Content uploaded by Asako K. Matsumoto
Author content
All content in this area was uploaded by Asako K. Matsumoto on Feb 10, 2018
Content may be subject to copyright.
First findings of decapod crustacea in the hadal zone
A.J. Jamieson
a,
, T. Fujii
a
, M. Solan
a
, A.K. Matsumoto
b
, P.M. Bagley
a
, I.G. Priede
a
a
Oceanlab, University of Aberdeen, Main Street Newburgh, Aberdeenshire AB41 6AA, UK
b
Ocean Research Institute, University of Tokyo, 1-15-1, Minamidai, Tokyo 164-8639, Japan
article info
Article history:
Received 21 July 2008
Received in revised form
30 October 2008
Accepted 6 November 2008
Available online 17 November 2008
Keywords:
Decapoda
Crustacea
Benthesicymus crenatus
Hadal zone
Trenches
Pacific Ocean
abstract
Since the first major hadal sampling efforts in the 1950s, crustaceans of the order
Decapoda have been thought absent from the hadal zone (60 00–11,000 m) with no
representatives documented 45700 m. A baited video lander deployed at 60 07, 6890
and 7966 m in the Kermadec Trench, 8798 and 9729 m in the Tonga Trench (SW Pacific),
6945 and 7703 m in the Japan Trench and 5469 m in the Marianas region (NW Pacific)
has now revealed a conspicuous presence of the Benthesicymid prawn Benthesicymus
crenatus Bate 1881. Decapods were observed at all sites except at 7966m in the
Kermadec Trench and the two Tonga Trench sites, making the deepest finding 7703 m in
the Japan Trench, 2000 m deeper than previously thought. These natantian decapods
were readily attracted to fish bait and, rather than feeding on the bait itself, were
observed preying upon smaller scavenging amphipods. These are the first observations
of predation in the hadal zone. In less than 10h of bottom time, 12 observations of 10
individuals were documented at 6007 m and 5 observations of 3 individuals were
documented at 6890 m in the Kermadec Trench. In the Japan Trench at 6945 m 29
observations of 20 individuals were documented whilst only one individual was seen
at 7703 m. Two individuals were observed in the abyssal Marianas Region (5575 m).
Also, in the Kermadec Trench, individual caridean prawns (Acanthephyra spp.) were
observed at 6007 and 6890 m, proving categorically that the crustacean order of
Decapoda is represented in the hadal zone.
&2008 Elsevier Ltd. All rights reserved.
1. Introduction
Decapod crustaceans have long been considered
to have no representatives in the hadal zone
(6000–11,000 m), a conclusion based largely on the
returns of a series of hadal trawls during the Galathea
and Vitjaz expeditions in the 1950s (Wolff, 1960, 1970).
Although 4700 species of invertebrates and fish were
described from the 33,000 individuals that were recov-
ered from 80 hadal trawls on the Galathea, not a single
decapod was found (Wolff, 1970). The apparent absence of
the decapods was considered to be due to the physiolo-
gical limitations of hydrostatic pressure; the deepest
decapod (Parapagurus sp.) was recorded at 5160 m. Since
then, the deepest findings of decapods are 4785, 4986,
5060, 5413, 5440, 5700 m (Tiefenbacher, 2001;Haedrich
et al., 1980;Gore, 1985b;Bouvier, 1908;Domanski, 1986;
Kikuchi and Nemoto, 1991, respectively). There are how-
ever some tentative reports of natantian decapods at
hadal depths (Peres, 1965;Hessler et al., 1978), but it is
still generally agreed that decapods have no hadal
representatives (Blankenship et al., 2007; Herring, 2002).
In contrast to the wealth of information on shallow
water decapod crustaceans, little is known about the
vertical and horizontal spatial distribution of decapods at
abyssal depths (Tiefenbacher, 2001). At these depths the
scavenging community primarily comprises of lysianas-
soid amphipods and macrourid fishes, both highly
efficient necrophages often observed to rapidly intercept
and consume simulated carrion falls (Ingram and Hessler,
Contents lists available at ScienceDirect
journal homepage: www.elsevier.com/locate/dsri
Deep-Sea Research I
ARTICLE IN PRESS
0967-0637/$ - see front matter &2008 Elsevier Ltd. All rights reserved.
doi:10.1016/j.dsr.2008.11.003
Corresponding author. Tel.: +44 1224 274410; fax: +44 1224274402.
E-mail address: a.jamieson@abdn.ac.uk (A.J. Jamieson).
Deep-Sea Research I 56 (2009) 641–647
1983;Priede et al., 1991). Amongst these more conspic-
uous abyssal scavengers, decapod crustaceans are often
observed, albeit less frequently and perhaps less predic-
tably (Henriques et al., 2002;Janßen et al., 2000;Kemp
et al., 2006). The galatheid crab Munidopsis spp., for
example, is often observed in relatively high numbers at
large carrion falls, such as cetacean carcasses, although
they do not appear for several days (Jones et al., 1998;
Kemp et al., 2006). On shorter time scales, where smaller
baits are consumed within 24 h, the highly active and fast
moving natantian decapod Plesiopenaeus sp., thought to be
a facultative scavenger, is frequently observed but in
lower numbers and often during periods of low fish
activity (Henriques et al., 2002;Thurston et al., 1995).
The aim of this paper is to describe the first observa-
tions of highly active decapod crustaceans in the hadal
trenches of the Pacific Ocean.
2. Methods
2.1. Lander operations
A hadal-rated baited video lander was deployed in the
Southern Hemisphere in the Kermadec trench (at 6007,
6890, 7966 m) and in the Tonga Trench (at 8798 and
9729 m) during July 2007. In the Northern Hemisphere it
was deployed in the Japan Trench (6945 m) and abyssal
Marianas Region (5469 m) during October–November
2007. One further deployment was made in the Japan
Trench (7703 m) in October 2008. The lander was pre-
programmed to record 1min of video every 5 min in
MPEG2 format (704 576 pixels) using a custom-built
video camera (12kCAM-V-1; Oceanlab, UK; NETmc Marine
Ltd., UK). The video system was positioned 1 m off the
seafloor with the camera and twin 50 W lamps facing
vertically down and focussed on an area of seafloor
68 cm 51 cm (0.35 m
2
). One kilogram of bait (blue-fin
tuna, Thunnus thynnus) was secured on a stainless-steel
bar in the centre of the field of view. Pressure (decibars)
and temperature (degree Celsius) were recorded every
30 s throughout using an SBE-39 sensor (Seabird Electro-
nics, USA). Pressure was converted to depth following
Saunders (1981).
As there are presently no current meters available with
sufficient operational depth ratings, it was not possible to
accurately measure current velocity, but estimates were
made by tracking of particles in the water resuspended by
activity of bottom-fauna.
2.2. Study sites
The Japan and Kermadec/Tonga Trench systems exhibit
the characteristic V-shaped topography of many trench
systems with soft sediment on the Oceanic plate side
(to the East) and rockier steep slopes on the continental
plate side (to the West). The Kermadec and Tonga trenches
are open to the incursion of cold, deep Antarctic water and
bottom temperatures are between 1.2 and 1.8 1C making
this trench one of the coldest in the world (Belyaev, 1989).
These trenches are oligotrophic as they lie under the
South Pacific Subtropical Gyre (SPSG) province which has
an average primary production rate of 87gC m
2
yr
1
(Longhurst et al., 1995). The low export of primary
production to the seafloor is likely to increase the
significance of carrion falls as sources of food for
the benthic food web (Blankenship and Levin, 2007). In
contrast to the Kermadec and Tonga trenches, the Japan
Trench, situated in the Kuroshio Province (KURO), which
has higher surface primary production rate (193 g C m
2
yr
1
(Longhurst et al., 1995) and has generally higher
bottom temperatures. The Abyssal Marianas Region site
(5469 m), situated to the east of the Southern Marianas
Trench, is the most oligotrophic of the three sites (primary
production rate ¼82 g C m
2
yr
1
;Longhurst et al., 1995).
2.3. Data analysis
Each minute of MPEG2 video sequence was manually
analysed. Individual decapods were identified using still
images obtained from the MPEG2 sequence in conjunction
with the video sequences. Differences in body size, shape
and carapace markings were sufficient to unambiguously
distinguish each conspecific. Body lengths were measured
in cm by comparison with the in situ scale bar in the field
of view. Over ground swimming speeds were recorded in
cm per second (cm s
1
) and converted to body lengths per
second (BL s
1
). To test the statistical significance for
differences, one-way ANOVA was conducted to compare
the measurements for body length (cm) and swimming
speed (cm s
1
and BL s
1
) across three regions (Japan
6945 m; Kermadec 6007 and 68 90 m; Marianas 5469 m).
All the statistical analyses were performed using SPSS
(SPSS Inc., USA) and the ‘R’ statistical and programming
environment (R Development Core Team, 2005).
3. Results
During deployments at 5469 m in the Marianas Region,
6007 and 6890 m in the Kermadec Trench and 6945 and
7703 m in the Japan Trench, the Benthesicymid prawn
Benthesicymus crenatus Bate 1881 was readily attracted to
bait and apparently undeterred by lander structures or
illumination (Fig. 1). On video, its appearance is similar to
that of the common deep-sea Aristaeid prawn Plesiope-
naeus armatus, perhaps more commonly seen at baits
(Janßen et al., 2000;Thurston et al., 1995). However, the
resolution of the camera was sufficiently high to allow
B. crenatus to be distinguished from P. armatus (shorter
rostrum), although we recognise that both species are
known to co-inhabit in deeper areas of the South Pacific
Ocean (Poupin, 1998).
At 60 07 m in the Kermadec Trench, 12 sightings of
B. crenatus were recorded during 4 h 27 min bottom time.
Based on the length and carapace markings, these were
of 10 individuals, mean (7S.D.) body length 2273.9 cm.
From 6890 m, 4 sightings were recorded between 2 h
42 min and 6 h 02 min post touchdown of 2 individuals
(mean body length ¼22.4 cm72.6 S.D.). At 6945 m in the
Japan Trench, 29 sightings of 20 individual B. crenatus
were made (mean body length 15.3 cm72.9 S.D.) during
ARTICLE IN PRESS
A.J. Jamieson et al. / Deep-Sea Research I 56 (2009) 641–647642
the 9 h 26 min bottom time (first arrival ¼10 min). On the
Abyssal Marianas location, amongst a succession of the
scavenging marcrourid fish Coryphaenoides yaquinae, two
sightings were made of one individual decapod (body
length ¼23.5 cm, arrival time 1 h 02 min). At 7703 m in
the Japan Trench, one individual was briefly observed
traversing the field of view of the camera at 4h 01 min
after touchdown. The individual was swimming too close
to the camera to determine body length or swimming
speed with confidence. The mean body length for the
Kermadec Trench decapods (n¼14) was significantly
larger than that of the Japan Trench decapods (n¼20)
(one-way ANOVA, F
(1, 32)
¼40.1, po0.0001; Fig. 3).
Single Caridean prawns were observed approaching
the bait up-current towards the end of the 6077 and
6890 m deployments in the Kermadec Trench (Fig. 1).
The individual in deployment 1 (body length ¼13.2 cm)
arrived at 3 h 40 min post touchdown and in deployment 2
the individual (14 cm) arrived at 5 h 50 min. Because
of their small body size, however, it was not possible
to confirm their identity with confidence beyond that
of the family Acanthephyra. Other than swimming speeds
of 5.7 and 6.6 m s
1
, respectively, no other information
was obtained and they are simply mentioned here for
reference.
No further sightings of decapods were made beyond
6890 m in the Kermadec Trench (7966 m) or at the Tonga
Trench locations (8798 and 9729 m). A summary of these
observations and environmental characteristics is given in
Table 1.
The presence of scavenging amphipods was recorded
at all sites and increased in numbers with depth. The
resolution of the video camera did not permit positive
identification, however, Blankenship et al. (2006) docu-
mented 4 species captured in traps, from the Kermadec
and Tonga Trenches (Eurythenes gryllus,Scopelocheirus
schellenbergi,Hirondellea dubia and Uristes sp. nov.).
The three deepest sites from the Kermadec and Tonga
Trench deployments (where decapods were absent) were
dominated by extremely large numbers of amphipods.
Based on the large numbers of H. dubia reported by
Blankenship et al. (2006) these can be assumed with some
confidence to be H. dubia. The species of amphipods from
the Japan Trench and Marianas Region are not known.
3.1. Decapod swimming speeds
At the Kermadec 6007 and 6 890 m sites, the typical
behaviour of B. crenatus was to approach the bait at mean
over ground swimming speeds of 7.4 cm s
1
71.8S.D. and
6.9 cm s
1
71.6S.D., respectively (translating as 0.34 BL
s
1
70.08 S.D. and 0.35 BL s
1
70.11 S.D.), stopping
abruptly once in close proximity to the bait, temporarily
interacting with the bait, and leaving the immediate
vicinity. Of the 12 sightings at 6007m, 7 individuals were
seen approaching the bait against the current. Of those
that were observed leaving (10 ind.), 7 left cross-current,
2 drifted back down-current and 1 continued swimming
up-current. Of these 12, each stopped at the bait (never
away from or off the bait) for a mean (7S.D.) duration of
22.8720.2 s. At 6890 m, 3 of the 4 observations showed
individuals approaching the bait from upstream. Only one
was observed leaving (backwards and down-current),
stopping at the bait for 9 s. The general behaviour of
B. crenatus in the Japan Trench was similar to that in the
Kermadec Trench, approaching the bait up-current, stop-
ping in close proximity to the bait and leaving. Of the
18 approaches recorded, 17 were up-current and 1 down-
current. They approached the bait at a mean (7S.D.)
swimming speed of 6.972.0 cm s
1
(0.49 BL s
1
70.17
S.D.). Seventeen approaches were seen to stop on the
bait, but for a shorter time (6.8 s73.8 S.D.) compared to
those in the Kermadec Trench. However, in contrast to the
Kermadec sites, in the Japan Trench individuals showed
a strong tendency to leave the bait and exited the field of
ARTICLE IN PRESS
Fig. 1. The natantian decapod Benthesicymus crenatus (A) and caridean prawns (B and C) in the Kermadec Trench 46000 m (scale bar ¼20 cm).
A.J. Jamieson et al. / Deep-Sea Research I 56 (2009) 641–647 643
view up-current; of the 20 exits observed, 15 exited up-
current, 4 cross-current and 1 down-current. On physical
contact with the bait during approach, one individual
B. crenatus initiated a tail-flip escape response at a speed
of 82.7 cm s
1
. The single individual seen twice in the
Marianas region approached the bait up-current both
times with a mean swimming speed of 7.7cm s
1
72.0 S.D.
(0.33 BL s
1
) and one occasion stayed at the bait for 7 s. On
the second occasion it responded to a large macrourid
with a tail-flip escape response, exiting the area at a burst
swimming speed of 147.1 cm s
1
. Although no statistically
significant difference was found when the mean swim-
ming speeds (cm s
1
;Fig. 4a) were compared, there is a
significant difference when expressed in body lengths per
second (BL s
1
;Fig. 4b).
3.2. Decapod predation behaviour
The decapods preyed upon on scavenging amphipods
already present on the bait, rather than the bait itself.
The presence of amphipods at the bait was consistent
throughout all the locations with first arrival times
of 20 min or less at all locations. In the Kermadec Trench
at 6007 and 6890 m, Japan Trench at 6945 m and the
Marianas Region at 5469 m sites the maximum density of
actively swimming amphipods was 40ind. m
2
(3 h 50 min),
102.9 ind. m
2
(5 h 45 min) 28.6 ind. m
2
(2 h 58 min) and
94.3 ind. m
2
(7 h 47 min), with corresponding bait cover-
age of 60%, 410%, 410% and 410%, respectively. At the
next deepest site in the Kermadec Trench (7966 m) 100%
bait coverage was reached in 2 h 35 min with the density
of actively swimming amphipods reaching 1028 ind. m
2
.
Amphipod densities within the field of view increased
further at the deeper sites at 8798 and 9729m in the
Tonga trench, with peak swimming amphipod densities of
1542.9 and 2914.3 ind. m
2
, respectively and 100% bait
coverage at 1 h 15min and 0 h 45min post touchdown.
The confirmation of predation of amphipods by
B. crenatus was made difficult because the mouth is ventrally
located and seldom in view. In one instance, however,
an individual was observed removing and handling a large
lysianassoid amphipod (20 mm body length). The dec-
apod, 20 cm in length, approached the bait up-current
with its pereiopods trailing below and outwards (Fig. 2A).
It rapidly decelerated once contact was made with the bait
(Fig. 2B). It then reached down, clasped and shuffled its
pereiopods for 3 s while drifting slightly down-current
away from the bait (Fig. 2C). During this time, the
distinctive orange body of the amphipod clearly con-
trasted against the red coloured underside of the decapod.
At the same time, the location on the bait previously
occupied by the large amphipod became visible and
vacant (Fig. 2D). The decapod remained stationary for a
ARTICLE IN PRESS
Table 1
Deployment locations and environmental characteristics with decapod lengths, speeds, occurrences, arrival and staying times at all stations.
Mariana
5469
Kermadec
6007
Kermadec
6890
Japan
6945
Japan
7703
Kermadec
7966
Tonga
8798
Tonga 9726
Latitude 18149.2
0
S26143.9
0
S26148.7
0
S40115.3
0
N36114.8
0
N26155.0
0
S24108.1
0
S24116.4
0
S
Longitude 149150.6
0
E175111.3
0
W175118.1
0
W144130.8
0
E142149.7
0
E175130.7
0
W175111.0
0
W175109.3
0
W
Depth (m) 5469 6007 6890 6945 7703 7966 8798 9726
Primary production rate
(g C m
2
yr
1
)
82 87 87 193 193 87 87 87
Temperature (1C) 1.5 1.2 1.3 1.8 1.9 1.5 1.6 1.8
Estimated current velocity (cm s
1
)5–9 8–9 10–14 3–7 2–4
aaa
Month year Nov ‘07 July ‘07 July ‘07 Oct ‘07 Sept ‘08 July ‘07 July ‘07 July ‘07
Bottom time (hh:mm) 08:55 04:27 06:02 09:26 07:00 10:07 09:05 08:44
Benthesicymus crenatus
Mean body length (cm7S.D.) 23.5 22.073.9 22.472.6 15.372.9
b
–––
Over ground swimming speed
(cm s
1
7S.D.)
7.7 7. 471.8 6.971.6 6.972.0
b
–––
Burst swimming speed (cm s
1
) 147.1 – – 82.7 – – – –
Total number of sightings 2 19 4 29 1 – – –
Number of individuals 1 10 2 20 1 – – –
Mean sightings per hour 0.2 4.7 0.7 3.1 0.14 – – –
First arrival time 01:02 00:41 02:42 00:10 04:01 – – –
Mean bait interaction time (s) 7 22.8 720.2 9 6.8 73.8 0 – – –
Acanthephyra sp.
Body length (cm) – 13.2 14.0 – – – – –
Swimming speed (cm s
1
) – 5.7 6.6 – – – – –
Total number of sightings – 1 1 – – – – –
Number of individuals – 1 1 – – – – –
Mean sightings per hour – 0.2 0.2 – – – – –
First arrival time – 03:40 05:50 – – – – –
a
Indicates that current speed was too low to estimate with confidence, therefore close to 0cm s
1
.
b
Indicates that the length and speed of the decapod was immeasurable because of its swimming altitude.
A.J. Jamieson et al. / Deep-Sea Research I 56 (2009) 641–647644
further 5 s with its forward appendages held tightly
against its ventral surface, before swimming away cross-
current. As the lander had sunk slightly into the sediment,
the area around where the amphipod had been was close
to the sediment-water interface and so ruled out the
possibility that the amphipod had been knocked off or
relocated.
4. Discussion
The observations described here confirm that indeed
decapods do inhabit the hadal zone to at least 7703 m, but
the mechanisms dictating their occurrence and distribu-
tion at this time are unresolved. The behaviour of the
natantian decapods is similar to other baited camera
observations on the shallower abyssal plains. One study,
where the numbers of decapods (P. armatus) were high
was in the Arabian Sea at 40 00–4500 m (Janßen et al.,
2000). Although they were the first to arrive at bait
(within 1 h), they did not appear in the same place in
consecutive images and no loss of bait was visible when
only decapods were present. Similarly, only 40% of
individuals were seen in contact with the bait. This also
suggests that the decapods may have been exploiting the
temporarily high density of amphipods rather than
feeding at the bait itself. In the Atlantic (4000–5000m)
small clusters of P. armatus have been viewed directly on
the bait; however, in both instances still photography
could not confirm predatory behaviour (Thurston et al.,
1995). The lack of bait consumption by decapods in this
study and others (Janßen et al., 2000) suggests exclusive
dependency on carrion falls is unlikely. Stomach contents
from abyssal specimens have comprised phytodetritus,
small bivalves and ground-up crustacean parts (Domanski,
1986;Thurston et al., 1995), further evidence of facultative
necrophagy and possibly active epibenthic predation,
also suggested by these observations and others (Gore,
1985a, b). However, the possibility of some of the stomach
contents being derived from undigested food within the
gut of a swallowed amphipod is not discussed but may be
possible.
Thurston et al. (1995) also reported peaks in decapod
numbers during periods of low current speeds, a relation-
ship reinforced by Domanski (1986), who suggested that
natantian decapods utilise weak currents as a low-energy
mechanism to search for food. In this study, the decapods
in the eutrophic Japan Trench tended to continue foraging
up-current rather than perhaps preserving valuable
energy by drifting cross-current like those in the oligo-
trophic Kermadec Trench, where current speeds are
higher. This continual up-current foraging may only be
possible in the Japan Trench because of its location under
high surface productivity where other food sources
are readily available, as eutrophic trenches are known to
support a higher benthic biomass (Jumars and Hessler,
1976). This may explain why this population exhibits a
significantly higher body length per second swimming
speed (Fig. 4b) as the conservation of energy is less
important. In the oligotrophic Kermadec Trench, where
food is sparser and current speeds are higher, the
tendency to conserve energy by swimming slower and
ARTICLE IN PRESS
Fig. 2. Predation of amphipods by B. crenatus at 6007 m in the Kermadec Trench. (A) Decapod approaching bait with pereiopods out and amphipod
feeding at bait (arrowed). (B) Decapod stopped and grappling with amphipod, just visible under decapod’s body (arrowed). (C) Confirmation that the
amphipod has been removed as the decapod backs off (arrowed). (D) The decapod leaves with pereiopods clasped underneath and amphipod is no longer
at the bait (scale bar ¼20 cm).
A.J. Jamieson et al. / Deep-Sea Research I 56 (2009) 641–647 645
drifting cross-current are perhaps more important.
The larger body size (Fig. 3) may also be an adaptation
to this low-food environment (compared to the eutrophic
Japan Trench), a foraging strategy supported further in the
bait interaction times. Those in the Kermadec Trench
spent approximately three times longer in close proximity
to the bait (and associated amphipods) than those in
the Japan Trench, again suggesting a varying degree
of importance of individual food-falls between the two
trenches. The early arrival times, particularly in the Japan
Trench (10 min) may reflect a high decapod population
density as opposed to highly efficient food-fall intercep-
tion (Sainte-Marie and Hargrave, 1987;Priede at al 1990),
a view also supported by Christiansen and Martin (2000).
B. crenatus have been described from French Polynesia
at 430 0 m around the Tuamoto Archipelago (Poupin, 1998)
and the North West Pacific close to Japan as deep as
5700 m (Kikuchi and Nemoto, 1991). These new findings
place the geographical distribution of B. crenatus further
west into Polynesia in the Kermadec Trench, confirm their
distribution in the North West Pacific and bridge the gap
between the two with sightings in the Micronesian
Marianas Region near Guam. It can be confirmed that
the geographical distribution of B. crenatus currently
known is the west Pacific Ocean between 401N and 261S
with a bathymetric range of 3530 m (Kikuchi and Nemoto,
1991) to 7703, 200 0 m deeper than previously thought.
5. Conclusion
Decapod crustaceans are active at hadal depths,
inhabiting the upper slopes of the Pacific Ocean Trenches
in both the Northern and Southern hemisphere. They are
capable of preying upon the abundant scavenging amphi-
pod community that thrives at these depths and form a
major component of the hadal food web (Blankenship et
al., 2006;Blankenship and Levin, 2007), thus adding a
new element to the hadal food web as a top predator in
the upper trenches. Although the upper limit of the hadal
zone was originally, and to all intents and purposes still is,
defined as 60 00 m, Wolff (1960) recommended 6800–
7000 m as a better reflection of the change in faunal
composition. As the decapods described in this study,
known from as shallow as 3530m are found beyond the
limits of Wolff’s recommendation, these individuals are
ARTICLE IN PRESS
25
20
15
10
Japan Kermadec Marianas
Region
Body length (cm)
Fig. 3. Box plots of the body size (cm) of Benthesicymus crenatus in three
regions. (The centre of the box is the median. The notch in the box
represents an approximate 95% confidence interval for the median. The
top and bottom of the box represent the 3rd and the 1st quartiles,
respectively. The capped whiskers extend to the maximum and
minimum values.)
10
8
6
4
Japan Kermadec Marianas
Region Region
Japan Kermadec Marianas
0.8
0.7
0.6
0.5
0.4
0.3
0.2
Swimming speed (BL s-1)
Swimming speed (BL s-1)
Fig. 4. Box plots of (a) the over ground swimming speed (cm s
1
) and (b) the speed expressed in body length per second (BL s
1
), of Benthesicymus
crenatus in three regions. In (b), the mean value for Japan Trench (n¼20) was significantly larger than that for Kermadec Trench (n¼16) (one-way
ANOVA, F
(1, 34)
¼10.4, po0.01).
A.J. Jamieson et al. / Deep-Sea Research I 56 (2009) 641–647646
non-endemic but at the deeper limits of their habitat that
happens to cross the abyssal–hadal transition zone.
The absence of decapods at 7966 m in the Kermadec
Trench (and the deeper sites in the Tonga Trench) and the
single sighting at 7703 m in the Japan Trench compared to
the larger numbers at the shallower sites suggest that
7700 m may be close to their maximum depth. The
limiting factor to their bathymetric range remains unclear.
The lack of physical specimens prohibits any conclusion
based on hydrostatic pressure as a limiting factor as
suggested by Wolff (1970). The reason decapods have not
been found at hadal depths previously is that they appear
to be extremely difficult to physically catch. The baited
camera method was successful in finding them at these
depths but does not provide the necessary physical
specimen to conclude any adaptations that could explain
why they survive at such high pressure nor any biological
tolerances that still limit them to around 80 00 m. Never-
theless, because of the limited number of hadal sampling
efforts it is unlikely that these observations of decapods
are in fact the deepest, but they do however, highlight
an oversight in our knowledge of deep-sea zonation.
For an entire and relatively conspicuous order of Crustacea
to remain undiscovered from such a large depth zone
emphasises the need for further exploration of these hadal
trenches with appropriate techniques in order to reveal
the true structure of the hadal community.
Acknowledgements
We are thankful to Prof. H.-J. Wagner (University of
Tubingen, Germany) and Prof. H. Tokuyama (University of
Tokyo, Japan) for supporting this work. We thank the crew
and company of the FS Sonne (SO194),RV Hakuho-Maru
(KH07-3) and the RV Kairei (KR07-16). This research was
funded jointly by the Natural Environmental Research
Council (UK) and the Nippon Foundation (Japan) with
additional support from the Sasakawa foundation (Japan)
and the University of Aberdeen (UK).
References
Belyaev, G.M., 1989. Deep-Sea Ocean Trenches and their Fauna. Nauka
Publishing House, Moscow, 385pp (Translated to English by Scripps
Institution of Oceanography, USA, 2004).
Blankenship, L.E., Yayanos, A.A., Cadien, D.B., Levin, L.A., 2006. Vertical
zonation patterns of scavenging amphipods from the Hadal zone of
the Tonga and Kermadec Trenches. Deep-Sea Research I 53, 48–67.
Blankenship, L.E., Levin, L.A., 2007. Extreme food webs: foraging strategies
and diets of scavenging amphipods from the ocean’s deepest 5
kilometers. Limnology and Oceanography 52 (4), 1685–1697.
Bouvier, E.L., 1908. Crustaces decapodes (peneides) provenant des
campagnes de ‘‘Hirondelle’’ et de la ‘‘Princess Alice’’ (1886–1907).
Result. Camp. Scient. Prince Albert I 33, 1–122.
Christiansen, B., Martin, B., 2000. Observations on deep-sea benthope-
lagic nekton at two stations in the northern Arabian Sea: links to
organic matter supply. Deep-Sea Research II 47, 3027–3038.
Domanski, P., 1986. The near-bottom shrimp faunas (Decapoda:
Natantia) at two abyssal sites in the Northeast Atlantic Ocean.
Marine Biology 93, 171–180.
Gore, R.H., 1985a. Abyssobenthic and abyssopelagic penaeoidean shimp
(families Aristaeidae and Penaeidae) from the Vebezuela Basin,
Carribean Sea. Crustaceana 49, 119–138.
Gore, R.H., 1985b. Bright colours in the realm of eternal light. Sea
Frontiers 31, 264–271.
Haedrich, R.L., Rowe, G.T., Polloni, P.T., 1980. The megabenthic fauna
in the deep sea south of New England, USA. Marine Biology 57,
165–179.
Henriques, C., Priede, I.G., Bagley, P.M., 2002. Baited camera observations
of deep-sea demersal fishes of the northeast Atlantic Ocean at
15–281N off West Africa. Marine Biology 141, 307–314.
Herring, P.J., 2002. The Biology of the Deep Ocean. Oxford University
Press, UK.
Hessler, R.R., Ingram, C.L., Yayanos, A.A., Burnett, B.R., 1978. Scavenging
amphipods from the floor of the Philippine Trench. Deep-Sea
Research II 25, 1029–1047.
Ingram, C.L., Hessler, R.R., 1983. Distribution of scavenging amphipods
from the central North Pacific. Deep-Sea Research 30, 683–706.
Janßen, F., Treude, T., Witte, U., 2000. Scavenger assemblages under
differing trophic conditions: a case study in the deep Arabian Sea.
Deep-Sea Research II 47, 2999–3026.
Jones, E.J., Collins, M.A., Bagley, P.M., Addison, S., Priede, I.G., 1998.
The fate of cetacean carcasses in the deep sea: observations on
consumption rates and succession of scavenging species in the
abyssal north-east Atlantic Ocean. Proceedings of the Royal Society B
265, 1119–1127.
Jumars, P.A., Hessler, R.R., 1976. Hadal community structure: implications
from the Aluetian Trench. Journal of Marine Research 34, 547–560.
Kemp, K.M., Jamieson, A.J., Bagley, P.M., McGrath, H., Bailey, D.M., Collins,
M.A., Priede, I.G., 2006. Consumption of a large bathyal food fall, a six
month study in the north-east Atlantic. Marine Ecology Progress
Series 310, 65–76.
Kikuchi, T., Nemoto, T., 1991. Deep-sea shrimps of the genus Benthesi-
cymus (Decapoda: Dendrobranchiata) from the western North
Pacific. Journal of Crustacean Biology 11 (1), 64–89.
Longhurst, A., Sathyendranath, S., Patt, T., Caverhill, C., 1995. An estimate
of global primary production in the ocean from satellite radiometer
data. Journal of Plankton Research 17 (6), 1245–1271.
Peres, J.M., 1965. Apercu sur les resultats de deux plongees effectuees
dans le ravin de Puerto Rico parle bathyscaphe Archimede. Deep-Sea
Research 12, 883–891.
Poupin, J., 1998. Crustacea Decapoda and Stromatopoda of French
Polynesia. Atoll Research Bulletin 451.
Priede, I.G., Smith Jr., K.L., Armstrong, J.D., 1990. Foraging behaviour of
abyssal grenadier fish: inferences from acoustic tagging and tracking
in the North Pacific Ocean. Deep-sea Research 37, 81–101.
Priede, I.G., Bagley, P., Armstrong, J.D., Smith, K.L., Merrett, N.R., 1991.
Direct Measurement of active dispersal of food-falls by abyssal
demersal fishes. Nature 351, 647–649.
Sainte-Marie, B., Hargrave, B.T., 1987. Estimation of scavenger abundance
and distance of attraction to bait. Marine Biology 94, 431–443.
Saunders, P.M., 1981. Practical conversion of pressure to depth. Journal of
Physical Oceanography 11 (4), 573–574.
Thurston, M.H., Bett, B.J., Rice, A.L., 1995. Abyssal megafaunal necrofages:
latitudinal differences in the Eastern North Atlantic Ocean. Inter-
nationale Revue der gesamten Hydrobiologie 80 (2), 267–286.
Tiefenbacher, L., 2001. Recent samples of mainly rare decapod crustacean
taken from the deep-sea floor of the southern Wesrt Europ Basin.
Hydrobiologia 449, 59–70.
Wolff, T., 1960. The hadal community, an introduction. Deep-Sea
Research 6, 95–124.
Wolff, T., 1970. The concept of the hadal or ultra-abyssal fauna. Deep-Sea
Research 17, 983–1003.
ARTICLE IN PRESS
A.J. Jamieson et al. / Deep-Sea Research I 56 (2009) 641–647 647
