HABITAT REQUIREMENTS of RING-NECKED PHEASANT HENS (Phasianus Colchicus)

Home , Lagopus, Mallard, Perdix, Tympanuchus

HABITAT REQUIREMENTS OF RING-NECKED PHEASANT HENS (Phasianus colchicus)

ON FARMLAND IN LOWER AUSTRIA DURING NESTING AND BROOD REARING

THOMAS HOESMAN BLISS

(Under the Direction of John P. Carroll)

ABSTRACT

Ring-necked pheasants (Phasianus colchicus) are an important game species in Austria where populations have declined precipitously during the last half century. Given the lack of knowledge of populations within Austria, this research was conducted to determine habitat use and survival of pheasant hens during the breeding season and for broods during the first 21 days after hatching. Hen pheasants establish home ranges around set-aside and wetland habitats and nest in set aside. Brooding hens prefer to use set aside habitat and that game crop positively affects brood survival. Predation is the main reason for loss of hens and broods. Therefore, in order to increase the population of pheasants I suggest increasing the availability of set aside, wetland, and game crop habitats while incorporating supplemental feeding and predator control.

INDEX WORDS: Ring-necked pheasant, Austria, Brood, Habitat Use, Home Range, Survival

HABITAT REQUIREMENTS OF RING-NECKED PHEASANT HENS (Phasianus colchicus)

ON FARMLAND IN LOWER AUSTRIA DURING NESTING AND BROOD REARING

THOMAS HOESMAN BLISS

B.S., University of Tennessee: Chattanooga, 1997

A Thesis Submitted to the Graduate Faculty of The University of Georgia in Partial Fulfillment

of the Requirements for the Degree

MASTER OF SCIENCE

ATHENS, GEORGIA

2004

Thomas Hoesman Bliss

HABITAT REQUIREMENTS OF RING-NECKED PHEASANT HENS (Phasianus colchicus)

ON FARMLAND IN LOWER AUSTRIA DURING NESTING AND BROOD REARING

THOMAS HOESMAN BLISS

Major Professor: John P. Carroll

Committee: Karl V. Miller William Palmer

Electronic Version Approved:

Maureen Grasso Dean of the Graduate School The University of Georgia December 2004

DEDICATION

I dedicate this thesis to my loving wife, Angela, who had the strength to allow me to pursue this endeavor and for her patience and understanding during my time in Austria.

ACKNOWLEDGEMENTS

The funding for this project was a joint effort between Gutsverwaltung Hardegg, Austria,

The Game Conservancy Trust, Fordingbridge, England, and the University of Georgia Warnell

School of Forest Resources, USA. I personally want to thank the following people.

John Carroll, my major professor, who took me on as graduate student and gave me the wonderful opportunity to work in Austria. Also, for his patience and positive outlook even during the frustrating periods encountered while conducting research abroad.

To Roger Draycott, for his friendship and support while conducting research. I appreciated the much needed change of pace during your visits. Additionally, for your invaluable knowledge of pheasants which helped me out tremendously in the field and while writing.

To Maximilian and Alexandra Hardegg, for their friendship and for welcoming a total stranger into their home. Also, for their love of the outdoors which without this research would not have been possible.

Finally, I want to thank Karl Pock, for his friendship and patience in helping out an

American, who has no grasp of the German language, get adjusted to life in Seefeld. Also for his valuable knowledge of wildlife which was imparted to me, and his ability to get my mind off work when I needed a break.

TABLE OF CONTENTS

Page

ACKNOWLEDGEMENTS...... v

CHAPTER

1 STATUS AND DISTRIBUTION OF RING-NECKED PHEASANTS ...... 1

LITERATURE REVIEW...... 1

LITERATURE CITED...... 4

2 HOME RANGE, HABITAT USE, AND SURVIVAL OF HEN PHEASANTS IN

LOWER AUSTRIA ...... 9

INTRODUCTION...... 9

STUDY AREA...... 10

METHODS...... 12

RESULTS...... 16

DISCUSSION ...... 19

LITERATURE CITED...... 21

3 HOME RANGE, HABITAT USE, AND SURVIVAL OF PHEASANT BROODS IN

LOWER AUSTRIA ...... 40

INTRODUCTION...... 40

STUDY AREA...... 41

METHODS...... 42

RESULTS...... 46

DISCUSSION ...... 48

LITERATURE CITED...... 51

4 MANAGEMENT IMPLICATIONS FOR RING-NECKED PHEASANTS IN

LOWER AUSTRIA ...... 69

MANAGEMENT IMPLICATIONS...... 69

LITERATURE CITED...... 72

APPENDICES ...... 74

A Patagial tag, date captured, weight (g), tarsus (mm), condition index (weight/tarsus),

feather shaft (mm), age, capture location, and cover at trap of hen pheasants on

Seefeld Estate, Lower Austria, Austria 2002 ...... 74

B Patagial tag, date captured, weight (g), tarsus (mm), condition index (weight/tarsus),

feather shaft (mm), age, capture location, and cover at trap of hen pheasants on

Seefeld Estate, Lower Austria, Austria 2003 ...... 76

C Models considered to determine the effect of dispersal (A), condition index (B), age

(C), agriculture (D), set aside (E), woodland (F), amount edge within the home

range (G), game crop (H), and wetland (I) upon the survival of pheasant hens on

Seefeld Estate, Lower Austria, Austria 2002 – 2003 ...... 78

D Models considered to determine the effect of condition index of hen (A), age (B),

game crop (C), wetland (D), amount edge within the home range (E) nesting

habitat (F), woodland (G), agriculture (H), and set aside (I) upon survival of

pheasant broods on Seefeld Estate, Lower Austria, Austria 2001 – 2003 ...... 81

vii

E Example of a small breeding season 100% MCP home range (2.7 ha) using #1577.

Observe mixture of preferred hen habitats, set aside and wetland, along with male

territory habitat, woodland edge and coppice. Seefeld Estate, Lower Austria,

Austria 2002-2003...... 84

F Example of a large breeding season 100% MCP home range (264 ha) using #1595.

Notice low amount of male territory habitat, woodland and coppice, within the

home range. Seefeld Estate, Lower Austria, Austria 2002-2003 ...... 85

G Example of hen dispersal using #1570 showing the 100% MCP home range during

the breeding season on Seefeld Estate, Austria during 2002 ...... 86

H Example of a small 100% MCP brood home range (2.6 ha) using #1512. Home range

is comprised of well managed set aside, game crop and wetland habitats which

provide food and cover. Seefeld Estate, Lower Austria, Austria 2002-2003...... 87

I Example of an average 100% MCP brood home range (11 ha) using #1573, which

contains a good mixture of set aside, game crop, wetland, and agricultural land.

Notice that majority of locations lie within game crop. Seefeld Estate, Lower

Austria, Austria 2002-2003 ...... 88

J Example of a large 100% MCP brood home range (44 ha) using #1568. Observe that

the home range is dominated by agricultural land which increased the area covered

in order to find preferred set aside habitat. Seefeld Estate, Lower Austria, Austria

2002-2003...... 89

viii

CHAPTER 1

STATUS AND DISTRIBUTION OF RING-NECKED PHEASANTS

LITERATURE REVIEW

The Common (or Ring-necked) pheasant (Phasianus colchicus) is native from the shores

of the Black and Caspian seas east to Manchuria and Korea and to Taiwan and Japan (Hill and

Robertson 1988, Johnsgard 1999). Its name is derived from the Phasis Valley in Colchis, an area

within the Republic of Georgia, where it was found by Jason and the Argonauts who is believed

to have brought it to Europe around 1300 BC (Robertson 1997). Hill and Robertson (1988)

consider it one of the most widely introduced bird species by man and its present distribution

covers over 50 countries on 6 continents (Long 1981).

Unlike many other large gamebirds, such as the Greater Prairie-Chicken (Tympanuchus cupido) (Westemeier 1980), and Lesser Prairie-Chicken (Tympanuchus pallidicinctus) (Crawford

1980, Geisen 1998) in the United States, and the Great Bustard (Otis tarda) in Europe (Glutz et al. 1973, Robertson 1997), whose distributions and populations have diminished with agriculture expansion, pheasants have thrived and increased their range with the expansion of agriculture

(Robertson 1997). However, intensification of agriculture practices in the latter half of the 20th century throughout Europe has had a negative impact upon wild pheasant populations (Hill and

Robertson 1988, Tapper 1999, Csanyi 2000), and other bird species associated with agricultural lands (Tucker and Heath 1994, Tucker 1997, Potts 1997). The use of machinery and pesticides increased rapidly after World War II, and in England the number of pesticide sprayers increased from 2,000 to 51,000 during 1944 – 1962, and field size expanded from 6.5 ha to 11.0 ha (Potts

1986). Similar trends have also been observed throughout Europe and the United States.

Austria is no exception to agricultural intensification and has seen a decline in pheasant

populations throughout much of the latter part of the 20th century. The Austrian landscape, like

the rest of Europe, is dominated by agriculture which covers 80% of the land and is the base of

their economy (Molterer 1997). Lower Austria (Neiderösterrich), a province in northeast

Austria, has the highest concentration of farms and the greatest amount of land under production

(Molterer 1997). This region has seen a reduction in harvest of pheasants from a high of 18 per

100 hectares in 1972 to 4 per 100 hectares in 2000 (Draycott et al. 2002). Unlike Great Britain

and North America, where pheasant research has been extensive (Giudice and Ratti 2001), little

is known about the life history and habitat requirements of wild pheasant populations in Austria.

Studies on pheasants have suggested 4 crucial periods that affect abundance: 1) winter

survival, 2) male territories and breeding habitat, 3) nest habitat and success, 4) and brood habitat

and survival. The only previous research conducted in Austria examined winter habitat use and

survival on Seefeld Estate, in Lower Austria (Anderson 2002). Lack of winter habitat and

weather can cause over winter losses of 82% (Dumke and Pils 1973, Perkins et al. 1997), but

over winter losses at Seefeld were only 3% (Anderson 2002). This suggests that winter habitat is

sufficient and observations found that reed grass (Phragmites communis) and woodland coppice

were used extensively on Seefeld Estate during winter (Anderson 2002), and is similar to what

Robertson et al. (1993) found in England and to studies by Perkins et al. (1997) and Gabbert et

al. (1999) in the U.S. Seefeld, however, is exceptional where these habitats have been maintained and restored.

Breeding territory counts have been collected on Seefeld Estate from 1992 – 2004, where territories are established along the edges of woodland, hedge rows, and edges of wetlands

(Draycott et al. 2002), and follows the techniques described by Hill and Robertson (1988).

Females are attracted to territories where they form harems (Robertson et al. 1993) and feed under the protection of the territorial male to build up fat reserves for egg-laying (Draycott et al.

2002). On Seefeld, the number of territories has increased from 10/km2 in 1992 to 19 /km2 in

2004, but few non-territorial males observed which suggests that breeding densities are not yet

limited by habitat (Robertson et al. 1993).

Currently, limited data is available on nest habitat and success and brood habitat use and

success in Austria. Studies on nesting in other areas suggest that initial nests are located in

weedy areas (Baxter and Wolfe 1973, Snyder 1984, Warner and Joselyn 1986, Clark et al. 1999)

and residual cover (Olsen 1977) that provide sufficient cover and protection from predation.

Hens in England prefer to place initial nests in residual cover and woodlands with gradual shift

to cereal fields (Hill and Robertson 1988), whereas in the U.S. crops, such as alfalfa (Medicago

satvia) and growing winter wheat (Snyder 1984, Whiteside and Guthery 1983) are often

preferred.

Brood habitat can limit pheasant populations if suitable nesting cover is available and

winter survival is high (Warner 1979, Robertson 1996). Broods use habitats that contain

abundant arthropods, such as rough grass and weedy strips in England (Hill and Robertson

1988), and grassland and legumes in the United States (Nelson et al. 1990, Riley et al. 1998). In

the United Kingdom increased use of pesticides reduced the diversity of weeds and a 50%

decrease in insect abundance (Southwood and Cross 1969). Insects comprise a high percentage

of the diet of pheasant chicks (Hill 1985), and are necessary for proper development of gray

partridge (Perdix perdix), red grouse (Lagopus l. scotius), and red legged partridge (Alectoris

rufa) (Cross 1966, Hudson et al. 1994).

Home range size also affects the survival of pheasant broods. Fragmentation of suitable brood habitat increases the distance that chicks must move while foraging, which causes more energy to be expended and increases mortality (Hill 1985). Fragmentation increases use of road sides and edge habitat that can increase depredation (Warner 1984, Riley et al. 1998). Home ranges of broods range from 2-11 ha in the United States (Kuck et al. 1970, Hanson and

Progulske 1973, Warner 1979) and 4.8 ha in England (Hill 1985).

Given the paucity of information on pheasant populations during the breeding season in

Austria, the purpose for this study is to provide information about the habitat use and success of pheasant hens while nesting and rearing broods on Seefeld Estate, Austria. The specific objectives of this research are.

1. Estimate survival of pheasant hens during the breeding season.

2. Determine home range and habitat use of pheasant hens prior to nesting.

3. Determine nest habitat use, nest success, and habitat preference for nests.

4. Estimate home range and habitat use by hens with broods.

5. Estimate survival and the influence of habitat on brood survival.

LITERATURE CITED

Anderson, B.C. 2002. Habitat use and nesting ecology of Ring-necked Pheasant (Phasianus

colchicus) on a landscape dominated by agriculture in Lower Austria. Thesis, University

of Georgia, Athens, Georgia, USA.

Baxter, W.L., and C.W. Wolfe. 1973. Life history and ecology of the ring-necked pheasant in

Nebraska. Nebraska Game and Parks Commission, Lincoln, Nebraska, USA.

Clark, W.R., R.A. Schmitz, and T.R. Bogenschutz. 1999. Site selection and nest success of

ring-necked pheasants as a function of location in Iowa landscapes. Journal of Wildlife

Management 63:976-989.

Crawford J.A. 1980. Status, problems, and research needs of the lesser prairie-chicken. Pages 1-

7 in Vohs P.A. and Knopf F.L., editors. Proceedings prairie grouse symposium.

Oklahoma State University, Stillwater, Oklahoma, USA.

Cross, D.A. 1966. Approaches toward an assessment of the role of insect food in the ecology of

gamebirds, especially the partridge (Perdix perdix). Dissertation, University of London,

London, United Kingdom.

Csànyi, S., 2000. The effect of hand-reared pheasants on the wild population in Hungary: a

modeling approach. Hungarian Small Game Bulletin 5:71-82.

Draycott, R.A.H., K. Pock, and J.P. Carroll. 2002. Sustainable management of a wild pheasant

population in Austria. Zeitschrift fur Jagdwissenschaft 48:346-353.

Dumke, R.T., and C.M. Pils. 1973. Mortality of radio-tagged pheasants on the Waterloo

Wildlife Area. Wisconsin Department of Natural Resources. Technical Bulletin 72,

Madison.

Gabbert, A.E, A.P. Leif, J.R. Purvis, and L.D. Flake. 1999. Survival and habitat use by Ring-

necked Pheasants during two disparate winters in South Dakota. Journal of Wildlife

Management 63:711-722.

Geisen, K.M. 1998. Lesser Prairie-Chicken. Birds of North America 364:1-20.

Giudice, J.H., and J.T. Ratti. 2001. Ring-necked Pheasant. The Birds of North America

572:1-31.

Glutz von Blotzheim, U.N., K.M. Bauer, and E. Bezzel. 1973. 247-281 in Handbuch der Vögel

Mitteleuropas (Volume 5). Akademische Verlagsgesellschaft, Frankfurt am Main,

Germany.

Hansen, L.E. and D.R. Progulske. 1973. Movements and cover preferences of pheasants in

south Dakota. Journal of Wildlife Management 37:454-461.

Hill, D.A. 1985. The feeding ecology and survival of pheasant chicks on arable farmland.

Journal of Applied Ecology 22:645-654.

_____, and P.A. Robertson. 1988. The pheasant: ecology, management and conservation.

Blackwell Scientific, London, United Kingdom.

Hudson, P., F. Booth, M. Hurley. and D. Howarth. 1994. Problems with red grouse chick

survival. Game Conservancy Annual Review 25:120-122.

Johnsgard, P.A. 1999. The pheasants of the World. 2nd edition. Smithsonian Institute Press,

Washington, D.C., USA.

Kuck, T.L., R.B. Dahlgren, and D.R. Progulske. 1970. Movements and behaviour of hen

pheasants during the nesting season. Journal of Wildlife Management 34:626-630.

Long, J.L. 1981. Introduced birds of the world: the worldwide history, distribution, and

influence of birds introduced to new environments. University Books, New York, USA.

Molterer, W. 1997. Agriculture in Austria - managing in harmony with nature. Austrian

Information 50 (11):1-8.

Nelson, D.R., R.O. Kimmel, and M.J. Frydendall. 1990. Ring-necked Pheasant and Gray

Partridge brood habitat in roadsides and managed grasslands. Perdix V:103-112.

Olsen, D.W. 1977. A literature review of pheasant habitat requirements and improvement

methods. Utah State Department of Natural Resources. Publication number 77-7, Salt

Lake City, Utah, USA.

Perkins, A. L., W.R. Clark, T.Z. Riley, and P.A. Vohs. 1997. Effects of landscape and weather

on winter survival of ring-necked pheasants hens. Journal of Wildlife Management

61:634-644.

Potts, G.R. 1986. The Partridge: pesticides, predation and conservation. William Collins Sons

& Co. Ltd. London, United Kingdom.

_____, 1997. Cereal farming, pesticides and Grey Partridges. Pages 150-170 in Pain, D.J. and

M.W. Pienkowski, editors. Farming and birds in Europe: The common agricultural

practices and its implications for bird conservation. Academic Press, London, United

Kingdom.

Riley, T.Z., W.R. Clark, D.E. Ewing, and P.A. Vohs. 1998. Survival of ring-necked pheasant

chicks during brood rearing. Journal of Wildlife Management 62:36-44.

Robertson, P.A., M.I.A. Woodburn, W. Neutal, and C.E. Bealey. 1993. Effects of land use on

breeding pheasant density. Journal of Applied Ecology 30:465-477.

_____, 1996. Does nesting cover limit abundance of Ring-necked Pheasants in North America?

Wildlife Society Bulletin 24:98-106.

_____, 1997. A natural history of the Pheasant. Swan Hill Press. Shrewsbury, United

Kingdom.

Snyder, W.D. 1984. Ring-necked pheasant nesting ecology and wheat farming on the High

Plains. Journal of Wildlife Management 48:878-888.

Southwood, T.R.E., and D.J.Cross. 1969. The ecology of the partridge III. Breeding success

and the abundance of insects in natural habitats. Journal of Animal Ecology 38:497-509.

Tapper, S.C. 1999. A question of balance: game animals and their role in the British

Countryside. The Game Conservancy Trust. United Kingdom.

Tucker, G.M., and M.F. Heath. 1994. Birds in Europe: Their conservation status. Birdlife

Conservation Series Number 3. Cambridge: Birdlife International.

_____, 1997. Priorities for bird conservation in Europe: the importance of the farmland

landscape. Pages 79-116 in Pain, D.J. and M.W. Pienkowski, editors. Farming and birds

in Europe. Academic Press, London, United Kingdom.

Warner, R.E. 1979. Use of cover by pheasant broods in east-central Illinois. Journal of Wildlife

Management 43:334-346.

_____, 1984. Effects of changing agriculture on ring-necked pheasant brood movements in

Illinois. Journal of Wildlife Management 48:1014-1018.

_____, and G.B. Joselyn. 1986. Response of Illinois Ring-necked Pheasant populations to block

roadside management. Journal of Wildlife Management 50:525-532.

Westemeier, R.L. 1980. Greater Prairie Chicken status and management 1968-1979. Pages 8-

17 in Vohs P.A. and Knopf F.L., editors. Proceedings prairie grouse symposium.

Oklahoma State University, Stillwater, Oklahoma, USA.

Whiteside, R.W., and F.S. Guthery. 1983. Ring-necked Pheasant movements, home ranges, and

habitat use in west Texas. Journal of Wildlife Management 47:1097-1104.

CHAPTER 2

HOME RANGE, HABITAT USE, AND SURVIVAL OF HEN PHEASANTS IN LOWER

AUSTRIA

INTRODUCTION

During spring, common or ring-necked pheasant (Phasianus colchicus) hens begin to

disperse from winter habitats in order to find mates, nest sites, and brood rearing habitat (Dumke

and Pils 1973). Several studies have shown habitat selection and reproduction success (Clark et

al. 1999), as well as disturbance factors (Warner and Etter 1985), are important in understanding the dynamics of pheasant populations. Throughout Europe, pheasant populations have generally decreased since WWII (Hill and Robertson 1988, Tapper 1999, Csànyi 2000), and this is attributed to loss of habitat by agricultural intensification (Jarvis and Simpson 1978, Hill 1985,

Potts 1991). These changes are apparent in harvest data from Lower Austria, where the harvest rate by sport hunters has decreased from a high of 18 pheasants harvested per 100 ha in 1972 to 4 per 100 ha in 2000 (Draycott et al. 2002).

As agricultural intensification has increased habitat fragmentation has also increased.

This can have a negative impact upon pheasant populations (Haensly et al. 1987, Schmitz and

Clark 1999), and influence the size of home ranges (Whiteside and Guthery 1983). Some studies have shown that composition of habitat within home ranges does not affect survival (Perkins et al. 1997), but that some landscape features can have a negative affect upon survival (Schmitz and Clark 1999), and nest success (Haensly et al. 1987, Clark et al. 1999).

Pheasants have a territorial defense harem polygynous mating system. Research in the

United Kingdom has demonstrated that lack of male territories during the breeding season can influence dispersal rates and distances of hens (Robertson 1996). These territories are often established in open habitats along the edges of woodlands, hedge rows, and wetlands (Hill and

Robertson 1988, Robertson et al. 1993). Hens do not choose males based on nesting habitat within the territory (Ridley 1983, Göransson et al. 1990), but are influenced by the type of habitats within the male territory (Ridley 1983) where hens feed in order to build up fat reserves for nesting (Draycott et al. 2002).

The quality of nesting habitat can also influence productivity of pheasant populations and habitats that are regularly used for nesting include fallow areas, cereal grains, hay fields, pastures, row crops, residual cover, hedgerows, and woodlands (Robertson 1996). Management of nesting habitat can increase the probability of successfully hatching a clutch, but may not lead to overall population increases, and thus individual farm management in areas of poor regional management might increase the dispersal and distribution of pheasants rather than increase production (Burger 1988).

Despite the importance of pheasants as a game species throughout Austria, no information is currently available about habitat use by hens during the breeding season.

Therefore, I examined hen pheasants during the breeding season to determine home range and habitat preference, estimate nest success and habitat preference, and determine survival and the effects of habitat and landscape features upon survival.

STUDY AREA

Seefeld Estate is a 2,400 ha farm located in northeast Austria in the state of

Neiderösterich (Lower Austria) on the Czech Republic border, approximately 150 km north of

Vienna in the town of Seefeld-Kadolz (Figure 2.1). The estate has been farmed by the Hardegg

family since the 15th century and much of the property sits upon reclaimed marsh lands. It is a

linear property and runs about 9 km east to west and ranges from 0.5 km – 3 km south to north.

Land use on Seefeld Estate is dominated by annual crops which comprise 72% of the land area.

Winter wheat is the most common crop, and as farm management practices have improved,

yields have increased from 3 tons/ha in the 1960’s to 5 tons/ha in the 1990’s. Barley, sugar

beets, and rape are also important crops. Vineyards for wine production are common on the hills

north and east of the farm. The largest indoor pig production facility, in Austria, is also located

on the property. The remaining 28% of the estate is coppice, game crop, set aside, wetland

shrub, woodland, and vineyard (Table 2.1).

The Pulkau River runs through the estate and provides water for irrigation through a

series of ditches and center pivot irrigation. This small river was channelized from the 1950’s to

1990’s, but recent management has been to restore the meandering flow and adjacent wetlands.

Current management strategies focus on intensive predator control with an emphasis on red fox

(Vulpes vulpes), and crows and magpies (Corvid spp.). Supplemental feed, utilizing wheat grain in hoppers, is provided during the winter and used in the spring to enhance the quality of male breeding territories. The estate sits at an elevation of 190 m above sea level. Climate is mid- continental with mean annual precipitation of 480 mm per year, with approximately 160 mm falling in May and June. The temperature range in summer is 6 to 37 C and in winter is -25 to 5

C. The surrounding area is comprised of a series of small villages, wine production areas, and small family farms with an average field size of 14.6 ha. These farms cover 80% of the land area outside of villages (Molterer 1997).

METHODS

Hens were captured using walk in funnel traps during 1 March – 10 April in 2002 and

2003. Traps were 1 m x 0.5 m x 0.5 m with 2.8 cm nylon webbing; funnels were created using

2.3 cm chicken wire fencing. Captured hens were fitted with a 6g necklace collar (Holohil© model RI-2B) in 2002 and a 9.9g necklace collar (Holohil© model RI-2B) in 2003. Weight and

tarsus length were measured to determine a condition index (weight/tarsus length) for each hen

(Robertson et al. 1985). An aluminum patagial tag was fixed to the right wing for individual

identification. All pheasants were handled and tagged according to guidelines established in the

American Ornithologists’ Union Report of the Committee (American Ornithologists’ Union

1988) and the University of Georgia Institutional Animal Care and Use Committee (Permit

#A3437-01).

Age was established by measuring the shaft diameter of the 10th primary (1st U.S.) wing

feather at the base of the barbs and is 92% accurate in estimating age (Greenburg et al. 1972).

Feathers were allowed to dry for 24 hours before measuring. Hens with a shaft diameters greater than or equal to 2.8 mm were classified as adults (hatched prior to previous breeding season) and all others were considered yearlings (hatched during previous breeding season).

Radio-telemetry was conducted using a portable receiver (Telonics® model TR-2) and a

handheld yagi antenna to locate radio-tagged hens 3 times weekly. Radio-tagged hens were

observed from time of capture until 10 August, depredation, or signal loss in both 2002 and

2003. Hens were observed from > 15-30 m and slowly circled to ascertain exact location and

habitat. Locations were spot mapped and UTM coordinates were determined from 1:50,000

Bundesamt für Eich-und Vermessungswesen (BEV) (Federal Office of Metrology and

Surveying) topographical map.

Aerial color ortho-photographs of Seefeld Estate and the surrounding area were taken by fixed-winged aircraft by BEV at a scale of 1:15,000 in 1999. Photographs were projected in the

Gauss-Kruger datum and used Österreichischen Landesvermessung (MGI) coordinate system.

The ortho-photographs were scanned using a Microtek ScanMaker 9600XL scanner and saved as

GeoTiff images for import into ArcView Version 3.2. The GeoTiff images were digitized in

ArcView and each polygon was assigned a value based on habitat type. The habitat map of

Seefeld was then re-projected into WGS 84 datum with UTM zone 33 coordinates using

ShapeWarp 2.2 ArcView Extension (McVay 1998) which had a root mean square error (RMSE) of 5.6 m. Xtools Extension (Delaune 2003) in ArcView was used to calculate the total hectares of the study area and for each habitat.

The minimum convex polygon (MCP) (Mohr 1947) and the 95% adaptive kernel

(Worton 1989) were used to estimate home ranges using Animal Movement Extension Version

2.04 (Hooge and Eichenlaub 2000). Bootstrapping with replacement was conducted using

Animal Movement Extension Version 2.04 (Hooge and Eichenlaub 2000) to determine minimum number of locations needed to estimate the MCP home range. Bootstrapping results suggested that >20 locations were sufficient for estimating MCP home ranges. A minimum of 20 locations was also used to estimate 95% home range (Kenward 2001). Home ranges include all locations observed from time of capture except for locations recorded while incubating or accompanied by a brood.

I compared home ranges between year and age classes using analysis of variance

(ANOVA) for unequal sample size (Sokal and Rohlf 1969) using PROC GLM (SAS Institute

1999).

I used compositional analysis (Aebischer et al. 1993) to determine habitat preference at

the 2nd and 3rd order (Johnson 1980) for both the MCP and 95% adaptive kernel home ranges using BYCOMP.SAS (Ott and Hovey 1997). Habitat proportions within home ranges were determined using Geoprocessing Wizard and XTools in ArcView. I calculated the amount of edge within the MCP home range using Patch Analyst 3.1 (Rempel and Carr 2003). Significance of the Wilks’ λ was calculated using randomization on 1,000 runs of the data. I then ranked habitat preferences by a series of paired t-tests.

I combined all the habitats available on the study area into 4 functional units: 1) agriculture, including all row crops and vineyards, 2) set aside, including planted grasslands and game crops, 3) wetland shrub, and 4) woodland, including wooded areas, coppice, and windbreaks. Pheasants captured in 2002 and 2003 were combined to have >10 hens per habitat

as suggested by Aebischer et al. (1993). Missing values at 2nd and 3rd order analysis were

replaced following guidelines established by Aebischer et al. (1993).

Weekly survival during 1 March through 10 August was calculated by the Kaplan-Meier method (Kaplan and Meier 1958) with staggered entry (Pollock et al. 1989) using known fate model (S[t]) in Program Mark (White and Burnham 1999) on the logit scale for both 2002 and

2003. The (S[t]) model was chosen a priori to ascertain which weeks during the breeding season had an affect upon survival. Hens that died or were lost within 2 weeks of capture were removed from the study.

Covariates were chosen to estimate their influence upon survival. Non-habitat variables

considered were age, condition index, and dispersal. Habitat variables included amount of edge

(m/ha) within home range, and proportion of agricultural, game crop, set aside, woodland, and

wetland habitats within the 95% adaptive kernel home range. Adaptive kernel was selected over

MCP because the kernel estimates the probability that a particular habitat was used and as few as

10 locations can be used (Kenward 2001). To determine the effects of covariates upon survival

two models, survival constant (S[.]) and group (S[g]), (groups representing 2002 and 2003), were

chosen a priori because covariates were not measured over time. To evaluate the importance of

covariates, Akaike’s Information Criterion (AICc) was used to determine which models fit best

(Anderson et al. 2001). A slope was determined for each covariate by model averaging along

with the unconditional standard error (SE) and 95% confidence interval (CI) (Anderson et al.

2001).

Locations, from time of capture until nesting or last location if nesting was not observed,

were used to determine if dispersal occurred using RANGESV (Kenward and Hodder 1996). A

stochastic detector of 250 m (Schmitz and Clark 1999) and a probability level of 5% were

selected to reduce the chance of dispersal being detected prematurely or erroneously as

recommended by Kenward and Hodder (1996). If dispersal was detected, the arithmetic mean of

locations before and after the dispersal date was calculated to estimate distance of dispersal

(White and Garrott 1990). The mean date of dispersal was also calculated. Dispersal between

age and year was compared using ANOVA in Proc GLM (SAS Institute 1999) for uneven

sample size to test if dispersal was equal between age and year.

If dispersal was not detected the arithmetic mean of locations before and after the

calculated mean date of dispersal was used to determine the average distance moved from

wintering area to breeding location.

Nest success was estimated using the Mayfield method (Mayfield 1961) for initial, 2nd, and 3rd nests, and for initial nests located in each habitat. Nest habitat preference was estimated

by χ2 analysis (SAS Institute 1999) to test if nesting habitat selection was random. If χ2 was

significant, Bonferroni Z-test was used to test avoidance or preference for each habitat. The

standardized index (Manley et al. 1993) was used to estimate nest habitat preference if all

habitats were equally available.

I measured density of vegetation at each nest site using the visual obstruction method (R.

Draycott, pers. comm.) in which the nest is viewed from 1 m directly above the nest and from the

side at 0.5 m at kneeling height. Visual obstruction of the nest was then ranked on a scale of 1 to

4, with 4 representing 75-100% visual obscuration. Mann-Whitney U-test for ranked data (SAS

Institute 1999) was used to determine if visual obstruction at successful and unsuccessful nests

were different.

RESULTS

Ninety-one pheasant hens were radio-tagged during 2002 – 2003 and were monitored

from 1 March – 10 August each year. The age ratio was evenly distributed with 44 yearlings and

46 adults (Figure 2.2). Mean weight of captured hens was 1014.8 g (+ 12.5 SE) with a mean

condition index of 16.9 (+ 0.18 SE). In 2002, hens (n = 43) had a mean weight of 1049.5 g (+

18.3 SE) and condition index of 17.4 (+ 0.25 SE) and 48 hens captured in 2003 had an average

weight of 983.8 g (+ 16.1 SE) and a condition index of 16.4 (+ 0.24 SE). The condition of hens was significantly different between year (F1,89 = 67.16, P = 0.0089) and age (F1,89 = 9.48,

P=0.0028) (Table 2.2).

Minimum convex polygon home range size averaged 56.05 ha (+ 6.50 SE, n = 62) and

had mean edge density of 134.71 m/ha (+ 5.02 SE). Home range for yearlings averaged 61.39 ha

(+ 8.26 SE, n = 27) and 51.92 ha (+ 9.63 SE, n = 35) for adults, and these were not different

(F1,60 = 0.52 ,P = 0.47). I found a difference between years (F1, 60 = 9.34, P = 0.003) with an

average of 78.70 ha (+ 12.85 SE, n = 25) and 40.74 ha (+ 4.98 SE, n = 37), during 2002 and

2003, respectively (Table 2.3). Adaptive kernel home range size averaged 63.17 ha (+ 9.05 SE,

n = 62).

nd Habitat use was not random at either the 2 (Wilks’ λ = 0.65, F3,59 = 10.38, P < 0.0001)

rd or 3 (Wilks’ λ = 0.19, F3,59 = 80.81, P < 0.0001) order using MCP. Adaptive kernel results

nd rd were similar for both 2 (Wilks’ λ 0.4686, F3,57 = 21.54, P < 0.0001) and 3 order (Wilks’ λ =

0.2180, F3,57 = 68.18, P < 0.0001) results (Figure 2.3).

Habitat rankings for MCP at the 2nd order revealed that set aside habitat ranked highest

and woodland habitat ranked lowest (Table 2.4.a). Third order rankings stated wetland habitat ranked highest while agricultural land ranked lowest (Table 2.4.b). Ranking of habitats for adaptive kernel at the 2nd and 3rd order were identical to results for the MCP.

Kaplan Meier survival estimates were 63.3% (+ 8.7% SE, n = 33) and 60.5% (+ 8.25%

SE, n = 40) during 2002 and 2003, respectively. The majority of hen loss occurred during 5

April – 5 July during both years (Figure 2.4). All covariates tested had confidence intervals that

included zero suggesting that none of the covariates had an effect on survival (Table 2.5).

I detected dispersal for 41.5% (32/77) of hens; 45.2% of yearlings, and 41.9% of adults.

Hens that dispersed moved approximately 1447.32 m (+ 144.94 SE) from their wintering

locations to nesting area. Mean dispersal date was 11 April (+ 2.6 days SE). Adults that dispersed moved an average of 1418.23 m (+ 218.65 SE, n = 18) and yearlings averaged 1484.73

m (+ 184.22 SE, n = 14). No difference was detected (F1,30 = 0.05, P = 0.82). Mean dispersal in

2002 was 1853.18 m (+ 226.74 SE, n = 16) compared to 1041.17 m (+ 116.02 SE, n = 16) in

2003 and these were statistically different (F1,30 = 10.16, P = 0.003). Hens that did not disperse moved an average 200.10 m (+ 27.90 SE) from their wintering area to their breeding areas (Table

2.6).

Fifty-one initial nests were observed in 2002 and 2003 with a mean initiation date of 16

May (+ 1.7 SE) and a hatch success rate of 48.7% (+ 6.59 SE). Initial nests were located in 4

habitats; 23 in agricultural fields with a success rate of 54.5% (+ 11.08 SE), 19 in set aside with a

51.1% (+ 11.30 SE) success rate, 5 in wetland with a success rate of 26.4% (+ 11.31 SE), and 4

in woodland with a success rate of 34.9% (+ 16.25 SE) (Table 2.7).

Twenty-eight initial nests were unsuccessful. The primary cause of nest loss was

predation by mammalian and avian predators which were responsible for 75.0% (21/28) of nests

lost. The remaining nests were loss to abandonments caused by natural or human disturbance.

Nest scores were obtained from 22 successful and 22 unsuccessful nests. Visual obstruction at successful nests had a mean top down score of 2.8 (+ 0.2 SE) and a side view score of 3.3 (+ 0.1

SE), while unsuccessful nests had an average top down score of 2.4 (+ 0.3 SE) and a side view

score of 2.9 (+ 0.2 SE) and I found no difference between successful and unsuccessful nests

(Mann-Whitney U, Z-value 1.033, P = 0.301). Nests in each habitat were not compared to

random locations.

Habitat selection for nesting was not random (χ2 = 386.4, 3 df, P < 0.0001). Among

available habitats, I found that use of agriculture and set aside were different with set aside being

used more than availability and agriculture less then availability (Figure 2.5).

The standardized index revealed that if all habitats were equally available then set aside

would contain 61.2% of the nests, wetland shrub 25.5%, woodland 11.7%, and agriculture 1.6%.

Nineteen 2nd nests were observed in 2002 and 2003 and had a success rate of 31.2% (+ 6.7 SE).

Four 3rd nests were found in 2003 and all were unsuccessful.

DISCUSSION

Home range sizes of hen pheasants on Seefeld Estate were smaller than those studied in

Wisconsin and Maryland (56 ha compared with 146 and 235 ha) (Gates and Hale 1974, Smith et al. 1999), but were comparable to the 90 day range (May-July) found in Iowa of 47 ha (Schmitz and Clark 1999). Home ranges observed in 2002 were significantly greater than in 2003 and could be related to an unusually hot and dry summer. Dispersal was detected in 41.5% of tagged hens and distance traveled is similar to other studies, but contrary to other studies that observed greater movement of yearlings than adults, our findings suggested that home range and dispersal between yearlings and adults were not different (Gates and Hale 1974).

My analysis of habitat showed that at the 2nd order (comparing habitat proportions within home ranges to the study area) set aside habitat ranked highest. This corroborates other studies which show a preference of grassland areas within home ranges (Dumke and Pils 1973, Warner et al. 1987, Schmitz and Clark 1999). Whiteside and Guthery (1983) also found that preferred

habitats in home ranges resembled nesting habitat. Third order analysis (comparing proportion

of radio-locations in each habitat to proportions within the home range) revealed that wetland

shrub was heavily used and could indicate use of this habitat for escape and loafing cover. At

Seefeld, many males used the tall reeds (Phragmites communis) at the edge of wetlands for

breeding territories which would also attract females. It is also notable that wetland areas are

also preferred during winter months to escape inclement weather which aided in high winter

survival at Seefeld Estate (Anderson 2002).

Breeding season survival is similar to other studies (Riley and Schulz 2001). Proportions

of habitats within home ranges were not good measures to use to estimate survival and

corroborate the results of Perkins et al. (1997), but suggest that survival could be better in

agricultural lands than set aside. This could be an effect of patch size rather than habitat cover as

studies in Iowa showed that larger blocks have less edge and therefore are used less by predators

(Schmitz and Clark 1999), but no relationship between edge and survival was found at Seefeld

Estate. Although predator removal on Seefeld is quite intensive, the lack of predator control off

the estate combined with the high edge to area ratio of the estate suggests that re-colonization of predators occurs quickly from surrounding areas (Riley and Schulz 2001).

Nest initiation rate was 100% of radio-tagged hens which is greater than estimates in southern England (Hoodless et al. 1999). The high rate suggests that condition of hens entering the breeding season is good and that supplemental feeding during the winter and spring is having a positive affect. Draycott (et al. 1998) demonstrated that fat reserves on pheasants were maintained with supplemental feeding during the winter and spring. It might also be related to the lack of rearing and releasing of penned birds on Seefeld, which is commonly done in the

U.K. Nest success is 48.7% and is similar to studies which range from 19 – 60% in North

America (Giudice and Ratti 2001) and in Europe (Hoodless et al. 1999). One exception is

Mallard Island, North Dakota where nest success was 90%. This was an area where predator control was undertaken (Carroll and Sayler 1990).

Set aside is preferred nesting habitat at Seefeld Estate. Although more nests were found in agricultural crops, mainly winter wheat, set aside was strongly selected in proportion to its availability. Set aside plots that are properly managed allow for residual cover to remain and make it attractive for nesting. Boyd and Richmond (1980) observed that first nests were located in old-field habitats and Snyder (1984) found that pheasants prefer areas with tall, dense herbaceous cover whether it is new growth or residual cover. Carroll and Sayler (1990) observed that nests in grass habitat had more cover and were preferred over cropland. Winter wheat is not

preferred nesting habitat (Olsen 1977), but can provide suitable nesting habitat in years when

environmental conditions are right for growth and early structure is available when nesting

begins (Snyder 1984).

My research showed that management of wild pheasants on Seefeld Estate has produced

conditions that promote high survival and productivity of wild pheasants. High winter survival

(Anderson 2002) and supplemental feeding allow pheasants to enter the breeding season in

excellent condition. Home ranges were smaller than other studies and are centered around set

aside habitat which is also preferred nesting habitat. Dispersal distances were not extreme

suggesting that individual hen pheasants were not avoiding habitats on the estate or high

population densities. Nest success is similar to many other studies and could be enhanced by

increasing the availability of set aside throughout the area. Breeding season survival is good and

is enhanced by the use of wetland areas as escape cover and predator control. These breeding

season management practices enable populations to increase the chances of hatching a nest due

to high initiation rates and allow hens the ability to survive to the fall with a good chance of surviving until the following breeding season.

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Table 2.1. Description of habitats found on Seefeld Estate and surrounding areas in Lower

Austria.

Habitat Description Agriculture Arable lands dominated by cereal grains (i.e. winter wheat and (row crops) barley), but also includes lucerne (alfalfa), maize (corn), Austrian pea, rape, and sugar beet.

Coppice Areas of hardwood forest that are cut and allowed to stump sprout. Wood here is used mainly for firewood or posts for the vineyards. Most common species is false-acacia (Robinia pseudacacia)

Game crop Arable land planted in mixtures of cereals, hemp, lucerne (alfalfa), native forbs, and sunflowers to provide cover for wildlife.

Set aside Arable land that is designated as long-term (multiple years) or rotational (single year) set aside that are planted in cocksfoot (orchard) grass (Dactylis glomerata) or native species and are partially subsidized by the European Union.

Vineyard Arable lands, usually on hill sides, planted in grape vines and sometimes intermixed with other row crops.

Wetland Land that is adjacent to permanent of seasonal water which includes irrigation ditches and the bank of the Pulkau River, usually dominated by reed grass (Phragmites communis)

Woodland Mature forest, dominated by hardwoods that are thinned for timber, fuel, and for hunting purposes.

Urban Areas that include villages, farm buildings, storage facilities, wine cellars, and road ways.

Table 2.2. Weight (+ SE) and condition index (+ SE) of juvenile and adult radio-tagged

pheasant hens captured from 1 March – 10 April at Seefeld Estate, Lower Austria, Austria, 2002-

2003.

_____Weight______Condition Index__ n Mean SE Mean SE 2002 a 43 1049.5 18.3 17.4 0.25 Yearling 21 1013.8 27.9 16.9 0.39 Adult b 21 1096.2 18.85 17.95 0.27

2003 a 48 983.8 16.1 16.4 0.24 Yearling b 22 945.5 25.1 15.85 0.34 Adult 26 1016.2 18.9 16.9 0.30

All Yearlings c 43 978.8 19.0 16.3 0.26 All Adults c 47 1051.9 14.5 17.4 0.22

Categories with the same letter are significantly different at alpha = 0.05.

Table 2.3. Mean minimum convex polygon and adaptive kernel home ranges (ha) of radio-

tagged pheasant hens during 1 March – 10 August by year and age at Seefeld Estate, Lower

Austria, Austria, 2002-2003.

____100% MCP______95% Kernel____ n Mean SE Mean SE Year 2002a 25 78.70 12.85 95.93 18.72 2003a 37 40.74 4.98 41.03 6.39 Age Yearling 27 61.39 8.26 82.46 17.34 Adult 35 51.92 9.63 48.28 8.23

All home ranges 62 56.05 6.50 66.17 9.05

Categories with the same letter are significantly different at alpha = 0.05.

Table 2.4a,b. Habitat ranking matrix of 4 defined habitat types based upon 2nd order (A) and 3rd order (B) compositional analysis. Higher ranking indicates greater use compared to availability.

Within the matrix, (+) signifies that the row habitat is preferred over the column habitat, whereas a (-) signifies the opposite. Significant difference between habitats (P < 0.05) is indicated by

(+++) or (---).

Habitat Woodland Set aside Wetland Agriculture Rank Woodland . ------0 Set aside +++ . + +++ 3 Wetland +++ - . +++ 2 Agriculture +++ ------. 1

Habitat Woodland Set aside Wetland Agriculture Rank Woodland . +++ - +++ 2 Set aside --- . --- +++ 1 Wetland + +++ . +++ 3 Agriculture ------. 0

Table 2.5. AICc value, delta AICc, slope (β), and 95% CI of non-habitat and habitat covariates upon survival of radio-tagged pheasant hens at Seefeld Estate, Lower Austria, Austria, 2002-

2003. Inclusion of zero in the 95% CI means there is no significant slope.

AICc Delta AICc Slope 95% CI Pheasant covariates Dispersal 246.26 0.00 0.602 -0.333 1.537 Condition index 246.60 0.34 -0.325 -0.970 0.319 Age 247.43 1.17 -0.078 -0.897 0.741 Habitat covariates Agriculture (%) 245.65 0.00 1.236 -0.542 3.014 Set aside (%) 245.78 0.13 -1.512 -3.669 0.644 Woodland (%) 246.68 1.03 -1.972 -6.098 2.155 Edge (m/ha) 247.20 1.55 -0.144 -0.645 0.357 Game crop (%) 247.46 1.81 -0.714 -5.868 4.440 Wetland (%) 247.46 1.81 0.624 -6.779 8.026

Table 2.6. Distance (m) from center of winter locations to center of breeding locations for radio- tagged pheasant hens that dispersal was detected and for hens that no dispersal was detected at

Seefeld Estate, Lower Austria, Austria, 2002-2003.

n Mean SE Dispersal detected Yearling 14 1484.7 184.22 Adult 18 1418.2 218.65 2002 16 1853.2 226.74 2003 16 1041.2 116.02 Both years combined 32 1447.3 144.94 No dispersal detected Yearling 19 225.0 58.75 Adult 25 182.7 23.81 2002 21 241.7 52.08 2003 24 163.7 24.59 Both years combined 45 200.1 27.90

Table 2.7. Mayfield estimates of initial nest success (%) by radio-tagged pheasant hens for each habitat and for all habitats combined at Seefeld Estate, Lower Austria, Austria, 2002-2003.

Habitat Number of nests Mayfield estimate SE Agriculture 23 54.5 11.08 Set aside 19 51.1 11.30 Wetland 5 26.4 11.31 Woodland 4 34.9 16.25

All habitats combined 51 48.7 6.59

Figure 2.1. Map of Austria (Magellan Geographix 2003) and habitat map of Seefeld Estate and surrounding area, Lower Austria, Austria.

20 Juvenile Adult

15 er of hens b 10 Num

0 2.1 2.2 2.3 2.4 2.5 2.6 2.7 2.8 2.9 3.0 3.1

Wing shaft diameter (mm) `

Figure 2.2. Age distribution of radio-tagged hen pheasants (n = 90) during spring (1 March – 10

April) on Seefeld Estate, Lower Austria, Austria 2002 – 2003. The line denotes separation of juvenile and adult hens based upon 10th primary shaft diameter.

Woodland

Wetland Study area Home range Radio locations Set aside

Agriculture

0 20406080100

Proportion

Figure 2.3. Habitat proportions within the study area, 100% MCP home range (+SE), and radio locations (+SE) of 63 radio-tagged hens from 1 March – 10 August, Seefeld Estate, Lower

Austria, 2002 – 2003.

1.0

0.9 al

iv 0.8 rv Su 0.7

0.6

0.5 Mar Apr May Jun Jul Aug

Date B.

1.0

0.9

ival 0.8

rv Su 0.7

0.6

0.5 Mar Apr May Jun Jul Aug Date

Figure 2.4. Kaplan-Meier survival estimates (+SE), modified for staggered entry, of radio- tagged pheasant hens from 1 March – 10 August in (A) 2002 (n = 33) and (B) 2003 (n = 40) at

Seefeld Estate, Lower Austria, Austria.

Woodland

Wetland Available Nesting

Set aside

Agriculture

0 20406080100

Proportion

Figure 2.5. Percent of available habitat vs. nesting habitat (+95% CI) by 51 hen pheasants on

Seefeld Estate, Lower Austria, 2002 – 2003. Note CIs that do not overlap with availability indicated significantly more or less use of that habitat for nesting.

CHAPTER 3

HOME RANGE, HABITAT USE, AND SURVIVAL OF PHEASANT BROODS IN

LOWER AUSTRIA

INTRODUCTION

Studies of population dynamics and abundance of common or ring-necked pheasants

(Phasianus colchicus) have suggested that nesting and brood habitat is critical to recruitment and could be limiting pheasant populations in Europe and the U.S. (Chiverton 1994, Warner et al.

1999), and that brood survival is one of the most important and least understood components of pheasant life history (Warner et al. 1984, Hill and Robertson 1988). Although pheasants are widely distributed across Europe, wild populations have declined during the past half-century

(Hill and Robertson 1988, Tapper 1999, Csànyi 2000), especially as farming has shifted to row crop agriculture and adopted more intensive farming practices (Jarvis and Simpson 1978, Hill

1985, Potts 1991).

The first 14 days after broods hatch is considered to be the most critical age class relative to variation in survival (Hill 1985, Meyers et al. 1988, Riley et al. 1998). Increased use of pesticides, shifts in crops grown and changes in field management have reduced availability of weedy plants and insects important for chick growth and development (Potts 1980, Hill 1985,

Sotherton et al. 1985, Rands 1986, Sotherton and Robertson 1990). Studies have suggested that broods prefer home ranges consisting of weedy areas and grassland (Hill 1985), and that home range size is negatively correlated with insect abundance and survival of pheasant (Hill 1985), gray partridge (Perdix perdix), and red-legged partridge (Alectoris rufa) broods (Green 1984).

The quality of nesting habitat can also influence chick recruitment because broods that hatch earlier are generally larger and have a greater chance of survival (Riley et al. 1998). The loss of fallow areas, residual cover, hedgerows, and woodlands for nesting (Robertson 1996) and its separation from brooding areas can increase the area that must be covered by broods in order to find food (Warner 1984). Increased movements increase the chance that more edge will be encountered which can negatively affect the survival of pheasant hens due to increase use by predators (Schmitz and Clark 1999). Also, the condition of hens which is improved by supplemental feeding during winter and spring (Draycott et al. 1998) could affect the hatch weight of chicks which is known to have an effect upon survival (Riley et al. 1998)

Lower Austria, Austria, like the rest of Europe, has also seen a precipice decline in harvest rates of wild pheasants over the past 30 years (Draycott et al. 2002). Currently, little information is available on pheasant populations and specifically brood habitat use and survival in Austria. Given that farming practices in Austria are different than in Great Britain and North

America where most pheasant research has been conducted, I examined pheasant broods to determine home range size, habitat preference, survival, and the affect of habitat and landscape features upon survival.

STUDY AREA

Seefeld Estate is a 2,400 ha farm that has been farmed by the Hardegg family since the

15th century. The estate is located in the state of Neiderösterich (Lower Austria), Austria and is about 150 km northeast of Vienna in the town of Seefeld-Kadolz on the border of the Czech

Republic (Figure 3.1). Seefeld Estate covers approximately 9 km from east to west and ranges from 0.5 km – 3 km north to south. A majority of the estate property sits upon reclaimed marsh lands of which 72% of the area is planted in annual crops. The dominant crop is winter wheat

and currently yields an average of 5 tons/ha. Other important crops are summer and winter barley, sugar beets, and oil seed rape. Wine production facilities are also on the property and vineyards are found on the hills north and east of the farm. The largest indoor pig production facility in Austria is also located on the property. The remaining 28% of the estate is coppice, game crop, set aside, wetland shrub, woodland, and vineyard (Table 3.1).

Center pivot irrigation is made possible by water from the Pulkau River, which runs the length of the estate, and is distributed by a series of open ditches. Wetland areas are being established along the river by restoring the meandering flow of the Pulkau, which was channelized during the 1950’s. Wheat is provided to pheasants as supplemental feed by a series of grain hoppers spaced throughout the estate to provide food during winter and improve the quality of male breeding territories in spring. The main predators on the estate are the red fox

(Vulpes vulpes) and crows and magpies (Corvid spp.); which are intensively controlled throughout the year. Seefeld estate has a mid-continental climate and an elevation of 190 m above sea level. Approximately 480 mm of precipitation fall per year, with May and June receiving approximately 160 mm. Temperatures range from 6 to 37 C in summer to -25 to 5 C in winter. Small villages, private vineyards, and family farms surround the estate. Family farms cover approximately 80% of the land outside the villages and have an average field size of 14.6 ha (Molterer 1997).

METHODS

Pheasant hens were captured from 1 March – 10 April in 2002 and 2003 using walk in funnel traps. Funnel traps were constructed using 1 m x 0.5 m x 0.5 m wooden frames with 2.8 cm nylon webbing; funnels were created using 2.3 cm chicken wire fencing. Hens were fitted with a 6 g necklace collar (Holohil© model RI-2B) in 2002 and a 9.9 g necklace collar (Holohil©

model RI-2B) in 2003. The condition of each hen was estimated by the condition index

(Robertson et al. 1985) by calculating a ratio of weight and tarsus length (Robertson et al. 1985).

Aluminum patagial tag was attached to one wing for individual identification. Captured pheasants were handled and tagged according to guidelines established in the American

Ornithologists’ Union Report of the Committee (American Ornithologists’ Union 1988) and the

University of Georgia Institutional Animal Care and Use Committee (Permit #A3437-01).

Thirty-five hens were captured in the spring of 2001 on Seefeld Estate, using the same techniques, in conjunction with another study (Anderson 2002). Data collected on broods were incorporated to increase sample size when estimating brood home range, habitat preference, and survival.

Hens were located 3 times per week using a portable receiver (Telonics© model TR-2) and a handheld yagi antenna. Once a hen had nested the nest site was located and monitored every other day. If a hen was located off the nest, I walked into the site to determine clutch size or nest fate. As the projected hatch date approached, nests were monitored daily until hatch.

Once the nest hatched it was examined to determine the number of chicks that hatched. I estimated hatch success by dividing the total number of eggs that hatched by the total number of eggs laid in successful nest. Broods were located twice daily for the first 21 days. I determined exact location and habitat by slowly circling the birds from a distance of >15-30 m. Since chicks were not tagged, a hen was considered to have lost a brood if a brood caution call or brood gathering call, as summarized by Guidice and Ratti (2001) were not heard during consecutive observations or if the hen died.

Aerial color ortho-photographs of Seefeld Estate and the surrounding area were taken at a scale of 1:15,000 in 1999. The photographs were scanned using a Microtek ScanMaker 9600XL

scanner and saved as GeoTiff images for import into ArcView Version 3.2. Once the images were imported into ArcView they were digitized and each polygon was assigned a value based on habitat type. The habitat map of Seefeld was then re-projected using ShapeWarp 2.2

ArcView Extension (McVay 1998) into WGS 84 datum with UTM zone 33 coordinates and had a root mean square error (RMSE) of 5.6 m. Proportions of habitats within the study area were calculated using Xtools Extension (Delaune 2003) in ArcView.

The 100% minimum convex polygon (MCP) (Mohr 1947) and 95% adaptive kernel

(Worton 1989) were constructed to estimate brood home range using locations observed during the first 21 days after hatch in Animal Movement Extension Version 2.04 (Hooge and

Eichenlaub 2000) in ArcView 3.2. Bootstrapping with replacement was performed in Animal

Movement Extension Version 2.04 (Hooge and Eichenlaub 2000) to estimate the minimum number of locations needed to construct MCP home ranges. Results from bootstrapping indicated that >20 locations were necessary to estimate MCP home ranges. Adaptive kernel home ranges were also constructed using a minimum of 20 locations (Kenward 2001). Once brood home ranges were calculated, they were separated by year and age and compared using analysis of variance (ANOVA) for unequal sample size (Sokal and Rohlf 1969) using PROC

GLM (SAS Institute 1999) to determine if home ranges differed between years or age.

In the United Kingdom, it has been suggested that hens nest on the edge of their breeding home range and then raise the brood back toward the interior of their breeding range (R.

Draycott, pers. comm.). To examine if brood home ranges are incorporated into the hens home range before nesting, the home range of hens with broods was constructed using locations from time of capture until nesting. The amount of home range overlap was calculated according to

criteria established by Mizutani and Jewell (1998). Distance from nest site to arithmetic mean of brood home range was calculated to estimate brood movement.

To determine habitat proportions within each home range I used Geoprocessing Wizard and Xtools extension in ArcView. The amount of edge within each MCP home range was calculated using Patch Analyst 3.1 (Rempel and Carr 2003). I then used compositional analysis

(Aebischer et al. 1993) to estimate habitat preference at the 2nd and 3rd order (Johnson 1980) for both the MCP and 95% adaptive kernel home ranges using BYCOMP.SAS (Ott and Hovey

1997). Wilks’ λ was calculated by running 1,000 iterations of the data to determine if habitat use was not random. Habitat preferences were then ranked by a series of paired t-tests.

To control bias habitats were combined into 4 categories as recommended by Aebischer et al. (1993); 1) Agriculture, includes all row crops and vineyards, 2) set aside, which includes planted grasslands and game crop, 3) wetland shrub, and 4) woodland, includes wooded areas, coppice, and wind breaks. Missing values at the 2nd and 3rd order were replaced according the criteria established by Aebischer et al. (1993).

I estimated brood survival during the first 21 days after hatching using Kaplan-Meier method (Kaplan and Meier 1958) via the known fate model in Program Mark (White and

Burnham 1999) on the logit scale for years 2001, 2002, and 2003. Since broods were left censored to have the same start time, survival constant (S[.]) was used.

Covariates were then used within the model to determine their affect upon brood survival.

Non-habitat variables include condition of hen at time of capture and age. Landscape variables include nesting habitat and proportion of agricultural land, game crop, set aside, wetland, woodland, and amount of edge (m/ha) within each home range. Habitat proportions for survival analysis were calculated using the 95% adaptive kernel since it estimates the probability that a

habitat was used and as few as 10 locations can be used (Kenward 2001). To estimate habitat proportions for broods with less than 10 locations the arithmetic mean was calculated and then buffered by the average kernel home range. To ascertain the effect of covariates on survival constant (S[.]) and by group (S[g]) were chosen a priori since broods were left censored and covariates were not measured over time. Akaike’s Information Criterion for small sample size

(AICc) was used to determine which models fit best and to assess the effect of each covariate upon survival (Anderson et al. 2001). A slope (β), unconditional standard error (SE), and 95% confidence interval (CI) for each covariate was calculated by model averaging. Covariates whose CI include zero are determined not to have an influence upon survival.

Broods were considered successful if the hen was observed with > 1 chick 21 days post hatch when flush counts were conducted to approximate the number of chicks that survived.

RESULTS

In 2002 and 2003, first nests were the most productive resulting in 85% (23/27) of the broods, which had a mean clutch size of 10.3 eggs (+ 0.46 SE, n = 42) and a hatch date of 12

June (+ 2.3 SE). Re-nesting attempts were responsible for 15% (4/27) of broods which had a clutch size of 8.4 eggs (+ 0.85 SE, n = 14) and a hatch date of 28 June (+ 7.6 SE). Hatch success of initial nests was 82.4% with an average brood size of 8.8 (+ 0.65 SE) chicks. Re-nests had a hatch rate of 53.1% with only 6.5 (+ 1.85 SE) chicks per brood (Table 3.2).

Thirty-six broods were observed in the three years with 9 in 2001, 15 in 2002, and 12 in

2003. Twenty-eight broods survived to day 21 and had a mean MCP home range of 11.1 ha (+

2.13 SE) and a kernel estimate of 14.6 ha (+ 2.45 SE). Home range of successful broods was not different between years (F2,25 = 1.99, P = 0.16) or age of the hen (F1,26 = 0.02, P = 0.90) (Table

3.3). Mean distance from nest location to arithmetic mean of brood home range for broods in

2002 and 2003 was 217.96 m (+ 33.0 SE, n = 21) and 44.9% (+ 7.88% SE, range 0 – 100%) of brood home ranges overlap the breeding home range of the hen (Table 3.4).

Habitats on Seefeld were not available in equal proportions and my compositional analysis of MCP home ranges showed that habitat use was not random at the 2nd (Wilk’s λ =

rd 0.59, F3,30 = 6.92, P = 0.001) or 3 order (Wilk’s λ = 0.44, F3,30 = 12.66, P < 0.0001) (Figure

3.2). At the 2nd order agriculture ranked highest among the habitats considered (Table 3.5a).

Third order MCP compositional analysis indicated that set aside was most preferred (Table

3.5b). Adaptive kernel results at the 2nd order indicated that habitat use was not random (Wilk’s

λ = 0.6577, F3,30 = 5.20, P = 0.0052) and rankings were identical to MCP results (Table 3.6a).

Third order kernel analysis indicated that habitat use was not random (Wilk’s λ = 0.6372, F3,30 =

5.69, P = 0.0033) and habitat rankings indicated that wetland habitat was favored (Table 3.6b).

Survival analysis showed that brood success for the first 21 days was 74.4% (+ 15.6%

SE), 91.9% (+ 7.8% SE), and 65.7% (+ 13.8% SE) in 2001, 2002, and 2003, respectively (Figure

3.3). Flush counts suggested that chick survival was 38% with a mean of 3.4 chicks (+ 0.51 SE) per brood in 2002, but was higher at 62% survival with an average of 5.9 (+ 0.77 SE) chicks per brood in 2003. Seven broods were lost during the 3 years, with losses occurring between 2 – 17 days after hatching and mean brood loss occurring 11 days after hatch (+ 2 days SE). Foxes were responsible for the loss of 4 broods. The remaining 3 broods were lost to a mammalian predator, exposure, and to crop harvest, respectively. One brood was lost due to failure of the radio-tag while the hen was incubating and fate was not determined.

None of the non-habitat covariates considered had an affect upon survival (Table 3.7) and the only habitat covariate that had a positive effect upon brood survival was game crop (Table

3.7). Increasing amounts of set aside appeared to have a negative effect on chick survival (Table

3.7). All of the other habitat covariates measured had a 95% CI that included zero. Although not significant, the slope for woodland suggested a negative relationship between percent woodland and survival, whereas the slope for wetland suggested a positive relationship with brood survival (Table 3.7).

DISCUSSION

The results of my study suggest that nesting habitat, clutch size and hatch success are not limiting pheasants at Seefeld Estate. The clutch size of initial nests, 10.8 eggs, is similar to studies in England (range 10.7 - 11.6) (Hoodless et al. 1999) and the U.S. (range 10.1 – 13.0)

(Hammerstrom 1936, Knot et al. 1943, Gates and Hale 1975, Penrod et al. 1986, Clarke and

Bogenshutz 1999). The pattern of smaller clutch size in re-nests is analogous to observations in other studies (Gates and Hale 1975, Penrod et al. 1986, Carroll and Sayler 1990, Clarke and

Bogenshutz 1999). A hatch rate of 82.4% coupled with a mean hatch date of 12 June allowed initial brood size to average 8.8 chicks. This is comparable to populations in Iowa that had a hatch date of 15 June (Riley et al. 1998) and brood size is similar to studies in the U.S. which range from 7.8 to 10.8 (Linder et al. 1963, Gates and Hale 1975, Warner 1984). Work on red grouse (Lagopus lagopus) established if initial clutch size and hatch rate are sufficient, then either diet and or habitat are likely to limit recruitment (Hudson et al. 1994).

My home range estimates suggest that broods were near the upper range observed in the

U.S. of 2 – 11 ha (Kuck et al. 1970, Hanson and Progulske 1973, Warner 1979) and greater than

4.8 ha observed in England (Hill 1985). One possible reason for differences in home range values from England is because broods were followed for 21 instead of 14 days. Hanson and

Progulske (1973) demonstrated that the home range of broods increases as the chicks mature.

Compositional analysis at the 2nd order indicated that agricultural fields were incorporated into home ranges and corroborates previous studies which found that broods were often located in cereal crops (Warner 1979, Hill 1985, Enck 1986). The structure of cereal crops is attractive to broods since it provides cover from predators and allows for easy movement

(Aebischer and Blake 1994). Green (1984) observed that grey partridge broods were often found within the 5 m of the field edge where insects and weeds are more prevalent (Chiverton 1994).

Third order habitat analysis suggests that set aside habitat was preferred. Properly managed set aside has a greater diversity of plants which attracts invertebrates and produces small seeds which are necessary for chick development (Aebischer and Blake 1994) which complements previous research that located broods in weedy areas and undisturbed grasslands (Warner 1979,

Hill 1985, Riley et al. 1998). Compositional analysis also suggests that wetland habitat is an important component of brood home ranges indicating that it could be used as escape cover. I also determined that the home range of the hen during the breeding season does not seem to influence the brood home range. This is due to a shift in hen behavior from locating areas for breeding and nesting to finding habitats with little ground litter (Nelson et al. 1990) and an abundance of insects.

My findings demonstrated that brood loss occurred during the first 17 days after hatching and is consistent with results in England (Hill 1985) and the U.S. (Meyers et al. 1988, Riley et al. 1998). Principle loss of broods was due to predation by mammals, with minor loss attributed to exposure and harvest. Mammalian predation has been implicated in other studies (Riley et al.

1994, Riley et al. 1998) and the fox has been identified as the dominant predator upon pheasants in Iowa (Riley and Schulz 2001). Predator removal studies have shown that recruitment increases during these periods, but falls to pre-treatment levels once removal stops (Chesness et

al. 1968, Jensen 1970). Avian predators have been implicated in other studies (Carroll and

Sayler 1990), but were not observed during this study. Loss to exposure is hard to attribute to any one factor but weather, chick weight, and abundance of high protein insects can all affect the ability of broods to survive (Ryser and Morrison 1954, Hill 1985, Hudson et al. 1994, Riley et al.

1998).

I found brood survival during the first 21 days ranged from 62.3% - 91.1% and broods that incorporated game crop into their home range had 100% survival. This corroborates findings in Iowa and Illinois where survival has been correlated to the abundance of grassland

(Warner et al. 1984, Riley et al. 1998). In England, important arthropods are more abundant in set aside than in cereal crops (Sotherton et al. 1998) and also dictated the size of brood home ranges (Hill 1985). Also, game crops likely offer concealment from predators. My results also suggest that wetland areas positively influence chick survival, since they are not sprayed with pesticides and also provide cover once crops are harvested. My results corroborate findings by

Hill (1985) that woodlands have a negative effect upon brood survival. Woodland edge has been shown to influence habitat use of partridges (Dudzinski 1992) and pheasants (Wasilewski 1986) due to increased number of predators.

Average flush counts indicate that chick survival is similar to ranges observed in the U.S.

(Gates and Hale 1975, Warner et al. 1984, Carroll and Sayler 1990, Riley et al. 1998) and greater than the 32.5% reported by Hill (1985) in England. Given the inaccurate and bias results of flush counts (Riley et al. 1998) it is important to examine chick survival separately from brood survival since 91% of the broods observed in 2002 survived, but flush counts indicated that only

38% of chicks survived.

Although farming practices are different in Austria, my results suggest that chick recruitment can be enhanced through proper management of set aside and increasing the availability of game crop. It is important to understand the requirements that pheasants need for each period of their life history and how they relate to one another. Through proper management of all habitats, supplemental feeding, and predator control, broods will have an increased chance of survival and ability to adapt to poor weather conditions which cannot be controlled.

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Perdix perdix. Journal of Applied Ecology 33:865-978.

Warner, R.E., 1979. Use of cover by pheasant broods in east-central Illinois. Journal of

Wildlife Management 43:334-346.

_____, S.L. Etter, G.B. Joselyn, and J.A. Ellis. 1984. Declining survival of ring-necked pheasant

chicks in Illinois agricultural systems. Journal of Wildlife Management 48:82-88.

_____, 1984. Effects of changing agriculture on ring-necked pheasant brood movements in

Illinois. Journal of Wildlife Management 48:1014-1018.

_____, P.C. Mankin, L.M. David, and S.L. Etter. 1999. Declining survival of ring-necked

pheasant chicks in Illinois during the late 1900s. Journal of Wildlife Management

63:705-710.

Wasilewski, M. 1986. Population dynamics of pheasants near Rogów, central Poland. Ekologia

Polska 34:669-680.

White, G.C. and K.P. Burnham. 1999. Program MARK: Survival estimation from populations

of marked animals. Bird Study Supplement 46: 120-138.

Worton, B. J. (1989). Kernel methods for estimating the utilization distribution in home range

studies. Ecology 70: 164-168.

Table 3.1. Description of habitats found on Seefeld Estate and surrounding areas in Lower

Austria.

Coppice Areas of hardwood forest that are cut and allowed to stump sprout. Wood here is used mainly for firewood or posts for the vineyards. Most common species is false-acacia (Robinia pseudacacia)

Game crop Arable land planted in mixtures of cereals, hemp, lucerne (alfalfa), native forbs, and sunflowers to provide cover for wildlife.

Vineyard Arable lands, usually on hill sides, planted in grape vines and sometimes intermixed with other row crops.

Wetland Land that is adjacent to permanent of seasonal water which includes irrigation ditches and the bank of the Pulkau River, usually dominated by reed grass (Phragmites communis)

Woodland Mature forest, dominated by hardwoods that are thinned for timber, fuel, and for hunting purposes.

Urban Areas that include villages, farm buildings, storage facilities, wine cellars, and road ways.

Table 3.2. Clutch size, hatch date, and brood size of initial nests and re-nests of ring-necked pheasants at Seefeld Estate, Lower Austria, Austria during 2002 – 2003.

Mean Brood Size Clutch Mean Hatch Date at Hatch Initial nests 10.3 (n = 42) 12 June (n = 23) 8.8 (n = 23) 2002 9.7 (n = 21) 12 June (n = 12) 8.7 (n = 12) 2003 11.0 (n = 21) 12 June (n = 11) 8.9 (n = 11)

Re-nests 8.4 (n = 14) 28 June (n = 4) 6.5 (n = 4) 2002 8.1 (n = 8) 27 June (n = 3) 7.0 (n = 3) 2003 8.7 (n = 6) 30 June (n = 1) 5.0 (n = 1)

Table 3.3. Mean minimum convex polygon and adaptive kernel home ranges (ha) of radio- tagged pheasant broods during the first 21 days after hatch by year and age at Seefeld Estate,

Lower Austria, Austria during 2001 - 2003.

___100% MCP______95% Kernel___ n Mean SE Mean SE Year 2001 6 5.45 1.72 8.07 2.74 2002 12 15.62 4.09 19.88 4.72 2003 10 9.16 2.74 12.27 2.91 Hen age Yearling 12 10.82 3.24 14.31 3.37 Adult 16 11.37 2.92 14.87 3.58

All broods 28 11.13 2.13 14.63 2.46

Table 3.4. Percentage of brood home range (ha) located inside the breeding home range (ha) of ring-necked pheasant hens at Seefeld Estate, Lower Austria, Austria 2002 – 2003.

Hen Id Breeding HR (ha) Brood HR (ha) Brood HR overlap (ha) % overlap 1512 27.40 2.63 2.63 100.00% 1517 30.08 13.07 12.95 98.26% 1524 17.50 2.69 2.55 90.14% 1525 16.58 7.97 1.18 2.19% 1547 13.56 13.73 4.56 11.03% 1552 58.29 15.12 14.74 95.04% 1556 22.10 6.54 6.34 93.74% 1566 9.27 4.91 0.00 0.00% 1567 117.75 5.48 4.97 82.06% 1568 53.75 44.13 27.56 39.00% 1573 70.21 11.05 9.17 68.89% 1576 49.31 8.36 5.12 37.52% 1577 2.70 6.11 0.30 0.24% 1579 39.01 30.28 18.19 36.09% 1584 19.06 17.66 14.18 64.46% 1587 84.48 5.66 0.17 0.09% 1593 42.77 6.94 1.30 3.52% 1595 264.27 11.83 4.86 16.86% 1596 20.96 8.78 1.96 4.98% 1598 26.88 45.87 22.52 24.10% 1599 101.82 8.13 7.06 75.33%

Average 51.80 13.19 7.73 44.93% SE 11.76 2.49 1.59 7.88%

Table 3.5a,b. Habitat ranking matrix of 4 defined habitat types based upon 2nd order (A) and 3rd order (B) compositional analysis of MCP home ranges. Higher ranking indicates greater use compared to availability. Within the matrix, a (+) means that the row habitat is used relatively more than the column habitat, whereas a (-) means the opposite and a +++ or --- mean that they are different at (P < 0.05)

Habitat Woodland Set aside Wetland Agriculture Rank Woodland . ------0 Set aside +++ . + - 2 Wetland +++ - . - 1 Agriculture +++ + + . 3

Habitat Woodland Set aside Wetland Agriculture Rank Woodland . --- + + 2 Set aside +++ . +++ +++ 3 Wetland - --- . + 1 Agriculture - --- - . 0

Table 3.6a,b. Habitat ranking matrix of 4 defined habitat types based upon 2nd order (A) and 3rd order (B) compositional analysis of 95% adaptive kernel home ranges. Higher ranking indicates greater use compared to availability. Within the matrix, a (+) means that the row habitat is used relatively more than the column habitat, whereas a (-) means the opposite and a +++ or --- mean that they are different at (P < 0.05)

Habitat Woodland Set aside Wetland Agriculture Rank Woodland . ------0 Set aside +++ . + - 2 Wetland +++ - . - 1 Agriculture +++ + + . 3

Habitat Woodland Set aside Wetland Agriculture Rank Woodland . - - +++ 1 Set aside + . - +++ 2 Wetland + + . +++ 3 Agriculture ------. 0

Table 3.7. AICc value, delta AICc, slope (β), and 95% Confidence Interval (CI) of non-habitat and habitat covariates upon 21 day survival of radio-tagged pheasant broods at Seefeld Estate,

Lower Austria, Austria during 2001 - 2003. Inclusion of zero within the 95% CI means there is no significant slope.

AICc Delta AICc Slope 95% CI Pheasant hen covariates Condition index 80.247 0.000 0.187 -0.308 0.683 Age 80.289 0.042 0.382 -1.222 1.986 Habitat covariates Game crop (%) 74.516 0.000 609.037 472.284 745.790 Wetland (%) 75.735 1.219 36.578 -9.696 82.851 Edge (m/ha) 75.995 1.479 -0.347 -0.929 0.235 Nest location (in sa) 78.489 3.973 -1.221 -2.921 0.479 Woodland (%) 78.777 4.261 -14.586 -32.924 3.753 Agriculture (%) 80.558 6.042 -1.128 -5.265 3.009 Set aside (%) 80.801 6.285 -0.219 -3.604 3.167

Figure 3.1. Map of Austria (Magellan Geographix 2003) and habitat map of Seefeld Estate and surrounding area, Lower Austria, Austria.

Woodland

Wetland Study Area Home range Radio locations Set aside

Agriculture

0.00.20.40.60.81.0

Proportion

Figure 3.2. Habitat proportions within the study area, 100% MCP home range (+SE), and radio- locations (+SE) for 28 radio-tagged pheasants broods at Seefeld Estate, Lower Austria during

2002 – 2003.

1.0 0.9 al v 0.8 rvi

Su 0.7 0.6 0.5 5101520

Days after hatching B.

1.0 0.9 al

iv 0.8

Surv 0.7 0.6 0.5 5101520

Days after hatching

1.0 0.9

iv 0.8 rv

Su 0.7 0.6

0.5 5101520

Days after hatching

Figure 3.3. Kaplan-Meier survival estimates (+ SE) of radio-tagged pheasant broods during (A)

CHAPTER 4

MANAGEMENT IMPLICATIONS FOR RING-NECKED PHEASANTS

IN LOWER AUSTRIA

MANAGEMENT IMPLICATIONS

My results from Seefeld Estate demonstrate that careful management of set aside, game crop, and wetland habitat in conjunction with supplemental feeding and predator control is important for the survival of pheasant populations during spring and summer in Austria. Set aside habitat was used extensively by hen pheasants during the spring and summer for nesting, while game crop was important for brood survival. Wetland, which is also important during winter, is incorporated into the home ranges of hens and broods showing that it may also be important escape cover outside of winter.

I found that hens preferred home ranges that contained set aside and wetland habitats in proportions greater than their availability, and were most often found in set aside. The availability of set aside around wintering and breeding areas, which is used for nesting, reduced the need for females to travel long distances from wintering areas to breeding areas. No difference was detected between the dispersal distance of juveniles and adults which is contrary to other studies (Gates and Hale 1974). The proximity of necessary habitats allowed hens to have home ranges smaller than was observed in Maryland for the same time period (Smith et al.

1999), and supports studies which found that larger home range and increased edge (Schmitz and

Clark 1999), where predator traffic is higher (Warner and Joselyn 1986, Andren 1995), could negatively affect survival. Survival was constant during the 2 years (60.5 – 63.3%), and I found that hen losses coincided with peak nesting and brood rearing periods which corroborates

other studies that found depredation of hens to be greatest during incubation (Riley and Schulz

2001).

Set aside was the preferred nesting habitat since the residual cover provides structure and cover before cereal crops were sufficiently high. Although more nests were located in cereal fields, I found that if all habitats were equally available set aside would contain over 60% of nests. Presence of set aside allowed for nesting to peak by 15 May with a hatch date of 12 June.

This allows broods more than 3 weeks to develop before harvest begins in mid- to late-July. In

Iowa, nests that hatched earlier had heavier chicks and higher chick survival than nests that hatch later than the mean hatch date (Riley et al. 1998). Successful re-nests hatched 2 weeks after initial nests indicating that supplemental feeding at Seefeld Estate kept hens in good condition.

This allowed hens to re-nest quickly compared to observations in the United Kingdom where hens often had low fat reserves (Hoodless et al. 1999).

Home range estimates for broods were at the upper end of what had been reported in

England and the U.S.A. (Kuck et al. 1970, Hanson and Progulske 1973, Warner 1979, Hill 1985) and were primarily composed of crop land, but compositional analysis showed that set aside and game crop were preferred habitats. Broods that incorporated game crop in their home range had a 100% survival during the first 21 days. Properly managed game crop has low amounts of ground debris and low stem density which makes it easier for broods to move around and feed.

The mixtures of plant species found in game crop plantings provide cover and attract insects which are important to the diet of pheasant chicks (Hill 1985). It could be that the excellent brood cover found at Seefeld combined with warmer and drier weather reduced the feeding and chilling impacts on pheasant chicks observed in other regions. I also found that woodland

habitat could have negative effect upon brood survival. Therefore, I advocate the placement of game crop away from woodland edges to decrease the chances of predation.

My research indicated that predation was caused mainly by mammalian predators during nesting and brood rearing. Despite extensive and intensive predator control on the study area, it appears that predators are able to filter in from adjacent areas very quickly. Future research on nest predation, by video surveillance, needs to be conducted so that nest predator species can be targeted. Also more detailed research on chick survival needs to be conducted. Broods were considered successful if at least 1 chick was observed with the hen at the end of the first 21 days.

Therefore marking of chicks at time of hatch, as performed in Iowa (Riley et al. 1998), would enable detailed estimates of chick survival to be derived that can be used in development of a population model. Brood surveys need to be conducted yearly in the summer after harvest to determine brood size which can be used as an index to the health of the population (Riley and

Riley 1999).

Research conducted on Seefeld Estate could be implemented across Lower Austria to increase and stabilize pheasant populations. Many of the habitat management tools used on

Seefeld Estate are subsidized via European Union agricultural programs, therefore would not be difficult for others to implement. Therefore I suggest that a management plan incorporating habitat management, predator control, and supplemental feeding be implemented across the region. Results from this study suggest that with proper management that approximately 15% of the landscape should be in set aside, game crop, and wetland to allow for survival of hens, nests, and broods. Development of breeding territories, through creation of shrubby field buffers, can not be forgotten since its availability has been shown to limit populations (Robertson 1996).

This research, in conjunction with winter work, on Seefeld Estate (Anderson 2002) allowed an understanding of the habitat requirements needed for pheasant populations.

However, additional research needs to be conducted to better understand the configuration of habitats, nest predators, chick survival, and if supplemental feeding is related to chick condition and survival. Through continued research and understanding of pheasant populations at Seefeld

Estate and Lower Austria will facilitate better management within the area and throughout

Eastern Europe where little is known about pheasant ecology and management.

LITERTURE CITED

Anderson, B.C. 2002. Habitat use and nesting ecology of Ring-necked Pheasant (Phasianus

colchicus) on a landscape dominated by agriculture in Lower Austria. Thesis, University

of Georgia, Athens, Georgia, USA.

Andren, H., 1995. Effects of landscape composition on predation rates at habitat edges. Pages

225-255 in L. Hansson, L. Fahrig, and G. Merriam, editors. Mosaic landscapes and

ecological processes. Chapman and Hall, London, United Kingdom.

Gates, J.M., and J.B. Hale. 1974. Seasonal movement, winter habitat use and population

distribution of an east central pheasant population. Wisconsin Department of Natural

Resources Technical Bulletin 76, Madison.

Hanson, L.E., and R.F. Progulske. 1973. Movements and cover preferences of pheasants in

South Dakota. Journal of Wildlife Management 37:454-461.

Hill, D.A. 1985. The feeding ecology and survival of pheasant chicks on arable farmland.

Journal of Applied Ecology 22:645-654.

Hoodless, A.N., R.A.H. Draycott, M.N. Ludiman, and P.A. Robertson. 1999. Effects of

supplemental feeding on territoriality, breeding success and survival of pheasants.

Journal of Applied Ecology 36:147-156.

Kuck, T.L., R.B. Dahlgren, and D.R. Progalske. 1970. Movements and behavior of hen

pheasants during the nesting season. Journal of Wildlife Management 34:626-630.

Riley, T.Z.,, W.R. Clarke, D.E. Ewing, and P.A. Vohs. 1998. Survival of ring-necked pheasant

chicks during brood rearing. Journal of Wildlife Management 62:36-44.

_____, and S.P. Riley. 1999. Temporal comparison of pheasant brood size in the Midwest.

Wildlife Society Bulletin 27:366-373.

_____, and J.H. Schulz. 2001. Predation and ring-necked pheasant population dynamics.

Wildlife Society Bulletin 29:33-38.

Robertson, P.A., 1996. Does nesting cover limit abundance of ring-necked pheasants in North

America? Wildlife Society Bulletin 24:98-106.

Schmitz, R.A., and W.M. Clark. 1999. Survival of ring-necked pheasant hens during spring in

relation to landscape features. Journal of Wildlife Management 63:147-154.

Smith, S.A., N.J. Stewart, and J.E. Gates. 1999. Home ranges, habitat selection and mortality of

ring-necked pheasants (Phasianus colchicus) in North-central Maryland. American

Midland Naturalist 141:185-197.

Warner, R.E., 1979. Use of cover by pheasant broods in east-central Illinois. Journal of

Wildlife Management 43:334-346.

_____, and G.B. Joselyn. 1986. Responses of Illinois ring-necked pheasants to block roadside

management. Journal of Wildlife Management 50:525-532.

APPENDICES

Appendix A. Patagial tag, date captured, weight (g), tarsus (mm), condition index

(weight/tarsus), feather shaft (mm), age, capture location, and cover at trap of hen pheasants on

Seefeld Estate, Lower Austria, Austria 2002.

Tag Date Weight Tarsus Condition Feather Age Capture location Cover UTMx UTMy 1523 10/ Mar 02 1080 64.0 16.9 2.7 Y 593337 5397479 WS 1524 2/ Mar 02 1100 67.0 16.4 2.8 A 590412 5396640 WS 1525 2/ Mar 02 910 60.1 15.1 2.6 Y 587995 5395990 WS 1535 31/ Mar 02 1120 58.6 19.1 2.9 A 592239 5396845 C/SA 1538 31/ Mar 02 1140 64.6 17.6 2.8 A 592615 5397586 WS 1540 31/ Mar 02 1040 63.1 16.5 2.3 Y 593612 5397774 WS 1541 10/ Apr 02 1300 62.4 20.8 2.6 Y 586126 5397664 SA 1542 3/ Apr 02 960 58.7 16.4 2.2 Y 586685 5397126 W 1543 30/ Mar 02 1300 63.8 20.4 2.7 Y 586685 5397126 W 1544 11/ Mar 02 1060 57.7 18.4 2.8 A 590412 5396967 WS 1545 28/ Mar 02 1160 60.4 19.2 2.8 A 593612 5397774 WS 1546 31/ Mar 02 980 63.7 15.4 2.7 Y 594313 5396845 C/CL 1547 10/ Mar 02 1100 62.0 17.7 2.9 A 597337 5397479 WS 1549 28/ Mar 02 1120 61.0 18.4 2.8 A 594313 5396845 C/CL 1550 10/ Mar 02 1040 57.2 18.2 2.7 Y 597471 5396997 WS 1565 31/ Mar 02 1020 60.0 17.0 2.5 Y 592615 5397586 WS 1567 3/ Mar 02 1000 58.0 17.2 2.8 A 589904 5394339 WS 1568 3/ Mar 02 980 58.0 16.9 2.7 Y 589116 5396314 WS 1569 6/ Mar 02 980 60.1 16.3 2.8 A 590412 5396967 WS 1570 6/ Mar 02 1040 56.4 18.4 2.8 A 589027 5395278 WS 1571 4/ Mar 02 1060 59.9 17.7 2.8 A 589116 5397314 WS 1572 4/ Mar 02 1100 66.1 16.6 2.9 A 587990 5395980 WS

Appendix A. (continued).

Tag Date Weight Tarsus Condition Feather Age Capture location Cover UTMx UTMy 1573 2/ Mar 02 860 60.6 14.2 2.4 Y 589116 5396314 WS 1574 1/ Mar 02 820 58.8 13.9 2.1 Y 589116 5396314 WS 1575 1/ Mar 02 1040 59.0 17.6 2.2 Y 589116 5396314 WS 1583 5/ Apr 02 1020 59.4 17.2 2.8 A 593612 5397774 WS 1584 5/ Apr 02 1100 63.2 17.4 2.8 A 586685 5397126 W 1585 29/ Mar 02 1120 57.5 19.5 2.9 A 593612 5397774 WS 1587 23/ Mar 02 1100 60.2 18.3 2.4 Y 593612 5397774 WS 1588 11/ Mar 02 820 54.3 15.1 - - 589545 5395377 WS 1589 10/ Apr 02 1360 64.4 21.1 2.8 A 593612 5397774 WS 1590 28/ Mar 02 1160 62.8 18.5 2.6 Y 594313 5396845 C/CL 1591 10/ Mar 02 1000 58.2 17.2 2.8 A 589545 5395377 WS 1592 27/ Mar 02 940 61.8 15.2 2.2 Y 592239 5396845 C/SA 1593 18/ Mar 02 1040 61.1 17.0 2.9 A 597337 5397479 WS 1594 8/ Mar 02 880 57.8 15.2 2.4 Y 590412 5396967 WS 1595 30/ Mar 02 1160 61.1 19.0 2.8 A 594313 5396845 C/CL 1596 6/ Mar 02 880 52.4 16.8 2.4 Y 588281 5395375 WS 1597 7/ Mar 02 920 55.1 16.7 2.1 Y 588281 5395375 WS 1598 12/ Mar 02 1220 63.4 19.2 2.9 A 597337 5397479 WS 1599 27/ Mar 02 1040 63.4 16.4 2.5 Y 593612 5397774 WS 1600 2/ Mar 02 1020 62.7 16.3 2.8 A 588281 5395375 WS - 1/ Mar 02 - - - - 590412 5396967 WS - 29/ Mar 02 1040 59.0 17.6 2.4 Y 592615 5397586 WS

Appendix B. Patagial tag, date captured, weight (g), tarsus (mm), condition index

(weight/tarsus), feather shaft (mm), age, capture location, and cover at trap of hen pheasants on

Seefeld Estate, Lower Austria, Austria 2003.

Tag Date Weight Tarsus Condition Feather Age Capture location Cover UTMx UTMy 1372a 2/ Mar 03 960 56.3 17.05 2.9 A 589219 5397421 WS 1501 28/ Mar 03 1040 59.7 17.42 3.0 A 589033 5396353 WS 1503 9/ Mar 03 1100 61.4 17.92 3.0 A 589908 5394394 WS 1505 9/ Mar 03 940 61.0 15.41 2.5 Y 588693 5396097 SA 1506 9/ Mar 03 900 63.3 14.22 3.0 A 593471 5396982 WS 1507 9/ Mar 03 940 57.4 16.38 2.4 Y 593658 5397730 WS 1508 6/ Mar 03 1100 60.1 18.30 2.9 A 586553 5394965 W 1509 6/ Mar 03 1000 59.6 16.78 3.0 A 586553 5394965 W 1512 14/ Mar 03 920 61.5 14.96 2.6 Y 593658 5397730 WS 1513 9/ Mar 03 900 60.3 14.93 2.6 Y 593658 5397730 WS 1514 10/ Mar 03 1200 62.3 19.26 3.1 A 586536 5397613 V 1516 24/ Mar 03 980 60.1 16.31 2.4 Y 589219 5397421 WS 1517 6/ Mar 03 960 62.3 15.41 2.8 A 591627 5396969 WS 1518 9/ Mar 03 900 60.4 14.90 2.7 Y 586447 5397587 GC 1519 7/ Mar 03 960 61.4 15.64 2.8 A 586536 5397613 V 1520 8/ Mar 03 960 62.1 15.46 2.7 Y 586536 5397613 V 1521 5/ Mar 03 820 56.9 14.41 2.8 A 589039 5396358 WS 1522 10/ Mar 03 700 56.0 12.50 2.2 Y 591687 5396965 WS 1526 8/ Mar 03 1020 58.7 17.38 2.5 Y 589721 5396720 W 1527 19/ Mar 03 1140 59.4 19.19 2.7 Y 585785 5396036 SA 1529 9/ Mar 03 980 59.8 16.39 3.1 A 591687 5396965 WS 1530 30/ Mar 03 1100 66.0 16.67 2.2 Y 592265 5397595 WS 1531 17/ Mar 03 1020 59.0 17.29 2.8 A 593658 5397730 WS 1532 10/ Mar 03 980 58.0 16.90 2.9 A 586536 5397613 V 1533 14/ Mar 03 720 56.7 12.70 2.3 Y 591687 5396965 WS 1536 6/ Mar 03 880 53.9 16.33 2.6 Y 591627 5396969 WS 1539 17/ Mar 03 880 58.4 15.07 2.4 Y 585914 5396338 WS 1551 5/ Mar 03 1160 65.6 17.68 2.6 Y 589219 5397421 WS

Appendix B. (continued).

Tag Date Weight Tarsus Condition Feather Age Capture location Cover UTMx UTMy 1552 10/ Mar 03 980 60.3 16.25 2.5 Y 593658 5397730 WS 1553 7/ Mar 03 1040 62.1 16.75 2.9 A 586573 5394975 W 1555 10/ Mar 03 1040 59.2 17.57 3.1 A 593658 5397730 WS 1556 1/ Mar 03 980 60.2 16.28 2.6 Y 589219 5397421 WS 1557 18/ Mar 03 920 56.2 16.37 2.9 A 589219 5397421 WS 1558 5/ Mar 03 1080 58.5 18.46 2.2 Y 589898 5394404 WS 1559 18/ Mar 03 1060 61.1 17.35 2.8 A 589219 5397421 WS 1560 8/ Mar 03 980 61.5 15.93 2.7 Y 589219 5397421 WS 1562 4/ Mar 03 960 57.3 16.75 2.6 Y 591687 5396965 WS 1563 3/ Mar 03 1040 61.8 16.83 2.8 A 591687 5396965 WS 1566 11/ Mar 03 900 61.0 14.75 2.4 Y 591687 5396965 WS 1575 18/ Mar 03 1140 60.0 19.00 2.8 A 589219 5397421 WS 1576 12/ Mar 03 1000 59.1 16.92 2.9 A 593658 5397730 WS 1577 10/ Mar 03 1100 58.4 18.84 2.8 A 589033 5396353 WS 1579 7/ Apr 03 1040 61.4 16.94 2.5 A 592265 5397595 WS 1580 9/ Mar 03 1040 61.3 16.97 2.8 A 586536 5397613 V 1581 1/ Apr 03 1100 59.4 18.52 3.0 A 592265 5397595 WS 1582 6/ Mar 03 760 59.3 12.82 3.0 A 586531 5397603 V 1586 1/ Apr 03 1120 61.0 18.36 3.0 A 592265 5397595 WS n/a 12/ Mar 03 780 53.9 14.47 2.3 Y 589033 5396353 WS a indicates hen tagged in spring 2001.

Appendix C. Models considered to determine the effect of dispersal (A), condition index (B), age (C), agriculture (D), set aside (E), woodland (F), amount edge within the home range (G), game crop (H), and wetland (I) upon the survival of pheasant hens on Seefeld Estate, Lower

Austria, Austria 2002 – 2003.

A. Model AICc Delta AICc AICc Weight Estimate SE {S(.) disp} 244.6017 0.0000 0.2949 0.6267 0.4722 {S(.) disp+cond} 245.9057 1.3040 0.1537 0.5404 0.4829 {S(.) edge+disp} 246.4828 1.8811 0.1151 0.5669 0.5011 {S(.) age+disp} 246.5472 1.9455 0.1115 0.6258 0.4722 {S(g) disp} 246.6085 2.0068 0.1081 0.6269 0.4722 {S(g) cond+disp} 247.9007 3.2990 0.0567 0.5373 0.4833 {S(.) age+disp+cond} 247.9104 3.3087 0.0564 0.5421 0.4832 {S(g) edge+disp} 248.4957 3.8940 0.0421 0.5675 0.5015 {S(g) age+disp} 248.5605 3.9588 0.0407 0.6259 0.4722 {S(g) age+cond+disp} 249.9065 5.3048 0.0208 0.5390 0.4835

B. Model AICc Delta AICc AICc Weight Estimate SE {S(.) cond} 245.2467 0.0000 0.3104 -0.3589 0.3191 {S(.) disp+cond} 245.9057 0.6590 0.2233 -0.2740 0.3281 {S(g) cond} 247.2198 1.9731 0.1157 -0.3712 0.3253 {S(.) age+cond} 247.2546 2.0079 0.1137 -0.3563 0.3242 {S(g) cond+disp} 247.9007 2.6540 0.0823 -0.2840 0.3359 {S(.) age+disp+cond} 247.9104 2.6637 0.0819 -0.2685 0.3339 {S(g) age+cond} 249.2298 3.9831 0.0424 -0.3680 0.3300 {S(g) age+cond+disp} 249.9065 4.6598 0.0302 -0.2783 0.3407

C. Model AICc Delta AICc AICc Weight Estimate SE {S(.) age} 246.4672 0.0000 0.2343 -0.1093 0.4125 {S(.) age+disp} 246.5472 0.0800 0.2251 -0.1047 0.4129 {S(.) age+cond} 247.2546 0.7874 0.1580 -0.0197 0.4190 {S(.) age+disp+cond} 247.9104 1.4432 0.1138 -0.0393 0.4194 {S(g) age} 248.4774 2.0102 0.0857 -0.1089 0.4170 {S(g) age+disp} 248.5605 2.0933 0.0823 -0.1035 0.4184 {S(g) age+cond} 249.2298 2.7626 0.0589 -0.0250 0.4203 {S(g) age+cond+disp} 249.9065 3.4393 0.0420 -0.0444 0.4213

Appendix C. (continued).

D. Model AICc Delta AICc AICc Weight Estimate SE {S(.) ag} 244.5738 0.0000 0.5327 1.2556 0.8693 {S(.) edge+ag} 246.5527 1.9789 0.1980 1.1730 0.9934 {S(g) ag} 246.5674 1.9936 0.1966 1.2649 0.8740 {S(g) edge+ag} 248.5558 3.9820 0.0727 1.1879 1.0050

E. Model AICc Delta AICc AICc Weight Estimate SE {S(.) sa} 244.7305 0.0000 0.5162 -1.5381 1.0673 {S(.) edge+sa} 246.5227 1.7922 0.2107 -1.3807 1.1310 {S(g) sa} 246.6589 1.9284 0.1968 -1.6118 1.1057 {S(g) edge+sa} 248.5558 3.8253 0.0762 -1.4459 1.1793

F. Model AICc Delta AICc AICc Weight Estimate SE {S(.) wd} 245.6382 0.0000 0.5056 -2.0379 2.0330 {S(.) edge+wd} 247.2893 1.6511 0.2214 -1.6898 2.1574 {S(g) wd} 247.5983 1.9601 0.1897 -2.1824 2.1329 {S(g) edge+wd} 249.2465 3.6083 0.0832 -1.8363 2.2450

G. Model AICc Delta AICc AICc Weight Estimate SE {S(.) edge} 245.8398 0.0000 0.1652 -0.1970 0.2312 {S(.) edge+disp} 246.4828 0.6430 0.1198 -0.0913 0.2522 {S(.) edge+sa} 246.5227 0.6829 0.1174 -0.1102 0.2353 {S(.) edge+ag} 246.5527 0.7129 0.1157 -0.0459 0.2595 {S(.) edge+wd} 247.2893 1.4495 0.0800 -0.1457 0.2405 {S(.) edge+ws} 247.5717 1.7319 0.0695 -0.2827 0.2733 {S(.) edge+gc} 247.7637 1.9239 0.0631 -0.2015 0.2329 {S(g) edge} 247.8483 2.0085 0.0605 -0.1979 0.2321 {S(g) edge+sa} 248.4925 2.6527 0.0439 -0.1017 0.2391 {S(g) edge+disp} 248.4957 2.6559 0.0438 -0.0908 0.2532 {S(g) edge+ag} 248.5558 2.7160 0.0425 -0.0417 0.2629 {S(g) edge+wd} 249.2465 3.4067 0.0301 -0.1467 0.2399 {S(g) edge+ws} 249.5819 3.7421 0.0254 -0.2835 0.2738 {S(g) edge+gc} 249.7772 3.9374 0.0231 -0.2014 0.2336

Appendix C. (continued).

H. Model AICc Delta AICc AICc Weight Estimate SE {S(.) gc} 246.4740 0.0000 0.4799 -0.6733 2.6289 {S(.) edge+gc} 247.7637 1.2897 0.2518 -0.7777 2.6085 {S(g) gc} 248.4780 2.0040 0.1762 -0.7008 2.6528 {S(g) edge+gc} 249.7772 3.3032 0.0920 -0.7797 2.6429

I. Model AICc Delta AICc AICc Weight Estimate SE {S(.) ws} 246.5317 0.0000 0.4591 -0.2627 3.3388 {S(.) edge+ws} 247.5717 1.0400 0.2729 2.0941 3.9586 {S(g) ws} 248.5409 2.0092 0.1681 -0.2423 3.4034 {S(g) edge+ws} 249.5819 3.0502 0.0999 2.1366 4.0218

Appendix D. Models considered to determine the effect of condition index of hen (A), age (B), game crop (C), wetland (D), amount edge within the home range (E) nesting habitat (F), woodland (G), agriculture (H), and set aside (I) upon survival of pheasant broods on Seefeld

Estate, Lower Austria, Austria 2001 – 2003.

A. Model AICc Delta AICc AICc Weight Estimate SE {S(.) cond} 79.1783 0.0000 0.5191 0.2136 0.2304 {S(g) cond} 81.0132 1.8349 0.2074 0.1575 0.2462 {S(.) age+cond} 81.1923 2.0140 0.1896 0.2026 0.2677 {S(g) age+cond} 82.8225 3.6442 0.0839 0.0646 0.3165

B. Model AICc Delta AICc AICc Weight Estimate SE {S(.) age+nest} 79.2473 0.0000 0.3288 0.3202 0.7722 {S(.) age} 79.7783 0.5310 0.2521 0.4244 0.7682 {S(g) age} 80.8300 1.5827 0.1490 0.6041 0.7801 {S(.) age+cond} 81.1923 1.9450 0.1243 0.0712 0.8954 {S(g) age+nest} 81.8216 2.5743 0.0908 0.4901 0.7862 {S(g) age+cond} 82.8225 3.5752 0.0550 0.4727 1.0061

C. Model AICc Delta AICc AICc Weight Estimate SE {S(.) gc} 73.8348 0.0000 0.4006 562.9608 0.0000 {S(.) edge+gc} 74.1812 0.3464 0.3369 706.7263 0.0000 {S(g) gc} 75.5682 1.7334 0.1684 550.1875 0.0000 {S(g) edge+gc} 76.7321 2.8973 0.0941 560.7566 0.0000

D. Model AICc Delta AICc AICc Weight Estimate SE {S(.) ws} 75.4179 0.0000 0.3018 36.4257 24.7621 {S(g) ws} 75.4344 0.0165 0.2993 35.2744 21.3196 {S(.) edge+ws} 75.8560 0.4381 0.2424 38.7630 25.6559 {S(g) edge+ws} 76.7321 1.3142 0.1564 35.9768 21.8927

Appendix D. (continued).

E. Model AICc Delta AICc AICc Weight Estimate SE {S(.) edge+gc} 74.1812 0.0000 0.4305 -0.3922 0.2735 {S(.) edge+ws} 75.8560 1.6748 0.1863 -0.3585 0.2592 {S(g) edge+gc} 76.7321 2.5509 0.1202 -0.2805 0.2836 {S(g) edge+ws} 76.8820 2.7008 0.1115 -0.2017 0.2526 {S(.) edge} 79.0030 4.8218 0.0386 -0.3567 0.3161 {S(.) edge+wd} 79.2053 5.0241 0.0349 -0.2823 0.3365 {S(.) edge+ag} 80.0919 5.9107 0.0224 -0.6290 0.4334 {S(g) edge} 81.0002 6.8190 0.0142 -0.2204 0.3194 {S(.) edge+sa} 81.0018 6.8206 0.0142 -0.3897 0.3897 {S(g) edge+wd} 81.2237 7.0425 0.0127 -0.1798 0.3403 {S(g) edge+ag} 81.9123 7.7311 0.0090 -0.5612 0.4545 {S(g) edge+sa} 82.9794 8.7982 0.0053 -0.2794 0.4059

F. Model AICc Delta AICc AICc Weight Estimate SE {S(.) nest} 77.4007 0.0000 0.5695 -1.2962 0.8409 {S(.) age+nest} 79.2473 1.8466 0.2262 -1.2687 0.8435 {S(g) nest} 80.1810 2.7803 0.1418 -0.9811 0.9108 {S(g) age+nest} 81.8216 4.4209 0.0624 -0.9052 0.9192

G. Model AICc Delta AICc AICc Weight Estimate SE {S(.) wd} 77.8192 0.0000 0.4707 -14.4692 8.4039 {S(.) edge+wd} 79.2053 1.3861 0.2354 -13.5291 8.9548 {S(g) wd} 79.4521 1.6329 0.2081 -15.6624 10.7632 {S(g) edge+wd} 81.2237 3.4045 0.0858 -15.5112 11.1838

H. Model AICc Delta AICc AICc Weight Estimate SE {S(.) ag} 80.0663 0.0000 0.3432 -0.1995 1.3888 {S(.) edge+ag} 80.0919 0.0256 0.3388 -1.9483 2.2736 {S(g) ag} 81.3385 1.2722 0.1817 -0.4469 1.4505 {S(g) edge+ag} 81.9123 1.8460 0.1364 -2.3337 2.5001

Appendix D. (continued).

I. Model AICc Delta AICc AICc Weight Estimate SE {S(.) sa} 79.9250 0.0000 0.4385 -0.6112 1.4584 {S(.) edge+sa} 81.0018 1.0768 0.2560 0.2809 1.9473 {S(g) sa} 81.3951 1.4701 0.2103 -0.3217 1.5330 {S(g) edge+sa} 82.9794 3.0544 0.0952 0.4728 2.0668

Appendix E. Example of a small breeding season 100% MCP home range (2.7 ha) using #1577. Observe mixture of preferred hen habitats, set aside and wetland, along with male territory habitat, woodland edge and coppice. Seefeld Estate, Lower Austria, Austria

2002-2003.

Appendix F. Example of a large breeding season 100% MCP home range (264 ha) using #1595. Notice low amount of male territory habitat, woodland and coppice, within the home range. Seefeld Estate, Lower Austria, Austria 2002-2003.

Appendix G. Example of hen dispersal using #1570 showing the 100% MCP home range during the breeding season on Seefeld

Estate, Austria during 2002.

Appendix H. Example of a small 100% MCP brood home range (2.6 ha) using #1512. Home range is comprised of well managed set aside, game crop and wetland habitats which provide food and cover. Seefeld Estate, Lower Austria, Austria 2002-2003.

Appendix I. Example of an average 100% MCP brood home range (11 ha) using #1573, which contains a good mixture of set aside, game crop, wetland, and agricultural land. Notice that majority of locations lie within game crop. Seefeld Estate, Lower Austria,

Austria 2002-2003.

Appendix J. Example of a large 100% MCP brood home range (44 ha) using #1568. Observe that the home range is dominated by agricultural land which increased the area covered in order to find preferred set aside habitat. Seefeld Estate, Lower Austria, Austria

2002-2003.