Size does matter : the eco-evolutionary effects of changing body size in fish

This is a self-archived version of an original article. This version may differ from the original in pagination and typographic details. Author(s): Title: Year: Version: Copyright: Rights: Rights url: Please cite the original version: In Copyright http://rightsstatements.org/page/InC/1.0/?language=en Size does matter : the eco-evolutionary effects of changing body size in fish © Canadian Science Publishing, 2020 Accepted version (Final draft) Ahti, Pauliina Anna; Kuparinen, Anna; Uusi-Heikkilä, Silva Ahti, P. A., Kuparinen, A., & Uusi-Heikkilä, S. (2020). Size does matter : the eco-evolutionary effects of changing body size in fish. Environmental Reviews, 28(3), 311-324. https://doi.org/10.1139/er-2019-0076 2020 Draft Size does matter – the eco-evolutionary effects of changing body size in fish Journal: Environmental Reviews Manuscript ID er-2019-0076.R2 Manuscript Type: Review Date Submitted by the Author: 20-Apr-2020 Complete List of Authors: Ahti, Pauliina; University of Jyväskylä , Department of Biological and Environmental Science Kuparinen, Anna; University of Jyväskylä , Department of Biological and Environmental Science Uusi-Heikkilä, Silva; University of Jyväskylä , Department of Biological and Environmental Science Is this manuscript invited for consideration in a Special Issue? : Not applicable (regular submission) Keyword: body size, fisheries, environmental change, evolution, life-history trait https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 1 1Size does matter – the eco-evolutionary effects of changing body size in fish 2Pauliina A. Ahti1*, Anna Kuparinen1 & Silva Uusi-Heikkilä1 31 Department of Biological and Environmental Sciences, P.O. Box 35, 40014 University of 4Jyväskylä, Finland 5*Corresponding author: Pauliina A. Ahti, Department of Biological and Environmental 6Sciences, P.O. Box 35, 40014 University of Jyväskylä, Finland, telephone: +358-41-49274459, e-mail: [email protected] 8Word count: 10534 9 10 11 12 13 14 15 16 17 18 19 20 21 Page 1 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 2 22 Abstract 23 Body size acts as a proxy for many fitness-related traits. Body size is also subject to 24 directional selection from various anthropogenic stressors such as increasing water 25 temperature, decreasing dissolved oxygen, fisheries, as well as natural predators. Changes in 26 individual body size correlate with changes in fecundity, behaviour, and survival, and can 27 propagate through populations and ecosystems by truncating age and size structures and 28 changing predator-prey dynamics. In this review, we will explore the causes and 29 consequences of changing body size in fish in the light of recent literature and relevant 30 theories. We will investigate the central role of body size in ecology by first discussing the 31 main selective agents that influence body size: fishing, increasing water temperature, 32 decreasing dissolved oxygen, and predation. We will then explore the impacts of these 33 changes at the individual, population and ecosystem levels. Considering the relatively high 34 heritability of body size, we will discuss how a change in body size can leave a genetic 35 signature in the population and translate to a change in the evolutionary potential of the 36 species. 37 38 Keywords: body size, fisheries, environmental change, evolution, life-history trait, plasticity 39 40 41 42 43 44 Page 2 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 3 45 Introduction 46 Historically, before the influence of modern human and industrialisation, most species 47 responded to the changing climate by shifting ranges (Cabanne et al. 2016). In the present 48 day, as anthropogenically driven, rapidly changing climate is accompanied by a range of 49 other selective agents, biological responses are likely to be more complex (Parmesan et al. 50 2000). Threats such as increasing temperature (Barnett et al. 2005), acidification (Orr et al. 51 2005), reduced dissolved oxygen (Garcia et al. 2005), pollution (Moore 2008), overfishing 52 (Jackson et al. 2001), and habitat destruction (Nordhaus et al. 2018) are posing serious 53 challenges to our oceans, often acting in concert. To survive the change, species have three 54 options: disperse to a more suitable habitat, adjust through phenotypic plasticity, or adapt 55 through evolutionary change. How well an organism can respond to a change depends largely 56 on its plasticity and evolutionary potential (Hoffmann and Willi 2008). 57 Conventionally, research has focused on either ecological responses such dispersal or 58 phenotypic plasticity, or evolutionary responses such as genetic adaptation, and this 59 distinction was based on the assumption that ecological and evolutionary responses occur at 60 vastly different time scales. However, it is now widely documented that evolutionary change 61 can occur over ecologically relevant timescales (Stockwell et al. 2003; Carroll et al. 2007), 62 and that these processes are not independent of each other (Hanski 2012). Consequently, to 63 fully understand the ecological and evolutionary processes in nature, we need to move 64 beyond just documenting the ecological or evolutionary change, towards exploring the impact 65 of the change in a wider concept. 66 Many natural and anthropogenic stressors are inducing selection pressure on a key ecological 67 trait: the body size of an organism. Body size, like many other traits, has both plastic 68 (Crozier and Hutchings 2014) and genetic (Mousseau and Roff 1987; Garcia De Leaniz et al. Page 3 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 4 69 2007) components. It acts as an easy-to-measure proxy for many life-history characters such 70 as lifetime reproductive success (Barneche et al. 2018), size and age at maturity (Jensen 71 1996), and survival (White et al. 2013). Many functions in an organism’s life, including 72 energetics, resource use, susceptibility to predators, species interactions and several 73 population parameters are functions of its body size (Werner and Gilliam 1984). 74 The question whether organisms have the evolutionary potential to respond to anthropogenic 75 stressors fast enough is becoming increasingly more important. Unlike terrestrial organisms, 76 fishes, being immersed in water, are in an intimate contact with their physical and chemical 77 environment making them particularly sensitive to any changes in their environment (Cossins 78 and Crawford 2005). Compared to air, oxygen availability in the water is much lower and due 79 to the higher viscosity of water, more effort is needed to utilise it (Verberk et al. 2011). 80 Increasing water temperature and decreasing oxygen content, even without the pressure from 81 fisheries, have been predicted to lead to sharp declines in fish body sizes (van Rijn et al. 82 2017). Regardless of the cause, changes in body size occur at the level of an individual, and 83 scale up to population, community and ecosystem levels (Pörtner 2002; Pörtner and Peck 84 2010). The nature of the change, whether it is plastic or genetic, determines whether the 85 change is likely to be reversible or lead to adaptation. The more the trait under selection 86 changes, and the more links there are between the trait variation and ecological interactions, 87 the more important the role of contemporary evolution to ecological processes becomes 88 (Hairston et al. 2005). 89 Here, we review the recent literature on the causes and consequences of changing body size 90 in fish and its links to ecological and evolutionary processes. While there is a large selection 91 of important and relevant literature focusing on body size in fish (Uusi-Heikkilä et al. 2008; 92 Devine et al. 2012; Enberg et al. 2012; Audzijonyte et al. 2013; Heino et al. 2013, 2015; Diaz 93 Pauli and Heino 2014; Diaz Pauli and Sih 2017; Hollins et al. 2018), much of it tends to focus Page 4 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 5 94 on fishing as a selective agent. We believe it is necessary to expand our investigations into 95 how different selective agents (i.e. not only fishing) affect body size, a key measure in 96 ecology and a central theme in several ecological theories. To achieve this, we draw attention 97 to not only fishing, but also to what we consider to be the main (or at least relatively well98 studied) abiotic factors i.e. increasing temperature and dissolved oxygen, as well as a major 99 biotic factor i.e. predation, in driving selection on body size. Other agents exist as well, such 100 as pH, water chemistry, primary production, light, and mate choice, and more studies are 101 needed to understand their role in selection. Additionally, there is currently a knowledge gap 102 on how different selection agents or stressors interact, and we call for more research effort on 103 the synergistic effects of multiple stressors. It should also be noted, that a single selective 104 agent may select for various traits such as body size, growth rate, and natural mortality 105 schedules. The majority of the current literature is descriptive, and fails to consider the far106 reaching effects of changing body size. Given the rapidly changing environment, and the 107 various selection pressures imposed by it, it is crucial to investigate the role of changing body 108 size in a wider perspective beyond fisheries science, to understand how those changes may 109 affect the entire ecosystem over ecological and evolutionary time. 110 With body size, we refer to metrics such as asymptotic length, size at age, body mass, as well 111 as growth rate. Although size at age is a state, and growth is a process, these metrics are 112 closely linked, as growth is necessary in order to reach a certain size (Enberg et al. 2012). 113 Also, size at age can be considered a proxy for growth (Heino et al. 2015). We will first 114 provide an overview of the main ecological theories behind the mechanistic functioning of 115 different selective agents, namely increasing temperature coupled with decreasing dissolved 116 oxygen, fishing, and predation. We will then use examples from relevant literature to discuss 117 how changes in body size can influence other life-history traits of an individual fish, and Page 5 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 6 118 propagate through different levels of biological organisation through to population and 119 ecosystem levels, at ecologically and evolutionarily relevant time scales (Fig.1). 120 121 The ecological theories behind different stressors as selection agents 122 In this section, we summarise the selection agents we focus on, namely: temperature, 123 dissolved oxygen, harvesting and natural predators, and provide an overview of the main 124 ecological theories behind the selective mechanisms (Fig. 2). 125 Temperature and dissolved oxygen 126 Temperature is known to affect organisms at community, population and individual levels. 127 This effect is embodied by three rules: the Bergmann’s rule, (Bergmann 1847), the James’ 128 rule (James 1970), and the Temperature – Size –rule (Atkinson 1994). The Bergmann’s rule 129 predicts that, at the community level, organisms in cold habitats are larger than those in 130 warmer habitats. While not universal (Belk and Houston 2002; Ashton and Feldman 2003; 131 Fisher et al. 2010), many taxa including birds and mammals (Meiri et al. 2003) as well as 132 marine (Fernández-torres et al. 2018; Saunders and Tarling 2018) and freshwater (Knouft 133 2004; Daufresne et al. 2009; Rypel 2014) fishes are known to conform to Bergmann’s rule. 134 This rule is known to apply particularly to freshwater species in cold waters (Rypel 2014). 135 However, at least for freshwater fish, this pattern may be disrupted by non-native species 136 (Blanchet et al. 2010). At the population level, James’ rule (James 1970) predicts that within 137 species, populations with smaller body size tend to be found in warmer habitats. As with 138 Bergmann’s rule, James’ rule is not universal, but does hold for some fishes in the marine 139 (Pörtner et al. 2008; Cappo et al. 2013) as well as freshwater (Daufresne et al. 2009) 140 environments. Temperature is a particularly important aspect in the life of ectothermic 141 organisms, such as most fishes, as they rely on external source for body heat. The wellPage 6 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 7 142 established temperature-size –rule (Atkinson 1994) predicts that at lower temperatures 143 organisms grow slower but reach larger size at maturity (or inversely at higher temperatures 144 organisms have higher growth rate but lower final size). Indeed, it has been proposed that 145 shrinking body size is one of the universal responses to global warming, alongside with range 146 shifts and life cycle mismatches (Daufresne et al. 2009; Sheridan and Bickford 2011). 147 Declines in body size has been observed in marine (Daufresne et al. 2009; Cheung et al. 148 2012; Baudron et al. 2014) as well as freshwater (Baudron et al. 2011; Forster et al. 2012) 149 fishes, ranging from warm water to cold water species in every ocean basin as well as in 150 freshwater environments (Daufresne et al. 2009; Cheung et al. 2012). ). 151 Coupled with increasing water temperature is a decreased level of dissolved oxygen. Warm 152 water, whether marine or fresh, holds less oxygen than cold water, and water mass can 153 become under-saturated with oxygen due to excess organic carbon and other anthropogenic 154 processes (Rabalais et al. 2010). Oxygen limitation together with increasing temperature may 155 have synergistic effects on aquatic life, as limited oxygen can further impair the tolerance for 156 warmer temperatures (Pörtner and Knust 2007; Verberk et al. 2016). Warm water causes two 157 opposing effects: on the one hand warmer water holds less oxygen, on the other hand warmer 158 temperature increases the anabolic oxygen demand of fish, complicating the oxygen need vs. 159 oxygen supply -equation. Following this imbalance, it has been suggested that the surface 160 area of the gills cannot meet the oxygen demand of large bodies, a phenomenon explained by 161 the Gill-Oxygen Limitation (GOL) theory (Pauly 1981; Pauly and Cheung 2018). 162 Consequently, small body size may become favourable. However, while the trend of 163 decreasing body size with increasing temperature and decreasing oxygen is evident in both 164 marine (Pörtner and Knust 2007) and freshwater (Andrews et al. 1973; Pedersen 1987; Pauli 165 et al. 2017) environments, the underlying mechanisms of the GOL theory are currently 166 debated. It has been suggested that the scaling of gills with body mass, which the GOL theory Page 7 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 14 315 et al. (2001) showed that guppies in the high-predation environment grew faster, but also 316 reached higher asymptotic lengths than those inhabiting low-predation sites. Reznick et al. 317 (2001) suggested that the higher growth rate and higher asymptotic length were in part due to 318 high-predation areas also having higher resource availability. This could be caused by abiotic 319 factors such as the higher light level in the area resulting in higher primary productivity, or it 320 could be a consequence of predators lowering prey biomass thus freeing more nutrients per 321 capita (Reznick et al. 2001). Compared to selection induced by fisheries, natural selection 322 often operates in the opposite direction, favouring larger size (Olsen and Moland 2011). The 323 fitness landscape then becomes dynamic, moving towards the direction of the stronger 324 selection force (Edeline et al. 2007). 325 Over the last two decades, experimental studies have started to reveal the genetic basis of 326 changing body size. However, while experimental studies allow for the control of 327 confounding factors, they lack the density dependence present in nature and thus alone cannot 328 be directly translated to describe processes taking place in the nature. Most of this 329 experimental evidence comes from harvesting studies on both freshwater and marine/ 330 brackish species, and the results have indicated no differences in the mechanistic changes as a 331 result of harvest depending on the salinity of the water. Following positively size-selective 332 harvest (i.e., large individuals removed from the population, selection similar to most 333 fisheries) of the Atlantic silverside (Menidia menidia) over four generations, Conover and 334 Munch (2002) reported significantly lower body weight and slower growth rate of the 335 positively size-selected line. Later, using the same selection lines, Salinas et al. (2012) 336 showed that some of these changes were reversible, some partially reversible, and some did 337 not show any signs of reversibility after five generations of the cessation of fishing. While 338 this common garden approach suggested that some of the changes were likely to be genetic, 339 the direct evidence for genetic changes caused by size-selective harvesting was provided over Page 14 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 15 340 a decade later. Van Wijk et al. (2013) subjected guppies to three generations of positively and 341 negatively size-selective harvest and quantified the phenotypic and genetic responses to 342 selection. The phenotypic responses to directional selection were similar to those observed by 343 Conover and Munch (2002): positively size-selective harvest led to a reduction in adult body 344 size. Van Wijk et al. (2013) used 17 candidate loci to study genetic changes in response to 345 size-selective harvesting, and demonstrated changes in seven of them. At least three of their 346 candidate loci were associated with body size. Following a similar harvest design over five 347 generations, an experimental study on zebrafish (Danio rerio) provided further evidence on 348 declining body size and changes in associated life-history characters after intensive 349 directional selection (Uusi-Heikkilä et al. 2015). Uusi-Heikkilä et al. (2015) utilised 384 350 single nucleotide polymorphisms (SNPs) and identified 21 outlier SNPs responding to 351 directional selection, thus demonstrating contemporary evolutionary changes in the 352 experimentally exploited populations. With the same experimental selection lines, Uusi353 Heikkilä et al. (2017) studied harvest-induced changes in gene expression by sequencing the 354 transcriptomes of individual fishes. They demonstrated that size-selective harvesting had 355 changed the expression of over 4 000 genes and some of these changes were associated with 356 changes at the sequence level. Importantly, they showed that the changes in gene expression 357 were slow to reverse. Further, in 2019, Therkildsen and colleagues showed that the 358 phenotypic responses to harvest selection may in fact be underlain by divergent genomic 359 shifts, meaning fishing may potentially cause genomic changes comparable to distinct 360 populations in nature (Therkildsen et al. 2019). Experimental studies such as these can further 361 our understanding of the nature of the changes, and help us monitor and manage fish 362 populations in an efficient way. 363 Changes in growth rate Page 15 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 16 364 The fight for survival begins from the egg phase, and continues through the larval stage into 365 adulthood and eventual mortality. This survival is often non-random with respect to size. The 366 ‘bigger is better’ –hypothesis suggests that fast and large-growing larvae may better avoid 367 predators. The size-selective mortality of the sprat Spratelloides gracilis provided evidence 368 for this hypothesis, as selection was operating against small and slow growing individuals 369 (Meekan et al. 2006). Allain et al. (2003) reported similar trends in anchovies (Engraulis 370 encrasicolus) and attributed survival to faster larval growth rate. While juvenile stage is of 371 crucial importance as it determines the future cohort sizes, juvenile growth rates can be 372 altered by size selective harvest (Conover and Munch 2002; Walsh et al. 2006), changing 373 temperatures (Meekan et al. 2003; Baudron et al. 2011, 2014; Rogers et al. 2011), and 374 predation (Reznick et al. 2001; Heins et al. 2016). 375 Body size and growth rate are closely associated, and for animals with indeterminate growth 376 this close association is often described by the von Bertalanffy equation. There is a strong 377 negative correlation between the von Bertalanffy (vB) growth parametres = asymptotic 378 length, and k = the intrinsic individual growth rate, suggesting a presence of trade-off. 379 Following this, if asymptotic length decreases, the growth rate would be expected to increase 380 (and vice versa). However, depending on the selective agent, selection on body size may 381 influence growth rate differently. 382 Baudron et al. (2011) used over three decades of length and age data to fit the von Bertalanffy 383 growth model and investigated the growth response of North Sea haddock (Melanogrammus 384 aeglefinus) to different temperature scenarios. They showed that increasing temperature was 385 linked with a decrease in L∞. This finding was coupled with an increase in the growth rate 386 during early life stages (Baudron et al. 2011), in line with the vB growth model (where L∞ 387 and intrinsic individual growth rate correlate negatively), and as predicted by the 388 Temperature-Size –rule. A similar increase in juvenile growth rate with an increasing Page 16 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 17 389 temperature appears to be relatively common, as it has also been detected in the North Sea 390 cod (Rindorf et al. 2008), Northeast Arctic cod (Michalsen et al. 1998), and Norwegian 391 coastal cod (Otterlei et al. 1999),, and holds true across spatial scales (Brander 1995). In 392 some cases, temperature may select directly for increased juvenile growth rate. Thresher et al. 393 (2007) examined otolith data dating back to 1861, and reported a significant change in the 394 growth rate of six out of eight studied species. Three of these species were shallow water 395 fishes (< 250 m), and thus had been subjected to increasing water temperatures over the past 396 century, and showed increasing juvenile growth rates. The remaining three species were 397 deep-water species, and temperature recontructions inferred from deep-water corals fitted the 398 growth curves of the older, deep-water species which showed no incearese in juvenile growth 399 rates (Thresher et al. 2007). More recently, Morrongiello et al. (2019) were the first to report 400 fishing-induced shifts in thermal reaction norms in marine fish, highlighting the potential 401 synergistic effects of harvest and changing temperature. By fishing out the largest invididuals 402 that have a high thermal capacity, we might be weakening the species’ adaptive potential to 403 respond to climate change (Morrongiello et al. 2019). 404 While the fish in the study by Morrongiello et al. (2019) attained faster adult growth rates 405 (partially due to release from density dependence as a result of fishing), a typical selection 406 pressure from fisheries tends to select towards lower growth rate (Enberg et al. 2012). 407 Empirical studies of the Alpine freshwater whitefish Coregonus palaea (Nusslé et al. 2009), 408 Coregonus albellus, and Coregonus fatioi (Nusslé et al. 2011), all subject to steady and heavy 409 fishing pressure, have shown declines in the average adult (but not juvenile) growth rate as 410 well as significant selection differentials over several generations. Studies in the laboratory 411 have provided further evidence for changes in growth rate. Experimental harvesting studies 412 have revealed declines in juvenile growth rate following positively size-selective harvest over 413 four generations (Conover and Munch 2002). The parental harvest extended its influence to Page 17 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 18 414 the next generation, causing changes also in the larval growth rates. The larvae of positively 415 size-selected parents evolved to grow slower than the larvae of negatively size-selected 416 parents, potentially subjecting slow-growing larvae to higher predation pressure in nature 417 (Conover and Munch 2002). 418 It is thought that the natural mortality of many fish populations has been increasing in the 419 recent decades (Gislason et al. 2010). Following the presence of life history trade-offs in 420 nature (Jensen 1996; Douhard et al. 2015), it could be that increased growth rate early in life 421 is associated with increased mortality late in life. Indeed, accelerated growth rate is known to 422 be linked to shorter lifespan in many species, (Metcalfe and Monaghan 2003), and a recent 423 meta-analysis suggested a link between growth rate and natural mortality in fish (Gislason et 424 al. 2010). The link between faster growth rate and higher natural mortality (or shorter life425 span, which in a controlled laboratory environment corresponds to higher natural mortality) 426 has also been shown experimentally in three-spined sticklebacks (Lee et al. 2013). By 427 inducing changes in fish growth rate, we may be indirectly altering fish natural mortality 428 rates, too. 429 Big and bold? Linking body size and behaviour 430 Body size is known to correlate with many behavioural traits, which can enhance or reduce 431 individual fitness. For example, after five generations of harvesting, small Atlantic silverside 432 (Menidia menidia) showed reduced willingness to forage under a threat of predation 433 compared to the large ones, potentially reducing individual energy flow and therefore 434 reducing fitness (Walsh et al. 2006). Later, using the same experimental populations, Salinas 435 et al. (2012) showed that the changes in food consumption did not recover after the cessation 436 of fishing. This kind of change in food consumption is likely to influence the population 437 growth rate and productivity (Conover and Munch 2002). Uusi-Heikkilä et al. (2015) Page 18 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 19 438 investigated behavioural changes in juvenile zebrafish, and showed that negatively size439 selected fish (i.e. small individuals removed) were more active, explorative and bolder than 440 positively size-selected fish (i.e. large individuals removed), and that this shift in behaviour 441 was accompanied by genetic changes (but see Sbragaglia et al. 2019). Although not 442 demonstrated by the zebrafish study, it can be speculated that larger fish have higher energy 443 demand and therefore they are more active and bolder in search of food (Réale et al. 2010). 444 This in turn could make them more vulnerable not only to natural predators but also to fishing 445 gear. 446 Fishing gear selection is a two-way street. While behaviour can influence the vulnerability to 447 fishing, selective fishing can cause behavioural changes. As an example, Sutter et al. (2012) 448 focused on behaviour instead of size, and showed that aggression, intensity of parental care, 449 and reproductive fitness positively correlated with vulnerability to angling. Similarly, 450 physiological traits such as anaerobic swimming performance and metabolic demand are 451 known to be linked to vulnerability to fishing (Killen et al. 2015). Fishing can also be the 452 cause of behavioural changes, as the gear retention probability depends on the fishing gear in 453 question, and the selection curves are often non-linear with respect to size (Kuparinen et al. 454 2009). Since size and behavioural traits are linked, and fishing often selects a certain size 455 regime, the selection likely affects behavioural traits too. Obviously, vulnerability also 456 depends on the fishing gear at hand. Behavioural changes associated with body size have also 457 been studied in the context of fish farming, as it provides a semi-natural laboratory to study 458 contemporary evolution. Farmed fish that have been selected for higher growth rates are 459 expressing changes in boldness (Biro et al. 2004). The fish under selection tend to take more 460 risks while foraging, grow faster and survive at a lower rate than their wild conspecifics (Biro 461 et al. 2004; Biro and Post 2008). Page 19 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 20 462 In addition to fishing and fish farming, behavioural characters may be indirectly impacted by 463 temperature. Given that fishes are mainly ectothermic, increasing water temperature will 464 likely affect metabolic rates, which in turn may alter feeding behaviour and activity. Scott et 465 al. (2017) showed that this kind of response is size dependant. On the Great Barrier Reef in 466 Australia, the common coral trout (Plectropomus leopardus) shows more active foraging 467 behaviour during hotter months than colder months, but when the water temperature exceeds 468 30 ºC, foraging frequency declines. This may suggest the presence of a temperature 469 threshold, beyond which P. leopardus cannot compensate the increased basal metabolic rate 470 with increased foraging activity (Scott et al. 2017). They also showed that larger individuals 471 spent more time completely inactive in increasing temperatures compared to their smaller 472 conspecifics (Scott et al. 2017). This kind of behavioural change may have far-reaching 473 effects, from non-sufficient foraging and abnormal spawning-related movements to predator474 prey relationships (Scott et al. 2017). Studying the same species, Johansen et al. (2015) 475 demonstrated, that while P. leopardus, a predatory species, may be able to increase food 476 intake in response to increased metabolic demand due to increased water temperature, the 477 lower trophic levels may not be able to provide this energy. This highlights the importance of 478 considering the wider ecosystem, not just one species, when predicting the influence of 479 natural or anthropogenic stressors. 480 An interesting link between changing sea temperature, body size and activity level was also 481 discovered by van Rijn et al. (2017). They studied 74 fish species across space in the 482 Mediterranean Sea, ranging from small bottom-associated gobies to large pelagic tunas, and 483 demonstrated that body size decreased with increasing temperature as predicted by the 484 Temperature-Size –rule. Additionally, they found that the activity level of the species was 485 strongly correlated with the decrease in body size, so that large, active species showed the 486 strongest decline in size in response to increasing temperature (van Rijn et al. 2017). This Page 20 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 21 487 highlights the complexity of several selective agents acting simultaneously. As discussed 488 above, fisheries selection may select for a certain type of behaviour or activity level, and as 489 shown here, activity level is linked to body size response to warming. This provides an 490 interesting connection between fishing, changing temperature, activity level and body size, 491 and warrants for further research into the potential synergistic effects of multiple stressors. 492 Changes in reproduction 493 A commonly used proxy for individual fitness is the lifetime reproductive success (LRS), 494 which can simply be described as the total number of offspring an individual produces during 495 its lifetime (Mousseau and Roff 1987). The reproductive output of an individual tends to 496 increase with body size (Hixon et al. 2014), thus contributing to the individual fitness. In a 497 meta-analysis of 342 species of marine fishes from 15 different orders, Barneche et al. (2018) 498 quantified this increase, and estimated how selection towards smaller body size may reduce 499 fecundity. They controlled for phylogenetic nonindependence and scaled female mass and 500 reproductive energy output, including fecundity, egg volume and egg energy, to show that 501 large females reproduce disproportionally more than small females. Similar trend has been 502 seen in experimental studies, where positively size-selected harvest has led to reduction in 503 fecundity (lower spawning probability, fewer eggs both absolutely and relatively) in zebrafish 504 (Uusi-Heikkilä et al. 2015). Similarly, a study on the Atlantic silverside subjected to five 505 generations of size-selective harvest showed marked reductions in egg volume, larval size at 506 hatching, larval growth rate, and larval survival probability (Walsh et al. 2006). The 507 correlation of body size and fecundity has been documented in various fish species both intra508 (Dick et al. 2017) and inter-specifically (Savage et al. 2004b; Hayward and Gillooly 2011). It 509 is important to note, however, that the strength of the correlation depends on the species in 510 question (Barneche et al. 2018), and that like other animals in the wild (Nussey et al. 2013), 511 at least some fish experience actuarial (Beverton et al. 2004; Uriarte et al. 2016) and Page 21 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 22 512 reproductive (Reznick et al. 2006; Benoît et al. 2018) senescence, meaning that the 513 reproductive capacity of fish may not increase indefinitely with size. Therefore, predictions 514 about reproductive capacity based on body size alone should be made with caution. 515 The mechanisms of how larger body size benefits the reproductive output vary, however, and 516 some are related to behaviour. In coho salmon (O. kisutch), larger female body size has been 517 associated with an advantage in mate and territory selection (Fleming and Gross 1994). 518 Bigger size appears to be beneficial, as it allows for increased egg production, better access to 519 nesting territories, and improved nest digging (Fleming and Gross 1994). Further, the 520 changes in body size are often accompanied with associated changes in population age 521 structure. The truncation of age structure can potentially enhance the effects of declining 522 body size at individual and population level. As an example, longevity may be advantageous 523 as it can increase population recruitment variability (in timing), particularly in long-lived fish 524 species, thereby making the population less vulnerable in unstable environments (Longhurst 525 2002). Maternal age can also contribute to the fitness of offspring. The larvae of older black 526 rockfish (Sebastes melanops) showed over three times faster growth rate than the larvae of 527 younger rockfish, and they survived starvation better (Berkeley et al. 2004). Since body size 528 and reproductive output are often positively correlated (Barneche et al. 2018), changes in 529 individual body size can have multigenerational impacts, truncate the population age and size 530 structure, and potentially lower the population growth rate. 531 532 How do changes in body size translate to population dynamics? 533 The intrinsic rate of per-capita population growth rate (r) can be thought of as a population 534 level analogue to LRS. Although r cannot be directly translated from LRS, body size is a 535 strong correlate for both (Denney et al. 2002; Anderson et al. 2008). Indeed, Uusi-Heikkilä et Page 22 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 23 536 al. (2015) modelled their results from the zebrafish studies, and found that in the absence of 537 fishing, positively size-selected (large individuals removed) population had a slower 538 population growth rate than negatively and randomly size-selected populations. In their 539 modelling study, Audzijonyte and Kuparinen (2016) showed that earlier maturation that 540 evolved during fishing, reduced r after fishing had been halted. They also showed that 541 decreased adult body size reduced the reproductive output and increased adult natural 542 mortality. However, species’ age-specific survival and trophic position was also important in 543 determining r: a species inhabiting offshore environment and exposed to few predators did 544 not suffer, in terms of population growth rate, as much from reduced body size as a species 545 living near shore and being intensively predated by other fish. A density dependent 546 population response to fishing was predicted by the model of Dunlop et al. (2015), who 547 observed that as fishing began, r first declined as the largest individuals were removed from 548 the reproductive pool, but once density dependent processes started regulating the population, 549 r and reproductive output increased. After moratorium, the opposite occurred: at first r 550 increased temporarily, but was eventually counter-acted by density dependent processes, 551 leading to reduction in r (Dunlop et al. 2015). This kind of delayed response can explain how 552 evolution may impede stock recovery after harvest (Dunlop et al. 2015). 553 The form of selection will affect the outcome of the selection: stabilising selection favours 554 the intermediate phenotype, and disruptive selection favours the extreme phenotypes. 555 Directional selection occurs when selection favours one phenotype over another so that the 556 frequency of this phenotype in a population becomes higher than before selection. On the one 557 hand, studies have shown that stabilising selection can lead to decreasing phenotypic and 558 genetic variation (Olsen et al. 2009), which is known to reduce population fitness (Reed and 559 Frankham 2003). On the other hand, if the selection is disruptive, it may increase the genetic 560 variance in a population (Edeline et al. 2009), potentially increasing population fitness. Thus, Page 23 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 30 709 use. In the lakes with no pike present, perch utilised both the hypolimnion and epilimnion 710 habitats and thuswere also found below the thermocline. The authors attributed these 711 differences to the differences in the size when perch reached size refuge, illustrating how 712 size-dependent predation can feed back to the ecosystem function by altering the habitat use 713 of the species (Persson et al. 1995). Growth rate and body size may even determine who 714 becomes a prey and who is a predator. A study by Nilsson et al. (2019) documented a 715 predator-prey role reversal for pike and stickleback. While pike is usually a predator and 716 stickleback is a prey, they provided evidence that juvenile pike mortality is size-dependant 717 and corresponds to stickleback gape limitation (Nilsson et al. 2019). In the presence of 718 increasing water temperature or human induced changes in body size and growth rate, this 719 kind of predator-prey role reversals may add to the hindrance of population recovery. 720 While size-dependent predation can have ecological effects on the environment as described 721 in the above study by Persson et al. (1995), the cause and effect is likely more complex. Size722 related evolutionary and ecological changes can also drive changes in the ecosystem, and 723 these may further feed back into the ecosystem. In their experimental study, Palkovacs et al. 724 (2009) compared the effects of ecological (invasive guppies, P. reticulata), evolutionary 725 (different guppy life-histories as a result of different predation pressure) and co-evolutionary 726 (coevolution of guppies and the small fish Rivulus) agents driving ecosystem change in 727 Trinidadian river ecosystems. They concluded that guppy evolution and guppy-Rivulus 728 coevolution caused significant changes in the surrounding ecosystem, while guppy invasion 729 did not. Another example of evolutionary change affecting the ecosystem comes from the 730 gape-limited predatory alewife (Alosa pseudoharengus), which has diverged in its foraging 731 traits (Palkovacs and Post 2009). Anadromous alewives are capable of preying on larger prey 732 than their landlocked counterparts, and this has in turn significantly affected the size structure 733 of its prey, crustacean zooplankton (Palkovacs and Post 2009). These studies provide Page 30 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 31 734 evidence that evolutionary changes in one species can have an impact on the entire 735 ecosystem. Human-induced evolutionary changes on various species and their effects on 736 ecosystems have further been reviewed by Palkovacs et al. (2011) and Fraser (2013). 737 Trophic cascade 738 The tale of Atlantic cod is an iconic example of how changes have propagated through 739 different levels of biological organisation in nature. The intensely fished northern cod 740 population in southern Labrador and eastern Newfoundland collapsed in the early 1990s, and 741 despite a decades long moratorium, the stocks have not fully recovered (Neuenhoff et al. 742 2019; Sguotti et al. 2019). The reasons for the lack of recovery are complex, and include 743 several hypotheses such as changing sea water temperatures, predation by seals, continued 744 fishing, and life-history changes due to fishing (Swain et al. 2011). Olsen et al. (2004) 745 showed that the fishery collapse was preceded by a rapid phenotypic change toward younger 746 age and smaller size at maturity, and the study strongly suggested there was an evolutionary 747 component behind this change. It is likely that this has contributed to the slow recovery of the 748 stocks (Olsen et al. 2004), and similar trends have been recorded in many other top predators 749 (Hutchings and Baum 2005). The collapse of cod and other large predator populations in the 750 northwest Atlantic marine ecosystem led to a cascade involving four trophic levels (Frank et 751 al. 2005). Following the collapse of large predators, the abundance of smaller fish and 752 benthic macroinvertebrates increased markedly. The increase in smaller fish in turn 753 influenced the abundance of zooplankton and phytoplankton, followed by a response in 754 nitrate concentrations. This top-down trophic cascade has likely further impeded the recovery 755 of the cod population by altering food webs (Frank et al. 2005). 756 Similarly, in the Northeast Pacific, marked declines in the abundance of the oldest and largest 757 individuals of Chinook salmon (O. tshawytscha), and changes in the age and size structure of Page 31 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 32 758 the population coincide with a decline in the length-at-age of the fish (Ohlberger et al. 2017). 759 Intriguingly, while the decrease in the length-at-age was clear, the cause of it was not. 760 Ohlberger et al. (2017) hypothesised that the changes could be a result of size-selective 761 harvesting, changing temperatures, interspecific competition, or, more likely, predation by 762 large apex predators such as the resident killer whale, the numbers of which have increased 763 following a harvest ban in the 1970s. The loss of the oldest and largest individuals of the 764 Chinook salmon population are of concern because it can have a negative impact on the long765 term viability of the population (Ohlberger et al. 2017). Another ecosystem-level example 766 comes from the Northwest Atlantic, where the combination of size-selective harvest and 767 changing climate caused reductions in the body sizes of functional top predators, which in 768 turn contributed to the increase in prey biomass (Shackell et al. 2009). This demographic 769 change was also weakly associated with a decline in zooplankton and an increase in 770 phytoplankton abundance, indicating a cascading effect through the food web involving three 771 trophic levels (Shackell et al. 2009). 772 As discussed, not only fishing, but also changing water temperature has been associated with 773 shifts towards smaller body size in fish, affecting populations and communities. A 774 comprehensive meta-analysis by Daufresne et al. (2009) demonstrated an increase in the 775 proportion of small-sized species with increasing temperature, in terms of both species 776 richness and abundance (Daufresne et al. 2009). The latter result includes a latitude-related 777 component, and is in line with not only the temperature-size rule, but also ecogeographical 778 rules such as the Bergmann’s rule or the James’ rule (Daufresne et al. 2009). Warming779 induced changes in body size and resulting consequences at the population and community 780 level have also been reviewed by Ohlberger (2013). These are important studies showing how 781 the changing climate can have an impact at individual, population and community level, and 782 even change the species composition of an ecosystem. Page 32 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 33 783 The explicit demonstration of changes propagating through different levels of biological 784 organisation is challenging, but in addition to empirical examples discussed above, modelling 785 and microcosm studies are showing similar trends. In a modelling study, Kuparinen et al. 786 (2016) used a network model to demonstrate how declining body size together with advanced 787 maturation may magnify fluctuations in fished populations. They showed that this fluctuation 788 can extend to other species in the ecosystem, and may continue long after fishing has ceased 789 (Kuparinen et al. 2016). A microcosm study on predator-prey (rotifer – algae) interactions 790 showed that the prey evolution affected the growth of predator population more than the 791 actual abundance of prey, and that the changes in predator abundance in turn affected the 792 prey evolution, suggesting an eco-evolutionary feedback loop (Becks et al. 2012). While 793 these particular results cannot be directly extrapolated to fish communities, changes in fish 794 body size are likely to influence other species too, especially when predation is gape-limited. 795 Indeed, examples of parallel evolution in fishes such as the cichlids, threespine sticklebacks, 796 and guppies are illuminating the tight linkage there exists between the genetics of an 797 individual and the ecological environment surrounding it (Elmer and Meyer 2011). 798 799 Conclusion 800 Changes in body size at an individual level can propagate through different levels of 801 biological organisation. Given the current rapid environmental change, organisms will have 802 to adjust, adapt or move, probably quicker than ever before. Adaptation requires evolutionary 803 change, and evolutionary change requires building material: genetic variability. Evolutionary 804 change can occur at ecologically relevant time scales, and therefore it is vital for conservation 805 biologists and decision makers to understand both the ecology of an organism, and the 806 genetic architecture that enables evolution. Changes in body size and correlated life history Page 33 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 34 807 characters following harvest, changing temperature and dissolved oxygen, as well as 808 predation have been demonstrated empirically (Nusslé et al. 2009, 2011; Baudron et al. 2014; 809 Hunter et al. 2015; Ohlberger et al. 2017), experimentally (Reznick et al. 1990; Conover and 810 Munch 2002; van Wijk et al. 2013; Uusi-Heikkilä et al. 2015), as well as through 811 mathematical modelling (Kuparinen et al. 2016). Consequently, the outcomes of these 812 changes can vary from behavioural changes (McCormick 2009; Sutter et al. 2012) to changes 813 in reproductive output (Hixon et al. 2014), and can extend their influence to the next 814 generation (van Wijk et al. 2013), and the entire ecosystem (Frank et al. 2005) regardless of 815 whether the change is genetic or not (Bolnick et al. 2011). The genetic basis of the change 816 matters, however, because first, genetic change allows for the evolutionary adaptation to the 817 changing environment, and second, evolutionary change may be hard if not impossible to 818 reverse (Salinas et al. 2012; Uusi-Heikkilä et al. 2017). Should the changes be indeed genetic, 819 and therefore likely hard to reverse, the consequences that extend from declining body 820 through to population size and stability (Audzijonyte and Kuparinen 2016) to ecosystem 821 functions (Frank et al. 2005; Ohlberger et al. 2017), may also be hard to reverse. 822 In addition to reversibility, understanding the genetic basis of the changes occurring in nature 823 is vital from the biodiversity management perspective. From the multiple stressors affecting 824 our oceans today, some are easier controlled than others. Managing fisheries selection 825 regimes, for instance, is easier and quicker to control than changing climate. If we utilise the 826 genetic knowledge and fishing methods that conserve variation, we may be maintaining a 827 higher genetic variability (Edeline et al. 2009), as well as population stability (Bolnick et al. 828 2011). Higher genetic variability, in turn, is what forms the basis for evolutionary potential. 829 In this way, we may better conserve the biodiversity not only now, but also in the future. 830 Page 34 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 35 831 Acknowledgements 832 This work was supported by funding from the Emil Aaltonen Foundation (PAA), the 833 Academy of Finland AK (grant no. 317495), SUH (grant no. 325107), Discovery Grant from 834 the National Sciences and Engineering Research Council of Canada (NSERC to AK), the 835 European Research Council (COMPLEX-FISH 400820 to AK), and the Finnish Cultural 836 Foundation (SUH). The authors thank the two anonymous referees as well as the editing 837 team. 838 839 840 841 842 843 844 845 846 847 848 849 850 851 Page 35 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 36 852 853 References: 854 Agrawal, A.A. 2001. Phenotypic Plasticity in the Interactions and Evolution of Species. 855 Science. 294: 321–327. doi:10.1126/science.1060701. 856 Allain, G., Petitgas, P., Grellier, P., and Lazure, P. 2003. The selection process from larval to 857 juvenile stages of anchovy (Engraulis encrasicolus) in the Bay of Biscay investigated by 858 Lagrangian simulations and comparative otolith growth. Fish. Oceanogr. 12(4–5): 407– 859 418. doi:10.1046/j.1365-2419.2003.00245.x. 860 Allendorf, F.W., and Hard, J.J. 2009. Human-induced evolution caused by unnatural 861 selection through harvest of wild animals. PNAS. 3: 9987–9994. doi:10.17226/12692. 862 Anderson, C.N.K., Chih-hao, H., Sandin, S.A., Hewitt, R., Hollowed, A., Beddington, J., 863 May, R.M., and Sugihara, G. 2008. Why fishing magnifies fluctuations in fish 864 abundance. Nature. 452: 835–839. doi:10.1038/nature06851. 865 Andrews, J.W., Murai, T., and Gibbons, G. 1973. The influence of dissolved oxygen on the 866 growth of channel catfish. Trans. Am. Fish. Soc. 102(4): 835–838. doi:10.1577/1548867 8659(1973)102<835. 868 Arandia-Gorostidi, N., Huete-Stauffer, T.M., Alonso-Saez, L., and Moran, X.A.G. 2017. 869 Testing the metabolic theory of ecology with marine bacteria: different temperature 870 sensitivity of major phylogenetic groups during the spring phytoplankton bloom. 871 Environ. Microbiol. 19(11): 4493–4505. doi:10.1111/1462-2920.13898. 872 Ashton, K.G., and Feldman, C.R. 2003. Bergmann’s rule in nonavian reptiles: turtles follow 873 it, lizards and snakes reverse it. Evolution. 57(5): 1151–1163. doi:10.1111/j.0014874 3820.2003.tb00324.x. Page 36 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 37 875 Atkinson, D. 1994. Temperature and organism size - a biological law for ectotherms. Adv. 876 Ecol. Res. 25: 1–58. doi:10.1016/S0065-2504(08)60212-3. 877 Audzijonyte, A., Barneche, D.R., Baudron, A.R., Belmaker, J., Clark, T.D., Marshall, C.T., 878 Morrongiello, J.R., and van Rijn, I. 2019. Is oxygen limitation in warming waters a valid 879 mechanism to explain decreased body sizes in aquatic ectotherms? Glob. Ecol. 880 Biogeogr. 28(2): 64–77. doi:10.1111/geb.12847. 881 Audzijonyte, A., and Kuparinen, A. 2016. The role of life histories and trophic interactions in 882 population recovery. Conserv. Biol. 30(4): 734–743. doi:10.1111/cobi.12651. 883 Audzijonyte, A., Kuparinen, A., and Fulton, E.A. 2013. How fast is fisheries-induced 884 evolution? Quantitative analysis of modelling and empirical studies. Evol. Appl. 6(4): 885 585–595. doi:10.1111/eva.12044. 886 Bailey, K.M., Ciannelli, L., Hunsicker, M., Rindorf, A., Neuenfeldt, S., and Mo, C. 2010. 887 Comparative analysis of marine ecosystems: workshop on predator – prey interactions. 888 Biol. Lett. 6: 579–581. doi:10.1098/rsbl.2010.0326. 889 Bailey, N., Bailey, D.M., Bellini, L.C., Fernandes, P.G., Fox, C., S, H., Holmes, S., Howe, J., 890 Hughes, S., Magill, S., McIntyre, F., McKee, D., Ryan, M.R., Smith, I.P., Tyldesley, G., 891 Watret, R., and Turrell, W.R. 2011. The West of Scotland marine ecosystem: A review 892 of scientific knowledge. Mar. Scotl. Sci. Rep. 0911: 1–288. 893 Bailly, D., Cassemiro, F.A.S., Agostinho, C.S., Marques, E.E., and Agostinho, A.A. 2014. 894 The metabolic theory of ecology convincingly explains the latitudinal diversity gradient 895 of Neotropical freshwater fish. Ecology 95(2): 553–562. doi: 10.1890/13-0483.1. 896 Barneche, D.R., Kulbicki, M., Floeter, S.R., Friedlander, A.M., and Allen, A.P. 2016. 897 Energetic and ecological constraints on population density of reef fishes. Proc. R. Soc. B Page 37 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 38 898 283: 20152186. http://dx.doi.org/10.1098/rspb.2015.2186. 899 Barneche, D.R., Robertson, D.R., White, C.R., and Marshall, D.J. 2018. Fish reproductive900 energy output increases disproportionately with body size. Science. 360: 642–645. doi: 901 10.1126/science.aao6868. 902 Barnett, T.P., Pierce, D.W., AchutaRao, K.M., J, G.P., D, S.B., Gregory, J.M., and 903 Washington, W.M. 2005. Penetration of human-induced warming into the world’s 904 oceans. Science. 309: 284–287. doi:10.1126/science.1112418. 905 Barson, N.J., Aykanat, T., Hindar, K., Baranski, M., Bolstad, G.H., Fiske, P., Jacq, C., 906 Jensen, A.J., Johnston, S.E., Karlsson, S., Kent, M., Moen, T., Niemelä, E., Nome, T., 907 Næsje, T.F., Orell, P., Romakkaniemi, A., Sægrov, H., Urdal, K., Erkinaro, J., Lien, S., 908 and Primmer, C.R. 2015. Sex-dependent dominance at a single locus maintains variation 909 in age at maturity in salmon. Nature 528: 405–408. doi:10.1038/nature16062. 910 Baudron, A.R., Needle, C.L., and Marshall, C.T. 2011. Implications of a warming North Sea 911 for the growth of haddock Melanogrammus aeglefinus. J. Fish Biol. 78(7): 1874–1889. 912 doi:10.1111/j.1095-8649.2011.02940.x. 913 Baudron, A.R., Needle, C.L., Rijnsdorp, A.D., and Tara Marshall, C. 2014. Warming 914 temperatures and smaller body sizes: Synchronous changes in growth of North Sea 915 fishes. Glob. Chang. Biol. 20(4): 1023–1031. doi:10.1111/gcb.12514. 916 Becks, L., Ellner, S.P., Jones, L.E., and Hairston, N.G. 2012. The functional genomics of an 917 eco-evolutionary feedback loop: Linking gene expression, trait evolution, and 918 community dynamics. Ecol. Lett. 15(5): 492–501. doi:10.1111/j.1461919 0248.2012.01763.x. 920 Belk, M.C., and Houston, D.D. 2002. Bergmann’s rule in ectotherms: A test using freshwater Page 38 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 39 921 fishes. Am. Nat. 160(6): 803–808. doi:10.1086/343880. 922 Benoît, H.P., Swain, D.P., Hutchings, J.A., Knox, D., Doniol-Valcroze, T., and Bourne, C.M. 923 2018. Evidence for reproductive senescence in a broadly distributed harvested marine 924 fish. Mar. Ecol. Prog. Ser. 592: 207–224. doi:10.3354/meps12532. 925 Berkeley, S.A., Chapman, C., and Sogard, S.M. 2004. Maternal Age as a Determinant of 926 Larval Growth and Survival in a Marine Fish, Sebastes Melanops. Ecology 85(5): 1258– 927 1264. doi:10.1890/03-0706. 928 Bergmann, C. 1847. About the relationships between heat conservation and body size of 929 animals. Goett. Stud. (Original in German) 1:595–708. 930 Bernatchez, L. 2016. On the maintenance of genetic variation and adaptation to 931 environmental change: considerations from population genomics in fishes. J. Fish Biol. 932 89(6): 2519–2556. doi:10.1111/jfb.13145. 933 Beverton, R.J.H., Hylen, A., Østvedt, O.J., Alvsvaag, J., and Iles, T.C. 2004. Growth, 934 maturation, and longevity of maturation cohorts of Norwegian spring-spawning herring. 935 ICES J. Mar. Sci. 61(2): 165–175. doi:10.1016/j.icesjms.2004.01.001. 936 Biro, P.A., Abrahams, M. V., Post, J.R., and Parkinson, E.A. 2004. Predators select against 937 high growth rates and risk-taking behaviour in domestic trout populations. Proc. R. Soc. 938 B Biol. Sci. 271(1554): 2233–2237. doi:10.1098/rspb.2004.2861. 939 Biro, P.A., and Post, J.R. 2008. Rapid depletion of genotypes with fast growth and bold 940 personality traits from harvested fish populations. Proc. Natl. Acad. Sci. 105(8): 2919– 941 2922. doi:10.1073/pnas.0708159105. 942 Blanchet, S., Beauchard, O., and Tedesco, P.A. 2010. Non-native species disrupt the 943 worldwide patterns of freshwater fish body size: implications for Bergmann’s rule. Ecol. Page 39 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 46 1082 Heino, M., Baulier, L., Boukal, D.S., Ernande, B., Johnston, F.D., Mollet, F.M., Pardoe, H., 1083 Therkildsen, N.O., Uusi-Heikkilä, S., Vainikka, A., Arlinghaus, R., Dankel, D.J., 1084 Dunlop, E.S., Eikeset, A.M., Enberg, K., Engelhard, G.H., Jørgensen, C., Laugen, A.T., 1085 Masumura, S., Nusslé, S., Urbach, D., Whitlock, R., Rijnsdorp, A.D., and Dieckmann, 1086 U. 2013. Can fisheries-induced evolution shift reference points for fisheries 1087 management? ICES J. Mar. Sci. 70(4): 707–721. doi:10.1093/icesjms/fst077. 1088 Heino, M., Díaz Pauli, B., and Dieckmann, U. 2015. Fisheries-Induced Evolution. Annu. 1089 Rev. Ecol. Evol. Syst. 46(1): 461–480. doi:10.1146/annurev-ecolsys-112414-054339. 1090 Heins, D.C., Knoper, H., and Baker, J.A. 2016. Consumptive and non-consumptive effects of 1091 predation by introduced northern pike on life-history traits in threespine stickleback. 1092 Evol. Ecol. Res. 17(3): 355–372. 1093 Hixon, M.A., Johnson, D.W., and Sogard, S.M. 2014. BOFFFFs: on the importance of 1094 conserving old-growth age structure in fishery populations. ICES J. Mar. Sci. 71(8): 1095 2171–2185. doi:10.1093/icesjms/fst200. 1096 Hoffmann, A.A., and Willi, Y. 2008. Detecting genetic responses to environmental change. 1097 Nat. Rev. Genet. 9(6): 421–432. doi:10.1038/nrg2339. 1098 Hollins, J., Thambithurai, D., Köeck, B., Crespel, A., Bailey, D.M., Cooke, S.J., Lindström, 1099 J., Parsons, K.J., and Killen, S.S. 2018. A Physiological perspective on fisheries-induced 1100 evolution. Evol. Appl. 11: 561-576. doi:10.1111/eva.12597. 1101 Hooper, D.U., Chapin, F.S., Ewel, J.J., Hector, A., Inchausti, P., Lavorel, S., Lawton, J.H., 1102 Lodge, D.M., Loreau, M., Naeem, S., Schmid, B., Setälä, H., Symstad, A.J., 1103 Vandermeer, J., and Wardle, D.A. 2005. Effects of biodiversity on ecosystem 1104 functioning: A consensus of current knowledge. Ecol. Monogr. 75(1): 3–35. 1105 doi:10.1890/04-0922. Page 46 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 47 1106 Hunter, A., Speirs, D.C., and Heath, M.R. 2015. Fishery-induced changes to age and length 1107 dependent maturation schedules of three demersal fish species in the Firth of Clyde. 1108 Fish. Res. 170: 14–23. doi:10.1016/j.fishres.2015.05.004. 1109 Hutchings, J.A., and Baum, J.K. 2005. Measuring marine fish biodiversity: Temporal 1110 changes in abundance, life history and demography. Philos. Trans. R. Soc. B Biol. Sci. 1111 360(1454): 315–338. doi:10.1098/rstb.2004.1586. 1112 Hutchinson, W.F., Van Oosterhout, C., Rogers, S.I., and Carvalho, G.R. 2003. Temporal 1113 analysis of archived samples indicates marked genetic changes in declining North Sea 1114 cod (Gadus morhua). Proc. R. Soc. B Biol. Sci. 270(1529): 2125–2132. 1115 doi:10.1098/rspb.2003.2493. 1116 Jackson, J.B.C., Kirby, M.X., Berger, W.H., Bjorndal, K.A., Botsford, L.W., Bourque, B.J., 1117 Bradbury, R.H., Cooke, R., Erlandson, J., Estes, J.A., Hughes, T.P., Kidwell, S., Lange, 1118 C.B., Lenihan, H.S., Pandolfi, J.M., Peterson, C.H., Steneck, R.S., Tegner, M.J., and 1119 Warner, R.R. 2001. Historical overfishing and the recent collapse of coastal ecosystems. 1120 Science. 293(5530): 629–637. doi:10.1126/science.1059199. 1121 Jakobsdóttir, K.B., Pardoe, H., Magnússon, Á., Björnsson, H., Pampoulie, C., Ruzzante, 1122 D.E., and Marteinsdóttir, G. 2011. Historical changes in genotypic frequencies at the 1123 Pantophysin locus in Atlantic cod (Gadus morhua) in Icelandic waters: Evidence of 1124 fisheries-induced selection? Evol. Appl. 4(4): 562–573. doi:10.1111/j.17521125 4571.2010.00176.x. 1126 James, F. 1970. Geographic size variation in birds and its relationship to climate. Ecology 1127 51:365–390. 1128 Jensen, A.L. 1996. Beverton and Holt life history invariants result from optimal trade-off of 1129 reproduction and survival. Can. J. Fish. Aquat. Sci. 53(4): 820–822. doi:10.1139/f95Page 47 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 48 1130 233. 1131 Johansen, J.L., Pratchett, M.S., Messmer, V., Coker, D.J., Tobin, A.J., and Hoey, A.S. 2015. 1132 Large predatory coral trout species unlikely to meet increasing energetic demands in a 1133 warming ocean. Sci. Rep. 5: 1–8. doi:10.1038/srep13830. 1134 Killen, S., Nati, J., and Suski, C. 2015. Catchability of individual fish by trawling is driven by 1135 capacity for anaerobic metabolism. Proc. R. Soc. B Biol. Sci. 282: 20150603. doi: 1136 10.1098/rspb.2015.0603. 1137 Kleiber, M. 1932. Body size and metabolism. Hilgardia 6(11): 315 - 353. 1138 Knouft, J.H. 2004. Latitudinal variation in the shape of the species body size distribution : an 1139 analysis using freshwater fishes. Oecologia 139: 408–417. doi:10.1007/s00442-0041140 1510-x. 1141 Kozlowski, J., and Konarzewski, M. 1998. Is West, Brown and Enquist’s model of allometric 1142 scaling mathematically correct and biologically relevant? Funct. Ecol. 18: 283–289. 1143 doi:10.1111/j.0269-8463.2004.00830.x. 1144 Kozlowski, J., and Konarzewski, M. 2005. West, Brown and Enquist’s model of allometric 1145 scaling again: the same questions remain. Funct. Ecol. 19: 739–743. doi:10.1111/j.13651146 2435.2005.01021.x. 1147 Kuparinen, A., Boit, A., Valdovinos, F.S., Lassaux, H., and Martinez, N.D. 2016. Fishing1148 induced life-history changes degrade and destabilize harvested ecosystems. Sci. Rep. 1149 6(1): 22245. doi:10.1038/srep22245. 1150 Kuparinen, A., and Hutchings, J.A. 2016. Genetic architecture of age at maturity can generate 1151 either directional or divergent and disruptive harvest-induced evolution. Philos. Trans. 1152 R. Soc. B Biol. Sci. 372: 20160035. doi:10.1098/rstb.2016.0035. Page 48 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 49 1153 Kuparinen, A., Kuikka, S., and Merilä, J. 2009. Estimating fisheries-induced selection: 1154 Traditional gear selectivity research meets fisheries-induced evolution. Evol. Appl. 2(2): 1155 234–243. doi:10.1111/j.1752-4571.2009.00070.x. 1156 Lee, W.S., Monaghan, P., and Metcalfe, N.B. 2013. Experimental demonstration of the 1157 growth rate-lifespan trade-off. Proc. R. Soc. B Biol. Sci. 280:20122370. 1158 doi:10.1098/rspb.2012.2370. 1159 Lefevre, S., McKenzie, D.J., and Nilsson, G.E. 2017. Models projecting the fate of fish 1160 populations under climate change need to be based on valid physiological mechanisms. 1161 Glob. Chang. Biol. 23(9): 3449–3459. doi:10.1111/gcb.13652. 1162 Longhurst, A. 2002. Murphy’s law revisited: longevity as a factor in recruitment to fish 1163 populations. Fish. Res. 56(2): 125–131. doi: 10.1016/S0165-7836(01)00351-4. 1164 Lush, J. L. 1937. Animal Breeding Plans, Collegiate Press Inc, Ames, IA. 1165 MacArthur, R.H., and Pianka, E.R. 1966. On optimal use of a patchy environment. Am. Nat. 1166 100(916): 603–609. doi:10.1086/282454. 1167 Mainka, S.A., and Howard, G.W. 2010. Climate change and invasive species: Double 1168 jeopardy. Integr. Zool. 5(2): 102–111. doi:10.1111/j.1749-4877.2010.00193.x. 1169 Marty, L., Dieckmann, U., and Ernande, B. 2015. Fisheries-induced neutral and adaptive 1170 evolution in exploited fish populations and consequences for their adaptive potential. 1171 Evol. Appl. 8(1): 47–63. doi:10.1111/eva.12220. 1172 McCormick, M.I. 2009. Behaviourally mediated phenotypic selection in a disturbed coral 1173 reef environment. PLoS One 4(9): e7096. doi:10.1371/journal.pone.0007096. 1174 Meekan, M.G., Carleton, J.H., McKinnon, A.D., Flynn, K., and Furnas, M. 2003. What 1175 determines the growth of tropical reef fish larvae in the plankton: Food or temperature? Page 49 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 50 1176 Mar. Ecol. Prog. Ser. 256: 193–204. doi:10.3354/meps256193. 1177 Meekan, M.G., Vigliola, L., Hansen, A., Doherty, P.J., Halford, A., and Carleton, J.H. 2006. 1178 Bigger is better: Size-selective mortality throughout the life history of a fast-growing 1179 clupeid, Spratelloides gracilis. Mar. Ecol. Prog. Ser. 317: 237–244. 1180 doi:10.3354/meps317237. 1181 Meiri, S., Dayan, T., and Aviv, T. 2003. On the validity of Bergmann’s rule. J. Biogeogr. 30: 1182 331–351. doi:10.1046/j.1365-2699.2003.00837.x. 1183 Metcalfe, N.B., and Monaghan, P. 2001. Compensation for a bad start: Grow now, pay later? 1184 Trends Ecol. Evol. 16(5): 254–260. doi:10.1016/S0169-5347(01)02124-3. 1185 Metcalfe, N.B., and Monaghan, P. 2003. Growth versus lifespan: Perspectives from 1186 evolutionary ecology. Exp. Gerontol. 38(9): 935–940. doi:10.1016/S05311187 5565(03)00159-1. 1188 Michalsen, K., Ottersen, G., and Nakken, O. 1998. Growth of northeast Arctic cod in relation 1189 to ambient temperature. ICES J. Mar. Sci. 55: 863–877. doi: 10.1006/jmsc.1998.0364. 1190 Miller T.J., Crowder L.B., Rice J.A., Marschall, E.A. 1988 Larval size and recruitment 1191 mechanisms in fishes: toward a conceptual framework. Can. J. Fish. Aquat. Sci. 1192 45:1657–1670. 1193 Moore, C.J. 2008. Synthetic polymers in the marine environment: A rapidly increasing, long1194 term threat. Environ. Res. 108(2): 131–139. doi:10.1016/j.envres.2008.07.025. 1195 Morrongiello, J.R., Sweetman, P.C., and Thresher, R.E. 2019. Fishing constrains phenotypic 1196 responses of marine fish to climate variability. J. Anim. Ecol. 88(11): 1645–1656. 1197 doi:10.1111/1365-2656.12999. 1198 Mousseau, T.A., and Roff, D.A. 1987. Natural selection and the heritability of fitness Page 50 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 51 1199 components. Heredity. 59(59): 181–197. doi:10.1038/hdy.1987.113. 1200 Murawski, S.A. 2010. Rebuilding depleted fish stocks: the good, the bad, and, mostly, the 1201 ugly. ICES J. Mar. Sci. 67(9): 1830–1840. doi:10.1093/icesjms/fsq125. 1202 Neuenhoff, R.D., Swain, D.P., Cox, S.P., McAllister, M.K., Trites, A.W., Walters, C.J., and 1203 Hammill, M.O. 2019. Continued decline of a collapsed population of Atlantic cod 1204 (Gadus morhua) due to predation-driven Allee effects. Can. J. Fish. Aquat. Sci. 76(1): 1205 168–184. doi:10.1139/cjfas-2017-0190. 1206 Nilsson, J., Flink, H., and Tibblin, P. 2019. Predator–prey role reversal may impair the 1207 recovery of declining pike populations. J. Anim. Ecol. 88: 927–939. doi:10.1111/13651208 2656.12981. 1209 Nordhaus, I., Roelke, D.L., Vaquer-Sunyer, R., and Winter, C. 2018. Coastal systems in 1210 transition: From a ‘natural’ to an ‘anthropogenically-modified’ state. Estuar. Coast. 1211 Shelf Sci. 211: 1–5. doi:10.1016/j.ecss.2018.08.001. 1212 Nussey, D.H., Froy, H., Lemaitre, J.F., Gaillard, J.M., and Austad, S.N. 2013. Senescence in 1213 natural populations of animals: Widespread evidence and its implications for bio1214 gerontology. Ageing Res. Rev. 12(1): 214–225. doi:10.1016/j.arr.2012.07.004. 1215 Nusslé, S., Bornand, C.N., and Wedekind, C. 2009. Fishery-induced selection on an Alpine 1216 whitefish: Quantifying genetic and environmental effects on individual growth rate. 1217 Evol. Appl. 2(2): 200–208. doi:10.1111/j.1752-4571.2008.00054.x. 1218 Nusslé, S., Bréchon, A., and Wedekind, C. 2011. Change in individual growth rate and its 1219 link to gill-net fishing in two sympatric whitefish species. Evol. Ecol. 25(3): 681–693. 1220 doi:10.1007/s10682-010-9412-3. 1221 Ohlberger, J. 2013. Climate warming and ectotherm body size - from individual physiology Page 51 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 52 1222 to community ecology. Funct. Ecol. 27(4): 991–1001. doi:10.1111/1365-2435.12098. 1223 Ohlberger, J., Ward, E.J., Schindler, D.E., and Lewis, B. 2017. Demographic changes in 1224 Chinook salmon across the Northeast Pacific Ocean. Fish Fish. 19: 533–546. 1225 doi:10.1111/faf.12272. 1226 Olsen, E.M., Carlson, S.M., Gjøsæter, J., and Stenseth, N.C. 2009. Nine decades of 1227 decreasing phenotypic variability in Atlantic cod. Ecol. Lett. 12(7): 622–631. 1228 doi:10.1111/j.1461-0248.2009.01311.x. 1229 Olsen, E.M., Heino, M., Lilly, G.R., Morgan, M.J., Brattey, J., Ernande, B., and Dieckmann, 1230 U. 2004. Maturation trends indicative of rapid evolution preceded the collapse of 1231 northern cod. Nature 428: 4–7. doi:10.1038/nature02453.1. 1232 Olsen, E.M., and Moland, E. 2011. Fitness landscape of Atlantic cod shaped by harvest 1233 selection and natural selection. Evol. Ecol. 25(3): 695–710. doi:10.1007/s10682-0101234 9427-9. 1235 Orr, J.C., Fabry, V.J., Aumont, O., Bopp, L., Doney, S.C., Feely, R.A., Gnanadesikan, A., 1236 Gruber, N., Ishida, A., Joos, F., Key, R.M., Lindsay, K., Maier-Reimer, E., Matear, R., 1237 Monfray, P., Mouchet, A., Najjar, R.G., Plattner, G.-K., Rodgers, K.B., Sabine, C.L., 1238 Sarmiento, J.L., Schlitzer, R., Slater, R.D., Totterdell, I.J., Weirig, M.-F., Yamanaka, Y., 1239 and Yool, A. 2005. Anthropogenic ocean acidification over the twenty-first century and 1240 its impact on calcifying organisms. Nature 437(4): 681–686. doi:10.1038/nature04095. 1241 Otterlei, E., Nyhammer, G., Folkvord, A., and Stefansson, S. O. 1999. Temperatureand size1242 dependent growth of larval and early juvenile Atlantic cod (Gadus morhua): a 1243 comparative study of Norwegian coastal cod and northeast Arctic cod. Can. J. Fish. 1244 Aquat. Sci. 56: 2099–2111. Page 52 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 53 1245 Palkovacs, E.P., Kinnison, M.T., Correa, C., Dalton, C.M., and Hendry, A.P. 2011. Fates 1246 beyond traits: ecological consequences of human-induced trait change. Evol. Appl. 5: 1247 183–191. doi:10.1111/j.1752-4571.2011.00212.x. 1248 Palkovacs, E.P., Marshall, M.C., Lamphere, B.A., Lynch, B.R., Weese, D.J., Fraser, D.F., 1249 Reznick, D.N., Pringle, C.M., and Kinnison, M.T. 2009. Experimental evaluation of 1250 evolution and coevolution as agents of ecosystem change in Trinidadian streams. Philos. 1251 Trans. R. Soc. B Biol. Sci. 364: 1617–1628. doi:10.1098/rstb.2009.0016. 1252 Palkovacs, E.P., and Post, D.M. 2009. Experimental evidence that phenotypic divergence in 1253 predators drives community divergence in prey. Ecology 90(2): 300–305. 1254 doi:10.1890/08-1673.1. 1255 Palstra, F.P., and Ruzzante, D.E. 2008. Genetic estimates of contemporary effective 1256 population size: What can they tell us about the importance of genetic stochasticity for 1257 wild population persistence? Mol. Ecol. 17(15): 3428–3447. doi:10.1111/j.13651258 294X.2008.03842.x. 1259 Parmesan, C., Root, T.L., and Willig, M.R. 2000. Impacts of Etreme Weather and Climate on 1260 Terrestrial Biota. Bull. Am. Meteorol. Soc. 81(3): 443–450. 1261 doi:http://dx.doi.org/10.1108/17506200710779521. 1262 Pauli, B.D., Kolding, J., Jeyakanth, G., and Heino, M. 2017. Effects of ambient oxygen and 1263 size-selective mortality on growth and maturation in guppies. Conserv. Physiol. 5: 1–13. 1264 doi:10.1093/conphys/cox010. 1265 Pauly, D. 1981. The relationships between gill surface area and growth performance in fish: a 1266 generalization of von Bertalanffy’s theory of growth. Meeresforschung 28: 251–282. 1267 Pauly, D., and Cheung, W.W.L. 2018. Sound physiological knowledge and principles in Page 53 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 54 1268 modeling shrinking of fishes under climate change. Glob. Chang. Biol. 24(1): e15–e26. 1269 doi:10.1111/gcb.13831. 1270 Pedersen, L. 1987. Energy budgets for juvenile Rainbow Trout at various oxygen 1271 concentrations. Aquaculture 62: 289–298. doi:10.1016/0044-8486(87)90171-2 1272 Persson, L., Andersson, J., Wahlström, E., and Eklöv, P. 1995. Size-specific interactions in 1273 lake systems: predator gape limitation and prey growth rate and mortality. Ecology 1274 77(3): 900–911. doi:10.2307/2265510. 1275 Pinsky, M.L., and Palumbi, S.R. 2014. Meta-analysis reveals lower genetic diversity in 1276 overfished populations. Mol. Ecol. 23(1): 29–39. doi:10.1111/mec.12509. 1277 Pörtner, H.O. 2002. Climate variations and the physiological basis of temperature dependent 1278 biogeography: systemic to molecular hierarchy of thermal tolerance in animals. Comp. 1279 Biochem. Physiol. Part A 132: 739–761. doi:10.1016/S1095-6433(02)00045-4. 1280 Pörtner, H.O., Bock, C., Knust, R., Lannig, G., Lucassen, M., Mark, F.C., and Sartoris, F.J. 1281 2008. Cod and climate in a latitudinal cline: Physiological analyses of climate effects in 1282 marine fishes. Clim. Res. 37(2–3): 253–270. doi:10.3354/cr00766. 1283 Pörtner, H.O., and Knust, R. 2007. Climate change affects marine fishes through the oxygen 1284 limitation of thermal tolerance. Science. 315: 95–98. doi:10.1126/science.1135471. 1285 Pörtner, H.O., and Peck, M.A. 2010. Climate change effects on fishes and fisheries: Towards 1286 a cause-and-effect understanding. J. Fish Biol. 77(8): 1745–1779. doi:10.1111/j.10951287 8649.2010.02783.x. 1288 Rabalais, N.N., Díaz, R.J., Levin, L.A., Turner, R.E., Gilbert, D., and Zhang, J. 2010. 1289 Dynamics and distribution of natural and human-caused hypoxia. Biogeosciences 7: 1290 585–619. doi:10.5194/bg-7-585-2010. Page 54 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 55 1291 Rahel, F.J., and Olden, J.D. 2008. Assessing the effects of climate change on aquatic invasive 1292 species. Conserv. Biol. 22(3): 521–533. doi:10.1111/j.1523-1739.2008.00950.x. 1293 Ratner, S., and Lande, R. 2001. Demographic and evolutionary responses to selective 1294 harvesting in populations with discrete generations. Ecology 82(11): 3093–3104. 1295 doi:10.1890/0012-9658(2001)082[3093:DAERTS]2.0.CO;2. 1296 Réale, D., Garant, D., Humphries, M.M., Bergeron, P., Careau, V., and Montiglio, P.O. 2010. 1297 Personality and the emergence of the pace-of-life syndrome concept at the population 1298 level. Philos. Trans. R. Soc. B Biol. Sci. 365(1560): 4051–4063. 1299 doi:10.1098/rstb.2010.0208. 1300 Reed, D.H., and Frankham, R. 2003. Correlation between fitness and genetic diversity. 1301 Conserv. Biol. 17(1): 230–237. doi:10.1046/j.1523-1739.2003.01236.x. 1302 Reusch, T.B.H., Ehlers, A., Hämmerli, A., and Worm, B. 2008. Ecosystem recovery after 1303 climatic extremes enhanced by genotypic diversity. Proc. Natl. Acad. Sci. 102(8): 2826– 1304 2831. doi:10.1073/pnas.091092498. 1305 Reznick, Butler, and Rodd. 2001. Life-History Evolution in Guppies. VII. The Comparative 1306 Ecology of Highand Low-Predation Environments. Am. Nat. 157(2): 126. 1307 doi:10.2307/3079254. 1308 Reznick, D., Bryant, M., and Holmes, D. 2006. The evolution of senescence and post1309 reproductive lifespan in guppies (Poecilia reticulata). PLoS Biol. 4(1): 0136–0143. 1310 doi:10.1371/journal.pbio.0040007. 1311 Reznick, D.A., Bryga, H., and Endler, J. 1990. Experimentally induced life-history evolution 1312 in a natural population. Nature. 346: 357 – 359. doi:10.1038/346357a0. 1313 Reznick, D.N., Shaw, F.H., Rodd, F.H., and Shaw, R.G. 1997. Evaluation of the rate of Page 55 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 62 1451 1452 1453 Figure 1. A summary of how different external stressors may affect fish body size, and how 1454 the change in body size or associated life-history traits may propagate through different levels 1455 of biological organisation. The superscript denotes an example study on the topic. 1) Hunter 1456 et al. 2015; 2) Nusslé et al. 2009, 2011; 3) Conover and Munch 2002; 4) van Wijk et al. 2013; 1457 5) Uusi-Heikkilä et al. 2015; 6) Uusi-Heikkilä et al. 2017; 7) Walsh et al. 2006; 8) Salinas et 1458 al. 2012; 9) Biro et al. 2004, Biro and Post 2008; 10) Uusi-Heikkilä et al. 2008; 11) Uusi1459 Heikkilä et al. 2015; 12) Barneche et al. 2018; 13) Dick et al. 2017; 14) Savage et al. 2004; 1460 15) Fleming and Gross 1994; 16) Berkeley et al. 2004; 17) Hixon et al. 2014; 18) 1461 Audzijonyte and Kuparinen 2016; 19) Dunlop et al. 2007; 20) Hauser et al. 2002; 21) Pinsky 1462 and Palumbi 2014; 22) Hutchinson et al. 2003; 23) Jakobsdóttir et al. 2011; 24) Olsen et al. 1463 2009; 25) Reznick et al. 1997; 26) Edeline et al. 2007; 27) Olsen et al. 2004; 28) Frank et al. 1464 2005; 29) Ohlberger et al. 2017; 30) Shackell et al. 2009; 31) Daufresne et al. 2009; 32) 1465 Kuparinen et al. 2016; 33) Becks et al. 2012; 34) Elmer and Meyer 2011; 35) Heins et al. 1466 2016; 36) Reznick et al. 2001; 37) Persson et al 1995; 38) Nilsson et al. 2019. 1467 1468 Figure 2. 1469 A summary of the main ecological theories behind the discussed stressors as selection agents. 1470 1) Bergmann 1847; 2) James1970; 3) Atkinson 1994; 4) West et al. 1997; 5) Pauly 1981; 6) 1471 Lefevre et al. 2017; Audzijonyte et al 2019 7) Lush 1937; 8) Miller et al. 1988; 9) MacArthur 1472 and Pianka 1966. 1473 Page 62 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft 63 1474 1475 1476 1477 1478 Page 63 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft A summary of how different external stressors may affect fish body size, and how the change in body size or associated life-history traits may propagate through different levels of biological organisation. The superscript denotes an example study on the topic. 1) Hunter et al. 2015; 2) Nusslé et al. 2009, 2011; 3) Conover and Munch 2002; 4) van Wijk et al. 2013; 5) Uusi-Heikkilä et al. 2015; 6) Uusi-Heikkilä et al. 2017; 7) Walsh et al. 2006; 8) Salinas et al. 2012; 9) Biro et al. 2004, Biro and Post 2008; 10) UusiHeikkilä et al. 2008; 11) Uusi-Heikkilä et al. 2015; 12) Barneche et al. 2018; 13) Dick et al. 2017; 14) Savage et al. 2004; 15) Fleming and Gross 1994; 16) Berkeley et al. 2004; 17) Hixon et al. 2014; 18) Audzijonyte and Kuparinen 2016; 19) Dunlop et al. 2007; 20) Hauser et al. 2002; 21) Pinsky and Palumbi 2014; 22) Hutchinson et al. 2003; 23) Jakobsdóttir et al. 2011; 24) Olsen et al. 2009; 25) Reznick et al. 1997; 26) Edeline et al. 2007; 27) Olsen et al. 2004; 28) Frank et al. 2005; 29) Ohlberger et al. 2017; 30) Shackell et al. 2009; 31) Daufresne et al. 2009; 32) Kuparinen et al. 2016; 33) Becks et al. 2012; 34) Elmer and Meyer 2011; 35) Heins et al. 2016; 36) Reznick et al. 2001; 37) Persson et al 1995; 38) Nilsson et al. 2019. 338x190mm (96 x 96 DPI) Page 64 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews Draft A summary of the main ecological theories behind the discussed stressors as selection agents. 1) Bergmann 1847; 2) James1970; 3) Atkinson 1994; 4) West et al. 1997; 5) Pauly 1981; 6) Lefevre et al. 2017; Audzijonyte et al 2019 7) Lush 1937; 8) Miller et al. 1988; 9) MacArthur and Pianka 1966. 338x190mm (96 x 96 DPI) Page 65 of 65 https://mc06.manuscriptcentral.com/er-pubs Environmental Reviews