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Captive Breeding

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 178 Captive Breeding
area between the tip of the rostrum and the blowholes, referred to as
a “continuous” callosity (found only rarely in southern right whales).
Identification can be confounded since the whale lice on and around
the callosities are mobile, sometimes masking the true edge of the cal-
losity. However, by using additional distinctive features, including the
three-dimensional configuration of the callosities, additional scars or
marks, and crenulations on the lower lip, North Atlantic researchers
have still been able to reliably identify each individual. Photographic
C catalogs of identified right whales, primarily of their callosity patterns,
are maintained for the North Atlantic Ocean (by the New England
Aquarium, Boston, MA), the North Pacific Ocean (the US National
Marine Fisheries Service), and in the numerous places in the Southern
Hemisphere, including off of South America near Peninsula Valdez,
Argentina and off the coast of Brazil, off the coast of South Africa,
and off of New Zealand’s sub-Antarctic Auckland Islands (Best et al.,
2001). Additional collections of photographs of individual right whales
based on callosities and other natural markings exist in various institu-
tions around the world.
See Also the Following Article
Right Whales
References
Best, P. B., Bannister, J. L., Brownell, R. L., and Donovan, G. P. (2001).
Right whales: Worldwide Status. Journal of Cetacean Research and
Management(Special Issue 2), 309.
Hamilton, P. K., and Martin, S. M. (1999). “A Catalog of Identified
Right Whales from the Western North Atlantic: 1935 to 1997.” New
England Aquarium, Boston, MA.
Kraus, S. D., Moore, K. E., Price, C. A., Crone, M. J., Watkins, W. A.,
Winn, H. E., and Prescott, J. E. (1986). The use of photographs to
identify individual northern right whales (Eubalaena glacialis).
Reports of the International Whaling Commission, Special Issue 10,
145–151.
Matthews, L. H. (1938). Notes on the southern right whale, Eubalaena
australis. Discovery Report, 17, 169–182.
Omura, H., Ohsumi, S., Nemoto, T., Nasu, K., and Kasuya, T. (1969).
Black right whales in the North Pacific. Sci. Rep. Whales Research
Institute, Tokyo 21, 1–78.
Payne, R., and Dorsey, E. M. (1983). Sexual dimporphism and aggres-
sive use of callosities in right whales (Eubalaena australis). In
“Communication and Behavior of Whales” (R. Payne, ed.), pp. 295–
329. AAAS Selected Symposium 76, Westview Press, Boulder, CO.
Payne, R. S., and Rowntree, V. J. (1984). Southern right whales: A pho-
tographic catalogue of individual whales seen in waters surrounding
Peninsula Valdes, Argentina. Unpublished Report available from
Center for Long Term Research, Inc., 191 Weston Road, Lincoln,
MA 01773.
Payne, R., Brazier, O., Dorsey, E.M., Perkins, J.S., Rowntree, V.J.,
and Titus, A. (1983). External features in southern right whales
(Eubalaena australis) and their use in identifying individuals. In
“Communication and Behavior of Whales” (R. Payne, ed.), pp. 371–
445. AAAS Selected Symposium 76, Westview Press, Boulder, CO.
Rowntree, V. (1983). Cyamids: The louse that moored. Whalewatcher
(J. American Cetacean Society) 17(4), 14–17.
Rowntree, V. (1996). Feeding, distribution, and reproductive behavior
of cyamids (Crustacea: Amphipoda) living on humpback and right
whales. Can. J. Zool. 74, 103–109.
Tomilin, A. G. (1957). Mammals of the USSR, Vol. 8: Cetacea.
Translated by the Israel Program for Scientific Translations. Available
from National Technical Information Service, 5285 Port Royal Road,
Springfield, VA 22161.
Captive Breeding
TODD R. ROBECK, JUSTINE K. O’BRIEN AND
DANIEL K. ODELL
I. Marine Mammals in Captivity
A
nimals have been held in captivity in one form or another for
hundreds, if not thousands, of years. Private collections turned
into “public” collections. A private animal collection at Schloss
Schönbrunn, Vienna, Austria, was opened to the public in 1765 and is
considered to be one of the first modern zoos. The first marine mam-
mals to be held in captivity may have been polar bears (Ursus mar-
itimus) and various pinnipeds. Reeves and Mead (1999) provide an
excellent overview of marine mammals in captivity. Harbor porpoises
(Phocoena phocoena) may have been held as early as the 1400s, polar
bears since about 1060, and walruses (Odobenus rosmarus) since
1608. As with terrestrial animals, marine mammals were held in pri-
vate collections.
However, most pinnipeds and some sirenians were not held in
captivity until the late 1800s and early 1900s. Being considerably
more difficult to capture, transport, and maintain, cetaceans, with
few exceptions, have only been held in captivity since the mid-1900s.
According to Reeves and Mead (1999), 4 species of sirenians, 33
pinnipeds, 51 cetaceans, the polar bear, and the sea otter (Enhydra
lutris) have been held in captivity. Of these, 2 species of sirenians, 22
pinnipeds, 15 cetaceans, the polar bear, and the sea otter have repro-
duced in captivity. Among these, however, only a few species, such
as the polar bear, California sea lion (Zalophus californianus), harbor
seal (Phoca vitulina), bottlenose dolphin (Tursiops truncatus), and
killer whale (Orcinus orca), have enough numbers and have been
reproductively managed with enough production to be considered
part of a successful captive breeding program.
Successful captive breeding of any marine mammal requires a
combination of an appropriate habitat, adequate nutrition and a
social structure that is conducive toward successful reproduction. It
becomes obvious when analyzing the history of successful births and
survivorship of these species in captivity that early animal managers
had little thought or, in some cases, knowledge of the requirements
necessary for the development of successful breeding programs. In
contrast, past records of breeding and survivorship with recent trends
beginning in the mid-1980s where captive breeding successes have
equaled or, in some cases, surpassed the best scientific estimates of
wild population breeding and survivorship, and one can see just how
far the captive marine mammal community has evolved. Detailed
censuses of captive marine mammals in North America (Asper et al.,
1990; Andrews et al., 1997) have shown the increasing numbers of
captive-bred marine mammals, particularly California sea lions, har-
bor seals, and bottlenose dolphins. In 1995, 70% of the California
sea lions, 56% of the harbor seals, and 43% of the bottlenose dol-
phins on display in North American facilities were captive-born. This
compares with 3%, 4%, and 6%, respectively, in 1975.
II. Why Breed Marine Mammals in Captivity?
A. Legal Necessity
In the “early days,” when animals were just being displayed as
“curiosities,” it was easier to collect replacements when animals died.
In most countries today this is simply not possible. Despite only a
 Captive Breeding
few species of marine mammals being threatened or endangered,
they and their habitats are protected by a myriad of national and
international laws and regulations. Although a number of countries
are considering regulation of the minimum conditions (e.g., pool size
and volume, water quality, food quality and handling, medical care)
under which marine mammals may be held in zoos, marine parks, or
research facilities, apparently only the USA has such regulations in
place. Even though the natural habitat of most marine mammals can-
not be duplicated in captivity, the trend is toward larger, more com-
plex, habitat-oriented displays and exhibits. Together, the various laws
and regulations have reduced the collection of wild marine mammals
and have eliminated smaller facilities that did not have the financial
resources to adequately provide for their animals as required by law.
The just-mentioned laws and regulations favor captive breeding
programs. A successful captive breeding program eliminates costly
field expeditions and animal transports.
B. Maintaining/Enlarging a Captive Population
A successful captive breeding program can provide animals for
other institutions with adequate holding facilities but without the
financial resources to maintain a breeding colony or (if even pos-
sible) to collect wild animals. Captive-born animals have a known
medical history and, to some extent, are imprinted on their keep-
ers. Captive breeding programs have, out of necessity, reduced the
impact on wild populations of marine mammals. Professional organi-
zations such as the Association of Zoos and Aquariums (AZA) have
established studbooks and other programs to assist with the manage-
ment of captive animal breeding colonies. For example, studbooks
track individual animals from birth to death and their reproductive
histories. Computer programs are used to pair animals to optimize
genetic diversity. In the USA, formal studbooks exist for the beluga
whale (Delphinapterus leucas), common bottlenose dolphin, Florida
manatee (Trichechus manatus latirostris), gray seal (Halichoerus
grypus), harbor seal, northern fur seal (Callorhinus ursinus), and
polar bear and several others are under development. Similar stud-
books are in place on other continents. The AZA also hosts a Marine
Mammal Taxon Advisory Group whose function is to promote man-
aged captive breeding of marine mammals.
C. A Breeding Program as a Scientific Resource
A successful captive breeding program is a unique scientific
resource in that it allows one to document, in great detail, various
aspects of reproductive behavior, reproductive physiology, and the
subsequent birth, growth, and development of the offspring. This
is particularly valuable for cetaceans, which are typically difficult to
study in great detail in their natural habitat due to various environ-
mental factors and the fact that they spend most of their lives under-
water. It is, however, most important to recognize that studies on
captive marine mammals, even in the best breeding colonies, do not
and cannot replace field studies. Both types of studies (i.e., laboratory
and field) are necessary to fully describe the biology of a species.
Routine components of a proper animal husbandry program include
regular physical exams, collection of blood, urine, and fecal samples,
body measurements, and body weights. These samples and data are vir-
tually impossible to collect from wild marine mammals. Consider, e.g.,
what it would take to get a daily urine sample from a wild bottlenose
dolphin or killer whale! Captive animals are easily conditioned to pro-
vide urine samples and to station for blood sampling, body measure-
ments, and so on. Figures 1, 2, and 3 illustrate the kinds of observations
that can be made and the kinds of data that can be gathered.
179
C
Figure 1 Birth of a killer whale (Orcinus orca) at SeaWorld
Florida. Photo credit: SeaWorld Florida.
D. A Breeding Program as a Conservation Resource
A successful captive breeding program may provide the physical
and human resources necessary to save some species of marine mam-
mals in imminent danger of extinction (Ralls and Meadows, 2000).
These resources may allow us to maintain a viable gene pool until the
habitat can be restored or other reasons for endangerment are elimi-
nated. While such an undertaking is certainly honorable and the right
thing to do, the magnitude of the job should not be underestimated
and there are, at the present time, obvious limits based in good meas-
ure on the sheer size of the animals. For example, the population
of right whales (Eubalaena glacialis) in the western North Atlantic
Ocean is about 350 animals and may be decreasing due to human
activities. These animals may reach lengths of 18 m and weights on
the order of 20 tons. No facility in existence today (or likely to be in
the foreseeable future) could hold a breeding group of right whales.
However, threatened or endangered marine mammals such as the
Hawaiian and Mediterranean monk seals (Monachus schauinslandi
and M. monachus, respectively) and the river dolphins (families
Platanistidae, Iniidae, and Pontoporiidae) could be maintained in
viable captive breeding colonies. The importance of taking immedi-
ate action on behalf of these species is demonstrated by the recent
extinction of a small cetacean, the baiji (Lipotes vexillifer). If a cap-
tive breeding colony had been established, this species would still be
present today and this colony may have provided potential animals for
reintroduction to the wild.
III. Assisted Reproductive Technologies (ART)
The domestic animal industry [cattle (Bos spp.), pigs (Sus spp.),
horses (Equus caballus)] long ago realized that it is much more effi-
cient to move genetic material (e.g., semen) to different facilities
than it is to move the animal. Methodologies for semen collection,
preservation, and transportation, along with methods for artificial
insemination (AI), induction of superovulation, and embryo collec-
tion and transfer, were developed and are in widespread use world-
wide today. Some of these techniques have been applied successfully
to endangered animals. And recently, AI has been successfully devel-
oped in three species of marine mammals; killer whales, bottlenose
dolphins, and Pacific white-sided dolphins (Lagenorhynchus obliqui-
dens) (Robeck et al., 2003, 2004, 2005).
What then are the needs for the development and application of
ART to marine mammals? The most obvious ART that should be
 180 Captive Breeding

50,000

False
40,000 pregnancy?
Pregnancy Pregnancy Pregnancy
Progesterone (pg/ml)

30,000
C
20,000

Estrus Estrus
Calf cycling
10,000 Calf cycling
born
born Calf
born

0
1500 1900 2300 2700 3100 3500 3900 4300 4700 5100 5500
Age (days)

Figure 2 Serum progesterone of killer whale (Orcinus orca) from birth through sexual matu-
rity and calving and showing estrous cycling and pregnancies. Progesterone levels were “zero”
until after 1900 days of age.

600

500
Total length (cm)

400

300
⫺0.001X
Y ⫽ 539.60 (e⫺0.76 e )

200
0 1000 2000 3000 4000 5000 6000
Age (days)

Figure 3 Killer whale (Orcinus orca) growth (length) curve as an example of data
that can be gathered from captive-bred animals.

developed and the one that is most likely to have an immediate impact
on the genetic management of captive marine mammals is AI. Once
developed in all captive species, AI can provide an immediate mecha-
nism for marine mammal managers to increase the genetic fitness of
their respective populations without having to rely on the transporta-
tion of animals between facilities. Wild animal population studies have
shown that dolphins develop strong social ties to other animals and
that these bonds can be maintained throughout the life of the animals
(Connor et al., 2000). To what extent these bonds are important for
the health of these animals can only be speculated, but it seems pru-
dent for managers who have groups of compatible animals to carefully
consider the impact of removing or introducing individuals on the
population social dynamics. Further, bringing animals from other loca-
tions may expose the new population of animals to bacterial or viral
organisms, which they have no natural resistance against.
The development of AI, the most common assisted reproductive
technology, for commercial use began in the 1950s. The successful
application of AI and other ART to domestic species and humans was
 Captive Breeding
in part because of their accessibility for research into their reproduc-
tive mechanism or reproductive physiology. Critical importance for
the successful application of ART in any species is the determina-
tion of how reproductive hormones, particularly luteinizing hormone
(LH), relate to ovulation (Sorensen, 1979).
Once ARTs were developed in domestic species, it was naively
believed that they could easily be transferred to exotic species.
However, relatively little success was realized with exotic animals until
the late 1970s when an endocrinological breakthrough occurred. This
breakthrough was the ability to analyze reproductive hormones in
urine. This technology was successfully used to characterize the endo-
crinological events in a wide range of exotic species, including for the
first time in marine mammals; the killer whale (Walker et al., 1988).
Other smaller cetacean species proved more difficult to train for urine
collection, and detailed endocrinological evaluations would have to
wait until a technique was developed for training urine collection in
bottlenose dolphins (Lenzi, 2000). This simple technique has since
revolutionized urine collection and the subsequent endocrine evalu-
ation of cetaceans (whereas Pinnipeds are yet to undergo such exten-
sive evaluation). As a result, nearly 15 years after the first published
report of urinary hormones in killer whales, endocrine characteristics
have now been reported in four additional cetacean species.
Although the capability to characterize the endocrine cycle is criti-
cal, for it to be useful to predict the timing of the actual insemination,
it must first be related to ovulation. For deduction of the temporal
relationship between urinary hormones and ovulation in cetaceans,
two important techniques had to be developed. This first was an assay
system that could rapidly detect the LH surge—a brief and dramatic
hormonal surge that in most mammalian species precedes ovulation
at a regular interval. The second was to develop a method to allow
for consistent observation of ovarian activity using transabdominal
ultrasonography. Brook (2001) was the first to develop a technique
to accomplish this in the bottlenose dolphin. This simple technique
which relies on understanding the anatomical location of the ovaries in
relationship to the abdominal musculature has since been successfully
utilized to consistently locate ovaries in many other captive marine
mammals, including the killer whale, beluga, the Pacific white-sided
dolphin (Fig. 4), and the false killer whale (Pseudorca crassidens).
Finally, by combining this ultrasound technique with rapid LH assay
systems, landmark research was completed that allowed characteriza-
tion of the timing between the LH surge and ovulation (Robeck et al.,
2004, 2005). This information then opened the door for the develop-
ment and application of AI in cetaceans.
Once ovulation timing could be predicted, the optimum timing of
insemination had to be systematically determined for each species.
However, before any AI attempts could be made the collection of
semen had to be developed. Currently, collection of semen on a rou-
tine basis has been achieved only in a few marine mammal species,
and as one would expect, this list correlates with the species where
AI has now been accomplished. The first species where semen collec-
tion was successfully trained was the bottlenose dolphin (Keller, 1986).
Bottlenose dolphins seem to have consistently elevated libidos and thus
were relatively easily trained to provide semen via manual stimulation.
Since the success with bottlenose dolphins, four killer whales and three
Pacific white-sided dolphins and one beluga have been trained to pro-
vide semen on a regular basis. Obviously, to reach the full potential of
AI, all genetically valuable males should be trained to provide semen.
Once the semen has been collected, it must be stored temporar-
ily for immediate use or permanently by cryopreservation. Semen
cryopreservation has been successfully accomplished by different
methods (straws, pellets, or cryovials) in all of the species for which
181
Skin
1 cm Blubber
Muscle
Follicle
Ovary C
Figure 4 Sonographic image of an ovary of a Pacific white-sided
dolphin (Lagenorhynchus obliquidens) showing a follicle.
semen has been obtained (Schroeder and Keller, 1990; Robeck and
O’Brien, 2004; Robeck et al., 2004; O’Brien and Robeck, 2007).
Although cryopreserved semen provides a long-term supply of
genetic material from a particular animal and is the only method
currently available to store semen for long (greater than a few days)
periods of time, when used with AI it has approximately half the fer-
tilizing life span of fresh semen. As a general rule, frozen–thawed
semen must be deposited in the uterus and at close proximity to ovu-
lation. Fresh, cooled semen, having a longer fertilizing life span than
frozen–thawed semen, can be placed with less accuracy, generally in
the true cervix or distal uterine body, and with a greater time inter-
val prior to ovulation. Thus, when attempting to develop AI, first
attempts are generally made using fresh cooled semen.
All of the successful AIs in marine mammals have been per-
formed using an endoscope to deposit either fresh or frozen–thawed
semen into the uterus (uterine body or the horn). Before this could
be accomplished, a thorough understanding of female reproductive
tract anatomy was essential. Despite the close phylogenetic relation-
ship between captive delphinids, significant variation exists in their
reproductive tract morphology. For example, bottlenose dolphins have
a pseudocervical vaginal fold or flap that must be traversed before
encountering the true cervix. Knowledge of the anatomy of the bot-
tlenose dolphin would do little to help placement of semen into the
uterine body of a closely related member of the delphinid family, the
Pacific white-sided dolphin. This species has a cervix which is com-
posed of a series of three annular folds that present anatomical barri-
ers to the placement of semen. The killer whale, another member of
the delphinid family, has a completely different arrangement consist-
ing of two longitudinally ridged cervices aligned in series.
Once the semen has been deposited and ovulation has occurred,
how do you determine if you have had success? With the applica-
tion of ultrasound for pregnancy detection (Williamson et al., 1990),
more and more marine mammal practitioners have become aware
of the fact that all species of captive delphinids can exhibit variable
periods of false pregnancies. These periods of elevated progesterone
are currently endocrinologically indistinguishable from pregnancy.
Thus, pregnancy can only be confirmed with the use of ultrasound.
In the bottlenose dolphin, the conception rate after AI using
frozen–thawed semen is 65–70%. AI in this species has been taken
a step further by the development of sperm-sorting technology, a
 182 Captive Breeding
C collection using electro-ejaculation, then release back into the wild
Figure 5 The world’s first female bottlenose dolphin (Tursiops
truncatus) born at SeaWorld San Diego (California) after using AI
with X-selected spermatozoa. Photo credit: Mike Aguilera, SeaWorld
San Diego.
sex predetermination method (Johnson et al., 1989). The ability to
skew the sex ratio through insemination of females with X chro-
mosome-bearing (female) spermatozoa helps to alleviate space
and social problems that often occur with multiple adult breeding
males. AI of sex-sorted, cryopreserved sperm could also contribute
to improved genetic management of captive populations in the same
way as conventionally frozen sperm. Further, small populations can
be replenished at a faster rate with higher numbers of females than
males. Development of this technology for application to bottlenose
dolphin population management has led to the world’s first cetacean
pregnancy using AI with sex selected then cryopreserved sperm
(O’Brien and Robeck, 2006). The technology has been integrated
into SeaWorld’s multi-site reproductive and genetic management
program for this species, with five sex-selected (female) calves being
born to date (O’Brien and Robeck, unpublished data; Fig. 5). Sex
sorting of previously frozen spermatozoa, derived from ejaculates or
post-mortem epididymal samples can also be accomplished provid-
ing species-specific modifications are developed in controlled stud-
ies (O’Brien et al., 2004, 2005). The diverse applications of sperm
sorting in conjunction with AI have great implications for the genetic
and social management of captive cetaceans, particularly those which
are endangered with extinction.
IV. Challenges for the Future
If self-sustaining populations are to be developed, then all repro-
ductively capable animals must contribute to the gene pool. Producing
viable offspring is the primary goal of any captive breeding program,
but it must be managed to avoid overpopulation of the facility and
to prevent or minimize inbreeding. This may require separate facili-
ties for adult males, preparturient females, and females with new off-
spring. Breeding may be regulated simply by separating the sexes or
by neutering (physically or chemically) both males and females.
Currently, many populations of marine mammals are kept in
small groups or same-sex groups. Therefore, a large portion of repro-
ductively mature animals within these groups are not reproducing.
These animals are functionally excluded from contributing to the
collective captive gene pool. Therefore, valuable financial resources
and pool space are being used for a minority of the available genetic
lines. This inefficient use of animal resources must be corrected
before long-term population stability can be achieved. Judicious
use of contraception and continued development of AI in all captive
marine mammal species would help managers optimize utilization of
the population’s genetic pool. However, if the maximum utilization of
genetic resources does not result in a predictable stable population,
then genetic infusions from wild stocks will be necessary. ART may
provide another answer to this future dilemma if semen (and pos-
sibly oocytes) can be collected from wild animals that are inciden-
tally or purposely killed in fisheries or subsistence hunting activities.
Alternatively, temporary capture of wild males, followed by semen
after appropriate monitoring, could also represent a means of obtain-
ing valuable genetic material for potential infusion into the captive
population using ART. As an alternative to the ART solution or until
ARTs such as AI and IVF were perfected, mangers could “borrow”
adult males from wild populations for 1–2 years for breeding pur-
poses and then return them. This, of course, involves considerable
expense and there is no guarantee that any given animal would breed
successfully.
See Also the Following Articles
Captivity ■ Genetics for Management ■ Marine Parks and Zoos
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Captivity
PETER CORKERON
I. The Debate
T
he debate over the ethics of marine mammals in captivity is
essentially about cetaceans (whales, dolphins, and porpoises)
because other marine mammals such as seals and sea lions do
not inspire the same passion as whales and dolphins do. The ongoing
debate over whether cetaceans should be kept in captivity is relatively
recent in contrast to the history of human/marine mammal interac-
tions. Human interest in marine mammals was initially based on the
commercial value of killing seals and whales for oil, meat, and hides.
The larger animals represented a greater profit, so small marine mam-
mals such as dolphins were mostly considered to be pests to fisher-
men. Occasional reports of marine mammals being kept in captivity as
curiosities are scattered throughout history: polar bears (Ursus mar-
itimus) were kept by Scandinavian rulers prior to the Middle Ages; a
killer whale (Orcinus orca) that had been live stranded was kept and
used for sport by Roman guards during the first century ad; and seals
were kept in menageries by the eighteenth century. In the mid-1800s,
P. T. Barnum displayed belugas (Delphinapterus leucas) and common
bottlenose dolphins (Tursiops truncatus) in his New York museum for
a short time, and in the late 1800s, the Brighton Aquarium in England
displayed harbor porpoises (Phocoena phocoena) for several months.
A new era of modern marine mammal exhibits began in the late 1930s
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when the Marine Studios at Marineland opened in Florida, USA
(Reeves and Mead, 1999).
At first, marine mammal facilities were quite popular. During
the 1950s, 1960s, and 1970s, the number of aquaria and zoological
parks displaying marine mammals increased rapidly to meet pub-
lic demand, especially in Europe, North America, and Australasia.
Simultaneously, technology and methods for the capture, transport,
and maintenance of marine mammals improved with increasing
knowledge and experience. Scientists took advantage of the ability
of the animals at these facilities to conduct groundbreaking research
C
on dolphin acoustics, human/dolphin communication, dolphin brain
function, hearing and echolocation, and behavior. Soon the public
became more familiar with dolphins through shows at aquaria and
from “Flipper,” a popular TV show, in which a dolphin was portrayed
as a free-ranging family friend. With this heightened awareness, the
public began to understand that dolphins were not large fish, but
intelligent and friendly marine mammals. However, the image trans-
formation inspired—at least in part by animals at marine mammal
facilities—would soon become a public relations nightmare for the
industry that helped to make these animals popular.
II. The Impact
In developed nations, the boom in aquaria and oceanaria experi-
enced through the 1970s came to a near halt during the mid-1980s
due to the growing debate over keeping cetaceans in captivity.
Pressure from non-government organizations (NGOs) and changes in
public opinion forced the closure of some existing facilities and pre-
vented some new facilities from being opened. While the 1990s saw a
decline in the number of facilities for keeping cetaceans in Australia
and some parts of Europe, but the number of captive facilities
increased in other parts of the world, notably in Asia, Latin America,
and the Caribbean, where there was little or no domestic agitation
against oceanaria.
Compared with most terrestrial mammals, marine mammals are
expensive and difficult to maintain in captivity. They require a good
deal of logistical support, such as high-quality food sources, special-
ized veterinary care, large enclosures, and expensive water-quality
maintenance systems. Cetaceans and sirenians (manatees, Trichechus
spp., and dugongs, Dugong dugon]), being wholly aquatic, present
greater logistic difficulties than any other marine mammals. A few
species predominate at these facilities because they have shown
greater success in captivity, and because they are relatively easy to
capture. Of these animals, those most often used in public perform-
ances include common or Indo-Pacific (Tursiops aduncus) bottlenose
dolphins, belugas or killer whales, and California sea lions (Zalophus
californianus), whereas phocids such as harbor seals (Phoca vitu-
lina), sea otters (Enhydra lutris), and sirenians are more typically
maintained in non-performing exhibits.
Most modern facilities in developed nations maintain high stand-
ards of operation and animal care. However, many facilities, particu-
larly those in less developed nations, fall well short of such standards
and internationally, new captive facilities are opening mostly in less
developed nations. There is concern that capture and holding facilities
that are effectively unregulated exist, particularly in southeast Asia and
parts of Latin America and the Caribbean, supplying animals to sup-
port the growth of the aquarium industry in these regions.
Even if only for short periods of time, almost every species of
marine mammal other than most of the great whales have been main-
tained in captivity. As an anomaly, two young gray whales (Eschrichtius
robustus) were kept in captivity for about 1 year each, and a wealth of