In a more extensive study using 5 male and 5 female beagles, BEACH & .... Stand: Maintains erect posture for at least 5 seconds when mounted or in- vestigated.
INTERINDIVIDUAL
ASSOCIATIONS
IN DOGS
by BURNEY
J. LE BOEUF 1)
(Department of Psychology, University of California, Berkeley, Calif., U. S. A.) (With 7
Figures)
(Rec. 3I-1-i967)
CONTENTS I. Introduction .269 II. Subjects, Materials and Methods.270 III. Results ..275 A. Heterosexual Interactions.275 i. Normative data from the anestrous, estrous, and post-estrous periods
2. 3. 4. 5. 6. 7.
.. of females .275 The response of individual females to individual males . . . Behavior in the circle when females were in estrus . A comparisonof present and previous mating preferences Behavior in the circle when females were in nonestrus .. .282 Temporal changes during estrus . . . .285 Individual differences among males
8. Differences
in the testing condition, tethered vs roving
B. Intrasexual Interactions i. Female-femalebehavior 2. Male-male behavior C. Normative Comparisons .. IV. Discussion .. Summary .. References
.
.
277 277 28I 282
286
..286 ..286 ..288 ..290
.29I .292
.293
Resume ...294
i) Present address: Crown College, University of California, Santa Cruz, California, U.S.A. This research was conducted at the Field Station for Research on Animal Behavior in Berkeley, California and was supported by United States Public Health Service Grant MH-o4noo to Dr FRANKA. BEACH.The work reportedis a condensation of a Ph.D. thesis submittedto the University of California at Berkeley. I wish to thank Drs P. R. MARLERand G. E. FRENCH for reading and commenting on the original manu-
script and particularly,Dr FRANKA. BEACH,who overseeredthe work from its inception to the writing of the final draft.
INTERINDIVIDUAL
ASSOCIATIONS IN DOGS
269
I. INTRODUCTION Numerousallusionsto individualpreferences,friendships,personalbonds, sympathiesand antipathies,etc. can be foundin the literatureon the domestic dog. These citationsemanatefrom a broadsourcewhichincludesdog owners, breeders, veterinarians HARROP,
(WHITNEY,
I947; HANCOCK & ROWLANDS, I949;
I96o), and well-knowninvestigatorsof behavior(ENDERS, I945;
TINBERGEN, I951; LORENZ, I952; KINSEY
et al,
I923;
BEACH, I96I).
The
most frequentreportdeals with strong attractionsbetweenparticularmembers of the oppositesex when the femaleis in estrus. Some authoritiesclaim the male shows preferencesfor certain females while others report that it is the femalethat selects and refuses to mate with certainmales. Whichever the case may be, one thing is certain: all of these referencesare based on subjective impressionsor observations,which are unsubstantiatedby experimentaldata. Two studiespresentquantitativedata.FULLER & DuBuIs (I962) reported sexual behaviorscores for a single fox terrier female observed separately with two males of the same breed during portions of her first estrus. Attemptsto mate the bitch with a litter mate were unsuccessful,but when a strangemale experiencedas a stud was introducedcopulationwas effected within two minutes. In a more extensive study using 5 male and 5 female beagles,BEACH& LE BOEUF (in press) observed differences in each female's readiness to matewith different masculinepartnersover two estrousperiods.Each female matedreadilywith somemalesbut repeatedlyrejectedthe copulatoryattempts of others. Females attemptingto prevent mounting barked, growled, and snappedat males or attackedand bit them.The same males were not always rejected by all females; a male preferred by one female was frequently rejectedby another.A female'spreferenceswere consistentfrom her first estrous period througha subsequentone six monthsto a year later. The foregoing study was not designed specifically to investigate the phenomenaof selectivematingand it is probablethat certainfeaturesin the testing procedureobscuredthe full magnitudeof female preferences. In some tests the mannerin which the animalswere releasedmade it possible for a genital lock to occur duringthe first few secondsafter the start of a test. This had two results. The femalewas tied before she had time to look the male over, so to speak, and once the animalswere locked, no further scoringof preferentialbehaviorwas possible.When copulationdid not occur during the first few seconds of a test, the female exerted obvious control over the matingsituation,copulatingreadilywith somemaleswhile repeatedly
BURNEY J. LE BOEUF
270
rejecting others, usually for the entire I5-minute observation period. Furthermore, each female was permitted to lock just once a day each day of her estrous period. After a lock, no further tests were conducted on that day with other males. Consequently when the female locked with the first male tested one could not say what might have resulted if a different male had been used. As a result of this procedure, all males were not tested on the same days with the estrous bitch and some males copulated more frequently than others. Lastly, rejection behavior by the female was the only measure of individual preferences employed. The primary purpose of the present experiments was to extend the study of BEACH& LE BOEUFand, specifically, to investigate interindividual asisociations during, before, and after mating in a test situation which maximized the frequency of opportunities for interaction. The procedure was designed to facilitate the expression and description of individual preferences by: (a) permitting all males to have equal time with estrous females and vice versa, (b) alleviating the difficulties presented by occurrence of the genital lock, (c) permitting one animal in a test pair to control the frequency and nature of interaction and, in effect, choose among several partners, and (d) measuring preferences, in several different ways. In addition, the nature and frequency of associations of males with males and of females with females was investigated. SUBJECTS, MATERIALS AND METHODS S u b j e c t s'. The subjects (Ss) were 9 pure bred, 3-year-old Beagles of which 5 were males and 4 were females. These same animals were used in the study by BEACH& LE BOEUF (in press). They lived together in a 3/4 acre enclosed field (see Figure i) from the time they were weaned to the beginning of the study, with only occasional separation of the sexes when females were in estrus. During this experiment when the females were not being tested, they were caged and the males ran free in the field. The Ss were fed after the completion of a day's tests. S etti ng. Tests were conducted in a fenced enclosure measuring approximately I,700 sq. ft which formed part of the unit where the Ss lived. Observations were made from a small cabin on the premises which was designed specifically for the purpose. The cabin contained recording instruments and afforded the observer an excellent view of the testing area (see Figures i and 2).
~~~~~~~~~~~
-
Figure
2.
~~~~~~~~~~~~~~~~~
The testing area. A tethered animal is restricted to the area surrounded b small circle (the large circle is not relevant to the present study).
.. ... .......
_....:::::: .: .....................................................................:...:::.
Figure 3. A female wearing a chastity belt.
INTERINDIVIDUAL
ASSOCIATIONS IN DOGS
27I
CABIN STAKE
AND
ENCLOSED
CIRCLE AREA
TESTING
N
/
Figure
i.
CAGES
_
A schematic of the living unit and the enclosure where tests were conducted. The field area measures approximately 34 acre.
P r o c e d u r e.
The behavior of the following pair combinations was observed in the same test situation: (a) heterosexual pairs before, during, and after the female was in estrus, (b) female pairs before, during, and after one of them was in estrus, and (c) male-male pairs. The test situation provided a setup in which one animal could completely avoid contact with the other or terminate contact at any time. One S was tethered to a stake by a 5 ft chain attached to a fitted harness, and thus resitricted to a circular area with a diameter of approximately I2 ft (see Figure 2). The perimeter of thiscircular area was outlined with gypsum to facilitate observations and scoring behavior. The other S, the roving animal, was released into the testing enclosure from the observer's cabin situated 20 ft from the tethering stake. The roving animal was allowed 5 minutes in the field, the duration of each test. It is reported that the type of interactions observed between an estrous female and a male dog do not change substantially if the test duration is extended beyond 5 minutes to 30 minutes (BEACH & LE BOEUF, in press). Daily tests with each pair of Ss included two observations, one while the animal was tethered and one while it was roving free. Behavioral
items
recorded.
The following gross measures were recorded: (a) whether or not the roving animal went to the tethered animal and made physical contact, (b) the time spent by the roving animal in the circle with the tethered animal, and
272
BURNEY J. LE BOEUF
(c) the numberof entrancesand departuresinto and out of the circleby the rovinganimal.Behavioralinteractionsbetweenthe rovingand the tetheredS in the circle were described with categories similar to those used by BEACH & LE BOEUF(in press). In this schema, shown in Table i, an attempt was
made to reduce behavioralinteractionsto discrete stimulus-responseunits or episodes. The term "episode" was used to refer to the initiating act of one animal plus the response of the other. TABLE I Categories of social and sexual behavior in dogs Stimulus Units of Behavior* Code Description of Behavior I Approach: moves to within 2 feet 2 Partial investigation: brings nose within i foot of anogenital region or makes momentary contact with that area 3 Investigation: sniffs and/or licks anogenital area for at least 5 seconds 4A Pre-mount contact: partial mount, forepaws on back, chin over back 4B Prance: forepaws extended, body thrown back on haunches, head cocked to one side; barking and vigorous tail wagging may be involved Present: hindquartersdirected to the other animal and tail is deviated laterally 4C
5
Mount:forelimbclaspwithoutpelvicthrusts
6 7
Clasp and thrust: Same as 5 with several pelvic thrusts Behazior of intromission: same as 6 but thrusts are more rapid and include pelvic oscillation, "steppingand treading" of the hindlegs
Response Units of Behavior** Code Description of Behavior Ai Stand: Maintains erect posture for at least 5 seconds when mounted or in-
vestigated A2
B C
Stand with Tail Deviation: same as Ai plus lateraldeviationof tail Face: whirls aroundto face other animalwhen investigatedor mounted Sit: the nonerect female does not rise when mounted or investigated
D
Avoid: Movesor pullsaway,sits or lies downto avoidmountor investigation
E
Threat: barks, growls, snaps, lunges or any combinationwhich prevents the initiation or continuationof behavior by the other animal Attack: chase, drive, bite or any combination
F
The
estrous
period.
A female's estrous period was defined and determined by a combination of behavioral signs, genital swelling and/or bleeding, and vaginal smears * Malesdisplayall behaviorsexcept4C. Femalesdisplayall behaviorsdescribed.Any activitynot directedto the rear of the other animalbut ratherto the head or side is suffixed by an "H"or an "S". ** All responseunits are shownby the female.The male respondsto a femaleby standing(AI) or by movingaway (D). FemalesterminateAi and A2 responseswith B, D, E, or F, or they standuntil dismounted.
INTERINDIVIDUAL
ASSOCIATIONS IN DOGS
273
taken before each day's tests. A femalewas classifiedas being in behavioral estrus on those days in which the following conditions were satisfied: (a) swelling with or without vaginal discharge,(b) a vaginal smear of the late proestrousor estrous type, i.e., primarilycornified epithelium,with or without red blood cells, and with but few nucleatedepithelialcells, (c) the female "stood",deviatedher tail laterallyand permitteda male to mount, clasp her hindquartersand execute severalpelvic thrusts.With the first two conditionssatisfied,one mountwith clasp and thrustsby one male, in which the female stood and deviatedher tail, was sufficient to classify the female as being in estrus on that day. The behavioralcriterion of standing and "curlingthe tail", alone, has frequentlybeen employedas an indicationof estrus in the bitch
(WHITNEY,
I947; HANCOCK & ROWLANDS, I949; GRA-
HAM, I96I; ASDELL, I966).
One female, Blanche,was tested while in naturalestrus. The other three femaleswere broughtinto estrus by means of 2 intramuscularinjectionsof estrogen1) (.38 mglkg of estradiol benzoate, Schering "Progynon",3.3 mg/ccin sesameoil), spaced48 hours apart.The hormonewas administered during the anestrousperiod, 4-6 months after a female's previous natural estrus. It was known from previouswork that the bitch usually comes into estrus I-5 days after the second injection and remains,in estrus for approximatelyIOdays. Special equipment. In heterosexualtests it was importantto maintainthe female stimulus to each male in as constanta conditionas possible.To do this, and in order to run several combinationsof tests per day, the occurrenceof complete copulationwith insertionand genital lock was prevented.This was effected by fitting eachwith a "chastity"belt whichcoveredthe entireanogenitalarea and preventedmales from achievinginsertion (see Figure 3). The belt was made of thin vinyl plastic, and small holes in the transparentmaterial permittedthe femaleto urinateand allowedfor the transmissionof olfactory and gustatorycues to the male. After a brief period of adaptation,females becameaccustomedto wearingthe belt. It did not impedethe movementsof the femalenor did it produceany apparentdecreasein her interestin the male nor the male'sinterestin the female. It was also necessaryto adaptbothmalesand femalesto the harnessesand the tether. This was accomplishedduring the same 6-day pre-experimental i) The hormoneswere suppliedby Dr PRESTONL. PERLMAN of the Biological Research Department, Schering Corporation,Bloomfield, New Jersey.
274
BURNEY
J. LE BOEUF
period during which females were adapted to the belts. Harnesses were worn at all times; chastity belts were installed for tests only. Observation
periods
and their
duration.
Tests involving estrous females were started 8 days before the female received her first estrogen injection and continued without interruption (except for a few days when the weather conditions made observation impossible) until the ioth day after the female was last mounted. The 8 observation days prior to injections were called the anestrous period; days with mounts in which the female stood and deviated the tail constituted the estrous period; and the io days following the last day of estrus was called the postestrous period. Anestrus and post-estrus combined were called nonestrus. Combinations
or groups
of Ss tested.
Since tests with each S included observations when the animal was tethered and when he or she was roving free, and since each female was tested during estrus and nonestrus, several combinations'of sex and female "reproductive" conditions resulted. These combinations and their abbreviations (called groups) are shown in Table 2 and will be referred to in a later section. TABLE
2
Combinations of subjects tested Groups
Roving Animal
Tethered Animal
M-EF EF-M M-NEF NEF-M NEF-EF EF-NEF NEF-NEF M-M
male estrous female male nonestrous female nonestrous female estrous female nonestrous female male
estrous female male nonestrousfemale male estrous female nonestrous female nonestrous female male
Time
and order
of testing.
Two females were brought into estrus at the start of the experiment while the other two females served as anestrous Ss. When testing was completed on the first two females, they served as anestrous Ss and a third female, used formerly as an anestrous S, was brought into estrus. The last S came into natural estrus during the post-estrous' tests of the first two females tested. Heterosexual pairs and female-female pairs were observed on the same days with the former always tested first in the day. All tests involving
INTERINDIVIDUAL
ASSOCIATIONS IN DOGS
275
femaleswere completedbefore observationof male-malepairs were initiated. Each male was tested IO times with each other male over a period of 30 days. To control for possibleorder effects in the daily tests, individualsof one sex were run first to individualsof the other sex on alternatedays. For example,an estrous female was run to males before they were run to her on one day and the procedurewas reversedon the following day. When two females were being tested on the same day, they were alternatelyrun first and second. Within each group, the order of release or tetheringwas varied in a way which preventedsystematicerror.
RESULTS HETEROSEXUAL INTERACTIONS
Normative post-estrous
data f rom the anestrous, periods of females.
estrous,
and
Table 3 shows the percentageof days that the roving dog visited the tethered dog and initiated a behavioralepisode. Males visited females on 93.lYo of the days that the latterwere in estrus, an increaseover the number of days females were visited when they were in anestrus (78.6%) and post-estrus(72.0%). Similarly,femalesvisited males more frequentlywhen they were in estrus (76.2%) than in anestrus (27.8%) or post-estrus (4255%). Under each female condition,males visited females more often than females visited males. The percentageof total test time spent by the roving dog in the circle with the tethereddog is shown in Table 4. The findings representedin this table supportthe conclusionssuggestedby Table 3. Accordingto these two measures, the sexes are generally most attractedto each other when the female is in estrus. Detailed examinationof Table 3 and 4 shows that one female was far more responsiveto males than were the other three females.Kate'svisits to males totaled 89%, ioo% and ioo% of the tests during her anestrous, estrous, and post-estrous periods, respectively. The response of the other three
females was much lower during their respectiveperiods. Kate also spent more time with males,once she visited them,than did the other females.The magnitude of these individual dif ferences precludes further normative analyses. A more meaningful approach, for our purposes, is to examine the response of individuals, the source of the variation. BehaviourXXIX
I8
TABLE 3 Percentage of days that the roving dog visited the FEMALES
~~~~Tethered
Days MALES CLARK
75
JOHN
12
EDDIE,
62 88
KEN BROADTUS
MEANS
i8 E
8 A
100
8
io PA
100
78
Roving i8 1o E_P 0
100
12
22
20
12
83
100
40
12
100
I10 90
i00
90
25
II
0
100
25
i00
8o
700]
17.2
100
167.495.6
BLANCHE
SPOT
PEGGY
Tethered 8 12 I0 E P A
8 A
100
100
38
50 83
12
I0
E
PA
I17
0 30
30
0
20
0
100
100
100
67
50
83
0 0
100
50 7
Te g A
Roving 8 A
12
I0
E
P
42
0
100
30
89 33
89
40
0
92
20
70
12
5
0
100
40
0
50
0
90
100
100
0
83
0
i00
i00
90
0
100
50
100
100
100
12
100
40
100
182.0
90.0
84.0
4.8
70.0
140]
i00
88 88
Roving
Tethered 12 8 I0 P E
63.2 36.0] 82.8 86.6
44.0]
0 7I.8 20.0
82.2
TABLE 4 Percentage of total test time spent by the roving dog in tethered dog FEMALES
MALES
PEGGY Tethered Roving E A P1 A E
CLARK
2-7
JOHN
I125
KEN
5 33
BROADUS
21
EDDIE,
MENS A= Anestrus
30
72
12
4
I
0
3
82
II
E = Estrus
0
A
12
I
I
I
84
0 14
E__Pi 6
4 15
9
8
-
Ken
$-*9
Broadus
z
X
t
9
7/I I
63.6
O
8
4
7
4
2/62
9 8
7 8
3 2
I0
0
15
0
37177 7/83
7
2 0
2
i5o/i66 I 6/38
90.4
0
0 0
2
I0
56
o
II/I58
7.0
I
9
49
4
9 e
Day of No. MCT Total No. Female NegativeEpisodes MCT Days MCTs SolicitationsProportion Percent
Ist
Tethered -
Clark
9
_1 8 -9
9
John
Q
Eddie Ken
Broadus X 8
8
9 9 < 8
0
io/i6i
6.2
4 0 i6
2/I4I
I14
0/27
0 3.3
I 3 0 7 0
2/61 i1/26
0/80 32/i65 6/1iI2
6/85
3.8 0 194
54
2
i1/64
7.I i.6
I 32
491464
io.6
1I /458
24
called "threat" and "'attack", respectively. Negative episodes are those in which the female responded with threat or attack. Tables 5, 6, 7, and 8 show the response of each female to each male for 3 categories of behavior: the time and frequency of MCTs, the frequency of solicitation behavior, and the percentage of negative episodes. The data in these tables show that Peggy, Spot, and Blanche, each reacted positively to some males and negatively to others, i.e., they showed preferences. This is
280
BURNEY
J. LE BOEUF
best indicated by considering one fenmalein detail. For example, Peggy clearly preferred Broadus and Eddie to Clark and Ken. She vis,ited Broadus and Eddie every day that she was in estrus and spent the majority of her test time with them when they were tethered (Table 3). On the contrary, she rarely visited Clark and Ken and when she did she remained with them for only a short time (Table 4). Table 5 shows that on these few visits, Ken never mounted. Clark mounted Peggy for 3 days but not until the 14th day of her i8-day estrous period. On the other hand, Peggy's visits to Broadus and Eddie resulted in numerous mounts as early as Day 2 and 3, and for a period of i6 and 15 days, respectively. When Peggy was tethered and the males ran free, Ken and Clark did not mount until the 14th day of her estrous period. Once again, Peggy's response to these two males contrasted sharply with her response to Broadus and Eddie, who mounted sooner, on more days and more times per day. Peggy solicited Eddie and Broadus frequently, particularly when they were tethered and she was free. She rarely solicited Clark and never solicited Ken. The proportion of negative episodes to total episodes initiated by the male when he was roving indicates that Clark and Ken did not mount frequently because they were prevented from doing so by the female. Twenty-seven percent of the episodes initiated by Clark and 76%vof those initiated by Ken, Peggy actively repulsed with threats and attack. Markedly lower percentages on the same measure were obtained by the other 3 males: John Eddie (2.8%), and Broadus (0.5%). (3.2%), The response of the other females can be compared to Peggy's (Tables- 6, 7, and 8). Both Spot and Blanche were most positive to Broadus and somewhat less so to John. Spot, like Peggy, was most uncooperative with Clark and Ken. She did not stand to be mounted and she rejected them 9 out of every iO times that they initiated an episode. Blanche was positive to Ken, in that he mounted often and was at times solicited, but she also rejected him frequently. The most interesting point is that Blanche displayed the greatest aversion to Eddie, one of Peggy's favorites. This suggests that the antipathy elicited by a male does not depend on an attribute peculiar to him to which all females respond, e.g., his outward appearance. Eddie obviously affected Peggy and Blanche dif ferently. Kate did not show the antipathies shown by other females. She visited each male every day that she was in estrus and spent from 47% to 68% of her time with them. With Kate, no one male ranks far above or far below any other on any of the measures shown in Table 8. Preferences were suggested in still another type of behavior shown by the female which some investigators have called "teasing" (WHITNEY, 1947).
INTERINDIVIDUAL
28I
IN DOGS
ASSOCIATIONS
A female in estrus often terminates an investigation or mount by running away from the male. The male usually gives chase and after a short run, the female comes to an abrupt halt and may stand once again. If the male does not follow her, the female often returns to the male and solicits his attention (FULLER & DuBuis, I962; LE BOEUF, I967). Females frequently ran to, away from, and back to tethered males in this study and these data are shown in Table 9. The measure is the mean number of circle entrances by the female on days in which she visited tethered males. TABLE 9 Mean number of circle entrances on days in which female visited the male Roving Females Tethered Males Clark John Eddie Ken Broadus Means A
=
E = P =
A I
Peggy P E
A
Spot E P
Blanche A E P
A
Kate E
P
I .5
0
0
I
0
0
I
0
1.3
2.5
I.,
I.8
I
0
3.2
I
0
4.2
I
1.2
2.5
1.3 1.1
I
2.5
T.1
0
3.4
I.5
I
I
0
L.I
2.2
2
I
0
0
I.7
0
0
2.9
0
1.2
2.8
1.2
I
3.0
I
0
4.4
I .4
I
4.2
I
I .4
2.1
1.2
I.2
2.0
0
2.7
o.8
0.4
2.7
0.4
T.2
2.4
1.2
.6
anestrus estrus post-estrus
All females ran in and out of the circle most frequently when they were in the estrous condition. The mean number of circle entrances during this period for each female, with the exception of Kate, reflects the same preferences for particular males which were evident in the measures previously discussed. The conclusion is that females "'teased"preferred males only, or put more objectively, they returned to preferred males more often after leaving them than they did to nonpreferred males. >A comparison e r en c e s.
of present
and
previous
mating
pref-
The various differential responses of estrous females to certain males suggest preferences which are remarkably similar to those shown previously by these same females to the same males during their first and second natural estrous periods. Table io, adapted from BEACH & LE BOEUF (in
282
BURNEY
J. LE BOEUF
press), shows the percentage of male acts (symbols i through 4A in Table i) to which femalesrespondedwith threator attack.Scores from both estrous periods are combinedto form the rejectioncoefficients shown. The similarityand consistencyof present and previous mating preferencesare even more surprisingwhen one considersthe numerousdifferences already pointedout betweenthe two studies, and the fact that a span of two years separatedthe first estrousperiods from those observedin the presentstudy. The presentresultsalso suggestthata female's,preferencesare not attenuated by the artificialinductionof estrus with exogenoushormones. TABLE io Rejection coefficients from two estrous periods for each female with each male (adaptedfrom Beach and LeBoeuf, in press) Female Male
... . . .... ..... .... .... ..... ... .. ..... ...... .. .... ..... . ..... ... .. ................
Clark John Eddie Ken
Broadus
Peggy Spot Blanche Kate ............ .... . . .. . . ....................................................................... 33
6i
55
5
10 0
IO
I5
2
I2
8i 0
33 4
58 59 6
30
12
o
in the circle when females were in nonBehavior estrous. Only a small number of episodes took place during the anestrousand post-estrousperiods of each female. No mountingoccurredand solicitation was rareduringtheseperiods.A few negativeepisodeswere observedduring anestrusbutthey were distributedamongseveralmales.No malewas rejected during post-estrus.More will be said about the anestrousand post-estrous periodsin the next section. during estrus. Temporal changes In additionto displayingindividualbiases to males, females differed in more general ways amongst themselves. Each female responded to the administrationlof exogenoushormoneswith different latenciesto the onset of estrus and its duration(see Figures 4-7). Kate not only failed to show preferences,but was more "social"than other females when she was not in estrus (see Tables 3, 4 and 9). Peggy solicitedfrequentlywhen she was in estrus while Spot solicitedonly rarely.Spot respondednegativelyto most
I
80
70
-
z
-
50
-
I
I
i
I
I
MOUNTS BY MALES
A-
SOLICITATION BY FEMALE
X-
NEGATIVE EPISODES
~~ ~~~~~~~A I
U
40
8
283
DOGS
PEGGY 0-
60
IN
ASSOCIATIONS
INTERINDIVIDUAL
r
A
30
30
/
