American Journal of Primatology 3R25-37 (1995)

Model Life Table for Captive Chimpanzees

BENNETT DYKE', TIMOTHY B. GAGE', PATRICIA L. ALFORD3, BRENT SWENSON4,
AND SARAH WILLIAMS-BLANGERO'
'Department of Genetics, Southwest Foundation for Biomedical Research, San Antonio,
Texas; 'Department of Anthropology, State University of New York at Albany, Albany;
'University of Texas M.D. Anderson Cancer Center Science Park, Department of Veterinary
Science, Bastrop; 4Yerkes Regional Primate Research Center, Emory University,
Atlanta, Georgia

Mortality statistics from three captive populations of chimpanzees (Pan

troglodytes) were combined to generate standard model life tables for each
sex in this species. The model is compared to an estimate of survivorship
of a group of wild animals, and is applied to an incomplete data set to
illustrate how the model may be used t o extend estimates of mortality
statistics to missing older ages. o 1995 Wiley-Liss, Inc.

Key words: mortality, chimpanzees, survivorship

INTRODUCTION
High quality mortality statistics for nonhuman primates are difficult to ob-
tain. In the wild, ages are usually not known with precision, and deaths often must
be inferred from prolonged absences from populations whose boundaries may be
poorly defined. Data for captive animals are usually more complete, although
culling and interference with social and breeding structures can seriously distort
estimates of underlying vital rates. Most troublesome, however, is the fact that
both wild and captive population sizes are extremely small by demographic stan-
dards (N 5 lO,OOO), which results in considerable variability of the measures
computed from them. Over the past few years we have been developing methods to
circumvent these deficits in nonhuman primate mortality data.
Our ultimate goals are twofold. First, we wish to develop standardized mor-
tality measures for individual breeding populations that can be used in epidemi-
ological monitoring, modeling future growth and costs, etc., at the colony level.
Second, we are interested more generally in understanding the contribution of
mortality t o the life history of the species. A common demographic tool for both
these purposes is the model life table. Model life tables consist of a series of age-
specific death rates separated uniformly by small regular differences in the entries
for each age. Although they do not represent the rates found in any specific pop-
ulation, they are based on empirically derived population statistics and are usually
constructed as a family of tables encompassing a wide range of known demographic
experience. Because entries in these tables are regular, complete, and easy to

Received for publication October 12, 1993; revision accepted January 14, 1995.
Address reprint requests to Dr. Bennett Dyke, Department of Genetics, Southwest Foundation for Bio-
medical Research, San Antonio, TX 78228.

0 1995 Wiley-Liss, Inc.

26 I Dyke et al.
control, they are used to interpolate and smooth mortality statistics derived from
data that are biased, incomplete, or based on samples of inadequate size. Model
tables are also useful as standards against which to compare observed rates, and as
a source of plausible approximations for simulation and other modeling studies.
The best known human model life tables are those of Coale and Demeny [1983],
which are based on a survey of hundreds of national demographic registries and
censuses collected over the past century. We have developed model life tables for
the larger Old World monkeys [Gage and Dyke, 1988,19931 and the smaller New
World monkeys [Dyke e t al., 19931. In this paper we present model life tables for
chimpanzees. We consider this effort to be of particular importance because of the
seriousness of the management and financial problems associated with lifetime
maintenance of a large number of animals already in U.S. colonies, and the need
to plan for long-term survival of self-sustaining breeding groups.

METHODS
With vital statistics records for no more than a few thousand chimpanzees ever
in captivity in Europe and North America, we clearly do not have enough data on
chimpanzee mortality to construct multiple tables directly from registries and
censuses as has been done for humans. Our approach is first to develop a mortality
standard by fitting a mathematical model to a n aggregate of high quality vital
statistics data taken from several populations of the species of interest. Next, this
standard which represents the generalized pattern of mortality characteristic of
the species, is used with simple logit regression methods to smooth and extend
mortality schedules of individual populations in which numbers of deaths may be
small, particularly in the older ages.

Development of the Mortality Standard

Development of a mathematical model of chimpanzee mortality is similar to
that reported in Gage and Dyke [1988, 19931 and Dyke et al. [19931. The method
depends on the following assumptions:
1. For any given species there is a n underlying schedule of sex- and age-
specific probabilities of dying t h a t is the result of the interaction of intrinsic bio-
logical determinants and extrinsic environmental factors.
2. Death rates observed in any population represent a sample of a universe in
which some underlying mortality pattern is operating and observed mortaIity
statistics can be used to estimate the underlying schedule. A corollary to this
assumption is that the better the quantity and quality of data available, the more
closely the observed pattern reflects underlying mortality.
3. Under a variety of circumstances, a n estimate of the underlying schedule is
preferable to a life table computed directly from the data. This is particularly true
when critical data points are missing from observed data.
The construction of mathematical mortality models has a long history. The
best-known model is that of Gompertz [18251, who found that a simple exponential
function fit the distribution of deaths after childhood in human populations. This
equation was modified in 1867 by Makeham [Makeham, 18671, who found that
incorporating a n additive constant to the Gompertz function improved the fit. The
Gompertz and Makeham functions have been used extensively in actuarial studies,
where the principal concern has been determining costs of life insurance for adults.
Although these functions have occasionally been applied to nonhuman popula-
tions, they do not serve as models of mortality over the entire life span, since they
 Chimpanzee Life Table / 27
TABLE I. Population Characteristics
Deaths Living population
Population Total records Males Females Total Males Females Total
I 485 64 60 124 103 135 238
I1 320 40 27 67 109 134 243
I11 683 176 171 347 85 123 208
Total 1.488 280 258 538 297 392 689

ignore deaths in the earliest ages. Early attempts were made to develop models
that include effects of both early and late mortality IThiele 18711, but these equa-
tions remained unusable because they had no simple analytic solutions. It was not
until computerized numerical analysis became available that solutions to complex
equations such as these became practicable.
In 1979, Siler published a three-component competing hazards model which he
applied to the analysis of mortality in a series of wild animal populations [Siler,
19791. The rationale for the equation was the 1935 suggestion of Pearl and Miner
that in most organisms, three components of mortality can be distinguished [Pearl
& Miner, 19351. These are shown algebraically in Equation 1

mt = aleblt az + agebst (1)

where m, is the death (hazard) rate due to the additive effects of the three compo-
nents a t age t, the a parameters represent the force of mortality due to each of the
three components a t birth, while the b parameters express the rate at which each
mortality component changes with age. The three constituents making up the
right-hand side of the equation are (subscript 1) a component representing imma-
ture mortality that declines exponentially with age, (subscript 2) a residual com-
ponent representing a constant force of mortality that is independent of age, and
(subscript 3 ) a senescent component representing mortality that increases expo-
nentially with age. To put equation (1)in historical perspective, Siler added the
immature component to the Gompertz-Makeham equation to make it cover the
entire life span.
Our method consists of fitting the Siler equation to observed mortality data
that are as complete as possible, and which extend over the entire age span. These
data should come from as many sources as possible so that the standard will not be
based on the idiosyncrasies of any single mortality schedule. In practice, this
means that at this stage, statistics must come from captive colonies.
Data used in this analysis were extracted from records of three major breeding
colonies in the United States. Although the primary assignment of the animals in
these colonies has been to breeding programs, some have been used as subjects in
biomedical and other research projects a t various times in their lives. However, no
deaths attributable to research or experimental procedures were found in the
records, and we have not attempted to correct for any biases that might be intro-
duced by differential use or caging. Table 1 summarizes the data.
A period mortality analysis was done for each colony using a software package
that applies a longitudinal mortality analysis using right-censored, left-truncated
animal colony data [Dyke, 19891. Of the 1,488 animals for which records were
available, 142 were excluded because of an unknown birth date, gender, or cause
of exit from a colony, leaving 1,346 on which analyses were based. Of these, 1,073
were known to have been colony-born (not necessarily in their current colony),
28 / Dyke et al.

0.0 ; I I I I I I
0 10 20 30 40 50 60

Fig. 1. Sex-specific proportions surviving by age for three colonies.

with the remaining 273 presumably captured in the wild. Sex-specific proportions
surviving by age for each of these populations are shown in Figure 1.
Three points are worth noting:
1. There is considerable variability between populations with respect to both
levels of survival and the maximum ages to which the curves extend.
2. The individual curves tend to vary erratically from age to age.
3. Male survival rates are lower than those of females.
To reduce some of the effects of small sample size, we combined the data from
the three colonies, keeping the sexes separate, and reanalyzed. We then fit the
Siler model to each of the resulting survivorship distributions using the nonlinear
optimization algorithm MODFIT LMcIntosh and McIntosh, 19801.
Using the Standard to Estimate Mortality in Individual Populations
One limitation of fitting the Siler model directly to data taken from individual
populations is that reliable survivorship statistics over the entire life span are
required. This means that the model cannot be used with fragmentary mortality
data (for example, in recently founded populations that contain no older animals,
or small colonies where by chance deaths have not been observed at some ages). In
these cases, we have taken a n indirect approach, using a standard mortality table
based on a fit of the Siler model in combination with the logit model life table
technique of Brass [1971]. This method is based on Brass’ observation of the ap-
 Chimpanzee Life Table I 29
TABLE 11. Parameter Values of the Siler Model
a, b, a? a- b, RMSE*
Males 5.8611 31.0920 0.0249600 0.000094935 0.17331 0.01289
Standard deviation 3.2142 17.3280 0.0009146 0.000096840 0.02817
Females 2.8640 21.3110 0.0166880 0.000052596 0.14691 0.01555
Standard deviation
~~
1.0472 8.0052 0.0004601 0.000031767 0.01348
*RMSE = root mean squared error

proximate linearity between the logit transforms of survivorship schedules of spe-

cies with similar life spans. This relationship makes it possible to fit a standard
schedule to observed data with ordinary linear regression having the form
logit(1,) = A + B . logit(l,,) (2)
where t is age, I, are the age-specific proportions surviving observed in the popu-
lation, I,, is the same statistic taken from the standard model (see Table 111), and
the A parameter is the constant of the regression, which reflects the difference in
level of mortality (or expectation of life) between the observed and standard sur-
vivorship schedules. B is the slope of the regression, which depends on the ratio of
mortality during the early years to that occurring toward the end of life, holding
level of mortality constant.
RESULTS
Mortality Standard
Table I1 gives the final parameter estimates and their standard errors for the
fit of the Siler model to the combined male and combined female mortality sched-
ules, respectively. The standard errors are in general quite large because of the
relatively small numbers in each age class and because of the ill-conditioned na-
ture of the solution of the fit [Gage and Dyke, 19861. On the other hand, the fit of
the model is quite good, as can be seen from Figure 2a and b which show fitted
distributions of proportions surviving superimposed on the observed aggregate
values for males and females, respectively.
The standard model curves fit quite well, and as can been seen from the
figures, generally fall between the dotted lines in the figures representing 95%
confidence bands defined by points lying 1.96 standard errors above and below
observed proportions surviving from birth to each successive age. Standard errors
of proportions surviving are given by

where 1, is the proportion surviving from birth to age t, q iis the probability of dying
during the yearly interval beginning at age i, niis the number exposed to risk of
dying, and dithe number dying during the interval [see Lee, 1980, p. 911.
Acceptable goodness of fit is also indicated by the low RMSE values (repre-
senting mean squared errors between observed and fitted points for the series) in
Table 11: 0.013 and 0.016 for males and females, respectively. Model curves are
considerably smoother than the observed survivorships, and they extend well into
the older years.
Model life tables for each sex, computed from the composite 1, values, and
including a column of logits (that is, Z,) are shown in Tables IIIa and IIIb.
30 / Dyke et al.

a Males

L
0
n
0
L
a

0 10 20 30 40 50 60

Age

1.o
b Females

0.8

u
l
c
.-
------- Observed
.->
>
L
3 0.6 Fitted
rn
rn
C
.-0
0.4
0
CT
2
a
0.2

0.0 I I I I I I i
10 20 30 40 50 60 70

Age
Fig. 2. a: Distributions of proportions surviving from the Siler model superimposed on schedules for males
computed from aggregate values from three colonies. Dotted lines represent 95%confidence bands. b Distri-
butions of proportions surviving from the Siler model superimposed on schedules for females computed from
aggregate values from three colonies. Dotted lines represent 95%confidence bands.
 Chimpanzee Life Table / 31
TABLE IIIa. Standard Chimpanzee Model I ife Table (Males)

0.000 1.0000 0.1618 2.1121 0.0766 20.7573 20.7573 Infinity

0.083 0.8382 0.0150 0.1819 0.0693 20.6807 24.6720 -0.8225
0.167 0.8256 0.0031 0.0368 0.0687 20.6114 24.9648 -0.7774
0.250 0.8231 0.0022 0.0259 0.0685 20.5427 24.9581 -0.7687
0.333 0.8213 0.0021 0.0251 0.0684 20.4742 24.9287 -0.7626
0.417 0.8196 0.0021 0.0251 0.0682 20.4058 24.8975 -0.7568
0.500 0.8179 0.0021 0.0251 0.0681 20.3376 24.8661 -0.7510
0.583 0.8162 0.0021 0.0251 0.0679 20.2695 24.8347 -0.7453
0.667 0.8145 0.0021 0.0251 0.0678 20.2015 24.8032 -0.7397
0.750 0.8128 0.0021 0.0251 0.0677 20.1337 24.7717 -0.7341
0.833 0.8111 0.0021 0.0251 0.0675 20.0661 24.7401 -0.7285
0.917 0.8094 0.0021 0.0251 0.0674 19.9986 24.7084 -0.7230
1.000 0.8077 0.0490 0.0251 1.5759 19.9312 24.6766 -0.7175
3.000 0.7682 0.0491 0.0251 1.4986 18.3553 23.8953 -0.5990
5.000 0.7305 0.0492 0.0252 1.4250 16.8567 23.0764 -0.4985
7.000 0.6945 0.0494 0.0253 1.3547 15.4317 22.2190 -0.4107
9.000 0.6602 0.0497 0.0255 1.2876 14.0770 21.3221 -0.3321
11.000 0.6274 0.0501 0.0257 1.2233 12.7894 20.3855 -0.2605
13.000 0.5959 0.0507 0.0260 1.1616 11.5661 19.4089 -0.1942
15.000 0.5657 0.0516 0.0265 1.1022 10.4045 18.3930 -0.1321
17.000 0.5365 0.0528 0.0271 1.0447 9.3023 17.3392 -0.0731
19.000 0.5082 0.0545 0.0280 0.9886 8.2577 16.2499 -0.0163
21.000 0.4805 0.0569 0.0293 0.9336 7.2690 15.1288 0.0391
23.000 0.4531 0.0602 0.0311 0.8790 6.3354 13.9809 0.0940
25.000 0.4258 0.0650 0.0336 0.8240 5.4564 12.8131 0.1494
27.000 0.3982 0.0717 0.0372 0.7678 4.6324 11.6342 0.2065
29.000 0.3696 0.0810 0.0422 0.7093 3.8646 10.4551 0.2669
31.000 0.3397 0.0941 0.0494 0.6474 3.1553 9.2887 0.3323
33.000 0.3077 0.1122 0.0595 0.5809 2.5078 8.1495 0.4054
35.000 0.2732 0.1373 0.0737 0.5089 1.9269 7.0534 0.4893
37.000 0.2357 0.1716 0.0938 0.4309 1.4181 6.0170 0.5883
39.000 0.1952 0.2177 0.1221 0.3480 0.9871 5.0559 0.7081
41.000 0.1527 0.2786 0.1618 0.2629 0.6392 4.1845 0.8566
43.000 0.1102 0.3567 0.2171 0.1811 0.3762 3.4142 1.0444
45.000 0.0709 0.4529 0.2928 0.1097 0.1951 2.7528 1.2866
47.000 0.0388 0.5650 0.3937 0.0556 0.0855 2.2040 1.6051
49.000 0.0169 0.6854 0.5214 0.0222 0.0298 1.7678 2.0326
51.000 0.0053 0.8011 0.6682 0.0064 0.0076 1.4409 2.6167
53.000 0.0011 0.8960 0.8116 0.0012 0.0013 1.2166 3.4264
55.000 0.0001 0.9584 0.9202 0.0001 0.0001 1.0831 4.5586
57.000 0.0000 1.0000 2.0000 0.0000 0.0000 0.0000 6.1488

Estimating Mortality in Individual Populations

To illustrate how the Standard model may be used to estimate a mortality
schedule for an individual population, we used the logit regression method to fit
the l,, values given in Tables IIIa and b to the 1, values for each sex of one of the
populations used in making up the standard. Results are shown in Figure 3a and
b, where again, dotted lines represent 95% confidence bands. Both curves are
extended beyond the maximum observed ages.
32 / Dyke et al.
TABLE IIIb. Standard Chimpanzee Model Life Table (Females)

0.000 1.0000 0.1069 1.3549 0.0789 29.4441 29.4441 Infinity

0.083 0.8931 0.0201 0.2436 0.0737 29.3652 32.8792 - 1.0615
0.167 0.8752 0.0046 0.0552 0.0728 29.2915 33.4692 -0.9738
0.250 0.8712 0.0019 0.0232 0.0725 29.2187 33.5398 -0.9557
0.333 0.8695 0.0015 0.0178 0.0724 29.1462 33.5215 -0.9482
0.417 0.8682 0.0014 0.0169 0.0723 29.0738 33.4879 -0.9425
0.500 0.8670 0.0014 0.0168 0.0722 29.0015 33.4518 -0.9372
0.583 0.8658 0.0014 0.0168 0.0721 28.9293 33.4152 -0.9320
0.667 0.8645 0.0014 0.0167 0.0720 28.8572 33.3785 -0.9268
0.750 0.8633 0.0014 0.0167 0.0719 28.7852 33.3417 -0.9216
0.833 0.8621 0.0014 0.0167 0.0718 28.7133 33.3049 -0.9166
0.917 0.8609 0.0014 0.0167 0.0717 28.6415 33.2680 -0.9115
1.000 0.8597 0.0330 0.0168 1.6911 28.5698 33.2311 -0.9065
3 .OOO 0.8314 0.0330 0.0168 1.6353 26.8787 32.3297 -0.7978
5.000 0.8039 0.0331 0.0168 1.5813 25.2434 31.3992 -0.7056
7.000 0.7774 0.0332 0.0169 1.5289 23.6621 30.4389 -0.6252
9.000 0.7516 0.0333 0.0169 1.4782 22.1331 29.4485 -0.5535
11.000 0.7266 0.0334 0.0170 1.4289 20.6549 28.4275 -0.4887
13.000 0.7023 0.0336 0.0171 1.3810 19.2261 27.3758 -0.4291
15.000 0.6787 0.0339 0.0172 1.3344 17.8451 26.2935 -0.3739
17.000 0.6557 0.0343 0.0174 1.2889 16.5107 25.1809 -0.3221
19.000 0.6332 0.0348 0.0177 1.2444 15.2218 24.0388 -0.2730
21.000 0.6112 0.0354 0.0180 1.2008 13.9774 22.8683 -0.2262
23.000 0.5896 0.0363 0.0185 1.1577 12.7766 21.6710 -0.1811
25.000 0.5682 0.0375 0.0191 1.1151 11.6188 20.4494 -0.1372
27.000 0.5469 0.0390 0.0199 1.0724 10.5038 19.2065 -0.0941
29.000 0.5255 0.0412 0.0210 1.0294 9.4314 17.9463 -0.0511
31.000 0.5039 0.0440 0.0225 0.9856 8.4019 16.6739 -0.0078
33.000 0.4817 0.0478 0.0245 0.9404 7.4163 15.3953 0.0366
35.000 0.4587 0.0528 0.0271 0.8932 6.4759 14.1177 0.0828
37.000 0.4345 0.0596 0.0307 0.8430 5.5827 12.8495 0.1318
39.000 0.4086 0.0686 0.0355 0.7892 4.7397 11.6003 0.1849
41.000 0.3806 0.0804 0.0419 0.7305 3.9505 10.3804 0.2436
43.000 0.3500 0.0962 0.0505 0.6663 3.2200 9.2011 0.3096
45.000 0.3163 0.1168 0.0620 0.5956 2.5537 8.0737 0.3854
47.000 0.2793 0.1438 0.0775 0.5185 1.9581 7.0096 0.4739
49.000 0.2392 0.1787 0.0981 0.4356 1.4396 6.0192 0.5786
51.000 0.1964 0.2234 0.1257 0.3490 1.0040 5.1116 0.7044
53.000 0.1525 0.2794 0.1624 0.2625 0.6550 4.2941 0.8574
55.000 0.1099 0.3484 0.2109 0.1815 0.3926 3.5716 1.0458
57.000 0.0716 0.4306 0.2744 0.1124 0.2110 2.9464 1.2810
59.000 0.0408 0.5248 0.3558 0.0602 0.0986 2.4182 1.5789
61.000 0.0194 0.6272 0.4569 0.0266 0.0385 1.9845 1.9620
63.000 0.0072 0.7308 0.5758 0.0092 0.0119 1.6410 2.4615
65.000 0.0019 0.8261 0.7037 0.0023 0.0027 1.3815 3.1204
67.000 + 0.0003 1.0000 2.0000 0.0004 0.0004 1.1936 3.9958

DISCUSSION
Mortality Standard
Development of a standard model mortality schedule for chimpanzees differs
from our previous work on monkey mortality in that the data used are taken from
 Chimpanzee Life Table / 33

a Males

1.o
b
0.8

v)
c
.-0
C 0.4
0
P
2
a
0.2

0.0 I I I I I I
0 10 20 30 40 50 60 70

Age
Fig. 3. a: Fitted standard survivorship schedule for chimpanzee males, superimposed on a schedule computed
directly from colony vital statistics. Dotted lines represent 95% confidence bands. Brass logit method. b Fitted
standard survivorship schedule for chimpanzee females, superimposed on a schedule computed directly from
colony vital statistics. Dotted lines represent 95% confidence bands. Brass logit method.
34 I Dyke et al.
a single species. As noted previously, however, there is considerable variability
between data sets in spite of species identity. This heterogeneity is principally the
result of differences in colony history. Since most colonies tend to be founded with
young animals, the presence of older animals is largely a function of the age of the
colony (these data include animals born from as much as 80 years ago to the
present). Furthermore, the mortality statistics used here have been aggregated
over the entire life of each colony, which means that a proportion of deaths re-
corded for older colonies took place before reductions in mortality brought about by
modern husbandry methods and the introduction of medical technology (including
antibiotics). The somewhat erratic fluctuation from age to age exhibited by all the
distributions shown in Figure 1 is typical of primate colony populations where
numbers of vital events are small, even when they are aggregated over time.
Another difference between the chimpanzee and monkey standard models is
that unlike either the Old World or New World monkeys, chimpanzees exhibit a
sizeable residual component: a, = 0.02496 for males, and a, = 0.016688 for fe-
males. It is not clear what age-independent causes of mortality affect captive
chimpanzees that would not also be present in monkey colonies. It is tempting to
think that chimpanzee mortality patterns resemble those of humans more than
monkeys, and in fact, the a, values are typical of some primitive human groups and
most industrialized nations prior to 1960. However, values for the residual com-
ponent approach zero in other human population isolates and industrial nations
since 1960 [Gage and Dyke, 19861. The residual component is the only one that
shows a statistically significant difference between the sexes (P < 0.001). An a,
value higher for males than for females is atypical of human populations, but we
are reluctant to speculate on an interpretation without more data from other
primates, as well as chimpanzees.
The standard mortality model and its life tables represent an averaging of
many sources of variability, some of which may be quite idiosyncratic, which leads
us to some caution in attempting our goal of understanding chimpanzee mortality
at the most general level. Thus, while the standard represents the data upon which
it is based, some care should be exercised in interpreting it too generally as rep-
resentative of this species. It is of interest, however, to compare this work with that
of Mode [1988] who, using a more elaborate mathematical model applied to rather
fragmentary data, estimated a survivorship schedule for female chimpanzees from
the Gombe reserve. This comparison is shown in Figure 4.
Much of the difference between the curve based on the wild population and
either of the two model schedules can be accounted for by differences in mortality
during the first year of life (only about 67% of wild infants survive the first year,
compared to 80 to 85% in the captive models). In fact, were infant mortality re-
duced by half in the wild population, its survivorship would closely resemble the
model curves well into the adult years, as can be seen from the similarities in
slopes until the mid thirties. Not enough data are available for reliable estimates
of mortality in wild chimpanzees to do more than surmise that captivity might also
have a protective effect for animals older than 40 years. The mortality schedule
estimated by Mode [19881 may be more appropriate for use in modeling studies of
wild chimpanzees than our standard schedules. On the other hand, the standard
schedules may be useful in studies of wild populations if used for mortality in a
manner analogous to the natural fertility schedule [Henry, 19611,which is thought
to represent the maximum reproductive potential of modern non-contracepting
human populations. The consistent sex differences in mortality we find in captive
colonies also suggests that increased mortality of males might be a factor in wild
populations as well.
 Chimpanzee Life Table / 35

1'011 -------
Gombe females

Male Standard

-.-.---Female Standard

0 10 20 30 40 50 60

Age

Fig. 4. Gombe female survivorship schedules vs. male and female standard schedules.

Estimating Mortality in Individual Populations

The extension of schedules beyond ages for which individual population data
are observed is probably the most important use of these models. In cases where
the standard and observed schedules are not too divergent, the Brass logit method
may be appropriate, since mortality data currently available for most chimpanzee
colonies are fragmentary (particularly at the older ages), and thus cannot be used
directly with the Siler model. Figure 3b shows the advantages of this method,
where estimates of proportions surviving are extended quite dramatically beyond
observed values. Expectation of life at birth for females calculated from the ob-
served data is 23.09 years, while that based on the fitted schedule is 44.88 years,
clearly a more realistic estimate for the future of this colony.
Our rule of thumb for determining the adequacy of a fitted model is that (1)
estimated proportions surviving should be distributed within 95% confidence in-
tervals in an unbiased fashion (that is, more or less equally distributed above and
below corresponding observed values), and (2) the fitted survivorship should ap-
proach zero at an age thought to be appropriate for the species in question (both
conditions are met quite adequately in the fit shown in Figure 3). Use of the
standard schedules guarantees that the second requirement is fulfilled, but the
requirement of no bias may be more diffkult to meet. Exactly how to deal with a
poor fit of the standard schedule depends on the extent to which precision is re-
quired. In some cases ad hoc methods such as combining parts of the observed and
fitted schedules may be attempted. Before this approach is taken, however, one
should be certain that the fitted distribution by itself (however approximate) will
not suffice.
As time passes, more data on older animals will become available, which will
improve our knowledge of long-term survival of chimpanzees. At some point it will
36 / Dyke et al.
no doubt be advantageous to reformulate the standard model, but the present
version provides for any colony a plausible mortality schedule extending through
the end of the life span for this species. Such a schedule is detailed enough for
purposes of modeling future colony structure, and for actuarial projections of fu-
ture costs of colony maintenance.
CONCLUSIONS
1. A standard mortality schedule for chimpanzees has been developed from the
vital statistics data of three captive colonies, using the three-component competing
hazard model of Siler [19791.
2. Model mortality may often be substituted for incomplete observed schedules
even when the data are truncated (that is, when mortality statistics are available
for only the younger ages), using the simple regression method of Brass [1971],
provided that the fit to the observed fragment is close and unbiased.
3. Despite some obvious differences in level of mortality, a survival distribu-
tion estimated for wild animals exhibits some similarities in shape to the model
schedules based on captive colonies.
4. As more chimpanzee vital statistics data are collected, the mortality stan-
dard developed here may require modification. Used with care in its present state,
however, it should be adequate for demographic and actuarial projections.
ACKNOWLEDGMENTS
This study was supported by NIH Grants RR 02229 (to B.D.), HD 25346 (to
T.B.G.), and RR 08122 (to S.W.B.). Populations studied are supported by NIH grant
RR 00165 to the Yerkes Regional Primate Research Center; NIH grant RR 03589
and NIH contract 263-88-C-0248 to the University of Texas M.D. Anderson Cancer
Center Science Park; and NIH contract NO-1 HB-27091 and institutional funds at
the Southwest Foundation for Biomedical Research. All animal facilities of the
populations studied are accredited by the American Association of Accreditation of
Laboratory Animal Care. We thank Drs. Anthony Comuzzie and John Ely for their
comments on the manuscript.
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