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Integrating Behaviour Into Wildlife Conservation: the Multiple Ways That Behaviour Can Reduce Ne Laura L

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Integrating Behaviour Into Wildlife Conservation: the Multiple Ways That Behaviour Can Reduce Ne Laura L Biological Conservation 95 (2000) 303±315 www.elsevier.com/locate/biocon Integrating behaviour into wildlife conservation: the multiple ways that behaviour can reduce Ne Laura L. Anthony a, Daniel T. Blumstein a,b,c,* aDepartment of Biological Sciences, Macquarie University, Sydney, NSW 2109, Australia bDepartment of Psychology, Macquarie University, Sydney, NSW 2109, Australia cThe Cooperative Research Centre for the Conservation and Management of Marsupials, Macquarie University, Sydney, NSW 2109, Australia Received 10 June 1999; received in revised form 19 August 1999; accepted 16 October 1999 Abstract There has been a recent interest in integrating an understanding of behaviour into conservation biology. Unfortunately, there has been no paradigm for such a process. Without a clear framework for integration, conservation biologists may have diculties recognising how behavioural knowledge can help solve real-world conservation problems. Eective population size (Ne) is a key demographic parameter used to understand population viability. A variety of behaviours and behavioural traits impact Ne, yet their importance for conservation is under-appreciated. We suggest that identifying behavioural traits that aect Ne provides a paradigm for integrating behavioural biology into conservation biology. Behaviour can aect Ne through at least three dierent mechanisms: reducing N Ð the population size; reducing r Ð the population growth rate, and/or by increasing reproductive skew. We discuss how nine common behavioural traits can reduce Ne, and suggest how an understanding of these traits may inform management of both free-living and captive animals. # 2000 Elsevier Science Ltd. All rights reserved. Keywords: Behaviour and conservation; Eective population size; Population viability 1. Introduction knowledge of animal behaviour to conservation pro- blems, there is no clear framework to help conservation Conservation biology is a crisis discipline aimed at biologists identify the speci®c cases when they should be saving biodiversity (Soule , 1986). A common and concerned about behaviour, nor which behaviours they important approach to saving biodiversity has been to should be concerned about. preserve patches of habitat in order to maintain ecosystem- The number of individuals in a population, N,isa level processes which, in turn, preserves populations of ®rst approximation of endangerment. However, other species (Franklin, 1993). With these broad objectives and factors in¯uence the likelihood of a population going methods, conservation biologists have paid relatively extinct over time. For instance, variation in the number little attention to how an individual animal's behaviour of breeding individuals, variation in breeding success, can help save biodiversity. Recently, a number of beha- the ratio of breeding males to females, as well as other vioural biologists have written reviews and book chapters factors in¯uence the maintenance of genetic variation in on the role of behaviour in conservation arguing that a a population (Falconer, 1989). Genetic variation in¯u- fundamental understanding of behavioural processes ences long term sustainability because genetic variation can contribute to conservation biology (Caro and Durant, is required to combat any negative eects of inbreeding, 1995; Curio, 1996; HoÈ glund, 1996; Lima and Zollner, and to allow evolutionary adaptation to an ever-changing 1996; Ulfstrand, 1996; Clemmons and Buchholz, 1997; environment. To compensate for the inadequacy of N Strier, 1997; Caro, 1998a,b; Sutherland, 1998). Despite alone in predicting the likelihood of a population per- this recognition that it may be important to apply sisting over time, population geneticist's have developed the concept of the eective population size, Ne which better re¯ects the likelihood of a population persisting * Corresponding author. Tel.: +61-2-9850-9440; fax: +61-2-9850- 9231. over time (Gilpin and Soule , 1986). Ne is an estimate of E-mail address: [email protected] (D.T. Blumstein). the theoretical number of breeding individuals assuming 0006-3207/00/$ - see front matter # 2000 Elsevier Science Ltd. All rights reserved. PII: S0006-3207(00)00037-9 304 L.L. Anthony, D.T. Blumstein / Biological Conservation 95 (2000) 303±315 they behave in an ideal way. Ne models an ideal popu- From this we clearly see that Ne decreases by either lation with the following properties: the population is decreasing the number of breeding individuals or by split into sub-populations where there is no migration skewing breeding sex ratios. between sub-populations, generations do not overlap, Population viability analysis (PVA) is an important the number of breeding individuals is the same for all tool for managers because it provides an estimate of generations and sub-populations, mating is at random the viability and sustainability of a population. PVAs and includes a random amount of self-fertilisation, there model the eect of certain biotic (fecundity, age of is no selection, and mutation is assumed to be unim- senescence) and abiotic factors (habitat availability) portant (Falconer, 1989). Ne aects population viability on Ne. It is through the calculation of Ne that a PVA by increasing homozygosity and decreasing the number predicts population persistence. Ne is therefore a cen- of non-selected alleles. The loss of variation is com- tral parameter determining population viability in pounded by an increase in linkage disequilibrium Ð PVAs. nonindependent assortment of alleles Ð which reduces As we will discuss in a number of following examples, the frequency of novel gene combinations. Ne is in¯u- individual behavioural strategies in¯uence how individuals enced by factors that halt the passing of gametes to the respond to habitat modi®cation, hunting, fragmenta- next generation. Falconer (1989) identi®ed six factors tion, corridor construction, reduced resource quality, which in¯uence Ne: (i) exclusion of closely related matings, and resource ¯uctuation. Understanding factors that (ii) skewed sex ratios, (iii) unequal generation size, (iv) in¯uence behaviour over short time scales provides vital unequal family size, (v) inbreeding, and (vi) overlapping information for those developing more accurate popu- generations. lation models as well as for those charged with managing A number of behavioural traits either directly or populations. It is therefore surprising that most PVA indirectly in¯uence Ne by changing demographic para- models ignore behavioural variation (Derrickson et al., meters that contribute to Ne. We de®ne behaviour 1998). broadly and recognise a hierarchy that begins with the Ne.s central importance in PVA models suggests that neurobiological, genetic, and physiological processes the best strategy for integrating behaviour into con- that underlie observed motor patterns, includes the servation will involve identifying behaviours and beha- functional integration of those motor patterns into vioural traits that impact Ne. Ne has already been behaviours, as well as the integration of behaviours into recognised as one of a series of ways in which beha- behavioural traits. For instance, infanticide Ð a complex vioural ecology can contribute to conservation biology behavioural trait where adults kill young of their own (Parker and Waite, 1997; Caro, 1998a). Our approach species Ð directly in¯uences Ne by reducing the popu- diers in that we view that identifying and modeling the lation size, while another behavioural trait, reproductive ways in which behaviour in¯uences Ne as the perhaps suppression of adults, reduces the number of breeding the single most important way in which knowledge of individuals and may decrease a population's rate of animal behaviour can contribute to wildlife conserva- increase Ð r Ð therefore reducing Ne. To integrate tion. behaviour into wildlife conservation, we must under- When dealing with threatened populations, Ne is stand how behaviour skews the operational sex ratio-the commonly quite low. Using knowledge of animal beha- ratio of breeding males to females. Behavioural traits viour to design management regimes may only margin- can in¯uence the operational sex ratio in a number of ally increase Ne, yet this may be all that is needed to ways. For instance, mature animals may be prevented ensure population viability. from mating by dominant conspeci®cs or there may be Behaviour can aect Ne through its eects on N; r, active mate choice mechanisms that prevent certain and reproductive skew. N; r, and reproductive skew animals from reproducing. Behaviour contributes to in¯uence Ne in at least ®ve ways by their solitary and predation risk and skewed operational sex ratios emerge combined eects (Fig. 1). In this paper we discuss nine via dierential mortality and survival Ð a `double common behavioural traits that either directly or indir- whammy' for species already in danger of extinction. ectly aect Ne. We focus on reproductive suppression, Wright (1938) illustrated how skewed sex ratios in¯u- sexually-selected infanticide, mechanisms of mate- ence Ne. Speci®cally, choice, mating systems, social plasticity, dispersal, migration, conspeci®c attraction, and reproductive 1 behaviours which require special resources. These N 1 e 1 1 behavioural traits are found in many taxa. By discussing 4N 4N these, we hope to illustrate how behaviour can help m f inform conservation biology and how a wildlife man- ager might go about determining whether knowledge of where Nf is the number of breeding females
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