OUP CORRECTED PROOF – FINAL, 05/24/12, SPi - Life Sciences

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1 OUP CORRECTED PROOF FINAL, 05/24/12, SPi CHAPTER 14 The role of behavioural variation in the invasion of new areas Ben L. Phillips and Andrew V. Suarez Overview Behaviour determines the rate at which invasive species spread, as well as the impact they have on natives. When behaviour varies between individuals (as it almost always does), then the mean behaviour is often less important than the extremes of behaviour. The rate at which a species spreads, for example, is governed primarily by the most extreme dispersers. Similarly, individuals of native species that are extreme in their behaviour may be more, or less, likely to suffer impact from a given invasive species. Thus, we argue, an understanding of behavioural variation is critical if we are to understand the long-term impacts of invasive species in a changing world. (Blackburn and Duncan 2001; Simons 2003; Tingley 14.1 Introduction et al. 2010). Very few species pass easily through all Biological invasions are now widely considered to three filters. be a form of global change (Vitousek et al. 1996). In Importantly, the same filtering process that addition to severe economic consequences (Pimental applies to taxa also applies to individuals within an et al. 2000), the introduction of species into new invasive taxon (in fact, it applies to the individuals environments can have devastating ecological first, and the taxon only as an after-effect). The indi- impacts (Mack et al. 2000; Parker et al. 1999). It is viduals that are transported, that form established something of a relief then that we dont have a lot populations, and that become invasive are all poten- more invasive species, particularly given that every tially non-random subsets of the population from year probably hundreds to thousands of species are which they originated (Phillips et al. 2010a). Thus, introduced outside their natural range. Luckily, population-level variation in traits associated with very few of these go on to establish, spread, and invasion success is a key determinant of the success, have large ecological impacts. Why is this so? It or otherwise, of a particular invasion. turns out that the different stages of the invasion Invasive species do not, however, occur in a vac- processtransportation across dispersal barriers, uum. As an invader spreads, its interactions with establishment of a population at the new location, resident species become more widespread and the and post-establishment spreadact as strong phe- success of the invader, as well as that of the invaded, notypic filters, winnowing out the majority of spe- may well depend on these interactions. Variation is cies. Thus, a non-random subset of all taxa are important here too. Introduced species often inter- transported, a non-random subset of these are act negatively with native species (either directly, as then capable of establishment, and then a non- predators, competitors, and parasites; or indirectly, random subset of these become spreading invaders through habitat modification). So the variation in Behavioural Responses to a Changing World. First Edition. Edited by Ulrika Candolin and Bob B.M. Wong. 2012 Oxford University Press. Published 2012 by Oxford University Press.

2 OUP CORRECTED PROOF FINAL, 05/24/12, SPi T HE R OL E O F B EH AV I O U R A L VA R I AT I O N I N T H E I NVA S I O N O F N EW A R EA S 191 traits mediating these interactions will determine individuals through space (and into new space), which native individuals survive and which do not: and population growth fills space (including newly natural selection in action. colonized space) with individuals. The rate at which Many of the traits associated with invasion and a population spreads, then, is determined almost interspecific interaction are behavioural. In this entirely by these two fundamental processes. This chapter, we focus primarily on behaviour during said, however, a surprising array of spread dynam- the post-establishment stage of invasion: spread. ics are possible (including lags, accelerating, pulsed, We discuss the processes that lead to population and constant spread rates) depending upon how spread, and some of the behaviours that facilitate these fundamental processes are executed (Hastings those processes. We also discuss the role of behav- et al. 2005; Johnson et al. 2006). iour in determining the outcomes of interspecific If there is variation in a population for traits interactions between introduced and resident spe- affecting dispersal or population growth rate, things cies. In both cases we focus not just on behaviours, get more complex. In addition to the two funda- but on the effect of behavioural variation. We will mental processes of spread (dispersal and popula- see that behavioural variation not only matters, but tion growth), two unavoidable consequences of that it is central to understanding both the process spread allow trait variation to come into play of invasion, and the impact of invaders on natives. (Phillips et al. 2010a). The first unavoidable conse- quence is that, on the invasion front, individuals are spatially assorted by dispersal ability. This is known 14.2 Behaviours influencing the process as the Olympic village effect, or spatial sorting of spread (Phillips et al. 2008a; Shine et al. 2011), and leads to assortative mating by dispersal ability and runaway 14.2.1 The mechanics of spread selection for increased dispersal on the invasion Populations spread through space as a function front (Travis and Dytham 2002; Hughes et al. 2007; of two fundamental processes: dispersal and popu- Fig. 14.1). The second unavoidable consequence of lation growth (Skellam 1951). Dispersal moves range advance is that individuals on the expanding 4.5 (a) (b) 50 log(mean daily displacement) 4.0 Spread rate (km/year) 40 3.5 30 3.0 20 10 2.5 1960 1980 2000 0 500 1000 1500 Year Distance from invasion source (km) Figure 14.1 The accelerating rate of invasion of cane toads Rhinella marina across northern Australia (a), is associated with evolutionarily increased dispersal rates (b) shown here as an increase in mean daily displacement of animals collected from populations spanning the earliest introduction point (zero on the x-axis) through to the still-expanding invasion front. This is a pattern increasingly seen in the rate of spread of many invasive species. Error bars are one standard error. Data redrawn from Phillips et al. 2006, 2008a.

3 OUP CORRECTED PROOF FINAL, 05/24/12, SPi 192 BEHAV IOUR AL R E S PON S E S TO A C HAN GI NG W O R LD edge are in an environment with a very low density Another cheap technique is to hitch a lift. Small of conspecifics relative to established populations. species that allow other, typically much larger, spe- In such an environment, traits that increase repro- cies to effect their movement may get around very ductive rate or decrease Allee effects are favoured cheaply indeed. Some of the best examples of such (Burton et al. 2010). facilitated dispersal come from epidemiology, With these two fundamental processes and two where diseases are spread rapidly across an area by unavoidable consequences, we now have a frame- their animal vectors (e.g. spread of West Nile virus, work with which to examine the influence of behav- Loss et al. 2009; Seidowski et al. 2010). Of course iour, and behavioural variation, on spread. humans also make excellent targets for facilitated dispersal. Human-mediated jump dispersal is not only responsible for creating new foci of invasion 14.2.2 Dispersal behaviour during spread well away from established populations, but may Dispersal, the permanent movement of an animal also be the primary means by which some species from its birth place to its place of first reproduction spread at the landscape level (e.g. Argentine ants (sensu Howard, 1960), carries strong costs, but Linepithema humile, Suarez et al. 2001; snails Xeropicta despite this, is almost ubiquitous across all living derbentina, Aubry et al., 2006). organisms (Clobert et al. 2001). Dispersal, therefore, Finally, in cases where dispersal is active, must also carry strong rewards, and these include increased efficiency can be gained by straightening the avoidance of inbreeding and kin competition, the movement path (Barto et al. 2009): individuals escape from parasites and pathogens, as well as the that disperse in a relatively straight line will, ulti- subtler advantages of colonizing newly available mately, displace much further than individuals fol- patches in an extinction prone landscape (e.g. lowing a more tortuous path. Indeed, a tendency to Hamilton and May 1977; Van Valen 1971; Gandon follow straight paths is a clear attribute of the most and Michalakis 2001; Weisser et al. 2001). Thus, rapidly spreading populations of invasive cane most organisms disperse, but getting from A to B toads Rhinella marina in northern Australia (Alford can be done in an astonishing number of ways. It et al. 2009; Phillips et al., 2008a). can be done passively, such as a windborne seed, actively, such as a kangaroo hopping, or in an amal- 14.2.3 Behaviour and population growth gam of the two. during spread The most successful invaders are those that spread over large areas very quickly. Invasive spe- In broad terms, population growth is the net result cies will therefore tend to have behaviours that gen- of the number of births in a population minus the erate long-distance dispersal relatively cheaply number of deaths. The number of births and deaths, (Barto et al. 2009). Typically, such dispersal strate- however, is a gross summary of the myriad behav- gies involve some form of passive or active/passive iours, morphologies, and physiologies that allow combination. In terrestrial realms, flight is perhaps individuals in a population to survive, grow, and the most obvious amalgam of active and passive reproduce. In this light, then, almost any behaviour dispersal: although getting airborne might be ener- has some influence on population growth. For logis- getically expensive, the ability to harness the move- tical reasons then, we focus here on behaviours that ment of air once aloft can disperse airborne animals have an obvious bearing on the colonization of new over vast distances. Thus, many of the most rapid areas: behaviours that allow individuals to survive and famous invasions have involved flying species, and quickly capture resources for growth and repro- such as the house finch Carpodacus mexicanus, house duction in a new environment. Although there has sparrow Passer domesticus, the European starling been no definitive laundry list of behavioural char- Sturnus vulgaris, the Eurasian collared dove acteristics that convey invasion success, it has been Streptopelia decaocto, and the gypsy moth Lymantria suggested that some degree of sociality along with dispar (Elton 1958; Veit and Lewis 1996). high levels of aggression, activity (foraging/search-

4 OUP CORRECTED PROOF FINAL, 05/24/12, SPi T HE R OL E O F B EH AV I O U R A L VA R I AT I O N I N T H E I NVA S I O N O F N EW A R EA S 193 into the spread process. Importantly, plastic Boldness Boldness Activity responses will have an almost instantaneous effect on spread rate (as they occur as a direct response to the environment), whereas evolved responses will take longer (several to many generations) to play Aggression Aggression Activity out. Plastic responses may or may not increase Figure 14.2 Relationship between foraging activity, intraspecific spread rate, whereas evolved responses will likely aggression, and boldness in the invasive signal crayfish Pacifastacus always increase spread rate. leniusculus. All correlations are significant (p

5 OUP CORRECTED PROOF FINAL, 05/24/12, SPi 194 BEHAV IOUR AL R E S PON S E S TO A C HAN GI NG W O R LD or as a consequence of longer term selection for ability, and r-selection (driven by low conspecific individuals that are better at finding mates in situa- density), which drive selection for increased disper- tions of low conspecific density, is unknown. sal and reproductive rates (Burton et al. 2010). Thus, Plastic responses might also occur as accidents of during invasion, behaviours will evolve to increase a changed environment during invasion. For exam- dispersal and, if possible given trade-offs with dis- ple, many frog species alter both their behaviour persal, increase reproductive rates. Our brief survey and developmental rate as tadpoles in response to of behavioural traits that facilitate invasion (above) the presence of aquatic predators (Kiesecker and gives us an idea as to the potential suite of traits that Blaustein 1997; Relyea 2001). One of the conse- might evolve in this way during invasion. quences of low conspecific density and assortment Empirical evidence for changed dispersal behav- by dispersal on the edge of an invading population iour during invasion is rapidly accumulating. is that coevolved predators often get left behind Insects on expanding range edges have been shown (Phillips et al. 2010d). If this happens, then the cues to invest more in dispersal than their conspecifics in driving the typical predator response may be absent, older, established populations (Hughes et al. 2007; and a morphologically and behaviourally predator Simmons and Thomas 2004; Lotard et al. 2009), free phenotype will emerge (even if predators, and invasive cane toads in northern Australia have albeit unfamiliar ones, are still present). Typically, evolved increased dispersal rates on the invasion tadpoles in the absence of predators spend more front relative to their conspecifics from older, estab- time active, and have higher growth rates than con- lished populations (Phillips et al. 2010b; Phillips specifics in the presence of predators (e.g. Thiemann et al. 2008a). and Wassersug 2000; Relyea 2001). Higher individ- Empirical evidence for the evolution of behav- ual growth rates resulting from tadpoles behaving iours facilitating population growth is, however, in a predator free manner could, conceivably, lead weaker. Higher growth rates have been found in to higher population growth rates and faster spread, invasion front populations of cane toads (Phillips although this net effect would be an accidental 2009), but the behavioural correlates of this higher byproduct of an animal in an unfamiliar environ- growth rate, if any, remain to be elucidated. The ment. Alternatively, of course, increased predation increased sensitivity of invasion front starlings to rates driven by inappropriate responses to unfamil- the calls of their conspecifics (Rodriguez et al. 2010) iar predators could depress population growth and might be plastic or evolved (or both), we simply do slow spread. The effects of plasticity could, conceiv- not know yet. But as an adaptation to reducing ably, either accelerate or decelerate an invasion. Allee effects and increasing population growth rate on an invasion front, we might expect it to be an evolved shift. Generally, the importance of compar- 14.3.2 Evolved responses ing newly colonized and older established popula- Traditionally, we have thought about evolution in tions inside the invasive range is only recently being invasive species as being about adaptation to their appreciated. So the field is open for behavioural new environment (Mack et al. 2000). While this ecologists to start investigating behavioural shift in standard adaptive process must indeed be happen- response to range shift. ing after an invasive population has become estab- Another intriguing aspect of evolution during lished, it is increasingly apparent that evolutionary invasive spread is that it may bring together multi- processes on the invasion front itself are driven by ple behavioural traits into sets. Being such a strong additional forces (Phillips et al. 2010a). Invasion directed selective pressure, and being directed at fronts can capture rare, and even deleterious, muta- dispersal and reproductive rates (complex traits tions, drive them to high frequency, and then spread typically driven by multiple behaviours), we would them across large areas as the invasion front moves expect range shift to start accumulating individuals through space (Travis et al. 2007). Coupled with this with multiple behaviours that increase dispersal are the processes of assortative mating by dispersal and reproductive rates, whilst minimizing Allee

6 OUP CORRECTED PROOF FINAL, 05/24/12, SPi T HE R OL E O F B EH AV I O U R A L VA R I AT I O N I N T H E I NVA S I O N O F N EW A R EA S 195 effects (Travis et al. 2010; Shine et al. 2011). In than 22 species of native mammal were lost from Australian cane toads, individuals from the invasion the Australian continent following the arrival of the front grow faster, move more often, move further invasive cat and fox (McKenzie et al. 2007). Similarly, when they do move, and follow straighter paths in Lake Victoria, the introduction of the Nile perch than their trailing conspecifics, all of which con- Lates niloticus led to the extinction of around 200 spires to increase their net dispersal and reproduc- species of native cichlid fish (Ogutu-Ohwayo 1999). tive rates (Phillips et al. 2008a; Alford et al. 2009; In New Zealand, the introduction of Pacific rats Phillips 2009). Similarly, we might expect all traits Rattus exulans led to the extinction from the main- that influence dispersal and reproductive rates to land of many species of birds and lizards (Towns become associated in invasion front populations. et al. 2007). These examples of extinction point to Given that many of the traits associated with ani- situations where native species simply didnt have mal personality (e.g. aggression and boldness: appropriate behavioural variation (in both evolved Duckworth 2008; Duckworth and Badyaev 2007; Sih and plastic senses) to cope with the arrival of a new et al. 2004b) may also be associated with dispersal predator. Potentially, however, there may be many and/or reproduction, range shifts can, over time, more (unreported) instances where natives can assemble populations with distinct personalities modify behaviours in response to predation, and so (e.g. aggressive, risk taking, and highly active). Why persist. For example, Hoare et al. (2007) show that, animals exhibit distinct personalities is a current on New Zealand islands colonized by Pacific rats, focus amongst behavioural ecologists (Dingemanse native geckoes Hoplodactylus duvaucelii rapidly shift et al. 2010; Sih et al. 2004b). It may be that selection their habitat preferences to avoid interactions with for dispersal (Cote et al. 2010) during invasion is a the predatory rats. Unfortunately, these subtler currently under-appreciated way for animal person- examples (where native species are not extirpated, alities to be assembled. Future work exploring this but survive through behavioural shifts) are possibility would likely be rewarding. Such work understudied. would focus strongly on behavioural variation, and The impacts of introduced species are also likely how it varies through invasion history. to be density dependent (Yokomizo et al. 2009). Therefore, behaviours that influence the density of species in introduced populations may dispropor- 14.4 Behavioural variation and the tionately influence their impact. For example, impacts of invasive species on natives intraspecific competition is often cited as a primary Invasive species are typically of broad interest mechanism that regulates relative abundance (e.g. because of their perceived impacts on native spe- Ryti and Case 1988; Comita et al. 2010), so highly cies. Almost all the impacts of invasive species on territorial species that exclude conspecifics from natives are likely mediated by behaviour. Both the large areas are not likely to become high-impact behaviour of the invader, and that of the native, invasives. In contrast, social species that do not determine the outcome. Where the outcome of an exhibit discrete or exclusive territories may attain interaction is negative (either for the native or the higher densities and may also avoid Allee effects invader, or both), we might expect to see rapid evo- that inhibit population spread. While an association lution of behaviour. Thus, species invasions offer between intraspecific aggression and impact has immense opportunities for behavioural ecologists not been explored across many taxa, it has been interested in the evolution (and coevolution) of suggested as a mechanism for the success of inva- behaviour. Invasive species often represent a novel sive ants (Holway and Suarez 1999). Many intro- selective force in the environment that native spe- duced populations of highly invasive ants are cies need to respond to through behavioural means unicolonialspatially separate nests behave as (Ashley et al. 2003). one colony across the entire population (Holway Many of the biggest impacts from invasive ani- et al. 2002); a situation that is not always the case in mals occur when the invader is a predator. More their native range. In unicolonial populations where

7 OUP CORRECTED PROOF FINAL, 05/24/12, SPi 196 BEHAV IOUR AL R E S PON S E S TO A C HAN GI NG W O R LD intraspecific territorial behaviour is absent or cies toxicity (Phillips and Shine 2007). Non-toxic reduced, colonies often exhibit higher resource frogs are simply grabbed and swallowed, whereas retrieval rates, and greater brood and worker pro- frogs which secrete a sticky glue or which are neu- duction relative to native species that exhibit terri- rotoxic are envenomated and released to die, after torial behaviour (Holway et al. 1998). The origin which the snake then tracks the meal down and and maintenance of unicoloniality in ants may pro- ingests it. Snakes wait around ten minutes between vide insight into the role of behavioural variation in bite and consumption for the gluey frogs but wait the success and impact of invasive species more much longer (typically around 40 minutes) before generally, and is currently the subject of intense tasting and, ultimately, consuming the neurotoxic debate and study (Helantera et al. 2009). species. When these snakes first encounter a toad, Behavioural variation across environmental con- the response of individuals varied: some snakes texts may also be important in influencing the out- treat the toad as if it was non-toxic, while others comes of competition with invasive species. For treat them as if they were gluey, neurotoxic, or example, by hanging lights in abandoned Second something that is ultimately inedible (Hagman et al. World War aircraft hangers in Hawaii, Petren and 2009). Almost all death adders that consume a toad colleagues (1993) were able to experimentally dem- die, thus, a snakes classification of this new prey onstrate how increasing the local abundance of item (and the subsequent behaviours that flow from insects influenced competitive interactions between that classification), strongly defines the impact that two species of gecko: the invasive Hemidactylus fre- the arrival of toads will have on that individual. natus and the resident Lepidodactylus lugubris. The This correlation between behaviour and impact larger, more aggressive H. frenatus was able to for- suggests strong selection operating on prey prefer- age more effectively and monopolize clumped ence in this species (Phillips et al. 2010c), and indeed resources in open areas relative to the less aggres- at least one Australian snake species appears to sive L. lugubris. However, the effects of competition have evolved behavioural avoidance of toads as between the species disappeared if resources were prey due to this strong selection pressure (Phillips not clumped or if complex topography was added and Shine 2006). around the light sources to provide safe refuges for These examples show the critical importance of the less aggressive species to hide in (Petren and the behaviour of both native and invasive species in Case 1998; Fig. 14.3). mediating the invasives impact. Importantly, how- Another well-studied example of behavioural ever, they also demonstrate the critical importance mediation of impacts concerns the arrival, in of behavioural variability (both plastic and evolved) Australia, of the toxic cane toad. In Australia, toads in determining the long-term outcome of that occur in very high densities, and show no territori- impact. Behavioural plasticity or rapid evolved ality. They are a predator, but their major impact responses may be integral for native species to per- appears to be on native predators that mistake toads sist in invaded landscapes. as palatable prey. Numerous native predators, which were nave to the toxins in the toads skin, 14.5 Conclusion and future directions have been fatally poisoned following the arrival of toads. Despite the simplicity of the impact mode Through both theory and example, we have dem- poisoningimpacts have varied tremendously, onstrated the importance of behaviour to both the both within populations as well as between closely spread and impact of invasive species. Spread rate related species (Shine 2010). Much of this variation (governed by dispersal and population growth) is, in impact is driven by subtle differences in the ultimately, an outcome of the behaviour of the natives prey handling behaviour. For example, invader. Similarly, the impact of the invader depends death adders Acanthophis praelongus in northern on the interaction between its behaviour and that of Australia handle different prey species (all of which nearby native species. The central importance of are frogs) differently, depending upon the prey spe- behaviour in these cases is not particularly surpris-

8 OUP CORRECTED PROOF FINAL, 05/24/12, SPi T HE R OL E O F B EH AV I O U R A L VA R I AT I O N I N T H E I NVA S I O N O F N EW A R EA S 197 0.04 Li+Hf, complex topography 0.03 Li+Li, complex topography 0.02 LI body condition 0.01 (mean SE) 0 0.01 0.02 Li+Li, simple 0.03 topography 0.04 Li+Hf, simple topography 0.05 0 10 20 30 40 50 60 70 Days Figure 14.3 The distribution of resources (clumped versus dispersed) and the degree of habitat complexity influences the outcome of interspecific competition between invasive Hemidactylus frenatus Hf, and resident Lepidodactylus lugubris Li, geckos. When insect resources are clumped at light sources, and the space around the lights is open, the larger more aggressive Hemidactylus can monopolize access to insects. However, if resources are not clumped and/or baffles are placed around the lights creating refuges for the geckos, Lepidodactylus are able to forage more effectively, evidenced by an increase in body condition. Figure reprinted from Petren and Case 1998, with permission. Copyright (1998) National Academy of Sciences, U.S.A. ing, but consideration of behavioural variation References (either plastic or evolved) can lead to surprising Alford, R. A., Brown, G. P., Schwarzkopf, L., Phillips, B. L., results. For example, early predictions of the spread and Shine, R. (2009). Comparisons through time and rate of cane toads across northern Australia were space suggest rapid evolution of dispersal behaviour in rendered wildly inaccurate by the rapid evolution an invasive species. Wildlife Research, 36, 238. of increased dispersal behaviour in this species Amiel, J. J., Tingley, R., and Shine, R. (2011). Effects of (Phillips et al. 2008b). Similarly, many predictions of relative brain size on establishment success of inva- deleterious impacts of invasive species on natives sive amphibians and reptiles. PLOS ONE 6: e18277. may prove to be fleeting as native species exhibit doi:10.1371/journal.pone.0018277. behavioural flexibility in response to the invader Ashley, M. V., Willson, M. F., Pergams, O. R. W., ODowd, (e.g. Langkilde 2009). D. J., Gende, S. M., and Brown, J. S. (2003). Evolutionarily The future, thus, should be about variation. When enlightened management. Biological Conservation, 111, an invader arrives, the average behaviour often 11523. Aubry, S., Labaune, C., Magnin, F., Roche, P., and Kiss, L. turns out to be much less important than extremes (2006). Active and passive dispersal of an invading land of behaviour. Extreme dispersers come to dominate snail in Mediterranean France. Journal of Animal Ecology, the vanguard of an invasive population, and 75, 80213. extreme behaviours may be associated with indi- Barto, K. A., Phillips, B. L., Morales, J. M., and Travis, J. viduals most or least at risk from the presence of an M. J. (2009). The evolution of an intelligent dispersal invader. The challenge for behavioural ecologists is strategy: biased, correlated random walks on patchy thus clear. We need to understand behavioural vari- landscapes. Oikos, 118, 30919. ability; within and between individuals and, within Blackburn, T. M. and Duncan, R. P. (2001). Establishment and between populations. We need to understand patterns of exotic birds are constrained by non-random both the genesis of such variation as well as its envi- patterns in introduction. Journal of Biogeography, 28, ronmental correlates. Only by understanding this 92739. Burton, O. J., Travis, J. M. J., and Phillips, B. L. (2010). variation can we hope to predict behavioural Trade-offs and the evolution of life-histories during responses into the future, in a changing world. range expansion. Ecology Letters, 13, 121020.

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