Zoo Biology 20:169–183 (2001)

Interactions of Target Population Size,

Population Parameters, and Program
Management on Viability of Captive
Populations
Joanne M. Earnhardt,* Steven D. Thompson, and Emily A. Marhevsky
Department of Conservation and Science, Lincoln Park Zoo, Chicago, Illinois

When established conservation programs expand and evolve, management prac-

tices may become inconsistent with program goals. In the past decade, the Ameri-
can Zoo and Aquarium Association expanded species conservation programs by
increasing the number of Species Survival Plans (SSP) and establishing more
than 300 new Population Management Plan (PMP) programs. However, limited
space in captive breeding facilities forces a competition among SSPs and less
intensively managed PMPs. Regional Collection Plans establish priorities and
allocate space accordingly by setting target population size for each species; spe-
cies of high conservation priority (SSPs) are allocated space at the expense of
lower priority species (PMPs). Because population size and genetic composition
interact to impact population viability, target population size is a significant fac-
tor to a population’s prospects for long-term survival. We examined four popula-
tion parameters (current population size, target population size, current gene
diversity, and mean generation time) for 46 mammalian SSPs and 17 PMPs.
Relative to SSPs, PMPs combine smaller current and target population sizes,
lower levels of current gene diversity, and shorter mean generation times than
SSPs. Thus, the average PMP population can expect to lose gene diversity more
rapidly than the average SSP population. PMPs are projected to lose 10% or
more of their founding gene diversity, within only 2 years. In contrast, the aver-
age SSP population is projected to lose 10% in 40 years. Populations with small
current or target population sizes require intensive management to avoid extinc-
tion. More intensive genetic management of populations typically designated as
PMPs, through recruitment of potential founders and equalization of founder
representation, could increase gene diversity and improve viability. Less rigor-
ous population management should be reserved for populations whose long-term
survival is either secure or that can be readily replenished from the wild. Be-
cause PMP populations need intense genetic management similar to that cur-
rently in effect for SSPs, there should be neither a management-level distinction
between programs nor an arbitrary difference in space allocated to programs.
Zoo Biol 20:169–183, 2001. © 2001 Wiley-Liss, Inc.

*Correspondence to: Joanne M. Earnhardt, Department of Conservation and Science, Lincoln Park
Zoo, 2001 North Clark Street, Chicago, IL 60614. E-mail: joanne@lpzoo.org
Received for publication August 12, 2000; Accepted January 18, 2001.

© 2001 Wiley-Liss, Inc.

170 Earnhardt et al.
Key words: Regional Collection Plan (RCP); genetic and demographic management;
extinction; Population Management Plan (PMP); Species Survival Plan (SSP)

INTRODUCTION

Captivity is often seen as an oasis from the inter- and intra-specific competi-
tion that occurs in the wild. However, species do compete in captivity; as in the wild,
this competition occurs because resources and spaces in captive breeding facilities
are limited. The American Zoo and Aquarium Association (AZA) mandates its Taxon
Advisory Groups (TAGs) to develop Regional Collection Plans (RCPs), which iden-
tify goals for each captive-held species. TAGs prioritize all species in a taxon by
using various criteria, which generally include conservation status, educational value,
research needs, and population characteristics (e.g., initial population size, availabil-
ity, and husbandry) [Hutchins et al., 1995, 1998]. Within an RCP, each population is
assigned a specific role or function. These roles—Species Survival Plans© (SSP),
Population Management Plans (PMP), Display/Education/Research Populations, or
phase-out—reflect a decreasing intensity of demographic and genetic management.
Based on the categorizations and on an estimate of the space available for this taxon
within AZA and cooperating institutions, a TAG then establishes a target population
size for each species in the RCP. A consequence of this process is that species win or
lose in the space allocation contest; some species are phased out or omitted from the
plan, whereas others are allocated many or few of the limited spaces. For example,
based on RCP criteria developed by the Prosimian TAG [Porton, 1999], the Coquerel’s
mouse lemur program is given a high priority (SSP), whereas the fat-tailed dwarf
lemur is proposed as a phase-out; thus, space in captivity is allocated to Coquerel’s
mouse lemurs (200 spaces) at the expense of fat-tailed dwarf lemurs (0 spaces).
Competition for space in zoos threatens three primary goals of AZA programs:
conservation of wild populations, no removal of specimens from the wild, and de-
velopment of self-sustaining captive populations [Hutchins et al., 1998; Hutchins, in
press]. Self-sustaining captive populations are important because many captive popu-
lations represent insurance policies against extinction or they act as ambassadors for
conservation of wild populations. Given the potential importance of captive popula-
tions and the limited space available in captivity, AZA’s captive management pro-
grams should make optimal use of the space available to house and preserve as many
species as possible [Conway, 1986; Soulé et al., 1986; Hutchins et al., 1995, 1998].
Populations that have a poor viability prognosis under diverse management scenarios
should be given strong consideration as phase-outs so that the space they occupy can
be used to expand the populations of other species. This article explores how current
genetic and demographic structure and target population sizes imposed by RCPs might
affect the relative viability of AZA’s managed populations. In particular, we evaluate
whether the goal to develop self-sustaining captive populations is consistent with the
relationship between population structure and current management strategies.
Target population sizes impact the management of captive breeding facilities
and AZA programs at two levels. First, population size determines the number of
specimens available for display. If the population is too small, there may not be enough
specimens to meet exhibit demands. Conversely, if the population is too large, other
species may be unnecessarily reduced in number or crowded out altogether. Second,
population size is directly related to population viability—the probable risk of extinc-
 Population Viability for Managed Programs 171

tion within a given period. Smaller populations are more vulnerable to extinction
from loss of genetic diversity and from random demographic events and catastrophes
(e.g., disease) [Goodman, 1987; Shaffer, 1987]. Even though the effect varies from
population to population, inbreeding will depress individual fitness, and thus a
population’s potential for growth, setting up the possibility for an “extinction vortex”
when small inbreeding populations become smaller, more inbred, and increasingly
susceptible to extinction [Gilpin and Soulé, 1986].
Relative to most wild populations, populations in captive breeding facilities are
small (e.g., mammal SSPs average fewer than 200 specimens). Many RCPs have set
target population sizes for PMPs and new SSPs at 100 or fewer. These small targets
seem to be based either on so-called “magic numbers” that are presumed to ensure
wild population viability [Franklin, 1980; Gilpin and Soulé, 1986; Lande and
Barrowclough, 1987], on the presumption that populations of almost any size can be
managed to avoid extinction, or on a practical assessment of available space coupled
with the desire to maximize species richness in the zoo environment.
The level of management and determination of target population sizes varies
between SSPs and PMPs. SSPs were initiated in 1981, and the science of SSP manage-
ment is fairly well established [Ballou and Foose, 1996]. SSP management involves
rigorous demographic and genetic analyses that lead to systematic specimen-by-speci-
men recommendations. Most SSP programs base target population size on demographic
and genetic goals such as maintaining a specified level of gene diversity for a set
period (e.g., 90% or greater of founding gene diversity for 100 years). PMPs were
conceived in the early 1990s as a less rigorous approach to population management.
Demographic and genetic analyses lead to generalized PMP recommendations that
may or may not entail a systematic assessment of every individual in the population.
Many PMP programs lack the rigorous analysis required to project changes in gene
diversity and to scientifically establish target population sizes. Although the goals
associated with any given target population size are often not stated, we assume that
most PMP programs have as an implicit goal the creation of a self-sustaining popula-
tion [Carter, 1999; Lyon and Pate, 1999; Brown, 2001].
For either type of program, managing a population toward viability depends on
factors such as current gene diversity, current population size, effective population
size, population growth, and mean generation time. If the values for these factors
were similar between SSP and PMP populations, the viabilities of PMP populations
would be similar to those of SSP populations. In this study, we evaluate 1) the effect
of four population parameters—current population size, mean generation time, cur-
rent gene diversity , and recommended target population sizes on future loss of gene
diversity and 2) the future loss of gene diversity for PMP populations relative to SSP
populations. In addition, we consider the conflict between the level of management
and the population viability for PMP relative to SSP programs.

METHODS
Population Size Terminology
In the coordinated management of captive populations, size interacts with pro-
grams at several different junctures. To clarify the varied uses of size terminology, we
define population size, effective population size, and target population size. Popula-
tion size (N) is the number of specimens in the defined population at a specified point
172 Earnhardt et al.

in time. Effective population size (Ne) is a genetic metric that represents the size of a
randomly breeding population that would lose genetic diversity (or have the same
rate of inbreeding) as the population under scrutiny. Target population size (Ktarget) is
the number of specimens (spaces) allocated by a TAG through its RCP. When space
is allocated scientifically to develop target population size, the numerical value incor-
porates specific genetic and demographic objectives [Hutchins et al., 1998].
Population Viability
To characterize viability of AZA-managed populations, we establish profiles
for mammalian SSP and PMP populations. Because loss of gene diversity (GD) re-
sulting from genetic drift occurs according to GDt = GD0 × {1 – 1/(2N)t} [Lacy,
1995], we chose to evaluate current GD, mean generation time (T), and current popu-
lation size (N). The values for these parameters and for target population size (Ktarget)
are extracted from SSP and PMP program reports published in AZA’s 1997 Annual
Report on Conservation and Science (ARCS) [Hodskins, 1997]. When one or more
of these parameters are missing from ARCS, we substitute values from either an
approved RCP (e.g., the target population size) or from an electronic studbook data-
base (e.g., population size, GD, mean generation time) maintained in Single Popula-
tion Analysis and Record Keeping Software, SPARKS (version 1.42) [ISIS, 1999].
Mean generation time (T) is determined for the period 1980–1997. Because SPARKS
constructs its life tables by assigning one half of each captive birth each to the male
and female life tables, T is calculated averaging the male and female values for T
from the SPARKS Life Table Report. For GD, data for January 1, 1997 are exported
from SPARKS to GENES (version 11.9) [Lacy, 1999], software that calculates the
GD on a given date. It should be noted that the average profile includes 9 of 46 SSP
and 6 of the 17 PMP populations that are currently less than 90% GD (Tables 1 and
2). Mammalian species are exclusively selected because these taxa have published
RCPs and a larger sample of SSP and PMP programs than other classes. Species
with population sizes of 25 or less are omitted from the process because many of
these populations were new to the management programs and either have not yet
initiated population management or have de-emphasized maintenance of gene diver-
sity. Although its current population size is 34, the Jaguar SSP is excluded from our
analyses because it is a new program that has made specialized genetic decisions.
Based on these criteria, we produce a profile for mammal SSP populations (46 spe-
cies) and PMP populations (17 species) consisting of the average for each parameter
(mean generation time, GD, current population size, and target population size).
We compare the relative viability of SSP and PMP populations using a genetic
measure—the length of time until a population’s GD falls to 90% of that present in
the presumptive wild population from which the founders are derived. This 90%
level is referred to as the viability risk threshold. Maintenance of GD above a 90%
threshold for a given time has become a viability standard in captive breeding pro-
grams [Soulé et al., 1986; Willis and Wiese, 1993]. Gene diversity (expected het-
erozygosity [Lacy, 1995]), is reduced by inbreeding and genetic drift. In particular,
captive populations that lose significant amounts of GD relative to that present in the
wild can expect to experience increasing reproductive problems including lowered
conception rates, higher infant mortality, and lower litter/clutch size [Ralls et al.,
1979; Ralls and Ballou, 1983; Lacy, 1997]. These reproductive problems can reduce
population size even further and decrease population viability (increasing the prob-
 Population Viability for Managed Programs 173
TABLE 1. Species Survival Plan (SSP) species and population parameters
SSP species N Ktarget T % GD
(1) Acinonyx jubatus 286 300 5.9 96.9
(2) Addax nasomaculatus 264 250 7.5 86.48
(3) Aonyx cinerea 113 NA 7 93.7
(4) Bos gaurus 95 95 7.97 88.9
(5) Callimico goeldii 127 200 4.88 94.98
(6) Canis lupus baileyi 101 240 7.05 82
(7) Canis rufus 259 330 6.13 90.4
(8) Ceratotherium simum 120 NA NA NA
(9) Cercocebus torquatus 37 NA NA NA
(10) Cercopithecus neglectus 57 125 NA 82.5
(11) Choeropsis liberiensis 42 70 13.32 96.1
(12) Chrysocyon brachyurus 97 100 5.69 94.2
(13) Colobus angolensis 40 75 NA 94
(14) Colobus guereza 266 250 NA 98
(15) Dendrolagus matschiei 83 75 8.01 91.5
(16) Diceros bicornis michaeli 73 90 15.15 96.8
(17) Diceros bicornis minor 38 80 NA 96
(18) Elephas maximus 155 NA 22.79 97.02
(19) Equus grevyi 227 90 11.30 97.6
(20) Eulemur macaco flavifrons 32 100 NA 87.6
(21) Eulemur macaco macaco 98 100 NA 94.22
(22) Gorilla gorilla gorilla 343 400 17.93 98.8
(23) Helarctos malayanus 56 NA 11.02 90.6
(24) Lemur catta 362 350 9.49 94.7
(25) Loxodonta africana 135 NA 15.73 90.3
(26) Lycaon pictus 34 100 NA 78
(27) Macaca silenus 217 NA 11.95 96.9
(28) Melursus ursinus 46 NA 9.68 92.38
(29) Mirza coquereli 52 200 7.04 89
(30) Mustela nigripes 299 240 2.42 87.93
(31) Nycticebus pygmaeus 58 206 6.99 95.8
(32) Oryx dammah 364 380 7.56 95.9
(33) Oryx leucoryx 228 250 8.17 91.8
(34) Pan paniscus 55 NA 20.85 92.38
(35) Pan troglodytes 247 200 21.84 98.7
(36) Panthera leo krugeri 45 NA 6.23 93.56
(37) Panthera tigris altaica 104 160 7 96.4
(38) Panthera tigris sumatrae 57 100 7.91 90.7
(39) Panthera uncia 250 300 7.42 96.5
(40) Pongo pygmaeus 237 250 20.36 99.3
(41) Pteropus rodricensis 227 NA NA NA
(42) Rhinoceros unicornis 43 NA 19 91.7
(43) Saguinus oedipus 246 200 NA 98
(44) Tremarctos ornatus 80 100 11.11 94.6
(45) Varecia variegata rubra 215 200 NA 89.7
(46) Varecia variegata variegata 223 200 NA 95.2
(1)–(46) Hodskins, 1997; additional data from (1) Marker-Kraus, 1997; (2) Correll, 1994; (4) Morris,
1994; (5) Warneke, 1996; (6) Siminski, 1996; (7) Waddell, 1997; (11) Thompson, 1995; (12) Matern,
1998; (15) Rodden, 1997; (16) Foose, 1998a; (18) Keele, 1997; (19) Fischer, 1998; (22) Wharton,
1996; (23) Frederick, 1998; (24) Villers, 1997; (27) Gledhill, 1996; (28) Wachsberg, 1999; (29) Baker,
1996; (30) Russell, 1997; (32) Rost, 1998; (33) Brown, unk; (35) Fulk, 1997; (36) Pfaff, 1998; (38)
Brady 1998; (39) Varsik, 1997; (42) Foose, 1998b; (44) Rosenthal, 1998.
N, number of animals—current year. Ktarget, target population size. T, mean generation time (years).
% GD, fraction of gene diversity retained. NA, not available.
174 Earnhardt et al.
TABLE 2. Population Mangement Plan (PMP) species and population parameters
PMP species N Ktarget T % GD
(1) Ateles fusciceps robustus 106 106 15.94 97.14
(2) Callithrix geoffroyi 88 100 4.72 94.47
(3) Erythrocebus patas 46 75 9.79 95.58
(4) Gazella dama ruficollis 153 185 4.4 NA
(5) Gazella spekei 33 50 3.54 77.98
(6) Leopardus [Felis] pardalis 76 75 6.59 88.98
(7) Litocranius walleri walleri 86 150 4.78 88.52
(8) Loris tardigradus 47 NA 6.39 90.74
(9) Macaca fuscata fuscata 198 50 9.77 96.92
(10) Macropus parma 62 NA 4 94.34
(11) Madoqua guentheri 73 75 4.49 83.27
(12) Madoqua kirkii—Type A 49 75 3.58 90.37
(13) Panthera pardus orientalis 55 100 NA 81.69
(14) Phascolarctos cinereus adustus 72 NA 7.21 93.35
(15) Pithecia pithecia 114 100 12.85 96.37
(16) Proteles cristatus 32 50 4 88.25
(17) Ursus maritimus 150 150 NA NA
(1)–(17) Hodskins, 1997.
N, number of animals—current year; Ktarget, target population size; T, mean generation time (years); %
GD, fraction of gene diversity retained; NA, not available.

ability of extinction). In this study, we use the term viability in reference to popula-
tion persistence to the 90% risk threshold at a specific future time, and we distin-
guish self-sustaining as population persistence that has no associated time component.
For many SSPs (e.g., black lemur, black rhinoceros, callimico, cheetah, chimpan-
zee), maintenance above the viability risk threshold is a high priority and a stated
goal of their program [Hodskins, 1997].
SSP and PMP populations are compared using independent t-tests with
Bonferroni corrections, for population size, target population size, mean genera-
tion time, and founder genome equivalents (FGE). GD and FGEs are functionally
equivalent measures of gene diversity (GDt/GD0 = 1 – 1/(2 FGE); [Lacy, 1995]).
Because GD is expressed as a proportion, FGE is more appropriate for statistical
analyses; GD for each species was converted to FGE. We use Spearman rank cor-
relations to assess the relationships of parameters within and between SSPs and
PMPs. We expect that target population size should be inversely related to mean
generation time and GD and to be roughly independent of current population size
because a larger target population size would be a logical management strategy to
retard the loss of GD and offset short mean generation time. All analyses are con-
ducted using STATISTICA [StatSoft, 1997, Version 5] with, where appropriate,
pairwise deletion of missing data.
Using CAPACITY [Ballou, 1999, version 4], a program that models the theo-
retical changes in GD over time, we model the average (±1 SD) SSP and PMP popu-
lation projected trajectories for 100 years. For these analyses, the population growth
rate, lambda (λ), is held constant at 1.05 (rapid growth), there is neither immigration
nor emigration, and the ratio of effective population size to population size (Ne/N)
ratio is held constant at 0.30. For many captive populations, the Ne/N ratio is 0.2 –
0.3 [Balmford et al., 1996] (R. Lacy, unpublished observations). To assess the poten-
tial contribution of genetic management to these populations, we examine the effects
 Population Viability for Managed Programs 175

of increasing the ratio of effective population size to population size (Ne/N) from 0.3
to 0.4 to simulate management aimed at minimizing inbreeding and equalizing founder
representation.
To characterize the interaction of the average PMP population parameters with
the future loss of gene diversity, we systematically vary current population size, cur-
rent (initial) GD, mean generation time, and target population size. Using CAPAC-
ITY seeded with the mean of each parameter, and a Ne/N of 0.3, we project the time
to: a) the viability risk threshold (90% GD) and b) a lower risk threshold (85% GD).
For example, current population size is varied in increments of 50, 100, 150, 200,
250, and 300, whereas the other three parameters (current GD, mean generation time,
and target population size) are held constant at their respective mean. Then, mean
generation time is varied in increments of 5, 7, 9, 11, 13, and 15 (while current
population size, current GD, and target population size are held constant). This sys-
tematic variation is repeated for the remaining parameters.

RESULTS
Data are available in ARCS for 46 SSP and 17 PMP programs (Tables 1 and 2).
Although there is great variability among the programs (Table 3), PMP populations
have lower means and narrower ranges for all four parameters than SSP populations.
T-tests indicate that PMPs and SSPs differ significantly in population size (t = 3.394,
df = 57.6, P = 0.006), target population size (t= 4.688, df = 47.9, P = 0.000), and
mean generation time (t= 2.881, df = 38.4, P = 0.032). Thus, SSP populations are
larger, have higher target population sizes, and longer mean generation times.
For SSPs and PMPs, significant correlations exist for 1) current population
size and target population size, 2) current population size and GD, and 3) mean
generation time and GD (Table 4). However, the strength of the correlations among
variables varies between SSPs and PMPs. This pattern may reflect historical differ-
ences in genetic and demographic management between the two programs. For ex-
ample, the current population size and target population size of SSP programs may
be more closely associated because management goals have been reached and/or
because space limitations have resulted in RCPs that have expediently let target popu-
lation size (Ktarget) equal actual population size (N). The weaker correlation between
target population size (Ktarget) and population size (N) for PMPs suggests that, be-

TABLE 3. Comparisons of population parameters for American Zoo and Aquarium Association–
managed populations
SSP PMP
Mean SD n Range Mean SD n Range
a
Population size (N) 148.54 102.14 46 32–364 84.71 46.43 17 32–198a
(specimens)
Target population size 188.1 96.21 34 70–400a 95.78 41.15 14 50–185a
(Ktarget) (specimens)
Gene diversity (% GD) 92.97 4.74 43 78–99.3 90.53 5.86 15 78–97.1
Mean generation time 10.68 5.49 33 2.4–22.8a 6.8 3.73 15 3.5–15.9a
(T) (y)
a
Significantly different at P < 0.05.
SSP, Species Survival Plans; PMP, Population Managment Plans.
176 Earnhardt et al.
TABLE 4. Correlations among population parameters for American Zoo and Aquarium
Association–managed populations
SSP PMP
Target Target
Population population Gene Population population Gene
size size diversity size size diversity
Target population size 0.809a 0.555a
(Ktarget) 34 14
Gene diversity 0.394a 0.17 0.586a 0.249
(% GD) 43 34 15 12
Mean generation –0.114 –0.162 0.364a 0.484 0.33 0.763a
time (T) 33 24 33 15 12 14
a
Significant at P < 0.05.
SSP, Species Survival Plans; PMP, Population Management Plans.

cause most PMPs are relatively young, there has been less opportunity to reach goals
and/or to adjust target population size with respect to available space. The stronger
correlation between mean generation time (T) and GD in PMPs is also consistent
with a lack of population management. If a population is managed, GD should be
weakly associated with mean generation time. Similarly, GD should be less strongly
correlated with the current population size in a managed than an unmanaged popula-
tion: SSPs have a weaker correlation between these parameters than PMPs. The means
for each of the parameters are unlikely to be independent, given the association among
the parameters.
The average PMP population loses gene diversity more rapidly than the aver-
age SSP population (Fig. 1); an average SSP population drops to a 90% viability
risk threshold in a 40-year time frame, whereas the average PMP falls to this thresh-
old in 2 years. In addition, the range (±1 SD) of potential loss is much larger for the
PMP population than the SSP. At 100 years, the PMP range is 28.7–86.1% GD and
the SSP range is 60.7–90.7% GD. This result is expected given the mean values for
the parameters for each program (Table 3). Because SSP programs minimize losses
in GD through active management (Thompson, unpublished observations), these
projections may be overly pessimistic for SSP programs. We increase the Ne/N from
0.3 to 0.4 (theoretically attainable with intense management); this increases the time
to decline to a 90% threshold for SSPs from 40 to 53 years and for PMPs from 2 to
3 years.
Varying parameter values for the average PMP population produces different
patterns for the standard viability risk threshold (90% GD) and the lower level of
85% GD (Fig. 2). For the standard 90% GD level, increases in current population
size and mean generation time have only a slight positive effect, and there is no
effect from an increase in target population size. PMP populations are so constrained
by the low current GD (90.5%, Table 3) that even substantial changes in other pa-
rameters cannot substantially extend the time to the viability risk threshold. The only
change in parameters that produces a substantial improvement is increasing the cur-
rent percent GD. However, when the GD threshold is shifted to 85%, the average
PMP population does not cross this lower threshold until 24 years, in contrast to the
2 years for the standard viability risk threshold. In addition, there is a more substan-
tial time gain with increasing target population sizes (Fig. 2d).
 Population Viability for Managed Programs 177

Fig. 1. Projections for loss of gene diversity (GD) in Species Survival Plan programs (a) and in
Population Management Plan programs (b). The arrows indicate the time at which the projected GD
crosses the 90% viability risk threshold.

DISCUSSION
During the two decades since SSPs were established, AZA’s vision for species
conservation programs has evolved to encompass broader goals. When the concept
for Species Survival Plans was developed in 1981, the primary goal was creation of
 178
Earnhardt et al.
Fig. 2. Comparisons of quantitative variations in population parameters for the average Population Management Plan population: mean generation
time (a), current population size (b), current gene diversity (c), and target population size (d). Open squares indicate the time (years) for the population
simulations to decline to 85% gene diversity; closed diamonds indicate the time for the population simulations to decline to 90% gene diversity, the
viability risk threshold.
 Population Viability for Managed Programs 179

self-sustaining captive populations for endangered, charismatic, high-profile verte-

brates, especially those that could be used for reintroduction programs. Scientific
application of genetic and demographic management based on studbook data would
secure these self-sustaining populations by creating viable population sizes and slow-
ing the loss of GD. Since the initiation of the SSP programs, many SSPs have proved
successful at attaining the proposed larger population sizes. However, other changes
in AZA species conservation programs occurred in these decades as well: 1) TAGs
expanded AZA programs to include strategic multispecies plans (RCPs); 2) more
than 300 PMP programs were created for managed non-SSP species including a broad
range of populations with characteristics that differ from the original SSP popula-
tions; and 3) concerns over space constraints require target population sizes for ev-
ery species program. However, many of the practices dating from the exclusive
management of SSP populations have not been reevaluated to conform to the expan-
sion and evolution of AZA programs. This deficiency can be seen in our compari-
sons of SSP and PMP populations, in the inconsistencies of target population sizes,
and in the application of levels of genetic management.
Comparison of SSP and PMP Populations
Genetic and demographic management of populations influences the popula-
tion parameters: population size, mean generation time, and current GD. PMPs, a
relatively recent addition to cooperative programs, generally lack long-term goals
and management; their smaller population size, shorter mean generation time and
lower GD, and the correlations among these parameters reflect that lack of manage-
ment. For example, population growth is frequently recommended when manage-
ment is initiated; thus, the longer SSP management period has had more opportunity
for population growth. In addition, because most PMP populations have been in zoos
for many years, many generations have elapsed without the benefit of genetic man-
agement. Because gene diversity is lost with each generation, by the time the proto-
PMP population becomes a PMP, there is much less genetic variation compared to
SSP populations.
If the type of species designated as a PMP program differs from those in SSP
programs, the demographic and genetic population parameters may differ as well.
SSP species are usually charismatic megavertebrates that have higher conservation
priorities and are popular with visitors (and thus in high demand for display). These
larger, charismatic species such as elephants, great apes, and large cats have long
mean generation times that help stem the rate at which gene diversity is lost. Con-
versely, PMP species are more likely to be smaller-bodied individuals with shorter
mean generation times that predispose them to lose GD at higher rates than SSP
species. In addition, because PMP species are less charismatic than SSP species, the
incentive to provide space for individuals and resources for importation is less pow-
erful. A commitment to SSP species occurs despite the higher costs associated with
larger vertebrates [Balmford et al., 1996]. Thus, the perceived value of PMP popula-
tions may also contribute to their lower viability relative to SSP populations.
Setting Target Population Size
TAGs have typically assigned larger target population sizes to SSP programs
and smaller target population sizes to PMP programs (Table 3). An SSP generally
has explicit long-term goals of maintaining gene diversity for a given time frame;
180 Earnhardt et al.

thus, target population sizes are frequently scientifically determined for these pro-
grams. On the other hand, PMP populations rarely have the benefit of either long-
term goals or scientifically determined target population sizes. Thus, no scientific
basis exists for setting the target population size of a PMP lower than that of an SSP.
Recognizing that the past loss of GD for PMP populations places many popu-
lations perilously close to or below the 90% GD threshold, and because 90% GD is a
benchmark, not an absolute value that is comparable across species, we lower the
threshold to 85% GD and consider parameters in relation to this new threshold. Us-
ing the standard 90% level, increases in target population size do not increase time
to the viability risk threshold. Increases in GD (that could potentially be attained
through intense genetic management) are the optimal strategy. Using the lower level
of 85% GD, both increases in GD and target population size substantially increase
the time to the new threshold and benefit the long-term prospects of the population.
Unfortunately, tolerable levels of GD for a program cannot be predetermined, and
any reduction only increases the risk of extinction.
When developing an RCP, TAGs strive to balance species richness—the num-
ber of species to be managed under the plan—against available space. Because small
target population sizes permit a greater number of cooperative management programs
within the limited space of captive breeding facilities, there is a temptation to keep
population sizes low. The nature of this trade-off between population size, available
space, viability, and species richness is not complex. For example, if 600 potential
spaces are allocated equally as target population sizes for 12 average SSP programs,
each program could expect a 90% risk threshold after 12 years (Table 5). In contrast,
three SSP programs with target population sizes of 200 would not reach a 90% risk
threshold until 43 years. With smaller target population size comes the penalty of
lower viability. Spreading this elevated risk among many species, rather than con-
centrating on few species with lower risk per species, sacrifices long-term conserva-
tion and education goals for short-term gains in variety on display. An evaluation of
space that simultaneously considers the needs of multiple species could lead to more
optimal use of space while setting realistic target population sizes.

Demographic and Genetic Management of Stochasticity

All populations in AZA cooperative programs are small (<500) [Franklin, 1980].
Small populations are, in general, less viable than large populations because
stochasticity at either the demographic or genetic level can drive small populations
to extinction. Population managers of cooperative programs reduce stochastic threats
by altering, either singly or in combination, the factors contributing to these risks.
For example, the common remedy for demographic stochasticity is an increase in

TABLE 5. Allocation of 600 spaces among varying numbers of programs

Spaces per species No. of species Years to 90% GDa
50 12 12
100 6 24
150 4 34
200 3 43
a
Based on the average Species Survival Plan profile.
GD, gene diversity.
 Population Viability for Managed Programs 181

size; therefore, target population sizes should be as large as possible to minimize the
effects of random, elevated numbers of deaths or extremes of sex ratio bias. In-
creases in size benefit populations at the genetic level as well by slowing genetic
drift and reducing the probability of inbreeding. Unfortunately, populations in cap-
tive breeding programs are small because space is limited; they cannot completely
escape the common threats to small populations.
Rigorous genetic management may provide the greatest potential for long-term
viability of PMP populations. Because genetic management of pedigreed populations
can minimize inbreeding and loss of gene diversity, there is a need for genetic man-
agement of small populations and the smaller the population, the greater the need for
intensive management. In many unmanaged captive populations such as PMPs, in-
equalities in reproductive success for individuals have led to great disparity between
over- and underrepresented individuals. Genetic management can move to equalize
the representation, which translates into a gain in gene diversity. Many unmanaged
populations also have potential founders that have not produced offspring for
nonbiological reasons (e.g., inappropriate exhibits or social groupings, opportunity to
mate). Identification of these individuals and actions to remedy the situation could
provide new genetic material for populations. The equalization of representation and
the conversion of potential founders to founders would require detailed population
analysis, specimen-by-specimen recommendations, and full participation by holding
institutions—actions typical of SSP management. Yet the cost of genetic management
may be inexpensive in relation to the gain in viability of populations. The efficiency
of genetic management is especially high if it is compared to the hefty costs of alter-
natives such as increases in importations, assisted reproductive technology, increases
in the number and/or size of exhibits, or increases in off-exhibit holding spaces.
In summary, the existing practices of linking differing levels of genetic man-
agement and differing levels of target population sizes with program classifications
(e.g., SSP vs. PMP) have no scientific basis. These practices probably result from
the combination of rapid AZA program expansion and the failure to reassess pro-
gram criteria and goals. Unfortunately, these practices create the potential for failure
as managers strive to develop viable populations.

CONCLUSIONS
1. The average values for mean generation time, current population size, and
current gene diversity are lower for mammal PMPs than SSPs. This difference im-
plies that PMP populations will lose genetic diversity faster than SSP populations.
2. Target population size is not scientifically allocated to cooperative programs.
Regional Collection Plans routinely set target population sizes for PMP programs
that are smaller than SSP program targets. Small target population sizes allow more
AZA cooperative programs, but they decrease the probable viability of each pro-
gram. As RCPs are developed for other classes (birds, reptiles and amphibians), TAGS
could benefit from the example of mammal RCPs.
3. Given the limited space and resources for captive breeding programs, com-
petition among programs is inevitable. Conflicts in goals for target population sizes
may best be resolved by consistently integrating three factors into determination of
target population sizes: RCP goals (e.g., optimization of space), members’ exhibit
needs, and genetic and demographic goals (i.e., viability) for each species.
182 Earnhardt et al.

4. The smaller a population, the greater the intensity of genetic management

needed to meet viability goals. By unintentionally coupling genetic management
with program level designation (SSP or PMP), many AZA programs have inad-
vertently reversed the requisite intensity of management. PMP populations, which
are often smaller than SSP populations, would benefit from more intensive genetic
management.

ACKNOWLEDGMENTS
We thank those studbook keepers who provided us with copies of their elec-
tronic databases. Without their diligence in maintaining those databases, neither AZA’s
population management efforts nor this type of comparative study would be pos-
sible. Our thanks also to those TAGs that provided us with final or draft RCPs.
Earlier versions of the manuscript were much improved through comments by Lisa
Faust, Sadie Ryan, and three anonymous reviewers.

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