Bull Mar Sci. 93(0):000–000. 2017
Bulletin of Marine Science
© 2017 Rosenstiel School of Marine & Atmospheric Science of
the University of Miami
Fishing in the dark: the science and management
of recreational sheries at night
Steven J Cooke 1 *
Robert J Lennox 1
Shannon D Bower 1
Andrij Z Horodysky 2
Melissa K Treml 3
Erin Stoddard 4
Lisa A Donaldson 1
Andy J Danylchuk 5
ABSTRAC T.—Recreational ﬁshing is a popular activity
around the globe, generating billions of dollars in economic
beneﬁt based on ﬁsheries in marine and inland waters. In
most developed countries, recreational ﬁsheries are managed
to achieve diverse objectives and ensure that such ﬁsheries
are sustainable. While many anglers ﬁsh during daylight
hours, some target ﬁsh species during the night. Indeed,
sensory physiology of some species makes them vulnerable to
capture at night, while being more diﬃcult to capture during
the day. However, night creates a number of challenges for
recreational ﬁsheries assessment and management. In some
jurisdictions, ﬁshing is prohibited at night (through both
eﬀort and harvest controls) or there are speciﬁc restrictions
placed on night ﬁsheries (e.g., no use of artiﬁcial lights). Here,
we summarize the science and management of recreational
ﬁsheries at night covering both inland and marine realms.
In doing so, we also provide a review of diﬀerent angling
regulations speciﬁc to night ﬁsheries across the globe, as
well as the basis for those regulations. We discuss the extent
to which there is both need and opportunity to actively
manage anglers who are targeting ﬁsh at night and how this
diﬀers from ﬁsheries that occur during lighted periods. We
provide two case studies, one for white sturgeon (Acipenser
transmontanus Richardson, 1836) and one for walleye
[Sander vitreus (Mitch ill, 1818)], for which nig httime closures
have been used as a ﬁsheries management tool to control
eﬀort and harvest (illegal harvest in the case of the sturgeon
case study). Based on the synthesis, we conclude that natural
resource management agencies should decide if and how they
need to manage recreational ﬁsheries at night, recognizing
the practical challenges (e.g., compliance monitoring, stock
assessment) with doing so in the dark.
1 Fish Ecology and Conservation
Department of Biology and
Institute of Environmental
Science, Carleton University, 1125
Colonel By Dr., Ottawa, Ontario,
K1S 5B6, Canada.
2 Department of Marine
and Environmental Science,
Hampton University, Hampton,
3 Minnesota Department of
Natural Resources, St. Paul,
4 Fish and Wildlife Section,
Governme nt of British Columbia ,
Surrey, British Columbia, V3R
5 Department of Environmental
Conservation, University of
Amherst, Massachusetts 01003.
* Corresponding author email:
Section Editor: John F Walter, III
Date Submitted: 4 January, 2016.
Date Accepted: 19 August, 2016.
Available Online: 28 November, 2016.
Fas t Track
Bulletin of Marine Science. Vol 93, No 0. 20172
Recreational ﬁshing is deﬁned as ﬁshing of aquatic animals (mainly ﬁsh) that do
not constitute the individual’s primary resource to meet basic nutritional needs and
are generally not sold or otherwise traded on export, domestic, or black markets (UN
FAO 2012). It is a popular activity around the globe, estimated to be practiced by ap-
proximately 10% of the global population (Arlinghaus and Cooke 2009, Arlinghaus
et al. 2015b). On an annual basis, as many as 40 billion ﬁshes may be captured by
recreational ﬁshers, of which more than half are released (Cooke and Cowx 2004).
Recreational ﬁshing yields numerous beneﬁts around the globe, not the least of which
is generation of tens of billions of dollars of direct and indirect economic activity
(Arlinghaus and Cooke 2009, Tufts et al. 2015). A variety of gear types can be used in
recreational ﬁsheries, but the dominant one is rod and reel (i.e., angling). Although
relative to commercial ﬁsheries, the eﬀects of recreational ﬁshing on global ﬁsh de-
cline and the environment are regarded as more benign (Cooke and Cowx 2006,
Lewin et al. 2006), there are certainly examples of ﬁsh population declines and even
collapse attributed to recreational ﬁshing (see Post et al. 2002). Increasingly, recre-
ational ﬁshing targets species or populations that are declining, which is creating a
number of management challenges (Coleman et al. 2004, Cooke et al. 2016).
Given the importance of recreational ﬁshing, it is not surprising that in many juris-
dictions, particularly in developed countries, governance structures exist to support
the sustainable management of recreational ﬁsheries. Typically underpinning such
management eﬀorts are science-based ﬁshery assessments. In developing countries
and emerging economies, science capacity is often lacking and governance struc-
tures (in terms of policy instruments) fail to provide natural resource agencies with
the tools and support needed to actively manage ﬁsheries. At the core of recreational
ﬁsheries management are traditional harvest control regulations such as bag limits
and size limits (Johnson and Martinez 1995). However, eﬀort controls are gaining
in popularity (e.g., protected areas, seasonal closures, Cox et al. 2003). Recreational
ﬁsheries management is often regarded as a partnership between government and
various stakeholder groups through formal or informal co-management agreements
(UN FAO 2012). With adequate regulations related to harvest and eﬀort control,
along with requisite habitat protection (see Lapointe et al. 2014), most recreational
ﬁsheries can be managed to achieve multiple beneﬁts.
Nighttime (and its associated darkness) is omnipresent around the globe and many
ﬁshes can certainly be captured during nocturnal periods, reﬂecting species-speciﬁc
diﬀerences in sensory physiology and feeding activity. Quantifying the number of
anglers who ﬁsh at night has a number of practical challenges (e.g., safety and logis-
tics of working on or near water at night). From an enforcement perspective, night
and its associated darkness can provide “cover” for those that intend to not comply
with regulations. From a science and management perspective, the vast majority of
staﬀ eﬀort is focused on daytime periods. Here, we provide the ﬁrst synthesis on the
science and management of recreational ﬁsheries at night. First, we describe ﬁsh-
ing at night from the perspective of a ﬁsh, exploring how species-speciﬁc sensory
physiology and biology contributes to vulnerability to capture. Next, we characterize
the state of night ﬁshing, identifying examples of speciﬁc tactics used to target ﬁsh
at night. en, we summarize the science and assessment of ﬁshing at night needed
to support ﬁsheries management. Finally, we explore strategies used to manage ﬁsh-
ing at night with a particular focus on policy compliance challenges using several
case studies where night-speciﬁc management regulations have been implemented.
Cooke et al.: Recreational shing in the dark 3
With increasing recreational ﬁshing eﬀort on a global basis, it is our hope that our
synthesis will provide managers with information to achieve recreational ﬁsheries
sustainability by managing ﬁsheries around the clock, not just during daylight. We
are global in our approach, covering marine and inland recreational ﬁsheries, but
limit our review to recreational angling (i.e., ﬁshing via hook and line). We recognize
that depending on latitude (e.g., polar regions) and season, night and darkness are
not always aligned, but for the purpose of our synthesis, we take night to imply dark-
ness at least in a relative sense compared to daytime periods.
F N F’ P
Predatory gameﬁshes exhibit species-speciﬁc diurnal rhythms in both sensory
physiology and feeding activity (Reebs 2002). Physiological adaptations of gameﬁsh-
es to low light levels, including overcast conditions, crepuscular periods, and night,
may explain why catches of many species peak at these times. Midday clouds can
drop aquatic light intensities by one to two orders of magnitude; during crepuscular
periods, intensity can change roughly tenfold every 10 min (Fig. 1A). Natural noc-
turnal light levels are a million to a billion times dimmer than those at high noon,
depending on moon phase (Warrant 1999, Johnsen 2012). Many predatory ﬁshes
forage visually, using rod photoreceptors during scotopic (dim/dark) conditions to
increase sensitivity and form monochromatic images, and cone photoreceptors un-
der photopic (bright) conditions to form high-resolution, contrasting images of prey.
Nocturnal foragers have large eyes, a high rod:cone ratio, slow vision, poor acuity,
prevalent tapeta lucida, high luminous sensitivity (Warrant 1999, Horodysky et al.
2008), and/or may have enhancements in chemosensory and mechanosensory sys-
tems for food search (Pohlmann et al. 2004). Examples of such ﬁshes include walleye
[Sander vitreus (Mitchill, 1818)], adult brown trout (Salmo trutta Linnaeus, 1758) and
bull trout [Salvelinus conﬂuentus (Suckley, 1859)], channel catﬁsh [Ictalurus puncta-
tus (Raﬁnesque, 1818)], weakﬁsh [Cynoscion regalis (Bloch and Schneider, 1801)], and
swordﬁsh (Xiphias gladius Linnaeus, 1758). By contrast, predators of daylight hours
have smaller eyes, higher cone:rod ratios, faster vision, better acuity, wider chromatic
sensitivity, and moderate luminous sensitivity (Horodysky et al. 2008). Examples of
these ﬁshes include boneﬁsh (Albula spp.), striped bass (Morone saxatilis ( Wa lbaum,
1792)], yellow perch [Perca ﬂavescens (Mitchill, 1814)], and northern pike (Esox lucius
Linnaeus, 1758). Of course, the latter species can still be captured under low light
Within a species, luminous sensitivity can be extended under falling light levels
as permitted by physical and physiological bounds by widening pupils, increasing
temporal and spatial summation of ganglion cells, and/or via circadian retinomo-
tor movements that withdraw the pigment epithelium protecting rod photoreceptors
from daylight (Fig. 1B, Warrant 1999). However, because of unavoidable tradeoﬀs,
physiological responses that increase sensitivity come at the cost of slower temporal
resolution, reductions in acuity due to reduced spatial summation, and constrained
chromatic sensitivity (Horodysky et al. 2010). Under natural low-light conditions,
diurnal predatory ﬁshes may be forced to cease visual foraging when image forma-
tion is impaired, and turn increasingly to encounter-based chemosensory, acous-
tic, and/or mechanoreceptive cues to locate and track prey as per species-speciﬁc
adaptations and abilities (Hara and Zielinski 2006). Some dim-light and nocturnal
Bulletin of Marine Science. Vol 93, No 0. 20174
foragers such as burbot [Lota lota (Linnaeus, 1758)] and ﬂathead catﬁsh [Pylodictis
olivaris (Raﬁnesque, 1818)] may cue predominantly on chemosensory cues (Døving
and Gemne 1965, Hinkens and Cochran 1988, Daugherty and Sutton 2005), which
are dependent on water ﬂow, and may be less aﬀected by aquatic photodynamism.
Crepuscular periods are brief photodynamic windows enveloping the night in
which the solar elevation is low, light intensity and spectra change rapidly, and many
Figure 1. Mechanistic examination of light conditions, ecophysiological processes, and behav-
ioral strategies during crepuscular periods. (A) Changes in light intensity during dawn and dusk.
(B) Mechanistic pathways (blue ar rows) of changes in light intensity at dusk on physiology and
behavior, with feedbacks (dashed gray arrows). (C) Effects of low solar elevation and changing
light intensities characteristic of crepuscular periods on prey visual contrast and behavioral for-
aging strategies of a predator (following Johnsen 2003, Johnsen and Sosik 2003).
Cooke et al.: Recreational shing in the dark 5
prey countershading and camouﬂage strategies can be counteracted by predators
and exploited by anglers (Fig. 1C) (Johnsen 2003, 2012). It is thus not surprising that
much ﬁshing eﬀort is exerted, and angling success experienced, at these times. Light
intensity cha nges by roughly 2 log units between 0° and 5° of solar elevation, as the su n
is near the horizon (Johnsen 2003, 2012). Below the horizon, light changes by 104–107
units from the time of ﬁrst/last light (−18°) to sunrise/sunset (i.e., 0°), and is intense-
ly dominated by shorter (UV and blue) wavelengths (Warrant and Johnsen 2013).
Once the sun is >18° below the horizon (i.e., true night), the blue twilight is replaced
by dimmer and redder starlight, airglow, and zodiacal light (new moon), by a dim
spectrum that resembles slightly red-shifted daylight in spectral composition (full
moon), or a combination (intermediate moon phases) (Warrant and Johnsen 2013).
At low solar elevation, the rising or setting sun can illuminate the lateral ﬂanks of
animals to a much higher degree than the overhead noon sun. Viewing backgrounds
away from the low-elevation sun are dark/shaded, whereas those into the sun are
bright (Johnsen and Sosik 2003). When viewed away from the sun, dark-ﬂanked prey
become slightly less cryptic than at noon, but the ﬂanks of mirrored, light-colored,
countershaded prey contrast strongly against the dark background (Fig. 1C; Johnsen
2003, Johnsen and Sosik 2003). Conversely, when viewed into the plane of the low-
elevation sun, dark-ﬂanked and countershaded prey contrast strongly against the
bright background, and mirrored and light-colored prey experience better crypsis
(Fig. 1C; Johnsen 2003). Mirrored organisms can never be completely cryptic when
backlit by the sun because this requires the physical impossibility of a reﬂectance >1
(Johnsen and Sosik 2003). In fact, both mirrored and light-ﬂanked prey block sun-
light, leaving silhouettes that are darker than the veiling spacelight.
During crepuscular periods, predators can increase the visibility of prey by search-
ing in circular patterns relative to the low solar elevation, creating background opti-
cal mismatches (Fig. 1C). Predators can then drive prey toward the surface, where
the asymmetry of the aquatic light ﬁeld will be most pronounced (Johnsen 2003).
Interestingly, countershading coloration patterns that are eﬀective at noon can leave
prey highly conspicuous at dawn and dusk, as either their dorsum or ventrum will
contrast strongly against the optical background into or away from the sun. Finally,
predators transition between circling and encounter-rate strategies when light be-
comes a factor limiting image formation (early in dawn or late into dusk). Once all
sunlight is extirpated, the natural conditions of true night can impede schooling
and visual foraging in many ﬁshes, depending on moon phase (Helfman 1993, Fréon
et al. 1996). Diurnal game ﬁshes such as largemouth bass [Micropterus salmoides
(Lacépède, 1802)] may be able to visually forage under a full moon’s light intensity,
but not under starlight typical of a new moon (McMahon and Holanov 1995).
Objects viewed from below block downwelling light from the night sky and may
be silhouetted against the surface (Johnsen 2003), thus anglers ﬁshing under waxing,
waning, and new moons often opt for large, dark, water-displacing lures to attract
ﬁsh to the silhouette, sound, and vibration. Others select odoriferous baits that gen-
erate a chemical plume to stimulate olfactory and gustatory systems. Chemical light
sticks, where legal, may also be added to bait in an attempt to enhance catchability. In
commercial ﬁsheries, Hazin et al. (2005) compared the catch-per-unit eﬀort (CPUE)
of squid-baited hook baskets illuminated by light sticks to those without light sticks
for catching swordﬁsh with an artisanal longline vessel ﬁshing at 30–150 m depth.
Hazin et al. (2005) found that using a light stick on alternating hooks (i.e., on three
Bulletin of Marine Science. Vol 93, No 0. 20176
out of six hooks) signiﬁcantly increased CPUE relative to using no light stick or a
light stick on every hook. Similar evaluations of light sticks in recreational ﬁsheries
For a variety of sensory and environmental reasons, some species of ﬁshes be-
come active at night (Emery 1973, Munz and McFarland 1977; Fig. 2). ere is diel
variation in catchability with sampling gears (e.g., electroﬁshing, netting; Pope and
Willis 1996); however, diel variation in CPUE with recreational ﬁshing gear has not
been well studied. Yet, some anglers like to go ﬁshing in the evenings or early in the
morning before daybreak, suggesting that ﬁshing during crepuscular periods and at
night is productive. In some specialized ﬁsheries, anglers will speciﬁcally wait for
nightfall to go ﬁshing. Although the ﬁshing can be rewarding, ﬁshing at night is
logistically challenging depending on the target species, particularly due to visibility
and navigational issues. However, urbanization has led to the installation of artiﬁcial
lights along coasts and embayments, which shine into the water (Nightingale et al.
2006; Fig. 3). Such lighting attracts baitﬁsh (Ben-Yami 1976, 1988) and insects, which
in turn draws predatory ﬁshes close to shore (Browder 2012). At night, anglers can
target these artiﬁcially lit areas. For example, anglers often target common snook
[Centropomus undecimalis (Bloch, 1792)] that follow baitﬁsh into the shallow, illu-
minated areas. Some ﬁshing guides explicitly mention “ﬁshing under lighted docks”
in their advertising materials emphasizing how artiﬁcial lighting (in this case, light
pollution) can be exploited by anglers.
Sometimes ﬁshing is best without light, especially when target species have evolved
to feed in darkness and/or are photophobic. Nightingale et al. (2006) described how
weakﬁsh forage only below 0.5 lux and anglers avoid ﬁshing during the full moon
because their targets are inactive. For other species, feeding/vulnerability can be en-
hanced during full moon phases when visual predators have more light with which to
perceive potential food items (Fig 2). However, ﬁsh feed using many diﬀerent senses
(see above, Pavlov and Kasumyan 1990) meaning that visual cues are not entirely
necessary for catching ﬁsh. New et al. (2001) ablated the eyes of muskellunge (Esox
masquinongy Mitchill, 1824) and found that they used somatosensory cues to inform
their angles of prey attack. Benthic feeding species such as catﬁshes (Siluriformes)
feed at night using olfactory and gustatory cues, sweeping the benthos with external
tastebuds (such as on barbels) to inform feeding (Atema 1971). Anglers can ﬁsh at
night for these benthivores with passively ﬁshed set lines by sinking baited hooks to
the benthos and waiting for ﬁsh to ingest the bait, generally hooking themselves (of-
ten in the throat or stomach because the hook is ingested with the bait). Fishing with
set lines is illegal in some jurisdictions, particularly because set lines can increase the
probability of deep hooking and mortality of ﬁsh that are captured. To indicate when
a ﬁsh strikes, tools such as bells or alarms can be ﬁxed to the rod. Electronic bite
alarms are marketed to carp (Cyprinus carpio Linnaeus, 1758) anglers that ﬁsh from
shore at night so that when they fall asleep with their bait set, the battery-powered
alarm will sound to indicate a strike. Setting baits under ﬂoats or bobbers that are
reﬂective or glow-in-the dark can also increase strike detection in the dark (Johnson
2013). Some manufacturers produce ﬁshing rods that have tips intended to glow at
night (often in the presence of black light) to facilitate strike detection. e angling
Cooke et al.: Recreational shing in the dark 7
industry (including the outdoor media) are acutely aware of the market for night ﬁsh-
ing with many books, videos, television segments and magazine articles on the topic.
ere are also a number of charters advertised as being speciﬁc to ﬁshing at night
(e.g., ﬁshing oﬀ head-boats oﬀ the shores of North Carolina and South Carolina for
deepwater reef ﬁsh; ﬁshing for swordﬁsh oﬀ the Atlantic coast of Florida).
One of the oddest night-speciﬁc ﬁsheries issues that emerges is for specialized carp
angling where it is common to place ﬁsh captured at night in “carp sacks” to hold the
ﬁsh until the daylight when photographic opportunities are better. However, during
Figure 2. Night shing under natural and anthropogenically-inuenced conditions. Human ar-
ticial lighting can increase nocturnal light intensities to within 104 units of high noon, leading
to changes in sh aggregation, available sensory modalities, foraging strategies, and catchabil-
ity (q). Management strategies for nat ural and anthropogenically-inuenced nocturnal sheries
should consider spatiotemporal proper ties, terminal gears, and size and bag limits. SS = species
specic. Senses are: audition (A), gustation (G), mechanoreception (M), olfaction (O), and vision
Bulletin of Marine Science. Vol 93, No 0. 20178
retention in the carp sack, the ﬁsh become quite vigorous so it is necessary to inten-
tionally air expose the carp (often by hanging them from a tree in the carp sack) to
induce some level of physiological exhaustion so that the ﬁsh can be held for photos.
Although this practice may seem to be one that would be deleterious to ﬁsh, research
on the topic suggests that carp are extremely robust to both carp sack retention and
prolonged air exposure such that there is negligible mortality and rapid recovery
from the associated stress (Rapp et al. 2012).
Figure 3. Two cate gories of anth ropoge nic arti c ia l light, wit h inuences on aqu at ic habit at s. (A)
general illumination of the urban night sky can increase aquatic light up to 10,000 times brighter
than the new moon, enabling visual foraging by piscivores such as cutthroat trout (Mazur and
Beauchamp 2006). (B) Point source illumination typical of docks, piers, bridges, marinas, and
waterfront restaurants. Light is far more limited and concentrated by point sources, increasing
asymmetries of prey contrast under the light and predator crypsis in the shadow lines. Both arti-
cial light conditions increase nocturnal foraging and catchability of predators that would not be
able to forage visually under natural conditions.
Cooke et al.: Recreational shing in the dark 9
N S A
Where ﬁsheries management exists globally, the general governing principle is
that the management strategies follow a science-based approach. Diﬀerences among
target species and the behaviors anglers employ to catch ﬁsh vary widely among ﬁsh-
eries, such that research to identify species-speciﬁc impacts due to recreational ﬁsh-
ing have been recommended, particularly for catch-and-release (C&R) ﬁshing (e.g.,
Cooke and Suski 2005). Similarly, it cannot be assumed that conditions that aﬀect
daylight ﬁshing apply broadly to night ﬁshing. Yet, night ﬁshing is often explicitly
excluded from ﬁsheries assessment surveys (e.g., Brouwer et al. 1997, Smallwood et
al. 2006, Zeller et al. 2007), including the Marine Recreational Information Program
(MRIP) of the US National Marine Fisheries Service that did not include night sam-
pling in their surveys until 2013. is lack of inclusion may reﬂect the position that
night ﬁshing is not widely popular. In a study of the Majorca Island recreational
ﬁsheries, nighttime anglers represented only 2.4% of ﬁshing activity (Morales-Nin et
al. 2005), yet in a survey of angling behaviors in the South African shore ﬁshery, 54%
of anglers interviewed indicated that they participated in night ﬁshing, and 34% of
their ﬁshing activity took place at night (Brouwer et al. 1997), indicating that popu-
larity of the practice is globally variable. erefore, the dearth of available literature
on night angling survey results speaks to the presence of a knowledge gap, and likely
speaks to the challenges in conducting such surveys, rather than to a lack of interest
or need. Researchers may look to studies documenting impacts of devices and behav-
iors commonly used to target ﬁsh at night to inform research priorities, but it must
be determined whether these results apply to ﬁshing at night.
As discussed in the earlier sections, ﬁsh biology and behavior is inﬂuenced by diel
patterns. Diel migrations, whether from benthic to littoral zones, from oﬀshore to
inshore regions, or vertical migrations in the water column, can result in diﬀerences
in species composition between day and night (Bassett and Montgomery 2011). is
suggests that there potentially may be signiﬁcant diﬀerences in expected outcomes
of recreational ﬁshing behaviors. For example, the increased presence of predators
in a nocturnal community may result in an increase in post-release predation after a
C&R event because predation rates can increase at night (Danilowicz and Sale 1999).
In a study of recreational bycatch aﬀecting the critically endangered gray nurse shark
(Carcharias taurus Raﬁnesque, 1810) in Australia, no diel patterns in hooking were
found, though authors noted that C. taurus was the only predator in the area taking
bait at night (Robbins et al. 2013), a ﬁnding that also raises the potential implication
of diel patterns in recreational bycatch. Tropical mangrove estuaries are predomi-
nantly comprised of nocturnal ﬁsh (Ley and Halliday 2007), and a third of ﬁsh fauna
in any ecosystem may be nocturnal (Helfman 1978, cited in Bassett and Montgomery
2011), supporting the idea that conditions for night ﬁshing may be diﬀerent, and
species assemblages at night may diﬀer. Further, diel variations in catchability have
been noted for some species (Benoít and Swain 2003), which could potentially impact
recommendations for catch limits.
Night ﬁshing may result in diﬀerent species-speciﬁc impacts due to changes in key
angling variables, such as extended handling times and air exposure as a result of
reduced visibility in darkness. Rates of deep hooking, injury, and post-release mor-
tality may also be tied to reduced visibility as anglers may be slower to register bites,
particularly if using “passive” techniques such as bobbers (Lennox et al. 2015) or set
Bulletin of Marine Science. Vol 93, No 0. 201710
lines. Moreover, handling and unhooking times can increase at night as a result of
poor visibility. Diﬀerences in angling methods between day and night could result in
diﬀerent hooking mortality rates for released ﬁsh that are independent of diﬀerence
in handling time due to poor visibility. Anecdotally, night ﬁshing involves more use
of artiﬁcial lights and scent-based attractants than day ﬁshing. ere is much vari-
ability among species in response to light (i.e., diﬀerences among and within species
according to life stage) and there is a high degree of plasticity in these responses
(Nightingale et al. 2006), which could inﬂuence the extent to which anglers using
light can directly or indirectly impact populations. Further study of recreational
ﬁshing at night can inform regulations for night ﬁshing; for example, the use of circle
hooks may be warranted to reduce deep hooking associated with using passive ﬁsh-
ing techniques at night (Cooke and Suski 2004).
Diﬀerences in angling communities and angler behavior at night should be anoth-
er integral component of night surveys, including attempts to understand motivation
and external relationships with other users. For example, Arlinghaus (2005) noted
that there might be conﬂict among nighttime recreational ﬁshers in areas where
these activities overlap with some types of commercial ﬁshing (e.g., those that use
fyke nets). Diﬀerences may also exist within the angling community; in the Maldives,
recreational ﬁshing is not popular among locals, focusing mainly on tourists, yet
locals do participate in recreational night ﬁshing (FAO Fisheries and Aquaculture
Department 2009), which suggests that angling communities may exhibit diel varia-
tion in composition in some areas. is conclusion is supported by the suggestion to
relax the ban on night ﬁshing in urban Berlin, Germany, as a way to promote urban
ﬁshing experiences, because night ﬁshing is more popular with urban than rural
anglers (Arlinghaus et al. 2008). To some extent, this pattern may be driven by the
prevalence of anthropogenic illumination.
ere are challenges inherent in conducting surveys of night ﬁsheries, including
considerations of safety and unintended contributions of safety and research gear
to study outcomes. Safety considerations, both perceived and actual, have been sug-
gested as one of the driving factors in a lack of night studies (Smallwood et al. 2011).
In addition to reduced visibility constraining safe operation of equipment, increased
activity of land- or water-based predators (e.g., crocodiles) at night is also a con-
cern in some areas. e use of surveys and interviews conducted during the day can
be used to gather information regarding angler behaviors and perspectives, and for
some ﬁsheries, creel surveys can safely be performed at night. Roving creel surveys
were used at night in a study of a prawn ﬁshery in New South Wales, Australia, where
researchers were able to identify prawn ﬁshers because of artiﬁcial light bobbers af-
ﬁxed to the scoop nets they used (Reid and Montgomery 2005).
New technologies, such as the use of remote and infrared cameras, may be help-
ful in alleviating some of the safety concerns associated with night surveys. Remote
cameras using infrared to observe shore-based angling activities at night found that
camera placement was integral to ensuring that the number of people in a party could
be identiﬁed, and to identifying which activity types were taking place (Smallwood
et al. 2011). Conversely, a study conducted to identify night assemblages found that
use of infrared light (as opposed to white light) resulted in improved surveys because
infrared light allowed researchers to distinguish among individuals more eﬀectively
(Harvey et al. 2012). In a study comparing underwater assessment techniques us-
ing bait and infrared video to conduct underwater surveys, the authors found that
Cooke et al.: Recreational shing in the dark 11
olfactory-driven species arrived at video sites sooner, whereas non-olfactory driven
species were captured more readily in traditional underwater survey techniques (us-
ing scuba and/or snorkel; Bassett and Montgomery 2011). e authors concluded
that the type of survey will yield diﬀerent species-speciﬁc encounter and catch-
ability rates depending upon the sensory capabilities of the organisms (Bassett and
In addition to new technologies, more traditional methods may prove suitable for
night surveys, though diel diﬀerences in eﬃciency should be tested. When electro-
ﬁshing for smallmouth bass (Micropterus dolomieu Lacépède, 1802), Paragamian
(1989) suggested ﬁshing at night would improve gear eﬃciency and catch numbers,
because CPUE was higher. Questions regarding night ﬁshing activities might also
represent an opportunity to invest more fully in sources of local knowledge for as-
sessment (Hamilton et al. 2012). Concerns about using local knowledge include po-
tential for recollection bias, that such information has been devalued as being purely
anecdotal, and that integration into formal assessment methodologies is challenging
(Johannes and Neis 2007), but these concerns can be addressed by approaching the
collection of local knowledge in a scientiﬁc and veriﬁable way (e.g., see Arlinghaus
and Krause 2013). With such concerns accounted for, local ﬁsher knowledge can help
to close gaps in scientiﬁc understanding (Johannes and Neis 2007), and can be useful
in identifying likely research priorities and safety concerns.
Fisheries management activities can often be categorized as managing habitat,
managing people, and managing ﬁsh(es) (Krueger and Decker 1999, Arlinghaus et al.
2015a). Here we brieﬂy discuss the relevance of night to those three elements of rec-
reational ﬁsheries management. We also provide two recent high-proﬁle case studies
that involved regulating recreational angling activities for white sturgeon (Acipenser
transmontanus Richardson, 1836) and walleye [Sander vitreus (Mitchil l, 1818)].
Managing people is one of the more common recreational ﬁsheries management
strategies as it relates to elements of angler access, eﬀort, and harvest. Questions re-
garding diel diﬀerences in angler behavior can inform management decisions related
not only to outcomes for ﬁshes, but issues of compliance, enforcement, and even
promoting the practice of angling. For example, diﬀerences in compliance with ﬁsh-
ing regulations among night anglers could be a factor in informing the need for more
enforcement at diﬀerent times of day. Enforcement and compliance monitoring is in-
herently more diﬃcult (and dangerous) at night. Of course there are developments in
night vision goggles and aircraft or drone-based night imaging [e.g., forward looking
infra-red (FLIR) thermal imaging] that do provide enforcement staﬀ with some tools
for peering into the dark. Motivations for angling may also diﬀer at night, impact-
ing which management or enforcement strategies are likely to be successful. Anglers
who prefer to ﬁsh at night have expressed a desire to avoid increasing boat traﬃc,
warm temperatures, and to increase catch rates that may decrease in times when
ﬁsh are subjected to higher amounts of angling pressure (Quinn 2014). Some anglers
have even indicated preferences related to the phases of the moon, believing catch-
ability of their target species to be inﬂuenced by moonlight (Quinn 2014).
Regulations surrounding night ﬁshing are also variable; for example, the activity
is permitted in some areas of Portugal but prohibited in others such as the Parque
Bulletin of Marine Science. Vol 93, No 0. 201712
Natural do Sudoeste Alentejano e Costa Vicentina (Veiga et al. 2010). Night ﬁshing is
banned entirely in Greece, but is widely permitted in Cyprus, where licenses are only
required if ﬁshers intend to spearﬁsh at night (Pawson et al. 2008). In the Back Bay
National Wildlife Refuge (and indeed in all such refuges) in the Virginia, USA, night
ﬁshing activities were banned (see USFWS 2009). However, local angling groups lob-
bied successfully for opening limited night ﬁshing opportunities for striped bass (M.
saxatilis). A special lisence was required to fund the additional staﬀ time (for as-
sessment, management, and enforcement) to ensure that the ﬁshery was properly
regulated and monitored. A practical aspect of any eﬀorts to limit nighttime ﬁshing
involves deﬁning “nighttime” in a manner that is enforceable. Typically, nighttime
periods are identiﬁed relative to “published” sunrise and sunset periods (e.g., a clo-
sure from dusk until dawn starting 1 hr after sunset until 1 hr before sunrise). Other
common regulations relevant to night involve placing restrictions on speciﬁc gears.
For example, use of artiﬁcial lights (for ﬁsh attraction) are prohibited in many juris-
dictions. Also typically restricted are lures/baits that contain a light source, but lures
that “glow” (e.g., using glowing paint) tend to be allowed.
M C S: L F R S N F
C.—e Fraser River is a large river system in British Columbia (BC), Canada,
that originates near the Alberta border and drains a signiﬁcant portion of the prov-
ince. e lower Fraser River comprises the >180 km section from its mouth upstream
to Hells Gate in the Fraser Canyon, and supports large populations of all ﬁve species
of Paciﬁc salmon (Oncorhynchus spp.), steelhead [Oncorhynchus mykiss (Walbaum,
1792)], coastal cutthroat trout [Oncorhynchus clarki (Richardson, 1836)], bull trout,
and white sturgeon. e Lower Mainland, which includes the lower Fraser and BCs
largest metropolitan city (Vancouver), also supports BC’s largest human population.
e number of federal and provincial ﬁshery enforcement staﬀ is small relative to
the size of the human population, the extent of the ﬁsheries, and area to enforce. e
lower Fraser currently supports important cultural and multi-million dollar First
Nations (FN), commercial and recreational salmon ﬁsheries, and a multi-million
dollar recreational C&R white sturgeon ﬁshery.
e lower Fraser River is split into two jurisdictions: the river is designated as tidal
downstream of the CPR rail Bridge at Mission BC, and non-tidal upstream of the
bridge. Fisheries and Oceans Canada (DFO) manages and regulates all ﬁsheries in
tidal waters. FN, recreational and commercial Paciﬁc salmon ﬁsheries, in both tidal
and non-tidal waters, are also managed by DFO. Tidal and non-tidal nighttime an-
gling closures on the lower Fraser, and some tributaries, were implemented by DFO
to better manage the recreational salmon ﬁsheries, including the sockeye salmon
[Oncorhynchus nerka (Walbaum, 1792)] ﬁshery. e nighttime closure includes 1 hr
after sunset until 1 hr before sunrise, and was implemented in 2002.
e white sturgeon ﬁshery on the lower Fraser has been a C&R only ﬁshery since
the early 1990s, and has grown signiﬁcantly since the late-1990s. However, recent
studies (Nelson et al. 2014) indicated that the population was not growing as ex-
pected. e province has had concerns with respect to sturgeon night ﬁshing for
more than a decade because white sturgeon typically feed in the dark, making them
vulnerable to capture by angling at night. However, darkness is also the primary
time when poachers operate on the lower Fraser. Due to its high value for its ﬂesh
and its eggs, white sturgeon can bring large sums in the illegal trade market, and due
Cooke et al.: Recreational shing in the dark 13
to the size of the Lower Mainland human population, the potential market is large.
Poaching for sturgeon in the lower Fraser is conducted by angling, setline, or net.
Nighttime poaching is typically from shore by angling, but has also been conducted
by boat and with other methods. e province has been concerned about the han-
dling of white sturgeon in the C&R ﬁshery for more than a decade, with evidence
that there is risk of injury and mortality, especially when handling large adult ﬁsh
which can be much harder to handle in the dark. Further, it was brought to the at-
tention of provincial ﬁsheries staﬀ by enforcement during the consultation that a
sturgeon angler died in 2013 when a large sturgeon pulled him oﬀ a bridge onto an
abutment while he was ﬁshing in the dark.
In 2013, after several years of scoping the issue with stakeholders, the province de-
cided to initiate formal consultation on the potential implementation of a nighttime
closure to sturgeon ﬁshing on non-tidal waters of the lower Fraser River, lower Pitt
River, and Harrison River for the better management and protection of the species,
and for the safety of anglers. Federal and provincial enforcement staﬀ also recom-
mended this closure as being the only way to eﬀectively ensure that nighttime stur-
geon poaching could be enforced. Upon further consultation with legal, regulatory,
and stakeholder advisors, it was determined that it would be necessary to consult on
a total ﬁshing closure rather than a sturgeon only night ﬁshing closure. e extent of
the nighttime sturgeon ﬁshery at the time was unknown, but ﬁsheries and enforce-
ment staﬀ had observed that the majority of sturgeon angling occurs by boat during
daylight hours. Also, numerous nighttime sturgeon poaching enforcement cases had
recently proceeded to conviction, even with extensive education of the general public
and anglers of the importance of protecting white sturgeon.
A number of concerns were identiﬁed during stakeholder consultation on the pro-
posed lower Fraser nighttime closure, including concern that this would take “eyes
and ears” oﬀ the river to watch for poachers, that enforcement was inadequate to en-
sure compliance, and that the closure should pertain to both tidal and non-tidal wa-
ters. Provincial ﬁsheries staﬀ indicated that they expected DFO to mirror the change
for tidal waters. On April 1, 2015, the nighttime regulatory closure to all ﬁshing in
non-tidal waters of the lower Fraser, lower Pitt, and Harrison rivers came into eﬀect
with the timing of the closure extending from 1 hr after sunset to 1 hr before sunrise,
which is consistent with other recreational night closures, and the provincial hunt-
To date, DFO has not mirrored the nighttime ﬁshing closure for the tidal waters
of the lower Fraser River and Pitt River. Communication on social media appeared
to be limited as a consequence of the closure, and no recent communications with
regard to the closure have been received by provincial ﬁsheries staﬀ, which suggests
that anglers and angling guides have adjusted their activities around the closure.
Monitoring eﬀorts are underway to identify compliance with the regulatory change
and to assess the population-level responses.
M C S: M L L W N F
C.—Mille Lacs Lake is a 53,620 ha lake in north central Minnesota and is one
of Minnesota’s most important walleye (S. vitreus) ﬁsheries averaging 3 million hours
of angling pressure annually (Jensen 2013). Public interest in Mille Lacs management
dates back to the late 1940s with concerns about declining catch rates and increased
ﬁshing pressure. e ﬁrst documented concern over night ﬁshing occurred in 1961
Bulletin of Marine Science. Vol 93, No 0. 201714
after decreased angling success was noted the previous year. In response to numer-
ous stakeholder requests over several decades, the Minnesota Department of Natural
Resources (MNDNR) enacted a night ﬁshing ban in 1984 from 22:00 to 06:00 hrs for
the ﬁrst 4 wks of the open water season, which begins in early May. e next year, a
size restriction limiting harvest of walleye >508 mm (total length) was also imple-
mented. ese regulations remained unchanged through 1996. e primary intent
of the night closure was to redistribute harvest over the ﬁshing season rather than
reduce total harvest.
From 1984 to 1996, the median night harvest was about 15,000 kg (range 5000–
50,000 kg) comprising about 7% of the total annual angler harvest, including esti-
mated hooking mortality (Reeves and Bruesewitz 2007). In 1997, Mille Lacs became
a shared ﬁshery between state licensed anglers and Ojibwa (Chippewa) tribal ﬁshers.
From 1997 to 2013, the total allowable annual harvest of walleye was determined by
a ﬁxed exploitation policy using age-structured stock assessment model estimates
of total population biomass and averaged 200,000 kg (harvested ﬁsh and hooking
deaths). Tribal ﬁshers declared a ﬁxed quota each year, on average 25%–30% of the
total allowable harvest, with the remainder allocated to state recreational anglers.
e state recreational angling ﬁshery was managed using size-based regulations and
bag limits to remain within allocation. During this period, the spring night ﬁshing
ban remained in eﬀect while 10 diﬀerent size-based regulations and two diﬀerent
bag limits, along with mid-season changes to either more or less size restrictive regu-
lations, were implemented to control harvest.
Despite intensive management, the population did not increase (Venturelli et al.
2014). In 2014, a suite of alternative regulations was presented to stakeholders and
the open water season-long night ﬁshing closure was the most supported additional
restriction, followed by mandatory use of circle hooks and a more restrictive sea-
son-long night closure (18:00–06:00 hrs). What became evident is that night ﬁsh-
ing regulation is one management tool and it is unlikely to work in isolation unless
Table 1. Research needs specic to recreational sheries science and management at night.
• Identify sh habitat needs at night to ensure that critical habitats are protected and to inform
various enhancement and restoration activities
• Determine the extent to which light attracts different life-stages and species to determine the
relevance of regulations that ban light attraction and to exploit light to improve night assessment
activities (e.g., as is done with larval light traps)
• Identify survey designs that accurately quantify catch and effort over 24 hrs given that without
accurate quantication of catch and effort by day and night, management cannot be effective
• Examine the potential for selective effects of night vs day shing (are we catching the “same”
sh by day and night?)
• Characterize the “articial light food web” to understand how light pollution inuences key
sportssh and their prey (e.g., exigent need to study the sh–articial light–foraging relationship)
• Determine if sh handling and associated injury, stress, and mortality are elevated at night in the
context of catch-and-release shing
• Evaluate the extent to which post-release predation is mediated by night
• Conduct social science surveys to understand angler perspectives on night shing and associated
regulations (usually bans)
Cooke et al.: Recreational shing in the dark 15
combined with other tools (e.g., bag and slot limits and seasonal closures). Also rele-
vant is that all of these management tools rely on projections of anticipated outcomes
that do not necessarily occur due to interannual variability in catch rates and ﬁshery
conditions. Long-term monitoring to assess ﬁsh population responses to regulatory
changes as well a human dimensions work to evaluate stakeholder perspectives are
underway. What is clear is that night-speciﬁc regulations expand the toolbox for
It is evident from our review that recreational ﬁshing at night is popular, but not
universally so. e sensory and foraging ecologies of some species provide anglers
with unique opportunities to access ﬁsh during the night. To that end, the ﬁshing
industry has developed a variety of products intended to facilitate ﬁsh capture at
night. In general, less is known about the ecology and biology of ﬁshes at night, partly
driven by the inherent challenges of studying ﬁsh in darkness. Fisheries management
eﬀorts can speciﬁcally target the night, often in the form of temporal closures or gear
restrictions. When such management eﬀorts are enacted, there may be additional
resource needs and associated costs that need to be considered by natural resource
management agencies, particularly related to assessment and compliance monitor-
ing at night. e two case studies we presented exemplify high-proﬁle ﬁsheries for
which night time ﬁshery closures have been applied in an eﬀort to reduce directed
harvest (walleye), poaching (white sturgeon), and poor ﬁsh handling (both). e bio-
logical eﬀectiveness of these closures is still being assessed (e.g., did ﬁsh populations
respond as expected), but signiﬁcant eﬀort is also being devoted to understanding
stakeholder perspectives and compliance.
With eﬀorts by some anglers to be alone when ﬁshing, one might anticipate that
night ﬁshing may become more popular in the future as some anglers attempt to
avoid the masses that may ﬁsh during the day. We encourage the ﬁsheries manage-
ment community to think creatively about how nighttime recreational ﬁshing can
be promoted, but in a manner that is supported by eﬀective stock assessment and
management. ere are a number of outstanding research needs that were identiﬁed
throughout the review (see Table 1). Moving forward, we anticipate that the rec-
reational ﬁshing community may have more opportunities for ﬁshing in the dark
provided that management agencies can address the signiﬁcant assessment and com-
pliance monitoring challenges such that they are not “managing in the dark.”
Cooke is supported by NSERC and the Canada Research Chairs Program. Danylchuk is
supported by the National Institute of Food & Agriculture, US Department of Agriculture,
the Massachusetts Agricultural Experiment Station and Department of Environmental
Conservation. Horodysk y is supported by the NOAA Living Marine Resources Cooperative
Science Center and the NSF Educational Partnership in Climate Change and Sustainability.
We thank the Bulletin of Marine Science for organizing the Fish at Night conference and pro-
viding us the opportunit y to develop this paper.
Bulletin of Marine Science. Vol 93, No 0. 201716
Arlinghaus R. 2005. A conceptual framework to identify and understand conﬂicts in recre-
ational ﬁsheries systems, with implications for sustainable management. Aquat Res Cult
Dev. 1(2):145–174. http://dx.doi.org/10.1079/ARC200511
Arlinghaus R, Bork M, Fladung E. 2008. Understanding the heterogeneity of recreational an-
glers across an urban–rural gradient in a metropolitan area (Berlin, Germany), with im-
plications for ﬁsheries management. Fish Res. 92(1):53–62. http://dx.doi.org/10.1016/j.
Arlinghaus R, Cooke SJ. 2009. Recreational ﬁshing: socio-economic importance, conservation
issues and management challenges. In: Dickson B, Hutton J, Adams B, editors. Recreational
hunting, conservation and rural livelihoods: science and practice. Oxford: Blackwell
Publishing. p. 39–58.
Arlinghaus R, Krause J. 2013. Wisdom of the crowd and natural resource management. Trends
Ecol Evol. 28(1):8–11. http://dx.doi.org/10.1016/j.tree.2012.10.009
Arlinghaus R, Lorenzen K, Johnson BM, Cooke SJ, Cowx IG. 2015a. Managing freshwater ﬁsh-
eries: addressing habitat, people and ﬁsh. In: Craig J, editor. Freshwater ﬁsheries ecology.
UK: Blackwell Science. p. 557–579.
Arlinghaus R, Tillner R, Bork M. 2015b. Explaining participation rates in recreational ﬁshing
across industrialised countries. Fish Manag Ecol. 22(1):45–55. http://dx.doi.org/10.1111/
Atema J. 1971. Structures and functions of the sense of taste in the catﬁsh (Ictalurus natalis).
Brain Behav Evol. 4(4):273–294. http://dx.doi.org/10.1159/000125438
Bassett DK, Montgomery JC. 2011. Investigating nocturnal ﬁsh populations in situ using baited
underwater video: with special reference to their olfactory capabilities. J Exp Mar Biol Ecol.
Benoít HP, Swain DP. 2003. Accounting for length-and depth-dependent diel variation in
catchability of ﬁsh and invertebrates in an annual bottom-trawl survey. ICES J Mar Sci.
Ben-Yami M. 1976. Fishing with light. FAO Fishing Manuals. Farnham Surrey (UK): Fishing
News Books Ltd.
Ben-Yami M. 1988. Attracting ﬁsh with light. FAO Training Series no. 14. Rome: FAO.
Brouwer SL, Mann BQ, Lamberth SJ, Sauer WHH, Erasmus C. 1997. A survey of the South
African shore-angling ﬁshery. S Afr J Marine Sci. 18(1):165–177.
Browder R. 2012. Fishing lights at night. In-Fisherman. Available from: http://www.in-ﬁsher-
Coleman FC, Figueira WF, Ueland JS, Crowder LB. 2004. e impact of United States recre-
ational ﬁsheries on marine ﬁsh populations. Science. 305(5692):1958–1960. http://dx.doi.
Cooke SJ, Cowx IG. 2004. e role of recreational ﬁshing in global ﬁsh crises. Bioscience.
Cooke SJ, Suski CD. 2004. Are circle hooks an eﬀective tool for conserving marine and fresh-
water recreational catch-and-release ﬁsheries? Aquat Conserv. 14(3):299–326. http://
Cooke SJ, Suski CD. 2005. Do we need species-speciﬁc guidelines for catch-and-release rec-
reational angling to eﬀectively conserve diverse ﬁshery resources? Biodivers Conserv.
Cooke SJ, Cowx IG. 2006. Contrasting recreational and commercial ﬁshing: searching for com-
mon issues to promote uniﬁed conservation of ﬁsheries resources and aquatic environ-
ments. Biol Conserv. 128(1):93–108. http://dx.doi.org/10.1016/j.biocon.2005.09.019
Cooke SJ, Hogan ZS, Butcher PA, Stokesburry MJW, Raghavan R, Gallagher AJ, Hammerschlag
N, Danylchuk AJ. 2016. Angling for endangered ﬁsh: conservation problem or conservation
action? Fish Fish. 17:249–265. http://dx.doi.org/10.1111/faf.12076
Cooke et al.: Recreational shing in the dark 17
Cox SP, Walters CJ, Post JR. 2003. A model-based evaluation of active management of recre-
ational ﬁshing eﬀort. N Am J Fish Manage. 23(4):1294–1302. http://dx.doi.org/10.1577/
Danilowicz BS, Sale PF. 1999. Relative intensity of predation on the french grunt, Haemulon
ﬂavolineatum, during diurnal, dusk, and nocturnal periods on a coral reef. Mar Biol.
Daugherty DJ, Sutton TM. 2005. Diel movement patterns and habitat use of ﬂathead catﬁsh in
the lower St. Joseph River, Michigan. J Freshwat Ecol. 20(1):1–8. http://dx.doi.org/10.1080
Døving KB, Gemne G. 1965. Electrophysiological and histological properties of the olfactory
tract of the burbot (Lota lota L.). J Neurophysiol. 28(1):139–153.
Emery AR. 1973. Preliminary comparisons of day and night habits of freshwater ﬁsh in Ontario
lakes. J Fish Res Board Can. 30(6):761–774. http://dx.doi.org/10.1139/f73-131
FAO Fisheries and Aquaculture Department. 2009. Fishery and aquaculture country proﬁles:
the Republic of Maldives. Rome: FAO Fisheries and Aquaculture Department. Available
Fréon P, Gerlotto F, Soria M. 1996. Diel variability of school structure with special refer-
ence to transition periods. ICES J Mar Sci. 53(2):459–464. http://dx.doi.org/10.1006/
Hamilton RJ, Giningele M, Aswani S, Ecochard JL. 2012. Fishing in the dark-local knowl-
edge, night spearﬁshing and spawning aggregations in the Western Solomon Islands. Biol
Conserv. 145(1):246–257. http://dx.doi.org/10.1016/j.biocon.2011.11.020
Hara TJ, Zielinski BS. 2006. Fish physiology: sensory systems neuroscience. New York: Elsevier
Harvey ES, Butler JJ, McLean DL, Shand J. 2012. Contrasting habitat use of diurnal and noctur-
nal ﬁsh assemblages in temperate Western Australia. J Exp Mar Biol Ecol. 426–427:78–86.
Hazin HG, Hazin FHV, Travassos P, Erzini K. 2005. Eﬀect of light-sticks and electralume at-
tractors on surface-longline catches of swordﬁsh (Xiphias gladius Linnaeus, 1959) in the
southwest equatorial Atlantic. Fish Res. 72(2–3):271–277. http://dx.doi.org/10.1016/j.
Helfman GS. 1978. Patterns of community structure in ﬁshes: summary and overview. Environ
Biol Fishes. 3:129–148. http://dx.doi.org/10.1007/BF00006313
Helfman GS. 1993. Fish behaviour by day, night, and twilight. In: Pitcher TJ, editor. Behaviour
of Teleost Fishes. 2nd ed. London: Chapman and Hall. p. 479–512.
Hinkens E, Cochran PA. 1988. Taste buds on the pelvic ﬁn rays of the burbot, Lota lota (L.). J
Fish Biol. 32(6):975. http://dx.doi.org/10.1111/j.1095-8649.1988.tb05441.x
Horodysky AZ, Brill RW, Warrant EJ, Musick JA, Latour RJ. 2008. Comparative visual function
in ﬁve sciaenid ﬁshes. J Exp Biol. 211(22):3601–3612. http://dx.doi.org/10.1242/jeb.023358
Horodysky AZ, Brill RW, Warrant EJ, Musick JA, Latour RJ. 2010. Comparative visual function
in four piscivorous ﬁshes inhabiting Chesapeake Bay. J Exp Biol. 213(10):1751–1761. http://
Jensen EJ. 2013. Completion report: large lake assessment report for Mille Lacs Lake 2011.
Minnesota Department of Natural Resources, St. Paul. Available from: http://ﬁles.dnr.state.
Johnson BM, Martinez PJ. 1995. Selecting harvest regulations for recreational ﬁsheries: op-
portunities for research/management cooperation. Fisheries. 20(10):22–29. http://dx.doi.
Johannes RE, Neis B. 2007. e value of anecdote. In: Haggan N, Neis B, Baird IG, editors.
Fishers’ knowledge in ﬁsheries science and management. Paris: UNESCO Publishing. p.
Johnsen S. 2003. Lifting the cloak of invisibility: the eﬀects of changing optical conditions on
pelagic crypsis. Integr Comp Biol. 43(4):580–590. http://dx.doi.org/10.1093/icb/43.4.580
Bulletin of Marine Science. Vol 93, No 0. 201718
Johnsen S, Sosik HM. 2003. Cryptic coloration and mirrored sides as camouﬂage strategies
in near-surface pelagic habitats: implications for foraging and predator avoidance. Limnol
Oceanogr. 48(3):1277–1288. http://dx.doi.org/10.4319/lo.2003.48.3.1277
Johnsen S. 2012. e optics of life: a biologist’s guide to light in nature. Princeton NJ: Princeton
Johnson D. 2013. Best bobbers for night ﬁshing. In-Fisherman. Available from: http://www.in-
Krueger CC, Decker DJ. 1999. e process of ﬁsheries management. In: Kohler CC, Hubert
WA, editors. Inland ﬁsheries management in North America. 2nd ed. Bethesda: American
Fisheries Society. p. 31–59.
Lapointe NW, Cooke SJ, Imhof JG, Boisclair D, Casselman JM, Curry RA, Langer OE,
McLaughlin RL, Minns CK, Post JR, et al. 2014. Principles for ensuring healthy and produc-
tive freshwater ecosystems that support sustainable ﬁsheries. Environ Rev. 22(2):110–134.
Lennox RJ, Whoriskey K, Crossin GT, Cooke SJ. 2015. Inﬂuence of angler hook-set behaviour
relative to hook type on capture success and incidences of deep hooking and injury in a
teleost ﬁsh. Fish Res. 164:201–205. http://dx.doi.org/10.1016/j.ﬁshres.2014.11.015
Lewin WC, Arlinghaus R, Mehner T. 2006. Documented and potential biological impacts of
recreational ﬁshing: insights for management and conservation. Rev Fish Sci. 14(4):305–
Ley J, Halliday JA. 2007. Diel variation in mangrove ﬁsh abundances and trophic guilds of
northeastern australian estuaries with a proposed trophodynamic model. Bull Mar Sci.
Mazur MM, Beauchamp DA. 2006. Linking piscivory to spatial-temporal distributions of
pelagic prey ﬁsh with a visual foraging model. J Fish Biol. 69(1):151–175. http://dx.doi.
McMahon TE, Holanov SH. 1995. Foraging success of largemouth bass at diﬀerent light inten-
sities: implications for time and depth of foraging. J Fish Biol. 46(5):759–767. http://dx.doi.
Morales-Nin B, Moranta J, Garcı’a C, Tugores MP, Grau AM, Riera F, Cerda’ M. 2005. e
recreational ﬁshery oﬀ Majorca Island (western Mediterranean): some implications for
coastal resource management. ICES J Mar Sci. 62(4):727–739. http://dx.doi.org/10.1016/j.
Munz FW, McFarland WN. 1977. Evolutionary adaptations of ﬁshes to the photic environ-
ment. In: Crescitelli F, editor. Handbook of sensory physiology vol 7/5: the visual system in
vertebrates. p: 193–275.
Nelson TC, Gazey WJ, Robichaud D, English KK, Mochizuki T. 2014. Status of white sturgeon
in the lower Fraser River: report on the ﬁndings of the Lower Fraser River white sturgeon
monitoring and assessment program 2013. Summary report. Sidney BC: LGL Limited.
Available from: http://www.frasersturgeon.com/media/LFRWS_Summary_2013.pdf
New JG, Fewkes LA, Khan AN. 2001. Strike feeding behavior in the muskellunge, Esox
masquinongy: contributions of the lateral line and visual sensory systems. J Exp Biol.
Nightingale B, Longcore T, Simenstad CA. 2006. Artiﬁcial night lighting and ﬁshes. In: Rich C,
Longcore T, editors. Ecological consequences of artiﬁcial night lighting. Washington, DC:
Island Press. p. 257–276.
Paragamian VL. 1989. A comparison of day and night electroﬁshing: size structure and catch
per unit eﬀort for smallmouth bass. N Am J Fish Manage. 9(4):500–503. http://dx.doi.
Pavlov DS, Kasumyan AO. 1990. Sensory principles of the feeding behavior of ﬁshes. J Ichthyol.
Pawson MG, Glenn H, Padda G. 2008. e deﬁnition of marine recreational ﬁshing in Europe.
Mar Policy. 32(3):339–350. http://dx.doi.org/10.1016/j.marpol.2007.07.001
Cooke et al.: Recreational shing in the dark 19
Pohlmann K, Atema J, Breithaupt T. 2004. e importance of the lateral line in nocturnal pre-
dation of piscivorous catﬁsh. J Exp Biol. 207(17):2971–2978. http://dx.doi.org/10.1242/
Pope KL, Willis DW. 1996. Seasonal inﬂuences on freshwater ﬁsheries sampling data. Rev Fish
Sci. 4(1):57–73. http://dx.doi.org/10.1080/10641269609388578
Post JR, Sullivan M, Cox S, Lester NP, Walters CJ, Parkinson EA, Paul AJ, Jackson L, Shuter B.
2002. Canada’s recreational ﬁsheries: the invisible collapse? Fisheries. 27(1):6–17. http://
Quinn S. 2014. Night ﬁshing largemouth bass. In-Fisherman. Available from: http://www.in-
Rapp T, Hallermann J, Cooke SJ, Hetz SK, Wuertz S, Arlinghaus R. 2012. Physiological and be-
havioural consequences of capture and retention in carp sacks on common carp (Cyprinus
carpio L.), with implications for catch-and-release recreational ﬁshing. Fish Res. 125–
Reebs SG. 2002. Plasticity of diel and circadian activity rhythms in ﬁshes. Rev Fish Biol Fish.
Reeves KA, Bruesewitz RE. 2007. Factors inﬂuencing the hooking mortality of walleyes caught
by recreational anglers on Mille Lacs, Minnesota. N Am J Fish Manage. 27:443–452. http://
Reid DD, Montgomery SS. 2005. Creel survey based estimation of recreational harvest of
penaeid prawns in four southeastern Australian estuaries and comparison with commercial
catches. Fish Res. 74(1–3):169–185. http://dx.doi.org/10.1016/j.ﬁshres.2005.03.007
Robbins WD, Peddemors VM, Broadhurst MK, Gray CA. 2013. Hooked on ﬁshing? Recreational
angling interactions with the critically endangered grey nurse shark Carcharias taurus in
eastern Australia. Endang Species Res. 21:161–170.
Smallwood CB, Beckle y LE, Sumner NR. 2006. Shore-based recreational angling in the Rottnest
Island Reserve, Western Australia: spatial and temporal distribution of catch and ﬁshing ef-
fort. Pac Conserv Biol. 12(3):238–251. http://dx.doi.org/10.1071/PC060238
Smallwood CB, Pollock KH, Wise BS, Hall NG, Gaughan DJ. 2011. Quantifying recreational
ﬁshing catch and eﬀort: a pilot study of shore-based ﬁshers in the Perth Metropolitan area.
Fisheries Research Report No. 216. Final NRM Report - Project No. 09040. Department of
Fisheries. Available from: http://www.ﬁsh.wa.gov.au/Documents/research_reports/frr216.
Tufts BL, Holden J, DeMille M. 2015. Beneﬁts arising from sustainable use of North America’s
ﬁshery resources: economic and conservation impacts of recreational angling. Int J Environ
Stud. 72(5):850–868. http://dx.doi.org/10.1080/00207233.2015.1022987
USFWS. 2009. Recreational Fishing Management Plan. Back Bay National Wildlife Refuge.
Virginia: US Department of the Interior. Fish and Wildlife Service. Available from: http://
Veiga P, Ribeiro J, Goncalves JMS, Erzini K. 2010. Quantifying recreational shore angling catch
and harvest in southern Portugal (north‐east Atlantic Ocean): implications for conser-
vation and integrated ﬁsheries management. J Fish Biol. 76(9):2216–2237. http://dx.doi.
UN FAO. 2012. Recreational ﬁsheries: FAO technical guidelines for responsible ﬁsheries. No.
13. Rome. 176 p. (written under contract by R Arlinghaus, SJ Cooke, and B Johnson)
Venturelli P, Bence J, Brendan T, Lester N, Rudstam L. 2014. Mille Lacs Lake walleye blue rib-
bon panel data review and recommendations for future data collection and management.
Prepared for Minnesota DNR. Available from: https://fwcb.cfans.umn.edu/sites/fwcb.
Warrant EW. 1999. Seeing better at night: life style, eye design and the optimum strategy of
spatial and temporal summation. Vision Res. 39(9):1611–1630. http://dx.doi.org/10.1016/
Bulletin of Marine Science. Vol 93, No 0. 201720
Warrant EW, Johnsen S. 2013. Vision and the light environment. Curr Biol. 23:R990–R994.
Zeller D, Booth S, Davis G, Pauly D. 2007. Re-estimation of small-scale ﬁshery catches for
US ﬂag-associated island areas in the western Paciﬁc: the last 50 years. Fish B-NOAA.