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This is to certify that the
thesis entitled

Changes in the Distribution of Michigan’s Flying Squirrels

presented by
Robert Russell Skillen

has been accepted towards fulfillment
of the requirements for the

MS. i degree in

 

Zoology

fl! é E 6306? g: .
Major Professor’s Signature

2%A; 2001’:

Date

 

 

MSU is an Affirmative Action/Equal Opportunity Institution

LIBRARIES
IGAN STATE UNIVERSITY
ErgTIiI‘ANSING, MICH 48824-1048

. M
v.—'——--——'—- r -

v‘ ‘v .
-u ' .

PLACE IN RETURN BOX to remove this checkout from your record.
10 AVOID FINES return on or before date due.
MAY BE RECALLED with earlier due date if requested.

 

DATE DUE

DATE DUE

DATE DUE

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

2/05 cJClRC/DateDuejndd-p. 15

 

CHANGES IN THE DISTRIBUTION OF MICHIGAN’S FLYING SQUIRRELS
By

Robert Russell Skillen

A THESIS
Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of
MASTER OF SCIENCE
Department of Zoology

2005

ABSTRACT

CHANGES IN THE DISTRIBUTION OF MICHIGAN’S FLYING SQUIRRELS
By

Robert Russell Skillen

In light of growing concern for boreal mammalian Species at the southern edge of
their range, the current and historical ranges of flying squirrels (Glaucomys) in Michigan
were examined. The available data clearly indicate a dramatic northward advance of the
southern flying squirrel (G. volans) in the Lower Peninsula and an eastward advance in
the Upper Peninsula. While nonhem flying squirrels (G. sabrinus) persist in the eastern
Upper Peninsula, recent intensive trapping efforts indicate that populations may be
declining in other parts of the historical range, particularly the northern Lower Peninsula.
January isotherms coinciding with the northern range limit of the southern flying squirrel
proposed by Muul (1968) and Stapp et al. (1991) appear not be valid in Michigan.
However, changes in other climatic measures more indicative of winter severity and in
the distribution of conifer forests may be influencing the recent changes in the

distn'bution of flying squirrels in Michigan.

ACKNOWLEDGEMENTS

I would like to thank the members of my graduate advisory committee— Dr.
Barbara Lundrigan, Dr. Philip Myers, Dr. Kelly Millenbah, and Dr. Laura Smale— for
their support, understanding, and mentoring. I also thank Sean Maher and Michelle
Smith for field assistance, the Oscoda US. Forest Service Ranger Station personnel, the
staff of Seney National Wildlife Refuge, my beautiful wife Jennifer Skillen and
Genevieve Nesslage for GIS support and review, and the Michigan Animal Damage
Control Association for donating specimens. Funding for this project was provided by a
Michigan Department of Natural Resources 2002 Natural Heritage Nongame Wildlife
grant and the MSU Department of Zoology. Funding for a poster version of this thesis
was supported by the MSU Graduate School and presented at the 2004 meeting of the

American Society of Mammalogists.

iii

TABLE OF CONTENTS

LIST OF TABLES ................................................................................... vi
LIST OF FIGURES ................................................................................. vii
INTRODUCTION .................................................................................... 1
METHODS ............................................................................................ 6
Glaucomys Distribution ..................................................................... 6
Climate and Range ........................................................................... 8
Habitat Associations ....................................................................... 10
Michigan Land Cover Change ............................................................ 12
RESULTS ............................................................................................ l7
Glaucomys Distribution .................................................................... 17
Climate and Range ......................................................................... 17
Habitat Associations ....................................................................... 23
Michigan Land Cover Change ............................................................ 26
DISCUSSION ....................................................................................... 30
Glaucomys Distribution .................................................................... 30
Climate and Range .......................................................................... 31
Habitat Associations ....................................................................... 32
Michigan Land Cover Change ............................................................ 34
Recommendations .......................................................................... 36
LITERATURE CITED ............................................................................. 38
APPENDD< I
a. Georeferenced location data for Glaucomys captured in Michigan (North
American Datum 1927) ............................................................... 43
b. Original location information associated with each record from Appendix
Ia ......................................................................................... 60
c. Notes corresponding to each record from Appendix Ia ........................... 77
APPENDIX 11
List of US and Canadian natural history museums contacted and data
received ...................................................................................... 94
APPENDIX III

Sites trapped during summer 2002 field season. All localities are NAD27.........98

APPENDIX IV
Nest boxes surveyed for over-wintering Glaucomys from December 2002 to
February 2003. All locations are NAD27 ............................................. 100

Table 1:

Table 2:
Table 3:

Table 4:

Table 5:

Table 6:

Table 7:

LIST OF TABLES

List of literature sources and home range sizes for Glaucomys sabrinus

and Glaucomys volans ............................................................. ll
Reclassification of 1978 MIRIS land cover classes ........................... l3
Reclassification of 2001 IFMAP/GAP land cover classes ................... 16

2001 IFMAP/GAP land cover associated with buffers of Michigan
Glaucomys records from 1999-2003 with a maximum error 5 26
meters ............................................................................... 24

1992 Landsat Thematic Mapper National Land Cover associated with
buffers of Michigan Glaucomys records from 1990-1994 with a
maximum error 5 26 meters ...................................................... 25

Sixteen categories of land cover change from 1978 to 2001 (based on
reclassifications from Tables 2 & 3) ............................................. 27

Reclassification of 1978 MIRIS and 2001 IFMAP/GAP land cover
classes ............................................................................... 28

vi

Figure 1:

Figure 2:

Figure 3:

Figure 4:

Figure 5:

Figure 6:

LIST OF FIGURES

Decadal distribution of Glaucomys volans capture locations in Michigan
(N = 316). Pre-1954 triangular points represent three 1923 outliers; open
circle point represents two records from 1939 ................................ l8

Decadal distribution of Glaucomys sabrinus capture locations in
Michigan (N = 198) ............................................................... l9

Decadal distribution of both Glaucomys sabrinus ( O , N = 198) and
Glaucomys volans ( D , N =316) capture locations in Michigan ............ 20

Relative abundance index of G. sabrinus as a percent of total Glaucomys
captures north of 44° north latitude in five year intervals from 1878 to
2003 ................................................................................. 21

Trends of number of J anuarys per N 0AA Cooperative Weather Station
with January average daily temperature 5 -6.7°C (a), January minimum
daily temperature s -15°C (b), and the number of days per year s -15°C
(c). For all figures, ( + ) indicates a significant increase, ( - ) a

significant decrease, and ( O ) is not significantly different from zero

(p S 0.05). Although the number of years of data available for each
individual weather station vary, data overall range from January 1,

1900 to December 31, 2003 ....................................................... 22

Change in conifer land cover from 1978 to 2001 in Michigan. From left
to right, conifer land cover gain, loss, and unchanged. . . 29

vii

INTRODUCTION

Recent syntheses of numerous studies of wild plants and animals attribute shifts in
phenology (timing of events such as egg laying, flowering, or migration) and range
(toward the poles or higher elevations) to climate change (Parmesan and Yohe 2003;
Root et al. 2003). These studies examined over 1,500 Species; of those species showing
statistically Significant changes in either phenology or range, over 80% are shifting either
their phenology or range in a direction predicted by global warming.

In Michigan, Myers et al. (in press) have documented a northward shift in the
range of the white-footed mouse (Peromyscus leucopus), and a coincident decline in
captures of its northern congener, the woodland deer mouse (Peromyscus maniculatus
gracilis). Myers et al. (in press) suggest that climatic warming may be responsible for
the distributional changes in these species.

Based on recent and historical capture records, Myers and colleagues (personal
communication) have also speculated that another pair of closely related species with
overlapping ranges in Michigan, the southern flying squirrel (Glaucomys volans) and the
northern flying squirrel (Glaucomys sabrinus), are experiencing changes in their
distribution similar to the pattern documented for Peromyscus. The historical range maps
drawn for Glaucomys in Michigan by Burt (1969), Baker (1983), and Kurta (1995)
illustrate some of these changes. All three authors depict G. volans as ubiquitous
throughout the Lower Peninsula (LP). However, in the Upper Peninsula (UP), G. volans
has advanced eastward through time. Specifically, in Burt's (1969) map, they occur only
in the southernmost UP County, Menominee; in Baker's (1983) map, their range extends

through Menominee, Marquette and Houghton Counties; and in Kurta's (1995) map, G.

volans occurs through most of the mid-section of the UP, with the eastern boundary at
approximately the western border of Seney National Wildlife Refuge and the western
boundary at the western borders of Houghton and Iron Counties. These changes in the
distribution of G. volans in Michigan were summarized by Wells-Gosling (1982), who
noted that the records available at the time “may indicate a northward range extension for
this species [G. volans] in the Upper Peninsula”. However, Wells-Gosling (1982)
believed this difficult to substantiate at the time due to the limited representation of
Glaucomys in museum mammal collections.

Burt (1969), Baker (1983), and Kurta (1995) illustrate the range of G. sabrinus to
consistently include all of the UP and the northern two-thirds of the LP (excluding the
“thumb” region). However, the low number of G. sabrinus captures from the LP and the
western half of the UP in the 1980s and 1990s led Myers and colleagues (personal
communication) to believe that a more detailed examination of the current and historical
ranges of Glaucomys in Michigan was warranted.

Although I know of no studies that examine changes in the distribution of
Glaucomys in Michigan beyond those mentioned here, changes in the distribution of
Glaucomys have been documented in neighboring regions, including Pennsylvania
(Mahan et al. 1999), Quebec (Oxley and Gall 1977; Youngman and Gill 1968), and
Ontario (J. Bowman, Ontario Ministry of Natural Resources, personal communication).
Mahan et al. (1999) have documented a decline in G. sabrinus and Oxley and Gall
(1977), Youngman and Gill (1968), and J. Bowman (Ontario Ministry of Natural
Resources, personal communication) have noted northward range expansions of G.

volans. Four factors that may influence changes in the distribution of Glaucomys have

been proposed by these and other authors: habitat fragmentation or destruction (Mahan et
al. 1999), parasite transmission (Wetzel and Weigl 1994), competition between G.
sabrinus and G. volans (W eigl 1978), and climate change (Muul 1968; Stapp et al. 1991).

Mahan et al. (1999) focused on habitat change as a possible cause for the decline
in G. sabrinus captures in Pennsylvania. Studies have shown G. sabrinus to prefer mature
conifer forests and occasionally mature mixed and pure stands of deciduous forest, all
within fairly close proximity to water (Carey et al. 1999; Cotton and Parker 2000;
Harestad 1990; Payne et al. 1989; Ransome and Sullivan 1997; Weigl 1978; Wells-
Gosling and Heaney 1984). Mahan et al. (1999) have documented fragmentation and
loss of old growth forests associated with the decline in G. sabrinus captures.

' Wetzel and Weigl (1994) suggest that G. sabrinus populations may be negatively
affected by the transmission of the cold intolerant parasite Strongyloides robustus from
the more resistant G. volans in areas of range overlap where the two species may come
into close contact. The recent discovery that both species of Glaucomys occasionally
share the same dens during the winter months (Lavers 2004) may be an important factor
when considering parasite transmission, although den sharing by both species of
Glaucomys has not been documented in Michigan.

A study by Weigl (1978) indicates that G. volans may out-compete G. sabrinus.
Specifically, G. volans was found to be more aggressive in defending its home area and
typically controlled nests when paired in laboratory behavioral trials with G. sabrinus.
Weigl (1978) speculated that by breeding earlier in the year, G. volans may occupy a
limited supply of nest cavities in the wild first. However, it is not clear that a limited

supply of nest cavities negatively impacts G. volans (Brady et al. 2000) or G. sabrinus

(Carey 2002) population densities. Both Brady et al. (2000) and Carey (2002) found that
supplementary nest boxes did not result in a Si gnificant increase in population densities of
either Glaucomys species as compared to control populations.

Additionally, Muul (1968), and more recently Stapp et al. (1991), have proposed
the existence of thermal barriers that may impose physiological limitations on the ability
of G. volans to occupy northern regions of the United States and southern Canada.
Specifically, Muul (1968) proposed that the northern range limit of G. volans “agrees
fairly well with the 20°F (-6.7°C) isotherm of average January temperature” and Stapp et
al. (1991) that “the northern boundary nearly coincided with the -15°C isotherm of
January minimum daily temperature”. Stapp et al. (1991) argue that at -15°C, G. volans
nesting either in aggregations (multiple squirrels nesting together) or singly in cavity
nests require 18.82 kcal, or 2.5 x Basal Metabolic Rate (BMR), daily. They speculate
that at this extreme maximum of daily energy expenditure, G. volans may not be able to
store enough food to survive the winter. This relationship of the northern range boundary
coinciding with 2.5 x BMR also holds true for several bird species (Root 1988). Stapp et
al. (1991) admit that it is not clear why this relationship exists and that it could be “purely
coincidental”. However, given the paucity of information on the physiological
limitations of G. volans, this provides a useful starting point for examining the
relationship between climatic variables and the geographical range limits of this squirrel.
If such thermal barriers exist and the range of G. volans has shifted northward in
Michigan, there should be evidence that these thermal barriers have shifted northward in

Michigan as well.

The objectives of this study were to 1) describe distributional changes in G.
sabrinus and G. volans in Michigan, USA. over the last ca. 100 years, 2) determine
whether changes in biologically important climatic variables coincide with changes in
Glaucomys distributions, 3) describe the habitats associated with Glaucomys capture
locations and 4) determine whether changes in critical habitat coincide with changes in

Glaucomys distributions.

METHODS

Glaucomys Distribution

To evaluate the distributional changes of Glaucomys in Michigan, collecting data
(i.e., species, date, and locality; Appendix I) were solicited from North American natural
history museum mammal collections (Appendix 11). These data were supplemented by
live-trapping records from Dr. Philip Myers (University of Michigan) and his students,
mist netting records from Dr. Allen Kurta (Eastern Michigan University), and Specimens
collected by animal damage control workers and subsequently donated to the Michigan
State University Museum (Appendix 1). Although a variety of biases inevitably exist in
the methods of and reasons for collection of individual specimens, these records are the
only sources of information available. Therefore, it is assumed that, in the aggregate,
these records are a reasonable representation of the past and present distribution of
Glaucomys in Michigan.

Trapping (AUF # 03/02-04400, MDN R Scientific Collectors Permit # SC1169)
was conducted specifically for this study during May-August 2002 in the northern LP
(defined to be north of 44° north latitude, excluding the “thumb” region) and central UP
(in and around Seney National Wildlife Refuge). These regions were believed to include
the zone of range overlap between G. volans and G. sabrinus. Individual trapping sites
within these regions were chosen based on the availability of appropriate habitat (mature
conifer, deciduous, or mixed forests) on public lands (Appendix III). Trap lines consisted
of 40 traps, alternating LFA (3 x 3.5 x 9") with XLF-lS (4 x 4.5 x 15") ShermanTM live
traps; traps were set approximately 8 meters apart. All trap lines were set for at least two

consecutive nights (up to five consecutive nights in some areas) at all locations and total

trapping effort was 72 trap lines and 8440 total trap nights (Appendix HI). However,
1772 traps were found shut without captures for an effective total of 6668 trap nights.
Traps were placed in lines instead of a grid pattern to maximize the number of flying
squirrel territories that could be sampled per unit effort. All traps were baited with oats
and sunflower seeds. In addition to trapping, nest boxes erected mainly for wood ducks
by the US. Forest Service in Alcona County were surveyed (under the same permits as
trapping) from December 2002 to February 2003 for over-wintering flying squirrels
(Appendix IV).

The localities of all Glaucomys captures from trapping and nest box surveys were
determined using a Garmin eTrexTM handheld Global Positioning System (GPS) unit set
to North American Datum 1927 (NAD27). Trapping data from other studies and locality
data from all specimens were georeferenced using protocols defined by the Mammal
Network Information System (MaN IS; http://dlp.cs.berkeley.edu/manis/) using NAD27.
Records were imported into ArcView GIS 3.2 (Environmental Systems Research Institute
1992-1995) and transformed to the Michigan Georef coordinate system for analysis.

Rather than reporting absolute numbers of squirrels captured, the relative
abundance of G. sabrinus and G. volans was documented as the ratio (#G. sabrinus)/(#G.
sabrinus+G. volans) in five year intervals from 1878 to 2003. Absolute numbers were
not used because the trapping effort (number of trap-nights) of surveys was often
unknown and inconsistent from survey to survey. It should be noted that changes in
relative abundance over time will be misleading if methods used in earlier surveys (e.g.,

type of trap, bait, habitats trapped) differed from those used recently and those methods

were biased towards one of the species. However, there is no evidence to suggest that

this is the case.

Climate and Range

Daily temperature data (datasets 3200, 3206, and 3210) covering the period from
January 1, 1900 to December 31, 2003, and Cooperative Weather Station localities were i
downloaded from the National Oceanographic and Atmospheric Administration (NOAA)
National Climate Data Center websites (ftp://ftp.ncdc.noaa.gov/publdata/inventories/
COOP.TXT, http://www4.ncdc.noaa.gov/ cgi—win/wwcgi.dll?wwDI~GetCity~USA) for
the entire state of Michigan. These data were used to calculate three temperature
measures per year per Cooperative Weather Station: the January average of daily

minimum temperature, the January average of average daily temperature, and the number

of days per year at or below the daily minimum of -15°C.

NOAA was unable to provide accurate information as to the datum used to
georeference each of their Cooperative Weather Stations. However, they indicated that
the majority (88.6%) of their station locations nationwide are NAD83 (J. Arnfield,
NOAA, personal communication). Therefore, NAD83 was assumed for all Michigan
weather station locations. The data were transformed to the Michigan Georef coordinate
system and, as each station was listed by county, each location was checked to determine
if it was actually located within the county listed. As all stations were accurate to at least
the county level, the potential error due to inaccuracies in location data were considered

negligible as trends in climate were analyzed at the statewide level for this study.

In addition to datum inaccuracies, multiple locations were occasionally identified
with the same Cooperative Weather Station number. Cooperative Weather Stations were
assigned the same number only if they were within 2 horizontal miles or 100 vertical feet
of one another (http:l/lwf.ncdc.noaa.gov/oa/climate/stationlocator.html). Since the most
recent location data were accurate to seconds (older location data were accurate to
minutes), only the most recent location data for any particular NOAA Cooperative
Weather Station were used for these analyses. J anuarys with fewer than thirty-one days

of data and years with an incomplete record of the number of days minimum temperature

was 5 -150C were excluded from these analyses. Known errors, as cited in NCAA
metadata (http://www4.ncdc.noaa.gov/ol/documentlibrary/datasets.html#'I‘D9767B) and
errors discovered during data processing (mainly duplicate records) were corrected before
analysis.

The January average daily temperature isotherm of —6.7°C and the January
minimum daily temperature isotherm of -15°C were evaluated for change over time using
logistic regressions for each weather station. The independent variable in each regression

was time (year), and the dependent variable was either the number of J anuarys where the
January average daily temperature was 5 -6.7°C or the number of Januarys where the

January minimum daily temperature was S —15°C. The slopes of these regressions were
assessed to determine if temperature was significantly increasing over time (negative
slope), decreasing over time (positive slope), or not significantly different from zero (pg

0.05) for each weather station. Simple linear regressions were run with time (year) as the

independent variable and the number of days per year that were 5 —15°C as the

dependent variable. These regression slopes were assessed using the same methods as

those described for January temperatures. The number of J anuarys/years of data per
station varied; stations with fewer than six J anuarys/years of data were dropped from the
regression analyses. Program “R” (Ihaka and Gentleman 1996) was used for all

regressions.

Habitat Associations

Classified land cover data for the entire state of Michigan from the 2001
IFMAP/GAP Landsat Thematic Mapper imagery (Michigan Department of Natural
Resources 2003) were obtained from the Michigan Department of Natural Resources
(MDN R) and the 1992 National Land Cover Landsat Thematic Mapper data (V ogelmann
et al. 2001) were downloaded from the USGS National Land Cover Characterization web
site (http://1andcover.usgs.gov/natllandcover.aspl). The 1992 data were re-projected from
Albers equal area conic (NAD83) to the Michigan Georeference coordinate system. All
of the land cover data used are raster files with 30x30 meter pixels. The time periods of
data collection varied; the 1992 land cover data were collected between 1989 and 1995,
and the 2001 land cover between 1997 and 2001.

To determine habitat associated with Glaucomys capture locations, a literature
review was conducted to estimate a typical home range size for both G. sabrinus and G.
volans (Table l). A buffer with a radius of 160 meters (or a circle with an area of 8.002
ha) was chosen as a reasonable home range size for both species as this value falls within
range of most of the home ranges calculated by the studies listed in Table 1. Each 160

meter buffer was centered on each georeferenced specimen location. Only specimen

10

Table 1. List of literature sources and home range sizes for Glaucomys sabrinus and
Glaucomys volans.

 

 

Species Range (ha) Mean (ha) Location Source

G. sabrinus 4.9-12.6 8.7 Pennsylvania Weigl & Osgood 1974

G. sabrinus 4.9-7.1 5.9 North Carolina Weigl & Osgood 1974

G. sabrinus 6-31 - Alaska Mowrey & Zasada 1982
G. sabrinus 3.4-4.9 - Oregon Witt 1992

G. sabn'nus 5.1-6.7 - Oregon Martin & Anthony 1999
G. volans 1.6-2.0 - Missouri Schwartz &Schwartz 1959
G. volans 0.53 - New York Madden 1974

G. volans 1.89-3.49 - Maryland Gilmore & Gates 1985

G. volans 2.45 2.45 ($0.88 SE) Maryland Bendel & Gates 1987

G. volans 11.3-24.8 16 ($6.0 SD) New Hampshire Fridell & Litvaitis 1991

G. volans 4.0-16.0 9.0 Q25 SE) Arkansas Stone et al. 1997

G. volans 2.76-44 - Arkansas Taulman and Seaman 2000

 

ll

localities that were recorded with a maximum error _<_ 26.0 meters were analyzed
(calculated by MaN IS protocols; see Appendix Ia). Additionally, only Glaucomys
capture locations that were recorded within two years of a particular land cover year were
used, i.e. Glaucomys captures from 1990-1994 were associated with the 1992 land cover,
and captures from 1999-2003 with the 2001 land cover. Although land cover data are
available statewide for 1978, none of the Glaucomys capture location data from 1976-
1980 met the maximum error criteria set above as none were recorded using GPS.

The raster land cover data were converted to vector and the buffered capture
locations were used to clip habitat data from the associated land cover. Clipped habitat
data were used to calculate the percentages of each habitat type associated with each

species of Glaucomys.

Michigan Land Cover Change

The 2001 land cover and the 1978 Michigan Resource Information System
(Michigan Department of Natural Resources 1999; collected between 1978 and 1979)
land cover were used to determine change in conifer land cover statewide from 1978 to
2001. Original land cover classes were reclassified to conifer, deciduous, mixed forest,
or other (see Table 2 for 1978 and Table 3 for 2001). To assess land cover change over
time, the map calculator function of ArcView GIS 3.2 was used to add the 1978 land
cover reclassification to the 2001 land cover reclassification. For example, when a pixel
reclassified as conifer from the 1978 land cover was added to the reclassified 2001 land
cover, four outcomes were possible: unchanged (conifer to conifer), conifer to deciduous,

conifer to mixed, or conifer to other.

12

Table 2. Reclassification of 1978 MIRIS land cover classes.

 

 

Grid code ha Label Reclassification
0 2059 Other
1 100 6 Residential Other
1110 496 Multi-Family-Medium to High Rise Other
1120 12029 Multi-Family-Low Rise Other
1 131 486328 Single Family Duplex Other
1132 97254 Single Family Duplex Low Density Other
1150 8886 Mobile Home Park Other
1200 6113 Commercial Services Institutional Other
1210 6477 Central Business District Other
1220 3325 Shopping Center Mall Other
1230 121 Strip Commercial Other
1240 31567 Neighborhood Business Other
1260 41421 Institutional Other
1 300 45843 Industrial Other
1380 8 181 Industrial Park Other
1400 949 Transportation Communication Utilities Other
1410 16242 Air Transportation Other
1420 1 89 1 Rail Transportation Other
1430 528 Water Transportation Other
1440 35084 Road Transportation Other
1450 787 Communication Facilities Other
1460 18062 Utilities Waste Disposal Other
1 700 479 Extractive Other
1711 5155 Metallic Mineral Quarry Other
1712 245 Nonmetallic Mineral Quarry Other
1713 40059 Open Pit Other
1714 8 Other Open Pit Other
1715 3273 Sand and Gravel Other
1721 378 Metallic Other
1722 5829 Underground Extractive Other
1731 8 Extractive Wells Salt Other
1732 6 Oil Wells Other
1733 6 Waste Disposal Other
1734 2197 Wells Other
1900 3312 Open Land and Other Other
1930 52515 Outdoor Recreation Other
1940 8305 Cemeteries Other
2100 4141331 Cropland Other
2200 103306 Orchards Vineyards Other
2300 5465 Confined Feeding Other
2400 144342 Permanent Pasture Other
2900 9173 Other Agriculture Other
3100 669164 Herbaceous Openland Other

 

l3

Table 2 (cont’d).

 

3200 53991 1 Shrubland Other
3300 487 Pine or Oak Opening (Savanna) Other
4000 59 Forested Land Mixed
4100 27491 Broadleaved Forest (Generally Deciduous) Deciduous
41 1 1 2 Basswood Deciduous
41 12 126 Beech Deciduous
41 1 3 89 Cherry Deciduous
41 14 55 Elm Deciduous
4115 1904234 Northern Hardwood Deciduous
4116 2158 Red Maple Deciduous
4117 21782 Sugar Maple Deciduous
4118 209003 Undifferentiated Northern Hardwood Deciduous
4119 1 White Ash ‘ Deciduous
4121 ' 165 Black Oak Deciduous
4122 790176 Central Hardwood Deciduous
4123 106068 Red Oak Deciduous
4124 56452 Undifferentiated Oak/Hickory Deciduous
4125 58 White Oak Deciduous
4131 1156900 Aspe Birch Deciduous
4132 3245 Bigtooth Aspen Deciduous
4133 214 Trembling Aspen Deciduous
4134 270658 Undifferentiated Aspen/White Birch Deciduous
4135 1277 White Birch Deciduous
4141 32 Ash Deciduous
4142 67223 Aspen Deciduous
4143 23 Balm-of-Gilead Deciduous
4144 3 Cottonwood Deciduous
4145 42 Elm Deciduous
4146 897545 Lowland Hardwood Deciduous
4147 13781 Soft Maple Deciduous
4148 27561 Undifferentiated Lowland Hardwood Deciduous
4149 138 White Birch Deciduous
4200 36 Coniferous Forest Conifer
421 1 134100 Jack Pine Conifer
4212 637323 Pine Conifer
4213 69287 Red Pine Conifer
4214 4458 Scotch Pine Conifer
4215 3353 Undifferentiated Pine Conifer
4216 8646 White Pine Conifer
4221 137 Balsam Fir Conifer
4222 16 Black Spruce Conifer
4223 19 Hemlock Conifer
4224 165582 Other Upland Conifer Conifer
4225 3058 Undifferentiated Upland Conifer Conifer
4226 1279 White Spruce Conifer

 

14

Table 2 (cont’d).

 

4231 454 Balsam Fir Conifer
4232 290 Balsam Fir/White Spruce Conifer
4233 1209 Black Spruce Conifer
4234 21654 Cedar Conifer
4235 7 Jack Pine Conifer
4236 627716 Lowland Conifer Conifer
4237 1022 Tamarack Conifer
4238 84441 Undifferentiated Lowland Conifer Conifer
4291 15 Blue Spruce Conifer
4292 27430 Christmas Tree Plantation Other
4293 156 Scotch Pine Conifer
4294 437 Undifferentiated Christmas Tree Plantation Other
4295 5 White Pine Conifer
5 100 20292 River Other
5200 292016 Lake Pond Other
5300 29568 Reservoir Other
5400 267 Great Lakes Other
6000 1770 Wetlands Other
61 10 104512 Wooded Wetland Mixed
6120 479102 Shrub/Scrub Wetland Other
6200 9 Nonforested Wetlands Other
6210 24519 Aquatic Bed Wetland Other
6220 168115 Emergent Wetland Other
6230 1891 Flats Other
6240 7 Deep Marshes Other
7000 269 Barren Other
7200 4547 Beach Riverbank Other
7300 7601 Sand Dune Other
7400 8023 Bare Exposed Rocks Other

 

15

Table 3. Reclassification of 2001 lFMAP/GAP land cover classes.

 

 

Code ha Class (Label) Reclassification
1 219353 Low Intensity Urban Other

2 140935 High Intensity Urban Other

3 3707 Airports Other

4 425223 Road/Parking Lot Other

5 23824 Non-Vegetated Farmland Other

6 1612563 Row Crops Other

7 2132315 Forage Crops/Non-Tilled Herb. Ag. Other

9 73543 Orchards/VineyardS/Nursery Other

10 1106399 Herbaceous Openland Other

12 393140 Upland Shrub/Low Density Trees Other

13 31510 Parks/Golf Courses Other

14 1737133 Northern Hardwood Assoc. Deciduous
15 627457 Oak Association Deciduous
16 1031460 Aspen Association Deciduous
17 16450 Other Upland Deciduous Deciduous
18 431669 Mixed Upland Deciduous Deciduous
19 837983 Pines Conifer
20 150033 Other Upland Conifers Conifer

21 82056 Mixed Upland Conifers Conifer
22 785004 Upland Mixed Forest Mixed

23 591817 Water Other

24 733021 Lowland Deciduous Forest Deciduous
25 934763 Lowland Coniferous Forest Conifer
26 52492 Lowland Mixed Forest Mixed

27 45613 Floating Aquatic Other

28 628714 Lowland Shrub Other

29 1 10584 Emergent Wetland Other

30 313352 Mixed Non-Forest Wetland Other

31 54385 Sand, Soil Other

32 2627 Exposed Rock Other

33 14 Mud Flats Other

35 20718 Other Bare/Sparsely Vegetated Other

 

16

RESULTS

Glaucomys Distribution

The combination of efforts to acquire current and historical Glaucomys capture
records for Michigan resulted in 198 records for G. sabrinus and 316 records for G.
volans. These distributional data are presented by decade for G. volans (Figure 1) and G.
sabn'nus (Figure 2). It is clear that the range of G. volans has advanced northward in the
LP and eastward in the UP (Figures 1 and 3). The relative abundance of G. sabrinus in
the northern LP and the UP decreased from a range of 84-100% for the time intervals

before 1984 to <44% after 1984 (Figure 4).

Climate and Range

The regressions of Michigan January temperatures against time (N = 7
significant, N = 164 not significant for January average daily temperature, Figure 5a; and
N = 2 significant and N = 174 not Significant for January minimum daily temperature,
Figure 5b) indicate that the trend for the majority of stations is not different from zero.

This suggests that neither the January average daily temperature isotherm of —6.7°C nor

the January minimum daily temperature isotherm of —15°C have changed in Michigan in

the last ca. 100 years. Regressions of number of days per year per weather station that

were 5 —150C against time (N = 178, Figure 5c) indicate that winters are becoming less

severe in some areas of the LP and UP where G. volans has recently expanded its range

(Figure 1).

17

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21

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22

Habitat Associations

The 2001 land cover classes associated with the buffered capture location
records for G. volans from 1999-2003 (N = 39) are presented in Table 4. The five
deciduous classes (northern hardwood, aspen, and oak associations; lowland and mixed
upland deciduous forests) cover 56.2% of the 39 combined buffers for G. volans. The
four coniferous classes (pines, lowland conifer, mixed upland conifer, and other upland
conifer) cover 18.1%, and the two mixed forest classes (upland and lowland) cover 8.7%.
The 1992 land cover classes associated with the buffered capture location records for G.
volans from 1990-1994 (N = 6) resulted in 85.8% deciduous forest, 8.1% mixed forest,
and 3.2% conifer forest cover (Table 5).

The 2001 land cover classes associated with the buffered capture location
records for G. sabrinus from 1999-2003 (N = 13) are presented in Table 4. If the conifer,
deciduous, and mixed forest classes are grouped as for G. volans (described above),
deciduous forest classes cover 38.4%, conifer forest 22.8%, and mixed forest 10.0% of
the thirteen combined buffers for G. sabrinus. The 1992 land cover classes associated
with the buffered capture location records for G. sabrinus from 1990—1994 (N = 3)
resulted in woody wetlands covering 37.2%, mixed forest 29.7%, conifer forest 22.0%,

and deciduous forest 9.7% (Table 5).

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25

Michigan Land Cover Change

Conifer land cover changes in Michigan were evaluated using the
reclassifications of the 1978 and 2001 land covers (in Tables 2 and 3 respectively). These
reclassifications were added creating sixteen unique categories that describe land cover
change (T able 6). Conifer forests increased from 11.7% of the total land cover in 1978 to
13.1% of the total land cover in 2001 (Table 7). Approximately 51% of the conifer land
cover that existed in 1978 remained in the same locations throughout Michigan in 2001
(Table 6 and Figure 6). Deciduous forest land cover decreased from 36.9% of the total
land cover in 1978 to 29.8% of the total land cover in 2001 (Table 7). However,
approximately 69.3 % of the deciduous land cover that existed in 1978 remained in the

same locations in 2001 (Table 6).

26

Table 6. Sixteen categories of land cover change from 1978 to 2001 (based on
reclassifications from Tables 2 & 3).

 

 

Unchanged from Changed from Changed to ha % Unchanged from
1978 to 2001 (1978) (2001) 1978 to 2001

Conifer - - 1005418 51 .0
- Other Conifer 333012 -
- Deciduous Conifer 633953 -
- Mixed Conifer 4 -
Total Conifer - - 1972387 -
Deciduous - - 1956593 69.3
- Other Deciduous 7 15 129 -
- Conifer Deciduous 153571 -
- Mixed Deciduous 20 -
Total Deciduous - - 28253 1 3 -
Mixed - - 2 0.0
- Other Mixed 158291 -
- Conifer Mixed 174434 -
- Deciduous Mixed 496210 -
Total Mixed - - 828937 —
Other - - 6508096 69.5
- Conifer Other 412664 -
- Deciduous Other 2439179 -
- Mixed Other 33 -
Total Other - - 9359972 -

 

27

Table 7. Reclassification of 1978 MIRIS and 2001 IFMAP/GAP land cover classes.

 

 

Reclassification % of Total
1978 2001
Conifer 1 1.7 13.1
Deciduous 36.9 29.8
Mixed 0.7 5.5
Other 50.7 51.7

 

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29

DISCUSSION

Glaucomys Distribution

Changes in the distribution of Glaucomys have been documented in Pennsylvania
(Mahan et al. 1999), Quebec (Oxley and Gall 1977; Youngman and Gill 1968), and
Ontario (1. Bowman, Ontario Ministry of Natural Resources, personal communication).
In Michigan, there is convincing evidence that G. volans has advanced northward in the
LP and eastward in the UP over the last ca. 100 years (Figure .1). Of special note are
three G. volans that were captured in the northern LP near the city of Elmira (Otsego
County) in 1923 (#469-471; Appendix land marked by triangles in Figure 1). Although
the identification of these three specimens has been verified by the respective museum
staff, these records represent significant outliers as G. volans was not captured again in
the northern LP until 1986 (see record #337, Appendix I). Nineteen G. sabrinus were
captured between 1923 and 1986 in various localities throughout the same region. From
these data it seems unlikely that G. volans populations were well established in the
northern LP much before 1986.

The available Glaucomys distribution data suggest that in regions where G. volans
is advancing, G. sabrinus is retreating. Intensive trapping has recently been conducted in
areas where G. sabrinus were previously known to occur. However, these efforts have
predominantly resulted in G. volans captures. As an example, only three G. sabrinus
captures have been recorded for the northern LP within the last five years. For the same
time period and region, forty-one G. volans were captured. Similarly, trapping conducted

during the summer of 2003 by P. Myers and B. Lundrigan at the Huron Mountain Club

30

(Marquette County), an area where all previous captures recorded were G. sabrinus,
resulted in nineteen G. volans captures and only one G. sabrinus. Although trapping
efforts for my study were focused on habitats believed to be appropriate for G. sabrinus,
it is unknown whether other trapping efforts had habitat biases. The lack of G. sabrinus
captures is never a sure indicator that the species is not present. However, it is reasonable
to assume that capture of both Glaucomys species is equally likely using the trapping
methods typically employed by researchers in the field (e.g., ShermanTM live traps).
Therefore, the available data indicate that G. sabrinus populations are declining in
Michigan.

Due to the geographic isolation of Michigan in all directions except the south in
the LP and the west in UP, the isolated pockets of G. sabrinus that remain in the state
should be considered “island” populations, as no northern refugia are available. The
geographic isolation of Michigan’s G. sabrinus populations curtails gene flow between
these populations and the larger metapopulation, which depending on population
densities, could in time lead to inbreeding depression. Significant genetic differences
have been found between the populations of G. sabrinus in the LP and UP (Arbogast
1999), and the loss of these populations could mean a significant loss of genetic diversity

for the species as a whole.

Climate and Range
It has been speculated that winter temperature extremes prevent G. volans from
occupying habitats north of its present distribution (Muul 1968; Stapp et al. 1991). No

correspondence was found between changes in the distribution of G. volans in Michigan

31

(Figure l) and changes in the January isotherms described by Muul (1968) or Stapp et al.
(1991) (Figures 5a and 5b respectively). It should be noted that both Muul (1968) and
Stapp et al. (1991) used different and more limited sources of climate data than did this
study. Muul (1968) used 1941 Michigan weather maps to describe isotherms (US. Dept.
of Agriculture Yearbook 1941), and Stapp et al. (1991) used national weather maps from
1970 for the US. and 1974 for Canada (Canada Dept. of Energy Mines and Resources
1974; United States Geological Survey 1970). The climate data for this study ranged
from January 1, 1900 to December 31, 2003.

January temperatures alone may not be a good indicator of winter severity, and
may therefore be less biologically meaningful to G. volans than temperature variables
that incorporate more information. For this reason, the number of days 5 -15°C per year
per weather station were also evaluated for significant trends. This analysis revealed a
correspondence between the number of days _<_ ~15°C per year per weather station (Figure
5c) and changes in the distribution of G. volans over time (Figure 1). Overall, there are
more weather stations with significant warming trends than cooling trends and stations
with these significant warming trends are located in those areas of the northern LP and

UP where G. volans has recently expanded its range.

Habitat Associations
Although the 1992 land cover data are classified into fewer land cover types
(and are therefore coarser) than the 2001 land cover data, both are 30x30 meter

resolution. By aggregating the classes of both land covers into the broad categories of

32

conifer, deciduous, and mixed forest, general comparisons between the two datasets are
possible.

Glaucomys volans is thought to be associated with mature deciduous and
mixed forests, while avoiding pure stands of conifer (Dolan and Carter 1977; Gilmore
and Gates 1985; Ivan and Swihart 2000; Sonenshine and Levy 1981; Taulman and
Seaman 2000; Weigl 1978). Habitat associated with G. volans capture locations (T ables
4 and 5) fits well with the associations described in the literature. Glaucomys volans was
highly associated with deciduous forests (56.2% for the 2001~land cover and 85.8% for
the 1992 land cover). Mixed forests were expected to represent a larger proportion of the
habitat association than conifer, which held true for the 1992 land cover (8.1% vs. 3.2%
respectively), but not for the 2001 land cover (8.7% vs. 18.1% respectively).

G. sabrinus is thought to be associated primarily with mature conifer forests
and occasionally with mature mixed and pure stands of deciduous forest, all within fairly
close proximity to water (Carey et al. 1999; Cotton and Parker 2000; Harestad 1990;
Payne et al. 1989; Ransome and Sullivan 1997; Weigl 1978; Wells-Gosling and Heaney
1984). In somewhat of a departure from the literature, the G. sabrinus records associated
with the 2001 land cover (T able 4) revealed that the total conifer forest (22.8%)
represented a lower percentage than the total deciduous forest (38.4%), while the total
mixed forest at 10% is comparable to the 8.7% calculated for G. volans. The G. sabrinus
records associated with the 1992 land cover (T able 5) supported the association of G.
sabrr'nus with wet habitats, i.e. a relatively high percentage (37.2%) of the records were
associated with woody wetlands, as compared to the less than one tenth of one percent for

G. volans. However, this result was somewhat surprising as the metadata for the 1992

33

land cover describes the woody wetlands class as “Areas where forest or Shrubland
vegetation accounts for 25-100 percent of the cover and the soil or substrate is
periodically saturated with or covered with water” (httpzlllandcover.usgs.gov/
natllandcover.aspl). It is highly unlikely that G. sabrinus would be associated with
shrublands as forests are quintessential to their gliding lifestyle (W ells-Gosling and
Heaney 1984). Additionally, mixed forests (29.7%) were a stronger component than
conifer forests (22.0%) for the G. sabrinus records associated with the 1992 land cover.
However, these results may be skewed by the low sample size (N = 3).

Based on the habitat requirements described for G. sabrinus in the literature,
conifer forests were expected to represent a higher percentage of the combined buffers
for each capture location. One possible explanation for this disparity is that placing
perfectly circular buffers around Glaucomys capture locations does not accurately
represent the actual home range and habitat used by an individual squirrel. Another
possibility is that habitat preferences differ geographically. Genetic data indicate
significant genetic differences between eastern and western subspecies of G. sabrinus
(Arbogast 1999). Additionally, much of the research on the habitat associations of G.
sabrinus has been conducted in the Pacific Northwest and may not adequately describe

habitat associations for the species as a whole.

Michigan Land Cover Change
Mature conifer forests have been described as important habitat for G. sabrinus in
the literature; changes in land cover between 1978 and 2001 were thus examined to

determine if land cover has changed in Michigan in ways that are either beneficial or

34

detrimental to the species. Although statewide 1992 land cover data were available, these
data were not used to assess changes in land cover. The l9-year time span between the
data collection periods for the 1978 and 2001 land covers was believed to provide a better
assessment of long term habitat change. Only 2 years elapsed between the data collection
periods for the 1992 land cover (1989-1995) and the 2001 land cover (1997-2001).
Although 10 years elapsed between the 1978 land cover (1978-1979) and the 1992 land
cover (1989-1995), this was not considered sufficient for a robust assessment of land
cover change.

A key factor to consider when analyzing land cover change between 1978 and
2001 is that for the 1978 land cover, forested lands are defined as those which . .are at
least 10% stocked by forest trees of any size” (MDNR, 1981; http://www.crs.msu.edu/
pdf/lclu/CUI.pdf) and for the 2001 land cover, forested lands are defined as those which
have at least 25% canopy cover (http://www.dnr.state.mi.us/spatialdatalibrary/sd12/
land_use_cover/200l/[FMAP_lp_landcover.htrn) which equates to 16.7% stocked by
forest trees of any size (MDNR, 2001; http://www.rsgis.msu.edu/pdf/lclu/
Michigan_LC_LU_ Classification_System_2000.pdf). By these definitions it clear that
any gains in forest land cover from 1978 to 2001 are likely to be true gains, while losses
may be partly a function of the more conservative 2001 definition of forested lands.

Conifer forests increased from 11.7% of the total land cover in 1978 to 13.1% in
2001 (Table 7). While this may benefit G. sabrinus, it is important to note that as little as
51% of the conifer land cover that existed in Michigan in 1978 remained in the same
locations through 2001 (Table 6 and Figure 5). The gain in conifer land cover from 1978

to 2001 may benefit G. sabrinus if it represents an increase in the percentage of mature

35

conifer forests. However, if this gain is the result of a high rate of turnover of conifer
forests, this may be detrimental to the species.

The status of Michigan’s conifer forests as appropriate habitat for G. sabrinus
requires further investigation. Habitat variables important to G. sabrinus such as stand
age, the amount of down woody debris, and the distribution of fungi (a food source)

cannot be determined from satellite imagery.

Recommendations

More formal studies of the habitat requirements of G. sabrinus in Michigan
may reveal departures from those described in the literature. In order to more accurately
determine habitat use by both species of Glaucomys in Michigan, radio telemetry‘studies
such as those conducted by Stone et al. (1997) and Taulman and Seaman (2000) should
be considered. However, the primary emphasis of future research should be placed on
locating and monitoring G. sabrinus populations within the state. Such monitoring
should include genetic analysis of the remaining G. sabrinus populations, which may
reveal much about their current viability. Second, the region of the UP from the western
border of Seney National Wildlife Refuge eastward should be monitored for the
possibility of a continued advance of G. volans into that region.

To better understand the relationship between climate and the northern range limit

of G. volans, populations at the extreme northern edge of their range should be

monitored, particularly those within close proximity to weather stations. If, as Stapp et
al. (1991) propose, -15°C is physiologically meaningful to G. volans in terms of limiting

the northern range boundary, it may be instructive to compare the number of consecutive

36

days 5 —150C per year per weather station to the range expansion of G. volans in
Michigan over time.

Future efforts to capture flying squirrels should focus on either the use of nest
boxes specifically designed for flying squirrels (Althoff and Althoff 2001; Mahan et al.
1999) or traps placed in trees (Taylor and Lowman 1996), which although more labor
intensive, have been found to be more effective than ground trapping.

To the best of my knowledge, no current management strategy exists for G.
sabrinus or G. volans in the state of Michigan. Additionally, as these species are
currently not considered endangered, threatened, or game species, I could not find any
legislation regulating the take of flying squirrels in Michigan. Based on the findings of
this research, it would be advisable to implement a state sponsored monitoring plan for G.
sabrinus populations before federal intervention is required. A first step in such a plan
may include the requirement that all incidental take of flying squirrels, including their
capture locations, be reported to the MDNR and, where possible, specimens donated to
the nearest natural history museum. Currently, the only requirement for the incidental
take by commercial trappers is to. report that a “squirrel” was caught with no indication of

species or location.

37

LITERATURE CITED

ALTHOFF, D. R, AND P. S. ALTHOFF. 2001. Monitoring southern flying squirrel
populations with nest boxes, Ohio Journal of Science 101 :2-1 1.

ARBOGAST, B. S. 1999. Mitochondrial DNA phylogeography of the new world flying
squirrels (Glaucomys): implications for Pleistocene biogeography, Journal of
Mammalogy 80:142-155.

BAKER, R. H. 1983. Michigan Mammals. Wayne State University Press, Detroit.

BENDEL, P. R., AND J. E. GATES. 1987. Home range and rnicrohabitat partitioning of
the southern flying squirrel (Glaucomys volans), Journal of Mammalogy 68:243-
255.

BRADY, M. J ., T. S. RISCH, AND F. S. DOBSON. 2000. Availability of nest sites does
not limit population size of southern flying squirrels, Canadian Journal of
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CANADA DEPT. OF ENERGY MINES AND RESOURCES. 1974. The National Atlas
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CAREY, A. B. 2002. Response of northern flying squirrels to supplementary dens,
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CAREY, A. B., J. KERSHNER, B. L. BISWELL, AND L. DOMINGUEZ DE TOLEDO.
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COTTON, C. L., AND K. L. PARKER. 2000. Winter habitat and nest trees used by
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1086.

DOLAN, P. G., AND D. C. CARTER. 1977. Glaucomys volans, Mammalian Species
No. 78:1-6.

ENVIRONMENTAL SYSTEMS RESEARCH INSTITUTE, I. 1992-1995. ArcView GIS
3.2.

38

FRIDELL, R. A., AND J. A. LITVAI'TIS. 1991. Influence of resource distribution and
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Journal of Zoology 69:2589-2593.

GILMORE, R. M., AND J. E. GATES. 1985. Habitat use by the southern flying squirrel
at a hemlock-northem hardwood ecotone, Journal of Wildlife Management
49:703-710.

HARESTAD, A. S. 1990. Nest site selection by northern flying squirrels and Douglas
squirrels, Northwestern Naturalist 71:43-45.

HAVEMAN, J. R., AND W. L. ROBINSON. 1976. Northward range extension of the
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IHAKA, R., AND R. GENTLEMAN. 1996. R: A language for data analysis and
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IVAN, J. 8., AND R. K. SWIHART. 2000. Selection of mast by granivorous rodents of
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KURTA, A. 1995. Mammals of the Great Lakes Region. The University of Michigan
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squirrel, Glaucomys volans. M.S., Acadia University, Wolfville.

MADDEN, J. R. 1974. Female territoriality in a Suffolk county, Long Island, population
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MAHAN, C. G., M. A. STEELE, M. J. PATRICK, AND G. L. KIRKLAND, JR. 1999.
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MARTIN, K. J ., AND R. G. ANTHONY. 1999. Movements of northern flying squirrels
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MICHIGAN DEPARTMENT OF NATURAL RESOURCES. 2003. Integrated Forest
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Remote sensing image. Time period of content 1997-2001., Lansing.

39

MOWREY, R. A., AND J. C. ZASADA. 1982. Den tree use and movements of northern
flying squirrels in interior Alaska and implications for forest management, Pp.
351-356 in Fish and wildlife relationships in old-growth forests; proceedings of a
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Masters, Ashland.

MUUL, I. 1968. Behavioral and physiological Influences on the distribution of the flying
squirrel, Glaucomys volans, Pp. 1-66 in Miscellaneous publications Museum of
Zoology, University of Michigan, Ann Arbor.

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97

Sites trapped during summer 2002 field season. All localities are NAD27.

Appendix III

 

 

Date Trap line Start Finish
Latitude Longitude Latitude Longitude

5/7-5/10/02 1 45.114578 -84.426192 45.118581 -84.389490
5n-5/10/02 2 45.125600 -84.397852 45.123250 -84.397658
5/7-5/10/02 3 45.125485 -84.398 179 45.123315 -84.397707
5/7-5/10/02 4 45.128251 -84.404935 45.127215 -84.407306
5/7-5/10/02 5 45.127228 -84.407227 No Data No Data
5/21—5/24/02 1 44.169317 -84.833245 44.168491 -84.832963
5/21-5/24/02 2 44.169432 -84.832431 44.168230 ~84.832588
5/21-5/24/02 3 44.169304 -84.834388 44.168244 -84.834006
5/21-5/24/02 4 44.169244 84.832016 44.168184 -84.831848
5/21-5/24/02 5 44.167586 -84.839368 44.165759 —84.838768
5/21-5/24/02 6 44.167599 -84.83945 1 44. 166558 -84.842497
5/21-5/24/02 7 44. 167635 -84.839465 44.167968 -84.841014
5/28-5/31/02 l 45.263083 -84.430839 45.259486 -84.432740
5/28-5/31/02 2 45.261871 -84.430816 45.259079 -84.431336
5/28-5/31/02 3 45.299302 -84.425989 45.299789 -84.422056
5/28-5/31/02 4 45.299686 -84.426095 45.300065 -84.421883
5/28-5/31/02 5 45.296222 -84.428992 No Data No Data
5/28—5/31/02 6 45.281397 -84.422405 45.281636 -84.427537
5/28-5/31/02 7 45.281813 -84.421003 45.280099 84.417223
6/10-6/14/02 l 44.617968 -83.763305 44.614716 -83.762792
6/10-6/14/02 2 44.617712 -83.763791 44.614623 -83.765766
6/10—6/14/02 3 44.666560 -83.838452 44.665356 -83.842618
6/10—6/14/02 4 44.617777 -83.706396 44.618395 -83.701593
6/10-6/14/02 5 44.617385 -83.706588 44.617908 -83.701855
6/10—6/14/02 6 44.573372 -83.784222 44.572627 -83.785832
6/18-6/21/02 1 44.572297 -85.335715 44.571492 -85.333485
6/18—6/21/02 2 44.568669 -85.368610 44.571910 -85.369837
6/18-6/21/02 3 44.568018 -85.370212 44.564656 -85.369331
6/18-6/21/02 4 44.567960 ~85.378572 44.568123 -85.375287
6/18-6/21/02 5 44.568528 -85.378613 44.569668 -85.37S933
6/18-6/21/02 6 44.5421 10 -85.409041 44.542777 -85.404246
6/18-6/21/02 7 44.542392 ~85.409572 44.542304 -85.414001
6/22-6/24/02 1 45.040163 -83.559207 45.042152 -83.558334
6/22-6/24/02 2 45.039401 -83.558647 45.041405 -83.557741
6/22-6/24/02 3 45.009329 -83.51 1692 45.007477 -83.51 1712
6/22-6/24/02 4 45.009227 -83.512077 45.007479 -83.5 1 1976
7/1 1-7/12/02 1 46.330577 -86.65 8022 46.327895 -86.660159
7/1 1-7/12/02 2 46.330275 ~86.656891 46.3271 18 -86.657598
7]] 1-7/1 2102 3 46.342255 -86.548035 46.339618 -86.544600
7/1 1-7/12/02 4 46.342393 -86.547707 46.344340 -86.549702

98

7/ 1 1 -7/ 1 2/02
7/ 1 1 -7/ 1 2/02
7/ 1 3-7/ 14/02
7/ 1 3-7/14/02
7/1 3-7/ 14/02
7/13—7/14/02
7/ 1 3-7/14/02
7/ 1 3-7/ 14/02
7/ 15-7/17/02
7/ 1 5-7/ 1 7/02
7/ 1 5-7/ 1 7/02
7/15-7/17/02
7/24-7/25/02
7/24-7/25/02
7/24-7/25/02
7/27-7/29/02
7/27-7/29/02
7/27-7/30/02
7/27-7/30/02
7/3 1-8/ 1/02

7/3 1-8/ 1/02

7/3 1-8/ 1 /02

7/3 1 -8/ 1 /02

8/14-8/ 15/02
8/14-8/15/02
8!] 4-8/ 15/02
8!] 6-8/ 1 8/02
8/16-8/ 1 8/02
8/16-8/1 8/02
8/19-8/2 1/02
8/19-8/2 1/02
8/1 9-8/21/02

MN—wNI—UN-‘AWN—me—ANHAWN—GM#WN—O\M

46.350661
46.349982
46. 1 54932
46. 1 55096
46. 1 38080
46. 1 37992
46. 165853
46. 1 67 1 17
45.988540
45.989604
46.027927
46.027855
46.282720
46.282722
46.236144
46.465 395
46.463474
46.210177
46.210616
46.287630
46.287600
46.216560
46.216610
46.324194
46.323903
46.292130
46. 1 86028
46.185 138
46. 153860
46.074685
46.162925
46. 168129

-86.632689
-86.632 123
-86.546543
-86.545 137
-86.5 89208
-86.5 89069
-86.648499
-86.6465 1 1
—86.685322
-86.686062
-86.697060
-86.696029
-86.078945
-86.079325
-86.241 1 18
-86. 125652
-86. 123 134
-86.001959
-86.001 81 1
-85.939210
-85.938680
-85.968700
-85.965290
-86.449554
86.448667
-86.452903
-86.497762
—86.497534
-86.403676
86.459082
-86.448599
-86.45 1275

99

46.352102
46.348457
46.150847
46.151978
No Data

46.136778
46.163199
46.164020
45.985846
45.987241
46.029199
46.025670
46.280775
46.284885
46.236953
46.463634
46.462013
46.211487
46.212671
No Data

No Data

No Data

No Data

46.326188
46.324450
46.293418
46.187843
46.184889
46.153005
46.072548
46.162836
46.165852

-86.636759
-86.628510
-86.547407
-86.545118
No Data

-86.585051
-86.646576
-86.644325
-86.681402
-86.682977
-86.701823
-86.692548
-86.077613
-86.083615
-86.236694
-86. 123313
-86.121266
-85.998648
-85.999420
No Data

No Data

No Data

No Data

-86.447743
-86.444577
-86.454917
-86.500144
-86.494604
-86.401380
-86.459142
-86.445412
-86.452369

APPENDIX IV

Nest boxes surveyed for over-wintering Glaucomys from December 2002 to February

2003. All locations are N AD27.

 

Date

1211612002
1211612002
1211612002
1211612002
12/16/2002
1211612002
1211612002
1211612002
1211612002
1211612002
1211612002
1211612002
1211612002
1211612002
1211612002
1211612002
1211612002
1211712002
1211712002
12117/2002
12117/2002
1211712002
1211712002
1211712002
1211712002
1211712002
1211712002
1211712002
1211712002
1211712002
12/17/2002
1211712002
1211712002
1211712002
1211812002
12118/2002
1211812002
1211812002
1211812002
12118/2002
12118/2002

Type

Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck

Latitude

43.4362 17
43.436346
43.436200
43.502625
43.502488
43.502808
43.503488
43.503319
43.504674
43.504575
43.504415
43.500621
43.500531
43.499871
43.499935
43.498653
43.498731
43.433347
43.433436
43.432563
43.432565
43.432530
43.432625
43.432504
43.432636
43.486584
43.486166
43.486009
43.485768
43.485789
43.485336
43.485250
43.6131 10
43.613083

' 43.413213

43.449575
43.764003
43.764448
43.764812
43.786168
43.785944

Longitude Notes

85.665993
-85.665956
~85.666342
-85.5796 1 9
8557928 1
-85.578384
-85.577525
-85.575879
-85.570732
-85.570676
-85.570477
—85.587459
-85.587320
-85.589210
-85.589286
-85.592360
-85.592504
-85.665637
-85.665715
85.665278
-85.663151
-85.662983
-85.661456
-85.660901
-85.660157
85.640145
-85.642277
-85.642382
-85.644485
-85.644613
-85.646598
~85.646628
-85.480661
-85 .480746
-83.359881
-83.4 18696
-82.892274
-82.892109
-82.892354
-82.821684
~82.820820

100

Appendix I#

1 211 812002
1 211 812002
1 211 812002
1 212 1 12002
1 212 112002
1 212 112002
1 212 112002
1 2/2 1 [2002
1 212 112002
1 212112002
1 212 112002
1 212 112002
1 212 112002
1 212 112002
1 212 112002
1 212 112002
1 2/2 112002
1 212 112002
1 212 112002
1 212 112002
1212 112002
1 212 112002
1 212212002
1 212212002
1212212002
1 212212002
1 212212002
1 212212002
1 2122/2002
1 212212002
1 212212002
1 212212002
1 212212002
1 212212002
1 212212002
1 212212002
1212212002
1 212212002
1212212002
1 W2212002
1 212212002
1212212002
1 212212002
1/3/2003

1 [312003
11312003
1/3/2003
111 612003

Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck

43.785947
43.783280
43.782292
44.405668
44.405 347
44.405 1 27
44.365249
44.36568 1
44.365469
44.265954
44.265863
44.265459
44.2653 19
44.265229
44.265273
44.256368
44.2563 12
44.255083
44.238743
44.255746
44.237689
44.237271
44.664177
44.664202
44.665782
44.665735
44.561940
44.562101
44.561280
44.766451
44.454009
44.454082
44.455504
44.652092
44.456562
44.456674
44.456200
44.430744
44.430918
44.433956
44.433912
44.435906
44.430136
44.220274
44.2213 10
44.221326
44.222272
44.452724

-82.820439
-82.920223
-82.920052
-85.730920
-85.73 1 1 1 6
-85.73 1084
-85.820878
-85.827 101
-85.827 197
-85 .9 1235 1
-85.91 2491
-85.928609
-85.928696
-85 .938362
-85.938391
—85.9395 19
-85.939202
-85.933526
-85.783090
-85.935360
-85.918903
-85.919633
-84. 141219
-84. 141420
-84. 147000
-84. 147307
-83.806400
—83.806397
-83.805 362
-83.719687
-83.680378
-83.6807 10
-83.687320
~84.1 16670
-83.693286
-83.693264
-83.675223
-83.452868
-83.452647
83.459760
-83.459706
-83.465672
-83.460574
-85.7 13083
-85.708601
-85 .708277
-85.695740
-83.852249

101

11 1612003
11 1 612003
111 612003
111 612003
111612003
111 712003
111 712003
111712003
111 712003
111712003
1/1712003
111712003
111 712003
111712003
111712003
111712003
111712003
111712003
111712003
111712003
111712003
111712003
111712003
111712003
1117/2003
111712003
111 812003
111 812003
1118/2003
111812003
111 812003
111 812003
111812003
111 812003
111 812003
111 812003
111 812003
111 812003
111812003
111812003
111 812003
1123/2003
1123/2003
1123/2003
1123/2003
1123/2003
1123/2003
1123/2003

Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck

44.453 108
44.453385
44.452483
44.457049
44.457979
44.448552
44.446962
44.4469 16
44.445374
44.4465 14
44.445896
44.445940
44.444840
44.448231
44.448140
44.461287
44.459199
44.461280
44.432752
44.433576
44.433992
44.435077
44.434150
44.43571 1
44.435050
44.433947
44.437473
44.438792
44.440484
44.441497
44.442108
44.436165
44.585561
44.584714
44.582877
44.587012
44.588085
44.589004
44.587790
44.588281
44.587 195
44.570568
44.571846
44.572602
44.571806
44.574721
44.576082
44.576024

83.850468
83.846507
83.847414
83.806144
83.805503
83.82528 1
83.823665
83.823063
83.823674
83.826846
83.82255 1
83.82125 1
83.822262
83.823217
83.824401
83.824629
83.826485
83.826352
83.858476
83.856909
83.858142
83.858382
83.859507
83.861 130
83.861667
83.861676
83.862809
83.862845
83.863075
83.863096
83.861752
83.862935
83.884600
83.882406
83.882544
83.885563
83.883879
83.885377
83.882394
83.881562
83.882888
83.883 195
83.882499
83.883557
83.883780
83.871778
83.869797
83.868309

102

1 12312003
1 12312003
1 12312003
1 12312003
1 12412003
1 12412003
1 12412003
1 12412003
1 12412003
1 12412003
1 12412003
1 12412003
1 12412003
112412003
1 12412003
112412003
112412003
112412003
112412003
112512003
112512003
1125/2003
112512003
112512003
1 12512003
112512003
112512003
1125/2003
112512003
112512003
112512003
112512003
112512003
112512003
112512003
112512003
21612003

21612003

21612003

21612003

21612003

21612003

21612003

21612003

21612003

21612003

21612003

21612003

Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck

44.5753 1 8
44.541 109
44.540582
44.541463
44.55401 1
44.552347
44.55291 1
44.552196
44.551 691
44.552101
44.551660
44.553748
44.554180
44.552734
44.554664
44.555227
44.554769
44.554138
44.555096
44.574008
44.571992
44.571759
44.572130
44.577014
44.578320
44.579364
44.581647
44.581865
44.577394
44.576704
44.574969
44.575376
44.576916
44.577278
44.578301
44.576306
44.578199
44.579290
44.582886
44.582155
44.582172
44.582644
44.582888
44.583421
44.586792
44.587305
44.587344
44.580983

83.869686
83.87 178 1
83.873917
83.873493
83.885461
83.910286
83.885458
83.884602
83.882926
83.880090
83.879548
83.876418
83.873546
83.873321
83.875353
83.87641 1
83.8771 10
83.878192
83.879507
83.774975
83.773361
83.771639
83.771094
83.759796
83.760523
83.761900
83.763106
83.761353
83.758364
83.756815
83.757347
83.758835
83.730829
83.731979
83.733887
83.731221
83.693657
83.693365
83.694379
83.693503
83.692510
83.692802
83.693627
83.69421 1
83.700386
83.699249
83.702545
83.702984

103

2 Glaucomys, caught 1 G. volans

3 Glaucomys, caught 1 G. volans

Caught 3 G. volans

228

229

230, 231, & 232

21612003
21612003
21712003
21712003
21712003
21712003
21712003
21712003
21712003
21712003
21712003
21712003
21712003
21712003
21712003
21712003
21712003
2f712003
21712003
21712003
21812003
21812003
21812003
21812003
21812003
21812003
21812003
211 312003
2/ 1 312003
211 312003
211 312003
211 312003
211 312003
211 312003
211 312003
211 312003
21 1 312003
211 312003
211 312003
211 312003
211 312003
211 312003
211 312003
211 412003
211 412003
211412003
211 412003
2114/2003

Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Wood duck
Owl

Wood duck

44.580253
44.579032
44.580128
44.580294
44.579613
44.579488
44.578992
44.578349
44.577735
44.574929
44.575519
44.575322
44.575656
44.575865
44.577831
44.579654
44.579835
44.600824
44.600790
44.601049
44.599520
44.600124
44.600206
44.600012
44.598837
44.600487
44.596426
44.638332
44.636957
44.637255
44.638156
44.635881
44.639683
44.640869
44.640528
44.641892
44.641859
44.638644
44.594137
44.594454
44.594993
44.595735
44.595202
44.674916
44.670523
44.668690
44.674692
44.674734

83.703046
83.704523
83.682757
83.683633
83.683965
83.683203
83.682961
83.682560
83.682946
83.683251
83.682448
83.680651
83.681225
83.683498
83.679190
83.681854
83.680551
83.735181
83.741 132
83.740270
83.730739
83.731 154
83.732642
83.732397
83.731380
83.728519
83.760181
83.741967
83.741236
83.739803
83.737707
83.736040
83.736305
83.733316
83.735448
83.738332
83.738707
83.739201
83.782614
83.782736
83.784500
83.783398
83.782174
83.744910
83.745709
83.745271
83.741539
83.733365

104

Sciurus carolinensis

Glaucomys found week before

Sciurus niger

Sciurus carolinensis

Sciurus carolinensis

211 412003
211 412003
211412003
211 412003
211 412003

Wood duck
Owl
Wood duck
Owl
Wood duck

44.673844
44.671029
44.668744
44.668254
44.670534

83.730842
83.7291 10

83.736748 10 Glaucomys, caught 3 G.

83.733401
83.737612

volans 233, 234, & 235

 

105

n1111111111111111111111112111n

31