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Seasonal patterns in the nocturnal distribution and behavior of the mesopelagic fish Maurolicus muelleri at high latitudes


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Acoustic scattering layers (SL) ascribed to pearlside Maurolicus muelleri were studied in Masfjorden, Norway, using upward-looking echo sounders cabled to shore for continuous long-term measurements. The acoustic studies were accompanied by continuous measurements of surface light and supplemented with intermittent field campaigns. From autumn to spring, young M. muelleri formed an SL in the upper similar to 75 to 150 m in the daytime, characterized by migration to near-surface water near dusk, subsequent 'midnight sinking', followed by a dawn ascent before a return to the daytime habitat. Light levels were similar to 1 order of magnitude lower during the dawn ascent than for ascent in the afternoon, with the latter terminating before fish reached upper layers on similar to 1/3 of the nights from late November to mid-April. Adults showed less tendency of migration during autumn and winter, until the SLs of young and adults merged in late spring, and thereafter displayed coherent migration behavior. The midnight sinking became progressively deeper from autumn to winter but was strongly reduced from mid-May when the darkest nocturnal light intensity (PAR) at the surface was above 10(-3) mu mol m(-2) s(-1). The pearlside took on schooling in upper waters during the even lighter nights in early June, with minimum light of similar to 5 x 10(-3) to 10(-1) mu mol m(-2) s(-1) at the surface. Nocturnal schooling ceased in early July, and midnight sinking reappeared in mid-August. We suggest that the strong variation in nocturnal light intensity at high latitudes provides changing trade-offs between visual foraging and avoiding predators and hence varying time budgets for feeding in the upper, productive layers.
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Mar Ecol Prog Ser
Vol. 521: 189–200, 2015
doi: 10.3354/meps11139 Published February 17
Mesopelagic fish are widely distributed in the
world’s oceans (Gjøsæter & Kawaguchi 1980, Dal-
padado & Gjøsæter 1988, Irigoien et al. 2014). These
fish act as a trophic link between zooplankton
(Shreeve et al. 2009) and commercially valuable and
other predators (Giske et al. 1990, Potier et al. 2007,
Doksæter et al. 2008). They may also play a signifi-
cant role in the carbon transport from productive
upper layers to food-deprived deeper layers (Her -
nández-León et al. 2010, Bianchi et al. 2013, Irigoien
et al. 2014). Diel vertical migration (DVM) is common
among both zooplankton and fish, usually explained
as trade-offs between avoiding predators and forag-
ing (Rosland & Giske 1994, Pearre 2003). Various
studies have shown that light acts as proximate ini-
tiator for DVM (Kampa & Boden 1954, Kampa 1970),
and whether DVM behavior relates to the preference
for a specific absolute intensity or isolume (Sweatt &
Forward 1985), the rate of change in the light inten-
sity (Ringelberg 1995, Cohen & Forward 2009), or the
preference for a range of light intensities (Staby &
Aksnes 2011) has been debated.
Most focus on the mesopelagic fish DVM in rela-
tion to light has been on light conditions during the
© Inter-Research 2015 ·*Corresponding author:
Seasonal patterns in the nocturnal distribution
and behavior of the mesopelagic fish Maurolicus
muelleri at high latitudes
Perdana K. Prihartato1, Dag L. Aksnes2, Stein Kaartvedt1, 3,*
1King Abdullah University of Science and Technology, Red Sea Research Center, Thuwal 23955-6900, Saudi Arabia
2Hjort Centre for Marine Ecosystem Dynamics, Department of Biology, University of Bergen, Norway
3University of Oslo, Department of Biosciences, PO Box 1066 Blindern, 0316 Oslo, Norway
ABSTRACT: Acoustic scattering layers (SL) ascribed to pearlside Maurolicus muelleri were stud-
ied in Masfjorden, Norway, using upward-looking echo sounders cabled to shore for continuous
long-term measurements. The acoustic studies were accompanied by continuous measurements
of surface light and supplemented with intermittent field campaigns. From autumn to spring,
young M. muelleri formed an SL in the upper ~75 to 150 m in the daytime, characterized by migra-
tion to near-surface water near dusk, subsequent ‘midnight sinking’, followed by a dawn ascent
before a return to the daytime habitat. Light levels were ~1 order of magnitude lower during the
dawn ascent than for ascent in the afternoon, with the latter terminating before fish reached upper
layers on ~1/3 of the nights from late November to mid-April. Adults showed less tendency of
migration during autumn and winter, until the SLs of young and adults merged in late spring, and
thereafter displayed coherent migration behavior. The midnight sinking became progressively
deeper from autumn to winter but was strongly reduced from mid-May when the darkest noctur-
nal light intensity (PAR) at the surface was above 10−3 µmol m−2 s−1. The pearlside took on school-
ing in upper waters during the even lighter nights in early June, with minimum light of ~5 ×10−3
to 10−1 µmol m−2 s−1 at the surface. Nocturnal schooling ceased in early July, and midnight sinking
reappeared in mid-August. We suggest that the strong variation in nocturnal light intensity at
high latitudes provides changing trade-offs between visual foraging and avoiding predators and
hence varying time budgets for feeding in the upper, productive layers.
KEY WORDS: Behavior · Diel vertical migration · Light levels · Mesopelagic · Acoustics
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Mar Ecol Prog Ser 521: 189–200, 2015
day (Baliño & Aksnes 1993) or differences between
day and night (O’Driscoll et al. 2009, Klevjer et al.
2012). However, light levels also vary at night. Fish
vertical distribution, predation, and foraging activi-
ties can significantly be affected by nocturnal lights
in relation to moonlight (Gliwicz 1986, Benoit-Bird et
al. 2009) but also by seasonal cycles in the nocturnal
light climate (Sameoto 1989, Rasmussen & Giske
1994, Kaartvedt 2008). Masfjorden, Norway, is
located at a high latitude (~60° N) and represents a
relatively deep, protected, semi-enclosed water
body, providing a natural laboratory to study meso-
pelagic fish. Maurolicus muelleri, or pearlside, which
is the focus species of this study, is the prevailing
acoustic target in the upper ~200 m of Masfjorden
during daytime, being distributed shallower during
the night (Giske et al. 1990, Baliño & Aksnes 1993,
Rasmussen & Giske 1994, Kaartvedt et al. 1998,
Aksnes et al. 2004, Staby & Aksnes 2011, Staby et al.
2011). However, not all parts of the population
appear to take part in DVM at all times (Staby et al.
Several studies have indicated that the vertical dis-
tribution of pearlside is related to a certain range of
light levels (Baliño & Aksnes 1993, Kaartvedt et al.
1996, Staby & Aksnes 2011). Pearlside appear to take
advantage of the so-called anti-predation window
(Clark & Levy 1988) and forage in upper waters at
dusk and dawn. Particularly juveniles migrate to the
surface in the afternoon and spend a short period
near the surface, followed by midnight sinking and a
subsequent dawn rise before returning to their day-
time depth (Giske et al. 1990, Baliño & Aksnes 1993,
Rasmussen & Giske 1994, Staby et al. 2011, 2013).
According to the anti-predation window hypothe-
sis (Clark & Levy 1988), small planktivorous fish seek
out intermediate light levels that are sufficient for
obtaining food while at the same time being suffi-
ciently low to be relatively safe from visually search-
ing piscivores. ‘Midnight sinking’ follows as it
becomes too dark for foraging at night. However, at
high latitudes, the timing of the anti-predation win-
dow in upper waters may change, as nights get much
lighter during spring and summer. This pattern may
prevent vertically migrating fish from entering sur-
face layers for feeding if there is too much light
(Sameoto 1989, Kaartvedt 2008). On the other hand,
the dusky light levels of summer nights might cause
an extended, shallow anti-predation window, per-
mitting foraging in likely food-rich upper waters
throughout the night. Some previous studies in Nor-
way have suggested that midnight sinking may be
skipped entirely in summer (Rasmussen & Giske
1994, Kaartvedt et al. 1998, Staby et al. 2011). M.
muelleri may even initiate schooling in upper layers
during light summer nights (Kaartvedt et al. 1998).
However, such reports are mostly from short-term
observations, and light levels associated with
switches between the different behaviors have not
been established.
In this study, we take advantage of the unique
opportunity for a long-term study of M. muelleri
offered by a deep locality in close proximity to land.
We applied upward-looking echosounders cabled to
shore that provided continuous data on vertical
migration behavior from autumn throughout the sub-
sequent summer. This research was accompanied
with continuous measurements of surface light and
intermittent field campaigns, including measure-
ments of water-column light extinction. We here
address the seasonal patterns in DVM behavior of M.
muelleri, emphasizing the nocturnal distribution,
particularly focusing on the relatively light summer
Acoustic measurements
The study was carried out at a ~370 m deep loca-
tion in Masfjorden, Norway (~60° 50’ N, ~5° 30’ E),
from 7 October 2010 to 15 August 2011. Continuous
acoustic measurements were made near the location
of former studies (Staby et al. 2011, Dypvik et al.
2012a,b; see Fig. 1a). We used 3 upward looking
SIMRAD EK60 split beam echo sounders (7.1° beam
widths), mounted on the bottom (38 kHz; ~370 m)
and on rigs anchored to the bottom, floating at
~250 m (120 kHz) and ~90 m (200 kHz) (Fig. 1). Echo
sounder depths were selected to give high- resolution
data of different segments of the water column. The
echo sounders were calibrated at the surface using
standard methods (Foote et al. 1987). Pulse lengths
and ping rates were 128 µs and 1 to 2 pings s−1
(200 kHz), 256 µs and 1 to 2 pings s−1 (120 kHz), and
512 µs and 1 ping s−1 (38 kHz), respectively.
The submerged transceivers were kept in pres-
sure-proof casings and cabled to shore for power and
transmittance of digitized signals to laptop comput-
ers, where data were stored in raw format for later
analysis. The laptops were connected to the Internet
for delivery of real-time echograms via web-based
interfaces as well as to remotely control the echo
sounders. This linkage allowed remote access for
restarting the echo sounders after periods of power
Prihartato et al.: Nocturnal distribution and behavior of Maurolicus muelleri
failures, which occurred due to a periodically unsta-
ble electrical line.
Observed DVM patterns were largely similar at 38
and 120 kHz. This similarity between the 2 frequen-
cies suggests that the observed DVM patterns can be
ascribed to fish because most plankton will not be
detected at 38 kHz with the settings applied here
(Kaartvedt et al. 2008). Moreover, Maurolicus muel-
leri mainly occurs in the upper ~250 m of the water
column. Hence, we only present data from the float-
ing echo sounders at 120 and 200 kHz, although we
also refer to findings at 38 kHz. Data were obtained
for 276 d and 269 d for 120 kHz and 200 kHz, respec-
Processing and visualization of acoustic data
Echograms were visualized using MATLAB (ver-
sion 2012b). Seasonal patterns of DVM were pre-
sented as monthly averaged daily 24 h echograms,
with bin size of 2 min ×5 cm, for the section of ~0 to
250 m. We organize the monthly echograms into 4
seasonal categories: autumn (October to November),
winter (December to February), spring (March to
May), and summer (June to August).
To display specific behavior patterns of organisms
forming nighttime acoustic scattering layers (SLs), 4
high-resolution examples of SLs were chosen from
both echo sounders to represent events that occurred
in the course of the registration period: interrupted
ascent (6 to 8 January), termination of midnight sink-
ing (14 to 16 May), schooling behavior at light sum-
mer nights (18 to 20 June) and resumption of mid-
night sinking (14 to 16 August). This selection was
done for the whole days at 120 kHz, and with finer
resolution for better visualization at 200 kHz.
The seasonal patterns of nocturnal scattering were
visualized by importing all nocturnal data from
120 kHz and 200 kHz into respective echograms
spanning the whole study period. Night was defined
as the period after sunset and before sunrise at local
time (UTC + 1 h; UTC + 2 h during daylight saving
time from 28 March to 30 October 2011). Echo data
were intermittently affected by different sources of
noise, so prior to importing the data for final analysis,
supervised noise removal was conducted to exclude
‘bad echoes’ using the LSSS software (Korneliussen
et al. 2009). This process involved filtering of a 9-point
moving average window to remove irregular spikes.
The output value of this process was the scattering
area (SA), which was then translated into logarithmic
value of backscattering volume (SV; Knudsen 1990).
SAwas integrated at a resolution of 15 min ×2 m. The
upper 0–4 m was deleted to avoid surface-generated
Composition of scattering layers
SLs in Masfjorden corresponding to those referred
to in this paper have repeatedly been ascribed to M.
muelleri. Previous studies conclude that the young-of
the year form SLs in the upper 100 m from autumn to
spring, and the adults stay at greater depths (Baliño
& Aksnes 1993, Rasmussen & Giske 1994, Staby et al.
2011). Results from extensive trawling with a 100 m2
pelagic trawl at the start (7 October 2010) and end
(15 August 2011) of the present study were in accor-
dance with these previous findings, with daytime
Fig. 1. (A) Map of Masfjorden depicting locations of the 3 echo sounders. The upward-looking echo sounders were mounted
on the bottom ~370 m (38 kHz) and floating in the water column at ~250 m (120 kHz) and 90 m (200 kHz). (B) Sketch of the
experimental set up
Mar Ecol Prog Ser 521: 189–200, 2015
catches in the upper 150 m being completely domi-
nated by M. muelleri. As the identity of these SLs is
already well documented through previous work,
details of trawl catches are not presented for the pur-
pose of this paper. Additional plankton targets were
apparent, mostly at 200 kHz.
Light measurements
Photosynthetically active radiation (PAR, 400 to
700 nm) was continuously recorded about 2 m above
the surface (referred to as surface irradiance) from 10
December 2010 to 15 August 2011 with a calibrated
LI-190 quantum sensor and data stored on a LI-1400
data logger. Measurements were averaged and
stored every 15 min. The lower threshold of the sen-
sor was 0.0001 µmol m−2 s−1. The surface irradiance at
the darkest period at night (minimum nocturnal light
levels) was higher than this sensitivity threshold from
28 April until records ended on 15 August 2011. Prior
to April 28, the sensitivity of the sensor was not ade-
quate to measure light during night. A few inci-
dences of relatively high nocturnal light measured
during winter were removed from the dataset be -
cause these incidents were ascribed to artificial light-
ing at the shore station where the logging was con-
We used the results of surface irradiance as a proxy
for the irradiance at the top of the SL (as visible in the
high-resolution 200 kHz echogram when using an SV
threshold of −75 dB) when the SL approached the
surface in the morning (right before dawn descent)
and evening (dusk ascent). However, sometimes
interrupted ascents of the SL were observed in the
afternoon, i.e. the top of the SL was then located well
below the surface. In these cases, results from under-
water light measurements were used to approximate
the ambient irradiance at the top of the SL (explained
Underwater irradiance was measured using a
RAMSES ACC hyper-spectral radiometer (Trios-opti-
cal sensors, Oldenburg, Germany) around noon on
26 January, 22 February, 11 April, 16 June, and 16
August 2011. These measurements were taken at 1,
5, and 10 m and then every 10 m down to ~90 m
depth. For each of these depths, we calculated the
attenuation coefficient (Kz)for downwelling irradi-
ance (PAR) between the surface (i.e. 1 m depth) and
the depth (z) in question according to the following
Kz= −ln(E1/Ez)/(z − 1) (1)
where E1and Ezare the measured downwelling irra-
diance at 1 and zm respectively. Simultaneous regis-
tration of surface irradiance was obtained for each of
the underwater measurements so that the Kzesti-
mates were based on simultaneous measurements
of E1and Ez. As noted above, for the interrupted
ascents, we approximated the ambient irradiance
(ETSL), on the order of magnitude, for the depth (ZTSL)
where the upward migration of the SL halted accord-
ing to Eq. (2):
ETSL =E0exp[−KZTSL] (2)
where E0is the measured surface irradiance at the
time of the interrupted ascent, and Kis the estimated
attenuation coefficient at the date closest to the inter-
rupted ascent and for the depth closest to ZTSL.
In accordance with the anti-predation window
hypothesis (and previous studies on pearlside), we
hypothesize that M. muelleri will exploit dim light to
forage while concurrently avoiding visually search-
ing piscivores (Clark & Levy 1988, Rosland & Giske
1994, Staby et al. 2013). The extension of the anti-
predation window (by migration) is according to this
hypothesis important for their foraging and survival
success. M. muelleri (particularly juveniles) appear
to mainly forage in upper waters during dusk and
dawn (Staby et al. 2011), and we used the time dur-
ing which surface light values were between 10−3
and 1 µmol m−2 s−1 as a rough proxy to assess how the
duration of potential foraging in near-surface waters
would vary throughout the year. These values were
selected based on the ranges measured in the course
of this investigation and also encompassing the val-
ues given for the top of the Maurolicus SL at any time
of day and year by Staby & Aksnes (2011).
Nocturnal light
Surface light could be detected even during the
darkest part of the night from 28 April to the end of
the registration period (15 August 2011). The mini-
mum measured nocturnal irradiance spanned 2
orders of magnitude, from 0.00016 µmol m−2 s −1 on
28 April to 0.061 µmol m−2 s−1 on 22 June. Thereafter,
it decreased to the end of the records, yet with some
variation between nights (Fig. 2).
Surface light was always above the registration
threshold when the upper SL of Maurolicus muelleri
appeared close to the surface at dusk and dawn
(Fig. 3). Due to the shallow distribution, these light
Prihartato et al.: Nocturnal distribution and behavior of Maurolicus muelleri
levels approximately represent light at the top of the
near-surface scattering layer (SL), although the
migrations sometimes stopped at a few meters depth,
apparently hampered by a pycnocline (acoustically
visible and likely related to low-salinity surface
layer). Light levels proved to differ significantly
between dusk and dawn (Mann-Whitney Utest, p <<
0.001), regardless of season. On average, the surface
light at initiation of descent in the morning was rela-
tively stable at 0.032 µmol m−2 s−1. This was an order
of magnitude lower than the average light level
when fish reached the surface layers in the afternoon
(0.21 µmol m−2 s−1). However, as will be outlined
below, on a number of nights, fish interrupted their
ascents in the afternoon and did then not experience
these high light intensities (Fig. 3B).
Behavioral pattern of M. muelleri scattering layers
In autumn and winter, at least 2 SLs ascribed to M.
muelleri were recorded, referred to here as the deep
scattering layer (DSL) and shallow scattering layer
(SSL), often structured into several sub-layers, which
to some extent are masked by the monthly average
presented in Fig. 4. The DSL was situated at 150 to
200 m day and night, with some but only a limited
range of vertical migration subsequent to October
and until March/April. Thereafter, all prevailing
acoustic scatter in the upper ~200 m performed
coherent DVM until records ended in August.
The younger M. muelleri, which formed the SSL,
stayed at depths of ~75 to 125 m during daytime but
were located progressively deeper through the regis-
tration period, so that daytime depth reached 150 m
in spring (Fig. 4). These M. muelleri always carried
out DVM, ascending to the surface at dusk followed
by ‘midnight sinking’ with a subsequent ascent at
dawn before descending to their daytime depths.
The nocturnal distribution of the SSL subsequent to
the descent from near-surface waters at dusk deep-
ened throughout autumn to winter, from the upper
~30 to 50 m early in the registration period toward 70
to 100 m at the end of winter and in early spring,
thereafter reversing to a shallower distribution
(Figs. 4 & 5).
Fig. 2. Minimum nocturnal surface
irradiance (PAR) as measured with a
LiCor quantum meter and fish behav-
ior derived from echograms. There
are no light measurements prior to 28
April due to insufficient sensitivity of
the PAR sensor. Dots are measured
minimum light values, with the line
representing the moving average (n =
23). Color and symbol depict fish
behavior on the respective nights
Fig. 3. Downwelling irradiance at the
surface when the upper SL of Mauroli-
cus muelleri is located closest to the sur-
face. (A) Prior to descent in the morn-
ing; (B) after ascent in the afternoon.
Blue points represent the calculated
underwater light intensity at the top of
the SL during interrupted ascent
Mar Ecol Prog Ser 521: 189–200, 2015
Fig. 4. Monthly averaged echograms at 120 kHz, displaying 24 h images for the upper ~280 m. Averages are based on the fol-
lowing numbers of days: Oct (26 d), Nov (30 d), Dec (21 d), Jan (29 d), Feb (19 d), Mar (30 d), Apr (30 d), May (29 d), Jun (30 d),
Jul (17 d), and Aug (15 d). Color scale refers to backscattering strength (SV) values (dB). Time is given in UTC (local stan-
dard time − 1 h)
Fig. 5. Echograms depicting the nocturnal backscatter for the whole study period at (A) 120 kHz and (B) 200 kHz. Gaps in the
datasets are from periods without records. Color scale refers to backscattering strength (SV) values (dB)
Prihartato et al.: Nocturnal distribution and behavior of Maurolicus muelleri
From late autumn to spring, there was an asyn-
chrony between the dusk and dawn ascent. In the
afternoon, individuals forming the SSL repeatedly
arrested their ascent at various depths (mean = 24.6 ±
17.4 m) before reaching upper layers (e.g. Fig. 6A).
Such interrupted ascents were recorded on 46 of the
158 dates from 27 November 2010 to 18 April 2011,
particularly frequently in March. The estimated
underwater irradiance at the top of the SL during
interrupted ascent ranged between 10−3 and 10−1
µmol m−2 s−1 and was about 1 to 2 orders of magni-
tude lower than during the afternoons they contin-
ued to the surface (Fig. 3B). Interrupted ascents were
never recorded in the morning, so that dawn rise to
surface waters was documented on every date dur-
ing the same period.
A change in the nocturnal distribution took place
from mid-May. The midnight sinking behavior then
tended to cease, with a subsequently increased por-
tion of the nocturnal SL inhabiting near-surface
waters (upper ~25 m) throughout the night (Fig. 6B).
The minimum surface irradiance at night at the time
of initiation of this behavior was ~10−3 µmol m−2 s−1
(Figs. 2 & 7A,B). The structure of the nocturnal SLs
subsequently changed in early June. On the first
dates of the month, fish were schooling upon reach-
ing the surface early at night, while schooling per-
sisted throughout the short nights in mid-summer
(Figs. 2, 6C & 7C,D). During this time, minimum noc-
turnal surface irradiance was mostly between ~5 ×
10−3 and 10−1 µmol m−2 s−1, while the fishes reached
upper waters at levels of ~1 µmol m−2 s−1 (Fig. 7C,D).
Schooling subsided in early July. Fish thereafter con-
tinued to stay in upper layers throughout night until
mid-August (Fig. 6D), when midnight sinking re -
appeared as nocturnal surface irradiance apparently
(irregular light-data at this time) again dropped
toward 10−3 µmol m−2 s−1 (Figs. 2 & 7E).
The duration of periods with nocturnal surface
light between 10−3 and 1 µmol m−2 s−1 (i.e. a coarse
proxy for the nocturnal extension of the anti-predator
window in upper layers) varied through the year
Fig. 6. Selected echograms for 3 consecutive days representing different diel migration patterns. (A) Interrupted ascent in win-
ter (6−8 January 2011), (B) termination of midnight sinking in late spring (14−16 May), (C) schooling in mid-summer (18−20
June), and (D) resumption of midnight sinking in late summer (14−16 August). Color scale refers to backscattering strength
(SV) values (dB)
Mar Ecol Prog Ser 521: 189–200, 2015
(Fig. 8) and roughly comprised the time fish resided
in upper waters. It was relatively short (~2 h) during
winter and early spring, as confined to dusk and
dawn periods. The time-window for this light range
increased in mid-May, being 5 to 6 h in late May and
early June. The period thereafter became shorter as
night became shorter toward the end of June (~4 h),
before subsequently increasing with increasing
length of the night until the end of the registration
period in mid-August, when the duration would drop
due to resumption of midnight sinking.
We have unveiled seasonal variations in DVM-
behavior of scattering layers (SLs) ascribed to Mau-
rolicus muelleri (sampling from this study and pre -
vious sampling; see references in ‘Material and
methods’) by exploring long-term, high-resolution
acoustic data. Emphasis has been on the nocturnal
vertical distribution in a high-latitude ecosystem
where nocturnal light conditions vary strongly
throughout the year. Individuals in the deeper SL
Fig. 7. Echograms of the upper 40 m showing
nocturnal SL (200 kHz) with corresponding sur-
face light intensities depicting (A,B) termination
of midnight sinking (15−16 May), (C,D) school-
ing behavior (18−19 June) and (E) resumption of
midnight sinking (15 August; E). Color scale
refers to backscattering strength (SV) values (dB)
Fig. 8. Nocturnal extension of the
duration of the anti-predation win-
dow, with grey circles depicting
observations and the line indicating
the moving average (n = 23). The
antipredation window relates to
intermediate light levels sufficient for
obtaining food, while at the same
time being sufficiently low to offer
relative protection from visually
searching piscivores. We selected the
periods with surface light values
being between 10−3 and 1 µmol m−2
s−1 based on the light ranges meas-
ured between Maurolicus muelleri
reaching the surface in the afternoon
and initiation of midnight sinking
Prihartato et al.: Nocturnal distribution and behavior of Maurolicus muelleri
(adult part of the population) did not appear to
migrate extensively during fall and winter, yet the
mid-water vertical distribution was somewhat shal-
lower at night. Corresponding patterns emerge from
previous studies (Giske et al. 1990, Staby & Aksnes
2011). Different behavior in juveniles and adults has
been interpreted as different trade-off between for-
aging and predator avoidance in the 2 groups
(Rosland & Giske 1994), as also found for other
pelagic taxa (De Robertis 2002, Pearre 2003). Adults
seem to have no or negative growth during winter
(Rosland & Giske 1997) yet may forage in their day-
time depth on plankton, such as overwintering cope-
pods (Bagøien et al. 2001). By largely remaining in
deep water, adult pearlside strongly increase their
probability of survival to the spawning season in
spring (Rosland & Giske 1997).
Individuals of the shallowest SL always carried out
DVM, as was also the case for the adults during
spring and summer, again in accordance with previ-
ous studies (Staby et al. 2011). However, the migra-
tion pattern varied and comprised migrations with
and without midnight sinking, interrupted ascents in
the evening and shifts from occurrence in diffuse
nocturnal SL through most of the year to nocturnal
near-surface schooling behaviors during the lightest
part of the year.
Behavioral responses to nocturnal lights
It is well established that the vertical distribution of
pearlside is related to light intensity in daytime and
during diel vertical migrations (Baliño & Aksnes
1993, Rasmussen & Giske 1994). We here unveil how
changes in nocturnal light levels also affect the verti-
cal distribution and behavior of pearlside. We largely
refer to surface light, and it must also be noted that
the underwater light intensity we have estimated
involves interpolation of the light attenuation coeffi-
cient over relatively large time periods. This interpo-
lation has obviously reduced the accuracy of our esti-
mates of underwater light but nevertheless provides
useful information concerning the order of magni-
tude of the ambient light intensities experienced by
the fishes.
For the whole period of measurements, estimated
light intensity at the top of the SL typically spanned 4
orders of magnitude (10−4 to 1.6 µmol m−2 s−1; the lat-
ter being surface value when the fish reached their
shallowest distribution). The additional light extinc-
tion from top to bottom of the SL should also be con-
sidered. Therefore, although the vertical positioning
of M. muelleri is tightly related to light, this is not a
fixed light value. In line with Staby & Aksnes (2011),
our results suggest that pearlside follow preferred
ranges of light intensity rather than a constant fixed
isolume and that these ranges span several orders of
magnitude. This finding concurs with conclusions
from studies of mesopelagic fish in other systems
(Roe 1983, Benoit-Bird et al. 2009).
At shorter time scales, the vertical distribution
appears to be coupled to a narrower range of light
levels (Baliño & Aksnes 1993, Staby & Aksnes 2011),
although there were notable variations from day to
day. This variation particularly related to afternoons
with interrupted ascents, but other aspects of the
behavior also did not fully match the variation in light
levels (e.g. Fig. 2). Some of this variation might
reflect methodological constraints, as interpretations
from echograms were subjective, but evidently other
factors than light will also affect behavior. Finally,
our calculation of underwater light intensity has not
accounted for variations in Kcaused by variations in
the radiant field, such as the change in the angle of
incoming sunlight during the day, cloudiness, and
wave action.
Within days, the surface light intensity was on
average ~1 order of magnitude weaker when the SL
reached the surface during the morning ascent than
during the corresponding ascent in the afternoon.
This pattern corroborates findings by Staby & Aksnes
(2011). The weaker light when fish reached the sur-
face in the morning would be in line with fish being
motivated for early feeding after a long night without
foraging. In contrast, this behavior is contrary to the
assumption that hungry fish would be more prone to
undertake risky behavior (Dill 1983) and therefore
extend the duration of their dawn ascent in suppos-
edly food-rich waters into higher light intensities.
Alternatively, this behavior might have a pure physi-
ological cause if dark-adapted fish are more efficient
in detecting prey at low light levels in the morning
than in the evening (this hypothesis might relate both
to the pearlside and their predators). The dark-to-
light adaptation in the retina is indeed much more
rapid than the light-to-dark adaptation (Ferwerda et
al. 1996). This hypothesis has been suggested as one
possible explanation for the emergence of small
planktivorous coral fishes from their nocturnal shel-
ter in the morning at lower light levels than the levels
at which they return to their shelter in the evening
(Rickel & Genin 2005). Note however, that the trade-
off between feeding and predator avoidance is very
different in that setting. Small planktivorous fish on
coral reefs are safest in bright light since they spot
Mar Ecol Prog Ser 521: 189–200, 2015
their predators by sight and can retract to nearby
shelters when threatened as well as when light
decreases (Holbrook & Schmitt 2002). Mesopelagic
fish, in contrast, rely on hiding in dim light.
Interrupted ascent
Interrupted ascent behavior was frequently
recorded from late autumn to mid-March yet at dif-
ferent depths. In spring, the estimated underwater
irradiance at the top of the SL during interrupted
ascent was about 1 to 2 orders of magnitude lower
than during the afternoons that the fish continued to
the surface (Fig. 3). This pattern might suggest some
relation to predator avoidance behavior. Previous
studies have shown instantaneous diving responses
among mesopelagic fish to the presence of predators
(Kaartvedt et al. 2012), and such responses were also
recorded for M. muelleri during this study (e.g. Godø
et al. 2014). However, there were no systematic
records of excessive numbers of predators associated
with such events, and the interrupted ascents were
recorded coherently by all echo sounders, located
several hundred meters away from each other. Alter-
natively, as the copepod Calanus ascends from over-
wintering in winter and early spring, it is possible
that satiation following feeding during the ascent
reduces motivation for further migration during this
time of the year (Staby et al. 2011). However, such
reaction to satiation would expectedly be an individ-
ual response (Pearre 2003) and not the population
response indicated in the echograms. Migrations
might also be stopped by gradients in temperature
and salinity, yet such gradients are not expected at
depth and would not explain the consistent differ-
ence between dawn and dusk ascents. Overall, the
most likely reason for the interrupted ascents in the
afternoon seems to have some relation to perceived
risk of predation, but this behavior is still poorly
Midnight sinking
M. muelleri does not forage in darkness, and mid-
night sinking apparently takes place when the condi-
tions are too dark for visual detection of prey (Giske
et al. 1990). Accordingly, midnight sinking was initi-
ated as nights became darker in late summer (Fig.
7E). Resumption of midnight sinking in August is also
documented by Staby et al. (2011). In parallel to the
darker nights in August, there may have been
changes in the distribution of potential prey, as the
copepod Calanus finmarchicus tends to start des cen -
ding for overwintering during this time of the year,
with potential effects on the diel migration pattern of
mesopelagic fish (Kaartvedt et al. 2009, Dypvik et al.
The nocturnal distribution became deeper
through fall and winter (Fig. 5). Giske & Aksnes
(1992) suggested that pearlside was seeking warm
temperature at night for more rapid digestion of the
afternoon meal, in this way maximizing growth (cf.
Wurtsbaugh & Neverman 1988). We do not have
temperature profiles through winter to correlate
with the observed distributions, but the subsurface
temperature maximum will become progressively
deeper due to cooling from above (Bagøien et al.
2001). This pattern would be in accordance with
the observations. Results from Staby et al. (2011)
did not fully support the importance of temperature
profiles for pearlside that perform midnight sinking
behavior, and these authors referred to predator
avoidance from visually searching piscivores as an
alternative explanation. We could not measure sur-
face light during the dark winter nights due to lack
of instrument sensitivity, but light intensities at the
upper SL during midnight sinking would have
been 3 to 5 orders of magnitude less than surface
light (based on measurements of extinction), i.e.
<10−8 µmol m−2 s−1. In early spring, when nocturnal
surface light could be detected, midnight sinking
occurred when this light decreased to ~10−3 µmol
m−2 s−1, then translating into <10−6 µmol m−2 s−1 at
the depth of midnight sinking. Gadoids appear to
be the most important predators on M. muelleri in
Masfjorden (Giske et al. 1990, Staby & Aksnes
2011). Ryer & Olla (1999) showed that juveniles of
the gadoid walleye pollock could forage success-
fully on Artemia at very low light intensities (5 ×
10−7 µmol m−2 s−1). This result suggests that mid-
night sinking of M. muelleri actually may be bene-
ficial to avoid nocturnal predators.
Termination of midnight sinking
From mid-May to mid-August, M. muelleri re -
mained in upper layers throughout the nights, with
limited evidence of midnight sinking. This time rep-
resents a period of the year when the concentration
of zooplankton peaks in upper waters of Masfjorden
(Aksnes et al. 1989, Rasmussen & Giske 1994). Sea-
sonally fluctuating vertical distribution and abun-
dance of zooplankton appear to affect the migration
Prihartato et al.: Nocturnal distribution and behavior of Maurolicus muelleri
patterns of the mesopelagic fish in this system
(Staby et al. 2011, Dypvik et al. 2012b), yet as visual
predators, M. muelleri need sufficient light to see
their prey. The termination of midnight sinking
occurred when the minimum nocturnal surface light
exceeded 10−3 µmol m−2 s−1. We interpret the shift in
behavior as a sign that the pearlside now had suffi-
cient light for foraging throughout the night (Ras-
mussen & Giske 1994, Kaartvedt et al. 1998). The
light summer nights therefore represented a marked
increase in the time available for foraging in upper
waters (Fig. 8). This result suggests that summer
may be a particularly important period for growth,
not only due to higher plankton concentrations and
warmer surface waters at this time but also due to
the seasonal light cycle. The importance of the sea-
sonal light cycle at high latitudes has been reported
for fish in other settings. Suthers & Sundby (1996)
found enhanced growth rates of cod larvae at high
latitudes in summer, which they ascribed to the long
period for visual foraging at times with midnight
Schooling in light summer nights
As nights became even lighter toward mid-sum-
mer, M. muelleri took on schooling in upper layers.
This pattern occurred from early June until ceasing
the first half of July concordant with minimum noc-
turnal surface light above 5 ×10−3 µmol m−2 s−1 (Figs.
2, 6 & 7). The pearlside often schooled at 5 to 20 m
depth, with light intensities at these depths being 1 to
2 orders of magnitude lower than at the surface.
Schooling is well acknowledged as an anti-predator
strategy (Magurran 1990), and mesopelagic fish
taking up schooling behavior have previously been
reported for fish being chased by tuna during day-
light hours (Alverson 1961, Marchal & Lebourges
The change in nocturnal anti-predator behavior
shows behavioral flexibility, which may permit
extension of the oceanic habitat available for M.
muelleri. However, there are likely limitations for the
seasonal variation in light conditions that the meso-
pelagic fish can handle because they appear to be
scarce in the polar regions. Kaartvedt (2008) sug-
gested that this could relate to the extreme light
climate at high latitudes; very light summer nights
with midnight sun will prevent the fish from seeking
upper layers in cover of darkness, while total dark-
ness in winter hampers their feeding during that
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Editorial responsibility: Alejandro Gallego,
Aberdeen, UK
Submitted: May 12, 2014; Accepted: November 28, 2014
Proofs received from author(s): January 23, 2015
... Fish of the genus Maurolicus have a world-wide distribution (Rees et al., 2020) and are known for their relatively high abundance in the upper mesopelagic (Gauthier et al., 2014;Escobar-Flores, 2019). Pearlsides have a distinct vertical migration behaviour which is strongly influenced by season (Prihartato et al., 2015; and ontogeny (Giske et al., 1990;Baliño and Aksnes, 1993;Staby et al., 2013), and characterized by immediate reactions to changes in ambient light (Baliño and Aksnes, 1993;Staby and Aksnes, 2011), s.a. Supplementary Figure S1. ...
... We deployed three upward-looking SIMRAD EK60 split-beam echo sounders (7.1 beam angle) in Masfjorden ($60 50 0 N, $5 30 0 E), from 7 October 2010 to 15 August 2011 (s.a. Prihartato et al., 2015). The submerged transceivers were kept in pressureproof casings and cabled to a shore station for power supply and data storage. ...
... Surface photosynthetically active radiation (PAR, 400-700 nm) was measured continuously with a calibrated LI-190 quantum sensor (lower sensitivity threshold of 1 Â 10 À4 mmol m À2 s À1 ) from 10th December 2010 to the end of the study period (see Prihartato et al., 2015 for details). On five days in 2011 (26th January, 22nd February, 11th April, 16th June, and 16th August), water column PAR (400-700 nm, with a resolution of 3.3 nm) was measured around noon with a RAMSES ACC hyper-spectral radiometer (Trios-optical sensors, Oldenburg, Germany). ...
Variability of mesopelagic scattering layers is often attributed to environmental conditions or multi-species layer composition. Yet, little is known about variation in behaviour among the individuals forming scattering layers. Based on a 10 months high-resolution dataset from stationary echosounders in a Norwegian fjord, we here assess short-term and long-term behaviour of a single mesopelagic fish species, the pearlside Maurolicus muelleri. The daytime vertical extension of the monospecific pearlside scattering layers spanned four orders of magnitude ambient light in the autumn and winter and less than one order of magnitude in summer. While the main layers tracked relatively stable light levels over daytime, some individuals actively crossed light gradients of up to 1.5 orders of magnitude. This included individuals that moved between scattering layers, and apparently bold individuals that made regular upward excursions beyond the main population distribution. During the daytime, M. muelleri mitigated the risk of predation by forming tight groups in the upper scattering layer and, at light levels >10−6 µmol m−2 s−1, by instantly diving into deeper waters upon encounters with predators. Our observations suggest that individual, and probably state-dependent, decisions may extend the pearlsides’ vertical distribution, with implications for predator–prey interactions.
... Mesopelagic fish and other diel vertical migrants are usually expected to feed near the surface at night. How the organisms actually spend the night, though, varies between species (Simard et al., 1985;Giske et al., 1990;Balino and Aksnes, 1993;Pearre, 2003), seasons (Staby and Aksnes, 2011;Prihartato et al., 2015), and states (Simard et al., 1985;Pearre, 2003;Staby et al., 2012). Despite studies indicating this variability among mesopelagic fish (Torgersen, 2001;Kaartvedt et al., 2008;Olivar et al., 2017;Solberg and Kaartvedt, 2017), quantitative data of their individual behavior is largely lacking. ...
... Twilight migrations, where animals ascend to the ocean surface around dusk and dawn, but return to intermediate depths during night (midnight sinking) are common among pelagic taxa and repeatedly documented for acoustic scattering layers (Tarling et al., 2002;Valle-Levinson et al., 2014). Potential reasons for such behavior are manifold, but include reduced activity after satiation (Cushing, 1951;Rudjakov, 1970), temperature selection for energy optimization (Giske and Aksnes, 1992), prey distribution (Torgersen et al., 1997;Valle-Levinson et al., 2014) and predator avoidance (Torgersen et al., 1997;Staby, 2010;Staby and Aksnes, 2011;Prihartato et al., 2015). Still, we know very little about the individual behavior and activities resulting in such commonly observed population distribution patterns. ...
... Juveniles (<1 year) instead maximize growth by feeding on plankton in shallower waters during twilight (Giske et al., 1990;Bagøien et al., 2001). The juveniles form a shallow scattering layer with a strong diel migration pattern with ascent to the surface in the afternoon, subsequent midnight-sinking during a non-feeding period at night (Giske and Aksnes, 1992;Bagøien et al., 2001) and a dawn ascent in the morning before returning to daytime depth (Staby and Aksnes, 2011;Prihartato et al., 2015). Juveniles are a main prey for e.g., blue whiting and saithe (Giske et al., 1990). ...
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Nocturnal migration of mesopelagic fish into surface waters is well-documented. Yet, although there is increasing evidence of individual-based deviations from average population migrations and of the importance of small-scale individual behavior for predator-prey interactions and energetic cycling, little is known about what mesopelagic animals do when in upper waters. Using high-resolution data from an upward-facing, moored, split-beam echosounder we analyzed the night-time individual vertical swimming behavior of pearlsides (Maurolicus muelleri) over one winter. The population behavior is characterized by migration to the surface after sunset, “midnight-sinking” and another migration to the surface in the morning, followed by return to the daytime habitat. Focusing on individuals unveiled diverse behavioral patterns during different phases of the migration. After ascending to upper layers at dusk, M. muelleri leaves the surface waters, not by sinking, but by actively swimming in a step-wise pattern characterized by relocations alternated by pauses. Following the descent, vertical swimming is sustained at lower levels. Around midnight, the vertical swimming direction changes from predominantly downward to upward. Several hours before dawn, the fish start ascending toward the surface in a step-wise pattern. During population ascent in the afternoon and descent in the morning, some individuals at the fringes of schools migrate without intermittent pauses. This study documents the feasibility of using submerged, stationary echosounders in unveiling the individual behavior of mesopelagic fish.
... It is well established by numerous studies that seasonal rhythms govern various vital functions of fish (Bulow et al. 1981;Eliassen and Vahl 1982;Zapata et al. 1992;Gozdowska et al. 2006;Prihartato et al. 2015;etc.). Along with the alternation of seasons, temperate fish experience changes in mean temperature, amounts of oxygen, food availability large enough to present organisms with a physiological challenge (Clarke 1993). ...
... Summer GSH peaks appear to be not only GST-related. As the spawning is accomplished, the bream leaves the nearshore zones and moves to the deep in search for areas rich with benthos (Poddubny 1971). Active feeding governed by the need for energy reserve accumulation that will cover vital functions later when food consumption is ceased, stimulates intensive digestion. ...
... Toward the end of autumn, after abrupt decline of water temperature before the ice-covered period, the bream ceases feeding (Poddubny 1971). At this time, fish metabolism is adjusted to a stable state preparing for the wintering period (Shulman and Love 1999). ...
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... Micronekton are the dominant component of the SL organisms that inhabit the twilight zone, and they account for a significant proportion of DVM that feed on zooplankton near the surface. Their distribution can vary considerably depending on the time of day, area, season, and water mass characteristics (Prihartato et al., 2015;Annasawmy et al., 2019;Geoffroy et al., 2019;Cisewski et al., 2021). They are considered a potential harvestable resource because of their huge biomass and nutritional quality (protein and marine lipids), and some species can be suitable food sources for humans. ...
... In this study, MOCNESS (200 µm mesh size of the net) was used to target zooplankton. (Olivar et al., 2012;Dypvik and Kaartvedt, 2013;Gauthier et al., 2014;Davison et al., 2015;Prihartato et al., 2015;Gjøsaeter et al., 2017;Knutsen et al., 2017;Annasawmy et al., 2018Annasawmy et al., , 2019Annasawmy et al., , 2020. Each sampling gear has its fishing selectivity. ...
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In global oceans, ubiquitous and persistent sound scattering layers (SL) are frequently detected with echosounders. The southwest Indian Ocean has a unique feature, a region of significant upwelling known as the Seychelles-Chagos Thermocline Ridge (SCTR), which affects sea surface temperature and marine ecosystems. Despite their importance, sound SL within and beyond the SCTR are poorly understood. This study aimed to compare the characteristics of the sound SL within and beyond the SCTR in connection with environmental properties, and dominant zooplankton. To this end, the region north of the 12°S latitude in the survey area was defined as SCTR, and the region south of 12°S was defined as non-SCTR. The results indicated contrasting oceanographic properties based on the depth layers between SCTR and non-SCTR regions. Distribution dynamics of the sound SL differed between the two regions. In particular, the diel vertical migration pattern, acoustic scattering values, metrics, and positional properties of acoustic scatterers showed two distinct features. In addition, the density of zooplankton sampled was higher in SCTR than in the non-SCTR region. This is the first study to present bioacoustic and hydrographic water properties within and beyond the SCTR in the southwest Indian Ocean.
... However, our predictions of daytime foraging bouts into surface waters match observations where regular diel vertical migration patterns break up when daylight lasts for most of the day (Dietz, 1962). The model also does not allow for the emergence of alternative predator-avoidance behaviours, such as schooling, which has been observed for M. muelleri in light summer nights (Prihartato et al., 2015). Such behavioural adaptations could help extend distributions into more seasonal light environments. ...
... in the north-eastern basins of the Norwegian and Iceland Seas. An exception is Mueller's pearlside, Maurolicus muelleri (order: Stomiiformes), which is abundant in many fjords and near the coast of western Norway (e.g.Kaartvedt et al., 1998;Prihartato et al., 2015;Staby et al., 2013). Myctophids, exclusively represented by B. glaciale, are the dominant mesopelagic fish taxon in the Norwegian and Iceland Seas, but crustaceans make ...
Aim Mesopelagic fishes have a near-global distribution in the upper 1,000 m from tropical to sub-Arctic oceans across temperature regimes. Yet, their abundance decreases poleward and viable populations seem excluded from high latitudes. Why? Location North Atlantic between 50–85°N, with implications for high-latitude oceans globally. Time period Present-day. Major taxa studied Diel vertically migrating (DVM) mesopelagic fishes. Methods We use a mechanistic, state-dependent life-history model to characterize DVM mesopelagic fishes. This model links light-dependent encounters and temperature-dependent physiology, allowing optimal DVM strategies to emerge. We run the model along a latitudinal gradient with increasing seasonality in light and track individual fitness-related measures, that is, survival and surplus energy, through the annual cycle to make predictions about population consequences. Results Mesopelagic fishes thrive in the oceans’ twilight zone, and many are dependent on periods of darkness for safe foraging near the surface, before migrating back to depth during daytime. When daylight lasts for 24 hr during the Arctic summer, these fish are trapped in deep waters void of prey because it is never safe to forage in the shallow waters where zooplankton prey are found. Hence, they are left with two poor options, starvation at depth or depredation while foraging. Our model predicts surplus energy, vital for reproduction and growth, to halve from 50–85°N and annual survival to drop by two-thirds over a narrow range of 10° of latitude around the Arctic Circle. Thus, low recruitment and high predation mortality during summer make polar waters population sinks for mesopelagic fishes because of the extreme seasonality in light. Main conclusions At high latitudes, foraging mesopelagic fishes are exposed to sunlight in upper waters also at night. This makes them easy prey for visual predators, which limits their poleward distribution. Our findings highlight the importance to think beyond temperature to explain high-latitude range limits.
... They form daytime acoustic scattering layers that are shallower than what is normally termed as the DSL. Moreover, Maurolicus inhabits upper waters throughout the relatively light summer nights at high latitudes (Rasmussen and Giske, 1994;Prihartato et al., 2015). In darker nights, M. muelleri appears to stop feeding (Rasmussen and Giske, 1994), displaying "mid-night sinking" after migrating to surface waters at dusk and before a subsequent dawn ascent providing favourable light conditions (Prihartato et al., 2015). ...
... Moreover, Maurolicus inhabits upper waters throughout the relatively light summer nights at high latitudes (Rasmussen and Giske, 1994;Prihartato et al., 2015). In darker nights, M. muelleri appears to stop feeding (Rasmussen and Giske, 1994), displaying "mid-night sinking" after migrating to surface waters at dusk and before a subsequent dawn ascent providing favourable light conditions (Prihartato et al., 2015). So while e.g. ...
By definition, the mesopelagic twilight zone extends from 200 to 1000 m depth. Rather than confining the twilight zone to a certain depth interval, we here propose a definition that covers absolute light intensities ranging from 10−9 to 10−1 μmol quanta m−2 s−1. The lowest intensity of this twilight habitat corresponds to the visual threshold of lanternfishes (Myctophidae). The highest intensity corresponds to the upper light exposure of pearlsides (Maurolicus spp.), which have a unique eye adapted to higher light intensities than the lanternfishes. By this definition, the daytime twilight habitat extends deeper than 1000 m in very clear oceanic water, while may even be largely located above 200 m in very murky coastal waters. During moonlit nights in clear water, the twilight habitat would still extend deep into the mesopelagic depth zone, while becoming compressed toward the surface in dark nights. Large variation in night light, from 10−3 μmol quanta m−2 s−1 during moonlit nights to 10−8 μmol quanta m−2 s−1 in dark overcast nights, implies that division of light into night- and daylight is insufficient to characterize the habitats and distributional patterns of twilight organisms. Future research will benefit from in situ light measurements, during night- as well as daytime, and habitat classification based on optical properties in addition to depth. We suggest some pertinent research questions for future exploration of the twilight zone.
... copepods, krill, mesopelagic fish, squids, and jellyfish. There can be huge variations in realized DVM patterns, both spatially (Klevjer et al. 2016) and temporally (Prihartato et al. 2015), and are likely due to the different trade-offs faced by migrants (Ohman 1990;Ohman and Romagnan 2016;). The carbon draw-down mediated by actively migrating organisms has been widely explored, especially for zooplankton (Longhurst et al. 1990;Archibald et al. 2019;Kelly et al. 2019), and more recently also for fish (Davison et al. 2013;Pinti et al. 2021b) and jellyfish (Luo et al. 2020;Pinti et al. 2021b). ...
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The biological carbon pump transports photosynthetically fixed carbon from surface waters to depths. It removes carbon from the atmosphere and sequesters it in the deep ocean, playing an important role in global climate regulation. As the biological carbon pump is directly related to biological processes, it is heavily influenced by the biomass and trophic interactions between populations in the ecosystem. However, behavioral responses and adaptations to predation risk change trophic interactions, potentially having larger impacts than direct effects on trophic interactions and population abundances. Thus, predation risk may play an important role in shaping the biological carbon pump's strength (how much carbon leaves the euphotic zone) and efficiency (what fraction of detritus reaches a certain depth without being degraded). Except in the case of active carbon transport by vertically migrating organisms, this role of risk is not generally recognized. Here, we synthesize the existing knowledge on the consequences of anti‐predation responses on the biological carbon pump. First, we consider a generic anti‐predation response and investigate the different direct, indirect, and cascading effects that the response can induce. Then, we focus on pelagic anti‐predation responses and detail how they can specifically alter the different components of the pump. Finally, we discuss points to consider in biological carbon pump studies and highlight directions for future research. In particular, there is a need for more quantitative research to evaluate the importance of anti‐predation responses in shaping the biological carbon pump.
... During reverse DVM, animals occupy shallower water depths during the day and deeper waters at night (Martin and Christiansen, 2009;Dypvik et al., 2012). Twilight DVM involves an ascent from midwaters at sunset and a descent shortly after dusk, followed by a second ascent to the surface at dawn and subsequent descent 4 during the day (Valle-Levinson et al., 2014;Prihartato et al., 2015). Some migrant species do not migrate every day (Sutton and Hopkins, 1996; or at specific ontogenetic stages (Badcock and Merrett, 1976;Hulley, 1984). ...
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We examined the diel vertical migration (DVM) behavior and vertical spatial structure of sound-scattering layers (SLs) at two seamounts (Condor and Gigante) in the Azores and in surrounding open-waters. Active acoustic data were recorded day and night during nine cruises conducted in spring, summer and autumn between 2009 and 2011. SLs were permanent features with two main layers, shallow scattering layers (SSLs) and deep scattering layers (DSLs). Over seamount plateaus, SSLs aggregated close to the seafloor during the day and in slightly shallower waters at night. Backscatter intensity on plateaus varied little between day and night and was consistently higher than in SSLs above slopes or in open-waters. DSLs found over slopes and open-waters migrated towards the surface at dusk and returned to their daylight depths at dawn but an intense DSL persisted overnight in deep open-waters. Variograms showed that SSLs and DSLs were not uniformly distributed but amount of spatial heterogeneity varied between seamount plateaus, slopes and open-waters, and day-night periods. Taken together, these findings suggest that (1) Condor and Gigante Seamounts host a resident micronekton community, (2) there is an influx of vertically migrating organisms over the plateaus at night, and (3) seamount topography affects the DVM behavior of the DSL. Physical processes, such as Taylor cap effect, along with topographic blockage and lateral advection of migrating organisms, may play a role in transporting micronekton from open-waters to seamount plateaus. These findings are critical to understanding the ecology of seamount communities and the function of seamount habitats in oceanic ecosystems.
... However, if distributions are less tightly linked with the optical environment (Siegelman-Charbit and Planque, 2016) and cold waters currently constrain mesopelagic fish toward the poles (Proud et al., 2017); then mesopelagic fish may invade a future Arctic Ocean. There is some mesopelagic fish (Maurolicus muelleri) switching to schooling in upper layers during light Norwegian summer nights (Kaartvedt et al., 1998;Prihartato et al., 2015). Such behaviour might facilitate further northward extension if other conditions like temperature became more favourable. ...
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Climate change drives fish and plankton species ranges toward the poles, often related to warmer waters mediating geographic distributions via changes in vital rates. Yet, the distribution of fish may also be governed by less acknowledged mechanisms. Ice limits access to air for physostomous fish filling their swimbladders at the surface. We hypothesize that release of ice constraints may facilitate northward expansion of physostomes, with implied impact on their zooplankton prey. On the other hand, even in a changing Arctic, the extreme high-latitude photoperiod will persist. The abundance of mesopelagic fish is low in the Arctic Ocean. Feeding conditions may be inferior during the darkness of winter and in light summer nights. If the photoperiod is constraining distributions, biogeographic boundaries of mesopelagic fish may be relatively unaffected by climate change. Alternatively, if low temperatures are their main constraint, we hypothesize that northward extensions in a warmer ocean may be detrimental to key Arctic copepods as we argue that their current success relates to low mortality during overwintering in the absence of mesopelagic fish. It is therefore essential to discriminate the role of the light climate at high latitudes from those related to temperatures for assessing future biogeographic boundaries. © International Council for the Exploration of the Sea 2018. All rights reserved.
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Diel vertical migration of fish and other metazoans actively transports organic carbon from the ocean surface to depth, contributing to the biological carbon pump. Here, we use a global vertical migration model to estimate global carbon fluxes and sequestration by fish and metazoans due to respiration, fecal pellets, and deadfalls. We estimate that fish and metazoans contribute 5.2 PgC/yr (2.1-8.8PgC/yr) to passive export out of the euphotic zone. Together with active transport, we estimate that fish are responsible for 20% (9-29%) of global carbon export, and 32% (18-43%) of oceanic carbon sequestration, with forage and deep-dwelling mesopelagic fish contributing the most. This essential ecosystem service could be at risk from unregulated fishing on the high seas.
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A cycle of zooplankton density that fluctuated in phase with the moon was observed throughout 1982-1983 in Cahora Bassa Reservoir on the lower Zambezi, in southeastern Africa. Despite constant birth rates, densities of four cladoceran and two copepod species, as determined from vertically hauled plankton net samples taken every 2-6 d, fluctuated over one order of magnitude. The pattern followed by each species included an exponential increase in population density from the last quarter of the moon through the new moon and the first quarter, till the full moon, then a sudden decrease resulting in lowest numbers during the moon's last quarter. The cycle was shown to be induced by predation. Much higher death rates between the full moon and the last quarter were caused by the abundant Tanganyikan sardine Limnothrissa miodon. As seen from an examination of gut contents, sardines crop zooplankton most efficiently on nights when the full or nearly full moon rises after sunset, i.e., when zooplankton approach the surface during darkness and become suddenly vulnerable in the first light of the rising moon. After the last quarter, zooplankton density is low, the moon gives little light, the fish shift to alternate food resources, and zooplankton populations grow exponentially again. I suggest that the moon phase cycle in zooplankton is a global phenomenon, but, previously uninterpreted, has been seen only as distracting @'random@' variations in seasonal density patterns. I also suggest that similar prey-predator interactions might have been responsible for selecting for and fixing intrinsic monthly rhythms in behavior and physiology of animals with long life-spans.
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Marine ecosystem acoustics (MEA): quantifying processes in the sea at the spatio-temporal scales on which they occur. – ICES Journal of Marine Science, doi: 10.1093/icesjms/fsu116. Sustainable management of fisheries resources requires quantitative knowledge and understanding of species distribution, abundance, and prod-uctivity-determining processes. Conventional sampling by physical capture is inconsistent with the spatial and temporal scales on which many of these processes occur. In contrast, acoustic observations can be obtained on spatial scales from centimetres to ocean basins, and temporal scales from seconds to seasons. The concept of marine ecosystem acoustics (MEA) is founded on the basic capability of acoustics to detect, classify, and quantify organisms and biological and physical heterogeneities in the water column. Acoustics observations integrate operational technologies, platforms, and models and can generate information by taxon at the relevant scales. The gaps between single-species assessment and ecosys-tem-based management, as well as between fisheries oceanography and ecology, are thereby bridged. The MEA concept combines state-of-the-art acoustic technology with advanced operational capabilities and tailored modelling integrated into a flexible tool for ecosystem research and monitoring. Case studies are presented to illustrate application of the MEA concept in quantification of biophysical coupling, patchiness of organisms, predator– prey interactions, and fish stock recruitment processes. Widespread implementation of MEA will have a large impact on marine monitoring and assessment practices and it is to be hoped that they also promote and facilitate interaction among disciplines within the marine sciences.
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Throughout the ocean, countless small animals swim to depth in the daytime, presumably to seek refuge from large predators. These animals return to the surface at night to feed. This substantial diel vertical migration can result in the transfer of significant amounts of carbon and nutrients from the surface to depth. However, its consequences on ocean chemistry at the global scale have remained uncertain. Here, we determine the depths of these diel migrations in the global ocean using a global array of backscatter data from acoustic Doppler current profilers, collected between 1990 and 2011. We show that the depth of diel migration follows coherent large-scale patterns. We find that migration depth is greater where subsurface oxygen concentrations are high, such that seawater oxygen concentration is the best single predictor of migration depth at the global scale. In oxygen minimum zone areas, migratory animals generally descend as far as the upper margins of the low-oxygen waters. Using an ocean biogeochemical model coupled to a general circulation model, we show that by focusing oxygen consumption in poorly ventilated regions of the upper ocean, diel vertical migration intensifies oxygen depletion in the upper margin of oxygen minimum zones. We suggest that future changes in the extent of oxygen minimum zones could alter the migratory depths of marine organisms, with consequences for marine biogeochemistry, food webs and fisheries.
Fish should respond to low levels of food availability by altering their behavior to ensure higher feeding rates, larger feeding territories and broader diets. The gastric sensation of hunger and its rate of change may act as appropriate cues to food availability; hunger-motivated changes in feeding behavior can produce all of these predicted effects. Juvenile coho salmon Oncorhynchus kisutch alter their behavior in an adaptive manner when faced with variable degrees of threat of competition from territorial intruders, and of risk of predation. Learning is an important mechanism providing behavioral flexibility, and changes in fish feeding behavior with experience are summarized. A graphical model shows that these changes can result in training biases and food specialization. Learning also results in increased feeding rates.-from Author
Diel vertical migration (DVM) is a characteristic behavioural pattern performed by zoo- plankton in which their vertical distribution changes over the 24-h day. Here the proximate control of zooplankton DVM is reviewed. Light has emerged as the major proximate cue controlling DVM behaviour and the understanding of zooplankton visual physiology and the light-mediated behaviour underlying DVM is expanding. Field and laboratory evidence exist to support each of the three major hypotheses for the exogenous role of light in DVM: (1) preferendum or isolume, (2) absolute intensity threshold, and (3) relative rate of change. Light may also play an endogenous role in DVM by entraining circadian rhythms in vertical movement or activity. This appreciation of the role of light has improved modelling efforts into the causes and consequences of DVM. The most important recent advance in the study of DVM is the recognition that this behaviour is a phenotypic response in many species and is most commonly activated by chemical cues (kairomones) from fish predators. High levels of kairomones signal high levels of predation pressure, and DVM-related photobehaviours, such as swimming responses on relative rates of irradiance change, are altered such that migration occurs and zooplankton achieve a refuge from visual predators. © R.N. Gibson, R.J.A. Atkinson, and J.M.D. Gordon, Editors Talyor & Francis.
Mesopelagic fishes occur in all the world's oceans, but their abundance and consequently their ecological significance remains uncertain. The current global estimate based on net sampling prior to 1980 suggests a global abundance of one gigatonne (10(9) t) wet weight. Here we report novel evidence of efficient avoidance of such sampling by the most common myctophid fish in the Northern Atlantic, i.e. Benthosema glaciale. We reason that similar avoidance of nets may explain consistently higher acoustic abundance estimates of mesopelagic fish from different parts of the world's oceans. It appears that meso pelagic fish abundance may be underestimated by one order of magnitude, suggesting that the role of mesopelagic fish in the oceans might need to be revised.
The plankton outburst during the so-called late winter bloom in subtropical waters was studied in relation to lunar illumination in the Canary Island waters. Nutrient enrichment by mixing and dust deposition promoted a bloom of phyto- and zooplankton. Mesozooplankton biomass increased as the winter mixing progressed but peaked in every full moon and decreased thereafter because of the effect of predation by interzonal diel vertical migrants (DVMs). The pattern was similar to the one described in lakes due to predation by fishes and confirms that this phenomenon is important in the sea. The estimated consumption and subsequent transport of epipelagic zooplankton biomass by DVMs after every full moon is on the order of the mean gravitational export and is an unaccounted flux of carbon to the mesopelagic zone that may play a pivotal role in the efficiency of the biological pump.
In the Norwegian fjord Masfjorden, different developmental stages of the mesopelagic planktivore Maurolicus muelleri form vertically separate sound scattering layers (SSLs) and in late autumn display different diel vertical migration (DVM) behaviour. Post-larvae and juvenile fish perform normal crepuscular DVM, whereas the large majority of adults remain at depth throughout the diel period. In this study we examined the stomach contents of juvenile and adult fish caught at different times and depths during a 24-h period in autumn. The different DVM behaviour of these two SSLs in addition to a shallow layer believed to be composed of post-larvae is explained with a model for visual foraging in aquatic environments that uses gradients in vertical light intensity and copepod density and size as input variables. Field data revealed that vertically migrating juveniles distributed at a higher ambient light intensity and on average consumed 25 times more copepods than non-migrating adult fish. The model showed that juveniles experienced a 15 times higher prey encounter rate and a higher level of predation risk than non-migrating adults, and that the energetic benefits for post larvae and juveniles from prolonged feeding in a nearly constant and brighter environment outweigh the associated predation risk. The model also suggests that the visual detection range of piscivore predators is relatively more limited by the turbid surface water than that of their prey, which provide the post-larva and juvenile life-stages of M. muelleri a window of reduced visual predation near the surface.