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Bell & Howell Information and Learning 300 North Zeeb Road, Ann Artxjr, Ml 48106-1346 USA 800-521-0600 UMT
GROUND BEETLE ABUNDANCE AND DrVERSUY PATTERNS
\%TTHIN MDŒD-OAK FORESTS SUBJECTED TO PRESCRIBED BURNING
IN SOUTHERN OHIO
DISSERTATION
Presented in Partial Fulfillment of the Requirements for
the Degree Doctor of Philosophy in the Graduate
School of The Ohio State University
By
Robert Christopher Stanton, B.A., M.S. *****
The Ohio State University
2000
Dissertation Committee: iroyed by Dr. David J. Horn, Advisor
Dr. Ralph EJ. Boemer Advisor Dr. Norman F. Johnson
Dr. Deborah H. S tinner Department of Entomology UMI Number 9982983
UMI*
UMI Microform 9982983 Copyright 2000 by Beil & Howell Information and teaming Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code.
Bell & Howell Information and Learning Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor, Ml 48106-1346 ABSTRACT
The deciduous forests of the eastern United States have been dominated by oak
(Quercus) species for the past 10.000 years. However, over the last 50 years the oak component of the regeneration of many of these forests has declined dramatically.
Prescribed burning may improve oak regeneration, but the effects of this practice on other aspects of these communities are largely unknown.
To better understand the ecological response of oak-dominated communities to prescribed burning, the USD A Forest Service initiated an ecosystem management project in 1994 in southern Ohio. Four study sites in Vinton and Lawrence counties were subdivided into three treatment units. The treatments were “frequent” fire (burned annually), “infrequent” fire (burned every third year), and “control” (not burned). Fires were set during the spring (dormant season) of 1996 through 1999.
In order to investigate the impacts of prescribed burning on surface-active invertebrates, ground beetles (Coleoptera: Carabidae) were monitored via pitfall and light trapping along dry ridge tops. Carabids are well suited as indicators of environmental conditions and are often used to monitor changes within a community.
Overall carabid abundance and flightless carabid abundance recorded by pitfall trapping decreased during the course of this study, but there were no significant changes related to prescribed burning. Seasonal trends in carabid abundance also were not significantly affected by prescribed burning.
No significant effects of prescribed burning were detected on carabid species
richness, diversity, or evenness. Year-to-year changes in these indices were related to
changes in one very abundant species {Synuchus impunctatus) and to the low number of
carabids collected in 1999.
Comparisons of the carabid species collected at each study site via pitfall trapping
suggested that distinct assemblages existed in Vinton County, northern Lawrence County,
and southern Lawrence County. These different assemblages were most likely related to
differences in soil characteristics and understory vegetation among these three areas.
Results from other aspects of this research showed that prescribed burning alters
the local environment in areas such as soil chemistry, leaf litter biomass, and understory
vegetation. Despite these changes, this study indicated that low intensity spring fires do
not significantly impact surface-active carabid populations on xeric, upland sites in oak-
dominated forests.
This research project allowed forest carabid populations to-be monitored for five
consecutive years. This information is important to better understanding carabid
communities in deciduous forests and their response to prescribed burning management
practices.
lU Dedicated to tny students—past and future:
You have been a constant source of encouragement and hope.
IV ACKNOWLEDGMENTS
I would lika to first acknowledge Ralph Boemer, Norm Johnson, Deb Stinner, and
Dave Horn for serving on my committee. They held me to higher standards and pushed me closer to my full potential. I would especially like to thank Dave Horn for being patient and understanding with all of my last-minute activities.
I would also like to thank the USDA Forest Service and Northeastern Forest
Experiment Station for making this research possible. Elaine Kermedy Sutherland, David
Hosack, and especially Todd Hutchinson were instrumental in helping this study reach its completion. Todd is a good friend and colleague and I wish him luck in the future.
Foster Purrington and his expertise as a carabidologist allowed this 5-year study to be completed in approximately five years. I will miss working with him, the other
residents of the greenhouses, and the many field and lab workers who have assisted over the years.
Most importantly, I would like to thank my families. Both the Stantons and
Taylors have helped in facilitating this work and I appreciate their assistance. But it has
been Trish, Alex, and Ben who have sacrificed the most in order to support me. I’m sure
I do not realize all that has been given up to allow me to reach this point, but I hope they
know that this work was undertaken and accomplished with each of them in mind. VTTA
October 9, 1968 ...... Bom - Cincinnati, Ohio
1991...... B.A. English, Wittenberg University
199 4...... M.S. Entomology and Plant Pathology, The University of Tennessee
1995 - present...... Graduate Teaching, Research, and Administrative Associate, The Ohio State University
PUBLICATIONS
Research Publications
1. F.F. Purrington, R.C. Stanton, and D.J. Horn. 1999. Ground beetle range extensions: Six new Ohio records (Coleoptera: Carabidae). Great Lakes Entomol. 32:47-49.
2. F.F. PiuTington and R.C. Stanton. 1996. New records of five ground beetles from Ohio (Coleoptera; Carabidae). Great Lakes Entomol. 29:43-44.
3. R.C. Stanton, J.F. Grant, P.L.Lambdin, L.R. Barber, and S.E. Schlarbaum. 1993. Preliminary investigations of arthropod species diversity in a northern red oak seedling seed orchard. Proc. 38* So. Nurserymen's Assoc. Res. Conf. 38: 172-174.
FIELDS OF STUDY
Major Field: Entomology
VI TABLE OF CONTENTS
Page Abstract...... ü
Dedication ...... iv
Acknowledgments ...... v
Vita...... vi
List of Tables ...... ix
List of Figures ...... xii
Chapters:
1. General Introduction ...... l
Oak Regeneration and Fire ...... I The Ecosystem Management Project ...... 5 Invertebrates as Indicators ...... 16 Ground Beetles and Monitoring Methods ...... 19
2. Effects of Prescribed Burning on Carabid Abundance and Seasonality in Mixed-Oak Forests of Southern Ohio ...... 34
Introduction ...... 34 Materials and Methods ...... 37 Results...... 40 Discussion ...... 45
3. Effects of Prescribed Burning on Carabid Species Richness and Diversity in Mixed-Oak Forests of Southern Ohio ...... 61
Introduction ...... 61 Materials and Methods ...... 63 Results...... 64 Discussion ...... 73
vu 4. Carabid Assemblages in Mixed-Oak Forests of Southern Ohio...... 84
Introduction ...... 84 Materials and Methods ...... 86 Results...... 87 Discussion ...... 97
5. General Discussion ...... 106
Appendix ...... 109
List of References ...... 150
VIU LIST OF TABLES
Table Page
1 Locations and ownership of the four study areas chosen for the ecosystem management project ...... 9
2 Years in which prescribed fire treatments were administered to treatment units ...... 15
3 Dates of each prescribed fire for each study site ...... 15
4 Criteria for selecting indicator taxa (from Rodriguex et al. 1998)...... 18
5 Dates when pitfall traps were operated and corresponding collecting efforts ...... 29
6 Fire treatments applied to each treatment unit ...... 38
7 Overall carabid abundance collected by pitfall trapping on each treatment unit ( 1996 data are adjusted to reflect a more even sampling effort) ...... 41
8 Analysis of variance for overall carabid abundance (adjusted and log transformed) ...... 43
9 Analysis of variance for spring carabid abundance (adjusted) ...... 49
10 Analysis of variance for summer carabid abundance (adjusted and log transformed) ...... 49
11 Analysis of variance for autumn carabid abundance (adjusted and log transformed) ...... 49
12 Abundance of Synuchus impunctatus collected by pitfall trapping (1996 data are adjiti .::d to reflect a more even sampling effort) ...... 52 ix Table
13 Analysis of variance for overall carabid abundance without Synuchus impunctatus (adjusted and log transformed) ...... 54
14 Analysis of variance for overall flightless carabid abundance (adjusted and log transformed) ...... 59
15 Carabid species richness collected by pitfall trapping on each treatment unit (1996 data are adjusted to reflect a more even sampling effort) ...... 65
16 Analysis of variance for pooled carabid species richness (adjusted and log transformed) ...... 68
17 Carabid diversity for pitfall trapping data using the Shannon Index 1996 data are adjusted) ...... 70
18 Analysis of variance for pooled Shannon Index scores (adjusted) 72
19 Carabid evenness for pitfall trapping data using Shaimon Evenness (1996 data are adjusted) ...... 74
20 Analysis of variance for pooled Shatmon Evenness scores (based on adjusted data) ...... 76
21 Estimates of “true” species richness using the Chao ( 1984) estimator ( 1996 data are adjusted) ...... 77
22 Analysis of variance for pooled Chao estimated species richness 79
23 Carabid species collected every year by pitfall trapping ( 1995-1999) ...... 88
24 Carabid species captured at all four study sites by pitfall trapping 90
25 Carabid species captured at all four study sites by light trapping 91
26 Complementarity values for each site assemblages based on pitfall data... 92
27 Complementarity values for each site assemblages based on light trapping data ...... 94
28 Carabid species unique to each study site ...... 99 Table Page
29 Species dominance on the Arch Rock units ...... LOI
30 Species dominance on the Bluegrass Ridge units ...... 102
31 Species dominance on the Watch Rock units ...... 103
32 Species dominance on the Young’s Branch units ...... 104
Ai Precipitation totals recorded at the Vinton Experimental Forest (Hosack, unpublished)...... 110
A2 List of carabid species collected by pitfall trapping ...... 112
A3 List of carabid species collected by light trapping ...... 114
A4 Regional carabid assemblage collected by pitfall trapping ...... 117
A5 Carabid assemblage captured by pitfall trapping at Arch Rock ( 1996 data are adjusted) ...... 119
A6 Carabid assemblage captured by pitfall trapping at Bluegrass Ridge ( 1996 data are adjusted) ...... 121
AT Carabid assemblage captured by pitfall trapping at Watch Rock ( 1996 data are adjusted) ...... 123
A8 Carabid assemblage captured by pitfall trapping at Young’s Branch ( 1996 data are adjusted) ...... 125
A9 Carabid assemblages captured by light trapping ...... 127
XI LIST OFHGURES
Figure
1 Map of Ohio showing the locations of Arch Rock (AR), Bluegrass Ridge (BR), Watch Rock (WR), and Young’s Branch (YB) ...... 8
2 Topographic map of the Arch Rock treatment units ...... II
3 Topographic map of the Bluegrass Ridge treatment units ...... 12
4 Topographic map of the Watch Rock treatment units ...... 13
5 Topographic map of the Young’s Branch treatment units ...... 14
6 Average carabid abundance by prescribed burning treatment and year.. 42
7 Average spring abundance of carabids by treatment and year ...... 46
8 Average summer abundance of carabids by treatment and year 47
9 Average aummn abundance of carabids by treatment and year 48
10 Precipitation levels recorded at the Vinton Experimental Forest (Hosack, unpublished) ...... 55
11 Average abundance of flightless carabids by treatment and year 58
12 Average carabid species richness by treatment and year ...... 67
13 Average Shannon Diversity for carabids by treatment and year 71
14 Average carabid evenness by treatment and year ...... 75
15 Average estimated species richness (Chao estimator) by treatment and year ...... 78
16 Principle Components Analysis of pitfall carabid data ...... 95
xii Figure Page
17 Carabid assemblages from pitfall trapping at each study site ...... 96
A1 Pitfall collection data from Arch Rock, 1995 ...... 130
A2 Pitfall collection data from Bluegrass Ridge, 1995 ...... 131
A3 Pitfall collection data from Watch Rock. 1995 ...... 132
A4 Pitfall collection data from Young’s Branch, 1995 ...... 133
A5 Pitfall collection data from Arch Rock, 1996 ...... 134
A6 Pitfall collection data from Bluegrass Ridge, 1996 ...... 135
AT Pitfall collection data from Watch Rock, 1996 ...... 136
A8 Pitfall collection data from Young’s Branch, 1996 ...... 137
A9 Pitfall collection data from Arch Rock, 1997 ...... 138
AlO Pitfall collection data from Bluegrass Ridge, 1997 ...... 139
A11 Pitfall collection data from Watch Rock, 1997 ...... 140
A12 Pitfall collection data from Young’s Branch. 1997 ...... 141
A13 Pitfall collection data from Arch Rock, 1998 ...... 142
A14 Pitfall collection data from Bluegrass Ridge, 1998 ...... 143
A15 Pitfall collection data from Watch Rock, 1998 ...... 144
A16 Pitfall collection data from Young’s Branch, 1998 ...... 145
A17 Pitfall collection data fmm Arch Rock, 1999 ...... 146
A18 Pitfall collection data from Bluegrass Ridge, 1999 ...... 147
A19 Pitfall collection data from Watch Rock, 1999 ...... 148
A20 Pitfall collection data from Young’s Branch, 1999 ...... 149
xm CHAPTER I
GENERAL INTRODUCTION
The oak component of forests found in the upland regions of the eastern United
States has been in decline for the past 50 years. Most of these forests lack sufficient amounts of established oak seedlings, known as advanced oak regeneration, to maintain oak dominance in the overstory. Prescribed burning of the understory is one method that may help improve oak regeneration, but ± e consequences of this practice on other aspects of the oak forest community are largely unknown. In this study, ground beetle populations were monitored in mixed-oak forests subjected to prescribed burning, and compared to unbumed controls, to help determine the effects of burning on forest floor invertebrates and evaluate prescribed burning as a silvicultural tool.
Oak Regeneration and Fire
The hardwood forests that cover much of eastern North America are largely composed of oak {Querciis) species (Keyser et al. 1996). Oak-hickory (jQuercus-Carya) forest is the most common forest type in the eastern United States, occupying about 46 million hectares and dominating the dry upland regions (Braun 1972, Powell et ai. 1993, Sander et ai. 1983). Pollen records indicate that this dominance has existed in the eastern states for nearly 10,000 years (Delcourt and Delcourt 1987, Webb
1981). In southern Ohio, the status of oak species as the major component of these deciduous forests is known to predate European colonization (Crow 1988, Gordon 1969).
It is widely held that this oak dominance developed, and has been perpetuated by. frequent fires (Abrams 1992, Abrams and Nowacki 1992, Brown 1960, Lorimer 1984,
Olson 1996). Many of these fires are believed to have been intentionally set by Native
Americans (Cooper 1961, Day 1953, Genevan 1992, Little 1974) and early European settlers (Abrams 1992, Pyne 1982. Van Lear and Waldrop 1989) in order to attract wildlife, encourage specific vegetation, and improve travel and visibility (Little 1974,
Van Lear and Waldrop 1991). To whatever extent the fires were natural or anthropogenic, intentional or accidental, they are closely associated with the composition of present-day oak forests (Abrams 1992, Johnson 1993, Sutherland 1997).
However, over the past 40 to 50 years, researchers have observed a decline in the oak component of the overstory of these forests due to a dramatic decrease in advanced oak regeneration (McGee et al. 1995. Lorimer 1992). As a result of this decline, more shade-tolerant, mixed mesophytic species have increased (Abrams 1992, Abrams 1998,
Crow 1988, Lorimer 1984), such as American beech {Fagus grandifolia), red maple
(Acer rubrum), sugar maple (Acer saccharum), and yellow-poplar (Liriodendron
tuUpifera).
Several explanations have been suggested for this change in forest regeneration,
including increased atmospheric deposition (Hutchinson et al. 1999, Solomon et al.
1987), changes in timber harvesting (Hutchinson et al. 1999, Johnson 1993, Little 1974), and the widespread Ore suppression policies instituted in the 1930s and 1940s (Abrams
1992, Crow 1988, Pyne 1984. Robertson and Heikens 1994). Other possibilities include gypsy moth (Lymantria dispar) defoliation in the northeastern states and oak diseases
(Ammon et al. 1989, Johnson 1993). In some forests, this shift from oak-hickory to beech-maple may be due to natural succession.
The concern over the loss of oaks is based on the fact that the more mesic tree species lack the economic and ecological value of oaks (Brose and Van Lear 1998,
Keyser et ai. 1996). Beech sawtimber does not have the economic value of oak, and maples and yellow-poplar do not provide as many benefits to wildlife as oaks (Keyser et al. 1996, Loftis 1990a).
Because oak decline has been traced to the reduction of oak regeneration, several solutions for improving oak regeneration have been recommended. These suggestions include the shelterwood harvesting system (Hannah 1987, Loftis 1990b), a prescribed burning regime (Abrams 1992. Lorimer 1992, Van Lear 1990, Van Lear and Watt 1992), and a combination of shelterwood harvesting and a form of disturbance, such as herbicide application (Loftis 1990b, Lorimer et al. 1994) and prescribed fire (Hannah 1987).
However, it has been prescribed biuming alone to improve oak regeneration that has received the most attention from researchers.
Prescribed burning has been defined as “fire set under plaimed conditions to accomplish specific management objectives” (Van Lear and Waldrop 1991). It has been widely accepted and used to regenerate other tree species, such as western conifers
(Patterson 1992) and southern pines (Duryea and Dougherty 1991). Aside fix>m maintaining certain species and eliminating undesirable ones, prescribed fires have also
3 been used as a management tool to reduce the risk of wildfire and enhance species diversity (Whelan 1995). Although this method has been widely applied, public perception of the technique has been negative (Taylor and Daniel 1984), but Manfredo et al. (1990) demonstrated that as knowledge about the technique increases, public support of the fires also increases.
The idea that prescribed burning would facilitate oak regeneration is based on the tenet that oaks are fire-resistant species (Lorimer 1985) that developed under a frequent fire regime (Patterson 1992). Oak seedlings accentuate their root development more than their shoot growth, while the mixed mesophytic species emphasize shoot growth (Brose and Van Lear 1998, Johnson 1993. Kolb et al. 1990). Therefore, oaks are more capable of resprouting firom the root-cellar after top-kill by fire whereas seedlings of other species are often killed by fire (Loftis 1990b, Van Lear and Waldrop 1988). The larger root systems of oaks also allow for rapid growth in height upon release (Sander 1971, Sander
1972). In addition, oaks have thick bark and can quickly compartmentalize cambium that has been damaged by fire (Abrams 1992). Maple species have a thinner bark, which also contributes to their susceptibility to fire (Abrams i992). So, as summarized by Johnson
( 1993), prescribed fires can promote oak regeneration by reducing the number of fire- sensitive understory competitors, by reducing overstory density by killing trees with thin, fire sensitive bark, and by killing oak stems which increases the rootrshoot ratio of those trees that survive by resprouting.
hiitial smdies, based on one-time prescribed bums, suggested that prescribed fire did not encourage oak regeneration (Johnson 1974, Teuke and Van Lear 1982, Wendell and Smith 1986). More recently, multiple applications of prescribed fire have been shown to improve oak regeneration^ as a follow-up to shelterwood harvesting (Brose and
Van Lear 1998, Brose et al. 1999, Keyser et al. 1996) and on their own (Barnes and Van
Lear 1998, Merritt and Pope 1991). Brose et al. ( 1999) and Teuke and Van Lear ( 1982)
report that prescribed burning also improves the form and growth rate of oak sprouts.
Today, prescribed burning is viewed as a promising tool for increasing the oak
component of the advanced regeneration pool (Brose and Van Lear 1998). However, no
studies prior to this one have been conducted to determine the other effects this practice
may have on the various components within the oak forest communities of the eastern
United States.
The Ecosvstem Management Project
The interest in utilizing prescribed burning to improve oak regeneration and the
remaining questions regarding its use (Barnes and Van Lear 1998) led to a long-term
research project initiated by the USDA Forest Service, Northeastern Forest Experiment
Station, in 1994. The research objective of this project was to determine the ecological
response of oak-dominated communities in southern Ohio to frequent and infrequent
prescribed burning regimes (Sutherland, pers. comm.). Because this study sought to
determine the response of the entire community to fire, and not just oak regeneration, it
was known as an ecosystem management (EM) project.
The two main goals of the EM project were to 1.) determine appropriate
prescribed underbuming regimes as management tools in restoring the structure and
function and much of the composition (fire-adapted flora) to the mixed-oak forests of
southern Ohio, and 2.) design and implement a monitoring program of fire effects and
5 ecosystem sustainability (Sutherland, pers. comm.). The first goal refers to the lack of knowledge regarding the appropriate fire frequencies needed to restore and maintain oak forests. Fire frequency in this region of the United States is not well known, mostly due to a lack of old-growth trees to study (Sutherland 1997). Guyette and Cutter ( 1991) reported an average fire interval of 4.2 years between 1700 and 1810 in a Missouri oak savanna. In southern Illinois, fire frequency since the 1870s was highest between 1915 and 1935 (Robertson and Heinkens 1994). In a smdy of an area near the EM project,
Sutherland (1997) reported average fire intervals of 3.6 years for small fires and 7.5 years for large ones since the 1850s.
The second goal of the EM project relates to monitoring fire effects on mixed-oak communities. Several parameters of these forests were selected to be monitored,
including soil dynamics and belowground productivity, understory vegetation,
regeneration, overstory vegetation, vertebrate fauna and invertebrate fauna. Few studies
have focused on the effects of fire on any one of these aspects of deciduous forests. This
is the first known study to consider the effects of fire on each of these components of the
community.
The study sites for this project were located in Vinton and Lawrence counties on
the unglaciated Allegheny Plateau of southern Ohio. The climate of the region is humid-
continental, having a mean armual temperature of 11.3° C, mean annual precipitation of
1024 mm, and an average of 158 frost-fise days (Hutchinson et al. 1999). The
topography consists of numerous ridges and steep drainages, with elevations from 200 to
315 m (Horn, pers. comm.). At the time European settlement of southern Ohio began (around 18(X)), 95% of this area was covered by oak forests (Sutherland 1997). Deforestation of the immediate area occurred as the pig iron industry thrived between the 1850s and 1900, peaking during the 1880s (Sutherland 1997, Willard 1916). During this time, nearly all forests were clearcut for use as charcoal in the iron smelters or furnaces (Stout 1933). By the turn of the century, most of the iron industry had moved to Wisconsin and Miimesota and the forests have since regenerated to oak dominance, with limited logging (Griffith 1991,
Hutchinson et al. 1999, Stout 1933).
In 1994, four smdy sites were chosen (Arch Rock, Bluegrass Ridge, Watch Rock, and Young’s Branch) in this area (Figure 1) based on forest composition and age, soil type, and land ownership (Table 1). All four sites ranged in area from 75 to 110 ha within larger blocks of continuous forest and were composed of relatively undisturbed,
second-growth (80 to 120 years old) mixed-oak forests (Hutchinson et al. 1999). The overstory of these forests was comprised of > 70 % oak species (Quercus alba, Q.
velutina, Q. rubra, and Q. prinus). The understory, typical of today’s eastern oak forests,
is dominated by mixed mesophytic species (e.g. Acer saccharum, A. rubrum, and Nyssa sylvatica) (Hutchinson et al. 1999).
Underlying the smdy sites were sandstones and siltstones of Mississipian and
Pennsylvanian origin, although some areas of interbedded limestone were found within
the Bluegrass Ridge site. At Arch Rock, Watch Rock, and Young’s Branch, the major
soil association was Steinsburg-Gilpin Association. At Bluegrass Ridge, soils were of the
Upshur-Gilpin-Steinsburg Association. All of these soils were mostly silt loam alflsols
with moderate acidity and low water-holding capacity (Hutchinson et al. 1999).
7 CM
lYB
BR
Figure 1. Map of Ohio showing the locations of Arch Rock (AR), Bluegrass Ridge (BR), Watch Rock (WR) and Young’s Branch (YB). Study Site Countv Latitude/Longitude Ownership
Arch Rock Vinton SQnZ'N. 82°23'W Mead Paper Corp.
Bluegrass Ridge Lawrence 38°36'N, 82°31'W Wayne National Forest (Ironton District)
Watch Rock Vinton 39° I m , 82°22'W Mead Paper Corp.
Young’s Branch Lawrence 38°43'N, 82°4IW Wayne National Forest (Ironton District)
Table 1. Locations and ownership of the four study areas chosen for the ecosystem management project (from Hutchinson et al. 1999). Each of the four study sites was subdivided into three treatment units, approximately 30 ha each, as a randomized complete block (Figures 2,3,4, and 5). The treatments were “frequent” fire (burned annually), “infrequent” fire (burned every third year), and “control” (not burned).
Preliminary pre-treatment data were collected for the EM project in 1994 and
1995. In 1996, all eight frequent and infrequent treatment units were burned. The four frequent units were rebumed in 1997,1998, and 1999. The four infrequent units were rebumed in 1999 (Table 2).
Each prescribed bum was a spring fire, set during March or April when fire conditions were appropriate (Table 3). Spring (March and April) and aummn (October and November) are the natural fire seasons in Ohio (Sutherland 1997) as well as most of
the eastern United States (Brose and Van Lear 1998, Robertson and Heikens 1994). One
springtime prescribed fire has been reported to be as effective as three winter bums in encouraging oak regeneration (Barnes and Van Lear 1998) and often result in moderately
intense fires in oak forests (Brose and Van Lear 1998. Brose et al. 1999). Keyser et al.
( 1996) initiated a summer (August) fire in Virginia that was of moderate intensity. In
general, spring is considered to be the preferred season for prescribed burning in oak-
dominated stands (Brose and Van Lear 1998, Brose et al. 1999).
Drip torches containing a mixture of diesel fuel and kerosene were used to set
strip-head fires, at various intervals (Hutchinson, pers. comm.). In most cases, flames
burned uphill and /or were aided by wind. Forest patches that did not bum were not
reignited.
10 ARCH ROCK Frequent
Control
Infrequent
# Trapping Locations I I Rrelines /\/ 20-foot contours
400 800 Mete IS
Figure 2. Topographic map of the Arch Rock treatment units.
II BLUEGRASS RIDGE
Frequent
Infrequent
Control
# Trapping Locations IN [~~| Rrelines /'\/ 20-foot contours A
400 800 Meters
Figure 3. Topographic map o f the Bluegrass Ridge treatment units.
12 WATCH ROCK
Infrequent
Frequent Control
IN 0 Trapping Locations Rrelines / V 20-foot contours A
400 800 Meters
Rgure 4. Topographic map of the Watch Rock treatment units.
13 YOUNG'S BRANCH
Control
Frequent Infrequent
4^ Trappfng Locations Rrelines / V 20-foot contours
400 800 Meters
Figure 5. Topographic map o f the Young’s Branch treatment units.
14 Treatment Unit 1994 1995 1996 1997 1998 1999
Frequent no no yes yes yes yes
Infrequent no no yes no no yes
Control no no no no no no
Table 2. Years in which prescribed fire treatments were administered to treatment units.
Study Site 1996 1997 1998 1999
Arch Rock Apr IB Apr 2 Apr 6 Mar 26
Bluegrass Ridge A p ril. 12 Mar 21 Mar 30 Mar 29
Watch Rock Apr 19,21 Apr 3 Apr 6 Mar 27
Young’s Branch Apr 18,19 Mar 27 Mar 29 Mar 30
Table 3. Dates of each prescribed fire for each study site.
15 The effects of these fires on the various components of the mixed-oak community have been studied by a large team of researchers from agencies and instimtions such as the USD A Forest Service, The Nature Conservancy, Ohio Division of Natural Areas and
Preserves, The Ohio State University, and Ohio University. The research presented here is a portion of the efforts made to determine the effects of these fires at these sites.
Invertebrates as Indicators
Invertebrate species constitute a significant portion of this mixed-oak community.
To determine the impacts of prescribed burning on ail invertebrates would be nearly impossible considering the immense diversity of taxa and the time, labor, funding, and taxonomic expertise necessary for such an undertaking. Therefore, research efforts needed to be concentrated on a limited number of taxa most likely to provide the most information relating to the effects of the fire treatments.
Of the many invertebrate groups on which to focus, those that occupy or are active on the forest floor are most likely to be impacted by periodic surface fires.
Commonly collected invertebrates from the forest floor of these study areas include millipedes (Diplopoda), spiders ( Araneae), and insects (Insecta). Of the insects, several
families of beetles (Coleoptera), moths (Lepidoptera), flies (Diptera), and wasps
(Hymenoptera), as well as crickets (Gryllacrididae) and ants (Formicidae), are frequently encountered. Selecting one or several groups of invertebrates to accimitely represent all
invertebrates is a difficult, if not impossible, task.
The concept of selecting a representative or characteristic group is a conventional procedure in ecological smdies but has received increased attention over the past 15 years
16 (Dufirene and Legendre 1997). Various terms have been used for these groups, such as focal groups, predictor sets, target taxa, priority taxa, surrogate taxa, and indicator taxa
(Oliver and Beattie 1996), but “indicator group/taxa” is now the most commonly used.
Criteria for selecting appropriate indicator taxa have been proposed by several authors (Noss 1990, Pearson 1994. Pearson and Cassola 1992). These criteria (Table 4) were summarized by Rodriguez et al. ( 1998). Other methods of identifying indicator groups, such as ordinations and indices, have also been suggested (Dufrene and Legendre
1997).
Few terrestrial invertebrate groups have been identified and utilized as suitable
indicators for monitoring and inventory studies (Foremen et al. 1993, Oliver and Beattie
1996). Currently, the most commonly used indicator groups are butterflies (Brown 1991,
Kremen 1992), ants (Andersen 1990, Majer 1983, York 1994), spiders (Hopkins and
Webb 1984), and ground, tiger, and scarab beetles (Hutcheson 1990, Klein 1989, Pearson
and Cassola 1992, Refseth 1980). Of these groups, ground beetles (Family Carabidae)
are the most utilized indicator taxa in northern and temperate regions.
Of the criteria listed in Table 4, ground beetles easily meet the first five criteria.
Not much data exist on carabids in terms of the sixth criterion (Pearson and Cassola
1992) and the economic importance of these beetles, thus far, has been based on their role
as biological control agents. In general, most researchers agree that carabids are suitable
and informative indicators for many monitoring and inventory studies (Butterfield et al.
1995, Maelfait and Desender 1990, Rykken et al. 1997, Thiele 1977).
For these reasons, as well as the fact that a carabid systematist was readily
available for this study, ground beetles were chosen as an indicator group for this study.
17 1. Well-known and stable taxonomy, so that species and populations can be readily determined.
2. Well-understood biology and natural history
3. Easily surveyed populations
4. Broad geographic distribution and wide range of habitat types (at higher taxonomic levels, such as family and genus)
5. Specialization within narrow habitats (at lower taxonomic levels, such as species and subspecies)
6. Patterns reflect those observed in other taxonomically related and unrelated taxa
7. Potential economic importance
Table 4. Criteria for selecting indicator taxa (from Rodriguez et al. 1998).
18 Ground beetle populations were monitored for any changes that occurred after prescribed fire treatments with the idea that these changes may indicate the impacts of prescribed burning on other forest floor invertebrates.
Ground Beetles and Monitoring Methods
Ground beetles form a very large, popular group of insects that has received enormous attention from both professional scientists and amateur collectors. Nearly every aspect of their biology, and how they relate to larger fields such as ecology and evolution, has been studied to some degree, especially in Europe (Thiele 1977). Ground beetles first appeared in our current body of scientific literature in 1602 (Lindroth 1979), well before Linnaeus introduced binomial nomenclature in 1758. Today, they are one of the most-studied arthropod groups (Desender et al. 1994). A complete synthesis of all of the scientific work done on these insects is not possible; however several important features of these organisms relevant to this research will be introduced.
Taxonomically, ground beetles are placed in the coleopteran family Carabidae.
Although there is still some debate (Crowson 1981, Knisley and Schultz 1997), most carabidologists also include tiger beetles as a subfamily (Cicindelinae) within the
Carabidae (Desender et al. 1994, Erwin 1979). This family contains more than 40,000 described species, including about 2,000 described species of tiger beetles (Knisley and
Schultz 1997), classified into 86 tribes (Lovei and Sunderland 1996). The Carabidae is considered to be the third largest beetle family, behind the Staphylinidae and
Curculionidae, in North America (Borror et al. 1989). Despite this immense species
19 richness, Carabidae is one of the best-studied families of arthropods (Desender et ai.
1994).
It is generally accepted that carabids emerged during the early Tertiary period from wet, tropical habitats (Erwin 1979a). They later radiated to drier envirorunents and more extreme latitudes and altitudes (Lovei and Sunderland 1996). Currently, ground beedes and tiger beetles are found worldwide, occurring on every continent except
Antarctica (Noonan et al. 1992) and are common in nearly every habitat with the exception of deserts (Lovei and Sunderland 1996). Some species have evolved to fill specialized habitats such as caves, edges of glaciers, forest canopies, and self-constructed turmels in tropical sands and soils (Erwin 1985, Lovei and Sunderland 1996). Carabid species richness, however, reaches its highest levels in tropical regions (Erwin 1985).
Despite the species radiation exhibited by ground beetles, carabid morphology is fairly conserved and adheres to an easily recognized, generalized body plan (Lovei and
Sunderland 1996, Thiele 1977). The family is placed in the suborder Adephaga on the basis of the presence of six abdominal ventrites, hind coxae that divide the first visible abdominal sternum, pygidial defense glands in the adult, and liquid-feeding mouthparts in the larvae (Borror et al. 1989, Lawrence and Britton 1991). The adults of nearly every species have relatively long, thin cursorial legs with the primitive 5-5-5 tarsal formula, notopleural sutures, and filiform antennae (Borror et al. 1989, Evans 1986). Larvae are campodeiform, with well-developed legs, antennae, and mandibles (Lovei and
Sunderland 1996).
Most carabid species are voracious predators (Thiele 1977) that himt or scavenge on the soil surface or in leaf litter and hide under leaves, stones, logs, and tree bark
20 (Evans 1986). The majority of adults kill and disjoin their prey using their well- developed mandibles (Lovei and Sunderland 1996), although preoral digestion is not uncommon (Loreau 1986). Other carabid species are polyphagous (feeding on both live prey and carrion), phytophagous, or omnivorous (Lovei and Sunderland 1996, Thiele
1977).
LarocheUe (1990) surveyed the literature for dietary reports and found that approximately 73% of the 1054 species covered were carnivorous, 19% omnivorous, and
8% phytophagous. Gut-dissection studies, conducted in Europe and New Zealand, have revealed a wide array of food choices. Among the insect prey reported in these studies are aphids, lepidopterans (adults and larvae), fly larvae, heteropterans, beetle larvae, springtails, hymenopterans, and insect eggs. Other invertebrate prey items include spiders, opilionids, enchytraid worms, lumbricid worms, nematodes, centipedes, miUipedes, and moUusks (Hengeveld 1980, PoUet and Desender 1987, Sunderland et al.
1995). A study of soil-dwelling carabids in forests outside Moscow adds psocopterans and isopods to this list (Gryuntal and Sergeyeva 1994). Some riparian carabids reportedly feed on emerging stoneflies and other aquatic organisms (Hering and Plachter
1997). Plant material found within the digestive tracts of carabids includes spores, fungal hyphae, seeds, and pollen (Hengeveld 1980, PoUet and Desender 1987, Sunderland et al.
1995).
From laboratory studies, Currie et al. (1996) reported that two species of
Pterostichus exhibit cannibalism and congeneric predation. Ovaska and Smith (1988) demonstrated that two forest species on Vancouver Island consumed juvenile salamanders.
21 Many carabid larvae are also active foragers, feeding on live prey, carrion, and plant material. Some species exhibit very specialized life histories, such as ectoparasitoids on other invertebrates and symbiotic relationships with ants and termites
(Erwin 1979b).
Of the carabid species that have been studied, very few are considered to be specialists. Most species are opportunistic feeders, preying on whatever food sources are readily available (Allen 1979). One notable exception is Calosoma sycophanta, a large carabid that feeds primarily on forest caterpillars, especially the gypsy moth, Lymantria dispar (Lovti and Sunderland 1996, Thiele 1977). However, nearly all carabid feeding studies have been conducted in the northern hemisphere on local species, so results are biased to these species (Lovei and Sunderland 1996).
Another feature of some carabids that deserves mention is flightlessness. Many species have lost the ability to fly, a feature that has evolved on multiple occasions within the Carabidae (Darlington 1943, Roff 1994). Many other species exhibit wing dimorphism in which some individuals within a species possess functional wings and others individuals do not. The expression of dimorphism may be influenced by environmental conditions (Aukema 1991) or by residence time within a stable habitat
(den Boer et al. 1980, Niemela and Spence 1991). Of the species that are capable of flight, many prefer to run and/or hide rather than fly (Thiele 1977).
This inability, or reluctance, to fly is important to the fact that many carabid species are found within specific habitats and do not occur outside of narrowly-defined environmental conditions (Noonan et al. 1992). For example, in a smdy of the carabid fauna on New England mountains. Bell (1992) identified four distinct
22 climatological/vegetation zones. The carabids collected in this region also showed well- marked zonation, dividing up into distinct assemblages according to climate and vegetation. Many similar examples of habitat fidelity exhibited by ground beetles exist in the literature (Luff 1990, Luff et ai. 1992, Pizzolotto and Brandmayr 1990, Rushton et al.
1990).
Because many carabids demonstrate this specific and consistent preference in habitat, the presence or absence of certain carabid species can be used to characterize a habitat (Lovei and Sunderland 1996) and often indicates environmental trends within that habitat (Thiele 1977). This concept of carabids serving as “environmental indicators”
(Thiele 1977) gained support during the 1970s from the many studies showing that carabid abundance is significantly reduced by insecticide applications in agricultural fields (Basedow 1990, Croft and Brown 1975, Kulman 1974, Vickerman and Sunderland
1977). These observations led to the use of carabids as bioindicators of other forms of environmental pollution and/or disturbance (Freitag 1979). More recently, carabid censuses have been used to assess the extent of environmental disturbances
(Mossakowski et al. 1990, Rodriguez et al. 1998), ecological condition or quality (Casale
1990, Van Essen 1994), and the geographical/climatological history of a vast array of habitats (Ervynck et ai. 1994, Kavanaugh 1979). Other studies have employed carabids to indicate and monitor areas in need of management (Butterfield et al. 1995. Desender and Maelfait 1991, Eyre and Luff 1990, Pizzolotto 1994) and conservation (Desender et al. 1991, Eyre and Rushton 1989, Heijerman andTurm 1994, Horvatovich 1994).
Kremen et al. (1993) and Pearson (1994) divided these and similar studies into two
23 general categories—"monitoring studies" that evaluate changes over time and “inventory smdies” that record distributional patterns (Rodriguez et al. 1998).
Despite the heavy use of carabids as indicators of various habitat conditions, there is still limited debate as to their value as bioindicators because little has been done to critically assess the adequacy of indicator groups in general (Lovei and Sunderland
1996). For example, one taxonomic group may not adequately represent the trends of other groups (Lawton et al. 1998).
Another widely debated issue, that most carabidologists agree on nonetheless, relates to the methods of sampling for ground beetles. Four basic collection methods have been used to smdy carabids—hand collecting, soil or litter removal, blacklight (or ultraviolet) traps, and pitfall traps. The latter method has received widespread criticism since its introduction: however it continues to be the most widely used sampling technique (Digweed et al. 1995).
Historically, carabids had been exclusively collected by hand (Thiele 1977) and no other methods were used until Hertz ( 1927) and Barber (1931) first used tin cans to collect insects. These cans were the precursors to today’s pitfall traps, originally known as Barber traps. However, widespread use of this collecting method did not occur until the mid-1950s (Thiele 1977).
The other collecting methods have received limited use. Sod and litter removal involves collecting a specific amount of soil and/or leaf litter and either soaking it in water to isolate the beetles (Dunn 1983, Gryuntal 1982, Spence and Niemela 1994) or drying out the sample in Tullgren or Berlese funnels to extract the beetles (Borror et al.
1989). Blacklight traps capture carabid species that are attracted to light but this method
24 has only recently been employed in ecological studies (Kadar and Szel 1995, Kadar and
Szentkiralyi 1997, Matalin 1996, Purrington 1996, Yahiro et al. 1997).
The original pitfall traps consisted simply of tin cans buried in the ground.
Carabids moving along the ground would fall into the cans and would be unable to escape. These traps would now be referred to as “live traps,” because there was no killing agent inside the cans. Carabids, and other arthropods, captured in these traps remained alive assuming no within-trap predation occurred (Mitchell 1963).
Evenmally, killing and preserving agents were added to the traps to prevent escape and decomposition of specimens. Originally, formalin, picric acid, and water were used. Formalin and picric acid are no longer in wide use due to associated health hazards (Weeks and McIntyre 1997). Water is not a very effective killing or preserving agent and has been shown to repel some carabid species (Weeks and McIntyre 1997).
Concentrated brine is also a poor preservative (Lemieux 1998, van den Berghe 1992).
Ethylene glycol (commonly used as automotive antifreeze) is now the most frequently used killing and preserving solution. However, it is extremely toxic to mammals (Hall 1991) and one study claims that it may attract certain carabids. resulting in biased samples relating to speciesrspecies and malerfemale ratios (Holopainen 1990).
Recently, propylene glycol, which is non-toxic, has been introduced to the market. Initial studies indicate that propylene glycol is just as effective as ethylene glycol in the number of beetles captured and the condition in which they are preserved (Weeks and McIntyre
1997).
Other modifications have also been made to the original pitfall trap (Southwood
1978). Barriers, or “drift fences,” are often used to guide beetles toward the traps and
25 increase the number of beetles collected (Durkis and Reeves 1982) and ramps can facilitate capture in some habitats (Bostanian et al. 1983). Baiting the traps has also been practicecb but has not been shown to increase the number of carabids collected (Novak
1969, Tardiff and Dindal 1980). General guidelines for using pitfall traps have been summarized by van den Berghe ( 1992), although modifications are continually being suggested (Lemieux and Lindgren 1999).
The reasons behind the popularity of pitfall traps are many. Fundamentally, they are inexpensive, do not require a large amount of labor, and result in high numbers of captured specimens that could not be achieved by hand collecting (Thiele 1977). Other advantages include continuous sampling over a 24-hour period or entire season, simultaneous sampling on multiple sites, and a collection that is independent of the skill of the collector (Bostanian et al. 1983).
However, there has been long-standing opposition to the use of pitfall-collected data. Initial arguments centered on the fact that pitfall data cannot serve as accurate estimates of population density because they simply record carabid activity (Thiele
1977), which creates problems with data interpretation (Briggs 1961, Mitchell 1963,
Southwood 1978, Turnbull 1973). Potential problems and sources of error that have been associated with pitfall trapping include varying levels of activity by different species
(Thiele 1977), different microclimate conditions near the traps (Honek 1988), the number, size, and placement of traps (Adis 1979), trap material (Luff 1975), surrounding vegetation (Greenslade 1964), carabid secretions and pheromones (Luff 1986), and killing agent (Holopainen 1990). Some researchers state that these factors influence the results so much that ecological inferences are difficult to make (Thiele 1977). Several
26 studies conducted to address these issues have concluded that pitfall data are unreliable and should not be used extensively (Halsall and Wratten 1988, Topping and Sunderland
1992),
Despite this opposition, many researchers support the use of pitfall traps
(Breymeyer 1966, Gist and Crossley 1973, Uetz and Unzicker 1976, Uetz 1977). Mark- and-recapture studies suggest that pitfall traps give a reliable estimate of yearly fluctuations in carabid densities (van der Drift 1951, den Boer 1986). Other studies report that data pooled &om an entire season of activity give an accurate picture of local abundance of individual species, especially in forested habitats (Baars 1979, den Boer
1986, Luff, 1982). However, Spence and Niemela (1994) do not recommend comparisons of pitfall data across habitat types. Currently, the majority of carabidologists agree that pitfall data are useful, but that conclusions should be made cautiously, incorporating the limitations of the data (Eyre and Luff 1990).
Despite the early debates and qualifications regarding the use of pitfall traps, the technique continues to be widely used and is generally accepted today (Digweed et al.
1995, Spence and Niemela 1994). One reason for this continued use may be that no reasonable alternatives have been suggested or made available (Spence and Niemela
1994).
Di this study, carabid populations were sampled via continuous pitfall trapping. A linear transect of 12 traps was established within each treatment unit. Traps were placed approximately 10 m apart—the exact location depending on natinal obstacles such as
fallen trees, rocks, and thick understory vegetation.
27 Traplines were positioned along dry ridge tops as near the center of each unit and in as similar a habitat as possible. Trapline locations were based on ready access as well as attempts to reduce edge effects. Therefore, trapline placement near the center of each treatment unit was not always possible (Figures 2,3,4, and 5). especially when dry ridge tops were situated near the edge of a treatment unit.
Each individual trap consisted of two plastic cups, with a diameter of 12 cm and a depth of 14 cm, one stacked inside the other. Holes were dug in the soil using a standard golf hole “cup cutter.” Cups were placed in the holes so that the rims of both cups were flush with the soil surface. The outer cup served to maintain the holes and facilitate the emptying of traps. This cup typically became embedded in the ground and was difficult to remove by the end of each season. Drainage holes were punched in the bottom of each outer cup so that rainwater and runoff could drain and the inner cup would not float up above the soil surface.
Inner cups were filled with approximately 150 ml of ethylene glycol. A 20 cm x
20 cm masonite rain shield was placed approximately 5 cm over each trap, supported by various sticks and rocks. A 45 cm x 45 cm piece of hex netting (“chicken wire”) was staked down over each trap assembly with four 20 cm-long aluminum gutter nails. The hex netting helped to keep the rain shields positioned over the traps and deterred trap disturbance by mammals.
Pitfall traplines were established at each study site as quickly as was logistically
possible after each fire (Table 5). In 1995,1996, and 1997, traps were emptied and reset at weekly intervals. In 1998 and 1999, traps were emptied approximately every two
weeks. Trap contents were poured through a fine mesh strainer and deposited into a
28 Year Studv Site Date of Fire Traps Set First Pick-up Traps Removed # Collectine Davs
1995 AR — 10-May 18-May 28-Sep 141
BR — 11-May 24-May lO-Oct 152
WR — 10-May 18-May 28-Sep 141
YB — 11-May 24-May lO-Oct 152 1996 AR 18-Apr 28-Apr 9-May 24-Oct 179 BR 11,12-Apr 20 Apr 8-May 23-Ocl 186 WR 19,21-Apr 28-Apr 9-May 24-Oci 179 YB 18,19-Apr 27-Apr 8-May 23-Oct 179 1997 AR 2-Apr 24-Apr 9-May 24-Sep 153 BR 21-Mar 24-Apr 9-May 24-Sep 153 WR 3-Apr 24-Apr 9-May 25-Sep 154 YB 27-Mar 24-Apr 9-May 24-Sep 153 1998 AR 6-Apr 25-Apr 7-May 24-Sep 152 BR 30-Mar 23-Apr 7-May 23-Sep 153 WR 6-Apr 25-Apr 7-May 24-Sep 152 YB 29-Mar 23-Apr 7-May 23-Sep 153
Table 5, Dates when pitfall traps were operated and corresponding collecting efforts. (continued) Table 5 (conlinued)
Year Studv Site Date of Fire Traps Set First Pick-UD Traps Removed # Collectine Davs
1999 AR 26-Mar \ 22-Apr 6-May 21-Sep 152 BR 29-Mar 21-Apr 6-May 21-Sep 153 WR 27-Mar 22-Apr 6-May 21-Sep 152 YB 30-Mar 21-Apr 6-May 21-Sep 153
W o plastic bag labeled with the date and location. Each plastic bag held 12 traps-worth of material.
Once the traps were emptied, plastic bags were placed in a cooler with an ice pack and transported to The Ohio State University in Columbus, OH. Samples were kept at ca.
-10° C in a standard chest freezer until they could be processed.
Traplines were maintained into autumn, depending on available labor (Table 5).
Ideally, traps would have been operated until the first hard frost, but this was not always possible. Traps were located in the same hole each year, except when fallen trees or eroded holes necessitated digging a new hole. Newly dug holes were located within 1 m of the original trap location.
Each pitfall sample was processed individually in the lab. Beetles, ants, and spiders were removed and placed into 60 ml plastic containers in 70 % ethanol to await identification. In some years, millipedes and snail shells were also removed. Organic debris, such as leaves and twigs, was disposed and the remainder of the sample stored in a 250 ml glass jar in 70 % ethanol.
Carabid species determinations were made by Foster F. Purrington (Depts. of
Entomology and Evolution, Ecology, and Organismal Biology at The Ohio State
University), following Bousquet and LarocheUe (1993). Purrington is a weU-known carabid systematist, having identified carabids for ecological studies in Iowa, Ohio,
Michigan, and Massachusetts. Voucher specimens of each species were soaked in water for 24 hours, pinned, and labeled for eventual deposit in the Museum of Biological
Diversity at The Ohio State University.
31 In addition to pitfall traps, a single 8-watt ultraviolet light trap was also operated within each treatment unit. The light traps were mainly used to assess treatment effects on other insects, such as moths, but did capture a large number of carabids. Light traps were operated approximately one night/week in 1995, 1996, and 1997 and one night/two weeks in 1998 and 1999.
Light traps were hung from low branches, approximately 1-2 m off the ground and were powered by a 12-volt portable, rechargeable battery. Batteries were wired to a timer that turned the light on at 9:00 pm and off at 5:00 am. Beetles, and other insects, approaching the light were intercepted by clear, plexiglass baffles that caused the insects
to fall through a funnel and into a 18.95-liter (5-gallon) plastic bucket. Ethyl acetate and
vapona (2,2-dichlorovinyl dimethyl phosphate) were used as killing agents inside the
bucket.
Samples were collected within 10 hours of the light’s turning off. Buckets were
emptied into plastic bags labeled with the date and location. Plastic bags were
transported in a cooler to the freezer in Columbus. As samples were sorted, beetles were
placed into 60 ml plastic cups in 70 % ethanol until identified.
These procedures were followed for each study area and treatment unit from 1995
to 1999. However, light trapping was discontinued in 1998 and 1999 at the Bluegrass
Ridge-A treatment unit (control) and pitfall trapping on this unit was discontinued in
1999.
These monitoring methods resulted in a five-year data set of carabid activity in
southern Ohio mixed-oak forests. Few long-term studies of carabid populations in North
American deciduous forests have been published. In the following chapters, trends in
32 carabid abundance, diversity, and assemblages will be presented and discussed. This information will help meet the goals of the EM project and increase our knowledge of carabid biology in southern Ohio’s mixed-oak forests.
33 CHAPTER 2
EFFECTS OF PRESCRIBED BURNING ON CARABID ABUNDANCE AND
SEASONALITY IN MDŒD-OAK FORESTS OF SOUTHERN OHIO
INTRODUCTION
The USDA Forest Service initiated an ecosystem management project (EM project) in southern Ohio in 1994. The objective of this long-term project was to determine the ecological response of mixed-oak forests in southern Ohio to two prescribed burning regimes. One goal of the EM project involved determining which regime, or prescribed burning treatment, was most appropriate for restoring and maintaining the structure, function, and composition of mixed-oak forests in southern
Ohio.
Multiple components of the mixed-oak community, such as soil dynamics, understory and overstory vegetation, resident bird populations, and several invertebrate taxa were monitored for impacts related to the fires. Groimd beetles (Coleoptera:
Carabidae) were one such taxon that was monitored beginning in 1995. Carabids are
34 well-suited for this kind of study because they are abundant, species-rich, easily collected, readily identified, and their biology is relatively well-known (Thiele 1977).
This family of beetles has been used in other studies to monitor the enviromnental impacts of management practices and ecological disturbances (Mossakowski et al. 1990,
Niemela et al. 1993, Lenski 1982. Rodriguez et al. 1998, Rushton et al. 1990). In this study, the effects of prescribed burning treatments on carabid abundance and seasonality were monitored to improve our understanding of the broader impacts of fire on the forest floor of mixed-oak communities.
The effects of fire on insects and other invertebrates have not been well studied
(McCoy 1987, McCullough et al. 1998). Most studies on the topic relate to the use of prescribed burning to control insect pests (Brennan and Hermann 1994, Komarek 1971,
Miller 1979). Of the studies that have been conducted on arthropod response to fire, most have been in prairie and other grassland habitats or coniferous forests. No studies have been published on the effects of fire on macroinvertebrates from a deciduous forest habitat.
Responses of prairie and grassland arthropods to periodic burning have been reviewed by Warren et al. (1987) and Reed (1997). Both of these reviews report that arthropod response varies depending on the taxon. Van Amburg ( 1981) also reported positive, negative, and neutral responses by arthropods to a single spring bum in tallgrass prairie. A study of belowgroimd arthropods in tallgrass prairie revealed no differences in total arthropod densities in aimually burned vs. unbumed sites (Seastedt 1984).
In sandplain grassland on Nantucket Island, Dunwiddie (1991) reported that a single April bum resulted in decreased spider abimdance, increased Orthoptera and
35 Homoptera abundance, and no consistent trends for Hymenopetra, Diptera, Lepidoptera, and Coleoptera. A study of oak savanna communities in Minnesota also foimd arthropod response to vary, but that arthropods in general, including beetles, were not greatly
impacted (Siemann et al, 1997). In Florida sandhill communities burned at intervals of one to seven years, no direct effects of fire on beetle abimdance, including carabids. were
detected (McCoy 1987).
Insect responses to fire in northern and boreal forests have been summarized by
Wikars ( 1997) and McCullough et al. ( 1998). These reports suggest that insect response
to fires in these habitats vary as well. A series of fire-effect studies in Australian
eucalypt forests found that spring and autumn fires reduced soil arthropod diversity
(Neuman and Tolhurst 1991, Springett 1976). However, low intensity autunm fires did
not have a discernable effect on the abundance of forest floor arthropods, especially
beetles (Collett 1998, Neumann et al. 1995).
Few studies have been conducted on the effects of fire on carabid abundance
specifically. In shrub steppe vegetation, burning had little impact on Calosoma luxatum
(Rickard 1970). Harris and Whitcomb (1971) reported that some carabid populations
increased and some decreased in abundance in pine stands burned annually vs. unbumed
stands in Florida. In boreal forests, wildfire does not seem to affect carabid species
richness, but species composition (McCullough et al. 1998) and relative abundance
(Holliday 1984, Holliday 1991, Richard and Holliday 1982) maybe altered. Holliday
(1992) found fewer individuals in burned areas vs. unbumed areas in spruce-aspen forests
of Manitoba. In tallgrass prairie, a spring bum was found to increase carabid abundance
(Van Amburg et al. 1981). None of these studies, however, had pre-fire data on carabid
36 abundance to compare to post-fire data. Even fewer studies have focused on changes in seasonal abundance of carabids due to fire, although Holliday (1991) suspected that seasonal patterns were not affected by burning.
The effects of fire on carabids in oak forests have not been reported, but this information is needed by forest managers and other researchers who weigh the use of prescribed burning with its side effects (Collett 1998). The objectives of this study were to 1.) document the abundance and seasonality of carabids existing on the forest floor prior to prescribed burning treatments, 2.) monitor carabid abundance and seasonality on each treatment unit, and 3.) evaluate the effects of prescribed burning on carabid abundance and seasonality.
The experimental hypothesis tested was that prescribed burning would cause changes in overall carabid abundance and/or seasonality in the burned areas. The null hypothesis was that the fires would have no effect on the beetle populations.
MATERIALS AND METHODS
The EM project was conducted in 80 to 120 year-old, mixed-oak forests in southern Ohio (Figure 1). These relatively undisturbed, second-growth forests are described in Chapter 1. Four study sites were chosen, ranging in size from 75 to 100 ha
(Table 1). The sites were known as Arch Rock (AR), Bluegrass Ridge (BR), Watch Rock
(WR), and Young's Branch (YB).
Each site was subdivided into three treatment units (approximately 30 ha each), creating a 4x3 randomized complete block. The fire treatments applied to each study area (Table 6) were “fiequenf ’ fire (burned annually), “infiequentf’ fire (burned every
37 Study Site Treatment Unit Treatment
Arch Rock A frequent B control C infrequent
Bluegrass Ridge A control B frequent C infrequent
Watch Rock A frequent B control C infrequent
Young’s Branch A control B frequent C infrequent
Table 6. Fire treatments applied to each treatment unit.
38 three years), and a “control” (unbumed). All fires were springtime bums, occurring in
March or April of 1996 through 1999, when fire conditions were optimal (Table 3). Each prescribed bum treatment generally resulted in low intensity surface fires, with flame heights less than 50 cm (Hutchinson, unpublished). However, fire intensity and
temperature varied within the treatment units due to topography, fuel distribution, and
microclimate. In a given year, small patches (ca. 10-50 m") of these forests did not bum.
Carabid populations were sampled via continuous pitfall trapping. A linear
transect of 12 traps, each trap approximately 10 m apart, was established along dry ridge
tops within each treatment unit (Figures 2 ,3 ,4 , and 5).
Each trap consisted of two white, plastic cups, with a diameter of 12 cm at the rim
and a depth of 14 cm, stacked together. The cups were placed together in the ground so
that the rims were even with the soil surface. Drainage holes were punched in the bottom
of the outer cup so that rainwater could drain and the inner cup would not float above the
soil surface.
Ethylene glycol (ca. 150 ml) was added to the inner cup as a killing agent and
preservative. A 20 cm x 20 cm masonite board was supported above each trap and held
in position by a 45 cm x 45 cm piece of hex netting (chicken wire). Traps were emptied
weekly in 1995,1996, and 1997 and every two weeks in 1998 and 1999.
Trap lines were operated &om late April/early May to late September/October
(Tables 5 and 6), averaging 146.5 trapping days in 1995, 180.75 days in 1996,153.25
days in 1997,152.5 days in 1998, and 152.5 days in 1999. In order to even out the
sampling effort in 1996 for multi-year comparisons, data from the last two collecting
dates in 1996 were omitted, resulting in a more comparable season of collecting and
39 sampling effort ( 152.75 average trapping days after adjustment of data). The Bluegrass
Ridge-A (control) treatment unit was not sampled in 1999.
Abundance and seasonality data were entered into Excel spreadsheets. Statistical analysis of these data was performed using S AS (S AS 6.12 for Windows; SAS 1996) to determine if significant differences existed in carabid abundance among fire treatments.
A mixed, factorial analysis of variance model was used in which ‘treatment’ was a fixed variable and ‘site’ and ‘year’ were random variables. Abimdance data were checked for normality with the Kolmogorov-S mimov Normality Test and, when necessary, were normalized with a log transformation before analysis.
RESULTS
Overall Abimdance
Over the five years of this study, a total of 18,346 adult ground beetles was collected by pitfall traps. When 1996 data were truncated to reflect a more even sampling effort, the total drops to 16,967 carabids (Table 7). Overall carabid abundance in each year declined sharply from 1996 to 1999 (Figure 6). No significant differences
(P< 0.05) in overall carabid abundance (using data adjusted for sampling effort) pooled over the five years were found among treatments. However, differences in carabid
abimdance among years were significant, especially in 1999 compared to previous years
(Table 8).
Only seven species constituted more than 3% of the 5-year total of carabids collected and only three species constituted more than 10%. The proportion of these
40 Site Unit 1995 1996 1997 1998 1999 Totals
Arch Rock A 932 746 389 102 33 2202 B 465 573 396 178 53 1665 C 685 430 247 137 64 1563
Bluegrass A 277 289 327 270 1163 Ridge B 316 317 559 392 37 1621 C 419 560 392 184 22 1577
Watch Rock A 430 395 162 60 11 1058 B 255 440 171 69 14 949 C 548 552 293 166 42 1601
Young’^s A 334 318 177 202 33 1064 Branch B 263 348 153 273 70 1107 C 243 685 185 255 29 1397
Totals 12 5167 5653 3451 2288 408 16967
Table 7. Overall carabid abundance collected by pitfall trapping on each treatment unit (1996 data is adjusted to reflect a more even sampling effort).
41 700
600 FREQ CONT 500 INFREQ 2CO XI 400 s "o 300 * I 200
100
0
1995 1996 1997 1998 1999 Year
Kgure 6. Average carabid abundance by prescribed burning treatment and year.
42 Source ^ F P treatment 2 0.41 0.6780 year 4 32.48 0.0001 treatment*year 8 0.30 0.9616
Table 8. Analysis of variance for overall carabid abundance (adjusted and log transformed).
43 common species among the overall species pool was sufficiently small that analyzing
their numbers separately was not warranted. Their abundance patterns followed that of
overall abundance.
Seasonal Abundance
In 1995 (prefire), carabid abundance reached its highest levels in June and again
in late September/early October on all 12 units (Figtires A1-A4 of Appendix). These 12
seasonal abundance curves were similar in that they demonstrated synchronous seasonal
peaks in abundance. In 1996, carabid abundance also attained its highest levels in June
and late September on all units and seasonal abimdance curves remained similar among
sites (Rgures A5-A8). In 1997, seasonal abundance curves were less synchronous
(Figures A9-A12). Carabid abundance was greatest in May rather than June on some
units. However, the late September peaks in abundance were still synchronized on all 12
treatment units. In 1998 and 1999, seasonal abundance curves were asynchronous
throughout the trapping year and completely lacked the higher levels of abundance in
September (Figures A13-A20).
hi some cases, the peaks in the abundance data were misleading. Some peaks at
season’s end were exaggerated because the trapping period was longer than previous
trapping periods (i.e. Figure A l). In other cases, carabid abundance was so low that two
or three beetles created a peak (i.e. Figure A19). So, comparison of seasonality curves
should be done cautiously when overall numbers are low. There were obvious
differences in these curves among years and within years, but there did not appear to be
any consistent trends related to fire treatments.
44 If overall carabid abundance is viewed seasonally as “Spring” (April, May),
“Summer” (June, July, August), and “Autumn” (September, October), there again did not appear to be any trends related to fire treatments (Figures 7-9). These seasonal divisions were arbitrary but similar to defined seasonal divisions used in other components of the
EM project (Hutchinson, pers. comm.).
Carabid abundance during spring (Figiure 7) was pooled for the five study years, and analyzed for treatment differences (no transformation of the data was necessary). No significant differences (P < 0.05) were found among treatments, but significant differences did exist among years (Table 9).
Carabid abundance during the summer months (Figure 8) was analyzed in the same maimer as the spring data, however a log transformation was needed to normalize the data. Again, no significant differences (P < 0.05) were found among treatments in the log transformed, pooled data. Significant differences did exist, however, among study years (Table 10).
Carabid abundance (log transformed) during the autumn months reflected the same results (Figure 9). No significant differences (P < 0.05) were detected among treatments but did exist among years (Table 11).
DISCUSSION
Pitfall Trapping
The use of pitfall traps to record and monitor carabid abundance must be done cautiously (Digweed et al. 1995). In actuality, these traps record carabid activity rather
45 80
70 - FREQ CONT 60 - INFREQ 2CO S S 40 - *5 %
20 -
10 -
1995 1996 1997 1998 1999 Year
Hgure 7. Average spring abundance of carabids by treatment and year.
46 500
FREQ 400 - CONT INFREQ
2 300 n S 8 •5 200
5 < 100
0 -
1995 1996 1997 1998 1999 Year
Rgure 8. Average summer abundance o f carabids by treatment and year.
47 500
FREQ 400 CONT INFREQ
2 300 sS •5 200
5 < 100
0 -
1995 1996 1997 1998 1999 Year
Rgure 9. Average autumn abundance of carabids by treatment and year.
48 Source ^ F P treatment 2 4.00 0.0789 year 4 2.89 0.0364 treatment*year 8 0.27 0.9700
Table 9. Analysis of variance for spring carabid abundance (adjusted).
Source FP treatment 2 2.14 0.1983 year 4 30.01 0.0001 treatment*year 8 0.80 0.6047
Table 10. Analysis of variance for summer carabid abundance (adjusted and log transformed).
Source FP treatment 2 0.15 0.8644 year 4 82.54 0.0001 treatment*year 8 0.27 0.9712
Table 11. Analysis of variance for autumn carabid abundance (adjusted and log transformed).
49 than abundance, but results can be used as estimates of abundance (Kharfaoutli and Mack
1991, Niemela et al. 1992). Continuous pitfall trapping results summed over the entire year provide a more accurate estimate of the size of carabid populations because they are less influenced by short-term, seasonal effects (Baars 1979, den Boer 1977).
In 1998 and. 1999. pitfall traps were emptied every two weeks rather than weekly.
This change in procedure should not have affected the number of carabids collected and most likely did not result in any differences in capture rates. Gryuntal (1982) reported no difference between emptying traps every seven days vs. once a month.
No immediate “digging-in effect” was observed in this study. This term refers to the fact that carabid catches are often highest just after the pitfall traps are placed in the ground (Digweed et al. 1995, Greensiade 1973).
Overall Abundance
This study captured an average of 3670 beetles/year. In a 2-year study of carabids in North Carolina oak-hickory and pine-oak-hickory forests, Lenski ( 1982) collected
4,727 carabids between June and October. A 1-year study in upland oak forests of
Michigan captured 2,019 carabids between April and October (Liebherr and Maher
1979). In 1990 and 1991,171 carabids were pitfall trapped in an upland mixed-oak forest in eastern Ohio (MacLean and Usis 1992). These other smdies contained fewer study sites and utilized periodic, rather than continuous, sampling methods. Direct comparisons of these results are not possible due to unequal sampling efforts among studies.
50 No significant differences were found in overall carabid abundance pooled over the five years among prescribed burning treatments. It is possible that the prescribed fires were of such low intensity and occurred at a time of year when few species were active that the beetles were unaffected by the fires. Beetles that were active at the time of the fires could have avoided the heat by burrowing 2 cm below the soil surface, climbing trees, or retreating to unbumed patches. Hansen ( 1986) reported that carabids survived the combustion phase of a range fire in Utah. It is doubtfiil that carabids were killed by the fires and other individuals re-colonized the burned units by the time pitfall traps were set.
Although no significant differences were found among treatments, there were significant differences among years, documenting large year-to-year changes in carabid abundance. The drastic decline in abundance between 1996 and 1999 (Figure 6) is striking, but not a direct result of prescribed burning. These changes are most likely the result of some other environmental stress acting on this indicator group.
Several possible explanations exist for this dramatic decrease in carabid abundance. One explanation could be that the large year-to-year changes in the abundance of specific species are normal fluctuations. Few studies have monitored carabid populations longer than two years, so little information is available regarding year-to-year fluctuations in carabid abundance (den Boer 1985, Holliday 1991).
Much of the decline in overall carabid abundance was attributed to the decline of a single species, Synuchus mpunctatus. This species dominated samples in 1995 and
1996, especially at the Vinton county sites, but was not collected at all by 1999 (Table
12). In an 11-year study of carabids in the forests of Manitoba, Holliday (1991) found
51 Study Area Unit 1995 1996 1997 1998 1999
Arch Rock A 565 429 42 0 0 B 126 301 79 2 0 C 78 181 8 0 0
Bluegrass A 41 30 15 2 - Ridge B 31 1 0 0 0 C 9 78 28 0 0
Watch Rock A 239 263 22 0 0 B 125 330 45 0 0 C 283 377 63 2 0
Young’s A 92 88 24 4 0 Branch B 76 93 10 0 0 C 57 151 9 0 0
Total 12 1722 2322 345 10 0
Table 12. Abundance of Synuchus impunctatus collected by pitfall trapping ( 1996 data are adjusted to reflect a more even sampling effort).
52 population levels of this species to peak approximately every three to four years.
Therefore, the decline in the abundance of this species during this study may have been a natural fluctuation.
Because Synuchus impunctatus constituted more than 25% of all carabids collected in this study, carabid abundance was re-analyzed in the absence of this species.
Log-transformed data were pooled over five years and no significant differences (P <
0.05) were found among treatments (Table 13), but did exist among years.
Another possible explanation for the decline in overall carabid abundance could relate to soil moisture levels. Precipitation totals for 1994 to 1999 recorded at the Vinton
County sites (Table Al) show that yearly totals flucmated from year-to-year (Figure 10).
1999 was widely considered to be a drought year in southern Ohio; however the total precipitation for that year was not unusual. Rainfall totals from May of each year (Figure
10), however, show a sharp decline between 1995 and 1999, which may have resulted in lower soil moisture and caused a subsequent decline in carabid populations and/or activity. Hot, dry conditions in Minnesota in 1980 vs. 1981 were used to explain lower activity-densities for some carabid species (Epstein and Kulman 1990).
Another explanation for the decline in overall carabid abundance could be over trapping or depletion of the local carabid fauna. Pitfall trapping without replacement has the potential to deplete local carabid populations (Digweed et al. 1995). With traps situated in the same location each year and beetles removed from these locations from
May to October for five years the local populations might have been impacted.
53 Source F P treatment 2 0.84 0.4765 year 4 22.20 0.0001 treatment*year 8 0.40 0.9106
Table 13. Analysis of variance for overall carabid abundance without Synuchus impunctatus (adjusted and log transformed).
54 Yeariy total May only 160 -
140 -
« 120 -
® 100 - 6 c 80 - ® “ 60-
40 -
20 -
1995 1996 1997 1998 1999 Year
Rgure 10. Précipitation levels recorded at The Vinton Experimental Forest (Hosack, unpublished).
55 Over-trapping of local carabid populations has been suspected in other pitfall studies
(Carrington, per. comm., Digweed et al. 1995, Luff 1975, Ostbye et al. 1978). Some researchers took precautions such as periodic sampling and live trapping to minimize the depletion of the carabid fauna being studied (Holliday 1992, Lenski 1982). At the time of this study two similar studies were being conducted in mixed-oak forests in West
Virginia and Kenmcky. Both of these studies show an increase in carabid abundance from 1997 to 1998 (Buss, pers. comm., Carrington, pers. comm.) while this study shows a dramatic decrease.
It is also possible that the decrease in carabid abundance was caused by a combination of natural fluctuations, low soil moisture, and trapping pressure. Certain
species may have been undergoing normal flucmations, while other species were
impacted by the low moisture, and other species were more sensitive to the continuous
trapping. To determine how each species may have been impacted, however, would be
difficult based on these data. In any case, prescribed burning treatments did not appear to
impact carabid abundance.
Seasonal Abundance
This study was not specifically designed to study seasonal trends in carabid
populations; however the data can be partitioned to give reasonable approximations of
seasonal abundance. Research conducted on the soil layers of these sites indicate that
soils on the burned units increase in temperature and dry mote rapidly than unbumed
units (Dress et. al 1999). It is reasonable to expect that these differences in soil
characteristics could have an impact on the seasonal abundance or activity of carabids.
56 However, no significant difierences were found among treatments during each of the three seasons studied. It is possible that the seasonality data (Figures AI-A20) are biased due to the low numbers of carabids caught, especially in the 1998 and 1999.
One of the most striking features of the seasonality data is the decline in carabid abundance during autumn. Much of this change can be traced to the decline in Synuchus impunctatus (Table 12). This species typically is most abundant in late September and dominated collections in 1995 and 1996, but was not collected at all in 1999.
Abundance o f Flightless Species
Synuchus impunctatus is a flightless species commonly found in forested habitats
(Lindroth 1961-1969). This species, as well as other flightless carabids, are more likely to reflect an impact of prescribed burning than flightworthy species because their reduced dispersal abilities are likely to restrict them to the treatment units. An analysis of the abundance of flightless species (log transformed for normality), however, revealed the same results as for overall carabid abundance (Figure 11). In the data pooled from the
five study years, no significant differences were found in flightless carabid abundance among prescribed burning treatments, but significant differences were found between years (Table 14). Possible explanations for these results have been discussed above.
Other EM Project Results
Results from other portions of the EM project indicate that these prescribed fires do impact the local environments in which the carabids exist. For example, leaf litter mass is significantly lower on the frequently burned units than on the other treatment
57 600
500 FREQ CONT INFREQ 400 HI *o % 200 I 100 -
0 -
1995 1996 1997 1998 1999 Year
Rgure 11. Average abundance o f flightless carabids by treatment and year.
58 Source ^ F P treatment 2 L03 0.4135 year 4 22.66 0.0001
treatment*year 8 0.54 0.8188
Table 14. Analysis of variance for overall flightless carabid abundance (adjusted and lo transformed).
59 units (Bœmer et al., in press). Diversity of understory vegetation has increased on the burned units vs. the controls (Hutchinson, unpublished). In terms of vegetation cover, the frequently burned units contain significantly more forbs and grasses and fewer seedlings and shrubs than the other treatments ( Artman, unpublished).
Despite these changes in the local habitat caused by prescribed burning, carabid abundance and seasonality do not appear to be impacted by the fires. These results may indicate that the carabid fauna in these mixed-oak forests is fire-tolerant and/or fire- avoiding.
Conclusions
Overall, the carabid abundance recorded in this study suggests that low intensity prescribed bums set in early spring do not significantly impact carabid activity on dry upland sites. However, these populations were seemingly impacted by other environmental variable(s), possibly low rainfall or trapping pressure resulting in declines of the local fauna.
Because prescribed burning does not significandy impact carabid abundance on dry upland sites, it is possible that populations of other beetles active on the forest floor of dry sites are not impacted either. These results suggest that prescribed burning may
not have devastating effects on carabid populations on dry sites and should continue to be considered as a management tool for restoring and maintaining mixed-oak forests in southern Ohio.
60 CHAPTERS
EFFECTS OF PRESCRIBED BURNING ON CARABID SPECIES RICHNESS AND
DIVERSITY IN MIXED-OAK FORESTS OF SOUTHERN OHIO
INTRODUCTION
In order to determine the ecological response of mixed-oak forests to prescribed burning, the USDA Forest Service initiated a long-term, multi-disciplinary project. This project, known as the ecosystem management project (EM project), began in 1994 and was conducted in mature, second-growth oak forests in southern Ohio (Figure I).
Multiple components of the mixed-oak community, including ground beetles
(Coleoptera: Carabidae), were monitored for impacts related to the fires. Carabids are commonly used in ecological studies as indicators of environmental conditions (Thiele
1977) and were well suited to help determine the impacts of prescribed burning on the invertebrate fauna occupying the forest floor.
Liformation regarding the effects of fire on surface-dwelling insects is lacking, especially 6om deciduous forests (Harris and Whitcomb 1974, McCullough et al. 1998).
Studies that have focused on the impacts of fire on insect abundance, particularly
61 carabids and other beetles, are reviewed in Chapter 2. These studies were conducted in a variety of habitats and report a variety of responses. Few of these studies have looked specifically at changes in beetle species richness and diversity as a result of fire.
In the Florida sandhill habitat, McCoy ( 1987) found little difference in the species richness of ground-dwelling beetles, including carabids, due to periodic burning. In
Australian eucalypt forest, low intensity fires had no effect on taxon richness of surface- active arthropods, including beetles (Collett 1998). Holliday (1992) reported that intense fires and/or the corresponding habitat changes that followed caused many carabid species to become locally extinct, but after a 10-year period following wildfire found no differences in species richness between burned and unbumed spruce stands.
No studies have been published on the effects of fire on surface-dwelling beetles
firom deciduous forests, yet this information is needed by forest managers and other
researchers for informed decision-making regarding prescribed burning (Collett 1998).
The objectives of this study were to 1.) document the species richness and diversity of carabids existing on the forest floor prior to prescribed burning treatments, 2.) monitor
species richness and diversity on each treatment unit, and 3.) evaluate the effects of
prescribed burning on carabid species richness and diversity.
The experimental hypothesis tested was that prescribed burning would cause
changes in overall carabid species richness and/or diversity on the burned units. The null
hypothesis was that the fires would have no impact on carabid species richness and/or
diversity.
62 MATERIALS AND METHODS
The EM project involved four study sites, ranging in size from 75-100 ha. located within 80-120 year-old, mixed-oak forests in southern Ohio (Table I). These sites, known as Arch Rock (AR), Bluegrass Ridge (BR), Watch Rock (WR), and Young's
Branch (YB), were subdivided into three treatment units, creating a 4x3 randomized complete block. The fire treatments (Table 6) were “frequent” fire (burned each spring),
“infrequent” fire (burned every third spring), and a “control” (unbumed).
Carabid populations were sampled via a linear transect of 12 traps established
along dry ridge tops within each treatment unit. Trapping was conducted from late
April/early May to late September/October (Table 5), averaging 146.5 trapping days in
1995, 180.75 days in 1996, 153.25 days in 1997, 152.5 days in 1998, and 152.5 days in
1999. Sampling effort in 1996 was reduced for multi-year comparisons by omitting the
last two collecting dates (referred to as “adjusted” data), which resulted in a more
comparable season of collecting and sampling effort ( 152.75 average trapping days). The
Bluegrass Ridge-A treatment unit was not sampled in 1999.
In addition to pitfall traps, a single 8-watt ultraviolet light trap was also operated
within each study block. Light traps were operated approximately one night/week in
1995, 1996, and 1997 and one night/two weeks in 1998 and 1999. Traps were hung
approximately 1-2 m off the ground and were powered by a 12-volt portable,
rechargeable battery. Light trapping was discontinued at Bluegrass Ridge-A in 1998 and
1999.
Many of the species collected in this study (approximately 50 %) were captured in
low numbers (five year totals < 20). These results are common in insect faunal studies
63 (Colwell and Coddington 1994). Due to this high number of “rare” species, Chao’s
(1984) estimator was used to estimate the “true” number of species for each unit in each year. This estimator (5/*j is a non-parametric method that was specifically developed for estimating true species richness of an assemblage based on the number of rare species in the sample. This estimator performs well on data sets with a high number of relatively rare species (Colwell and Coddington 1994). The Chao estimator is calculated as 5/* =
‘îobs + (a"/26), where 5obs is the observed number of species in a sample, a is the number of “singletons,” and b is the number of “doubletons.”
Species richness data and all calculations were entered into Excel spreadsheets.
Statistical analyses of these data and calculations were performed using SAS (SAS 6.12 for Windows; SAS 1996) to determine if significant differences existed in carabid species richness and diversity among fire treatments. A mixed, factorial analysis of variance was used in which ‘treatment’ was a fixed variable and ‘site’ and ‘year’ were random variables. All data were checked for normality with the Kolmogoro v-S mimov Normality
Test (except where noted) and were normalized with a log transformation when necessary.
RESULTS
Species Richness from Pitfall Traps
A total of 67 species was collected in this study by pitfall trapping (Table 15). In general, the number of species collected each year remained fairly constant, averaging
46.8 species/year. However, in 1999 only 29 species were collected.
64 Site Unit 1995 1996 1997 1998 1999 Total
Arch Rock A 23 24 31 20 7 37 B 25 25 23 21 10 35 C 26 20 24 19 7 35
Bluegrass A 14 14 18 18 — 22 Ridge B 14 15 20 15 11 31 C 17 17 22 22 7 28
Watch Rock A 23 25 21 15 8 30 B 21 21 19 16 8 29 C 24 27 23 24 7 37
Young’s A 13 18 21 20 12 28 Branch B 21 21 19 16 11 27 C 22 30 19 16 10 36
Total 12 45 49 49 44 29 67
Table 15. Carabid species richness collected by pitfall trapping on each treatment unit (1996 data is adjusted to reflect a more even sampling effort).
65 Three species collected by pitfall trapping were newly recorded for Ohio. These species were Carabus sylvosus, Cyclotrachelus incisus, and Piesmus siibmarginatus. A complete list of species collected by pitfall traps is shown in the Appendix (Table A2).
The mean (adjusted) carabid species richness collected from pitfall traps for each fire treatment in each year remained fairly constant within all three treatments until the decline in 1999 (Figure 12). No significant differences (P< 0.05) in carabid species richness (adjusted, pooled, and transformed) were found among treatments. There were, however, significant differences in species richness among years (Table 16).
Species Richness from Light Traps
A total of 99 species was collected by light trapping in this study (Table A3). Of these species,Agonum albicrus, Lebia collaris. Pentagonica and flavipes, Stenolophus dissimilis were newly recorded for Ohio.
Several problems were experienced while using the light traps. In a number of cases, technical problems resulted in the lights’ not deploying and carabids were not collected. In other cases, inspection of the samples suggested that lights did not remain lit for equal amounts of time on each unit. The result of these problems was that light trap data could not be analyzed quantitatively. Due to unequal sampling effort, and no reasonable method to equalize the effort, only qualitative comparisons could be made among units, treatments, or years.
However, the light traps collected carabids that were not captured by pitfall trapping. For this reason, and due to the continuing moth study, light trapping was
66 30
FREQ CONT 25 - INFREQ
S § 20 8- Q- *o * © 15
10 -
1995 1996 1997 1998 1999 Year
Figure 12. Average carabid species richness by treatment and year.
67 Source ^ F P treatment 2 0.66 0.5502 year 4 45.05 0.0001 treatment*year 8 1.10 0.3882
Table 16. Analysis of variance for pooled carabid species richness (adjusted and log transformed).
68 continued each year of the study, despite the limitations of the data, in order to generate
more complete species lists. Of the 67 species collected by pitfall trap and the 99 species
collected by Light trap, 18 species ( 12.2%) were collected by both methods.
Carabid Diversity
Carabid diversity (Table 17) was calculated for the pitfall trapping data using the
Shannon Index (H’). This index is one of the most commonly used measures of diversity
(Lande 1996, Magurran 1988). When these index scores are normally distributed it is
acceptable to use analysis of variance to compare the diversity of different communities
or units (Magurran 1988).
Carabid diversity scores (Figure 13) did not deviate significantly from normal (P
= 0.08 with the Ryan-Joiner W-test for Normality) and was compared among treatments
and years using a mixed model analysis of variance. Index scores (based on adjusted
data) were pooled over the five years. No significant differences (P < 0.05) were found
among treatments, but significant differences were detected among years (Table 18).
Carabid Evenness
Evenness refers to the equality of relative abundances of species within a sample.
The Shannon Diversity index incorporates evenness into its calculations, but an
additional measure of evenness (£) can be computed. In this case, E = H ’f In S whereS is
the total number of species in the sample. This evenness score ranges from 0 to 1.0, with
1.0 representing equal abundance of all species (Magurran 1988).
69 Site Unit 1995 1996 1997 1998 1999
Arch Rock A 1.613 1.543 2.729 2.112 1.644 B 2.102 1.731 2.273 2.365 1.618 C 1.903 1.806 2.592 2.146 0.814
Bluegrass A 1.815 1.839 1.869 1.831 — Ridge B 1.859 0.938 1.707 1.441 1.876 C 1.348 0.998 1.737 2.261 1.707
Watch Rock A 1.735 1.355 2.559 2.346 1.894 B 1.993 1.049 2.364 2.453 1.946 C 1.770 1.276 2.365 2.426 0.879
Young’s A 1.768 1.713 2.258 2.311 2.147 Branch B 2.347 1.994 2.462 2.108 2.013 C 2.424 2.115 2.579 2.173 1.829
Table 17. Carabid diversity for pitfall trapping data using the Shannon Index ( 1996 data are adjusted).
70 CONT INFREQ
1s o c c m co
Figure 13. Average Shannon Diversity for carabids by treatment and year.
71 Source ^ F P treatment 2 0.46 0.6491 year 4 12.63 0.0001 treatment*year 8 1.19 0.3317
Table 18. Analysis of variance for pooled Shannon Index scores (adjusted).
72 Overall carabid evenness (£) was calculated (Table 19) for the pitfall trapping data. The Anderson-Darling Normality Test determined these evenness scores to be normally distributed, allowing the use of analysis of variance (Magurran 1988). Carabid evenness (Figure 14) was analyzed using a mixed model of analysis of variance. In the adjusted and pooled data, no significant differences were found among treatments but were detected among years (Table 20).
Chao Estimates
Chao estimates of species richness were calculated from the adjusted pitfall trapping data (Table 21). When graphed (Figure 15), these calculated values reflect similar trends as the actual species richness collected (Figure 12), although with greater numbers.
Chao estimates were pooled over the five years, log transformed, and analyzed with a mixed model analysis of variance. No significant differences were found among treatments, but significant differences did exist among years (Table 22).
DISCUSSION
Carabid Diversity Studies
More studies of carabid faunas have been conducted in Europe than in North
America (Epstein and Kulman 1990). Of those conducted in North America, most have focused on the species richness or diversity of a particular area containing several habitats rather than focusing on one particular habitat For example, Purrington ( 1996)
73 Site Unit 1995 1996 1997 1998 1999
Arch Rock A 0.514 0.486 0.795 0.705 0.845 B 0.653 0.538 0.725 0.777 0.703 C 0.584 0.603 0.816 0.729 0.418
Bluegrass A 0.688 0.697 0.647 0.633 — Ridge B 0.704 0.346 0.570 0.532 0.782 C 0.476 0.352 0.562 0.731 0.877
Watch Rock A 0.553 0.421 0.841 0.866 0.911 B 0.655 0.345 0.803 0.885 0.936 C 0.560 0.387 0.754 0.763 0.452
Young's A 0.689 0.593 0.742 0.771 0.864 Branch B 0.771 0.655 0.836 0.760 0.839 C 0.784 0.622 0.876 0.784 0.794
Table 19. Carabid evenness for pitfall trapping data using Shannon Evenness (1996 data are adjusted).
74 FREQ CONT INFREQ
(o 0.9 - (Dc œ 0 .8 - til 0.5 - 0.3 9 9951996 1997 1998 19991995 Year Hgure 14. Average carabid evenness by treatment and year. 75 Source ^ F P treatment 2 1.38 0.3211 year 4 10.97 0.0001 treatment*year 8 0.88 0.5388 Table 20. Analysis of variance for pooled Shannon Evenness scores (based on adjusted data). 76 Site Unit 1995 1996 1997 1998 1999 Arch Rock A 24.1 25.0 44.5 35.1 9.0 B 31.1 26.7 24.6 30.0 11.5 C 30.2 31.5 33.0 25.1 9.3 Bluegrass A 16.0 14.7 27.0 20.0 — Ridge B 15.0 19.0 32.3 16.0 29.0 C 23.0 23.3 23.5 30.2 8.0 Watch Rock A 41.0 23.6 25.2 21.3 8.0 B 22.2 22.5 27.0 20.2 10.7 C 27.0 32.8 39.0 44.3 15.0 Young’s A 17.5 24.3 37.0 22.7 30.0 Branch B 29.0 21.0 23.2 16.2 12.0 C 26.5 35.4 21.3 17.0 22.5 Table 21. Estimates of “true” species richness using the Chao ( 1984) estimator ( 1996 data are adjusted). 77 FREQ 35 - CONT INFREQ © 30 - o <0 o % "O B I 1 10 - 1995 1996 1997 1998 1999 Year Figure 15. Average estimated species richness (Chao estimator) by treatment and year. 78 Source ^ F P treatment 2 1.38 0.32 II year 4 10.97 0.0001 treatment*year 8 0.88 0.5388 Table 22. Analysis of variance for pooled Chao estimated species richness. 79 reported 102 carabid species from Nanmcket Island and Will et al. (1995) listed 241 species from the island region of western Lake Erie and nearby mainland sites. When surveys concentrate on one habitat, species richness is lower due to the lack of habitat diversity. Lenski ( 1982) collected 43 species of carabids in forested and clearcut areas of the Blue Ridge Mountains in North Carolina. In mixed-oak-maple-hickory forests, Carrington (unpublished) collected 69 species. In Michigan, 37 species were collected in upland oak forests (Liebherr and Mahar 1979). At least 462 carabid species are known from Ohio (Usis and MacLean 1998). Extensive carabid surveys have been conducted in the northern half of the state (MacLean and Usis 1992, Purrington et al. 1989, Usis and MacLean 1998. Will et ai. 1995) but no published surveys exist from the southern half. In Stark county, 66 carabid species were captured from several habitats, including upland forest (Purrington et al. 1989). One hundred and one species were collected from marsh and forest habitats in Carroll county (Usis and MacLean 1998). In unglaciated Columbiana county, MacLean and Usis (1992) collected 49 species from three forest sites. Considering these published studies, the 67 species collected by pitfall trapping and 99 species collected by light trapping in this study seems relatively high, although this study collected for a longer time period than the other studies. Seven of the species collected in this study were state records for Ohio (Purrington and Stanton 1996, Purrington et al. 1999). Species Richness No significant differences in carabid species richness were detected among prescribed burning treatments in this study. The fixes occurred at a time of year when 80 few species were active and most were still overwintering in places protected from fire, such as below ground. Fires were of such low intensity that it is likely the beetles were not directly affected by the heat of the fires. It is possible tnat a few rare species were negatively affected, but that their abundance in the collections was too low to reflect the change. Overall carabid abundance was also unaffected by prescribed burning treatments (Chapter 2). Carabid Diversity The Shannon index is one of the most popular and widely used diversity measures (Lande 1996, Magurran 1988). In this study, H’ suggests that prescribed burning treatments do not significantly impact carabid diversity indices calculated from pitfall data. A source of error, however, can come from failure to include all species from a community in a sample (Magurran 1988), although this information is often not known (Lande 1996). In 1999, it is doubtful that all species of the communities were represented in the samples due to low overall numbers. Therefore, the 1999 calculations are not comparable to those of previous years. Carabid Evenness Although the Shannon index incorporates evenness into its calculations, additional measures of evenness can be calculated. Evenness values for this study indicate that prescribed burning treatments do not have a measurable effect on carabid diversity. These calculations operate under the same assumption as the Shannon index 81 (that all species are represented in a sample), so it is likely that 1999 values are compromised, as noted above. Chao Estimates The Chao estimator was used to determine if there were any effects of prescribed burning treatment on estimated species richness because so many species occurred as singletons and doubletons. No significant effects of prescribed burning were detected. The estimated species richness values for 1998 and 1999 were based on low numbers of carabids. These low numbers most likely did not represent an accurate picture of these carabid communities; therefore the Chao estimates are most likely misleading as well. Light Trap Data Determining the effects of prescribed burning on carabid species richness, diversity, and evenness is based solely on pitfall trap data. However, the results of this study show that the carabid fauna in these forests is much more species-rich than is recorded by pitfall trapping alone. Light trapping sampled a completely different carabid fauna, with only a 12.2 % overlap in species composition. Usis and MacLean (1998) collected 41 species in pitfall traps, 51 species in light traps, and 9 species (8.9%) with both methods. The authors indicated that differences in the two faunas reflected habitat preferences, flight capabilities, and sampling effort. Liebherr and Mahar ( 1979) also collected additional carabid species by utilizing additional collecting methods. Therefore, surveys aimed at complete carabid faimas need to utilize a variety of collecting methods. 82 Conclusions Based on the data recorded by pitfall trapping, low intensity spring bums do not appear to significantly affect the species richness, diversity, or evenness of surface-active carabids on dry upland sites. These results are somewhat surprising given that the fires have been found to significantly impact other aspects of these mixed-oak communities, such as soil pH and other properties (Boemer et al. 1998, Morris et al. 1998), leaf litter biomass and understory vegetation composition (Hutchinson, pers. comm.), and several bird species (Artman, pers. comm.). Although prescribed burning treatments did not appear to impact this carabid fauna, the beetles did appear to be impacted by some other environmental variable(s) in 1999. It is possible that other carabid faunas, such as those collected by light traps or those existing in the lower, mesic sites, might have been impacted by the fires. Because the fires did not affect carabid diversity, as measured by the Shannon index and measures of evenness, it is very possible that prescribed burning does not significantly impact other surface-active macro-invertebrates on dry sites. Preliminary results on spiders (Bradley and Shone, pers. comm.), scarab beetles (Smith, pers. comm.), silphid beetles, moths, and parasitic Hymenoptera (Horn, pers. comm.) from the EM project appear to confirm this conclusion, although longhomed beetles increase on fiequently and infrequently burned units (Osborne, pers. comm.). 83 CHAPTER 4 CARABID ASSEMBLAGES IN MDŒD-OAK FORESTS OF SOUTHERN OHIO INTRODUCTION The ecosystem management project (EM project) initiated by the USDA Forest Service in southern Ohio in 1994 provided the opportunity to monitor the carabid fauna of mixed-oak forests over a 5-year period. Studies of forest carabid assemblages are rare (Niemeia et ai 1992) and very few of these studies have monitored carabid populations for more than two years. Goulet (1974) sampled the populations of two carabid species for four years in various habitats in Alberta. Holliday (1991) sampled carabids in the same locations for 11 years after a forest Ere, but not on a continual basis as in this study, den Boer (1985) presented carabid data from 26 years of sampling. These long-term studies (3 or more years) allow for a more complete picture of carabid assemblages and changes than do one or two-year ‘snap shot’ studies. One of the primary goals of this study was to determine the impact of prescribed burning on carabid populations in order to address the overall objective of the EM project. The data collected indicate that low intensity spring fires do not significantly impact carabid abundance (Chapter 2) or species richness, diversity, or evenness (Chapter 84 3). With these results in mind, the dynamics of these carabid assemblages can be focused upon without the complicating factors of prescribed burning effects. In general, carabid assemblages are moderately species-rich, usually composed of 10-40 active species/habitat/season (Lovei and Sunderland 1996). Carabid populations are relatively stable in comparison to the variablity of other insects (Lovei and Sunderland 1996), but can exhibit large, year-to-year flucmations in abundance, dominance (relative abundance), and distribution (den Boer 1985, Holliday 1991, Niemela et al. 1992, Niemela et al. 1993). Spatial distribution of carabids within a habitat is rarely random and the causes of these aggregated distributions are not well understood (Luff 1986). Most researchers do not agree on which factors are responsible for the variations in distribution of carabid populations. On the regional scale (within the geographic range of a species), variation in distribution has been explained by species-specific adaptations to environmental conditions (Duffene and Legendre 1997, Niemela and Spence 1994). On the smaller local scale (within a habitat), causes of variation are a source of debate. Some researchers suggest that the distributions of carabids are influenced by soil moisture and other differences in the microhabitat (Epstein and Kulman 1990, Niemela et al. 1992, Thiele 1977). Others (Lenski 1984, Loreau 1986) consider interspecific competition to be important in structuring carabid assemblages, although convincing evidence is lacking (Lovei and Sunderland 1996). A third explanation has been related to behavioral responses, such as mating behavior and pheromones, that cause an aggregation in pitfall traps (Niemela et al. 1992). Geiger ( 1966) concluded that it is not possible to identify one factor as being responsible for the distribution of carabid species. 85 Niemela and Spence ( 1994) suggested that carabid distribution is determined by different factors on different scales and the effects of these factors do not necessarily translate between scales. Population numbers of carabids flucmate also, although little work has been done on the causes, den Boer ( 1985) concluded that Qucmation patterns in the population of a species vary based on the dispersal power of that species. In this chapter, the carabid assemblages sampled in this study are discussed on three spatial scales. The scales considered are regional, site, and local. The regional (or ecoregional) scale for this smdy refers to the unglaciated western Allegheny Plateau that extends into southern Ohio. All four smdy sites (Arch Rock, Bluegrass Ridge, Watch Rock, and Young’s Branch) are located within this region. On this scale, all carabids collected were grouped into one assemblage for dry forested sites. On the site scale, each smdy site is an individual unit (not subdivided into smaller treatment units). Therefore, four carabid assemblages were formed, one for each smdy site. On the local scale, each treatment unit (Arch Rock-A, Arch Rock-B, Arch Rock-C, ...etc.) is an individual unit, so 12 assemblages were considered. MATERIALS AND METHODS The EM project was conducted in 80-120 year old, mixed-oak forests in southern Ohio (Figure 1). Four smdy sites. Arch Rock (AR), Bluegrass Ridge (BR), Watch Rock (WR), and Young’s Branch (YB), were chosen ranging in size from 75 to 100 ha. Each area was subdivided into three units, approximately 30 ha each. 86 Carabid populations were sampled via continuous pitfall trapping and periodic light trapping. A linear transect of 12 pitfall traps, approximately 10 m apart, was established along dry ridge tops within each treatment unit (Figures 2-5). One 8-watt light trap was also operated within each unit. Light trapping was discontinued at Bluegrass Ridge in 1998 and 1999 and at Young's Branch in 1999. Further details of sampling protocols are described in Chapters 1 and 2. ‘Complementarity’ was calculated to show the distincmess of carabid assemblages at the site scale (Colwell and Coddington 1994). Complementarity (C) is calculated as C = Ujk/ whereSjk = the number of species in one assemblage (5)) + the number of species in the other assemblage (5*) - the number of species in both assemblages (V^t). Ujk =Sj^Sk-2 Vyt The scale for this measure is 0-100%, with 0% indicating identical assemblages and 100% indicating completely different ones. In order to determine if distinct assemblages truly existed and to identify the major source of variation, principle components analysis was performed on the pitfall trap data, after a natural log transformation. RESULTS Regional Scale At the larger regional scale, the list of carabids identified in this study by five years of pitfall trapping for the western Allegheny Plateau contained a total of 67 species (Table A4). Of the 67 species collected, 21 (31.3%) occurred every year (Table 23). A total of 99 species was collected by light trapping (Table A3), but year-to-year comparisons were inappropriate due to uneven sampling effort among years. The 87 SPECIES Apenes lucidulus Carabus goryi Cyclotrachelus incisus Cymindis limbatus Dicaelus dilatants Dicaelus elongatus Dicaleus politus Dicaelus piirpuratus Galerita bicolor Galerita janus Myas coracinus Notiophilus aenea Pasimachus punctulatus Pterostichus adoxus Pterostichus atratus Pterostichus sayanus Pterostichus stigicus Pterostichus tristis Rhadine caudata Sphaeroderus stenostomus lecontei Trichotichnus autumnalis Table 23. Carabid species collected every year by pitfall trapping (1995-1999). 88 majority of species (88 %) was not captured by pitfall trapping. These results indicate that the overall assemblage is larger than is reflected by pitfall trapping alone. Site Scale At the site scale, four groups of carabids were collected over the five years by pitfall trapping (Tables A5-A8). Of the 67 total species, only 11 (16.4%) occurred at least once at all four sites (Table 24), suggesting that differences existed in the assemblages among sites. The most species-rich groups were found at Arch Rock and Watch Rock (46 species each), followed by Young’s Branch (39 species), and Bluegrass Ridge (38 species). Four additional groupings of carabids were identified by light trapping (Table A9). Of these species, 34 (34.3 %) occurred at all four sites (Table 25), although sampling effort within each site varied over the course of the study. These results also indicate that differences existed among study sites. Based on these data. Watch Rock had the highest number of species (79), followed by Arch Rock (72), Young’s Branch (67), and Bluegrass Ridge (40). It is possible that these values reflect the uneven sampling effort spent at each site. Based on complementarity values calculated from the pitfall data for the four assemblages (Table 26), the Arch Rock and Watch Rock assemblages were the most similar (C = 33%), which is not surprising given that these areas are ca. 1 km apart. The next similar assemblage to these two was Young’s Branch, which had a C value of 40% compared to the Vinton county assemblages. Young’s Branch and Bluegrass Ridge were 89 SPECIES Apenes lucidulus Chlaenius emarginatus Cyclotrachelus convivus Dicaelus politus Galerita bicolor Myas coracinus Pterostichus tristis Rhadine caudata Sphaeroderus stenostomus lecontei Synuchus imptmctatus Trichotichnus autumnalis Table 24. Carabid species captured at all four study sites by pitfall trapping. 90 Species Agonum aeruginosian Amphasia sericea Apenes sinuatus Bembidion affine Calleida viridipennis Chlaenius tricolor Clivina americana Clivina bipustulata Colliuris pensylvanica Coptodera aerata Cymindis limatus Cymindis platicolis Dromius piceus Harpalus compar Harpalus erythropus Lebia analis Lebia atriventris Lebia grandis Lebia solea Lebia tricolor Lebia viridis Lebia viridipennis Notiobia terminata Oodes amaroides Platynus tenuicollis Plochionus timidus Selenophorus hylacis Selenophorus opalinus Stenolophus lecontei Stenolophus ochropezus Trichotichnus dichrous Trichotichnus vulpeculus Zuphium americanum Table 25. Carabid species captured at all four study sites by light trapping. 91 Arch Bluegrass Watch Young’s Site Rock Ridge Rock Branch Arch Rock 50% 33% 40% Bluegrass Ridge 50 % 50 % 46% Watch Rock 33% 50% 40% Young’s Branch 40 % 46 % 40 % Table 26. Complementarity values for each site assemblage based on pitfall data. 92 slightly less similar (C = 46%) and the biggest difference (C = 50%) was between the Vinton county assemblages and Bluegrass Ridge. Complementarity values were also calculated for the four assemblages recorded by light trapping (Table 27). These results were very similar to the pitfall trapping results. Arch Rock and Watch Rock were the most similar (C = 34 %), although the Young’s Branch assemblage was also very similar. The Bluegrass Ridge assemblage was again the most distinct grouping of species. Principle components analysis of the pitfall trap data resulted in an eigenvalue of .283 for axis 1 and an eigenvalue of .113 for axis 2. These values roughly correspond to 28% of the variation in species composition of assemblages explained by axis 1 and 11% of the variation explained by axis 2. Points along axis 1 appear to be grouped by study site and along axis 2 by year (Figine 16). This analysis shows that at least three distinct assemblages exist (Figure 17), namely the Bluegrass Ridge, Young’s Branch, and Vinton County groupings. The analysis also indicates that individual units were moving up axis 2, which is a reflection of the low numbers captured by this trapping method in 1998 and 1999. Local Scale At the local scale, 12 harvests were identified by pitfall trapping (Tables A5-A8). Species composition of these groups ranged from 22 species at Bluegrass Ridge-A to 37 species at Arch Rock-A (Table 15). Light trap data at the local scale was not included due to numerous mechanical problems that made sampling effort uneven and unit-to-unit comparisons invalid. 93 Arch Bluegrass Watch Young’s Site Rock Ridse Rock Branch Arch Rock 53 % 34 % 35 % Bluegrass Ridge 53 % 55 % 49% Watch Rock 34% 55% 40% Young’s Branch 35 % 49 % 40 % Table 27. Complementarity values for each site assemblage based on light trapping data. 94 yuC'M WH'C'M MCM ARBW BR-C-M ARAM WRBH VBUH YBCM YB«M VWAM YBAM fN ARC-M WRC#W%BW BR-C-M V) • • BRB-M • ARBW YBCBI YBCM A R C , " YBBM WR'B'VS Y B C M • YBAM BRB-M W H A », • C MT^M BR •vmwi BM BR-C'»» BH A M BRBMBRB», A RB»» W H A M A R B W »'f • Y B A », • • AHA*, YBAM * • BH'A'M YBAM AHA'»» ARAM Axis I Figure 16. Principle Componenls Analysis of pitfall carabid data. WRB VB-I WR- vo A R -SM • AR ÀwR-c-n l-M lAH-A-1 l-A'M Ax Figure 17, Carabid assemblages from pilfall trapping at each study siti DISCUSSION This study was designed to evaluate the effects of prescribed burning treatments on carabid abundance and diversity. Although these results provide some insight into the carabid communities existing on these dry upland sites, the data cannot accurately define the entire community or accurately reflect relative abundance, especially for the rare species. Regional Scale Species turnover (beta diversity) on this scale has been suspected in other studies (Holliday 1991, Purrington 1996). The pitfall trapping results from this study (Table A4) suggest that species composition changed from year-to-year, but in nearly every case the species gained and lost were rare and collected in very low numbers (< five individuals/year). A notable exception is Pterostichus relictus, which was not rare. Therefore, these changes from year-to-year most likely reflect “pseudo-turnover” because a species may not have become locally extinct but simply was not captured in a given year by this trapping method. Species turnover may have occurred within this assemblage, but data from pitfall and light traps are insufficient to support this conclusion. Site Scale The pitfall data indicate that differences existed in the carabid assemblages among sites. The Arch Rock assemblage contained six species unique to that site. The Bluegrass Ridge assemblage had seven unique species, and both Watch Rock and 97 Young’s Branch had five unique species (Table 28). These species were rare, collected in very low numbers (Brachinus americanus was an exception), and may not acmally have been as unique as the data indicate. Although these species were rare, it is interesting to note their apparent site-specific occurrence. Niemela et al. ( 1993) found that carabid assemblages differed between various forested sites, but these differences were attributed to forest age. Although much of the differences among site assemblages was attributed to rare species, the distinctness of each assemblage appears to be supponed. Possible explanations for these differences are suggested by the results of other research conducted at these same sites. The underlying geology at Arch Rock, Watch Rock, and Young’s Branch is similar, composed of Allegheny group sandstones and shales, with some interbedded limestone. However, the geology at Bluegrass Efidge is composed of Conemaugh group sandstones, shales, and clay shales, with more interbedded limestone (Morris and Boemer 1998). The soils differ among these sites as well (due, in part, to the geological differences). The Arch Rock, Watch Rock, and Young’s Branch sites all contain silt loams; however the Bluegrass Ridge site has more sandy soils and higher pH, Ca, Mg, and CarAl than the other sites. These differences have been attributed to the greater amounts of underlying limestone foimd at Bluegrass Ridge (Morris and Boemer 1998). 98 Arch Rock Amara aenea Brachinus tenuicollis Calosoma scrutator Harpalus erythropus Pterostichus moestus Scaphinotus unicolor heros Bluegrass Ridge Anisodactylus nigerrimus Bembidion pedicellatum Brachinus alternons Brachinus americanus Clivina bipustulatus Platynus hypolithos Trichotichnus dichrous Watch Rock Amara impuncticollis Cyclotrachelus freitagi Harpalus longicollis Pterostichus permundus Trichotichnus vulpeculus Young's Branch Agonum placidum Agonum punct^orme Bembidion quadrimaculatum opposition Calosoma wilcoxi Paratachys pumilus Table 28. Carabid species unique to each study site. 99 Variation in species composition of understory vegetation also exists among sites. Arch Rock and Watch Rock are the most similar in species composition, Bluegrass Ridge is the most distinct, and Young’s Branch is intermediate (Hutchinson et al. 1999). Carabid assemblages, as recorded by both pitfall trapping and light trapping, exhibit these same patterns. The principle components analysis confirmed that site-to-site differences were the main source of variation in these assemblages. This result is not surprising given the physical and vegetational differences among sites (described above). The results of this analysis also further supports the conclusion that low intensity surface fires do not have a measurable impact on carabid abundance or species richness on dry ridgetops. Local Scale On the local scale, species richness (as recorded by pitfall trapping) of these assemblages remained fairly constant, with the exception of 1999. The species composition of these assemblages varied, but as discussed above, these differences were largely due to the variable occurrence of rare species. Therefore, few conclusions can be drawn from these data. However, the most common one or two species in each assemblage also varied (Tables 29 — 32). These data show that the different assemblages are often dominated by different species and these dominating species tend to change hrom year to year. Little information is known regarding year-to-year changes in the relative abundance of carabid species. It is possible that these changes are a result of normal population fluctuations. 100 Year Arch Rock-A Arch Rock-B Arch Rock-C 1995 S. impunctatus (60.6%) G. bicolor (293%) G. bicolor (45.8%) G. bicolor (9.9 %) S. impunctatus (27.1%) P. punctulatus ( 19%) 1996 S. impunctatus (57.5%) S. impunctatus (523%) S. impunctatus (42.1%) EL caudata ( 12.6%) G. bicolor (173%) G. bicolor (18.8%) 1997 R. caudata ( 163%) G. bicolor (283%) G. bicolor (183%) P. tristis (12.1%) S. impunctatus ( 19.9%) P. punculatus ( 17 %) 1998 D. elongatus (373%) G. bicolor (253%) P. punctulatus (283%) R. caudata (18.6%) EL caudata (213%) R. caudata (19.7%) 1999 R. caudata (36.4%) P. punctulatus (52.8%) P. punctulatus (79.7%) Table 29. Species dominance on the Arch Rock units. 101 Year Blueerass Ridse-A Bluegrass Ridee-B Blueerass Ridee-C 1995 G. bicolor (453%) P. atratus (45.6%) P. atratus (64.9%) S. impunctatus ( 14.8 %) G. bicolor (10.8%) C. emarginatus (103%) 1996 G. bicolor (43.9%) P. atratus (77.6%) P. atratus (73.8%) P. tristis ( 15.6%) G. bicolor (6.0%) S. impunctatus ( 13.9%) 1997 G. bicolor (47.4%) P. atratus (41.1%) P. atratus (60%) P. tristis (12.8%) G. bicolor (233%) S. impunctatus (7.1 %) 1998 G. bicolor (493%) G. bicolor (59.7%) P. atratus (353%) D. teter/S.s. lecontei (113%) P. atratus (18.4%) G. bicolor (13.6%) 1999 G. bicolor (32.4%) P. atratus (31.8%) Table 30. Species dominance on the Bluegrass Ridge units. 102 Year Watch Rock-A Watch Rock-B Watch Rock-C 1995 S. impuncatus (55.6%) S. impunctatus (49%) S. impunctatus (51.6%) G. bicolor (14.9 %) C. goryi (9.4%) G. bicolor (12.4%) 1996 S. impunctatus (66.6%) S. impunctatus (75%) S. impunctatus (213%) P. tristis (8.6%) P. tristis (73%) P. tristis (63%) 1997 N. aeneus (14.8%) S. impunctatus (263%) S. impunctatus (213%) R. caudata/ N. aeneus (17%) C. goryi (17.1 %) S. impunctatus 113.6%) 1998 R. caudata (25%) C. goryi (17.4%) P. punctulatus (30.1%) D. elongatus/ D. politus/ C. goryi (13.9%) S.S. lecontei (13.3%) D. teter ( 13.0%) 1999 P. punctulatus (36.4%) C. goryi (28.6%) P. punctulatus (78.6%) Table 31. Species dominance on the Watch Rock units. 103 Year Young's Branch-A Young's Branch-B Young's Branch-C 1995 S. impuncatus (273%) S. impunctatus (28.9%) S. impunctatus (51.6%) P. tristis (26.6 %) G. bicolor ( 19%) G. bicolor (20.6%) 1996 G. bicolor (29.6%) G. bicolor (35.6%) G. bicolor (33.9%) S. impunctatus (27.7%) S. impunctatus (26.7%) S. impunctatus (22%) 1997 G. bicolor (27.1%) G. bicolor ( 19.6%) S. impunctatus (213%) P. tristis (20.3%) R. caudata ( 15.7%) S.s. lecontei (13 %) 1998 G. bicolor (32.7%) T. auumnalis (35.2%) G. bicolor (31%) P. tristis (13.9%) G. bicolor ( 17.9%) T. autumnalis ( 153%) 1999 A. lucidulus/ T. autumnalis (27.1%) T. autumnalis (44.8%) T. autumnalis (24.2%) Table 32. Species dominance on the Young’s Branch units. 104 Conclusions The carabid assemblages identified by pitfall trapping from dry upland sites on the local, site, and regional scales in this smdy all exhibit a high degree of variation from year-to-year; however most of this variation was due to low trapping frequencies of rare species. Despite the problems presented by rare species, three distinct carabid assemblages (Vinton County, Young’s Branch, and Bluegrass Ridge) were identified for dry sites on the site scale. These assemblages correspond to physical and vegetational variation among these same sites. Light trap data may also reflect these site differences (Table 28), although conclusions based on light trap data are tentative due to uneven sampling effort. Additional research is needed to better understand the dynamics of carabid communities, especially to address accurate sampling of rare species and analysis of species assemblages. A better understanding of carabid communities would allow for further interpretation of fire effects as well as increasing our knowledge of complex communities such as mixed-oak forests. 105 CHAPTERS GENERAL DISCUSSION This study originally proposed to address four objectives (Stanton 1996, unpublished). These objectives were to I) document the diversity and abundance of carabid species existing on the forest floor prior to controlled burning, 2) monitor these species on burned and unbumed units following prescribed burning, 3) evaluate the effects of the bums on carabid species richness, and 4) extrapolate the overall effects of prescribed burning on the forest floor community. In order to meet the first objective, fieldwork began in 1995 to define the carabid abundance and species richness on each treatment unit. These prebum data were important in that few studies on the impacts of fire contain prefire information. Carabid abundance and diversity were largely determined based on the results of pitfall trapping. Results from this method must be interpreted cautiously, as they measure carabid activity more so than actual carabid abundance. In this study, pitfall traps operated continuously fix>m late spring to early autumn, capturing over 5100 carabids representing 45 species in 1995. Although these data provided information on the carabid assemblages at each study site, they did not provide a complete picture of the fauna, as evidenced by the distinctly different carabid faunas collected by light trapping. 106 In order to fully document the diversity of carabids in an area, multiple collection techniques are required. The second objective required continuing the sampling procedure begun in 1995. Continuous pitfall trapping was conducted for four additional years, resulting in a long term data set. Few published smdies have monitored carabid populations for more than two years. These data illustrate a sharp decline in carabid abundance from 1996 to 1999. Species richness remained fairly constant from 1995 to 1998, but then underwent a decline in 1999. The cause(s) of these declines are unknown, although possible explanations include normal flucmations within species, adverse environmental conditions (such as low seasonal rainfall), trapping pressure, and combinations of these factors. In any case, the declines did not appear to be fire-related. The third objective requires statistical analyses of the species richness data. These analyses concluded that prescribed burning treatments did not have a significant effect in carabid species richness, diversity, or evermess. Although species richness did not vary much among years, species composition and dominance did vary from year to year. Due to the high number of rare species, conclusions regarding species turnover and carabid dynamics are limited, however it does appear that three distinct assemblages existed (Vinton County, Yoimg’s Branch, and Bluegrass Ridge). These assemblages correspond to physical and vegetational differences among these three areas. The fourth objective required carabids to serve as indicators for other forest floor taxa. Because it was determined that prescribed burning did not significantly impact the carabids captured on dry sites in terms of abundance and diversity, it is tempting to state 107 that the fires probably did not impact other forest floor invertebrates. However, no evidence is available that suggests that carabids as a whole can reflect the dynamics of other invertebrates. Additional work 6om this project, however, suggests that the fires did not greatly impact other members of the forest floor community, such as spiders, scarab beetles, and silphid beetles. Despite the potential limitations of the sampling method and the indicator role of carabids, these data support the conclusion that, in general, low intensity spring bums do not greatly impact most carabid species on xeric ridgetops. No other study has been conducted for this amount of time on carabid assemblages in oak forests of Ohio and is therefore valuable in characterizing carabid communities in deciduous forests and their response to prescribed burning. 108 APPENDIX 109 Year Month Precipitation (cm) Yearly Total fcmi 1994 Jan 17.04 Feb L0.I9 Mar 18.47 Apr 18.62 May 5.64 Jun 8.26 Jul 10.36 Aug 19.05 Sep 3.40 Oct 3.53 Nov 5.31 Dec 7.82 127.69 1995 Jan 11.89 Feb 5.05 Mar 4.19 Apr 5.44 May 20.04 Jun 7.87 Jul 4.67 Aug 13.72 Sep 4.29 Oct 11.63 Nov 4.80 Dec 6.91 100.50 1996 Jan 15.44 Feb 7.82 Mar 14.40 Apr 9.37 May 19.08 Jun 14.35 Jul 12.01 Aug 5.00 Sep 1131 Oct 4.42 Nov 12.34 Dec 8.59 134.33 Table A l. Precipitation totals recorded at the Vinton Experimental Forest (HosacK impublished). (contmned) no Table A l (continued) Year Month Precipitation (inches) Yearly Total 1997 Jan 8.69 Feb 11.13 Mar 18.49 Apr 4.34 May 12.45 Jun 8.38 Jul 7.75 Aug 9.47 Sep 2.90 Oct 3.48 Nov 8.61 Dec 4.19 99.88 1998 Jan 11.79 Feb 10.29 Mar 9.27 Apr 20.09 May 7.04 Jun 19.25 Jul 8.69 Aug 3.02 Sep 4.01 Oct 7.09 Nov 5.61 Dec 5.54 111.69 1999 Jan 16.97 Feb 9.73 Mar 8.43 Apr 6.99 May 4.70 Jun 2.18 Jul 8.18 Aug 13.69 Sep 2.64 Oct 7.95 Nov 10.95 Dec 8.26 100.67 III Species Agonum placidum (Say) Agonum pimctiforme (Say) Amara aenea (DeGeer) Amara impuncticollis (Say) Anisodactylus nigerrimns (Dejean) Apenes lucidulus (Dejean) Apenes sinuatus (Say) Bembidion pedicellatum LeConte Bembidion quadrimaculatum oppositum Say Brachinus alternons Dejean Brachinus americanus (LeConte) Brachinus tenuicollis LeConte Calathus gregarius (Say) Calathus opaculus LeConte Calosoma scrutator (F.) Calosoma wilcoxi LeConte Carabus goryi Dejean Carabus sylvosus Say state record Chlaenius emarginatus Say Chlaenius platyderus Chaudoir Cicindela sexguttata F. Cicindela unipunctata F. ClMna bipustuiata (F.) Cyclotrachelus convivus (LeConte) Cyclotrachelus freitagi Bousquet Cyclotrachelus incisus (LeConte) state record Cymindus americanus Dejean Cymindus limbatus Dejean Cymindus neglectus Haldeman Cymindus platicollis (Say) Dicaelus ambiguus Laferte-Senectere Dicaelus dilatatus Say Dicaelus elongatus Bonelli Dicaelus furvus Dejean Dicaelus politus Dejean Dicaelus purpuratus Bonelli Dicaelus teter Bonelli Galerita bicolor (Drury) Galerita Janus (F.) Table A2. List of carabid species collected by pitfall trapping. (continued) 112 Table A2 (continued) Species Harpalus calignosus (F.) Harpalus erythropus Dejean Harpalus longicollis LeConte Myas coracinus (Say) Notiophilus aeneus (Herbst) Notiophilus novemstriatus LeConte Paratachys pumilus (De jean) Pasimachus punctulatus Haldeman Piesmus submarginatus (Say)------state record Platynus hypolithos (Say) Pterostichus adoxus (Say) Pterostichus atratus (Newman) Pterostichus lachrymosus (Newman) Pterostichus moestus (Say) Pterostichus permundus (Say) Pterostichus reiictus (Newman) Pterostichus sayanus Csiki Pterostichus stygicus (Say) Pterostichus tristis (Dejean) Rhadine caudata (LeConte) Scaphinotus andrewsii mutabilis (Casey) Scaphinotus unicolor Harrisheros Selenophorus opalinus (LeConte) Sphaeroderus stenostomus lecontei (Chaudoir) Synuchus impunctatus (Say) Trichotichnus autumnalis (Say) Trichotichnus dichrous (Dejean) Trichotichnus vulpeculus (Say) 113 Species Acupalpus indistinctus Dejean Acupalpuspartiarius (Say) Aculpalpus pumilus Lindroth Agonum aeruginosum Agonum albicrus------state record Agonum fidele Agonum galvestonicum Casey Agonum harrisii LeConte Agonum lutulentum Agonum mutatum Agonum punctiforme (Say) * Agonum striatopunctatum Dejean Agonum tenue Amphasia sericea (Harris) Anatrichis minuta (Dejean) Anisodactylus carbonarius Apenes lucidulus (Dejean) * Apenes sinuatus (Say) * Badister maculatus LeConte Badister ocularis Casey Badister reflexus LeConte Bembidion affine Say Bembidion impotens Bembidion patruele Bembidion rapidum Bradycellus lugubris (LeConte) Bradycellus nigriceps Bradycellus rupestris (Say) Bradycellus tantillus (Dejean) Calathus opaculus LeConte * Calleida viridipennis (Say) Calosoma scrutator (F.) * Calosoma wilcoxi LeConte * Chlaenius emarginatus Say * Chlaenius sericeus Chlaenius tricolor Dejean Cicindela punctulata Cicindela sexguttata F. * Table A3. List of carabid species collected by light trapping. (Species marked with a * were collected by light trap and pitfall trap) (continued) 114 Table A3 (continued) Species Clivina americana Dejean Clivina bipustuiata (F.) * Clivina dentipes Dejean Clivina impressefrons Colliuris pensylvanica (LeConte) Coptodera aerata Dejean Cymindis limbatus Dejean * Cymindis platicollis (Say) * Dromius piceus Dejean Dyschirius erythrocerus LeConte Elaphropus ferrugineus (Dejean) Elaphropus xanthopus (Dejean) Galerita bicolor (Drury) * Harpalus compar LeConte Harpalus erythropus Dejean * Harpalus longicollis LeConte * Harpalus pensylvanicus Lebia analis Dejean Lebia atriventris Say Lebia collaris Dejean------state record Lebia fuscata Dejean Lebia grandis Hentz Lebia omata Say Lebia pulchella Dejean Lebia solea Hentz Lebia tricolor Say Lebia viridis Say Lebia viridipennis Dejean Lebia vittata (F.) Leptotrachelus dorsalis (F.) Loxandrus velocipes Casey Loxandrus vitiosus Allen Morion monilicomis (Latreille) Notiobia terminata (Say) Oodes amaroides Dejean Oodes brevis Panagaeus fasciatus Say Paratachys proximus (Say) Paratachys scitulus (LeConte) Patrobus longicomis (Say) Pentagonica flavipes (LeConte) ------state record (continued) 115 Table A3 (continued) Species Pentagonica picticomis Bates Perigona nigriceps (Dejean) Platynus cincticollis Platynus tenuicollis (LeConte) Plochionus timidus Haldeman Poecilus chalcites (Say) Schizogenius lineolatus (Say) Selenophorus hylacis (Say) Selenophorus opalinus (LeConte) * Stenocrepis cuprea Stenolophus comma (F.) Stenolophus conjunctus (Say) Stenolophus dissimilis Dejean------state record Stenolophus lecontei (Chaudoir) Stenolophus ochropezus (Say) Tetraleucus picticomis (Newman) Trichotichnus autumnalis (Say) * Trichotichnus dichrous (Dejean) * Trichotichnus vulpeculus (Say) * Zuphium americanum Dejean 116 Soecies 1995 1996 1997 19981999 Total Agonum placidum I 1 Agonum punctiforme I 1 Amara aenea I I Amara impuncticollis I I Anisodactylus nigerrimus L 1 Apenes lucidulus 100 30 13 20 20 183 Apenes sinuatus 2 2 1 5 10 Bembidion pedicellatum 1 1 Bembidion quadrimaculatum oppositum 1 1 Brachinus altemans I 4 5 Brachinus americanus I 4 49 12 66 Brachinus tenuicollis I 1 Calathus gregarius 2 22 28 52 Calathus opaculus 5 4 5 14 Calosoma scrutator 2 I I 5 Calosoma wilcoxi 2 2 Carabus goryi 147 86 148 90 12 483 Carabus sylvosus 43 17 39 5 104 Chlaenius emarginatus 87 16 30 25 158 Chlaenius platyderus 78 20 48 146 Cicindela sexguttata 5 2 I 8 Ciciiuiela unipunctata 1 2 5 4 12 Clivina bipustuiata I 1 Cyclotrachelus convivus 27 23 23 27 100 Cyclotrachelus freitagi I I I 3 Cyclotrachelus incisus 8 14 10 II 1 44 Cymindus americanus 11 6 8 4 29 Cymindus limbatus 4 2 I I 1 9 Cymindus neglectus 2 3 1 6 Cymindus platicollis I I 1 2 5 Dicaelus ambiguus 36 71 33 57 20 217 Dicaelus dilatatus 3 6 12 14 4 39 Dicaelus elongatus 7 18 23 58 6 112 Dicaelus furvus I 3 7 3 14 Dicaelus politus 132 161 97 143 7 540 Dicaelus purpuratus 33 31 34 44 28 170 Dicaelus teter 25 30 28 58 141 Galerita bicolor 1071 934 656 690 24 3375 Galerita Janus 8 11 9 3 2 33 Harpalus calignosus 1 1 2 Harpalus erythropus I I 2 Harpalus longicollis 2 2 Myas coracinus 131 87 99 30 5 352 Notiophiltts aeneus 104 78 240 43 22 487 Notiophilus novemstriatus 1 1 Paratachys pumilus I 1 Pasimachus punctulatus 263 141 149 128 121 802 Piesmus submarginatus 3 5 5 1 14 Platynus hypolithos 1 1 Table A4. Regional carabid assemblage collected by pitfall trapping. (continued) 117 Table A4 (continued) Soecies 1995 1996 1997 1998 1999Total Pterostichus adoxus 34 31 71 26 4 166 Pterostichus atratus 419 901 466 137 18 1941 Pterostichus lachrymosus 22 16 9 2 49 Pterostichus moestus I 1 Pterostichus permundus I 1 Pterostichus reiictus 66 76 21 163 Pterostichus sayanus 2 10 15 23 I 51 Pterostichus stygicus 2 37 98 25 1 163 Pterostichus tristis 374 513 252 39 3 1181 Rhadine caudata 85 299 266 200 31 881 Scaphinotus andrewsii mutabilis 9 4 2 15 Scaphinotus unicolor heros I 1 Selenophorus opalinus 3 1 4 Sphaeroderus stenostomus lecontei 137 74 107 114 1 433 Synuchus impunctatus 1722 3134 345 10 5211 Trichotichnus autumnalis 33 40 17 174 62 326 Trichotichnus dichrous 1 1 Trichotichnus vulpeculus I 1 Total 7162 9028 5448 4286 2407 18346 118 1995 1996 1997 1998 1999 Siississ A B C A B C A B C AB C B Amara aenea 1 Apenes lucidulus 5 4 6 3 3 5 2 3 1 2 Bracfilnus tenuicollis 1 Calathus gregarius 2 17 2 21 2 1 Calathus opaculus 2 1 2 4 Calosoma scrutator 2 1 1 Carabus goryi 11 10 11 1 5 7 11 6 14 1 6 1 Carabus sylvosus 31 1 10 8 2 4 32 4 2 1 1 Chlaenius emarginatus 2 1 2 2 1 1 Chlaenius platyderus 1 Cicindela sexguttata 1 2 Cicindela unipunctata 1 1 2 1 2 1 Cyclotrachelus convivus 1 3 1 1 2 Cyclotrachelus incisus 6 3 5 9 5 2 so Cymindus americanus 1 1 1 Cymindus limbatus 2 1 1 Cymindus neglectus 1 1 Cymindus platicollis 1 1 Dicaelus ambiguus 2 3 4 2 1 1 1 1 9 5 2 Dicaelus dilatatus 2 1 Dicaelus elongatus 3 3 4 6 38 2 4 Dicaelus furvus 1 Dicaelus politus 11 5 16 11 10 5 17 2 7 11 8 8 Dicaelus purpuratus 1 4 2 1 3 1 2 Dicaelus teter 6 3 2 5 3 2 1 1 5 2 Galerita bicolor 92 137 314 32 100 81 27 112 45 1 45 25 Galerita janus 1 1 2 1 1 Table AS. Carabid assem b lage captured by pitfall trapping at Arcfi Rock (1996 data are adjusted). (continued) Table AS (continued) 1995 1996 1997 1998 1999 Species A BC A B C A BC A BC A BC Harpalus erythropus 1 1 Myas coracinus 11 22 16 6 3 6 11 12 7 2 4 2 Notiophilus aeneus 27 20 13 10 10 13 44 43 27 1 8 3 7 5 2 Pasimachus punctulatus 4 64 130 2 27 68 14 35 42 8 22 39 2 28 51 Piesmus submarginatus 2 2 1 Pterostichus adoxus 5 2 11 2 3 1 13 10 3 7 3 1 Pterostichus atratus 1 Pterostichus lachrymosus 20 2 14 1 6 3 1 Pterostichus moestus 1 Pterostichus reiictus 34 24 2 33 16 8 11 1 Pterostichus sayanus 1 4 1 4 7 6 13 1 o Pterostichus stygicus 1 1 Pterostichus tristis 71 13 34 69 31 16 47 17 14 1 1 i 1 Rhadine caudata 7 9 7 94 28 29 64 32 30 19 38 27 12 2 6 Scaphinotus andrewsii mutabilis 2 Scaphinotus unicoior heros 1 Sphaeroderus stenostomus iecr 21 10 8 2 11 3 3 15 5 2 13 3 Synuchus impunctatus 565 126 78 429 301 181 42 79 8 2 Trichotichnus autumnalis 1 1 1 1 7 1 Total 932 465 685 746 573 430 389 396 247 102 178 137 33 53 64 # of species 23 25 26 24 24 19 31 23 24 20 21 19 7 10 7 1995 1996 1997 1998 1999 Species A B C A BC A B C A B C A B C Anisodactylus nigerrimus 1 Apenes lucidulus 13 5 2 1 1 1 — Apenes sinuatus 1 2 1 — 3 2 Bembidion pedicellatum 1 — Brachinus alternans 1 3 1 -- Brachinus americanus 1 4 1 44 4 9 3 — Calathus opaculus 1 -- Carabus goryi 2 2 10 9 — Chlaenius emarginatus 2 33 44 i4 3 1 1 12 8 1 7 3 — Chlaenius platyderus 1 — Cicindela sexguttata 1 — Clivina blpustulatus 1 — N Cyclotrachelus convivus 4 1 1 2 I 4 5 3 8 2 1 — Cymindus americanus 1 1 2 — Dicaelus ambiguus 1 Dicaelus dilatatus 2 3 1 6 2 1 5 — 3 Dicaelus elongatus 1 4 3 — 1 Dicaelus furvus 1 1 2 4 2 — 2 1 Dicaelus politus 5 17 15 2 7 23 2 4 18 14 17 21 — Dicaelus purpuratus 5 2 2 1 1 2 -- 2 Dicaelus teter 3 6 10 31 — Galerita bicolor 126 34 38 127 19 14 155 130 21 133 234 25 -- 12 1 Galerita Janus 1 2 2 2 1 1 — 1 Myas coracinus 13 15 18 13 5 4 12 13 10 2 6 4 -- 1 Notiophilus aeneus 4 2 20 5 2 -- Pasimachus punctulatus 1 —* Platynus hypolithos 1 -- Table A6, Carabid assemblage captured by pitfall trapping at Bluegrass Ridge (1996 data are adjusted). (continued) Table A6 (continued) 1995 1996 1997 1998 1999 Speçips A BC A B C A BCA B C A B C Pterostichus adoxus 1 4 16 1 12 Pterostichus atratus 144 272 2 246 413 1 230 235 72 65 - Il 7 Pterostichus sayanus 1 3 2 3 -- Pterostichus stygicus 1 12 93 4 1 20 1 -- 1 Pterostichus tristis 36 15 6 45 7 6 42 9 14 4 2 - Rhadine caudata 13 2 39 6 3 32 4 5 17 8 12 - 1 Scaphinotus andrewsii mutabilis 1 il 1 1 -- Sphaeroderus stenostomus lecontei 15 6 1 13 II 7 31 7 -- Synuchus impunctatus 41 31 9 30 1 78 15 28 2 -- Trichotichnus autumnalis 10 2 4 7 3 6 14 - 3 6 Trichotichnus dichrous 1 -- (y Total 277 316 419 289 317 560 327 559 392 270 392 184 _ 37 22 # of species 14 14 17 14 15 17 18 20 22 18 15 22 " Il 7 1995 1996 1997 1998 1999 Species A BC A BC A BC A B c: B Amara Impuncticollis 1 Apenes lucidulus 4 2 3 2 2 2 1 2 Apenes sinuatus 1 Calathus gregarius 1 2 2 Calathus opaculus 1 1 Carabus goryi 19 24 46 11 6 9 14 9 50 8 12 23 Carabus sylvosus 1 1 3 1 Chlaenius emarginatus 1 1 1 1 1 1 1 Cicindela sexguttata 1 1 Cicindela unipunctata 1 1 1 Cyclotrachelus convivus 2 1 2 2 1 B Cyclotrachelus freitagi 1 1 1 Cyclotrachelus Incisus 2 1 5 1 1 3 Cymindus americanus 1 5 2 2 1 2 3 1 Cymindus limbatus 1 Cymindus neglectus 1 1 1 Cymindis platicollis 1 Dicaelus ambiguus 6 3 4 2 5 1 1 5 2 Dicaelus elongatus 3 3 8 1 2 2 8 1 4 Dicaelus politus 10 7 6 2 12 8 3 5 5 2 9 12 Dicaelus purpuratus 1 9 4 1 1 2 6 3 8 4 5 6 Dicaelus teter 3 3 5 5 3 3 2 4 4 1 9 6 Galerita bicolor 64 16 68 21 16 24 21 9 27 3 8 22 Galerita janus 1 2 2 4 2 1 1 Table A7. Carabid assemblage captured by pitfall trapping at Watcfi Rock (1996 data are adjusted). (continued) Table A7 (continued) 1995 1996 1997 1998 1999 Species A BC A B C A B C A B c: A BC Harpalus caliginosus 1 Harpalus longicollis 2 Myas coracinus 3 2 3 5 2 2 6 3 11 2 4 2 Notiophilus aeneus 8 3 21 5 1 16 24 29 30 3 2 5 1 l Notiophilus novemstriatus 1 Pasimachus punctulatus 4 16 43 4 5 32 7 6 45 5 4 50 4 2 33 Piesmus submarginatus 2 1 1 Pterostichus adoxus 4 2 2 2 5 4 3 4 1 1 Pterostichus atratus 1 Pterostichus lachrymosus 1 N Pterostichus permundus 1 Pterostichus reiictus 3 2 1 Pterostichus sayanus 1 1 1 Pterostichus stygicus 2 Pterostichus tristis 24 20 37 34 32 36 14 21 18 1 1 Rhadine caudata 14 1 18 7 14 22 10 12 15 4 7 1 1 Scaphinotus andrewsii mutabilis 1 Selenophorus opalinis 3 Spheroderus stenostomus lecontei 13 6 11 4 6 1 15 2 2 7 8 1 Synuchus impunctatus 239 125 283 263 330 377 22 45 63 2 Trichotichnus autumnalis 4 4 1 1 2 1 4 2 2 Trichotichnus vulpeculus 1 Total 430 255 548 395 440 552 162 171 293 60 69 166 II 14 42 # of species 23 21 24 22 20 27 21 19 23 15 16 24 8 8 7 |W 5 | ‘)% l‘J')7 I'J'W ■Specius A H (’ A » C A H C A H C A I) Afuniitn pliiciduin Ayimiiiii punciiriiMiie Apunus liicidulii.s 17 25 H 4 5 .1 3 2 2 5 5 3 12 Bümtruliun quiitl. opposilui)) 1 ( ’iiliisoiiiii wiliiixi 1 1 C.'miihiis j5uiyi II III 2 (> 12 3 8 23 V () 15 Chliiunius uiniirginiiius 2 4 3 1 3 2 5 3 Chlaenius platyderus 7 71 2 17 15 32 Cicindela sexgullala 1 1 Cicindela unipunciala 1 Cyeoiraehelus convivus 6 5 2 7 .1 I 5 1 1 12 2 2 Cymindus americanus 1 1 1 1 1 Cymindus limhaius 2 1 Cymindus platicollis 1 I Dicaelus amhiüuus 1 8 0 II 48 1 7 7 1 25 18 I II I Dicaelus dilalalus 1 1 1 4 1 2 2 I Dicaelus elongatus 1 2 1 1 1 Dicaelus politus 1 17 21) 1 24 54 5 17 12 II 24 (> I I Dicaelus purpuratus •1 5 5 1 7 1 1 () 3 11 II I « 1 Dicaelus teter 1 1 5 1 2 2 1 Cialerita hieolor 82 5t) 511 VI 124 232 48 It) 31 IV 7V (lalerita janus 1 Harpalus caligiiiosus 1 Myas coracinus 2 II 12 1 1 13 5 0 2 2 Notiophilus aeneus 1 5 2 2 V V I V III 5 5 4 Baratachys pumilus Pasimachus punctulatus 2 2 Table A8. Carabid assemblage captured by pillall trapping at Young's Branch (1996 data are adjusted), (continued) Table A8 (continued) 1995 1996 1997 1998 1999 Species A BC A B C A BC A B c: A BC Piesmus submarginatus 1 1 1 1 Pterostichus adoxus 4 3 4 1 4 13 3 1 1 1 3 Pterostichus atratus 1 2 Pterostichus reiictus 1 1 1 Pterostichus stygicus 2 1 3 Pterostichus tristis 89 8 21 66 7 2 36 12 8 28 2 Rhadine caudata 16 4 12 25 16 6 16 24 15 21 16 16 3 2 3 Scaphinotus andrewsii mutabilis 1 5 1 1 Selenophorus opalinus 1 Spheroderus stenostomus lecontei 20 16 10 8 9 16 8 17 24 11 9 21 Synuchus impunctatus 92 76 57 88 93 151 24 10 9 4 Trichotichnus autumnalis 3 9 19 14 2 2 14 96 39 8 19 13 Total 334 263 243 318 348 685 177 153 185 202 273 255 33 70 29 # of species 13 21 22 18 21 29 21 19 19 20 16 16 12 11 10 Bluegrass Young’s Soecies Arch Rock Ridee Watch Rock Branch Acupalpus indistinctus X X Acupalpus partiarius X X Acupalpus pumilus X Agonum aeruginosum X XX X Agonum albicrus X X Agonum fidele X X Agonum galvestonicum X Agonum harrisii X X Agonum lutulentum X X Agonum mulatum X Agonum punctiforme X Agonum striatopunctatum X Agonum tenue X Amphasia sericea X XXX Anatrichis minuta X Anisodactylus carbonarius X Apenes lucidulus X X Apenes sinuatus X XXX Badister maculatus X XX Badister ocularis X X Badister reflexus X X X Bembidion affine X X XX Bembidion impotens X X X Bembidion patruele X XX Bembidion rapidum X X X Bradycellus lugubris X Bradycellus nigriceps X Bradycellus rupestris X X Bradycellus tantillus X X X Calathus opaculus X Calleida viridipeimis X X X X Calosoma scrutator X X X Calosoma wilcoxi X Chlaenius emarginatus X X Chlaenius sericeus X X Chlaenius ofcolor X XX X Cicindela punctulata X Cicindela sexguttata X X X Clivina americana X XX X Clivina bipustuiata X XX X Clivina dentipes X X Clivina imptesseSrons X X Colliuris pensylvanica X XX X Table A9. Carabid assemblages captured by light trapping. (continued) 127 Table A9 (continued) Bluegrass Young's Soecies Arch Rock Ridee Watch Rock Branch Coptodera aerata X X X X Cymindis limbatus X X X X Cymindis platicollis X X X X Dromius piceus X X XX Dyschirius erytiiocerus X Elaphropus ferrugineus X Elaphropus xanthopus X X Galerita bicolor X Harpalus compar X X X X Harpalus erythropus X X XX Harpalus longicollis X Harpalus pensylvanicus X X XX Lebia analis X X XX Lebia atriventris X X XX Lebia collaris X Lebia fuscata X X X Lebia grandis X X X X Lebia omata XX XX Lebia pulchella X XX Lebia solea X X X X Lebia tricolor X X XX Lebia viridis X X X X Lebia viridipennis X X X X Lebia vittata X Leptotrachelus dorsalis X Loxandrus velocipes X X X Loxandrus vitiosus X X X Morion monilicomis X Notiobia terminata X X X X Oodes amaroides X X X X Oodes brevis X Panagaeus fasciatus X Paratachys proximus X X Paratachys scitulus X X X Patrobus longicomis X Pentagonica flavipes X Pentagonica picticomis X X Perigona nigriceps X X P la n u s cincticollis X X X Platynus tenuicollis XX X X Plochionus timidus X X X X (continued) 128 Table A9 (continued) Bluegrass Young's Species Arch Rock Ridee Watch Rock Branch Poecilus chalcites X X X Schizogenius lineolatus X Selenophorus hylacis X X XX Selenophorus opalinus X X XX Stenocrepis cuprea X Stenolophus comma X X X Stenolophus conjunctus X X X Stenolophus dissimilis X Stenolophus lecontei X XXX Stenolophus ochropezus X XXX Tetraleucus picticomis X X Trichotichnus autumnalis X XX Trichotichnus dichrous X X X X Trichotichnus vulpeculus X X X X Zuphium americanum X X X X Total 72 40 79 67 129 5 00 FREQ 400 - CONT INFREQ 300 - 2 g 200 - o 100 - May Jun Jul Aug Sep Oct 1995 Figure A l. Pitfall collection data from Arch Rock, 1995. 160 140 - FREQ CONT 120 - INFREQ 100 - (0 TJ 1 80 - 8 60 - w 40 - 20 - May Jun Jul Aug Sep Oct Nov 1995 Figure A2. Pitfall collection data from Bluegrass Ridge, 1995. 260 FREQ CONT 200 - INFREQ 160 - (0 ■ o g 100 - *5 60 - JunMay Sep OctAugJul 1996 Figure A3. Pitfall collection data from Watch Rock, 1995. 140 FREQ 120 - CONT INFREQ 100 - ■8 80 - 15 2 g 60 - % 40 - 20 - JunMay Jul Aug SepOct Nov 1995 Figure A4, Pitfall collection data from Young’s Branch, 1995. 300 FREQ 250 - CONT INFREQ 200 - 0) T3 150 - S 8 *5 100 - 50 - May Jun Jul Aug OctSep Nov 1996 Figure A5. Piifall collection data from Arch Rock, 1996. 140 120 - FREQ CONT INFREQ 100 - w 80 - 1g 60 - 40 - 20 - May Jun Jul Aug Sep Oct Nov 1996 Figure A6, Pitfall collection data from Bluegrass Ridge, 1996, 250 FREQ CONT 200 - INFREQ 150 - tn ■o 1g 100 - o\ *6 50 - May Jun Jul Aug Sep Oct Nov 1996 Figure A7. PitfaJl collection data from Watch Rock, 1996. 160 140 - FREQ CONT INFREQ 120 - 100 - V) "O 80 - S 60 - *6 40 - 20 - May Jun Jut Aug Sep Oct Nov 1996 Figure A8, Pitfall collection data from Young’s Branch, 1996. FREQ CONT 60 - INFREQ S w 00 20 - May Jun Jul Aug Sep Oct 1997 Figure A9. Pilfull collection data from Arch Rock, 1997. 160 n 140 - CONT 120 - INFREQ 100 - (J) s ■8 80 - 8 *5 60 - sow 40 - 20 - 0 - May Jun Jul Aug Sep Cet 1997 Figure A 10. Pitfall collection data from Bluegrass Ridge, 1997. FREQ 60 - CONT INFREQ 50 - w 40 - S g 30- ê 20 - 10 - May Jun Jul Aug Sep Cet 1 9 9 7 Figure Al 1. PItfull collecllon cima from Watch Rock, 1997, FREQ CONT 30 - INFREQ 0) S 20- S May Jun Jul Aug Sep Oct 1997 Figure A 12, Pitfall collection data from Young’s Branch, 1997. FREQ 30 - CONT INFREQ 25 - 20 - S o 15 - 10 - May Jun Aug Sep OctJul 1998 Figure A 13. Pitfall collection data from Arch Rock, 1998. 100 FREQ 80 - CONT INFREQ 60 - g 40 - 20 - JunMay Jul Aug Sep Oct 1998 Figure A 14. Pilfull collcciion dulu from Blucgruss Ridge, 1998. FREQ 25 - CONT INFREQ 20 - 15 - O 10 - JunMay Jul Aug OctSep 1998 Figure A 15, Pitfall collection data from Watch Rock, 1998. FREQ CONT INFREQ w S 40 S May Jun Aug Sep OctJul 1 9 9 8 Figure A 16. Pitfall collection data from Young’s Branch, 1998. FREQ 25 - CONT INFREQ 20 - 0) S 15 - S S; *5 10 - May Jun Jul Aug Sep Cet 1999 Figure A 17. Pitfall collection data from Arch Rock, 1999. 14 - FREQ CONT 12 - INFREQ 10 - 8 *5 May Jun Jul Aug Sep Oct 1999 Figure A 18. Pilfull collection dulu from Bluegruss Ridge, 1999, 12 - FREQ CONT INFREQ 10 - CO "O !o 2 8 % JunMay Jul Sep OctAug 1999 Figure A 19. Pitfall collection data from Watch Rock, 1999, 16 14 FREQ CONT INFREQ 12 10 (0 *D ï 8 S 6 ê *6 4 2 0 May Jun Jul Aug Sep Oct 1999 Figure A20. Pilfall colleciion data from Young’s Branch, 1999. LIST OF REFERENCES Abrams, M.D. 1998, The red maple paradox. BioScience 42:346-353. Abrams, M.D. 1992. Fire and the development of oak forests. 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