Behavioral Beology Vol 6 No, 2: 199-208 Investigating structure and temporal scale in social organizations using identified individuals Hal Whitehead Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1, Canada ‘Studies of individually identified animals can produce substantial data sets containing information on the structure and temporal scale of social organizations, However, methods of analyzing such data are not well established, Important features of a sorial ‘organization are revealed by plotting the rate of persistence of the associ 1s between pairs of individuals over a range of, time lags (lagged association rate). The consistency of long-term relationships can be characterized using the rate of association of pairs of individuals between their first and last observed associations (intermediate association rate). A hierarchical series of ‘models featuring exponentially decaying lagged association rates may be fitted to these data. This technique reurieved the ‘essential parameters of five simulated social organizations and, when used on real data, portrayed the essential features of the patterns of temporal change in relationships between animals. The method should be especially useful for analyzing fission fusion societies containing 10-10,000 individually identifiable animals. Ky words: association, individual identification, social ‘organization, temporal scale, [Bea Kol 6:199-208 (1995)] AR Secate more of social organization is vial if we are to gain understanding of the ecological pressures affect ing sociality (Myers, 1983). However, our current quantitative methods of describing social organization are relatively crude, and rarely incorporate the significant dimension of temporal studies of captive or wild populations generate large *s of daca on associations between known identified individuals. A standard and frequently employed procedure is to calculate indices of association between each pair of incl viduals (Cains and Schwager, 1987). Maximum spanning trees, cluster analyses, and multidimensional scaling are often used to assist interpretation of these resulting association ma- tices (Morgan et al., 1976; Penzhorn, 1984). A simple way of incorporating temporal changes in relationships into such e- scriptions i to produce different association matrices, and dis plays of them, for different time periods (e.g., Lott and Minta, 1983; Morgan et al, 1976). ‘This approach suffers from several drawbacks. The time in- tervals must be long enough for sufficient data to be collected within each (0 provide a reasonably accurate association ma- trix. This changes in social organization over sill time in= tervals are not examined. Second, the common techniques tused to uncover social organization from association matrices become unvieldy with data sets containing hundreds or ‘thousands of individuals, Nonmetric multidimensional scaling and some of the preferred clustering techniques, such as a= ferage linkage clustering, are often hard to implement on, large datasets, and the resulting plots or dendrogram usially contain (oo much information to be assimilated easily. How ever large data sets from many animals, especially when col lected over fong time periods, potentially contain very de- ied information about social organization, Finally, and pethaps most importantly, this approach does not quantify hhow the relationships between animals change over time. Received 4 August 1998; revised 22 March 1004 i 15 June 1904 1045 2240/8/$5.00 © 1995 Internacional Society for Behavioral Fcaogy I aceepiance There is no standard way in which the variability of social relationships over different time periods can he described. Here, I develop a technique which uses data on associations between identified individuals to investigate how relationships between animals change over time, The methods of displaying information on social organization used by Underwood (2981) and Myers (1983) inspired the development of this technique, a form of which has been used by Whitchead et al. (1991)'to examine social organization in female sperm whales (Piyscer macrocphalus). The technique produces a simple portrayal of the principal temporal elements of the social organization of a population. In many cases, it is possible to fit quantitative hierarchical models to the data based on exponential decay in the prob- ability of an association persisting between two individuals. ‘The models allow several different mechanisms of association and disassociation to operate at the same time between pairs of individuals within a population (e.g, constant companion: ships, temporary feeding associations, passing “tourist” rela. tionships, death). These models will be appropriate in the many cases when the probability of disassociation is indepen- dent of the duration of an association, Included are most in- stances when disassociation might be triggered by the behay- jor of a third animal, or an unpredictable environmental event, or when the animal’ behavior towards one another does not change systematically during the length of the asso- ciation. Excluded are situations where association and disa- sociation are related to external cycles, where animals tend to associate for a fixed time (eg., an estrous or breedling period), ‘or change their behavior toward one another as the associa. tion progresses (c.g., a courting pair). However, these models would appear wo be appropriate approximations for many cog- nitively advanced animals which are flexible in their behavior and live in fissionsfusion societies. ‘The method models the temporal duration of associations, bewicen individuals, and does not give a full description of the society; This has the advantage that many forms of social organization are included, However, once a particular set of parameters has been selected by the method described here, further tests may be required to elucidate deuails (eg. distin.200 ishing between casual acquaintances within a society and ‘migratory individuals passing through). Twill descrihe the method and show how it works on sim ulated data from five different social organizations, demon: strate its use on real data, and then discuss its advantages and limitations. For those interested in using the method. Appr dix A steps through a simple example and Appendix B shows how the method can be modified to be used on data in which not all associates of an identified individual were necessarily recorded. Analyzing the temporal scales of a social organization The data T assume that a population is observed during several short periods of time, These need not be regularly spaced but Should be at integer multiples of some time unit. During each observation period a number of “key individuals” are ob- served (not all Key individuals in the population need be ob- served each period), and the identities of the associates ofthe key individuals are recorded. Key individuals can be recorded, as associates of other key individuals and often all individuals in a population may be considered key. However, there are circumstances in which itis useful to restrict the animals that will be considered key (see below). I will initially assume that all current associates of key individual are recorded, but the method can be modified to permit the incomplete recording, of all asociates (see Appendix B). The method of determin- ing associates could be spatial (e.g, "seen within x meters of key individual"), temporal (eg., “Seen within x min of key individual”), or behavioral (e.g, “groomed key individual ‘during observation period”) “Thus the data set consists, for each observation period, of te identities of the Key individuals seen during the period and, for each key individual, the identities ofits associates, Sotection of key individuals Sometimes it may he illustrative 10 consider one class of in- dividuals (e.g., males) as key individuals, and another as as- sociates (eg, females). In this way it is possible to examine how relationships berween individual males and individual fe- males change with time. Another reason to restrict the selection of Key individuals may be to reduce bias caused by oversampling animals large groups. Suppose two animals are considered to be associated HM they are members of the same group, and that group size varies, Associations between members of large groups will of ton be less persistent than those between members of small groups. If, during any sampling period, groups are encoun- tered with probabilities proportional to their size, then asso- ciations between members of kirge groups will be preferen- tially sampled. Then the mean persistence of randomly chosen associations between members of the population vill be underestimated. In this case restricting the number of key individuals chosen from any group © the minimum group ‘sie will remove bias, We may be more interested in the persistence of the aver age association of a randomly chosen’ member of the pops lation than the persistence of a randomly chosen association benseen members of the population. If groups are sampled with the same probability regardless of size and all members ‘of each sampled group are considered Key, then we examine the persistence of randomly chosen associations. To examine the persistence of the average association of a randomly cho- sen member of the population, the number of key individuals ‘chosen from any group should be restricted to the minimum size of a group in the population, Hchavioral Ecology Vol. 6 No. 2 aged association rates For any discrete length of time, d, and for each key individual, A, the pairs of sampling periods that A was identified «time units apart, ate listed (indexed by J): (ura tana * 5 Caw thas ts vo Caja bya + so.» Iethe timber of assoc ates that were seen with Key individual A at both time f,;,and, time ye * dis (A, j, d), then the total number of repeat associations in the daia set after time Tags of dis ao = LY ana ‘This will include some repeat associations wviee, if, for in stance, both A and B were observed as key individuals and associating with each other at times fand (+ d ‘Let gd) be the probability that if Bis an associate of A at time & them iti an associate d units later. Fassume that ge) is independent of t, so that the process is stationary in the Satisitcal sense. Ifthe number of asociates of Key invidval at time Fs NA, ), then the expected number of repeat associates observed with key individual A at both times ys and tat is NA, f,.)-e(@). Thus, the expected number fof repeat associations alter lags of is BCU] = EY MA, dete @ ‘Then, combining Equations (1) and (2), the probability of association with a lag of d, g(«), can be estimated from: Pee) TONGA ay aa (a) is the association rate between individuals after a ime lag d from a previous association, I call g(@) the “lagged as- sociation rate.” The lagged association rate is, in many eon- texts, the same as Underwood's (1981) “proportion of com panions remaining,” and the “reassociation rate” of Whitehead et al. (1991). (1 have been shown that “reassocia- sion rate” is a confusing term in this context, and so I suggest its replacement with “lagged association rate. Plots of f(a) against d, such as in figure $ of Underwood. (1981) and on the time axis of figure 3 of Myers (1983) and, Figures 1-4 of this article can reveal much about the social ‘organization of a population. If £(¢) is roughly constant over a certain range of d, then this suggests that there is litle ‘change in the rate of association ver this time scale, On the other hand, if g(a) falls dramatically, then there is significant disassociation aver this Gime scale. Distinguishing betwen reassociations and permanent ‘companions Plots of lagged association rates against time lag showing a similar shape, with a fairly constant, but nonzero, g(a) for large d, can be produced by very different models of social ‘organization. For instance Figure Ie both show falls in the lagged association rate, {(d) over periods of 1-30 days, fol lowed by constant, nonzero g(a). But (as explained below) Figure Te represents data from a simulated population in which pairs of individuals with casual relationships repeatedly reassociated by chance in a stall population, whereas Figure 1d represents a population in which there were permanent companions who associated constantly with one another. How can these situations be distinguished? Ifa population estimate of the number of potential associ ates of a key individual, P is available (eg, from mark-recap- ture methods on the identification data), then an estimate of the lagged association rate for all lags d in the case of com- pletely random association isWhitehead + Analyzing social organizations te | Lagged association rate Time lag in days aa = MP “ where Ais the mean number of associates of a key individual in any observation period. This “null” association rate can be compared with the lagged association rates calculated from, the data as described above. In many cases F the number of potential associates ofa key individual, will simply be the pop lation size minus one. Additionally, or aernatively when the number of potential associates is not known, the association rates of pairs of indi- viduals during observation periods benween their first and last recorded associations can be examined (Whitehead et al, 1991), An “intermediate association rate" is calculated in a similar way to the lagged association rate, except that only time periods between the first and last recorded association, ‘of cach diad are considered. An intermediate association of lag d between Key individual A and individual 2 is recorded, if B was observed associated with Ad time units after their first association (and 2d time units before their last associa- Figure 1 Lagged aswciation rates (@) cover diferent time lage caleue lated from simulated observa. tions of members of five social ‘organizations. In each ease the theoretical lagged sociation rate is shown by the curve, For simulations (e) and (a), inter Imeeliate association rates are shown by &. tion) or d time units before their last association (and 2d time Units after theit first association). Let L(d) be the number of such intermediate associations of lag d, ‘The number of po- tential intermediate associations of lag d, Q(d), is calculated bby summing for each Key individual 4, and each associate 2, the number of occasions that A was observed d time units alter the first association of B with A or d time units before the last association of Bwith A (which ever interval is smaller) ‘The intermediate association rate is then Kd) = 1/0 ‘The intermediate association rate is an estimate of the probe ability that associates remain associated between theit frstand last identifications together. It will approximate 1.0 if asoct ations with long lags are between members of permanent sroupings which do not disassociate between observed asso- iations (c.., simulation d below; Figure 1d). Alternatively, if long-term reassociations often follow periods of separation 6)