ABSTRACT

EARLY DEVELOPMENT OF BEHAVIOUR PATTERNS
IN GROUP AND ISOLATION REARED BOBWHITE
QUAIL (COLINUS VIRGIANIUS)

BY

Mary Lee Nitschke

Results of studies on socially reared and isolated
birds in species closely related to the bobwhite quail
suggested that the isolate birds develop the same patterns
evidenced in the socially reared birds, but show these
later in some instances and with greater variation in the
pattern and sequence of development. The present study is
an attempt to (1) provide a baseline of early develOpment of
the behaviour of the bobwhite, a non-domestic precocial
galliform, under laboratory controlled conditions with
minimal human contact or intervention and under a natural
lighting regime; (2) extend previous findings on the
effects of isolation with the galliforme birds by observing
a new species having a longer period of develOpment before
reaching maturity than those previously studied; and (3) to
describe components of behaviour classes not previously

reported in eXperimental studies.

Mary Lee Nitschke

During the first 29 days post-hatch the isolate
birds were housed individually under tactual and visual
isolation conditions. From day 30 on the isolate birds
were combined into a social living unit. The socially
reared birds were housed communally from the outset. All
birds were maintained on ad lib water and food on a
natural diurnal light cycle, virtually without human
contact. Observations were made from a darkened, adjoining
room and data recorded electromechanically and electro-
magnetically.

The results obtained extend and support previous
findings on the effect of isolation. In addition, this
set of observations provided descriptions of the following
behaviours which have been previously unreported for this
Species reared in the laboratory: (1) distinct forms of
Care of the Body Surface components, (2) components of the
dusting pattern occurring on wire, (3) the age of onset of
crouching and "cautious" postural patterns, (4) the
Cubicle Pacing and Sustained Tactual Contacting patterns
as a persistent behaviour of the isolated subjects, (5) an
attempt to characterize the synchronous versus the frag-
mented nature of the topography of the behaviour patterns
as they differed for the two groups, and (6) the progression
and development of components of the "roosting disk"
Resting Group Structure, a species-Specific pattern for

which the bobwhite quail is famous.

Mary Lee Nitschke

In conclusion, the results of the present study,
taken in conjunction with those of other studies on the
galliformes, indicate that generally isolation may retard
but does not prevent the develOpment of social behaviours.
One exception was found, the Ring Nesting Resting Group
Structure never appeared in the isolate group even after

having been combined and living together 45 days.

EARLY DEVELOPMENT OF BEHAVIOUR PATTERNS
IN GROUP AND ISOLATION REARED BOBWHITE

QUAIL (COLINUS VIRGIANIUS)

 

BY

Mary Lee Nitschke

A THESIS

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

MASTER OF ARTS
Department of Psychology

1971

AC KNOWLEDGMEN'I‘ S

The author wishes to thank Dr. Stanley C. Ratner
for his patient guidance and direction of this thesis.
I would also like to express my appreciation to Dr.
Terrence M. Allen and Dr. Robert K. Ringer for their
technical assistance and the enthusiastic encouragement
they provided for my work in this area.

Several of my peers assisted as extra observers
during the study. Gratitude for this goes to Suzanne
Power Marshall, Peter L. Borchelt, Robert E. Otis, and
John Rosenketter.

Finally, this thesis is dedicated to Rusty who

taught me the most about Bobwhite quail.

ii

TABLE OF CONTENTS

ACKNOWLEDGMENTS . . . . . . .

LIST OF TABLES . . . . . . . .

LIST OF FIGURES . . . . . . .

INTRODUCTION . . . . . . . .

METHOD 0 O O O O O O O I C

General Organization . .
Subjects
Apparatus

Phase I . .
Subjects .
Apparatus .

Procedure . . . .

O O O O O
O
O
O O O O O
;

Phase II: Procedure
Phase III: Procedure

RESULTS 0 O O O I O O O O 0

Definitions of Behaviour Classes .
Outline of Behaviour Classes . .
Measurement . . . . . . .
Classes Common to Both Groups .
Classes Unique to Isolate Group .
Classes Unique to Social Group .

Descriptions of Behaviour . . .
Behaviour of the Social Group .
Behaviour of the Isolate Subjects

Comparisons Between Social and Isolate Groups

DISCUSSION . . . . . . . . .
Behaviour of the Social Group . .
Behaviour of the Isolate Subjects .
Summary . . . . . . . . .

BIBLIOGRAPHY . . . . . . . .

APPENDIX C O O O O O O O O O

Page
ii

iv

15

15
16
19
21
21
22
22
26
27

3O

31
31
32
32
37
39
40
4O
69
91

105
105
109
112
114

117

Table

1.

LIST OF TABLES

Page

Behaviour classes with respective response

patterns of social and isolate groups

showing age of onset (lst occurrence),

median onset age, and day the behaviour

had been shown at least once by each group

member (100% onset day) . . . . . . . 42

Range and median frequency of allOpecking

encounters of a series of observations for
each week of age in a group of socially
reared bobwhite quail . . . . . . . 68

Frequency of allOpecking encounters observed

in the isolate reared group (N=7) on the
day of assembly into a social living unit . 90

iv

LIST OF FIGURES

Figure Page
1. Design sketch of basic living unit . . . . 20

2. Onset of (1) general preening; (2) oil gland
preening . . . . . . . . . . . . 45

3. Mean frequency per day of general preening
bouts in Social and Isolate groups over
a 4 week period . . . . . . . . . 46

4. Mean frequency per day of Bill Wiping in
Social and Isolate groups over a 4 week
periOd O O O O O O O O O O O 0 48

5. Onset of AlIOpreening in Social and Isolate
groups . . . . . . . . . . . . 51

6. Onset of Flying in Social and Isolate groups 53

7. Initial appearance of wing quills in Social
and Isolate groups . . . . . . . . 54

8. Onset and duration of predominate structures
of group resting patterns observed in the
Social and Isolate groups throughout the

StUdy O O O O O O I O O O O O O 5 7
9. Onset of freezing in social and isolate
reared groups of bobwhite quail . . . . 60
10. Onset of the Cautious Alert Posture . . . 63
ll. Onset of Crouching in Social and Isolate
groups . . . . . . . . . . . . 65
12. Onset of Running in Social and Isolate groups 75
13. Onset of Jumping in Social and Isolate groups 76

14. Onset of Frolic with object Locomotion
pattern in Social and Isolate groups . . 80

Figure

15.

16.

17.

18.

19.

20.

21.

22.

23.

24.

Mean frequency per day of incidence of
Sustained Tactual Contacting in seven
Isolate subjects over 4 weeks of
isolation . . . . . . . . . . .

Mean frequency per day of three rates of
Cubicle Pacing by Isolate group subjects
over 4 weeks of isolation . . . . . .

Onset of Sophisticated Preening in Social
and Isolate groups . . . . . . . .

Onset of Bill Wiping in Social and Isolate
groups 0 O O O O O O O O O O O

Onset of Dust-Bathing in Social and Isolate
groups . . . . . . . . . . . .

Onset of Popping in Social and Isolate groups

Onset of Flit-pOpping in Social and Isolate
groups I O O O O O O O O O O O

Onset of Generalized Pecking in Social and
Isolate groups . . . . . . . . . .

Onset of Frolic Locomotion pattern in Social
and Isolate groups . . . . . . . .

Initial appearance of tail feather shafts in
Socially reared and Isolation reared groups

vi

Page

82

83

93

94

95

96

97

98

100

102

INTRODUCTION

Natural history and semi-technical observational
reports on the bobwhite quail recommend it as an excellent
preparation for the study of many of the major classes of
consummatory behaviours. Stoddard's (1931) monograph
stands to date as the classic source of background informa-
tion on the behavior of the bobwhite quail. In addition to
being one of the most pOpular of the upland game birds of
widespread distribution throughout the United States, its
varied and distinctive behavioural repertoire offers a
challenge to our understanding of the behaviour of a non-
domestic member of the galliformes. As a precocial bird
it is also an excellent laboratory animal and is now
beginning to be utilized as a subject in eXperimental
behavioural investigations. The lack of behaviour work
with this animal to date is surprising since this species
has many attributes that would recommend it for behaviour
research, in addition to its being a hardy and easily
handled animal when reared in groups and requiring
minimal maintenance and space.

As a good preparation for studying several classes
of behaviour in a social species the bobwhite exhibits

1

powerful examples of behaviour classes that have received
little intensive attention from experimental investigators.
In the present instance the classes of locomotion, care of
the body surface, resting, and predator defense (behaviour
of the prey animal) are primary examples. Although not
specifically considered here, the bobwhite also exhibits
clear and often elaborate examples of the classes of
sexual behaviour, care of the young, nesting, and intra-
specific communication possessing representatives of each
of the classes of calls named by Collias (1960). Thus,
smithin the aves, the bobwhite could profitably be investi-
gerted as a species that offers an integrated picture of a
nwrjority of the consummatory classes of behaviour (with
thxa possible exception of fighting) as listed by Ratner

(Deanny and Ratner, 1970).

The bobwhite quail is a small variegated brown and
budff colored precocial galliforme, native to the United
Startes. The chick hatches into the company of 7-15 other
tier "feathered bombshells" (Bent, 1963) each weighing on
th£3 average of 6 grams and covered with natal down. At
orue day of age the brood begins foraging eXpeditions under
tine direction of both parents. For the next two weeks or
SC) the chicks spend 75% of their time being brooded by one
Of the parents until they become thermoregulatory and begin
acquiring juvenile plumage. The daily activity of the

family for the next several weeks over the summer consists

primarily of an active early morning foraging period
followed by bouts of maintenance activities such as
preening, dusting and resting, until a few hours before
sunset when another active feeding period begins and con-
tinues until the group gathers together for the night
(Stoddard, 1931).

One of the striking aspects of a quail family or a
quail covey (a collection of quail larger than the family)
is their highly integrated group behaviour. The group
appears to act in concert as a unit, particularly with
regard to daily activity and predator defense patterns.

A typical observation made by Stoddard (p. 18) illustrates
this point:
. . . every little while some imaginary alarm
causes every bird to freeze and remain absolutely
motionless for periods of seconds or minutes,
when suddenly, as if at a given signal . . . all
relax and go about their business.

The information on development of behaviour patterns
in the bobwhite chick is sketchy. Stoddard (1931) mentions
that the first week the chicks jump and make flying leaps
in pursuit of insects. At two weeks chicks will "flush;
when disturbed. He describes this as: "All members
freeze to the ground with feet firmly placed, only to
spring into the air with a thunderous roar . . . ." This
perhaps is the origin of the title "feathered bombshell."

He noted a group of quail hatched in captivity and reared

with a brooder, never having been in contact with adult

quail, take up the circular roosting formation character-
istic of bobwhite at four weeks of age. This formation is
described as: ". . . heads out and bodies pressed so
tightly together that the tails are forced skyward, and

the sides are in close contact" (p. 45). At eight weeks
the chicks are entirely in juvenile plumage and the
sexually dimorphic coloration is evident. At fifteen weeks
their plumage is indistinguishable from that of the parents
without close visual inspection of the primaries.

Another source of general information on the
behaviour of the bobwhite is a more recent paper by Stokes
(1967). The book by Stoddard represents a conservation-
ist's orientation but contains careful description of
several of the behaviour patterns observed in the field,
with a focus on social and parental behaviours within the
covey. Stokes (1967) utilizes Stoddard's descriptions but
focuses his three year investigation on the function of
agonistic, sexual and parental behaviour patterns and
vocalizations of the mature animal. Although other recent
investigators (Garreffa, 1969) have also studied the
behaviour of this species, most have concentrated on a
specific experimental problem utilizing the mature bird and
minimal attention has been directed to the very young
animal and the early deve10pment of its behavioural

repertoire.

Recently, Borchelt (1970) provided initial experi-
mental information on the deve10pment of fear displays in
the bobwhite. This study suggests that responses to
predators appear sequentially as (1) crouching, (2) freez-
ing, and (3) immobility or fear display. Borchelt found
freezing first appeared in this context in 50% of his
birds between 10-19 days; with immobility appearing most
reliably at about the age (roughly two weeks) that the bird
begins to achieve physical independence from the nest.

Prior to this, the only deve10pmental data on fear
responses and a basic motor coordination in precocial birds
relative to other birds was provided by Nice (1962). She
proposed a stages-of-development scheme with motor
patterns changing throughout the following stages:

(1) post-embryonic, (2) preliminary, (3) transition,

(4) locomotory, and (5) socialization. However, definite
age limits for these stages and precisely defined criteria
for identifying the behaviours associated with the stages
were not provided; nor is this information available for
the bobwhite quail.

An initial step in the preparatory stages of the
comparative analysis of behaviour, as proposed by Ratner
(Denny and Ratner, 1970), is the consideration of the
question "what does this organism do?" He suggests that
an analysis and understanding of consummatory behaviour

requires the investigator to be familiar with the natural

history and the specific behavioural repertoire of the
animal being studied. Thus, identification and character-
ization of behaviour as a prerequisite for a powerful
functional classification system requires careful descrip-
tive work. When this body of information is not available
an eXploratory descriptive study is necessary to provide
baseline data before an investigator can legitimately
proceed to ask incisive questions with regard to choosing
powerful variables for an experimental analysis of classes
of behaviour.

It is generally recognized that species, age and
prior eXperience are powerful variables influencing the
process of behavioural deve10pment in a vertebrate
organism. In reviewing the comprehensive paper on social
behaviour in the mature bobwhite by Stokes (1967), the
question arises as to the origin of these behaviours and
how the various components fit into a classification of
patterns.

One technique for investigating the origin of
social behaviours is to restrict aspects of the social
experience of an animal and discern the effect of this
restriction of stimulation. An extensive literature
exists on the effects of isolation as a rearing experience
in a variety of social species. However, if we accept
that species is a powerful variable, we are obligated to

begin an investigation at that level of organization

before proceeding to other questions of variables assumed
to have a causal effect on behavioural processes.

Within the galliformes, a large body of literature
exists on the social behaviour of adult domestic fowl, as
evidenced by the reviews of Wood-Gush (1955) and Guhl
(1953). However, few studies have concentrated on the
development of behaviour from hatching and the few that
have were often aimed toward an analysis of a specific
behavioural pattern, e.g., sexual or aggressive. Thus,
there are few studies which provide the necessary data for
making direct comparisons between other birds and the
early deve10pment of the bobwhite. Those available for
comparison purposes are studies using domestic fowl (Guhl,
1958; Hartford, 1967; Ratner, 1965), or Japanese quail
(Farris, 1964). Although these researchers were primarily
.interested in aggressive or sexual behaviour, they
aapproached the problem from a developmental standpoint and
'their results provide some basis for looking at the present
{problem as part of a larger scheme.

Guhl (1958) raised various groups of chickens
(White Leghorns, White Rocks and commercial hybrids) in
Lbartial isolation from other chicks. His primary purpose
Vvas to determine whether partial isolation would influence
flhe age at which these chicks formed peck orders when
€Jrouped. He described the sequence of behaviour patterns

<>ccurring in his birds as: generalized pecking, escape

reaction, frolicking, sparring, aggressive pecking,
avoidance behaviours, and fighting. The isolation reared
birds formed a peck order after being assembled into a
group within short (8-48 hour) periods. Guhl concluded
that the early experience of such behaviours as frolicking,
pecking, and fighting were not essential to rapid formation
of social organization. However, observations were not
conducted on the behaviour of the chicks while they were
isolated. Therefore, as Ratner (1965) suggested, the
possibility exists that Guhl's isolates did perform some
of the behaviours in question prior to their assembly into
groups. One should also pay attention here to the type of
isolation employed in this instance, as a bird is a highly
visual animal. Partial isolation was defined as chicks
that could see other chicks but could not make tactual
contact with them. This partial isolation may have provided
considerable stimulation of an important nature during the
"isolate" rearing period for these chicks.

In elaborating the schedule of behaviour deve10p-
Inent reported by Guhl, Ratner (1965) attempted to deter-
Inine: (l) the sequence of deve10pment of social behaviours
in a normal group, and (2) the effects of isolation on
loehaviour development during and after a period of isola-
tion. He observed White Plymouth Rock and White Leghorn
cockerels reared together and chicks raised in visual

isolation from other chicks. His results confirmed Guhl's

observations of frolicking, sparring, aggressive and
generalized pecking, and fighting; the findings differed
only in age of behaviour onset. In addition, he provided
the initial description of frolicking with an object, food
dominate and food submit, and the juvenile fight. He

found that isolation led some behaviours to be retarded in
age of occurrence and that the patterns of frolic and
frolic with object were indeed exhibited by some isolate
birds during the isolate rearing condition. When the
isolated chicks were assembled at 70 days of age aggressive
interaction began almost immediately. Although the tOpog-
raphy of the pecking pattern appeared less organized in

the isolates, they formed a stable peck order within a few
days. Following assembly most birds then showed all of the
behaviours shown by the normal group, and the two groups
were indistinguishable in terms of social behaviour patterns
within several days.

Ratner defined the age of onset for a behaviour
pattern as the day the first bird in the group showed this
behaviour. It would perhaps be useful to extend this
information to look at onset of behaviour pattern as the
day at which at least 50% of the birds within a group show
a behaviour pattern, and to gain an idea of the variation
within a group of subjects by looking at the range between

initial day of onset and the day at which a particular

10

behaviour had been shown by every member of the group at
least once.

Farris (1964) studied the behaviour development of
young Japanese (Coturnix) quail under both social and
isolated rearing conditions and reported differences in
development in terms of day of onset between the two
groups on dashing, bill wiping, jumping, jumping, popping,
brief flight, hOpping, dusting, and cricket calling. He
reports "freezing" occurring on day one post-hatch in
response to "slight disturbances" and later evolving into
or combining with the "alert stance" on day 12. In general,
he found there were few differences between the social and
isolate birds in the course of behaviour deve10pment except
for sexual behaviour. The differences referred to above
are attributed to differences in the size of the home
environment for the two groups.

This brief review of isolation studies in the
galliformes indicates that the appearance of components
of social behaviour patterns generally may be retarded-
altered-delayed in subjects experiencing restricted or
subnormal stimulation, but rarely does this treatment
prevent the occurrence of the response components them-

selves.

11

Present Study

 

Methodological Considerations

 

In the studies cited above, several procedural
points should be called to the reader's attention.

1. Guhl (1953), Ratner (1965), and Hartford (1967)
all studied the domestic fowl. As stated by Ratner, ". . .
in general, domestic species represent poor preparations
for the purpose of uncovering behaviours that relate to
consummatory activities" (Denny and Ratner, 1970, p. 212).
Thus, one of the objectives of the present study was to
provide a descriptive baseline of the early deve10pment
of the bobwhite quail, a non-domestic precocial galli-
forme, under laboratory controlled conditions.

2. Other laboratory studies have employed a
variety of lighting regimes. Borchelt (1970), Hartford
(1967), Ratner (1965), Farris (1964) and Guhl (1953) all
employed a 24 hour continuous light procedure. Of the
above only the Borchelt study utilized a warm white type
of lighting. Since the amount of time a bird is exposed
to light is known to have significant effects as a stimulus
for the pituitary to release gonadotrOphins (Sturkie, 1965),
and since the lighting cycle has been shown to alter the
reproductive activity and general level of behavioural
activity in the bobwhite (Kirkpatrick, 1955), lighting is
a significant variable in the environment of this species.

Continuous lighting is thought to produce an agitated

12

state of behaviour in birds (Menaker, 1969). Other effects
of continuous lighting in the domestic fowl are an increase
in the size of the eye and diminished visual acuity when
subjected to continuous light over long periods of time
(Sturkie, 1965). The type of light to which the bird is
exposed, cool white versus warm white, also has been
demonstrated to have considerable effects on the repro-
ductive physiology and the neuro-endocrine system of the
fowl (Sturkie, 1965). Cool white light alters the repro-
ductive processes and due to its effect on gonadal and
thyroid hormones, Ringer suggests in Sturkie (1965) that
it influences the birds weight as well as the structure of
its feathers. Although data on these specific points are
not available for the bobwhite, the weight of the evidence
suggests that continuous lighting of a cool white type
would be a less than desirable choice of living conditions
to impose on a group of birds for a behavioural study.
Thus, the present study specifically inCluded as one of
its objectives looking at the behaviour of this galliforme
under a natural diurnal cycle of warm white simulated
daylight.

The third objective of the present study was to
provide an environment for the subjects which did not
include their being continually exposed to human inter-
‘vention or contact. The studies cited immediately above

all included conditions in which the bird could View the

13

eXperimenter while data was collected. One of the inten-
sively studied aspects of altered stimulation in early
experience studies with birds is the phenomenon of imprint-
ing (Denny and Ratner, 1970). Considering the possibility
that imprinting could occur if the bird is permitted visual
contact with the eXperimenter, this study was designed to
eliminate that particular possibility. In addition to
this, handling of the subjects was kept to an absolute
minimum.

A fourth objective in the present study was to
provide living space for the subjects that would prevent
crowding, and that would allow sufficient room for the
subjects to perform many of their natural behaviours.
Previous studies using social and isolated groups provided
only minimal living space for the subjects. Hartford
(1967), Ratner (1965), Farris (1964), and Guhl (1953) all
failed to provide cages large enough for the birds to fly
within the cage, and often only "commercially" sufficient
space was provided for the socially reared birds.

These same four investigators also obtained their
subjects from sources where there was no control of their
experience until up to 24 hours following hatching. Since
previous work has shown (Vince, 1966) that differential
pre-hatching conditions, together versus separated,
produces differential stimulation for the avian embryo,

this variable should be controlled in a study comparing

14

group reared with isolation reared subjects. As a fifth
objective, the present study attempted to impose the
differential treatment of groups prior to pipping of the
egg in order to more closely define the different rearing
conditions for the two groups of subjects.

As previously mentioned, the bird is a highly
auditory animal and this consideration suggests that
auditory stimulation should be treated as a variable.
However, both in the studies previously discussed and the
present study, this form of stimulation was not controlled.

In summary, the present study had as its focus to
provide an experimental ethogram and descriptive baseline
of the early deve10pment of the bobwhite quail, a non-
domestic precocial galliforme, under laboratory controlled
conditions with minimal human contact and under an appro-
priate lighting regime. Consideration is also given to
extending previous findings on the effect of isolation and
the development of behaviour patterns by defining the
sequence in which the components of a behaviour class
developed in terms of the median onset day for the group
and by observing a species previously not studied in this
manner. The results will be discussed in the tradition
of earlier ontogeny studies (Bolles and Woods, 1964;

Nelson, 1965).

METHOD

General Organization

 

This study can be considered to have three phases
that are defined by living conditions and sizes of groups.
Phase I involved two living conditions: (1) social, with
birds housed together; and (2) visually and tactually
isolated, with each bird living within an isolate cubicle.
Phase I ended when the birds were 29% days of age. At
the beginning of Phase II visual and tactual isolation
conditions were removed and the isolate group assembled
into their own social living unit by removing the parti-
tions that defined the isolate cubicles. This resulted in
the two subject groups now having identical cages. At the
initiation of Phase II the behavioural taxonomy protocols
also became identical for the two groups. This phase
followed Phase I immediately and continued until the birds
were 33 days of age. During I and II each bird was observed
five minutes at each of two daily observation periods.
Phase III was marked by the removal of seven randomly
selected subjects from the social group, in order to
equalize the N for the two groups. This phase continued

until the birds were 75 days old. The observation schedule

15

16

later in Phase III was altered to include only one observa-
tion each two—three days.

The procedure of Phase I for the two separate groups
will be treated first. Then the procedure of Phases II and

III will be considered.

Subjects

 

The subjects for this experiment, bobwhite quail

(Colinus virginianus), were obtained from the seven year

 

old Michigan State University Poultry Science quail colony.
Four hundred and fifty-seven eggs were incubated. Forty
eggs were randomly selected from undisturbed incubator
compartments prior to pipping and divided into two groups
by alternately designating each egg S or I. This selection
was made at the beginning of the twelve hour peak of
hatching for the total group. The eggs designated as S
were then placed in the standard group hatching tray and
returned to the incubator. Each egg designated as I was
placed into an individual bin within an identical tray and
also returned to the incubator. These latter trays were
partitioned into separate compartments by fitted metal
inserts and were placed into the incubator in a manner that
prevented a newly hatched occupant from viewing other
chicks. The thirty-two surviving chicks were hatched
within twelve hours of each other; fifteen of the twenty

S or social eggs hatched and seventeen of the I or isolate

eggs hatched. Within two hours of hatching each quail was

l7

weighed, leg banded and returned to the appropriate tray
in the incubator. All subjects remained there until
transferred to their respective living units.

Thus, the subjects in the social group experienced
the presence of other quail from prior to completed hatch-
ing to the end of the study; while the isolate group sub—
jects were visually and tactally separated from other
quail from the pre-hatched state and maintained in this
manner until the initiation of Phase II.

Throughout the study all bobwhite were maintained
on ad lib water and food (Michigan State University Quail
Breeder 25% protein, dry mash, King Milling Company).

Food and water trays were filled on alternate evenings
following the scheduled observation period. Water trays
were removed and cleaned twice each week. A diurnal light
cycle of 14% hours of light and 9% hours of darkness was
in effect continuously. The lights in each living unit
were turned on a 6:00 a.m. and remained on until 8:30 p.m.

Following their arrival at the Psychology Research
Building at one day of age, the subjects were never per-
Initted to view the eXperimenter and were maintained without
.human contact with three exceptions: Phase I--all subjects
‘were individually removed from their unit, weighed,
:rebanded, and returned; Phase III--at two different times
all subjects were handled briefly or experienced disturbance

(noncomitant with equalizing the group sizes and with

l8

checking and loosening leg bands. All of the above Opera-
tions and all maintenance activities (cleaning, changing
water, filling food trays, and checking cage temperatures)
were carried out either in darkness or under minimal
illumination provided by one 15 watt, warm white,
fluorescent light covered with a dark blue filter and
located approximately two meters behind the living cages.
All observations were made from an adjoining darkened
observation room fitted with one-way vision glass. No
extra eliciting stimuli were presented to any subject at
any time during the course of the study.

The sex composition of the two groups remained
unknown until sexual dimorphic coloration made positive
identification possible around 10 weeks of age. Composi-
tion of the groups as determined at the close of the
study was as follows:

Phase I: Social N = 14; 10 females, 3 males,

1 unknown (died before
positive identification

made).

Isolate N 3 females, 4 males

II
\I
‘0

Phase II: No change

Phase III: Social N = 7 5 females, 1 male,

1 unknown

‘0

Isolate N

ll
4

No change.

 

 

 

52295

feede
for t
table
meter
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preve
while
remai
previ
remai

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ident
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the b

enclo
diame
serve.

15 Wa:

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19

Apparatus

 

The birds lived in cages that provided floor,
feeder, and water space well above the minimum requirements
for this species. These living units rested on adjacent
tables in an experimental observation room 5 x 2.5 x 2.8
meters. The cage fronts faced a series of one-way vision
windows, each 22 x 51 cm. with the level of the cage floor
drOpped 10 cm. below the bottom edge of the glass to
prevent the subjects from viewing their reflected image
while walking about in the cage. The experimental room
remained unilluminated throughout the study except as
previously noted. The room temperature and humidity
remained essentially stable around a mean of 27.5°C and
41 rh, respectively.

The two living units, each 45 x 45 x 155 cm.,
identical except for the removable partitions described
for the isolation cubicles, were constructed as follows:
the back and sides were 10 mm. unpainted plywood board,
the floor was 6 mm. hardware cloth, with the tOp and front
enclosed with 12 mm. hardware cloth. Tin troughs, 75 mm.
diameter x 153 cm. length, resting on the floor surface
served as water trays at the back of each cage and food
trays at the front (see Figure l).

Cage illumination was provided by three 45 cm.

15 watt, warm white fluorescent lamps (GE F15 TlZ/WW)

attached to a center light bar running the full length

20

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.mcmsonu cw» BE mu Amy

4|

so om

EU mv

.o003SHa as OH Irv .mnaz as NH lac .muaz as o lac
.QHSQ mEmH Apv .Hmn whommsm usmfla ADV .Dflcs Hmumms Any
"pas: msfl>fla Damon mo noumxm cmflmwnln.a musmflm

EU mmH

JL

 

21

of the top of the unit. Each cage contained three lamps,
providing a range from 25 to 150 lux illumination within
a unit with approximately equal values between the cages.
Heat was available from four 38 cm. strip-type poultry
brooder heaters secured in a 75 mm. tin reflector trough
suspended beneath each cage. Temperature regulation was
effected by raising, lowering, or altering the angle of
this heater unit. The heating units were turned off
when the subjects were 36 days of age; shortly after the
initiation of Phase III.

Recording equipment consisted of the following:
an electromagnetic tape recorder (Norelco 101), one ten
pen event recorder (Esterline-Angus) Operated by a manual

key board, and one luminous dial clock.

Phase I

Subjects

At one day of age all thirty—two bobwhite were
transferred from the Poultry Science Building to the
eXperimental room in the Psychology Research Building and
placed into their respective living units. A total of
fifteen birds were housed together in the social cage.
One subject from this group died of an undetermined cause
on day 14.

Of the seventeen isolate group subjects; ten were

immediately placed into the isolate living unit; with each

22

bird occupying one complete cubicle. The remaining seven
subjects were maintained in an alternate isolation cage.
Within four days, ten of the isolate subjects (some from
each isolate cage) died of undetermined causes.

Thus from day 14 through day 29 the social group
contained 14 birds. From day 5 through day 29 there were

a total of seven isolated quail available for observation.

Apparatus

 

For both subject groups, each living unit within
the cages provided a heat gradient ranging from 30-26°C
at the water tray to 44-33°C on the floor directly over
the heating unit at mid-cage. Temperatures were lowered
gradually over the 29 day period and the lower figures
above denote the absolute degree ranges at day 29.

Unique to this phase was the isolation cubicles
defined by opaque partitions (.63 cm. masonite board)
fitted flush between the front and back walls within the
living cage housing the Isolate subject group. Thus each
cubicle provided a floor space of 270 square centimeters
(45 x 45 x 15 cm.); housing one bird in a manner that
prevented direct visual or tactual commerce with another

quail.

Procedure

 

Phase I is defined by the living conditions of the

:aubjects; social versus isolate, and by the age of the

23

subjects; including data from day 1 through day 29. The
social group birds were communally housed in one observa-
tion cage. The isolate group subjects were housed in an
identical cage, except for the partitions that defined the
isolation unit. Neither group could view subjects in the
other group although both groups experienced the same
auditory array, including noises from the hallway and
vocalizations from any bird within the eXperimental room.

Each bird was observed five minutes at each of two
daily observation periods. The initial and terminal stages
of the light cycle were scheduled observation periods.
Data from earlier investigators (Kirkpatrick, 1955),
indicated that these were periods of considerable activity
for young precocial birds. The order of observation of
individuals within a group was randomized by following a
random number table keyed to a subject identification
number. The observation order for the two groups followed
a counterbalanced design. For example, a three day block
of six successive data collection periods would be as
follows:

1. 6:00 a.m.: Begin with the isolate group,
following each bird for five minutes, with the individual
subject order determined by the random number table
designation for that observation number; following this the

social group birds would be observed similarly.

24

2. 6:30 p.m.: Begin by observing the social
group and following completion of this, the isolate group
data would be collected;

3. 6:00 a.m.: The order would be social, isolate;

4. 6:30 p.m.: The order would be isolate, social;

5. 6:00 a.m.: The order would be isolate, social;

6. 6:30 p.m.: The order would be social, isolate.
This counterbalanced schedule was followed throughout the
observations.

On day 9 following the a.m. observation the cage
lights were turned off in both units and the blue light
condition utilized while all subjects were individually
removed from their respective units, weighed, rebanded, and
returned to their cages. This was the only instance of
the subjects eXperiencing human contact during this phase
of the study.

On day 22 an all day observation period was
scheduled with two other observors alternating with the
regular eXperimenter and reliability data were obtained at
that time. Interobservor agreement was sufficient to allow
accurate deliniation of behavioural patterns.

All observations were made from a darkened
adjoining-room through a one-way Vision glass. Response
categories were arrived at on the basis of initial
observation during the first sixteen sessions and were

altered as necessary to accommodate new behaviours. These

25

were recorded on tape and later transferred to protocal
sheets and inspected to arrive at a data grouping that
would provide a functional taxonomy of the most frequent
distinctive (in most cases mutually exclusive) behaviour
patterns for each of the two subject groups. Beginning
with observation seventeen, a ten pen event recorder
equipped with a manually Operated finger key board was used
to collect data; with verbal subject identifications and
additional comments being recorded simultaneously on tape
by the eXperimenter. Responses were scored (by depressing
the apprOpriate key) on the event recorder in two ways,
either discretely or continuously. If a response occurred
in discrete units (e.g., drinking), then the apprOpriate
key was depressed each time an instance of this response
occurred. A continuous response is one which took an
indefinite amount of time (e.g., resting); the apprOpriate
key was pressed throughout the duration of a continuous
response. Thus, in addition to noting the frequency of a
behaviour within a general behaviour class, these patterns
could often be broken into their component parts and both
antecedent conditions and subsequent effects identified
sequentially and temporally. Behaviours were recorded in
the order of occurrence with frequency, duration and time
interval information available on the event recorder

paper and subject identification and other information
available from the tape. Specific behaviour categories

are first defined and then described in the results section.

26

In addition to scoring the behaviour categories;
detailed notes were kept on the feather deve10pment of the
subjects. Feather deve10pment onset was characterized in
terms of visibility of the feather shafts (still encased
within the cuticle) to the observer, from the observation
booth.

There was also a data log kept of unusual or
uncontrollable conditions occurring throughout the course
of the study (e.g., fire alarm warning system being tested
in the building), and the reactions of the subjects to
these events. In instances where events of this nature
provided fortuitious data collection Opportunities; these

will be identified in the results section.

Phase II: Procedure

 

One hour prior to the scheduled morning observation
when the subjects were 29% days of age, food and water
trays were filled in both living units. The cage top
retaining hooks were removed from both units and the
isolate living cage was prepared for removal of the parti-
tions that provided isolation cubicles. Shortly before
6:00 a.m. the tOp was removed and replaced on the Social
unit cage to provide some disturbance for the subjects in
this group. Following this the tOp was removed from the
Isolate cage, and screen wire strips were positioned over
the cubicles to prevent subjects from escaping while the

partitions were pulled. The tOp was secured into place

27

and it became identical to the Social living unit. These
manipulations were carried out in subdued, blue filtered
light provided by one 15 watt bulb located two meters
behind the cages, with minimal disturbance to the subjects,
that is, no subjects were handled by the eXperimenter.
Within a few minutes the cage lights came on for the
regularly scheduled observation.

For this observation the regular schedule was
altered. The newly assembled isolate group of subjects
was observed for two hours continuously. A regular observa-
tion of the social group followed. Another 30 minute
observation was made at noon that day, followed by another
extra observation at mid-afternoon. The regularly
scheduled observation was then carried out in the early
evening. These data from the extra observations were
kept separate from the daily observation log. Following
the initial day of Phase II, the regular observation
schedule used during Phase I was in effect until the

initiation of Phase III.

Phase III: Procedure

 

When the birds were 33 days of age the group sizes
were equalized by removing seven subjects from the Social
Group; thus each group would now contain seven subjects.
Ideally the birds to be removed in order to equalize the
N for the two groups should be identified by choosing from

a random number table keyed to colored leg bands. However,

28

since it was important to prevent the subjects from viewing
the human experimenter, a compromise procedure was
utilized. The experimenter reached into the cage picking
up one bird at a time until seven subjects had been
removed. This was carried out in total darkness without
the group of birds being disturbed. The cage was divided
into three sections and a bird was removed from each
section alternately until the required N was complete.
The extra subjects were immediately transferred to a hold-
ing cage at the far end of the room and maintained there
until the completion of the study. With the exception of
carrying out the removal of the subjects in total darkness,
the other aspects of the manipulations carried out here
duplicated those used in Phase II. That is, both cages
were disturbed, extra observations were carried out
immediately following the institution of this phase.

Following the initial day of Phase III the observa-
tion schedule was altered to include one complete observa-
tion each 2—3 days during the week with the counterbalanced
design continued until the birds were 75 days old and the
study was terminated. At this time determination of sex
was easily made by visual inspection of the beak and
plumage of each bird.

The birds were not weighed at this time because
they were scheduled to become subjects for another study;

in which the other eXperimenters wished to utilize the

29

"no—human-contact" aspect of the history of these

subjects.

RESULTS

Behaviour classes and their components were identi—
fied from three general sources: (1) the taxonomy of F
consummatory behaviours presented by Ratner (Denny and
Ratner, 1970); (2) previous literature describing these
or similar behaviours (Stokes, 1967; Ratner, 1965; Farris,

1964; Stoddard, 1931); and (3) pilot work by the present

 

author involving observation of this species in the labora-
tory and the field.

An Outline of Behaviour Classes introduces the
Measurement section. Following this the behaviours are
defined as they were scored for the purposes of this study.
The definitions of behaviours that are general to the aves
are not referenced, for example, flying, but are described
here as the author observed them to occur in the present
study. Behaviour patterns that are essentially identical
to those observed in other species by other investigators
are so indicated within the definition of that pattern.
Behaviour patterns that are described for the first time
in an experimental study of this species are indicated by
an asterisk and those first experimentally defined by the

present author are also followed by the author's name.

30

31

Following the definitions of the behaviour classes,
the descriptions of behaviour for the two groups is found.
Data is presented in tables and figures illustrating the
onset of each behaviour pattern for the individuals in the
two groups: Social N = 14, Isolate N = 7. No statistical

significance tests are apprOpriate for these data. 5

Definitions of Behaviour Classes

 

Outline of Behaviour Classes

 

I. NUTRITION

 

A. Feeding
B. Drinking

II. CARE OF THE BODY SURFACE

A. Preening
1. General
2. SOphisticated*
3. Oiling*
4. Allopreening*
B. Bill Wiping*
C. Dusting*

III. LOCOMOTION

. Running

Jumping

POpping

Flying
Flit-popping*
Frolic*

. Frolic with Object*

OWL‘UUOWS’

IV. DAILY ACTIVITY

A. Generalized Pecking
B. Resting
C. Resting Group Structure*
1. Huddle-Heap
2. Huddle-Space
3. Linear
4. Ring or 1/2 Circle
D. Cubicle Pacing*

32

V. POSTURAL PATTERNS

Crouching*

Freezing

Cautious Alert*

Sustained Tactual Contacting*

0(3U1>

VI. ALLOPECKING
A. Body
B. Head-Nares-Beak
C. Toe Picking

VII. FEATHER DEVELOPMENT

A. Wing
B. Tail
Measurement

 

Each major class of behaviour will be descriptively
identified and the components of that class listed and
described as they were scored in this study.

As this study focuses on the deve10pment of
behavioural patterns; those classes of behaviour that are
common to both groups of subjects will be described first.
Following this the behaviour classes that are unique to a

group will be treated.

Classes Common to Both Groups

 

I. NUTRITION
A. Feeding: the act of ingesting food; pecks
into food tray.
B. Drinking: ingesting of water by pecking

into the water tray.

 

33

II. CARE OF BODY SURFACE
A. Preening: the act of a bird's trimming,
dressing, or cleaning its feathers with its beak, in one of
the following patterns:

1. general preening: a soft picking or nibbling
movement directed at the feathers, characterized by orienta—
tion and movement of the bill and head toward a body area.
The intensity and duration varies widely but the form
(tOpography) of this movement is unmistakable.

2. *SOphisticated preening: a general preening
movement directed to a specific feather shaft and character-
ized by the feather being passed between the mandibles, which
"nests" the barbules. This movement may be accompanied by a
greater frequency of bill Opening and closing during the time
the bill is directed to the base of the feather (Nitschke).

3. *oil gland preening: a directed movement bring-
ing the bill to the dorsal base of the tail, accompanied by
raising of feathers in the dorsal caudal tract as the beak
tugs on the nipple of the urOpygial gland or oil gland;
following this the beak is directed to various body areas as
the bird oils the feather surfaces, legs or feet. Another
component of this pattern, frequent but variable, is a vigor-
ous rubbing of the beak in the area of the gland, character—
ized by rapid movements of the head from side to side
(Nitschke).

B. *Bill Wiping: bird's head is directed to

the floor surface at various angles; beak contact with the

 

34

surface is evident as both sides of the mandible are brushed
against the surface in any one full sequence. This may be
repeated several times in succession with minimal accompany-
ing body movement or locomotion, after Farris (1964).

C. "Dusting": bird rests on ventral body sur-
face with wings partially extended while body is "rolled" to
one side as the contralateral foot executes scratching move-
ments. Simultaneously feathers in the caudal, alar and capi-
tal tracts are raised, as both wing and tail primary feathers

spread and close several times in succession. The bird

 

U'

typically rolls from side to side repeating these movements
and pecking rapidly at the sustaining surface between shifts
in position. Shifts in position may or may not involve
vigorous scratching with alternate feet. Head and neck
extension may accompany the dusting pattern; the head is also
"rolled" or brought into contact with the surface in a rub-
bing motion. This resembles the bill wipe movement except
that the neck is twisted in such a way that the head feathers
contact the substratum. In a soft medium, e.g., dry mash in
a food tray, a body sized depression may be formed as a

result of this activity, after Stokes (1967).

III. LOCOMOTION
A. Running: terrestrial and rapid movement
of the bird in an upright position; head slightly extended
from a "streamlined" body, i.e., the feathers compressed

against the body and tail feathers trimmed.

35

B. Jumping: bird springs vertically from
floor, somewhat forward; wing movement may or may not be
invclved in ascent or descent. When wing movement is in-
volved it does not appear to be precisely synchronous. A
shallow crouch may precede jumping but is not a reliable
predictor of this behaviour. Compared to popping the
tOpography of this pattern is jagged in appearance.

C. *Popping: bird springs directly upward in
rapid vertical ascent with body streamlined on ascent; at tOp
of upward spring, wings Open, tail flares and bird is postu—
rally in correct attitude to initiate forward flight. Flight
may or may not follow. The smooth but "explosive" character
of this movement serves to make it quite distinguishable in
form from other locomotor patterns, after Farris (1964).

D. Flying: an aerial, forward propelling,
coordinated wing movement pattern that may be preceded by
any or none of the above patterns.

E. *Flit-popping: the popping pattern, but
characterized by a distinctive deep crouch preceding the
ascent (Nitschke).

F. *Frolic, or wing-flutter-run: a short burst

of running accompanied by wing movements (after Ratner, 1965).

IV . DAILY ACTIVITY
A. Generalized pecking: pecking movements
directed at inanimate objects not within the food tray; nail

heads, feces, etc. (after Ratner, 1965).

36

B. Resting: bird remains motionless with eyes
closed in one of three positions: (1) lying on side with
head and feet stretched out; (2) squatting with undersides
touching the floor with head withdrawn; or (3) standing up-
right, often on one foot, with head withdrawn (after Farris,
1964).

C. Resting Group Structure:

1. *huddle-heap: three or more birds touching in a
squatting posture and stacked into a heap such that at least
one bird does not occupy floor space (Nitschke).

2. *huddle and spacing: a huddle group in which some
birds at the edge of the group occupy floor space near the
group but may or may not be touching members of the huddle
group (Nitschke).

Contacting the environment (Denny and Ratner, 1970)
or investigatory behaviour (Scott, 1958) were classes of be-
haviour that were not scored in this study because of their
non-specifiable characteristics with regard to the nature of

the present problem.

V. POSTURAL PATTERNS
A. *Crouching: bird's breast is near or touch—
ing the floor; the neck and head are withdrawn, with the body
posterior extended vertically and the legs in a flexed posi-
tion beneath the body (Nitschke).
B. Freeze: bird momentarily ceases all observ-
lable movement and remains motionless for a minimum of 15

sseconds, in whatever posture characterized termination of the

 

37

previously ongoing behaviour at the onset of the freeze
posture. This pattern may be followed by a cautious alert
posture once movement is initiated.

C. *Cautious alert posture: bird stands up-
right with legs straight beneath the body, the tail lowered
with the head and neck extended forward, giving the bird an
erect and stretched-out appearance. Generally only the head
makes noticeable movements; shifting of a foot position may
be observed, although infrequently. On occasion, a bird may

exhibit this pattern following the freeze posture, and in

 

I-‘vc
I

this instance may take a few steps while maintaining this

posture (Nitschke).

Classes Unique to Isolate Group

 

V. POSTURAL PATTERNS
D. *Sustained tactual contacting: the general

configuration of this pattern involves the bird touching a
vertical surface, usually a partition, with at least one wing,
and the beak or head may also be in contact with the surface.
The typical intense expression of this pattern is a "pushing"
against the partition such that the feathers are noticeably
re-arranged during the extent of the contact and the bird
appears to be "flat" on one side. This form of contacting
may also be observed in the squatting or standing resting
posture or pacing pattern.

In "pushing" the bird's toes are hooked into the wire
floor 2-5 cm. away from the partition with which it is in

contact, with the legs defining an angle less than 90 degrees

38

with the bird's body. This gives the appearance of a con-
siderable contact pressure being sustained between the bird

and the vertical surface in the standing position (Nitschke).

IV. DAILY ACTIVITY
D. *Cubicle pacing: bird travels the length
of the cubicle, crossing the mid-line in continuous and suc- ..
cessive trips, walking rapidly with the body upright and the D
head slightly extended (Nitschke). This is often accompanied

by contact between the beak or breast area and a vertical

 

‘I .'.‘ . .

cage surface throughout the traversal of the cubicle. Three
movement rates were defined by the frequency with which the
bird crossed the mid—line of the cage and traversed not less
than one half of the possible area on any one trip:

1. Low Frequency = less than or equal to 5 trips during
one observation period (5 minutes)

more than 5 and less than 15 trips

2. Medium Frequency

"frantic" pacing = 15 trips or more
each minute for two or more minutes
of the observation period.

3. High Frequency

Classes Unique to Social
Group Through Phase I

VI. ALLOPECKING
A. Body pecking: pecks delivered to body
(usually dorsal-caudal tract) area of another bird of suffi—
cient force to elicit at least a recoil response on the part
of the recipient.
B. Head or nares pecking: a peck directed at

the head, nares, or eye of another bird of sufficient force

39

to elicit a recoil and usually avoidance from the bird
receiving the peck.

C. Foot or toe pecking: pecks directed at the
toes or lower tarsus of another bird, may often involve pur-
suit of the victim by the bird delivering pecks and attempted

escape on the part of the victim.

II. CARE OF BODY SURFACE t}
A. Preening:
4. *AllOpreening: a general preening pattern

directed at the head or body area of another bird; may

 

1" ._

involve pecking bits of food from feathers, but this cannot
be determined by visual observation. The pattern is dis-
tinguished from body pecking by observing the reaction of
the recipient, i.e., there is no recoil or avoidance move-

ment. This pattern may occur frequently during huddling.

III. LOCOMOTION
G. *Frolic with object: frolic while carrying
some object, e.g., a feather, in the beak (after Ratner,

1965). This pattern may elicit pursuit by other birds.

IV. DAILY ACTIVITY
C. Resting group structure: in addition to the
basic resting postures and structures previously described,
the social group exhibited two patterns never observed in the

isolate group quail, as follows:

40

3. *Linear: three or more birds squatting in a
resting posture touching each other laterally but with
every bird occupying his own floor space. The birds may or
may not be facing the same direction but the configuration
appears linear (to be one of a line) (Nitschke).

4. *Ring or half-circle configuration of a resting
group: several birds in the linear structure form a con—
tinuous line that may take the shape of a semi-circle or
complete circle, with the heads of all the participating

birds oriented in the same direction and the tails of these

 

birds defining the interior of the circle (after Stoddard,

1931).

Descriptions of Behaviour

 

Behaviour of the Social Group

 

General description of typical observations.--A
typical pattern of activity observed for the birds in this
group would be as follows:

1. Within a few minutes after the onset of light
most birds would have arisen from their resting huddle and
begun to drink, feed, or engage in brief bouts of general
preening; a 2-3 minute feeding bout invariably marked th

beginning of their day.

- 41

2. The next prolonged activity was Care of the
Body Surface.

3. Within ten minutes the synchrony of the group's
activity was less evident as bouts of feeding, preening,
drinking, moving about the cage, generalized pecking, and
resting were interspersed with each other. The quail
spent various amounts of time feeding or climbing in and
out of the food tray for much of the first hour before
settling down as a group for their first morning nap.

As shown in Table 1, it can readily be seen that
the birds were all eating and drinking well under the
laboratory conditions of this study at one day of age.

The evening observation differed only slightly from the
above description. Often the group would be found resting
or scattered in small groups throughout the cage preening
and dozing. At least one and usually several group feeding
bouts would be noted; these also would occur alternately

or be interspersed with other general activities.

Incidents of allOpecking encounters were typically uni-
lateral and were observed to be interspersed throughout the
other activities and appeared to be integrated into the
daily pattern. These encounters were brief and never
resulted in fighting. A few of the Locomotion and Postural
Pattern behaviours occurred in isolated instances in a

single bird; however, the more usual incidence of these

 

42

 

 

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onwmum

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mcHommm can OHOOOI

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msHummm

mconmm UONHHmumcwo
>9H>HBO< NHH<O

44

behaviours was a group of birds exhibiting the pattern in

response to some identifiable condition or stimulation.

Onset of behaviours in the different consummatory
classes.--Care of the Body surface activities came in
early. The sequence in which these components developed
as indicated by the median onset day for the group is as
follows: General Preening (day 2); Bill Wiping (day 6);
Sophisticated Preening (day 15); Oil Gland Preening (day
17); Dusting (day 19); Allopreening (day 36). Of the
class Care of the Body Surface, general preening is by far
the most frequent. It tended to occur at any time during
an observation and in a variety of locations within the
cage, as well as in and out of huddles. As Figures 2 and
3 show, this preening pattern developed rapidly in the
social birds and was a pOpular activity.

SOphisticated preening appeared as the feathers
developed into the first juvenile feather structures and
usually occurred in bouts having a longer duration than
that of general preening. Typically this pattern did not
occur in the mornings until after the first feeding bouts
were finished and the group was beginning to nestle down
into huddles.

The pattern for oiling or oil gland preening, like
that for sophisticated preening, was of bouts of long

duration once initiated. Oiling developed almost

.OQHOOOHQ UQmHm HHO HNV

ON

P

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min 338 CID

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45

 

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queo 18d eArqunan

46

.OOHHOQ xmmz O m Hm>o mmsoum mDmHomH Ocm
HOHoom OH mason mcHnmem Hmumqmm mo MOO Hmm hocmsvmhm qm02||.m OHOmHm

mxmmz OH mom

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

m H
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9
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n
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OH w
A
HOHoom .l; .NN
ON
OHMHomH I

47

simultaneously with sophisticated preening and tended to
occur at the same times; the two were often engaged in
alternately by an individual bird. These patterns were
more separated in time from other activities such as
feeding or generalized pecking than the pattern of general
preening, that is, they were not as likely to be inter-
spersed with concurrent activities.

Bill wiping also develOped within the first week
and as shown in Figure 4 was algo a fairly pOpular form
of activity. It was observed to occur at any time during
an observation and was typically quite brief and inter-
spersed with a variety of other behaviours. Bill wiping
always occurred while the bird was in an upright position
and the surface contacted was the cage floor. It appeared
to follow generalized pecking most frequently when a bird
had pecked at a fresh feces. It also followed drinking,
feeding, pecking another bird, and receiving a peck from
another bird, as well as occurring in instances where no
previous pattern appeared to be related to the event, such
as running up and down the cage, popping, and preening
activities. Its ubiquitious and general occurrence suggests
that it is more properly classes as a form of Care of the
Body Surface than as a "greeting gesture to other chicks"
as suggested by Spalding (1873).

Dusting or dust bathing was not an activity I had

anticipated observing in this study. However, the birds

48

.OOHHOQ Mmm3 O m Hm>o masonm OHOHOOH
can HOHoom CH OQHOHB HHHm mo MOO Hmm mocmsvam cmeII.O OHOOHM

mxmma CH OOH

 

 

 

 

 

 

 

 

 

 

v m N H
J [I‘ll—Ulla
vfi w
E
vm u
ONH "M
w
ill. fimH %
rON W
.ON [u
.3 .m
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MHUO
H . w . - [I Hmm k

 

 

OHOHOmH I

49

found the food troughs suitable dusting sites; and only
complete bouts in this location were scored as a dust bath.
The scratching and shuffling components of this behaviour
also occurred on the wire cage floor directly over the
heating unit strips. This suggests that the condition
eliciting this pattern needs further investigation, as
Schein (1968) implies that this pattern will not occur on
wire. The dusting pattern was infrequent compared to the
other Care of the Body Surface activities.

AllOpreening occurred most frequently when the
birds were in a huddle or when an individual approached
an already formed huddle in which some of the birds were
engaged in preening. No obvious appetitive display compon-
ents were observed that appeared to function as a solici-
tation to initiate allOpreening on the part of the bird
receiving the preening. Proximity seemed to be a suffi-
cient stimulus for the behaviour to occur. However,
once initiated, the recipient would often extend the neck
and turn the head so that the feathers between the mandibles
and the ear were well exposed and readily available to the
other bird. Allopreening bouts were generally briefer in
duration than either of the more elaborate forms of auto
preening; SOphisticated or oil gland preening. The possi-
bility exists that hits of dry mash in the feathers of
another bird would elicit this behaviour; however, it was

not possible to determine this by visual inspection under

50

the conditions of this study. The form of allOpreening
was overwhelmingly unilateral in occurrence, with mutual
preening being observed only five times throughout the
course of the study. This might be taken as supporting
the suggestion that food bits served to elicit this
behaviour. However, the relatively late onset of this
behaviour pattern as shown in Figure 5 would suggest that
the food bits were probably not the critical factor. In
addition to this, feeding off the back of a bird (that had
been covered with food as a result of a quail taking a
dust bath in the food tray, or scratching in the food) was
observed to occur within the first week and had a
tOpography that distinguished it as clearly different

from an allOpreening behaviour. Farris (1964) observed
mutual preening frequently in the Japanese quail, but no
comparable data exist for the bobwhite.

The sequence of deve10pment of the basic Locomotion
patterns in the group using the median onset day was as
follows: Running (3), POpping (3), Jumping (9), Frolic
or wing-flutter-run (10), Frolic-with-object (l3), Flit-
pOp (l4), and Flying (20). These patterns were evident at
various times during an observation and, once they were
well develOped in all of the subjects, appeared to occur
most frequently as a result of social facilitation. This
was particularly true of the patterns of pOpping, flit-

pOpping, flying, and frolic-with-object. Once flying was

51

.mmsoum OHOHomH cam HOHoom QH OQHQOOHQOHHH mo ummco:l.m OHOOHM

Omm mo mmma

 

OHOHOOH .
HOHoom AU

.hm

21.1180 18d 8111:7191:an

 

 

52

initiated by an individual, often within seconds, the
entire cage would be a flurry of birds flit-pOpping and
flying from one end of the cage to the other. It was
impossible to discern what had initially stimulated the
first bird when this occurred. However, the daily fre-
quency of popping appeared to be related to the arousal
state of the group. This inference was suggested by the
fact that when workmen were repairing heating ducts in the
ceiling of the building during a series of observations
the birds appeared to startle easily and often this was
followed by the cage suddenly exploding into a group of
"popping" birds.

The frolic-with-Object pattern often developed
into a sequence of activity involving various numbers of
birds over a continuous 3-4 minute period. This pattern
tended to occur most frequently late in the morning
observation period or early in the evening period and
usually stOpped as abruptly as it began.

Shortly after the wing primaries were achieving
some length (see Figures 6 and 7) the quail began flying
the length of the cage and on several occasions a bird
flying overhead elicited crouching in a bird on the cage
floor. Unfortunately due to the flurry of activity within
the cage at these times it was impossible to obtain a
consistent measure of this relationship, but it should prove

a fruitful area for further investigation.

53

.mmsoum OHOHomH OOO HOHoom OH OOHNHM mo uOmOOII.O OHOOHO

mom mo mhma
HN ON OH OH OH OH OH

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.h
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54

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OOO mo mama
ON NN ON OH OH OH NH OH O O

n P D v D p r P L

 

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.mm
OOH

 

'1

55

Generalized pecking had been observed in each of
the birds in this group by the third day of age. This was
a very frequent behaviour and was likely to occur any time
a bird was up and moving about the cage. Included in this
category of behaviour was a component of the "competitive
pecking" pattern described by Farris (1964) as "two or
more birds pecking at the same object." The most fre-
quently pecked "object" in this instance was a fresh feces
on the cage floor. Individual birds pecking the cage wall
was by contrast an infrequent occurrence.

Resting as a class of behaviour occurred several
times within any one observation, particularly during the
first two weeks of age. The postures of lying on one side
or standing upright were less typical of the resting
behaviour of these subjects than was the squatting with
the underside touching the floor. A group of birds resting
appeared to have considerable eliciting strength as a
stimulus to approach. If an individual bird had been part
of a resting group and the others in the group moved to a
new location in the cage, the bird left alone, upon
awaking, would arise and run rapidly to join another
resting group. Since shifts in position of the individuals
in a resting group were frequent, particularly during the
first 10-14 days, the composition of the resting group was

continually changing concommitant with a change in

56

location within the cage. Thus, a sleeping bird might

often go to sleep in a group and awaken to find himself

alone.
As noted in Figure 8 the Resting Group Structure

went through several major changes in predominant forma-

tion patterns. The first few days the group of birds

most resembled a pile of bodies stacked into a corner,
with individuals continually changing position within the

heap. Toward the latter part of the first week, this heap

of birds began to flatten out somewhat as fewer birds
attempted to crawl onto the top of the pile; instead they
vwould more frequently join a resting group by approaching
tflie edge and nesting down near a bird on the periphery or
airtempt to crawl in between their mates and eventually
sxmcceed in inserting their body into the group and gaining

scnne floor space. The onset of this Huddle-Spacing struc-

tixre also marked a change in the location of resting groups
axvay from the ends of the cage next to the plyboard walls,

toward the middle or Open areas of the cage, but still
Ipxrimarily positioned over the heating units. Depending
<311 the general level of activity within the cage, there
Unjught be one or more of either of these groups formed at

Eirry time during an observation as the quail interspersed

Short resting bouts with most other activities. The

Idirlear Resting Group Structure became evident toward the

57

 

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OOHHOOH OOOHO mo mOHOHoOHHm OHOOHEOOOHQ mo OOHHOHOU OOO HOmOO .O OHOmHm

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58

end of the second week and was also typically observed to
occur over the heating units. One of the distinguishing
aspects of this pattern was that the birds less frequently
attempted to crawl in between other resting birds; rather
a bird would approach a resting group and settle next to
the side of a bird bringing his wing area into contact
with the bird laterally and sink into the squatting
posture.

Although the above patterns were observed at
various times throughout an observation, the Ring or
Nesting Circle Resting Group Structure was most frequently
noted at the beginning of either a morning or evening
observation and sometimes during the evening observation.
This pattern was also somewhat different than the others
in that it appeared to be more stable or, once formed
involved fewer position shifts among the constituents,
and at full intensity involved all of the animals within
the cage when it was observed. Once the nesting circle
broke up at the beginning of a morning observation it
rarely reformed during that period at full intensity,
that is with all the group participating. However, there
were several instances of smaller groups forming the Half-
Circle configuration for brief periods during both
scheduled observations. When smaller groups formed the

pattern it most frequently took the 1/2 circle form, which

59

concurs with Rosene's (1969) statement that due to the
positioning of individuals within a "roosting disk" the
minimum number required to form the complete circle is

six birds. Also lending support to this suggestion is

the fact that one of the few changes in behaviour observed
to occur directly in relation to the initiation of Phase
III of this study, the removal of seven subjects from this
group, is that the full circle resting structure was
observed less frequently immediately following the Phase
III manipulation than previously. However, it did not
cease to occur among the remaining seven subjects.

It is impossible at this point to rule out the
possibility that removal of birds from a group is a dis-
ruptive or interfering event with regard to a social or
group behaviour pattern. The question of number of
animals certainly suggests itself as a variable to be
investigated in further research of this nature.

The sequence of deve10pment of the behaviours
referred to as Postural Patterns was: Freezing, Cautious
Alert, and Crouching. As shown in Figure 9 the freezing
pattern occurred in all the birds in the group on day 7.
Since there was no attempt at a systematic manipulation
of conditions that might elicit freezing in the bobwhite,
the day of onset shown here may only be incidental

developmental information. Other investigators (Stokes,

.OOm OH HOOOH HO How mmOHOOHHOE mOHOEOO
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mo mmsoum UOOOOH OHOHOOH OOO HOHoom OH OOHNOOHO mo HOmOo .O OHOOHm

O04 mo OOOQ
ON ON ON mN HN OH. OH OH MH Hm O

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60

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queo zed eArqunmna

 

 

 

61

1967; Bent, 1963; Stoddard, 1931) report freezing to occur
at one day of age in response to auditory stimulation. The
occasion for freezing on day 7 in the present data was a
metal flashlight accidentally dropped on a metal stool in
the recording booth. The striking aspect of this pattern
for the social group was the synchrony of the behaviour;
every subject in the social cage froze for 90 seconds and
all the subjects appeared to terminate the pattern at the
same instant. That is, the birds appeared to begin move-
ment as abruptly as they had ceased movement. Prior to

the onset of the freeze, there was a momentary flurry of
activity within the cage that was so rapid and brief that
it was impossible to describe or attempt to measure. This
very likely could have been the escape-fleeing behaviour
(Guhl, 1958) or dash-to-cover (Stokes, 1967) described by
other investigators as the initial response to a "strange
sound." Freezing was observed to occur in this synchronous
pattern at various times throughout the study and in most
instances it could be identified as a response to unusually
loud noises occurring in the hallway outside the experi-
mental room, such as a soft-drink bottle kicked down the
hall. The most striking occurrence of this pattern was a
freeze of four minutes in duration that was initiated by
the janitor opening the door to the experimental room
rapidly and banging it into the table on which the living
units rested. This occurred during an evening observation

in which I was recording this group.

62

The Cautious Alert Posture appears to be similar
to the "Alert" stance described by Farris (1964) in the
Japanese Quail. The conditions eliciting this Cautious
Alert pattern appeared to be sudden changes in the general
level of stimulation or sudden alterations in the environ-
ment; it was frequently seen when a drop of water was
splashed onto a heater covering, producing a brief
"sizzle." As illustrated in Figure 10 this pattern
appeared in a few birds before the Freezing pattern. It
appears that it may be related to this general class of
behaviour. Once the Cautious posture had developed in all
of the birds in the group; it exhibited a topography of
synchrony analogous to that described for the freezing
pattern. On day 20 one of the cage lights went out in the
social living unit; there was an immediate flurry of
activity as the quail scattered to the end of the cage
and went into the Cautious Posture as a group. All of the
subjects oriented toward the other end of the cage while
maintaining this posture for 75 seconds and making only
small twisting movements of the head. This was followed
by an abrupt transition to a more "relaxed" posture and
the birds began to move about the cage. The other link
to the freezing pattern was that the cautious posture was
observed to follow an incidence of group freezing
behaviour on three occasions. However, this was not

observed at every incidence of freezing. The present

63

.OOOOHOOmmO =HOo OOOOHOHHO= H5O HOOHO OO UHHO OOH OOH>HO
.OHOBHOH OOOOOHxO OOOO OOO OOOO OHH3 .OOHOSOH HHOH OOO HOOHOHHO mOOH OHHB
HOOHHQO mOOOHm OHHm .OOOHmom HOOHO mOOHHOOU OOH mo HOmOOII.OH OHOOHm

OOO mo OMOQ
om ON ON ON NN ON OH OH OH OH O O O

o O u o . . b b b x P p p a
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OHOHOOH Av .O
HOHoom Av .OH
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Hom
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53

 

 

 

 

31180 18a SATQEIHUIHD

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64

observations do suggest that it might be profitable to
approach these patterns as one class of consummatory
behaviour and consider the possibility that the cautious
posture may be a post-consummatory component.

Although previous investigators have suggested
that Crouching is a response related to fear in the bob-
white (Borchelt, 1970; Stokes, 1967) no direct data on
this question is available. It is included here, Figure
11, as a Postural Pattern because of the distinctive
postural topography which characterizes it and because
no firm data are available to suggest what class of events
might consistently elicit this pattern in the immature
quail. It was not observed to be necessarily a social or
group behaviour in the manner described for the previous
postural patterns. This is not to say that it never
appeared in several birds in any one instance. It was
most frequently observed during a period when the cage was
very active or the birds appeared to be aroused and there
were a variety of Locomotion patterns occurring such as
pOpping and flying. Since these behaviours appeared to be
socially facilitated and often involved several birds in
rapid-fire sequences of behaviours, it was impossible to
isolate for certain the pattern or class of events that
might be eliciting the crouching pattern. In several
instances it was observed to occur when a cage mate flew

over the head of a quail standing on the floor, but this

65

.mmOOHO OHOHOOH OOO HOHoom OH OOHOOOOHU mo HOmOOII.HH OHOOHO

OOO mo OOOQ
om LN ON ON ON ON ON ON NN HN ON OH OH OH OH OH OH OH NH

- s b p p n p n n p p b In 5 n + p P p

OHOHomH o - OH
388 O . Hm

 

O HO

. mm
O03

 

 

queo Jed eAraeInmna

66

was not found to be a consistent elicitor of the crouching
pattern.

Allopecking was observed within the first few days
of age in a few birds and had been observed in each of the
subjects by day 12. This class of behaviour was divided
into three distinct types: body pecks, head or nares
pecks, and foot or toe pecks. All three forms were pri-
marily unilateral and only rarely involved pursuit of the
pecked bird. Reciprocal or mutual pecking was so infre-
quent (from 0-4) that it will not be considered further
except to note that fighting with pecks (Ratner, 1965)
and bill-fighting (Stokes, 1967) was essentially absent
from the consistent behaviour patterns of these birds.

The frequency of allOpecking was measured by counting the
number of occurrences as pecking encounters. This defines
each entry or each pecking event as an encounter between
two birds in which pecks were delivered in one instance
of pecking not separated by an intervening behaviour on
the part of either bird: (1) the bird delivering the
pecks walking away, going to drink, pecking the floor, or
(2) the bird receiving the pecks avoiding the pecker
successfully for a minimum of four seconds (one space on
the event recorder paper). This means that the measure
representing frequency of allOpecking shows not number of
individual pecks delivered, but number of pecking

encounters between subjects.

67

AllOpecking tended to occur sporadically through-
out either of the daily observations and in all locations
within the living unit. The condition which led one
bird to begin pecking another could not be determined with
certainty, there appeared to be no consistent postures,
cage location, or measured behaviours that were reliably
related to the initiation of a pecking encounter. The r3
data notes suggest that it might prove fruitful to investi-
gate the relationship between bill wiping and drinking and 9

subsequent receipt of pecks. On several occasions various

 

numbers of the quail would engage in what can only be .
described as "playing" in the water trough; jumping in

and out of it, sticking a foot in and kicking, or swishing
their beaks back and forth in the water. This usually led
others to engage in similar behaviour. Pecking encounters
of the types observed at other times frequently followed
these water games. It is quite likely that in this
situation water drOplets were the elicitors of pecking.
The form and descriptive criteria of allOpecking would
necessarily lead one to identify these behaviours as
aggressive pecks as described by Ratner (1965). However,
even though it appeared to be this type of pecking that
was observed, it is curious that there was never an
instance of a bird being visibly wounded as a result of

a pecking encounter and raw or bloody spots were never

observed to occur in this group of quail. Pilot work

68

with this species and observation of the quail in the
Borchelt (1970) study highlight this as a somewhat
unusual set of data. However, Stoddard (1931) and Stokes
(1967) suggest that "peaceful relations" between bobwhites
in a group are the rule rather than the exception at all
times of the year except for the mating season when
mature males engage in fighting. As can be seen in

Table 2 the frequency of allOpecking in this group was
never very high. Body allOpecking was so infrequent that

it is not shown (0-3 encounters per observation).

TABLE 2.--Range and median frequency of allOpecking
encounters of a series of observations for each week of
age in a group of socially reared bobwhite quail.

 

 

 

 

Head-Nares-Eye Pecking Toe-Foot Pecking

Range Median Range Median
Week 1 0-20 6 0-13 4
Week 2 7-26 13 3-25 21
Week 3 3-38 9 2-44 12
Week 4 5-31 16 2-42 13
Week 5 3-13 13 2- 7 6
Week 6 2-10 6 l- 5 1

Week 7 2- 3 3 1- 8 8

 

69

BehaviOur of the Isolate Subjects

Behaviour during isolation.--

(1) General description of a typical observation:
The first two weeks the behaviour of the isolate subjects
could be characterized as showing two distinctively pre-
dominant patterns, either long periods of sleeping or
continual movement such as running the length of the
cubicle. The running pattern was so striking that a
separate measurement category was develOped to differentiate
a distinct form of this behaviour, namely that of Cubicle
Pacing. Following light onset in the mornings some sub-
jects would remain in their resting posture and others
would begin immediately to pace the length of their
cubicle, from the food tray to the water tray. Feeding
and drinking bouts were quite brief in duration, as were
general preening episodes. When the birds were resting
they slept directly over the heating units and maintained
contact with a cubicle partition. The topography of this
contacting behaviour and its ubiquitous occurrence in
each of the subjects led to the deve10pment of another
separate measurement category in an attempt to further
«characterize the general pattern of behaviour in these
:isolated birds. This pattern, Sustained Tactual Contacting,
\vas observed to occur throughout both observation periods

Enid was not restricted to periods when the birds were

70

sleeping. In fact it appeared to be a behaviour pattern
that was engaged in by a bird concomitant with all other
activities except those whose form directly prevented its
occurrence, such as feeding or flying. The most pOpular
location for this behaviour, as with resting, was also
directly over the heating unit.

Although Cubicle Pacing and Sustained Tactual
Contacting continued to be predominant behaviours during
the third and fourth weeks, more time was spent now
engaging in feeding, drinking, preening, and other forms
of Locomotion. In addition to the contacting and pacing
patterns, all the behaviour patterns observed in the
social birds (excepting those that isolation conditions
prevented from occurring) were seen in at least some of
the isolate subjects during isolation with one exception.
The frolic-with-object Locomotion pattern was never
observed in an isolate subject during the Phase I condi-
tion of isolation.

(2) Onset of behaviours in the different measure-
ment classes: Following the outling in Table 1 it can be
seen that the isolates were feeding and drinking at one
ciay of age. Feeding bouts of more than a few seconds at
zany one episode were unusual in these subjects. It
zippeared that most of their ingestion of mash from the
:fiood tray took place a few pecks at a time, in between

fmacing trips. The second week three of the isolates

71

developed a pattern of vigorous scratching in the food
tray. For several days this behaviour was so frequent
that their food troughs had to be refilled daily instead
of every other day, in contrast to the typical procedure.
At times this scratching behaviour was accompanied by
feeding though not consistently. Even during these
scratching-feeding bouts, the feeding behaviour was a
pattern of isolated individual pecks interspersed with
relatively longer scratching episodes. Two of these
three birds were in adjacent cubicles but the third quail
that consistently scratched his food tray empty was at
the other end of the living unit. Eventually all of the
Ibirds were observed to scratch in their food but only
'these three evidenced the behaviour at the intensity
ciescribed above and the pattern all but disappeared in
tflnese birds the latter part of the third week.

It was difficult to make precise notes on all
(irrinking behaviour because the subjects would often
aiztempt to stick their beaks through the wire between the
<311bicle partitions in the water trough. However, distinct
cirinking behaviour was also observed in each of the birds,
lit) addition to the above pattern.

Care of the Body Surface patterns began showing
11E> the first week in some birds. As illustrated in Table

1- comparing columns 1 and 4, there was considerable

I

72

variation in the day of onset for the individuals in this

group. Although general preening appeared in all of the
birds within the first week, it was not a predominant
aspect of their daily activity and was typically brief in
duration, on the order of a few seconds at any one instance,

when it was shown. The second to third weeks it increased

considerably both in duration of individual bouts and as
shown in Figure 3 in frequency of occurrence. Bill wiping
was the next behaviour to develop in this class and its
tOpography with respect to both duration and frequency
xuas quite similar to that described for general preening.
In addition to this, abortive or fragmented bill wipes
xwere observed in these subjects, that is, a brief down-
xward swipe in only one direction and failing to contact
1:he surface of the cage floor, was a pattern observed in

1:hese subjects. During the first 10 days this pattern

“Has almost as frequent as the complete bill wipe pattern

sczored as an incidence of bill wiping, as defined in the

Inezasurement section. It appeared that bill wiping most

‘txrpically followed drinking in the isolate subjects but
trlis was definitely not the only time at which it occurred.
Early in the third week the sophisticated preening
Pattern appeared to be well developed in these birds.
C=<Danaring Figures 7 and 16 it can be seen that this

E?€it:tern appeared to develop as the wing primaries

d€3‘JelOped. Most of the sophisticated preens were directed

73

to the wing primaries and were most likely to occur in
association with a resting period. Though it was observed
in each of the subjects during Phase I it was not a
typical pattern of their daily activity. Oil gland preen-
ing, which was observed in only two subjects before the
third week and reached 100% onset the last day of isola-
tion (see Figure 2) was by contrast with SOphisticated
preening a rare occurrence. It was never seen in more than
two birds during any one observation period and when seen
it was not often exhibited along with the SOphisticated
preening pattern. The oiling tended to be relatively
brief or fragmented, that is, often only one part of the
body would be "annointed" and only a few tugs at the oil
gland would be in evidence at one sitting.

The dust bathing pattern was observed in only two
of the seven birds during isolation and occurred in the
food trough. A pattern of scratching on the cage floor
was also evidenced in these same two animals. The curious
thing about this scratching component here is that it was
observed to precede settling down into a resting posture,
and was not followed by the kick and shuffle components
clearly evidenced in the full dusting pattern.

Locomotion patterns developed in the isolated
subjects in the following order: running, jumping,

frolicking, pOpping, flying and flit-pOpping. Running and

74

jumping both develOped within the first few days and were
behaviour patterns that characterized the activity of these
subjects much of the time when they were not resting or
engaged in a continual pacing pattern. As shown in

Figures 12 and 13 these patterns developed rapidly in this
group of subjects. The two patterns were often observed

to occur alternately in an individual bird exhibiting a fit
quality that can best be described as similar in nature

to escape behaviour. An individual bird would often spend

the majority of one observation unit (five minutes) engaged

 

in short runs in the cubicle, alternated with jumping
against the partition, climbing in and out of the food
tray and attempting to stick its head through the hardware
cloth cage front, running with its beak in contact with
the partition and stOpping every few inches to jump up
against the partition. Another interesting component of
the running pattern seen in this context was that the birds
often ran "looking up" while in motion. The running
pattern observed as a part of this escape type behaviour
was differentiated from the pacing pattern by the start
and stOp quality and the intervening jumping behaviour.
During the second week the jumping against the partition
decreased as pOpping began to appear and became part of
the running activity. Also about this time another strik-

ing pattern began to be evident, that is, individual birds

75

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76

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77

began showing fairly consistent patterns of integrated forms
of locomotion and other activities. These patterns were so
consistent for an individual animal that they could best

be described as a form of stereotype. An example of one

of these patterns exhibited by subject #1 is as follows:
The bird would suddenly dash to the water tray, peck the
partition where it was notched near the trough, go to the
Opposite side of the cubicle, jump up and peck the second
bolt head, run along the ledge just inside the outer cage
wall to the front of the cage, cross to the Opposite side
of the cubicle and peck the cage partition, run alongside
the partition, at times stOpping to jump against it

though not always, till the bird reached the water trough
and would again peck the partition notch and repeat the
cycle. This pattern would often occur 5-10 times without
interruption and would cease as abruptly as it began.

Other than the stereotyped form of behaviour no other
regularities about its occurrence were discernable to the
experimenter. Once it became a regular feature in the
activity of that bird it continued throughout the isolation
period. Another subject developed a regular pattern of
chasing one wing tip and would often turn circles con-
secutively for several seconds while attempting to peck

the protruding feather.

78

POpping often appeared to be elicited by auditory
stimulation from an adjacent cubicle where another subject
was pOpping or jumping. However, vibratory stimulation
could also be involved in this instance since all the
isolate subjects shared a common wire floor and pOpping
in these subjects was a fairly dramatic movement; at
thirteen days of age they were popping fifteen inches
above the floor. Flying developed in the isolate subjects
in the third week and also seemed to be related to the
activity of the bird in an adjacent cubicle. These
subjects were capable of flying from the front to the
back of their cubicle and would often make several of
these flights in quick succession, particularly if the
adjacent bird continued to fly or jump or pOp during the
interim. However, it was also a typical occurrence for
one bird to be flying or jumping and the adjacent bird to
continue sleeping or preening or feeding with no notice-
able change in behaviour. By contrast with the previous
forms of locomotion discussed, frolicking and flit-popping
were observed in these birds only occasionally, that is
they were never observed in more than two birds during
any one observation and were not observed to occur
regularly within the series of observations during the
Phase I isolation conditions. Noteable here, due to its

absence, is the frolic-with-object pattern of activity.

79

As shown in Figure 14, this pattern was not observed in
any subject in the isolate group until the initiation of
Phase II of the study.

The behaviour class Resting comprised a major
portion of the behaviour of the isolates during the first
week. It was not unusual for a subject to sleep through
his scheduled observation period and still be sleeping
after the other isolate birds had been observed. All
three forms of resting postures were observed in these
subjects but the standing upright posture in conjunction
with the Sustained Tactual Contacting pattern best
characterizes the nature of their resting behaviour during
this period. Although the "pushing" component of this
contacting pattern was the more dramatic instance of the
contacting behaviour, all the forms of resting involved
contacting a partition. It is interesting to note that
the subjects at either end of the isolate living unit
rested next to their partitions rather than next to the
wooden cage walls. This contact resting pattern typically
involved some preliminary movements of a type that can
only be described as a form of rubbing the body surface
against the partition several times and often from alter-
nate directions, before settling into a resting posture
with one side of the body pressed against the partition.
As mentioned previously this consistently occurred directly

over or very near to the heating unit location. It is

80

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queo lea eArqunan

81

possible that these masonite partitions retained suffi-
cient heat from the heating units to provide a warm
surface and thus had considerable eliciting value for
these quail. If this were the case, it would tend to
support Spalding's (1873) contention that the first
response of the newly hatched chick is to maintain contact
with a warm surface. As can be seen in Figure 15, the
daily incidence of occurrence of this contacting pattern
remained high over the four week period of isolation.

The behaviour pattern that appeared to compete
most effectively with the resting and contacting pattern
was that of Cubicle Pacing. As shown in Table l, by the
third day all of the isolate subjects were exhibiting
this pattern of traversing at least 1/2 of the length of
the cubicle (45 cm) in continual and successive trips.

The topography of this behaviour had a ritual-like
appearance and at its most intense eXpression, that of

the high frequency pace or a minimum of fifteen trips each
minute for at least two minutes of a five minute observa-
tion unit, can only be characterized as a "frantic" type
behaviour. Figure 16 illustrates the relative mean
frequency per day of the incidence of the three types of
pacing for subjects in this group over the four week
period. The continual and frequent occurrence of this
pacing behaviour might be considered to be so pervasive in

the daily activities of these subjects that it might be

82

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83

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84

useful to consider it as a competing response system, when
looking at the relative infrequent occurrence of Care of
the Body Surface behaviour patterns.

Another aspect of the pacing pattern was the per-
vasive and apparent uninterruptable nature of this
activity. On day 3 the fire alarm bell was tested in the
experimental building during an observation. On the first
ring the social subjects huddled into a corner and
remained quiet during the following five rings. In con-
trast to this the isolate subjects appeared to exhibit
the "scatter" pattern very briefly on the first ring and
immediately returned to pacing in the cubicles. On day 7
which was the first recorded freeze for the social birds,
when a flashlight was dropped in the recording booth, the
isolate jubjects--apparently exposed to the same stimula-
tion--only continued to pace. On day 9 in response to a
soft-drink bottle being kicked in the hallway, the social
subjects all froze as a group for a 25 second period.
Three of the isolate subjects, one on each end and one

near the middle of the living unit, never stOpped pacing;

the other four isolates paused momentarily but began moving

before any subject in the social group made a movement.

The first clear incidence of freezing in the isolate group

was a 30 second freeze in subject #10 on day 17. On day 19

an incident previously described for the social group, that

of a light bulb going out, to which the social group

 

 

85

"exploded" to one end of their cage, was apparently the
stimulus for freezing in five of the isolates. Subjects

3 and 4 froze for 30 seconds; subjects 7, 8, and 10 froze
for 45 seconds and the other two subjects only became
inactive for a short period. It is interesting to note
that freezing as a synchronous behaviour of the group does
not appear in the isolate subjects during isolation.

The Postural Pattern of crouching in the isolate
birds very clearly shows up during isolation as seen in
Figure 11. On day 16 the first crouch was observed to
occur in this group and appeared to be in response to
another isolate subject flying in his cubicle although
these were not adjacent birds. Within a few days three
of the isolate birds began a pattern of crouching prior
to initiating flight. The more usual pattern after this
time appeared to be one of a subject crouching in response
to noises in the hallway or those made by other quail in
the experimental room. The Cautious Alert posture appeared
under conditions similar to those described for the social
birds but in contrast was only observed to occur in single
subjects at any one incidence of the behaviour while the

birds were isolated.

Behaviour after being assembled.--Removal of the

 

partitions that provided the isolate cubicles was the

occasion for continual "conversational cheeping" (Stoddard,

86

1931) among the isolate subjects but no distress calling
was heard as this procedure was carried out by the
experimenter.

Following light onset in the newly assembled group
the first noticeable behaviour was a Cautious Alert
posture in the subjects. Most subjects maintained this
posture while slowly swinging their body or head or both
through an arc of about 180 degrees. Subsequently some
of the subjects began walking about the cage while main-
taining the extended neck component of the posture. This
went on for several minutes until a subject popped, which
resulted in the group of birds eXploding into a flurry of
pOpping and flying quail. This would abruptly cease and
all the subjects run to a cage wall and pile into the
Huddle-Heap. Very soon one or two birds would venture out
from the huddle and start walking across the cage, again
in the cautious type posture, and soon the other birds
would leave the huddle and traverse the cage until one
bird would pOp and the cage would again eXplode and return
to the Huddle-Heap Structure. One of the dramatic aspects
of these huddles was the evident Sustained Tactual Contact
posture observed here; the pushing component of that
pattern was so forceful that birds on the interior of the
huddle were continually being squeezed to the top of the
pile. These birds would then begin again at the floor,

also pushing against their cage mates in the huddle. This

87

resulted in a continually revolving group of birds and was
the characteristic form of the Huddle-Heap seen in this
group of subjects for the next few days. The carry over of
this pushing component into the huddle structure was not
the only incidence of the contact pattern observed in the
newly assembled birds. Another pattern observed in these
birds that was quite similar was what I began calling a
"safe" position and involved a bird or two or three birds
exhibiting the contact type pressing against a surface
but with their head stuck into the corner between the cage
wall and one of the troughs. Within the first week the
Huddle and Spacing Group Resting Structure also began to
show up in this group. It should be pointed out here that
these were the only two Group Resting Structure patterns
observed in this group. The birds had the same environ-
mental living conditions as the other, socially reared,
group and remained together until they were 75 days old
and yet were never observed to form either the Linear,
Ring of 1/2 Circle formations that were a regular aspect
of the other group's resting behaviour throughout this
period under these conditions.

Within the class Locomotion, the frolic-with-
object made its first appearance in this group of
subjects on the combination day, as illustrated in Figure
14. Several of the other behaviour patterns that had

been observed in some of the isolates during Phase I,

88

but had not reached 100% onset during that time, were now
observed in the remainder of the group of birds after
their assembly. These patterns were: Dusting, POpping,
Frolicking, Crouching, Freezing, and the Cautious Alert
Posture.

The regularly scheduled observation procedure was
altered on the first day of Phase II to include a con-
tinuous two hour observation following assembly and two
other short observations the same day, one at noon and the
other at 3:00 p.m. followed by the regular evening
observation. Throughout all of these observations the
level of activity within the isolate cage remained high.
Feeding, drinking, general preening, running, flit-pOpping,
pOpping, flying, crouching, and generalized pecking were
observed in all of the subjects during each of the observa—
tion units mentioned above. Sleeping was not observed in
any of the birds until the noon observation and sophisti-
cated preening, oiling, frolicking, and bill wiping
remained infrequent until the 3:00 observation.

Approximately 30 minutes after the initial light
onset and the beginning of these birds' eXperience at
living in tactual and visual contact with conspecifics,
the cautious exploratory and huddling behaviour decreased
considerably and the birds began to move about the cage
independently and allOpecking immediately became the

predominate behaviour pattern for the group as they

89

started running wildly all over the cage pecking each
other. A pecking encounter was typically unilateral and
although the pecks appeared to be delivered with con-
siderable force there were no wounded animals at the end
of the day. It was often difficult to distinguish if a
bird that was being pecked avoided the pecker or if
another bird nearby provided a stimulus to approach and
the bird moved off to peck some other bird. Adding to
this confusion was the fact that very few pursuits were
involved in these pecking encounters. The name of the
game appeared to be "if a quail is anywhere near--peck it."
The pecking organization was equally multilateral with one
exception. There was one bird that was nearly always
alone and just generally didn't appear to be involved in
any of the group's activities. This bird seldom pecked
other birds and was seldom pecked by other birds but
didn't appear to avoid a bird that approached.

A new form of allOpecking that had not previously
been observed in the other group showed up toward the end
of the morning observation and although it was never very
frequent (a maximum of five encounters in one observation
unit) its tOpography was impressive. This was a pattern
called tail feather pulling and several times involved
the recipient being pulled off balance when it occurred.

This pattern occurred only sporadically in encounters

90

(from 0-3 in one observation) after day 30 to the end of
the study. Another allOpecking behaviour that exhibited
this same sporadic pattern of occurrence was the body peck.
When observed in these birds it also appeared to be
delivered with considerable force and often would knock
the recipient off his feet. However, this allOpecking
pattern did not appear until day 31.

Table 3 below lists the raw frequency of encounters
data for the allOpecking behaviour observed in these sub-
jects on the initial day of Phase II. By comparing this
with the data presented in Table 2 for the fifth week of
allOpecking behaviour in the socially reared group, the

constrast is readily apparent.

TABLE 3.--Frequency of allOpecking encounters observed in
the isolate reared group (N=7) on the day of assembly into
a social living unit.

 

 

Time of Observation Toe-Foot Head-Eye-Nares
6:00 - 7:00 a.m. ll 84
7:00 - 8:00 a.m. 16 106

12:00 -12:35 5 70
3:00 - 3:35 12 72

7:25 - 8:00 p.m. O 13

 

91

Given the generally high frequency of head pecking
throughout the day on day 30, the figure for the evening
observation is so low that one is tempted to surmise that
the birds were too exhausted to continue pecking, particu-
larly since the next morning's observation of this group
resulted in a frequency tally of head pecks of 91. This
high frequency of head pecking then began to flucutate
through the remainder of Phase II, ranging from 31 to 91
with a median frequency for the series of observations of
73.

Feet pecking remained low throughout the remainder
of the study with a range of frequencies from 0-12.

From the beginning of Phase III to the end of the
study, days 36-75, the incidence of head pecking encounters
fluctuated even more extremely than before, having a range
from 8-78, with a median frequency over this series of
observations of 39.

Comparisons Between Social
and’Isolate Groups

 

 

Comparisons between the two groups can be made
generally along three broad divisions: (l) Behaviour
patterns shown by subjects in both groups during the
period of differential rearing conditions, Phase I;

(2) Group or Social interaction patterns, or patterns that
appear immediately upon subjects being assembled into a

group; and (3) Behaviour patterns unique to a group of

92

subjects eXperiencing the same rearing conditions for the
first 30 days of age. These will be discussed in order.

1. Behaviours shown by subjects in both groups
during Phase I (not including those patterns prevented by
the imposed living conditions) included all the measured
classes with the exception of one locomotion pattern that
will be discussed later. The general patterns that are
evident upon examination of Table l are: (a) the later
median onset day for the Isolate group subjects in given
behaviour category with (b) a correspondingly greater
variation of range values (compare column 1 to column 4)
for that behaviour category. In the Care of the Body
Surface class of consummatory behaviours the Sophisticated
Preening is the only exception to this pattern (see Figure
17).

The class of Postural Patterns, Generalized Pecking
behaviour, and the appearance of the cuticle shafts for
both wing and tail feathers follows this pattern. In the
Locomotion class, the pattern of a greater variation in
range is evident throughout; however, the median onset day
is earlier for the isolates than the social group in the
jumping and flying categories but is identical for the
running category. Figures 2, 3, 5, 6, 9, ll, 14, and 18-22
illustrate these two patterns visually for the above

mentioned classes of behaviours shown by both groups.

93

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99

2. Group or social interaction behaviours, or,
patterns that appear immediately upon assembly into a group
included the following: the frolic-with-object pattern
of Locomotion behaviour. As defined in the measurement
section, the frolic pattern is an obviously related com-
ponent of the frolic-with—object pattern and as shown in
Figure 23 the frolic pattern did occur in the isolate sub-
jects during isolation. This makes it curious that the
frolic-with-object pattern does not show up in the isolate
group, see Figure 14, until after assembly. This would
suggest that while isolation may not prevent this pattern
neither does it encourage its eXpression. Since there
was not a control group of birds that remained isolated
during Phases II and III it is impossible to rule out the
possibility that age can be a factor. However, the
appearance of this pattern immediately following assembly
would seem to support the suggestion that this is a social
behaviour and thus requires the presence of at least one
other animal to initiate the pattern, or to function as an
elicitor to signal the appetitive components of the inter-
action. Previous work (Ratner, 1965) showed that the
pattern would occur in chickens isolated for 70 days, but
only occurred in four out of fourteen birds.

Figures 5 and 17 show the comparative onset of
sophisticated preening and of allOpreening in the two groups

of quail. The close correspondence of development of this

100

 

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101

pattern within the groups would suggest that a maturational
view of this data might be a useful interpretation of these
results although the data on feather deve10pment (see
Figures 7 and 24), would not necessarily support that
interpretation.
The Resting Group Structure patterns, shown in

Figure 8, illustrate that apparently the first formation E_“
of resting postures in a newly assembled group are the i
more or less undifferentiated huddle type. As pointed out

previously, it is interesting that while the isolate group

 

had the same environmental conditions, in terms of light E-
and heat, they did not develOp the typical bobwhite
resting group formations. The data from the social group
in this case point out that it is apparently not necessary
to be raised in a nest with the parent quail present, for
the Species-specific nesting or roosting formation to
develop in a group of bobwhite young.

The differences in allopecking behaviours, as
shown in Tables 2 and 3, illustrate one of the more
striking differences in the occurrence of the same pattern
in the two groups. However, considering the natural
history of this species and the age of these subjects,
it is not necessarily surprising that a definite dominance
order was not established in either of the groups under

these living conditions.

102

 

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convoy AHHMHoom CH mummnm Hngmmm HHmu mo moqmummmmm HMHHHGHII.HN mhsmHm

mom yo mama
mN mN ¢N NN ON mH mH vH NH 0H

.o

.h

mumHOmH . .VH
.HN
Hmfloom 0 .mm
.mm
.mv
.om
.hm
.vm
.Hh
.wb
.mm
.mm
.OOH

dues 18d aArqetnan

 

103

3. Behaviour patterns unique to one or the other
of the two groups included the following: (a) those unique
to the isolate subjects included Cubicle Pacing as a
predominant form of daily activity during isolation (see
Figures 15 and 16), and the Sustained Tactual Contacting
that contained components that carried over into the other
phase of the study, adding a related component during
Phase II, that of the "safe" position; the deve10pment of
two forms of allOpecking that differed vastly from the
forms observed in the social group; the ritual-like
stereotopy patterns that were observed during isolation;
and the fragmented nature of the tOpography of several of
the maintenance patterns, specifically those of bill
wiping, oiling and the correspondence of oiling with the
sophisticated preening pattern. (b) The patterns unique
to the social group included the previously mentioned
Resting Group Structures of Linear, Ring, and 1/2 Circle
formations; the smoother and more coordinated patterns of
the Care of the Body Surface activities, and the corre-
sponding higher frequency of occurrence of these behaviours
as shown in Figures 3 and 4; and perhaps most striking is
the apparent synchrony of the individual subjects within
the group when engaged in various of the measured behaviour
patterns but particularly the Postural Patterns.

To conclude, further work should attempt to

delineate the parameters of isolation for a social species

104

such as the bobwhite. Because it is a non-domestic species
it should be a good preparation for probing what variables
are involved in signaling group behaviours and perhaps
skied some light on whether or not visual and tactual
isolation without removal of conspecific auditory stimula-
tion is a stressful condition that serves to facilitate
the appearance of competing response patterns in isolated f

conspecifics that normally live in a group.

C“

 

 

D I SCUSSION

' The present study, in contrast to much of the
previous laboratory work on behaviour deve10pment in
precocial galliformes, had as one of its explicit objectives
to provide a laboratory controlled environment for the
subjects which more closely approximated natural living
conditions. Specifically, observations involved minimal
human contact between the subjects and the experimenter
and regular diurnal light cycles were provided by simulated
daylight or warm white illumination. In accord with the
focus of the study an emphasis was placed on identifying
and describing behaviour patterns as they appeared in
these subjects in an attempt to provide baseline data on
the behaviour of this species living under these laboratory

conditions.

Behaviour of the Social Group

 

Six classes of behaviour patterns were identified
and described for the group of birds reared together. The
sequence in which the components of a behaviour class
develOped was defined in terms of the median onset day
for the group. For a summary of this schedule of develop-

ment see Appendix A. The present findings suggest that

105

 

106'

certain behaviour patterns are a useful index to the age
of the bird. As the young quail matures more complex and
elaborate forms of activities become a part of his
behavioural repertoire. The schedule of deve10pment of
the behaviour classes follow generally the report given
by Stoddard (1931).

It is not surprising that several differences were f“
found in the behaviour patterns of this species compared .
to the domestic fowl and the Japanese quail. In contrast 1
to domestic fowl, the bobwhite is a natural gallinaceous

bird; in contrast to the Japanese quail, it has a longer

 

rum. i \‘JU

period of deve10pment before reaching maturity, and it is
a species that is non-territorial and is not known to
fight in the wild except during the mating season
(Errington, 1968; Stokes, 1967; Stoddard, 1931).

Where comparisons of the data between the group
and isolated birds is possible, both groups of subjects in
the present study showed onset of Care of the Body Surface
activities earlier than that reported for the domestic
fowl by Ratner (1965) but evidenced a later onset for the
comparable Locomotion categories of frolic and frolic-
with-object. Both Guhl (1958) and Ratner (1965) report
sparring and fighting occurring within four weeks of age
in the domestic chicken with corresponding developments

of dominance hierarchies. In addition to this Ratner

107

reports that bilateral pushing at the food tray develOped
into food-dominate and food-submit patterns of interactions
between birds and that threats and juvenile fights were
observed within this period. Although aggressive allo-
pecking developed early in the bobwhites and bilateral
pushing was observed; threats, fights, dominate and submit
patterns as described by Ratner were conspicuously absent
from their patterns of behaviour. It is also of interest
that the bobwhite in this study were never observed to
suffer wounds as a result of the allOpecking that did
occur, whereas in another study of bobwhites (Borchelt,
1970) several subjects were severely wounded from pecking
encounters. However, differences between the two studies
in the lighting cycle, access to food, space available per
bird, and handling of the subjects may well make direct
comparisons impossible to interpret in this instance.

In a deve10pmental study of the behaviour patterns
of social and isolate Japanese quail Farris (1964)
described many of the same behaviour patterns observed in
the present study. ‘Behaviour patterns included in his
descriptions that were apparently similar in tOpography
for these two species were the following: Resting
postures, Generalized pecking, Competitive pecking,
AllOpecking, Freezing, Alert, Escape scattering or fleeing,
General preening, Bill wiping, Dusting, Feeding, Drinking,

Walking, Running, Jumping, Popping, Flying, Frolic, and

 

108

Frolic-with-object. With the exceptions of Resting and
Nutrition which were the same, and Popping and the Alert
posture which were earlier for the bobwhite; the socially
reared Coturnix showed an earlier median onset of the
patterns listed above. The isolated Coturnix showed earlier
median onsets for all of the patterns in comparison to the
isolate group of bobwhites in the present study. Farris F‘=
reports that in general there were few differences between E
his two groups in age of onset in the above behaviour

categories and the differences found are attributed to

 

differences in the available living space for the two E
groups. In contrast to this the birds in the present
study were not subject to comparable living Space
restrictions, for example, each isolate subject had suf-
ficient space to fly and did make brief flights within
the isolate cubicle. Therefore, available living space
is not felt to be of primary importance relating to the
differences observed in these two groups of bobwhite.
The Coturnix is a species whose natural history differs
considerably from that of the bobwhite; they are highly
territorial, do not live in coveys, show minimal care of
the young, and are sexually mature in 28 days. Since
there are not only these species differences but also
methodological differences between Farris' study and the
present study direct comparison of the data is difficult

to interpret.

109

The synchrony of the behaviour of individuals in
the social group, the general level of activity, and the
daily routine of the social group appeared to be basically
similar to that described by Stoddard (1931) for wild
groups of bobwhite. Specifically there were early and
late feeding bouts, no fighting, and several sessions of
Resting and Care of the Body Surface activities observed m1
between these early morning and late afternoon-early
evening feeds. The present results suggest that laboratory

conditions do permit the controlled observation of the

 

behaviour of this species without severe disruption of

natural and species-specific behaviour patterns.

Behaviour of the Isolate Subjects

 

During isolation the general behavioural deve10p-
ment of the birds in this group paralleled that observed
in the socially reared quail in that they showed the same
behaviour patterns (with a few exceptions). However, the
median onset was typically at a later age for the isolates.
These results are generally in agreement with other
investigations of the effects of isolation in the avian
galliformes. Ratner (1965), similarly found that the
isolated birds did exhibit the same individual behaviour
patterns in isolation that socially reared fowl exhibited

but showed them at a later age.

 

110

However, the birds in this group also exhibited
some patterns unique to the isolated subjects. In
particular, the Cubicle Pacing Pattern and the Sustained
Tactual Contacting pattern were conspicuous and carried
over into their behaviour after they were assembled. The
pervasive nature of these patterns and the consistency
with which they were seen as an integral part of the daily
routine of these subjects, and their absence from the
behaviour of the social group, suggest that tactual and
visual contact during the first month of age is of major
importance in the behaviour of the bobwhite quail. In the
present study the relative frequency with which these
patterns were engaged in by the isolate subjects, compared
to other behaviour patterns, such as Care of the Body
Surface behaviours suggests that it might be useful to
consider how the pacing and contact patterns may have
interferred or competed with the other classes of behaviour.
That haptic stimulation from conspecifics is important to
the normal deve10pment of the bobwhite is further suggested
in these data by the absence of the linear, ring or half—
circle group resting structure in the isolate birds even
after having been assembled.

The behaviour of the isolate birds immediately
following their assembly into a social living environment
was very similar to that described by Ratner (1965) for

the parallel condition in the domestic fowl and lends

111

further support to the idea that tactual and visual isola-
tion for a contact species of the galliformes is a powerful
behavioural variable. In addition to this, the striking
synchrony of group behaviour patterns observed in the
socially reared subjects and its continual absence in the
isolate group is a dimension of organization that should
prove useful as a variable with which to assess the effects 7‘5
of isolation. f
To conclude, the results of the present study,

taken in conjunction with those of other studies on the

 

galliformes, indicates that isolation may retard but does t
not prevent the deve10pment of maintainance and social
behaviour patterns in the isolated subjects in general,
with two exceptions: (l) the species-Specific Resting
Group Structure or roosting formation of the linear, ring
or half-circle pattern, and (2) the group synchrony of
Postural patterns such as freezing and the cautious alert.
These patterns never appeared in the isolate group subjects
in the laboratory even after having been combined into a
social group and living together 45 days. However, before
further studies undertake a comparison of social versus
isolate rearing conditions as a determinant of later
behavioural development, the author feels that a parametric
study of isolation in this Species is necessary to provide
the basis for making adequate interpretations that rest on

a functional classification system. As indicated in the

112

literature (Stokes, 1967) the bobwhite is a highly

auditory animal. in addition, the present results indicate
that haptic stimulation may prove to be a critical compo-
nent in the deve10pment of this bird. Further, it is
generally accepted that avians are also very dependent

upon their visual systems. Thus future research is

planned to attempt to separate these three forms of fax
stimulation in an attempt to deliniate what are the 1
actual variables being manipulated, in terms of the

sensory systems of the animal, in an isolation study.

 

1w

Summary

The present study provided an initial description
of several of the behaviour patterns of the very young
bobwhite quail raised in the laboratory under the condi-
tions described previously. The data collected confirm
the reports offered by Stoddard (1931) of behaviour pat-
terns of this species in a natural environment. The
results obtained extend and support previous findings on
the effect of visual and tactual isolation in a member of
the gallinaceous birds in that these isolation conditions
may retard but do not prevent the development of those
behaviour patterns typically measured in this group of
animals.

In addition to the above, this set of observations
provided descriptions of several behaviours previously

unreported for this species reared in the laboratory and

 

113

attempted to characterize their patterns. These include
the following: (1) distinct forms of preening, general,
SOphisticated, and oiling; (2) components of the dusting
pattern occurring on a bare wire floor and its possible
relation to the intensity of thermal stimulation; (3) the
age of onset of crouching as a Postural pattern in this
Species; (4) the Cubicle Pacing and Sustained Tactual Con- ran
tacting patterns as a persistent behaviour pattern of the f
isolated subjects (Farris, personal communication) pre-

viously observed but not described, measured or reported;

 

(5) an attempt to characterize the synchronous versus the

[I
l

fragmented nature of the tOpography of the group behaviour
patterns as they differed for the two groups; and (6) the
progression and deve10pment of components of the "roosting
disk" Resting Group Structure for which the bobwhite is
famous for to sportsmen and naturalists throughout the
United States.

Throughout this report an emphasis is placed on
describing the behavioural repertoire of this species
maintained under these laboratory conditions in an attempt
to add to our understanding of the behaviour of a non-
domestic animal and one which offers considerable promise
as an excellent laboratory preparation for studying several
of the classes of consummatory behaviours and the variables

determining the form in which they are likely to occur.

BIBLIOGRAPHY

114

25.32.!

 

31.7

BIBLIOGRAPHY

Bent, A. C. Life Histories of North American Gallinaceous
Birds. New York: Dover Publications, 1963.

 

Bolles, R. C., and Woods, P. S. The ontogeny of behaviour
in the albino rat. Animal Behaviour, 1964, EEJ
427-441.

 

Borchelt, P. L. DevelOpment of fear displays (immobility)
in the bobwhite quail. Unpublished masters thesis,
Michigan State University, 1970.

IA} 1.1::

Collias, N. E. An ecological and functional classification
of animal sounds. In W. E. Lanyon and W. N. Tavolga
(Eds.), Animal Sounds and Communication. t
Washington, D.C.: American Institute Biological
Science, 1960.

 

Denny, M. R., and Ratner, S. C. Comparative Psychology.
Homewood, Ill.: Dorsey Press, 1970.

Errington, P. L. Of Predation and Life. Ames, Iowa: Iowa
State University Press, 1967.

Farris, H. E. Behavioural development, social organiza-
tion, and conditioning of courtship behaviour in
the Japanese quail, Coturnix coturnix Japonica.
Unpublished doctoral dissertation, MIChigan State
University, 1964.

Garreffa, L. F. Group membership and group size as
determinants of aggressive behaviour between pairs
of male bobwhite quail (Colinus virginianus).
Comm. Behav. Biol., 1969, Ali, 69-72.

I

 

Guhl, A. M. The deve10pment of social organization in the
domestic chick. Animal Behaviour, 1958, 6, 92-111.

 

Guhl, A. M. Social behaviour of the domestic fowl. Tech.
Bull., 1953, 13, Kansas Agricultural Exp. Station,
Manhattan, Kansas.

115

116

Hartford, A. The behaviour development of group-raised and
isolation-raised female domestic fowl. Unpublished
masters thesis, Michigan State University, 1967.

Kirkpatrick, C. M. Factors in photoperiodism of bobwhite
quail. Physiological Zoology, 1955, 28, 255-264.

Menaker, M. Biological clocks. BioScience, 1969, 19,
no. 18.

Nelson, J. E. Behaviour of Australian PterOpodidae
MegachirOptera. Animal Behaviour, 1965, 13, 544-557.

 

Nice, M. M. DevelOpment of behaviour in precocial birds.
Trans. Linnaean Soc. of New York, 1962, 8, 158-169.

Ratner, S. C. Comparisons between behaviour deve10pment
of normal and isolated domestic fowl. Animal
Behaviour, 1965, 13, 497-503.

 

Rosene, W. The Bobwhite Quail, its Life and Management.
New BrunswiCk, N.J.: Rutgers University Press,
1969.

Schein, M. W. Dust bathing in birds. In A. W. Stokes (Ed.),
Animal Behaviour in Laboratory and Field. San
Francisco: Freeman, 1968.

Scott, J. P. Animal Behaviour. Chicago: University of
Chicago Press, 1958.

 

Spalding, D. Instinct. Macmillan's‘Magazine, 1873, 21,
282-293. Reprinted British J. Animal Behaviour,
1954, 2, 2-11.

Stoddard, H. L. The Bobwhite Quail. New York: Charles
Scribner's Sons, 1931.

Stokes, A. W. Behaviour of the bobwhite quail, Colinus
Virginianus. Auk, 1967, 84(1), 1-33.

 

Sturkie, P. D. Avian Physiology. Ithaca, New York: Cornell
University Press, 1965.

 

Vince, M. A. Potential stimulation produced by avian
embryos. Animal Behaviour, 1966, 14, 34-41.

WOod-Gush, D. G. M. The behaviour of the domestic chicken:
a review of the literature. British J. Animal
Behaviour, 1955, 3, 81-110.

 

 

APPENDIX

117

 

 

11

SEQUENTIAL APPEARANCE OF COMPONENTS OF THE MEASURED
BEHAVIOUR CLASSES SHOWN BY MEDIAN ONSET DAY FOR
TWO GROUPS OF BOBWHITE QUAIL

Social

 

NUTRITION

CARE OF BODY SURFACE
General Preening
Bill Wiping
Sophisticated Preening
Oiling
Dusting
AllOpreening

LOCOMOTION
Running
Popping
Jumping
Frolic
Frolic-object
Flit-popping
Flying

ACTIVITY
Resting
Huddle-Heap
General Pecking
Huddle-Space
Linear
Ring and Half Circle

POSTURAL
Freeze
Cautious Alert
Crouching

ALLOPECKING
Body
Head-Nares
Toe-Tarsus

FEATHER SHAFT APPEARANCE
Wing
Tail

15
17
19
36

NH
U1U'lrb-Ni-‘l-4

\ICDQ

12
12

Isolate

 

NUTRITION

CARE OF BODY SURFACE
General Preening
Bill Wiping
Sophisticated Preening
Oil Gland Preening
AllOpreening
Dusting

LOCOMOTION
Running
Jumping
Frolic
Popping
Flying
Flit-popping
Frolic-object

ACTIVITY
Resting
Cubicle Pacing
General Pecking
Huddle-Heap
Huddle-Space
Linear, Ring

POSTURAL
Sustained Contact
Freeze
Crouching
Cautious Alert

ALLOPECKING
Body, Head, Toe

FEATHER SHAFT APPEARANCE
Wing
Tail

15
24
32
36

16
20

 

 

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69 0120