The Impact of Catch-and-Release Management Practices on Fish

John B. Hume
+44 (0)7745 550 116 lampreydoctor@gmail.com http://lampreydoctor.wix.com/jbhumebio Twitter @thatlampreyguy

A report commissioned for the IBIS Knowledge Transfer Programme 2013

A project supported by the European Union's INTERREG IVA Programme managed by the Special EU Programmes Body

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The impact of catch-and-release management practices on fish

INTRODUCTION..1 METHODS.4 LETHAL EFFECTS OF CATCH-AND-RELEASE MANAGEMENT PRACTICES.5 Total mortality5 Does body size matter?..............................................................................................................7 Anatomical hook location...7 The impact of hook type.9 Treble vs. single hooks...9 Small vs. large hook sizes10 Circle vs. J-hooks.11 Barbed vs. barbless hooks12 The influence of bait type.13 Can line-cutting reduce mortality rates of deeply hooked fish?..............................................14 The effect of air exposure & handling times16 The influence of capture depth.18 The impact of barotrauma20

Can venting mitigate the effects of barotrauma?.....................................................................21 SUB-LETHAL EFFECTS OF CATCH-AND-RELEASE MANAGEMENT

PRACTICES23 The impact of C&R management practices on physiology..23 The influence of water temperature.25 The impact on post-release growth..26 Retention vs. immediate release following capture..28 The impact of capture on post-release behaviour29 The effects of capture in nets31 Other impacts of C&R management practices.32 THE EFFECTS OF CATCH-AND-RELEASE MANAGEMENT PRACTICES ON IBIS PRIORITY-MANAGEMENT SPECIES.33 CASE STUDY - BROWN TROUT Salmo trutta33 CASE STUDY ATLANTIC SALMON Salmo salar...34 CASE STUDY NORTHERN PIKE Esox lucius..37 CASE STUDY SHARKS (Selachimorpha spp.)..39 CASE STUDY SKATES (Rajidae spp.)...42 REFERENCES.43

INTRODUCTION Wild populations of both freshwater and marine fishes are heavily exploited by recreational fisheries, usually considered to be those where fishing is conducted by individuals for sport or leisure, and sometimes also for personal consumption (Cooke & Cowx, 2006; Cowx et al., 2010). Globally, 220 to 700 million people are believed to participate in these fisheries (Arlinghaus et al., 2013) and global trends of both fisheries catches and participation are generally increasing (Cooke & Cowx, 2004; 2006). This pressure on fish stocks can be just as intense as commercial exploitation (Post et al., 2002), thus affecting populations in a similar manner and requiring dedicated research, policy and management initiatives (Cowx et al., 2010; Arlinghaus et al., 2013). To ensure these fisheries resources are sustainable, concerted actions must be adopted by both individuals and stakeholders (Arlinghaus et al., 2002), one example being the Code of Conduct for Sustainable Fisheries (CCRF) by the United Nations Food and Agricultural Organisation (FAO, 1995). Recently, the European Inland Fisheries Advisory Commission (EIFAC) and European Angling Alliance (EAA) highlighted the need for an international Code of Practice (CoP) for recreational fisheries (Arlinghaus et al., 2012), and this provides a general collection of best practice guidelines for the sustainable development and maintenance of those fisheries (EIFAC, 2008; Arlinghaus et al., 2010). In its simplest form, the capture and subsequent release (catch-and-release, hereafter C&R) of an individual fish is the most basic of applied outcomes from such CoP guidelines, and is now widespread in global recreational fisheries (Arlinghaus, 2007; Ferter et al., 2013) having been discussed in at least three symposia since 1977 (Cooke & Schramm, 2007). Although harvest regulations for both recreational and commercial fisheries require some form of C&R, a low post-release survival rate could render any regulations ineffective (Pollock & Pine, 2007; Ferter et al., 2013) and conflict can arise between those groups who practice voluntary C&R and those mandated by regulation (Arlinghaus, 2007; Goodyear,
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2007). Conflict can also arise between recreational fisheries and non-fishery related aquatic biodiversity conservation objectives (Cowx et al., 2010), including practices detrimental to the health of non-target fish species (Hewlett et al., 2009). The sub-lethal effects of C&R could be equally important when considering the longterm resilience of fish populations (Arlinghaus et al., 2007) particularly in the context of a heightened awareness of fish welfare issues in contemporary society (Huntingford et al., 2006; Arlinghaus et al., 2012). The definition adopted by the EIFAC CoP was good welfare means that an individual fish is in good health, with its biological systems functioning properly and with no impairment of fitness, thus it is relevant when, for example, a fish has been captured but released with an injury to the mouth. If recovery of the fish following the C&R event is rapid and objectively measurable states of fitness have not declined (e.g., fecundity, quality of gametes, growth) (Cooke & Suski, 2005), then the impact of C&R on the individuals welfare is negligible (Arlinghaus et al., 2010). The welfare of fish in a recreational fishery context is, however, a complex and frequently contentious issue and is reviewed elsewhere (Cooke & Sneddon, 2007; Arlinghaus et al., 2009; Diggles et al., 2011). To provide guidance to recreational fisheries the EIFAC CoP outlined recommendations for appropriate behaviour and techniques that would act to minimise negative impacts on fish welfare during C&R events and maximise post-release survival. For example, the duration of landing should be minimised (Meka & McCormick, 2005); air exposure should be minimised (Ferguson & Tufts, 1992); as should injury (Arlinghaus et al., 2008), all of which can be achieved through alterations to tackle and gear as well as angler behaviour. Angling should also be avoided during warm weather or the reproductive period (Cooke & Suski, 2005). If fish are being held for a period of time as opposed to immediate release, then any devices, such as keep-nets or live-wells, should provide sufficient space and high water quality throughout (Arlinghaus et al., 2010). Although these generalities have
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their limitations, they do, however, provide some level of protection to all species (Cooke & Suski, 2005). Ultimately, the success of any fisheries regulations, and in particular C&R management practices, depends on ensuring a high rate of post-release survival is achieved through reductions to both sub-lethal effects and mortality rates (Pelletier et al., 2007; Pollock & Pine, 2007). Mortality impacts the demographic characteristics of fish populations and, by extension, fisheries managers impact the population by altering those rates of mortality. Therefore, estimates of mortality as a result of C&R management practices can and should be integrated with existing fisheries management strategy (Cooke & Schramm, 2007; Pollock & Pine, 2007; Ferter et al., 2013). The effects of implementing C&R practices are not, however, immediately observable, as was the case with European grayling Thymallus thymallus in northern Sweden for example, which required a decade of no-take angling to see positive effects on population demographics (Nslund et al., 2005). Managers must, therefore, remain patient. Additionally, although the sub-lethal effects of C&R practices are generally of lesser concern to fisheries managers, academics would contend that sub-lethal effects, in particular alterations to a fishs physiology and behaviour, are more likely to provide mechanistic reasons underlying post-release mortality and should, therefore, draw greater attention from managers (Cooke & Schramm, 2007; Pelletier et al., 2007). In order to evaluate the potential effectiveness of C&R management practices to priority-management marine and freshwater fish species in the Integrated Aquatic Resources Management Between Ireland, Northern Ireland and Scotland (IBIS) study area, factors that influence post-release mortality and sub-lethal effects have been reviewed from the primary literature. This is followed by more specific accounts of the efficacy of C&R management practices for six priority fish species or groups in the IBIS study area: Atlantic salmon Salmo

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salar, brown and sea trout Salmo trutta, northern pike Esox lucius, skate (Rajidae) spp. and shark (Selachimorpha) spp.

METHODS ISI Web of Knowledge (Thompson Reuters) was used to identify peer-reviewed literature that examined the efficacy of catch-and-release (C&R) management practices and its applicability to IBIS priority-management marine and freshwater fish species. Irrelevant articles were removed by reading abstracts and full articles where necessary, and excluded if it was not evident that the article contained quantitative data on the short- and long-term conservation value and/or economic value of C&R management practices. A combination of search terms was used to maximise the number of relevant articles. Articles must have included in their title, abstract or keywords one of the following search terms: catch and release management AND fish, Salmo salar AND catch and release. Salmo trutta AND catch and release, Esox lucius AND catch and release, skate AND catch and release, shark AND catch and release. All articles resulting from this search were collated into a database for further consideration. The literature search was supplemented by a single-pass reading of each article in the database to find additional relevant cited literature. This literature search resulted in a total of 340 peer-reviewed articles, 46% (n = 157) of which were abstracts only.

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LETHAL EFFECTS OF CATCH-AND-RELEASE MANAGEMENT PRACTICES Total mortality Estimates of C&R-induced mortality can vary widely depending on a variety of intrinsic and extrinsic factors, and is both highly species- and fishery-specific (Cooke & Suski, 2005), making general conclusions undesirable and potentially misleading. However, knowledge of the number of fish that die as a result of C&R practices is essential for basic fisheries management (Cooke et al., 2006). A meta-analysis of 274 studies found average mortality as a result of C&R practices to be 18% and, although highly skewed, mortality distributions were remarkably similar for both marine and freshwater taxa (Bartholomew & Bohnsack, 2005). Within this study, and ratified elsewhere (Cooke et al., 2006), seven mortality factors were found to be significant: anatomical hook location, bait type, removal of deep-set hooks (i.e., from the gut, oesophagus), J-hooks, capture depth, water temperature and handling times. In some fisheries release rates can be as high as 100% (e.g., course fisheries in western Europe) and, globally, approximately 60% of fish captured by recreational anglers are released (Cooke & Cowx, 2006). However, an unknown proportion of these subsequently die post-release (Cook et al., 2002). The following table presents the short-term (< 48 hours post-release) total mortality rates for a range of freshwater and marine species that support active recreational fisheries, and which were captured using standard hook-and-line methodology. Mortality rates of fish species following a C&R event vary widely; with an average of 18% across freshwater and marine species.

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Species Acanthopagrus butcheri Centropomus undecimalis Centropristis striata Cynoscion nebulosus Cynoscion regalis Epinephalus morio Glaucosoma hebraicum Lutjanus campechanus Macquaria ambigua Macquaria novemaculeata Megalops atlanticus Micropterus dolomieu Micropterus punctulatus Micropterus salmoides

Common name black bream common snook black sea bass spotted seatrout weakfish red grouper west Australian dhufish red snapper golden perch Australian bass tarpon smallmouth bass spotted bass largemouth bass

Mortality Citation Rate (%) 15.5 Grixti et al. (2008) 2.1 4.7 4.6 - 11 2.6 20 51 20 - 49.1 0 0-6 3.7 0 8.9 8.5 1.42 - 38 Taylor et al. (2001) Bugley & Shepherd (1991) Murphy et al. (1995); Stunz & McKee (2006) Malchoff & Heins (1997) Burns et al. (2012) St John & Syers (2005) Render & Wilson (1994); Burns et al. (2012) Hall et al. (2010) Hall et al. (2009); Dowling et al. (2010) Edwards (1998) Jackson & Willis (1991); Hartley & Moring (1995); Dunmall et al. (2001) Muoneke (1992) Meals & Miranda (1994); Hartley & Moring (1995); Kwak & Henry (1995); Weathers & Newman (1997); Neal & Lopez-Clayton (2001); DeBoom et al. (2010) Millard et al. (2003) Schill (1996) Lyle et al. (2007) Fabrizio et al. (2008); Broadhurst et al. (2012) Muoneke (1992) Brobbel et al. (1996); Dempson et al. (2002) Boyd et al. (2010) Tsuboi et al. (2002) Persons & Hirsch (1994) Reeves & Bruesewitz (2007); Schramm et al. (2010) Vecchio & Wenner (2007) Veiga et al. (2011) OToole et al. (2010) Veiga et al. (2011)

Morone saxitilis Oncorhynchus mykiss Platycephalus bassensis Pomatomus saltatrix Pomoxis annularis Salmo salar Salmo trutta Salvelinus leucomaenis Salvelinus namaycush Sander vitreus

striped bass rainbow trout sand flathead bluefish white crappie Atlantic salmon brown trout white-spotted charr lake trout walleye

31 16 1-6 8 - 25 9.3 8 12 5 6.7 24 0 - 54

Sciaenops ocellatus red drum 2 Sparus aurata gilthead seabream 11.7 Sphyraena barracuda great barracuda 0 Spondyliosoma black seabream 2.8 cantharus Tetrapturus albidus white marlin 12.5 Horodysky & Graves (2005) Thunnus thynnus Atlantic bluefin tuna 1.7 Stokesbury et al. (2011) Table 1. Selection of studies in which mortality rate following a C&R event has been estimated. 7" "

Does body size matter? The size of an individual fish is not consistently correlated with its rate of mortality following a C&R event, as some studies have shown larger fish to suffer greater mortality than smaller individuals, or the opposite effect, but the majority of studies fail to find any effect of body size on mortality rates (reviewed in Bartholomew & Bohnsack, 2005; Alos et al., 2009). The size of an individual, however, may lend itself to greater mortality rates through certain intrinsic factors such as alterations in foraging strategy. For example, among Trachynotus ovatus there is a tendency for larger specimens to become deeply hooked, which may increase post-release mortality rates as a result of injury (Alos, 2009) and a similar trend is seen in captured black bream Acanthopagrus butcheri (Grixti et al., 2007). The body size of a fish can affect the likelihood of mortality following a C&R event; however, the cause of such an effect differs between species.

Anatomical hook location In general, fishes hooked in critical body areas (e.g., gut, oesophagus, gills) suffer higher mortality rates compared to those hooked in the mouth (Bartholomew & Bohnsack, 2005; Butcher et al., 2006; Alos, 2008; Alos et al., 2008; Grixti et al., 2008). Bleeding, often a sign of severe trauma is correlated with anatomical hook location (Domeier et al., 2003; Prince et al., 2007; Mapleston et al., 2008; Grixti et al., 2010). For example, among groupers, bleeding was more likely to occur when a specimen was hooked in the gills (66.7%) and gut (40%) compared with the mouth (5%) (Bacheler & Buckel, 2004). However, the extent of any such bleeding varies among species (Rudershausen et al., 2007). Capture-related injury is the primary source of mortality in black bass Micropterus spp. for example, and has been related

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to the level of angler experience, hook type and bait type (Siepker et al., 2007). Among derbio Trachynotus ovatus fisheries 85% of fish hooked in the oesophagus, gills or gut died within two hours of capture (Alos et al., 2008) and suffered greater mortality compared to those hooked in the mouth (Alos, 2009). A similar effect was seen in silver perch Bidyanus bidyanus (van der Walt et al., 2005), striped bass Morone saxitilis (Millard et al., 2003), lake trout Salvelinus namaycush (Persons & Hirsch, 1994), spotted seatrout Cynoscion nebulosus (Murphy et al., 1995), yellowfin bream Acanthopagrus australis (Broadhurst et al., 2005), red drum Sciaenops ocellatus (Flaherty et al., 2013), snapper Pagrus auratus (Grixti et al., 2010), sand flathead Platycephalus bassensis (Lyle et al., 2007) and walleye Sander vitreus (Reeves & Bruesewitz, 2007), with specimens hooked in the gut or oesophagus suffering greater mortality rates compared to those hooked in the mouth or gills. St John & Syers (2005) found that 70% of west Australian dhufish Glaucosoma hebraicum died following hooking in the gut, as did 69% of striped bass Morone saxitilis (Millard et al., 2003). In this latter species, a deeply-hooked fish was 12-times more likely to die as a result of capture compared to a shallow-hooked specimen, strikingly similar to red porgy Pagrus pagrus, which were 11-times more likely to survive if hooked in the lip compared to deep locations (Overton et al., 2008). Contrary to these findings, however, DeBoom et al. (2010) found that largemouth bass Micropterus salmoides exhibited similar levels of mortality when experimentally deep-hooked or hooked in the mouth, and Pope & Wilde (2010) also found similar results for this species in field trials. This suggests that some species are better able to tolerate injuries related to deep hooking. The likelihood of hooking a fish in a critical location varies depending on the species foraging strategy, gear type and angler experience. (Schill, 1996; Dunmall et al., 2001; Cooke et al., 2012). For example, sparids have a tendency to be hooked more frequently in the mouth or jaw (Veiga et al., 2011). In some species, such as the roman Chrysoblephus
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laticeps, larger individuals were found to be more susceptible to being hooked in the gut compared to smaller specimens (Gotz et al., 2007) and the probability of deep-hooking in black bream Acanthopagrus butcheri increased with fish length (Grixti et al., 2007). Generally, deeply hooked fish require greater handling times and this could contribute the greater mortality rates seen among such specimens (Diggles & Ernst, 1997; Grixti et al., 2010). In both annular Diplodis annularis and striped seabream Lithognathus mormyrus, 54% and 92% of deeply hooked cases were unhooked with difficulty and was the most important factor explaining mortality rates over a 4-5 hour observation period (Alos et al., 2009). The rate of removal of deeply-set hooks decreased significantly in larger specimens of black bream Acanthopagrus butcheri, with a 0.8 less chance of hook removal for every one centimetre increase in fish length (Grixti et al., 2008). The evidence strongly suggests that fish suffer increased mortality rates following a C&R event when hooks are set in anatomical locations other than the mouth.

The impact of hook type Treble vs. single hooks. DuBois et al. (1994) found that treble hooks caused fewer mortalities in northern pike Esox lucius than did single large hooks, and single hooks also caused significantly greater mortality rates (5.1%) in yellow stripey Lutjanus carponotatus and the wire-netting cod Epinephelus auoyanus compared to treble-hooked lures (0.4%) (Diggles & Ernst, 1997). However, no differences in mortality rates were found between these hook types in cutthroat trout Oncorhynchus clarkii or common snook Centropomus undecimalis populations (Bartholomew & Bohnsack, 2005), as well as several trout species (DuBois & Dubielzig,
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2004). Ayvazian et al. (2002), however, found that treble hooks caused significantly higher mortality of tailor Pomatamus saltatrix compared to four other hook types, including single hooks, and may have caused extensive lacerations and other physical trauma (OToole et al., 2010; Mandelman et al., 2012). Treble hooks could act to reduce mortality rates by being less likely to set in critical anatomical locations compared to single hooks, but as they generally take longer to remove compared to single hooks (OToole et al., 2010) this positive effect may be offset (Gjernes et al., 1993). Only four of 49 fishery agencies in North America recommend the use of single hooks over treble hooks (Pelletier et al., 2007) suggesting that, at least in that region, fishery managers believe treble hooks to be less injurious to fish. The evidence is that single and treble hooks result in different outcomes following a C&R event; with a reduced likelihood of mortality when using treble hooks as a result of shorter handling times.

Small vs. large hook sizes. In a general sense, larger hooks capture larger specimens (Alos et al., 2008). However, in some specialised fisheries, such as common carp Cyprinus carpio, small hooks tend to capture larger specimens (Rapp et al., 2008). Hook size can be a significant predictor of mortality rates following a C&R event. For example, small hooks (size 1/0) were found to account for 25% mortality in captured blue cod Parapercis colias compared to zero mortality when using large hooks (size 6/0) (Carbines, 1999), and smaller hooks (sizes 14 and 12) caused greater mortality of derbio Trachynotus ovatus compared to larger hooks (sizes 10, 8 and 6) (Alos et al., 2008a). Large hook sizes were, however, found to result in increased incidences of deeply hooked fish in some studies (Alos et al., 2008b; Mapleston et al., 2008; Rapp et al., 2008) but the opposite effect was seen in a mixed grouper fishery, where small
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hooks resulted in more deeply hooked specimens (2 = 8.99, d.f. = 3, P = 0.03) (Bacheler & Buckler, 2004). Black bream Acanthopagrus butcheri are 6.6-times more likely to be deeply hooked on small hooks compared to large hooks (Grixti et al., 2007). No effect of hook size on mortality was found in striped bass Morone saxitilis or cutthroat trout Oncorhynchus clarkii (Bartholomew & Bohnsack, 2005), suggesting that the effect of hook size on mortality rates is species specific. The evidence suggests that hook sizes can affect mortality rates following a C&R event; however, the effect varies between species.

Circle vs. J-hooks. Circle hooks are more likely to become set in the corner of fishes mouths during capture thus reducing handling time, while J-hooks are more likely to be swallowed and become set in the gut or oesophagus (Domeier et al., 2003; Cooke & Suski, 2004; Meka, 2004; Bartholomew & Bohnsack, 2005; Beckwith & Rand, 2005; Prince et al., 2007). For example, there was a significant correlation (2 = 119.71, d.f. = 1, P < 0.001) between the anatomical hooking location and unhooking time of painted comber Serranus scriba caught using J-hooks (Alos, 2008). In a mixed grouper fishery gut hooking was significantly higher when using J-hooks compared to size 12/0 circle hooks (2 = 21.78, d.f. = 3, P < 0.001) (Bacheler & Buckler, 2004). However, in North America only 5 of 49 surveyed fisheries agencies recommended the use of circle hooks, indicating a lack of adherence with current scientific opinion in this region (Pelletier et al., 2007), perhaps as a result of the perceived reduced hooking and landing efficiencies of circle hooks (Cooke et al., 2003), and that not all studies agree hook type affects anatomical hooking location (Alos, 2009) or hook retention (Jones, 2005; van der Walt et al., 2005).
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The proportion of Atlantic Istiophorus albicans and Pacific sailfish I. platypterus hooked in the mouth using circle hooks was 85%, compared to 25% with J-hooks, and resulted in deeply hooked fish on only 2% of occasions compared to 46% when using Jhooks (Prince et al. 2002) leading to fewer injuries when the fishery employed circle hooks. Circle hooks also more frequently penetrated the lip region of white seabass Atractoscion nobilis (73%) compared to J-hooks (41%) (Aalbers et al., 2004). There were fewer mortalities among bluefin tuna Thunnus thynnus (Skomal et al., 2002), sand flathead Platycephalus bassensis (Lyle et al., 2007), seabream Diplodis annularis (Alos et al., 2009), red drum Sciaenops ocellatus (Vechio & Wenner, 2007), striped bass Morone saxitilis (Millard et al., 2005) and Chinook salmon Oncorhynchus tshawytscha (Grover et al., 2002) when caught using circle hooks compared to J-hooks, and striped bass Morone saxitilis suffer greater mortality when caught using J-hooks as a result of frequent deep hooking (Bartholomew & Bohnsack, 2005). Hooking injuries as a result of deeply-set J-hooks can cause delayed mortality in those specimens with trauma such as severed oesophagi (Burns et al., 2012; Flaherty et al., 2013). However, no effect of hook type on mortality was seen in white seabass Atractoscion nobilis (Aalbers et al., 2004), red drum Sciaenops ocellatus (Flaherty et al., 2013), yellowfin tuna Thunnus albacares or summer flounder Paralichthys dentata populations captured, although circle hooks did generally result in fewer critical anatomical hooking locations (reviewed in Bartholomew & Bohnsack, 2005). Large circle hooks, but not small ones, have been found to reduce the incidence of deep-hooking relative to using J-hooks in red drum Sciaenops ocellatus fisheries (Beckwith & Rand, 2005). Even among pelagic longline fisheries the odds of post-release survival for many target and bycatch species were found to be significantly higher if circle hooks were employed (Horodysky & Graves, 2005;

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Carruthers et al., 2009; Graves & Horodysky, 2010; Serafy et al., 2012) and this holds true for a variety of circle hook models (Graves & Horodysky, 2008). Despite variation among studies due to factors such as hook size, fishing style and feeding mode of the target species (Cooke et al., 2012; Walter et al., 2012), the evidence is that mortality rates appear to be consistently lower when circles hooks are used compared to J-hooks (Cooke & Suski, 2004; Wilson & Diaz, 2012).

Barbed vs. barbless hooks. Barbless hooks are easier to remove when compared to barbed hooks (DuBois & Dubielzig, 2004; Meka, 2004), significantly reducing air exposure for some species such as rock bass Ambloplites rupestris (Cooke et al., 2001; Pelletier et al., 2007) and yellow stripey Lutjanus carponotatus (Diggles & Ernst, 1999). Barbed hooks are, therefore, believed by some authors to be the most significant determinants of injury and/or mortality following a C&R event (reviewed in Muoneke & Childress, 1994). However, Cooke et al. (2001) found no differences in mortality rates of rock bass Ambloplites rupestris captured on barbed vs. barbless jigs or barbed vs. barbless worms and no delayed mortality was observed after five days. Neither were there significant differences in the mortality rates of several trout species captured on single barbless hooks compared with barbed singles or trebles (DuBois & Dubielzig, 2004) or in bluegills Lepomis macrochirus experimentally hooked in the oesophagus (Robert et al., 2012). Barbed hooks do, however, in general cause higher rates of injury compared to barbless hooks, for example, in rainbow trout Oncorhynchus mykiss fisheries (Meka, 2004). This was not found to be the case though for walleye Sander vitreus, which exhibited similar

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levels of hook-related damage when captured using barbed vs. barbless live-bait hooks (Reeves & Staples, 2011). In a meta-analysis by Schill & Scarpella (1998) it was concluded that the use of barbless hooks played no role in reducing post-release mortality rates among salmonids, and that barbed hook restrictions appear to be more of a social issue among anglers and managers rather than based on scientific evidence. There is little empirical evidence supporting the widely held belief that barbless hooks result in a reduced likelihood of mortality following a C&R event for most fish species.

The influence of bait type Specific studies relating to effect of bait type on anatomical hook location are not common, despite the fact that natural baits are widely believed to lead to greater incidences of deep hooking as fish are more likely to ingest those compared with artificial baits or lures (Schill, 1996; Diggles & Ernst, 1997; Nelson, 1998; Cooke et al., 2001; Dunmall et al., 2001; Stunz & McKee, 2006), of which they may be more wary (Diggles & Ernst, 1997). Natural baits have been found to lead to greater mortality rates in some fish species when compared with artificial lures or flies, such as cutthroat trout Oncorhynchus clarkii and rainbow trout Oncorhynchus mykiss, but not in others such as striped bass Morone saxitilis, common snook Centropomus undecimalis (Muoneke & Childress, 1994; Bartholomew & Bohnsack, 2005) or black bass Micropterus spp. (Siepker et al., 2007). Using beach worms was found to result in a significant increase in the probability of deep hooking among sand whiting Sillago cilliata, therefore resulting in greater rates of mortality (Butcher et al., 2006). In a mixed-species marine recreational fishery bait type was found to have a significant effect on hooking location, with 26.9% (n = 537) of fish captured

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using worm bait being deeply hooked, whereas 14.1% (n = 214) of fish captured using shrimp as bait were deeply hooked (Alos et al., 2009). Rock bass Ambloplites rupestris were also found more likely to be deeply hooked when captured using worms compared to a lure (Cooke et al., 2001). The probability of mortality in Australian bass Macquaria novemaculeata as a result of capture using natural bait was found to be 12-times higher than for those captured using artificial lures (Hall et al., 2009). The evidence suggests that, in general, natural baits have the potential to result in a greater likelihood of mortality following a C&R event compared with artificial lures.

Can line-cutting reduce mortality rates of deeply hooked fish? Cutting the line on a deeply-set hook is likely to significantly improve survivorship of a fish following capture, as attempts to retrieve the hook could cause additional internal injuries (Bartholomew & Bohnsack, 2005; Grixti et al., 2008; Grixti et al., 2010). In North America 90% (n = 44) of surveyed fishery agencies recommended cutting the line when a fish was deeply hooked (Pelletier et al., 2007). Deeply-hooked, experimentally angled Trachynotus ovatus that had the line cut experienced a 50% reduction in post-release mortality, although this was not a statistically significant effect (Alos, 2009), and a similar non-significant effect was seen among deeply-hooked silver perch Bidyanus bidyanus (van der Walt et al., 2005). In black bream Acanthopagrus butcheri cutting the line on deeply-hooked fish resulted in a 92.7% (n = 105) delayed (1-72 hours) survival rate, compared to 69.5% (n = 51) in those where the hook was removed, despite almost identical initial (< 1 hour) survival rates (84.7% vs. 83.6%) (Grixti et al., 2008). In addition, there was little evidence found among 27 recreationally-caught Australian species that cutting the line negatively impacted recapture rates (Wilde & Sawynok, 2009) and, although survival rates were not found to be
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significantly affected by line-cutting in deeply hooked walleye Sander vitreus (Reeves & Bruesewitz, 2007), most studies agree line cutting has a positive effect on survival rates following C&R events. In deeply hooked snapper Pagrus auratus 13% of 59 specimens were able to shed the hook in less than three days (Grixti et al., 2010), fewer than the evacuation rate seen in deeply-hooked white-spotted charr Salvelinus leucomeinis (32.9%, n = 82 hooks in 72 fish) (Tsuboi et al., 2006) and masu salmon Oncorhynchus masou masou (70-100% in 21 days) (Doi et al., 2005). Yellowfin bream Acanthopagrus australis were also found to be able to shed ingested nickel-plated J-hooks in 5-56 days and had oxidised the hook to c. 94% of the original weight and often broken into two pieces (Broadhurst et al., 2007). These results suggest that, for some species, line-cutting for those specimens deeplyhooked is a viable management practice that reduces the likelihood of mortality following a C&R event.

The effect of air exposure & handling times Several recent studies have concluded that air exposure is a significant factor affecting survival rates and stress condition in fish experiencing a C&R event (Bartholomew & Bohnsack, 2005; Cooke & Suski, 2005; Arlinghaus et al., 2007; Gingerich et al., 2007; Wedemeyer & Wydoski, 2008). Removing a fish from water following capture is stressful as it deprives them of oxygen at a critical point when they must recover following exertion, and this can lead to extended recovery times (Cooke et al., 2001) and, in some cases, permanent tissue damage (Cooke & Suski et al., 2005). Most surveyed fishery agencies in North America (84%, n = 41) strongly emphasise that fish should be brought in as quickly as

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practical to avoid unnecessary struggling of the fish (Pelletier et al., 2007). The cardiovascular system of fish is heavily impacted during air exposure and the duration of a C&R event generally correlates positively with the magnitude of physiological disturbance and subsequent recovery times (Cooke & Suski, 2005). For example, when rock bass Ambloplites rupestris where exposed for either 30 or 180 seconds their cardiac output, stroke volume and heart rate required significantly longer to return to base levels following longer durations of air exposure (Cooke et al., 2001). A strong correlation between the time taken for cardiovascular variables to return to normal and the duration of air exposure was also noted in smallmouth bass Micropterus dolomieu (Cooke et al., 2002). Mortality rates tend to be increased by exposing fish to air following capture. For example, rainbow trout Oncorhynchus mykiss exposed to air following 60 seconds of exhaustive exercise had a mortality rate of 72%, compared to 12% mortality of individuals exhaustively exercised but not exposed (Ferguson & Tufts, 1992). Longer air exposure, and greater overall handling times were significant predictors of the loss of equilibrium in bonefish Albula vulpes but were not directly related to the risk of subsequent predation (Danylchuk et al., 2007). Bonefish exhibited significant increases in plasma glucose, haematocrit and blood lactate when exercised experimentally for four minutes compared to resting values, and can take up to 10-times longer to regain equilibrium than individuals exposed to air for just one minute (Suski et al., 2007). In bluegill Lepomis macrochirus fish exposed for 960 seconds during experimental treatments suffered higher levels of mortality, and did so faster, than those exposed for 480 and 240 second treatments, indicating a linear relationship between the duration of air exposure and mortality rates (Gingerich et al., 2007). Largemouth bass Micropterus salmoides also exhibited a positive correlation between the duration of air exposure, length of time to regain equilibrium and increased glucose concentrations, suggesting C&R events could have had a significant impact on their post18" "

release behaviour (White et al., 2008). Among nesting black bass Micropterus spp. exposed to simulated tournament angling conditions, which include extended handling times when they are weighed, there is a significant trend towards the abandonment of broods as a result of increasing the duration of treatment times (Hanson et al., 2007). A similar effect on brood abandonment was also seen in the congeneric species Micropterus dolomieu exposed to simulated tournament conditions (Hanson et al., 2008). The magnitude of physiological disturbance on tournament-caught walleye Sander vitreus was also found to be strongly positively correlated with the duration of air exposure during the weigh-in process (Killen et al., 2006). Brook trout Salvelinus fontinalis exposed for 120 seconds or more suffered from severe reductions in swimming performance (equivalent to 75%) (Schreer et al., 2005). However, handling time was not found to be a significant predictor of post-release survival in coho salmon Oncorhynchus kisutch captured using seine nets, neither did it appear to affect subsequent migration rates following release (Raby et al., 2012). In a perhaps extreme case air exposure for periods up to five minutes had no significant effect on survival rates of silver perch Bidyanus bidyanus (van der Walt et al., 2005), suggesting a species-specific tolerance for air exposure. There is strong evidence to suggest that a period of air exposure, and exacerbated by periods of exercise prior to landing the fish, increases the likelihood of mortality during a C&R event. However, there are significant differences between species.

The influence of capture depth In conjunction with anatomical hook location Bartholomew & Bohnsack (2005) considered capture depth to be the most important factor influencing the mortality rates of fish released following a C&R event. In general, increased depth of capture results in a variety of injurious
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conditions as a result of barotrauma, which can subsequently increase the mortality rate (Diggles & Ernst, 1997; Alos, 2008). In painted comber Serranus scriba mortality rates increased as capture depth increased, and fish captured deeper than 16 m had a 50% chance of experiencing some form of barotrauma (Alos, 2008). Only 2% of sauger Sander canadensis captured at depths of less than 9 m suffered mortality, compared to 33% of those captured at depths of 9-24 m (Meerbeek & Hoxmeier, 2011). Mortality rates among west Australian dhufish Glaucosoma hebraicum increased from 21% at capture depths of 0-14 m to 86% at depths greater than 45 m (St John & Syers, 2005). Similarly, more than 60% of gag Mycteroperca microlepis captured at 36 m depth or greater, and 95% of red grouper Epinephelus morio captured at depths greater than 41 m, exhibited distended stomachs (Bacheler & Buckel, 2004). The survival rates of red grouper captured at depths greater than 44 m was less than 33% (Wilson & Burns, 1996). Red snapper Pagrus auratus exhibited very little mortality when captured at depths of less than 30 m (0-2%), but this increased rapidly to 39% mortality at depths of 30-44 m and 55% between 45 and 59 m (Stewart, 2008). This indicates a general positive relationship between capture depth and the incidence of barotrauma and associated mortality. However, annular seabream Diplodus annularis captured at depths of down to 42 m exhibited no correlation between mortality and capture depth (Alos et al., 2009) and a linear relationship between capture depth and signs of barotrauma was not noted in experimentally captured red emperor Lutjanus sebae (Brown et al., 2010). Survival rates of yelloweye rockfish Sebastes ruberrimus were not significantly influenced by capture depths ranging 1872 m (Hochhalter & Reed, 2011). The evidence suggests that even among those groups susceptible to barotrauma from capture at great depths there is much inter-individual variation, generating conflicting results between studies (Rudershausen et al., 2007).
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The impact of barotrauma Barotrauma, which results from a rapid ascension from depth, can include a variety of conditions that negatively impact the mortality rates of fish following a C&R event (Rummer & Bennett, 2005; Schreer et al., 2009), and includes the formation of gas bubbles in tissue that can result in clotting of the blood, embolisms and haemorrhaging (Alos, 2008; Butcher et al., 2012) as well as a range of other physiological alterations (Morrissey et al., 2005). Various injuries can also result from the expansion of the gas bladder, including protrusion of the intestines through the anus and the stomach through the mouth, or protrusion of eyes (exophthalmia) (Brown et al., 2010; Butcher et al., 2012). In extreme cases of exophthalmia mean ( S. E.) eye displacement can be as much as 60.9 5.1% of total body length (Rogers et al., 2011). Gas bladder expansion can also render fish positively buoyant, preventing them from returning to depth and exposing them to predation (Shasteen & Sheehan, 1997; Bartholomew & Bohnsack, 2005; St John & Syers, 2005; Alos, 2008; Campbell et al., 2010). Additionally, ripe females experiencing expanded gas bladders may release their ova, therefore compromising their future reproductive potential (Roach et al., 2011). The susceptibility of species to these effects is a function of their anatomy. Some fishes have a pneumatic duct connecting the gas bladder and digestive tract (physostomous species) and these species can regulate the volume of their gas bladder rapidly compared to physoclistous species, which lack this duct (Bartholomew & Bohnsack, 2005; Pelletier et al., 2007). In snapper Pagrus auratus clinical signs of barotrauma become apparent at capture depths of 11 m and 19% (n = 315) of specimens captured at depths of down to 60 m exhibited a distended coelomic cavity, with none showing signs of subcutaneous gas bubbles or exophthalmia (Butcher et al., 2012). The presence of a ruptured gas bladder (84.4%) was the only clinical sign positively associated with capture depth in this study. Among experimentally angled red snapper Lutjanus campechanus captured at depths of 30-50 m 53%
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(n = 63) of individuals exhibited external signs of barotrauma (42% stomach eversion; 12% exophthalmia; 2.5% gut expulsion) (Diamond & Campbell, 2009). In the west Australian dhufish Glaucosoma hebraicum injuries resulting from barotrauma (bubbles in the eyes, exophthalmia, everted stomach and enlarged gas bladder) increased in both frequency and severity with increasing capture depth (St John & Syers, 2005). The effects of barotrauma are highly species-specific as evidenced by studies of mixed fisheries capturing a range of species at the same depths under the same environmental conditions with similar gear types (e.g., Rudershausen et al., 2007).

Can venting mitigate the effects of barotrauma? Venting relieves internal pressure by puncturing the expanded gas bladder of a positively buoyant fish with a hypodermic needle, preventing that fish from floating once released - a condition that could render them susceptible to predatory attack or increasing stress (Siepker et al., 2007). In North America the techniques to do so, and advice provided, by fishery agencies is inconsistent and this could reflect the uncertainty of the usefulness of this technique in the scientific literature (Pelletier et al., 2007; Brown et al., 2010). In painted comber Serranus scriba that experienced excess gas as a result of rapid decompression mortality rates were halved following venting, although this effect was not statistically significant (Alos, 2008). Snapper Pagrus auratus were able to immediately regain control of their buoyancy following venting, and were able to swim faster than untreated fish (Butcher et al., 2012) and mortality rates among west Australian dhufish Glaucosoma hebraicum did not appear to be negatively affected by venting (St John & Syers, 2005). Smallmouth bass Micropterus dolomieu that experienced barotrauma and were subjected to venting did not suffer any mortality, and their subsequent mean daily movement following treatment was not
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affected (Nguyen et al., 2009). There is some indication that the Lutjanus spp. coral reef fishery could benefit from venting as the recapture rate of some species appeared to be positively affected by the technique (Sumpton et al., 2008). Venting yellow perch Perca flavescens was found to have a significant positive effect on survival rates for up to three days post-release, and mark-recapture studies indicated long-term survival was not impacted (Kinery et al., 1996). However, in a meta-analysis of studies (n = 17) investigating the effects of venting on fish following a C&R event Wilde (2009) concluded that the technique may in fact be detrimental as a conservation measure for both marine and freshwater fisheries as it appears harmful to fishes captured from deep waters. In an experimental study of the effects of barotrauma on Australian bass Macquaria novemaculeata the only mortalities recorded (n = 4) were due to venting, most likely caused by severe internal injury (Roach et al., 2011). Bruesewitz & Coble (1993) found no evidence that the mortality rates of burbot Lota lota that exhibited expanded gas bladders were positively affected by venting, and a similar non-effect was noted in largemouth bass Micropterus salmoides (Lyle, 1992) and red snapper Lutjanus campechanus (Render & Wilson, 1994). The evidence is that despite mitigating against the effects of barotrauma, venting may also cause significant harm to fish and is unlikely to reduce the likelihood of mortality following a C&R event.

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SUB-LETHAL EFFECTS OF CATCH-AND-RELEASE MANAGEMENT PRACTICES The impact of C&R management practices on physiology During a C&R event a fish will undergo a combination of aerobic and anaerobic exercise that result in physiological changes, such as depletion of energy stores, accumulation of lactate and cardiac and osmoregulatory disturbance (Cooke et al., 2001; Cooke & Suski, 2005; Siepker et al., 2007; Suski et al., 2007). All of these may have sub-lethal effects on postrelease survival rates as the fish may or may not recover (Cooke et al., 2004; Arlinghaus et al., 2007). However, as a result of the enormous variation in capture methodology, environmental variables, angler experience, etc. the physiological consequences of C&R practices for fish are extremely context dependent (reviewed in Cooke et al., 2013). Laboratory based studies of the sub-lethal effects of C&R, despite their commonality (Cooke et al., 2004), are not always directly relevant to what occurs in the wild and novel wild-based methodologies are better able to provide more meaningful measures of stress and the correlation of mortality rates with post-release physiology (Siepker et al., 2007; Cooke et al., 2013). For example, using telemetry (Cooke et al., 2002; reviewed in Donaldson, 2008) or blood sampled from fish immediately following capture in the wild is more likely to be representative of the natural stress response of an individual than is blood sampled from a fish artificially stressed in a laboratory (Skomal, 2007). In a study of experimentally angled largemouth bass Micropterus salmoides heart-rate telemetry was able to show that higher maximal heart rates were consistently seen in wild vs. laboratory-exercised fish, and that those rates were independent of water temperature and could persist for as long as 300 minutes post-release (Cooke et al., 2004). The duration of capture of largemouth bass Micropterus salmoides was positively correlated with increases in plasma glucose, chloride, osmolarity and lactate levels
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(Gustaveson et al., 1991; White et al., 2008) and similar effects were noted in captured Atlantic salmon Salmo salar (Thorstad et al., 2003), striped bass Morone saxitilis (Ridley et al., 1997), tigerfish Hydrocynus vittatus (Smit et al., 2009) and red drum Sciaenops ocellatus (Gallman et al., 1999). In smallmouth bass Micropterus dolomieu an increase in lactate and a decrease in energy stores was more severe for individuals that were angled for two minutes compared to those angled for 20 seconds (Kieffer et al., 1995) and was always higher in exercised fish compared to controls (Hanson et al., 2008). In Atlantic cod Gadus morhua capture methodology had a profound effect on an individuals physiology, with those caught on a longline exhibiting 13- to 14-times greater serum cortisol concentrations, and higher serum glucose and plasma lactate levels compared to those caught on a jig (Mandelman et al., 2012). However, some authors advise against collecting blood samples in the field due uncontrolled variables (Diamond & Campbell, 2009) and a lack of control samples (OToole et al., 2010). Holding fish in tanks or cages for observation following a C&R event may also add additional stresses and negatively impact on an individuals physiology. In sea bream Pagrus major captured using hook-and-line methodology the plasma cortisol levels (mean S.E.) of resting fish (12.21 g ml-1; n = 12) were significantly lower compared to those sampled from captured fish, and increased as capture duration was extended from 10 minutes to three hours (up to 104.1 34.8 g ml-1; n = 8) (Chopin et al., 1996). In common carp Cyprinus carpio retained in so-called carp sacks following simulated capture, plasma cortisol and plasma glucose concentrations were significantly higher than control fish, and retention also had a significant negative effect on plasma lactate, blood-pH and osmolality (Rapp et al., 2012). A similar effect was also noted in walleye Sander vitreus (Killen et al., 2006) and largemouth bass Micropterus salmoides (Suski et al., 2003) that were being weighed following tournament angling, with large increases in white muscle and plasma lactate concentrations
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and significant cardiac disturbance. Wild coho salmon Oncorhynchus kisutch captured using beach seines also experienced significant changes in their plasma physiology as a direct function of handling time (Raby et al., 2012) as did several commercially caught pelagic fishes retained on longlines such as sharks, tunas and billfishes (reviewed in Skomal, 2007; Fabrizio et al., 2008). However, in the case of sand whiting Sillago cilliata captured using hook-and-line methodology and released into sea cages no significant differences were detected in the mean ( S.E.) plasma glucose concentrations between six wild (3.42 0.32 mmol l-1) and 22 experimental (4.13 0.23 mmol l-1) fish following seven days of holding (Butcher et al., 2006). Several salmonids were also found not to exhibit physiological disturbance as a result of retention in net-pens following a C&R event (Wedemeyer & Wydoski, 2008), suggesting some species can return to resting physiological rates more quickly than others. The evidence suggests that a C&R event significantly elevates stress hormone levels, cardiac output and disrupts osmoregulatory capacity in fish; all of which may negatively impact mortality rates. However, some species appear physiologically more tolerant of such disturbances.

The influence of water temperature At higher water temperatures there is less oxygen available, yet a fishs respiratory rate increases resulting in physiological stress during a C&R event (Muoneke & Childress, 1994; Cooke & Suski, 2005). This can lead to inflated mortality rates in warmer water conditions due to limitations in the maximal cardiovascular performance as fish approach their maximal metabolic rate and an exposure to greater oxygen debts following exercise (Cooke & Suski,

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2005). For example, Atlantic salmon Salmo salar exhibited low levels of mortality when captured at temperatures between 8C and 18C, but mortality rates associated with C&R events increased exponentially at temperatures higher than 18C (Thorstad et al., 2003). In a meta-analysis of 83 peer-reviewed articles 70% suggested that warm water conditions (relative to a particular species thermal tolerance) contributed to both increased indices of stress and subsequent mortality rates (Gale et al., 2013). In addition, higher water temperatures have been found to result in an increased prevalence of secondary infection following injury (Muoneke, 1992). However, approximately half of the studies examining recovery rates following C&R events indicate that warmer temperatures facilitate or expedite recovery time (Gale et al., 2013) and relative surface temperatures appear to have a negligible effect on post-release mortality (Alos, 2009). The evidence suggests a complex interaction between water temperatures and fish physiology that is highly context dependent; however, in general warm water temperatures appear to increase the likelihood of mortality following a C&R event.

The impact on post-release growth Normal feeding behaviours can be interrupted by C&R events, impairing the growth of an individual and potentially the reproductive capacity of that individual and its population (Cooke & Schramm, 2007; Siepker et al., 2007). But results of bioenergetics models appear inconsistent. A compromised feeding ability, caused by either injury, stress or a feeding delay following C&R, can negatively affect the growth of black bass Micropterus spp. (Siepker et al., 2006) and rainbow trout Oncorhynchus mykiss (Meka & Margraff, 2007). However, in experimental studies of rainbow trout, where individuals were hooked in the mouth, no change in growth was recorded (Pope et al., 2007). Neither were differences in growth rate
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observed in captured vs. un-caught: largemouth bass over a 40-day period (Pope & Wilde, 2004), white-spotted charr Salvelinus leucomaenis over a 50-day period (Tsuboi et al., 2001), white seabass Atractoscion nobilis over a 90-day period (Aalbers et al., 2004), or in largemouth bass over an 11-month period (DeBoom et al., 2010). Despite the prevalence of deep-hooking as a result of some hook-and-line methodologies in one particular study of yellowfin bream Acanthopagrus australis that had experimentally ingested a hook (n = 17) no significant differences were noted in their ability to digest and assimilate food compared to control fish (n = 20) (Broadhurst et al., 2007). No effect of air exposure was noted in the growth of zander Sander lucioperca exposed for 0 240 seconds following simulated angling (Arlinghaus & Hallerman, 2007), suggesting even the most severely injurious implications of C&R practices do not always result in compromised growth. In some species the number of times an individual fish has been captured, as well as water temperature, have both been implicated in the degree to which growth rate was reduced (Siepker et al., 2007). In a long-term (27 years) mark-recapture study of largemouth bass Micropterus salmoides 1050 individuals were tagged and released to monitor growth performance following C&R events (Cline et al., 2012). In this study some individuals were captured one to six-times each season (1-98 day intervals) and exhibited a post-release weight-loss period (up to six days post-release), but a subsequent compensatory growth period (six to 27 days post-release), resulting in negligible impact on growth patterns overall. The evidence suggests that multiple C&R events do not necessarily result in compromised long-term growth.

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Retention vs. immediate release following capture In recreational fisheries, catches are typically released at or close to the site of capture, allowing the fish to return to its original position. However, in competitive angling competitions, such as with North American black bass Micropterus spp., fish are retained in either a keep-net or live-well and transported to a central location to be weighed (Siepker et al., 2007). Following the weigh-in, competitors catches are usually released near to the weigh-in site, as opposed to the original capture location and this can further exacerbate fitness costs to the individual (Siepker et al., 2009). Despite recent improvements aimed at reducing the mortality of black bass retained during tournaments (e.g., continuous water flow in live-wells, aeration, early starts in warm weather, reduced catch limits) some events still exhibit mortality rates exceeding 50% of captured fish (Suski et al., 2006; Siepker et al., 2007). Mortality rates may also be increased when live-wells are employed as the result of an increased prevalence of largemouth bass virus (LMBV) (Siepker et al., 2007), which has been shown to transmit from infected to uninfected juvenile bass through shared water alone (Grant et al., 2005). A variety of other pathogenic bacteria have also been identified from tournament-caught black bass (Steeger et al., 1994). In largemouth bass Micropterus salmoides captured and retained during tournaments nest abandonment was far higher (90%) than those fish captured and immediately released (33%) (Diana et al., 2012) and the same effect on nest abandonment was seen in the congeneric Micropterus dolomieu (Hanson et al., 2008). Smallmouth bass also rapidly disperse following tournament capture, moving an average 1,475 m away from the release site (Kaintz & Bettoli, 2010). Among common carp Cyprinus carpio captured using hookand-line methodology and retained in so-called carp sacks, fish retained for less than 9 hours moved significantly shorter distances compared to control groups released immediately (Rapp et al., 2012).
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The evidence suggests that fish should be released as close to the point of capture as possible, and retained for as short a period of time as possible, to ensure they are best able to return to their original location.

The impact of capture on post-release behaviour The behaviour of fish following a C&R event could have a strong effect on any subsequent mortality or sub-lethal effects, especially if that behaviour is atypical. In a review of posttournament dispersal of captured black bass Micropterus spp. it was noted that smallmouth bass Micropterus dolomieu exhibited greater dispersal distances (mean 7.3 km) and returned more frequently to their capture location (mean 37%) than did largemouth bass Micropterus salmoides (mean dispersal distance = 3.5 km; 14% return rate) (Wilde, 2003). Dispersal rates for these species did not differ between fishing locations, and across all studies fish took between four days and two years to return to their capture sites (Wilde, 2003; Wilde & Paulson, 2003). Black bass that had been captured using hook-and-line methodology in general exhibit a trend for greater swimming distances and take longer to return to nests than in control experiments (Hanson et al., 2007 and references therein; Kaintz & Bettoli, 2010). Additional impacts on population structure following C&R events are those which affect the reproductive success of captured individuals. This is exemplified by those species exhibiting parental care, such as black bass Micropterus spp., and which are frequently angled for during the spawning season, leading to increased nest abandonment rates and depressed reproductive output (Siepker et al., 2007). Largemouth bass Micropterus salmoides captured prior to spawning also produced smaller offspring, indicating more wide ranging effects of C&R practices on the population as a whole (Ostrand et al., 2004). In simulated and wild angling studies there is a general trend for increased nest/brood abandonment in
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black bass as a result of any angling practice compared to control groups (Phillip et al., 1997; Ridgway & Schuter, 1997; Suski et al., 2003; Hanson et al., 2008; Siepker et al., 2009; Hanson et al., 2010; Diana et al., 2012). However, population level studies of black bass populations exploited by tournament fishing have so far produced negligible, or even neutral, effects (Siepker et al., 2007). This would suggest C&R practices in this fishery have a minimal effect on the population size overall. Similarly, common snook Centropomus undecimalis do not appear to be prevented from reproducing even if captured and released from spawning aggregations (Lowerre-Barbieri et al., 2003). Although predation following a C&R event is commonly reported this source of postrelease mortality has rarely been quantified (Donaldson et al., 2010). For example, a bonefish Albula vulpes released following capture was selectively predated by lemon sharks Negaprion brevirostris, despite aggregating with un-caught bonefish (Danylchuk et al., 2007) and in areas where shark numbers were high 39% of bonefish were attacked within 30 minutes of release (Cooke & Phillip, 2004). Similarly, a single black marlin Makaira indica was predated by a shark following release from capture on hook-and-line gear, although seven other tracked individuals in this study exhibited typical behaviours (Pepperell & Davis, 1999). Red snapper Lutjanus campechanus that had been captured by hook-and-line methodology, vented and then released infrequently (5.6%) exhibited erratic swimming behaviour and were exposed to predation close to the release site (30 60 m), possibly as a function of the C&R event or the swimming behaviour itself (Campbell et al., 2010). The evidence suggests that although short-term behaviour may be altered following a C&R event many species do not exhibit long-term effects. This could suggest fish populations are not severely impacted by the disturbance of small numbers of individuals even during sensitive periods of the life cycle, such as spawning.

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The effects of capture in nets Sea bream Pagrus major captured in 10 x 2 m trammel nets exhibited elevated cortisol levels after 10 minutes and these remained significantly higher than resting levels for up to 18 hours of retention in the net (Chopin et al., 1996). Delayed mortality rates of 11% (n = 11) in these fish occurred between eight and 18 days later, while 28% (n = 28) of fish died in the nets. These nets also caused scale removal as well as deep and superficial cuts and resulted in greater levels of stress, injury and mortality compared to hook-and-line capture (Chopin et al., 2006). Spotted seatrout Cynoscion nebulosus mortality as a result of capture in runaround gill nets in Florida was found to be 28%, significantly greater than individuals captured using hook-and-line (4.6%) (Murphy et al., 1995). Southern flounder Paralichthys lethostigma suffered greater short-term mortality (< 72 hours) in commercial gill nets set in the summer (70-78%) compared to autumn fisheries (13-26%) (Smith & Scharf, 2011). The survival rates of lake trout Salvelinus namaycush released from commercial gill nets in Lake Superior in winter was estimated at 21% (range = 18-24%) and in spring 31% (range = 36-36%) and, in general, trout larger than 25 cm total length had greater survival rates compared to smaller individuals (Gallinat et al., 1997). This is much lower than the survival rates of black bream Acanthopagrus butcheri captured using commercial gill nets in the Gippsland Lakes, Australia. Here, total survival rate (initial survivorship plus delayed survivorship) of black bream (c. 6,000 from 372 nets) was estimated as 90.8% ( 3.8% S. E.) (Grixti et al., 2010). There is some indication that C&R methodology using nets may have a greater negative impact than that of hook-and-line methodology on some species, especially in warm water conditions.

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Other impacts of C&R management practices An interesting impact of C&R management practices on fish populations that has rarely been explored is the risk of cumulative mortality as a result of multiple C&R events on a single fish. This effect is particularly important for long-lived or heavily exploited species. Bartholomew & Bohnsack (2005) produced a cumulative mortality model for red grouper Epinephelus morio that indicated that at high encounter rates post-release mortality reached near certainty for long life-spans (72% cumulative probability following 10 C&R events). However, if individual fish learn or express avoidance behaviour following a C&R event then their susceptibility to capture will become lower. Thus, even fisheries enforcing 100% C&R will experience a decline in catch rates over time as a result of decoupling from fish density and/or angler effort. For example, in an experimental study of five lakes in British Columbia, Canada stocked with rainbow trout Oncorhynchus mykiss a sustained fishing effort (8 anglerhours day-1 ha-1) quickly depressed catch rates after just seven to 10 days (Askey et al., 2006). Models fitted to those data suggested that the population contained a group of readily catchable fish that were rapidly captured and then released, but subsequently avoided secondary capture. The same effect was also seen among common carp Cyprinus carpio in stocked ponds, with decreased catches of individuals following experience of just a single C&R event (Raat, 1985). These results compliment experimental work with largemouth bass Micropterus salmoides that suggested vulnerability to capture by angling is actually a heritable trait in this species (Phillip et al., 2009; Redpath et al., 2009). Anglers are not, however, universally attracted to fisheries with high catch rates (Johnston et al., 2011) and so these trends may not necessarily be detrimental to participation numbers. In fact, some studies have suggested that anglers will adjust effort relative not only to perceived fish quality (trophy specimens, hard fighting, etc.) but also in relation to travel time, resulting in poor quality fisheries close to population centres (Post & Parkinson, 2012).
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Some anglers, such as those targeting trophy specimens, will be negatively impacted by C&R management practices in some cases as post-release mortality rates can depress recruitment to older year classes for certain species such as brook trout Salvelinus fontinalis (Risley & Zydlewski, 2010). Handling techniques, another potentially causative agent of post-release mortality for captured fish, are highly species specific but injury as the result of scale or mucus removal is often attributed to handling the fish with dry hands, or damage to the gills when held by the operculum (Pelletier et al., 2007). Currently there are no scientific studies examining the effect of handling or orientation on captured fish.

THE EFFECTS OF CATCH-AND-RELEASE MANAGEMENT PRACTICES ON IBIS PRIORITY-MANAGEMENT SPECIES

CASE STUDY - BROWN TROUT Salmo trutta Brown trout Salmo trutta (hereafter termed trout) are frequently targeted by recreational anglers in small streams and rivers through to dedicated fishing ponds and up to large lake systems and even estuaries and coastal waters. Such is their socio-economic importance (Cooke & Cowyx, 2006) that hatchery-reared trout are regularly stocked in order to supplement wild populations (e.g., Hesthagen et al., 1999), particularly in areas exposed to high intensity fishing pressure (Arlinghaus et al., 2002). This practice often runs contrary the desires of conservation managers who wish to protect localised populations (Araguas et al., 2008). From a fishery perspective the impact of this management practice may be seen as either positive or negative, depending on the response by anglers. If fishing effort remains
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constant then a greater density of fish (mixed wild and stocked) could reduce the chance that any particular individual would be captured, and perhaps is more beneficial to wild fish that may be less susceptible to capture compared to stocked fish. If, however, anglers are attracted by higher catch rates then a greater density of fish may result in an overall increase in fishing pressure. For example, in a mark-recapture study Baer et al. (2007) found that recently stocked trout accounted for 67-83% of all fish caught in a German river fishery over a two year period, and that individual stocked trout were captured in less than 160 days. In central Spain, angling pressure has negatively impacted trout populations by decreasing density, biomass, fecundity and brood stock relative to unexploited rivers elsewhere (Cooke & Cowx, 2006). However, it is unlikely hook-and-line fisheries are as capable of depleting trout stocks as rapidly or as comprehensively as the use of gillnets, as was seen in Finland (Syrjnen & Valkeajrvi, 2010). In Montana, United States where trout are non-native, mortality rates related to C&R angling using single hooks were less than 5%, and were unrelated to water temperatures (Boyd et al., 2010). Osmoregulatory and metabolic disturbances in another nonnative trout population were not found to be significantly impacted by C&R practices in the western United States (Wedemeyer & Wydoski, 2008).

CASE STUDY ATLANTIC SALMON Salmo salar Many Atlantic salmon Salmo salar (hereafter termed salmon) populations undertake extensive migrations into freshwater systems to spawn, and some subsequently leave the system as a kelt following successful spawning (Haltunnen et al., 2010). Different components of these populations exhibiting variation in migration timing can be differentially exploited by anglers in some river systems (Thorley et al., 2007). This can be problematic as
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during the migratory period salmon do not feed and are exposed to a wide range of ambient water temperatures during different seasons (Brobbel et al., 1996). Therefore, these factors have the potential to exacerbate the sub-lethal physiological effects of C&R practices and potentially result in inflated post-release mortality rates (Tufts et al., 1991). For example, kelts are exhausted during a C&R event more quickly (mean S. E., 187 11 s) than prespawning salmon (320 26 s) (Brobbel et al., 1996). White muscle phosphocreatine levels were decreased to a greater extent in kelts (57%) compared to pre-spawning salmon (25%), and lactate levels were also lower in kelts (22.1 1 mmol L-1) compared to pre-spawning salmon (40.6 3.6 mmol L-1) (Brobbel et al., 1996). Despite these findings, however, no kelts in this study died while 12% of pre-spawning salmon died within 12 hours of postrelease. Glycogen concentrations in grilse (one sea-winter fish) can be depressed by as much as 73% during a C&R event and be as low as 44% of resting levels as long as 12 hours postrelease (Booth et al., 1995). In addition, this study also found that the lactate concentrations of the white muscle of grilse were significantly greater than those of multi-sea-winter salmon and, elsewhere, lactate concentrations increased in a linear fashion with the angling duration (Thorstad et al., 2003). The influence of water temperature could have a strong impact on C&R-related mortality as salmon angled at 8C were found to have a resting heart rate of 40 0.28 (mean S. E.) beats-per-minute (BPM), at 16.5C it was 66.9 0.5 BPM and at 20C was 72.3 0.4 BPM, with heart rates remaining elevated for up to 16 hours post-release (Anderson et al., 1998). The impact of water chemistry on physiological changes during C&R events is unclear, but water hardness (CaCO3 mg L-1) appears to have very little effect on exhaustively exercised salmon (Kieffer et al., 2002). Mortality rates of salmon captured in Newfoundland using hook-and-line methodology were 8.2% (Dempson et al., 2002) and 4% of kelts captured in Norway suffered
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mortality following a C&R event (Haltunnen et al., 2010). On the Kola Peninsula, Russian Federation approximately 11% of angled salmon that were released survived to be recaptured, with up to 10% being recaptured a second time and 0.5% three times (Whoriskey et al., 2000) suggesting hook-and-line methodology results in low levels of post-release mortality. However, in a trap-net fishery in the Bay of Bothnia average mortality rates as a result of capture were 11% (4-21%) indicating this could be a more injurious technique for populations overall (Siira et al., 2006). In spite of this Siira et al. (2006) go on to suggest (derived from modelling data) that cumulative mortality following three capture events for any given individual was less than 7%, and that maximum cumulative mortality for the total population following repeated C&R events could be as low as 2% overall. Commercial trawling appears highly injurious to salmon populations, as a Finnish lake pelagic trawl killed significantly more salmon than either brown trout Salmo trutta or pike-perch Stizostedion lucioperca (Jurvelius et al., 2000), despite using the same gear and in similar environmental conditions. In a wide-scale study of salmon anglers in Norway, Thorstad et al. (2003) found that fish were hooked in the mouth on 93% of occasions, while 7% were deeply-hooked. Those deeply-hooked fish were characterised as being in bad or very bad condition compared with fish hooked in the mouth (2 = 124.5, d.f. = 1, P < 0.01). These salmon tended to move downstream (83%, n = 25) and migrated an average of 2.2 km (range 0.13-5.1 km) in the first week post-capture (Thorstad et al., 2006). Salmon that have recently entered freshwater (< 1 week) and subjected to a C&R event do not appear to be strongly negatively affected, as 17 of 18 captured and released fish were subsequently (1 3.5 months) located on spawning grounds in a study by Thorstad et al. (2007). However, it was found that they took an average of 34 days (range 0-94 days) to migrate just one kilometre, indicating a significant delay in migration may be caused by C&R practices.
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The post-release behaviour of salmon does not appear to be altered drastically as a result of C&R practices, as Norwegian kelts were found to migrate similar distances (> 30 km) and at similar speeds to control fish, although there was a short delay in river descent in captured fish (Haltunnen et al., 2010). However, in a comparative study between hook-andline and gillnet capture methodologies of upstream migrating salmon found that fish captured in gillnets moved up to 17.5 km day-1 back downstream, while those captured using hookand-line moved up to 2.1 km day-1 (Mkinen et al., 2000). These fish did not resume their migration for up to 28 days post-release. It appears that in salmon a short-term downstream movement is typical following a C&R event, and that typical migration patterns resume following a short delay (Jensen et al., 2010). In a recent study of an entire population of salmon from a river in Qubec, Canada it was found that 20% of multi-sea-winter fish were captured and subsequently released, and that these fish were at least one of the parents for 22% of the offspring from that river that were genotyped (Richard et al., 2013). This suggests that the captured fish played an important role in the populations total reproductive output, despite reducing their chances of mating (on an individual level) by 12%, confirming previous studies that suggested the viability of gametes in salmon that have experienced a C&R event are not negatively affected (Booth et al., 1995).

CASE STUDY NORTHERN PIKE Esox lucius Northern pike Esox lucius (hereafter termed pike) populations are a popular target of anglers in both Europe and North America and C&R management practices are an important factor in many fisheries (Arlinghaus & Mehner, 2004; Margenau et al., 2008; Arlinghaus et al., 2010; Jansen et al., 2013), especially given the susceptibility of the species to overexploitation (Pierce et al., 1995; Post et al., 2002; but see Kuparinen et al., 2012).
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Exhaustive exercise (60 seconds) results in muscle lactate concentrations increasing to double the rate of resting fish, but these levels return to normal after one hours rest (Arlinghaus et al., 2009). Both phosphocreatine and muscle ATP levels decline as a result of exhaustive exercise (70% & 66% respectively), returning to normal after one hour (for phosphocreatine) and six hours (for muscle ATP) (Arlinghaus et al., 2009). Exhaustive exercise also caused significant increases in plasma glucose, potassium and sodium concentrations. Interestingly though, additional air exposure (up to 300 seconds) was found to cause only minimal levels of additional physiological disturbance, despite its significant effect on post-release behaviour (Arlinghaus et al., 2009). Following air exposure of up to 300 seconds pike spend a greater proportion of time remaining stationary (vs. active swimming) and can take up to 15 minutes to exhibit movement, typically moving less than 50 metres in the first hour post-release (Arlinghaus et al., 2008; Arlinghaus et al., 2009). Despite moving rapidly away from the release site the day following capture (typically 1.5 2.5 km) pike will return to capture sites in less than three weeks (Arlinghaus et al., 2009). Pike that had been captured using hook-and-line artificial lures and had the lines cut (vs. C&R following hook removal) moved significantly less in the first hour post-release, spent significantly more time resting, took significantly longer to initiate movement and then moved farther from the capture site the day after release (Arlinghaus et al., 2008). This movement pattern would appear to be common to pike populations in general (Klefoth et al., 2008). The foraging strategy of pike following a C&R event appears to be significantly impacted. For example, the growth rates of pike following a C&R event were significantly smaller than those of uncaptured pike (44%) (Stlhammar et al., 2012). However, pike released back into groups were found to initiate feeding faster compare to those released alone suggesting some mitigation may be achievable.

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Short-term mortality of pike appears generally low and not significantly influenced by bait type. For example, initial mortality was found to average 2.4% ( 1.5 S.E) when using a range of natural baits and artificial lures (Arlinghaus et al., 2008), and was just 1% when angled with size 4 treble hooks baited with live fish (DuBois et al., 1994). However, the use of so-called pike hooks (or Swedish hooks) in conjunction with live-bait was found to result in 33% mortality, mainly associated with deeply set hooks, and would appear to be highly injurious to pike populations (DuBois et al., 1994). In a congeneric species, the muskellunge Esox masquinongy, the use of live baits resulted in a 22% mortality of those fish hooked in the gut, most likely from extensive internal trauma (Margenau, 2007). In a hoop net fishery (Ontario, Canada) more than 80% of captured pike exhibited jaw damage, scale loss and fin damage when retained for between two and six days (Colotelo et al., 2013). In this study 25% of pike retained in nets for six days (17C) suffered mortality while 5.3% died if held for just two days.

CASE STUDY SHARKS (Selachimorpha spp.) Sharks are typically top-level predators in both marine and freshwater environments and their life history traits (long life span, slow maturity and low reproduction rates) make them especially vulnerable to exploitation, leading to global concern for their management and conservation (Martin, 2005; Lynch et al., 2010; Afonso et al., 2011; Hsu et al., 2012). Much of this concern is related to the difficulty in assessing post-release mortality or sub-lethal effects caused by physiological disturbance, particularly for pelagic or migratory species (reviewed in Skomal, 2007; Marshall et al., 2012). Longline fisheries for other target species frequently result in the unwanted bycatch of sharks (reviewed in Molina & Cooke, 2012) and may lead to exceptionally high mortality
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rates usually as a factor of the gear type used (Diaz & Serafy, 2005; Gilman et al., 2008; Afonso et al., 2012; Bromhead et al., 2012; but see Yokota et al., 2006). For example, in an Atlantic pelagic longline fishery blue Prionace glauca, silky Carcharhinus falciformis and oceanic whitetip Carcharhinus longimanus sharks suffered greater mortality with J-hooks compared to circle hooks, while the night shark Carcharhinus signatus had 100% mortality on both hook types and the nurse shark Ginglymostoma cirratum had 0% mortality on both (Afonso et al., 2011). When captured on J-hooks these species were frequently hooked in the oesophagus or gut, although circle hooks captured more sharks. However, in a mixed pelagic fishery Afonso et al. (2012) found no significant differences between circle and J-hooks in regards to the catchability or mortality rates in a variety of shark species, and Yokota et al. (2006) drew the same conclusion when studying a single species, the blue shark Prionace glauca. During trawling, gummy sharks Mustelus antarcticus were found to suffer very high rates of mortality (up to 87%) while Port Jackson sharks Heterodontus portusjacksoni were not killed using the same gear for the same tow duration (Frick et al., 2010). Spiny dogfish, or spurdog, Squalus acanthias are frequently targeted by northwest Atlantic fisheries and average mortality rates among those captured in otter trawls (24% 6% S. D) are similar to those captured by hook-and-line methodology (29% 12%), although the blood parameters of trawled individuals appear to be more perturbed compared to those captured on hook-andline (Mandelman & Farrington, 2007). In a study of immediate C&R mortality among tracked Atlantic sharpnose sharks Rhizoprionodon terraenovae mortality (10%) did not appear to be correlated with anatomical hook location as the one individual that died (< 45 minutes post-release) was hooked in the gills, while four specimens were gut hooked and tracked for up to 309 minutes (Gurshin & Szedlmayer, 2004). The sand tiger shark Carcharias taurus appears particularly susceptible to being hooked in the gut (87.4%, n =
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184) as they were found to generally consume prey in a single piece (93.7%), leading to injuries to the oesophagus, gut and pericardium (Lucifora et al., 2009). Sharks are believed to experience prolonged recovery periods following anaerobic exercise compared with teleosts as elasmobranchs lack several physiological mechanisms found among the bony fishes. In gummy sharks Mustelus antarcticus experimentally gillnetted for 60 minutes intramuscular lactate concentrations appeared to be the most useful predictor of post-release mortality, as those individuals in poorest condition also exhibited the highest concentrations (Frick et al., 2012). At 180 minutes, gummy sharks exhibited up to 70% mortality when captured in gillnets, while Port Jackson sharks Heterodontus portusjacksoni appeared to be much more resilient to the same capture stress (100% survivorship) (Frick et al., 2010). In common thresher sharks, plasma lactate and haematocrit increased significantly during C&R events and was positively related to the duration of the angling event (Heberer et al., 2010) and may be similar in additional species (Hyatt et al., 2012). However, in juvenile sandbar sharks, experimental C&R angling using hook-and-line methodology had less of an impact on blood-oxygen delivery than was expected, due to an apparent species-specific physiological reaction (Brill et al., 2008). In this study a 21% increase in haematocrit was seen in angled sandbar sharks, a result of c. 28% increase in mean red blood cell volume (approximately twice the value seen in rainbow trout Oncorhynchus mykiss) and not recorded in other shark species. It seems unlikely such a response is confined only to a single shark species and further investigation of sub-lethal effects of C&R on sharks is required (e.g., Manire et al., 2001; Marshal et al., 2012). Air exposure times of 10 minutes were found to exacerbate physiological disturbance in both gummy Mustelus antarcticus and Port Jackson sharks Heterodontus portusjacksoni, although it was not the main factor contributing to mortality rates (Frick et al., 2010). Similar additive effects of handling times on physiological disturbance were noted among
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bonnethead Sphyrna tiburo, lemon Negaprion brevirostris and bull sharks Carcharhinus leucus (Hyatt et al., 2012). The post-release behaviour of sharks appears to be minimally affected by C&R management practices (Stevens et al., 2000; Hight et al., 2007), particularly those that are captured using hook-and-line methodology (Mysyl et al., 2011). For example, the long-term mortality of blue sharks Prionace glauca following longline capture was modelled using blood samples from tracked sharks and predicted that those specimens landed in a healthy condition (defined by blood chemistry) had a greater than 95% chance of survival postrelease (Moyes et al., 2006). Common thresher sharks Alopias vulpinus that experienced a C&R event did not exhibit unusual movement patterns following capture (Heberer et al., 2010) and Atlantic sharpnose sharks Rhizoprionodon terraenovae do not alter movement patterns following C&R events, regardless of anatomical hooking location (Gurshin & Szedlmayer, 2004).

CASE STUDY SKATES (Rajidae spp.) Skates (Rajidae) are, like sharks, elasmobranchs and as such they too are slow growing, late maturing and exhibit a generally low fecundity, although this is higher than in sharks (Dulvy et al., 2000; Ellis et al., 2011). Such life history traits make them vulnerable to exploitation, even under low intensity effort (Cedrola et al., 2005). This group has historically, and is currently, a commercially important group in both the U.K. and elsewhere in Europe and North America (ICES, 2009). In the U.K. since the 1950s there has been a general decline in the landings of skates (tonnes), a reduction in the abundance of larger species (with a concomitant increase in smaller species), and the local extirpation of several skate species

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(common Dipturus batis, long-nosed Dipturus oxyrhinchus and white Rostroraja alba) (Dulvy et al., 2000). In the Bristol Channel, U.K. beam trawls of a short duration (< 2 hours) resulted in a mean 13% mortality rate of three Raja spp. after two days, while a typical commercial trawl (2.7 4.3 hours) resulted in 45% mortality of four species (Enever et al., 2009). In a study of the impact of commercial trawling gear on four northwest Atlantic skate species (n = 1288) initial mortality was found to be low (< 1%), rising to 15% (range 8 59%) after 72 hours (Mandelman et al., 2013). However, in this study 44% of all specimens landed were classified as injured, ranging from minor trauma and diminished vigour through to deep-tissue trauma and appearing moribund, suggesting that delayed post-release mortality among many skate species is likely to be higher than initial mortality.

REFERENCES Aalbers, S. A., Stutzer, G. M. & Drawbridge, M. A. (2004). The effects of catch-and-release angling on the growth and survival of juvenile white seabass captured on offset circle and Jtype hooks. North American Journal of Fisheries Management 24: 793-800. Afonso, A. S., Hazin, F. H. V., Carvalho, F. et al. (2011). Fishing gear modifications to reduce elasmobranch mortality in pelagic and bottom longline fisheries off Northeast Brazil. Fisheries Research 108: 336-343. Afonso, A. S., Santiago, R., Hazon, H. et al. (2012). Shark bycatch and mortality and hook bite-offs in pelagic longlines: interactions between hook types and leader materials. Fisheries Research 131-133: 9-14.

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Allen, M. S., Walters, C. J. & Myers, R. (2008). Temporal trends in largemouth bass mortality, with fishery implications. North American Journal of Fisheries Management 28: 418-427. Allen, M. S., Walters, C. J. & Myers, R. (2008). Temporal trends in voluntary release of largemouth bass. North American Journal of Fisheries Management 28: 418-427. Als, J. (2008). Influence of anatomical hooking depth, capture depth, and venting on mortality of painted comber (Serranus scriba) released by recreational anglers. ICES, Oxford Journals: 1620-1625. Als, J. (2009). Mortality impact of recreational angling techniques and hook types on Trachynotus ovatus (Linnaeus, 1758) following catch-and-release. Fisheries Research 95: 365-369. Als, J., Arlinghaus, R., Palmer, M. et al. (2009). The influence of type of natural bait on fish catches and hooking location in a mixed-species marine recreational fishery, with implications for management. Fisheries Research 97: 270-277. Als, J., Cerd, M., Deudero, S. et al. (2008). Influence of hook size and type on short-term mortality, hooking location and size selectivity in a Spanish recreational fishery. Journal of Applied Ichthyology 24: 658-663. Als, J., Palmer, M. & Grau, A. M. (2009). Mortality of Diplodus annularis and Lithognathus mormyrus released by recreational anglers: implications for recreational fisheries management. Fisheries Management and Ecology 16: 298-305. Als, J., Palmer, M., Grau, A. M. et al. (2008). Effects of hook size and barbless hooks on hooking injury, catch per unit effort, and fish size in a mixed-species recreational fishery in the western Mediterranean Sea. ICES, Oxford Journals: 899-905.
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Anderson, W. G., Booth, R., Beddow, T. A. et al. (1998). Remote monitoring of heart rate as a measure of recovery in angled Atlantic salmon, Salmo salar (L.). Hydrobiologia 371/372: 233-240. Araguas, R. M., Sanz, M., Fernndez, R. et al. (2008). Genetic refuges for self-sustained fishery: experience in wild brown trout populations in the eastern Pyrenees. Ecology of Freshwater Fish 17: 610-616. Arlinghaus, R. & Hallermann, J. (2007). Effects of air exposure on mortality and growth of undersized pikeperch, Sander lucioperca, at low water temperatures with implications for catch-and-release fishing. Fisheries Management and Ecology 14: 155-160. Arlinghaus, R. (2007). Voluntary catch-and-release can generate conflict within the recreational angling community: a qualitative case study of specialised carp, Cyprinus carpio, angling in Germany. Fisheries Management and Ecology 14: 161-171. Arlinghaus, R., Cooke, S. J. & Cowx, I. G. (2010). Providing context to the global code of practice for recreational fisheries. Fisheries Management and Ecology 17: 146-156. Arlinghaus, R., Cooke, S. J. & Potts, W. (2013). Towards resilient recreational fisheries on a global scale through improved understanding of fish and fisher behaviour. Fisheries Management and Ecology 20: 91-98. Arlinghaus, R., Douglas Jr., B. T., Cooke, S. J. et al. (2012). Benefits and risks of adopting the global code of practice for recreational fisheries. Fisheries 37: 165-172. Arlinghaus, R., Klefoth, T., Cooke, S. J. et al. (2009). Physiological and behavioural consequences of catch-and-release angling on northern pike (Esox lucius L.). Fisheries Research 97: 223-233.

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Arlinghaus, R., Klefoth, T., Gingerich, A. J. et al. (2008). Behaviour and survival of pike, Esox lucius, with a retained lure in the lower jaw. Fisheries Management and Ecology 15: 459-466. Arlinghaus, R., Klefoth, T., Kobler, A. et al. (2008). Size selectivity, injury, handling time, and determinants of initial hooking mortality in recreational angling for northern pike: the influence of type and size of bait. North American Journal of Fisheries Management 28: 123134. Arlinghaus, R., Matsumua, S. & Dieckmann, U. (2010). The conservation and fishery benefits of protecting large pike (Esox lucius L.) by harvest regulations in recreational fishing. Biological Conservation 143: 1444-1459. Arlinghaus, R., Schwab, A., Cooke, S. J. et al. (2009). Contrasting pragmatic and sufferingcentred approaches to fish welfare in recreational angling. Journal of Fish Biology 75: 24482463. Askey, P. J., Richards, S.A., Post, J. R. et al. (2006). Linking angling catch rates and fish learning under catch-and-release regulations. North American Journal of Fisheries Management 26: 1020-1029. Ayvazian, S. G., Wise, B. S. & Young, G. C. (2002). Short-term hooking mortality of tailor (Pomatomus saltatrix) in western Australia and the impact on yield per recruit. Fisheries Research 58: 241-248. Bacheler, N. M. & Buckel, J. A. (2004). Does hook type influence the catch rate, size, and injury of grouper in a North Carolina commercial fishery? Fisheries Research 69: 303-311.

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Bacheler, N. M., Hightower, J. E., Burdick, S. M. et al. (2010). Using generalised linear models to estimate selectivity from short-term recoveries of tagged red drum Sciaenops ocellatus: effects of gear, fate, and regulation period. Fisheries Research 102: 266-275. Bacheler, N. M., Hightower, J. E., Paramore, L. M. et al. (2008). An age-dependent tag return model for estimating mortality and selectivity of an estuarine-dependent fish with high rates of catch and release. Transactions of the American Fisheries Society 137: 1422-1432. Baer, J., Blasel, K. & Diekmann, M. (2007). Benefits of repeated stocking with adult, hatchery-reared brown trout, Salmo trutta, to recreational fisheries? Fisheries Management and Ecology 14: 51-59. Barker, R. J., Taylor, P. H. & Smith, S. (2002). Effect of a change in fishing regulations on the survival and capture probabilities of rainbow trout in the upper Rangitikei River, New Zealand. North American Journal of Fisheries Management 22: 465-473. Bartholomew, A. & Bohnsack, J. A. (2005). A review of catch-and-release angling mortality with implications for no-take reserves. Reviews in Fish Biology and Fisheries 15: 129-154. Beckwith Jr., G. H. & Rand, P. S. (2005). Large circle hooks and short leaders with fixed weights reduce incidence of deep hooking in angled red drum. Fisheries Research 71: 115120. Bendock, T. & Alexandersdottir, M. (1993). Hooking mortality of chinook salmon released in the Kenai River, Alaska. North American Journal of Fisheries Management 13: 540-549. Benoit, H. B., Swain, D. P., Niles, M. et al. (2010). Incidental catch amounts and potential post-release survival of winter skate (Leucoraja ocellata) captured in the scallop dredge fishery in the southern Gulf of St. Lawrence (2006-2008). Canadian Science Advisory Secretariat Research Document 31: 1-21.
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Benot, H. P., Hurlbut, T., Chass, J. et al. (2012). Estimating fishery-scale rates of discard mortality using conditional reasoning. Fisheries Research 125-126: 318-330. Booth, R. K., Kieffer, J. D., Davidson, K. et al. (1995). Effects of late-season catch and release angling on anaerobic metabolism, acid-base status, survival, and gamete viability in wild Atlantic salmon (Salmo salar). Canadian Journal of Fisheries and Aquatic Sciences 52: 283-290. Boyd, J. W., Guy, C. S., Horton, T. B. et al. (2010). Effects of catch-and-release angling on salmonids at elevated water temperatures. North American Journal of Fisheries Management 30: 898-907. Brill, R., Bushnell, P., Schroff, S. et al. (2008). Effects of anaerobic exercise accompanying catch-and-release fishing on blood-oxygen affinity of the sandbar shark (Carcharhinus plumbeus, Nardo). Journal of Experimental Marine Biology and Ecology 354: 132-143. Broadhurst, M. K., Butcher, P. A. & Cullis, B. R. (2011). Post-released mortality of angled sand mullet (Myxus elongates: Mugilidae). Fisheries Research 107: 272-275. Broadhurst, M. K., Butcher, P. A. & Cullis, B. R. (2012). Catch-and-release angling mortality of south-eastern Australian Pomatomus saltatrix. African Journal of Marine Science 34: 289295. Broadhurst, M. K., Butcher, P. A., Brand, C. P. et al. (2007). Ingestion and ejection of hooks: effects on long-term health and mortality of angler-caught yellowfin bream Acanthopagrus australis. Diseases of Aquatic Organisms 74: 17-26. Broadhurst, M. K., Gray, C. A., Reid, D. D. et al. (2005). Mortality of key fish species released by recreational anglers in an Australian estuary. Journal of Experimental Marine Biology and Ecology 321: 171-179.
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Broadhurst, M. K., Uhlmann, S. S. & Millar, R. B. (2008). Reducing discard mortality in an estuarine trawl fishery. Journal of Experimental Marine Biology and Ecology 364: 54-61. Brobbel, M. A., Wilkie, M. P., Davidson, K. et al. (1996). Physiological effects of catch and release angling in Atlantic salmon (Salmo salar) at different stages of freshwater migration. Canadian Journal of Fisheries and Aquatic Sciences 53: 2036-2043. Bromhead, D., Clarke, S., Hoyle, S. et al. (2012). Indication of factors influencing shark catch and mortality in the Marshall Islands tuna longline fishery and management implications. Journal of Fish Biology 80: 1870-1894. Brown, I., Sumpton, W., McLennan, M. et al. (2010). An improved technique for estimating short-term survival of released line-caught fish, and an application comparing barotraumarelief methods in red emperor (Lutjanus sebae Cuvier 1816). Journal of Experimental Marine Biology and Ecology 385: 1-7. Bruesewitz, R. E. & Coble, D. W. (1993). Effects of deflating the expanded swim bladder on survival of burbot. North American Journal of Fisheries Management 13: 346-348. Bugley, K. & Shepherd, G. (1991). Effect of catch-and-release angling on the survival of black sea bass. North American Journal of Fisheries Management 11: 468-471. Burns, K. M. & Froeschke, J. T. (2012). Survival of red grouper (Epinephalus morio) and red snapper (Lutjanus campechanus) caught on J-hooks and circle hooks in the Florida recreational and recreational-for-hire fisheries. Bulletin of Marine Science 88: 633-646. Butcher, P. A., Broadhurst, M. K. & Brand, C. P. (2006). Mortality of sand whiting (Sillago cilliata) released by recreational anglers in an Australian estuary. ICES Journal of Marine Science 63: 567-571.

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Butcher, P. A., Broadhurst, M. K., Hall, K. C. et al. (2012). Assessing barotrauma among angled snapper (Pagrus auratus) and the utility of release methods. Fisheries Research 127128: 49-55. Campana, S. E., Joyce, W., Francis, M. P. et al. (2009). Comparability of blue shark mortality estimates for the Atlantic and Pacific longline fisheries. Marine Ecology Progress Series 396: 161-164. Campbell, M. D., Tolan, J., Strauss, R. et al. (2010). Relating angling-dependent fish impairment to immediate release mortality of red snapper (Lutjanus campechanus). Fisheries Research 106: 64-70. Candy, J. R., Carter, E. W., Quinn, T. P. et al. (1996). Adult chinook salmon behavior and survival after catch and release from purse-seine vessels in Johnstone Strait, British Columbia. North American Journal of Fisheries Management 16: 521-529. Carbines, G. D. (1999). Large hooks reduce catch-and-release mortality of blue cod Parapercis colias in the Marlborough Sounds of New Zealand. North American Journal of Fisheries Management 19: 992-998. Carlson, A. J. & Isermann, D. A. (2010). Mandatory catch and release and maximum length limits for largemouth bass in Minnesota: is exploitation still a relevant concern? North American Journal of Fisheries Management 30: 209-220. Carruthers, E. H., Schneider, D. C. & Neilson, J. D. (2009). Estimating the odds of surviving and identifying mitigation opportunities for common bycatch in pelagic longline fisheries. Biological Conservation 142: 2620-2630.

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Carter, D. W. & Liese, C. (2012). The economic value of catching and keeping or releasing saltwater sport fish in the southeast USA. North American Journal of Fisheries Management 32: 613-625. Cedrola, P. V., Gonzlez, A. M. & Pettovello, A. D. (2005). Bycatch of skates (Elasmobranchii: Arhynchobatidae, Rajidae) in the Patagonian red shrimp fishery. Fisheries Research 71: 141-150. Chopin, F. S., Arimoto, T. & Inoue, Y. (1996). A comparison of the stress response and mortality of sea bream Pagrus major captured by hook and line and trammel net. Fisheries Research 28: 277-289. Cline, T. J., Weidel, B. C., Kitchell, J. F. et al. (2012). Growth response of largemouth bass (Micropterus salmoides) to catch-and-release angling: a 27-year mark-recapture study. Canadian Journal of Fisheries and Aquatic Sciences 69: 224-230. Coggins Jr., L. G., Catalano, M. J., Allen, M. S. et al. (2007). Effects of cryptic mortality and the hidden costs of using length limits in fishery management. Fish and Fisheries 8: 1962010. Colotelo, A. H. & Cooke, S. J. (2011). Evaluation of common angling-induced sources of epithelial damage for popular freshwater sport fish using fluorescein. Fisheries Research 109: 217-224. Colotelo, A. H., Raby, G. D., Hasler, C. T. et al. (2013). Northern pike bycatch in an inland commercial hoop net fishery: Effects of water temperature and net tending frequency on injury, physiology, and survival. Fisheries Research 137: 41-49. Cooke, S. J. & Cowx, I, G. (2004). The role of recreational fishing in global fish crises. BioScience 54: 857-859.
52" "

Cooke, S. J. & Cowx, I, G. (2006). Contrasting recreational and commercial fishing: Searching for common issues to promote unified conservation of fisheries resources and aquatic environments. Biological Conservation 128: 93-108. Cooke, S. J. & Phillip, D. P. (2004). Behaviour and mortality of caught-and-released bonefish (Albula spp.) in Bahamian waters with implications for a sustainable recreational fishery. Biological Conservation 118: 599-607. Cooke, S. J. & Schramm, H. L. (2007). Catch-and-release science and its application to conservation and management of recreational fisheries. Fisheries Management and Ecology 14: 73-79. Cooke, S. J. & Sneddon, L. U. (2007). Animal welfare perspectives on recreational angling. Applied Animal Behaviour Science 104: 176-198. Cooke, S. J. & Suski, C. D. (2004). Are circle hooks an effective tool for conserving marine and freshwater recreational catch-and-release fisheries? Aquatic Conservation: Marine and Freshwater Ecosystems 14: 299-326. Cooke, S. J. & Suski, C. D. (2005). Do we need species-specific guidelines for catch-andrelease recreational angling to effectively conserve diverse fishery resources? Biodiversity and Conservation 14: 1195-1209. Cooke, S. J., Bunt, C. M., Ostrand, K. G. et al. (2004). Angling-induced cardiac disturbance of free-swimming largemouth bass (Micropterus salmoides) monitored with heart rate telemetry. Journal of Applied Ichthyology 20: 28-36. Cooke, S. J., Danylchuk, A. J., Danylchuk, S. E. et al. (2006). Is catch-and-release recreational angling compatible with no-take marine protected areas? Ocean and Coastal Management 49: 342-354.
53" "

Cooke, S. J., Nguyen, V. M., Murchie, K. J. et al. (2012). Scientific and stakeholder perspectives on the use of circle hooks in recreational fisheries. Bulletin of Marine Science 88: 395-410. Cooke, S. J., Phillip, D. P., Dunmall, K. M. et al. (2001). The influence of terminal tackle on injury, handling time, and cardiac disturbance of rock bass. North American Journal of Fisheries Management 21: 333-342. Cooke, S. J., Phillip, D. P., Schreer, J. F. et al. (2000). Locomotory impairment of nesting male largemouth bass following catch-and-release angling. North American Journal of Fisheries Management 20: 968-977. Cooke, S. J., Raby, G. D., Donaldson, M. R. et al. (2013). The physiological consequences of catch-and-release angling: perspectives on experimental design, interpretation, extrapolation and relevance to stakeholders. Fisheries Management and Ecology 20: 268-287. Cowen, L., Trouton, N. & Bailey, R. E. (2007). Effects of angling on Chinook salmon for the Nicola River, British Columbia, 1996-2002. North American Journal of Fisheries Management 27: 256-267. Cowx, I. G., Arlinghaus, R. & Cooke, S. J. (2010). Harmonising recreational fisheries and conservation objectives for aquatic biodiversity in inland waters. Journal of Fish Biology 76: 2194-2215. Cox-Rogers, S. (2004). 1999 coho catch and release mortality studies (marine recreational) in northern British Columbia. Canadian Manuscript Report of Fisheries and Aquatic Sciences 2676: 1-12. Danylchuk, S. E., Danylchuk, A. J., Cooke, S. J. et al. (2007). Effects of recreational angling on the post-release behaviour and predation of bonefish (Albula vulpes): the role of
54" "

equilibrium status at the time of release. Journal of Experimental Marine Biology and Research 346: 127-133. DeBoom, C. S., van Landeghem, M. M., Wahl, D. H. et al. (2010). Effects of four hook removal techniques on feeding, growth, and survival of deeply hooked largemouth bass. North American Journal of Fisheries Management 30: 956-963. Dedual, M. (1996). Observed mortality of rainbow trout caught by different angling techniques in Lake Taupo, New Zealand. North American Journal of Fisheries Management 16: 357-363. Dempson, J. B., Furey, G. & Bloom, M. (2002). Effects of catch and release angling on Atlantic salmon, Salmo salar L., of the Conne River, Newfoundland. Fisheries Management and Ecology 9: 139-147. Diamond, S. L. & Campbell, M. D. (2009). Linking sink or swim indicators to delayed mortality in red snapper by using a condition index. Marine and Coastal Fisheries: Dynamics, Management, and Ecosystem Science 1: 107-120. Diana, M. J., Larsen, A. L., Siepker, M. J. et al. (2012). Effects of tournament compared with catch and release angling on nest abandonment of largemouth bass. North American Journal of Fisheries Management 32: 832-837. Diaz, G. A. & Serafy, J. E. (2005). Longline-caught blue shark (Prionace glauca): factors affecting the numbers available for live release. Fisheries Bulletin 103: 720-724. Diewert, R. E., Nagtegaal, D. A. & Patterson, J. (2002). Results of a marine recreational chinook and coho catch and release mortality study conducted in the lower Strait of Georgia during 2001. Canadian Manuscript Report of Fisheries and Aquatic Sciences 2625: 1-32.

55" "

Diggles, B. K. & Ernst, I. (1997). Hooking mortality of two species of shallow-water reef fish caught by recreational angling methods. Marine and Freshwater Research 48: 479-483. Diggles, B. K., Cooke, S. J., Rose, J. D. et al. (2011). Ecology and welfare of aquatic animals in wild capture fisheries. Reviews in Fish Biology and Fisheries 21: 739-765. Diodati, P. J. & Richards, R. A. (1996). Mortality of striped bass hooked and released in salt water. Transactions of the American Fisheries Society 125: 300-307. Doi, T., Nakamura, T., Yokota, M. et al. (2005). Destiny of hooks remaining in the body of Japanese charr Salvelinus leucomaenis and masu salmon Oncorhynchus masou masou. Nippon Suisan Gakkaishi 71: 348-353. Domeier, M. L., Dewar, H. & Nasby-Lucas, N. (2003). Mortality rate of striped marlin (Tetrapturus audax) caught with recreational tackle. Marine and Freshwater Research 54: 435-445. Donaldson, M. R., Arlinghaus, R., Hanson, K. C. et al. (2008). Enhancing catch-and-release science with biotelemetry. Fish and Fisheries 9: 79-105. Donaldson, M. R., Raby, G. D., Nguyen, V. D. et al. (2013). Evaluation of a simple technique for recovering fish from capture stress: integrating physiology, biotelemetry, and social science to solve a conservation problem. Canadian Journal of Fisheries and Aquatic Sciences 70: 90-100. Douglas, J., Brown, P., Hunt, T. et al. (2010). Evaluating relative impacts of recreational fishing harvest and discard mortality on Murray cod (Maccullochella peelii peeli). Fisheries Research 106: 18-21.

56" "

Dowling, C. E., Hall, K. C. & Broadhurst, M. K. (2010). Immediate fate of angled-andreleased Australian bass Macquaria novemaculeata. Hydrobiologia 641: 145-157. DuBois, R. B. & Dubielzig, R. R. (2004). Effect of hook type on mortality, trauma, and capture efficiency of wild stream trout caught by angling with spinners. North American Journal of Fisheries Management 24: 609-616. Dubois, R. B., Margenau, T. L., Scot, S. R. et al. (1994). Hooking mortality of northern pike angled through ice. North American Journal of Fisheries Management 14: 769-775. Dulvy, N. K., Metcalfe, J. D., Glanville, J. et al. (2000). Fishery stability, local extinctions, and shifts in communication structure in skates. Conservation Biology 14: 283-293. Dunmall, K. M., Cooke, S. J., Schreer, J. F. et al. (2001). The effect of scented lures on the hooking injury and mortality of smallmouth bass caught by novice and experienced anglers. North American Journal of Fisheries Management 21: 242-248. Edwards, G. R., Neumann, R. M., Jacobs, R. P. et al. (2004). Impacts of small club tournaments on black bass populations in Connecticut and the effects of regulation exemptions. North American Journal of Fisheries Management 24: 811-821. Edwards, R. E. (1998). Survival and movement patterns of released tarpon (Megalops atlanticus). Gulf of Mexico Science 16: 1-7. Ellis, J. R., Morel, G., Burt, G. et al. (2011). Preliminary observations on the life history and movements of skates (Rajidae) around the Island of Jersey, western English Channel. Journal of the Marine Biological Association of the UK 91: 1185-1192. Enever, R., Catchpole, T. L., Ellis, J. R. et al. (2009). The survival of skates (Rajidae) caught by demersal trawlers fishing in UK waters. Fisheries Research 97: 72-76.

57" "

Fabrizio, M. C., Scharf, F. S., Shepherd, G. R. et al. (2008). Factors affecting catch-andrelease mortality of bluefish. North American Journal of Fisheries Management 28: 533-546. Farrell, A. P., Gallaugher, P. E. & Routledge, R. (2001). Rapid recovery of exhausted adult coho salmon after commercial capture by troll fishing. Canadian Journal of Fisheries and Aquatic Sciences 58: 2319-2324. Farrell, M. P., Gallaugher, P. E., Fraser, J. et al. (2001). Successful recovery of the physiological status of coho salmon on board a commercial gillnet vessel by means of a newly designed revival box. Canadian Journal of Fisheries and Aquatic Sciences 58: 19321946. Ferguson, R. A. & Tufts, B. L. (1992). Physiological effects of brief air exposure in exhaustively exercised rainbow trout (Oncorhynchus mykiss): Implications for catch and release fisheries. Canadian Journal of Fisheries and Aquatic Sciences 49: 1157-1162. Ferter, K., Borsch, T., Kolding, G. et al. (2013). Angler behaviour and implications for management catch-and-release among marine angling tourists in Norway. Fisheries Management and Ecology 20: 137-147. Flaherty, K. E., Winner, B. L., Vecchio, J. L. et al. (2013). Spatial and size distribution of red drum caught and released in Tampa Bay, Florida, and factors associated with post-release hoking mortality. Gulf and Caribbean Research 25: 29-41. Fredricks, K. T., Meinertz, J. R., Ambrose, R. D. et al. (2012). Feeding response of sport fish after electrical immobilization, chemical sedation, or both. North American Journal of Fisheries Management 32: 679-686. Frick, L. H., Reina, R. D. & Walker, T. I. (2010). Stress related physiological changes and post-release survival of Port Jackson sharks (Heterodontus portusjacksoni) and gummy
58" "

sharks (Mustelus antarcticus) following gill-net and longline capture in captivity. Journal of Experimental Marine Biology and Ecology 385: 29-37. Frick, L. H., Walker, T. I. & Reina, R. D. (2010). Trawl capture of Port Jackson sharks, Heterodontus portusjacksoni, and gummy sharks, Mustelus antarcticus, in a controlled setting: effects of tow duration, air exposure and crowding. Fisheries Research 106: 344-350. Frick, L. H., Walker, T. I. & Reina, R. D. (2012). Immediate and delayed effects of gill-net capture on acid-base balance and intramuscular lactate concentration of gummy shark, Mustelus antarcticus. Comparative Biochemistry and Physiology, Part A 162: 88-93. Gale, M. K., Hinch, S. C., Eliason, E. J. et al. (2011). Physiological impairment of adult sockeye salmon in fresh water after simulated catch-and-release across a range of temperatures. Fisheries Research 112: 58-95. Gale, M. K., Hinch, S. G. & Donaldson, M. R. (2013). The role of temperature in the capture and release of fish. Fish and Fisheries 14: 1-33. Gallinat, M. P., Ngu, H. H. & Shively, J. D. (1997). Short-term survival of lake trout released from commercial gill nets in Lake Superior. North American Journal of Fisheries Management 17: 136-140. Gallman, E. A., Isely, J. J., Tomasso, J. R. et al. (1999). Short-term physiological responses of wild and hatchery-produced red drum during angling. North American Journal of Fisheries Management 19: 833-836. Gilman, E., Clarke, S., Brothers, N. et al. (2008). Shark interactions in pelagic longline fisheries. Marine Policy 32: 1-18.

59" "

Gingerich, M. J., Cooke, S. J., Hanson, K. J. et al. (2007). Evaluation of the interactive effects of air exposure duration and water temperature on the condition and survival of angled and released fish. Fisheries Research 86: 169-178. Gjernes, T., Kronlund, A. R. & Mulligan, T. J. (1993). Mortality of Chinook and coho salmon in their first year of ocean life following catch and release by anglers. North American Journal of Fisheries Management 13: 524-539. Goeman, T. J. (1997). Walleye mortality during a live-release tournament on Mille Lacs, Minnesota USA. North American Journal of Fisheries Management 11: 57-61. Goetz, A., Kerwath, S. E., Attwood, C. G. et al. (2007). Comparison of the effects of different linefishing methods on catch composition and capture mortality of South African temperate reef fish. African Journal of Marine Science 29: 177-185. Goodyear, C. P. (2000). Factors affecting robust estimates of the catch and release mortality using pop-off tag technology. International Commission for the Conservation of Atlantic Tunas Collective Volume of Scientific Papers 51: 968-975. Goodyear, C. P. (2007). Recreational catch and release: resource allocation between commercial and recreational fishermen. North American Journal of Fisheries Management 27: 1189-1194. Grant, E. C., Inendino, K. R., Love, W. J. et al. (2005). Effects of practices related to catchand-release angling on mortality and viral transmission in juvenile largemouth bass infected with largemouth bass virus. Journal of Aquatic Animal Health 17: 315-322. Graves, J. E. & Horodysky, A. Z. (2010). Asymmetric conservation benefits of circle hooks in multispecies billfish recreational fisheries: a synthesis of hook performance and analysis of blue marlin (Makaira nigricans) postrelease survival. Fishery Bulletin 108: 433-441.
60" "

Graves, J. E., Horodysky, A. Z. (2008). Does hook choice matter? Effects of three circle hook models on postrelease survival of white marlin. North American Journal of Fisheries Management 28: 471-480. Grixti, D., Conron, S. D. & Jones, P. L. (2007). The effect of hook/bait size and angling technique on the hooking location and the catch of recreationally caught black bream Acanthopagrus butcheri. Fisheries Research 84: 338-344. Grixti, D., Conron, S. D. & Morison, A. (2010). Post-release survival of recreationally caught snapper, Pagrus auratus, in Port Phillip Bay, south-eastern Australia. Fisheries Management and Ecology 17: 1-9. Grixti, D., Conron, S. D., Morison, A. et al. (2008). Estimating post-release survival and the influential factors for recreationally caught black bream (Acanthopagrus butcheri) in the Glenelg River, south eastern Australia. Fisheries Research 92: 303-313. Grixti, D., Morison, A. & Bell, J. D. (2010). Undersized Acanthopagrus butcheri caught and released from commercial gill nets show high survival rates in the Gippsland Lakes, southeastern Australia. North American Journal of Fisheries Management 30: 723-731. Gurshin, C. W. D. & Szedlmayer, S. T. (2004). Short-term survival and movements of Atlantic sharpnose sharks captured by hook-and-line in the north-east Gulf of Mexico. Journal of Fish Biology 65: 973-986. Gustaveson, A. W., Wydoski, R. S. & Wedemeyer, G. A. (1991). Physiological response of largemouth bass to angling stress. Transactions of the American Fisheries Society 120: 629636.

61" "

Hall, K. C., Broadhurst, M. K. & Butcher, P. K. (2012). Post-release mortality of angled golden perch Macquaria ambigua and Murray cod Maccullochella peelii. Fisheries Management and Ecology 19: 10-21. Hall, K. C., Butcher, P. A. & Broadhurst, M. K. (2009). Short-term mortality of Australian bass, Macquaria novemaculeata, after catch-and-release angling. Fisheries Management and Ecology 16: 235-247. Haltunnen, E., Rikardsen, A. H., Thorstad, E. B. et al. (2010). Impact of catch-and-release practices on behaviour and mortality of Atlantic salmon (Salmo salar L.) kelts. Fisheries Research 105: 141-147. Hansen, L. P. (1990). Exploitation of Atlantic salmon (Salmo salar L.) from the River Drammenselv, SE Norway. Fisheries Research 10: 125-135. Hanson, K. C., Cooke, S. J., Suski, C. D. et al. (2007). Effects of different angling practices on post-release behaviour of nest-guarding male black bass, Micropterus spp. Fisheries Management and Ecology 14: 141-148. Hanson, K. C., Gravel, M-A., Redpath, T. et al. (2008). Latitudinal variation in physiological and behavioral responses of nest-guarding smallmouth bass to common recreational angling practices. Transactions of the American Fisheries Society 137: 1558-1566. Hartley, R. A. & Moring, J. R. (1995). Differences in mortality between largemouth and smallmouth bass caught in tournaments. North American Journal of Fisheries Management 15: 666-670. Heberer, C., Aalbers, S. A., Bernal, D. et al. (2010). Insights into catch-and-release survivorship and stress-induced blood biochemistry of common thresher sharks (Alopias

62" "

vulpinus) captured in the southern California recreational fishery. Fisheries Research 106: 495-500. Henderson, I. M. (2009). Optimising recreational harvests of blue cod: the effects of catchand-release mortality and size-selectivity. Fisheries Research 99: 184-195. Hesthagen, T., Jonsson, B. & Skurdal, J. (1989). Survival, exploitation and movement of takeable size brown trout, Salmo trutta L., in a Norwegian river. Aquaculture and Fisheries Management 20: 475-484. Hewlett, N. R., Snow, J. & Britton, J. R. (2009). The role of management practices in fish kills in recreational lake fisheries in England in Wales. Fisheries Management and Ecology 16: 248-254. Hight, B. V., Holts, D., Graham, J. B. et al. (2007). Plasma catecholamine levels as indicators of the post-release survivorship of juvenile pelagic sharks caught on experimental drift longlines in the Southern California Bight. Marine and Freshwater Research 58: 145-151. Hochhalter, S. J. & Reed, D. J. (2011). The effectiveness of deepwater release at improving the survival of discarded yelloweye rockfish. North American Journal of Fisheries Management 31: 852-860. Hoffman, G. C., Coble, D. W., Frie, R. V. et al. (1996). Walleye and sauger mortality associated with live-release tournaments of the Lake Winnebago system, Wisconsin. North American Journal of Fisheries Management 16: 364-370. Horodysky, A. Z. & Graves, J. E. (2005). Application of pop-up satellite archival tag technology to estimate postrelease survival of white marlin (Tetrapturus albidus) caught on circle and straight-shank ("J") hooks in the western North Atlantic recreational fishery. Fishery Bulletin 103: 84-96.
63" "

Hsu, H. H., Joung, S. G. & Liu, K. M. (2012). Fisheries, management and conservation of the whale shark Rhincodon typus in Taiwan. Journal of Fish Biology 80: 1595-1607. Hutt, C. P. & Bettoli, P. W. (2007). Preferences, specialization, and management attitudes of trout anglers fishing in Tennessee tailwaters. North American Journal of Fisheries Management 27: 1257-1267. Hyatt, M. W., Anderson, P. A., ODonnell, P. M. et al. (2012). Assessment of acid-base derangements among bonnethead (Sphyrna tiburo), bull (Carcharhinus leucus), and lemon (Negaprion brevirostris) sharks from gillnet and longline capture and handling methods. Comparative Biochemistry and Physiology, Part A 162: 113-120. Hyvrinen, P., Leppniemi, V., Johansson, K. et al. (2008). Stress and survival of small pikeperch Sander lucioperca (L.) after trawling and chilling. Journal of Fish Biology 72: 26772688. Jackson, J. J. & Willis, D. W. (1991). Short-term mortality of smallmouth bass caught during a live-release tournament at Lake Oahe, South Dakota. Prairie Naturalist 23: 201-204. Jansen, T., Arlinghaus, R., Als, T. D. et al. (2013). Voluntary angler logbooks reveal longterm changes in a lentic pike, Esox lucius, population. Fisheries Management and Ecology 20: 125-136. Jenkins, T. M. (2003). Evaluating recent innovations in bait fishing tackle and technique for catch and release of rainbow trout. North American Journal of Fisheries Management 23: 1098-1107. Jensen, J. L. A., Haltunnen, E., Thorstad, E. B. et al. (2010). Does catch-and-release angling alter the migratory behaviour of Atlantic salmon? Fisheries Research 106: 550-554.

64" "

Jensen, O. P., Gilroy, D. J., Hogan, Z. et al. (2009). Evaluating recreational fisheries for an endangered species: a case study of taimen, Hucho taimen, in Mongolia. Canadian Journal of Fisheries and Aquatic Sciences 66: 1707-1718. Jensen, O. P., Ortega-Garcia, S., Martell, S. J. D. et al. (2010). Local management of a highly migratory species: the effects of long-line closures and recreational catch-andrelease for Baja California striped marlin fisheries. Progress in Oceanography 86: 176-186. Jiang, H., Pollock, K. H., Bownie, C. et al. (2007). Tag return models allowing for harvest and catch and release: evidence of environmental and management impacts on striped bass fishing and natural mortality rates. North American Journal of Fisheries Management 27: 387-396. Johnston, F. D., Arlinghaus, R., Stelfox, J. et al. (2011). Decline in angler use despite increased catch rates: anglers response to the implementation of a total catch-and-release regulation. Fisheries Research 110: 189-197. Jokikokko, E., Kallio-Nyberg, I., Jutila, E. et al. (2006). Effect of origin, sex and sea age of Atlantic salmon on their recapture rate after river ascent. Journal of Applied Ichthyology 22: 489-494. Jones, T. S. (2005). The influence of circle hooks on the capture efficiency and injury rate of walleyes. North American Journal of Fisheries Management 25: 725-731. Jurvelius, J., Riikonen, R., Marjomki, T. J. et al. (2000). Mortality of pike-perch (Stizostedion lucioperca) and landlocked salmon (Salmo salar m. sebago) caught as by-catch in pelagic trawling in a Finnish lake. Fisheries Research 45: 291-296.

65" "

Kaimmer, S. M. & Trumble, R. J. (1998). Injury, condition, and mortality of Pacific halibut bycatch following careful release by Pacific cod and sablefish longline fisheries. Fisheries Research 38: 131-144. Kaintz, M. A. & Bettoli, P. W. (2010). Dispersal of smallmouth bass from a simulated tournament weigh-in site. North American Journal of Fisheries Management 30: 976-982. Keniry, M. J., Brofka, W. A., Horns, W. H. et al. (1996). Effects of decompression and puncturing the gas bladder on survival of tagged yellow perch. North American Journal of Fisheries Management 16: 201-206. Kerstetter, D. W. & Graves, J. E. (2006). Survival of white marlin (Tetrapturus albidus) released from commercial pelagic longline gear in the western North Atlantic. Fishery Bulletin 104: 434-444. Kerstetter, D. W. & Graves, J. E. (2008). Postrelease survival of sailfish caught by commercial pelagic longline gear in the southern Gulf of Mexico. North American Journal of Fisheries Management 28: 1578-1586. Kieffer, J. D., Rossiter, A. M., Kieffer, C. A. et al. (2002). Physiology and survival of Atlantic salmon following exhaustive exercise in hard and softer water: Implications for the catch-and-release sport fishery. North American Journal of Fisheries Management 22: 132144. Killen, S. S., Suski, C. D., Cooke, S. J. et al. (2006). Factors contributing to the physiological disturbance in walleyes during simulated live-release angling tournaments. Transactions of the American Fisheries Society 135: 557-569. Kitamura, S., Ikuta, K., Shikama, T. et al. (2005). Effects of catch-and-release (C&R) regulation in the Yukawa River. Bulletin of Fisheries Research Agency 15: 1-10.
66" "

Klefoth, T., Kobler, A. & Arlinghaus, R. (2008). The impact of catch-and-release angling on short-term behaviour and habitat choice of northern pike (Esox lucius L.). Hydrobiologia 601: 99-110. Klefoth, T., Kobler, A. & Arlinghaus, R. (2011). Behavioural and fitness consequences of direct and indirect non-lethal disturbances in a catch-and-release northern pike (Esox lucius) fishery. Knowledge and Management of Aquatic Ecosystems 403 (11). Kuparinen, A., Alho, J. S., Olin, M. et al. (2012). Estimation of northern pike population sizes via mark-recapture monitoring. Fisheries Management and Ecology 19: 323-332. Kuparinen, A., Klefoth, T. & Arlinghaus, R. (2010). Abiotic and fishing-related correlates of angling catch rates in pike (Esox lucius). Fisheries Research 105: 111-11. Kwak, T. J. & Henry, M. G. (1995). Largemouth bass mortality and related causal factors during live-release fishing tournaments on a large Minnesota Lake. North American Journal of Fisheries Management 15: 621-630. Lee, D. P. (1992). Gas bladder deflation of depressurized largemouth bass. North American Journal of Fisheries Management 12: 662-664. Lindsay, R. B., Schroeder, R. K., Kenaston, K. R. et al. (2004). Hooking mortality by anatomical location and its use in estimating mortality of spring Chinook salmon caught and released in a river sport fishery. North American Journal of Fisheries Management 24: 367378. Lowerre-Barbieri, S. K., Vose, F. E. & Whittington, J. A. (2003). Catch-and-release fishing on a spawning aggregation of common snook: does it affect reproductive output? Transactions of the American Fisheries Society 132: 940-952.

67" "

Lucifora, L. O., Garca, V. B. & Escalante, A. H. (2009). How can the feeding habits of the sand tiger shark influence the success of conservation programmes? Animal Conservation 12: 291-301. Lyle, J. M., Moltschaniwskyj, N. A., Morton, A. J. et al. (2007). Effects of hooking damage and hook type on post-release survival of sand flathead (Platycephalus bassensis). Marine and Freshwater Research 58: 445-453. Lynch, A-M. J., Sutton, S. G. & Simpfendorfer, C. A. (2010). Implications of recreational fishing for elasmobranch conservation in the Great Barrier Reef Marine Park. Aquatic Conservation: Marine and Freshwater Ecosystems 20: 312-318. Mkinen, T. S., Niemel, E., Moen, K. et al. (2000). Behaviour of gill-net and rod-captured Atlantic salmon (Salmo salar L.) during upstream migration and following radio tagging. Fisheries Research 45: 117-127. Malchoff, M. H. & Heins, S. W. (1997). Short-term hooking mortality of weakfish caught on single-barb hooks. North American Journal of Fisheries Management 17: 477-481. Mandelman, J. W. & Farrington, M. A. (2007). The estimated short-term discard mortality of a trawled elasmobranch, the spiny dogfish (Squalus acanthias). Fisheries Research 83: 238245. Mandelman, J. W., Cicia, A. M., Ingram Jr., G. W. et al. (2013). Short-term post-release mortality of skates (family Rajidae) discarded in a western North Atlantic commercial otter trawl fishery. Fisheries Research 139: 76-84. Mandelman, J. W., Morrison, R. A., Cavin, J. M. et al. (2012). The blood chemical status of Atlantic cod Gadus morhua following capture by jig and demersal longline with differential hook removal methods. Journal of Fish Biology 81: 1406-1414.
68" "

Manire, C., Hueter, R., Hull, E. et al. (2001). Serological changes associated with gill-net capture and restraint in three species of sharks. Transactions of the American Fisheries Society 130: 1038-1048. Mapleston, A., Welch, D., Begg, G. A. et al. (2008). Effects of change in hook pattern and size on catch rate, hooking location, injury and bleeding for a number of tropical reef fish species. Fisheries Research 91: 203-211. Margenau, T. L. (2007). Effects of angling with a single-hook and live bait on muskellunge survival. Environmental Biology of Fishes 79: 155-162. Margenau, T. L., AveLallemant, S. P., Giehtbrock, D. et al. (2008). Ecology and management of northern pike in Wisconsin. Hydrobiologia 601: 111-123. Margenau, T. L., Gilbert, S. J. & Hatzenbeler, G. R. (2003). Angler catch and harvest of northern pike in northern Wisconsin lakes. North American Journal of Fisheries Management 23: 307-312. Marshall, H., Field, L., Afiadata, A. et al. (2012). Hematological indicators of stress in longline-captured sharks. Comparative Biochemistry and Physiology, Part A 162: 121-129. Martin, R. A. (2005). Conservation of freshwater and euryhaline elasmobranchs: a review. Journal of the Marine Biological Association of the U.K. 85: 1049-1073. Meals, K. O. & Miranda, L. E. (1994). Size-related mortality of tournament-caught largemouth bass. North American Journal of Fisheries Management 14: 460-463. Meerbeek, J. R. & Hoxmeier, R. J. H. (2011). Winter catch-and-release hooking mortality of saugers below Lock and Dam 3 of the Mississippi River. North American Journal of Fisheries Management 31: 197-202.

69" "

Meka, J. M. & Margraf, F. J. (2007). Using a bioenergetic model to assess growth reduction from catch-and-release fishing and hooking injury in rainbow trout, Oncorhynchus mykiss. Fisheries Management and Ecology 14: 131-139. Meka, J. M. & McCormick, S. D. (2005). Physiological response of wild rainbow trout to angling: impact of angling duration, fish size, body condition, and temperature. Fisheries Research 72: 311-322. Meka, J. M. (2004). The influence of hook type, angler experience, and fish size on injury rates and the duration of capture in an Alaskan catch-and-release rainbow trout fishery. North American Journal of Fisheries Management 24: 1309-1321. Millard, M. J., Mohler, J. W., Kahnle, A. et al. (2005). Mortality associated with catch-andrelease angling of striped bass in the Hudson River. North American Journal of Fisheries Management 25: 1533-1541. Millard, M. J., Welsh, M. A., Fletcher, J. W. et al. (2003). Mortality associated with catch and release of striped bass in the Hudson River. Fisheries Management and Ecology 10: 295300. Molina, J. M. & Cooke, S. J. (2012). Trends in shark bycatch research: current status and research needs. Reviews in Fish Biology and Fisheries 22: 719-737. Morrissey, M. B., Suski, C. D., Esseltine, K. R. et al. (2005). Incidence and physiological consequences of decompression in smallmouth bass after live-release angling tournaments. Transactions of the American Fisheries Society 134: 1038-1047. Moyes, C. D., Fragoso, N., Musyl, M. K. et al. (2006). Predicting postrelease survival in large pelagic fish. Transactions of the American Fisheries Society 135: 1389-1397.

70" "

Muoneke, M. I. (1992). Hooking mortality of white crappie, Pomoxis annularis Rafinesque, and spotted bass, Micropterus punctulatus (Rafinesque), in Texas reservoirs. Aquaculture and Fisheries Management 23: 87-93. Murphy, M. D., Heagey, R. F., Neugebauer, V. H. et al. (1995). Mortality of spotted seatrout released from gill-net or hook-and-line gear in Florida. North American Journal of Fisheries Management 15: 748-753. Mysyl, M. K., Brill, R. W., Curran, D. D. et al. (2011). Postrelease survival, vertical and horizontal movements, and thermal habitats of five species of pelagic sharks in the central Pacific Ocean. Fishery Bulletin (Seattle) 109: 341-368. Nslund, I., Nordwall, F., Eriksson, T. et al. (2005). Long-term responses of a streamdwelling grayling population to restrictive fishing regulations. Fisheries Research 72: 323332. Neal, J. W. & Lopez-Clayton, D. (2001). Mortality of largemouth bass during catch-andrelease tournaments in a Puerto Rico reservoir. North American Journal of Fisheries Management 21: 834-842. Nelson, K. L. (1998). Catch-and release mortality of striped bass in the Roanoke River, North Carolina. North American Journal of Fisheries Management 18: 25-30. Nguyen, V., Gravel, M-A., Mapleston, A. et al. (2009). The post-release behaviour and fate of tournament-caught smallmouth bass after fizzing to alleviate distended swim bladders. Fisheries Research 96: 313-318. Nyberg, P., Degerman, P. & Sers, B. (1996). Survival after catch in trap-nets, movements and growth of the pikeperch (Stizostedion lucioperca) in Lake Hjalmaren, Central Sweden. Annales Zoologici Fennici 33: 569-575.
71" "

OToole, A. C., Danylchuk, A. J., Suski, C. D. et al. (2010). Consequences of catch-andrelease angling on the physiological status, injury, and immediate mortality of great barracuda (Sphyraena barracuda) in the Bahamas. ICES, Oxford Journals: 1667-1675. Orciari, R. D. & Leonard, G. H. (1990). Catch and release management of a trout stream contaminated with PCBs. North American Journal of Fisheries Management 10: 315-329. Orsi, J. A., Wertheimer, A. C. & Jaenicke, H. W. (1993). Influence of selected hook and lure type on catch, size, and mortality of commercially troll-caught chinook salmon. North American Journal of Fisheries Mangement 13: 709-722. Ostrand, K. G., Siepker, M. J. & Cooke, S. J. (2006). Capture efficiencies of two hook types and associated injury and mortality of juvenile muskellunge angled with live baitfish. North American Journal of Fisheries Management 26: 622-627. Overton, A. S., Zabawski, J. & Riley, K. L. (2008). Release mortality of undersized fish from the snapper-grouper complex off the North Carolina coast. North American Journal of Fisheries Management 28: 733-739. Pankhurst, N. W. & Dedual, M. (1994). Effects of capture and recovery on plasma levels of cortisol, lactate and gonadal steroids in a natural population of rainbow trout. Journal of Fish Biology 45: 1013-1025. Parker, B. R., Schindler, D. W., Wilhelm, F. M. et al. (2007). Bull trout population responses to reductions in angler effort and retention limits. North American Journal of Fisheries Management 27: 848-859. Pawson, M. G., Brown, M., Leballeur, J. et al. (2008). Will philopatry in sea bass, Dicentrarchus labrax, facilitate the use of catch-restricted areas for management of recreational fisheries? Fisheries Research 93: 240-243.
72" "

Pelletier, C., Hanson, K. C. & Cooke, S. J. (2007). Do catch-and-release guidelines from state and provincial fisheries agencies in North America conform to scientifically based best practices? Environmental Management 39: 760-773. Pepperell, J. G. & Davis, D. L. O. (1999). Post-release behaviour of black marlin, Makaira indica, caught off the Great Barrier Reef with sportfishing gear. Marine Biology 135: 369380. Persons, S. E. & Hirsch, S. A. (1994). Hooking mortality of lake trout angled through ice by jigging and set-lining. North American Journal of Fisheries Management 14: 664-668. Philipp, D. P., Cooke, S. J., Claussen, J. E. et al. (2009). Selection for vulnerability to angling in largemouth bass. Transactions of the American Fisheries Society 138: 189-199. Philipp, D. P., Toline, C. A., Kubacki, M. F. et al. (1997). The impact of catch-and-release angling on the reproductive success of smallmouth bass and largemouth bass. North American Journal of Fisheries Management 17: 557-567. Pierce, R. B., Tomcko, C. M. & Schupp, D. H. (1995). Exploitation of northern pike in seven small north-central Minnesota lakes. North American Journal of Fisheries Management 15: 601-609. Pine III, W. E., Martell, S. J. D., Jensen, O. P. et al. (2008). Catch-and-release and size limit regulations for blue, white, and striped marlin: the role of postrelease survival in effective policy design. Canadian Journal of Fisheries and Aquatic Sciences 65: 975-988. Pollock, K. H. & Pine III, W. E. (2007). The design and analysis of field studies to estimate catch-and-release mortality. Fisheries Management and Ecology 14: 123-130.

73" "

Pope, K. L. & Wilde, G. R. (2004). Effect of catch-and-release angling on growth of largemouth bass, Micropterus salmoides. Fisheries Management and Ecology 11: 39-44. Pope, K. L. & Wilde, G. R. (2010). Survival of foul-hooked largemouth bass (Micropterus salmoides). Journal of Freshwater Ecology 25: 135-139. Pope, K. L., Wilde, G. R. & Knabe, D. W. (2007). Effect of catch-and-release angling on growth and survival of rainbow trout, Oncorhynchus mykiss. Fisheries Management and Ecology 14: 115-121. Post, J. R. & Parkinson, E. A. (2012). Temporal and spatial patterns of angler effort across lake districts and policy options to sustain recreational fisheries. Canadian Journal of Fisheries and Aquatic Sciences 69: 321-329. Pratten, D. J. & Shearer, W. M. (1985). The commercial exploitation of sea trout, Salmo trutta L. Aquaculture and Fisheries Management 1: 71-89. Prince, E. D., Snodgrass, D., Orbesen, E. S. et al. (2007). Circle hooks, J hooks and dropback time: a hook performance study of the south Florida recreational live-bait fishery for sailfish, Istiophorus platypterus. Fisheries Management and Ecology 14: 173-182. Raat, A. J. P. (1985). Analysis of angling vulnerability of common carp, Cyprinus carpio L., in catch-and-release angling in ponds. Aquaculture and Fisheries Management 16: 171-187. Raby, G. D., Donaldson, M. R., Hinch, S. G. et al. (2012). Validation of reflex indicators for measuring vitality and predicting the delayed mortality of wild coho salmon bycatch released from fishing gears. Journal of Applied Ichthyology 49: 90-98.

74" "

Rapp, T., Cooke, S. J. & Arlinghaus, R. (2008). Exploitation of specialised fisheries resources: The importance of hook size in recreational angling for large common carp (Cyprinus carpio L.). Fisheries Research 94: 79-83. Rapp, T., Hallermann, J., Cooke, S. J. et al. (2012). Physiological and behavioural consequences of capture and retention in carp sacks on common carp (Cyprinus carpio L.), with implications for catch-and-release recreational fishing. Fisheries Research 125-126: 5768. Redpath, T. D., Cooke, S. J., Arlinghaus, R. et al. (2009). Life-history traits and energetic status in relation to vulnerability to angling in an experimentally selected teleost fish. Evolutionary Applications 2: 312-323. Reeves, K. A. & Bruesewitz, R. E. (2007). Factors influencing the hooking mortality of walleyes caught by recreational anglers on Mille Lacs, Minnesota. North American Journal of Fisheries Management 27: 443-452. Reeves, K. A. & Staples, D. F. (2011). Relative hooking mortality among walleyes caught on barbed and barbless octopus hooks and barbed jigs. North American Journal of Fisheries Management 31: 32-40. Render, J. H. & Wilson, C. A. (1994). Hook-and-line mortality of caught and released red snapper around oil and gas platform structural habitat. Bulletin of Marine Science 55: 11061111. Richard, A., Dionne, M., Wang, J. et al. (2013). Does catch and release affect the mating system and individual reproductive success of wild Atlantic salmon (Salmo salar L.)? Molecular Ecology 22: 187-200.

75" "

Ricks, B. R. & Maceina, M. J. (2007). Effects of catch-and-release tournament fishing on dispersal and population metrics of black bass in Lake Martin, Alabama. Balancing Fisheries Management and Water Uses for Impounded River Systems (American Fisheries Society Symposium) 62: 293-308. Ridgway, M. S. & Schuter, B. J. (1997). Predicting the effects of angling for nesting male smallmouth bass on production of age-0 fish with an individual-based model. North American Journal of Fisheries Management 17: 557-567. Ridley, C. A., Isley, J. J. & Tommasso, J. R. (1997). Effects of supplemental feeding on physical characteristics and physiological responses to angling of striped bass. North American Journal of Fisheries Management 17: 791-794. Risley, C. A. L. & Zydlewski, J. (2010). Assessing the effects of catch-and-release regulations on a brook trout population using an age-structured model. North American Journal of Fisheries Management 30: 1434-1444. Roach, J. P., Hall, K. C. & Broadhurst, M. K. (2011). Effects of barotrauma and mitigation methods on released Australian bass Macquaria novemaculeata. Journal of Fish Biology 79: 1130-1145. Robert, J. J., Larocque, S. M. & Cooke, S. J. (2012). Influence of hook size and style on short-term survival of deeply hooked bluegills. North American Journal of Fisheries Management 32: 869-874. Rogers, B. L., Lowe, C. G. & Fernndez-Juricic, E. (2011). Recovery of visual performance in rosy rockfish (Sebastes rosaceus) following exophthalmia resulting from barotrauma. Fisheries Research 112: 1-7.

76" "

Rudershausen, P. J., Buckel, J. A. & Williams, E. H. (2007). Discard composition and release fate in the snapper and grouper commercial hook-and-line fishery in North Carolina, USA. Fisheries Management and Ecology 14: 103-113. Rummer, J. L. & Bennett, W. A. (2005). Physiological effects of swim bladder overexpansion and catastrophic decompression on red snapper. Transactions of the American Fisheries Society 134: 1457-1470. Scarnecchia, D, L. & Stewart, P. A. (1997). Implementation and evaluation of a catch-andrelease fishery for paddlefish. North American Journal of Fisheries Management 17: 795799. Schill, D. G., Griffith, J. S. & Gresswell, R. E. (1986). Hooking mortality of cutthroat trout Salmo clarkii Bouvieri in a catch-and-release segment of the Yellowstone River, Yellowstone National Park, Wyoming, USA. North American Journal of Fisheries Management 6: 226232. Schill, D. J. & Scarpella, R. L. (1997). Barbed hook restrictions in catch-and-release trout fisheries: a social issue. North American Journal of Fisheries Management 17: 873-881. Schill, D. J. (1996). Hooking mortality of bait-caught rainbow trout in an Idaho trout stream and a hatchery: Implications for special-regulation management. North American Journal of Fisheries Management 16: 348-356. Schneider, J. C. & Lockwood, R. N. (2002). Use of walleye stocking, antimycin treatments, and catch-and-release angling regulations to increase growth and length of stunted bluegill populations in Michigan lakes. North American Journal of Fisheries Management 22: 10411052.

77" "

Schramm, H. L., Vondracek, B., French, W. E. et al. (2010). Factors associated with mortality of walleyes and saugers caught in live-release tournaments. North American Journal of Fisheries Management 30: 238-258. Schreer, J. F., Gokey, J. & DeGhett, V. J. (2009). The incidence and consequences of barotrauma in fish in the St. Lawrence River. North American Journal of Fisheries Management 29: 1707-1713. Schreer, J. F., Resch, D. M., Gatley, M. L. et al. (2005). Swimming performance of brook trout after simulated catch-and-release angling: Looking for air exposure thresholds. North American Journal of Fisheries Management 25: 1513-1517. Schultz, R. D. & Robinson, D. A. (2002). Exploitation and mortality rates of white bass in Kansas reservoirs. North American Journal of Fisheries Management 22: 652-658. Serafy, J. E., Orbesen, E. S., Snodgrass, D. J. G. et al. (2012). Hooking survival of fishes captured by the United States Atlantic pelagic longline fishery: Impact of the 2004 circle hook rule. Bulletin of Marine Science 88: 605-621. Shasteen, S. P. & Sheehan, R. J. (1997). Laboratory evaluation of artificial swim bladder deflation in largemouth bass: Potential benefits for catch-and-release fisheries. North American Journal of Fisheries Management 17: 32-37. Siepker, M. J., Cooke, S. J., Wahl, D. H. et al. (2009). Individual reproductive success of largemouth bass and smallmouth bass subjected to different components of competitive angling events. Transactions of the American Fisheries Society 138: 818-825. Siepker, M. J., Ostrand, K. G., Cooke, S. J. et al. (2007). A review of the effects of catchand-release angling on black bass, Micropterus spp.: implications for conservation and management of populations. Fisheries Management and Ecology 14: 91-101.
78" "

Siira, A., Suuronen, P., Ikonen, E. et al. (2006). Survival of Atlantic salmon captured in and released from a commercial trap-net: potential for selective harvesting of stocked salmon. Fisheries Research 80: 280-294. Skomal, G. B. (2007). Evaluating the physiological and physical consequences of capture on post-release survivorship in large pelagic fishes. Fisheries Management and Ecology 14: 8189. Smale, M. J., Booth, A. J., Farquhar, M. R. et al. (2012). Migration and habitat use of formerly captive and wild raggedtooth sharks (Carcharias taurus) on the southeast coast of South Africa. Marine Biology Research 8: 115-128. Small, I. (1987). The performance of a small catch-and-release fishery stocked with rainbow trout, Salmo gairdneri Richardson, and thoughts on stocking density. Aquaculture and Fisheries Management 18: 291-308. Smit, N. J., Howatson, G. & Greenfield, R. (2009). Blood lactate levels as a biomarker for angling-induced stress in tigerfish Hydrocynus vittatus from the Okavango Delta, Botswana. African Journal of Aquatic Science 34: 255-259. Smith, D. R., Burnham, K. P., Kahn, D. M. et al. (2000). Bias in survival estimates from tagrecovery models where catch-and-release is common, with an example from Atlantic striped bass (Morone saxitilis). Canadian Journal of Fisheries and Aquatic Sciences 57: 886-897. Smith, P. A. (2002). The relationship between stock and catch and the effect of bait on catch as determined from a UK recreational catch and release fishery. Fisheries Management and Ecology 9: 261-266.

79" "

Smith, W. E. & Scharf, F. S. (2011). Postrelease survival of sublegal southern flounder captured in a commercial gill-net fishery. North American Journal of Fisheries Management 31: 445-454. St John, J. & Syers, C. J. (2005). Mortality of the West Australian dhufish, Glaucosoma hebraicum (Richardson 1845) following catch-and-release: The influence of capture depth, venting and hook type. Fisheries Research 76: 106-116. Stlhammar, M., Linderfalk, R., Brnmark, C. et al. (2012). The impact of catch-and-release on the foraging behaviour of pike (Esox lucius) when released alone or into groups. Fisheries Research 125-126: 51-56. Steeger, T. M., Grizzle, J. M., Weathers, K. et al. (1994). Bacterial diseases and mortality of angler-caught largemouth bass released after tournaments on Walter F. George Reservoir, Alabama-Georgia. North American Journal of Fisheries Management 14: 435-441. Steinhart, G. B., Marschall, E. A. & Stein, R. A. (2004). Round goby predation on smallmouth bass offspring in nests during simulated catch-and-release angling. Transactions of the American Fisheries Society 133: 121-131. Stensland, S. (2012). Typology of landowners in Norwegian salmon angling: attitudes towards river owner organisations and management actions. Fisheries Management and Ecology 19: 273-282. Stephen, J. A. & Harris, P. J. (2010). Commercial catch composition with discard and immediate release mortality proportions off the southeastern coast of the United States. Fisheries Research 103: 18-24.

80" "

Stevens, J. D., West, G, J. & McLoughlin, K. J. (2000). Movements, recapture patterns, and factors affecting the return rate of carcharhinid and other sharks tagged off northern Australia. Marine and Freshwater Research 51: 127-141. Stewart, J. (2008). Capture depth related mortality of discarded snapper (Pagrus auratus) and implications for management. Fisheries Research 90: 289-295. Stokesbury, M. J. W., Neilson, J. D., Susko, E. et al. (2011). Estimating mortality of Atlantic bluefin tuna (Thunnus thynnus) in an experimental recreational catch-and release fishery. Biological Conservation 144: 2684-2691. Stunz, G. W. & McKee, D. A. (2006). Catch-and-release mortality of spotted seatrout in Texas. North American Journal of Fisheries Management 26: 843-848. Sumpton, W., Mayer, D., Brown, I. et al. (2008). Investigation of movement and factors influencing post-release survival of line-caught coral reef fish using recreational tag-recapture data. Fisheries Research 92: 189-195. Suski, C. D., Cooke, S. J., Danylchuk, A. J. et al. (2007). Physiological disturbance and recovery dynamics of bonefish (Albula vulpes), a tropical marine fish, in response to variable exercise and exposure to air. Comparative Biochemistry and Physiology, Part A 148: 664673. Suski, C. D., Killen, S. S., Morrissey, M. B. et al. (2003). Physiological changes in largemouth bass caused by live-release angling tournaments in southeastern Ontario. North American Journal of Fisheries Management 23: 760-769. Suski, C. D., Phelan, F. J. S., Kubacki, M. F. et al. (2002). The use of sanctuaries for protecting nesting black bass from angling. Black Bass: Ecology, Conservation & Management (American Fisheries Society Symposium) 31: 371-378.
81" "

Suski, C. D., Svec, J. H., Ludden, J. B. et al. (2003). The effect of catch-and-release angling on the parental care behavior off male smallmouth bass. Transactions of the American Fisheries Society 132: 210-218. Syrjnen, J. & Valkeajrvi, P. (2010). Gillnet fishing drives lake-migrating brown trout to near extinction in the Lake Pijnne region, Finland. Fisheries Management and Ecology 17: 199-208. Talmage, P. J. & Staples, D. F. (2011). Mortality of walleyes angled from the deep waters of Rainy Lake, Minnesota. North American Journal of Fisheries Management 31: 826-831. Taylor, R. G., Whittington, J. A. & Haymans, D. E. (2001). Catch-and-release mortality rates of common snook in Florida. North American Journal of Fisheries Management 21: 70-75. Thorley, J. L., Youngson, A. F. & Laughton, R. (2007). Seasonal variation in rod recaptures rates indicates differential exploitation of Atlantic salmon, Salmo salar, stock components. Fisheries Management and Ecology 14: 191-198. Thorstad, E. B., Hay, C. J., Nsje, T. F. et al. (2004). Effects of catch-and-release angling on large cichlids in the subtropical Zambezi River. Fisheries Research 69: 141-144. Thorstad, E. B., Nsje, T, F. & Leinan, I. (2007). Long-term effects of catch-and-release angling on ascending Atlantic salmon during different stages of spawning migration. Fisheries Research 85: 316-320. Thorstad, E. B., Nsje, T. F., Fiske, B. et al. (2003). Effects of hook and release on Atlantic salmon in the River Alta, northern Norway. Fisheries Research 60: 293-307. Tsuboi, J. & Morita, K. (2004). Selectivity effects on wild white-spotted charr (Salvelinus leucomaenis) during a catch and release fishery. Fisheries Research 69: 229-238.

82" "

Tsuboi, J., Morita, K. & Ikeda, H. (2006). Fate of deep-hooked white-spotted charr after cutting the line in a catch-and-release fishery. Fisheries Research 79: 226-230. Tsuboi, J., Morita, K. & Matsuishi, T. (2002). Effects of catch-and-release angling on growth, survival and catchability of white-spotted charr Salvelinus leucomaenis in wild streams. Nippon Suisan Gakkaishi 68: 180-185. Tufts, B. L., Tang, Y., Tufts, K. et al. (1991). Exhaustive exercise in wild Atlantic salmon (Salmo salar): Acid-base regulation and blood gas transport. Canadian Journal of Fisheries and Aquatic Sciences 48: 868-874. van der Walt, B., Faragher, R. A. & Lowry, M. B. (2005). Hooking mortality of released silver perch (Bidyanus bidyanus) after capture by hook-and-line fishing in New South Wales, Australia. Asian Fisheries Science 18: 205-216. Vecchio, J. L. & Wenner, C. A. (2007). Catch-and-release mortality in subadult and adult red drum captured with popular fishing hook types. North American Journal of Fisheries Management 27: 891-899. Veiga, P., Conalves, J. M. S. & Erzini, K. (2011). Short-term hooking mortality of three marine fish species (Sparidae) caught by recreational angling in south Portugal. Fisheries Research 108: 58-64. Vincent-Lang, D., Alexandersdottir, M. & McBride, D. (1993). Mortality of coho salmon caught and released using sport tackle in the Little Susitna River, Alaska. Fisheries Research 15: 339-356. Walter, J. F., Orbesen, E. S., Liese, C. et al. (2012). Can circle hooks improve western Atlantic sailfish, Istiophorus platypterus populations? Bulletin of Marine Science 88: 755770.
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Waters, J. R. & Huntsman, G. R. (1986). Incorporating mortality from catch and release into yield-per-recruit analyses of minimum-size limits. North American Journal of Fisheries Management 6: 463-471. Weathers, K. C. & Newman, M. J. (1997). Effects of organizational procedures on mortality of largemouth bass during summer tournaments. North American Journal of Fisheries Management 17: 131-135. Webb, J. H. & Youngson, A. F. (1992). Reared Atlantic salmon, Salmo salar L., in the catches of a salmon fishery on the western coast of Scotland. Aquaculture and Fisheries Management 23: 393-397. Wedemeyer, G. A. & Wydoski, R. S. (2008). Physiological response of some ecologically important freshwater salmonids to catch-and-release fishing. North American Journal of Fisheries Management 28: 1587-1596. White, A. J., Schreer, J. F. & Cooke, S. J. (2008). Behavioural and physiological responses of the congeneric largemouth (Micropterus salmoides) and smallmouth bass (M. dolomieu) to various exercise and air exposure durations. Fisheries Research 89: 9-16. Whoriskey, F. G., Prusov, S. & Crabbe, S. (2000). Evaluation of the effects of catch-andrelease angling on the Atlantic salmon (Salmo salar) of the Ponoi River, Kola Peninsula, Russian Federation. Ecology of Freshwater Fish 9: 118-125. Whoriskey, S., Arauz, R. & Baum, J. K. (2011). Potential impacts of emerging mahi-mahi fisheries on sea turtle and elasmobranch bycatch species. Biological Conservation 144: 18411849. Wilde, G. (2009). Does venting promote survival of released fish? Fisheries 34: 20-28.

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Wilde, G. R. & Paulson, L. J. (2003). Movement and dispersal of tournament-caught largemouth bass in Lake Mead, Arizona-Nevada. Journal of Freshwater Ecology 18: 339342. Wilde, G. R. & Sawynok, W. (2009). Effect of hook removal on recapture rates of 27 species of angler-caught fish in Australia. Transactions of the American Fisheries Society 138: 692697. Wilde, G. R. (2003). Dispersal of tournament-caught black bass. Fisheries 28: 10-17. Wilde, G. R., Shavlik, C. E. & Pope, K. L. (2002). Initial mortality of black bass in BASS fishing tournaments. North American Journal of Fisheries Management 22: 950-954. Wilson, J. A. & Diaz, G. A. (2012). An overview of circle hook use and management measures in United States marine fisheries. Bulletin of Marine Science 88: 771-788. Wilson, R. R. & Burns, K. M. (1996). Potential survival of released groupers caught deeper than 40 m based on shipboard and in-situ observations, and tag-recapture data. Bulletin of Marine Science 58: 234-247. Yergey, M. E., Grotheus, T. M., Able, K. W. et al. (2012). Evaluating discard mortality of summer flounder (Paralichthys dentatus) in the commercial trawl fishery: developing acoustic telemetry techniques. Fisheries Research 115-116: 72-81. Yokota, K., Kiyota, M. & Minami, H. (2006). Shark catch in pelagic longline fishery: Comparison of circle and tuna hooks. Fisheries Research 81: 337-341. Zischke, M. T., Griffiths, S. P. & Tibbetts, I. R. (2012). Catch and effort from a specialised recreational pelagic sport fishery off eastern Australia, Fisheries Research 127-128: 61-72.

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