Survey
* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project
* Your assessment is very important for improving the workof artificial intelligence, which forms the content of this project
F I S H and F I S H E R I E S , 2014, 15, 593–615 Latitudinal shifts in coral reef fishes: why some species do and others do not shift David A Feary1, Morgan S Pratchett2, Micheal J Emslie3, Ashley M Fowler1, Will F Figueira4, Osmar J Luiz5, Yohei Nakamura6 & David J Booth1 1 School of the Environment, University of Technology, 123 Broadway, Sydney, NSW 2007, Australia; 2ARC Centre of Excellence for Coral Reef Studies, James Cook University, Townsville, Qld. Q4811, Australia; 3Australian Institute of Marine Science, PMB No.3, TMC, Townsville, Qld. 4810, Australia; 4School of Biological Sciences, University of Sydney, Sydney, NSW 2006, Australia; 5Department of Biological Sciences, Macquarie University, Sydney, NSW 2109, Australia; 6Graduate School of Kuroshio Science, Kochi University, 200 Monobe, Nankoku, Kochi 783-8502, Japan Abstract Climate change is resulting in rapid poleward shifts in the geographical distribution of many tropical fish species, but it is equally apparent that some fishes are failing to exhibit expected shifts in their geographical distribution. There is still little understanding of the species-specific traits that may constrain or promote successful establishment of populations in temperate regions. We review the factors likely to affect population establishment, including larval supply, settlement and post-settlement processes. In addition, we conduct meta-analyses on existing and new data to examine relationships between species-specific traits and vagrancy. We show that tropical vagrant species are more likely to originate from high-latitude populations, while at the demographic level, tropical fish species with large body size, high swimming ability, large size at settlement and pelagic spawning behaviour are more likely to show successful settlement into temperate habitats. We also show that both habitat and food limitation at settlement and within juvenile stages may constrain tropical vagrant communities to those species with medium to low reliance on coral resources. Correspondence: David A Feary, School of the Environment, University of Technology, Sydney, 123 Broadway, NSW 2007, Australia Tel.: +61 2 9514 4068 Fax: +61 2 9514 4079 E-mail: david. [email protected] Received 10 Sep 2012 Accepted 6 Mar 2013 Keywords Climate change adaptation, global warming, range shifts, temperate reef, tropical reef fishes, tropical vagrant Introduction 594 Scope of this review 594 Section 1 Extrinsic factors regulating arrival of tropical vagrants 596 Oceanographic factors 596 Large-scale ocean currents 597 Ocean eddies 598 Section 2 Intrinsic factors constraining latitudinal population movement 598 Environmental constraints to distributional shifts 598 Population density and latitudinal distribution 599 Life-history traits associated with vagrancy 600 Correlations between species-specific traits in predicting vagrancy 603 © 2013 John Wiley & Sons Ltd DOI: 10.1111/faf.12036 593 Tropical vagrant fishes D A Feary et al, Resource constraints to vagrant success 603 Habitat association and settlement preferences 604 Dietary preferences and functional groups 605 Section 3 Future research needs 606 Conclusions 607 Acknowledgements 608 References 608 Supporting Information 615 Introduction A central premise of biogeography is that the natural distribution of a species is at least partly governed by climate (Guisan and Zimmermann 2000; Parmesan et al. 2005; Soberon 2007; Sunday et al. 2012). Therefore, we can expect to see major changes in the distribution of species associated with changes in global climate (IPCC 2007). The fossil record (Davis et al. 2002; Carnaval and Moritz 2008) and recent observed trends (Parmesan et al. 1999; Thomas and Lennon 1999; Hickling et al. 2006; Burrows et al. 2011) attest to increased incidence of natural populations showing both range expansion and contraction during periods of rapid climate change. Metabolism, growth, reproduction and ultimately survival of all organisms are tightly linked to temperature, and there are both upper and lower limits within which organisms can survive (Hurst 2007). During warming periods, organisms move poleward either to escape deleterious effects of high temperatures at low latitudes or to take advantage of high-latitude locations which they could not otherwise tolerate (Parmesan and Yohe 2003; Hoegh-Guldberg and Bruno 2010). To date, poleward shifts in species geographical distribution are most apparent for terrestrial plants and animals (Parmesan and Yohe 2003; Parmesan 2006; Burrows et al. 2011). However, there is increasing evidence of range shifts among marine fishes and invertebrates, especially at high latitudes in the Northern Hemisphere (Dulvy et al. 2008; Nye et al. 2009; Stefansdottir et al. 2010). This bias in recorded range shifts (between hemispheres and among latitudes) is consistent with higher rates of temperature change recorded in Northern Hemisphere, high-latitude regions (IPCC 2007). However, large-scale changes in ocean temperature are also taking place at low latitudes, characterized by a widening of the tropical belt (Seidel et al. 2008; Lu et al. 2009). Such changes 594 have been linked to poleward expansions in the distribution of many tropical organisms (Booth et al. 2007, 2011; Madin et al. 2012). The distribution of tropical marine organisms has been substantially affected by changes in both ocean temperatures and major ocean currents (Seidel et al. 2008; Lu et al. 2009). Since 1900, surface waters associated with western boundary currents (e.g. Gulf Stream, Agulhas Current) have increased in temperature 2–3 times faster than the global mean surface ocean warming rate (Wu et al. 2012). Such warming of surface currents has also been associated with a poleward shift in the extent of these boundary currents and substantial change in the distribution and extent of tropical benthic species (Oviatt 2004; Helmuth et al. 2006; Yamano et al. 2011). Such poleward shifts in warm currents have also had important effects on tropical reef fish larvae, which can frequently be transported 100– 1000s of kilometres from tropical to temperate latitudes (e.g. Booth et al. 2007; Figueira et al. 2009; Hirata et al. 2011; Soeparno et al. 2012). Scope of this review There is increasing evidence that climate change is leading to rapid changes in marine species’ distributional envelope, with such range shifts expected to increase in strength and intensity as global climatic conditions change (Booth et al. 2011; Madin et al. 2012). While there is mounting evidence that benthic marine communities are showing substantial changes in their distribution and boundaries (Greenstein and Pandolfi 2008; Yamano et al. 2011), relatively less is known of the potential impacts on the associated fish communities (Munday et al. 2008a, 2009; but see Burrows et al. 2011). Although climate-mediated shifts in the geographical range of temperate and subtropical fish species are expected, and indeed have been shown in several regions (Sorte et al. 2010; Last © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 Tropical vagrant fishes D A Feary et al., et al. 2011), tropical fish species may be particularly sensitive to increasing temperatures as they exist in a relatively thermostable environment (Hoegh-Guldberg et al. 2007). Most tropical fishes may not live within environments that are close to their lethal thermal limits (Mora and Ospına 2001), but we can expect that elevated ocean temperature may have substantial effects on individual performance, with potential implications for population shifts (Munday et al. 2008a, 2009). Despite this, there is little predictive framework to understand whether species-specific traits may structure range shifts within tropical fish assemblages and the potential changes this may have in the composition and structure of tropical fish communities (Munday et al. 2008a,b). Indeed, there is mounting evidence to suggest that marine organisms have shown average geographical shifts associated with climate-mediated global thermal shifts from 1.5 to 5 times faster than their terrestrial counterparts (termed ‘climate velocity’, sensu Burrows et al. 2011). The question then is what are the factors that may allow tropical coral reef species to track ‘climate velocity’ closely? Within this review, we argue that species-specific demographic traits (e.g. body size, post-larval duration, size at settlement, reproductive behaviour) are likely to be important in structuring such geographical shifts in coral reef fish species. An understanding of how such traits may differ between tropical fish species is lacking in tropical reef science and is the focus of this review. Within this review, we focus solely on marine ray-finned fishes (Class Actinopterygii) that breed within tropical coral reef habitats (dominated by reef building, scleractinian reefs), and their larvae show or have shown settlement into temperate reef habitats (hereafter termed ‘tropical vagrants’). Although there is increasing awareness that other groups of mobile tropical organisms are showing shifts in their latitudinal distribution (i.e. Elasmobranchii: Last et al. 2011), these studies are outside the scope of this review. All of the tropical vagrants identified in this review, despite being found within temperate habitats, show substantial reductions in abundance associated with low winter-water temperatures (e.g. Choat et al. 1988; Francis et al. 1999; Booth et al. 2007; Figueira and Booth 2010). However, as global sea surface temperature increases are associated with climate warming scenarios (IPCC 2007), we predict that for a suite of tropical vagrants being advected into high-latitude regions, survival, adaptation and population development may occur rapidly (Figueira and Booth 2010; Booth et al. 2011; Madin et al. 2012; Soeparno et al. 2012). We examine both the intrinsic and extrinsic processes that likely influence the spatial, temporal and taxonomic biases in the species-specific structure of tropical vagrant populations (Fig. 1). It is assumed a priori that extrinsic processes (i.e. oceanographic factors) will interact with inherent differences in the life histories of fishes to determine which species settle, as well as when and where (see also Munday et al. 2009). Therefore, we first examine the oceanographic factors that may facilitate physical connectivity between tropical and temperate regions. Determining the intrinsic processes that may facilitate or constrain latitudinal movement is expected to be key to understanding which tropical species may be increasingly susceptible to sustained and ongoing climate change (Cowen et al. 2000, 2006; Jones et al. 2005, 2009a; Almany et al. 2007a; Hobbs et al. 2010, 2012). Therefore, within this review, we examine the potential physiological constraints to successful settlement and recruitment of tropical vagrants in temperate regions. We then consider pre-settlement mechanisms associated with successful advection of tropical vagrants into high-latitude temperate regions. Cheung et al. (2010) in their global analysis on the redistribution of marine fishes suggested that range shifts are likely to be constrained by specific resource (prey or habitat) requirements for some fishes. Thus, we also examine the potential importance of resource requirements in successful settlement and survival of tropical vagrants within temperate habitats. Finally, albeit not exhaustive, we highlight the array of research questions that will be vital in understanding the role of increasing climate change in structuring the range expansion of tropical coral reef fishes into temperate environments. A total of 360 species of tropical fishes (within the Class Actinopterygii) from 55 different families have been recorded settling into temperate regions (hereafter termed ‘tropical vagrants’) (Supplementary Data S1). Despite this, vagrants still appear relatively uncommon within families (Fig. 2). Tropical vagrants include a disproportionate number of species from the families Acanthuridae, Balistidae, Chaetodontidae, Cirrhitidae, Labridae, Lutjanidae, Mullidae, Pomacentridae and Scaridae (Fig. 2). In comparison, several common tropical families are extremely under-represented within © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 595 Tropical vagrant fishes D A Feary et al, Figure 1 Schematic of factors potentially limiting the range expansion of tropical fishes in temperate habitats during three stages of the expansion process (natal, larval and novel environments). Factors discussed and/or tested in this study are indicated by the section number in parentheses. vagrant surveys, including the Apogonidae, Callionymidae, Gobiesocidae, Gobiidae, Serranidae, Syngnathidae and Tripterygiidae (Fig. 2). It is important to note that the majority of these under-represented families predominantly hold small, relatively cryptic species (see Munday and Jones 1998), which may result in such species being missed in vagrant surveys, rather than not tending to be vagrants (see Section 2). However, as the majority of families that hold vagrants are also relatively small-bodied species, and the majority of vagrant survey data comprise surveys of new recruit and early-stage juveniles (~2–3 cm TL), other ecological traits may also be important in structuring vagrant success (see Section 2). Section 1: Extrinsic factors regulating arrival of tropical vagrants Oceanographic factors Most marine organisms have a bipartite life history, characterized by a sessile (often reproductive) 596 phase and highly dispersive (often larval) phase (Leis et al. 2011). As our knowledge regarding the physiology and behaviour of larval marine organisms grows, it is becoming increasingly apparent that contrary to initial speculation, many larvae possess complex behavioural abilities (including swimming and orientation behaviour) (Leis and Carson-Ewart 1998; Leis et al. 2002, 2003, 2007). Indeed, relatively high levels of local larval retention and self-recruitment (settlement of locally spawned larvae: up to 40–60%) have been shown (Jones et al. 1999, 2005; Swearer et al. 1999, 2002; Thorrold et al. 2001; Almany et al. 2007a; Harrison et al. 2012). However, the corollary of this means that 40–60% of a reef fish population may be exported long distances away from local reef area (Leis et al. 2011). In addition, while larval marine organisms, and fish in particular, can attain impressive swimming and orienting abilities, the onset of this ability may be species specific and form gradually (i.e. swimming ability: Fisher et al. 2000) or relatively rapid (i.e. orientation: Paris and Cowen 2004; Dixson et al. 2012). © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 Tropical vagrant fishes D A Feary et al, Riegl, B.M., Purkis, S.J., Al-Cibahy, A.S., Abdel-Moati, M.A. and Hoegh-Guldberg, O. (2011) Present limits to heat-adaptability in corals and population-level responses to climate extremes. PLoS ONE 6, e24802. Rodolfo-Metalpa, R., Reynaud, S., Allemand, D. and Ferrier-Pages, C. (2008) Temporal and depth responses of two temperate corals, Cladocora caespitosa and Oculina patagonica, from the North Mediterranean Sea. Marine Ecology Progress Series 369, 103–114. Roughan, M., Macdonald, H.S., Baird, M.E. and Glasby, T.M. (2011) Modelling coastal connectivity in a Western Boundary Current: seasonal and inter-annual variability. Deep-Sea Research Part Ii-Topical Studies in Oceanography 58, 628–644. Roughgarden, J. (2009) Is there a general theory of community ecology? Biology and Philosophy 24, 521– 529. Russell, B.C. (1971) A preliminary annotated checklist of fishes of the Poor Knights Islands. Tane 17, 81–90. Sale, P.F. (ed.) (1991) The Ecology of Fishes on Coral Reefs. Academic Press, New York, 754 pp. Sale, P.F. (ed.) (2002) Coral Reef Fishes. Dynamics and Diversity in a Complex Ecosystem. Academic Press, San Diego, 549 pp. Scott, A. and Harrison, P.L. (2008) Larval settlement and juvenile development of sea anemones that provide habitat for anemonefish. Marine Biology 154, 833–839. Seidel, D.J., Fu, Q., Randel, W.J. and Reichler, T.J. (2008) Widening of the tropical belt in a changing climate. Nature Geoscience 1, 21–24. Shanks, A.L. (2009) Pelagic larval duration and dispersal distance revisited. Biological Bulletin 216, 373–385. Soberon, J. (2007) Grinnellian and Eltonian niches and geographic distributions of species. Ecology Letters 10, 1115–1123. Soeparno, Nakamura, Y., Shibuno, T. and Yamaoka, K. (2012) Relationship between pelagic larval duration and abundance of tropical fishes on temperate coasts of Japan. Journal of Fish Biology 80, 346–357. Sorte, C.J.B., Williams, S.L. and Carlton, J.T. (2010) Marine range shifts and species introductions: comparative spread rates and community impacts. Global Ecology and Biogeography 19, 303–316. Sponaugle, S. and Grorud-Covert, K. (2006) Environmental variability, early life-history traits, and survival of new coral reef fish recruits. Integrative and Comparative Biology 46, 623–633. Sponaugle, S., Cowen, R.K., Shanks, A. et al. (2002) Predicting self-recruitment in marine populations: biophysical correlates and mechanisms. Bulletin of Marine Science 70, 341–375. Sponaugle, S., Boulay, J.N. and Rankin, T.L. (2011) Growth- and size-selective mortality in pelagic larvae of a common reef fish. Aquatic Biology 13, 263–273. 614 Stefansdottir, L., Solmundsson, J., Marteinsdottir, G., Kristinsson, K. and Jonasson, J.P. (2010) Groundfish species diversity and assemblage structure in Icelandic waters during recent years of warming. Fisheries Oceanography 19, 42–62. Stevens, G.C. (1989) The latitudinal gradients in geographical range: how so many species co-exist in the tropics. American Naturalist 133, 240–256. Stillman, J. and Somero, G.N. (2000) A comparative analysis of the upper thermal tolerance limits of eastern Pacific porcelain crabs, genus Petrolisthes: Influences of latitude, vertical zonation, acclimation, and phylogeny. Physiological and Biochemical Zoology 73, 200–208. Stobutzki, I.C. and Bellwood, D.R. (1994) An analysis of the critical swimming abilities of pre- and post-settlement coral reef fishes. Journal of Experimental Marine Biology and Ecology 175, 275–286. Stobutzki, I.C. and Bellwood, D.R. (1997) Sustained swimming abilities of the late pelagic stages of coral reef fishes. Marine Ecology Progress Series 149, 35–41. Sunday, J.M., Bates, A.E. and Dulvy, N.K. (2012) Thermal tolerance and the global redistribution of animals. Nature Climate Change 2, 686–690. doi:10.1038/NCLIMATE1539. Suthers, I.M. (1998) Bigger? Fatter? Or is faster growth better? Considerations on condition in larval and juvenile coral-reef fish. Australian Journal of Ecology 23, 265–273. Swearer, S.E., Caselle, J.E., Lea, D.W. and Warner, R.R. (1999) Larval retention and recruitment in an island population of a coral-reef fish. Nature 402, 799–802. Swearer, S.E., Shima, J.S., Hellberg, M.E. et al. (2002) Evidence of self-recruitment in demersal marine populations. Bulletin of Marine Science 70, 251–271. Sweatman, H. (1988) Field evidence that settling coralreef fish larvae detect resident fishes using dissolved chemical cues. Journal of Experimental Marine Biology and Ecology 124, 163–174. Syahailatua, A., Roughan, M. and Suthers, I.M. (2011) Characteristic ichthyoplankton taxa in the separation zone of the East Australian Current: larval assemblages as tracers of coastal mixing. Deep-Sea Research Part Ii-Topical Studies in Oceanography 58, 678–690. Thomas, C.D. and Lennon, J.J. (1999) Birds extend their ranges northwards. Nature 399, 213. Thorrold, S.R., Latkoczy, C., Swart, P.K. and Jones, C.M. (2001) Natal homing in a marine fish metapopulation. Science 291, 297–299. Thresher, R. (1984) Reproduction in Reef Fishes. TFH Publications, Neptune City. Thresher, R.E. and Brothers, E.B. (1989) Evidence of intra- and inter-oceanic regional differences in the early life history of reef-associated fishes. Marine Ecology Progress Series 57, 187–205. © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 Tropical vagrant fishes D A Feary et al, On the west coast, the Leeuwin Current actually seems to promote it (Condie et al. 2011). Thus, the highly coherent nature of these large-scale boundary currents, which may effectively transport larvae over great distances, may also impede the ability of those larvae to reach viable inshore habitat. Ocean eddies Recirculating current features, or eddies, have commonly been proposed as mechanisms that may retard, rather than encourage, long-distance transport of larva from a natal area (Sponaugle et al. 2002). For instance, eddies forming in the wake of islands may concentrate eggs or larvae (e.g. Wing et al. 1998) preventing their dispersal. Such recirculation may then result in the selfrecruitment of larvae, as has been suggested for the Tortugas Gyre and other spin-off eddies of the Florida Current (Lee et al. 1992, 1994, 1995; Limouzy-Paris et al. 1997). However, while eddies retain larvae near a natal site, they may also be extremely important for larval transport where individuals are entrained in large-scale ocean currents, but may also be important in trapping larvae when they arrive at new locations. Due to a complex mix of factors (e.g. topography, density gradients), eddies are commonly shed from ocean currents and may migrate, transporting the water masses they contain. For example, warm-core eddies shed by the Gulf Stream north of Cape Hatteras have been implicated in the transport of larvae towards coastal waters of not only cosmopolitan temperate fish larvae (Hare and Cowen 1996; Hare et al. 2002), but also tropical fish (Hare et al. 2002). The EAC separates from the east Australian coast at ~32°S latitude, shedding a series of cold- and warm-core eddies (Ridgway and Dunn 2003; Choukroun et al. 2010). The tropical water masses contained within the EAC and its resultant eddies have been shown to contain larval fish assemblages uniquely different from the surrounding waters (Keane and Neira 2008; Syahailatua et al. 2011). The dynamic nature of the eddy field south of the separation zone results in episodic patterns of tropical larval recruitment (Booth et al. 2007). A similar process occurs within the Leeuwin Current on the west Australian coast, where eddies greatly facilitate on-shelf transport of water masses (Condie et al. 2011). 598 Section 2: Intrinsic factors constraining latitudinal population movement Environmental constraints to distributional shifts Temperature is one of the primary variables regulating aquatic species bioenergetics (Kitchell et al. 1977). Ambient temperatures substantially determine physiological processes such as feeding, respiration, faecal egestion rates and ultimately growth (von Herbing 2002; Domenici et al. 2007). Even within their natal ranges, newly recruited fishes will experience a range of temperatures, both spatially and temporally, due to behavioural activities and in situ environmental fluctuations (Danilowicz 1997; Sponaugle and Grorud-Covert 2006; Abesamis and Russ 2010). Such changes in temperature can have substantial effects on fish physiology, defining species thermal geographical boundaries, even within relatively open-water habitats. We can predict that for warm-adapted tropical vagrants within cooler waters, thermal optima for physiological mechanisms may exist for limited periods during the year (i.e. warm summer months), and therefore, key bioenergetic parameters may only be maximized for a short time frame. Below, we examine the potential importance of physiological constraints to successful latitudinal extension in tropical reef fish populations. Environmental temperature is one of the most important physical variables affecting the performance of ectotherms (Hazel and Prosser 1974; Hurst 2007). Therefore, we can expect that one of the most important factors in structuring the success of tropical vagrants within temperate environments will be species-specific thermal limits for minimum environmental temperature (Attrill and Power 2002; Dulvy et al. 2008; Poertner and Farrell 2008). Virtually, all aspects of the behaviour and physiology of ectotherms are sensitive to environmental temperature (Hazel and Prosser 1974; Poertner and Farrell 2008). Therefore, we can expect that among tropical vagrants, the predominant source of mortality in temperate environments will be an individual’s inability to maintain cellular and organismal homoeostasis during winter (Hurst 2007). Although there is an increasing array of studies determining the upper thermal physiological limits within tropical fishes (Munday et al. 2008b; Donelson et al. 2011, 2012), there is comparatively little information on lower thermal limits. The majority of studies © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 Tropical vagrant fishes D A Feary et al., examining overwintering success have focused on temperate freshwater or estuarine fishes, examining survival within lakes or estuaries that have a partial or full winter freeze of surface waters (e.g. Hurst and Conover 2001; Pratt and Fox 2002). Recent work has shown that tropical fishes may be able to effectively withstand water temperatures much lower than predominantly found in tropical latitudes. Eme and Bennett (2008) found that water temperatures of ~15 °C were the critical thermal minimum for eight Indo-Pacific damselfishes (Pomacentridae), while Figueira et al. (2009) showed that for the tropical Indo-Pacific sergeant (Abudefduf vaigiensis, Pomacentridae), water temperatures needed to be 17 °C to cause substantial population loss within field surveys. Figueira et al. (2009) also demonstrated in aquaria trials that individuals of the Indo-Pacific sergeant were able to withstand water temperature down to 16 °C. However, such thermal ability may vary among tropical fish families and even species. Visual surveys of tropical vagrants in south-eastern Australia revealed that populations of four damselfish species tolerated temperatures down to 17 °C, while two butterflyfish species populations tolerated water temperatures down to 19 °C (Figueira and Booth 2010). In corroboration with this, low sea surface temperatures (SSTs) at tropical/ temperate transition zones also support the prediction that tropical fishes may be able to withstand low winter-water temperatures. For example, at the Solitary Islands [northern New South Wales (NSW)], winter SSTs regularly drop to 16.6 °C (Malcolm et al. 2011). Despite this, these islands support a large breeding colony of the tropical Threespot damselfish (Dascyllus trimaculatus, Pomacentridae) (HA Malcolm pers comm), while more than 50% of the species within the Solitary Islands are regarded as tropical (Malcolm et al. 2010). Population density and latitudinal distribution The likelihood of tropical fish larvae dispersal into temperate regions is likely to be associated with population abundance: that is, species that are extremely rare at the current latitudinal extremes of their distribution, or may be non-reproductive at these limits, will ultimately have lower numbers of larvae available to disperse away from source locations than species that are exceedingly abundant. Although there is limited data on the rarity and/or fecundity of species throughout their range, we can potentially use species population abundance at southern limits of coral development as a proxy for these processes. We can expect, therefore, that tropical vagrants may be among those with the highest population abundance on tropical reefs. In comparison, species that are relatively rare, or locally uncommon, are unlikely to have high numbers of larvae exported to reefs distant from natal sources (Jones et al. 2002). One of the most common macro-ecological patterns reported is a positive correlation between local abundance of a species and geographical range (Lawton 1999; Roughgarden 2009). However, there is little evidence to suggest that tropical species abundance is associated with geographical range, with numerous accounts of both spatially restricted and non-restricted fishes with little difference in total abundance (Jones et al. 2002; Hobbs et al. 2010, 2012). To examine whether the abundance of tropical reef fish species within the natal habitat [southern Great Barrier Reef (GBR)] was an important predictor of tropical vagrant occurrence within the adjacent temperate reef habitat (NSW), we used a logistic regression analysis and compared the total abundance of 24 butterflyfishes, 19 surgeonfishes and 42 damselfishes surveyed within the Swain sector (southern GBR) with the total abundance of these same species observed within NSW over the same 5-year period (2003, 2004, 2005, 2007 and 2009). This analysis found that species abundance in the southern GBR was not significantly associated with species abundance in NSW (P = 0.334). Overall, there was little similarity in tropical vagrant abundance between the Swain sector and NSW sites; although the most abundant tropical vagrant butterflyfishes (surveyed in NSW) showed some of the highest densities within the Swain group (i.e. Black butterflyfish (Chaetodon flavirostris, Chaetodontidae) and the Threadfin butterflyfish (Chaetodon auriga, Chaetodontidae)), one of the most abundant butterflyfish within the Swain sector (i.e. Blackback butterflyfish (Chaetodon melannotus, Chaetodontidae) was the least abundant species in tropical vagrant surveys within NSW. Tropical reef fish populations may be widely distributed, encompassing the entire latitudinal extent of coral reefs. Conversely, they may have truncated distributions, being predominantly found in a narrow band near the equator or occurring only in high-latitude regions (Choat and Robertson © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 599 Tropical vagrant fishes D A Feary et al, 2002; Jones et al. 2002; Emslie et al. 2010, 2012; Cheal et al. 2012). Such differences in the latitudinal extent of coral reef fishes may be an important predictor in understanding vagrancy within coral reef fishes, and we found latitudinal extent to be significantly associated with vagrants (P = 0.000). Indeed, there is increasing interest in understanding the mechanisms responsible for Rapoport’s rule, the increase in the latitudinal extents of occurrence of species towards higher latitudes (Stevens 1989). Although there is evidence to suggest that species upper thermal limits may show a geographical variation (Stillman and Somero 2000), recent work has shown that lower lethal temperatures decline with latitude (Gaston et al. 1998, Gaston and Chown 1999, Addo-Bedaiko et al. 2000). Therefore, although there is little data to examine this (see Eme and Bennett 2008; Figueira et al. 2009), we can predict that differences between vagrant and non-vagrant fishes in their latitudinal extent (i.e. abundance in tropical/temperate transition region) may be associated with a wider thermal tolerance for lower temperatures within vagrants (Addo-Bedaiko et al. 2000). However, more important in predicting the extent of vagrancy among coral reef fishes may be the geographical distance between a population’s tropical source and its temperate sink (Sorte et al. 2010). We can expect that species with distributions truncated in low latitudes are unlikely to form vagrant communities, while those with distributions that are closer to temperate regions are more likely to form them (Munday et al. 2008a). To test the role of latitude, and therefore distance from source to sink in structuring tropical vagrant communities, we examined the proportional abundance of butterflyfishes (as a percentage of total abundance) surveyed in both high-latitude (Capricorn Bunker group) and low-latitude (Lizard Island/Cooktown Sector) regions (using data from Emslie et al. 2010) and compared it with butterflyfishes surveyed throughout NSW (sensu Booth et al. 2007) (Table 1). Tropical vagrants surveyed in NSW were more likely to show higher abundances in high-, than low-latitude regions (Table 1). Of the seven butterflyfish species surveyed within temperate NSW waters, five species (Sunburst butterflyfish (Chaetodon kleinii, Chaetodontidae), Black butterflyfish, Speckled butterflyfish (C. citrinellus, Chaetodontidae), Raccoon butterflyfish (Chaetodon lunula, Chaetodontidae) and Blackback butterflyfish) were predominantly 600 the most important butterflyfish species (as a percentage of total abundance) surveyed in the highlatitude region. In comparison, the majority of tropical vagrants were not abundant in the lowlatitude region, although the Threadfin butterflyfish and Raccoon butterflyfish were relatively important (Table 1). In corroboration with this, recent research has shown that there is very little exchange in surface circulation between the northern (north of 18°S) and southern GBR (Choukroun et al. 2010), further supporting the potential importance of high-latitude populations as sources for tropical vagrants. Life-history traits associated with vagrancy Among species, a positive relationship has been shown between pelagic larval duration (PLD) and dispersal distance (e.g. Shanks 2009). We may, therefore, expect taxa with long PLDs to be more prevalent in expatriated larval assemblages, especially further from tropical sources (Booth and Parkinson 2011). However, this prediction may be overly simple as the phylogeographical literature has shown that reef fish have a great deal of flexibility in their larval dispersal characteristics. For example, recent work strongly suggests dual strategies in many tropical fish species: that is, localized dispersal that ensures retention in the immediate natal area and long-distance dispersal to geographically distant areas (Jones et al. 1999, 2005, 2009b; Planes et al. 2009). In addition, dispersal distance has also been shown to be independent of PLD for numerous species in multiple locations; there are many fish species with relatively low PLDs, which regularly cross the East Pacific barrier each way (c. 5000 km of deep water separating the Eastern from the Central Pacific) (Lessios and Robertson 2006; Leis et al. 2011), while others with relatively high PLDs can show exceptionally high levels of self-recruitment (i.e. Vagabond butterflyfish (Chaetodon vagabundus, Chaetodontidae) Almany et al. 2007a). Despite this, taxa with relatively longer PLDs (e.g. Chaetodontidae) were relatively more common than taxa with relatively shorter PLDs (e.g. Pomacentridae) among a suite of tropical vagrant species in south-eastern Australia (Booth et al. 2007). To examine whether PLD is associated with successful long-distance dispersal of vagrants, where possible, we compared the mean PLD (in days) of tropical vagrant (n = 109) and tropical non-vagrant fishes © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 Tropical vagrant fishes D A Feary et al., (n = 214). We found no relationship between vagrants and mean PLD, with both high and low PLD values found for both vagrant and nonvagrant species (P = 0.782). Life-history theory predicts relationships between numerous life-history traits and body size of fishes (Hutchings et al. 2012). Among these is a positive relationship between body size and lifetime reproductive output, with larger fishes producing far more gametes and higher lifetime reproductive output than smaller-bodied fishes (Weatherly 1972; Thresher 1984). We can expect then that such size-mediated differences in reproductive output may substantially increase the probability of Table 1 Distribution of Chaetodontidae as a percentage of total abundance between Lizard Island/Cooktown and Capricorn Bunker sectors (Emslie et al. 2010). Species in bold denote species that have been surveyed within NSW waters (sensu Booth et al. 2007). Species Lizard Island/ Cooktown Sector Species Swain sector H. polylepis C. meyeri C. reticulatus C. ephippium C. auriga C. bennetti F. longirostris C. lunula F. flavissimus C. punctatofasciatus C. pelewensis C. ulietensis C. vagabundus C. ornatissimus C. speculum C. lunulatus C. plebeius C. baronessa C. unimaculatus C. rafflesi C. aureofasciatus C. kleinii C. trifascialis C. lineolatus C. mertensii C. melannotus C. rostratus C. rainfordi C. citrinellus C. flavirostris 83 73.9 72.4 70.1 57.5 52.6 51.3 49 41.7 40.7 40.1 38.9 38.3 35.8 33.6 33 25 23.3 22.7 20.1 19.8 17.1 15.8 14.6 14.3 13.1 11.6 7.5 4.9 0.8 C. trifascialis C. kleinii C. flavirostris C. citrinellus C. lunula C. melannotus C. rainfordi C. unimaculatus C. reticulatus F. flavissimus C. speculum C. ornatissimus C. plebeius F. longirostris C. lineolatus C. pelewensis C. lunulatus C. auriga C. vagabundus C. baronessa C. ephippium C. rafflesi C. aureofasciatus C. mertensii C. rostratus H. polylepis C. ulietensis C. punctatofasciatus C. meyeri C. bennetti 63.6 49.5 38.8 26.6 16.3 15.3 15.1 13.2 12.2 11.4 11.2 10.2 8.6 7.7 7.5 6.2 6 5.4 3.4 2.5 1.2 0.9 0.1 <0.1 0 0 0 0 0 0 long-distance larval dispersal (Leis 1993; Munday and Jones 1998). Therefore, to determine whether body size is associated with vagrants, where possible, we compared the maximum body size (cm TL) of vagrant vs. non-vagrant tropical fishes (n = 341 vagrants, n = 4060 non-vagrants). Logistic regression of vagrant potential as a function of body size illustrates that fishes with larger body size are significantly more likely to have expatriated larval assemblages than smaller-sized fishes (P = 0.000). In fact, there is a threshold at which species are more likely to show vagrancy: within tropical fishes with body size larger than 10.95 cm TL, 12% are tropical vagrants, while within fishes smaller than 10.95 cm TL, only 0.02% are vagrants (Fig. 3). Coral reef fishes vary in the degree of parental care invested in their offspring. Broadcast spawners release gametes into the water column with very little investment, while demersal spawners invest vast resources into building and defending nest sites where eggs are laid and protected until hatching (Thresher 1984). We can expect that such differences in spawning mode may influence larval dispersal (Thresher 1984; Brogan 1994; LoYat et al. 2006). Pelagically spawned larvae are immediately subject to oceanographic processes, which may disperse them further than demersally spawned larvae (Thresher 1984). However, demersally spawned larvae have longer incubation times, are larger-bodied and have higher developed sensory and locomotor systems (Thresher 1984; Figure 3 We identified the threshold of body size effect on vagrancy by fitting a binary recursive partitioning, which splits the data successively and selects the split that maximally distinguishes the response variable above and below a given value. © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 601 Tropical vagrant fishes D A Feary et al, Leis 1993). Therefore, demersally spawned larvae may have a greater ability to control their position within the water column, thereby influencing the potential for dispersal (Leis and Goldman 1984; Leis 1993; Brogan 1994; Lo-Yat et al. 2006). To examine whether spawning mode is associated with vagrancy, we compared the reproductive strategies of 4665 species of tropical fish (n = 330 vagrant species, n = 4335 non-vagrant species) and classified them into one of three distinct reproductive guilds, each exhibiting different levels of parental care: (i) non-guarders (low care: pelagic spawner), (ii) guarders (moderate care: guarding and caring for eggs spawned into a demersal nest) and (iii) bearers (high care: brooding). This analysis showed that both high and medium levels of parental care were negatively associated with vagrant potential, while low levels of parental care were positively associated with vagrant potential (Fig. 4). Species-specific differences in larval swimming behaviour have been implicated in the dynamics of dispersal within a range of reef fish larvae (Stobutzki and Bellwood 1994, 1997; Leis et al. 1996; Leis and Carson-Ewart 1997, 2003; Fisher 2005). Although there is considerable variation among taxa, recent work on late-stage, or settlement-competent, larvae of coral reef fishes shows them to be outstanding swimmers both in terms of swimming speed and endurance (Fisher 2005; Fisher et al. 2005; Leis et al. 2011). Although species-specific differences in swimming ability may be important Figure 4 Vagrant potential as a function of parental care type (See Supplementary Data S2). 602 for retention of fishes within natal waters (Leis et al. 2011), such differences may also be associated with potential long-distance dispersal. If swimming ability is associated with potential dispersal, we can expect tropical vagrants to have greater larval swimming ability than non-vagrants. Therefore, we compared the potential swimming ability of tropical vagrants vs. non-vagrants using (Ucrit), the average critical speed (cms 1) as a proxy for swimming ability. Ucrit data were available for a total of 31 tropical vagrants vs. 32 non-vagrants, mostly from the Pomacentridae, Apogonidae and Chaetodontidae (using Fisher et al. 2005; Hogan et al. 2007; Leis et al. 2011). A logistic regression of vagrant potential as a function of average critical speed indicated that fishes with higher swimming ability were more likely to show vagrancy than those with lower swimming ability (P = 0.015). There is a substantial literature showing that factors operating within the early life history of coral reef fishes (e.g. feeding, growth rate, size) may have substantial flow-on effects to juvenile growth and ultimately survival (Brunton and Booth 2003; Hoey and McCormick 2004; McCormick and Hoey 2004). One of the most important factors in determining the survival of early history stages of coral reef fishes is size at settlement, as survival and longevity of new recruits are positively associated with increasing new settler body size (Sponaugle and Grorud-Covert 2006; Sponaugle et al. 2011). We can expect, therefore, that tropical vagrants may have larger size at settlement than non-vagrants. Therefore, to examine whether body size at settlement is an important predictor of the diversity of tropical vagrant fishes, we compared the size at settlement of tropical vagrant and non-vagrant damselfishes (using Kerrigan 1996; Thresher and Brothers 1989; Wellington and Robertson 2001; Wellington and Victor 1989). Logistic regression of vagrant potential as a function of size at settlement illustrated that fishes with larger size at settlement were more likely to have expatriated larval assemblages than species with smaller-sized settlers (P = 0.045). In fact, of the 51 species of damselfish for which average size-at-settlement data were available (n = 15 tropical vagrants, n = 36 tropical nonvagrants), seven species (Bicolor chromis (Chromis margaritifer, Pomacentridae), Nagasaki damsel (Pomacentrus nagasakiensis, Pomacentridae), Sapphire damsel (Pomacentrus pavo, Pomacentridae), © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 Tropical vagrant fishes D A Feary et al., Speckled damselfish, (Pomacentrus bankanensis, Pomacentridae), Ward’s damsel (Pomacentrus wardi, Pomacentridae), Neon damsel (Pomacentrus coelestis, Pomacentridae) and Yellowtail demoiselle (Neopomacentrus azysron, Pomacentridae)) settled between 13 and 15 mm TL, which was in the top 17% of values for settlement size. In comparison, the majority of non-vagrant Pomacentrid species larvae settled at sizes below 13 mm TL. Correlations between species-specific traits in predicting vagrancy Within this review, we have focused on examining whether single species-specific traits may be useful in predicting vagrant potential. However, there is also the need to examine whether there are correlations between traits. For example, temperature is a major factor determining PLD (O’Connor et al. 2007) and latitudinal range, thus confounding single analyses looking at either of these two. In addition, body size may co-vary with abiotic factors (e.g. temperature); warmer water species tend to have smaller body size than colder water species because of oxygen and thermal capacity limitation (Pauly 1997). However, when undertaking correlation analyses encompassing a number of species traits, there are several caveats that need to be addressed. Firstly, although coral reef fishes are among the best-studied teleost assemblages, there remain considerable gaps in our knowledge of their biology and ecology (Pratchett et al. 2008b). Therefore, there will be always limited information on specific traits within this fauna. For example, when comparing the Ucrit values between vagrant and non-vagrant fishes, our analyses comprised 31 species of vagrant fishes (encompassing 9 families) and 32 non-vagrant fishes (encompassing nine families). Secondly, there is a substantial bias in the phylogenetic extent of research undertaken on coral reef fish communities, with the butterflyfishes and damselfishes the most well-researched groups (Sale 1991, 2002). Therefore, for any particular trait, the potential analysis between vagrants and non-vagrants may be affected by the limited phylogenetic extent of the sample; the potential for a phylogenetic confound is then potentially high. Despite these caveats, the need to develop multiple trait correlations will be important in developing the range of traits that will aid in predicting vagrancy within coral reef fishes. Therefore, we undertook a logistic regression using three traits in which there was sufficient data available: latitudinal range, body size and reproductive mode [all data sourced from Fishbase (Froese and Pauly 2012)]. For this analysis, there were 290 species (encapsulating surgeonfishes, butterflyfishes and damselfishes, 97 species were vagrants) for which values for all three traits were available (with ‘family’ as a random factor in the binomial GLMM). This analysis showed that only latitudinal range was significant in predicting vagrancy (P <0.000, z = 6.307). We then extended this analysis and included all data available on PLD (which reduced the available dataset to 129 species, 61 species were vagrants). As PLD and reproductive mode were highly correlated (Spearman’s correlation = 0.8), we excluded reproductive mode from this analysis. However, the results also showed that latitudinal range was the only significant factor that predicts vagrancy (P <0.000, z = 3.735). Therefore, although such correlative analyses at present are limited by available data, and potentially have a phylogenetic bias, they will be useful in developing a predictive framework for understanding vagrancy within tropical reef fishes. We have shown that the present latitudinal range of tropical reef fishes may be a suitable predictor of vagrancy: the higher the latitudinal range, the more likely that the species will have expatriated larval assemblages. Resource constraints to vagrant success We can expect that tropical range shifts are likely to be limited by species-specific resource requirements (Munday et al. 2008a; Cheung et al. 2010). In particular, for tropical fishes, temperate reefs will lack a range of settlement substrates, settlement cues or specific dietary components found on tropical coral reefs (Harriott and Banks 2002). Temperate reef habitats are not devoid of scleractinian corals (Rodolfo-Metalpa et al. 2008; Lien et al. 2012), and it is possible that tropical reefbuilding corals may become increasingly established beyond their normal latitudinal limits due to sustained increases in ocean temperature (Precht and Aronson 2004; Greenstein and Pandolfi 2008; Yamano et al. 2011). Until this happens, however, it is likely that the lack of tropical corals, the resources associated with a coral reef or the types of habitat available within a coral reef will significantly limit the successful recruitment and thereby the range extensions of coral reef fishes into temperate environments. © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 603 Tropical vagrant fishes D A Feary et al, Approximately 10% of coral reef fishes can be classified as coral dependent at some part of their life stage (Pratchett et al. 2008b). These fishes include obligate coral settlers or dwellers (Munday et al. 1997; Gardiner and Jones 2005; Feary et al. 2007b) and corallivores (Pratchett 2005). As reductions in coral cover nearly always cause corresponding declines in the abundances of such coral-dependent species (Wilson et al. 2006; Feary et al. 2007a; Emslie et al. 2011), we predict that reliance on coral resources is likely to constrain the shifts of these species into temperate reef habitats (Fig. 5). In addition, some reef fish families as a whole are more reliant on live coral cover than others, with a higher proportion of species in diverse reef fish families such as the butterflyfish (Chaetodontidae), cardinalfish (Apogonidae) and gobies (Gobiidae) known to be closely associated with live coral cover (Pratchett 2005; Pratchett et al. 2008b). If the degree of live coral cover is important in structuring settlement, or early survival of these families, we can expect these families to show the lowest levels of abundance and diversity within temperate reef habitats. Habitat association and settlement preferences Proportional abundance We can predict that the availability of specific coral types at settlement will not be the only 90 GBR 80 NSW 70 60 50 40 30 20 10 0 Low Med High Coral association Figure 5 Coral habitat association between tropical vagrants and tropical non-vagrants (using Wilson et al. 2006; Froese and Pauly 2012) comparing fishes surveyed within the Swain sector within the Australian Institute of Marine Science Long Term Monitoring Project (AIMS LTMP unpublished data) and tropical vagrant surveyed throughout NSW (D.J. Booth, unpublished data). Coral habitat association is divided into three categories: (i) those that settle, dwell or feed on the reef (‘high’ association), (ii) those that are associated with the reef structure (‘medium’ association) and (iii) those that are not associated with the reef (‘low’ association). 604 factor limiting the success of tropical vagrants in colonizing temperate reef habitats. There are a range of factors, independent of live coral cover, which can be used to potentially predict the successful settlement and recruitment of tropical reef fish assemblages (Caley et al. 1996; Booth and Wellington 1998; Leis and McCormick 2002). These factors include, but are not limited to, the availability of suitable trophic resources (Booth and Hixon 1999), habitat complexity (Holbrook et al. 2002b), prior resident density (Sweatman 1988; Booth 2004) and the composition of predator assemblages (Beukers-Stewart and Jones 2004; Beukers-Stewart et al. 2011). Understanding the role of such factors in structuring species-specific settlement in tropical assemblages will be vital in predicting potential vagrant species. Tropical species that show distinct settlement preferences (i.e. settlement habitat specialists) would be expected to be much more limited in their ability to utilize habitats and environments beyond their normal latitudinal limits, compared with more generalized settlement resource use (i.e. settlement habitat generalists). For example, within a range of group-forming planktivorous damselfishes (e.g. Dascyllus complex), specific habitats (i.e. structurally complex corymbose corals) are fundamental to successful individual settlement (Holbrook et al. 2002a,b) and post-settlement survival (Holbrook and Schmitt 2002). For these species, we can predict that the lack of these specific habitats may result in low or non-existent settlement of this species complex within temperate reef environments. In support, there is little evidence to suggest that any of the Dascyllus species found within the GBR [i.e. Whitetail dascyllus (Dascyllus aruanus, Pomacentridae), Blacktail humbug (Dascyllus melanurus, Pomacentridae) and Threespot dascyllus (Dascyllus trimaculatus, Pomacentridae)] settle and survive in habitats within temperate NSW (Booth et al. 2007), despite being found in relatively high abundances throughout the southern limits of the GBR and within subtropical coral reef habitats in northern NSW (Scott and Harrison 2008). Morphologically, temperate benthic reef systems are substantially different from their tropical counterparts (Ebeling and Hixon 1991; Kingsford and Battershill 1998), and we can predict that structural differences in benthic communities between temperate and tropical ecosystems will substantially affect the composition of tropical vagrants © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 Tropical vagrant fishes D A Feary et al., settling in temperate reefs. Shallow temperate reefs, especially within south-east Australia, form a mosaic of habitats, dominated by large stands of laminarian and fucoid algae (e.g. Ecklonia radiata, Sargassum species) and relatively barren rocky substrata with low coralline turfing or crustose algal populations (termed ‘urchin barrens’) (Underwood et al. 1991). We may predict that tropical vagrant larvae may show avoidance behaviour towards habitats dominated by macroalgae, due to both the physical movement of such habitats associated with wave action and relatively low levels of topographical complexity (and therefore fine-scale microshelter for new settlers) (Kingsford and Carlson 2010). In fact, recent evidence suggests that settlement of tropical vagrants is closely associated with urchin barren habitats, specifically within finescale cracks and crevices where urchins have cleared algae (H.J. Beck, unpublished data). Such habitats are also preferred settlement habitats within a range of temperate reef fish species (Kingsford and Carlson 2010). Dietary preferences and functional groups Although the availability of habitat resources may be important in constraining vagrant success within temperate systems, resource requirements will be further constrained by specific dietary requirements. Obligate coral-feeding butterflyfishes, for example, will be unlikely to recruit into habitats devoid of extensive cover of preferred coral species (Pratchett et al. 2008a). Accordingly, the overwhelming majority of Chaetodon butterflyfishes recorded in surveys of vagrant fishes along the NSW coast are non-coral or facultative coral feeders, including non-coral benthic invertebrate feeders (Threadfin butterflyfish, Crochet butterflyfish (C. guentheri, Chaetodontidae), Vagabond butterflyfish), soft coral specialists (Blackback butterflyfish) and generalist benthic foragers (Speckled butterflyfish, Black butterflyfish, Sunburst butterflyfish and Raccoon butterflyfish). Obligate coral-feeding butterflyfishes have been sighted (e.g. Blueblotch butterflyfish (Chaetodon plebeius, Chaetodontidae)); however, they are exceptionally rare within surveys (D.J. Booth, unpublished database) despite their high relative abundance in the southern GBR (i.e. Swain sector). This suggests that the availability of specific prey will strongly constrain range extensions for highly specialized reef fishes, but three mechanisms may occur: these fishes may have limited larval dispersal to reefs devoid of coral, or not settle on these reefs, or experience rapid early post-settlement mortality if settlement occurs on reefs devoid of coral. For some tropical reef fish species, differences in the availability of specific prey resources may not necessarily constrain settlement and immediate survivorship (Booth et al. 2007; Figueira et al. 2009), but the lack or limited availability of certain resources may lead to marked physiological changes (e.g. growth and development) or reduced fitness. For example, Pratchett et al. (2004) found little change in the population abundance of an obligate coral-feeding butterflyfish, the Oval butterflyfish (Chaetodon lunulatus, Chaetodontidae), two years after extensive coral loss on reefs in the central GBR. However, there were significant declines in the physiological condition of populations, likely to have resulted from declines in the quantity and quality of available coral prey (Pratchett et al. 2004). Likewise, Feary et al. (2009) found little effect on individual persistence following experimental coral loss within groups of two planktivorous damselfish species (Goldtail demoiselle (Chrysiptera parasema, Pomacentridae) and Blacktail humbug). However, growth rates of both species (quantified over a 29-day period) were directly related to percentage live coral cover; individuals within colonies with reduced live coral exhibited slower morphometric growth than those within colonies with high live coral cover (Feary et al. 2009). Therefore, although tropical fishes may be able to successfully settle and persist within suitable temperate reef habitats (Booth et al. 2007; Figueira et al. 2009), the full impact on each species population following such settlement may take months to years to become apparent. Thus, instead of an immediate reduction in the abundance of tropical vagrants following settlement, populations may maintain their numbers over a relatively prolonged time period (Choat et al. 1988). For example, the Lord Howe Island butterflyfish (Amphichaetodon howensis, Chaetodontidae) is a relatively common subtropical species found within the northern islands of the Poor Knights Islands, New Zealand. Although paired individuals of the Lord Howe Island butterflyfish have been consistently surveyed since the early 1970s (e.g. Russell 1971), there is still no evidence to suggest species replenishment is associated with local reproduction. Rather, population replenishment appears to be due solely to the © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 605 Tropical vagrant fishes D A Feary et al, continual immigration of larvae from tropical and subtropical locations. Section 3: Future research needs Aside from established topics of research on traits that may facilitate or impede range shifts of tropical reef fish species (e.g. larval biology and ecology, post-settlement resource use), there are several other key areas of future research needed to understand range shifts among tropical species. This is predominantly because tropical fishes provide some of the most extreme examples in geographical range expansion (Booth et al. 2007; Figueira et al. 2009) compared with other perciform fishes. The most important topics, addressed in turn, are behavioural ecology and biomechanics, habitat use of subtropical reefs, physiology, predation and competition with temperate residents. This is by no means a comprehensive list of potential future research topics, but we present these topics with the hope of stimulating even more research on tropical vagrant fishes. There is still little understanding of the importance of subtropical (i.e. marginal) coral reefs in providing a ‘habitat refuge’ from the impacts of climate change for coral reef fish communities. Although this review has focused on temperate reef habitats, there is considerable overlap in tropical and subtropical reef fish community structure (Malcolm et al. 2007, Malcolm et al. 2010). We may expect that tropical vagrants that successfully utilize subtropical reef systems (i.e. are able to survive and reproduce) may be more likely to settle and survive within temperate reef systems (see Section 2). To date, research on the use of subtropical reefs by tropical vagrants has predominantly focused on how the structure and composition of these populations vary between tropical and subtropical reef systems. Such studies have shown that while there is considerable overlap in community structure between these systems, marginal reefs typically have lower abundance and, more notably, lower diversity than tropical reef systems (Feary et al. 2010; Pratchett et al. 2012). Such differences in community structure may be partly associated with subtropical reefs having greater seasonal variation in oceanographic factors than their lower-latitude counterparts (Riegl et al. 2011; Feary et al. 2012). In particular, seasonal variations in seawater temperature of up to 12 °C are relatively common on 606 subtropical and high-latitude reefs (i.e. Japan: 16– 28 °C; Lord Howe Island: 17–27 °C; Gulf of Oman: 22–32 °C). Therefore, understanding how such extremes in oceanographic variables affect the resource acquisition (i.e. feeding rates, resource use and competition) (Pratchett et al. 2012), and energy demands and physiology (e.g. metabolic rates and swimming abilities) of tropical fishes on such reefs may not only provide a mechanistic basis for the factors that structure tropical fish communities on these marginal reefs, but will provide important insights into how reef fish families may acclimate or adapt to environmentally extreme ecosystems. Successful establishment of certain tropical reef fishes within temperate regions may be associated with specific dietary requirements (see Section 2 above). For tropical herbivorous fishes, the often high abundance of micro- and macroalgae within temperate reef habitats means that ‘food’ availability per se may not necessarily constrain survivorship (Booth et al. 2007; Figueira et al. 2009). However, the lack of, or limited availability of, specific tropical trophic resources (i.e. warm-water algal species with which the vagrants have historically co-occurred) may lead to marked physiological reductions in fishes condition (Pratchett et al. 2004). In particular, we may expect that for tropical fishes evolved to utilize specific tropical algae, the use of temperate algal species may affect their physiological condition. This may happen directly through novel chemical defences of the new foods or reduction in assimilation rates, or it may be mediated by changes in gut microflora, a key consideration for almost all herbivorous consumers, including humans (e.g. recent studies of the human gut ‘microbiome’: Nicholson et al. 2012). An established gut microbiota is essential for the healthy physiological and immunological development of most animals, with changes in biota having substantial effects on nutrient utilization and therefore physiological performance. Therefore, although tropical fishes may be transported to and successfully settle within temperate habitats (Booth et al. 2007; Figueira et al. 2009), to persist they must be able to consume dietary resources that are suboptimal or novel. Species-specific differences in tropic ecology within these habitats (i.e. associated with both feeding and nutrient assimilation) may then have substantial effects on population persistence and, ultimately, success of these vagrants. © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 Tropical vagrant fishes D A Feary et al., Predation is a major source of mortality for early life stages of tropical fish (Hixon and Beets 1993; Planes and Lecaillon 2001; Doherty et al. 2004), and minimizing lethal interactions with predators is critical to successful population establishment in temperate areas. Predation within the first 7 days accounts for up to 78% mortality of recently settled juveniles in tropical reef fishes (Doherty and Sale 1986; Victor 1986; McCormick 1998), and mortality during this period is likely to have a disproportionate effect on the size of tropical populations (critical period hypothesis, sensu Suthers 1998). Therefore, we can expect that tropical fish species that are relatively successful at avoiding temperate predators, both during and immediately following settlement, are more likely to survive in temperate systems. This will be important given that for tropical vagrants, all temperate ‘predators’ within temperate environments will be novel, with the potential for substantial predation risk within temperate environments. However, the traits that may influence predation risk of juvenile tropical reef fish still remain poorly understood (McCormick 1998). Selective mortality has been linked to variation in numerous morphological and physiological traits (Hixon 1991, 2011). Although contradictory evidence exists (Holmes and McCormick 2009, 2010), numerous studies have reported enhanced survival of recruits and juveniles that were larger for a given age (Sponaugle and Grorud-Covert 2006; Sponaugle et al. 2011). Tropical vagrant species with larger size at settlement may therefore be more likely to succeed within temperate environments. To test whether size at settlement differs between tropical vagrants and temperate residents, where possible we compared the log mean size at settlement of tropical vagrant (n = 15) and temperate residents (n = 41). We found that there was no substantial difference between groups; log mean size at settlement (mm) (95% CI) for tropical vagrants was 2.28 (0.055), while the log mean size at settlement for temperate residents was 2.47 (0.065). Predator recognition by juvenile reef fish is a vital factor affecting the outcome of predator–prey interactions (Almany and Webster 2004; Almany et al. 2007b), with higher mortality observed in individuals with reduced ability to identify potential threats (McCormick and Holmes 2006; Loennstedt et al. 2012). The majority of research focusing on tropical fishes has examined the effect of prior experience on subsequent predator recognition (learned responses) and has indicated that individuals previously exposed to visual and chemical cues associated with predators have a greater chance than naive individuals of surviving predatory encounters (Lonnstedt et al. 2012; Manassa and McCormick 2012). However, the likelihood that initial (non-learned) abilities to recognize predators may vary among tropical vagrants suggests that predation risk during initial predatory encounters will also substantially differ among species settling in temperate environments. Although based on a relatively small subset of tropical vagrant species, recent work has shown that there are substantial changes in the behaviour of tropical species when exposed to temperate predators, including reduced feeding rates and increased sheltering (H.J. Beck, unpublished data). We predict that tropical vagrants that are initially better at recognizing temperate predators, or learn to recognize them more rapidly, may experience reduced predation risk during settlement and early recruitment and are able to successfully establish temperate populations. Despite the increasing abundance of tropical fish species within temperate reef systems, little is known of their competitive abilities in temperate environments. Recent evidence suggests that relative activity levels and feeding rates of tropical range-shifting species compared with temperate residents are dependent on habitat availability and temperature fluctuations (H.J. Beck, unpublished data). Therefore, research into competitive interactions between temperate and tropical species and how these interactions may be influenced and mediated by temperature and habitat availability is necessary. As space limitation is important in determining juvenile abundance in numerous siteattached tropical fish species (Hixon and Beets 1989; Munday et al. 2001; Holbrook and Schmitt 2002; Bonin et al. 2009), we hypothesize that competition for temperate macroalgal resources both during and following settlement may have substantial consequences for the abundance and diversity of tropical range-shifting species within temperate ecosystems and the potential impact on the temperate resident fishes. Conclusions An understanding of both the extrinsic and intrinsic processes that are likely to influence the spatial, temporal and taxonomic biases in tropical © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 607 Tropical vagrant fishes D A Feary et al, vagrant community structure will be vital in predicting the repercussions for tropical fishes with increased changes in global climate. It can be assumed that extrinsic processes (e.g. currents, environmental temperature) will interact with inherent differences in the life histories of fishes to determine what species settle into temperate regions, when these species settle and where. However, intrinsic processes constraining the movement and population success of tropical vagrants are key to understanding which species will move and which will not and which species will become susceptible to sustained and ongoing climate change. This review has shown that tropical species with viable populations near the latitudinal margins of reef development (high-latitude reefs) will be expected to show substantial changes in distribution. In this respect, we can also expect that adaptation to changes in local water temperature may occur more rapidly in small-bodied, short-lived species, where selection can operate over a large number of generations. Larger species, with much longer generation times, may have reduced ability to adapt to temperate regions. To our knowledge, successful reproduction in tropical vagrants has not occurred within temperate regions, with all observed population expansion driven solely by larval input from tropical sources (however, see Kokita 2004). However, with increasing warming of waters, we can expect that the physical variables constraining reproduction will reduce (i.e. increased growth rates in individuals, development of reproductively active individuals) and viable breeding populations of tropical vagrants will increase within temperate regions. Acknowledgements DAF was funded by the University of Technology, Sydney, under the Chancellors Postdoctoral Fellowship scheme. OJL was supported by a doctoral fellowship grant provided by the Sydney Institute of Marine Science. Thanks to Nicolas Bailly (Fishbase) for providing data. This paper is contribution 87 from the Sydney Institute of Marine Science. References Abesamis, R.A. and Russ, G.R. (2010) Patterns of recruitment of coral reef fishes in a monsoonal environment. Coral Reefs 29, 911–921. 608 Addo-Bediako, A., Chown, S.L. and Gaston, K.J. (2000) Thermal tolerance, climatic variability and latitude. Proceedings of the Royal Society London B 267, 739–745. Almany, G.R. and Webster, M.S. (2004) Odd species out as predators reduce diversity of coral-reef fishes. Ecology 85, 2933–2937. Almany, G.R., Berumen, M.L., Thorrold, S.R., Planes, S. and Jones, G.P. (2007a) Local replenishment of coral reef fish populations in a marine reserve. Science 316, 742–744. Almany, G.R., Peacock, L.F., Syms, C., McCormick, M.I. and Jones, G.P. (2007b) Predators target rare prey in coral reef fish assemblages. Oecologia 152, 751–761. Attrill, M.J. and Power, M. (2002) Climatic influence on a marine fish assemblage. Nature 417, 275–278. Beukers-Stewart, B.D. and Jones, G.P. (2004) The influence of prey abundance on the feeding ecology of two piscivorous species of coral reef fish. Journal of Experimental Marine Biology and Ecology 299, 155–184. Beukers-Stewart, B.D., Beukers-Stewart, J.S. and Jones, G.P. (2011) Behavioural and developmental responses of predatory coral reef fish to variation in the abundance of prey. Coral Reefs 30, 855–864. Bonin, M.C., Srinivasan, M., Almany, G.R. and Jones, G.P. (2009) Interactive effects of interspecific competition and microhabitat on early post-settlement survival in a coral reef fish. Coral Reefs 28, 265–274. Booth, D.J. (2004) Synergistic effects of conspecifics and food on growth and energy allocation of a damselfish. Ecology 85, 2881–2887. Booth, D.J. and Hixon, M.A. (1999) Food ration and condition affect early survival of the coral reef damselfish, Stegastes partitus. Oecologia 121, 364–368. Booth, D.J. and Parkinson, K. (2011) Pelagic larval duration is similar across 23A degrees of latitude for two species of butterflyfish (Chaetodontidae) in eastern Australia. Coral Reefs 30, 1071–1075. Booth, D.J. and Wellington, G. (1998) Settlement preferences in coral-reef fishes: effects on patterns of adult and juvenile distributions, individual fitness and population structure. Australian Journal of Ecology 23, 274– 279. Booth, D.J., Figueira, W.F., Gregson, M.A., Brown, L. and Beretta, G. (2007) Occurrence of tropical fishes in temperate southeastern Australia: role of the East Australian Current. Estuarine Coastal and Shelf Science 72, 102–114. Booth, D.J., Bond, N. and Macreadie, P. (2011) Detecting range shifts among Australian fishes in response to climate change. Marine and Freshwater Research 62, 1027–1042. Brogan, M.W. (1994) Distribution and retention of larval fishes near reefs in the Gulf of California. Marine Ecology Progress Series 115, 1–13. Brunton, B.J. and Booth, D.J. (2003) Density- and sizedependent mortality of a settling coral-reef damselfish © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 Tropical vagrant fishes D A Feary et al., (Pomacentrus moluccensis Bleeker). Oecologia 137, 377–384. Burrows, M.T., Schoeman, D.S., Buckley, L.B. et al. (2011) The pace of shifting climate in marine and terrestrial ecosystems. Science 334, 652–655. Caley, M.J., Carr, M.H., Hixon, M.A., Hughes, T.P., Jones, G.P. and Menge, B.A. (1996) Recruitment and the local dynamics of open marine populations. Annual Review of Ecology and Systematics 27, 477–500. Carnaval, A.C. and Moritz, C. (2008) Historical climate modelling predicts patterns of current biodiversity in the Brazilian Atlantic forest. Journal of Biogeography 35, 1187–1201. Cheal, A., Emslie, M., Miller, I. and Sweatman, H. (2012) The distribution of herbivorous fishes on the Great Barrier Reef. Marine Biology 159, 1143–1154. Cheung, W.W.L., Lam, V.W.Y., Sarmiento, J.L. et al. (2010) Large-scale redistribution of maximum fisheries catch potential in the global ocean under climate change. Global Change Biology 16, 24–35. Choat, J.H. and Robertson, D.R. (eds.) (2002) Age-Based Studies on Coral Reef Fishes. Academic Press, San Diego. Choat, J.H., Ayling, A.M. and Schiel, D.R. (1988) Temporal and spatial variation in an island fish fauna. Journal of Experimental Marine Biology and Ecology 121, 91–111. Choukroun, S., Ridd, P.V., Brinkman, R. and McKinna, L.I.W. (2010) On the surface circulation in the western Coral Sea and residence times in the Great Barrier Reef. Journal of Geophysical Research-Oceans 115, 1–13. Condie, S.A., Mansbridge, J.V. and Cahill, M.L. (2011) Contrasting local retention and cross-shore transports of the East Australian Current and the Leeuwin Current and their relative influences on the life histories of small pelagic fishes. Deep-Sea Research Part Ii-Topical Studies in Oceanography 58, 606–615. Cowen, R.K., Lwiza, K.M.M., Sponaugle, S., Paris, C.B. and Olson, D.B. (2000) Connectivity of marine populations: open or closed? Science 287, 857–859. Cowen, R.K., Paris, C.B. and Srinivasan, A. (2006) Scaling of connectivity in marine populations. Science 311, 522–527. Danilowicz, B.S. (1997) A potential mechanism for episodic recruitment of a coral reef fish. Ecology 78, 1415–1423. Davis, C.C., Bell, C.D., Mathews, S. and Donoghue, M.J. (2002) Laurasian migration explains Gondwanan disjunctions: evidence from Malpighiaceae. Proceedings of the National Academy of Sciences of the United States of America 99, 6833–6837. Dixson, D.L., Munday, P.L., Pratchett, M. and Jones, G.P. (2012) Ontogenetic changes in responses to settlement cues by Anemonefish. Coral Reefs 30, 903–910. Doherty, P.J. and Sale, P.F. (1986) Predation on juvenile coral reef fishes: an exclusion experiment. Coral Reefs 4, 225–234. Doherty, P.J., Dufour, V., Galzin, R., Hixon, M.A., Meekan, M.G. and Planes, S. (2004) High mortality during settlement is a population bottleneck for a tropical surgeonfish. Ecology 85, 2422–2428. Domenici, P., Claireaux, G. and McKenzie, D.J. (2007) Environmental constraints upon locomotion and predator-prey interactions in aquatic organisms: an introduction. Philosophical Transactions of the Royal Society B-Biological Sciences 362, 1929–1936. Donelson, J.M., Munday, P.L., McCormick, M.I. and Nilsson, G.E. (2011) Acclimation to predicted ocean warming through developmental plasticity in a tropical reef fish. Global Change Biology 17, 1712–1719. Donelson, J.M., Munday, P.L., McCormick, M.I. and Pitcher, C.R. (2012) Rapid transgenerational acclimation of a tropical reef fish to climate change. Nature Climate Change 2, 30–32. Dulvy, N.K., Rogers, S.I., Jennings, S., Stelzenmuller, V., Dye, S.R. and Skjoldal, H.R. (2008) Climate change and deepening of the North Sea fish assemblage: a biotic indicator of warming seas. Journal of Applied Ecology 45, 1029–1039. Ebeling, A.W. and Hixon, M.A.. (1991) Tropical and temperate reef fishes: comparison of community structures. In: The Ecology of Fishes on Coral Reefs (ed. P.F. Sale). Academic Press, San Diego, CA, pp. 509– 563. Eme, J. and Bennett, W.A. (2008) Low temperature as a limiting factor for introduction and distribution of Indo-Pacific damselfishes in the eastern United States. Journal of Thermal Biology 33, 62–66. Emslie, M.J., Pratchett, M.S., Cheal, A.J. and Osborne, K. (2010) Great Barrier Reef butterflyfish community structure: the role of shelf position and benthic community type. Coral Reefs 29, 705–715. Emslie, M.J., Pratchett, M.S. and Cheal, A.J. (2011) Effects of different disturbance types on butterflyfish communities of Australia’s Great Barrier Reef. Coral Reefs 30, 461–471. Emslie, M.J., Logan, M., Ceccarelli, D.M. et al. (2012) Regional-scale variation in the distribution and abundance of farming damselfishes on Australia’s Great Barrier Reef. Marine Biology 159, 1293–1304. Feary, D.A., Almany, G.R., Jones, G.P. and McCormick, M.I. (2007a) Coral degradation and the structure of tropical reef fish communities. Marine Ecology Progress Series 333, 243–248. Feary, D.A., Almany, G.R., McCormick, M.I. and Jones, G.P. (2007b) Habitat choice, recruitment and the response of coral reef fishes to coral degradation. Oecologia 153, 727–737. Feary, D.A., Burt, J.A., Bauman, A.G., Usseglio, P., Sale, P.F. and Cavalcante, G.H. (2010) Fish communities on the world’s warmest reefs: what can they tell us about the effects of climate change in the future? Journal of Fish Biology 77, 1931–1947. © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 609 Tropical vagrant fishes D A Feary et al, Feary, D.A., Burt, J.A., Cavalcante, G.H. and Bauman, A.G.. (2012) Extreme physical factors and the structure of Gulf communities. In: Coral Reefs of the Gulf: Adaptation to Climatic Extremes (eds B. Riegl and S. Purkis). Springer, NL, pp. 163–170. Feary, D.A., McCormick, M.I. and Jones, G.P. (2009) Growth of reef fishes in response to live coral cover. Journal of Experimental Marine Biology and Ecology 373, 45–49. Figueira, W.F. and Booth, D.J. (2010) Increasing ocean temperatures allow tropical fishes to survive overwinter in temperate waters. Global Change Biology 16, 506–516. Figueira, W.F., Biro, P., Booth, D.J. and Valenzuela, V.C. (2009) Performance of tropical fish recruiting to temperate habitats: role of ambient temperature and implications of climate change. Marine Ecology Progress Series 384, 231–239. Fisher, R. (2005) Swimming speeds of larval coral reef fishes: impacts on self-recruitment and dispersal. Marine Ecology Progress Series 285, 223–232. Fisher, R., Bellwood, D.R. and Job, S.D. (2000) Development of swimming abilities in reef fish larvae. Marine Ecology Progress Series 202, 163–173. Fisher, R., Leis, J.M., Clark, D.L. and Wilson, S.K. (2005) Critical swimming speeds of late-stage coral reef fish larvae: variation within species, among species and between locations. Marine Biology 147, 1201–1212. Francis, M.P., Worthington, C.J., Saul, P. and Clements, K.D. (1999) New and rare tropical and subtropical fishes from northern New Zealand. New Zealand Journal of Marine and Freshwater Research 33, 571–586. Froese, R. and Pauly, D. (2012) FishBase. World Wide Web electronic publication. www.fishbase.org, version (08/2012). Gardiner, N. and Jones, G.P. (2005) Habitat specialisation and overlap in a guild of coral reef cardinalfish (family Apogonidae). Marine Ecology Progress Series 305, 163–175. Gaston, K.J. and Chown, S.L. (1999) Elevation and climatic tolerance: a test using dung beetles. Oikos 86, 584–590. Gaston, K.J., Blackburn, T.M. and Spicer, J.I. (1998) Rapoport’s rule: time for an epitaph? Trends in Ecology and Evolution 13, 70–74. Greenstein, B.J. and Pandolfi, J.M. (2008) Escaping the heat: range shifts of reef coral taxa in coastal Western Australia. Global Change Biology 14, 513–528. Guisan, A. and Zimmermann, N.E. (2000) Predictive habitat distribution models in ecology. Ecological Modelling 135, 147–186. Hare, J.A. and Cowen, R.K. (1991) Expatriation of Xyrichtys novacula (Pisces, Labridae) larvae – evidence of rapid cross-slope exchange. Journal of Marine Research 49, 801–823. Hare, J.A. and Cowen, R.K. (1996) Transport mechanisms of larval and pelagic juvenile bluefish (Pomato- 610 mus saltatrix) from South Atlantic Bight spawning grounds to Middle Atlantic Bight nursery habitats. Limnology and Oceanography 41, 1264–1280. Hare, J.A., Churchill, J.H., Cowen, R.K. et al. (2002) Routes and rates of larval fish transport from the southeast to the northeast United States continental shelf. Limnology and Oceanography 47, 1774–1789. Harriott, V.J. and Banks, S.A. (2002) Latitudinal variation in coral communities in eastern Australia: a qualitative biophysical model of factors regulating coral reefs. Coral Reefs 21, 83–94. Harrison, H.B., Williamson, D.H., Evans, R.D. et al. (2012) Larval export from marine reserves and the recruitment benefit for fish and fisheries. Current Biology 22, 1023–1028. Hazel, J.R. and Prosser, C.L. (1974) Molecular mechanisms of temperature compensation in poikilotherms. Physiological Reviews 54, 620–677. Helmuth, B., Mieszkowska, N., Moore, P. and Hawkins, S.J. (2006) Living on the edge of two changing worlds: forecasting the responses of rocky intertidal ecosystems to climate change. Annual Review of Ecology Evolution and Systematics 37, 373–404. von Herbing, I.H. (2002) Effects of temperature on larval fish swimming performance: the importance of physics to physiology. Journal of Fish Biology 61, 865–876. Hickling, R., Roy, D.B., Hill, J.K., Fox, R. and Thomas, C.D. (2006) The distributions of a wide range of taxonomic groups are expanding polewards. Global Change Biology 12, 450–455. Hirata, T., Oguri, S., Hirata, S., Fukami, H., Nakamura, Y. and Yamaoka, K. (2011) Seasonal changes in fish assemblages in an area of hermatypic corals in Yokonami, Tosa Bay, Japan. Japanese Journal of Ichthyology 58, 49–64. Hixon, M.A. (1991) Predation as a process structuring coral-reef fish communities. In: The Ecology of Fishes on Coral Reefs. (ed. P.F. Sale). Academic Press, San Diego, CA, pp. 475–508. Hixon, M.A. (2011) 60 years of coral reef fish ecology: past, present, future. Bulletin of Marine Science 87, 727–765. Hixon, M.A. and Beets, J.P. (1989) Shelter characteristics and Caribbean fish assemblages – experiments with artificial reefs. Bulletin of Marine Science 44, 666–680. Hixon, M.A. and Beets, J.P. (1993) Predation, prey refuges, and the structure of coral reef fish assemblages. Ecological Monographs 63, 77–101. Hobbs, J.-P.A., Jones, G.P. and Munday, P.L. (2010) Rarity and extinction risk in coral reef angelfishes on isolated islands: interrelationships among abundance, geographic range size and specialisation. Coral Reefs 29, 1–11. Hobbs, J.-P.A., Jones, G.P., Munday, P.L., Connolly, S.R. and Srinivasan, M. (2012) Biogeography and the structure of coral reef fish communities on isolated islands. Journal of Biogeography 39, 130–139. © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 Tropical vagrant fishes D A Feary et al., Hoegh-Guldberg, O. and Bruno, J.F. (2010) The impact of climate change on the world’s marine ecosystems. Science 328, 1523–1528. Hoegh-Guldberg, O., Mumby, P.J., Hooten, A.J. et al. (2007) Coral reefs under rapid climate change and ocean acidification. Science 318, 1737–1742. Hoey, A.S. and McCormick, M.I. (2004) Selective predation for low body condition at the larval-juvenile transition of a coral reef fish. Oecologia 139, 23–29. Hogan, J.D., Fisher, R. and Nolan, C. (2007) Critical swimming speed of settlement-stage coral reef fishes from the Caribbean: a methodological and geographical comparison. Bulletin of Marine Science 80, 219–231. Holbrook, S.J. and Schmitt, R.J. (2002) Competition for shelter space causes density-dependent predation mortality in damselfishes. Ecology 83, 2855–2868. Holbrook, S.J., Brooks, A.J. and Schmitt, R.J. (2002a) Predictability of fish assemblages on coral patch reefs. Marine and Freshwater Research 53, 181–188. Holbrook, S.J., Brooks, A.J. and Schmitt, R.J. (2002b) Variation in structural attributes of patch-forming corals and in patterns of abundance of associated fishes. Marine and Freshwater Research 53, 1045–1053. Holmes, T.H. and McCormick, M.I. (2009) Influence of prey body characteristics and performance on predator selection. Oecologia 159, 401–413. Holmes, T.H. and McCormick, M.I. (2010) Size-selectivity of predatory reef fish on juvenile prey. Marine Ecology Progress Series 399, 273–283. Hurst, T.P. (2007) Causes and consequences of winter mortality in fishes. Journal of Fish Biology 71, 315–345. Hurst, T.P. and Conover, D.O. (2001) Activity-related constraints on overwintering young-of-the-year striped bass (Morone saxatilis). Canadian Journal of ZoologyRevue Canadienne De Zoologie 79, 129–136. Hutchings, J.A., Myers, R.A., Garcia, V.B., Lucifora, L.O. and Kuparinen, A. (2012) Life-history correlates of extinction risk and recovery potential. Ecological Applications 22, 1061–1067. Hutchins, J.B. (1991) Dispersal of tropical fishes to temperate seas in the southern hemisphere. Journal of the Royal Society of Western Australia 74, 79–84. Hutchins, J.B. and Pearce, A.F. (1994) Influence of the current on recruitment of tropical reef fishes at rottnest island, Western Australia. Bulletin of Marine Science 54, 245–255. IPCC (2007) Climate change 2007: the physical science basis. In: Contribution of Working Group I to the Fourth Assessment. Report of the Intergovernmental Panel on Climate Change. (eds S. Solomon, D. Qin, M. Manning, Z. Chen, M. Marquis, K. Averyt, M.M.B. Tignor, H.L.M. Jr and Z. Chen). Cambridge University Press, Cambridge, UK, pp. 996. Jones, G.P., Milicich, M.J., Emslie, M.J. and Lunow, C. (1999) Self-recruitment in a coral reef fish population. Nature 402, 802–804. Jones, G.P., Caley, M.J. and Munday, P.L. (2002) Rarity in coral reef fish communities. In: Coral Reef Fishes. Dynamics and Diversity in a Complex Ecosystem (ed. P.F. Sale), Academic Press, San Diego, pp. 81–102. Jones, G.P., Planes, S. and Thorrold, S.R. (2005) Coral reef fish larvae settle close to home. Current Biology 15, 1314–1318. Jones, G.P., Almany, G.R., Russ, G.R. et al. (2009a) Larval retention and connectivity among populations of corals and reef fishes: history, advances and challenges. Coral Reefs 28, 307–325. Jones, G.P., Russ, G.R., Sale, P.F. and Steneck, R.S. (2009b) Theme section on “Larval connectivity, resilience and the future of coral reefs”. Coral Reefs 28, 303–305. Keane, J.P. and Neira, F.J. (2008) Larval fish assemblages along the south-eastern Australian shelf: linking mesoscale non-depth-discriminate structure and water masses. Fisheries Oceanography 17, 263–280. Kerrigan, B.A. (1996) Temporal patterns in size and condition at settlement in two tropical reef fishes (Pomacentridae: Pomacentrus amboinensis and P. nagasakiensis). Marine Ecology Progress Series 135, 27–41. Kingsford, M. and Battershill, C. (1998) Studying Temperate Marine Environments. A Handbook for Ecologists., Canterbury University Press, Christchurch. Kingsford, M.J. and Carlson, I.J. (2010) Patterns of distribution and movement of fishes, Ophthalmolepis lineolatus and Hypoplectrodes maccullochi, on temperate rocky reefs of south eastern Australia. Environmental Biology of Fishes 88, 105–118. Kitchell, J.F., Stewart, D.J. and Weininger, D. (1977) Applications of a bioenergetics model to yellow perch (Perca flavescens) and walleye (Stizostedion vitreum vitreum). Journal of the Fisheries Research Board of Canada 34, 1922–1935. Kokita, T. (2004) Latitudinal compensation in female reproductive rate of a geographically widespread reef fish. Environmental Biology of Fishes 71, 213–224. Last, P.R., White, W.T., Gledhill, D.C. et al. (2011) Longterm shifts in abundance and distribution of a temperate fish fauna: a response to climate change and fishing practices. Global Ecology and Biogeography 20, 58–72. Lawton, J.H. (1999) Are there general laws in ecology? Oikos 84, 177–192. Lee, T.N., Rooth, C., Williams, E., McGowan, M., Szmant, A.F. and Clarke, M.E. (1992) Influence of Florida Current, gyres and wind-driven circulation on transport of larvae and recruitment in the Florida Keys coral reefs. Continental Shelf Research 12, 971–1002. Lee, T.N., Clarke, M.E., Williams, E., Szmant, A.F. and Berger, T. (1994) Evolution of the Tortugas Gyre and its influence on recruitment in the Florida Keys. Bulletin of Marine Science 54, 621–646. Lee, T.N., Leaman, K., Williams, E., Berger, T. and Atkinson, L. (1995) Florida current meanders and gyre © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 611 Tropical vagrant fishes D A Feary et al, formation in the southern straits of Florida. Journal of Geophysical Research-Oceans 100, 8607–8620. Leis, J.M. (1993) Larval fish assemblages near indo-pacific coral-reefs. Bulletin of Marine Science 53, 362–392. Leis, J.M., Sweatman, H.P.A., Reader, S.E. (1996) What the pelagic stages of coral reef fishes are doing out in blue water: Daytime field observations of larval behavioural capabilities. Marine and Freshwater Research 47, 401–411. Leis, J.M. and Carson-Ewart, B.M. (1997) In situ swimming speeds of the late pelagic larvae of some IndoPacific coral-reef fishes. Marine Ecology Progress Series 159, 165–174. Leis, J.M. and Carson-Ewart, B.M. (1998) Complex behaviour by coral-reef fish larvae in open-water and near-reef pelagic environments. Environmental Biology of Fishes 53, 259–266. Leis, J.M. and Carson-Ewart, B.M. (2003) Orientation of pelagic larvae of coral-reef fishes in the ocean. Marine Ecology Progress Series 252, 239–253. Leis, J.M. and Goldman, B. (1984) A preliminary distributional study of fish larvae near a ribbon coral-reef in the great barrier-reef. Coral Reefs 2, 197–203. Leis, J.M. and McCormick, M.I. (2002) The biology, behavior, and ecology of the pelagic, larval stage of coral reef fishes. In: Coral Reef Fishes: Dynamic and Diversity in a Complex Ecosystem (eds P.F. Sale). Elsevier, San Diego, CA, USA, pp. 171–199. Leis, J.M., Carson-Ewart, B.M. and Cato, D.H. (2002) Sound detection in situ by the larvae of a coral-reef damselfish (Pomacentridae). Marine Ecology Progress Series 232, 259–268. Leis, J.M., Carson-Ewart, B.M., Hay, A.C. and Cato, D.H. (2003) Coral-reef sounds enable nocturnal navigation by some reef-fish larvae in some places and at some times. Journal of Fish Biology 63, 724–737. Leis, J.M., Hay, A.C., Lockett, M.M., Chen, J.-P. and Fang, L.-S. (2007) Ontogeny of swimming speed in larvae of pelagic-spawning, tropical, marine fishes. Marine Ecology Progress Series 349, 255–267. Leis, J.M., Siebeck, U. and Dixson, D.L. (2011) How Nemo finds home: the neuroecology of dispersal and of population connectivity in larvae of marine fishes. Integrative and Comparative Biology 51, 826–843. Leis, J.M., Sweatman, H.P.A. and Reader, S.E. (1996) What the pelagic stages of coral reef fishes are doing out in blue water: Daytime field observations of larval behavioural capabilities. Marine and Freshwater Research 47, 401–411. Lessios, H.A. and Robertson, D.R. (2006) Crossing the impassable: genetic connections in 20 reef fishes across the eastern Pacific barrier. Proceedings of the Royal Society B-Biological Sciences 273, 2201–2208. Lien, Y.T., Fukami, H. and Yamashita, Y. (2012) Symbiodinium Clade C dominates zooxanthellate corals 612 (Scleractinia) in the temperate region of Japan. Zoological Science 29, 173–180. Limouzy-Paris, C.B., Graber, H.C., Jones, D.L., Ropke, A.W. and Richards, W.J. (1997) Translocation of larval coral reef fishes via sub-mesoscale spin-off eddies from the Florida current. Bulletin of Marine Science 60, 966–983. Loennstedt, O.M., McCormick, M.I., Meekan, M.G., Ferrari, M.C.O. and Chivers, D.P. (2012) Learn and live: predator experience and feeding history determines prey behaviour and survival. Proceedings of the Royal Society B-Biological Sciences 279, 2091–2098. Lonnstedt, O.M., McCormick, M.I. and Chivers, D.P. (2012) Well-informed foraging: damage-released chemical cues of injured prey signal quality and size to predators. Oecologia 168, 651–658. Lo-Yat, A., Meekan, M.G., Carleton, J.H. and Galzin, R. (2006) Large-scale dispersal of the larvae of nearshore and pelagic fishes in the tropical oceanic waters of French Polynesia. Marine Ecology Progress Series 325, 195–203. Lu, J., Deser, C. and Reichler, T. (2009) Cause of the widening of the tropical belt since 1958. Geophysical Research Letters 36, L03803. Madin, E.M.P., Ban, N.C., Doubleday, Z.A., Holmes, T.H., Pecl, G.T. and Smith, F. (2012) Socio-economic and management implications of range-shifting species in marine systems. Global Environmental Change-Human and Policy Dimensions 22, 137–146. Malcolm, H.A., Davies, P.L., Jordan, A. and Smith, S.D.A. (2011) Variation in sea temperature and the East Australian Current in the Solitary Islands region between 2001–2008. Deep-Sea Research Part Ii-Topical Studies in Oceanography 58, 616–627. Malcolm, H.A., Gladstone, W., Lindfield, S., Wraith, J. and Lynch, T.P. (2007) Spatial and temporal variation in reef fish assemblages of marine parks in New South Wales, Australia - baited video observations. Marine Ecology Progress Series 350, 277–290. Malcolm, H.A., Smith, S.D.A., and Jordan, A. (2010) Using patterns of reef fish assemblages to refine a Habitat Classification System for marine parks in NSW, Australia. Aquatic Conservation-Marine and Freshwater Ecosystems 20, 83–92. Manassa, R.P. and McCormick, M.I. (2012) Social learning and acquired recognition of a predator by a marine fish. Animal Cognition 15, 559–565. McBride, R. (1996) On the rarity of banded butterflyfish in the Mid-Atlantic. Underwater Naturalist 23, 18–20. McBride, R.S. and Able, K.W. (1998) Ecology and fate of butterflyfishes, Chaetodon spp., in the temperate, western North Atlantic. Bulletin of Marine Science 63, 401–416. McCormick, M.I. (1998) Condition and growth of reef fish at settlement: is it important? Australian Journal of Ecology 23, 258–264. © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 Tropical vagrant fishes D A Feary et al., McCormick, M.I. and Hoey, A.S. (2004) Larval growth history determines juvenile growth and survival in a tropical marine fish. Oikos 106, 225–242. McCormick, M.I. and Holmes, T.H. (2006) Prey experience of predation influences mortality rates at settlement in a coral reef fish, Pomacentrus amboinensis. Journal of Fish Biology 68, 969–974. Mora, C. and Ospına, A.F. (2001) Tolerance to high temperatures and potential impact of sea warming on reef fishes of Gorgona Island (tropical Eastern Pacific). Marine Biology 139, 765–769. Munday, P.L. and Jones, G.P. (1998) The ecological implications of small body size among coral reef fishes. Oceanography and Marine Biology Annual Review 36, 373–411. Munday, P.L., Jones, G.P. and Caley, M.J. (1997) Habitat specialisation and the distribution and abundance of coral-dwelling gobies. Marine Ecology Progress Series 152, 227–239. Munday, P.L., Jones, G.P. and Caley, M.J. (2001) Interspecific competition and coexistence in a guild of coraldwelling fishes. Ecology 82, 2177–2189. Munday, P.L., Jones, G.P., Pratchett, M.S. and Williams, A.J. (2008a) Climate change and the future for coral reef fishes. Fish and Fisheries 9, 261–285. Munday, P.L., Kingsford, M.J., O’Callaghan, M. and Donelson, J.M. (2008b) Elevated temperature restricts growth potential of the coral reef fish Acanthochromis polyacanthus. Coral Reefs 27, 927–931. Munday, P.L., Leis, J.M., Lough, J.M. et al. (2009) Climate change and coral reef connectivity. Coral Reefs 28, 379–395. Nakazono, A. (2002) Fate of tropical reef fish juveniles that settle to a temperate habitat. Fisheries Science 68, 127–130. Nicholson, J.K., Holmes, E., Kinross, J. et al. (2012) Hostgut microbiota metabolic interactions. Science 336, 1262–1267. Nye, J.A., Link, J.S., Hare, J.A. and Overholtz, W.J. (2009) Changing spatial distribution of fish stocks in relation to climate and population size on the Northeast United States continental shelf. Marine Ecology Progress Series 393, 111–129. O’Connor, M.I., Bruno, J.F., Gaines, S.D., Halpern, B.S. and Lester, S.E. (2007) Temperature control of larval dispersal and the implications for marine ecology, evolution, and conservation. Proceedings of the National Academy of Sciences USA 104, 1266–1271. Oviatt, C.A. (2004) The changing ecology of temperate coastal waters during a warming trend. Estuaries 27, 895–904. Paris, C.B. and Cowen, R.K. (2004) Direct evidence of a biophysical retention mechanism for coral reef fish larvae. Limnology and Oceanography 49, 1964–1979. Parmesan, C. (2006) Ecological and evolutionary responses to recent climate change. Annual Review of Ecology Evolution and Systematics 37, 637–669. Parmesan, C. and Yohe, G. (2003) A globally coherent fingerprint of climate change impacts across natural systems. Nature 421, 37–42. Parmesan, C., Ryrholm, N., Stefanescu, C. et al. (1999) Poleward shifts in geographical ranges of butterfly species associated with regional warming. Nature 399, 579–583. Parmesan, C., Gaines, S., Gonzalez, L. et al. (2005) Empirical perspectives on species borders: from traditional biogeography to global change. Oikos 108, 58– 75. Pauly, D. (1997) Geometrical constraints on body size. Trends in Ecology and Evolution 12, 442–443. Planes, S. and Lecaillon, G. (2001) Caging experiment to examine mortality, during metamorphosis of coral reef fish larvae. Coral Reefs 20, 211–218. Planes, S., Jones, G.P. and Thorrold, S.R. (2009) Larval dispersal connects fish populations in a network of marine protected areas. Proceedings of the National Academy of Sciences of the United States of America 106, 5693–5697. Poertner, H.O. and Farrell, A.P. (2008) Physiology and climate change. Science 322, 690–692. Pratchett, M.S. (2005) Dietary overlap among coral-feeding butterflyfishes (Chaetodontidae) at Lizard Island, northern Great Barrier Reef. Marine Biology 148, 373– 382. Pratchett, M.S., Hoey, A.J., Feary, D.A., Bauman, A., Burt, J. and Riegl, B. (2012) Functional composition of Chaetodon butterflyfishes at a peripheral and extreme coral reef location, the southern Persian Gulf. Marine Pollution Bulletin. DOI:10.1016/j.marpolbul.2012.10. 014 [Epub ahead of print]. Pratchett, M.S., Wilson, S.K., Berumen, M.L. and McCormick, M.I. (2004) Sublethal effects of coral bleaching on an obligate coral feeding butterflyfish. Coral Reefs 23, 352–356. Pratchett, M.S., Berumen, M.L., Marnane, M.J., Eagle, J.V. and Pratchett, D.J. (2008a) Habitat associations of juvenile versus adult butterflyfishes. Coral Reefs 27, 541–551. Pratchett, M.S., Munday, P.L., Wilson, S.K. et al. (2008b) Effects of climate-induced coral bleaching on coral-reef fishes: ecological and economic consequences. Oceanography and Marine Biology Annual Review 46, 251–296. Pratt, T.C. and Fox, M.G. (2002) Influence of predation risk on the overwinter mortality and energetic relationships of young-of-year walleyes. Transactions of the American Fisheries Society 131, 885–898. Precht, W.F. and Aronson, R.B. (2004) Climate flickers and range shifts of reef corals. Frontiers in Ecology and the Environment 2, 307–314. Ridgway, K.R. and Dunn, J.R. (2003) Mesoscale structure of the mean East Australian Current System and its relationship with topography. Progress in Oceanography 56, 189–222. © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 613 Tropical vagrant fishes D A Feary et al, Riegl, B.M., Purkis, S.J., Al-Cibahy, A.S., Abdel-Moati, M.A. and Hoegh-Guldberg, O. (2011) Present limits to heat-adaptability in corals and population-level responses to climate extremes. PLoS ONE 6, e24802. Rodolfo-Metalpa, R., Reynaud, S., Allemand, D. and Ferrier-Pages, C. (2008) Temporal and depth responses of two temperate corals, Cladocora caespitosa and Oculina patagonica, from the North Mediterranean Sea. Marine Ecology Progress Series 369, 103–114. Roughan, M., Macdonald, H.S., Baird, M.E. and Glasby, T.M. (2011) Modelling coastal connectivity in a Western Boundary Current: seasonal and inter-annual variability. Deep-Sea Research Part Ii-Topical Studies in Oceanography 58, 628–644. Roughgarden, J. (2009) Is there a general theory of community ecology? Biology and Philosophy 24, 521– 529. Russell, B.C. (1971) A preliminary annotated checklist of fishes of the Poor Knights Islands. Tane 17, 81–90. Sale, P.F. (ed.) (1991) The Ecology of Fishes on Coral Reefs. Academic Press, New York, 754 pp. Sale, P.F. (ed.) (2002) Coral Reef Fishes. Dynamics and Diversity in a Complex Ecosystem. Academic Press, San Diego, 549 pp. Scott, A. and Harrison, P.L. (2008) Larval settlement and juvenile development of sea anemones that provide habitat for anemonefish. Marine Biology 154, 833–839. Seidel, D.J., Fu, Q., Randel, W.J. and Reichler, T.J. (2008) Widening of the tropical belt in a changing climate. Nature Geoscience 1, 21–24. Shanks, A.L. (2009) Pelagic larval duration and dispersal distance revisited. Biological Bulletin 216, 373–385. Soberon, J. (2007) Grinnellian and Eltonian niches and geographic distributions of species. Ecology Letters 10, 1115–1123. Soeparno, Nakamura, Y., Shibuno, T. and Yamaoka, K. (2012) Relationship between pelagic larval duration and abundance of tropical fishes on temperate coasts of Japan. Journal of Fish Biology 80, 346–357. Sorte, C.J.B., Williams, S.L. and Carlton, J.T. (2010) Marine range shifts and species introductions: comparative spread rates and community impacts. Global Ecology and Biogeography 19, 303–316. Sponaugle, S. and Grorud-Covert, K. (2006) Environmental variability, early life-history traits, and survival of new coral reef fish recruits. Integrative and Comparative Biology 46, 623–633. Sponaugle, S., Cowen, R.K., Shanks, A. et al. (2002) Predicting self-recruitment in marine populations: biophysical correlates and mechanisms. Bulletin of Marine Science 70, 341–375. Sponaugle, S., Boulay, J.N. and Rankin, T.L. (2011) Growth- and size-selective mortality in pelagic larvae of a common reef fish. Aquatic Biology 13, 263–273. 614 Stefansdottir, L., Solmundsson, J., Marteinsdottir, G., Kristinsson, K. and Jonasson, J.P. (2010) Groundfish species diversity and assemblage structure in Icelandic waters during recent years of warming. Fisheries Oceanography 19, 42–62. Stevens, G.C. (1989) The latitudinal gradients in geographical range: how so many species co-exist in the tropics. American Naturalist 133, 240–256. Stillman, J. and Somero, G.N. (2000) A comparative analysis of the upper thermal tolerance limits of eastern Pacific porcelain crabs, genus Petrolisthes: Influences of latitude, vertical zonation, acclimation, and phylogeny. Physiological and Biochemical Zoology 73, 200–208. Stobutzki, I.C. and Bellwood, D.R. (1994) An analysis of the critical swimming abilities of pre- and post-settlement coral reef fishes. Journal of Experimental Marine Biology and Ecology 175, 275–286. Stobutzki, I.C. and Bellwood, D.R. (1997) Sustained swimming abilities of the late pelagic stages of coral reef fishes. Marine Ecology Progress Series 149, 35–41. Sunday, J.M., Bates, A.E. and Dulvy, N.K. (2012) Thermal tolerance and the global redistribution of animals. Nature Climate Change 2, 686–690. doi:10.1038/NCLIMATE1539. Suthers, I.M. (1998) Bigger? Fatter? Or is faster growth better? Considerations on condition in larval and juvenile coral-reef fish. Australian Journal of Ecology 23, 265–273. Swearer, S.E., Caselle, J.E., Lea, D.W. and Warner, R.R. (1999) Larval retention and recruitment in an island population of a coral-reef fish. Nature 402, 799–802. Swearer, S.E., Shima, J.S., Hellberg, M.E. et al. (2002) Evidence of self-recruitment in demersal marine populations. Bulletin of Marine Science 70, 251–271. Sweatman, H. (1988) Field evidence that settling coralreef fish larvae detect resident fishes using dissolved chemical cues. Journal of Experimental Marine Biology and Ecology 124, 163–174. Syahailatua, A., Roughan, M. and Suthers, I.M. (2011) Characteristic ichthyoplankton taxa in the separation zone of the East Australian Current: larval assemblages as tracers of coastal mixing. Deep-Sea Research Part Ii-Topical Studies in Oceanography 58, 678–690. Thomas, C.D. and Lennon, J.J. (1999) Birds extend their ranges northwards. Nature 399, 213. Thorrold, S.R., Latkoczy, C., Swart, P.K. and Jones, C.M. (2001) Natal homing in a marine fish metapopulation. Science 291, 297–299. Thresher, R. (1984) Reproduction in Reef Fishes. TFH Publications, Neptune City. Thresher, R.E. and Brothers, E.B. (1989) Evidence of intra- and inter-oceanic regional differences in the early life history of reef-associated fishes. Marine Ecology Progress Series 57, 187–205. © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 Tropical vagrant fishes D A Feary et al., Underwood, A.J., Kingsford, M.J. and Andrew, N.L. (1991) Patterns in shallow subtidal marine assemblages along the coast of New South Wales. Australian Journal of Ecology 6, 231–249. Victor, B.C. (1986) Larval settlement and juvenile mortality in a recruitment-limited coral-reef fish population. Ecological Monographs 56, 145–160. Weatherly, A.H. (1972) Growth and Ecology of Fish Population. Academic Press, London. Wellington, G.M. and Robertson, D.R. (2001) Variation in larval life-history traits among reef fishes across the Isthmus of Panama. Marine Biology 138, 11–22. Wellington, G.M. and Victor, B.C. (1989) Planktonic larval duration of one hundred species of Pacific and Atlantic damselfishes (Pomacentridae). Marine Biology 101, 557–567. Wilson, S.K., Graham, N.A.J., Pratchett, M.S., Jones, G.P. and Polunin, N.V.C. (2006) Multiple disturbances and the global degradation of coral reefs: are reef fishes at risk or resilient? Global Change Biology 12, 2220–2234. Wing, S.R., Botsford, L.W., Ralston, S.V. and Largier, J.L. (1998) Meroplanktonic distribution and circulation in a coastal retention zone of the northern California upwelling system. Limnology and Oceanography 43, 1710–1721. Wu, L.X., Cai, W.J., Zhang, L.P. et al. (2012) Enhanced warming over the global subtropical western boundary currents. Nature Climate Change 2, 161–166. Yamano, H., Sugihara, K. and Nomura, K. (2011) Rapid poleward range expansion of tropical reef corals in response to rising sea surface temperatures. Geophysical Research Letters 38, L04601. Supporting Information Additional Supporting Information may be found in the online version of this article: Data S1. Full list of all tropical reef associated fish species (Family and species) that have been identified as utilising temperate reef habitats within larval, sub-adult or adult phase. Data S2. Methods for logistic analyses. © 2013 John Wiley & Sons Ltd, F I S H and F I S H E R I E S , 15, 593–615 615