Introduction

Home , Lasiurus, Nycticeius

Foraging Habitat Selection by Ohio Bats: An Examination between Eastern Second

Growth Forest, Eastern Old Growth Forest, and Pasture Land

A thesis presented to

the faculty of

the College of Arts and Sciences of Ohio University

In partial fulfillment

of the requirements for the degree

Master of Science

Richard T. Carter

March 2008 2

This thesis titled

Foraging Habitat Selection by Ohio Bats: An Examination between Eastern Second

Growth Forest, Eastern Old Growth Forest, and Pasture Land

RICHARD T. CARTER

has been approved for

the Program of Environmental Studies

and the College of Arts and Sciences by

Donald B. Miles

Professor of Biological Sciences

Benjamin M. Ogles

Dean, College of Arts and Sciences

Abstract

CARTER, RICHARD T., M.S., March 2008, Environmental Studies

Foraging Habitat Selection by Ohio Bats: An Examination Between Eastern Second

Growth Forest, Eastern Old Growth Forsest and Pasture Land (55 pp.)

Director ofThesis: Donald B. Miles

Bats are an ideal group of species to evaluate the effect of land management practices because their nesting and feeding habitats are often different and spatially separate. The presence or absence of bat species may indicate habitat quality. The use of different habitat types by different bat species was detected using echolocation calls. Five bat species were detected in SE Ohio. Myotis lucifugus was found in significantly higher numbers in the areas of Old Growth forest that areas thinning into Pasture habitat (eco- tone). M. lucifugus was also found to have significantly higher counts of feeding buzzes in this habitat type. Eptesicus fuscus was found in significantly higher numbers in the

Pasture habitat it was also found to be hunting in significantly high numbers in the

Pasture habitat. Lasionycteris noctivagans was found in significantly high numbers in the

Old Growth, Eco-tone, and Pasture habitats. L. noctivagans was not significantly found to

hunt in any particular habitat type.

Approved: ______

Donald B. Miles

Professor of Biological Sciences

Acknowledgements

First and foremost I would like to thank my advisor, Dr. Donald Miles, for his support and help without which this thesis would not have been possible. I would also like to thank the rest of my research committee, Dr. Kim Brown and Dr. John Zook, for their support and expertise which made this thesis possible. Thank you to Macdonald

Burgess for providing the topographic maps of the study site area. Thanks to my family and friends who urged me on when things looked like they were starting to get unbearable. And last but not least to Sarah who accompanied me on trips to my study site bringing along food and good cheer.

Table of Contents

Page

Abstract ...... 3

Acknowledgements ...... 4

List of Tables ...... 6

List of Figures ...... 7

List of Maps ...... 8

Introduction ...... 9

Study Site ...... 14

Materials and Methods ...... 16

Results ...... 20

Discussion ...... 23

References ...... 32

Appendix A: Tables and Figures ...... 35

Appendix B: Maps ...... 53

List of Tables

Table Page

1. Dates, description, and coordinates of sampled sites ...... 36

2. Bat species surveyed for at Dysart Woods ...... 37

3. Results from discriminant function analysis ...... 39

4. Average number of pulses per call for each species ...... 40

5. General activity counts of species in habitat type ...... 41

6. Feeding activity counts of species in habitat type ...... 42

7. Presence and feeding counts of all species surveyed for over the entire study period ..51

List of Figures

Figure Page

1. Varying clutter habitats of the forest ...... 35

2. Myotis lucifugus activity in all habitat types ...... 43

3. Eptesicus fuscus activity in all habitat types ...... 44

4. Lasionycteris noctivagans activity in all habitat types ...... 45

5. Myotis lucifugus feeding activity in all habitat types ...... 46

6. Eptesicus fuscus feeding activity in habitat types ...... 47

7. General and feeding activity time at half hour intervals for Myotis lucifugus in the eco- tone habitat ...... 48

8. General and feeding activity at half hour intervals for Eptesicus fuscus in the pasture habitat ...... 49

9. General activity at half hour intervals for Lasionycteris noctivagans ...... 50

List of Maps

Map Page

1. Entire study area ...... 53

2. Northern section of old growth forest ...... 54

3. Southern section of old growth forest ...... 55

Introduction

It is known that bats are not opportunistic foragers. Instead they exhibit behavior that indicates a preference for certain areas during foraging. The flight style and echolocation system used by bats are factors that are most likely to be responsible for a bats particular choice in foraging area (Neuweiler 1993). Variation in prey availability in

relation to habitat structure favors distinctive feeding morphology and behavior of bat

species. The interaction between prey distribution and complexity of habitat has been the

major factor in the evolution of different flight styles and echolocation system. The

diversity of echolocation styles as well as morphological diversity has allowed different

bat species to exploit the range of available biotopes (an area of uniform environmental

conditions). The biotopes exploited by bats may vary from completely open spaces with

fast moving prey that relies on speed rather than camouflage to avoid predation to areas

of thick vegetation with much slower moving cryptic prey. The specific echolocation and

flight styles used to exploit these differing biotopes include: bats with long slender wings

(high aspect ratio, low wing loading) that are fast flying using long range narrow

bandwidth echolocation calls (ideal for completely open spaces with fast flying prey that

are not relying on camouflage), and bats with shorter, broader wings using slow or

hovering flight (low aspect ratio, high wing loading) using short range broadband

echolocation which provides a higher resolution and thus more information (ideal for

areas around and in foliage where prey is slower and more cryptic due to the high clutter

background) (Norberg and Rayner, 1987; Neuweiler, 1993) (figure 1).

The call of a bat foraging in an open space will be one with a narrow frequency bandwidth (a narrow range of frequencies contained with in the call). This narrow bandwidth results in a more focused, energy concentrated beam and aids detection in open spaces. The focused beam is needed due to the larger distances between the bat and the potential prey item; a higher intensity focused beam will travel farther than one with a large bandwidth (Simmons et al, 1978). The focused beam will however have less ability to resolve target location compared to one that has a large bandwidth and therefore once the bat starts getting closer to the prey item it typically begins to emit a call with a wider bandwidth and thus is better able to pinpoint the precise location of the prey item. In contrast the call of bats that are hunting close to or gleaning of foliage typically have a wide bandwidth and therefore a high resolution necessary to identify a prey item close to foliage. The call’s energy can be spread over a wider bandwidth because a bat foraging in cover is relatively close to its prey, and does not need its call to travel as far as an open- habitat foraging bat.

Echolocation calls can be measured and the pattern can be used to characterize each individual pulse of the call (usually a frequency modulation that is a function of change in frequency over time). Measurements such as maximum frequency, minimum frequency, and duration of pulse are most often used to characterize an individual pulse in an echolocation call (Brodgers, et al., 2004; Murray et al, 2001; Wund, 2006). The

Brodgers (2004) and the Wund (2006) studies suggest that high clutter species tend to emit calls that were made up of pulses with higher starting frequencies, shorter durations, and greater band widths (which is a function of a higher beginning frequency and a lower

end frequency). In comparison low clutter species emitted calls composed of pulses with

lower starting frequencies, longer durations and lesser bandwidths. There is evidence

that bats in the same genus have similar calls (Jones and Holderied, 2007).

To accurately evaluate the effect of a different type of land management on

biodiversity it is necessary to do it on a species by species specific basis as well as at the community level (Patriquin and Barclay, 2003). This is particularly important with regard to bat species due the diversity of habitat needed by the different species of bat that may share a common site. With the increasing amount of habitat change (i.e. forest fragmentation, deforestation, forest degradation) the availability and quality of foraging habitat for bat species is being brought in to question. The effects of these habitat changes on bat assemblages are poorly known. The use of previous methods in documenting bat species in a locality is time consuming and often impractical. I used an alternative method to survey for bat species in a habitat mosaic.

My study relies on the ability to measure variation in echolocation calls among

species to better understand which bat species are exploiting the Old growth forest,

Second growth, Pasture land, and the Eco-tone habitat that make up the Dysart Woods

landscape. This study will also provide a bat species inventory of the Dysart Woods

Laboratory.

Old growth forest typically has a different physical structural complexity than

Second growth forest. Old growth forest is typified by larger natural gaps in the canopy

due to large tree falls, this provides for a heterogeneous over-story and a markedly varied

under-story (McCarthy, 1995). These characteristics are common, but not defining

characteristics (see study site section for defining characteristics of mixed oak eastern

Old growth) but are rather characteristics that tend to naturally occur in mixed oak

eastern Old growth. Such gaps in the under- and over-story are not common in most

Second growth forests. Canopy gaps are a feature that some non-clutter bat species may

take advantage of in their foraging. Dysart Woods is an example of Old growth

surrounded by both Second growth forest and pasture land. This area provides a

landscape mosaic with undisturbed areas as well as areas with varying degrees of

disturbance. Such variation makes Dysart a perfect study site to compare four land usage

types [Second growth forest, Old growth forest, Old growth/Pasture (Eco-tone), and

Pasture land].

My study is based on earlier work reported from the U.K. (Vaughan et al., 1997).

This study used acoustical recording methods to quantify habitat use (woodlands,

pastures, above lakes/rivers) of different bat species in the south west of Britain. Their

primary findings suggest that there were individual preferences between species as to which habitat they would hunt in/over. Based on their identification of bats habitat use, these authors suggested specific land management techniques aimed at preserving the habitat that is needed by specific bat species.

A more recent study by Ford et al., (2005) was carried out in the Central

Appalachians in West Virginia. The presence of bat species was related to forest

measures such as proximity to riparian areas, forest canopy width gap, and forest canopy

cover. Small-bodied species with a high start frequency (short pulse and large bandwidth)

(Myotis septentrionalis and Myotis sodalis) were found in forest regions with greater

canopy cover and closed riparian habitat while the small-bodied habitat generalists

(Myotis lucifugus and Pipistrellus subflavus) were primarily associated with larger water courses with open spaces. The larger bodied bats that make use of lower frequencies

(Eptesicus fuscus and Lasiurus cinereus) made use of open areas while the medium bodied bat with medium and high-frequency echolocation characteristics (Lasiurus borealis) was not associated with any particular habitat type, this probably being due to the fact that it is such a habitat generalist. A high point of the Ford study was the use of

Anabat acoustic technology that allowed accurate identification of bat species within their habitat.

A third study (Brooks and Ford, 2005) was conducted in central Massachusetts in

forest landscape that contained six different habitat types. Like the previous study (Ford

et al, 2005) large bodied bats such as Eptesicus fuscus were found significantly more

often in open canopy settings. Unlike Ford (2005), Brooks and Ford (2005) found the

small bodied generalist Myotis lucifugus to be ubiquitous in all habitat types. Myotis

septentrionalis was found in closed canopy riparian habitats in both studies. All three of

the above mentioned studies were successful in showing the power of using some form of

acoustical recording technique as a means to identify individual bat species in a particular area or habitat. Brooks and Ford (2005) and Ford et al. (2005) distinguish between small, medium, and large bodied bats and the correlation between these body sizes and the type of foraging habitat selected by the species. In this study I chose to make the distinction between fast flying bats and agile bats which can be correlated with wing shape and wing loading. This is because a bats agility or speed and thus foraging habitat choice is not

14 simply a matter of body size. Some big bodied bats can be quite agile flyers. In general , however, less agile bats tend to have long slender wings (low wing loading- less lift and less resistance) and slower more agile bats tending to have shorter broader wings (high wing loading- more lift and more resistance) (Norberg and Rayner, 1987). The correlation found by Brooks and Ford (2005) and Ford et al. (2005) may be related to the choice of bat species and the focus on body size rather than flight style, a correlation may be reflected in a small subset of bat species but not one that can be considered a rule.

Although my study considers many of the same species, my main distinction is between agile species (high wing loading) and fast species (low wing loading). The use of acoustic identification methods in my study will establish the presence of a bat species in a particular habitat as well as whether or not a particular species is foraging in a particular habitat type. A weakness of acoustic studies is the inability to estimate density of individuals. I am testing the hypothesis that there is greater bat diversity in the Old growth forest as well as more bat activity in the Old growth forest compared to the surrounding Second growth forest, Eco-tone, and Pasture land.

Study Site

The study was conducted at Ohio University Dysart Woods Laboratory located in

Belmont County, Ohio, which is a 20 hectare tract of old growth forest. Dysart Woods is classified as a mesophytic deciduous forest with cool, moist ravines and upland slopes that supports a high biological diversity (McCarthy, 1995). The Dysart Woods fit the profile of eastern mixed oak Old growth (no direct, stand initiating agriculture or logging

15 disturbance; trees that are near the maximum size and age for the species site; trees that exhibit low growth rates; trees that have boles without large branches or branch scares below the crown; trees that have flat top, spreading crowns; and stands that are all aged) surrounded by a mixture of Second growth forest and Pasture land (White and White,

1996).

The sites where data collection took place can be located by the use of a grid system that has been set up in Dysart as part of earlier studies (Burgess, 2006). The grid points are approximately 6 meters apart and can be located on the ground by the presence of stakes and ID tags. Four different habitat types were distinguished between and sampled. Second growth was represented by young small trees in relatively close association with one another. The Second growth canopy was relatively closed with very few or no openings. The Old growth forest was represented by much larger trees that were more widely spaced resulting in a more open spaced under-story. The old growth canopy contained large gaps created by tree falls. The Eco-tone habitat was an area were

Old growth forest thinned out into Pasture land resulting in large trees more widely spaced than the Old growth forest with no forest like canopy. The Pasture land was cleared of all trees and was used for cattle grazing or hay production (Refer to table 1 for site description and coordinates and refer to Appendix B for maps of site locations).

Seven species of bat were surveyed for. The species were selected based on the likelihood of their presence (Belwood, 1998) in Ohio as well as the availability of their sample calls in already established bat call libraries. Myotis sodalis (the Indiana Bat) was

the only species that I could not survey for due to the unavailability of sample calls.

(Refer to table 2 for list of species, flight and echolocation styles, and basic biology).

Materials and Methods

I employed digital recording technology to estimate species richness. Bat calls

were digitally recorded using Anabat II (Titley Electronics). Each unit contained an

ultrasonic microphone, amplifier, and a digital signal divider (Waldren, 2000). These

Anabat units were connected to laptop computers in the field which digitally stored calls

and saved then with a time and date stamp to a hard disk. Calls were then archived in the

laboratory on a Gateway p.c. hard drive and sorted into site location files. Data collection

took place from April until September, 2006

Within the four land use types described earlier the Anabat units were set up in

varying types of forest structure e.g.: along forest edges, in open pasture, within forest gaps, or within the under-story (Second or Old growth). When the units were placed in gaps they were positioned on a tripod on the ground at one end of the gap with the microphone pointing towards the far end of end of the gap. These gap units were angled so that the receiving cone area was almost entirely within the gap and not pointing out above the canopy (< 45○). The units placed in contiguous Second growth forest were also

placed on tripods on the ground and were angled as to have the majority of the receiving

cone below the bulk of the foliage. Units placed in fields were angled at forty five degrees with respect to the ground in order to have the maximum amount of receiving cone pointing in to the open space. All units at all sites were positioned approximately

1.5 meters off the ground, no units were placed in the canopy. Call identification was

correlated with location to make it possible to tell what species prefer specific forest

structure within the general forest setting.

Bat Species Identification

Three call characteristics were used to analyze each pulse: highest frequency in

the pulse, lowest frequency in the pulse, and duration of call pulse (Murray et al., 2001).

The characteristics were measured by using the Microsoft program Analyze, version 2.3,

Jolly (1999), calls with less than five pulses were not analyzed or recorded as per Murray et al. (2001). The recorded calls were assigned to a particular species based both on qualitative and quantitative techniques. The qualitative techniques are the same that are outlined by O’Farrell et al. (1999) with the major distinction being made between the

Myotis group and a group made up of a mixture of other genera (Eptesicus, Lasionycteris,

Lasiurus, Nycticeius, and Pipistrellus). These two groups are distinguishable by examining the highest and lowest frequencies of the pulses that make up a call. The

Myotis group tends to have a pulse that begins at a higher frequency (mid 50’s (KHz) and above) while the mixed genera (Eptesicus, Lasionycteris, Lasiurus, Nycticeius, and

Pipistrellus) group tends to have a beginning frequency that ranges from the mid 50’s

(KHz) down to the low 30’s (KHz) range. The mixed genera group can then be further divided by comparing the relative uniformity of calls. For example Lasionycteris noctivagans has a quite uniform call with little variation in minimum frequency while

Lasiurus cinereus tends to have one that fluctuates in minimum frequency. Averages for these characteristics were obtained from Murray et al. (2001). We developed a

classification function to statistically separate species based on call variables. Known

calls from an already established call library (UNM Bat Call Library) were used to

develop the classification function. A discriminate function analysis was run on the bat

call library to ensure that the selected call variables (highest frequency, lowest frequency,

and duration) did show variation between species; we employed a quadratic discriminate

function [(Proc Discrim), SAS, version 9.1] to accomplish this. We then entered

unidentified calls into the discriminate function to statistically identify the species to

known calls from the UNM bat call library. Pulses were only assigned to be a particular

species if they had a 0.7 or greater probability of being that species. Calls were only

assigned to a particular species if there was an obvious majority of pulses belonging to a particular species, otherwise they were considered to be unknown. Unknown calls were analyzed based on a visual reference (O’Farrell et al.,1999; Keinath, 2005). Using both qualitative and quantitative techniques allowed for greater precision in the assigning of calls to particular species.

The feeding buzzes were counted by converting the digital calls in to audio files

that played at an audible frequency using Anamusic, version 3.4 (Corben 2000) and by

listening to the recordings. The feeding buzzes or terminal capture phase were typically

clearly audible at the end of certain calls as the dramatic increase of pulse rate (Britton

and Jones, 1999). Using the audible version of the feeding buzz provided more precision

in identification compared to analyzing an entire call visually with Analook, version 4.8

(Corben, 2000) and trying to isolate the feeding buzz from the rest of the call. The

audible version provided a higher degree of clarity. The counts of identified species were

tallied for all sites. The average number of pulses per call was calculated for any species detected in a particular habitat type. This would give an indication of in which habitats each species was spending the most time. The assumption was that a higher average number of pulses per call of a species in one habitat type would indicate a species spending more time in that habitat type and not merely passing through, while a lower average number of pulses per call of the same species in a different habitat would

indicate that a species was just passing through and echolocating to navigate rather

hunting for prey. Bats that are in an area and are searching for prey may emit

significantly more pulses in order to accomplish a thorough search of that area, while a

bat just passing through an area may emit fewer pulses due to it spending less time in the

area. This, along with the presence of feeding buzz detections will give an indication of which habitat types are being used as hunting grounds and which habitat types are merely being passed through on the way to other hunting grounds.

Statistical Analysis

For each species a Chi-square analysis was carried out to determine whether or

not the counts within the habitat types were statistically occurring by chance or whether

the occurrences were not due to chance alone. The same was carried out for each species

that was detected emitting a feeding buzz. A one way, nonparametric ANOVA

(Wilcoxon) was run on sites within each habitat type in order to tell whether or not there

was a significant difference between sites within a habitat type, a difference which may

occur due to some sites of a particular habitat type being in the north facing forest and

some being in south facing forest. This would indicate whether or not significant activity

was limited to a particular site within a certain habitat type or present at more than one other site within that same habitat type. The bat calls and feeding buzzes of species that showed a significant preference for one or more habitat types were sorted in to a time table with half hour increments using the time stamps on the call files. When plotted this data gave an idea as to when the peak activity time and peak feeding time is for each species in each of the habitat types where they showed significant activity. A one way nonparametric ANOVA (Wilcoxon) was run on the average number of pulses per call for each species for each habitat type in which the species was detected.

Data Collection

Data sampling began at dusk and continued for approximately 4 hours (depending

on battery life). The choice and order of site sampled was random with no site being

samples more than once. It was difficult to sample all habitat sites during every sampling

night due to unforeseen technical difficulties, however at least two of the habitat types

being sampled during most sampling nights. Sampling took place once every two weeks

as conditions permitted. The topographic maps used were obtained from a previous

hydrology study done in Dysart Woods (Burgess, 2006).

Results

Bat Activity and Species Identification

The results for the discriminant function analysis of the known calls from the

UNM bat call library can be found in table 3. A total of 4012 pulses was from detected

304 calls (table 4). Five species of bat were identified (Proc Discrim function in SAS)

over all the habitat types with Myotis lucifugus being the species with highest number of counts recorded (124 counts). The other identified species included Eptesicus fuscus (73

counts), Lasionycteris noctivagans (67 counts), Myotis septentrionalis (5 counts), and

Lasiurus cinereus (4 counts) (Table 5).

Spatial Variation in Bat Activity

Bat calls were recorded in all habitat types as well as all sampled sites over the

course of the survey. There were a total of 31 unidentified calls (3 - Second growth

habitat, 3 - Old growth habitat, 1 - Eco-tone habitat, and 24 - Pasture habitat) (table 5). Of

the five bat species identified four were recorded emitting feeding buzzes with Myotis

lucifugus emitting the most (38 buzzes). The other species that showed feeding behavior

included Eptesicus fuscus (22 buzzes), Lasionycteris noctivagans (4 buzzes), and

Lasiurus cinereus (2 buzzes). There were a total of 13 feeding buzzes that belonged to

calls that were unidentified (1 – Second growth habitat, 2 – Old growth habitat, 4 – Eco-

tone habitat, and 6 – Pasture habitat) (table 6 and 7).

Habitat Preference

Of the five species identified, 3 showed significant preference for one or more particular habitat type (table 5). Myotis lucifugus while being detected in all habitat types was most significantly found in the Eco-tone with 84 counts (table 5 and 7, figure 2) (χ2 =

47.75, df = 3, P< 0.001). Eptesicus fuscus was detected in all habitat types but was most

significantly found in the Pasture land with 63 counts (table 5 and 7, figure 3) (χ2 =

115.69, df = 3, P< 0.001). Lasionycteris noctivagans was detected in all sites but had the highest numbers in the Old growth (19 counts), Eco-tone (23 counts), and Pasture land

(24 counts) sites (table 5 and 7, figure 4) (χ2 = 16.73, df = 3, P < 0.001). Myotis

septentrionalis and Lasiurus cinereus were rarely detected and could not be shown to

statistically favor any particular habitat type, they however were each only found in one

habitat type, Second growth and Pasture respectively (table 1).

There was no significant average number of pulses per call for any species in any

habitat type: Eptesicus fuscus (χ2 = 3, df = 3, P < 0.3916), Lasiurus cinereus (χ2 = 3, df =

3, P < 0.3916), Lasionycteris noctivagans (χ2 = 3, df = 3, P < 0.3916), Myotis lucifugus

(χ2 = 3, df = 3, P < 0.3916), Myotis septentrionalis (χ2 = 3, df = 3, P < 0.3916)

Two species showed significance preferences in feeding behavior (table 6).

Feeding Myotis lucifugus were found in the Old growth, Eco-tone, and Pasture sites but

showed a significant preference to the Eco-tone with 28 feeding buzzes being recorded

(χ2 = 21.38, df = 3, P < 0.001) (figure 5). Feeding Eptesicus fuscus were only recorded in

the pasture sites with 22 feeding buzzes being recorded (χ2 = 26.4, df = 3, P < 0.001)

(figure 6). Both Lasionycteris noctivagans and Lasiurus cinereus were detected feeding

in the pPasture type habitat but not in significant numbers. There was no significance

between counts from any sites within a particular habitat type (χ2 = 10.4, df = 8, P>0.232

(Wilcoxon)).

Phenology

The three species that showed significant presence in specific habitats were

plotted by using the counts of those species in the particular habitat which were then

sorted in to the half hour increments in which they were recorded. Myotis lucifugus had

significantly high numbers in the Eco-tone habitat with counts beginning to rise at 21:00

23 and peaking between 22:00 and 22:30 and then tapering off between 23:00 and 23:30 and remaining low after that (figure 7). Eptesicus fuscus was found to have significant activity in the Pasture habitat with activity beginning to increase after 20:30 and peaking between 21:00 and 21:30 and then tapering off between 21:30 and 22:00 and remaining relatively low after that (figure 8). Lasionycteris noctivgans was found to have significantly high activity in Old growth, Eco-tone, and Pasture habitats. The activity in the Old growth habitat peaked between 20:30 and 21:00 and then began tapering off between 21:00 and 21:30 and stayed relatively low after that (figure 9). The activity in the Eco-tone habitat began peaking between 21:00 and 21:30 and started tapering off between 21:30 and 22:00 and stayed relatively low after that (figure 8). The activity in the pasture habitat started to rise between 20:30 and 21:00 and peaked between 21:00 and

21:30 and then began tapering off between 21:30 and 22:00 and stayed relatively low after that (figure 8).

The two species that showed a significant preference in their feeding sites were plotted the same way as the general activity counts. The feeding activity of Myotis lucifugus began to rise between 21:30 and 22:00 and peaked between 22:00 and 22:30 and began tapering off between 22:30 and 23:00 and stayed relatively low after that

(figure 7). The feeding activity for Eptesicus fuscus began to increase just after 20:30 and peaked between 21:00 and 21:30 and began to taper off between 21:30 and 22:00 with no activity after that (figure 8).

Discussion

Activity and Habitat Preference

My data suggest that Myotis lucifugus while being found in all habitat types and being found to forage in three of the four habitat types generally preferred the Eco-tone habitat, which afforded greater gap size than the Old growth sites while not being as open as a field or pasture. This pattern is in agreement with Ford et al. (2005) and Brooks and

Ford, (2005) who found that Myotis lucifugus tended to prefer more open spaces compared to that of other Myotis species but not the wide open spaces that the faster less agile bats tend to prefer. Although there was a tendency for Myotis lucifugus to show significant preference to the more open wooded areas this did not stop it from exploiting other habitat types. This pattern agrees with Brooks and Fords (2005) assessment that

Myotis lucifugus while showing a preference for more open spaces can be a generalist and may exploit many habitat types. My study indicates that Myotis lucifugus tends to prefer larger gaps in which to forage with the prime habitat and gap sizes being in the areas that consist of Old growth forest that is merging in to pasture habitat but with pasture land being preferred over pure Old growth forest (table 1). This selection by

Myotis lucifugus of the larger gaps, which are created as a result of old growth forest that is thinning in to pasture land, may be due to the likelihood these gaps act as insect funnels in and out of the forest. The bats possibly optimize their foraging by utilizing these naturally formed funnels as foraging habitat. In a third study by Jung et al. (1999), involving old growth Pine forests, it was apparent that Myotis lucifugus and Myotis septentrionalis tend to spatially separate for feeding. Although I failed to detect any feeding activity by Myotis septentrionalis in the Second growth habitat it was the only habitat type in which I detected their activity. This pattern may suggest that there is

indeed a spatial separation in the foraging activity of Myotis lucifugus and Myotis

septentrionalis. These two species may separate their foraging habitat along the basis of

gap size in the forest canopy with Myotis septentrionalis preferring small gap size (5 and

6 in figure 1) and Myotis lucifugus preferring much larger gaps that merge in to pasture

land (pink label 2, light blue label 3, and dark blue label 1 in figure 1). Myotis

septentrionalis was found to prefer relatively closed canopies in a West Virginian

Appalachian study (Owen et al., 2003). It was suggested that land management practices

that promoted closed canopies will promote and improve Myotis septentrionalis foraging

habitat. This preference by Myotis septentrionalis for more closed forest was also noticed

in the Brooks and Ford, (2005) study which described Myotis septentrionalis as being a high clutter species. The preference of forest habitats by these two species is due to their larger bandwidths [calls that will attenuate (weaken) quickly due to the energy of the call

being divided into many frequencies] which are more effective in smaller areas where the

sound is not traveling as far but can achieve a higher resolution (due to the many frequencies) and therefore be more precise in prey targeting. Myotis septentrionalis emits

calls with the widest bandwidth of all Ohio Myotis species. This call pattern suggests that

they may be more effective hunters in the much smaller aerial gaps where high resolution

is a selective advantage but long range is not (Belwood, 1998; Murray et al., 2001). M.

lucifugus tend to emit the narrowest bandwidth of all the Myotis species found in Ohio.

This call type is likely to be more suited for more open gap forest habitat where the distance that the sound waves are traveling are farther than that of the small gaps in

Second growth forest, this species however does posses the ability to adjust the

26 bandwidth of its echolocation calls which may allow it to exploit other habitats with differing gap sizes. Generally however, M. lucifugus prefers larger gap forest habitats

(Brooks and Ford, 2005). This alteration of call may allow a greater effective range of the call or it may allow a higher resolution but a shorter effective range. The differing pattern of echolocation between Myotis septentrionalis and Myotis lucifugus may result in differing prey detection ranges between the two species. Myotis septentrionalis might be expected to have a shorter perceptual distance compared to that of Myotis lucifugus

(Brodgers et al., 2004). Differences between Myotis lucifugus and Myotis septentrionalis also extend to the pattern of prey capture that they employ. Myotis lucifugus are known to be aerial insectivores which would do better in open spaces while Myotis septentrionalis is known to be a gleaner species and would do better in an environment with many surfaces off which to glean insects (Belwood, 1998; Patriquin and Barclay, 2003). These differences in feeding strategy may be ultimate driving force in the spatial separation of foraging habitats between these two species.

Eptesicus fuscus was detected in all habitat types but with the over whelming number of detections being the Pasture habitat with all the feeding activity in the Pasture habitat (dark blue label 1 in figure 1). These findings are in agreement with Ford et al.

(2005) and Brooks and Ford (2005) which found that Eptesicus fuscus prefers open areas in which to forage. This habitat choice may be attributed to the fact that it is a fast flying less agile bat with a narrower bandwidth call, effective in open areas.

Lasionycteris noctivagans was detected in all habitat types with relatively equal counts in the Old growth (pink label 2 and light blue label 3 in figure 1), Eco-tone and

Pasture habitats (dark blue label 1 in figure 1) and a single count in the Second Growth

habitat. Feeding activity was only detected for Lasionycteris noctivagans in the Pasture

habitat (dark blue label 1 in figure 1). Feeding activity limited to Pasture habitat is in

agreement with Patriquin and Barclay (2003) that found that Lasionycteris noctivagans

only foraged in clear cut areas of the forest and no where else. However the relatively

high numbers of vocalization counts in the Old growth and the Eco-tone habitat coupled without feeding activity in those same areas is unexpected. This observation is consistent with the idea that this bat species is one that is a fast flying less agile species.

Lasionycteris noctivagans predictably makes use of open foraging habitat where it uses narrow bandwidth calls that are best suited for long rang detection. The high counts of

Lasionycteris noctivagans in areas where no feeding activity was recorded maybe the result of individuals passing through the area on their way to and from the more open feeding sites.

Lasiurus cinereus was only detected in the Pasture habitat (dark blue label 1 in

figure 1), this was not significantly high. Of the four detections, three of the calls

contained feeding buzzes indicating that Lasiurus cinereus does feed in the more open

Pasture habitat types. This observation is in agreement with Ford et al., (2005) who also

found Lasiurus cinereus to be an open habitat forager. Again these observations are

consistant with the idea that this species is a fast flying, less agile one that makes use of

narrower bandwidths while foraging.

The lack of significance for any of the average number of pulses per call for any

species in any habitat type may suggest that average pulses per call is an inaccurate way

of telling if a bat species is spending any significant amount of time in a particular habitat

type or just merely passing through. It is possible that an individual bat has to emit a

certain amount of pulses per call in order to gather enough information about its

surroundings whether or not it is merely navigating its way through a habitat type or

whether it is actively searching for and pursuing prey. The number of pulses needed by a

navigating bat and a hunting bat may not be significantly different. Further investigations

correlating call average with actual behavioral observations would have to be carried out

in order to verify if the number of pulses per call vary between hunting and navigating

bats.

Phenology

The calls for the significant habitat types for each species were plotted to indicate

at what time they were most active in identified habitat sites. Myotis lucifugus was found

to have the most significant activity in the Eco-tone habitat which began to increase between 21:00 and 21:30 and peaked between 22:00 and 22:30 and then tapering off between 22:30 and 23:00. This finding agrees with Belwood, (1998) who claims that

Myotis lucifugus emerges at late dusk to begin their foraging. The feeding activity follows much the same pattern as would be expected with the activity beginning to rise between 21:00 and 21:30, peaking between 22:00 and 22:30 and beginning to taper off between 22:30 and 23:00.

Eptesicus fuscus was found in significant numbers in the Pasture habitat with

activity beginning to start increasing between 20:30 and 21:00, peaking between 21:00

and 21:30, and starting to decrease between 21:30 and 22:00. This agrees with Belwood,

(1998) who claims that Eptesicus fuscus are the first bats to emerge each night from their roosts in order to forage. The feeding activity time line for this species looks much the same as the general activity time line with the highest amount of activity occurring between 21:00 and 21:30 and then quickly tapering off soon after this.

Lasionycteris noctivagans was found in significant numbers in old growth, Eco-

tone, and Pasture habitats. This species has its peak activity in the old growth habitat

between 20:30 and 21:00. This activity dies down between 21:00 and 21:30. It is at this

point that the activity peaks in both the Eco-tone and Pasture habitats. This peak dies

down in the Eco-tone habitat between 21:30 and 22:00 but only dies down in the Pasture

habitat between 22:00 and 22:30. Although there was no evidence of where this species

was actively foraging, the activity pattern of this species (in comparison to that of Myotis

lucifugus and Eptesicus fuscus) suggests that these species may be partitioning the

resources found in these three habitat types.

It is notable that Lasionycteris noctivagans only entered the Pasture habitat once the activity of Eptesicus fuscus had decreased dramatically (21:00 – 21:30) (figure 7, 8).

The Pasture activity of Lasionycteris noctivagans did not die down suddenly. This was unlike the Eco-tone habitat where the decrease in activity of Lasionycteris noctivagans coincides with the increase of Myotis lucifugus activity (21:30-22:00) (figure 6, 8). The activity and potential use of the old growth habitat by Lasionycteris noctivagans prior to its move in to the Eco-tone and Pasture habitats may be allowing these bats to avoid competition with other species and time their movement into the Pasture habitat so that competition from other species is minimized.

The increased activity in the Pasture land by Eptesicus fuscus and Lasionycteris noctivagans as the summer progresses may be as a result of certain insect species increasing their activity in the open habitats in the later summer months. The combined decrease in activity in the old growth and increase of activity in Pasture land by Eptesicus fuscus and Lasionycteris noctivagans may indicate these species are leaving the Old growth and spending more time in the Pasture land habitats as the summer progresses.

The decrease in activity of Lasiurus cinereus in the Pasture habitat as the summer progressed may be due to the increased competition from Eptesicus fuscus and

Lasionycteris noctivagans, it is not clear as to what habitat the species shift to later in the summer.

The existence of a gradient of habitat with regard to clutter enables the various bat species to develop foraging niches which will in turn reduce competition between the species. The high clutter of the Second growth habitat provides species such as Myotis septentrionalis an optimal area in which to hunt, a habitat that it has evolved to exploit through a specific echolocation and feeding technique. The medium clutter habitats of the

Old Growth and Eco-tone allow Myotis lucifugus, a medium clutter species, and a similar selective niche to forage and feed. While the open habitat of the Pasture type provides the low clutter species like Eptesicus fuscus a habitat that they can exploit and feed in. This clutter gradient is also represented in a temporal scale as well with certain species moving through the gradient and making use of different habitats at different times of night. The loss of this gradient would result in some of the species being forced in to the same niches and increasing direct competition between species. The presence of Dysart

Woods with the surrounding second growth forest and pasture lands provides the widest gradient for bat species in the local area.

These findings force me to reject my hypothesis that the regions of Old growth

forest would have higher bat diversity and more bat activity. It appears that the Old

growth forest is part of a clutter gradient that is, for the most, part used equally by

different species

Dysart woods integrity may be compromised due to the long wall mining that is

already underway beneath the forest. Long wall mining is a technique used to remove

coal from a seam by removing panels of the coal seam that are hundreds of feet wide and

thousands long. Long wall mining causes ground movement and strata displacement over

large areas approximately the size of the removed panels. These movements cause the

land surface to “sag” or “subside” which can cause damage to land structures. The

movements can also cause rock strata to fracture, bend, or shear which often alters the

pattern of ground water flow (Pennsylvania Environmental Protection Agency, 2005).

The alteration of ground water flow may alter the forest structure due to the fact that the

trees in this region most likely rely on ground water as part of their water budget during

the growing season (Burgess, 2006).The loss of these woods would result in a loss of

habitat gradient. Forest management and conservation practices should focus on creating

a heterogeneous habitat that contains a range of forest patches that are high in clutter,

have medium clutter, and are low in clutter. This will allow for spatial and temporal

separation of bat species while also providing habitat types for all species.

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www.msb.unm.edu/mammals/batcall/html/calllibrary.html

Appendix A- Tables and Figures

Figure 1: Varying areas of the forest that may be exploited by different species based on their echolocation type and amount of clutter. Dark blue label 1 represents open areas exploited by fast less agile species that utilize longer slender wings, pink label 2 and light blue label 3 represents less open areas that are exploited by slower but more agile species with wings that are broader and shorter. Red label 4 and yellow label 5 represent areas with the most clutter and are exploited by species that are the slowest but most agile having the broadest and shortest wings Habitats of varying clutter within the forest setting

Corben and O’Farrell, 1999

Dates, description, and coordinates of sampled sites Table 1: Site number with category of habitat type, brief site description, coordinates relevant to maps 2 and 3, and date the site was sampled

Date Site Habitat type Description Coordinates sampled

Old growth 1 forest Smaller canopy opening/ disturbance SW of K-13 07/26/2006 Old growth 2 forest Canopy opening/ disturbance G-4 05/24/2006 Old growth 3 forest Canopy opening/ disturbance I-5 16/08/06 Sparsely spaced old growth 4 Eco-tone becoming pasture E of F-3 07/07/2006 5 Pasture Field with few trees D-6 07/07/2006 6 Pasture Open field with no trees SE of J-6 08/16/2006 7 Second growth Canopy break/ disturbance S of road, N of I-10 05/08/2006 Thick under-story with no canopy 8 Second growth gaps E of F-10 07/26/2006 Thick under-story with no canopy 9 Second growth gaps I-6 07/07/2006

Bat species surveyed for at Dysart Woods Table 2: Species, flight style, common name, echolocation style, and basic biology (Belwood, 1998) of bat species that were surveyed for at Dysart Woods in 2006. Species Wing Flight style Abundance Echolocation Foraging (common type (foraging Characteristics name) habitat type, refer to fig 1) Eptesicus Long Faster less Common Narrow Foraging fuscus(Big slender agile flight bandwidth peak Brown Bat) wing during (Menzel (dark blue second et al., label 1) hour after 2005) sunset Lasionycteris Long Faster less Rarely Narrow Emerge noctivagans slender agile flight encountered bandwidth early, (Silver Haired wing hunt over Bat) (dark blue woodland label 1) ponds and along the edge of woodlots Lasiurus Long Faster less Rarely Narrow Emerge cinereus slender agile flight encountered bandwidth later in (Hoary Bat) wing the (Menzel (dark blue evening, et al., label 1) forage in 2005) the open Nyticeius Shorter Slower more Rarely Broader Emerge humeralis broader agile flight encountered bandwidth early, (Evening Bat) wings feed (Menzel (pink label 2 among et al., and light blue trees 2005) label 3) Pipistrellus Shorter Slower more Rarely Broader Among subflavus broader agile flight encountered bandwidth the first (Eastern wings to Pipistrelle) (Menzel (pink label 2 emerge, et al., and light blue feed at 2005) label 3) tree top level and forest edges Myotis Shorter Slower more Common Broader Emerge

38 lucifugus broader agile flight bandwidth at late (Little Brown wings dusk, Bat) (pink label 2 thought and light blue to hunt label 3) over water often following the same pursuit patterns Myotis Short Slow/hovering Uncommon Broad Emerge septentrionalis broad flight bandwidth just after (Northern wings dusk to Long-Eared (red label 4 glean Bat) and yellow prey off label 5) surfaces

Results from discriminant function analysis Table 3: Proportional loadings for the three pulse characteristics along three axis including the Eigenvalue, Wilks’ Lambda, and F value provided by a discriminate function analysis. Loadings Variable Axis 1 Axis 2 Axis 3 Highest freq 0.891 0.317 -0.322 Lowest freq 0.997 0.055 0.035 Duration -0.804 0.581 0.127 ______Eigenvalue 3.908 Wilks' Lambda 0.111

FNVM, DEN 335.5 p= 0.0001

Average number of pulses per call for each species Table 4: Total number of pulses, total number of calls, and average number of pulses per call for each species in each habitat type in which they were detected Habitat type Species Number of Total Average number calls number of of pulses per call pulses Second Myotis 5 44 8.8 Growth septentrionalis

Lasionycteris 1 5 5 noctivagans

Myotis lucifugus 10 82 8.2

Eptesicus fuscus 1 7 7

Old Growth Lasionycteris 19 162 8.5 noctivagans

Eptesicus fuscus 8 79 9.9

Myotis lucifugus 11 88 8

Eco-tone Eptesicus fuscus 1 38 38

Lasionycteris 23 205 8.9 noctivagans

Myotis lucifugus 84 1000 11.9

Pasture Lasionycteris 24 243 10.1 noctivagans

Lasiurus cinereus 4 39 9.75

Eptesicus fuscus 63 1097 17.4

Myotis lucifugus 19 375 19.7 .

General activity counts of species in habitat type Table 5: The counts for each species detected sorted into the habitat types in which they were detected and the percent of total calls found within the particular habitat type. The asterisks (*) indicates counts that were significantly high (Chi squared). Counts of unknowns are not represented by percentages Second Growth Old Growth Eco-tone Pasture (20) (41) (110) (134) % of % of % of count % of tot count tot count tot count tot Lasiurus 0 0 0 0 0 0 4 100 cinereus

Lasionycteris 1 1.5 19 28.4 23 34.3 24 35.8 noctivagans * * *

Eptesicus 1 1.4 8 9.7 1 1.4 63 87.5 fuscus *

Myotis 10 8.1 11 8.9 84 67.7 19 15.3 lucifugus *

Myotis 5 100 0 0 0 0 0 0 Septentrionalis

Unknown 3 - 3 - 1 - 24 -

Feeding activity counts of species in habitat type Table 6: Indicates the counts of feeding buzzes for the four species that were detected feeding as well as the percents of the feeding buzzes for each species found in each habitat type out of the total number of feeding buzzes detected for that species. The asterisks (*) indicates counts that were significantly high (Chi squared). Counts of unknowns are not represented by percentages Second Growth Old Growth Eco-tone Pasture (1) (7) (32) (40) count % of tot count % of tot count % of tot count % of tot Lasiurus 0 0 0 0 0 0 3 100 cinereus

Lasionycteris 0 0 0 0 0 0 4 100 noctivagans

Eptesicus 0 0 0 0 0 0 22 100 fuscus *

Myotis 0 0 5 13.2 28 73.6 5 13.2 lucifugus *

Unknown 1 - 2 - 4 - 6 -

Figure 2: Indicates the counts of Myotis lucifugus in all habitat types. Note the relatively high counts in the eco-tone habitat while also being found in all other habitat types to some degree.

Myotis lucifugus activity in all habitat types

90 85 80 75 70 65 60 55 50 45

count 40 35 30 25 20 15 10 5 0 Second Growth Old Growth Eco-tone Pasture habitat type

Figure 3: Indicates the counts of Eptesicus fuscus in all habitat types. Note the relatively high numbers in the pasture habitat.

Eptesicus fuscus activity in all habitats

count40

0 Second Growth Old Growth Eco-tone Pasture habitat type

Figure 4: Indicates the counts of Lasionycteris noctivagans in all habitat types. Note the relatively high numbers in Old Growth, Eco-tone, and Pasture habitats.

Lasionycteris noctivagans activity in all habitat types

70 65 60 55 50 45 40 35

count 30 25 20 15 10 5 0 Second Growth Old Growth Eco-tone Pasture habitat type

Figure 5: Indicates the feeding buzz counts for Myotis lucifugus in all habitat types. Note the generalist activity with significantly higher numbers being found in the Eco-tone habitat.

Myotis lucifugus feeding activity in all habitat types

10 feeding buzz counts feeding

0 Second Growth Old Growth Eco-tone Pasture

Figure 6: Indicates the feeding buzz counts for Eptesicus fuscus in all habitat types clearly showing the only site were it was detected to be feeding.

Eptesicus fuscus feeding activity in all habitat types

10 feeding buzz counts feeding

0 Second Growth Old Growth Eco-tone Pasture

Figure 7: General activity and feeding counts for Myotis lucifugus plotted over the half hour intervals that they occurred in the Eco-tone habitat.

General and feeding activity time line for Myotis lucifugus in the eco-tone habitat

30 feeding 25 general counts 20

15 10

0 20:00- 20:30- 21:00- 21:30- 22:00- 22:30- 23:00- 23:30- 20:30 21:00 21:30 22:00 22:30 23:00 23:30 24:00

Figure 8: General and feeding activity counts for Eptesicus fuscus plotted over the half hour intervals that they occurred in for the pasture land habitat.

General and feeding activity for Eptesicus fuscus in the pasture habitat

25 feeding 20 general counts

0 20:00- 20:30- 21:00- 21:30- 22:00- 22:30- 23:00- 23:30- 20:30 21:00 21:30 22:00 22:30 23:00 23:30 24:00

Figure 9: General count data for Lasionycteris noctivagans plotted in the half hour interval that they occurred in for Old Growth, Eco-tone, and Pasture habitats.

General activity time line for Lasionycteris noctivagans

50 45 40 35 Old Growth 30 25 Eco-tone 20 counts 15 Pasture 10 5 0

:30 :00 :30 :00 :30 :00 :30 :00 -20 -21 -21 -22 -22 -23 -23 -24 :00 :30 :00 :30 :00 :30 :00 :30 20 20 21 21 22 22 23 23

Presence and feeding counts of all species surveyed for over the entire study period Table 7: The detection and feeding presence of all species surveyed for all habitat types over the entire course of the study period. ● refers to a detection of an individual of a certain species, ◙ refers to a feeding detection of an individual of a certain species, - refers to no detection of presence or feeding, × refers to sites that were not sampled on that date due to technical difficulties, values included with individual detection and feeding detection symbol in parenthesis represent the count of that activity for that date. Species Date Habitat type Second growth Old growth Eco-tone Pasture Lasiurus 08/05/06 - × × × cinereus 24/05/06 × - × × 07/07/06 - × - (◙3) (●3) 26/07/06 - - × × 16/08/06 - - × (●1)

Lasionycteris 08/05/06 - × × × noctivagans 24/05/06 × (●10) × × 07/07/06 (●1) × (●23) (●9) 26/07/06 - (●8) × × 16/08/06 - (●1) × (◙4) (●15)

Eptesicus 08/05/06 - × × × fuscus 24/05/06 × (●7) × × 07/07/06 (●1) × (●1) (●1) 26/07/06 - (●1) × × 16/08/06 - - × (◙22) (●62)

Myotis 08/05/06 - × × × lucifugus 24/05/06 × - × × 07/07/06 (●10) × (◙29) (●84) (●1) 26/07/06 - (◙1) (●11) × × 16/08/06 - - × (◙5) (●18)

Myotis 08/05/06 (●2) × × × septentrionalis 24/05/06 × - × × 07/07/06 (●3) × - - 26/07/06 - - × × 16/08/06 - - × -

Nycticeius 08/05/06 - × × × humeralis 24/05/06 × - × × 07/07/06 - × - - 26/07/06 - - × × 16/08/06 - - × -

Pipistrellus 08/05/06 - × × × subflavus 24/05/06 × - × × 07/07/06 - × - - 26/07/06 - - × × 16/08/06 - - × -

Appendix B- Maps Entire study area Map 1: Map of old growth Dysart woods (in green loop) including second growth (area outside of green loop inside bold yellow border

Burgess, 2006

Northern section of old growth forest Map 2: Northern section of old growth forest. Including coordinate grid, foot path (red), access road (black), stream (blue), and sites where Anabat units were set up (◘)

Burgess, 2006

Site 4 ◘

◘ Site 2 Site 3 ◘

◘ Site 5 ◘ Site 9

◘ Site 6 ◘ ◘ Site 7

◘ Site 8

Southern section of old growth forest

Burgess, 2006

◘ Site 7

◘ Site 8

◘ Site 1

Map 3: Southern section of old growth forest. Including coordinate grid, foot path (purple), access road (black), stream (blue), and sites

where Anabat units were set up (◘)