michael r. frisina
irgaref frisina

Montana State Library

iiiiiiiiiiii ii i rfwiiiu

3 0864 1006 0678 2

weather
wi(d?fe

volume ii:up(and game birds

an annotated bibliography

Michael R. Frisina
R. Margaret Frisina

A PUBLICATION OF THE HABITAT SECTION
WILDLIFE BUREAU

Cover page and title page photography copyright 2003 Michael R. Frisina.

This document was prepared under Project W-154-R. Copyright 2008 Montana Department of
Fish, Wildlife & Parks. All rights reserved. Printed in the United States of America. Permission to
reproduce or copy any portion of this report is granted on condition of full credit to Montana
Department of Fish, Wildlife & Parks and the authors/photographers. Not for sale/resale.

Rough Writers ^Tt+e&o

Robert L. Eng

Terry Lonner photo

Weather & Wildlife Volume II: Upland Game Birds

is dedicated to Dr. Robert L. Eng in recognition of his

extensive research and publication on the

life history of upland game birds.

%

fy ^ ^

Digitized by the Internet Archive

in 2012 with funding from

Montana State Library

http://archive.org/details/weatherwildlifea37fris

Michael R. Frisina copyright 1999

Table of Contents

Definitions

ix

Foreword

xi

Preface

xiii

Acknowledgements
Introduction

XV

xvii

Bibliography
Avian Taxa

1
251

Author Index

253

Topic Index

273

^

% ^te***

'Much of wildlife management is in the nature

of insurance against future

probable contingencies"

R. T. King 1937 Ruffed Grouse Management

"The ups and downs of any animal population

are just as natural as changes in the weather.

It is even fair to say, dependent on

changes in the weather. The weather furnishes them

a poor house or a good house, depending on how much

cover is grown by the rains which may

or may not fall. It freezes or bakes them. "

R. Dahlgren 1967 The Pheasant Decline

J.

> >' <c> ^

VII

Definitions

Climate: the average weather conditions throughout the seasons over a fairly wide or
very extensive area of the earth's surface and considered over many years (usually 30
to 35 years) in terms of climatic elements, climatology, local climate,

MACROCLIMATE, MESOCLIMATE, MICROCLIMATE, MICROCLIMATOLOGY.

Weather: a general term for the conditions prevailing in the atmosphere-1 , especially in
the layer near the ground (toposphere), over a short period of time (in contrast to
climate) or at a specific time, at any one place, and as affecting human beings.
Temperature, sunshine, pressure and wind, humidity, amount of cloud, precipitation
(rain, sleet, hail, snow), the presence of fog or mist are all taken into account.

Clark, Audrey N. 1998. Dictionary of Geography. Penguin Books, New York.

4-

K fc* >f*

vfc

•'■■/■ - ( • .

^^m:^J*?'r' r

7Wz*^ , V-, i \

X

■**

Michael R. Frisina copyright 2003

Foreword

In recent years, much ado has been made about climate change, using models that are
complex, incomplete and not well understood. Most people, however, readily recognize
the effects of daily and seasonal weather on their lives.

So it is also in the lives of wild animals — weather effects their movements, growth and
survival. Wildlife management pioneers suggested that the effects of weather should be
incorporated into the new science of wildlife studies and management. We have been
very slow to do so.

This document is an effort to encourage working field biologists to better understand and
incorporate weather and its effects into their wildlife and habitat management
recommendations and actions. My hope is this publication will establish weather in its

% f>"^^

rightful place as a major factor in professional wildlife management decisions as well
in the public's understanding of weather impacts on wildlife populations.

ffi^} \ far

Steve Knapp

HABITAT SECTION MANAGER

XII

% ;M*fc*>ffc

c *->

^:

»

1. •

-y

Michael R. Frisina copyright 1999

Preface

Weather & Wildlife, Volume II: Upland Game Birds emphasizes literature describing the
influence of weather-related events these birds rather than addressing the larger topic of
global climate change. While the bibliography contains a few papers related to modeling,
our focus is on the direct and indirect influence of weather events on the life cycle,
ecology, and evolution of upland game birds.

While not comprehensive, this bibliography of over 500 entries, combined with additional
references in the introductory material, provides the wildlife manager, researcher, and
student with an extensive cross-section of literature on the subject. Research from

^ ^ ^

XIII

around the world was sought based upon an extensive database search including,
among others, JSTOR, BioOne, Biological and Agricultural Index, Wildlife and Ecology
Studies World Wide, Zoological Record and Biological Abstracts.

Weather & Wildlife, Volume II: Upland Game Birds is a companion to Weather & Wildlife,
Volume I: Large Ungulates.

JH

% fxfc*>f*

Roberta Bomar photo.

Acknowledgements

The authors appreciate the help offered by the following individuals and libraries in
obtaining literature contained in this bibliography:

Dr. Jon Swenson, Professor, Agricultural University of Norway

Renne Library, Montana State University-Bozeman

Mansfield Library, University of Montana-Missoula

Montana State Library-Helena; particularly Marjorie Smith, Client Services.

** ^ t*

xv

R. Margaret Frisina copyright 2006

Introduction

Weather & Wildlife:Volume /emphasized research papers defining the relationship
between weather and large ungulates; the 562 references in Volume II illustrate the
relationship between weather and upland game birds. Together Volumes I and II provide
the reader with a source of 957 publications on the subject of weather and wildlife. Our
hope is that wildlife managers and researchers will find the bibliography useful in making
science-based decisions for the conservation of upland game birds. Since the inception
of the profession, wildlife managers have struggled with how to account for varying and
unpredictable impacts of weather patterns on wildlife. W&W II is intended to provide a
resource for the professional, student and generalist in considering this important
variable in upland game bird management.

4

f \ itf* ^=*C

XVII

c£^

Bibliography

"Strong habitat is the best and only
practicable defense against extremes of weather.

Hamerstrom, Mattson & Hamerstrom - 1957
A Guide to Prairie Chicken Management

001

Adler, P. H., D. Roach, W. K. Reeves, J. P. Flanagan, M. E. Morrow, and J. E.
Toepfer. 2007. Attacks on the endangered Attwater's prairie-chicken
(Tympanuchus cupido attwateri) by black flies (Diptera: Simuliidae) infected with
an avian blood parasite. Journal of Vector Ecology 32(2):309-312. Abstract: With
fewer than 50 birds remaining in the wild, Attwater's prairie-chicken {Tympanuchus
cupido attwateri) is critically endangered. Individuals of this species on the Attwater
Prairie Chicken National Wildlife Refuge, Colorado Co., TX have been attacked in
successive winters, 2005-2006, by the blood-feeding black fly Cnephia ornithophilia.
Attwater's prairie-chicken is a previously unreported host for Cnephia ornithophilia.
Molecular screening indicated that about 15% of 13 blood-fed flies sampled from
captured Attwater's prairie-chickens carried parasite of the genus Leucocytozoon that
can cause a debilitating avian malaria-like disease. If blood feeding or transmission of
the disease agent becomes a threat to the birds, particularly in years of lean food
supply or harsh winter, management of Cnephia ornithophilia should be considered.
Key words: prairie chicken, severe winter, nutrition, insects, disease

002

Aldrich, J. W. 1963. Geographic orientation of American Tetraonidae. Journal of
Wildlife Management 27(4):529-545. Notes: "In the ruffed grouse, we find a classic
example of the correlation of intensity of coloration with the amount of moisture in the
environment, as well as with the density and background shades of the vegetation
cover, themselves correlated with moisture. Thus the darkest ruffed grouse are in the
lush forests of the Olympic Peninsula rain forest and the palest in the relatively arid
aspen groves of the Great Basin Mountains of Utah and the low hills rising out of the
shortgrass prairies of the central states and provinces." Key words: ruffed grouse,
precipitation, humidity, morphology

003

Aldridge, C. L. 2003. Understanding the decline — is there hope for Alberta's sage-
grouse? Biodiversity Makes It Work; Newsletter of the Alberta NWAMP
Partnership. Ducks Unlimited, Edmonton, Canada. Notes: Aldridge concluded that a

% VHt&ip

^?& -'rar- ^^ c£Zi

large mid-May snow storm and three days of continuous cold, heavy rain in 2002
resulted in the lowest-ever-recorded nest success for the population in southeastern
Alberta: only 5% of 37 nest attempts were successful (13.5%), due to eggs being frozen
early in incubation. Key words: sage-grouse, snow, temperature, precipitation,
reproduction

004

. 2000. Reproduction and habitat use by sage-grouse {Centrocercus

urophasianus) in a northern fringe population. M.S. Thesis, University of Regina,
Regina, Canada. Notes: Pages 33-34 — "Climate. Although sage-grouse are fairly
robust birds, harsh climatic conditions at the northern edge of the species' range may
affect populations. Short summers and particularly harsh winters likely reduce the ability
of individuals to find enough food in winter months and decrease lipid reserves
necessary for reproduction and possibly lower overwinter survival. There is a positive
relationship between spring precipitation and sage-grouse productivity. During the
1980s, spring precipitation was considerably lower than the long-term average. This
likely contributed to decreased productivity and survival. The effects of other limiting
factors may be compounded during drought conditions. For example, consistent cattle
stocking rates during the drought of the 1980s may have resulted in a substantial loss of
vegetative cover, perhaps lowering nest success, increasing predation, and possibly
lowering overwinter survival. Impacts may have been particularly severe in more moist
habitats, which supply important herbaceous growth during nesting and brood rearing.
The attraction of cattle to these areas was probably increased during drought
conditions, which may have decreased brood survival." Page 84 — "I did not observe a
shift in habitat used by broods, which typically occurs due to changing dietary
requirements of chicks... .Spring precipitation was above average in both years of my
study, resulting in increased vegetation growth. Thus, broods likely had increased food
resources available to them in sagebrush habitat, allowing them to remain in sagebrush
uplands. In dry years, broods should have to move from sagebrush uplands to more
mesic sites. However, mesic wetland type habitats are generally limited and may not be
available for broods in dry years. Thus, brood survival may be even lower during times
of drought." Page 85 — "In years when precipitation is average or below average, forb
cover may be below that required to provide suitable brood habitat. The lack of a shift in
brood habitat between early and late-brood rearing in my study suggests that
differences in the availability of forbs did not exist, at least in wet years. However, the
limited cover provided by forbs at brood locations despite high spring precipitation
suggests that key brood habitat in moist wetlands and drainages may be limiting in
southeastern Alberta, even in moist years." Page 85 — "Virtually all research on brood
habitat use has found that areas with forbs are selected and that a shift to more mesic
sites occurs after broods reach six weeks of age. Sage-grouse broods in my study did
not select use sites based on forb availability and no shift in habitat use occurred. Mesic
areas with nutrient rich forbs may be limiting and forbs as a food resource may be even
more limiting during drier years. Lack of suitable moist drainages for broods to forage in
may also be a factor contributing to low chick survival and poor recruitment." Key
words: sage-grouse, precipitation, drought, grazing, vegetation, productivity,
mortality, movement, habitat use, recruitment

4*

fci t& t*

005

. 1998. Status of the sage-grouse (Centrocercus urophasianus) in Alberta.

Alberta Environmental Protection, Wildlife Management Division, and Alberta
Conservation Association, Wildlife Status Report No. 13, Edmonton, Canada.

Notes: Page 14 — Climate. Summarizes research of weather impacts on sage-grouse.
The author notes that the drought of the 1980s may have limited productivity and
contributed to a population decline. He also states that the impacts of other limiting
factors may be compounded during drought and goes on to cite stocking rates during
drought that may have resulted in substantial loss of vegetative cover possibly
impacting nesting success, predation and winter survival. Key words: sage-grouse,
drought, temperature, precipitation, population dynamics, grazing, vegetation

006

and R. M. Brigham. 2003. Distribution, abundance, and status of the greater

sage-grouse, Centrocercus urophasianus, in Canada. Canadian Field-Naturalist
117:25-34. Notes: Page 30 — "Overgrazing has long been suggested as one of the main
reasons for declining sage-grouse numbers. The removal of vegetation cover by cattle
can have an impact on sage-grouse populations, either by reducing habitat suitability or
by increasing the exposure of birds and nests to predators or extreme weather, all of
which decrease survival and nest success... heavy grazing during drought conditions
could intensify these effects... grazing may simply decrease the carrying capacity of
sage-grouse habitat, especially in years with below average annual precipitation." Page
31 — "Climate. Although sage-grouse are large robust birds, harsh climatic conditions at
the northern edge of the species/ range likely affect populations. Short summers and
particularly harsh winters likely reduce the ability of individuals to find enough food in
winter months, especially given the low abundance of sagebrush in Canada. This would
result in decreased lipid reserves that are necessary for reproduction and possibly
reduce overwinter survival. There is a positive relationship between spring precipitation
and sage-grouse productivity. Years with below-average spring moisture result in less
vegetation growth, apparently reducing sage-grouse nest success and reducing the
availability of lush vegetation that is an important dietary component, especially for
chicks. During the 1980's, spring precipitation was considerably below the long-term
average. This likely contributed to decreased productivity and resulted in reduced chick
survival in Alberta. However, cold, wet spring precipitation events (rain or snow) can
also result in increased nest failures. The effects of other limiting factors may be
compounded during drought conditions. For example, consistent cattle-stocking rates in
Canada during the droughts of the 1980's may have resulted in a substantial loss of
vegetative cover, perhaps lowering nest success, increasing predation, and possibly
lowering overwinter survival. The impacts may have been particularly severe in more
moist habitats, which supply important herbaceous growth during nesting and brood
rearing. The probable increased attraction of cattle to these areas during drought
conditions may decrease chick survival." Key words: sage-grouse, precipitation,
drought, temperature, vegetation, reproduction, mortality, chick survival

i f*^**

<gj ^ <£^> d£>

007

Allen, D., editor. 1956. Pheasants in North America. Stackpole Company and
Wildlife Management Institute, Washington, D.C., USA. Notes: Chapter 2 — Chinese
Pheasants in the Northwest: Page 56 — "Although a general reduction in bird
populations is apparent and explainable, wide fluctuations within this pattern are difficult
to account for. There are many indicators that they are correlated with weather cycles or
variations in temperature and rainfall during the nesting season." Page 61 — "During
periods of heavy and prolonged snowfall, there is evidence that a supplement of grit
may be of greater value to pheasants than other foods." Chapter 3 — Pheasant in
California: Page 105 — "All available evidence on pheasant populations and
reproduction for this area points to the fact that success hinges upon weather conditions
during spring and early summer." Page 107 - "It is believed that... indirect effects of
spring rainfall upon nesting by influencing farming practices is more important than
direct effect upon clutch size, hatch, or chick survival (Sacramento Valley rice belt).
Page 1 10 - San Joaquin Valley Habitat - 'Detailed studies of limiting factors have not
been made in this region, but low rainfall and unfavorable agricultural conditions
generally are believed to be basic." Imperial Valley - "Low rainfall is the basic reason for
poor nesting conditions." Chapter 4 — Pheasants in the Arid Southwest: Page 161 — u
Moisture-temperature relationships and land use practices that severely reduce cover
and food seem to explain the lack of success of the ringneck in the south and southeast
portions of the region." Page 171 — "...predation becomes serious during blizzards,
when hundreds, even thousands, of pheasants may be driven to the insecure shelter of
small wooded areas. Under such conditions, losses may be severe." "...blizzards. .loss
seems to result from the combination of low temperatures, heavy snow, and high wind;
and death actually is caused by suffocation rather than freezing. The greatest mortality
always occurs in cover-deficient areas, where birds roosting in stubble, or in other thin
cover, receive the full force of the wind." Page 171 — " Unusually heavy, wet snow in
May or early June may destroy many nests, forcing extensive renesting and late
hatching peaks. The obvious approach to meeting the contingencies of winter extremes
is the creation of more and better cover." Chapter 5 — Pheasants of the Plains and
Prairies: Page 210 - "...increased rainfall resulted in more abundant cover in the form
of sweet clover, sunflower, and other herbaceous species." Page 211 - "...population
fluctuation and shifts appear to be explainable on the basis of environmental changes."
"Highest loss came from freezing and choking during severe weather." "Mortality from
predators and from accidents is associated with winter storms and may be considered
as indirect storm losses... following a severe storm they are more susceptible to
predation since they may be encumbered by balls of ice and snow, and because
escape cover is filled with snow." Chapter 6 — Ringnecked Pheasants in the Great
Plains — Page 289 — "Hunger alone, prior to any physical evidence of starvation, is a
stress factor. When such stress is coupled with any one or a combination of extreme
cold, high wind, rain, pursuit by predators, reduction of cover, or deep snow, it may
render birds more vulnerable or even cause death." Page 320 - "There is a good
possibility that spring weather conditions may have their most significant effect through
the timing of such farm operations as hay mowing or sod plowing." Page 322 - "It
seems that local pheasant abundance may be explained by the occurrence of weather

4>

^ <C> **

extremes, but rangewide comparisons must of necessity rely on means.... it seems that
high temperature alone is not detrimental and that relative humidity or a
temperature:moisture ratio may be of importance." Page 324 - ". . .these ideas appear to
be important relative to the pheasant and weather: 1) cold and wet spring weather that
may be detrimental in one part of the Lake States range may be beneficial in another; 2)
High temperatures alone do not limit distribution; 3) Some relationship between
temperature and moisture governs suitability of pheasant range and perhaps yearly
productivity; 4) What to pheasants are ideal climatic conditions are not necessarily
those measured by weather stations." Chapter 8 — The Pheasant in the Northeastern
States: Page 400 — "...a 20%. ..decrease was [caused by] the combined result of
blizzard conditions in January and February climaxed by a severe ice storm in March,
with ice glaze persisting as much as a week in much of the pheasant range." Page 405
- "...the pheasant decline in New York started when weather conditions could not be
considered adverse, unless because of drouth, but the most rapid decline was
associated with cold and wet springs, probably aggravated by a hard winter." Key
words: ring-necked pheasant, temperature, precipitation, drought, snow, ice,
wind, humidity, mortality, population dynamics, habitat modification, predation,
productivity

008

Andreev, A. V. 1990. The winter biology of Siberian spruce grouse {Falcipennis
falcipennis) in the Priamurie. Zoologicheskii Zhurnal 69(3):69-80. Abstract: The
biotopes preferred by the Siberian spruce grouse in winter are concentrated in a zone
between spruce and larch forests, particularly close to mossy swamps. In the spruce
forests grouses feed and settle for night in the snow on margins of the mossy swamps.
The daylight activity of the birds starts and ends at the illumination intensity of 5 to 7 lux;
the daylight time the grouses spend in the tops of Picea jezoensis changing one to two
trees a day. The birds spend feeding about forty percent of the daylight time (in January
it is approximately 4 hours). The dominant food is coniferous needles. When feeding
intensively the grouse consumes every minutes from 37 to 73 needles mainly from the
branch ends in the spruce top. An average digestivity of the coniferous needles is 0.28;
the daily food needed in January is 89 g of dry and 1 50 g of wet food for the bird's
bodyweight of about 700 g. The total flight distance does not exceed 100 m per day and
their duration, 10 sec/day. The duration of one flight is 2.5 sec. At the air temperature
above -20 degree, the birds prefer spruce tops at night and dig in snow if it is cold. The
anthropogenic landscape transformation in the grouse biotopes resulted in a drastic
growth of the hazel hen population. It is assumed that in a number of regions the
competitive relations with the hazel may suppress the grouse population. Key words:
Siberian spruce grouse, solar radiation, snow, temperature, movement, behavior

009

. 1977. Temperature conditions in snow cavities of the hazel grouse

{Tetrastes bonasia kolymensis But.). Soviet Journal of Ecology 8(5):454-455.

Notes: The results of the measurements show that the temperature in the cavity can
change from -15 to -0.3 deg and depends little on the temperature of the outside air.
How the bird buried itself had much greater influence on the temperature in the cavity:

4*

H ^ ^

\~- C*~^-J^}> C J

the length, the bends in the entrance tunnel, and, in particular, the presence of an
opening between the ceiling of the chamber and the snow plug, closing the chamber
from the tunnel. If the plug was sufficiently tight, then even during a very strong frost the
temperature in the cavity could approach 0°...The maintenance of the high temperature
in the cavity is provided for not only by the insulating properties of snow and the heat
given off by the bird. Of important significant is the flow of heat coming from the soil
surface. It also depends on the air temperature and the thickness of the snow. From
November to March hazel grouse pass a large part of the day in the snow. Thus, in
spite of the severe climatic conditions they winter under highly mild microclimatic
conditions (it is only necessary that the thickness of the snow be more than 25 cm).
Specifically these conditions permit the smallest representative of tetraonid birds to
winter successfully at northern latitudes, being nourished by comparatively low-quality
branch forage. Key words: hazel grouse, temperature, snow, behavior, nutrition

010

Applegate, R. D. and T. Z. Riley. 1998. Lesser prairie chicken management.
Rangelands 20:13-15. Note: Winter cover can be particularly important in periods of
drought when cover, food, and moisture from leafy vegetation are limited. "There is little
free water across most of the range of lesser prairie chickens, and leafy vegetation may
be the only moisture available in winter during periods of drought. An extended drought
over several years can be devastating to prairie chickens. Flocks will travel great
distances to find sources of food and moisture, but mortality rates increase dramatically
when sources are difficult to locate." Key words: prairie chicken, drought,
vegetation, nutrition, movement

on

Archibald, H. L. 1976. Spring drumming patterns of ruffed grouse. Auk 93(4):808-
829. Abstract: Drumming of 7 male ruffed grouse [Bonasa umbellus] was monitored
from 7-56 days with modified automatic radio-tracking equipment during spring 1970.
Grouse generally drummed at the same times of day, but certain individuals drummed
at shorter intervals than others consistently throughout the day. The highest drumming
frequencies within a day tended to occur at the beginning of a drumming period and an
average of 38.7 min before sunrise. During the first 2 % wk of the drumming season,
nocturnal drumming was consistently recorded for all radio-marked males except under
windy or inclement weather conditions. Within about 5 days, drumming became nearly
exclusively diurnal and continued so thereafter. Peak drumming periods coincided with
the full moons in April and May. Estimated dates of copulation of 3 hens coincided with
the drumming peak in April. The 2nd peak may have been associated with remating of
hens that had lost their initial clutch. Five abiotic factors accounted for approximately
67% of the variation in the seasonal distribution of the average daily total number of
drums; the most important of these factors were moon fullness (seasonal variation) and
kilometers of wind/day (day-to-day variation). Two possible forms of communication
between adjacent males were identified: drumming shortly after the start of a neighbor's
drum and duplication of the interval between successive drums of a neighbor. Stimulus-
response functions for 2 neighboring grouse showed a tendency for the drums of 1 bird

.*

% Wt&ik

c^>

to occur either within 10 s or between 50 and 130 s after the drums of the other bird.
Key words: ruffed grouse, behavior, wind

012

Armbruster, J. W. 1994. Early season nesting success of mourning doves
(Zenaida macroura) in central Illinois. Transactions of the Illinois State Academy
of Science 87(1-2):37^1. Summary: Early season mourning dove {Zenaida macroura)
nests are subjected to cold weather which may reduce fledging success. In a residential
setting in central Illinois, a wind storm destroyed 50% of all nests and fledging success
was only 13.7% because of the storm and low hatching success of eggs that remained
in the nests. Although early season nests may often have low reproductive success, if
they are successful, they may result in fledglings that can breed by the end of the
season. Key words: mourning dove, wind, mortality, reproduction

013

Aulie, A. 1976. The pectoral muscles and the development of thermoregulation in
chicks of willow ptarmigan (Lapopus lagopus). Comparative Biochemistry and
Physiology 53A:343-346. Abstract: 1. The 02 consumption, EMG from m.pectoralis
major and body temperatures during cold exposure were measured and the weights of
pectoral muscles, leg muscles and liver recorded in ptarmigan chicks of different ages.
2. Up to the second day the pectoral muscle made up ~1 .8% of the body weight. From
the third day their mass increased faster than the rest of the body and was 1 1 % on the
twelfth day. 3. The shivering response was weak until the second day and increased
about 6-fold from the third day. 4. It was a clear connection between the increase in
pectoral muscle mass, maximal 02 consumption (m102/g hr) and improved resistance to
cold. Three factors seem to be of vital importance in the development of
thermoregulation in ptarmigan chicks: (1) The chick responds to cold exposure with
thermo-conserving behavior, remaining calm and fluffed up. (2) Activation of its skeletal
muscles strongly through shivering. In most chicks both these factors seem to appear
between the second and third day after hatching. From this stage, a further
improvement of the thermoregulatory capacity seems to depend on (3) the mass
increase of the pectoral muscles. Key words: willow ptarmigan, temperature,
thermoregulation, behavior

014

and P. Moen. 1975. Metabolic thermoregulatory responses in eggs and

chicks of willow ptarmigan {Lagopus lagopus). Comparative Biochemical
Physiology 51A:605-609. Abstract: (1). The Co2 production in eggs and chicks of
willow ptarmigan has been measured during cold exposure. (2) Fourteen-day-old
embryos showed no sign of thermoregulation. In the fully developed embryo a 10°C
drop in ambient temperature decreased the metabolic rate with only 26 percent,
indicating that a thermoregulatory response is present. (3) Newly hatched (wet) chicks
were unable to increase the metabolism when exposed to 29°C in spite of increased
muscular activity. Half a day old chicks increased the metabolism with 45 percent at this
temperature. No sign of shivering could be observed on the first day and the non-
shivering thermogenesis is thought to be caused by the increased muscular activity in

% K^^

£Z^> c^

connection with the flight behavior. (4) Some of the 2- and 3-day-old chicks balled up
and shivered when exposed to cold and became less hypothermic (35-0°C) than the
one that did not shiver (31 .2°C) when exposed to 19°C for 20 min. (5) Within 5 days
after hatching all the chicks balled up and shivered when exposed to cold and they
became only slightly hypothermic (35.8-38.3°C) when exposed to 19°C for 20 min. The
lower critical temperature in the 5- to 7-day-old chicks was about 30°C. (6) Brooding is
thought to be a more important mechanism than shivering in maintaining a high and
constant body temperature in the willow ptarmigan during the first week after hatching.
Key words: willow ptarmigan, temperature, thermoregulation, reproduction,
morphology

015

Austin, D. E. and L. W. DeGraff. 1975. Winter survival of wild turkeys in the
southern Adirondacks. Proceedings of the National Wild Turkey Symposium 3:55-

60. Abstract: The survival and behavior of an isolated population of wild turkeys
(Meleagris gallopavo silvestris) was investigated in the southern Adirondack Mountains,
1966-73. Five winters had mild or average weather conditions and the turkey survival
averaged 75 percent. Two winters were severe and the survival was only 55 percent.
Although the wild turkeys were adaptable to extreme winter weather conditions, the
severe winters caused sufficient mortality to limit population growth. Key words: wild
turkey, severe weather conditions, mortality, population dynamics

016

Back, G. N., M. R. Barrington, and J. K. McAdoo. 1987. Sage-grouse use of snow
burrows in northeastern Nevada. Wilson Bulletin 99(3):488-490. Notes: The authors
note that the type of roost used by sage-grouse varies with snow conditions and identify
two distinct burrow types: "snow forms" and "snow burrows", the first being shallow
depressions or open bowls in the snow. They observed 83 snow burrows at a number
of locations; in the majority of cases, the burrowing occurred within one week of
snowfall. Minimum temperatures during the periods of snow-burrowing use were <-10°C
in all but one case. The type of burrow varied with snow conditions. Drift burrows were
used when drifts provided the only deep snow in which to burrow. Open-snow burrows
only were observed when soft, dry snow >25 cm was available. In over 75% of the
precipitation events, the daily minimum temperature was at least 10°C lower for several
days following precipitation. Increased loses of hazel grouse and black grouse have
occurred when snow cover was lacking and temperatures were extreme. Key words:
sage-grouse, snow, temperature, behavior

017

Baines, D. 1996. Seasonal variation of lek attendance and lekking behaviour by
male black grouse Tetrao tetrix. Ibis 138(2): 177-1 80. Abstract: Eleven black grouse
Tetrao tetrix leks in the Scottish Highlands were visited at dawn once every week for a
year. Apart from July and three leks which were unoccupied in winter, males visited leks
throughout the year. Total numbers of males attending leks peaked in March in one
study area (six leks) and in April in the second (five leks). The number of males present
and the proportion of leks occupied varied seasonally. The proportion of males at leks

J*
4

^ fcfc t*

cCi

was at a maximum in April (80% of males) when all leks were attended, with a
secondary peak in September when 60% of leks had males present. Males spent the
most time displaying in April. The optimal conditions for attendance were calm, dry
mornings just after dawn. Most reliable counts of numbers of males were those made
between the last week of March and the end of the first week of May. Key words:
black grouse, wind, precipitation, behavior

018

. 1991. Factors contributing to local and regional variation in black grouse

breeding success in northern Britain. Ornis Scandinavica 22(3):264-269. Notes:
"Differences in gamebird chick mortality between years have been frequently explained
by extrinsic factors such as cold, wet weather or shortage of insect food. Findings from
this and other studies have led me to produce three hypotheses regarding factors which
could reduce black grouse breeding success and which may contribute to the decline in
both numbers and range of black grouse recorded in Britain: (1) A tendency for wetter,
cooler weather in June; (2) an increase in the number of egg and chick predators; and
(3) a decline in the abundance of preferred insect foods. Adverse weather will tend to
increase the need for brooding at the expense of foraging time. Cold, hungry chicks are
often noisy and may attract predators, whilst hens with reduced foraging time may have
to feed more intensively and as a result be less vigilant. Ground predators that hunt by
scent may even be more effective after rain. In June 1989, the weather at and after
hatching was warm and dry. Chick survival was high and production was 2.7 juveniles
per hen. But in the cold, wet June of 1990, more hens were broodless and only 1 .7
juveniles per hen were reared, a 37% reduction. Key words: black grouse,
temperature, precipitation, population dynamics, behavior, predation, mortality,
productivity, chick survival

019

, P. Warren, and M. Richardson. 2007. Variations in the vital rates of black

grouse Tetrao tetrix in the United Kingdom. Wildlife Biology 13 Supplement
1:109-116. Abstract: In the United Kingdom, black grouse Tetrao tetrix are in severe
decline with only 6,500 displaying males in 1995-1996 and a range retraction of 28%
between 1972 and 1991. Recent declines have been greatest in central and southern
Scotland and parts of Wales and contrast with relative stability in northern England. We
compare the demography of black grouse in three regions: North Wales, northern
England and the Scottish Highlands. Patterns in annual fecundity, measured as
fledglings per breeding female, were correlated between regions, suggesting that
annual weather patterns common across regions may be a key determinant of breeding
success. Site related effects such as habitat quality or management were also
significant in northern England and North Wales. Male population growth rates at leks
were positively correlated with fecundity in the previous year. Fecundity was highest in
North Wales and the Scottish Highlands at 1.7 chicks per female in August compared to
1.3 in northern England. Variations in the annual fecundity of radio-tagged females were
linked to differences in brood survival rather than clutch survival, which did not differ
among years. We found a non-significant trend for juvenile survival to be lower in North
Wales (0.18) than in either northern England (0.65) or the Scottish Highlands (0.56).

4

)X fc* **

Similarly, annual adult survival also tended to be lower in North Wales (0.44) than in
either northern England (0.70) or the Scottish Highlands (0.66). Predation was the main
cause of death in all regions, with red fox Vulpes vulpes and raptors being the chief
predators in North Wales and the Scottish Highlands and stoat Mustela erminea in
northern England. The last 10 years have seen the implementation of a series of black
grouse recovery projects in the UK. An understanding of the limiting demographic stage
in each project area is critical before appropriate remedial management prescriptions
can be implemented. Key words: black grouse, fecundity, weather patterns

020

Bakken, G. S. 1990. Estimating the effect of wind on avian metabolic rate with
standard operative temperature. Auk 107:587-594. Abstract: I developed a simplified
procedure to estimate the effect of wind on avian energy use rates from published and
unpublished studies of 10 passerine and 7 nonpasserine species. Below the lower
critical temperature, energy-use rates of passerines resting in the dark can be estimated
as that of a bird of the same species in a metabolism chamber set to the standard
operative temperature, Tes of the habitat, defined as Tes = Tb - (1 + 0.26Vu)(Tb - Te).
Wind speed is u, and Tb is body temperature. Operative temperature (Te), is ordinarily
close to air temperature for birds resting at night, but Te can include the effects of
thermal and solar radiation. The 95% confidence interval for predictions of the average
metabolic rate of a passerine is ±9.3% for air speed up to 4 m/s and temperatures
below the thermal neutral zone. The procedure also appears valid for some, but not all,
nonpasserines. Key words: ruffed grouse, Gambel's quail, wind, temperature,
metabolism, technique

021

Ballard, W. B. and M. C. Wallace, editors. 2005. Changes in land use patterns and
their effects on Rio Grande turkeys in the Rolling Plains of Texas and southwest
Kansas. Final Report to the Texas Parks and Wildlife Department and the National
Wild Turkey Federation, Texas Tech University, Lubbock, USA. Notes: This report
contains a discussion of the effects of precipitation and cover on Rio Grande turkey
nesting ecology in the Texas panhandle and southwestern Kansas. It states, in part,
"No relationship was found between pre-nesting precipitation and nesting rate of first
attempts, or nesting success. Nesting season precipitation had no relationship to
nesting success, nor did the number of days with precipitation during nesting had no
relationship to nesting success. Days since precipitation for depredated nests did not
conform to any distribution pattern. However, almost 30% of nests were depredated on
days with measurable precipitation. Managers can use local weather data to determine
the probability of nests being successful; this may allow for estimates of recruitment into
the fall populations. Managers should develop habitat management plans that maximize
visual obstruction available for potential nesting habitat." Key words: turkey,
precipitation, vegetation, habitat use, predation, population dynamics, habitat
manipulation

10 jj

*4*

f \ ^ ^

<£^>

022

Barnett, J. K. and J. A. Crawford. 1994. Pre-laying nutrition of sage-grouse hens
in Oregon. Journal of Range Management 47(2):1 14-1 18. Notes: "Production of
sage-grouse, measured by chicks/hen and average brood size, decreased in 1991. This
corresponded to a decrease in forbs eaten and decrease in nutrient content of the
composite diet. Many factors contribute to successful reproduction, including predation,
weather, and nutrition. Our study revealed that forbs are an important source of protein
and other nutrients, and consumption of forbs increased nutritional status of the hens.
Differences in amounts of forbs consumed between 1990 and 1991 likely was related to
decreased availability of forbs between the 2 years. This change in abundance of sage-
grouse foods perhaps was related to the 40% reduction from normal precipitation for the
year preceding the 1991 sample. Key words: sage-grouse, precipitation, vegetation,
nutrition, habitat use

023

Battazzo, A. M. and E. Greene. 2006. Winter survival and habitat quality of greater
sage-grouse in south Phillips County, Montana. Annual Conference Wildlife
Society 13. Abstract: Factors influencing sage-grouse (Centrocercus urophasianus)
populations vary across the species/ range, requiring conservation strategies to be
developed from geographically-specific data. We studied two populations of sage-
grouse hens in Phillips County, Montana to understand how winter conditions and
habitat quality influence sage-grouse winter survival rates and habitat use. We radio-
collared 148 hen sage-grouse and located individuals one to three times per week.
Observations took place from December 15-March 15 during 2005 and 2006 winter
seasons. We use locations to generate population-level winter home ranges using
minimum convex polygon (MCP) and kernel techniques. Home ranges were stratified
into low, medium, and high-use categories, and habitat quality (defined as percent
sagebrush cover and protein composition) was assessed in each strata. If we observed
high winter survival, we predicted fitness consequences would be apparent in the early
stages of reproduction. Our analyses examined associations between climatic
(temperature and wind speed), habitat (sagebrush cover and protein composition),
group (density and study site), and individual (age and sex) covariates and winter
survival, nest initiation, and clutch size. Results from analysis of population-level winter
range habitat quality were compared with current conservation recommendations. We
tested the hypothesis that levels of use (low, medium, and high) ordinally follow levels
of habitat quality (poor, marginal, and good), and investigated the effects of climatic
conditions on winter movements and distribution. We will summarize our results and
discuss management implications with respect to sage-grouse winter habitat
conservation. Key words: sage-grouse, temperature, wind, vegetation, habitat use,
reproduction, movement, distribution

024

Baumann, D. P. Jr., W. E. Mahan, and W. E. Rhodes. 1995. Effects of Hurricane
Hugo on the Francis Marion National Forest wild turkey population. Proceedings
National Wild Turkey Symposium 7:55-60. Abstract: The Francis Marion National
Forest (FMNF) is an important area for wild turkeys {Meleagris gallopavo silvestri) in

% fVfc*^

11

^k -'car- ^^ c^>

South Carolina. On 21 September 1989, Hurricane Hugo, a category IV storm, struck
the South Carolina coast, and the strongest winds swept across the FMNF. Over 1
billion board feet of timber were damaged or destroyed. To determine the effects of
Hurricane Hugo on the FMNF wild turkey population, we examined spring turkey
harvest and reproduction pre- and post-Hugo. Prior to Hugo, the spring turkey harvest
increased at a mean annual rate of 18% on the FMNF and 25% statewide. Following
the storm, the harvest declined 22% per year on the FMNF, whereas the statewide
spring harvest increased 4% annually. Mean number of hens with poults (P = 0.07),
brood size (P = 0.008), gobblers observed (P = 0.018), total turkeys observed (P =
0.01 1 ), and recruitment ratio (P = 0.006) have declined since the storm. The negative
habitat alterations from Hurricane Hugo that occurred on the FMNF were responsible
for the decline in the wild turkey population. Key words: wild turkey, wind, habitat
modification, productivity, reproduction

025

Baumgartner, F. M. 1939. Studies on the distribution and habits of the sharptail
grouse in Michigan. Transactions of the North American Wildlife Conference
4:485-490. Notes: Page 486 — "The birds generally roost fairly close together in snow
beds beneath scattered trees or clumps of bushes and brush... During mild weather the
birds seem to prefer to feed in areas where green herbaceous plants, persistent fruits,
or grain are not covered over by deep snow; during stormy weather or on extremely
cold days sharptails have been found to retire to the edge of white birch and aspen
thickets. At such times their diet appears to consist entirely of buds and catkins. The
period of daily activity has revealed some surprising variations dependent upon weather
conditions. On bright, warm days, sharptails were observed to leave their snow roosts
shortly after sunrise, spend a few minutes in a sketchy courtship display, and then fly to
feeding grounds. By the middle of the morning, after their crops were stuffed, they either
bury themselves beneath the snow or sit out in some open spot, apparently to enjoy the
rays of the sun. Active feeding is resumed again in mid-afternoon and lasts until dusk.
The contrast between this period of 8 to 10 hours and the very limited activity on
extremely cold or stormy days is quite marked. On such days the sharptails do not
leave their roosts until 8:00 to 9:00 a.m. Immediately they fly to a nearby feeding station
or grain field or commence to fill their crops with birch or aspen buds. By 9:30 or 10:00
a.m. they are back into their snow forms for a 22-hour period of inactivity not emerging
again until the following morning." Page 487 — "During the heat of the day and at night
the cocks and hens spend most of their time feeding or resting in the edges of willow
thickets or beneath the aspens and tag alders along the swamp and stream borders."
Page 488 — "We have found that the birds are extremely difficult to flush during the heat
of the day. However, in the early morning and again toward evening small flocks feed
on the plants and fruits of strawberry, the leaves of clover, dandelions, and other herbs,
and to a considerable extent on insects... [when] "heavy snow and stormy weather in
December and January make it impossible for the grouse to obtain grain and weed
seeds or sufficient quantities of green plants... they move to the white birth and aspen
thickets and take up a diet of buds and catkins for the winter." Key words: sharptailed
grouse, snow, temperature, solar radiation, behavior, habitat use, census

12 j,

4-

H fc* **

026

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, USA.

Notes: Page 18 - "Factors that cause changes in nesting chronology - Climate:
Regression analyses were performed using temperature and precipitation
measurements in the following forms: monthly mean, monthly high, monthly low, and
monthly deviation from normal. All of these measurements were shown to exert an
influence on the mean date of hatch, but the clearest example of this influence was
noted in a multiple regression of the total deviation from normal of temperature and
precipitation for the months of February, March, April, May, and June. In this analysis, a
significant rvalue (0.619) was obtained. Warmer temperatures were associated with an
earlier mean hatching date, while above normal precipitation delayed hatching. Using a
standard partial regression analysis on the above rvalue, it was calculated that 33.5
percent was attributable to temperature, while 66.5 percent was attributable to
precipitation. It appeared from this analysis that in south-central Nebraska, seasonal
precipitation deviation from normal had more effect on the mean date of hatch than did
temperature deviation from normal. The earliest hatching peak occurred during a year
classified by the Weather Bureau as "warm and dry", while the latest occurred during a
"cold and wet" year. No relationship between degree-days and mean date of hatch was
shown using analysis of variance for the regression (P>.05). A non-significant F value
(P>.05) using analysis of variance for the regression indicated that no relationship
existed between percent of possible sunshine (Lincoln Station) and mean date of
hatch." Key words: wild turkey, temperature, precipitation, reproduction

027

Beasom, S. L. 1973. Ecological factors affecting wild turkey reproductive success
in south Texas. Dissertation, Texas A&M University, College Station, USA. Unable
to obtain for abstract.

028

and O. H. Pattee. 1980. The effect of selected climatic variables on wild

turkey productivity. Proceedings National Wild Turkey Symposium 4:127-135.

Abstract: Net reproductive success of Rio Grande wild turkey (Meleagris gallopavo
intermedia) was monitored yearly between the months of June and August from 1968
through 1977 on 2 study locations in south Texas by way of repetitive road transect
counts. Rainfall and temperature data were compiled from U. S. Climatological Records
and soil moisture data were calculated from these parameters. Between-year turkey
production fluctuated from 0-576 poults per 100 hens on the Santa Gertrudis area near
Kingsville and from 8-443 on the Encino area near Falfurrias. The maximum R2
determination indicated that 97.3% of the variability in annual rate of turkey productivity
on Santa Gertrudis was explainable by a model composed of soil moisture storage the
previous August, combined total precipitation for the previous September and October,
and total precipitation for the current March. Similarly, 98.3% of this variability on Encino
was explainable by a model composed of combined total precipitation for the previous
August and September, soil moisture storage in the previous September, and soil

*""* <o t*

X i_~,

<£^>

moisture storage in the current May. Key words: Rio Grande turkey, precipitation,
soil moisture, productivity

029

Beck, T. D. 1. 1977. Sage grouse flock characteristics and habitat selection in
winter. Journal of Wildlife Management 41(1):18-26. Abstract: Sage grouse
(Centrocercus urophasianus) were studied in North Park, Colorado, during the winters
of 1973-74 and 1974-75. Distribution was plotted from sightings of 199 flocks and 17
single birds, totaling 5,080 grouse. Only 50 percent of the 1,252 km2 of lands dominated
by sagebrush (Artemisia spp.) sustained winter use by grouse because of snow depth,
steepness of slope, and sagebrush disturbance. Nearly 80 percent of the use occurred
in 7 areas comprising less than 7 percent of the total area. Sexes segregated; males
formed more unisexual flocks. Flocks were the dominant social unit and contained less
than 50 individuals in 88 percent of all observations. Flocks containing more than 50
percent females were larger than male flocks and used denser sagebrush stands for
feeding and loafing. Roosting and feeding sites had similar vegetal and physical
characteristics. Sixty-six percent of flocks were on slopes less than 5 percent, and only
13 percent were on slopes greater than 10 percent. Sixty-two percent of 2,350 grouse in
1973-74 and 61 percent of 1,984 grouse in 1974-75 were females. Key words: sage
grouse, snow, habitat use

030

Bell, L. A., S. D. Fuhlendorf, and M. A. Patten. 2003. Brood survivorship in relation
to microclimate/habitat use of lesser prairie chickens. Proceedings of the Prairie
Grouse Technical Council 25:16. Abstract: Chicks have a high surface-area-to-
volume ratio, so maintaining their internal body temperature can be difficult. Thus, apart
from predation, relative humidity, ambient temperature, and exposure to wind are
factors that influence survival of chicks. We studied the relationship between
survivorship and microhabitat use for lesser prairie chicken {Tympanuchus
pallidicinctus) broods from hatch to reproductive stages. Results will indicate the effect
heat stress and other variables have on brood movements, microhabitat selection, and
survivorship Key words: lesser prairie chicken, thermoregulation, humidity,
temperature, wind, heat stress, movement, habitat use, chick survival

031

Bendell, J. F. 1955. Disease as a control of a population of blue grouse,
Dendragapus obscurus fuliginosus (Ridgway). Canadian Journal of Zoology
33(3):1 95-223. Abstract: A population of blue grouse was studied on its summer range
at Quinsam Lake, Vancouver Island, to determine the factors of importance in
population control. The population was stable with a density of 0.40 adult males and
0.78 yearling and adult females to the acre. A life table was constructed from band
returns on the basis of a stable population. This disclosed that 80% of the chicks died in
the first three months after hatching and that the death of 31% of the adults occurred
each year. The survivorship curve was negatively "J"-shaped. Mortality rates appeared
constant and independent of age after the first year of life. Space, weather, food,
predators, and disease are considered as factors capable of population control. Six

14 j^

4

H fa* ^

c^

parasites were recorded as new for this host. Two, Plagiorhynchus formosus and
Dispharynx nasuta, occurred commonly and almost exclusively in the chicks, where
they caused extreme damage to gut tissues. Parasitism by these helminthes was an
important mortality factor in chicks and a major cause of population stability. Key
words: blue grouse, weather, disease, mortality

032

and L. I. Bendell-Young. 2006. Eggs of spruce grouse dry at a faster rate

than those of ruffed grouse. Canadian Journal of Zoology 84:1688-1692. Abstract:
We measured the rate of water loss and pore density of eggs of spruce grouse
(Canachites canadensis canace (L, 1766)) and ruffed grouse (Bonasa umbellus togata
(L, 1766)) from different parts of their range in Ontario. Eggs were dried in enclosed
glass jars over Drierite® and in paper trays in open air at room temperature and
humidity. Eggs were weighed to the nearest 0.01 g every 2-4 days and change in mass
was measured as water loss. Pores of shells were counted (pores/cm2) in the blunt,
middle, and pointed sections of the egg. Eggs of spruce grouse lost water at a fast rate
in Drierite® and in open air and had a greater density of pores than eggs of ruffed
grouse. Rates of water loss were constant and varied inversely with ambient humidity,
with the difference between species greatest in open air. Eggs late in incubation of
ruffed grouse dried at a faster rate than those early in incubation in Drierite®. The
adaptation of eggs of each grouse to the moisture of the nest may help explain their
distribution, density, and habitat and nest-site selections, as well as behavioral aspects
of the nesting hen. Both, especially the spruce grouse, may be good indicators of
climate change. Keywords: spruce grouse, ruffed grouse, humidity, temperature,
index

033

Bennitt, R. and H. V. Terrill. 1940. Possible temperature factors in north central
pheasant distribution. Transactions of the North American Wildlife Conference
5:428-432. Notes: Page 429— It seems very likely that lethal temperatures may occur
under natural conditions. The egg temperature in ground-nesting birds would depend
most upon the temperature of the ground and the degree of exposure to the sun... it is
possible that the pheasant's occasional practice of leaving the nest during the hottest
part of the day may raise the temperature of the eggs if the air and ground temperatures
are unusually high. Key words: ring-necked pheasant, temperature, solar radiation,
behavior, mortality

034

Berry, J. D. and R. L. Eng. 1985. Inter-seasonal movements and fidelity to
seasonal use areas by female sage-grouse. Journal of Wildlife Management
49(1 ):237 -240. Notes: "Although snowstorms appeared to induce migration to wintering
areas, movement was often completed in advance of snow accumulation that would
cover food resources. One hen moved onto the wintering area, returned to the
summering area, and moved back toward the wintering area before finally returning to
the breeding ground. This uncharacteristic midwinter return to a summering area was
associated with moderating winter weather. It is possible that extrinsic factors, primarily

H ^ ^

M Ajv i. ..l. 15

c^

moisture conditions and the resulting distribution of succulent vegetation, played a
major role in summer distribution, but only after the initial brood rearing phase is
completed. Migration to the winter range was observed in the absence of severe winter
conditions. The annual return to specific wintering areas by adult hens, regardless of
apparent weather or food conditions, may introduce recruits within a population to
traditional areas. When seasonal key areas for a population are separated to this
extent, numerous land owners are often involved. In such cases, agencies responsible
for wildlife resources are obligated to assess year-long rather that seasonal
requirements and to look at large ecologically defined land units rather than
economically or politically delineated parcels of land." Key words: sage-grouse, snow,
temperature, vegetation, movement, habitat use, habitat management

035

Blake, C. S. 1970. The response of sage-grouse populations to precipitation
trends and habitat quality in south central Idaho. Annual Conference Western
Association of State Game and Fish Commissioners 50:452-462. Notes: The author
states that a reduction in annual precipitation lead to a downward trend in sage-grouse
numbers lasting for ten years. "It was obvious that a reduction of succulent food plants
and early desiccation of vegetation was denying hens and broods the use of important
summer ranges. Reproductive success was low and for several years over 55% of the
hens observed on brood count routes were without young." Key words: sage-grouse,
precipitation, vegetation, nutrition, reproduction, drought

036

Blanchett, P., J.-C. Bourgeois, and S. St-Onge. 2007. Winter selection of roost
sites by ruffed grouse during daytime in mixed nordic-temperate forests, Quebec,
Canada. Canadian Journal of Zoology 85(4):497-504. Abstract: We determined the
categories of roost sites used by ruffed grouse (Bonasa umbellus (L, 1766)) during
daytime in winter from 245 radiotelemetric locations of 26 adult females. We conducted
our study in the Reserve Faunique de Portneuf, located in a mixed nordic-temperate
softwood-hardwood forest in Quebec, Canada. We evaluated the effects of weather,
snow, and habitat variables on the incidence of snow burrowing, tree roosting, and on-
snow roosting using mixed multinomial models, ANOVA, and logistic regressions. The
best logistic regression model of snow burrowing probability was identified using the
Akaike path. The incidence of each category of roost sites was 41 .2% tree roosts,
36.3% snow burrows, and 22.4% on-snow roosts. Coniferous canopy closure and depth
of fluffy snow were the variables that influenced roosting behavior the most. Probability
of snow burrowing increased with compaction depth and decreased with coniferous
cover. Probability of tree roosting increased with temperature. On-snow roosts had a
denser lateral obstruction than snow burrows, whereas tree roosts had a greater
coniferous basal area, stem density, and canopy cover than snow burrows. Stand type
also influenced the incidence of each category of roost sites, snow burrows dominating
in deciduous stands and tree roosts dominating in mixed and coniferous stands. Key
words: ruffed grouse, snow, temperature, behavior, habitat use, snow condition

16 j,

H fc* **

cCi

037

Boggs, C, E. Norris, and J. B. Steen. 1977. Behavioral and physiological
temperature regulation in young chicks of the willow grouse Lagopus lagopus.
Comparative Biochemistry and Physiology A 58(4):371-372. Abstract: The length of
brooding and foraging periods was measured for a brood of 8 grouse chicks and
parents kept in a 255 m2 open air enclosure. The chicks had shorter foraging periods
and longer brooding, the poorer the weather. Foraging periods became longer and
brooding shorter as the chicks grew older. At 8 days of age the heaviest chick was
thermally independent at ambient temperatures above 15°C. The duration of foraging is
apparently determined by the degree of hypothermia. Key words: willow grouse,
temperature, thermoregulation, foraging, behavior

038

Bohl, W. H. 1957. Chukars in New Mexico 1931-1957. Bulletin No. 6. New Mexico
Department of Game and Fish, Sante Fe, USA. Notes: Page 41 - Feeding, Weather
Tolerances: No adverse snow conditions have been experienced at three Turkish trial
areas during the study. From Turkey there is information that may illustrate what effects
heavy snow may have on the chukar should these conditions occur late in the trial
areas: 'Chukers stand winter snows well, so long as adequate food is available. If it is
not, they will move either to the edges of the villages or down into the lower valleys,
returning to higher ground in the spring. It is doubtful however, if they can survive any
extended periods of snow covered habitat if the snow remains for long to a depth
exceeding 8-12 inches, unless considerable areas are swept bare by wind, or on
southern exposures, are uncovered by the winter sun.' Key words: chukar partridge,
snow, wind, nutrition, movement, solar radiation

039

Boos, M., J. -P. Boidot, and J. -P. Robin. 2005. Body condition in the Eurasian
woodcock wintering in the west of France: practical study for wildlife
management during cold spells. Wildlife Biology in Practice 1(1):15-23. Abstract:
The Eurasian woodcock (Scolopax rusticola) is one of the most widespread species of
the Scolopax genus in temperate regions. However, population levels can be greatly
affected by harsh cold spells that lead woodcocks to starve by exhaustion of their body
fuels. To better understand the vulnerability of woodcocks to such climatic conditions it
is of a major importance to determine the amount of their body reserves (both lipids and
proteins) throughout the wintering season. This was performed on 55 individuals
collected by hunters in the western part of France during two consecutive winters under
mild weather conditions. Body reserves, that can be mobilized, were determined as the
difference between the total amount of lipids and proteins minus the values obtained on
starved individuals found dead during previous cold spells. Overall, body reserves did
not significantly change over both winters (p>0.40), the maximal mean value (1539 [plus
or minus] 1 17 kJ) being however reached in January. Storing body fuels would not
adversely affect wing and power loading, suggesting that the amount of body reserves
would agree with the "starvation-predation trade-off". If woodcocks sit through a cold
spell, their mean survival time (or fasting endurance) would be 6.5 [plus or minus] 0.5
days; 25 to 40% of the birds would have a life expectancy of 7-9 days, and about 8 to

f v ^ ^

\ ***•

C*~S^-^J? <_ _b

L©£

17% less than 5 days. On the contrary, if woodcocks immediately leave their wintering
quarter, they would be able to perform a trip of 740 [plus or minus] 50 km, 20 to 40% of
the woodcocks being able to fly over 750 km. Body mass explains only 47 to 57% of the
fasting endurance and flight autonomy variations, and therefore we recommend a
further carcass analysis to accurately estimate body condition. These results
underscore the suitability of determining the state of body reserves for practical cases of
population management and hunting policy during cold spells. Key words: Eurasian
woodcock, temperature, mortality, nutrition

040

Botsford, L. W., T. C. Wainwright, J. T. Smith, S. Mastrup, and D. F. Lott. 1988.
Populaiton dynamics of California quail related to meteorological conditions.
Journal of Wildlife Management 52(3):469-477. Abstract: Demographic responses of
California quail (Callipepla californica) to precipitation-related variables vary among
locations with different mean rainfall. With 23 years of data from a California quail
population in a semiarid region, we determined a positive response of reproductive
success (juv/ad) to precipitation during the previous winter. Computed correlations with
soil moisture content and actual evapotranspiration were significant but not as high as
with precipitation. Correlations with mean monthly temperature, days/month <0C, and
days/month >38C were not significant. Attempts to account for possible lower
reproductive capability of first-year breeders did not improve statistical relationships.
The number of adults each year was positively correlated with the number of juveniles
in the previous year. The number of juveniles produced each year was correlated with
the number of adults in that year only when the effect of precipitation was removed; the
relationship was then linear. The interannual fluctuations in population numbers resulted
from low adult survival and the influence of precipitation on recruitment through
unknown mechanisms. Keywords: California quail, precipitation, fecundity,
temperature, recruitment

041

Boughton, R. V. 1937. Endoparasitic infestations in grouse, their pathogenicity
and correlation with meteoro-topographical conditions. Technical Bulletin 121,
University of Minnesota Agricultural Experiment Station, Minneapolis, USA.

Notes: Page 39 — "The third cause, meteorological and topographic factors, is in the
writer's opinion the most important. It is well known that the ova of many parasites when
outside the host require moisture before development can proceed. Lack of moisture,
on the other hand, reduces or prevents development, and after a limited time the ova
may fail to hatch even when placed in a suitable environment. In addition to moisture,
other important factors affecting the percentage of ova that hatch are temperature, type
of soil, and vegetation.... These experiments show... that the development of nematode
ova depend upon a number of physical factors, the most important of which are
moisture, type of soil, temperature and coverage.... The important factor with regard to
the soil is its ability to retain or lose water." Page 46 — "In the case of the ruffed grouse,
a definite correlation between the degree of parasitism and the meteorological and
topographical factors appear to occur. Infestation in the different zones outlined appears
to depend upon the mean temperature, mean precipitation, soil type, and coverage, the

18 \^

i

>->' kr* **

maximum infestation taking place where the mean temperature and precipitation are
highest, where the humus content of the surface soil is greatest, and where abundant
coverage occurs... That the parasites have played a secondary part in the death of the
birds is probable. Accompanying the increase in infestation, there has been a definite
increase in weight difference between parasitized and non-parasitized birds. The
parasites may have played a part in reducing the vitality of the birds to a point where
secondary diseases could set in." Key words: grouse, ruffed grouse, temperature,
precipitation, topography, soil, solar radiation, vegetation, disease, mortality,
population dynamics, parasites

042

Bousquet, K. R. and J. J. Rotella. 1998. Reproductive success of sharp-tailed
grouse in central Montana. Prairie Naturalist 30(2):63-70. Abstract: We estimated
nest success and survival of broods and chicks for plains sharp-tailed grouse
(Tympanuchus phasianellus jamesi) on a large grassland in central Montana. We
collected productivity data from 24 different radio-marked females during 1994-95. Nest
success varied between years (P = 0.02) and was high (0.93, n = 12) in 1994 when
yellow sweetclover (Melilotus officinalis) was dense and moderate (0.56, n = 18) in
1995. Mean hatching date was 18 June, and the average successful nest produced
1 1 .3 chicks/nest (SE = 1 .0, n = 21 nests). Chick survival to 56 days of age varied by
year (P = 0.05) and was higher in 1994 (0.44, SE = 0.15) than in 1995 (0.09, SE = 0.10)
when extensive, cold, wet weather was suspected of causing high chick mortality. Key
words: sharp-tailed grouse, temperature, precipitation, mortality, chick survival

043

Bradbury, J. W., S. L. Vehrencamp, and R. M. Gibson. 1989. Dispersion of
displaying male sage grouse I. Patterns of temporal variation. Behavioral Ecology
and Sociobiology 24(1):1-14. Abstract: The distribution of lek sizes was examined in
each of three populations of sage grouse in eastern California [USA]. Peak seasonal lek
sizes collected over a 35-year period were found to co-vary among the three sites
indicating that some global environmental or demographic features modulated male
attendance in any given year. Despite these annual variations, the ranks of the three
populations with regard to mean lek size remained stable. In all three populations, there
was a persistent excess of small and large leks, compared to random settlement on the
same number of sites, and a consistency in the ranking by size of particular sites in
successive years. The sequential phenology of lek site occupation in each population
was correlated with recolonization of habitats surrounding central wintering refuges
each spring. Some lek sites utilized for display in early spring were regularly abandoned
prior to the onset of mating as more peripheral leks became active. On top of
population, site, and seasonal variations in lek size, pronounced daily fluctuations in
attendance were common. Multivariate regressions indicated than an average 36% of
the daily variation in male numbers was correlated with weather variables, female
attendance levels, and prior raptor harassment. Several outcomes of the analyses
support the notion that dispersion of males is partly determined by male settlement on
current female traffic patterns (hotspot settlement). The analyses also suggest that
display is sufficiently costly that variations in male attendance are in part a result of

'% fv^t*

19

§®>\ fyjmjfc ^^

conflicts between strutting and thermoregulatory expenditures. Key words: sage
grouse, temperature, thermoregulation, behavior, movement

044

Braun, C. E., J. W. Connelly, and M. A. Schroeder. 2005. Seasonal habitat
requirements for sage-grouse: spring, summer, fall and winter. Pages 38-42 in N.
Shaw, M. Pelland, and S. B. Monsen, compilers. Sage-grouse habitat restoration
symposium, Boise, Idaho. RMRS-P-38, Rocky Mountain Research Station, Fort
Collins, USA. Abstract: Sage-grouse (Centrocercus minimus, C. urophasianus) are
dependent upon live sagebrush {Aremisia spp.) for all life processes across their entire
range. This paper describes habitats used by sage-grouse as documented in the
scientific literature. The leaves of sagebrush are eaten by sage-grouse throughout the
entire year and comprise 99 percent of their winter diets. Spring (late March through
May) habitats are those with intermixed areas of taller (40 t 80 cm) sagebrush with
canopy cover of 15 to 25 percent and taller (>18 cm) grass/forb cover of at least 15
percent. Sites used for display have shorter vegetation, frequently few or only short
sagebrush plants, but with taller, more robust sagebrush within 100 to 200 m that is
used for escape cover. Nesting cover mimics that used overall during spring but with
clumps of tall (> 50 cm), dense (about 25 percent) live sagebrush and abundant forbs (>
10 to 12 percent cover). Early brood rearing areas are those within 200 m (initial 3 to 7
days posthatch) to 1 km (up to 3 to 4 weeks posthatch) of nest sites. Forbs and taller (>
18 cm) grasses are important for broods; forbs provide succulent foods, grasses
provide hiding cover, and the grass/forb mixture supports insects used by chicks.
Summer use areas are those with abundant succulent forbs with live, taller (> 40 cm),
and robust (10 to 25 percent canopy cover) sagebrush useful for cover. These areas
continue to be used into fall when sage-grouse move to higher benches/ridges where
they forage on remaining succulent forbs such as buckwheat (Eriogonum spp.) and
switch to more use of sagebrush leaves. Winter (early December to mid-March) use
areas are often on windswept ridges, and south to southwest aspect slopes as well as
draws with tall, robust live sagebrush. Height (25 to 35 cm) of sagebrush above the
surface of the snow in areas used in winter is important, as is canopy cover (10 to 30
percent). Management of habitats used by sage-grouse should initially focus on
maintaining all present use areas. Practices to enhance sagebrush habitats to benefit
sage-grouse are reviewed, as is the need to annually monitor sage-grouse numbers
along with systematic monitoring of the health of sagebrush ecosystems. Key words:
sage-grouse, snow, vegetation, nutrition, habitat use

045

, R. W. Hoffman, and G. E. Rogers. 1976. Wintering areas and winter

ecology of white-tailed ptarmigan in Colorado. Special Report Number 38,
Colorado Division of Wildlife, Denver, USA. Abstract: Aspects of the ecology of
white-tailed ptarmigan (Lagopus leucurus) in winter were periodically investigated in
Colorado from 1964 to 1975. Ptarmigan were found to select areas dominated or co-
dominated by willow (Salix spp). Areas utilized in winter most frequently were drainage
basins at or above treeline and stream courses below treeline from 2,591 to 3,810 m
elevation (8,500 to 12,500 ft) where food (willow) and roosting sites (soft snow) were

20 ^

<±

f > *c« >f*

^1/^

readily available. Partial sex segregation occurred from late October through mid-April,
with males wintering at higher elevations, frequently krummholz areas at treeline.
Flocks of females most commonly selected large drainage basins and stream courses.
Arrival and departure were weather-related and varied with year. Flock size was stable
among months and years, with male flocks being smaller in number (<15 birds) than
female flocks (20 to 40 birds). Within individual wintering areas, essentially all sites
were utilized each winter, with use depending upon snow depth and wind action. Adult
ptarmigan exhibited greater affinity for wintering areas where they were banded than did
subadults. Wintering areas have been altered and reduced in size through reservoir
construction, development of roads, mining activity, domestic livestock grazing, and
increased recreational use and development. Maps of known wintering areas are
presented. Key words: white-tailed ptarmigan, snow, wind, habitat use, movement

046

, J. W. Connelly, and M. A. Schroeder. 2001. Seasonal habitat requirements

for sage-grouse: spring, summer, fall, and winter. Pages 38-42 in N. L. Shaw, M.
Pellant, and S. B. Monsen, compilers. Sage-grouse habitat restoration
symposium proceedings, Boise, Idaho. Notes: Page 40 — "Sagebrush height
(>20cm, but usually >30 cm, above the surface of the snow) is important as is the
robust structure of live sagebrush. Sage-grouse use a variety of sites in winter including
windswept ridges with open stands of sagebrush to draws with dense stands. Quality of
the snow can be important because sage-grouse are known to use snow roosts and
burrows. Aspect is also important with south and southwest slopes most used in hilly
terrain. Leaves of live, vigorous sagebrush plants provide >99% of the foods eaten
during the winter period (early December until early to mid-March). Generally, winter is
a time of body mass gain, although severe winter conditions over prolonged intervals
can reduce the amount of area available for foraging and cover. Overall movement
during winter may be extensive and home ranges can be large. Key words: sage-
grouse, snow, wind, topography, habitat use, vegetation, nutrition

047

and R. K. Schmidt Jr. 1971. Effects of snow and wind on wintering

populations of white-tailed ptarmigan in Colorado. Pages 238-250 in A. O.
Haugen, editor. Proceedings of the snow and ice in relation to wildlife and
recreation symposium, Iowa State University, Ames, USA. Abstract: Studies of
wintering populations of white-tailed ptarmigan (Lagopus leucurus) were initiated in
Colorado in the fall of 1966 and were continued through December 1970. Objectives
were to describe the habitat and environmental features typical of major wintering
areas, determine composition of winter flocks, and delineate behavior and movements
of individual birds and flocks in relation to changes in environmental conditions.
Ptarmigan were found to prefer sites where willow (Salix spp.) was a major component
of the vegetation. Partial segregation of sexes occurred in winter with males preferring
higher, more windswept areas than females. Willow density on upland sites was sparse,
and availability was a result of wind action. Female ptarmigan preferred lower areas
near or below tree line where tall, dense willow stands occurred. While willow was the
most important component of wintering areas, wind and snow depth were most

H fc* **

,>■/.*

~- <C>^j^~^^Jp C j

important in determining availability. Snow quality was of extreme importance to
ptarmigan in winter, with most overnight roosts completely under the snow surface.
Ptarmigan were observed to move up to 7 miles from fall to winter use sites and up to 1
mile from feeding areas to sites with snow suitable for roosting. Key words: white-
tailed ptarmigan, snow, wind, nutrition, habitat use, behavior, vegetation,
movement, snow condition

048

, T. Britt, and R. O. Wallestad. 1977. Guidelines for maintenance of sage-
grouse habitats. Wildlife Society Bulletin 5(3):99-106. Notes: The authors stated the
extent of seasonal movements in sage-grouse vary with the severity of winter weather,
topography and vegetative cover. They note that during winter, sage-grouse may be
restricted to less than 10 percent of the sagebrush-dominated lands within a given area.
Suitability of sagebrush manipulation is closely linked to seasonal precipitation and forb
production Key words: sage-grouse, precipitation, habitat manipulation, habitat
use, nutrition

049

Braunisch, V. and R. Suchant. 2007. A model for evaluating the 'habitat potential'
of a landscape for capercaillie Tetrao urogallus: a tool for conservation planning.
Wildlife Biology 13 Supplement 1:21-33. Summary: Most habitat models developed
for defining priority conservation sites target areas currently exhibiting suitable habitat
conditions. For species whose habitats have been altered by land use practices, these
models may fail to identify sites with the potential of producing suitable habitats, if
management practices were modified. Using capercaillie Tetrao urogallus as an
example, we propose a model for evaluating the potential of ecological conditions at the
landscape level to provide suitable habitat at the local scale. Initially, we evaluated the
influence of selected landscape parameters on the structural characteristic of vegetation
relevant to capercaillie. Then we used capercaillie presence data and an ecological
niche factor analysis (ENFA) to identify landscape and land use variables relevant to
capercaillie habitat selection. We also studied the effect of scale on predictive model
quality. Despite high variance, correlations between landscape variables and forest
structure were detected. The greatest influence on forest structure was recorded for
climate and soil conditions, which were also found to be the best predictors of
capercaillie habitat selection in the ENFA. The final model, retaining only two landscape
variables (soil conditions and days with snow) and three land use variables (proportion
of forest, distance to roads and forest-agricultural borders), explained a high degree of
capercaillie habitat selection, even before considering patch size and connectivity. By
restricting the analyses to areas with stable subpopulations and a set of relatively stable
landscape variables capable of explaining habitat quality at a local scale, we were able
to identify areas with long-term relevance to conservation of capercaillie. Key words:
capercaillie, model, snow, conservation

22 _tf

4-

>'>" <o >f*

c£^b

050

Bridges, A. S., M. J. Peterson, N. J. Silvy, F. E. Smeins, and X. B. Wu. 2001.
Differential influence of weather on regional quail abundance in Texas. Journal of
Wildlife Management 65(1):10-18. Abstract: Although weather variables are known to
influence quail abundance in some habitats, most studies have addressed only limited
geographic areas and indices to weather conditions. The few replicated studies
addressed relatively similar climate zones. We used 21 years (1978-98) of quail
abundance data collected by the Texas Parks and Wildlife Department (TPWD)
biologists to address the relationship between both simple precipitation and Palmer
drought indices and northern bobwhite (Colinus virginianus) and scaled quail (Calipepla
squamata) abundance in 6 ecological regions of Texas. Three 12-month Palmer indices
were more highly correlated with changes in northern bobwhite abundance in the South
Texas Plains ecological region than was raw precipitation alone. The 12-month Modified
Palmer Drought Severity Index (PMDI) was correlated (rs > 0.78, P < 0.001 ) with the
mean number of northern bobwhites visually observed per survey route in the Rolling
and South Texas Plains ecological regions, while a 12-month, raw precipitation index
was correlated (rs = 0.64, P = 0.002) with northern bobwhite abundance in only the
South Texas Plains. The PMDI and raw precipitation were correlated (rs = 0.67, P <
0.001 and rs > 0.57, P < 0.007, respectively) with the mean number scaled quail
observed per survey route in the Edwards Plateau, South Texas Plains, and Trans-
Pecos Mountains and Basins ecological regions. There was no relationship (P> 0.437)
between changes in quail abundance and the PMDI or raw precipitation in the Gulf
Prairies and Marshes physiographic region, where precipitation was relatively high. The
monthly PMDI was a better indicator of changes in both northern bobwhite and scaled
quail abundance among years than was monthly precipitation alone. Both monthly and
12-month precipitation-based weather indices were more correlated with changes in
northern bobwhite and scaled quail abundance among years in relatively dry as
opposed to wet ecological regions. Our approach should help wildlife biologists and
managers better account for annual variability in quail productivity in semi-arid
environments so that long-term population trends can be better elucidated. Key words:
bobwhite quail, scaled quail, drought, index, precipitation, population dynamics

051

Bridgman, C. L. 2002. Habitat use, distribution and conservation status of the
mikado pheasant (Syrmaticus mikado) in Taiwan. Dissertation, University of
Tennessee, Knoxville, USA. Abstract: To evaluate the conservation status of Taiwan's
Mikado pheasant, Syrmaticus mikado (Phasianidae), I test the similarity of preferred
habitat to primary and secondary forest, develop models of habitat availability within
Taiwan, and examine population trends within two locations inside Yushan National
Park. The characteristics of locations with pheasant activity were most similar to
secondary forest: high shrub stem counts and low canopy and leaf litter coverage. None
of these variables were applicable to geographic information systems analysis. To the
known extent of range and area of occupancy, I compared a model based on the
habitats described in field guides. This model underestimated extent of range,
confirmed that 39% of the pheasant's range is protected inside parks and reserves, and
suggested that Taiwan potentially has 6477 km2 of habitat available to S. mikado.

% Kfc*>f*

23

% m <& 9

Within Yushan National Park, there may be as many as 10,000 S. mikado. The small
home ranges (<0.86 km2) and the lack of movement across the 400 m separating the
study sites imply limited gene flow between populations and poor ability to colonize
suitable habitat. There were 58 pheasants per km2 in the primary forest site and 48 in
the secondary forest sites. At the primary forest site, the population appeared stable. At
the secondary forest site, the population declined 65% from 3.56 pheasants
encountered per day in 1989-1992 to 1.24 in 1996-1999 due to poor productivity
because of increase numbers of nest predators (from 1 .4 predators per day to 2.2) and
the cooler weather conditions during hatching. In 1989-1991, most rainfall was in the
last week of June, but rainfall was evenly distributed throughout May and June of 1996-
1998. Poaching, a threat to adult pheasants, increased during this time from 0.6
incidents per day to 0.23. As the pheasant lives at elevations naturally disturbed by
landslides, tolerance for disturbance would be adaptive. Because of the poaching, the
limited nature of the pheasant's distribution, and until the population decline is identified
as indicating a general trend or part of a cyclic pattern in the pheasant's population
dynamics, I recommend S. mikado be considered vulnerable to the risks of extinction.
Keywords: Mikado pheasant, temperature, precipitation, productivity, landslide

052

Brittas, R. 1988. Nutrition and reproduction of the willow grouse Lagopus lagopus
in central Sweden. Ornis Scandinavica 19(1):49-57. Notes: Page 55 -".. .arctic
monocotyledons like cotton grass begin growth and reach peak concentrations of
nutrients sooner after the snow-melt than do dicotyledons. Since the birds start laying
eggs closer to the snow-melt in late springs, this may indicate that a supply of cotton
grass is of greater importance to grouse in such springs, than in early ones. When the
snow disappears early, dicotyledons like bilberry probably have increased their food
quality to a larger extent before the egg-laying period." Key words: willow grouse,
snow, reproduction, nutrition, vegetation

053

Brown, D. E. 1979. Factors influencing reproductive success and population
densities in Montezuma quail. Journal of Wildlife Management 43(2):522-526.

Notes: This study analyzes harvest statistics and correlates them with climatological
information to determine the effects, if any, of weather on Montezuma quail populations
in Arizona. Correlations analysis showed a positive correlation between reproductive
success and summer precipitation. The highest positive correlation coefficient tested
was with the percentage of young quail in the harvest and the preceding summer's June
through August precipitation (a=0.55, P<0.10). Except in 1977, the maximum
reproductive success measured was no more than 78% young in the bag and it appears
that summer precipitation in excess of about 23 cm (9 inches) was superfluous and
possibly even served to depress the percentage of young birds. There was a negative
correlation coefficient between winter (October-March) precipitation and the following
season's reproductive success (a^-0.73, P<0.02). There was a positive linear
relationship between hunting success and the summer precipitation of the year previous
to the summer preceding the hunt plus the population density (as measured by hunting
success)(r=0.53; P<0.10). These data suggest that survival is normally more important

24 j^

H <C> ^

0-"x

c£^>

in determining Montezuma quail population levels than reproductive success.... the
estimated annual survival rates also had a positive correlation with the previous year's
summer precipitation (a=0.60; P<0.05). Lack of either food or grass production brought
on by inadequate precipitation or the removal of grasses by livestock would then reduce
overwinter survival and result in lower population densities. It is easy to see how
continued heavy grazing, particularly during summer droughts, results in the extirpation
of Montezuma quail. Deep snows and long periods of subfreezing temperatures are
also known or believed to have caused high mortality, both locally and generally.
Montezuma quail, precipitation, snow, temperature, reproduction, population
dynamics, vegetation, grazing

054

. 1978. Grazing, grassland cover and gamebirds. Transactions of the North

American Wildlife and Natural Resources Conference 43:477-485. Conclusions —
"California quail, Gambel's quail and mountain quail evolved within comparatively
moderate climates having winter-spring rainfall and a dry growing season. These so
called "Mediterranean" species are characterized by "boom or bust" fluctuations in
reproductive success which is an evolutionary adaptation to annual variations in
precipitation related conditions. These scrub-adapted birds are thus able to rapidly
exploit favorable situations and adjust to less favorable ones. Just the opposite is true
for the grassland gamebirds that evolved with "continental" or tropical climatic regimes.
Prairie chickens, bobwhite, Montezuma quail, sharp-tailed grouse and scaled quail all
display less pronounced variations in reproductive success than Mediterranean species.
Reproductive success in at least 3 of these species in the Southwest is related to
summer precipitation which, within the range of these species, is less variable than
winter rainfall. Continental species are adapted to changes in overwinter survival and
population fluctuations are, at least partially, responses to changes in perennial grass
and/or food production. The reduction of either of these entities decreases the survival
rate (and population level) by increasing the incidence of predation and starvation. The
removal of herbaceous vegetation by livestock deprives these grassland birds (which
are all more or less cryptically colored) of their principal cover — residual
grasses... .Even a conservative utilization of forage in the neighborhood of 20 to 40
percent could be highly detrimental to grassland birds during drought periods because it
could remove that percentage of the bird's cover habitat for the next year." Key words:
prairie chickens, bobwhite quail, Montezuma quail, sharp-tailed grouse, scaled
quail, California quail, Gambel's quail, precipitation, nutrition, predation, drought,
vegetation

055

. and R. H. Smith. 1980. Winter-spring precipitation and population levels of

blue grouse in Arizona. Wildlife Society Bulletin 8(2): 136-1 41. Abstract: Multiple
regression analysis showed an interdependent positive relationship between winter-
spring precipitation, density, reproductive success and subsequent survival rates of blue
grouse (Dendragapus obscurus) in Arizona. With the exception of one year, the
percentage of juvenile grouse observed in August was consistently high and close to a
mean of 65%. Variations in survival are probably more important in determining grouse

f* fc* **

4*

.>">«

£2^ cC±

population levels in Arizona than changes in reproductive success. Precipitation,
hunting success, and survival data suggest that most mortality occurs between spring
and autumn. Annual variations in mortality are thought to be influenced by variations in
the spring growth of herbaceous vegetation and density of grouse. Key words: blue
grouse, precipitation, fecundity, mortality, vegetation

056

Bruce, J. R. 2008. Greater sage-grouse movements and habitat use during winter
in central Oregon. M.S. Thesis, Oregon State University, Corvallis, USA. Notes:
"Winter habitat used by sage-grouse has, in other states, been reported to be
influenced by the amount of big sagebrush exposed above the snow, and these past
studies had snow when they measured sagebrush as being either 0.25-0.46 m above
the snow, or having a total sagebrush height of 0.41-0.56 m. These contrast strongly
with our reported average canopy height range of 0.25 to 0.75 m. We suggest that his
larger range may be that our study lacked enough snow to cover sagebrush, and the
period we measured sagebrush height in plots used by sage-grouse extended into large
winter/early spring when females tended to begin using taller sagebrush... Wyoming big
sagebrush was still used during our study even with little snow as well as presence of
low sagebrush within the study site. ..We determined that this sage-grouse population is
a resident population, as opposed to a migrant one. . .however, it is possible that this
population of sage-grouse could be described as migrant if more severe winter weather
caused grouse to move longer distances in order to find appropriate food and
cover... Based on a year of limited snow cover, we recommend management strategies
for winter habitats that restore or enhance a mosaic of low, mountain, and Wyoming big
sagebrush with sagebrush heights ranging from 0.25 m to 0.75 m tall in areas that have
a large percentage of flat ground and/or areas having no greater than 10% slope and
provide northeast aspects... the time period of our study had a greater range of
temperatures, and was much drier than average overall. Although this study found that
most sage-grouse were located in low sagebrush, various types of sagebrush habitat
are necessary to provide differing sagebrush heights and cover during winter as snow
depth may be a limiting factor to sage-grouse survival during winters with more
precipitation... As sage-grouse seek different levels of cover or different compositions of
sagebrush communities for food and shelter, they will require a landscape large enough
to provide such habitats. Effective sage-grouse management efforts to create or
maintain habitat diversity on extensive tracts large enough to encompass a mosaic of
habitat types for different seasons is essential for the long-term conservation of sage-
grouse populations. Key words: sage-grouse, snow, temperature, vegetation,
habitat use, movement

057

Brunjes, J. H. IV. 2005. The population biology and landscape ecology of Rio
Grande wild turkeys in the Rolling Plains of Texas and Kansas. Dissertation,
Texas Tech University, Lubbock, USA. Notes: Page viii — "Researchers investigating
survival of wild turkeys traditionally have assumed mortalities within the first 14 days
may be capture-related, and have excluded those data from analyses. Few have
explored ways to reduce mortality during this period. In 2000, we initiated a long-term

26 j^

si*

* > t* ><=*

radio telemetry study of the ecology of Rio Grande wild turkeys (M. g. intermedia) in the
southern Great Plains. During 2000-2002, we captured and outfitted 667 turkeys with
backpack-style radio transmitters. We recaptured 123 previously transmittered birds for
a total of 780 14-day survival periods. Sixty-seven birds (8.5%) died < 14 days post
capture and were considered capture-related mortalities. Male mortality (13.4%) was
greater than female (5.8%) mortality (P = 0.001). Birds captured in the afternoon had
higher (P = 0.035) mortality rates (11.6%) versus morning (8.0%) or mid-day (7.1%)
captures. We found no differences in mortality among study sites (P = 0.14), years (P =
0.27), age class for males (P = 0.38) or females (P + 0.99), or capture method (P =
0.64). We found no relationship between weather conditions and 14-day postcapture
survival of turkeys with the exception of precipitation 48 hours post capture (P = 0.01).
We recommend minimizing handling of males and avoiding afternoon captures to
reduce capture-related mortalities." Key words: Rio Grande turkey, precipitation,
capture mortality

058

Burdukov, G. N. and V. M. Kozlov. 1979. Experiment in measuring specific loading
of animals moving through snow. Ekologiya 1:107-109. Abstract: A method is
presented for determining specific loading of animals moving over snow. Specific
loading was studied in the pine marten, otter, wolverine, white hare, squirrel, lynx,
ermine, mink and several species of grouse. A spring-controlled durometer with
alternating punches was used to measure snow resistance (hardness). Paw penetration
was measured and reproduced by the durometer. This method was more precise and
readily performable than previous static methods. Specific loading data help determine
an animal's potential for locomotion. Keywords: grouse, snow, movement,
technique

059

Burkepile, N. A., K. P. Reese, and J. W. Connelly. 2004. Modeling greater sage-
grouse chick survival in southeast Idaho. Meeting of the Western Agencies Sage
and Columbian Sharp-tailed Grouse Technical Committee 24:8. Abstract: Sage-
grouse populations have been declining throughout their range. As a result of this
decline we initiated a 4-year study to determine what reproductive parameters were
limiting greater sage-grouse productivity. During 1999-2002, we radio-marked greater
sage-grouse hens and monitored nesting activity. After eggs hatched, we radio-marked
one-day-old chicks and monitored survival to 10 weeks post-hatch. Nest success
ranged between 41-51% and did not differ (PO.001) between years. From 1999-2001 ,
chick survival ranged between 20-25% and did not differ (PO.001) between years.
However, in 2002 chick survival was higher (35%, P>0.10) than the previous 3 years. In
all years, the highest mortality occurred during the first 3 weeks post-hatch. Proportional
hazard models indicated that increased May-June precipitation had a positive influence
on chick survival. Along with weather, vegetative cover (i.e. grass height, grass and forb
cover) also had a positive influence on chick survival. Our results indicate that greater
sage-grouse populations are negatively influenced by drought conditions through
reduced chick survival, likely mediated through reduced vegetative cover. Key words:
sage-grouse, precipitation, drought, chick survival, mortality, vegetation

) V 4e> t*

■% **"

060

Burnett, L. E. 1905. The sage-grouse, Centrocercus urophasianus. Condor 7:102-
105. Notes: Page 15 — "After feeding they hide either on the feeding ground or at some
distance from it where the sage is large enough to screen them from enemies and the
rays of the sun. ..Hail storms often kill large numbers when they strike the places of
hiding. When their feathers are drenched with rain, the birds are often unable to rise,
and at such times have been killed with a stick." Key words: sage-grouse,
temperature, precipitation, hail, mortality, habitat use, solar radiation

061

Buss, I. O. and C. V. Swanson. 1950. Some effects of weather on pheasant
reproduction in southeastern Washington. Transactions of the North American
Wildlife Conference 15:364-378. Summary: "During 1948 the months of March
through July were abnormally wet and cool, whereas the same months in 1949 were
relatively dry and warm. The wet spring of 1948 terminated in flood conditions during
late May. Plotting 71 hatching dates for 1948 showed that most nests hatched from
June 15 to 28, whereas 117 hatching dates for 1949 showed that most nests hatched
between May 28 and June 4, over two weeks earlier than the hatching peak for 1948."
Keywords: ring-necked pheasant, temperature, precipitation, productivity

062

Cain, J. R., S. L. Beasom, L. O. Rowland, and L. D. Rowe. 1982. The effects of
varying dietary phosphorus on breeding bobwhites. Journal of Wildlife
Management 46(4):1 061 -1065. Summary: This study was undertaken to determine if
the availability of dietary phosphorus could be a factor in the variability of bobwhite quail
reproductive success. Phosphorus content of the soil, and thus the forage, is commonly
deficient in southwestern quail ranges. Further, phosphorus deficiency can be
aggravated during periods of inadequate moisture. Although phosphorus deficiency can
be magnified during drought periods, these findings suggest that a phosphorus-deficient
diet for at least 90 days prior to and including the normal egg-laying period may be a
contributory factor, but is not singularly responsible for causing periodic reproductive
failures in bobwhite quail. Key words: bobwhite quail, precipitation, drought, diet,
fecundity, soil

063

Call, M. W. 1974. Habitat requirements and management recommendations for
sage-grouse. Technical Note 330, U.S. Department of the Interior, Bureau of Land
Management, Denver. Notes: Page 5 — "The oft mentioned decimating factors of
unfavorable weather, increased predation, hunting and disease may have been of
significance in localized areas, but were relatively unimportant in the overall decline in
sage-grouse number." Page 10 — "The primary factor involved with sage-grouse is that
precipitation be adequate to provide for succulent forbs and grasses amongst or near
their sagebrush habitat." Page 12 — " ...In winter, snow takes care of their moisture
requirements, either directly or as it melts and provides free water on warm days.

28 jx

4

^ta^

>>"x

c£^

Precipitation distribution, amounts, and seasonal occurrence, all affect sage-grouse
distribution to some extent. Where late spring and early summer precipitation is
abundant and widespread, the development of succulent vegetation usually induces a
wider distribution of grouse, whereas in drought years they congregate in areas where
free water is available." "Deep snow may cover the spring and summer ranges forcing
the birds to migrate to some distant area for winter and to return for nesting as snow
depths decrease. Where grouse nest and raise their broods on sage-covered slopes on
in mountain valleys at high elevations, they usually must migrate to the desert floors or
other low elevations to find exposed sagebrush for food during the winter. In many
areas, summer and winter areas may be as much as 20 or more miles apart." Page
17 — "In late August at high elevations, or in September or October in some lower areas,
the meadows dry and the incidence of frost increases, leading to drying or killing of the
foliage of forbs. The incidence of sagebrush consumption increases at that time.
Increased moisture content may make sagebrush more palatable than during summer."
Page 17 — "Sage-grouse live almost exclusively on the leaves of sagebrush during the
winter. In Montana when snow depth exceeded 12 inches, sage-grouse were restricted
to taller sagebrush stands, a relatively small percentage of the total range available to
them in a normal (lesser snow) winter... As weather moderated in February, activities
shifted to more open stands of sagebrush." Page 20 — "Wintering birds respond to snow.
Snow depth forces the birds to lower elevations and appears to be a factor determining
the actual wintering site for a flock. "Deep snow limits the availability of food.... In some
areas, sage-grouse occupy windswept sagebrush ridges in winter." Page 26 — "In
wintering habitats, there is little place for fire. Retention of sagebrush is essential on
winter ranges. Even tall, decadent sagebrush, not useful for nesting or brooding, may
be important during severe winters when most other sagebrush could be covered by
snow." Key words: sage-grouse, precipitation, snow, frost, fire, wind, vegetation,
habitat use, movement

064

and C. Maser. 1985. Wildlife habitats in managed rangelands — the Great

Basin of southeastern Oregon, Sage-grouse. General Technical Report PNW-187,
Pacific Northwest Forest and Range Experiment Station, Portland, USA. Notes:
Abiotic factors-Climate. The highest densities of sage-grouse occur where precipitation
averages 25-38 cm (10-16 in) per year. Marginal populations occur in areas of lesser
precipitation. Page 10 — Grouse consumption of sagebrush intensifies as meadows and
foliage of forbs dry and the incidence of frost increases, in late August at high elevations
or in September and October in lower areas. Wintering Habitat: As snow begins to
accumulate on their summer-fall ranges, sage-grouse start moving to lowlands or other
sites, such as windblown ridges, where their needs for forage and cover can be met
throughout the winter. The extent of seasonal movements varies with the severity of
winter weather, topography, and vegetative cover. Sedentary populations meet all their
seasonal requirements in the same general area and, as winter approaches, simply
change their use areas from meadows to sagebrush, because they live almost entirely
on the leaves of sagebrush in winter. Page 1 1 — In seeking wintering areas, grouse
initially select areas with the most palatable sagebrush; if those areas become covered
with snow, they shift to available sagebrush. Page 12 — Habitat surveys conducted
during the summer may give the impression of vast acreages of sagebrush available as

% f>'^^

29

*IA.

<£^ cCi

winter range for grouse. Observations during winter, however, reveal that much of the
habitat is not available because of snow depth or it is unsuitable for other seasons. For
example, when snow depth in Montana exceeds 12 in., sage-grouse were restricted to
taller sagebrush stands. Only 7 percent of the range was available when snow depth
exceeded 1 1 .8 inches. Sage-grouse in Idaho moved to taller sagebrush types when
snow depth reached about 13 in. Key words: sage-grouse, snow, precipitation,
frost, wind, vegetation, diet, movement, habitat use

065

Campbell, H. 1968. Seasonal precipitation and scaled quail in eastern New
Mexico. Journal of Wildlife Management 32(3) :641 -644. Abstract: Spring-summer
rainfall was positively and significantly correlated with subsequent hunter success on
scaled quail {Callipepla squomata) in eastern New Mexico during 1957-66. It is
reasonable to assume that hunter success is related to quail population density in a
direct, but not necessarily proportional, way. On this assumption it appears that the
quail population of eastern New Mexico is importantly influenced and perhaps controlled
by spring-summer rainfall. No statistically significant correlation was found between fall-
winter precipitation and hunter success. Hence, it appears that, unlike the situation in
Arizona with respect to Gambel's quail (Lophortyx gambelii), fall-winter precipitation had
no important influence on the scaled quail population studied. Key words: scaled
quail, Gambel's quail, precipitation, population dynamics

066

, D. K. Martin, P. E. Ferkovich, and B. K. Harris 1973. Effects of hunting and

some other environmental factors on scaled quail in New Mexico. Wildlife
Monograph 34, The Wildlife Society, Washington, D. C, USA. Summary: Quail
breeding success on the study areas was strongly related in a positive way with spring-
summer precipitation, and the July-August component of the precipitation was
especially important. There was no statistical relationship whatever between scaled
quail breeding success and the fall-winter precipitation which Swank and Gallizioli found
essential for successful production of Gambel's quail in Arizona. It is unlikely that
seasonal precipitation per se exerts an important influence on quail populations, but
rather that precipitation controls some other factor which does exert an important or
controlling influence. This factor probably is nutritional, most likely the food, vitamins,
minerals, etc., produced by a diversity of forbs, all of which depend for their
development and fruiting on adequate rainfall during definite seasons of the year. The
influence of seasonal rainfall in producing crops of insects useful to quail as food may
also be of great importance. Climatographs suggest the possibility that there may have
been a negative causal relationship between comparatively high summer temperatures
and high quail production, but that there was no relationship between winter
temperatures and quail production. Key words: scaled quail, precipitation,
temperature, productivity, nutrition

067

Campbell-Kissock, L, L. H. Blankenship, and L D. White. 1984. Grazing
management impacts of quail during drought in the northern Rio Grande Plain,

30

4-

>'>" fe ^

Texas. Journal of Range Management 37(5): 442-446. Abstract: Relationships
between the abundance of 2 quail species and range site and grazing management
during drought were evaluated in the northern Rio Grande Plain of Texas. Clay loam
range sites provided better nesting cover and greater abundance of forbs for quail than
sandy loam and shallow ridge range sites. Foliar cover and aboveground standing crop
of grass were greater on the 3 range sites within the short duration and deferred
rotation systems as compared with the yearlong system. During drought, grazing
systems provided better nesting and protective cover for quail than yearlong grazing.
Key words: quail, drought, vegetation, grazing, soil

068

Cardona, C. J., A. Ihejirika, and L. McClellan. 2002. Haemoproteus lophortyx
infection in bobwhite quail. Avian Diseases 46(1):249-255. Abstract: This report
chronicles recurring outbreaks of Haemoproteus lophortyx infection in captive bobwhite
quail. Clinically, the signs of infection included reluctance to move, ruffled appearance,
prostration, and death. These signs were associated with parasitemia, anemia, and the
presence of large megaloschizonts in skeletal muscles, particularly those of the thighs
and back. The average cumulative mortality for flocks experiencing outbreaks was over
20%. In a typical outbreak, mortality rose when the birds were 5-6 weeks of age,
peaked in 8- to 10-week-old quail, and declined rapidly when the quail were 9-11 weeks
old. Outbreaks occurred exclusively between the months of May and October, and
warm weather was determined to be a risk factor for H. lophortyx mortality. This
protozoan most likely overwinters in native California quail in the area and is transmitted
to quail on the ranch by an insect vector that emerges in warm weather. Infection of the
large population of naive bobwhite quail on the ranch leads to amplification of H.
lophortyx, resulting in epidemics in successive flocks. Key words: bobwhite quail,
temperature, disease, mortality, parasites

069

Carpenter, J. E. 2007. West Nile virus and parasites in greater sage-grouse
{Centrocercus urophasianus) populations. M.S. Thesis, University of Alberta,
Edmonton, Canada. Notes: "Because development of C. tarsalis appears closely tied
to temperature and moisture regimes, local differences in temperature and moisture will
influence the annual risk presented by WNv. Early virus activity, detected from wild bird
surveillance and mosquito monitoring, coupled with local moisture and temperature
conditions are critical indicators of WNv risk." Key words: sage-grouse, temperature,
precipitation, disease, insects

070

Case, R. M. and R. J. Robel. 1974. Bioenergetics of the bobwhite. Journal of
Wildlife Management 38(4):638-652. Abstract: Ten bobwhites (Collnus virginianus) of
each sex were individually confined under controlled, simulated seasonal conditions
(10- and 15-hour photoperiods, temperatures from 0 to 40 C) in order to quantify energy
requirements (gross energy intake, existence energy, and excretory energy). Quails
confined under a 15-hour photoperiod had significantly (P < 0.005) greater energy
requirements than those under a 10-hour photoperiod. There was no difference (P >

f V ^ ^v

X **"

4^> cc^

0.05) between males and females at 10 hours, nor between males and nonlaying
females at 15 hours, for energy requirements. The above energy variables decreased
with increasing temperature, in a linear manner, except for egg-laying females (changed
quadratically). Existence energy of quails increased 108 percent between 0 and 30 C
for 10-hour and 124 percent for 15 hour photoperiods. Body weights changed
curvilinearly with temperature, at both photoperiods, in each sex. Males were heavier
than females at 10 hours; at 15 hours the females were heavier. Body weight accounted
for less than 1 percent of the total variation about energy variables. Males survived
higher temperatures than females at both photoperiods (mean upper lethal temperature
43.7 C vs. 42.6 C at 10 hours and 44.7 C vs. 41.0 C at 15 hours). There was no
predictable pattern for efficiency of feed utilization. The highest efficiency (76 percent)
was at the 10-hour photoperiod. Productive energy for egg-laying females decreased at
both high and low temperatures and was maximum (13.800 kcal/bird-day) at 25 C.
None of our egg measurements correlated significantly (P > 0.05) with body weight of
quails. Key words: bobwhite quail, temperature, bioenergetics

071

Cattadori, I. M., D. T. Haydon, and P. J. Hudson. 2005. Parasites and climate
synchronize red grouse populations. Nature 433:737-741. Abstract: There is
circumstantial evidence that correlated climatic conditions can drive animal populations
into synchronous fluctuations in abundance. However, it is unclear whether climate
directly affects the survival and fecundity of individuals, or indirectly, by influencing food
and natural enemies. Here we propose that climate affects trophic interactions and
could be an important mechanism for synchronizing spatially distributed populations.
We show that in specific years the size of red grouse populations in northern England
either increases or decreases in synchrony. In these years, widespread and correlated
climatic conditions during May and July affect populations regionally and influence the
density-dependent transmission of the gastrointestinal nematode Erichostrongylus
tenuis, a parasite that reduces grouse fecundity. This in turn forces grouse populations
into synchrony. We conclude that specific climatic events may lead to outbreaks of
infectious diseases or pests that may cause dramatic, synchronized changes in the
abundance of their hosts. Key words: red grouse, temperature, humidity,
precipitation, disease, parasites, fecundity, mortality, population dynamics

072

Cedarleaf, J. D., S. D. Worthen, and J. D. Brotherson. 1982. Weather conditions in
early summer and their effects on September blue grouse Dendragapus obscurus
harvest. Great Basin Naturalist 42(1):91-95. Abstract: Relationships of temperature
and precipitation to the reproductive success of blue grouse (D. obscurus) were
investigated. Maximum and minimum temperatures followed similar patterns during the
years 1976-1981 and showed no patterns relative to hatching success. Precipitation
data was variable. When significant amounts of precipitation fell during the last three
weeks of the hatching period, chick survival and thus recruitment were adversely
affected. Precipitation occurring at the end of the hatch period probably reduces the
September harvest of birds. Key words: blue grouse, temperature, precipitation,
reproductive success, chick survival, recruitment

32 ^

H 1& >f*

073

Chamberlain, E., R. D. Drobney, and T. V. Dailey. 2002. Winter cover height and
heat loss: is taller better? Proceedings of the National Quail Symposium 5:157.

Abstract: Previous studies have demonstrated that roost site selection affects energy
requirements for thermoregulation in several avian species; however, the influence of
microhabitat characteristics on heat loss has not been evaluated for northern bobwhites
(Colinus virginianus). One frequently measured microhabitat feature that is commonly
thought to influence the thermal characteristics of avian ground roost sites is cover
height. We simultaneously measured thermoregulatory energy expenditure of
bobwhites across a range of low ambient temperatures (-24° to 14° C) in 3 cover
heights (0 cm, 46 cm, 124 cm) using 3 heated taxidermic mounts. Predicted metabolic
rates (PMR) were derived on the basis of power consumption of the taxidermic mounts.
Predicted metabolic rate for each vegetation height was linearly related to ambient
temperature, and decreased significantly (P < 0.769) among the 3 vegetation heights
across a range of environmental conditions. These findings suggest that under the
conditions occurring during our field measurements, thermoregulatory energy
requirements of bobwhites was essentially independent of vegetation height at the
roost, and primarily are a function of conductive rather than convective heat loss. Key
words: bobwhite quail, temperature, thermoregulation, vegetation

074

, , and . 2000. Vegetation and thermal characteristics of

bobwhite nocturnal roost sites in native warm-season grass. Proceedings of the
National Quail Symposium 4:58. Abstract: Native warm-season grass (NWSG) has
been widely promoted as wildlife habitat, but little empirical evidence is available to
support its value for most wildlife species. One justification for a conversion to NWSG is
the high thermal quality of cover resulting from the height and structure of the
vegetation. Because vegetation cover is an important factor contributing to bobwhite
winter survival, we predicted that they should select roost sites with superior thermal
characteristics during winter when energy requirements for thermoregulation are
greatest. In this 3-year study we used data derived from roost sites (/7=166) obtained
from radio-marked quail to compare the relative use of NWSG and 5 other habitat
types, and the micro-habitat characteristics of winter roost and random sites on an area
intensively managed for quail in Missouri. Of the 6 habitats used for roosting, most
locations (51 .2%) were in old-field habitats. NWSG ranked third with 17% of the
locations. Our findings indicated that roost site selection may be influenced to a greater
extent by the micro-habitat characteristics of a site rather than by habitat type. Two
micro-habitat features that were of particular importance in habitats used most by quail
were litter cover and canopy cover. These habitat features are valuable in reducing
conductive and convective heat loss. Key words: bobwhite quail, thermoregulation,
vegetation, habitat use, micro-habitat

075

Charette, J. 2000. On the diet of the American woodcock (Scolopax minor) during
drought periods. In: D. A. Stroud and N. C. Davidson, editors. Wader Studies

% M 4e> t*

33

£Z^> c£^>

Methods: Practical Papers Published in the Wader Study Group Bulletin Part 2:
Feeding, census and survey techniques. Unable to obtain copy for abstract.

076

Choate, T. S. 1963. Habitat and population dynamics of white-tailed ptarmigan in
Montana. Journal of Wildlife Management 27(4):684-699. Abstract: The ecology and
structure of a population of white-tailed ptarmigan (Lagopus leucurus leucurus), residing
during the summer near Logan Pass, Glacier National Park, were studied from early
June to mid-September, 1959-62. Although ptarmigan were observed in a wide variety
of habitats within the alpine zone, they showed a preference for areas having moderate
rock cover, plentiful snow or water, and short, young vegetation. During the breeding
season, 21 males and 14 females used the study area (7.08 acres of ptarmigan
habitat). Territorial males returned to the same sites year after year and occasionally
had the same mates. Yearling males rarely established and mated in their first year, but
yearling females bred successfully. Females occasionally returned to the same
breeding areas. Few females born on the study area returned to it, but many males did.
Clutch size averaged 5.2 eggs (range 3-9). Nest parasitism was rare. Renesting
occurred occasionally in phenologically early years. The percent of successful females
varied from 35 to 82. Poor success was often caused by inclement weather. Hatching
success was 85.5 percent. Brood size at flight age ranged from 3.25 to 3.47. Despite a
large variation in natality, the size of the adult population at Logan Pass varied little.
Average adult annual mortality was 29 percent, lower than that reported for other
galliform species. Chick mortality was 35-44 percent by the time of dispersal. Mortality
of immature birds was 63 percent by the following spring. A life table based on these
average mortalities indicates a maximum life-length of 15 years, a mean longevity of
3.02 years, and an annual mortality rate of 42.1 percent. The Logan Pass population did
not show the fluctuation in numbers reported for other species of ptarmigan and
suggested for the tundra biome. Key words: white-tailed ptarmigan, snow, mortality

077

Christensen, G. C. 1970. The chukar partridge: its introduction, life history, and
management. Biological Bulletin No. 4, Nevada Department of Wildlife, Reno,
USA. Notes: In a section titled "Inimical Factors", the author discussed the effects of
climate Pages 43-46 — "Deep, persistent snow can completely cover food in higher
areas that causes the chukars to migrate to lower elevations where food is available.
Chukars are able to scratch beneath brush when snow is only an inch or two deep. In
1951 chukars in the Peavine Mountain area northwest of Reno were forced onto
highways near Reno and into the town, obtaining food uncovered by plows along
roadways. One specimen showed near depletion of fat reserves. Other birds showed no
distress. Chukar losses due to snowfall were recorded in some areas. There can be
severe losses to local chukar populations as a result of heavy snows. Prolonged cold
temperatures following heavy snows compound the problem. In terms of precipitation,
recently hatched chicks can be killed as a result of exposure to rains." "Factors affecting
reproduction" Pages 38-39 — "Precipitation is discussed as a factor in determining
population dynamics directly and indirectly through nutrition. In arid or semi-arid regions
it is important that the precipitation occur at the proper time of year to facilitate vegetal

34 j,

4>

IX ^ ^

growth and production culminating in a seed crop. Localized thundershowers can have
a considerable influence in some areas resulting in pockets of good bird production and
other areas exhibiting poor production. Temperature and wind as well as soil condition
(frozen or not) play important roles in determining the effectiveness of precipitation. In
Nevada population trends show chukars exhibit boom and bust patterns correlated with
drought conditions and lack of nutritious food. During the severe drought years of 1953
and 1954 the adult:young ratios in Washoe County were 100/63 and 100/11
respectively, showing the production had almost reached a standstill. Food conditions
were so poor that adult birds were found to be subsisting on dry rootstocks and stems
of grass. When a population bottoms out it seems to require a minimum of three good
production years, back to back, to bring the population to a peak again. In addition to
affecting chukar production indirectly through the food chain, climatic conditions can
also directly affect the success of the hatch. Abnormally heavy precipitation in May and
early June can cause chick morality and in some instances adverse climate may result
in nest loss. Keywords: chukar partridge, precipitation, drought, temperature, soil,
productivity, mortality, vegetation, nutrition, population dynamics

078

.1958. The effects of drought and hunting on the chukar partridge.

Transactions of the North American Wildlife Conference 23:331-341. Summary:
The precipitation pattern and effectiveness has a direct bearing upon the annual food
crop which is produced in chukar habitat. The condition of the range follows in turn as
the determining factor in chukar production. During a three-year drought in Nevada
there was a decided decrease in chukar production and decline in population. With the
advent of proper range conditions the populations made a quick recovery. Key words:
chukar partridge, drought, nutrition, productivity

079

Christiansen, T. 2005. Greater sage-grouse job completion report, June 1, 2005-
May 31, 2006. Wyoming Game and Fish Department, Cheyenne. Notes: Page 7 —
"Results of this and prior monitoring suggest sage-grouse populations in the Wyoming
(sic) were at their lowest level ever recorded in the mid-1990s. Grouse numbers then
responded to increased precipitation during the late 1990's with some individual leks
seeing three fold increases in the number of males counted between 1997 and 1999.
The return of drought conditions in the early 2000's led to decreases in chick production
and survival and therefore population declines, although the population did not decline
in mid-1 990's levels. Improved habitat conditions due to timely precipitation in 2004 led
to high chick production and survival. This resulted in 2006's counts and surveys having
the highest recorded average males per lek since 1978. Key words: sage-grouse,
precipitation, temperature, vegetation, population dynamics

080

Church, K. E. and W. F. Porter. 1990. Population responses by gray partridge to
severe winter conditions. Pages 295-303 in K. E. Church, R. E. Warner, and S. J.
Brady, editors. Proceedings Perdix Symposium V: gray partridge and ring-necked
pheasant workshop. Minnesota Department of Natural Resources, Mankato, USA.

>V fe« **

^

Abstract: We compared 3 methods of estimating changes in the abundance of gray
partridge (Perdix perdix), and described population responses during a severe winter in
New York. Change-in-ratio analysis and biweekly census provided similar estimates of
winter mortality, whereas changes in mean covey sizes underestimated loss due to
migration and extinction. We estimated 58% population loss over a 2-week period in
early February. During this period subadult partridge were 3 times less likely to survive
than adults. Evidence from radio-tagged partridge indicated loss was primarily due to
predation by great horned owl (Bubo virginianus) or red-tailed hawk (Buteo
jamiacensus), not starvation. We concluded overwinter survival may limit gray partridge
populations, and recommend supplemental feeding to minimize the risk of predation.
Key words: gray partridge, ring-necked pheasant, severe winter, mortality,
predation

081

Churchwell, R., J. T. Ratti, and F. Edelmann. 2004. Comparison of fall and winter
food habits for sympatric chukar and gray partridge in Hells Canyon of Idaho and
Oregon. Northwest Science 78(1):42^47. Abstract: Chukar and gray partridge are
abundant and sympatric in Hells Canyon, Idaho and Oregon. We collected crops (birds
collected during the hunting season) from both species to compare fall and winter food
habits, which allowed for comparison of food selection, estimation of niche overlap, and
identification of important food resources. We collected 143 chukar and 112 gray
partridge crops during the 1999 and 2000 hunting seasons (late September through
early January). We identified 21 items consumed by chukar and 16 items consumed by
gray partridge. Both bird species consumed similar foods, with an 80% dietary overlap.
Dominant food items for both species included prairiestar root nodules and vegetative
parts from various grasses and forbs. Food items differed between years, which may
have reflected variation in weather and plant cycles. This is the first study of gray
partridge from a non-agricultural environment, and the first rigorous comparison of food
habits for sympatric chukar and gray partridge population in North America. Key words:
chukar partridge, gray partridge, weather, vegetation, diet

082

Clarke, J. A. and R. E. Johnson. 1992. The influence of spring snow depth on
white-tailed ptarmigan breeding success in the Sierra Nevada. Condor 94(3):622-
627. Abstract: The relationship between spring snow depth and breeding success of an
introduced population of white-tailed ptarmigan (Lagopus leucurus altipetens) in the
Sierra Nevada, California, was studied from 1982 through 1987. Yearly spring snow
depth varied from 50.8 cm to 424.2 cm. Hatch dates were later in years of deep snow
(P<0.05) but brood size showed no relation to snow depth. Nesting success, chick
survival, and brood success (all of which contribute to breeding success) were
negatively correlated with snow depth as was breeding success (P<0.05). The number
of paired ptarmigan in the study area varied from 22 (in 1987) to 46 in 1985 which was
the year with the least snow; however, no significant relationship existed between
breeding numbers and snow depth. Successful reproduction of white-tailed ptarmigan in
the Sierra Nevada appears to be strongly affected by snow, potentially due to its

36 \|

<ir

H' fc* ^

c^>

influence on the availability of resources such as nest sites, food, and cover. Key
words: white-tailed ptarmigan, snow, reproduction

083

Coates. P. S. and D. J. Delehanty. 2008. Effects of environmental factors on
incubation patterns of greater sage-grouse. Condor 110(4):627-638. Abstract: Birds
in which only one sex incubates the eggs are often faced with a direct conflict between
foraging to meet metabolic needs and incubation. Knowledge of environmental and
ecological factors that shape life-history strategies of incubation is limited. We used
continuous videography to make precise measurements of female greater sage-grouse
(Centrocercus urophasianus) incubation constancy (percentage of time spent at the
nest in a 24-hour period) and recess duration. We used an information-theoretic
approach to evaluate incubation patterns in relation to grouse age, timing of incubation,
raven abundance, microhabitat, weather, and food availability. Overall, sage-grouse
females showed an incubation constancy of 96% and a distinctive bimodal distribution
of brief incubation recesses that peaked at sunset and 30 min prior to sunrise. Grouse
typically returned to their nests during low light conditions. Incubation constancy of
yearlings was lower than that of adults, particularly in the later stages of incubation.
Yearlings spent more time away from nests later in the morning and earlier in the
evening compared to adults. Video images revealed that nearly all predation events by
common ravens (Corvus corax), the most frequently recorded predator at sage-grouse
nests, took place during mornings and evenings after sunrise and before sunset,
respectively. These were the times of the day when sage-grouse typically returned from
incubation recesses. Recess duration was negatively related to raven abundance. We
found evidence that incubation constancy increased with greater visual obstruction,
usually from vegetation, of nests. An understanding of how incubation patterns related
to environmental factors will help managers make decisions aimed at increasing
productivity through successful incubation. Key words: sage-grouse, solar radiation,
microhabitat, weather, predation, incubation

084

Cobb, D. T. and P. D. Doerr. 1997. Eastern wild turkey reproduction in an area
subjected to flooding. Journal of Wildlife Management 61(2):313-317. Abstract: We
used cohort analyses and population cohort matrices to model a wild turkey {Meleagris
gallopavo silvestris) population under perturbed (i.e., man-induced flooding on 3-year
intervals) and unperturbed (i.e., non-flood) conditions. The net reproductive rate (R0) of
a cohort in which reproduction in the hatching-year (HY) age class was perturbed by
flooding dropped to 0.460 from R0 = 1 .383 for unperturbed cohorts. The R0 of cohorts in
which only after-hatching-year (AHY) age classes were exposed to flooding was > 1 .0.
Cohort analyses demonstrated the importance to the population of nesting by HY hens
and the significant effects on cohort reproductive potential of exposing the HY age class
to flooding. Evaluation of population cohort matrices also suggested that flooding on a
3-year interval precludes sufficient reproduction to maintain this wild turkey population.
Key words: turkey, reproduction, flood

— ** A — T^' / _ t I ^

\ H^^

eg, ^ £^> d^

085

Coggins, K. A. 1998. Relationship between habitat changes and productivity of
sage-grouse at Hartt Mountain National Antelope Refuge, Oregon. M.S. Thesis,
Oregon State University, Corvallis, USA. Notes: "After early studies (1989-1991),
climatic conditions and land-use practices changed at HMNAR. Livestock grazing was
eliminated from HMNAR in 1991 and during 1989-1997 there was an upward trend in
crop year precipitation (total precipitation from 1 September-30 June). Because of these
changes a study was initiated to test the relationships inferred from earlier studies. Two
sample periods (1989-1991 and 1995-1997) were compared to determine the collective
effect of increased crop year precipitation and reduced livestock grazing on key habitat
components in 4 cover types and on sage-grouse productivity measures... .Herbaceous
vegetation increased in all cover types during 1995-1997 compared with 1989-1991 at
HMNAR. Increased herbaceous vegetation during the second time period was probably
related to above average crop year precipitation in conjunction with the absence of
livestock grazing during 1992-1997. Results of my study indicated that mean nest
initiation and renesting rates increased concordantly with spring forbs, which increased
2- to 3-fold in several key cover types between periods. Cover type use by pre-laying
hens did not change between time periods; therefore, increases in nest initiation and
renesting rates apparently were related more to changes in habitat components (spring
forbs) in these cover types rather than changes in use of cover types by grouse.
Greater availability off spring forbs at HMNAR likely provided hens more opportunity to
consume protein rich foods, and ultimately, to become physiologically prepared to
initiate nesting and renesting, if necessary. I also found that mean nesting success
increased concomitantly with increases in tall grass cover available during
spring.... greater residual herbaceous cover in sage-grouse nesting habitat during spring
increased the chances of nest success by providing scent, physical, and visual barriers
to predators.... Results of this study revealed that sage-grouse productivity increased
concomitantly with increased herbaceous vegetation at HMNAR and confirmed the
relationship of certain hebaceous habitat components to reproductive success of sage-
grouse... data from Oregon hunter wing returns indicated sage-grouse productivity was
at record high levels throughout Oregon in 1996 and 1997. During 1989-1991, 35% of
hunter harvested sage-grouse were immatures, whereas during 1995-1997, 47% were
immatures. These results possibly reflected widespread changes in herbaceous
vegetation related to increased precipitation in conjunction with reduced livestock
stocking rates... Herbaceous vegetation and, ultimately, sage-grouse productivity
increased concurrently with increased crop year precipitation and removal of livestock
grazing at HMNAR." Key words: sage-grouse, precipitation, grazing, vegetation,
nutrition, reproduction

086

Connan, R. M. and D. R. Wise. 1994. Further studies on the development and
survival at low temperatures of the free living stages of Trichostrongylus tenuis.
Research in Veterinary Science 57(2):21 5-219. Abstract: To assess the ability of the
free living stages of Trichostrongylus tenuis, a pathogen of red grouse, to survive the
winter in significant numbers, the temperatures prevailing during the autumn, winter and
spring of 1991-1992 on the North Yorkshire moors were simulated in an incubator into

38 ^

) \ ^ ^

Q>

which replicate cultures of T. tenuis eggs were placed at intervals. September and early
October eggs developed into larvae which survived over winter and were infective the
following spring. Few November to February eggs survived to become larvae but early
March eggs were reasonably successful. Continuous temperatures of -15 degree C
were lethal to infective larvae within 12 days, but significant numbers of larvae survived
temperatures of -10 degrees C or higher for up to three weeks and remained infective.
Key words: red grouse, temperature, precipitation, disease, parasites

087

Connelly, J. W., K. P. Reese, R. A. Fischer, and W. L. Wakkinen. 2000. Response
of a sage-grouse breeding population to fire in southeastern Idaho. Wildlife
Society Bulletin 28(1):90-96. Notes: "...we... concluded that breeding population
declines became more severe in years following fire. Prescribed burning negatively
affected sage-grouse in southeastern Idaho and should not be used in low-precipitation
sagebrush habitats occupied by breeding sage-grouse." Key words: sage-grouse,
fire, precipitation, reproduction

088

Conover, M. R. and J. S. Borgo. 2009. Do sharp-tailed grouse select loafing sites
to avoid visual or olfactory predators? Journal of Wildlife Management 73(2):242-
247. Abstract: Grouse should seek loafing sites hidden from predators; however, good
hiding sites from predators that use vision to locate prey differ from good hiding sites
from predators that use odor to locate prey. We compared characteristics of control
sites to sites used for loafing by sharp-tailed grouse (Tympanuchus phasianellus) to
determine whether selection of loafing sites was more influenced by the need to hide
from visual or olfactory predators. Sites used for loafing were similar to control sites in
characteristics that would help hide a grouse from visual predators (i.e., visual
obstruction, lateral visibility, visual obstruction, cover height, and surface roughness),
but loafing sites differed from control sites in characteristics that would help hide a
grouse from olfactory predators (i.e., greater updrafts, wind velocities, and atmospheric
turbulence). Key words: sharp-tailed grouse, wind, predation, topography, habitat
use, behavior

089

Coon, R. A., P. D. Caldwell, and G. L. Storm. 1976. Some characteristics of fall
migration of female woodcock. Journal of Wildlife Management 40(1):91-95.

Abstract: Nine female woodcock (Philohela minor) were radio-tagged in central
Pennsylvania before fall migration to monitor premigratory and migratory movements.
Within 15 days of departure, 5 of the birds moved 0.8 to 8.0 km from their normally used
area, but the remaining 4 did not move. In 1973 five marked woodcock began migration
between 30 November and 8 December. In 1974, four birds departed between 18 and
29 November. Departures coincided with high pressure centers approaching from the
north and west or low pressure centers retreating to the north and east, or both. Eight of
the 9 woodcock departed 2.5 or more hours after sunset and at least 7 left before
midnight. Two hatching-year birds were tracked for up to 201 km SSW of the study area

i >*rte*>f*

39

Cgj: ^ ^2^ <£J>

during 2 nights. Their air speeds (mean ± SD) were 36 ± 2 and 45 ± 3 km/h. Key
words: American woodcock, barometric pressure, movement, behavior

090

Cope, M. G. 1992. Distribution, habitat selection and survival of transplanted
Columbian sharp-tailed grouse in the Tobacco Valley, Montana. M. S. Thesis,
Montana State University, Bozeman, USA. Notes: Page 35 — Use of grasslands in
winter was attributed to the fact that snow cover was minimal, and food may not have
been a limiting factor. Complete snow cover on the Tobacco Plains is rare, so this
assumption might also be applied to the Tobacco Valley." Key words: sharp-tailed
grouse, snow, habitat use

091

Copelin, F. F. 1963. The lesser prairie chicken in Oklahoma. Oklahoma Wildlife
Conservation Department Technical Bulletin No. 6, Oklahoma City, USA. Notes:
Page 37 — Brood Range. Broods seemed to be more mobile in dry years when grass
and shinnery cover was sparse than during wet seasons when cover was dense. During
the summer of 1956, a severe drought year when population density was very low, one
marked brood was observed in 13 places in July and August, and they covered a
minimum enclosed area of 256 acres. The greatest distance between points of
observation was 1 .43 miles. On the other hand, in 1 959 when rainfall was near the long
term mean and vegetation was dense, three marked broods were found in no more than
three places, and on a smaller range of about 160 acres. Habitat Use — In summer,
during hot weather, lesser prairie chickens apparently required adequate shade. During
summer they were found only in pastures, never in cultivated fields. In the shin oak type
of vegetation prairie chickens gathered in the shade of oak motts, but only when it was
very hot. In sand sagebrush and mixed-grass prairie types they moved into the shade of
sagebrush, skunkbrush, sand plum, ragweed, and other bushy plants and forbs. The
dependence of prairie chickens on good shade in summer is indicated by the following
correlations between weather, soil moisture, and habitat use: t. Insolation (solar
radiation). Prairie chickens were found in motts only on clear days. Motts provided
better shade than shinnery. Z Air Temperature. More prairie chickens were found in
motts on hot days than on cool days. A direct correlation existed between the numbers
of birds in motts and daily maximum temperatures, except during late August soon after
a 1 .02 inch rain when ground moisture was high. 3. Wind velocity. There appeared to
be little correlation between wind velocity and the number of prairie chickens in motts,
except winds above 30 miles per hour may have reduced usage slightly. 4^ Soil
moisture (three inches below the ground surface). More prairie chickens were found in
motts when the ground was dry than when it was moist... It is important to recognize
that birds used motts only when both temperature was high and ground moisture was
low. Page 42 — Finding and trapping prairie chickens was less rewarding when either
temperature or soil moisture was not optimum.... I believe herein is the reason lesser
prairie chickens occur only in regions with brushy vegetation. The need of shade in
summer is critical when temperatures near or exceed 100°F. Apparently, grassy
vegetation does not provide sufficient shade, particularly during extreme droughts when
temperatures are highest and grass cover is sparse. Page 47 — The change in winter

40 j,

4-

f > fcL, >f*

feeding habits apparently was influenced by increased production of acorns, grass
seed, and forb seed in the pastures. This year (1957) was the first wet year following a
long drought. Vegetative ground cover had been sparse. With increased moisture, forbs
and legumes prospered and produced an abundance of seed. Page 51 — One important
component of lesser prairie chicken habitat is "brushy" vegetation. Shinnery oak, sand
sagebrush, plum and skunkbrush commonly provide shade in summer. Sagebrush and
skunkbrush provide protection from strong winds in winter. Lesser prairie chicken range
does not extend to grasslands where brush is absent. Sagebrush leaves and shin oak
acorns are eaten by prairie chickens. Key words: lesser prairie chicken,
precipitation, temperature, drought, wind, vegetation, habitat use, nutrition, soil
moisture, census, movement, habitat modification, solar radiation

092

Coup, R. N. and P. J. Pekins. 1999. Field metabolic rate of wild turkeys in winter.
Canadian Journal of Zoology 77(7):1075-1082. Abstract: We investigated the winter
bioenergetics of eastern wild turkeys (Meleagris gallopavo sylvestris) by measuring
standard metabolic rate (SMR) and existence metabolism (EM) of captive turkeys and
field metabolic rate (FMR) of free-ranging turkeys. Mean SMR and EM were
0.511 ±0.040 ml_ 02g"1 h-1 and 499.7±17.7 kJ kg body mass 0734d 1 (mean ±SE) as
measured by indirect respirometry and food consumption, respectively. FMR was
measured with doubly labeled water and was 10.5% higher in juvenile (0.976±0.039 L
C02 kg 0734h"1) than adult turkeys (0.883±0.034 L C02 kg"0734h 1); their FMR:SMR ratios
were 1.74 and 1.58, respectively. Juvenile turkeys weighed less and had less body fat
(13.5%) than adults (18.9%). Mean FMR was lowest in 1996, when ground forage was
unavailable and weather was more windy and cold than in 1995, when ground forage
was available and the turkeys' activity and range were greater. Turkeys reduced FMR in
1996 by restricting movement and range, and using proximate shelter and supplemental
food. We predict that juvenile turkeys are at an energetic disadvantage when food
availability is restricted because of their higher FMR, lower body and fat masses, and
higher activity costs than adults. Key words: eastern wild turkey, bioenergetics,
temperature, wind, vegetation, habitat use

093

Crawford, J. A. 1978. Factors affecting California quail populations on the E. E.
Wilson Wildlife Area, Oregon. Murrelet 59(1):7-13. Summary: Flush censuses and
transect counts revealed that California quail preferred early successional stages and
open habitat. Long-term effects of weather and succession also were examined;
succession was the only variable (r= 0.58) which was related significantly to the
population decline. Rate of harvest did not affect quail populations from 1958 to 1974.
Because of the rapid rate of succession, California quail were closely associated with
early successional stages and open habitat. Key words: California quail, weather,
habitat use, vegetation

094

Criddle, N. 1930. Some natural factors governing the fluctuation of grouse in
Manitoba. Canadian Field-Naturalist 44(4):77-80. Abstract: The author suggests that

% f >'' fe* ^

41

<£2& -"car- ^^> <^>

young birds starve from lack of insects during cold and rainy weather rather than
succumb directly to rain and cold. An annually repeated census of 2 areas in Manitoba
from 1914 to 1929 shows a periodic fluctuation in the numbers of ruffed grouse and
various other birds, peaks of abundance occurring in 1915 and 1924. A graph shows
fluctuations in sharp-tailed and ruffed grouse and grasshoppers in correlation with
rainfall in Aweme, Manitoba, from 1895 to 1929. Grasshopper abundance is generally
preceded or accompanied by abnormally dry seasons. The peaks of grouse abundance,
in turn, are preceded or accompanied by grasshopper outbreaks. Grasshoppers are
apparently necessary food for young sharp-tailed grouse. The ruffed grouse, being
more of a woodland species than the sharp-tailed, seems to be less affected than the
latter by grasshopper prevalence. Key words: sharp-tailed grouse, ruffed grouse,
temperature, precipitation, drought, insects, nutrition, population dynamics

095

Crissey, W. F. 1947. Influence of weather. Pages 299-306 in G. Bump, R. W.
Darrow, F. C. Edminster, and W. F. Crissey, editors. The ruffed grouse; life
history, propagation, management. New York State Conservation Department,
New York, USA. Notes: Page 300 — "Weather is a basic influence on grouse and other
wildlife species through its effect on the environment in which they live; its effect is
largely indirect and specific relationships are therefore difficult to determine; Page
301 — "direct losses from weather conditions seem negligible in New York, but on one
occasion a severe cloudburst and flood before the chicks were six weeks old apparently
took a heavy toll"; Page 303 — "although the onset of the reproductive season is
primarily controlled by the progressively increasing length of daylight in the spring,
minor variations in nesting dates from year to year seem related to the average
minimum temperature during early April; there is apparent correlation between
temperature and precipitation during the three weeks immediately following hatching
and the brood mortality occurring during the latter part of the summer"; Page 304 — "the
degree of over-winter loss among adult grouse seems to be affected to some extent by
the severity of the weather during March"; Page 306 — "there appears to be some
degree of agreement between the occurrence of low temperatures during March and
June and periods of grouse scarcity which have been recorded in the past." Key
words: ruffed grouse, precipitation, flood, temperature, solar radiation, mortality

096

Dahlgren, R. 1974. Pheasant mortality and production in South Dakota related to
antibodies for western and eastern viral encephalitis. American Midland
Naturalist 91(1):237-241. Notes: Page 240— Sizes of insect vector populations would,
of course, be influenced by weather patterns. Success of pheasant reproduction has
also been shown to be dependent on weather patterns. Since the extent of arbovirus
diseases is dependent on weather and vector populations, WEE and EEE may be
influenced by the same weather patterns which are associated with ups and downs in
pheasant populations. These interrelationships are too complex to be analyzed easily.
However, the relationship of disease to population fluctuations in pheasants has neither
been thoroughly investigated nor discussed. While further research is necessary to
determine if WEE and EEE are casually or causally related to success of pheasant

42

% ryi&n*

reproduction, we think the correlations presented in this paper given sufficient reason
for the wildlife biologist to consider diseases as a factor that may be involved in
population regulation. Keywords: ring-necked pheasant, weather patterns, disease,
insects, population dynamics

097

.1967. The pheasant decline. South Dakota Department of Game, Fish &

Parks, Pierre, USA. Notes: Page 5-". ..such things as blizzards, pesticides,
herbicides, fertilizers, fall plowing, corn chopping, an increase in alfalfa, more livestock
grazing, less cover for pheasants to live in, predators, cold springs, low spring humidity,
too much hot weather at hatching time, and diseases — all tend to lower pheasant
numbers." "Blizzards kill pheasants. A good example was the blizzard of March, 1966.
About 85 percent of our pheasants in several counties in north central South Dakota
were killed. When blizzards strike, they not only reduce the winter population, but the
next hatching season is not enough to replace birds lost during the winter." Page 10-
"... years of late springs have been poor pheasant years... low April temperatures
correlate with our "low" pheasant years, especially before 1959." Page 1 1 - "...the
nesting or brooding instinct depends on other factors, one of which is weather. When
April temperatures are below average, egg laying goes on, but serious nesting is
delayed. Thus less time is left for brooding a clutch. Late springs mean fewer broods
are produced before the nesting cycle is thrown out of gear in July, and results in a poor
pheasant year... The stress of laying extra eggs in years of late springs resulted in poor
pheasant years and high adult pheasant mortality." Page 12 - "Really poor pheasant
reproduction occurred in 1950, 1959, and 1964 when temperatures rose to 94 degrees
or higher in both May and June.... cool May and June temperatures are an indicator of
how many pheasant young will be produced. Freezing and flooding... affect pheasant
nesting... dips occurred 4 to 5 weeks after severe thundershowers." Page 13 - "One of
the Department's biologists voiced the opinion in 1964 that low humidity during
pheasant nesting was one of the causes for a poor hatch." Page 16 - "A one-year
drought in our modern day and age has a disastrous effect on the pheasant population,
not only for that year, but for the next." Page 28 - "The ups and downs of any animal
population is just as natural as changes in the weather. It is even fair to say, dependent
on changes in the weather.' The weather furnishes them a poor house or a good house,
depending on how much cover is grown by the rains which may or may not fall. It
freezes or bakes them. Normally they can take such conditions admirably well — but
they are vulnerable during reproduction." Page 29 - "In 1959, a drought year with a
substantial pheasant decline, we tested pheasant blood and found over half of our birds
had been infected with the virus [equine encephalitis]. In 1960 and 1961, good
pheasant years, we found very few pheasants reacting to the blood tests." Key words:
ring-necked pheasant, temperature, precipitation, flood, reproduction, mortality,
disease, drought, thundershower, snow, humidity

098

Dale, F. H. 1942. Influence of rainfall and soil on Hungarian partridges and
pheasants in southeastern Michigan. Journal of Wildlife Management 6(1):17-18.

Summary: From the differences in habitat requirements of the two species, it seems

4-

*->' ^ ^c

43

Cx~*s~^P <_ _b

likely that the increase in pheasants and the decrease in Hungarians on lake-bed soils
of southeastern Michigan in the latter part of the decade 1920-39 was influenced largely
by the heavier summer precipitation of that period. It also appears likely that the early
success of the Hungarian partridge in this area, on soils of a texture generally regarded
elsewhere as unfavorable to the species, was made possible by a succession of years
with less than average rainfall. Key words: Hungarian partridge, pheasant,
precipitation, productivity, soil

099

Dalke, P. D. 1943. Effect of winter weather on the feeding habits of pheasants in
southern Michigan. Journal of Wildlife Management 7(3):343-344. Notes: Dalke
states that snow appeared to not exert significant impact on pheasant feeding in
southern Michigan in 1929-33. They fed along sheltered spots in marshes or along
steep ditch banks where grass was available. The author states "A sleet storm which
covers all vegetation with ice is not a real hazard to feeding unless the coating is an
inch or more in thickness, or unless it is followed by heavy snow." Dalke found
pheasants dug out food covered with ice using their bills; in fact, in March 1933 these
birds pecked through nearly an inch of ice to eat corn on the cob available in crop fields.
When ice reached Va inch thick, it does restrict the kinds of food available. The birds
then shifted their feeding to berries which remained ice-free on one side. "In one case a
flock of nine pheasants discovered and fed exclusively upon frost grade throughout a
severe sleet storm." One negative impact is the "balling up" of pheasant tails by slush
and ice, making flight difficult or impossible, and walking to search for food becomes
very difficult. "Temperature does... seem to have a pronounced effect on mobility. The
birds prefer to stand in protected fence rows of shrubs or herbaceous growth rather
than walk about and expose themselves... Rainstorms do not keep pheasants under
cover unless the showers are very heavy." Key words: pheasant, snow,
precipitation, ice, temperature, movement, habitat use, nutrition

100

, D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. F. Schlatterer. 1963.

Ecology, productivity and management of sage-grouse in Idaho. Journal of
Wildlife Management 27(4):81 1-841. Abstract: A study of the seasonal movements,
productivity, and management of sage-grouse {Centrocercus urophasianus) was
undertaken by the Idaho Cooperative Wildlife Research Unit from August, 1952, to May,
1960, on an area in Fremont and Clark counties in Idaho, directly west of Yellowstone
National Park. Nineteen individual strutting grounds 1/10=10 acres in size were located
along 12 miles of the Red Road. Summer brood range was found to be 13-27 miles
north and northeast of the Red Road strutting grounds. Flocks of sage-grouse began
migrating west and southwest in October and November and traveled 30-50 miles,
depending upon the depth of the snow. Winter concentrations were usually found where
snow was less than 6 inches deep. Key words: sage-grouse, snow, habitat use,
movement

44 j^

4>

^ ^ ^

?®£ 'wP ^

101

, , , , and . 1960. Seasonal movements and breeding

behavior of sage grouse in Idaho. Transactions of the North American Wildlife
Conference 25:396^*07. Abstract: A study of seasonal movements and strutting
ground behavior of sage grouse was undertaken in 1952 in an area comprising parts of
three counties in southeastern Idaho directly west of Yellowstone National Park. The
study area is mostly high sagebrush plains with scattered irrigated farming, and
livestock grazing is the principal use of the sagebrush-grass lands. Sage grouse travel
50 to 100 miles from the summer range, westerly and southwesterly, to winter ranges
which have only 3 to 6 inches of snow for short periods. Adult males arrive on their
strutting grounds on the Red Road area north of St. Anthony during late March while
snow is still on the ground. The breeding population is at a maximum on the Red Road
strutting grounds from April 7 to 21 during most years. Marking of males and females
show movements up to 3.3 miles. Females are more likely to move to other strutting
grounds than males. The annual high-count of males on strutting grounds is a useful
population trend tool, but may, in some years, fail to provide the game manager with the
means of forecasting the annual crop of sage grouse in some areas. The numbers of
grouse returning to the strutting grounds in 1959 indicated a high yield of subadult birds.
Adverse weather conditions at the time when 75% of the eggs were hatching resulted in
the lowest population of young birds in the 7 years of the study. Brood-count routes at
least 20 miles in length give reliable results until about July 30; however, there is only a
small decline in the average brood size after July 15. By the third week in July
information is available for reliable forecasting of the annual crop of sage grouse which
will be available to hunters in the September hunting season. Key words: sage
grouse, snow, mortality, population dynamics

102

Davies, R. G. and A. T. Bergerud. 1988. Demography and behavior of ruffed
grouse in British Columbia. Pages 78-121 in A. T. Bergerud and M. W. Gratson,
editors. Adaptive strategies and population ecology of northern grouse,
University of Minnesota Press, Minneapolis, USA. Notes: Page 110 — Role of the
extrinsic environment in population change. Survival of chicks was higher in years when
the weather was warm in June, the time when the chicks were only 2-3 weeks old.
Several studies of grouse have now shown that growth is improved with warm springs.
Chick survival was not correlated with mean or mean-maximum temperatures in May.
Page 113 — In summary, results support a hypothesis that approximately 50% of the
population change was caused by spring weather and chick survival, and 50% by
changing, adult survival rates than may result from predation. Ruffed grouse use snow
as cover to escape goshawks... the greatest decline occurred in the winter of 1972-73,
which had the least snow, consistent with the explanation of the need for snow by ruffed
grouse to escape goshawks. Page 115 — Hypotheses for the different mortalities of gray
hens in colder springs are: (1) physiological stress during incubation and early brood
rearing and (2) differential mortality relative to predation. Page 116— Weather not only
may affect birds directly but may also affect plant growth and hence concealment from
predators. Key words: ruffed grouse, temperature, snow, predation, nutrition,
vegetation

♦->' ^ Nf*

«,'-. <£^ii <Q>

103

Davis, J. R. 1976. Management for Alabama wild turkeys. Federal Aid in Wildlife
Restoration Project W-35, Special Report No. 5. Alabama Department of
Conservation and Natural Resources, Game and Fish Division, Alabama,
Montgomery, USA. Notes: Page 9 - "Severe flooding and other adverse weather
factors cause nest losses... Certain conditions such as adverse weather, disease,
predation and abandonment of poults by the hen are some of the explanations for this
loss of poults (52%)." Key words: turkey, flood, mortality, severe weather

104

Davison, V. E. 1949. Bobwhites on the rise. Charles Scribner's' Sons, New York,
USA. Notes: Page 5 — "Although bobwhites might be very plentiful and afford good
shooting for several years (in Michigan), hard winters sometimes wipe them out over
large areas or reduce their numbers so drastically that it took as much as a decade for
them to become reestablished. It is quite evident that in the bobwhite quail we have a
species whose distribution to the north is delineated by climate. Snow may cause
mortality in several ways. As it piles up in depth over weed seeds and grains, or when
accompanied by low temperatures and high winds, it may bury a covey completely and
imprison them in a cell where they die either from starvation or suffocation. It seems
justifiable that snow and ice are the most destructive climatic agents to the bobwhite
quail on the northern edge of its range. They operate either by cutting off the birds' food
supply or by killing them directly when accompanied by strong winds and low
temperatures. It is the exceptional year which kills the quail, and it has been estimated
that such a winter in the north central region can be expected every 4 to 7 years." Key
words: bobwhite quail, snow, ice, temperature, wind, nutrition, mortality,
distribution

105

Devers, P. K. 2005. Population ecology of and the effects of hunting on ruffed
grouse (Bonasa umbellus) in the southern and central Appalachians.
Dissertation, Virginia Polytechnic Institute and University, Blacksburg, USA.

Notes: "Model 16 received the more support than other models and indicated nest
success was influenced by the interaction between mast production the previous fall
and mean minimum temperature in April and May. Nest success has been found to be
positively correlated with mean minimum temperature in other studies. During cold
springs females may be required to feed more often and for longer periods to meet their
energetic requirements. During these feeding bouts, eggs are exposed to cold
temperatures and predators which may decrease hatchability and increase nest
predation.' Key words: ruffed grouse, temperature, vegetation, nutrition, behavior,
predation, model

106

, D. F. Stauffer, G. W. Norman, D. E. Steffen, D. M. Whitaker, J. D. Sole, T. J.

Allen, S. L. Bittner, D. A. Buehler, J. W. Edwards, D. E. Figert, S. T. Friedhoff, W.

46

^

^

H t* **

d2i

W. Giuliano, C. A. Harper, W. K. Igo, R. L. Kirkpatrick, M. H. Seamster, H. A. Spiker
Jr., D. A. Swanson, and B. C. Tefft. 2007. Ruffed grouse population ecology in the
Appalachian Region. Wildlife Monograph 168:1, The Wildlife Society, Washington,
D.C., USA. Abstract: We posit ruffed grouse in the Appalachian region exhibit a clinal
population structure characterized by changes in life-history strategies. Changes in life
history strategies are in response to gradual changes in forest structure, quality of food
resources, snowfall and accumulation patterns, and predator communities. Key words:
ruffed grouse, snow, life history

107

Doerr, P. D., L. B. Keith, D. H. Rusch, and C. A. Fischer. 1974. Characteristics of
winter feeding aggregations of ruffed grouse in Alberta. Journal of Wildlife
Management 38(4):601-615. Notes: Page 608 -"When sufficient soft snow is present
(about 6-8 inches [15.2-20.3 cm]) grouse may burrow into the snow to roost
immediately after feeding, thereby moderating the effects of severe cold and wind." Key
words: ruffed grouse, snow, temperature, wind, behavior

108

Doherty, K. E. 2004. Fall movement patterns of adult female American woodcock
(Scolopax minor) in the western Great Lakes region. Thesis, University of
Minnesota, Minneapolis, USA. Notes: Decreasing daily low temperature was
negatively correlated with movement distances, with >2/3 of movements >500 m
occurring when the daily low temperature was above the median low temperature of
2.4°C. This suggests that woodcock make fewer large movements to conserve energy
when decreasing temperatures increase metabolic demands. Conservation of energy
has been documented during a drought in Maine, where woodcock ceased to make
flights to nocturnal roosting areas in conditions of low food availability... The positive
relationship of soil porosity was linked to the interaction between rain and porosity,
suggesting woodcock responded to this interaction by making large movements into
new foraging areas that were previously too dry to adequately provide them with their
primary food source, earthworms. The combination of rain and porosity interacting
together exhibited the greatest positive relationship to movement than any of the
predictive variables... woodcock made large movements into previously unused areas
after precipitation events. The strong influence of the interaction of soil porosity and rain
also suggests that habitat use is influenced by prevailing conditions that affect habitat
quality at a particular time. Key words: American woodcock, precipitation, habitat
use, movement, behavior, soil

109

Dorney, R. S. 1959. Relation of hunting, weather, and parasitic disease to
Wisconsin ruffed grouse populations. Dissertation, University of Wisconsin-
Madison, Madison, USA. Notes: Page 64 — "In summation, it appears that low
populations of ruffed grouse respond to warm spring temperatures by having above-
average production and reduced differential adult female mortality (more equal adult
sex ratios): conversely, cold May weather results in low production and high differential
adult female loss. Laying and incubation temperatures (May) appear to be more

% f\^^

47

ti^x^-^tb C- — >

important than pre-laying (April) or hatching (June) temperatures. A possible
explanation for the difference between high and low population-temperature
relationships will be presented later in this report." "...a lightly crusted snow that ruffed
grouse cannot penetrate, through sublimation in cold weather, can crystallize and again
become "soft", allowing birds to burrow under it. Exposure, cloud cover and wind as well
as temperature all effect the qualities of snow." Page 65 — "...mild and open conditions
in northern Wisconsin typically lead to crusted snow, with little or no bare ground
present. General snow conditions in the north, therefore, appeared to be correlated with
above-average fall and winter mortalities computed from flush counts and age ratios.
One exception to the above observations must be mentioned. In Otter Creek in the
winter 1957-58, with little snow present, total annual mortality measured by spring age
ratios and territorial cock counts was very low. Apparently some compensatory factor or
factors must have been present to counteract any unfavorable influence from warm
winter weather." Page 67 — "As far as this study could ascertain, summer weather (June
15-'August) exerts little influence on chick survival." Page 76 — "Adult females are more
heavily parasitized. Furthermore, adult female mortality is closely related to May
temperatures for the low populations only. The poor correlation for high populations
suggests that additional density-dependent factors must be involved in this adult female
loss... This parasite (grapeworm) could be one density-dependent factor involved, since
this nematode has a known devitalizing effect on penned galliformes....This increased
parasitism could interact with the physiological stresses of egg laying and incubation to
accelerate female losses under nesting temperatures that would not ordinarily cause
mortality in low populations." Page 87 — "In summation, my observations appear to
indicate that an interplay of spring weather and parasitic disease are related to
population fluctuations." Page 88 — "A warm May was characterized by above-average
production and an almost even fall adult sex ratio; a cold May, by low production and a
loss in adult females; high populations were more sensitive to May temperatures than
low populations ." Key words: ruffed grouse, temperature, precipitation, snow,
disease, parasites, population dynamics, mortality, reproduction

110

and C. Kabat. 1960. Relation of weather, parasitic disease and hunting to

Wisconsin ruffed grouse populations. Wisconsin Conservation Department
Technical Bulletin No. 20. Wisconsin Conservation Department, Madison, USA.

Notes: Page 31 --"Factors Affecting Population Levels — Weather". ...it appears that
ruffed grouse respond to warm spring temperatures by having higher fall age ratios and
more equal adult sex ratios; conversely, cold May weather results in low production and
high differential adult female loss. Laying and incubation temperatures (May) appear to
be more important than pre-laying (April) or hatching (June) temperatures. Low
populations respond to favorable May temperatures better than do high populations
since summer chick loss is about 12 percent in "low" years compared to 23 percent in
"high" years. Winter: "General snow conditions in the north appeared to be correlated
with above-average fall and winter mortalities computed from flush counts and age
ratios. [One exception is described.] The possibility that density-related factors may be
involved in over-winter loss cannot be excluded. It appears that above average winter
temperatures may be unfavorable to grouse in northern Wisconsin. The exact
mechanism of this relationship must await more detailed ecological research. Summer:

48 y

4*

>-> ^ **

"As far as we could ascertain, summer weather (June 15-August) exerts little influence
on chick survival. Losses due to cold and rainy weather immediately following hatching
might logically be expected." Key words: ruffed grouse, temperature, snow,
precipitation, productivity, mortality, sex ratio, age ratio,

111

and A. C. Todd. 1960. Spring incidence of ruffed grouse blood parasites.

Journal of Parasitology 46(6):687-694. Summary: Occurrence of blood parasites in
Wisconsin ruffed grouse was examined. May temperatures in 1954 during the peak of
the breeding season were extremely cold, in contrast to a warm May in 1955. The
percentage of red blood cells containing Haemoproteus gametocytes was 2.4 times
higher in late May 1954 as compared to levels at the same time in 1955. Perhaps the
abnormally cold temperatures directly and/or indirectly may have acted as an additional
stress causing a more severe "relapse" in 1954. The specific mortality and production
during the 2 years 1954 and 1955 were as follows. In 1954, the spring breeding
population was at a low point in the 10-year period, 1949-1958. Weather was extremely
cold in May with a 6-inch snowfall interrupting cock drumming early in the month.
Production of young was poor (average summer brood size 6.8) and adult sex ratios in
fall considerably distorted, 58 males:42 females, suggesting a heavy summer hen loss.
In 1955 the spring was strikingly different than that of 1954, with above average
temperatures. Production of young was high with summer brood sizes averaging 8.5.
Adult female loss in summer was low as indicated by a fall sex ratio of 52 males:48
females. Yet, adult male survival over the 12-month period to spring 1956 was
considerably poorer with a calculated mortality of 48 percent, double that of the
preceding year. The increased adult female mortality in 1954 could conceivably be due
to the combined effect of unfavorable weather in May for nesting, and infection with
blood protozoa. Factors unimportant in years of low population density may become
limiting at high densities. Key words: ruffed grouse, temperature, snow, disease,
parasites, mortality, population density, sex ratio

112

Doucette, D. R. and S. Reebs. 1994. Influence of temperature and other factors on
the daily roosting times of mourning doves. Canadian Journal of Zoology
72:1287-1290. Abstract: From November 1992 to February 1993, observations were
made during 30 departures and 30 arrivals at a mourning dove (Zenaida macroura)
roost in Moncton, New Brunswick, Canada. Our objective was to identify the effect of
cold on the timing of roosting flights in this species, a recent addition to the local
wintering fauna. The effect of other environmental factors was taken into account by
including them, along with temperature, in a multiple regression analysis. Doves left the
roost later relative to sunrise (i) on longer days, (ii) on cloudy mornings, (iii) when fewer
birds were using the roosts, (iv) on colder mornings, and (v) when winds were high.
They returned to the roost later relative to sunset (i) on colder evenings and (ii) in clear
weather. Late arrivals on colder days represent an unusual finding. Anatomical and
behavioral considerations suggest that mourning doves cannot reduce heat loss as
substantially as other species; therefore, late arrivals on cold evenings may reflect the
more important role of energy gain through extended foraging required to survive the

% f \ ^ N^C

49

% m & &

long winter night. Key words: mourning dove, temperature, clouds, wind, behavior,
thermoregulation

113

Drovetskii, S. V. 1997. Spring social organization, habitat use, diet, and body
mass dynamics of hazel grouse Bonasa bonasia in northeastern Asia. Wildlife
Biology 3(3/4):251-259. Abstract: Hazel grouse Bonasa bonasia encounter more
severe climates in northeastern Asia than anywhere else in their range, yet the time
between snow melt and laying is shorter here than elsewhere. Birds were able to lay
early because they moved to follow the phenology of snow melt, changing their diet as
they did so. Before snow melt, habitat distribution and diet were the same as in winter.
As soon as the first snow-free patches occurred on terraces, grouse moved there and
fed on thawed berries, which were available in great quantities on these terraces only.
This habitat shift coincided with the shift in diet; winter foods such as twigs, buds and
catkins decreased from 75 to 3%, whereas the proportion of berries increased from 6 to
70%. Females began to lay a few days after all snow had melted, returning to riparian
habitats where the variety and abundance of plants were greatest. Foliage increased
from 27 to 72% of the diet, and the proportion of berries declined from 70 to 19%. In
winter, Asian hazel grouse primarily occur in flocks. In spring males guard their mates
rather than their territories; they follow females up to a few kilometers from their riparian
breeding habitats, as these move on to terraces to gain weight for egg laying. Such food
related movements between habitats have not been reported for this species in Europe.
Key words: hazel grouse, snow, diet, habitat use, movement

114

. 1992. Materials on the ecology of the hazel grouse Tetrastes bonasia in

south Magadan Oblast in winter. Zoologicheskii Zhurnal 71(4):45-59. Abstract:
Influence of weather parameters on biotopical distribution, feeding content, food quality
and hazel grouse population structure were investigated by the author from October
1988 till March 1989 in the Yana River valley [Russia] (59°59'N, 150°18'E). Generally,
hazel grouse occurred in mixed forests in river valleys (43% of encounters; n = 339).
Usage of other biotopes was less common and depended on weather conditions. Alder
(Alnaster fruticosus) catkins were the most important food item (68% of dry mass of
crop content; n = 55). In very cold periods birds fed on chosenia {Chosenia arbutifolia)
twigs (67%). Flock size depending on the habitat size was 5, 8-5, 6 birds (n = 58) in
average. 1988/89 wintering population consisted of adult birds mainly (88,6%, n = 79).
Key words: hazel grouse, temperature, habitat use, behavior

115

and S. Rohwer. 2000. Habitat use, chick survival and density of Caucasian

black grouse Tetrao mlokosiewiczi. Wildlife Biology 6(4):233-240. Abstract: We
surveyed Caucasian black grouse Tetrao mlokosiewiczi residing on the Lagonakskiy
Ridge, northwestern Caucasus, Russia, in July-July 1998, and on the Magisho Ridge in
July 1999. Weather affected habitat use by brood hens: on sunny days 18 broods were
encountered in meadows, six in ravines, and three in pine forests; on wet days broods
moved to the ridge tops (n = 9) and only one was observed in pine forest. Subadult

50 ^

4*

ft <CSr **

<£^>

males used the same habitats as females with broods. Seven of eight adult males were
encountered in ravines; only one was encountered in a meadow. Habitat use by adult
males was not affected by weather and differed from habitat use by females and
subadult males. Well-camouflaged females, chicks and subadult males used relatively
open, food-rich habitats, whereas black adult males preferred ravines, where nutrition
was poor, but where tall grass protected them from aerial predators. One nest with five
hatched eggs was found. In broods an average of one chick per 10 days was lost. A
goshawk Accipiter gentilis killed one adult female. Our density estimate of 2.3
adults/km2 for the Lagonakskiy Ridge was similar to densities reported elsewhere
(2.3±1.2; n = 7). Key words: black grouse, temperature, precipitation, habitat use,
behavior

116

Duck, L. G. 1943. Seasonal movements of bobwhite quail in northwestern
Oklahoma. Journal of Wildlife Management 7(4):365-368. Notes: Page 367 - "There
appears to be a greater shift of birds from uplands to bottomlands and dunes during
winters of severe weather. The winter of 1939-40 was one of the most severe
experienced in northwestern Oklahoma for some years and considerable mortality of
bobwhite quail occurred... The general trend of the shift... [is] from poor cover to good
winter range on the approach of cold weather... observations here and elsewhere
indicate that the richness of the habitat in relation with severity of winter weather
determines the proportion of birds forced to move.... seasonal movements are
intensified during winters of severe weather, indicating that the movement is forced."
Keywords: bobwhite quail, temperature, behavior, movement, habitat quality,
mortality, severe weather

117

Duke, G. E. 1966. Reliability of censuses of singing male woodcocks. Journal of
Wildlife Management 30(4):697-707. Abstract: Factors affecting the courtship
performance of woodcocks (Philohela minor) and the accuracy of annual singing ground
survey for the U. S. Fish and Wildlife Service were studied in 1963 and 1964.
Woodcock courtship calls were counted at 2-min. intervals, alternating each day
between several areas of differing woodcock density. At the same time, cooperators
made the standard singing ground survey through the study area each evening.
Climatic conditions were measured for each woodcock performance. The most stable
courtship activity occurred from about April 15 to May 15 and appeared to be affected
only by extremes in climatic conditions. The performance seemed to be initiated by light
intensities averaging 2.1 footcandles, so that starting times were earlier in relation to
official sunset when the cloud cover was heavier. Male activity increased briefly with the
coincident appearance of many broods. As the density of performing males increased,
the level of preening activity per bird significantly decreased; courtship flight activity was
less affected. As density increased, the probability of hearing individual males was
reduced. The predawn performance was deemed unsuitable for surveys. Poor hearing
ability of survey cooperators lowered the survey results significantly. The inclusion of
flight songs was not detrimental to the daily tally, nor was the survey significantly

f >* fc* >f*

<£2& -"car- 4MttMf> ^>

affected by stage of the moon. Recommendations for singing ground survey changes
are presented. Key words: woodcock, census, clouds, solar radiation

118

Dunn, P. O. and C. E. Braun. 1986. Late summer-spring movements of juvenile
sage-grouse. Wilson Bulletin 98(1):83-92. Abstract: Late summer to early spring
movements of radio-marked juvenile sage-grouse (Centrocercus urophasianus) were
studied on Cold Spring Mountain, northwestern Colorado, from July to February 1981-
82 and August to May 1982-83. Movements were analyzed from 118 locations (N=8
grouse) during July-November 1981 and 213 locations (A/=10 grouse) during August-
November 1982. Grouse steadily moved away from capture sites until November each
year when they moved to winter-use sites. Movements to wintering areas in late
November were related to snowfall and subsequent availability of sagebrush. Maximum
one-way distance to wintering areas was 30.3 km (A/=4 radio-marked grouse). Sage-
grouse generally followed topographic features and avoided areas without sagebrush
cover, although they were capable of long-distance (23 km) movements over areas
without shrub cover. During spring recruitment there appeared to be roving groups of
males, probably yearlings, that spent much of the breeding season displaying near
females away from traditional leks. Key words: sage-grouse, snow, movement,
vegetation

119

Duriez, O., H. Fritz, F. Binet, Y. Tremblay, and Y. Ferrand. 2004. Individual activity
rates in wintering Eurasian woodcocks: starvation versus predation risk trade-
off? Association for the Study of Animal Behavior, published online 30
November, Elsevier Ltd. Abstract: Wintering birds face a trade-off between starvation
and predation risk. In Eurasian woodcocks, Scolopax rusticola, habitat use may reflect
this trade-off because meadows, where most birds spend the night, are characterized
by a higher risk of predation and a higher biomass of food (earthworms) than the
woods, used by day. We monitored activity of 34 woodcocks fitted with tiltswitch
radiotags. Young birds were more active than adults, probably because they were less
efficient at foraging. In general, nocturnal activity was inversely correlated with air
temperature and with daylight foraging activity, suggesting some compensatory
mechanism, modulated by thermoregulatory constraints. Individual activity patterns
differ, and we classified woodcocks according to three main wintering strategies:
'always', 'sometimes' or 'never' visiting fields at night. The decision to fly to fields at
night seemed to be taken every evening, according to the amount of daylight foraging
activity in woods and the air temperature. After feeding in a rich patch of food on a mild
day, woodcocks did not have to risk going to meadows. Conversely, in patches of fewer
food resources or at lower temperatures or both, woodcocks could not meet all their
energy requirements without going to fields at night (where there was always sufficient
food) and eventually, changing their diurnal sites. Therefore, the trade-off between
feeding and predation risk depends on how efficiently birds find a rich patch of food in
the forest and exploit it optimally during the day. Key words: woodcock, temperature,
thermoregulation, habitat use, nutrition, predation

52 ^

fa l& ^

120

Dusek, G. L., C. D. Eustace, and J. G. Peterson. 2002. The ecology and status of
sage-grouse in Montana. Intermountain Journal of Science 8(2):67-81. Abstract:
We describe the ecology and status of the greater sage-grouse (Centrocercus
urophasianus) in Montana as part of an effort to develop a species conservation plan.
Sage-grouse are primarily associated with big sagebrush (Artemisia tridentata)-
grassland although the original range has been greatly reduced or fragmented by a
variety of human uses and activities. Efforts by the State's wildlife agency to delineate
distribution of sage-grouse in Montana during the 1960s and 1970s suggested that
sage-grouse occupied about 4.4 million ha in eastern and southwest Montana although
more recent efforts to assess sage-grouse habitat suggest occupied habitat could be as
much as 10.9 million ha. Findings from studies during that period suggested that
yearlong distribution and movements reflect regional or local conditions. That is, sage-
grouse tend to be nonmigratory in eastern Montana, where close interspersion of
seasonal habitats rarely requires large movements, and migratory in the intermountain
valleys of southwest Montana. Habitat requirements of sage-grouse vary seasonally, in
terms of structure and composition, to accommodate successful breeding and brood
rearing and over-winter survival. Yearly precipitation patterns, in addition to habitat
quality, can affect nesting success and chick survival. Data from statewide wing
collections suggest that productivity of sage-grouse declined from an average of 2.63
juveniles/hen during 1962-1979 to an average of 2.08 juveniles/hen during 1980-1992;
drought conditions were more frequent during the latter period. An estimate of mortality
of sage-grouse during the first year of life approaches 85 percent of which about two-
thirds occurs prior to the opening of the upland bird hunting season in September.
Sage-grouse populations in southwestern Montana have declined from the 1960s
through the 1980s following a period of large-scale sagebrush manipulation and
conversion of native range to cropland. Numbers of birds remain relatively abundant
throughout areas of central and eastern Montana that continue to support large,
unfragmented stands of big sagebrush. Several state-initiated programs offer incentives
to private landowners to maintain or enhance habitat quality for sage-grouse and other
wildlife species. Key words: sage-grouse, drought, precipitation, vegetation,
habitat quality, population dynamics, habitat manipulation

121

Edminster, F. C. 1947. The ruffed grouse. Macmillan, New York, USA. Abstract:
Largely the results of extensive field studies on about 3,000 acres of abandoned
farmland and 2d-growth beech-birch-maple-hemlock forest near Ithaca, New York, from
1930 to 1941. Following a popularized general account of the ruffed grouse's life history
are presentations of basic ecological and management data. Shelter is considered the
probable principal factor limiting abundance in the northeastern states. For optimum
benefits, overgrown land or slashings and uneven aged hardwoods, mixed woods and
coniferous woods must be thoroughly interspersed with all points within 300 ft. of an
edge. Extensive areas of continuous woodlands may be improved by properly spaced
clear-cut areas. Protection from grazing, half-cutting small trees, light release cuttings of
conifers, woods border establishments, creation and maintenance of 30-ft wide woods
roads, slashings, preservation of selected "weed tree" species and provision of

% &(&**

53

<f5& -*isr- *££$> c£^

drumming and dusting places are among the forest management measures suggested
for use where consistent with the primary economic values of the area. The range
carrying capacity, rather than predators, disease or other apparently limiting factors,
generally seems to determine grouse populations. Carrying capacities vary from 4 to 20
acres per bird. Ten to 100-acre refuges are occasionally warrantable. While abnormal
weather conditions may prove disastrous to young grouse, average early-season losses
are still not fully explained. All notable grouse declines in New York since 1890 occurred
after 2 years of severe February-March snow and cold conditions followed by a very
cold June. The details however were not uniform and the declines do not seem to follow
a truly cyclic pattern. No authentic case is known of grouse being killed in snow roosts
by imprisoning ice, despite hearsay to the contrary. Red and gray foxes, the great
horned owl, 2 weasels and several accipitrine hawks take many grouse, but predation is
not a limiting factor. Experts in predator control show definitely that such control is
costly and of negative value; the "vermin" shooting hunter is a detriment to grouse
conservation. A number of diseases and parasites affecting grouse are discussed, but
disease is not considered to be a general primary morality agency. Grouse foods,
especially the plants, are discussed in detail but deaths due to starvation were found
negligible. Both chicks and adults are independent of open water. Keywords: ruffed
grouse, snow, temperature, ice, mortality

122

Edwards, W. R. 1972. Quail, land use, and weather in Illinois, 1956-70.
Proceedings of the National Quail Symposium 1:174-181. Abstract: Bobwhite quail
(Colinus virginianus) populations were at a 15-year high in 1968 and 1969 in Illinois.
Analysis indicated that quail abundance over the years from 1956 through 1970 was
significantly correlated with changes in land use and weather. Adverse effects
associated with increased acreages of row crops and reduced acreages of oats
appeared to be offset by aspects of land use, favorable to quail, that were associated
with reduced acreages of harvested hay. Quail also appeared to be adversely affected
by heavy snow, above-normal rainfall in late winter and spring, heavy summer rains,
and drought in summer and fall. Key words: bobwhite quail, precipitation, snow,
drought, mortality

123

, P. J. Mikolaj, and E. A. Leite. 1964. Implications from winter: spring

weights of pheasants. Journal of Wildlife Management 28(2):270-279. Abstract:
Ecological implications regarding reproductive condition were obtained from weights of
pheasants (Phasianus colochicus) killed accidentally in Ohio from January through
June, 1961 . Weights of female pheasants averaged 35.0 ounces. Hens had a low mean
weight of about 31 ounces in early January and a high of about 37 ounces in early May.
Weights of cocks had a mean of 44.8 ounces but did not exhibit as great a
proportionate weight change, although similar seasonal trends were evident. Mean
weights of both sexes declined during the nesting season. On the basis of data from
this and other studies, it was postulated (1 ) that in some years the reproductive success
of pheasants is partially controlled by the severity of winter weather and its subsequent
effect on the physical condition of hens in early spring, and (2) that there is an

54 jj

4>

>*< (& **

c£^

increasing probability of excessive hen mortality and lowered reproductive success as
weights in early May fall below about 37 ounces. Key words: ring-necked pheasant,
severe weather, mortality, reproduction, nutrition

124

Eiberle, K. 1987. Effects of air temperature and precipitation on the annual bag of
grouse {Tetraoninae). Cahiers d'Ethologie Appliquee 7(2): 109-1 28. Abstract: The
present paper deals with the effects of the weather on the annual bag of capercaillie,
black grouse, hazel hen and ptarmigan in the canton of Grison (Switzerland). The
investigation covers the period 1 91 9-1 979, comprising 41 , 44 or 59 annual bags,
depending on the game species concerned. The results showed that under the given
general climate the grouse bag was influenced by climatic factors to a remarkably high
degree. The effects of the weather during early rearing and the long-term living
conditions in winter, in particular, proved to be important key factors in shaping the
pattern of abundance dynamics in this central alpine region. The bird species involved
showed different sensitivity with respect to the individual climatic factors. Key words:
capercaillie, black grouse, hazel grouse, ptarmigan, precipitation, temperature,
abundance

125

Eierl, K. 1980. Effect of the weather on the kill of some game species in the
canton of Graubuenden Switzerland. Zeitschrift fur Jagdwissenschft 26(3):142-
153. Abstract: How the kill of some game species [fox, marten, badger, rock partridge,
ptarmigan, heathcock, hazel grouse, hare and marmot] in the canton of Graubuenden
was affected by annual mean temperature and annual total precipitation was studied
using multiple correlations. A statistically significant relationship between meteorological
factors and kill was found only in the case of those game species on which
meteorological elements had an immediate and great effect on population development.
How effective and important the meteorological factors are depends on the general
character of the climate and the location of the habitats typical of a particular kind of
species. The influence of the weather as compared to that of predators was dominant in
the canton of Graubuenden. The influence of predators on prey populations should not
be overestimated because a considerable amount of ecological compensation must be
expected. The fact that low precipitation was advantageous to many game species is
typical for the characteristic climate existing in the canton of Graubuenden. The lack of
habitats at optimum heat level which is due to high altitude is not compensated for by
the continental character of the climate. Keywords: hazel grouse, heathcock,
ptarmigan, rock partridge, temperature, precipitation

126

Einarsen, A. S. 1945. Some factors affecting ring-necked pheasant population
density. Murrelet 26(1):2-9. Summary: The excessively wet nesting season in 1942
adversely affected hatching and chick survival. The weather at hatching time in 1942
was wetter than at any previous hatching season, and may have been a vital factor in
reducing hatches for that year. Another effect of weather upon chicks is the scarcity of
insects due to the cool and damp climate. Few dead chicks were found during any

)->' ^ n£*C

4-

period, however. Key words: ring-necked pheasant, precipitation, temperature,
chick survival, hatching

127

. 1946. The importance of weather data in wildlife management. Murrelet

27(2):28-33. Notes: Einarsen discusses the importance of temperature and precipitation
in relation to pheasant production and survival as well as to other species. He states
"...in much of our game habitat the customary records are not as satisfactory as records
where both weather and habitat conditions are observed... The effects of weather on
wildlife are too great to be overlooked or relegated to a place of minor importance. The
establishment of weather-data collecting stations in correlation with wildlife and range
management programs is essentially a sound practice. The facilities of existing weather
stations can be more useful if their programs are altered to include supplemental
observation points and the addition of certain pertinent data in problem areas." Key
words: ring-necked pheasant, temperature, precipitation, reproduction, weather
data, vegetation

128

Eng, R. L. 1954. Upland game bird survey and investigations; Sage-grouse brood
studies, 1953. Job Completion Report, Project No. W-38-R-5, Volume V, No. 1,
Montana Fish and Game Department, Helena. Summary: (1) Brood studies were
continued in 1953 on established routes on the same trend areas as used in 1952.
Additional spot counts were made by project personnel and brood count data was
collected by deputy wardens in their districts. (2) Temperature and precipitation data for
the trend areas show adverse conditions consisting of excessive rains and cool
temperatures which prevailed throughout the initial incubation and hatching peak in
1953. This condition is in contrast to the mild dry weather in 1952. (3) Low production
was observed on the trend areas in 1953 as determined by a decrease of 16 percent in
the breeding population, an increase of 65.7 percent in the percentage of adults in the
populations during the brood season, and a 10.6 percent decrease in the average brood
size. (4) Yearly differences in production appear to be reflected to a greater extent in
the percent of successful females and/or the percent of adults in the population
throughout the brood season than the average brood sizes. (5) Comparison of
production data from the trend areas to that from a similar habitat experiencing no
adverse weather during 1953 further illustrates the importance of favorable weather to
yearly production of sage-grouse. Key words: sage-grouse, temperature,
precipitation, productivity, population dynamics

129

and P. Schladweiler. 1972. Sage grouse winter movements and habitat use

in central Montana. Journal of Wildlife Management 36(1):141-146. Abstract:
Movements and habitat use by sage grouse (Centrocercus urophasianus) were studied
in central Montana during the winters of 1965-66 and 1966-67. Two and three female
sage grouse were radio-equipped and tracked during the two respective winters. Winter
ranges of the five instrumented females ranged from approximately 2,615 to 7,760
acres. A4-square-mile primary study area, containing over half of the relocations of the

56 j^

4>

^^^

five instrumented birds, was separated into two big sagebrush (Artemisia tridentata)
canopy cover classes on 1 6-inch: 1 -mile aerial photographs. Fifty-five percent of the
primary study area was in the more dense (over 20 percent canopy coverage) and 45
percent in the less dense (under 20 percent canopy coverage) category. Observed use
of the two canopy coverage classes was significantly (P < 0.01 ) different, a decided
preference for the more dense stands being indicated. The characteristics of central
Montana sage grouse winter areas (large expanses of dense sagebrush with little if any
slope) make them prime targets of sagebrush control programs. Removal of sagebrush
from these areas would greatly reduce their capacity to support wintering sage grouse.
Key words: sage grouse, habitat use, snow, vegetation

130

Erikstad, K. E. 1986. Relationship between weather, body condition, and
incubation rhythm in willow grouse Lagopus lagopus lagopus. Fauna Norvegica
Seris C Cinclus 9(1):7-12. Abstract: The incubation of 18 willow grouse Lagopus
lagopus lagopus hens was studied during two breeding seasons. Incubation constancy
of hens averaged 95.5±1 .9% (range 90.8-97.5), number of recesses per day 3.4±1 .1
(range 2.5-8) and the mean length of recesses 19.2±10.3 min. (range 10.9-30.6).
Yearling hens weighed less than adults and tended to leave the nest more often and for
longer periods. Hens in good physical condition took few recesses and lost about 30%
of their body weight. Hens in poor physical condition took more recesses and lost little
or no weight. Hens in good condition preferred to leave the nest during the night and
early morning when ambient temperatures were less favorable. Two captive willow
grouse hens kept in large outdoor enclosures and given additional food increased their
incubation constancy and lost less weight than normal hens. Incubation behavior in
willow grouse is suggested partly to have evolved to minimize the exposure of eggs to
predators, but this behavior conflicts with the energy demands of the hen. Key words:
willow grouse, temperature, behavior, predation

131

. 1985. Growth and survival of willow grouse chicks in relation to home

range size, brood movements and habitat selection. Ornis Scandinavica
16(3):181-190. Summary: Spacing, movements and mortality of willow grouse Lagopus
I. lagopus broods were studied using radio-telemetry during three breeding seasons on
a small island in northern Norway. Chicks survived worse during two cold summers with
few insects than in a warm summer with many. In cold weather, many died at 3-5 days.
Broods selected forests, bogs and fens, and shore vegetation which supported the
largest number of insects. Mean differences between years in spacing and brood
movements were small. Within years, growth was slower and survival lower among
those broods with the greatest mobility and largest home ranges. Broods occupying
small home ranges restricted their movements to areas especially rich in insects,
whereas longer movements were made in poorer habitats. There appeared to be a
dominance hierarchy among broods which effectively prevented them from occupying
the same areas. Key words: willow grouse, temperature, movement, mortality,
insects, nutrition, behavior

-¥ A ^ t . . ,-l. 57

<j, ^>i ^ ^

132

and R. Andersen. 1983. The effects of weather on survival, growth rate and

feeding time in different sized willow grouse broods. Ornis Scandinavica
14(4):249-252. Abstract: During cold and wet summers 0-8 day old chicks from small
(2-6 chicks) willow grouse Lagopus lagopus lagopus broods grew faster and survived
better than chicks from large broods (8-14 chicks). Chicks from large broods spent
longer time being brooded by the hen and had consequently shorter time available for
feeding than chicks from small broods. These results agree with the finding that cold
and wet weather prevents sufficient food intake in willow grouse chicks. Key words:
willow grouse, temperature, precipitation, nutrition, population dynamics,
behavior

133

and . 1982. The influence of weather on food intake, insect prey

selection and feeding behavior in willow grouse in northern Norway. Ornis
Scandinavica 13(3):176-182. Summary: During a warm and dry summer crops of 0-27
day-old chicks contained nearly twice as much food by dry weight as during an
extremely wet and cold summer. Weather did not influence the ratio of insects and
plants in the diet but greatly influenced the insect prey selection. During the warm
summer chicks selectively fed on larvae (80%), while in the cold summer they fed
mainly on smaller insects like Cicadinae, Aphidinae, and adult Diptera. A drop in mean
ambient temperature from 19.5 to 9°C and rain reduced the available feeding time for 2-
5 day-old chicks by about 60%. This reduced feeding time was caused by shorter
feeding periods (mean of 20.8 and 7.8 min) and longer resting periods during evening
and night. Evidence is also presented that in bad weather young chicks avoid habitats
with dense vegetation (which supports the best food supply) to avoid becoming wet.
Extremely cold and wet weather is suggested to be critical in preventing sufficient food
intake in willow grouse chicks. Key words: willow grouse, temperature,
precipitation, nutrition, behavior, habitat use

134

Errington, P. L. and F. N. Hamerstrom Jr. 1938. Observations on the effect of a
spring drought on reproduction in the Hungarian partridge. Condor 40(3):71 -73.

Summary: "On the whole, we doubt that the adverse conditions (drought) of 1934
resulted in any drastic change in the population of the Hungarian partridge in
northwestern Iowa, though they were not without effect. The ecological picture seems to
be essentially one of retardation and decreased productivity of the nesting season
rather than one of ultimate failure.... one of the three breeding seasons (1934) during
which notes were taken was characterized by extreme spring drought that continued
until early June. The spring drought of 1934 naturally was attended by scanty growth of
ground cover. Dry grass and weed clumps of the previous year afforded initial
concealment for many nests, but 'short pastures' forced the farmers to graze their stock
along roads, fence rows, and borders of marshes, with resulting detriment to the nesting
habitats of the birds frequenting such places. Midsummer rainfall was followed by some
recovery of vegetation and improvement of environmental conditions for the partridges.
So far as we can see, the nesting losses of 1934 differed from those of 1933 and 1935

58 ^

4

H to, ^

WW °

principally in the increased scale upon which they occurred, and this in turn may be
attributed in large measure to the exceptionally unsatisfactory status of nesting habitats
during the period of drought. On the whole, we doubt that the adverse conditions of
1934 resulted in any drastic change in the population of the Hungarian partridge in
northwestern Iowa, though they were not without effect. The ecological picture seems to
be essentially one of retardation and decreased productivity of the nesting season
rather than one of ultimate failure. Key words: Hungarian partridge, drought,
vegetation, precipitation, productivity

135

Erwin, M. J. 1975. Comparison of reproductive physiology, molt, and behavior of
the California quail in two years of differing rainfall. M.S. Thesis, University of
California, Berkeley, USA. Unable to obtain for abstract.

136

Etter, S. L., R. E. Warner, G. B. Joselyn, and J. E. Warnock. 1988. Pages 111-127 in
D. L. Hallett, W. R. Edwards, and G. V. Burger, editors. Pheasants: symptoms of
wildlife problems on agricultural lands. North-central Section of The Wildlife
Society, Bloomington, USA. Abstract: Survival of wild ring-necked pheasants
(Phasianus colchicus) in east-central Illinois was studied during 1962-1972 — a period of
abrupt change in agriculture and declining pheasant abundance. During the phase-out
of the Federal Feed Grain Program of the 1960's, increasing farm disturbances and
reduced interspersion of prime cover types rendered the fall-winter landscape less
hospitable for pheasants. In general, pheasants were found to have variable year-to-
year patterns of survival and reproduction, reflecting complex interactions of weather,
characteristics of cover, farming practices, and pheasant movements. Dispersal was
similar between age classes. However, fall-to-early winter survival of juvenile hens was
typically only one-half to two-thirds that of adult hens. Differential survival between
juvenile and adult hens was confined to fall and early winter. For a juvenile hen, the
later the date of hatch, the lower its probability of survival from autumn into winter.
Likelihood of survival was not a simple function of physiological condition as expressed
by body weight, but apparently related to how pheasants responded to changing
weather and farming disturbances in fall, with older, relatively experienced birds more
likely to cope with these changes. Had juvenile hens survived as well as adults,
numbers of pheasants in winter would have been nearly one-third higher. Key words:
ring-necked pheasant, mortality, winter, behavior, nutrition, habitat modification

137

Eustace, C. D. 2002. Sage-grouse hatching success and chronology for south-
central Montana. Intermountain Journal of Sciences 8(2):82-93. Abstract: A recent
short-term fluctuation in sage-grouse (Centrocercus urophasianus) abundance for
south-central Montana was attributed to dramatically reduced productivity. This period
of low productivity was associated with a peak hatching date 2 weeks earlier than
normal. Molt patterns for adult hens indicated the early hatch was due to a shift in
breeding and nesting and not reduced survival during June. These molt patterns also
indicated a significant attempt at renesting by hens > 2 years of age during periods of

1 fVfc*^

59

low productivity. To test the influence of climatic conditions on hatching chronology, I
selected years that were at least one standard deviation above or below the long-term
average for hatches occurring before 1 June. I concluded that early hatches occurred
during warm dry springs, which were also associated with drought conditions. Warm dry
springs also may result in earlier dates for spring green-up and grasshopper hatches. I
interpret early hatches as an adaptation by sage-grouse hens to take the best possible
advantage of cover and forage conditions during the first critical weeks of life for their
chicks Key words: sage-grouse, temperature, precipitation, drought, productivity,
population dynamics

138

Evans, C. A. and S. D. Schemnitz. 2000. Temperature and humidity relationships
of scaled quail nests in southern New Mexico. Proceedings of the National Quail
Symposium 4:116-118. Abstract: We observed unmarked and radio-marked (20
females/1994; 9 females and 11 males/1995) scaled quail (Callipepla squamata) during
the nesting season in the Chihuahuan Desert of southern New Mexico. In 1994, pairing
was completed by early April. Clutch size averaged 13.8 ±1.7 (n=7). Nests were
located an average 216 ± 13.8 m from permanent water. All 97 chicks disappeared from
radio-marked pairs by 16 July. In 1995, all radio-marked females and 6 of the radio-
marked males were paired by mid-April. Clutch size averaged 10.3 ± 1.3 in nests (n=8)
that averaged 545 ± 1 .7 m from permanent water. Almost half of the hatched chicks
(49.6%) fledged in 1995. Nest temperature never exceeded 34°C, while ambient
temperatures reached >43°C. Nest humidity averaged 23%, while ambient humidity
averaged 12%. Notes: Page 117/118 — We hypothesize that the long period of high
temperature exacerbated by drought conditions was primarily responsible for
reproductive failure on JER in 1994. Key words: scaled quail, drought, temperature,
productivity, mortality

139

Evans, K. E. 1968. Characteristics and habitat requirements of the greater prairie
chicken and sharp-tailed grouse — a review of the literature. Conservation
Research Report No. 12. Rocky Mountain Forest and Range Experiment Station,
Fort Collins, USA. Notes: Greater Prairie Chicken Page 8 — "Decimating Factors:
Weather - Weather conditions during nesting are important in determining the number
of young birds available for the fall hunting season. A series of torrential cloudbursts
followed by a long, cold rainy period during the first two weeks of June will cause heavy
brood mortality. Although weather significantly influences the distribution and activities
of prairie chickens, to adequately appraise its part in controlling population numbers is
difficult." Sharp-tailed Grouse Pages 16-17 — " Decimating Factors: Weather - Weather
probably has very little effect on survival of adult sharptails. Sharptails exist in extremely
cold conditions near the Arctic Circle, and in areas of the Great Basin where hot dry
summers are common. Undoubtedly, some adult birds are lost during hailstorms,
blizzards, and ice storms. This loss probably does not greatly reduce the population.
Weather conditions during early June are important in determining brood success.
Torrential cloudbursts, and long, cold rainy spells, during the early brood season
probably reduce the nesting success of sharptails as with all ground nesters. Key

60 j

4>

*->' ^ ^

words: prairie chicken, sharp-tailed grouse, precipitation, temperature, mortality,
cloud burst

140

and A. N. Moen. 1975. Thermal exchange between sharp-tailed grouse

(Pedioecetes phasianellus) and their winter environment. Condor 77(2):160-168.

Summary: Winter energetics of sharp-tailed grouse includes both heat production and
heat loss factors. This report deals with factors influencing nonevaporative heat loss by
grouse during fall and winter on the northern Great Plains. Maintenance metabolic rate,
weight change features, effective surface area, thermal transfer properties,
thermoregulatory behavior, wind velocity, and ambient temperature were analyzed to
facilitate the prediction of energy requirements for homeothermy. The equation
representing nonevaporative heat exchange between a sharp-tailed grouse and its
environment is Q = (KhAh + Kb Ao) ATf. Q is heat exchange (Kcal); Kh and Kb are
thermal conductance values for the head and body, respectively; Ah and Ab are the
effective surface area of the head and body, respectively; AT is the temperature (C)
gradient between deep body temperature and ambient temperature; and f is time.
Thermal conductance values change with wind velocity up to approximately 8 mph and
can be predicted with the following two equations: Kh = 6.50 - [(2.84=VUw)2/10] and Kb
= 2.12 - [(2.84 - VUw)2/10], where \Jw is the wind velocity in mph. Behavioral
thermoregulation involves the selection of habitats and postures to regulate heat
balance. The series of habitats and postures available contains gradients of ambient
temperatures and wind velocities that greatly influence nonevaporative heat dissipation.
Grouse can vary the effective surface area of the neck and heat from 5 to 20% of the
maximum surface area. Maximum surface area (m2) was determined to be (7.46W(g)065)
x 1 0^. A 1 000-g grouse at an ambient temperature of -20°C and an 8 mph wind velocity
in exposed areas could utilize the available habitat and posture gradients to regulate
nonevaporative heat loss between 142 Kcal/day (1.9XBMR) and 319 Kcal/day
(4.3XBMR). Grouse can reduce heat loss by adding insulation such as nocturnal
roosting in the snow or dense vegetative cover. Such responses also reduce wind
velocities over a bird's surface. Overhead cover reduces net radiative heat loss by
protecting a bird's surface from exposure to the night sky, which, when clear, is colder.
Wind velocity gradients also exist through the vertical and horizontal planes of grouse
habitats." Key words: sharp-tailed grouse, wind, temperature, thermoregulation,
behavior, vegetation, bioenergetics

141

Evard, J. O. 1996. Winter weather and pheasant populations and harvests in
northwestern Wisconsin. Research Report 171, Wisconsin Department of Natural
Resources, Grantsburg, USA. Abstract: A study was conducted during 1982-91 in
northwest Wisconsin to examine the effects of winter weather on spring ring-necked
pheasant (Phasianus colchicus) populations and subsequent fall harvests and to
determine pheasant harvest characteristics. The 502 mile2 study area was located in St.
Croix and southern Polk counties, on the northern limit of pheasant range. Winter
weather records were obtained from an Amery weather station, and a winter food
availability index was developed by examining a random sample of 10% of the study

»->' ^ ^

\ *>»■•

area. Crowing pheasant cocks were censused throughout the study area, including
Waterfowl Production Areas (WPAs) and Conservation Reserve Program (CRP) lands.
Hunter bag checks were conducted during the first 2 days of the hunting season, while
other hunters were asked to maintain season-long hunting diaries. Little food, in the
form of standing corn, was available to pheasants in most winters. There was a
significant negative relationship between winter severity and spring pheasant indices.
Crowing cocks declined following winters with prolonged deep snow and subzero
temperatures and increased following mild winters. There was a relationship between
winter weather and spring crowing-cock trends and between spring crowing-cock trends
and fall pheasant harvests. Trends in hunter bag checks during the first 2 days of the
hunting season predicted total season bags. Inexpensive hunter diaries provided
accurate trends in pheasant harvests and hunter success. Key words: ring-necked
pheasant, snow, temperature, nutrition, index, hunting

142

Fallis, A. M. and C. E. Hope. 1950. Observations of ruffed grouse in southern
Ontario with a discussion of cycles. Canadian Field-Naturalist 64(2):82-85.
Abstract: Observations on [Bonasa umbellus] in isolated woodlots suggested that egg
laying was related to weather conditions, especially temperature. Egg laying was not
prolonged by removal of eggs from a nest. Destruction of nests was not increased as a
result of human trails made to them. Second clutches appeared to be the exception
rather than the rule following destruction of eggs that were being incubated. The
irregularity of cyclical fluctuation in grouse populations, and the need for the
accumulation of more quantitative data to explain these periodic fluctuations is
discussed. Key words: ruffed grouse, temperature, nesting

143

Fames, P. E. 1991. A scaled index of winter severity. Proceedings of the Western
Snow Conference 59:12-15. Introduction: Various methods have been used to
represent winter severity and usually include temperature and some measure of
snowfall or winter precipitation. Most represent winter severity as some departure from
normal or as percent of average. Likewise, most use mean monthly temperatures or
monthly snowfall or precipitation. In many cases, the mean monthly temperature does
not fully explain the stresses that are imposed on wildlife during extremely cold periods
that persist for periods less than a month. Also, it is difficult for non-technical people and
even some professionals to quantify how severe or mild a given winter might be. The
scaled index of winter severity (IWS) was developed to assist both wildlife managers
and the general public better understand how mild or severe a given month or season
might be or might have been compared to the historical variations. An index of -4
represents the most severe condition while a +4 represents the mildest condition. An
index of 0 represents average or 50 percent probability of occurrence. Key words:
snow, temperature, index

144

Fields, T. L., G. C. White, W. C. Gilbert, and R. D. Rodgers. 2006. Nest and brood
survival of lesser prairie-chickens in west central Kansas. Journal of Wildlife

62 ^

4-

H (& ^

Management 70(4):931-938. Abstract: We evaluated the effect of habitat use and other
sources of variation on survival of lesser prairie-chicken (Tympanuchus pallidicinctus)
and greater prairie-chicken (Tympanuchus cupido) nests and broods. Daily nest and
brood-survival probabilities were a function of a quadratic time trend, and both declined
as the season progressed. Daily nest survival was negatively associated with nest age,
and daily brood survival was positively associated with brood age. Lastly, broods
tended by adult females had higher daily survival rates than broods reared by subadult
females. The probability of a nest surviving from 10 May to 1 June was 0.72 (SE =
0.06). The probability of a brood surviving from 1 June to 30 July (hatch to 60 days
posthatch) was 0.49 (SE = 0.19) and 0.05 (SE = 0.06). The probability of a brood
surviving from 1 June to 30 July (hatch to 60 days posthatch) was 0.49 (SE = 0.19) and
0.05 (SE = 0.03) for broods reared by adults and subadults, respectively. Although
nesting females and females with broods were using Conservation Reserve Program
grasslands, there appeared to be no benefit to nest and brood survival during our study.
Instead, age of the nest and brood, timing during the season, age of the brooding
female, and precipitation during brooding were more important predictors of survival.
Further experimentation is needed to determine the mechanisms responsible for
decreased nest and brood survival throughout the season. Results from such research
could be used to formulate management strategies to improve nest and brood survival.
Key words: prairie-chicken, precipitation, productivity

145

Fischer, R. A., K. P. Reese, and J. W. Connelly. 1996. Influence of vegetal
moisture content and nest fate on timing of female sage grouse migration.
Condor 98(4):868-872. Summary: Sage grouse most likely obtain the majority of their
water from vegetation, because free-standing water is not readily available or when
available is only occasionally used. Sage grouse inhabit areas having low annual
precipitation. We hypothesized that (1) cumulative annual precipitation and temperature
influence timing of spring and summer vegetal dessication, (2) vegetal moisture content
would be higher in plants collected at sites used by sage grouse than at random sites,
and (3) annual timing of sage grouse migration is related to vegetal moisture content
and nest fate. Timing of both migration and vegetal moisture content varied among
years. During drier years grouse tended to initiate migration earlier in the summer.
During 1991 (a wet year) >12% of females remained on the study area at least through
late July. Our data indicated that there was a threshold of moisture content that
provided a cue to birds to initiate summer migration. During all three years the largest
proportional increase in female migration occurred during the largest decline in vegetal
moisture, when moisture amounts in vegetation declined to approximately 60% water.
Although there was a strong correlation between moisture in vegetation and onset of
migration, our study design precluded us from ruling out possible endogenous factors.
Successful female sage grouse also moved to summer range later than unsuccessful
females, presumably because of brood care. However, during the extremely dry
summer of 1992, even successful females initiated migration early in the summer.
Although timing of migration varied among years, nest success was similar among
years. Our study indicates timing of plant dessication can vary annually with
precipitation. Our data also indicate that during wet summers, fewer grouse moved to
summer range. However, 34 years of precipitation data suggest that there have been

fa tC't* f*

-* Xv- i ... 63

<£^>

few years with precipitation that would have provided succulent vegetal growth
throughout the summer sufficient to attenuate migrational behavior. Key words: sage
grouse, precipitation, vegetation, behavior, movement

146

Flake, L. D., C. P. Lehman, A. P. Leif, M. A. Rumble, and D. J. Thompson. 2006.
The wild turkey in South Dakota. South Dakota Department of Game, Fish and
Parks, Pierre, South Dakota Agriculture Experiment Station, Brookings, USA.

Notes: Chapter 4 - "Snow depths exceeding about 12 inches can essentially stop
movements of wild turkeys on the ground. Wild turkeys can sustain several days of
severe weather, provided high-energy food sources are readily available, but without a
concentrated food supply, birds soon face starvation. Consequently, migrations from
summer to winter ranges may occur in mountainous terrain where snow accumulations
are common. In the central Black Hills, Merriam's turkeys began dying one week after
snow accumulations of about 11 inches, despite mild conditions and eventual snowmelt
within about a week. For approximately four days following this storm, birds stayed in
sunny areas and made no effort to obtain food. Similarly, juvenile hens in the southern
Black Hills began dying within 9 days following 9 inches of snowfall followed by
persistent snow cover. Merriam's turkeys in the Black Hills were found frozen in roost
trees and on the ground below a roost following nighttime conditions of approximately -
30°F with 40 mph winds. Wild turkeys, however, can withstand extremely cold nighttime
temperatures if they have high-energy food available. Eastern and Rio Grande turkeys
in northeastern South Dakota survived temperatures of -30 to -40°F, high winds, and
deep snow as long as cereal grains were available at farmsteads or in windswept crop
fields." Key words: Merriam's turkey, Rio Grande turkey, snow, wind, temperature,
nutrition, mortality, movement

147

Flanders, B. L. 2002. Prairie grouse production on Valentine National Wildlife
Refuge: the effects of weather and grassland management. M.S. Thesis, Colorado
State University, Fort Collins, USA. Abstract: Valentine National Wildlife Refuge,
located in the Sandhills of Nebraska, maintains permanent populations of two prairie
grouse species, greater prairie chicken {Tympanuchus cupido pinnatus) and plains
sharp-tailed grouse {Tympanuchus phasianellus jamesi). A greater prairie-chicken
population increase beginning in the mid-80s was partially equated to increased
vegetative cover on the refuge through improved grassland management, as well as to
favorable weather conditions in the region throughout that time. However, there was no
indication of a similar increase in prairie grouse fall production during the same time
period. Therefore, my goal was to use historical data to assess how vegetation and
weather factors affected production in the fall. ...I assessed the effectiveness of specific
fire and grazing treatments in increasing vegetation visual obstruction, a determinant in
the quality and availability of escape and thermal cover... I then evaluated the validity of
bird harvest age ratios as an index of production.... Next I used linear multiple
regression methods to evaluate the importance of various vegetation and weather
factors on sharp-tailed grouse production. Model-averaged production estimates
provided reasonable predictions of actual production indices, although prediction

64

4*

H ^ ><=*

cci

intervals were large. The most useful predictor variables according to cumulative AlCc
weights were wall weather variables, emphasizing the significant influence of weather
on sharp-tailed grouse production. As hypothesized, 'May Average Temperature', 'June
Average Temperature', and 'Cumulative Precipitation through July 31' were positively
correlated with sharp-tailed grouse production, while 'June Number of Days 35C(95F)'
and 'June Number of Days of Precipitation >2.54 mm (0.10 in)' were negatively
correlated with sharp-tailed grouse production. 'Cumulative Precipitation through July
31' was the largest predictor of sharp-tailed grouse production. Finally, I assessed area
effects using the nearby Samuel "R. McKelvie National Forest, which is more
completely grazed and grazed at a heavier stocking rate than Valentine National
Wildlife Refuge. Averaged across years, the Valentine model over-predicted the real
sharp-tailed grouse production indices on Samuel R. McKelvie National Forest by 0.77
juveniles per adult. Notes: Under Management Implications, Flanders states "It is
apparent that a large amount of the variability in sharptail production from year to year
is influenced by weather. However, that does not mean that managers are rendered
helpless in terms of managing for sharptail populations. Manipulating habitat to increase
vegetative cover may help alleviate negative weather effects that can impair sharptail
production.... In drought years, there may not be adequate vegetation growth to provide
escape cover and shelter from extreme weather events. Therefore managing
grasslands for good standing residual cover provides a sort of insurance policy that
chicks will have sufficient shelter in all years. Of course, weather will always have
effects on extraneous factors, such as insect abundance, that are generally beyond a
manager's control. Poor shelter, low insect abundance, and high predator numbers can
cause sharptail production to crash. Ensuring that chicks have adequate shelter and
escape cover in all years helps buffer sharptail production in years of low insect
abundance... Reductions in stocking rate and/or annually grazed acreage would result
in less vegetation disturbance and increased standing residual cover for sharptails and
other prairie grouse. Key words: sharp-tailed grouse, greater prairie chicken,
precipitation, temperature, vegetation, grazing, habitat modification

148

Flanders-Wanner, B. L, G. C. White, and L. L. McDaniel. 2004. Weather and prairie
grouse: dealing with effects beyond our control. Wildlife Society Bulletin 32(1 ):22-

34. Abstract: We used multiple-linear-regression methods to simultaneously assess
effects of vegetative disturbance and weather on the production of sharp-tailed grouse
(Tympanuchus phasianellus) on Valentine National Wildlife Refuge (NWR) in Nebraska
using a long-term data set of harvest-age ratios as production indices. After developing
the model, we plotted the model-averaged predictions of sharp-tailed grouse production
indices for Valentine NWR against actual sharp-tailed grouse production indices for our
reference area, Samuel R. McKelvie National Forest (NF), Nebraska. Model-averaged
estimates of production provided reasonable predictions of actual production indices on
Valentine NWR, although prediction intervals were large. The most useful predictor
variables according to cumulative Akaike's Information Criterion weights were weather
variables, emphasizing the significant influence of weather on sharp-tailed grouse
production. As hypothesized a priori, "May Average Temperature," "June Average
Temperature," and "Cumulative Precipitation from 1 January-31 July" were positively
correlated with sharp-tailed grouse production, while "June Number of Heat Stress

*~>r fc* **

-* A.Vi ... 65

<£2& -"car- ■^ZJ^ cC^

Days" and "June Number of Days of Precipitation > 2.54 mm" were negatively
correlated with sharp-tailed grouse production. The drought index, Cumulative
Precipitation from 1 January-31 July, explained the most variation in sharp-tailed grouse
production indices. The model developed on Valentine NWR over predicted sharp-tailed
grouse production indices on Samuel R. McKelvie NF by 0.77 juveniles per adult, when
averaged across years. Further experimentation is needed to support our hypothesis
that vegetative disturbance on Samuel R. McKelvie NF is negatively affecting sharp-
tailed grouse production at its current levels. Key words: sharp-tailed grouse,
precipitation, temperature, productivity, brood survival, model

149

Fleming, K. K. and W. F. Porter. 2007. Synchrony in a wild turkey population and
its relationship to spring weather. Journal of Wildlife Management 71(4):1192-
1196. Abstract: Synchrony is an important component of wildlife population dynamics
because it describes spatial pattern in temporal population fluctuations. The strength
and spatial extent of synchrony can provide information about the extrinsic and intrinsic
forces that shape population structure. Wild turkey (Meleagris gallopavo silvestris)
populations undergo annual fluctuations, possibly due to variation in weather during the
reproductive season. To determine if spring weather plays a role in synchronizing wild
turkey populations, we used a modified Mantel-type spatial autocorrelation procedure to
measure the synchrony in fall wild turkey harvest data collected in 443 townships from
1990 to 1995 and compared this to the pattern of synchrony in spring weather variables
(May rainfall and temp) over the same period. We measured correlation using
Spearman correlation coefficients between the total fall harvests from 1990 to 1995 for
each pair of townships, and sorted pairs into 6 50-km distance intervals. We calculated
a mean correlation coefficient for each interval and estimated its P-value using
resampling. We found moderately significant synchrony in the fall harvest (rs = 0.12-
0.34, P<0.008) among township paired <150 km apart, but no significant synchrony
beyond this distance. In contrast, both May temperature (r= 0.82-0.90, P<0.001) and
rainfall {r= 0.49-0.76, P<0.001) were strongly synchronized across all 6 distance
intervals. Visual inspection of time series in the wild turkey fall harvest suggests that
populations may be synchronized in some years when weather promotes high
reproductive success (i.e., a synchronized growth peak) and asynchronous in other
years. Knowledge of the spatial dynamics of wild turkey populations will aid wildlife
managers in estimating population change, setting harvest quotas, and managing
habitat. Keywords: Merriam's turkey, temperature, precipitation, reproduction

150

Formozov, A. N. 1964. Snow cover as an integral factor of the environment and its
importance in the ecology of mammals and birds. Boreal Institute for Northern
Studies, Occasional Publication No. 1, University of Alberta, Edmonton, Canada.

Notes: Formozov devotes several chapters to the description of various types of snow
and its modification through the interaction of other meteorological factors. Page 36 -
"Observations show that for many animals a thick layer of snow which is covered by
nast [ice] is sometimes more dangerous than relatively deep and fluffy snow." Page 42
- "There are indications that black grouse, when followed by goshawk {Accipiter

66 j^

4<

^^^

>®£ <fir> ^

gentilis) plunge into the fluffy snow, digging in fast and deep, as they do in the evening
when they hide for the night." Page 46 - "...ptarmigan obtained their food form under
the snow and partly even from under the icy crust which usually covers the parts of the
tundra which had been blown free of snow by wind. It appeared that the ptarmigan in
this case used the services of reindeer which dug the snow with their feet." Citing
Romanov 1934, "...I often saw how grey partridge had fed at the places where hares
had grazed." Page 47 - For a long time it has been known that flocks of snow
partridges (Tetraogallus caucasicus) congregate in winter at the places where turs
(Capra caucasica) have grazed, an adaptation which can be explained by the same
ease of securing food on the places where the snow cover has been even partly
destroyed by the strong hoofed animals. The above described phenomena of temporary
symbiosis actuated by the winter presence of deep or compact snow cover is a good
example of how specific and finely attuned some biotic relationships are. Undoubtedly
the destruction of deer or hares would aggravate the wintering conditions of ptarmigan
or partridge, and during difficult winters their die-off would be more freguent and
complete." Page 73 - ".. .northern Caucasian pheasants. ..are sharply diminished in
numbers after hard winters, but their numbers are re-established after 2 or 3 easy
years." "There is a marked decrease in the numbers of Manchurian pheasants in
Primoria and Ussuri Krai after winters with increased snowiness." Key words: snow,
temperature, wind, Manchurian pheasant, snow partridge, ptarmigan

151

Forrester, N. D., F. S. Guthery, S. D. Kopp, and W. E. Cohen. 1998. Operative
temperature reduces habitat space for northern bobwhites. Journal of Wildlife
Management 62(4):1 506-1 511. Abstract: High operative temperatures may cause wild
animals to avoid habitat space-time, leading to thermal fragmentation of habitat. We
evaluated thermal fragmentation of habitat space for northern bobwhites (Colinus
virginianus) in a subtropical-subhumid portion of Texas during June 1994-August 1995.
Based on data from 606 random points, 405 flushing points, and 237 landing points,
bobwhites avoided habitat space-time with operative temperatures >39°C during the
hottest period (Jul-Sep). The estimated proportion of habitat space-time avoided
exceeded 0.50 during all seasons and reached maximums of 0.65 for flushing points
(Mar-Jun) and 0.74 for landing points (Jul-Sep). Habitat management that fosters lower
operative temperatures near the ground may increase the abundance of bobwhites in
tropical and subtropical environments. Key words: bobwhite, temperature, habitat
modification, habitat use

152

Francis, W. J. 1970. The influence of weather on population fluctuations in
California quail. Journal of Wildlife Management 34(2):249-266. Abstract: Age ratios
of quail {Lophortyx californicus) in San Luis Obispo County, California, were available
for 14 consecutive years. Ratios were based on hunter-shot samples averaging 672
birds per year. Weather data were compiled and a multiple linear regression of quail
productivity on selected weather parameters revealed a close relationship (P< 0.01).
Quail productivity seemed to be a function, in order of importance, of (1) soil moisture in
late April calculated from temperature and rainfall data, (2) proportion of breeding

X fVfec^

67

w **

females over 1 year old, and (3) the seasonal rainfall from September to April. Three
geographically isolated wild quail populations and one penned population were
observed during two breeding seasons that were very different in productivity. In 1963,
quail produced many young. The breeding period was characterized by intense activity
and persistence of breeding effort extending, in captive birds, to production of second
broods. Vegetation that year included many annual forbs growing vigorously during the
breeding season. In 1964, production of quail was very low on all areas. The birds
seemed to lack reproductive drive, and breeding effort terminated early. Forb vegetation
that year was sparse. Key words: California quail, temperature, precipitation,
vegetation, behavior, nutrition, productivity

153

. 1968. Temperature and humidity conditions in potential pheasant nesting

habitat. Journal of Wildlife Management 32(1):36-46. Abstract: Measurements of
temperature and humidity within different vegetative cover types in Illinois pheasant
range were made, and absolute humidity parameters calculated. Preferred nesting
cover of pheasants (Phasianus colchicus) has been found to be strip cover (such as
fencerows and roadsides) and hayfields. Within these cover types summertime
maximum temperatures remained lower than in other cover types. Saturation deficits
were also lower in the preferred cover types than in other cover types, and were very
high in cornfields and certain grasses. Success of pheasant reproduction appears to be
related to placement of nests in vegetative types offering a favorable microclimate.
Temperature and humidity variations among cover types exceed the mean geographical
variation over a wide area. Key words: pheasant, humidity, temperature,
reproduction, habitat use

154

. 1967. Prediction of California quail populations from weather data. Condor

69(4):405-410. Summary: A method is presented by which the productivity (proportion
of subadults in the fall population) of California quail in San Luis Obispo county,
California, can be predicted from the age-ratio of females, and two weather parameters
(1 ) the soil moisture at the end of April computed by the Thomthwaite
evapotranspiration and water-balance method, and (2) the total seasonal precipitation
from September through April. Tests on independent data verify the predictive value of
the method. Keywords: California quail, index, productivity, precipitation,
evapotranspiration

155

. 1965. The effect of weather on population fluctuations in the California

quail (Lgopus phortyx calif ornicus). Dissertation, University of California,
Berkeley, USA. Abstract: "Discussion: ...The multiple linear regression analysis of the
data for the Shandon area pins down with considerable accuracy the factors that vary in
correlation with reproductive success, i.e., the age ratio of the breeding females, the soil
moisture in the spring, and the total seasonal rainfall. The first of these is so strongly
supported by the observed breeding behavior of birds in captivity, and is also so clearly
independent of the other factors that little discussion is called for... In wild quail, dark

68 ^

4

^fe^

cloudy days in February and March might similarly delay breeding and enable the first
year hens to reach a physiological condition that would permit more effective breeding.
Such light conditions are so closely tied up with rainfall conditions that the effect would
be included in other weather parameters; there does not seem to be any clear indication
from the regression analysis that this is important, the correlation between reproduction
and the temperature range in early spring (measuring the degree of cloudiness) not
being highly significant. The soil moisture calculated for the end of April, the most
important factor in the regression equation, serves to measure the accumulated effect
of the amount and distribution of rainfall as modified by the temperature regime. It is
more useful to think of this parameter as an index of the combined weather effect than
as an accurate measure of the actual soil moisture which could be measured at a given
locality. The latter varies considerably with soil type, exposure, relief, land use, and
perhaps other factors; the former depends only on the rainfall and temperature regime
which is similar over a wide area, in just the same way that variations in quail
populations are similar. It should be noted that this parameter is relatively constant
throughout the early spring, so that it is not merely at the end of April that a high or low
value may have an effect on quail reproduction; the value for any decade depends
strongly on that for the previous decade, as well as on the rainfall during the decade.
The importance of this parameter in the multiple regression equation enables us to say
with confidence that our hypothesis is supported by the data, i.e., that 'soil moisture
storage during the breeding season maintains green growth that stimulates breeding.'
This hypothesis was of course based on numerous reports linking reproductive success
with green growth. We should then attribute production of gonadotropins to some factor
in the vegetation that is available in the early breeding season of California quail. The
precise nature of the link is not shown by this study. Examination of specimens and
observations of the vegetation during my study show that lack of food to the point of
emaciation cannot be a factor in reproductive failure in these populations. Birds which
failed to breed had fat deposits, and green vegetation was available even under the
poor conditions of 1964 in the Shandon area. It is evident that the deficiencies must be
qualitative, and that some factors which are available under normal weather conditions
are lacking in a regime characterized by a low calculated oil moisture. These factors
may include proteins found only in rapidly growing young shoots of new plants, any of
various vitamins, or nutritional factors found only in certain species of plants used as
food by quail. The third factor in the regression equation is the total seasonal rainfall,
which is negatively correlated with the reproduction index. Since it is positively
correlated with the calculated soil moisture storage (r=0.874), this is to be interpreted as
meaning that an excess of rainfall over that required to give the maximum soil moisture
storage will tend to reduce the rate of reproduction somewhat. For example, both 1952
and 1958 had high values of calculated soil moisture in the Shandon area; 1958 had 5
inches more rainfall than did 1952, but a lower immature:adult ratio of 2.46, compared
to 4.30 in 1952. Of the difference, 1.13 can be assigned to the female age ratio (1952
having almost all adults in its breeding population), leaving 0.64 attributed to the greater
rainfall in 1958. This is a relatively large figure, amounting to one-fourth of the total
reproductive rate for this year, and about one-half of the standard deviation of the
immature:adult ratio. The distribution and amounts of rainfall were similar in both years
until mid-winter, but in 1958 heavy rains continued up to the first part of April, while
1952 had appreciable rains only in January and March. The mechanisms by which

% ^y^^k

69

c^

these weather factors affect quail reproduction can only be conjectured. More sunshine
and longer effective day lengths occurred in 1952, but the correlation of reproduction
with early spring temperature range was relatively low; it is possible that light and
sunshine will have an effect on reproduction only in those years when other factors are
favorable. It seems more likely that such differences are related to the plant species
which are abundant under the different rainfall regimes, and to the times at which they
germinate and when they are growing most vigorously... Observations in the present
study, both in the field land in the Behavior Laboratory enclosure, showed striking
differences in the vegetation composition, years of good quail reproduction being those
with an abundance of annual forbs, including legumes, known to be important as quail
foods. Species and growth stage of these plants may be more important to quail
nutritionally than the production of food in quantity. Still another effect of vegetation,
consistent with the heavy growth of grass in the Behavior Laboratory enclosure in 1964
when reproduction was poor, is the observation by Linsdale (1947) of the 'reduction of
California quail which came with the increased density of the ground cover', referring to
dense growth of grass on the Hastings Reservation. To sum up the considerations
discussed above, it appears that the temperature and rainfall regime in a given year is
responsible for the production, at the beginning of the quail breeding season, of highly
nutritious plant foods which, under optimum conditions, supply all the elements (protein,
vitamins, and other) necessary to vigorous reproductive behavior. An appropriate
photoperiod will trigger the beginning of the reproductive season; if nutrients are present
in an adequate amount, gonadotropic secretions of the pituitary will lead to full gonadal
development, as evidenced by pairing, aggressive behavior, vocalizations, mating, etc.,
followed by nesting and egg-laying. Normal environmental stimuli, and perhaps gonadal
feed-back to the pituitary, will then stimulate the production of prolactin and other
hormones associated with incubation and with parental behavior; the latter are more
readily elicited in experienced females than in those breeding for the first time. With
deficiencies of essential nutrients, the pituitary will produce subnormal amounts of both
gonadogropins and prolactin, and both reproductive and parental behavior will be
expressed at a low level, resulting in poor hatches of eggs and poor survival of young."
Key words: California quail, solar radiation, precipitation, temperature, clouds,
vegetation, nutrition, reproduction, soil moisture, index

156

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(2):71 1-722. Abstract: Severe winter weather in
the Northern Great Plains of North American can alter availability of winter cover and
cause increased mortality of ring-necked pheasants (Phasianus colchicus). We
monitored pheasant survival and habitat use via radiotelemetry during the second most
severe winter in eastern South Dakota since 1892. We captured and radiomarked 489
female ring-necked pheasants at the onset of the 1996-97 winter and monitored
survivors through spring at 3 sites in eastern South Dakota. We also monitored 58
female ring-necked pheasants at the same sites during 1995-96 winter, a winter
characterized by below average temperature and average snowfall (winter severity
rank: 35th). Survival of radiomarked hens in 1995-96 (0.61 [SE = 0.07]) was higher (P <
0.001 ) than that in 1996-97 (0.03 [SE = 0.02]). Mortality due to predation was higher (P

70 ^j

4>

>> fc* 1*

±±l*

c£^

< 0.042) than mortality due to weather in both winters. Mortality due to weather did not
differ (P= 0.787) between winters. However, 31 of 41 deaths occurred during blizzard
periods in 1996-97, indicating severe weather increased the vulnerability of pheasants
to predation. Radiomarked hens showed the highest preference for tall grass (> 75 cm),
cattail (Typha spp.) wetland, and corn food plot habitats in winter 1995-96, and early
winter 1996-97. Shelterbelt and corn food plot ranked highest for pheasants that
survived to the second half of the 1996-97 winter. We conclude that shelterbelt and food
plot habitats are essential to the survival of pheasants in eastern South Dakota during
extreme winter weather conditions. Key words: ring-necked pheasant, temperature,
snow, predation, habitat use, mortality, severe weather

157

Gates, R. J. 1983. Sage-grouse, lagomorph, and pronghorn use of a sagebrush
grassland burn sites on the Idaho National Engineering Laboratory. M. S. Thesis,
Montana State University, Bozeman, USA. Notes: Page 24 — "...burn and control
sites were probably snow-free or had minimal snow cover for a longer period in the
winter 1980-81 than in the other 2 winters. This may at least partially explain the
relatively higher grouse pellet density which was observed in 1980-81." Page 44/47 — "
Grouse observations per km of transect declined 72% between the summers of 1980
and 1982. The ratio of juveniles to adult hens observed on the transect indicated low
production of young during the springs of 1980-82. Precipitation during May, the normal
month of peak hatching on the INEL was 2.9 and 1 .8 cm above the 20 year average in
1980 and 1981 . Also, daily temperatures averaged 0.7C and 1 .7C below normal in May
of 1980 and 1981. ..Despite below average precipitation during the spring of 1982
production was still estimated to be low; however, the small number of hens and broods
observed in 1982 may not have accurately reflected reproductive success." Page 47 —
"...during the years of this study in which precipitation during the first 6 months was
above average, grouse moved to summer ranges earlier than in a year when
precipitation was below average. In spite of lower precipitation, succulent forbs were
observed in sagebrush grassland through most of the summer in 1982; whereas during
1980 and 1981 most forbs became desiccated by the end of June. Abnormally high
precipitation (1.3 cm above average) during July and August 1982 may have prolonged
the season of forb growth into July and August 1982." Page 62 — "Since fecal pellet
densities on the burn area varied inversely with the depth and persistence of snow
cover over 3 years, grouse may have concentrated on portions of the study area other
than burn site as snow depth increased. The low pellet density observed in 1982, after
the winter of greatest snow cover, may also have in part reflected the declining
population trend." Key words: sage-grouse, precipitation, snow, vegetation, habitat
use, movement, nutrition, reproduction, habitat manipulation

158

Gatti, R. C, R. T. Dumke, and C. M. Pils. 1989. Habitat use and movements of
female ring-necked pheasants during fall and winter. Journal of Wildlife
Management 53(2):462-475. Abstract: We studied diurnal habitat use and movements
of 56 female ring-necked pheasants (Phasianus colchicus) on private and public lands
in southern Wisconsin from September through April 1968-71 using radio telemetry.

>"V 4c„« -**

% ±»"

Cgj ^ ^ c£J>

Habitat use of female pheasants was a function of month, year, female age, and snow
cover depth, but not female survival through April. Females preferred food patches and
brush, and avoided pastures and active croplands in all 7 months. Shrub-carr wetlands
were preferred by females in all months except October, and marshes were preferred in
all months except January and February. Retired croplands were preferred from
October to December, but avoided in January. Upland hardwoods were avoided in most
months except for periods of deep snow cover. Monthly home-range sizes averaged 32
ha. Home-range size during 10-day periods peaked in late October through early
November, and declined to a low in early January through early February. Juveniles
had larger ranges than adults, and preyed-upon females had larger ranges than
surviving females. Home-range size was positively related to corn harvesting and
pheasant hunting pressure, and negatively related to snow cover depth. Females began
a directional move (mean = 1,150 m) to winter cover in late September. Distance to
winter cover was also a function of female age, female fate, corn harvest, and snow
cover depth. Heavy hunting pressure appeared to temporarily suppress the use of food
patches and strip cover, and accelerate the final move to winter cover, but did not cause
females to move off public lands. Females remained in winter cover from early
November through early February, using a variety of areas and habitats. The
abundance of wetland cover in winter may have been responsible for a lack of winter
concentration and independent moves between adults and juveniles. Key words: ring-
necked pheasant, snow, habitat use, movement

159

Gefell, D. J. 1990. An exploration of the influence of environmental factors on
variation in wild turkey populations. M. S. Thesis, State University of New York,
Syracuse, USA. Abstract: The 2 primary objectives of this study were: (1 ) to explore
the utility of harvest data for assessing absolute and relative wild turkey abundance,
and (2) to assess the correlations of wild turkey density and rate of change in density
with potentially limiting weather, hunter pressure, and land use factors. The first
objective was addressed using wild turkey spring and fall harvest data from townships
in southern New York State during 1969-1981. Two methods of assessing absolute
abundance, and 4 potential indices of relative abundance were examined. Neither
technique for estimating absolute abundance was found to be reliable. However, 1
method of indexing relative wild turkey abundance successfully passed tests of
accuracy and logical consistency. The second objective was addressed by using the
turkey density index to compare how weather, hunter pressure, and land use patterns
each influenced historical township-level wild turkey population variability across
southern New York. Keywords: wild turkey, weather, population dynamics, index

160

Gerstell, R. 1939. Certain mechanics of winter quail losses revealed by laboratory
experimentation. Transactions of the North American Wildlife Conference 4:462-
467. Conclusions: (1 ) That the characteristic huddling habit of the bobwhite quail may
at least in part represent an instinctive reaction which tends to reduce the heat loss from
the bodies of the various individuals which make up any given covey. (2) That, at least
within certain limits, the ability of a covey of quail to withstand low environmental

72 ^

4>

H tefc ^

c£^>

temperatures is directly proportional to the size of that covey. (3) That the body
temperature of individual quail may drop more than 25° F below normal without the
bird's suffering a breakdown of its thermal regulatory system which would result in
death. In closing, it seems fitting to call to the game manager's and sportsmen's
attention the practical importance of the results obtained even from the few experiments
so far completed. This lies in the indication that throughout the northern portion of the
quail range, the bobwhite coveys should not be shot down to a point where their
chances of winter survival are seriously endangered simply by too great a reduction in
the average size of the bevies. Key words: bobwhite quail, temperature, biothermal
regulation, behavior, hunting

161

Gill, R. B. 1966. A literature review on the sage-grouse. Special Report No. 6.
Department of Game, Fish and Parks, Game Research Division and Cooperative
Wildlife Research Unit, Denver, USA. Notes: Page 23 - "Decimating Factors,
Weather. Patterson studied nesting under contrasting weather conditions and
concluded that little or no nesting mortality occurred as a result of rain or snow during
the incubation period. However, periods of cold rain, sleet, or snow during or
immediately following the hatch caused mortality among chicks. Concluded from his
studies in Idaho that above-average rainfall during the months of May and June
depressed production of sage-grouse. Schlatterer continued this study and found that
adverse weather reduced the average brood sizes of birds hatched during the bad
weather. Pyrah reported that high nesting success and brood survival in Wyoming
occurred one year after periods of above average rainfall. He concluded that while
adverse weather depressed production in the year that it occurred, it increased
production the following year." Key words: sage-grouse, precipitation, snow, sleet,
productivity

162

.1966b. Weather and sage-grouse productivity. Outdoor Facts No. 37, Game

Information Leaflet, Colorado Game, Fish and Parks Department, Denver, USA.

Summary: The primary effect of weather on gallinaceous game birds is manifested in
nesting success and survival of young chicks. The two most important weather factors
reported in the literature were temperature and precipitation during the nesting and
early brood rearing season. Investigations conducted in the Lake John area of North
Park, Colorado, indicated that temperature was the most important single factor
affecting sage-grouse productivity. Variations in temperature alone accounted for 68
percent of the variability in strutting male counts the following spring, and the combined
effects of temperature and precipitation accounted for 87 percent of the variation.
Knowledge of the relationship between temperature and production permitted the
calculation of a prediction formula from which reliable predictions of strutting male
counts can be made. In general it was found that when the mean average temperature
for the period April-June exceeded 44.5°F, good production resulted. Also it was found
that the best production resulted from spring seasons which had a mean average
temperature above 45.0T and two or more inches of total precipitation. A scale for
evaluating the combined effects of temperature and precipitation was constructed from

f V ^ ^

% $s?> <Q> &

known and predicted values... One interesting sidelight in connection with these data is
that when spring counts of strutting males were correlated with hunter harvest data on
the previous year, no significant correlation resulted (^0.324). From this it was
apparent that present levels of hunting pressure have had little or no effect on sage-
grouse populations in the Lake John area; instead, the most important factors affecting
these populations were found to be the combined effects of temperature and
precipitation during nesting and early brood rearing. Key words: sage-grouse,
temperature, precipitation, productivity, index, population dynamics

163

Giuliano, W. M. and R. S. Lutz. 1993. Quail and rain: what's the relationship?
Proceedings of the National Quail Symposium 3:64-68. Abstract: We used
Christmas Bird Count reports in conjunction with precipitation data from 9 locations in
Texas, to investigate relationships between rainfall and northern bobwhite (Colinus
virginianus) and scaled quail (Callipepla squamata) abundance. Regional differences in
northern bobwhite abundance could not be predicted by precipitation regimes, whereas
scaled quail abundance was negatively correlated with fall and winter rainfall.
Differences in rainfall patterns were not significantly correlated with year-to-year
changes in northern bobwhite and scaled quail abundance. Key words: bobwhite
quail, scaled quail, precipitation, population dynamics

164

Gjerde, I. and P. Wegge. 1987. Activity patterns of capercaillie, Tetrao urogallus,
during winter. Holarctic Ecology 10(4):286-293. Summary. Winter activity patterns of
16 radio-marked capercaillie, Tetrao urogallus, were studied during 1981-83 at
Varaldskogen, SE Norway. Activity was confined to the light hours of the day, closely
following the photoperiod. Diel distribution showed a major peak near sunset during
every month. A second peak was found in the morning. This peak was comparable to
the evening peak in early and late winter, but nearly disappeared in midwinter. Ambient
temperature rather than photoperiod seemed to be the important proximate factor
responsible for the changes in morning activity. By postponing activity to the afternoon,
capercaillie avoids energy expenditure during the coldest morning hours. Total daily
activity (TDA) during the period November-April averaged about 3 hours for both sexes.
TDA of cocks was U-shaped with the lowest values (2.0 hours) in December-January,
whereas TDA of hens was fairly stable during November-March with mean value of 2.7
hours, increasing abruptly to 4 hours in April. TDA during December-January is believed
to express the time used for feeding in pine trees. The significantly higher TDA of hens
(26%) compared with cocks during this period may be explained by a higher relative
heat loss of hens (body weights: hens 2.0 kg, cocks 4.3 kg). The increased level of
activity during early and late winter among cocks was probably due to display and
related territorial behavior, whereas increased activity of hens during April probably was
caused by a shift from feeding in pines to more time-consuming selective feeding on the
ground. Key words: capercaillie, temperature, behavior

74 ^

4>

H to* ^

cci

165

Glover, F. A. 1948. Winter activities of wild turkey in West Virginia. Journal of
Wildlife Management 12(4):41 6-427. Notes: Under "Summary" the author notes: "The
daily range was reduced markedly by deep snows. At times the birds moved just
between the roosting sites and the feeding areas. During the periods of deep, soft snow
the turkeys traveled through the trees or flew short distances. Turkeys commonly
scratched through snow 12 inches deep to obtain food." Key words: Eastern wild
turkey, snow, movement, habitat use

166

Gobeille, J. E. 1992. The effects of fire on Merriam's turkey brood habitat in
southeastern Montana. M. S. Thesis, Montana State University, Bozeman, USA.

Notes: Page 42 — Poult mortality was difficult to assess due to inability to observe very
young poults in herbaceous cover. However, some broods appeared to suffer heavy
mortality due to wet spring weather... hen #1177 lost 70% of her poults during the first 3
weeks after hatching when 3.5 cm of rain occurred in the Ekalaka Hills. The majority of
this fell in 2 storms over a 4-day period. Key words: Merriam's turkey, precipitation,
mortality

167

Goddard, A. D. 2007. Reproductive success and habitat selection of sharp-tailed
grouse {Tympanuchus phasianellus) in the Peace Reiver region, northeast British
Columbia. M.S. Thesis, University of Northern British Columbia, Prince George,
Canada. Abstract: Sharp-tailed grouse {Tympanuchus phasianellus) are relatively
abundant across the Peace River region. Local biologists and landowners suggest,
however, that populations have been steadily declining over the past several decades,
and little is known regarding the current population status and habitat requirements of
the species. Success during the reproductive season is often the most important factor
limiting population growth of sharp-tailed grouse. To effectively manage populations and
the habitats required by sharp-tailed grouse, it is important that baseline data describing
patterns of reproductive success and habitat selection are collected. Using 15-g radio
collars to monitor female sharp-tailed grouse during 2 consecutive breeding seasons, I
quantified the success of first and re-nest attempts, the survival of offspring to 35 days
of age, and I determined patterns of habitat selection during the nesting and brood-
rearing periods. Nesting success in 2004 and 2005 was 41% and 50%, respectively,
and was positively influenced by habitat conditions at the nest site including increased
shrub and living cover. Seventy-five percent of females successfully reared a minimum
of 1 chick to 35 days of age. Only 35% of chicks, however, survived the same period.
Survival of offspring was positively related to pre-hatching and negatively related to
post-hatching weather conditions as well as the distance travelled by a brood during the
first week after hatching. Breeding sharp-tailed grouse selected habitats in a
hierarchical fashion, selecting habitats at the landscape, patch, and site scales. At the
landscape scale, breeding females selected areas with non-forest cover-types between
550-650 m in elevation. At the scale of the nest site, females selected sites with greater
overhead cover, shrub and grass cover, taller vegetation, and sites with more residual
vegetation than random. During the early part of the brood-rearing period (offspring 0-

-y 75

^

r* 1& ^k

£»~»~^0 C — )

14 days of age), females with broods selected for agricultural habitats rather than
natural habitats. This trend was not observed, however during the late brood-rearing
period (15-49 days of age). In general, habitat selection choices made by breeding
sharp-tailed grouse corresponded with increased fitness in the form of greater
reproductive success. Key words: sharp-tailed grouse, productivity, habitat use,
vegetation

168

Goldstein, D. L. 1984. The thermal environment and its constraint on activity of
desert quail in summer. Auk 101(3):542-550. Abstract: I measured the thermal
environments experienced by Gambel's quail (Callipepla gambelii) during the
summertime in the Clorado Desert by using estimates of standard operative
temperature (Tes). Simultaneously with these measurements, I monitored the activity
pattern of quail. The thermoneutral zone of freshly captured quail extends to 44°C, yet
Teo in sunlit areas may exceed this upper critical temperature for more than 10 hours of
a hot day. Even in the shade, Teo is at the upper limits of thermoneutrality for much of
the day. At these temperatures the birds maintain a body temperature that is very close
to their upper lethal limit. The activity pattern of Gambel's quail on hot summer days is
bimodal, with morning and late afternoon foraging separated by a long quiescent period
during midday. The time of onset and the duration of this inactive period varies directly
with Teoand foraging occurs throughout the day on cool, overcast, summer days. This
suggests that the thermal environment constrains midday activity. A bimodal activity
pattern during the winter is probably a response to predation pressure from avian
raptors. During the summer, however, it is advantageous for growing juvenile quail to
maximize foraging, and the thermal environment is the most important factor shaping
the activity schedule. Key words: Gambel's quail, thermoregulation, temperature,
behavior

169

. 1983. Effect of wind on avian metabolic rate with particular reference to

Gambel's quail. Physiological Zoology 56:485-492. Abstract: The metabolic rate of
Gambel's quail (Callipepla gambelii) increases linearly with increasing wind speed at
both 10 and 20 C. These results are compared with data from the literature for seven
other avian species. The square root of wind speed, though often used as the
independent variable, does not provide the best description of metabolic rate in wind for
most species and temperatures; however, presentation of all data in a common form
does reveal patterns among and within species. The effect of convective heat loss on
metabolic rate — that is, the slope of metabolism (in watts, W) on the squared root of
wind speed (m/s) — increases as mass increases. This slope also increases within a
given species as ambient temperature (Ta) decreases. These relationships are the
result of relative changes in surface area, thermal conductance, and the temperature
difference driving heat flux. The slope of the regression of metabolism on the square
root of wind speed [b, in W/(m/s)1/7] may be described as b = .0092M.66AT.32, where M
is mass in grams and AT is the difference between the lower critical temperature and Ta
(in °C). This equation predicts the metabolic rate of a bird at any wind speed in

76 _>,

4-

f\ ^ ^

(^

temperatures below thermoneutrality. Key words: Gambel's quail, wind,
temperature, thermoregulation

170

Gore, H. G. 1973. Land-use practices and Rio Grande turkeys in Texas.
Proceedings of the National Wild Turkey Symposium 2:253-262. Abstract:
Populations of Rio Grande turkeys (Meleagris gallopavo intermedia) in Texas are now
closely correlated with areas of least human activity located where mean annual rainfall
is 16 to 20 inches. Turkey range and populations are further restricted by overgrazing,
human occupation and disturbances, and destruction of habitat. In general, turkeys
have declined in direct proportion to the increase in human occupation of the land. Most
lands in Texas are privately owned with no restrictions on use. Both Rio Grande and
eastern turkeys (M. g. silvestris) were abundant in Texas in 1870 when the population
of 818,500 Texans owned 4.5 million head of livestock. Today's 12 million people and
3.5 million head of livestock (including 3.5 million goats and 4.0 million sheep) have
made inroads into turkey habitat that cannot be readily corrected. Vegetative
associations that once produced an abundance of food for turkeys have been subjected
to brush-control programs or have been replaced by various invader species of limited
value to turkeys. The economic importance of livestock points to even greater
decreases in turkey habitat as human populations and demands increase. Removal of
brush and timber is not the only reason that the number of turkeys has decreased,
because properly managed open range is extensively utilized by turkeys for nesting and
feeding. It is destruction of winter roost sites and large-scale removal of mast-producing
species that presently cause remnant flocks to change their ranges-changes that more
often than not result in conditions that make the survival of turkeys impossible.
Transplanting of wild-trapped birds to ancestral ranges is credited with increasing the
number of turkeys within their present range. Widespread precipitation and improved
range conditions following periods of drought have also increased local turkey
populations. The future of the Rio Grande turkey hinges on whether limited habitat can
be maintained or improved under the present system of agricultural economics. Land-
use practices that presently promote increases in the number of turkeys are rotated
grazing systems, decreases in numbers of goats and sheep, planting of small grains in
winter food plots, minimal human disturbance, and supplement feeding during critical
periods of drought. Key words: Rio Grande turkey, eastern turkey, precipitation,
drought, habitat modification, grazing

171

Graham S. A. and G. Hesterberg. 1948. The influence of climate on the ring-
necked pheasant. Journal of Wildlife Management 12(1):9-14. Summary: The author
applied the "bioclimatograph" to explore the failure of pheasant to thrive in southern
North America while they succeeded further north. From the discussion of
meteorological influences it seems entirely possible that the southern distribution of
pheasants in the United States may be due to intense spring insolation of the exposed
eggs during oviposition. However, it must not be inferred that temperature is the only
controlling factor. If the insolation hypothesis proves to be sound, spring and early
summer temperatures on exposed surfaces would mark the limits of possible

Jrf 77

*4>

^ k^ 1*

^SS -"Jar- <£^b> cO±

distribution toward the south. But within the favorable area other limiting factors would
operate to make specific localities either favorable or unfavorable. Key words: ring-
necked pheasant, temperature, insolation, precipitation, distribution

172

Grandjean-Thomsen, A. 1994. The influence of some factors on the display
activity of black grouse (Tetrao tetrix) in Denmark. Dansk Ornitologisk Forenings
Tidsskrifft 88(2):85-90. Abstract: The present work identifies some factors influencing
the display activity of the black grouse on an arena in central Jutland, studied in 1974-
75. The activity varies seasonally, with a peak period from April to the end of May.
Strong winds damp the activity, whereas decreasing temperature apparently has a
stimulating effect. It was not possible to demonstrate any effect of cloud cover, but rain
interrupted the display. The presence of hens on the display ground has a strong
stimulating effect on the displaying cocks. Overflying curlews (Numenius arquata) have
a similar effect, apparently because they are taken for black grouse hens in the twilight
of the morning. Key words: black grouse, wind, temperature, precipitation,
behavior

173

Gratson, M. W. 1988. Spatial patterns, movements, and cover selection by sharp-
tailed grouse. Pages 158-192 in A. T. Bergerud and M. W. Gratson, editors.
Adaptive strategies and population ecology of northern grouse. Wildlife
Management Institute and University of Minnesota Press, Minneapolis, USA.

Notes: Page 182 — Generally, flocks were largest when snow was on the ground but
was less than 18 cm, a depth beyond which birds could no longer obtain ground foods
easily but at which they could snow-burrow. Thus, mean flock size decreased when
there was a reduced availability and distribution of ground foods, and also when there
was increased snow-burrowing opportunities. These changes were statistically
significant for males from conditions of no snow to those with snow less than 18 cm and
for females from conditions of snow less than 18 cm to those greater than this depth.
Page 189 — At snow depths greater than 18 cm, flocks were generally smaller than
when snow was present but less than 18 cm. Food availability and distribution, certainly
the latter, decreased with snow depth, if at all; however, more birds did not join flocks
when snow was deep. Rather, mean flock size decreased. Flock sizes were related to
snow-burrowing opportunities. At snow depths that allowed grouse to snow-burrow,
birds left flocks. When depths were reduced and they had to roost exposed on the
snow, birds joined others, and mean flock sizes increased. These data appear
consistent with the general hypothesis that grouse join flocks when they are most
conspicuous, on top of the snow. When sharp-tailed grouse snow-burrowed, the
benefits of increased vigilance owing to flocking should have been fewer, and as
predicted by the general antipredator models, birds left flocks. Key words: sharp-tailed
grouse, snow, behavior, habitat use, predation

174

Gray, B. T. and H. H. Prince. 1988. Basal metabolism and energetic cost of
thermoregulation in wild turkeys. Journal of Wildlife Management 52(1):133-137.

78 ^

^

>->' fee >f*

Abstract: Metabolism of eastern wild turkeys (Meleagris gallopavo silvestris) was
measured as a function of temperature during the winters of 1983-84, 1984-85, and the
summer of 1984. Basal metabolism (mL oxygen/g/hr) did not differ between sexes
within seasons, but was higher for juveniles during winter and adults in summer than for
adults in winter. Metabolic costs for thermoregulation below the lower critical
temperature (Tk) for females was greater than for males during each season. Extended
periods of deep powder snow combined with low temperatures are considered a major
limiting factor for wild turkey populations. During periods of deep powder snow mobility
is greatly reduced and feeding is restricted to the immediate vicinity of the roost. If no
food is readily available, turkeys must fast. The more rapid rise in energy expenditure
below Tk in females than males during periods of fasting would result in depletion of
female's lipid reserves at a greater rate. Key words: eastern turkey, bioenergetics,
snow, temperature, nutrition, population dynamics, mortality

175

Green, R. E. 1984. The feeding ecology and survival of partridge chicks {Alectoris
rufa and Perdix perdix) on arable farmland in East Anglia. Journal of Applied
Ecology 21:817-830. Summary: (1) Broods of grey and red-legged partridges were
radio-tracked for 20 days after hatching and their diet determined from feces collected
at nocturnal roosts. (2) Grey partridge chicks foraged in cereal fields and red-legged
partridge chicks in cereal, carrot and sugar-beet fields. Chicks of both species fed on
arthropods, weed seeds, leaves, flowers and cereal grain but grey partridge chicks were
more insectivorous than red-legged partridges. Both species preferred to feed at the
edges of fields, where arthropods and weeds are more abundant, and among cereals
they preferred winter wheat fields which had the highest densities of arthropods and
grass seeds. (3) Periods of brood activity and resting were monitored by continuous
recording of radio signal strength. Activity was reduced during periods of low
temperature and rain or dew. (4) Surveys of chick survival rates and food supplies on
different farms showed that the survival of grey partridge chicks increased with
increasing density of prey arthropods. Red-legged partridge chick survival was
influenced by the abundance of arthropod prey and grass seeds. (5) Mean survival
rates for partridge chicks estimated annual for samples of farms in East Anglia showed
that survival increased with June and July temperature for both species. It is uncertain
to what extent weather affects chick survival directly via its effect on chick activity rather
than indirectly via effects on food supplies. Key words: grey partridge, red-legged
partridge, temperature, precipitation, dew, behavior, nutrition, insects

176

Gregg, L. 1984. Population ecology of woodcock in Wisconsin. Technical Bulletin
No. 144, Department of Natural Resources, Madison, USA. Notes: Page 19 —
"Although predators were responsible for the destruction of eggs in many nests, the fate
of some of those nests might have already been sealed. Because the woodcock is a
very early nester, inclement weather conditions, including cold temperatures and
significant snowfalls, are not unusual during the nesting season. Such adverse weather
has been known to result in deserted clutches when eggs are frozen or buried under
snow... although weather conditions caused the loss of many nests which were then

% Hteof*

79

subsequently destroyed by predators, it was seldom possible to document such losses.
During the 1979 nesting season, however, a late snowstorm provided an opportunity to
gain information on the effect of weather on nesting success. A 6-inch snowfall within
the study area on 5 May was judged to have forced hens to abandon those nests
having little or no overhead cover. Of 17 nests under surveillance at the time of the
storm, 9 were found inactive upon inspection 8 May. The eggs were missing from 6
nests, shells from eggs destroyed by predators remained in 2 nests, and abandoned but
intact eggs were found in only 1 nest. There is little doubt that the impact of this storm
was unusually severe because snow depth was sufficient to cover nearly all of the
ground vegetation surrounding the nests, thus making the hen or the eggs very visible
to passing predators. Nevertheless, the rapid disappearance of the eggs from those
nests points up the difficulties involved in discriminating between weather and predation
as causes of nest losses. Page 20 — Fate of individual eggs in completed clutches was
dependent upon environmental factors, such as weather and predators, and also upon
intrinsic factors such as fertility. Page 38 — Year-to-year variations in average brood size
were smaller than those recorded for nesting success but were still large enough to
indicate that chick survival varied between years. Annual variations in weather
conditions were believed to be primarily responsible for observed differences in brood
size, but an examination of weather records provided little evidence of a relationship. If
weather conditions during some years caused a large number of hens to lose their
entire brood, however, such losses would not necessarily be reflected in smaller
average brood sizes during those years since it is impossible to identify the proportion
of hens which had lost their brood. Page 42 — Causes of Mortality, Weather. Although
several records of woodcock mortality resulting from cold waves exist for the wintering
grounds, such losses are more difficult to detect on the breeding grounds. Within our
study areas, inclement weather caused significant losses of nests and occasionally
even young chicks, but was believed to be a much more important mortality factor than
indicated by any carcass counts... we recorded two instances of weather-related
starvation among males in the spring. Males, because of their smaller body size and
associated greater heat loss, are less well equipped than females to withstand cold
weather. Body weights of adult males are at their lowest point during the spring
courtship period, and energy reserves in the form of body fat would not be available to
sustain them if food became unavailable. Thus, any mortality resulting from severe
spring weather would involve primarily male birds and could be a factor in maintaining
the preponderance of females among adult woodcock. Key words: American
woodcock, temperature, snow, productivity, mortality, predation, nutrition,
population dynamics, sex ratio

177

Gregg, M. A. 2006. Greater sage-grouse reproductive ecology: linkages among
habitat resources, maternal nutrition, and chick survival. Dissertation, Oregon
State University, Corvallis, USA. Notes: Page 40 - "Reduced availability and
consumption of forbs during 2002 may have been related to late winter snow that
covered my study area during the early collection period, which perhaps delayed forb
growth. In contrast, my study area was snow free during the early collection period in
2003, forb phenology was advanced, and availability and consumption of forbs was
greater than I observed in 2002. Variation in availability and consumption of forbs

80 ^

4>

>->" fc* **

between years was also related to ovary weight of my hens. Follicle development was
advanced for hens collected during the early period in 2003 compared with 2002 and
may have been related to greater nutrition because of increased consumption of forbs
during 2003." Key words: sage-grouse, snow, vegetation, nutrition, productivity

178

Gross, A. O. 1930. Progress report of the Wisconsin prairie chicken investigation.
Report of the Wisconsin Conservation Commission, Madison, USA. Abstract:
Important factors which control the numbers of grouse in Wisconsin are periodic cycles,
weather conditions, fires, encroachment of agriculture upon nesting and feeding
grounds, hunting, predators, diseases, and parasites. Prior to 1887 there were no laws
limiting hunting of prairie grouse in Wisconsin. The open season has gradually
decreased from 3 months in 1887 to 5 days in certain counties in recent years. The bag
limit has been reduced to 5 birds. Estimates place the numbers of prairie chicken at
54,000, and the sharp-tailed grouse at 55,000, or very roughly one prairie chicken and
one sharp-tailed grouse to each square mile of land in Wisconsin. The prairie chickens
undergo distinct migrations in the northern portion of their range. There were 6 internal
and 6 external parasites of varying importance found in Wisconsin grouse. The dreaded
poultry disease "blackhead" was found in 3 of the prairie chickens examined. The
organic food of the birds examined was made up of 84 kinds of vegetable matter,
constituting 72 percent, and 82 kinds of animal matter, making up 28 percent of the
food. Gravel constituted 6 percent of the combined organic and inorganic contents.
Experimental winter feeding stations established by planting plots of ground of one to
two acres in buckwheat and other grains to be used by the birds during times when the
natural food is covered with ice and snow proved an excellent conservation measure.
Forty-six nests were used in the life history study. The average number of eggs per set
was 1 1 .5. The average measurements of 1 00 eggs was 44.86 by 33.59 mm; weight,
21.86 grams. The incubation period was 23 days. Destruction of nests by predators is
one of the serious problems of conservation of the prairie chicken. Hybrids between
prairie chickens and sharp-tailed grouse are common. Erythrim and albinism occur.
Courtship and behavior of the prairie chicken is identical with that of the heath hen,
confirming their subspecific relationship. Key words: prairie chicken, sharp-tailed
grouse, snow, ice, mortality

179

Gruys, R. C. 1993. Autumn and winter movements and sexual segregation of
willow ptarmigan. Arctic 48(3):228-229. Abstract: Willow ptarmigan {Lagopus lagopus
alexandrae) in northern British Columbia leave their breeding areas during autumn and
winter. The movements differ between males and females. In this study I examine the
causes and extent of these differences. Ptarmigan did not leave their breeding grounds
immediately after the breeding season, but remained on or near their territories until
December. After chicks fledged, part of the population moved uphill from their
territories. Coincident with moult into winter plumage, ptarmigan moved farther from
their territories. Both movements were probably to areas with better protection against
predators. After moulting, all tagged males and half of the tagged females returned to
their territories, and males resumed territorial display. Ptarmigan remained on their

H (& **

territories until increasing snow cover depleted cover, forcing them to leave. Males left
the breeding grounds later than females and returned earlier in spring. In winter females
moved farther than males, supporting the reproductive strategy hypothesis, but
segregation was not complete. Sexual segregation may not be related to migration
alone, but could occur at any time ptarmigan are in flocks. Key words: willow
ptarmigan, snow, habitat use, movement

180

Gullion, G. W. 1970. Factors influencing ruffed grouse populations. Transactions
of the North American Wildlife and Natural Resources Conference 35:93-105.

Notes: Page 94 — "In northern Minnesota I believe we must consider ruffed grouse to be
chionophiles, or "snow lovers" and that their overwinter welfare depends in large degree
upon snow conditions... 20 inches of snow with a hard crust can be as deleterious as 6
inches of soft snow with no crust, when the temperatures remain in subzero (°F) ranges
for prolonged periods." Page 95 — "During the 10 years of adequate study only in 1963-
64 was the overwinter loss substantially greater than a mean of about 55 percent. This
we attribute to increased predation associated with poor snow conditions... Poor snow
conditions in 1967-68 resulted in a poor production season in 1968 and 1969 proved
only moderately successful." Page 96 — "We believe that when environmental
temperatures drop much below 20°F they are below the thermoneutral range of this
grouse and that the birds must increase their metabolic rates to maintain body
temperatures... snow depth and quality is a most important factor affecting year-to-year
changes in northern Minnesota's ruffed grouse populations — but it is not the entire
story." Page 97 — "We have found at Cloquet that the color of a ruffed grouse's tail
indicates the bird's probable longevity. Over the past 13 years male grouse with red
tails have lived an average of only 69 percent as long as the birds with gray tails (1 1 .4
months vs 16.6 months — significant at 99.5% level; 275 df). Furthermore, this
differential survival appears to be related to environmental stress. It is during winters of
unfavorable snow conditions that the red birds show the greatest losses. In good snow
years, such as 1964-65, the red birds survived as well as the gray birds." Key words:
ruffed grouse, snow, temperature, thermoregulation, color-phase, wind, fire,
habitat use, habitat alteration

181

Gunderson, P. 1990. Nesting and brood rearing ecology of sharp-tailed grouse on
the Charles M. Russell National Wildlife Refuge, Montana. M.S. Thesis, Montana
State University, Bozeman, USA. Abstract: Spring and summer habitat use was
studied for plains sharp-tailed grouse (Tympanuchus phasianellus jamesi) during 1988
and 1989. The study area spanned 104 km2 in south Valley County, Montana. Thirty-
four sharptails with transmitters were followed. Males used 5 display grounds, slightly
raised from surrounding topography and interspersed with big sagebrush {Artemisia
tridenata), from the start of field seasons in April, until early June. Twenty-seven nests
were located over 2 years and used to characterize nesting habitat vegetatively and
topographically. Most nests (63%) were located in junipers (Juniperus sp.). Canopy
height over nests averaged 47 cm, and screening cover height was near 18 cm.
Hatching occurred in early to mid-June. Success of nesting hens was high (65%), but

82 y

H ter ^

■>"^

c£^

predators destroyed nearly 30%. Four hens nested in both years, and another renested
in 1989. Fidelity to nesting areas was shown by 2 hens. Due to weather and predation,
no chicks were alive after 2 weeks in 1988. Adults centered summer home ranges
around Juniper, Shale, and Lakeshore cover types. In 1989, 5 hens with broods were
followed until late August. Vegetative measurements were recorded at 105 locations
sites and at areas within Juniper, Shale, and Lakeshore cover types, along with Poa
sandbergii/Sarcobatus vermmiculatus. Cover type use by broods differed between
early, middle, and late summer in 1989. Brood habitat contained significantly more
grass than at random points. Brood home ranges (433 ha) were larger than for adults
without broods (274 ha). Two hens were followed during both summers. Hen 1788 used
the same summer area in both years. Only 14 chicks were raised in 1989. Low overall
production was caused by weather and predation. Lack of production in 1988 was
evidenced by census and trapping data collected in 1989; numbers of displaying males
was reduced 33% from 1988 to 1989, and most birds (88%) captured in 1989 were
adults. Page 60 — Later hatching in 1989 was possibly caused by a 5-day rainstorm in
late April which may have delayed female attendance on dancing grounds. Rain and
snow have inhibited dancing activity in Minnesota and North Dakota. Weather
conditions during the breeding period may modify the peak of hatching in any given
year. Egg hatchability in 1988 (81%) was lower than 1989 (100%). This was probably
due to the extremely high temperatures experienced during the 1988 hatching period.
Cooler temperatures prevailed in 1989 and hatchability was comparable to other
studies. Page 63 — Nests in 1989 were more screened from view by vegetation than in
1988. Height-density pole readings were also significantly higher in 1989. This increase
in vegetation was associated with a 7.1 cm increase in precipitation totals during April
and May in 1989. In 1989, both vegetation height and effective height increased
significantly throughout incubation. The decrease in vegetation height and effective
height, in 1988, was most likely caused by lack of grass and forb growth nests and
natural reduction of residual vegetation. Lack of new growth may have contributed to
the death of an entire clutch. This nest, in residual grasses, became increasingly
exposed to the sun as incubation progressed. Page 65 — In 1988, high daily
temperatures in June were over 32 C (90 F) and 38 C (100 F), 22 and 6 times,
respectively. Ground temperatures on the shale soils were probably considerably
higher. Two chicks were found dead within 10 m of nests and 10 chicks died while
exiting their eggs. In 1989, temperatures during the early brood period were mild and no
loss of entire broods was recorded. Weather conditions in early June are important in
determining brood success. Although hot weather was a major cause of early chick
mortality in this study, cold, wet weather is reported more often as having detrimental
effects on young gallinaceous birds. Page 66 — It appears that young in 1988 were
initially affected by excessive heat prior to normal predation. Key words: sharp-tailed
grouse, productivity, temperature, precipitation, vegetation, predation, habitat
use, behavior, movement

182

. 1989. 1988 sharp-tailed grouse production in the Missouri River Breaks.

Proceedings Annual Meeting of the Montana Chapter of The Wildlife Society 27:6-

7. Notes: Gunderson found that sharp-tailed grouse production was poor. He states:
"Despite high nest success, 50 percent of the nests in my study had eggs that didn't

i f>'<c^

83

£^ c^

hatch. Eight partially pipped eggs were also found in three separate successful nests.
The chicks either became desiccated or overheated before they could escape the shell.
The high mortality at hatching appeared unusual since I have not located published
reports of similar losses. On two separate occasions, despite successfully exiting the
shell, a newly-hatched chick was found dead within ten feet of the nest." Page 7 - "The
peak of hatch occurred in the first week and a half of June. The average daily high for
the first ten days of June was 96.4°F, including 8 days over 90° and 4 over 100°." Key
words: sharp-tailed grouse, temperature, drought, mortality

183

Guthery, F. S. 1985. Bobwhite and turkey management during drought. Pages 31-
36 in R. D. Brown, editor. Livestock and wildlife management during drought,
Caesar Kleberg Wildlife Research Institute, Kingsville, USA. Summary: A "mini-
drought" struck south Texas in 1984. Many areas received less than ten inches of rain
from January through September. In fall 1984, we measured bobwhite densities on 16
sites scattered from Victoria to Zapata. The lowest density was 1 bird/100 acres on an
overgrazed pasture in Zapata County. The highest was 190 birds/100 acres on a 1 ,600-
acre pasture in Hidalgo County. The results show that good bobwhite densities can be
carried through short droughts if managers are willing to plan and make some
sacrifices. Late fall and late spring rains are more important to bobwhites than rains at
other times. Research by the Texas Parks and Wildlife Department shows that chick
production and fall rains go hand in hand. The same is true of poult production in
turkeys. Drought or no, you have to address habitat structure first. Key words:
bobwhite quail, wild turkey, drought, precipitation, productivity, habitat quality

184

, J. J. Lusk, D. R. Synatzske, J. Gallagher, S. J. DeMaso, R. R. George, and

M. J. Peterson. 2002. Weather and age ratios of northern bobwhites in south
Texas. Proceedings of the National Quail Symposium 5:99-105. Abstract:
Understanding the effects of weather on quail reproduction in semiarid environments
requires simultaneous consideration of temperature and precipitation data. Therefore,
we used neural modeling to assess the interactive effects of summer (Jan-Aug)
temperatures (monthly means of daily maxima) and seasonal precipitation (totals) on
age ratios (juvenile/adult) of northern bobwhites (Colinus virginianus) in south Texas
based on data collected during 1940-97 (n=35,23 years missing). Age ratios increased
with June temperature. Ratios were insensitive to mean maximum daily temperature in
July up to 36°C, when they began to decline rapidly. Ratios were insensitive to August
temperatures. Ratios increased in an asymptotic manner with fall (Sep-Nov), spring
(Mar-May), and summer precipitation, and were least sensitive to fall precipitation and
most sensitive to spring precipitation. Based on our analysis, temperature and
precipitation influenced bobwhite production in a complex, nonlinear manner that
seemed to contain thresholds and asymptotes. Low temperatures can ameliorate the
negative effects of drought, and high temperatures can suppress the positive effects of
precipitation. The apparent asymptotic effect of precipitation, given temperature,
illustrates that assumed linearity between precipitation and production may lead to

84 ^

% rafter ><*

CC}

errors of interpretation and expectation for production in a particular year. Key words:
bobwhite quail, temperature, precipitation, productivity

185

, A. R. Rybak, S. D. Fuhlendorf, T. L. Hiller, S. G. Smith, W. H. Puckett Jr.,

and R. A. Baker. 2005. Aspects of the thermal ecology of bobwhites in north
Texas. Wildlife Monographs No. 159, The Wildlife Society, Washington, D.C., USA.

Abstract: We studied the thermal ecology of northern bobwhites (Colinus virginianus) to
better understand the role of temperature in the field behavior of these birds. We
obtained descriptive data on thermal aspects of the landscape; bobwhite selection for
Normalized Difference Vegetation Index (NDVI; vegetation biomass) classes and cover
associations relative to their thermal properties; and thermal conditions at nests, mid-
day coverts, and roosts. We collected data on a 796-ha area in the Texas Rolling Plains
during May 2000-July 2003 using satellite imagery, black-bulb temperature probes,
mortality- and temperature-sensing radiotransmitters, and continuous-recording video
cameras for nest observations. Linear models of black-bulb temperature (Tbb) as a
function of air temperature (Ta) at a base weather station explained 42-70% of the
variation in Tbb in 10 NDVI classes during daylight and 78% during night in summer (all
NDVI classes; July 2001). During February 202, Ta explained 38-92% of the variation
during day and 89% of the variation at night. The linear models provided a means of
qualitatively assessing thermal space on the landscape as Ta changed and of predicting
Tbb in NDVI classes. At Ta = 42°C, 100% of the 796-ha landscape under study had
predicted Jbb > 39°C, the approximate threshold leading to hyperthermia in bobwhites.
Based on 9,287 radiolocations of 217 bobwhites, bobwhites selected for all NDVI
classes in mixed-shrub cover (sandplum, [Prunus angustifolia]; fragrant sumac, [Rhus
aromatica]) on an annual and seasonal basis. If Ta was <35°C, the approximate upper
critical temperature, the operative temperature (Te) experienced by bobwhites exceeded
Ta in 699 of 818 simultaneous readings (n = 24 bobwhites with thermal transmitters)
and the difference between Te and Ta increased as air temperature declined. Data from
video cameras indicated thermal stress (i.e., gular flutter) in 25 of 26 incubating
bobwhites. Gular flutter began at an average Ta of 30.4±0.2°C SE (n = 158) and total
bouts of gular flutter averaged 87 minutes/bird/day after 16 June. Data from thermal
radiotransmitters indicated 91.3±6.1% of incubating adults were in thermal stress at Ta
> 35°C. Temperature of nest contents averaged about 30°C and incubating bobwhites
appeared to protect nest contents more rigorously from hyperthermia than from
hypothermia. Mid-day covert selection (NDVI 6, mixed-shrub cover; n = 58) during
summer reduced bobwhite exposure to Tbb> 30°C by an average of 1,6000.7 heating-
degree minutes in comparison with random points (n = 58) during 1200-1600 hours.
The roosting disc appeared to confer energetic advantages at Ta < 16.2°C and the
advantage increased as Ta declined. Our results were consistent with the heat
hypyothesis on reproduction variability because we observed thermal stress at low Ta
(> 22.1 °C) in incubating birds and we projected lethal Tbb (53.2°C) at high Ta(45°C) in
NDVI classes used by nesting bobwhites. Our results reaffirm that temperatures on the
landscape are a legitimate concern for bobwhite managers. The goal of habitat
management vis-a-vis temperature is to insure well-distributed mid-day coverts for
thermal refugia in summer and ground cover of sufficient height and density to prevent
heat excess in the near-ground environment. Key words: bobwhite quail,

^ *C« **

-X,

S~*^,' »N"' s^*\, y~^

% w* 4& ^

temperature, habitat use, habitat management, biothermal regulation, behavior,
vegetation

186

, M. J. Peterson, and R. R. George. 2000. Viability of northern bobwhite

populations. Journal of Wildlife Management 64(3):646-662. Abstract: Quail
populations are declining in North America. Hence, we modeled the viability of northern
bobwhite (Colinus virginianus) populations subject to weather catastrophes and harvest
to help focus management and research efforts to reverse the decline. We examined
northern-latitude (winter weather catastrophes) and southern-latitude (summer weather
catastrophes) populations separately, because winter survival is lower and density-
dependent production is stronger in northern- than in southern-latitude populations.
Under a criterion of quasiextinction at < 14 birds (1 covey), the demographic capacity
required for at least a 95% probability of persistence for 100 years was about 100 birds
with summer weather catastrophes, about 500 birds with winter weather catastrophes,
and about 800 birds with both winter and summer weather catastrophes. Given
assumptions underlying the model, populations subject to summer weather
catastrophes were sustainable under a < 30% harvest rate if demographic capacity in
autumn was about 700 birds. Populations subject to winter weather catastrophes could
persist at < 40% harvest rates and a demographic capacity of about 400 birds. Northern
populations were more vulnerable to extinction in the absence of harvest, whereas
southern populations were more vulnerable to extinction in the presence of harvest.
Key words: bobwhite quail, model, weather catastrophes

187

, N. D. Forrester, K. R. Nolte, W. E. Cohen, and W. P. Kuvlesky Jr. 2000.

Potential effects of global warming on quail populations. Proceedings of the
National Quail Symposium 4:198-204. Abstract: Populations of scaled quail
(Callipepla squamata) and northern bobwhites (Colinus virginianus) have declined in
North America coincident with global warming. We speculate on a cause-effect relation
between global warming and quail declines. Quail are sensitive to operative
temperatures > 38.7°C, which commonly occur under natural conditions in southern
latitudes. Based on empirical results, the laying season for quail may be reduced by as
much as 60 days because of high temperatures. We provide mechanistic models that
show how reduction in length of the laying season suppresses per-capita annual
production. Global warming could be associated with declining quail populations
through suppression of reproduction; it also could exacerbate the effects of habitat loss
and fragmentation. These possibilities should be explored in field and laboratory
research. Key words: scaled quail, bobwhite quail, temperature, reproduction,
productivity, habitat loss/fragmentation, model

188

Hamerstrom, F. N. Jr. 1939. A study of Wisconsin prairie chicken and sharp-tailed
grouse. Wilson Bulletin 51(2):105-120. Notes: The author notes the effects of weather
on breeding behavior. The dancing of the Columbian sharp-tailed grouse is correlated
with light conditions. Hamerstrom, based on his study, believed the prairie chicken

86 ^

4-

H <c> **

#*

mating is regulated by light and temperature. He found that (a) late winter displays
occurred on days warmer than usual, (b) early spring displays occurred for an hour or
two after sunrise on clear bright mornings with temperatures a few degrees below
freezing, but not on raw cloudy mornings which were five or six degrees warmer, (c)
during the early part of the main booming season booming occurred all day on cloudy
days; later, in warmer weather, cloudy days did not produce the day-long response, (d)
during the height of the season booming occurred from the first faint trace of light until
an hour or two after sun-up, again for about two hours before dark; there was less
activity on unusually hot days than on moderate ones, (e) on those hot days and toward
the end of the season generally, there was more activity in the cooler early morning
than in the evening, and morning booming stopped earlier than usual, and (f) at the end
of the season, although cocks continued to come to their regular stations morning and
evening, booming was heard only on occasional clear frosty mornings and stopped as
soon as the air grew warmer shortly after sunrise. Light rains did not deter booming
cocks; heavy rains silenced them completely. Key words: sharp-tailed grouse,
prairie chicken, temperature, precipitation, clouds, behavior, frost

189

, O. E. Mattson, and F. Hamerstrom. 1957. A guide to prairie chicken

management. Technical Wildlife Bulletin Number 15, Wisconsin Conservation
Department, Madison, USA. Notes: Page 79 - Weather. The only real defense against
extremes of weather is good habitat.... the prairie chicken can be saved only by
guaranteeing to it a place to live. Above all else, this means habitat management. Key
words: prairie chicken, weather extremes, habitat management

190

Hammond, M. C. 1941. Fall and winter mortality among Hungarian partridges in
Bottineau and McHenry counties, North Dakota. Journal of Wildlife Management
5(4):375-382. Notes: "..there [is a] decided tendency for the partridges to include a
graveled, or at least a graded, road within their winter range, especially during periods
of moderately deep snow." "Even during periods of deep snows and extremely low
temperatures, all partridges examined were in very good physical condition, while
pheasants in the same localities were often found to be thin and sometimes, apparently,
were starving." "Winter conditions are quite variable in the portion of North Dakota
studied, and years of relatively mild weather as well as those of extreme cold occur.
Even though the snow was 20 inches in depth at times (in areas protected from drifting),
winds kept it piled in drifts, and the stubble fields had a covering that averaged not more
than 5 or 6 inches. Field observations disclosed that partridges were able to dig through
these snow depths and find waste grain." "Yeatter found evidence that wet weather
combined with certain soil conditions resulted in mudballing of young chicks." Key
words: Hungarian partridge, snow, temperature, habitat use, nutrition, soil

191

Hannon, S. J. and K. Martin. 2006. Ecology of juvenile grouse during the
transition to adulthood. Journal of Zoology 269:422-433. Notes: Page 423— The
primary causes of early juvenile mortality were cold wet weather, predation, lack of food

>•> ^ ^

\ ^"

^<S\~ cL^J^ c _b

and low quality of the chick or hen. When cold wet weather occurs shortly after hatch,
juvenile mortality can be high. Chicks are not able to thermoregulate fully until they are
8-10 days old, and during inclement weather must reduce foraging time to brood. In
poor weather, the lower availability of insects may reduce foraging success, resulting in
poor chick growth. Chicks may also be more susceptible to predation after cold wet
weather. Before achieving adult body mass, juveniles have higher energy demands
because of growth and moulting, poorer insulation and higher surface/volume ratio than
adults. In species where males are much larger than females, male chicks have higher
mortality rates than females in years with poor weather and/or low food production. This
has been attributed to the higher energy requirements and growth rates of males. For
prairie grouse, drought conditions may also reduce juvenile survival. However, several
studies found little or no link between chick mortality and weather. Page 429 — There
appear to be two main 'bottlenecks' for juvenile survival. The first is over the first 2
weeks of life when chicks are dependent on the hen for thermoregulation, habitat
selection and protection from predators. For species where males are much larger than
females, males suffer proportionately higher mortality than females in years with poor
weather and/or food. Key words: grouse, precipitation, temperature, habitat use,
insects, nutrition, mortality, thermoregulation, drought, predation

192

Haroldson, B. S. and R. O. Kimmel. 1990. Winter bait-trapping of gray partridge as
influenced by snow cover. Pages 159-164 in K. E. Church, R. E. Warner, and S. J.
Brady, editors. Proceedings Perdix V: Gray partridge and ring-necked pheasant
workshop. Minnesota Department of Natural Resources, Mankato, USA. Abstract:
Gray partridge {Perdix perdix) were trapped in south-central Minnesota during the
winters of 1984 through 1988 using walk-in bait traps. Trapping success ranged from
0.00 birds per trap-day in 1985 and 1987 to 0.52 birds per trap-day in 1988. Success
increased during periods with complete snow cover and decreased with patchy snow
conditions. Snow cover should be present for several weeks before tapping begins. Key
words: gray partridge, snow, technique

193

Haroldson, K. J. 1995. Energy requirements for winter survival of wild turkeys.
Proceedings of the National Wild Turkey Symposium 7:9-14. Abstract: As wildlife
managers expand the range of wild turkeys (Meleagris gallopavo) beyond ancestral
northern limits, information on the tolerance of wild turkeys for severe winter weather
becomes increasingly important. I used predictive models based on time-energy
budgets to estimate winter food requirements of wild turkeys. An average 4.23 g wild
turkey would require 1 1 .3 kg of a mixed diet during a 120-day winter with a mean
temperature >1 1°C. Winter food requirements would increase by 2.4 kg/bird for every
10°C drop in mean winter temperature. Because wild turkeys in northern climates often
supplement natural foods with corn during winter, I estimated size of corn food plots
needed to sustain wild turkeys based on average winter temperature. Key words: wild
turkey, temperature, model, nutrition

88 j,

f \ <c> ^

194

, R. O. Kimmel, M. R. Riggs, and A. H. Brener. 2006. Association of ring-
necked pheasant, gray partridge and meadowlark abundance to Conservation
Reserve Program grasslands. Journal of Wildlife Management 70(5):1276-1284.

Notes: Page 1282 — In Minnesota, statewide partridge indices peaked during a
widespread drought in 1987-1990, declined when the drought ended in 1991, and fell to
a low during the extremely wet summer of 1993... We believe that spring and summer
precipitation trends heavily influenced the strong year effect we observed. Key words:
gray partridge, precipitation, drought, population dynamics

195

, M. L. Svihel, R. O. Kimmel, and M. R. Riggs. 1998. Effects of winter

temperature on wild turkey metabolism. Journal of Wildlife Management
62(1):299-305. Abstract: We used indirect calohmetry to measure the effects of air
temperature (Ta), age class, and body mass on metabolic rates of 9 adult and 7 juvenile
female eastern wild turkeys (Meleagris gallopaco silvestris) during winter. Previous
studies produced disparate results on this important aspect of winter ecology of wild
turkeys. Standard metabolic rates (SMRs) of adult and juvenile hens were not different
(P = 0.122) and averaged 28.69 ml_ 02-min"1.bird"1. Wild turkey metabolism increased
with decreasing Ta (P < 0.001 ) below the lower critical temperature (T!c) of 10.9°C.
Metabolic rates were not related to body mass (P = 0.571), and age-specific metabolic
rates were not distinguishable (P = 0.998). We estimated that a flock of 20 hens would
need to find 400 g/day of additional food to meet thermoregulatory demands for each
10°C drop in Ta below 10.9°C. Key words: turkey, temperature, thermoregulation,
nutrition

196

Harper, H. T., B. H. Harry, and W. D. Bailey. 1958. The chukar partridge in
California. California Fish and Game 44(1):5-50. Notes: Page 49 — "The so-called
'wet' years in the desert and semi-desert areas appear to be more favorable to chukar
reproduction than any other factor, especially when precipitation occurs during the late
winter and spring months.... During the hot, dry summer months chukars are seldom
observed more than one mile from water. After the first cool days and the first rains in
early fall, chukars disperse from their summer haunts near water and forage into range
not frequented during the summer." Key words: chukar partridge, precipitation,
temperature, behavior, habitat use

197

Hawkins, A. D. 1937. Winter feeding at Faville Grove 1935-1937. Journal of Wildlife
Management 1 :62-69. Notes: "The summer of 1 935 was ideal for wildlife. Food of all
sorts was plentiful and cover was lush. Nesting was unusually successful and game
birds were abundant, but winter brought the deepest snow and most intense cold of the
century. As a result the plentiful food supply was buried under several feet of snow, and
the birds died. In the summer of 1936 a record drought severely curtailed both food and
cover. The diminished crop of ragweed and acorns suffered a further loss of 10 per cent

> \ ^ ^

^ ±2%, i . L- 89

,>"x

c^>

from insect attack. Nesting results were poor due to the drought. Winter brought no
snow and little severe cold, but the fields were sealed by a blanket of ice on January 6
and remained so, without a break, for two months. Few birds died on fed areas like
Faville Grove; but on some unfed areas quail and Hungarian partridges perished in
large numbers." Key words: quail, Hungarian partridge, snow, precipitation, ice,
temperature, nutrition, mortality

198

Hays, D., M. Tirhi, and D. Stinson. 1998. Washington state status report for the
sage-grouse. Washington Department of Fish and Wildlife, Olympia. Notes: The
authors provide a synthesis of weather impacts on sage-grouse and its impacts on
populations. They state in part "Habitat that provides good shrub and grass cover for
nesting and wintering allows grouse to increase despite predation"; they also note the
negative impacts of overgrazing and fire on sage lands. Key words: sage-grouse,
weather, population dynamics

199

Healy, W. H. and E. S. Nenno. 1985. Effect of weather on wild turkey poult
survival. Proceedings of the National Wild Turkey Symposium 5:91-101. Abstract:
Weather conditions are partly responsible for annual variations in poult production, but
the specific effects of weather on the survival of eastern wild turkey poults (Meleagris
gallopavo silvestris) are not well documented. Understanding the relationships between
weather and poult survival might permit predicting fall populations from the preceding
spring weather. Weather conditions and poult survival were monitored daily during 2
breeding seasons. The impact of weather on poult mortality included both age-specific
and random effects. Rain and low temperatures (3.8 cm, 7-8° C) over an 18-hour period
produced mortality in 12- and 15-day-old broods, but not in 4- and 6-day-old broods.
One hen abandoned her nest during this storm. The disappearance of 8 of 11 poults in
another brood coincided with a heavy thunderstorm that occurred about 1 hour after the
birds left the nest. Predation and accidents were additional causes of poult mortality.
Systematic brood counts provide the best means for predicting fall populations because
spring weather accounts for only part of the annual variation in productivity. Key words:
eastern wild turkey, precipitation, thunderstorm, temperature, mortality, chick
survival

200

Heath, B., R. Straw, S. Anderson, and J. Lawson. 1996. Proceedings of the sage-
grouse workshop, Pinedale, Wyoming, USA. Notes: "All large-scale sagebrush
control projects, regardless of type, adversely affect sage-grouse populations. The
combination of sagebrush eradication and drought is devastating to sage-grouse
populations. During drought, ungulate numbers need to be reduced to diminish the
impact to sagebrush rangelands." Page 5 - "Impacts such as drought, fire, and heavy
livestock grazing greatly reduce the availability of forbs and insects that are required by
sage-grouse chicks. Connelly stated that drought impacted early brood rearing habitat
by reducing important food forbs, furthermore, when drought conditions persist certain
plant species are not produced at all. Connelly's research in Idaho indicated that fire in

90

4>

K kr« **

?©£ 4^ > ^

the Artemisia tridentata wyomingensis type degraded early brood rearing habitat by
reducing sagebrush cover and insect abundance (especially ants). When the affects of
fire are coupled with drought, it dramatically lowered chick survival. It was universally
agreed that sage-grouse use sagebrush exclusively for food during the winter. Because
of the variability in winter snow depths, managers need to maintain different heights and
coverages of sagebrush to ensure sage-grouse use. The use of a particular area is
dependent on the severity of the winter. Typically, sage-grouse will move to "critical
winter habitat" once every 8-1 0 years and these areas are of extreme importance.
These critical winter areas provide sage-grouse with food and cover when most of the
surrounding areas are snow covered. Page 6 - "Mike Schroeder indicated that
windswept, sagebrush ridges often provide winter habitat for sage-grouse because they
provide sagebrush for food and cover even during the harshest winters. Additionally,
they provide snow drifts that sage-grouse can roost in to escape extreme cold." Page 8
- "Connelly stated that since 1986, with the exception of 1989 and 1993, Idaho has
been experiencing a drought. During this time some sage-grouse populations have
declined 60-70%. Drought, in association with other factors, was thought to contribute to
these population declines. During prolonged periods of drought, herbaceous cover is
not produced or is produced at lower levels. The problem is compounded when grazing
pressure is not adjusted. The reduction in herbaceous cover results in lower nesting
success and lost clutches in the 2 week period after hatch. It appears that chicks are
dying from starvation as a result of the lack of herbaceous forage and/or are being
predated when they are forced to move long distances through sparse cover to obtain
food." Key words: sage-grouse, snow, precipitation, drought, nutrition, mortality,
range modification

201

Heffelfinger, J. R. and R. J. Olding. 2000. Montezuma quail management in
Arizona. Proceedings of the National Quail Symposium 4:183-190. Abstract: The
Montezuma quail (Cyrtonyx montezumae mearnsi) has substantially different habitat
requirements than other quails found in the U.S. They inhabit evergreen oak woodlands
of mountain ranges in the southwest and feed primarily on underground bulbs and
tubers. Populations respond to summer precipitation because the vegetation which
provides food and cover for Montezuma quail flourishes after the summer rains.
Moderate to heavy grazing increases availability of Montezuma quail food plants, but
resultant lack of cover precludes use of such sites. Montezuma quail avoid areas with
greater than 50% forage utilization by ungulates. As with other Arizona quail species,
hunting has been shown to have limited or no impact on the population level during the
following years. Birds may be depleted in localized areas temporarily, but available
habitat is re-occupied when pre-nesting dispersal occurs. Annual pre- and post-hunt
flush counts were conducted 1988-1996 by the Arizona Game & Fish Department,
United States Forest Service, volunteers, and local quail hunters. Average covey size
decreased during the hunting season, but the magnitude of the decrease was similar in
unhunted populations. Montezuma quail populations fluctuate in response to habitat
and weather conditions. A state-wide hunter questionnaire program estimated total
harvest trends for Arizona. In addition, wing collection barrels had been placed in
heavily hunted areas from 1981 to 1996 to obtain hunter-effort information and sex/age
characteristics of the harvest. Data from these wings indicate average percentage of

>->' tc« f*

\ *»"

^ -<&- <£fr ^

juveniles in the harvest was higher for Montezuma quail (mean - 74.4%, range = 55.9-
84.9%) than other Arizona quail species such as Gambel's (mean = 65.6%, range = 23-
77%). Hunters harvested an average of 2.2 Montezuma quail per day. In 3,107 hunter-
days during this period, only 13 (0.4%) resulted in a limit of birds. Three of these limits
occurred in 1996 when the bag limit was reduced from 15 to 8 Montezuma quail. Key
words: Montezuma quail, precipitation, vegetation, cover, nutrition, hunting

202

, F. S. Guthery, R. J. Olding, C. L. Cochran Jr., and C. M. McMullen. 1999.

Influence of precipitation timing and summer temperatures on reproduction of
Gambel's quail. Journal of Wildlife Management 63(1): 154-1 61. Abstract: We

analyzed the influence of rainfall and temperature patterns on Gambel's quail
(Callipepla gambelii) to better understand variability in call counts and reproduction,
based on data collected in Arizona during 1978-96, midwinter (Dec-Jan) precipitation
invoked a stronger calling response than early-winter (Oct-Nov) or late-winter (Feb-Mar)
rainfall. Reproductive failure (< 1 juv/ad) was associated with low rainfall in October-
March and high mean daily temperatures during June-July. Moderate production (1-2
juv/ad) occurred under low rainfall in winter, if June-July temperatures were cool. For
any rainfall pattern, higher temperatures in July were associated with lower age ratios.
Key words: Gambel's quail, temperature, precipitation, census, productivity,
population dynamics, age ratio

203

Hejl, S. J., J. Verner, and R. P. Bala. 1988. Weather and bird populations in true fir
forests of the Sierra Nevada, California. Condor 90(3):561-574. Abstract: We
monitored bird communities of true fir forests at 51 study sites in the western Sierra
Nevada during the breeding seasons of 1983, 1984, and 1985. The summer of 1983
followed the El Nino winter of 1982-1983, with the greatest snowfall on record. The
summers of 1984 and 1985 followed winters with moderate snowfalls. Bird species
richness (BSR) and average total count (ATC) increased linearly from 1983 to 1985.
The relative abundances of 12 common species increased from 1983 to 1985;
abundances of two common species did not differ among years. Twenty-eight
uncommon species increased and four decreased in numbers from 1983 to 1985. Bird
response patterns differed between lower-elevation white fir (Abies concolor) forests
and upper-elevation red fir (A magnifica) forests. Bird numbers were similar in both
habitats in 1983 but greater in white fir than in red fir in 1984 and 1985. Although
abundances of the common species increased in both habitats in both years, those of
uncommon species did not increase substantially in red fir until 1985. We suggest that
bird numbers were depressed in 1983, but not atypically so for true fir forests. Numbers
of permanent residents are often limited by frequent but unpredictable winters with
excessive snowfall (Beedy 1982, Granholm 1982, Raphael and White 1984). Numbers
of migrants, as in the case of this study, are sometimes affected similarly. White fir and
lower-elevation forests may harbor "source" populations for red fir "sink" populations
during periods of resource stress Key words: mountain quail, blue grouse, snow, El
Nino, habitat use

92 ^

)*( kr<* **

<£^

204

Henderson, C. W. 1971. Comparative temperature and moisture responses in
Gambel and scaled quail. Condor 73:430-436. Summary: Various responses to high
temperature and low moisture were studied in the laboratory in Gambel quail (Lophortyx
gambelii) and scaled quail (Callipepla squamata). The minimum daily water
requirements of the two species were found not to differ significantly at 28-29°C.
Evaporative water loss rates were similar at 25, 30, and 35° C. At ambient temperatures
of 40 and 45° C, Gambel quail lost significantly higher percentages of water through
evaporation than did scaled quail. The response of body temperature to increasing
ambient temperatures was, in general, the same for both species, and indicated that the
thermoneutral zone was probably between 25 and 35° C. Both species were able to
tolerate an ambient temperature of 40° C without ill effects, but at 45° C Gambel quail
survived better than scaled quail. Gular fluttering was exhibited by Gambel quail when
body temperature exceeded 43° C, but was not observed in scaled quail at any
experimental temperature. Oxygen consumption values were similar at 30° C, but at 40°
C the values were significantly higher for Gambel quail. Both species were able to
dissipate all metabolic heat while experiencing hyperthermia at 40° C. These results
indicate that Gambel quail are better adapted, physiologically, to hot, arid environments
than are scaled quail. This is quite plausible since Gambel quail inhabit extreme desert
areas while scaled quail are confined to more mesic areas with more moderate
temperatures. Scaled quail are better adapted with regard to water economy when
ambient temperatures do not exceed 40° C. Key words: Gambel's quail, scaled
quail, temperature, hydration

205

Hennessy, T. E. and L. Van Camp. 1963. Wintering mourning doves in northern
Ohio. Journal of Wildlife Management 27(3):367-373. Notes: Winter Injuries -Winter
injuries to doves in the form of frozen toes and losses of plumage have been reported
by several workers. On this study, 26 of 36 birds trapped in Huron County during the
winter of 1961-62 had one or more frozen toes. Some of the birds, probably adults, had
lost nails from the digits and had one or more enlarged, club-like toes. After a severe ice
storm on February 25, 1961, which covered the ground and vegetation with an inch of
ice, 8 doves from a flock of 60 had lost their tails completely. Others had lost some tail
feathers. At ground roosts, after such weather, Van Camp has found dead doves frozen
to their perches. Several doves, trapped after a wet snow followed by a sharp drop in
temperature, had balls of frozen snow on their lower breast feathers. Some birds
exhibited tears up to 2.5 cm in diameter in the skin of their breasts. It is unlikely that the
weight of the snow caused the tears. Probably they were self-inflicted when doves were
freeing themselves from perches to which they had become frozen. Key words:
mourning dove, temperature, ice, snow, mortality

206

Herman, C. M. 1969. The impact of disease on wildlife populations. BioScience
19(4):321-330. Notes: "A possible relationship between the intensity of coccidian
infection of California quail and food habits was observed some years ago. During the
dry summer months, when intensity of coccidian oocyst output was at its lowest, the

% f \ ^ ^

93

Cgj: ^ ^^ <£>

birds subsisted mainly on seeds. During the months of heaviest rainfall, the birds fed
mainly on leafy plants and had many more coccidial oocysts in their feces." Key words:
California quail, precipitation, diet, vegetation, disease, mortality

207

Herman-Brunson, K. M. 2007. Nesting and brood-rearing habitat selection of
greater sage-grouse and associated survival of hens and broods at the edge of
their historic distribution. M.S. Thesis, South Dakota State University, Brookings,
USA. Notes: Page 76 — "Exposure to wet and cold weather can also reduce survival of
chicks. I found high mortality to chicks exposed to rain and cold weather immediately
after hatch. Greater precipitation in 2005 resulting in increased herbaceous cover and
delayed plant desiccation may have resulted in higher survival of chicks >5-6 weeks to
1 January in 2005." Key words: precipitation, temperature, mortality, vegetation,
nutrition

208

Hermann, M. F. 1980. Spruce grouse habitat requirements in western Montana.
Dissertation, University of Montana, Missoula, USA. Abstract Note: "Climatic
differences between the 2 years somewhat limited the availability of berry-producing
plants. Thus, in 1977, the birds supplemented their diets with lodge pole pine,
ponderosa pine, Douglas-fir, and Engelmann spruce needles coupled with grasses,
sedges, and forbs." Key words: spruce grouse, temperature, vegetation, diet

209

Hernandez, F., J. D. Vasquez, F. C. Bryant, A. A. Radomski, and R. Howard. 2002.
Effects of Hurricane Bret on northern bobwhite survival in south Texas.
Proceedings of the National Quail Symposium 5:87-90. Abstract: The impacts of
intense storms such as hurricanes on wildlife rarely are documented. We had the
opportunity to monitor the impact of Hurricane Bret on northern bobwhite {Colinus
virginianus) survival and reproduction in Brooks County, Texas. On 23 August 1999,
Hurricane Bret struck our study area, which received >45 cm of rain and experienced
wind gusts >160 km/h. We documented the survival of bobwhite adults (n=82), broods
(/7=15), and nests (n=4) via radiotelemetry before and after the hurricane. Only 1 1
(13%) adult bobwhites were killed, with 4 killed directly from exposure to the hurricane.
Broods experienced higher mortality, with 7 (47%) broods killed during the hurricane.
Six of the 7 dead broods were <1 week old. Sizes of the 8 surviving broods were
reduced from a mean brood size of about 1 1 chicks prior to the hurricane to a mean
size of 4 after the hurricane (P=0.01 ). Of the 4 nests monitored, 3 were depredated and
eggs in 1 nest hatched the weekend of the storm. Hurricanes may negatively impact the
survival of young (i.e., <2 weeks old) bobwhite broods. Key words: bobwhite quail,
wind, hurricane, mortality, predation

210

Herzog, P. W. 1980. Winter habitat use by white-tailed ptarmigan in southwestern
Alberta. Canadian Field-Naturalist 94(2):1 59-162. Abstract: Winter habitat and

94 jj

4*

H fefc ^

J"."*-

cCi

movement of white-tailed ptarmigan (Lagopus leucurus) were investigated in Alberta
during January and February of 1977 and 1978. The critical feature influencing habitat
use appeared to be the availability of food, mainly willow (Salix spp.). In 1977, most
sightings of ptarmigan occurred in alpine cirques where low-growing willows remained
snow-free. Alpine willows were completely snow-covered in 1978, when 99% of
ptarmigan sightings occurred along stream courses and willow flats in subalpine forest,
2.5-7.5 km from cirque habitats. Food availability, determined by snow accumulation,
may be an important factor influencing the migration of ptarmigan. Page 161 — I suggest
that the geographic differences in migration of willow and sage-grouse may be due to
variations in snow accumulation between different areas. Key words: white-tailed
ptarmigan, snow, vegetation, nutrition, behavior, habitat use, movement

211

Hewitt, D. G. and R. L. Kirkpatrick. 1997. Ruffed grouse consumption and
detoxification of evergreen leaves. Journal of Wildlife Management 61(1):129-139.

Abstract: Ruffed grouse (Bonasa umbellus) in the southeastern United States
commonly eat evergreen leaves during winter. Despite their abundance, these leaves
rarely make up >50% of the diet. We fed diets containing 20 and 40% mountain laurel
(ML) (Kalmia latifolie) and Christmas holly fern (CHF) (Polystichum acrostichoides) to
captive ruffed grouse to determine the bird's ability to exist on these forages and to
investigate detoxification of secondary plant compounds. Grouse consuming diets with
20% of test forages performed similar to grouse eating a control diet. Diets with 40% of
the test forages caused reduced intake and birds consuming the CHF diet were unable
to maintain body mass. Conjugate-based detoxification systems were stimulated by
both test forages, although detoxification strategies varied between forages and levels
of forage in the diet. Although grouse appear unable to exist solely on evergreen
leaves, these forages probably contribute to ruffed grouse winter survival by remaining
available during snow accumulation and by decreasing foraging times due to high
intake rates. Key words: ruffed grouse, snow, nutrition, vegetation

212

Hillman, C. N. and W. W. Jackson. 1973. The sharp-tailed grouse in South Dakota.
Technical Bulletin Number 3, South Dakota Department of Game, Fish and Parks,
Pierre, USA. Notes: In the segment titled Limiting Factors the authors address effects
of weather on sharp-tailed grouse. They state these birds in South Dakota "...endure
cold temperatures and high winds with little known adverse effects. During periods of
deep snow, birds burrow or roost beneath the snow which provides insulation from cold
temperatures and wind. Grouse resort to tree budding when deep snow covers their
preferred foods. During mild winters the birds are located throughout the prairie, but
during extreme weather conditions, the birds retreat to sheltered areas such as river
bottoms or windbreaks. Severe blizzards, which may cause extensive pheasant losses,
do not greatly affect grouse populations. In many areas a cold, wet spring is detrimental
during the nesting and brood rearing periods. In South Dakota, however, wet spring
weather appears favorable to the grouse hatch. Age ratios obtained from fall bag
checks indicate better reproduction during relatively wet years as compared to hot, dry
years. Reproduction the following year is probably increased because of the additional

\ f^ffc.*^

95

<j$£ ^ ^ ££>

birds in the breeding population and more residual cover resulting from the extra
rainfall. Hail storms or local hard spring rains may be detrimental to broods in localized
areas." Page 58 — "Weather conditions over large areas influence habitat conditions
which in turn may cause fluctuations in sharptail numbers." Key words: sharp-tailed
grouse, temperature, snow, wind, precipitation, hail, behavior, mortality,
vegetation, habitat use, reproduction

213

Hjorth, 1. 1966. Influence of abiotic factors upon the display of the black grouse
Lyrurus tetrix. Var Fagelvarld 25(4):289-314. Abstract: The following is a brief outline
of a preliminary statistical investigation of data from 1962-1964 concerning the
correlation of display activity of the black grouse with weather and ground conditions.
The field work was done on the Gyamossen bog, about 7 m to the west of Jassjo in the
central part of the highlands of southern Sweden. At those periods of the year, during
which the black grouse had display activity, daily barograph and thermohygrograph
recordings were taken. The weather situation (clouds, precipitation, rime, mist, wind,
etc.) was written down 4 times/day. Light evaluations were made with a luxmeter every
10 min during the lek observations. Some of the factors which influence the display
were viz. motivation, ground conditions, weather, light, activity of other cocks and hens,
territorialism. Thus, sign stimuli belonging to display behavior emanate partly from the
biotic and partly from the abiotic environment. Here only the abiotic factors are treated.
Activities belonging to lek behavior are released by a stimuli-chain, in which the different
links offer new factors in each situation on the cocks' way from the nightquarters till
arrival at the arena. In spring, when motivation is at a high level, the cocks normally
ignore weather conditions (except light). Then the birds come to the arena even when
there are strong winds or when rain is falling. Therefore an analysis of the significance
of the weather factors must exclude the spring data. During the rest of the year abiotic
factors had strong influence upon the display activity. This report contains data from
September 1 to March 15 only. The black grouse begins the morning display later when
snow covers the ground of its nightquarters. When the ground is bare he flies to the
arena early at dawn. If, however, the hollows of the arena bog are covered with frozen
slush, the cocks leave the place rather quickly, returning at sunrise. The snow itself did
not have a negative influence on the display motivation, but lek was hindered when the
loose snow-covering exceeded 6 cm. Display motivation was often manifested in the
tree-tops on those occasions. A thick snow-covering also affected the territorialism on
the arenas. Then the territory boundaries (well-known to each territorial cock in bare
ground conditions) cannot be recognized and more real fighting occurred as a result. In
this report Chi2-tests were used to test the relationship between air pressure
tendencies, amount of clouds, temperature tendencies and wind velocities respectively
and estimates of the display intensities observed. There was apparently a relationship
between increasing air pressure and display activity, while decreasing pressure did not
seem to have much significance. Low morning temperatures (ranging from -20°C to
about 2°C) were more favorable to lek than higher ones. In the latter case the cocks
display over a wide range of their habitat. Low temperature conditions seem to favor
aggregation and territorialism on the arena. Temperature tendency had a strong effect
on lek activity. A lowering of temperature stimulates the cocks to lek activity while rising
or unchanged conditions did not have any significant effect. The influence of falling

96 ^

4>

H fee ^

l&- 4^> Q>

temperature was much stronger in mild weather than in cold weather. The velocity of
wind near the ground affected the grouse more in open places, where the arena is
situated, than in the forest, where the birds have nightquarters. Thus, in the forest,
velocities over 3 m/sec counteracted display activity, but in the arena values over 2
m/sec made the cocks give up lek activity entirely. Clear weather was more favorable to
lek than was a cloudy sky. In this method of correlation the influence of other weather
factors than that treated cannot be estimated. In addition, meteorological factors were
not freely varied but were dependent on each other. The results presented must not be
taken as examples of obligate relationships. The weather factor which seemed to favor
display or lek could have varied in accordance with some other meteorological quantity,
which also stimulated the cocks. It was evident, however, that all weather conditions,
favorable to display, were to be found in one and the same meteorological situation, viz.
the high pressure condition. When all positive factors, occurred the same morning, lek
will always follow. Key words: black grouse, wind, precipitation, ice, snow,
barometric pressure, clouds, behavior

214

Hoffman, D. M. 1973. Some effects of weather and timber management on
Merriam's turkeys in Colorado. Proceedings of the National Wild Turkey
Symposium 2:263-271. Abstract: The climate of the range occupied by Merriam's
turkey (Meleagris gallopavo merriami) in Colorado is usually not severe, because flocks
move from high summer range to preferred winter range at lower elevations.
Measurements of snowfall and precipitation within two preferred winter areas are listed.
Occasionally, as in the 6-week period from early February 1964 to mid-March 1964,
frequent heavy snows and high winds resulted in unusual winter stress on turkeys,
unless food conditions are exceptionally good. Methods of harvesting timber affect
selection of roost sites and the establishment and maintenance of forest openings.
Selective cutting that removed approximately half of the old growth in stands of uneven-
age ponderosa pine (Pinus ponderosa) on two summer roost sites did not appear to
deter turkeys from using these sites for roosting. There were 17 other roost sites (9
winter and 8 summer sites) in unlogged tracts of old-growth timber. No roost sites were
found in second-growth timber after clear-cutting operations. Key words: Merriam's
turkey, snow, precipitation, wind, habitat modification

215

. 1963. The lesser prairie chicken in Colorado. Journal of Wildlife

Management 27(4):726-732. Notes: "A major reduction in lesser prairie chicken range
and numbers apparently coincided with the dust-bowl condition of the 1930's. Drought
and land-use practices undoubtedly contributed to this reduction." "Appropriate
grassland management is required for the survival and increase of the lesser prairie
chicken. The decline in number of lesser prairie chickens in Colorado has been
attributed to pasture depletion during the severe drought of the 1930's." "Best counts
were obtained on clear, calm mornings, when the cocks were most active on the
grounds. On mornings with strong winds or rain, the cocks were usually far less active."
Key words: prairie chicken, drought, wind, precipitation, census, vegetation,
population dynamics, behavior

^ fc* t*

if A;v i 97

216

. 1962. The wild turkey in eastern Colorado: a research and management

study of Merriam's wild turkey in eastern Colorado. Technical Publication No. 12.
Colorado Department of Game and Fish, Denver, USA. Notes: Page 8 - "Findings
also indicate that turkeys will desert an area when water becomes scarce. In the winter
when snow is available, the flocks will eat snow and drink from melting snow so that
open water is not needed as in the warmer seasons." "Most roosts are selected in
locations sheltered from high winds." Page 15 - "Predation plays an important role in
disposing of sick and weak birds and is probably more beneficial than harmful except
where predator populations become excessively large or the wild turkeys become weak
from lack of food caused by deep snow or other adverse conditions." Page 17- "Two
instances in which adverse weather affected wild turkey populations were recorded
during this study. Purgatoire River Flood: During late July 1954 a flood from heavy rains
east of Trinidad moved down the Purgatoire River. Ranchers in the Higbee area
reported that a strip of land approximately a half mile wide was flooded. Mr. W. W.
Zimmerman stated that he and several other local ranchers pulled four live wild turkey
hens from the flood waters near a wide curve in the main channel above Higbee bridge.
Several other hens could be seen, but could not be reached. Those rescued were
completely exhausted, but after resting awhile, they wandered off. Later the same party
pulled three dead hens from the waters of the main channel and two more from an
irrigation ditch. The number of hens caught in the flood at this time of year indicates that
the hens were probably attempting to hatch a second setting of eggs. Much of the
nesting cover in this area is found along the main river bottom where a flood of this
intensity can be dangerous. Late Spring Snows: The late heavy snow during May 1955
undoubtedly ruined many wild turkey nests on the eastern slope mountain ranges
although the turkeys in much of the Lower Purgatoire River and Mesa de Maya areas
were apparently unaffected. The snow, which accumulated to depths varying between 3
and 5 feet in most mountainous areas came when most hens are normally nesting. The
effect of this storm was a reduction in the number of broods in the mountain areas.
Many broods were much later and much smaller than usual, indicating that many hens
were forced to renest in order to bring off any young. Two carcasses and seven ruined
nests were reported in Huerfano and Las Animas counties following these heavy
snows." Key words: Merriam's turkey, snow, flood, mortality

217

Hoglund, N. H. 1980. Winter ecology of the willow grouse Lagopus lagopus
lagopus. Swedish Wildlife Research Viltrevy 11(5):249-270. Abstract: The
Loevhoegen area, situated on the border between Dalarna and Haerjedalen at latitude
62°N, was studied during mid- and late winter, 1966-1968. The grouse's food consisted
almost exclusively of twigs, buds and birch catkins. The birds always fed walking on the
snow. When not occupied by eating, the grouse roosted, always on the ground and
preferably in snow burrows. The night burrows were completely closed, dug below the
surface of the snow during January and February. In March, grouse begin to roost for
the night in open burrows sbd pits in the snow. The time spent by the grouse in the
burrow could be estimated by counting the pellet-shaped feces in it. The diurnal rhythm

98 ^

4>

^ ^ fer ^

<^>

in the activity of the grouse, the alternation between food gathering and roosting, was
adapted to light and weather conditions. A strong coherence within the flocks was
noted. The activity of the grouse was very concordant, during food gathering as well as
during roosting. Under snow conditions, with available nutrition, the grouse population
remains stationary. Otherwise, it moves down to lower situated areas with more readily
accessible food, especially the hens. Key words: willow grouse, snow, light,
behavior

218

Holloran, M. J., B. J. Heath, A. G. Lyon, S. J. Slater, J. L. Kuipers, and S. H.
Anderson. 2005. Greater sage-grouse nesting habitat selection and success in
Wyoming. Journal of Wildlife Management 69(2):638-649. Abstract: Nesting habitat
degradation and its negative effect on nesting success might contribute to the recent
population and distributional declines of greater sage-grouse (Centrocercus
urophasianus) throughout North America. We used radiotelemetry to locate greater
sage-grouse nests in 7 different areas of central and southwestern Wyoming between
1994 and 2002; we studied each area for 2 to 4 years. Using binary logistic regression,
we compared microsite vegetal data collected at nests (n = 457) and random (n = 563)
sites and successful (n = 21 1 ) and unsuccessful (n - 238) nests to test hypotheses
concerning greater sage-grouse nesting habitat selection and vegetal conditions
associated with nesting success. We used Akaike's Information Criterion (AICC) and
model averaging to make inference about the weighted support for the importance of
individual habitat variables through the comparison of sets of competing models.
Selected nest sites were located in areas with increased total shrub canopy cover
(relative importance [Rl] = 1.00), residual grass cover (Rl = 0.47) and residual grass
height (Rl = 0.77) compared to random sites. Successful nests had increased residual
grass cover (Rl = 0.43) and height (Rl = 0.48) relative to unsuccessful nests.
Additionally, annual nest success rates (i.e., above vs. below our study's average) were
related to the preceding year's spring (Apr-May, Rl = 0.44) and winter-early spring (Jan-
Jun) precipitation (Rl = 0.32). Correct classification rates for weighted average models
that we derived through the 3 comparisons were between 60 and 70%, suggesting the
variables adequately differentiated between plot types. However, high model selection
uncertainty (i.e., the total number of models included in the sets of AICc-selected
models) suggested that nest site selection and nesting success may be influenced by
factors not considered in the modeling process. Management strategies that protect
dense sagebrush stands and enhance residual grass cover and height within those
stands should be used to maintain nesting habitat and increase nesting success of
greater sage-grouse. Key words: sage-grouse, precipitation, habitat use, model,
vegetation

219

Homan, H. J., G. M. Linz, and W. J. Bleier. 2000. Winter habitat use and survival of
female ring-necked pheasants in southeastern North Dakota. American Midland
Naturalist 143:463-480. Abstract: From 1992 to 1995 we used radiotelemetry to
monitor winter habitat selection and survival of female ring-necked pheasants
(Phasianus colchicus) in southeastern North Dakota. We captured 100 birds at nine

% ;MV<tf*>f*c

99

sites in six study blocks centered on cattail-dominated {Tyha spp.) semi-permanent
wetlands. Pheasants showed nonrandom habitat use at two hierarchical scales. At the
second-order scale (23-km2 blocks) semi-permanent wetlands were preferred during
two winters in which habitat selection could be assessed (1992-1993 and 1994-1995).
An additional second-order preference for grass-covered uplands was shown during the
mild 1994-1995 winter. At the third-order scale (home-range) pheasants preferred the
edges of wetlands in 1992-1993 and 1994-1995. The central portions of wetlands were
preferred in 1992-1993 and used proportionately in 1994-1995. The average winter
survival rate was 0.41, with rates ranging from 0.04-0.86 and differing significantly
among winters. Survival was lower during early winter and midwinter periods for birds
weighing less than 1090 g and for birds captured in semi-permanent wetlands under
private ownership. A 1°C increase in the mean weekly maximum temperature
decreased the probability of death by 0.06 and a 2.5 cm increase in new snow raised
the probability of death by 0.08. Key words: ring-necked pheasant, temperature,
snow, mortality

220

Hoodless, A. N. and J. C. Coulson. 1998. Breeding biology of the woodcock
Scolopax rusticola in Britain. Bird Study 45:195-204. Abstract: A total of 449 nest
record cards collected by the British Trust for Ornithology during 1945-89 was used to
examine timing of nesting and brood production of the woodcock. Daily chick mortality
rates were estimated from recaptures of 26 woodcock broods during an intensive study
in 1988-92. Clutches were initiated between March and July, with a mode at the end of
March. On average, egg laying commenced earlier in England (median first-egg date 8
April) than in Scotland (20 April) and there were negative relationships between first-
egg dates and mean March air temperatures, both between years and regionally within
years. Nest survival during egg laying and incubation was 41 ± 1% and the mean
number of clutches hatched per female alive in March was 0.89 (95% CL 0.47-1 .58).
Chick survival until fledging was estimated as 56 ± 8%, resulting in a mean annual
production of 1 .80 ± 0.72 fledged young per female alive at the start of the breeding
season. A small second peak in the distribution of first-egg dates in mid-May indicates
that in some years females may attempt to rear second broods. Key words:
woodcock, temperature, reproduction, productivity

221

Hornell-Willebrand, M., V. Marcstrom, R. Brittas, and T. Willebrand. 2006.
Temporal and spatial correlation in chick production of willow grouse Lagopus
lagopus in Sweden and Norway. Wildlife Biology 12(4):347-355. Notes: The authors
believe that variation in annual production of willow grouse is largely affected by
external factors, stating "Most likely, weather plays an important role in large-scale
synchronization, but other factors such as food quality and predation are also known to
be important. Of course none of these factors are independent, and they may interact in
different ways. Weather seems to be the only likely candidate explanation for the long-
distance correlations in the harvest data, but neither Hammarstrom nor Haakenstad
found any evidence that weather was affecting the juvenile ratio in the harvest data.
Brittas showed a positive relationship with spring weather, digestibility of food,

100 ^

4<

K <o **

cCi

abundance of microtine rodents and the breeding success of willow grouse at site No.
1 . The two years with the lowest breeding success were years with poor willow grouse
nutrition and condition together with a decline in small mammal populations. Extremely
bad weather during the first week after hatching has been shown to increase chick
mortality. Slagsvold found a positive correlation between a high mean temperature in
the middle of June and the breeding success of willow grouse. Myrberget et al showed
a similar relationship, with warm weather during the early chick period having a positive
effect on survival and growth rate of chicks. However, many studies have not found any
correlation between chick survival and weather. It is difficult to evaluate weather data
without an understanding of the potential mechanism between weather and breeding
success. Climate changes have strong impacts on wildlife species at high elevations,
and higher temperatures have the potential to alter the amount of alpine and subalpine
habitat and to increase alpine fragmentation because of rising subalpine tree line. Long-
term monitoring programs for alpine and subalpine wildlife, like the Swedish willow
grouse census, will be important when even small increments in warming are shown to
significantly impact habitat quality and quantity for breeding and migration in other
Lagopus species. Key words: willow grouse, precipitation, temperature,
reproduction, mortality, nutrition

222

Hornfeldt, B., T. Hipkiss, and U. Eklund. 2001. Juvenile sex ratio in relation to
breeding success in capercaillie Tetrao urogallus and black grouse T. tetrix. Ibis
143:627-631. Notes: Page 629 — "...many studies indicate the importance of weather to
breeding success in capercaillie. Frequent rain, especially around the time of hatching,
and late snowmelt affect breeding success detrimentally in capercaillie, but rain around
the time of hatching was found not to affect black grouse breeding success. Adverse
weather conditions affect juvenile male capercaillie more severely than females, since
males have higher energy requirements even under normal conditions, and this might
also be the case for black grouse. Additionally, the increased food requirements of
males may force them to forage during adverse weather, while females may be able to
take shelter." Key words: capercaillie, black grouse, precipitation, snow,
temperature, productivity, sex ratio, behavior

223

Host, P. 1942. Effect of light on the moults and sequences of plumage in the
willow ptarmigan. Auk 59:338-403. Summary: Over several years the author kept
between thirty and sixty willow ptarmigan, in an attempt to raise them in captivity on a
large scale. Observations on these birds, especially during the breeding season,
showed a marked synchronization of the plumage cycle with the different phases of
reproduction. Smaller numbers of the birds were used at different times for experiments
to test the effects of light and temperature on plumage and reproduction. Birds kept at
normal, low temperatures during the winter months, November to February, were
exposed to artificial light day and night. They responded to these changes in light
conditions by showing different signs of reproductive activity, most marked in a female
that laid a normal clutch of eggs in the last days of December and the beginning of
January. While all the control birds kept their winter plumage unchanged during this

f>' ^ N^C"

4

<£^ cCi

period, all birds that were exposed to light developed different phases of spring
plumage and summer plumage, thus indicating that light and not temperature is the
main controlling factor for both the reproductive cycle and the development of spring
and summer plumages in this species. One of the birds, a male, that showed fully
developed spring plumage by the beginning of February, had day-length again reduced
to seven hours a day. It responded to this by moulting and developing new white
feathers, and within two months had completely changed from spring plumage directly
into a new winter plumage. The winter plumage was developed in spite of the fact that
the bird was kept at temperatures considerably higher than normal at the time of year
when winter plumage develops in nature. Five ptarmigan, that had the normal day-
length reduced at the beginning of August, responded to this by developing winter
plumage one month earlier than the control birds which developed this plumage at the
normal time. These experiments indicate that the development of fall plumage and
winter plumage also are controlled by light conditions, and are caused by the reduced
length of day in fall and winter. Several observations from the different experiments are
mentioned, showing that even very small changes in light conditions can produce
marked effects on plumage development. Key words: willow ptarmigan, solar
radiation, temperature, reproduction

224

Hubalek, Z., J. Halouzka, and Z. Juoicova. 2003. Host-seeking activity of ixodid
ticks in relation to weather variables. Journal of Vector Ecology 28(2):15-165.

Abstract: Ixodid ticks were monitored in a temperate deciduous broad-leaved forest in
South Moravia (Czech Republic). Relative abundance of the ticks collected before noon
(10:00-12:00 h) was compared to several weather variables (air and soil temperatures,
relative humidity, precipitation, wind speed, and derived values) using the Pearson
correlation coefficient. The tick numbers were found to be most closely related to the
amplitude of the soil (-5 cm) temperature between 07 h and 14 h (TSamp, in Ixodes
ricinus), and the soil temperature (TS) at noon (in Haemaphysalis concinna) or in the
morning {Dermacentor reticulates). While a growing amplitude of TSamp caused an
increased host-seeking activity of /. rincinus and H. concinna, it suppressed the activity
of D. reticulates, a tick species mainly occurring in colder seasons of the year in Central
Europe. The air temperature (TA) and relative humidity (RH) were also closely related
to the tick activity, whereas rainfall and wind speed remained largely uncorrected with
the activity of the three tick species. Multiple linear regression on several variables
(TSamp, TA, TS, TA-TS, RH) explained 48% of the variance in /. ricinus, 47% in H.
concinna, and 38% in D. reticulates. Predictive two-variable regression models of
relative abundance in host-seeking ticks were based on morning temperature (TA or
TS) and morning RH as the most important environmental factors: they explained 32%
(/. ricinus), 39% (/-/. concinna), and 35% (D. reticulates) of the variance. Key words:
temperature, soil, precipitation, humidity, wind, insect, behavior

225

Hudson, P. J., M. Cattadori, B. Boag, and A. P. Dobson. 2006. Climate disruption
and parasite-host dynamics: patterns and processes associated with warming
and the frequency of extreme climatic events. Journal of Helminthology

102 jj

4>

» te, ^

<^>

80(2):175-182. Abstract: Levels of parasitism and the dynamics of helminth systems is
subject to the impact of environmental conditions such that we may expect long term
increases in temperature will increase the force of infection and the parasite's basic
reproduction number, R-O. We postulate that an increase in the force of infection will
only lead to an increase in mean intensity of adults when adult parasite mortality is not
determined by acquired immunity. Preliminary examination of long term trends of
parasites of rabbits and grouse confirm these predictions. Parasite development rate
increases with temperature and while laboratory studies indicate this is linear, some
recent studies indicate that this may be non-linear and would have an important impact
on R-O. Warming would also reduce the selective pressure for the development of
arrestment and this would increase R-O so that in systems like the grouse and
Trichostrongylus tenuis this would increase the instability and lead to larger disease
outbreaks. Extreme climatic events that act across populations appear important in
synchronizing transmission and disease outbreaks, so it is speculated that climate
disruption will lead to increased frequency and intensity of disease outbreaks in parasite
populations not regulated y acquired immunity. Key words: grouse, disease,
parasites, temperature, climate change

226

Huempfner, R. A. and J. R. Tester. 1988. Winter arboreal feeding behavior of
ruffed grouse in east-central Minnesota. Pages 122-157 in A. T. Bergerud and M.
W. Gratson, editors. Adaptive strategies and population ecology of northern
grouse. Wildlife Management Institute and University of Minnesota Press,
Minneapolis, USA. Notes: Page 128 — The mean numbers of grouse per km observed
each day during evening budding were 9.0, 1.8, and 2.2 for the three winters,
respectively. About half of this 82% reduction can be explained by a decline in the
ruffed grouse population based on fall flushing rates. The remainder was apparently
caused by variations in snow depth, snow quality, and duration of snow cover among
winters, and the actual timing of winter predation. The generally deep and more
favorable snow profile in 1973-74, compared with 1972-73, resulted in a slightly higher
daily observation rate in 1973-74, even though the grouse population was apparently
lower. Page 139 — When snows were crusty, total daily activity was increased as grouse
walked on the snow surface primarily in search of hazel catkins. This pattern changed
from April to June for both male and female grouse at Cedar Creek. Both sexes showed
increased daily activity during this interval, even though they began activity later and
terminated it earlier relative to sunrise and sunset, respectively. Our explanation for this
change from an early morning and late evening activity pattern in winter is based on the
elevated light levels in the woodlot in winter owing to snow cover and the absence of
leaves. It may be that daily activity of ruffed grouse begins and ends at similar light
levels year round, but that these levels occur at different times relative to sunrise and
sunset based on the presence or absence of snow and leaf cover... consistently cold
temperatures in data periods 8-10 appeared to cause equally consistent and early peak
feeding. A warmup during data periods 11-13 appeared to cause a shift in peak budding
to almost 8 minutes later. A gradual warming trend in 1972-73 from data periods 11 to
14 was also accompanied by a shift in peak budding from 32.5 minutes to 9.5 minutes
before sunrise. It appeared that ruffed grouse may have shifted the period of peak
morning feeding to compensate for a potentially large, negative energy balance during

% »'^>t*

103

/~H' *fO ^*\. y~^

% l@> <^ <9

particularly cold portions of the winter. Seiskari (1962) noted that feeding activity of
penned black grouse and capercaillie increased with cold weather and high barometric
pressure. Seiskari's data suggest that earlier morning feeding appeared to follow low
temperatures for each species. Unlike the morning period, peak evening budding was
poorly correlated with temperature. Key words: ruffed grouse, snow, temperature,
behavior, census, habitat use

227

Huffman, R. T. 2005. The effect of precipitation on Rio Grande wild turkey nesting
ecology in the Texas panhandle and southwestern Kansas. M. S. Thesis, Texas
Tech University, Lubbock, USA. Abstract: Nesting activity of Rio Grande wild turkeys
(Meleagris gallopavo intermedia) was monitored from 2000 through 2004 at 3 sites in
the Texas Panhandle and 1 site in southwestern Kansas. During this time 360 adult
female and 282 juvenile female wild turkeys attempted 396 nests. A total of 129 (33%)
nests successfully hatched out !Y1 (sic) live poult. Nesting rate did not vary among sites
(P = 0.2871) or years (P=0.2453), but did range from a low of 15% to a high of 58%. No
relationship was found between pre-nesting precipitation (September-March) and
nesting rate of first nest attempts (P = 0.207), or nesting success (P - 0.8495). Nesting
season precipitation (April-July) had no relationship to nesting success (P = 0.1542),
nor did the number of days with precipitation during nesting had no (sic) relationship to
nesting success (P = 0.106). Days since precipitation for depredated nests (moisture-
facilitated nest depredation hypothesis) did not conform to any distribution pattern.
However, almost 30% of nests were depredated on days with measurable precipitation.
Using data from 269 nests stepwise logistic regression selected 3 variables into a
model predicting nesting success. Pre-nesting precipitation (PRE) and number of
rainfall events during nesting (EVENTS) had a positive relationship to the success of a
nest while the week of nest initiation (WEEK) had a negative relationship. More
precipitation during pre-nesting, more rainfall events during nesting, and initiation of a
nest early in the nesting season increased the probability of a nest being successful.
Height of visual obstruction did not differ between successful nests and unsuccessful
nests. Mean height of visual obstruction at successful nests and unsuccessful nests
was 0.5 m. Regardless of nesting outcome nest sites had greater height of visual
obstruction than paired random plots. This relationship held among all years and all
sites except at the Matador Wildlife Management Area in 2001 . In 2003 and 2004
females selected for greater height (0.4-0.5 m) of visual obstruction and avoided lower
height (0. 1 -0.2 m) of visual obstruction (P < 0.001 ). In 2004 nests sites with trees in the
nesting area were selected for while nest sites without trees were avoided across all
study sites (P < 0.001 ). Managers can use local weather data to determine the
probability of nests being successful; this may allow for estimates of recruitment into the
fall populations. Managers should develop habitat management plans that maximize
visual obstruction availability for potential nesting habitat. Key words: Rio Grande
turkey, precipitation, reproduction, productivity, vegetation, predation

228

Hungerford, C. R. 1964. Vitamin A and productivity in Gambel's quail. Journal of
Wildlife Management 28(1):141-147. Abstract: The influence of available natural

104 j,

)'>'• ^, ^

#*

sources of vitamin A on reproduction of Gambel's quail (Lophortyx gambelii) in
southeastern Arizona was investigated in a 5-year study. Birds were observed and
collected auring critical seasons on three basic types of quail range. The amount of liver
vitamin A in a sample of 177 birds was found to vary with rainfall and the resulting green
plant growth. Differences in covey behavior, reproductive activity, and a size variation in
reproductive organs was associated with differences in vitamin A storage. Vitamin A or
a closely associated substance derived from green plant material apparently acts as a
stimulator which influences the rate of breeding in this desert quail. Key words:
Gambel's quail, precipitation, fecundity, nutrition, habitat quality, vegetation

229

Hupp, J. W. 1987. Sage-grouse resource exploitation and endogenous reserves in
Colorado. Dissertation, Colorado State University, Fort Collins, USA. Abstract:
Winter habitat use and foraging ecology of sage-grouse {Centrocercus urophasianus)
were studied in the Gunnison Basin, Colorado between 1985 and 1986. Sage-grouse
foraging activity (A/=157 feeding sites) was not proportionally distributed (PO.001 )
among topographic features. Topographic distribution of feeding activity was influenced
by physiographic variation in shrub structure of mountain big sagebrush {Artemisia
tridentata vaseyana) relative to snow depth. Most foraging (45-64% of feeding sites)
occurred in drainages and on southwest slopes where sagebrush exposure above snow
was maximized. Sage-grouse rarely foraged (1-4% of feeding sites) on northeast
aspects with slopes >5° because exposed sagebrush was not widely available.
Distribution of foraging was not influenced by topographic variation in crude protein or
monoterpene concentrations of mountain big sagebrush. Within feeding sites, sage-
grouse did not selectively forage on plants with high crude protein or low monoterpene
concentrations. Sagebrush structural characteristics at 87 winter feeding sites were
compared to 100 random locations. Sagebrush structural measures were not useful to
identify winter habitat in the mesic terrains where sagebrush removal is most likely to
occur. Spring lipid reserves of adult male sage-grouse were determined from carcass
analysis of 96 individuals collected from 2 Colorado populations between 1983 and
1985. Spring lipid reserves were affected by winter severity. Males depleted lipid
reserves during the courtship season. Males in Jackson County mobilized 125-130 g of
fat during courtship while males in Gunnison County used an average of 66 g. Strutting
display rates of adult males were quantified in Jackson and Gunnison counties in 1986.
Slower display rates, lack of evening display, greater variance in male attendance at
leks suggest reduced energetic investment in courtship among males in Gunnison
County. However, behavioral differences could not be attributed to unequal size of
endogenous reserves as lipid deposits (A/=10 adult males/population) during early
courtship were similar between populations in 1986. Lipid catabolism likely provides
<10% of adult male energetic needs during courtship. Lipids may primarily be mobilized
during early courtship when male displays are most vigorous due to the presence of
females on leks, and when male reproductive success is determined. Key words:
sage-grouse, vegetation, snow, habitat use, nutrition

i fate***

105

> • -,■ Q

230

and C. E. Braun. 1991. Geographic variation among sage-grouse in

Colorado. Wilson Bulletin 103(2):255-261. Notes: "Winter severity affects body
condition of sage-grouse and may cause body mass to vary. The winter of 1983-84 was
especially severe in Colorado, and lipid reserves and body mass of adult males were
lower than in other years. Carpal length also varied among years and suggests that
growth and wear of sage-grouse remiges may be influenced by environmental
conditions." Key words: sage-grouse, severe weather, condition

231

and . 1989. Topographic distribution of sage-grouse foraging in

winter. Journal of Wildlife Management 53(3):823-829. Abstract: We studied
sagebrush (Artemisia spp.) exposure above snow and topographic distribution of sage-
grouse (Centrocercus urophasianus) foraging sites in winter (Jan-Mar) in the Gunnison
Basin, Colorado. Sage-grouse feeding activity {n = 157 foraging sites) was not
proportionally distributed among 5 topographic categories (P < 0.001 ). Most (46 and
75% of foraging sites in 1985 and 1986, respectively) feeding activity occurred in
drainages and on slopes with south or west aspects. Use of slopes with north or east
aspects was less than expected. Distribution of sage-grouse feeding activity was
influenced by topographic variation in snow depth and mountain big sagebrush (A.
tridentata vaseyana) exposure above snow. During a severe winter in 1984, <10% of
the sagebrush vegetation in the Gunnison Basin was exposed above snow and
available to sage-grouse. During milder winters in 1985 and 1986, exposure of
sagebrush was 84 and 79%, respectively. We recommend that sagebrush be
maintained in drainages and on slopes with south or west aspects. Key words: sage-
grouse, snow, temperature, habitat use, nutrition, vegetation

232

and . 1989. Endogenous reserves of adult male sage-grouse during

courtship. Condor 91 :266-271 . Abstract: Lipid reserves of 1 1 6 adult (>1 year of age)
male sage-grouse {Centrocercus urophasianus) were evaluated in two Colorado
populations during lek attendance between 1983 and 1986. Lipid reserves following
winters (November-March) with snowfalls <124 cm were larger (P<0.001) than reserves
following winters with snowfalls >160 cm. Lipid reserves during early courtship were
larger than reserves during late courtship (P<0.001). Males catabolized lipids during
courtship but did not use breast muscle protein. Catabolism of lipids likely provides <5%
of male energetic requirements during courtship. An adaptive advantage to fat
deposition before breeding may exist if males primarily mobilize lipids during the peak
period of female lek attendance when male reproductive success is determined, or
during periods when thermoregulatory costs are high due to low ambient temperatures
or wind Key words: sage-grouse, wind, temperature, snow, bioenergetics

233

Ivacic, D. L. and R. F. Labisky. 1973. Metabolic responses of mourning doves to
short-term food and temperature stresses in winter. Wilson Bulletin 82(2):182-

106 \,

N^

H fea, t*

0>"^

c^>

196. Summary: Winter-acclimatized mourning doves were subjected to a photoperiod
and an ambient temperature range designed to simulate conditions of a severe winter
storm, common in central Illinois. Metabolic rates were recorded for two daily cycles of
decreasing temperatures (10°C to -18°C) in darkness and increasing temperatures (-
18°C to 10°C) in light; the doves were fasted throughout the 44-hour treatment. Juvenile
doves had higher metabolic rates than adults, and females had higher rates than males,
but neither difference was statistically significant (P > 0.05). The metabolic rates of
doves were greater during exposure to decreasing ambient temperatures and darkness
than during exposure to increasing ambient temperatures and light, the difference being
statistically significant (P < 0.05). The doves responded metabolically more to the
directional temperature gradient than to either actual ambient temperature or light. The
consumption of oxygen by doves did not follow a straight line increase with concurrent
declines in ambient temperatures. Oxygen consumption was not only less at -18°C than
at -10°C (in darkness), but also diminished with extended exposure at -18°C. These
observations suggested that mourning doves employed a physiological mechanism
(perhaps reduced body temperature) to decrease their metabolic expenditures at low
ambient temperatures; the mechanism permits the conservation of energy and thus
augments survival. The potential survival ability of doves exposed to simulated winter
storms was related to sex and age attributes. Adult doves (particularly males), because
of their age or greater weight, or both, were better equipped physiologically than
juveniles (particularly females) to survive the stresses of low ambient temperatures
without food. Thus, among mourning doves wintering in the northern and central U.S.,
the autumn-to-spring changes in sex ratios and in age ratios, which usually favor males
and adults, respectively, probably reflect the differential survival abilities of the sex and
age cohorts. Key words: mourning dove, temperature, mortality

234

Jackson, A. S. and R. DeArment. 1963. The lesser prairie chicken in the Texas
Panhandle. Journal of Wildlife Management 27(4):733-737. Abstract: Trends in
populations of lesser prairie chicks (Tympanuchus pallidicinctus) in the Texas
Panhandle were investigated by censusing drumming grounds annually on two study
areas during a 10-year period, 1952-62, for comparison with data from a census of the
same areas in 1942. Severe drops in populations came in 1952. The decline was
triggered by onset of a major drought lasting through 1956, but populations did not
increase during a series of good rainfall years starting with 1957. Changing land-use
practices are responsible for keeping lesser prairie chickens at low population levels in
the Texas Panhandle. The more important of these are overgrazing of cattle range,
particularly in dry weather, resulting in displacement of the tall grasses; accelerated
programs of aerial spraying with herbicides for brush control; and combine harvesting of
grain sorghum in place of storage by stacking and shocking in the field. Key words:
prairie chicken, drought, precipitation, population dynamics, vegetation, range
condition

235

Jarvis, J. M. 1973. The Parker Mountain sage-grouse study. Proceedings of the
Annual Conference of the Western Association of State Game and Fish

}x t* t*

J< 1>W i ... 107

Commissioners 53:345-352. Abstract: This was a three-year study of sage-grouse
(Centrocercus urophasianus) on the Parker Mountain in south central Utah. The
population trend was downward during the study because of poor production. The dry
warm springs of 1971 and 1972 allowed for early mating and nesting but the lack of
succulence indirectly increased brood mortality. The lack of production affected the
harvest so the hunter effort diminished as the population diminished. Observations of
individually color marked birds indicated cocks were loyal to one strutting ground in
succeeding years. That hens orient more toward nesting habitat than strutting grounds
for nest sites. Twenty-four percent (24%) of the population was harvested annually. Key
words: sage-grouse, temperature, precipitation, nutrition, vegetation, mortality,
production

236

Jenkins, D., A. Watson, and G. R. Miller. 1967. Population fluctuations in the red
grouse Lagopus lagopus scoticus. Journal of Animal Ecology 36(1):97-122. Notes:
The "browning" of heather by frost was found to be correlated with poor breeding with
supporting the hypothesis that breeding success depended on the condition of the hens'
food in early spring. The data from all study areas since 1960 confirm this hypothesis.
Variations in the heather index parallel variations in hatching date and clutch size in all
four years, and in 1962-64 they were also related to the clutch size, the dimensions of
eggs, the weight of chicks at hatching and the chicks' survival. An "intermediate" year
was related to early breeding and large clutches, but with poor chick survival, fairly
small eggs, and light chicks. The authors conclude early nesting and large clutches may
have been determined by food quality in early April with some other factor affecting egg
size and chick survival, perhaps new plant growth. They analyzed the weather records
from 1962-65 between the start of heather growth and egg laying. There were more
frosts in 1965, which might have delayed heather growth. Using available data, there is
a potential correlation between the first week of heather growth and breeding success.
Key words: red grouse, frost, vegetation, nutrition, reproduction

237

Johnson, D. E. 1999. Ruffed grouse productivity and habitat selection at the base
of the Beartooth Plateau in southcentral Montana. M.S. Thesis, Montana State
University, Bozeman, USA. Notes: Page 60 — "Likely causes of [chick] mortality
include weather, predation, physiological disturbance, and starvation." Keywords:
ruffed grouse, predation, chick mortality, nutrition, weather

238

Johnson, R. A. 1927. The ruffed grouse in winter. Auk 44(3):31 9-321. Notes: The
author states that the more severe the weather, the less active are the ruffed grouse.
Also that the grouse is reluctant to leave roosting places on dark and stormy days. On
bright and sunny winter days, however, the ruffed grouse perch in conifers to take
advantage of the sunlight. In severe weather the grouse feeds in the very early morning
and in the late evening just before dark. In severe weather, after some snow has
accumulated, the ruffed grouse favors a berth in the snow. Key words: ruffed grouse,
temperature, snow, behavior

108 j^

^

) > ^ ^

239

Jonas, R. J. 1968. Adverse weather affects Merriam's turkey reproduction in
Montana. Journal of Wildlife Management 32(4):987-989. Abstract: A 37-inch
snowfall was recorded in the Long Pines study area in southeastern Montana on May 1 ,
1967, near the end of egg laying and beginning of incubation for most Merriam's turkeys
(Meleagris gallopavo merriami). Compared with normal years (1961-62-63), most egg
laying or incubation was delayed or interrupted during the first 10 days in May, 1967,
and no successful hatching occurred May 1-20. The population was maintained by late
nesting or renesting. There was no evidence of renesting by hens that lost broods
during May 1-20, 1967. Key words: turkey, snow, reproduction

240

. 1966. Merriam's turkeys in southeastern Montana. Montana Fish and Game

Department, Technical Bulletin No. 3, Helena, USA. Notes: Page 15 — "Differences in
productivity may have been related to precipitation. For the three months critical to
nesting, hatching and rearing success (April, May and June), deviations in precipitation
from the long-term mean were as follows: 1961,-2.03"; 1962 +2.01" and 1963 +6.30".
The dry spring of 1961 had been preceded by two abnormally dry years. Possibly most
effects of precipitation in the dry years (1959, 1960, 1961) and the wet 1962 were
indirect through influence upon vegetation and hence cover. Cover was poor in 1961
and excellent in 1962. It was also superior in 1963; however, four heavy snows
occurred at intervals of about one week in April (one 17 inches) and rainfall was 4.81
inches above the average for June. Weather may have had a direct adverse affect upon
reproductive success that year. Further evidence of this is apparent from some late
hatching dates for 1963, resulting from retarded first nests or re-nesting."; Page 16 —
"Following three dry years, superior cover in 1962 ma have made formerly submarginal
habitats suitable for turkeys. Thus improvement rather than deterioration of the habitat
may promote turkey emigration."; Page 18 — "Weather appeared to control the time of
movement from summer range."; Page 20 — "Apparently weather was the primary factor
determining the time of departure from the summer range.";Page 22 — "During periods
of snowy or windy weather, turkeys apparently roosted all day or restricted their
movements to 100 acres or less. Temperatures down to -10°F did not appear to restrict
the birds as they fed or loafed on the bare slopes. During periods of warm weather,
birds often moved up into the ponderosa pine forest even into one or two feet of snow
when the slopes below were comparatively bare. They moved down the slopes to
roost."; Page 23 — "During periods of snowy or windy weather, turkeys apparently
roosted all day or restricted their movements to 100 acres or less. Temperature down to
-10°F did not appear to restrict the birds as they fed or loafed on the bare slopes.
During periods of warm weather, birds often moved up into the ponderosa pine forest
even into one or two feet of snow when the slopes below were comparatively bare.
They moved down the slopes to roost."; Page 25 — "Percentages of the hatch occurring
June 1 or after were 17.1 percent, 45.0 percent and 45.2 per cent for 1961, 1962 and
1963 respectively. Percentages of juveniles retaining the eighth juvenile wing-primary
during the fall hunting season in 1 961 , 1 962 and 1 963 were 37. 1 per cent, 40.7 per cent
and 44.7 per cent respectively. This corroborates data in Table 17 indicating

% *">'<o**

109

% $fc <m> &

progressively later hatching dates for 1961, 1962 and 1963. This information suggests a
direct relationship to weather, especially precipitation, for April, May and June for those
years (Table 18). The year of 1961 was a dry "normal" year. Higher percentages of
hatches occurring later in the wetter 1962 and 1963 suggests delay in nesting or re-
nesting. Some of the disruption may have been due to cooler temperatures
accompanying the rain and snow. At the beginning of the peak hatching period May 21,
1963, a temperature of 18°F was recorded."; Page 29 — "Late fall rains in 1961 brought
about a 'greening up' of the vegetation and most of the crop contents that winter
consisted of green grass blades. Effects of above average rainfall the spring and
summer of 1962 were evident in crop contents the following winter."; Page 31 — " During
the record year for precipitation (1963), grasses and shrubs were so tall, even under the
usually heavy grazing, poults had difficulty flying from the ground when flushed."; Page
31 — "Low precipitation during winter months generally results in a light snow cover.
Relatively heavy rains and some cool temperatures in spring may delay nesting, but all
evidence indicates that this is no serious threat to productivity. Moisture on the
vegetation early in the morning is common in June. Apparently this is no particular
problem to poults 10 to 15 days old that have been seen feeding in moisture-soaked
vegetation on several occasions." Key words: Merriam's turkey, precipitation, snow,
temperature, wind, productivity, movement, vegetation, nutrition, behavior

241

Jorgensen, E. and A. S. Blix. 1988. Energy conservation by restricted body
cooling in cold-exposed willow ptarmigan chicks? Ornis Scandinavica 19(1):17-

20. Summary: Metabolic heat production (M) and body temperature (Tb) in relation to
ambient temperature (Ta) were measured in 72 7-9 day old willow ptarmigan chicks. M
increased linearly with falling Ta in the 6 to 35°C range and can be described by the
equation: M(Wkg"1) = -0.66 Ta(°C) + 37.4 (r= 0.792). Upon cold exposure, M rapidly
rose to a stable level, which was maintained in spite of a linear decrease in Tb. The drop
in Tb was similar at all Ta below 27°C when exposure lasted for only 10 minutes, while
the drop in Tb was significantly higher at 6°C compared with 21 °C when exposure lasted
for 30 minutes. These results indicate that M is chiefly influenced by peripheral thermal
inputs, and that willow ptarmigan chicks do not utilize their full thermoregulatory
capacity to maintain Tb. Apparently, they do instead use it to restrict body cooling,
whereby feeding time will be less dependent on Ta. Key words: willow ptarmigan,
thermoregulation, behavior, temperature

242

. and . 1985. Effects of climate and nutrition on growth and survival of

willow ptarmigan chicks. Ornis Scandinavia 16(2):99-107. Summary: Effects of
ambient temperature (Ta), and food availability and quality on growth and survival of
willow ptarmigan chicks Lagopus I. lagopus were investigated in the laboratory. Under
standard food conditions, growth increased with increasing Ta. Only a persisting sub-
zero Ta affected survival. Because of changes in the chicks' behavior a 67% reduction
of food availability did not affect their growth and survival. When food was available only
22% of the time, however, all chicks died within 5 days after hatching, irrespective of Ta.
Even small reductions in food quality strongly affected growth and survival. Food intake

110

4-

H<o>f*

#

O

was always correlated with body weight, irrespective of age, Ta, and food quality. These
results indicate that young ptarmigan chicks are unable to compensate for increased
energy expenditure and reduced food quality with increased food intake. Therefore the
availability of high quality food seems to be crucial for their growth and survival, while
low ambient temperatures perse seem to be of little consequence. Key words: willow
ptarmigan, temperature, behavior, nutrition, mortality

243

Jurgenson, P. B. 1968. A study of fluctuations of populations of the hazel grouse.
Pages 75-81 in Z. D. Spurin, chief editor. Birds of the Baltic region: ecology and
migrations. Israel Program for Scientific Translations [from Russian]. Notes: Page
80 - "Our main conclusion, based on the differences and similarities of the fluctuations
of populations of the Siberian and Manchurian hazel grouse in various parts of the
Northern Hemisphere, is that there are different environmental factors which have a
similar effect and act to a considerable extent simultaneously. Such factors which act
similarly over enormous areas can only be climatic factors." Key words: hazel grouse,
climate, population dynamics

244

Kabat, C, D. R. Thompson, and F. M. Kozlik. 1950. Pheasant weights and wing
molt in relation to reproduction with survival implications. Technical Wildlife
Bulletin Number 2, Pittman-Robertson Project 9-R, Wisconsin Conservation
Department, Madison, USA. Notes: Page 12 — Whatever the basic cause is that brings
on molt and the regaining of the weight lost during the breeding season, it appears to be
linked with the time the adult hen becomes broody, incubates and hatches her clutch.
Hence the occurrence of adverse weather or other factors which delay reproduction will
also set back the date of the post-nuptial molt and the post-breeding season weight
recovery. It is possible then that the survival of adult hens which are behind in physical
development, when fall and winter conditions prevail, could be handicapped
accordingly. The exact way in which delayed reproduction could affect the survival of
adult hens or their subsequent reproductive efforts is unknown, but probably it would be
an increased susceptibility to mortality factors of all types that is involved. Page 15 —
Prolonged egg laying due to nest destruction or other factors inhibiting the development
of broodiness constitutes an abnormal stress that may result in an increased
susceptibility to mortality causes of all types. Thus hens bringing off late broods or not
becoming broody may have a higher incidence of such diseases as the avian-leucosis
complex. . .Delay of the hatching date schedule by adverse weather or some other factor
may result in both the production of a relatively small crop of young birds and an
increase in the mortality rate of adult hen pheasants. Key words: ring-necked
pheasant, adverse weather, productivity, mortality, predation, disease, population
dynamics

245

Kaczor, N. W. 2008. Nesting and brood-rearing success and resource selection of
greater sage-grouse in northwestern South Dakota. M.S.Thesis, South Dakota
State University, Brookings, USA. Notes: The second-ranked model of nest success

** t* ^

(AICC weight = 0.15) included grass height, litter, daily precipitation, and a 1-day lag
effect of precipitation. Although daily precipitation had a positive influence on nest
success (B = 29.45 SE = 40.35), and the 1-day lag effect negatively influenced nest
success (8 = -1.89 SE = 0.77), neither variable improved the top model and were only
present due to being combined with grass height and litter. The third and fourth ranked
models included daily precipitation, and bird age, respectively, but they were also
combined with grass height and litter. Nest success varied 14.8% between years (37.7
± 7.3% in 2006 compared to 525 ± 7.2% in 2007). However, adding a year affect to the
top model did not improve model fit. Key words: sage-grouse, nesting success,
precipitation, model

246

Kane, D. F., R. O. Kimmel, and W. E. Faber. 2007. Winter survival of wild turkey
females in central Minnesota. Journal of Wildlife Management 71(6):1800-1807.

Abstract: There is interest in expanding eastern wild turkey {Meleagris gallopavo
silvestris) populations north of their current range. We hypothesized that winter survival
and food availability are primary determinants in setting the northern extent of wild
turkey distribution. To test our hypothesis, we translocated wild turkey females north of
their present range into central Minnesota, USA, and compared survival in areas with
supplemental food in the form of corn food plots versus areas with no supplemental
food. During 2 winters with below-average snow, winter survival was higher for females
with supplemental food. In one winter with above-average snow depths, survival was
extremely low even with supplemental food. Supplemental food could augment survival
during mild winters if wildlife managers arrange with farmers to, annually, retain
standing corn near roosting habitat, but food plots may only partially offset effects of
deep snow. Managers should critically evaluate northern habitats, long-term costs of
sustained feeding, and potential outcomes of concentrating animals and introducing
wild animals into new ecosystems. Winter survival may delimit the northern range of
wild turkeys, though annual survival rates may also be important and need further
research. Key words: turkey, snow, nutrition, distribution

247

Kasprzykowski, Z. 2002. Decline of the black grouse Tetrao tetrix population in
east-central Poland. Vogelwelt 123(5):253-258. Abstract: The study was conducted in
the Mazovian and South-Podlasian Lowlands, east-central Poland, covering ca. 13% of
the area of Poland. The data for 1975-2000 are based on the author's own observations
and material gathered by the regional card-index of the Mazovian Society for the
Protection of Fauna. In addition, in 1989-2000 population abundance was surveyed on
8 plots. In the mid-1980s, 96 leks were known, mostly from the eastern part of the
region. Over a period of 10 years (by 1985) leks disappeared from central and southern
parts of the Mazovian Lowland. Black grouse abandoned over 60% of the known
refuges. Early in the 1990s this process continued, and lekking grouse were recorded
mainly from the northern part of the region. Surveys in 1997-2000 revealed only 8 leks.
This means that over a period of 25 years, the number of leks decreased by 92%. A
detailed analysis of changes in numbers was possible for northern portion of the grouse
population. At the end of the 1980s, the number of birds varied between 1 and 6 on five

112 j^

^

> > k# H*

plots; they were absent from the remaining three plots. A comparison of numbers for
1990 and 1998 showed a decline from 101 to 17 individuals, that is by 83%, over a
period of 8 years. The number of males declined by more than 80% and that of females
by about 70%. The mean annual rate of decline was 10-13%. The number of individuals
per lek decreased with the total population. Early in the 1990s, groups of 8-20 (8.2 on
average) lekking males were noted, whereas in 1998, only four males were present on
the biggest lek, and only single birds were observed on the remaining leks (x = 1.8
male/lek). The most important factors causing this decline are assumed to be increasing
predation, mainly by foxes, hunting on leks and the weather conditions in winter (thin
snow cover). Co-occurrence of pheasants and changes in the natural habitat are
believed to be of less importance in the study region. Key words: black grouse, snow,
population dynamics

248

Kennamer, J. E., D. H. Arner, C. R. Hopkins, and R. C. Clanton. 1975. Productivity
of the eastern wild turkey in the Mississippi delta. Proceedings of the National
Wild Turkey Symposium 3:41-47. Abstract: Eastern wild turkey (Meleagris gallopavo
silvestris) productivity was determined during the summers of 1968-72 in the Mississippi
River Delta, Mississippi. Highest productivity occurred in 1969 and the lowest in 1970.
Production was not correlated with rainfall but it appeared to be influenced by extreme
river flood conditions. An inverse relationship was found between percentage of hens
with poults and total number of adults observed per field day during August. Key
Words: Eastern wild turkey, precipitation, flood, productivity

249

Keppie, D. M. 1977. Snow cover and the use of trees by spruce grouse in autumn.
Condor 79(3):382-384. Summary: I investigated whether snow in autumn affects the
tendency of spruce grouse (C. c. franklinii) to spend more time in trees. The significant
increases of birds in trees from August to September in 1971 and 1972 occurred only
after the first snowfall. Within months, birds were usually in trees more frequently on
days with greater snow cover than on days with lesser snow cover. In 15 of 19 cases,
proportionally more birds were in trees on days when a greater extent of snow was
present and differences were significant in 13 cases. The proportion of birds in trees on
snow-free days increased significantly between successive months in only two of five
possible instances. Spruce grouse were seen more frequently in trees as the autumn
progressed, and long before snow cover became permanent. The frequency of birds in
trees did not begin to increase until after the first snowfall. Grouse were found in trees
more often on days of major snow cover. The proportion of birds in trees on snow-free
days generally did not increase through autumn, although snow-free days became less
common as autumn progressed. This latter point suggests that after being in trees on a
day with much snow cover, grouse did not tend to remain in trees on a succeeding day
with less snow present. Pendergast and Boag (Condor 73:437-443, 1971) speculated
whether heavy mortality, particularly of young, might occur at the start of winter if
grouse were suddenly forced onto a conifer diet because of heavy snows, without
previous conditioning of the gastrointestinal tract to the new diet. This probably does not
occur, at least in regions with autumn snow. Data from Alberta suggest that periods of

f >f fc* **

i~- c*~*^-j^}> c b

temporary snow accustom the grouse to aboreal life. My initial interpretation of my data
was that snow causes birds to take to trees more frequently, enabling them to begin
heavy feeding on conifer browse, thus changing the size of the gastrointestinal tract.
With a delay in snowfall, the chain of events might be delayed, as well as in areas
without autumn snow. If this sequence does occur, ineffective dietary changeover might
cause mortality in regions that are snow-free in autumn. Key words: spruce grouse,
snow, habitat use, diet, mortality

250

and J. Towers. 1990. Using phenology to predict commencement of

nesting of female spruce grouse (Dendragapus canadensis). American Midland
Naturalist 124(1):164-170. Abstract: Dates for loss of snow cover and beginning of
plant growth were evaluated with respect to median dates for commencement of egg-
laying and hatch of cultures by female spruce grouse {Dendragapus canadensis) in
three widely separated localities in Canada. Female spruce grouse begin to lay eggs at
a time when environmental conditions are predictable as witnessed by low variation in
calendar dates among years in phonological evens. Among the events that we
monitored, the dates for first flowers of blueberry (Vacinium angustifolium) and trailing
arbutus (Epigaea repens) and the dates for 50% snow cover were correlated best with
median dates for commencement of egg-laying and hatch of clutches. Median nesting
dates can be predicted well by these events. There is now circumstantial evidence that
timing of plant development influences commencement times for egg-laying. Key
words: spruce grouse, vegetation, phenology, index, reproduction

251

Kerwin, M. L. 1971. The status, behavior, and ecology of the sage-grouse in
Saskatchewan. M.S. Thesis, University of Saskatchewan, Regina, Canada. Notes:
Kerwin concluded that unusually cold and wet conditions in the spring of 1970 reduced
nesting success, compared to the more normal temperatures and dry conditions of
1971. Key words: sage-grouse, temperature, precipitation, reproduction

252

Kessler, F. W. 1959. A study of the effects of weather on breeding behavior, nest
establishment, egg laying, and hatchability of the ring-necked pheasant.
Dissertation, Ohio State University, Akron, USA. Summary: Territorial behavior
occurred later in 1956 when the spring was colder and wetter. The clutch size was
smaller during wet breeding seasons. Incubating hens spent a greater amount of time
off the nest during the early and late stages of incubation than during the intermediate
period. Average egg temperatures were lower during 1956, the year with more rainfall.
Winds of 15 to 18 mph during the early part of the breeding season forced pheasants
into small areas of heavy cover with consequent strife among cocks. This strife resulted
in some of the territorial cocks becoming submissive and remaining so throughout the
season. At an ambient temperature of 70°F, winds of 10-20 mph acting for a period of
30 min. caused a differential egg temperature decrease of 2°-8° F between eggs in
exposed nests and eggs in nests sheltered from the wind. Data from this study
indicated that the duration of rainfall had no noticeable effect on incubation

114

H kr* **

±11'

<^>

temperatures. The amount of rainfall did not noticeably influence the egg temperatures
of the pheasants in this study. The number of periods of precipitation may under certain
conditions influence the number of times a hen leaves the nest. Key words: ring-
necked pheasant, temperature, precipitation, wind, habitat use, behavior,
reproduction

253

Kienzler, J. M., T. W. Little, and W. A. Fuller. 1995. Effects of weather, incubation
and hunting on gobbling activity in wild turkeys. Proceedings of the National Wild
Turkey Symposium 7:61-67. Abstract: The setting of spring wild turkey (Meleagris
gallopavo) hunting seasons has been influenced by tradition, gobbling, and hen
vulnerability. Knowing the peaks of gobbling and the beginning of incubation is
important in setting spring hunting seasons. We were interested in (1) determining the
effects of hunting and weather factors on gobbling activity, (2) quantifying daily and
seasonal trends in the intensity of gobbling activity, and (3) determining the relationship
of chronology of incubation and gobbling activity. Early morning gobbling activity was
monitored daily from mid-March through early June on two areas in south-central Iowa,
1978-81. Although no linear trend of gobbling activity and hunter density could be
detected (P = 0.87), the presence of hunters depressed gobbling counts (P<0.001).
Temperature and light intensity were also related to gobbling counts (P<0.01).
Precipitation the previous 12 hours and wind were inversely related to the counts
(P<0.01). Although gobbling activity was usually consistent between years, the
chronology of nesting did not appear to strictly coincide with gobbling every year. After
sunrise, within-day patterns of gobbling were similar before and during the hunting
season. Before the hunting season started, high average counts were relatively higher
prior to sunrise, however. Hunting depressed gobbling counts at all times of the day.
Hunting was estimated to be responsible for part of the late-April dip in gobbling activity
usually attributed to nesting. Key words: wild turkey, temperature, solar radiation,
wind, behavior, hunting

254

King, R. T. Ruffed grouse management. 1937. Journal of Forestry 35(6):523-532.

Notes: Winter cover for ruffed grouse must provide protection from extreme weather
conditions and roosting places safe from predators. In the northern part of their range,
snow ordinarily meets both of these requirements. Where snow occurs to a depth of
twelve inches or more and is not crusted the birds deliberately dive into it; they not only
roost under the snow at night, but we have found that they spend a great part of each
day under its protection when temperatures are near or below zero. On our grouse
ranges there is always a sufficient depth of snow to provide excellent protection, and it
is usually not crusted to the extent that the birds cannot utilize it. There are, however
years in which a sufficient depth of snow has not fallen by the time the deciduous trees
have lost their leaves; under these conditions, the birds are exposed to the effects of
low temperatures and bad storms during the winter which put a crust on the snow that
no bird can break through. This condition also forces them to roost in trees. At such
times when snow protection is not available the next best protection is provided by
clumps or fringes of balsam, spruce, or cedar. Pines do not furnish good winter cover

H ^ ^

-*

ii^9

except as young dense stands. At about 15 years of age they outgrow their usefulness
in this respect. If snow could be depended upon each winter all winter long, there would
be no need to worry about coniferous cover. Unfortunately it can not be depended upon
for the reasons just mentioned. It is, therefore, essential that coniferous cover be
provided, even though this cover is necessary only once in every several years. Much
of wildlife management is in the nature of insurance against future probable
contingencies. A two-day ice storm one in five years can do away with all the increase
built up during that period unless the management measures have taken account of and
provided for the ice storm. Key words: ruffed grouse, snow, temperature, ice,
vegetation, habitat use

255

Kirby, A. D., A. A. Smith, T. G. Benton, and P. J. Hudson. 2004. Rising burden of
immature sheep ticks {Ixodes ricinus) on red grouse (Lagopus lagopus scoticus)
chicks in the Scottish uplands. Medical and Veterinary Entomology 18:67-70.

Abstract: The sheep tick Ixodes ricinus (L.) (AcarUxodidae) is an ectoparasite of major
economic and pathogenic importance in Scotland. Its distribution in the Scottish uplands
is assumed to be governed by the abundance and distribution of its definitive hosts
(deer and sheep) and climatic variables such as temperature and rainfall. As the
numbers of its major host in Scotland, red deer, have increased dramatically and
climatic conditions have become more favourable, the level of parasitism could have
been expected to rise. We use data gathered from tick counts on over 4000 red grouse
chicks Lagopus lagopus scoticus Latham (Galliformes:Tetraonidae) in various
experiments over the past 19 years to ascertain whether the intensity and prevalence of
parasitism has been increasing. From 1985 to 2003 the average tick burden of a
parasitized red grouse chick has grown from 2.60±1.12 ticks per chick to 12.71l1.44.
Over this period the percentage of chicks of a given brood parasitized has also
increased from 4±2% to 92±3%. The possible implications of this increase in parasitism
for red grouse production are discussed. Key words: red grouse, temperature,
precipitation, disease, insects

256

Klaus, S. 2007. A 33-year study of hazel grouse Bonasa bonasia in the Bohemian
Forest, Sumava, Czech Republic: effects of weather on density in autumn.
Wildlife Biology 13 Supplement 1:105-108. Abstract: The only long-term census of
hazel grouse Bonasa bonasia in Central Europe was conducted during 1972-2004 on a
100-km2 area of the Bohemian Forest, Sumava, Czech Republic. To obtain a density
index in October, I recorded indirect signs of hazel grouse, such as dust bathing sites,
feathers, droppings or tracks, and reactions to a whistle that imitates the male territorial
song along fixed routes covering 80 km in total. During the 33 years of counting, I found
no statistically significant trend in the fluctuating numbers. I also looked for the influence
of weather on hazel grouse abundance. Analysis by stepwise multiple regression
suggested that six weather variables were correlated with the annual rate of change of
hazel grouse density in autumn, explaining 44% of the variation in density: mean
temperature and total precipitation in April (highly significant positive correlations) and
mean temperature and total precipitation in May and September (significant negative

116 ^

4>

»' kr* **

i/< ^

*<

correlations). The results for April and May seem to indicate a positive effect of rainfall
and temperature in April on reproductive success in the prelaying period, but a negative
effect of rainfall in May on chick survival. Key words: hazel grouse, temperature,
precipitation, density, reproduction

257

Klongan, E. D. 1971. Effects of some Iowa winters on pheasants. Pages 268-278
in: A. O. Haugen, editor. Snow and ice in relation to wildlife and recreation
symposium. Iowa Cooperative Wildlife Research Unit, Iowa State University,
Ames, USA. Abstract: Severe winter conditions are an important factor influencing
pheasant population levels in Iowa. Significant snowfall in combination with low
temperatures and strong winds exerts the greatest stress on the birds. Mortality usually
occurs in a short time from exposure and freezing complications, with starvation
exerting little or no importance. Effects of some of the most severe winters in Iowa
history on pheasants are described. Rapid changes in farming technology in northern
Iowa are resulting in limitations from lack of safe nesting cover to assume an equal, or
even greater, role than winter cover in determining pheasant numbers. Relationships
between location of food sources and winter cover are also becoming increasingly
important. Winter feeding programs would have done little or nothing to prevent the
types of winter pheasant losses described, though increasingly intensive farming may
have an effect here, too. Key words: ring-necked pheasant, snow, temperature,
wind, mortality, nutrition, habitat modification

258

Knapton, R. W. 1980. Winter mortality in a gray partridge population in Manitoba.
Canadian Field-Naturalist 94(2): 190-1 91. Notes: "The storm, the worst on record in
the area since March 1966, continued unabated from noon on 10 January 1975 until
noon on 12 January, and covered all of southern Manitoba and southeastern
Saskatchewan. Temperatures varied between -16 and -25°C, wind speeds from 55 to
65 km/h, with gusts to 100 km/h, and 16.5 cm of snow fell at Lyleton. The result was a
38% loss in total numbers of partridges, covey 2 being particularly hard hit as its
numbers declined from 7 to 2 birds. No further losses were noted for the duration of the
winter. The reduction in numbers was almost certainly a result of the storm, and not of
other factors such as emigration. ..thus, one storm was responsible for 70% of the total
over-winter loss of partridges on the study area." Key words: gray partridge, snow,
wind, temperature, mortality

259

Kobriger, J. 1989. Effects of weather conditions on sharp-tailed grouse brood
surveys. Prairie Grouse Technical Council Conference 18:14. Abstract: Sharp-tailed
grouse brood routes have been conducted in North Dakota since 1977. Weather
conditions during brood surveys were recorded. A numerical rating for weather was
devised using temperature, wind, cloud cover, and dew. Routes were graded from 1 to
12, 1 being very poor while 12 was excellent. A significant correlation was found
between both sharptails/100 miles (R2 = .95, P= <05) and sharptail broods/100 miles

)■* fc* **

4-

(R2 = .98, P = <01 ) and weather conditions. Key words: sharptail grouse,
temperature, wind, clouds, dew, humidity, census

260

Korhonen, K. 1989. Heat loss of willow grouse (Lagopus lagopus I.) in a snowy
environment. Journal of Thermal Biology 14:27-31. Abstract: 1. The amount of heat
lost by willow grouse (Lagopus I. lagopus L) in a snowy environment was measured
using a heat flow sensor. 2. During digging of the burrow, the heat flux density from
chest and feet increased to 1 100-1500 Wm 2 in only a few minutes. The heat loss from
the back was less (250-1 000 Wm'2). 3. As the burrow became warm, the heat flux
density decreased from the chest, back and legs to ca 1 15-130 Wm"2. 4. Digging in the
snow created a state of cold stress in the grouse, which was also expressed in
accelerated heartbeat and strong shivering at short intervals. 5. A snowy environment
increases heat loss from grouse walking on the surface or digging into the snow,
although snow also has the opposite, insulating, effect of efficiently protecting birds in
warmed, snow burrows from the cold. Key words: willow grouse, snow, temperature,
thermoregulation, behavior

261

. 1987. The effect of short-term fasting on certain blood parameters and on

glycogen storage in pectoral muscles of willow grouse Lagopus lagopus lagopus
L. Comparative Biochemistry and Physiology A 88(4):677-682. Abstract: (1) The
blood plasma glucose, cholesterol, total protein, urea, and uric acid contents of willow
grouse raised in a bird yard in northern Finland were measured after fasts of 12, 24,
and 48 hours in mild and severe cold weather. (2) There were significant differences
between the sexes in the urea and total protein contents of normal blood (12 hr fast). (3)
In all the blood parameters studied, changes occurred after 24 and 48 hour fasts, which
were greater or took place more rapidly in severe cold than in milder weather. Key
words: willow grouse, temperature, blood parameters, nutrition

262

Kothmann, H. G. and G. W. Litton. 1975. Utilization of man-made roosts by turkey
in west Texas. Proceedings of the National Wild Turkey Symposium 3:159-163.

Abstract: The Rio Grande turkey (Meleagris gallopavo intermedia) has extended its
range into 1.4 million hectares (ha) of semi-arid scrub mesquite prairie in west Texas. A
well established and increasing population exists where the southern edge of the High
Plains meets the Edwards Plateau and Trans-Pecos. Population fluctuations are
proportional to range conditions and rainfall patterns. Natural roost sites are absent, but
man-made structures such as utility line and poles, oil storage tanks, and windmill
towers provide adequate substitutes. Key words: Rio Grande turkey, precipitation,
habitat condition

263

Kozicky, E. L., G. O. Hendrickson, P. G. Homeyer, and R. Nomsen. 1955. Weather
and fall pheasant populations in Iowa. Journal of Wildlife Management 19(1):136-

118 j^

<l>

K fc* 1*

cc±

142. Summary: (1) An attempt was made to determine the relationship of the fall
roadside census from 1936 through 1952 within Iowa's primary pheasant range and
winter and spring mean temperatures and mean total precipitation. (2) The rate of
pheasant production was apparently related to changes in habitat and population level.
(3) Two months of continuous subnormal temperatures from December through
February apparently influenced the subsequent fall pheasant population. (4) Cold
temperatures in May and/or June were not conducive to an increase in fall pheasant
populations. (5) A warm March and/or April did not have any discernible benefits to
pheasant production over a normal and/or cold corresponding bimonthly period. In fact,
a warm March and April accompanied by a cold May, 1946, was followed by a decrease
in the fall pheasant population. (6) Above normal mean total precipitation in conjunction
with low temperatures were only evident in years of decreasing fall pheasant
populations; with high temperatures, precipitation had no apparent adverse effect on fall
pheasant populations. (7) Normal spring weather prevailed during years in which the fall
pheasant population remained the same or increased, depending to an undetermined
extent on the level of the pheasant population. (8) The present analysis merely
represents the weather-pheasant relationships for the past 17 years. Some
permutations of temperature and precipitation have been examined only once, and
many not at all. Key words: ring-necked pheasant, precipitation, temperature,
population dynamics

264

Krementz, D. G. and J. J. Jackson. 1999. Woodcock in the southeast. Natural
history and management for landowners. University of Georgia College of
Agricultural and Environmental Sciences, Bulletin No. 1183, Cooperative
Extension Service, Athens, USA. Notes: "Weather and Worms: Weather affects
worms. If surface soil is too cold or too hot, worms go deep. A soil temperature of 50° to
68°F is about right for most of the woodcock's favorite worms. During the southern
summer, the surface temperature is too hot for the woodcock's favorite worms. We
wonder if that is one reason why they migrate north to find better feeding. When surface
soil dries out, worms also go deep. Woodcock cannot get worms from hard, dry soil.
Nor can woodcock feed in soil that is too wet. Surface soil moisture of abut 20 percent
to 50 percent is best for woodcock and worms. Occasionally, widespread, extremely
cold winter temperatures in the South have caused soil to freeze and woodcock to
move. In most years, however, it is flooding on the winter grounds that causes
relocations of wintering woodcock. Their movements are generally local; at most, a few
miles. During these times, woodcock suffer much predation because the better covers
are often inundated with water and woodcock are forced into poor-quality cover. As
floods recede, the woodcock fly back to their old haunts. The worms survive the
flooding." Key words: American woodcock, temperature, precipitation, flooding,
movement, behavior, nutrition, predation

265

Labisky, R. F., C. A. Harper, and F. Greeley. 1964. Influence of land use, calcium,
and weather on the distribution and abundance of pheasants in Illinois. Biological

fV ^ ^

119

Notes No. 51, Illinois Natural History Survey, Urbana. Notes: Page 12 — Pheasants
and Weather:

266

Laperriere, A. J. 1972. Seasonal precipitation influence on mourning dove
breeding populations in Iowa. Journal of Wildlife Management 36(3):979-981.
Abstract: An inverse relationship was found between June-July rainfall in Iowa and
percentage of change in the Iowa mourning dove (Zenaidura macroura) call-count index
the following year. Indications are that unfavorable seasonal rainfall during the past
decade may be involved in the recent decline in breeding populations of mourning
doves in Iowa. Keywords: mourning dove, precipitation, fecundity

267

Larsen, J. A. 1957. Influence of weather upon the ruffed grouse population of the
Cloquet Experimental Forest, Minnesota. M. S. Thesis, University of Wisconsin-
Madison, Madison, USA. Unable to obtain for abstract.

268

and J. F. Lahey. 1958. Influence of weather upon a ruffed grouse

population. Journal of Wildlife Management 22(1):63-70. Summary: A statistical
technique developed by Fisher (1924) has been employed to demonstrate that about 50
per cent of the variability of the ruffed grouse population of the Cloquet Experimental
Forest in Minnesota during the years 1927-55 is associated with corresponding annual
changes in the distributional pattern of maximum temperature. The curve depicting the
effect of a degree above average of maximum temperature throughout the year shows
that warm days in spring and summer tend to be associated with a high grouse
population the following April, and that warm days during winter tend to be associated
with a low grouse population the following April. Key words: ruffed grouse,
population dynamics, temperature

269

Larson, M. A. 2005. Identifying plots for surveys of prairie chickens in Minnesota.
Pages 39-43 in Wingate, P. J., R. O. Kimmel, J. S. Lawrence, and M. S. Lenarz,
editors. Summaries of Wildlife Research Findings 2005. Minnesota Department of
Natural Resources, Fish and Wildlife Division, St. Paul, USA. Notes: To explore
potential improvements in surveys of greater prairie chickens (Tympanuchus cupido
pinnatus) in Minnesota, I developed this study to determine landscape-scale
characteristics associated with plots of land occupied by prairie chicken leks, and to
evaluate potential within-year sources of variation in the probability of detecting a prairie
chicken lek, if one is present. The study area consisted of nearly the entire range of
prairie chickens in northwest Minnesota... Wind speed and cloud cover were negatively
correlated with the probability of detecting a lek. Key words: greater prairie chicken,
wind, clouds, census

120 ^

^

H kr* **

<Ci

270

Latham, R. M. 1947. Differential ability of male and female game birds to
withstand starvation and climatic extremes. Journal of Wildlife Management
11(2):139-149. Summary: A series of experiments with six different species of game
birds, which included 1,332 individuals, pointed toward a significant difference between
the sexes in their ability to withstand fasting and climatic extremes. Among polygamous
species, the females proved the stronger; and among monogamous species, the
reverse was the case. Key words: Hungarian partridge, bobwhite quail, ring-neck
pheasant, wild turkey, nutrition, temperature, wind, mortality

271

. 1976. Complete book of the wild turkey. Stackpole Company, Harrisburg,

Pennsylvania USA. Notes: Page 78-79 — "When deep snows cover the food on the
ground, turkeys will regularly go to springs and spring runs to scratch for whatever food
may be available underwater. During this wading and feeding, ice may form on the wet
feathers of breast, belly and tail. This heavy coating of ice will often cause feathers to
pull or break off, leaving the bird without adequate protection against cold. These "bare
breasted" birds probably suffer considerably and perhaps die if exposed to long periods
of intense cold and high wind. Ice storms in themselves have been blamed for such
losses. When freezing rains cover all branches and twigs so that a twig the size of a
lead pencil becomes as large around as a hoe handle, the turkey roosting in the open
may become so weighted with ice, particularly on its tail feathers, that it is unable to fly
or even walk without great difficulty. At the very least, this has a tendency to weaken the
bird and make it less resistant to stress. Also, on the ground or on the roost it becomes
easy prey to its natural enemies." Key words: turkey, snow, ice, temperature,
precipitation, wind, nutrition, predation, mortality

272

Lauckhart, J. B. 1957. Animal cycles and food. Journal of Wildlife Management
21(2):230-234. Abstract: Some rodent fluctuations apparently can be accounted for by
population buildup, food depletion, starvation, and recovery of food base. Malnutrition
from use of poor foods is commoner than starvation. Malnutrition may lead to disease,
low reproduction, and less survival of young, all of which appear in cyclic declines.
There is evidence that buds and twigs of northern trees, food supply of various
herbivores such as grouse, are very low in nutrients, so that a slight change could
cause malnutrition. A cyclic change that does affect northern trees is the seed cycle. It
seems that trees in the north must store food for an average of about 3.5 years before
they can produce seed crop. In this period their twigs and buds apparently become
more nutritious, but become suddenly less so after seed formation. This might explain
cycles of bud and twig eaters, and why these animals do not peak the same years as
seed eaters. Interacting seed cycles could cause longer animal cycles. Weather can
prevent a seed crop in any one year and thus lengthen a cycle. Key words: grouse,
vegetation, weather, nutrition, disease, fecundity

* v k* <*"

% »*"

•>*''

cCi

273

Lehman, C. P. 2005. Ecology of Merriam's turkeys in the southern Black Hills,
South Dakota. Dissertation, South Dakota State University, Brookings, USA.

Notes: "Successful nests were on steeper slopes, had greater visual obstruction, and
greater total ground level vegetation and shrub cover than unsuccessful nests.
Precipitation during the incubation period was also an important variable predicting nest
success. Survival of poults <14 days posthatch decreased during cold periods and
during or immediately following periods of rainfall. Monitoring rainfall and temperature
data during May and June may provide an index of annual nest success and poult
recruitment Keywords: Merriam's turkey, precipitation, temperature, index,
reproduction

274

, M. A. Rumble, L. D. Flake, and D. J. Thompson. 2008. Merriam's turkey

nest survival and factors affecting nest predation by mammals. Journal of
Wildlife Management 72(8):1765-1744. Abstract: Nest success is an important
parameter affecting population fluctuations of wild turkeys (Meleagris gallopavo).
Factors influencing mammalian predation on turkey nests are complicated and not well
understood. Therefore, we assessed nest hazard risk by testing competing hypotheses
of Merriam's turkey (M g. merriami) nest survival in a ponderosa pine (Pinus
ponderosa) ecosystem during 2001-2003. We collected nesting information on 83
female Merriam's turkeys; annual nest success averaged 50% for adult females (range
= 49-59%) and 83% for yearling females (range = 75-100%). Proportional hazard
modeling indicated that precipitation increased the hazard of nest mortality. However,
estimated hazard of nest predation was lowered when incubating females had greater
shrub cover and visual obstruction around nests. Coyotes (Canis latrans) were the
primary predator on turkey nests. We hypothesize that precipitation is the best predictor
of nest survival for first nests because coyotes use olfaction effectively to find nesting
females during wet periods. Temporally, as the nesting season progressed,
precipitation declined and vegetation cover increased and coyotes may have more
difficulty detecting nests under these conditions later in the nesting period. The
interaction of concealment cover with precipitation indicated that nest hazard risk from
daily precipitation was reduced with greater shrub cover. Management activities that
promote greater shrub cover may partially offset the negative effects of greater
precipitation events. Key words: Merriam's turkey, precipitation, predation,
vegetation, habitat management

275

, L. D. Flake, M. A. Rumble, and D. J. Thompson. 2008. Merriam's turkey

poult survival in the Black Hills, South Dakota. Intermountain Journal of Sciences
14(4):78-88. Abstract: We investigated poult survival from hatching to 4 wks of age for
Merriam's wild turkey {Meleagris gallopavo merriami) poults in the southern Black Hills,
South Dakota, We estimated survival from 841 poults reared by 57 radio-marked wild
turkeys (r»=52 adult females, n=5 yearling females). Survival of poults to 4 wks
posthatch averaged 33 percent with 54 percent of the mortality occurring in the first 7
days after hatching. Merriam's turkey poult survival in the southern Black Hills was low

122 ^

Nk

K ^ >^

c^>

compared to Merriam's populations found elsewhere in the entire current range.
Survival of poults increased with age, fewer precipitation events, and fewer extreme
cold and wet events. The interaction of age of poults with cold and wet events through
15 days posthatch indicated that younger poults were more susceptible to cold and wet
weather events than older-aged poults. We observed several poults <3 days of age that
apparently died from hypothermia. A fine-scale based weather index that uses
individual weather stations for specific areas occupied by turkeys may be a valuable
tool for managers to estimate production in Merriam's turkeys if survey or radio
telemetry data are not available. Key words: Merriam's turkey, precipitation, chick
survival, index

276

Lehman, V. W. 1953. Bobwhite population fluctuations and Vitamin A.
Transactions of the North American Wildlife Conference 18:199-246. Summary and
Conclusions: "In this paper the results of analysis of 270 quail livers for Vitamin A are
considered against a background of field data on population fluctuations, breeding, food
habits, parasites, cover, and intra-specific tolerance on southwestern rangeland. The
study period (1949-1951) included both "normal" times and the unusually cold winter of
1950-1951 which followed a "dry" summer and fall. Winter mortality was heavy, and
Vitamin A reserves in the time of crisis were lower (average 132 I.U.'s per gram of liver
and 236 I.U.'s per liver) than at any other time. Averages of Vitamin A reserves per
gram of liver and per liver seemingly identified periods of greatest abundance (as
September, 1949) and scarcity (March, 1951). The Vitamin A reserves of individual
birds, however, showed a wide range of variation at all seasons. Under adverse
conditions, the ability to manufacture and store Vitamin A may be influenced by sex and
age. As crisis approached in 1950, old males, old females, and young-of-the-year
appeared able to maintain Vitamin A reserves, and life itself, in the order named.
Fluctuations in average Vitamin A reserves, one collection period to the next, were
sometimes coincidental with changes in diet. Rises in average Vitamin A levels were
recorded with increased consumption of weed seeds or mast. Reductions in Vitamin A
levels were sometimes concurrent with high consumption of grass seed, prickly pear
fruit, and insects (principally termites). Marked Vitamin A improvement on diets
importantly "greens" occurred only after the material was largely buds, flowers, and
dough-stage seeds as contrasted to emergent sprouts and leaves There was not a
significant correlation between body weight and Vitamin A reserves. At the time of
winter die-off, low Vitamin A reserves (40 I.U.'s or less per liver) occurred in 21 per cent
of the study specimens weighing over 150 grams and in 25 percent of the quail
weighing less than 150 grams. Symptoms of distress, i.e., ruffled plumage, rhinitis,
decreased alertness, weak flight, heavy parasite infestations, etc., were not confined to
the birds of light weight. It did not appear that high Vitamin A reserves were necessary
for awakening the breeding urge. General breeding effort, however, was definitely
associated with lush range and its frequent accompaniment of above-average feed.
Quail, especially females, quickly attained high Vitamin A reserves after generous
rainfall, and most successful breeding occurred after heavy May rains in both 1949 and
1950. With verdant range of short duration in 1951, survival of young was only 1.3 per
adult as compared to 2.6 per adult in 1950, and 3.9 per adult in 1949." Key words:

^ 123

N^

^v^**

^ tj^\ ^^ ^

bobwhite quail, precipitation, vegetation, nutrition, population dynamics,
mortality

277

. 1949. Bobwhite quail reproduction in southwestern Texas. Journal of

Wildlife Management 10(2):111-123. Summary: Bobwhite renesting was influenced
importantly by rainfall. Hatchability was high (about 94%) in spring, but decreased
(71%) as the weather warmed. Successful renestings produced only about half as many
chicks as spring layings. Important factors regulating quail reproduction in southwestern
Texas are rainfall, through its effect on nesting activity, and coyote predation, because
of its severity except in mid-summer. Quail are limited by overgrazing on many ranches.
Keywords: bobwhite quail, precipitation, productivity, renesting, behavior,
grazing

278

Lehtinen, S. A. 1983. Movements and habitat use of ruffed grouse in the Bridger
Mountains, Montana. M. S. Thesis, Montana State University, Bozeman, USA.

Notes: Page 85 — Ruffed grouse preferred coniferous forests for tree roosting. No
changes in habitat selection were observed between day and night by tree-roosting
grouse. The preference for dense coniferous cover as tree roosting sites could be
expected, since it offered better protection from inclement winter weather and predators
than the other habitats. Page 86 — Topographic Preference - In winter the aspects most
often associated with birds were south and west, while southeast, north, and southwest
were all avoided. Hungerford found ruffed grouse on the upper parts of south-facing
slopes during winter in Idaho. In Stauffer's study, higher elevations were preferred
during winter. Grouse were always associated more with south and east aspects,
although in summer and fall they could be found across all aspects. Slopes used were
generally less than 20°, which he considered as gentle terrain. Both of the above
authors believed that the higher hillsides were warmer than areas below. This seems to
be a likely explanation for the preference of hillsides. The solar radiation on those
aspects that face the sun for the longest part of the day would cause snow to melt
during sunny days, even in mid-winter, uncovering more shrubs for feeding grouse. The
density of fruit-bearing plants might also be higher on south-facing slopes than on those
facing north. Key words: ruffed grouse, snow, solar radiation, temperature,
behavior, habitat use, diet, vegetation

279

Lennerstedt, 1. 1966. Egg temperature and incubation rhythm of a capercaillie
(Tetrao urogallus L.) in Swedish Lapland. Oikos 17:169-174. Abstract: A thermistor
was mounted in one of the eggs in a capercaillie's nest, and the remaining space of the
egg was filled with paraffin. Automatic registration of the electrical resistance every sixth
minute throughout the last twelve days of the incubation period made calculation of the
egg temperature possible. During incubation the temperature showed only minor
variations, usually within the limits 34-35°C. The minimum egg temperature during
absence was on average 27.7°C, the lowest recorded value being 21 °C. During three

124 j^

>->" fc* >(=*

J/„

c£^

days the weather conditions were severe with snow-fall and the average of the
minimum egg temperatures was 2.4°C lower than that for the other days. As long as the
female behaves in a normal way. i.e. the length of the absent periods are within the
normal limits, even severe weather conditions do not seem to affect the embryonic
development. The diurnal rhythm of incubation and absence was calculated from the
temperature values. There was no concentration of periods of absence to any particular
part of the day except five hours in the afternoon, 18.00-23.00, when there always
occurred a period of uninterrupted incubation. These five hours obviously represent a
period of rest in the diurnal rhythm of certain activities such as feeding. Key words:
capercaillie, temperature, reproduction, behavior

280

Leopold, A. 1937. The effect of the winter of 1935-36 on Wisconsin quail.
American Midland Naturalist 18(3):408-416. Summary: (1) The winter of 1935-36
greatly constricted the range and population density of Wisconsin quail. Mortality on
measured samples ranged from 30 to 83 per cent. (2) The lethal effect is ascribed to 40
days of continuous cold and snow. The previous winter of 1934-35, as measured by a
"hardness test," was only 15 percent as severe; 1917-18, 72 per cent. (3) Fat well-fed
quail died in numbers, a phenomenon hitherto unrecorded. Some of these had ice
under the wings. Many were imprisoned in snow. (4) Attempts to develop a size-index
from body measurements, and an emaciation-index from weight:length ratios, were
unsuccessful. Key words: quail, snow, temperature, ice, mortality, nutrition

281

Leopold, A. S. 1977. The California quail. University of California Press, Berkeley,
USA. Notes: In Chapter 9, "Rainfall as a factor affecting reproductive success", Leopold
provides a synthesis of research on the direct and indirect impacts of precipitation upon
the reproduction and population dynamics of California quail. The timing of rain and
temperature fluctuations as well as the type of vegetation promoted in the environment
exert influences on fecundity, survival, and recruitment. Key words: California quail,
precipitation, temperature, vegetation, nutrition, reproduction, population
dynamics

282

Leopold, B. D., L. A. Brennan, W. Rosene, and G. A Hurst. 1993. Long-term trends
of northern bobwhite populations in the southeastern U.S.: the role of abiotic
factors. Proceedings of the National Quail Symposium 3:185. Abstract: We
assessed the potential influence of precipitation variation and drought-severity on long-
term trends of northern bobwhite population indexes using data derived from the
Christmas Bird Count (1961-89) in the southeastern U.S., and harvest data (number of
bobwhite bagged per unit effort) from Groton Plantation (1957-89) and Oakland Hunting
Club (1927-87) in South Carolina. We calculated long-term yearly drought-severity
indices to simultaneously scale precipitation, average temperature, water holding
capacities of soil, and evapo-transpiration, and used these data as independent
variables in regression analyses of long-term bobwhite population indices. Drought-
severity indices were correlated (P < 0.5) with long-term bobwhite population trends and

X Hfc*>f*

125

<^>

explained approximately 50% of the year-to-year variation in population changes.
Variation in population indices not explained by drought-severity indexes is apparently
the result of biotic factors associated with changes in land use. Key words: bobwhite
quail, precipitation, drought, temperature, evapotranspiration, soil, index

283

Leptich, D. J. 1992. Winter habitat use by hen pheasants in southern Idaho.
Journal of Wildlife Management 56(2):376-380. Notes: Wildlife managers should
promote a change in management direction to one that encourages tall, erect, perennial
herbaceous and, where it does not interfere with farming or maintenance, woody
vegetation on these sites. Furthermore, because this cover type may be flattened or
filled with snow and become unusable during severe winter weather, it is important that
habitat managers provide a good interspersion of other preferred cover types
throughout the agricultural ecosystem. Remnant tracts of sagebrush provide pheasants
with important winter loafing and escape cover. Habitat managers should work to
ensure a good distribution of sagebrush tracts up to 1 km wide across the agricultural
ecosystem and provide firebreaks or green stripping to protect them from fire. Key
words: ring-necked pheasant, vegetation, habitat management, snow

284

Lewis, J. C. 1963. Observations on the winter range of wild turkeys in Michigan.
Journal of Wildlife Management 27(1):98-102. Abstract: The daily home range of 21
flocks of 12 wild turkeys {Meleagris gallapavo) was studied in Allegan County, Michigan,
during the winter of 1957-58. The area of the daily home range seemed to vary
inversely with snow depth and directly with temperature; however, positive relationships
were somewhat obscured by other influencing factors. Twenty-nine measurements of
the area contained within the limits of the daily movements of turkey flocks varied from
2 to 160 acres and averaged 49 acres. The home range used throughout the winter
averaged 683 acres for 8 flocks of gobblers only, 435 acres for 7 flocks of hens only,
and 492 acres for 6 flocks of mixed sexes. Flocks remained individually identifiable
throughout the winter either by flock composition or the discreteness of flock ranges.
The range of 86 percent of 132 turkeys bordered streams or lakes, and 74 percent of 19
winter roosts were found there. It was hypothesized that an efficient and economical
census of turkeys in winter could be based on a search along streams and borders of
lakes for turkeys and for their signs in snow. Key words: turkey, snow, temperature,
census

285

, J. W. Ault, III, V. J. Heller, and J. A. Morrison. 1976. Delayed molt of

primary feathers of mourning doves during winter. Journal of Wildlife
Management 40(1 ):1 84-1 87. Notes: "Presumably, severe winter weather and the
associated food shortages were factors slowing the molt of immature mourning doves in
Oklahoma. The winter of 1972-73 was the more severe of the study period and the one
having the higher percentage of the population showing indications of a prolonged molt.
Also during January 1973, a heavy mortality of doves coincided with 6 days of low
temperatures and snow and ice covering food supplies. Severe weather and food

126 ^

^

)-V ^ ^

shortages may magnify the delay in molting patterns, but they may not be the only
factors responsible for the delay in Oklahoma. Biologists should be cautious about
estimating hatching dates of doves by using birds harvested or trapped in late fall and
winter." Key words: mourning dove, snow, temperature, nutrition, behavior,
technique

286

Lewis R. A. and M. A. Degner. 1982. Mortality of nestling blue grouse owing to
inclement weather. Canadian Field-Naturalist 96(2):218-219. Notes: "Young blue
grouse usually leave the nest within a day of hatching, and our finding indicates that
inclement weather can cause mortality during this period. Ours was the only instance
[on Vancouver or Hardwicke Island] in which nestlings died as a result of adverse
weather. Therefore, in most cases, cool, wet weather does not affect survival of nestling
blue grouse in coastal British Columbia." "Although the direct cause of death was
almost certainly cool, wet weather, poor cover at the nest site may have contributed
indirectly to the mortality." Key words: blue grouse, precipitation, wind,
temperature, mortality, reproduction, severe weather

287

Ligon, J. S. 1946. History and management of Merriam's wild turkey. New Mexico
Game and Fish Commission. University of New Mexico Press, Albuquerque, USA.

Notes: Page 43 - "Cold rains are seriously detrimental to young turkeys, because of the
direct effects of wet and chill; furthermore, if the storms are too extended or continuous,
the loss of feeding time results in weakening of the young, which renders them less
resistant." "...dense spruce and other conifers are a most important element of turkey-
nesting habitat in high country in that they furnish shelter for hens and young during
frequent rain or sleet squalls after the rainy season begins. Spruce gives the best
protection (from rain, sleet, and snow), as the branches, in addition to being heavily
foliaged from top to bottom, extend out in more or less flat fans, providing a maximum of
protection. However, continued cold and blowing rains may be seriously destructive to
young turkeys, regardless of all natural protective cover." Key words: Merriam's
turkey, precipitation, temperature, sleet, snow, wind, nutrition, habitat use

288

Lindstrdm, J. 1996. Weather and grouse population dynamics. Wildlife Biology
2(2):93-99. Abstract: One paradigm in the biology of game animals is that short-term
fluctuations in population densities can be explained with variations in weather. A
number of empirical models have been produced supporting this view. However,
validation of such models has often been lacking or insufficient. Two methods for
checking the validity of such models are presented. The first method is to derive a
model for one population and test it against another data set. The second method is to
evaluate the forecasting power of the model. For these purposes, the relation between
36 weather variables and population parameters of capercaillie Tetrao urogallus, black
grouse T. tetrix, and hazel grouse Bonasa bonasia was studied using population data
(1964-1984) from three adjacent provinces in Finland. Total population size, number of
juveniles and population growth rate were used as dependent variables. Prior to the

^ l& +*

^
4

% m <& 9

analyses, the population data were In-transformed and detrended. Stepwise regression
analysis was used with province-specific weather data as explanatory variables. These
models were then used to make forecasts one year ahead for each species and
province, and the prediction was tested against observed data. Transferred models
from the other provinces were also used. The requirement for a good empirical model is
that it should be possible to use the model on similar problems in nearby areas.
Stepwise regression analyses yielded reasonable fits in most cases (hivin R-2 ranging
between 0.2-2.0). However, a model from a given province invariably produced a poor
fit when applied to another province. Forecasting the population dynamics was only
occasionally successful, and was not directly related to the fit of the models. The results
suggest that it may often be hazardous to use weather data for predicting population
fluctuations of game species, especially for management purposes. This conclusion
was further strengthened by demonstrating that using 36 province-specific white noise
variables, it was possible to build models with fit and forecasting properties essentially
equal to those of the weather-based models. Key words: capercaillie, black grouse,
hazel grouse, model, index

289

, E. Ranta, and H. Linden. 1996. Large-scale synchrony in the dynamics of

capercaillie, black grouse and hazel grouse populations in Finland. Oikos 76:221-
227. Abstract: We analyzed 1964-1983 data on population numbers of capercaillie,
black grouse and hazel grouse in 1 1 provinces in Finland. The three species are known
to display cyclic dynamics with a periodicity of approximately six years. Synchrony in
species-specific population fluctuations was scored by calculating cross correlations
with zero lag among all the provinces. Provincial data were also used to assess the
relation between climatic factors and synchrony in the population fluctuations. To
enable the statistical analysis, the number of the 12 climatic variables was reduced to
three non-correlating principal components. A bootstrap method was used both for
parameter estimation and for assessing statistical significance level when comparing
the tetraonid synchrony with the climate data. For all three species we found large-scale
synchrony in population fluctuations. The level of synchrony decreased, however, with
geographic distance among the provinces. When a partial correlation analysis is done
correcting for the weather-derived principal components, correlations with synchrony
and distance remained with two of the three principal components. When corrected for
distance, there is correlation between synchrony and climate for capercaillie and black
grouse. We conclude that spatial phenomena, such as climatic homogeneity within
closely located areas and dispersal of individuals are responsible for maintaining large-
scale synchrony. Key words: capercaillie, black grouse, hazel grouse, temperature,
snow, precipitation, population dynamics

290

Long, C. R. 2007. Pre-breeding food habits and condition of ruffed grouse and
effects on reproduction in the central and southern Appalachians. M.S. Thesis,
West Virginia University, Morgantown, USA. Notes: Grouse were collected
throughout the northern states and it is likely that the lack of snow and warming
temperatures enabled grouse in some areas to forage on herbaceous vegetation more

128 jj

4>

>-> ^, ^

c£^

easily. Key words: ruffed grouse, temperature, snow, behavior, nutrition,
reproduction

291

Loneux, M. 2003. Are climate changes the cause of the decline of the black
grouse? Levende Natuur 104(3):104-107. Abstract: Population data and
meteorological records performed for several decades have allowed us to model the
climates influence on black grouse (Tetrao tetrix) population dynamics. Results
obtained in several European protected areas where the species breeds still nowadays
have shown the important role of some climatic parameters during the life cycle of the
bird to explain the observed fluctuations. The common factors involved in the modeling
for the different geographical sites studied in West-Europe suggest that global climate
warming can explain the declining trend of the species. On one hand the winters are
more and more rainy and less snowy due to the warming of the minimal temperature,
while black grouse is adapted to rough snowy conditions. On the other hand, rainfalls
are more and more abundant during key times of this species brooding period. However
the new climatic trend cannot fully explain the general decline of black grouse observed
even in protected areas in Western Europe. The species is suffering from the global
climate change in addition to attacks on its habitat and tranquility. At short term and
local level decisions and actions of managers to improve and maintain the quality and
carrying capacity of the typical habitats used by the species during its whole life cycle
will play major role for the short term survival of this emblematic bird in the west
positions of its distribution. On longer term, the future of the small isolated populations
of the species will depend on its ability to face the changing environment. Key words:
black grouse, temperature, precipitation, population dynamics, climate change

292

, J. Lindsey, and J. C. Ruwet. 1997. Influence of the climate on the dynamics

of the black grouse (Tetrao tetrix) population in the Belgian "Hautes-Fagnes"
from 1967 to 1996. Cahiers d'Ethologie 17(2-4):345-386. Abstract: This study
involves demographic data on the varying population size of black grouse living in the
Hautes-Fagnes obtained during yearly spring censuses on their arenas. These are
related to data on local meteorological conditions having a known influence on the
survival of these birds during the various periods of the life cycle of the species. The
data were modeled by Poisson multiple regression using the GLIM4 software. The
dynamics studied cover a period of thirty years of censuses (1967-1996). The
predictions from the best statistical model follow the observed values exceptionally well.
The variables best explaining the changes in population size of the black grouse in the
Hautes-Fagnes are, in addition to the mother population: the climate during the previous
two winters as measured by the mean of the minimum temperatures from 1 November
to 31 March (negative effect); the climate close to the period of hatching, measured by
the mean minimum temperature during the three weeks starting on 16 June (positive
effect) and the total precipitation during the three weeks starting on 1 June (negative
effect); the climate during incubation, measured by the total precipitation during four
weeks starting on the 19 and 25 May (negative and positive effects, respectively); and
the climate during September, measured by the total precipitation during this month

i f*r^^

129

o^ £}g^ ^^> <£>

(negative effect). All of these data involve the year preceding that being studied. Black
grouse benefit from cold winters, standing badly the mild winters. This relationship is
explained by the low level of their winter metabolism, adapted to a winter life based on
long rests in igloos and short periods looking for food of low energy value. Winter acts
as an elimination trial for the young grouse who do not yet have the adult weight. Later,
this is also a crucial period of the life cycle of adults. Our results unquestionably confirm
this for the population in Hautes-Fagnes. Black grouse benefit from a warm dry summer
climate during the periods covering the first weeks of life of the chicks. For the total
precipitation over the four weeks covering the incubation period, a subsequent study will
look at shorter periods to attempt to determine at what date the effect is reversed. A
rainy September is bad for the grouse. When the predictions are far from the observed
values, other variables than the climatic ones used in the model must dominate. The
model predicts well the direction of the variation observed, but diverges more or less
from the exact value in five of the 30 years, in one direction or the other (1980, 1983,
1986, 1990, 1996). It is reassuring that this climatic model does not agree perfectly with
the observations because that would mean that all other factors are negligible,
contradicting many publications on the subject. If it seems reasonable to ignore the
effects of illness in favor of factors involving "quality of habitat" or "disturbances", the
factor, "predation", thought to be negligible in a variable population, may have had more
impact the last four years because of the higher density of foxes (an opportunistic
predator) and the low density of grouse (the lowest point since 1967). For the poorly
predicted years, we should look for events or phenomena that could have produced
effects, positive or negative: fires, specific management activities or lack of
management, direct or indirect disturbances by walkers or their dogs, skiers,
photographers. This type of information requires archives regularly fed by the facts
noticed by foresters, wardens, naturalists as well as local inhabitants who are often in
the field. Key words: black grouse, temperature, snow, precipitation, climate
change, population dynamics, model

293

Longcore, J. R., D. G. McAuley, G. F. Sepik, and G. W. Pendleton. 2000. Survival
of female American woodcock breeding in Maine. Proceedings of the American
Woodcock Symposium 9:65-76. Notes: Excluding birds that died from entanglement,
only 2 of 7 (28%) females (1 each in 1988 and 1989) died within 16 days of being radio-
marked, which is within the period of most severe weather in spring. Page 71 —
Females, because of larger body size and associated heat dynamics are better able to
withstand sub-zero temperatures and sparse foods in spring than are males. No
females starved during this study, but two males did. In Maine in 1989, when the
minimum, mean daily temperature in April was the lowest (-2.5°C) among years. Page
72-... "In 1986 when the mean, minimum daily temperature in April was the highest
among years, survival was the lowest, but based on a sample of 1 1 birds. Although
deep frost caused by lack of snow and sub-zero temperatures in early winter and spring
snowstorms can limit food availability and cause females to delay nesting, survival of
females in 1989 was not adversely affected. This high survival rate may have been
related to reduced nesting effort. Furthermore, the energetic advantages of larger body
size and not engaging in energetically costly display flights, as do males, probably
contributes to higher survival rates for females than for males, especially in years with

130 jj

>•>' k* a*

inclement spring weather." Key words: American woodcock, temperature, snow,
frost, mortality, nutrition, behavior

294

, , , and . 1997. Survival of breeding male American

woodcock in Maine. Canadian Journal of Zoology 74:2046-2054. Abstract: We
radio-marked 150 male American woodcock (Scolopax minor) during 1987-1989 and
estimated period survival for 1 April - 15 June. Survival varied from 0.690 (1989) to
0.924 (1988), with a 3-year mean (95% confidence interval) of 0.789 (0.693-0.885).
Woodcock were killed by raptors (r?=14, 53.8%), mammals (n=1, 3.8%), or unknown
predators (n=5, 19.2%); six deaths (23.1%) were from miscellaneous causes, including
three (11. 5%) from entanglement in the transmitter harness. A composite survival
estimate based on telemetry studies for the breeding, postbreeding, and wintering
periods was 0.471 (0.789 x 0.923 x 0.647). The calculated survival rates were 0.881 for
the spring migration period and 0.853 for the combined hunting and fall migration
period. In a proportional hazards model, body mass at capture was not related to
survival. Forest type (hardwood versus conifers) affected survival (P<0. 01 6), which was
lower for woodcock using mostly conifer sites. Survival was related positively to mean
snow depth in December (P<0.038), negatively to snow depth in April (P<0.046), and
positively to minimum temperature in December (P<0.054) and April (P<0.066) in some
analyses. Key words: American woodcock, snow, temperature, habitat use,
mortality

295

Lowery, D. K. 1999. Relationships among wild turkey hens, predators, and
environmental conditions on Tallahala Wildlife Management Area, Mississippi
(Meleagris gallopavo). M. S. Thesis, Mississippi State University, Starkville, USA.

Abstract: Wild turkey (Meleagris gallopavo) hen nest success, survival, and recess
movements (time away from nest) were studied on Tallahala Wildlife Management Area
in central Mississippi, 1991-1995. Additionally, microhabitat and weather variables were
collected at nest sites and were compared between successful and unsuccessful nests.
Nesting rate and nesting success averaged 42% and 19%, respectively. Major cause of
nest failure was predation (87%, n = 27). Hen survival rates differed significantly among
years and averaged 50.5%. Predation was the major cause of hen mortality accounting
for 50% of all known mortalities. Hen recess movements (time away from nest) did not
differ in duration or frequency between successful and unsuccessful hens. However,
successful hens had more days without any recess movement than unsuccessful hens
(P = 0.017). Successful hens had significantly less cumulative rainfall (P = 0.002) and
fewer rainfall events (P =0.002) during incubation. Annual nesting success was
significantly correlated (r = 0.90, P = 0.03) with the number of rainfall events from
March-June of each year. Known dates of predation of hens (n = 19) had significantly
fewer days since rain (P = 0.004) and more rain the day of predation (P = 0.003) than
random dates during the same period. It rained on or just prior to the date of predation.
Micro-habitat variables for nest sites were similar to what has been reported.
Percentage grass at ground level and percentage canopy closure were significantly
different between successful and unsuccessful hens. Percentage grass averaged

% &(&**

131

% ?©>> ^ <£>

56.13% for successful and 24.58% for unsuccessful nests (P = 0.003). Canopy closure
averaged 62.45% for successful and 73.45% for unsuccessful (P = 0.043). Other
variables were similar for both groups. Management implications were also made. Key
words: wild turkey, precipitation, microhabitat, reproduction, vegetation

296

Lu, X. and G. M. Zheng. 2001. Habitat selection and use by a hybrid of white and
Tibetan eared pheasants in eastern Tibet during the post-incubation period.
Canadian Journal of Zoology 79:319-324. Abstract: We investigated habitat selection
and use by a recently discovered hybrid of the white eared pheasant (Crossoptilon
crossoptilon) and Tibetan eared pheasant (Crossoptilon harmani) in the forests of
eastern Tibet (93°39'E, 32°24'N) during the post-incubation period in 1995. The
frequency of encountering molted feathers was used as an indicator of the relative
abundance of eared pheasants in order to analyze patterns of habitat selection and use.
Forests on south-facing slopes, dominated by the holly leaf-like oak {Quercus
aquifolioides) and Tibetan juniper (Sabina tibetica), were the habitats preferred by eared
pheasants. North-facing slopes with coniferous forest, which is the most preferred
habitat of eared pheasant species in other areas, were completely avoided, probably
because moisture-heat conditions that are beyond the birds' physiological tolerance.
We conclude that climatic conditions are the main determinant of macrohabitat selection
by eared pheasant species. In preferred habitats, oak and juniper woodland accounted
for a larger proportion of home ranges of family flocks. Daily movements of a flock might
cover a large altitudinal range, from the base of the mountain to the area above tree
line, with an apparent preference for sites that can be used for foraging and dusting.
Key words: Tibetan eared pheasant, white-eared pheasant, temperature,
humidity, habitat use, behavior

297

Ludwig, G. X., R. V. Alatalo, P. Helle, H. Linden, J. Lindstrom, and H. Siitari. 2006.
Short- and long-term population dynamical consequences of asymmetric climate
change in black grouse. Proceedings Royal Society B 273:2009-2016. Abstract:
Temporal asymmetry in patterns of regional climate change may jeopardize the match
between the proximate and ultimate cues of the timing of breeding. The consequences
on short- and long-term population dynamics and trends as well as the underlying
mechanisms are, however, often unknown. Using long-term data from Finland, we
demonstrate that black grouse (Tetrao tetrix) have responded to spring warming by
advancing both egg-laying and hatching. However, early summer (the time of hatching)
has not advanced, and chicks have to face colder post-hatching conditions.
Demonstrating that these conditions are critical to post-hatching survival, we show that
chicks are increasingly suffering higher mortality because they hatch too early.
Consequently, breeding success and population size has severely declined over the
past four decades. Finally, we modeled the impact of this particular climate change
scenario on population dynamics and show that the mismatch can further explain the
observed collapse of cyclic fluctuations. Because the evolutionary response of grouse is
lagging behind the novel selective pressures, seasonally asymmetric climate change is
likely to constitute an important determinant of future short- and long-term changes in

132 ^

H fee ^

ccb

the dynamics of black grouse populations. Key words: black grouse, population
dynamics, temperature, mortality, breeding success

298

Lusk, J. J., F. S. Guthery, R. R. George, M. J. Peterson, and S. J. DeMaso. 2002.
Relative abundance of bobwhites in relation to weather and land use. Journal of
Wildlife Management 66(4):1040-1051. Abstract: Weather and land use are important
factors influencing the population dynamics of northern bobwhites {Colinus virginianus)
in Texas and elsewhere. Using an artificial neural network, we studied the effects of
these factors on an index of bobwhite abundance (hereafter, index) in 6 ecoregions in
Texas. We used roadside-count data collected by the Texas Parks and Wildlife
Department (TPWD) during 1978-1997. Weather variables were June, July, and August
mean maximum temperatures, and winter (Dec-Feb), spring (Mar-May), summer (Jun-
Aug), and fall (Sep-Nov) rainfall. We also included the proportion of county area in
cultivation, the number of livestock per hectare of noncultivated land, and the previous
year's bobwhite count in the analyses. The data were partitioned into training and
validation data sets prior to analyses. The neural model explained 65% of the variation
in the training data (n = 72) and 61% of the variation in the validation data (n = 17). The
most important variables contributing to network predictions were July temperature, fall
rainfall, cattle density, and the previous year's bobwhite count. State-level simulation
results indicated that the bobwhite index decreased with increasing June temperature
and livestock density. The bobwhite index increased with July and August temperature,
fall rainfall, and the previous year's bobwhite count. Bobwhite abundance increased
with the proportion of county area in cultivation up to approximately 20% cultivation and
then declined. Winter, spring, and summer rainfall had little effect on the bobwhite
index. Although many relationships appeared approximately linear or were decelerating,
proportion of county area in cultivation and livestock density on noncultivated land
showed strongly curvilinear responses. Therefore, cultivation up to approximately 20%
of county area was beneficial, but the benefits disappeared as cultivation increased
beyond this level. Further, at low livestock densities, between 0.15 and 0.40 head/ha,
small increases in head/ha resulted in a decrease in the bobwhite index of
156.4%/head/ha. The results also indicated that a potential bias might exist in the
survey protocol resulting in artificially inflated counts under some weather conditions.
Key words: bobwhite quail, temperature, precipitation, census, abundance, index

299

, , and S. J. DeMaso. 2001. Northern bobwhite {Colinus virginianus)

abundance in relation to yearly weather and long-term climate patterns.
Ecological Modeling 146(1-3):3-15. Abstract: We used multilayered, backpropagation
neural network to investigate the relative effects of yearly weather and long-term climate
patterns on the abundance of northern bobwhites (Colinus virginianus; hereafter,
bobwhite) in Oklahoma, USA. Bobwhite populations have been declining for several
decades across the United States, and predicted global climate change might
accelerate the rate of decline. We were interested in whether bobwhite abundance was
more responsive to yearly precipitation and temperature, or to annual deviations from
long-term mean climate patterns. We used roadside count data collected over a 6-year

y A>* / . . .. l- 133

% >*((?***

^ $jfe <Q>

*

period (1991-1997) by the Oklahoma Department of Wildlife Conservation as a
measure of bobwhite abundance. We standardized quail counts among counties by
calculating the standard normal deviate for each county. Weather data were obtained
from weather stations closest to the roadside-count route. We had 280 training cases
and 68 test-validation cases. Two data sets were constructed; one using yearly weather
data (actual rainfall and temperature) and the second using annual deviations from
long-term mean values. We conducted simulation analyses to determine the nature of
the relationship between each dependent variable and the standardized bobwhite
counts. A neural network with eight neurons was most efficient for the yearly weather
data, counting for 25% of the variation in the training data. The adjusted sum-of-squares
for this model was 2.42. A four-neuron network was selected for the deviation-from-
normal data set, accounting for 23% of the variation in the training data. The adjusted
sum-of-squares for the deviation model was 1.44, indicating it performed better than the
model for yearly weather patterns. Deviation from long-term mean July and August
temperatures combined contributed 31.5% to the climate network's predictions, and
deviations from mean winter, spring, and summer precipitation combined contributed
42.8% to the network's predictions. As July temperature increased over the long-term
mean, the number of bobwhites counted increased over the route mean, but the
relationship decelerated at high July temperatures. Predicted increases in bobwhites
counted were highest when August temperatures were below the mean and decreased
rapidly for all temperatures greater than the mean. Predicted bobwhite counts increased
asymptotically as winter rain increased over the long-term mean, but were greatest at
mean spring-rainfall amounts and at below average amounts of summer rainfall. We
conclude that the absolute changes in yearly weather pattern predicted by some global
change models will not have as great an impact on bobwhite abundance as will the
magnitude of the deviations of these values from the climate bobwhites are adapted to
in this portion of their range. Keywords: bobwhite quail, temperature, precipitation,
population dynamics, model, climate change

300

McAdoo, J. K. and G. N. Back. n.d. Sage-grouse habitat requirements. Fact Sheet
FS-01.-44. University of Nevada, Cooperative Extension, Reno, USA. Notes: Winter
Habitats - Winter habitat for sage-grouse varies according to weather conditions. But
regardless of weather, the presence of sagebrush for food and cover is the common
denominator. Sage-grouse feed almost exclusively on sagebrush leaves at this time of
year. At least 10 to 12 inches of sagebrush must be exposed above the snow to allow
feeding by sage-grouse. Although big sagebrush dominates sage-grouse diets in most
portions of this bird's range, other sagebrush species are also eaten. In northern
Nevada, low sagebrush is used until it is covered with snow, at which time the birds
move to big sagebrush areas. Sagebrush is also important in winter for cover. Sage-
grouse will roost in open, low sagebrush sites on clear, calm nights above 10°F.
However, on windy nights or during snowstorms the birds seek out taller shrubs in
areas with 20% or more canopy cover. On cold nights when powdery snow is available,
sage-grouse will burrow in the snow to conserve energy. These birds will fly more than
5 miles between winter-feeding and snow roosting sites during very cold weather (below
10°F)... sage-grouse use other habitats, like wet meadow areas, and increase their

134 j^

4>

>>' k> **

#

c£^

dependence on these habitats in mid to late summer during drought years. Key words:
sage-grouse, snow, temperature, wind, habitat use, vegetation, drought

301

McAtee, W. L., editor. 1945. The ring-necked pheasant and its management in
North America. American Wildlife Institute, Washington, D.C., USA. Summary: This
book presents the status of pheasants in various regions of the U.S. as follows:
Northeast: In Vermont, towns having the least annual snowfall have the most
pheasants. Occasional 3- to 4-inch snows and sleet storms that may coat the cover for
days, or subzero spells down to -30°F, alone do not ordinarily restrict pheasant
increase. As a rule, if good food and cover are available, the birds can be counted upon
to hold their own against the elements in the northeast without unreasonable losses.
Excessive snowfall, followed by heavy crusting may seal in some of the birds.
Prolonged sleet storms, and protracted subzero weather will eventually take a toll.
These conditions have a cumulative effect, gradually sapping the vitality of the birds,
and making them more susceptible to disease and predation. Ice storms, aside from
depriving the birds of food, may cause ice to form under their wings or freeze their tail
feathers to the ground. Extreme winter conditions conceivably can result in catastrophe
for the pheasant populations. Spring freshets, or other floods, cause nest losses in
some locations. Some swales are avoided or only partly utilized (for roosting) because
air drainage from slopes above makes them frost pockets, several degrees colder than
surrounding areas. Prevailing winter winds may drift in much of the swale with deep
snow for more or less extensive periods. Other part-time roosts are flooded in winter
thaws. Ohio: Deep snows and sleet sometimes make grain, weed seeds and other
foods relatively unavailable to pheasants. Such conditions, prolonged, may result in
starvation. Rain sometimes floods and destroys pheasant nests in low places. However,
a rainy period may delay the average date of hay mowing for a week or more at the
height of the pheasant nesting season and, in so doing, permit thousands of eggs to
hatch that would otherwise be destroyed. In winter pheasants commonly frequent
drainage ditches where they find shelter from the wind. In the thick growths of cattails
and sedges found in the bottoms of many of the ditches, there are often hollows and
channels through the snow. These are utilized by pheasants in severe winter weather.
Michigan: There is a lag in the time of day when the birds leave their roosting sites in
search of food and a hastening of their return to the roosts, both of which are
associated with lower temperature and diminished light intensity. High wind velocity
leads them to search persistently for windbreaks, such as the protected sides of trees,
heavy brush cover, lower spots protected by sedges, and weedy places. It is generally
believed that pheasants seek marshlands largely for cover, but the temperature factor
may also be an important secondary influence for this choice, for it has been
demonstrated that the winter temperature may be 19° higher among the marsh sedges
than in the grass and mixed herbaceous cover in more elevated locations. Cover is
valuable not merely in providing shelter from enemies or from adverse climatic
conditions, but in giving a sense of security that may conserve nervous and muscular
energy which would otherwise be expended in unnecessary skulking from dangers that
do not exist. Concerted crowing and fighting of cocks appear to be affected by
temperature, being conspicuous during warm periods and almost lacking during cold
spells. The pheasant's movements off of the roost are retarded by decreased

% H'^>^

135

Cgj ^ ^ Q>

temperature, especially during periods of strong winds. When temperatures remain
below 10° to 15°F, pheasants usually do not leave marsh cover to feed in nearby
cornfields, but take materials available within the margins of the marsh, which may
provide only a starvation diet. On very cold days, pheasants linger near or within the
roosting sites, and frequently do not leave the roost at all. Northern prairie states:
Prairie winds, while covering much ground with drifts, leave large spaces exposed, so
this bird is unlikely to go hungry. It can endure fairly protracted fasting in cold weather
and quickly recover lost weight when food is again available. There is evidence that
sheer severity of the winter climate alone may delimit pheasant populations on the
northern edge of their prairie range. In northern Iowa a severe winter caused a large
die-off. A large part of the loss... was attributed to freezing and choking. Birds, caught in
drift storms and blizzards away from dense escape cover, almost invariably turn their
tails to the wind and crouched on the snow. The body feathers of such unfortunate
pheasants were ruffed and the driven snow was packed under the feathers. Body heat
melted the snow and the severe cold caused the water to freeze and thus encase the
birds in ice. Many froze to death. Survival was highest in the flocks that roosted in
dense cover of willows and groves adjacent to an available food supply that required
little ranging to obtain. Nebraska: Weather affects carrying capacity of sandhill habitat.
Dry years cut down on the winter food supply. Extremely wet years also are detrimental
as the marshes are filled with water, freeze over as soon as winter arrives, and exclude
pheasants from normally important feeding grounds. Winter kill of the birds is always
very severe during periods of heavy snowfall on areas that have a limited food supply.
Heavy snows hardened by winds and subzero temperatures prevents the birds from
getting to the ground for food, and winter kill occurs within 15 days after such weather
begins. Pheasants also become inactive at subzero temperatures, do less foraging, and
may soon succumb to freezing. Intermountain Irrigated Region: There is no question
that near the surface of the (irrigated) ground and among cultivated plants the
temperature and particularly the humidity have been markedly affected. Winter snow
conditions appear to be especially important, with light to moderate snowfall favoring
high populations. Snow conditions are apparently a more serious factor than differences
in elevation and winter temperature. It was evident that the pheasants became more
dependent upon food other than waste grain as the snow cover increased (they ate
seeds and fruits of native plants). Pacific Northwest: After a snowfall of 3 feet, that
remained for about 10 days, numerous kills were made on pheasants by hawks and
other predators both in cover and in the open. Rains apparently have little effect upon
adult ring-necked pheasants. It is undesirable to take inventories during the first fall
rains, as the pheasants then remain in seclusion for a week or 10 days. After that period
of adjustment, they ignore the rain and carry on normally. Weather... definitely affects
survival of newly-hatched and immature birds. Birds hatched during periods of
extremely wet or cold weather do not survive. Wet, chilly weather during the hatching
period is directly reflected in a lessened number reaching maturity. Key words: ring-
necked pheasant, precipitation, snow, ice, wind, temperature, nutrition, predation,
mortality, behavior, habitat use

302

McAuley, D. G., J. R. Longcore, D. A. Clugston, W. H. Teman, and G. F. Sepik.
2006. Survival of American woodcock broods and chicks in Maine. In 10th

136 jj

4

>^^^

cCi

American Woodcock Symposium:25, Michican Department of Natural Resources
and the United States Fish and Wildlife Service. Abstract: During 1986-1989, 89
female American woodcock (Scolopax minor) were radio-marked during the period 1
April-30 June at Moosehom National Wildlife Refuge in Maine. Forty-six broods made
up of 190 chicks were followed for a 21 -day period to determine survival. Brood
survival, the probability of fledging >1 chick, during the 21-day period ranged from 0.339
to 1.000. Survival of chicks varied from 0.142 to 0.944. Survival rates differed among
years. Preliminary estimates indicate that survival of chicks and broods from after
second year (ASY) females differed from second year (SY) females. Differences
between original nests and re-nests and effects of weather were investigated. Key
words: American woodcock, weather, reproduction, mortality

303

McCabe, K. F. and L. D. Flake. 1985. Brood rearing habitat use by wild turkey
hens in south-central South Dakota. Proceedings of the National Wild Turkey
Symposium 5:121-131. Notes: Page 127 — "Adverse weather may have been a factor
in the low recruitment. During both years of the study, late snowfalls may have caused
poor initial nesting success because of abandonment. Also, prolonged rains and cool
weather occurred in June of both years, when any initial hatch of poults was most
vulnerable to adverse conditions. ..".Page 128 — "South-facing slopes also provide the
benefit of early morning sunlight, useful in burning off dew quickly, which reduces the
hazard of poults being dampened and chilled. South-facing woodlands likely were
selected over south-facing grasslands because the 48% average canopy cover could
provide protection from aerial predators, shelter against rain and wind while giving
ready access to open drying areas wherever trees are sparse, and shade from heat
(which also may concentrate insects)." Key words: Merriam's turkey, snow,
temperature, precipitation, productivity, solar radiation, dew, habitat use,
vegetation

304

McCabe, R. A. and A. S. Hawkins. 1946. The Hungarian partridge in Wisconsin.
American Midland Naturalist 36(1):1-75. Notes: Page 9 — "The true effect of climate is
often obscured by averages. Adverse climatic conditions become lethal in three ways,
suddenness, severity, and duration, all of which may be absorbed in averages. Climatic
conditions that in themselves would have no effect on wildlife may be combined to
produce a climatic complex that is detrimental. For example, rain or temperatures just
below freezing are of no importance separately, but a temperature drop to below
freezing during a rain may cause serious food shortage by coating both food and grit
with ice."; "Weather Mortality — The partridge is a hardy bird and it takes a combination
of adverse weather conditions such as snow, wind, and prolonged low temperatures to
bring about mortality. Two such combinations occurred during the winters of 1935-36
and 1 936-37. . .the first winter brought very deep snow and continued sub-zero
temperatures, and many partridges died. In January of the second winter, rain followed
by freezing temperatures covered southeastern Wisconsin with a sheet of ice which
remained for almost two months. Food and grit were difficult to obtain, and again many
partridges perished. It was also during this winter that loss through egress took place.

^

i f>'^^

137

<C^ oJ&_ ££}, c^

Roosting habits vary somewhat with weather conditions. On mild winter nights the
partridge covey may roost as a loose group, but on cold nights roost in a ring as
compact as that of the bobwhite. On one occasion... a covey of partridges roosted in
grouse fashion by plunging into a snowdrift. Feeding habits, like roosting, conform to
weather changes. When snow covers a field, they often tunnel, in one case to a depth
of twelve inches, to reach the feed." Key words: Hungarian partridge, snow, wind,
ice, precipitation, temperature, mortality, behavior, nutrition

305

McClure, H. E. 1942. Mourning dove production in southwestern Iowa. Auk
59(1):64-75. Notes: Page 66 - "No new nests were built after September 17, 1938, and
after September 24, 1939. Although a storm of August 30 served only to truncate the
September peak, a three-day cold rain and wind from September 13 through
September 1 7 brought a halt to nest building in 1 938. Hot winds from September 4 to
15, 1939, stopped nest building except for three unsuccessful attempts made after
September 15. Continued warm and calm weather after these disruptive spells failed in
both years to stimulate the birds to more nesting, although some were seen courting.
Captive birds showed a cessation of activity at this time, too. Wind and heavy storms
are the greatest decimating factors of nests, eggs and young. During 1938 there were
but few severe storms, while in 1939 the season was a series of blasts. On June 7 a
hailstorm occurred during which hailstones weighting as much as one-half pound fell.
Nests were not only knocked out by wind and rain, but parents and young were killed on
the nests by direct blows." Key words: mourning dove, precipitation, temperature,
wind, hail, behavior, mortality

306

McCrow, V. P. 1982. Gray partridge habitat use and nesting biology in north-
central Iowa. Dissertation, Iowa State University, Ames, USA. Abstract: Gray
partridge (Perdix perdix) movements, habitat use, and nesting were studied from 1975
to 1977 in north-central Iowa, a region of intensive row-crop agriculture. Use of 4 habitat
classes by 9 birds monitored by radio telemetry was compared with habitat availability
over 4 seasonal periods. Strip cover was used at higher than expected frequencies in
all periods. Partridge utilized the periphery of fields at a relatively high frequency while
use of the central portion was less than expected. Three partridge broods utilized
soybeans at a high frequency and corn at a relatively low frequency through the 1st 4
weeks of life, but use of corn increased from 5 to 8 weeks of age. Hay and strip cover
were used at a relatively high frequency through 2 weeks of age as were the edges of
fields. Average overall activity range for 6 mated birds that were monitored over 4 or
more months was 1.93 km('2) (range = 0.84 to 3.66 km('2)). Paired partridge occupied
relatively restricted ranges from the pre-nesting through the nesting period, but activity
ranges increased in the late summer. An unmated male followed a different pattern of
range size distribution with extensive wandering in the spring and a large activity range
followed by a general reduction in range size through the summer. Birds occupying
habitat with relatively high interspersion of cover types tended to have smaller overall
activity ranges. Over all years, 23 partridge nests and 29 ring-necked pheasant
(Phasianus colchicus) nests were found. Rates of nest success were 0.26 for partridge

138 jj

4

H to, >«*

±1*'

£^

and 0.41 for pheasants. Mean clutch size for 9 completed partridge nests was 14.9
eggs. In contrast to pheasants, partridge made little use of drainage ditchbank, railroad
righty-of-way, or grass waterway cover for nesting; 83% of the partridge nests were
found in roadside and fenceline cover. Only 2 pheasant nests and no partridge nests
were found in hayfields. Spring-to-fall gain was more closely associated with an index to
pair success than the mean brood size. Low pair success was affected by weather, re-
nesting effort, and the scarcity of stable and protected nesting cover. Protection of
roadside nesting cover from disturbance should benefit partridge populations in
northern Iowa. Keywords: gray partridge, ring-necked pheasant, weather,
productivity, vegetation, habitat use

307

McGowan, J. D. 1969. Starvation of Alaskan ruffed and sharp-tailed grouse
caused by icing. Auk 86(1):142-143. Summary: In 1968 thick accumulations of ice cut
off food supplies and roosting sites of sharp-tailed and ruffed grouse. A die-off resulted.
Exposed vegetation near Minto, Alaska was coated with about !4 inch of ice which
persisted for 5 weeks. Dead grouse were first noticed about 4 weeks after the heavy
glaze occurred. Less severe icing occurring near Fairbanks did not appear to affect
local ruffed grouse. Return to very low temperatures following glazing could make the
effect of icing more severe in interior Alaska than in temperate areas, especially if
glazing caused heavy crusting of snow, and thus prevented snow roosting. Key words:
ruffed grouse, sharp-tailed grouse, ice, temperature, nutrition, mortality

308

Mackenzie, J. M. D. 1952. Fluctuations in the numbers of British tetraonids.
Journal of Animal Ecology 21(1):128-153. Abstract: Data on red grouse, Lagopus
scoticus, are given for some 80 moors and the oldest records go back to 1834. Peaks
are shown, and some records of disease outbreaks back to 1797 are noted. Data on
other tetraonids, Tetrao urogallus, Lyrurus textrix, Lagopus mutus, from other estates in
central Scotland from 1866 to 1949 are given. L. scoticus fluctuates throughout its
range, the period being irregular, between 3 and 10 years but mostly 5 or 6. There is a
general synchrony which is not however exact. Peak years may differ even on adjoining
moors. It is suggested that this synchrony is brought about because, while crashes on
each moor are due to the interaction between the birds and their immediate
environment, there is occasionally a very good or very bad year which brings big areas
into line. The critical density may differ in the same area in different periods and varies
widely from place to place. Where density is high, strongylosis (infestation by the
nematode, Trichostrongylus pergracilis) is often but not always associated with crashes.
Cycles with much the same rhythm and dates occur in areas where it is unknown, when
bad nesting seasons are thought to be the limiting factor. Some of the worst crashes
occur when they are associated with strongylosis. A possible explanation is that once
the critical density is exceeded or resources are reduced below requirements (perhaps
only temporarily) the particular agency which reduces numbers may vary. The other
three tetraonids show rises and falls more or less synchronously with grouse. Duplicate
peaks sometimes occur at intervals of 2 years only. Species other than grouse
sometimes have interpolated peaks in long intervals. The only common factor likely to

)'V ^ Nf*

-* IJW i

Igf ?<§fe- ^ ^

cause these events is food, which is affected by weather. Peak periods of 2 or 3 years
are suggested, during which most peaks occur. Blackgame bags from south Scotland
from 1850 and 1949 are given and agree in general with data from central Scotland. An
increase in warmth and wetness in the northern climate may be the ultimate cause of
the long-term variations. Key words: black grouse, red grouse, temperature,
precipitation, nutrition, disease, mortality, population dynamics

309

McLean, R. G. 2006. West Nile Virus in North American birds. Ornithological
Monographs 60:44-64, American Ornithologists' Union, USA. Notes: Page 50 - "In
contrast to epidemic conditions in the western United States, WNV activity in areas east
of the Rocky Mountains was reported to be significantly less in 2004 than in 2003, likely
because of unfavorable weather for WNV transmission. Climatic data for 2004 showed
a significantly cooler and wetter summer, particularly compared with 2003, in the
eastern region of the United States, whereas weather in the western region — including
Arizona and California -- where there was increased WNV activity — did not change from
the previous five years. A combination of a wet and cool summer along with other
factors can greatly reduce mosquito production and activity, lengthen the extrinsic
incubation period of the virus in the vector, and affect reproduction and populations of
insect-eating birds, all of which could reduce WNV transmission and lower the number
of infected birds, equines, and humans in the temperate areas of the eastern United
States. However, reduced WNV transmission in southeastern states such as Florida in
2004 was attributable to summer drought." Page 53 - "Mortality of greater sage-grouse
(Centrocercus urophasianus, a declining and threatened species in western North
America) caused by WNV was documented in free-ranging populations in Montana,
Wyoming, and Alberta in 2003. Of 22 radiomarked females from four study sites that
could be tested, 18 (82%) died from WNV infection. In addition, serum collected from
112 greater sage-grouse from those areas after the outbreaks were all antibody-
negative, which suggests a low survival rate following WNV infection. Experimental
studies confirmed the high susceptibility and mortality in greater sage-grouse from WNV
infection: 100% mortality in second-year birds, with a 3.7-day mean survival time." Page
59 — "Climate and seasonal weather affect the winter survival, spring initiation, summer
intensity and diversity, and the intensity and distribution of local or regional mosquito-
borne virus activity. Climate appears to have less effect on winter survival and early-
spring initiation of WNV than of other mosquito-borne viruses." Key words: sage-
grouse, precipitation, temperature, drought, insects, disease, mortality

310

MacMillan, 1. 1. 1964. Annual population changes in California quail. Journal of
Wildlife Management 28(4):702-711. Abstract. A population of California quail
(Lophortyx californicus) in San Luis Obispo County, California, was kept under study
and management from 1946 to 1962. Observations and age ratios obtained during fall
hunting have indicated wide fluctuation of reproductive success, age ratios varying from
6-81 percent young. In general, years of high rainfall correlated with high reproduction;
dry winters were followed by reduced breeding. The key factor in reproductive rate was
the drive to keep nesting and renesting throughout the summer, resulting presumably

140 ^

4*

Hr t* f*

cCb

from a physiological preconditioning whose nature and origin is unknown. Reproductive
drive was shown to be independent of environmental conditions during the actual
nesting season, appearing to be related more to early breeding season weather
conditions that control annual plant growth and production of quail food. Key words:
California quail, precipitation, fecundity, nutrition

311

MacMullan, R. A. and L. L. Eberhardt. 1953. Tolerance of incubating pheasant
eggs to exposure. Journal of Wildlife Management 17(3):322-330. Summary:
Incubating pheasant eggs were subjected to low temperatures and simulated rain for
various lengths of time. From the tolerance data thus obtained, an equation has been
set up to approximate the relationship between temperature, stage of incubation, and
length of exposure as affecting survival. Conclusions: 1) As exposure temperature is
lowered, length of time pheasant embryos can safely tolerate exposure decreases. 2)
Embryos are progressively more vulnerable to exposure as incubation progresses. 3)
Alternate cooling and incubating do not cause higher mortality than a similar elapsed
time of continuous cooling. 4) Exposure to simulated rainfall or flooding reduces the
survival threshold below that of simple cooling to approximately the same temperatures.
5) Death from chilling evidently occurs during the exposure. "After-effects" are of minor
importance. 6) Newly-hatched chicks exposed to lowered temperatures are much more
vulnerable than eggs in any stage of incubation. The tolerance shown by pheasant eggs
in these experiments was considerably greater than popularly supposed. For much of
the incubation period, tolerance is great enough that widespread mortality from
unseasonal cold spells would not likely occur in the wild. There are two possibilities for
widespread mortality of chicks or eggs in the wild, however, that warrant further
consideration and study: 1) Changes of habits of hens during cold, wet weather could
affect survival of eggs in the wild. At 'normal' spring temperatures, hens apparently do
not remain off the nest long enough to allow lethal chilling. A change in habits of hens
during cold weather at late stages of incubation, when eggs are more vulnerable,
however, could allow lethal exposure. 2) Chicks appeared markedly less tolerant to cold
than eggs in any stage of incubation. While the experiment with chicks was limited, it
suggested that chicks are highly dependent upon the brooding hen for warmth.
Conceivably, severe cold, especially in conjunction with precipitation, could cause
widespread mortality, if it occurred at a time when a large proportion of the chick
population was vulnerable. Key words: ring-necked pheasant, temperature,
precipitation, flood, behavior, mortality, behavior

312

McNeil, J., H. Yin, and R. McNeil. 2007. Analysis of the climate in the greater sage-
grouse range of southwestern Saskatchewan and southeastern Alberta. Report
prepared for Prairie Farm Rehabilitation Administration, Agriculture and Agri-
Food Canada; Grasslands National Park, Parks Canada; Saskatchewan
Watershed Authority; and the Interdepartment Recovery Fund. LandWise Inc.,
Lethbridge, Alberta, Canada. Notes: This report discussed the potential effects of cold
wet weather and hot dry weather on sage-grouse. Page 36 "Correlations between
extreme weather and sage-grouse decline" states: "Sage-grouse numbers based on lek

% f>rfc*>f*

141

. I /^

counts declined dramatically after the spring of 1988, corresponding to the exceptionally
hot and dry conditions or drought in 1987 and 1988. The greater frequency of extreme
weather events in May and June in the 1990s and beyond may have contributed to the
continued but more gradual decline in sage-grouse populations since 1989. For
example, the recent population decrease may be in part related to the greater frequency
of cold and wet spring conditions in the study area between 1999 and 2004." Page 40:
"The exceptionally hot and dry conditions in 1988 were followed by a dramatic decline in
sage-grouse counts in the following spring. This severe decline may be directly related
to the extremely hot and dry weather in 1989. Insecticide use to control high
grasshopper populations associated with drought conditions in the mid to late 1980s
may also have contributed to the decline. Some insecticides used in Saskatchewan at
that time are highly toxic to birds, and the decrease in insect populations may also have
had a detrimental effect, particularly on sage-grouse chicks. The exceptionally hot and
dry conditions that culminated in 1988 were followed by several years with exceptionally
cold and wet springs, including 1991, and in particular, 1999, 2002, 2003 and 2004. The
recent cold and we conditions may have contributed to the more gradual decline in
sage-grouse numbers since 1998." Key words: sage-grouse, drought, precipitation,
temperature, pesticide, mortality, population dynamics, index

313

Manzer, D. L. and S. J. Hannon. 2007. Survival of sharp-tailed grouse
Tympanuchus phasianellus chicks and hens in a fragmented prairie landscape.
Wildlife Biology 14:1:16-25. Notes: The authors state death from exposure appeared
to be associated with periods of heavy precipitation. They speculate that heavy losses
in a single year presumably could occur if a prolonged period of wet and cold weather
persisted (2-3 days) through the early stages of chick growth before chicks are able to
thermoregulate. Key words: sharp-tailed grouse, precipitation, temperature,
mortality

314

Marcstrom, V. and N. H. Hoglund. 1980. Factors affecting reproduction of willow
grouse Lagopus lagopus in two highland areas of Sweden. Swedish Wildlife
Research Viltrevy 11(7):285-314. Abstract: Reproductive success in willow grouse is
investigated in relation to weather conditions during the chick period, time of breeding,
population fluctuations in small mammals and other factors. Unfavorable weather,
especially sleet and snow, had a drastic effect on survival of young chicks. Less than
30% young willow grouse in the autumn population could always be referred to poor
posthatching weather. Successful reproduction was more usual in early than in late
springs. An apparent correlation often existed between small mammal numbers and
reproductive success in willow grouse, although some obvious exceptions occurred.
The timing of small mammal declines was most likely of importance for the number of
predators present in spring to feed upon eggs and young of grouse. The situation could
be complicated by the fact that all small mammal species did not always decline in the
same time, and to the same extent. The average age composition in autumn was 54%
young willow grouse both at Lovhogen and Ammarnas. Available studies on population
dynamic in tetraonids seem to show that many factors are necessary to explain all the

142 ^

4>

>->" t* **

complex population changes. Key words: willow grouse, snow, ice, sleet, chick
survival, reproductive success, predation

315

Marjakangas, A. 1990. A suggested antipredator function for snow-roosting
behavior in the black grouse Tetrao tetrix. Ornis Scandinavica 21(1):77-78.

Summary: Several studies suggest that grouse conserve heat in cold winter weather by
snow-roosting. However, black grouse frequently snow-roost even in mild weather when
it may not be necessary for energetic reasons, and in moist snow when there is the risk
of being imprisoned after freezing of the snow. Thus, snow roosting may also have
other important functions for black grouse, such as avoidance of predators. Key words:
black grouse, snow, predation, behavior

316

. 1986. On the winter ecology of the black grouse Tetrao tetrix in central

Finland. Acta Universitats Ouluensis Series A Scientiae Rerum Naturalium
Biologica, 29:1-87. Abstract: The snow roosting behavior of the black grouse and the
effects of artificial feeding were studied at three feeding sites in Oulu (75°N) and
Ylivieska (64°N), W Finland, in the winters of 1976/1977-1981/1982. The black grouse
roosted regularly in snow burrows when there was at least 27 cm of readily penetrable
snow, but were able to penetrate relatively hard crusts within the snow. Under adverse
burrowing conditions they tended to prefer the micro- and macrohabitats with the
deepest snow available. The thickness of the roofs of their closed cavities averaged not
more than about 10 cm irrespective of the snow conditions. This together with some
other structural features of their burrows may be related to the avoidance of ground
predators. In relative terms, the closed cavities were high, and those excavated by the
females 8.7% wider than those of the males, enabling more effective ptiloerection. The
black grouse roosted exclusively in closed burrows at -3°C and below, and also in
milder weather, in which case cavity temperatures were well above zero, as shown
experimentally. The flushing distance from closed burrows showed diurnal variation,
being longer for flocks than for solitary grouse. The birds in large flocks roosted closer
to each other than those in small ones. When able to roost in closed burrows, they
restricted their activity In approximately 1 h 25 min in the morning. The black grouse fed
on oats daily, the mean distance of the roosting sites from the feeding site being only
591 m. The sex ratio of the feeding flocks was skewed in favor of males. The birds
spent 62.8% of their diurnal activity at the feeding site. Adult males occupying territories
devoted 43.2% of their individual activity to display throughout the winter, but only
29.9% to feeding on oats, although this was compensated for by a high pecking rate
and prolonged activity. A dominance rank order prevailed among the non-territorial
birds, the juvenile females being the most subordinate, but there were no appreciable
differences between the individual time budgets of the males and females. No ranking
was evident in trees, and the females fed at an equal rate to the males. The rate of dry
matter intake was superior when feeding on oats and lowest with pine. The average
composition of the diet was 86.1% oats, 4.6% birch and 9.3% pine. Particularly in late
winter, the black grouse browsed heavily on bare pine twigs and needles. The mineral
content of the oats, especially Ca and Mn, was lower than that of their natural winter

_y 143

<P

P* fc* **

^c^ ^

food, and consequently the soft tissues of the birds which had fed on oats contained
significantly less of some of the minerals than did those of the control birds and their
caeca were significantly shorter. Possibilities for improving feeding practices are
discussed. Key words: black grouse, snow, behavior

317

, H. Rintamaki, and R. Hissa. 1984. Thermal responses in capercaillie and

the black grouse roosting in snow. Physiological Zoology 57(1):99-104. Abstract:
Thermal responses were studied in the capercaillie and the black grouse roosting in
closed snow burrows in northern Finland. For capercaillie, the air temperature within the
burrow declined with the distance from the bird but was as high as 1 1C adjacent to the
bird when snow temperatures varied between -7.5 and 11. 5C. The body temperature of
the burrowed capercaillie was significantly lower than that of those roosting on the snow
surface, indicating lessened alertness. In a comparison of the responses of the two
species, the body temperature of burrowed black grouse was significantly higher than
that of similarly situated capercaillie. The burrowed black grouse shivered occasionally
but weakly during the nest. Measurements of pectoral muscle electromyographic
activity (EMG) were not taken for capercaillie. The results support the hypothesis that in
winter, under suitable snow conditions, these birds are able to shelter in, or close to, a
thermoneutral microenvironment. Keywords: capercaillie, black grouse, snow,
thermoregulation

318

Marks, J. S. and V. S. Marks. 1988. Winter habitat use by Columbian sharp-tailed
grouse in western Idaho. Journal of Wildlife Management 52(4):743-746. Abstract:
We studied habitat use by Columbian sharp-tailed grouse (Tympanuchus phasianellus
columbianus) during 3 winters in western Idaho. Grouse were closely associated with
mountain shrub and riparian cover types, the only cover types that provided food and
escape cover regardless of snow depth. Fruits of Douglas hawthorn (Crataegus
douglassi) and buds of Saskatoon serviceberry (Amelanchier alnifolia) and common
chokecherry (Prunus virginiana) were the main winter foods. Hawthorn buds apparently
were not eaten by grouse. Because hawthorn fruits were not available throughout the
winter, plants producing palatable buds or catkins should be present to provide food
during late winter and in years when fruit production is poor. Key words: Columbian
sharp-tailed grouse, snow, vegetation, nutrition

319

Marra, R. P., S. Griffing, C. Caffrey, A. M. Kilpatrick, R. McLean, C. Brand, E. Saito,
A. P. Dupuis, L. Kramer, and R. Novak. West Nile Virus and wildlife. BioScience
54(5):393-402. Notes: Page 399 — "Wet springs and hot, dry summers may also
facilitate WNV epidemics. Such was the case in the New York City area in 1999 and in
Colorado in 2003, when the production of mosquitoes in higher than normal numbers
was followed by maximized rates of viral replication within mosquitoes (because of
warm temperatures), leading to shortened transmission and amplification cycles.
Torrential rains at summer's end can also lead to heightened amplification by causing

144

% *>'<0^

an increase in ovipositioning sites." Key words: temperature, disease, West Nile
Virus

320

Marshall, A. J. 1949. Weather factors and spermatogenesis in birds. Proceedings
of the Zoological Society of London Series A 119:711-716. Unable to obtain for
abstract.

321

Marshall, W.H. 1965. Ruffed grouse behavior. BioScience 15(2):92-94. Summary:
The authors believe that the paramount factors affecting ruffed grouse behavior during
cold periods involve both a quest for habitats to minimize energy loss and for foods of
high calorigenic value. During 1963 the lack of snow precluded roosting at night in snow
burrows, one means by which grouse avoid heat loss. Instead, this bird was forced to
remain exposed to the winter night sky which, at Cloquet, Minnesota, is often a clear
and an intensely cold radiation sink. By utilizing the spruce-fir stands for roosting at
night, this bird selected a habitat wherein energy leakage by radiation was limited. By
feeding rapidly on the buds of clones of staminate aspen trees for short periods of time
the birds choose a very rich source of food while minimizing energy loss. Key words:
ruffed grouse, temperature, snow, behavior, habitat use

322

and M. S. Jensen. 1937. Winter and spring studies of the sharp-tailed

grouse in Utah. Journal of Wildlife Management 1(3/4):87-99. Notes: The shifts to
and from the maple-chokecherry type parallel changes in snow depth and in elevation.
Roosting — Cover requirements... apparently varied largely with snow depths and crust
conditions. When there was no snow the birds were found using both the sage types
and the weed-grass type. As snow covered up the latter, they shifted to the sage types
and even when the sage was covered, they would use this type, if there was no crust,
by diving or burrowing into the snow often until they were in the branches of the sage
brush. Heavy snow crusts in January and February caused the birds to roost in bushes
that were above the snow; since maple and chokecherry were the only ones present on
the area, they were the ones used. Feeding — Availability of food apparently influenced
the food habits greatly. Only once has evidence been obtained that the sharptail digs or
scratches for food; thus, as snow depths increase or decrease, certain foods in turn
lose or maintain availability. The heights of the important foods above the ground are
charted with snow depths for the winter period... attention is called to the correlation of
changes in foods eaten with snow depths. Key words: sharp-tailed grouse, snow,
snow condition, habitat use, feeding, behavior, vegetation

323

Martin, K. and K. L. Wiebe. 2004. Coping mechanisms of alpine and arctic
breeding birds: extreme weather and limitations to reproductive resilience.
Integrated Comparative Biology 44:177-185. Synopsis: As ground nesting
homeotherms, alpine and arctic birds must meet similar physiological requirements for

i fv^^c

145

% #> o 9

breeding as other birds, but must do so in more extreme conditions. Annual spring
snowfall and timing of snow melt can vary by up to 1 month and daily temperatures near
the ground surface vary from below freezing to over 45°C in alpine and arctic habitats.
Species breeding in these environments have various behavioral, physiological, and
morphological adaptations to cope with energetically demanding conditions. We review
the ways birds cope with harsh and variable weather, and present data from long term
field studies of ptarmigan to examine effects of spring weather on reproduction. In
variable but normal spring conditions, timing of breeding was not influenced by snow
melt, snow depth or daily temperatures in the alpine, as breeding did not commence
until conditions were generally favorable. Arctic ptarmigan tended to vary breeding
onset in response to spring conditions. Generally, birds breeding in alpine and arctic
habitats suffer a seasonal reproductive disadvantage compared to birds at lower
latitudes or elevations because the breeding window is short and in late years, nest
failure may be high with little opportunity for renesting. Coping mechanisms may only be
effective below a threshold of climactic extremes. Despite strong resilience in fecundity
parameters. When snowmelt is extremely delayed breeding success is greatly reduced.
Alpine and arctic birds will be further challenged as they attempt to cope with
anticipated increases in the frequency and severity of weather events (climate varia
bility), as well as general climate warming. Key words: ptarmigan, snow,
temperature, reproduction

324

Martinson, R. K. 1963. The relationship of weather to productivity in upland game
birds. Pittman-Robertson Project W-67-R-4, Phase B, Upland Game
Investigations, Job No. 17, North Dakota State Game and Fish Department,
Bismark, USA. Abstract: The relationship of annual weather patterns to productivity
among sharp-tailed grouse, ruffed grouse and European partridges in North Dakota was
investigated by comparing age ratios from those species with weather data. The various
weather parameters used in the analyses included: (1) temperature, (2) precipitation,
(3) Setzer Indices and (4) Hammond Method indices. Age ratios among sharp-tailed
grouse from the western part of the state showed positive correlation with April plus
May precipitation; those from the eastern part of the state with March plus April
precipitation. In general, high age ratios among sharp-tails were found in years that had
relatively high spring precipitation. Age ratios from ruffed grouse exhibited positive
correlation with April plus May plus June temperatures; i.e., the higher rates of
productivity among ruffed grouse occurred in years when spring and early summer
temperatures were warmer than average. European partridge age ratios showed
negative correlation with indices of coolness calculated for periods during June and
July. That is, the higher age ratios among partridges were observed in years that had
relatively warm periods in June and July. The decline which occurred in sharptail,
partridge and pheasant populations in 1959 and the ten-year ruffed grouse "cycle" were
discussed with reference to the affects of weather. Notes: At the present time it appears
that weather, among all density-independent factors, has perhaps the most profound
impact on upland game bird populations occupying static habitat. If this is so, the game
manager with a knowledge of climatological phenomena and their influence on
populations may be able to foretell or explain changes in the abundance of game birds.

146 jj

4<

H (& **

cCb

Key words: sharp-tailed grouse, ruffed grouse, European partridge, precipitation,
temperature, age ratio, productivity, index

325

and C. R. Grondahl. 1966. Weather and pheasant populations in

southwestern North Dakota. Journal of Wildlife Management 30(1):74-81. Abstract:
High productivity and survival of ring-necked pheasants {Phasianus colchicus) were
correlated with high rainfall and cool temperatures in May and June during an 8-year
period in southwestern North Dakota. These findings differed markedly from those of
several earlier studies in the less arid Midwestern United States where cool, wet
weather in the spring generally affected pheasant populations adversely. General
trends in the pheasant population could be explained on the basis of precipitation in
May and June. The population was relatively high during the mid-1950s when wet and
dry May and June periods occurred in alternate years but declined markedly after 2
consecutive years of spring and summer drought. The population remained at a static
low during 1960 through 1963 when wet and dry spring and summer periods again
occurred alternately. Key words: ring-necked pheasant, temperature, precipitation,
drought, population dynamics

326

Merchant, S. S. 1982. Habitat use, reproductive success and survival of female
lesser prairie chickens in two years of contrasting weather. M.S. Thesis, New
Mexico State University, Las Cruces, USA. Abstract: Spring and summer habitat use,
home ranges, reproductive success, and survival of female lesser prairie chickens
(Tympanuchus pallidicinctus) in Roosevelt and Lea counties, New Mexico, are
contrasted between 1979 and 1980. In 1979, weather was near optimal for prairie
chicken reproduction and survival, whereas in 1980, drought conditions existed. Effects
of these conditions are discussed.... The increased size of home ranges in 1980 was
attributed to effects of the drought... All indices to reproductive success were lower for
1980 than for 1979. These included the percent of females nesting, percent of females
renesting, percent of successful nests, number of brood observations, and mean brood
size. The lower reproductive success in 1980 was attributed to direct and indirect
influences of the drought. Spring and summer survival of females was lower in 1980,
probably due to increased vulnerability to predators, which was associated with the
stressful drought conditions. Key words: lesser prairie chicken, drought,
precipitation, vegetation, habitat use, predation, reproduction

327

Meriggi, A., D. Montagna, D. Zacchetti, C. Matteucci, and S. Toso. 1990.
Population dynamics of the gray partridge in relation to agriculture and weather
in northern Italy. Pages 241-253 in K. E. Church, R. E. Warner, and S. J. Brady,
editors. Proceedings Perdix V: gray partridge and ring-necked pheasant
workshop. Minnesota Department of Natural Resources, Mankato, USA. Abstract:
Car censuses were carried out on a 175-km2 area of north Italy from spring 1984 to fall
1988. Seasonal and annual changes in gray partridge density, recruitment and mortality
were estimated. The average density was 4.2 pairs/km2 (SE = 0.39) in spring and 3.4

-¥ \^ , - . , l. 147

% H ^ ^c

Cgf ^ <£^> ^b

broods/km2 (SE = 0.29) in summer. Average brood size was 12.8 (SE = 2.73) at
hatching and 8.2 (SE = 0.33) when full grown. Chick mortality rate from hatching to 30
days old averaged 42.4%. Brood size was constant after 30 days post-hatching. Spring
density was closely correlated with the percentage of winter cereals (wheat and barley)
and the length of hedgerows. The high rate of recruitment is followed in winter by a high
rate of mortality (64.3%), as the population was reduced to the levels of the previous
spring. We hypothesize cereals determine the carrying capacity of the study area for
gray partridge, as a food source and alternative nesting cover. Key words: gray
partridge, mortality, winter weather

328

Meunier, J. 2005. Fall migration chronology and habitat use of the American
woodcock in the western Great Lakes region. M.S. Thesis, University of
Wisconsin-Madison, Madison, USA. Conclusions: "Result of my investigation
indicated that there are many factors that likely contribute to the onset of woodcock
migration including factors often described by observations of woodcock in the process
of migration, namely synoptic weather variables of low pressure systems like barometric
pressure changes and northwest wind directions. My research indicates that these
weather variables may be of secondary importance to photoperiod, which served as the
initial cue and in all years demonstrated the greatest influence on woodcock migration.
Photoperiod likely serves to synchronize the Zugstimmung, the initiation of migration of
woodcock such that woodcock at northern latitudes initiate migration prior to woodcock
at lower latitudes despite proximate weather in different locales... Once migration
begins, local weather influenced the rate of migration." Key words: American
woodcock, barometric pressure, wind, photoperiod, movement, behavior

329

, R. S. Lutz, K. E. Doherty, D. E. Andersen, E. Oppelt, and J. G. Bruggink.

2006. Fall diurnal habitat use by adult female American woodcock in the western
Great Lakes Region. 10th American Woodcock Symposium:11. Michigan
Department of Natural Resources and United States Fish and Wildlife Service.

Abstract: We assessed how habitat structure and food availability influenced use of
cover types at the habitat patch and home-range scales by adult female (after hatch
year) American woodcock in Michigan, Minnesota, and Wisconsin from 2002 through
2004. We also investigated use of alder (Alnus spp.) as a staging cover prior to fall
migration. We selected this cover to investigate seasonally changing use of cover types
based on past observations. Seasonal changes in cover type use could have important
ramifications for woodcock management intended to provide or improve woodcock
habitat. We measured edge proximity, stem density and earthworm abundance at
woodcock locations and paired these locations to random locations at the micro-habitat
scale (2002 data); 20 m from use locations within the same stand. We also compared
edge proximity with paired use and random locations at the home range scale (2003
and 2004 data): >35<200 m from use locations across cover types and investigated
habitat selection at this scale. Adult female woodcock (n^ 39) used a variety of cover
types and the percent of total use changed among years and states. We found the
greatest frequency of alder use across all states in 2003, a drought year. We found no

148

% l-x^^

difference between alder use during the entire fall period and early fall (P=0.9) or late
fall (P=0.7) indicating that alder was not used as staging cover prior to migration. We
found that structural habitat features were more important than food resources to
habitat selection at both the micro-habitat and home-range spatial scales. Cover types
most heavily used by woodcock often had the lowest earthworm abundances. We did
find evidence that woodcock selected for edge proximity at both home-range and micro-
habitat scales, especially in mature cover types. Woodcock also selected habitats with
higher shrub densities and fewer mature stems in young cover types than we found at
random. Key words: American woodcock, drought, habitat use

330

Middleton, A. D. 1936. The population of partridges (Perdix perdix) in Great
Britain during 1935. Journal of Animal Ecology 5:252-261. Notes: "...general
observation indicates that the heavy rainfalls and lack of sunshine during June and July
have been the cause of a great deal of mortality among young partridges over most of
England." Key words: partridge, precipitation, solar radiation, mortality

331

Miller, D. A., G. A. Hurst, and B. D. Leopold. 1997. Factors affecting gobbling
activity of wild turkeys in central Mississippi. Proceedings of the Annual
Conference Southeast Association of Fish and Wildlife Agencies 51:352-361.

Abstract: Call counts for a number of gamebirds (e.g., northern bobwhite [Colinus
virginianus] and wild turkey [Meleagris gallopavo]) have been used to index population
levels and trends and to document species presence or absence. Call counts for wild
turkeys have been used for these purposes, but gobbling activity has not been related
quantitatively to population size, reproduction, weather, male age structure, or hunting
variables. Consequently, we examined these factors as they affected gobbling activity
on Tallahala Wildlife Management Area, Bienville National Forest, in central Mississippi,
from 1984 to 1995. Using multiple linear regression, we determined that within-year
gobbling activity was related to hunter effort, days into call count period, wind velocity,
year, and dewpoint. Among years, gobbling activity was related to hunter effort and
hunter success. An index to proportion of 2-year-old gobblers in the population was
correlated to an increased number of gobblers heard, but not number of calls heard.
Gobbling activity was influenced by a complex interaction of population and
environmental conditions that cannot easily be modeled. In central Mississippi, gobble
call counts were not related to gobbler population size, and their applicability in other
areas warrants examination. Key words: turkey, wind, dewpoint, behavior, census

332

Miller, M. R., C. L. Stemler, J. L. Yee, and D. S. Blankenship. 2001. Differences in
mourning dove productivity among three time periods at Gray Lodge Wildlife
Area, California. California Fish and Game 87(3):93-1 01. Abstract: We used
published and unpublished data to compare mourning dove, Zenaida macroura,
recruitment among the 1949-50 (early), 1979-80 (middle), and 1992-95 (late) periods in
riparian habitat at Gray Lodge Wildlife Area, California, to document any differences
that corresponded to the long-term decline of doves in California (as estimated by the

x

\ *^"^>^

149

Cgc 0fe <££ d£

(J. S. Fish and Wildlife Service's annual Call-count Survey). Estimates of true nest
success, fledglings produced per nesting attempt, and fledglings produced per active
dove pair, were highest in the early period, lowest in the middle period, and low to
moderate in the late period (nest success: 44-56% early, 25-29% middle, 24-25% late;
fledglings/nest attempt: 1.20-1.44 early, 0.78-0.91 middle, 0.81-1.03 late; fledglings/pair:
5.93-6.58 early, 2.403.40 middle, 5.00-5.60 late). In the early period, high production
coincided with the warmest and driest spring weather and an active predator control
program, conditions thought to boost dove production. However, based on banding
estimates of survival, fledgling production/pair during each period should have been
more than adequate to increase breeding populations (2.5 requires)(with the exception
of 1980). Therefore, continued population decline in California must result from
landscape-scale habitat alteration, poor survival, or both, and not recruitment; or high
recruitment at Gray Lodge was not representative of California nesting doves (or
riparian habitats) in general; or the three periods during which data were collected at
Gray Lodge were too infrequent relative to the 47-year time period to detect long-term
change in recruitment. Initiation of a wing-collection survey to monitor harvest age ratios
and an operational banding program to estimate survival would ultimately provide more
predictable data sets for analyses of long-term trends in annual population status than
would additional analyses of historical nesting data. Key words: mourning dove,
temperature, precipitation, productivity

333

Mitchell, G. J. 1977. The effects of spring and summer weather on Hungarian
partridge productivity in southern Alberta. Pages 201-209 in G. D. Kobriger,
editor, Proceedings Perdix I Hungarian Partridge Workshop, Dickinson, North
Dakota, USA. Notes: ...correlation analysis shows that, with but two exceptions, all of
the correlations involving temperatures and the hours of bright sunshine are positive,
while those involving rainfall, the number of days with rain, and the number of days with
frosts are negative. These relationships and the brood size and weather data for 1961
and 1953, indicate that reproductive success of the Hungarian partridge in Alberta is
enhanced by warm and dry spring-summer weather, and impaired by rainy and cool
weather during the nesting and rearing season... It appears that weather factors have
both a pre-hatching and post-hatching influence on the productivity of the Hungarian
partridge in Alberta. This productivity — expressed in terms of mean brood size — is
highest in years with above average temperatures and hours of bright sunshine; few
days with frost; and below-average amounts of rainfall in the May to July period.
Conversely, productivity is lowest in those years characterized by cool and moist
summer weather. During the pre-hatching period, the relatively high correlations found
between brood size data and the values for temperatures and rainfall in May indicates
that the minimum daily temperatures and the amounts of rainfall are important factors
influencing Hungarian partridge productivity. Key words: Hungarian partridge,
precipitation, solar radiation, fog, frost, temperature, reproduction, productivity

334

Montagna, D. and A. Meriggi. 1991. Population dynamics of grey partridge (Perdix
perdix) in northern Italy. Bollettino di Zoologia 58(2):151-155. Abstract: Population

150

4>

>->• i& **

dynamics of grey partridge was studied from 1982 to 1986 in northern Italy in order to
evaluate fluctuations in density and the relationships between population parameters
and climatic factors. The general trend was a clear decrease both in spring and summer
density after 1984, weighting 34.6% and 22.2%, respectively, mainly due to severe
winters. The reproductive success and the recruitment were negatively correlated with
spring density. Brood size significantly decreased through the rearing season and chick
mortality occurred mainly within the first 30 days after hatching in 1985, while in 1984 it
was distributed over the whole growing period. It was also significantly related to the
hatching period, earlier broods showing higher mortality. Chick mortality was strongly
related to weather conditions in spring, which negatively affected the abundance and
distribution of insects. Adult losses from spring to summer were found to be negatively
related to reproductive success and positively to spring density. Both density-dependent
and climatic factors appeared responsible for a control of population levels. Severe
winters and cold springs represented limiting factors; however, the population showed
ability to recover by increasing reproductive success and recruitment. Key words: grey
partridge, temperature, mortality, population dynamics, insects, severe winter

335

Moritz, W. E. 1988. Wildlife use of fire-disturbed areas in sagebrush steppe on the
Idaho National Engineering Laboratory. M. S. Thesis, Montana State University,
Bozeman, USA. Notes: Page 55 — "Although this area does not appear to be critical
winter range, removal of any sagebrush probably reduced its suitability as winter
habitat. Because sage-grouse distribution on winter range has been reported to vary
with snow depth, higher winter use of this fire scar may have been a result of mild
winters occurring both years." Page 59 — "The precise use of prescribed burning to
increase sage-grouse populations required detailed knowledge of sage-grouse
movements related to seasonal weather patterns plus a detailed vegetation inventory of
the entire area utilized. ..the best management practices is a maintenance of sagebrush
habitats known to be utilized by sage-grouse." Page 85 — "Seasonal use and distribution
of most mobile species on the INEL appears to be related to seasonal precipitation
patterns. Snow depth and summer rains appear to follow storm tracks which may be
related to the locations of nearby mountain ranges. While weather monitoring stations
currently exist on the area, a map of precipitation patterns and storm tracks would be of
great benefit to investigations of animal movements, especially in situations where both
migratory and non-migratory populations appear to coexist." Key words: sage-grouse,
precipitation, snow, vegetation, habitat manipulation, movement, habitat use,
weather mapping

336

Morrow, M. E., R. S. Adamcik, J. D. Friday, and L. B. McKinney. 1996. Factors
affecting Attwater's prairie-chicken decline on the Attwater Prairie Chicken
National Wildlife Refuge. Wildlife Society Bulletin 24(4): 593-601. Abstract: We

examined the association between changes in the number of Attwater's prairie-chicken
(Tympanuchus cupido attwateri) on the Attwater Prairie Chicken National Wildlife
Refuge from 1972 to 1993 and changes in refuge management practices (burning and
grazing). We also examined the relationship of prairie-chicken population increases and

% H>,^

151

decreases on the refuge to rainfall and off-refuge prairie-chicken populations. Burning
within the prairie-chicken's core habitat on the refuge and variability in grassland
structure were directly correlated (P < 0.05) with increases and declines in prairie-
chicken populations. Refuge population increases and declines were inversely
correlated (P< 0.1) with average April rainfall/event, May absolute departure from long-
term average rainfall, May number of rainfall events, annual absolute departure of
rainfall from the long-term average, and the annual number of rainfall events. Refuge
population increases and declines were directly correlated (P < 0.1) with off-refuge
populations, although the off-refuge population decline began 4 years earlier than on-
refuge Key words: prairie-chicken, precipitation, land use

337

and N. J. Silvy. 1982. Nestling mortality of mourning doves in cental Texas.

Proceeding of the Annual Conference Southeast Association of Fish and Wildlife
Agencies 36:537-542. Abstract: Mourning dove (Zenaida macroura) nests were located
on the Texas A&M University Campus from February-October 1981. Mortality differed
between the various stages of the nesting cycle, and in different months within each
stage. The number of days in which wind speed exceeded 27.6 km/h (15 kts) each
month explained 34% of the variability (P=0.10) associated with total nesting mortality.
Mean monthly temperature was positively correlated (P<0.10) with mortality occurring
during the 1st nestling week and negatively correlated during the 2nd week after
hatching. Loss of nestlings was correlated (PO.05) with an index to blue jay
(Cyanocitta cristata) production and total monthly rainfall (P=0.10). Key words:
mourning dove, wind, temperature, precipitation, mortality

338

Moss, A. 1985. Rain, breeding success and distribution of capercaillie Tetrao
urogallus and black grouse Tetrao tetrix in Scotland. Ibis 128:65-72. Abstract: The
number of chicks reared by capercaillie was inversely related to the number of days
with rain during and just after their hatching. However, rain had no detectable effect on
the breeding success of black grouse on the same study area. The breeding distribution
of capercaillie in Scotland is more limited than that of black grouse, which extends into
rainier areas. It is speculated that sexual selection for large size in capercaillie has
indirectly caused their chicks to be more vulnerable to rain and, consequently, has
limited their distribution to less rainy areas. Key words: capercaillie, black grouse,
precipitation, productivity, population dynamics

339

Moss, R. and A. Watson. 2001. Population cycles in birds of the grouse family
(Tetraonidae). Advances in Ecological Research 32:53-111. Abstract: Population
cycles of boreal herbivores such as voles, hares and birds of the grouse family
(Tetraonidae) were an early problem in ecology. Here we systematize present
understanding of population cycles in tetraonids. Cycles have not been recorded in all
grouse species, and non-cyclic populations occur in all grouse species known to show
cycles. Much knowledge about cycles comes from time series of measures of
population abundance. The fluctuations in such series have properties such as period,

152 j^

4>

*-> t* **

#

*I/"

cCi

amplitude, symmetry, and degree of synchrony with other time series of the same and
other species. Each property shows temporal and spatial variation, and is used to
delineate hypotheses about causes of cycles. Cyclic time series show delayed density
dependence, as expected from models. From species known to show cycles, we give
three examples of non-cyclic populations that showed direct density dependence. Each
was in fragmented or degraded habitat. Much anecdotal evidence suggests that grouse
cycles occur mostly in large tracts of fairly homogeneous habitat, and that fragmentation
of such habitat is associated with disappearance of cycles. This has been attributed to
increased dispersal out of fragmented habitat patches into population sinks, and to
increased predation from generalist predators. We distinguish between processes that
drive cycles and those that determine cycle period. Commonly reported periods are 3-4,
6-7 and 10 years. Different populations of the same species show different periods, and
sympatric species with different mean vital rates fluctuate together with the same
period. Hence cycle period is not a necessary consequence of a particular set of mean
vital rates. Period is characteristic more of location than of species. There is no simple
large-scale relationship between cycle period and latitude or longitude. One example, of
a 10-year rock ptarmigan cycle going together with a 10-year weather cycle, suggests
that local weather might affect period. Documented extremes in the amplitude of cycles
in population density vary from about two-fold to about six-fold. Most documented
tetraonid population cycles are fairly symmetrical, with decline and increase phases in
any one fluctuation taking similar number of years. Fluctuations of spatially separate
parts of cyclic populations have shown partial synchrony that diminishes with distance
between the parts. Explanations of this include the effects of weather and dispersal.
Dispersal might be of the cyclic species, or its resources, or one of its natural enemies
or competitors. A consequence of dispersal in some cyclic model populations is
travelling waves of population density, so far confirmed for species of grouse, hare and
vole. The main documented demographic cause of changes in tetraonid breeding
numbers is variation in the recruitment of young birds into the breeding population. In
seven studies of two species, including cycles with periods of 3-4, 7-8 and 10 years, this
happened in two stages. First, during cyclic declines the birds' breeding success' was
lower, that is fewer young were reared to independence per adult. Second, a smaller
proportion of these independent young survived the winter to be recruited into the
breeding population. However, mean breeding success differed among populations
within species, such that cyclic increases in some populations occurred with lower
breeding success than cyclic declines in other populations. Key words: grouse,
weather, population dynamics

340

and J. Oswald. 1985. Population dynamics of capercaillie in a north-east

Scottish glen. Ornis Scandinavica 16(3):229-238. Abstract: During a 9-year study of
capercaillie almost twice as many female as male chicks were reared, but there were no
more hens than cocks in the adult population. Both the proportion of male chicks in late
summer each year, and mean brood size, were related to the chicks' condition in July.
Condition was related to the number of days with rain in early June, during and just after
hatching. The breeding density of hens each year varied between 4.0 and 8.0 km2 but
was related neither to their density in the previous year nor to the production of female
chicks in the intervening summer. The best predictive model was simply that breeding

% Hk<***

153

densities would return to their mean each year. It was inferred that breeding densities
were determined by density-dependent losses and gains involving emigration and
immigration. Autumn densities of young and old birds were not related to breeding
densities and breeding success, but were inversely related to the proportion of broods
which disappeared, and probably emigrated, during the summer. Key words:
capercaillie, precipitation, population dynamics

341

, , and D. Baines. 2001. Climate change and breeding success: decline

of the capercaillie in Scotland. Journal of Animal Ecology 70(1):47-61. Summary:
(1 ) The hypothesis that climate change caused lower breeding success was
investigated in this study. (2) Temperature usually rose during April. There was no trend
in mean April temperature during the study (1975-99) but there was a progressive
cooling in mid-April relative to the rest of the month, such that the normal April warming
was increasingly delayed. (3) Hens reared more chicks when the temperature rose
more in early April. It is suggested that this stimulated timely plant growth, so improving
the laying hens' plane of nutrition and the viability of their chicks. (4) Hens also reared
more chicks when late May was warmer and early June was warmer and had fewer rain
days. Young chicks may have foraged more successfully in warm dry conditions.
However, neither temperature nor rain days in late May or early June showed any trend
during the study. (5) Increasingly protracted spring warming seems to have been a
major cause of the decline of the capercaillie in Scotland. Key words: capercaillie,
temperature, precipitation, population dynamics, vegetation

342

, A. Watson, R. A. Parr, I. B. Trenholm, and M. Marquiss. 1993. Growth rate,

condition and survival of red grouse Lagopus lagopus scoticus chicks. Ornis
Scandinavica 24(4):303-310. Summary: An index of condition was developed for wild
red grouse chicks aged 10-30 days. This was derived from the difference between the
observed weight and that expected from the chick's age, or stage of feather
development. To measure it, the chicks needed to be caught only once. Variations in
condition among different broods of wild chicks were greater than variations among
broods of captive chicks in the same years, probably because the wild chicks'
environment was harsher and more variable. Within years, the condition of wild chicks
was not related to their brood size. Among years, mean condition was correlated both
with breeding success and with the biomass of green heather food available in spring.
This was because the chicks' condition and survival were poor for two years following
an episode of severe weather damage to the heather. Key words: red grouse, index,
nutrition, fecundity

343

, , , and . 1993. Caecal threadworms Trichostrongylus

tenuis in red grouse Lagopus lagopus scoticus: effects of weather and host
density upon estimated worm burdens. Parasitology 107(2):199-209. Abstract:
Trichostrongylus tenuis eggs were counted in feces from individually marked wild red
grouse for 8 years. Egg counts varied seasonally and annually. In some years, a

154 ^

4>

» <0 >f*

sudden increase in mid-April was consistent with delayed maturation of larvae which
had overwintered in the birds in a hypobiotic state. A more gradual increase in summer
was probably due to uninterrupted maturation of larvae ingested then. Despite 30-fold
year-to-year variation in mean egg counts, relative differences in egg counts among
known individuals within years tended to persist across years. Rainfall in previous
summers explained much of the year-to-year variation in egg counts, probably because
parasite recruitment was greatest during wet summers. Grouse density was only weakly
related to worm egg counts. The data were not consistent with the hypothesis that the
cyclic-type population fluctuation in red grouse numbers observed at the time of this
study was caused by the parasites. Key words: red grouse, disease, precipitation,
population dynamics, parasites

344

Mossop, D. H. 1988. Winter survival and spring breeding strategies of willow
ptarmigan. Pages 330-378 in A. T. Bergerud and M. W. Gratson, editors. Adaptive
strategies and population ecology of northern grouse. Wildlife Management
Institute and University of Minnesota Press, Minneapolis, USA. Notes: Page 337 —
Wind, as manifested by windchill and blowing snow, is probably the chief factor of
winter severity. The windchill at Chilkat Pass was low enough that humans often found
it difficult to function. However, in the microclimates that ptarmigan occupied, the birds
virtually never exposed themselves to these extremes in wind. Even in the worst
weather, ptarmigan were found with full crops and did not forgo feeding. I found no
evidence that the formation of a surface crust prevented snow-roosting. Snow
sufficiently loose to excavate was always available in the wind shadows of shrubs. Page
340 — Food actually became more available on the Chilkat Pass as winter passed and
the snow level rose. Birds gained access to higher and higher levels of the shrub crown,
where abundance and possibly nutritional content of buds and twigs were better. I
recorded an increase in browse availability in food tallies immediately after blizzards.
Page 347 — The physical link between vegetative cover and snow roosts is also
maintained; snow roosts of willow ptarmigan are normally dug close to or in shrub
cover. Birds roosting near shrubs gain advantage because predator search paths are
partly blocked. Page 352 — When birds left the Highland area in February, adequate
food remained available. Birds actually moved to an area with more heavily browsed
shrub resources. Further, they shared this food with more birds from many adjacent
ranges. They did not make this move until late winter, after the worst weather had
passed. Accumulation of the snowpack was the only weather parameter that I could
causally relate to movement. As the accumulated snow reached about 1 m, leaving only
1 m of overhead shrub cover, birds began to leave upland sites and move to tall
patches of riparian willow cover at lower elevations... cover to escape predators (both
detection and once pursued, i.e. in the broad sense) is a sufficient explanation for this
late-winter migration. When the snowpack had reached about 1 m on unprotected
ground, drifting snow had inundated shrubs on the pass well into their crowns. This left
ample food, in the form of shrub tips above the snow, but cover was lacking. Larger
riparian shrubs at lower elevations were sought by the ptarmigan — this was the winter
migration. Key words: willow ptarmigan, snow, wind, habitat use, vegetation,
predation, microclimate, movement, behavior

** l& *k

JV 1>V l . L- 155

4-

<QJr -"CST- <£~^}> cC^

345

Moynahan, B. J. 2004. Landscape-scale factors affecting population dynamics of
greater sage-grouse (Centrocercus urophasianus) in north-central Montana,
2001-2004. Thesis, University of Montana, Missoula, USA. Abstract: Populations of
greater sage-grouse (Centrocercus urophasianus) have declined by 69-99%.
Information on population dynamics of these birds at a landscape scale is essential to
informed management. I radio-collared 243 female sage-grouse, monitored 287 nests
and 115 broods, and measured 426 vegetation plots at 4 sites during 2001-2003 in a
3,200 km2 landscape in north-central Montana, USA. My objective was to examine the
relationship between nest success, brood survival, and hen survival rates, habitat
conditions, environmental variables, and hen characteristics. I used program MARK to
model (1) daily survival rates of nests and broods and (2) seasonal and annual survival
of hens. Nest survival varied with year, grass canopy cover, daily precipitation with a 1-
day lag effect, and nesting attempt. The best-approximating model of brood survival
included effects of brood age and year, indicating substantial annual variation. Hen
survival analyses indicated that survival varies by season within years and by year
within seasons, that nesting hens have higher breeding-season survival than non-
nesting hens, and that individuals at one site had lower hunting-season survival than
hens at other sites. I observed considerable variation in hen survival. Low annual
survival in 2003 is a result of the compounded effects of a West Nile virus outbreak in
August of that year and a severe winter of 2003-2004. My findings underscore the
importance of large-scale approaches to conservation of sage-grouse habitats and to
maintenance and recovery of sage-grouse populations. Management for hen survival
must address hunting pressure and identification and conservation of important
wintering areas. Maintaining quality habitat and a high proportion of adult hens will
maximize potential for population growth when environmental conditions are favorable.
Key words: sage-grouse, precipitation, temperature, snow, disease, mortality,
model, habitat

346

, M. S. Lindberg, J. J. Rotella, and J. W. Ward. 2007. Factors affecting nest

survival of greater-sage grouse in north central Montana. Journal of Wildlife
Management 71():1773-1787. Abstract: We studied greater sage-grouse (Centrocercus
urophasianus) in north central Montana, USA, to examine the relationship between nest
success and habitat conditions, environmental variables, and female sage-grouse
characteristics. During 2001-2003, we radiomarked 243 female greater sage-grouse,
monitored 287 nests, and measured 426 vegetation plots at 4 sites in a 3,200-km2
landscape. Nest survival varied with year, grass canopy cover, daily precipitation with a
1-day lag effect, and nesting attempt. In all years, daily survival rate increased on the
day of a rain event and decreased the next day. There was temporal variation in nest
success both within and among years: success of early (first 28 d of nesting season)
nests ranged from 0.238 (SE = 0.080) in 2001 to 0.316 (SE = 0.055) in 2003, whereas
survival of late (last 28 d of nesting season) nests ranged from 0.276 (SE = 0.090) in
2001 to 0.418 (SE = 0.055) in 2003. Renests experienced higher survival than first
nests. Grass cover was the only important model term that could be managed, but
direction and magnitude of the grass effect varied. Site, shrub and forb canopy cover,

156 ^

<^

^ <0 ^

cCi

and Robel pole reading were less useful predictors of nest success; however, temporal
and spatial variation in these habitat covariates was low during our study. We note a
marked difference between both values and interpretations of apparent nest success,
which have been used almost exclusively in the past, and maximum-likelihood
estimates used in our study. Annual apparent nest success (0.56) was, on average,
53% higher than maximum-likelihood estimates that incorporate individual,
environmental, and habitat covariates. The difference between estimates was variable
(range = +8% to +91%). Management of habitats for nesting sage-grouse should focus
on increasing grass cover to increase survival of first nests and contribute to favorable
conditions for renesting, which should be less likely if survival of first nests increases.
Key words: sage grouse, precipitation, nesting success

347

, M. S. Linberg, and J. W. Thomas. 2006. Factors contributing to process

variance in annual survival of female greater sage-grouse in Montana. Ecological
Applications 16(4): 1529-1 538. Abstract: Populations of greater sage-grouse
{Centrocercus urophasianus) have declined by 69-99% from historic levels, and
information on population dynamics of these birds at a landscape scale is essential to
informed management. We examined the relationships between hen survival and a
suite of landscape-scale habitat and environmental conditions. We radio-marked 237
female sage-grouse and measured 426 vegetation plots during 2001-2004 at four sites
in a 3200-km2 landscape in north-central Montana, USA. We used program MARK to
model monthly survival rates for 1 1 seasonal intervals. There was strong support for the
best-approximating model (AlCc weight = 0.810), which indicated that (1) hen survival
varied by season within years' and by year within seasons, (2) nesting hens had higher
nesting-season survival than non-nesting hens, and (3) individuals at one site had lower
hunting-season survival than at other sites. We observed considerable variation in hen
survival. Process variation was 0.255, with an expected range of annual survival of 0.12
to 1 .0. The ratio of process to total variation was 0.999, indicating that observed
variation was real and not attributable to sampling variation. We observed a nearly
fourfold difference in maximum and minimum annual survival, ranging from 0.962±
0.024 (mean ± SE) for nesting hens in 2001-2002 to 0.247 ± 0.050) for non-nesters in
2003-2004. Low annual survival in 2003 resulted from the compounded effects of a
West Nile virus outbreak in August and a severe winter in 2003-2004. Increased hen
mortality associated with severe winter weather contrasts with prior beliefs that sage-
grouse populations are typically unaffected by winter weather conditions and
underscores the importance of protecting winter sagebrush (Artemisia spp.) habitats.
Key words: sage-grouse, severe weather, vegetation, mortality

348

and J. Walker. 2004. Literature review of Montana upland game bird biology

and habitat relationships as related to Montana Fish, Wildlife & Parks upland
game bird habitat enhancement program. Personal Services Contract: FWP-
050046. Montana Department of Fish, Wildlife & Parks, Helena. Summary: The
authors provide an overview of the grouse species, pheasants, turkey and partridges.
Includes a literature cited. Key words: upland birds, weather

f^f ^ ^

,>"^

<£Sh cC^

349

Mussehl, T. W. and F. W. Howell, editors. Game Management in Montana.
Montana Fish and Game Department, Helena, USA. Notes: In Chapter 15, Sharp-
tailed grouse, R. L. Brown notes that "...snow, rain, wind and temperature extremes
greatly influence the welfare of grouse, depending of course on condition of the habitat
sheltering grouse from the elements. Strong prairie winds and snow are an ever
changing stress in the winter. By roosting behind rough topography or in deep snow,
sharp-tails find protection from the weather as well as predators. In Chapter 18,
Pheasants, the authors write Winter is a critical time for pheasants. Artificial feeding,
however, is not justified, since the birds seldom need additional food. When they do,
feeding concentrates them and exposes them to diseases, predators and highway
traffic. If feeding must be stopped during a critical period, the pheasants are less likely
to survive than if they had not been fed at all. Heavy losses occur during severe
blizzards. Birds caught without suitable cover face the storm. Their nostrils and mouths
fill with ice and they suffocate. Sometimes whole flocks in small cover patches are
buried under deep drifts. After a severe winter, hens in poor condition may die because
of the additional stress imposed by nesting. Recovery of the population after these
losses is usually rapid if food and cover are adequate. Chapter 19, Hungarian Partridge,
the authors note: "Factors responsible for... losses include adverse weather.." among
other factors. "Optimum conditions for a flourishing population of Hungarian partridge
are a cool, moderately dry climate and mixture of cultivated and non-cultivated land."
Chapter 21 , Chukar Partridge - "While the chukar successfully inhabits areas of sparse
rainfall, they are heavily dependent on free water within close proximity. Periods of
drought or low-water levels will cause birds to cluster around available water sources.
Perhaps the most critical limiting factor is snow conditions during the winter months.
Populations may be seriously affected by heavy or prolonged snow covering feeding
and roosting areas. In many localities where chukars initially survived and bred, severe
winters gradually eliminated them. Keywords: sharp-tailed grouse, pheasant,
Hungarian partridge, chukar partridge, snow, temperature, drought, mortality,
habitat use, behavior, topography

350

Myhre, K., M. Cabanac, and G. Myhre. 1975. Thermoregulatory behavior and body
temperature in chicks of willow grouse (Lagopus lagopus lagopus). Poultry
Science 54:1174-1179. Abstract: Cloacal temperatures (Tcl) of outdoor living captive
willow grouse chicks {Lagopus lagopus lagopus) were found to increase from 39.4 ±
0.5°C the first day to 40.3 ± 0.5°C the twelfth day after hatching. Average Td of adults
was 40.7 ± 0.3°C. When left alone for 30 min. in a controlled test environment providing
temperatures ranging from 21 °C to 46°C, the one-day-old chicks preferred significantly
higher ambient temperatures than eight-day-old birds. Td was significantly lower in the
chicks tested the day after hatching than in the older chicks. It is concluded that the
chicks' thermoregulatory behavior is essential for maintaining homeothermia, and that
the birds' thermoregulatory set-point is low the day after hatching and climbs to adult
level during the first week. Key words: willow grouse, temperature,
thermoregulation, behavior

158 jx

4>

>'> te **

cCi

351

Myrberget, S. 1988. Demography of an island population of willow ptarmigan in
northern Norway. Pages 379-419 in A. T. Bergerud and M. W. Gratson, editors.
Adaptive strategies and population ecology of northern grouse. Wildlife
Management Institute, Washington D.C. and University of Minnesota,
Minneapolis, USA. Notes: Page 401 — Failure of eggs to hatch. There was a strong
tendency, however, for hatching failure to be large in years with much snow remaining
in May and delayed egg-laying... Chick Mortality — For 7 years Myrgerget et al found that
the annual survival of chicks was significantly correlated with the mean temperature
before hatching... no significant relationship was found between the chick survival rate
and the mean ambient temperature during the 10 days following the average hatching
date. ..Page 408 — In spring there is a gradual shift in the diet of ptarmigan on Tranoy
from woody plants to evergreen heath species and finally to herbs. Availability of the
plants forming the ground layer to tetraonids, and their chemical composition, are
affected by the date of snowmelt and by the weather in the spring, factors that vary in a
random manner...Page 411 — Environment of chicks. Cold and rainy weather reduce
the available time in which small ptarmigan chicks are able to feed, because they are
dependent on the hen for warmth. Cold weather may also reduce the availability of
insects that chicks eat. But no relationship existed between the chick mortality rate and
the air temperature just after hatching. This agrees with results obtained for other
Lagopus populations. In some areas, however, extremely unfavorable weather
conditions, such as snow or sleet showers, have resulted in the death of many
ptarmigan chicks. ..On Tranoy, a relationship was found between the weather before
hatching and the mortality rate of chicks, in agreement with Slagsvold's hypothesis.
However, a close relationship between the amount of insects during the early stages of
the life of the chicks and mortality rate was not apparent.. .Page 412 — At Tromso,
annual variations in the survival of willow ptarmigan chicks kept in captivity were
apparently caused by variations in plant phenology, but in such a way that more chicks
died when the spring was early and warm, because of an early onset of the lignification
of bilberry leaves. In some years there may also have been a shortage of certain
vitamins in the food available to chicks... the relationship between the spring weather,
the chicks' food, and survival rate... would seem more complex than suggested by
Slagsvold...Page_414 — My conclusion is that the poor productivity of ptarmigan in vole
crash years occurs because the predator situation, the quality of the food supply, and
the weather in the chick period are likely to be unfavorable in such years. Key words:
willow ptarmigan, snow, temperature, reproduction, nutrition, vegetation,
productivity

352

, R. Blom, and K. E. Erikstad. 1977a. Effect of a bad production year for

willow grouse on the size of the breeding population the following year. Fauna
30(2):88-97. Abstract: Data are given on the number of breeding Lagopus lagopus in
1975 and 1976 in the Tranoy study area, northern Norway. The total autumn population
in 1975 and the breeding population in 1976 (17 pairs/km2) were less than in any year
since 1960. The winter mortality rate for adult grouse hens in 1975-1976 was about

H <C# t*

4

cC±

average for the area, 42.5%. The winter loss of immature birds was estimated at 83%,
which is slightly higher than the average (75%), but in fact, the value for this particular
winter is somewhat imprecise. The possible higher rate of mortality of immatures in the
winter of 1975-1976 may have been due to the particularly poor physical condition of
the young grouse in the autumn of 1975 as a result of an extremely unfavorable
environment for chicks that summer. Key words: willow grouse, reproduction,
severe winter, mortality, nutrition, chick survival

353

, K. E. Erikstad, and T. K. Spidso. 1977b. Variations from year to year in

growth rates of willow grouse chicks. Astarte 10(1):9-14. Abstract: Data are given
for increase of total weights and wing lengths of chicks of willow grouse, Lagopus
lagopus, for 7 years in a study area in northern Norway. There was a significant
difference in growth rates for different years and a positive correlation between
variations in annual growth rates and survival rates of the chicks. High temperatures
before hatching had a positive influence on the growth and survival of the chicks. The
weather during the early chick period may modify growth rates for small chicks. Key
words: willow grouse, temperature, chick survival

354

, C. Norris, and E. Norris. 1975. Grit in Norwegian Lagopus spp. Norwegian

Journal of Zoology 23:205-212. Abstract: There was found to be a decrease from
summer to winter in the quantity and weight of grit, and an increase in the size of the
individual stones, from Lagopus lagopus and L mutus in Norway. These changes are
related to changes in the snow cover. In chicks, the size, roundness, weight, and the
amount of grit were all found to increase with age. The grit consisted of hard minerals,
poor in calcium. Key words: willow grouse, snow, grit

355

Naugle, D. E., C. L. Aldridge, B. L. Walker, K. E. Doherty, M. R. Matchett, J.
Mcintosh, T. E. Cornish, and M. S. Boyce. 2005. West Nile Virus and sage-grouse:
what more have we learned? Wildlife Society Bulletin 33(2):616-623. Abstract:
Evidence suggests that risk of infection was low in 2004 because unseasonably cool
summer temperatures delayed or reduced mosquito production. Moreover, mortalities
occurred 2-3 weeks later in 2004 than in 2003, and the shift to later timing was
consistent between years at sites where WNv reduced survival both years. Key words:
sage-grouse, disease, temperature, West Nile Virus

356

Neave, D. J. and B. S. Wright. 1969. The effects of weather and DDT spraying on a
ruffed grouse population. Journal of Wildlife Management 33(4):1015-1020.

Abstract: A study of population dynamics illustrated the effects of temperature and
pesticide spraying on a ruffed grouse (Bonasa umbellus) population in central New
Brunswick. May and June temperatures were related to the date of nest initiation, to a
partially developed egg loss, to fall age ratios and to annual kill fluctuations. A

160 j^

4>

^ fete >f*

synergistic effect of chemical spraying with DDT (in oil at the rate of 0.5 lb/0.7gal/acre or
0.25/0.5gal/acre) and temperature is suggested in the partially developed egg loss.
Spraying was also correlated with immature survival as significant differences were
found in June brood sizes in sprayed and unsprayed areas and in fall age ratios,
suggesting complete and partial brood losses. Key words: ruffed grouse,
temperature, population dynamics, habitat manipulation

357

Nelson, O. C. 1955. Afield study of the sage-grouse in south-eastern Oregon with
special reference to reproduction and survival. M. Sc. Thesis, Oregon State
College, Corvallis, USA. Notes: Page 53 — "Comparable weather data was recorded
for the study year 1954. Total precipitation for April was .62 inch, part of which was in
the form of snow amounting to 2.0 inches in depth. Total precipitation for May was 1.25
inches, part of which was 3.0 inches of snow. The month of April was mild with no
critical storms. The temperature for this month never dropped below 15°F and had a
29.3° average minimum. The highest temperature recorded for the month was 71°, the
average maximum being 56.3°. Thus, weather conditions during April favored a
successful nesting period for sage-grouse.... In making a comparison of April and May
of 1954 with the same months for the years 1941 to 1953, it appears that the weather
conditions for the study year were much more conducive to successful nesting on the
basis of more mild conditions with less extremes in temperature." Page 82 — "Weather
conditions, as they affect brood survival, are frequently given as causes for losses. On
May 29, shortly after the peak of hatching, .70 inch of rain and one inch of snow, sleet,
and hail fell on the study area. This was accompanied by strong winds and cold
temperatures. On June 5, similar conditions occurred. No observations were made or
data collected that indicated any harmful effect of such weather conditions on brood
survival. Brood counts following such critical periods showed no noticeable decrease in
brood sizes. The species inhabits areas exhibiting extremes in temperatures and
weather conditions; however, the brooding instincts of the the sage-grouse appear
strongly developed. Thus, it seems likely that losses are at a minimum in this respect."
Page 90 — "Causes for... movement into and out of the meadow areas appeared to be
the result of a combination of factors. On June 10, 1.18 inches of rain fell. This was the
last appreciable amount of precipitation for the remainder of the summer. It is believed
that the sage-grouse began to move into the more preferable meadow areas because
of the shortages of water and green foods from most of the area as conditions became
drier. The meadows had much to offer the birds in the area at this time such as water,
greens, and a plentiful supply of insects for the young in particular. However, in the
absence of any rainfall, these benefits gradually disappeared." Key words: sage-
grouse, precipitation, snow, temperature, sleet, hail, wind, mortality, nutrition,
movement, habitat use, productivity

358

Nestler, R. B. 1946. Vitamin A, vital factor in the survival of bobwhites.
Transactions of the North American Wildlife Conference 11:176-195. Notes: "Of 45
wild quail shot or trapped this winter in Pennsylvania, Maryland, Virginia and Alabama,
at least 31 per cent, or nearly one third, had not enough vitamin A stored in their livers

X l^V t . L- 161

% )->f ^> ^c

**

to help them survive more than 4 weeks of a deficiency of this factor in their diet.
Imagine the effect of a thick blanket of snow lasting for a long period. Most of all,
sources of vitamin A would be made unavailable. While carnivores can obtain their
vitamin A from the tissues of other animals, quail, in winter at least, depend largely on
carotene in plants. Unless they have stored away a plentiful supply in their livers their
case would become desperate." "Less carotene is manufactured in plants during a hot,
dry period than during a cool, wet season. Combine a droughty summer and fall with a
severe winter of prolonged snow, as was the case in Ohio in 1944-45, and the result is
an ideal condition for vitamin A deficiency. Birds will die either directly or indirectly from
avitaminosis. Weakness, impaired eyesight, lack of alertness, and loss of speed causes
them to succumb quickly to predation and severe weather. Thus, submarginal, or even
marginal, intakes of vitamin A, while sufficient to keep birds from dying from
avitaminosis, yet many underline their constitutions to such an extent that death results
anyway from other causes. Also death of a part of a covey from vitamin A deficiency
may so reduce the size of the group; that the survivors may perish from lack of
protection from cold during huddles. Key words: bobwhite quail, nutrition, snow,
drought, vegetation, predation, temperature, disease

359

, J. B. DeWitt, and J. V. Derby Jr. 1949. Vitamin A storage in wild quail and

its possible significance. Journal of Wildlife Management 13(3):265~21. Summary:
Research on penned bobwhite quail concluded that during winter especially, Vitamin A
apparently stands on par with carbohydrates as a factor that could decide the fate of
wild quail and might be a very important factor in population fluctuations in quail. In
winter quail depend largely on carotene in plants. Unless they have stored away a
plentiful supply in their livers, their chances for survival under adverse weather
conditions are much reduced. Less carotene is manufactured in plants during a hot dry
period than during a cool wet season. Drought-affected plants are lower in carotene
than plants grown under normal weather conditions. Combine a droughty summer and
fall with a severe winter of prolonged snow, as was the case in Ohio during 1944-45
and Iowa during 1936-37 (an estimated quail loss of 88%), and the result seemingly is
an ideal condition for vitamin A deficiency. Birds may die either directly or indirectly from
avitaminosis. Weakness, impaired eyesight, lack of alertness, and loss of speed cause
them to succumb to predation and severe weather. Key words: bobwhite quail,
nutrition, disease, mortality, predation, drought, temperature, vegetation

360

Nguyen, L. P, J. Hamr, and G. H. Parker. 2003. Survival and reproduction of wild
turkey hens in central Ontario. Wilson Bulletin 115(2):131-139. Abstract: Recent
success of Eastern wild turkey (Meleagris gallopavo silvestris) reintroductions across
southern Ontario has prompted wildlife managers to investigate the potential of
extending the northern limit of this subspecies' range. We monitored the survival and
reproduction of introduced wild turkeys on the Precambhan Shield in central Ontario
during 1999-2001. Mean annual survival of 39 radio-tagged hens was 0.288±0.057 SE.
Summer and winter survival rates differed between the first and second years of the
study. Spring and fall survival rates did not differ significantly between years.

162 _>,

4-

>->' (& **

cCb

Reproductive parameters that characterized the population included a nesting rate of
0.588, mean clutch size of 10.0 eggs/nest success of 0.500, hatching rate of 0.81, hen
natality rate of 1.18 female hatched/female, poult survival of 0.54, and fall recruitment of
0.63 juvenile females/breeding hen. Success of the pilot wild turkey introduction in
central Ontario was compromised by high predation, low numbers of introduced birds,
and a prolonged period of deep snow during 2000-2001. Key words: turkey, snow,
mortality

361

Norman, G. W., T. M. Fearer, and P. K. Devers. 2004. Factors affecting wild turkey
recruitment in western Virginia. Proceeding of the Annual Conference Southeast
Association Fish and Wildlife Agencies 58:248-262. Abstract: Annual recruitment of
eastern wild turkeys (Meleagris gallopavo silvestris) should be closely monitored to
regulate fall turkey seasons and reduce risk of over-harvest. However, previous studies
have not encompassed the spatial or temporal scales needed to produce models that
can consistently predict recruitment over a large region. Our objective was to assess
the ability of using long-term data sets of sex-age ratios, oak (Quercus spp.) mast, and
weather variables to forecast annual wild turkey recruitment in western Virginia. We
conducted a thorough literature search on factors believed to be limiting reproduction
and developed a series of 14 a priori models and 1 a posteriori model to predict
recruitment. We used fall harvest ratios of juveniles per adult female, averaged over 26
western Virginia counties, during 1973-2002 as an index to annual recruitment and
investigated the relationship of recruitment to age structure of the population, oak mast
production in the previous fall, and spring weather. We considered impacts of different
weather severity measures and investigated effects of deviation from mean, 90%, and
75% quartile values on recruitment. Our best model (u>9=0.812) predicting recruitment
incorporated May and June rainfall and March temperatures at the 75% quartile scale.
This model accounted for a significant amount of variation in recruitment residuals
(R2=0.50, R2ajd=0.44). Monitoring these selected weather parameters offers managers
the ability to predict significant changes in recruitment annually. Key words: eastern
wild turkey, precipitation, temperature, recruitment, production, model

362

Northrup, R. D. 1991. Sharp-tailed grouse habitat use during fall and winter on the
Charles M. Russell National Wildlife Refuge, Montana. M.S. Thesis, Montana State
University, Bozeman, USA. Notes: Pages 33-34 — "In this study, birds were active on
display grounds through January 1990 of winter 2; however, display did not occur
during January or February 1989. Snow accumulations and/or cold air temperatures
apparently prevented such activity until early March that year." Page 34-"Grouse
habitat selection was also affected at least in part by weather conditions. Heavy snow
and/or cold temperatures presumably forced grouse to use the juniper type almost
exclusively for cover and food during winter 1 . Of the cover types used by sharptails in
this study, juniper undoubtedly provided the best security cover and shelter from harsh
winter weather." Page 38 — "A number of sharptails roosts during winter 1 consisted of
snow burrows and shallow snow depressions. Fresh drifts created from wind and large
junipers or broken topography typically provided sufficient snow depths for

i f>"^^

163

<£5& -"car- <£Z^h> c£^b

burrowing... as fresh drifts became available, grouse immediately switched to burrows.
However, because snow developed a crust within days of a snowfall, depression snow
roosts were the most common." Page 40 — "Sharptail management objectives should
include maintaining and/or enhancing shrub cover which provides protection and food
especially during periods of harsh winter weather." Key words: sharp-tailed grouse,
snow, temperature, wind, topography, behavior, habitat use, vegetation, habitat
manipulation

363

Norton, M. A. 2005. Reproductive success and brood habitat use of greater prairie
chickens and sharp-tailed grouse on the Fort Pierre National Grassland of central
South Dakota. M.S. Thesis, South Dakota State University, Brookings, USA.

Notes: Chick survival was higher in 2005 than the previous two years, probably
attributable to increased precipitation and warmer temperature patterns. PC broods
utilized habitats with a mean vegetation height of 32 cm (n=8), and with ground cover of
18% grass and 10% forbs in 2004. During 2005, PC broods used habitats with a mean
vegetation height of 38 cm (a?=10) and 18% grass and 5% forb ground cover. ST broods
used habitats with a mean vegetation height of 43 cm during both 2004 (n=8) and 2005
(n=9). High nesting success and survival rates found during this study are likely
attributed to the vast grassland ecosystem and the implemented rotational grazing
regime, both of which had multiple positive influences. Predators had to spend more
time searching to find a nest or bird to prey on in larger unfragmented grasslands. The
large expanse of grassland is maintained by a grazing regime that provided an ample
food source of invertebrates and a mosaic of habitats needed for brood rearing hens.
Key words: greater prairie chicken, sharp-tailed grouse, temperature,
precipitation, vegetation, nutrition, predation, habitat manipulation

364

Novoa, C, A. Besnard, J. F. Brenot, and L. N. Ellison. 2008. Effect of weather on
the reproductive rate of rock ptarmigan Lagopus muta in the eastern Pyrenees.
Ibis 150(2):270-278. Abstract: Understanding the effects of climate on avian life history
traits is essential if we wish to predict the demographic consequences of expected
climatic changes. We investigated the influence of weather conditions on the
reproductive success of rock ptarmigan Lagopus muta in the eastern French Pyenees,
one of the southernmost areas inhabited by the species. Reproductive success was
estimated in early August between 1997 and 2006 by counting adults and well-grown
chicks with pointing dogs. The number of young per adult varied from 0.08 to 0.72.
Using Poisson regression and Akaike's information criterion, we selected the best
model explaining the effect of weather (date of snowmelt and, for both laying/incubation
and post-hatching periods, mean minimum and maximum temperatures, monthly rainfall
and number of days with rain) on the proportion of young in August. Reproductive
success was positively associated with early appearance of snow-free ground, and date
of snowmelt alone was the model that best explained annual variation in reproductive
success. Other models, which included a negative effect of rainfall, particularly after
hatching, also had some support. Hence, both pre-laying and post-hatching weather
conditions influenced reproductive success of rock ptarmigan in the eastern French

164 j^

4>

H fe&r >f*

c^>

Pyrenees. On a continental scale, reproductive success of alpine populations of rock
ptarmigan is consistently lower than that of northern populations. This difference in
productivity may be partly correlated with climatic conditions observed along an arctic-
alpine gradient, the amount and variation of rainfall being greater in southern alpine
areas than elsewhere in the species' range. Key words: rock ptarmigan, rain, snow,
fecundity, productivity

365

Oakleaf, R. J. 1971. The relationship of sage-grouse to upland meadows in
Nevada. M. S. Thesis, University of Nevada, Reno, USA. Unable to obtain for
abstract.

366

Oberlag, D. F., P. J. Pekins, and W. W. Mautz. 1990. Influence of seasonal
temperatures on wild turkey metabolism. Journal of Wildlife Management
54(4):663-667. Abstract: We measured standard metabolic rate (SMR) and the effects
of temperature on metabolism of adult eastern wild turkeys (Meleagris gallopavo
silvestris) during each season of the year. Female SMR was similar throughout the
year, and male SMR was lower during spring. Female SMR exceeded male SMR each
season and was significantly (P < 0.05) higher during summer and autumn. Both sexes
had similar lower critical temperatures (Tk), but females lost heat faster below Tk. Low
winter Tk (about -15C) indicated that thermoregulatory costs are probably minimal for
turkeys at normal winter temperatures in south-central New Hampshire. Key words:
Merriam's turkey, temperature, metabolism, thermoregulation

367

Osmolovksya, V. I. 1966. Reasons for changes in the number of tetraonid birds.
Byul. Mosk. Obshchest Ispyt prir Otd Biol 71(5):61-70. Abstract: There are different
causes that affect the changes in the number of tetraonids (Tetraonidae) in different
geographic zones. On the basis of 733 questionnaires, it has been established that cold
springs are of decisive importance in northern taiga environments. Further south the
first role belongs to the anthropogenic factors of mass deforestation, immoderate
hunting, agriculture and forestry methods unfavorable for game birds and forest fires in
sparsely populated areas of Siberia. Populations of different species of tetraonids react
to certain changes in the environmental conditions in a varying way. The number of
capercaillie (Tetrao urogallus L.) decreases mainly owing to deforestation, while the
black grouse (Lyrurus tetrix L), which inhabits more open biotopes, is affected by a
much greater number of factors (changes in weather, activity of the beasts of prey,
grazing of cattle, poisoning by chemicals, etc.). The number of hazel hen (Tetrastes
bonasia L.) is under the influence of cold springs and deforestation. Tetraonids, as an
object of hunting, can be preserved only under a reasonable combination of the
interests of forestry, agriculture and hunting. Key words: capercaillie, black grouse,
hazel grouse, temperature, population dynamics

*

I Kfc***

165

<9^ ^ar- <£^> ^

368

Overton, C. T., R. A. Schmitz, and M. L. Casazza. 2005. Post-precipitation bias in
band-tailed pigeon surveys conducted at mineral sites. Wildlife Society Bulletin
33(3):1 047-1054. Abstract: Many animal surveys to estimate populations or index
trends include protocol prohibiting counts during rain but fail to address effects of rainfall
preceding the count. Prior research on Pacific Coast band-tailed pigeons {Patagioenas
fasciata monilis) documented declines in use of mineral sites during rainfall. We
hypothesized that prior precipitation was associated with a short-term increase in use of
mineral sites following rain. We conducted weekly counts of band-tailed pigeons at 19
Pacific Northwest mineral sites in 2001 and 20 sites in 2002. Results from regression
analysis indicated higher counts < 2 days after rain (11. 31 ±5. 00% [mean±SE])
compared to > 3 days. Individual index counts conducted < 2 days after rain were
biased high, resulting in reduced ability to accurately estimate population trends.
Models of band-tailed pigeon visitation rates throughout the summer showed increased
mineral-site counts during both June and August migration periods, relative to the July
breeding period. Our research supported previous studies recommending that mineral-
site counts used to index the band-tailed pigeon population be conducted during July.
We further recommend conducting counts > 3 days after rain to avoid weather-related
bias in index estimation. The design of other population sampling strategies that rely on
annual counts should consider the influence of aberrant weather not only coincident
with but also preceding surveys if weather patterns are thought to influence behavior or
detection probability of target species. Key words: band-tailed pigeon, census,
precipitation

369

Palmer, W. E., S. R. Priest, R. S. Seiss, P. S. Phalen, and G. A. Hurst. 1993.
Reproductive effort and success in a declining wild turkey population.
Proceedings of the Annual Conference Southeast Association of Fish and
Wildlife Agencies 47:138-147. Abstract: We monitored reproductive effort and success
of a wild turkey (Meleagris gallopavo) population on a public wildlife management area
in Mississippi for 9 years using telemetry and other indices. Adult hens (A/=143) had a
nesting rate of 72.7% (range 54-100) and was greater (P=0.0001) than the nesting rate
(26.7%) of juvenile hens (A/=15). Annual nest success of 104 nests of adult hens
averaged 30.8% (range 0-62) and poult survival to >50 days was 22.7% for 27 broods
(A/=203 poults). Clutch size averaged 9.1 (SE=0.54) and 6.7 (SE=1.1), and hatchability
was 93% and 100%, for first and second clutches, respectively. During this period,
turkey population estimates and indices (gobbler harvest, hen ancillary observations)
declined 250%-350%. Low reproduction was due primarily to high predation of nests
and poults and appeared to have caused the population decline. However,
environmental factors (i.e., food, rainfall) appeared to significantly impact reproduction
in some years. Reproduction was greatest in 1992 following a distemper outbreak
among some nest/poult predator populations in 1991. Key words: wild turkey,
precipitation, reproduction

166 j^

4>

H ^ **

370

Panek, M. 1992. The effect of environmental factors on survival of grey partridge
(Perdix perdix) chicks in Poland during 1987-89. Journal of Applied Ecology
29:745-750. Summary: I.The survival of grey partridge chicks was estimated from the
brood sizes in several areas of Poland, and related to weather conditions, abundance of
insect food, occurrence of weeds, and the use of herbicides. 2. Geometric mean brood
size in different areas and years ranged from 7.5 to 1 1.2, with a mean of 9.3, and the
estimated chick survival rate ranged from 42 to 71%, with a mean of 56%. 3. Chick
survival rates increased with mean temperatures and decreased with increasing
numbers of rainy days in June. 4. Chick survival rates increased with numbers of plant
bugs, but they had no significant effect after removal of the effect of weather. Numbers
of plant bugs increased with temperature. 5. Density and species diversity of
dicotyledonous weeds in cereals were relatively high, and difference in weeds between
areas were not in accordance with differences in the abundance of insects selected by
partridges. The application of herbicides in Poland so far has not had a marked indirect
negative effect on the survival of partridge chicks. Presumably the occurrence of
permanent cover in crop fields has a positive effect on chick survival. Key words: grey
partridge, temperature, insects, nutrition, precipitation, vegetation, habitat
modification, chick survival

371

. 1990. Factors influencing winter mortality of gray partridge in western

Poland. Pages 304-314 in K. E. Church, R. E. Warner, and S. J. Brady, editors.
Proceedings Perdix V: gray partridge and ring-necked pheasant workshop,
Minnesota Department of Natural Resources. Abstract: I surveyed gray partridge
{Perdix perdix) coveys near Czempin, Poland twice a month during December through
February, 1985-1988 (n = 225 surveys). Mortality rate per half a month was calculated
from a decrease in covey sizes. Mortality was lower when there were 2 to 7 cm of soft
snow than in the absence of snow (P < 0.01 ). However, it was higher during periods of
hard snow cover than soft snow(P< 0.001) and in the absence of snow (P< 0.001).
The rate of mortality increased when the depth of hard snow increased from 0 to 10 cm.
No correlation was found between mortality rate and air temperature, but in the
presence of hard snow MR increased with declining air temperature in the preceding
half-month period. Mortality was higher on large than on small crop fields (P < 0.05). A
negative correlation was found between mortality rate and the distance of coveys from
the forest. The rate of mortality was higher when densities of coveys exceeded the
mean value than when it was below the mean (P < 0.01). Total winter mortality
averaged 35%, and ranged from 32 to 42%. Key words: gray pargridge, snow,
temperature, mortality

372

Parmalee, P. W. 1955. Some factors affecting nesting success of the bobwhite
quail in east-central Texas. American Midland Naturalist 53(1):45-55. Abstract:
Locally bobwhite (Colinus virginianus) abundance is controlled by conditions within the
immediate habitat, primarily adequate food and cover, although a large number of
additional factors are continually affecting quail numbers to a somewhat lesser degree.

i &*&**

167

££$> cCb

Geographically, bobwhites appear to be limited by extremes in temperature and
precipitation and by such factors that accompany each of these conditions. Climate, in
addition to its geographic influence, is of such significance as to produce periodic or
seasonal fluctuations in quail numbers in regions where bobwhites have successfully
established themselves and are filling a particular ecological niche. The influence of
climate on bobwhite populations in the post oak region during 1950 and 1951 has
become evident from data based on temperature and precipitation records, nesting
success, and sex and age ratios. Key words: bobwhite quail, temperature,
precipitation, distribution, productivity, sex ratio, age ratio

373

Paterni, M. J. 1979. Winter habitat selection by female spruce grouse (Canachites
canadensis franklinii) in a mixed coniferous forest. M. S. Thesis, University of
Montana, Missoula, USA. Abstract: Captured spruce grouse were fitted with solar
powered radio transmitters and/or coded leg bands. The movements, areas used, social
groupings, and food habits of radioed and banded birds were monitored during the fall
and winter seasons. Meteorological conditions were monitored to evaluate the effects of
extreme temperatures, winds, and snow conditions on habitat selection of grouse. Sites
used by grouse were analyzed to determine the horizontal and vertical structure of
vegetation. Areas utilized by observed grouse were compared to unused available
areas on the basis of vegetation species composition, basal area, relative density,
canopy cover, and the vertical distribution of foliage. The topographic diversity of spruce
grouse habitat in the intermountain west influences not only vegetation community
structure and distribution, but also thermal and radiant energy regimes. Over the winter
period spruce grouse activity is concentrated within areas subject to tolerable
microclimatic conditions. Ridges, upper slope positions, and southerly aspects are
selected over drainage bottoms and northerly aspects. In coincidence with the
physiographic features selected, certain forest vegetation communities with distinct
structures (horizontal and vertical), and furthermore, specific sites exhibiting distinct
structures were selected for daytime activities and night roosting. Needles of ponderosa
pine (Pinus ponderosa) were major food consumed by spruce grouse during the winter
period." On pages 34-35, Paterni states "...short movements within activity areas were
apparently influenced by changing weather. Wind conditions appeared to be the major
factor stimulating movement. Radio-marked spruce grouse exposed to winds sufficiently
intense to ruffle the bird's feathers often flew to sites protected from the wind. Given
equivalent intensities, gusty winds did not stimulate movement to the same extent as
constantly flowing winds. The longest movement witnessed under gusty wind conditions
(n=3) was approximately 250 m (813 ft). In most cases spruce grouse were able to find
suitable protection from the wind by shifting their position within the roost tree or clump
of trees being used. Precipitation and fluctuating temperatures did not apparently
influence spruce grouse movement during the feeding-loafing period beyond limited
position adjustment within a given tree. During periods of precipitation spruce grouse
often roosted beneath clumps of needles in the roost tree, and exhibited no difficulty in
finding adequate cover at feeding-loading sites. On clear, cold days wintering spruce
grouse appeared to take advantage of direct solar radiation by roosting at positions on
or in the canopy of the roost tree which was directly exposed to the sun's rays. On
warm, clear days, the grouse often perched in the shade of clumps of needles, the bole

168

3/
4

^ fcfc **

of the tree, or other tree canopies." Key words: spruce grouse, temperature, solar
radiation, wind, behavior, habitat use, movement

374

Patten, M. A., D. H. Wolfe, E. Shochat, and S. K. Sherrod. 2005. Effects of
microhabitat and microclimate selection on adult survivorship of the lesser
prairie-chicken. Journal of Wildlife Management 69(3): 1270-1 278. Abstract:
Populations of the lesser prairie-chicken (Tympanuchus pallidicinctus), an endemic
grouse of the south-central United States, have declined precipitously. This species
occurs in short- and mixed-grass prairies with sandy soils. Apart from perennial grasses
of short stature, prairie-chicken habitat is characterized by dryland shrubs of the sand
shinnery community, particularly the shinnery oak (Quercus havardii) and sand
sagebrush (Artemisia filifolia). We measured microhabitat and microclimate
characteristics at bird-centered and random points at the southwestern (New Mexico)
and northeastern (Oklahoma) edges of the species' range. We estimated survival by
locating radio-tagged prairie-chickens (n - 544) from April 1999 to June 2003. We found
that lesser prairie-chickens used sites within the sand shinnery community that had a
higher cover and greater density of shrubs (ANOVA: P<0.0001). Microclimate differed
substantially between occupied and random sites (MANOVA: P<0.001), and prairie-
chicken survival was higher in microhabitat that was cooler, more humid, and less
exposed to the wind. Survivorship was higher for adults that chose microhabitat with a
higher cover of shrubs and grasses and a higher density of vegetation. Survivorship
was higher for prairie-chickens that used sites with >20% cover of shrubs than for those
choosing 10-20% cover; in turn, survivorship was higher for prairie-chickens choosing
10-20% cover than for those choosing <10% cover (Cox regression: P<0.05). Whereas
vegetation may recover following moderate habitat disturbance, land managers
applying herbicides or otherwise removing shrubs should understand the potentially
negative effects of reduced shrub cover on adult survivorship of lesser prairie-chickens.
Key words: lesser prairie-chicken, temperature, humidity, wind, shelter, habitat
use, habitat modification, mortality, microhabitat, microclimate, predation

375

Patterson, R. L. 1952. The Sage-grouse in Wyoming. Wyoming Game and Fish
Commission. Sage Books, Inc., Denver, USA. Summary: Page 54 — "The nature of
the environment has placed heavy demands upon the physiological system of the sage-
grouse. Long winters, dry summers, high elevations, heavy snowfall and ground
blizzards all have provided critical hazards in the search for food, shelter and water."
Page 55 — "Snow conditions naturally altered the ground feeding patterns. In winter,
birds may be forced to feed on the exposed tips of sagebrush branches or compelled to
pick the leaves from small prostrate bushes on windswept ridges. In Utah sage grouse
were documented digging holes in the snow to reach the tips of sagebrush leaves.
Observations in Wyoming have revealed the importance of antelope in making snow-
covered sagebrush available to sage chickens as a result of digging for their own food
supply." Page 67 — "Evidence indicated hens retired to the foothills region and more
abundant water supplies during unusually dry seasons. On the basis of their behavior, it
was felt that a series of drought years would likely telescope sage-grouse populations

% f>"^^

169

into areas less likely to exhibit the effects of continued drought as expressed in the
vegetation and available surface water. It would, in effect, not only result in a restriction
of sage-grouse distribution, but would cause a noticeable decrease in numbers on sub-
marginal ranges." Page 72 — "Another outbreak of coccidiosis occurring among sage-
grouse in Wyoming in 1939. This loss, which began in June and continued until
September, again involved young birds. It was his (Scott) opinion that the drought years
concentrated the grouse on available water, thereby creating a situation favorable to the
transmission of the coccidium." Page 115 - Weather and its effect upon nesting and
brood survival - "Hatching success in 1949 and 1950 was 92.1 per cent and 91.4 per
cent, respectively. No nests under observation were deserted or failed to hatch as a
result of low temperatures or snow conditions which prevailed during the 1950 nesting
season, even though weather of this nature generally is supposed to be extremely
unfavorable to successful sage-grouse nesting seasons." Page 118 - "Juvenile
mortality was high (during this period) due to several consecutive days of cold rain,
sleet, and snow, accompanied by low temperatures." Page 138 -"Hail storms occur
frequently in parts of Wyoming and may at times kill young birds over local areas.
Weather apparently exerts no serious effects upon sage-grouse once they have passed
the vulnerable early stages of their growth." Page 152 - "On days with strong winds,
rain, snow storms, and low barometric pressure, the activity of birds was greatly
reduced and many failed to appear on the strutting grounds at all... maximum strutting
activity, accompanied by the largest number of cocks on the grounds, occurred on
mornings with clear, fair weather, irrespective of low temperatures and the amount of
snow on the ground." Page 179 - "Local residents have reported to me that sage-
grouse on occasion dig into deep-crusted snow and roost in these open holes." Page
184 - "Honess reported the number of sage chickens wintering in the (Jackson Hole)
area was related to snow depths which, of course, regulated the amount of exposed
sagebrush." Page 189 - "Movements of sage-grouse onto their winter ranges was
correlated with snow conditions and the availability of sagebrush. In the absence of
open water, snow furnished the moisture requirements for sage-grouse in the winter."
Page 208 - "Sage-grouse have also been observed to regularly visit partially frozen
streams in Eden Valley during the late fall months in order to drink through holes in the
ice." Key words: sage-grouse, wind, rain, snow, barometric pressure, drought,
behavior, mortality, parasites, disease, nutrition, reproduction

376

Pedersen, A. 0., 0. Overrein, S. Unander, and E. Fuglei. 2005. Svalbard rock
ptarmigan (Lagopus mutus hyperboreus): a status report. Report No. 125,
Norwegian Polar Institute, Norway. Notes: Page 14 — "Climate: The large fluctuations
in numbers seen in most ptarmigan populations seem to be caused by a combination of
biotic factors and climatic variation. The systematic component of the fluctuation (i.e.
cycles) most likely originates from biotic interactions (self regulation through e.g. social
behavior, predator-prey or host-parasite interactions). Also regular fluctuation in sun-
spot activity has been invoked as cause of cycles in some northern vertebrate species.
In willow ptarmigan and rock ptarmigan climate events like bad weather during the
breeding season compromising breeding success is thought to be a vital factor in
synchronizing the population dynamics at different locations, often called the "Moran
effect"... "Poor climate conditions as heavy snow fall during egg laying caused delayed

170 ^

^

f V ^, ^

hatching in Svalbard rock ptarmigan, but how important such conditions are in shaping
population dynamics is at the present unknown." Key words: rock ptarmigan, snow,
sun spots, reproduction

377

Pedersen. H. Chr. and J. B. Steen. 1979. Behavioral thermoregulation in willow
ptarmigan chicks Lagopus lagopus. Ornis Scandinavica 10(1):17-21. Summary:
The effect of behavior on the Tb of ptarmigan chicks was studied in captive and wild
birds of different ages and at different ambient temperatures (Ta). Newly hatched chicks
show inter-brood coordinated periods of brooding and browsing. During browsing Tb
decreases; during brooding it returns to normal. At Ta from 7 to 10°C, the lowest
recorded value was 34.3°C. Post browsing Tb showed little variation with Ta but
increased with age. No hypothermia was found in 3-week-old chicks. The duration of
each browsing period increased both with increasing Ta and with age. If prevented from
brooding, the chicks start to utter distress calls when their Tb drops below 34°C and they
become lethargic at Tb below 32°C. The sensory input which triggers the chicks' return
to brooding (the hen) appears to be the decreased Tb rather than signals from the hen.
Ta near freezing appear to be a limiting factor for ptarmigan chicks during their first week
of life, especially if the food supply is scarce. Key words: willow ptarmigan,
temperature, behavior, nutrition

378

Pekins, P. J. 1988. Winter ecological energetics of blue grouse. Dissertation, Utah
State University, Logan, USA. Abstract: This study investigated relationships
between blue grouse (Dendragapus obscurus) winter habitat and blue grouse adaptive
strategies and overwinter survival. Blue grouse metabolism, physical characteristics of
use-trees and roost sites, microclimatic parameters at roost sites, daily winter energy
costs, and specific energy-saving behaviors were studied. Blue grouse have a relatively
low standard metabolic rate (SMR: 0.835 ml 02 g 1 hr 1) and lower critical temperature
in comparison to other northern tetraonids. Metabolic rate did not significantly increase
from -5 to -20 C. Ambient temperature (Ta) and the square root of wind speed were
significantly correlated with metabolic rate... Wind speeds were reduced by 63 and 85%
at diurnal and nocturnal roosts, respectively. Douglas-firs provided exposure to solar
radiation, protection from wind, and a food source during the day. Subalpine firs
increased protection from the wind and provided near maximum canopy coverage at
night. Nocturnal subalpine roost sites provided a 10% net energy savings in comparison
to diurnal Douglas-fir roost sites. The average field metabolic rate (FMR) of blue grouse
measured with doubly-labeled water was about 1 .4 times SMR, and was not influenced
by weather. The proportion of FMR due to basal metabolism was twice that commonly
assumed for other species, indicating the importance of microhabitat selection and
relative inactivity by blue grouse in minimizing energy costs. Other energy saving
behaviors included sunning, snow roosting, and walking uphill instead of flying. Habitat
selection by blue grouse, particularly microhabitat selection of roost sites, reflects active
choices designed to minimize energy costs imposed by winter weather. This selection
indicates the importance of Douglas-fir and subalpine fir for overwinter survival. Key

H' fc* *k

4*

Ofe -"far- c^CClib cCb

words: blue grouse, temperature, wind, habitat use, biothermal regulation,
behavior, vegetation

379

, J. A. Gessaman, and F. G. Lindzey. 1997. Microclimatic characteristics of

blue grouse Dendragapus obscurus roost-sites: influence on energy expenditure.
Wildlife Biology 3(3/4):243-250. Abstract: Energetic models which incorporate
environmental measures have demonstrated that significant thermoregulatory savings
are accrued from nocturnal winter roost-sites, usually from reduced wind speed and
radiated heat loss. Because blue grouse Dendragapus obscurus occupy high elevation,
snow-bound coniferous stands in the Rocky Mountains during winter, selection of a
favorable microhabitat is likely their primary thermoregulatory behavior. Therefore, we
measured the microclimatic conditions at diurnal and nocturnal roost-sites of blue
grouse to determine whether their choice of roost-sites reflects thermoregulatory
behavior. Temperature, wind speed and solar radiation were measured at 17 diurnal
Douglas-fir Pseudotsuga menziesii and 17 nocturnal subalpine fir Abies lasiocarpa
roost-sites and compared to those of an open control site in Logan, Utah, 1985-1986.
Temperature varied <2°C between the roost-sites and the control site. Wind speed was
significantly lower in 15 of 17 diurnal (mean = 0.71 m/sec) and all nocturnal roost-sites
(mean = 0.24 m/sec) than in the control site (mean = 1 .75 m/sec). Wind speed was
reduced >75% at all but one nocturnal roost-site. Solar radiation at the diurnal roost-
sites (mean = 51 W/m2) was significantly lower than at the control site (201 W/m2);
however, five roost-sites had maximum values >90% of the control maximum. Douglas-
fir roost-sites had significantly greater solar radiation, diurnal, and nocturnal wind speed
than subalpine fir roost-sites. Reduction of convective heat loss was the major
thermoregulatory contribution of both diurnal and nocturnal roost-sites. Diurnal roost-
sites also afforded measurable radiant energy and, presumably, grouse could track the
sun in roost trees to maximize such heat input. Daily energy costs predicted from
metabolic equations incorporating temperature and wind speed were below the
metabolizable energy intake of captive blue grouse. Application of the average
microclimatic conditions from both roost trees to an energetic model revealed that a
blue grouse would realize a 50% greater reduction in convective heat loss, and a 10%
greater net energy savings, by roosting overnight in a subalpine fir rather than a
Douglas-fir. This difference may explain why blue grouse show affinity to subalpine firs
for nocturnal roosting, and points to the energetic importance of specific coniferous
habitats to wintering blue grouse. Key words: blue grouse, solar radiation, wind,
habitat use, thermoregulation, microclimate

380

, , and . 1994. Field metabolic rate of blue grouse during winter.

Canadian Journal of Zoology 72:227-231. Abstract: We measured the field metabolic
rate (FMR) of seven free-ranging and two captive blue grouse (Dendragapus obscurus)
with doubly labeled water. Average carbon dioxide production (1.016±0.088 L CO2 kg "
0734 h-1) of free-ranging grouse was 8% higher but not significantly different (P<0.05)
from captive grouse (0.944±0.058 L CO2 kg"734 h-1). Ambient temperature was not
correlated with FMR (P = 0.268). The mean fat content of free-ranging blue grouse was

172 ^

>> *& ^

39 g (3.4%), which was equal to the energy equivalent of about 3 x daily standard
metabolic rate (SMR). The FMR of free-ranging grouse averaged 657±62 kj/d or 1.6 x
SMR. The FMRs of free-ranging blue grouse averaged about 25% below FMRs
predicted from allometric equations; most were 35-40% below those predicted. We
suggest that there is little energetic constraint on blue grouse during winter because
they are able to maintain a positive energy balance by minimizing energy costs through
effective thermoregulation, microhabitat selection, and reduced activity. Key words:
blue grouse, temperature, metabolism, thermoregulation, habitat use, behavior

381

, , and . 1992. Winter energy requirements of blue grouse.

Canadian Journal of Zoology 70:22-24. Abstract: We measured the effects of
temperature (Ta) on the metabolic rate of 6 blue grouse (Dendragopus obscurus) with
indirect respiration calorimetry. The standard metabolic rate was 0.812 L 02 (kg0734)"1
and was 24% higher than that predicted allometrically. The lower critical temperature
(T|C) of fasted grouse was -5°C; metabolism occurred linearly below -5°C. The heat
increment associated with a Douglas-fir (Pseudotsuga menziesii) diet lowered the T,c by
5°C. From -5 to -20°C, the metabolism of fasted and fed grouse increased by 30 and
12%, respectively. A positive winter energy balance was predicted for blue grouse from
estimates of the field metabolic rate and the consumption and assimilation rates of a
Douglas-fir diet. Keywords: blue grouse, thermoregulation, nutrition

382

Pelgren, E. C. and J. A. Crawford. 1997. Blue grouse Dendragopus obscurus
recruitment and weather relationships in northeastern Oregon, USA. Wildlife
Biology 3(3/4). Not able to obtain for abstract.

383

Pendlebury, C. J., M. G. MacLeod, and D. M. Bryant. 2004. Variation in
temperature increases cost of living in birds. Journal of Experimental Biology
207:2065-2070. Abstract: The effect of temperature variability on laying birds was
studied experimentally, using Japanese quail. Two aspects of temperature variability
were investigated: the effects of regular daily variation in temperature, and of a sudden
change in temperature. Both of these may become more common as a consequence of
climate change. These manipulations were carried out at two levels of food supply.
Energy expenditure increased with higher daily temperature variation, and also after a
sudden change in temperature, taking several days to settle to a constant level.
Manipulating daily temperature variation also resulted in smaller eggs being laid under
more variable temperatures, when food quality was also low. The results demonstrate
that day-to-day variation in temperature, as well as mean temperature, affects energy
expenditure, which can have consequences for egg production. Key words: Japanese
quail, temperature, climate change, biothermal regulation, productivity

% f>^>f=*

173

Ojl ^Jife <£C^i c^i

384

Pepin, D. and M. Fouquet. 1992. Factors affecting the incidence of dawn calling in
red-legged and grey partridges. Behavioural Processes 26:167-176. Abstract:
Seasonal variations in the frequency and the timing of calling in red-legged (Alectoris
rufa) and grey (Perdix perdix) partridges, and their relationships with biological and
environmental factors, were investigated during 89 morning surveys. In winter, when
birds were living in social groups, the vocal activity of both species decreased. Most
calling began before sunrise. Yet many variations in the timing of first calls were
observed when sunrise was prior to 0530 h, i.e. when the daylength was above 13
hours. On the contrary, when daylength was shorter, calling activity always began
approximately 45 minutes before sunrise. Red-legged partridge calls were more
particularly stimulated when sunrise was between 0530 h and 0700 h, with no wind,
when moonrise was between 000 h and 0900 h, and when cloud cover was low. Key
words: grey partridge, red-legged partridge, behavior, solar radiation, wind,
clouds

385

Pepper, G. W. 1972. The ecology of sharp-tailed grouse during spring and
summer in the aspen parklands of Saskatchewan. Wildlife Report No. 1,
Saskatchewan Department of Natural Resources, Regina, Canada. Notes: Page 31
- "Very hot weather resulted in heavier-than-normal woody cover selection and on cool,
heavily overcast days broods were often found all day in very heavy herbaceous
vegetation. After heavy rains or on cool sunny days broods were more likely to be found
in lighter than usual cover." Key words: sharp-tailed grouse, temperature, clouds,
precipitation, solar radiation, behavior, habitat use

386

Perez, R. M., J. F. Gallagher, and M. C. Frisbie. 2002. Fine scale influence of
weather on northern bobwhite abundance, breeding success, and harvest in
south Texas. Proceedings of the National Quail Symposium 5:106-110. Abstract:
Weather plays a substantial role in annual changes in populations of northern bobwhite
(Colinus virginianus) within and among ecological regions. Few studies have tested this
relationship within the confines of specific sites. We examined the fine scale influence of
annual (12-month), seasonal (6-month), and monthly Modified Palmer Drought Severity
Indices (PMDI) and raw precipitation on abundance, breeding success, and harvest of
northern bobwhites on 2 sites in south Texas. We used 18 years (1984-01 ) of roadside
census, juvenile:adult ratios, and harvest records from the Chaparral Wildlife
Management Area (CMWA) in La Salle County and 15 years (1984-99) of juvenile:adult
ratios and harvest records from a private property in Brooks County (BCP) to examine
relationships and trends with weather variables. Bobwhite abundance was correlated
(r>0.53, P<0.023) with 12- and 6-month sums of precipitation and PMDI. Breeding
success was correlated (r>0.53, P<0.023) with 12-month precipitation for both sites and
was correlated (/=0.53, P=0.040) with 6-month precipitation for BCP only. Harvest
variables for CWMA were correlated (a>0.54, P<0.022) with 12- and 6-month PMDI,
while BCP harvest/ha was correlated (r=0.54, P=0.027) with the 12-month precipitation
sum. Monthly correlates with precipitation increased from spring to summer until July

174 j^

^-

VH fc* tk

% m <m? 9

when they became negatively related to rainfall on both sites. Monthly PMDI correlates
became increasingly important from spring through summer including July. Our findings
account for at least part of the annual variation in northern bobwhite abundance in south
Texas and provide information useful in understanding of the influence of weather at
fine spatial scales. Key words: bobwhite quail, precipitation, drought, index,
population dynamics, reproduction

387

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 pheasant hens. Journal of Wildlife
Management 61(3):634-644. Abstract: We studied populations of ring-necked pheasant
hens (Phasianus colchicus) on a diverse landscape in Palo Alto County, Iowa and on an
intensively farmed area in Kossuth County, Iowa to examine the hypothesis that
differences in habitat composition influence winter survival. We captured hen pheasants
by nightlighting or bait trapping on each site and relocated them with telemetry from
December through March, 1989-94. We determined home range area, habitat use, and
movements during the day. We estimated survival using Kaplan-Meier statistics and
related survival to habitat and weather covariates using parametric models. In both
landscapes, home ranges were located in areas with more grass habitat than the
surrounding landscape (P< 0.05). Home range area averaged 76 ha and daily
movements averaged 251 m at Palo Alto, whereas these variables were 96 ha and 270
m at Kossuth. Survival from 27 November to 1 April ranged from 0.23 in 1993 on
Kossuth to 0.96 in 1990 on Palo Alto but the average difference in survival between
sites among years was not different from zero (P = 0.77). Parametric hazards models
indicated that measures of weather and temperature were predictive of mortality, but
that small scale habitat use and daily movements were not. Winters with extensive
snow cover and cold temperatures periodically reduce survival even where the
distribution of small blocks of cover such as idle grass, woody cover, and wetlands is
considered adequate by managers. Key words: ring-necked pheasant, snow,
temperature, mortality, habitat use

388

Perry, R. F. 1946. An appraisal of pheasant abundance in New York State during
1945 and some of the factors responsible for the recent decline. Transactions of
the North American Wildlife Conference 11:141-155. "Discussion of limiting factors —
The most important factors controlling the quality of the environment are shelter, food,
and weather. The whole structure rests on the soil which basically determines the
vegetative cover (food and shelter) and on the climate. Surveys and field observations
have indicated that whatever forces that caused the sudden decline in pheasant
abundance since 1941 have been operative principally during the spring and summer
period. This fact has been particularly obvious in areas where appreciable breeding
populations failed to produce expected numbers of young. Observations during this
period for the past several years have shown that unusually cold, wet weather has often
been associated with such failures.... known seasons of poor pheasant hunting coincide
to a marked degree with years which were abnormally cold and wet during the breeding
and rearing season... it appears quite possible that adverse weather during the spring

\ K<0**

175

and summer may have been to a large extent responsible for the recent "crash" in
pheasant abundance." Key words: ring-necked pheasant, mortality, precipitation,
temperature, productivity

389

Petersen, L. R., R. T. Dunke, and J. M. Gates. 1988. Pheasant survival and the role
of predation. Pages 180-196 in D. L. Hallett, W. R. Edwards, and G. V. Burger,
editors. Pheasants: symptoms of wildlife problems on agricultural lands. North-
central Section of The Wildlife Society, Bloomington, IN, USA. Abstract:
Fluctuations in pheasant abundance are caused primarily by variable rates of mortality,
with reproductive performance playing a subsidiary role. This paper examines seasonal
and annual (fall-to-fall) survival with emphasis on predation. Mean annual survival of
hens of 30-35% appears normal. Wisconsin pheasant studies suggest a long-term
decline in survival and abundance, and an increase in productivity. Low survival has
resulted from habitat loss, and operates through increased predation. Important
predators of adult pheasants are the red fox, red-tailed hawk, and great horned owl.
Predator numbers vary spatially and temporally. High numbers of red foxes have
persisted since the 1940's in nearly all Midwestern states. In addition, the red fox
expanded westward into the prairie states in the 1950's to early 1960's. Red-tailed hawk
and great horned owl numbers were relatively stable in the 1950's and 1960's, but
generally increased 1965-85 in the plains and prairie states, the Great Lakes states,
and those parts of New England that support pheasants. Predation rates are related to
snow depth in winter. Unfavorable weather combined with poor habitat occasionally
results in excessive predation in spring. Predator-prey ratios are likely higher in
Wisconsin than in the more intensively cultivated and highly simplified agricultural
habitats of the plains and prairie states. Predation is, at best, imperfectly density
dependent. It appears that severe winters lead to loss of body condition and increased
mortality due to predation. Attainment of weights favorable for reproduction depends on
the time of winter break-up, and on favorable pre-nesting temperatures. Reproduction
success relates in part to the physiological condition of hens at the start of reproduction
as well as weather and spring cover conditions as they influence predation. Key words:
ring-necked pheasant, snow, habitat condition, predation, nutrition, reproduction

390

Peterson, M. J. and N. J. Silvy. 1994. Spring precipitation and fluctuations in
Attwater's prairie-chicken numbers: hypotheses revisited. Journal of Wildlife
Management 58(2):222-229. Abstract: Two related hypotheses argue that greater than
normal precipitation during May alone or spring (Mar-Jun) leads to decreased Attwater's
prairie-chicken {Tympanuchus cupido attwateri) breeding success, whereas less than
normal precipitation during these periods leads to increased breeding success. These
hypotheses have been accepted by wildlife managers and, seemingly because of
observer expectancy bias, have been used to explain annual variation in Attwater's
prairie-chicken numbers. We demonstrate that neither hypothesis is supported by
available data. Similarly, alternative hypotheses that May or spring flooding, the date in
May when maximum precipitation occurs, or precipitation variability among spring
months drives spring breeding numbers also were not supported. We found, however,

176 ^y

^

K fc* **

that breeding success in spring can drive proportional changes in breeding numbers the
following spring. Keywords: prairie-chicken, precipitation, temperature, population
dynamics

391

Piao, Z.-J. 1997. Reproductive success of hazel grouse at Changbai Mountain.
Acta Zoologica Sinica 43(3):279-284. Abstract: The reproductive success of hazel
grouse (Bonasa bonasia) was studied at Changbai Mountain from 1986 to 1995. Radio-
tracking data showed that the rate of reproductive success was 20.9% (n = 4), and the
survival rate of chicks at the time of brood dissolution was 37.5% (n = 3). Field
observations on broods during May to August showed that the brood chick numbers
decreased with their daily age, with the regression equation of y = 9.916-
0.185x+0. 001 x2(P<0. 0001). Analysis for the effects of weather during April to August on
the reproductive success of hazel grouse indicated that the IRS (index of reproductive
success) was correlated with the weather during early chick period. The reproductive
success was significantly (P<0.005, P<0.025) negatively correlated with the
precipitation and the days of precipitation in June during the young chick period,
suggesting that precipitation had an adverse effect on chick survival. We found that
parasitism by the tick Ixodes persudactus was one of the main factors causing death of
the chicks. Key words: hazel grouse, precipitation, disease, mortality, parasites

392

Porkert, J. 1991. Hoarfrost deposits as a factor contributing to the extinction of
tetraonids in the eastern Sudetes. Ornis Scandinavica 22(3):292-293. Notes:
"Hoarfrost, snow and ice deposits on vegetation affect tetraonids directly in the following
ways: (a) food consumption is hindered, (b) metabolic rates are disturbed, and (c) toxic
substances found in frost and snow are consumed. The birds are also affected indirectly
because (d) the structure of the habitat and the quality of plant foods deteriorate as an
effect of increasing acidity and high contents of toxic substances in the hoarfrost. The
thawing to body temperature of the winter food, together with hoarfrost, snow and ice, in
a bird's crop requires considerable energy. Energy losses may not be compensated for
by eating more food or by using richer energy sources. In the presence of heavy
hoarfrost, larger amounts of frost are consumed during each activity period because
hoarfrost, ice and snow are consumed simultaneously with the food. In periods with
wind and crusty snow, birds have difficulty in digging depressions in the snow, and
thereby lose considerable energy. Keywords: grouse, ice, snow, hoarfrost,
pollution, nutrition, wind, thermoregulation

393

Porter, W. F. and D. J. Gefell. 1995. Influences of weather and land use on wild
turkey populations in New York. Proceedings of the National Wild Turkey
Symposium 7:75-80. Abstract: We assessed the influence of weather and land use of
the population dynamics of wild turkeys (Meleagris gallopavo) during a 12-year period in
southern New York. Wild turkey abundance and rate of population change were
indexed using hunter effort and harvest records from fall hunting across 256 townships.
Winter and spring weather conditions were assessed using data collected at National

-* 4 ^< i . ..^ 177

% >->"<c^**

^■{SN' *d*~*^h> <_ — _)

Oceanic and Atmospheric Administration (NOAA) weather stations. Land use was
assessed using data from New York's Land Use and Natural Resources (LUNR) survey
conducted in 1970. Population abundance was generally below the long-term mean,
with periodic eruptions. On an annual basis, weather and land use were poor predictors
of population dynamics. However, on medium time intervals (3-4 years), weather and
land use accounted for 19 to 95% of the variation in population abundance and rate of
change. We hypothesize that weather factors are poor predictors in annual intervals
because positive and negative influences of different factors are seldom in synchrony.
When favorable conditions are coincident among several weather factors, rapid
population growth occurs. The magnitude of population growth appears related to land
use. Key words: wild turkey, weather, population dynamics

394

Porter, W. F., G. C. Nelson, and K. Mattson. 1983. Effects of winter conditions on
reproduction in a northern wild turkey population. Journal of Wildlife
Management 47(2):281-290. Abstract: Nine aspects of the reproductive performance of
female wild turkeys (Meleaghs gallopavo) were measured in Minnesota during 1975,
1977, and 1978. Sixty-eight of 75 (90%) radio-tagged females attempted to nest,
renesting was common, and eggs in nests of 67% of the females ultimately hatched.
Clutch size averaged 1 1 .7 eggs, hatching success was 82% and 37% of the young
produced survived to late summer. Winter severity, measured in terms of impacts of
site-specific conditions on late-winter body weights of females and mortality rates,
varied within 14 different winter flocks. Females weighing less than 4.3 kg were less
likely to survive to breed, less likely to nest, and had a natality rate of 2.6
females/female in the breeding population. Heavier females had higher survival and
nesting rates and natality was 4.3. Strong correlations were observed between
survivorship within winter flocks and reproductive performance of females that survived
to breed. Severe winter conditions appeared to reduce egg hatching success among
yearling females and recruitment of young among adults. Population modeling
demonstrated that winter mortality and impaired reproduction performance can result in
a significant decline in the population. At least 2 breeding seasons are necessary for
population recovery. Impacts on hunting harvest quotas are discussed. Key words:
turkey, temperature, snow, population dynamics, nutrition

395

Potapov, R. L. and A. V. Andreev. 1982. Time and energy budgets in wintering
Tetraonidae. Congressus Internationalis Omithologici XVIII:409-412. Summary:
Daily energy budgets in 6 species of the family Tetraonidae, wintering in North
Palearctic at Ta-20° - 44° were determined in two or three ways, including
simultaneously measured time budget. The low level of winter's DEB is the result of: (1)
low level of activity - the birds spend only 1-4 hours daily for obtaining the food; (2) the
birds spend much time (20-23 hours daily) in snow burrows - the thermal refuges with
constant Ta of -4° - -5°; (3) the low lower critical To. Using the thermal refuges,
tetraonid birds may regulate average daily Ta near -7° instead of -10° - -60° in open air.
Key words: grouse, snow, temperature, biothermal regulation, behavior

178 ^

4-

f* ^ ffc

(£^,

396

Potts, G. R., S. C. Tapper, and P. J. Hudson. 1984. Population fluctuations in red
grouse Lagopus lagopus analysis of bag records and a simulation model. Journal
of Animal Ecology 53(1):21-36. Abstract: A time-series analysis of numbers of red
grouse (L. lagopus) shot per annum was carried out for 52 moors in northern England
[UK] where data were available for at least 20 consecutive years during the period
1870-1977. Correlograms were constructed and auto-correlation coefficients were
tested for statistical significance. In the most representative sub-sample of moors, 83%
of the series had significant negative coefficients at T + 2 or T + 3 years or both. A
mathematical model was fitted to the fluctuations to describe their quasi-cyclic nature
and the average cycle-length was 4.84 ± 0.086 yr field observations and data from trials
with captive grouse were combined to construct a simulation model of a red grouse
population; its main features were: an inverse logistic curve relating mean numbers of
the parasitic nematode Trichostrongylus tenuis in adult red grouse to their breeding
success; T. tenuis accumulation by young red grouse to steady state levels (the
numbers of worms accumulated per bird varied according to a combined effect of the
density of grouse prior to the breeding season and the worm burden of these grouse); a
logistic curve relating to the proportion of grouse shot with the density of grouse
available; and an annual survival rate of the non-shot population inversely proportional
to the density of old grouse. The simulation model successfully captured mean levels
and other population parameters, but it fit well to observed fluctuation patterns only
when stochastic elements were introduced to represent known effects of weather. The
model showed that red grouse cycles could be caused by effect of T. tenuis working
together with stochastic elements and a time delay arising form the uptake of worms.
Key words: red grouse, disease, weather, population dynamics, parasites, model

397

Prellwitz, D. M. 2002. Greater sage-grouse at Bowdoin National Wildlife Refuge,
Montana. Intermountain Journal of Science 8(2):1 15-116. Notes: Page 115-
"Refuge employees, while conducting annual waterfowl nesting studies, nest drag from
160-510 ha of upland grass habitat, and occasionally flush lone sage-grouse in front of
the drag. One sage-grouse nest was found on 6 June 1988. The nest of seven eggs
was found in dense grass cover beneath one of only a few plains silver sagebrush
plants in the area. The nest hatched at least four eggs by 27 June during one of the
hottest months on record when daytime highs were >32°C on 22 days with 9 of those
days >38°C. Although this age grouse nest survived despite the above-normal daytime
temperatures, several nests of other species dehydrated and were lost. Total
precipitation in 1988 was 24.4 cm, far below the long-term average of 31 .9 cm. Key
words: sage-grouse, temperature, precipitation, habitat use, vegetation,
productivity

398

Priest, S. R. 1995. Relationships between wild turkey hens, environmental factors,
and predation during the wild turkey reproductive period on Tallahala Wildlife
Management Area (Mississippi, Meleagris gallopavo, Procyon lotor). M. S.Thesis,
Mississippi State University, Starkville, USA. Abstract: Racoon (Procyon lotor) home

% ^ffc&r^

179

*±f'

(-otv^^jJ3 C J

range size, habitat use, survival rates, and effects of meteorological events on wild
turkey (Meleagris gallopavo) hen nest losses from predation were studied on Tallahala
Wildlife Management Area (TWMA) in central Mississippi, 1991-1992. Data from 33
transmittered raccoons and 103 turkey hens were used in analyses. Raccoon seasonal
home range sizes were significantly larger during the wild turkey nesting period and
generally were smallest during the post-nesting period. Composite home range sizes
were consistent with results of other recent studies of raccoons conducted in the
Southeast. Bottomland hardwood forests were preferred by raccoons during all periods.
However, during the turkey nesting period raccoon habitat use changed to include more
mature pine (Pinus spp.) stands and pine regeneration areas and less hardwood.
Disease seemed to be a major mortality factor of raccoons during 1 991 . Illegal trapping
was the major cause of death during 1992. Survival rates were 0.53 and 0.69 for 1991
and 1992, respectively. Although total amount of rainfall during the wild turkey nesting
period was not correlated with nesting success, pattern of rainfall was important.
Predation on turkey nests often occurred after rains preceded by at least 2 days of dry
weather. Management implications and recommendations for future research are
discussed. Key words: wild turkey, precipitation, predation, mortality

399

Pullianen, E. and P. S. Tunkkari. 1991. Responses by the capercaillie (Tetrao
urogallus) and the willow grouse (Laogpus lagopus) to the green matter available
in early spring. Holarctic Ecology 14(2):156-160. Abstract: Food matter eaten in the
first snowless spots early in the spring by capercaillie Tetrao urogallus and willow
grouse Lagopus lagopus was studied in Finnish Forest Lapland. When the snow
disappears, both species change to feeding mainly on the plants available in the
snowless spots, although male capercaillie does not exploit this nutritious diet to the
same extent as female capercaillie and willow grouse. Selection in favor of nitrogen and
phosphorus-rich food items, i.e. Betula pubescents, male catkins (Eriophorum
vaginatum), flower buds, and Equisetum spp., stems and tips, seems to be
characteristic especially of the female capercaillie, whose winter diet is poor in these
elements. Female capercaillie also feeds on more plant species or different parts of
plants at one time than willow grouse, while no differences were recorded between
male and female willow grouse in the composition of their spring food. The crowberry
Empetrum hermaphroditum, almost regularly produces a bountiful berry crop in northern
Finalnd, and since it overwinters well, it represents an easy source on energy and water
available almost every spring. The quality and/or the quantity of the green food matter
available early in the spring may fluctuate considerably and be of importance for short-
term fluctuations in the populations of these tetranoid species. Key words:
capercaillie, willow grouse, snow, temperature, vegetation, phenology, nutrition

400

. 1978. Behavior of a willow grouse Lagopus lagopus lagopus at the nest.

Ornis Fennica 55(4):141-148. Abstract: The behavior of a nesting willow grouse pair
was studied mainly with the aid of a field television system with remote control in
eastern Finnish Lapland (67°N44'N, 29°37'E) in continuous daylight. During the laying
period the hen covered the eggs with plant matter. Incubation started before the laying

180 j^

4

^ fc* ^

cCi

of the penultimate egg. Only the hen incubated. The hen could be off the nest at any
time of the day and the nest was left unattended 5.7% of the time. Snowfalls occurred
during the incubation period. In continuous daylight the willow grouse is active for 24 h.
In cold weather the frequency of settling movements remained high, while preening,
shifting, resettling and nest-building activities decreased. Exceptionally high frequencies
of preening, feeding (catching) and nest-building were recorded during the hatching of
the chicks, which points to displacement activity. The period from the beginning of
intensive incubation to departure of the brood was about 22 days. Key words: willow
grouse, temperature, behavior

401

Queal, L. M., J. A. Norman, K. Sexson, and G. J. Horak. 1971. Population
dynamics of prairie chicken in grassland and cropland areas. Pittman-Robertson
Project W-23-R, Job No. D-3, Kansas Forestry, Fish and Game
Commission.Topeka, USA. Notes: Page 64 — Effects of Rainfall on Brood Survival.
There appears to be a correlation between variations in the annual precipitation
patterns and variation in prairie chicken population density. The precipitation pattern
during the latter part of May and into June, when young are susceptible to inclement
weather, seem particularly important. Page 67 — Effects of Weather on Hunting
Success. Temperatures below 30°F occurred only twice during 1964 prior to the hunting
season when hunting success was at the lowest.... During both days of the opening
weekend in the low year of 1964 and in the high of 1966, fog was prevalent until 8:30
A.M. or 9:00 A.M. In 1966, when the temperatures dipped to the lower teens prior to the
season, certain populations of insects and living vegetation were frozen, lowering the
availability of food in the pastures; prairie chickens then began using feed fields where
food was more plentiful. Key words: prairie chicken, precipitation, temperature, fog,
mortality, hunting, habitat use, movement

402

Quinn, N. W. S. and D. M. Keppie. 1981. Factors influencing growth of juvenile
spruce grouse. Journal of Canadian Zoology 59(9):1635-1855. Abstract: The
influences of date of hatch, and age and prelaying body weight of brood female on the
growth rate of juvenile spruce grouse (Canachites canadensis) were studied in central
New Brunswick in 1977 and 1978. Because of differential timing of hatch of broods of
adult and yearling females, it was not clear whether a difference in juvenile growth rates
during 5-14 days of age in 1977 was related to date of hatch, age of brood female, or
both. Differences in juvenile growth rates within and between years apparently were not
influenced by body weight of brood females prior to egg laying. Results suggest that
posthatch factors are more important in determining growth rate than a prehatch or
"maternal" influence. Notes: Page 1793 — The marked difference in growth of juveniles
during the 2 years of this study probably was due to much warmer, drier weather
throughout the summer of 1978 than 1977, which allowed chicks to divert more
metabolizable food energy to growth. Faster early growth of late-hatched than early-
hatched juveniles also has been reported for blue grouse (Redfield 1978). Redfield
speculated that the relation between hatch date and growth was due to changing
abundance of insects used as food. Faster early growth of late-hatched juveniles could

\ fo^yfk

181

t£^> c^

be due to any factor such as food supply or temperature which influences the
partitioning of food energy. Key words: spruce grouse, temperature, precipitation,
morphology, insects, nutrition

403

Rabe, D. L. 1981. Habitat and energetic relationships of American woodcock in
Michigan. Dissertation, Michigan State University, East Lansing, USA. Abstract:
Studies were undertaken to examine (1) the role of interspersion and food availability on
habitat utilization by breeding woodcock (Philohela minor), (2) the proximal cues of
habitat used to locate feeding sites and the foraging strategies used to capture
earthworms (Lumbricidae), and (3) the impact of weather on food availability and
bioenergetics of breeding woodcock. During the springs of 1978 and 1979, 23 aspen
(Populous tremuloides) community habitat complexes were censused for singing males
and intensively searched with a pointing dog to locate nests, broods, and solitary birds.
At the same time, data were collected on habitat structure and earthworm abundance.
Thirty-two solitary birds and 31 broods were located during 78 h of searching in the two
years. Singing male woodcock used 17 and 20 of the habitat complexes in 1978 and
1979, respectively. Three of the complexes were never used by woodcock during the
study. Between-year comparisons of each habitat complex revealed that use by singing
males and solitary birds was much more consistent than brood use. Numbers of broods
using a complex were correlated (P < 0.10) with numbers of singing males in 1979.
Earthworm abundance was correlated (P < 0.10) with brood use in both years;
correlations to males were weaker. Structural measures of the habitat complexes (i.e.,
size, shape, and interspersion) were not consistently correlated with any woodcock use
between the years. In a series of laboratory experiments, the foraging behavior of live-
trapped adult woodcock was examined in order to test specific hypotheses about the
proximal cues of habitat used to select feeding sites and foraging strategies used to
capture earthworms. Foraging trials were conducted in a circular arena where the birds
were allowed to probe among eight soil trays containing various soil types, soil
moistures, and earthworm size classes and densities. Color, which tends to be
correlated with soil types and moisture regimes preferred by earthworms, was found to
be the primary proximal cue used by woodcock for selecting feeding sites in these
experiments. Birds concentrated searching effort in areas of relatively high prey density,
but exhibited no size selectivity. Following the capture of one earthworm, birds tended
to concentrate additional searching in the immediate area. This non-random search
pattern seems to account for the greater efficiency of capture when prey are
aggregated Key words: American woodcock, moisture, soil, nutrition,
bioenergetics

404

, H. H. Prince, and E. D. Goodman. 1983. The effect of weather on

bioenergetics of breeding American woodcock. Journal of Wildlife Management
47(3):762-771. Abstract: Simulation modeling was used to investigate the impact of
weather on the bioenergetics of breeding and postbreeding American woodcock
(Scolopax minor). Using air temperature and precipitation as inputs, the model
calculates the daily energy requirements of an adult female woodcock and chicks along

182 j,

4

K kr« t*

with the availability of their primary food, earthworms (Lumbricidae). Energetics were
modeled from previous studies on woodcock and related species. Earthworm
availability was modeled from field data collected in northern Michigan. Results suggest
that the greatest potential for weather-related stress on woodcock occurs during the
brood-rearing period, with nesting being the 2nd most critical time. When simulated
earthworm availability during the brood period was compared with reproductive success
data, it indicated that the impact of spring weather on earthworm availability is a
significant factor affecting chick survival. Key words: American woodcock,
temperature, precipitation, bioenergetics, nutrition, reproduction, model

405

Raitt, R. J. and R. E. Genelly. 1964. Dynamics of a population of California quail.
Journal of Wildlife Management 28(1):127-141. Abstract: Dynamics of a population of
California quail (Lophortyx californicus) were studied for 8 years, 1950-57, by capture-
recapture methods. Fall population size varied between 25 and 140. Productivity varied
between 56.5 immatures:100 adults and 222 immatures:100 adults as indicated by fall
age ratios. Reproductive success appears to have been promoted by early onset of the
breeding season and a minimum of fog and rain during the hatching season. The
composite overall mortality rate was 71 percent, and life tables indicate only slight
variation with age. The success of the breeding effort was probably the major factor in
determining fall population size. Seasonal changes in the type and degree of mobility —
correlated with changes in social behavior — are described. Population reductions were
carried out in two winters, and in both years the population had returned to normal size
by autumn. Immigration of birds from surrounding areas, rather than compensatory
changes in mortality and productivity, appears to have been responsible for the
recovery. Key words: California quail, precipitation, fog, reproduction,
productivity

406

Rand, A. L. 1947. Clutch size in the spruce grouse and theoretical considerations
of some factors affecting clutch size. Canadian Field-Naturalist 61:127-130.

Abstract: The available data indicate that clutch size in Canachites canadensis is 4-7,
which is fewer than indicated in standard texts. There is no geographic variation in
clutch size in Canada. Clutch size may be affected by food abundance, cyclic
abundance, spring weather changes, age of bird, interrupted nesting and by whether
the bird is a determinate or indeterminate layer. With the red grouse of Europe the
number of eggs laid is said to be 4 to 9 in cold wet springs; 6 to 12 in very favorable
seasons, and 7 to 8 in average years. Key words: spruce grouse, red grouse,
weather, reproduction

407

Ratti, J. T. and J. H. Giudice. 2001. Assessment of chukar and gray partridge
populations and habitat in Hells Canyon. Technical Report Appendix E.3.2-7,
Hells Canyon Complex FERC No. 1971. Report to Environmental Affairs, Idaho
Power Company, Boise, USA. Summary: This report contains a detailed literature
review on the chukar and gray partridge. Subsections 4.5.4 Wintering provides a

H fc* >f*

X

?#

£^> <£^>

discussion of the impact of weather on these two species. Key words: chukar
partridge, gray partridge, precipitation, snow, wind, habitat use, vegetation

408

Remington, T. E. and C. E. Braun. 1985. Sage-grouse food selection in winter,
North Park Colorado. Journal of Wildlife Management 49(4):1 055-1 061. Notes:
"Sage-grouse use seasonal habitats that differ in plant composition and structure, and
that may be widely separated. Winter habitat is probably the most limited seasonal
habitat and thus may be the most critical. This study has shown that winter use areas
have a specific species/subspecies composition and unique chemical characteristics in
addition to topographic and structural features. Identification and protection of these
areas is essential if sage-grouse populations are to be maintained. Locating stands of
palatable sagebrush (e.g., Mountain big sagebrush) is not enough, because snow cover
can make large expanses of sagebrush unavailable to sage-grouse in winter. Within
wintering areas, particular protection should be given to stands of palatable
species/subspecies of sagebrush. Identification of big sagebrush to subspecies is
essential because use differs greatly. Use and importance of Mountain big sagebrush
may increase in winters of heavy snowfall because of its generally taller growth form.
Use and preference of species/subspecies of sagebrush should be investigated locally,
particularly where composition differs from that encountered in this study. Key words;
sage-grouse, snow, diet, nutrition, habitat management, vegetation

409

Rice, S. M., F. S. Guthery, G. S. Spears, S. J. DeMaso, and B. H. Koerth. 1993. A
precipitation-habitat model for northern bobwhites on semiarid rangeland.
Journal of Wildlife Management 57(1):92-102. Abstract: Models that predict wildlife
responses to natural and human perturbations are useful in research, extension, and
management. Thus, we developed a priori and tested a precipitation-habitat model for
predicting autumn density of northern bobwhites (Colinus virginianus) on semiarid
rangeland. Variables in the model included the proportion of an area usable based on
dispersion of woody cover, Thornwaite's index of precipitation effectiveness, proportion
of sand-sized particles in the soil, proportional coverage of woody cover and forbs, and
proportional exposure of bare ground. Observed densities were a linear function of
modeled densities based on pooled data from the Rio Grande Plains (n = 16) and Gulf
Coast Prairies, Texas (n = 6)(r2 = 0.34, 20 df, a = 0.0021). The model differed from
species-habitat models by using an environmental correlate of bobwhite abundance
(precipitation effectiveness), which allowed incorporation of quail population history into
model output. The model provides an analytical framework for identifying habitat
deficiencies on semiarid rangeland and for predicting bobwhite population response to
remedial management. Key words: bobwhite quail, precipitation, model, population
dynamics, index

410

Rich, T. 1985. Sage-grouse population fluctuations: evidence for a 10-year cycle.
Technical Bulletin 85-1, Bureau of Land Management, Idaho State Office, Boise,
USA. Abstract: Thirty-two years of counts of sage-grouse (Centrocercus urophasianus)

184 ^

4>

H t* >f*

cCb

males on leks in southern Idaho revealed major population peaks about every 10 years.
Lek counts from northern Utah and western Nevada and harvest data from Idaho and
Utah all show strong synchrony in this cycle. Individual lek counts within each region
tended to fluctuate together despite being located in different habitats or at different
elevations. Eight of nine monthly weather variables having significant correlations with
lek counts in Idaho were precipitation, rather than temperature, variables. However,
weather variables were not useful in predicting mean lek counts through linear-
regression analysis. In Idaho 79% of the variation in mean lek count could be explained
by knowing the mean lek count of the previous spring and the juvenile/adult ratio in the
preceding autumn's harvest. About 65% of the variance in the juvenile/adult ratio could,
in turn, be explained by knowing the amount of precipitation in July and August. There
is no evidence that sage-grouse populations decline as a result of being alternate prey
in a predator-prey system involving coyotes (Canis latrans) and cycling black-tailed
jackrabbit {Lepus californicus) populations. Key words: sage-grouse, precipitation,
temperature, population dynamics

411

Riley, T. Z., W. R. Clark, E. Ewing, and P. A. Vohs. 1998. Survival of ring-necked
pheasant chicks during brood rearing. Journal of Wildlife Management 62(1 ):36-
44. Abstract: Survival of chicks is an important and poorly understood component of
ring-necked pheasant (Phasianus colchicus) population dynamics. We implanted
transmitters in day-old chicks (n = 332) with brooding hens (n = 117) during 1990-94 in
northern Iowa and calculated survival to 28 days of age. We contrasted survival among
years and between an area in Palo Alto County with >25% grassland habitat and an
area in Kossuth County with <10% grassland. Average survival (S2s) of pheasant chicks
in Palo Alto County was 0.46± 0.1 1 and did not differ (P = 0.355) from Kossuth County
(S28 = 0.37±0.13, P = 0.039). Weasel (Mustela erminea), red fox (Vulpes Vulpes), and
mink (Mustela vison) together accounted for >85% of the mortality. Twenty-three chicks
died of exposure on days when at least a trace of rainfall fell, and 11 of 23 (48%) died
on days when rainfall was >0.6 cm (mean = 0.86 cm, range = 0.68-3.38 cm). Age of the
hen did not influence chick survival. Chick mortality rate was increased by 2.3% for
each day chicks hatched after the median date of hatch (15 Jun) and was decreased by
10% for each gram of mass above the average chick mass at hatch (18.5±0.13 g).
Habitat management to improve chick survival of pheasants on agricultural landscapes
should emphasize perennial grass and legume cover dispersed among crop fields.
Grassland cover should remain undisturbed, particularly early in the nesting season (15
April-1 Jun), to improve the chances of successful first-nest attempts. Key words: ring-
necked pheasant, precipitation, mortality, temperature, habitat use, chick survival

412

Ritcey, R. 1995. Status of the sharp-tailed grouse (columbianus subspecies) in
British Columbia. Wildlife Working Report No. WR-70, Ministry of Environment,
Lands and Parks, Wildlife Branch, Victoria, British Columbia, Canada. Notes: Page
8 — Weather, disease and starvation. There is no evidence that severe weather is a
significant direct mortality factor affecting "Columbian" sharp-tailed grouse. However, its
indirect effects may be more profound. Years of low precipitation may reduce

\ f>"^^

185

production of vegetation needed for security or nesting cover, leading to increased
predation of adults and chicks. Key words: sharp-tailed grouse, precipitation,
vegetation, predation, mortality

413

Ritcey, R. W. and R. Y. Edwards. 1963. Grouse abundance and June temperatures
in Wells Gray Park, British Columbia. Journal of Wildlife Management 27(4):604-
606. Abstract: A study of weather records and the annual grouse kill in Wells Gray Park,
British Columbia, showed a relationship between June weather and the fall kill. A
positive correlation coefficient of 0.913 was found to exist between the fall grouse kill
and the mean maximum June temperature. The grouse involved were chiefly ruffed
grouse (Bonasa umbellus). It seems apparent that high temperatures in late spring and
early summer are an important factor in the reproductive success of grouse. Key
words: ruffed grouse, temperature, reproduction

414

Rivera-Milan, F. F. 1997. Seasonal and annual changes in the population density
of Zenaida doves in the xerophytic forest of Guanica, Puerto Rico. Journal of
Field Ornithology 68(2):259-272. Abstract: The Zenaida dove (Zenaida aurita) is
common and widely distributed in the three major life zones (dry, moist, wet) of Puerto
Rico. The xerophytic forest of Guanica may be a key habitat for the reproduction of
Zenaida doves in the dry zone. Point-counts (six 3-min stations) were conducted from
1986-1996 and nest-counts (46 0.1 -ha strip-transects) from 1987-1992 to study the
changes in the density (D) of Zenaida doves and their active nests in the xerophytic
forest. From 1987-1992, the mean D of Zenaida doves was 4.83/ha and the mean D of
active nests was 1.66/ha. The mean D of active nests was higher in areas dominated by
semi-deciduous vegetation than in areas dominated by deciduous and thorn-scrub
vegetation. Point-counts and strip-transects suggested a population decline from 1987-
1992, with 1989 having the lowest D of Zenaida doves and active nests. The rainfall of
the first 6 mo of the year explained 72-80% of the variation in 6 from point-counts and
strip-transects from 1987-1992. Point-counts suggested a significant population decline
from 1986-1996, with 1989 and 1995 having the lowest counts of Zenaida doves. The
rainfall of the first 6 mo of the year explained a significant proportion (51%) of the
variation of point-counts from 1986-1996. To detect 10-25% differences (a=0.05,
power=0.80) in the annual mean D of Zenaida doves and their active nests, sampling
32 3-min stations and 79 0.1-ha strip-transects monthly from April-June is
recommended. These sample sizes are large enough to detect positive or negative
trends of 10% through linear regression analysis. The conservation and management of
the xerophytic forest may be of critical importance for maintaining current population
levels of Zenaida doves in the dry zone of Puerto Rico. Key words: Zenaida dove,
precipitation, population dynamics

415

and F. C. Schaffner. 2002. Demography of Zenaida doves on Cayo del

Agua, Culebra, Puerto Rico, Condor 104:587-597. Abstract: The demography of
Zenaida doves (Zenaida aurita) was studied on Cayo del Agua, Culebra, Puerto Rico.

186 j^

^

>->'• ^ ^

We collected capture-recapture and reproductive success data and monitored annual
changes in the density of ground nests in 1987-1993 and 2000. Models with time-
specific apparent survival and constant capture rates and constant apparent survival
and time-specific capture rates were equally parsimonious, with the former having 1.5
times more support from the data. Rainfall of the first six months of the year was
positively related to nest density, and crab density was negatively related to nesting
success and the number of doves fledged per nest. Crabs are the main predators of the
dove nests on Cayo del Agua. Models, parameterized with field data, were used to
simulate full and reduced stochastic variation in environmental and demographic
conditions, and predict annual changes in population size. High recruitment (birds or
births + immigrants) offset high losses (deaths + emigrants) in all instances. Our field
data suggest that Zenaida doves suffered an ecological crunch between 1989 and
1990, when weather (a hurricane followed by a drought), food availability, and nest
predation interacted, lowering the number of locally fledged doves that survived the
hatching year (1989-1990) and reproductive success (1990). Under severe conditions,
population size and recovery mainly depend on immigration. Apparent survival returned
to pre-hurricane levels between 1990 and 1991, and reproductive success was about
average in 1991. Key words: dove, wind, precipitation, drought, nutrition,
predation, mortality, productivity, model

416

Robel, R. J. 1969. Movements and flock stratification within a population of
blackcocks in Scotland. Journal of Animal Ecology 38(3):755-763. Notes: (Page
759) — Blackcocks roosted far more frequently in the forest than on the moor. Roosting
in trees was most commonly observed during rainy nights in the winter months, while
roosting amongst the heather on the forest floor was most frequent when deep snow
covered the moor. Only when the weather was dry and mild did blackcocks normally
roost on the open moor. Key words: blackcock, precipitation, temperature, snow,
habitat use, behavior

417

Roberts, S. D. 1997. Relationships between weather and the annual dynamics of
wild turkey populations in New York (Meleagris gallopavo, reproductive).
Dissertation, State University of New York, Syracuse, USA. Abstract: Reproductive
success can be highly variable within wild turkey (Meleagris gallopavo silvestris)
populations, and success in some regions appears to be primarily associated with
annual fluctuations in nest or poult survival. Identification of factors causing variation in
these parameters could result in new indices to productivity. Radio-telemetry data
obtained from wild turkeys in southcentral New York during 1990-93 were analyzed
using logistic regression to: (1) examine relationships between 14 weather variables
and daily nest survival, and (2) investigate the influence of heating-degree-days and
deviation from normal precipitation on survival of poults to 25-days posthatch. In
addition, linear regression analyses were used to examine relationships between May
precipitation and fall wild turkey harvest in southwestern New York during 1972-95.
Daily nest survival was negatively associated with cumulative departure from normal
seasonal rainfall (P = 0.003) and daily precipitation (P = 0.07), and positively associated

i ^H^^k

187

<£*^*-f--vj3 C _>

with heating-degree-days (P - 0.01 ). Predicted and observed rates of daily nest survival
were not highly correlated (r= 0.31), but rates based on 10- or 20-day means were
more highly correlated (r= 0.76 and r= 0.93, respectively). The proportion of poults
surviving the 25 days posthatch was negatively associated with mean daily heating-
degree-days during the first week post hatch (P = 0.0001 ) and negatively associated
with mean deviation from normal daily precipitation during the second week posthatch
(P = 0.08). Survival rates within broods were not highly associated with weather-based
predictions of brood survival (r= 0.46, n = 26), but observed and predicted estimates of
annual poult survival were similar. Annual changes in reported harvest were negatively
associated (P<0.0001) with the loge of annual changes in May precipitation. Weather-
based indices can be a viable alternative to brood surveys: a simple linear regression
model correctly predicted observed annual trends (increase vs. decrease) in fall harvest
for 19 of 24 years. Predictability was similar for a more complex model that accounted
for autocorrelation of error terms. Data indicate that nest success may be driving annual
trends in New York's wild turkey population. Key words: eastern wild turkey,
temperature, precipitation, reproduction, index, model

418

, J. M. Coffey, and W. F. Porter. 1995. Survival and reproduction of female

wild turkeys in New York. Journal of Wildlife Management 59(3):437^447. Abstract:
Annual fluctuations of northern wild turkey (Meleagris gallopavo silvestris) populations
often are attributed to high winter mortality. However, studies conducted in agricultural
environments have demonstrated that seasonal survival can be highest during winter,
suggesting other factors are more important to annual population change. We examined
survival and reproduction of female eastern wild turkeys in south-central New York
during 1990-93 and conducted a sensitivity analysis to determine the relative
importance of demographic parameters to annual population change. Seasonal survival
rates (n = 238) were 0.800 for spring, 0.855 for summer, 0.834 for fall, and 0.873 for
winter. Fall survival rates varied by years (P < 0.01) and were higher during years of
above average hen success. Annual survival rates averaged 0.498, and crude annual
mortality rates (M) averaged 0.321 for predation and 0.117 for poaching, hunting, and
wounding combined. Subadult females had lower nesting rates (P = 0.002, n = 201 ),
lower renesting rates (P = 0.001 , n = 1 15), and lower hen success rates (P = 0.02, n =
196) than adult females. Nest success averaged 37.9% (n = 323) and was highest (P =
0.005) during years with average to below average May rainfall. We observed annual
variation in nest success (P = 0.001 ), hen success (P = 0.003), and hatching rates (P =
0.04). Poult survival averaged 40% (n = 605) and did not vary among years (P = 0.74).
Nest success was the primary factor contributing to annual population change. We
suggest that annual fluctuations of northern populations in mixed agricultural and
forested environments rarely result from variability of annual survival and may result
from variability of annual nest success and poult survival. Northern populations
subjected to infrequent severe winters in mixed agricultural and forested environments
likely would benefit more from enhancement of nesting and brood-rearing habitat. Key
words: eastern turkey, precipitation, population dynamics, reproduction,
vegetation

188 j,

^

K (& >f*

cCi

419

and W. F. Porter. 1 998a. Relation between weather and survival of wild

turkey nests. Journal of Wildlife Management 62(4):1492-1498. Abstract:
Recruitment can be highly variable within eastern wild turkey {Meleagris gallopavo
silvestris) populations and appears primarily associated with annual fluctuations in nest
survival. Identification of the causes of variation in nest survival could lead to the
development of new indices to productivity. We used logistic regression to examine
relations between 14 weather variables and daily survival of wild turkey nests ( n =
3,332 nest-days from 206 nests) in south central New York from 1 May to 9 June 1990-
93. Daily nest survival was associated negatively with cumulative departure from normal
seasonal rainfall (P = 0.003) and daily precipitation (P = 0.07) and was associated
positively with heating degree-days (P = 0.01 ). Although estimates of daily nest survival
predicted by the logistic regression model were not correlated highly with observed
estimates (r= 0.31), a strong positive correlation was observed between predicted and
observed probabilities of 20-day nest survival (r = 0.93). Key words: turkey,
precipitation, temperature, nest survival

420

and . 1998b. Influence of temperature and precipitation on survival of

wild turkey poults. Journal of Wildlife Management 62(4):1499-1505. Abstract: New
methods for determining annual trends in wild turkey (Meleagris gallopavo) populations
are desirable, especially when heavily hunted populations are managed with limited
resources. Cost-effective indices to population trends could result from an increased
understanding of relations between environmental factors and reproductive success.
We examined the influence of heating degree-days (HDD) and deviation from normal
precipitation on the 25-day survival of eastern wild turkey poults (M.g. silvestris) in
southcentral New York during 1991-93. The proportion of poults surviving to 25 days
posthatch was associated negatively with mean HDD/day during the first week
posthatch (P < 0.001) and associated negatively with mean deviation from normal daily
precipitation during the second week posthatch (P = 0.08). Observed survival rates of
poults within broods were not highly associated (r= 0.46, n - 26) with weather-based
predictions of survival within broods, but observed and predicted estimates of annual
poult survival were similar. This similarity suggested weather might be a good predictor
of annual rates of poult survival. Key words: turkey, index, temperature,
precipitation, productivity, chick survival

421

Robinson, J. D. 2007. Ecology of two geographically distinct greater sage-grouse
populations inhabiting Utah's west desert. M.S. Thesis, Utah State University,
Logan, USA. Abstract: "...nesting success was higher in 2005 than 2006. Brood
success was similar for the two years. The ratios of chicks per successful brood were
higher in 2005 than 2006, for both sites, Ants (Formicidae) were the most abundant
arthropod available to sage-grouse within the Sheeprock Watershed. I attribute these
differences to precipitation. The spring of 2005 had twice the 30-year average spring
precipitation, coming after a 5-year drought. However, there were no differences in
vegetation at brood and random sites between years for either population. Chick

j^ 189

^

f V ^ ^

1§f y$> ^^ ^

recruitment in both populations was lower than reported in the literature. Sage-grouse
survival rates for the Sheeprock and Deep Creek Watershed populations are lower and
higher, respectively, than most published reports. Sage-grouse conservation strategies
in both areas should emphasize enhancing existing brood-rearing habitat and protecting
critical seasonal winter habitat." Key words: sage-grouse, precipitation,
reproduction

422

Roersma, S. J. 2001. Nesting and brood rearing ecology of plains sharp-tailed
grouse (Tympanuchus phasianellus jamesi) in a mixed-grass/fescue ecoregion of
southern Alberta. M.S. Thesis, University of Manitoba, Winnipeg, Canada. Notes:
(Page 93): "Weather conditions can be a major factor in brood mortality. Several
researchers have identified extended periods of wet cold weather as a cause of brood
loss. Hens in this study experienced a significant loss of broods due to similar
conditions in mid June of 1998. During this time 3 broods were lost to exposure, which
constituted 37.5 percent of the total broods. In addition to 1 predation and 2
unconfirmed loss, brood survival in 1998 was 0.25. These results parallel those
documented by Brousquet and Rotella who in one year reported a 0.30 brood survival
rate due to inclement weather. Hence, the presence of dense cover to escape
inclement weather conditions is essential to ensure sufficient year-to-year recruitment of
juvenile individuals into the population." (Page 94): "...I feel that the habitats available to
brood rearing hens on the Milk River Ridge mitigated any further brood losses due to
the effects of weather." Key words: sharp-tailed grouse, temperature, precipitation,
reproduction, vegetation, population dynamics

423

Rogers, G. E. 1968. The blue grouse in Colorado. Technical Publication No. 21,
Colorado Game, Fish and Parks Department, Denver. Notes: Pages 44-45: Weather
and vegetative relationships. Personnel making upland game-bird brood counts are
usually instructed to run census trends only when weather conditions are clear, calm,
and dry. However, while establishing sage-grouse trends it was noted that counts were
often high following a storm since birds moved to open areas to escape wet vegetation.
In 1961, while less than 10% of trend-route travel was under wet or rainy conditions,
one of the seven broods was observed while rain was falling, five of seven were
observed while vegetation was wet from rain, four of seven when cloud cover exceeded
50%; and five of seven while the sun was not shining. Page 46: With equal coverage in
1963 under wet and dry conditions, one of three broods was observed in damp
vegetation, light showers having occurred early in the morning; two of three were
observed under clear, dry conditions; and none were observed during wet and cloudy
conditions. The average wind velocity was recorded at the start and end of the census
route and at each observation. In only one instance during 3 years was a brood
observed when wind velocity exceeded 14 miles-per-hour. Page 59: The peak of hatch
probably occurs during the latter part of June. Optimum time of day for brood
observations varied, partly due to moisture conditions; early morning, late evening, and
from 2:00 to 5:00 PM were most productive. No broods were observed where shrubs

190 ^

^

» t& >£*

?©>> <mfr O

were absent. Key words: blue grouse, precipitation, clouds, wind, population
survey, vegetation, movement

424

. 1964. Sage-grouse investigations in Colorado. Technical Publication

Number 16, Game Research Division, Colorado Game, Fish and Parks
Department, Denver. Notes: Page 21 — Seasonal Distribution. "In some instances,
sage-grouse begin strutting on their winter range and gradually move to strutting areas
at higher elevations as snow and mud disappear, thus the birds may use 2 or 3 grounds
during a single season. The use of alternate strutting grounds was observed in the
Gunnison area." Page 27 — "Sagebrush growth from 7 to 15 inches high were preferred
for feeding, nesting, and roosting, with the taller plants being used for nesting, shade,
and escape cover." Page 48 — Factors Affecting Reproductive Success: Several authors
found weather to be a factor in nest destruction or desertion; two researchers found
weather had little, if any, effect on nesting sage-grouse. Key words: sage-grouse,
snow, soil, solar radiation, vegetation, behavior, habitat use

425

Rolley, R. E., K. F. Kubisiak, R. N. Paisley, and R. G. Wright. 1998. Wild turkey
population dynamics in southwestern Wisconsin. Journal of Wildlife Management
62(3):91 7-924. Abstract: We studied the population dynamics of eastern wild turkeys
(Meleagris gallopavo silvestris) in southwestern Wisconsin during 1987-94 to better
understand the effects of variation in reproduction and fall harvest on growth of a
northern population. We combined yearly estimates of reproduction, survival, and
population size in a stochastic population model to estimate the finite rate of increase of
the population. Our model indicated recruitment during 1989-92 was inadequate to
offset observed mortality: the finite rate of increase was 0.831 ± 0.097 (mean ± SE).
The simulated trend was corroborated by declines in spring harvest success and
observations of turkeys by deer hunters. We illustrate that population growth is affected
by variation in both reproduction and fall harvest rates. The population decline was
apparently caused by reduced recruitment in combination with reduced hen survival
associated with initiation of fall hunting. The decline in recruitment was associated with
cold and wet spring weather (r^= 0.92, P < 0.001). Analysis of 103 years of weather
data suggested the low level of recruitment observed during 1989-92 may be
representative of the long-term recruitment potential of this population. More
conservative harvest strategies may be appropriate for northern populations. Key
words: eastern turkey, precipitation, temperature, population dynamics,
recruitment, model, hunting

426

Roseberry, J. L. 1964. Some responses of bobwhites to snow cover in southern
Illinois. Journal of Wildlife Management 28(2): 244-249. Abstract: An uncommonly
long period of snow coverage early in 1960 (23 consecutive days with 3-16 inches of
snow) afforded an opportunity to observe responses of bobwhites (Colinus virginianus)
to adverse weather conditions. Daily contact was maintained with four coveys on the
1,600-acre Carbondale Research Area in southern Illinois. Roosting and loafing sites

% ^>>^

191

£Z^> cC^

were shifted from open to woody cover, especially clumps of Japanese honeysuckle
(Lonicerajaponica). As waste grain in harvested fields was covered by snow, feeding
was generally confined to small patches of unharvested corn and soybeans adjacent to
woody cover. Coveys were more sedentary during the snow coverage. Daily
movements varied from 70 to 600 yards, averaging 265 yards. Three late winter ranges
averaging 23.7 acres were reduced during the snow to include only woody cover and
unharvested cropland; one covey utilized only 3.3 acres for a period of 7 days. An
estimated 29 birds perished out of a pre-snow population of 162; losses appeared to
correlate with quality of winter range, especially the availability of agricultural grains.
Key words: bobwhite quail, snow, habitat use, movement, mortality, habitat
quality

427

. 1962. Avian mortality in southern Illinois resulting from severe weather

conditions. Ecology 43(4):739-740. Summary: A prolonged period of snow coverage
in early 1960 increased avian mortality in parts of southern Illinois. Meadowlarks and
bobwhites constituted 45% of 112 avian deaths observed. Starlings and other
blackbirds comprised 20% of the recorded mortality. The severe weather may have had
an adverse effect on bobwhite fecundity during the subsequent breeding season. A
large amount of available corn and soybeans possibly prevented mortality from being
much greater. Key words: bobwhite quail, snow, fecundity, mortality

428

and W. D. Klimstra. 1972. Some aspects of the dynamics of a hunted

bobwhite population. Proceedings of the National Quail Symposium 1:268-282.

Abstract: Multiple regression analysis showed that annual rates of productivity were
significantly influenced by the combined effect of breeding density, length of snow cover
during the previous 2 winters, and amounts of prenesting rainfall. It was not clear
whether 2 severe winters caused the apparent cyclic regularity or merely accentuated
the lows. Key words: bobwhite quail, snow, precipitation, productivity, severe
weather

429

Rosene, W. 1969. The bobwhite quail: its life and management. Rutgers
University Press, New Brunswick, USA. Summary: Chapter 9 Natural Mortality,
contains a lengthy discussion under the subheading Weather as a Factor. The author
notes the differences in the impacts of weather events in terms of severity and/or
combinations of events, e.g. rain and low temperature leading to icing, etc. He also
relates these occurrences to impacts depending on the lifecycle stage of the bobwhite
quail in several areas of the United States. Key words: bobwhite quail, temperature,
precipitation, ice, snow, wind, clouds, behavior, mortality, habitat use,
reproduction

192 ^

4

^ te ^

cCi

430

Rowland, M. M. and M. J. Wisdom. 2002. Research problem analysis for greater
sage-grouse in Oregon. Final report. Oregon Department of Fish and Wildlife; U.
S. Department of the Interior, Bureau of Land Management, Oregon/Washington
State Office; and U. S. Department of Agriculture, Forest Service, Pacific
Northwest Research Station, Portland, USA. Summary: On page 15, the authors
review the influence of weather and climate upon sage-grouse stating: "Weather
patterns frequently have been linked to sage-grouse abundance and nest success,
primarily through the influence of moisture and temperature on abundance and
phenology of herbaceous plants used as forage and cover. Studies in a Wyoming big
sagebrush community in central Oregon showed wide fluctuations in both forb and total
herbaceous production, as well as plant species numbers, in response to variation in
annual precipitation. Willis et al (1993) found no relationship between long-term trends
in sage-grouse productivity and precipitation. Weather, however, may influence timing
of seasonal movements in sage-grouse as well as diets. Call and Maser (1985)
reported that sage-grouse densities in Oregon were greatest in areas of 25-40 cm of
annual precipitation. Further analyses of relationships between long-term trends in
sage-grouse productivity and weather should be conducted to better understand these
interactions." A citations list is provided. Key words: sage-grouse, precipitation,
temperature, vegetation, productivity

431

Rozycki, A. L. M. Keller, and T. Buczek. 2007. Numbers and habitat preferences of
the hazel grouse Bonasa bonasia in the Lasy Parczewskie Forest. Notatki
Ornitologiczne 48(3):151-162. Abstract: The study was carried out in 2002-2004 in the
Lasy Parczewskie Forest (eastern Poland) covering c. 7500 ha by the annual territory
mapping method performed in two stages: in the autumn-winter season the whole area
was walked over by several people in extended order and then in early spring the prior
localized sites of the hazel grouse were verified. In total, 243 observations of birds were
made and evidence of their occurrence — feces, feathers or track — recorded. The
species abundance was estimated at 104-116 territories, yielding a density of 1.4-1.6
territories/km2 which made this population one of the most numerous in the belt of
central Poland. A clear increase in the hazel grouse numbers was noted relative to the
early 1990s. The paper discusses hypotheses that associate the increasing trend with a
few factors which are not mutually exclusive: (1 ) advantageous habitat changes
connected with formerly open peat-bogs getting overgrown with birches due to drainage
of the terrain (land reclamation in the 1960s and 1970s); (2) introduction by foresters of
the spruce into the understory; (3) a series of warm winters in a dozen or so years
preceding the study period; (4) favorable meteorological conditions in the season of
young rearing, and (5) a decline in the goshawk {Accipiter gentilis) abundance. The
basic food items during winter are birch and hazel buds. The sheltering function is
played by spruce undergrowth or, alternatively, pine stickstands, especially ones with a
hazel understory. Tree stands of younger age category (20-40 years) and old-growth
pine forest with dense hazel underbrush (coverage level of 0.5-0.7) are decidedly more
preferred. Typical sites where the hazel grouse occurs are ecotone zones between
young birchstands and old-growth pine forest with spruce in the understory. The more

^ 193

% r>(<c*^<

% i$> O 9

mosaic in character the vegetation cover( diverse species composition and age
structure) and surface features the more attractive the site to hazel grouse. Key words:
hazel grouse, temperature, reproduction

432

Rumble, M. A., F. Wakeling, and L. D. Flake. 2003. Factors affecting survival and
recruitment of female Merriam's turkeys. Intermountain Journal of Science 9:26-

37. Notes: Based on personal observations, we conclude that predation and weather
are important contributors to early mortality of Merriam's turkey poults. Key words:
Merriam's turkey, mortality, predation, weather

433

and S. H. Anderson. 1996. Microhabitats of Merriam's turkeys in the Black

Hills, South Dakota. Ecological Applications 6(1):326-334. Abstract: Merriam's
turkeys (Meleagris gallopavo merriami) are associated with ponderosa pine (Pinus
ponderosa) forests in the western United States, but are not native to the ponderosa
pine forest of the Black Hills, South Dakota. The Black Hills population was established
by transplanting birds from New Mexico and Colorado between 1948 and 1951. Despite
being outside its original range, this population provides a unique opportunity to assess
mechanisms of habitat selection because the age of the population is known and
literature indicates that it is more productive than other populations. We studied
microhabitats of Merriam's turkeys in the Black Hills between 1986 and 1991. We found
few differences in microhabitats among diurnal time periods or between sexes. Turkeys
preferred southern exposures during winter. Production of pine seed, a major food item
of turkeys, differed among years. There was a strong relationship between abundance
of pine seeds and microhabitats selected by turkeys. Basal area of microhabitats
between October and March was positively correlated with annual ponderosa pine seed
production. Abundance of ponderosa pine seeds at turkey microhabitats during this
period was at least four times the estimated average annual production. Management
prescriptions for ponderosa pine of basal area < 1 8 m2/ha will reduce winter habitat for
turkeys. Summer habitats are more compatible with timber management goals for
ponderosa pine in the Black Hills. Key words: turkey, habitat use, habitat
management, temperature, microhabitat

434

and R. A. Hodorff. 1993. Nesting ecology of Merriam's turkeys in the Black

Hills, South Dakota. Journal of Wildlife Management 57(4):789-801. Abstract:
Merriam's wild turkeys (Meleagris gallopavo merriami) were introduced to the Black
Hills approximately 40 years ago, and recently population estimates show a large and
stable population. Until now, few studies have evaluated nesting ecology of Merriam's
turkeys, and none occurred in predominantly pure ponderosa pine (Pinus ponderosa)
forests. Thus, we studied nesting and nest habitat factors that influence population
productivity using a hierarchical approach in the Black Hills, South Dakota. In contrast
to other studies, yearling Merriam's turkey hens showed a high propensity to nest. Nest
survival for adult hens did not differ (P= 0.18) from yearlings, but adult hen success
was higher (P = 0.03). April-June precipitation was positively related (R2 = 0.93, P <

194 ^

<^

H to* ^

0.01) to the number of nest attempts. Primary nest predators were mammals and
American crows (Corvus brachyrhynchos; hereafter referred to as crows). Among
macrohabitats (third-order habitats), there were no (P = 0.45) patterns of nest site
selection. Among microhabitats (fourth-order habitats), hens selected small sites (< 5 m
across) with obstructed view of the nest and vegetation averaging 2.3 dm tall. Few
microhabitat differences occurred between successful and unsuccessful nests, and
those that did were related to higher (P = 0.02) survival of third nest attempts. Our data
indicate that availability of nest habitat was not limiting turkey populations in this area.
Key words: turkey, precipitation, productivity

435

Ruth, J. M. 1972. Influence of weather on movement and habitat use of hen
pheasants during brood rearing. M.S. Thesis, South Dakota State University,
Brookings, USA. Notes: The objective of this study was to determine the effect of
selected weather conditions on the movement and habitat use of hen pheasants during
the brood-rearing period. Habitat-use indices were compiled to determine habitat
preferences. An index for each weather class was prepared for each of the 3 years of
study. These indices were then combined and a weighted mean index value calculated
for each weather class. Alfalfa was the preferred cover type under all conditions. Under
low barometric pressure conditions, alfalfa, corn and residual cover were the preferred
habitat types with indices over 1 .0. Pasture and ditches had indices of 0.7 while all other
cover types were 0.5 or less. Under high pressure conditions the same four types
remained preferred, but residual cover was preferred over corn. Analysis of wind
velocity showed three major shifts in preference. During precipitation, preference for
residual cover and treerow and farmstead increased markedly, while preference for
alfalfa dropped rather sharply during periods of precipitation. No significant relationships
were found between distance of bird movement and any of the weather factors tested.
Normal daily fluctuations of weather elements did not affect pheasant activities to a
large degree. Under conditions of precipitation, high wind velocity, and low barometric
pressure the preference index for alfalfa was much lower. With these same conditions
an increase in use of corn and residual cover was noted. This seems to be a logical
shift in usage because these two cover types offered more protection from the
elements. Key words: ring-necked pheasant, precipitation, wind, barometric
pressure, habitat use, movement, index

436

Ryser, F. A. and P. R. Morrison. 1954. Cold resistance in the young ring-necked
pheasant. Auk 71:253-266. Summary: The development of cold resistance was
studied in young ring-necked pheasants. Although living in a hot thermal environment
the body temperature of the chick is lower than that of the adult, and the regulated level
of body temperature gradually rises during the first few weeks of life in a brooder to the
adult level. During this period (2 to 15 days), exposure to moderate cold (20°) results in
substantial losses in body temperature (2 to 4°). When two- to three-day-old chicks are
repeatedly chilled by 30-mnute exposures at 20°, the development of cold resistance is
impaired, and they experience a high rate of mortality. Chicks of seven days and older
are not adversely affected by repeated exposures at 20°, which instead improve their

y 195

% r>ikr*^<

c^2>

resistance to cold. Key words: ring-necked pheasant, temperature,
thermoregulation, mortality

437

Saab, V. A. and J. S. Marks 1992. Summer habitat use by Columbian sharp-tailed
grouse in western Idaho. Western North American Naturalist 52(2):166-173.

Abstract: We studied habitat use by Columbian sharp-tailed grouse {Tympanuchus
phasianellus columbianus) in western Idaho during 1983-85. Vegetative and
topographic measurements were recorded at 716 locations of 15 radio-tagged grouse
and at 180 random sites within the major vegetation/cover types in the study area. The
mean size of summer home ranges was 1 .87 ± 1 .14 km2. Of eight cover types identified
in the study area, individual grouse used the big sagebrush {Artemisia tridentate) cover
type more than or in proportion to availability, the low sagebrush (A. arbuscula) in
proportion to availability, and avoided the shrubby eriogonum (Eriogonum spp) type.
Characteristics of the big sagebrush cover type that sharp-tailed grouse preferred
include moderate vegetative cover, high plant species diversity, and high structural
diversity. Grouse used areas of dense cover (i.e., mountain shrub and riparian cover
types) primarily for escape cover. Compared with random sites, grouse selected areas
with (1) greater horizontal and vertical cover, (2) greater canopy coverage of forbs
typically decreased by livestock grazing, (3) greater density and canopy coverage of
arrowleaf balsamroot (Balsamorhiza sagittata), and (4) greater canopy coverage of
Bluebunch wheatgrass (Agropyron spicatum) in the big sagebrush cover type in 1984
and the low sagebrush cover type in 1985. The importance of the native perennials
arrowleaf balsamroot and Bluebunch wheatgrass became apparent during a drought
year when many exotic annuals dried up and provided no cover. Overall grouse
selected vegetative communities that were least modified by livestock grazing. Key
words: sharp-tailed grouse, vegetation, habitat use, grazing, drought,
temperature

438

Sandercock, B. K., K. Martin, and S. J. Hannon. 2005. Life history strategies in
extreme environments: comparative demography of arctic and alpine ptarmigan.
Ecology 86(8):21 76-21 86. Abstract: Arctic and alpine habitats are extreme
environments characterized by short breeding seasons, cold temperatures, limited food
availability, and potentially high predation rates. Stringent ecological conditions are
likely to have important consequences for the evolution of life history traits, but direct
empirical tests are few. We compare the demography of three populations of ptarmigan
on an environmental gradient spanning alpine, subalpine, and arctic habitats. Female
willow ptarmigan (Lagopus lagos) and white-tailed ptarmigan (L. leucurus) breeding at
subalpine and alpine sites had smaller clutches and lower probabilities of nesting
success, fledging success, and renesting than willow ptarmigan nesting at a low-
elevation arctic site. Annual fecundity, measured as female fledglings per breeding
female, did not overlap among the three populations and was ranked: alpine (0.40 ±
0.08, mean±SE, 95% CI = 0.26-0.58) < subalpine (1.33 ± 0.10, 1.13-1.54) < arctic
(2.04 ± 0.18, 1.68-2.39). There was a nonsignificant trend for apparent survival rates
(<D) of breeding females to vary in the opposite direction: alpine (0.46 ± 0.04) >

196 ^

4>

fa <C& f*

subalpine (0.43 ± 0.03) > arctic (0.37 ± 0.06). Population growth rates predicted
significant declines for the alpine population (A = 0.65 ± 0.07, 95% CI = 0.52-0.79), but
not the subalpine (A = 1.00 ± 0.07, 0.86-1.14) or arctic populations (A = 1.13 ± 0.20,
0.78-1.54). The adjusted estimates of survival necessary to sustain a stationary
population indicated that actual variation in female survival was more pronounced than
the observed rates: alpine (0.71) > subalpine (0.43) > arctic (0.33). Together, the
fecundity and survival values provide evidence that even congeneric populations can
exhibit a continuum between high reproductive and survivor life history strategies.
Variation in ptarmigan life history traits was consistent with population differences in
predation rates on eggs and breeding females, and it was not related to duration of the
breeding season, climatic conditions, or food availability. Ptarmigan demography also
covaried with body size, but not in the predicted pattern. Overall, the life history
strategies of ptarmigan are consistent with our current understanding of the impacts of
environmental factors upon life history variation in passerine songbirds. Key words:
willow ptarmigan, white-tailed ptarmigan, fecundity, reproduction, temperature

439

Saniga, M. 2002. Nest loss and chick mortality in capercaillie {Tetrao urogallus)
and hazel grouse (Bonasa bonasia) in west Carpathians. Folia Zoologia 51(3):205-
214. Notes: In the years with very cold weather during May (heavy snowfall), nests were
destroyed by snow cover and abandoned (20%). Nine clutches (6%) were found
abandoned, their hens probably having been predated by goshawk, golden eagle, or
Ural owl, or by some of the mammalian predators: lynx, red fox, marten. Predation
pressure on capercaillie and hazel grouse nests decreased significantly during the
incubation period. A decrease in nest losses during the incubation period was expected
as, at the time of egg-laying, there only birds of prey and owls are breeding. Capercaillie
and hazel grouse nest quite early in the spring prior to the onset of breeding of other
resident and migratory birds. Thus the predation pressure on the two forest-dwelling
grouse species is much higher in the first half of May than later when the forest habitat
is inhabited by 53-59 breeding bird species. A second factor is that nests placed on the
ground at the beginning of May may be exposed to predators until the vegetation has
adequately developed. Like the study of Angelstam on black grouse, I recorded a
pronounced increase in capercaillie hen mortality during the nesting period than hazel
grouse hens. This was not surprising, as the larger capercaillie females mate about 10-
15 days earlier than the hazel grouse hens. Presumably, they are therefore even more
dependent on access to the limited snow-free patches and emerging new vegetation,
thus putting themselves at a high risk of predation. When rodent density crashed during
winter, the predation pressure on capercaillie and hazel grouse nests increased
significantly during the incubation period of the following year. Nest losses in hazel
grouse were much higher in years when the weather conditions during May and June
were unsuitable - heavy rains or snowfalls. Monthly mortality of hazel grouse chicks
was 10-12% in the Ural region and was correlated with the weather conditions during
late spring and early summer. Key words: capercaillie, hazel grouse, snow,
precipitation, predation, mortality

* >' t* **

x **"

e

■% ?©>> <m^ $>

440

Savage, D. E. 1969. Relation of sage-grouse to upland meadows in Nevada.
Nevada Fish and Game Commission. Transactions California-Nevada Section The
Wildlife Society:8-17. Abstract: Meadow utilization by sage-grouse was studied for two
summers at three locations in Nevada. The principal methods used included
observation and collection of birds and vegetational description of the meadow and
surrounding sagebrush vegetation types. Movements to the meadow areas were
apparently stimulated by the desiccation of vegetation in the sagebrush types.
Meadows were used almost exclusively by hens and young birds with only a few cocks
being observed in the drier of two summers studied. A pattern of use was evident while
the birds were on the meadows. Variations in use, however, occurred with varying
climatological conditions. Segregation of cocks, hens with chicks and hens without
chicks was evident, but the degree of segregation was also dependent upon
climatological conditions. Hens and chicks, however, would congregate only at those
portions of meadows having some meadow vegetation. Bare portions of the meadows
or water sources lacking the desired vegetation would be ignored by this group. Food
habits of the various ages and sexes of birds showed a high preference for certain
species of succulent forbs by young birds and hens. The meadow, as a sole source of
the desired food species, became a necessity to these groups. Consumption of forbs
decreased with age in the young birds and by fall with the older birds. Movement away
from the meadows was associated with the decreasing temperatures and precipitation
of fall Keywords: sage-grouse, precipitation, temperature, vegetation, habitat
use, movement

441

Sayre, M. W. and N. J. Silvy. 1993. Nesting and Production. Pages 81-104 in T. W.
Baskett, M. W. Sayre, R. E. Tomlinson, and R. E. Mirarchi, editors. Ecology and
management of the mourning dove. Wildlife Management Institute, Stackpole
Books, Harrisburg, USA. Notes: Page 101 - "Weather. Weather is thought to be an
important source of nest loss. It is not uncommon to find broken dove eggs on the
ground under nests after thunderstorms or periods of high winds. In Texas, Morrow and
Silvy (1982) found significant correlations between monthly estimates of nest mortality
and the number of days in which wind speed exceeded 15 knots per hour (27.6 km/hr).
Mean monthly temperature also was positively correlated with nest success, indicating
that below-normal temperatures may adversely affect nesting success. Yahner (1983)
found that 7 percent of nest loss in a Minnesota shelterbelt could be attributable to
weather conditions". Key words: mourning dove, thunderstorms, wind,
temperature, solar radiation, mortality

442

Schemnitz, S. D. 1961. Ecology of the scaled quail in the Oklahoma Panhandle.
Wildlife Monographs 8:3-47. Notes: Page 40 — "During the winter period, the birds
remained closely associated with the protective shelter provided by shrubs and man-
made cover." The author also notes the scaled quail used shrubs for shelter from the
sun Key words: scaled quail, temperature, habitat use, vegetation

J*
4

^ftefc**

cCi

443

. 1964. Comparative ecology of bobwhite and scaled quail in the Oklahoma

Panhandle. American Midland Naturalist 71(2):429-433. Abstract: Bobwhite quail
were encountered primarily in the riparian habitat in Cimarros County while the scaled
quail occupied parts of the short grass, sandsage grassland, and pihon-juniper
vegetation types. During a severe drought period, 1954-56, scaled quail populations
remained at a high level while bobwhite quail numbers diminished. The early winter diet
of bobwhite and scaled quail was similar. Scaled quail mingled in larger coveys during
the winter period than did bobwhites. Key words: bobwhite, scaled quail, drought,
population dynamics, behavior

Schmidt, P. A., J. A. Hanf, and E. B. Groshens. 1994. Relationship between
precipitation and number of males on leks in central Oregon: a model to predict
number of sage-grouse. Proceedings of the Western States Sage and Columbian
Sharp-tailed Grouse Workshop 29. Unable to obtain copy for abstract.

445

Schneegas, E. R. 1967. Sage-grouse and sagebrush control. Transactions of the
North American Wildlife and Natural Resource Conference 32:270-274. Notes:
Page 274 — "The decline of sage-grouse populations after 1930 was probably caused by
unfavorable changes in vegetative types resulting from drought and overuse by
livestock." Key words: sage-grouse, drought, vegetation, habitat use, grazing

446

Schroeder, M. A. 1997. Unusually high reproductive effort by sage-grouse in a
fragmented habitat in north-central Washington. Condor 99:933-941. Summary:
The annual variation in date of nest initiation observed in this study of almost 2 weeks
was similar to that recorded in other studies. The variation in nest initiation appeared to
be related to annual variation in weather; cold, snowy winters and/or cold, wet springs
appeared to result in later dates of nest initiation. Key words: sage-grouse,
temperature, snow, precipitation, reproduction

447

Schroder, W., J. Schroder, and W. Scherzinger. 1982. Uber die rolle der Witterung
in der populationsdynamik des Auerhuhns (Tetrao urogallus). Journal of
Ornithology 123(3):287-296. Summary: This study examines the hypothesis that
random yearly variation in weather conditions can cause long term cycles in capercaillie
populations. Observations in captivity as well as in the field indicate that air temperature
and precipitation during the first few weeks after birth have an effect on the survival of
chicks. The station Hohenpeiflenberg of the German Weather Service has
measurements of such values for the past 200 and 100 years respectively. The effect of
these weather factors over many years was studied in a population simulation. The
results support the hypothesis that phases of favorable and unfavorable population

* l^f i . t. 1"

% Hte***

<gf ^gjf- ^^ <£>

development can occur, caused merely by random year to year variation in the weather
after hatching of capercaillie chicks. The simulated population trends correspond to
historical records of increase and decrease, expansion and decline of the species. Key
words: capercaillie, temperature, precipitation, mortality, model, population
dynamics

448

Schwertner, T. W., M. J. Peterson, and N. J. Silvy. 2005. Effect of precipitation on
Rio Grande wild turkey poult production in Texas. Proceedings of the National
Wild Turkey Symposium 9:10-15. Abstract: Precipitation can strongly influence the
population dynamics of gallinaceous birds in semiarid regions. Little is known, however,
about the interaction of precipitation and Rio Grande wild turkey {Meleagris gallopavo
intermedia] RGWT) production in Texas, particularly across broad spatial and temporal
scales. We compared RGWT production data with precipitation and drought data across
5 ecological regions of Texas for 1976-2000. Poult production was positively correlated
with both the June Modified Palmer Drought Severity Index (PMDI) and September-
June raw precipitation in all ecological regions. We found weaker correlations with June
raw precipitation in all ecological regions except the Post Oak Savannah, and with
cumulative September-June PMDI in the Edwards Plateau, Cross Timbers and Prairies,
and South Texas Plains. Our results indicate that poult production is more influenced by
cumulative weather effect over several months than by individual rainfall events,
suggesting that direct precipitation-induced mortality does not substantially affect
RGWT production in Texas. Further, precipitation data provides managers with an
inexpensive, effective indicator of RGWT production in Texas. Key words: Rio Grande
turkey, precipitation, drought, production, Palmer Drought Severity Index, index

449

Scott, T. D. 1937. Snow-killing of the bobwhite. Wilson Bulletin 49(1):21-27.

Summary: Reasonably healthy bobwhites may perish through imprisonment by drifting
snow. Exposure to cold, high winds and snow may kill reasonably healthy bobwhites.
Cover subject to heavy drifting is not ideal. Key words: bobwhite quail, snow,
temperature, wind, mortality

450

Sedvarsky, W. D. 1988. Reproductive ecology of female greater prairie-chickens
in Minnesota. Pages 193-239 in A. T. Bergerud and M. W. Gratson, editors.
Adaptive strategies and population ecology of northern grouse. Wildlife
Management Institute, Washington D.C. and University of Minnesota Press,
Minneapolis, USA. Summary: Page 239 — The poor drainage of the study area and the
early summer peaks in precipitation were believed to contribute to poor brood survival.
Exposure to precipitation and cool temperatures were affected by hatching dates,
characteristics of brood habitats, and extent of movements as influenced by disturbance
and proximity of nest sites to brood cover. Climatic factors may affect chick survival
directly or indirectly by affecting insect populations. Key words: greater prairie-
chicken, temperature, precipitation, population dynamics, mortality, insects,
habitat use

200 j,

II

>-> fc* **

451

Seiler, T. P., R. D. Drobney, and T. V. Dailey. 2002. Use of weather variables for
predicting fall covey calling rates of northern bobwhites. Proceedings of the
National Quail Symposium 5:91-98. Abstract: A newly developed technique for
estimating fall northern bobwhite (Colinus virginianus) density is currently being
employed in parts of the United States. One aspect of this technique involved predicting
morning covey calling rates (i.e., the proportion of coveys that call on a given morning).
We monitored 60 radiomarked coveys, a total of 229 covey observations, to determine
whether or not each covey called. Calling rates were evaluated in relation to date, year,
area, temperature, relative humidity, barometric pressure, barometric status, cloud
coverage, and wind speed. We used logistic regression to test 9 a priori models as
predictive models of bobwhite covey calling behavior. Models were compared using
Akaike information criteria (AlCc) values to determine the relative importance of 6
different variables (wind speed, date, temperature, cloud coverage, barometric
pressure, and relative humidity). An exploratory analysis was then conducted to find the
best predictive model using the best subsets model selection procedure. Standard
errors of the coefficients in the best models were calculated using a traditional
bootstrapping technique. We found an overall calling rate of 78%. Wind speed and date
were the most influential of the 6 variables used in a priori model tests. Nine of the 19
exploratory models fit the data reasonably well. The best model included area and wind
speed as independent variables, and was a better model than the best a priori model.
There was a difference in calling rates between areas, and as a consequence, we
recommend caution in application of our models to new areas. Key words: bobwhite
quail, wind speed, temperature, clouds, barometric pressure, humidity, behavior,
census, model

452

Selas, V. 2006. Patterns in grouse and woodcock Scolopax rusticola hunting
yields from central Norway 1901-24 do not support the alternative prey
hypothesis for grouse cycles. Ibis 148:678-686. Abstract: According to the
alternative prey hypothesis, autumn populations of ground-nesting game birds fluctuate
in synchrony with vole numbers because generalist predators that mainly eat voles
switch to alternative prey, such as eggs and chicks, when vole numbers decline. In
hunting statistics from Nord-Trondelag, central Norway, 1901-24, annual fluctuations in
the number of willow grouse Lagopus lagopus and western capercaillie Tetrao
urogailus, but not of woodcock Scolopax rusticola, were positively related to vole
numbers in the current year. Both woodcock and grouse indices were related to hunting
indices of Goshawk Accipiter gentilis and to weather variables assumed to influence the
birds' survival or reproduction, suggesting that the indices actually reflected local
population levels. Synchronous vole and grouse fluctuations are consistent with the
alternative prey hypothesis (although predator densities were low in the early 1900s),
but the asynchronous woodcock fluctuations refute the hypothesis. Rather, because the
woodcock does not feed on plants utilized by voles and grouse, I suggest that food
quality is the ultimate factor for the synchrony in vole and grouse numbers in Norway.
Key words: willow grouse, capercaillie, woodcock, reproduction, nutrition

x

f>f <&.* x*

201

<2& ^Jsfe. dZ^j> cCi

453

. 2001. Autumn population size of capercaillie Tetrao urogallus in relation to

bilberry Vaccinium myrtillus production and weather: an analysis of Norwegian
game reports. Wildlife Biology 7(1 ):1 7-25. Summary: From a study area in Aust-
Agder, southern Norway, game reports from 1920-1978, supplemented with autumn
counts carried out during 1968-1984, were used to determine whether the autumn
population size of capercaillie Tetrao urogallus showed no increase, a slight increase or
a strong increase compared to the population size the previous year. Based on the
mast depression hypothesis, it was predicted that adverse weather conditions should
have less influence on the reproduction and thus also on the autumn population size in
post-mast years of bilberry Vaccinium myrtillus than in other years, because of a higher
food quality and therefore also a higher body condition of the birds. In a logistic
regression model, only the bilberry index and the June-September temperature of the
previous year contributed significantly to explain the status of the capercaillie
population. A negative effect of high summer temperatures in the previous year was
highly significant when analyzing post-mast years separately, possibly because bilberry
plants were less depressed after a high seed crop if summer temperatures and thus
primary production were high. Only when years with high bilberry production were
analyzed separately, did I find effects of weather conditions which could be assumed to
have direct impacts on breeding success, such as snow conditions in spring and
precipitation in early summer. Key words: capercaillie, productivity, nutrition,
temperature, snow, precipitation

454

. 2000. Population dynamics of capercaillie Tetrao urogallus in relation to

bilberry Vaccinium myrtillus production in southern Norway. Wildlife Biology
6(1):1-11. Abstract: Forester T. Grasaas' data on numbers of capercaillie Tetrao
urogallus cocks and hens observed at leks and clutch sizes in Vegarshei, southern
Norway, during 1953-1962 (high population level) and 1969-1978 (low population level)
were analyzed with regard to bilberry Vaccinium myrtillus production, autumn population
indices and snow conditions in spring. From the mast depression hypothesis, it was
predicted that the number of capercaillies counted at leks and the mean clutch size
should be high after high seed crops (masts) of bilberry, usually produced at intervals of
3-5 years. In stepwise regression models, both the bilberry index of the preceding year
and the autumn population index one (hens) or two (cocks) years earlier contributed to
explain the mean number of capercaillies counted at leks during 1970-1978.
Capercaillie clutch sizes were highest in years with early thaw, but the effect was
significant only for the period 1969-1978. For this period, there also was a positive
effect of the bilberry index and a negative effect of the autumn population index of the
previous year. It is concluded that the synchronous population fluctuation of grouse and
voles in Norway and Sweden cannot be explained by the alternative prey hypothesis
alone, and that food quality should also be considered as a possible contributing factor
when analyzing population fluctuations of grouse and other herbivorous species. Key
words: capercaillie, snow, population dynamics, vegetation, diet

202

-¥
4

*^fte***

cCi

455

Seiskari, P. 1962. On the winter ecology of the capercaillie, Tetrao urogallus and
the black grouse, Lyrurus tetrix, in Finland. Riistatieteellisia Julkaisuja 22:3-119.

Abstract: The present investigation deals with the winter habitat requirements of two
sympatric species and the factors influencing them in Finland. Attention has mainly
been devoted to the feeding ecology and feeding activities in winter. The capercaillie
and the black grouse have very specialized habitat requirements in winter. The ultimate
cause of these is adaptation to a single food plant. The food plants seem to act as an
important proximate factor in habitat selection, too. Openness of the environment in the
shelter habitat is required by the black grouse and north Finish capercaillie, since they
roost in the snow. The southern capercaillie seem to seek shelter among the branches
of the spruce, however, as this tree species is important in the shelter habitat in winter.
The male capercaillie selects as feeding plants pines of a certain structure and chooses
its habitat on this basis. The feeding habitat requirements of the female are based on
structural features of the environment. The black grouse selects its habitat according to
the amount of its favorite food, catkins, and the edge effect also influences habitat
selection. The capercaillie and the black grouse have clearly become adapted to
different niches, as have the different sexes of the capercaillie, owning to great
morphological sex differences. This is reflected in many differences in activities and
habitats. The shift to winter food is more clearly correlated with temperature than with
snow cover. As a rule, most of the ground is still under snow at the time the birds begin
to feed on the ground in the spring. Of the external factors influencing the winter feeding
the shortness of the light period puts a limit to feeding. The capercaillie takes more food
in late than in mid-winter, probably owing to the lengthening of the light period. There is
no comparable phenomenon in the black grouse, probably because a filled crop suffices
to satisfy the food requirements of one day, whereas the contents of a filled crop seem
to be insufficient for the capercaillie. Feeding activity does not become bimodal until the
end of Feb., which seems to mean that the birds have time to fill their crops twice a day.
Short days reduce the amount of food consumed by less than half that consumed
during long days, since the capercaillie feeds throughout the day during short days, and
there probably is a continuous food stream from the crop to the gizzard. The feeding
activity of both species shows a rise in March. The feeding activity is correlated with
changes in the weather. A change in weather stimulates the birds' general activity, even
in the night. Feeding activity is slight on such days but as a rule is intensified thereafter,
whereas general activity is reduced. Feeding activity is stimulated by cold weather and
high pressure, whereas warm weather and low pressure reduce it. The winter food
supplies of the capercaillie and the black grouse are of a different nature, since the
replacement of the food supply of the former is influenced by consumption, whereas the
replacement of catkins, which are the favorite food of the black grouse, is independent
of consumption. Perhaps this explains why the reserves of the capercaillie are markedly
larger than those of the black grouse. The adequacy of the food supply is no proximate
ecological factor for the capercaillie, although the quality of the food maybe reflected in
fluctuations of density. Fluctuations in the amount of catkins can cause density
fluctuations in the black grouse. But the same factors (climatic factors) which favor the
food plants may be important as proximate factors influencing the habitat requirements
of both species. In the winter community to which the capercaillie and the black grouse
belong different animal species have adopted niches in different stages of the natural

j^ 203

i Kfc***

^h <£^

forest succession. Many species have become adapted to the pioneer stages, whereas
only a single species, the capercaillie, feeding on the vegetative parts of plants, has
become adapted to one of the 2 types of climax (pine and spruce climax). In
explanation it is suggested that the climax stage is a more hazardous food resource
because of its slow recovery after catastrophes. The effects of man on the winter
ecosystem is great, especially as he alters the natural succession. It is the species
adapted to the several stages on which silvicultural practices are concentrated that are
most strongly affected. Forestry seems to leave the pioneer stages almost unaltered,
whereas the middle-aged stages deviate greatly from the natural stages. The alteration
in the structure of the forests which has taken place during recent decades has had a
visible effect on the populations of capercaillie and the black grouse. Key words:
capercaillie, black grouse, solar radiation, habitat use, habitat modification

456

Semenov-Tyan-Shanskii, O. I. 1959. Ecology of grouse. Tr Oaplandsk Gos
Zapovednika 5:1-319. Abstract: The numbers of young in grouse populations
(particularly wood grouse) vary from year to year, depending on the weather during the
breeding season. Air temperature and precipitation during the hatching period have the
highest significance. "Peak" years and depression years usually coincide with the
widespread area from Scotland to the Urals. Embryonic deaths from unfavorable
meteorological conditions during incubation under natural conditions was investigated
(in Lapland and Pechoro-llych Forest Preserves) by means of special autographs. It
was apparent, that during unfavorable weather the setting hens abandon the nests.
More rarely, the eggs freeze at the beginning of incubation. The abandoned nests and
addled eggs are preserved in the forest for a long time and play some role in the winter
nutrition of marten and foxes, so that in the winter after a failure of tundra partridge
nesting, the significance of this food increases. Key words: wood grouse, air
temperature, precipitation, nest success, mortality

457

Sepik, G. F., W. H. Goudy, and D. R. Dessecker. 2000. Factors influencing
recruitment and condition of American woodcock in Minnesota. American
Woodcock Symposium 9:96-100. Abstract: Hunting regulations for American
woodcock (Scolopax minor) are based on information on the status of the population
from data collected during the previous hunting season and the following spring. No
surveys, however, measured the status of the population through spring and summer
prior to the hunting season. A model was developed to predict recruitment to the fall
population based on weather during the spring and summer. Also determined was
whether the condition of birds in fall reflected weather conditions prior to the hunting
season. The model was based on Minnesota weather and the recruitment index
(immatures-adult female). The model predictor was April mean minimum temperature.
The model was tested using Maine and Wisconsin data and the predicted and
estimated values for the recruitment index were correlated. The weights of birds shot in
early October differed among years and were correlated with September mean
maximum and mean minimum temperatures. The frequency of feather retention of adult
females was correlated with the recruitment index. Weather during April had the

204 jj

4-

H <<> >f*

cC^

greatest influence on the recruitment index. Key words: American woodcock,
temperature, index, model, recruitment, nutrition

458

, R. B. Owen, and T. J. Dwyer. 1983. The effect of drought on a local

woodcock population. Transactions of the Northeast Section of The Wildlife
Society 40:1-8. Unable to obtain for abstract.

459

Shaw, J. L. 1988. Epidemiology of the caecal threadworm Trichostrongylus tenuis
in red grouse (Lagopus lagopus Scoticus Lath.). Dissertation, University of
Aberdeen, UK. Abstract: The prevalence of Trichostrongylus tenuis in red grouse was
high and the distribution of parasites in their hosts highly aggregated. The prevalence
and intensity of threadworms were greater in old birds than in young ones. There was
no relationship between the body condition of individual grouse and their burden in adult
worms. On the moor, T. tenuis eggs did not develop to third-stage infective larvae
during the winter. In the summer, when development did occur, yields of third-stage
larvae were dependent on temperature. Third-stage larvae moved laterally through the
heather but were relatively short-lived and very few survived over winter. The proportion
of larvae, ingested by captive grouse, which developed to adult worms varied with
season and between individuals. Red grouse acquired little or no effective immunity to
reinfection after challenge with infective larvae. Consequently, caecal threadworms
produced a chronic infection in grouse: mature worms survived for over two years with
very little mortality. Parasite fecundity decreased as the worms aged but not with
intensity of infection. The transmission of T. tenuis is presumably influenced mainly by
the density of infective larvae, which in turn was affected by weather, particularly
temperature. A major regulatory constraint on this parasite is host mortality. Mortality,
however, is unlikely to be linearly related to parasite intensity because developing
larvae were more pathogenic than adult worms. Furthermore, there seemed to be no
important intensity-dependent processes acting to regulate T. tenuis in red grouse
populations. Key words: red grouse, temperature, disease

460

Sheldon, W. G. 1967. The book of the American woodcock. University of
Massachusetts Press, Amherst, USA. Notes: The author found that stormy weather,
including high winds with or without heavy rains, downpours with no wind, and
temperatures below freezing curtailed breeding activity of these birds, particularly
relevant to census and marking efforts. Windy nights with an air movement of more than
five miles per hour yielded poor bird catches. Still, hot, humid evenings following
afternoon thunder showers created conditions which were most consistently productive
of birds and often stimulated insect activities. Woodcock were also active on quiet
evenings, with a light drizzle or fog. Clear nights with full moon caused erratic behavior
and few birds were captured. Many woodcock die in years marked by periods of
prolonged freezing in the wintering grounds. There are fewer records of woodcock
deaths under unfavorable weather conditions during migration. Extreme cold and heavy
snow in spring undoubtedly cause woodcock mortality since the birds then are in

j, 205

N^

^ ^ ^

relatively poor condition and under the stress of breeding. Such losses have not been
detected more often because, unlike the concentrations on the wintering ground, the
birds are scattered over far-flung migration lanes. It is unlikely that heavy fall freezes
cause appreciable mortality, since the birds can move south. Sustained cold, rainy
weather during the height of the hatch probably causes some losses of downy chicks.
Some recently hatched chicks have been killed by a heavy rainstorm. It is doubtful if
drought causes woodcock deaths in their breeding grounds, since they will readily feed
on all types of invertebrates other than earthworms in summer. Key words: American
woodcock, precipitation, snow, temperature, fog, humidity, mortality, census,
technique, drought

461

Shelford, V. E. and R. E. Yeatter. 1955. Some suggested relations of prairie
chicken abundance to physical factors, especially rainfall and solar radiation.
Journal of Wildlife Management 19(2):233-242. Notes: The authors summarize the
construction and use of thermohydrograms to examine the impacts of solar radiation
and precipitation on the fecundity of prairie chickens and other game birds. They note
"The effect of temperature is in part controlled by the moisture present. Two factors
which operate in this way have been called paired factors." Also "The possible effect of
weather on cover, food, and dissemination of parasites should not be neglected."
Shelford suggests a relationship between the prairie-chicken population in northwestern
Indiana and sunshine paired with rainfall, noting "In the case of prairie chickens in
Illinois, the usual effects of sunshine and rain hold good only so long as there is plenty
of cover." Key words: prairie chicken, precipitation, temperature, clouds, solar
radiation, fecundity, technique

462

Sherfy, M. H. 1992. The influence of season, temperature and wind speed on
sage-grouse metabolism. Thesis, University of New Hampshire, Durham, USA.

Abstract: We used indirect respiration calorimetry to measure the metabolism of six
adult sage-grouse (Centrocercus urophasianus) during winter, spring, and summer.
During winter the metabolic rate of fed birds was higher (P<0.05) than that of fasted
birds. The standard metabolic rate (SMR) of females (0.692 ml_ 02. g"1 . h"1) was higher
than of males (0.583 ml_ 02 . g"1 . h 1) in winter; in both sexes SMR was higher in winter
than in summer. Females' SMR was lower (P = 0.0001) in spring than in winter. Lower
critical temperatures of both males and females were substantially lower in winter (-
0.6°C, -4.8X) than in summer (19.9°C, 14.8°C). Although seasonally elevated, the
SMR of sage-grouse in winter is low in comparison with that of other galliforms with
northern distributions. Thermoregulation during a winter night at -10°C would result in
minimal (<5%) expenditure of endogenous reserves by either sex. Thermoregulation
and SMR in winter are more energetically costly to female sage-grouse than to males,
and may necessitate increased behavioral thermoregulation by females. Seasonal
change in SMR differs between the sexes, and is probably influenced by the energetic
demands of the breeding season. Key words: sage-grouse, temperature,
thermoregulation, behavior, wind speed

206 ^

>4>

>->" l& f*

c£±

463

and P. J. Pekins. 1995. Influence of wind speed on sage-grouse

metabolism. Canadian Journal of Zoology 73:749-754. Abstract: We measured the
effect of wind speed on the metabolic rate of six adult sage-grouse (Centrocercus
urophasianus) with indirect respiration calorimetry at ambient temperatures above,
near, and below the lower critical temperature. There was a significant effect (P < 0.05)
of temperature on metabolic rate at all wind speeds, and a significant effect (P< 0.05) of
wind speed on metabolic rate for temperatures less than or equal to 0°C. Wind speed
had a more pronounced effect on metabolism at temperatures below the lower critical
temperature for sage-grouse. Metabolic rates measured at wind speeds of > 1 .5 m/s
were significantly higher than those measured at wind speeds <1 .5 m/s. Multiple
regression analysis of wind speed (u; m/s) and temperature (Ta; °C) on metabolism
(MR; ml_ 02 gA-1 hA -1 ) yielded the equation MR = 0.0837 (u) - 0.0248 (Ta) + 0.5444.
The predicted cost of thermoregulation at conditions of -5°C and u = 1.5 m/s was about
1 .5 times standard metabolic rate; half the increase was due to wind. Measurements of
wind speed in sagebrush (Artemisia spp.) stands indicate that such habitat effectively
reduces wind speeds to less than 1 .5 m/s. Microhabitat value should be recognized in
the management of sagebrush stands. Key words: sage-grouse, wind, metabolism,
habitat use, vegetation, wind speed

464

and . 1994. The influence of season, temperature, and absorptive

state on sage-grouse metabolism. Canadian Journal of Zoology 72:898-903.

Abstract: We used indirect respiration calorimetry to measure the metabolism of six
adult sage-grouse (Centrocercus urophasianus) during winter, spring, and summer.
During winter the metabolic rate of fed birds was higher (F<0.05) than that of fasted
birds. The standard metabolic rate (SMR) of females (0.692 mL 02gA-1hA-1) was
higher than of males (0.583 mL 02 gA-1 hA-1 ) in winter; in both sexes SMR was higher
in winter than in summer. Females' SMR was lower (P = 0.00001 ) in spring than in
winter. Lower critical temperatures of both males and females were substantially lower
in winter (-0.6°C, -4.8°C) than in summer (14.9°C, 14.8°C). Although seasonally
elevated, the SMR of sage-grouse in winter is low in comparison with that of other
galliforms with northern distributions. Thermoregulation during a winter night at -10°C
would result in minimal (<5%) expenditure of endogenous reserves by either sex.
Thermoregulation and SMR in winter are more energetically costly to female sage-
grouse than to males, and may necessitate increased behavioral thermoregulation by
females. Seasonal change in SMR differs between the sexes, and is probably
influenced by the energetic demands of the breeding season. Key words: sage-
grouse, temperature, metabolism, thermoregulation, behavior

465

Shields, P. W. and D. A. Duncan. 1966. Fall and winter food of California quail in
dry years. California Fish and Game 52(4):275-282. Abstract: Fall-winter diet of
California quail, Lophortyx californica, was studied on the San Joaquin Experimental
Range near O'Neals, California during November, December, and January of 1960-

% fXfe*^

207

1§f ?©£ ^^ <£>

1963. Results were compared with a 1937 study in the same area. Crops were
collected from 375 California quail during the regular hunting seasons. They were
grouped by month, and 171 chosen at random and analyzed. Diet components were
reported by month and by year, and separated into seeds and leafage. Important
differences between current results and results of the 1937 study are shown and
discussed. Varying weather conditions and resultant botanical composition caused wide
variations in quail diet. The year 1937 and preceding years were wet; the period 1960-
63 was rather dry. In 1937, seed of filaree and turkey mullein formed more than 50
percent of quail diet in November and December, but made up only 6% in the 3-year
period 1960-1963. Filaree was abundant in both periods, but early rains in 1960-1963
caused general plant germination that reduced availability of this seed. Turkey mullein
was very scarce in 1960 and 1961 and seed was not available. Far more green leafage
was taken in November and December of 1960-1962 than in 1937. This was a result of
earlier rains, general plant germination, and availability of green material in 1960-1962.
Other important differences are discussed. Key words: California quail, precipitation,
vegetation, diet

466

Sieux, J.-S. and W. Delvingt. 1997. The hazel grouse (Bonasa bonasia) in western
Ardennes: habitat, conservation measures and integration in a forestry
management. Aves 34(4): 185-1 94. Abstract: The population of the hazel grouse has
been declining for several decades in the Ardennes region and in the surrounding
areas, largely as a result of the disappearance of copses (simple or standards).
Intensive forestry and hunting management have eliminated the thick diverse
understory necessary for the species. Recent field surveys in western Ardennes have
shown that the species requires a high diversity and density of understory shrubs for
feeding purposes as well as protection against predators and bad weather. Measures
have to be taken if the species is to survive in this country, especially the maintenance
of low thickets in areas where traditional forestry does not pay. Key words: hazel
grouse, shelter, severe weather, vegetation

467

Siitari, H., J. Viitala, and M. Hovi. 2002. Behavioral evidence for ultraviolet vision
in a tetraonid species: foraging experiment with black grouse Tetrao tetrix.
Journal of Avian Biology 33(2):199-202. Abstract: In addition of wavelengths visible to
humans (400-700 nm), many birds are able to detect near ultraviolet light (320-400 nm).
Most studies of ultraviolet (UV) vision in birds have concentrated on the importance of
UV vision in intraspecific signaling, especially in passerine birds. However, birds may
also use UV vision for other purposes, e.g. foraging. We performed a laboratory
experiment to test whether a tetraonid species, black grouse Tetrao tetrix, could detect
the difference between UV-reflecting and non-UV-reflecting food items (two color
morphs of bilberry (Vaccinium myrtillus). Black grouse preferred UV-reflecting berries
when UV light was used for illumination, but showed no preference in the absence of
UV light. This observation establishes a potential UV sensitivity in this species; such a
sensitivity should be considered in behavior experiments with this species. Key words:
black grouse, UV light, behavior, feeding

208 ^

^ te* **

cd

468

Siivonen, L. 1948. Decline in numerous mammal and bird populations in north-
western Europe during the 1940's. Riistatieteellisia Julkaisuja 2:1-26. Abstract: An
exceptionally long decline, which started at the turn of the 1930's into the 1940's and is
still continuing in many areas in respect to numerous species of mammals and birds,
has been found to have occurred in Finland, Sweden, Norway, Germany, and England.
The reason for the decline has not been the same for all spp. One group is comprised
of those spp of a southern character that during the last decades have spread to the
north. Such spp are the tawny owl, partridge, pheasant, field hare, polecat, blue
titmouse, blackbird, hedgehog and roe deer. The other group is composed of spp of a
northern character, such as the woodpeckers, capercaillie, blackgame, willow grouse,
red grouse and varying hare. The spp belonging to the southern group have not been
able to withstand the increasing thickness of the snow layer and severe cold weather
during the winters of 1939-40, 1941-42, and 1942-43; and 1946. Spp belonging to the
2d group suffered a decline caused by a deviation in the short-term fluctuations in
numbers. Normally those yrs ending in the numbers 0 or 1 , in 3 or 4, as well as in 6 and
7 are peak yrs for those spp in n-w Europe. Those yrs ending in 2, 5, or 8 are normally
yrs of minima. In the 1930's the fluctuations in numbers of the spp were generally
normal. In 1939, however, normally a peak yr, the stock declined to 1/7 of the preceding
minimum (1934) and in certain cases even to 1/30 of the preceding maximum (1937).
As a result of this deviation the theoretical peak in 1940 (41) did not appear. This kind of
deviation is not unique, as it can well be compared to the so-called great deviations
which continue over 2-3 consecutive cycles. A similar deviation occurred in the 1880's.
The decline during the 1940's might have been considerably alleviated by measures
that aimed at the protection of the increasing stock of 1939. Key words: pheasant,
partridge, willow grouse, capercaillie, snow, temperature, population dynamics

469

Sime, C. A. 1991. Sage-grouse use of burned, non-burned, and seeded vegetation
communities on the Idaho National Engineering Laboratory, Idaho. M. S. Thesis,
Montana State University, Bozeman, USA. Notes: Page 42 — "Movements in 1988
and 1989 varied with precipitation. In extreme drought years like 1988, sage-grouse
were forced to leave Tractor Flats, retreating to alternate spring/summer range because
palatable forbs had desiccated by July. However, in more normal years such as 1989,
sage-grouse could meet life history requirements on Tractor Flats." Page 45 — "During
years of below normal moisture and limited forb growth such as 1988, plant phenology
is either accelerated or truncated. Thus more forbs may be removed by early livestock
grazing before the peak of grouse hatch." Page 53 — "...this tool [fire] would be most
effective in areas where adequate annual precipitation enables rapid revegetation and
in areas with a limited potential for invasion of the disturbed sites by weedy undesirable
species." Key words: sage-grouse, precipitation, vegetation, movement, habitat
use, habitat manipulation

X ^fter^

209

470

Sisson, L. 1976. The sharp-tailed grouse in Nebraska. A research study. Nebraska
Game and Parks Commission, Lincoln, USA. Notes: Under the section II Population
Studies, the author writes - Page 20 — "Regression analysis was used to evaluate the
joint relationship between sharptail numbers (standard score) and values of several
weather variables during spring counts on the Bessey area... significant terms in
equation were the first- and second-order terms of time of spring, temperature, and
relative humidity (P<0.05). The first order term of wind was significant (P<0.05), as were
the products (interactions) of time of spring and temperature, and temperature and wind
(P<0.05). Both terms of relative humidity were significant with no significant interactions
with other variables... numbers of birds... decreased at higher temperatures in
combination with low wind velocities. A possible explanation for these phenomena was
that, while unusually warm mornings were normally windy, a calm, warm morning was
often associated with changes in weather, particularly an approaching storm.
Experience indicated that attendance and courtship activity usually decreases under
such conditions... precipitation had an obvious effect on attendance and courtship
activity. During mornings with light rain, display activity decreased and fewer birds were
present." Weather variables subjected to analysis included temperature, relative
humidity, wind and cloud cover. Key words: sharp-tail grouse, temperature,
humidity, wind, clouds, behavior

471

Slagsvold, T. 1975. Production of young by willow grouse Lagopus lagopus in
relation to temperature. Norwegian Journal of Zoology 23:269-275. Abstract: The
number of young willow grouse (Lagopus lagopus) per two adults bagged during the
autumn shoots 1963-72 in Norway were compared with the temperatures prevailing
during various periods throughout the year. A significant correlation was found with
June temperatures, the period 7-16 June yielding the highest correlation coefficient.
Earlier data on the fluctuations in grouse populations (1873-1963) also lend support to
the hypothesis of a positive influence of high temperatures in June. Higher
temperatures lead to lower energy demand for body maintenance for the incubating hen
and, at the same time, accelerate the phonological development of those plants and
insects which are important as food for both hen and chicks. Key words: willow
grouse, temperature, population dynamics, nutrition, bioenergetics

472

and T. Grasaas. 1979. Autumn population size of the capercaillie Tetrao

urogallus in relation to weather. Ornis Scandinavica 10(1):37-41. Summary: Annual
fluctuations in the autumn population size of the capercaillie Tetrao urogallus at
Vegarshei in South Norway, 1920-70, were analyzed statistically in relation to various
meteorological factors to discover short-term effects. The most important weather
variables which coincided with a high population size were: an early snowmelt in spring,
only a few days of rain at the time of egg-hatching at the beginning of June and only a
few days of rain in July. These factors may have operated through their influence on the
production of young. The results are discussed in relation to data on the species from

210

H (& 1*

other countries. Key words: capercaillie, snow, temperature, precipitation,
productivity

473

Smith, J. T. 2003. Greater sage-grouse on the edge of their range: leks and
surrounding landscapes in the Dakotas. M.S. Thesis, South Dakota State
University, Brookings, USA. Notes: Page 48 — "Counts [lek] should be made from
sunrise to 8:00 A.M. on days with low to moderate wind (<10 km per hr) and no
precipitation." Key words: sage-grouse, wind, precipitation, census

474

Smyth, K. E. and D. A. Boag. 1984. Production in spruce grouse Dendragapus
canadensis franklinii and its relationship to environmental factors and population
parameters. Canadian Journal of Zoology 62(11):2250-2257. Abstract: The extent to
which a relationship between selected environmental and population parameters, and
annual production in a population of spruce grouse (D. canadensis franklinii) was
investigated through correlation analysis of data spanning 16 yr. Production of young
grouse varied between 0.0 and 1 .4 young, aged 4 or more weeks, per resident female
over the period of this study. There was no evidence that differences in annual
productivity were related to variation (maximum of 17 days) in the onset of reproduction,
a function of spring temperatures. Nevertheless, weather conditions during incubation
and the immediate posthatch period did exhibit a significant correlation with productivity;
cold, wet conditions during incubation were associated with years of poor productivity.
The same conditions during the immediate posthatch period were associated with years
of good survival of young (at least to 19 years of age). Clutch and brood sizes of
yearlings were smaller than those of adults; hence, production could be influenced by
the proportion of the population in each age category. An inverse relationship between
the density of resident females in spring and their annual production, believed to be
important in this population, was reexamined with data form an additional 6 yr. The
relationship, although still inverse, was no longer significant. Key words: spruce
grouse, temperature, precipitation, productivity

475

Snyder, W. D. 1985. Management procedures for ring-necked pheasants in
Colorado. Special Report Number 59, Project W-37-R, Colorado Division of
Wildlife, Denver, USA. Notes: Page 3 — The section of particular interest in relation to
weather is titled Predicting Population Trends Based on Environmental Factors. Over a
45-year interval, a general relationship was evident between harvest and precipitation.
Productivity was influenced by influence of rain and temperature on agricultural crops
(wheat). "Thus, precipitation received in a given year influences wheat growth that year
while the amount stored as subsoil moisture in summer fallow impacts wheat growth the
next year. Better wheat growth usually results in taller stubble which is carried as
potential nesting cover into the 3rd spring. Therefore, precipitation in any one year can
influence pheasant production over a 3-year period." Snyder then explains how
information on precipitation and temperature can be used to predict productivity and
population trends well in advance of hunting seasons. He further notes that "...one year

% f\fc*>f*

211

of below average precipitation may not have major impacts on pheasants, but a series
of drought years or a combination of drought and severe winter loss can be devastating.
Precipitation also influences the quality of cover in roadsides, waste areas, and other
habitats much in the same way that it impacts wheat. Thus, weather is the primary
agent affecting pheasants and their habitat in Colorado and can be used as a general
predictor of population trends." Key words: ring-necked pheasant, temperature,
precipitation, soil moisture, vegetation, productivity, index

476

. 1978. The bobwhite in eastern Colorado. Technical Publication No. 32.

Project W-37-R. Colorado Division of Wildlife, Denver, USA. Notes: The author
discussed weather influences on bobwhites in the section titled Environmental
Influences (page 50). Impacts include winter snow stress and winter mortality, and the
interaction of combinations of conditions. He also discussed the interaction of hunting
and snow stress and impact of precipitation on reproduction. Snyder specifically notes
that lack of brush cover in quantities that won't be buried by snow interacts to exclude
bobwhite from much of eastern Colorado. Key words: bobwhite, snow, precipitation,
temperature, mortality, stress, vegetation

477

Spidso, T. K., O. Hjeljord, and J. G. Dokk. 1997. Seasonal mortality of black
grouse Tetra tetrix during a year with little snow. Wildlife Biology 3(3/4):205-209.

Abstract: The seasonal mortality of black grouse Tetrao tetrix was studied in
southeastern Norway (60°26'N, 10°45'E), in a study area dominated by up to 80%
Norway spruce Picea abies. Modern forestry with clear-cuttings has been practiced, and
50% of the forest was younger than 30 years. Fourteen males and seven females were
captured on a lek in spring 1991 and equipped with radio transmitters. During the
following 12 months, 72% of the birds were killed, mostly by predators. No birds died
from capture through July. In autumn, predation was higher, with four black grouse
being killed. Mortality was highest during winter, with 58% of the birds dying. The
goshawk Accipiter gentilis was the most important predator during winter. High
predation by goshawks in winter 1991-92 may have been the result of black grouse
being more vulnerable to predation because limited snowfall precluded snow roosting,
or of an invasion by goshawks. Therefore, snow roosting may be an important
antipredator behavior in black grouse. The effect of increased adult mortality is
discussed in relation to chick production. Key words: black grouse, snow, predation,
mortality, habitat modification

478

Stanford, J. A. 1972. Bobwhite quail population dynamics: relationships of
weather, nesting, production patterns, fall population characteristics, and harvest
in Missouri quail. Proceedings of the National Quail Symposium 1:115-139.

Abstract: For 25 years Missouri has investigated bobwhite quail (Colinus virginianus)
behavior, production, and population response to 4 major types of weather. Ten
population parameters are examined annually to compare effects of Normal, Wet-
Deluge, Snow-Cold and Drought weather years on quail populations. Different types of

212 ^

^-

H te ^

c£^

weather are related to varying annual quail abundance by affecting productivity and
survival and influencing relative levels of annual harvest and hunter interest. Normal
and Wet-Deluge years yield favorable fall quail populations and satisfactory hunting.
Years having winters of severe snow and cold have high breeder losses, low
production, and reduced hunting success. In years having high temperature and
drought in spring and summer, quail reproduction is inhibited, resulting in high losses of
eggs and young, greatly reduced fall bird crops, and below-par hunting for many
hunters. Recovery from weather-caused population lows usually occurs within 2 or 3
years after favorable weather conditions return. Reliable techniques for sampling have
been developed to yield indices of annual production and hunting success. Production
curves show the value of data on the distribution of peaks in hatching for understanding
annual production and fall population levels of quail in Missouri. Such data form the
basis for setting annual hunting regulations of bobwhite harvest. Key words: bobwhite
quail, precipitation, snow, temperature, drought, production, abundance

479

. 1971. Bobwhite quail population responses — losses and recovery — to

excessive snowfall and low temperatures. Pages 203-237 in A. O. Haugen, editor.
Proceedings of the snow and ice in relation to wildlife and recreation symposium,
Iowa State University, Ames, USA. Abstract: Within the snow portion of the bobwhite
quail (Colinus virginianus) range, the winter period is the most critical time in the bird's
life. Severe snowfall causes exceptionally high and dramatic quail losses. In Missouri,
such losses occur on the average of once per 1 1 years. The "big snow" of 1960, its
effects on quail and the post snow recovery period, is illustrative of severe winter-quail
relationships. Heavy snow buries food and cover causing abnormal bird movement with
resulting high mortality from starvation and predation. Quail body weights decline as
much as 64%. Mortality, varying with vegetative types, agriculture, and habitat quality,
may reach 80-90%. Subsequent first hunting season success is very poor. Public
relations problems must be met with skill and firmness. Winter feeding, while fulfilling
human desires, accomplishes little. Both fed and unfed quail populations recovered
equally fast or within three nesting seasons. An animal's abundance within its
geographical range is dependent upon habitat quality and annual climatic occurrences.
Man can, to a varying degree, determine the course of habitat quality. On the effects of
climatic factors, he exerts no control except, by the management of habitat quality he
may alleviate damaging effects of severe winter weather. Within the snow portion of the
bobwhite quail range, winter and its related snowfall, is the most critical period in the life
of the bird. While weather of other seasons may impose drastic limitations on quail
production and survival, it is the winter-snow time when direct losses to potential
breeders occur. And it is the overly severe winter when such losses become
exceptionally high and dramatic. Within its optimum climatic range, quail thrive well in
the niche provided by moderate degrees of agriculture, woodland disturbance and the
resulting open ground. The annual bird surplus resulting from favorable habitat-climatic
conditions usually provides unexcelled game bird hunting. Bobwhite quail abundance or
scarcities result from one or several conditions which are categorized as either long-
term or short-term occurrences. Category number one pertains to the habitat and is
concerned with land use and vegetation control. These vegetative-land use conditions
which hinge on plant composition and time are classed long-term habitat factors.

% f*^^

213

% m <q> <$

Annual climatic conditions which affect overwintering potential breeders, production
season success and young bird survival are short-term occurrences. Results are
measured in annual bird crops which may vary considerably from year to year. Man has
no control over these factors, but his management of habitat factors can reduce bird
losses and hasten bird recoveries from climatic caused quail lows. This paper applies to
short-term production factors, which as climatic emergencies, affect annual quail
abundance. While the adverse effects of either (a) drought-high temperatures or (b)
snow-cold share notoriety as decimators of quail, the specific subject selected here
pertains to the effects of snow-cold on quail populations with special emphasis from
Missouri investigations. Key words: bobwhite quail, temperature, snow, mortality,
predation, nutrition, habitat quality, habitat modification, vegetation

480

Steen, J. B., O. Andersen, A. Saebo, H. Chr. Pedersen, and K. E. Erikstad. 1988.
Viability of newly hatched chicks of willow ptarmigan Lagopus I. lagopus. Ornis
Scandinavica 19(2):93-96. Notes: The authors document that the mortality of newly
hatched willow ptarmigan chicks was due to reduced chick viability caused by
inadequate maternal nutrition and unfavorable weather conditions during incubation.
Chemical composition of available plants prior to egg-laying indicated the quality of
maternal food; air temperature and humidity were the weather conditions of concern
during the reproductive period. Heavy rainfall during the days following mean hatching
date evidently took a far higher toll on chicks. Egg water loss was related to increase in
conductance and ambient vapor pressure. The authors state their results "demonstrate
that the poor reproductive results in 1982 was associated with eggs of poor quality.
They were smaller and had more porous shells than in the other years." Key words:
willow ptarmigan, mortality, egg quality, vegetation, nutrition, precipitation,
humidity

481

Steen, J. B., H. Steen, N. Chr. Stenseth, S. Myrberget, and V. Marestrom. 1988.
Microtine density and weather as predictors of chick production in willow
ptarmigan Lagopus I. lagopus. Oikos 51(3):367-373. Abstract: We have evaluated to
what extent biotic (predation) and abiotic (weather) factors influence annual chick
production in willow ptarmigan (Lagopus lagopus lagopus). An index of predation was
derived from data on microtine density; weather conditions from daily records of
temperature and precipitation. Stepwise regression analysis between observed chick
production on two Scandinavian study sites (Tranoy and Lovhogen) for periods of 23
and 19 years, and independent variables characterizing predation and weather
conditions suggested which variables were the most important in determining chick
production. The resulting model explains between 56% and 25% of the annual variation
in chick production; the calculated values deviating on average between 1 .14 and 0.95
checks/2 adults from the observed ones. Predation was found to be the dominant
factor, accounting for 41% and 25% of the annual variation in chick production in the
two areas. We suggest that year-to-year variation in reproductive success is, in general,
influenced to a greater degree by biotic than by abiotic factors, possibly because the

214 j^

4>

fci <o **

c£^

predators "co-adapt" to the ptarmigan. Key words: willow ptarmigan, temperature,
precipitation, predation, model, productivity

482

Stewart, P. A. 1967. Hooting of Sitka blue grouse in relation to weather, season
and time of day. Journal of Wildlife Management 31(1):28-34. Abstract: The hooting
behavior of a population of Sitka blue grouse {Dendragapus obscurus sitkensis) was
studied on Mitkof Island in southeastern Alaska, chiefly by counting hoots at different
times and under different weather conditions. Hooting was first heard on April 7 and last
on June 12, but a longer hooting season could be assumed for larger populations.
Hooting was maximum on April 26 when 627 hoot-series were heard in 1 hour. Hooting
began to decline 1 week after its peak. The level was high from April 24-May 6, and
lower but still relatively high during May 7-15. Hooting then declined markedly; during
some counting periods there was no hooting. Eight seconds was the shortest interval
observed between hooting of individual birds. Early in the season, hooting was inhibited
by inclement weather; after peak activity was reached, weather seemed to have no
effect. Hooting started in the morning 58-95 min before sunrise and continued until 19-
65 min after sunset, with a lull between 5:00 and 7:00 A.M. Key words: blue grouse,
behavior, precipitation, snow, clouds, fog, technique, index

483

Stiver, S. J. 1984. Himalayan snowcocks: Nevada's newest upland bird. Cal-Neva
Wildlife Transactions:55-57. Notes: Page 56 - "In the winter, snowcocks in their
native range are reported to descent to lower elevations, especially if snowfall is
heavy.... snowcocks are reported to prefer big game concentration areas during the
winter." Key words: Himalayan snowcock, snow, habitat use

484

Stoddard, H. L. 1931. The bobwhite quail: its habits, preservation and increase.
Charles Scribner's Sons, New York. Notes: Page 60 - "Other habits-Quail behavior in
relation to weather. When storms are threatening they may feed off and on all day,
keeping their crops full. On hot, clear days they are apt to be lazy, and if food is plentiful
most of their feeding will be carried on in the cool of the morning and very late in the
evening, and once located, they usually 'lie' well to the dog. In windy weather they
frequently fly badly, and scent is mixed up by the air currents, while, as previously
mentioned, on wet days they are inclined to run and only flush when compelled to, a
fact taken advantage of in by-gone days when netting was in vogue." Page 123 - "The
character of the food of the bobwhite varies with its availability, and availability is
governed by the weather, by fruiting habits of plants from which food is obtained, by
season, by environment, and by other factors. During years of either severe drought or
excessive rainfall the seed crop of many plants is affected. Furthermore, when swampy
areas dry up, this makes available many foods that are little used at other times. Plants
such as the pine seed abundantly once in several years. Berries and insects are more
abundant during the warmer months, and seeds during the late fall and winter. Food
plants are not equally abundant over the whole range. It is readily seen that these
several constantly changing factors greatly vary the availability of food." Page 200 -

f \ ^ ^

j<

£Zh> c^>

"Loss due to the elements. There is a general belief among rural people that many quail
chicks are 'drowned out' during heavy rains. As torrential rainstorms, or cloudbursts, do
occur at times and with such severity that newly hatched chicks could scarcely escape,
it is probable that considerable loss sometimes is occasioned in restricted areas.
Experience in the field and in the artificial propagation of these birds, however, would
indicate that actual drowning of quail chicks is exceptional and largely confined to such
as are caught in ditches and low spots. Quail chicks frequently get into ditches and
become lost from the brood, and all such would drown in ordinary rainstorms. Where
one or both parents are with the broods on well-drained ground, the chicks are
protected perfectly during the heaviest rains. The experience of the Investigation would
indicate that one important reason why very wet years are poor seasons for quail
reproduction is that considerable destruction and desertion of nests are occasioned by
torrential rainstorms. If rains are cold and of very long duration, a comparatively rare
thing in the South, quail chicks also become wet and chilled; chilled chicks seldom
recover in captivity, and the same may be expected afield. Chicks that are flushed on
cold or wet nights in the field, by cats, dogs, or other roving enemies, undoubtedly
perish before they can be assembled, for a baby quail in the natal down quickly gets
wet to the skin, and is chilled when running about in dew-laden grass... Severe
hailstorms undoubtedly cause a loss in the breeding season, especially in the rare
instances where the hailstones are very large, but we have no detailed data on their
effect on bobwhites...Very wet summers.. .are unfavorable to developing chicks in many
ways, all of which contribute to the dwindling of the broods. While the main damage is
undoubtedly caused by the destruction and desertion of nests, the young chicks have
their vitality lowered by wet and chilling; constant wetting may cause their death either
directly or indirectly by making them more susceptible to disease. The general belief
that wet summers are unfavorable to quail multiplication undoubtedly is only too well
founded. Drought, if extreme, may be equally detrimental. Not only do eggs spoil, but
the food supply in the shape of small fruits, insects, and succulent vegetation is much
reduced." Page 223 - "(2) The elements. Some loss to growing birds may be expected
annually from meteorologic conditions, which may reach serious proportions during
seasons either of abnormal rainfall or of intense drought. A season of normal rainfall or
one somewhat dried than average, but not dry enough seriously to affect the food
supply, is the most favorable to bobwhite increase, and fluctuations due to the elements
must be expected and allowed for in the shooting." Key words: bobwhite quail,
precipitation, drought, temperature, hail, nutrition, mortality, productivity

485

Stokkan, K.-A. 1992. Energetics and adaptations to cold in ptarmigan in winter.
Ornis Scandinavica 23(3):366-370. Summary: Ptarmigan Lagopus spp inhabit arctic,
subarctic and temperate alpine regions where winter is typically cold and snowy. Low
ambient temperatures, however, have little effect on their energy budget. Ptarmigan
have a highly insulative white winter plumage which contributes to a very low heat loss.
They normally evade exposure to severe cold by roosting in the snow, where ambient
temperature usually exceeds their lower critical temperature. Ptarmigan have a diurnal
activity pattern and each day collect and store enough food in their crop-sack to last the
long winter night. Their digestive system shows seasonal changes in length which
apparently adapt the birds to a low quality winter diet. With the exception of high-arctic

216 _^

4

^ fee >f*

c£^

species, ptarmigan do not store substantial amounts of fat in the winter. The Svalbard
ptarmigan living at 77-80°N prepare for winter by depositing large stores of fat (up to
35% of body mass; BM) in autumn. Concurrently there is a marked reduction in their
daily energy expenditure which permits such fattening to occur despite the fact that food
intake is simultaneously reduced. Key words: ptarmigan, snow, temperature,
behavior, thermoregulation

486

Straw, J. A. Jr., D. G. Krementz, M. W. Olinde, and G. F. Sepik. 1994. American
woodcock. Pages 97-114 in T. C. Tacha and C. E. Braun, editors. Migratory shore
and upland game bird management in North America. International Association of
Fish and Wildlife Agencies, Washington, D.C., USA. Notes: Migration chronology is
affected by wintering latitude and weather, especially strong cold fronts. Temperature
and moisture influence food availability and, thus, selection of wintering areas by
woodcock. During wet winters, woodcock winter more extensively in Texas than during
normal or dry winters. Most mortality during winter is attributed to predation, although
prolonged cold temperatures may result in localized mortality due to starvation. Key
words: woodcock, temperature, precipitation, behavior, nutrition, mortality

487

Stribling, H. L. and P. D. Doerr. 1985. Nocturnal use of fields by American
woodcock. Journal of Wildlife Management 49(2):485-491. Abstract: Nocturnal
behavior of the American woodcock (Scolopax minor) was studied during the winters of
1978-82 in agricultural crop fields in North Carolina. Woodcock preference for cutover
soybean fields could be attributed to at least two factors: earthworm (Oligochaeta)
quality and thermal advantages offered by soybean field microtopography. Over 75% of
the woodcock sampled in soybean fields fed during the night, and earthworms made up
99% of the food ingested (wet weight). Availability of earthworms (wet weight) among
cutover soybeans, disked corn, and winter wheat fields was similar (P > 0.05). The
percent protein of earthworms found in soybean fields was greater (P < 0.05) than in
other field types (corn and wheat). The parallel ridge/furrow microtopography of
soybean fields offered thermal advantages to woodcock. The plant debris that
accumulated between the furrows prevented the soil from freezing and allowed
woodcock access to earthworms on cold nights. Key words: American woodcock,
thermal cover, nutrition, habitat use, microtopography

488

Summers, R. W., R. E. Green, R. Proctor, D. Dugan, D. Lambie, R. Moncrieff, R.
Moss, and D. Baines. 2004. An experimental study of the effects of predation on
the breeding productivity of capercaillie and black grouse. Journal of Applied
Ecology 41:513-525. Summary: (1) The capercaillie Tetrao urogallus and black grouse
Tetrao tetrix are declining in the UK, and low breeding success has been identified as
the key factor in the decline of the former. To investigate possible causes, breeding
productivity was studied in relation to predation, weather, vegetation changes and deer
numbers over an 11 -year period (1989-99) within native pinewood at Abernethy Forest,
Scotland. The abundance of predators (crows Corvus corone and red foxes Vulpes

+ t^kr*^

217

e

<£C^ c£^>

vulpes) were experimentally manipulated in 1992-96 by culling. Productivity (chicks
reared per female) was compared between forests with and without experimental
predator management. (2) During predator control, the number of breeding crows was
reduced from 10 pairs to one. The attempted reduction in red fox abundance was
unsuccessful; only small numbers of adults were killed, and neither scat nor den counts
declined significantly. (3) Predation on artificial nests containing six hen eggs and a hen
egg filled with wax was measured as an index of predator activity from 1991 to 1999.
Predation was lowest during the last three years of predator control, 1994-96. Predators
could sometimes be distinguished by signs on depredated eggs. Predation on artificial
nests by crows was highest during 1991-93. However, after predator removal stopped
in 1997 few crows returned, and increased predation on artificial nests did not involve
increased sign of crow predation. Pine marten Maries martes numbers increased during
the study period and became significant predators of artificial nests. (4)The total number
of capercaillie eggs and nests depredated by crows was estimated from the number of
depredated capercaillie eggs found and the proportion of crow-predated hen eggs in
artificial nests. The values ranged from 18 to 158 eggs over 3 years, equivalent to 3-23
capercaillie nests year"1. (5) Capercaillie productivity was low (<1 chick per female)
during 1989-93 and 1997-99 but higher during 1994-96. Compared with nine other
forests in Scotland, changes in capercaillie productivity at Abernethy were different.
Productivity at Abernethy was negatively related to June rainfall, and to the minimum
daily predation rate on artificial nests by crows. There was also a significant interaction
in that capercaillie were most productive when low rainfall coincided with low predation
by crows on artificial nests. (6) The productivities of black grouse and capercaillie were
positively correlated, but greater in the former. As in capercaillie, black grouse
productivity was negatively related both to June rainfall and the minimum daily
predation rate on artificial nests by crows, and there was an interaction. (7) Synthesis
and applications. The long-term increase in crows and red foxes and the predicted
increase in rainfall in Scotland may have negative effects on capercaillie and black
grouse. In the short term, control of crows is likely to improve productivity. In the long
term, increased woodland size and some reversal of fragmentation might decrease the
access to woodland of predators associated with the interface between farmland and
woodland Keywords: capercaillie, black grouse, predation, precipitation

489

Swank, W. G. and S. Gallizioli. 1954. The influence of hunting and of rainfall upon
Gambel's quail populations. Transactions of the North American Wildlife
Conference 19:283-297. Notes: Page 287— "In the desert country, the range of the bird
in question, production of green vegetation in the spring is primarily dependent upon the
quantity of rainfall received during the preceding winter months. A ratio of more than
two young to one adult occurred only during those years when the accumulated rainfall
from December through April amounted to more than the average of 5.58 inches." Page
294 — "Information on fluctuations of the Gambel's quail population in Arizona over the
past 13 years indicates that the amount of rainfall during winter months, December
through April, is the factor limiting abundance." Key words: Gambel's quail,
precipitation, productivity

218 ^

4*

>->" (& ^

490

Swanson, D. L. and D. P. Weinacht. 1997. Seasonal effects of metabolism and
thermoregulation in northern bobwhite. Condor 99(2):478-489. Abstract: Seasonal
differences in metabolism and cold hardiness are common among small passerine
birds. However, seasonal adjustments of metabolism and insulation are less well
studied in nonpasserines and in larger birds. We measured basal metabolic rate (BMR),
metabolic response to temperature, and maximal capacity for thermogenesis (peak
cold-induced oxygen consumption, V02sum) in late spring/summer and winter in outdoor
captive northern bobwhite (Colinus virginianus) near the northern boundary of their
natural range, to determine whether seasonal adjustments in metabolism are a
component of acclimatization in this species. Mass, BMR and regression equations
describing diurnal and nocturnal metabolic response to temperature were not
significantly different between seasons. After metabolic tests below thermoneutrality,
cloacal temperature (Tb) was not dependent on ambient temperature (Ta) at either
season, and nocturnal Tb did not differ significantly between seasons. However, after
metabolic tests below thermoneutrality, diurnal Tb was significantly greater in summer
(38. 9± 1 . 1°C) than in winter (37.7 ±1.1 °C). Although body mass of winter birds was
significantly greater than their body mass in late spring, maximal thermogenic capacity
did not differ significantly on a seasonal basis, and winter bobwhite were only marginally
more cold tolerant than late spring birds under severe cold stress. For individual birds
tested in both winter and late spring, individual ranging of V02sum was not consistent
between seasons (i.e., birds with a high V02sum in winter did not necessarily have a high
V02sum in late spring). These data suggest little seasonal adjustment of metabolism or
insulation in the northern bobwhite, a relatively large nonpasserine species native
mainly to regions with relatively mild winter climates. Key words: bobwhite quail,
thermoregulation, temperature

491

Swearingin, R. M. 2007. Winter roosting ecology of Rio Grande wild turkeys in the
Rolling Plains of Texas. M.S. Thesis, Texas Tech University, Lubbock, USA.

Notes: We examined possible relationships between roost/flock counts and climatic
variables. We predicted that daily low temperature and length of photoperiod would
explain a significant amount of the variation in foraging and roosting flock sizes
throughout the year. Wild turkey attendance at traditional winter roost sites was not
correlated with daily precipitation (/=0.139, n = 101, P=0.164), daily snowfall (^0.139,
n=101, P=0.164), length of photoperiod (n= -0.091, n=101, P=0.368), or daily maximum
temperature (a=0.177, d=101, P=0.077). We did find a significant correlation between
roost size and minimum temperature (n= -0.245, n=1 1 , P=0.014). We found no
relationship between wild turkey flock sizes and daily precipitation (r=0.008, n=3047,
P=0.640) or daily snowfall (^0.028, n=3047, P=0.128). However, we did find weak
negative correlations between wild turkey flock size and maximum daily temperature (r=
-0.290, A7=3047, P=0.000), minimum daily temperature (r= -0.323, n = 3047, P=0.000),
and length of photoperiod (r= -0.367, n=3047, P=0.000). We suggest that daily
precipitation and daily snowfall may have had little effect on wild turkey flocking
behavior. Most (70%) rainfall on the study sites occurred between April and August,
there were few rain and snowfall events during our period of interest. We found that

% f>'^t*

219

e

o^ ^§^ £Z^> c£j>

minimum temperature influences flock size and that minimum temperature, maximum
temperature, and length of photoperiod influence roost size. Although these correlations
were significant, these variables explained little of the variation in flock and roost size.
Even though we observed low correlation coefficients, we believe that these variables
may have a greater influence on winter turkey concentrations in combination. There
may also be an effect of time lag influencing winter concentrations. Since cold
temperatures impact wild turkey concentrations, roosts should be counted during the
coldest portion of the specified peak period for the greatest likelihood of the highest
densities of wild turkeys using major roosts. Key words: turkey, snow, precipitation,
temperature, behavior, census

492

Swenson, J. E. 1991 . Evaluation of a density index for territorial male hazel
grouse Bonasa bonasia in spring and autumn. Ornis Fennica 68(2):57-65.

Abstract: A density index for territorial male hazel grouse Bonasa bonasia in spring and
autumn is presented and evaluated. One whistles with a hunter's whistle every 30
seconds for 6 minutes from census points located at 150-m intervals within the area to
be censused, and responding hazel grouse are counted. Counts were conducted
throughout days with little or no wind. The number of counted males was significantly
and linearly correlated with the number of known territorial males on an intensive study
area. There, 82% of the territorial males responded and were counted. Response rate
appeared to be independent of density, based on counts in areas with a 20-fold
variation in densities. When censuses were repeated, both results were similar. Within
the conditions of these counts, no effects of weather or date were found. However,
hazel grouse responded less at midday. Using this method, one can count hazel grouse
at a rate of about 5 minutes per ha along transects and 5-9 minutes per ha on blocks of
habitat, depending on plot size. I recommend that censuses be conducted during 4-5
weeks prior to laying in spring and during 4-5 weeks after brood dissolution in autumn.
Key words: hazel grouse, index, technique, wind, temperature, census

493

.1986. Differential survival by sex in juvenile sage grouse and gray

partridge. Ornis Scandinavica 17(1):1417. Summary: Studies of capercaillie Tetrao
urogallus and black grouse T. tetrix in northern Europe have found that juvenile males
of these sexually size dimorphic species suffer higher morality than juvenile females
during adverse conditions. This difference may be due to the more rapid growth rates
among males. Differences in the juvenile survival of the dimorphic sage-grouse
Centrocercus urophasianus and the monomorphic gray partridge Perdix perdix were
studied in western North America. Juvenile male sage grouse survive less well than
juvenile females during years unfavorable for juvenile survival and in poorer habitats.
Juvenile male gray partridge showed little or no such trend. These results are consistent
with those obtained in the European studies. Pressures on sexual selection may have
led to a growth rate in juvenile males of highly dimorphic grouse species which is near
the upper limit of that which can be sustained by their ecological niche. Key words:
sage-grouse, gray partridge, mortality, genetics, habitat quality

220 ^

^

>> <0 **

cCb

494

. 1985. Seasonal habitat use by sharp-tailed grouse, Tympanuchus

phasianellus, on mixed-grass prairie in Montana. Canadian Field-Naturalist 99:40-

46. Abstract: Upland grass and upland crops were first and second, respectively, in
observed use by sharp-tailed grouse (Tympanuchus phasianellus) during all seasons.
Hardwood draws were used for foraging for fruits and berries in autumn and for buds in
winter. Winter use of habitats varied with snow depth; during periods of deep snow, the
grouse used hardwood draws more and upland grass and crops less. Upland grass was
selected for placement of arenas. A mosaic of upland grass with Skunkbush sumac
(Rhus trilobata) and riparian hardwood draws seemed to comprise optimum habitat; but
small upland winter wheat fields interspersed throughout the area and within 500 m of
hardwood draws provided an apparently preferred food source. Key words: sharp-
tailed grouse, snow, habitat use, behavior

495

and B. Olsson. 1990. Hazel grouse night roost site preferences when snow-
roosting is not possible in winter. Ornis Scandinavica 22(3):284-286. Abstract
Hazel grouse Bonasa bonasia typically roost in snow burrows at night in winter, but this
is often not possible in maritime climates. The alternative, roosting in trees, was studied
in south-central Sweden during three winters. Data from 64 roost sites showed that
hazel grouse preferred to roost in Norway spruce Picea abies, the tree species on the
area providing the greatest amount of vertical cover. Roost trees were consistently
shorter than the surrounding canopy trees and birds roosted low in the trees. Roosting
sites had a significantly higher density than expected of both spruces and total trees.
Thus, hazel grouse roost sites had more vertical and horizontal cover than expected.
This type of site probably provided thermal benefits to roosting grouse. Key words:
hazel grouse, behavior, thermoregulation, temperature, wind, snow, habitat use

496

, L. Saari, and Z. Bonczar. 1994. Effects of weather on hazel grouse

reproduction: an allometric perspective. Journal of Avian Biology 25(1):8-14.

Summary: We analyzed the effects of weather during prelaying, incubation, and early
chick periods on the reproductive success of hazel grouse Bonasa bonasia in
southwestern Finland (14 years) and southern Poland (6 years). Reproductive success
was most clearly correlated with weather during the prelaying period in both areas, with
days of precipitation correlating negatively and temperature positively. We suggest that
the availability of nutritionally rich food and the ability of the females to obtain it during
the prelaying period, when the females are rapidly gaining mass and forming eggs,
determine in large part the reproductive success of hazel grouse. A literature review
suggested that this is generally true for small grouse species. The larger species, which
have greater endogenous reserves and invest relatively less in their clutches, are less
affected by prelaying weather conditions. Chicks of larger species have higher growth
rates and energy requirements, however, which is probably the reason why the
reproductive success of the larger species is more affected by weather during the early
chick period. Key words: hazel grouse, precipitation, temperature, productivity

y 221

n^

>X fc* ^

OSr -"car- ^0^ cO±

497

Szaro, R. C. and R. P. Balda. 1986. Relationships among weather, habitat
structure, and ponderosa pine forest birds. Journal of Wildlife Management
50(2):253-260. Abstract: Avian community structure during the breeding season in a
ponderosa pine (Pinus ponderosa) forest of northern Arizona was influenced by
weather and a series of timber harvest treatments. Fewer birds and bird species were
present after a winter with the heaviest snowfall on record and low temperatures than
after milder winters. Bird density was greater (P<0.05) on the light and medium cut plots
than on the untreated plot. A cluster analysis of bird densities over plots and the 3-year
study period indicated treatment effects were more important in determining bird
community composition than weather effects. Key words: mourning dove,
temperature, precipitation, snow, habitat use

498

Tacha, T. C. and C. E. Braun, editors. 1994. Migratory shore and upland game bird
management in North America. International Association of Fish and Wildlife
Agencies and Fish and Wildlife Service, Lawrence, Kansas, USA. Abstract: White-
tipped doves (Leptotila verreauxi) are large tropical columbids native to southern Texas,
Mexico, Central America, and portions of South America. They have been a legally
hunted game species in the United States since 1984. Their range in the U.S. is a 14-
county area of southern Texas. White-tipped doves are sedentary and do not migrate.
In southern Texas they nest in native brush and citrus orchards. The annual breeding
population has been stable in Texas for the past 10 years, except for declines during
years following major freezes that adversely affect citrus and native brush nesting
habitat. A limited harvest of 2 white-tipped doves per day is allowed during the special
white-winged (Zenaida asiatica) and mourning dove (Z. macroura) dove hunting
seasons. Improvements in harvest surveys are needed. Other management and
research needs include more detailed breeding population surveys in the northern part
of their range, and additional life history data. Chapter 7 — American Woodcock: Notes -
Page 100 - "Weather factors such as temperature and moisture influence food
availability and, thus selection of wintering areas by woodcock. The abundance and
distribution of woodcock in Louisiana... varies with winter severity. Furthermore, during
wet winters, woodcock winter more extensively in Texas than during normal or dry
winters. Most mortality during winter is attributed to predation, although prolonged cold
temperatures may result in localized mortality due to starvation." Key words: white-
tipped dove, American woodcock, temperature, ice, moisture, nutrition,
distribution, mortality, population dynamics

499

Tautin, J., P. H. Geissler, R. E. Munro, and R. S. Pospahala. 1983. Monitoring the
population status of American woodcock. Transactions of the North American
Wildlife and Natural Resources Conference 48:376-388. Notes: Page 381 --
"(Calling) counts should not be made during heavy rain, snow, high wind, or
temperatures below 40°F (4°C). Normally, weather has little effect on courtship
performances, but the conditions listed above result in curtailed activity." Page 386 --
"Does the Singing-Ground Survey provide information useful or management

222 ^

4

^ t& >f*

purposes? We believe it does and that our proposed new analytical methods will
enhance its capabilities. Although the studies of the 1970s did not conclusively answer
the question, certain empirical information suggests that the answer is affirmative. The
blizzard that struck the northeast in early April 1982 was predicted to impact adversely
on woodcock that had already returned to their breeding grounds. Subsequent results
from the singing ground survey showed a decrease of 20.3 percent from 1 981 ,
suggesting that the survey is sensitive to between year population changes." Key
words: American woodcock, census, precipitation, snow, wind, temperature,
behavior, index

500

Tavecchia, G., R. Pradel, F. Grossmann, C. Bastat, Y. Ferrand, and J.-D. Lebreton.
2002. Temporal variation in annual survival probability of the European woodcock
Scolopax rusticola wintering in France. Wildlife Biology 8(1):39-48. Abstract: The
Eurasian woodcock Scolopax rusticola is an important quarry species hunted all over its
range. Some authors have reported local declines in both wintering and breeding
woodcock numbers. In order to investigate whether these possible declines are the
result of a negative trend in survival, we analyzed 3,312 recoveries of 15,839
woodcocks ringed in France during 14 consecutive winters (1984/85-1997-98). We
distinguished between winter (October-February) and summer (March-September)
recoveries in order to estimate survival and recovery rate separately for each period
because selective pressures during these two periods are likely to be different. Survival
varied according to year during both winter and summer. Winter survival probability
covaried positively with mean winter nocturnal temperature and ranges from 0.74 (SE =
0.057) during the winter of 1985/86 to 0.83 (SE = 0.042) during the winter of 1994/95.
Mortality of first-year birds was 22% higher than that of adults in any year. Results from
a second analysis in which we compared survival of birds ringed during 1991-1997 in
the three main woodcock wintering areas along the French Atlantic coast suggested a
threshold effect of weather conditions. Mean winter survival covaried with temperature
and rainfall mainly in the northernmost regions where weather conditions are more
severe. We did not find any particular trend in survival probability that could explain the
possible declines in woodcock numbers. However, the generally low adult annual
survival, and the negative influence of stochastic events such as severe winter
conditions might drive populations to a level from which it would be difficult to recover.
Results of a two-age-class demographic model are discussed together with implications
for management. Key words: Eurasian woodcock, temperature, precipitation,
population dynamics, model

501

Taylor, J. S., K. E. Church, and D. H. Rusch. 2000. Habitat and weather effects on
northern bobwhite brood movements. Proceedings of the National Quail
Symposium 4:153-157. Abstract: We observed radio-marked northern bobwhite
(Colinus virginianus) broods (adults with chicks < 1 days old; n = 12) in Kansas during
1991-94 to test effects of weather (temperature and precipitation) and macrohabitat
(composition, relative diversity, and mean distance to grassland) variables on brood
home range size and daily movements at large (28.5 km2), intermediate (3.14 km2), and

-* 223

i FV'^t*

% #> <Q> 9>

small (about 0.14 km2) spatial scales surrounding habitats available for broods.
Principal component analyses followed by stepwise multiple linear regression indicated
neither weather nor habitat influenced (P> 0.1) home range size at the large and
intermediate scales. However, the principal component representing mean distance to
grassland and percent cropland within the home range (i.e., at a small scale) was
positively related to home range size. Neither temperature nor habitat influenced daily
distance of movements. We concluded that brood mobility was independent of
landscape-scale features, but that habitat management at smaller spatial scales could
influence movements. To create optimal habitat for bobwhite, managers should
consider relationships among habitat attributes and the movement of individuals,
including the spatial scales at which these relationships are most important. Key
words: bobwhite quail, temperature, precipitation, behavior, movement, habitat
manipulation

502

Theberge, J. B. and G. C. West. 1973. Significance of brooding to the energy
demands of Alaskan rock ptarmigan chicks. Arctic:138-148. Abstract: Rock
ptarmigan (Lagopus mutus) chicks are brooded periodically during the first few days of
life; longer in cold and rainy weather. Computed minimum foraging time in adverse
weather conditions is 96 minutes/24 hours. Crop analysis and calorimetry of the 6 major
food items show that a full crop may contain up to 0.47 kcals. Energy requirements
were calculated for both an 18-gram chick and a 30-gram chick. The 18-gram chick
required between 34 and 50 crop loads per 24 hours. With 96 minutes foraging time,
and the observed pecking rates, this was considered possible. The 30-gram chick
required twice as much foraging time but since it was approaching homeothermy, it was
tentatively concluded that neither was that chick being handicapped by brooding.
Vagaries in early survival of rock ptarmigan chicks, therefore, are not due to difference
in post-hatch weather. Key words: rock ptarmigan, temperature, precipitation,
nutrition, thermoregulation

503

Thomas, V. G. 1987. Similar winter energy strategies of grouse, hares and rabbits
in northern biomes. Oikos 50(2):206-212. Abstract: Neutral fat and protein reserves
were measured in three species of grouse, cottontail rabbits, snowshoe hares and
European hares collected during the mid to late winter period in their natural habitats.
Neutral fat levels were measured in arctic hares from the Canadian high arctic during
late summer. Sharp-tailed grouse, spruce grouse and ruffed grouse contained,
respectively, 2.2, 2.3 and 1.0 g of fat per 100 g fat-free body weight, regardless of sex,
year or location. Cottontail rabbits, arctic hares, snowshoe hares and European hares
had, respectively, 4.1, 2.5, 2.3-4.1 and 6.9 g of fat per 100 g fat-free body weight. The
low values for fat and protein reserves are similar to those reported for other grouse and
leporid species in different parts of North America and Scandinavia. Collectively they
indicate that species in both taxa have little metabolic resistance to winter fasting and
must feed regularly to maintain energy balance. Grouse, hares and rabbits in winter
habitats share similar energetic tactics of acquiring small energy reserves, feeding
regularly for short periods and emphasize energy conservation rather than energy

224 ^

^

» te ^

c£^

acquisition. A theoretical model relates the size of energy reserves in these species to
the probability and duration of fasts due to severe winter weather, the changing quality
of diets, and the probability of predation while amassing reserves prior to winter and
using them during winter. Key words: sharp-tailed grouse, spruce grouse, ruffed
grouse, bioenergetics, severe weather

504

Thompson, D. J. 2003. Roosting habitat and poult survival of Merriam's turkeys in
the southern Black Hills of South Dakota. M.S.Thesis, South Dakota State
University, Brookings, USA. Abstract: Poult survival rates were insignificantly higher
during 2002 (P>0.10). The increased invertebrate abundance during 2002, coupled with
cold wet weather during poult development in 2001 , accounted for the slight variation
between poult survival between years in the southern Black Hills. Key words:
Merriam's turkey, temperature, precipitation, mortality, nutrition, chick survival

505

Thompson, D. R. 1950. Foot-freezing and arrestment of post-juvenal wing molt in
the mourning dove. Wilson Bulletin 62(4):212-213. Notes: The author notes that
mourning doves were captured that had badly frozen feet in the winter of 1949-50. The
frozen distal phalanges dropped off; had these birds needed to forage for their own food
during their convalescence they would probably have died of starvation. Arrestment of
the post-juvenal molt was apparent in all three of the birds held captive following foot
freezing. The arrest of molt varied between the birds. These birds were described as
"short, ragged, and faded dull brown, lacking entirely the sheen of the pearly gray new
feathers." If birds of late-hatched broods do not molt the outer primaries before the
arrival of cold weather the molt may be arrested or suspended. Key words: mourning
dove, temperature, frost, molt

506

Thompson, K. M., M. J. Holloran, S. J. Slater, J. L. Kuipers, and S. H. Anderson.
2006. Early brood-rearing habitat use and productivity of greater sage-grouse in
Wyoming. Western North American Naturalist 66(3):332-342. Notes: Page 339 -
"We found virtually no relationship between weather and productivity. It is possible that
short-term, extreme weather conditions (e.g. heavy rainfall, severe cold spells)
influenced productivity; but these occurrences were not detectable using annual
weather data. However, the trends we did observe were consistent; all weather
variables were positively associated with our two measures of productivity. Warm and
dry conditions appeared to be more favorable for productivity than cold and wet
conditions." Keywords: sage-grouse, precipitation, temperature, productivity

507

Thompson, F. R. Ill and E. K. Fritzell. 1988. Ruffed grouse metabolic rate and
temperature cycles. Journal of Wildlife Management 52(3):450-453. Abstract: We
measured the effects of temperature and wind on ruffed grouse {Bonasa umbellus)
metabolic rates in an open-circuit respiration system with a closed-circuit wind tunnel.

% f>"^^c

225

Metabolic heat production was linearly related to operative temperature (Te) below a
lower critical temperature of 1 .5 ± 0.99 (SE) C. Standard metabolic rate was 3.2 ± 0.1 1
W (66.1 kcal/day) for a grouse of mean weight (607 g). Metabolic rate was related
linearly to wind speed0 5. The regression predicting net metabolic heat production
(metabolic heat production - evaporative cooling [M - E]) was 3.1589 - 0.1 176Te +
0.4141 (wind speed)0 V = 0.98, P<0.001 ), where Te = 0 to -20C and wind speed < 3
m/second. Mean high daytime cloacal temperature (40.6 ± 0.24 C) was > mean low
nocturnal cloacal temperature (38.3 ± 0.54 C) (P - 0.006). Key words: ruffed grouse,
temperature, wind, metabolic rate

508

and . 1988b. Ruffed grouse winter roost site preference and

influence on energy demands. Journal of Wildlife Management 52(3):454-460.

Abstract: We located winter roosts of ruffed grouse (Bonasa umbellus) in Missouri with
radiotelemetry and determined roost type preference. Thermostatic energy demands in
4 roost types were measured with a heated taxidermic mount calibrated with metabolic
rates of captive grouse. Ruffed grouse preferred to roost in the canopies of eastern red
cedar {Juniperus virginiana) and avoided roosting in deciduous cover. Roost sites had
higher woody stem densities (mean = 5,494 stems/ha) than random plots (mean =
4,236 stems/ha). We predicted ruffed grouse metabolic rates (±0.75%), at ambient
temperatures (Ta) of -20 to 0 C, from power (Pm) used by the heated taxidermic mount.
Standard operative temperature (Tes) was elevated 7.3, 2.6, 2.9, and -0.3 C in snow
roosts, cedar tree roosts, cedar ground roosts, and deciduous roosts, respectively,
above that at an open site for Ta of -20 to 0 C. When wind speed was 3 m/second in the
open, Tes was elevated a mean of 12.9, 7.0, 6.7, and 2.5 C, in snow, cedar tree, cedar
ground, and deciduous roosts, respectively. These increases in Tes resulted in a 33, 19,
18, and 6% reduction in metabolic rate in snow roosts, cedar tree roosts, cedar ground
roosts, and deciduous roosts, respectively, from that in the open at -20 to 0 C and 3
m/second wind speed. About 40% of the elevation in Tes resulted from reduced
convection inside roost sites and 60% from a more favorable radiation balance in
roosts. Low coniferous vegetation provided thermal benefits that may be important
because snow roosts are rarely available. Key words: ruffed grouse,
thermoregulation, behavior, wind, snow, temperature, habitat use, solar radiation,
vegetation

509

Thompson, W. L. 1993. Ecology of Merriam's turkeys in relation to burned and
logged areas in southeastern Montana. Dissertation, Montana State University,
Bozeman, USA. Abstract: Merriam's turkeys (Meleagris gallopavo merriami) were
captured, equipped with radio transmitters, and monitored via telemetry to study their
ecology in relation to burned (1988 wildfire) and logged areas in southeastern Montana,
1989-1991. Neither burned nor logged areas appeared to restrict seasonal movements
of monitored turkeys during spring and fall. However, males and subadult hens in the
burned study division moved farther than those in the unbumed during spring/fall. Hens
tended to nest more in burned areas with each subsequent year following the fire. Nest
success was highest during the spring with largest amounts of precipitation. High levels

226

4>

» te ^

c£Tb

of precipitation during spring likely promoted understory plant growth and increased
nesting cover especially where fire opened the canopy to more sunlight and increased
nutrient release to soils. Only 12 of 49 nests occurred in logged areas. Clearcut logging
may be more detrimental to nesting habitat in xeric (for example, southeastern
Montana) than mesic regions because of slower vegetational growth and lower habitat
structural diversity. Nests generally occurred in cover >6. 5 dm high and on >20%
slopes. Summer home range sizes were not correlated with percent burned or logged
areas that they contained. Roost trees within burned areas were larger in diameter than
those in unbumed areas. More "large" trees in severely burned areas may have
become attractive to roosting turkeys through loss of dense canopy and simplified
branch structure. Turkeys usually roosted in the taller (mean = 18m) and larger
diameter (mean = 41cm dbh) trees available. On average, roosts occurred on 25%
slopes with easterly or northeasterly aspects. Severely burned trees used for roosting
may be lost over time because of their breakage and falling. No roosts occurred in
forest stands subjected to any type of timber harvesting scheme. Survival rates of
monitored turkeys were similar between burned and unbumed areas. Overall, habitat
quality for turkeys likely was lowered during the first year or 2 following the fire, but
should generally increase (except roosting habitat) in subsequent years. The effects of
logging on turkey habitat in southeastern Montana remain unclear; however, logged
areas were rarely used by monitored turkeys during this study. Key words: Merriam's
turkey, precipitation, fire, habitat use, vegetation

510

Toepfer, J. E. 1988. The ecology of the greater prairie chicken as related to
reintroductions. Dissertation, Montana State University, Bozeman, USA. Notes:
Page 34 — "In the fall, cocks displayed only during the morning on clear, calm mornings.
During stormy or very cold days in winter the birds fed only in the morning... wild prairie
chickens typically feed twice a day except during cold winter weather when they fed
only once.: Page 49/50 — "Adult cocks returned to the booming grounds in mid-
September to re-establish their territories about the time of the first hard frost... [a] cock
was observed displaying on a regular basis even on snow..." Page 71 — "The decline [in
weight] in winter was due to lower ambient temperatures and snow cover and reduced
availability of food and increased energy demands. Reductions in food and cover
caused daily movements and ranges to expand which thus increased energy demands
caused by lower temperatures." Page 73 — "Immatures and adult hens appeared to be
more sensitive to the rigors of winter than adult cocks." Page 73/76 — "Reasons for
increased weight loss observed in the winter of 1976 are unknown. Weather differences
between the years may have a greater effect on hens and immature cocks since they
are generally smaller and more mobile than adult cocks." Page 78 — "The quantity and
distribution of winter food and cover arte critical to survival as they directly influence
daily movements and hence weight loss. Weight gains, during fall and on early winter
peak, imply that food and cover in the fall are also very important." Page 79 — "The rapid
increase in mean weights observed in prairie chickens during the early spring
correspond to warming temperatures and the loss of snow cover, which increased
distribution, quantity and quality of food and cover." Page 80 — "After snow melted
radioed prairie chickens... were also attracted to the green plants which started to grow
early in cultivated fields because of warmer soil conditions created by the disturbed

^ 227

^

rY <o ^

<QS~ -"taC' t^ZSS? r7"^

soil." Page 82 — "A relationship seemed to exist between the occurrence of green-up
and onset of breeding since both were usually associated with warm springs." Page
175 — "Winter was the period when wild prairie chickens were most vulnerable and
suffered highest mortality." Page 201 — "Seasonal variation in flushing distances was
believed to be related to security provided by vegetative cover. Mean flushing distances
were greatest in winter when cover was limited by snow and shortest during the
summer when vegetation provided the greatest security." Page 218 — "Snow cover
restricted the availability of food causing both cocks and hens to form larger flocks."
Page 225 — "The number of movements made per day by radio-tagged prairie chickens
varied with and within the seasons and was influenced most by the distribution of food
and cover, breeding activities, sex, age, and weather conditions." Page 232 — "The large
home ranges in winter were related to snow cover, which reduced the availability of
food and cover causing individuals to increase their home ranges to meet their daily
needs. Page 233 — "Home range sizes increased in fall as temperatures declined and
birds began feeding in the agricultural fields in the mornings and evenings." Page 235 —
"Seasonal home ranges of both cocks and hens and immatures and adults were most
comparable during summer and winter in the absence of breeding behavior. During
these periods habitat conditions appeared to have the greatest influence on the size of
a bird's home range." Page 237 — "The fluctuations in weekly [home] ranges during
winter were due to snow cover which changed food and cover conditions." Page 250 —
"Resident radio-tagged prairie chickens made movements that covered relatively large
areas during periods when they were adjusting to new or changing conditions. These
extended irregular movements in residents occurred... in winter when new snow
covered a food source and birds made irregular movements to locate a new
one. . ."Page 269 — "Mean DD movements remained high and fluctuated throughout the
winter as birds adjusted their movement patterns whenever fresh snow altered food and
cover conditions. The magnitude and patterns of daily movements during winter were
comparable between the sexes and ages and reflected the strong influence that habitat
distribution had on movements." Page 274 — "...twice daily feeding pattern stopped in
winter during periods of sub-zero weather. During periods of cold weather radio-tagged
birds and their flock members remained in roosts longer in the A.M., fed in agricultural
fields later or during midday, flew to roosts earlier than normal (as early as 1400 hrs),
and remained there until the following day." Page 275 — "Visual observations were
obtained of radio-tagged birds and flock members in snow burrows several hours before
they normally would have gone to roost. Reduced activity during cold weather is
believed to be an energy conservation mechanism. Several days of very cold weather
are necessary for prairie chickens to initiate this behavior... radio-tagged prairie
chickens have been known to remain in snow burrows after a heavy snow storm for 36
hours." Page 283 — "The first permanent snow cover greater than 6 cm caused the
cocks to abandon their booming grounds and center their activities around a food
source... in fall, adult hens moved away from the booming grounds with the first heavy
frosts as they began to feed in agricultural areas." Page 334 — "Transplanting birds
during winter because they are easy to catch will only compound the high natural
mortality that resident prairie chickens experience at this time." Page 342 — "The
variation in plant species identified in seasonal cover reflected their structural form and
their ability to provide cover at different times of the year, particularly the ability to
withstand the weight of snow." Page 247 — "Winter was the most difficult season for

228 j^

» k> **

prairie chickens because snow reduced the amount and availability of food and cover.
Each new snowfall changed conditions and altered the distribution of food and cover
which caused prairie chickens to increase their movements and energy demands.
Habitat types which provided cover and food above the snow were utilized most by
birds... The snowfence effect of... willows caused an accumulation of snow on the lee
side that was used for snow burrowing. This accumulation of snow also kept some of
the leeward vegetation from being covered by snow." Page 350 — "Some habitat types
used at other times of the year were not used during winter because of snow cover.
Upland grass was used during periods without snow, but used very little during winter."
Page 351 — "The only true shrub habitat used by the prairie chickens was Spirea in
central Wisconsin which made up only 0.8% of day use and 8.3% of night use.
However, shrub habitat may play an important role in providing cover when snow
becomes crusted or unusually deep reducing the amount of wetland vegetation and
grass-forb." Page 357 — "In this study birds... made no movements to shrubs or trees for
shade during hot weather. Instead of woody cover the radio-tagged prairie
chickens... used taller clumps of grass and grass-forbs for shade." Page 360 — "Frost
and changing temperatures during fall caused the quality of upland grass to deteriorate
as night cover. Changing weather conditions also caused cover to open up and allow
access to some cover that was too dense to use in summer." Page 367 — "Habitat use
trends at night were influenced by snow and cold temperatures." Page 369 —
"Vegetation that projected above the snow was critical in providing prairie chickens with
screening cover and protection from the snow and wind during winter.... Classes I and II
cover were used by prairie chickens when loafing in snow depressions or when the
snow was deep enough to permit snow burrowing." Page 373 — "During winter, snow
altered the [vegetation] structure and permitted access to much of the taller
vegetation... even during summer when temperatures were warm, radio-tagged prairie
chickens shifted from shorter more open cover to faller.... vegetation which made up
79.5% of the night roosting cover." Page 376 — "Use of vegetation by height classes
also varied within the seasons and was related to the development of vegetation and
weather conditions. Page 381 — "Much of the grass and sedge cover in the Class I V+
vegetation category was reduced to Class III or II category by weathering in fall and
snow cover in winter." Page 467 — "In Missouri, where the winters are milder and snow
of lesser amount and duration, the bulk of the prairie chickens' diet consisted of native
plants. Cultivated grains were used most when snow covered native vegetation. Deep
persistent snow cover and the inability to subsist entirely on browse may have been the
barrier that originally isolated the prairie chicken from the sharptail." Key words: prairie
chicken, snow, temperature, wind, frost, solar radiation, vegetation, habitat use,
distribution, movement, soil, habitat quality, mortality, behavior, transplanting,
plant phenology, plant structure

511

Trautman, M. B., W. E. Bills, and E. L. Wickliff. 1939. Winter losses from starvation
and exposure of waterfowl and upland game birds in Ohio and other northern
states. Wilson Bulletin 51(2):86-104. Conclusions: Winter mortality of waterfowl and
some species of upland game birds, caused by starvation, exposure to low
temperatures, or a combination of both, occurs in widely scattered localities in the
northern United States. Such mortality generally takes place during or immediately

\ frkr***

229

igy ?®Ti <££> ^

following severe sleet or snow storms when low temperatures freeze the surface water
shutting off the usual food supply. During the average stress period apparently only
those birds die which are weakened by disease, parasitism, lead poisoning, mechanical
injury such as are made by gunshot wounds, or ice formation over eyes, in bill and
nostrils. In extreme stress periods, and especially when the accumulated, adverse
effects of winter are most pronounced, mortality of normal birds may occur from
starvation and exposure. Key words: ruffed grouse, ring-necked pheasant,
bobwhite quail, snow, temperature, ice, mortality, nutrition

512

Tsuji, L. J. S. 1992. Snowfall causes lek movement in the sharp-tailed grouse.
Wilson Bulletin 104(1):188-189. Summary: Observations made on two leks suggest
that stability of territories on leks can be affected by unusual environmental
circumstances. In one case, flooding caused the lek to move 30.5 m from its location
noted the previous day; in the other, a snowstorm covered existing landmarks on the
muskeg with the lek forming and reforming in three different locations in the same
morning. Although the location of the lek changed in these two cases, the actual
infrastructure of the lek appeared unchanged. In other words, individuals in the lek
maintained their positions relative to one another. Key words: sharp-tailed grouse,
snow, flood, habitat use, behavior

513

Uhlig, H. G. and R. W. Bailey. 1952. Factors influencing the distribution and
abundance of the wild turkey in West Virginia. Journal of Wildlife Management
176(1):24-32. Notes: Under "Summary" the authors state — "The turkey kill and mast
conditions correlate with minimum temperatures occurring during May. The greater the
extent of below normal temperatures for that month, the poorer the crop of mast and the
lower the reported turkey kill of the same year." Key words: Eastern wild turkey,
temperature, vegetation, hunting

514

Vander Haegen, W. M. 1992. Bioenergetics of American woodcock during the
breeding season on Moosehorn National Wildlife Refuge, Maine. Dissertation,
University of Maine, Orono, USA. Abstract: Bioenergetics of female American
woodcock (Scolopax minor) was studied from 1987-1989 at Moosehorn National
Wildlife Refuge, Maine. A model of daily energy expenditure was developed from
laboratory-derived data on metabolic rates; from data on activity and microclimates
collected in the field; and from body component analysis of collected birds. Energy
demands incurred by female woodcock on the breeding grounds were highest during
the pre-nesting (60.3 kcal/day) and laying (89.1 kcal/day) periods. Availability of food
(earthworms - Lumbricidae) is normally sufficient during these periods, but shortages
such as the one caused by persistent soil frost in spring of 1989 can delay nesting and
affect productivity. Female woodcock feed throughout the diel period prior to incubation,
obtaining nutrients for reproductive tissues and to store fat for use during incubation.
Incubating females spend only 8% of the day active and used endogenous reserves to
supplement energy derived by feeding, losing about 75% of their body fat over

230 ^

> > fa. ^

c£^>

incubation. In March and April 1989, energy intake was too low to initiate egg
production as nesting did not occur until the frost melted and earthworm availability
returned to normal, 3-4 weeks later than the typical nesting date in Maine. Woodcock
chicks are not homeothermic until 15-20 days old, and there is an inverse relationship
between air temperature and brooding requirements. At air temperatures typical of the
brood period, a drop in mean air temperature of (illegible) can result in a 40% decrease
in time spent active, with a concomitant loss of foraging time. Rainfall also increases the
brooding requirements of chicks, reducing by 30% the time spent activity by chicks
(illegible) 10 days old. Reduced foraging time lowers both energy intake by the female
and her ability to feed the chicks. Lack of snow cover, and freezing temperatures,
influence the depth of soil frost and can reduce both food availability in spring and
woodcock productivity. In addition, weather during the brood period and condition of the
female at the end of incubation play important roles in determining the number of
offspring produced. Thus, habitat management should strive to provide high earthworm
biomass in a variety of suitable feeding sites to ameliorate the effects of weather. Key
words: American woodcock, temperature, frost, precipitation, snow, soil,
nutrition, productivity

515

, W. B. Krohn, R. B. Owen Jr., J. R. Longcore, and G. F. Sepik. 1993. Effects

of weather on earthworm abundance and foods of the American woodcock in
spring. Proceedings of the American Woodcock Symposium 8, U. S. Fish and
Wildlife Service Biological Report 16:26-31. Abstract: Earthworms composed greater
than 90% (dry weight and volume) of food consumed by 48 American woodcocks
(Scolopax minor) collected during the springs of 1987-89 on the Moosehorn National
Wildlife Refuge, Maine. The availability of earthworms seemed related to amount of
snow in winter and persistence of frost in the soil in spring. When earthworms were less
available, the intake of food by woodcocks declined and the relative importance of litter-
inhabiting prey (Coleoptera and Araneae) increased. Reduced intake of food lowered
the body mass of female woodcocks in 1989 and evidently caused a 3-4 week delay in
nesting. Results suggest that conditions in winter and spring affect availability of food
and subsequently influence body mass, time of nesting, and reproductive success of
woodcock. Key words: American woodcock, snow, frost, temperature, nutrition,
reproduction

516

, M. W. Sayre, and W. E. Dodge. 1989. Winter use of agricultural habitats by

wild turkeys in Massachusetts. Journal of Wildlife Management 53(1): 30-33.
Abstract: We examined winter habitat use by eastern wild turkeys (Meleagris gallopavo
sylvestris) in western Massachusetts during 1984 and 1985. Use-availability analyses
revealed that flocks used cropland, pasture, and softwood habitats more, and
hardwoods, mixed woods, and abandoned fields less than expected (P < 0.05). Flocks
spent 54% of the diurnal period in cropland and pastures and spent more time feeding
on manure spread on fields than on any other single food source. During period of deep
snow, turkeys restricted their movements to < 20 ha, used softwood stands and
adjacent cropland and pastures, and fed largely on manure. Decline of dairy farming in

Jtf 231

N^

^ fc* ^

c^>

Massachusetts may adversely affect local turkey populations. Key words: turkey,
snow, habitat use, behavior

517

Van Kooten, G. C, A. J. Eagle, and M. E. Eiswerth. 2007. Determinants of
threatened sage-grouse in northeastern Nevada. Human Dimensions of Wildlife
12:53-70. Abstract: We examined potential human determinants of observed declines in
greater sage-grouse (Centrocercus urophasianus) populations in Elko County, Nevada.
Although monitoring of sage-grouse has occurred for decades, monitoring levels have
not been consistent. This article contributes to the literature by normalizing grouse
counts by the annual effort to count them, performing regression analyses to explain the
resulting normalized data, and correcting for sample selectivity bias that arises from
years when counts were not taken. Our findings provide some evidence that cattle
grazing contributes to a reduction in sage-grouse populations, but this result should be
interpreted with caution because our data do not include indications about the timing
and precise nature of grazing practices. Annual variations in weather appear to be a
major determinant after statistically controlling for human interactions with the
landscape, suggesting that climate change is a key potential long-run threat to this
species. Keywords: sage-grouse, climate change, population dynamics

518

Vehrencamp, S. L., J. W. Bradbury, and R. M. Gibson. 1989. The energetic cost of
display in male sage-grouse. Animal Behavior 38:885-896. Abstract: The energetic
expenditure of displaying male sage-grouse, Centrocercus urophasianus, was
measured for 18 individuals in the field using the doubly labeled water technique. Daily
energy expenditure increased significantly with increased display rate, increased time
spent on the lek, and decreased ambient temperature. Daily energy expenditure for the
most vigorously displaying males was two times higher than for a non-displaying male
and four times higher than basal metabolic rate. Estimates of the instantaneous rate of
energy expenditure during display ranged from 13 x 9 to 17 x 4 times basal metabolic
rate. The effort devoted to display differed markedly among males and was correlated
with certain other male characteristics. Males that attended leks were in better condition
(higher body weight relative to size) than non-attenders, but among lek attenders
condition was negatively correlated with increased display effort. Key words: sage-
grouse, temperature, behavior

519

Voronin, R. N. 1991. Variability in the lay size of Lagopus lagopus in the
Bolshezemelskaya Tundra, Russian SFSR, USSR. Ekologiya 1:68-72. Abstract:
Dynamics of the lay size of the willow grouse (L. lagopus) were studied from 1970 to
1984 in the Bolshezemelskaya Tundra. In different years the average lay varied from
6.09±0.39 to 1 0.85±0. 1 5 eggs. A series of biotic and abiotic factors serves as the basis
for these variations (i.e., physiological status of individuals, the structure of the
population, and weather conditions). Key words: willow grouse, productivity,
weather

232 ^

4

fv<^^

520

Vysotsky, V. G. and I. V. Iljinsky. 2004. Method of forecasting changes in the
abundance of woodcock (Scolopax rusticola) based on weather conditions on the
wintering grounds. Proceedings of the Zoological Institute, Russian Academy of
Sciences 300:165-174. Notes: Page 165 — "It is known that woodcock winter survival in
France covahed positively with mean winter nocturnal temperature. We... tested the
hypothesis that annual survival rates depend on winter precipitation and temperature
within woodcock main winter range." Page 1 72 — "Average monthly air temperature for
November-February in United Kingdom, France, and Italy is an important index for bird
annual survival process. It is known that temperature affects directly activity (and hence
availability) of many species of earthworms, which are the basis of woodcock diet.
Temporal variability in earthworm availability may be an important determinant of
woodcock winter survival. Moreover, annual variation in woodcock abundance during
breeding in north-western Russia may well be explained by winter severity in the main
part of the wintering area." Key words: woodcock, temperature, precipitation, index,
model, mortality, nutrition

521

Walker, B. L., D. E. Naugle, K. E. Doherty, and T. E. Cornish. 2007. West Nile Virus
and greater sage-grouse: estimating infection rate in a wild bird population. Avian
Diseases 51:691-696. Summary: Understanding impacts of disease on wild bird
populations requires knowing not only mortality rate following infection, but also the
proportion of the population that is infected. Greater sage-grouse (Centrocercus
urophasianus) in western North America are known to have a high mortality rate
following infection with West Nile virus (WNv), but actual infection rates in wild
populations remain unknown. We used rates of WNv-related mortality and
seroprevalence from radiomarked females to estimate infection rates in a wild greater
sage-grouse population in the Powder River basin (PRB) of Montana and Wyoming
from 2003 to 2005. Minimum WNv-related mortality rates ranged from 2.4% to 13.3%
among years and maximum possible rates ranged from 8.2% to 28.9%. All live-captured
birds in 2003 and 2004 tested seronegative. In spring 2005 and spring 2006, 10.3% and
1.8%, respectively, of newly captured females tested seropositive for neutralizing
antibodies to WNv. These are the first documented cases of sage-grouse surviving
infection with WNv. Low to moderate WNv-related mortality in summer followed by low
seroprevalence the following spring in all years indicates that annual infection rates
were between 4% and 29%. This suggests that most sage-grouse in the PRSB have
not yet been exposed and remain susceptible. Impacts of WNv in the PRB in the near
future will likely depend more on annual variation in temperature and changes in vector
distribution than on the spread of resistance. Until the epizootiology of WNv in
sagebrush-steppe ecosystems is better understood, we suggest that management to
reduce impacts of WNv focus on eliminating man-made water sources that support
breeding mosquitoes known to vector the virus. Our findings also underscore problems
with using seroprevalence as a surrogate for infection rate and for identifying competent
hosts in highly susceptible species. Key words: sage-grouse, temperature, insects,
disease

% f\^>^

233

c£^

522

Wallestad, R. 1975. Life history and habitat requirements of sage-grouse in
central Montana. Montana Department of Fish and Game, Helena, USA. Notes:
Under the heading Limiting Factors, the author gives a brief treatment of weather.
Wallestad notes "The success or failure of sage-grouse in a particular year has
generally been attributed to weather conditions during hatching. Cold, rainy weather
were the conditions usually blamed for poor productivity." He cites research by
Wallestad and Watts that showed no correlation between productivity and rainfall during
the hatching period, and an inverse correlation between productivity and rainfall during
the egg-laying period. Heavy rain (greater than 1 inch) during the egg-laying period
caused a late hatch resulting in poor productivity (less than 400 juveniles:100 females).
Total spring precipitation, as it potentially affected spring greenup of vegetation, further
explained variations in productivity. Even if rainfall was optimum during the egg-laying
period, production would be poor if total spring precipitation during the growing season
was inadequate for necessary plant growth (les than 3 inches from mid-April thorough
mid-June). They found no correlation between temperature and productivity. Key
words: sage-grouse, precipitation, temperature, productivity, vegetation

523

and R. C. Watts. 1972. Factors effecting annual sage-grouse productivity

in central Montana. Small Game Research W-120-R-3, Montana Department of
Fish and Game, Helena, USA. Abstract: Analysis of 10 years (1962-1971) of sage-
grouse population and selected climatic variables have identified several factors
apparently affecting sage-grouse productivity in central Montana. Years with a late
hatch have significantly poorer productivity than years with a normal hatch. Total rainfall
or storm intensity during the egg laying period is inversely correlated to productivity.
However, a lack of moisture necessary for spring green-up of vegetation can cause
poor productivity. There is also a correlation between productivity and percent of
successful yearling females in the population. None of the other climatic or population
variables analyzed affected productivity. Key words: sage-grouse, precipitation,
productivity

524

Wambolt, C. L, A. J. Hapr, B. L. Welch, N. Shaw, J. W. Connelly, K. P. Reese, C. E.
Braun, D. A. Klebenow, E. D. McArthur, J. G. Thompson, L. A. Torell, and J. A.
Tanaka. 2002. Conservation of greater sage-grouse on public lands in the western
U.S.: implications of recovery and management policies. PACWPL Policy Paper
SG-02-02. Policy Analysis Center for Western Public Lands, Caldwell, USA. Notes:
Page 5 — "In years of high precipitation, succulent forbs may persist in sagebrush
uplands and change bird movements... In very dry years, sage-grouse may use summer
habitat until November. Grouse may also move to higher elevations containing moist
sites." Page 6 — "...fires in sagebrush habitats may decrease insects used by grouse as
food.... In some areas, meadows are important summer habitat for sage-grouse
because they provide an abundance of succulent forbs. These areas are especially
important during drier summers.... Desiccation and frost kill forbs in summer foraging
areas... .Unlike other seasonal habitats that have two or more components important to

234 ^

^

H ^ ^

.1/^

<^

sage-grouse, the only critical habitat component important during winter is sagebrush.
Sage-grouse are totally dependent upon sagebrush habitats for food and cover
throughout the winter period." Page 7 — "...low sagebrush and black sagebrush also
provide important winter habitats when snow depth allows grouse access to these
relatively low-growing shrubs. Sagebrush canopy cover in sage-grouse winter habitats
ranges from 12 to 43%, and sagebrush height above snow ranges from 8 to 22 inches
(20 to 56 cm) throughout the species' range... .On a landscape scale, sage-grouse
winter habitats should allow grouse access to sagebrush under all snow
conditions... There is little, if any, evidence that severe winter weather affects sage-
grouse populations unless sagebrush cover has been eliminated or significantly
reduced. However, without adequate sagebrush leaves available for winter forage, body
mass may decrease and spring breeding displays may be reduced." Page 25 — "
Drought: When drought is severe on most allotments, grazing regimes are already, or
should be, altered to comply with the agency standards and guidelines. Public grazing
lessees often feel that grouse advocates are singling them out unfairly. They feel that
many factors affect the grouse and, until other factors are addressed as well, grazing
should not be further restricted beyond what is required to meet standards and
guidelines. Grouse advocates and most environmentalists counter by pointing out that if
grouse habitats were 'optimal' then the grouse population should be able to withstand
drought periods without such serious declines." Key words: sage-grouse,
precipitation, fire, frost, snow, vegetation, nutrition, habitat use, reproduction,
drought, grazing

525

Wang, G., N. T. Hobbs, H. Galbraith, and K. M. Giesen. 2002. Signatures of large-
scale and local climates on the demography of white-tailed ptarmigan in Rocky
Mountain National Park, Colorado, USA. International Journal of Biometeorology
46:197-201. Abstract: Global climate change may impact wildlife populations by
affecting local weather patterns, which, in turn, can impact a variety of ecological
processes. However, it is not clear that local variations in ecological processes can be
explained by large-scale patterns of climate. The North Atlantic Oscillation (NAO) is a
large-scale climate phenomenon that has been shown to influence the population
dynamics of some animals. Although effects of the NAO on vertebrate population
dynamics have been studied, it remains uncertain whether it broadly predicts the impact
of weather on species. We examined the ability of local weather data and NAO to
explain the annual variation in population dynamics of white-tailed ptarmigan (Lagopus
leucurus) in Rocky Mountain National Park, USA. We performed canonical correlation
analysis of the demographic subspace of ptarmigan and local-climate subspace defined
by the empirical orthogonal function (EOF) using data from 1975 to 1999. We found that
two subspaces were significantly correlated on the first canonical variable. The Pearson
correlation coefficient of the first EOF values of the demographic and local-climate
subspaces was significant. The population density and the first EOF of local-climate
subspace influenced the ptarmigan population with 1-year lags in the Gompertz model.
However, the NAO index was neither related to the first two EOF of local-climate
subspace nor to the first EOF of the demographic subspace of ptarmigan. Moreover,
the NAO index was not a significant term in the Gompertz model for the ptarmigan
population. Therefore, local climate had stronger signature on the demography of

* 1^< / .. . , 235

% K<c^-

**

ptarmigan than did a large-scale index, i.e., the NAO index. We conclude that local
responses of wildlife populations to changing climate may not be adequately explained
by model that project large-scale climatic patterns. Key words: white-tailed
ptarmigan, NAO, index, model, population dynamics

526

Ward, J. M., D. J. McCafferty, G. D. Ruxton, and D. C. Houston. 2007. Thermal
consequences of turning white in winter: a comparative study of red grouse
Lagopus lagopus scoticus and Scandinavian willow grouse L-l. lagopus. Wildlife
Biology 13(2):120-129. Abstract: The red grouse Lagopus lagopus scoticus differs from
the willow grouse L. L. lagopus of mainland Europe in not developing a white winter
plumage. Previous studies have suggested that plumage coloration in birds can have
important consequences for heat transfer through the feather layer. We examined the
thermal consequences of plumage coloration in both subspecies of grouse. There were
no differences in feather density, plumage depth or thermal resistance of the plumage
between dark rufous Scottish and white Scandinavian grouse. In still air, heat gained
from simulated solar radiation was greater through dark than through white plumage.
However, in wind there was no difference in heat load between dark and white
plumages. Our study suggests that there may be a tradeoff between thermal and
camouflage benefits of plumage color for grouse in the wild. Key words: red grouse,
willow grouse, thermoregulation, solar radiation, wind, morphology

527

Warner, R. E. and L. M. David. 1982. Woody habitat and severe winter mortality of
ring-necked pheasants in central Illinois. Journal of Range Management
46(4):923-932. Abstract: Ring-necked pheasant (Phasianus colchicus) populations
declined 44-82%, primarily from exposure to precipitation and severe wind chill, in the
severe winters of 1976-77 and 1977-78 on 5 study areas in east central Illinois.
Pheasant abundance, estimated by cock-call counts, and woody vegetation were
measured on 45 1,036-ha subunits during 1976-78. Multiple regression analyses
indicated no relationship between pheasant abundance on the 45 subunits before or
after severe winter weather, and the abundance, growth forms, or arrangement on the
landscape of woody vegetation. Declines in cock calls, 1976-78, were a mathematical
function of pheasant densities prior to severe winter weather (r = 0.94, P < 0.001 ). Key
words: ring-necked pheasant, temperature, wind, precipitation, mortality

528

Watson, A., R. Moss, and P. Rothery. 2000. Weather and synchrony in 10-year
population cycles of rock ptarmigan and red grouse in Scotland. Ecology
81(8):2126-2136. Abstract: Rock ptarmigan (Lagopus mutus) on two adjacent
submassifs, the red grouse (Lagopus lagopus scoticus) on lower ground between them,
showed largely synchronous ~10-year cycles during a ~50-year study on the infertile
Cairngorms massif of Scotland. Adult birds of both these Lagopus species were
counted along transect walks. Both species showed the very low mid-1 940s trough
previously recorded for tetraonids in much of northwest Europe. Each of five
subsequent peaks in all three populations fell within a year of one another, and 1-2 year

236 ^

^

>->' fc* >f*

c£^>

after cyclic high June temperatures at a nearby village. Troughs were less synchronous.
A model with lagged June temperatures and fourth-order delayed density dependence,
with no input from observed bird numbers after the first 4 years, gave a good postdiction
of rock ptarmigan numbers on the bigger submassif for 49 years, suggesting a weather
cycle entraining on rock ptarmigan cycle. However, June temperatures had little
explanatory value for rock ptarmigan numbers on the smaller submassif. Indirect
evidence suggested that synchrony between the two rock ptarmigan trajectories may
have been due partly to emigration from the bigger to the smaller submassif. The
population trajectory of red grouse resembled that of rock ptarmigan on the smaller
submassif more closely than the two rock ptarmigan trajectories resembled one other.
Hence synchrony depended more on local circumstances than on species. Key words:
rock ptarmigan, red grouse, temperature, population dynamics, behavior

529

, , and S. Rae. 1998. Population dynamics of Scottish rock ptarmigan

cycles. Ecology 79(4): 1 174-1 192. Abstract: Scottish rock ptarmigan (Lagopus mutus)
showed unstable population dynamics, with density-dependent features differing
between areas on rich and poor soils. The study was conducted on arctic-alpine land on
four submassif, three of nutrient-poor granite and one of richer schist. Ptarmigan
numbers cycled with a period usually of -10 yr, but sometimes 6 yr. Delayed density
dependence was detected in populations on granite submassif, but not on schist,
despite significant cyclicity there. Spring densities and the number of young reared per
brood were higher on schist than on granite. Population change from one spring to the
next was related to intervening breeding success on both granite and schist, but to
summer loss only on granite. The same factors that determined variations in clutch size
among years also largely affected breeding success. Population change, breeding
success, summer loss, and clutch size showed delayed density dependence on granite,
but not on schist, where numbers fluctuated more erratically. On granite, but not schist,
early plant growth was followed by larger clutches, breeding success was related to
June air temperature, and June temperature tended to peak 1-2 yr before ptarmigan
peaks. Cyclic June temperature may have entrained the timing of ptarmigan cycles on
granite. Results on both schist and granite were inconsistent with expectations from
predator-prey hypotheses for cycles. In years of rising numbers, hens laid more eggs
and reared more chicks per extra egg laid than in years of decline, and the proportion of
surviving young recruited into the spring population was probably greater. Our
hypothesis is that Scottish rock ptarmigan show unstable dynamics, driven by intrinsic
positive feedback between recruitment in one year and the next, regulated by negative
feedback between recruitment and density, and modified by the physical factors of
weather and soil fertility that affect female nutrition and breeding success. Key words:
rock ptarmigan, temperature, population dynamics, soil

530

Watts, C. R. 1973. Factors affecting annual sage-grouse productivity in central
Montana. Western States sage-grouse Workshop Proceedings 8:49. Abstract:
Analysis of 10-years (1962-71) of wing data has identified several factors apparently
affecting sage-grouse (Centrocercus urophasianus) productivity in central Montana.

f \ ^ ^k

^

Years with a late hatch have significantly poorer productivity than years with a normal
hatch. Total rainfall or storm intensity during the egg laying period is inversely correlated
to productivity. However, a lack of moisture necessary for spring green up of vegetation
can cause poor productivity. Productivity and percent of successful yearling females in
the population is directly correlated. None of the other climatic or population variables
analyzed affected productivity. Key words: sage-grouse, precipitation, productivity

531

Wayne, A. T. 1899. Destruction of birds by the great cold wave of February 13 and
14, 1899. Auk 26:197-198. Notes: "The woodcock arrived in countless thousands. Prior
to their arrival I had seen but two birds the entire winter. They were everywhere and
were completely bewildered. Tens of thousands were killed by would be sportsmen, and
thousands were frozen to death. The great majority were so emaciated that they were
practically feathers and of course were unable to withstand the cold." [The cold wave
which struck the coast of South Carolina was the severest recorded for 200 years
including deep snow up to 5 in. and drifts 2 ft. deep with cold temperatures.] Key
words: woodcock, snow, temperature, mortality

532

Wegge, P. and L. Kastdalen. 2007. Pattern and causes of natural mortality of
capercaillie, Tetrao urogallus, chicks in a fragmented boreal forest. Annales
Zoologici Fennici 44:141-151. Abstract: Over a 3-year period, we equipped 115 newly
hatched capercaillie chicks in 29 broods with small radio transmitters in southeast
Norway. Besides monitoring the fate of the chicks, we measured the abundance of
microtine rodents and insect food and, together with weather records, we examined the
observed mortality in relation to these factors. On average, 57% of the chicks died
within the first month of life. Mortality was highest during the first 3 weeks, coinciding
with the period when chicks feed almost exclusively on insects, depend heavily on their
mother for maintaining body warmth, and cannot fly well. Predation was by far the most
important proximate cause of mortality, accounting for 90% of all observed losses. Only
7% of the losses could be ascribed to direct effects of cold and wet weather, all
recorded during the first 8 days of life. However, because predation losses were also
quite high during and immediately after heavy rainfalls, adverse weather probably
predisposed the chicks to mammalian predation. Most chicks were killed by mustelids,
mainly pine martens; low numbers of red fox due to sarcoptic mange probably explains
the low proportion taken by this predator. Among known predation losses (n = 40),
goshawks accounted for a minimum of 25%. Chick mortality during the first month
varied markedly (29%-83%) among the three years. It was highest in the year when
June weather was exceptionally wet and cold; the same year when the density of
microtine rodents and food abundance in terms of insect larvae were highest. Net
production in late August was poorly related to chick loss during the first month
posthatch Key words: capercaillie, temperature, precipitation, predation, chick
survival

238 y

>>' (& **

<£

533

Weigand, J. P. 1980. Ecology of the Hungarian partridge in north-central Montana.
Wildlife Monographs 74:3-106. Notes: Under the subheading "Effects of Weather",
Weigand states: "Findings of the current study generally agreed that weather
conditions, particularly amounts of precipitation, in late spring-early summer influenced
partridge production. A dry spring results in early nesting and yields above average
production unless heavy and/or prolonged rainfall is received during the first 3 weeks
after peak hatching periods. The latter conditions will effectively reduce anticipated
population gains. Above average precipitation in April probably will delay or interfere
with early nesting, but low precipitation received after hatching will permit greater chick
survival and reasonably good, overall production. Heavy rainfall during nesting and
hatching probably precludes any reasonable degree of successful production for that
year." Key words: Hungarian partridge, precipitation, temperature, productivity

534

. 1977. The biology and ecology of Hungarian (European gray) partridge

(Perdix perdix L.) in northcentral Montana. Dissertation, Montana State
University, Bozeman, USA. Notes: On page 176 under "Effects of Weather" Weigand
discusses the impacts of precipitation and temperature on nesting and brooding. He
found that in 1969 some incubated nests were adversely affected in June and renesting
occurred. Spring of 1970 was the wettest period in 5 years. Above normal precipitation
in April/May and below normal temperatures apparently delayed nesting. Mean brood
sizes were 1% below the 5-year mean in July and 11% in August. In 1971 a dry early
brooding period favored chick survival and the result was above normal chick
production. "Findings of the Montana study generally agreed that weather conditions,
particularly amounts of precipitation, in late spring-early summer influenced partridge
production. A dry spring results in early nesting which yields above-average production
unless heavy and/or prolonged rainfall is received during the first three weeks after
peak hatching periods. The latter conditions will effectively reduce anticipated
population gains. Above-average precipitation in April will probably delay or interfere
with early nesting but low precipitation received during post-hatching will permit greater
chick survival and reasonably good overall production. Heavy rainfall during nesting and
hatching probably precludes any reasonable degree of successful production for that
year." Page 245 - "Shelterbelts and tree groups were in greatest association during
Dec-Feb (>95%) and least in October (60%). Low use of shrubs in winter was attributed
to these types filling in with snow, thereby severely limiting their value as protective
cover. Page 265 — "Partridge proximity to woody cover I winter appeared to be
influenced by weather and ground conditions. Partridge were less mobile early in milder
than average (1969-70) and average (1970-71) winters than in a severe winter (1971 -
72)... recurrence of severe winter conditions caused the birds to resume close
association." Key words: Hungarian partridge, temperature, snow, precipitation,
habitat use, reproduction, population dynamics

535

and R. G. Janson. 1976. Montana's ring-necked pheasant. Montana

Department of Fish and Game, Helena, USA. Summary: Under Decimating Factors,

239

t f^<tf*^

% m <o> 9

the authors present a discussion of weather and its impact on productivity. Among the
weather factors that exert such impacts are snow depth, precipitation, hail, temperature,
and man-made weather alterations. Key words: ring-necked pheasant, temperature,
snow, hail, precipitation, mortality, productivity, habitat use

536

Whitaker, D. M. 2003. Ruffed grouse (Bonasa umbellus) habitat ecology in the
central and southern Appalachians. Dissertation, Virginia Polytechnic Institute
and State University, Blacksburg, USA. Notes: Page 32 — "Weather-related variables
influenced roost site selection by grouse. Although snow depths were never sufficient to
allow snow burrowing, ground roosting increased when any snow cover was present.
Precipitation during the night also influenced the incidence of ground roosting, with the
smallest proportion of ground roosts observed during nights having rainfall, and the
greatest proportion observed during nights having snowfall. Grouse shifted from
predominantly using above ground roosts at temperatures above freezing to using
ground level roosts when temperatures fell below 0°C. However this relationship may
have resulted from the association of snow cover with cold weather, and no difference
was detected when observations were restricted to nights having no snow cover. Air
temperature did not influence the proportion of grouse roosting in evergreen
vegetation." Page 34 — "Ground roosting was frequent on nights when temperatures fell
below freezing and least common on nights having rainfall, when leaves were wet.
Because snow cover was always shallow, ground roosts in snow typically extended
down into the leaf litter. Snow cover is associated with low temperatures, and it may be
that the combination of these factors contributes to a general superiority of ground
roosts during cold weather. Finally, grouse in the region are red-phased, so are
inconspicuous to predators when buried flush with the similarly colored leaf litter. Given
the behavioral parallel to snow burrowing, the development of such leaf-roosting
behavior seems reasonable." Key words: ruffed grouse, snow, temperature,
precipitation, behavior, habitat use

537

. and D. F. Stauffer. 2003. Night roost selection during winter by ruffed

grouse in the central Appalachians. Southeastern Naturalist 2(3):377-392.

Abstract: In northern regions, ruffed grouse (Bonasa umbellus) conserve considerable
energy during winter by burrowing under snow cover to roost. When conditions are
unsuitable for snow burrowing grouse almost invariably roost in conifers. We studied
selection of winter night roosts by ruffed grouse in western Virginia, a region where
snow accumulations are rare and transient. Grouse almost always used ground roosts
when snow was present even though snow was never deep enough for snow
burrowing. When snow was absent grouse did not show any clear preference in roost
microsite type, and were found roosting in and under deciduous and evergreen trees
and shrubs, in brush piles, and in leaf litter. We hypothesize that this ambivalence to
conifers was due in part to persistent accumulations of fallen oak leaves, which likely
afford grouse good thermal cover and concealment. Grouse were frequently found at
low elevations during daytime, but rarely roosted in bottoms. This suggests daily

240 ^

^

>->" fc&. ^

elevational movements, possibly to avoid cold air settling in low-lying areas during night.
Key words: ruffed grouse, snow, temperature, behavior, thermoregulation

538

Whiting, R. M. Jr., D. A. Haukos, and L. M. Smith. 2005. Factors affecting January
reproduction of American woodcock in Texas. Southeastern Naturalist 4(4):639-
646. Abstract: Scolopax minor (American Woodcock) populations have been declining
for the past several decades. Weather conditions have been hypothesized to affect
reproductive efforts in February and March in the southern United States, but similar
influences on January breeding activities are unknown. We used a 17-year harvest data
set from eastern Texas to examine the influence of several temperature and
precipitation measures, female body mass, and forest habitat type on the occurrence of
reproduction by woodcock in January. Only adult females exhibited characteristics of
breeding activity. There was annual variation (0-29%) in the occurrence of breeding
adult females, but local temperature and precipitation measures were not correlated
with January reproduction. More breeding woodcock were harvested in open sapling
stands than other habitats. Environmental factors other than monthly temperature and
precipitation may influence January reproductive efforts of woodcock in eastern Texas.
We suggest investigation of timing of fall migration and condition of the birds upon
arrival on wintering grounds as potential factors influencing reproductive efforts in
January. Key words: American woodcock, temperature, precipitation,
reproduction

539

Wiebe, K. L. and K. Martin. 1995. Ecological and physiological effects on egg
laying intervals in ptarmigan. Condor 97:708-717. Abstract: Birds lay eggs at
different rates and at different times of the day. Some species also show skipped days
during the laying sequence, "laying gaps," but patterns of egg laying have not been well
studied. We compared timing of laying during the day, laying gaps, and laying rates
(number of eggs/day) for white-tailed ptarmigan (Lagopus leucurus) and willow
ptarmigan (L. lagopus). Both species tended to lay eggs near midday, but willow
ptarmigan had fewer laying gaps and faster laying rates than white-tailed ptarmigan.
The variation in laying rates among individuals was greater for white-tailed ptarmigan
which had a bimodal distribution if inter-egg intervals, long (mean = 44 hr) and short
(mean = 26 hr). Laying gaps were not associated with spring body condition but severe
spring storms seemed to cause some laying delays. The patterns and frequency of
gaps observed within and between species may be the result of nutrient constraints on
egg formation in conjunction with physiological mechanisms which control a time-
window for egg laying. Key words: white-tailed ptarmigan, willow ptarmigan, severe
weather, reproduction

540

Wiley, E. N. II and M. K. Causey. 1987. Survival of American woodcock chicks in
Alabama. Journal of Wildlife Management 51(3):583-586. Abstract: Survivability of
American woodcock (Scolopax minor) chicks hatched in Alabama was investigated from
March through April 1984-85. Ten broods (36 chicks) were radio tracked intermittently

\ f*te*t»c

241

Q

between hatching and brood breakup (at 32 days). Eleven chicks died during this
period, 10 after fledging (15 days). The most common cause of mortality was predation
(4 avian, 2 mammalian, and 2 unknown). One death was accidental and 2 died of
unknown causes. Weather conditions apparently did not result in any chick losses. The
pre-fledging survival probability (0.95) possibly indicates a high survival rate for
Alabama-hatched chicks prior to fledging. The most vulnerable interval of the brood-
rearing period occurred after fledging (16-32 days) when the survival probability
decreased to 0.67 but may have been influenced by study methods. Key words:
American woodcock, mortality, weather conditions, chick survival

541

Williams, G. R. 1954. Population fluctuations in some northern hemisphere game
birds (Tetraonidae). Journal of Animal Ecology 23(1 ):1 -34. Notes: Page 26-".. .it
does seem that weather systems must play an important part. But, so far, all attempts to
find a climatic factor correlated with the population changes have failed. Chitty has
suggested how random variations in weather might tend to correlate cycles: in
widespread favorable years, peak populations which had already approached their
maximum densities might be unable to reach much higher levels; whereas sparser
populations would be increasing rapidly towards the maximum possible for each
particular environment — thus all populations would tend to approach their respective
peaks together. On the other hand, if a widespread spell of severe weather were to
result in mortality rates which increased with population density, then all populations
would be reduced to similarly low levels. The effects of these two extreme conditions
should be the promotion of some degree of synchrony. Key words: weather extremes,
population dynamics

542

Williams, H. W. and A. W. Stokes. 1965. Factors affecting the incidence of rally
calling in the chukar partridge. Condor 67:31-43. Summary: The purpose of this
study was to measure the effects of physical and biotic factors on rates of rally calling in
chukar partridge. Calling was most frequent around sunrise and sunset with little calling
in between. Of the physical factors, light intensity and precipitation had the greatest
effect on calling. The four factors of light intensity, temperature, wind, and rain
accounted for 45% of the variation in morning calling and 54% of the variation in
evening calling. Low light intensities associated with cloudiness during the day did not
always result in increased calling rates. Wind velocities ranged from 0 to 45 mph;
however, there were only 47 sampling periods with velocities in excess of 10 mph. At
these times the chukars normally remained close to cover and did not call. The
decrease in calling with increasing winds could be due to either the depressing effect of
wind on the birds or the inability of a person to hear chukar calls above the noise of the
wind. Rainfall normally caused the chukars to move to cover and stop calling. When
unusually motivated, chukars may continue to call despite wind and rain. Key words:
chukar partridge, wind, precipitation, temperature, census, clouds

242 ^

*>' ^ f*

543

Wilson, W. B., M. J. Chamberlain, and F. G. Kimmel. 2005. Survival and nest
success of female wild turkeys in a Louisiana bottomland hardwood forest.
Proceedings of the Annual Conference Southeast Association Fish and Wildlife
Agencies 59:126-134. Abstract: Survival of female wild turkeys (Meleagris gallopavo)
influences turkey productivity. Although patterns of survival and productivity have been
extensively researched in most forested landscapes, little information is available for
female turkeys in bottomland hardwood systems, although importance of these systems
is widely recognized. Therefore, we captured and radiomarked 39 female wild turkeys in
a bottomland hardwood forest in south-central Louisiana during 2001-2004. Mean
annual survival was 0.67. Survival was greatest during preincubation (1.00) potentially
because of increased habitat sampling and movement during this period. Fall-winter
survival was high (0.93), likely attributable to stable foraging resources and a lack of
illegal and legal harvest during this period. Lowest survival occurred during incubation
(0.75) and brood-rearing (0.83), primarily as a result of increased risks of predation
associated with nesting and brood rearing. Nest initiation rates (33%) were among the
lowest reported, likely attributable to high nest loss from predation and flooding prior to
completion of laying. Nest success of females reaching onset of incubation was 38%.
Our findings suggest that the wild turkey population on our study site balances
exceptionally low productivity with relatively high adult female survival. To ensure
sustainable populations of wild turkeys, managers should monitor relationships between
survival and productivity. Specific to our study site, improvements in nesting habitat may
be needed to increase nest success and recruitment. Key words: turkey, flooding,
reproduction, recruitment

544

Windingstad, R. M., S. M. Kerr, R. M. Duncan, and C. J. Brand. 1988.
Characterization of an avian cholera epizootic in wild birds in western Nebraska.
Avian Diseases 32(1):124-131. Summary: Avian cholera killed an estimated 2,500
birds in western Nebraska and eastern Wyoming from 28 November 1985 to late
January 1986. Wild mallards (Anas platyrhynchos) suffered the most losses. Other wild
waterfowl, wild turkeys (Meleagris gallopavo), a few domestic fowl, and a bald eagle
(Haliaeetusleucocephalus) also died. Pasteurella multocda serotype 1 was the
predominant isolate from these carcasses. Cold, wet weather persisted throughout the
outbreak, but daily losses in the flock of 50,000 mallards using the area were low.
Pasteurella multocida was isolated from nasal swabs of 35 of 37 cattle from a feedlot in
which many of these mallards were feeding. Eighty percent of the cattle isolates had
antigenic characteristics of serotype 3 or serotype 3 with cross-reactivity. Isolates from
wild mallards, wild turkeys, and the bald eagle were virulent to game-farm mallards
when inoculated subcutaneously, but P. multocida isolates from cattle were not. Key
words: turkey, disease, temperature, precipitation, mortality, Avian cholera

545

Wiseman, D. S. and J. C. Lewis. 1981. Bobwhite use of habitat in tallgrass
rangeland. Wildlife Society Bulletin 9(4):248-255. Notes: Page 250— "Coveys often
rested in tall shrub habitat for several hours during midday." Page 251 — "Coveys also

*~> fc* ^

M A <V i . L- 243

Cgj ?}jfe £^> ^

showed preference (24% of use) for short shrubs that made up only 9% of the home
ranges. Coveys used short shrubs for resting and for protection from weather and
predators... during snow, rain, and prolonged cold some coveys spent most of the day in
short shrubs and in December and January often roosted there." Key words: bobwhite
quail, snow, precipitation, temperature, habitat use, vegetation, behavior

546

Wisinski, C. L. 2007. Survival and summer habitat selection of male greater sage-
grouse {Centrocercus urophasianus) in southwestern Montana. M.S. Thesis,
Montana State University, Bozeman, USA. Abstract: During the 20th century, greater
sage-grouse (Centrocercus urophasianus) populations in North America have declined
by 69-99%. In southwest Montana little is known about the factors leading to declines in
sage-grouse populations; as a result, there are strong concerns regarding sage-grouse
population trends and habitat quality. I used radio-marked male sage-grouse to obtain
known-fate survival data and provide locations for habitat analyses. From 2001-2005,
45 male sage-grouse were instrumented and monitored. The estimated annual survival
rate was 0.34 (95% CI: 0.21 toO 0.47). In 2004 and 2005, I measured vegetation
characteristics at 78 habitat plots (43 used, 35 available). I used logistic regression to
model habitat selection; grass height was an important predictor of use, but the
negative relationship between the probability of a site being used and grass height was
opposite of what I had predicted. Further research is needed to determine whether this
behavior is adaptive. An important predictor of use in my exploratory analysis was solar
radiation index (SRI). The negative effect of SRI on habitat selection may have been a
result of sampling only during summer daylight hours, and SRI may have acted as a
proxy for microsite temperature. Further research is needed over a wide variety of
conditions to determine how habitat components relate to or interact with each other,
habitat selection by sage-grouse, and survival of sage-grouse. Key words: greater
sage-grouse, solar radiation, index, habitat use, vegetation

547

Withers, P. C. and T. M. Crowe. 1980. Brain temperature fluctuations in helmeted
guineafowl under semi-natural conditions. Condor 82:99-100. Notes: Page 100-
...the scatter in Tbr for the guineafowl indicates that environmental variables, such as air
temperature, radiation, convection and rain, probably influence Tbr greatly. The highest
Tbr's (41.8±SE 0.1 °C, r?=9) were consistently recorded between 09:00 and 13:00 when
the aviary was in direct sunlight. The lowest Tbr's were 38.6±0.4°C (n=16) during the
night and particularly after rain. The Tbr's during the day, in diffuse but not direct
sunlight, were intermediate at 39.9±0.2°C (n=32). Brain temperature was significantly
higher (ANOVA test, P<0. 001 ) during the day with direct solar radiation than at other
times, but Tbr was not significantly correlated with either shade or sunlight air
temperature. Although Tbr was significantly lower with diffuse radiation (P<0.001) and
during the night (P<0.001), Tbr was not correlated with shade air temperature. Free-
ranging guineafowl are clearly able to maintain higher Tbr, through physiological or
behavioral means, than birds in the laboratory... These data for the Tbr regulation of
Helmeted Guineafowl under semi-natural conditions, although preliminary, clearly
indicate that: (1) guineafowl under semi-natural conditions have a markedly labile Tbr;

244 ^

^

>->" fc* >f*

±Ll'

(2) Tbr is generally greater for free-ranging guineafowl than for birds in the laboratory,
reflecting the importance of incident solar radiation and diffuse radiation, and normal
behavior patterns; (3) Tbr of guineafowl under semi-natural conditions can be elevated in
diffuse, and particularly in direct, solar radiation; we believe that the radiation effect is to
directly warm the head and neck rather than influence blood flow patterns to the head;
(4) postural adjustment scan result in rapid changes in Tbr because of altered heat
exchange with the environment. Key words: helmeted guineafowl, solar radiation,
rain, temperature, behavior, biothermal regulation

548

Woodward, J. K. 2006. Greater sage-grouse (Centrocercus urophasianus) habitat
in central Montana. M.S. Thesis, Montana State University, Bozeman, USA. Notes:
Page 52 - "I also observed larger flocks on my study site when weather was cold.
Flocks ranged from 1 to >150 birds during the two winters. I noticed flocks flushed when
the observer was at a farther distance in the winter than was the case during nesting
and brood rearing season. This suggests grouse would use cover for hiding from
predators in nesting and brood rearing, but in the winter sage-grouse relied on one
another to acknowledge predators and quickly leave the area." Key words: sage-
grouse, temperature, behavior

549

Wright, R. G., R. N. Paisley, and J. F. Kubisiak. 1996. Survival of wild turkey hens
in southwestern Wisconsin. Journal of Wildlife Management 60(2):31 3-320.

Abstract: Successful restoration of wild turkeys (Meleagris gallopavo) to Wisconsin has
stimulated increased interest in hunting and a need for specific management
information. We determined survival and causes of mortality for 224 radiomarked hens
during 1988-94 in Vernon County, Wisconsin under a relatively liberal fall harvest
regime. Annual survival ranged from 43.1 to 66.0 and averaged 52.7% (SE = 4.8).
Seasonal survival averaged 72.2 (SE = 2.9), 81.3 (SE = 3.3) and 89.3% (SE = 2.5)
during recruitment (16 Mar to 14 Jul), post-recruitment (15 Jul to 21 Nov), and winter
(22 Nov to 15 Mar), and was lower (P = 0.001) during recruitment than during winter.
Predation accounted for 71.2% of all deaths and was highest during recruitment.
Localized starvation occurred during a winter with 49 consecutive days of deep snow
and cold temperatures. Cumulative fall hunter densities were about 3.5/km2 of woodland
and the fall harvest rate of radioed hens averaged 7.3%. Turkey densities declined
during the study and more conservative fall harvests may be necessary. Key words:
turkey, snow, temperature, nutrition, mortality

550

Wunz, G. A. and A. H. Hayden. 1975. Winter mortality and supplemental feeding of
turkeys in Pennsylvania. Proceedings of the National Wild Turkey Symposium
3:61-69. Abstract: During a 12-year study in north-central Pennsylvania, wild turkeys
{Meleagris gallopavo silvestris) starved in four winters when extended periods of deep
powder snow prevented their foraging for food. More than half the turkeys at higher
elevations died, even when supplemental food was provided. Turkey populations
usually recovered in one breeding season and appeared more dependent upon the

i fX^*^

245

<£=S& -CSs'- <£~^h> cC^b

previous summer's reproductive success than upon the mildness of the preceding
winter or the number of breeders available. Key words: Eastern wild turkey, snow,
nutrition, population dynamics

551

Yahner, R. H. 1981. Avian winter abundance patterns in farmstead shelterbelts:
weather and temporal effects. Journal of Field Ornithology 52(1):50-56. Notes:
"Use of shelter belts by this species [pheasants] at the Rosemount Station was not
correlated with weather or temporal factors in winter 1979, yet snow depth was
important in winter 1980. I suggest that mean snow depth in 1979 (0.46 m) plus
additional depth resulting from drifting in certain sectors of all shelterbelts, virtually
eliminated access to low-lying sources of food and cover in shelterbelts during the 1979
winter. In contrast, decreased amounts of snow in shelterbelts during winter 1980 (0.10
m) conceivably permitted ring-necked pheasants to more easily obtain food and/or
cover near ground level and in leaf litter with reduced time-energy costs because a
smaller portion of shrubs and herbaceous plants were buried in snow compared to the
previous winter... in years of reduced snow accumulation use of shelterbelts by ring-
necked pheasants may be largely a function of snow depth; when snow depths are
excessive, use of shelterbelts perhaps is contingent on other factors, such as annual
fluctuations in population densities or proximity to alternate food sources. Key words:
ring-necked pheasant, snow, wind, habitat use, feeding, shelter

552

Yeatter, R. E. 1950. Effects of different preincubation temperatures on the
hatchability of pheasant eggs. Science 112(2914):529-530. Notes: "It seems
probable that vulnerability of pheasant embryos to air temperature during the laying
period has an important influence in limiting the southern distribution of pheasants.
Pheasants reported breeding locally in the southern Pacific Coast and Rocky Mountain
regions may be predominantly of southern Asiatic origin, and possibly thus more
tolerant of higher temperatures." Key words: ring-necked pheasant, temperature,
reproduction, species range

553

Yom-Tov, Y., Y. Benjamini, and S. Kark. 2002. Global warming, Bergmann's rule
and body mass: are they related? The chukar partridge (Alectoris chukar) case.
Journal of Zoology (London) 257(4):449-495. Abstract: Using museum specimens
collected in Israel during the second half of the 20th century, no support was found for
the hypothesis that body mass and tarsus length of chukar partridges Alectoris chukar
has changed as a result of global warming. Body mass showed fluctuations during the
year, reaching a maximum in late winter and spring and a minimum in summer.
Bergmann's rule predicts that in warm-blooded animals, races from warm regions will
be smaller than races from colder regions, and a wider explanation states that body size
is positively related to latitude. Because of its topography and varied climate, Israel
provides a unique opportunity to separate partly the effect of latitude from that of
ambient temperature, thus testing if Bergmann's rule is related to latitude or to climatic
variables. We found that body mass (and marginally also tarsus length) declined

246

4

^ te« **

significantly with decreasing latitude in accordance with the wider explanation of
Bergmann's rule, but ambient temperature explained a much smaller fraction of the
variation in body mass than latitude. These results weaken the traditional explanation to
Bergmann's rule that a heat conservation mechanism causes the latitudinal size
variation. Key words: chukar partridge, temperature, morphology, global warming

554

York, D. L. and S. D. Schemnitz. 2003. Home range, habitat use, and diet of
Gould's turkeys, Peloncillo Mountains, New Mexico. Southwestern Naturalist
48(2):231-240. Abstract: Our study documents home range, habitat use, and diet of
Gould's turkey (Meleagris gallopavo mexicana) in the Peloncillo Mountains of New
Mexico and Arizona. Fieldwork began in 1989 and consisted of 2 field seasons from
May to August, with periodic winter and spring forays. The study was conducted during
a drought period with only 58% of average rainfall. Combined annual home range
(mean = 4,385 ha, SE - 1 ,845) of radio-equipped Gould's hens was similar to home
ranges of other wild turkey subspecies that inhabit arid regions of the western United
States. Preferred Gould's turkey habitat consisted of pinyon-juniper woodland with an
abundance of pinyon ricegrass (Piptochaetium fimbriatum). In addition, 3 riparian
habitat types were used disproportionately to their availability. A food habits analysis
showed that the diet of turkey in our study consisted primarily of juniper (Juniperus
deppeana) and manzanits (Arctostaphylos pungens) fruit, though mustard forbs
(Cruciferae spp.) and pinyon ricegrass also were utilized. Periodic drought combined
with livestock grazing in the Peloncillo Mountains most likely is a major factor
determining forage availability and subsequent habitat use by this turkey population. A
combination of natural and anthropogenic factors might result in fluctuations in the
Gould's turkey population. Keywords: Gould's turkey, drought, precipitation,
habitat use, vegetation, diet, population dynamics

555

Young, J. A. 1981. Chukar partridge. Rangelands 3(4):166-168. Notes: Page 168—
"Winter kill is to be periodically expected throughout the northern range of chukars in
North America. Although chukars are very tolerant of cold weather, long periods of deep
snows on feeding slopes can result in starvation." Key words: chukar partridge,
temperature, snow, motality

556

Zablan, M. A., C. E. Braun, and G. C. White. 2003. Estimation of greater sage
grouse survival in North Park, Colorado. Journal of Wildlife Management
67(1 ):1 44-1 54. Abstract: We estimated survival rates of greater sage grouse
(Centrocercus urophasianus) in North Park, Colorado, USA, from band-recovery data of
6,021 birds banded during spring, 1973-1990, with recoveries through 1993. Average
annual adult female survival (S = 0.59, SE + 0.01 1 ) was greater than average adult
male survival (S = 0.37, SE = 0.007), and average subadult (<1 yr old at time of
banding) female survival (S = 0.77, SES = 0.030) was greater than average subadult
male survival (S = 0.63, SE = 0.034). Four weather covariates (spring and winter

y A^ / ... 247

% fv^,^

cgy y§f- <^ d£

precipitation and temperature) did not contribute to predicting annual survival. Key
words: sage grouse, precipitation, temperature, survival prediction

557

Zbinden, N. and M. Salvioni. 2004. Bedeutung der Temperatur in der
fruehen Aufzuchtzeit fuer den Fortpflanzungserfolg des Birkhuhns Tetrao
tetri x auf verschiedenen Hoehenstufen im Tessin, Suedschweiz (The
importance of temperature during the early chick-rearing period for the
reproductive success of black grouse Tetrao tetrix at different altitude
levels in the Ticino, southern Swiss Alps. Der Ornithologische Beobachter
101(4):307-318. Abstract: In their first days of life grouse chicks are very
susceptible to outside temperature because thermoregulation has not fully
developed. From 1981 to 202 the relationship between reproductive success of
black grouse and temperature was studied in the southern Swiss alps. Data
showed that at different altitude levels temperature conditions in different time
periods were particularly important: In northern Ticino (at the highest altitude)
reproductive success depended on the mean daily temperature of five-day
period 41 (20 — 24 July), in the central Ticino that of five-day period 39 (10 — 14
July) and in southern Ticino (at the lowest altitude) temperature of five-day
period 37 (30 June to 4 July). At the altitude levels of the home ranges of Black
grouse hens temperature in these three five-day periods reached a mean of 1 1 .4
tO 11.8 degrees C. In northern Ticino five-day period 41 lies in the warmest
season. In central and southern Ticino temperatures continue to grow after the
five-day period most relevant for reproductive success. A comparison of the
temperature conditions in the southern Swiss alps with those in Finland
indicated different time constraints for alpine and northern birds: in Finland,
before the start of egg-laying only one month with temperatures above 0
degrees C and thus increased assimilation of food plants is available to the
hens, whereas in northern Ticino this period reaches more than two months.
When spring arrives late Finnish hens are not able to build up sufficient reserves
and lay eggs from which weak chicks hatch. On the other hand, temperature
conditions in Finland after hatching are favorable for chick rearing. In the
southern Swiss alps the opposite is the case. Hens have enough time to build up
reserves before egg laying. In cold summers, however, temperature after
hatching is often not high enough to enable a high number of chicks to survive
the first critical days. Key words: black grouse, temperature, mortality

558

Zimmerman, G. S., R. R. Horton, D. R. Dessecker, and R. J. Gutierrez. 2008. New
insight to old hypotheses: ruffed grouse population cycles. Wilson Journal of
Ornithology 120(2):239-247. Abstract: We examined factors hypothesized to influence
ruffed grouse {Bonasa umbellus) population cycles by evaluating 13 a priori models that
represented correlations between spring counts of male ruffed grouse drumming

248 ^

4

H I& +*

displays and these factors. We used AICC to rank the relative ability of these models to
fit the data and used variance components analysis to assess the amount of temporal
process variation in ruffed grouse spring counts explained by the best model. A
hypothesis representing an interaction between winter precipitation and winter
temperature was the top-ranked model. This model indicated that increased
precipitation during cold winters (soft snow cover for roosting) was correlated with
higher grouse population indices, but that increased precipitation during warm winters
(snow crust effect) was correlated with lower spring counts. The highest ranked model
(AICC weight = 0.45), explained only 17% of the temporal process variation. The number
of migrating northern goshawks (Accipiter gentillis), which has been correlated with
grouse cycles in previous studies, does not adequately explain, by itself, the variation in
annual population indices of ruffed grouse. Other factors not considered in our analysis,
such as endogenous mechanisms, parasites, or interactions among factors may also be
important, which suggest that mechanisms mediating the ruffed grouse cycle still
require investigation. Key words: ruffed grouse, precipitation, snow, temperature,
model, population dynamics

559

and R. J. Gutierrez. 2006. The influence of ecological factors on detecting

drumming ruffed grouse. Journal of Wildlife Management 71 (6):1 765-1 772.

Abstract: We surveyed drumming ruffed grouse (Bonasa umbelus) to estimate the
probability of detecting an individual and we used Bayesian model selection to assess
the influence of factors that may affect detection probabilities of drumming grouse. We
found the average probability of detecting a drumming ruffed grouse during a daily
survey was 0.33. The probability of detecting a grouse was most strongly influenced by
the temperature change during a survey and its interaction with temperature at the start
of the survey. Although the best model also included a main effect of temperature at the
start of surveys, this variable did not strongly correlate with detection probabilities.
Model assessment using data collected at other sites indicated that this best model
performed adequately (i.e., positive correlation between observed and predicted values)
but did not explain much of the variation in detection rates. Our results are useful for
understanding the historical drumming index used to assess ruffed grouse populations
and for designing auditory surveys for this important game bird. Key words: ruffed
grouse, census, technique, model, temperature, index, behavior

560

Zwickel, F. C. 1977. Local variations in the time of breeding of female blue grouse.
Condor 79(2): 185-1 91. Abstract: Timing of breeding of female blue grouse
(Dendragapus obscurus) was studied in 2 populations living about -3 km apart. Timing
of breeding varied among years, between populations, and between adults and
yearlings. Variations in time of breeding among years showed some correlation with
annual differences in April weather. Breeding was about 1 wk later on the area that was
higher in elevation, cooler, wetter, had more snowpack and held its snowpack longer in
spring. Most yearling females copulated only after most adult females were beginning to
nest. The period of copulation was up to 10 wk long in adult females but only up to 7 wk
long in yearlings. Most females were on breeding range only about 2-3 wk prior to

jj 249

% f>>>>f*

copulation. While there was a 3-4 wk difference in timing of loss of snow between
areas, there was only a 1 wk difference in timing of breeding. Differences in timing of
breeding between age classes may relate to differences in physiological maturity,
amount of time on the breeding range prior to copulation, harassment of yearlings by
adults or interactions among new recruits (yearlings) themselves. Proximate control of
the reproductive cycle of females appears most subject to local ecological conditions.
The breeding period of males has likely been selected to cover the entire receptive
period of females, including annual, geographical and age-class variations. Keywords:
blue grouse, temperature, snow, precipitation, breeding

561

1967. Some observations of weather and brood behavior in blue grouse.

Journal of Wildlife Management 31(3):563-568. Abstract: Two broods of wild blue
grouse {Dendragapus obscurus fuliginosus) were observed for a total of nearly 27 hours
in 1962, each on two consecutive days, the first day in each case being cold and wet
and the second warmer and drier. These broods, plus another for which some data are
presented, experienced what is generally considered to be inclement weather for young
gallinaceous birds. There was no apparent correlation between periods of brooding and
occurrence of rain. Rain and cold, as described here, did not seem to injure chicks
directly but did increase the amount of time spent in brooding, and thus shortened the
time available for feeding. This too had no apparent effect on the young. It is suggested
that young blue grouse are able to cope with the kinds of conditions noted here. Key
words: blue grouse, precipitation, temperature, behavior, nutrition

562

and J. F. Bendell. 1967. Early mortality and the regulation of numbers in

blue grouse. Canadian Journal of Zoology 45(5):817-851. Notes: In trying to explain
the variations in the annual pattern of mortality, it is impossible to overlook the fact that
the peak of hatch in 1962, the year of poor survival, was later than in other years. The
peak of hatch is, of course, determined prior to, or during, the laying season. No
correlations with this delay and general weather conditions in the spring were found.
This does not mean, however, that weather is not involved, for there may be
interactions between reproduction and weather which were not detected. Key words:
blue grouse, weather, reproduction

250 ^

^

fa te ^

Avian Taxa

Common Name

Scientific Name

American woodcock

Scolopax minor

Attwater's prairie chicken

Tympanuchus cupido attwateri

Band-tailed pigeon

Columba fasciata

Black grouse

Tetrao tetrix

Blue grouse

Dendragopus obscurus

Bobwhite quail

Colinus virginianus

California quail

Callipepla californica

Capercaillie

Tetrao urogallus

Caucasian black grouse

Tetrao mlokosiewiczi

Chukar partridge

Alectoris chukar

Common snipe

Capella gallinago delicata

Columbian sharp-tailed grouse

Tympannuchus phasianellus columbianus

Desert quail

Callipepla gambelii

Eastern wild turkey

Meleagris gallopavo silvestris

Eurasian woodcock

Scolopax rusticola

European woodcock

Scolopax rusticola

Gambel's quail

Callipepla gambelii

Gould's turkey

Meleagris gallopavo mexicana

Gray partridge

Perdix perdix

Greater prairie chicken

Tympanuchus cupido

Greater sage-grouse

Centrocurcus urophasianus

Hazel grouse

Bonasa bonasia

Helmeted guineafowl

Numida meleagris

Himalayan snowcock

Tetraogallus himalayensis

Hungarian partridge

Perdix perdix

Lesser prairie chicken

Tympanuchus pallidicinctus

Merriam's turkey

Meleagris gallopavo merhami

Mikado pheasant

Syrmaticus mikado

Montezuma quail

Cyrtonyx montezumae

Mourning dove

Zenaida macroura

Prairie chicken (Lesser)

Tympanuchus pallidicinctus

Red grouse

Lapopus lagopus scoticus

Red-legged partridge

Alectoris rufa

Ring-necked pheasant

Phasianus colchicus

Rio Grande turkey

Meleagris gallopavo intermedia

Rock ptarmigan

Lagopus muta

Ruffed grouse

Bonasa umbellus

Sage-grouse

Centrocercus urophasianus

Scaled quail

Calipepla squamata

Sharp-tailed grouse

Tympanuchus phasianellus

Sitka blue grouse

Dendragapus obscurus sitkensis

Snow partridge

Tetraogallus caucasicus

Spruce grouse

Dendragapus canadensis

Tibetan eared pheasant

Crossoptilon harmani

White-eared pheasant

Crossoptilon crossoptilon

White-tailed ptarmigan

Lagopus leucurus

Wild turkey

Meleagris gallopavo silvestris

Willow ptarmigan

Lagopus lagopus

Zenaida dove

Zenaida macroura

-X

m *" ^fc 4 ■ j _^

N^

f * *> ^

251

im

<£^>

Michael R. Frisina copyright 1969

Author Index

A

Adamcik, R. S.

336

Adler, P. H.

1

Alatalo, R. V.

297

Aldrich, J. W.

2

Aldridge, C. L.

3,4,5,6,355

Allen, D.

7

Allen, T. J.

106

Andersen, D. E.

329

Andersen, 0.

480

Andersen, R.

132,133

Anderson, S.

200

Anderson, S. H.

218,433,506

Andreev, A. V.

8,9,395

Applegate, R. D.

10

Archibald, H. L.

11

Armbruster, J. W.

12

Arner, D. H.

248

Aulie, A.

13,14

Ault III, J. W.

285

Austin, D. E.

15

B

Back, G. N.

16,300

Bailey, R. W.

513

Bailey, W. D.

196

Baines, D.

17,18,19,341,488

Baker, R. A.

185

Bakken, G. S.

20

Bala, R. P.

203

% ^f<CSr^

253

c£^

Balda, R. P.

497

Ballard, W. B.

21

Barnett, J. K.

22

Barrington, M. R.

16

Bastat, C.

500

Battazzo, A. M.

23

Baumann Jr, D. P.

24

Baumgartner, F. M.

25

Baxter, W. L.

26

Beasom, S. L.

27,28,62

Beck, T. D. 1.

29

Bell, L. A.

30

Bendell.J. F.

31,32,562

Bendell-Young, L. 1.

32

Bennitt, R.

33

Benton, T. G.

255

Benjamini, Y.

553

Bergerud, A. T.

102

Berry, J. D.

34

Besnard, A.

364

Bills, W. E.

511

Binet, F.

119

Bittner, S. L.

106

Blake, C. S.

35

Blanchett, P.

36

Blankenship, D. S.

332

Blankenship, L. H.

67

Bleier, W. J.

219

Blix, A. S.

241,242

Blom, R.

352

Boag, B.

225

Boag, D. A.

474

Boggs, C.

37

Bohl.W. H.

38

Boidot, J. -P.

39

Bonczar, Z.

496

Boos, M.

39

Borgo, J. S.

88

Botsford, L. W.

40

Bourgeois, J.-C.

36

Boughton, R. V.

41

Bousquet, K. R.

42

Boyce, M. S.

355

Bradbury, J. W.

43,518

Brand, C.

318

Brand, C. J.

543

Braun, C. E.

44,45,46,47,48,118,230,231,232,408,498,524,556

Braunisch, V.

49

254

X Mfc***

Brener, A. H. 194

Brennan, L. A. 282

Brenot, J. F. 364

Bridges, A. S. 50

Bridgman, C. L. 51

Brigham, R. M. 6

Britt, T. 48

Brittas, R. 52,221

Brotherson, J. D. 72

Brown, D. E. 53,54,55

Bryant, D. M. 382

Bryant, F. C. 209

Bruce, J. R. 56

Bruggink, J. G. 329

Buehler, D. A. 106

Brunjes IV, J. H. 57

Buczek, T. 431

Burdukov, G. N. 58

Burkepile, N. A. 59

Burnett, L. E. 60

Buss, I. O. 61

C

Cabanac, M. 350

Caffrey, C. 319

Cain, J. R. 62

Caldwell, P. D. 89

Call, M. W. 63,64

Campbell, H. 65,66

Campbell-Kissock, L. 67

Cardona, C. J. 68

Carpenter, J. E. 69

Casazza, M. L. 368

Case, R. M. 70

Cattadori, I. M. 71,225

Causey, M. K. 540

Cedarleaf, J. D. 72

Chamberlailn, M. J. 73,74,543

Charette, J. 75

Choate, T. S. 76

Christiansen, T. 79

Christensen, G. C. 77,78

Church, K. E. 80,501

Churchwell, R. 81

Clanton, R. C. 248

Clark, W. R. 387,411

Clarke, J. A. 82

Clugson, D. A. 302

4>

}X 1& ^

255

*!/>

<£$> c£^

Coates, P. S.

83

Cobb, D. T.

84

Cochran Jr, C. L.

202

Coffey, J. M.

318

Coggins, K. A.

85

Cohen, W. E.

151,187

Connan, R. M.

86

Connelly, J.W.

44,46,59,87,145,253

Conover, M. R.

88

Coon, R. A.

89

Cope, M. G.

90

Copelin, F. F.

91

Cornish, T. E.

355,521

Coulson, J. C.

220

Coup, R. N.

92

Crawford, J. A.

22,382

Crawford, J. E.

93,100,101

Criddle, N.

94

Crissey, W. F.

95

Crowe, T. M.

547

D

Dahlgren, R.

96,97

Dailey, T. V.

73,74,451

Dale, F. H.

98

Dalke, P. D.

99,100,101

David, L. M.

526

Davies, R. G.

102

Davis, J. R.

103

Davison, V. E.

104

DeArment, R.

234

Degner, M. A.

286

DeGraff, L. W.

15

DeMaso, S. J.

184,298,299,409

DeWitt, J. B.

359

Delehanty, D. J.

83

Delvingt, W.

466

Derby Jr., J. V.

359

Dessecker, D. R.

457,558

Devers, P. K.

105,106,361

Dobson, A. P.

225

Dodge, W. E.

516

Doerr, P. D.

84,107,487

Doherty, K. E.

108,329,355,521

Dokk, J.G.

477

Dorney, R. S.

109,110,111

Drobney, R. D.

73,74,451

Drovetskii, S. V.

113,114,115

256

^ 4-Vi^ .

<i< f " rv

**

cCb

Doucette, D. R.

112

Duck, L. G.

116

Dugan, D.

488

Duke, G. E.

117

Dumke, R. T.

158

Duncan, D. A.

465

Duncan, R. M.

544

Dunke, R. T.

389

Dunn, P. 0.

118

Dupuis, A. P.

319

Duriez, 0.

119

Dusek, G. L.

120

Dwyer, T. J.

458

E

Eagle, A. J.

517

Eberhardt, L. L.

311

Edelmann, F.

81

Edminster, F. C.

121

Edwards, J. W.

106

Edwards, R. Y.

413

Edwards, W. R.

122,123

Eiberle, K.

124

Eierl, K.

125

Einarsen, A. S.

126,127

Eiswerth, M. E.

517

Eklund, U.

222

Ellison, L. N.

364

Eng, R. L.

34,128,129

Erikstad, K. E.

130,131,132,133,352,353,480

Errington, P. L.

134

Erwin, M. J.

135

Etter, S. L.

136

Eustace, C. D.

120,137

Evans, C. A.

138

Evans, K. E.

139,140

Evard, J. 0.

141

Ewing, E.

411

F

Faber, W. E.

246

Fallis, A. M.

142

Fames, P. E.

143

Fearer, T. M.

361

Ferkovich, P. E.

66

Ferrand, Y.

119,500

Fields, T. L.

144

Figert, D. E.

106

-* -4-V i *A*

4< 7 i k<± ^

257

£Z^> cCb

Fischer, C. A.

107

Fischer, R. A.

87,145

Flake, L. D.

146,156,274,275,303,432

Flanagan, J. P.

1

Flanders, B. L.

147

Flanders-Wanner, B. L.

148

Fleming, K. K.

149

Formozov, A. N.

150

Forrester, N. D.

151,187

Fouquet, M.

384

Francis, W. J.

152,153,154,155

Freidhoff, S. T.

106

Friday, J. D.

336

Frisbie, M. C.

386

Fischer, R. A.

Fritz, H.

119

Fritzell, E. K.

507,508

Fuglei, E.

376

Fuhlendorf, S. D.

30,185

Fuller, W. A.

253

G

Gabbert, A. E.

156

Galbraith, H.

525

Gallagher, J.

184

Gallagher, J. F.

386

Gallizioli, S.

489

Gates, J. M.

157,389

Gatti, R. C.

158

Gefell, D. J.

159,393

Geissler, P. H.

499

Genelly, R. E.

405

George, R. R.

184,186,298

Gerstell, R.

160

Gessaman, J. A.

379,380,381

Gibson, R. M.

518

Gilbert, W. C.

144

Gibson, R. M.

43

Giesen, K. M.

525

Gill, R. B.

161,162

Giudice, J. H.

407

Giuliano, W. W.

106

Giuliano, W. M.

163

Gjerde, 1.

164

Glover, F. A.

165

Gobeille, J. E.

166

Goddard, A. D.

167

Goldstein, D. L.

168,169

258

\ fakr*^

Goodman, E. D. 404

Gore, H. G. 170

Goudy, W. H. 457

Graham, S. A. 171

Grandjean-Thomsen, A. 172

Grasaas, T. 472

Gratson, M. W. 173

Gray, B.T. 174

Green, R. E. 175,488

Greeley, F. 265

Greene, E. 23

Gregg, L. 176

Gregg, M. A. 177

Griffing, S. 319

Grondahl, C. R. 325

Groshens, E. B. 444

Gross, A. O. 178

Grossman, F. 500

Gruys, R. C. 179

Gullion, G. W. 180

Gunderson, P. 181,182

Guthery, F. S. 151,183,184,185,186,187,202,298,299,409

Gutierrez, R. J. 558,559

H

Halouzka, J. 224

Hamerstrom, F. 189

Hamerstrom Jr, F. N. 134,188,189

Hammond, M. C. 190

Hamr, J. 360

Hanf, J. A. 444

Hannon, S.J. 191,312,438

Harper, C. A. 106

Harper, H. T. 196

Hapr, A. J. 524

Haroldson, B. S. 192

Haroldson, K, J. 193,194,195

Harper, C. A. 106,265

Harper, H. T. 196

Harris, B. K. 66

Harry, B. H. 196

Haukos, D. A. 538

Hawkins, A. D. 197

Hawkins, A. S. 304

Hayden, A. H. 550

Haydon, D. T. 71

Hays, D. 198

Healy, W. H. 199

4<

f * *c* ^

259

,>"x

dL^± <^>

Heath, B.

200

Heath, B. J.

218

Heffelfinger, J. R.

201,202

Hejl, S. J.

203

Helle, P.

296

Heller, V. J.

284

Henderson, C. W.

204

Hendrickson, G. 0.

263

Hennessy, T. E.

205

Herman, C. M.

206

Herman-Brunson, K. M.

207

Hermann, M. F.

208

Hernandez, F.

209

Herzog, P. W.

210

Hesterberg, G.

171

Hewitt, D. G.

211

Hiller, T. L.

185

Hillman, C. N.

212

Hipkiss, T.

222

Hissa, R.

317

Hjeljord, 0.

477

Hjorth, 1.

213

Hobbs, N. T.

525

Hodorff, R. A.

434

Hoffman, D. M.

214,215,216

Hoffman, R. W.

45

Hoglund, N. H.

217,314

Holloran, M. J.

218,506

Homan, H. J.

219

Homeyer, P. G.

263

Hoodless, A. N.

220

Hope, C. E.

142

Hopkins, C. R.

248

Horak, G. J.

401

Hornfeldt, B.

222

Homell-Willebrand, M.

221

Horton, R. R.

558

Host, P.

223

Houston, D. C.

526

Hovi, M.

467

Howard, R.

209

Howell, F. W.

349

Hubalek, Z.

224

Hudson, P. J.

71,225,255,396

Huempfner, R. A.

226

Huffman, R. T.

227

Hungerford, C. R.

228

Hupp, J. W.

229,230,231,232

260

^ 4 v l^

<*- f * *v

**

cC^>

Hurst, G. A.

I

Igo, W. K.

282,331,369

106

Ihejirika, A.

68

Iljinsky, 1. V.

520

Ivacic, D. L.

233

J

Jackson, A. S.

234

Jackson, J. J.

264

Jackson, W. W.

212

Janson, R. G.

535

Jarvis, J. M.

235

Jenkins, D.

236

Jensen, M. S.

322

Johnson, D. E.

238

Johnson, R. A.

239

Johnson, R. E.

82

Jonas, R. J.

239,240

Jorgensen, E.

241,242

Joselyn, G. B.

136

Juoicova, Z.

224

Jurgenson, P. B.

243

K

Kabat, C.

110,244

Kaczor, N. W.

245

Kane, D. F.

246

Kark, S.

553

Kasprzykowski, Z.

247

Kastdalen, L.

532

Keith, L. B.

107

Keller, M.

431

Kennamer, J. E.

248

Keppie, D. M.

249,250,402

Kerr, S. M.

544

Kerwin, M. L.

251

Kessler, F. W.

252

Kienzler, J. M.

253

Kilpatrick, A. M.

319

Kimmel, F. G.

543

Kimmel, R. 0.

192,194,195,246,543

King, R. T.

254

Kirby, A. D.

255

Kirkpatrick, R. L.

106,211

Klaus, S.

256

Klebenow, D. A.

524

Klimstra, W. D.

428

I HfcL***

261

S $fe <Q> <9

Klongan, E. D.

257

Knapton, R. W.

258

Kobriger, J.

259

Koerth, B. H.

409

Kopp, S. D.

151

Korhonen, K.

260,261

Kothmann, H. G.

262

Kozicky, E. L.

263

Kozlik, F. M.

244

Kozlov, V. M.

58

Kramer, L.

319

Krementz, D. G.

264,486

Krohn, W. B.

515

Kubisiak, J. F.

549

Kubisiak, K. F.

425,549

Kuipers, J. L.

218,506

Kuvlesky Jr, W. P.

187

L

Labisky, R. F.

233,265

Lahey, J. F.

268

Lambie, D.

488

Laperriere, A. J.

266

Larsen, J. A.

267,268

Larson, M. A.

269

Latham, R. M.

270,271

Lauckhart, J. B.

272

Lawson, J.

200

Lebreton, J.-D.

500

Lehman, C. P.

146,273,274,275

Lehman, V. W.

275,276

Lehtinen, S. A.

277

Leif, A. P.

146,156

Leite, A.

123

Lennerstedt, 1.

279

Leopold, A.

280

Leopold, A. S.

281

Leopold, B. D.

282,331

Leptich, D. J.

283

Lewis, J. C.

284,285,545

Lewis, R. A.

286

Ligon, J. S.

287

Linberg, M. S.

346,347

Lindstrdm, J.

288,289,297

Linden, H.

289,297

Lindsey, J.

292

Lindzey, F. G.

379,380,381

Linz, G. M.

219

262

% **<CV

>f*

Little, T. W.

253

Litton, G. W.

262

Loneux, M.

291,292

Long, C. R.

290

Longcore, J. R.

293,294,302,515

Lott, D. F.

40

Lowery, D. K.

295

Lu, X.

296

Ludwig, G. X.

297

Lusk, J. J.

184,298,299

Lutz, R. S.

163,329

Lyon, A. G.

218

M

McAdoo, J. K.

16,300

McArthur, E. D.

524

McAtee, W. L.

301

McAuley, D. G.

293,294,302

McCabe, K. F.

303

McCabe, R. A.

303,304

McCafferty, D. J.

526

McClellan, L.

68

McClure, H. E.

305

McCrow, V. P.

306

McDaniel, L. L.

148

McGowan, J. D.

307

Mcintosh, J.

355

Mackenzie, J. M. D.

308

McKinney, L. B.

336

McLean, R.

319

McLean, R. G.

309

MacLeod, M. G.

383

McMillan, 1. 1.

310

McMullen, C. M.

202

MacMullan, R. A.

311

McNeil, J.

312

McNeil, R.

312

Mahan, W. E.

24

Manzer, D. L.

313

Marcstrom, V.

221

Marcstrom, V.

314

Marestrom, V.

481

Marjakangas, A.

315,316,317

Marks, J. S.

318,437

Marks, V. S.

318

Marquiss, M.

342,343

Marra, R. P.

319

Marshall, A. J.

320

4> f i ^rv

**

263

% $fe <Q? <9

Marshall, W. H.

321,322

Martin, D. K.

66

Martin, K.

191,323,458,539

Martinson, R. K.

324,325

Maser, C.

64

Mastrup, S.

40

Matchett, M. R.

355

Matteucci, C.

327

Mattson, K.

394

Mattson, 0. E.

189

Mautz, W. W.

366

Merchant, S. S.

326

Meriggi, A.

327

Meunier, J.

328,329

Middleton, A. D.

330

Mikolaj, J.

123

Miller, D. A.

331

Miller, G. R.

236

Miller, M. R.

332

Mitchell, G. J.

333

Moen, A. N.

140

Moen, P.

14

Moncrieff, R.

488

Montagna, D.

327,334

Moritz, W. E.

335

Morrison, J. A.

285

Morrison, P. R.

436

Morrow, M. E.

336,337

Moss, A.

338

Moss, R.

339,340,341 ,342,343,488,528,529

Mossop, D. H.

344

Moynahan, B. J.

345,346,347,348

Munro, R. E.

499

Mussehl, T. W.

349

Myhre, G.

350

Myhre, K.

350

Myrberget, S.

351,352,353,354,481

N

Naugle, D. E.

355,521

Neave, D. J.

356

Nelson, G. C.

394

Nelson, 0. C.

357

Nenno, E. S.

198

Nestler, R. B.

358,359

Nguyen, L. P.

360

Nolte, K. R.

187

Nomsen, R.

263

264

\ f^tei***

c£^

Norman, G. W.

361

Norman, J. A.

401

Norris, C.

354

Norris, E.

37,354

Northrup, R. D.

362

Norton, M. A.

363

Novak, R.

319

Novoa, C.

364

0

Oakleaf, R. J.

365

Oberlag, D. F.

366

Olinde, M. W.

486

Olding, R. J.

201,202

Olsson, B.

495

Oppelt, E.

329

Osmolovskya, V. 1.

367

Oswald, J.

340,341

Owen Jr, R. B.

458,515

Overrein, 0.

376

Overton, C. T.

368

P

Paisley, R. N.

425,549

Palmer, W. E.

369

Panek, M.

370,371

Parker, G. H.

360

Parmalee, P. W.

372

Parr, R. A.

342,343

Paterni, M. J.

373

Pattee, 0. H.

28

Patten, M. A.

30,374

Patterson, R. L.

375

Pedersen, A. 0.

376

Pedersen, H. Chr.

377,480

Pekins, P. J.

92,366,378,379,380,381 ,463,464

Pelgren, E. C.

382

Pendlebury, C. J.

383

Pendleton, G. W.

293,294

Pepin, D.

384

Pepper, G. W.

385

Perez, R. M.

386

Perkins, A. L.

387

Perry, R. F.

388

Petersen, L. R.

389

Peterson, J. G.

120

Peterson, M. J.

50,184,186,298,390,448

Phalen, P. S.

369

4 fxv***

265

Piao, Z.-J. 391

Pils, C. M. 158

Porkert, J. 392

Porter, W. F. 80,149,393,394,418,419,430

Pospahala, R. S. 490

Potapov, R. L. 395

Potts, G. R. 396

Pradel, R. 500

Prellwitz, D. M. 397

Priest, S. R. 369,398

Prince, H. H. 174,404

Proctor, R. 488

Puckett Jr, W. H. 185

Pullianen, E. 399,400

Purvis, J. R. 156

Pyrah, D. B. 100,101

Q

Queal, L. M. 401

Quinn, N. W. S. 402

Rabe, D. L. 403,404

Radomski, A. A. 209

Rae, S. 529

Raitt, R. J. 405

Rand, A. L. 406

Ranta, E. 289

Ratti, J.T. 81,407

Reebs, S. 112

Reese, K. P. 59,87,145,524

Reeves, W. K. 1

Remington, T. E. 408

Rhodes, W. E. 24

Rice, S. M. 409

Rich, T. 410

Richardson, M. 19

Riggs, M. R. 194,195

Riley, T. Z. 10,387,411

Rintimaki, H. 317

Ritcey, R. 412

Ritcey, R. W. 413

Rivera-Milan, F. F. 414,415

Roach, D. 1

Robel, R. J. 70,416

Roberts, S. D. 417,418,419,420

Robin, J.-P. 39

Robinson, J. D. 421

266

\

H <o **

Rogers, G. E.

45,423,424

Rodgers, R. D.

144

Roersma, S. J.

422

Rohwer, S.

115

Rolley, R. E.

425

Roseberry, J. L.

426,427,428

Rosene, W.

282,429

Rotella, J. J.

42,346

Rothery, P.

528

Rowe, L. D.

62

Rowland, L. 0.

62

Rowland, M. M.

430

Rozycki, A.

431

Rumble, M. A.

146,274,275,432,433,434

Rusch, D. H.

107,501

Ruth, J. M.

435

Ruwet, J. C.

292

Ruxton, G. D.

526

Rybak, A. R.

185

Ryser, F. A.

436

s

Saab, V. A.

437

Saari, L.

496

Saebo, A.

480

Saito, E.

319

Salvioni, M.

557

Sandercock, B. K.

438

Saniga, M.

439

Savage, D. E.

440

Sayre, M. W.

441,516

Schaffner, F. C.

416

Schemnitz, S. D.

138,442,443,554

Schetzinger, W.

447

Schladweiler, P.

129

Schlatterer, E. F.

100,101

Schmidt, P. A.

444

Schmidt Jr, R. K.

47

Schmitz, R. A.

368

Scherzinger, W.

447

Schneegas, E. R.

445

Schroder, J.

447

Schroder, W.

447

Schroeder, M. A.

44,46,446

Schwertner, T. W.

448

Scott, T. D.

449

Seamster, M. H.

106

Sedvarsky, W. D.

450

% ^^^

267

*11'

£2^ <^>

Seller, T. P.

451

Seiskari, P.

455

Seiss, R. S.

369

Selas, V.

452,453,454

Semenov-Tyan-Shanskii, 0. 1.

456

Sepik, G. F.

293,294,302,457,458,486,51 5

Sexson, K.

401

Shaw, J. L.

459

Shaw, N.

524

Sheldon, W. G.

460

Shelford, V. E.

461

Sherfy, M. H.

462,463,464

Sherrod, S. K.

374

Shields, P. W.

465

Shochat, E.

374

Sieux, J.-S.

466

Siltari, H.

297,467

Siivonen, L.

468

Silvy, N. J.

50,337,390,441,448

Sime, C. A.

469

Sisson, L.

470

Slagsvold, T.

471,472

Slater, S. J.

218,506

Smeins, F. E.

50

Smith, A. A.

255

Smith, J. T.

40,473

Smith, L. M.

538

Smith, R. H.

55

Smith, S. G.

185

Smyth, K. E.

474

Snyder, W. D.

475,476

Sole, J. D.

106

Spears, G. S.

409

Spidso, T. K.

353,477

Spike Jr, H. A.

106

St. Onge, S.

36

Stanford, J. A.

478,479

Stanton, D. C.

100,101

Stauffer, D. F.

106,537

Steen, H.

480,481

Steen, J. B.

37,376,481

Steffen, D. E.

106

Stemler, C. L.

332

Stenseth, N. Chr.

481

Steward, P. A.

482

Stemler, C. L.

332

Stinson, D.

198

Stiver, S. J.

483

268

** te «*

G>

Stoddard, H. L.

484

Stokes. A. W.

542

Stokkan, K.-A.

485

Storm, G. L.

89

Straw Jr, J. A.

486

Straw, R.

200

Stribling, H. L.

487

Suchant, R.

49

Summers, R. W.

488

Svihel, M. L.

195

Swank, W. G.

489

Swanson, C. V.

61

Swanson, D. A.

106

Swanson, D. L.

490

Swearingin, R. M.

491

Swenson, J. E.

492,493,494,495,496

Synatzske, D. R.

184

Szaro, R. C.

497

T

Tacha, T. C.

498

Tanaka, J. A.

524

Tapper, S. C.

396

Tautin, J.

499

Tavecchia, G.

500

Taylor, J. S.

501

Tefft, B. C.

106

Teman, W. H.

302

Terrill, H. V.

33

Tester, J. R.

226

Theberge, J. B.

502

Thomas, J. W.

347

Thomas, V. G.

503

Thompson III, F. R.

507,508

Thompson, D. J.

274,275,504

Thompson, D. R.

244,505

Thompson, J. G.

146,524

Thompson, K. M.

506

Thompson, W. L.

509

Tirhi, M.

198

Todd, A. C.

111

Toepfer, J. E.

1,510

Torell, L. A.

524

Toso, S.

327

Towers, J.

250

Trautman, M. B.

511

Tremblay, Y.

119

Trenholm, 1. B.

342,343

I HfcL^

269

r

Tsuji, L. J. S.

512

Tunkkari, P. S.

396,397

u

Uhlig, H. G.

513

Unander, S.

376

U

Van Camp, L.

205

Van der Haegen, W. M.

514,515,516

Van Kooten, G. C.

517

Vasquez, J. D.

209

Vehrencamp, S. L.

43,518

Verner, J.

203

Viitala, J.

467

Vohs, P. A.

387,411

Voronin, R. N.

519

Vysotsky, V. G.

520

W

Wainwright, T. C.

40

Wakeling, F.

432

Wakkinen, W. L.

87

Walker, B. L.

355,521

Walker, J.

348

Wallace, M. C.

21

Wallestad, R.

522,523

Wallestad, R. 0.

48

Wambolt, C. L.

524

Wang, G.

525

Ward, J. M.

526

Ward, J. W.

346

Warner, R. E.

136,527

Warnock, J. E.

136

Warren, P.

19

Watson, A.

236,339,528,529

Watts, R. C.

523,530

Wayne, A. T.

531

Wegge, P.

164,532

Weigand, J. P.

533,534,535

Weinacht, D. P.

490

Welch, B. L.

524

West, G. C.

502

Whitaker, D. M.

106,536,537

White, G. C.

144,148,536

White, L. D.

67

Whiting Jr, R. M.

538

Wickliff, E. L.

511

Wiebe, K. L.

323,539

270

>4*

to &

<£Z^h> c^>

**

Wiley II, E. N.

540

Willebrand, T.

221

Williams, G. R.

541

Williams, H. W.

542

Wilson, W. B.

543

Windingstad, R. M.

544

Wisdom, M. J.

430

Wise, D. R.

86

Wiseman, D. S.

545

Wisinski, C. L.

546

Withers, P. C.

547

Wolfe, C. W.

26

Wolfe, D. H.

374

Woodward, J. K.

548

Worthen, S. D.

72

Wright, B. S.

356

Wright, R. G.

425,549

Wu, X. B.

50

Wunz, G. A.

550

Y

Yahner, R. H.

551

Yeatter, R. E.

461,552

Yee, J. L.

332

Yin, H.

312

Yom-Tov, Y.

553

York, D. L.

554

Young, J. A.

555

Z

Zablan, M. A.

556

Zacchetti, D.

327

Zbinden, N.

558

Zheng, G. M.

296

Zimmerman, G. S.

557,559

Zwickel, F. C.

560,561,562

)">' <o ^k

X i*,,

R. Margaret Frisina copyright 2003

TOPIC INDEX

A

Abundance
Age Ratio
American woodcock

Attwater's prairie chicken
Avian cholera

B

Band-tailed pigeon
Barometric pressure
Behavior

Bioenergetics

*4

124,298,478

110,202,324,372

89,108,176,264,293,294,302,328,329,403,404,457,

460,487,498,499,51 4,51 5,538,540

1

544

368

89,213,328,375,435,451

8,9,11,13,16,17,18,25,33,36,37,43,47,88,89,105,

107,108,112,114,115,116,130,131,132,133,135,

140,145,152,160,164,168,172,175,181,185,188,

210,212,213,215,217,222,224,226,238,240,241,

242,252,253,260,264,277,273,279,285,293,296,

301 ,304,31 1 ,31 5,31 6,321 ,322,328,331 ,344,349,

350,362,373,375,377,378,380,384,385,395,400,

41 6,424,429,443,451 ,462,464,467,470,482,485,

486,491,494,495,499,501,508,510,512,516,518,

528,536,537,545,547,548,559,561

70,92,140,174,232,403,404,471,503

')'>' ^> ^

273

*JJ*

<£^> cCb

Biothermal regulation 160,185,378,383,395,547

Black grouse 17,18,19,115,124,172,213,222,247,288,289,291,

292,297,308,315,316,317,338,367,416,455,467,
477,488,557

Blood parameters 261

Blue grouse 31 ,55,72,203,286,378,379,380,381 ,423,482,560,

561,562

Bobwhite quail 50,54,62,68,70,73,74,104,116,122,151,160,163,

183,184,185,186,187,209,270,276,277,282,298,
299,358,359,372,386,409,426,427,428,429,443,
449,451 ,478,484,490,501 ,51 1 ,545,560

Breeding 560

Breeding success 297

Brood survival 148

c

California quail 40,54,93,1 52,1 54,1 55,206,280,31 0,405,465

Capercaillie 49,124,164,222,279,288,289,317,338,340,341,367,

399,439,447,452,453,454,455,468,472,488,532
Capture mortality 57

Census 25,91 ,1 1 7,202,21 5,226,259,269,284,298,331 ,368,

423,451 ,460,473,491 ,492,499,542,559
Chick survival 6,18,30,42,59,72,126,199,237,275,314,352,353,

370,411,420,504,532,540
Chukar partridge 38,77,78,81,196,349,407,542,553,555

Climate change 225,291 ,292,299,383,51 7

Cloudburst 139

Clouds 112,117,155,188,213,259,269,384,385,423,429,

451,461,470,482,542
Columbian sharp-tailed grouse 318
Color phase 180

Condition 230

Conservation 49

Cover 201

D

Density 111,256

Dew 175,259,303

Dew point 331

Diet 62,64,81 ,1 13,206,208,249,276,408,454,465,554

Disease 31 ,41 ,68,69,71 ,86,96,97,1 09,1 1 1 ,206,225,244,255,

272,308,309,319,343,345,355,358,359,375,391,
396,459,521,544

Distribution 23,1 04,1 77,246,372,498,51 0

Drought 4,6,7,10,35,50,54,59,62,67,77,78,91,94,97,120,122,

134,137,138,170,182,183,191,194,200,215,234,
282,300,309,312,325,326,329,349,358,359,375,
386,437,443,445,448,460,478,484,524,554

274 ^

4>

+>i fc* **

cCi

E

Eastern viral encephalitis

Eastern wild turkey 165,170,174,199,248,361,417,418,425,513

Egg quality 480

El Nino 203

Eurasian woodcock 39,500

European partridge 324

Evapotranspiration 154,282

F

Fecundity 19,40,55,62,71,228,266,272,310,342,364,427,438,

461

Feeding 321,467,541

Fire 87,180,509,524

Flood 84,95,97,103,216,248,264,311,512,543

Fog 333,401,405,460,482

Foraging 37

Frost 63,64,188,236,293,333,505,510,514,515,524

G

Gambel's quail 20,54,65,1 68,1 69,202,204,228,489

Genetics 493

Global warming 553

Gould's turkey 554

Gray partridge 80,81,175,192,194,259,306,327,333,370,371,384,

407,493

Grazing 5,53,67,85,147,170,277,437,445

Greater prairie chicken 147,269,363,450

Greater sage-grouse 546

Grouse 41 ,58,1 91 ,225,272,339,392,395

H

Habitat 345

Habitat condition 262,389

Habitat loss/fragmentation 187

Habitat management 34,185,189,274,283,362,363,370,408,433

Habitat modification 7,21,24,48,91,120,136,147,151,157,170,180,214,

257,335,356,374,455,469,477,479,501

Habitat quality 1 1 6,1 20,1 83,228,426,479,493,51 0

Habitat use 4,21,22,23,25,29,30,34,36,44,45,46,47,48,56,60,63,

64,74,88,90,91,92,93,99,100,108,113,114,115,119,
129,133,151,153,156,157,158,165,167,173,179,
180,181,185,190,191,196,203,210,212,218,226,
229,231 ,249,252,254,278,287,294,296,300,301 ,
303,306,321,322,326,329,335,344,349,357,362,
373,374,378,379,380,385,387,397,401 ,407,41 1 ,
416,424,426,429,433,435,437,440,442,445,450,
455,463,469,483,487,494,495,497,508,509,510,
51 2,51 6,524,534,535,536,545,546,551 ,554

275

J/

^' ^ fc* t*

*•/*

^^ cCb

Hail

60,212,305,357,484,535

Hatching

126

Hazel grouse

9,113,114,124,125,243,256,288,289,367,391,431,

439,466,492,495,496

Heat stress

30,

Heathcock

125

Helmeted guineafow

547

Himalayan snowcock

483

Hoarfrost

392,460

Humidity

2,30,32,71,97,153,224,259,296,374,4501470,480

Hungarian partridge

98,134,190,197,270,304,333,349,533,534

Hunting

141,160,201,253,401,425,513

Hurricane

209

Hydration

1

Ice

204

7,99,1 04,1 21 ,1 78,1 97,205,21 3,254,271 ,280,301 ,

304,307,314,392,429,498,51 1

Incubation

83

Index

32,50,141,143,154,155,159,162,250,273,275,282.

Insects

288,298,312,324,342,386,409,417,420,435,448,

457,475,482,492,499,520,521,525,526,559

1 ,69,94,96,1 31 ,1 75,191 ,224,255,309,334,370,402,

450

Japanese quail

L

Landslide

383

51

Land use

336

Lesser prairie chicken

30,91,326,374

Life history

106

Light

217

M

Manchurian pheasant

150

Merriam's wild turkey

146,149,166,214,216,240,273,274,275,287,303,

366,432,504,509

Metabolism

20,366,380,463,464,507

Microclimate

344,374,379

Microhabitat

74,83,295,374,433

Microtopography

487

Mikado pheasant

51

Model

49,105,148,186,187,193,218,245,288,292,299,345,

361 ,396,404,409,41 5,41 7,425,447,451 ,457,481 ,

500,520,525,558,559

Moisture

403,498

Molt

505

276

Jt
4

>->" fee ^

c^>

Montezuma quail 53,201

Morphology 2,14,402,526,553

Mortality 4,6,7,1 5,1 8,31 ,33,39,41 ,42,55,59,60,68,71 ,76,77,

80,95,97,101,103,104,109,110,111,116,121,122,
123,131,135,138,139,146,156,166,174,176,178,
182,191,197,199,200,205,206,207,209,212,216,
21 9,221 ,233,235,242,244,249,257,258,270,271 ,
276,280,286,293,294,297,301,302,304,305,307,
308,309,31 1 ,312,31 3,327,330,334,337,345,347,
349,352,357,359,360,371 ,374,375,387,388,391 ,
398,401,411,412,415,426,427,429,432,436,439,
441,447,449,450,456,460,476,477,479,480,484,
486,493,498,510,51 1,520,527,531 ,535,540,544,
549,555,557

Mountain quail 54,203

Mourning dove 12,112,205,233,266,285,305,332,337,441,497,505

Movement 8,1 0,23,30,34,38,43,45,47,56,58,63,64,89,91 ,99,

100,108,113,116,118,131,145,146,158,165,179,
1 81 ,21 0,240,264,327,334,343,356,372,401 ,423,
426,435,440,469,501,510

N

NAO 525

Nest survival 142,419

Nesting success 245,346,456

Nutrition 1,9,10,22,35,38,39,44.46,47,48,52,54,66,77,78,85,

91,94,99,102,119,123,131,132,133,134,141,146,
152,155,174,175,176,177,190,191,193,195,197,
200,201,207,210,211,221,228,229,231,235,236,
237,240,242,246,257,261 ,264,270,271 ,272,276,
280,281 ,285,287,290,293,301 ,304,307,308,31 0,
31 8,342,351 ,352,357,358,359,363,370,375,377,
381 ,389,392,394,399,402,403,404,408,41 5,452,
453,457,471,479,480,484,486,487,498,502,504,
511,514,515,520,524,549,550,561

Palmer Drought Severity Index 448

Parasites 41 ,68,71 ,86,1 09,1 1 1 ,225,343,375,391 ,396

Partridge 330,468

Pesticide 312

Photoperiod 328

Pollution 392

Population dynamics 5,7,15,18,21,41,50,53,65,71,77,79,94,96,101,109,1

20,128,132,137,159,162,163,174,176,194,198,202,
21 5,234,243,244,247,263,268,276,281 ,289,291 ,
292,297,299,308,31 2,325,334,338,339,340,341 ,
343,356,367,386,390,393,394,396,409,410,414,
41 8,422,425,443,447,450,454,468,471 ,498,500,

f ->- ^ ^

S #> ^ <9

51 7,525,528,529,534,541 ,550,554,558

Prairie chicken 1,10,54,336,390,401,461,510

Precipitation 2,3,4,5,6,7,17,18,21,222,26,28,35,40,41,42,48,50,

51 ,53,54,55,57,59,60,61 ,62,63,64,65,66,69,71 ,72,
77,79,85,86,91,94,95,97,99,108,109,110,115,120
122,124,125,126,127,128,132,133,134,137,139,
144,145,147,148,149,152,154,155,157,161,162,
163,166,170,1771,172,175,181,183,184,188,191,
194,196,197,199,200,201,202,206,207,213,214,
21 5,21 8,221 ,222,224,227,228,234,235,240,245,
248,251 ,252,255,262,263,264,266,271 ,273,274,
275,276,277,281 ,282,286,287,289,291 ,292,295,
298,299,301,303,304,305,308,309,310,311,311,
313,324,325,326,330,332,333,335,336,337,338,
340,341 ,343,345,346,361 ,363,364,368,369,370,
372,375,385,386,388,390,391 ,397,398,401 ,402,
404,405,407,409,410,411,412,414,415,416,417,
41 8,41 9,420,421 ,422,423,425,428,430,434,435,
439,440,446,447,448,450,453,456,460,461,465,
469,472,473,474,475,476,478,480,481,482,484,
486,488,489,491 ,496,497,499,500,501 ,502,504,
506,509,514,520,522,523,524,527,530,532,533,
534,535,536,538,542,544,545,547,554,556,558,
560,561

Predation 7,18,21,54,80,83,88,102,105,119,130,156,173,176,

191 ,209,227,237,244,264,271 ,274,301 ,314,31 5,
326,344,358,359,363,374,389,398,412,432,439,
477,481,488,532

Productivity 4,7,18,24,28,51,61,66,77,78,110,134,137,138,144,

148,152,154,161,162,167,176,177,181,183,184,
187,202,220,222,227,235,240,244,248,277,303,
306,324,332,333,338,351 ,357,361 ,364,372,383,
388,397,405,415,420,428,430,434,448,453,472,
474,475,478,481,532

Ptarmigan 124,125,150,323,485

Q

Quail 67,197,280

R

Range condition 235

Range modification 200

Recruitment 4,40,72,361 ,425,457,543

Red grouse 71,36,236,255,308,342,343,396,406,459,528

Red-legged partridge 175,384

Re-nesting 277

Reproduction 3,6,12,14,23,24,26,35,53,72,82,84,85,87,97,109,

123,127,149,153,155,157,187,212,220,221,223,
227,236,239,250,251 ,252,256,273,279,281 ,286,

278 ^

K fe, >&

290,295,302,314,323,326,333,351,352,369,375,
376,386,389,404,405,406,41 3,41 7,41 8,421 ,422,
429,431 ,438,446,452,51 5,524,534,538,539,543,
552,562

Ring-necked pheasant 7,33,61,80,96,97,99,123,126,127,135,141,153,156,

158,171,219,244,252,257,263,283,301,306,311,
325,349,387,388,389,41 1 ,435,436,468,475,51 1 ,
527,535,551,552

Rio Grande turkey 28,57,146,170,227,262,448

Rock ptarmigan 364,376,502,528,529

Ruffed grouse 2,11,20,32,36,41,94,95,102,105,106,107,109,110,

111,121 ,1 42,1 80,21 1 ,226,237,238,254,268,278,
290,307,321 ,324,356,41 3,503,507,508,51 1 ,536,
537,559

s

Sage-grouse 3,4,5,6,1 6,22,23,29,34,35,43,44,46,48,56,59,60,63,

64,69,79,83,85,87,100,101,118,120,128,129,137,
145,157,161,162,177,198,200,218,229,230,231,
232,235,245,251 ,300,309,31 2,335,345,346,347,
355,357,375,397,408,41 0,421 ,424,430,440,445,
446,462,463,464,469,473,493,506,517,518,521,
522,523,524,530,548,556

Scaled quail 50,54,65,66,138,163,187,204,442,443

Severe weather 15,103,116,123,156,230,286,347,428,466,503,539

Severe winter 1,80,334,352

Sex ratio 110,111,176,222

Sharp-tailed grouse 25,42,54,88,90,94,139,140,147,148,167,173,178,

181,182,188,212,259,307,313,322,324,349,362,
363,374,385,412,422,437,470,494,503,512

Shelter 466,551

Sleet 161,287,314,357

Siberian spruce grouse 8

Snow 3,7,8,9,1 6,25,29,34,36,38,44,45,46,47,49,52,53,56,

58,63,64,76,82,90,97,99,100,101,102,104,106,107,
109,110,111,113,118,121,122,129,141,143,146,
150,156,157,158,161,165,173,174,176,177,178,
179,180,190,192,197,200,203,205,210,211,212,
213,214,216,217,219,222,226,229,231,232,238,
239,240,246,247,249,254,257,258,260,271,278,
280,283,284,285,287,289,290,292,293,294,300,
301,303,304,313,314,315,316,317,318,321,322,
323,335,344,345,349,351,354,358,360,362,364,
371 ,376,387,389,392,394,395,399,407,408,41 6,
424,426,427,428,429,439,446,449,453,454,460,
468,472,476,477,478,479,482,483,485,491,494,
495,497,499,508,510,511,512,514,524,534,535,
536,537,545,549,550,551,555,558,560

Snow condition 36,47,322

* X>w i . ^ 279

% ¥*(*?***

$fc ?«jfe <Q> Q

Snow partridge 150

Solar radiation 8,25,33,38,41 ,60,83,91 ,95,1 1 7,1 55,223,253,278,

303.330.333.373.379.384.385.424.441 .455.461 ,
508,510,526,547

Soil 41,62,67,77,108,190,224,282,403,424,510,514,529,

546
Soil moisture 28,91,155

Spruce grouse 32,208,249,250,373,402,406,474,503

Stress 476

Sun spots 376

Survival prediction 556

T

Technique 20,1 92,285,460,461 ,482,492,559

Temperature 3,5,6,7,8,9,13,14,16,18,20,23,25,26,30,32,33,34,36,

37,39.40,41 ,42.43,51 ,53,56,60,61 ,66,69,70,71 ,72,
73,77,86,91 ,92,94,95,97,99,1 02,1 04,1 05,1 07,1 09,
110,111,112,114,115,119,121,124,125,126,127,
128,130,131,132,133,137,138,139,140,141,142,
143,146,147,148,149,150,151,152,153,155,156,
160,162,164,168,169,171,172,174,175,176,180,
181,182,184,185,187,190,191,193,195,196,197,
199,202,204,205,207,208,212,219,220,221,222,
223,224,225,226,231 ,232,233,235,238,240,241 ,
242,251,252,253,254,255,256,257,258,259,260,
261 ,263,264,268,270,271 ,273,278,279,280,282,
284,285,286,287,289,290,291,292,293,294,296,
297,298,299,300,301,303,304,305,307,308,309,
311,312,319,321,323,324,325,332,333,334,335,
341 ,345,349,350,351 ,353,355,356,357,358,359,
361 ,362,363,364,367,370,371 ,372,373,374,377,
378,380,383,385,387,388,390,395,397,399,400,
401 ,402,404,41 0,41 1 ,41 3,41 6,41 7,419,420,422,
425,430,431 ,433,436,437,438,440,441 ,442,446,

447.449.450.451 .453.456.457.459.460.461 .462,
464,468,470,471 ,472,474,475,476,478,479,481 ,
484,485,486,490,491,492,495,496,498,499,500,
501,502,504,505,506,507,508,510,511,513,515,
51 8,520,521 ,522,527,528,529,531 ,532,533,534,
535,536,537,538,542,544,545,547,548,549,552,
553,555,556,557,558,559,560,561

Thermal cover 487

Thermoregulation 1 3,14,30,37,43,73,74,1 12,11 9,1 40,1 68,1 69,1 80,

191,195,241,260,317,350,366,379,380,381,392,
436,464,485,490,495,502,526,537

Thundershower

Thunderstorm 199,440

Tibetan eared pheasant 296

Topography 41,46,88,349,362

280 ^

N^

*'>" fee ^

Turkey

U

Upland birds

V

Vegetation

w

Weather

Weather catastrophes
Weather extremes
Weather patterns
West Nile Virus
White-eared pheasant
White-tailed ptarmigan
White-tipped dove
Willow grouse

Willow ptarmigan

Wind

Wind speed
Winter weather
Woodcock
Wood grouse

z

Zenaida dove

15,21,24,26,84,92,103,159,183,193,195,239,246,
253,270,271 ,284,295,331 ,360,369,393,394,398,
419,420,433,434,491,516,543,544,549,550

348

4,5,6,10,21,22,23,34,35,41,44,46,47,52,53,54,55,

56,59,63,64,67,73,74,77,79,81,85,91,92,93,102,

105,118,120,127,129,140,145,147,152,155,157,

167,177,181,185,201,206,207,208,210,211,212,

215,218,227,228,229,231,234,235,236,240,250,

254,272,274,276,278,281,283,295,300,303,306,

31 8,322,326,335,341 ,344,347,351 ,358,359,362,

363,370,378,397,407,408,41 2,41 8,422,423,424,

430,437,440,442,445,454,463,465,466,469,475,

476,479,480,508,509,51 0,51 3,522,524,545,546,554

31,81,83,93,159,198,237,272,302,306,339,348,393,

396,406,432,519,540,562

186

189,541

18,96

319,355

296

45,47,76,82,21 0,438,525,539

498

37,52,130,131,132,133,217,221,260,261,314,350,

352,353,354,399,400,452,468,471 ,51 9,526

13,14,179,223,241,242,344,351,377,438,480,481,

539

7,11,12,17,20,23,24,30,38,45,46,47,63,88,91,92,

104,107,112,140,146,169,172,180,212,213,214,

215,224,232,240,252,253,257,258,259,269,270,

271 ,286,287,300,301 ,304,305,328,331 ,337,344,

357,362,373,374,375,378,379,384,392,407,415,

423,429,435,441,449,470,473,492,495,499,507,

508,510,526,527,542,551

451,462,463

327

117,1 1 9,220,452,486,520,531

456

414,415

4>

^>i i&k ^<

281

500 copies of this public document were published
at an estimated cost of $12.09 per copy, for a total
cost oi^ $6,045.00, which includes $6,045.00 for
printing and $0.00 for distribution.