Dominance Relationships and Agonistic Behavior of Canada Geese in Winter PDF

Dominance Relationships and Agonistic Behavior of Canada Geese in Winter Author(s): Dennis G.
Raveling Reviewed work(s): Source: Behaviour, Vol. 37, No. 3/4 (1970), pp. 291-319 Published by: BRILL Stable URL: http://www.jstor.org/stable/4533358 . Accessed: 26/01/2013 16:44
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DOMINANCE
RELATIONSHIPS AND AGONISTIC BEHAVIOR OF CANADA GEESE IN WINTER by

DENNIS G. RAVELING 1) 2)
Wildlife ResearchLaboratory, SouthernIllinois University,Carbondale, (Cooperative Illinois,U.S.A.)

(With 9 Figures) (Rec. 15-VII-197o)
INTRODUCTION The fact that many animals exist in close, intraspecific, competitive situations with regard to many essentials of life has stimulated a variety of studies and explanations of the functions and evolution of aggression and dominance orders. Studies of dominance in various species of geese
have been reported by JENKINS (I944), COLLIAS(I950a), HANSON (I953), BOYD (I953); LORENZ (I959, I966) and FISCHER (1965) mentioned
certain dominance aspects or implications regarding geese. Past studies were based on observation of captive or semi-captive birds, or on partially colormarked families or unmarked wild birds. This study is based on repeated observations of the same color-marked individuals and families of Canada geese (Branta canadensis) in the wild, free-living state during winter. Radiotelemetry techniques enabled locations, movements, and behavior of specific individuals to be recorded. This paper reports results of victories and losses in aggressive conflict situations and describes the postures associated with those results. An attempt is made to integrate an understanding of the function and evolution of agonistic postures in relation to the stable, functional dominance hierarchy exhibited by Canada geese. I) This studywas financed mainlyby the NationalScienceFoundation (GB-623)with
additional support from the CooperativeWildlife Research Laboratory,Southern Illinois Director. Many persons and agencies cooperatedin this University, Dr W. D. KLIMSTRA, work. I wish to thank again Messrs W. E. CREWS, W. W. COCHRAN, L. A. MEHRHOFF, offered helpful criticism of the manuscript.Results formed a portion of a Ph.D. dissertation presented to the Department of Zoology, Southern Illinois University. 2) Present address: Canadian Wildlife Service, I4-A Garry Street. Winnipeg I, Manitoba, Canada.
R. G. PERSONIUS, H. C. HANSON,and D. W. WARNER. and Drs W. D. KLIMSTRA, H. BOYD
292
DENNIS G. RAVELING
METHODS This study was conducted at Crab Orchard National Wildlife Refuge, Williamson County, Illinois where approximately 40,000 Canada geese (B. c. interior; cf. HANSON & SMITH, 1950: 77) spend a large part of the winter. The inviolate portion of the refuge encompasses 22,000 acres including 2,600 acres of Crab Orchard Lake where the geese roost and 5,000 acres of cropland (corn and soybeans) and 2,800 acres of pasture where the geese feed. During the winters of 1963-64 and I964-65, 77 geese were radio- and color-marked. These included all or parts of IO families, 2 pairs, and 35 yearlings. Color marking involved dyeing the white portions of the abdomen and cheek patches various colors (cf. BOYD, 1952; KOZLIK et al, 1959) and attachment of colored /4-inch nasal disks (LINDMEIER &
JOHNSON, 1958; BARTONEK & DANE, 1964). Observations of the activities
of the flock and of radio- and color-marked geese were recorded daily from late September to mid-March during the two winters of this study. Further description of Crab Orchard, details of the recognition and capture of families and other social classes of geese, their permanency, local roost and movement patterns and behavior, and discussion of the techniques of color-marking and radio-tracking are provided in RAVELING (I969a, b, c). Terms. Immature son-James
from hatching (June) until the return to nesting areas (HudBay lowland muskeg, HANSON & SMITH, 1950: 79) the
following April. Yearling - from the time of return to nesting areas for their second summer of life until their return to nesting areas the following spring. Adult - from the beginning of the third summer of life and older. Family - any association of two or more geese resulting from a pair bond or parentage-progeny-sibling relationships. Families of three or more represent an adult pair with their offspring or surviving members of such an association. Behavior was not affected by color-marking procedures (RAVELING, I969a). The transmitter was usually concealed in the breast feathers and held in place by a neck and a body loop. The radio caused more than normal preening on the breast area but did not affect movements, flight ability or participation in any of the activities observed to occur with unmarked geese
(RAVELING, i969a).
Since the history of the marked geese of this study before trapping was unknown, it is not possible to be certain that the young of these families
AGONISTIC
BEHAVIOR
OF CANADA
GEESE IN WINTER
293
were in fact hatched and raised by the adults they were in company with when trapped. However, the result was the same in that the study animals behaved as a unit in all their daily winter activities (see RAVELING, I968a, I969a-c). PART I DOMINANCE RELATIONSHIPS
For the present discussion distinction is made only between fights and threat-chase behaviors. Fights refer to an occurrence when two geese rushed at each other, grabbed each other's feathers at the base of the neck, rose erect, and beat each other with their open, bent wings. Besides superiority in a fight the other category of victories referred to here includes all forms of threat and chasing when these movements were linked with avoidance or active escape on the part of a submitting goose.
RESULTS
situation of aggressive behavior. ecological In general the geese at Crab Orchard roosted on the main lake during the night and the mid-day. They flew to nearby fields to feed in early morning and late afternoon. Lengths of feeding periods were extended when overcast sky conditions prevailed. Canada geese were gregarious in all their daily activities during winter but frequency and intensity of aggression fluctuated widely. Conflicts were most frequent at the roost lake in two situations: (a) just after geese arrived from their morning feeding period and were moving to and establishing their loafing areas, and (b) when geese became more active in the hour or two before flying out to feed in the afternoon. Conflicts were most numerous and intense during feeding activity in fields. General TABLE I Numbers of aggressive encounters between Canada geese observed in different situations (October 3-1I, 1964)
Number of 15 minute observation periods
9 2 5
Situation
Lake shoreline (midday, p.m.) VWheat
Estimated numbers of geese observed

(go - I6o)** i30
I25*
Numbers of threats and chases

23
Numbers of fights
0.4
(12 - 36) 50
(o - 2) 3
(a.m. feeding period)

Millet (a.m. feeding period) * Average, ** Range.
(IIo & I50)

I 5 (Ioo - I30)
(44 & 55)

I17 (74 - I56)
(2 & 4)
4 (2 5)
294
DENNIS G. RAVELING
To compare the magnitude of conflicts in different situations, approximately I00-I50 geese were isolated in the field of view of a spotting scope and all attack-fleeing incidents observed in 15 minute periods were noted. These data demonstrate that more conflicts occurred when geese were in an area where food was concentrated (millet) than when it was evenly dispersed (wheat), or absent (lake shoreline) (Table I). Even greater frequency and intensity of encounters among the geese were observed when they were feeding in corn fields but it proved virtually impossible to keep track of a relatively constant number of geese and record all conflicts. While not quantitatively documented, the above relationship of frequency of conflicts prevailed all winter. Relationship rank order. of social status to aggression and
The opportunity to observe different families and single geese of known age and sex made it possible to compare success of these animals in aggressive situations. Data presented in Table 2 are pooled results from all radio-marked TABLE 2 Results of observations of 516 aggressive encounters involving radio- and color-marked Canada geese
Social status Single immatures Single yearling females
Single yearling males
N 3 9
Io
Total number of conflicts 20

29 64
Number of victories o (o%) 5 (17%)

i6 (25%)
* *
(2I%)
Single adult female

Single adult males Total all singles Pairs
I
3 26 6
9
I3 135 39
I
7 29
I2
(3I%)
Families of 3 Families of 4 Families of 5

Family of 6
7 7 4
I
57
92
32
(56%)
I8i
12
69 (75%) I49 (82%)

9
*
* Percent not calculatedwhen total conflicts <20.
geese. These data, therefore, are totals derived from observations of relatively few conflicts of each individual goose or family. The small samples did not allow statistical comparisons of variations of success within most categories. However, for families of five and four, Chi-square tests for two independent
samples (SIEGEL, 1956: IO4-I07) indicated non-significant differences in
ratios of success in encounters of different families within each category
AGONISTIC
BEHAVIOR
OF CANADA
GEESE IN WINTER
295
TABLE 3. Rank order of five, single, captive Canada geese as determined by threat and peck victories during six hours of observation
Losses Bird no. A A 3-year old or older male X B 8 C
II
D 16
E 7
B*
2-year old X 6 38 4
male
C* 2-year old male X 10 9
>
D*
2-year old X 35
female
E*
2-year old female * Age known because geese were trappedprevious year as yearlings. X
that were combined to make up totals in Table 2 (X2 - 4.94, 3 d.f., p>o.Io; x2 -= 0.32, 6 d.f., p>o.IO, respectively for families of five and four). Numbers of different individuals or groups contributing to totals in Table 2 vary from the actual number of individuals or families that were radio-marked. The same individuals may be represented in observations of different sized groups because of temporary separation of family members or deaths. Using observations of an adult pair and one immature as a family of three even though there were two other offspring that were separated at the time of a particular observation is believed to be valid as dominance position is directly and immediately related to numbers of individuals in a
family at any one time (JENKINS, I944; COLLIAS, I950a; BOYD, I953: IIO, FISCHER, 1965: 270; see below).
Results of aggressive encounters and defeats (Table 2) reveal that success was related directly to family size. Data for single geese indicate considerable differences of success in conflicts depending upon age and sex. Single immatures were the most submissive geese and rarely initiated an encounter, but were often continuously "busy" avoiding other geese. Among geese held
296
DENNIS
G. RAVELING
captive during this study single adults dominated single yearlings and individuals in both these age classes dominated single immatures. Within any age class, single males were dominant over single females. Dominance order was quantified as to victories and losses for five unrelated, single geese that had been held in captivity for a year (Table 3). in aggressive encounters. participation Intrafamily Identification of unmarked families depends upon unity of family action in Triumph Ceremonies (exaggerated head and neck motions often with
raucous honking see FISCHER, I965; LORENZ, I966; RAVELING, I967),
attack-escape situations, and "purposeful" movements. While this procedure often yields correct interpretations, chances for error are considerable. It was relatively rare that an entire family entered into aggressive postures or all members came together in a Triumph Ceremony. Of 325 encounters involving marked pairs and families in which the activities of all members of TABLE 4 Interrelationships of members of pairs and families of radio- and color-marked Canada geese with respect to initiation and participation in victorious conflicts
Group and number of victories * Pairs Io Families of 3
25
Individual of group Adult male Adult female Adult male

One of the others Adult male
Number of victorious conflicts initiated 6 (6o%) 4 (40%) I9 (76%)
Number of victories in which all members participated 3 (30o%)
I 6 (25%) 45 (71%)
(4%)
Families of 4
63 Families of 5
I49
One of the others Adult male

One of the others
5 (8%) I8 (29%) 112 (75%)

12 37 (25%) (8%)
Totals 247
Adult male One of the others
I82 (74%)
21
(8.5%)
65 (26%)
* Numbers of conflicts may vary from totals presented elsewhere; data in this table represent conflicts in which activity of all members of group were recorded.
AGONISTIC
BEHAVIOR
OF CANADA
GEESE IN WINTER
297
the family were recorded, only 33 (ca IOpercent) involved unified action of all members of the family (Tables 4 and 5). The greater the intensity of an encounter the greater was the unity of family participation. Often a goose (or geese) submitting to one or more threatening members of a family appeared as a single or small family but actually was a portion of a larger family. For example, in 32 percent of the losses of marked pairs and families only the adult male received or noticeably reacted to a threat or chase (Table 5). Unified action of all members of a family occurred twice as often in defeat as in victory (Tables 4 and 5). TABLE 5 Interrelationships of members of pairs and families of radio- and color-marked Canada geese in losses to more dominant geese
Group and number of losses * Pairs
II
Individual of group reacting as loser in conflicts Only adult male Adult female Both Only adult male One or both of the others
All 3
Number of losses 2 (I8%) 5 (45%) 4 (37%) 5 (33%) 8 (53%)

2 (13%)
Families of 3
I5
Families of 4 20 Families of 5
32
Only adult male One or more of the others All 4 Only adult male
One or more of the others
6 (30%)
II (55%)
3 (i5%) 12 (38%)
17 (53%)
All 5
Totals Only adult male
3 (9%)
25 (32%)
78
One or more of the others

All members of a group
41 (53%)
12 (I5%)
* Numbers of conflicts may vary from totals presented elsewhere; data in this table represent conflicts in which activity of all members of group were recorded. I sometimes observed an immature of a family avoid another threatening goose (or group) while the rest of its family made no move to attack or avoid the aggressor. Sometimes the gander of a family would chase a goose (or group) that had threatened one of his family. Immatures of a family could be as successful as their gander at chasing away other geese, even adults and families, if the immature was near its gander. Some defeats indicated for families in Tables 2 and 5 were a result of an Behaviour XXXVII I9
298
DENNIS
G. RAVELING
immature being threatened while the gander did not respond so that the percentage of victories in Table 2 under-emphasizes the high success of family ganders in dominating other geese. Ganders were involved in 82.5 percent of victorious aggressive encounters (Table 4, initiation plus participation) but reacted directly in only 47 percent of defeats (Table 5, reactions of adult male only plus reaction by all of a family). The numbers of young in a family were directly related to dominance position, but rank order was most clearly defined by the action of the gander in dominating or submitting. All groups had a higher success ratio in encounters in which the gander was involved (Table 6) as compared to all conflicts combined regardless of which family member was involved (Table 2). TABLE 6 Percent success of aggressive encounters involving the adult male of radioand color-marked pairs and families of Canada geese of and number Group conflictsin whichadult malewas involved Pairs
I5
Numberof victories 9 (6o%)

i9 (70%)
Families of 3
27
Families of 4 59 Families of 5 I39
50 (85%) 124 (89%)
Dominance of fighting
relation between
as determined from observations families. of unmarked ganders
Participation of all members of a family in an aggressive encounter was directly related to the intensity of display and failure of threatening postures to cause avoidance by another individual or family. In 516 encounters (Table 2) of marked geese, fighting occurred only I4 times (2.7 percent). Nine of these 14 fights involved the gander of a family of five, two fights each involved ganders of families of four and three, and one the gander of a pair. Of 33 occasions in which all members of a marked family were distinctly united in attack and/or escape, 14 of these observations occurred during the fights described above. Intense Triumph Ceremonies involving all members of the victorious family followed the most vigorous chases and fights. Thus, the most reliable way to identify numbers of geese in unmarked families is to observe two groups displaying prior to, and the victors after,
AGONISTIC
BEHAVIOR
OF CANADA
GEESE IN WINTER
299
the most vigorous aggressive encounters. The outcome of 26 such fights observed in I964-65 between ganders of unmarked families is presented in Table 7. These results reveal only one deviation from the expected occurrence of the largest family always winning. TABLE 7 Relationship between size of family and victories in 26 fights between ganders of unmarked families lof Canada geese Losses Familysize*
8
8
7 6
5
4
7 X
5 I
3
2
I
3
5
2
2
2 I
3 2
I I
* Size of familiesidentified because of high intensityof displays beforethe fights. Effects of separation of family members on dominance position. When one of the marked immature males of a family of six was separated on one occasion he became a very submissive, avoiding goose. He was threatened and pecked I times and he never threatened other geese. Prior to being separated and after being reunited this individual was one of the most active and aggressive immatures seen in this study, but only in the presence of his family. A family of four had been observed in 43 wins and 15 losses in aggressive encounters. When an immature was separated the remaining three were observed in four victories and five losses. Too few encounters of other families affected by separation were observed to allow statistical comparisons. However, the changes in general disposition were striking and conclusive in demonstrating the effects of separation and reunification on dominance. The change was best observed in a highly aggressive family of five at a time when the adult pair and one immature were separated from the other two young. The gander then avoided geese and was less alert. At a point when
300
DENNIS
G. RAVELING
a fight seemed inevitable this gander turned away and the family of three was put to rout by a gander of a family of four. No family of four was ever observed to defeat this family when they were intact. The next day one of the separated immatures rejoined the family and the increase in activity and aggressiveness was notable; the gander was observed making nine successful attacks and in the most intense encounter he thrashed a gander of a family of four. Finally, the third immature was reunited and the family resumed its previous high position in the rank order. Aggressive conflicts per unit time.
Data provided by radio- and color-marked geese clearly demonstrate that families of four and five were involved in the most conflicts (Table 8). There are no significant differences in the rates of conflict of all categories of singles or among singles, pairs, and families of three (X2 = .93; 4 d.f., p > 0.70; X2 = 0.28, 2 d.f., p > 0.80, respectively). Families of four were involved in more conflicts per unit time than families of three (X2 = 5.22, i d.f., p < 0.05) but there was no difference between families of four and five (X2 = 0.63, I d.f., p > 0.80). Families of four and five were involved in approximately twice as many encounters as smaller groups and singles. Since most conflict observations were obtained when geese were at the roost lake the ratios are probably biased as actual conflicts per day would probably show much more than a twofold increase for large families if more feeding period conflict records were available (cf. Table I). TABLE 8 Aggressive encounters per unit time observed to occur with different social status radio- and color-marked Canada geese. Number of
Group Single immatures
Single yearling females
Lengthof time of observations

(hours) i8.6
I7.4 48.2 10.2 12.0 o06.4
Conflictsper
hour
I.I
conflicts
20
29
Single yearling males

Single adult female Single adult males Total all singles Pairs
64 9 13 135 39
1.6 1.3
0.9 I.I 1.3 1.2
33.8
Families of 3
Families of 4 Families of 5 Family of 6
57
92 i82
12
49.9
43.6
72.0
I.I
2.1 2.5
34
3.6
AGONISTIC BEHAVIOR OF CANADA GEESE IN WINTER
301
Exceptions
to the usual
dominance
hierarchy.
No pair formation was known to have occurred among any of the marked geese. However, behavior was observed that I interpreted as preliminary to pairing ("courtship") and as proof of the culmination of pair bond formation among some unmarked geese. Such occurrences were rare and took place in February or March just before spring migration. Descriptions of postures involved are provided elsewhere (RAVELING, 1967: 57-6I). While it differs in some aspects of form and time relations as compared to greylag geese (Anser anser) I believe the essential functions and results with regard to enabling association and effects in social relations and dominance are as
described for greylags by HEINROTH (I9II: 622); LORENZ (I959, I966),
and especially FISCHER(I965). Many ganders of large families seemed to avoid fighting with ganders of newly formed pairs. Often a family gander was observed to start Headpumping and Rolling (Agonistic associated display - see Part II) but the newly mated male did not usually stop and display, or when he did it was very brief. Instead he went into attack almost immediately and the family gander turned away from this onslaught. Other exceptions to the usual dominance order occurred in "artificial" situations. The normal pattern of family members remaining close to each other while feeding with the gander performing many threats and chases was altered at baited trap sites or when corn was spread to supplement food availability for the geese. In those situations food was concentrated and large numbers of geese attempted to feed in a relatively small area. Initially the ganders of families engaged in enormous amounts of hostile behavior but often geese continued to move toward the food and "family-defense" became physically impossible. When this happened, geese crowded body to body and aggressive behavior was reduced most commonly to vigorous pecking alternating with attempts to eat. Family unity was almost always disrupted. Intrafamily aggression.
& JAHN,1959) but Downy siblings may fight among themselves (COLLAS when the Triumph Ceremony display becomes fully coordinated, attack among
siblings is inhibited (FISCHER, 1965: 264). LORENZ (1935) commented on
the compatability of young geese within a family without there being a rank order. The following data were derived from 169 hours of observation of marked families during which the activity of each family member was recorded. Adult males were seen pecking one of their young on four occasions, and adult
302
DENNIS G. RAVELING
females pecked one of theirs on three occasions. Pecks between siblings were observed six times. Pecks within a family were less forceful and not followed by escape or attack behavior which occurred when non-family birds were pecked. Intrafamily pecks occurred when one individual moved its head right next to the other when feeding, although this often happened without resulting in pecking. These pecks were of very low intensity and, except on one occasion, the pecked goose either swung its head away slightly or never moved. The behavior of the pecking goose was ambivalent, as exhibited by intention movements of pecking and withdrawing, or very rapid withdrawal and hesitation after a peck had been delivered. It was clear that aggression within the family was effectively inhibited. DISCUSSION investigations. noted that geese are mostly grazers and that manifestations of social rank seem to be absent in the wild but develop in confinement where food is localized. However, conflicts associated with rank position were noted during all aspects of daily activities of the geese during this study, i.e., while grazing or even "loafing" during periods of relative inactivity. Since victories in aggressive encounters were related directly to family size, this order should correspond to the dominance hierarchy of Canada geese, i.e., large family > smaller family > pair > single. JENKINS (I944) reported that families dominated pairs and that single geese were lowest in a rank order observed in a partially captive flock of blue and snow geese (Anser caerulescens) and Canada geese. For B. c. interior, HANSON (I953) concluded that the rank order was family > pair > unmated adults and yearlings; BOYD (I953: Ioo) noted the same in white-fronted geese (Anser albifrons) as well as an increase in success of the largest families. Data presented here suggest a rigid rank order in which a family's position is directly influenced by the total numbers in the family and that a difference of even one in family size is highly influential in their rank relations. BOYD (1953: 93, IOO) found similar relations in white-fronted geese, but his data on victories and losses of various social status families reveal many instances in which supposedly larger families were dominated by smaller groups or even singles. The discrepancy between the rigidity of the rank order observed in this study and the greater frequency of smaller families dominating larger families in BOYD'S(I953) study is probably explained by a bias introduced in my
DELACOUR & MAYR (1945: 8-io)
Comparison
to other
303
method of capturing the animals. Cannot-net traps (DILL & THORNESBERRY,
I950) were used to catch the geese but in locations where only a few geese roosted. Conflicts over a small amount of bait in front of the net eventually left only one family in position to be trapped by virtue of the intensity and success of a family's gander in keeping all other geese away. The suggestion is that only highly aggressive ganders and their families were caught and marked. Subsequent research on large samples of color, neck-banded geese taken from trapping efforts during the molt or large cannon-net catches has revealed a large range of variation in aggressive inclinations by individual ganders regardless of family size (RAVELING, unpublished data). While the general theme of family size as the most important correlate and probable determinant of success in hostile encounters is still true, and still especially noticeable in ganders that do much chasing and fighting, there are some family ganders who rarely engage in this activity. A further correlate of a lack of aggressiveness by some ganders is a more loose family association where family members repeatedly separate and rejoin (RAVELING, unpublished data). In contrast, the families in this study were together on 96 percent of all observations (RAVELING, I969a).
Other factors which could partially explain the difference between BOYD'S (1953) study and this one in numbers of smaller families dominating larger families is the lack of marked birds in BOYD'S(1953) study. Because of this it would be difficult to record all subtle or low intensity actions observed as related to family size because united action of an entire family is relatively rare (Tables 4 and 5). Conditioning is often very important in influencing aggressive behavior (cf. SCOTT, 1958: IO-I7), but in geese both the rank of the remaining group and separated individual(s) of families are most dependent upon physical proximity of related birds (JENKINS, 1944; FISCHER, I965: 270; BOYD, 1953: IIO; COLLIAS,I95oa). The same result occurred in Canada goose families during this study when separation and reunification were observed. Establishment flock. and function of rank order in a large
Since the ability of geese to be victorious in aggressive encounters is related directly to the numbers of individuals cofa family and this dominance ability can change immediately upon separation and reunification of related geese, important questions arise about the mechanisms causing geese to be more or less successful? How do other geese recognize individuals and groups that are superior or inferior to them in fighting ability?
304
DENNIS G. RAVELING
In addition to causing dispersion of individuals, a major function of aggressive behavior and rank orders is to promote stability in a group. Fighting and hostile behavior leads to less fighting because an individual learns to recognize individuals which have defeated or can defeat him and those which he can defeat. Once this is learned, these geese can coexist without fighting because then intention movements of attack (threats) suffice to maintain the benefits of being superior (e.g., access to food, space) and constantly reinforce individuals' positions with respect to each other. Because of the size of many wild goose flocks and what superficially appear to be shifting movements or mixing, some workers have denied that individual recognition and rank order are important in limiting aggression. BOYD
(I953: III) concluded this for white-fronts and WYNNE-EDWARDS (I962:
136-138) pointed out the impossibility or impracticability of rigid rank orders in huge flocks of birds because of a finite ability to recognize "faces". Such conclusions apply only partially to Canada geese. Geese were not distributed in roosting and feeding areas in a random fashion (RAVELING, I969b). Habit was important in influencing the locations of geese, particularly families. The flock at Crab Orchard consisted of many subflocks within which the activity and use area of a particular family was limited. Families regularly used shoreline areas at the lake that contained only a few hundred geese and many of these must have been the same individuals. Since singles and pairs were so submissive and avoiding, a stable rank order could exist, for example, if the Ioo or 150 family ganders of a group of Iooo geese regularly using a particular roost site had learned to recognize each other. Geese can probably recognize dozens and possibly hundreds of other individuals. FISCHER (I965: 267) noted that even goslings could recognize large numbers of geese and even where these geese were in relation to their family in the rank order. While I believe that stable rank orders can exist based largely on individual recognition, behavior differences between individuals and families in large flocks are probably most important in allowing initial recognition of superiority. At Crab Orchard thousands of geese representing several subflocks often fed in the same field. A rank order depending completely upon individual recognition in these situations is not plausible. Great increases in intensity and frequency of aggressive behavior occurred while feeding (Table I). As compared to non-feeding periods, fighting increased approximately tenfold whereas threatening and chasing increased only two to five times. While these data are limited, it is believed that the disproportionate increase in fighting partially reflects the fact that many families unfamiliar with each other were brought together. I suggest that fighting in winter usually occurs only between ganders
305
which are unfamiliar with each other and have not established a dominance relationship. However, when thousands of geese were feeding in one area, the increase in fighting observed could not possibly have equalled the increase in the numbers of family ganders unfamiliar with each other. This fact also suggests that recognition of rank superiority was accomplished by some means other than individual recognition. Nineteen of the 26 fights reported in Table 7 occurred between ganders with families of equal size or a disparity of one. These data further substantiate that many geese must recognize superior families since fighting usually occurred among ganders which were essentially equally aggressive. In other words single ganders did not fight family ganders, ganders of pairs seldom fought ganders of families of four or larger, ganders of families of four seldom fought ganders of families of six and so on.
WYNNE-EDWARDS (I962: I40) noted for many species the lack of correla-
tion of dominance with size and apparent strength. This is true in Canada geese as some older, larger males have no young and are submissive to younger, smaller males with broods. For example, one marked adult male of IO lbs, 3 oz (larger than average; cf. RAVELING, I968b) with greatly extensor of the enlarged portions carpometacarpus, (indicative of an older adult male; HANSON, 1962) was a gander of a family of three. The behavior of this bird was typical for that size group and he was not highly aggressive, nor more often victorious than other ganders in families of three. Another gander in this study weighed 8 lbs, 14 oz (less than average) and had only modestly enlarged and still feathered knobs of the carpometacarpus. He was probably 212 or 312 years old, but he was a gander of a family of five and was as successful as ganders of other families of five in winning encounters. I suggest that the mechanisms which allow recognition and avoidance of fighting with ganders of large families by ganders of smaller families and which allow the gander of the larger family to win a fight when one occurs are associated with the role of increased numbers of young in directly stimulating increased intensity of threat display and fighting ability. The vigor of display of a gander and the intensity and amount of participation of his family are related to situations in which mild threats do not cause avoidance but instead lead to pre-attack behavior by other geese so that a fight becomes imminent. The actions of the ganders involved change from mild threat to ritualized threat serving postures (Erect, Head-pumping, and Rolling - see Part II) which usually suffice to prevent a fight because one family moves away, especially if the gander of the other family starts to attack. FISCHER(I965: 302) concluded that Rolling served to maintain rank order without severe fighting because intense display and actual performance of
306
DENNIS G. RAVELING
the activity connected to a display are often largely mutually exclusive. While this appears true it does not explain why the Rolling gander of a smaller family avoids or loses to the Rolling gander of a larger family. Stereotyped displays of ducklings and castrated ducks can be caused by
testosterone treatment (PHILLIPS & MCKINNEY, I962). Much research has
demonstrated the correlation of aggressive behavior and social rank with

androgen production (e.g., see reviews by GOLLIAS,I95ob and DAVIS, I964). HANSON (1953) suggested that pairs of Canada geese with broods might
have higher "hormone" levels through the winter than unmated geese or pairs without young. Since dominance and aggressive success fluctuate immediately upon separation and reunification of family members it appears that numbers of a brood serve to stimulate in a rather quantum fashion aggressive behavior related to androgen level which is permissive to success in "bluffs" and fights. Apparently geese with high motivation (more young) and, thus, fighting ability, display more intensely and this is recognized by lower ranking geese. The fact that ganders of newly formed pairs were dominant over family ganders supports rather than contradicts the above explanation. The formation of a pair bond, the rather continuous Triumph Ceremony activity of a new pair, and the female's unusual aggressive and inciting behavior (RAVELING, I967: 58) were apparently powerful stimuli causing the male to be in one of the most aggressive states possible and, therefore, able to attack any goose. The normally much more aggressive family ganders would begin displaying but were then dominated by the male of a newly formed pair which usually went immediately into attack. These observations support the suggestion that quantitative differences in the vigor of threat display and attack intensity exist between ganders and that these differences account for most of the submission of lower ranking geese. Since dominance implies benefits, families of geese are served best by aggressive behavior and in increasing magnitude with increased size of family. In long-lived species in which the young are dependent on or at least closely associated with their parents for a relatively long period of time considerable survival values are attached to behaviors that maintain success:ful pairs and their offspring. Immature geese in families are assured food, space, and relative freedom from constant attack or defeat as opposed to immatures without parents and adults without young. With limited food in a captive flock, JENKINS (I944) observed that the dominant fed first. JENKINS(1944) and HANSON(1953) noted how dominance of pairs with young could be important in times of food shortages for insuring the maintenance of successfully producing individuals and their offspring.
BEHAVIOUR
XXXVII,
(I970):
306
PLATE VIII
Figure I. Common attitude of a Canada goose fleeing from attack.
. .......,,j,. .. :
Figure 2. Submissive attitude of a Canada goose.
BEHAVIOUR
XXXVII,
(I970):
306
PLATE IX
Figure 3. Erect postures of Canada geese. Extreme position is illustrated by the goose being attacked. The goose immediatelyto the right of the attacker illustrates more closely the usual attitude before attacking or fleeing (or resuming other activity).
Figure 5. Triumph Ceremonybetween a pair of Canadageese. The male (left) is in the Cackling posture.
BEHAVIOUR
XXXVII,
(I970):
306
PLATE X
Figure 7. Bent-neck posture of a Canada goose.
Figure 8. Forward posture of a Canada goose.
BEHAVIOUR
XXXVII,
(1970):
307
XI PLATE
Figure 9. Attack by a family of 6 Canada geese. The gander (I) is transforming from Rolling movements to attack a gander (A) of another family (remainder of family is out of the photo but were identified in display before the attack). Other members of the attacking family (2, 3, 4, 5, plus a sixth out of the frame to the right) exhibit varying Bent-neck and Forward postures with the head high or low. Note the sleeked neck feathers of attacking geese with extended head and neck in contrast to erected neck feathers of the escaping goose (A). Note also the Erect posture of B in contrast to general unalertness of other geese in the vicinity.
AGONISTIC BEHAVIOR. OF CANADA GEESE IN WINTER
307
PART II. -
AGONISTIC BEHAVIORS
It was established in Part I that benefits of dominance among geese were usually obtained not by fighting but by recognition of actions which serve the functions of threat and submission. Stereotyped postures with threat functions (and other displays serving as social signals) often represent the resultant of the simultaneous activation of two or more incompatible tendencies to behave in relatively mutually exclusive manners, e.g. attack or
flee (cf. TINBERGEN, I952a, I954, I959, I964; HINDE, I953; BASTOCKet al., I954; ANDREW, I956; VAN IERSEL & BOL, 1958).
This "conflict hypothesis" of motivation analysis has been clearly demonstrated in Canada geese. BLURTONJONES (I960) experimentally pro-
duced stimuli eliciting uninhibited attack, uninhibited escape, and both situations simultaneously with captive, tame geese. His results demonstrated that certain postures exhibited by Canada geese in aggressive situations were related to varying amounts of conflicting tendencies to attack and escape at
the same time. BLURTON JONES (1960) used the terms Erect, Head-pumping,
Bent-neck, and Forward to describe four agonistic postures. Other descriptions of some threat and attack behavior of Canada geese are provided by BALHAM (I94: 128-137, I58-I59), COLLIAS & JAHN (I959), and KLOPMAN
(I96I, I968). I choose to follow the terminology of BLURTON JONES (I960) and FISCHER (I965) rather than KLOPMAN(I968).
RESULTS
Descriptions
and Interrelationships.
Fleeing. This is the obvious escape of one goose from another (Figure I). Usually the neck is erect and the head held high with neck feathers erected. Submissive Posture. The neck is curled and the bill points down and away from other geese and almost touches the breast, and neck feathers are somewhat erected (Figure 2). This attitude is especially prevalent among single geese as they move among other geese or are being approached by other geese. This posture is frequently assumed by a goose after it has fled from another individual. A goose in this posture is often pecked or threatened by other geese, but not attacked vigorously. Erect Posture. This position is almost identical to the alert position geese assume when disturbed, except that the body is usually more erect in the agonistic situation
308
DENNIS
G. RAVELING
(Figure 3). The angle of the neck may be vertical or tilted away from or towards other nearby geese. The Erect posture is similar to fleeing and often neck feathers are raised. Sometimes, however, the neck feathers are sleeked and the body feathers erected. The Erect posture is a response to activities of another goose (or groups) in the near vicinity upon which the Erect goose fixates. This posture often gives way to fleeing, especially when neck feathers are raised and the neck tilted away from nearby geese which are exhibiting more elaborate displays. Many times nothing further develops and the goose resumes a normal carriage and continues the activity in which it was engaged previously. The Erect posture may transform to more obvious displays (see below) or attack, especially if the Erect goose has sleeked neck feathers and erected body feathers. Head-pumping. Repeatedly lowering and raising the head causes a bobbing motion (Figure 4). The neck is often vertical but it may be tilted towards another goose while the pumping continues. This display is initiated when approaching other geese or as a response to other geese which are approaching or exhibiting hostile displays. Nothing further may develop, or Head-pumping may transform directly or by transition into Bent-neck, Forward, or Rolling postures or even attack. Head-pumping may also transform to the Erect posture or escape.
Figure 4. Head-pumping postures of a Canada goose. The head is repeatedly lowered and raised (adapted from BLURTON JONES,I960).
Rolling. This is the most complex display of Canada geese. At low intensity the head may be shaken in a manner almost indistinguishable from preflight
Head-tossing
(RAVELING,
I969C). This involves a vertical lifting of the bill
while the neck is erect. At higher intensity the neck and head are waved forward and back and from side to side in an arc-like movement with the bill pointing up and the head being vigorously shaken from side to side. The
309
white cheek patches are thereby displayed prominently. Strident honking is characteristic of the display. Wing-flicks or Wing-shaking (MCKINNEY, 1965) may accompany the highest intensity performances. Rolling may alter with Cackling between Triumph Ceremony partners (Figures 5 and 6) and Bent-neck and Forward postures "aimed" at other geese. Thus the head may swing from rotary Rolling movements to horizontal Cackling to Forward threatening and back to Rolling. Rolling and Cackling are two components of the Triumph Ceremony of geese and terminology follows FISCHER (I965). This behavior is associated with a variety of situations and functions. The
Figure 6. Triumph Ceremony between a pair of Canada geese illustrating Rolling postures with the male on the left. (Adapted from FISCHER, I965: 279).
discussion here refers only to the hostile or threat function. Further description of the Triumph Ceremony of Canada geese is provided elsewhere
(RAVELING, I967: I8-33).
Rolling is initiated when approaching other geese or as a response to approach or nearness of another Erect, Head-pumping, Rolling or chasing goose. It is often a performance between two ganders of different families just prior to attack or fighting. Sometimes nothing further develops but when attacks occur they are of the highest intensity. Rolling may develop from Erect or Head-pumping postures and it may transform to the Erect posture. Bent-neck Posture. The head is pointed at another goose with the bill often angling slightly down, and the neck is coiled back to varying degrees (Figure 7). The Bentneck posture is frequently directed at nearby geese in the Submissive or Erect attitude, or already fleeing. Bent-neck positions often develop from Erect or Head-pumping postures or as the goose lowers its head to drink or eat, but then turns slightly towards another goose. Bent-neck occasionally transforms to Head-pumping, Erect, or Forward postures, or direct attack.
3IO
DENNIS
G. RAVELING
Forward Posture. The head and neck are extended horizontally with the neck more or less straight and directly oriented at another goose (Figure 8). This posture is given in essentially the same situations as the Bent-neck and usually develops from the Bent-neck and may revert to it. Attack. Except for fleeing and often the Submissive position, the above descriptions of agonistic postures refer to geese that are essentially stationary, although hesitant movements toward or away from other geese occur with all the postures. A goose which moves rapidly toward another goose is considered as attacking. Actually it is more common to observe the Forward posture while an individual is pursuing a fleeing goose than when stationary. Attacks occur in a gradation of intensities and intermediates between the described postures are common, especially the degree of neck coiling of the attacker. However, attacks can be considered in three categories: (a) low intensity - walking towards another goose while in a Bent-neck, Forward, or intermediate posture; intermediate intensity - running towards another goose, often in a Bent-neck posture but usually more in a Forward position; high intensity - running at a goose with head high or low, the neck in varying coiled positions, and, often, with the wings spread slightly (Figure 9). As with Bent-neck and Forward postures given while stationary, low and medium intensity attacks are usually directed at geese that are already fleeing or in the Submissive attitude. High intensity attacks are often those situations where two ganders rush at each other, usually subsequent to Rolling. High intensity attacks by the victor may continue after the defeated goose has turned and is fleeing. These types of attack and fighting almost always involve ganders of families. After more vigorous attacks and always after a fight, the victorious gander turns and enters a Triumph Ceremony with one or more members of his family. DISCUSSION and function of motivation, evolution, Origin, behaviors. agonistic Since displays serving threat functions are often considered resultants of the conflict to attack and escape at the same time, comparison of threat postures to actual attack and fleeing is a method of analysis for discovering the balance or predominance of underlying motivations and origin of a
AGONISTIC
BEHAVIOR
OF CANADA
GEESE IN WINTER
311
behavior (cf. TINBERGEN, I952a, I959, I964).
Some hostile actions may be
obscure if they have been ritualized from the original movement or are displacement activities which appear irrelevant but have been formalized in a new context having signal value. Submissive Attitude. Comparison of this "Appeasement" posture to other positions shows that it is antithetical to attack. The tendency to escape is involved as indicated by the raised neck feathers and avoiding behavior of the goose. The Submissive posture is characteristic of single geese and their predisposition to aggressive behavior is extremely low. The Submissive attitude is, however, different from the usual fleeing behavior and since attack tendencies do not seem to be involved, the probable motivations resulting in this behavior are complex. Regardless of how avoiding a goose is, the power of attraction to other geese and entrance into the activities of a flock prevent single geese from
isolating themselves. I suggest that the Submissive attitude results mainly
from the conflicting tendency to approach (but not attack) and be with other geese and flee from them at the same time. Support for this suggestion is derived from three sources: (a) The behavior usually results from the tendency of a single to join and stay with conspecifics even though it may be rather constantly threatened or chased. (b) This attitude is similar to the manner in which approach and pair formation is initiated in other goose
species (cf. FI;SCHER, I965: 275) and probably in Canada geese (RAVELING,
1967: 57). (C) The Submissive posture and "listless" avoiding attitude while at the same time approaching many geese is the most characteristic behavior of individuals separated from and "searching" for their Triumph Ceremony partner(s).
FISCHER (I965: 257) described the unceasing searching behavior of geese separated from their mates and families and LORENZ (I959: 216, 1966: 207)
gave particularly graphic descriptions of the state of these animals. Such behavior was clearly demonstrated on the occasion that an immature male of a marked family of six was split from the rest of his family. He was observed six times for approximately 7 hours while separated and a great deal of this time was spent in continually walking among large numbers of loafing and sleeping geese. He was often threatened, but not vigorously chased. He was almost continually in the Submissive posture and for 8 days he did not fly out to feed. Upon reunification with his family, he abruptly became again one of the most active and aggressive immatures observed during this study. The Submissive posture seems to have originated from the intention
312
DENNIS G. RAVELING
movement of withdrawing the head from contact with other geese. The movement has apparently been ritualized and serves the functions of identifying single geese, allowing approach, habituation, and ultimately pair formation. This behavior also serves to prevent violent attacks against these individuals, probably because the attitude contains little or no aggressive components. The continuing approach of these lowest ranking individuals close to other geese, however, causes them to be threatened a great many times since any paired or family geese are dominant over them. Erect Posture. This position with neck feathers raised often gives way to fleeing and the posture is similar to escape and can be considered as the intention movement of escape, but in conflict with a lesser tendency to remain and/or attack. BLURTON JONES (1960) found that the Erect posture could be elicited only when a tendency to attack coexisted with a strong tendency to flee, but not when attack was prevented when no escape causing stimuli were present. When sleeked neck feathers and raised body feathers are characteristic of the Erect posture they represent intention elements of attack. BALHAM
(1954: I37) and COLLIAS & JAHN (I959) also noted that erected neck
feathers were related to alarm and escape while oppressed neck feathers and erected body feathers were associated with aggressive behaviors. When attack tendencies predominate this posture often transforms into other actions representing higher levels of attack or conflict motivations (Head-pumping, Rolling) or into attack itself. I interpret the Erect posture as one which represents a balance between attack and escape tendencies. Either motivation can predominate depending upon the behavior of other nearby geese. The position has originated from intention movements of fighting and escaping but since it often is not followed by either such activity it represents a relatively "weak" conflict
(cf. TINBERGEN, I952b).
The Erect posture is exaggerated in the sense that the angle of the body can be extreme and the posture can be held for relatively long periods of time. The behavior appears to have signal fuction as other geese avoid, or react with similar postures, an individual in this position. Thus, this posture serves threat, rank order establishment and maintenance functions. Head-Pumping. This display appears to represent an almost perfect balance between a simultaneous tendency to escape and attack. When the head is lowered the posture is almost identical to the Bent-neck and incipient attack. I consider
313
this lowering and "aiming" of the head to be an intention movement of attack. When the neck is then straightened and the head held high the posture is similar to escape and is interpreted as an intention movement of escape. The behavior of a Head-pumping goose is ambivalent and either attacking and fleeing can occur. Whether or not a goose will transform to more intense display, attack, or flee can usually be predicted from the rapidity and extent of the Head-pumping in relation to the angle of the neck. If pumping is intense the chances for attacking or fleeing increase and the neck slants toward or away from another goose and Head-pumping transforms into more prevailing attack or escape tendencies, respectively. Although I interpret Head-pumping as originating from intention movements of attack and escape, I suggest that this posture is ritualized and that pumping itself is communicative of attack rather than just the lowered head component. BLURTON JONES (I960) observed Head-pumping only in situations containing simultaneous attack and flee stimuli. Other geese respond with avoidance or display and thus Head-pumping also serves threat and rank order functions. Rolling. Rolling contains motor patterns and vocalizations indicating that this behavior is the result of a variety of conflicting motivations (FISCHER, I965). Especially important appears to be the spatial relationship of a gander to his mate and family. Intrusion of another high ranking gander or family on these boundaries may result in violent attack. Rolling often serves as the highest intensity threat of geese and is responded to by other geese with avoidance or comparable Rolling. The motivation and the origin of all the components of Rolling are not clearly understood. However, it does seem clear that the exaggerated head and neck movements are derived from preflight Head-tossing which represents a displacement resulting from the conflict of leaving and remaining with Triumph Ceremony partner(s) (RAVELING, I969c). The behavior is highly
conspicuous.
In Part I, I suggested that geese recognize superiority on the basis of type and intensity of threat. The occurrence of Head-pumping and Rolling prior to attack or escape by the marked geese demonstratedthe much greater amount of posturing engaged in by ganders of families as compared to pairs and single geese (Table 9). There was a non-significant difference in amount of Head-pumping by ganders of different size families but Rolling was observed more often in families of 5 than in families of 4 (x2 = 2.81, p < o.IO). These limited data provide some confirmation of the hypothesis that threat intensity is related to family size which thereby accomplishes recognition of rank Behaviour XXXVII 20
314
DENNIS G. RAVELING
position. In addition to recording category of threat, quantification of relative intensity within each display by different social status individuals and families is needed. Studies of vocalizations are also needed. TABLE 9 Variation among different social status radio- and color-marked Canada geese in the prevalence of two different intensity displays serving threat functions Number of Number Numberof conflictspreceded of conflictspreceded by Rolling, at least conflicts by Head-pumping by the gander
I35 39 57 92 3 2 i6 23
(2.2%)
Group
Single geese Pairs Families of 3 Families of 4
Totals
3 (2.2%) (5.1%) 17 (30%) 29 (32%)
(5.1%) (28%) (25%)
-2 I (I.8%) 6 (6.5%)
Families of 5
I8i
36 (20%)
24 (13%)
60 (33%)
Bent-neck and Forward Postures. These positions represent intention movements of attack and the Forward usually results from the Bent-neck. When another goose is not already fleeing or does not begin to escape in response to Bent-neck or Forward postures, the behavior of the threatening goose often changes to Erect and/or Headpumping or into full attack. LORENZ (1959: 206, 1966: IOI) interpreted the
amount of neck coiling as indicating the balance of attack and escape and
thus the Bent-neck was interpreted as containing more fear than the Forward posture. BLURTONJONES (I960) demonstrated that these postures do indeed
commonly occur in situations where attack and fleeing stimuli were simultaneously present and that the Forward posture indicated a more strongly attack motivated individual. He also demonstrated that Bent-neck and Forward postures were displayed in situations where attack was thwarted by other than an escape situation.
While fear or escape tendencies may be conflicting in some occurrences
of Bent-neck and Forward displays, it appears that these behaviors often result from the conflict of the tendency to attack and continue their present activity at the same time. Escape tendencies need not be involved. The Bentneck is often observed to alternate with feeding, preening, loafing, or travelling about. This conclusion is further supported by the fact that when Bent-neck or Forward threats are directed at a goose which does not flee but may begin displaying itself, the behavior changes to Head-pumping and other postures described above that more clearly represent a conflict of attack and escape.
AGONISTIC BEIIAVIOR OF CANADA GEESE IN WINTER
315
Therefore, I conclude that Bent-neck and Forward postures, except possibly as associated with high intensity attacks, are not displays that are important in serving recognition and establishment of the dominance hierarchy in Canada geese. These movements, however, would serve to maintain and reinforce spatial separation among geese that already recognize each other. Dominance over singles and possibly pairs may be achieved by these postures because singles usually avoid any type of threatening.
SUMMARY The dominance relationships and associated agonistic postures of Canada geese were studied during the winters I963-64 and I964-65 by observationof the behavior of radioand color-marked families and individuals living in a large, wild flock in southern Illinois. Part I I. Behavior was not significantly altered by color-marking and attachment of transmitters. 2. Hostile encounters occurred during almost all activities. They varied widely in frequency and intensity and were especially associated with feeding. 3. Success in hostile encounters was directly related to family size, i.e., large family> smaller family>pair> single. 4. Unified action by all members of a family occurred in 8.5 percent of victories and 15 percent of defeats. 5. All members of a family shared equal dominancestatus but the success of a family in the rank order was most dependentupon the gander. 6. Only once in 26 fights between unmarked family ganders did the gander of the largest family lose. 7. Dominance position of family individuals decreased immediately upon separation, and increasedupon reunificationof family members. 8. Large families were engaged in significantly more conflicts per unit time than were singles, pairs, and small families. 9. Exceptions to the usual dominance hierarchy occurred after pairs were newly formed. The gander of a newly formed pair could dominatefamily ganders. o1. Intrafamily aggression was rare and of low intensity. T . Fights rarely occurred; threats and chases were common. I2. In some instances, rank orders based upon individual recognition could exist. However, stable rank orders in most large flocks appear to be based on recognition of different postures and levels of intensity of threat. 13. 'The dominanceorder of geese yields benefit in terms of food and space acquisition and freedom from defeat in aggressive encounters for the pairs and their young in direct relation to those most successful at raising a brood. Part TI 1. Postures associated with attack or fleeing or simultaneoustendenciesto do both are described. These include actual fleeing or attack, Submissive attitude, Erect, Headpumping,Rolling, Bent-neck, and Forward postures. 2. The Submissiveattitude is exhibited mostly by single geese and probablyresults from the conflicting tendency to approach (but not attack) and flee from other geese at the same time. This posture functions to identify single geese, allow approach,habituation, and ultimately pair formation, and inhibits violent attack.
316
DENNIS G. RAVELING
3. The Erect posture may take either the form of intention movements of escape or attack and represents an ambivalentmotivation between these two tendencies. 4. Head-pumpingcontains alternating intention movementsof attacking and fleeing and represents almost a perfect balance between these two tendencies but is of higher intensity and ritualizationthan the Erect position. 5. Rolling is a complex portion of the Triumph Ceremonybut also serves as the most intense threat of Canadageese and is highly ritualized.The spatial relationshipsof a gander to his mate and family appearmost importantin motivating Rolling. Intrusion of another high ranking gander or family on those boundariesmay result in violent attack. 6. Erect, Head-pumping,and Rolling serve as three different intensity threats which are recognized by other geese and serve to maintain and establish the rank order of geese without undue fighting. 7. Bent-neck and Forward postures may occasionally represent conflicting attack and flee tendencies but often appear to represent a conflict of attack and remain doing another activity such as feeding or preening. These postures serve to maintain and reinforce a rank order but are probably not very important in initial establishment of rank. REFERENCES R. remarks on behaviour in conflict situations, with special ANDREW, J. (I956). Some reference to Emberiza spp. - Brit J. Anim. Behav. 4, p. 41-45. BALHAM, R. W. (I954). The behavior of the Canada goose (Branta canadensis) in Manitoba. - Ph.D. dissertation. Univ. Missouri, Columbia.229 pp. BARTONEK, J. C. & DANE, C. W. (I964). Numbered nasal discs for waterfowl. J. Wildl. Mgmt. 28, p. 688-692.
BASTOCK, M. D., MORRIS, D. & MOYNIHAN, M. (I954). Some comments on conflict and BLURTON JONES, N.
Canada geese. - Wildfowl Trust Ann. Rept. II, p. 46-52. BOYD, H. (1952). Notes on colour marking of geese. - Wildfowl Trust Ann. Rept. 4, p. 14-16. - (I953). On encounters between wild white-fronted geese in winter flocks. CoLLIAS,
thwarting in animals. - Behaviour 6, p. 66-84. G. (196o). Experiments on the causation of the threat postures of
birds and mammals. -
Behaviour 5, p. 85-129. N. E. (I95oa). Some variations in grouping and dominance patterns among
Zoologica 35, p. 97-II9.
(I95ob). Hormones and behaviorwith special reference to birds and the mechanisms of hormone action. - In: A Symposiumon Steroid Hormones. E. S. GORDON (ed.).
University of Wisconsin Press. Madison, p. 277-329.
--
DELACOUR, J. & MAYR, E. (I945). The family Anatidae. - Wilson Bull. 57, p. 3-55. FISCHER, HELGA (I965). Das Triumphgeschrei der Graugans (Anser anser). Z. Tierpsychol. 22, p. 247-304. HANSON, H. C. (I953). Inter-family dominance in Canada geese. - Auk 70, p. II-I6.
L. R. (I959). Social behavior and breedingsuccess in Canadageese (Branta & JAHN, canddensis) confined under seminaturalconditions.- Auk 76, p. 478-509. DAVIs, D. E. (1964). The physiological analysis of aggressive behavior. - In: Social behavior and organization among vertebrates. W. ETKIN(ed.). Univ. of Chicago Press. Chicago, p. 53-74. f.
Nat. Hist. Surv. Biol. Notes 49. 15 pp. R. H. (I950). Canada geese of the Mississippi Flyway: with special - & SMITH, reference to an Illinois flock. - Illinois Nat. Hist. Surv. Bull. 25, p. 67-210.
(I962). Characters of age, sex, and sexual maturity in Canada geese. -
Illinois
AGONISTIC
BEHAVIOR
OF CANADA
GEESE IN WINTER
317
Anatiden. - Verhand. Tnter.Ornithol. Kongr. 5, p. 589-702. R. A. (I953). The conflict between drives in the courtship and copulationof the HINDE, Chaffinch. - Behaviour 5, p. I-3I. IERSEL, J. J. A. VAN & BOL, A. C. A. (1958). Preening of two tern species. A study of displacement activities. - Behaviour 13, p. 1-88.
JENKINS, D. W. (I944).
0. (I9II). Beitrage zur Biologie, namentlich Ethologie und Psychologie der HEINROTH,
for determining migration routes. - Calif. Fish and Game 45, p. 69-82. P. & JOHNSON, L. L. (I958). Marking ducks for research. - Flicker 30, p. 98-99. K. (I935). Der Kumpanin der Umwelt des Vogels. - J. f. Orithol. 83, p. 137LORENZ,
LINDMEIER, J.
Territory as a result of despotism and social organization in geese. - Auk 6I, 30-47. R. B. (I96I). The greeting ceremonyof Canadageese. - Mag. of Ducks and KLOPMAN, Geese 12, p. 6-9. - (1968). The agonistic behavior of the Canada goose. - Behaviour 30, p. 287-319. F. M., MILLER, A. W. & RIENECKER, W. C. (I959). Color-markingwhite geese KOZLIK,
214; p. 289-413. (I959). The role of aggression in group formation. -
In: Group processes: Trans-
McKINNEY, F. (I965). The comfort movements of Anatidae. - Behaviour 25, p. 120-220. PHILLIPS, R. E. & McKINNEY, F. (1962). The role of testosteronein the displays of some
actions of the fourth conference. B. SCHAFFNER (ed.). Josiah Macy. Jr. Foundation. N.Y., p. 181-252. - (1966). On aggression. - Harcourt, Brace & World, Inc. N.Y. 306 pp.
ducks. Anim. Behav. 10, p. 244-246.
RAVELING,
D. G. (I967). Sociobiology and ecology of Canada geese in winter. Ph.D. dissertation. Southern Illinois Univ. Carbondale.213 pp. (I968a). Can counts of group sizes of Canadageese reveal populationstructure?In: Canadagoose management.(R. L. HINE & C. SCHOENFELD, eds.). Dembar Educ. Res. Serv. Inc., Madison, Wisc. p. 87-91. (I968b). Weights of Branta canadensisinterior during winter. - J. Wildl. Mgmt.
32, p. 412-414.
(I969a). Social classes of Canadageese in winter. - J. Wildl. Mgmt. 33, p. 304-318. (I969b). Roost sites and flight patterns of Canada geese in winter. - J. Wildl.
(I969c). Preflight and flight behaviorof Canadageese. - Auk 86. p. 671-681. SCOTT, J. P. (I958). Aggression. - Univ. of Chicago Press. Chicago. 149 pp. SIEGEL, S. (I956). Nonparametricstatistics for the behavioral sciences. - McGraw-Hill
TINBERGEN,
Mgmt. 33, P. 319-330.
Book Co., Inc. N.Y. 312 pp. N. (I952a). "Derived" activities; their causation, biological significance, origin and emancipation during evolution. - Quart. Rev. Biol. 27, p. 1-32. - (I952b). A note on the orgin and evolution of threat display. - Ibis 94, p. 160-162. (1954). The origin and evolution of courtship and threat display. - In: Evolution as a process. A. C. HARDY,J. S. HUXLEY, and E. B. FORD (eds.). Allen and Unwin, Ltd. London. pp. 233-250. Behaviour 15, p. 1-70.
- (959). Comparativestudies of the behaviourof gulls (Laridae): A progress report.
(I964). The evolution of signaling devices. - In: Social behavior and organization among vertebrates. W. ETKIN (ed.). Univ. Chicago Press. Chicago. pp. 206-230. WYNNE-EDWARDS, V. C. (I962). Animal dispersion in relation to social behavior. Hafner Co. N.Y. 653 pp.
3I8
DENNIS
G. RAVELING
RfSUMP La predominance des parentes et I'associationdes po,stureshostiles des Oies du Canada, ont ete'etudiees durant les hivers de I963 a I964 et de I964 a 1965, par l'observationdu comportement des families, marquees par la ,,radio" ou par la couleur et d',individus appartenanta un grand troupeau sauvage du sud de l'Illinois. 1ere Partie T. Le comportementn'etait pas significativement altere en marquantles sujets par la couleur et en leur attachant des radio-emetteurs. 2. Des rencontres hostiles se sont produitespresque toujours durant toutes les activites, variant largement dans Ia frequence et l'intensite. Elles etaient surtout associees a l'alimentation. 3. Le succes des rencontres hostiles etait directement lie a la taille de la famille, par exemple: une grande famille> une plus petite famille> un couple> ou unique. 4. L'action unifee de tous les membresde la famille s'opere dans 8.5% des victoires et
de 15% des echecs.
5. Tous les membres de la famille partagent egalement la position autoritaire; mais le succes d'une fanille quant a l'ordre de son rang depend surtout du jars. 6. Dans 26 combats entre families de jars non marques, seulement une fois les jars de la plus grande famille ont echoue. 7. La position predominantedes individus diminue immediatementapres la separation d'avec la famille et augmente par la reunification des membres avec la famille. 8. Les grandes familles etaient significativement engagees dans de plus grand conflits que les individus isoles, les couples et les petites familles. 9. Exceptions a la predominancehierarchique habituelle, se sont produites apres la formation de nouveaux couples. Le jars d'une famille recemmnent formee, peut dominer une famille de jars. To. L'agression intrafanille etait rare, et sont intensite tres faible. IT. Les combats se sont rarement produits, mais les menaces et les poursuites etaient frequentes. 12. Dans quelques cas, l'ordre des rangs, fonde sturla reconnaissancedes individus peut exister, bien que l'ordre le plus stable dans les grandes bandes soit fonde sur la reconnaissancede differentes postures sur les degres de l'intensite de la menace. 13. L'ordre de dominancedes Oies donne un benefice en nourriture, en espace, et en liberte a la suite des rencontres agressives des couples et de leurs petits et se tient en relation directe avec l'aptitudea elever une nichee. 2eme Partie i. Les postures associees a l'attaque, ou a 1'envol ou aux deux sont decrites. Celles-ci comprennentl'envol ou I'attaque.Attitude soumrise, erigee, gonflant la tete, roulant, cou penche et tete portee en avant. 2. L'attitude somllise est surtout prise par les Oies isoles et resulte souvent d'une tendance a s'approcherdes autres Oies, non pour attaquer mais pour les eloigner. Cette position a pour but d'identifier les Oises isolees, permettre l'approche,la familiarisationet finalement la formation du couple et empecherune attaque violente. 3. L'attitude erigee prend soit la forme des mouvements avec l'intention de s'echapper ou d'attaquer,et represente une motivation ambivalente entre ces deux tendances. 4. Le comportementdit ,,pompage de la tete" consiste alternativementen des mouvements avec l'intention d'attaquer ou de s'enfuir et repre'sentepresque un parfait equilibre entre ces deux tendances; mais il est d'une intensite et d'une ritualisation plus haute que ,,la posture herissee".
319
5. L'attitudedite de ,,roulement"est un segment complexe de la ceremoniedu triomphe; elle figure la menace la plus intense qu'exprimentles Oiescdu Canada.Les relations spatiales d'un jars avec sa femelle et sa famille semblent tres importantes dans la motivation de l'attitude ,,roulement".L'intrusion d'un jars d'un rang social eleve ou d'une famille sur ces frontieres peut provoquerune attaque violente. 6. L'attitude ,,tete gonflee" et ,,roulement herissee' servent de menaces differentes qui sont reconnuespour telles par les autres Oies, etablissent et maintiennentl'ordre hierarchiquedes Oies sans bataille excessive. 7. ,,Cou penche" et ,,tete en avant" representent eventuellement une attaque ou une tendance a s'enfuir, mais souvent semblent representer une attitude d'attaque, se combinant a d'autres activites telles que la prise de nourriture ou le lissage des plumes. Ces poses aident a maintenir et a renforcer l'ordre hierardhique, maisl elles sont moins importantesque d'autres dans l'etablissementinitial du rang.

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Dominance Relationships and Agonistic Behavior of Canada Geese in Winter Author(s): Dennis G.

RELATIONSHIPS AND AGONISTIC BEHAVIOR OF CANADA GEESE IN WINTER by

Wildlife ResearchLaboratory, SouthernIllinois University,Carbondale, (Cooperative Illinois,U.S.A.)

Estimated numbers of geese observed

Numbers of threats and chases

(a.m. feeding period)

(IIo & I50)

(44 & 55)

Total number of conflicts 20

Number of victories o (o%) 5 (17%)

Single adult female

Families of 3 Families of 4 Families of 5

69 (75%) I49 (82%)

* Percent not calculatedwhen total conflicts <20.

ratios of success in encounters of different families within each category

Individual of group Adult male Adult female Adult male

Number of victorious conflicts initiated 6 (6o%) 4 (40%) I9 (76%)

Number of victories in which all members participated 3 (30o%)

One of the others Adult male

5 (8%) I8 (29%) 112 (75%)

Adult male One of the others

Number of losses 2 (I8%) 5 (45%) 4 (37%) 5 (33%) 8 (53%)

One or more of the others

Numberof victories 9 (6o%)

Families of 4 59 Families of 5 I39

50 (85%) 124 (89%)

as determined from observations families. of unmarked ganders

Lengthof time of observations

Single yearling males

AGONISTIC BEHAVIOR OF CANADA GEESE IN WINTER

AGONISTIC BEHAVIOR OF CANADA GEESE IN WINTER

method of capturing the animals. Cannot-net traps (DILL & THORNESBERRY,

AGONISTIC BEHAVIOR OF CANADA GEESE IN WINTER

demonstrated the correlation of aggressive behavior and social rank with

Figure I. Common attitude of a Canada goose fleeing from attack.

Figure 2. Submissive attitude of a Canada goose.

Figure 7. Bent-neck posture of a Canada goose.

Figure 8. Forward posture of a Canada goose.

AGONISTIC BEHAVIOR. OF CANADA GEESE IN WINTER

I969C). This involves a vertical lifting of the bill

AGONISTIC BEHAVIOR OF CANADA GEESE IN WINTER

behavior (cf. TINBERGEN, I952a, I959, I964).

Some hostile actions may be

AGONISTIC BEHAVIOR OF CANADA GEESE IN WINTER

(5.1%) (28%) (25%)

AGONISTIC BEIIAVIOR OF CANADA GEESE IN WINTER

birds and mammals. -

(I962). Characters of age, sex, and sexual maturity in Canada geese. -

214; p. 289-413. (I959). The role of aggression in group formation. -

In: Group processes: Trans-

Mgmt. 33, P. 319-330.

- (959). Comparativestudies of the behaviourof gulls (Laridae): A progress report.

AGONISTIC BEHAVIOR OF CANADA GEESE IN WINTER

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