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This is to certify that the

thesis entitled

Sichuan, Ring-necked, and F1 Hybrid Hen Pheasant
Survival and Reproduction in Southern
Michigan Habitats

presented by

John Alan Niewoonder

has been accepted towards fulfillment
of the requirements for

Master of Science degree in Fish. & Wildl.

 

 

Q.

Major professor

Date 3/30/615—

0-7639 MS U is an Affirmative Action/Equal Opportunity Institution

 

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MSU I. An mm Action/Emil Opportunity Intuition
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”3-0.1

 

SICHUAN, RING-NECKED, AND F1 HYBRID HEN PHEASANT SURVIVAL
AND REPRODUCTION IN SOUTHERN MICHIGAN HABITATS

BY

John Alan Niewoonder

A THESIS

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

MASTER OF SCIENCE

Department of Fisheries and Wildlife

1995

ABSTRACT

SICHUAN, RING-NECKED, AND F1 HYBRID HEN PHEASANT SURVIVAL
AND REPRODUCTION IN SOUTHERN MICHIGAN HABITATS

BY

John Alan Niewoonder

Genetic differences in plumage, behavior and habitat
preferences have been demonstrated between Sichuan and ring-
necked pheasants. Survival and productivity of Sichuan, ring-
necked, and.F1 hybrid hens hatched and reared in captivity and
released in Barry and Eaton counties were evaluated. Four
release sites with varied. habitat. were selected and 96
Sichuan, 88 ring-necked and 76 F1 hybrid radio-collared hens
were released in early April of 1993 and 1994.

Survival probabilities of hybrids (0.350) were over
double the values for ring-necked (0.162) and Sichuan females
(0.105). Avian and mammalian predators killed 54% and 19% of
the hens, respectively. Clutch size of first nest attempts and
nest success for hybrids were intermediate between.Sichuan.and
ring-necked females. The heterotic effects of greatest
survival by hybrid females resulted in seasonal production of
3.1 chicks per hen released compared with 1.5 and 0.8 chicks

per hen for ring-necked and Sichuan hens, respectively.

ACKNOWLEDGEMENTS

This study was funded through the Wildlife Conservation
and.Restoration.Act Pittman-Robertson Project Number W-127-R.
All phases of the study were undertaken in conjunction with a
long-term.pheasant reSearch effort initiated by the Rose Lake
Wildlife Research Station.

This project required the efforts of many individuals.
Thanks are extended. to :members of the Rose Lake staff
including: Glenn Belyea, Greg Bragdon, Tom Cooley, Dave Creed,
Stephanie Hogle, Paula Somes, Bill Scullon and Bruce Warren
for assistance in handling of birds, instruction of telemetry
techniques, and necropsy work.

I would like to thank and acknowledge Dr. Harold H.
Prince, my major advisor, for his guidance, patience and
support. Additional thanks to Dr. Donald L. Beaver, and Dr.
Scott R. Winterstein for serving on my graduate committee.
Special thanks go to David R. Luukkonen for serving as
Michigan DNR.representative and co-investigator on this study
and for his invaluable assistance throughout the project.

Additional appreciation is extended to Carol Baumann,
Linda Briggs, Cheryl Cunningham, Dave Denomme, Chad.Fox, Anita

Moon, Steve Moore, Dave Morton, Lori Neely, Mark Smith and

iii

Michael Whitt for assistance with field work in 1993. Mark
Boersen, Dave Dortman, Elliot Grondin, Leslie Jagger, Mike
Kurncz, Mark Ledebuhr and Mike Parker are acknowledged for
assistance with field work in 1994.

Thanks to Cathy Flegel for assistance with analysis and
report preparation, Lori Neely for data entry and countless
hours of digitizing and to Gary Roloff for assistance with
GIS.

Thanks are also due to fellow graduate students Linda
Briggs, Jennifer Dorset-Derby, Cathy Flegel, Mike Monfils,
Timothy R. VanDeelen, Michael Whitt and especially Charlotte
E. Lawrence for assistance, support and friendship.

Finally, I would like to thank my parents Marvin and
Johanna, my brothers and sisters-in-law, and most of all my
wife Kristin for all their love, support and prayers over the

years.

iv

TABLE OF CONTENTS

LISTOFTABLESOOOOOOOOOOOOOOOOOOOOOOOOOOOOOO0.0... v1
LISTOFFIGURES.0.00000000000COOOOOOOOOOOO00...... Vii

INTRODUCTIONOOOOOOOOO00.0.0000...0.0.0....00...... 1

.b

METHODS.O.C.0.0000000000COOOOOOOOOOOO0.00.00.00.00

Study area...................................
Stocks.......................................
Rearing......................................
Release......................................
Dispersal....................................
Survival.....................................
Mortality....................................
Nesting......................................

Hid
o<3a3m~40\m4>

RESULTS...OOCCOOOCOOOOOOOOO00.000.000.000....00... 13

Land use/habitats............................ 13
Dispersal.................................... 15
Survival..................................... 16
Mortality.................................... 20
Nesting...................................... 22
Production................................... 27

DISCUSSIONOOOOOCOOOO0......OOOOOOOOOOOOOOOOOOOOOOO 30

LITERATURECITEDOO ..... OOOOOOOOOOOOOOOOOOOOO0.0... 38

LIST OF TABLES

Land use for the 4 study sites during
the 1993 field season............. ........

Dispersal distances (km) of Sichuan,
ring-necked, and Sichuan x ring-necked
F1 hybrid hen pheasants, combined over
all 4 study sites, during the 1993 and
1994 field seasons........................

Survival probabilities(i SE) of Sichuan,
ring-necked and F1 hybrid hen pheasants
for the 214 day study period(1 April -

31 October)...............................

ring-necked and F1 hybrid hen pheasants
during specific time periods.. ....... .....

Daily survival probabilities + SE of Sichuan,

Average clutch size (i SE) of first and
second nests of Sichuan, ring-necked,
and Sichuan x ring-necked F1 hybrid hen
pheasants, combined over all 4 study
sites, during the 1993 and 1994 field
seasons...................................

Nest success rates for all nests for
Sichuan, ring-necked, and Sichuan x
ring-necked hybrid hen pheasants,
combined over all 4 study sites, during
the 1993 and 1994 field seasons...........

Number of chicks hatched by race and nesting

attempt during the 1993 and 1994 field
seasons 0. OOOOOOOOOOOOOO 00.0.00... ........

vi

14

16

17

20

25

26

29

LIST OF FIGURES

1. Location of study areas in Barry and
Eaton counties of southern Michigan.
(T= Thornapple, B= Baltimore, K=
Kalamo/Maple Grove, 0= Oneida/Benton) .....

2. Sichuan, ring-necked and F1 hybrid
hen survival(1 April - 31 October)
during the 1993 and 1994 field
seasons........................... ........

3. Causes of mortality for Sichuan,
ring-necked and F1 hybrid hen
pheasants during the 1993 and 1994
field seasons. Other category
included farm equipment, exposure
and malnutrition.................... ......

4. Percentage of first nest initiations
for all hen pheasants, combined over
all study areas, during the 1993 and
1994 field seasons................. .......

vii

18

21

23

INTRODUCTION

Survival probability of female common pheasants
(Phasianus colchicus) is a function of weather conditions,
predator abundance, life stages, habitat preferences, habitat
availability and_ habitat juxtaposition. Pheasants select
distinct habitat types for prenesting, nesting, brood rearing,
fall assembly, and wintering (Hanson and Progulske 1973,
Whiteside and Guthery 1983, Myers et a1. 1988, Gatti et al.
1989, Leptich 1992). Availability of quality habitat that
meets the year-round requirements of pheasants is the key to
successful pheasant reproduction and survival.

Habitat loss and change has resulted in the decline of
ring-necked pheasants in southern Michigan that has occurred
over the past 40 years (Prince et al. 1988). The release of a
race of pheasant that can utilize a broad spectrum of habitat
types may result in a resurgence in numbers despite this
trend.

There are 32 races of the common pheasant in Asia.
Differences in these races have developed due to isolation and
uniqueness of native habitats (Delacour 1977, Johnsgard 1986) .

Some habitats in China that are used by common pheasants are

2
comparable to habitats in Michigan that are not being used by
ring-necked pheasants (P. c. torquatus).

It was determined, based on habitat utilization and
accessibility of native populations, that Strauch's pheasant
(P. c. strauchi) was the best suited for an introduction to
Michigan (Prince et a1. 1988). This sub-species is being
called the Sichuan pheasant by the Michigan Department of
Natural Resources (MDNR). It is believed that Sichuan
pheasants occupy a different habitat niche than the subspecies
currently established in North America.

The current strategy of the "Sichuan Project" is to
establish Sichuan pheasants in habitat in Michigan now void or
nearly void of common pheasants, and to maximize sizes of
foundingpopulations. Nearly all pheasant introductions in the
United States had European game farm background, having been
imported there from China hundreds of years earlier (Squibb
1985). In Europe, pheasants 'were ‘undoubtedly' exposed. to
inbreeding and artificial selection that may have decreased
genetic diversity and diminished the ability to survive in
wild conditions. Sichuan pheasants imported to Michigan were
also exposed to game farm conditions, however, this was for
only a short time and precautions were taken to prevent
inbreeding and other negative effects. It was hoped that
genetic input from the wild Sichuan line could help increase
genetic diversity of Michigan's pheasant population as well as

provide pheasants which utilize different habitats.

3
Investigation of the impact of the introduction of Sichuan
pheasants on ring-necked pheasant populations established from
previous programs can provide a baseline for research on the
importance of each subspecies and the hybrid condition to the
development of robust populations of pheasants.

This study is based on the premise that survival and
reproductive success of Sichuan, ring-necked, and Sichuan x
ring-necked F1 hybrid pheasants should be similar because of
the common genetic background. This study compares survival
and reproduction of the 2 races and their hybrids released in

common environments representative of southern Michigan.

METHODS

Stu rea

The release sites were selected to provide a range of
habitats suitable for comparison of the 3 lines of pheasant.
All sites were located in Barry and Eaton counties in southern
Michigan. Four township-sized areas (9,325 ha) were used as
release sites and included Baltimore township (T. 2 N., R. 8
W.) and Thornapple township (T. 4 N., R. 10 W.) in Barry
county, Oneida/Benton townships (T. 4 N., R. 4 W. S 1/2 and T.
3 N., R. 4 W. N 1/2) in Eaton county, and Kalamo/Maple Grove
townships (T. 2 N., R. 7 W. E 1/2 and T. 2 N., R. 6 W. W 1/2)
on the Barry-Eaton county border (Figure. 1).

Habitat and land use for each study area was evaluated
using the Geographical Information System (GIS) PC ARC/INFO.
Maps were generated from color-infrared aerial photos (36" x
36", scale = 1:14,500) by tracing habitat boundaries with fine
tipped permanent markers. Habitat boundaries were entered into
the data base on a 12" X 12" digitizing tablet. Ground

truthing was done during the spring and summer of 1993.

        

‘ BARRY EATON
T — i
‘ —‘ O

 

 

 

 

 

 

 

 

 

 

Fig. 1. Location of study areas in Barry and Eaton counties of southern Michigan
(T =Thornapple, B=Baltimore, K=KalamolMaple Grove, 0=Oneida/Benton).

Slacks

Four groups of 50 chicks, representing all breeding
combinations including Sichuan, ring-necked, and Sichuan x
ring-necked F1 reciprocal crosses were reared together and
maintained in outdoor pens throughout the entire year.
Sichuans were hatched from eggs laid by birds hatched from
eggs taken directly from Sichuan Province. Ring-necks were
hatched from eggs layed by Michigan ring-necked pheasant
stocks maintained by the Michigan DNR. The hybrids are the F1
products of reciprocal crosses between the Sichuans and the
Michigan ring-necks. Throughout the rearing process, care was
taken to provide a similar environment for all pheasants used

in this study.

Rearing

Eggs were collected daily and were cleaned and placed in
a cooler until they were ready to be set. Eggs were set each
Friday during the breeding season to synchronize hatching
dates. Incubation occurred until hatching on the 24th day,
after which chicks were moved into brooder rooms where they
remained for one week. For the next 5 weeks chicks were
allowed to move back and forth from the broader room to a set
of outdoor acclimation pens. Pheasants were then moved into
large flight pens where they remained throughout the winter
until spring (Bruce Warren, Mich. Dep. Nat. Resour., pers.

commun.).

Release

Necklace style radio telemeters (Lotek Engineering, Inc. ,
Newmarket, Ontario, Canada) weighing approximately 15 grams
and equipped with 15" antennas and 9 hour mortality sensors
were affixed to the hens in mid-March of both 1993 and 1994.
Pheasants were held in pens for 1 week after the transmitters
were attached to allow'the birds to become accustomed.to them.
Each of the four sites received 12 Sichuan, 10 ring-necked,
and 7 hybrid hens in 1993 and 12 Sichuan, 12 ring-necked and
12 F1 hybrid hens in 1994. No hens from 1993 survived long
enough to nest in 1994. This provided a total of 96 Sichuan,
88 ring-necked, and 76 hybrid radio-collared hens for the 2
years of this study. In addition, 10 non-radioed Sichuan.cocks
were released at each site on both years.

The release schedule was similar for both 1993 and 1994.
On 1 April, hen pheasants were released on Oneida/Benton and
Kalamo/Maple Grove sites. Two days later, releases were made
on the Baltimore and Thornapple sites. Specific release sites
were chosen on the basis of providing adequate escape cover
and were located as nearly as possible to the center of the
study areas. Releases were made before sunrise using brush
covered poultry crates. Crates were positioned adjacent to
protective cover and observed from a distance to ensure that
predators did not interfere with release. Pheasants were

allowed to exit the crates by walking into the surrounding

8
vegetative cover. This was done to minimize the stress of

handling and to facilitate a more natural and safe dispersal.

Dispersal

Dispersal distances were measured at 4 weeks and again at
8 weeks after release. Distances were measured as the
Euclidean distance from the release site to the center of the
grid cell (100 m X 100 m) in which the hen was located at the
end of each of the 2 time periods. Dispersal distance was
compared between the 3 pheasant lines using the Kruskal-Wallis
test. When significant differences were found, the Mann-

Whitney U test was used to determine pairwise differences.

Survival

All birds were located at least 3 times per week until
mid May when birds were then monitored 4 to 6 times per week.
Pheasants were monitored from early April through October by
technicians equipped with radio receivers, and hand-held, 3-
element directional antennas.

Survival was estimated over a 214 day period (1 April -
31 October). In addition, since pheasant survival is likely to
be different during various times of the year, survival was
estimated over 3 periods: the first month following release (1
April - 30 April), the peak nesting period (1 May - 31 July),
and the postbreeding period (1 August - 31 October).

Nine hour mortality sensors allowed early detection of

9

dead pheasants. When 1 or more days elapsed between locations
and mortality had occurred, pheasants were considered dead on
~the day after they were last found to be alive. Days between
locations rarely exceeded 2 days surrounding a mortality.

Survival was estimated using the Kaplan-Meier method
(Kaplan and Meier 1958). This method calculates a survival
probability estimate at the time of death for each radio-
tagged animal being monitored. Kaplan-Meier allows staggered
entries of individuals and the ability to censor individuals.
Individuals or observations may be censored because of radio
failure, radio loss, emigration from the study area, or
because the animal survives past the end of the study period.
The Kaplan-Meier method requires 3 assumptions: 1) the
censoring mechanism is random, 2) survival times are
independent among individuals, and 3) a random sample of
animals is obtained. Problems with the first assumption may
have caused us to overestimate actual survival. On several
occasions, damaged radios were found on dead hen pheasants. It
appears that automobiles, farming equipment and some predators
are capable of damaging transmitters when killing pheasants.
Generally when this happens, radios are not found and
individuals become censored when they should have been
considered dead. This problem may have also contributed to the
relatively high number of censored individuals in this study.
I believe that, due to the nature of pheasant behavior and the

source of pheasants used in this study, assumptions 2 and 3

10
did not. present. problems in ‘this study. Differences in
survival functions were determined using a log-rank test

(Pollock et al. 1989).

mug

The mortality sensors on the transmitters allowed us to
retrieve carcasses. quickly following mortality. Several
' observations could be made in the field which indicated cause
of death. Tooth.marks on transmitter or antenna, chewed bones,
sheared feathers, presence of mammalian tracks, scat, and
odor, buried remains and proximity of active dens indicated
mammalian predation. Straight or triangular beak marks on
transmitter or antenna, intact skeleton with mmscle tissue
missing, plucked feathers, and presence of raptor whitewash or
pellets were clues indicating avian predators (Rabe et a1.
1988).

Remains were frozen and later taken to Rose Lake Wildlife
Research Center for necropsy. Observations made in the field
and results of the necropsy usually allowed us to determine

cause of death.

Nesting

Incubating females and their nest sites were located with
the aid of radio-telemetry. Locations of nests were determined
and flags were placed in the general area of the nest.

Direction and distance of the nest in relation to the flags

11
were recorded. Care was taken not to disturb the birds or to
influence their behavior in any way.

Nest initiation dates were estimated by multiplying the
number of eggs by 1.3 (number of days required to lay 1 egg
(Campa et. al. 1987)) and backdating from the first day of
incubation. Initiation dates of first nests were then
converted to Julian days and compared using the nonparametric
Kruskal-Wallis test.

Clutch size was determined by visiting the nest while the
hen was off feeding or after eggs hatched or were abandoned.
Clutch size was compared between the 3 lines using the
nonparametric Kruskal-Wallis test.

Nest success ‘was estimated. by ‘the computer' program
Micromort (Heisey and Fuller 1985) which uses the modified
Mayfield model (Mayfield 1961, 1975). This model estimates a
daily survival rate from the number of days that each nest.was
known to be at risk and the number of nests that were
destroyed during that particular time period. This rate can
then be applied to the total number of days that each nest was
at risk including both the laying and incubation periods
(Miller and Johnson 1978). Daily survival probabilities may
differ between the egg laying and incubation stages, however,
we were unable to estimate survival during incubation. By
backdating, we can estimate nest success without ignoring
nests that.were not.detected.because they’were destroyed at an

early stage. If nests that are destroyed before they are

12

detected are not accounted for, nest success will tend to be
overestimated. Nest success rates of the 3 lines were compared
using a 2 'test (Hensler’ and. Nichols 1981). Nests were
considered successful if 1 or more eggs hatched. Destroyed
nests were considered unsuccessful on the day after they were
last observed. In all cases, nests were visited nearly every
day during incubation.

Multiple study areas, a large initial number of radio
tagged pheasants, and random sampling were used in this study.
This was done to achieve power and type I error rates that
would help ensure correct conclusions and completion of a
sound study. P values of g 0.10 were considered significant
for results in this study. Nonparametric statistics were
applied for dispersal distance, date of nest initiation for
first nests, and clutch size analysis as I was unable to

assume normal distributions for these comparisons.

RESULTS

u ab' t

Most of the land on the study areas is privately owned.
Farms, farm fields and woodlots are relatively small with a
high degree of fragmentation, creating a mosaic landscape
consisting of large amounts of linear shaped habitats. The
areas consisted of relatively small amounts of land enrolled
in the Conservation Reserve Program (CRP) . CRP land often
provides quality nesting habitat and winter cover for
pheasants. The Oneida/Benton site contained just 57 ha (<1 %
of the area) of CRP land, the Kalamo/Maple Grove site had 796
ha(8.5% of the area), the Baltimore site had 392 ha (4.3%),
and the Thornapple site contained just 183 ha (2.4%). In spite
of the lack of CRP land in these areas (< 5% of total area on
all sites), there existed varying amounts of idle land that
was not enrolled in the CRP program.

Land use and available habitat varied among release
sites. Most noticeably, the Oneida/Benton site had the highest
percentage of land in agricultural use (58%) while the
Baltimore site had the lowest (21%). Conversely, the Baltimore
site had the greatest area covered by woody plant species
(shrub areas, woodlots, and woody fence.rows) at 55% while the

Oneida/Benton site had woody cover over just 25% of its area.

13

14
The Kalamo/Maple Grove and Thornapple sites consisted of
similar amounts of cropland and woody cover and were

intermediate between Oneida/Benton and Baltimore (Table 1).

Table 1. Land use for the 4 study sites during the 1993 field
season.

 

Oneilflent gallM.G. Balgimore orn le
Landuse ha % ha % ha % ha %
Corn 2592 29 1086 12 771 9 1481 19
Soybeans 1512 17 784 8 189 2 454 6'
Wheat 702 8 126 48 1 175
Hay 345 4 1084 12 839 9 802 10
Herbaceous 594 1675 18 1289 14 1165 15
Shrubs 442 5 732 7 1479 16 581 8
Forest 1774 20 2792 30 3511 39 2050 27
Em.Wet1and 5 <1 107 l 178 67 1
Other 992 10 1022 11 730 8 956 12
Total 8958 9408 9034 7741

 

15

Dispersal

Dispersal of hen pheasants from release sites was
immediate and most birds appeared to move independently of
each other. Several birds did remain in groups of 2 or 3 for
a short time following release. Combining data from the 1993
and 1994 field seasons, pheasants moved an average of 1.4 i
0.1 km (n= 105) from the release sites after 4 weeks. Sichuan,
ring-necked, and hybrid dispersal distances were different
(Kruskal-Wallis, P= 0.017) during this time. Pairwise analysis
revealed that ring-necked and hybrid dispersal distances were
similar (Mann-Whitney U, P= 0.661), while Sichuans tended to
disperse shorter distances than both ring-necks (Mann-Whitney
U, £= 0.066) and hybrids (Mann-Whitney U, P= 0.010). After 8
weeks, hens had moved an average of 1.7 i 0.1 km (n= 77) from
the release sites. For this time period, dispersal distances
were similar (Kruskal-Wallis, P= 0.857) between the 3 lines

(Table 2).

16

Table 2. Mean dispersal distances (km) 1; SE of Sichuan, ring-
necked, and Sichuan x ring-necked hybrid hen pheasants,
combined over all 4 study sites, during the 1993 and 1994
field seasons.

 

Time since release

 

 

 

Pheasant line n 4 weeks n 8 weeks
Sichuan 35 1.1 i 0.1 24 1.3 i 0.2
Ring-necked 29 1.8 i 0.3 20 i 0.3
Hybrid 41 1.5 i 0.2 33 1.8 i 0.3
Combined 1.4 i 0.1 1.7 i 0.1
Survival

Survival probabilities were similar between years for
each of the 3 lines (log-rank test, largest.x2= 1.08, 1 df, P;
0.305). Combined over the 1993 and 1994 field seasons,
survival for the entire 214 day period was higher for hybrids
than for both ring-necks (log-rank test, x2: 7.67, 1 df, P:
0.006) and Sichuans (log-rank test, x2= 12.42, 1 df, P=
0.0004). Ring-neck and Sichuan survival was similar over this
time period (log-rank test, x2= 0.33, 1 df, P = 0.57) (Table
3, Figure 2). Of the 260 hen pheasants released, 182 died and

were found during the study, 48 were censored, and 30 were

still alive at the end of the study period.

17

Table 3. Survival probabilities (i SE) of Sichuan, ring-necked
and F1 hybrid hen pheasants for the 214 day study period (1
April - 31 October).

 

Pheasant line 1993 1994 Combined
Sichuan 0.11(i0.06) 0.10(:o.o7) 0.11(¢o.05)
Ring-necked O.17(i0.08) 0.16(10.07) 0.16(i0.05)

Hybrid 0.31(io.11) 0.37(i0.08) 0.35(io.07)

 

18

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19

Although low sample sizes prevented analysis of survival
on each of the 4 study sites, pheasant survival curves
appeared to differ somewhat between the sites. Ring-neck
survival was higher on the Oneida/Benton site, which was the
most intensively farmed and lower on the Baltimore site which
has the most woody cover. Sichuan survival, while relatively
low on all 4 sites was higher on the Baltimore site. Hybrid
pheasants, with relatively high survival on all study sites,
exhibited their highest survival on the Baltimore site as
well. For all 3 lines of pheasant, the Thornapple site
appeared to have the lowest survival of all sites.

Combining 1993 and 1994, survival for the first 30 days
after release was higher for hybrids (70%) than for ring-necks
(55%) (log-rank test, x2= 2.93, 1 df, 2= 0.087) and for
Sichuans (50%) (log-rank test, x2= 5.98, 1 df, _13 = 0.014)
which were similar (log-rank test, x2= 0.58, 1 df, 13 = 0.450) .
During the peak nesting season (1 May - 30 July), although not
significantly different, hybrid survival (58%) appeared to be
slightly higher than ring-neck survival (47%) (log-rank test,
x2= 1.24, 1 df, 13 = 0.266) and Sichuan survival (39%) (log-
rank test, x2= 1.94, 1 df, 2 = 0.164) which were again similar
(log-rank test, x2= 0.03, 1 df, P = 0.86). During the post-
breeding period (1 August - 31 October), hybrid, ring-neck and
Sichuan survival was not significantly different (76%, 58%,
and 60%, respectively) (log-rank test, highest x2: 0.33, 1 df,

g = 0.57).

20
Since the 3 periods are of different lengths of time, the
survival differential during these 3 time periods is more
clearly represented by daily survival rates. Daily survival
probability was lowest for all pheasants during the initial 30
days. Also, note that the greatest advantage for hybrids

occurred during the first 30 days following release (Table 4).

Table 4. Daily survival probabilities 1; SE of Sichuan, ring-necked, and F1
hybrid hen pheasants during specific time periods.

 

Pheasant Initial 30 Peak Post

line daysa nesting breedingc Total
Sichuan 0.97710.004 0.99010.006 0.99430.007 0.99010.003
Ring-neck o.9eo:o.oos 0.99210.007 0.99410.007 0.99110.002
Hybrid 0.98810.005 0.99410.006 0.99710.006 0.99510.001
Total 0.981.110.002 0.99210.003 0.99510.004 0.99210.001

 

° 1 April - 30 April.
b 1 May - 31 July.
C 1 August - 31 October.

Moitality

Cause of death was due mainly to predation from.avian and
mammalian predators (Figure 3). Causes of mortality did not
appear to be different between the 3 lines. The primary avian
predators identified by direct observation and location and
condition of the carcass were the red-tailed hawk (Buteo

jamaicensis) and.theigreat.horned.owl (Bubo Virginianus). The

21

 

 

 

 

 

 

 

 

RING-NECK ..

 

 

(n=54)

 

 

 

PHEASANT LINE

 

 

 

COMBINED
(n=] 72)

 

 

 

 

 

 

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0 1o 20 30 4o 50 60
PERCENTOFMORTALITY

 

 

Immom Don-ER

I AVIAN PREDATOR I MAMMALIAN PREDATOR 3523;?

 

AUTCMCBILE

 

 

Fig. 3. Causes of mortality for uncersored Sichuan, ring-necked and F1 hybrid hen
pheasants during the 1993 and 1994 field seasons. Other category included farm

equipment, exposure, and malmmition.

70

22
red fox (Vulpes vulpes) was identified as the most common
mammalian predator. Other factors, including farming activity,
road kills, exposure and malnutrition also contributed to
pheasant mortalities. For several birds, the information was

not sufficient to determine cause of death.

Mm

Initiation dates of all nest attempts ranged from 11
April to 27 July. The peak of first or initial nests occurred
in late April and early May (Figure 4). Comparison of first
nest initiation dates by female type, indicate that Sichuan
hens.may tend.to start their first.nests slightly earlier than
both ring-necks and hybrids. Mean nest initiation date was 4
May (i 6 days) for Sichuan hens, 6 May (i 7 days) for hybrid
hens and 11 May (i 7 days) for ring-necked hens. These dates
were not significantly different (Kruskal-Wallis, 2; 0.738).

One hundred-twenty nests were attempted during 1993 and
1994. Eighty-nine (74%) of these were first nests. Although
the 75 hybrid females represented only 29% of hens released in
this study, they accounted for 50 (42%) of all nest attempts.

We were able to determine clutch size for 86 of the 120
nests attempted. Although there was not a significant
difference (Kruskal-Wallis, 2; 0.115) between clutch sizes of
first nests for the 3 pheasant lines, it appeared that ring-

necks may have slightly larger initial clutch sizes than both

 

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hybrids and Sichuans (Table 5). Combining all nesting
attempts, no statistical difference was detected between the

3 lines (Kruskal-Wallis, P = 0.46).

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26

Nest success, calculated using the Mayfield model, was
37.1% for all nests during the 1993 field season. Sichuans
were less successful than both ring-necked hens (z= 1.774, g
= 0.077) and hybrid hens (z= 2.032, 2 = 0.042) which were
similar (z= 0.210, P = 0.834) in 1993. In 1994, overall nest
success was 30.6%. That year, nests of all races were equally
successful (highest z= 0.637, 2 = 0.522). Combined over both
years, 32.3% of all nests were successful. Sichuan hens were
less successful at nesting than ring-necked hens (z= 1.652, 2
= 0.099) and equally successful as hybrid hens (z= 1.428, 2,=
0.153) . Ring-neck hens and hybrid hens exhibited similar

success rates (z= 0.412, P =

0.682) (Table 6).

Table 6. Nest success rates for all nests for Sichuan, ring-
necked, and Sichuan x ring-necked hybrid hen pheasants,
combined over all 4 study sites, during the 1993 and 1994
field seasons.

 

 

 

 

1993 1994 Total
Race n Success n Success Success
Sichuan 20 0.19 20 0.26 0.22
Ring-
necked 16 0.46 14 0.36 0.40

Hybrid 17 0.50 33 0.31 0.36

 

27

Specific nest predators were difficult to identify.
Potential nest predators in this area include red fox, gray
fox (Urocyon cinereoargenteus), coyotes (Canis latrans),
raccoons (Procyon lotor), striped skunks (Mephitis mephitis),
opposums (Didelphis virginianus), mink (Mustela vison),
thirteen- lined. ground squirrels (Spermophilus
tridecemliniatus) and American crows (Corvus brachyrhynchos).
Nest predators destroyed 49 of 120 nests attempted. Farming
activity destroyed 9 nests, 5 hens were killed away from their
nests, and 4 hens abandoned their nests for unknown reasons.

Ninety-one percent (n=11) of Sichuan hens, 87.5% (n=8) of
ring-neck.hens, and 100% (n=13) of hybrid.hens that were still
living 1 month after their first nest was destroyed attempted
a second nest. No Sichuan hens (n=1), 50% (n=2) of ring-necked
hens, and 60% (n=5) of hybrid.hens that lived at least 1 month
after loss of their second nest attempted a third nest.
Renesting accounted for 25% of Sichuan broods, 18% of ring-
necked broods and 38% of hybrid broods. In addition, 1
Sichuan, 1 ring-necked, and 1 hybrid hen attempted to renest

after a successful nest had hatched earlier in the breeding

season.
r duc ' n

Combining all 3 lines, an estimated 225 chicks were

produced in 1993 and 212 chicks in 1994. Of these 437 chicks,

28
288 (65.9%) were from first.nesting attempts. Second and third
nesting attempts contributed 133 chicks (30.4%) and 16 chicks
(3.7%) respectively. Combining data from both years, the 96
Sichuan hens produced 76 chicks (0.79 chicks per hen
released), the 87 ring-necked hens produced 131 chicks (1.51
chicks per hen released) and the 75 hybrid hens hatched 230

chicks (3.07 chicks per hen released) (Table 7).

29

Table 7. Number of chicks hatched by race and nesting attempt
during the 1993 and 1994 field seasons.

Sichuan Ring-necked. lHybrid Total

 

1993
First nest 22 79 42 143
Second nest 8 12 53 73
Third nest 9 0 0 9
1994
First nest 22 24 99 145
Second nest 15 16 29 60
Third nest 0 0 7 7
Combined
First nest 44 103 141 288
Second nest 23 28 82 133
Third nest 9 0 7 16
All nests 76 131 230 437

Chicks/hen 0.79 1.51 3.07 1.69

 

DISCUSSION

The habitat present on the 4 study areas is a diverse
mixture of row crops, hay fields, idle fields, woodlots, and
scrub-shrub zones. The high degree of fragmentation that
ocCurs in these areas may result in habitat that provides for
year around needs of pheasants without requiring large
movements .in order to survive and reproduce. However, the
nature of this mosaic landscape pattern also provides large
amounts of linear shaped habitats. These types of habitats may
provide predators with an advantage for capturing prey and
locating nests that they may not have in other parts of North
America's pheasant range.

While ring—neck and hybrid dispersal distances were
slightly greater than Sichuan hen dispersal, distances
traveled from release sites by all 3 lines were comparable to
dispersal distances of pheasants found in the literature.
Warner(1988) reported that dispersal of released pheasants
ranged from 1.5 to 3 km. Wilson et al. (1992) and Rabe et al.
(1988) measured dispersal distances of 0.8 to 2.1 km.

Survival for all 3 lines in this study fell within or
below the ranges reported in the literature for wild trapped

hen pheasants and near or above survival rates of pen raised

30

31
pheasants. Survival rates of wild trapped hen pheasants ranged
from 20% to 55% (Dumke and Pils 1973, Snyder 1985, Penrod et
al. 1986, Wilson et al. 1992, and Lief 1994). For pen raised
birds, spring to fall survival rates ranged from 2-26% (Ellis
and Anderson 1963, Jarvis and Engbring 1976, Haensly et al.
1985, and Lief 1994).

Hybrid hen pheasant spring to fall survival rates were
considerably greater than the < 10% survival rates reported
for Sichuan x ring-necked hybrids in Pennsylvania (Johnson
1992) and were near the annual survival rate of wild hens of
30 - 35% suggested by Peterson et al. (1988) as the minimum
long term rate of a self—sustaining population. Sichuan and
ring-necked hen pheasant survival rates, however, were well
below this level. Populations with survival rates lower than
30% were assumed to be declining, while those with rates
greater than.35% were considered.to be increasing (Peterson et
al. 1988).

Low survival rates occurring in this study can be
partially explained by the fact that hand-reared pheasants
were studied. Hill and Robertson (1988) stated that hand-
reared pheasants tend to suffer heavy losses immediately after
being released and that these losses can reach a magnitude of
65% within 1 week and 80% during the first month. Hill and
Robertson (1988) also stated that hand-reared birds
consistently demonstrated a higher mortality rate than wild

birds, due to a greater vulnerability to predation. As a

32

consequence, fewer females survive the breeding season to
raise chicks than for their wild counterparts. Without the
benefits provided by the natural rearing of the hen,
unfamiliarity with wild food Sources and natural predators
result in low survival rates. Inbreeding depression, genetic
defects and lack of competitive ability in the offspring may
also contribute to low survival and productivity of hand
reared pheasants (Woodward et al. 1983). Sixty-one percent of
all deaths in this study occurred during the first month
following release.

Although hybrid daily survival was higher during the
first month following release, the peak nesting season and.the
post-breeding season, the greatest difference between hybrid
daily survival probability and daily survival probability of
Sichuan and ring-necked hens occurred during the first month
following release. It is not clearly understood why hybrid
survival is so much higher for the initial month following
release, however, it appears that.this time period is the most
crucial for hand-reared.pheasants. Increasing survival during
the first month following release may be the key to improving
success of future hand-reared pheasant releases.

Cause of death was attributed mainly to avian predators
especially red-tailed hawks and great horned owls. Mammalian
predators, mainly red fox, played a substantial but lesser
role. Road kill, death by farm implement, exposure, and

malnutrition were the cause of only a small fraction of total

33
deaths.

Pheasants seldom make up a major portion of an individual
predator's diet, however they are frequently eaten by a
variety of predators. In Wisconsin, 11 mammals and 9 raptors
are recorded as preying on pheasants (Wagner et al. 1965) .
Hessler et al. (1970) reported that red-tailed hawks, great
horned owls, red fox and mink (Mustela vison) commonly preyed
on pheasants with avian predators presenting the most serious
threat. . Rabe et al. (1988) found that mammalian predators
killed 28%, avian predators 12%, other, including motor
vehicle and farm related 7% and unknown 53% of pen reared
Sichuan hens in a study done in Michigan. In England and
Ireland, where avian predators capable of killing ring-necked
pheasants are rare, foxes are reported to be the predominant
predators. Hill and Robertson ( 1988) found in Ireland that
foxes accounted for 64% of all mortalities and 93% of all
predator related mortalities of pen raised pheasants.

Although several studies (Boag 1972, Herzog 1979, Johnson
and Berner 1980, Hines and Zwickel 1985) have commented on the
effects that radio transmitters have on bird behavior and
survival, this problem has become less of a concern as
technology has allowed reduction of transmitter weight.
Transmitter weight was less than the 2% of hen body weight
above which Warner and Etter (1983) reported an effect on
survival. Marcstrom et al. (1989) stated that necklace radio

transmitters, at 2 to 3% of body weight, are the most suitable

34
option for studies of pheasant survival. Birds observed in
pens for a short time following radio tagging did not appear
to have difficulty adjusting to the transmitters.

Comparison of first nest initiation dates by female type
indicate that although no statistical differences were
detected, Sichuan hens may tend to start their first nests
slightly earlier than both ring-necks and hybrids. These dates
were slightly later than the mean initiation date of 1 May for
first nests of ring-neck.hens that Dumke and.Pils (1979) found
in Wisconsin.

Mean clutch sizes for first nests (Sichuan= 8.3, ring-
necked= 10.6, hybrid= 9.2) demonstrate the genetic differences
between the 3 lines, which appear to be based on additive
genetic variance. Clutch sizes were equal to or slightly less
than clutch sizes commonly reported in the literature. For
ring-necks, Dumke and Pils (1979) reported a mean clutch size
of 11.8 eggs for first nests and 11.5 for second nests in
Wisconsin. Anderson (1964) found a mean of 10.2 eggs in
Illinois. In Michigan, Luukkonen (1993) reported mean clutch
sizes of first nests of 11.9 for wild trapped ring-necked
hens. .

Gates and Hale (1975) reported a range of 13- 46% nest
success, with an average of 24.6% for 18 studies of ring-
necked pheasants. This was lower than the 40% nest success
rate found for ring-necks in this study. In Michigan,

Luukkonen (1993) reported a higher nest success rate of 50%

35
for wild trapped ring-necks over a 3 year period. For Sichuan
hens, the 22% success rate that.we found was lower than the 40
% rate reported for Sichuans in Michigan by Padding (1988).
Hybrid nest success (36%) was similar to the 31% and 33%
success reported for Sichuan x ring-neck hybrids in
Pennsylvania (Johnson 1992).

Occurrence of renesting following predation of first
nests was higher for all 3 races than was found in the
literature. Dumke and Pils (1979) found that 68% of ring-
necked hens renested after their first nest was destroyed or
abandoned. However, the 41% that renested after the second
nest was destroyed in that study was much higher than was
found in this study. They also found that, similar to this
study, renesting efforts produced 40% of all broods in a
breeding season.

Perhaps the best measure of the fitness of an introduced
animal to its new'habitat is the number of young produced.plus
the survival and reproductive capacities of these young.
Spring releases of adult hens in Oregon (Jarvis and Engbring
1976) resulted in production of just 0.05 poults fledged per
female released. Ellis and Anderson (1963) also reported 0.05
young per hen in Illinois. Haensly et al. (1985) reported
production of 0.18- 0.28 young per female in Oregon. Brittas
et al. (1992) reported production of 1.0- 1.7 chicks fledged
per hen in a growing pheasant population on an island in

Sweden. We were unable to measure chick survival to fledge or

36

the subsequent reproductive abilities of these offspring but
did obtain information on number of chicks hatched by race
during the first breeding season following release. Sichuan
hens hatched 0.79, ring-necked.hens 1.51, and hybrid hens 3.07
chicks per hen released during this study. The lower
production exhibited by the Sichuan hens can be attributed to
a combination of low adult survival, small clutch sizes, and
low nest success rates. The greater production achieved by the
hybrid hens can be attributed mainly to the high adult
survival which allowed more hens to survive until the nesting
period.

In southern Michigan habitats, Sichuan hens survived at
a lower rate, had smaller clutch sizes and had lower nest
success probabilities than ring-necked and hybrid hens. The
low adult survival and nest success of Sichuan hens may be due
to habitat preferences that increase susceptibility of hens to
predators and nests to predation.

Hybrid hen reproductive parameters (clutch size and nest
success) were intermediate between ring-necked hens and
Sichuan hens, which seems to indicate additive genetic
expression. Hybrid adult hen survival, meanwhile was much
higher than both ring-necked and Sichuan hen survival in both
1993 and 1994. This heterotic response of reduced
vulnerability to predators with correspondingly higher
survival requires additional analysis for clarification and

may be the result of habitat selection. Sichuan and ring-

37

necked pheasants interbreed readily and the hybrid condition
should make up a significant segment of pheasant populations
where Sichuans have been released. Information on how long the
heterotic effects persist in a natural population is very
important in determining future pheasant release strategies.

The survival probabilities and reproductive success of
known Sichuan x ring-necked hybrid hen pheasants relative to
pure Sichuan and Michigan ring-necks shed light on factors
affecting success of releases of Sichuan pheasants in
Michigan. The release of F1 hybrid pheasants may also provide
managers with another tool to enhance the recreational

benefits from local pheasant populations.

LITERATURE CITED

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released in southern Illinois. J. Wildl. Manage. 28: 254-
264.

Boag, D. A. 1972. Effect of radio packages on behavior of
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Brittas, R. , V. Marcstrom, R. E. Kenward, and M. Karlbom.
1992. Survival and breeding success of reared and wild
ring-necked pheasants in Sweden. J. Wildl. Manage.
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Campa, H., III, M. L. Rabe, P. I. Padding, E. J. Flegler, Jr.,
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and . 1979. Renesting and dynamics of nest
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38

39

Haensly, T.F., S.M. Meyers, J.A. Crawford and W.J. Castillo.
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40

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