EXPERIMENTAL STUDIES OF THE EFFECTS OF GRAZING BY
CAPTIVE GOSLINGS OF THE LESSER SNOW GOOSE
ON THE VEGETATION OF A SUBARCTIC SALT MARSH
by
DAVID S. HIK
DEPARTMENT OF BOTANY
A thesis submitted in conformity with
the requirements for the degree of
Master of Science
in the University of Toronto
May 1988
ACKNOWLEDGEMENTS
This is not the sort of project that I could have
taken on single-handed and I am indebted to many people
for their help over the past two years. I am particularly
grateful to Prof. Bob Jefferies for his guidance and
support, for sharing his insights about pattern and
process in grazing ecosystems, and for teaching me how to
hurry up and wait. Dr. Roger Ruess introduced me to the
world of microbes and provided helpful advice on many
aspects of this research. Thanks Bwana!1
Excellent field assistance was provided by Janet
Curran, Joan Montagnes, Tony lacobelli, Alison Jefferies,
Paul McKuniak, Evan Cooch and Martin Morgan. I extend
special thanks to Helen Sadul, Peter Kotanen, and Iris
Frey for their help, friendship and support throughout
four month field seasons, and in Toronto. Larry Flanagan
answered many of my questions about plant physiology. I
would be remiss not to acknowledege my mob of sixty
goslings; without their "cooperation" this project would
have not been possible.
Many of the staff members and students of La Perouse
Bay and the Churchill Northern Studies Center provided
logistical support and other help when required. I would
particularly like to thank Dr. Fred Cooke for providing me
with the opportunity to work with snow geese in the first
place, and Dr. Rocky Rockwell for his generous assistance
and for his efforts in maintaining the productive and
relatively sane atmosphere at LPB during the past four
summers.
11
Drs. Spencer Barrett and Sara Shettleworth provided
many helpful comments during committee meetings. Dr. Dolf
Harmsen shared his insights about the interactions between
geese and plant communities at La Perouse Bay. Thank you.
Thanks also to Valerie Anderson who provided many cups of
tea.
Funding for this research was provided by the
Canadian Wildlife Service (Environment Canada), the
Department of Indian and Northern Affairs (Canada), an
Orville Erickson Memorial Scholarship from the Canadian
Wildlife Federation, and NSERC grants to R.L. Jefferies.
Finally, I thank my parents who encouraged me
constantly.
Ill
SUMMARY
1. The effects of grazing by captive goslings of the
Lesser Snow Goose on the vegetation of the La Perouse Bay
salt marsh were investigated. Goslings fed on swards of
Puccinellia phryqanodes; net above-ground primary
production (NAPP) and forage quality (amounts of nitrogen
and carbon) were measured over the course of the season.
2. Grazing early in the season resulted in increased
NAPP of swards of Puccinellia, as predicted by the
herbivore-optimization model. The greatest enhancement of
production occurred at low to moderate levels of grazing
compared to production in ungrazed swards. At higher
levels of grazing production was reduced. Enhancement of
NAPP following grazing was detected only early in the
season; the ability of the plants to recover from the
effects of grazing decreased later in the season.
3. Repeated grazing episodes on the same swards of
Puccinellia over the course of the summer demonstrated
that swards could maintain high rates of production, and
high nitrogen content of shoots even when they were grazed
late into the season. Swards of Puccinellia grazed at
approximately 24 day intervals throughout the summer and
swards grazed at approximately 12 day intervals on four
occasions had the largest increase in NAPP compared to
ungrazed swards. Increases in NAPP were achieved over a
wide range of grazing periods and intervals between
grazing episodes.
IV
4. NAPP was 30 to 40% less in swards of Puccinellia where
faeces were removed compared to corresponding values for
swards where the faeces remained. Faeces provided a
source of soluble nitrogen that could be readily taken up
by plants. However, the amount of nitrogen contributed by
the faeces to the total amount of nitrogen accumulated in
the above-ground biomass of grazed swards of Puccinellia
was less than 10%. Up to 44% of total nitrogen
accumulated in the above-ground biomass of grazed swards
could not be accounted for.
5. Nitrogen mineralisation processes which occur in
goose faeces may be important in maintaining the nitrogen
balance of the salt marsh. Rates of microbial respiration
and the net mineralization of nitrogen in fresh faeces
from gosings grazing Carex subspathecea were higher than
corresponding values for faeces derived from Puccinellia
phryqanodes and Calamagrostis deschampsiodes. However
the mineralisation efficiency (net mineralisation rate /
respiration rate) was similar in faeces derived from Carex
and Puccinellia. The ratios were considerably higher than
the corresponding rates in faeces derived from
Calamaqrostis. The digestibility of Calamagrostis shoots
was considerably lower than that of Carex and Puccinellia
shoots based on analysis of acid-detergent fibre.
6. The interactive and dynamic processes of nitrogen
cycling between plants, geese, sediments and microbes lead
to enhanced primary production in this grazed system. A
suite of interactions exist which extend the influence of
plant-herbivore interactions beyond the immediate
participants, and involve all trophic levels in this
grazing system.
VI
TABLE OF CONTENTS
PAGE
ACKNOWLEDGEMENTS i
SUMMARY iii
LIST OF TABLES xi
LIST OF FIGURES xiv
LIST OF ABBREVIATIONS xvii
CHAPTER 1: GENERAL INTRODUCTION - THE PROCESS OF
HERBIVORY 1
1.1 Introduction to herbivory 1
1.2 Positive feedbacks and nutrient cycling in
grazing ecosystems 4
1.3 Responses of plants to grazing 7
1.3.1 Plant defenses to the effects of herbivory 9
1.2.3 Productivity and growth rate of plants in
relation to herbivory 12
1.4 Compensatory responses 13
1.4.1 Intrinsic mechanisms 15
1.4.1.1 Photosynthesis 15
1.4.1.2 Resource Allocation 16
1.4.1.3 Morphological Plasticity 18
1.4.2 Extrinsic mechanisms 19
1.4.2.1 Water-use efficency 19
1.4.2.2 Salivatory Hormones 20
1.4.2.3 Fertilization 21
1.5 Grazing and below-ground biomass
and production 23
1.6 Principle Objectives 26
Vll
CHAPTER 2: DESCRIPTION OF STUDY SITE AND SPECIES 29
2.1 Description and history of the La Perouse
Bay grazing system 29
2.2 Biology of Puccinellia phyrganodes 37
2.3 Biology of Lesser Snow Geese 40
2.3.1 Distribution 40
2.3.2 Growth and reproduction 41
2.3.3 Foraging behavoir and digestive physiology 43
2.4 The use of captive goslings in experimental
studies 46
CHAPTER 3: WITHIN SEASON INCREASES IN THE NET ABOVE-
GROUND PRIMARY PRODUCTION OF A SALT MARSH
FORAGE GRASS: A TEST OF THE HERBIVORE
OPTIMIZATION MODEL 50
3^  1 Introduction 50
3.2 Methods 53
3.2.1 Study Site 53
3.2.2 Experimental trials 54
3.2.3 Sampling and Data Analysis 56
3.3 Results 58
3.3.1 Changes in above-ground biomass following
grazing 58
3.3.2 Increase in NAPP of swards of Puccinellia
following grazing 63
3.3.3 Forage quality and nitrogen content of shoots ... 65
3.3.4 Effect of faeces on the growth of swards of
Puccinellia following grazing 70
Vlll
3.3.5 Effect of faeces on NAPP 73
3.3.6 Nitrogen content of above-ground biomass
following grazing in the presence and absence
of faeces 76
3.3.7 Accounting for nitrogen accumulated in shoots ... 80
3.4 Discussion 85
CHAPTER 4: NET ABOVE-GROUND PRIMARY PRODUCTION OF
PUCCINELLIA PHRYGANODES IN RESPONSE
TO MULTIPLE GRAZING EPISODES BY LESSER
SNOW GEESE: THE IMPORTANCE OF THE TIMING
OF GRAZING DURING THE SUMMER 91
4.1 Introduction 91
4.2 Methods 94
4.2.1 Study Site 94
4.2.2 Experimental trials 95
4.2.3 Sampling and Data Analysis 98
4.3 Results 99
4.3.1 Changes in NAPP of swards of Puccinellia
phryganodes in response to grazing 99
4.3.2 Amounts of nitrogen and carbon in above-
ground biomass of Puccinellia phryganodes 104
4.3.3 Nitrogen accumulated in above-ground biomass
of Puccinellia phryganodes 106
4.4 Discussion 110
ix
CHAPTER 5: MICROBIAL NITROGEN MINERALISATION PROCESSES
IN A GRAZED SALT MARSH - POSITIVE FEEDBACKS
BETWEEN SNOW GOOSE FORAGING AND NITROGEN
CYCLING 121
5.1 Introduction 121
5.2 Methods 124
5.2.1 Study Site 124
5.2.2 Experimental trials 125
5.2.3 Measurement of rates of microbial respiration
and rates of nitrogen mineralization in fresh
faeces 126
5.2.4 Determination of soluble nitrogen 127
5.2.5 Carbon and nitrogen determinations 128
5.2.6 Neutron activation analysis 128
5.2.7 Acid detergent fibre analysis 129
5.2.8 Statistical analysis of data 130
5.3 Results 130
5.3.1 Feeding behaviour of goslings 130
5.3.2 Chemical composition of plants of Puccinellia
phryqanodes, Carex subspathecea and
Calamaqrostis deschampsiodes 132
5.3.3 Chemical composition of faeces from goslings
grazing on Puccinellia phryganodes, Carex
subspathecea and Calamagrostis deschampsiodes ... 135
5.3.4 Microbial respiration and nitrogen
mineralisation during the incubation of faeces .. 136
5.3.5 Digestibility of the three forage types 143
5.4 Discussion 143
XCHAPTER 6: GENERAL DISCUSSION - HERBIVORY AS AN
ECOSYSTEM PROCESS AND COEVOLUTION IN
GRAZING SYSTEMS 151
6.1 Overview of results 151
6.2 The limitations of perturbation experiments 152
6. 3 Ecosystems as dynamic units 154
6.4 Ecosystem responses to herbivory and disturbance .. 156
6.5 Nutrient cycling in grazing ecosystems 160
6.6 Arctic breeding geese and community structure 163
6.7 Coevolution in grazing systems 165
LITERATURE CITED , , , . , , „ , , , , . 169
xi
LIST OF TABLES
PAGE
TABLE 2-1. Summary of precipitation (rainfall and
snowfall water equivalent), bright sunshine hours, and
mean high and low temperatures for May, June, July, August
and September in 1986 and 1987, and yearly totals 32
TABLE 3-1. Dates on which grazing experiments were
conducted at La Perouse Bay, Manitoba and mean gosling
weights and number of goslings grazed per plot for each
experiment
TABLE 3-2. Summary of results of a two-way ANOVA for
increases in above-ground biomass in Experiments 2, 3, 4,
and 5 (Fig. 3-2)
TABLE 3-3. Cumulative net above-ground primary production
(g DWT * m"1 ) for each harvest of experimental plots in
Experiments 1 to 5
TABLE 3-4. Summary of results of a 2-way ANOVA of amounts
of nitrogen expressed as a percentage (%N) of the dry
weight of above-ground biomass in Experiments 1 to 5 (Fig.
3-3)
TABLE 3-5a. Summary of results of 3-way and 2-way ANOVA
of changes in above-ground biomass of Puccinellia
phryqanodes following grazing in Experiments 6 and 7 (Fig.
3-5)
TABLE 3-5b. Summary of results of 3-way and 2-way ANOVA
of changes in above-ground biomass of Puccinellia
phryqanodes following grazing in Experiments 8 and 9
55
62
64
67
72
75
Xll
TABLE 3-6. Cumulative net above-ground primary production
(g DWT * m"* ) for each harvest of grazed plots in
Experiments 6 and 7, and 8 and 9 77
TABLE 3-7. Summary of results of 3-way and 2-way ANOVA of
amounts of nitrogen expressed as a percentage of the dry
weight of above-ground biomass of Puccinellia phryganodes
for Experiments 6 and 7
TABLE 3-8. Summary of net amounts of nitrogen accumulated
in above ground biomass of swards of Puccinellia
phryganodes in Experiments 6 and 7 (Fig. 3-8) and
calculation of the proportion of nitrogen available from a
number of sources to account for the total nitrogen
accumulated after 60 days of regrowth
TABLE 4-1. Days on which experimental plots containing
swards of Puccinellia phryganodes were grazed and/or
sampled during 1987 at La Perouse Bay, Manitoba, on a 12-
day cycle, 24-day cycle and late season only grazing
schedule. The mean weight of goslings (g), length of the
grazing bout, and number of goslings grazed per plot is
also shown .......
79
84
97
TABLE 4-2. Amount of above-ground biomass of swards of
Puccinellia phryganodes consumed by goslings of the Lesser
Snow Goose in experimental plots in 1987 103
TABLE 4-3. Amount of nitrogen (g N * m~* ) in faeces of
goslings of Lesser Snow Geese deposited on grazed swards
of Puccinellia phryganodes in experimental plots in 1987.. 109
xiii
TABLE 5-1. Gosling weight (a), pecking rate (b) and
defecation rate (c) of goslings of the Lesser Snow Goose
feeding on three forage types (Puccinellia phryganodes,
Carex subspathacea and Calamagrostis deschampsiodes), in
the three feeding experiments , 131
TABLE 5-2. Elemental composition of the above-ground
biomass of Puccinellia phryganodes, Carex subspathacea and
Calamagrostis deschampsiodes forage consumed by goslings
in the three grazing experiments 133
TABLE 5-3. Elemental composition of faeces of goslings
grazing on Puccinellia phryganodes, Carex subspathacea and
Calamagrostis deschampsiodes forage in the three grazing
experiments, and amounts of H20 and NH£ -N, and pH of
fresh faeces 137
TABLE 5-4. Microbial respiration between 0 and 12 hours,
and 0 and 24 hours, net nitrogen mineralization between 0
and 24 hours, and mineralization efficiency
(mineralization/respiration) of faeces of goslings grazing
on Puccinellia phryganodes, Carex subspathacea and
Calamagrostis deschampsiodes forage in the three grazing
experiments 141
TABLE 5-5. Digestibility of Puccinellia phryganodes,
Carex subspathacea and Calamagrostis deschampsiodes forage
based on estimates of the ratio of acid detergent fibre .. 144
XIV
LIST OF FIGURES
PAGE
FIG. 1-1. Three possible effects of herbivory on plant
growth (the herbivore optimization model) 14
FIG. 2-1. Map of of Hudson Bay showing the location of La
Perouse Bay, Manitoba 30
FIG. 2-2. Daily maximum and minimum temperatures at
Churchill, Manitoba during the summer of 1986 (a) and 1987
(b) 31
FIG. 2-3. Diagram of the growth habit of of Puccinellia
phryganodes (Trin.). Scribn. and Merr. The positioning of
an axillary shoot is magnified 39
FIG. 2-4. Growth of captive goslings of the Lesser Snow
Goose at La Perouse Bay in 1987. Increases in mean weight
(ing) following hatch (23 June 1987) 48
FIG. 2-5. Comparison of pecking rate of wild and captive
goslings at La Perouse Bay 49
FIG. 3-1. Increase in above-ground biomass of
Puccinellia phryqanodes following grazing on 23 June, 1986
by goslings of the Lesser Snow Goose in Experiment 1 (mean
values of 4 plots) 59
FIG. 3-2. Incease in above-ground biomass of Puccinellia
phryganodes following grazing by goslings in Experiment 2,
2 July 1986 (a); Experiment 3, 13 July 1986 (b) ;
experiment 4, 1 August 1986 (c); Experiment 5, 10 August
1986 (d) 61
XV
FIG. 3-3. Total nitrogen and carbon content as a
percentage of the dry weight of above-ground biomass of
Puccinellia phryganodes following grazing by goslings in
Experiments 1 (a) , 2 (b) , 3 (c) , 4 (d) , and 5 (e)
FIG. 3-4. Net amount of nitrogen (g * m"a ) accumulated
in the regrowth of above-ground biomass of swards of
Puccinellia phryganodes following grazing in Experiment 1
(a); Experiment 2 (b); Experiment 3 (c); Experiment 4 (d);
and Experiment 5 (e)
FIG. 3-5. Increase in above-ground biomass of
Puccinellia phryganodes following grazing by goslings of
the Lesser Snow Goose in Experiment 6 with faeces
remaining (a), and Experiment 7 with faeces removed (b) ..
FIG. 3-6. Increase in above-ground biomass of
Puccinellia phryganodes following grazing by goslings of
the Lesser Snow Goose in Experiment 8 with faeces
remaining (a), and Experiment 9 with faeces removed (b) ..
FIG. 3-7. Total nitrogen and carbon content as a
percentage of the dry weight of above-ground biomass of
Puccinellia phryganodes following grazing by goslings in
Experiments 6 (a,c) and 7 (b,d)
FIG. 3-8. Net amounts of nitrogen (g * m~*) accumulated
in the regrowth of biomass of swards of Puccinellia
phryganodes following grazing in Experiment 6 (a), and
Experiment 7 (b) ,
66
69
71
74
78
81
XVI
FIG. 3-9. Relationship between length of the grazing
period and the amount of above-ground biomass removed as a
percentage of the initial amount present (% offtake) (a).
Relationship between % offtake and the amount of
nitrogen present in faeces (mg N * m~a) deposited on plots
(b) 83
FIG. 4-1. Cumulative net above-ground primary production
of swards of Puccinellia phryganodes grazed by goslings of
the Lesser Snow Goose for each treatment during the summer
of 1987 101
FIG. 4-2. Total nitrogen (a) and carbon (b) content as a
percentage of the dry weight of above-ground biomass of
Puccinellia phryganodes during the summer of 1987 105
FIG. 4-3. Net amounts of nitrogen (g N * m"1 ) accumulated
in the regrowth of above-ground biomass of swards of
Puccinellia phryganodes during the summer of 1987 108
FIG. 4-4. Responses of NAPP to increases in grazing
intensity and the changes in the interval between
successive grazing bouts 117
XVI1
LIST OF ABBREVIATIONS
ANOVA:
NAPP:
Puccinellia:
analysis of variance
net above-ground primary production
Puccinellia phyrganodes
CHAPTER 1: GENERAL INTRODUCTION - THE PROCESS
OF HERBIVORY
1.1 Introduction to herbivory
Herbivory has been pervasive in determining the
structure and dynamics of plant and animal communities,
populations and individuals (Crawley 1983; Howe and
Westley 1988). The effects of herbivory are evident at
all levels of biological organization. Herbivory affects
primary and secondary production, energy flow, nutrient
cycling, and the structure and composition of plant
communities (Harper 1969, 1977; Crawley 1983; Vitosek
1984). A suite of processes exist, which extend the
influence of plant-herbivore interactions beyond the
immediate participants to other members of the community
(see Owen-Smith 1987; Ruess and McNaughton 1987). The
effects of the interactions are evident at both
evolutionary and ecological time scales.
It is estimated that at least 10% of the net above-
ground primary production of terrestrial ecosystems is
consumed by vertebrate and invertebrate herbivores.
(Odum, Connell and Davenport 1962; Teal 1962; Chew and
Chew 1970; Chew 1974; Crawley 1983). In some communities
(eg. grazing systems) this proportion may be considerably
higher, approaching 100% (Dyer et al. 1982; Cargill and
Jefferies 1984b; Coley, Bryant and Chapin 1985;
McNaughton 1985). There is a paucity of research on
below-ground herbivory, but estimates of the consumption
of plant biomass range from 6% to 30% of net below-ground
production (Andersen 1987). It is unlikely that any
plant is completely immune from the effects of herbivory.
Herbivores and plants have shared a long
evolutionary history (Friis, Chaloner and Crane 1987).
Herbivory has probably been a strong selection force on
the patterns of leaf growth that have evolved in plants
(Mooney and Gulmon 1982). Virtually all leaves have
traits which, to some degree, reduce the intensity of
herbivory (eg. lignification, silification, secondary
compounds, reduced palatability). Similarly, herbivores
display a wide range of feeding morphologies and
v
behaviours, which have probably evolved in response to
the evolution of their forage plants. From the time of
the Cretaceous when the herbivore community was dominated
by dinosaurs that were generalists in their foraging
behaviour, browsing has been a common form of herbivory.
It was not until the mid- to late Eocene (45 mybp),
following world-wide cooling of climate and the radiation
of annual or biennial reproductive strategies in plants
and the presence of plants with perenating buds at or
below ground level, that large mammalian grazers became
common (Stebbins 1981; Wing and Tiffney 1987).
The graminoid (grasses and sedges) plant form, in
particular, is believed to have coevolved with large
ungulate grazers (Webb 1977, 1978; Stebbins 1981; Mack
and Thompson 1982; Coughenour 1985; Herrera 1985; Wing
and Tiffney 1987). Many graminiods possess
morphological traits that permit them to tolerate
repeated or partial defoliation (Hyder 1972; Dahl and
Hyder 1977; Briske 1986). Grasses and sedges usually
have their shoot apical meristems located at or below
ground level and have leaves which elongate from
intercalary meristems. These characteristics protect
meristems from herbivore damage and allow regrowth of
clipped leaves to occur (Mack and Thompson 1982; Crawley
1983; Coughenour 1985).
The question of whether or not herbivory is "good"
for plants is a controversial issue in ecology
(McNaughton 1983, 1986a,b; Westoby 1985, 1986; Belsky
1986a, 1987; Crawley 1987). Compensatory growth
responses of plants following defoliation by herbivores
may result in increases in fitness, or more often, in net
primary production (McNaughton 1986). The effects of
herbivory at the level of the individual, population,
community and ecosystem, and interactions between these
levels of organization will in turn influence the outcome
of a particular plant-herbivore interaction.
Herbivores act as agents of natural selection,
leading to differential performance of plant genotypes in
response to herbivory. Plants in grazed graminoid
communities frequently have prostrate leaves and shoots,
shortened internodes, and increased rates of leaf
production and of tillering (Turesson 1922; Stapleton
1928; McNaughton 1979a; Gray and Scott 1980; Crawley
1983; Detling and Painter 1983; Sadul 1987; Waisel 1987;
Jefferies 1988a). This implies modification of the
genetic structure of the population in response to the
overall effects of herbivory. Intensive herbivory is
likely to increase the mortality of the more suseceptible
genotypes, leading to an increase in the frequency of
HI
grazing-tolerant or resistant genotypes (see Wilbur 1976;
Snaydon 1980). Few studies have shown an increase in the
genetic fitness of individual plants following
defoliation (Belsky 1986a, 1987), and most claims of an
increase in genetic fitness are unconvincing (for example
see Owen 1978; Boscher 1979; Boucher and Sork 1979).
Paige and Whitham (1987) have recently shown that
individuals of Ipomopsis aqgreqata browsed by deer or elk
have nearly 2.5 times the relative fitness of intact,
ungrazed plants based on seed output, but the mechanism
is unclear.
From the point of view of the herbivore, enhancement
of forage availability and of the nutritional quality of
forage following grazing is a desirable attribute of
grazed swards. In grasses where propagation is achieved
primarily by vigorous vegetative clonal growth, fitness
measured in terms of seed production may be mostly
irrelevant (Crawley 1983). Changes in net above-ground
primary production may have the greatest influence on
plant-grazer interactions in these systems.
1.2 Positive feedbacks and nutrient cycling
in grazing ecosystems
Feedback processes can regulate the flow of
materials and the effect of various physical and biotic
processes in ecosystems (DeAngelis, Post and Travis
1986). When a population or ecological community is
perturbed slightly from "equilibrium", the balance of
positive and negative feedback mechanisms in the system
act to counteract the perturbation and restore the system
to "equilibrium" (assuming a stable equilibrium in the
first place) (DeAngelis, Post and Travis 1986).
Alternatively, feedback mechanisms may cause the system
to change, if the balance between positive and negative
influences is upset. The magnitude of various feedback
processes in the ecosystem determine the outcome, and as
in most processes involving more than one step, the one
which is rate-limiting is often the most important.
Nutrient cycling in ecosystems is often thought of
as a feedback process. In terrestrial ecosystems,
herbivores accelerate rates of nutrient mineralization
above those in their absence (Arman, Hopcraft and
McDonald 1975; Woodmansee 1978; Schowalter, Hargrove and
Crossley 1986; Ruess and McNaughton 1987). Nutrient
limitations on plant growth can be offset by the rapid
recycling of essential elements within an ecosystem and
these processes may be enhanced by herbivores. The uptake
of mineral nutrients by plants is often limited by the
rate of mineralization processes in the soil.
Nitrogen is a critically limiting nutrient for plant
growth (Lee, Harmer and Ignaciuk 1983; Stewart et al.
1983; Sprent 1987). Many studies have shown that it is
the most important nutrient limiting growth in tundra
ecosystems (Ulrich and Gersper 1978; Shaver and Chapin
1980), and in salt marshes (Pigott 1969; Stewart, Lee and
Orebamjo 1973; Valeila and Teal 1974; Cargill and
Jefferies 1984a).
A large number of processes are involved in nitrogen
inputs, transfers and losses in natural ecosystems (Clark
1981; Sprent 1987). Pathways of nitrogen cycling involve
all trophic levels, including live and dead plant
biomass, microbes (bacteria, actinomycetes, fungi), soil
animals (arthropods, nematodes, protozoans), invertebrate
grazers (grasshoppers), vertebrate grazers (large and
small mammals, birds), and chemical processes in the soil
and atmosphere. These processes include the
mineralization of nitrogen from dead organic matter (eg.
plant litter, animal faeces and carcasses), to ammonia
(ammonification) or nitrate (nitrification). Ammonium
ions (NH£ ) may be in either an exchangable or soluble
form, and depending upon conditions of temperature, pH
and moisture may volatilized as a gas (NHj). Ammonia
volatilization and denitrification of nitrate to gaseous
nitrogen (N^  and N^O) consitute pathways of nitrogen loss
from ecosystems. Leaching of organic and and inorganic
nitrogen following rainfall or snowmelt may also result
in losses from the system. Nitrogen may be immobilized
by the uptake of ammonium and nitrate by plants and soil
micro-organisms. In addition, free-living or symbiotic
cyanobacteria may fix atmospheric nitrogen.
A close coupling of the activities of the soil
microflora and the growth and nutrition of plants in
natural ecosystems exists (Fenchel and Blackburn 1979;
Lee, Harmer and Ignaciuk 1983). In fact, most plants are
dependent on the microbial breakdown of soil organic
matter for their nitrogen supply. Microbial activity
determines the rate of release of ammonium and nitrate to
higher plants, and any environmental factor which affects
the microorganisms will indirectly determine the
availability of these ions for plant growth. For
example, factors such as low temperatures, high soil
acidity (low pH), and anaerobic conditions may depress
nitrification (Aziz and Nedwell 1979; Fenchel and
Blackburn 1979; Lee, Harmer and Ignaciuk 1983).
1.3 Responses of Plants to Grazing
Grazing by animals affects the structure of plant
populations (Crawley 1983). In a series of mostly
discussion papers published in Forum in Oikos, Owen
(1980), Owen and Wiegert (1976, 1981, 1982a,b, 1983) and
Petelle (1982) suggested that there are a wide variety of
coevolved mutualisms between plants and herbivores. They
suggest that plants exploit consumers and that the
outcome of this arrangement is an increase in the genetic
fitness of plants. However, this hypothesis was
challenged on the grounds that most theoretical and
experimental evidence did not support it (Stenseth 1978,
1983; Silvertown 1982; Thompson and Uttley 1982; Herrera
1982, 1985; Belsky 1986a; Dirzo 1984; Lam and Dudgeon
1985) .
There is a considerable body of "suggestive"
evidence which supports the general idea that herbivory
results in increased net primary production, but there
are few studies which provide "real" evidence
demonstrating that this increase occurs, and also provide
evidence of the mechanism which accounts for this
8regrowth following grazing. Collectively however, the
evidence suggests that increases in production may occur
following grazing under some circumstances in both
terrestrial (Cook, Stoddart and Kissinger 1958; Vickery
1972; Dyer 1975; McNaughton 1976, 1979; Prins, Ydenburg
and Drent 1980; Cargill and Jefferies 1984b), and aquatic
algal and coral ecosystems (Bjorndahl 1980; MacDonald
1985; Bergquist and Carpenter 1986; Sterner 1986;
Carpenter 1986; Power 1987). On the other hand, plants
also produce a wide variety of secondary chemicals in
response to herbivory (Rhoades and Gates 1976; Rhoades
1979, 1983, 1985; Feeny 1980; Harborne 1982; Crawley
1983) . Therefore, the responses of plants to herbivory
may be considered under two general, operational
headings; 1) structural, phenological and chemical
defenses, and 2) production and growth responses. These
responses are not mutually exclusive (see Westoby 1980;
Bryant 1987; Coley 1988; Georgiadis and McNaughton 1988;
van der Meijden, Wijn and Verkaar 1988), however one type
of response generally dominates in a particular species.
A broad distinction between the two responses, based
on differences in the morphology between woody shrubs or
herbs which are generally characteristic of browsing
systems, and graminiods which dominate terrestrial
grazing systems, can be made. Graminoids have evolved a
number of defenses to grazers including secondary
chemicals, but they are generally more tolerant of
grazing than woody shrubs or herbs (Coughenour 1985).
The terms "defensive" and "tolerance" need to be
treated with some degree of caution because their
definitions can be interpreted a number of ways.
Defenses may have a negative impact on consumers, or they
may actually reduce net losses in terms of tissue biomass
or fitness (Schultz 1988). Defenses can be considered
avoidance mechanisms that reduce the probability of
defoliation, while tolerance mechanisms facilitate
regrowth following defoliation (Briske 1986). Tolerance
does not necessarily imply increased production or
fitness as a consequence of herbivory.
1.3.1 Plant defenses to the effects of herbivory
Defenses have important influences on the feeding
preferences of herbivores (Harper 1969, 1977; Rosenthal
and Janzen 1979; Crawely 1983; Sinclair and Smith 1984;
Robbins et al. 1987). Structural defences function by
making herbivory difficult, damaging or energetically
expensive. In graminoids common structural defences
include large amounts of silica, and indigestible lignins
and cellulose (McNaughton and Tarrants 1983; Coughenour
1985). These defences may also reduce the concentrations
of nutrients in plant tissues relative to the amount of
structural material (Mattson 1980; Crawley 1983). Other
avoidance mechanisms include reduced tissue accessibility
(ie. decreased leaf length and angle, and leaf/culm
ratio), awns or spines of the inflorescence, and changes
in leaf tensile strength at the individual level (Briske
1986), and interspecific associations with less palatable
species within the community (McNaughton 1978). Feeny
10
(1976, 1980) and Rhoades and Gates (1976) have also
suggested that ephemerality or rarity may reduce damage
to individual plants by herbivores. Asynchrony between
the phenology of plant growth and forage demand may limit
the size of a herbivore population within an area
(Crawley 1983; Drent and Prins 1987).
Many plants produce antiherbivore chemicals
(Fraenkel 1959; Harborne 1982; Crawley 1983). Plants or
tissues with a slow growth rate tend to be protected with
high concentrations of "quantitative" defences, such as
tannins and polyphenols, which act in a dosage-dependent
manner by inhibiting digestion (Feeny 1976, 1980; Rhoades
and Gates 1976; Coley 1988). "Quantitative" defenses
tend to be carbon-based and are not very labile (Coley,
Bryant and Chapin 1985). Plants or tissues with rapid
growth rates tend to be protected with low concentrations
of "qualitative" defences, such as alkaloids, cyanogenic
glycosides and terpenes. These toxins are usually
nitrogen-based and turn over rapidly (Feeny 1976, 1980;
Rhoades and Gates 1976; Harborne 1982; Coley, Bryant and
Chapin 1985)
Different patterns of phytochemistry have been
explained in terms of resource availability (Mooney and
Gulmon 1982; Coley, Bryant and Chapin 1985; Bazzaz et al.
1987) . Resource-limited plants with slow growth rates
and long lived tissues tend to develop "quantitative"
defences. Although these compounds are initially
expensive to produce, maintenance costs are low. Plants
that are not resource limited, which are fast-growing,
and have short-lived tissues tend to be protected with
11
low concentrations of "qualitative" defences. The most
obvious exception to these generalizations are graminoids
which are depauperate of defensive chemicals (Crawley
1983; Coughenour 1985). Jung, Batzli and Seigler (1979),
in a survey of the phytochemistry of arctic graminoids,
found that alkaloids were present only in small
concentrations (< 1 ppt); tannins, saponins, cyanogens
and anthraquinones were not detected.
Induced chemical defences generally are produced in
response to tissue damage (Rosenthal and Janzen 1979;
Crawley 1983; Coley, Bryant and Chapin 1985). These
generally are "qualitative" defences, and are usually
restricted to damaged areas. Induced "quantitative"
defences (tannins and resins) may also appear more
gradually (Rhoades 1979). Induced chemical defenses are
a form of phenotypic plasticity of plants, in response to
the effects of herbivory. Georgiadis and McNaughton
(1988) have shown that defoliation can induce
cyanogenisis in Cynoden plectostachyus, an African C<.
grass, and suggest that this trait evolved as a herbivore
deterent.
Carroll (1988) has suggested that fungal endophytes
may protect host plants from herbivores which are
poisoned by the mycotoxins produced by the endophytes.
In particular, many grasses are infected by fungal
endophytes which produce physiologically-active alkaloids
in the tissues of their host (Clay 1988). These fungal
endophytes are of the genus Acremonium (mainly in C,
grasses) or Balansia (mainly in C. grasses, but also
some C, grasses). Their hyphae occur intercellularly in
12
leaf and stem and root tissue, and they produce no
external morphological changes to the host plant (Clay
1988) .
Many infected grasses are known to be toxic to
livestock, however the effects of endophyte-infected
grasses on the foraging and diets of nondomestic and
insect grazers are generally unknown (Clay 1988).
Greater growth and seed production of infected plants has
been reported for perennial ryegrass and tall fescue in
the absence of grazing (Latch, Hunt and Musgrave 1985;
Clay 1987). The outcome of the interactions between
chemical defenses, fungal endophytes, herbivory, and
intrinsic and extrinsic compensatory growth mechanisms
may have profound consequences not just for the
individual plant but also for the structure and
composition of the plant community.
1.3.2 Productivity and growth rate of plants
in relation to herbivory
The growth responses of plants to defoliation as a
result of grazing can be grouped into three categories:
1) negative response, or undercompensation; 2) exact
compensation; and 3) enhancement, or overcompensation
(the herbivore optimization model; see Fig. 1-1). The
type of response observed depends upon the intensity and
frequency of grazing, the timing of grazing during the
season, and the phenology of plant growth (McNaughton
13
1979a; Dyer et al. 1982; Jefferies 1988a). Grazing can
affect both above- and below-ground production of grazed
plants. Clipping by itself is almost always detrimental
(Dahl and Hyder 1977; Lee and Bazzaz 1980; Sadul 1987),
but compensatory morphological and physiological
responses may enhance above-ground production. These
responses include changes in photosynthesis,
transpiration, nutrient uptake and translocation, and
rates of tissue turnover. All of these processes may
interact with each other.
A difference exists in the type of growth responses
observed between grazing and browsing systems, (Bryant,
Chapin and Klein 1983). The regrowth of woody plants
(browsing systems) is often limited by the availability
of resources and by the destruction of apical meristems.
There is a heavy reliance on chemical (usually carbon-
based) defences. In comparison, regrowth in many
graminoids (grazing systems) is much faster, since growth
rates and turnover rates of leaves are much faster and
meristems are generally protected from damage.
1.4 Compensatory Responses
A critical assumption of the herbivore optimization
model (Fig. 1-1) is that some plant species possess a
strong potential for compensatory regrowth following
grazing. A variety of mechanisms may account for
increased primary production following grazing.
Intrinsic mechanisms that operate at the level of
14
Fig. 1-1. Three possible effects of herbivory on plant
growth (or fitness). 1) a decline as the intensity of
herbivory increases; 2) compensation up to a threshold
level of herbivory, after which growth declines with
increasing herbivory; 3) enhancement of growth at moderate
levels of herbivory above that of ungrazed plants. At
higher intensities growth declines. This is the herbivore
optimization curve (after McNaughton 1979, 1983a; Dyer et
al. 1982; Jefferies 1988a).
EFFECT
ON
PLANT
GROWTH
PLANT
DEATH
(UN6RAZED
PLANTS)
INTENSITY
OF HERBIVORY
15
individual plants and extrinsic mechanisms that involve
processes at the community or ecosystem level have been
proposed to account for an increase in production
(McNaughton 1983a,b).
1.4.1 Intrinsic Mechanisms
1.4.1.1 Photosynthesis
Photosynthetic CO., assimilation by leaves is
affected by the nutrition of the plant, the light regime
during growth, leaf age, water stress and many other
environmental and physiological factors. Both an increase
and a decrease in photosynthetic rates in tissue
remaining or produced after grazing have been observed
(King, Wardlaw and Evans 1967; Gifford and Marshall 1973;
Detling, Dyer and Winn 1979; Caldwell et a1. 1981;
Painter and Detling 1981; Detling and Painter 1983; Nowak
and Caldwell 1984; von Caemmerer and Farquhar 1984;
Wallace, McNaughton and Coughenour 1985). In Bouteloua
gracilis, Detling, Dyer and Winn (1979) measured both
increased net photosynthetic rates within three days of
defoliation, and an increase in the proportion of new
photosynthate allocated to the production of leaves.
Gifford and Marshall (1973) suggested that decreases in
mesophyll resistance to CO- diffusion in leaves of Lolium
remaining after partial defoliation may lead to increased
photosynthetic rates. A more likely explanation is that
16
partial defoliation causes a proportional increase in
both ribulose-1,5-bisphosphate (RuBP) carboxylase-
oxygenase activity, and RuBP regeneration capacity (von
Caemmerer and Farquhar 1984). These authors measured a
significant increase in CO.-assimilation rate of leaves
of Phaseolus vulqaris eight days after partial
defoliation, which was accompanied by an increase in
chlorophyll concentration per unit leaf area and by an
increase in in-vitro RuBP-carboxylase activity.
A number of graminoid species show enhanced
photosynthetic rates when partial defoliation occurs
(Caldwell et aJL. 1981; Wallace, McNaughton and Coughenour
1985). However, Detling and Painter (1983) found no
difference in photosynthetic rates or stomatal
conductances between two ecotypes (one from grazed sites,
one from an exclosed site) of Agropyron smithii following
clipping. Similarly, Nowak and Caldwell (1984) did not
record a difference in gas exchange rates in clipped and
undipped leaves of the same age of plants of A.
desertorum and A. spicatum. The overall significance of
increased rates of photosynthesis in leaves following
grazing in relation to compensatory growth is unclear.
1.4.1.2 Resource Allocation
Herbivory frequently alters patterns of resource
allocation. A number of studies have reported that the
proportion of photosynthate allocated to the production
of new leaf area increases after grazing (Ryle and Powell
17
1975; Detling, Dyer and Winn 1979; Detling et al. 1980).
Richards (1984) reported an increase in allocation of
current photosynthate from roots to shoots. While
foliage herbivory often increases upward translocation of
assimilates at the expense of root growth, grazing of
roots may reverse this pattern (Crawley 1983; Andersen
1987). However, the ability of plants to support above-
ground production with below-ground reserves, and vice
versa, is not entirely clear (see Section 1.4).
A distinction must be made between carbon
(carbohydrate) and nutrient (particularly nitrogen and
phosphorus) limitation of growth. Although well
fertilized plants may be carbon-limited, under natural
conditions plant growth is frequently limited by nutrient
availability (Harper 1977). This is particularly
pronounced in nutrient-deficient habitats. The regrowth
of leaves following defoliation requires substantial
quantities of mineral nutrients and carbohydrates.
Calculations of nutrient budgets of tundra graminoids at
Barrow, Alaska, suggest that leaf regrowth following
defoliation depletes plant nitrogen and phosphorus
reserves more rapidly than carbohydrate reserves (Chapin
1977) .
Herbivory often results in increases in the quality
of forage. Translocation of stored nutrients from the
below-ground biomass and increased uptake of inorganic
nutrients can lead to high concentrations of nitrogen and
other nutrients and soluble (non-structural)
carbohydrates in the developing leaves of grazed plants
(Ydenberg and Prins 1981; Cargill and Jefferies 1984b;
18
McNaughton and Chapin 1985; but see Ruess 1984).
However, while grazing may result in changes in resource
allocation patterns, new growth cannot be sustained if a
resource is limited. Eriophorum vaginatum and other
graminoids frequently grow in nutrient-limited habitats
in the Arctic. Sequential leaf development is associated
with the movement of nutrients from senescing leaves to
developing leaves, thereby minimizing annual nutrient
requirements from the soil (Jonasson and Chapin 1985).
Under these conditions removal of leaves by herbivores
restricts further leaf development. The nutrient pool
has to be replenished (by fertilization), and the plants
must be able to assimilate these nutrients at a rate
proportional to the grazing intensity, or else new growth
will cease.
1.4.1.3 Morphological Plasticity
Grazing is known to increase rates of tillering in
graminoids (Stapleton 1928; Peterson 1962; Younger 1972;
Laude 1972; Belsky 1986; Kotanen and Jefferies 1987;
Bazely and Jefferies 198N). These responses could be due
to the removal of apical dominance if the herbivore has
removed the apical bud. At the community level,
defoliation results in the opening of the canopy and
increased light penetration which leads to rapid changes
in the growth rates of the surrounding plants as they
exploit the light gaps. In grasslands, these responses
are related to the morphology of graminoids, which
19
protect meristems from damage during grazing (Hyder 1972;
Dahl and Hyder 1977; Briske 1986).
At the level of the individual tiller the
probability of defoliation may be reduced as a result of
the existence of a number of so called avoidance
mechanisms (Branson 1953). In species where shoots are
not erect, so that the apical meristems are close to the
ground or else are surrounded by vegetative tillers, the
meristems are not likely to be destroyed and hence plant
growth can be sustained following defoliation. Growth is
likely to be most rapid from intercalary meristems,
followed by newly developed leaf primordia, and least
rapid from newly initiated axillary buds (Briske 1986).
Leaf replacement following grazing is largely a function
of the number, source and location of meristems of each
plant. In Puccinellia phryganodes, grazing by Lesser
Snow Geese resulted in twice as many axillary tillers
produced per main tiller in grazed plants compared to the
number in ungrazed plants (Bazely and Jefferies 198N).
The clonal growth of graminoid vegetation permits rapid
regrowth following grazing. Morphological plasticity is
an important mechanism by which grazed plants can sustain
production when grazed.
1^4.2 Extrinsic mechanisms
1.4.2.1 Water-use efficency
McNaughton (1979, 1983a,b, 1985) has suggested that
grazing leads to the conservation of soil moisture by
20
reducing transpirational water losses, because of reduced
leaf area of grazed plants. The validity of this
observation has been questioned by Belsky (1986a), based
on the results of studies by Younger (1972) and
Ellison (1960). Hodgkinson and Baas Becking (1977) have
shown that defoliation during periods of low water
availability accelerate plant death, because root growth
is impaired as a result of lack of assimilates, and new
roots cannot extend into the lowered water table.
Detling and Painter (1983), and Nowak and Caldwell (1984)
found no difference in photosynthetic water-use efficency
between defoliated and undipped plants. Cox, Parr and
Plant (1988) found that frequent clipping reduced water-
use efficiency of plants of Lolium perenne in the first
year but that clipping had no effect on established
swards. These contradictory results suggest that grazing
has a very unpredictable influence on water- use
efficency at the level of the individual and community,
and that this is not likely to be an important
compensatory mechanism, except perhaps in arid regions.
Since the response of plants is linked to plant
architecture, any generalizations are difficult.
1.4.2.2 Salivatory hormones
The stimulatory effects of salivatory hormones on
grasses has been investigated as a possible compensatory
mechanism (Reardon 1972; Dyer and Bokhari 1976; Dyer
1980; Dyer et al. 1982; McNaughton 1985). Classes of
chemicals recognized as growth factors or regulators in
21
plants which are involved in molecular, cellular, and
physiological activities may be transfered across trophic
levels from herbivores to plants. Experiments with
thiamine and bovine saliva suggested that direct
application of saliva stimulates plant growth (Reardon,
Leinweber and Merrill 1972). However, in a subsequent
publication (Reardon, Leinweber and Merrill 1974), no
stimulation of plant growth was reported. Johnson and
Bailey (1972) showed that saliva did not promote the
growth of two Festuca species, and Detling et al. (1980,
1981) found that bison saliva did not lead to an increase
in plant growth or photosynthetic physiological activity
of Avena sativa and Bouteloua gracilis.
McNaughton (1985) detected increased growth of
thiamine treated leaves of the undipped grasses,
Sprobolus ioclados and S. pyramidalis, but thiamine did
not promote the growth of defoliated plants beyond that
of ungrazed plants. Thiamine application also resulted
in a decreased concentration of leaf nitrogen (McNaughton
1985). In summary, the influence of salivatory hormones
on the growth of grazed plants resulting in enhanced
production is not established as a compensatory
mechanism.
1.4.2.3 Fertilization
Perhaps the most important extrinsic factor
responsible for the enhancement of plant growth following
grazing is the effect of fertilization from faeces and
22
urine (see Section 1.2). Herbivores play an important
role in nutrient cycling in most ecosystems (Mattson and
Addy 1975; Bjorndal 1980; McNaughton, Coughenour and
Wallace 1982; Ruess and McNaughton 1984; Bazely and
Jefferies 1985; Schowalter, Hargrave and Crossley 1986).
Excretion of soluble nutrients by herbivores accelerates
the availability of nutrients that normally would be
limited by the rate of plant litter decomposition. This
accelerated transfer of nutrients is essential to sustain
the growth of grazed swards (McKendrick et al. 1980;
Coppock et al. 1983; Jefferies 1988a). It is likely to
be of particular significance in systems which are
intrinsically nutrient-limited. Grazers may also
indirectly affect the balance between the net
mineralization and immobilization of nutrients in soils
by altering the dynamics of the soil microbial processes
(Ruess and McNaughton 1987; Chapter 5).
Nutrient availability affects the suitability of
plants as forage for herbivores. Most plants generally
experience nutrient flushes early in the growing season
and the have highest uptake capacity at that time (Chapin
1980). Because grazing is coincident with the return of
nutrients to the soil from faeces and urine, plants may
be able to maintain a high rate of uptake of nutrients
for a long period during the growing season. In addition
uptake rates of nutrients may be considerably higher in
grazed plants, than those in ungrazed plants (see Ruess,
McNaughton and Coughenour 1983).
A number of authors have recently suggested that the
effects of fertilization are "blatantly group
23
selectionist" (Belsky 1987; Crawley 1987). This arguement
is based on the observation that the effects of
defoliation affect only some of the plant species in the
community, while nutrients recycled in faeces are
available to all plants. However, in grazing systems
very few plants on a "grazing lawn" escape defoliation
(McNaughton 1984; Jefferies 1988a). Effects of herbivory
evident at one level of organization (ecosystems,
communities, populations or individuals), may not be as
obvious at another level. Fertilization by faeces and
urine has effects at the level of the individual plant as
well as at the level of the entire grazing ecosystem.
All of the effects of fertilization need to be considered
before concluding that this mechanism has no influence on
individual selection.
1.5 Grazing and Below-Ground Biomass and Production
Grazing may have effects on root growth and the
activities of roots (Milthorpe and Davidson 1977;
Andersen 1987). Root growth is reduced in many plants
following clipping of above-ground tissue (Crider 1955;
Hodgkinson and Baas Becking 1977; Detling, Dyer and Winn
1979). Excessive grazing causes a dramatic decline in
root growth, and the rate of shoot development is
reduced. However, clipping of culms only affected root
growth of individual ramets (Crider 1955). Since
selective clipping is typical of the natural grazing
patterns of many animals (Stephens and Krebs 1986), these
24
results suggest that the effects of grazing on root
growth are more complicated (and possibly not as severe)
as many studies might indicate.
In many tundra plants there is a large source of
fixed carbon, energy and nutrients below-ground. This
source is available for plant growth following herbivory.
Chapin and Slack (1979) suggest that defoliation of
increasing intensity produces the following responses: 1)
decreased root growth, 2) decreased activity of existing
roots, and finally 3) increased root mortality. However,
in perennial plants which have large below-ground
reserves, root function may diminish only after below-
ground nutrient reserves are depleted. This emphasizes
the need for nutrient input following grazing if
production is to be sustained in nutrient-limited
environments.
In two tundra graminoids, Eriophorium vaginatum and
Carex acpaatilis, moderate experimental defoliation
stimulated root respiration and phosphate absorption
(Chapin and Slack 1979). The was the result of lowered
root phosphorus levels following defoliation, as nutrient
reserves were reallocated to support shoot growth.
However, the investigators noted that Carex aquatilis has
a large perennial root system and is less sensitive to
defoliation than Eriophorum which produces a new set of
roots each season.
Cargill and Jefferies (1984b) did not detect a
difference in the below-ground biomass of grazed plants
of Puccinellia phyrganodes and Carex subspathecea
compared to ungrazed plants. However, this does not imply
25
that the rate of turnover of biomass between grazed and
ungrazed sites is not different. Measurement of small
changes in live below-ground biomass is very complicated
because it is difficult to separate live and dead roots.
The extensive root systems of established plants may
contain sufficient carbohydrates to support root growth
and maintenence following grazing, until enough
photosynthate is produced and translocated below-ground.
Below-ground herbivory is also likely to be
important but its effects are poorly understood (Andersen
1987; Seartedt, Ramundo and Hayes 1988). Some plants
respond to the removal of root tissue by increasing the
growth rate of the remaining roots and by initiating new
roots (Andersen 1987). Because there is often a
simultaneous decrease in shoot growth proportional to the
degree of root loss, below-ground herbivores may have a
major role in determining the energy and nutrient flow
through the community (Andersen 1987). In a recent study
of the dynamics of the interactions between soil
arthropods and the prairie grass, Andropogon gerardii,
Seartedt, Ramundo and Hayes (1988) have shown a strong
positive correlation between the intensity of above-
ground herbivory and the density of below-ground
herbivores. The results of these recent experiments re-
enforces the concept that the effects of herbivory are
evident at all levels of community organization.
1.6
26
Principle Objectives
The general goal of this study was to investigate
the response of Puccinellia phyrganodes, a stoloniferous
salt marsh grass, to different levels of grazing by
captive goslings of the Lesser Snow Goose (Chen
caerulescens caerulescens). The effect of goose faeces
on nitrogen cycling and rates of nitrogen mineralization
in faeces were also examined. Previous studies of the
effects of snow geese on production, nutrient cycling and
composition of the grazed salt marsh at La Perouse Bay,
Manitoba, since 1978, provided the framework for
conducting experiments in this study (see Jefferies 1988a
for an overview). The herbivore optimization model,
which predicts that grazing may lead to the enhancement
of net primary production under certain conditions,
provided the conceptual framework (see McNaughton 1979,
1983a,b; Hilbert et al. 1981; Dyer et al. 1982; Jefferies
1988a). The major objectives and rationale for these
experiments are listed below.
1. An experimental test of the prediction of the
herbivore optimization model, that moderate levels of
herbivory lead to the greatest enhancement of NAPP of
Puccinellia phryganodes grazed by geese.
Cargill and Jefferies (1984b) found that grazing by
Lesser Snow Geese led to increases in NAPP of grazed
Puccinellia under field conditions. However the
relationship between the length of the grazing period and
27
the increase in NAPP could not be established. This can
be accomplished by controlling the duration of grazing on
swards of Puccinellia in experimental plots by using
captive goslings.
2. An examination of the effects of the timing of grazing
within a season and the interval between grazing events,
on the type of response exhibited by swards of
Puccinellia.
The ecological time frame over which enhanced NAPP
occurs in response to grazing is rarely discussed
(Jefferies 1988a). The capacity of the plants to recover
from the effects of grazing is dependent upon the timing
and intensity of herbivory during the season, and the
phenology of plant growth. A number of experiments were
conducted to examine the capacity of swards of
Puccinellia to recover from grazing at different times
during the summer. In particular, the effects of
repeated partial defoliation at different intervals
during the season were examined.
3. The determination of the relative effects of clipping
and fertilization by goose faeces on the NAPP of swards
of Puccinellia.
Fertilization by goose faeces is essential for the
increase in NAPP of Puccinellia following grazing (Bazely
and Jefferies 1985). The relative effects of clipping
and fertilization at different levels of grazing may
28
change the type of growth response observed. Grazing
experiments where faeces either remained or were removed
from grazed swards of Puccinellia were conducted to
examine this question.
4. An examination of microbial-mediated nitrogen
mineralization processes in goose faeces derived from
three graminoid forage species, Puccinellia phryganodes,
Carex subspathacea and Calamagrostis deschampsiodes.
Soluble nitrogen of goose faeces is rapidly leached
to the sediments within 48 hours and is taken up by
plants (Bazely and Jefferies 1985).s The microbial
biomass present in goose faeces may also contribute to
the net nitrogen balance in the marsh as a result of the
net mineralization of organic nitrogen present in faeces.
Rates of net nitrogen mineralization and microbial
respiration in fresh goose faeces derived from goslings
grazing on three forage species were measured on three
occasions during the summer.
29
CHAPTER 2: DESCRIPTION OF STUDY SITE AND SPECIES
2.1 Description and History of the La Perouse Bay
Grazing System
La Perouse Bay is located on the west coast of Hudson
Bay, approximately 25 km east of Churchill, Manitoba (58* ,
24' N, 94", 24' W) (Fig. 2-1). It lies within the belt of
coastal tundra that extends along Hudson Bay in this
region. The treeline is approximately 10 km inland. The
duration of the snow-free period averages four months,
from the end of May until the end of September. Snow melt
usually occurs in late May, at which time the entire area
is flooded. Minimum temperatures are consistently greater
than 0°C in June, July, August and September (Fig. 2-2;
Table 2-1). Maximum temperatures average 10°to 15"C, but
can vary from below 0" C to 30°C, within the same 24 h
period. Annual precipitation is approximately 450 mm, of
which approximately 60% falls during the snow-free period.
Over 50% of the total number of hours of bright sunshine
occur during this period (Table 2-1).
The La Perouse Bay colony of Lesser Snow Geese (Chen
caerulescens caerulescens (L.) Gundl.) currently numbers
about 8000 nesting pairs, although tens of thousands of
migrants pass through the area each spring and autumn.
Snow Geese are the dominant and most important herbivore
of graminoid vegetation at La Perouse Bay. Canada Geese
(Branta canadensis) are also resident during part of the
summer, but at much lower numbers (< 100). Mammalian
herbivores which occur infrequently in the area are
30
Fig. 2-1. Map of of Hudson Bay showing the location of La
Perouse Bay, Manitoba.
w^"-'. '.-iiS ;^: • I v u j i v i kRankin
Inlet %__ .^ inlet
HUDSON BAY
^Churchill
Perouse
Ifx Bay
, Cape Churchill
• '.'•'iiiilVaii'. • • '•
iFort Severn Islands?;^:;
»^«"---. • . • - - .-.<>
'.iiii^  Poste-de-la
Baleine
Moosonee(y>ip
Superior '•'$
31
Fig. 2-2. Daily maximum and minimum temperatures at
Churchill, Manitoba during the summer of 1986 (a) and 1987
(b).
3
0
-
2
0
-
1
0
-
-1
0
-2
0
I
 
I
M
 
J
 
J
 
A
 
S
M
ONTH
30
 
-
-1
0
-2
0
M
M
ONTH
32
TABLE 2-1. Summary of precipitation (rainfall and
snowfall water equivalent), bright sunshine hours, and
mean high and low temperatures for May, June, July, August
and September in 1986 and 1987, and yearly totals. Data
compiled from Churchill, Manitoba monthly climatological
summary (Environment Canada, Churchill Weather Office).
1986
MAY JUNE JULY AUG. SEPT.
Precipitation
(mm)
Rainfall 29.6 103.0 48.8 74.7 40.8
Snowfall 0 15.3 0 0 2.4
Water
Equivilent
Bright 287.9 223.6 247.0 233.1 73.7
Sunshine
Hours
Mean High ( °C) 6.0 10.5 14.9 15.6 7.5
Mean Low (°C) -3.7 1.0 5.3 6.0 1.2
1987
MAY JUNE JULY AUG. SEPT.
Precipitation
(mm)
Rainfall 3.6 58.7 37.0 66.0 42.0
Snowfall 65.4 0.9 0 0 0.2
Water
Equivilent
Bright 196.7 251.0 339.6 233.5 76.0
Sunshine
Hours
Mean High ( °C) 1.7 10.9 17.6 13.8 10.1
Mean Low (°C) -6.2 0.9 6.6 5.9 4.3
YEAR
TOTAL
306.0
191.9
1821.8
YEAR
TOTAL
225.3
204.8
1831.9
33
caribou (Rangifer tarandus) and meadow voles (Microtus
pennsylvanicus). A few species of birds also feed on
graminiod vegetation for short periods (both residents
(eg. Willow Ptarmigan, Lagopus lagopus albus) and migrants
(eg. Snow Buntings, Plectrophenax nivalis nivalis)) (see
Savory 1976; Bazely 1987).
Salt marshes, such as those at La Perouse Bay, are a
common physiographic feature along the southern and
western coastlines of Hudson Bay. The rate of isostatic
uplift along this coast is estimated to be between 0.5 and
1.2 m per century (Hunter 1970; Andrews 1974; Hansel et
al. 1983; Scott, Hansel and Fayle 1987). This high rate
of uplift combined with the flatness of the land (0.5 to
1.0 m in elevation per km) results in the appearance of an
estimated 100 m to 200 m of new shoreline every decade
(Martini 1982). In many areas this newly emergent land
is utilized by breeding geese (Gaston et al. 1987).
The vegetation of these extensive intertidal flats
and emergant sediments at La Perouse Bay is dominated by
Puccinellia phyrqanodes (Trin.) Scribn. and Merr., a
stoloniferous arctic salt marsh grass. In less saline
sites and where freshwater drainage streams dissect the
tidal flats Carex subspathecea Wormsk., a rhizomatous
sedge, dominates the vegetation, and often Puccinellia and
Carex occur in mixed swards. These two species acccount
for over 90% of the above-ground standing crop (Bazely and
Jefferies 1986). Other species present at low frequencies
are Triglochin palustris L., and a number of
dicotyledonous species including Potentilla egedii
Wormsk., Ranunculus cymbalaria Pursh., Stellaria humifusa
34
Rottb., and Plantago maritima L. The soft sediments of
freshwater streams are dominated by Hippuris tetraphylla
L. . The total area of the salt marsh is about 5.4 km1
(Cargill and Jefferies 1984b).
Inland from the intertidal flats the vegetation is
dominated by several species of willow (especially Salix
brachycarpa Nutt. and Salix phylicifolia L. ssp.
planifolia (Pursh) Hiitonen), shrub birch (Betula
glandulosa Michx.), and Myrica gale L. which establish on
frost-heave mounds. Ground cover is composed of Elymus
mollis Trin., Carex subspathecea, Festuca rubra L.,
Calamagrostis deschampsiodies Trin., Dupontia fisheri R.
Br., and a number of other graminoids and herbs (see
Jefferies, Jensen and Abraham 1979). Low-lying ground and
small pools in this zone are often very saline (Jefferies,
Jensen and Abraham 1979; lacobelli and Jefferies
unpublished), and may be colonized by the halophyte
Salicornia borealis Wolff and Jefferies (Wolff and
Jefferies 1987). The willow areas are the primary zone of
nesting of Lesser Snow Geese (Hik 1986; Jackson, Hik and
Rockwell 1988).
Further inland freshwater ponds and sedge meadows are
dominated by Carex ao^iatilis Wahl., Carex X flavicans Nyl.
and Eriophorum angustifolium Honck.. Many other graminoids
also occur in these fresh-water habitats. Saturated
carpets of moss (mainly Drepanocladus uncinatus (Hedw.)
Warnst.) surround the edges of many ponds. This zone is
also used by nesting and feeding geese, although not as
heavily as the saltmarsh.
35
Growth of salt marsh graminiod species is strongly
nitrogen- limited (Cargill and Jefferies 1984a; Frey
1988) . The accelerated cycling of nitrogen via goose
faeces bypasses normal decomposition processes, and
provides a renewable source of nitrogen for plant growth
during the summer. Fertilization by goose faeces
increased the net above-ground primary production of
Puccinellia by up to 105%, compared to ungrazed swards;
approximately 80% of the annual NAPP was consumed by the
geese (Bazely 1984; Cargill and Jefferies 1984b; Bazely
and Jefferies 1985). Grazing also prevents the
accumulation of litter and limits the formation of a
canopy. Because of this, nitrogen-fixing cyanobacteria
are able to colonize the surface of the sediment between
grazed shoots, thereby increasing the absolute input of
nitrogen and replacing nitrogen which the geese export
south at the end of each summer (Bazely and Jefferies
1988). Grazing by the geese also retards changes in
community compositition, which becomes evident when salt-
marsh vegetation is protected from grazing by the erection
of exclosures (Bazely and Jefferies 1986).
In spring, before the onset of above-ground plant
growth, geese grub for roots and rhizomes of graminiod
plants in the salt marsh (mainly Puccinellia phyrganodes),
which creates patches of disturbed sediments (Jefferies
1988a,b). Approximately 40% of the shoots of Puccinellia
are removed by the geese. Small patches are generally
recolonized by the remaining plants. Large patches on the
other hand, may be seriously and irreversibly damaged by
secondary erosion (Jefferies 1988a,b). Grubbing of the
36
graminoids under and around willow vegetation leads to the
rapid death of willows by exposing their roots (lacobelli
and Jefferies, unpublished). In the freshwater sedge
meadows geese eat the swollen basal portions of shoots of
carices in spring. Up to 1600 shoots per m2 may be
removed (Kotanen 1987, 1988). Intensive shoot pulling of
developing shoots of Carex aquatilis by geese in the
spring has been implicated in the production of moss
carpets (Jefferies and Kerbes 1985; Kotanen 1987;
Jefferies 1988a,b). Leaves of plants of both salt and
fresh water marshes are grazed during the summer.
The La Perouse Bay Snow Goose colony is one of the
most southerly breeding grounds and was probably
established only recently. Small numbers of geese were
first documented nesting in the area during the summers of
1953 (Wellein and Newcomb 1953) and 1957 (Foster 1957).
In 1943 no geese were observed nesting in the area
(Lumsden, cited in Cooke 1981), and there was no evidence
of nesting in previous centuries (Cooke 1981). Hanson et
al. (1972) reported no nesting in 1962, but at least 2500
pairs of geese nested in 1963, and successful nesting by
large numbers of geese has been documented every year
since then. This large increase in 1963 may be explained
by a group of nesting birds, destined for breeding
colonies further north, which were forced to nest at La
Perouse Bay due to adverse weather conditions (see
Geramita and Cooke 1982 for recent documentation of a
similar immigration). This original group of immigrants
formed the nucleus of the colony which continued to
persist as their offspring returned. This is a
37
consequence of strong female natal philopatry (Abraham
1980; Cooke and Abraham 1980). Between 1968 and 1987 the
colony increased in size from 1200 to 8000 breeding pairs,
reaching a peak of 9700 in 1983. Recent estimates of
colony size have been confounded by the geographic
expansion of nesting areas beyond the boundaries of the
traditional colony. Detailed studies of the breeding
biology, behaviour, genetics and ecology of Lesser Snow
Geese at La Perouse Bay have been conducted continuously
since 1968 (see Cooke et al. 1982), and of the
interactions between geese and local plant communities
since 1978 (see Jefferies 1988a).
2.2 Biology of Puccinellia phyrganodes
Puccinellia phryganodes (Trin.) Scribn. and Merr. is
the dominant grass species on the salt marsh flats at La
Perouse Bay. The Puccinellia genus contains generally
caespitose or stoloniferous, yellowish-green, smooth
grasses found in alkaline or saline environments (Porsild
and Cody 1980). P. phryganodes is a halophyte that
dominates arctic and sub-arctic coastal marshes and it has
a circumpolar distribution (Polunin 1940; Hulten 1968;
Porsild and Cody 1980). It is the primary colonist of
intertidal mud flats and unconsolidated silt in arctic
coastal areas (Jefferies 1977).
P. phryganodes is a stoloniferous species that forms
caespitose or densely tufted mats (Fig. 2-3). It often
38
occurs in pure swards, or in association with another
common graminoid, Carex subspathacea Wormskj., a
rhizomatous sedge. Grazed swards of P. phyrganodes at La
Perouse Bay rarely exceed 2 cm in height (Cargill 1981),
although ungrazed plants may grow to 15 cm (Polunin
1940; personal observation). Typical of many grasses, the
apical meristem is close to the ground in Puccinellia.
This highly active region of growth produces leaf
primordia, which eventually form nodes, on opposite sides,
just beneath the meristem (Langer 1979). Each primordium
is completely meristematic at the beginning of its
development, but soon is restricted to an intercalary
meristem at its base. Puccinellia forms extra-axillary
(extravaginal) shoots which act as stolons and have the
capability of rooting at their nodes. Puccinellia
contains low amounts of crude fiber and silica, no known
secondary defensive compounds and has a high concentration
of nitrogen in its tissue (Cargill and Jefferies 1984b).
Thus it is particularly high quality goose forage.
P. phryganodes is a sterile triploid (2n = 21) in the
Hudson Bay region, which rarely flowers and is not known
to set seed (Polunin 1940; Bowden 1961; Dore and McNeill
1980; Jefferies and Gottlieb 1983; Sadul 1987). Pollen is
sterile. P. phryganodes reproduces vegetatively by
extensive clonal. propagation, however dispersal mechanisms
are effective. Ice rafting, tidal action and spring
runoff all are means by which pieces of the plant are
transported around the saltmarsh. Broken shoots and
leaves of P. phryganodes are able to root in soft
sediments and rapidly colonize soft sediments and
39
Fig. 2-3. Diagram of the growth habit of of Puccinellia
phryqanodes (Trin.). Scribn. and Merr. The positioning of
an axillary shoot is magnified.
99.
'
O
40
disturbed sites (Jefferies, unpublished).
Sadul (1987) has detected considerable
electrophoretic variation amoung plants from La Perouse
Bay. This variation in the population appears to be
maintained by frequent abiotic and biotic disturbances,
followed by recolonisation (Sadul 1987; Jefferies 1988a).
There is also considerable morphological variation among
individuals. This variation may reflect the fact the
there is selection for genets with slow growth rates in
ungrazed sites and selection for genets with faster growth
rates under grazed conditions (also see Gray and Scott
1980) .
2.3 Biology of Lesser Snow Geese
2.3.1 Distribution
Snow geese are members of the tribe Anserini, in the
family Anatidae, the ducks, geese and swans. Currently
two subspecies of snow geese are recognized: the Lesser
Snow Goose (Chen caerulescens caerulescens (L.) Gundl.)
and the Greater Snow Goose (C. c. atlantica Kennard)
(Godfrey 1979; Owen 1980). The two subspecies are
distinguished on the bases of their sizes and
distribution. In addition, Lesser Snow Geese display a
conspicuous "white" or "blue" plumage polymorphism. The
blue phase is rarely observed in the Greater Snow Goose
41
(Cooke and Cooch 1968) . The Lesser Snow Goose is the
subspecies occuring at La Perouse Bay, Manitoba.
The eastern Arctic population of Lesser Snow Geese
numbers about two million birds (Boyd, Smith and Cooch
1982). This population has been increasing rapidly over
the past two decades (Maclnnes and Kerbes 1987; Cooch
1988). The geese winter on the Gulf coast of Texas and
Louisiana, and are becoming increasingly common in grain
fields in the midwest States along the Central and
Missisippi flyways. These geese nest in colonies in the
south and west coasts of Hudson Bay, the southwest coast
of Baffin Island, Southhampton Island, and the smaller
islands in the Foxe Basin and Hudson Bay (Gaston et al.
1987) .
Snow geese are usually monogamous and females return
with their mates to their natal colonies to breed (Cooke,
Maclnnes and Prevett 1975). The consequences of this
female natal philopatry on the genetic structure of the La
Perouse Bay population is approximately 50% gene flow
(from males) each generation (Rockwell and Cooke 1977;
Rockwell and Barrowclough 1987). Female snow geese
display strong local philopatry at La Perouse Bay,
generally nesting within 500 m of their previous nest site
(Abraham 1980; Cooke et al. 1983; Hik 1986).
2.3.2 Reproduction and growth
The arrival of snow geese on the colony is generally
coincident with snow melt (Abraham 1980; Davies and Cooke
42
1983). Females begin to lay eggs soon after arriving.
Nests are generally located on snow-free willow hummocks
(Abraham 1980; Jackson, Hik and Rockwell 1988). Modal
clutch size is four, however there has been a systematic
decline in clutch size (about 0.5 egg) over the past 15
years (Cooch et al. 198N). The duration of the
incubation period is approximately 23 days and goslings
hatch in late June or early July. During this period
incubating females may lose up to 20% of their body
weight (approx. 400 g; Ankney and Maclnnes 1978).
Hatching is synchronous, occurring at all nests within a
two week period (Findlay and Cooke 1982). Following
hatch, adults and goslings graze intensively on the salt-
marsh vegetation. At this time the population of geese on
the colony may exceed 30,000. Both adults and goslings
are flightless during this period. Their demand for
forage is very high. In a six week period the goslings
increase in weight from 80 g to over 1500 g, and females
regain weight lost during incubation. By fledging (six
weeks) the size of each brood has been reduced to
approximately two goslings (Cooke et al. 1985; Rockwell,
Findlay and Cooke 1987), a consequence of predation by
jaegers, gulls, Arctic foxes, wolves and polar bears, and
disease. Obtaining sufficient quantities of high quality
forage may also be a problem in some years, leading to
increased gosling mortality.
43
2.3.3 Foraging behaviour and digestive physiology
The diet of Snow Geese is dominated by graminoids.
On the wintering grounds and on the migration flyways
their diet consists primarily of the roots and rhizomes of
plants of coastal marshes. The most important forage
plants include species of Scirpus, Spartina, Distichlis,
and Typha (Lynch, O'Neil and Lay 1947; Burton, Hudson and
Bragg 1979; Owen 1980; Smith and Odum 1981; Smith 1983;
Bedard, Nadeau and Gauthier 1986). In the past decade
waste rice and corn, and cereal stubble on farmland along
the flyway and Gulf coast have become important sources of
forage during the migrations and during the winter months
(Reed 1976; Bedard, Nadeau and Gauthier 1986). Grubbing
for forage during this time of year is often destructive
(Lynch, O'Neil and Lay 1947; Smith and Odum 1981; Smith
1983). In spring this destructive grubbing also occurs on
the breeding grounds, as the geese often arrive prior to
melt and the commencement of plant growth (Jefferies
1988a,b).
Many studies have indicated that the critical factor
determining the success of breeding is the amount of food
available to parents for building up body reserves prior
to breeding (see Perrins 1970; Sinclair 1977; Teunissen,
Spaans and Drent 1985). Thus forage aquisition during the
spring migration is critical for breeding Snow Geese (see
Wypkema and Ankney 1979; Davies and Cooke 1983; Hamman,
Andrews and Cooke 1986) . The preferred foods during this
time include the roots, rhizomes and young shoots of marsh
plants. Prevett, Marshall and Thomas (1979, 1985) found
44
that migrating geese in James Bay consume about equal
proportions of below- and above-ground tissue of grasses
(mainly Puccinellia phryganodes), sedges, Triglochin spp.
and Equisetum spp.. Geese feed selectively on the parts
of plants which have the highest nitrogen content (Thomas
and Prevett 1980, 1982).
During the summer at La Perouse Bay geese feed
primarily on two saltmarsh graminoids, Puccinellia
phryganodes and Carex subspathacea. Calamaqrostis
deschampsiodies is also frequently eaten. In freshwater
areas Carex X flavicans and Carex aquatilis are common
forage sedges. These areas are intensively used from the
end of June following hatch until mid-August when the
birds begin their southward migration. Although goose
families are initially limited to feeding areas within a
few kilometers of their nest, as the goslings increase in
size families may travel further (up to 50 km) in search
of forage (McLaren and McLaren 1982; R.F. Rockwell,
unpublished data for LPB).
Geese spend 70-80% of their time feeding during the
summer (Sedinger and Raveling 1986; Lessels 1987;
Rockwell, unpublished data), and are characterised as
having continuous flow digestive systems (Sibly 1981;
Penry and Jumars 1987). Geese have small gut capacities
and relatively high metabolic rates and thus must maintain
high rates of forage intake to meet their metabolic
requirements (Demment and Van Soest 1983). Survival and
reproductive success is critically dependent on a
continual supply of energy and nutrients. Geese tend to
choose food that is relatively easy to digest, and which
45
has few mechanical or biochemical defences against
herb ivory. The optimal strategy for geese apparently is
to maximise the rate of intake of energy or more often,
some limiting nutrient (Sibly and Calow 1986).
The daily intake of an adult goose weighing 1500 to
1800 g is estimated to be 350-520 g dry weight of forage
(Cargill and Jefferies 1984b). The throughput time (or
rate of passage) in geese is estimated to be 30 to 150
minutes (Owen 1975; Burton, Hudson and Bragg 1979), and
adult geese defecate about every 5 minutes (Cargill and
Jefferies 1984a). One consequence of this rapid
throughput is that the efficency of digestion is poor
(Mattocks 1970; Demment and Van Soest 1983). Buchsbaum,
Wilson and Valiela (1986) estimate that Brant and Canada
Geese digest 36-39% of organic matter and 28% of
cellulose. These values are similar to estimates for
Lesser Snow Geese which retain 34% of the dry weight of
forage (Cargill and Jefferies 1984a). Protein digestion
is considerably higher (61-80%), however geese, like most
herbivores, have difficulty obtaining sufficient nitrogen
in their diet (Mattson 1980; Buchsbaum, Wilson and Valiela
1986; Sedinger and Raveling 1986).
Feeding preferences of geese are correlated with
forage digestability and the nitrogen content of forage
(Harwood 1974, 1977; Owen 1975, 1976; Owen, Nugent and
Davies 1977; Boudewijn 1984; Sedinger and Raveling 1984,
1986) . Buchsbaum, Valiela and Swain (1984) report that
Canada Geese avoided feeding on plants with a high
phenolic content. It is generally agreed that the
digestive physiology and feeding behaviour of geese
46
maximizes protein intake at the expense of digestive
efficiency (Sibly 1981; Sibly and Calow 1986; Buchsbaum,
Wilson and Valiela 1986), however food intake is still
limited by the processing rate (Sedinger and Raveling
1988) .
A number of potentially pathogenic parasites
(associated with feeding) are also prevelent in Lesser
Snow Geese at La Perouse Bay (Wobeser 1981; Gajadhar,
Cawthorn and Rainnie 1982; Rainnie 1982). These include
renal coccidia (Eimeria spp.), gizzard nematode
infections, typhlitis infections and a blood protozoan
parasite (Leucocytozoon simondi). These diseases are
important in the context of food resources because the
first three are transmitted via faeces. Contamination of
the limited snow free area by infected birds arriving on
the colony in spring, combined with increased
susceptability to infection at this time may result in the
increased prevalence of disease in the population (Rainnie
1982). The population dynamics of snow geese are
influenced by these parasites, which in turn are dependent
upon the colonial feeding habits of the geese.
2.4 The use of captive goslings in experimental studies
Pattern and process in natural ecosystems can only be
understood by experimentally manipulating various
components of the system. This is particularly true of
grazing systems where there are many levels of
47
interactions at all trophic levels. The experiments
discussed in the following chapters were conducted using
captive goslings. This permitted control of the level of
grazing on specific plots of vegetation and control of
nutrient input from faeces.
Goslings were collected within hours of hatch from
the nests of wild geese. The goslings were returned to
the camp and raised in captivity. Guidelines for the
experimental handling of animals in research issued by the
University of Toronto, and permit requirements for
migratory waterfowl issued by the Canadian Wildlife
Service were strictly adhered to. Goslings were raised on
a diet of Puccinellia and Carex aquatilis, supplemented by
Purina 8815 Duck Growena or dry dog food. Diet was not
supplemented on the day prior to experiments.
Goslings double in weight approximately every 7 days
(Fig. 2-4). Asymtopic weight of about 1600 g is reached
about six weeks after hatch. Differences in the growth
rate of wild and captive raised goslings have been
observed in other studies (Aubin, Dunn and Mclnnes
unpublished), but the composition of body tissues and the
feeding behavior of wild and captive goslings does not
appear to differ. Pecking rates of wild goslings are not
significantly different than those measured for captives
(Fig. 2-5). Similar seasonal patterns in growth and
pecking rate were observed by Sedinger and Raveling
(1988). Captive goslings behave much like wild goslings
in most respects, however they are much easier to
manipulate experimentally.
48
Fig. 2-4. Growth of captive goslings of the Lesser Snow
Goose at La Perouse Bay in 1987. Increases in mean weight
(ing) of between 40 and 8 goslings were determined on a
number of occasions following hatch (23 June 1987) .
G
O
SL
IN
G
 
W
EI
G
H
T 
(G
RA
M
S)
N3 O
"
o O o 5 w O
1_
o O) o
O o
_
J
O o o
J
 
L
J
 
L
01 o o
o o o
49
Fig. 2-5. Comparison of pecking rate of wild (o) and
captive (•) goslings at La Perouse Bay. Pecking rates for
captive goslings were determined for Puccinellia
vegetation during the summer of 1987. Rates for wild
goslings were measured on mixed Puccinellia-Carex sward
(see Lessels 1987). Lines fitted by eye.
*
(/
k
A.
o
53
P
E
CK
IN
G
 
R
A
T
E
 
(P
EC
KS
 
*
? 
.
 
§
s
ro O O
o 0 M 2! O M t~
\
00
A. O Wl
-
O
\
I I '•
50
CHAPTER 3: WITHIN SEASON INCREASES IN THE NET ABOVE-GROUND
PRIMARY PRODUCTION OF A SALT MARSH FORAGE GRASS: A
TEST OF THE HERBIVORE-OPTIMIZATION MODEL
3.1 INTRODUCTION
Plant-herbivore interactions cannot be considered in
isolation from the ecosystems in which they occur.
Herbivores affect primary and secondary production, energy
flow and nutrient cycling, and the structure and
composition of plant communities (Harper 1969; Crawley
1983; Vitousek 1984). The herbivore optimization model
(Dyer 1975, McNaughton 1979, 1983a,b; Hilbert et al. 1981;
Dyer et al. 1982; Jefferies 1988a), predicts that one of
the effects of herbivory on plant production is the
enhancement of net primary production of forage plants
above that of ungrazed plants, at moderate levels of
herbivory. Circumstantial evidence in support of this
prediction has come primarily from studies of the
responses of terrestrial graminoid communities (McNaughton
1976, 1979; Prins, Ydenburg and Drent 1980; Cargill and
Jefferies 1984b) and aquatic algal and coral communites
(Ogden and Lobel 1978; Bjorndahl 1980; Bergquist and
Carpenter 1986; Carpenter 1986; Power, 1987) to herbivory.
The generally applicability of these results remains
highly controversial (Stenseth 1978, 1983; Belsky 1986a,
1987; McNaughton 1986a; Crawely 1987).
51
Intrinsic mechanisms that operate at the level of
individual plants and/or extrinsic mechanisms that involve
processes at the ecosystem level have been proposed to
account for the increase in net primary production of
forage plants at moderate intensities of herbivory
(McNaughton 1983a,b). These include photosynthetic
compensation, reallocation of resources for growth,
changes in morphology and leaf turnover rates, and
fertilization by faeces and urine.
We conducted a critical test of the herbivore
optimization model using captive goslings of the Lesser
Snow Goose (Chen caerulescens caerulescens L.) which
grazed on graminoid vegetation of a sub-arctic salt marsh.
Plant growth in this marsh is strongly nitrogen-limited
(Cargill and Jefferies 1984a). Previous studies have
shown that grazing by snow geese significantly increased
the net above-ground primary production (NAPP) of a
stoloniferous salt-marsh grass, Puccinellia phryganodes
(Trin.) Scribn. and Merr., and a rhizomatous sedge, Carex
subspathacea Wormsk., from 30% to 105%, depending upon the
year, compared to corresponding values in ungrazed swards
of these two species (Cargill and Jefferies 1984b; Bazely
1984). A consequence of the intense grazing is that plant
tissue is converted into either goose biomass or faeces,
instead of accumulating as live plant biomass or litter.
Nitrogen released from goose faeces accelerates the supply of
this element to the sediments. In addition, patches of
sediment (0.5 cm diameter) are colonized by cyanobacteria
that contribute to the long-term input of nitrogen, as a
result of nitrogen fixation (Bazely and Jefferies 1988).
52
Detailed demographic studies have shown that plants
of Puccinellia phryganodes are capable of increasing the
production of shoots and leaves in resonse to grazing
(Bazely and Jefferies 198N). In contrast, plants of Carex
subspathacea increase leaf production following grazing
(Kotanen and Jefferies 1987). The persistence of the
grazed graminoid swards is strongly dependent on the
colonial feeding behaviour of the geese (Bazely and
Jefferies 1986). There is selection for biotypes of
Puccinellia with faster growth rates under grazed
conditions (Sadul 1987). The ability of both Puccinellia
and Carex to show rapid growth responses within the season
compensates for the adverse effects of defoliation per se
on net above-ground primary production (Sadul 1987;
Jefferies 1988a). A continual supply of high quality
forage is produced during a period when it is most
required by the geese.
Although it is evident that grazing by snow geese
leads to enhanced NAPP of the grazed sward,
interrelationships between intensity of clipping,
fertilization from faecal nitrogen, and forage regrowth
and quality, can only be determined experimentally under
controlled conditions. We have used captive goslings to
investigate the dynamics of the overall effects of grazing
(grazing, addition of faeces, trampling) on plant
production on the salt marsh flats at the La Perouse Bay.
In this study the following questions have been addressed.
1. What is the effect of increasing periods of grazing
(grazing intensity) on NAPP and forage quality?
53
2. Does grazing affect NAPP and forage quality in the
same way throughout the summer?
3. If grazing results in the enhancement of NAPP what
mechanism(s) account for this, and how is this related to
the intensity of herbivory? Faeces may be either
left on or removed from the grazed plots to assess
the importance of nutrients in faeces to the
regrowth of vegetation following grazing.
3.2 METHODS
3.2.1 Study site
Over 8000 pairs of Lesser Snow Geese breed each
summer at La Perouse Bay, 25 km east of Churchill,
Manitoba, Canada, (58°, 24' N, 94°, 24' W), on the west
coast of Hudson Bay. Following the hatch of goslings in
late June or early July, the geese forage intensively on
the vegetation of the intertidal salt marsh, dominated by
either Puccinellia phryganodes, a stoloniferous grass, or
Carex subspathacea, a rhizomatous sedge which is abundant
in less saline areas. The intertidal grazing lawn is
about 2.5 cm high. There are over 4000 shoots of these
species per square meter and very few shoots (< 5%)
escaped the effects of grazing (Kotanen and Jefferies
1987; Bazely and Jefferies 198N). By the second week of
August the geese begin their autumn migration and leave
the salt marsh. Permanent snowfall and the cessation of
54
plant growth occurs in late September. A complete
description of the study area is provided in Jefferies,
Jensen and Abraham (1979) and Jefferies (1988a).
3.2.2 Experimental trials
All experiments were conducted using captive
goslings. The goslings were removed from nests of wild
geese within 24 hours of hatching and were raised in
captivity in the research camp. Goslings were fed a diet
of natural foods (mainly Puccinellia phryganodes and Carex
aquatilis) , supplemented with a daily ration of Purina
8815 Duck Growena, or dry dog food, except on the days
prior to an experiment. Prior to each experiment the
weight of the goslings was determined (Table 3-1).
Depending upon the number of goslings required for each
experiment, those which were greater than 1 S.E. (larger
or smaller) than the mean in weight were excluded from the
grazing trials.
Experimental plots were established prior to each
experiment on selected areas of saltmarsh vegetation
dominated by Puccinellia. Each plot was 1m x 1m. Four
replicates for each grazing treatment were established in
a completely randomized block design. On the morning of
the experiment, goslings were randomly assigned to a plot
and allowed to graze for a specific period. The number of
goslings per plot and the period of grazing was adjusted
during the season as the goslings increased in weight and
their foraging efficency increased (Table 3-1). Grazing
times varied from 0 to 150 minutes at intervals of 15 or
55
TABLE 3-1. Dates on which grazing experiments were
conducted at La Perouse Bay, Manitoba and mean gosling
weights and number of goslings grazed per plot for each
experiment.
Experiment
1
2
3
4
5
6/7
8/9
date
23/6/86
2/7/86
13/7/86
1/8/86
10/8/86
1/7/87
11/8/87
mean gosling wt (g)
130
371
539
1273
1420
247
1478
goslings per plot
4
4
3
3
2
4
2
56
30 minutes. The extreme grazing levels (120 and 150
minutes) exaggerated the overall effects of grazing well
beyond natural levels. At the end of the grazing trials
all plots were exclosed with chicken wire to prevent
further grazing by wild geese.
Five experiments were conducted in 1986 (23 June; 2,
17 July; 4, 11 August). The feeding activity of the
goslings was monitored at intervals of 10 or 15 minutes
during each experiment. Goslings spent at least 90% of
their time feeding. In 1987 Experiments 6/7 and 8/9 were
conducted on 1 July and 12 August respectively. Paired
plots (1m x 1m) were established on a Puccinellia sward.
Both sets of plots received .identical grazing treatments,
however in one set of plots faeces were removed at
intervals of ten minutes during the experiments. This was
to ensure that nitrogen was not transfered from the faeces
into the plants or sediments. At the end of the
experiments the number of faeces deposited on each plot
was counted, and some of these were randomly selected, and
returned to the laboratory for determinations of their
water content and dry weight, and amounts of soluble and
total nitrogen present in individual droppings.
3.3.3 Sampling and data analysis
Above-ground biomass was sampled by removing a 7.5 x
7.5 cm turve at random from each of the plots before
grazing, after grazing, and at intervals of approximately
12 days for the duration of the summer. The above-ground
57
biomass was removed by clipping the sward at ground level.
Since approximately 90% of the biomass was live, dead
material was not removed prior to biomass or nitrogen
determinations. The plant material was washed and then
dried at 60-80°C for 24 hr. Dry weights of the material
were determined using a Mettler (PK-300) electronic
balance. Determinations of amounts of carbon and nitrogen
present in dried faeces and plant tissues were made using
a LECO series 600 CHN autoanalyser (St. Joseph, MI., USA).
All material was ground (20 mesh size) in a Wiley mill and
appropriate quantities (approx. 100 mg) were preweighed
before combustion in the analyser. Soluble nitrogen of
fresh goose faeces was determined in 1987 using the
phenol-sodium hypochlorite method (Solorzano 1969) after
extraction over 12 h with a solution of 1 M KC1.
Approximately 6 g of fresh goose droppings were suspended
in 40 ml of solution.
Analysis of Variance (ANOVA) was used to assess all
treatment effects. Data sets were transformed prior to
analysis if variance was heterogenous. Significant
interaction terms of two-way ANOVA's were investigated
using trend analysis (Kirk 1982). Using this approach,
the between-group sum of squares can be partitioned into a
number of trend components (linear, quadratic, cubic)
reflecting orthogonal trend contrasts. The efficacy of
the trend components was measured by CJA, the strength of
association. This statistic was calculated in conjunction
with the trend analyses (Kirk 1982; Keppel 1982). The
magnitude of the effect measured by cj* is considered to be
large when GJ*> 0.15, and moderate when to* > 0.06 (Keppel 1982)
58
3.3 RESULTS
3.3.1 Changes in above-ground biomass following grazing
The ability of Puccinellia swards to regrow following
grazing late in the spring (23 June 1986) is shown in Fig.
3-1. At 30, 60 and 90 minutes of grazing significant
increases in above-ground biomass compared to that of
ungrazed plots were detected after 24, 36 and 48 days of
regrowth. At the extreme grazing levels (120 and 150
minutes), no enhancement of growth is evident, possibly
because of damage to meristems as a result of grazing and
trampling. The results of a two-way ANOVA indicate that
both main effects are significant (harvest time: F[5,108]
= 66.98, p < 0.001; grazing treatment: F[5,108] = 11.76, p
< 0.001). In addition the time x treatment interaction is
significant (F[25,108] = 2.15, p < 0.004). This
interaction can be explained by the change in the shape of
the growth response curves from a linear to an approximate
quadratic function 24, 36 and 48 days after grazing. Over
half of the variance of an a posteriori trend analysis
(Kirk 1982) can be explained by a quadratic term after 24,
36 and 48 days of regrowth (71%, 85% and 53% respectively;
W1> 0.15) .
The ability of Puccinellia to recover from the
effects of grazing apparently decreases over the course
of the summer (Fig. 3-2). With each successive experiment
the maximum amount of regrowth of the Puccinellia sward
occurs in plots which have received shorter periods of
grazing. In Experiment 2 (2 July) the greatest amount of
59
FIG. 3-1. Increase in above-ground biomass of Puccinellia
phryganodes following grazing on 23 June, 1986 by goslings
of the Lesser Snow Goose in Experiment 1 (mean values of 4
plots). Each plot was 1 m2 in which four goslings grazed.
Error bar is Tukey's honestly significant difference.
Before grazing (x); after grazing (A); 12 days regrowth (n);
24 days regrowth (•); 36 days regrowth (o); 48 days regrowth
o5
°
O D O
 
o>
TI
 
o
O X) > o ? CO
01
 
-
o
A
B
O
VE
-G
R
O
U
N
D
 
BI
O
M
AS
S 
(G
-D
W
T-
M
'2)
-
»
• 
ro
 
c>
j
0
0
0
o
01 o
o
o
\0
o
CD O
1
60
regrowth of vegetation occurred in plots which had been
grazed for 15, 30 and 60 minutes, whereas in Experiment 3
(17 July), the standing crop was highest in the plots
grazed for 15 and 30 minutes. As in Experiment 1, the
time, treatment and interaction terms of the ANOVA are
significant for both these experiments (Table 3-2).
Although there is tendency for the curves of above-ground
biomass to become unimodal, the quadratic term is not
significant for any interval following grazing in
Experiments 2 and 3. The proportion of the variance
explained by a quadratic term in Experiment 2 after 24 and
36 days regrowth is only 14% and 18% respectively (co1 =
0) . In Experiment 3 the variance explained by the
quadratic term after 24 and 48 days regrowth is 12% and
18% respectively ( &*•= 0) .
Significant regrowth of the Puccinellia in excess of
values for the standing crop of ungrazed vegetation was
not detected for grazing Experiments 4 and 5 conducted in
August 1986. Although in these two Experiments the plots
were sampled only at 12 and 24 days, the last harvest was
taken just two or three weeks before "freeze-up",
consequently further significant increases in biomass were
unlikely to have occurred. In Experiments 4 and 5, the
time x treatment interaction term is not significant,
although both the main effects (time treatment) show
significant differences (Table 3-2). These results
reflect the amount of biomass consumed by grazing and not
the regrowth of Puccinellia following grazing.
61
FIG. 3-2. Incease in above-ground biomass of Puccinellia
phryganodes following grazing by goslings in Experiment 2,
2 July 1986 (a); Experiment 3, 13 July 1986 (b);
Experiment 4, 1 August 1986 (c); Experiment 5, 10 August
1986 (d). Values shown are mean values of 4 plots for
Experiments 2, 3, 4 and 3 plots for Experiment 5. Each
plot was 1 mx, in which 4 goslings grazed in Experiment 2,
3 goslings grazed in Experiments 3 and 4, and 2 goslings
grazed in Experiment 5. Error bar is Tukey's honestly
significant difference. Before grazing (x); after
grazing (A); 12 days regrowth (D); 24 days regrowth (•);
36 days regrowth (o); 48 days regrowth (•); 60 days
regrowth (v).
110
too
90
80
CO
CO
O
03
Q
O
cr
o
UJi
00
60
50
40
30
20 -
10 -
130
120
110
I
ti
CO
CO
O
CD
10O
90
80
O
cr
o
UJi
CD
60
50
A
B
O
V
E
-G
R
O
U
N
D
 
B
IO
M
A
S
S
 
(G
-D
W
T
 
M
'*
)
2 o o o N Z CD I
 
8
O
"
O m 5 o o fl I«
A
B
O
V
E
-G
R
O
U
N
D
 
B
IO
M
A
S
S
 
(G
-D
W
T-
M
"2
)
AB
OV
E-
GR
OU
ND
 
BI
OM
AS
S 
(G
-D
W
T-
M'
2)
"
O rn 2 O 8«
p
T> m § 
»
o o INI O —
 
<
£l
in
AB
OV
E-
GR
OU
ND
 
BI
O
M
AS
S 
(G
-D
WT
-M
-2)
K 
"
 
*
 
01
9
 
o
 
o
 
o
cr
62
TABLE 3-2. Summary of results of a two-way ANOVA for
increases in above-ground biomass in Experiments 2, 3, 4,
and 5 (Fig. 3-2). Main effects are harvest times during
the summer (T), and grazing treatment (G), and the time x
treatment interaction (TG).
Exp 2
Exp 3
Exp 4
Exp 5
Effect
T
G
TG
T
G
TG
T
G
TG
T
G
TG
F-ratio
135.66
12.94
2.79
25.02
17.60
2.10
8.20
7.91
1.12
11.23
11.43
1.87
df
5,108
5,108
25,108
4,90
5,90
20,90
3,72
5,72
15,72
3,48
5,48
15,48
P
<.001
<.001
<.001
<.001
<.001
<.020
<.001
<.001
>.353
<.001
<.001
>.051
63
3.3.2 Increase of NAPP of swards of Puccinellia
following grazing
Cumulative NAPP of Puccinellia phryqanodes for each
grazing treatment was calculated by summing positive
increments of above-ground biomass between harvests for
each of the five experiments. There were significant
increases in the NAPP of the swards following grazing, in
Experiments 1, 2, and 3 (Table 3-3). In Experiment 1 the
greatest increases in NAPP occurred in plots which were
grazed for 30, 60 and 90 minutes. Cumulative NAPP in these
plots was approximately 40% greater than that in the
ungrazed plots. Corresponding values for NAPP in plots
which were grazed for 120 or 150 minutes were less than
the mean value for ungrazed plots.
A similar pattern of increases of NAPP is evident in
Experiment 2, in which the largest increases occurred
where plots were grazed for 15, 30 and 60 minutes (35-45%
greater than that of ungrazed plots). There were no
significant increases in NAPP beyond 36 days, irrespective
of the grazing treatment. In Experiment 3 the greatest
increase was evident in plots grazed for 15 and 30
minutes (the values were approximately 70% greater than
the value of ungrazed plots). The NAPP of grazed swards
was not significantly greater than that of ungrazed swards
in Experiments 4 and 5 after 24 days of regrowth.
64
TABLE 3-3. Cumulative net above-ground primary production
(g DWT * m~a) for each harvest of experimental plots in
Experiments 1 to 5. Calculated as positive increases in
dry weight of above-ground biomass between successive
harvests. Significant differences in total NAPP between
that of ungrazed plots (control) and values for grazed
plots for each treatment (p < 0.05) are indicated (*).
Harvest
(days)
EXP 1
EXP 2
EXP 3
EXP 4
EXP 5
12
24
36
48
12
24
36
60
12
24
48
12
24
12
24
0
16
16
19
32
30
30
46
46
9
11
11
1
15
1
7
15
33
46
73
73*
4
37
37*
9
23
7
19
Period
30
12
27
47
54*
32
51
83
83*
8
45
45*
0
13
1
5
of grazing (min)
60 90 120
26
36
48
57*
28
46
76
76*
13
25
25
17
19
4
5
21
40
40
55*
14
25
45
57
15
26
26
20
29
9
9
9
20
39
39
15
34
60
60
8
19
19
6
6
10
13
150
16
16
28
32
65
3.3.3 Forage quality and nitrogen content of shoots
Grazing early in the season (Experiment 1) not only
resulted in increased NAPP at intermediate grazing
treatments, but also led to higher amounts of nitrogen
present in above-ground tissue compared to corresponding
values for ungrazed plots. Fig. 3-3a indicates that the
nitrogen content of above-ground biomass was significantly
higher in the grazed plots than in ungrazed plots.
Although the nitrogen content of plant tissue declined
over the course of the summer (from about 4.5% to 2.8%),
the rate of decline in shoots from grazed plots was less
than that for shoots from the ungrazed plots, mainly
because no decline occurred in the first 12 days following
grazing. Statistical analysis revealed no significant
interaction between sampling time and grazing treatment
(Table 3-4), but both of the main effects were
significant. Carbon content of plant tissues did not
change over the course of the season, and there were no
differences in the amounts of carbon in tissues between
swards subject to the various grazing treatments (Fig. 3-
3a) .
The pattern of changes in amounts of nitrogen and
carbon as a percentage of the dry weight for the different
treatments in Experiments 2 and 3 were similar to those
for Experiment 1 (Fig. 3-3b,c). However, the nitrogen
contents of the shoots, prior to grazing, declined
throughout the season (4.5%, 2.8%, 2.5% for Experiments 1,
2, and 3 respectively). In Experiment 2 neither of the
main effects were significant (Table 3-4).
66
FIG. 3-3. Total nitrogen (% N) and carbon (% C) content
as a percentage of the dry weight of above-ground biomass
of Puccinellia phryganodes following grazing by goslings
in Experiments 1 (a), 2 (b), 3 (c), 4 (d), and 5 (e),
(mean values are shown, n = 3 for Experiment 1; n = 2 for
Experiments 2 to 5). Above-ground biomass with a nitrogen
content significantly greater than that from ungrazed
swards at each harvest are circled (P < 0.05; SNK
multiple range test). Control (ungrazed): 0 mins
grazing (o); 15 mins grazing (v); 30 mins grazing (n);
60 mins grazing (•); 90 mins grazing (A); 120 mins
grazing (•); 150 mins grazing ( + ).
5 4
Z
 
3
5*
 
2 1 0 4 3
#
 
2 1 0 4 3
Z #
 
2 1 0 4 3
?
 
2 1 0 4 3
2
1 0
*5D
B
m If] o* CV n Sr a* Ov
a
12
 
24
 
36
 
48
DA
YS
 
OF
 
RE
GR
OW
TH
i 60
O O
45
|-
40 50 45 40
D
*
• 8 i
• A § • 1
a A 8* • i
OS + 1
• •
 
A
o ? J.
50
 r
O
45 50
O
45 50
o
45 40
3* 12
24
36
4
8
60
DA
YS
 
OF
 
RE
GR
OW
TH
67
TABLE 3-4. Summary of results of a 2-way ANOVA of amounts
of nitrogen expressed as a percentage (%N) of the dry
weight of above-ground biomass in Experiments 1 to 5 (Fig.
3-3). Main effects are harvest times during the summer
(T), and grazing treatment (G), and the time x treatment
interaction (TG).
Exp 1
Exp 2
Exp 3
Exp 4
Exp 5
Effect
T
G
TG
T
G
TG
T
G
TG
T
G
TG
T
G
TG
F-ratio
20.06
2.59
0.83
1.67
1.26
0.82
20.06
2.59
0.83
24.29
9.02
1.08
1.09
4.39
1.17
df
4,60
5,60
20,60
4,30
5,30
20,30
3,24
5,24
15,24
2,18
5,18
10,18
2,18
5,18
10,18
P
<.001
<.035
>.672
>.183
>.308
>.679
<.001
<.035
>.672
<.001
<.001
>.428
>.360
>.009
>.372
68
In Experiment 4 there were slight declines in
amounts of nitrogen as a percentage of the dry weight in
shoots in all plots during the course of the Experiment,
but the decrease was relatively small (0.5%) (Fig. 3-3d).
In Experiment 5 no differences in the amounts of nitrogen
present in shoots from swards subject to a particular
grazing treatment were evident (Fig. 3-3e; Table 3-4).
The nitrogen content of the above-ground biomass averaged
about 2.5% of the dry weight in both Experiments 4 and 5.
Amounts of carbon as a percentage of the dry weight showed
no significant time, treatment or interaction effects for
any Experiment (2-way ANOVA; p > .20). Carbon content of
tissue averaged approximately 45% of dry weight throughout
the season in all experiments.
The cumulative net amounts of nitrogen present in
above-ground biomass from the different plots are shown in
Fig. 3-4 for Experiments 1 to 5. These results were
obtained by multiplying the increment of live, above-
ground biomass for each interval between successive
harvests by the average nitrogen content of above-ground
biomass during that period. In general, the cumulative
net amounts of nitrogen were greatest in shoots from plots
which received the intermediate grazing treatments. Late
in season (Experiments 4 and 5) the rate of accumulation
of nitrogen in shoots was lower than that measured earlier
in the season for comparable periods (Experiments 1 and
2). In those treatments in which the goslings grazed the
swards for 120 and 150 minutes the swards contained low
amounts of nitrogen, because the NAPP in these plots was
low (Table 3-3).
69
FIG. 3-4. Net amounts of nitrogen (g * itr*) accumulated
in the regrowth of above-ground biomass of swards of
Puccinellia phryganodes following grazing in Experiment 1
(a); Experiment 2 (b); Experiment 3 (c); Experiment 4 (d);
and Experiment 5 (e). Control (ungrazed): 0 mins grazing
(O); 15 mins grazing (v); 30 mins grazing (n); 60 mins
grazing (o); 90 mins grazing (A); 120 mins grazing (•);
150 mins grazing (+).
N
E
T
 
A
M
O
U
N
T
S
 
(g
) 
O
F
 
N
IT
R
O
G
E
N
 
IN
 
A
B
O
V
E
-G
R
O
U
N
D
 
R
E
G
R
O
W
T
H
 
O
F
 
S
W
A
R
D
S
 
(M
2
)
OF
 
PU
CC
IN
EL
L/
A 
FO
LL
O
W
IN
G
 
GR
AZ
IN
G
ro
CD
70
The amount of nitrogen available in above-ground biomass
per unit area from moderately grazed plots after 36 and 48
days was almost twice that of ungrazed (control) plots in
Experiments 1 and 2 (Fig. 3-4a,b). This difference was
smaller in the other Experiments (Fig. 3-4c,d,e).
3.3.4 Effect of faeces on the growth of swards of
Puccinellia following grazing
Although the overall effects of grazing can
significantly increase the subsequent growth of Puccinellia
(Fig. 3-1 and 3-2), the experimental design and the results
do not allow the separation of the relative effects on plant
growth of the addition of nutrients from faeces or those
caused by clipping. On 1 July 1987 two further grazing
Experiments were conducted. The two Experiments were
similar, except that while the geese were grazing faeces
were removed from one set of plots every ten minutes
(Experiment 7). In the adjacent group of paired plots the
faeces remained (Experiment 6). The results show that
clipping without the concomitant deposition of faeces did not
lead to increased growth evident when faeces remained in
plots (Fig. 3-5, Table 3-5a). Clipping alone did not result
in increases in production beyond that of ungrazed plots.
This is most clearly indicated by the significant faeces x
treatment interaction (p < .017; Table 3-5a).
After 60 days of regrowth following grazing the
standing crop in Experiment 6 was approximately 160 g DWT
71
FIG. 3-5. Increase in above-ground biomass of
Puccinellia phryganodes following grazing by goslings of
the Lesser Snow Goose in Experiment 6 with faeces
remaining (a), and Experiment 7 with faeces removed (b)
(mean value for four plots (1 m2) grazed by 4 goslings).
Error bar is Tukey's honestly significant difference.
Before grazing (x); after grazing (A); 12 days regrowth
(D); 22 days regrowth (•); 37 days regrowth (O); 48 days
regrowth (•); 60 days regrowth (v)•
O)
o
(0
o
O
D
o
N
Z
Ol
o
o -
Ol
o
8
ABOVE-GROUND BIOMASS
(G-DWT-M-2)
Ol
o
CD
O O
Ul
o
CO J.
o o
Ol
o
Ol
o
--J
o
10
o
CD
O
73
* m"a in plots which were grazed for 39, 60 and 90
minutes, compared to a corresponding value of
approximately 130 g DWT * irr3- in Experiment 7. The curve
of standing crop of Puccinellia for the different
treatments in Experiment 6, 48 days after grazing (Fig. 3-
5a) , approximates to a quadratic form ( cj*> 0.09). This
type of growth response occurred 24 days later than that
measured in the previous year (Fig. 3-1). The difference
in the timing of response is most probably due to seasonal
and site differences.
On 11 August 1987 a similar grazing Experiment with
faeces present (Experiment 8) and faeces removed
(Experiment 9) was conducted. The results indicated only
a slight increase in the growth of swards of plots that
were grazed for 15, 30 and 45 minutes, compared with
growth in corresponding plots from which faeces were
removed (Fig. 3-6; Table 3-5b). The difference (95 g DWT *
m"1 compared to 85 g DWT * m'*) was not significant.
After 65 days following grazing (mid-October) the above-
ground biomass had decreased by almost 30%. The faeces x
treatment interaction was not significant (p > .281; Table
3-5b), hence the onset of winter precluded the
establishment of significant differences in the growth of
swards with respect to the different treatments.
3.3.5 Effect of faeces on NAPP
The NAPP of plots with faeces (Experiment 6) showed
significant increases compared to values for plots from
which faeces were removed (Experiment 7; Table 3-6). At
74
FIG. 3-6. Increase in above-ground biomass of
Puccinellia phryganodes following grazing by goslings of
the Lesser Snow Goose in Experiment 8 with faeces
remaining (a), and Experiment 9 with faeces removed (b)
(mean value of 4 plots (1 m2) grazed by 2 goslings). Error
bar is Tukey's honestly significant difference. Before
grazing (x); after grazing (A); 12 days regrowth (a); 24
days regrowth (•); 65 days regrowth (+).
ABOVE-GROUND BIOMASS (G-DWT-M" 2 )
01
t)
m
2 oO
o
N
O O
O3 __
"— 01
o<\ \
s
A
o
Ol
o 01o o 8 8
o
Ol
o
O)
o
CD
O 8 8
75
TABLE 3-5b. Summary of results of 3-way and 2-way ANOVA
of changes in above-ground biomass of Puccinellia
phryqanodes following grazing in Experiments 8 and 9 (Fig.
3-6). Main effects are faeces (F), harvest time (T), and
grazing treatment (G).
Exp 8/9
Exp 8
Exp 9
Effect
F
T
G
FT
FG
TG
FTG
T
G
TG
T
G
TG
F-ratio
1.56
31.23
13.27
0.65
1.26
1.71
0.24
7.95
6.19
1.22
8.44
4.78
1.16
df
1,180
4,180
5,180
4,180
5,180
20,180
20,180
4,72
5,72
20,72
4,72
5,72
20,72
P
<.001
<.001
>.634
>.281
<.025
>.999
<.001
<.001
>.279
<.001
<.002
>.326
76
intermediate grazing levels NAPP was 30 to 40% greater in
Experiment 6; the highest values for NAPP were detected
in plots where the goslings grazed for 30, 60 and 90
minutes. The values were approximately 35% greater than
that for ungrazed swards. The values of NAPP of swards
subject to the different grazing treatments in Experiment
7 were not significantly different from each other, or
from the NAPP of ungrazed plots after 60 days of regrowth
(Table 3-6). There were no differences between the NAPP
of plots in Experiments 8 and 9, irrespective of the
treatment. In both of these experiments, however, NAPP
was largest for swards which had been grazed, and lowest
for ungrazed plots (Table 3-6).
3.3.6 Nitrogen content of above-ground biomass following
grazing in the presence and absence of faeces
There were no significant differences in the nitrogen
(or carbon) content of above-ground biomass of grazed
plants in the presence or absence of faeces (Experiments 6
and 7; Fig. 3-7; Table 3-7). As in Experiment 1, the rate
of decrease in the amount of nitrogen as a percent of the
dry weight was least in tissue from grazed plots where
faeces remained. Nevertheless, no significant differences
in the amount of nitrogen as a percent of the dry weight
in plant tissues from Experiments 6 and 7 were detected (p
> .713; Table 3-7). Initial amounts of nitrogen, which
were about 2.8%, decreased to about 1.5% after 60 days.
77
TABLE 3-6. Cumulative net above-ground primary production
(g DWT * m'1 ) for each harvest of grazed plots in
Experiments 6 and 7, and 8 and 9. Significant differences
in total NAPP between the ungrazed (control) plots and
plots which received the different grazing treatments (p <
.05) are indicated (*).
Harvest Period of grazing (min)
(days)
EXP 6
EXP 7
EXP 8
EXP 9
12
22
37
48
60
12
22
37
48
60
12
24
65
12
24
65
0
24
38
39
51
67
39
39
42
56
64
0
5
5
5
1
3
3
30
26
39
56
68
91*
29
29
29
48
64
15
23
25
25*
14
14
14
60
38
46
65
106
106*
13
13
38
59
61
Period
30
16
30
30*
22
24
24*
90
40
48
64
85
101*
23
27
51
67
78
of grazing
45
19
27
27*
14
15
15
120
31
35
54
74
82
8
21
39
57
57
(min)
60
17
17
17
11
11
11
150
30
30
42
55
72
16
20
37
44
61
75
11
12
12
15
15
15
78
FIG. 3-7. Total nitrogen and carbon content as a
percentage of the dry weight of above-ground biomass of
Puccinellia phryganodes following grazing by goslings in
Experiments 6 (a,c) and 7 (b,d), (mean values given, n =
2). Above-ground biomass with a nitrogen content
significantly greater than that from ungrazed swards at
each harvest are circled (p < 0.05; SNK multiple range
test). Control (ungrazed): 0 mins grazing (o); 30 mins
grazing (n); 60 mins grazing (•); 90 mins grazing (A);
120 mins grazing (•); 150 mins grazing (+).
NITROGEN CONTENT (% DRY WEIGHT)
ro tji
<0
(f)
O roPO
m
H
Q)
CARBON CONTENT (% DRY WEIGHT)
O
in
O roro
m
o3i
a* y*m
+DO,
o
n o
•f •
DO
79
TABLE 3-7. Summary of results of 3-way and 2-way ANOVA of
amounts of nitrogen expressed as a percentage of the dry
weight of above-ground biomass of Puccinellia phryganodes
for Experiments 6 and 7. Main effects are faeces (F),
harvest time (T), and grazing treatment (G).
Effect F-ratio df
Exp 6/7 F
T
G
0.14
121.63
7.67
1,72
5,72
5,72
.713
<.001
<.001
ALL INTERACTION TERMS NON-SIGNIFICANT (p > .60)
Exp 6 T
G
TG
70.56
6.52
0.83
5,36
5,36
25,36
<.001
<.001
.689
Exp 7 T
G
TG
54.65
2.43
0.62
5,36
5,36
25,36
<.001
.054
.897
80
The amount of carbon in the tissues increased slightly
from 42.5% to 44.5% of the dry weight after 60 days.
The amount of nitrogen deposited in the above-ground
biomass in Experiments 6 and 7 was calculated by
multiplying the increment of live, above-ground biomass in
each interval between successive harvests by the average
total nitrogen content of above-ground biomass during that
period (Fig. 3-8). In both experiments the amount of
nitrogen accumulated in the above-ground biomass in the
ungrazed control after 60 days was approximately 1.5% of
the dry weight. In the presence of faeces all swards
accumulated more nitrogen per unit area than the
corresponding value for ungrazed swards (Fig. 3-8a). In
the absence of faeces the total amount of nitrogen per
unit area present in grazed swards was less than that for
ungrazed swards (Fig. 3-8b).
3.3.7 Accounting for nitrogen accumulated in shoots
The amount of above-ground biomass consumed by the
grazers was not linearly related to the length of the
grazing period (Fig. 3-9a). The amount of above-ground
biomass consumed by the geese (% offtake) levelled off at
about 28% of the initial standing crop after 90 minutes of
grazing. Continued grazing and trampling probably damaged
meristems. The input of nitrogen from faeces in the
Experimental plots was significantly and positively
related to the length of the grazing period and the amount
of live biomass comsumed (% offtake) (Fig. 3-8a;
81
FIG. 3-8. Net amounts of nitrogen (g * m~*) accumulated
in the regrowth of biomass of swards of Puccinellia
phryganodes following grazing in Experiment 6 (a), and
Experiment 7 (b). Control (ungrazed): 0 mins grazing
(O); 30 mins grazing (a); 60 mins grazing (•); 90 mins
grazing (A); 120 mins grazing (•); 150 mins grazing (+).
NET AMOUNTS (g) OF NITROGEN IN
ABOVE-GROUND REGROWTH OF SWARDS (M2)
OF PUCCINELLIA FOLLOWING GRAZING
p -» -» ro no
o 01 b bi b en
I
82
r = 0.856, p < 0.001), and ranged from 32 ing N - g DWT *
nr* to 192 mg N - g DWT * m'1.
The amount of nitrogen was 1.71 + .10% of the dry
weight of faeces. Approximately 34.4% was in a soluble
form (mainly NH£ -N). It is possible to calculate the
total input of faecal nitrogen into each grazed plot based
on the number of faeces deposited during the grazing
period, the dry weight of faeces, and the nitrogen content
of faeces. The net amount of nitrogen which is
incorporated in above-ground biomass (g * m'* ) directly
from faecal nitrogen, or from the translocation of
nitrogen from below-ground reserves or as a result of
nitrogen fixation by cyanobacteria on the sediment
surface, can be calculated (Experiment 6). From these
values the corresponding values of the total amount of
nitrogen accumulated in above-ground biomass (g * m~*) in
plots where all these sources of nitrogen are available,
except the presence of faeces (Experiment 7), can be
subtracted. If the amounts of nitrogen (g * m'*) in
swards where faeces remained are in excess of the values
for amounts in swards in the absence of faeces the
difference may be balanced by the nitrogen supplied in the
faeces (Table 3-8). In fact, the differences in amount
are in excess of that predicted from assuming that all the
nitrogen in faeces is available for plant growth (Table 3-
8) . The values of of amounts of nitrogen unaccounted for
range from 1.1% to 44.3% of the nitrogen deposited in the
above-ground biomass in Experiment 6 (14 to 1016 mg N * m"*)
83
FIG. 3-9. Relationship between length of the grazing period
and the amount of above-ground biomass removed as a
percentage of the initial amount present (% offtake) (a).
Bars are mean + SEM, n = 2); and the relationship between %
offtake and the amount of nitrogen present in faeces (mg N *
m~A) deposited on plots (b).
TO
TA
L 
N 
DE
PO
SI
TE
D 
ON
 
PL
OT
S 
(m
gN
-M
'2)
%
 
O
FF
TA
KE
o
^ O m
o
 
8 
o
 
8 
8
 
8
T
l m 33 O O o -n CD N Z CD
ot
84
TABLE 3-8. Summary of net amounts of nitrogen accumulated
in above ground biomass of swards of Puccinellia
phryganodes in Experiments 6 and 7 (Fig. 3-8) and
calculation of the proportion of nitrogen available from a
number of sources to account for the total nitrogen
accumulated after 60 days of regrowth.
A) Total N deposited in above-ground biomass (Experiment 6
with faeces remaining).
B) Total N available from faeces (assuming 100%
mineralization and no ammonia volatilization).
C) Total N deposited in above-ground biomass (Experiment 7
with faeces removed); attributable to below-ground
reallocation and N fixation at the sediment surface and
net mineralisation of organic nitrogen in the sediments in
the absence of faeces.
D) Amount (and percentage) of nitrogen (A) unaccounted for
by (B) and (C).
nig N * m
A)
B)
C)
D)
0
1144
0
1673
0
(0%)
30
1759
33
1270
456
(26%)
~* at each grazing treatment
60
2296
58
1222
1016
(44%)
90
2042
107
1557
378
(19%)
120
1751
185
1160
406
(23%)
150
1328
191
1123
14
(1%)
85
3.4 DISCUSSION
The main conclusions of these Experiments are, 1)
that grazing by goslings of the Lesser Snow Goose leads to
an increase of NAPP at low to moderate levels of grazing
early in the season, compared to that of ungrazed swards;
2) that the ability of the plants of regrow following
grazing declines as the season progresses; 3) that
moderate levels of grazing maintain forage quality (amount
of nitrogen as a percentage of the dry weight of tissue)
during the season compared to the corresponding
percentages for ungrazed plants; 4) that the increase in
NAPP at moderate grazing levels above that of ungrazed
swards is dependent upon goose faeces; without faeces
there is no increase; and 5) the input of nitrogen from
faeces is insufficient to account for the accumulation of
nitrogen (g * m"*) in swards which have regrown following
grazing. The presence of faeces probably influences
microbial growth and rates of net mineralization of
nitrogen in sediments (Chapter 5). This conclusion is
based on the magnitude of the difference in the amount of
nitrogen (g * m"3-) in above-ground biomass between grazed
swards in which faeces remained, and similar swards where
the faeces were removed. This difference is greater than
the amount of nitrogen supplied as faeces.
Early in the season, grazing by goslings leads to the
enhancement of above-ground growth of swards of
Puccinellia phryganodes in a pattern consistent with the
predictions of the herbivore optimization model. To our
knowledge this is the first experiment conducted in a
86
terrestrial system where the results provide a critical
test of the model. The mechanisms which appear to account
for the regrowth of Puccinellia are the addition to the
sediments of soluble nitrogen from faeces which is rapidly
taken up and used by the plants (Bazely and Jefferies
1985), and the net mineralization of nitrogen in faeces
and sediments which results in the further release of
inorganic nitrogen for plant growth (Chapter 5). The
reduced ability of the vegetation from heavily grazed
plots to show increased NAPP is likely to be a
consequence of the severe damage to meristems from
excessive grazing and trampling, and to inputs of soluble
nitrogen (largely as ammonium ions) from fresh faeces,
which may approach toxic levels (Bazely 1984).
Grazing late in the summer failed to produce results
which support the herbivore optimization model (Fig. 4-
2c,d). Rather, grazing resulted in amounts of NAPP similar
to amounts measured in the ungrazed swards. The results
appear to indicate that enhancement of NAPP was generally
limited to swards which were grazed early in the season
when there was still sufficent time for regrowth to take
place during the remainder of the season relative to the
growth phenology of the plants. The patterns of growth may
have precluded high rates of regrowth late in the season.
This decline in growth is coincident with the southward
migration of the geese, possibly reflecting the lower
nitrogen content of the above-ground tissue by this time.
In August, plants may have already begun to store
carbohydrates and nutrients in below-ground tissue, rather
than produce new above-ground tissue. Nevertheless,
87
grazing in the late season does stimulate some production
of new tissue. In Experiments 8 and 9, the NAPP of swards
in grazed plots was greater than that of swards of
ungrazed (control) plots after only 24 days of regrowth
(Table 3-6), even though the total above-ground biomass
was considerably less, compared to earlier in the season.
Our results, and those of Cargill and Jefferies
(1984b) demonstrate increased production of Puccinellia
swards. Similar results have been obtained in
experimental studies when leaves of individual shoots of
plants of this grass which have been grown in pots are
clipped and/or nutrients are added to the soil (Sadul
1987). The collective experimental evidence demonstrates
the importance of an adequate supply of nitrogen in order
to achieve enhanced production of swards of Puccinellia
following grazing. It is also clear, however, that the
efficient cycling of nitrogen necessary to sustain plant
growth in this system involves a number of processes well
removed from the immediate plant-herbivore interaction.
Previous studies have suggested that grazing by
Lesser Snow Geese results in the selection for genets of
Puccinellia which have higher growth rates than plants
which grow in ungrazed sites (Sadul 1987). While
fertilization from faeces may be required in order to
enhance production, clipping by itself also may stimulate
production in Puccinellia, by selecting for plants with
faster growth rates that would be inherently more tolerant
of grazing. There is some evidence for this, particularly
in Experiments conducted late in the growing season
(Experiment 8 and 9; Fig. 3-6), where grazing both in the
88
presence and in the absence of faeces increased NAPP above
that of ungrazed swards.
Not all plant species have the ability to respond to
herbivory as rapidly or effectively as Puccinellia
phryganodes. This response is dependent upon the ability
of Puccinellia to rapidly take up nutrients from the
sediments. The stolons are on the surface of the sediments
and the majority of the root system is confined to the top
five centimeters of sediment, hence nutrients which are
leached from faeces are likely to be rapidly absorbed by
the roots or stolons. Nitrogen applied on the surface of
the sediments is rarely available for uptake by deeper
roots, but is rapidly taken up at the surface (Marion,
Miller and Black 1987). Plants of Carex subspathacea
begin to senesce earlier in the season compared to
Puccinellia (Kotanen 1987), which precludes regrowth
following grazing except early in the summer (see Cargill
and Jefferies 1984b). Swards of two coarser, turf-forming
grasses common in willow sites in the upper marsh,
Calamagrostis deschampsiodes and Festuca rubra, do not
show increases in growth, within the season, following
grazing (Frey 1988), possibly because they are not able to
utilize nutrients available in faeces as rapidly as
Puccinellia.
In swards of Puccinellia 90% of the total plant
biomass is below-ground (Cargill and Jefferies 1984b).
However, Cargill and Jefferies (1984b) detected no
differences in below-ground biomass between grazed and
ungrazed swards. This does not imply that the rates of
turnover were the same. The differences in above-ground
89
biomass and NAPP shown in Fig. 3-5 and Table 3-6 also
indicate that the redistribution of biomass from below- to
above-ground alone cannot account for the enhanced growth
of grazed shoots. The presence of faeces was essential in
order to achieve the response.
The results of Experiment 6 and 7 provide critical
evidence that the faeces are essential for the herbivore
optimization model to operate at La Perouse Bay. There
are no differences in the amounts of nitrogen as a
percentage of the dry weight of shoots from the different
treatments, however there are significant differences in
NAPP, and thus in the total amount of nitrogen accumulated
in above-ground tissue (g * m"*). Thus the total amount
of high quality forage available to the geese is twice as
high in the presence of faeces compared to swards where
faeces were removed.
In a few other grazing systems similar increases in
NAPP have been reported. In the Serengeti, McNaughton and
his collegues have demonstrated that grazing by large
ungulates leads to increases in NAPP (McNaughton 1979,
1984, 1985a,b; McNaughton, Coughenour and Wallace 1982;
McNaughton, Wallace and Coughenour 1983; Ruess, McNaughton
and Coughenour 1983; Coughenour, McNaughton and Wallace
1985b). The recycling of nutrients from faeces and urine
in this system appears to be essential for regrowth to
occur (Ruess and McNaughton 1984, 1987, 1988). In coral
reef systems grazing by herbivores may also lead to
increases in production at moderate levels of herbivory
(Carpenter 1986). Coral reefs are highly productive
systems surrounded by seas low in nutrients, thus tight
90
nutrient cycling is imperative (Carpenter 1986; Sprent
1987) . Herbivores (sea urchins, limpets, fish) graze
intensively on "algal turf" communities which dominate the
surface of reefs. These algae are one of the primary
producers in the reef ecosystem (Odum and Odum 1955).
Herbivores excrete nitrogen into waters surrounding the
reefs, which provides a readily available source of
ammonium to support the growth of algae and cyanobacteria.
Grazing by sea urchins in particular increases the overall
rate of reef productivity by facilitating the flow of
nutrients required for growth (Carpenter 1986).
Goose faeces are essential for the regrowth of
Puccinellia following grazing. Goose faeces differ from
some other types of herbivore faeces in that they
generally have no adverse effects on the vegetation they
are deposited on. Cow pads, for example, provide a source
of nutrients for grazed plants, but represent a local
disaster, leading to death, for the plants they bury
(Harper 1977). Goose faeces may also have effects on the
nitrogen budget of the salt marsh other than the direct
contribution of nitrogen. These processes are critical to
the sustained regrowth of Puccinellia following grazing,
and will be discussed in the greater detail in the
following Chapters.
91
CHAPTER 4: NET ABOVE-GROUND PRIMARY PRODUCTION OF PUCCINELLIA
PHRYGAONODES IN RESPONSE TO MULTIPLE GRAZING EPISODES BY
LESSER SNOW GEESE: THE IMPORTANCE OF THE TIMING
OF GRAZING DURING THE SUMMER
4.1 INTRODUCTION
The response of plants to grazing depends upon the
phenology of plant growth in relation to the season, and
the frequency and the severity of defoliation of the
grazed sward. Experimental evidence suggests that
regrowth of graminoid plants following grazing, and
therefore the longevity and persistence of a particular
plant-herbivore interaction, depends upon the rapid return
of nutrients to the soil via faeces and urine (Petersen,
Lucas and Woodhouse 1956; McKendrick et al. 1980;
McNaughton 1983a,b; Ruess and McNaughton 1984, 1987;
Bazely and Jefferies 1985, 1988). The phenology of plant
growth may be altered by the timing of grazing during the
summer, and the ability of plants to respond to inputs of
nutrients following defoliation (Chapin 1980).
There is considerable interest in a model of grazing
responses which suggests that, at moderate levels of
herbivory, net above-ground primary production of forage
plants may be increased compared to that of ungrazed
plants (the herbivore optimization model; McNaughton 1979,
1983a,b; Hilbert et al. 1981; Dyer et al. 1982; Jefferies
1988a). Evidence in support of the type of response
predicted by the model comes mainly from studies of
grazing of terrestrial graminoid communities and aquatic
92
algal and coral communities (McNaughton 1976, 1979; Prins,
Ydenburg and Drent 1980; Cargill and Jefferies 1984b;
Ogden and Lobel 1978; Bergquist and Carpenter 1986;
Carpenter 1986; Power 1987; Chapter 3). This model
predicts the overall effects of grazing intensity on
production but does not address the outcome of the
interaction between the frequency and severity of
defoliation over the course of the season on plant
regrowth.
A number of models of the stability of plant-grazer
systems exist (Noy-Meir 1985; Parsons, Johnson and Harvey
1988), and these have been of value in managed
agricultural grazing systems where the objective is to
find the "optimum" balance between stocking density, the
duration of the period of grazing (continuous or
intermittent), and plant regrowth (Stoddart and Smith
1955; Parsons, Johnson and Harvey 1988). In natural
grazing systems the conditions necessary for increases in
plant growth to occur following grazing are not well
understood (Jefferies 1988a).
Seasonal effects in relation to the phenology of
plant growth and the timing of the foraging activities of
grazers are extremely important in determining the outcome
of plant-grazer interactions. In the Arctic, the
foraging patterns of a number of herbivores shows a strong
seasonal pattern. Many grazers are migratory (geese,
caribou) and the production of plant biomass is limited to
a brief period during the summer. At other times of the
year most plant biomass is frozen and buried beneath snow.
Nevertheless, consumption of total net above-ground
93
primary production (NAPP) in these environments can be in
excess of 80% of the available biomass (Cargill and
Jefferies 1984b; Bliss 1986).
Plant growth in the subarctic La Perouse Bay salt
marsh is strongly nitrogen-limited (Cargill and Jefferies
1984a). Previous studies have shown that grazing by
Lesser Snow Geese (Chen caerulescens caerulescens L.)
significantly increased the net above-ground primary
production of a stoloniferous salt-marsh grass,
Puccinellia phryganodes (Trin.) Scribn. and Merr., and a
rhizomatous sedge, Carex subspathacea Wormsk., from 30 to
105%, depending upon the year, compared to corresponding
values for ungrazed swards of these two species (Cargill
and Jefferies 1984b; Bazely 1984). A consequence of the
intense grazing is that plant tissue is converted into
either goose biomass or faeces, instead of accumulating
as live plant biomass or litter. Nitrogen released from
goose faeces accelerates the supply of this element to
the sediments. In addition, patches of sediment between
grazed shoots (0.5 cm diameter) are colonized by
cyanobacteria that contribute to the long-term input of
nitrogen, as a result of nitrogen-fixation (Bazely and
Jefferies 1988).
Detailed demographic studies have shown that plants
of Puccinellia phryganodes are capable of increasing the
production of shoots and leaves in response to grazing
(Bazely and Jefferies 198N). There is selection for
biotypes of Puccinellia with faster growth rates under
grazed conditions (Sadul 1987). The ability of both
Puccinellia and Carex to show rapid growth responses
94
within the season compensates for the adverse effects of
defoliation per se on above-ground net primary production
(Sadul 1987; Jefferies 1988a). A continual supply of high
quality forage is produced during a period when it is
most required by the geese.
As discussed previously in earlier experiments
(Chapter 3), the foraging activities of the goslings of
Lesser Snow Geese resulted in increased NAPP of
Puccinellia phryqanodes early in the season, but as the
season progressed the ability of grazed swards to recover
declined. Nitrogen from faeces was responsible, in part,
for these increases in NAPP and for maintaining the high
nitrogen content of grazed swards relative to ungrazed
swards (Chapter 3). In this study captive goslings have
been used to examine the effects of multiple grazing bouts
on swards of Puccinellia phryqanodes throughout the
growing season. Swards were grazed on 12 or 24 day cycles
at the same intensity of grazing (proportion of above-
ground biomass removed), in order to examine the effects
of the (seasonal) timing of grazing and of the interval
between successive grazing bouts on NAPP, the phenology of
plant growth and the quality of forage.
4.2 METHODS
4.2.1 Study site
Over 8000 pairs of Lesser Snow Geese breed each
summer at La Perouse Bay, 25 km east of Churchill,
95
Manitoba, Canada, (58°, 24X N, 94°, 24' W), on the west
coast of Hudson Bay. Following the hatch of goslings in
late June or early July, the geese forage intensively on
the vegetation of the intertidal salt marsh for 6 to 8
weeks before migrating south. The vegetation is dominated
by Puccinellia phryqanodes and Carex subspathacea, which
is abundant in less saline areas. The intertidal grazing
lawn is approximately 2 cm high and very few shoots (< 5%)
escape the effects of grazing (Kotanen and Jefferies 1987;
Bazely and Jefferies 198N). There are over 4000 shoots of
these species per square meter. After the geese leave the
marsh, plant growth continues in September before the
arrival of permanent snow. A complete description of the
study area is provided in Jefferies, Jensen and Abraham
(1979) and Jefferies (1988a).
4.2.2 Experimental trials
All experiments were conducted using captive
goslings. The goslings were removed from nests of wild
geese within 24 hours of hatching and were raised in
captivity in the research camp. Goslings were fed a diet
of natural foods (mainly Puccinellia phryqanodes and Carex
aquatilis), supplemented with a daily ration of Purina
8815 Duck Growena, or dry dog food, except on the days
prior to an experiment. Prior to each experiment the
weight of the goslings was determined (Table 4-1).
Experimental plots were established at the beginning of
the summer on selected areas of saltmarsh vegetation
96
dominated by Puccinellia phryganodes. The plots were
exclosed with chicken wire to prevent any grazing by wild
geese. Each plot was 1.1 m x 1.1 m. Four replicates for
each of the ten grazing treatments were established in a
completely randomized block design. On the morning of the
grazing bouts (6, 18 July; 3, 12, 23 August; 3 September
1987), goslings were taken to the grazing site and allowed
to "acclimate" to the vegetation in holding pens for 1 h.
They were then randomly assigned to a plot and allowed to
graze for a specific period. This process was repeated
until all plots had been grazed. Fresh water was provided
in small plastic bowls during the grazing bouts.
Grazing bouts were conducted at approximately 12 day
intervals (Table 4-1). The number of goslings per plot
and the period of grazing was adjusted over the season, as
the goslings increased in weight and their rate of intake
of vegetation increased, in order to achieve the removal
of approximately 25% of above-ground biomass (% offtake)
during each grazing bout. Grazing times varied over the
course of the summer from 45 to 15 minutes and 4 to 2
goslings were placed in a plot (Table 4-1). Faeces
remained in the plots and were counted at the end of each
experiment. Two days before an experiment was conducted,
goslings were allowed to graze plots of similar
vegetation, in order to determine the conditions necessary
(ie. number of birds, length of grazing peroid) to achieve
a 25% offtake.
Swards of Puccinellia were grazed at intervals of
approximately 12 or 24 days (Table 4-1). For the 12 day
cycle, plots were grazed on 1, 2, 3, 4, 5, or 6 occasions.
97
Table 4-1. Days on which experimental plots containing
swards of Puccinellia phryqanodes were grazed and/or
sampled during 1987 at La Perouse Bay, Manitoba, on a 12
day cycle, 24 day cycle and late season only grazing
schedule. The mean weight of goslings (g), length of the
grazing bout, and number of goslings grazed per plot is
also shown. Grazed plots were sampled before and after
grazing bouts.
Treatment
(# of bouts)
(of grazing)
Date
12 day cycle
6
5
4
3
2
1
0
24 day cycle
3
2
late season
3
mean
weight (g)
grazing time
Dates grazed
6/7
g
g
g
g
g
g
s
g
g
s
499
45
18/7
g
g
g
g
g
s
s
s
s
s
and
3/8
g
g
g
g
s
s
s
g
g
s
880 1200
30 40
sampled
13/8
g
g
g
s
s
s
s
s
s
g
1478
25
(g) or
23/8
g
g
s
s
s
s
s
g
s
g
1841
20
sampled (s)
3/9 15/10
g
s
s
s
s
s
s
s
s
g
1876
15
s
s
s
s
s
s
s
s
s
s
(mins)
# goslings
per plot
98
For the 24 day cycle, plots were grazed on 1, 2, or 3
occasions (Table 4-1) . The same plots for the "ungrazed
control" and for those plots where one bout of grazing
occurred were used in the comparisons for both the 12 and
24 day cycles. One further treatment was established
where grazing by goslings was not initiated until 13
August (late season schedule). Grazing was continued in
these plots until the end of the season (3 September) at
12 day intervals. Swards in plots which were grazed on 23
August and 3 September extended the period of grazing up
to 3 weeks after wild geese had left the marsh,
exaggerating the overall effects of grazing on the
vegetation beyond levels observed under natural
conditions.
4.3.3 Sampling and data analysis
Above-ground biomass was sampled by removing a 7.5 x
7.5 cm turve at random from each of the plots before
grazing, and after grazing from the grazed plots. The
above-ground biomass was removed by clipping the sward at
ground level. The plant material was washed and then dried
at 60-80°C for 24 hr. Since 90% of all material was live,
standing dead biomass was not removed prior to weighing. A
number of fresh faeces were collected from the gosling
holding pen adjacent to the experimentsl plots for
determination of their dry weight and their carbon and
nitrogen contents. Dry weights were determined using a
Mettler (PK-300) electronic balance. Determinations of
amounts of carbon and nitrogen present in dried faeces and
99
plant tissues were made using a LEGO series 600 CHN
autoanalyser (St. Joseph, MI . , USA). All material was
ground (20 mesh size) in a Wiley mill and appropriate
quantities (approx. 100 mg) were preweighed before
combustion in the analyser. All above-ground tissue was
pooled for the analyses; leaves and shoots were not
separated. Analysis of Variance (ANOVA) was used to
assess the effects of the treatment.
4 .3 RESULTS
4.3.1 Changes in NAPP of swards of Puccinellia
phryganodes in response to grazing
Net above-ground primary production of plots of
grazed swards of Puccinellia phryganodes was influenced by
the number and timing of the grazing bouts during the
season (Fig. 4-1). Where swards of Puccinellia were
grazed approximately once every 12 days (Fig. 4-la), those
grazed on 3 or 4 occasions had a higher total NAPP than
the ungrazed (control) swards or swards grazed on 1 or 2,
or 5 or 6 occasions (2-way ANOVA: F = 2.388, df = 6,105,
p < 0.033). The cumulative NAPP of all swards increased
significantly over the course of the summer (F = 132.730,
df = 4,105, p < 0.001), but the interaction between season
and treatment was not significant (F = 0.344, df = 24,105,
p > 0.998). No significant differences in cumulative
NAPP among the grazing treatments were apparent until 23
100
August. By the end of the summer (3 September) ungrazed
swards had a lower value for cumulative NAPP than all
grazed swards (Fig. 4-la).
For plots grazed on a 24-day cycle (Fig. 4-lb), the
cumulative NAPP of the sward grazed on three occasions was
significantly greater than that of swards grazed on 1 or 2
occasions, and the ungrazed sward (F = 4.578, df = 3,60, p
< 0.006). Increases in cumulative NAPP between treatments
over the course of the season were significant (F =
106.830, df = 4,60, p < 0.001), but the interaction between
season and treatment was not (F = 0.638, df = 12,60, p <
0.802). In the sward which was left ungrazed until 13
August (late season schedule), the cumulative NAPP was
similar to the plot grazed on 3 occasions at 24-day
intervals earlier in the season.
The initial amount of above-ground biomass present in
all plots at the beginning of the experiment (6 July) was
54.6 + 3.5 g DWT * nr*. By the end of the summer (3
September) the mean above-ground standing crop in the
seven treatments for the swards on the 12-day grazing
cycle varied significantly (116.5, 119.3, 111.2, 103.1,
82.7, 58.8, 61.6 g DWT * m'a for swards grazed on 0
(control), 1, 2, 3, 4, 5 and 6 occasions respectively).
For the 24-day grazing cycle and the plots grazed late in
the season the mean above-ground biomass on 3 September
was 116.5, 119.3, 106.4, 88.6, and 78.9 g DWT * nT*
respectively for swards grazed on 0 (control),1,2 and 3
occasions, and those grazed late in the season. The
above-ground biomass of swards in plots sampled on 15
October was lower than that sampled in early September for
t_01
Fig. 4-L. Curnulative net above-ground primary production
of swards of Puccinellia phrvqanodes grazed by goslings of
the Lesser snow Goose for each treatment during the summer
of Lg87 (rnean value of 4 plots i each plot was L. 1 x 1' l-
m ). a) NAPP of swards grazed at approximately L2 day
intervals. Ungrazed control (+); one grazing bout (O); two
grazing bouts (a); three qrazing bouts (o); four grazing
bouts (r); five grazing bouts (l); six grazing bouts (l).
b) NAPP of swards grazed at approxirnately 24 day intervals
and only late in the season. ungrazed control (+); one
grazing bout (o); two grazing bouts (v); three grazing
bouts (v); three grazing bouts late in the season (*).
II
rEbc)
26Er'
+l
E
Fits1
Al
as
3Rt
CUMULATIVE NET ABOVE.GROUND PRIMARY
PRODUCTION ( G-DWT, M'' )
CUMULATIVE NET ABOVE. GROUND PRIMARY
PRODUCTION ( G.OWT. M.?)
'atnTl sTr{? 1e ue6aq .f,1uo ddVN Jo sluauarnseau acuTs
'paunsuoc ddvN Tefol aql Jo a6eluac.rad aqq Jo aleuT?sa
aq1 uT papnTcur 1ou sT (dtnf g) 6u1ze:6 Jo lnoq lsrTJ eq1
6u1:np s6u11so6 aq1 
^q 
paumsuoc sseuroTg 
- 
eZv) suolsecco
E uo pazerb spJpAs aq1 uT 1saq61q seat alcr{c 6uyze:6 dep
?Z aq1 uT s6ug1so6 aq1 lIq paunsuoc ddVN aq1 Jo abeluac:ad
aqfi '(Z-V aTqe;) a1c.dc ^dep ?,I aql roJ sauTl 9 ro E
paze:6 sp.rEns aq1 uT AEL ralo <>1 ,rauruns aq1 6u1:np acTAl
paze:6 sp.rpns ar{l uT ZL.7T urorJ ,s1noq 6u1ze:6 Jo Jagrrnu
aql I{1Tr{ pasearcur sluaurTJadxa aq1 6u1.rnp s6u11so6 ar{1 ^f,q
paunsuoc ddVN Te o? ar{1 Jo abeluec:ad aq1 ,Te:aua6 uf
'ZgZ rllalerurxo:dde TTTIs spn palouar sseuoTq Jo e6eluac:ad
aldl '.raaaato11 
.:aq61q sen sseuoTq Jo lunoue TeTXTuT ar{l
a.raqn uosees aql uT aleT r(1uo paze.r6 atraA qcTr{A s1o1d aq1
ur ldacxa '(sseuotq puno:6-aloqp Jo Z9Z lnoqe) 1uau1:adxa
qcea .roJ lnoq 6u1ze:6 qopa 6u1:np T_ut * ,ll{O 6 gT 01 ZT
pa6e:ane paunsuoc sseuroTq Jo lunoue aq; .6u1ze:6 a.roJaq
luasa:d sseuoTq puno.r6-aloqe Jo lunoue aql uojrJ butzezb
6u1lro11o3 fi1a1e1paruur1 ss€urorq puno:6-aloqp Jo lunoure ar{1
6ut1ce:1qns dq paleTncTpc sEA sTr{tr 
- 
z-n aTqe& uT urtoqs sT
6u1ze:r6 Jo lnoq qcee 6u1:np pup Jaururns aq? .ralo s6ur1so6
z(q peunsuoc sseuroTq puno:6-aloqe Jo lunoure aq;
'appu 1ou a:an sp.rooa.r a^TlelTluenb z(1a1eun1.roJun
lnq '.rauuns aq? uT alpT paze:6 a:on spJei*rs a.rar{r'r sqold
aI{1 uT spJ{ raqolcg uT luapTle salpaT J{au Jo raqumu 1sa6:e1
aq6 
. 
(f-t 
.6T,f) :auruns aq1 uT .raTT.ree sauTX Z ro T
.f,1uo paze:6 sp.rpr'rs uT pup sp-reAs pazeJ6un aq1 u1 1sa1ea:6
pup 'Jauruns aql uT a?pT peze:6 sp:ens aq1 uT lseaT spn
:aqualdag e uo luasa:d sseuoTq aql r{1Tr'r pa:eduoc ssEluoTq
puno:6-aloqp uT uoTlcnpal aql ,:aaaaog -s1uaurlpa.rl TTp
zor
v' L9
e'eT
L'TV
0
0
L'VI
z'62
9'?V
I'9 L
6'8L
Z'IT L'LZ L'NZ0
00
0 6'9?,
0
o
0
0
0
oe
uoseas axPT
0'9T Z
8'ZT E
aTs.dc .f,ep tz
00
'.TT 
T
T'OT Z
Z'LT g
I'Ll V
E'8T 9
6'E 9
aTc^c J{ep zT
00
00
oo
00
o0
0 8'zI
9'T 9'6
0
0
0
0
9'8
0'87
g'87
9'07
g'rT
0
0
0
9'Zr
6'gT
?'97
s'tT
0
0
1'Zr
8'Zr
L'ET
?. TT
t'e T
6/Es/E?,s/zrs/eL/87L/e
paunsuoc
ddvN
Te101 Z
( t-u + .lMo 6) paunsuoc sseuoTq puno.r6-aAoqe 
Jo lunouv
(6u1zer6 ;o) (s1noq 3io #)
lUaUlearJ,
'(r(tnf g uo paunsuoc sseuoTq opnTcuT lou op sanTpl
asaql) uAor{s osTe sT raururns aq1 6u1:np s6u11so6 aq1 trq
paunsuoc sen qcrr{A '(1"-t 
.6Ta) pa:nseaur davN Tplol aq1 Jo
uotl:odo;rd aq,1, 'eg6T uT s1o1d TeluaurT.radxa u1 asoog rrrous
rassaT aq1 Jo s6u11so6 ,{q paurnsuoc G$5EE6E@ ffi-tlfffi
Jo spreAs Jo sspuroTq puno:6-aloqp Jo lunourg 'Z-t aTqe,tr
eoT
pazer6un uroJJ slooqs .roJ sanTen 6ulpuodsa.rJoc ueql :a1ea:6
slooqs aql uT ua6o:1;u Jo slunoue peq uoseas aq1 uT aleT
paze:6 sloTd ar{1 pue suoTsecco g pup Z uo paze:6 sloId aql
'(ez-v '6T.r) alcdc ,{ep zT ar{1 uo paze:6 s1o1d uorJ sxooqs
uT leq1 uEr{1 ssaT r{11e:aua6 a:an 1q61aar .d:p aq1 Eo abeluac:ad
p sp sloor{s uT ua6o.r11u gro slunoue aq1 'uoseas aq1 uT alpT
paze:6 asoql pue alcr(c dep vz e uo pazeJ6 s1o1d aq1 uI
'(ez-v '6Tg) :aununs aq1 Jo :apuTeurar aql JoJ pauTeluTeul
sen acuaf,aJJTp sTr{,1 'sprens paze:6un aql uoJJ slooqs
uT ua6o:11u Jo slunoure aq1 upql :aq61q r(11uec1gr1u61s
aran rauurns aq1 6u1:np saurT1 9 pue 'g 'V pazez6 s1o1d
ruo.rJ ETre-ffiE Jo slooqs aq1 uT ua6o:11u Jo lunoup aq?
(+sn6nv e ) burzezb Jo lnoq prTql aq? -ralJg '.raqo1co Jo
aTppTur aql pue .raqueldas Jo 6u1uu16aq aq1 uaanlaq I11q611s
pasea.rcuT sp.reAs paze:6un pup pazp:6 qloq Jo sloot{s
aq1 uT ua6o:11u Jo lunoup aq& 'Gtt'O < 'd '6V'ge = Jp
'V9L'O = ,!I :uorlcereluT 1uaulpaJ1 x auTl ITOO'O > d '6i'9
= JP 'C?T'9 = S :?uaur1€ar1 jTOO'O > d '6V'g = Jp '9Z6'6
- .{ :auTl iVAONV Aem-Z) :aqualdag u1 uala Ze'Z eloqp
pauTeual lnq 'uoseas aql Jo asJnoc aq1 :elo r(11uec1E1u61s
asearcap 1ou pTp 1q61ai-r .d:p aq1 gro a6eluac:ad
e se ua6o:1Tu Jo lunoue aq1 alc.dc dep ZT ar{1 uo s1o1d
paze:6 aq1 uI 
. 
@z-t 
.6T,{) :aqualdag ;o 6u1uu16aq aql
.dq s1o1d (1o:1uoc) paze:6un aq1 u1 ZEL'T d1a1eu1xo:ddB
o? pasparcap lnq 'sluaurlee-r1 TTp -roJ ZL'Z lnoqp
(f,tnf g) I11e141u1 sei"r 1q61aaa ^6:p ar{1 Jo uo11:odo:d B se
'sseuorq puno:6-aloqe aq? uT ua6o:11u Jo lunoue aq,tr
sapouEbfiqd
puno:6-aloqe uT uoqJec
eTTTauTscnd Jo
pue ua6oflTu
?OT
sseuoTq
JO s?UnOUIV 7,'e'?
t_05
Fig. 4-2. Total nitrogen (a) and carbon (b) content as a
percentage of the dry weight of above-ground biomass of
Puccinellia phrvctanodes during the summer of 1987 (mean
values are showni n = 2). Above-ground biomass with a
nitrogen content significantly greater than that from the
ungrazed swards at each harvest are circled (p < 0.05t sNK
multiple range test). ungrazed control (+); Swards grazed
on L2 day cycle: one grazing bout (o); two grazing bouts
(A); three grazing bouts (o); four gtazing bouts (r); five
qrazlng bouts (r); six grazing bouts (o). swards grazed on
24 day cycle and late in the season only: two grazing
bouts (v); three qrazing bouts (v); three gtazing bouts
Iate in the season (*).
NITROGEN CONTENT (% DRY WEIGHT)
i.'PNs! oo'o'6,5,
I
m
a
-t
o
{m
(!@
!
CAREON CONTENT (% DRY WEIGHT)
:ia!a3iB38
c6)r
u>
:E
-
m
o
-l
o
m
(o
co
{
saleaT lnau Jo lualuoc uaboJlTu ar{1 ueql ssaT dlqeqo:d
sT sspurorq puno:6-aloqe pelood ar{1 Jo lualuoc ua6o:1;u
aq& 'sspuroTq puno:6-anoqe uT ua6o:q1u Jo uoTleTnuncop
+au aq1 Jo sanTea abe.rarre a.re saleurTlsa asar{1 ,saaeal aql
lsn[ 1ou pue sseuoTq puno:6-aloqp Tp1ol .roJ pauTurJalep sT
ua6o:11u Jo lunoue aq? acuTs 
.po1.rad sTrll 6u1:np 1q61an
d:p aq1 ;o a6eluac:ad p se slooqs aq1 uT ua6o:X1u gro
lunoue aq1 
^q 
luau1pa.r1 6u1ze.r6 qcea :og slsan.rer{ uaaAlaq
sspuroTq Jo sluauarcuT alrssaccns 6u1I1d111nu ^dq palelncTp3
arp sanTel asaqJ, 
.€-? .6T,f uT uAor{s sT sseuorq puno:6
-aloqe aq1 uT paleTnurncce ua6o-r11u Jo lunoup aq,l
sapoaeTFF
sseuoTq puno:6-aaoqp uT
eTTTauTccnd Jo
paleTnuncce ua6orlTN E-e.n
'(qz-v '6T,q) slooqs 3lo 1q61aar d:p aq1
Jo Z9'Lv .d1a1eru1xo:ddB 01 pasea:cuT sXuaurlea:1 6u1ze:6 TTe
uto.rJ spJerqs uT uoqrpc Jo lunoure aq1 .raqolco uI . (qz_V
'61,{) raqrualdas pue 1sn6ng ,r(1np u1 s1salrpr{ TTp :oJ puE
'1o:1uoc paze:6un aq1 6ugpnTcuT ,s1uaur1eer1 TTe ;oJ *tt
I1a1eulxo:dde pa6e:a.rre , sseruorq puno:6-aloqe ar{1 Jo 1q61arr
^f,rp ar{? ;o ebeluac:ad p se ,uoqtec Jo lunoue aq;,
'anssTl Jo
1q61an zkp aq1 go aFeluac:ad e sp ua6or11u Jo lunoue .ranol
p ppq .d11e:auab uospas aq1 uT a?eT paze:6 spJeJqs uorJ
sloor{S 
. 
(Eee.O
luau?ea.rl x auTl iOgO.O > d 'gZ'e = Jp '916.Z =
,jI :luaurlea-rl ITOO.O > d 'gZ'g: Jp ,099.6 = d :aurTl jVAOI{\a
.tten-Z) acuo r{1uo paze:6 aJaA r{cTr{/{ sp:EAs luo.rJ pup sp.rplts
907
Jo lunourp aq& 
.aTcIc 6u1ze:6 .ttep Zl aq+ .roJ suoTsecco
9 uo paze:6 s4o1d :oJ t-ur 1 H 6 EZS.O 01 ,acuo Iluo paze:6
s1o1d u1 !-ln + N 6 2ZT.O uro.rJ a6ue: sanTEA asardf,, 
.sacapJ
uT pallsodap ua6o:1Tu Jo lunoue aql pue slnoq 6u1ze:6
Io raqunu aq1 uaaAlaq dlqsuolqeTa.r .rpauTT aa11lsod e
sT a.rar{[, 'e-n aTqef,, uT u/'roqs sT ( r-nr + N 6) s1o1d paze:6
aql uo sacapJ se pa1Tsodap ua6o:11u Jo lunoue ar{,1
'spJer'rs
paze:6un aq1 upr{1 ua6o:11u Jo slunorue :a6:BT pa?eTnluncce
uospas aq? uT aleT pazetb spJpi*rs aql '1e:aua6 u1 
.1sn6ng
eZ trq luaprla aJaA sprEAS paze:6 pue (1o:1uoc) paze:6un
Jo sseuoTq ar{l uT ua6o:11u Jo slunourp uT saouaJaJJTp
?uecTJTuFTS '(qe-f '6T.{) suolsecco E pue Z uo paze:6
a.ran I{cTqA spJprts uT 1sa1ea:6 araA sseuroTq puno:6-aloqe
uT paleTnuncce ua6orlTu Jo slunoue ar{X ,a1c.f,c dep VZ p
uo paz€r6 s1o1d aq? :og 
.s6u11so6 aq1 ,{q suolsecco g pup
'V 'e uo paze:6 a.raA r{cTqn s1o1d uorJ sp.re/{s aq1 uT aran
:aqualdag g ,{q paleTnurncce uaboJlTu Jo sXunorue 1saq61q
eq& '(E-r '6T.{) sp.rerrs paze.r6 aql roJ ueql ssaT sen
sprpr'rs paze:6un u1 paltsodap ua6o:1Tu aql auTl sTql ralJg
'1sn6ng 91 o1 dn s?uaurleazq buyzetb TTe JoJ JeTTurTs ser'l
sseuoTq puno:6-aloqe aq1 uT palefnurncce ua6orlTu Jo lunoue
1au aq1 
, (et-? -6T,E) a1c.dc 6u1ze.r6 .dep zI aq1 -rod
'poqlau sTrll 6u1sn
uosEas aql uT a?eT slooqs aq1 ,{q dn ua>1e1 ua6o:11u 3o
lunourp lau aql ale1ncTec 01 alqlssod lou sT 1T a:oJa-raql
'sseuo;q ut saspaJcur ou aran a.raql '(ez-t 
.61,{) :aquraldag
uT leq1 ueqX :eq61q .d11eu16:eu ser'r sseuoTq puno:6
-aloqe Jo Xualuoc ua6o:?Tu aq1 pue , (uo11en:asqo leuos:ad)
raqolog ur luapT^a ser'lr qlno:6 arau q6noq11g 
.auoTp
LOT.
l_08
rig. 4-3. Net amounts of nitrogen (g N * m-1) accumurated
in the regrowth of above-ground bionass of swards of
Puccinellia phrvqanodes during the surnmer of Lgg7. a)
NAPP of swards grazed at intervals of 12 days. Ungrazed
control (+); one grazing bout (O); two grazing bouts (a);
three grazing bouts (o); four grazing bouts (r); five
grazing bouts (r); six grazing bouts (r). b) NApp of
swards grazed at 24 day intervals and only late in the
season. Ungrazed control (+); one grazing bout (o); two
grazing bouts (v); three grazing bouts (v); three grazing
bouts late in the season (*).
I' e{
NET AMOt'NTS OF NITROGEN IN ABOVE.GROUND
R€GROWTH OF SWARDS OF PIJCCINELA'q (G.N.M.')
NET AMOUNTS OF NITROGEN IN ABOVE.GROUND
REGROWTH OF SWARDS OF PUCCINELLIA (G. N.M.2)
I
Im
(oo6- c(, l,l
v
m
a{
o
m
(o
o
I
C.
c
e
co
cg,
8ee'
EgZ'
LLE'
0
LZT'
z9z'
89V'
EO9'
VIL'
t.zg'
o
0
0
0
o
0
TTT'
o
860'
0
0
0
o
0
9TT'
LNI'
0
0
LEI'
6eT'
9eT'
T9T'
8tT'
vzT'
9TT'
0
LZt'
9ZT'
0?T'
6?T'
8TT'
6TT'
?80'ZIT'znr' 00e
uoseas aleT
z
e
aTc{c .f,ep tZ
o
T
z
e
v
I
9
aTcd3 .dPp ZT
O TET'
0 g9T'
00
o0
00
o 68T'
TtT' 08T'
9?T' ?LI'
LVT' e8T'
0
0
6/ee /ez,s/zr8/EL/8r L/e
f e10,l(r-nr + N pallsodap ua6o:1ru6) Jo lunouv(6u1ze:6 Eo) (s1noq gio #)
lUAlulea.IJ,
'L86r ur s1o1d rpluaurradxa ur Fan-TE@ ffiffiE go
sprplcs paze:6 uo palTsodap asaag l'Ious JassaT ;o s6u11so6
Jo sacaeJ uT ( r_u + N 6) ua6o.r1tu Jo lunourv . E_t afqp,tr
607
.s/tpp ZI Jo Tp^.raluT 6utze:6 e uo paseq ,suorsecco
t pup E uo paze.r6 sp.rpAs ur 1se1ea:6 a.ra/{ ssplrrorq
puno:6-a^oqe aq? uT s1ue1d .dq palelnurncce ua6o:11u Eo
slunoue ar{tr 
. 
spJel[s paze:6 3:o sseuoTq puno:6_a^oqe aql uT
ua6o.r1tu go sTaAaT ar{1 pue ddVN 6u1ule1sns .roJ TeT1uassa
araA d11ua:edde saoaeJ uT ua6o:11u go s1ndu1 . (t_f .61.{)
uoseas aql uT aleT paze:6 spJelus roJ lseaT se& Jaqolco pue
:aquraldas uaaAlaq sseuoTq puno:6_aloqe aql uT asea:cap aqa
's1noq 6u1ze:6 uaanleq Tp^ra1uT dep l? ao ZT aq? rar{1Ta
01 1cafqns s1o1d :ogr ,JaAaAog .s.dep V?, Jo Te^.ra1uT
ue 1e suoTsecoo or'rX .ro auo uo paze.r6 s1o1d ueql ddVN fp?o1
:aq61q e peq ,uospas aql Jo pua ar{1 1e Iluo suoTspcco
e uo pazezb asoql pup ,sz(ep tr7 f,,za/.e acuo suoTsecco g
uo pazerb s1o1d ,uos1:eduoc .f,g ..raununs aq? lnoq6no:q1 s.dep
zr d:aaa .f,1a1eur1xo:dde pazer6 sp.reAs Jo pue ,uoseas aql Jo
6u1uut6aq aq? 1e dluo paze:6 sp.re/vrs Jo ,spreats paze:6un go
uoT+cnpo.rd puno:6-aaoqe o1 pa:eduroo ddVN .raq6gq e pa^or{s
1sn6ng ;ro 6u1uu16aq aq1 TTlun paze:6 sprpl,rs .qllo:6 1ue1d
go .f,6o1ouaqd aql 01 a^rlpTa.r ,sqnoq 6u1ze:6 a^Tssaccns
uaarrlaq TeA-ra?uT ar.If pue uoseas aq1 6ur:np 6urze:6
Jo aurl ar{1 uo luapuadap sT @ofu6r:Td 
-E-mT-t-ne JO Sp.rp''\s Jo qlrrno:6a.r aql 1eql Aoqs slTnsa.r aqtr
NOISSNCSICI ?'V
. 
(e-A aTqe&) r_rr + N
?'0 ueql SSAT Sp/rl, uospes aql uT aleT paze:6 :o alcdc .dep
?z aq1 uo pazer6 sprens uo palTsodap sacapJ ur ue60:11u
OTT
dn p11nq o1 suals pup salpaT paTqeua eleq r[eu ,:auurns aq1
Jo qcnu .roJ uoTleTToJap Jo ecuasqp aq1 uT 1cpluT pauTpua.r
r{oTqr*r ,eare aceJ.rns JeaT c11aq1uz(soloqd aq1 ,uol1gppe
uruoseas aq? uT 31er rTlun qlltro:6 uTelsns o? luarcrJJns
uaaq a^er{ rleu ua60r1Tu ctue6:ou1 Jo asear€J pue ua60:1tu
Jo uoTlpzTTe-rauTu 1au 6ur11nsal aqfi 
. 
(feet ,{Tazeg)
I{6Tr{ ate sarn4e:adua1 luaurpas uar{A. aruTl e le sluauTpas
uT ua6o.r11u Jo uoTlezTTerauTu 1au aq1 snql pue ,qlrrno:6
TeTqo.rcTu alenTlce .ro au1:d .deu au11 sTql 1e sacaeJ
uT paplao:d ua6o:1ru 1nq ,rea1c .d1a:11ua Xou a:e sTql roJ
suosPa.r ar{& 
.tauuns ar{1 Jo JTeq ral1ET ar{1 uT suoTsecco
e uo paze.r6 a.raA qcTr{A s1o1d u1 lspaT aJaA .laqualdag
.f,1:ea u1 sanTel qlTrrr pa:eduoc raqolcg uT do:c 6u1pue1s
puno:6-aloqp Jo sasparcap q6noq11e , (dTng alET 01 pTur)
q1no.r6 1ue1d :1ead Jo por.rad ar{+ 01 palTuTT sen 6u1ze:6
uar{rrt 1saq61q seA .raqualdag z(1:ea o1 dn sp.Jre/qs paze:6
Jo ddVN 
.luaprla a.ran bul.zetb 6u1,rro11ogl uolqelabaA Jo
q1to.r6a: aq1 uo qllto:6 1ue1d ;o .66o1ouaqd aq? pup slcaJJa
Teuospas qloq Jo acue1:odurt aq;, 
. 
Telo? ar{l Jo ZgZ
lnoqE 1p luplsuoc .f,1aaT1pTar 1da{ spA lnoq 6u1ze:6 qcpa
6ut:np paloual sspluorq Jo Aunoue aq;, .sTerrJaluT lua.raJJTp
1e slnoq 6u1ze:6 Jo Jaqunu 6u1sea:cur ue o? pa?ca[qns
ffifr-ffi6 go spreAs Jo r{1^or6e: aq1 pauTuexa .fipn1s
sTql 'pa1e611saAuT a.ran uorlela6al ar{1 Jo r{1Ao.r6a: aq1
uo 6ulzetb Eo spol:ad 1ua-reJJrp Jo slcaJJa aql r{cTqa uT
(g :aldeqc aas) laporu uoTlezTuTldo aJolrqJaq aq1 Jo slsal
.raTTrea a>tTTun 
.spJens paze:6 Jo ddVN aql uT sasearouT
uT lTnsa.r 6urze.r6 ;o sTaAaT alerpa'uraXuT 1pr{l , (e
:aldeqc) Tapou uoTlezrurrldo-a:olrqraq ar{1 Jo suo11c1pa:d
aq1 1:oddns luaut:adxa sTrll Jo slTnsa.r aq,L
TTT
sp're,o' aql JO Sseurorq puno'6-a^oqp ar{f uT palernunccp
ua6oJlTu eql Jo TTv .ger pue gz uearrleq sT :oJ
palunocce aq FEIF ?er{1 (sp:ens peze:6un Jo N alTleTnunc
- N sacaeJ sp'e^s paze.r6 Jo N aallelnrunc) ua6o:1ru gro
s?unoue aATleTnlunc aq1 suoTsecco ? pue , e 
., z 
, 
T UO paze:6 aJaA 
1pq1 sprp^s u. sp'p^s peze:6 Jo ddvN aql UT lUasa:d
ua6o:11u Jo slunoue Jo sanTe^ asaql uroJJ palcprlqns
sT sprE,[S (1o:1uoc) paze:6un Jo ddVN ar{1 uT pale:odjrocur
sT 1eq1 ua6o:11u Jo lunoue eq1 .ra1JE sabeluac.rad asaql
alpTncTpcal 01 sT uor?eTncTec sTql uT dals puocas aqtr,
'.t{lanrlcadsa: ggz pue *zz aze suoTsecco n
pue g uo paze:6 s1o1d Jo ddvN ar{1 uT palernuncce 
ua6oJlTu
Jo slunour' lau ar{1 Jo a6eluac:ad e s€ sacaeJ uT 
luasa:d ua6o:1;u 4q,1, . (tst) ac1,rt1 :o (ZD 
acuo .d1uo paze:6 s1o1d
uT lspaT pup , (Zgt) suorspcco E pue , (gge) suolsecco g uo
paze:6 s1o1d Eo spre^S uT 1sa6:e1 sT uolq:odo.rd 
s1q1 a1c.dc
6u1ze:6 dep zr aq1 roJ '(s reldeq3 jN86T seTiraJJaf pu.
tTH 
,sseng aas) sassof uoTlezTTTfeToA ou a.re aJaql 
1eql pue pazTTeraurur sr 
sacaeJ eq1 uT ua6o:11u aql Jo tOgT ler{1
aunss. suoTlerncrec asaqtr' 'pa?etncr.c aq uec 
ddvN ar{f uT luasa:d ( 
l-ur * 6) ua60:11U Jo sfunoure lau a^TfeTnunc
aql Jo a6eluac:ad e se , (r-ru I b) s1o1d u1 palrsodap
socapJ uT luasa:d ua6o:11u Jo lunoue Tplo? aqtr
. 
suoTlTpuoc
TpJn+eu Japun spJpns paze-r6 ;o su:a11ed qluo:6 ar{l r{1TA
acuerJea Jp are uoseas aq? 01 uorleTeJ uT 6u1ur11 
s?T pup aaoqe 
pa1:oda: asuodse: q1,rto:6 Jo addl aql q?oq 
1eq1 os 'paze:6 ,{11aeaq 
uaaq alpq spJpi*ls aql relJe fsn6nv d1:ea ut
rlsrEIII aql eaeaT asaa6 aq1 .d11ensn .parJncco uoTlpTToJap
acuo qlnor6a: :oE ,t:essacau sele:pfqoqrec Jo se^Jasar
ZTT
Japun uorlcnpo:d .d:eu1:d 1au ,,6u1z;ur1do,, uT ?sa.raluT
sT araql ,:e1nc11:ed u1 ,sualsr{s TernlTnct:6e u1
paze:6 a.re s1up1d a:eqn s?uaurrrau*" "=";1":r1.:::::r-"::
'ugTleTToJap 6ularo11og e1115-ffia Jo spueuap qXrrao:6
aq1 laau o1 alenbape upq? aJour are ua6orlTu gro sallddns
aspc qcTqA uT ,palrsodap ale sacaeJ aJaqA s1o1d paze.r6 ffi
Jo sluauTpas uT rncco .dBur 6urul:d s1q1 ,aaoqe pauoTluaur
sp '1uaaa .f,ue u1 
.par{sTTqelsa aq 01 sureura.r lTcrJap aq1
jtoJ slunoccE rIcTr{J'l sluaurpas uT ua6o.r11u Jo uoTlpsTTe.rauTur
1au raql.rnJ o1 6urpeaT q?Ao-r6 1elqo:crur salplT?ce
Jo 'saul:d s1o1d paze:6 asaql uT sacapJ uro.rJ paseafaJ
ua6o:11u aql .raqlaq6 
.uospes aq1 uT alET paze:6 asoql pue
'suorsecco Z pue E uo pazeeb sprens :ogl .{1aa11cadsa: ZZg
pue tgg 'ZEt a:e slTcTJap aq; .sprpns paze:6un Jo sseuorq
aq1 uT ler{l aloqe pue raao s1o1d paze:6 u1 ua6o.r11u
Jo slunoure a^TleTnurnc aq1 JoJ lunoccp 01 luaTcTJJnsuT a:e
(6u1ze:6 uoseas a?eT pue alcXc dep nd sluauTradxa asaql
ur saoapJ Jo lualuoc ua6o: Tu aq1 Jo sanTpl 1eq1 salecTpuT
salpurrlsa Jo 1as puocas sTq& 
.^f,1al11cadsa: g8? pup
Zgv 'ZLg aurocaq sa6eluacrad asaql ,pa1ce:1qns sT sp.rens
paze:6un ;o sseurorq puno:6-a^oqe aql uT luasa:d ua6o:11u
Jo lunoue a^TlpTnurnc aql uar{& 
. 
ggT spA uoseas aql
ur aler paze:6 afai'I spJeI'Is aJar{/q a6eluec:ad 6ulpuodsa::oc
aq& 'sdep n?, Jo sTeAJaluT 1E suoTsecco z pup g uo paze:6
spJE&ls Jo sspurorq puno:6-a^oqe uT palernurncce ua60r1Tu
Telo1 aql Jo ,d1an11cadsa: ?ET pup g8T roJ pa1unoocp
sacaeJ uT luasa:d ua6o:1ru ,sluaut:adxa .raqlo aq1 uI
.suoTlPTncTPC asAql UO
paseq .roJ palunocce aq pTnoc suorspcco g pup E uo pazetb
CTT
.6u1ze:6 Jo acuanTJuT
aq1 xapun pal:asqo uoTlela6aa aq1 Jo ddVN uT aspaJcuT
arll roJ lunocce 01 .raAsue elqeldacce ?solu aqX sapTrro:d
ualsds aql uTrtllrrr 6u11c.dc 1ue1:1n1.1 
.pauuTr{1 e:e .daq1 uar{A
s1ue1d do:c go spuels auros uT s:ncco sE ,acuerpe:;r1 uT
sa6ueqc Jo lTnsa: aql aq louuec sp.reAs paze:6 pue paze:6un
uaanlaq ddVN uT acua.raJJTp ar{1 acuaH 
- 
(ggef ueq6no:g)
0'e Jo ssacxa uT sT IwI aq1 a.rar{A srncco Illensn sTq,l
'ddouec aq1 q6no:ql acuprpe.r.rT Jo uorlenual?e ur lTnsa.r
o1 .fi1aq1Tun arp sanTpl qcns 
. 
(feet rllazeg) E.f paacxa
1ou saop @f, go sprens paze:6un pue pazp:6 ;o
xapul parv JpaT ar{1 qsreu lTps r(eg asno.rgd eT aq1 1v
.paze:6
sT p.rens ar{l aJaqn (r_>tlr + 1_6 6) salp; qry,ro:6 alTleTar
I{6Tr{ anarr{ce 01 slueTd a6e:ogr ar{l saTqeua sTq& . rldouec
aql uTr{lTiq sTaAaT TTp 1p pauTelsns aq uEc pa.Ie JpaT
lTun :ad stsaqlur(soloqd Jo sa1p.r umurTxeu 1eq1 os ,ddouec
aq1 qbnoJql s.rncoo aoupTper-rT Jo uoTlenuallp aTl1TT
Xpr{4 sa.rnsua 6urze.z6 Jo acuanTJuT erdl Jepun l\fi AoT ar{1
pue do:c 6u1pue1s Jo lunoure rrror r1aa11e1ar aql ,Jraaanoll
'(geOt darr:eg pup uosur{of ,suos:e6) sp:ens paze:6un
uT sanTel 6urpuodsaf,.roc r{1Trn pa:edruoc ,rto1 sT (ffff)
xapul ea.rv JeaT aql a.raqn salTs le r'roT a.re auTl uaag6
p 1e do-rc 6utpuels eq? pu€ 'p:errs gro zur :ad slsaqlu.dsoloqd
Jo lunoue Tp?o1 aq1 ,as:noc ag .paleTqcp aJe eare
JeaT lTun :ad slsaqlur(so1or{d Jo sale.r q6Tq suoTlTpuoo qcns
.rapun '(r(douec aql r{leauaq puno:6 ;o ea:e lTun :ed seaeal
Jo eare) xapul ea.rg JeaT i"roT .d1an11e1a: e 6u1u1e1u1eru ^f,q
p.reAs paze:6 e uT palarqcp ualJo sT .ralau :ad uo;lcnpord
d:eu1:d puno:6-aloqp 1au unurTxel,{ .6utze.r6 snonuTluoo
?TT
ddvN Jo sanTp^ lsaq6Tr{ aq1 01 paT r{cTr{A r{1Ao.r6al Jo porrad
aq1 01 I1a1eu1xo:dde spuodsa:Joc lnq , (Hger sar.raJJaf
pue .dlazeg) z{eg asno:gd er 1p sabpas pue sasse:6 r{s;plu
lTps .roJ pa^Jasqo suorlerToJap paleada: uaa,qlaq pot:ad .dep
ZT ar{1 upq? :e6uo1 .d11q611s sr po1:ad sTq,t . (eeOf .TE la
luerg igg6T .daa:eg pup uosur{of ,suos:e6 Jgg6T 6uruua6 pue
suos:e6) s.dep ge pue VT uaar'rlaq sr slnoq bulzezb uaa$1aq
uo11e1a6an Jo qlato:6a: ;o TpA^re1uT ar{l uar{A ,,paz1u11do,,
eq 01 pual (ffiartd ffiTToT) sse:6ad: Jo ,{1T1;q11sa61p
pue alpr qlro:6 ,sasec q?oq uT ,uol1cnpo:d ur saspaJour
1sa1ea.r6 aql 01 ppaT suralled 6ulze:6 1ua11Tur.ra1uT ro
snonuTluoc Jar{laqA 01 sp alpqap auos sT araql aTTr{f{
'uoTlcnpo:d p:eats luanbasqns uo uoT?eTToJap Jo slcaJJa ar{l
6u1:ou61 .dq pa::ncu1 a-re sto.rJa aTqp.raprsuoc ,a1e:lsuouap
(gaer) suos:ed pue uosur{of sV .6utze:6 .dq palcaJJpun
sT uoTlcnpo:d 1eq1 saunsse osTp (Srct) :1ap_.doN sTapou sTr{
uJ 'sa.ro^Tq.rar{ Jo uorle:6rururr .ro uoT?epe.r6ap le?Tqer{ sE
qcns 'acupq-rnlsTp r{q lasdn .dlrsea aq z(eur sualsds 6utze:6
Te.rnleu uT z(111qe1s , (ggOt .Tp 1a luerg j8B6T rtaa:eg
pup uosur{of ,suos:e6 jgg6T 6uruua4 pup suosred) suals^ds
6urze:6 TprnfTncl:6e pa6eueu u1 pTTe,, ate suoTldtunsse
asaql aTTq$ 
.parTo.r?uoc sT sTeurTue ar{1 Jo luAua^our
aq1 Xeql pue ,aloT st uals.ds e urr{1gaa .d11aua6o.ra1aq
TeluauuorT^ua 1pq1 ,11eus sT (a:oarq:aq pup 1ue1d) salcads
Jo 
.raqunu aq1 1eql paurnsse pup ,uorldurnsuoc a:olTq.rar{
puP qlico-r6 1ue1d uaa/q1aq suoTlceJaluT aldurrs pa-raprsuoc
sas.f,1eue srg'suorleral addl da-rd-:o1epa.rd pr.ralToA-e{1or
uo paspq sualsds 6u1ze-r6 Jo ,{lTTTqe?s aq1 Jo slsdleue
uE paluasa.rd (gtOt) :1an_.dop .aToqn e se suralsr(s ilulzezb
aqr.rcsap 01 p.reAJoJ 1nd uaaq alpq sTapou Jo .raqunu V
ETT
6utzer6 luelsuoc e 6urunsse ,a1cr(c 6u1ze:6 dep ZT ar{1 .roJ
ddVN Jo aler aq1 uo slnoq 6ulzeeb Jo Jaqunu eq? 6u1sea:cu1
Jo slcaJJa ar{l Ieldslp d11ec1qde:6 gr 'a'p7_t .6T,t
. 
(cV-V 
.6T,f ) 1sn6ng-pTur uT alTs aq?
aleaT asaa6 aq1 fTlun sauTTcap uaq? pue ,s6u11so6 Jo rlclpq
6u1no11oE z(1a1e1paruu1 1sa1ea:6 s1 r{eg asno:gd er ?e preAs
ffiF-h" aqX uo (re) butzetb ;o I11suaf,uT ar{,tr . (auTf
uaqo:q) sp.rpAs paze:6 Jo 1eq1 ueql uospas aql uT raTT.rpa
auTfcap o1 su16aq I11e.raua6 (auTt pTTos) sp:ears paze:6un
Jo ddvN Jo a?er aqtr 
. 
(qv-v 
.6T,r) uoseas ar{f uT Ja?Er
6utsea:cep pue ,:auuns d1:ea u1 6ur::ncco qlrrto:6 plde:
lsollr aql qlTrtr ,uoseas aq1 ralo saEueqc (p-dep + ddVN)
qlto.rb Jo a1p.r ar{f,, -sprens paze:6 .d1lneaq ro pazp:6un o1
pa-reduroc (rc) 6urze:6 Jo sra^ar ale.rapou 1e paspa.rcur sr
ddVN !eV-? 
.6T.f uT unor{s sr Tapou uorlezrurTldo aro^Tq.raq
ar{J, 
.zlcuanba:gr 6urze:6 .ro (fe '.bur.zez6 go pol:ad)
.f,11sua1u1 6u1ze:6 pue ,uoseas 6urrrao:6 ar{1 Jo a6e1s aq1
'ddvN gro qde:6 TeuorsuauTp aatql e qbnorql suoTlcas_ssoro
se Jo 1q6noq1 aq upc (t-p'q)V-V .6T,f uT unords sal.rnc
aq& '.dcuanba:3r pup (6urze:6 3io po1:ad) .d11sua1u1 6urze:6
JO UOTIeUTqluOC qCeA JOJ palsnIlSUOC Aq Uec uospas aq1
ssoJce (,-dep + t_lu + ,I,OO 6) ddVN Jo alpr aq1 .roJ sa^f,nc
asuodsa: crlsr.ralce.rpqc 'ua1sr{s 6u1ze:6 .t{eg asno:a6 pr
aq1 :oJ Tapou uoTlezrurlldo_a:o^Tq.req ar{l Jo 1xa?uoc aq1 uT
paTTapou aq uec ,q1no:6 queld uo uoseas aq1 .ra^o 6utze:6
3o 
.f,cuanba:g: pue .61gsua?uT butzezb Jo lcaJJa aq,t
.luaJaJJTp Aq r(eu sp:errs
aq1 Jo sasuodsa: aq1 ,t11sualur 6urzB:6 :aq61q e lV .lnoq
6u1ze:6 qcea roJ 6u1ze.r6 Jo poTrad paxlJ auo 01 dldde rluo
sltnsal asaql ',:aaa.uo11 'aJaq pa1:oda: sluaurredxa aq1 ur
9TT
1,1,7
Fiq. 4-4. Responses of NApp to increases in grazing
intensity and the changes in the interval between
successive grazing bouts. The herbivore optinization nodel(a); seasonal patterns of rates of NApp (g Dwr * m-a *
day-l ) (solid line: ungrazed sward; broken line: grazed
sward) (b); seasonar pattern of grazing intensity (cr) and
use of salt marsh forage by geese (c); seasonal patterns
of rates of NAPP in sitrards where 25? of above_ground
biomass is removed during each grazing bout (cr = o.2s)
and grazed once every L2 days on 2 occasions (d); on 4
occasions (e); and on 6 occasions (f); Seasonal pattens of
rates of NApp i.n swards where zsz of above-ground biornass
is removed during each grazing bout (cr = o.2s) and grazed
once every 24 days where applicable: on L occasion (il; on
2 occasions (h); and on 3 occasions (i); seasonal patterns
of rates of NApp of swards where grazing intensity and/or
the freguency of grazing bouts is increasing (eg. cr = 0.4(i); Gr = 0.6 (k); GI = 0.8 (I)). The verticat arrows
indicate the difference in NApp * day-r compared to
ungrazed swards.
NOSV3S
o o
+ +
NOSV3SNOSV35
NOSv3S
z
!
T
Ao
z
'o
T
C\
o
+ NOSV3S+NOSVSS
z
.I
!
01 slueTd :og s}taaA x1s ,tlalputxo:ddp Jo po1:ad e saptao:d
lsnbng-pTur uT asaa6 aql Jo a:n1;edap aqfi 
. 
sluaua.rtnba:
TeuoTlT.r1nu tTar{l JoJ r{srelu lTps aql Jo uol1e1a6aa
aq1 uodn r{1lrreaq z(1a: asaa6 aq1 ,pa6pe1gl aneq s6u11so6
aql a.roJaq pue po1:ad lTnou ssa11q6113l sTr{l 6u1:nq . (e
-? '61,r) abe.rog .d+11enb r{6Tq Jo slunoue a6:e1 a:1nba: daql
uaq/'t auTl p 1p asaa6 aql 
^q 
pazTTTln sT rloTr{i*r ,sseurolq
puno:6-aloqe aql uT ua6o:1tu gro slunoue aql uTeXuTeur
seop asaa6 aql ^dq 6urze:6 ,:aAano11 . (f-E .6lA) 1sn6ng
uT pazTTear are sacua.raJJTp 1uec13l1u61s ,rt1ng u1 spreAs
paze:6un Jo 1eq1 o1 pa:eduoc sprpAs peze.r6 Jo ddVN ar{1 uT
sesea.rcuT luecrJTubls u1 lfnsar lou saop 6urze:6 q6noqlTV
'1sn6nq d1:ea u1 uorle:61u p.reAq?nos .rTar{1, uo qs.rpur
lTes aq? alea1 asaa6 ar{1 TTlun ?,uapTla d1a:11ua aq 1ou Aeu
uo11e1a6aa ar{1 uo asaa6 aq1 
^q 
6urze:6 Jo slcaJJa aqJ,
'(,{1aal1cadsa:
T'{'lV-v 6l.f ) sprpas aseql Jo r{1nojr6 :ood 6ul1uasa:de:
'pessa;dap aq ,(talTt lsollr pTnon sal.rnc asuodsa: asaql
'sasea.rcur slnoq 6u1zer6 Eo dcuanba:; aql jro d11sua1ug
6u1ze.r6 sg 
.p:ens paze:6un aq? Jo 1eq1 o1 pa.:eduroc
pa^de1ep sr ddvN Jo e1e.r aql uT auTTcap aq1 uospas aql
o?uT :aleT sanuTluoc 6u1ze:6 se ,r(lpuocag .slnoq 6u1ze:6
Jo Jaqunu aql uodn spuadap ddVN Jo a?pr aql ,.d11s.r1g
'a1c.dc 6u1ze:6 .dep tZ pue ZI aq1 qloq :oJ 1ua:edde
arp puar1 or'r,1, 
.a{e?JJo Z9z lo ^dltsualur 6ulze:6 luelsuoc
e 1p 'suorsecco (f ) g pue (q) Z, '(b) T uo sTel-raluT
dBp VZ 1e 6urzezb 1o slcaJJa ar{l Aor{s T,r{,bV-V .6TA
'(qt-l '6Tg) p-reats paze:6un aq1 01 pa:eduoc ddvN Jo aler
ar{1 uT asearcur eq1 alecTpuT sAor:p TecT1rarr ar{& . (s1noq
9 :I !s?,noq ? :a !s?,noq Z :p) a{e?JJo *Ez 3:o tr11sualuT
8TT
aq? qlTr,r op 01 6u1q1ou seq .,(1a>1TT ra/vtsue aq,l .1ng11ua1d
sr pue ,{111enb q6Tr{ Jo sT TTT?s Ilddns pooJ ar{1 uar{ru aruTl
e 6u1:np ,1sn6ng go 6u1uuT6eq aq1 le qsreu lTes aql aleaT
asaab aq1 .f,qn Jo uoTlsanb aq1 asTpr slTnsal asarlfi
.po1:ed sTql 6u1:np ddVN ?caJJp saop lnq
'anss11 puno.r6-aloqp ar{1 Jo lualuoc ua6o.r11u aql lcaJJe
1ou saop rauurns aq? uT aleT 6utze:6 panuTluo3 . (eZ
-t '61,r) uoseas aq1 uT d1:ea butzetb dq sp:ens paze:6 go
1pq1 o1 pa.reduroc ddVN asparcur pue a6e:og rTaql gro .d111enb
aql uTeluTeur o? aTqe osTe a.re asaa6 aq,l .6u1ze:6 .dq
pa.do:1sap a-re sualsT.rau lecrde aq? aouo uoseas aql uTr{?Ti*r
qlaro:6a.: gio alqedec 1ou a:e daql asnpcaq , (ggef sarJaJJaf
pue .f,1azeg) asaa6 ar{1 jroJ a6e:og aTqelTns lou e:p r{cTqn
sarcads snouoparrl0clp .dq peceld"Tp are sarcads prourue.rg
'6u1ze:6 Jo acuasqe aql ur luapT^a arp uorllsodruoc salcads
uT sa6ueqc pTdpg 
. 
(geOf sar:aJJaf pue z(1azeg) asaa6 aq1
Jo rnorlpqaq 6ulpaa3r TeTuoToc aql uo luapuadap d16uo-r1s
sT sp;pns pTourureJ6 pazetb aq1 Jo acualsTs:ad aq;
.pa.rJnoco,,dn-azaaJg:,, a.roJaq sTI{l
op 01 6uo1 se alpr.l 1ou pTnon .raqualdag .d1:ea pup 1sn6ng
aleT ur paze:6 sprei"rs 
. 
( 086T u1deq3 pue .raAer{S jOg6T
uazaT& pue Jar{c.rv !BL6I :ads:ag pue qcrJTn !gL6I uazaT,l
pup uosuqof !gL6T s6ulTTTg pup .raler{S !EL6T .Te 1a ssTTg
j 896T dauool4 pue sburrrrg) sploulue:6 crlc.re ur sseuorq
puno:6-noraq uT salJasar ale.rpdqoq:ec Jo uAop 6u1z(e1
aq1 .roJ 1ue1:odurT sT po1:ad sTrl,l .uoseas 6utarollog aq1
qlto:6 1:oddns 01 pue .raluTn.ralo 01 saa.rasal a1e:p.dqoq.rec
pue 1uaT.r?nu luarcrJJns a:1nbce 01 s?ueTd aq1 ro3:
Terluassa aq rteu 1sn6nv aler ur s.rnc3o rlcrr{rc sprEl[s pezetb
Jo q1Ao.r6a: aqg, .6u1ze:6 Jo slcalJa ar{4 luorJ :alocatr
6TT
' (q'egg6t sa1:aggag) salTs 6u16e:og
,{au puTJ o1 asaa6 aq1 6u1c:ogr pue uals,{s 6ul zetb sTt{l
6u1pe-rbap d1r,to1s a.re uoT1e1a6arr, qs.reur lfes ar{1 uo sleu
1e61e pue 6u1:nocs-acr .dq pasnec acueqrnlsTp pazTTpcoT
pue asaa6 aq1 .f,q 6u1qqn:6 6u1:ds 3io slcaJJa ar{1 '"*T1 6uo1
e .ralo A11n;ssaocns pauTpluTpur aq uec uoTlcpraluT al{l pup
.f,:e1uaurl1dmoc arp qll'ro:6 1ue1d 3lo Ibolouaqd eq? pup asaa6
aq1 ^f,q 6u16e:og Jo su:a11ed ar{1 JT uang 'ua1sr(s 6u1zer6
qsreu ?Tps-asoo6 nous ar{1 Jo ,t+TtTqeas aq1 pue 'q1no:6
1ue1d uo 6urzetb uoseas alpl Jo s?caJJa uual-6uo1 aq1
oluT s1q61su1 auos eplao:d TTTl,r 886T Jo Jauuns aq1 6u1:np
s1o1d TpluauT.radxa g:o 6u11drues rar.{1:ng 'uoTlp1a6an at11
:oJ r(:aaocar Jo poTJad e s1 .rnoT^eqaq sTr{1 Eo acuanbasuoc
lcaJTpur aq,tr 'TeATA.rns luanbasqns JoJ 1ue1:odu1
aq .(eu dlgr 01 aTqe a:e s6u11so6 aq1 aouo spare 6u16e1s
pue r(erra.f,1g ar{1 Jo uotle:o1dxa '(Oget sauulcpl{ pup ?lalard)
:ead auo .roJ pauTe1uTeu sT puoq 6u1:ds;gio-luared aq1 pue
salcads pan11-6uo1 .d1en11eTa.r e sT asoo6 Aous aql acuTs
'osTV '(Af Of ratpg) uoT+Tpe.rl uoT1er61u pue poTradoloqd
6u1sea:cap r{1TA paleTcossp sasuodsa: uo11e:61u aleuT
t{lTA raqle: 1nq 'a6e:og: aq1 Jo d+TtTqetTple :o ,{111enb
ozr
uoaqbT\raNcN pue sser\t ti86t olITTet[ pula u]eu.tpN :T86T .Te
F uTr+og) sassaco.rd uollezrrersur'r reTqo.rcrru 6urlouo.rd ^f,q
suals,tsoce Jo sassacord uolllsodruocap pue sla6pnq e,uaT.rlnu
aq1 loaJ;e KllcarTpuT osTe ^6.eu sa:olTqraq ,6urlcAcar
1uaT.rlnu uo slcaJJa lca.rTp rTar{l 01 uoTlTppe uI . (Bgeet
sar.raJJaf ie 86T 
.Te 1a >1coddo3 jTg6T aleoTg ,O8GT .Te la
1cr:puaycp) palTurTT-ua6o:1ru st sXelTqeq asar{4 uT qry,ror6
1ue1d se 'paulelsns aq 01 sT sp.re&s paze:6 Jo q1l1or6a.: aq1
JT slelTqpq Texnlpu ur Terluasse Illuanba:g sT 6u11c.f,car
sTq,I '(SeOf sarraJJaf pue r(1azeg !J'q,regg6T uolq6negcq)
(:a++Tt) 1e1:a1eru 1ue1d peap uorJ sluarrlnu Jo aspaTar
at{l Jo dals 6u111ur11-a1p: aq1 6urssedlq .f,qa:aq1 ,sseuorq
queld u1 luasa:d sluarrlnu 6u11c.dca: pue 6ulssacord
.f,q s1ua6e bupzyTpJauru sp +ce r(eu sa:oATqJaH
'sacaeJ pue auTrn uT ?PlTqeq
aql 01 pauJn?a.r sT paze:6 a.re r{orrllt slue1d ur ua6o:11u aq1
JO UorlJodo.rd a6.re1 E i srpurrue dq +earqeq p uro:J pa^ourar
sT anssTl 1ue1d ur ua6o:1ru aq1 Jo auos ,(TuO .elarcxa
uT jro .ra1?rt uT ua6o:11u c1ue6:o Jo uor?EzTTerauT_ur 1au
Jo alpr ar{1 ,{q palTurTT ualJo sT s1ue1d Jo r{lno.rg . (faer
1ua:dg JTS6T a1eo1g) s1ue1d trq dn ua{e1 aq r(eur qcTqA
suuroJ oluT pa+raluoc aJe spunoduroc ua6o:1Tu 1pq1 TTos
aq1 uT sT 1I 
. 
TTos aql 01 luauraTa sTr{1 Jo u.rnlaf arll osTe
pue ?uETd aq1 o1 TTos aq1 uo:J ua6o:q1u Jo AoTJ ar{+ uo
luapuadap sT qyro:6 1ue1d sr.uals.dsoca Ter.rlsar:a1 uI
NOITCNOOUTNI T'E
9NTTCI'C NSgoEIIN ONV 9NI9\rlIOd ESOO9 IiONS
N8gl{,tgg sxcvgqggJt EAI,LISOd - HSU\fi{ ,LTVS qgZ\ltl9
v Nr sgssgcoud Nor&vzrrvugNrl{ Ng9ou,trN Tvrgoucrn : s u
TZT
sa@E-6Til{d ffi@E 'sse:6 qsreu-lTes sno:aJTuoTols
e Jo (aavH) uoTlcnpo:d d:eu1:d puno:6-aaoqe 1au
aql pasea:cuT I11uec131u61s esaag rqous rassaT .f,q 6u1ze:6
Apql unor{s arrer{ saTpn?s snoTla.rd '(ergOf saT.raJJaf
pue 1116re3) palTurTT-ua6o:11u I16uo:1s sT ('t suacs-ctruaBc
suacsaTnraec uaqc) asaag AouS .rassaT gro r{uo1oc
6u1paa.rq e sT a.raq? araqA r{eg uospnH Jo lseoc uralsaA aql
uo .fieg asno.;rgd eT le qsrplu lTes aq1 uo qlno:6 lueTd
'(eeet uosdl111qd pue uosqTg aas) seca€J urorJ sluaT.rlnu
Eo 6ug1cr{ca: dq paulelsns aq louuec q1nor6 lueTd 'sa11s
,auTrleTr urorJ paqe:edas .d11e11eds sT prptls 6u1ze:6 aq1
a.raqn sa?Ts uI 'p:pAs paze:6 aql uo sacaeJ Jo uoTlTsodap
aq1 04 pa{uTT I16uo:1s sT saTlTunuuoc pTouTue:6 paze:6 u1
6u11c.dc ua6o:11H 'suralsrlsoca Te:n?eu uT 6u11cdc ua6o:11u
gro .f,ceclJJa aql acuanTJur r(eru uorlcafas a6e:og '(tggt
'11anssog pue {JeTC) sadlq a6e:og pu€ TTos lua.raJJTp
6uoue zt:ea sa11:ado:d esaql acuTs 'aleJlsqns aq1
Jo oT1p.r N:C aq? pue TTos aq+ Jo Ud'TTos ar{1 Jo lualuoc
ralpn se sJo?ceJ qcns uodn luapuadap are 'sluaurlpas aq1
uT rncco r{cTqA'sassaco:d uollezTTTqounuT-uoTlezTTeJauTu
paleTpau -TeTqo.rclur .due1^1' (peef uolq6negcn)
s:aze:6 ar{X Jo .rnoTler{aq 6u16e:og aq1 uodn luapuadap
aq osTe r(eur saqoJcTur pup saJo^TqJar{ 's1ue1d uaaAxaq
6utldnoc luarrlnu aql Jo acua1sTs:ed pue 1ua1xa aq,l
'sluaT.Ilnu IoJ >[uTs e sP pue
acrnos luaT:1nu 1au p qloq se suoTlcunJ sspuroTq IPTqo:cTlu
TTos aqf,, '(egor uolq6nepcl^I pue ssang) s1ue1d .f,q dn ua{tl
Xeql ueql :a1ea:6 sauTl Tpranas aq r(eu sseuloTq TpTqo.rcTul
aq1 q6no.rq1 s?uaT.r?nu Jo xnTJ Tenuue aLIL '(Heor
saTraJJaf pup XTH 'sseng ,896T saT.ralJaf pue dTazeg '.Lg6I
?ZT
lueTd pue 6uTTcIc luar;1nu pue JnoTler{aq 6u16e:og: uaar'rlaq
>tuTT ar{l auTulexa o? Jap:o uT 'pau1r,u-ra1ap araA a6e:ogi Jo
sad.f,l aarql paze:6 qcTr{i*r asoog AouS :assaT ar{1 Jo s6ur1so6
ruo:J sacaeJ r{sarJ uT qgro:6 TpTqo.rcTur Jo scyureu.dp aq1
pue ua6o:1Tu Jo uoTlpzTTe.rauTur lau 'uorllsoduocap TpcapJ
.dpn1s sTttl uI 
. 
s1ue1d 01 aTqeTTerre ua6o:1ru Eo dlddns
aq16u11e.IaTaccez(qp:eatsE@EaI{1JoddvNpacuPqua
ol sppaT asaa6 &ous aq1 .f,q 6u;ze:6 leq1 luaprla sT 1I
'asaa6 aq1 trq parlnba:
lsout sT 1T uaqA por.rad e 6u1:np pacnpo:d s1 a6e:oE d111enb
I{6Tr{ Eo dlddns TenuTluoc V 
. 
(Heef saT.raJJaf pue ^f,1azeg
! Lg61' Tnpps ! Lg6T sarraJJaf pup uaueloy) uoTlcnpo:d uo
as 
=Ad 
uoTleTToJap Jo slcaJJa asralpe arl? JoJ salesuaduroc
uoseas aql uTqlTr-r sasuodsa: qlaao:6 plde: Aor{s o1 Fffi pue
EffiT@n-e Jo ,{lTTTqp aq,t '(Aeef sar.raJJaf pue ITazeg)
sprpAs paze:6un ur 1pq1 o1 pa:eduoo spJpr'rs paze.r6 uT
raq6lq .611uec1g1u61s s1 uoT?exTJ Jo a1e.r aqf,r 
. 
sluauTpas
aq1 Jo acpJrns aq1 uo 6u1a11 Br:alcpqouer(c dq ua6o:11u
cr:aqdsoulp Jo uorlexrJ ar{1 ltq pacueTpq.ralunoc .dlqeqo:d
sT 1:odxa sTql Jo r{cnu , (qtget saT.raJJaf pue ttT6:eC) :Bar(
qcea sseuroTq asoo6 oluT pale:od:ocuT aq deu qs.reur aql urorJ
t-ul + N 6 Z se'qcnur se q6noq11g 
.sluauTpas aql 01 luauraTa
sTrll ;o Ilddns aql sale:aTacce sacaeJ asoo6 uroJJ paspaTar
ua6o:111.1 'ra11TT .ro sseuroTq aueTd aaTT se bulleTnunocp
Jo p€a1suT 'sacae; Jo sseurorq asoo6 :aq1Ta oXuT pa+raluoc
sT anssTl 1ue1d 4eq1 sT 6u1ze:6 asua?uT ar{+ Jo acuanbasuoc
V '(leOr Ilazeg iq?86T saTraJJaf pue 1116:e3) salcads
onl asaql Jo spJpns paze:6un u1 sanTel 6ulpuodsaJJoc
o1 pa:eduoc ?oOT :aao r(q , .>1sur:o1r1 ptcpT{@-
ffi 'a6pas snolpurozrrlJ e pue , .::ap pue .uqT.rcs ( .uT:,r)
EZT
sTt{,I '@ Ff5oTffie',ffi s; sa+Ts AoTTT1,l
asaql uT sse-r6 uouluoc lsour ar{J, .uoT?e1a6aa aq1 aleuTurop
('xI{cTI{ ffiTfrEUE=fF m1as) qcrrq pue (.++nH e(Ire5@e-fq
x-FS z(1u1eu) sAoTTT/q qnrr{s ,1aaa1 apTl r{6Tr{ aqq puo.daq
'qs:pur :addn ar{1 uI 
. 
(qreOf saTraJJaf puq 1116:e3) asaa6
aql ^q paurnsuoc sr rlsrpur lTes ar{1 Jo uoTlcnpord .f,:eru1:d
puno:6-aloqe ar{1 Jo *Og 
^Taqerulxo:ddg 
.q61q ruc g.Z
lnoqp sr uAeT 6u1ze:6 TepT+raluT ar{J, . (Neet saTraJfag pue
.6,1azeg I lg6T saTraJJac pue uaueloy) bulze:6 trq palcaJJeun
are (*s >) slooqs r'\al d:aa pue .ra?,aur a:enbs :ad salcads
asaql Io sq.ooqs OOO? .ralo arp areql . (e fet ureqerc${
pup uasuaf ,sar:ag:gag) spo.re auTTps ssaT uT luepunqr
sr I{cTt{A E-ett-E-@'nF ffi pue F5}6TEbTfrcI €-ffi
.dq paleuruop sr r{s-reur lTes ar{l Jo uol1e1a6a^ ar{f,, .r{sJprrr
+Tes rPpr+ arl? Jo uorlela6a^ aql uo .dlarrtsualur a6e:o3
s6u11so6 pue slTnpe oOo,OE o1 dn ,dTnf d1:ea ro aun1.
elpT uT s6ur1so6 Jo qcleq aq1 ralJv .deg uospnH Jo +seoc
lsa^ ardl uo , (U tlZ 'o16 ,N t?Z ,"gg) ,epeue3 ,eqo1;ueg
,11rqc:nr{C Jo lsea uq EZ ,deg asno.rgd pT ?p rauruns
qcea paa.rq aseag Aous rassaT 3lo s:ged ooog ralo
alTs trpn1s T.z.g
sctoH,tgn z'9
's1ue1d as:r{1 asaq? uro:J pa^T.rap sacapJ uo-rJ
ua6or11u c1ue6:ou1 Jo aseeTa.r aql oA uoTleTa.r uT salcads
a6e:o3 ae.rq1 ar{1 Jo acuel:odur1 arrTleTal ar,ll alpcTpuT
slTnsar aIIJr .,teg asnorsrd pT 1e sAeTJ TepT? ar{l uo qlto:6
nzr
pue sale.r buTIcad . (1sn6nV eZ ,gt) sluauT:adxa o.ar1 lspT
ar{l uT (Telo1 ZT) pasn araA s6ur1so6 :noJ ,tTuO .s1o1d
Teluarur:adxa aq1 uT (sTTsoTffi @ ffauTc."nd)
salcads aarql aq1 Jo qcea Jo spreiqs uo paze:6 s6u11so6
xTs (dtnf et) T 1uau1:adxg uI .srnoq g .d:aaa dlalerutxo:dde
uorlela6a^ paze.r6un 6uruTeluoc s101d Aeu 01 pa^ou a:ar{
s6u11so6 aq,1, 
.d1:e1n6al papT^o:d sen ral.A qsa:J .sacaeJ
Jo uorlcatToc aql 01 :o1:d s.rnor{ oT 01 g roJ uo11e1a6aa
aql o? ,a?eurTcce, 01 paAoTTe aJaA s6u11so6 aqg,
. 
(1sn6nV eZ :e luaur:adxg
llsn6ng gT :Z luauT:adxg :dTnf Ct :T luarur:adxg) Lg6;- uT
suorspcco aa:r{l uo palcnpuoc-a.raA sluarul:adxg .urg x lug
dlaleutxo:dde sear 1o1d qcpg .@ FTTsj-1-6Eft,ep5
ro 'Ea-5EETE-ffi xa.ffi:o 'sapouE@A ffiE rar{1ra
dq paleuTurop uorleqa6al qsJeur-lTes Jo spaJp paloaTas
uo par{srTqelsa a.ran s1o1d Te1uaurT.radxa Jo Jaqunu V
'paurul;ra1ap spA s6ur1so6 aql Jo 1q61en aq1 1uau1:adxa
qcea aroJag .luaurTJadxa ue o1 :orrd sr(ep aq1 uo ldacxa
'pooJ 6op r(:p :o ,euaaao:g {cnq ETgg euT.rnd Jo uoTle: d11ep
E qlTiq paluauarddns '(srrT1effi ffib pue Fa]6TEl6E-(d
effi.ffif, rtluleu) spooJ Te.rnleu Jo laTp € paJ
atrai'I s6u11sog 'duec qc-rpasar aql uT r11rr11dec ur pasre.r
aJa/q, pue Eurqclpq Jo s.rnoq tz uTqlTn asaa6 pTTA Jo s?sau
luoJJ pa^ouar a:a' s6u11so6 aqg, - Lg6r JO Ja'n,'ns aql 6ur:np
s6ur1so6 aar4dec 6ursn palcnpuoc aJa& sluarul:edxa TTV
sTeT:1 Teluaurrtadxg Z. ?,. g
'(q'eggot
sa1:aEg:ag) palTuTT a:e a6e:og: Jo sac.rnos .raqlo ;o sarlddns
uaqn asaa6 aql ,tq paze:6 st sse:6 6u1ru:o3:_Jrn1 ,ras.reoc
9Zr
'coT'o T z'6r Jo arnlpredual luelsuoc p ?e palcnpuoc
a-raA sluaurr-radxa aa:q1 TTe .roJ suoTleqncur .1uaru1:adxa
qcea 6ur:np dn 1as a.raA (1uasa:d sacaeJ ou) re[ Torluoc
auo pue (sTts-.T@ pue *E:t6 ,@Re) sluaurlparl
aaJr{1 aq? Jo r{cea Jo sa1ec11da.r oAJ, .re[ sse16 arlTT 8.0
e ur aEuods paualsTolu p uo paceld pue paq6TaA araA a6e:oE
;o ad^l qcea uo..rJ sacapJ qsa:J Jo 6 oZ ET r{laleurrxo:ddV
'luaurJadxa aq1 Eo 6u;u16aq ar{X lp appur a:rar'r
a6e:ogr g:o ad.dl qcea :oE gd pup lualuoc .ralen TpcaeJ Jo
suorleuTuJalaq 'r{:o1e:oqe1 eql 01 pau:n1al aJaA s6u11so6
.dq paze.r6 s1o1d aql uo:J pa1caTToc sacaeJ r{sa.r{
sasaPJ r{sarJ uT uoTlPzTTe.rauTu
ua6o:1ru Jo sale.r pup saXeJ uo11e:1dsa.r
TeTqo.rcTur Jo selpJ Jo luaua.rnseahl t.z.g
'/tloTaq paqTJssap
sp pauTuuralap ara^ TeT:a1pu aql uT luasa:d suor c1ue6:ou1
Jo sfunoue pup sanssTl 1ue1d asaql Jo slualuoo uoq:ec
pue ua6o-rlTu aqf,' '-rq vz roJ c oo8 le parrp pu. palcarroc
serq saTcads aq1 Jo r{cpa Jo sseuorq puno.r6_aaoqg . stnoq 0T
01 8 snolaa:d aql .roJ bulzetb uaaq peq s6u11so6 ar{1 ralJe
'r{ oorz pue q oooz ueaA?aq palcarroc aJa}r sacaeJ qsa'd
'sadd1 a6e:oE aa.rql ar{1 Jo d+TfTqT+sa61p aq1 Jo acuaprla
lcaJTpur saptao:d sJolecrpur o,o1 asaql Jo luaua.rnspaur
':aqTJ se 1q61aaa .,t:p rTar{1 g:o a6eluac:ad q6Tr{ e ure?uoc
s1ue1d a6e:o3l uar{A r{6Trt ^fi11e:aua6 a:e saler uoT?pcaJap
pup Aor a-re sale: 6ut>1cad aculs .?uaurT.radxa qcea 6u;:np
suorspcco oA1 lsear le uo paurnlralap aJaA sa?pJ uorlecaJap
9Zr
. 
uoTlnTos
alTrorqcod.dq-10uaqd nau qcea roJ pa:eda.rd a.raA apr.rorr{c
urnTuorulne Jo suorlnfos p.repuels r'raN . (OZ_cadS TuTur)
ralauoloqdo:1cads pTaTJ e 6u1sn pauTurJalap spr,r uru gzg lp
slcerlxa pa.ralTTJ JO aCupqtosqe ar{J, .uoTlnTos JO Tur o? uT
papuadsns aJaic s6utddo:p asoo6 qsa:J Jo 6 g rTafeurlxo:ddg
'T!lX I{ T Jo uoTlnTos p r{1T/tl srnoq zT :o3 uoT+cprlxa
ralJe' (6ga;- ouez:o1og) poqlaur alTjro1r{codr{q urnTpos-Touaqd
aq1 6u1sn pauTur^ralap spA sacaeJ asoo6 qsa.rJ go ua60:1ru
arqnros 'uoTlezrre.raurur ua60:11u 1au Jo arnseau e se/{
acuaraJJTp ar{1 iq te .ra?Jp pue luaur:adxa aql Jo 1:pls aql
1e pa.rnseau spn sacaeJ uT luasa:d unTuoluure aTqnTos
ua6o:1tu atqnTos Jo uorleururJalaq V.Z.g
'(raer ppeT pue 
uosuT{uaf) sseuroTq TeTqo.rcrur Jo uoTXeuTurralap
roJ suoTsJaluoc p:epuels uo paseq sT uorssa.rdxa sTr{,1
secaeJ ;Ma b/ (9 + 9. o + 1upr1r1 Tur) = socopJ LMg b/c_ loc 6ru
: sT pa:Tdsa.r loc Jo lunoue aql 01 L. g ol e .8 uro:J ud
aq1 a6ueqc 01 pasn 1up.r1T1 Jo sa.rlTTTTTTrrr lro.rJ uorsJaluoc
aq; 
. 
(:a1atu Hd 9gT TapoI^I Aaunccg .raqslg) TCH N EO.O
rllTlt aldures (S:1; aAnTTp e 6ur1e:lT1 .dq pauTurralap seA
pog:ad q ZT qcea 6u1:np HopN .dq padde:1 tOc Jo lunoure aq&
'q zT .ra+Jp pabueqc araA sde:1 HoeN aq,tr .)trep aq1 uT r{ vz
JoJ palpqncuT areA s.raqueqc asaq,tr .uoT1p:Tdsa: TETqoJcTur
uroJJ palTrap ,roc de:1 01 s.zpf paleas aq? apTsuT paceld
spA TeTA uoTleTTTluTcs sse16 Trr OZ p uT HOeN NT Jo Tur ET
LZI
rfe:-euue6 aq1 gro sarlrsualuT aqfi . (1nc11cauuoC ,uap1:aq
'sarras OOTg 
,:o1ca1ap (Tt)aC p:JaqupC) :alauro:1cads
.f,er-euue6 e 6u1sn spuocas ooz :oJ palunoc ara,o sadolosl
-orper pacnpo:d z(11e1c1grTl:e aql 
^q 
pa?1Tura s.(e: eruue6
aq1 'saldues 1ue1d .roE sa?nuTu Z pue ,saldues fecapJ JoJ
uoTlpTpe.rrT 6u1rrro11ogl sa?nuTru 8 Jo aurT? .de1ep p ralJg .!D{
Z +e salnuTur g roJ r-s + g_llc + suo.rlnaU,,0T x T Jo xnTJ
uorlnau e z(q pa?eTpBr:T airaA saldueS . TeT,, aualr(q1ad1od
(uc T-T x g.Z) TTplus p uT pa6e4ced sen aldures VqNI
qceg 'suoTleTppJ.rT TTe rOJ pasn seA oluo.:rof,, Jo rllts:aaru61
aql 1e rolcpar gyodt{oTs aq& . (WHf ) slsIleup uoTle^Tlce
uo.Ilnau Teluaulr.IlsuT 6u1sn (,-,fUO 6 + alorurfl urnTcTec
pup 'apT:o1qc ,urntsselod ,unlpos ,urnlsau6eur Jo slunoue
roJ pasdleue ajran s1ue1d pue sacaeJ Jo saldues paTJq
sts.dleue uoT?elTlce uo:lnaN g. Z. g
,r1ra11cnpuoc ffii-':'l:'":: 1:l:]::.::; =":""","11;" TeluautaTa j3oog6 Xe pau.rnq araA seldrueg ..ras.d1eue
ar{1 ur uorlsnquoc eroJaq paq6lairra:d sen (6ur oor -xo:dde)
alduesqns e pue TTTru traTTU p uT (az1s qsaur oZ) puno:6
sEr'r TeTralpur TTV . (VSn ,.Ir{ ,qdaso4 .1S) :asz{1eueo1ne
NH3 009 sar-ras ocET e 6u1sn apeu a.raA sanssrl 1ue1d pue
sacapJ paTrp g:o saldurps Jo lqbgarr d:p aq1 Eo a6eluac:ad
e sp ua6o:11u pue uoq.rpc Jo sXunoue Jo sluauernsealil
suoTleurnlralap ua6o:1tu pue uoqJeC g.Z.g
8ZT
O0T x [ (.{qV TecaeJ g / agg 1ue1d g)-r] = tr1TT1q;1sa61p *
:sen fr+ltTqT+sa61p a?eTncTec
o? pasn uorssa:dxa aqJ, 
. 
(geet eTaTTeA pue uosTTl{
'urneqsqcng) sacaeJ o1 pa:eduoc sluefd uT (gqV) aJqTJ
1ua6:a1ap pTcE Jo oTle: aq? uo paseq a:aA ,I+TtTqTlsablp
Jo suoTleTncTeC 
.oTJeXuO ,qd1an9 ,poo,{ pue arnllncg:6g
go I:1s1uT!{ oT.rpluo , sa1:oleroqeT poog-1:6g llq pasdleue
a.ran saldures luBTd pue sacapJ paT-rp Jo (eeOf lsaos
upA pup luauuaq) a:qTJ 1ua6:a1ap pTce Jo saTlTluenO
sasr{1eue arqTJ 1ua6:a1ep pTcV L.Z.g
.6ur1unoc pup uoT?eTpprJT uaanlaq
sadol0sr anllcedsar eq1 Jo r(ecap aJTT-JTpr{ foJ palcaJ.roc
aJaA slunoc TTv 
. 
sp.repuels aq1 uro.rJ pauTp?qo :aqunu aq1
t{lTn1 slunoc Jo .raqunu aql Eur:eduroc ,(q pa^rrap sp ,1uaua1a
qcea roJ clglcads :olceJ uoTs.ra^uoc p sT X pue ,pa.rnseaur
6u1aq s.de.r euue6 ar{1 JoJ .ralauoJlcads ar{1 Jo 6ulpea-r
puno:6:1cpq aql sT g pue 6urpea: >1ead aq1 sT d a.rar{i{
(X x sseu aldtues) / (g-a) = uoTlerluacuoc
! eTnnr.ToJ ar{1
.dq suoglerluacuoc oX pel.raluoo a.ran slunoc uoTXeTppE
'a^oqe paqTJcsap spoqlau aq1 rq pasdleue pup pa:eda:d aJaA
sp.repuels TeluauaTa , suorle:luacuoc Tecruar{c 01 s?unoc
uorlPTpeJ +ra^uoc o,l 'suot6a-r 1e:1cads palcaras r(11enueur
Jo uoTle:6a1u1 :asr(1eue crleuolnp xq pauTurJa1ap aralc s>1ead
6Zr
'uospas aq? ssorce sa?e.r uoTlpcaJap uT sacua.TaJJTp
acuecTJTubTS OU a.ra/q aJaq,r '(t-g arqB,l) sluaurJadxa oarl
AsrTJ aq1 uT Effid@f, :o *EE :o; aler o1 pa:eduoc
aran uorlecaJap Jo saleg 
. 
(f_E aTqe&) luaulearl
r(ue :oE uoseas er{1 Jo asJnoc aq1 ralo I11uec1gr1u61s.
abueqc lou pTp saler Fu1>1ca6 .*-+lF=ffi Jo sapeTq
pup suals lca.ra ,JJT1s aq1 6u111q dllncTJJTp peq s6u11so6
4er{1 pa?ecTpuT suoT?plrasqg 
. 
(f-E aTqe&) sluaut:adxa
ea.rq1 TTe uT ffi :o xarec uo 6u1ze:6 s6u11so6 o1
pa:eduroc saler 6u1>1cad Jar'ror peq 
- 
rEeruffi uo 6utpaag:
s6u11so9 
. 
(f -g aTqp;) luaut:adxa qopa .roJ adr{l a6e:og:
qcea uo aze.r6 01 uopueJ Ap pelcaTas sdno:6 aa:r{1 aql Jo
s6ur1so6 go 1q61eA aq1 ur sacua-raJJrp 1upc1g:1u61s ou aJart
arar{1 : spol:ad f pluaurradxa aa.rql aql JoJ Ilaallcadsa:
6 ZngT pue V6VT 'gLt pa6e-rale 1q6larrr 6u11so9
s6ut1so6 Jo .rorleqaq 6utpaag T 
. 
t. E
S.ITNSSU E'E
'dpn1s sTrtl Jo suoTsnTcuoc uTeu aq1 lcaJJE
1ou saop srVAoNV aq1 Jo uregac uT s:o.rJa If pup I adzk
Jo acuasa:d .f,1a>1r1 aq;, 
.AoT sen slsa1 TecT?sT1p?s aq1
3:o -ranod eq1 ',sluaur.radxe aq1 Jo l'!eJ e ur /'lor sear saldues
Jo raqurnu aql asnecag 
. 
(teOf JTqoU pue Tptos) sa:npaco:d
TpcT?sT1e1s pJppuels 6u1sn pazr{1eue a.ran pfeq
Plep ;o sgsdleue TecT?sT?p1s g.Z.g
oeT
OT 'T
+oT'?
++06'5
++TT 'OT
E0'?
+++6L'OZ
9t'o
6z'r
EO'7
0E'€ = I'2,9.
g'zt 6'6
TT' Zd 9'TT
6'Zg. O'6
SFSTFEuleftj
Z8'0 = OT '2,{
6' Zg. E'6L
g'2.1. L' LL
9r'zg, g'nL
FT=so'r6ffiTe.;
9Z'o = 9'zL
9'L
L'L
L'9
@
( l-e I seoaeJ)
ZZ'T = TT'eJ
7'L
5'8
r'6
xa.rPc'dxg
alE.r uorlecaJap (c
g
z
T
++90'6 - OI'Zl
g. ?TT
9'9ZI
8'007
eTTTaurccnd
( ,-u1ut *
09'0 = OT '2,{
t'eTT e
g'ezl z
O''TT T
xaffi 'dxg
s>1cad) aler 6ug>1cad (q
+++29'T9T = OT'Zd +++9?'OL = OT'Zd +++ZZ.gE = OI'Zg,
g'zI 9I.8T LELI ET6T
g'ZE, ESTT cSET LIiT
sr' zg. 
-toL T98 TgL
FTTs6-:ffiup-
-ffitlt.ccne
e
z
T
xa.rPc
(6) 1q61aat
'dxg
6u11so6 (e
(fOO'O > d'+++ iTO'O > d'+,t JgO.O > d r+)
sulnToc unop pue siqoJ sso:ce elep JoJ uaa16 a:e ,srVAoNV
r(er.r-auo uo paseq ,sanlea g . (1sn6nV gZ e .dxg tlsn6nv
gT Z 'dxg ldTnf gt T .dxg) sluaurT:adxa 6utpaag: aarr{l
ur '(@ FT=Ffi@5 pue EE@dffinE xeffi
'@bEabE'tfd @E) sadrl a6e.roE aarql uo Fulpaag
asoo9 AouS -rassaT aq1 Jo s6u11so6 gro (c) a1p.r uoTleoaJap
pue '(q) aler 6u1>1cad , (e) 1q61an 6u11so9 .T-9 aTqe,t
TCT
' k-s aTqe,l) 3-f1-sl$T6- :og asoqX se a6:e1 sp
acTA11souITeaIansanTeAaq1puP,:e11ur1sa.IaA@d
pue ffi Jo sseuroTq puno:6-aloqp aq1 uT luasa:d
unTcTec Jo slunoug 'sluaurrJadxa aarq? aq1 uT ?uapTla
seJ{ saTcads aa:ql aq1 .roJ s?uauaTa asaql Jo slunoue aq? uT
puaJl lualsTsuoc ou 's11nsa: aq1 uT uoTleTJpA aTqe:apTsuoc
sea aJaql q6noq+TV 'satcads aarql TTE uT reTruTs a:an
untsau6eu pup unTssplod Jo slunoiug '(z-s aTqefi) uoseas
aql lnoq6no:q1 F1{56Eberue1e3 u1 lseeT pue e111au1ccna
uT 1sa1ea:6 aJe apTroTr{c pue urnTpos Jo s?unourg
'(z-S aTqe&) z pup T sluauT.radxg uaarrrlaq teg r(q pasea.rcuT
1nq 'I luaut:adxg uT lsal{oT sen oTle: N: C aql @e
puB *Effi ut 
. 
g pup T sluaurTjradxg u1 ETTso-t6@
uT 1saq61q sei'r oTleJ N:C aq;, 'uospas aq1 Jo as;noc
aq1 6u1:np asearcur 01 papual 1q61an .d:p aq1 go a6eluac:ad
e se uoqrpc Jo slunouv 
. 
(z-s aTqp&) STASo-rFeffi
.ro ffi-ffi6 :og sanTel Fu:puodsaJ:oc o1 pa:eduoc lqblarrt
^f,:p aq1 g:o abeluac:ad e se uoqJec Jo slunourp 1saq61q
aq1 pauTe4uoc xarec 
.5!{5o-:-6-- :ogl z luaurT.radxg
uT pasearcuT lnq 'ffi-[- pue *Effi u1 uoseos
aq1 .rano pasee.rcep lualuoc ua6o:11N 
. 
sluaurTJadxa aaJr{l
TTe uT FTasoT5E-rue- pue *ffi roJ sanTel o1 pareduoc
1q61arra .fi:p aq1 gro a6eluac:ad e se ua6o:11u Jo slunoue
1saq61q aq1 peq 
-e@ffi 
. 
(Z-s aTqe&) sluaru;.radxa
6utpaagl ar{1 uT pasn salcads Auefd earql ar{1 Jo uorltsodluoc
TpcTuaqc aq1 uT punoJ aran sacua.raJJTp luecTJTu6Ts
@- f6erue1e3pue
Pa€:t11edEqns xa:ec 'sa@6E[d ffid
Jo slueTd Jo uolllsodruoc TpcTuraqC Z.e.g
ZET
cLI gVV
66 VOV
z9I LO9
6E'TZ 9 e'67
tT'8T 98'87
gL'TZ 88'TT
Z9'Z = t,'Zg. +++LB'9tZ = EtZg.
E'zE (er') eo'z (to') Ez'z
E'z,tr. (ez') zE'z (go') et,'z
E'?,,8 (lr') 08'T (go') 69'E
+++69'ggT = e'ZE
(eo') vvz e
(eo') zv z z
(ro') zE'E r
vtz
LVI
602
T.9'OZ
t8'oz
zz'eI
e
z
TPN
g
z
T N:C
?e 'T
87'0
+++ZZ '98t
vv'I
++e0'?7
++86'eZ
++tO'ZE : I'Z[.
t.' z,s. Gv') v' e v
E'zE (st') z'zv
?'zE (tz') z'68
9L'T = Et Z&
(ce') o's?
(ro') 6'ev
(er') L'zv
9T'0 = Et?^1
(sg') T'vv
ftz') z'v? (tz') 
B'e ?
e
z
T
F11so=@e111au1ccrrdxarec
'dxg
luauraTg
'(ngS f ) ueau are paluasa.rd sanTe1 
. 
(fOO.O > d ,+++ jTO.0
> d'{.+ JEO.O > d '+) sunToc uAop pue sr,roJ sso.roe plep JoJ
uaa16 a:e 'sTVAONV .derra-auo uo paseq ,sanlea g . (sluaruala
.Iaqlo TTp-roJ T:rr lNpueSJoJ ?=u) .1_,1&Q 6 + lorur/
sp passa:dxa sluauraTa -raq?o JoJ sanTe^ isseuorq puno:6
-a^oqp ar{1 Jo 1q61an d.rp aq1 ;o a6eluac.rad e se pa1:roda:
arp sanTel N pup C 
.sluaurJadxa butzezb aaJr{1 aql uT
s6u11so6 .f,q paurnsuoc a6e:og @ FTASotEe-uffi
pue EEE!{E'ffiS ffi5 ,FeFEIiEbTGrq-et @fre Jo sseuorq
puno:6-aloqe ar{1 Jo uorllsoduoc TeluauaTg . z_9 aTqp;,
eeT
997
9Vr
9Zr
?OT
ezT
ETT
Eg9
ne9
Ee9
67,2
962
LTE
9LI
T9e
z9z
9L
097
097
L8t,
99n
VTL
8TE
LET
?,29
z9z
I6Z
tTz
L8T
r0r
?OT
999
E89
et8
9ez
€6e
T9C
g
z
T
€
z
T
g
z
T
g
z
T
eTTTauTccnd
'dxg
xalec luauaTg sFso-.:ffieT$
teT
panuTluoc z-g aTqe,L
uT 1sa6:eT a.raru suoT apT.roTr{c pue unTpos Jo slunouv
'(e-S aTqe,l) Z luauTradxg
roJ ldacxa '5fffiTbEEffi :ogl 1sa6.re1 .d11e:aua6 serrt
sacapJ aql Jo oTlp.r N:O ar{;, 'salcads aarql TTe Jo s1ue1d
uto.IJ pa^T:ap sacaPJ uT uoseas eql Jo asJnos ar{1 ra^o
uoq.rec Jo lunoue eq? uT asea:cuT e spi*r a:ar{& ' ( g-g aTqefi)
luecTJTu6rs 1ou seln acuareJJTp sTr{l 1nq 'E-fi$'mffift]i
:o FE uo pal qcTrdn s6u11so6 uo.rJ sacaEJ roJ
s1Tnsa.I6u1puodsa.IJoco1pa:edruoc@uo:Jpa^T:ap
sacaeJ uT punoJ arai'r uoqJec Jo s?unoue lsaq6Tr{ aq,l
.-sT1so=6ffi
roJ ZLI pue ffi-if!ffif, :og Z6z '*EE :oJ *zz ajraA sanTpl
asaql sluau1:adxa aa.rq? TTe jralo pabe:eag 's6u11so6
aq1 z(q paurelar sei'r ua6orlTu qcnu Aoq Jo JolecTpuT
q6no: p sT (a,roqe z-g aTqe;,) anssTl 1ue1d puno.r6-aaoqp
ar{l uT lunoue aq1 01 pa:eduroc sacaeJ uT 1q61an d:p aq1
gio a6eluac:ad e se ua6o:1Tu Jo ?unoue aql uT asea:oap ar6,
'ao.rnos a6e:og aql Jo aarlcadsaJJT uosees ar{1 Jo asJnoo
aI{? Jarro I11uec1g:1u61s paspaJcur osTe secaeJ Jo lualuoo
ua6o:1ru aqg, 
. 
sTTsoTSerneTEc uo pal r{cTr{A sprTq uo.:rJ
saoaeJ uT anTel aq? 01 pa:eduoc a6e:og: Effi@E
pue xa.reC uroirJ parrT.rap sacaeJ uT sen 1q61a.,a r{:p aq1
;o a6eluac.rad e sB ua6o:1Tu Jo lunoup lsaq6gq aqg, 
. 
(e-g
aTqe,I) sluaulr.radxa 6utpaa3l arll uT pasn salcads lueTd aa.rql
aq1 uo 6urze:6 s6u11so6 aq1 uo-rJ sacaeJ aq1 Jo uolllsodruoc
TecTuraqc aq? uT punoJ araA sacuaraJJTp lupcTJTubTS
@ F?Tso:bEffi pue
xarec' ffi ouebF-r{el _ETtfr-ir!ffie
s6ut1so6 uo.rJ sacaeJ Jo uoTXTsodruoc
eacern@-
uo 6ulze:6
TecTual{c g'g'9
9CT
ffid uroJJ sacapJ suor1eqncur TecaeJ aq1 Jo srnoq
zI lsjrTJ aql uT :alaAor{ ,E-.r-+G:i6@TE5 :o Effi-ffif,
luro.rJ palT.rap sacapJ uT s.rnoq tZ :alo uo11e:1dsa.r
TErqo-rcTur Jo sale.r uT sacuaraJJrp Aupcl;1u61s ou
a.ran a:aq;, 
. 
(q,et-g aTqe,t) Fl1E-o E :o E@ffi
paze:6 qcTr{A s6urlso6 uorJ sacaeJ uT uer{1 ,mi
uo pal qcTqn s6u11so6 urorJ sacaeJ u1 raq6lq t?f pup
tgg 'a6pJaaB uo 
,aJaA Ilaallcadsa: pol.rad rnoq tZ E pue
.rnor{ ZI e .ralo uotqerldsa: TpTqo.rcTur Jo saler aqtr
sacaeJ Jo uorleqncuT aq1 6u1:np uoljezrTpJaurur
ua6o:11u pup uoTle:tdsa.r TpTqo:cTn ?. e 
.9
'(e-g aTqe,I) t.4 pue t.9 uaanlaq paT.rel pup ,s1uau1:adxa
oAl 1s:TJ aql JoJ paJnseau tr1uo sei*r sacaeJ ar{1 Jo
Hd aq,tr '(e-g aTqe,t) luaraJJTp I11uec1g1u61s 1ou a.raA sad.f,1
abe:og aarq? aql urorJ palrrap sacaeJ uT ,(_,fmO 6 + N- fit"l
ua6o:11u afqnTos Jo slunoue ar{J, 
.ffiEFE :o *EEg gro
slualuoc aq? uer{1 .ranoT Zv ,a6e.rarre uo ,spA F--TlE-Tffi
Jo lualuoc ;ralpn ar{J, 
. 
(e-g aTqe,I) salcads aa.rq? aq1 uoJJ
pa^T.rap secaEJ ar{1 Jo ?ueluoc .ralel\ eq1 uT sacua:aJJTp
1uec1g1u61s aJar'r aJar{1 ralanor{ ,Jauruns ar{1 Jo asrnoc aql
relo d11uec13l1u61s raJJTp ?ou pTp salcads a6e:og aarql aql
Jo qcPa uo.rJ pa^Trap sacaeJ ar{1 Jo lualuoc Jalp/q aqtr'
'(e-g aTqe&) FTasor6-
pue Effi-ffif, uo pal qcrqA s6ur1so6 Jo sacaeJ ur slunoue
aq1 r{1TA pa:eduroc xaJec uroJJ perrT:ap sacaeJ uT 1sa1ea.r6
osTp a.raA unTcTpc pup urnTsau6eu ,unlsselod go slunouq,
'(e-g aTqe,t) ETlEl6EEre1ec- uo pal qcTr{A s6u11so6 Jo sacaeJ
uT lsar'roT pue ,abe:o3l ffi uro.rJ palTrap sacaeJ
9eT
++99'VZ
nn'z
++TO'LE
?e '0
tz'o
rz'9
+'(Il'lv
E' zg. (go')
E'z,tr (or')
E'zE (ro')
++8L'08
e '2,!. (oo')
t'zg. (oz')
E'zt (so')
v6'vz
zv' 6T
89'TE
= EtzE
89'T
9T' Z
9Z'r
= Etzt
6'TV
8'Tt
6'6e
C6'LI
LE'VZ
98'n?,
++08'82 = C'ZE
(ro') 6e'z
(ro') zL'T
(ao') Le'r
z9'67
ET'?,2
T9'02
tg'Q = E'28.
(go') Bo'z
Qz') rG'r
(ro' ) z6'r
e
z
T
t
z
T
V9'O: E'2,9. +++L?'9ZT = e'ZI
(oo') 6'zv (eo') e'ov e
(ar') 6'rn (oo') s'Tr z
(eg') t.'rn (n') r'6e r
FflsoT@PTTTauTccndxaJec
'dxg
luauaTg
'(WgS t) upau are paluasa:d sanTpA 
. 
(1OO.O > d ,++f
iTo'O > d'++, J9O.O > d '+) sunToc uAop pup sJqo: ssoJcE
elep .roJ uaat6 a:e ,sTVAONV ,ter,t-auo uo paseq ,sanlerr g
'(gd pue 'N- luH ,ozHg :oJ e = u JslueuaTa Jer{1o TTp roJ
T = rr JN pue C roJ Z : u) 
.|-&MO 6 + lourrf sp passa:dxa
slueuraTa .raq1o JoJ sanTpA i sseuoTq puno:6-aloqe ar{l Jo
1q61an .(:p aq1 ;o a6eluac:ad e sE pa+roda.r a:e sanTel N
pue c 'suoTleqnour aq1 Jo 6u1uu16aq aq1 1p sacaeJ r.{sarJ
f,oJ paurmralap sacaeJ gro gd pue , (sacee; l_ .1,Ma 6 + N- 3HH
6tu) H- tgH p.t= (t) OZH Jo slunoure pup sluarul:adxa 6u1ze:6
aa.rql aq1 uT a6e:ogr @ F!{5oT6E@frj pue
eaceq-e@irs xarec 's@Gl6tr:{d E]ffru-ccnE uo 6u1ze:6
s6u11so6 Jo sacaeJ Jo uoTlTsodruoc TpluauaTg E-E afqe,l
LI.I
8L'?
LC'O
LT'T
+8?'6
09'z
++?9'67
z8'I = 9'zt
9'z,t (or' ) ?T' E
9'zs. (gr') gE'z
9' ?,8 (tg' ) gv'z
9?'Z = 9'ZE
s'?,8 (o'r) e'?B
9'zE (ot') T'rB
9'z^tr (zz') B'Eg
LT'Z = 9' z,tr
(rs' ) ro' ?
(oe' ) Es'z
(sg') e6'z
ZE'Z = 9'ZE
(or') o'sB
(er') o'eg
(gc') s'Lg
T9'T = 9' z$
(lr') Lt'z €
(gr') 8s'z z
(re') 6L'r r firu
Z€,'O = 9'Z[.
(or') o'sB e
(t'e) 6'68 z
(ez') z'ss r oz*z
6V
9VT
L6
?OT
TET
n6
9t.n
98V
9eE
Lev
00e
vov
66
9TT
z9T
L6
89
9CT
6ZT
LZZ
897
8LZ
99?
91L
OTL
8ZL
zvol
9LZ
LZV
T9r
?87
9rT
9Lt
6nE
LIZ
6ie
90v
98V
?E8
66t
69V
9LL
9ZZ
e6z
9Ze
g
z
T
g
z
T
e
z
T
e
z,
T
€
z
T
6n
P3
TC
ETEso-r@e@dxaJPc
'dxg
luauaTg
8eT
panuT?uoc g-E aTqP,tr
(er') oe'e
(rr' ) sv L
(zt') Ls's
(rr') zz'L
Gz') Ez' L
(or') ve 'L
z
T
sr?soJbpupTeCeTTTaurccnd
'dxg
luaueTg
panuTluoc
'(pl-g arqe,r)
-sTTs-o-fffimT6 
ulorJ pa^Trap sacaeJ o1 pa:eduroc EJJfr-ffiJ
:o *EF5 uo.rJ pa^Trap sacaeJ uT r{6Tq se acTAl ,a6e:aae
uo 'sp/q (dcualclJJe uoT?ezrTeJaulu ua6o:1Tu) pa:rdsa:
Y
'oc Jo lTun :ad pazrTpJauTu ua6o:1ru Jo lunoue aq,l
'(cl-E aTqe[) (Z quaurt:adxg) ,{1n4 ug palcaTToc sacaeJ .roJ
lsaAoT pup '(e quarll:adxg) uospas aq? uT alef pa?caTToc
sacaeJ roJ 1saq61q .f,11e:eua6 a:an uoTlezTTeirauTu ua6o:1ru
lau Jo saleu 
-ffi5 pue e1ffr-ffiE ,sTTg[jI6€ffiTE5
uroJJ paATJap sacaeJ ;o ua6o:11u Tp1ol ar{1 Jo
Alaallcadsa: Zz.sz pue t9.eT 'ZL.L Jo alpr uoTlezTTetreuru
E oJ lualenrnba sT sTr{,tr . (c7_g aTqe&) slueurr:adxa
aa:q1 aq1 ra^o pa6e:ane ,FTiS pue ,eTTT@e
'FflFa-@-J uo 6u1 zetb s6u11so6 urorJ sacaeJ .roJ ' 
rllaallcadsa: |-Q VZ + f _sacapJ fiMCI 6 + N_ 3gU o* L6. v o1
z9'z of TE.T uro-rg pa6ue: srnoq ?z paASpT 1eq? suorlpqncur
uo paspq saoapJ Jo salpJ uorlezrTeJauTur ua6o:1ru 1ag
'(q'pr-g arqp,r) palcalap aJaA uo11e:1dsa: Tprqo-rcrur
Jo saler uo (sluaur:adxg aa.rq? aq1 uaanlaq) slcaJJa
Teuospas 1uecrg1u61s oN 
.paspaTa.r araA sale.r+sqns
d.ro1e:rdse: se qlato:6 TpTqo:cTur plde: 01 paT r{cTr{/*t
pa.rrncco sTTac Jo sts.dlolne 1eq1 6ug1sa66ns ,uotleqncut
Jo srnoq zT lsfTJ aq1 roJ paprosa-r saler aql aTqnop upql
arou a.raA s.rnoq ?Z uo peseq uolqe:ldsar Jo sale5 . (el
-9 aTqe,I) sTl,soTffiUTeO uo pal qcTqA s6ur1so6 ;o sacapJ
. 
utql uolqe:1dsa.r Jo sa?eJ Ja/'IoT peq d11ua?srsuoc s1ue1d
0tT
*98'eT = E'zg. 99'g = E,z& LE.I = e'zt
oz'e = E'zt (rr.o) tz.gt (et.z) re.et Gz.z) oz.rz
+++96'66 = E'Zg. (ee'o) erEr (ug'o) tg.e (re.r) eo. &
0T'9 = e'zg ftz.t) za-e (st.r) n- n (re.e ) zt.zz
sTa-65ffi xffi
( 1-:Q nZ + ;-sacapJ ,trt{O 6 + ?OC 6,u) (tZf - oa) uolqe:1dsag (q
e
?,
T
'dxu
e
z
r
z8'n
+++t,L'nge
5C'g
9L'L = e'Zt
(gz'o) at'z
(ero) ve'z
(ero) ee'r
0t'0 = E'2,1.
(ro'r ) zs't
(ro 'o ) eo 'r
(ee'o) gt't
E9'f = e'zt
(go'o) rr I
(zt'o) oa'v
(za'o) sg'e
e'za
E'?&
e 'z.l
ETTFoT@
( r-r{ zr + f -sacapJ ,luo 6 + Eoc
xa:e3 
.dxg
uotle:1dsag (e
eTTTauTccnd
6lu) (zre - 01)
'(Wgs f ) ueeu a-re paluasa:d sanTeA . (roo.O
> d '+++ JTO.O > d '++ 3EO.O > d ,+) sunToc uAop pup
sfi,or sso.rcp ElEp JOJ Uarr16 a:e ,srvAoNv 
^ear_auo 
uo pasPq
'sanlea g 
. 
(E 'Z ,T) sluauT.radxa 6ulze:6 aa:ql aq?
roJ suoTlpqncuT 6u1:np a6e:og @ ETForEEu6g65
pue EEceq{E@s xEE ,saF-oTE-6TrTat ffie uo 6u1ze:6
s6u;1so6 Jo sacapJ Jo , (p) (uo11e:ldsa:/uollpzrTp.rauTur)
dcualclg;Ja uoT?ezrTe:auTur pue , (c) r{ tZ, pup
0 uaaAlaq uoT?€zTTe.rauTur ua6o:1tu 1au , (q) q ?Z pup O pue
'(e) q zr pue o uaanlaq uorxerldsa: Tprqo.rcrl^I -?-g arqp,tr
T?I
(ro'o) tr.o
(eoo'o) ?o'o
(eo'o) oo'o
(oo'o) eE'o
(ro'o) gr o
(to'o) zt'o
(ro'o) ez.o
(ro'o) eo.o
(rro) oe.o
ffi5
E
z
T
'dxg
e
z
T
+90'eZ
+€T'ZT
+LT'ZI
sFE-r@-.; -Pm@ilE
('iH=3$zxr,l;i"?3;:Tjt$r3 ;"r5jg?.gH"ITil-"3gS"iTi, 
(p
T0.9 = e'Zg. ++ZE.eg = E'Zg. Lg.9 = E,Zi
E'z& (ts.o) oz-z (ge.o) az.v (rr.o) to.g
e'zt (eo-o) tg-o (zo.o) sz.t (oE.o) tE.z
E'zs. (re 
.o) rc.o (ao.o) eo.z (le.r ) rc.g
STTs6T6-EETE'
( 1-rt VZ + r-sacaeJ ,trMq 6 + N- 3Hn Or)4TFTf6n-e FE5 
.dxs
uorlezTTejrauTur ua6o:311r1 (c
ztT
panurluoc t-g aT
lua.raJJrp e aleq deu s6u11so6 aalldec aq; .atrpc qlTr't,
appu aq pTnor{s ualsds Te.rnleu aql uT suoTlTpuoc aql 01
suoTleTode.rlxa aroJajraql pue ualsds TpTcTJT1:e up uT aceld
6u1>1e1 sassacord luasa:da: sluauT.radxa asaq1 Jo slTnsaJ
aq+ lua1xa auos otr 
.sacaeJ r{sarJ uT uoTlEzTTerauTur
ua60-r1ru ?au Jo sale.r pue qlro:6 Tprqo:crur Jo sclureu.f,p
aq1 pup a6e:oE ar{l Jo I+ltlqT+sa61p aq1 ,s6u11so6 aq1 glo
.rnoTaer{aq bulpaag eq1 palcaJ;E salcads a6e:o3i aarql aq1 Jo
uor4rsodruoc Tecruraqc ar{A uT sacua.raJJTp luecTJTu6Ts
NOISSnJSIC V'9
'(eeOf lsaos uel pue luauulaq) luasa:d ec111s Jo lunoue
aql se qcns ,dlTtTqTlsa6lp 1ua:edde AcaJJe 1q6Tur r{cTrIA
sJolceJ -raq10 Jo acuanrJur ar{1 .roJ palcarJoc 10u afaJq
sanTel asaql pup ,aldues auo Jo aJqTJ Tp1o1 aq1 uodn paspq
saleuTlsa apnrc dluo a:e ^d111tq11sa61p Jo saleurrlsa asaql
1pq1 pasrseqdua aq lsnu 1I .e luaurlradxg uT appu aq pfnoc
sFFrirFETF Jo d1TTTq11sa61p Jo alpurrlsa ou ra^aAor{
'uoseas aql .ralo I+TfTqTasa6lp u; auTTcep p aq o1 s:eadde
araqt salcads earq? TTe uI . (s_g aTqe&) 
-- TbEia-ffi pue
FE6' o1 pa:eduoc E]fr-ffif, u1 qsaqblq s1 I+TfTqT+sa61p
'1e:aua6 u1 
.sanssrl 1ue1d uT 1er{l o1 pe:eduroc sacapJ
uT luasa.rd arqTJ 4ua6:a1ap pTce Jo s?unoue Jo stsr(1eue
aq1 uodn paseq a:e .d1111q11sa6rp a6e:o3r Jo saleuTlsg
sad.ttl a6e:og: aarql ar{l Jo ,{fTtTqT+sa61q 9. e . E
CVT
9'tr
9'6
T'OT
o' ?z
9' LT.
?'e r
8'tz
t'9?,
g.ET
n' Lz
9'rg
uPaIrI
g 'dxg
z 'dxg
T 'dxtr
ffi5sT-fT@ E-TTF-ffie
I+TtTqTrsa6lp g
.(r = u) s1ue1d asaql 
uo 6urzez6 asoog Aous -rassar aq? Jo s6ur1so6
.dq pacnpo:d sacaeg pue slueTd Jo sspuroTq puno:6_a^oqp
Jo aJqTJ 1ua6:a1ap pTce Jo oTler er{? Jo sa?euTlsa uo paseq
a6e:oE @ FTas-rffitE5 pue EETEi[fEffis **6
'FapbTAETEA ffie Jo dlTTTql1sablq .s_E aTqe.tr
vnI
paTdnoc ,asaa6 Jo s?uaue.rrnba: TeuoTlTJlnu a6:e1 aq;,
. 
uoseas aq1 6ug:np sadz(1
abe:ogl luaraJJTp az{TT1n r(aq1 se ,rIeg asnoJsrd eT Ae asaa6
ur -rncco deu saEueqc 1ec160101s,{qd pue recTuoleup r€TTurrs
'(Orct 66e:g puE uospnH ,uo1:ng) aJqTJ Jo slunorue q61q
6ulu1e1uoc pooJ Jo uoTlpzTTTln luaTcTJJa aql uT s+fnsa.r
r{cTq&l uolleqdepe ue luasa:da: deur spuno:6 6u1:alutar
rTaql uo asaag AouS rassaT Jo sp:ezz16 go slq6garrt
pue auTlsalur rrEurs ar{1 Jo r{16uar aq? ur sabueqc rpuospas
'ProrJ 1n6 3ro uolllsoduroc aql uT sabueqc 6ulpuodsarroc aq
osTP deu e:aq? pue '(gtet TaTTTI,I 1996T auolep) slaTp arqTJ
r{6Tr{ o1 paldepe s{cnp JoJ pa1:oda: uaaq ser{ (aso1n11ac
Jo uAop{earq Terra?ceq Jo afTS ar{1) aunTol Tecapc
uT sasea.rcul 
. 
(geor {TH) r{c+Er{ 6u1rrto11og po1:ad 6u1:ea:
-poorq aq1 6ut:np saqs:eu :alen_r{saJJ pue _1Tps uaanlaq
.d11uanba.rg alou osre asaa6 auos .uounuoc s1 sf{FoTffi
a:aqr[ ,sa11s pasodxa ssaT oluT alour dar{1 Furuaaa aq1
6ur:np 1nq 'sprens @e-xarpc uo dlaalsualxa a6e:oE
daqe .dep aq1 6u1:nq .^d6o1o1s,{qd an11sa61p rTaql lcaJJE
.deu qclqr'r auTl Jo spol:ad 1:oqs tr1arr11e1a.r .raao laTp uT
sa6ueqc 01 1ca[qns a.re rleg asnor-ad eT le asaa6 pTTAI
.selcads asaql uo paaJ
asaa6 prrA uar{A suorlTpuoc re.rn?eu .rapun fncco op salcads
aa.rrdl asaq? uo.rJ pa^Trap secaeJ uT palca1ap l(lgnglce
TeTqo-rcrur uT spuaJl TTe-ra,'o aq1 1eq1 4qnop 01 uoseal ou
sT aJaql 'raaaatoll 'IfTtlqllse6lp a6e:oE pue uoT?ezTTerauTu
ua6o:1tu ruo11e:rdsar TeTqorcTur uT sacua:aJJTp
aql oA palnqTJluoc alpr{ deur paslpJ a.raA s6ur1so6
ar{l rdcTqA Japun suorlrpuoc aqtr .pooJ Te1uauaTddns
o1 pasodxe 1ou asaa6 pTTA uer{l E:oTJ 1n6 TeTqorcTllr
IrT
sluauara 1upl-rodurT lsorr aql 1eq1 s1sa66ns saTcads a6e:o3r
aarql ar{? Jo r{cea uoJJ sacaeJ pue anss11 1ue1d puno:6
-aaoqe Jo uoTlTsoduroc TeluauraTa aq1 uT sacuaraJJTq
.aloqe palou se ,I111enb a6e:ogr u; asEarcap aql
roJ alesuaduoc dBru arqTJ .d:e1a1p u1 sasearcur o1 asuodsar
uT lcer1 TpuTlselulo:1se6 aq1 Jo .druoleue u1 sabueqc
'dlarrrleu:a?Tg 
. 
(eeOf 6u11arreg pue :a6uTpas aas) 6u16e:og
luads auTl Jo lunoure ar{1 6u1sea:cu1 .dq a6e.rog sTr{1
azTTTln dlanllcaJJa 01 aTqe aq .t{eru asaag . (HeOf saTraJJaf
pu€ uauelox ,seqray) sarcads a6e:og pa.r.raJard a:ou
ar{1 Jo acuesqe aq? uT r(1aa1sua1xa pasn s; @
ErTEor@dJ '(;uH 'lurod olur)tsg reau ,deg asnorgd er Jo
ql:tou uDt O9Z) .ran1g TTauuoCcn aq1 1p ,raAaAo11 . (qreggOt
sal:a;grag) qfT@ ra^o par-raJard a:B 
-ffi5ffi puB *6T15 r(eg asnorad eT 1v . (eeer pTaTTeA pue uneqsqcng
!?g6I UTertS pue efarTpA ,urneqsqcng !LL6T ,VL6T poorrueg)
sale:pdqoqrec uT q6Tq pue ua6of,1Tu uT r,roT a6e:ogl o?
pa:eduoc par.raJard aq TTTA a6e:oE go adz(1 sTr{f leq+ dtalTt
sT 1r 'a1e:p.fiqoqrec o1 pa.reduroc ua60r1Tu Jo slunoue a6re1
uT=luoc sanssrl aq1 ,s.rncao salcads a6e:oE Jo r{1^0.16 plde:
aql uar{A po1:ad raununs aq1 6u1:nq .salTToqplaur .d:epuocas
.ro ecTTTs ,atqTJ Jo slunorue q61q 6ulule1uoc spooJ 6ugploae
pue sluaura:rnba' rpuoTlT-r1nu 6u11aeur uaaJq,laq asruo.rduoc
€ 
saaToaur .dlqeqo:d asaa6 r(q a6e:ogl Jo uorlcaTas . (feef
.TaTTPA pue uneqsqcng it86T 6ul1aaeg pue :a6uTpas j0g6T
l1a^a-rd pup spluor{J, JO86T ?uaJO pue 6rnquapl ,su1:6 !LL6T
saTlpq pue 1ua6ng ,uarrg lgL6T uar"to !e L6I JJaTT) asaa6
Jo rnoT^pqaq 6u1paa3r a^TlcaTas .d:aa ar{1 roJ uoTX.upTdxa ue
saptao:d '(T80t rrqrs) saur? lndq6no:q? lroqs .d1aa11e1a:
.dq pazl:alcerpqc ,pooJ Eo burssaco:d pgde: .rTaql qlT/r
9rT
burlcalgar ,a6erog: 
-FTfli6_tr{EJ 
urorJ pa^Trap sacaeJ
uT /rroT aran uoTlpzTTpJaurur ua6o.r11u 1au Jo sale.r aq,I,
.paseaTa.r lou a.re qlno:6
TeTqo-rcTur :oJ sale:tlsqns aptno:d pup sluaXuoc :eTnTTac aq1
uo lcp uec qcrr{A sasealo:d pue saseToJpr(q 1eq1 os ,6ut1so6
aI{1 Jo 1n6 aq1 q6no:q1 a6essed 6u1.rnp ua{orq aq r(eur sse:6
aql Jo sTTac r*ral d1aat1e1a: ,osTV .,{11a11ce TeTqorcrllr
lcT:1sa.r. r(eu qcTr{A a6e.rog: ffi-i@ne urorJ palrrap
sacaeJ uT apTJoTrIc pue urnTpos Jo slunoure :aq6gq aql r{1TA
palpTcosse aq r(eu 11 1nq ,uirrou>1un s1 acua.raJJTp sTr{1 Jo
asnec aqtr' 'ffi uuoJ palT-rap sanrel bulpuodsarroc ueql
ssaT d11ueclggu61s a:aA Effi. uo.rJ palT:ep sacaeJ
uT uoTlpzTTErauru ua6o.r1Tu 1au ar{l Jo sale.r ,FTE| r{1TA
pa:eduoc pmfrTffiE Jo ,{1TTTq11sa61p .raq61q aq1 alldsag
'a1E.r uol1e.r1dsa: q61q p r{1TA pa?pTcosse sen sTrI[ .ffi
peze:6 qcTqn sburlso6 uroJJ sacaeJ uT ua6o:11u aTqnTos
Jo suoTle:luacuoc aq1 pacnpal alpr{ deru uoglpzTTTqouurT
TeTqo.rcTw 
.EffiurcEfi :o F-ra-Fo:-6Fuffi ,-6Ei
uo pazE:6 qc1qa, s6u11so6 uo.rJ sacaeJ uT suoT unruounue
aTqnTos Jo suoTlp.rluacuoc aq1 uT punoJ a:aA sacuaraJJTp
oN 'qryro:6 TeTqo.rcTur ?caJJe o1 s:eadde (satcads a6e:og
?uaraJJTp llro.rJ sacaeJ) a1e:1sqns aq1 Eo .fi111e$ "r{,1
'qlro:6
6u11so6 plde: Jo por.rad 1:oqs aq1 6u1:np a.rnlcn:ls auoq
pup lrrsTToqelaur u1a10:d ut sluaurara asaql Jo acuel.rodrul
aq1 lcage.r unTcTpc pue ua6o:11u Jo slunorue a6:e1 Jo
uoTlualag 
.anssTl 1ue1d ur sanTel o1 pa:eduoc sacapJ uT
paureluoc unTsau6eu pue suoT apT.roTqc ,unlpos Jo slunoue
ar{X uT sasearcur a:en aJar{1 ,1e:aua6 u1 .urnlsselod
pue 'unrclec ,uoq:ec ,ua6o:11u ere asaa6 aq1 dq pauTela.r
LNT
JoJ aTqelTns arP sacaeJ qsa.rJ uT suoTlTpuoc TecTuaqc
-TecTs&{d aq+ ,:alanog . (egeOt sa1:agEeg) suoT unruoururp
tIlTA paul:d araA ,{aql lT s?uaurTpas aq? uT pecnpo:d I1plde:
spn alErlTN 
.lTncTJJTp uoT?celap rTaql 6u1>1eru ,palTruTT
-alerlsqns r(1a:aaas r[lqeqord a.;te sluaurTpas uT sraTJT:lTN
'uoTxpcTJTflTu apnTcuT 1ou saop 1T acuTs alEurTlsarepun ue
dlqeqord s1 arnseaur sTr{,1 
. 
(Heet saT.raJJaf pue {TH ,ssang)
sacaeJ uT ua6o:1;u c1ue6:ou1 Jo uoTlerluacuoc aql uT
asparcuT lau ?upc1g1u61s E .roJ TeTlualod aq1 6u11uasa:da:
'.d1aal1cadsa: 1_r{ vz + 1_,trgo 6 + N- 3gU O* g.z pue ,,-e
vZ + r-&l{Q 6 * ?oc 6ru o'Zt aq pTno/q saler uoTlezrTerauTur
ua6o:1ru pup uorle:1dsa: feTqo.rcTllr J,gI Xe 1eq?
alpcTpuT sacapJ r{sarJ Jo suoTleqncuT r(:o1e:oqe"J . (Neor
saT.ralJaf pue >tTH ,ssang) (pazTtTaeToa) a:aqdsoulp aql 01
eruounup se lsor ueql rar{4er 'sacaeS qsarJ ar{l ur luasa.rd
saqo.rcTur r{q pazllTqoururT ,{telTt spA sluarul:adxa .rTar{1 uT
q g? ralJe (SeOt) sa1:a;Jef pue .dlazeg dq :ogl palunoccpun
ua6o:11u TecaeJ aTqnTos ar{1 Jo uolX:od ?upcrJTubTs V
. 
(eeef lsaos up4 pup luaunuaq)
r111enb :ood go a6e:ogi uo Furpaagr saro^Tqraq ur pa^rasqo
ualJo sT d+TfTql+sa61p a6e:og pup a1e.r a6essed uaa/qlaq
dgqsuolleTar asralur sTr{,1 .q6Tq araA sale: suoTlecaJap
lnq 
,FTEEET6@TEJ uo /vroT .61a.rr11e1ar araA s6u11so6 Eo
sale-r 6u11ca6 'asaa6 :o3: a3:nosal pooJ p se .d111enb aaol
Jo sT 1ue1d sTql lpr{1 1sa66ns ,salcads a6e.roE oA1 -raq1o
aql roJ sanreA 6urpuodsar.roc o4 pa:edruoc FFiF6EbEriffi
Jo anssTl Jo deTfTqTasablp i*roT ar{? pue oTler N:c r{6Tr{
aq1 'a6essed 1n6 6ur.rnp auTl uoTXualar r*roT aq1 ,anss;4
ar{1 Jo lualuoo ue6o:11u no1 .dlaaTlpTar aqtr .alerlsqns
z{111enb :ood e uo qlato:6 TeTqorcTur palTurl1_ua6o:1ru
ItT
pooJ lcaTas d14ua:edde s:a6e:ogi acuTs .uoT1e1a6aa paze:6un
o1 pa.reduoc uoTlelabaa paze:6 u1 palrsodap ate ua6o:11u
Jo slunoue :aq6rq 1eq1 paluauncop aler{ (qleof ) sa1:agrEag
pue 11r6.re3 pue (trct ,VL6T) poon:eg .suoTlTpuoc 6u16p.roJ
1eu11do 6u1u1e1urpur roJ r{:essacau osre sT spuno:6 6urpaa:q
rTaql uo asaa6 aql Jo JnoTlpr{aq 6u16e:ogl TeTuoToc aq;
'asaa6 aql z(q pa:gnba: lsou uaqA
aurTl p 1E uotlela6al paze:6 Jo qilto:bar aql s1:oddns pue
'sluaulpas ar{1 oluT ua6o:1tu Jo rrtoTJ aql Jo acueualureu
ar{1 uT }tuTT Tprluessa ue sluasa.rda: ua6o:11u TpcaeJ
crue6:o Jo uoTlpzrrprauTl^I 
.r{sJplr lTps ar{1 Jo aoupreq
ua6o.r11u TTeraAo aql uT lupcrJru6ls aq TTTA qgro:6 1ue1d
roJ luauaTo sTql Jo ,{?TTTqpTTeAe Jo alpJ aq1 sasearcuT
r{cTr{ic ssaco:d due , (e?goT sar.raJJaf pue 1116:e3) ua6o:1ru
Jo dlTTTqeTreAp ar{l ^dq palTurTT sT qry,ro.r6 1ue1d acuTg
'saqo-rcrru z(q uorlPzrrprauru ua60:11u pue uor?lsodruocap
TecaeJ salouo:d tr111enb r{6Trt go a6e:og .ua1sr(s aq1
uT saxnTJ ua6o:11u Jo sa1e.r pue sr{errar{1pd aq? r{loq lcaJJe
I{cTt{A uorlrrlnu aJolTq.raq Jo TaAaT aq1 puodeq slcaJJa spq
asaa6 aq1 .dq a6e:oE d111enb q6Tq Jo uoT?caTas aqtr
'(eeOf uolq6nepcl{ pue ssang) qlro:6 1ue1d
roJ ua6o:1tu clue6.rouT Jo ,{+TtTqetTele aq1 pue ,qs:eu lfps
aql uo uoTlpzTTe.reuru ua6o.r11u Xau pasparcuT ol alnqTluoc
rleur u.rn1 u1 rIoTr{A ,saceegi asoo6 u1 I11a111ce TpTqoJcrur
Jo sTaAaT r{6Tq eq1 .f,q pa?ETnurrls aq osTp .f,eu saqorcTur fTos
Jo r{1rqor6 aql 
. 
(feef uolqbnegchl pue ssang !ng61- aspuaJag
pue pllpsog) saqo.rcTur ,tq pazllTqouurT aq ,teru ua6o:X1u
Te.rauTut 
,suo111puoc burlTurTT luarJlnu rapun ,Jaaaaog
'q1ror6 1ue1d :ogl a?erlTu Jo acrnos luecTJlu6rs e
aplao:d .{eu sacaeE uT paXe.raua6 a1e:11u pue ,uol1ec1JTrlTu
6VT
. 
sa+Ts asaql uo uals/ts aql
Iuo.rJ ua6orlTu Jo sassoT ?au 6ur1eq:acpxa Ilqlssod ,pacnpa:
sr saqorcrlu r{q ua60.r1Tu Jo uorlpzrre.rauTur (srTsoJ6'euilsre3
'6a) :ood s1 I111enb a6e:ogr a.rar{A .spurA Jo sapTl
rIq paaoua: are sacapJ aq1 aroJaq ,sluaurpas aq1 01 luauraTa
sTr{1 Jo urnlal aql salelTTTCpJ sacaeJ qsaJJ uT saqorcTllr
trq ua6o:1Tu Jo uorlezrTeJauTut 1au pldeJ acuTs ,asaa6 pue
s1ue1d qloq 01 lTJauaq TeTlualod Jo a.re 6u1zez6 Jo slcaJJa
aq1 snr{tr 
.s1ue1d paze:6un JaAo AT 6u11ca1as .d11enurluoc
asaa6 aql uT slfnsa.r spJEAs paze:6 ut uollcnpo:d
^f,:eurl:d punor6-a^oqp lau paspa.rcuT pue ua6orlTu Jo
slunoue r{6Tq Jo acuaualuTpu aq1 , (gegt sqary pue suarddalS
J986T t'roTpC pue dlqrg) a{pluT ua6o:11u azTurrxeu o?
osT
s?unoure Eulpuodsar-roc aql ueq? :e6:e1 sear EESasfeirFQfrE
ffi5 uo pal r{cTr{A s6u11so6 uorJ pa^T.rap sacaeJ
ulorJ pazTTeraulur ua6oJlTu Jo ?unoure aq,tr (S . (C :aldeqC)
paaoural araA sacaeJ aq1 r{cTr1&t uorJ ?nq paze:6 a:an 
Xeql spJp/rs 
Jo sspuroTq puno:6_anoqe aql uT ?unourE aq1 upqf,
:a1ea:6 d11uec1g1u61s sprq pa?pTnurncce ua6orlru 
Jo ?unoup aqfi .sp:eAs 
aq? uo secaeJ uT pallsodap ua6o:11u dq :og
palunocce aq lou pTnoc sp:pr,rs paze:6 Jo sseurolq puno:6
-aloqe aql uT palpTnuncce ua6orlTu Jo lunoure ar{J (V . (V
:a+deqC) spr.,os paze:6un Jo 1eq1 a^oqp ddVN uT sasea:cur
.ra6:e1 ue^a pa^oqs sdep ?z dTaleurlxo:dde Jo sTe^raluT
le suoTspcco aarq? uo pazp.r6 sp:earg . spJpltls paze:6un
Jo leq+ o1 pa:eduoc ddVN Jo sanTpl a^TleTnunc q61q
paaoqs sdep zI Jo sTerrJaluT 1p suorspcco ? :o 
e uo paze:6
E-TrE-ffid Jo sp-reAs (E . (v pue g s:a1deq3) asaa6 aq1
dq pa:1nbar lsour spA ?T uor{tu po1:ad aq1 6u1:np d411enb
TPuorlT.rlnu q61q g:o a6e:o3 Jo uoT4cnpo:d aql uT palTnsar
r{oTr{A ,sp:er\s paze:6un u1 1pq1 upql slunoure :aq61q
lp spJel[s aql Jo lueluoc ua60:lTu aq1 paurplureu 6urze.rg
(Z 
. 
(e :a+deq3) passa:6o:d uoseas aql se peuTTcap
6urze:6 go slcaJJe aql uro.rJ .ralooal o? uoTlelabaa aq1
Jo d?TTTqe aql leq1 1nq ,uospas eql uT r.l.rea 
==FotrE'rftd ETrTS-ffid- Jo spr.As Jo ddvN ur sasparcur 
or par asoog ,nous 
rassar ar{+ Jo s6u11so6 .dq 6ulze:6 30 sra^ar aleJapou
+eq? (r ajre sluaur:adxa asaq? Jo slTnsa: uTeur aqtr
s+TnseJ Jo iller^Ja^o T.g
sna.[s^s gNrzvEg Nr
NV SV .l.uoArguaH _
NOr.lnTongoc
NOrSSnCsrq
T9T
oNv ssgcoud ng,IStrSOCA
TVIIANAg :9 Ug,trdYIIC
IlaaylcaJJa are s6u11so6 raununs Jo pua aql ,{q q6noq11e
's11npe ueql .raTTplus a.re daql acuTs afecs 1e11eds
.raTfpus p uo luapTla aq pual uo11e1a6al aq? uo s6u11so6
Jo slcaJJa ar{g' 
.asaa6 lTnpp upql (paqsTTqndun TTarrqcoU)
6ulpaaE arrTl arour puads pue (qEeof saT:aJJaf pup 1116:e3)
salpr uorleceJap :aq6gr{ aleq 01 pual s6u11so9 
.rnoT^eqaq
6u16ero; rTar{1 uT asaa6 lTnpe uo.rg raJJTp s6ul1sog
'(s6u11so6 p.ue s11npe) asaa6 pTTA Jo saT^T1ce 6ut6p.roJ
aq1 uo.rJ lTnsa.r qcTr{A asoql se aups aqx aq 1ou z{eur
uol1e1a6al aq1 uo s6u11so6 ,{q bulzezb Jo slcaJJa ar{,1 
-T
'sluaurT.radxa
asaql uT s6u11so6 aalldec Jo asn aq1 qlTA palpTcosse
saTlTncTJJTp .:o[eu aa:q1 are a.raq? ,ssa1aq1:alaN
'.f,:oalq.rard Jo slcaJJa aq1 6u1Ipn+s uaqA se6elueape
snoT^qo r(ueur saplao:d uotlelaba^ Jo sprp,}ls uo (sacaeJ
urorJ ua6o:1ru Eo 1ndul pue palourar sseuroTq Jo Xunoue)
6u;ze:6 go TaAaT ar{1 To:luoc o? .f,1111qe aqg, .parapTsuoc
uaaq lou arrer{ qceo:dde sTql Jo suoTXelTurTT ar{1
1nq ' v'Z uoT?cas uT passncsTp a.rar,r saTpnls TpluaurT.radxa
uT s6ur1so6 arrtldec 6u1sn go sa6eluplpe aqfi
s1ueurTJadxe uotleq:n1:ad Jo suoTlelTurTT aqfr Z.g
'6u1ze:6 6ulrrto11og q1i*ror6
1ue1d urplsns o1 ua6o:1Tu Jo sacJnos TpuoTlTppe aplrro:d
z(eu sluaurTpas aql pue secaeJ uT uoTlezTTerauTur ua6o.r11u
TPTqoroTr{ 
.(E .raldeqc) @ Sflso-f6-
eTTTaUTccnd uro:J pa^T.rap sacapJ uro.rJ
z9r
:o F5$6TEEf:iB
t{lT/r auop aq xsnu suorlTpuoc re.rnleu aqx 01 sluauTradxa
asaq? urorJ pauTplqo slTnsar Jo uoTleTode.rlxg . E
..Inor^Er{aq 6uT6eroJ uT sabueqc luecTJTubTs
<>1 peaT 01 .dTa{TTun sT ?T 1nq ,s6ug1so6 arrlldec 6u1n1oau1
sluaurr:adxa 1ec160101strqd Jo slTnsar aq+ lnoqp suracuoc
sasTp: urelqo.rd sTq,l 
. 
(eeet lsaos ueA pup luaruuaq)
sprTq pTTA Jo 1eq1 o4 pa.reduoc  cprl a^TXsabTp aq+
uT uoTlecTJTpollr uT lTnsar z{eu s1q,1, .a6e:o3l Te.rnleu ueql
a:qTJ ssaT pu€ s?uaT.r1nu Jo s?unoue q61q uTpluoc 1pql
spooJ TeTcTJT?:e dq paluauraTddns se,r s6ug1so6 aarldec
Jo laTp aql 1eq1 sT sTr{1 .roJ uospar aqtr .s6u11so6
arrlldec pup pTTA go .d6o1o1s.dqd ant1sa61p aq1 uaai*rlaq
lsTxa z(eur sacuaraJJTp .IaAaAor{ '(V.Z uollcag) palcalap
10u a-ra/{ s6ur1so6 aatldec puP prTA Jo saler 6u1>1cad
uT sacua:aJJrq 
.s6u11so6 pTTA Jo cTlsT.ralce.rpqc 1eq1
ueql r(60101s2(qd antlsa6Tp luaraJJTp p a^eq pue s6ur1so6
pTTA luo.rJ I11ua:a3iJrp a^eqaq r{eur s6u11so6 aa11de3 . z
.asaa6 l1npp 6u1sn pa?cnpuoc uaaq peq
sluaurJadxa aql g:t palca?ap aq 1ou pTnoA uol1e1e6al ar{1
JO SASUOdSa: :prrurrs 1eq1 a^arraq o? uospa.r ou sr aJar{1
'uo11e1a6aa aq1 uo asaa6 ?Tnpp rtq 6u1ze:6 Eo slcaJJa aq?
Jo uoTlenre^a lcaJTp ou apr^o:d s6u11so6 aarldec 6ursn
sluaurT.radxa gro slTnsa.r aq1 q6noq? uala ,a:oEa:aqg, . (g-e1t
'F 1a a>1oo3) JalpT s>tean x1s aFpalgi s6u11so6 aql aurrl
aq1 ^f,q 1:1 Ilalerulxo:dde 01 saspa.rcap orlpr sTr{1 ,.Tr?,
dlaleurlxo:ddB st r{cfeq 6urr.o11og lTnpe :ad s6u11so6 g:o
ragumu aq1 'uoTlTppe uI . (HeOt saT.raJJaf pup >tTH ,ssang)
slTnpe o1 pe.reduoc ua6oJlTu aTqnTos Jo s+unoue .ranoT
uTeluoc osre r(Bru s6ur1so6 Jo sacaeJ arl,tr .azTs lTnpe
e9T
lurod urnr:qTTTnba ue .Ipau ureura: o1 ualsds e Jo dcuapual
aql 01 sJaJa.r r(11ensn ,(aTtTqe+s Jo ldacuoc aqI,
'saTecs 1e:odua1
pue 1e11eds :a6:e1 1e xalduoc aq .deur suoglceraluT
1ec16o1o1q ,,a1durg,r 
.parepTsuoc aq osTe lsnur uotle:6a1u1
Jo sTaAaT .rap:o :aq61q ,suo11e1ndod ro sTEnpT^TpuT
uaaAlaq AoTJ 1ueT.rlnu Jo Ta^aT aql 01 pacnpa.r aq uEc
'6u11cdc ?uaTrlnu se qcns ,sassaco:d sualsr(soca q6noq11v
'(geOf 
.Tp 1a TTTaN,O) lulodpue1s cT?sTToq e uo.rJ
paaarl aq lsnur 1nq ,s1e11qeq pup sarcads Jo suoTlcafToc
a1du1s sp pazrTenldacuoc aq louuec sualsdsocg
slTun clueudp sp suals.dsocg t.. g
'razrTrpal Jo uorleclldde aq1
pup uoTleTToJap 01 slueTd Jo sasuodsar ar{1 r{pnls 01 rapro
uT XdN Eo 6eq E pue auTrlceu 6ulrroru p se pasn sT asoo6
aq1 'qceo:dde Tp?uauTradxa aq1 6u1.d;rldrulsraao .d1sso:6 ;o
{sTr ar{1 1V 
. 
asoo6 Aous aq1 Jo .rnoTneqaq 6u16e:ogr aql Jo
rpnls e 1ou sT 1T :bu'r.zez6 01 uoraexa6aa arll Jo sasuodsar
aql Jo .dpn1s p salToaur 1ca[o.rd sTrI? ?pr{1 paslseqdura
aq lsnur 1I 
. 
suoTle^Jasqo pTaTJ urorJ uoT?e[rroJuT
aTqeTTar uTelqo o+ ?TncrJJTp d:arr st 1T lnq suorlrpuoc
TeJnxeu Japun rncco ,{eru sluaurlpar? Tp?uaurTJadxa
aql uT pasn suoT?Tpuoc aqg, .asaa6 pTTA 6ulsn paJansue
aq 1ou pTnoc passeJppe 6uraq a:an r{cTqn suorlsanb
aq1 asneoaq ,s1o1d TeluauT:adxa u1 s6u11so6 arrlldec
6u1sn palcnpuoc a.reA aloqe pa1:oda: s4uaurrJadxa aqg,
'.d6o1o1q Teluaurradxa ut rualqo:d 1e:aua6 p sT sTr{,1 .a:pc
v9T
. 
(geOt 
.TE Aa TTTaNTO ,9867
sTler,tr pue lsod ,s11a6ugaq) acueq:n1sTp 01 +uaTTrsa-r
pue 6ul6ueqcun .d1aa11e1a: sT 1er{1 rualsr(s e uT lTnsar
s1ceqpaal aa11e6au pue a^rllsod uaaAlaq acueTeq aq? arar{A
suorlenlTs aqT.rcsap 01 u,^ral aa11d1:csap TnJesn p suTeua.r
1T rala/qoq ,r(lrunuuroc ,,urnT.rqT11nba,, 3lo ldacuoc aql
1:oddns 1ou saop r(:oaq1 1ec16o1oca luarrnC .d+TtTqels Jo
uotldac.rad .rno acuanTJuT acueq.rnlsTp Jo saTecs lua.raJJrp
.f,11uanbasuoc : a6ueqc sarqpr.re^ Teluauuo.rTAua Jaqlo pue
alpurTTc se a6ueqc sualsdsoog . (eeOf esnos pup 11auuo3)
,t+TtTqees unTrqTTTnba aao:d o1 pa:1nbar a.re po1:ad
6uo1 p .ralo palcaTToc plpp acurs ,ua1s.ds runTJqTTlnba,, up
se tualsdsoca ar{1 JapTsuoc o  lectlce:dur1 sT 1T ualJo
'Grcr
sueg:g) alEfs utnTrqTTTnba ue Jo acuateedde aq1 aag6 trar{,1
'arqels d1aa11e1al ureual pup a3ueq:n1srp roJ alesuaduroc
01 aTqe aq 01 pua1 luelsuoc Jo fualsls:ad ,1uat11sa.r are
r{crtlA suals.ds 'acueq.rnlsrp 6u1ruo1103 ale1s rauroJ slT 01
su:nla.r uals^ds p qcrqn qlTA elet aql pue ,rt1111qe1s lasdn
o1 pa:1nba.r acueqJnlsTp eq1 Jo azTs ar{A r{1oq 6u1u1ur:a1ap
uT 1ue1:odurr sT aTqpls sT urals.r{s e r{cTr{ru :alo uo16a:
ar{1 Jo apnllldtup aqfi . (srualsds r(a:d_:o1epa:d .aT) luyod
Terluac auros puno.re aleTTTcso pup z(1111qe1s 1ec11c^dc Eo
nr.roJ auos lTqTr{xa .f,6o1oca u1 sassaco.rd uollceJa?uT ,{uen
'(Eesr suel:g) acupq:nlsTp e ra?Je 1T o? urn?al 01 .ro
9ET
-AoTaq 01 aTqeTTpAp a6eroJ Jo .dlTTenb aq+ sasea.rcuT roJ
alqlsuodsa.r a.rp sassacord .raq10 pue s.rolcpJ TpluauuorT^ua
suslue6ro-o.rcTur TTos rr(:oa1q:aq puno:6_no1aq
pue puno:6-aaoqe uaaAlaq suorlseJa1uT , Snr{[ - sa.ro^Tqrar{
puno:6-noTaq asaql 01 aTqpTTe/'p ua6o:1gu Jo lunouE
aq1 6u1sea-rcuT dq s:aqrunu podo.rr{ue uT asee.rcur ar{x roJ
alqlsuodsa: aq z(eu , (Oget oTTTTaI^I pup raqv aas) ilulzezb
6ug,rto11ogr sloo.r luacsauas 6u1.z1uoroc saqoJcrur dq 11os
aq? uT ua6o:1ru clue6:ou1 Jo uoTlesTTTqouruT ar{& .sTpluTue
TTos slTJauaq 1pq1 rauueu E uT suoTlcpraluT TpTqorcTlu
-Tros-1ue1d aq1 Jo apnlrubeu aq1 e6ueqc sa.rolTqrar{
puno:6-aloqe leq1 unoqs aleq (geOf ) sar(eg pue opunurpg
'1pa1:eag 
,aldurexa :og .dllunururoc aql uT srusgueb:o Jar{1o
uo slcaJJa alTle6au :o aarllsod alpr{ deu slcaJJa lcarTpul
'(uollezTtTUaJ .aT) s?caJJa lcarTpuT 01 acuel:odur1
r(:epuocas Jo aq z(eu (anssrl Eo TeAouaJ .aT) .6:oa1q:aq
Jo slcaJJa lcarTp ,1eaa1 r(lrunuuoc aql 1V
'(eaor
dalare:3 ! LL6I '696T :ad:eg) salcads 1ue1d luauodruoc
ar{1 Jo saTlTtTqe a^TlTladuoc aq1 .ralTe ro ,sa1n6edo:d
1ue1d ;o les.radsrp alrlcaJJa ar{1 uT pTp ,s1ue1d 3ro
uoTlcnpo:da: pue q1r,ro:6 (a4e1nru11s :o) acnpar ,salTunuuoc
1ue1d Jo uoTlTsoduroc aq1 ralTe .f,Bur saroATq.raH . (eeOf
uolq6nepc4 ,696I :ad:eg) suorlTpuoc cTlorqe 6u111Baa:d
aq1 pue ,d.ron1q.rar{ 01 s1ue1d Eo salcads snor.rpl
aq1 Jo asuodsa: q1,rto:6 aq1 ,sa:oalqJar{ aql Jo .rnor^prlaq
aq1 uodn puadap TfTA saTlTununuoc Ternleu g:o z(1gs:a^Tp
pue uorllsodruoc aq1 uodn d:oalq:ar{ Jo slcaJJa arIL
acueqrnlsTp puE ,{:orrtq:aq o? sasuodsa: uals.dsocg ?.g
9ET
aues aq1 asn ?ou .deu seJo^TqJaq luaJaJJTp pue aTTqou
r1aa11e1a: a'p saro^Tqraq lsour acuTs .pa^ourar ss.luorq
Jo lunoure aql pue peze:6 s1 r{c?ed e r{cTr{r*r r{lTl'r dcuanba.rE
ard1 apnTcuT saTqeT-rea 1ue1:odtut , acueqtnsTp sTr{1
asTJalcp.rerdc ol :ap.ro uf .paurnsuoc sT r{cTr{r'r 1ue1d aq1
:oJ acueq:nlsTp Jo nr:oJ p sluasa:dar .d:oalq:ag . (sget
alTqil pue llaTcTd J g96T IaTrue.rC) salcads Jo luauacelda:
ro (eeeet sa1:agEap) uoTlezTuoTocar roJ saTXTunl.roddo
aptaord qcTqn saqcXed alea.rc detu pue , (ggOt saTraJJaf pue
dlazeg) uorssaccns p:elal osTe .deu 11 , (Hg6t sar.raJJaf
pue uauelox ,saq:ay jTBOT uuel^l pue uol.req$) uoTlcn:lsap
lelTqel{ oX speeT ualJo acueqrn?s1p q6nor{1Tv . (qrp886T
saT.raJJaf i986T aseqa pue uossardc J9g6T alTqA pup 11a{3Td
Jt86T esnos) s1p+Tqeq r(ueur u1 r11aua6oralaq reuorlela6aa
Jo ac:nos 1ue1:odur ue sT seJolTqJar{ r{q acupqrn1sTq
'uorlTsodap lsnp auroqpurrrl 01 irerrurs a1p.;r e 1p uorlenuoJ
TTos Jo ssaco-rd aq1 01 salnqTJluoc suar{cTT cTqlTTopua
6u1u1e1uoc {cor uo sTTpus r(q 6ulze-r6 ler{1 1:oda:
(net) loue:g pup sauof '>leqceqg ,alduexa :og .sassaco:d
ualsdsoca uo lcedrul 1ue1:odurr ue aler{ ue3 saJolTq.raq
'11eus sT paunsuoc uorlcnpo:d ^f,:eru1:d Eo lunoue Tp1ol aql
aJeqr'r serlrununuoc uT uang . (geef saT:aJJaf pup .dlazeg
i986T :a1uad:eC pup 1s;nb6.rag jeg6T r{?Turg) sassaco:d
urals.f,soca AcaJJe suoTleorJTpour asaql pue rsalllununuoc
.dueur u1 uolllsoduroc sarcads aq1 pue ,sal11unuuoc 1ue1d
Jo uorlcnpord r(:purr:d Aau aq1 .dglporu uec sa.rolTqrag
.sTenpT^TpuT pue uoTlelndod r.f,lrunururoc
aq1 lp r(:on;q.raq 01 sasuodsar uaaA?aq qsln6uTlsTp
01 1u.1-rodtul sT lT JaAaAor{ , (E .raldpqC aas) uorutuocun lou
dlqeqo:d a:e eddl sTr{1 Jo suoTlceJaluf .sa:o^Tq:aq puno:6
L9T
'JraAaanoll'uoTlpzTTTlral se qcns susTuer{cau d.rolesuaduoc
aTqplTns Jo acuasa:d aq1 ur uala ,uoglce:aluT
arolTgraq-1ue1d ar{1 Jo auoclno aq1 auTuualap deru slca;Ea
6u11due:1 Jo d1r:aaas aq1 Apql 1sa66ns saTpnXs asaqtr
Te/\T/\JnS
raTTTl pacnpar 01 anp Ilqeqo.rd 's1ue1d forluoc ar{X Jo
lEI{1 Jo *Ll pue t6 uaaA1aq saTcads arn4sed Ot Jo rlryto:b
aq1 pacnpa: daaqs .dq 6u1pea:1 leq1 s1:oda: (lgOf) puorupg
'Grct saTreqC) s1uaT:lnu a:1nbce 01 s1ue1d aq1 :oJ
atqeTTe^e TTos Jo autnTo^ pacnpa.r e sT araql suoT?enlTs
asaql uI 'rttTnE dolanap louuec sloo.r aql pup pa-ralexc
sT p:eAs aq1 sasec auos uI 
. 
(geOf uesuag) slueurTpas
aI{1 uT suoTlTpuoc cTxoup Jo luaudolalap aql pue arnlcnrls
TTos Jo ssoT ar{1 01 speaT ualJo (saqsreu lTes 
.aT)
tI6Tr{ sT arnlsrolr TTos ararln 6u1pea:1 osTV 
. 
s:aze:6 r(q
uollceduroo 01 enp paspa:cuT sTTos lsTour g:o :ar(e1 acpJ.rns
aqX Jo saTlTsuap tTnq aql 1eq? X:oda: (nOt) 1s:oqloqcg
pue puTit 'an11e6au srlearle XsourTp are s1ue1d 1enp1^TpuT
Jo TerrTrr:ns pue spJpns Jo uoT?cnpo:d aq1 uo s:aze:6
a6:e1 Jo slcagEa 6u11duer1 aq1 '(v.t uoTAcaS lsursTueqcaur
{ceqpaal aal1lsod) 6u1ze:6 6ulrrto11og: uollcnpo:d
uT sasea.rcuT 01 peaT .deu qclqA sursTuer{oau r{:olesuaduoc
Terlualod Jo acua?sTxa eq1 alldsaq 
.uoT?e1a6aa uo
6ul1drue:1 pue 6u1pea:1 Jo sXcaJJa aql are uoTlpzTTT+raJ
pue (6u1ze:6) 6u1dd1Tc r{1TA luelTurocuoc .ssaco:d {cpqpaaJ
aatle6au .ro a^TlTsod raqlTa qlTn paleTcosse aq deu
suaqs.fis 6u1ze:6 u1 acueq.rnlsrp ,:a11:ea pauoTluau sV
'1sTxa oq d1a>11T are .f,11sua1u1 pue
r{cuanba:; uT 6urd:err 6u1ze:6 Jo sTaAaT acuar:adxa leqX
saqcled Jo cTpsou e ,r{llunuruoc euo uTq?Ti,r a6e:o3r go sad^d1
897
(reer
suTqqals) sraze:6 uelleuureu ,a6:e1 Jo uoTleTpe: plde:
p paAoTTe qcTqA ,sasspJ6 go pea:ds aq1 ol paT alpuTTc
aq+ Jo 6uglooc apTA-pT.roA eql auacoEglg_auacog elET
ar{1 uT I1:e11ur1S 
.sa.rolTq:aq a6:eT Jo ssaccns panuTluoc
ar{1 parnoleJ urnl uT uorlela6al Jo adz(1 alTlcnpo:rd ,{1q61q
sTq& 
. 
(taat q?Turs-uono eas osle) :adso:d pTnoc s1ue1d
z[paar*r a:er{l[ suoTlTpuoc alea.rc 01 papual .dlsnoauelTnurrs
puP sqradsolEue .dpaan Jo ssaccns renpr^TpuT ar{1 parno^eJ
snoacelarC aq? 6u1:np spode:1a1 snorolTqraq a6:e1
rIq acueq:nlsTp TpuoTlela6aa pue uoTlepa.rd 1ue1d 1eq1
1sa66ns (eeet) daug3r1,t pup 6rTu .luelrodur r(1aua:1xa
osTe sT acupq.rn?sTp aTecs ,{:euorlnToAa ue uo
.q1r,ro:6 uo
lcaJJa tu:a1-6uo1 ,as:aapp ue uT lTnsar lou saop asuodsa:
p r{ons . (uorlecrununuoc leuos:ad ,uauoo-1 .n) (sugur
09 >) anrTl Uoqs E :oJ a6e:og se aTqeTTeAEun uaql 6u11eru
snr{l 
, 
saAeal aTqpssecce lsollr aql Jo a16ue aql acnpa.r
.611:e.rodua1 uana r{eu asaa6 .6q 6ul1durpJtr .uor1e1a6aa
go sad.64 lsour uo lcaJJa 6u11druer1 pa+TulTT e aler{ dlqeqo:d
'asaa6 sp qcns 'stazeeb rarrplus .saJolTq.raq alern6un
a6:e1 .6q pasnec .dlqeqo:d a:e 6ul1drue:1 Jo slcaJJa a.ra^as
lsou ardl ,ssalaql.ralaN . (AeOf uolq6negcn pue srper6:oag
aas osTp) sleru TEuoTc urroJ 01 slueTd aq+ paTqeua pup
palooJ daql arar{l\ ,puno:6 aql oluT sraTTTl slT pac:oJ pue
Trerr-c-F Eb-@ Jo .ralau.Tp u^o:c-loo.r aql pasEarcur
sa1e1n6un uecrrJg Iq 6u11due:? Aeql panoqs (qgeOf)
rrsrag 'aa11e6eu eq r(eu slcaE;a 6u11due:1 TTp lou
697
aql /tq palcaJJe d16uo:1s a:e sTros ur rncco qcTq/{
sassacord uotlezTTTqouurT-uoTlezTTerauTu ua6o:q1N
'(eeOt saT.raJJaf pue r(1azeg) sluautpas Jo aceJrns
aq? uo ro a:aqdsozTqJ ar{1 uT raq?Ta uoTlexTJ ua6o:1ru pue
'sassacord 1e1qo-roTlu Jo ?Tnsal e se sluaurTpes uT ua6o:11u
clue6:o Jo uoTlpzTTeJauTut 1au ,slueulpas aqX uT qlrrtorE
1ue1d :ogl uaborlru aTqeTTeAe ,sal-rasa: puno:6-AoTaq
apnTcuT ptnoc sac.rnos asaql .1uasa:d arp sacaEJ uarlA dluo
qlato:6 1ue1d :oE aTqeTTpAe aq lsnu ua6o:11u Jo sac:nos
:aq1o 
, (9 :aldeqC) pTTe^ ,{ttnE sT suolldurnsse asaql
Jo -raqlTau acuTs 'pazrreJauru sr sacaeJ ur ua60:11u
aTqpTT€Ae aql TTe leq1 pup ,uo11ez;TTlpfo^ pTuoluue 01 anp
sacapJ urorJ ua6o:1ru gro ssoT ou saurnssp gOT Jo aleurTlsa
sTI{1 ualg 
. 
(e-e aTqp&) sac.rnos TeoapJ uo.rJ fi11ca:1p auoc
a^eq pTnoc uoseas aq1 Jo asJnoc aql .ra^o anssT+ Aau aql
oluT pale:od:ocuT ua6o:11u aq? Jo g0T uer{1 ssaT ,buTzetb
Jo por-rad qsa6uol aq1 01 palca[qns a.raA r{crr{rt ffi
Jo sp.rens uT uala leq1 r"ror{s g :aldeqc uoJJ elep ar{fi
'saroATqrer{ Jo saTlT^T?ce aql
luoIJ slTnsal qcTqA ualsz(soca ue uTqlTitl sluauaTa TpTxuassa
gro 6u11c.dca: luaTcTJJa pue plde: aql 
^q lasJJo aq uec
qry'ro.r6 1ue1d uo suoTlplTlnTT luarJlnN . (13AT. uolq6nepc4
pue ssanu !gL61L aasueupooM lgL6I pTeuoqcn pup lJe.rcdoH
'ueu:g) acuasqe -rTaql uT :ncco r{cTr{r'r sa+eJ ar{l aloqe
uoTxpzTTp.rauTur lueT.r1nu Jo saleJ Jo uoTlpraTacce up uT
lTnsa.I sa.ro^Tq:aq Jo saTlT^Tfce ar{1 sualsr{s TeT:xsarJal
uI '(geef 
.Te F TTTaNro) ssaco:d {ceqpaal aatlrsod e se
gro 1q6noq1 ualJo sr suelsfrsoca uT 6u11cdc luaT.r1nN
sualsr(soca 6u1ze:6 u1 6u11ctrc luaTJ?nN g.9
o9T
p 01 aluTl u:n?a: .aT) ,:nolneqeq 6u16e:og ;o u:a11ed
aslca:d e AoTfoJ 01 aleq 1ou op esaa6 aq; .luaurleaJ1 aql
Jo alrlcadsar:1 ,:e11uts .d1peo.rq e:e ddVN alTleTnurnc Jo
sanTel aq1 ,saposrda 6u1ze:6 Jo s:equnu pue sarlTsua?uT
6u1ze.r6 3lo a6uer apTA e JaAo ,1et{1 1sa66ns p :aldeq3 Jo
slTnsa.r eqtr 
.uospos aql uT r[1:ee pJeAs ar{1 uo pallsodap
aJe sacaeJ luaTcTJJns paptao:d , sp:errls paze:6un Jo leq1
aloqe 'spreAs paze:6 Jo ddVN aq1 uT sasea.rcuT 6u1aalqce
roJ TpcTlTrc lou a:e asaa6 aql 
^q 
saposrda 6urze:6
aq1 Jo 6u1ur1q aq1 1eq1 sa11dru1 XT ,1sa66ns slTnsa.r
aq1 sp 'rleg asnorg)d eT ?e s.rncco d11n11ce TeTqo:cTur TTos
go 6u1ru1:d s1q1 ;1 
. 
(fAOf uolq6nepcl^I pue ssang) sTTos
ut ua6o.r?Tu Jo uoT?pzTTEJauTur 1au e ur slTnsa.r d1a1erur11n
pue 'a.laqAasra a?e.rado o1 u/vrou)t sT (s:ncco uorlezrrTqoururT
ueql raqleJ uoTlEsTTE.rauTu ?au 1pq1 os oTlpr
N:c aql gro 6u1:ano1 
.aT) e.roTJ TeTqo:cTllr TTos ar{1 Jo
:au1.rd e sp sacapJ Jo aTo.r aqJ, 
.saqorcTur dq pazTTTqouurT
sT pazrTp.rauTu sT qcTqA ua6o:1ru ar{1 Jo lsolu Xeql dtalTt
sT XT soTle.r qcns 1V 
. 
(peqsTtqndun uasuaf pup saTraJJaf
i 896T sarraJJaf pue .dTazeg) OZ uEqX :a1ea:6 s1 ,(eg
asno.rird eT 1e sfuauTpas Jo oTlp-r N:C aq; .r{6Tq sT or1e.r
N:C aq1 pue ltroT e.rp slunoue ua6oJlTu arar{A sluaurrpas uT
dolarrap 01 e-rorJ Terqo:crur rros aq1 6u11qeua uT 1ue1:odrur
aq .deu seoaeJ uT ua6o:11u Jo uorlezTTpraurur lau aql Jo
fTnsa.r e se TTos aql oluT ua6o:1tu c1ue6:ou1 Jo aseaTa.r
aq,tr '(g :a1deq3 iN86T saT.raJJaf pup >tTH ,ssang) sacaeJ
asoo6 ur sspuorq TpTqoJcTut TeTluelsqns p Jo acuasa:d aq1
salecrpur ,{eg asnofgd er uo.rJ acuapr^g . (E :a1deq3 ! Lg6r
uoltl6negcl^I pue ssanu ! vg61' Tnpd pup ppeT : ?g6T aspuarag
pup Ellesog) sJncco 6urze:6 a.raqA sacaeJ Jo uoTlTppe
T9T
i{;l
i
I
l
'acrnosa.;r palTurTT p Jo 6u11c,{car luaTcTJJa
al{l uT lTnsar stuslue6.ro luauoduoc aq1 uaaAlaq sassaco:d
{ceqpaeJ pue paaTuTT sT q1r'ro:6 1ec16o1oTq roJ aTqeTTP^e
ua6o:1gu gro lood eq1 'qs:eu XTes paze:6 aq1 a{TT lsnf
'6u1ze:6 6u1no11og e1a.:tcxa uT suoT unTuollnuP Jo a>1e1dn
luaTcTJJa aq? urorJ .ro 'e1:a1ceqoue.f,c .{q uollexTJ luorJ
d11ca:1p .raqlTa sauloc Jaar TPJoc aq1 Jo qlto:6 aq1 -roJ
ua6o:11N 'suals.ds asaql uT sJacnpo.rd r(reu1-rd 1ue1:odtu1
lsoru aql a:e qcTI{A ' (taet 1ua:dg) ae6Te uaa:6 dq pasn aq
rleru suol ulnTuollnue pala.roxg '(saqslE pue sale:qa1:arru1)
sa.roATqJal{ z(q paze:6 d1an11ce aJP sJaar Jo acPJ.rns aq1 uo
ae61e uaa.rb pue eTJalceqoue.6c 6u1x13r-ua6o:11u'uo111ppe
uI '(eeor s1::ng) rualsr(s aql uo.rJ lsoT sT I'I3Tr{A
uoTlarcxa ulnTuolulue 1au Aoqs 01 pual aeTTal{lueoox 6u11ce1
sTe.roS 'TP.roc aql apTslno uo.rJ ulnTuolulue aleTTuTsse
osT€ r(eur pue Te:oc lsoq eq1 
^q 
palaJcxa ulnTuollrue
azTTTln 1eql (sa1e11a6e1grou1p 3o se6e1s aTTlolu-uou)
apTTaqlueoox c11o1quIs ?soq sTeroc lsot{ 'pauracuoo sT
ua6o:11u sp JpJ se slen Jo Jaqulnu e uT palaTqce sT sTI{.1
'(geot :a1uad:ep) ueaco aq1 oA lsoT Xou a.re sluaTJlnu
1pq1 os '1e11uassa sT 6u11c.dc luaT:1nu ?q6Tfi 'I{s.rellt lTes
deg asno:a6 pT aq1 Xp sassacoird >lceqpaa; 01 saTlTJeTTutTs
rtueu Aor{s sua1s.f,s Jaa: Te:oc uT {ceqpaal AuaTrAnN
'suoTlTpuoc 6u1ze:6 go abue: apTA e .;tapun
sprens pTouTureJ6 aq1 Jo qlrqorF aq1 :oE sluar.rlnu aseaTa.r
01 sseuroTq TeTqo:cTlll TTos aq1 Jo d+TtTqe aqt uo luaXxa
a6:e1 e 01 puadap o1 s:eadde asaag Aous JassaT dq 6u1ze:6
Jo slcaJJa eq1 o1 urals.ds aldl Jo acuaTtTsa.r aql 'ddvN
r{6Tr{ alaTqce 01 sp:e/"rs ar{? roJ .rap.ro uT (qc+ed :e1nc11:ed
291
z{111enb r{6Tq Eo dlddns TenuT1uoc p uT s1Tnsar asaa6 aq1
Jo saTlTnTlce 6ut6e:ogr aq1 1pql uo11sa66ns aq1 1:oddns
aloqe pa1:oda: sluaurrJadxa aq1 Jo slTnsajr arLI,
'(anoqe Z pue T suolldo) aTTqou lou are trar{1 uar{r'r
spol:ad 6u1:np ^d111enb s11 pue .f,111TqeTT.E^e a6erog qloq
alndtueu uec r(aq1 aJeqn pue 'r{6Tq sT 
^lTTTqeTTpAE 
pooJ
a.ral{r'r spa:p ur paa:q lsnu asaa6 Aous ,a:ogaJaq,l . (gggf
tTH JO86I ueqerqg) :eeI qcee aATs lsau pue r{uo1oc 6u11sau
aups aq1 01 uJn?at o? pual pue z[:1edo11qd Teleu aTeuaJ
6uo:1s Ieldslp asaag AouS .ressaT '1e:aua6 u1 
. 
(OeOf
ua.reTc!{ pup uar€Tcry aes) pa+TuTt sT sacuelsTp a6.re1 ;aao
alour ol IlTTqe .rTar{1 os raununs aqX 6u1:np aTTqou ,(tq6Tq
1OU aJp asaag 
.sacJnosar pooJ ,'rau ()1 a^our r(11enu11uoc
01 sT prTql aq& 'asn daql a6e:oE aql Jo .f,111enb
aq1 aleTndlueu 01 sT puocas ar{f,, 
. 
TnJTluaTd sT pooJ ajrar{r-r
sea:e uT paa.rq 01 sT ISJTJ aq;, 
.sTq1 qsrlduocce uec ,(aql
1eql sr(Bn Jo Jaqunu p are atar{J, .a6e:og d111enb q6Tq
Jo saTlTluenb a6:e1 a:tnba: asaa6 6urpaa:q crlc.rg
'uor1e1a6aa uo abueqc crlplurTc pup ,spuelarol
deg uospnH aq1 Jo lJTTdn cTlplsosT ,6u16e:oE asoog /,tous
.rassaT Jo slcaJJa uaa/"l1eq uorl3p.ra1uT aql 01 uoTlpTar
uT seaJe Telseoc asaql go luaurdoTalap TeuoTlpla6aa
Jo su:a11ed aql pala:d:a1ut seq (qeAOt) sa1:agr3rag
'slceqpaal aatle6au .ro eAT?Tsod gio sut:al uT pa.rapTsuoc
aq upc saTlTnTloe asaql ,aaoqe pauorluau sV . (qeeOf
sat:agrgag) cTAcJV uprpeue3 u:alspa aq1 uT spupTlan
Telseoc Jo uorlela6arr Jo uoTlTsoduroc pup a:nlcn:ls aql
.f,3gporu asaag Aous JassaT Jo saTlTlTlce 6u16e:og aq,tr
arnlcn.rls ^f,11unuuoc pup asaa6 6u1paa:q cT?c:V g. g
e9T
aql r{loq r,lTJauaq'' upc sJazeJb luaJaJJTp ,{q a6pJoJ Jo
asn aql Jo Uorlp.rpdas reJoduel Aor{ Jo ardurexa up sapT^o-rd
osTp (eeet uop:og) unqu Jo pueTST aql uo (.,1 snmel sog)
aT+1pc r(q 6ul zezb raluTilr 6ulno11ogr ( .t sfife'f a snffi)
Jaap por .roJ aTqeTTpAe sse:6 go sseuoTq Jo slunoue
uT sasea.rcul 
. 
s.laze:6 a1e1n6un ar{1 Jo sa16o1oqd:oru
aq1 pup s:noTleqaq 6utpaa; lua-raJJTp aq1 ^dq paleqTTToeJ
sT (Seet .'gL6I uolq6negcp) spueTsse:6 lla6uajras eql
uo saJo^Tqraq a1e1n6un lsTTetaua6 3ro ruoTssaccns 6u1zpf6,,
aql 'alduexa :og 
.s:azp:6 luanbasqns JoJ uorlela6aa
ar{1 Jo .t{111enb aq1 acuanrJuT aroJa.raql pue 'bu1zez6 urorJ
-ra^ocal 01 sluprd Jo d+TtTqe aq1 lcaJSre .fieru suorlelndod
Tpurrup lua:aJJTp Jo luauralou aqJ, 
.qyro:6 lueTd paurelsns
.roJ sualqo:d salea.ro ac.rnosar aues aq1 6u1sn sarolTq:eq
Jo saTcads lua.raJJTp uaanlaq suorlce.raluT ualJg
'(feOf suTrd pue 1ua.rq) ^f,1ddns pooJ rTar{1 pue suoTlelndod
a.ro^Tqraq aql Jo sluaura^ou pup azTs ar{1 uaa/tllaq
dlqsuolleTal aT{1 uT r(:laurru.dsp ue ualJo sT arar{J,
'uoseas Eutanollogl aq1 rrro:6 o1 s1ue1d
aql Jo tr+TtTqe aql ssaJdap .deur uoseas aq? uT alef 6u1ze:6
panuTluoc 'uoseas 6u1ruo1103l aq1 qll'Io:6 1:oddns r{crr{A
'unop pTpT a.rp salJasa.r a1e:pr[qoq:pc pue luaTrlnu por:ad
sTr{1 6u1:np ralanor{ ,pauru:alap aq o1 1a.d spq p.rpns
6u1zezb aq? Jo ecupualuTeur aql pup slue1d a6erogr ar{?
Jo rlli'ror6 aql :oJ pot:ad sTq? Jo acupclE;u61s lecr6oloce
aq.l 'raururns aq1 uT JarT.rea 6urze:6 alTualuT Jo slcaJJa
aql urorJ .ralooal 01 uorlpla6ea aq1 sAoTle qsn6ng_pTur uT
asaa6 aql Jo a:n1:edap aq.l 's:ead luanbasqns ur a6e:ogr
Jo uoTlcnpo:d panurluoc aqX .roJ ursTuer{cau e aplao:d
osTE 6u1ze:6 3o su:a11ed eqg, ..raururns aq1 6u1:np a6e.rogr
v9I
fTp 01 pualxa .dBur suorlceraluT asaql Jo acuanTJuT aq1
':ana$oq aloqp palou sV 
.sualsr(soca Tp:nleu uT palcadxa
aq 01 are suoTlelndod uaan1aq suoTlcE:aluT aTqpls_Tsenb
.ro aTqels d11ec16o1ocg 
. 
(eeOf uaues4g) aruTl .t{:euotlnTo^a
uT cTlsTTenlnu auocaq I11:essacau ,rLroalq:aq se qcns
'suo11cera1ur clqdo:1 1eq1 aAaTTaq ol uoseal ou sT araql
'ssalaql.raaaN 
. 
(SeOt sTAaT lgL6T rrels aas) asec aql aq
s.6en1e 1ou r(eu sTr{1 pasn ursrTenlnur Jo uoT?TuTJap aql uodn
bulpuadap 'raAerrrog 
..d.ro6a1ec sTr{+ uT papnTcuT I11eru:ou
1ou sT r{:oarq:aq , (gg6T uazuef) uorlcp.raluT ruslueb:o
-o?-lusTue6:o gro ad.d1 uolulroc e aJe sursTTenlnru q6noq11g
'uoTlceJaluT aq1 Jo lTnsa.r p se pacuequa sT suorlelndod
q4oq Jo ssaulTJ ctlaua6 .ro r{1n0.16 ar{1 a-rarlA suollelndod
o/tll uaaA?aq suoTlcpJeluT anToAuT sursTTen?nI^I
sualsds 6u1ze:6 u1 uor?nTolaoC L.g
'sad^(1 ?elTqeq
luaJaJJTp uaanlaq asaa6 aTceuJeg Jo su:a11ed 6u1zez6
uT s?JTrls Teuoseas .rETTulTs pa1:oda: aleq (Seef ) 6:nquapl
pue suT.rd 
.suosees luaJaJJTp le d111enb q6Tr{ Jo pue
aTgeTTp^e .f,11pear rdloq sT qcTqA a6e:og aq1 Jo asn a{eu o?
.rap.ro uT '(te1t suTJd pue lua:O lVg6T .Te 1a uapJag uea)
6u1:ds u1 sa^par 01 TT.J aq? uT eraSTGG sFTE Eo
spaas uo.rJ slJTqs spueTraq?aN ar{1 uT (ffi ffiy) asaa6
Jo ?aTp ar{& 'a6e:ogr sp uoseas aql Jo asrnoc aql :a^o
s1:ed 1ue1d luaJeJJrp azTTTln sarcads aur's ar{1 Jo s:aze:6
alduexa raqloue uils:aze:6 aq1 pup satcads a6e:og
E9T
JO qaA AsnJJTp e se peAaTA aq ue3 sualsdsocg
'sassPr6 aql Jo uoTlecTJTs-ra^Tp
aql .f,q palcT.rlsal aq 01 sleadde sa:oaTqraq uerTpllruEu
a6:e1 Jo uorlnTona aql .roJ Terlualod ar{l asec sTr{1
uI 
. 
(eeOf dau;g:1;, pue 6ugg) d6o1o1q a^Tlcnpo:da: pue
.f,tuou6orsrqd rl:adso16ue ut sa6ueqc xnoqe 1q6no:q a6ueqc
cTlpurTTc TTlun ,sa:onlq:eq a6:e1 Jo uoTlecTJTsra^Tp
aq1 lsuTe6e palelTTTur aJnlcnrls TpuoTlpla6aa,r(:e11:ag,
aql uI 
.aloqp pauorluau se :adso:d pTnoc snrradsoT6ue
.f,paan aJaqn suoT?Tpuoc pa?pa:c pup srtr_radsor6ue
.dpaarrr go ssaccns TenpTATpuT ar{1 paJnoleJ dlsnoauelTnuTs
sa.rolrq.raq a6:e1 .dq acueq.rn?sTp pup uoT?epa:d 1ue1d
'snoacelaJD aq1 6u1.rnq 
..f,:e11:a6 pup snoacplarC aql
uT su:adsol6ue pue sa.rolTqJar{ ale.rqa+ral Jo uoTlce:a?uT
leco:drca: 6uo:1s p uaaq seq araql 1eq? unor{s aaer{
(fgof ) dauggl,t pue 6rTla -uor?nTolaoc asnJJTp 3lo alduexa
up sp pasn ualJo sT srr{1 pup ,.d:o1srq r(:euorlnTola
6uo1 e pa.rpqs alpq saJolTq.raq pue rtr;roJ qy'ro:6 plouFle:6
1eq1 lqnop aT?lTT sT e.raq1 ,alduexa :og . (f .t uollcag)
sTeurTue pup slupTd qloq Jo uorlnTola ar{l uo acuanTJur
puno;o:d e peq seq qcrqn ssaco.rd e s1 d:oa1q:ag
'(qgeOt e.ra::ag) uoT?nTolaoc ,asnJJTprr Jo
salduexa aq 01 pa.raprsuoc are sarcads Eo sdno:6 uaaaalaq
suoTlceJaXuT Jo sad.d1 aseql .f,ltunuuoc ar{1 Jo Talaf
aq1 XV 
. 
(geef sTne:tr pue lsod ,s11a6ugaq) sualsr(soca
uT sdool {ceqpaal anrlrsod bululruexa .f,q paureldxa
aq uec sualsr(s crlsrrenlnu Jo sa.rnleal .duel .suoTlprndod
uaaj'Ilaq suorlercosse oTlsrTpnlnur ur 11nsa: deu
r{cTqn ssaco:d {ceqpaal arr111sod p se go 1q6noq1 aq upc
lpq1 ssaco:d e sT uorlnTolaoc 
.dllunuuoc er{A Jo sTaAaT
997
palsrxa seq 1T poT.rad 1:or{s ^61aa11e1eJ aql uaa16 ,ua1sz(s
sTql uT ,1cpJ uI .uoTlnTolaoc ^d1dru1 r(11:essacau Aou
saop sTql lnq ,ua1sr{s aq1 Jo ,(ATtTqels aql 6utu1u.ra?ap uT
lred luauodtuoc qcea Jo uns aql ueql .ra1ea:6 sr aTor{ru aql
Jo uoTle:6a1u1 aq.l 
.urelsr{s ar{1 Jo sluauodruoc raqlo uodn
pup raqlo qcpa uo acuantJuT punogro:d e peq aleq arolTqJer{
pup lueTd a.raqn rualsds p uT slTnsar sTrI& .uoT1€zTue6:o
Jo sTaAaT snoT.rpl 6uoue 6u11c^f,c luaTr?nu Eo 6u11dnoc
1r{6T1 Jo raqlpr 1nq , (oeet 'gL6I uosTTt{ aas) uorloaTas
,,dno:6 ITEJ1' roJ acuaprna TpaJ ou sT e:aql .raAaAoH
'uoTXpTndod aq1 3:o TaAaT aql ?e 1ou lnq ,1aaa1 r(lrunuruoc
aql le luaprna sr uals.ds r{srplu lTes _ asoog AouS
rassaT ar{1 uT uorlnfolaoc leq1 pan6:e aq pTnoc 1I
'u:o1 6uo1 ar41 rano 1uec13:1u61s
aJou aJe acueq.rnlsTp acupqc Jo sJo?ceJ snoaue.rlxa
aql .reqlar{A Jo ,uorlnloaa uals.,tsoca Jo asJnoo ar{? buldeqs
uT alpuTutop s{cpqpaal Tpuralur raqlaq!\ pauruualap aq 01
suTEuaJ 1T suralsz(s asaql uI 
. 
(rear unpo pue ual1pd jo86T
'9L6T uosTT&) (uralsdsoca ,.61tunuuoc -aT) uoTlpzTue6:o Eo
sTaAaT :aq61q le uoTlnTola .roJ lunocce o1 sdool {cpqpaaJ
aarltsod uaaAlaq uo1111aduoc anTo^ur lpr{1 pasodo:d
uaaq a^eq susrueqcar{ '.rnc30 .61q1ssod .6eu suals.dsoca
aTor{A Jo ,sarcads 6urlceralur Jo sdno:6 uea/tlaq uoTlcaTas
'lenpralput aq1 Jo TaAeT aql uo 1ce 01 paurnsse .611eru:ou
sT uorlcaTas TeJnlpu aTTr{t{ .sTenpT^TpuT pup suorlelndod
uo acupq.rnlsrp Jo slcaJJa ar{? 6uluadurep .dq os op
z(eu sallTunuruoc pup sualsdsoca uTqlTi*r daTtTqe+s :a1ea:6
01 peaT rIcTr{A sassaco.rd . (teef unpg pue ua11e6) aToqA
E se ualsr(s aq1 Jo ,{lTtTqe+s pup .{cua1c1JrJa ar{? alouo:d
r(eru suolXceJalur asaql 
. 
aloqe pa1sa66ns se 
, 
s>lceqpaagl
L9I
.asaag Aous rassaT /(q
bu:.zetb rauulns Jo slcaJJa aq1 01 luaTTTsa.r dlaura:4xa sT
rlcrr{A pue ' (gte1' sueT.ro nsu$) z(cuelsuoc Jo aa:6ap r{6Tq e
slTqTxa qcTrIA ualsr[s 6u1ze:6 e Jo uor?p1ard:a1u1 aq1 .roJ
sTseq e sapT^o:d sassaco:d TaAaT .dllunuruoc pue ualsr{soca
go 6u11dnoc 1q6t1 sTr{,t 
.sTaAaT clqdo:1 lua.raJJTp
01 's1uedtc11:ed alerpaururT aqq puo.daq suoT?cp.ralur
aJolTqJaq-1ue1d Jo acuanTJuT ar{1 pua?xa r{cTqA ,1s1xa
suorlce.raluT Jo elrns V 
. 
(BggOf sa1:a;;ag) suoTlcp.raluT
a.rolTq.raq-1ue1d go suorle6rlsa^uT uT 6u11c.dc luaTrlnu Jo
saTpnls 6u1pn1cu1 Eo dlrssaceu aq? pue 6urze:6 go slcaJJa
aI{1 Jo I111e.rn1d aq+ uo pacpTd aq lsnur slseqdrug .rualsr{s
paze:6 sTrll uT uor1cnpo:d ^6:eru1:d pacuequa 01 peaT
saqoJcTur pue sluauTpas ,asaa6 ,s1ue1d uaaA?aq 6ul1crlc
ua6o.r11u glo sassaco.rd clrueur{p pup alTlcp.raluT aq,l
'asaa6 aql /tq 6u1ze.r6 go 6u1ru11 aq? pue ,Effi-ifTffiE
gro d6oloqdroru aq1 'uorlezrueb:o Jo sralar snorJpl uaanlaq
6u11c.f,c 1uaT.r?nu 3ro 6u11dnoc 1r{6T1 aq1 01 palnqTrllp
aq uec sualstrs asaql Jo acuarTTsa.r ar{tr, .aluTl 6uo1
p JoJ palsTxa-oc dlqeqo:d aaeq cTlcrg aql uT uo11e1a6aa
TPlspOC pue asaa6 rrrous 'Ja,1a1'Io11 .aTqpualun sT pa.rrncco
ser{ uorlnTo^aoc ?pq? i'raT^ aq1 , (s:ear( Oe r(1a1eru1xo:dde)
897
'zEE-Tes '7, 't{c:eEsa[ mTffi Pue ffi
'uoTlpTToJap 01 s1ueTd e:pun1 Jo sasuodsa: TecT6oTolsdqd
pup qlno.rg 
. 
(OeOf ) 
.t.t ,uaza1g, pue .S ,:aqc:q,
'T6z-Lgz 'E 'EETT6 
.a1:1e:d sse.r6-paxTur upcr.raurv r{lf,oN
p uT sn1e1s .ra?en 1ue1d Jo uorleTparu arolTqraq 1e11ua1od
Jo uoTlpnfe^g 
. 
(geet) 
.X.C ,6u111aq pue .S ,:aqc:g
'rLv-6gi 'fi 'm Elm
rassaT aTeual Jo acueu:ogr:ad alTlcnpo:da.r
luar:1nN 
. 
(efOf) 
.O.C ,sauulcen pup
'ZBZ-7LZ 'fr ,m m .asaa9 /qous .rassaT
azTs ue6:o an11sa61p pup 6urpaag 
. 
(ttAil
'asaag Aous
puP sa^JasaJ
'q'c '.(au>1ug
6ulpaa:q u;
'c 'c 'r[au>1u11
.69T-9gT ,E ,@EFEE
mTffi Fup ffi 
-suor1ela:d:a1u1 crlpurrTc pue
'1ea:1a: ;o sale: 3lo suralqo.rd ,sarluac 1es:adslp :1aaqs
-acT apTlua.rnpT ursuocsrl{ aqtr, 
. 
G.rct) 
.,f .f ,sna:pug
'9gz-I9z '6 'K66- Jo AaT^au X1:a1:Eni$ .sTpurrue
TeT.rossoJ Fulzlseqdua naraa; p :salTununuoc TpJnlpu
uT d:oarq.raq puno:6-AoTeg 
. 
(13et) 
.C.q ,uas:apug
'tzn-grv '-89
'@ F FffiT EiEEffi 
.sTTos lsaroJ oluT raJsue.rl
luac:ad pue r{ecep Jo alpls 6u1:nseau :uoTlTsodruocap
f,a?lTT 
. 
(OeOt) 
.n.f ,o1111aq pue .O.1' ,.raqg
'epeuPc
'uo1s6u1y,.d1ts:arrTun sruaang rslsaqg, .q.r{d
@ E 6,fef diF @ 
.(oeor)
qg&rc gun,tvug,tlT
69r
'asaag Aous
',{'X 'rueqe:qg
' 9 o L- e 69' TL' TiFTo-cg-
F FumoT 
. 
tazezb e Jo Teaoura.r aqX 01 asuodsa.r u;
saTlTunuuoc qs:eur lTes ;o do:c 6u1pue1s pue uoTlrsodruoc
aI{1 uT sa6ueqp 
. 
(geef ) 
.q.U ,sa;:aE;af pue .U.q ,r{1azeg
'EoL-E6s '4, ,m Fb-TTT€ Jo fEffiof .r{sreur
lTes paze:6 e uT qry'ro:6 1ue1d :oE ua6o:11u Jo acrnos V
:sacapJ asoog 
. 
(geOf ) 
.f .U ,sa1:eE;af pue .U.q ,z(1azeg
'z6T-06t
'EE 'FE[i6I; 'uo11e.r61u 6u1:ds 6u1:np sse:6 qsrpu lTes e
Jo saleaT uo FulpaaE s6ul1ung /{ous 
. 
(ee6f ) 
.U.O ,dlazeg
'eppueC 'o1uo:og,Eo d11s:a^Tun
e- 01
'u'q'dlazeg
'sTsarlf,, 'cs'r{ 
.asaag Aouslassal-
uo-T1Ea6ilqsJP!'I-?TeSJo SasuotlsE '(teor)
'eluealIsuuad
'6:nqspno:1s "cur ssou pue uosuTr{clnH ,uapnog . (11auung
'T'd pue uazsaT& 
.T.T ,:aTTTr{ -c.d ,u.oro:g .f :s:o11pg)
- 
- 
sTEIFG esrpg ffi Toiles'18 pjrpunl
TElseoE 5ql, :tu+sjffi-ffi . c.rv @ ur 
.ua1s.6s cgqdo:1
paspq-arolrq.raq aqtr . (OeOt) .q.g ,:a111o3 pue .V.,it
'e{TalTd 
,.irf .,{.S ,uea1ce14 ,.g.U ,a?Tt&l ,.o.g ,TTzleg
'uopuoT 'uo1q6no4g
@ Jo rFdFos @F;tr ffi
pue Jappofi 'luauaAol^I
'(erot) 'u'u ':a>1eg
'dd 'proJxo ,11aaa:1ce19 . (drreq
'869-988
'f 'v puE sar:aJJaf 
.T.u
:s.ro1Tpg) stuauffiT^us ffi ur sEEffi fe-i-T6-oTo-cs
uI 
.luauuorT^ua r{sJpurltes aql uT suoT?pllr.IoJsup.Il
ua6o.r11u TerqoJcTI^I 
. 
GtOt) 
.e.q ,llarrtpag pue .V.S '"Trv
OLT
'268-oL8 'm 'IffiEfrFT EFuIv AqJ,
'saTecs ursrue6Jo pup ,dllunururoc ,uals.dsoce Jo 6ulpunoEuo3
:6u1ze.rb g:o slcaJJa aq,L 
. 
(aeet) 
.C.V ,d>1s1ag
.999-T?B 'T7. ,FEbJocg- Jo 'fEffiof .pruezuel
'11a6ua:as aq? uT pueTsse:6 clesoru e uT sassaco:d
.filrunuuoc pue uolqelndo4 
. 
(qgget) 
.f .V ,r{{sTag
'm, 
,IE=TE=frFN
V ;s1ue1d lrJauaq
uesTJauv aq&
rLrontq:aq saoq
'268-0L8
'asuapTAa ar{1 Jo jtlaT/\a.r
'(egAet) 
.C.V,r(>1s1ag
'eluerrlr(suuad'6:nqst::eg,alod>1ce1g'ecTrauv
q1:oN lo supns FG G-saae ffi . (grct) .c..f ,aso:11ag
'gl-sg 'TZ ,7ffi'To-cg FdTTdtr Jo Tpuino] .uo11cnpo:d
deq uo asaa6 Aous :a1ea:6 .{q 6u1ze:6 6u1:ds Jo sl3aJJg
'(geOf ) 
.C ,:a1q1ne9 pue ,.V ,neapelq ,.f ,p:epag
. 
L9-ss '8, 
,3EiiSTGroF .acuaJap
pue uoTlcnpo:da: 01 sacrnosar 6u11eco11g . (ZeOt) ..f .t
'e>11a114 pue 'O'd 'daToC ' .U.N ,o11a1:egqp , -V.d 'zezzeg
'pa+lrurqns
'x6- r- ro mn'or 
.bu1ze:6 o1 asuodsar ur ,ffi6867@
ffie 'sse:6 snoraJTuols crlc.rv up Eo r{qder6oruap
looqs pue JeaT 
. 
(NeOf ) 
.t.U ,sa1:agiglaf pue .U.O ,r{1azeg
'ssa:d u1 '-gL ,tr56'fo-cg Jo furnoT
'I{sJpur lTps paze:6 e Jo .,truouoca ua6o:11u aq1 pue asaag
AouS rassaT 
. 
(AeOf ) 
.t.U ,sa1:a;giaf pue .U.q ,r(1azeg
ILI
'Ls-Es '5T '@ Fue x6o- TToE
' suoTleq.rnuad 01 asuodsa-r uorlEsrTrqouuT-uoTlesTTe.rauTur
aql .roJ suoTlpcTTdurT :uoTlTlsodurocap Jo uorlpTnba.r
luaTrlnu .ro I6.raug 
. 
(leOt) 
..f ,aspuaJag pue .g ,el1esog
'66e-69e
'i 'qTEEStS]Gi pue 7b-Tocg Jo AaT^au Tenuuv 'sualsr{soca
prpunl cTlcrv . (ttot) .a ,uepp1tr1 pue ,.v.fi.q ,pTaTJlTr{u
"u'U 'aAaTU "T'o 'aT11pd "n'g 'u11:no3 "c'T '="Tfg
'92-9 'dd
')t.rol AaN 'uo11e:od:oC 6u1qs11qnd unua1d 
. 
(uosspunurpng
'o :ro11pg) sapnffi utattpou le qcteasal 
- uI ',{11cedec 6urzI::eo a.rolTqrar{ pue uoTlcunJ ,e:n1cn:1s
rTaql :suralsr(soca cTlc:V 
. 
(geOf ) 
.C.t ,ssTTg
. 
?9T- LNT
'Tg 'trFTo-Tg 
- 
. 
seplm pTuoffi aTilrnl uaa:6 ar{A Jo
roT^eqaq 6urze:6 pue TeuoTlT.rlnN 
. 
(OeOf ) 
-V.X ,1qepu:o[g
'tCt-gT? 'dd 
.)tJotr AaN ,6e1:an-:a6u1:dg
'prpun;' 
- 
ue)tsep ue Jo fFffi uo-TTffi
FE uoTTeTS-ban ur 
. 
spTouTure:6 e:pun1 uT s.raTTTl
pue sXoo: Jo saleJ uotle:rdsa.r pue ,:arrou:n1 ,q1aro.r9
' (ef et) 'U'C '.raaeqg pue , .p.X ,uos:a1a6 , .q.M ,s6u1TTTg
-TB? '-E?'sltilEE TEcffi 
.s1ue1d aurdle 
""" 
"r;::: gro d6oloca aqtr . (egOt) .V.g ,r(auoo14 pue .q.1r1 ,sEuTTTTg
'o9eT-TEcT 'E ,@ .uoTlcnpo:d ztrerul:d
pup suor?eTndod uo1>1ue1do1r(qd uo sXoaJJg :r(:oa1q:raq
crlauuTT '(geor) 
.u.s ,:a1uad:ec pue .!{.v ,1s1nb6:rag
ZLI
.TrT-g9T '5 ,Fffiab@fr
ffieu 3o TpuJno1' 'bulzetb 01 sasse:6 go acup?sTsa:
6ullcagrge s:o1ceJ rrlau oi'rtr 
. 
(gggf ) 
.V.,f ,uosuerg
'epeueC 'eanellg
'9V 
.og :ade4 TeuoTsecco ,aclnrag aJTTpTTt{ ueTpeueC
'cTl3.rv UBTpeUeC uJalseg eql Jo asaa9 Aous .rassaT
aqf, 'Zg5T 
.9..{ ,qcoo3 pue , .f .g.g ,t{lTrus , .I{ ,p.,tog
'gzI-ezr 'A '@€ F Teruno,c up-TF@ .ffiEfre
pue ffi :Eacnlsa,jt aqTr; .AI .sasse:6 u:ar{Uou uo salou
crllrouoxel pue sJaqurnu aurosoluo.rqC 
. 
(tgOf ) 
.n'X ,uapnog
.9or-t6 ,F ,Iil;6:Fm
'asaag luaJg .dq salts Eulpaagr bur:ds Jo uoT?caTas
aI{1 uT d+TfTqTesa6lp Jo aTor ar{f,, 
. 
(lgOf ) 
.,f ,uflatapnog
'Etn-ovn 'F 'EE>tTo- 
.uorlcaJuT lcasuT o1 asuodsar uT
slnu go do:p I1:eg 
- 
(erct) 
.t.n ,>1:og pue .H.CI ,:aqcnog
'Lge-eLE 'dd 
.Xrotr i*raN ,suog
pue ^f,a116 uqof 
. 
(q+TnrS 
.q.,1 pup :aToog .tt.C :s:o11pg)
suorleflClo6 Tpruruel^I dffiT- 3o F5lEEEft' uf 
. 
sclrueuz(p
uorlelndod leuureu a6:e1 o1 pa{uTT a.rp sassaco:d rualsr(soca
AoII '(teOf ) 'f,'S'T 'ntr1 pue 
.n.f ,oTTTTan ,.g.q ,uT>tlog
'ggv-rgv 'E 'S'm 
.eTfsassp ET=d6fd6Effi
'1casu1 sno6eqdoilqd e 01 asuodsa: ur ffi.frd mfrffi IaaT
;o d6a1er1s alTlcnpo:da: paTJTpo!{ 
. 
(arct) 
.r ,:aqcsog
CLT
.e6g-98€ 'E ,X66-fEm .1up.rg cTlueTlv
pue asaa6 epeuep dq sluanlTlsuoc 1ue1d Jo ^.(lTTTqrAsa6TCI
'(geet) 
.f ,eTaTTpA pup .f ,uos111r1 ,.U ,uneqsqcng
- 
6P'-ete 
,6 , (u11:ag)
ET56=f6Gd 
.r{s:eu Telseoc e uT aseab ppeuec .(q 6ulpaagr uT
sluanlTlsuoc 1ue1d Jaq?o pue spunoduroc cllouaqd Jo aTor
aq,l '(tgef ) 
.,f ,u1eng pue '.I ,e1a11en ,.U ,uneqsqcng
'tLgT-vggI '6 'X661- 'etseTv uT iqoTfTA
JeaTlTaJ pue aJeq aor{sAous Jo suoTlceJalul . (996T)
'd '1edo:ny ,.& 'uasnelp "q'g 'pue1a1g ,.d.f ,1uer(:g
'ggt.-Lge 'di 'mi 'd:on1q:aq ale.rqax.ral
oX uor?eTa.r uT s1ue1d Tparoq Jo acupTpq luaTrlnu/uoq:e3
'(eeOf ) 'U'q 'uta1y pue 'III 'S'J 'u1deq3 "d.f 'luBd:g
. 
LzeT-6IEl 
,83' ,@-cf, .uorssaccns uTeTdpoolgr
pue acueTpq ?uar:1nu/uoqJec lupTd :suorlceJaluT
areg aor{sAous - i"roTTTI{ JeeTlTag 
. 
(teAt) 
.a.1' ,luer(:g
'Lge-zLE 'Z ,@Effi
Teffiof G@]=EE 
.a:n1sed 3o qyro:6a.r
Eo 
.d11sualuT Jo slcaJJg 
. 
(ggof)
Tam'lTrt3T=fr F
uo uoTleTToJap
'^l'U'ueq6no:g
. 
LZV-qZV 
.dd .eJ.raqupC ,acualcg glo
.duapecg upTfp:lsnv 
. 
(srue1111g 
.g.O pue qcuI"1 .t{.d ,ssop
'f 'd :srolTps) 6G;talruo5 pueT$ueu FuoTIEnAilA puo5etr
ffi Jo @ 
- 
:apun acrnoE :spum@ ur
'saT1T.ro1:d uollecoTTp pue suoTlp.raprsuoc 1ec16o1oqd.roru
:uoTleTToJap o1 asuodsar lueTd 
. 
(gSef) 
.q.g ,a1s1:g
VLT
'g9E-9?s
F5Tffi 
.saTlTunruuoc Te6Te Jaa:
slT pue ,{:oaTqraq 6uguol11rypd . (geOf )
. 
r{s.rer{
@&
'E
Teroc
'c'u
'sqctPrbouol^I
uo slcaJJa
'.ra1uad:e3
'989-699
'fr, 
,m F+=fd€ lo mHnror .r{sreu .lTEs crlcte_qns
P Jo uol1e1a6a^ aT{? uo asaa6 Aous Jassar fiq 6u1ze:6 30
slcaJJa aqtr '(qfeOt) .t.U ,sar:a;;af pue .I{.S ,1116.re3
.Beg-Lsg '8, ,m FAT=tf€ F frnoT .qsreur
lTps crlcre-qns p uT uorlcnpo:d .d.reurr:d ;o uoTlp?TuTf
luaT:1nN '(etgOt) 
.t.U ,sa1:a3lgaf pue .n.S ,1116:e3
'pppupC 'o1uo:o;, ;o r{11s:alTun ,stsaq;, .cS.n
Tf -mi-u- ue F 6-TI"+E%- dF 6' a,s€,a5 nofis
ry:rg- F ffi aqr, 
. 
(reor) .n.s ,1116:e3
. 
nz-vT
'68 '(ut1.rag) pil@o .sesse.lbqcunq 6ilfoi6E pT:eTuras
o/q1 uT uorlecoTTe acrnosal pup .dltcedec cllaqluztsolor{d
:.d:oa;q:atl qlTA 6u1do3 . (teef ) .S.U ,ca:nzg pue .S.U
'>leatog "V.C 
,Uosuqog ,.H.1' ,sp.reqcrg ,.n.n ,11anp1e3
'gEL-szL '6 'FEfr5@ ffim Jo TffiEr .asaao roous
rassaT dq sauozTr{: qsn:Tnq Jo uorXpzrTTln Jo .dcualc1g3rg
'(erct) 'q'a '66e:g pue ,.f.u ,uospng ,.V.g ,uo1:ng
'ggT-TET 'TL 'FbqoTg 5- .sTp:oc ;o .f,1a1:ea e .dq
+?HNgT Jo uoTleTTulTssp pue a:1e1d61 . (eeor) .g.u 's1::ng
-9?T 'Ti, 
, (u11:eg) eT@
uo lcaJJa slT pue s1ue1d
uT I111gqe1.ren . (eaer) .r
'e9T
'asaa6 epeueC ^dq 6ulpaag
aur.rn?sa Eo .d:1sluraqc aq1
'e1a11e4 pue 
.U ,uneqsqcng
ELT
, *;ffi=m;:lr:;: sdTr{suoTleTal I6:aug . (OfOf ) .g.V ,r,raq3 pue .r{.U ,n",{C
oTI{o
'oLE-6sE 'TL 'affi F '[EffiELe
aqf, 
.sc11a6:eua 01 alTlpu.ra?Te up :sualsdsoca
Jo srole1n6a.r sp sJaunsuoC - (Vrct) .n.U ,naqC
'6EZ-62? 
.dd
'>[:otr r'raN ,nog pue :ad:eg . (asBC .f .1' pup puoureTq .f
:s:olrpg) ffi EFlffi'5 ur .ecualsrxaoc pue r:o1s;q
'dlTTTqer:el acueqc :saTJoar{1 r{lgunuuoc unT.rqTTTnba
-uou :r'raTrrJarro 
. 
(gAef ) 
.f .f ,ase3 pue .T.d ,uossaq3
'o?T-LcT '6T ;6T un-T-friffi
Fo@ 'E*-Ei F@E mF=E .uoTlpror.ralap
p.Ielus uT JolcpJ e se 6u1pea:9, . GtAt) .g.V ,sa1:eq3
. 
spr ou T ure:6 e:pu"- r" : :; ::r' rT"r' rj:: 
t: 
:"1"m pue 'uo11d.rosqp sn.roqdsoqd ,q1aro.r6  oo.r uodn uoTlETToJap
Jo lcaJJg 
. 
(At*) 
.n />1ce1g pue .S.J ,u1deq3
'm 'E'cTF5{STSI Fue ffi
pTTA Jo uoTlTrlnu TerauTu aqJ,
'09z-t.ez
F r"la?CE Fn@ 
.s1ue1d
' (oee r) 'rf r 
. 
S.d 
,u1deq3
.99I_Z7I .dd ,p:o3xg ,11aaqce1g . (den .r{.U
::o11pg) uoTlTpg puz ,@ @fffi uf .srualsz{s
a.roATqreq-1ueTd . (f eOt) .f ,uolrre-1 pup .9 ,rta1q6ne3
.6-?, t6 r@
'1uo1qu(s cTlsTTenlnu o1 ua6oqled 1ua1eT uo-rJ :saleaT
pup suals uT salztqdopua 1e6ung . (eAef) .C ,11o::e3
9LI
'uoTlp.reda:d u1 'x6o-To-E FETW go @ior . (@=ae5
Eii55ffiE5 EE[i5) asaag Aous .rassar ur rplTlrns pue
.dllpuncag luapuadap-d11suap Jo acuapTAa aTqTssod . (Heet)
'J 'a1oo3 pue ',it'U '11arvr>1cog ,.g.q ,IueT ,.g.g ,qcoo3
'66s-E68 'm, ,ffiT5Ei
'asuaJap arolrqJaqTlue 1ue1d pue ,{1l1TqeTTpAe acJnosau
'(Seet) 'fff 's'.{ 'uldeqp pue 'd'f '1uer(:g "q'd '{a1o3
.9eg-TEE 'TL , (u11:ag) eTFTo- .asuaJap
a.rolTq:ar{Tlue Jo ad.d1 pup ?unoure aql uo auT aJTT JeaT
pue alpr qlro:6 1ue1d gro slcaJJg . (eeef ) .g.a ,da1o3
'9T-OT '6
'@m '16un; pue s1up1d uaarralaq ursTTenlnur alTsueJap
V :sasse:6 g:o sal.6qdopua 1e6ung 
. 
(eeet) 
.X ,r(eTC
. 
Z9e -89e 'T[, 
, (u11.rag) efffito5ao .EE.@ri?iF
echlsag pue aEIibT6? ffiTFT gro .,t6o1oTq buTTpaas pue paas
aql uo sal.dqdopua le6ungr Jo sloaJJg . (feOt) .X ,defC
-ffi 
EE-tTEoro-ffi 
.s?"'drrr#tffi
ruatsffis.@ .sat-t6ilffirm't.
'(teef ) 
.s:o11pg ,.1, ,llerrrssoU pue .g.,{ ,>[:eTC
. 
vz-eT 
,gg , (ur1oq>1co1s) qi€TTEd
F5J@.tr 
. 
(llerrassog 
.& pup >1:eTc .g.d :s:o11pg)
s1ceffi Tuarua@ prre FaTffi @sG'iocg ffioo-rd
TFaTtIS Fa66-4-11.1 TEffisa-r:aa ur .acuacTT crlaod
r{1TA paAaTA a1c^dc ua6o:1tu aq5 . (f eOf ) .g..{ ,>[:eTC
. 
ge e T- z?,eT ,EE ,@-cg .a1:1e.rd sse:61.roqs
uT g1-ue6o:1Tu Jo 6u11cdc Teu.ralul . (trct) .g..{ ,:t:eTc
LLI
.oTE-e 5? 'a ,)tnE m .asaa9
nous rassaT Jo suoTleTndod 6urpaa:q uear'rlaq AoTJ auag
'Gtet) 'a'r '11arra:6 pue "q'J 'sauulcen ,.d ,a>1oop
.r{llpuncagr
a^TlcaTas aqfi 
.III
asoog Aous lassaT
'V'f 'q11urg pue , .,{ .U
' L LT-seT' dE,,ii6'FE-rs-
1aN :urslqd:oM1od a6erunld Jo anTel
' (suecsaln:aec suaSsaTnffi Jasrrv)
ar{1 Jo sarpnls fuo1s1q aJTT 
. 
(SeOr)
'11an>1cog 
,.s.O ,r{e1pu;.{ ,.J ,a1oo3
'(ssa:d u1) 
.o6ec1q9 ,sse:4 o6ec1q3 Eo I11s.ra^Tun
' (:1co:g-uo11nTc .,I ::o11pg) ssa55ns' 6aliffiFoTffi ur
'asaag ltrous JassaT 
. 
(geOt) 
.TTa,rutcoU .J.U pue .J ,a1oo3
'oog-682 '-zz
'6FF7'a- 'EcffiFr5 ffi asoo6 aql ur ruslgd:oru.d1od
Jo scrlaue6 aqg, 
. 
(gg6f ) 
.g..{ ,qcoo3 pue .,rI ,a1oo3
.epeuPc
'uo1s6ury 
,r(11s:aaTun sruaanQ ,d6o1org Jo luau1:edaq
'1:o€ TE5ffi E = 1ca-E.: asoog lrious ffi €,snoE er
aq.tr '(Zeet) 'f 'U 'ulnbag pue ,.V ,e:npeg ,.J.U ,.6a1eag
"S'D 'dpTpuT.il 
,.C.f ,sarneg ,.d.X ,ueqe:qg ,.C.J ,a1oo3
'100T-866 ,n, ,6ffi:iud5 @
EtroFiif--ffi aql Jo @ 
.acuarradxa snoyaa:d
Jo aTo.r aqf,' :asaag Aous JassaT uT uoTlcaTas r(1t1eco1
pue lelTqeH 
. 
(OeOt) 
.A.X ,ueqe:qg pue .,jI ,aqoo3
'oe-tz 'fr, '=AEEg aqg,'TTTH s1q61uy ' (teet) '.f 'a:1oo3
' z Lz- Lez' Tz, =-q-afe-fb@ EET66'Fts . sluaur?Ea.rl
6u1dd11c snorreA o1 sse:6 lpaql,l palsa.rc Jo sasuodsag
'(egor) 'g'.{ 'ra6urssTX pue ,.v.T ,1:eppolg ,.u,c ,>tooc
8LI
paas n.,. u.**il :: ;-"ffi.T;:"T-T"-"";, sse sprpns sse:6ad: Teruua.rad Jo dcualclgJa asn-ra?en pue
asn raleu '(eeef) 'V'U '1ue14 pue 
.A1.,tr '::eg ,.U ,xoC
.OTT-9OT ,89
'(ut1:ag) AffotocaO '^f,:on1q:aq 01 uolleldepe Jo lTurTT
e - Jalen pup ua6o:1tu 'uo11e1ToJap luanba:; 01 (etpueT-rl
E5ruartf,) ploulure:6 uecrrJv ue Jo sasuodsau 
. 
(qgeot)
'T'T 'ace11e6 pue "f'S 'uo1q6negc4 '.9.r{ ,:nouaq6no3
.98-o8,E5,(ut1:ag)PT@.1uauuojtTAuapa1TuITT-.Ia1PIv1
pue luaT:lnu 'a:oa1q:aq e 01 (E-Tnpu efua::eqdm)
sse.r6-11e? uecT.rJV uB Jo suoTle?depV 
. 
(egeef )
'T'T 'ace11eiv1 pue "1' 'S 'uolq6nepcp '.9.n ,lnouaq6no3
'ggg-zEg 'Tt 'GFreg;
FSfiC' Fnffi]m aq1 F sTetuv 'sassaco:d 6u11cp:axuT
pup 'suo11e1dexa 'suo11e1depV :sarolTq:aq a6:et trq
6utzB:6 01 sasuodsa: proururpJg 
. 
(SaOf ) 
.g.n ,:nouaq6no3
ET56-i6-d 'z(1rs:anrp sarcads luerd pup :":-"-r' j:::l::l
pup sseuoTq XueTd puno:baloqe TeuospaserluT uo s6op
a1.r1e:d pauel-)tceTq Jo lcaJJg 
.I .a1.r1e.rd sse:6-pax1u
upcTraurq r{1f,oN p uT suoTlcp.raluT a:olTqraq-1uefd . (eeOf )
'I'n ':az(q pue '.9.f 'sTTTg ,.X.f ,6u111aq ,.T.q ,>lcoddop
'vzB-6BL'm,ImT
'acualsTs:ad :o .d1111qe1s 1ec16o1oca a6pn[
acuapTla uO '(eAOf ) 
.esnog 'd.tt pup .H.f
uPcTJaurv
ox papaau
'11auuo3
aq,l
6LT
'TL-99'E
'(ur1:ag) pf@ .bu1ze:6 6oq a1:1p.rd Jo sar:o?srr{
asralTp qlTA suoTleTndod sse:g Xpaq6 uralsal{ Jo sasuoda:
uoTlpTToJaq 
. 
(eeef ) 
.t.g ,:a1u1e6 pue .X.1' ,6u111eg
'sesue>1rg ,uo111::o14
'suollecTTqnd qcteasag {colsalTT TpuoTlpu:a?ul tcoruTtl
'se:orTq-.raE F €r+T5E-ffi brail rc "|rx-r_Ee5 aaTiFffi -@ 
f'pog' 
. 
(e eer) 
.r.a ,1saog ueA pue , -il.n ,luautuaq
'dd OeZ 'urTrag
'6e1:a11-.ra6u1.rdg 
.sualsr$ TeJnleN uT
'(geOt) 'C'C 's1ae:9, pue ,.t{ ,lsod
{ceqpaad a^T?rsod
' 'T'q 'sl1a6ugaq
'962-t6z 'n 'Fffi5'6EER
crlcrg pue slqbno:p aT.rTp.rd
aJTTpm F Teurnof 's6u1:ds
6urlsau Tenuuv 
. 
(eeaf)
:asaag AouS uT .d11a11cnpo:d
',!I 'a:1oo3 pue 
.D.f ,salaeq
. 
o6Z-L7Z 
.dd .raluag ,1uaua6eue14
a6ueg .dlarcog 
. 
(eaqasog 
.u .pg) i(6-oTof5m lueTd
Fu€TaEueE ur 
.suorlecTTduT luaua6eueur pu' .f,6o1oqd:our
leluaurdoTaleq 
. 
(trct) 
.H.A ,:apr{g pue .g.g ,Tr{eq
'dd Ez 'm'rroN '-mE @m 5TnTTnt-r=6E
Jo Tffi1=ffi salels pATTm 
.ssB:6 Jo uoTlpTToJap uorJ
6ut11nse: abeddo4s qlivlor6 loog 
. 
(ESet) 
.f ..{ ,:ep1:3
'89T-Z9T 'Z ,E6FFIE pue I5TTo-cs
E s@ asaro^rq-reH luero^auag '(reer) 
.r.n ,r{a1rre:3
'proJxg 
, 
suorlecTTqnd
cTJTluaTcs TTaA{cpTs 
- 
suoTEsffir FEIre@
--- 3o sJJtuEuIB aT[J 
.@_11 . (e e6r) .f .n ,r{alrrre:3
087
. 
L?T-I11" 
.dd .lqca.rp.roq ,r(ueduoJ 6u1qs11qn6
{un1' '^{ 'rO '(uopleug 
.M.U pue ,:a>1>1eg .1' ,Tapug uen .f
:srolTps) 5ffio"ffi ETleffrdoa pup F6}€-c'a-dF @ETssErS
trT a@1s1q ur 
.,{1ddns pooJ slT Jo rauoslrd e
se arolTq:aq ar{f,, 
. 
(eeOt) 
.,f .I{.H ,sur:6 pue .H.U ,1ua:g
'P/v\e1lo
'epeue3 a:n11ncr.r6g 'qcue;g qc:pasa5 ,92 qde:6ouol^I
'6-T=ffi Jo sasserg 
. 
(oeet) 
.r ,111agc1 pue .g.ft ,a.ro6
'E9T-T?T'dd'sllasnqcesser{,pue1:apung
'.cul saleroossv .raneuTs 
. 
(ueq>1n:eg 
-f pue oz:Tq
'u :srolTps) f6-o'tocs G-FeTffiE ?ueTd 6 55aTa56@ ur
'i*raTrr'Jarro cTJluacoT.f,qd V :.d:oa1q:ag 
- 
(leOt) 
-U 'ozrIe
'9v2,-6ez 'Z 'FFTo-cg- f@s Jo Teurnof 
-s:o1e1n6e:
qlro:6 TeTlualod :o; pATTEs uosTq uecT:aurg q1:oN
Jo uoTleururexg 
. 
(f eef ) 
.f .n ,:adq pue ,.V.C ,e111uog
"U'q 'UaqTTH "H'n 'r{a1su1e6 '.A.C 'ssog 
,.y.f ,6u111ag
'IE-92
'fr ' (u11.rag) eib-To- 
.E-TJ-J--TF enoTalnog' g:o qlro:6a:
uo e.rrTTes uosTq Jo slcaJJa TeTlue?od Jo suorleuTurexg
:suoTlceralur a.ronrq:aq-1ueTd 
. 
(OeOf ) 
.A.A
'uu11r1 pue "c '66a:g-:a1co:d ,.I.n ,:arIg ,.x.f ,6u111aq
.VEI-LZT 'fr , (u11:ag) ETF-oto-
'6utze:6 paleTnuTs 6uratollog F!1158F ffiatnog
Jo r{1rrror6a: pue ,uo11e:1dsa: loor ,s1saq1u.dso1oqd
laN '(arct) ',f'O 'uu1tr4 pue '.I.n ':adq ,.X.f ,6u111aq
T8T
pue PTralcPg
'uopuoT 'ssa:6 cTurapecv Te:i5ffi
'(erct) 'g',1, 'u:nq>1ceTg pue 
.,1 ,laqcuag
'sPxatr,'u11sng
'ssa.r6 uTlsng go z(11s:alTun 
. 
(uaaeg 
.H.d pue 1:aqTTg
'g'T 's:o1rpg) (uoTfTpg paslnag) Ft- FuE slupTd
F 6f AfrFaaoc u1sa:olTlrard lcasuT rTar{1 pup s1ue1d
uaaalaq uorlnTonaoc TpcruraqcoTg 
. 
(oeor) 
.a ,duaad
'gL-T 'TZ 'E6EEE Te-5Fp?otr 
.spue1a6up.r Jo uorssaccns
?ueTd uo 6u1ze:6 Eo aouanTJul 
. 
(OgOf) 
.t ,uos111g
'er-r 'Z '@seu TE}nTTncT-:FE
Jo TeurnoT p@az r'raN 
. 
salcads a:n1sed 91 Jo q1nor6 aq1
uo 6utpea:1 daaqs Jo slcaJJe auos 
. 
(VgAt) 
.g.O ,puotupg
'962-992 
.dd .puoqeTTo
'ueu:og 'ssar6 Eluor{eT{O ;o d11s:anTun 
. 
(:aqn:g 
.N.f pue
'T:dI '1' 'U 'sa1sg 'U.1' :s.rolTpg) spue@E pue ffi
uI 'spueTsse:6 u1 saJolTqrar{ Jo aToJ aq;, 
. 
kest) 
.m.q
'1:aq11H pue "c'o 'ueualop '.x.f ,6u111ag ,-I.r{ ,:ar(q
'L1sv-ges? '-LL 'TE-ld 
.@ Jo IFpcv
F6;1eN- ar{T Jo s6- 
.q1lao:6 1ue1d saloruo:d
JolcpJ q1aao:6 1eu:ap1da uerTeunuel^I 
. 
(ogot) 
.r.n ,.rar(o
'9ZL-6I L
'-zI 'X6_dFts FslTTfr Jo Ieurnof 
. 
(.r sreur eail sule:6
uroc Jo uoTXcnpo:d sseuorq uo (.t snac-ruaoqal sn;rTttr)
sprrcptcelq pa6u1A-pal Jo slcaJJa arltr, 
- 
Grct) 
.f .n ,:ar(O
z8r
'z60z
-EBoz '6E- '@ Jo Tprr:nqr EEFEIIF 
- 
(ffi,Tn:raec . 
c ffiE) asae6 Aous JassaT Jo unTpTccoc Teua.r
e Jo alcr{c aJTT aq1 Jo saTpnls Teluaurrradxg . (ZAet)
'f 'CI 'aTuuteg pue ' 
.f .U ,u:oq1rrre3 ,.V.V ,regpe[e9
'a6pr:que3'ssa:6 d11s:aatu61 a6p1:qupc .secu5fr-dsuoc
F5T6o-pTq ffi p-G sur.rtGo-.rm' F ftTF EiiE . (reor)
's.rolTpg 
.U.d ,aue:3 pue .9.M ,:auo1eq3 ,.n.g ursTT.rd
'g'U'.{'6:nq.re4,6:nq:ep gio .61rs:alrug rslsaqg, .cS.n
'*E Fm uo saTpnls E5Tb6-cs . (eeor) .q.r ,.f,a:g
-oLvT_ggb1 'm ,bffi]fti .sacuplsqns 1ue1d
.d:epuocas Jo aJlarp uasrp.r eqfi . (ASO1) .S.S ,1a>1uae:g
'zz'n ,1FT5b--Tg
aql uT buglsau aseag anTg
pTaTJI oTJeluo'cT1c.rv u.raqlnos
puP AouS 
. 
(fSOt) 
.g.r ,:a1sog
'66L-98L 'x,
anTPrr aalldepe aqg,
aSOOg JTIOUS :assaT
pue 's'c 'deTpuTd
'TO9-9BE 'E , (uloqqcols)
- 
EIfrE' TEcTffi 
. 
(11e,,assog 
.,tr pue >1rpTc .g..{
:s-rorrpgl s1ceffir 
"uarutffien 
pfrE FaTbEle- uaasrsocg
sassaco:a 'saFb trffilTfF teFaE ur . suals.,tsoca
Tp-rnlTnc1:6e u1 6u11c.f,c ua6o:1Tu uo seJolrqJar{
a6:e1 ,{q 6ulzeeb 1o slcaJJg . (f Aef ) S.f .I{ ,a1eo1g
'iioTFETE 
.z(uo:qcu.ds anllcnpo:da.r 3ro
'If ' (sua5ffiE€c sua5se1n=aec rasuv)
aT{l uT ^,tuo:qcu.6g . (ZeOf ) ..f ,a>1oo;r
e8T
'szs-Lrs 'fr, ,ffiTfii Tq661oTtr Jo Teurnof
_EiEffiE 
.uoTXpTToJap raTTTl TeTXua.raJJTp 6u1no11o3l
''urpT ffiro-TmE rurfrfrf u1 uoTlnqT.rlsTp aleTTuTssp
pue srsaqlulsoloqa 
.e L6T .C ,11eqs:e!{ pue .n.U ,p:oJJTO
'66I-Lgr 'TOea 
, (uopuo'1) fTcos TgfGE aq1
Jo suoTT5ffi T6-E6-SoTTqd 
.sasrogol 1ue16 r{q pasn
slelTqpq o? sacuaraJar qlTr"r TTolv e:qppTv Jo uoTlcnpo:d
I:eu1:d aq; 
.(eeef) .C ,uosd111Trtd pue .A.&.C ,uosq1g
'eluealdsuuad
'6:nqspno:1s 
,.cul ssog pup uosuTr{c1nH ,uapnoq
'(11auung 
.T.d pue uazsaT,tr .T.T ,-raTTTr{ .J.d ,urrro:g .f
:s.ro1Tps) n 7- sT-seqF7tr esrF :Etsffi To.rres
1e e.rpunl TElseoc eqr,- :ruatilffi 
- W uf 
.sluar.r1nu
rTar{l pue sTros ar{f 
. 
(OeOt) 
-S.a ,1ug1g pue ,.1' .U
'a1eps:eg 
,.V.S ,da1>1:eg ,-A ,:apuexalg ,.T.d ,:ads:ag
'9EOZ-T90 Z 'G
'dbo1o6j to TEurnof ue-FEffi 
.asaa6 Aous rassaT aTpuaJ
uT alnTosqe 1ou sT duoloc 6urpaa:q Teleu o1 r{111ap1gl
1eq1 acuaprng 
. 
(ZAffi) 
..{ ,a>1oo3 pue .!{.f ,e11ure:a9
'oEe-gtg 'TE 'so-m '@ uojou7S
'sse.l6 ctua6ouBdc e pue sJaze:6 uaan?aq suorlcera?ul
'(eegf ) 
.f's ,uolq6neNcl^I pue .f .N ,stper6:oag
'962-98z 'd, '- 'epeuec 'rteg
uospnH u.raqgou pue urseg axod Jo slseoc ardl uo 6u1paa:q
spJTq Jo sarcads :a6:eT Jo uoT?nqr:?sTp aq& 
. 
(nAt)
'V 'paag pue ,.9.J ,qcoo3 ,.U ,:a>1ceq ,.f .V ,uo1se9
?87
'tr6-- TPurTuV Jo
ffi) asaag
Teuospas-erluT pue
aq,tr '(geer) '.r
'58?-T8? 'F
Terrrnof '(ffiG@ ffiEsraec
rrous .rassaT uT uorlPTJe/\ azTs q31nTc
-.ra1uT uT eTsa:1p reTncTTToJ Jo aro.r
'a>1oo3 pue ' 'g 'sna:pug' "n'1. 'ueuueg
'LOT-68 'F' '18.t56-ToTm ffi 'suoTlpTndod
r{sTlTJg lro.rJ saldues 1ue1d-a16u1s uro:J paleuTlsa
uoTleT.reA 'r ('t:ea) 'spng @ - _liffiE go
z(pnls 1ec16o1ocaua6 V '(oeOr) 'U '11ocg pue 'f 'V 'r{e.r9
.68-62 '6 ,6@i ffi pue ssprg
'6u1ze:6 lua1?TuualuT pue 6u11co1s snonuTluoc rapun
uoTlcnpo:d a6eq:ar{ Jo uosl:edurop 
. 
(AeOf ) 
.V.O ,uo1s1g
pue "f '6rTx ",T. '11an:og, ",1.g 'ure:q1:eg ,.v.s ,1ue:g
'e gg-gzE 'Q, ,F-.r6E Jo fi.aTcos m-A-E@
atn Jo tpurnof '6u111nc a:o sTaAaT oi'r1 1e uoTlpTToJap
1e11:ed r{1Tr'r (.t B--rs-uaae;d eod) sse:6an1q Jo d11a11cnpo:d
pue r{cuaTcTJJa aa1111adruo3 
. 
(eeet) 
..{.t ,:aqe:g
'OT-T '-sz '@-cg Fa-TTdffi Jo Ter.rrnof
'acueuurog:ad raap pa.r uo 1cedul slT pue aTl1ec .dq
6utze:6 raap pa-r Jo uoTlelTTTcBJ 
. 
(eeef ) 
.C.f ,uop:og
'6ZE-ZZ,e
'0T '4uaurabeupll aJrTpTTtl lo Teu:nof 
.sexatr Jo ?seoc JTn9
aql uo asaa6 pTTA Jo slTqer{ poo;l 
. 
GVA7) 
.C.U ,:auaze1g
' oeT-gET '8, '5Tm 
. 
po1:ad
6u1:ea: poo.rq aq1 6u1:np esaa6 Aous :aqea:6 dq asn.
lelTqeg 
. 
(leef ) 
.f ,p:epag pue ,.tr ,p:epag ,,{-f ,xno.rTg
g8T
.ppeupC ,uopuo'1 ,o1:e1ug uJalsa14
go .f,11s:anTun .srsar{f,, .o.r{d .suac,saTnlaec .EE ffi'
aE1q to saT6-le- @6 eqr, 
- 
Grct) 
.c ,poon:eg
.gc-gT ,E ,@€-ffi
-
TE-EfiE5T56Y Jo Tenrnof pueTeaz r"laN 
.6u1ze.r6 snonuTluoc
Japun a:n1sed .raAoTc alTqn-sse.r6a^d.r e uT aEueqc 6u11ca;;e
srolceJ auros 
. 
(egof ) 
.itt.U ,ureq6no:g pue .A ,s1::eg
'uopuoT'ssa.r6 cTurapecv
's1rret,e F' I6o-1o-Ts uoTlefiieE ' (trct) 
.r.c ,:ad:eg
'If,o-tr'
,{aN 'ueneqd61 ,.6:o1e:oqe1 TpuoTleN ua^erftoo.rg . (qlTrus
'H'H pue TTaApool,t 
.n.9 :s.rolTpg) snr5l=G fTffi-cg
u! tr+11lqeqs FUE @TE ur 
.d11s:an1p Teuorleta6aa
uT uoTlepa:d Jo aTo: aqtr, 
. 
(eg6t) 
.t.f ,:ad:eH
.uopuoT ,ssar6 crurapecg .@--TE'
TeoTbolocg 01 uollcnpo:1u1 
. 
(ZeOf) 
.f ,au:oq:eg
'sac.Inosau
TprnleN go .d:1s1uT!{ oTrplug .dd e6 (afltpfTU) luaurl.redaq
qc:pasau 'E6f6ei Fs' uospru{ uffiros aE frT ffi-6
@ ,ooB ffir pue anE aql F q6Fa p.ffirE
EEET F @ ri6-r+ErT-6re - krct) -u.s
'uo1:o11 pue '.f 
.1' ,qcu.d1 ,.9.H ,uapsurn'1 ,.C.H ,uosue11
' go z-86T ,-ge ,5!ffi . XJTTdn r{ceaq paleraTacoe
iroJ lusTupqcau alqtssod e - eqolTupl^I ,11;qc:nqC Ap euoz
TepTlraluT aql uT luaudolalap lsorJe'\.rad 
. 
(e eOf ) 
.C
'epoqoag pue ,.U ,q1:og1ue1g ,.V.d ,l1ocs ,.e.I.U ,lasueg
987
aq1
'eppueC 'uo1s6u1x ,r{11s:aaTug sruaane ,s1saq6 .cS.g
'ffis 6rrTi5TF6T@ Jo erou aq1 6oE noG ffi
q E-*adffi' FG' @ Tpcor . (ee6r) .s.o ,tTH
.I7I-ZEI 't-V ,S:1TO -sa1e.rqa1:aa .dq
spaas Jo 1es:ads;p cTlsTTenlnu Jo asec eq1 :uoTlnTolaoc
TeurTue-?ue1d gro slueururralaq 
. 
(qgeef ) 
-n.C ,eraJ.rag
-9tt'fr
ue
'Sm; '.d11s:anTp Ipoq-ecTTTs Jo
:suorlerper razetb/sseeg 
. 
(SeOf)
'Ltn
uoTlela:d:a1u1
'n'C 'e:a::eg
v :uoTlnTo^aoc
'!{'c 'erartra11
'LqZ-VqZ 'ET ,so-mi .?uauuroc
pue ursrTpnlnu ,s.raze.t6 ,sasselg . (ZgAf )
'LOgZ-ZO9Z
'E 
,@ 16 @rof E-TFe-6 .sTspo uasap :e1od
e Jo s1ue1d e.rpunl aarql uo uoTlezTTTX.raJ Jo slcaJJE
'(fe6f) 
.f 'epoqoag pue ,.9 ,ueupaa:g '.g ,d:uag
. 
z8z-96T '-Iz 
,freT ffi Tpnuuv I6@-Tg' auTreI^I FE
XEaeT6oupaco 
. 
salerqa?:elur aurreur ^dq ae61e TepTlraluT
;o 6ulzerg 
. 
(eeOt) 
.g.U ,11ou1:eH pue .f .S ,su11neg
'EI?,-ZOZ
'dE 'S)aTb 
.suoTleTndod a.rolTq.raq Jo uorlpnlcnTJ ar{l
uT secuaJap 1ue1d Jo aTor ar{1 uO 
. 
(OeOf ) 
.g ,e[ol>1neg
. 
nEz
-EzZ '1fr 'trbo-ftEf Jo Tpurnof @E@ 
.ssaccns 6utlsau
asoog anTg 6u11cagr;e s.rolcpd 
- 
ftrct) 
.n.f ,,{aa:eg
'sE-8t 'fr 'FEffi56ffi 6ffiTm F FffioT .asaao Aous
:assaT Eo .d6o1oca 6urpaegr :euruns 
. 
(trct) 
.C ,poorrt:eg
Z8T
'LTE-nOe 'dd '{.rotr AaN 'ssa:6 cTuappcg
'(TTaXcn 'hI'O pue :abunol 'g'A :sroXTpg) sasserg
Jo uoTlezTllln pue f63-1oTg' €rttrr- urqryto:6 an11e1a6aa
o? uoTlpTa: uT uoTleTloJaq '(Zrct) 
.H.q ':ap.dg
'8?9
-L?s 'Z 'EffiT5Ei ffi Jo rpu-rnoT trmffi '^deg saupf
'r{ueq1g 1:og 1p lJTTdn TerceT6lso6 . '(OeOf ) ',f 'g ':a1ung
'eTuroJTTec'p:o;ue1g'ssar4 .[11s:aalug
proJuels 'slueTd .IeTncsPA aI{1 Jo TenueI^I V :saTJolTr.Ia,Ir
6u;-:ffi6@- pup e1sery Jo e:oTd '(ggGT) 
.g 'ua11ng
')t.rotr rtraN 'ssaf4
^f,1r:an1un p.roJxo '@ pue ffi Jo sd@]ffi!
TE-T5o-I6oE- '(eeor) 'c't '.da11sag pue 
.J.H 'ano11
'zv-Te '62
'@seg FmEnfrc;:68 3lo terrrnof ftT]'!ffinf 
.s1ue1d
TETuuaJad euoz pTre auos Jo rdlqo.r6 1oo: ur uoTlerToJap Jo
loaJJg '(trct) 
.g.lt ,6ut>1cag speg pue .C.X ,uosu1>16pog
'gT-?T 'E '(u11:ag) e 
.srsaqlodfiq uo11ez1u11do
buyzetb aql pue salejr qlrro:6 a^TleTau 
. 
(f eOt)
'I'n ':adq pue '.X.f ,6u111aq ,.n.q ,?JTnS ,.A{.q ,1:aqTTH
'Oee-gze 'dd 
.eAe11O 'sagpnlg utaqlroN
roJ saT?Ts:a^Tun upTpeueC Jo uoTlercossv 
. 
(uosuqog
'9'd pue surepv 
.d.l{ :srolTpg) geef ,6T-8T .raqualoN
'sarpnls utaql.roN uo asua.raJuoc luapnls TpuoT?eN aql
Jo sbuTpaacoJd :r{1.roN sreppupC uT r{creasa5 +uaDh+S ut
'eqolTuer{ u.raq1:ou ,uot1e1a6an qs:eu ?Tes Jo uoTlcnpo:d
aql pup asaag Jqous JassaT ,.f,:orrrq:ag . (ggOt) .S.q ,>{TH
887
'6LL-?nL 'Tg' 'trFog
to Terrrnof GEEEE 
.EEp6TE6frTd x eT.Tte- lueTd
Tenxase aq1 Jo suorleTndod uaarqlaq pup uTrIlTA uoTleTre^
cTlauag 
. 
(eeOf ) 
.q.f ,qaTT1?og pue .T.U ,sag:aEg:ag
. 
ssa.rd
uT 'spue1:aqlaN aq,tr ,lqca:1n .6:ffi 6-T}E6I6EA
6 mnfffiG E6..F@E aq1 g,o s@ ur .asaao
AouS rassaT .dq 6u16e:ogl o1 asuodsal uT uo11e4a6a^ Tplseoc
cTlcre uT ssaco:d pue ural?pd . (qeeOf ) .t.U ,sa1:agrgal
'T9e-Otg 
.dd .uopuoT ,TTpH pue uerudeqp . (uTef .X.S
pue qaTT1log .Cr.T :s.ro1Tps) @'@ ffi
uI 'q1uo:6 1ue1d :og sao.rnosa.r pue ,suo11cetaluT TeurTue
-1ue1d 'scresout TeuoTlela6aA 
. 
(eeeOt) 
.f .U ,sa1:agi;ag
.?,Lg-tgg '6 ,@-cg Jo
'fffijo1' 'ecTJaurv q1.roN cT?cJp uT salTs Telspoc euos 1p
saqs.reu lTes Jo uorlelaberr aqg, . (ttAt) .f .u ,sar:eggrag
'56-0t 'dd '{ro1
$aN 'ssa:6 .d11s:a.rrtun p.roJxo . (:aqcnog .H.o ::o11pg)
6'F'ntofE pup ffi @fr1rm F ffioT€ aE ur
'slrsTfen+nu Jo I:o1s1q Te-rn1eu ar{J, . (geOt) .H.O ,uazuel
.ssa:d ur
'6 '@ Jo TeffioT CF@ 
-asoog Aous .rassar ar{1
uT ssaccns alTlcnpo:da: uo lplTqeq 6u11sau Jo acuanTJur
aqfi '(eeOf ) 
.A.U ,11aan1cou pup .S.q ,{TH ,.T.S ,uos>1ceg
687
'6oz-eoz 'Z 'X6o-F-ca pue so-r-+pT@G [.
's4ue1d cTlcrv ;o zklslruaqcol.dqd aq1 u1 suralled 
. 
(AtAt)
'S'O ':a16a1g pue 'O.9 'TTzleg , .g .f-.H ,ulqceog
'sv-LE '63 '6-rEa-5aA
'>1reuuaq 'ua6u111p>ts 1E uorlela6al qsrplu-lTps uo 6ulzezb
daaqs pup eT?1ec Jo lcaJJa aqf,r 
. 
(Seef) 
.V ,uesual'
.11LV-1T? .dd ..tr.N ,:a>1caq . (ppB:
'N'f pup Tned 'v.g :s:olTpg) g aurnToA ,XfE-@Tg
TTOS UI '.IA/\OU.InI pue ?Uaua.Inseaur :TTOS Ur SSEIUOTq
TeTqo:cTI^I '(feOt) 'H'f 'ppe1 pue 'S'q 'uosu1>1uag
'09 'd 'pr:aqueC ,acuarcg gro .durapecv uerTe.rlsnv . (sueTTtTl{
'g'o pue qcur(1 
.u.d ,sso1'.f.d :sJolTpg) acua.r€m
pufF teuo@ Fuocas ffi Jo s6F-:d
- 
E$un actnosau : spueTSbueu ur 
. 
Tapour {cpqpaaJ
aarllsod e - uoTlela6aa p:punl uo 6urze:6 ;o slcaJJg
' (geof ) 'n's '1116:e3 pue ' 'U'q 'r(1azeg ' 'T'U 'sa1:a;gag
'ogtT-6gtT 'Ts 'F-s F
Tprrrnof EF@ 
.pqolrupt{ ,r(eg asnotad eT 1e uorlelabaa
uo asaa6 Jo 4caJJa aql pue ?uaudolaaap TpuoTlpla6aA
'(et;t) 'a'X 'ureqe:qg' pue 
.V ,uasuag ,.T-U ,sa1:aEEa1
.dd 8 .ecrl.ras aJTTpTTA! upTpeupc aql
o1 1:oda: peqsrTqndu6l 
-huTo-e 
- 
F &TuTm
Ttslspoc ar11 uT EoTFIalba-A aq1 uo asaac e' ffi-T6.etod
F ffi aq;, 
. 
(geof ) 
.u.u ,saq:ay pue .T.U ,salragi;ag
057
'pa?lTurqns 'XFt6Fg Jo Terrrnof
'asaa6 dq 6u16e:ogi Jo lTnse.r e se slelTqer{ pueTla/{
Jo uorlepa:6ap aq1 pue saTlTunruuoc ?ueTd Jo uorlcnrlsaq
' (NeOf ) 
.t.U ,sat.ra3lgiag , .hI.d ,uaueloy , .H.U saqrax
. 
TT6-906 
,66'T ,IEn- E ap 1a 5-rTo-To-ctr,p
an^euI- CTp@ affihTtr 
.r(eg uospnH lsat{ uo
lelTqer{ JTaql pue asaag luous rassaT 
- 
(Zeffi) 
.g-U ,saq:ay
'r(as:ag atap 'sJJTTc
poonal6ug 'TTeH acTlua.rd 
. 
(uoT+Tpg puocag).
sraqS.reasau E :s.T-F pue u- 
. 
(zeor) 
.g
'Lo?-ooz '€e, 'Xbb- 
.puoqeT{o Ter?uac uT ar:te:d sse.r6
Tfp? alrlpu p Jo uor?cnpo:d 1ue1d pue uorltsodruoc aq1
uo 6u1zer6 a1e:apour Jo slcaJJg 
. 
(tSOf) 
.U.U ,6u111ay
'8TE-TTE 'E 
, (u11rag)
U16o1oca9 
.f mnleufben urf.ro!ffi ,a6pas uolloo
aqX Jo acupTeq luar.rlnu JoJ luaudolalap JeaT lelluanbas
Jo aouecr3itubls 
. 
(geot) 
.S..r ,uldeqp pue .s ,uosseuo4
{ooqpueH
'ladday
'nLg-cLg '-zg ,ffiT5si TpurTuv Jo
'asnoquaa:6 aq1 u1 qllro:6a: sse:6
Jo acuanTJul 
. 
(Zrct) 
.g.C ,.da11eg
Teu.rnof ueTpeuec
uo p/\TTes auT^oq
pue 'V 'uolsuqoj
'LgE-gve 'm. ,F66- @Toffi
f o f Bu:noC 
. 
butzetb Japun q1r,ro:6 sse:6 ;io s1s.d1eue
TecTlaroaql V 
. 
(EAO1) 
.f.V ,suos:e6 pue .U.I ,uosuqog
'ELI-6IT 'fr, ' (ut1:ag) pTfr-cao
'E{seTV cTlcjrv ur su.roJ 1ue1d prpunl uT su:a11ed
stsaqlu.dsoloqd pue ,q1lro:6 JeaT ,uorlecoTTe sspluorq
puno:6aaoqg 
. 
Grct) 
.t.f ,uaza1f,, pue .V.O ,uosuqo4
T6T
T9 g-09 C
'EEm'rusTTpnlrrr' arolTq.raq-1ueTd Jo suoTlpcTTdurT
ssaulTd '(geOt) 'q 'uoabpnq pue 
.S.X.d 'rrrpT
'0?8
-gzg 'E 'FTffir{coTa pue i[6Tofe fF 'uoT?ezTf€rauTu
pue uoTlezTTTqouurT ua6o:11u 6u1:np TTos u1 ua6o:11u
-pTce ouTurp aTqnTos-pTce Jo uoTlnqTrlsTp pue I11nTlce
clru.dzua u1 sa6ueq3 '(e fOt) 'V'g '1ne6 pue 'N'f 'ppeT
xa.rPc Jo
puP JeaT
'gL6-Tg6 'dI 'XF-cg Jo Furnof 'ea€@s
s1ue1d paze:6un pue paze:6 gro Iqde:6omap looqs
aq,t '(eeOt) 'f 'U 'sa1:agr;af pup 't{'d 'uaueloy
'LV;,-ZVE 'dd 'eAe11O 'sa1pn1g
uJaI{lJoN JoJ saTlTsJaATUn uPrpeuPc Jo uoTleTcossv
'(uosuqog 
.g'd pue suppv .d'$ :s.roXTpg) geef ,6T-87
.raqualoN '$ETEI utarnroN uo acuaraJuoc ffils FGm
al11 Jo @-:d :qUoN srepeuec ur @El[ luepnls
ureqolTuehl 'r(eg asno:gld pT '.d:oa1q:aq 01 saTaTunuruoc
e6pas Jo sasuodsa: aqg, '(ggOf) 
.n.a ,uaueloy
'pppupC'o1uo:o,1, go .611s:aATuO 
.sTsaq; .cS.n .asaag
r.ro$ -rassaT Iq 6uT5-sroJ 01 ffi &s Eo sasuoclsar
qli'roT5 aq1 Jo uosT-r-Afc' V ' (eeor) .n.a ,uaueloy
'PTU.roJTTEC
'a1o3ls:1oorg 
. 
(uoTaTpA puocasl sacualcs FloT^eqag aql
roJ sffipaco-rd :uFo T€luaurfaffi 
. 
(zeot) 
.g.u ,)trTx
'gLz-Igz 'n 'm 
.Xear{r'r uT
a1E.I cllaqluztsoloqd uo uoTlpzTTT?n alPTTtuTsse Jo lcaJJg
'(nat) ',r't 'sueag pue ".tr'I ,rrlelp:eir1 ,.&.u ,6uTx
z6T
'orre1uo 'uopuo.1 'o1:e1ug u.ralsa!{ ;o r{11s:alTufl
'q'qd 7.@ TTauuoffi arta 1E ffi
'slsaqg,
anTg JoZffiocg 6u-Lp- rarnuns ' G tet) 'c 'g ' JJaTT
AaTAaE Tenuuv
EpPUEC
FG
'6e-62 'dd ')tJotr i'reN 'ssar6 .d11s:aalun p.roJxo
' (.raqcnog 'H'q ::o11ps) ii6, FTE pup f6|ocs TfrSTf1nm
F fffi-FE, eq,r, u1scT?upluas z(66os pue sldacuoc
dsT:c :lusrTenlnur pue s1so1qu.f,g '(gget) 'tt'g 's1na-1
pue f6-ofrffi 3ro
01 slueTd Jo
. 
GEr-rzr 'Z 
,EsT]ffilEEi
'sa:oATqJaq pue suaFoqled
sacuaJap TecTuraqc aqfi 'G6f.) 'q rulrra'1
'Eoz-68T '€L 'Eeprv- 'saTTTlupJ
asoog Aous f,assaT Jo .rorleqaq :slabpnq auTl pue
azTs poo.rq '1uarl1saau1 Te?uaJed '(fggf ) 'n'C 'slassa'1
'zzs-eTg 'ET 'X6-Fcs-
to Teurnof 'F€pf@ toffiqv 1ue1d Tenuue aql
Jo uorXcnpo:da: pue qlno:6 uo uo1111aduroc pup uoTleTToJap
Jo slcaJJg '(OgOt) 'V'.{ 'zezzeg pue 
.O',1 'aaT
'ZTI-96 'dd 'proJxo '11ar,t1ce1g '(uoslrog 'H'I pue
TTTaNcI{ 'S 'aaT 'V'1. : s.ro1Tpgl :o+ceU TE5f6-oF€g- ue S
u5ffi uI.sarlrunuuoc 1ue1d u1 rolcpJ 6u111ru11 e se
ua6o:1TN '(ceor) 'u '1n1ceu6r pue 'u ':au.reg "v'f 'aaT
'897
-Eer '&,'@E TffiTtnoT:SE Jo teu.rnor, @az r'raN
'sse:6az{: f eTuua:ad Jo q1aro.r6 aq1 lcaJgre 16un; clllqdopug
'(gegf) 'U'O 'eaerbsnlq pue ",{'l{ '?unH "n'C'g 'r{cleT
'uopuoT 'p1ou:q' p.reApg 
.nE# d6o1o1g ut saTpnls
'uoTlTpg pu7 'r'ror9 sasselg i"roH 'Gt*) 
.n'H.U ':a6uBT
e6T
'sTT-roT '-z?, 'F6:iFm 
.uoTlsa6Tp
asoTnTTao pue 6utpaaE asoogl 
. 
(f fet) 
.C.C ,s>1co11e11
'L6T-6LT '6i
'@ Jo sTeuuv 
.pJpun? cT?c.rv ue{seT.V ue uT 6urze:6
6utruual paleTnuTs o1 1:aqslg 
- 
u6ilnq' go sasuodsau
.Gtet) .C.lt ,s1rta1 pue ,.T.T ,uazaT,L r.1l.d ,s1aq11ep
'6zn-sr? '6TT '5frb-r-leE iEX$ 6F i.a $Tffi an^au
EaE@ alsTTernleN aT 
.r{eg uospnH Jo lseoc oTrelug
aql uo sa.rnleal 1ect6o1or{d:ouoag 
. 
(ZeOf ) 
.a.f ,rur1:e11
'vEz-oez '6T ,XFFcE cTec:etoE
'suralsr(soca pJpunl >1cossn1 uT NgT racerl JoJ uo1111adruo3
. 
(f eef ) 
.H.c ,:1ce1g pue .c.d ,.raTTTIit ,.n.g ,uo1:ep
'ees-629 '4, ,5ii6G6Effi
€mTE F Terrrnof 
. 
s{cnp p.rpTTpru uT a6essed pooJ
Jo aleir :uo1)tupTd Jo 1es:ads1o 
. 
(Egsr) 
.u.c ,auo1e11
. 
ELL-LgL 
,67
'1s11e:n1e11 
- addals ur uoTlnTo^g
FTE1g treTp,Puec
TTauuoccll le
aql Jo r{1Ao-r9
aqg, 'sTplrrueu paloor{ ,a6:e1 na3r qlTrc
'(ZAfi) 
.H.f uosdruoqJ, pue .U ,Icetl
'68-ee 'TbT ,]ffiAeN-
'OB6T-0?6T :saTJo?T.rJaf,r lsar*rr{lroN,rerrTg
r(uo1oc '(suac5aTn.rffi uarp) ,asoo9 /rlous
'(feOf ) 
.H.U ,saq:ay pue .q.C ,sauu1ce11
-os 'tr 
,+uEEa6EtiEE
JTn6 01 sle:>{snur
'(tvot)
EJTTPTTM JO
pue asae6 g:o
'l{'e 'de1 pue
'9L
TeffiT 'saqs.rpu lseoo
acuecTJTu6ls luauabeuep
',1 'TTaNro "f'f 'rlcudT
v6I
.v6-z6,ET ,@i
'bu1zetb .f,q atolgl .d6:aua Jo uoTle4TTTcp.{ :lsaaqapTTr,l
d:o1e:61tu t1a6ua:ag 
. 
(grct) 
.f .S ,uoXq6negcq
. 
nos-r6v 
,-v6 ,-@frE 63m .uTlenaax
.IOTralUT uT asaag Jyrous JessaT Jo uoTlnqT:1sTp jrauuns
pue uoTle:61n '(zear) .v.n ,ua:e'1c1a1 pue .T-d ,ua:e1c11
'srE-999 '4, '@Effi mTffi pup ffi 
.uor1e1a6aa
pue sTTos e.rpunl uo uoTlezTTTl.ral pue saro^rqraq
ueTTeuluteul Jo slcaJJa auos '(OgOf ) 
.C.1' ,uosueng
pue "U.X ,11a:aag ,.O.g ,TTzleg ,.q-f ,:1c1:puayc14
. 
ETT-6?
'ei ' (u1oq>1cols) 
-ffitr F5fffi 
. 
(lleatssos 
.& pue
{:eTc 
.9..{ :s.rolTpgl se-c luerueE-upn FG' sa-Tffi
uralsffi ffird 
.saTt6 EE@ ffiET ur
's1ros puefsser6 uT N pue c Jo sclrueu.fiq . (f eof ) .o.r
'ssnau pue ,.g.U ,aasueupooll ,.ll.H ,?unH ,.g.1{ ,TTTgclt
'9gT-ZET 'B , (ur1-rag) eTFTocaO .ae61e paze:6
Jar{lo Jo luauaouequa q1lao:6 pup r{sTJ suo^T1)tueldolIqd
6u1ze:6 e Jo luauacuequa q1Ao.r9 
. 
(SAOf ) 
.g.r{ ,pleuoqcpq
'z?,s-irg '6EE ,- i
'uoT?cnpo:d .f,:eur:d 1sa:ogr Jo s:o1e1n6a.r se slcasuT
sno6eqdo+r(qa 
- 
GtAt) 
.O.H ,dppg pue .f.t{ ,uos11eq
'TeT-6TT 'TT 's51-lI@E-lGi
Fup f6o1ocg Jo AaT^au Ten@ 
.lua?uoc ua6o:1;u
1ue1d 01 uoTXpTa: uT d:oalq:ag 
. 
(Oeef ) 
.1' .irl ,uos11e1^1
E6T
uecT.rauv
uT sTeurTuv
'988-998 'm,
eq&- 'uorlnTolaoc pue ,ur:o3t
:suApT 6u1ze:9 '(ueet) 
.f.S
,lm1PT
1ue1d 'spraq
'uolq6negcp
'FtfcfeiEbb-l,t
dcua6ulluoc
Jo aroJ aqtr'
'uolqFnBNcIaI
'oze-r6z 'EE
fe-cT61-fi 
. 
uoTlpzTueb:o ^dllunruuroc uT
pue s:o1ceJ Teluauuo:Tlua allsodruoc
:d6o1oca pueTsserb r1a6ua:ag 
. 
(ceeet) 
.1'.S
' g L9- L99
'dd 'uTTrag '6e1.ra4-:a6u1:dg 
. 
(:a16a1Z 
.H pue ,puoursg
'a'O'aTqoN 
.S.d,a6ueT .T.o :sro?Tpu) TffirT^rrg
fTffi-To-Tg Fup 't.E. aq1 01 sasuo@ -Tr f6_Tocs
ffi Te5i@ ur 
-r(:orr1q:aq Jo suorlecTTdurT
lecr6oloca pue 1ec16o1oTstrqa 
. 
(qeeOf) 
-f.S ,uo1q6negc4
'98t.-6?,e 'G 's-6m; 'd.roa1q:aq 01 asuodsa:
E se qyao:6 1ue1d tr:olesuadruoC 
. 
(eeeet) 
.C.S ,uo1q6negc4
'COT-28 'dd 'o6ec1q3 'ssa:4
o6ec1q3 Eo tr11s:a^Tun , (sr{+TJJrrg-uogoN .W pue .rTeTcuTS
'g'U'v :srolTpg) uralsl- ue 3:o 35$ffire :-gE6ffi! ur
'sclueur{p e.ro^Tqraq-pueTssp.rg 
. 
(qOeOf ) 
.n.S ,uoqq6nepcq
. 
EoL-T6e ,m ,TsT-TE=im 
- 
dqr,
't1a6ua:as aq1 uT sdlqsuotlpTa: a1e1n6un-ssp:g :ssaco:d
uoTlezTurTldo ue sp 6u1ze:9 
- 
Geta1) 
.C.S ,uolqbnegc14
to8-9og 'R 
,ffis .sp11n6
asuaJap 1ue1d Jo ssaua^TlcaJJa aq1 sacuanTJuT r(11a11ca1as
6u1paa; :sa1e1n6un 11a6ua.ras 
. 
(efef ) 
.f .S ,uolq6negcq
967
Jot{1AoI6uo;:]:;.::",T.;m;H:,::;:":]::j; :1xa1uoc uals.dsoca ue ur uotleldepp lueTd . (eeOt)
'g'n ':nouaq6noC pup ,.T.T ,ace11e6 ,.f -S ,uo1q6netr1c1n1
'euor{eT{O,ueu:o1,1 rssa.l6 euoqpT)to
3ro 
.t{1ls.ra^Tun 
. 
(ue>1un.rg 
.N.f pue a:r(I, .f .U :s:o1Tpg)
f66tocg Fue scTTpura?sifi : spueTssm pue sasse:e ur
'sua1s.,{soca pupTsse:b u1 sassacoJd arrllce.Ialul 
. 
(ZeOf )
'T'T 'ace11e6 pue , .g.r{ ,:nouaq6no3 , .1' . S ,uo1q6neNcI,I
'629I-LTII '6 ,XFF€g-
sploulue:6 tc uo uoTlpTToJap
slcaJJg' (geor)'rrrs'J
'su1e1d r1a6ua:as ar{l uo.rJ
pue uoTlT.rlnu sn.roqdsor{d Jo
'u1deq3 pue 'f 'S 'uolq6neNcl^I
.T6L-16L ,6T , (vsn) se-ffiT$ Jo Xruepecg
Fftffi aql ;:o F6Fp-rd 
.sa.ronTqrar{ lsuTe6e asua;rap
aTqTcnpuT ue roJ uoTlcaTas TprnleN :uor?pBcTJTcTTTs
JeaT sse.rg 
. 
(e ee f ) 
.t.C ,s1ue:-re& pup .f .S ,uolq6nepcq
.OLL-ggL 'M ,+sTre:iFT .
aqilsa-ro^Tqiraq pue s1ue1d u6 '(geot) 'r.s 'uolrdbnBNcI.I
'98V-8LV '6 , (uT1.rag)
e}6@Go 
.uoTlr.rlnu ctue6irouT pue ,1ecluaqc .d.rea11es
p 'uorletloJap r{q satl.rado.rd TpcTuer{c pup pTarz( sse:6
Jo uoT1e1n6a: alrlce;eluI 
. 
(qgeef) 
.C.S ,uo1q6negcry
'n6z-692
'99 's[dETfru-o14 Te-cT6{o-ctr 
. 
r1a6ua:as aq& :urals.dsoca
6u;ze:6 p Jo ,t6o1ocg . (eseot) .f .s ,uolqbnepcp
L6I
'TB?-6E? '-Eg
'X6_ftr-ca Jo Tpu:noT 'sqde:6 z(a:d-:o1epa:d Jo uor4eclldde
up :sualsds burzezb Jo dlTTTqelS '(gaef ) 'f ':1aq-r(og
'8Tg
-TTe 'E '(u11:ag) p-Tffi-To-cao 'acueraTol r[:orrlqraq .roJ
suoTlpoTTdurT :pTaTJ aql uT slsaqluz(soloqd r(rolesuaduoc
Jo lsa1 V '(teef ) 'n'n '11a.rrp1eC pup 'S'U '>1eno1,1
'gET-oET '6 '(u11:eg) e-Tffi '(sTsuepeued
rolsec) :aleaq .dq pa:a1Te urea.rls crlc.reqns p Jo
1a6pnq ua6o:1TN '(regr) 'n'c 'oTTTTaI^I pue 'f 'u 'ueurleg
'90z-86T '7e, '5cuaTcsi6E
'.d:oalq:aq 01 acua:aJa: uT uoTlcunJ pue a.rn1cnrls JeaT
uo slurprlsuoC '(ZgOt) 't'S 'uour1ng pue 
.V.H 'r{auoohl
'99?'-I- 'dd 'uopuo'1 'sq1:oru:a11ng 
. 
(su1..\1 
.g.f pup
ad:ot{1TTlt 'T'.{ :srolTpg) 55-sse=6 pue sTparac Jo qar'roi6
aE uf 'sasser6 u1 q1no.r6a: gro slcadse 1ecl6o1oTsr(qa
'(ggOt) 
.t.C 'uosptaeq pue .T.,it ,adroq111j,1
'90?-T9E 'fr 'EifdE:_6b-uo1a1 F-tT5-oTo-cs 
.qceo:dde 6u11aporu
sruals.tts e :p>1seTg ur e.rpun1 {cossnl ul-TEtr.TbE^ ffiToqd61-
uT sassaco:d TTos-?ueTd '(reef) 
.t ,l:enls pue
"A'S 's1:aqog '.O.M 'leqcag 
,.r{.9 'uo1:eg4 ,.9.V ,suT)tuTT
''f 'no:auuny 
,.M.o ,+raqTTH ,.u-x ,11a:arrg ,IrI .s.,jt
'ugdeq3 
,.!{ ,uasqocef-a{pTg ,.n.d ,JaTTTI{ ,.c.d ,raTfT}.l
'ErT-seT 'TE, 'FffiAbfeuet^t
em Jo Enrnof 'Iq11enb laTp 01 uoTlefa.r
uT sp.reTTpur go z[6o1oqd:ou 1ng '(grer) 
.U.n '.raTTTn
857
'z6t-Bgv 'Tz 'EbTlo- assaulTJ 1ue1d azTuTxeu
s.raunsuoc oq 
.(gtot) .c.u '1ra6a1A pue .E.q ,uano
EEZ-OeZ
aq1 uoirJ lTJeuaq
F-fT drroq
etlffi-1ffi 'raunsuoc e
'ff 'ml 'uaq? 1ea 1Br{1 sreuTue
,{eu s1ue1d atog '(OeOf ) 
..f .q 'uaAg
'TLZ-ggZ '|p, '="m 
.unffirrn-F
aql uT TTPJ-Jear TPuoseas uo -
Jo lcaJJa aq& ' (efef ) ',{'q 'uang
'f 'N 'uo1acu1:6 'ssaJ6 d11s:aaTug uolaouT.rd
'snra1sffi Jo Tffi5 TeST'@ Y '(geor) 'H'J',1
'ue11g pue 'g'1, 'apTeg "T'O 'srla6ugaq '.A.U 'TTTaNTO
'0ET-6ST 'dd 'an6eH aq;,'s:aqst1qn4'A'g {unf 'l{
'Iq '(11auuocc!{-aruoT 'H'u pue uaqqoq ue^ 
.I{.1{ :s:o11pg)
fEroTo-cg uT sAiI6@ 6-uTffi uf 
. 
sualsr(soca Tern?pu uT
^f,11:n1eru pue d111Tqe1s ,tr11s.raa1c . (grer) .u.c ,sue1:9
'es-6v 'e 'EEr{sT.{- Jo- K66--Tg FFffi=T^ug-
'saTxTununuoc JaaJ Te.roc uT suTr{c.rn pue ser{sTJ snoJo^Tq.raq
Jo aTor aqtr 
. 
rct1t) 
.t.S ,1eqo1 pue .C.f ,uap69
'vvv-tzv 'm 'FTEfim 
- 
eq& aaoualsTxa
roJ a166n:1s ueTurnJpq uTeTd ro scrlau.zaqr(c pue
lursTTPnlnI^I :UOTleZTuB6:o Ualsr{Socg 
. 
(ggar) 
-t ,uaues>1g
'oz,t,-I6z 'Q, '3@e=mlroi,i Tec-T661- 
. 
TToAv {o?anTus
uo r{11unuuoc Jaar Te.roc pJervrpuTru e Jo .f,1tar1cnpo;rd
pue arnlcn.r?s clqdo:; 
. 
(SgOf ) 
.a.g ,runp6 pue .,I-H ,unpg
'96-99 ':{i 'X6- 'sualsdsoce pTaTJ pTo Jo sluauodruoc
.raunsuoc .6:eu1.rd aa.rq? Jo AoTJ r(6.raua uolXelndo6
'QgAt) 'g't '1:oduazrpc pue '.9.C '11auuo3 ,.d.9 'unpo
667
'z9e
-T9€ 'fr 'FbEft;qo-ated- 
.sarolTqraqp6au Jo aTo: Tplo^Td
aq1 :suoTXcuTlxa auacolsreTd 
. 
(feOf) 
.H ,q11ug-uaarg
'gz-Tz 'Tz
's6JFm 'asaag aTcpureg 6unod .fiq uol1e1a6al pazTTrlraJ
-ua6o:11u pup sarcads sse:6 uaaAlaq uoTleuTurTJcsTq
'(ttOt) 
.n 'salaeq pue '.n ,1ua6ng ,.n ,uang
'p:oEsleg'pTiroM ar{1 Jo asaa9 pTTt{
-sTL 'fr '?[bEFEE p-5TTT€ 3o TerrrnoT
z[q pooJ :aluTn Jo uoTlcaTas aq;
'uopuoT
'(oeet) 'n 'uaAo
'6ZL
'asaag paluo.rJalTqM
'(gror) 'n 'uatrt6
'r9T-s9T 'dE,
'-uaur56@ SFfFm Jo [Effio1' 
.luaua6eueu asoo6 :a1ugn
roJ pupTsse;tb 6u1z11TUaJ pue 6u111n3 
- 
(StOt) 
-n ,uang
'gET-?ET 'Ti 'm; '.d6o1oca r(:euollnTola
uT uoTlpTncads :ogl paau ar{? pue ,,.d:a1sr(ru aTqpuTlnoe€rr
srurnreo 
. 
(ceor) 
.e .u ,1:a6a16 pue .,{.q ,uang
.9TT
-ETT 'Ti, 'ffi 'utsTTenlnu:eze.rb/ssezb uo a.roln :aarl
lnTea aq1 6u11eag 
. 
(qZAOf ) 
.e.U ,1:aba161 pue .,jI.g ,uaAg
'69Z--89z- 'EE
puP sasserc
'STF; 's1saq1od.f,q
uaa/tlaq uIsTTenlnI^I
'ffi; aursTTenlnu P araql sT :s:aze:6
'(eZget) 'C'U '1ra6ar6 pue ',jI'q 'uang
d:euollnTo^a up :s:aze:6 ::: :::=":
. 
(reot) 
.c-u ,1:a6a11r1 pue .,{.O ,uaag
00e
'm, 
,m=iFT uecTraurv ar.{I,
TEurTuP 6u11apo4 
. 
(eeer)
'96-69
'sJolcear TPcTuaqo se s1n6
'V'd 's:eunp pue 
.T.CI ,r(:uad
-ess 'Efr,Im+eN
cTlau:aq^f,c aqg,
'E68
uecTJaulv aql- 'suralsrtsoca Jo aJnlPu
'(teOt) 'a'g 'urnpg pue 
.C.g ,ua1led
'LZ-sr '6 '6G-iGi 5ffiT FG ffi 
.prpAs
paze:6 .d11euo11e1oJ p uT q1,ao:6 Jo ale: a6e:aae aq1 pue
qleap JeaT 's1saq1u.f,so1oqd uo r{1Jyro.rba.r ;o uoTlernp aq?,
Jo loaJJa ar{,1 
. 
(eeOf ) 
.O.a ,6utuua6 pue .f .V ,suos:e6
'69-6n
'6 '_ i ffi pue sserg 
.sse.r6 p Jo uoTleTfoJap
lual1Tuua?uT pue snonurluoc aq? Jo uost.reduoc TelueuapunJ
p apTlo.rd o1 pup uor?eTToJep lua1lTurra1uT ;o I11:aAas pue
rlcuanba:E uaaAlaq uorlceJalur ar{A azTurlldo o1 Tapou e Jo
asn '996T 
.V ,r{an:eg pue .U.I ,uosuqog , .f .V ,suos;e4
'lL
-89 'Ti, 'Fffi56Elijm a'6ueu Jo TerirnoT 
.sse:61eaqn u.ralsan
Jo r{1/qor6a.r pue srseqlustrsoloqd lau uo uoTleTToJap
Jo slceJJg 
. 
(f e6f ) 
.X.f ,6u111aq pue .T.g ,:a1uge4
'srr-ErT'rg's{dE=b@,'IF-'T5-oTo-cs
'u.ra11ed Jo scrueu'{p aq1 pue acueq.rnlsTq :sadecspuel
fppTlralul 
. 
(feef ) 
.V.S ,uraa1 pue .&.U ,au1e6
'grv-Lov 'm, 'lFfTEmT uecf:aruv eqt- 
.ua?ea 6u1aq
Eo a6elupape aq1 :d:on1q:aq uprTeunrpul o1 asuodsa.r uT
uoTlpsuaduoc:aao 
. 
(feOf ) 
.e .,f ,ueq11q6 pue .N.X ,a61e6
roz
. 
e/vrP110
'epeue3 Jo sunasnhl TeuoTf,pN 'sacualcg Te:nleN Jo urnasnr{
TeuoTleN 'epeuec %a"T:oTTffiE ffirnroN @. 5
Jo ffi reTnSsp1 
. 
(oeot) 
.r.u 'z(po3 pue .g.v ,p11s:o6
'epeue3 'eatellg ' VZ 
. 
oN saTras
1ec16o1oTg uT '26 'oN 'q]3-ffiE @ Jo utnasnl^I @m
'e+r{r{do+Bnr:ads pue elIqdopl:a16 
.I l:pd ,c11c.rg
uralseg ueTpeupc aq1 Jo r{uelog 
. 
(oror) 
.H ,u1un1o4
'99-92 'dd 'p.roJxo '11aru>1ce1g '(uosl:og
s?ueTd JouoTlTrlnNFEEffi arn Jo slcaffi
'H'I ::o11pg)
Te5lffiocs ur
aq,tr
'saqsreu lTes Tplseoc uT s1ue1d 6u1:a,,ao1gr Jo uorleuoz aq1
uo ua6o:11u fe.reuTur Jo ocuanTJul 
. 
(OgOf ) 
.O.C ,lXobTd
'>tJoI rtaN 'ssa.r6 cTurapecg
'STEftE ffi pue
. 
(seer) 
. 
(s:o+Tpa)
6BT-sBr 'ET 'FAI56-eEE d6ueu
5o of 'ssB:6 pearql-pue-aTpaau uT buyzezb f,aeaq
01 acuplsTsa.r 6ullcaEgre stolced 
. 
(Zget) 
.V-U ,uos:a1a6
'6nt-onn
'Ti '@n6I Xurouotf'Y 
.,{11111:ag a:n1sed uo lcaJJa s1T
pup aTl1ec 6ur6ue: ^d1aa:31 Jo elaroxa oJ uoTlnqTJlsTp aq,tr
'(ggOf ) 'U'm 'asnoqpoo$ pue 
.T.H ,secn1 ,.g.U ,uas.ra1a6
. 
LZT-gZ,t 
,ET ,qE; . s1ue1d pue
sJaunsuoc uaanlaq susTTpnlnur a.ror{ 
. 
(ZAffi) 
.n ,a11a1a6
'ggz-zvz 'Tt '- 'suospas
6ulpaa.rq sp:Tq 3lo 6uru1? aq& 
. 
(OaOf ) 
.n.C ,sur::a6
acueqJn?sTq
's'd 'a?TqM
Te.rnleN Jo
pue'v',1's
fFTocg
'11a{oTd
zoz
.Lrr-L?,mT,=a1mi @
XSroN'uab:aqslTds uralsan'spuno:r6 6u1paa:q
aq1 uo uoT1elToldxa pooJ 01 uoT?eTa: uT 'gfEd6ffiq
plueJg 'asoo9 aTcpu:eg aql Jo ssaccrls alTlcnpo:dag
'(leet) 'g'U '1ua:q pue "U'r{ 'uap:ag uea ,.1. 'do:d
'gZ"-ZZt 'gg '(u11:ag) affotopaO '(sTs6'cnaT ffi-)
asoo6 aTceuireq ar{1 Jo lJTqs lelTqer{ Tpuoseas e pue qgto:6
uo11e1aba6 '(EeAf ) 'C'U '6:aquapl pue ';,'H'H 'sugr6
'969-E89 '-62
'Em-ffie1q- 
-@tr 
Blcv 's?uaurT.radxa pue suoTXelrasqo
pTaTJ :6u16e1s 6u1:ds 6u1:np e6_ o6E1u- u1e1ue1d
eas pue f@! eluetg asaa6 ?uaJg Jo uoTlceraluT aq,tr
'(oeer) 'g'u '1ua:o pue "c'u '6:aquaptr "&'H'H 'su1:6
'e 9E-8sE '-6v 'TUffiE.EEbm SFtFm Jo Ter.rrnof
'r(eg saupf ?e aseag eppupc pue AouS Jo spooJ 6u1.rdg
'(SeOf ) 'g'n 'seuoq,1, pue ',iI'I '11eqs.re4 'd'f '11aaa:d
'zvL-sEL 'fr ,T[ffiE-6]e-b@ 6Jmm Jo teffio1'
'uot6a.r .t{eg sauref arll uT aseag Aous rassaT Jo spooJ TTp.{
'(5rct) 'C'n 'seuoq;, pue '.{'I '11eqs:e4 'd.f '11ana:6
'rL
'E'Eafe=bffiT 5ffiFm 'asaeb Aous uT sdno:6 TpTcos .raqlo
pue r{1tueg '(Oeef ) 
.O.C 'sauulcen puP .d.f ,11e/1'a.r4
'TrnossT!{ 'e1qurn1o3 
.fFT-cos TpcT6=t-_g 
- ffiN 
- 
fuue ffiTFrffi
pue ae61e 'qstglec pa.routtv :s:aze.:6
,{q luauacuequa acJnosa.r 1aN
'xcerlsgv '?uaurTpas
Jo slsaJJa 1ca:TpuT
'(eeOr) 'g'n ':aoo6
toz
.ssz-soz ,€t 'IffiffiF! 
- Affi 'r[:oaq1 lecr6oloca pue sclureur{p uollelndod a:onrqraq
uT acualaf a.r .rTaq[, : s1ue1d pue saJolTqJaq uaanlaq
suorlceraluT aATsuaJap-a^TSuaJJo' (Seef)'.{.O'sepeoqg
'oz,z-9gl 'dd '(a:n13c11 's't[
pue ouuao 'd'U :srolrpg) sua15fs F56@ pue TernleN
ffi saro- pue silleT? al@a uf 'rtrqsluaqc 1ue1d
pup scTrueu^f,p uotlelndod a:onlqraH '(CeOf ) ',f 'q 'sapeoqg
'(uazueg 'H'g pue Ter{luasog 'V'9
:srolTpg) salftoqelehl lueld fuppuocas qm 6ry
-ffiiif 
:sa:onTq:aH uf 'saJolTq.raq lsuTe6e sacuaJap
Tpcruraqo 1ue1d Jo uoTXnToAg '(e rct) ',f 'O 'sapeor{U
.9z-Tz'6,(u11:ag)ef@.adocs1:ad1oo:parro:dru1
ue l{lTiyt suor?pl:asqo pTeTJ : sasse:6qcunq uorr(do:6v or'r1
uT uoTleTToJap oA asuodse: loog '(AAet) 
.U.f 'sp.rer{cT6
'99T-gET 'n,
'pueTu.reJ pue UoT T:lnu 'asaag '. Grct)
'To?-oo? 'Tz 'iEffiE;6@T a6upu F Teriffi
pup errTTES TpurTue o1 eue.r6 sleoapTs glo asuodsag
'g'T'f '11t::a6 pue'T'C '.raqearu1e"1 "O'd
,IA6JrFTT},I
'v 'paau
'auTueTql
'ftrct)
'uop:eag
'gog-L6g 'Tt 'ffiEi 't. Eo
Teffiof 'sasse:6 uo eATTes auTloq Jo lcaJJa aq& 
. 
(7rct)
'g'T'f '111::a6 pue 'T'C ':aqaarural "O.d 'uop.leag
'uoole{ses'uer'raqcle)lses 3ro z(1ls:a^Tun
'par{'1a1't[ 6@Es e uo TsuacsaTnJapc 
.c.rasuvl
asaag AouS .rassa'I Jo saseasTq '(ZgSt) 
.f .q 'a1uu1eg
voz
'887
-g8T 'TF 's{To- 'eso.nffi EJ6-u--r-!1flS ut a1e1dn luarr}nu
uo acrnos ua6o:1Tu pup 6u1dd1Tc Jo slcaJJa TpXuaurT.radxa
zbulzetb pue luaualour luar:1nN 
. 
(leef) 
.m.U ,ssang
. 
s98-6e 8 
,6'F ,Im=nlEN
- 
58 ;asaa6 Aous uT azTs r{cln1c 1eu11do up araql
sI '(fggt) 
..f 'a>1oog pue .S.C ,.de1purg ,.,{.U ,11erru1cog
'ZZe-gIe '9E , (u11.rag) eT@ .dllpuncagi uo a6e pue
aluTl Jo acuanTJuT aqfi 'r '(ffi]ae5 @frffi
Fffi) asoog AouS rassaT eq? Jo saTpnls ^drolslq aJTT
' (eeOf ) 
..f ,a>1oo3 pue . S.C ,delputg , .J.U ,11aau1cog
'992-e?,2 
.dd .)t.rotr r'raN ,ssa:6
cTurapecg 
. 
(.daf{cng 
.d pup a{ooC .,{ :srolTpg) scTlauag
GFv uf 
. 
suoT?eTndod Jo a:nlcnrXs c11aua6 aql pue
AoTJ auag 
. 
(eAOt) 
..{.e ,q6nolclroJJpg pue ..{.U ,11aaanco6
. 
L6-rG ,rrt ,lmE:nm ft5TFW Affi
'(suacsaTnJaec suacsaruaec Jesuv) asoog rtous rassar ar{l
:sp.rTq clqd:otulp 6u11sau dlleluoToc uT uolleldepe TpcoT
pup r{oTJ auag 
. 
(ttn) 
..{ ,a>1oo3 pue ..{.U ,11aau1cog
'9T9T-909T 'E3-
'@g- ;uol1sa6Tp .ra11eu .f,:p u1 uoTlcnper :slupuTunr
1su1e6e s1ue1d Furpuagiap uT suruupl Jo aToU 
. 
(eeef)
'V'& 'r{a1ueg pue 'g'V 'ueu:e6eg ,.S ,aTon ,.,tr.C ,sulqqog
'ETZ-89T'bT,@q
'r[:1s1ueqc a.rolrqraqTlup 1ue1d ;o rt:oaq1
'Grct) 
.g.U ,sa1e3 pue .d.CI ,sapeoqg
sacue^pv
1e:aua6
luaCaU
e pJeAotrJ
902
,,6T-LLT,E,@
F AaT^au fuuf 
.sua1s^s palse:oJ uT r(:oa1q:ag . (gget)
'r1' 'V'q 'r(a1sso:C puB 'I\t'M 'aao:6:eg "q.,1 ':allenoqcg
'6-r '?m.'Em
s{cTrlc asno.rg pau Jo pooJ
aq,l 'sncTlocs 'T
aq& '(trct) 'r'c
sn?Io6Pii
'r[:oaes
'ppeue3 'o1uo:o;, ;o d11s:alTun 'sTsar{& 'cS'I{
Fue suoTaere ffi tfffiE 6 6Efze.rg
ffiog Aous ffi Jo slcaJug ffi '(eaef ) 'v'g 'Tnpes
'pallTuqns '(u11:ag) eT6oTocao 'qsreu lTEs cTlcreqns
p uT sjtossaco:d ua6o:1Tu se asaag AouS jrassaT Jo aTor
aq[ 
. 
(Neef ) 
.t.U ,sa1:a;graf pue .s.q ,{TH ,.U-U ,sseng
' T,Z-E?Z'-69' (u11:ag)
E-i6-o'To- 'a6pes uecT.:rJV lspg ue Jo a>1e1dn ua6o:11u
pue sasuodsa: qlno.rb aql uo uoTle.rluacuoc ua6o:11u
pue aornos ua6o:11u '6u1dd11c Jo sloaJJa aqfi 
. 
(eeef)
'g'n ':nouaq6noC pup "f 'S 'uo1q6ne5c11 '.U.U 'ssang
'pa1lTurqns, (uTT.rag)
E-i6-oT6G[ 'spueTsse:6 ueclrJv lspg uro:J ssoT luaT.r1nu uo
s1eurueu 6u1ze:6 a6:e1 Jo slcaJJa aq1 pue uorlezrTTleTol
eTuoluurg '(ggOf ) 
.C.S 'uolq6neNcl^I pup .M.U ,ssang
. 
OTT
-ToT '-6v 'S5iTo- 
.spueTsse.r6 11a6ua:as aq1 uT sassaco:d
TprqorcTur palpTn6a: .,{6:aua pup luarJlnu Jo soTureulp ar{X
pue 6u1ze:9 
. 
(feet) 
.C.S 'uolq6neNchl pue .l{.U ,ssang
'LgE-TEe '6 '(u11:ag) €-Tfficao
'suoTlcpJaluT a.ronTq:at{-?ueTd gio :aldnoc alTloruo:d
e se parn '(fget) .n.S 'uolq6neNcl^I pue .l{.U ,ssang
902
.650r-860r ,w, ,Ei5{. i
'1rasap e Jo 6u1:aqlean aql pue s{cor uT .(rorrlq:ag
'(feOf ) 'f '1oue:g pue 'g'C 'sauo1, ,.r{ ,leqceqg
'rE-Ev 'd, '@saE frTffi FiiE'- ft 
.epeuea
'eqo11uep '111qc:nq3 'au11aa:1 aq1 1e -sa6ueqc aleurTTc
asuodsa: pue suoTlelndod acn;rds alTqr*r Jo XuauqsTfqelsg
'(feOt) 'q 'a1.deg pue ,.C.I.U ,lasueg ,.V.d ,11ocg
'vzT-6IT 'TZ. 
, (u11:ag) eT@ .ale.r 6ulssaco.rd
pue d4llTqeTTe^p pooJ Jo saTor ar{1 .roJ suoTlecTTdurT
:s6u11so6 asoog epeueC 6u11>1cec Jo rnoTler{aq
6u16e.rog 
. 
(eeet) 
.C.O ,6u11aae6 pue .S.f ,:a6u1pag
'zorr-e 80T 'E ,X6_Fcg
ffi to tBurnor 
.s1ue1d pooJ rTaql Jo dlTfTqpTTp^e
pue d6olouaqd aql 01 uoTleTal uT asaag eppueC ^dq 6u11sau
go 6u1tu1,t '(ge6t) 'e 'O '6u11a,reg pue .s.1' ,ra6urpag
'908-962
'm ')tnv aq[, 
.asea6 6u11>1cec ;o s6u11so6 :og pooJ Jo
.(111enb pue dlllTqeTTe^e pooJ 01 uoTleTar uT r(1ra11ca1as
drelaTo '(taor) 'e 'o '6u11aaeu pue 
.s.1' ,:a6urpag
'gvz-Enz 'E' 'Etm; 'sTapour
lenldacuoc pue leclqde:6 racuapraa 1ec1:adrua :.d.roa1q:aq
a6e11ogr r(q sleurue TTos Jo saTlTsuap Jo uoTlpzrllrTxel^I
'(eeef ) 'C'q 'sadeg pue "V'U 'opunueg ,.U.1, ,1pa1seag
'ppeup3 /ene119 'sacualcg TernleN Jo umasnll
TEuoTleN 
.epeuec JoeJoTdeqL '(eeOt) 'f 'g 'uB66ocs
'6t-s? 'E 'uoT+fttonE FF 76-oTo-cs
r{q pacnpur sasuodsa: lueTd
uT spua.r& 'saJo^Tqraq
'(eeef ) 
.C.1' ,z11nqcg
LOZ
'OT?-EOn '6 '(u11:ag) eT6o-1o- asareq aoqsnous
Jo sacua:agia:d 6ulpaagr auTuualap spunoduoc r(:epuocas
1ue1d oq '(feOt) 'n'N'1' 'q11urg pue 'E'U'V '.rte1cu1s
'o6ec1q3 'ssar6 obecgq3 ;o r(11s.ra^Tun
aq& 'uralsiffs ue Jo sc-Tur-EE7fi TTT€@ ' g.rc;')
'(s:o11pg) 'n 'sqlTJJTre-uoXroN pup 'g'U'V 'rrpTcuTg
'obec1q3 'ssaJ6 o6eclqp g:o r(11s:alTun
'oTeffi' 
- 
5E '(trct) 's'u'v '.rrefcuTs
'6gz-egz '-8e 'E-ofm; 's:aze:6 pue sasser6
uaaAlaq usTTenlnu penTola oN '(Zeffi) 'U'f 'uarolranl;S
'uopuoT
'11an1ce1g 'qcriffitf€ X:- 1tr
Zb-dTo-cs TEc-T6-oF-|SXE ' (gee r) 'a 'no1e3
uv :sTeuTuv Jo
pue 'n'u 'ITqTs
'uopuoT'11an1ce1g' (no1e3
'd pue puasu/y\o[ 'u'c s.ro11pg) asn acrnoffi 01 q5po=il€
7ft-Ti:ref6 W :ffi'TecT66.rrcTFffi ur 
.uoTlecaJaq
pue uoT1sa61o go sa16a1e:19 
. 
(reor) 
-n-u ,,tTqTS
'9L9-299
'-Tg,m.rI}rto:6pueuoT1eTnunccE1uaTJ1nu:e.Ipun1
ue{spTv ue uT slluroJ qry"ro.r6 1ue1d snoT:rl ^f,q uorlezTTTl:aJ
o1 asuodsag 
. 
(OeOf ) 
.fff .S..,I ,urdeq3 pue .U.g ,:aaeqg
' 60 t-T o ? 'X '7ffiTo-cg ' PXseTv
'r*rotrreg 'utalsr(soca eJpunl laA e uT .ralourn? loo: pue
uoTlcnpo.rd loog 
. 
(StAt) 
.q.l{ ,s6ug11rg pup .U.9 ,:aaeqg
802
oz-r 'Tz '@-1e FTuatiT:Sffi =€ [1coE'
ar41 Jo ETEoTuilE 
.suoTlersosse c1urs1ue6:o 6uldgTsseTc
:oJ auraqcs pazTTe.raua6 V 
. 
(SfOt) 
.d.n ,::els
'vot-TL 'fi, '766T--s Jo @of
'.ro1ceJ cTloTq aql 01 uoTleTa.r uT saddloca (.t e+::r
sTfIffi') sse:6 ?ooJs)tcoc 
. 
GZar) 
.C.U ,uopaldelg
'a6p1.rque3'ssaJ6 z{11s:aa1ug abpl:que3
'-ft6 EeEom aq? Jo 76Tcg aq[, '(reor) 
.f .r ,1ua.rdg
'T6€-gEC'fr
'F5FG-is-F$ Fup7ffiTocfiF- ffi@ 
.saTlTunururoo
fernlpu ur acueqtnlsTp Jo aToJ ar{;, 
. 
(leef ) 
.a.m ,esnog
'TIB-66L 'fr, 'Xili1EffiIiffi; pup XFT
'poqlau alrroTqcod^dq-louaqd aq1 z(q s:a1ea Ternlpu
uT eTuoulup Jo uoTleuru.ralaq 
. 
(egOf ) 
.t ,ouez:o1og
'ocsTcueJd ups 
,.duBdruoC pue ueuaa:J .H.M - (uoTaTpg
puocag) @'TE 
- 
(reot ) 
.c..r ,3r1qog pue -,!I.u , Te{os
'oeT-r sr 'fr ,FqdEJEb@ TE5TEG' . sua1s,{s
Te.rnlfncl:6e u1 ,{qde:6otuap lueTd 
. 
(Oeef ) 
.U.U ,uoprIBus
'9oT-86 '-Zg
'trb-6- 'sar{s:eu lTes Telseoc uo asaae AouS fiq 6ulzezb
Jo slcaJJa aq,t 
. 
(teOt) 
.9.$ ,unpg pue ,III .f .,1 ,rI?TurS
'orz-voz 'o '76@E
cTlcjteToI{
1ue1d qs.rpur
'asaab Aous 6urze:6 .dq uolllsodruoc z(llunruruoc
lfes Jo uoTlpralTg '(eeef ) 'fff 'f 
.,t 'rIlTruS
602
')trotr r'raN 'TTTH-ne:gcI{ 
. 
uoTlTpa puocas ,TEffi56E@fr
5ffi '(ggOr) 
.q.V ,q11rug pue .V.T ,1:eppolg
'LZ-'t 'dd 'proJxg '11aaqce1g 
. 
(uos1:og 
.H.I
pue TTTaNcr{ 's 'aaT 'V. f : srolTpg) .roflceg TE-c]b6-ffi
ue se u5-6o:alrq ur 
.6u11cIc ua6o:11g . (e eot)
'd '11anog pue "N'V 'TeU ",1 'uo1sa.l6 '.d.q.A ,1:ena1g
. 
L?,7-TTZ 
.dd .proJxg ,11aau1ce1g
'(rVteq '1''V pue saT:eJJaf 'T'U :sro?Tpg) S@ug
@5 ET sessecord F-frEotoffi ur 
. 
salf,qdoleq 1ue1d
:aq61q u1 acup:aTo? 1Tes pue usTToqelau ua6o.r1TH 
- 
(atoil
'V'f 'aaT pue 'f 'peuqv "g ':aq:e'1 "U'g 'l:Br'ra1s
'svg-6Eg '-ZL 'TFT5-oJ@ AaN 
.uoTlpzTTTln
pue dlllTqeTTp^e a?er1TN 
.II sa4.f,qdo1eq Jo ursrToqeaau
ua6o:1111 '(grct) 
.oCrueqe:g pue .V.f ,aaT ,.U-9 ,X:enals
'Bo9-909 'ffi,' '- i 
. 
suoTleTndod 1e61e uo sAcaJJa
lca.rTpuT pue lcarTp ,saJolrq.raH 
. 
(geOt) 
.U.U ,:au:a1g
'uolacurJd'ssa:6 .d11s:err1u1 uolacuTrd
'Foaq[, ffiTffi{ 
. 
(geot) 
.u.f ,sqa:y pue .A.a ,suaqdalg
'€9T-Z9T ,fr ,-'tTO-
'r(6o1oca u1 bulaer"rpueq lnoqp luaururoc p :uoTlnTo^aoc
pup rusrTenlnlu ,sJazp:6 ,sasse:g . (ggOf ) .C.H ,q1asua1g
'9oE-662 'E 'E6TfEi assaulTJ 1ue1d
TenpT^Tpur azTurTxeu s.reze:6 oq 
. 
(efef ) 
.C.H ,q1asua1g
'g8-9L 'E3.
,@GtTPCTUelogJo sTeuuv 'saro^Tqraq
(reor) 'r'c 'su1qqa1g
T.rnossTN artl
uoTlnTo^aoc
oIz
pue sasse:6 3lo
'Tgt-Lg? 
.dd .{rotr AaN ,6e1.ra6-6up:dg . (uezsaT,tr .T.T
::o11pg) erpunl crlcre ffi up Jo @ 6T1:_nF6Td
Fue uo$+ei6--11 uf 
.q1r"ro:b luetd Erpunl Jo suoTle?TurTf
luar:1nu lueTd 
. 
(ef Ot) 
.f .a ,:ads:ag pue .V ,r{cT:Tn
. 
e TT-ooT 
,E ,EETFE:Efr . slrun 1ec16o1oca
se r(1ar:el pue satcads eqg, 
. 
QZA7) 
-e ,uossa.rng,
'Ppeuec
'r[.re61e3 'a1n1r1su1 TpaJog ar{J ,:a11ng .V.t{ trq pa+Tpg
'd::eg '$'1, pup JJo)tessI 
.g ,{q peleTsupJ,L .perbuluaT pue
r'rocsohl '.U.S.S.g aql Io acuaTcs ;o ^f,uapecg aq,l ,e1n111su1
TPCTuelog aq& 
.eJpunJ ar{1 Jo Ja^oc 
"ueTd 
aq? pue ptroM
Te@rV- artt Jo 
- 
s6o-TFT6! 
. 
(osor) 
.v.s ,AorTuroqxT,t
'?TT-e TT '-6t 'so-m 2bulzetb uro:J lTJauaq
sasse.r6 oO 
. 
(ZeOt) 
.e .n ,da111g pue .X ,uosduoqg,
'e9e-6Ee ,G , (uTTrag) pffot;aO .asaa6 6urpaa.rq
-u:ar{lirou Jo uorlr.rlnu eql uT (aeacelasTnbg) sTTplasJor{
Jo aTor aq; 
. 
(Zeffi) 
.a.f ,11ana:6 pue .9.A ,seuoq,1,
lo teu:nof 
.treg
TeuoTlT.rlnu aqtr'
.9e8-os8 ,+uffiA6Eu-Blt elTtprTlt
sauef 1e esaa6 01 sasse.r6-no::e Jo anTel
'(OeOf ) 
.a.f ,11arra:6 pue -g.A ,seuoq;,
'6TT-60T 'TI ,EEE . eas
uappe& aql uT 6u16e1s 6u1:ds 6u1:np saTlTunl:oddo Fulpaagr
TPnpT^TpuT
' (seor )
o1
'u
uoT?eTaJ uT luaJg uT ssaoons 6u1paa:g
'1ua.rq pue ,.9 ,sueBds ,.M ,uasstunag,
'nzg-vIg '& 'm 'e16:oa9 ;o rualsdsoca
qsreur lTEs er{1 uT AoTJ ^f,6:aug - (ZgOt) .n.f ,Tea,tr
ITZ
.Zes-o7g 'TL , (u11:ag) eT6'To5ao
'6utzB:6 Jo slcaJ;a alqrssod :slpaqn palecrlsauop
uT sw.roJ qlrrro:6 lcaJa Jo uoTlnTo^g 
. 
(reer) 
.l ,1as1e6
'6ZE-OZ,e 'bE 'EeueTE
sfrToaser{d Jo saleeT Jo .fiq1cedec cllaqludsoloqd aq1
uo (ZOC)d pacuequa 1e q1r,ro:6 pup ssa.rls xaleA nrra?-?roqs
'q1no:6 6u1:np acuaTper.rT uT se6ueqc ,uol1e1ToJap 1eg1:ed
Jo slcaJJg 
. 
(teOf ) 
.q.C ,:eqnb:eg pue .S ,trarauruapJ uol
'862-982
,F,@.s1saJoJ1ec1do:1u1uoT1e+TuITT1uaTI1nu
pue 6u11cdc 1ue1.rlnu ,TTpJrallTT - (leer) .n.a ,>1asno11n
'?Tg-LoE '6'trFcg
pa-TTTffi Jo TeurnJof 
.pupTssp.r6 a1e:adural € Jo uollcnpo:d
I:eu;:d 1eu pue 6utze.rg 
. 
(Zrct) 
.f .a ,r[:a>1c14
69T-89T 'T 
,6'.!ffi7E
's1ue1d lTJauaq 6urze:6 saop uar{M
FG ffiog ur spualf,
. 
(geOf ) 
.C.g ,:ee>1:a4
'e 9e -9Ee 'E ,E5m . sarolTqrar{ lsure6e a166u1s
aql uT sa16a1e:1s Xueld anTleu.ralTe ,qlruor6al pue acuaJaq
'(eaet) 'r'u ':ee>1:an pue 
.n ,u[Tu ,.g ,uap[Tan .rap uel
'Egg-Lng 'dd 
.)t:ol r'raN
'ssar6 cTurappcv 
. 
(uaane 
.H.t{ pue pTourrag -1' .lI :s:o11pg)
saEet[5'TeH go X6@E ur 
.uoT1p1a6aa qs:eu XTes uT
uoTlelTulTT luaTrlnN 
. 
ftrct) 
.n.f ,1eag, pue .I ,e1a11e4
zTz
'F5FEu5IE-GI pue X6o-rccs Jo AaT^au
1ea:6 aq1 pup ecTJauV q?nos 
.II
uT sale-rqalJal puuenes g:o d.rolsTq V
' ogzv-t 6e '6
Teruruv 'a6ueqc:aluT
ued 'pTJoirr itlau aq+
'(aear) 'q's 'qqa^r
enuue eod uT
.d6o1ocaua6
. 
osc-Esc 
,E' ,sc1TE@E7Gi pup Z5otocs
F AET^au Tenuuv 
. 
poTtaurv qlJoN 
. 
I 1:ed 
. 
pT:oA r-rau aql
uT sale.rqelral euuelps 3lo .d.ro1sTr{ v 
. 
(ttat) 
-o.s ,qqa6
'LEL-gzL 'T8 '1sT6-oJ@ AaN 
.spaq raAoTJ
pup sunel 6utllroq Jo luauuoJTnua oTesoul p ur .'f eruue eoa
uT uorlsaTas arrrldn:sTp JoJ acuapT^g 
.II .spaaA uAeT Jo
d6olocaua6 aq; 
. 
(qef Ot ) 
.A , s661:g pue .I. S ,>1crn:e1,1
.C7L-ITL ,TE- ,lsTSoToiXrId r"raN .spaq
Jar"roTJ pue suApT 6ul1rroq Jo luauuo.rrlua cresou e uT 
.T
uorlprlua:aJJTp uotlelndo6 
.f .spaan ur\e1 Jo
aq;, '(egZet) 
.O ,s66r:g pue .I.S ,>1c1aure6
'oEz
-zzz 'TL 
,@ Jo TeuJno1 ue- .sprourure.r6 uecl:;lg
lspg oA1 Jo uoTlcnpo.rd pue ,q1r"ro.r6 ,a6ueqcxa se6 aq1 uo
ua6o:1ru g:o sTaAaT rnoJ pue 6urdd11c gro slcaJJg . (geOf )
'g'n ':nouaq6noC pue "C'S 'uo1q6negc6 ,.T.T ,ace11e6
'gLz-zLz 'a
'(u11:ag) eT@,cao 
.saur6a: uoTlpzTTTUaJ pue 6urddTt"
luaraJJTp .rapun ,sarcads sse:6 VC e ,.T urnleroToc
ulncTued TEzTr{J.roc^dtuuou pue TezTqr-roc.dru go a6ueqcxa
se6 pue '.d6o1oqd.roru ,q1r"ro.r9 . (f eOt ) .t.t ,ace11e6
etz
oTuaI^I's6ururun3/u1rue[uag 
.s-!@5 pup
;io uoTlcaTes Te.rnleN art[ 
. 
(OeOf )
'o9er-8EeT 'm,'ffi-FE
Jo TaAaT ar.{1 uo uor1nTong 
. 
(gfef )
'XEbf- Jo Teu:nor snua6 aq1
ualas uT uorlnTola .d.ro1stq aJTT 
. 
Grct)
'vc '>[:Bd
ffiTi€T=n-6;i
. 
S.q 
,uosTTl{
'saTlTUnUnUOC
. 
S.O 
,uosf Tu
'onz-ezz '6
JO spaaJD[Trur
'r{'H':nqTTl4
6?eT-6eeT 'EE 
,- 
icT+EffiE
FE ffi 3lo Teurnof @EEJ 'eTlocs eAoN Jo lspoc
cTlupTlv ar{1 uo 'siuei@ snJeuroH ':a1sqo.1 upcTJauv aq?
10 acuepunqe aq1 pue'@ffi
'urqc:n eas aql Iq bulzezb alTlcn.rlsap uaaA?aq
dlqsuolleTag 
. 
(f eet) 
.tt.X ,uue4 pue .C.lt ,uo1:eqtr4
'r?6-ov6 'Tr
,lsTTe.In1eT
.IoJ ssaccns
'r,I '.f,qo1sair1
uecTJauv aqJ, 
.seJo^Tq:ar{ pTTA pue {colsa^TT
6u1ze:6 6utcuan13iu1 srusTuer{cahl 
. 
(geof )
'Tt8-Ot8 'm
'ilF!ftEnFF umTJaruv ar{& ls.raze.rb :ogl acrnosal pooJ aql
aao:durt .d11ensn 6u1ze-r6 .{neaq saog 
. 
(geet) 
.W ,dqolsa6
'698-e98 'Eg 'X6[6ffi Jo leurno,f 
.6u1dd11o :atJe s.raTfTl
pso=aqn1 
- lo Te^TA.irnS :sclrueudp uoT1p1ndod
raTTTl pup 1aua6 uaaarleq suorleTa5 
. 
(OAef ) 
.n ,r(qo1sair1
'g?,t-Lle 'Z ,sGffii rntpg Jo
Teurnof CTpffi 
.epeuec ,deg uospnH u.ra?seaqlnos ,er:ery
ellaT.ruaH adec 1e a6e alprlsqns pue AJTTdn 1e1ce161so6
'(OfOf ) 'A'C 'sata:pu11 pue 
.M.f ,uosp.reqcTg ,.f .d ,:aqqag
vTz
.eoE-?,62 .dd .>trotr itlaN
'ssar6 cTuapecg 
. 
(TTeXcn 
.W.C pue :aEunotr .g.A :s:o11pg)
sffi' F 6TF4im F-ue I6oT6Tfi aE ur .q1no:6a:
pue uoT?eTToJap gro .f,6o1oTs,{qa . (Zrct) .e.A ,:a6uno;
'Esv-Etv 'ET 'tr3{o-cs FSfffdE 3o p6rnof
'asaa6 aTceu.req dq d111enb poog: Jo uorlelndrueu aql pue
6utze:6 6u1:dg 
. 
(f eOf ) 
.A.H.H ,su1:6 pue .O.U ,6:aquaptr
'GTz-ET.z 'E 'lKb6T- Jo Teurnof ueTF:m5 .or.re1uo ,r(eg
saupf 1e 6u16e1s esaa6 Aous Jassar Jo sctueu.f,p a^Jasa.r
luaT:?nN '(AtAt) 
.O.C ,.f,au:1uq pue .d.C.U ,etualdr(tr1
'gzvT-ozvT '6 ,rcTo-s 5 @oc uEFEE$
.pcrJauv q?JoN u.ralseaql:ou uT (aecacelpodouaq3)
( ' t 's ) e-aedorna Effio5ffi pToTdTp Jo snlp?s
cTulouoxpl aq;' 
. 
(aAOt) 
.t.U ,se1:ag:g:af pue .S ,JJToU
.{Jotr rqaN ,ssa:6 unuaTd
'TAoJraletrl ptll,t Jo @ ' (teer) .v.e ,:asaqolq
'a6pr:qure3 ,ssa:4 ,{11s:arrru61 a6p1:qupC . (.U.a ,aue:p
pue 
.g.ll ,:auoleqg ,.n.g ursrrJd :s.:rolTpg) sacudnffi_c
E5TqoT-q -FaE pup s*ra@ffi Jo EuTET-o artr,
uf 'aurT1 q6no:q1 spode:1e1 sno.ro^Tqraq pup suradsolFue
Jo suoTlcp:a1uI 
. 
(f eOt) 
.H.e ,^6au;g1& pue .T. S ,6uTlt
'o8s-TLE 'Z ,acuaEi F to s-a-rffi Fuo-tl@
m a'q1 Jo suo-T1GEET .sarlrado:d TTos uo 6utze:6
Jo pue 6u1ze:6 :o3 tr+TtTqe+Tns uo sa11:ado:d lros go
acuanTJuT aqfi 
. 
(tgAt) 
.f .C ,1s:oqloqcg pue .d.9 ,puTU
9TZ