Variation in nature: its implications for zoo elephant management

Zoo Biology 25:161–171 (2006)

Commentary

Variation in Nature: Its Implicationsfor Zoo Elephant ManagementMichael Hutchins1–3�

1The Wildlife Society, Bethesda, Maryland2Center for Conservation and Behavior, Georgia Institute of Technology,Atlanta, Georgia3Graduate Program in Conservation Biology and Sustainable Development,University of Maryland, Bethesda, Maryland

Despite many advances in animal care and welfare over the past few decades, zooshave been criticized recently for the quality of their elephant managementprograms. More specifically, critics have argued that elephants live miserable livesin captivity and thus should not be kept in zoos. Poor health and reproductivesuccess, they say, are the result of the combined impact of: a lack of exercise;exposure to cold temperatures and disease; and stress due to the use of ‘‘brutal’’training techniques, chaining and inappropriate social environments. Everyone,including zoo professionals, seems to agree that improvements in zoo elephantmanagement are necessary and appropriate. However, there is considerabledisagreement on how to proceed and little information on which to base suchdecisions. One tactic that the critics have adopted in their efforts to promotechange is their frequent reference to ‘‘nature’’ as a yardstick for gauging theadequacy of zoo animal management and care. An argument is made that directzoo–wild comparisons are of questionable utility and may be invalid from ascientific perspective. Some critics talk about ‘‘nature’’ as if it represented a fixedset of rules by which captive managers must either abide by or risk diminishingthe health and welfare of their charges. However, many aspects of elephantnatural history vary greatly depending on prevailing environmental conditionsand elephants may be much more flexible than many critics are giving them creditfor. Thus, although zoo animal managers should look to nature for clues abouthow to best care for captive elephants, they should not feel constrained by them.This revelation is not intended as an excuse for poor elephant management or tosupport the status quo. On the contrary, a better approach is to develop realisticand biologically meaningful metrics that reflect the quality of elephant care and

Published online 15 March 2006 in Wiley InterScience (www.interscience.wiley.com).

DOI 10.1002/zoo.20087

Received 2 May 2005; Accepted 29 July 2005

�Correspondence to: Michael Hutchins, PhD, The Wildlife Society, 5410 Grosvenor Lane, Bethesda,

MD 20814-2144. E-mail: [email protected]

rr 2006 Wiley-Liss, Inc.

welfare and to use them to measure the success of evolving zoo elephantprograms. Zoo Biol 25:161–171, 2006. �c 2006 Wiley-Liss, Inc.

Keywords: elephants; zoos; variation in nature; captive management; animal care; animalwelfare; flexibility; metrics

‘‘Toiling in the treadmills of life we hide from the lessons of nature. We gazemorbidly through civilized fog upon our beautiful world.’’

John Muir (cited in Foster, 1993)

INTRODUCTION

Zoo exhibit design and animal care have advanced considerably over the pastfew decades. Contemporary animal exhibits tend to be comparatively larger andmore naturalistic (containing natural plantings, water features, rockwork, etc.);animal health is carefully monitored by well-trained veterinarians and enhancedusing advanced medical technologies; diets are formulated by expert nutritionists;there is an increased appreciation of behavioral considerations in animal manage-ment; and there is, no doubt, a heightened interest in and concern for animal welfare[Kleiman et al., 1996; Shepherdson et al., 1998; Hutchins and Smith, 2003]. As aresult, individuals of many species are living longer in modern zoos, behave morenormally and are generally perceived to have a better quality of life than they did inthe small, sterile, concrete and barred cages of the past.

The future of zoos and zoo animals would seem to be golden. However, zoosare undergoing a continual process of renewal and not all exhibits or species arecreated equal. In addition, some zoos are more advanced in their redevelopmentthan others; most retain at least a few older, inadequate exhibits they would prefer tosee demolished and even newer exhibits can become rapidly outdated as newapproaches to animal exhibition become available. Similarly, some species are doingbetter in zoos than others. Whereas some taxa are thriving, others continue toexperience a comparatively high incidence of mortality, disease, behavioralabnormalities and poor reproduction, evidence that their health and welfare mightbe compromised by captivity. Thus, the quality of animal care and zooredevelopment are closely linked, but this generates some fascinating questions forzoo professionals as they strive to improve: what standards of animal health andwelfare will be used to evaluate the adequacy of existing and new animal facilitiesand care programs? Because it is impossible for zoos to replicate nature precisely,how far do they need to go to provide acceptable levels of animal care? Furthermore,whose perception of ‘‘acceptable’’ will zoos adopt? To examine these importantquestions, elephants are used as an example.

ZOO ELEPHANT MANAGEMENT: THE CRITICS VIEW

Zoos have been facing growing criticism of their elephant managementprograms. More specifically, critics have argued that elephants live miserable lives incaptivity and thus should not be kept in zoos. Poor health and reproductive success,they say, are the result of the combined impact of: a lack of exercise; exposure to cold

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Zoo Biology DOI 10.1002/zoo

temperatures and disease; and stress due to the use of brutal training techniques, chainingand inappropriate social environments [Clubb and Mason, 2002; Farinato, 2004; Peoplefor the Ethical Treatment of Animals, 2005]. The finer points of zoo elephantmanagement will not be debated here. Some excellent programs undoubtedly exist(e.g., Disney’s Animal Kingdom, San Diego Wild Animal Park, Oakland Zoo, NorthCarolina Zoo), but there is broad consensus, from both within and outside theprofession, that zoos, in general, must improve their elephant programs or otherwisecease exhibiting these animals altogether [American Zoo and Aquarium Association,2001; Hutchins et al., in press]. Although these are important questions for contemporaryzoo managers, this article is focused on a recent strategy adopted by zoo critics intheir attempts to impose change and, more specifically, on the fundamental conceptbehind it. Of particular interest is their frequent reference to ‘‘nature’’ as a yardstickfor gauging the adequacy of zoo animal management and care. Some examples include:

‘‘yelephants in the wild can roam up to 50 miles a day.’’ ‘‘Elephants don’tbelong in urban zoo environments because they don’t have adequate space’’ [Smithcited in Vickroy, 2005].

‘‘A preserve of at least 2 square miles-1,280 acresywould be necessary to meetan elephant’s physical and social needs.’’ [Schobert, 2005].

‘‘A natural lifespan for an elephant is 60–70 years. In zoos they dieprematurely, on average at 34 years of age.’’ [In Defense of Animals, 2004].

‘‘You certainly don’t find elephants in the Arctic in the wild.’’ [Meyer cited inKershaw, 2005].

‘I doubt that there are many elephants in the wild that just fall down and die ofheart attacks.’’ [Katz cited in Anon, 2004b].

‘‘Wild female elephants don’t conceive until they are 18. The early breedingage in zoos puts mothers and babies at higher risk of death and illness.’’ [Clubb andMason cited in Anon, 2004a].

‘‘ydocumentation on wild elephants shows that their lives are characterizedby great physical activity on a daily basis, by life long bonds between stable femalegroups, by sophisticated communication, by a large behavioral repertoire, andby successful reproduction into the later years of their 60-plus year life span.’’[Farinato, 2004].

‘‘Just as polar bears don’t thrive in hot climates, Asian elephants shouldn’t livein small groups without many acres to roam. They clearly shouldn’t have to sufferwinters of the north’’ [Kagan cited in Detroit Zoological Institute, 2004].

ZOO–WILD COMPARISONS AND NATURAL VARIATION

Zoo professionals themselves have long argued that we must look to natureto find solutions to animal management challenges. Certainly, an understanding of

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a species’ basic biology, including its feeding ecology and diet, social organizationand behavior, reproductive biology and so forth are critical for developing successfulmethods of captive animal husbandry and care [Hediger, 1969; Kleiman et al., 1996;Hutchins, 2003]. However, making direct comparisons between wild and zoo animalsmay be a slippery slope, at least from a scientific perspective. The reason is the degreeof variation that exists in natural systems. Indeed, people’s perceptions of natureoften differ greatly from reality, perhaps because we are constantly trying to simplifyour complex world in an effort to better understand it. Yet, the range of naturalvariation is often astounding. I learned this lesson while studying primate anatomyin college. The textbook representations of arteries and nerves for a particularspecies were frequently incorrect when compared to those actually observed duringdissection. In some individuals, a branch could be missing altogether and, in others,there might be two or three branches instead of the expected one. The same is trueof many aspects of animal behavior and ecology. For example, variations in habitatquality are known to have significant impacts on animal behavior and sociality[Alcock, 2005]. Among marsh-nesting black-birds, for instance, the quality of amale’s territory (i.e., presence of cover, nesting sites and food) determines whatmating system he will adopt, that is, whether he will be monogamous or highlypolygynous [Orians, 1980]. Likewise, the distribution of resources in a particularhabitat can greatly affect animal movement patterns, range size and social behavior.When resources, such as food or mates, are more concentrated, animals do not haveto move as far to find them as they do when they are widely dispersed [Alcock, 2005].This, in turn may lead to higher rates of competition and aggression. Similarly,individuals of the same species, living in poor quality habitats (i.e., those with lessnutritious food, more competitors) typically have slower growth rates, higher ratesof mortality, reduced reproductive success and are less active when compared tothose living in high quality habitats (i.e., those with more nutritious food, lesscompetition). Geist [1978] has referred to these environmentally-induced variationsas ‘‘phenotypic syndromes.’’

Habitat quality can also affect reproductive strategies or parent–offspringrelationships. For example, uncertain environmental conditions may cause a femalemammal to fail to conceive or abort or resorb her fetus, thus resulting in theearly termination of parental investment [Bernds and Barash, 1979]. Similarly,environmental conditions can affect both the length and quality of parent-offspringrelationships post-partum. For example, Hutchins [1984] found that femalemountain goats (Oreamnos americanus) typically weaned their yearlings at thebirth of a new infant. However, when females failed to reproduce in consecutiveyears, presumably due to resource limitations, they continued to display a close,protective relationship toward their yearling offspring. Seasonal changes are anotherimportant source of environmental variation. For example, most people think ofmany cold-adapted mammals, such as mountain goats or polar bears (Ursus arctos)as living perpetually among snow and ice; however, during the summer monthsor in their southern-most ranges, these species can be subjected to verywarm temperatures and roam through green, vegetated habitats [Perry, 1966;Chadwick, 1983].

Many wild animals display remarkable flexibility in their behavior andtolerance for a wide range of social and environmental conditions. Although thisrange of flexibility certainly has its limits and may vary from species to species,

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nonetheless, it undoubtedly exists. If we want to understand what is ‘‘normal’’ fora particular species, then we cannot exclude the range of natural variation. Thus,if we are going to draw comparisons between wild and zoo animals, which wildpopulation or habitat, or time of the year, should we pick? What yardstick shouldwe use? Should it be the wild population living under ideal conditions, the one livingunder marginal or extreme conditions, or should we consider the average? The pointis that there is no ‘‘one nature’’ for a particular species, no simplified categorizationof the natural world that we can use as a model for either developing or evaluatingthe quality of zoo animal management programs. This is especially true for wideranging and widely distributed species, such as elephants. The real question thenbecomes what range of conditions is a particular species able to tolerate and stillfall within an acceptable range of possible welfare states (i.e., the well-beingof individuals in the population in question)?

How can we assess the range of conditions in which zoo elephants canbe expected thrive? How can this knowledge be used to plan for the future? Theseare critical questions facing the professional zoo community today. For example, weoften hear that wild elephants move up to 50 miles a day. Consequently, some critics,including some zoo professionals, have concluded that no urban zoo can everprovide the kind of space necessary to meet the basic needs of elephants in captivity[Stephens, 2004]. However, the range of daily movements by wild elephants variesconsiderably, depending on a number of factors, including the distribution of criticalresources (food, water, mates) and seasonal changes in climate (e.g., wet vs. drought)[Sukumar, 2003]. In fact, most mammals are energetically conservative and do notmove just for the sake of moving; wild lions, for example typically spend 90% oftheir time inactive [Schaller, 1972]. Although I believe that multi-acre habitats area must for future zoo elephant exhibits, the critical question becomes how muchspace is enough? The San Francisco Zoo was recently forced to close its elephanthabitat, based primarily on space considerations. Whereas closure of the outdatedexhibit was a responsible decision, the City Board of Supervisors also passedan ordinance preventing the zoo from having elephants in the future unless at least15 acres could be provided [Hampton, 2004]. However, oddly enough, the Board didnot specify the number of elephants that should be held at the facility, thus callinginto question the wisdom of having politicians make animal management decisionsat zoos. In addition, proponents of elephant sanctuaries, which have hundredsof acres for elephants to roam, claim that they are a better alternative than zoos[The Elephant Sanctuary, 2005]. Yet, elephants at smaller, but multi-acre facilities,such as Disney’s Animal Kingdom and the San Diego Wild Animal Park, seem to bedoing just fine based on any objective measure of welfare one chooses, includingsuccessful reproduction, activity patterns, lack of stereotypic behavior, and animalhealth and mortality [J. Lenhardt and R. Rieches, personal communication]. Onecould reasonably conclude therefore that the range of acceptable enclosure sizesfor elephants might lie in the 3–7-acre range, and not necessarily in the 15-acre plusrange. However, precise estimates are going to be difficult until more informationbecomes available. Objective measures that might help refine estimates of acceptableenclosure size include: incidence of foot problems [Csuti et al., 2001] and observedactivity levels, particularly locomotion, which is indicative of the amount of exerciseanimals are getting. Of course, the size of the enclosure will also be dependenton how many animals are projected to be in the herd. Furthermore, size is not



everything; the complexity of an enclosure and the care program provided(e.g., feeding strategies) are also important [Stoinski et al., 2000].

Weather is another sticking point. Critics have suggested that keepingelephants in cold climates is unnatural. Zoos in northern climates must keep theirelephants indoors much of the winter, they say, resulting in arthritis and footproblems that come from standing on cold, concrete floors [Clubb and Mason,2002]. But this conclusion is largely conjectural and based on anecdotal evidence.Wild Asian elephants are typically found in or at the periphery of lower elevations intropical and sub-tropical forest and are usually not subjected to cold temperatures.However, African elephants are another story; they are found at high elevations orin desert habitats, where temperatures can drop below freezing [Wells, 1929]. Andwho can forget mammoths and mastadons? Certainly, elephants, like many otherspecies kept in zoos, require shelter from severe weather, especially during the coldestwinter months. However, some of the most successful zoo elephant programsare found in northern climes (e.g., Syracuse, NY, Portland, OR, Columbus, OH,and Indianapolis, IN), suggesting that elephants may actually tolerate a muchwider range of temperatures than zoo critics give them credit for. Depending howlong elephants are expected to stay indoors during the winter, northern latitude zoos(and sanctuaries) probably need to construct larger indoor holding facilities toprovide elephants with improved opportunities for social interaction and exercise.Furthermore, larger outdoor facilities with varied terrain, combined with appro-priate feeding strategies and social enrichment, should give the animals ampleopportunity for movement and exploration during the vast majority of the year.An ability to walk on natural abrasive surfaces is important for foot health andregular exercise helps to lower weight and maintain strength and flexibility[Csuti et al., 2001]. As mentioned earlier, zoos will need to develop simple metricsfor measuring the success of such enclosures, including distances travelled per dayand the incidence of foot problems or need for foot trimming.

With regard to group size and composition, I agree that adult female elephantsneed to be maintained in larger groups; keeping them as singles or even pairs issimply not acceptable for these highly social animals (American Zoo and AquariumAssociation, 2001]. But how many is enough? Herd size in wild elephants varieswidely. The largest groups have been recorded in east African savannahs, wheremean group sizes of over 10 are common [Sukumar, 2003]. African and Asianelephant groups living in woodlands and dry forest, respectively, typically rangefrom 5–10 individuals. However, much smaller mean group sizes of less than fiveindividuals are the norm in African rain forests. Furthermore, group size can varywith season, with the largest groups being observed during the rainy season andsmallest during the dry [Sukumar, 2003]. This suggests that group size and stabilityare also influenced by environmental factors, such as food availability anddispersion. The current American Zoo and Aquarium Association standards callfor a minimum of three adult female elephants (and their offspring) to be housedtogether in the same facility [American Zoo and Aquarium Association, 2001].Sanctuaries claim that five should be the minimum [The Elephant Sanctuary, 2005],but some zoo critics have suggested that the number should be even higher. Onceagain, determining an optimum group size based on ‘‘nature’’ will be difficult, aswild elephants seem to be extremely flexible in this regard and have the ability toadapt to prevailing conditions. Group compatibility might be a more important

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factor than group size per se. An elephant maintained in a smaller, but compatiblegroup may, in fact, be better off than one in a larger, incompatible group.

On a related topic, critics have also suggested that wild female elephants arefound in large, highly stable and closely related matrilineal herds, whereas captivegroups tend to be composed of unrelated individuals that are frequently movedaround for the purposes of management and breeding [Clubb and Mason, 2002].Captive groups do tend be composed of unrelated individuals; but, stable captivegroups have been formed successfully from a nucleus of unrelated individuals [e.g.,Disney’s Animal Kingdom: J. Lenhardt, personal communication; Oregon Zoo:M. Keele, personal communication]. In fact, there is also great variation in thestability and relatedness of elephant groups across various habitats in nature[Sukumar, 2003]; in some populations the only stable unit observed was that of anadult female and her immature offspring, whereas in others, the layers of socialrelationships among adult females are complex and fairly stable [Whittemyer et al.,2005]. Of course, regardless of group size and composition, the bottom line shouldbe how the animals are behaving. If zoo elephants are exhibiting a broad range ofnatural social behaviors and not showing high levels of agitation, aggression,lethargy, or stereotypic movements, then one might reasonably conclude their socialneeds were being met.

A related question concerns the presence of young elephants in captive groups.Critics have suggested that zoos should not breed elephants as captivity isdetrimental to their welfare [The Elephant Sanctuary, 2005]. However, as Clubband Mason [2002] point out, the absence of animals in younger age classes is nottypical and therefore may diminish welfare. Most, although not all, free-rangingherds of adult female elephants contain young animals at various stages ofdevelopment [Sukumar, 2003]. Young elephants are a focus of activity for the groupand are, by themselves, an important source of social cohesion, activity andenrichment [Moss, 1988]. In addition, the presence of infants is what gives youngfemales an opportunity to learn to develop their maternal skills. The absence ofinfants in zoos has created an entire generation of females that has little or no care-giving experience and, as was the case for lowland gorillas (Gorilla g. gorilla) in the1970s and 80s, this could be negatively impacting overall reproductive success [Beckand Power, 1988]. The formation of larger, more stable breeding female herds could,therefore, begin to create conditions conducive to natural rearing. Furthermore,there is some indication that delaying female reproduction may contribute to thedevelopment of reproductive pathologies, which can interfere with successfulbreeding later in life [Brown et al., 2004]. Clearly, zoos need to breed femaleelephants earlier and more consistently and they are trying. However, zoo critics’ callfor a cessation of breeding by zoos is inconsistent with the concept that ‘‘nature’’should be the yardstick by which the adequacy of zoos (or sanctuaries) is measured.

Critics have also complained about the time that young are allowed to spendwith their mothers in zoos [Clubb and Mason, 2002]. Data available from one wildpopulation (Amboseili) suggest that offspring remain within 5 meters of theirmothers up to 8 years of age [Moss, 1988]. Cessation of nursing usually occurred onthe birth of a new infant (around 3 years). In some cases, both the young of the yearand older offspring were seen nursing simultaneously. However, inter-birth intervalsare known to vary between different populations, probably due to environmentalfactors [Sukumar, 2003]. The current American Zoo and Aquarium Association



standards state that young must remain with their mothers for a minimum of threeyears [American Zoo and Aquarium Association, 2001]; the European Associationof Zoos and Aquaria (EAZA) recommends 4–5 years [European Association of Zoosand Aquaria, 2004]. Once again, a precise estimate of an optimal time of removalwould be difficult, because it is quite possible that elephants may be flexible inthis regard too. If zoos wanted to err on the side of caution, however, they mightconsider keeping female offspring with their mothers indefinitely and removingmales when they are 5–8 years old. When zoo elephant groups must be split upto reduce population size, then perhaps entire matrilines (i.e., females and theirfemale offspring) could be moved together as opposed to a single individual[Hutchins et al., in press].

Also in dispute is the age at which adult female elephants should first be bred.Clubb and Mason [2002] state that zoo elephants are allowed to breed at youngerages than in the wild, 15 years on average, as opposed to ‘‘18–20 years’’ for wildfemales. They further contend that early breeding ‘‘may have serious consequences,’’including early reproductive senescence and increased mortality. Again, this is highlyspeculative, especially given the fact that age at first reproduction is highly variablein wild elephants and probably related to habitat quality; there are even reports ofAsian cows maturing as early as 7–9 years of age. The mean age of first calving insouthern India was around 17–18 years and age of first conception at 15–16 years.Mean age of sexual maturity in African cows is from 11–14 years, but can range from9–22 years [Sukumar, 2003]. Thus, the range of variation in wild populationsoverlaps that seen in captivity.

Finally, critics have suggested that the life spans of zoo elephants areconsiderably shorter than those of their wild counterparts. I will not spend muchtime on this topic here, as it is aptly covered by Wiese and Willis [2004]. Suffice to saythat their results contradicted an earlier, well-publicized study by Clubb and Mason[2002], which concluded that the average longevity of zoo elephants was significantlyshorter than that of wild populations for which data were available. As it turnsout, Clubb and Mason based their analysis of zoo elephant mortality exclusivelyon animals that had died and not on the entire population, both living and dead.This resulted in their greatly underestimating average life spans. Wiese and Willisalso pointed out that zoo critics often use maximal values to characterize the lifespanof elephants (e.g., 70 years), rather than averages (see above). Of course, this ispatently unfair and scientifically invalid, because such individuals represent extremecases (i.e., they are statistical outliers). How many humans live to be 120 years old?Not many. Indeed, contrary to popular belief, wild elephants die for many of thesame reasons that captive elephants do, including herpes viruses, tuberculosis andarterial disease [Sukumar, 2003]. The incidence of disease and mortality also variesby habitat and population. For example, Sikes [1968] found that arteriosclerosiswas much more common in wild African elephants that lived in habitats withnutritionally poor vegetation (it seems that, despite the protestations of critics, heartattacks may be well within the realm of possibility for wild elephants). The point isthat comparisons of the average life spans of wild and captive animals are fraughtwith many difficulties, again, related to the range of variation in nature, which canbe considerable. More specifically, which wild population should we choose to bethe focus of our comparisons with zoos? Elephants in high quality habitats are likelyto be in better physiological condition, have higher rates of survivability at all

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demographic stages, and live longer than elephants in low quality habitats [Sukumar,2003]. Similarly, should we choose to make our comparisons during times of droughtor just after the rainy season, when food is plentiful and nutritious? I do not bringthis up to argue that zoos are doing the best they can for elephants, as thereis obviously great room for improvement. This does illustrate, however, thelimitations of broad-scale zoo–wild comparisons.

CONCLUSIONS

Zoo professionals should look to nature for clues about how to best carefor captive animals, but they should not feel constrained by them [See Veasey et al.,1996]. Many zoo critics talk about ‘‘nature’’ as if it represented a fixed set of rules bywhich captive managers must either abide by or risk diminishing the health andwelfare of their charges. Yet, perfect replication of nature in zoos is not onlyimpossible, but frequently undesirable. For example, zoos are not in the business ofreplicating major die-offs due to floods, droughts, disease, parasites, starvation, andpredation. In addition, there is considerable variation in nature and many animalspecies are flexible in their ability to adapt to or tolerate a wide range of conditionsand circumstances. However, this degree of flexibility/tolerance is not unlimited andmay also vary between species. Consequently, zoo professionals should look at allthe information at their disposal to make the best possible decisions. It is a given thatprofessionally managed zoos should continually strive to improve animal care andwelfare; however, as they look to the future, zoos must also have reasonable andrealistic goals to strive for. In the case of elephants, several challenges remain and thetension of debate may ultimately prove beneficial. As zoos look to improve elephantcare, it is important that they study and learn from both their successes and failures.In addition, they should establish carefully designed metrics to help measuretheir progress; such metrics should be compatible with existing constraints and easyto collect and verify. Most importantly, zoo administrators and animal managersshould not be distracted by simplistic zoo–wild comparisons, as the underlyingconcept behind this thinking is fatally flawed. The world is actually much morecomplicated and, when it comes to successful animal management, the devil is in thedetails, not in the generalizations.

There are those who will argue that modern zoos should be required toreplicate the very best (i.e., optimal) conditions for animals in nature and thatevery individual should live to the maximum life span for its species. However, this isunrealistic. Modern zoos should not be in the business of setting longevity records;rather, they should take the best possible care they can of individual animals for theduration of their lives, no matter how long that may be. Regardless of what is doneto improve conditions for captive elephants, some individuals will live longer thanothers, as is the case whether in nature or captivity [Hutchins, in press]. If zoos are toadopt any fair measure of success, then it must be based on averages and standarddeviations and not on extremes. This is the essence of variation in nature, a pointthat zoo critics either fail to understand or purposely ignore to push their narrowpolitical agenda. In media debates, they continue to cite extreme optimal conditionsto make their case for banning elephants from zoos. Although dissent is important indemocratic societies, any attempt to support one’s argument with science is fair gamefor analysis and criticism. Zoos must continue to improve their elephant programs,



but the way ahead must take into account the range of variation that exists in natureand not be based on often biased and simplistic generalizations of what is and is not‘‘natural.’’ Keeping captive elephants in situations that replicate extreme marginalenvironments is clearly not acceptable, especially if it causes animals to suffer.Conversely, replicating extreme optimal environments may not be realistic. Zoosshould always strive for improvement in their animal care programs, but will likelyend up somewhere in the middle. The difficult issue will be about where to drawthe line between ‘‘acceptable’’ and ‘‘unacceptable.’’ This is an ethical, rather thana scientific, debate and it should be recognized as such.

ACKNOWLEDGMENTS

M. Keele, Assistant Director, Oregon Zoo and AZA Elephant TAG/SSP Chairreviewed and commented on an earlier version of this manuscript.

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Variation in nature: its implications for zoo elephant management

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