Playful Contact in Meerkats

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Ontogeny of Playful Contact in a Social Mongoose, the Meerkat, Suricata suricattaAuthor(s): C. Wemmer and M. J. FlemingSource: American Zoologist, Vol. 14, No. 1 (Winter, 1974), pp. 415-426Published by: Oxford University PressStable URL: http://www.jstor.org/stable/3881999.

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Amer. Zool., 14:415-426 (1974).Ontogeny of Playful Contact in a Social Mongoose, the Meerkat, Suricata suricatta

C. Wemmer and M. J. Fleming

Chicago Zoological Park, Brookfield, Illinois 60513synopsis. Playful contact interaetion within a captive family of six meerkats was ob?served using a rotating focal animal observation method over a 5-month period. Teninteraetion patterns are described. The mother interacted playfully with her familymembers at a low rate while the father and young interacted together far more thanwith the mother. The parents sniffed all family members about equally, but all offspringsniffed the father far more than they sniffed the mother. Dyadic interaetion sequencesbetween the youngsters incorporated two patterns that infrequently occurred in father-young sequences. Eighteen-week profiles of biting and forelimb contact displayed largeoscillations in rate for each animal, but there was little inter-individual concurrence ineach pattern and little similarity in profiles between patterns. Playful interaetion wasdepressed and fluctuations dampened following the birth of an ill-fated litter. Sniffingfluctuated over a smaller amplitude than either of the other two patterns. During the5-month observation period, only the father and one youngster were consistent in theirchoice of recipients of forelimb contact; the other three youngsters altered the propor?tion of contact delivered to their companions. The youngsters did not exhibit signifi-icant changes in the body targets of biting over the time period of the study.

INTRODUCTIONMeerkats are small mongooses restrictedto the desert and savannas of southern Af?

rica where they live in colonies of about25 or more animals (Michaelis, 1972). It isunknown how many families occupy acolony, but it is likely that many of theindividuals are related.

As Ewer (1963) has noted, the familyforms a social unit in which both parentsare of importance. She has shown thatmaternal behaviors in solitary species(grooming, guarding, defense, and playwith the young) are, in meerkats, sharedby the parents or exhibited primarily bythe male.

The mother's contribution during therearing of the young is largely restrictedto the nutritive role. The majority of socialinteraction occurs between members of agiven litter, their older brothers and sis-ters, and their father. Much of this inter?action, at least in captivity, is what would

We would like to thank Leland LaFrance forproviding photographs. Katherine LeClere occa?sionally substituted for one of us in data taking;dictated observation and manuscript were typed byMaria Bukowski and Mary Nichols.

be interpreted as play. Our purpose has notbeen to elucidate the physical or emotionalproperties of play, but to describe a varietyof patterns and relate them to the develop?ing social structure of a captive family.While the patterns of concern occur in avariety of contexts and undoubtedly emergefrom differing motivational states, theywere observed and analyzed in the sameway.This paper presents a quantitative de?scription of playful companion-directedcontact within a group of meerkats, andexamines some differences in the occurrenceof certain patterns with respect to age andsex.

materials and methodsThe subjects of this study were an adultmale and an adult female captured as

young adults in South West Africa andincorporated into the Brookfield collectionin 1970, a young adult female (S) born tothis pair in October 1972, and a female(AB) and three males (BP, LP, RP) born tothe pair on 22 February 1973. A third litterof four was born on 3 June 1973, but theseyoung were smaller than those of the pre-

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416 C. Wemmer and M. J. Flemingvious litter and died within 3 days. Obser?vations terminated when the young were 5months old. RP and LP were removed fromthe colony on 1 August 1973, and a fourthlitter now under study was born to theoriginal mother on 20 August 1973. Theadults were easily distinguished, but theyoung could be identified only by Nyanzoldye markings on the feet, legs, and tail.The group is housed in a glass-frontedcage (96 inches deep X 111 inches wide and84 inches high) containing a simulated con-crete termite mound having four systems oftunnels made of plastic PVC pipe (4 inchesdiameter) (Fig. la). Two short tunnels leadto nest boxes.The substrate is sand; water is providedad libitum, and food is presented at about3:30 every af ternoon. Live crickets andmice are fed twice a week.

Data were gathered using a rotating focalanimal method, and usually single 10-minobservation periods were made for eachanimal between the hours of 7:30 and10:00 am and 3:30 and 5:00 pm. Theamount of time a focal animal was notvisible to the observer was deducted fromthe 10-min period. The amount of obser?vation time for each animal varied from6 hr, 49 min (the mother) to 15 hr, 6 min(LP) and totaled 74 hr, 19 min for all indi?viduals. Table 2 lists the total focal obser?vation time for each animal. Observationsof ongoing social activities were recordedon a dictaphone or on data sheets sub-divided into behavioral categories. Wewere unable to consistently record the avail?ability of the focal animal's peers. A certainamount of time was spent by each animalout of sight in the tunnels, so it is safe tosay that probably all sender-receiver valuesare somewhat inflated. The sniffing andbiting data presented in Figure 4 are basedon both focal and nonfocal observationsfor all animals. All other data were com-piled only from the focal observations oneach animal. For each focal animal the re-cipients of its contact behaviors were re?corded as well as the body target to whichthe contact was directed. When the focalanimal received contact from a companion,the companion's identity was recorded, as

well as the specific target of the contact onthe focal animal's body.description of behavior patterns

The rate of interaetion in meerkats isoften rapid. Consequently, we employed acoarse-grained definition of those patternsthat were quantified. These, as well asothers seen in playful contexts, are listedbelow and illustrated in Figures 1-3. Themajority of the patterns have been filmed(Wemmer and LaFrance).

Grappling (Fig. \b, c). Both animalsstand bipedally and clasp the companionwith the forelimbs, about the neck, shoul-ders or chest. Each animal leans and pushesagainst its opponent while attempting tobite a part of the head or neck.

Wrestling (Fig. 2a, b). This is a ventro-ventral configuration in which one animalreclines on its side or back as its com?panion stands over it with its forefeet onthe floor or on the inferior animal's ven-trum or side. The reclining animal's bodymay be clasped with the forelegs if it issmall relative to the opponent.

Mounting (Fig. 2c, d). One animal sup?ports its forebody upon its companion'sback while clasping the sides somewherebetween the rib cage and the groin. Themounted animal's posture may vary fromreclining on its ventrum to standing onstiffly erect legs with or without the taildeflected laterally. Mount duration variesfrom less than a second to over 1 min.

Stiff-legged rocking gait. Described byEwer (1963) as a response to ground preda?tors, the gait is also seen occasionally asone animal approaches another, but with?out apparent growling. This is an exag?gerated bounding gallop in which the backis strongly arched. It is sometimes per?formed in place with little if any forwardprogression. A rapid rotational movementof the head and neck sometimes impartsan exuberant quality to the gait.

Head-rolling. Described by Ewer (1963)as inhibited biting, this pattern consists ofa single head twist to one side or a seriesof rapid rotations. The mouth is closedand the neck is usually extended. The pat-



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Social Play Ontogeny in Meerkats 417

FIG. 1. a, Meerkat cage containing simulated termitemound with burrows. b, Grappling between siblings.tern is performed during grappling, mount?ing, or clasping or as a response to theapproach of a companion. At times it ap?pears as an apparent response to facial con?tact (as when being pawed, bitten, orsniffed).

Side-pressing. The animal leans againstits companion with its neck, shoulder, orside while standing in one place or slowlywalking forwards. The animal may relin-guish its foot support and flop either on

c, A grappling animal biting its opponent's cheek.

top of or beside the other animal.Clasping (Figs. 2f, 3a). A part of a com-

panion's body is grasped simultaneouslywith the forelimbs. This pattern is a com?mon component of grappling, wrestling,and mounting, but it was only scored whenit occurred independently of these particu?lar body configurations.

Standing-on (Fig. 36). One animal sup-ports its forelegs on top of the torso of aquadrupedally standing or sitting com-



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418 C. Wemmer and M. J. Fleming

?ai

FIG. 2. a, Wrestling between siblings. b, Father-young wrestling; the father is being pawed in theface by the youngster. c, Father mounting his 6-month-old daughter while biting her neck. d, Sib-panion.

Pawing (Fig. 2e). A foreleg is extendedtoward and sometimes touches the com-panion's body. It is not always possible todiscern contact when the motion is fast.

ling mounting; notice the difference in the inferioranimal's stancc compared with c. e, A youngster paw?ing the father's muzzle. /, A uni-directional clasp byanimals in a bipedal stance.During wrestling a reclining animal mayalso repeatedly and alternately strike withthe forefeet at the body of its companion.

Biting (Figs. \b, c, 2c, $c). The durationand orientation of biting is highly variable



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FIG. 3. a, Clasping by animals in a quadrupedalstance. b, An animal standing on its sibling's back.c, A bite directed to the head and unaccompaniedby other forms of contact.

and largely dependent upon the orientationof the body.Virtually all parts of the body receivebites, but biting orientation depends uponthe body configuration of the interactinganimals. Table 1 summarizes some differ?

ences in the predominant biting targetsrelative to the body configuration of thetwo opponents. Not all bites, however, aredelivered by pairs of animals engaged inthe described configurations: one to severalanimals often stand on the sidelines anddeliver occasional bites with otherwise mini?mal contact (without engaging the com?panion with the limbs).

The body targets of sniffing can also beclearly discerned, but the pattern does notoccur under the diversified circumstancesof biting. Figure 4 compares the body tar?gets of these two relatively context-distinctpatterns. While there is a general similarityin the histograms, the greatest differenceoccurs in the regions of the neck, forelegs,and anogenital area. When the differencebetween the two histograms is tested withthe Spearman rank correlation coefficientthe null hypothesis must be accepted: thereis no relationship in the frequencies withwhich body regions are bitten or sniffed(rs = .1516). This supports the contentionthat biting and sniffing arise in differentcontexts, and both patterns will be usedto assess relationships within the group.

0 head , fleg ^ back side agneck shld belly hleg tailFIG. 4. The distribution of bites and sniffs to vari?ous body targets. Abbreviations: Fleg = forelegs (in?cluding feet); Shld. = shoulder; Hlg. = hindlegs(including feet); ag = anogenital region.



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420 C. Wemmer and M. J. Flemingtable 1. Variation in predominant biting targets relative to body configuration of opponents.

SNIFFING

AB(66)

(337)P AB(329)

MOUNTING,LASPING,TANDING-ON


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Social Play Ontogeny in Meerkats 421table 2. Overall frequencies and rates/min (in parentheses) of sniffing, forelimb contact (stand-on, clasp, mount, grapple), and biting for each family member.

appeared to treat her three brothers aboutequally.To test if there was a difference betweenthe three patterns in their distribution tocompanions (i.e., if the patterns were em?ployed differently), each individual's bite,sniff, and forelimb contact scores to itscompanions were tested in pairs with theKendall rank correlation coefficient. Onlybiting and forelimb contact were signifi?cantly correlated for all individuals exceptthe mother (P = .042, .008). In no individ?ual was biting and sniffing significantlycorrelated, and only in the young female(AB) was sniffing and forelimb contact sig?nificantly correlated (P = .042). The nega-

tive S value (1-8) for this particular testindicates the correlation is inverse, andthus, only the distribution of biting andforelimb contact to the companions arepositively correlated.Table 3 presents the distribution of eachfamily member's bites (except the mother)to the body parts of companions. To test thehypothesis that no difference exists betweenthe father and young in the proportion ofbites delivered to the body parts of theyoung, the data were retabulated and testedwith the Chi-square test for two inde?pendent samples. The difference in bitetarget distribution between the father andeach offspring was statistically significant

table 3. Biting target differences between members of a meerkat family. Figures are percentagesof the total number of observations (n) (ag = ano genital region).



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422 C. Wemmer and M. J. Flemingtable 4. Sniffing target differences between membersof a meerkat family.Sending andreceiving head trunk aganimals neck legs tail nBP all sibs 14 13 (5 33LP 11 7 0 18RP 16 11 7 34AB 14 6 7 27Total 55 37 20 112

(df 4, 5; P < .01). When biting target dis?tribution to male siblings was similarlytested between each male and the femalesibling no significant differences werefound. These results should be regarded astentative because the value for the r cate?gories were generated by two subjects.

Similar differences can be seen in the dis?tribution of sniffs between the male andthe infants (Table 4). Ranked in order ofmagnitude the siblings sniffed one another'shead-neck, trunk-legs, anogenital region-tail. The male's head and neck were forthe most part sniffed less by the youngstersthan the trunk and legs, but the male snif?fed the youngster's head-neck and trunk-legregions about equally. The difference is dis?tinct, but the reasons for it are unclear.In summary, the mother interacted withher family members at a low rate and with?out apparent preferences, while the fatherbit and held one male offspring more oftenthan the others. The offspring bit and heldone another more frequently than eitherparent, interacted with the father moreoften than the mother, and exhibited somepreferences among themselves. The parentssniffed all family members about equally,but all offspring sniffed the father morethan the mother or one another. Thereappear to be differences between the fatherand the offspring in the proportion of bitesreceived by the offsprings' body regions.

interaction sequencesPlayful interaction in meerkats shows

many variations in complexity. Simple in?teractions involve brief exchanges betweentwo animals, and complex bouts are pro?longed multidirectional exchanges of con?tact between three to five participants.Analysis of dyadic sequences of all be?haviors is formidable, but it is practicallyimpossible to analyze group participationbecause the actions of all animals are rarelyvisible at once.

Although limited by the recordingmethod, the contingencies of dyadic inter?action can be analyzed from the standpointof body configuration. Several differencesbetween father-young and sibling sequencesare evident in Figure 6. The absence ofgrappling from father-young sequencesclearly results from the size difference be?tween the participants. Grappling is essen?tially a contest of strength that requiresopponents of equal to subequal size. Thefather also exhibits far less mobility thanthe young; he rarely chases them once theydepart, and he departs from the scene ofinteraction far less than the young. In thisconnection the father's centripetal effectupon the young is notable. In the majorityof dyadic exchanges the youngster poten-tiates the interaction by approaching thefather, but the father usually initiates con?tact by clasping or mounting the younganimal. Wrestling proceeds when theyoungster rolls onto its shoulder and thenits back (postures termed Abwehrbuckeland Abwehrstellung byDiicker, 1962). Onceon its back the young meerkat's abdomen isstrongly flexed so that the belly is coveredby the rump and hindlegs (Fig. 2b). In thisposition the animal is able to fend off bitesby kicking, pawing, and feinting with openmouth.

In sibling interaction grappling is oftenthe initial configuration to follow approach(Fig. 6). Together with mounting and clasp?ing it frequently gives rise to wrestling, butas the opponents are matched in size eitheranimal may assume the inferior position.Departing animals are often pursued bythe companion.



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d-YOUNG SEQUENCESAPPROACH

DEPART

YOUNG-YOUNG SEQUENCESAPPROACH

MOUNT 23^? ? GRAPPLE CLASP

CHASE

As the young approached adult size thedyadic interaction with the father acquiredthe sequence characteristics of equal-sizedanimals by the ineorporation of grappling.Equal-sized opponents also are not as adeptat protecting the ventrum by strong abdom?inal flexion.

The above analysis is a rough approxi-mation of play sequencing based on a rela?tively small sample of sequences spread overa 1-month period when the young were 2 to3 months old. Other configurations alsooccur, but we were unable to describe themadequately, as their appearanee was brief,transitory, or highly variable. It is impor?tant to emphasize that the flow of behaviorbetween any two individuals involves split-second timing and that the temporal resolu?tion of a meerkat is clearly several orders ofmagnitude greater than that of the humanobserver.

FLUCTUATIONSAND ONTOGENTICCHANGES

Figure 7 presents three 18-week behav?ioral profiles for each family member. Sniff?ing for the most part occurred at a steadylow level amoung the young and mother,but was more variable in the father. Aslight peak in the male's sniffing a weekafter the birth of the ill-fated second litterwas possibly a response to the mother's post-partum estrus, though the birth of a subse?quent litter of three indicates she also cameinto heat at about week 17. Neverthelessthe mother was the recipient of most of themale's

sniffing during week 14. It is clearthat with the exception of the adult male,sniffing varies over a smaller range than theother two measures.

Fluctuations of considerable magnitudecharacterize both the biting and forelimbcontact curves, but the peaks and troughsdo not exhibit a strong concurrence eitherbetween individuals or the two measures.FIG. 6. Interaction sequences between the fatherand his young (male-young) and between siblings(young-young) based on 60 complete father-youngsequences and 51 complete sibling sequences re?corded during May 1973. Uncommon patterns havebeen deleted from the sequence, and hence percen?tage values for each pattern do not always total100%. Figures in parentheses represent the numberof cases from which percentage was calculated.



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Social Play Ontogeny in Meerkats 425limb contact suggest that sniffing has littlerelationship to play activities in all animalsexcept the father. In his case all three vari?ables have similar profiles.To test whether the proportion of con?tact delivered by given individuals to theircompanions change over time, each sender'scomposite scores for stand-on, clasp, mount,and grapple were combined into 3-weektotals for each receiving animal, and thedata were tested with the Friedman two-wayanalysis of variance (Table 5). Only thefather and one male youngster (RP) ex?hibited recipient rank totals that differedsignificantly (P < .01, < .02). The propor?tion of forelimb contact delivered by thesetwo animals to their companions was con?sistent over the period of observation. Inthe remaining three young, the proportionof companion-directed forelimb contactvaried from period to period.To test the hypothesis that there is nochange in the proportion with which dif?ferent body regions received sibling-directedbites, the total number of bites was tabu-lated for 3-week periods according to bodyregion (Table 6). Rank totals for differentbody regions are significantly different (P


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426 C. Wemmer and M. J. Flemingare established through playful activity asthe litter develops. This is supported by theobservation that the flow of contact be?tween some individuals changes throughtime.

When female AB was 9 months old themother meerkat came into estrus a weekafter giving birth to another litter. Bothfemales followed the father about the cageand copulation was judged to have occur?red in the nestboxes. In the midst of thissetting the mother and AB were discoveredengaged in a protracted fight. The patternsof play were executed with tenacity, bothanimals suffered bruises and cuts, and ABwas removed from the group. It is reason-able to hypothesize that AB was recognizedby the mother as a sexually mature femalevying for a limited and heretofore uncon-tested resource.

The father's centripetal effect upon theyoung deserves further mention. He isoften sought as a landmark about which theinteraction evolves. At other times an in-animate feature of the cage?often a con-cavity such as an empty food pan, watertrough, or short burrow?acquires a similarfocal interest to the young, and the arenaof interaction becomes spatially defined bythe locality of the object. The observationis subject to a variety of interpretations,but the defensive and competitive aspect ofthe patterns are obvious. Clearly strenuouscompanion-oriented contact is often preci-pitated by a referent. Playful interludes aswell as the episode between the mother andAB often have this feature in common.

Several time changes in behavior werenoted. Rates of biting and forelimb contactexhibited the greatest fluctuations in allanimals but the mother. Horwich (unpub?lished) has observed similar ontogenetic os?cillations for a variety of behaviors inseveral primate species, and interprets themto arise from emotional crises attendant tosocialization and integration into the group.The most pronounced depression in activity(biting and forelimb contact) coincided

with the birth of a litter. However, otherfluctuations probably resulted in part frominadvertent changes within the captive en?vironment such as heating failures withinthe building.Siblings exhibited no changes in bitingtargets to one another's bodies over the 18-week period. It is likely that if the orienta?tion of bites to companions differs at anytime in development it is during the earlyonset of sibling interaction, a period forwhich quantitative data are lacking. At thistime (age 4 to 6 weeks) the mechanics ofplayful interaction have a different quality.Body movement is jerky, and the animalslack the coordination to grapple. Standing-on and clasping are very common, but par-ticipants frequently lose balance and fallover.

Three of the youngsters displayed changesin the proportion of forelimb contactdirected to their siblings during the courseof the observations. This suggests that theavailability of the companions changed,that the preference of the subjects changed,or that both factors determined the chang?ing proportions of recipients. It appearsthat qualitative changes in interaction occurvery early in development. Quantitativechanges on the other hand probably occurthroughout the growth period of the youngand into their adulthood. Such changes areprobably responses to events within thecolony (i.e., births and deaths) and mani-festations of changing social relationships.

REFERENCESDucker, G. 1962. Brutpflegeverhalten and Ontoge-nese des Verhalten bei Surikaten (Suricata suri-catta Schreb, Viverridae). Behaviour 19:305-40.Ewer, R. F. 1963. The behavior of the meerkat

(Suricata suricatta Schreber). Z. Tierpsychol. 20:570-607.Michaelis, B. 1972. Die Schleichkatzen (Viverriden)Afrikas. Saugetierk. Mitt. 13:1-110.Wemmer, C., and L. LaFrance. (submitted). Playbehavior in the meerkat. 16mm black and whitefilm submitted to Encyclopedia Cinematographia.

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Playful Contact in Meerkats