Editorial The Emergence of Conservation Behavior

Conservation have begun to apply general prin- that use visual signals to make adaptive mate-choice deci- ciples of animal behavior to solve conservation prob- sions. As a result, fish may hybridize and suffer reduced lems. Four major edited volumes and a number of re- individual and the population itself may decrease. views published in the last decade have proposed the- Some argue that ultimate questions are most impor- oretical and empirical links between behavior and con- tant for conservation because they link variation in behav- servation (see the Animal Behavior Society’s Conserva- ior with fitness. However, understanding how proximate tion Committee website: www.animalbehavior.org/ABS/ processes affect survival is also crucial. For example, we Conservation/). To strengthen this multidisciplinary col- know that training captive-reared animals to recognize laboration, it is essential for conservation biologists to predators may increase post-release survival. Yet further understand the types of questions addressed by the field studies on the developmental contexts, presence of sensi- of animal behavior (proximate and ultimate) and the im- tive periods, and types of experiences required may yield plications of these questions for , the more efficient and effective training programs. conservation literature, and recovery teams. Conserva- From a management perspective, searching for the fit- tion biologists may be unfamiliar with the jargon of be- ness consequences of behavioral responses may provide havioral biologists, but lack of knowledge is no excuse to insights into the impacts of certain human activities on ignore animal behavior. We believe that behavioral biol- , but this does not necessarily translate into con- ogists have much to contribute to conservation biology servation gains. We believe that mechanistic research and that by involving conservation behaviorists too late in will increase the predictive power of our conceptual and conservation plans or recovery teams, we will continue mathematical models. In general, by searching for mech- to suffer avoidable setbacks, waste precious funds, and anisms, we search for a deeper understanding of how lose priceless animals. animals make adaptive decisions. And by taking this sort The Nobel Prize-winning ethologist Nikko Tinbergen of uniquely behavioral approach, we may gain important first outlined two main types of questions that could information about how to better manage populations. be asked about any behavioral phenomenon. Proximate For instance, a strategy to better manage the effects of questions focus on behavior’s development or immediate tourists on wildlife is to understand the proximate mech- causation. Ultimate questions focus on behavior’s evolu- anisms underlying the response of animals to humans. We tionary history or current adaptive utility. Thus, by defini- recognize the importance of documenting population- tion, conservation behavior should be viewed as a field of level responses to human disturbance. However, to re- study that encompasses multiple levels and incorporates duce negative effects on wildlife and increase the num- avariety of disciplines, including , , ber of species that can be viewed by tourists, managers behavioral , and . must know how to control rates of human visitation to Understanding social relationships that lead to fitness protected areas. To predict use in areas with dif- (ultimate questions) has clear links to conservation. For ferent levels of human visitation, we need to determine instance, infanticide by males is an adaptive reproduc- the number of tourists per unit time and area that do not tive strategy that needs to be considered in the manage- negatively affect the frequency of resource use (e.g., feed- ment of some species, such as bears and African lions. ing, nesting, and roosting sites) by target species. In due Managers typically view males as superfluous and model course, we must identify behavioral indicators of distur- populations based on the number of breeding females. bance and develop an understanding of the behavioral However, in species where males kill offspring sired by mechanisms that explain the tolerance of some, but not other males to encourage females to come into estrus and all, species to human disturbance. It is essential to rec- mate with them, any subsequent social disruption has a ognize these qualitatively different questions along with large effect on population size. Normally adaptive repro- their applied implications when assessing the value of ductive strategies have other consequences when individ- conservation behavior. uals encounter anthropogenic changes. For instance, as Habitat loss is a huge threat to , yet habi- water quality declines, it becomes more difficult for fish tat improvements are often made without regard to how 1175

Conservation Biology, Pages 1175–1177 Volume 18, No. 5, October 2004 1176 Editorial Blumstein & Fernandez-Juricic´ species perceive the habitat (a proximate question) or the dermic mounts, and playback of sounds through speakers behavioral consequences of such intervention. For in- to study the perceptual and assessment abilities of a vari- stance, adding nest boxes (i.e., increasing nesting habitat) ety of species. Through the use of simple stimuli, these is often done to increase reproductive success for cavity- techniques allow one to learn about how animals assess nesting birds. If the artificial nest boxes are too close risk. together, however, there may be unintended social con- Although managers must make tough decisions about sequences. Clumped nest boxes lead to an increased rate what can be done with animals prior to release, they must of egg dumping and conspecific brood parasitism, both of think critically about how much they can learn from these which reduce reproductive success. Hence, without at- studies to improve success rates. The recovery programs tention to behavioral details, such well-intentioned habi- forblack-footed ferrets (Mustela nigripes) and golden lion tat “improvement” programs may waste valuable funds tamarins (Leontopithecus rosalia)provide evidence that and have detrimental effects on the population under fully integrating behavioral biologists into the recovery recovery. process can make a difference. In both cases, paying atten- To develop more effective programs, conservation biol- tion to the environment these species were reared in and ogists must fully recognize the value of integrating prox- providing pre-release behavioral enrichment was essen- imate and ultimate behavioral studies. In recent years, tial for ultimate success. In the long run, it may be more this lack of integration has generated resistance toward cost-effective to incorporate behavioral biologists at the increasing the diversity of the conservation behavior lit- outset of and reintroduction programs erature and has constrained the use of behavioral data by rather than to wait until a “problem” emerges, animals recovery teams. die, and precious funds are spent. Based on our experiences and those of many of our Conservation behavior will not provide the solutions colleagues, it is difficult to publish papers on conserva- to all conservation problems, but we conservation behav- tion behavior, a problem not restricted to any particular iorists need to show how useful a behavioral perspective journal. When manuscripts on conservation behavior are can be. By understanding the type of questions conserva- submitted to conservation journals, negative reviews may tion behavior can address, one can better evaluate how highlight a lack of immediate applicability or a lack of a di- and when could benefit from behav- rect link between behavioral responses and fitness (e.g., ioral knowledge. Addressing both proximate and ultimate reproductive output, survival rates). Yet general questions questions of conservation behavior is essential to improv- about proximate behaviors that affect survival, mortality, ing our management decisions. Two research areas in par- or population size are all topics that are explicitly relevant ticular will not only help bridge the gulf between conser- to conservation biology. vationists and behaviorists but will also help research on When conservation behavior manuscripts are submit- conservation behavior to produce specific conservation ted to behavior journals, they may be rejected because of outcomes. an apparent lack of theoretical framework, or, if they are First, we must develop predictive models of behavior accepted, authors are asked to remove most references that are accessible to wildlife managers. Advances have to conservation. However, conservation behavior itself been made in theory and modeling habitat selec- can be theoretical. For example, predicting the behav- tion, wherein animal decisions have been modeled with ioral mechanisms underlying the vulnerability of a species increased sophistication. However, the sophistication to human impacts and predicting how long antipredator that comes with deeper understanding may impede ap- behaviors persist following isolation from predators are plication by its complexity. Behavioral biologists should behavioral considerations that are explicitly, theoretically strive to make their models accessible to others by identi- motivated. fying specific behavioral mechanisms. These mechanisms Afar greater threat to species survival than the diffi- should be the basis of bottom-up models that predict culties of publishing conservation behavioral findings is the behavior, movement, habitat use, and distribution of the tendency of recovery teams to wait too long to inte- species of conservation concern. grate behavioral biologists into their efforts. Such over- Second, we must develop an ultimate understanding of sights appear to be related to a lack of understanding of how and why species are vulnerable to human impacts. how behavioral studies are conducted. For instance, an Study of conservation behavior is much more than simply exciting and promising line of conservation behavior re- documenting whether or not humans affect wildlife. The search seeks to increase reintroduction success by eval- explosion in comparative biology and macroecology in uating the antipredator abilities of animals prior to re- the past decade offers a framework for future advances lease and then following their fates. If individuals with in predictive evolutionary models. Future studies should inferior antipredator behavior are preyed upon more examine how variation in life history and natural history frequently than those with superior abilities, then pre- explains variation in proneness to , tolerance release training may be used to prepare animals for re- to human disturbance, and tolerance to invasive species lease. For years, ethologists have used simple models, taxi- at local scales.

Conservation Biology Volume 18, No. 5, October 2004 Blumstein & Fernandez-Juricic´ Editorial 1177

Ignoring the diversity and methods of modern behav- that the exciting integrative field of conservation behav- ioral biology places conservation-dependent animals at ior will prosper. peril. Although we are excited that some wildlife man- agers and conservation biologists appreciate the exper- tise offered by behavioral biologists, we believe that Daniel T. Blumstein many still fail to appreciate the importance of proximate Department of Ecology and Evolutionary Biology, 621 Young Drive mechanisms. Some pigeonhole behavioral research as too South, University of California, Los Angeles, CA 90095–1606, U.S.A., species-specific and fail to recognize that behavioral biol- email [email protected] ogists can produce generalizable and predictive models. With an increased appreciation of what conservation be- Esteban Fernandez-Juricic´ haviorists have to offer, and a fundamental integration of Department of Biological Sciences, Peterson Hall 1-109, California State them into recovery teams, we believe that future recov- University–Long Beach, 1250 Bellflower Boulevard, Long Beach, CA ery efforts will be more successful and cost-effective, and 90840, U.S.A. email [email protected]

Conservation Biology Volume 18, No. 5, October 2004