiflorum X X 31 Trifolium sp. X X 32 Trisetum spicatum X 33 Vaccinum scoparium X X 34 Zigadenus venenosus X Species Richness 21 22 18

43

Table 6. Jaccard Coefficients for plant community structure between bedrock formations.

Formations 2006 2007 Combined Elbert + Ouray-Leadville 0.33 0.50 0.43 Elbert + Molas 0.38 0.29 0.34 Ouray-Leadville + Molas 0.32 0.37 0.33 All Formations 0.15 0.17 0.15

44

A)

B)

Fig. 1. General map of Colorado where the San Juan Mountains are (A) and a close up

map of the study area and Silverton (B). Silverton is located about 5km north of the study

area (orange dot). 45

Fig. 2. Topographic map and aerial photograph of the study area and site locations.

46

100 90 80 70 60 50 40 30 20

Cumulative Weight Percent Cumulative Weight 10 0 -5 Φ -4 Φ -2 Φ -1 Φ 0 Φ +1 Φ +2 Φ +3 Φ +4 Φ pan Grain Size

Fig. 3. Cumulative curve example for Site 9. 47

40 b 35 30

25 ab 20 a 15 10 5 Mean Percent Soil Moisture Mean Percent 0 Elbert Ouray-Leadville Molas

Fig. 4. Mean (± SEM) soil moisture in bedrock formations during week two, 2006.

25

20 b a ab 15

10

5 Mean Soil Temperature (°C) Temperature Mean Soil 0 Elbert Ouray-Leadville Molas

Fig. 5. Overall mean (± SEM) soil temperature in bedrock formations, 2006. 48

20 18 c 16 b 14 a 12 10 8 6 4

Mean Soil Temperature (°C) Mean Soil Temperature 2 0 Elbert Ouray-Leadville Molas

Fig. 6. Overall mean (± SEM) soil temperature in bedrock formations, 2007.

25.00 a 20.00 b c 15.00

10.00

5.00 Mean Soil Temperature (°C) Temperature Mean Soil 0.00 Elbert Ouray-Leadville Molas

Fig. 7. Mean (± SEM) soil temperature in bedrock formations, week four, 2006. Route:

site eleven to fifteen, site ten to six and site one to five.

49

25 b 20 b

15 a

10

5 Mean Soil Temperature (°C) Temperature Mean Soil 0 Elbert Ouray-Leadville Molas

Fig. 8. Mean (± SEM) soil temperature in bedrock formations, week six, 2006. Route: site

five to one, site six to ten and site fifteen to eleven.

20 18 b 16 ab 14 a 12 10 8 6 4

Mean Soil Temperature (°C) Mean Soil Temperature 2 0 Elbert Ouray-Leadville Molas

Fig. 9. Mean (± SEM) soil temperature in bedrock formations, week one, 2007. 50

20

18 b 16 ab 14 12 a 10 8 6 4

Mean Soil Temperature (°C) Mean Soil Temperature 2 0 Elbert Ouray-Leadville Molas

Fig. 10. Mean (± SEM) soil temperature in bedrock formations, week five, 2007.

30 y = 0.6093x + 12.651 25 R² = 0.4781 20

15 y = 0.3994x + 13.605 R² = 0.7337 10

Soil Temperature C° Soil Temperature 5

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Time

Fig. 11. Linear regressions for soil temperature and the relative time it was measured,

2006. Blue represents route: site five to site eleven, and red represents route: site eleven to site five. 51

25 y = 0.4687x + 10.78 20 R² = 0.5069 15 y = 0.0478x + 13.577 10 R² = 0.0128

5 Soil Temperature C° Soil Temperature

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Time

Fig. 12. Linear regressions for soil temperature and the relative time it was measured,

2007. Blue represents route: site five to site eleven, and red represents route: site eleven to site five.

40 c 35 30 25 b 20 15 10 5 a Mean Percent Detritus Cover Mean Percent 0 Elbert Ouray-Leadville Molas

Fig. 13. Mean (± SEM) detritus cover in bedrock formations, 2006. 52

50 b ab 45 40 35 30 25 a 20 15 10

Mean Percent Detritus Cover Mean 5 0 Elbert Ouray-Leadville Molas

Fig. 14. Mean (± SEM) detritus cover in bedrock formations, 2007.

3200 c b 3180 a 3160

3140

Mean Elevation (m) 3120

3100 Elbert Ouray-Leadville Molas

Fig. 15. Mean elevation of bedrock formations (mean ± SEM).

53

CHAPTER II

The Impact of Bedrock Formations on Carabidae (Coleoptera) Community Structure

Studies on how abiotic factors affect beetle communities in mountainous environments are rare in the literature. Most of the studies have been conducted in Europe, with other studies in Central and South America, Asia, South Africa, and Mexico (e.g., Focarile 1986, Thingstad

1987, Gereben 1995, Ottensen 1996, Sota 1996, Hosoda 1999, Lobo and Halffter 2000, Rüdiger

2001, Faïek 2004, Loffler and Finch 2005, Botes et al. 2007, Lobo et al. 2007). Most of these focused on altitude as a main abiotic factor, with a few studies that focused on soil factors and one on geomorphology. There were only a few studies in the United States for Coleoptera in mountain environments that related beetle occurrence to abiotic factors (e.g., Rieske and Buss

2001, Jones 2004, Apigian et al. 2006). None of the studies have examined how the geology of an area influenced beetle communities. Only one study on a specific population of an insect was performed that included geology (Garrison-Johnston et al. 2003) and another study that included how glacier geomorphology affected a specific beetle genus, Nebria (Coleoptera: Carabidae)

(Gereben 1995).

Most studies that have examined Coleoptera in Colorado were surveys within the mountains or of Colorado in general (Strecker 1878, LeConte 1878 and 1879, Wickham 1902,

Schmoller 1971a and 1971b, Kippenhan 1990, Heffern 1998). Other studies performed in

Colorado were based on populations of individual species which most dealt with Spruce beetle outbreaks (e.g., McCambridge and Knight, 1972, Veblen et al. 1991, Roovers and Rebertus

1993, Veblen et al. 1994, Kulakowski et al. 2003, Ryerson et al. 2003). Carabidae studies were 54 scattered through time and only certain localities have been intensely examined, specifically

Boulder County (Haubold 1951, Armin 1963, and Elias 1987). These studies mainly examined altitudinal effects on the distribution of Carabidae. Within the San Juan Mountains, only a few locality collections have been made (Strecker 1878, Wickham 1902, Elias 1987) for the family

Carabidae.

This study had two main objectives: First, it examined the influence geology had on the carabid community structure; and second, a preliminary assessment was performed to catalogue carabid species occurring in San Juan County. Carabids were used for the geological analysis since they live their life mostly in, or on, the ground, and as adults burrow into the soil for prey acquisition, food storage, concealment, and hibernation (Evans 1991). Carabid communities have been shown to be influenced mostly by soil properties (e.g., Thiele 1977, Paje and

Mossakowski 1984, Eyre et al. 1990, Holmes et al. 1993, Holopainen et al. 1995,Sanderson et al.

1995, and Ottensen 1996) and most soil properties tend to be influenced by geology (e.g., Plaster and Sherwood 1971, Bohn et al. 1985, Huggett 1995, Rahn 1996, Montgomery 1997, Burke

2002, Neff et al. 2006, Courjault-Radé 2007).

Materials and Methods

Site Location. The study area was in the San Juan Mountains of Colorado, several kilometers south of Silverton, a small mining town in southwest Colorado (Fig. 1). The Sultan Creek-Molas

Lake region was chosen for its several bedrock outcroppings that are mostly parallel to each other and occur at similar elevations at around 3,170 m (10,400 ft). The study area itself is an open meadow with remnants of the spruce-fir forest, just south of Sultan Creek and about a kilometer north of Molas Lake on the east side along Highway 550 (Fig. 2). The open meadow 55 borders the heavily forested area that lies on the Precambrian bedrock, within the Animas River watershed. The entire study area was about 400 m X 400 m.

Geology. The bedrock formations chosen for this study were Molas, Ouray-Leadville, and

Elbert, all sedimentary rocks. The Elbert formation is the oldest (Devonian Period) and composed of sandstone (quartz), siltstone, and paper thin shales. The Ouray-Leadville is two formations lumped together due to similar rock type (carbonates made up of limestone and dolomite). They are mainly composed of limestone and dolomite, with minor shale and sandstones and localized cherty areas. Both limestone and dolomite are made up of the minerals calcite and dolomite (CaCO3 and CaMg(CO3)2, respectively) which are structurally similar to

each other. The Ouray Formation was formed in the Devonian Period and the Leadville

Formation in the Mississippian Period. The Molas Formation is the youngest (Pennsylvanian)

and composed of shale, siltstone, and conglomerate. It is believed to be an ancient soil

(paleosol). These formations were deposited on top of each other during the Paleozoic Era.

Uplifting of the region caused these bedrock units to be tipped at an angle (≈ 10-25° W) and

formations striking at an N-S direction. Erosion and glaciation followed. Presently, the area is

made up of ridges, with a layer-cake topography of the bedrock units due to its geological

history.

Carabidae Methodology. Pitfall trapping was used to collect the insects. Pitfall trapping can

introduce biases from spacing of traps, plant structure effects, activity and density of a particular

species, body size, type of preservative, and trap avoidance (Greenslade 1964, Weeks Jr. and

McIntyre 1997, Lang 2000, and Ward et al. 2001). If abundance data is only analyzed, this is

actually more of a measure of activity density of a species instead of true abundance differences

between species. A very active predator searching for prey will be caught more often than a sit 56 and wait predator and the size of a population in a given area will also affect catches (Greenslade

1964 and Lang 2000). Small sized carabids may also be able to avoid traps by recovering their balance and escaping while on the lip of a trap (Greensdale 1964). Even with these downfalls, pitfall trapping provides a simple and inexpensive method that is most reliable in collecting a high number and wide range of ground dwelling invertebrates (Thiele 1977 and Ward et al.

2001).

A few of these biases were avoided for the most part in my study. Traps were open for an entire week, allowing collection of both diurnal and nocturnal carabids. The trap spacing used for this study (see below) was between two of the common spacings (1 m and 5 m) typically used in past studies (Ward et al. 2001). The 1 m and 5 m spacing of traps that Ward et al. (2001) used did not affect overall coleopteran abundance and species composition, but did affect species richness (1 m significantly lower than 5 m; 24 mean species compared to 29.7 mean species, respectively). The preservative used (see below) has been shown to collect a wide variety of species. Also, multiple quantitative indices besides abundance were used (see

Quantitative and Statistical Analyses section) to analyze the community. However, even with these measures, the species caught in this study does not likely represent the entire carabid fauna.

Nine pitfall traps were placed per site in a 5 m X 5 m grid (Fig. 3 and 4). A total of fifteen sites were set up. The sites were set up into three transects with five sites each. Each site was at least 50 m apart from another site, and 30 m from the contact zones of the bedrock formations and highway (exception for one site, Site 15, which was about 20m from highway and 25 m from contact zone due to a minor wetland interfering along the transect). Trapping was done each week for eight weeks total in 2006 (June 17 to August 11) and six weeks total in 2007

(June 24 to August 3). Traps were kept open for 72 - 120 hours. A 50:50 solution of propylene 57 glycol:water was used to kill and preserve insects. Propylene glycol has been shown to be quite effective at capturing a high number of species and wide variety of ground dwelling arthropods

(when compared to water and live traps) and is less toxic to non-target wildlife (Weeks Jr. and

McIntyre 1997). A small amount of dish soap was added into the solution to lower surface tension. Quinine was also added to deter mammals from eating the insects. Traps were filled to the 75 - 100 ml mark in a 250 ml cup (Corning Snap-Seal no. 1730 design; Fig. 5). The cup dimensions were 7.5 cm height and 7.6 cm diameter. Cups were left with no covering for most of the study period in 2006. During the last few weeks, site six had to be covered with rocks due to juvenile marmots digging up the cups and emptying the contents out. In 2007, traps were left with no covering throughout the summer, but some of the cups had chicken wire over them to prevent them from popping out of the ground. Traps were cleaned out and insects stored in 70% ethanol alcohol. Carabidae were identified by the author using museum specimens at Colorado

State University, Fort Collins and The University of Colorado, Boulder; by carabid specialists

Bob Davidson (Carnegie Museum of Natural History) and Foster Purrington (The Ohio State

University); and by using the Harvard Insect Collection website.

Quantitative and Statistical Analyses. Community structure for each sites and between formations was measured using species richness, abundance, frequency, similarity and species composition of community (McIntyre et al. 2001). Frequency was the number of times (0 - 14, total weekly appearances) a species appeared at a given site. Shannon-Weiner index (H’) was calculated to measure the overall biodiversity of sites and bedrock formations, taking into account species composition and relative proportion of each species. The higher H’ was, the greater the diversity was. Evenness (E) measured how similar the abundances of the species were within a site/formation. When evenness equals 1, all species abundances were the same, 58

H’ = –∑ pi ln (pi)

pi = proportion of species i to total abundance

E = H’/ln S

H’ = Shannon-Weiner index

S = species richness

whereas the closer the number was to zero, the less even (equal) a site was (mix of very abundant, and very rare species). ANOVAs were performed on these values and on the most abundant/frequent species between bedrock formations. If an ANOVA produced a significant P- value a Tukey test was performed to find which specific formations were significantly different from each other.

Relative abundance and frequency were calculated for each species per formation. This was performed to find the relative proportions of dominant species in each formation and to assess if there were any compositional differences between formations in dominant species. The

Jaccard Coefficient was used to find out if the carabids separated out into distinct communities

based on presence/absence only data for bedrocks formations.

Jaccard Coefficient: J = a/(a+b+c)

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 = 1 most similar, J = 0 least similar 59

A cluster analysis was performed using PAST software (Hammer et al. 2001) on the Jaccard coefficients to visually see how sites were connected to each other. The paired group algorithm, or UPGMA, was used. An analysis of similarity (ANOSIM) using Primer-E software (Clark and

Gorley 2006) was performed on the Jaccard Coefficients to see if the sites significantly clustered out based on the geology. Global R was calculated in ANOSIM which has a range of -1 to +1.

A zero represented samples identical to each other, +1 indicated samples were maximally different, and a negative value suggested samples between groups (e.g., bedrock in this study) were more similar to each other then within groups.

Results

Tables one and two summarize the abundance and frequency, respectively, of each carabid species found in each site. A total of 1,236 individuals were collected, representing 30 species. Abundance of individual species ranged from 1 to 561. Abundance at sites ranged from

19 individuals (site 11) to 253 individuals (site 12) and species richness ranged from 8 (sites 1,

12, and 13) to 15 (site 4) (Table 3). Total frequency of individual carabids throughout the study area ranged from 1 to 114. Total frequency at sites ranged from 13 (site 11) to 57 (site 9) (Table

3). Table three also summarizes Shannon-Weiner and evenness for each site. Site three had both the highest and most consistent biodiversity (H’ = 2.18 and E = 0.85) whereas site 12 had the lowest biodiversity and evenness, 0.63 and 0.30 respectively. At site 12, there was a high emergence of Amara quenseli in 2006 and 2007 and accounted for 168 of the 205 individuals and 38 of the 48 individuals collected, respectively, which lowered the evenness.

Species richness of the carabids was significantly different between formations (Table 4).

The Molas Formation was significantly lower, with a mean of nine species, than the Ouray- 60

Leadville Formation with a mean of 12 species (Fig. 6). The Molas Formation also had the lowest mean carabid frequency, Shannon-Weiner, and evenness but had the highest mean abundance, though not significantly different from the other formations (Fig. 7). The highest values were equal or switched between Elbert and Ouray-Leadville, depending on which measure of the carabid community was analyzed. Table five represents the absolute measurements of the carabid community between formations. Again, the Molas Formation was overall the lowest in all values except for abundance.

Of the most common carabids caught, Amara quenseli and Harpalus nigritarsi were the most abundant and frequent species throughout the study area (Tables 1 and 2). Fourteen of the species found were considered common (≥ 10 individuals). Seven species were caught only once throughout the study (Table 1). The Rhadine species may possibly be a new species.

Harpalus opacipennis and Harpalus ellipsis may be separated out correctly or may be mixed or may be only one of these species (pers. comm. R. Davidson).

ANOVAs run on the most abundant carabid species against formation can be found in

Table six. Cymindis cribricollis and Harpalus animosus were significantly different between bedrock formations. Cymindis cribricollis was significantly higher in the Ouray-Leadville then the Elbert Formation, whereas Harpalus animosus was significantly higher in the Elbert and

Ouray-Leadville when compared to the Molas Formation (Fig. 8). Harpalus animosus was completely absent from the Molas Formation. Other species that were absent from one of the formations (though not significantly) were Agonum placidum (Molas), Bembidion mutatum

(Ouray-Leadville), Harpalus laticeps (Molas) (Fig. 8).

ANOVAs were also run on the frequency of the most common species (Table 7) to reduce the effects of abundance (and potential influences of emergences) and see if 61 presence/absence over time was affected by bedrock. Carabus taedatus, Cymindis cribricollis,

Harpalus animosus, and Harpalus laticeps were significantly different between formations. The

Molas Formation was significantly lower or completely absent in frequency when compared to the other formations within these species except for Cymindis cribricollis in which the Elbert

Formation was significantly lower in frequency than the Ouray-Leadville Formation (Fig. 9).

Table eight represents the total relative abundance of the carabids found in each formation. Ninety percent of the carabid abundance was composed of eight, seven, and five species for Elbert, Ouray-Leadville, and Molas Formations, respectively (Table 8). The most dominant abundant species in all three formations was Amara quenseli, ranging from 30 - 60% of the total species composition. Three other species, Harpalus nigritarsis, Calathus ingratus and Carabus taedatus were also found in the top 90% of the fauna in all formations. Four carabid species in the top 90% were unique to a specific formation: Agonum placidum,

Bembidion dauricum, Cymindis cribricollis, and Bembidion mutatum. Overall, majority of the main composition of the formations shared similar dominant species.

Relative frequency of the carabids resulted in ten species in each formation dominating the composition with Amara quenseli and Harpalus nigritarsis the top two most frequent carabids (Table 9). The main species that composed the most abundance in each formation were also found composing the top 90% of the frequency, but in slightly different order. Using frequency drowned out the high emergence effects and added a few more species in the top 90% to each formation. The majority of the main species composition was still shared among the formations. Unique species in the top dominant 90% were mostly different from before:

Agonum placidum, Amara ellipsis, Harpalus opacipennis, and Notiophilus aquaticus. 62

Table ten shows the Jaccard coefficient of similarity for sites and table seven between bedrock formations. Similarity between individual sites ranged from 0.27 (sites 1 and 10) to

0.69 (sites 1 and 9). Overall similarity between bedrock formations ranged from 0.41 to 0.62

(Table 11). Thirty percent of carabids were shared (11 species) throughout the entire study area.

Table 12 represents the results for the analysis of similarity for bedrock units. Global R was

-0.419 which indicated sites between formations were more similar to each other then sites within formations. The carabids also did not significantly separate out into distinct communities based on presence/absence data due to bedrock (Table 12, Fig. 10). Sites separated out into three main groups (five branches if viewed in finer detail) based on the cluster analysis (Fig. 11).

Certain sites within bedrock formations clustered together (e.g., in the Molas Formation three of the five sites cluster together in one of the branches) but most sites, especially in the Elbert and

Ouray-Leadville, mixed together.

Discussion

The only community measurement that was significant between formations was species richness. Species richness was significantly lower in the Molas Formation compared to Ouray-

Leadville. This may be due to properties of the bedrock itself or possibly other reasons (e.g., road effects). The Molas Formation had the highest elevation, temperature, and detritus cover

(Chapter One), overall highest grain size but overall the lowest in plant species and soil pH (not significantly different from other formations). The higher mean grain size may have to do with the Molas Formation having fewer species than the Elbert and Ouray-Leadville Formations.

Higher species richness of carabids was caught in soils with finer sediment (with higher clay content) then coarser sediment in past studies (Thiele 1977 and Holopainen 1995). The clay 63 content percent was not measured due to time constraints (12 hours to gather raw data of one sample with hydrometer analysis technique), but should be studied at a later date.

The Molas Formation also parallels the road but sites were not adjacent to the road (20-

30m away). Fields and forests adjacent to roads have mixed affects on invertebrate and carabid species richness: increased species richness (Koivula 2005), decreased species richness (Haskell

2000), or mixed results or no significant changes between road and adjacent habitat (Vermeulen

1993 and Varchola and Dunn 1999). Sites in the Molas Formation were not considered adjacent, since they were 20-30m away. Koivula (2005) showed a significant difference in species richness just 25m away from the road compared to the adjacent road site in a boreal forest.

Future studies though should be included to make sure possible road affects are not influencing species richness. The Molas Formation appears in areas not parallel to the highway which can be sampled. Also, sites can be set up right next to the highway to sample carabids and compare to the original site locations.

Six individual species (Agonum placidum, Bembidion mutatum, Carabus taedatus,

Cymindis cribricollis, Harpalus animosus, and Harpalus laticeps) affiliated themselves with two bedrock formations and were either completely absent from the other formation (Molas or

Ouray-Leadville) or significantly lower in abundance/frequency in that formation. These species tend to be habitat generalists (alpine, subalpine, montane), but prefer to affiliate with open ground, meadows, sandy and/or gravelly soil/pits, dry to moderately moist soil, and usually sparse vegetation (Larochelle and Larviviere 2003). Geology may be playing a role in their habitat selection. Most of these species were found in the Elbert and Ouray-Leadville

Formations, which were made up of mostly fine sandy soils. Also, this area has exposed outcrops 64 with little to no soil development, allowing the area to stay more open with patches of sparse to no vegetation.

Bembidion mutatum was completely absent in the Ouray-Leadville but found in the

Elbert and Molas. This might be from trapping problems since this beetle is quite small or there might be something in the Ouray-Leadville Formations that was causing it to not be found. This species was found frequent enough in the 2006 season (trapped 7 out of 8 weeks and found in 7 sites) that trapping problems were most likely not the reason. Its habitat choices are moderate moisture in fine sandy soil, organic debris, mossy, and/or sparse vegetation (Larochelle and

Larviviere 2003). The Ouray-Leadville had sites that this species affiliates with: moderate moisture, moss, detritus, and low vegetation cover. These formations were different in type of rock compared to the other two formations, dolomite and limestone, which produce limy soils.

The chemistry of the soil may be inhibiting Bembidion mutatum. Future trapping and analyses of the geology that were not performed could possibly shed some light on the habitat preferences of this species.

Dominant species composition between bedrock were similar between formations, especially the top two species (Amara quenseli and Harpalus nigritarsis) and the general 90% species composition was also similar between formations in both abundance and frequency.

Four species (30% of the frequency composition) in the top 90% in the Ouray-Leadville

Formation were Harpalus species (Table 9). Quite a few Harpalus species in past studies have been known to be “Limestone” species in Swedish habitats due to the microclimate the limestone creates (Thiele 1977). Larochelle and Larviviere (2003) mentions nothing about the species found in this study affiliating themselves with limestone, but since a large number of limestone- loving species are Harpalus (Thiele 1977), it is possible my species may be a part of this group. 65

More research needs to be performed on these particular species to find out if they are affiliated with limestone.

The carabids did not separate out into distinct communities on formations based on species composition. There were three general groupings of the sites based on carabid species composition. Other variables (biotic and or abiotic) might be controlling how the carabid assemblages separated out in the study area.

There were only a few other carabid studies in Colorado that incorporated the subalpine zone. Five species (Haubold 1951) to 38 species (Armin 1963) were found in the subalpine in

Boulder County, Colorado. In the Rotger’s collection that Elias (1987) studied, he found 41 species in the subalpine zone collected over a large geographic location in Colorado. Thirty species were counted in this current study. The study area was a smaller area when compared to the previous studies, but yielded a count not too different from these. Widening the study to the whole county and incorporating other collecting techniques may yield more carabid species.

This study may have produced possibly new species localities for this county and a new species or two. When comparing other studies within San Juan County and Silverton locations, possibly eight species were already collected here (Elias 1987 and Wickham 1902). An in depth study of collections where material have not been published may yield more information if the other 20+ species will be new records or have been collected previously. The Rhadine species may be new since this genus is quite speciose (pers. comm. R. Davidson). It will need to be sent to taxonomists with collections that contain Rhadine species to compare to other specimens already identified.

66

Conclusion

Individual species were mainly affected by the bedrock formation they were on. These significant effects may possibly be due to the geology itself or other factors mentioned previously. The community structure and species composition, on the other hand, were mainly not affected by geology, except species richness between formations. Future studies need to be performed at other locations where the formations are exposed to see if these trends are found in other geographical areas. Also, specific species analyses need to be performed to find out which factor(s) control a species presence in specific formations (geological aspect or environmental).

There are other bedrock units in this area that should be added, too. This study used only sedimentary rocks. Igneous and metamorphic rocks are in this region too, along with glacial deposits. Perhaps community and species compositional differences occur based on other geological factors (e.g., general rock type: igneous, metamorphic, sedimentary), geomorphic, or environmental factors. This was also the first systematic carabid study performed in San Juan

County. Future surveys for Carabidae should incorporate the rest of the meadow and the rest of the county and include other habitats to compare to the meadow. These surveys will help obtain an overall picture of ground beetle communities living in these mountains. 67

References

Apigian, K.O., D.L. Dahlsten, and S.L. Stephens. 2006. Biodiversity of Coleoptera and the

importance of habitat structural features in a Sierra Nevada mixed-conifer forest.

Environmental Entomology. Vol. 35 (4): 964-975.

Armin, L.C. 1963. A study of the family Carabidae (Coleoptera) in Boulder County, Colorado.

Dissertation, University of Colorado.

Bohn, H., B. McNeal, and G. O’Conner. 1985. Soil chemistry. John Wiley and Sons, New

York.

Botes, A. M.A. McGeoch, and S.L. Chown. 2007. Ground-dwelling beetle assemblages in the

northern Cape Floristic Region: Patterns, correlates and implications. Austral Ecology.

Vol. 32 (2): 210-224.

Burke, A. 2002. Properties of soil pockets on arid Nama Karoo inselbergs-the effect of geology

and derived landforms. Journal of Arid Environments. Vol. 50 (2): 219-234.

Clarke, K.R. and R.N. Gorley. 2006. PRIMER v6: User Manual/Tutorial. PRIMER-E,

Plymouth.

Courjault-Radé P., M. Munoz, N. Hirissou and E. Maire. 2007. Geology, key factor for high

quality wine producton: An example from the Gaillac Appellation Region (Tarn, SW

France). XXXth World Congress of Wine, Budapest 10-16, June.

Elias, S.A. 1987. Colorado ground beetles (Coleoptera: Carabidae) from the Rotger Collection,

University of Colorado Museum. Great Basin Naturalist. Vol. 47 (4): 631-637.

Evans, M.E.G. 1991. Ground beetles and the soil: their adaptations and environmental effects.

In The environmental impact of burrowing animals and animal burrows. Ed. by P.S.

Meadows and A. Meadows. Clarendon Press, Oxford England. 119-132. 68

Eyre, M.D., M.L. Luff, and S.P. Rushton. 1990. The ground beetle (Coleoptera, Carabidae)

fauna of intensively managed agricultural grasslands in northern England and southern

Scotland. Pedobiologia. Vol. 34 (1): 11-18.

Faïek, E., P. Jay-Robert, J. Lumaret, and O. Piau. 2004. Composition and structure of dung

beetle (Coleoptera: Aphodiidae, Geotrupidae, Scarabaeidae) assemblages in mountain

grasslands of the Southern Alps. Annals of the Entomological Society of America. Vol.

97 (4):701-709.

Focarile, A. 1986. Altitudinal zonation and structural characteristics of phytosaprobic Cenoses

of Coleoptera in the Upper Seriana Valley Northern Italy Lombardy Bergamo District.

Giornale Italiano di Entomologia. Vol. 3 (14): 229-256.

Garrison-Johnston, M.T., J.A. Moore, S.P. Cook, and G.J. Niehoff. 2003. Douglas-fir

beetle infestations are associated with certain rock and stand types in the inland

northwestern United States. Environmental Entomology. Vol. 32 (6): 1354-1363.

Gereben, B.A. 1995. Co-ocurrence and microhabitat distribution of six Nebria species

(Coleoptera: Carabidae) in an alpine glacier retreat zone in the Alps, Austria. Arctic and

Alpine Research. Vol. 27 (4): 371-379.

Greenslade, P.J.M. 1964. Pitfall trapping as a method for studying populations of Carabdae

(Coleoptera). Journal of Animal Ecology. Vol. 33 (2): 301-310.

Hammer, Ø., D.A.T. Harper, and P. D. Ryan, 2001. PAST: Paleontological statistics software

package for education and data analysis. Palaeontologia Electronica 4 (1): 9.

http://palaeo-electronica.org/2001_1/past/issue1_01.htm

Harvard University MCZ Insect Collection. http://insects.oeb.harvard.edu/mcz/ 69

Haskell, D.G. 2000. Effects of forest roads on macroinvertebrate soil fauna of the southern

Appalachian mountains. Conservation Biology. Vol. 14 (1): 57-63.

Haubold, V.L. 1951. Distribution of the Carabidae (Coleoptera) of Boulder County, Colorado.

American Midland Naturalist. Vol. 45 (3): 683-710.

Heffern, D.J. 1998. A survey of the Cerambycidae (Coleoptera) or longhorned beetles of

Colorado (Insects of Western North America). Gillette Museum of Arthropod Diversity,

Dept. of Bioagricultural Sciences and Pest Management, Colorado State University, Fort

Collins Colorado.

Holmes, P.R., D.C. Boyce, and D.K. Reed. 1993. The ground beetle (Coleoptera: Carabidae)

fauna of Welsh peatland biotypes: Factors influencing the distribution of ground beetles

and conservation implications. Biological Conservation. Vol. 63 (2): 153-161.

Holopainen, J.K., T. Bergman, E.L. Hautala, and J. Oksanen. 1995. The ground beetle

fauna (Coleoptera: Carabidae) in relation to soil properties and foliar fluoride content in

spring cereals. Pedobiologia. Vol. 39 (3): 193-206.

Hosoda, H. 1999. Altitudinal occurrence of ground beetles (Coleoptera, Carabidae) on Mt.

Kurobi, central Japan, with special reference to forest vegetation and soil characteristics.

Pedobiologia. Vol. 43 (4): 364-371.

Huggett, R. 1995. Geoecology: An evolutionary approach. Routledge, London England.

Jones, M. E., T.D. Paine, M.E. Fenn, and M.A. Poth. 2004. Influence of ozone and nitrogen

deposition on bark beetle activity under drought conditions. Forest Ecology and

Management. Vol. 200 (1-3): 67-76. 70

Lang, A. 2000. The pitfalls of pitfalls: A comparison of pitfall trap catches and absolute density

estimates of epigeal invertebrate predators in arable land. Journal of Pest Science. Vol.

73: 99-106.

Kippenhan, M.G. 1990. A survey of the tiger beetles (Coleoptera: Cicindelidae) of Colorado.

Entomological News. Vol. 101 (5): 307-315.

Koivula, M.J. 2005. Effects of forest roads on spatial distribution of boreal carabid beetles

(Coleoptera: Carabidae). Coleopterists Bulletin. Vol. 59 (4): 465-487.

Kulakowski, D., T.T. Veblen, and P. Bebi. 2003. Effects of fire and spruce beetle outbreak

legacies on the disturbance regime of a subalpine forest in Colorado. Journal of

Biogeography. Vol. 30 (9): 1445-1456.

Larochelle, A. and M.C. Larviviere. 2003. A natural history of the ground beetles (Coleoptera:

Carabidae) of America north of Mexico. Pensoft Publishers, Sofia Bulgaria.

LeConte, J.L. 1878. The Coleoptera of the alpine regions of the Rocky Mountains. Bulletin of

the United States Geological and Geographical Survey of the Territories. Vol. 4 (2): 447-

480.

LeConte, J.L. 1879. The Coleoptera of the alpine Rocky Mountain regions - Part II. Bulletin

of the United States Geological and Geographical Survey of the Territories. Vol.5: 499-

520.

Lobo, J.M. and G. Halffter. 2000. Biogeographical and ecological factors affecting the

altitudinal variation of mountainous communities of coprophagous beetles (Coleoptera:

Scarabaeoidea): a comparative study. Annals of the Entomological Society of America.

Vol. 93 (1): 115-126. 71

Lobo, J.M., E. Chehlarov, and B. Gueorguiev. 2007. Variation in dung beetle (Coleoptera:

Scarabaeoidea) assemblages with altitude in the Bulgarian Rhodopes Mountains: A

comparison. European Journal of Entomology. Vol. 104: 489-495.

Loffler, J., and O-D. Finch. 2005. Spatio-temporal gradients between high mountain

ecosystems of central Norway. Arctic, Antarctic, and Alpine Research. Vol. 37: (4):

499-513.

McCambridge, W.F. and F.B Knight. 1972. Factors affecting spruce beetles during a small

outbreak. Ecology. Vol. 53 (5): 830-839.

McIntyre, N.E., J. Rango, W.F. Fagan, and S.H. Faeth. 2001. Ground arthropod community

structure in a heterogeneous urban environment. Landscape and Urban Planning. Vol.

52: 257-274.

Montgomery, C.W. 1997. Fundamentals of geology. Wm. C. Brown Publishers, Dubuque,

Iowa.

Neff, J. C., R. Reynolds, R.L. Sandford Jr., D. Fernandez, and P. Lamothe. 2006. Controls

of bedrock geochemistry on soil and plant nutrients in southeastern Utah. Ecosystems.

Vol. 9: 879–893.

Ottesen, P.S. 1996. Niche segregation of terrestrial alpine beetles (Coleoptera) in relation to

environmental gradients and phenology. Journal of Biogeography. Vol. 23: 353-369.

Paje, F. and D. Mossakowski. 1984. pH-preferences and habitat selection in carabid beetles.

Oecologia. Vol. 64: 41-46.

Plaster, R.W. and W.C. Sherwood. 1971. Bedrock weathering and residual soil formation in

central Virginia. GSA Bulletin. Vol. 82 (10): 2813-2826. 72

Rahn, P.H. 1996. Engineering geology: An environmental approach. Prentice Hall, Upper

Saddle River, New Jersey.

Rich, C., L. Walters, D. Stewart, R. Anderhalt, C. Onasch, D. May, and J. Evans.

Unpublished. Geologic map of the Molas Lake-Sultan Creek area, San Juan County,

Colorado. Bowling Green State University, Geology Department, Bowling Green, Ohio.

Rieske, L.K. and L.J. Buss. 2001. Influence of site on diversity and abundance of ground- and

litter-dwelling Coleoptera in Appalachian Oak–Hickory forests. Environmental

Entomology. Vol. 30 (3): 484-494.

Roovers, L.M. and A.J. Rebertus. 1993. Stand dynamics and conservation of an old-growth

Engelmann spruce-subalpine fir forest in Colorado. Natural Areas Journal. Vol. 13 (4):

256-267.

Rüdiger, K. 2001. Invertebrate succession on an alpine glacier foreland. Ecology. Vol. 82 (8):

2261-2278.

Ryerson, D.E., T.W. Swetnam, and A.M. Lynch. 2003. A tree-ring reconstruction of western

spruce budworm outbreaks in the San Juan Mountains, Colorado, U.S.A. Canadian

Journal of Forest Research/Revue Canadienne de Recherche Forestiere. Vol. 33 (6):

1010-1028.

Sanderson, R.A., S.P. Rushton, A.J. Cherrill, and J.P. Byrne. 1995. Soil, vegetation, and

space: An analysis of their effects on the invertebrate communities of a Moorland in

northeast England. Journal of Applied Ecology. Vol. 32: 506-518.

Schmoller, R.R. 1971a. Habitats and zoogeography of alpine tundra Arachnida and Carabidae

(Coleoptera) in Colorado. The Southwestern Naturalist. Vol. 15 (3): 319-329. 73

Schmoller, R.R. 1971b. Nocturnal arthropods in the alpine tundra of Colorado. Arctic and

Alpine Research. Vol. 3 (4): 345-352.

Soto, T. 1996. Altitudinal variation in life cycles of carabid beetles: Life-cycle strategy and

colonization in alpine zones. Arctic and Alpine Research. Vol. 28 (4): 441-447.

Strecker, H. 1878. Section II: Lepidoptera. In Annual Report upon Explorations and Surveys

in the Department of the Missouri being Appendix SS of the Annual Report of the Chief

of Engineers for 1878, by E.H. Ruffner. 1847-1867.

Thiele, H.U. 1977. Carabid beetles in their environments: A study on habitat selection by

adaptation in physiology and behaviour. Springer-Verlag, New York.

Thingstad, P.G. 1987. Pitfall trapping of the carabid fauna in alpine and subalpine habitats at

Saltfjellet North Norway. Fauna Norvegica Series B. Vol. 32 (2): 105-116.

Varchola, J.M., and J.P. Dunn. 1999. Changes in ground beetle (Coleoptera: Carabidae)

assemblages in farming systems bordered by complex or simple roadside vegetation.

Agriculture, Ecosystem, and Environment. Vol. 73: 41-49.

Veblen, T.T., K.S. Hadley, S.M. Reid and A.J. Rebertus. 1991. The response of subalpine

forests to spruce beetle outbreak in Colorado USA. Ecology (Washington D C). Vol. 72

(1): 213-231.

Veblen, T.T., K.S. Hadley, E.M. Nel, T. Kitzberger, M. Reid, and R. Villalba. 1994.

Disturbance regime and disturbance interactions in a Rocky Mountain subalpine forest.

Journal of Ecology. Vol. 82(1): 125-135.

Vermeulen, H.J.W. 1993. The composition of the carabid fauna on poor sandy road-side

verges in relation to comparable open areas. Biodiversity and Conversation. Vol. 2: 331-

350. 74

Ward, D.E., T.R. New, and A.L. Yen. 2001. Effects of pitfall trap spacing on the abundance,

richness, and composition of invertebrate catches. Journal of Insect Conservation. Vol.

5: 47-53.

Weeks Jr., R.D., and N.E. McIntyre. 1997. A comparison of live versus kill pitfall trapping

techniques using various killing agents. Entomologia Experimentalis et Applicata. Vol.

82: 267-273.

Wickman, H.F. 1902. The Coleoptera of Colorado. Bulletin of the State University of Iowa.

No. 5 (3): 217-310.

75

Table 1. Total abundance caught at each site for each carabid species.

Elbert Ouray-Leadville Molas Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Total Amara quenseli 21 38 26 23 12 29 41 6 26 82 1 206 45 5 0 561 Harpalus nigritarsis 13 22 12 30 16 24 5 9 25 12 2 38 5 27 18 258 Carabus taedatus 9 2 5 4 6 8 6 3 7 5 8 1 1 2 4 71 Amara laevipennis 0 4 14 7 0 10 0 16 9 1 2 0 1 4 1 69 Calathus ingratus 2 0 20 2 3 0 5 2 3 2 1 0 4 1 5 50 Bembidion dauricum 3 1 17 1 4 0 6 0 0 0 0 0 0 0 9 41 Cymindis cribricollis 0 2 1 1 1 5 6 2 5 1 1 2 1 1 2 31 Agonum placidum 0 0 19 0 4 1 0 0 1 1 0 0 0 0 0 26 Harpalus animosus 3 2 5 3 1 2 1 2 5 0 0 0 0 0 0 24 Bembidion mutatum 7 0 2 3 0 0 0 0 0 0 0 1 6 2 2 23 Harpalus laticeps 0 1 4 1 1 1 2 2 4 0 0 0 0 0 0 16 Harpalus opacipennis? 0 0 0 1 1 2 1 1 2 2 0 2 0 2 0 14 Amara ellipsis 0 0 1 1 0 0 0 1 3 2 2 0 0 0 0 10 Notiophilus aquaticus 1 1 1 0 0 0 1 0 1 0 0 1 0 1 3 10 Agonum cupreum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4 4 Amara merula 0 0 0 0 1 0 0 0 1 0 0 2 0 0 0 4 Nebria catenata 0 0 0 4 0 0 0 0 0 0 0 0 0 0 0 4 Pterostichus protractus 0 0 0 0 0 0 1 0 0 2 0 0 0 0 0 3 Amara (Bradytus) lindrothi 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 2 Amara littoralis 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 2 Calathus advena 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 2 Harpalus ellipsis? 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 2 Rhadine sp. 1 (nivalis grp) 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 2 Amara (Bradytus) sp. 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 Amara obesa 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 Amara sinuosa 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 Dyschirius dejeanii 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 Nebria gyllenhali 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 Pterostichus restrictus 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 Rhadine sp. 2? (nivalis grp) 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 total 59 75 127 83 52 83 77 45 93 111 19 253 65 46 48 1236 76

Table 2. Total frequency calculated for each carabid species at each site.

Elbert Ouray-Leadville Molas Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Total Harpalus nigritarsis 6 9 5 10 11 8 3 6 10 6 2 13 5 10 10 114 Amara quenseli 8 7 6 8 6 8 9 3 11 11 1 12 8 4 0 102 Carabus taedatus 7 2 4 4 6 5 5 3 6 5 4 2 1 2 3 59 Amara laevipennis 0 4 7 3 0 4 0 9 7 1 1 0 1 3 1 41 Calathus ingratus 2 0 8 2 3 0 3 2 3 2 1 0 2 1 4 33 Cymindis cribricollis 0 2 1 1 1 3 5 2 4 1 1 2 1 1 2 27 Bembidion dauricum 3 1 7 1 3 0 4 0 0 0 0 0 0 0 4 23 Harpalus animosus 3 2 3 3 1 2 1 2 5 0 0 0 0 0 0 22 Bembidion mutatum 1 0 2 2 0 0 0 0 0 0 0 1 5 2 2 15 Harpalus laticeps 0 1 3 1 1 1 2 2 3 0 0 0 0 0 0 14 Harpalus opacipennis? 0 0 0 1 1 2 1 1 1 2 0 2 0 2 0 13 Agonum placidum 0 0 6 0 3 1 0 0 1 1 0 0 0 0 0 12 Notiophilus aquaticus 1 1 1 0 0 0 1 0 1 0 0 1 0 1 3 10 Amara ellipsis 0 0 1 1 0 0 0 1 3 2 1 0 0 0 0 9 Agonum cupreum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4 4 Amara merula 0 0 0 0 1 0 0 0 1 0 0 2 0 0 0 4 Pterostichus protractus 0 0 0 0 0 0 1 0 0 2 0 0 0 0 0 3 Amara (Bradytus) lindrothi 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 2 Amara littoralis 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 2 Calathus advena 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 2 Harpalus ellipsis? 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 2 Rhadine sp. 1 (nivalis grp) 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 2 Amara (Bradytus) sp. 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 Amara obesa 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 Amara sinuosa 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 Dyschirius dejeanii 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 Nebria catenata 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 Nebria gyllenhali 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 Pterostichus restrictus 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 Rhadine sp. 2? (nivalis grp.) 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 total 31 31 54 40 39 35 37 32 57 34 13 35 25 27 33 523 77

Table 3. Carabidae values for species richness, abundance, frequency, and diversity calculated for each site.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Species Richness 8 10 13 15 13 10 13 11 14 11 9 8 8 10 9 Abundance 59 75 127 83 52 83 77 45 93 111 19 253 65 46 48 Frequency 31 31 54 40 39 35 37 32 57 34 13 35 25 27 33 Shannon-Wiener 1.73 1.42 2.18 1.93 2.04 1.72 1.72 1.95 2.08 1.06 1.85 0.63 1.14 1.51 1.85 Eveness 0.83 0.62 0.85 0.71 0.80 0.75 0.67 0.81 0.79 0.44 0.84 0.30 0.55 0.66 0.84

Table 4. Means and ANOVA results for species richness, abundance, frequency, and diversity within each formation.

Elbert Ouray-Leadville Molas P DF F Species Richness 12 ± 1.2 12 ± 0.7 9 ± 0.4 0.045 2,12 4.05 Abundance 79 ± 13 82 ± 11 86 ± 42 0.982 2,12 0.02 Frequency 39 ± 4 39 ± 5 27 ± 4 0.096 2,12 2.87 Shannon-Wiener 1.86 ± 0.13 1.71 ± 0.18 1.40 ± 0.23 0.235 2,12 1.64 Eveness 0.76 ± 0.043 0.69 ± 0.067 0.64 ± 0.101 0.516 2,12 0.70

Table 5. Absolute total of the carabid community measurements for each formation.

Elbert Ouray-Leadville Molas Species Richness 21 21 17 Abundance 396 409 431 Frequency 195 195 133 Shannon-Wiener 2.17 1.89 1.42 Eveness 0.71 0.62 0.50 78

Table 6. ANOVA statistics calculated for mean abundance differences between formations for the most common carabids.

Species P DF F Agonum placidum 0.289 2,12 1.38 Amara ellipsis 0.351 2,12 1.14 Amara laevipennis 0.268 2,12 1.47 Amara quenseli 0.729 2,12 0.32 Bembidion dauricum 0.391 2,12 1.02 Bembidion mutatum 0.182 2,12 1.97 Calathus ingratus 0.546 2,12 0.64 Carabus taedatus 0.272 2,12 1.46 Cymindis cribricollis 0.014 2,12 6.25 Harpalus animosus 0.020 2,12 5.47 Harpalus laticeps 0.089 2,12 2.98 Harpalus nigritarsis 0.859 2,12 0.15 Harpalus opacipennis? 0.082 2,12 3.11 Notiophilus aquaticus 0.531 2,12 0.67

Table 7. ANOVA statistics calculated for mean frequency differences between formations

for the most common carabids.

Frequency P DF F Agonum placidum 0.227 2,12 1.68 Amara ellipsis 0.191 2,12 1.91 Amara laevipennis 0.290 2,12 1.37 Amara quenseli 0.338 2,12 1.19 Bembidion dauricum 0.185 2,12 1.95 Bembidion mutatum 0.071 2,12 3.33 Calathus ingratus 0.560 2,12 0.61 Carabus taedatus 0.041 2,12 4.22 Cymindis cribricollis 0.025 2,12 5.09 Harpalus animosus 0.018 2,12 5.77 Harpalus laticeps 0.042 2,12 4.16 Harpalus nigritarsis 0.715 2,12 0.35 Harpalus opacipennis? 0.164 2,12 2.11 Notiophilus aquaticus 0.531 2,12 0.67

79

Table 8. Relative abundances of the species composition between each formation in order of most dominant to least. Line break represents 90% of the species composition in a formation (upper portion).

Elbert Ouray-Leadville Molas Relative Relative Relative Species Abundance Species Abundance Species Abundance Amara quenseli 30.30% Amara quenseli 44.99% Amara quenseli 59.63% Harpalus nigritarsis 23.48% Harpalus nigritarsis 18.34% Harpalus nigritarsis 20.88% Calathus ingratus 6.82% Amara laevipennis 8.80% Carabus taedatus 3.71% Carabus taedatus 6.57% Carabus taedatus 7.09% Calathus ingratus 2.55% Bembidion dauricum 6.57% Cymindis cribricollis 4.65% Bembidion mutatum 2.55% Amara laevipennis 6.31% Calathus ingratus 2.93% Bembidion dauricum 2.09% Agonum placidum 5.81% Harpalus animosus 2.44% Amara laevipennis 1.86% Harpalus animosus 3.54% Harpalus laticeps 2.20% Cymindis cribricollis 1.62% Bembidion mutatum 3.03% Harpalus opacipennis? 1.96% Notiophilus aquaticus 1.16% Harpalus laticeps 1.77% Bembidion dauricum 1.47% Harpalus opacipennis? 0.93% Cymindis cribricollis 1.26% Amara ellipsis 1.47% Agonum cupreum 0.93% Nebria catenata 1.01% Agonum placidum 0.73% Amara ellipsis 0.46% Notiophilus aquaticus 0.76% Pterostichus protractus 0.73% Amara merula 0.46% Harpalus opacipennis? 0.51% Notiophilus aquaticus 0.49% Amara (Bradytus) lindrothi 0.46% Amara ellipsis 0.51% Amara merula 0.24% Amara littoralis 0.23% Rhadine sp. 1 (nivalis grp) 0.51% Calathus advena 0.24% Amara sinuosa 0.23% Amara merula 0.25% Harpalus ellipsis? 0.24% Pterostichus restrictus 0.23% Amara littoralis 0.25% Amara (Bradytus) sp. 0.24% Calathus advena 0.25% Amara obesa 0.24% Harpalus ellipsis? 0.25% Nebria gyllenhali 0.24% Dyschirius dejeanii 0.25% Rhadine sp. 2? (nivalis grp) 0.24%

80

Table 9. Relative frequencies of the species composition between each formation in order of most dominant to least. . Line break represents 90% of the species composition in a formation (upper portion).

Elbert Ouray-Leadville Molas Relative Relative Relative Species Frequency Species Frequency Species Frequency Harpalus nigritarsis 21.03% Amara quenseli 21.54% Harpalus nigritarsis 30.08% Amara quenseli 17.95% Harpalus nigritarsis 16.92% Amara quenseli 18.80% Carabus taedatus 11.79% Carabus taedatus 12.31% Carabus taedatus 9.02% Bembidion dauricum 7.69% Amara laevipennis 10.77% Bembidion mutatum 7.52% Calathus ingratus 7.69% Cymindis cribricollis 7.69% Calathus ingratus 6.02% Amara laevipennis 7.18% Calathus ingratus 5.13% Cymindis cribricollis 5.26% Harpalus animosus 6.15% Harpalus animosus 5.13% Amara laevipennis 4.51% Agonum placidum 4.62% Harpalus laticeps 4.10% Notiophilus aquaticus 3.76% Harpalus laticeps 3.08% Harpalus opacipennis? 3.59% Agonum cupreum 3.01% Bembidion mutatum 2.56% Amara ellipsis 3.08% Bembidion dauricum 3.01% Cymindis cribricollis 2.56% Bembidion dauricum 2.05% Harpalus opacipennis? 3.01% Notiophilus aquaticus 1.54% Agonum placidum 1.54% Amara (Bradytus) lindrothi 1.50% Amara ellipsis 1.03% Pterostichus protractus 1.54% Amara merula 1.50% Harpalus opacipennis? 1.03% Notiophilus aquaticus 1.03% Amara ellipsis 0.75% Rhadine sp. 1 (nivalis grp) 1.03% Amara (Bradytus) sp. 0.51% Amara littoralis 0.75% Amara littoralis 0.51% Amara merula 0.51% Amara sinuosa 0.75% Amara merula 0.51% Amara obesa 0.51% Pterostichus restrictus 0.75% Calathus advena 0.51% Calathus advena 0.51% Dyschirius dejeanii 0.51% Harpalus ellipsis? 0.51% Harpalus ellipsis? 0.51% Nebria gyllenhali 0.51% Nebria catenata 0.51% Rhadine sp. 2? (nivalis grp.) 0.51%

81

Table 10. Jaccard coefficients calculated for carabid similarity between individual sites.

Site 2 3 4 5 6 7 8 9 10 11 12 13 14 15 1 0.50 0.62 0.44 0.40 0.29 0.50 0.36 0.38 0.27 0.31 0.45 0.45 0.50 0.55 2 0.64 0.47 0.44 0.54 0.53 0.50 0.50 0.31 0.36 0.38 0.38 0.43 0.46 3 0.65 0.53 0.53 0.53 0.60 0.69 0.56 0.47 0.40 0.50 0.53 0.57 4 0.47 0.56 0.47 0.63 0.53 0.44 0.41 0.35 0.44 0.56 0.41 5 0.53 0.63 0.50 0.59 0.41 0.29 0.40 0.31 0.35 0.29 6 0.44 0.62 0.60 0.50 0.36 0.38 0.38 0.43 0.27 7 0.50 0.50 0.41 0.29 0.40 0.31 0.44 0.38 8 0.67 0.57 0.54 0.36 0.46 0.50 0.33 9 0.56 0.44 0.47 0.38 0.50 0.35 10 0.54 0.36 0.46 0.50 0.33 11 0.31 0.55 0.46 0.38 12 0.45 0.64 0.42 13 0.64 0.55 14 0.58 15

Table 11. Overall Jaccard coefficients calculated for carabid similarity between bedrock formations.

Formations Ouray-Leadville Molas Elbert 0.62 0.52 Ouray-Leadville 0.41 Molas

82

Table 12. ANOSIM Output for the Jaccard coefficient.

Number R Significance Possible Actual > = Groups Statistic Level Permutations Permutations Observed Elbert - Ouray-Leadville -0.092 77% 126 126 126 Elbert - Molas -0.48 100% 126 126 126 Ouray-Leadville - Molas -0.63 100% 126 126 126 Global R -0.419 Significance 100%

83

Fig. 1. General view of study area.

Fig. 2. Map of study area showing bedrock formations and where sites were located. 84

Fig. 3. Pitfall trap set-up design.

Fig. 4. Site 12 showing the typical pitfall trap set-up. Yellow flags represent where the cups were located. 85

Fig. 5. Typical pitfall trap used in study area.

14 ab a 12

10 b 8

6

4

2

Mean Carabid Species Richness 0 Elbert Ouray-Leadville Molas

Fig. 6. Mean (± SEM) species richness in bedrock formations. 86

Fig. 7. Mean (± SEM) abundance, frequency, Shannon-Wiener Index, and evenness in bedrock formations. 87

Fig. 8. Individual carabid species mean (± SEM) abundance in formations.

88

Fig. 9. Individual carabid species mean (± SEM) frequency in formations.

89

Fig 10. Graphical Non-metric Multi-Dimensional Scaling output from the analysis of similarity (Table 8 shows numerical output) showing how sites within bedrock formations group.

90

12 14 15 1 5 7 3 9 8 4 6 2 10 11 13 1

0.9

0.8

0.7 LEGEND Sites Formation 1 Elbert 2 Elbert Jaccard Similarity 0.6 3 Elbert 4 Elbert 5 Elbert 6 Ouray-Leadville 7 Ouray-Leadville 8 Ouray-Leadville 0.5 9 Ouray-Leadville 10 Ouray-Leadville 11 Molas 12 Molas 13 Molas 0.4 14 Molas 15 Molas

0 3 6 9 12 15 Arbitrary Scale Representing Number of Sites in Analysis

Fig. 11. Cluster analysis of Jaccard coefficients for the Carabidae fauna. Circled areas represent the three main groups that branch out due to similar species composition.

91

CHAPTER III

Influences of Environmental Factors on a Carabidae Community in a Subalpine Meadow

Studies have shown that multiple abiotic and biotic factors (e.g., pH, plant structure and composition, soil density, soil water content, sediment sizes, geomorphology, soil temperature) play a role in Carabidae habitat preferences (e.g., Thiele 1977, Paje and D. Mossakowski 1984,

Eyre et al. 1990, Quinn et al. 1991, Holmes et al. 1993, Desender et al. 1994, Gereben 1995,

Holopainen et al. 1995,Sanderson et al. 1995, Ottensen 1996, Hosoda 1999, Ings and Hartley

1999, Magura et al. 2003, XiaoDong et al. 2003, Magura et al. 2005, Long and Medina 2006).

Several studies have incorporated multivariate analyses in mountainous communities for these

and other variables to explain Carabidae communities (e.g., Ottensen 1996, Rykken et al. 1997,

Humphrey et al. 1999, Magura et al. 2003, Gutierrez et al. 2004, Loffler and Finch 2005,

Apigian et al. 2006, Gobbi et al. 2007, Pohl et al. 2007, and Gonzalez-Megias et al. 2008). Yet,

these studies are still lacking in certain geographical regions, especially within the Rocky

Mountains, and none were performed in Colorado for Carabidae. In Colorado, most studies on

Carabidae communities in the Rocky Mountains are based on elevational gradients (Haubold

1951, Armin 1963, and Elias 1987). Temperature effects (Schmoller 1971a) and vegetation

cover and soil moisture (Schmoller 1971b) were also briefly studied in the alpine tundra. The

main objective of this paper was to find out which environmental variable(s) influenced the

carabid community of a subalpine meadow. This was the first study performed within the San

Juan Mountains of Colorado for Carabidae.

92

Materials and Methods

Site Location. The study area was in the San Juan Mountains of Colorado, several kilometers

south of Silverton, a small mining town in southwest Colorado (Fig. 1). The Sultan Creek-Molas

Lake region was chosen for its several bedrock outcroppings that are mostly parallel to each

other and occur at similar elevations at around 3,170 m (10,400 ft). The study area itself is an

open meadow with remnants of the spruce-fir forest, just south of Sultan Creek and about a

kilometer north of Molas Lake on the east side along Highway 550. The open meadow borders

the heavily forested area that lies on the Precambrian bedrock, within the Animas River

watershed. The whole study area was about 400 m X 400 m.

Carabidae Methodology. The carabids used for this project were from the same database from

Chapter Two. An in depth description of the collection methodology can be found in Chapter

Two. A short summary follows for the carabid collection. Pitfall trapping was used to collect

the insects. A total of fifteen sites were set up in a grid of three transects, five sites each. Nine

pitfall traps (250 ml Corning Snap-Seal cup, no. 1730 design) were placed in each site. Trapping

occurred weekly for eight weeks in 2006 (June 17 to August 11) and six weeks in 2007 (June 24

to August 3). A 50:50 solution of propylene glycol:water was used to kill and preserve insects.

Traps were cleaned out and insects stored in 70% ethanol alcohol. Carabidae were identified by

the author using museum specimens at Colorado State University, Fort Collins and The

University of Colorado, Boulder; by carabid specialists Bob Davidson (Carnegie Museum of

Natural History) and Foster Purrington (The Ohio State University); and by using the Harvard

Insect Collection website.

Biotic and Physical Variables Measured. Latitude and Longitude, elevation and aspect were

measured with a GPS unit for each site. The slope was estimated for each site with a Brunton 93

compass and board. The abiotic soil factors measured were: temperature, pH, moisture,

compressive strength, mean grain size and grain size sorting. The biotic soil factors measured

were: plant cover, height, and species richness and detritus cover and weight. In depth

methodology of how these variables were gathered can be found in Chapter One.

Quantitative and Statistical Analyses. The two seasons of carabid data were combined.

Species richness, abundance, frequency, Shannon-Weiner index (H’), and evenness (E) were

calculated for each site. Frequency was the number of times (0-14, total weekly appearances) a

species appeared at a given site. Shannon-Weiner index (H’) was calculated to measure the

overall biodiversity of the sites. The higher H’ was the greater the diversity was. Evenness (E)

measured how similar the abundances of the species are within a site. When evenness equals 1,

all species abundances were the same, whereas the closer the number was to zero, the less even

(equal) a site was (mix of very abundant, and very rare species).

H’ = –∑ pi ln (pi)

pi = proportion of species i to total abundance

E = H’/ln S

H’ = Shannon-Weiner index

S = species richness

ANOVAs (aspect, bedrock, and slope only) and linear regression analyses were run on these

measurements against the environmental variables to see if any variable(s) influenced these 94

parameters. The environmental variables over both years were also combined and averaged (if

multiple weekly measurements had been taken).

The influence of environmental factors on the carabid composition were studied using a

canonical correspondence analysis (CCA) (ter Braak 1986, Petillon et al. 2008, and Highland

Statistics Ltd Website 2009) performed with the software program PAST (Hammer et al. 2001).

The environmental matrix consisted of overall averages of the variables that were measured

multiple times throughout the two seasons (Table 1). The Carabidae matrix consisted of the total

frequency of the individual species at each site (Table 2). Frequency was used to reduce the

weight of highly abundant species and rare species. A permutation test with 1000 iterations was

performed to test for significance of the species and the environmental variables.

Moran’s I was calculated for abundance, species richness, and frequency of the Carabidae

fauna to see if spatial autocorrelation occurred at the scale I used for these measurements (Moran

1950, Sawada 2004, Levine 2005, and Purkis et al. 2006). The Moran’s I value ranges from +1

(clustering) to -1 (perfect dispersion). A value of zero lacks autocorrelation (values are random).

n = number of sites

xi = variable at a particular location

xj = variable at another location (where i ≠ j)

x = mean of the variables

Wij = a matrix of spatial weights

if two sites are adjacent, a 1 is used, if two sites are not adjacent a 0 is used 95

Sites were analyzed on how similar they were based on environmental characteristics

shared using the CCA and cluster analysis. The cluster analysis was performed with the PAST

software. Euclidean distance was performed using the Ward’s algorithm and Rho using the

paired group algorithm, the latter having a percent similarity.

Results

Carabid Community Measurements and Environmental Factors. Table three shows the

results of correlations (relationships) between the environmental factors with the carabid community structure and biodiversity measurements. Carabid species richness was significantly influenced by plant species richness and geology (F = 8.61, df = 1, 13, P = 0.012 and F = 4.05, df

= 2, 12, P = 0.045 respectively) (Figs. 2 and 3). Carabid species richness was positively correlated with plant species richness. The Molas Formation was significantly lower in carabid species then the Ouray-Leadville Formations. Carabid abundance and biodiversity measures (H’ and evenness) were significantly influenced by plant cover (F = 5.09, df = 1, 13, P = 0.042, F =

8.79, df = 1, 13, P = 0.011, and F = 12.97, df = 1, 13, P = 0.003 respectively) (Figs. 4-6).

Carabid abundance was negatively correlated with plant cover, whereas carabid biodiversity and evenness were positively correlated with plant cover. Carabid frequency was positively correlated with sediment sorting (F = 9.17, df = 1, 13, P = 0.010) (Fig. 7).

Site Similarity. Most sites clumped linearly in mainly two of the quadrants from the CCA results except for sites 13 and 15 (Fig. 8). Site 15 had the lowest pH, compressive strength and highest soil moisture. Site 13 had the highest detritus cover and soil temperature and lowest plant cover. When a 95% ellipse was calculated for these results, Site 15 did not occur in the ellipse. Site 15 was overall quite different from the rest of the sites within the study. The cluster 96

analyses performed (Fig. 9) showed that sites were > 80% similar to each other (Fig. 9b). The

sites break out into two main groups based on the environmental factors.

CCA Analysis on the Carabidae Community. Table four displays the results of the environmental variables in the CCA. The first axis was positively correlated with compressive strength and pH of soil and the second axis was positively correlated with plant cover and soil moisture (Fig. 10). Since the first two axes explained only 35% of the carabid community, the third axis was plotted (Fig. 11). The third axis was positively correlated with slope and aspect.

The first four axes explained 60.65% of the Carabidae fauna. The permutation results created a

Trace value of 1.045 and Trace p-value of 0.498; species were not linearly related to the

environmental variables.

The results for the carabid community in the CCA analysis can be found in Figures 12

and 13. In general, about half of the species plot on either side of axis one, and about two-thirds of the species plot on the positive side of axis two (higher plant cover and moisture levels). The rare species mostly plotted out away from the center where the majority of the species loosely clumped (Agonum cupreum, Amara sinuosa, Amara (Bradytus) lindrothi, Rhadine sp. 2?, and

Pterostichus restrictus) (Fig. 12). Rare species, if at an extreme location (or environmental variable) plotted out farther away. Of the more common carabids, Bembidion mutatum separated out due to low plant cover and moisture and to a lesser extent due to lower compressive strength and pH. Notiophilus aquaticus and Bembidion dauricum separated out somewhat from the rest of the carabids, too, based on lower compressive strength and pH (Axis 1). A few species appeared to tightly clump together (correlate with each other). This relationship was maximized in the third axis for Amara (Bradytus) sp., Calathus advena, and Dyschirius dejeanii (Fig. 13).

These three rare species shared a preference for a similar slope (7° – 10°) and a northern aspect. 97

Spatial Autocorrelation of Carabidae. Moran’s I values for the overall carabidae abundance,

frequency, and richness were -0.210, +0.046, and +0.158 respectively. The Moran’s I values

indicated that the pattern of these carabid measurements were random (values are dissimilar near

each other) in the study area, no autocorrelation was detected.

Discussion

All carabid community measurements [(species richness, abundance, frequency, and

biodiversity (H’ and E)] were significantly affected by at least one environmental variable, with

plant cover affecting the most measurements. Carabid species richness, abundance, and

biodiversity were mainly influenced by plants. The habitat structure and diversity in general has

been shown to be quite influential to carabid communities (Ings and Hartley 1999). When plant species richness increased carabid species richness also increased. This positive trend is well

known in the literature among plant-animal species relationships (Tews et al. 2004). Even

though carabids are mainly predators and scavengers, a higher diversity of plants can lead to

higher number of different food resources for these beetles (Landis et al. 2000 and Landis et al.

2005). Higher plant cover led to a higher overall biodiversity (Shannon-Wiener and Evenness).

Carabid abundance increased with less plant cover. Lower plant cover provides less resistance

for ground beetles to move on. A large number of carabid species tend to affiliate themselves

with open ground and very sparse to moderate vegetation (Larochelle and Larviviere 2003; see

discussion for some of the species found in this study that affiliate with this type of habitat in

Chapter two and below).

Geology and sediment sorting were the other two environmental variables that influenced

carabid species richness and frequency, respectively. Geology is discussed in great detail in 98

Chapter Two. The more poorly sorted the sediment was, the higher the carabid frequency. The

sorting ranged from 2.17 Φ – 4.19 Φ (in general terms, very poorly sorted sediments and

extremely poorly sorted sediments, respectively). These soils had a wide range of sizes

structuring the soil texture: gravel, sand, silt, and clay. Poorly sorted material can reduce

permeability of water, retaining it for longer times, thus increasing moisture levels (Montgomery

1997). Carabids have been shown to be influenced by soil moisture, which might explain the

higher frequency of carabid species for these sites. However, moisture did not show any

significant correlations with frequency or other carabid measurements taken. It did influence the

overall community composition of the carabids (see CCA discussion below).

The CCA results showed several environmental factors the carabid community was

affected by. In order of importance these are: compressive strength, pH, plant cover, soil

moisture, slope, and aspect. These factors explained about 48% of the community. A problem

arises in interpreting these factors. On all three axes plotted, two environmental factors were

positively correlated with each other, and their coordinates were very close (e.g., Axis one: 0.68

and 0.66 for compressive strength and pH, respectively). This may imply that both were

affecting the beetle community or one was while the other variable just correlated with the other.

In past studies though, compaction of soil (measured here as compressive strength) and pH has

been shown to affect diversity of carabids (Magura et al. 2003) and specific species preferences

(Paje and Mossakowski 1984). In quite a few studies, soil moisture content/humidity tended to

be one of the leading factors in structuring carabid communities (e.g., Thiele 1977, Holmes et al.

1993, Asteraki et al. 1995, Ottesen 1996). Soil moisture was the fourth most important factor in

this study, although it correlated strongly with plant cover, which also affected the carabids.

Both soil moisture and plant cover affect carabid communities and in turn affect each other: an 99

increase in plant cover leads to more moisture held in the ground due to reduction in evaporation

rates.

The species themselves in the CCA mostly clustered loosely near the origin of the axes in

three of the four quadrants. Several of the rare species plotted far away. This can be explained

by the fact that the rare species mostly appeared once or twice (and usually in the same site), and

if they had a specific environmental variable(s) that may be extreme or opposite of most other

species, they will plot out of the group as outliers. An example is Agonum cupreum, plotting at

an extreme negative value on axis one due to it being only found at site 15. Site 15 was shown to

be quite different from the rest of the sites. Three of the rare species correlated with each other

based on slope and aspect: Amara (Bradytus) sp., Calathus advena, and Dyschirius dejeanii.

Only a few common species appear to be affected differently than the rest of the

community, especially Bembidion mutatum, which was one of only three species in the lower left

quadrant and to a lesser extent Notiophilus aquaticus and Bembidion dauricum. Bembidion

mutatum is found in open areas (meadows, crops, pits, etc) with fine sandy soil with organic

debris, moderate moisture, and moss or sparse vegetation (Larochelle and Larviviere 2003).

Bembidion dauricum is an open ground species that is found in dry soils, fine sandy sediment, and low to no vegetation (Larochelle and Larviviere 2003). Notiophilus aquaticus is slightly

more general then these two. It is found in open areas (meadows, crops, pits, etc) that are

moderately dry, sandy to gravelly soils, and sparse to moderate vegetation (Larochelle and

Larviviere 2003). These three species generally affiliate themselves with open, sandy, and low

vegetation habitats, which the study area provides an ideal location for.

The environmental factors were not significant for a linear relationship with the carabid

community performed in the CCA. Thus, the species composition might be related nonlinearly 100 or spatially with the environmental factors. Spatial autocorrelation analyses resulted in a random distribution of values for carabid abundance, frequency, and species richness, so no autocorrelation was detected. Even though overall carabid frequency was not autocorrelated, the frequency of the individual species might be (which were used in the CCA). The individual species and the environmental factors were not examined for autocorrelation in this study. A future study should look at this in more detail regarding the possible spatial effects.

Conclusion

Carabid species richness, abundance, and biodiversity were mainly influenced by plant factors whereas carabid frequency was influenced by sediment sorting. Carabid species richness was also affecting by type of bedrock formation. The carabid species composition, based on

CCA, was influenced by compressive strength, pH, plant cover, soil moisture, slope, and aspect.

A few common species appeared to be affected differently than the rest of the community, whereas about half of the rare species separate out from the community. No spatial autocorrelation occurred for the general carabid community measurements taken, but the community composition may be spatially related to the environmental factors. Future analyses should be performed on only the common species for CCA to lessen the pull of the rare species and spatial autocorrelation tests on the individual species and environmental factors. Also, since this was a smaller scale study compared to other habitat analysis projects, future studies should incorporate a wider geographical area (the whole meadow surrounding Molas Lake) to test for correlations of these variables.

101

References

Apigian, K.O., D.L. Dahlsten, and S.L. Stephens. 2006. Biodiversity of Coleoptera and the

importance of habitat structural features in a Sierra Nevada mixed-conifer forest.

Environmental Entomology. Vol. 35 (4): 964-975.

Armin, L.C. 1963. A study of the family Carabidae (Coleoptera) in Boulder County, Colorado.

Dissertation, University of Colorado.

Asteraki, E.J., C.B. Hanks, and R.O. Clements. 1995. The influence of different types of

grassland field margin on carabid beetle (Coleoptera, Carabidae) communities.

Agriculture, Ecosystems, and Environment. Vol. 54: 195-202.

Desender, K. 1994. Carabid beetles: Ecology and evolution. Kluwer Academic Publishers,

Boston MA.

Elias, S.A. 1987. Colorado ground beetles (Coleoptera: Carabidae) from the Rotger Collection,

University of Colorado Museum. Great Basin Naturalist. Vol. 47 (4): 631-637.

Eyre, M.D., M.L. Luff, and S.P. Rushton. 1990. The ground beetle (Coleoptera, Carabidae)

fauna of intensively managed agricultural grasslands in northern England and southern

Scotland. Pedobiologia. Vol. 34 (1): 11-18.

Gereben, B.A. 1995. Co-ocurrence and microhabitat distribution of six Nebria species

(Coleoptera: Carabidae) in an alpine glacier retreat zone in the Alps, Austria. Arctic and

Alpine Research. Vol. 27 (4): 371-379.

Gobbi, M., B. Rossaro, A. Vater, F. De Bernardi, M. Pelfini, and P. Brandmayr. 2007.

Environmental features influencing carabid beetle (Coleoptera) assemblages along a

recently deglaciated area in the alpine region. Ecological Entomology. Vol. 32 (6):

682 – 689. 102

Gonzalez-Megías, A., J.M. Gomez, and F. Sanchez-Pinero. 2008. Factors determining

beetle richness and composition along an altitudinal gradient in the high mountains of the

Sierra Nevada National Park (Spain). Ecoscience. Vol. 15 (4): 429-441.

Gutierrez, D., R. Menendez, and M. Mendez. 2004. Habitat-based conservation priorities for

carabid beetles within the Picos de Europa National Park, northern Spain. Biological

Conservation. Vol. 115: 379-393.

Hammer, Ø., D.A.T. Harper, and P. D. Ryan, 2001. PAST: Paleontological statistics software

package for education and data analysis. Palaeontologia Electronica 4(1): 9pp.

http://palaeo-electronica.org/2001_1/past/issue1_01.htm

Harvard University MCZ Insect Collection. http://insects.oeb.harvard.edu/mcz/

Haubold, V.L. 1951. Distribution of the Carabidae (Coleoptera) of Boulder County, Colorado.

American Midland Naturalist. Vol. 45 (3): 683-710.

Highland Statistics Ltd Website. Accessed on April 2009. Hunting Spider Data, Canonical

Correspondence Analysis example. http://www.brodgar.com/spiders1.htm

Holmes, P.R., D.C. Boyce, and D.K. Reed. 1993. The ground beetle (Coleoptera: Carabidae)

fauna of Welsh peatland biotypes: Factors influencing the distribution of ground beetles

and conservation implications. Biological Conservation. Vol. 63 (2): 153-161.

Holopainen, J.K., T. Bergman, E.L. Hautala, and J. Oksanen. 1995. The ground beetle

fauna (Coleoptera: Carabidae) in relation to soil properties and foliar fluoride content in

spring cereals. Pedobiologia. Vol. 39 (3): 193-206.

Hosoda, H. 1999. Altitudinal occurrence of ground beetles (Coleoptera, Carabidae) on Mt.

Kurobi, central Japan, with special reference to forest vegetation and soil characteristics.

Pedobiologia. Vol. 43 (4): 364-371. 103

Humphrey, J.W., C. Hawes, A.J. Peace, R. Ferris-Kaan, and M.R. Jukes. 1999.

Relationships between insect diversity and habitat characteristics in plantation forests.

Forest Ecology and Management. Vol. 113: 11-21.

Ings, T.C. and S.E. Hartley. 1999. The effect of habitat structure on carabid communities

during the regeneration of a native Scottish forest. Forest Ecology and Management.

Vol. 119: 122-136.

Landis, D.A., S.D. Wratten, and G.M. Gurr. 2000. Habitat management to conserve natural

enemies of arthropod pests in agriculture. Annual Review of Entomology. Vol. 45: 175-

201.

Landis, D.A., F.D. Menalled, A.C. Costamagna and T.K. Wilkinson. 2005. Manipulating

plant resources to enhance beneficial arthropods in agricultural landscapes. Weed

Science. Vol. 53 (6): 902-908.

Larochelle, A. and M.C. Larviviere. 2003. A natural history of the ground beetles (Coleoptera:

Carabidae) of America north of Mexico. Pensoft Publishers, Sofia Bulgaria.

Levine, N. 2005. Chapter 4: Spatial distribution. CrimeStat Manual. 4.48-4.54.

Loffler, J., and O-D. Finch. 2005. Spatio-temporal gradients between high mountain

ecosystems of central Norway. Arctic, Antarctic, and Alpine Research. Vol. 37: (4):

499-513.

Long, J.W. and A.L. Medina. 2006. Consequences of ignoring geologic variation in

evaluating grazing impacts. Rangeland Ecology and Management. Vol. 59: 372-382.

Magura, T., B. Tothmeresz, and Z. Elek. 2003. Diversity and composition of carabids during

a forestry cycle. Biodiversity and Conservation. Vol. 12: 73–85. 104

Magura, T., B. Tothmeresz, and Z. Elek. 2005. Impacts of leaf litter addition on carabids in a

conifer plantation. Biodiversity and Conservation. Vol. 14: 475-491.

Montgomery, C.W. 1997. Fundamentals of geology. Wm. C. Brown Publishers, Dubuque,

Iowa.

Moran, P.A.P. 1950. Notes on continuous stochastic phenomena. Biometrika. Vol. 37 (1/2):

17-23.

Ottesen, P.S. 1996. Niche segregation of terrestrial alpine beetles (Coleoptera) in relation to

environmental gradients and phenology. Journal of Biogeography. Vol. 23: 353-369.

Paje, F. and D. Mossakowski. 1984. pH-preferences and habitat selection in carabid beetles.

Oecologia. Vol. 64: 41-46.

Petillon, J., A. Georges, A. Canard, J-C. Lefeuvre, J.P. Bakker, and F. Ysnel. 2008.

Influence of abiotic factors on spider and ground beetle communities in different salt-

marsh systems. Basic and Applied Ecology. Vol. 9: 743-751.

Pohl, G.R., D.W. Langor, and J.R. Spence. 2007. Rove beetles and ground beetles

(Coleoptera: Staphylinidae, Carabidae) as indicators of harvest and regeneration practices

in western Canadian Foothills Forests. Biological Conservation. Vol. 137: 294-307.

Purkis, S.J., S.W. Myint, and B.M. Riegl. 2006. Enhanced detection of the coral Acropora

cervicornis from the satellite imagery using a textural operator. Remote Sensing of

Environment. Vol. 101: 82-94.

Quinn, M.A., R.L. Kepner, D.D. Walgenbach, R.N. Foster, R.A. Bohls, P.D. Pooler, K.C.

Reuter, and J.L. Swain. 1991. Effect of habitat characteristics and perturbation from

insecticides on the community dynamics of ground beetles (Coleoptera: Carabidae) on

mixed grass rangeland. Environmental Entomology. Vol. 20 (5): 1285-1294. 105

Rykken, J.J., D.E. Capen, and S.P. Mahabir. 1997. Ground beetles as indicators of land type

diversity in the Green Mountains of Vermont. Conservation Biology. Vol. 11: 522–530.

Sanderson, R.A., S.P. Rushton, A.J. Cherrill, and J.P. Byrne. 1995. Soil, vegetation, and

space: An analysis of their effects on the invertebrate communities of a moorland in

northeast England. Journal of Applied Ecology. Vol. 32: 506-518.

Sawada, M. 2004. Global spatial autocorrelation indices-Moran’s I, Geary’s C and the General

Cross-Product Statistic. Research paper from the Laboratory for Paleoclimatology and

Climatology at the University of Ottawa.

http://www.lpc.uottawa.ca/publications/moransi/moran.htm

Schmoller, R.R. 1971a. Nocturnal arthropods in the alpine tundra of Colorado. Arctic and

Alpine Research. Vol. 3 (4): 345-352.

Schmoller, R.R. 1971b. Habitats and zoogeography of alpine tundra Arachnida and Carabidae

(Coleoptera) in Colorado. The Southwestern Naturalist. Vol. 15 (3): 319-329. ter Braak, C.J.F. 1986. Canonical correspondence analysis: A new eigenvector technique for

multivariate direct gradient analysis. Ecology. Vol. 67: 1167-1179.

Tews, J., U. Brose, V. Grimm, K. Tielborger, M.C. Wichmeann, M. Schwager, and F.

Jeltsch. 2004. Animal species diversity driven by habitat heterogeneity/diversity: The

importance of keystone structures. Journal of Biogeography. Vol. 31: 79-92.

Thiele, H.U. 1977. Carabid beetles in their environments: A study on habitat selection by

adaptation in physiology and behaviour. Springer-Verlag, New York.

XiaoDong, Y., L. TianHong, and Z. HongZhang. 2003. Species diversity of litter-layer

beetles in the Fengtongzhai National Nature Reserve, Sichuan Province. Acta

Entomologica Sinica. Vol. 46 (5): 609-616. 106

Table 1. Environmental variables* and their measurements taken at each site.

Site Bedrock UTM UTM Elevation Slope Aspect Soil Soil Soil Soil Soil Mean Grain Plant Plant % Plant Detritus Detritus Formation easting northing (ft) ° ° Temp Moisture Moisture pH Comp. Grain Sorting Species Cover Height % Cover Weight (°C) (Dry) (Wet) Strength Size Φ Richness (cm) (g) (kg/sq Φ cm) 1 Elbert 264181 4182693 10,369 ± 11 295 11.97 ± 16.43 ± 61.71 ± 6.24 ± 1.69 ± 3.23 2.17 6 50.63 ± 20.94 ± 14.22 ± 8.84 15 0.83 3.13 1.46 0.08 0.22 1.94 3.58 4.92 2 Elbert 264153 4182578 10,383 ± 0 0 14.69 ± 20.05 ± 63.33 ± 6.61 ± 2.66 ± 2.10 2.94 4 36.25 ± 11.83 ± 2.00 ± 25.29 24 0.50 2.76 1.92 0.07 0.35 1.02 0.51 1.00 3 Elbert 264159 4182514 10,405 ± 12 90 14.06 ± 29.95 ± 79.48 ± 5.93 ± 1.94 ± 0.93 3.10 10 65.94 ± 15.42 ± 5.44 ± 7.75 38 0.70 3.26 1.82 0.07 0.27 1.29 3.05 4.44 4 Elbert 264173 4182461 10,380 ± 0 0 12.56 ± 23.57 ± 70.38 ± 6.26 ± 1.73 ± 2.88 2.41 9 48.13 ± 17.75 ± 3.88 ± 2.22 17 0.43 3.46 1.94 0.09 0.29 2.31 2.27 2.88 5 Elbert 264209 4182398 10,395 ± 10 10 12.78 ± 37.35 ± 72.10 ± 6.39 ± 2.27 ± 3.07 2.76 9 51.88 ± 17.42 ± 8.06 ± 17.82 29 0.45 2.49 1.69 0.11 0.33 2.07 2.06 5.27 6 Ouray- 264042 4182650 10,439 ± 0 0 14.44 ± 28.50 ± 66.38 ± 6.39 ± 2.73 ± 2.23 2.37 8 56.88 ± 11.92 ± 19.63 ± 4.51 Leadville 33 0.37 1.89 2.03 0.06 0.29 2.63 1.01 8.67 7 Ouray- 264040 4182577 10,450 ± 7 321 11.81 ± 33.33 ± 66.38 ± 6.13 ± 1.52 ± 1.98 3.06 8 26.25 ± 14.58 ± 33.75 ± 32.15 Leadville 26 0.34 3.78 2.31 0.09 0.14 0.51 0.95 1.35 8 Ouray- 264088 4182506 10,438 ± 0 0 14.06 ± 31.19 ± 69.90 ± 6.43 ± 2.63 ± 2.32 2.44 8 83.13 ± 14.67 ± 28.88 ± 7.71 Leadville 18 0.35 2.60 1.80 0.13 0.21 3.29 1.44 11.98 9 Ouray- 264059 4182452 10,435 ± 15 95 17.11 ± 29.29 ± 72.90 ± 6.07 ± 2.15 ± -0.80 4.19 8 52.19 ± 17.83 ± 25.13 ± 59.1 Leadville 21 0.56 2.49 1.36 0.06 0.24 1.07 1.37 5.91 10 Ouray- 264048 4182375 10,480 ± 19 274 13.69 ± 10.80 ± 78.67 ± 6.06 ± 1.51 ± 2.92 3.10 9 38.13 ± 14.17 ± 11.72 ± 10.53 Leadville 22 0.63 2.02 2.44 0.16 0.27 4.13 0.29 4.01 11 Molas 263897 4182606 10,494 ± 0 0 13.81 ± 42.62 ± 80.86 ± 6.06 ± 1.90 ± 1.55 2.25 7 87.63 ± 11.08 ± 27.81 ± 10.8 25 0.29 2.15 1.87 0.15 0.22 1.47 0.89 5.24 12 Molas 263924 4182550 10,465 ± 7 329 16.50 ± 18.95 ± 59.05 ± 6.37 ± 2.67 ± 2.82 2.45 7 21.00 ± 14.50 ± 19.63 ± 18.66 38 1.00 3.16 1.56 0.09 0.37 1.16 1.85 3.14 13 Molas 263946 4182498 10,505 ± 14 122 18.31 ± 20.85 ± 60.57 ± 6.11 ± 1.92 ± 1.40 3.13 7 15.88 ± 15.25 ± 63.75 ± 11.27 30 0.51 3.27 1.24 0.08 0.21 0.99 0.45 2.70 14 Molas 263939 4182439 10,479 ± 0 0 14.03 ± 22.62 ± 56.00 ± 6.21 ± 2.02 ± 2.50 2.38 5 73.75 ± 9.75 ± 31.25 ± 6.48 19 0.65 1.84 1.03 0.12 0.12 1.84 0.82 4.18 15 Molas 263913 4182310 10,476 ± 5 100 13.86 ± 36.38 ± 84.10 ± 5.78 ± 0.99 ± 1.93 2.49 5 61.25 ± 15.92 ± 19.00 ± 4.5 13 0.24 2.49 2.28 0.06 0.09 0.51 1.32 2.59

*UTM are X and Y geographical coordinates Elevation ± number, represents the error range in elevation based off of the GPS unit. This is dependent on number of satellites, strength of signal. All other measurements with a reading ± number is the standard error Soil Moisture was divided up to represent the dry season (Dry) and monsoon season (Wet)

107

Table 2. Carabidae frequency data used for the CCA matrix.

Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Total Harpalus nigritarsis 6 9 5 10 11 8 3 6 10 6 2 13 5 10 10 114 Amara quenseli 8 7 6 8 6 8 9 3 11 11 1 12 8 4 0 102 Carabus taedatus 7 2 4 4 6 5 5 3 6 5 4 2 1 2 3 59 Amara laevipennis 0 4 7 3 0 4 0 9 7 1 1 0 1 3 1 41 Calathus ingratus 2 0 8 2 3 0 3 2 3 2 1 0 2 1 4 33 Cymindis cribricollis 0 2 1 1 1 3 5 2 4 1 1 2 1 1 2 27 Bembidion dauricum 3 1 7 1 3 0 4 0 0 0 0 0 0 0 4 23 Harpalus animosus 3 2 3 3 1 2 1 2 5 0 0 0 0 0 0 22 Bembidion mutatum 1 0 2 2 0 0 0 0 0 0 0 1 5 2 2 15 Harpalus laticeps 0 1 3 1 1 1 2 2 3 0 0 0 0 0 0 14 Harpalus opacipennis? 0 0 0 1 1 2 1 1 1 2 0 2 0 2 0 13 Agonum placidum 0 0 6 0 3 1 0 0 1 1 0 0 0 0 0 12 Notiophilus aquaticus 1 1 1 0 0 0 1 0 1 0 0 1 0 1 3 10 Amara ellipsis 0 0 1 1 0 0 0 1 3 2 1 0 0 0 0 9 Agonum cupreum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4 4 Amara merula 0 0 0 0 1 0 0 0 1 0 0 2 0 0 0 4 Pterostichus protractus 0 0 0 0 0 0 1 0 0 2 0 0 0 0 0 3 Amara (Bradytus) lindrothi 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 2 Amara littoralis 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 2 Calathus advena 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 2 Harpalus ellipsis? 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 2 Rhadine sp. 1 (nivalis grp) 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 2 Amara (Bradytus) sp. 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 1 Amara obesa 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 Amara sinuosa 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 Dyschirius dejeanii 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 1 Nebria catenata 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 Nebria gyllenhali 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 1 Pterostichus restrictus 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 Rhadine sp. 2? (nivalis grp.) 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 total 31 31 54 40 39 35 37 32 57 34 13 35 25 27 33 523

108

Table 3. P-values of the environmental factors and the corresponding carabid community and biodiversity measurements.

Environmental Variable Species Abundance Frequency Shannon- Evenness Richness Wiener Bedrock Formation 0.045 0.982 0.096 0.235 0.516 Elevation (ft) 0.110 0.999 0.091 0.102 0.203 Slope ° 0.999 0.374 0.271 0.847 0.777 Aspect ° 0.873 0.398 0.335 0.614 0.537 Soil Temp (°C) 0.291 0.209 0.793 0.124 0.137 Soil Moisture (Wet) 0.267 0.425 0.510 0.064 0.091 Soil pH 0.881 0.693 0.603 0.439 0.378 Soil Compressive Strength (kg/sq cm) 0.806 0.285 0.887 0.570 0.479 Mean Grain Size Φ 0.337 0.862 0.096 0.202 0.306 Grain Sorting Φ 0.080 0.603 0.010 0.710 0.782 Plant Species Richness 0.012 0.434 0.052 0.281 0.791 Plant % Cover 0.743 0.042 0.631 0.011 0.003 Plant Height (cm) 0.356 0.791 0.075 0.307 0.392 Detritus % Cover 0.164 0.490 0.163 0.346 0.578 Detritus Weight (g) 0.206 0.570 0.072 0.776 0.842

109

Table 4. CCA results of the first four axes for the environmental variables. Bold numbers represent the most important variable for that axis.

Variable AXIS 1 AXIS 2 AXIS 3 AXIS 4 Bedrock -0.328 -0.297 0.365 -0.248 Elevation -0.254 -0.265 0.199 -0.145 Slope -0.045 -0.231 -0.667 0.233 Aspect -0.065 -0.217 -0.636 -0.258 Soil Temp 0.081 -0.541 0.253 0.292 Soil Moist_D -0.382 0.612 0.266 0.022 Soil Moist_W -0.382 0.641 0.068 -0.053 pH 0.663 -0.124 0.082 -0.127 CS_Soil 0.684 -0.044 0.222 0.049 Grain_Size 0.041 -0.168 -0.225 -0.365 Grain_Sort 0.181 -0.060 -0.294 0.280 Plant_Spp 0.337 0.222 -0.465 0.340 Plant_Pc_Cover -0.052 0.679 0.583 0.000 Plant_Hgt -0.153 -0.032 -0.397 0.277 Detrit_Pc_Cover -0.142 -0.498 0.134 0.337 Detrit_Wgt 0.256 0.112 -0.179 -0.092 Eigenvalue % 18.68% 17.16% 13.00% 11.81% Total: 60.65%

110

Fig. 1. Topographic map of the study area and site locations.

16 y = 0.8468x + 4.5903 R² = 0.3983 14 12 10 8 6 4 2

Carabidae Species Richness 0 345678910 Plant Species Richness

Fig. 2. Linear regression analysis for carabid species richness versus plant species richness. 111

14 ab a 12

10 b 8

6

4

2 Carabidae Species Richness 0 Elbert Ouray-Leadville Molas

Fig. 3. Mean (± SEM) carabid species richness in bedrock formations.

300 y = -1.3568x + 151.95 R² = 0.2815 250

200

150

100

50 Carabidae Abundance 0 0 102030405060708090100 Percent Plant Cover

Fig. 4. Linear regression analysis for Carabidae abundance versus plant cover.

112

2.5 y = 0.0128x + 0.9971 R² = 0.4033 2.0

1.5

1.0

0.5 Shannon-Wiener Index Shannon-Wiener 0.0 0 102030405060708090100 Percent Plant Cover

Fig. 5. Linear regression analysis for Carabidae biodiversity versus plant cover.

1.0 y = 0.0054x + 0.4216 0.9 R² = 0.4995 0.8 0.7 0.6 0.5 0.4 Evenness 0.3 0.2 0.1 0.0 0 102030405060708090100 Percent Plant Cover

Fig. 6. Linear regression analysis for Carabidae evenness, versus plant cover.

113

60 y = 13.075x - 1.0819 R² = 0.4137 50

40

30

20

10 Carabidae Frequency

0 2.0 2.5 3.0 3.5 4.0 4.5 Sediment Sorting

Fig. 7. Linear regression analysis for Carabidae frequency versus sediment sorting.

114

1.6 1.2

0.8 11 0.4 3 8 7 0 15 5 9 126 Axis 2 Axis 10 -0.4 4 14 12 -0.8

-1.2 13 -1.6

-2 -2 -1.6 -1.2 -0.8 -0.4 0 0.4 0.8 1.2 1.6 Axis 1

Fig. 8. Sites and their distribution on the first and second axes for the CCA. Circle represents the 95% ellipse (similarity).

115

5 7 9 12 13 15 1 3 10 4 6 8 11 14 2 1 10 7 12 3 9 15 13 5 2 4 6 8 11 14

-60 0.95

-120 0.9

-180 0.85

-240 0.8

-300 0.75 Similarity Similarity

-360 0.7

-420 0.65

-480 0.6

-540 0.55

-600 0 1.6 3.2 4.8 6.4 8 9.6 11.2 12.8 14.4 16 0 1.6 3.2 4.8 6.4 8 9.6 11.2 12.8 14.4 16 a) Euclidian Cluster Analysis using Ward’s Algorithm b) Rho Cluster Analysis using Paired Group Algorithm

Fig. 9. Cluster analyses results of how sites are similar to each other based on all environmental variables.

116

0.64 Plant_Pc_Cover Moist_DMoist_W 0.48 LEGEND CCA Term Environmental Factor 0.32 Bedrock Bedrock Formation Elevation Elevation Slope Slope Plant_Spp Aspect Aspect 0.16 Temp Soil Temperature Moist_D Soil Moisture (Dry Season) Detrit_Wgt Moist_W Soil Moisture (Wet Season) 0 pH Soil pH Axis 2 Axis CS_Soil Soil Compressive Strength Plant_Hgt Grain_Size Mean Grain Size Grain_Sort CS_Soil Grain_Sort Grain Sorting Plant_Spp Plant Species Richness -0.16 pH Plant_Pc_Cover Plant % Cover Grain_Size Plant_Hgt Plant Height AspectSlope Detrit_Pc_Cover Detritus % Cover -0.32 BedrockElevation Detrit_Wgt Detritus Weight

-0.48 Detrit_Pc_Cover Temp -0.36 -0.24 -0.12 0 0.12 0.24 0.36 0.48 0.6 Axis 1

Fig. 10. CCA results* for the environmental variables, Axis 1 and 2.

*Longer the line (and more parallel) is with an axis, the more important that variable(s) is for structuring the carabid species composition. Axis 1 is most important followed by Axis 2.

117

Plant_Pc 0.48

Bedrock 0.32 LEGEND CCA Term Environmental Factor Temp Moist_D Bedrock Bedrock Formation 0.16 Elevation CS_Soil Elevation Elevation Slope Slope Detrit_Pc_Cover Aspect Aspect pH Moist_W Temp Soil Temperature 0 Moist_D Soil Moisture (Dry Season) Moist_W Soil Moisture (Wet Season) pH Soil pH

Axis 3 Axis CS_Soil Soil Compressive Strength -0.16 Grain_Size Mean Grain Size Grain_Sort Grain Sorting Detrit_Wgt Plant_Spp Plant Species Richness Grain_Size Plant_Pc_Cover Plant % Cover -0.32 Grain_Sort Plant_Hgt Plant Height Detrit_Pc Detritus % Cover Plant_Hgt Detrit_Wgt Detritus Weight -0.48 Plant_Spp

-0.64 SlopeAspect -0.48 -0.32 -0.16 0 0.16 0.32 0.48 0.64 Axis 2

Fig. 11. CCA results* for the environmental variables, Axis 2 and 3.

*Longer the line (and more parallel) is with an axis, the more important that variable(s) is for structuring the carabid species composition.

118

A_sinuosaP_restrictus

2.4 Rhadine_2? A_placidum 1.2 B_dauricum Bradytus_sp. H_laticeps D_dejeaniiC_advena A_ellipsisN_gyllenhali A_cupreum 11 H_animosusA_laevipennis C_ingratus 3 C_taedatus C_cribricollis8 Higher Plant Cover and Soil Moisture Soil and Cover Plant Higher 0 N_aquaticus15 57 9 P_protractus 124 106 Rhadine_1 H_nigritarsis14 12 A_quenseliH_opacipennis?H_ellipsis?A_obesa -1.2 A_merula Axis 2 Axis 13 N_catenata A_littoralis -2.4 B_mutatum -3.6

-4.8

-6 Lower Plant Cover and Soil Moisture Soil and Cover Plant Lower -7.2 B_lindrothi -7.2 -6 -4.8 -3.6 -2.4 -1.2 0 1.2 2.4 Lower Soil Compressive Strength and pH Axis 1 Higher Soil Compressive Strength and pH

Fig. 12. CCA results for Carabidae species and how they plot on Axis 1 and 2. For each species, Genus is initialized except for

Rhadine and species is fully written out. The most important environmental factors are shown for each axis and quadrant.

119

Rhadine_2? 4 P_restrictuA_sinuosa s

3 A_cupreum Rhadine_1 2 A_littoralis A_laevipennis N_catenataH_ellipsis? Eastern Aspect or None and No Slope No and None or Aspect Eastern N_gyllenhali 1 8 11 2 15 B_mutatum 14 4 N_aquaticus A_ellipsis 13 H_nigritarsis6 9 H_animosus 12 C_cribricollis 0 3 H_laticeps Axis 3 3 Axis B_lindrothi H_opacipennis? 101 57 C_taedatus A_quenseli C_ingratus -1 A_merula B_dauricum -2 A_placidum

-3 A_obesa P_protractus -4 D_dejeanii C_advena Bradytus_sp.

Northernand Western Aspect and Sloped -5 -7.2 -6 -4.8 -3.6 -2.4 -1.2 0 1.2 2.4 Lower Plant Cover and Soil Moisture Axis 2 Higher Plant Cover and Soil Moisture

Fig. 13. CCA results for Carabidae species and how they plot on Axis 2 and 3. For each species, Genus is initialized except for

Rhadine and species is fully written out. The most important environmental factors are shown for each axis and quadrant. 120

CHAPTER IV

Coleopteran Biodiversity within the Sultan Creek-Molas Lake Study Area in the San Juan

Mountains, San Juan County, Colorado

Only a few insect studies have been performed within the San Juan Mountains. One

paper was found on a survey performed by the Corps of Engineers, specifically for the San Juan

Mountain region in 1878, that included flora and fauna collections in which Coleoptera was one

of the main insects collected (Strecker 1878). No locality information was given for the beetles

collected, only for the other orders in the report. A few other papers list localities of beetles collected within the San Juan Mountains (Wickman 1902, Elias 1987, and Kippenhan 1990), but these articles were synthesis articles on multiple collections and surveys performed throughout

Colorado. The ecological insect studies that have been performed in this region were related to the spruce beetle outbreaks (Baker and Veblen 1990 and Ryerson et al. 2003) or the

Uncompahgre fritillary butterfly (Britten et al. 1994, Britten and Riley 1994). Other insect studies incorporating the San Juan Mountains for broader scale analyses were also found in the literature (DeChaine and Martin 2008 and Worrell et al. 2008). The main objective of this study was to perform a general biodiversity assessment of the soil-dwelling coleopteran fauna of a meadow in the San Juan Mountains and to note any general trends with their abundance.

Materials and Methods

Site Location. The field site was located in San Juan County, several kilometers south of

Silverton, a small mining town in southwest Colorado. The study area itself is an open meadow 121

with remnants of the spruce-fir forest with a mean elevation of 3,170 m (10,400 ft). The

meadow’s topography is made up of gently sloping ridges and valleys. It is just south of Sultan

Creek and about a kilometer north of Molas Lake on the east side along Highway 550 (Fig. 1).

The whole study area was approximately 400 m X 400 m.

Insect Collection. The beetles were caught via pitfall trapping. Nine pitfall traps were placed

per site in a 5 m X 5 m grid. A total of fifteen sites were set up. The sites were arranged into three transects with five sites each. Each site was at least 50 m apart from another site. Trapping was done each week for eight weeks total in 2006 (June 17 to August 11) and six weeks total in

2007 (June 24 to August 3). Traps were kept open for 72-120 hours. A 50:50 solution of propylene glycol:water was used to kill and preserve insects. A small amount of dish soap was added into the solution to lower surface tension. Also, quinine was added to deter mammals from eating the insects. Traps were filled up to the 75-100 ml mark in a 250 ml cup (Corning

Snap-Seal no. 1730 design). The cup dimensions were 7.5 cm height and 7.6 cm diameter. Cups were left with no covering for most of the study period in 2006. During the last few weeks, ite six had to be covered with rocks due to juvenile marmots digging up the cups and emptying the contents out. In 2007, traps were left with no covering throughout the summer, but some of the cups had chicken wire over them to prevent them from popping out of the ground. Traps were cleaned out and insects stored in 70% ethanol alcohol. The beetles were either pinned or stored in alcohol vials after sorting through and separating them out into morphospecies. Identification was performed by the author through guidebooks, internet websites, and museum collections at

The University of Colorado and Colorado State University, by specialists at the museums, or sent to other specialists through the mail. The following people helped identify the beetles: Daniel

Duran (Cicindelidae; Ph.D. student currently at Vanderbilt University), Bill Warner 122

(Scarabaeidae; Farnam Companies, INC), Dan Pavuk (Michigan State University-The Ohio State

University Extension), Donald E. Bright (Curculionidae and Scolytidae; Colorado State

University), Boris Kondratieff (Colorado State University), Virginia Scott (University of

Colorado), Shawn M. Clark (Chrysomelidae and several uncommon families; Brigham Young

University), Paul J. Johnson (Byrrhidae; South Dakota State University), Stewart Peck

(Leiodidae; Carleton University, Canada), Sam Wells (Elateridae; Bayer CropScience LP,

Fresno, CA), Bob Davidson (Carabidae; Carnegie Museum of Natural History), and Foster

Purrington (Carabidae; The Ohio State University).

Biotic and Physical Variables Measured. Latitude, Longitude, elevation and aspect were measured with a GPS unit for each site. The slope was estimated for each site with a Brunton compass and board. The abiotic soil factors measured were: temperature, pH, moisture, compressive strength, mean grain size, and grain size sorting. The biotic soil factors measured were: plant cover, plant height, plant species richness, detritus cover and detritus weight. In depth methodology of how these variables were gathered can be found in Chapter One.

Statistical Analyses. Total number of families, potential species, and abundance were calculated for the study area. General statistical analyses were performed on coleopteran abundance only to note any general trends due to the biological and physical variables collected for the 2006 and 2007 seasons. Linear regression analyses and one-way ANOVAs were performed depending if the variable was continuous or discrete. Site six was either completely removed from the analyses or weeks five through eight omitted for the 2006 analyses due to juvenile marmots destroying most of the site later in the season. Site three, week five was also removed in some of the analyses in 2006 due to a high emergence of two Scarabaeidae species. 123

In 2007, week four, site six, weeks five and six, sites seven and thirteen, were omitted in the analyses on weekly measured variables due to a high emergence of Staphylinidae.

Results

Coleopteran Biodiversity. A total of over 93,000 insects were collected in the two seasons.

Coleopteran abundance from this collection accounted for 7,316 specimens (adult and larvae).

In 2006, 3,726 beetles were collected whereas in 2007, 3,590 collected. At least 27 families

were identified with several specimens still unknown (Table 1). Over ninety-four percent of the

beetle fauna was composed of nine families (Staphylinidae, Carabidae, Curculionidae,

Chrysomelidae, Cicindelidae, Scarabaeidae, Mordellidae, Elateridae, and Byrrhidae) and beetle

larvae (5.2%). Seven of the nine top families were the highest in abundance in both years. Out

of the remainding 5.6% of the beetles, 2.8% were unidentified or unaccounted for. The other

2.8% of the beetles were composed of 18 families, with Scolytidae and Leiodidae accounting for

half of this percent. Most likely, some of the specimens unidentified may fall into these families

or others not on the list. An interesting observation throughout the seasons was with Family

Leiodidae. Leiodidae only occurred in the second half of the season in both years (weeks four

through eight in 2006, and weeks five and six in 2007).

Sixty-one species were completely identified with a total of ninety-four separated

correctly into morphospecies (Table 2). Amara quenseli quenseli and Cicindelida longilabris

were the most common species identified. There were also one or two staphylinid species that

were highly abundant (hundreds of individuals), but difficult to separate into morphospecies.

Other quite common species (> 100 individuals) in the meadow were: Panascopus sp. 1,

Flaviellus subtruncatus, Harpalus nigritarsis, Hypnoidus bicolor, Otiorhynchus ovatus, 124

Agoliinus canadensis, Otiorhynchus sulcatus, and Pachybrachis hepaticus hepaticus. Also, several other species were very abundant but were not identified down to species: one species of

Mordellidae, one species of a Chrysomelidae (Alticinae), and a few Staphylinidae species. A few interesting observations follow for some of the species caught. Hypnoidus bicolor was highly abundant in the Elbert and Molas Formations, but was extremely rare in the Ouray-

Leadville Formations with only six specimens caught. Panascopus sp. 1 was very common in the meadow, but was not found in any of the Museum collections visited in Colorado. Currently an expert is identifying this species. Panascopus sp. might be a rare species common at my site, a new state or county locality for the species, or possibly a new species. Flaviellus subtruncatus was quite common in the meadow but is considered very rare in collections (B. Warner pers. comm.). It was very common in the Elbert and Ouray-Leadville Formations, but rarely appeared in the Molas Formation (one specimen). Also, Agoliinus canadensis and Aegialia lacustris were found often in the meadow and are considered uncommon and/or unusual (B. Warner pers. comm.). Finally, the Rhadine species might be a new species since this genus is highly speciose

(R. Davidson pers. comm.).

Coleopteran Abundance Trends. In the 2006 season, the weekly total coleopteran abundance represented a typical bell-shaped curve throughout the season, low abundance in the early and late season whereas high abundance in the mid-season (Fig. 2). Plant cover, plant height and the week collect took place showed significant trends with coleopteran abundance in 2006 (Table 3).

Early season and end of the season mean coleopteran abundance were significantly lower than midway through the season (Fig. 3). Even though coleopteran abundance between sites was not

significantly different overall (Table 3), site 11 was significantly lower in abundance than site 12 125

(Fig. 4). Coleopteran abundance generally increased with lower percent plant cover and taller plants (Figs. 5 and 6).

In 2007 season, a near opposite trend occurred with coleopteran abundance caught per week, because high abundance was found in the early and end of the season with low abundance midway (Fig. 7). Plant cover, soil moisture, aspect, slope, collection site, and week collecting took place were significant for coleopteran abundance in 2007 (Table 4). During the season, coleopteran abundance caught in week three was significantly lower from week five (Fig. 8).

Site comparisons showed that site five was significantly higher than site 14 (Fig. 9). Also, half of the sites (1, 2, 3, 6, 8, 13 and 15) were lower in abundance when compared to site five (p = <

0.1, sites 10 and 11 had p-values of 0.054 and 0.053 respectively). Coleopteran abundance generally increased with lower plant cover and higher soil moisture (Figs. 10 and 11). Overall seasonal coleopteran abundance and seasonal moisture levels showed a strong positive correlation (Pearson’s = 0.921) with a p-value of 0.009 (F = 22.35, df = 1, 4) (Fig. 12). Also, north facing slopes had the highest mean abundance and was significantly higher from sites with no aspect (Fig. 13). Finally, slightly sloped sites (1-10°) had the highest mean abundance and were significantly higher than flat sites (Fig. 14).

Discussion

Coleopteran Biodiversity. This small scale study yielded a surprisingly large number of insects and diverse beetle fauna with using only pitfall traps. An estimate of possibly 120-130 species may occur in this meadow (based on the unknown families with specimens and potential number

of staphylinid species unidentified). A study performed in the Alberta Mountains within the

Willmore Wilderness Park produced 36 families and 318 species (Hilchie 2008). Most of these 126

sites were in the alpine and subalpine zones. This study was also on a larger scale and used

multiple collecting methods then this current study (e.g., pitfalls, pan, light traps, hand

collecting). Even though these two studies were at different scales, my small site led to 27

families being identified (possibly a few more) with just one trapping method.

This study was probably the first systematic collection of beetles within the San Juan

Mountains, especially in San Juan County, that includes site locations and multiple environmental variables. There might be new records for the county, Colorado, and the mountain region. As mentioned previously, I collected some species that are typically rare, and I also collected individuals that belong to families that have few entomologists working on them

(e.g., Byrrhidae and Leiodidae). Even well studied beetles, like the Cicindelidae, might lead to a

new county record. Cicindelida longilabris may be a new record, for San Juan County since the

county is not mentioned in Kippenhan’s (1990) survey article for this species. These species will

have to be checked out quite thoroughly with articles and museum collections to find out if they

are indeed new locality records. I may also have a few new species altogether which will need to

be sent off to taxonomic specialists for positive identifications.

Coleopteran Abundance Trends. Overall, beetle abundance was significantly affected by

aspect, plant cover, plant height, slope, soil moisture, site location, and week. Another study on

soil dwelling insects also showed a wide variety of variables affecting abundance: rainfall, soil

moisture, organic matter, soil and air temperatures, soil pH, relative humidity at the soil surface,

and potassium and phosphorus concentrations of the surface soil (Reddy and Venkataiah 1990).

Usually, general trends in abundance are seen with temperature and moisture, yet this was

generally not the case in my study. Soil moisture was significant in 2007, yet the R2 was only

6%, quite low due to a large amount of variation present in the data. The only strong 127

significance (R2 = 84%) occurred when the abundance was analyzed seasonally (per week). Soil

temperature had no affect on the abundance, although the air temperature was not analyzed.

Plant cover consistently affected abundance in both years. General trends in abundance

increased with less plant cover. Lower plant cover provides less resistance for soil dwellers to

move on the ground. Several of the most abundant families caught live specifically on the

ground and are quite active in low vegetation (Carabididae, Cicindelidae, Scarabaeidae, and

Staphylinidae). However, quite a few plant feeders were caught (Chrysomelidae and

Curculionidae) which may have lessened the strength of this trend (R2 = 11% in 2006 and R2 =

28% in 2007). Future analyses should be run on soil dwellers only and on specific families to

find trends.

Significant seasonal changes of the coleopteran abundance also occurred in both years

and were expected due to seasonality changes in insect communities due to emergences and die-

off/hibernation/migration. However, the change of pattern of coleopteran abundance itself was

opposite between seasons and quite unusual in year 2007. Year 2006 showed a typical bell

shaped curve with abundance. Usually in a season, low abundance occurs with an increase in the

middle season, than it drops again. The year 2007 has the opposite pattern. This might be

explained due to the snow season and later start of the monsoon rains. The 2006-2007 winter

had very late season snowfalls. The snow melt from this season may have influenced insect

emergences to occur earlier and peak when I started the trapping season. Then, due to the drought and potential initial emergence, abundance dropped off mid season. This definitely needs more testing with longer seasonal collecting of coleopterans and other insects, and multi- year analyses. Also, once the NOAA (National Oceanic and Atmospheric Administration) 128 website updates their historical records for Silverton (only goes up to 2005), the seasonal abundance trends can be studied with the precipitation and temperature data.

Conclusion

In conclusion, this study was possibly the first systematic coleopteran survey performed in the San Juan Mountains. A diverse fauna was collected in a relatively small sampling area, which yielded nearly a hundred species identified in twenty-seven families. Some of these species may possibly be new records for the county and the mountain region. Plant cover, plant height, soil moisture, aspect, slope, collection site, and week had significant effects on beetle abundance. These effects were different depending on year. Other factors not studied might be influencing abundance and should be analyzed with future studies (e.g., rainfall, snow season).

Future studies should also be performed in the rest of the open meadow and surrounding habitats in the Molas Lake region with multiple trapping methods to sample the overall coleopteran biodiversity and not just soil dwellers.

129

References

Baker, W.L. and T.T. Veblen. 1990. Spruce beetle outbreaks and fire in the nineteenth

century subalpine forests of western Colorado, U.S.A. Arctic and Alpine Research. Vol.

22: 65-80.

Britten, H.B., P.F. Brussard, and D.D. Murphy. 1994. The pending extinction of the

Uncompahgre fritillary butterfly. Conservation Biology. Vol. 8 (no. 1): 86-94.

Britten, H.B. and L. Riley. 1994. Nectar source diversity as an indicator of habitat suitability

for the endangered Uncompahgre fritillary, Boloria acrocnema (Nymphalidae). Journal

of the Lepidopterists Society. Vol. 48 (no. 3): 173-179.

DeChaine, E.G. and A. P. Martin. 2008. Historic cycles of fragmentation and expansion in

Parnassius smintheus (Papilionidae) inferred using mitochondrial DNA. Evolution. Vol.

58 (1): 113-127.

Elias, S.A. 1987. Colorado ground beetles (Coleoptera: Carabidae) from the Rotger Collection,

University of Colorado Museum. Great Basin Naturalist. Vol. 47 (4): 631-637.

Hilchie, G.J. 2008. A survey of the beetles (Coleoptera) in the western portion of Willmore

Wilderness Park. Prepared for Parks Resource Management Coordination Branch

Alberta Tourism, Parks and Recreation.

Kippenhan, M.G. 1990. A survey of the tiger beetles (Coleoptera: Cicindelidae) of Colorado.

Entomological News. Vol. 101 (5): 307-315.

Reddy, M.V. and B. Venkataiah. 1990. Seasonal abundance of soil-surface arthropods in

relation to some meteorological and edaphic variables of the grassland and tree-planted

areas in a tropical semi-arid savanna. International Journal of Biometeorology. Vol. 34:

49-59. 130

Ryerson, D.E., T.W. Swetnam, and A.M. Lynch. 2003. A tree-ring reconstruction of western

spruce budworm outbreaks in the San Juan Mountains, Colorado, U.S.A. Canadian

Journal of Forest Research/Revue Canadienne de Recherche Forestiere. Vol. 33 (no. 6):

1010-1028.

Strecker, H. 1878. Section II: Lepidoptera. In Annual Report upon Explorations and Surveys

in the Department of the Missouri being Appendix SS of the Annual Report of the Chief

of Engineers for 1878, by E.H. Ruffner. 1847-1867.

Worrall, J.J., L. Egeland, T. Eager, R.A. Mask, E.W. Johnson, P.A. Kemp, and W.D.

Shepperd. 2008. Rapid mortality of Populus tremuloides in southwestern Colorado,

USA. Forest Ecology and Management. Vol. 255: 686-696.

Wickman, H.F. 1902. The Coleoptera of Colorado. Bulletin of the State University of Iowa.

No. 5 (3): 217-310.

131

Table 1. Coleopteran families identified and the total number of individual specimens caught in each family at the Sultan Creek-Molas Lake study area.

Family 2006 2007 Total Percent Staphylinidae 386 1691 2077 28.39 Carabidae 992 244 1236 16.89 Curculionidae 342 435 777 10.62 Chrysomelidae 377 262 639 8.73 Cicindelidae 170 397 567 7.75 Scarabaeidae 443 107 550 7.52 Beetle Larvae 268 109 377 5.15 Mordellidae 200 60 260 3.55 Elateridae 212 46 258 3.53 Byrrhidae 137 31 168 2.30 Unaccounted for 24 88 112 1.53 Unknown 45 48 93 1.27 Scolytidae 35 28 63 0.86 Leiodidae 43 8 51 0.70 Cantharidae 14 16 30 0.41 Ptilidae 11 7 18 0.25 Cerambycidae 6 1 7 0.10 Coccinellidae 4 2 6 0.08 Buprestidae 1 4 5 0.07 Nitidulidae 4 1 5 0.07 Meloidae 1 3 4 0.05 Artematopidae 2 1 3 0.04 Clerididae? 2 0 2 0.03 Melyridae 1 1 2 0.03 Anobiidae 1 0 1 0.01 Cucujidae 1 0 1 0.01 Latridiidae 1 0 1 0.01 Melandryidae 1 0 1 0.01 Salpingidae 1 0 1 0.01 Silphidae 1 0 1 0.01

132

Table 2. Species identified and unidentified within each family at the Sultan Creek-Molas

Lake study area.

Family Species Abundance Anobiidae Species 1 1 Artematopidae Species 1 3 Buprestidae Chrysobothris breviloba 5 Byrrhidae Arctobyrrhus subcanus 55 Byrrhus cyclophorus 13 Byrrhus kirbyi 9 Morychus aeneolus 90 Porcinolus undatus 1 Cantharidae Podabrus sp. 1 14 Podabrus sp. 2 16 Carabidae Agonum cupreum 4 Agonum placidum 26 Amara ellipsis 10 Amara laevipennis 69 Amara (Bradytus) lindrothi a 2 Amara (Bradytus) sp. a 1 Amara littoralis 2 Amara merula 4 Amara obesa 1 Amara quenseli quenseli 561 Amara sinuosa 1 Bembidion dauricum 41 Bembidion mutatum 23 Calathus advena 2 Calathus ingratus 50 Carabus taedatus 71 Cymindis cribricollis 31 Dyschirius dejeanii 1 Harpalus animosus 24 Harpalus ellipsis? b 2 Harpalus laticeps 16 Harpalus nigritarsis 258 Harpalus opacipennis? b 14 Nebria catenata 4 Nebria gyllenhali 1 Notiophilus aquaticus 10 Pterostichus protractus 3 Pterostichus restrictus 1 Rhadine sp. 1 (nivalis grp) c 2 133

Rhadine sp. 2 (nivalis grp) c 1 Cerambycidae Gnathacmaeops pratensis 2 Neanthophylax mirificus 1 Pachyta lamed liturata 3 Species 1 1 Chrysomelidae Altica sp. 1 23 Altica sp. 2 68 Chaetocnema perturbata 1 Coleothorpa vittigera 11 Galeruca rudis 2 Graphops sp. 1 67 Longitarsus sp. 1 2 Pachybrachis hepaticus hepaticus 148 Phyllotreta cruciferae 1 Phyllotreta (one or more species) 4 Psylliodes sp. 1 3 Subfamily Alticinae (unidentified) 296 Unknown 13 Cicindelidae Cicindelida longilabris 567 Clerididae? Unknown 2 Coccinellidae Coccinella transversoguttata 2 Hippodamia convergens 3 Hippodamia signata 1 Cucujidae Species 1 1 Curculionidae Apion (one or more species?) 26 Ceutrorhynchus sp. 1 3 Otiorhynchus ovatas 175 Otiorhynchus sulcatus 149 Panscopus sp. 1 374 Peritaxia rugicollis 13 Species 1 2 Species 2 1 Species 3 1 Unknown 33 Elateridae Dalopius sp. 15 Hypnoidus bicolor 243 Latridiidae Species 1 1 Leiodidae Leiodes collaris 20 Leiodes longitarsus 23 Species 1 5 Species 2 3 Melandryidae Species 1 1 Meloidae Meloe laevis 4 Melyridae Species 1 2 Mordellidae multiple species? 260 134

Nitidulidae Meligethes sp. 5 Ptilidae Species 1 18 Salpingidae Species 1 1 Scarabaeidae Aegialia lacustris 94 Agoliinus canadensis 150 Dialytodius decipiens 1 Diplotaxis haydenii 35 Flaviellus subtruncatus 270 Scolytidae Crypturgus borealis 2 Hylastes gracilis 2 Pityokteines lasiocarpa 2 Pityophthorus sp. 1 (possibly absorus) 1 Pityophthorus sp. 2 1 Unknown 55 Silphidae Thanatophilus lapponicus 1 Staphylinidae Staphylinus sp. 1 3 Stenus sp. 1 39 Unknown 2035 Beetle Larvae 377 Unidentified Specimens d 93 Unaccounted For e 112 a Might be the same species b Might be the same or separate species c Might be the same species d Most are small beetles in vials that are difficult to key by the author e In 2006, most likely minor counting off with Staphylinids and Chrysomelids since author didn't write down number in vial In 2007, most likely a Staphylinid and/or Chrysomeldid vial(s) unaccounted for since large emergence of Staphs

135

Table 3. ANOVA and regression analyses performed for the 2006 variables against coleopteran abundance.

Variable P F DF Detritus Cover 0.743 0.11 1,40 Elevation 0.360 0.91 1,12 Wet ph 0.563 0.35 1,12 Plant Cover 0.028 5.20 1,40 Plant Height 0.007 8.08 1,40 Plant Species 0.291 1.22 1,12 Soil Compressive Strength 0.273 1.23 1,40 Soil Moisture 0.146 2.14 1,113 Soil Size 0.376 0.84 1,12 Soil Sorting 0.228 1.62 1,12 Soil Temperature 0.134 2.28 1,113

Aspect 0.448 0.96 3,10 Bedrock 0.723 0.33 2,11 Site 0.221 1.30 14,100 Slope 0.409 0.97 2,11 Week 0.000 16.36 7,107

136

Table 4. ANOVA and regression analyses performed for the 2007 variables against coleopteran abundance.

Variable P F DF Detritus Cover 0.683 0.17 1,13 Detritus Weight 0.265 1.36 1,13 Elevation 0.891 0.02 1,13 pH 0.198 1.86 2,12 Plant Cover 0.043 5.03 1,13 Plant Height 0.293 1.20 1,13 Plant Species Richness 0.470 0.55 1,13 Soil Compressive Strength 0.811 0.06 1,13 Soil Moisture 0.024 5.32 1,83 Soil Size 0.843 0.04 1,13 Soil Sorting 0.425 0.68 1,13 Soil Temperature 0.068 3.43 1,83

Aspect 0.006 7.15 3,11 Bedrock 0.924 0.08 2,12 Site 0.002 2.81 14,70 Slope 0.045 4.08 2,12 Week 0.004 3.77 5,79

137

Fig. 1. Topographic map of the study area and site locations.

900 800 700 600 500 400 300 200

Coleoptera Abundance 100 0 12345678 Week

Fig. 2. Total coleopteran caught each week in 2006 season. 138

60 b b 50 b b 40

30 a 20 a a a 10

Mean Coleopteran Abundance Mean Coleopteran 0 12345678 Week

Fig. 3. Mean (± SEM) coleopteran abundance caught each week in 2006 season.

60 * 50

40

30

20 *

10

Mean Coleopteran Abundance 0 123456789101112131415 Site

Fig. 4. Mean (± SEM) coleopteran abundance caught at each site in the 2006 season.

* The ANOVA did not show a significant value overall, but sites 11 and 12 were significantly different from each other, p-value of 0.013. 139

y = -0.3185x + 44.825 100 R² = 0.115 90 80 70 60 50 40 30

Coleoptera Abundance 20 10 0 0 20406080100 Percent Plant Cover

Fig. 5. Linear regression of coleopteran abundance for selected weeks that percent plant cover was measured in 2006.

100 y = 2.5334x - 8.3876 R² = 0.1681 90 80 70 60 50 40 30 20 Coleoptera Abundance 10 0 5 10152025 Plant Height (cm)

Fig. 6. Linear regression of coleopteran abundance for selected weeks that plant height was measured in 2006. 140

1100 1000 900 800 700 600 500 400 300 200 Coleoptera Abundance 100 0 123456 Week

Fig. 7. Total coleopteran caught each week in 2007 season.

60 b 50 ab 40 ab

ab 30 ab 20 a

10

Mean Coleopteran Abundance Mean Coleopteran 0 123456 Week

Fig. 8. Mean (± SEM) coleopteran abundance caught each week in 2007 season.

141

80 70 60 50 40 30 20 10 Mean Coleopteran Abundance 0 123456789101112131415 Site

Fig. 9. Mean (± SEM) coleopteran abundance caught at each site in the 2007 season.

* Site 5 is significantly different from site 14, p = 0.028

800 y = -4.5258x + 461.1 R² = 0.2792 700 600 500 400 300 200

Coleoptera Abundance 100 0 0 20406080100 Percent Plant Cover

Fig. 10. Linear regression of seasonal coleopteran abundance for percent plant cover in

2007. 142

120 y = 0.192x + 21.734 R² = 0.0602 100

80

60

40

20 Coleopteran Abundance 0 0 20406080100 Soil Moisture

Fig. 11. Linear regression of coleopteran abundance for weekly soil moisture content measured in 2007.

1200 y = 11.483x + 148.09 R² = 0.8399 1000

800

600

400

200 Coleopteran Abundance 0 0 20406080 Soil Moisture

Fig. 12. Seasonal correlation trend between coleopteran abundance and soil moisture levels in 2007.

143

700 b 600 500 400 300 ab

200 a ab 100 0 Mean Coleoptera Abundance Mean Coleoptera flat/none north east west Aspect

Fig. 13. Mean (± SEM) coleopteran abundance in different aspects in 2007.

600 b 500

400

300 ab

200 a 100

0 Mean Coleoptera Abundance Mean Coleoptera flat/none 1-10° 11-20° Slope

Fig. 14. Mean (± SEM) coleopteran abundance in different slopes in 2007 season.

144

GENERAL SUMMARY AND FUTURE STUDIES

One of the main objectives of this study was to catalog the coleopteran fauna of a subalpine environment within the San Juan Mountains of Colorado. This was the first systematic insect biodiversity study performed in a part of the San Juan Mountains (San Juan County). A diverse fauna was collected in a relatively small sampling area. The coleopteran abundance was

7,316 individuals, with 27 families, and nearly 100 species in which 61 species were completely identified. Some of these species may possibly be new records for the county, state, and this mountain region. The coleopteran abundance was analyzed against multiple environmental variables and was influenced by plant cover, plant height, soil moisture, aspect, slope, collection site, and week.

The second main objective was to analyze how the physical environment, especially the geology of the study area, influenced the community of the ground dwelling beetles. Three hypotheses were developed to help answer this. These are re-stated here and addressed, summarizing what was found.

Hypothesis 1: Geology influences the soil properties.

Elevation, detritus cover, and soil temperature were significantly distinct between formations. Temperature was the only soil parameter different between bedrock formations.

The soil temperature effects were most likely influenced by the sequence of collecting the measurements. The elevational differences were based on the geological history and depositional sequence of the formations (age). Detritus cover was an interesting difference and no known cause at this site was studied (e.g., microbrial, chemical). Other factors that tend to be 145 associated with geology that were not significantly different may have to do with using only sedimentary rocks or other factors (e.g., vegetation effects, rate of soil development). A few of these factors, specifically soil pH and sediment size and sorting, were not significantly different between formations, but were most likely affected by the geology (e.g., hydrothermal alteration, sharing a range of rock grain size types, like shale and siltstones).

Hypothesis 2: Bedrock formation influences the community structure of the Carabidae.

Carabid species richness was influenced by bedrock formation. Several species abundances were also influenced by bedrock formation (Agonum placidum, Bembidion mutatum,

Carabus taedatus, Cymindis cribricollis, Harpalus animosus, and Harpalus laticeps). The

Carabidae did not separate out into distinct communities based on bedrock formation.

Geological affects were more individualized due to a particular species instead on the community as a whole.

Hypothesis 3: Bedrock has a stronger effect on the Carabidae community and composition than other environmental factors.

Carabid community structure was mainly affected by botanical factors. Carabid species richness, abundance, and biodiversity were mainly influenced by plant factors whereas carabid frequency was influenced by sediment sorting. The carabid species composition based on CCA was influenced by compressive strength, pH, plant cover, soil moisture, slope, and aspect. 146

Overall, geology was less of a factor in the community structure (only affected species richness) and composition (sixth most important variable out of 16 variables on Axis 1 on the CCA).

Future studies should incorporate the rest of the meadow (Fig. 1) and a few other localities in which the bedrock formations occur in. Other units are also in the vicinity: metamorphic and igneous rocks (gneiss, quartzite, schist, slate, granite, igneous intrusion), other sedimentary rocks, and glacial deposits. The Carabidae community and environmental factors can be studied on a larger geographical area, incorporating multiple habitat and geological types.

Differences in the carabid community and environmental factors may be occurring at a different scale than formation: from localized bedding planes (e.g., shale, sandstone, conglomerate) to general rock type (igneous, metamorphic, and sedimentary). Also, specific species analyses need to be performed to find out what factor(s) are controlling why a species is found in specific formations (geological aspect or environmental).

The coleopteran fauna as a whole also needs more future work. Multiple trapping methods should be incorporated in the meadow to sample the overall coleopteran biodiversity and not only soil dwellers. Surveying in multiple habitats in the vicinity will also shed light on the diversity of the beetles living in this region. Also, other factors not studied for coleopteran abundance trends should be analyzed (e.g., rainfall, snow season).

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Fig. 1. Whole meadow in the Molas Lake vicinity.