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The Sizes of Elephant Groups in Zoos: Implications for Elephant Welfare


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This study examined the distribution of 495 Asian elephants (Elephas maximus) and 336 African elephants (Loxodonta africana) in 194 zoos, most of which were located in Europe (49.1%) and North America (32.6%). Cows outnumbered bulls 4 to 1 (Loxodonta) and 3 to 1 (Elephas). Groups contained 7 or fewer: mean, 4.28 (sigma = 5.73). One fifth of elephants lived alone or with one conspecific. Forty-six elephants (5.5%) had no conspecific. Many zoos ignore minimum group sizes of regional zoo association guidelines. The American Zoo and Aquarium Association recommends that breeding facilities keep herds of 6 to 12 elephants. The British and Irish Association of Zoos and Aquariums recommends keeping together at least 4 cows over 2 years old. Over 69% Asian and 80% African cow groups-including those under 2 years-consisted of fewer than 4 individuals. Recently, Europe and North America have made progress with some zoos no longer keeping elephants and with others investing in improved facilities and forming larger herds. The welfare of individual elephants should outweigh all other considerations; zoos should urgently seek to integrate small groups into larger herds.
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The Sizes of Elephant Groups in Zoos: Implications for Elephant Welfare
Paul A. Rees a
a School of Environment & Life Sciences, University of Salford, United Kingdom
Online Publication Date: 01 January 2009
To cite this Article Rees, Paul A.(2009)'The Sizes of Elephant Groups in Zoos: Implications for Elephant Welfare',Journal of Applied
Animal Welfare Science,12:1,44 — 60
To link to this Article: DOI: 10.1080/10888700802536699
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DOI: 10.1080/10888700802536699
The Sizes of Elephant Groups in Zoos:
Implications for Elephant Welfare
Paul A. Rees
School of Environment & Life Sciences,
University of Salford, United Kingdom
This study examined the distribution of 495 Asian elephants (Elephas maximus)
and 336 African elephants (Loxodonta africana) in 194 zoos, most of which were
located in Europe (49.1%) and North America (32.6%). Cows outnumbered bulls
4 to 1 (Loxodonta) and 3 to 1 (Elephas). Groups contained 7 or fewer: mean,
4.28 (D5.73). One fifth of elephants lived alone or with one conspecific.
Forty-six elephants (5.5%) had no conspecific. Many zoos ignore minimum group
sizes of regional zoo association guidelines. The American Zoo and Aquarium
Association recommends that breeding facilities keep herds of 6 to 12 elephants.
The British and Irish Association of Zoos and Aquariums recommends keeping
together at least 4 cows over 2 years old. Over 69% Asian and 80% African cow
groups—including those under 2 years—consisted of fewer than 4 individuals.
Recently, Europe and North America have made progress with some zoos no
longer keeping elephants and with others investing in improved facilities and
forming larger herds. The welfare of individual elephants should outweigh all
other considerations; zoos should urgently seek to integrate small groups into
larger herds.
Previous studies of elephants in the zoo populations have been concerned
with population management and the assessment of their possible future
contribution to conservation (Rees, 2003a; Wiese, 2000; Wiese & Willis,
2006). In spite of recent developments in artificial insemination (Hodgkins,
Correspondence should be sent to Paul A. Rees, School of Environment & Life Sciences and
Research Institute for the Built and Human Environment, Peel Building, University of Salford,
Salford, Greater Manchester, UK, M5 4WT. Email:
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2000; Schmitt, 1998), improved neonatal survival rates, and the demonstration
that the life span and longevity of zoo elephants are comparable to those
of elephants in the wild (Wiese & Willis, 2004), studies have generally
concluded that zoo elephant populations are not self-sustaining, except perhaps
the African (Loxodonta africana) population in North America (Olson & Wiese,
The survival of Asian elephants (Elephas maximus) in zoos will probably
depend upon the importation of additional individuals from range states, but there
has been considerable opposition to this (Hedges, Tyson, Sitompul, & Hammatt,
2006). Although the demographic characteristics of zoo elephant populations
have been studied in detail, there has been little consideration of the welfare
implications of keeping small social groups in captivity.
Almost 25 years ago, Eltringham (1984) suggested that zoos—to achieve
self-sustaining populations of elephants—would have to establish large breeding
groups and that this would require a specialized approach more like ranching
than zookeeping. Such groups would have a higher conservation value than
small groups and the potential to improve the welfare of elephants in the zoo.
Unfortunately, the zoo community has been slow to respond.
The complex nature of the family relationships that exist within elephant
populations has been well known from anecdotal evidence for 50 years (Car-
rington, 1958; Williams, 1951) and from the work of field scientists for more
than 30 years (Douglas-Hamilton & Douglas-Hamilton, 1978; Laws, Parker,
& Johnstone, 1975; McKay, 1973). Elephants live in family groups, led by a
matriarch and comprised of related adult cows and their offspring. These groups
are visited by adult bulls who spend much of their time alone or in bachelor
groups, except when in musth (Sukumar, 2003).
In spite of this well-established knowledge, many zoos have traditionally kept
elephants in solitary conditions or in small groups. There has been considerable
recent concern about the conditions in which elephants are kept in zoos (Clubb &
Mason, 2002). Although it has been argued that some welfare compromises are
essential if captive breeding programs are to succeed (Rees, 2003b), nevertheless,
a number of zoos have stopped keeping elephants and transferred their animals
to larger groups, notably in North America and Europe.
This study examines the social structure of the global zoo populations of
African and Asian elephants and considers the extent to which elephants are
currently being held alone and in inappropriate social groups.
Data on the number of elephants kept by zoos were taken from the database held
by the International Species Information System (ISIS). This database contains
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details of the animals kept by almost 650 member institutions from more than
70 countries. Most of the elephant records held by ISIS relate to traditional
zoological gardens and safari parks. The terms “zoo” and “ISIS zoo” are used
here to refer to any institution listed by ISIS.
Although there are other sources of information on captive elephants, such
as regional studbooks, they do not cover all countries. The ISIS database is a
“live” database; although it does not contain data on all zoos, it was considered
a reliable record of major zoo holdings at a fixed point in time.
This study analyzes data on the holdings of African (Loxodonta africana)
and Asian (Elephas maximus) elephants at 194 zoos on October 27, 2006
(Anonymous, 2006a, 2006b). The ISIS records for African elephants recog-
nized four subspecies: Loxodonta africana africana, L. a. knochenhaueri, L.
a. oxyotis, and L. a. cyclotis. This last group has recently been reclassified
as the separate species L. cyclotis and was represented by a single individ-
ual. Records for Asian elephants recognized four subspecies: Elephas maximus
maximus, E. m. indicus, E. m. hirsutus, and E. m. sumatranus. Data for all
subspecies have been concatenated so that they represent the two species L.
africana and E. maximus. Data collected from the ISIS database in 1999 for
Elephas (Rees, 2001a) have been compared with the October 2006 data for this
The ISIS database recorded individuals as male, female, and unknown (in
relation to sex). The ages of individuals were not recorded. Some of the following
analysis assumes that each zoo held its elephants as a single group, including
those that kept both species.
The Global Distribution of Zoos Holding Elephants
At the end of October 2006, a total of 194 zoos recorded by ISIS held 831
elephants (including 3 of unspecified sex). A total of 104 zoos held African
elephants, and 114 zoos held Asian elephants. Most zoos that held elephants
either kept the Asian species only (90 zoos) or the African species only (80 zoos).
Twenty-four zoos (12.4%) kept both species.
Most of the zoos holding elephants were located in North America or Europe.
A higher proportion of the zoos holding African elephants were located in North
America (51.0%) than in Europe (40.4%). This situation was reversed for zoos
holding Asian elephants, with 47.4% of zoos in Europe and 38.6% in North
America. The remaining zoos were distributed across Central America, South
America, Australasia, and Southeast Asia (particularly important for the Asian
species) with only Loxodonta in the listed African zoos (Table 1).
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The Global Distribution of ISIS Zoos Holding Elephants
(Anonymous, 2006a, 2006b)
Loxodonta africana Elephas maximus
Region Zoos %Zoos %
Europe 42 40.4 54 47.4
North America 53 51.0 44 38.6
Africa 2 1.9 0 0.0
Central America 2 1.9 2 1.8
South America 2 1.9 1 0.9
Southeast Asia 2 1.9 8 7.0
Australia 1 1.0 5 4.4
Total 104 100 114 100
The Global Distribution of Zoo Elephants
Most zoo elephants (81.7%) were located in Europe (49.1%) and North America
(32.6%). European and North American zoos held similar numbers of African
elephants (167 and 147, respectively, representing 93.5% of the total for this
species), but European zoos alone held 48.7% of all Asian elephants. Most of
the remaining Asian elephants (Table 2, Figure 1) were held by zoos in North
America (25.1%) and Southeast Asia (21.6%).
The Global Distribution of Elephants in ISIS Zoos (Anonymous, 2006a, 2006b)
Loxodonta africana Elephas maximus
Region Bulls Cows Sex Ratio Bulls Cows Sex Ratio
Europe 40 126 1:3.15 51 188 1:3.69
North America 22 125 1:5.68 23 101 1:4.39
Africa 2 4 1:2.00 0 0
Central America 1 3 1:3.00 0 4 0:4.00
South America 1 1 1:1.00 1 0 1:0
Southeast Asia 2 5 1:2.50 40 67 1:1.68
Australasia 0 3 0:3.00 4 14 1:3.50
Totals 68 267 1:3.93 119 374 1:3.14
aOne Loxodonta specimen whose sex was not recorded omitted. bTwo Elephas specimens whose
sex was not recorded omitted.
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FIGURE 1 Geographical distribution of individual zoo elephants.
Colombo Zoo, Sri Lanka, held 1 African and 69 Asian elephants. Most of
these animals were kept at the Pinnewela Elephant Orphanage (some distance
from the zoo itself), where they were able to associate in a large group. This
group of Asian elephants has been excluded from some of the following analysis
because it represents almost 14% of the total number of this species in the study
and therefore distorts the overall distribution.
Sex Ratio and Group Composition
Cow zoo elephants outnumbered bulls by almost four to one in Loxondata and
approximately three to one in Elephas. The sex ratio varied considerably on a
regional basis, but almost all regions possessed more cows than bulls (Table 2).
Five (4.4%) of the 114 zoos holding Elephas kept only bulls, 60 (52.6%) kept
only cows, and 49 (43.0%) kept both sexes. One zoo kept a single specimen
of Elephas of unrecorded sex. Overall, 54 zoos (47.4%) kept bulls; 109 zoos
(95.6%) kept cows.
Forty-eight (46.2%) of the 104 zoos holding Loxodonta kept bulls and 102
(98.1%) kept cows. Two zoos kept only bulls (1.9%), 56 (53.8%) kept only
cows, and 46 (44.2%) kept both sexes.
The mean number of bull African elephants held by each zoo (ND104)
holding this species was 0.65, (D0.90), and the mean number of cows was
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2.57 (D1.76). Zoos that kept Asian elephants (ND114) held a mean of 1.04
bulls (D2.85) and 3.28 cows (D4.14).
The modal number of elephants kept, for both Elephas and Loxodonta, was
zero bulls and two cows. If zoos that do not keep bulls are excluded, the modal
number of bulls was one for both species.
The Distribution of Group Sizes
Most group sizes were small, between 1 and 7 (Figure 2); the mean group size
was 4.28 (D5.73), including mixed-species groups, and ranged from 1 to
70 individuals (Colombo Zoo). There was no statistically significant difference
between the group sizes of Asian (xD4.34, D6.83) and African ( xD3.23,
D2.41) elephants (tD1.630, df D216, two-tailed, p>.10). More than
69% of Asian elephant cow groups and more than 80% of African elephant cow
groups consisted of fewer than 4 individuals (Figure 3).
The Distribution of Elephants Between Groups
Almost half of Asian elephants (47.9%) and 62.8% of African elephants were
held in groups of five or fewer individuals of the same species. When herds of
mixed species are considered, 52.6 % of all elephants were kept in groups of
five or fewer individuals of either the same species or in a mixed-species herd
(Figure 4).
FIGURE 2 Distribution of elephant group sizes. Note: The column labeled “both species”
indicates the frequency of group sizes when multi-species groups are included.
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FIGURE 3 Percentage cumulative frequency distribution of elephant group sizes. Elephas
and Loxodonta lines include only zoos that held cows of these species. The line labeled
“Both” includes all groups (single and multi-species) and both sexes.
The Global Distribution of Solitary Zoo Elephants
Twenty zoos held a single Asian elephant (4 bulls, 16 cows); 26 zoos held a
single African elephant (2 bulls, 24 cows), some of whom had companions of
a different elephant species. When these companions are taken into account,
only 22 zoos (11.3%) kept lone elephants: 1 bull and 12 cows of Loxodonta
and 2 bulls and 7 cows of Elephas. Nineteen (86.4%) of these solitary elephants
were located in Europe and North America.
Almost one fifth of all elephants were kept alone or with a single companion
of the same species. Forty-six elephants (5.5%) were being kept without any
companion of the same species and 112 (13.5%) with a single conspecific
companion. When companions of either species are taken into account, the
number of elephants considered to be kept alone approximately halved to 22
individuals; 114 were kept with a single companion of either species.
Changes in the Zoo Asian Elephant Population Between
1999 and 2006
The number of ISIS zoos holding Asian elephants fell from 135 in 1999 to 114
in 2006, a decrease of 15.6%. There was no statistically significant change in
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FIGURE 4 Percentage cumulative frequency distribution of elephants within different
group sizes. All lines exclude Colombo Zoo.
the mean group size between 1999 ( xD3.56, D2.95) and 2006 ( xD4.34,
D6.83) (tD1.13, df D247, two-tailed, p>.10). There was little change in
the distribution of group sizes between 1999 and 2006 (Figure 5). However, the
cumulative frequency curve for 2006 (Figure 6) is located slightly to the right
of that for 1999 and suggests a trend toward increased group sizes. There was
a decrease in the number of zoos holding just 1 or 2 elephants (Figure 7). In
1999, the three largest herds each consisted of 15 animals. By 2006, the largest
herd contained 69 animals (Colombo Zoo); however, only one of the herds of
15 remained.
Between 1999 and 2006, the total number of Asian elephants increased by
about 3% (from 481 to 495). However, the proportion of bulls increased from
18.1% in 1999 to 24.0% in 2006 (from 87 to 119 individuals).
This study examined the distribution of 495 Asian elephants and 336 African
elephants living in 194 zoos in October 2006. Most of these animals were
in zoos in Europe (49.1%) and North America (32.6%). Cow zoo elephants
outnumbered bulls by almost four to one in Loxodonta and approximately three
to one in Elephas. Most group sizes were small (between one and seven) with a
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FIGURE 5 Percentage cumulative frequency distribution of Asian elephant group sizes
(1999 and 2006). Colombo Zoo excluded from 2006 data.
FIGURE 6 Percentage cumulative frequency distribution of Asian elephants within dif-
ferent group sizes. Line X indicates the approximate position the curve might occupy if a
minimum group size of five is achieved.
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FIGURE 7 Change in Asian elephant group size distribution, 1999–2006. Excludes
Colombo Zoo.
mean of 4.28 (D5.73), including mixed-species groups; 19% of all elephants
were kept alone or with a single companion of the same species. Forty-six
elephants (5.5%) were being kept without any companion of the same species.
Modern zoos attempt to enrich the environments of their elephants with a vari-
ety of techniques and devices, many of which involve encouraging exploratory or
feeding behavior (Gilbert, 1994; Green, 1993; Haight, 1993). However, interac-
tion with other elephants provides the single most significant form of enrichment
to the lives of those animals kept in appropriately structured social groups (Rees,
In the wild, elephant social structure is complex. Laws et al. (1975) suggested
that the largest stable population unit (or the smallest discrete unit) averaged 5–
6 elephants in Murchison Falls Park, South in Uganda. The mean group size
recorded was 11.6 (range 2–29). Although the basic family unit has an average
of 3 members consisting of a cow and her 1–2 offspring (Spinage, 1994), huge
megaherds of several hundred and up to 1,000 individuals have been observed
in Africa (Laws et al., 1975).
Asian elephants live in family groups that typically contain one adult cow and
between one and five immature offspring (Sukumar, 1994). Larger groups may
contain two or more adult cows. Groups sometimes contain three generations.
A group of elephants with more than one adult cow has been called an
“extended family unit” (Douglas-Hamilton & Douglas-Hamilton, 1978) or a
“joint family” (Sukumar, 1994). Larger groups of related animals have been
called “kin groups” (Douglas-Hamilton & Douglas-Hamilton, 1978) or “bond
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groups” (Moss, 1988), the latter term not necessarily implying relatedness. Moss
and Poole (1983) have described a multitiered network of relationships within
the elephants in Amboseli—the population being divided into subpopulations
that are further divided into clans, then bond groups, family units, and mother-
offspring units. The complexity and flexibility of elephant social organization in
the wild have been discussed at length by Sukumar (2003).
For the purposes of comparing the size of elephant groups in European zoos
with those found in the wild, Clubb and Mason (2002) suggested that cow
elephants form stable groups of between 6 and 8 animals in the Asian species, on
average, and between 4 and 12 in the African species. However, Hutchins (2006)
has argued against using “nature” as a yardstick for measuring the adequacy of
zoo management because elephant society is highly variable with regard to group
size and composition and adapts to prevailing environmental conditions, such
as food supply. He suggests that—if elephants are exhibiting a broad range of
natural behaviors and not showing high levels of aggression, agitation, lethargy,
or stereotypic movements—it might be reasonable to conclude that their social
needs were being met, regardless of group size. Nevertheless, he concedes that
it is unacceptable to keep adult cows alone or in pairs.
Approximately 48% of all elephants in zoos are currently kept in social
groups of 5 or fewer animals (regardless of species composition). The larger we
assume the normal group size of wild elephant families to be, the greater the
number of zoos that will fail to achieve this. Almost 60% of zoos fail to achieve
a group size of 4 animals (of single or mixed species); this rises to about 91%
if the group size is 8 and to more than 95% if it is 12.
The European Endangered Species Programme (EEP), Species Survival Plan
(SSP) in North America, and Australian Species Management Program (ASMP)
recommend keeping elephants—as far as possible—in matriarchal intact family
groups (Conservation Breeding Specialist Group, 2004; European Association of
Zoos and Aquaria, 2004; Lees, 2004). In 2001, the American Zoo and Aquarium
Association (AZA) recommended that institutions should hold at least three cows
(AZA, 2001). At a meeting held in January 2005, the Elephant Directors of AZA
institutions agreed that breeding facilities should strive to hold 6 to 12 elephants,
with capabilities for 2 or more bulls, and that holding facilities should strive to
maintain 2 to 6 elephants, with capabilities for all bulls (Reed, 2005).
Clubb and Mason (2002) found that 61.1% of European zoos holding Asian
elephants kept three individuals or fewer (in 1999) and that 59.5% of zoos
holding African elephants housed four elephants or fewer (in 2001). In 2006,
64.0% of ISIS zoos held three or fewer Asian elephants and 80.8% of zoos held
four or fewer African elephants.
The British and Irish Association of Zoos and Aquariums recommends that
cows should be kept in groups of at least four animals more than 2 years old
(Stevenson & Walter, 2002). This analysis has shown that over 69% of Asian
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elephant cow groups and 80% of African elephant cow groups (including animals
under 2 years) consisted of fewer than four individuals (Figure 3).
Zoos’ elephant groups often consist of unrelated individuals. This is because
they have originated from a number of different sources, including logging
camps, other zoos, circuses, and the wild. Because of the risk of inbreeding,
zoos do not generally keep several generations together; in any event, relatively
few herds contain second-generation offspring bred in captivity.
In a zoo environment, the keeping of related animals together is a lower
priority than moving them between institutions for breeding purposes. However,
the formation of groupings of unrelated individuals is preferable to the keeping
of elephants alone or in very small groups. Even unrelated individuals may
form strong bonds. Friendships are important to elephants, and there is evidence
that they form special relationships with particular individuals in zoos (Garai,
1992). Allomothering is well known in elephants in the wild (Lee, 1987) and
is generally considered to involve close relatives of the mother. However, it is
also important in zoo elephants (Rapaport & Haight, 1987) where both related
and unrelated adult cows may act as allomothers (Rees, 2001a).
Contact with members of their own species is important if elephants are
to develop normal behaviors. There is some evidence that the development of
normal sexual behavior in juvenile bull elephants may depend upon exposure
to reproductively active adults (Rees, 2004); when given the opportunity, very
young calves show considerable interest in the sexual activity of adults (Rees,
2003c). This is not surprising in animals who are typical K-strategists and as
such make a large investment in each of their offspring (Southwood, 1981).
Zoo elephant social groups inevitably change due to births and deaths. How-
ever, in recent years they have also been changing because some zoos have
decided to stop keeping elephants, either following the death of their last indi-
vidual or because they have taken the decision, on welfare grounds, to move their
lone animals or small groups to join elephants in other zoos or in sanctuaries.
Such transfers may cause stress to the animals who are moved and to the
members of the receiving group; however, there is some evidence that this stress
is neither prolonged nor severe (Schmid, Heistermann, Ganslosser, & Hodges,
2001). Sometimes elephants are moved as a result of conflict within a group
(Rees, 2001a), and it has been suggested that group compatibility might be more
important to elephant welfare than group size (Hutchins, 2006). Veasey (2006)
has argued that single young cows should not be moved unless incompatibilities
occur and that new herds should be created from a nucleus of trios or more of
compatible, related elephants.
In the United Kingdom, London Zoo moved its three Asian elephants to
Whipsnade Park in 2001; Dudley Zoo and Longleat Safari Park sent their African
elephants to zoos in France in 2003. Zoos in Bristol, Edinburgh, Cricket St.
Thomas, and the Welsh Mountain Zoo no longer keep elephants. In the United
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States, Detroit Zoo, Frank Buck Zoo (Texas), Mesker Park Zoo (Indiana), Henry
Villas Zoo (Wisconsin), Chehaw Wild Animal Park (Georgia), Alaska Zoo, and
the Louisiana Purchase Gardens and Zoo all sent their animals to elephant
sanctuaries between 1998 and 2007. Sacramento Zoo and San Francisco Zoo no
longer keep elephants; others, like the Smithsonian’s National Zoological Park,
will not replace their remaining elephants. Some of these elephant movements
were the result of public pressure; in at least one case (London) elephants were
moved following the death of a keeper.
Since the listing of Asian elephants and most populations of African ele-
phants in Appendix I of the Convention on International Trade in Endangered
Species of Wild Fauna and Flora, 1973 (Convention on International Trade in
Endangered Species of Wild Fauna and Flora), it has become extremely difficult
for zoos to obtain elephants from their range states. There is opposition among
conservationists to the importation of elephants into nonrange states because the
conservation benefits of such movements are unproven (Hedges et al., 2006).
However, importation is still considered essential in the short term to increase
the reproductive potential of animals in the EEP, SSP, and the ASMP (Hutchins
& Keele, 2006; Wiese & Willis, 2006).
Some zoos have invested in new elephant facilities: Chester (England); Köln
(Germany); and, in the United States, the North Carolina and Pittsburgh (Penn-
sylvania) Zoos. These zoos intend to increase the size of their herds (A. Bloom-
smith, personal communication, April 20, 2007; Rees, 2001b). The facility at
Köln is particularly impressive—with an area of 20,000 m2—and is designed to
hold up to 15 cows and their offspring plus several bulls (Nogge, 2004). In the
United States, a number of AZA zoos are planning to upgrade from holding to
breeding facilities, creating 8 additional breeding facilities for Loxodonta and 12
for Elephas (Keele, 2006). These new facilities will undoubtedly deliver welfare
benefits to the individual elephants within the enlarged groups and increase
their potential to contribute to captive breeding programs. However, the scarcity
of suitable animals within the existing worldwide zoo populations will hinder
future attempts to form larger elephant groups, and there are significant legal,
political, and other barriers to further importation of animals from their range
states (Hedges et al., 2006; Hutchins & Keele, 2006).
In spite of the current attempts by some zoos—notably in the United States
and Europe—to create larger elephant herds, as yet there is little evidence
that zoos intend to establish very large elephant herds managed according to
the ranching methods envisaged by Eltringham (1984). However, it has been
proposed that regional elephant reserves should be developed in North America
particularly to increase zoos’ capacities to hold bulls (Keele, 2006).
Very large groups may not be essential for the social welfare needs of
elephants, but it remains to be seen if the planned breeding groups of up to
12 animals in AZA zoos will increase reproductive success sufficiently to pro-
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duce self-sustaining populations such as those modeled by Faust, Thompson, and
Earnhardt (2006) for Elephas. Unless reproductive rates increase significantly—
which will require significant changes to promote the sociosexual competence
of captive elephants—or more animals are imported, zoo elephant populations
face global extinction within a few decades (Olson & Wiese, 2000; Rees, 2003a;
Wiese, 2000).
A 2005 survey of 78 AZA-accredited zoos with elephants found that 40 zoos
planned to expand or build new elephant exhibits (AZA, 2007). However, this
transition will be difficult, and it will take time to raise the necessary funds.
There are few healthy, reproductively viable elephants available to grow captive
herds; in addition, some individuals are incompatible, often due to a history
of inadequate care in circuses or substandard zoos. These difficulties are com-
pounded by the poor reproductive performance of zoo elephants. This may be
due to the following:
1. Reproductive pathologies in cows (caused by delaying reproduction);
2. Lack of maternal competence;
3. Reproductive suppression;
4. Overnutrition and lack of exercise causing rapid fetal growth and dystocia;
5. A historical tendency to separate bulls from cows to prevent unwanted
male offspring (Hutchins, 2006; Hutchins & Keele, 2006).
Zoos that intend to continue keeping elephants need to be clear about their
conservation function. This is far from obvious. Some authorities still consider
zoos’ elephants a “reservoir for threatened in-situ populations” (Faust et al.,
2006); however, it is not at all clear that this can be justified in the context
of the current decline in zoo populations and their small size compared with
the relatively large number of elephants still remaining in the wild. Hutchins
and Keele have claimed that ex-situ elephant programs can assist field conserva-
tion through public education, technology development, training and technology
transfer, and fund-raising to support in-situ conservation initiatives. This may
be true for the best zoos; however, too many elephants are being kept in zoos
where their welfare is compromised by their lack of social contacts and their
indirect contribution to conservation is negligible.
The welfare of elephants in zoos is a legitimate concern of the public, govern-
ments, and the zoo community. However, much of the recent concern in Europe
has been generated by a report (Clubb & Mason, 2002) that was based on
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inaccurate data—including an apparent overestimate of the size of the European
elephant population of 17%—and has been highly criticized by the European
Elephant Group (Endres et al., 2004). If welfare concerns are to be taken seri-
ously, they must be based on hard evidence, accurate statistics, and appropriate
Many zoos are failing to comply with the minimum requirements for the
sizes of groups of captive elephants recommended by their own regional or-
ganizations. The overall position is likely to improve as more zoos invest in
new elephant facilities and further movements occur between zoos. Births and
future importations of elephants may also have beneficial impacts on group sizes.
Individual zoos need to assess their own position carefully and either phase out
the keeping of elephants or commit themselves to improving their facilities and
increasing elephant group sizes.
This study is based on data collected by the International Species Information
System that is made freely available to the public via the Internet. I am grateful to
two anonymous reviewers who suggested improvements to an early draft of the
manuscript and to Dr. M. A. Bloomsmith (Zoo/Wildlife section editor, Journal
of Applied Animal Welfare Science) for her helpful comments and guidance.
American Association of Zoos and Aquariums. (2001). AZA standards for elephant management
and care. Silver Spring, MD: American Association of Zoos and Aquariums.
American Association of Zoos and Aquariums. (2007). Top ten AZA elephant success stories:
2005 a banner year for elephants in AZA accredited zoos. Retrieved April 23, 2007, from
Anonymous. (2006a). ISIS abstracts: Elephas maximus. International species information system.
Retrieved October 27, 2006, from
Anonymous. (2006b). ISIS abstracts: Loxodonta africana. International species information system.
Retrieved October 27, 2006, from
Carrington, R. (1958). Elephants. London: Chatto & Windus.
Conservation Breeding Specialist Group. (2004). AZA elephant management strategic planning
workshop (Draft report). Apple Valley, MN: IUCN/SSC Conservation Specialist Group.
Convention on International Trade in Endangered Species of Wild Fauna and Flora. (1973, March
3). Her Majesty’s Stationery Office, Treaty Series Nos. 101 (1976), Cmnd.6647.
Clubb, R., & Mason, G. (2002). A review of the welfare of zoo elephants in Europe. A report
commissioned by the RSPCA. Oxford, UK: University of Oxford, Department of Zoology, Animal
Behaviour Research Group.
Douglas-Hamilton, I., & Douglas-Hamilton, O. (1978). Among the elephants. Glasgow, UK: William
Collins Sons & Co. Ltd.
Downloaded By: [Rees, Paul A.] At: 23:10 23 December 2008
Eltringham, K. (1984). Elephants in zoos. Biologist, 31(2), 108–111.
Endres, J., Haufellner, A., Haufellner, B., Schilfarth, J., Schilfarth, M., & Schweiger, G. (2004).
Documentation 2002: Elephants in European zoos and safari parks. European Elephant Group.
Münster, Germany: Schüling Verlag.
European Association of Zoos and Aquaria. (2004). Revised EAZA elephant TAG recommendations.
EAZA News, 47, 44.
Faust, L. J., Thompson, S. D., & Earnhardt, J. M. (2006). Is reversing the decline of Asian elephants
in North American zoos possible? An individual-based modeling approach. Zoo Biology, 25, 201–
Garai, M. E. (1992). Special relationships between female Asian elephants (Elephas maximus) in
zoological gardens. Ethology, 90, 197–205.
Gilbert, J. (1994). Elephant feeder balls. The Shape of Enrichment,3(4), 3–5.
Green, C. (1993). Enriching an elephant’s environment. The Shape of Enrichment, 2(1), 5–6.
Haight, J. (1993). Playing with their food—Ideas for elephants. The Shape of Enrichment,2(2), 9.
Hedges, S., Tyson, M. J., Sitompul, A. F., & Hammatt, H. (2006). Why inter-country loans will not
help Sumatra’s elephants. Zoo Biology, 25, 235–246.
Hodgkins, L. D. (2000). AZA annual report on conservation and science 1997–98 (Vol. II): Member
institution conservation and research projects. Silver Spring, MD: American Zoo and Aquarium
Hutchins, M. (2006). Variation in nature: Its implications for zoo elephant management. Zoo Biology,
25, 161–171.
Hutchins, M., & Keele, M. (2006). Elephant importation from range countries: Ethical considerations
for accredited zoos. Zoo Biology, 25, 219–233.
Keele, M. (2006, February). The future of elephants is upon us. Communiqué (American Association
of Zoos and Aquariums), 6–8.
Laws, R. M., Parker, I. S. C., & Johnstone, R. C. B. (1975). Elephants and their habitats. The
ecology of elephants in North Bunyoro, Uganda. Oxford, UK: Clarendon.
Lee, P. C. (1987). Allomothering among African elephants. Animal Behaviour, 35, 278–291.
Lees, C. (2004). Captive management plan for Asian elephant (Elephas maximus). Sydne y, Australia:
Australasian Regional Association of Zoological Parks and Aquaria.
McKay, G. M. (1973). Behaviour and ecology of the Asian elephant in southeastern Ceylon.
Smithsonian Contribution to Zoology, 125, 1–113.
Moss, C. J. (1988). Elephant memories. London: Elm Tree Books.
Moss, C. J., & Poole, J. H. (1983). Relationships and social structure of African elephants. In R. A.
Hinde (Ed.), Primate social relationships: An integrated approach (pp. 315–325). Oxford, UK:
Nogge, G. (2004). Elefanten in Köln [Elephants in Köln]. Zeitschrift des Kölner Zoo, 47(3), 95–108.
Olson, D., & Wiese, R. J. (2000). State of the North American African elephant population and
projections for the future. Zoo Biology, 19, 311–320.
Rapaport, L., & Haight, J. (1987). Some observations regarding allomaternal caretaking among
captive Asian elephants (Elephas maximus). Journal of Mammalogy, 68(2), 438–442.
Reed, M. C. (2005, March). From the president. Communiqué (American Association of Zoos and
Aquariums), 5.
Rees, P. A. (2000). Are elephant enrichment studies missing the point? International Zoo News, 47,
Rees, P. A. (2001a). Captive breeding of Asian elephants (Elephas maximus): The importance of
producing socially competent animals. In B. B. Hosetti & M. Venkateshwarlu (Eds.), Trends
in wildlife biodiversity, conservation and management (Vol. 1, pp. 76–91). Delhi, India: Daya
Publishing House.
Rees, P. A. (2001b) The history of the National Elephant Centre, Chester Zoo. International Zoo
News, 48, 170–183.
Downloaded By: [Rees, Paul A.] At: 23:10 23 December 2008
Rees, P. A. (2003a). Asian elephants in zoos fa ce global extinction: Should zo os accept the inevitable?
Oryx, 37(1), 20–22.
Rees, P. A. (2003b). RSPCA elephant welfare recommendations would compromise captive breeding
programmes. International Zoo News, 50, 86–90.
Rees, P. A. (2003c). Early sexual experience in the Asian elephant. International Zoo News, 50,
Rees, P. A. (2004). Some preliminary evidence of the social facilitation of mounting behavior in a
juvenile bull Asian elephant (Elephas maximus). Journal of Applied Animal Welfare Science, 7,
Schmid, J., Heistermann, M., Ganslosser, U., & Hodges, J. K. (2001). Introduction of foreign female
Asian elephants (Elephas maximus) into an existing group: Behavioural reactions and changes in
cortisol levels. Animal Welfare, 10, 357–372.
Schmitt, D. L. (1998, June). Report of a successful artificial insemination in an Asian elephant.
Proceedings of the Third International Elephant Research Symposium, Springfield, MO.
Southwood, T. R. E. (1981). Bionomic strategies and population parameters. In R. M. May (Ed.),
Theoretical ecology: Principles and applications (2nd ed., pp. 30–52). Oxford, UK: Blackwell
Scientific Publications.
Spinage, C. (1994). Elephants. London: T. & A. D. Poysner Ltd.
Stevenson, M. F., & Walter, O. (2002). Management guidelines for the welfare of zoo animals:
Elephants Loxodonta africana and Elephas maximus. London: British & Irish Association of
Zoos & Aquariums, Regent’s Park.
Sukumar, R. (1994). Elephant days and nights: Ten years with the Indian elephant. Oxford, UK:
Oxford University Press.
Sukumar, R. (2003). The living elep hants: Evolutionary ecolog y, behavior a nd conservation. Oxford,
UK: Oxford University Press.
Veasey, J. (2006). Concepts in the care and welfare of captive elephants. Intern ational Zoo Yearbook,
40, 63–79.
Wiese, R. J. (2000). Asian elephants are not self-sustaining in North America. Zoo Biology, 19,
Wiese, R. J., & Willis, K. (2004). Calculation of longevity and life expectancy in captive elephants.
Zoo Biology, 23, 365–373.
Wiese, R. J., & Willis, K. (2006). Population management of zoo elephants. International Zoo
Yearbook, 40, 80–87.
Williams, J. H. (1951). Elephant Bill. London: Rupert Hart-Davis.
Downloaded By: [Rees, Paul A.] At: 23:10 23 December 2008
... Despite the challenges currently facing Asian elephant breeding efforts [including many non-reproductive individuals , complicated logistics involved with bringing receptive mates together and/or artificial insemination (Schmitt, 2003;Hildebrandt et al., 2006;Schmitt, 2006;Kiso et al., 2007;Kiso, 2011), and elephant endotheliotropic herpesvirus hemorrhagic disease (EEHV-HD), a potentially fatal condition that has resulted in the mortality of a significant number of prepubertal animals (Richman et al., 1999;Long et al., 2015;Zachariah et al., 2018)], sustainability in the North American and European Asian elephant populations is improving (Nordin, 2017;Schmidt & Kappelhof, 2019). As breeding success increases, there will soon be a larger number of mature males housed in zoos and similar facilities than there has ever been (Hildebrandt et al., 2006;Wiese & Willis, 2006;Rees, 2009b). Historically, captive elephant populations that depended on importing wild specimens were female-biased because of the relative ease of housing, caring for, and handling female elephants (Wiese & Willis, 2006). ...
... Unfortunately, the sustainability of many ex-situ elephant populations is at risk (Wiese, 2000;Thitaram, 2012), primarily due to limited reproductive success with complicated breeding logistics and a small number of suitable candidates (Schmitt, 2006;Kiso et al., 2007;Brown et al., 2016), and fatal hemorrhagic disease associated with elephant endotheliotropic herpesvirus (Long et al., 2015;Zachariah et al., 2018). Historically, these captive populations have been female-biased, as female elephants were preferentially imported from the wild due to their tractability and often without the intent to establish captive breeding programs (Rees, 2009b). However, there is now decreased reliance on wild importations as breeding success is steadily improving (Wiese & Willis, 2006;Clubb et al., 2009); the European population of Asian elephants is already thought to be self-sustaining (Schmidt & Kappelhof, 2019). ...
In-situ and ex-situ populations of Asian elephants (Elephas maximus) are unsustainable in the long-term and require integrative approaches to address issues impacting their conservation status. Male elephants pose distinct challenges: they are more likely to engage in human–elephant conflict in the wild, and they require specialized care in zoos and other similar settings. Musth is a unique sexual state in elephants and is thought to be a major contributor to these challenges. However, our limited understanding of the nature of musth in Asian elephants limits our ability to implement animal-centered conservation strategies. The purpose of this dissertation was to characterize the behavioral and physiological variation of musth in wild and zoo-housed male Asian elephants in Sri Lanka and North America, respectively. This work was organized by four principal objectives: (1) identify intrinsic and extrinsic factors that influence behavioral activity around musth in wild and zoo-housed male elephants; (2) investigate the effects of age and musth on male social behavior in wild and zoo-housed elephants, and group formation in wild elephants; (3) utilize fecal samples collected longitudinally from zoo- housed elephants to determine factors that influence musth and three fecal hormone metabolites potentially associated with musth; and (4) integrate behavioral and hormonal datasets from wild and zoo-housed elephants to explore potential associations between behavioral and hormonal variation in males. To draw qualitative comparisons, data collection procedures were similar between in-situ and ex-situ elephant populations. In the first study, I identified characteristic behaviors that changed with the progression of musth in both wild and zoo-housed elephants, including increased activity, investigatory behavior, locomotion, and stereotypy (in zoo elephants), and decreased foraging. Behavior in zoo elephants was also influenced by factors such as age, space availability, and temperature. Based on these results, I proposed four behavioral stages of musth (non- musth, early musth, full musth, and post-musth) that can be reliably monitored with visible musth indicators [i.e., temporal gland secretions (TGS) and urine dribbling (UD)]. For the second study, I used a similar observational approach to identify factors that influence social behavior in male elephants. Results of this study showed that group formation in wild male elephants was associated with the interaction of age and musth status. Furthermore, broad categories of social behavior (aggression, prosocial behavior, dominance behavior, and submissive behavior) were correlated with changes in the social environment in wild and zoo-housed elephants, and these behaviors also were associated with age and musth status in zoo elephants. In the third study involving physiological variation of musth in zoo-housed elephants, I found strong associations of intrinsic and extrinsic factors with concentrations of fecal androgen metabolites (FAM) and fecal glucocorticoid metabolites (FGM). Additionally, the duration of musth episodes was negatively associated with body condition and positively associated with exposure to male conspecifics. TGS and UD activity also was correlated with changes in FAM and FGM concentrations. Finally, the fourth study demonstrated that many of the behavioral changes characteristic of musth also were associated with increased FAM concentrations (and increased active behavior and locomotion was positively related to FGM concentration in zoo-housed elephants), illustrating the multiple scales with which male elephants respond to changing internal and external environments. This dissertation emphasized the inherent variation associated with behavioral and physiological correlates of the heightened reproductive state of musth in Asian elephants, an endangered species with in-situ and ex-situ sustainability challenges. In identifying the intrinsic and extrinsic influences of this variation, we can develop informed approaches to conserve elephants as they respond to ever-changing internal and external environments.
... Elephants have complex behaviours and need for social interactions, as well as need for an extensive and enriched environment (2,3). In the wild, they have complex social structures consisting of related individuals (4,5,6), while in zoos elephant groups often consist of randomly grouped individuals with numbers below the recommended for breeding facilities (5,7). All of this leads to a difficulty in maintaining their welfare in captivity (2,3), which may result in stereotypic behaviour and health concerns (2,3,8,9). ...
... Elephants have complex behaviours and need for social interactions, as well as need for an extensive and enriched environment (2,3). In the wild, they have complex social structures consisting of related individuals (4,5,6), while in zoos elephant groups often consist of randomly grouped individuals with numbers below the recommended for breeding facilities (5,7). All of this leads to a difficulty in maintaining their welfare in captivity (2,3), which may result in stereotypic behaviour and health concerns (2,3,8,9). ...
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Providing good animal welfare standards is very important for wild animals in captivity, especially in zoos. Therefore, the aim of this study was to perform a welfare assessment of elephants in the Skopje Zoo using species-specific protocols. Two specific protocols were used for the welfare assessment and were combined for a more unified approach. These protocols focused on the assessment of the elephant's daytime behaviour, including Qualitative Behaviour Assessment (QBA), as well as the following sections: nutrition, physical health, environment, behaviour and management. Data was collected from the Asian and African elephants that were housed together, in the Skopje Zoo. Both species were observed for three consecutive days, during which time QBA was performed and they were observed on their daytime behaviour. Feeding (42% African, 34% Asian elephant) and anticipatory (35% African, 22% Asian elephant) behaviour were predominantly observed daytime behaviours of both elephants, as well as stereotypic behaviour (30%) of the Asian elephant. Concerning the QBA, 'uncomfortable' (296 Asian, 234 African elephants) and 'relaxed' (271 Asian, 280 African elephants) were the most dominant descriptors for both elephants. The collected data indicated that the elephants were in good physical health. Regular feeding enrichment, as well as environmental enrichment was recommended. There is room for improvement concerning the management practices of the elephants.
... Historically, captive elephant populations have been female-biased, as female elephants were preferentially imported from range countries due to their tractability and often without the intent to establish ex-situ captive breeding programs (Rees, 2009). However, there is now decreased reliance on wild importations, with populations relying almost entirely on ex-situ breeding efforts (Wiese and Willis, 2006;Clubb et al., 2009;Scherer et al., 2022). ...
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The sustainability of endangered Asian elephants in human care is threatened in part by low breeding success and concerns over individual animal wellbeing. Male elephants have received less research attention compared to females, yet males deserve special consideration due to their unique reproductive biology (particularly the sexual state of “musth”) and the complex interaction of physiological, environmental, and social pressures they face. We measured fecal androgen metabolites (FAMs), fecal glucocorticoid metabolites (FGMs), and fecal triiodothyronine metabolites (FT3s) collected weekly over approximately 12 months from 26 male Asian elephants housed in zoos across the US, hypothesizing that FAM, FGM, and FT3 concentrations would be associated with temporal correlates of musth and would vary further with intrinsic (musth status, age, body condition) and extrinsic (social environment) factors. The duration of each musth episode was positively associated with exposure to male conspecifics and negatively associated with body condition. Further, elevated FAM concentrations were associated with social exposure, age, and body condition, and FGM concentrations also varied with age and body condition. FT3 concentrations were not associated with any factor we measured. We also identified periods of lower FAM concentration than confirmed musth episodes (but still higher than baseline FAM concentrations) that we termed “elevated FAM episodes.” The durations of these episodes were negatively correlated with exposure to other male elephants. Together, these results provide evidence that hormone profiles (including those that are predicted to change around musth) vary significantly between male Asian elephants in a way that may be attributed to intrinsic and extrinsic factors. Studies like these serve to enhance the sustainability of ex-situ populations by providing wildlife managers with information to enhance the health, welfare, and reproduction of threatened species like Asian elephants.
... As more institutions house bulls socially, we hope our results provide insights into some of the demographic and life-history factors that may affect bull social interactions during the introductory period and beyond. Improving animal welfare is our "duty of care" [62] and as animal managers we can no longer disregard the evidence of bull elephant sociality [48,49,63]. Bull elephants often exhibit more agonistic behaviors than females [42], leading to common, and well-founded, concerns about socializing bulls in managed care. ...
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Wild bull Asian elephants spend time in all-male groups. Therefore, managers of ex situ populations increasingly house bulls together. We examined the social interactions of five bull Asian elephants at Denver Zoo, using instantaneous sampling to compare social interactions across adolescent and mature bulls, and bulls with a social history prior to the integration of this group compared to bulls with no social history. Both age and social history significantly affected bull behavior. Adolescent bulls exhibited more affiliative and submissive behaviors when housed with mixed-age and mature social partners compared to with only adolescents, and less non-contact agonistic behavior and less time in proximity to a conspecific with mixed-age groups compared to with only other adolescents. Mature bulls exhibited more affiliative behavior when they were with only adolescent bulls compared to only mature bulls, and more time in proximity to a conspecific and increased contact agonistic behavior with at least one adolescent compared to only mature bulls. Bulls in new social groups engaged in more affiliative, agonistic, and submissive behaviors, and spent less time in proximity, than when they were in previously established social combinations. As more institutions house bulls socially, our results provide insights into factors that may affect bull social interactions.
... In the U.S., the number of Association of Zoos and Aquariums-accredited facilities holding elephants has decreased [23]. However, the number of male elephants in the population has increased as a result of more breeding success and a 1:1 birth sex ratio [24]. ...
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Zoological institutions aim to continually improve the lives of the animals under their stewardship. To this end, bull elephants are now increasingly maintained in all-male groups to mimic social conditions observed in the wild. While cortisol is the most frequently used “stress” biomarker, secretory immunoglobulin A (sIgA) as a measure of health and positive affect, and the social hormone, oxytocin, are increasingly viewed as additional markers of welfare. The introduction of a pair of bull elephants to an existing group of three bull elephants at Denver Zoo presented an opportunity to assess sIgA, oxytocin and cortisol in response to the socialization process. In this study, sIgA varied greatly between individuals and did not correlate with cortisol but did correlate with salivary oxytocin. sIgA and oxytocin concentrations differed the most between social and solo situations during the introduction period compared to before bulls were introduced, and after a stable group had been formed. In contrast to findings in some species, sIgA and oxytocin were higher when housed alone than socially. Nonetheless, these results suggest that sIgA and oxytocin may be involved in social engagement and establishment of new social dynamics, and thus provide more insight into overall welfare states.
... These facilities are critically important to the continued existence of this species [3][4][5]. Historically, captive populations have been female-biased, as female elephants were preferentially imported from the wild due to their tractability [6]. However, there is now decreased reliance on wild importations as breeding success is steadily improving [7,8], and so the proportion of male elephants in ex-situ populations is increasing markedly [9]. ...
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In-situ and ex-situ Asian elephant populations are threatened with extinction, and male elephants pose unique challenges to long-term sustainability. The heightened sexual state of “musth” is accompanied by a suite of physical, behavioral and physiological changes. Furthermore, musth is unique to male elephants and requires special consideration when developing short- and long-term management strategies for elephants in the wild and in human care. The purpose of this study was to identify associations between fecal hormone metabolites [fecal androgen metabolites, FAM; fecal glucocorticoid metabolites, FGM; and fecal triiodothyronine (T3) metabolites, FT3] and visible musth indicators [temporal gland secretions (TGS) and urine dribbling (UD)], and behavioral changes around musth. From fecal samples collected non-invasively from wild elephants in Wasgamuwa National Park, Sri Lanka, and zoo-housed elephants in the United States, we hypothesized that (1) TGS and/or UD would be associated with changes in FAM, FGM, and/or FT3 concentrations; (2) variation in fecal hormone metabolites would be associated with increased locomotion and chemosensory behavior, and decreased foraging; and (3) relationships we identified would be similar between wild and zoo-housed elephants. We found that FAM concentrations changed significantly with TGS and UD activity in both wild and zoo elephants. Further while FGM concentrations were higher with increased TGS and UD in zoo elephants, the opposite pattern occurred in wild elephants. We did not identify substantial change in FT3 concentrations with TGS/UD activity. Behavioral changes in zoo elephants were significantly associated with FAM concentration as predicted, but these relationships were more difficult to identify in wild elephants due to lower sample availability. Further, FGM concentration was directly related to time spent locomoting in zoo elephants, but no other apparent association existed between FGM concentration with other behaviors in zoo elephants, or in any behaviors in wild elephants. Likewise, we did not report associations between FT3 and any behaviors we measured. This study contributes to our understanding of the complex response patterns that male Asian elephants exhibit around musth, and it provides another example of complementary in-situ–ex-situ research that can be directly applied to improve the well-being of elephants and other wildlife.
... If SB are a factor in the prevalence of captive elephant foot disease, the influences that increase the occurrence of abnormal behaviour, in addition to the actual SB, both need to be prevented in order to successfully lower the risk of disease and mortality. The potential stimuli causing SB includes low temperature, predictable routines, boredom, small group size , substrate type, and lack of space within an enclosure (Rees, 2004;Rees, 2009b;Rees, 2009a;Stroud, 2007). Sherwin, Harris and Harris (2008) describe many different SB including pacing, head bobbing, and route tracing with some observed in combinations. ...
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A copy of my original thesis submitted as part of my BSc (Hons) Animal Biology degree. The study was a meta-analysis discussing issues and care of, the feet of captive asian elephants, with an aim of collating the most recent information at the time of writing, so as to help better inform the then-current care of captive elephants.
... It is widely thought that behavioral restriction, including reduced foraging and social opportunities, can lead to stereotypic behavior in elephants and that these abnormal behaviors indicate compromised welfare [10,[22][23][24][25]. In a study across 194 institutions, 20% of all elephants were housed alone or with only one conspecific, contrary to management recommendations for females, which advise that they be housed in groups; recommendations for social housing of male elephants are not provided [26,27]. ...
Full-text available
There is a growing need for animal care institutions to house multiple bull elephants as the population increases due to transfers from private ownership and the births of male offspring in managed care. Elephants in North American, European, and Latin American zoos exhibit stereotypies—repetitive, fixed behaviors. Previous research demonstrated that housing Asian elephants alone increased stereotypic behavior. Therefore, for animals in managed care, social restriction can contribute to stereotypy and, by extension, reduce welfare. In this study, we examine how being alone affects stereotypic behavior by monitoring pacing and head-bobbing in individual bull Asian elephants at Denver Zoo when housed alone as well as with other bulls. Two young males arrived at Denver Zoo in September 2018 and joined an existing all-male group of three elephants that were previously socialized and housed together. From July 2018 to December 2019, we used instantaneous scan sampling to collect data on stereotypic behavior of focal bulls when they were housed alone and socially. The frequency of pacing and head-bobbing significantly decreased when the elephants were housed socially compared to when they were housed alone; these stereotypies were lower when elephants were housed with at least one other bull and were in close proximity to a conspecific. Additionally, pacing decreased as the proportion of affiliative behaviors increased, and the amount of agonistic behavior did not significantly affect stereotypic behavior. When housed alone, bulls in musth engaged in significantly more pacing behavior than when they were out of musth. Our results indicate that housing bull Asian elephants in all-male groups substantially improves their welfare by reducing stereotypies and provides a basis for future evidence-based management.
Zoos and aquariums are culturally and historically important places where families enjoy their leisure time and scientists study exotic animals. Many contain buildings of great architectural merit. Some people consider zoos little more than animal prisons, while others believe they play an important role in conservation and education. Zoos have been the subject of a vast number of academic studies, whose results are scattered throughout the literature. This interdisciplinary volume brings together research on animal behaviour, visitor studies, zoo history, human-animal relationships, veterinary medicine, welfare, education, enclosure design, reproduction, legislation, and zoo management conducted at around 200 institutions located throughout the world. The book is neither 'pro-' nor 'anti-' zoo and attempts to strike a balance between praising zoos for the good work they have done in the conservation of some species, while recognising that they face many challenges in making themselves relevant in the modern world.
The first recorded elephant in England arrived in 1255; a gift to King Henry III from King Louis IX of France. Elephants have been kept in zoos in the United States for over 200 years and, throughout the world, they are extremely popular with zoo visitors. Since the 1980s there have been captive breeding programmes for elephants in zoos but these have not been successful in creating self-sustaining zoo populations. Many people now believe that the welfare compromises endured by captive elephants outweigh any actual or potential benefits for elephant conservation that accrue from their being kept in zoos. Although there have been recent improvements in the longevity of elephants living in zoos, many zoos have given up keeping elephants and some have sent their animals to elephant sanctuaries. This case study considers the relative merits of the claims that elephants living in zoos have conservation and scientific functions and the counter-claims that, not only do they have no legitimate function, but they suffer significant welfare harms.
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Captive breeding programmes for the Asian elephant Elephas maximus have failed to establish self-sustaining zoo populations. Birth rates are low and calf mortality rates are high. The zoo population is widely dispersed, with few animals being moved on breeding loan. New techniques may increase birth rates but current predictions suggest demographic extinction within 50 years. It would be difficult to justify importing elephants from sustainable zoo reserves in Asia to participate in ex situ breeding programmes where reproductive success is low. Zoos should either urgently regroup animals to form breeding units, or accept that Asian elephants will die out in zoos and that funds should be diverted to in situ conservation projects.
The present study examined the extent to which the introduction of three female Asian elephants (aged 3, 11, and 27 years) into a group of 1.4 (1 male, 4 female) elephants at Münster zoo, Germany, affects the behaviour and urinary cortisol levels of the animals involved. At Münster, only the females were monitored — the bull was mainly kept separate. Behavioural observations were carried out before transfer and during the six-month period following transfer, and urine samples were collected regularly from each elephant during the whole observation period. All elephants showed behavioural changes to the process of introduction. The transferred animals increased their social behaviour after arrival in the foreign zoo. Two of them showed an increase in stereotypies and one a reduction in stereotypies. The elephants at Münster reacted with decreased frequencies of stereotypies and increased frequencies of social behaviour and manipulation/exploration behaviour. Six months after transfer, three of the four elephants at Münster and one of the three transferred elephants showed nearly the same behavioural activity pattern as before transfer. One female still showed elevated stereotypic behaviour. From the four elephants in which cortisol measurements could be reliably performed (two of the transferred elephants and two elephants at Münster), only one individual at Münster responded to the process of introduction with a short-term elevation in urinary cortisol levels. One elephant showed a negative correlation between locomotion and cortisol levels and one a positive correlation between stereotypies and cortisol levels. Taken together, the results suggest that transfer and introduction caused some stress responses in the elephants, but that stress was neither prolonged nor severe. Serious welfare problems may have been prevented through individual behavioural coping mechanisms and former experience with stressful situations.
Reviews life strategies and they are affected by size, longevity etc. Also discusses the 'natural enemy ravine'.