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Intercomparison of zooplankton (net) sampling systems: Results from the

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Progress in Oceanography 108 (2013) 1–42

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Progress in Oceanography
journal homepage: www.elsevier.com/locate/pocean

Intercomparison of zooplankton (net) sampling systems: Results from

the ICES/GLOBEC sea-going workshop
Hein Rune Skjoldal a, Peter H. Wiebe b,⇑, Lutz Postel c, Tor Knutsen a, Stein Kaartvedt d,1, Douglas D. Sameoto e
a
Institute of Marine Research, PO Box 1870, N-5017 Bergen-Nordnes, Norway
b
Woods Hole Oceanographic Institution, Woods Hole, MA, USA
c
Leibniz Institute for Baltic Sea Research, Warnemünde, Germany
d
Department of Biology, University of Oslo, PO Box 1066 Blindern, 0316 Oslo, Norway
e
Bedford Institute of Oceanography, Bedford, Nova Scotia, Canada

a r t i c l e i n f o a b s t r a c t

Article history: An inter-comparison and evaluation of methods for sampling and determination of zooplankton distribu-
Available online 29 October 2012 tion and biomass was conducted in a fjord environment (Storfjorden at Møre, Norway) from 2 to 13 June
1993. The inter-comparison was carried out with the German R/V ‘‘A. v. Humboldt’’ and the Norwegian
R/V ‘‘Johan Hjort’’ and involved a total number of 38 scientific personnel from 8 countries. The inter-com-
parisons included the MOCNESS, BIONESS, MultiNet, LHPR, OPC, CPR, Gulf-V, CalCOFI 1-m Ring Net, and
WP-2 net. In addition, acoustics data were collected with a Simrad EK500 echosounder operating on 4
frequencies (18, 38, 120, 200 kHz) and hydrographic, nutrient, phytoplankton, and meteorological data
were collected to characterize the environment in which the comparisons were made; environmental
changes were minor during the study.
The results of this study corroborate the results of earlier studies. Mesh size of the net had a major
influence on the biomass and species composition of the zooplankton community. There is a consistent
relationship between retention (or escapement through the mesh) and the width of the organisms; about
50% of the organisms escape through the mesh at a width equal to the mesh size. The effect of towing
speed on the extrusion of smaller organisms through the net can be substantial and adds to the loss
due to escapement. Active avoidance of the sampler is only important for the larger macrozooplankton
and not a significant problem for the mesozooplankton. Different vertical, oblique, and multiple open-
ing/closing net systems produced similar estimates of zooplankton when operated with comparable
mesh-sized nets and a sufficiently high mesh open area to mouth opening ratio. The choice of mesh size
is important and 150 lm mesh is possibly an optimal mesh size for use in coastal waters with neritic zoo-
plankton communities. Replicate tow variability was also consistent with earlier studies and reflects the
need to understand the interplay between the underlying patchiness of the zooplankton distributions
and the method of sampling. No single net is suitable to sample across the wide size range of zooplankton
from small mesozooplankton to macrozooplankton. Recent large interdisciplinary programs to assess
marine ecosystem structure and dynamics have recognized this through the use of nets designed to sam-
ple particular size fractions in combination with video and acoustic remote sensing techniques.
Ó 2012 Elsevier Ltd. All rights reserved.

1. Introduction oceanography (Wiebe and Benfield, 2003). Zooplankton sampling

Zooplankton constitutes a diverse group of animals that occur
suspended in the water column. They span a wide range in size
and swimming ability. Quantitative sampling of zooplankton has
posed a long-standing challenge and a large number of sampling
gear has been constructed up through the history of modern
⇑ Corresponding author.
E-mail address: pwiebe@whoi.edu (P.H. Wiebe).
1
Present address: King Abdullah University of Science and Technology, Red Sea
Research Center, Thuwal 23955-6900, Saudi Arabia.
variation and the evaluation and intercomparison of zooplankton
sampling equipment has taken place since quantitative sampling
techniques were introduced by Hensen (1887; 1895; Unesco,
1968; See Table 2 in Wiebe and Benfield (2003) for a summary
of comparisons). The primary sources of error are escapement,
avoidance, and patchiness.
Early on it was recognized that escapement of organisms
through the net mesh was due either to excess filtration pressure
causing extrusion or to the organisms being smaller than the mesh
opening (Kofoid, 1897; Lohmann, 1903). Small individuals may
pass through the net mesh and large individuals may sense and

0079-6611/$ - see front matter Ó 2012 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.pocean.2012.10.006
 Author's personal copy
2 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
escape the approaching net. Reducing the mesh size may increase
the retention of small organisms, but may lower the filtration effi-
ciency and increase the avoidance of large organisms. Avoidance of
the net mouth opening by larger, faster swimming animals also has
been recognized as a significant problem for decades (Clutter and
Anraku, 1968; Sameoto et al., 2000; Wiebe et al., 2004). Increasing
the towing speed may improve the catch efficiency of larger zoo-
plankton, but results in greater loss of small individuals by extru-
sion through the mesh of the net. There is thus a compromise
between mesh size and speed for the optimal sampling of a given
size range or group of zooplankton.
The importance of the non-random distribution of plankton in
the sea was debated for years before it was finally demonstrated
that ocean animal and plant distributions are patchy and the
patchiness is a major source of replicate tow variability (Herdman,
1921; Barnes and Marshall, 1951; Anraku, 1956; Wiebe and Hol-
land, 1968). Reducing error associated with plankton patchiness
is a matter of sampling design (Wiebe, 1971; Skjoldal et al., 2000).
It is recognized that harmonization and standardisation of
methodology is important to allow comparisons across different
ocean areas and time periods. This is particularly important in rela-
tion to investigations of global patterns and regional characterisa-
tions in global programmes such as GLOBEC (Global Ocean
Ecosystem Dynamics) and GOOS (Global Ocean Observing System).
Thus, ICES established in 1992 a Study Group on Zooplankton Pro-
duction (SGZP) that was transformed in 1994 into the Working
Group on Zooplankton Ecology (WGZE). The initial terms of refer-
ence were to address improvements and standardization of meth-
ods and to consider workshops to intercalibrate and evaluate
methods. The work resulted in the production of the ICES Zoo-
plankton Methodology Manual in 2000 (Harris et al., 2000).
A sea-going workshop was arranged from 2 to 13 June 1993 in
Storfjorden, Norway (Skjoldal et al., 1993; Fig. 1). The workshop
was a two-ship operation involving the Norwegian R/V Johan Hjort
and the German R/V A.v. Humboldt. The principal objective was to
intercompare, characterize, and evaluate the performance of gear
and techniques for quantitative description of zooplankton distri-
bution, biomass, and production. A total of 38 scientists and tech-
nicians from Canada, France, Germany, Iceland, Norway, Spain,
United Kingdom, and USA took part in the workshop. A wide range
of sampling gear and instruments was deployed.
This paper considers the comparisons of the WP2, Bongo,
BIONESS, MOCNESS, LHPR, MultiNet, Gulf-V, and CalCOFI net sys-
tems, especially with regard to collection of zooplankton biomass
data. All the underlying and supporting data including acoustic
data collected with echosounders, have been published in a
report with four CD-ROMs (Wiebe et al., 2002). Although the
study took place some time ago, the sampling gear used in the
intercomparison are those currently used in zooplankton research
globally and no other comprehensive comparison has been done.
Thus the results are important for ongoing and future zooplank-
ton research.
2. Methods
2.1. The site and sampling strategy
Storfjorden is a long and deep fjord located at Møre on the west
coast of Norway (Fig. 1). Storfjorden was in many respects ideal. It
was chosen as the site for the workshop due to its proximity to the
Norwegian Sea and the similarity in fauna, and because there ex-
isted a fair amount of background information from previous
investigations. Also, Storfjorden, like most Norwegian fjords, is
well protected from winds, and seas remained flat for the duration
of the cruise. This made handling of gear over the side and stern of
the vessels relatively easy. Thus for the 10-days duration of the
study variations between sampling devices were not a result of
poor sampling conditions. The sampling furthermore benefited
by occurring within a water column in which physical and biolog-
ical conditions changed only a little during the study.
A 5 nautical miles long sampling transect was chosen in the
outer part of Storfjorden where the bottom depth was about
400 m (62°23.80 N; 06°20.00 E–62°25.10 N; 06°30.50 E). The majority
of the work was carried out along this transect. R/V Johan Hjort
worked this section repeatedly (about 65 times) while doing
acoustical recordings at 4 frequencies and towing various sampling
gear at a series of stations numbered 386–396. R/V A.v. Humboldt
also worked this section repeatedly towing sampling gear. Further
details on the sampling stations and the samples collected are
found in Wiebe et al. (2002).
Sampling stations along the transect were occupied for deploy-
ment of vertically hauled zooplankton nets. R/V A.v. Humboldt also
conducted vertical profiling with a CTD equipped with fluores-
cence and oxygen sensors and a water bottle rosette for water sam-
pling. Vertical light profiles were obtained with a spectral
radiometer and continuous surface light measurements were made
with a pyroheliometer from the R/V Johan Hjort.
Zooplankton sampling was carried out for three main purposes:
– Description of the zooplankton and micronekton communities.
– Intercomparison of sampling gears.
– Studies of sampling variance and effect of net mesh size.
Zooplankton sampling events occurred over each 24 h day-
night period from 4 to 12 June 1993 (Fig. 2). Also shown are the
depth intervals sampled with the various gears.
Information on the zooplankton and micronekton communities
was obtained with echo sounders. Although the acoustical data are
quantitative, the information provided about the biological proper-
ties of the volume backscatterers is of a qualitative nature and
needs to be supplemented by zooplankton sampling. Exploratory
sampling was carried out on the first day (4 June) using different
zooplankton and micronekton nets towed around an acoustical
reflectance layer at about 160 m depth. In addition, tows were
made over the upper 200 m. The information gained from inspec-
tion of these samples guided the further design of the sampling
and experimental programme. For the overall description of the
zooplankton and micronekton communities in this paper, we have
used all available data including those from gear intercomparisons
or special studies.
The main gear intercomparison experiments were carried out
on 5, 6, 10, and 11 June. On 5 June, the whole water column from
about 400 m was sampled with various towed or vertically hauled
zooplankton nets (Fig. 2). On 6 June, comprehensive sampling was
carried out over the upper 100 m where most of the zooplankton
biomass was concentrated. This included repeated sampling with
a multiple net system (MOCNESS) with 6 hauls during the day
and 4 hauls during the night periods. On 10 June, intercomparison
of two multiple net systems (MOCNESS and BIONESS) was carried
out by repeated hauls between 200 and 300 m depth during day
and night periods, aiming to compare their efficiency in sampling
macrozooplankton that were expected to reside deep in the water
column during the day. On 11 June, repeated sampling over the
400 m water column was carried out during day and night periods.
For some analyses we have grouped data from this latter period
(10–12 June).
Supplementary sampling and intercomparison between Gulf-V
and Optical Plankton Counter (OPC) was carried out on 7 June. R/
V Johan Hjort was on another mission on 8–9 June. During this per-
iod R/V A. v. Humboldt carried out special studies on replicate var-
iance and effect of net mesh size.
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42 3

Fig. 1. Map of the study site showing the location of Storfjorden in Norway, and more detailed map with the sampling transects and station locations (CTD stations taken
from R/V A.v. Humboldt). Also plotted are the 100, 200, and 400 m depth contours. The zooplankton samples for gear intercomparisons were obtained along the 5 nm transect
(located between stations 9006 and 9007).

2.2. Zooplankton sampling gear
A total of 14 different zooplankton sampling instruments were
used in the workshop. These included four multiple opening and
closing net systems (1- and 10-m2 MOCNESS, 1-m2 BIONESS,
0.25-m2 MultiNet), three high-speed samplers (LHPR, Gulf-V,
CPR), one optical plankton counter (OPC), 4 vertically or obliquely
hauled nets (WP-2, 20- and 60-cm Bongo, 1-m Ring net (CalCOFI)),
and 2 micronekton trawls (MIK ring net, young-fish trawl). The
data from the CPR, OPC, 10-m2 MOCNESS and the 2 micronekton
trawls are not used in this analysis. Results of the OPC have been
reported by Wieland et al. (1997) and Halliday et al. (2001).
Some key characteristics of these sampling gears are summa-
rized in Table 1. Information on the deployment of the various
gears with respect to time, vertical range, and ship are given in
Fig. 2. Illustrations of the gears are given in Fig. 3. Further details
on the sampling gear and their deployment are given in the
following.
2.2.1. 1-m2 MOCNESS
The MOCNESS (Multiple Opening/Closing Net and Environmen-
tal Sensing System; Wiebe et al., 1976, 1985) is a net system based
on the Tucker Trawl principle (Tucker, 1951). The 1-m2 MOCNESS
was equipped with a 12-bit electronics system and weighed about
150 kg. It was operated on R/V Johan Hjort and deployed off the
stern of the vessel. Most of the tows were made with nets of
180 lm mesh netting. One series of tows was made with nets of
333 lm netting. The MOCNESS was towed obliquely with nets
sampling predetermined depth intervals. On 10 June, it was towed
up and down (towyoed) to obtain replicate samples from depth
intervals between 200 and 300 m (Fig. 2).
2.2.2. 1-m2 BIONESS
The BIONESS (Bedford Institute of Oceanography Net and Envi-
ronmental Sensing System) is a multiple net opening and closing
zooplankton sampler originally described by Sameoto et al.
(1979, 1980). It is a modern version of the net system originally de-
scribed by Bé et al. (1959). The nets on the BIONESS are arranged
horizontally one behind the other and are opened sequentially
with one net opening as one is closed. Each of the ten horizontal
dropping bars is lead filled and weighs approximately 25 kg. The
entire sampler has a mass in air of about 800 kg. The BIONESS
was operated from R/V Johan Hjort and deployed off the stern of
the vessel. The nets were made with 333 lm mesh netting. The
towing speed was 3–4 knots, compared to 1–2 knots for MOCNESS.
Otherwise the BIONESS was operated similar to MOCNESS, with
oblique and towyo tows. It was also equipped with a factory cali-
brated General Oceanics flow meter, pressure, and angle sensors,
and had data transmission to the control unit via conducting cable.
2.2.3. MultiNet
The MultiNet (HYDRO-BIOS Apparatebau GmbH, Kiel, Germany;
Weikert and John, 1981) is also a modern version of the Bé et al.
(1959) net system. The sampler consisted of a net frame with an
opening area of 0.25 m2, a pressure compensated motor unit
(3000 dbar) with battery housing, 5 nets (length 2.5 m, diameter
at the end 11 cm) with zip fasteners, five plastic net buckets with
side windows covered with sieve gauze, a V-fin depressor
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4 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42

Fig. 2. Time table of sampling events with gears and vertical ranges identified. BIO – BIONESS; MOC – MOCNESS; LHPR – Longhurst-Hardy Plankton Recorder; OPC – Optical
Plankton Counter; CPR – Continuous Plankton Recorder. See Table 1 for more information on the gears. Samples obtained with a young-fish trawl (YT), a MIK Ring-Net (MIK),
10-m2 MOCNESS (MOC-10), and a Hufsa pump are not presented here.

(22 kg), and a deck unit. The frame and bucket holder weighs
150 kg. It was operated in oblique tows from R/V A.v. Humboldt.
Depth was recorded online by an OTS CTD probe (ME Meerestech-
nik-Elektronik GmbH, Trappenkamp, Germany), which was
mounted on the net frame. The MultiNet was alternatively
equipped with 5 nets with 200, 100, or 55 lm mesh. The nets were
equipped with mechanical digital flow-meter. The number of rev-
olutions, multiplied by a calibration factor of 0.3 m per propeller
revolution and the net opening area gave the filtered amount of
water. The calibration factor is a linear function of towing speed
>0.5 kn. Comparisons of results with the towed distance, measured
by GPS on the ship, allowed a first order quality control. The open-
ing/closing device made it possible to collect plankton in five dif-
ferent depth levels during each oblique haul. The ship velocity
was 1.5 kn (0.7 m s1) and the winch hauling speed was
0.3 m s1, which resulted in a total speed for the net of about
1 m s1.
2.2.4. LHPR
The LHPR (Longhurst-Hardy Plankton Recorder; Longhurst
et al., 1966) is a towed plankton sampler (up to 6 knots) that takes
a series of samples on a single haul for studies of vertical or hori-
zontal distribution (e.g. Haury et al., 1976; Williams et al., 1983).
The system used in the workshop consisted of a modified
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
Fig. 3. Illustrations of the zooplankton samplers that were included in the sampling gear intercomparisons.
high-speed net frame (2.5 m in length, 76 cm diameter) in which a
conical net (200 lm mesh aperture) was mounted that ended in a
recorder unit. Above this net and frame was a fine mesh (53 lm)
conical net that also ended in a recorder box. The recorder unit
contained two rolls of filtering gauze which were threaded across
an intake tunnel and onto a single take-up spool. The gauze was
advanced by an electric motor at discrete time intervals to give a
sequential series of samples. A flowmeter mounted in a conical
nose-cone on the front of each net enabled volume filtered for each
sample to be determined. With a coarse mesh net inlet aperture of
35.6 cm diameter and a speed of about 3 knots, a 2 min advance
interval resulted in about 20 m3 of water being filtered. With a fine
mesh net inlet of 5.0 cm, volume filtered in 2 min was about 400 l.
The LHPR sampling was carried out from R/V A.v. Humboldt
(Halliday et al., 2001). Seven hauls were completed taking a total
of 175 samples to a maximum of 351 m depth. The Optical Plankton
Counter (OPC) was fitted to the LHPR on all hauls giving a data set
for comparison with the analysis results from the LHPR samples.
2.2.5. Gulf-V
The high speed plankton sampler ‘‘Nackthai’’ (Nellen and Hem-
pel, 1969), also known as Gulf-V sampler (Sameoto et al., 2000), is
5
a modified version of the Gulf III (Gehringer, 1952). The total
length of the case less equipment was about 240 cm. The conical
net had a length of 135 cm, with an upper diameter of 40 cm and
a lower diameter of 11 cm. The 20 cm diameter nose cone that
delimited the entrance is intended to give a high filtration effi-
ciency for the 300 lm mesh net. A mechanical GO-flowmeter (Gen-
eral Oceanics, Miami, FL, USA) was installed in the center of the net
opening area. The Gulf-V sampler was operated from R/V A.v. Hum-
boldt and used in combination with an Optical Plankton Counter
(OPC). It was towed in oblique hauls at a speed between 3 and 5
kn using a V-fin depressor of 22 kg mass.
2.2.6. WP-2 net
The WP-2 net was described in detail in the UNESCO zooplank-
ton manual (Working Party 2, 1968; see also Fraser, 1966). In many
respects, it is a modified Nansen Net with a cylindrical front sec-
tion of 95 cm length and a conical section of 166 cm length. The
net ring has a diameter of 57 cm (0.255 m2) and the nets normally
used in this study were 200 lm mesh for which it was originally
designed. For mesh comparison studies, the WP-2 was additionally
equipped with nets of 55 lm, 100 lm, and 400 lm. All the WP-2
nets had been exclusively manufactured for this exercise. Net
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6 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
material consisted of a simple locking weaved monofilament poly-
ester. The porosity of the 200 lm mesh of 50% was in the range of
the 55% stipulated by the ICES/SCOR/UNESCO working group on
standardization of zooplankton sampling methods at sea (Working
Party 2, 1968). It decreased to 37% in case of the 55 lm mesh
(Table 1).
The net was vertically lowered at 60 m min1 and towed at
45 m min1 as recommended (Working Party 2, 1968). The volume
of water filtered was estimated by use of pre-calibrated TSK (Tsu-
rumi-Seiki-Kosakusho, Co.) flow meter positioned ca. 14 cm from
the net rim (1/2 radius). On part of the hauls, a second flow meter
was placed in equivalent distance outside the rim. The ratio be-
tween the flow meters inside and outside the net opening gave
an estimate of the integrated filtration efficiency. It did not drop
below 90% in an experiment of repeated WP 2 hauls using a mesh
size of 200 lm. It decreased to 70% when using the WP 2 equipped
with 55 lm aperture in a mesh size experiment, indicating that
some clogging of the mesh was taking place.
Stratified sampling was performed applying a messenger-
releasable closing mechanism. The depth of the nets was estimated
by wire length used. Wire angles did not occur due to calm weath-
er conditions and use of a lead mass of 30 kg. A dual version of the
WP-2 net with two nets mounted side by side in a frame, was used
in one experiment to characterize replicate sampling variability.
2.2.7. Bongo nets, 20- and 60-cm diameter
The original Bongo Net was an opening/closing sampler that
used a double messenger system (McGowan and Brown, 1966)
and had 60 cm net diameter (Fig. 3). A Bongo system used here
was described by Posgay and Marak (1980). The two systems used
in this comparison were non-opening/closing dual net systems
with two nets mounted side by side in a metal frame. One system
had net opening of 20 cm diameter and the other 60 cm diameter.
Both types had the same cod end bucket with 11 cm diameter (Hy-
dro-Bios Apparatebau Kiel, Germany) and nets of 180 lm mesh.
The two Bongo net systems were operated from R/V Johan Hjort
fastened one above the other on the towing wire. They were towed
obliquely from the surface to maximum depth and then back to the
surface applying a 22 kg V-Fin depressor. Depth was determined
with a SCANMAR depth sensor that transmitted depth recordings
acoustically to a display on the ship. The amount of water filtered
was estimated using a Hydro-Bios digital flow-meter attached in
the center of the net mouth.
2.2.8. 1-m Ring net (CalCOFI)
The 1-m Ring net was constructed by Hydro-Bios Apparatebau
Kiel, Germany, and is similar to the 1-meter standard CalCOFI
plankton sampler (Ahlstrom, 1948). It consisted of a 1-m diameter
ring of stainless steel tubing, to which a cylinder and cone-shaped
net was attached. The net was equipped with 333 lm mesh nylon
gauze, similar to what the CalCOFI program used for many years.
The length of the cylindrical front part of the net was 1 m while
the conical part was 3 m. The diameter of the cod-end was
11 cm. The net was operated from R/V A.v. Humboldt with oblique
hauls using a V-Fin depressor of 22 kg mass. While the ship was
making 1–2 knots, a fixed amount of towing cable was paid out.
The net was held at depth for 30 s and then retrieved at a constant
rate (20 m min1). Ship speed was adjusted to maintain the towing
cable at a constant angle of approximately 45° to the vertical. The
amount of filtered water was estimated using a Hydro-Bios digital
flow-meter attached at the center of the net entrance.
2.3. Zooplankton sample treatment and analyses
A standard set of procedures to process the zooplankton sam-
ples were used throughout the cruise on both ships (Fig. 4). On
deck, nets were washed and the cod-end buckets were taken into
the ship’s laboratory.
The samples were divided with a Folsom plankton divider
(McEwan et al., 1954) on the R/V Humboldt and with a Motoda
splitter (Motoda, 1959) on the R/V Johan-Hjort into two fractions,
one preserved in a buffered 4% formalin solution for later species
identification and the second one for dry mass measurements.
Splitting errors for these two plankton splitters were similar in a
study by Van Guelpen et al. (1982). This standard procedure was
used for most samples except those from LHPR.
For most of the hauls, the LHPR samples from each depth inter-
val were preserved with formalin for subsequent species enumer-
ation and size measurements, as described by Halliday et al.
(2001). The samples from one haul (haul 6) were rinsed off the
plankton gauze and size fractioned, dried, and weighed following
the standard procedure described above.
2.3.1. Determination of dry mass
Although the current literature often refers to ‘‘dry weight’’ of
zooplankton, the correct term is ‘‘dry mass’’ according to the inter-
national agreement on units (S.I. = Système International d’Unités).
Following the lead of Postel et al. (2000), we use the term ‘‘mass’’ in
this paper. The sample portion for dry mass determination was
examined and individual krill, shrimps, and fish were removed.
Following species identification and length measurements, they
were transferred to 3 pre-weighed aluminum trays for krill,
shrimps, and fish, respectively. In some cases, these groups were
also picked out from the second sample portion prior to formalin
preservation and included with the specimens for biomass
determination.
The rest of the sample fraction was separated into 3 size frac-
tions by subsequent wet sieving through plankton mesh of 2000,
1000, and 180 lm. The sieves were Plexiglas cylinders (about
20 cm diameter and 10 cm height) with plankton gauze mounted
as a flat bottom. The sample was poured into the sieves and
washed by swirling, shaking, and lifting in a tray filled with about
2 l of seawater. This seawater containing the zooplankton that
passed through the coarser sieve was poured into the next sieve
and the process repeated. The zooplankton retained in the 3 sieves
were quickly rinsed with fresh water from a rinse bottle, and trans-
ferred with the help of a flat spoon into pre-weighed aluminum
trays.
The aluminum trays with zooplankton samples were frozen in a
freezer at about 20 C. Periodically during the course of the work-
shop, the samples were picked up by a chartered boat and taken to
a laboratory in a nearby technical institute (Ålesund University
College – AAUC). Here the samples were dried at about 60 C for
24 h or longer. After drying the samples were stored in a box with
desiccators and subsequently weighed.
2.3.2. Taxonomic analyses
The formalin-preserved samples have been analyzed by some-
what different procedures by 3 laboratories.
Selected samples from MOCNESS and BIONESS have been ana-
lyzed at the Institute of Marine Research (IMR) in Bergen, Norway,
following the routine procedure for taxonomical analysis at this
laboratory. This included identification to species or genus for most
plankton groups, with separate enumeration of copepodite stages
for Calanus spp. and some other calanoid copepods, and size groups
of krill, amphipods, shrimps, gastropods, chaetognaths, cteno-
phores, and medusae. Subsampling was used in an adaptive man-
ner depending on the abundance of specimens in the sample. Large
and less abundant forms (e.g. krill, amphipods, decapod larvae,
chaetognaths, siphonophores) were counted in the whole sample.
Medium-sized organisms (e.g. Calanus copepodites) were counted
in from ½ to 1/32 fraction. Small organisms (e.g. small copepods)
Table 1
List of sampling gears used in the Storfjorden gear intercomparisons study with key features detailed.

Gear Mesh size (lm) Mouth area (m2) Towing velocity Porosity Open area ratio Equivalent volume actually
(1 kn = 31 m/min) filtered MIN to MAX
(AVG ± STD, N) (m3)
1-m2 MOCNESS 180 1 1–2 kn 0.46 5.5 24–335 (144 ± 50; 174)
1-m2 BIONESS 333 1 2–3 kn 28–1725 (190 ± 188; 86)
MultiNet 200 0.25 1–2 kn (oblique haul), 0.4913 5.53 3–41(13 ± 10; 31)
45 m/min = 1.5 kn
(vertical haul)
100 0.25 1–2 kn (oblique haul), 0.4361 5.08 6–31 (15 ± 11; 3)
55 0.25 0.3735 4.24 3–31 (19 ± 12; 3)
Bongo net (Ø = 20 cm) 180 0.0314 23–33 (2)
Bongo net (Ø = 60 cm) 180 0.2826 148–196 (171 ± 23; 4)
WP-2 400 0.25 45 m/min = 1.5 kn 0.4359 5.30 13–14 (2)
200 0.25 0.4913 5.97 2–56 (9 ± 14; 34)
100 0.25 0.4361 5.24 0.7–12
55 0.25 0.3735 4.54 8–12 (2)
1-m Ring Net (CalCOFI) 333 0.785 1–2 kn 0.47 8.58 144–263 (208 ± 60; 3)
Gulf V 300 0.0314 4–5 kn 182–344 (243 ± 64; 5)
LHPR 200 0.10 3–4 kn 0.50 9.0 15

Total length of Net length Specific measures Flow-meter References

gear (m) (m)
1-m2 MOCNESS 6.0 6.0 Modified TSK–digital Wiebe et al. (1976, 1985)
flow-meter
1-m2 BIONESS Inside net/outside net Sameoto et al., 1980
pair of Digital flow-meters
MultiNet 2.50 Lower Ø = 0. 11 m Digital flow-meter HYDRO-BIOS HYDRO-BIOS Apparatebau,
Bongo net (20 cm diameter) – 2.50 Upper Ø = 0.20 m Apparatebau, Kiel, Germany, Kiel, Germany
Lower Ø = 0. 11 m in the center f = 3.33 U/m
Bongo net (60 cm diameter) – 2.50 Upper Ø = 0.60 m
Lower Ø = 0. 11 m
Author's personal copy

WP-2 – 3.11 Net: Upper Ø = 0.57 m; Digital flow-meter Tsurumi-Seiki Kosakusho Fraser (1966)
H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42

lower Ø = 0.11 m Co. Ltd., Yokohama, Japan (TSK),

14.25 cm from the rim
f = 6.4 ± 0.4 revolutions/m at 45 m/min
Cylindrical section: L = 0.95 m;
Conical section: L = 1.66 m
1-m Ring Net 4.0 4.0 Mechanical flow-meter General Oceanics, HYDRO-BIOS Apparatebau, Kiel,
in the center; f = 36.05 revolutions/m Germany
Gulf V 2.40 1.35 Cone: L = 0.5 m; top Ø = 0.2 m; Mechanical flow-meter General Oceanics, Nellen and Hempel (1969)
end Ø = 0.4 m; in the center; f = 36.05 revolutions/m
Net: top Ø = 0.40 m
End Ø = 0.11 m
LHPR 3.57 1.6 Icone = 0.356 m Longhurst et al. (1966) and
Williams et al. (1983)
7
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8 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42

Fig. 4. Flow chart diagram illustrating the treatment of zooplankton samples on the R/V Johan Hjort and the R/V A.v. Humboldt. The only difference was that a Folsom plankton
splitter was used on the Humboldt and a Motoda splitter (illustrated) was used on the Hjort.

were counted in from 1=4 to 1/1024 fraction. The degree of subsam-
pling was adapted so that in most cases more than 100 individuals
of the most common species or groups of medium and small
organisms were counted.
Samples from MULTINET, WP-2, Gulf-V, and 1-m Ring net (Cal-
COFI) were analyzed at the Institute of Baltic Sea Research in
Warnemünde, Germany (IOW), using the same taxonomic catego-
ries as for the MOCNESS and BIONESS samples analyzed by IMR.
Subsampling was used and all individuals were counted in a frac-
tion from 1/5 to 1/200. In a few cases the whole sample was ana-
lyzed. The degree of subsampling was adjusted to the density of
organisms in the sample so that usually between 300 and 700 indi-
viduals were counted in each case.
The LHPR samples were analyzed at Plymouth Marine Labora-
tory as described by Halliday et al. (2001). Zooplankton species
were identified to as detailed a taxonomic level as feasible and
counted. Mechanical damage by the sampler generally prevented
distinguishing between Paracalanus spp., Pseudocalanus spp., and
Microcalanus spp. and they were lumped into a single category
(Microcalanus spp.) since the majority were Microcalanus spp.
Copepodite stages were denoted from CI to the adult stage,
CVI. The depth information and flow data were used to
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
standardize the zooplankton counts to #/m3 for each 10 m inter-
val sampled.
Further details on taxonomic categories identified, subsam-
pling, and numbers counted are given in Wiebe et al. (2002) along
with the data.
2.3.3. Measurements of zooplankton size
Morphometric measurements were made on a number of zoo-
plankton taxa in samples obtained from the upper 100 m with MOC-
NESS. The measurements included length and width and/or height,
as detailed in Fig. 5 for the various taxa. Length measurements to
the nearest mm of individual specimens of krill, shrimps, and fish
were carried out for part of the material that was sorted for dry mass
determination. An extensive set of morphometric measurements of
zooplankton collected with LHPR was reported by Halliday (2001).
2.4. Acoustic sampling
Acoustic data were collected with a hull-mounted EK500 Sim-
rad echo sounder on R/V Johan Hjort, operating with 4 frequencies,
Fig. 5. Size measurements of zooplankton taxa. For the majority of taxa, D1 is length and D2 is width of the organisms.
9
18, 38, 120, and 200 kHz. These provided an indication of the ver-
tical distribution of the plankton and nekton, and were used to
guide the exploratory net sampling. The acoustical data are used
in this paper mainly for descriptive purposes and we attempt no
quantitative comparison between the acoustics and the net sam-
ples herein. Much of the acoustical data are contained in Wiebe
et al. (2002; attached CD-ROMs).
2.5. Determination of environmental variables
Meteorological, surface water property, and CTD/nutrients, phy-
toplankton species/fluorescence/downwelling light data were col-
lected on the R/V A. v. Humboldt and the R/V Johan Hjort using
standard methodologies. A brief summary of these data is pre-
sented below.
2.6. Data treatment and analyses
In the data analysis of the gear intercomparisons we have
distinguished between day and night periods. For the primary
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10 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42

comparisons we have used data collected with different gears on

the same date. We have also pooled data from neighboring dates
and over the whole sampling period. This introduces some uncer-
tainty due to possible temporal changes in the plankton commu-
nity over this period. Two-way Analysis of Variance (ANOVA)
using MATLAB software was carried out in comparisons of the ef-
fects of net mesh and gears. Similarity and hierarchical cluster
analyses were performed on some data sets using PRIMER software
after applying a log(x + 1) transformation and using Bray-Curtis
similarity resemblances (Clarke and Warwick, 1994).

3. The fjord environment

3.1. Environmental conditions and phytoplankton

The Storfjord is a glacially excavated flooded valley with steep

sides and a relatively flat bottom. The depth along the 5 nm sam-
pling transect varied from about 440 m in the western end to about
400 m in the eastern end. The transect was located outside the
opening to the Hjørundfjord, which is a side-branch to the Storfj-
ord. The Storfjord continues northwestwards through the Breisund
onto the shelf as a distinct trench with an outer sill depth of about
131 m (Dyb et al., 2003). There is also a connection to the sea
southwestwards through the Vartdalsfjord, but this is longer with
some narrow passages and a shallower sill at the shelf. The Storfj-
ord winds its way inland for another ca. 60 km from the main
study site to the head of the fjord in Geiranger. The maximum
depth of the fjord is about 680 m in the area of our hydrographic
sampling station 9017 (Fig. 1).
The weather conditions remained fairly calm and sunny to
partly cloudy during the study period. The wind speed was about
10 m s1 or less. At this latitude and time of the year, the days
are long and nights are short and not completely dark. Based on
the incoming solar radiation, the day period was defined as from
0530 to 2030 (local time), corresponding to incoming radiation of
about 30 lE m2 s1 or more. The night period was defined as
the time between 2230 and 0330, corresponding to incoming radi-
ation of about 2 lE m2 s1 or less. Dawn and dusk were defined as
the time periods between 0330 and 0530 and between 2030 and
2230, respectively.
Overall there were only slight temporal changes in the hydrog-
raphy (Fig. 6). Basically the water mass structure and properties in
terms of salinity and temperature remained unchanged during the
study period at the main study site. The salinity was low in the sur-
face layer, but increased rapidly to more than 34.5 below 30 m.
There was no clear pattern in the near surface variability among
the profiles taken during the study period. Below the surface layer,
there were only small differences between the profiles taken dur-
ing the course of the study. Temperature was around 11 °C in the
surface layer, decreasing to fairly uniform temperatures between
7 and 8 °C deeper in the water column. Apart from a slight warm-
ing of about 1 C of the very surface layer, temperature remained
virtually unchanged during the study period. The oxygen concen-
tration increased from the surface to maximum of ca. 8 ml l1 at
about 10–15 m. The oxygen concentration decreased sharply be-
low the maximum, but remained relatively high between 5.5 and
6.5 ml l1 throughout the deeper part of the water column. The
extinction coefficients were highest in the layer from about 10–
30 m, corresponding to the layer with the broad maximum in
in situ fluorescence. Over the course of the sampling, there was a
trend in Secchi depth readings to become shallower, decreasing
from 9 m on June 4 to 6 m on June 12. This probably reflected in-
creased particulate matter in the water column, as suggested by
an increase in chlorophyll fluorescence. The phytoplankton com-
munity in the upper 10 m and at 20 m was dominated by small
Fig. 6. Temporal development (as isopleth diagrams) in (A) temperature (oC), (B)
salinity, (C) oxygen concentration (ml l1), (D) nitrate concentration (lM l1), and
(E) in situ chlorophyll fluorescence (relative units) in the upper 50 m at the main
study site during the study period from 5 to 12 June.
unidentified flagellates and diatoms. The dominant diatoms were
Chaetoceros spp., Nitzschia pungens, Skeletonema costatum, Leptocyl-
indrus danicus, and unidentified centric diatoms.
3.2. Zooplankton community composition, distribution, and size
3.2.1. Taxonomic composition
The zooplankton community in the study area was dominated
by neritic species. Based on sampling with MOCNESS (180 lm),
the rank order of numerical abundance over the whole water col-
umn for taxonomic categories revealed that cladocerans (mainly
Evadne nordmanni but also Podon sp.) were the most common
group, making up almost 1/3 of the total number of individuals
(Table 2). The cladocerans were most abundant in the upper
50 m where they occurred with a mean density of about 1200 indi-
viduals m3. A higher value (about 5000 m3) was obtained with
WP-2 net in the upper 18 m (Fig. 7). At the average density, a ver-
tical haul with WP-2 net in the upper 20 m (0.25 m2 opening)
would collect 6000 individual cladocerans. A MOCNESS net hauled
through the same layer with a typical sampling volume of 300 m3
would collect 360,000 individuals.
The second most abundant taxa was Microcalanus sp., which
was abundant in the upper layer, but more numerically dominant
in the deeper layers (43–48% below 50 m). The third most common
group was Oithona spp., which occurred most commonly in the
upper layer. Together these three groups or species made up about
2/3 of the number of individuals in the water column.
Various larval forms (cirripedia, bivalves, cyphonautes, gastro-
pods, echinoderms) each made up from 1% to 5% and in sum about
12% of the numerical abundance over the water column. These
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42 11

Table 2
Rank order of abundance of zooplankton taxa for two MOCNESS tows with 180 lm mesh (Station Johan Hjort 394 MOCNESS hauls 4 and 5) based on their combined data in the 0–
400 m depth interval, and the abundance and percentage of total individuals for depth intervals within the upper 400 m water column.

Species/categories 0–400 m 0–50 m 50–100 m 100–200 m 200–400 m

3 3 3 3
#/m % of Total #/m % of Total #/m % of Total #/m % of Total #/m3 % of Total
Cladocera 162.01 32.396 1206.880 38.908 49.778 19.555 10.978 11.374 4.36 4.537
Microcalanus sp. 83.17 19.739 237.198 10.911 171.653 44.450 47.152 48.741 40.56 42.569
Oithona spp. 59.47 13.371 430.391 16.673 31.311 8.733 4.152 4.320 1.43 1.783
Cirripedia larvae 22.68 4.674 155.186 5.262 10.793 4.613 3.395 3.488 2.170 2.105
Bivalve larvae 15.87 3.558 120.102 4.681 1.523 0.694 1.473 1.507 0.60 0.743
Calanus spp. CI-III 15.11 3.239 115.687 4.191 3.088 0.954 0.664 0.681 0.19 0.211
Temora sp. 15.95 3.088 119.077 3.621 4.137 1.116 1.490 1.570 0.34 0.340
Metridia CIV-VI 11.50 2.745 7.477 0.359 5.243 1.528 10.326 10.866 14.66 18.957
Calanus spp. CIV-VI 11.66 2.555 12.484 0.493 2.887 0.754 6.014 6.318 16.47 18.779
Copepodnauplii 12.53 2.529 96.083 3.126 2.621 0.764 0.463 0.463 0.16 0.227
Acartia spp. 7.55 1.519 57.365 1.892 1.106 0.412 0.625 0.664 0.17 0.156
Cyphonautes 6.94 1.282 52.467 1.482 0.835 0.565 0.637 0.659 0.23 0.269
Gastropod larvae 5.67 1.238 40.456 1.479 3.650 0.787 0.434 0.459 0.09 0.108
Oncea spp. 5.10 1.219 25.047 1.067 5.243 1.528 1.315 1.341 1.98 2.406
Pseudocalanus CIV-VI 4.95 1.071 32.791 1.314 7.440 1.667 0.882 0.908 0.24 0.261
Pseudocalanus CI-III 4.42 1.021 29.473 0.987 2.065 0.387 0.045 0.045 0.11 0.154
Echinoderm larvae 3.60 0.760 18.569 0.576 7.323 2.600 1.230 1.242 0.11 0.119
Metridia spp CI-III 3.00 0.760 11.793 0.554 9.991 2.384 0.023 0.023 0.54 0.590
Maurolicus eggs 3.02 0.626 21.729 0.740 0.783 0.530 0.744 0.760 0.05 0.068
Ostracoda 1.86 0.455 3.160 0.164 2.636 1.448 0.587 0.590 1.98 2.537
Appendicularia 2.17 0.389 15.408 0.412 0.000 0.000 0.090 0.090 0.44 0.431
Decapod larvae 1.35 0.312 9.512 0.381 0.982 0.244 0.038 0.039 0.05 0.076
Other fish eggs 1.05 0.274 0.000 0.000 7.109 1.444 0.335 0.350 0.16 0.177
Other copepods 0.99 0.254 3.164 0.164 2.753 0.516 0.363 0.373 0.32 0.329
Euphausiid larvae 0.93 0.223 7.092 0.310 0.201 0.089 0.058 0.058 0.000 0.000
Scolecithricella minor 0.86 0.188 0.000 0.000 0.765 0.518 1.829 1.879 0.61 0.650
Polychaete larvae 0.59 0.161 4.741 0.246 0.000 0.000 0.000 0.000 0.00 0.001
Fish larvae 0.47 0.111 0.012 0.001 3.595 1.424 0.090 0.090 0.00 0.001
Siphonophores 0.27 0.057 0.006 0.000 0.005 0.001 0.093 0.093 0.48 0.530
Paraeuchaeta nor. CIV-VI 0.20 0.049 0.000 0.000 0.000 0.000 0.274 0.275 0.26 0.318
Others 0.21 0.044 0.024 0.001 0.149 0.098 0.142 0.143 0.31 0.318
Hydrozoa 0.16 0.043 0.067 0.002 0.767 0.148 0.082 0.082 0.07 0.097
Chaetognaths 0.11 0.029 0.009 0.000 0.000 0.000 0.351 0.352 0.04 0.056
Paraeuchaeta norv. CI-III 0.07 0.015 0.000 0.000 0.000 0.000 0.148 0.151 0.06 0.060
Centropages 0.02 0.004 0.000 0.000 0.070 0.047 0.000 0.000 0.03 0.029
Aglantha 0.01 0.003 0.076 0.003 0.000 0.000 0.003 0.003 0.00 0.004
Euphausiids <0.01 0.000 0.000 0.000 0.000 0.000 0.007 0.007 0.00 0.001
Total 465.50 2833.500 340.50 96.50 89.30

larval forms were most common in the upper layer, also in terms of
relative abundance.
Large calanoid copepods, in particular Calanus finmarchicus, are
usually dominant in zooplankton in the Norwegian Sea and adja-
cent shelf and fjord areas (Halvorsen et al., 2003; Melle et al.,
2004; Heath et al., 2004). We had therefore an expectation when
planning our study that we would be working in a Calanus domi-
nated zooplankton community. The copepods Calanus sp. and Met-
ridia sp., however, were present in relatively low densities. They
made up respectively about 6 and 3% of the numerical abundance,
with about 12 individuals m3 as an average for the whole water
column and around 15 individuals m3 in the deeper part (200–
400 m) for C. finmarchicus and Metridia spp. separately (Table 2).
The older copepodite stages (CIV-VI) were most abundant in the
deeper layer (200–400 m) where they together made up about
30% of the numerical abundance. Copepods (other than Microcal-
anus and Oithona spp.) made up about 18% of the numerical abun-
dance over the water column. At these densities a WP-2 net towed
vertically in the upper 100 m would catch about 300 individuals of
Calanus or Metridia, and a MOCNESS would typically catch about
3600 individuals of each of these species with a tow length corre-
sponding to 300 m3 of water filtered. The small copepods Oithona
spp. occurred with a density of about 400 individuals m3 in the
upper 50 m, while Microcalanus sp. were found with about 200
individuals m3 in the upper 100 m, as ‘‘seen’’ by the MOCNESS
(Table 2).
Krill was another group of zooplankton that we expected to be
relatively common in the Storfjord, and we carried out sampling
with MOCNESS and BIONESS to compare their effectiveness in cap-
turing krill and other macrozooplankton. Unfortunately, as with
Calanus, the density of krill was generally low in the study area.
In two MOCNESS hauls, the average density of krill samples was
less than 1 individual per 100 m3 (Table 2). It means that with a
typical haul (about 300 m3 volume filtered), the MOCNESS would
capture 3 or fewer krill. The highest density of krill was sampled
with night-time hauls with BIONESS in the upper 100 m (about
1 mg dry mass m3) which would correspond to approximately 1
krill per 10 m3 for Thysanoessa inermis 15–20 mm long (Dalpadado
and Skjoldal, 1996), but fewer for the larger Meganyctiphanes nor-
vegica, which was also present. In spite of these complications, the
results of this study should have broad applicability.
Larger forms were generally not common. Fish larvae, siphono-
phores, chaetognaths, and the large copepod Paraeuchaeta norvegica
were regularly caught, but they occurred at low numerical abun-
dance (<0.5 m3). Fish larvae were caught at 50–100 m whereas
the other groups mainly were collected below 100 m (Table 2).
The rank order of numerical abundance of taxa collected from
the upper 18 m with WP-2 net (200 lm) was in general agreement
with the results described in Table 2 (see Fig. 7). Microcalanus sp.
was not common in the upper 18 m (and does not appear in the
plot), while some small forms (notably Limacina sp., Fritillaria sp.
and polychaete larvae) were more common in this data set.
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12 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
Fig. 7. Rank order of abundance (A) and replicate variability (B; coefficient of variation = standard deviation/mean  100%) for species counts for 20 replicate samples from
the 10 hauls with double WP-2 net (200 lm) (A.v. Humboldt station 9012).
3.2.2. Vertical distribution of numerical abundance, biomass, and
acoustic backscatter
Vertical plots of total numerical abundance and total dry mass
were developed from the pooled data collected with all zooplank-
ton sampling gears during the study (Fig. 8A and B). The total
abundance ranged from <10 to about 20,000 individuals m3. The
highest abundance values occurred in the upper layer (above
30 m), with lower abundances (about 500 m3 or less) deeper than
100 m. The total dry mass ranged from 0.2 to 53 mg m3. The dry
mass followed a similar pattern to that of abundance, but with less
marked difference between the upper and the deeper layers. This is
reflected in higher ratios of total dry mass to total abundance
(‘average individual dry mass’) in the deeper layer (Fig. 8C).
Day and night biomass profiles obtained with MOCNESS
(180 lm) on 11–12 June show that combined about 50% of the total
biomass in the water column to 375 m was distributed in the upper
50 m (Fig. 9). The biomass was low in the 100–200 m depth layer
with 5–7% of the total biomass. The biomass was higher again be-
low 200 m with about 35% of the total biomass found in the 200–
375 m depth interval. The smallest size fraction (<1 mm) domi-
nated in the upper 150 m, whereas the intermediate fraction (1–
2 mm) dominated below 200 m. The total biomass in the water col-
umn was about 4.0 g dry mass m2, with little difference between
day and night. Of this total the smallest size fraction (<1 mm) made
up 38%, while the medium and largest fractions each made up 30%.
There were subtle changes between the day and night profiles,
indicating that there was some diel vertical migration. The biomass
in the 50–100 m layer was higher during night, indicating some
migration into this layer from below. One of three replicates of
the largest size fraction (>2 mm) in the day-time series at the
upper and deepest layers was markedly higher than the other
two and could have been outliers. Assuming this to be the case,
there is a pattern that the medium and largest size fractions (1–
2 mm and >2 mm) were higher in the upper layer during night.
More detailed profiles of zooplankton biomass in the upper
100 m obtained with MOCNESS (180 lm) on 6 June confirm the
pattern of some limited vertical migration (Fig. 10 A and B). The
smallest size fraction (<1 mm) shows very similar values between
day and night, being distributed in the upper 25 m with no indica-
tion of vertical migration. The medium and largest size fractions
(1–2 mm and >2 mm), however, showed higher values at night,
suggesting migration from below.
Acoustical recordings at different frequencies showed some
persistent vertical patterns. At all frequencies, there was an upper
scattering layer (SL) with peak concentrations at about 20–25 m,
i.e. centered at the base of the pycnocline, but with recordings
down to 40–60 m (Fig. 11). No indications of diel variations were
recorded for this layer (Fig. 11). Though clearly depicted, Sv values
were low and the SL originated from relatively weak or sparsely
distributed targets. Targets were probably large due to the reflec-
tion even at the lowest frequency.
Below 60 m was a zone almost devoid of backscattering. A dielly
migrating SL was located near 150 m during daytime, although
with erratic small-scale fluctuations in the vertical distribution.
Its nocturnal ascent terminated at ca 20 m (Fig. 11). Catches from
trawling targeted at this SL during different phases of the diel
migration cycle were always dominated by the small mesopelagic
fish Müller’s pearlside Maurolicus muelleri.
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
Fig. 8. Vertical plots of (A) total abundance, (B) total dry mass, and (C) ratio of total
dry mass to total abundance (‘average individual dry mass’) based on all
zooplankton sampling gears used along the 5 nm sampling transect during the
study where species counts were also made.
3.2.3. Size measurements of individual zooplankton
Based on the size measurements of length and width of the var-
ious zooplankton groups, species and stages (Table 3), the zoo-
plankton community was dominated by small forms. The largest
specimens measured were copepodites of Paraeuchaeta of 3 mm
length. The width of the copepods was typically about 1/3 of their
length. This varied somewhat with Calanus being the slimmest with
a width/length ratio of about 0.30, and Temora the most bulky with
a ratio of 0.47. The shape of groups other than copepods varied,
with high width/length ratios found for bivalve larvae, cyphona-
utes, cirriped nauplii, and the appendicularian Frittilaria borealis.
4. Results from gear comparisons
4.1. Replicate variability
Although the operating time required for the sampling put a
constraint on the number of replicates that could be accommo-
13
dated in each intercomparison exercise, a substantial number of
replicate tows and intercomparions between gears were com-
pleted. Replicate variance reflects the variances in zooplankton dis-
tribution and abundance, gear performance, and handling and
analyses of the zooplankton samples.
A replicate variability experiment was carried out from R/V A.v.
Humboldt on 9 June, using vertical hauls from 18 m to the surface
with a double WP-2 net. Ten replicate hauls were made during a
1.5 h period in the late afternoon. The coefficient of variation
(CV) (standard deviation (SD)/mean  100%) was about 17% for
both the total biomass and the smallest size fraction (<1 mm),
which made up most of the total (Table 4). The CV was larger
(70–120%) for the 1–2 mm fraction, which made up only about
3% of the total biomass. The left and right series of nets gave very
similar results both in terms of biomass and CV. The difference be-
tween the two nets in the pair was close to zero and the variability
(as SD) was similar to that of the series of nets. These results indi-
cate that the procedures followed for sampling, sample handling,
and analyses of dry mass produced consistent results with fairly
low variability.
The variability in the species counts from the taxonomic analy-
sis increased with decreasing numerical abundance of the taxa
(Figs. 7 and 12). Subsampling was used prior to counting. Follow-
ing the initial split of each sample into two halves (for biomass
and species counts, Fig. 4), a 1/50-portion of the preserved sample
was used for counting. The volume of water filtered was about
5 m3. Thus a species count of 5 individuals in the subsample corre-
sponds to an abundance of 100 individuals m3. A count of 50 cor-
responds to 1000 individuals m3 and a count of one in every
second sample on average, corresponds to 10 individuals m3.
The low numbers counted with the subsampling procedure no
doubt contributed much to the high variability associated with
low abundance.
The CV for the species counts for the 4 most abundant taxa
(800–7000 individuals m3) was 25–35% (Fig. 7). For the two cope-
pods, Temora spp. and Acartia spp. (CIV-VI), with abundance of
about 200 m3, the CV was about 75%.
The variability in samples of dry mass obtained with replicate
hauls with MOCNESS (180 lm) in the upper 100 m during the
day and night periods on 6–7 June (see Fig. 2) generally increased
from the smallest (<1 mm) to the largest (>2 mm) size fractions
(Fig. 10C and D). The average CV over the depth intervals in the
day and night profiles was 55%, 68%, and 118% for the <1, 1–2,
and >2 mm size fractions, respectively. The corresponding ranges
in the CV values were 7–121, 18–114, and 69–210% for the three
size fractions. The average CV% for the total biomass was 61%, with
a range of 6–121%. There was no clear relation between the CV and
the biomass level in the depth layers, although the CV tended to be
lower in the upper layer where most of the biomass was found,
during night-time. This difference in CV values between day and
night for the upper layers (above 37 m) could reflect a more patchy
distribution during daytime at the scale resolved by the sample
size.
Replicate hauls, each with replicate nets, were taken with MOC-
NESS (333 lm) and BIONESS (333 lm) in the depth intervals 200–
250 m and 250–300 m during day and night-time on 10 June (see
Fig. 2). In this depth layer, the variability was similar between
the smallest and medium size fractions (<1 mm and 1–2 mm), with
average CV values of 50–55% (Table 5B). For the largest size frac-
tion (>2 mm (not including fish, shrimp and krill)), the average
CV was about 70%. For the total biomass the average CV was about
55%. The CV values tended to be somewhat (but not significantly)
lower for the BIONESS than for the MOCNESS samples.
The variability in depth integrated biomass (per m2) as CV val-
ues showed no clear differences among the gears (Tables 5 and 7
and Fig. 13). The mean CV values over all gears and depth intervals
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14 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
Fig. 9. Vertical profiles of size-fractioned dry mass over the water column (0–375 m) obtained with MOCNESS (180 lm). (A) Day profile as mean values for 3 replicate hauls
(Johan Hjort station 394, MOC 3, 4 and 5) and (B) night profile (Johan Hjort station 395, MOC 1).
were 39% for the <1 mm size fraction, 34% for the 1–2 mm, 73% for
the >2 mm, and 32% for the sum total. The 60-cm Bongo net gave
very low CV values (<15%) for all size fractions. Since the number
of replicates was low (n = 4), this could be by chance. The CV values
for the largest size fraction (>2 mm) were greatest for the WP-2 net
and the MultiNet. This could reflect the smaller net openings and
sample size with these nets. The two pair-wise comparisons be-
tween MOCNESS (180 lm) and BIONESS (333 lm) showed lower
variability for the medium and largest size fractions (1–2 mm
and >2 mm) with BIONESS, but higher variability for the smallest
size fraction (Fig. 13B).
The variability in the determination of biomass of fish, shrimp,
and krill was generally large, with most values between 50% and
150% and average CV values of 71%, 116%, and 66% for the three
groups (Fig. 13B). The high variability reflected the low numbers
of these organisms in the samples, often with no individuals caught
in some of the replicates. The BIONESS had consistently lower CV
values for these groups than did the MOCNESS.
The CV values for all the intercomparisons between gears
showed no strong relationship with the zooplankton biomass level,
although CVs trended smaller with increasing biomass particularly
for the 1–2 mm size fraction (Fig. 14 A). The CV values for the larg-
est size fraction (>2 mm) were considerably higher than for the
smaller size fractions (<1 mm and 1–2 mm) for comparable bio-
mass levels. The CV values tended to be highest for the smallest
sample size and smaller with increasing volume filtered, most
notable for the largest size fraction (>2 mm; Fig. 14B).
The higher variability at the smaller sample size could reflect
more patchiness at the scale of sampling. With larger sample sizes,
due to larger net opening and especially the longer tow distance,
small-scale patches are to larger extent averaged out (Wiebe,
1971, 1972). The higher variability for the largest size fraction
(>2 mm) indicates more patchiness associated with the greater
mobility of these organisms compared to the smaller fractions.
The generally low CV values for the small and medium size frac-
tions (<1 and 1–2 mm) in the upper part of the water column (Ta-
ble 4, Fig. 13A) suggest that the inherent variability in the sample
processing (see Fig. 4) and weighing was moderately low.
4.2. Effects of mesh size
The gears that were intercompared had mesh sizes ranging
from 180 to 333 lm (Table 1). Some direct comparisons of nets
with different mesh size were carried out with the same gear, i.e.
with the WP-2, MultiNet, and MOCNESS.
WP-2 nets with 55, 100, 200, and 400 lm mesh were operated
from R/V A. v. Humboldt on 10 June, with two replicate vertical
hauls with each net from 100 m to the surface. The total biomass
caught was about half for the coarsest net (400 lm) compared to
the 200 lm net. The 400 lm net caught more of the largest size
fractions (1–2 mm and >2 mm), but considerably less of the 0.5–
1 mm fraction (Fig. 15). The total biomass was similar or higher
for the 100 lm net, but lower for the 55 lm net, compared to
the 200 lm net. There was no clear difference among these 3 nets
for the largest size fractions (1–2 mm and >2 mm). The 55 lm net
collected considerably less of the 0.5–1 mm fraction, especially the
first tow, perhaps because of net clogging, giving a lower total de-
spite the collection of a fair amount of biomass in the fractions
<0.2 mm (Fig. 15).
The species counts also revealed substantial differences be-
tween the nets of different mesh size (Fig. 16). The 400 and
200 lm mesh nets collected none or few copepod nauplii and
small copepods grouped as others. The 400 lm net collected none
or relatively few of the cladoceran Evadne nordmanni, the copepods
Oithona, Microcalanus, Temora, Acartia, and Oncaea, as well as bi-
valve larvae. A conspicuous feature for many taxa was a marked
decrease in catch from the 200 lm net compared to the finer
meshed nets. This was the case for the cladocerans Evadne and
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42 15

Fig. 10. Vertical profiles of size-fractioned dry mass in the upper 100 m obtained with MOCNESS (180 lm). (A) Day profile as mean values for 6 hauls taken during daytime on
6 June (Johan Hjort station 391). (B) Night profile as mean values for 4 profiles taken during night between 6 and 7 June (Johan Hjort station 392). (C and D) Coefficient of
variation (standard deviation/mean  100%) for the replicate hauls during day and night, respectively.

Podon, the copepods Oithona and Calanus spp. CIV-CVI, cyphona-
utes, and cirripedia larvae.
The MOCNESS was used with 180 lm and 333 lm mesh nets in
hauls between 200 and 300 m depth on 10 and 11 June. The
180 lm nets collected about twice as much total biomass as the
333 lm nets (Fig. 17). The 180 lm nets collected more in the
smallest size fraction (<1 mm), but the largest difference was for
the medium size fraction (1–2 mm) where the 180 lm collected
more than twice as much as the 333 lm nets. The 333 lm nets col-
lected more fish and shrimps.
The species counts also revealed clear differences between the
180 and 333 lm MOCNESS nets (Table 6). The 333 lm nets col-
lected very few of the small copepods Microcalanus sp., Oithona,
and Oncaea (Fig. 18). For the larger copepodite stages (CIV-CVI)
of Calanus and Metridia, the difference was not large, although
the 180 lm nets tended to collect more than the 333 lm nets.
The numbers collected were highest for the 333 lm nets for chae-
tognaths, Maurolicus eggs, and young copepodite stages of Calanus.
This is somewhat unexpected. For chaetognaths the difference ap-
pears to be real. They were counted for the whole preserved sam-
ple and the counts ranged from 13 to 188 for the 333 lm nets. For
Maurolicus eggs and young copepodites of Calanus, the result could
be an artifact due to low abundance and subsampling prior to
counting. The effect of mesh size on the catch of the smaller species
is clear in a plot of the ratio of the 333 lm mesh to 180 lm mesh
for copepods and other zooplankters vs. their mean individual
width (Fig. 19). Several high points have been corrected for outli-
ers. The 333 lm mesh has a diagonal of 470 lm. For individuals
with a width below this value, the ratio is generally less than
one and decreases rapidly with decreasing animal width. Individu-
als with a width larger than the diagonal are retained about equally
by both mesh sizes.
4.3. Comparison of gears in the upper 100 m – 5–6 June
During 5 and 6 June, sampling was carried out in the upper
100 m with WP-2 net (200 lm), 20- and 60-cm Bongo nets
(180 lm), MultiNet (200 lm), MOCNESS (180 lm), BIONESS
(333 lm), 1-m Ring Net (333 lm), and Gulf-V (300 lm). Depth-
integrated values of dry mass per m2 are summarized as mean val-
ues with variability (coefficient of variation) in Table 7 and shown
in Fig. 20. The depth-integrated values are based on one haul over
the 0–100 m depth interval (Bongo, 1-m Ring net, and Gulf-V), or
depth-stratified sampling over two (WP-2), 5 (MultiNet), or 8
depth layers (MOCNESS and BIONESS).
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16 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
Fig. 11. Incoming irradiance (A) and acoustical recordings at (B) 200, (C) 120, and (D) 38 kHz during a 24-h cycle on 6–7 June.
The finer meshed nets collected considerably more biomass in
the smallest size fraction (<1 mm) and in total than did the coarser
meshed nets (Fig. 20). There was a fair agreement in the biomass
levels obtained with the different finer meshed gears. The highest
biomass was obtained with the MultiNet (200 lm), followed by
WP-2 (200 lm). These two nets collected most biomass in the
<1 mm fraction. The high total catch with the MultiNet was partly
due to a high value in the >2 mm size fraction, where one of the
three replicates was exceptionally high. Removing this value as
an outlier reduces the mean for the >2 mm fraction to 101 mg
dry mass m2, which is compararable to the other nets, and the to-
tal to 1182 mg dry mass m2 (Fig. 20).
Of the two Bongos, the 60-cm net collected somewhat more
than the 20-cm net, which showed higher variability in the results
(Table 7).
There are three columns for MOCNESS (180 lm) in Fig. 20. The
first is the mean for the 6 replicate hauls made during day time on
6 June. The first haul gave exceptionally low biomass values for
several of the nets. It was noted a malfunctioning of the flowmeter
during this haul, and the volumes were estimated indirectly.
Removing this haul due to the uncertainty raises the mean biomass
values to 715 and 873 mg dry mass biomass m2 for the <1 mm
fraction and the total, respectively (Fig. 20). The second column
for MOCNESS is the mean for the 4 hauls made during night time
between 6 and 7 June. This is somewhat higher than for the day
samples, reflecting the migration of some zooplankton into the
upper 100 m from below (Fig. 10). The third column is one haul
made during 5 June.
Mean values for the depth-stratified samplers were also calcu-
lated for the 0–25 m and 25–100 m depth intervals (Fig. 21). The
values used for the MultiNet are with removal of the outlier for
the >2 mm size fraction (in the 0–25 m layer). The values were
quite similar for all three gears in the upper 25 m. For the 25–
100 m depth interval, MOCNESS collected less than half of what
WP-2 and MutiNet did for the <1 mm size fraction. This difference
was also reflected in the total (Fig. 21B). The day and night series
for MOCNESS gave very similar values, except for somewhat higher
night values for the 1–2 and >2 mm size fractions and the total for
the 25–100 m depth layer.
The BIONESS (333 lm), 1-m Ring net (333 lm), and Gulf-V
(300 lm) samplers produced considerably less biomass in the
smaller size fraction (<1 mm), which is reflected also in low totals
(Fig. 20). The 1-m Ring net in particular gave very low biomass in
the small fraction. BIONESS gave the highest value for the medium
size fraction (1–2 mm), while 1-m Ring net and Gulf-V gave the
lowest values. For the largest size fraction (>2 mm), there were
no clear differences, although WP-2 and 20-cm Bongo gave the
lowest values.
A two-way analysis of variance was used to test the significance
of the differences between size categories and sampling devices.
Systems that had three or more replicate samples were used in
the analysis (there were only two tows with the BIONESS, so it
was not included). For the MultiNet the outlier replacement de-
scribed above was used so that each net system had the same num-
ber of replicates. The differences between size fractions shown in
Fig. 20 are significant (p < 0.05) as are the differences between sam-
plers. As noted above, the main differences between the samplers
are that the 1-m Ring net and the Gulf-V samplers caught substan-
tially less plankton biomass, while the BIONESS and Bongo 20 cm
did so to a lesser extent. The WP-2, MultiNet, MOCNESS-180 lm,
and Bongo 60 cm biomasses were not significantly different.
The taxonomic composition of samples (depth-integrated num-
ber of individuals m2 for the upper 100 m) showed clear
differences among the gears (Fig. 22, Table 8). The coarser meshed
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42 17

Table 3
Individual size measurements in mm for zooplankton taxa (species, copepodite stages, and groups). D1 = length, D2 = widths; see Fig. 5.

Species (and stage) n D1 D2

Mean SD Min Max Mean SD Min Max
Evadne 20 0.67 0.12 0.46 0.87 0.26 0.05 0.17 0.38
Podon 9 0.60 0.21 0.40 0.94 0.29 0.11 0.18 0.47
Bivalve larvae 77 0.31 0.04 0.24 0.40 0.28 0.04 0.18 0.35
Cyphonautes 41 0.65 0.04 0.54 0.72 0.51 0.04 0.44 0.59
Cirriped nauplii 58 0.47 0.04 0.38 0.52 0.40 0.04 0.32 0.46
Cirriped cypris 15 0.63 0.03 0.56 0.68 0.28 0.03 0.26 0.35
Conchoecia 37 0.49 0.13 0.28 0.68 0.30 0.07 0.17 0.50
Frittilaria borealis 20 0.74 0.14 0.52 1.00 1.07 0.27 0.71 1.60
Echinoderm larvae 11 0.55 0.16 0.31 0.84 0.34 0.11 0.21 0.48
Gastropod larvae 28 0.35 0.11 0.24 0.64 0.23 0.07 0.14 0.40
Polychaet larvae 20 0.80 0.10 0.52 0.96 0.18 0.04 0.12 0.28
Copepod nauplii 18 0.42 0.10 0.22 0.56 0.20 0.03 0.14 0.26
Krill nauplii 2 0.47 0.45 0.49 0.26 0.26 0.26
Krill metanauplii 2 1.05 0.99 1.10 0.43 0.42 0.44
Decapod larvae 3 1.77 0.28 1.46 2.00 0.90 0.07 0.82 0.94
Pearlside eggs (Maurolicus muelleri) 29 1.43 0.07 1.31 1.55 0.98 0.05 0.90 1.08
Calanus finmarchicus – CI 16 0.66 0.04 0.61 0.78 0.22 0.02 0.19 0.24
Calanus finmarchicus – CII 7 0.96 0.06 0.85 1.03 0.27 0.04 0.19 0.31
Calanus finmarchicus – CIII 5 1.35 0.04 1.32 1.40 0.37 0.02 0.35 0.38
Calanus finmarchicus – CIV 16 1.85 0.13 1.55 2.03 0.56 0.06 0.45 0.66
Calanus finmarchicus – CV 23 2.45 0.12 2.23 2.63 0.72 0.03 0.65 0.78
Calanus finmarchicus – CVI females 13 2.56 0.15 2.33 2.81 0.80 0.05 0.72 0.90
Calanus finmarchicus – CVI males 70 2.40 0.28 1.73 2.88 0.72 0.08 0.48 0.90
Pseudocalanus spp. CI-III 4 0.54 0.04 0.49 0.59 0.19 0.00 0.19 0.19
Pseudocalanus spp. CIV-V 1 0.87 0.35
Pseudocalanus spp. CVI females 2 0.86 0.33
Pseudocalanus spp. CVI males 1 0.82 0.28
Microcalanus CIV-V 14 0.43 0.03 0.40 0.48 0.18 0.02 0.16 0.21
Microcalanus CVI females 17 0.47 0.03 0.40 0.52 0.21 0.01 0.19 0.23
Acartia spp. CIV-V 13 0.77 0.09 0.64 0.87 0.24 0.03 0.19 0.28
Acartia spp. CVI females 2 0.98 0.94 1.02 0.33 0.31 0.34
Acartia spp. CVI males 1 0.82 0.28
Temora spp. CI-III 7 0.38 0.06 0.30 0.49 0.20 0.03 0.16 0.26
Temora spp. CIV-V 10 0.60 0.07 0.48 0.68 0.28 0.03 0.24 0.35
Temora spp. CVI females 3 0.65 0.07 0.60 0.73 0.35 0.04 0.32 0.40
Temora spp. CVI males 2 0.66 0.64 0.68 0.27 0.26 0.27
Metridia spp. CI-III 3 1.09 0.18 0.88 1.19 0.40 0.08 0.31 0.44
Metridia lucens CIV 5 1.21 0.03 1.19 1.25 0.49 0.03 0.44 0.50
Metridia lucens CV 37 1.56 0.07 1.38 1.63 0.59 0.04 0.50 0.69
Metridia lucens CVI females 29 1.70 0.13 1.43 1.91 0.65 0.05 0.48 0.75
Metridia lucens CVI males 55 1.21 0.05 1.06 1.38 0.49 0.02 0.44 0.56
Metridia longa CIV 16 1.31 0.05 1.25 1.38 0.49 0.02 0.44 0.50
Metridia longa CV 91 1.78 0.07 1.69 2.00 0.67 0.04 0.56 0.75
Metridia longa CVI females 15 2.37 0.26 2.00 2.88 0.87 0.09 0.75 1.06
Metridia longa CVI males 9 1.69 0.07 1.56 1.75 0.65 0.05 0.56 0.69
Oithona spp. CIV-V 11 0.77 0.05 0.68 0.84 0.20 0.02 0.16 0.22
Oithona spp. CVI females 2 0.72 0.71 0.73 0.20 0.19 0.21
Oithona spp. CVI males 7 0.89 0.05 0.82 0.96 0.20 0.01 0.19 0.22
Oncaea spp. CVI females 14 0.68 0.03 0.62 0.73 0.21 0.01 0.19 0.22
Pareuchaeta spp. CIV-V 4 3.07 0.77 2.51 4.20 1.11 0.37 0.78 1.64

Table 4
WP-2 replicate biomass variability experiment in the 0–18 m depth interval on 9 June. A dual version of WP-2 with two nets mounted side by side in a frame was used in 10
replicate hauls. Results are given as mean and standard deviation of dry mass (mg m3) in size fractions and total, determined for the left and right nets separately, for the two
series combined (left and right), and for the difference between them (right minus left). One left sample was lost (n = 9).

Series n <1 mm 1–2 mm Total

Mean SD CV% Mean SD CV% Mean SD CV%
Left 9 39.06 6.68 17.1 1.27 0.87 69.0 40.38 7.10 17.6
Right 10 37.83 6.04 16.0 1.10 1.34 121.2 38.93 6.45 16.6
L+R 19 38.41 6.20 16.2 1.18 1.10 93.0 39.62 6.61 16.7
RL 9 0.23 7.43 0.13 0.90 0.10 7.59

1-m Ring net (333 lm) contained few or none of the smaller forms,
but gave comparable counts for the larger fish eggs, decapod lar-
vae, and older copepodites of Calanus to the other nets. The highest
counts were generally obtained with WP-2 (200 lm) and MultiNet
(200 lm). The two nets gave very similar counts for the most abun-
dant taxon, the cladocerans. MultiNet tended to have somewhat
higher values for some of the more abundant taxa, such as Clado-
cera, Oithona, bivalve larvae, and polychaete larvae.
The 1-m2 MOCNESS (180 lm) gave lower values than WP-2 or
MultiNet for some taxa notably cladocerans, appendicularians,
and polychaete larvae (Fig. 22, Table 8). There was only one
MOCNESS profile counted, and it is therefore difficult to test for
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18 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42

4.4. Comparison of WP-2, MultiNet, MOCNESS, and BIONESS over the

whole water column

Fig. 12. Plot of variability in species counts (as coefficient of variation = standard
deviation/mean  100%) vs. the determined numerical abundance for the various
taxa identified in the 20 replicate samples from the 10 hauls with double WP-2 net
(200 lm) (A.v. Humboldt station 9012).
statistical significance. Due to the low accuracy associated with
low abundance and low counts (see Figs. 7 and 12), some of the dif-
ferences, particularly for less abundant taxa, could be by chance.
A cluster analysis based on similarity across the taxonomic
counts showed that the 1-m Ring Net with coarser mesh was
markedly different while the other nets clustered at similarity of
70% or higher (Fig. 23). The replicate samples of the same gear clus-
tered at about 80–90% similarity, with the two MOCNESS replicates
in a separate cluster, while the WP-2 and MultiNet samples oc-
curred as a mixed group with no clear separation.
Vertical profiles over the whole water column (to about 375 m
depth) were obtained with WP-2 (200 lm), MultiNet (200 lm),
MOCNESS (180 lm), and BIONESS (333 lm) on 5 June. The portion
of the profiles in the upper 100 m has been included as replicates
in the comparisons described above (Figs. 21–23). The results for
the whole water column and the layer deeper than 100 m are sum-
marized in Table 7B and shown in Fig. 24.
Over the whole water column (0–375 m), the WP-2 net gave the
highest biomass for the smallest size fraction (<1 mm), while Mul-
tiNet gave somewhat lower value than for MOCNESS (Fig. 24 A).
For the medium size fraction (1–2 mm), MOCNESS collected con-
siderably more than WP-2 and MultiNet, leading also to a higher
total. This reflected mainly differences in sampling deeper than
100 m (Fig. 24 B).
The coarser meshed BIONESS (333 lm) gave less biomass than
MOCNESS (180 lm) for the smallest size fraction (<1 mm) and also
somewhat lower values for the medium (1–2 mm) fraction
(Fig. 24). BIONESS collected more of the largest fraction (>2 mm)
leading to a slightly higher total for BIONESS. This reflected primar-
ily catch of some larger shrimps with the BIONESS.
4.5. Comparison of MOCNESS, BIONESS, and LHPR over the whole
water column
Vertical profiles were obtained on 11 June with MOCNESS
(180 lm) and BIONESS (333 lm) from about 375 m depth and with

Table 5
Summary of biomass results (mg dry mass) in three size fractions, three groups, and the total, for samples obtained with various gears for the whole water column (0–375 m) and
0–100 m, 100–375 m, and 200–300 m depth layers on 10 to 12 June. A. Biomass per m2. B. Biomass per m3. Note that fish, shrimp and krill are included in the largest size fraction
(>2 mm) in A but not in B. MOC – MOCNESS; LHPR – Longhurst-Hardy Plankton Recorder; BIO – BIONESS.

Gear Depth (m) Size fraction

<1 mm 1–2 mm >2 mm Fish Shrimp Krill Total
n Mean CV% Mean CV% Mean CV% Mean CV% Mean CV% Mean CV% Mean CV%
(A) Biomass m2
MOC 180 Day 0–375 3 1615.2 37.7 1058.7 16.9 1039.1 64.6 64.7 96.8 49.6 129 42.2 76.7 3713 8.9
MOC 180 Night 0–375 1 1873 1346.5 745.7 46.7 0 67.1 3965.2
LHPR 200 0–300 1 1516.5 345 104.5 1986.8
LHPR 200 0–375 1 1574.4 413.2 136.7 2111
BIO 333 Day 0–375 2 945.2 53.7 599.1 4.3 456.1 29.6 79.1 2.8 92.3 63.5 93.1 49.2 2000.4 17.4
BIO 333 Night 0–375 1 1134.9 526.3 840.8 118.2 332.5 92.4 2502
WP-2 200 0–100 2 1237.1 130.9 32.8 1335.4
WP-2 400 0–100 2 319.4 192.9 111.5 623.9
MOC 180 Day 0–100 3 1173.4 34.8 167 19.7 618.1 148.3 0 0 0 1958.6 34.4
MOC 180 Night 0–100 1 1247.9 467 516.6 46.7 0 67.1 2231.5
LHPR 200 0–100 1 1006.9 192 56.9 1214
BIO 333 Day 0–100 2 842.7 230.8 123.7 0 0 0 1197.3
BIO 333 Night 0–100 1 998.6 202.2 428.5 118.2 0 92.4 1629.4
MOC 180 Day 100–375 3 441.8 46.8 891.7 17.1 421 60.3 64.7 96.8 49.6 129 42.2 76.7 1754.5 25.7
MOC 180 Night 100–375 1 625.1 879.5 229.1 0 0 0 1733.7
BIO 333 Day 100–375 2 102.5 368.2 332.4 79.1 92.3 93.1 803.1
BIO 333 Night 100–375 1 136.3 324 412.3 79.8 0 332.5 872.6
LHPR 200 100–375 1 567.5 221.2 79.8 897
MOC 180 Day 200–300 3 131.8 23.7 567.5 42.9 150.9 61.9 52.1 88.3 5.4 173.2 31.6 84.9 850.2 35.6
MOC 180 Night 200–300 1 288.3 481.8 119 0 0 0 889.1
MOC 333 Day 200–300 1 53.3 187.9 209.3 114 56.2 7.3 450.5
MOC 333 Night 200–300 1 51.6 227.1 108.7 13.7 37.6 1.4 387.5
BIO 333 Day 200–300 3 41.4 34.5 161.6 39.7 105.8 28.7 40.4 68.8 21 85.5 27.2 44.5 308.8 26.8
BIO 333 Night 200–300 2 44.8 132.8 152.5 16.8 109.6 0.6 330.1
(B) Biomass m3
MOC 333 Day 200–250 4 0.52 52.3 1.49 23.8 0.36 38.5 0.93 83.7 0.03 200 0.04 200 3.37 29.5
MOC 333 Day 250–300 4 0.54 66.3 2.27 93 0.27 78.8 1.35 83.2 1.09 183.4 0.11 200 5.64 98.4
MOC 333 Night 200–250 4 0.52 27.7 1.5 27 0.82 38.9 0 0.47 183.1 0 3.31 26.2
MOC 333 Night 250–300 4 0.51 77.5 3.04 81.4 0.3 99 0.27 200 0.29 132.9 0.03 200 4.44 83.7
BIO 333 Day 200–250 8 0.48 88.3 1.57 80.7 0.26 77.9 0.77 125 0 0.23 114 3.31 72.9
BIO 333 Day 250–300 10 0.44 43.7 2 62.5 0.12 89.7 0.68 197 0.38 162.2 0.19 207 3.82 58.4
BIO 333 Night 200–250 4 0.16 9.6 0.7 31.8 0.33 32.8 0.18 200 1.25 94.9 0.01 200 2.64 60.2
BIO 333 Night 250–300 4 0.14 70.3 1.32 21.3 0.1 87.2 0.49 61.9 0.61 87.9 0.01 200 2.67 7.7
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42 19

Table 6
Abundance (No. Ind. m3) of zooplankton taxa (individuals m-3) obtained with MOCNESS 180 lm and 333 lm and BIONESS 333 lm tows taken at night between 10 and 12 June
from 200–250 m and 250–300 m depth layers.

Species or group Width (mm) Depth (m) MOCNESS 180 lm MOCNESS 333 lm BIONESS 333 lm
n Mean Range n Mean SD Range n Mean SD Range
Total individuals 200–250 2 57.2 41–73 4 38.0 17.3 29–64 4 14.9 4.8 10–21
250–300 2 103.3 64–143 3 47.4 13.7 38–63 4 26.0 6.0 17–31
Microcalanus 0.18 200–250 2 18 4–32 4 0 0–0 4 0 0–0
250–300 2 31.9 21–43 3 0.09 0.16 0.0–0.3 4 0 0–0
Small copepodsa 0.2 200–250 2 3.8 1.3–6.4 4 0.66 0.48 0.21–1.33 4 0 0–0
250–300 2 6.9 6.6–7.2 3 0.22 0.26 0.0–0.5 4 0 0–0
Medium-sized copepodsb 0.28–0.35 200–250 2 0.32 0.25–0.40 4 1.11 1.48 0.32–3.32 4 0.03 0.04 0–0.07
250–300 2 1.40 0.69–2.1 3 0.64 0.42 0.36–1.12 4 0.19 0.14 0.0–0.30
Metridia CIV-VI 0.48 200–250 2 6.70 5.9–7.5 4 4.25 1.13 2.7–5.2 4 4.36 1.15 2.7–5.2
250–300 2 10.20 8.7–11.7 3 12.4 3.84 9.9–16.8 4 6.57 2.22 3.9–8.6
Calanus CIV-VI 0.56 200–250 2 13.20 11.9–14.5 4 8.79 1.18 7.3–10.1 4 4.85 1.79 3.0–7.3
250–300 2 32.30 17.8–46.9 3 23.3 5.94 18.8–30.0 4 17 4.29 11.5–21.9
Other large copepodsc >0.5 200–250 2 1.69 1.45–1.94 4 1.54 0.73 0.95–2.60 4 0.71 0.36 0.35–1.21
250–300 2 1.39 0.58–2.21 3 0.98 0.24 0.82–1.26 4 0.67 0.11 0.58–0.82
Cladocera 0.26 200–250 2 4.35 3.2–5.5 4 10.4 8.75 5.1–23.4 4 0.21 0.17 0.1–0.5
250–300 2 7.47 1.7–13.2 3 4.11 0.68 3.4–4.8 4 0.62 0.3 0.3–1.0
Ostracoda 0.39 200–250 2 3.77 0.8–6.8 4 2.74 0.64 1.9–3.3 4 1.77 0.6 0.9–2.2
250–300 2 2.76 2.4–3.1 3 0.87 0.77 0.1–1.7 4 0.4 0.33 0.2–0.9
Small invertebrate larvaed 0.23–0.28 200–250 2 3.66 2.8–4.5 4 3.3 2.33 1.8–6.8 4 0.08 0.09 0.03–0.22
250–300 2 4.53 1.4–7.7 3 1.67 0.54 1.2–2.3 4 0.14 0.13 0.00–0.30
Larger larvaee 0.34–0.9 200–250 2 0.41 0.0–0.8 4 0.51 0.61 0.0–1.3 4 0.06 0.05 0.0–0.1
250–300 2 0.85 0.4–1.4 3 0.3 0.25 0.1–0.6 4 0.11 0.08 0.1–0.2
Appendicularians 0.74 200–250 2 0.00 0.0–0.0 4 0 0.0–0.0 4 0 0.0–0.0
250–300 2 1.52 0.4–2.7 3 0.04 0.04 0.0–0.1 4 0 0.0–0.0
Siphonophores 200–250 2 0.20 0.1–0.3 4 1.98 2.02 0.3–4.5 4 2.14 1.02 1.5–3.7
250–300 2 1.35 1.1–1.6 3 1.36 0.64 0.7–1.9 4 0.15 0.18 0.1–0.4
Fish eggs 1 200–250 2 0.20 0.0–0.4 4 1.04 0.3 0.8–1.5 4 0.42 0.5 0.1–1.2
250–300 2 0.18 0.0–0.4 3 0.39 0.22 0.1–0.6 4 0.09 0.1 0.0–0.2
Chaetognaths 200–250 2 0.05 0.0–0.1 4 0.86 0.43 0.6–1.4 4 0.26 0.09 0.2–0.4
250–300 2 0.08 0.03–0.14 3 0.52 0.47 0.4–1.0 4 0.09 0.07 0.0–0.2
a
Oncaea, Oithona, copepod nauplii, Pseudocalanus CI-III, and Temora.
b
Acartia, Pseudocalanus IV-VI, and Metridia CI-III.
c
Scolecithricella minor, Paraeuchaeta, and others.
d
Gastropods, cirripeds, bivalves, and polychaetes.
e
Echinoderms, cyphonautes, and decapods.

LHPR (200 lm) from about 300 m (Fig. 2). Only one LHPR profile
was taken for biomass determination. Depth-integrated biomass
values are summarized in Table 5 and shown in Fig. 25. For com-
parison, the LHPR profile has been extrapolated to 375 m by using
the biomass concentrations at about 280–300 m also for the 300–
375 m layer.
The LHPR collected comparable biomass of the smallest size
fraction (<1 mm) to that collected by MOCNESS (180 lm), both
for the whole water column and for the upper and deeper layers
separately (Fig. 25). LHPR collected less biomass of the medium
(1–2 mm) and considerably less of the largest size fraction
(>2 mm), also in comparison with BIONESS. Over the whole water
column, the LHPR collected slightly over half the total biomass col-
lected by MOCNESS. The LHPR gave comparable total biomass to
that obtained with BIONESS (333 lm), reflecting higher collection
of the smallest fraction, particularly in the deeper layer (Fig. 25).
The species counts obtained in profiles of MOCNESS (180 lm),
BIONESS (333 lm), and LHPR (200 lm) over the water column
are shown are Fig. 26. BIONESS collected few or none of the small
copepods (Micro/Pseudocalanus, Oithona, Oncaea, Temora) and
bivalve larvae, whereas the counts were more comparable for the
larger forms like Calanus and Metridia. These results are broadly
consistent with those obtained in the previous comparisons for
the 200–300 m depth layer (see Figs. 18 and 19, Table 6).
The LHPR sampler gave higher counts for cladocerans, but lower
counts for smaller forms such as Micro/Pseudocalanus, Oithona, and
Oncaea compared to the MOCNESS (Fig. 26). It gave lower counts
also for some of the larger forms, e.g. Metridia, Calanus, cyphona-
utes, and echinoderm larvae. Most of the taxa showed lower counts
in the LHPR than in the MOCNESS samples. There is a tendency of
lower ratio for counts between the two gears with decreasing
width of the organisms (Fig. 27) but this trend is not statistically
significant.
There were no marked differences between the day and night
profiles either with MOCNESS or BIONESS (Fig. 25). This supports
the previous interpretation that vertical migration of mesozoo-
plankton was of limited extent. Comparing the finer meshed MOC-
NESS (180 lm) with the coarser meshed BIONESS (333 lm),
MOCNESS collected more of the smallest (<1 mm) and medium
(1–2 mm) size fractions, most notably in the deeper layer. This
was reflected in higher total biomass collected by MOCNESS.
4.6. Comparison of MOCNESS and BIONESS at 200–300 m
Repeated tows with MOCNESS and BIONESS, both equipped
with 333 lm meshed nets, were made between 200 and 300 m
depth on 10 June. For each tow, replicate nets were used in the
depth layers 200–250 m and 250–300 m (Fig. 2). Mean values
and variability (coefficient of variation) for dry mass (mg m3) of
3 size fractions, 3 macrozooplankton (or micronekton) groups,
and total, are summarized in Table 5B. Depth-integrated values
of biomass m2 for the 200–300 m layer are also summarized in
Table 5A and shown in Fig. 28. Here the values for BIONESS include
the section between 200 and 300 m in the profiles obtained on 11
June. Also included are the values for 200–300 m for MOCNESS
with 180 lm meshed net on 11 June.
BIONESS collected similar or somewhat less biomass in the
smallest size fraction (<1 mm) than did MOCNESS with 333 lm.
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20 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42

Fig. 14. Coefficient of variation (standard deviation/mean  100%) vs. biomass (A)
and volume filtered (B) for all the samples obtained in the inter-comparisons
between gears.

Fig. 13. Coefficient of variation (standard deviation/mean  100%) for determina-

tion of depth integrated biomass with different sampling gears. (A) 0–100 m depth
interval, 5–6 June. (B) Whole or part of water column, 10–11 June.

The night values for BIONESS for this fraction were lower than for
MOCNESS on 10 June (Table 5B), but a higher value on 11 June
made the mean for BIONESS fairly similar to MOCNESS (Fig. 28).
For the medium size fraction (1–2 mm), BIONESS tended to collect
less biomass than did MOCNESS (333 lm). The MOCNESS with
180 lm net collected considerably more than BIONESS or MOC-
NESS with 333 lm nets for these two size fractions (Fig. 28; see
also Fig. 17).
The largest size fraction (>2 mm) as shown in Table 5A and
Fig. 28 (depth-integrated per m2) includes the fish, shrimp, and
krill that are also shown separately. The variability is fairly high
for this fraction and for the three macrozooplankton or micronek-
ton components (see Figs. 13 and 14). There is also vertical migra-
tion that contributes to the variability. The MOCNESS and BIONESS
with 333 lm nets produced similar values for the >2 mm fraction
as did also the MOCNESS with 180 lm nets (Fig. 28). The highest
value was found for the day samples with MOCNESS (333 lm),
reflecting a high value for fish. There was a pattern that fish and Fig. 15. WP-2 mesh size comparisons for tows taken on 10 June. Biomass in size
krill were caught predominantly during day, probably reflecting fractions obtained in two replicate tows with nets of 55, 100, 200, and 400 lm mesh
upward migration out of the 200–300 m layer during night (see size.

Fig. 31). For shrimp there was no clear difference between day
and night. Excluding fish, shrimp, and krill, the remaining part of
the >2 mm fraction tended to be higher for the MOCNESS
(333 lm) than for BIONESS (Table 5B).
The MOCNESS (333 lm) collected somewhat higher total bio-
mass than did BIONESS. This reflected mainly the higher catch of
the 1–2 mm size fraction (Fig. 28).
Taxonomic analysis has been carried out for the night samples
obtained with MOCNESS (333 lm) and BIONESS (333 lm) (Ta-
ble 6). Mean values for the nets in each depth interval (200–
250 m and 250–300 m, n = 3 or 4) for species or groups are shown
in Fig. 18. The night samples were chosen as the case with lowest
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
Fig. 16. WP-2 mesh size comparisons for tows taken on 10 June. Species composition (average of two replicate hauls) of samples collected with nets of 55, 100, 200, and
400 lm mesh size. Same samples as in Fig. 15.
Fig. 17. Comparison of biomass in size fractions and categories (fish, krill, shrimp)
collected with MOCNESS nets with 180 lm and 333 lm mesh size for (A) 250–
300 m and (B) 200–250 m depth intervals.
probability of visual detection and escape by zooplankton from the
gear.
There was a clear difference in the taxonomic composition of
the samples collected by the two gears. Several of the smaller
forms were either not caught or caught in low numbers in BIONESS
21
(333 lm) while being caught in MOCNESS (333 lm). This was the
case for cladocerans, Oithona, Pseudocalanus, Temora, Acartia, young
copepodites (CI-CIII) of Calanus and Metridia, cyphonautes, and cir-
riped and bivalve larvae (Fig. 18A and B). Other and larger forms
were caught in more comparable numbers in the two nets. These
included ostracods, older copepodite stages (CIV-VI) of Calanus
and Metridia, Scolecithricella minor, other copepods, and siphono-
phores. The total number of copepods sampled was about 40% low-
er for the BIONESS compared to the MOCNESS (Fig. 18C). Eggs of
the small fish Maurolicus mülleri and chaetognaths were caught
less effectively in the BIONESS.
Cluster analysis of replicate samples based on species counts
generally separated the samples obtained with MOCNESS 180
and 333 lm and BIONESS 333 lm into separate groups, and also
separated the samples obtained from the two depth layers (200–
250 and 250–300 m) obtained with the same gear (Fig. 29). Excep-
tions were one of the deepest MOCNESS 333 samples (No. 8) which
had low similarity with the other replicates, and one of the MOC-
NESS 180 lm samples that grouped with the MOCNESS 333.
The ratio between species counts obtained with BIONESS and
MOCNESS, both equipped with 333 lm mesh, showed a clear
decreasing trend with decreasing width of the organisms, most
notably for copepods (Fig. 30). The figure suggests that escapement
by extrusion through the BIONESS net relative to the more slowly
towed MOCNESS starts for organisms with a width around 0.8–
1 mm. Calanus finmarchicus stages CI-III were not collected by the
BIONESS, while stage IV was collected at about 50% compared to
MOCNESS.
4.7. Comparison of MOCNESS and BIONESS for sampling of
macrozooplankton and micronekton
Individuals of krill, shrimps, and small fish were sorted from the
>2 mm size fraction of catches obtained with MOCNESS and
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22 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42

Table 7
Comparisons of dry mass (mg) values per m2 (mean, standard deviation, and coefficient of variation for n replicates) for three size fractions and the total for samples obtained
with various zooplankton nets in (A) the upper 100 m and (B) over the whole water column, during the first part of the study period (5–6 June). BON - BONGO (20 and 60 cm); MN
– MultiNet; MOC – MOCNESS; BIO – BIONESS; RN – 1-m Ring net; 180, 200 and 333 are mesh sizes in lm; Du - dusk, N - night.

Gear Depth (m) Size fraction

<1 mm 1–2 mm >2 mm Total
n Mean SD CV% Mean SD CV% Mean SD CV% Mean SD CV%
(A) Upper 100 m – 5–6 June
WP-2 200 0–100 4 840.5 295.1 35.1 89.9 43.2 48.1 42.5 58.5 137.7 972.9 361.7 37.2
BON20 0–100 4 579.9 415.2 71.6 128.3 78.6 61.2 18.3 12.4 68.0 726.5 492.8 67.8
BON60 0–100 4 775.8 114.3 14.7 104.3 12.4 11.9 66.6 7.0 10.6 946.7 109.3 11.5
MN 200 0–100 3 980.3 350.5 35.8 101.6 29.9 29.5 463.3 633.7 136.8 1545.2 747.7 48.4
MOC 180 0–100 6 646.7 292.0 45.2 70.9 38.4 54.2 67.6 66.9 99.0 773.2 375.8 48.6
MOC 180 0–100 5 714.8 268.0 37.5 78.8 37.1 47.1 79.6 67.3 84.6 873.2 318.6 36.5
MOC 180 0–100 N, 4 763.6 115.1 15.1 122.7 61.5 50.2 112.8 71.7 63.6 999.0 222.3 22.3
MOC 180 0–100 1 402.5 140.7 75.6 618.9
BIO 333 0–100 Du/N, 1 103.9 218.3 102.8 425.0
RN 333 0–100 3 22.1 11.7 53.1 52.7 21.6 41.0 61.3 25.3 41.2 136.1 41.0 30.2
Gulf-V 300 0–100 1 171.2 56.3 74.3 301.7
WP-2 200 0–25 4 601.4 152.8 25.4 40.5 37.5 92.5 14.6 29.1 200.0 656.5 174.4 26.6
MN 200 0–25 3 710.0 211.6 29.8 34.1 8.7 25.5 432.8 644.1 148.8 1176.8 623.8 53.0
MOC 180 0–25 6 570.1 291.3 51.1 32.1 23.7 73.7 39.0 48.2 123.7 641.2 336.4 52.5
MOC 180 0–25 5 641.1 261.2 40.7 35.2 25.0 71.1 46.8 49.5 105.8 723.1 301.9 41.7
MOC 180 0–25 N, 4 677.9 100.0 14.8 22.8 4.6 20.0 16.0 11.1 69.3 716.7 109.6 15.3
WP-2 200 25–100 4 239.1 149.0 62.3 49.4 30.8 62.3 27.9 30.2 108.1 316.4 198.1 62.6
MN 200 25–100 3 270.3 160.4 59.3 67.5 29.3 43.4 30.5 28.2 92.3 368.3 161.4 43.8
MOC 180 25–100 6 76.6 29.3 38.2 39.7 18.1 45.6 28.7 32.1 112.0 145.0 66.3 45.7
MOC 180 25–100 N, 4 85.7 15.4 18.0 99.9 62.4 62.4 96.7 61.1 63.2 282.4 133.1 47.1
MOC 180 25–100 1 135.8 83.0 327.1 545.9
BIO 333 25–100 1 53.2 95.8 71.0 220.0
(B) Whole water column – 5 June
WP-2 0–375 2 1265.2 309.7 111.8 1686.6
MN 200 0–375 2 473.5 481.8 106.0 1061.3
MOC 180 0–375 1 718.0 1628.8 161.2 2585.6
BIO 333 0–375 Du/N, 1 342.0 1515.7 817.3 2675.0
WP-2 200 100–375 2 182.4 217.4 36.0 435.7
MOC 180 100–375 1 315.5 1488.1 85.6 1966.8
BIO 333 100–375 Du/N, 1 238.1 1297.4 714.5 2250.0

BIONESS, and the biomass of these three groups were determined
separately (see Fig. 4). The data obtained over the study period
with each of these two multiple net samplers have been pooled
and are shown relative to sampling depth in Fig. 31. Reflecting gen-
erally low numbers, the variability in the biomass determination of
these groups was large (see Fig. 13 B) and there were many sam-
ples with zero catch.
BIONESS tended to catch these groups of macrozooplankton or
micronekton better than MOCNESS. This was reflected in higher
percentages of samples with catch and higher mean biomass
caught (Table 9).
Krill showed evidence of vertical migration and were caught in
the upper 100 m mainly at night-time, whereas the catches during
day-time were mainly obtained deeper than 100 m (Fig. 31A and
B). For the samples from deeper than 100 m during the day-time,
almost twice as many of the BIONESS hauls contained krill com-
pared to the MOCNESS hauls (45 vs. 26%) and the mean biomass
was three times higher (0.27 vs. 0.09 mg dw m3; Table 9). There
were few night-time hauls with BIONESS from the upper 100 m,
but 2 out of the three hauls contained krill with five times higher
mean biomass compared to MOCNESS (Table 9). Meganyctiphanes
norvegica was the predominant krill species caught with most indi-
viduals between 26 and 40 mm in length (Fig. 32).
Catches of shrimps were pelagic species of the genera Pasiphaea
and Sergestes, with Pasiphaea being the predominant. Both the spe-
cies P. tarda and P. multidentata were identified. Also the shrimps
showed evidence of vertical migration with catches displaced
somewhat shallower in the water column during night than during
the day (Fig. 31C and D). Few shrimps were caught from the upper
100 m either day or night. For the hauls deeper than 100 m,
BIONESS caught shrimps 1.5–2 times as frequent as did MOCNESS
(Table 9). The mean biomass of shrimps collected by BIONESS
was slightly higher than the MOCNESS value for the day-time
samples and four times higher than MOCNESS for the night-time
samples.
Catches of fish consisted mainly of northern lanternfish (Ben-
thosema glaciale), but also some individuals of Müller’s pearlsides
(M. muelleri) (3–5 cm) and herring larvae were caught. Fish were
most frequently caught deeper than 100 m, but some individuals
were also caught in the upper 100 m during night indicating verti-
cal migration (Fig. 31E and F). For the day-time series from deeper
than 100 m, the two gears gave comparable results both for fre-
quency (about 28%) and biomass (about 0.4 mg m3), while for
the night-time series the BIONESS had five times higher frequency
(38 vs. 7%) and three times higher mean biomass (0.22 vs.
0.08 mg m3) than did MOCNESS (Table 9).
With krill, shrimp, and fish caught more effectively with BION-
ESS than with MOCNESS, one could expect also a greater propor-
tion of large individuals to be caught if the ability to escape the
slower MOCNESS increases with the size of the individuals. There
was little evidence for this, however, in length frequency distribu-
tions in catches from the two gears (Fig. 32). This could partly be
due to the low number of individuals that form the basis for these
size frequency distributions, being 14–15 for the MOCNESS series
and 18–34 for the BIONESS series. The measured individuals were
obtained in BIONESS and MOCNESS samples taken on 10 and 11
June when the depth interval between 200 and 300 m or the whole
column was covered (see Fig. 2).
The size distributions of M. norvegica caught by BIONESS and
MOCNESS showed no clear difference, and while the highest
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42 23

Fig. 18. (A and B) Comparison of species composition in samples collected with MOCNESS 180 lm and 333 lm and BIONESS 333 lm mesh-sized nets for 200–250 m and
250–300 m depth intervals combined. (C) Comparison of the abundance of large, medium, and small sized copepods caught by the same nets. Size is based on width (D2; see
Fig. 5 and Table 3): large >0.4 mm; medium 0.2–0.4 mm; small <0.2 mm (see Table 6). The number of nets used to compute the average is given above each histogram.

relative numbers were found in the 31–35 mm range for BIONESS
vs. 26–30 mm for MOCNESS, the mean size was essentially the
same (Fig. 32 A). The shrimps ranged in size from 23 to 81 mm
in length with a tendency of bimodal size frequency distribution.
There was a small difference in the length frequency distributions
of Pasiphaea spp. caught with the two gears due to the fact that
BIONESS caught a few large individuals >70 mm vs. none for the
MOCNESS (Fig. 32B). The fish B. glaciale ranged in size from 21 to
52 mm length. Here the modal length was higher for BIONESS than
for MOCNESS, but MOCNESS caught more of the largest individuals
and the mean size was essentially the same (Fig. 32 C).
4.8. Comparison across all gears
The biomass values obtained during the latter part of the study
period (10–11 June) tended to be larger than those obtained during
the first part (5–6 June) (Tables 5 and 7), indicating an increase in
zooplankton biomass over the period. A direct comparison of
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24 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
Fig. 19. Ratio of abundance of copepods and other zooplankton taxa collected with
MOCNESS (333 lm)/MOCNESS (180 lm) vs. average individual width of the
organisms (from Table 3). Based on the data from 200 to 300 m shown in Fig. 18.
Four data points have been identified as outliers; regression line based on corrected
values (minus outliers). (1 = cladocera; 2 = Temora; 3 = Metridia; 4 = Calanus).
Fig. 20. Comparison of gears in the upper 100 m, 5–6 June. Dry mass (mg m2) in
size fractions and total. Effects of removal of two outliers are shown: 1 – MultiNet, 2
– MOCNESS (6 June). See Table 7 for further details.
biomass levels across the gears for the whole period is therefore
not attempted. However, there were no clear qualitative changes
in the zooplankton composition over this period. We have there-
fore used aggregated data on the ratio between biomass and
numerical abundance over the study period to compare and con-
trast the sampling performance of the different gears (Tables 10
and 11).
The total biomass divided by the total numerical abundance for
a sample has the unit of biomass per individual and can be taken as
an indicator of the average size of the individuals sampled. This
parameter is sensitive to the catch of both small and large individ-
uals, but in different ways. One or a few large individuals may add
substantially to the total biomass, but have little influence on the
total numbers. Catch of small individuals may on the other hand
contribute much to the numbers but little to biomass.
The ‘average biomass per individual’ generally increased with
depth (see Fig. 8C), reflecting the change in the zooplankton com-
munity between the upper and deeper layers (see Table 2). The
‘average biomass per individual’ obtained with WP-2 net with
200 lm mesh changed little in the upper 100 m (3.1–4.6 lg ind1),
Fig. 21. Comparison of WP-2, MultiNet, and MOCNESS (day and night series) in (A)
0–25 and (B) 25–100 m, 5–6 June. Biomass as dry mass (mg m2) in size fractions
and total.
but increased substantially below 200 m (33 lg ind1) (Table 10).
Mesh size between 55 and 400 lm with WP-2 hauls from the
upper 100 m had a moderate effect on the ‘average biomass per
individual’, changing from 3.7 to 7.4 lg ind1 from the smallest
to the largest mesh size (Table 10).
The ‘average biomass per individual’ for the different gears are
summarized in Table 11 with mean values and statistics given sep-
arately for the samples obtained from the upper 100 m and deeper
than 100 m. The means and statistics for the corresponding bio-
mass and abundance values are also included in Table 11. Note that
these latter mean values are not strictly comparable across gears
due to the strong vertical gradients in biomass and abundance,
with much of the biomass found in the upper 50 m and most of this
again concentrated to the upper 12 m (Figs. 8–10). The more than
two times higher mean biomass and abundance values for the WP-
2 net than for the other gears in Table 11 are thus largely reflecting
the higher number of short hauls taken in the upper 12 or 18 m
with this gear compared to the others.
The ‘average biomass per individual’ showed large variation
across the gears spanning a range of about a factor of 10 (Table 11),
which by comparison exceeded the range of variation found with
the different mesh sizes of the WP-2 net (Table 10). The lowest
‘average biomass per individual’ in the upper 100 m was found
with WP-2 (200 lm) followed by the MultiNet (200 lm), while
the largest average values were obtained with the Ring-Net and
BIONESS both equipped with 333 lm mesh (Table 11).
The Gulf-V sampler, despite its coarse mesh (300 lm), had the
third smallest ‘average biomass per individual’ in the upper
100 m and the lowest in the series from >100 m, although here
there was only a single Gulf-V tow (Table 11). The LHPR
(200 lm) sampler had intermediate values for ‘average biomass
per individual’, being about 1/3–1/2 of the mean values obtained
with MOCNESS (180 lm).
The minimum values for the ‘average biomass per individual’
were generally higher (by a factor of from <2 to >10) for the coarse
meshed nets (300 and 333 lm) compared to the finer meshed nets
(180 and 200 lm). The maximum values were also generally high-
er for the coarse meshed nets (by a factor of from <2 to about 8)
with two exceptions. The Gulf-V (300 lm) had low maximum va-
lue, which could reflect the low number of tows, while the
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
Fig. 22. Comparison of taxonomic composition in 0–100 m for MOCNESS (180 lm), MultiNet (MN, 200 lm), WP-2 (200 lm), and 1-m Ring Net (RN, 333 lm) 5–6 June.
MOCNESS with 180 lm mesh had high values comparable to or
exceeding those for BIONESS (333 lm), RingNet (333 lm) and
the MOCNESS with 333 lm mesh (Table 11). The range (from min-
imum to maximum) of average biomass per individual was there-
fore larger for MOCNESS with 180 lm, spanning almost two orders
of magnitude for the samples from the upper 100 m, compared to
one order of magnitude or less for the other gears (Table 11).
The ‘average biomass per individual’ metric is shown vs. the
opening area of the net and the towing speed in Fig. 33. The net
systems with large opening area have the highest values for this
metric, with the coarser meshed BIONESS (333 lm) having 20–
50% higher values than MOCNESS (180 lm). The high-speed sam-
plers with small opening area have comparable or somewhat larger
values than the WP-2 and MultiNet (200 lm). The Gulf-V sampler
(300 lm) with the smallest opening, has lower biomass per indi-
vidual value than the LHPR (200 lm), despite its coarser net.
5. Discussion
5.1. General comments about the sampling design, methods, and
nature of the community sampled
5.1.1. Methods and sample variance
In the gear intercomparison study, there are a number of factors
that contributed to variance and errors in the results. These include
25
the in situ structure of distribution of zooplankton (even, random,
or patchy), gear handling during and after sampling, calculation of
volume filtered (including use and calibration of flow meters),
sample transfer and processing from net to laboratory, splitting
and subsampling, and analyses for determination of biomass and
species composition.
A critical determinant of quantitative zooplankton sampling is
the relationship between the open mesh area of the net and the
area of the mouth opening. Previous studies have highlighted the
importance of having an open mesh area to mouth opening ratio
that exceeds 3.0 in order for a net to have filtration efficiency
greater than 85% (Smith et al., 1968). In fact, this ratio should ex-
ceed 5.0 or higher for most coastal and ocean sampling conditions
to attain this efficiency for the duration of a tow. The nets used in
this study adhered to this criterion.
We followed a procedure where each net sample was split in
two halves, one for biomass determination and the other for anal-
ysis of taxonomic composition (Fig. 4) during the steady calm con-
ditions in the fjord. The biomass samples were processed by
different teams on the two vessels, following the same procedures,
while drying and weighing were done by the same technicians at a
local laboratory. The biomass data from replicate sampling with a
double WP-2 net in the upper 18 m on 9 June gave a low variance
with a coefficient of variation (CV) of only 17% (SD as percentage of
the mean; Table 4). This would include environmental changes
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26 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42

Table 8
Summary of results of species counts of zooplankton groups for upper 100 m, collected with MOCNESS (180 lm), MultiNet (200 lm), WP-2 (200 lm), and 1-m Ring net (333 lm)
on 5–7 June.

Group of organisms Gear n Mean CV% Range

Total individuals MOC 180 2 133,627 76,000–191,000
MN 200 3 451,828 38.0 258,000–586,000
WP-2 200 4 315,663 28.4 227,000–439,000
Ring Net 333 3 6198 24.9 4400–7100
Cladocerans MOC 180 2 50,290 32,000–67,000
MN 200 3 185,904 42.1 96,000–241,000
WP-2 200 4 140,239 25.3 108,000–190,000
Ring Net 333 3 114 11.7 105–130
Small copepodsa MOC 180 2 38,250 22,000–54,000
MN 200 3 57,670 38.7 35,000–80,000
WP-2 200 4 31,961 31.5 24,000–47,000
Ring Net 333 3 7 173.2 0–21
Medium-sized copepodsb MOC 180 2 16,288 3700–28,900
MN 200 3 19,150 22.8 14,500–23,200
WP-2 200 4 12,867 26.3 9900–17,100
Ring Net 333 3 129 116.8 6–300
Large copepodsc MOC 180 2 1330 380–2280
MN 200 3 1772 78.9 840–3400
WP-2 200 4 2618 63.3 730–4700
Ring Net 333 3 782 41.0 440–1070
Small invertebrate larvaed MOC 180 2 9434 6700–12,200
MN 200 3 24,064 52.3 9600–32,400
WP-2 200 4 9947 40.4 6200–15,100
Ring Net 333 3 40 99.3 6–84
Larger invertebrate larvaee MOC 180 2 13,141 9600–16,700
MN 200 3 15,190 39.8 9200–21,300
WP-2 200 4 15,836 40.1 6500–19,900
Ring Net 333 3 254 37.5 150–330
Appendicularians MOC 180 2 2164 1100–3300
MN 200 3 16,988 26.1 13,000–22,000
WP-2 200 4 16,027 62.0 6400–29,000
Ring Net 333 3 0 0–0
Fish eggs MOC 180 2 680 560–800
MN 200 3 878 69.3 330–1500
WP-2 200 4 555 25.4 540–740
Ring Net 333 3 1238 21.7 1070–1550
a
Microcalanus, Oithona, Oncaea, Pseudocalanus CI-III, and copepod nauplii.
b
Acartia, Temora, Pseudocalanus IV-VI, Calanus CI-III, Metridia CI-III and Centropages.
c
Calanus IV-VI, Metridia IV-VI, Scolecithricella minor, Paraeuchaeta.
d
Polychaetes, gastropods, bivalves.
e
Cirripedia, echinoderms, cyphonautes.

during the sampling period and all sources of error in sampling,
sample treatment, and biomass determination. Low variance was
also associated with biomass samples from MOCNESS (180 lm)
in the upper layer with high biomass on 6–7 June, particularly
for the night time samples (Fig. 10C and D). These results show that
the sample processing and analysis of dry mass were associated
with relatively low errors, since the variance would also include
the effect from in situ variation on zooplankton density.
The variance in dry biomass determination could be expected to
be higher for samples containing low biomass, for two reasons.
First, errors associated with sample treatment (e.g. rinsing nets,
subsampling, and weighing) could be relatively higher when sam-
ple size was small. Second, larger samplers that filter more water
and produce larger samples would also integrate across smaller-
scale patchiness and thus possibly produce lower variance (Wiebe,
1971, 1972). Getting into larger-scale patchiness, however, could
have the opposite effect. The vertical net tows with smaller vol-
umes (<60 m3) generally produced somewhat larger CV than the
towed 1-m2 framed nets (>300 m3) for the total and smaller size
fractions (<1 mm and 1–2 mm; Fig. 14B). This was likely due to
in situ patchiness in zooplankton distribution, given the low vari-
ance associated with replicate sampling from the upper layer re-
ferred to above.
The variance for the largest size fraction (>2 mm) was consis-
tently much higher than for the smaller size fractions (<2 mm),
with CV values often larger than 100% (Figs. 13 and 14). This is
probably due to the greater mobility of the larger organisms,
allowing them to aggregate and be more patchily distributed com-
pared to the smaller zooplankton. The categories of fish, shrimp,
and krill were also associated with high relative variance
(Fig. 13B), but this was most likely due to the effect of the low
numbers of these organisms that were caught.
The species counts of the replicate WP-2 samples from the
upper 18 m on 9 June were associated with increasing variance
with decreasing abundance of taxa (Figs. 7 and 12). This is a typical
effect and generally related to the counting error, which depend on
the numbers of specimen counted (Cassie, 1971). It follows a Pois-
son distribution with confidence limits CL at probability level of
p
p < 0.05 calculated in the following way: CL [%] = 1.95 (100/ N).
The counting error is about ±10% with 400 individuals counted,
and it increases significantly at smaller numbers. In our case this
was largely an effect of subsampling, since only a 1/50-fraction
of the WP-2 samples was counted. An abundance of 100 individu-
als m3 in the sea corresponded to a count of five individuals after
subsampling, which is equal to a counting error of 90%. The gener-
ally high and increasing variance with decreasing abundance of
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42 27

Fig. 23. Similarity from cluster analysis of WP-2 net (200 lm), MultiNet (MN, 200 lm), MOCNESS (180 lm), and 1-m Ring net (RN, 333 lm) in the upper 100 m, based on
abundances per m2 of various taxonomic groups (c.f. Fig. 22) for all available samples (identified by station no. and replicate haul no.).

Fig. 24. Comparison of WP-2 (200 lm), MultiNet (200 lm), MOCNESS (180 lm)
and BIONESS (333 lm) for (A) whole water column and (B) 100–375 m depth
interval for tows made on 5–6 June. Biomass as dry mass (mg/m2) in size fractions
and total.

taxa was thus mainly an artifact of the procedure of sample treat-

ment and subsampling prior to counting. The samples from the
two ships were counted by different teams, according to the stan-
dard procedures used by IMR in Bergen and IWO in Warnemünde.
This may have influenced the results of gear comparisons with re- Fig. 25. Comparison of biomass (dry mass mg/m2) collected with MOCNESS
gard to taxonomic composition to some extent. However, in both (180 lm), BIONESS (333 lm) and LHPR (200 lm) over the whole water column (0–
cases the fraction of sample counted was adjusted according to 375 m) and in the 0–100 m and 100–375 m depth layers, for tows made on 11 June.
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28 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42

Fig. 26. Comparison of species composition in samples obtained with MOCNESS (180 lm), BIONESS (333 lm) and LHPR (200 lm) over the whole water column (0–375 m)
for profiles taken on 11 June.

Fig. 28. Comparison of biomass for MOCNESS (180 lm), MOCNESS (333 lm) and
BIONESS (333 lm) for samples obtained from the 200–300 m layer on 10 June.
Biomass as dry mass per m2 in size fractions, groups, and total; the groups (fish,
shrimp, krill) are included also in the >2 mm size fraction.

Fig. 27. Ratio of LHPR (200 lm)/MOCNESS (180 lm) abundance of copepods and
other zooplankton in 0–350 m tows vs. individual width of the organisms.
the numerical density of zooplankton so that a minimum number
of individuals was counted in each case (see Section 2.3.2).
5.1.2. Study design considerations
Diel vertical migration occurred to limited extent by the meso-
zooplankton as evidenced by the day and night profiles of zoo-
plankton biomass (Figs. 9 and 10). Larger fauna exhibited
pronounced vertical migration as shown by the deep scattering
layer, which rose from around 150 m at day-time to within the
upper 50 m at night (Fig. 11). We interpret the scattering layer to
consist mainly of Müller’s pearlsides (Maurolicus mulleri), which
was by far the dominant species caught when the layer was tar-
geted for sampling by a young-fish trawl (data not presented).
Macrozooplankton (or micronekton) in the groups of krill, pelagic
shrimps, and small fish also exhibited clear vertical migration, with
krill moving from 100 to 300 m during day into the upper 100 m at
night (Fig. 31).
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
Fig. 29. Similarity from cluster analysis of MOCNESS (180 lm), MOCNESS (333 lm) and BIONESS (333 lm) in the 200–300 m depth strata, based on abundances per m2 of
various taxonomic groups (c.f. Fig. 18) for all available samples (identified by haul no. and replicate net no.).
Fig. 30. Ratio of BIONESS/MOCNESS (333 lm) abundance of copepods and other
zooplankters in 200–300 m tows vs. width of the individuals (see Fig. 5 and
Table 3).
Change in zooplankton distribution from vertical migration was
one factor that we considered in the design of our study. Deploy-
ment of the various gears inevitably takes some time, and even
with sampling from two ships, this limited the possibility to do
simultaneous sampling with replicates. We have in our analyses
grouped samples into day and night categories for comparison of
the various sampling nets. We have also grouped samples over
two consecutive days to allow a broader comparison of gears as de-
scribed in the methods.
The consistency in our results suggests that any effects of short-
term changes were small and that the grouping of samples and
comparisons across gears that were deployed at somewhat differ-
ent times during the day and the experimental periods appears to
be acceptable. Over the study period, there was an increase in the
zooplankton biomass from around 1000 mg dry mass m2 to about
29
1500 mg m2 in the upper 100 m of the water column (Tables 5
and 7). This change appeared over a period of about 5 days, sug-
gesting an increase in zooplankton biomass of about 10% per day.
Grouping data from two days may therefore have introduced an er-
ror of this magnitude. An alternative experimental design, which
we did not pursue, would have been to do a pair-wise comparison
of gears deployed at the same time from the two ships. Whether
this would have been a better design is difficult to judge, given that
other sources of error may have been of larger importance than the
short-term changes in zooplankton abundance and distribution.
5.2. Major factors influencing the catch in plankton nets
5.2.1. Effects of mesh size
Mesh size has an obvious and well-known effect on the catch of
zooplankton, with smaller forms passing through coarser net, but
being collected on finer mesh (e.g. Saville, 1958; Vannucci, 1968;
Nichols and Thompson, 1991; Gallienne and Robins, 2001; Her-
man, 2005). Our experiments with different mesh size showed
very clear and consistent results. In the mesh size comparison with
vertical hauls of WP-2 nets in the upper 100 m, the biomass col-
lected dropped markedly from 200 to 400 lm mesh due to much
lower capture of small organisms in the <1 mm size fraction
(Fig. 15). These were the dominant species collected with the finer
meshed nets (Evadne, Oithona, and Microcalanus; Fig. 16). There
was not so much difference between the zooplankton caught in
the 100 and 200 lm nets, but the biomass collected dropped off
markedly with the 55 lm net. This was probably due to lower fil-
tration efficiency and a larger pressure wave in the front of the net,
which led to more organisms avoiding the net. Thus the dominant
forms (Evadne, Podon, Oithona, Microcalanus, Calanus, and cirriped
larvae) were collected in lower abundance in the 55 lm than in
the 100 or 200 lm nets. The main difference between the 100
and 200 lm nets was that small copepods (in the other copepods
category) and copepod nauplii were collected in the finer, but lar-
gely passed through the 200 lm mesh (Fig. 16).
The comparison of MOCNESS equipped with 333 and 180 lm
nets, sampling from the deep layer between 200 and 300 m depth,
revealed a very pronounced difference. The finer mesh collected
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30 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42

Fig. 31. Vertical distribution of the biomass of krill (A and B), shrimp (C and D) and fish (E and F) caught with BIONESS and MOCNESS from different depth intervals during
day and night. Mid-points of the depth interval sampled are plotted; open markers are zeros.

Table 9
Catches of krill, shrimp, and fish by MOCNESS and BIONESS in the upper 100 m and below 100 m. Frequency of samples with catch and biomass as dry mass per m3.

Depth Krill Shrimp Fish

Period Gear n % of Samples Dry mass (mg m3) % of Samples Dry mass (mg m3) % of Samples Dry mass (mg m3)
Mean SD Mean SD Mean SD
Upper 100 m
Day BIONESS 14 7 0.01 0.05 0 0.00 7 0.19 0.70
MOCNESS 55 2 0.00 0.03 2 0.00 0.01 5 0.02 0.17
Night BIONESS 3 67 0.98 0.93 0 0.00 33 0.79 1.36
MOCNESS 30 30 0.18 0.45 0 0.00 17 0.18 0.51
Below 100 m
Day BIONESS 40 45 0.27 0.42 30 0.30 0.85 28 0.43 0.94
MOCNESS 34 26 0.09 0.23 21 0.24 0.77 29 0.39 0.70
Night BIONESS 16 13 0.01 0.02 56 0.88 1.24 38 0.22 0.34
MOCNESS 14 7 0.01 0.03 29 0.21 0.49 7 0.08 0.29

more biomass in the <1 and 1–2 mm size fractions, with about
twice as high a total biomass as the 333 lm net (Fig. 17). This dif-
ference was due to low catch of small copepods (Microcalanus,
Oithona, Oncaea, Pseudocalanus, Temora) and bivalve larvae with
the 333 lm net (Figs. 18 and 19). There was a fairly consistent rela-
tionship between the relative catch in the coarser net (compared to
the finer) and the width of the organisms, with lower sampling
efficiency due to escapement through the mesh for organisms with
a width of <0.6 mm (Fig. 19). This corresponds approximately to
the size of C. finmarchicus copeodite stage CIV and Metridia lucens
CV (Table 3). Young copepodites of these or similar dominant cal-
anoid copepods in subarctic and boreal environments would there-
fore not be sampled representatively with a 333 lm net due to
escapement through the net. Similar conclusions have been
reached in other gear intercomparison studies (Pillar, 1984; Evans
and Sell, 1985; Nichols and Thompson, 1991; Gjøsæter et al., 2000).
5.2.2. Extrusion through the net mesh
Extrusion of animals through the net mesh was a strong deter-
minant of the performance of the net systems equipped with coar-
ser mesh netting. With faster towing speeds there is increased
filtration pressure on the meshes and increased escapement by
extrusion of the smaller animals through the meshes (Tranter
and Smith, 1968; Vannucci, 1968). The comparison of tows of
BIONESS and MOCNESS, both equipped with 333 lm meshed nets
but towed at different speeds (2–3 knots vs. 1–2 knots), with rep-
licate tows between 200 and 300 m depth, produced a clear differ-
ence that we ascribe to extrusion of organisms. The faster-towed
BIONESS gave about 25% less biomass compared to MOCNESS,
mainly for the two smaller size fractions (<1 and 1–2 mm)
(Fig. 28, Table 5). This difference was consistent over the day and
night comparisons and was statistically significant (ANOVA 2-
way; p = 0.03).
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42 31

collected by the BIONESS (Fig. 18, Table 6). Large copepods in con-
trast, were more comparable, although still collected in lower
numbers in BIONESS compared to MOCNESS (Fig. 18, Table 6).
The ratio of zooplankton taxa in BIONESS compared to MOCNESS
showed a clear relationship with width of the organisms
(Fig. 30), consistent with the effect of extrusion through the mesh
of smaller organisms. The results suggest that the loss of organisms
due to extrusion in BIONESS starts at a width of around 0.8 mm,
and that all organisms smaller than about 0.2–0.3 mm are lost
through the mesh. This would lead to undersampling of all but
the largest adult C. finmarchicus and Metrida, and no or very low
sampling of Oithona, Oncaea, Microcalanus, and young copepodite
stages of Acartia and Temora, by faster-towed BIONESS compared
to MOCNESS when both are equipped with 333 lm mesh (Table 3).
The LHPR sampler was included in comparisons with MOCNESS
(180 lm) and BIONESS (333 lm) in profiles over the whole water
column. Compared to MOCNESS with slightly finer mesh (180 vs.
200 lm), the LHPR produced comparable biomass values for the
smallest size fraction (<1 mm), but lower for the larger size fractions
(1–2 and >2 mm) (Fig. 25, Table 5). Over the whole water column,
LHPR gave slightly more than half the biomass collected with MOC-
NESS. There may at least be four contributing factors for this differ-
ence. The samples from LHPR for biomass determination was
treated differently from those from the other gears in that the zoo-
plankton were rinsed and scraped off the gauze used to collect them
in the LHPR cod end. Each sample was smaller compared to the
other gears because the water column was divided into more units
(about 10 m intervals). The difference in sample handling prior to
dry mass determination could have influenced the results. For in-
stance, larger individuals may have been squeezed on the gauze
and lost part of their biomass through leakage of oil droplets and
other body fluids. It is also possible that mechanical damage of spec-
imens collected on the gauze and transferred for further analysis
(Halliday et al., 2001) could have resulted in fragments that contrib-
uted to the biomass of the smallest size fraction (<1 mm). The three
other factors could have been the different mesh size, extrusion of
small zooplankton with the faster towed LHPR, and avoidance of
the smaller mouth opening by larger zooplankton. The latter factor
is considered in more detail below (see Section 5.2.3).
Mesh size selection and extrusion would lead to a loss of the
smallest zooplankton, and is opposite to the trend shown by the bio-
Fig. 32. Length frequency distributions of (A) Meganyctiphanes norvegica, (B) mass data in this MOCNESS/LHPR comparison. However, this could
Pasiphaea spp., and (C) Benthosema glaciale in catches obtained with BIONESS and be due to the handling of the samples for biomass determination.
MOCNESS.
The taxonomic counts were done on separate profiles and are inde-
pendent from the treatment used for getting biomass data. They
Table 10 would therefore not be influenced by such error. The counts show
Summary of ‘average biomass per individual’ (lg; total biomass/total abundance)
the largest difference between the LHPR and MOCNESS for the
collected with WP-2 net based on vertical tows to specific depths.
smallest copepods (Oithona, Oncaea, and Microcalanus/Pseudocal-
Depth (m) Mesh size (lm) Biomass per individual anus) and cyphonautes (Fig. 26). The ratio for taxa caught in LHPR
n Mean SD vs. MOCNESS showed a weak but not significant trend with width
18 200 19 3.77 0.98 of the zooplankton organisms (Fig. 27), indicative of an effect from
25 200 4 3.10 2.06 mesh size and/or extrusion in the LHPR compared to MOCNESS.
100 200 6 4.59 1.66 The ratio between net opening to mouth opening was 9 for LHPR
200 200 2 6.05 2.50 and 6 for MOCNESS and the towing speed was 3–4 knots and 1–2
390 200 2 32.74 2.38
100 55 2 3.73 2.50
knots respectively (Table 1). The higher opening ratio (by 50%)
100 100 2 3.78 1.24 was probably not sufficient to counter the effect of the even higher
100 200 6 4.59 1.66 towing speed (by 100% or more) with the LHPR, making it likely that
100 400 2 7.44 1.32 there was some extrusion of smaller zooplankton forms through the
mesh of the LHPR. In addition the net was somewhat coarser than
that used in the MOCNESS (200 vs. 180 lm). The design of the
The lower catch of biomass with BIONESS was reflected in an cod-end and the ratio of filtration area to mouth opening for the col-
even larger difference in numbers of zooplankton collected, with lecting gauze within the recorder box itself is also a factor for possi-
roughly half the total number of individuals caught by BIONESS ble extrusion with collection with LHPR (Haury et al., 1976).
compared to MOCNESS (Table 6). This difference was due largely In the comparisons between MOCNESS (180 lm), MultiNet
to few cladocerans, smaller copepods (Calanus CI-III, Metridia (200 lm), and WP-2 net (200 lm), there were some trends in
CI-III, Pseudocalanus, Temora, Acartia), and invertebrate larvae our data that were indicative of more extrusion in the MOCNESS
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32 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42

Table 11
Statistics (mean, standard deviation, minimum, and maximum for n replicates) for total biomass (mg dry mass m3), total abundance (number of individuals m3), and ‘average
biomass per individual’ (total biomass/total abundance; lg dry mass per individual) in samples collected with different gears from the upper 100 m and deeper than 100 m depth
layers.

Depth (m) Statistic Gulf V WP-2 MN LHPR MOC MOC BIO RN

300 lm 200 lm 200 lm 200 lm 180 lm 333 lm 333 lm 333 lm
Biomass (mg DM m3)
<100 m n 3 29 16 10 19 4 3
Mean 3.50 31.08 12.31 12.41 11.89 11.97 1.36
SD 3.30 14.57 14.17 12.86 14.85 11.89 0.41
Min. 1.49 1.64 0.69 3.21 0.84 1.99 1.03
Max. 7.30 53.37 47.42 40.90 51.51 28.36 1.82
>100 m n 1 4 8 20 12 7 20
Mean 0.76 1.33 3.43 3.33 6.30 4.40 2.91
SD 1.10 1.85 2.01 4.99 2.26 1.46
Min. 0.31 0.31 0.78 0.52 2.70 0.83
Max. 2.53 5.61 7.88 15.98 9.25 6.67

Abundance (No. Ind. m3)

<100 m n 3 30 20 10 19 4 4
Mean 378 8491 3499 1368 1176 668 247
SD 452 4405 5007 2206 1412 690 436
Min. 105 376 85 249 52 62 22
Max. 900 19,002 19,957 7359 3895 1384 900
>100 m n 1 4 8 20 12 7 20
Mean 84 67 132 130 105 42 19
SD 5 70 49 73 15 8
Min. 63 31 77 6 29 1
Max. 74 252 230 261 64 32

Biomass/no. individuals
<100 m n 3 29 16 10 19 4 3
Mean 11.9 4.0 5.9 15.0 23.3 28.5 47.0
SD 4.0 1.2 4.9 11.6 32.7 16.7 8.3
Min. 8.1 2.4 1.6 5.6 2.8 11.4 38.7
Max. 16.1 7.3 22.8 44.3 145.3 50.1 55.2
>100 m n 1 4 8 20 12 7 18
Mean 9.1 19.5 28.9 27.9 74.7 111.1 144.7
SD 15.4 19.2 19.0 55.9 60.5 59.8
Min. 4.6 10.0 7.0 15.4 64.6 74.0
Max. 34.4 62.1 84.2 218.5 243.6 292.2

(180 lm) compared to the other two nets. In the comparison of
biomass from the upper 100 m, MOCNESS collected about 20% low-
er biomass in the smallest size fraction (<1 mm) than did the WP-2
net and MultiNet (Table 7A, Fig. 20). The difference was most pro-
nounced for the depth layer 25–100 m where the biomass was
lowest (Fig. 21). The species counts were inconclusive in this com-
parison (Fig. 22). There were lower values for some taxa in the
MOCNESS samples (cladocerans, polychaete larvae, and appendi-
cularians), but this could possibly be due to the different proce-
dures used for the species counts in the two laboratories. Small
forms that showed clear differences in the mesh size comparisons
(e.g. copepod nauplii, Oithona, Oncaea, Microcalanus; see Figs. 16
and 18) did not show a clear difference between MOCNESS and
the other two nets (Fig. 22).
5.2.3. Avoidance
The issue of avoidance of the net by mobile forms of zooplank-
ton has been recognized for a long time and there have been many
studies addressing this problem (see Table 1 in Clutter and Anraku,
1968). Most of these studies have focused on fish larvae or krill,
which are macrozooplankton with mobility and size that extend
into the micronekton category of pelagic organisms. Surveys and
studies of fish larvae have been an important part of fisheries
investigations and there is a large body of literature addressing
sampling efficiency and avoidance for this group of organisms
(e.g. Bjørke et al., 1974; Brander and Thompson, 1989; Shima
and Bailey, 1994; Huse et al., 1996; Pepin and Shears, 1997; Potter
et al., 1990). Fish larvae use visual cues to detect and escape from
an approaching net in an avoidance reaction that can be seen as
part of their anti-predator behavior. There are several studies dem-
onstrating that avoidance is size-dependent, with larger fish larvae
or juveniles escaping the net more effectively than smaller larvae
(Brander and Thompson, 1989; Huse et al., 1996).
Several sampling gears have been designed specifically to catch
fish larvae representatively and quantitatively. These have been
either high-speed samplers with faster approach aiming to catch
the larvae by surprise, or large-opening nets which require a longer
swimming distance for the larvae to escape (e.g. Nash et al., 1998;
Methot, 1986). The Gulf sampler has a number of variants (Nash
et al., 1998) and there have been concerted efforts to examine
the efficiency of these (Anonymous, 1991; Schnack, 1992; Brander
et al., 1993).
Krill appear also to use visual cues to detect and avoid
approaching nets (Wiebe et al., 1982). ‘‘Blinding’’ them with strong
flashing light has been shown to substantially increase the catches
of krill species in the Northwest Atlantic (Sameoto et al., 1993) and
Antarctic krill (Wiebe et al., 2004). Our results suggest clearly that
the faster towed BIONESS collected krill and other macrozooplank-
ton (or micronekton) more effectively than the MOCNESS. This was
reflected in both higher frequency of samples with catch and high-
er average biomass of krill, shrimp, and fish (Table 9). The LHPR,
however, that was also towed at speeds exceeding 3 knots, did
not catch more large macrozooplankton than either the MOCNESS
or the BIONESS in tows close in space and time. Its smaller mouth
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
Fig. 33. ‘Average biomass per individual’ (total biomass/total abundance) obtained with different gears plotted vs. towing speed and mouth opening of the nets for (A) upper
100 m and (B) below 100 m depth.
opening may have resulted in more avoidance. Perhaps surpris-
ingly, the higher efficiency of catch was not associated with any
clear difference in the size frequency distribution of the three
groups collected with the two gears (Fig. 32). This suggests that
within the size range of organisms collected (2–8 cm) there was
no marked difference in size-selectivity associated with avoidance.
It is possible that the low numbers of organisms collected and the
pooled data from somewhat different sampling regimes for the two
gears (see Fig. 2) may have obscured any pattern in the data. How-
ever, this lack of a size frequency difference has been observed in
other studies (Wiebe et al., 1982, 2004; Sameoto et al., 1993).
The low density of krill and other macrozooplankton in the
study area made it difficult or impossible to address the issue of
avoidance for the organisms with the other nets used. Very few
krill were collected in the upper 100 m during day time (Fig. 31)
when most of the hauls and intercomparisons with the other gears
were made (see Fig. 2). Adult krill were not collected in any of the
hauls with WP-2 or MultiNet deployed over the water column. A
WP-2 net would filter about 100 m3 of water over a 400 m deep
water column, whereas the MultiNet hauled obliquely filtered
about 160 m3. With a density of krill considerably less than 1 per
100 m3 (as sampled by MOCNESS 180 lm; Table 2), there would
be a low likelihood of collecting them in a single tow with these
gears. The MultiNet was used four times over the whole water col-
umn and a few more hauls were made deeper than 100 m. The to-
tal volume of water filtered in these hauls was about 500 m3,
which suggest that the lack of catch of any krill could have been
by chance as well as the animals avoiding the net.
Our results provide limited information on possible avoidance
by copepods for the various nets. In the comparison between MOC-
NESS and BIONESS, large copepods were collected in lower num-
bers by the BIONESS (Table 6, Fig. 18). This is likely due to
33
extrusion through the net of the faster-towed BIONESS, as sug-
gested by the ratio of catch by the two gears (equipped with the
same mesh size 333 lm) vs. individual width (Fig. 30). In the com-
parisons between the MOCNESS (180 lm) and the other gears with
comparable mesh (WP-2, MultiNet, and LHPR; 200 lm), the data
did not show any clear differences that would suggest avoidance
as a factor. The large copepods (width >0.5 mm) were relatively
rare compared to small copepods (width about 0.2 mm) (by a fac-
tor of about 20 in the upper 100 m; see Table 6). They were there-
fore associated with fairly high counting errors with the
subsampling procedures followed (Figs. 7 and 12).
Copepods are instrumented with a sophisticated array of sen-
sory bristles and organs that detect vibrations and hydrodynamic
signals from their surroundings (Kiörboe, 2008 and references
therein). They may potentially jump in an escape response to avoid
being caught by a net like they would an approaching predator.
Small copepods can respond to danger by rapid escape within
3 ms with speeds exceeding 80 cm per second (Buskey et al.,
2002). However, such speeds are attained only over very short
range, and more sustained swimming speed is typically a few cm
per second. The reaction distance of copepods beginning an escape
response to hydrodynamic disturbances is typically of the order of
about 1 cm or less (Buskey et al., 2002; Kiörboe, 2008). The major-
ity of Acartia spp. individuals tested responded at a millimeter dis-
tance. Responses at distances of more than one centimeter were
not observed (Kiörboe, 2008). Towing velocities of most of the
gears are in the range of 1.5 kn or about 50 m min1 respectively
(Table 1). Consequently, avoidance by copepods is not to be ex-
pected, when comparing their reaction distance with the speed
of the approaching net. This is supported by video recordings that
showed that most copepods were in normal feeding orientation
and not in an escape posture 0.5 m in front of a MOCNESS
 Author's personal copy
34 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
(Broughton and Lough, 2006). Krill, on the other hand, appear to
use visual cues to detect a net and the evidence suggests that they
begin an avoidance reaction 1–2 m in front of a 1-m2 MOCNESS
and at a further distance in front of a 10-m2 MOCNESS (Wiebe
et al., 1982).
5.3. Inter-comparability and characteristics of zooplankton sampling
gear
5.3.1. Gear intercomparisons
At the sea-going workshop we compared several different ver-
tical, depth-stratified, and high-speed zooplankton sampling sys-
tems. Escapement and extrusion through the net have been
demonstrated to be main factors affecting the results in our study.
Avoidance of the net by mobile forms has not been so clearly ex-
pressed in our results, apart from the higher catch of krill and other
macrozooplankton (or micronekton) by the BIONESS compared to
MOCNESS. Likely, all gear used considerably under sampled the
mobile larger forms.
When nets with similar mesh size (180–200 lm) were com-
pared, they gave fairly similar estimates for zooplankton biomass
over depth intervals. The largest range of zooplankton nets were
compared for the upper 100 m on 5–6 June. These results showed
moderate differences between the sampling gear, with a range for
the total biomass (after correction for two outliers) from about 730
to 1200 mg dry mass m2 (Fig. 20, Table 7). The difference among
the gear was statistically significant. However, removing the Bon-
go-20 and 1-m Ring Net, which had the lowest biomass, from the
test, removed also the statistical difference among the remaining
gear (WP-2, MultiNet, MOCNESS (180 lm), and Bongo-60; ANOVA
results). The results for WP-2, MultiNet, and MOCNESS (180 lm) in
the upper 25 m (each with 3–6 replicates) were even closer, rang-
ing from about 650 mg dry mass m2 for WP-2 to about
800 mg m2 for MultiNet. In the same comparison the MOCNESS
(180 lm) gave lower biomass (for the smallest size fraction
(<1 mm) and the total) for the 25–100 m layer where biomass gen-
erally was lower (Fig. 21). In a different comparison over the whole
water column, with fewer replicates (1–2), MOCNESS gave higher
biomass (by about a factor of 2) compared to WP-2 and MultiNet.
This was due to higher biomass in the 1–2 mm size fraction below
100 m (Fig. 24, Table 7).
While the results for the biomass comparisons show some dif-
ferences, these are not consistent, but rather reflect variability in
the results. The results from the main comparison of the gears with
replicate hauls in the upper 100 m suggest that the WP-2, Multi-
Net, MOCNESS (180 lm), and Bongo-60 produce comparable re-
sults for size fractioned and total dry mass biomass of
zooplankton, within the limits of the errors in our procedures.
The most notable difference in the biomass comparisons was
the lower catch of larger size fractions (1–2 and >2 mm) in the
LHPR (Fig. 25), which could be an artifact due to sample treatment
since the LHPR samples were size-fractioned after being rinsed off
the plankton gauze (see Section 2.2). There was a tendency in the
data that MOCNESS collected somewhat more of the large biomass
size fraction (>2 mm) compared to the smaller nets (WP-2 and
MultiNet) (Tables 5 and 7, Figs. 21 and 24), although this trend
was not so pronounced. The comparison of total biomass vs. the to-
tal number of individuals collected (‘average biomass per individ-
ual’) showed that the MOCNESS consistently collected more
biomass relative to numbers than did the WP-2 and MultiNet
(Fig. 33). While part of this could be due to a counting artifact
(see below), it suggests that the MOCNESS collected more large
individuals and possibly less of the smallest forms due to extru-
sion. Gjøsæter et al. (2000) found a similar result in a comparison
of a WP-2 net vs. a MOCNESS (200 lm) from routine sampling in
the Barents Sea. They found no significant difference between the
two gears in terms of total biomass. However, the WP-2 net col-
lected more biomass in the smallest size fraction (<1 mm) while
the MOCNESS collected more biomass in the largest size fraction
(>2 mm). Similar results were obtained from the same area in com-
parisons between MOCNESS and vertical hauls with WP-2 or Juday
nets (Hassel et al., 1991; Skjoldal et al., 1992), although in these
cases the MOCNESS was equipped with a coarser net (333 lm).
The similar results obtained for biomass is supported by the
species counts for WP-2, MultiNet, and MOCNESS (180 lm) for
the upper 100 m from the same sample series that were also
broadly comparable (Fig. 22). The replicate samples from WP-2
and MultiNet clustered together at high similarity values, as did
the two MOCNESS samples. However, the MOCNESS samples dif-
fered significantly from the former (Fig. 23). This was due to lower
counts of cladocerans, polychaete larvae, appendicularians, and
Centropages in the MOCNESS samples (Fig. 22). It is likely that this
is an artifact due to differences in the counting procedures be-
tween IMR and IOW rather than real differences between the sam-
ples. The cladoceran E. nordmanni has high surface tension and
may adsorb to surfaces, which may cause bias in the subsampling
prior to counting. We do not know at present where the error lies.
The abundance of cladocerans was about 1.5  105 individuals m2
as obtained with WP-2 and MultiNet (Table 8). The mean length of
Evadne was 0.64 mm. Using equations for relationships between
length and carbon (Postel et al., 2007), and C/dry mass ratio of
0.45, gives a mean weight of 3 lg dry mass per individual. This
makes up a biomass of Evadne of 450 mg dry mass m2, or around
50% of the measured total biomass in the upper 100 m (Table 7). If
the difference in counts by the two laboratories was real one would
expect a clear difference also in the biomass for the MOCNESS vs.
the other gear.
Increasing speed and mouth opening of the net are two main
principles used to improve the catchability of zooplankton nets
(Wiebe et al., 1982). Higher speed requires an increase in the rela-
tive filtering area to avoid reduced filtration efficiency and in-
creased pressure wave in front of the net. A cone to reduce the
mouth opening is therefore used in the Gulf-V and LHPR samplers
to counter the effect of higher towing speed. The other approach is
to increase the mouth opening in an attempt to reduce the escape
distance of zooplankton relative to the size of the opening. This ap-
proach is taken with the MOCNESS and BIONESS samplers,
although they are designed for towing at different speeds (Table 1).
We have projected the aggregated results obtained with the
various sampling gears that were compared in our workshop onto
a diagram with the two axes of towing speed and net mouth open-
ing (Fig. 33). The parameter is the total biomass divided by the to-
tal number of individuals collected, being an indicator for average
biomass per individual. The results show a relatively clear pattern.
The systems with large net opening (BIONESS, MOCNESS, and 1-m
Ring Net) have higher ‘average biomass per individual’ than the
other gear, while the high-speed samplers (LHPR and Gulf-V) have
similar or somewhat higher values than the WP-2 and MultiNet. In
order to interpret these patterns it is necessary to examine the re-
sults in more detail.
The BIONESS tended to have somewhat higher biomass/individ-
ual ratio than the MOCNESS. This was due mainly to extrusion
through the net of the faster towed BIONESS relative to MOCNESS
equipped with the same mesh 333 lm. The number of individuals
collected with BIONESS was only about half the number collected
with MOCNESS 333 lm and reflected loss of small individuals (Ta-
ble 6, Fig. 18). The biomass was more similar, although somewhat
lower for the two smaller size fractions (<1 and 1–2 mm) for the
BIONESS (Fig. 28). Thus the loss of small individuals, which con-
tributed relatively little to biomass, was the main reason for the
higher biomass/individual ratio for BIONESS compared to MOC-
NESS 333 lm. In the comparison between MOCNESS equipped
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
with 333 lm and 180 lm, the difference was due mainly to
escapement through the net of smaller individuals through the
coarser meshed net (predominantly the small copepod Microcal-
anus; Table 6, Figs. 18 and 19). The biomass was also lower for
the 333 lm MOCNESS for the smallest size fraction (<1 mm; Ta-
ble 5, Fig. 28) but proportionately less so than for the numbers, giv-
ing higher biomass/individual ratio.
The MOCNESS (180 lm) was towed at approximately the same
speed as the WP-2 and MultiNet, but has a mouth opening four
times larger (1 vs. 0.25 m2; Table 1). The biomass/individual ratio
was about 3–6 times higher for MOCNESS (180 lm) than for the
two other nets, which were fairly similar for this metric (Fig. 33,
Table 11). For the upper layer, much of this difference was due to
lower counts of cladocerans, appendicularians, and polychaete lar-
vae for the MOCNESS samples (Table 6, Fig. 22). Differences in sub-
sampling and counting procedures between the two labs (IMR and
IOW) could have contributed to this difference. Removing these
taxa from the calculation reduces the difference, but MOCNESS
(180 lm) still has higher biomass/individual ratio compared to
WP-2 and MultiNet. For the deeper layer, the difference is due
mainly to higher biomass in the MOCNESS (180 lm), while num-
bers of individuals are more similar (Tables 7B and 11, Fig. 24).
Extrusion of small forms could have played a role for the larger bio-
mass/individual ratio in MOCNESS, but the evidence for this is not
clear (see Section 5.2.2). It is possible that MOCNESS collects pro-
portionally more large individuals than the other two nets with
smaller opening. Larger zooplankton forms that occur in low den-
sity may have a higher probability of being caught by the MOC-
NESS that filters more water compared to the smaller nets. Also,
it is possible that avoidance by larger forms could be more pro-
nounced for the smaller nets than for the MOCNESS.
5.3.2. Size-selective loss by escapement and extrusion through the
mesh when sampling zooplankton communities
One clear result in our study was the size-selective loss of zoo-
plankton organisms through the mesh due to escapement and
extrusion. The catch of zooplankton taxa in a coarser-meshed vs.
finer-meshed MOCNESS (333 lm vs. 180 lm) declined with
decreasing width of the organisms due to escapement through
the mesh (Fig. 19). In a similar fashion, the relative catch with
the faster-towed BIONESS compared to MOCNESS when both were
equipped with the same mesh (333 lm), declined with decreasing
width (Fig. 30). We have projected both these declining relation-
ships (the regression lines from Figs. 19 and 30) onto the upper pa-
nel in Fig. 34, and compared them with the size ranges and means
for copepodite stages of various species of copepods (data from Ta-
ble 3) (Fig. 34, lower panel).
The line for escapement through the 333 lm mesh of MOCNESS
(compared to 180 lm) starts to show some loss in the size range of
older copepodites of C. finmarchicus (CIV) and Metridia spp. (CV).
The size for 50% loss corresponds to the widths for older copepo-
dites of relatively small calanoids such as Temora and Acartia
spp., and to CIII for C. finmarchicus. The size (width) for loss is
shifted up by about 0.1–0.2 mm due to extrusion with the higher
towing speed of BIONESS, corresponding to a shift from the size
of CIII to that of CIV for C. finmarchicus (Fig. 34). The diagonal of
the 333 lm mesh is 470 lm (in the ideal case of no width of the
threads). This is somewhat less than the size of about 0.5–
0.6 mm where some loss through the mesh starts to occur (Figs. 19
and 34), suggesting that some extrusion takes place by squeezing
the zooplankton through the mesh in addition to the passive loss
due to size (width) alone. The width at 50% loss for the ‘‘escape-
ment line’’ (for MOCNESS 333 lm) in Fig. 34 is approximately sim-
ilar to the mesh size.
Early studies showed that organisms larger than the mesh size
could pass through plankton gauze due to a combination of
35
compressibility of the organisms and flexibility of the meshes (Sa-
ville, 1958; Vannucci, 1968; Bernhard et al., 1973). Nichols and
Thompson (1991) studied size selection using samples of 4 cala-
noid copepods collected with high speed plankton nets with differ-
ent mesh sizes from 61 to 270 lm deployed in the North Sea. They
fitted a logistic equation model to the data, where 50% retention
occurred for copepod width being equal to the mesh size, and
95% and 5% retention occurred at copepod width being 1.33 and
0.67 times the mesh size. This is a fairly steep escapement line with
loss starting at a width about equal to the diagonal of the square
mesh size and with almost complete loss at about 2/3 of the mesh
size. For a 180 lm mesh, this would correspond to loss starting at a
copepod width of about 240 lm and almost all passing through the
mesh at a width of 120 lm. Similar mesh selection experiments
using the Continuous Plankton Recorder (CPR) by Hays (1994)
showed good agreement with the model of Nichols and Thompson
(1991).
A hypothetical escapement line for a 180 lm mesh net has been
included in Fig. 34, extrapolated from the empirical escapement
line for MOCNESS with 333 lm mesh compared to 180 lm. The
hypothetical line could be for MOCNESS with 180 lm compared
to 100 lm mesh, assuming that other factors such as configura-
tions and filtration efficiencies were comparable. The hypothetical
line for a 180 lm mesh net indicates that loss by escapement
would start at a width corresponding to that of old copepodite
stages of relatively small calanoids such as Temora and of stage CIII
of C. finmarchicus. The width for 50% loss by escapement would
correspond to the size of CI for C. finmarchicus, about stage CIII-
IV for Acartia and Pseudocalanus, and older copepodites of Microcal-
anus, Oithona, and Oncaea (Fig. 34).
The empirical escapement line for 333 lm mesh used in MOC-
NESS (and the extrapolated and hypothetical line for 180 lm
mesh; Fig. 34) is less steep than the empirically based model equa-
tion of Nichols and Thompson (1991). The line for 180 lm mesh in
Fig. 34 corresponds to a drop-off from 100% retention at 260 lm to
0% retention at 80 lm (compared to about 250 and 110 lm for the
Nichols and Thompson’s model). The difference could be by
chance, since the escapement regression line in Fig. 19 was associ-
ated with an r2 of 0.62. There may, however, be real differences in
the escapement between the high-speed samplers and copepod
species in the previous studies (Nichols and Thompson, 1991;
Hays, 1994) and the MOCNESS sampler and broader range of taxa
used in our study. This may require more detailed and careful stud-
ies in the future. The important point to note here is the strong sen-
sitivity to mesh selection by the use of zooplankton sampling gear.
Due to the general steepness of the escapement line and the addi-
tional effect of extrusion, small changes in zooplankton commu-
nity composition and size distribution within species and stages,
as well as differences in towing speed and other factors (e.g. sea
state and heave) during operation of the gears, may have substan-
tial effects on the size and stage composition in the lower end of
the size range of zooplankton collected.
The WP-2 net was suggested as a standard net after careful
studies and analysis by a UNESCO working party in 1968 (Working
Party 2, 1968) and was designed to give good filtration efficiency
and minimal extrusion. It is a simple and robust net for obtaining
mesozooplankton biomass and species composition over the
euphotic zone or other ecological meaningful depth intervals. The
WP-2 net is commonly used in routine surveys to map the horizon-
tal distribution of zooplankton biomass, e.g. by IMR in the Barents
and Norwegian Seas (Skjoldal et al., 2000; Melle et al., 2004) and by
IOW in the Baltic Sea (Wasmund et al., 2010a,b) and off Africa
(Hernándes-Léon et al., 1999).
The depth-stratified samplers with large net mouth opening
(1-m2 MOCNESS and BIONESS) produce larger samples stratified
over a larger portion of the water column. They require a larger ship
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36 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42

Fig. 34. The empirical escapement line for 333 lm mesh used in MOCNESS relative to 180 lm mesh (from Fig. 19), and the empirical extrusion line for BIONESS compared to
MOCNESS when both were equipped with the same mesh (333 lm; from Fig. 30). Also shown is the extrapolated and hypothetical line for 180 lm mesh relative to 100 lm
mesh. B. Size ranges and means for the widths (D2) of copepodite stages of various copepod species. The figure illustrates the strong sensitivity to mesh selection by the use of
zooplankton sampling gear (see text for more details).

to operate from than the simpler WP-2 net, but may provide a profile
of samples for description of zooplankton biomass distribution in
and below the euphotic zone in a single haul. These sampling sys-
tems have an advantage in that they sample better into the size
range of macrozooplankton. This is because they filter a larger
volume than smaller nets and therefore catch more of large forms
that occur in low density. In addition, avoidance by mobile forms
may be less for these larger nets which therefore sample more
representatively.
The faster-towed BIONESS catches more macrozooplankton and
micronekton than does MOCNESS (Table 9). At the same time it
loses more of the small mesozooplankton, and our results show
that extrusion can be considerable even for relatively large cope-
pods (Figs. 18, 19 and 34). For sampling of mesozooplankton, the
slower-towed MOCNESS therefore seems to be the best alternative,
while BIONESS has its strength for sampling macrozooplankton.
However, avoidance, for instance by krill, can be a problem also
for the faster BIONESS (Sameoto et al., 1993). Since there is strong
evidence that avoidance behavior is related to the visual acuity of
individuals, use of a ‘‘blinding’’ flashing light on a net system towed
to reduce extrusion may be optimal.
5.4. Previous gear intercomparison studies
A main finding in our study was that most of the nets provided
estimates of zooplankton biomass and species abundance that
were broadly similar. There is a fairly large body of literature on
zooplankton sampling gear inter-comparisons summarized by
Wiebe and Benfield (2003) in their Table 2. The following high-
lights those relevant to the present work. Several of the previous
studies are in general agreement with this studies findings,
whereas in other studies significant disparities have been uncov-
ered as described below.
Hernroth (1987) did a comparison of the Nansen net and the
WP-2 net and found that the Nansen net was grossly inferior to
the WP-2 net because of the very small mesh area to mouth open-
ing ratio (<3.0) that resulted in substantially lower catches in re-
peated Nansen net replicate tows. Dicenta et al. (1976) compared
WP-2, Hensen, FAO ‘‘standard’’, Juday (-Bogorov modified), and
20- and 60-cm Bongo nets equipped with various mesh sizes from
200 to 500 lm. In comparisons across all gears (except WP-2) with
500 lm mesh there was no significant differences in total plankton
volume. From their original data (in their Annex 2) it appears that
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H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
Juday and Hensen nets produced somewhat less plankton volume
than did the Bongo Nets. The WP-2 net was used with finer mesh
(200 lm) than the other nets, but gave similar plankton volume to
that obtained with Juday and Hensen nets (250 and 300 lm).
WP-2 with 200 lm and Bongo with 300 and 500 lm nets were
compared by Pillar (1984) for sampling of meso- and macrozoo-
plankton in the Benguela Current. The WP-2 net gave similar val-
ues for larger mesozooplankton groups such as large copepods
and krill furcilia larvae, to those obtained with the coarser-meshed
Bongo nets. The latter gave much lower catches of small copepods
due to escapement, but higher catches of juvenile and adult krill,
which avoided the WP-2 net. The WP-2 net gave comparable zoo-
plankton biomass to a pump system used with the same mesh. Pil-
lar (1984) found no evidence of avoidance by copepods or krill
larvae in any of the gear.
Casas and Durbin (2006) compared catches of copepods from
Georges Bank obtained with 1-m2 MOCNESS equipped with
150 lm mesh nets and a zooplankton pump where samples were
collected on a 35 lm mesh net. The pump was more effective in
collecting copepodite stages, but not adults of Pseudocalanus spp.
and also more effective in collecting stage CI of C. finmarchicus.
The authors considered that small stages and species of copepods
escaped through the 150 lm mesh and that, due to clogging of fi-
ner meshed nets, other sampling approaches such as a pump are
required in order to quantitatively determine the age structure of
the younger copepod stages.
Brander and Thompson (1989) compared MOCNESS, LHPR, and
a pump system for sampling of herring larvae. MOCNESS and LHPR
were found to collect quite similar amounts of total plankton mea-
sured as settled volume. However, the LHPR collected considerably
more herring larvae, of larger average size, than did MOCNESS,
which was considered to reflect greater avoidance of the larvae
for the latter gear. A comparison of the larger (10 m2) MOCNESS
with a pelagic young fish trawl also suggested avoidance of larger
cod juveniles (>35 mm) with the slower-towed MOCNESS (Potter
et al., 1990). For small and less mobile cod larvae (up to 6.5 mm
in length), Solemdal and Ellertsen (1984) found that MOCNESS
and a pump system gave comparable and representative catches,
while a Gulf III sampler was more effective at catching larger lar-
vae. A similar result was obtained in a comparison of Gulf III, a
MIK Ring-net, and MOCNESS, where the latter collected smaller
capelin larvae than did the Gulf III and MIK (Huse et al., 1996).
Stehle et al. (2007) compared an LHPR (with 42 cm diameter
mouth opening and 280 lm mesh) to a 60 cm diameter Bongo
net (with 335 lm mesh). Total LHPR zooplankton biomass was sig-
nificantly higher than Bongo net biomass and with the exception of
appendicularians and cladocerans, abundances of copepods, mys-
ids, euphausiids, decapod larvae, amphipods, siphonophores,
hydromedusae, chaetognaths and fish eggs also were consistently
higher in the LHPR samples. These results are at variance with
the results in the present study.
Ohman and Smith (1995) compared a 1-m diameter ring net with
60-cm and 71-cm diameter Bongo nets equipped with 505 lm mesh
to assess the effects of a methodological change in the CalCOFI mac-
rozooplankton time series from the ring net to the Bongo. The Bongo
nets caught an average of about 40% more biomass than the ring net.
In a further study of these nets, Rebstock (2002) concluded that for
31 copepod species and stages as well as a total female category,
there were no significant differences between the nets. The higher
biomass caught by the Bongo nets was attributed to avoidance of
the ring net with a bridle by larger zooplankton such as euphausiids.
In an intercalibration study of the SCOR, NORPAC, and Bongo nets,
McKinnell and Mackas (2003) found that all three nets, properly
equipped with flow meters, provided similar estimates of zooplank-
ton biomass. There are several other studies comparing sampling
efficiency with Bongo nets. Colton et al. (1980) compared 61-cm
37
diameter Bongo nets with 253 and 333 lm mesh towed at 1.5 and
3.5 knots. Biomass was higher in the Bongo tows with 253 lm mesh
when towed at 3.5 knots, but there was no difference between the
mesh sizes for tows at 1.5 kts. However, many zooplankton species
were more abundant in the smaller mesh at both towing speeds.
Variation in zooplankton retention rates thus resulted from differ-
ences in mesh size and tow speed. A 20-cm diameter Bongo with
165 lm mesh nets was compared to a 61-cm diameter Bongo with
333 lm and 505 lm mesh nets (Anderson and Warren, 1991). Early
life stages of C. finmarchicus (CI and CII) were under-sampled by the
larger Bongo with 333 lm and 505 lm nets relative to the smaller
Bongo with 165 lm mesh, but larger stages (CIV to CVI) were
under-sampled by the smaller Bongo.
Cook and Hays (2001) compared the sampling efficiency of a U-
Tow high-speed sampler (resembling a CPR with 18 mm opening
and fitted with a similar Plankton Sampling Mechanism) with a
WP-2 net. The U-Tow seriously underestimated the abundance of
zooplankton (by about 70%), but gave similar species and size com-
position. This suggested that avoidance and extrusion with the
high-speed sampler with small mouth opening were not the main
reasons for the loss. Instead, much of the water probably flowed
through the sampler underneath the mesh, as suggested by pump-
ing experiments in the laboratory (Cook and Hays, 2001). Compar-
ing time-series data obtained with the CPR and WP-2 from the
North Sea showed much lower abundance of zooplankton obtained
with CPR (Clark et al., 2001). It is therefore possible that CPR and
similar high-speed samplers underestimate zooplankton abun-
dance (Cook and Hays, 2001).
Batten et al. (1999) compared data obtained with CPR and LHPR
in the Celtic Sea. Biomass for the two gears (measured as displace-
ment volume for LHPR samples and calculated from abundance
data for CPR) were broadly comparable, although the LHPR bio-
mass for the 0–50 m depth interval tended to be substantially
higher than the CPR biomass sampled at 7 m depth (Fig. 1 in Batten
et al., 1999).
The LHPR (200 lm) was compared to results obtained with an
OPC in the present study (results reported by Halliday et al.,
2001). The concentration of particles and total biovolume sampled
by the LHPR were consistently lower compared to that ‘‘seen’’ by
the OPC (on average about 25%). This could have been due to detec-
tion of flocculent aggregates and detritus by the OPC, and also un-
der-representation of small zooplankton (<0.35 mm ESD) due to
extrusion through the mesh in the LHPR (Halliday et al., 2001).
Heath et al. (1999) obtained higher counts for particles in the size
range of C. finmarchicus with an OPC than those collected with an
ARIES net sampling system with a 200 lm mesh. The OPC increas-
ingly overestimated the zooplankton density with decreasing con-
centration, with an overall average factor for the overestimate of
about 2.4 for the survey as a whole. This was ascribed to the detec-
tion of detrital aggregates by the OPC (Heath, 2002). Hopcroft
(2002) also found that the OPC consistently counted 2 times or
more particles than were collected with plankton nets (WP-2 and
Bongo nets) in the size range of sensitivity for the OPC. Copepods
made up 80–95% of the zooplankton in this study off California.
The OPC failed to detect some of the prominent peaks created by
the larger copepods in the size spectra for the net samples, sug-
gesting avoidance by these larger forms to the intake tunnel of
the OPC (Hopcroft, 2002).
Other studies have shown a general agreement between cope-
pod abundance obtained with net sampling and particle abun-
dance recorded with OPC (Herman, 1992, 2002, 2005; Zhou and
Tande, 2002). Grant et al. (2000) compared LHPR (200 lm) with
an OPC in the waters around South Georgia in the Southern Ocean.
With careful interpretation and calibration of the OPC data, a fairly
good agreement in the results obtained with the two gears was
found. The OPC sometimes gave higher counts, particularly on
 Author's personal copy
38 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
the ascent portion of hauls. More extrusion through the LHPR net
due to higher towing speed could have contributed to this discrep-
ancy (Grant et al., 2000). Baumgartner (2003) compared OPC and
MOCNESS for sampling of C. finmarchicus stage CV in the Gulf of
Maine and Scotian Shelf area. Using the OPC-derived particle abun-
dance in the 1.5–2 mm equivalent spherical diameter range, a
fairly strong correlation with net-derived abundance was found.
This OPC size range corresponds to the upper half of the size range
for stage CV of C. finmarchicus and was chosen to reduce the influ-
ence by smaller copepods and detrital particles. The regression line
for the OPC vs. MOCNESS comparison was suggested as a calibra-
tion to predict Calanus CV abundance from OPC counts. After
adjustment (multiplying with 2 to include also the lower half of
the size range for stage V), this empirical calibration line fitted well
the data of Heath et al. (1999) of wintering CV (and CIV) C. finmar-
chicus from the Faroe-Shetland Channel. Baumgartner (2003)
found evidence of avoidance of the OPC tunnel opening by Calanus
CV in his study.
GLOBEC arranged a workshop on OPC in 2001 and one of the
topics considered was calibration and comparison with net-tow
samples. It was recognized that there were several important fac-
tors to take into account in this respect. These include avoidance of
animals from nets and the OPC, escapement and extrusion through
nets, coincident-counts in OPC measurements, and relationship be-
tween plankton aspect sizes and OPC measured size (Zhou and
Tande, 2002). The latter can be complex as illustrated by an over-
estimate of about 70% in the equivalent spherical diameter re-
corded by OPC for copepods due to their appendages (Hopcroft,
2002). The issue of detrital particles and organic aggregates (mar-
ine snow) is important and may lead to overestimates and obscur-
ing of the abundance of zooplankton as recorded with an OPC.
Inter-comparisons of nets as a function of mouth opening, mesh
size, and towing speed etc. provides a means of understanding the
relationships between the catch of zooplankton that the different
gear provide. However, the gear in all the different configurations
provide an observed ‘‘filtered view’’ of the actual pattern of distri-
bution and abundance of the zooplankton (see Fig. 5 in Haury et al.,
1978). Organisms that are fragile such as larvaceans, radiolarians,
and other gelatinous taxa can be observed in situ with a Video
Plankton Recorder (VPR) and compared to net catches to see the
extent of the bias associated with nets (Benfield et al., 1996; Den-
nett et al., 2002; Hartfield et al., 2004; Remsen et al., 2004; Brough-
ton and Lough, 2006). The results of these studies are quite
consistent. Nets collect the relatively hard-body zooplankton effec-
tively while the more fragile organisms are significantly under-
sampled relative to the abundances observed optically. In addition,
for small copepods and larval stages of the larger ones, small ptero-
pods, and other taxa too small to be retained effectively by a par-
ticular size mesh, the optical systems provide estimates of
abundance higher than provided by the nets. However, most opti-
cal systems scan very small quantities of water for each image and
the total water scanned by an optical system is usually a very small
fraction of the total volume filtered by a net system towed in a
comparable way. Thus for relatively rare mesozooplankton and lar-
ger macrozooplankton, the very low probability of being observed
in the optical systems can result in their being present in the net
samples, but not observed optically (Benfield et al., 1996; Brough-
ton and Lough, 2006). Thus, future studies will require both optical
and net systems used together to characterize the distribution and
abundance of meso- and macro-zooplankton species.
6. Summary and recommendations
Retention efficiency of zooplankton nets and results of gear in-
ter-comparisons are influenced by the species and size
composition of the plankton. Our study was carried out with a
coastal zooplankton community dominated by small mesozoo-
plankton species with cladocerans and copepods (Microcalanus,
Oithona, Calanus, and Temora) making up most of the biomass in
the upper layer. Macrozooplankton such as krill were captured in
low abundance and mainly in the lower part of the water column.
The phytoplankton density was typical for a post-spring bloom,
early summer situation with relatively low biomass, and clogging
of the nets generally played a minor role as a source of error (ex-
cept for very fine meshed nets). Mesh size of the net had a major
influence on the estimated zooplankton biomass for the zooplank-
ton community in our study. A 400 lm WP-2 net caught only
about half as much biomass as a 200 lm net. The species compo-
sition was also markedly different, with small cyclopoid (Oithona,
Oncaea) and young stages of calanoid copepods (Acartia, Pseudocal-
anus, Temora) virtually not sampled at all in nets with 333 lm
mesh while being the numerically dominant forms collected with
180 lm mesh. Our results have confirmed many previous studies
on mesh selection and demonstrate that there is a fairly consistent
and robust relationship between retention (or escapement through
the mesh) and the width of the organisms. The rule-of-thumb is
that about 50% of the organisms escape through the mesh at a
width equal to the mesh size, and that loss starts to occur and is
almost complete (5% and 95% loss through the mesh) at organism
widths that are about 1/3 larger and 1/3 smaller, respectively, than
the mesh size.
It should be noted that the relationship for escapement or loss
through the mesh as a function of organism width is fairly steep
and goes from complete to no retention over a linear size range
of about 2/3 the mesh size. Thus for a 180 lm mesh net, this would
be a range of about 120 lm, from 120 to 240 lm width of the
organisms. The underlying theoretical line is a logistic function
(Nichols and Thompson, 1991), but it can broadly be approximated
with a straight line over most of the range (remembering that there
is a tapering-off in both ends). The steepness of the slope of the
escapement line is such that there is a transition from complete
to no retention over a size (width) range corresponding approxi-
mately to two copepodite stages (e.g. CIII of C. finmarchicus being
fully retained while CI largely escapes through a 180 lm mesh).
While most of the previous studies on mesh selection have been
with copepods, our results (Figs. 19 and 30) suggest that width
of the organisms is a key parameter determining escape also for
other groups.
We do not recommend that results on biomass or species
counts obtained with a specific mesh size are corrected for the ef-
fect of mesh size selection in general. However, the relationship
between escapement and width of the organisms appears to be ro-
bust enough to allow an estimate of the error associated with loss
of organisms in the small end to be made. For some specific pur-
poses, such as comparisons across different studies or studies of
the population dynamics and life cycles of copepods, it may be
appropriate and more correct to do a correction than not. This
needs to be evaluated in each single case. The reason why we do
not recommend correction in general is that escapement may be
influenced by a number of factors such as variation in towing
speed, ship heave during operation, and regional and spatial varia-
tion in the size frequency distribution of given life stages of organ-
isms such as copepodite stages.
The effect of towing speed on the extrusion of smaller organ-
isms through the net can be substantial, as demonstrated in our re-
sults for the comparison of the MOCNESS and BIONESS towed at
about 1.5 and 3 knots, respectively (Figs. 18, 27 and 34). Faster
towing of a given net can cause a shift-up in the escapement vs.
width, leading to a greater loss of organisms through the mesh. It
is therefore an important element in a quality assurance system
for zooplankton sampling to maintain a constant and correct
 Author's personal copy
H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42
towing speed. While properly calibrated and mounted flowmeters
help to correct for differences in volume filtered due to differences
in speed during tows, the effect of extrusion is an additional source
of error that is not accounted for.
Different vertical, oblique, and multiple opening and closing net
systems (WP-2, Bongo, MOCNESS, MultiNet) produced similar esti-
mates of zooplankton biomass when operated with comparable
mesh-sized nets (180–200 lm) in our study. The older literature
is broadly consistent with this finding, although there are several
cases where differences have been documented due to differences
in design configurations and operation of the gears. We note the
importance of a sufficiently high ratio between mesh opening area
and the mouth opening area of the net (R ratio in Table 1) to obtain a
high filtration efficiency without excessive pressure on the net and
pressure wave in front of the net (bucket effect). This was a ‘‘truth’’
established by early pioneers in zooplankton research (e.g. Smith
et al., 1968) and is still a valid point to remember. The choice of a
particular net depends in the end on the purpose of the study and
on practical considerations such as costs, size of the ship, and
equipment to handle the gear. Simple nets such as WP-2 or Bongo
may be suitable for regular surveys of an area, particularly in shal-
low coastal waters and for the upper water layer, e.g. the euphotic
zone. Multiple opening and closing net systems such as MOCNESS
and MultiNet have their strengths in providing stratified samples
over the water column and also in providing larger samples with
higher chances of catching larger but less abundant forms of macr-
ozoplankton. These net systems are relatively heavy and generally
require a large ship equipped for their operation.
The choice of mesh size is an important issue. As demonstrated
in Fig. 34 many small copepods have widths around 0.2 mm, for in-
stance older copepodites of Oithona, Oncaea, Microcalanus and
Acartia, and younger copepodites of Pseudocalanus, Calanus, and
Temora. The retention of these forms would be sensitive to changes
in mesh size in this range. A 150 lm net would be expected to col-
lect substantially more individuals of these small copepods than
would a 200 lm net. 150 lm is therefore possibly the preferred
mesh size to be used in coastal waters with neritic zooplankton
communities such as was the case in our study in Storfjorden.
150 lm was the net mesh used for the US GLOBEC zooplankton
studies on Georges Bank (Durbin and Casas, 2006). Clogging be-
comes an increasing problem with finer meshed nets and
150 lm may be a reasonable compromise between better collec-
tion of small copepods on the one hand and not having significant
clogging on the other. In spring bloom situations with chain-form-
ing diatoms and/or colony-forming Phaeocystis, clogging may be a
serious problem, but this would be the case also with a slightly
coarser mesh, such as 200 lm.
Nets with 180 or 200 lm mesh are used routinely by many lab-
oratories including IMR and IOW in their monitoring and research
in the Barents, Norwegian, North, and Baltic seas. In boreal and
sub-arctic waters where relatively large calanoid copepods pre-
dominate, such as various species of Calanus, these mesh sizes
may be an appropriate choice. It must be remembered, however,
that the youngest copepodite stages (e.g. CI of C. finmarchicus)
are not sampled representatively. There is little evidence to suggest
that copepods have an escape reaction that allows them to avoid
an approaching net. On the contrary, the reaction and escape dis-
tances appear to be so short that avoidance by copepods of even
relatively small nets is not a significant source of error. This is
probably also the case for most other forms of mesozooplankton.
For mesozooplankton, therefore, the main issue is escapement
and extrusion through the net and not active avoidance of the
net. An exception to this may be samplers with very small opening.
It has been suggested that avoidance of larger copepods to the
small opening of the OPC is a source of error for this instrument
(Hopcroft, 2002; Zhou and Tande, 2002). The CPR has also a tiny
39
opening, but is designed for high speed and high filtration effi-
ciency. CPR has been found to give much lower abundance esti-
mates in comparisons with other gears, but it is unclear how
much of this effect is due to passage of water through the sampler
without being filtered and the plankton retained (Cook and Hays,
2001). Avoidance of copepods and other mesozooplankton to small
opening samplers is one research topic that needs to be clarified.
For larger zooplankton forms, avoidance is an important source
of error and there is a large body of literature that documents this.
It is also well documented that high speed is one way of obtaining
higher sampling efficiency. Thus there are several studies showing
that LHPR or Gulf III are more effective in collecting mobile fish lar-
vae than are MOCNESS or vertical nets such as WP-2 (e.g. Brander
and Thompson, 1989). In our study the BIONESS caught consis-
tently more of the larger macrozooplankton (krill and shrimps)
than the slower MOCNESS. This was accompanied by a larger loss
of organisms in the small end due to extrusion. High-speed sam-
plers such as LHPR and Gulf-V counteract for this by having a nose
cone that reduces the opening of the net. This improves the filtra-
tion efficiency but reduces the volume of water filtered for a given
tow length. For organisms that occur with low abundance and den-
sity, such as krill in the present study, this may mean that they
have low probability of being caught with high-speed samplers
with small opening and low volume of water filtered.
No one single net is suitable to sample across a wide size range
of zooplankton from small mesozooplankton forms to macrozoo-
plankton. Small mesh nets are needed to capture the larval life
stages of many zooplankton species, but coarser nets need to be
used to capture the adults. In the small end of the size spectrum,
water bottles (e.g. 5–30 l Niskin bottles) may be an alternative
for quantitative sampling of microzooplankton such as eggs and
nauplii of copepods (e.g. Melle and Skjoldal, 1989, 1998). In the
high end of the spectrum, specially designed trawls to collect mac-
rozooplankton and micronekton (such as krill, amphipods, and pe-
lagic shrimps), offer promise for quantitative sampling of mobile
forms that may school and have patchy distribution at scales of
hundreds of meters or more (see trawl descriptions in Krafft
et al., 2010 and Heino et al., 2011). Optical technologies such as
the OPC and the VPR allow observations of zooplankton at fine
scales. Acoustical methods are powerful to record the spatial distri-
bution of zooplankton (particularly macrozooplankton, but also
mesozooplankton; e.g. Postel et al., 2007) and fish at scales from
one meter or less to several hundreds of meters in the vertical
and many kilometers in the horizontal.
A promising avenue for further work is to use a large mouth-
opening net system such as MOCNESS together with measures to
reduce avoidance of capture by using flashing light systems (Wiebe
et al., 1982; Sameoto et al., 1993; Wiebe et al., 2004; Wiebe, 2011).
In combination with video and acoustic remote sensing techniques
(e.g. Greene et al., 1998; Broughton and Lough, 2006), such systems
should be used to provide quantitative estimates of zooplankton
and micronekton required for assessments of ecosystem structure
and dynamics.
Acknowledgments
The data presented in this paper represents a small portion of
that produced by a large number of individuals in a number of
institutions who took part in the sea-going workshop. We thank
all these individuals for their contributions. A special thanks to
the technical staff from IMR and IOW for their skilled efforts in
sample processing and analyses. We also thank the Officers and
crew of the R/V A. v. Humboldt and the R/V Johan Hjort for their
excellent support during the Workshop-at-Sea. Finally we thank
the reviewers for many excellent suggestions for the improvement
of this paper.
 Author's personal copy

40 H.R. Skjoldal et al. / Progress in Oceanography 108 (2013) 1–42

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